United States

Department

of Agriculture

Forest Service

International Institute

of Tropical Forestry

San Juan, PR

Rocky Mountain

Research Station

Fort Collins, CO

General Technical

Report IITF-GTR-26

July 2004

Wildland Shrubs of the

United States and Its

Territories: Thamnic

Descriptions:

Volume 1

John K. Francis, Editor
The U.S. Department of Agriculture, Forest Service, International Institute of Tropical Forestry

operates in cooperation with the University of Puerto Rico, Río Piedras, PR, 00936-4984

——————

The use of trade or firm names in this publication is for reader information and does not

imply endorsement by the U.S. Department of Agriculture of any product or service

Abstract ______________________________________

Francis, John K. ed. 2004. Wildland shrubs of the United States and its Territories:

thamnic descriptions: volume 1. Gen. Tech. Rep. IITF-GTR-26. San Juan, PR: U.S.

Department of Agriculture, Forest Service, International Institute of Tropical Forestry, and

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station. 830 p.

A discussion of the general characteristics of shrubs as a life form and their distribution

within the United States is followed by 311 short monographs containing general descriptions, ranges, ecology, reproductive habits, growth and management, and benefits to

humans, animals, and the environment.

Keywords: Asteraceae, Rosaceae, Fabaceae, Cactaceae, woody plants, hardwoods, shrubs,

perennials

CAUTION:

PESTICIDES

Pesticide Precautionary Statement

This publication reports research involving pesticides. It does not

contain recommendations for their use, nor does it imply that the uses

discussed here have been registered. All uses of pesticides must be

registered by appropriate State and/or Federal agencies before they can

be recommended.

CAUTION: Pesticides can be injurious to humans, domestic animals,

desirable plants, and fish or other wildlife—if they are not handled or

applied properly. Use all pesticides selectively and carefully. Follow

recommended practices for the disposal of surplus pesticides and

pesticide containers.
Wildland Shrubs of the United States and Its

Territories: Thamnic Descriptions:

Volume 1

John K. Francis, Editor

i

Foreword 

Shrubs are an important life form that is difficult to circumscribe because of variations in 

individual species that may be classified as shrubs in some circumstances but as trees or 

herbs in others. Nevertheless the term shrub conjures up an image of a woody plant of 

limited stature; usually with mature specimens of one-third to twice human heights and 

often with multiple stems. Shrubs are important components in many kinds of habitats 

including a role as landscape dominants or icons as well as integrated components of 

other vegetation types. Whereas there are numerous national and regional handbooks on 

trees and grasses and floras of continental, regional, state, and local scales, handbooks of 

shrubs are limited to regional areas or to horticultural, agronomic, ornamental, or wildlife 

biology applications. This volume begins an effort to provide important characteristics 

for native and naturalized shrubs of the United States and its territories. Editor John 

Francis and other contributors provide descriptive, distributional, ecological, 

reproductive, growth, management, and use information on more than 300 shrub species 

in this volume; in all there are more than 5,000 shrub species in the United States and its 

territories. There is more work to do. 

I believe that it is appropriate that this effort was undertaken by the USDA Forest Service 

Institute of Tropical Forestry and the Rocky Mountain Station’s Shrub Sciences 

Laboratory. These institutions represent foci of significant shrub diversity. Shrub 

diversity is enhanced both by warm, mesic and tropical conditions and by aridity and 

diverse topography. The two institutions—one in the tropics and one in the continental 

Western United States desert--represent those contrasting situations. 

E. Durant McArthur 

Project Leader 

Shrubland Biology and Restoration Work Unit 

Shrub Sciences Laboratory 

Rocky Mountain Research Station 
ii 

Introduction

The evolution of the Wildland Shrub Manual begins a long time ago with the observation and 

description of plants; however, its direct ancestor is the U.S. Silvics Manual: “Silvics of North America” 

Volumes 1 and 2 (Burns and Honkala 1990a, 1990b) in which the editor participated as a species author. 

After this, a regional silvics manual developed at the International Institute of Tropical Forestry was 

published as single species’ separate publications and later translated into Spanish and compiled into a 

single volume, “Bioecología de Árboles y Exóticos de Puerto Rico y las Indias Occidentales” (Francis and 

others 2000). Besides the numerous botanical references cited in the following section and in the species 

descriptions, other important references were important contributing sources for this effort. The “WoodyPlant Seed Manual” (Schopmeyer 1974 and the update in progress, Bonner and Nisley 2003) is cited in 

many of the species descriptions. The “Fire Effects Information System” database (Fire Sciences 

Laboratory 2003) contributed a significant portion of the information used is a number of the mainland 

U.S. species in this volume. The Wildland Shrub Manual differs from its predecessors by the extensive use 

of Internet information sources that have not been available previously. 

Latin family, genus, species, and authority of plant names were checked against the “Plants” 

database (Natural Resources Conservation Service 2003). In a few cases, individual authors disagreed with 

the name given in the database and have cited their reasons for using the name in the form they did. 

Common names, for which there is really no central authority and which are likely to take another 

generation to stabilize, were left to the discretion of the authors. Common names cited were generally 

chosen from the many in existence for each species because they were more widely used or because they 

were unique or especially descriptive. 

Although somewhat simplified, the format of each shrub description generally follows that used in 

the silvical descriptions. Sections: General Description, Range, Ecology, Reproduction, Growth and 

Management, and Benefits, are followed by References cited. Each description carries a botanical drawing 

or a photo to aid in recognition. Because the number of shrubs to be described is so great, it was decided to 

make descriptions summaries rather than complete treatises of all information available. Also, to make the 

information useful to as wide an audience as possible, authors were encouraged to use wording that is as 

simple as possible without sacrificing technical accuracy. Although arranged in alphabetical order by genus 

name, a table of contents is provided to make the process of turning to an individual species easier. A 

glossary of technical terms and an index of common names are also provided. 

These descriptions have been previously published on the Internet (Francis 2003) and the number 

of shrub descriptions continues to be added to. The Internet allows revisions of existing descriptions so that 

Internet versions of the descriptions may vary somewhat from descriptions contained in this volume. 

Volume 1 of the Wildland Shrub Manual contains descriptions of 311 shrub species, a number determined 

more by the time permitted for the project than anything else. It is anticipated that the process of compiling 

information and describing shrubs will continue and other volumes will follow. The number of shrubs in 

the United States and its Territories easily justifies the publication of five or more volumes of descriptions. 

I sincerely hope that shrub scientists in the coming years will continue the effort of publishing thamnical 

descriptions to facilitate the management of our wildland shrub natural resource. 

References Cited 

Bonner, F.T., tech. coord. And R.G. Nisley, manag. ed. 2003. Woody plant seed manual. U.S. Department 

of Agriculture, Forest Service, Washington, DC. http://wpsm.net/ index.htm. [not paged]. 

Burns, R.M., and B.H. Honkala, eds. 1990a. Silvics of North America. Vol. 1, Conifers. U.S. Department of 

Agriculture, Forest Service, Agriculture Handbook 654. Washington, DC. 675 p. 

Burns, R.M., and B.H. Honkala, eds. 1990b. Silvics of North America. Vol. 2, Hardwoods. U.S. Department of 

Agriculture, Forest Service, Agriculture Handbook 654. Washington, DC. 877 p. 
iii

Fire Sciences Laboratory. 2003. Fire effects information system. U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Research Station, Ft. Collins, CO. http://www.fs.fed.us/database/feis/ [not 

paged]. 

Francis, J.K., ed. 2003. Wildland shrubs of the United States and its territories. U.S. General Technical 

Report IITF-WB-1. Department of Agriculture, Forest Service, International Institute of Tropical 

Forestry, San Juan, PR and Rocky Mountain Experiment Station, Shrub Sciences Laboratory, Provo UT. 

http://fs.fed.us/global/iitf/ wildland_shrubs.htm. [not paged]. 

Natural Resources Conservation Service. 2003. Plants database. U.S. Department of Agriculture, 

Washington, DC. http://plants.usda.gov/ [not paged]. 

Schopmeyer, C.S., tech. coord. 1974. Seeds of woody plants in the United States. Agriculture Handbook 

450. U.S. Department of Agriculture, Forest Service, Washington, DC. 883 p. 

Acknowledgments 

 I especially want to thank John A. Parrotta, then Team Leader at the International Institute of 

Tropical Forestry, USDA Forest Service, for his appreciation of the need to package species information 

for popular benefit and his encouragement to embark on the Shrub Manual project. I must also thank Ariel 

E. Lugo, IITF Director, for allowing me to dedicate almost 3 years nearly full time to shrub research and E. 

Durant McArthur, Project Leader, Shrub Sciences Laboratory, Rocky Mountain Research Station, USDA 

Forest Service, for graciously welcoming me to work at the Shrub Sciences Laboratory and furnishing 

considerable financial support to the project. These and some 36 other authors wrote thamnical descriptions 

to contribute to the book. More than 100 scientists participated in reviewing the manuscripts before they 

were edited. Isaac Nordlund helped with graphics and programming. I owe as much as any other to Louise 

Kingsbury, Director of Publishing Services for the Rocky Mountain Research Station, for personally and 

promptly editing every one of the manuscripts. 

 John K. Francis 

Research Forester, Retired 
iv 

Contents 

 Page 

Forward and acknowledgements i 

Introduction ii 

Contents iv 

Wildland shrubs 1 

Thamnical Descriptions 

Abrus precatorius 12 

Acacia farnesiana 14 

Acacia greggii 16 

Acacia retusa 19 

Acacia tortuosa 21 

Acer glabrum 24 

Acer spicatum 27 

Acnistus arborescens 30 

Aegiphila martinicensis 32 

Agave missionum 34 

Allamanda cathartica 36 

Amyris elemifera 38 

Aralia spinosa 40 

Ardisia elliptica 43 

Argythamnia candicans 45 

Artemisia arbuscula 47 

Artemisia bigelovii 50 

Artemisia californica 52 

Artemisia cana 57 

Artemisia filifolia 60 

Artemisia frigida 63 

Artemisia longiloba 66 

Artemisia lugoviciana 69 

Artemisia nova 72 

Artemisia pygmaea 75 

Artemisia ridgida 77 

Artemisia rothrockii 79 

Artemisia spinescens 83 

Artemisia tridentata 85 

Artemisia tripartita 88

Arundinaria gigantean 90 

Atriplex canescens 93 

Atriplex confertifolia 96 

Atriplex corrugata 99 

Atriplex cuneata 101 

Baccharis pilularis 103 

Barleria prionitis 105 

Batis maritima 107 

Bidens menziesii 110 

Bixa orellana 112 

Bocconia frutescens 115 

Bontia daphnoides 117 

Borrichia arborescens 119 

Bourreria virgata 121 

Brickellia californica 123 

Bromelia penguin 125 

Brunfelsia lactea 127 

 Page 

Caesalpinia bonduc 129 

Cajanus cajan 132 

Callicarpa americana 135 

Calotropis procera 137 

Capparis indica 140 

Capraria biflora 142 

Capsicum annuum 144 

Capsicum frutescens 147 

Casearia decandra 149 

Casearia sylvestris 151 

Cassine xylocarpa 154 

Castela erecta 156 

Ceanothus velutinus 158 

Celastrus orbiculatus 161 

Celastrus scandens 164 

Celtis reticulata 167 

Cephalanthus occidentalis 170 

Cercocarpus intricatus 173 

Cercocarpus ledifolius 175 

Cercocarpus montanus 178 

Cereus greggii 181 

Cestrum diurnum 184 

Cestrum laurafolium 186 

Chamaebiateria millefolium 188 

Chenopodium oahuense 190 

Chiococca alba 192 

Chromolaena geraniifolium 194 

Chromolaena odoratum 196 

Chrysobalanus icaco 199 

Chrysophyllum oliviforme 201 

Chrysothamnus nauseosus 203 

Chrysothamnus parryi 206 

Chrysothamnus viscidiflorus 209 

Cissampelos pareira 212 

Citrus aurantiifolia 214 

Clematis orientalis 217 

Clibadium erosum 219 

Clidemia cymosa 221 

Clidemia hirta 223 

Coccoloba krugii 225 

Coccoloba microstachya 227 

Coccoloba uvifera 229 

Coffea arabica 232 

Colubrina arborescens 235 

Comptonia peregrina 237 

Conocarpus erectus 240 

Corchorus hirsutus 243 

Cordia lima 245 

Cordia polycephala 247 

Cornus sericea 249 

Coursetia caribaea 252 

Crataegus douglasii 254 
v

Critonia portoricense 257 

Crossopetalum rhacoma 259 

Crotalaria lotifolia 261 

Crotalaria spectabilis 263 

Cuphea hyssopifolia 266 

Curatella americana 268 

Cytisus scoparius 275 

Dalbergia ecastaphyllum 278 

Dasiphora floribunda 280 

Dasylirion wheeleri 283 

Datura metel 286 

Dasmanthus virgatus 289 

Distictis lactiflora 291 

Dodonaea viscosa 293 

Dryas octopetala 295 

Duranta erecta 299 

Encelia farinosa 302 

Ephedra nevadensis 307 

Ephedra viridis 310 

Eriogonum fasciculatum 314 

Erithalis fruticosa 319 

Ernodea littoralis 321 

Erythroxylum areolatum 323 

Erythroxylum brevipes 325 

Eugenia foetida 327 

Eugenia pseudopsidium 329 

Euonymus alatus 331 

Euonymus americanus 333 

Euonymus atropurpureus 335 

Fallugia paradoxa 337 

Faramaea occidentalis 341 

Forestiera acuminata 343 

Forestiera segregata 345 

Fouquieria splendens 347 

Furcraea foetida 350 

Gesneria pedunculosa 352 

Gonzalagunia hirsuta 354 

Gossypium hirsutum 356 

Gouania lupuloides 359 

Grayia spinosa 361 

Guettarda elliptica 363 

Gundlachia corymbosa 365 

Gutierreza sarothae 367 

Gynerium sagittatum 370 

Hamelia patens 372 

Helicteres jamaicensis 374 

Hibiscus pernambucensis 376 

Holodiscus discolor 379 

Holodiscus dumosus 382 

Hyperbaena laurifolia 384 

Indigofera suffruticosa 386 

Isocoma tenuisecta 388 

Itea virginica 391 

Ixora coccinea 393 

Jacquinea arborea 395 

Jasminum fluminense 397 

Jatropha gossypiifolia 399 

Juniperus communis 401 

Juniperus osteosperma 404 

Kalmia latifolia 407 

Kaonophyllon polyodon 411 

Krameria ixine 412 

Lantana camara 414 

Lantana involucrata 417 

Larrea tridentata 419 

Lasiacis divaricata 425 

Leandra krugii 427 

Ledum groenlandicum 429 

Lepianthes peltata 432 

Lepidospartum burgessii 434 

Leucaena leucocephala 437 

Lindera melissifolia 440 

Lonicera japonica 442 

Lotus scoparius 445 

Ludwigia octovalvis 449 

Lupinus arboreus 451 

Lycium pallidum 453 

Macfadyena unguis-cati 456 

Machaonia portoricensis 459 

Mahonia aquifolium 461 

Mahonia repens 464 

Malvastrum americanum 467 

Mecranium latifolium 469 

Melochia nodiflora 471 

Melochia tomentosa 473 

Miconia impetiolaris 475 

Miconia prasina 477 

Miconia racemosa 480 

Mimosa aculeaticarpa 482 

Mimosa arenosa 485 

Mimosa ceratonia 487 

Mimosa pigra 489 

Mimosa pudica 492 

Mimulus aurantiacus 495 

Mitracarpus portoricensis 500 

Morella caroliniensis 502 

Morinda citrifolia 505 

Murraya exotica 508 

Myrica gale 511 

Neea buxifolia 514 

Odontonema cuspidatum 516 

Olyra latifolia 518 

Opuntia leptocaulis 520 

Palicourea crocea 524 

Parathesis crenulata 527 

Parkinsonia aculeata 529 

Parthenocissus quinquefolia 532 

Passiflora edulis 535 

Paullinia pinnata 538 

Pavonia spinifex 540 

Pennisetum purpureum 542 

Penstemon ambiguous 545 
vi 

Philadelphus lewisii 548 

Philadelphus microphyllus 551 

Phoradendron quadrangulare 553 

Phragmites australis 555 

Physocarpus malvaceus 558 

Picramnia pentandra 561 

Piper aduncum 564 

Piper amalago 567 

Piper swartzianum 569 

Piptocoma antillana 571 

Pisonia aculeata 573 

Pithecellobium unguis-cati 575 

Pluchea carolinensis 577 

Prosopis glandulosa 580 

Proustia vanillosma 584 

Prunus americana 586 

Prunus emarginata 589 

Prunus pumila 591 

Prunus virginiana 594 

Psidium guajava 597 

Psorothamnus scoparius 600 

Psychotria brachiata 602 

Psychotria deflexa 604 

Psychotria microdon 606 

Psychotria nervosa 608 

Purshia tridentate 610 

Quercus havardii 613 

Randia aculeata 617 

Rauvolfia viridis 619 

Rhododendron macrophyllum 621 

Rhododendron maximum 623 

Rhus copallinum 625 

Rhus glabra 628 

Rhus microphylla 631 

Rhus trilobata 634 

Ribes aureum 637 

Ribes cereum 641 

Ricinus communis 643 

Rondeletia inermis 645 

Rondeletia pilosa 647 

Rosa woodsii 649 

Rubus discolor 652 

Rubus parviflorus 655 

Rubus rosifolius 658 

Salix arctica 660 

Salix exigua 664 

Salix scouleriana 668 

Salvia apiana 671 

Salvia mellifera 676 

Sassafras albidum 681 

Schaefferia frutescens 684 

Schinus terebinthifolius 686 

Securidaca virgata 689 

Senecio flaccidus 691 

Senecio spartioides 694 

Senna atomaria 696 

Senna occidentalis 699 

Senna polyphylla 702 

Serenoa repens 704 

Serjania polyphylla 708 

Sesbania sericea 710 

Sida fallax 712 

Sida rhombifolia 714 

Smilax domingensis 717 

Solanum drymophilum 719 

Solanum dulcamara 721 

Solanum persicifolium 724 

Solanum torvum 726 

Sorbus scopulina 729 

Spermacoce verticillata 732 

Strumpfia maritima 735 

Styrax americanus 737 

Suaeda suffrutescens 739 

Suriana maritima 742 

Symphoricarpos albus 744 

Tabebuia haemantha 747 

Tamarix chinensis 749 

Tamonea boxiana 753 

Taxus brevifolia 755 

Taxus canadensis 758 

Tecoma stans 761 

Tetrapterys inaequalis 764 

Thrinax morrisii 766 

Toxicodendron radicans 769 

Toxicodendron rydbergii 772 

Trema lamarckianum 775 

Trichilia hirta 777 

Trichostigma octandrum 779 

Triumfetta semitriloba 781 

Turbina corymbosa 783 

Urena lobata 785 

Urera baccifera 788 

Vernonia albicaulis 790 

Vernonia borinquensis 792 

Vernonia proctori 794 

Vernonia sericea 796 

Waltheria indica 798 

Wedelia reticulata 801 

Zisiphus obtusifolia 803 

Zuckia brandegeei 806 

Conclusion 

Glossary of technical terms 808 

Common name index 812 

Authors 824 
1

 

 

Wildland Shrubs 

 According to the dictionary (Merriam-Webster Inc 1984), the word “shrub” came 

from the Old English scrybb wood, which is akin to the Norwegian skrubbebaer that means 

a dwarf hardwood species. The implication seems to be that a shrub is a dwarf tree--not far 

from the current usage of the word. The designation of this life form is driven by the need 

for a category between trees and herbs. 

 If a tree is “a woody perennial plant, typically large and with a single well-defined 

stem carrying a more or less definite crown” (Ford-Robertson 1971), then a shrub must 

necessarily be smaller. Most definitions require that a shrub should have more than one main 

stem caused by branching below or above the ground level (Allaby 1994, Viereck and Little 

1972). Other frequent qualifications include the need for the plant to be perennial and to be 

lignified (woody), at least in some of its parts (Allaby 1994, Ford-Robertson 1971, Orshan 

1989). Shrubs are distinguished from herbs in that herbs do not develop persistent woody 

tissue above ground (Ford-Robertson 1971). However, Lawrence (1955) admits that the 

term shrub is “not subject to precise circumscription”. 
2 

There are several reasons why shrubs cannot be precisely defined. Many species are 

trees in fertile habitat or favorable conditions and shrubs in difficult habitat. Certain species 

may grow as shrubs in large portions of their range and become trees in limited areas. Most 

large shrubs will produce an occasional individual with a clearly tree-like habit. It is not 

possible to define exactly what “large” is (referring to stem size). A number of species 

normally develop single stems but only reach stem sizes below that normally associated with 

trees. The degree of lignification is another source of ambiguity. Many annual herbs become 

quite lignified before dieing, and some perennial shrubs have little or no woody tissue. 

Finally, the concept of perennial looses much of its meaning in the humid tropics where a 

shrub may complete its life cycle in a year of continuous growth, and an herb may live 

without dieback for many years. 

 There are many subtypes of shrubs including dwarf shrubs, half-shrubs (sub-shrubs), 

cushion plants, woody vines (lianes and climbers), scrambling and prostrate forms, woody 

parasitic plants, and various forms of succulents and semi-succulents. A description of these 

forms can be found in Lawrence (1955), and a discussion of criterion for classification of 

shrub forms may be found in Orshan (1989). 

 The concept of the shrub layer, which is populated by functional shrubs, sidesteps 

the ambiguity of classifying species. In this system, the forest is divided into vertical layers 

(canopies or horizons). In their simplest form, the layers are: the tree layer, consisting of 

trees and tree-sized plants, the shrub layer, including all plants with any diameter at breast 

height (d.b.h.) that is less than some arbitrary upper d.b.h. limit such as 5.0 cm, and the herb 

layer, which does not reach breast height. The principal components of the shrub layer are 

actually young trees. This concept is useful in studies of wildlife habitat, plant ecology, and 

biomass and carbon distribution. 

 Because of their large size, influence on the environment, and considerable economic 

value, trees have received most of the attention in forest research and ecological 

descriptions. However, shrubs are more important than is generally realized, especially in 

terms of biodiversity. Little (1979) lists 679 species of trees native and naturalized in the 

continental United States. Another 60 native tree species with an additional number of 

naturalized species grow in Hawaii (Little and Skolmen 1989). Puerto Rico and the U.S. 

Virgin Islands support 547 native tree species (Little and others 1974) and at least 118 
3

species of naturalized exotics (Francis and Liogier 1991). With allowances for species 

represented in more than one area, the U.S. and its territories support about 1,300 tree 

species. Shrub species in the same area are much more numerous. 

 Estimating the number of shrub species in the U.S. and its territories is challenging 

because the life-form is subject to the perception of authors and compilers and not indicated 

or consistently identified in many of the checklists and floras. However, it is possible to 

count the shrub species indicated in the floras of several of the states or regions. Alaska has 

about 110 species of shrubs (Viereck and Little 1972), California about 800 species 

(McMinn 1951), Hawaii about 170 (Degener 1946, Degener 1957, Degener and Degener 

1963), Idaho about 160 (Davis 1952), Pennsylvania about 264 (Rhoads and Klein 1993), 

Puerto Rico about 540 (Liogier 1985, Liogeir 1988, Liogier 1994, Liogier 1995, Liogier 

1997), and tropical Florida about 250 (Long and Lakela 1971). These totals (with 

duplication eliminated) plus other miscellaneous sources (Abrams 1940, Abrams 1944, 

Abrams 1951, Abrams and Ferris 1960, Benson 1969, Correll and Johnston 1970, Everitt 

and Drawe 1993, Great Plains Flora Association 1986, Jones 1975, Morley 1969, Nelson 

1996) exceed 2800 species and seems to account for most of the shrub species in the United 

States and its territories. The Natural Resources Conservation Service (2003) “Plants” 

database recently made it possible to search and list shrub species by region within the 

United States and its territories (Puerto Rico and the U.S. Virgin Islands). Although sums for 

growth habits (life-forms) are higher than if each species were exclusively assigned to a 

single growth habit (eg. Salix exidua Nutt. appears as both tree and shrub), the figures are 

instructive, especially for comparing areas, life-forms, and taxonomic divisions. Searching 

all plant species that carry the growth habit designates, “shrub” or “sub-shrub,” yielded 

5,281 species for the U.S. and territories. There was a great deal of variation between 

political divisions (see figure below, not drawn to scale). California and Texas supported the 

greatest numbers of species with 1,311 and 1,300 respectively. North Dakota and Alaska 

had the lowest with 195 and 200 respectively. The U.S. Virgin Islands had the highest 

average shrub biodiversity with 527 shrub species in only 349 km2 of territory. 

 
4 

Several factors influence the amount of shrub diversity found within a political unit. 

Numbers of shrub species, as well as all plant species, increase as one travels from the harsh 

boreal regions to the humid tropics. Diversity of habitat, such as mountain peaks, deserts, 

riparian areas, and sea shore, is also very important. Southwestern deserts where shrubs are 

the dominant vegetation also tend to have high shrub diversity. 

 Using searches of the “Plants” database mentioned above, the shrubs of the U.S. and 

territories were found to be divided among 166 families. The most important families were: 

Asteraceae, 618 species; Rosaceae, 510 species; Fabaceae, 342 species; Cactaceae, 193 

species; Ericaceae, 189 species; Scrophulariaceae, 182 species; Rubiaceae, 165 species; 

Malvaceae, 148 species; Euphorbiaceae, 128 species; Lamiaceae, 124 species; 

Polygonaceae, 123 species; Companulaceae, 112 species; Boraginaceae, 106 species; 

Ramnaceae, 103 species; and 152 additional families with 82 to 1 species each. 

 Reaching or maintaining a high biodiversity is important to ecosystem health. 

Managers and the public are beginning to realize that subordinate species, as well as the tree 

canopy, are important. It will not be enough, as formerly called for in management plans, to 

establish a tree cover and wait for natural succession to fill in the subordinate species. 

Shrubs, which are near-climax in ecosystems such as subtropical deserts, are, thus, the 

principal species to be maintained or reestablished. In moister wildlands, wildlife managers 
5

have long sought to achieve a certain portion of the landscape in the “brush” stage, 

preferably in disbursed, irregular patches within more advanced forest. 

 The use of shrubs as well as all other types of native plants for reclamation and 

restoration of damaged sites is becoming a very important topic (Hansen 1989). Shrubs are 

planted as seedlings of various types and seeded using the same techniques employed with 

trees (Alder and Ostler 1989), except that densities must often be higher. Because it costs 

less, establishment by site manipulation and natural seeding and succession is preferred 

whenever possible. 

 Shrubs yield many benefits to humankind directly and indirectly. Berries and similar 

small fruits are the most important shrub-derived foods. There are many hundreds of kinds 

of edible, wild berrylike fruits throughout the world. Seasonally harvested and preserved, 

they were once very important to hunter-gatherer tribes and still are important in certain 

rural areas. All our commercial berries descended from wild shrubs, and their wild ancestors 

remain a source of genetic material for breeders. Wildland shrubs also furnish nuts, seeds, 

herbs, greens, and medicinal materials to rural peoples. 

 Fuel is another major direct benefit from shrubs. Although wood from shrubs is not 

present in quantities as great as tree wood in forests, its accessibility and ease of harvest 

have made it a very important fuel source in underdeveloped areas and during recreational 

camping, especially when collected by women and children. Shrubs are even harvested to 

make charcoal in areas with few trees. It has also been suggested that shrub stands could be 

harvested mechanically for industrial biomass fuel (Young and others 1989). 

 Humans benefit indirectly from shrubs through animals that eat them. Grazing 

domestic goats, camels, and, to a lesser extent, sheep and cattle obtain a large part of their 

forage from woody browse. Many wild ruminants preferentially browse shrubs even when 

grass is available. Others supplement their diet of grasses and forbs with forage from shrubs. 

A multitude of birds, mammals, amphibians, and reptiles obtain forage, mast (fruits and 

seeds), and insects from shrub hosts. The cover provided by shrubs is critical for a vast 

number of wild animals. 
6 

 

Finally, wild shrubs are an important esthetic component of our forests, prairies, and 

deserts. As greenbelts and semi-wild urban parks become more widespread, wild shrubs will 

be more appreciated and will be managed along with trees and herbaceous plants. 

 Shrubs also negatively impact humans. Undesirable shrubs often invade managed 

and semi-managed pastures, excluding the more desirable grass, and become weeds in 

croplands. They can temporarily suppress tree seedlings and slow reforestation. Shrubs, 

especially the many thorny and a few poisonous species, are the bane of cross-country hikers 

and woods workers. Forests with a heavy shrub layer are usually less visually pleasing than 

open forests. Shrub understories often are a vehicle for explosive fire spread in seasonally 

dry forest habitats. 

 In the early days of forestry, it seemed to be enough to protect and manage the forest 

as a whole. Later, it was realized that we needed detailed information on each of the tree 

species. To this end, a number of excellent references have been produced (Burns and 

Honkala 1990a, Burns and Honkala 1990b, Burns and others 1998, Francis and Lowe 1999). 

The time is coming when, if we are to manage the lesser species, we will need detailed 

information on each or, at least, the most important shrubs. The objective of this work is to 
7

provide an accessible reference of the biology, ecology, and management of individual 

species of shrubs. The study of the biology, ecology, and management of trees is called 

silvics, and the resulting descriptions are termed silvical descriptions. No such terms have 

come to general use for the study and description of shrubs. Perhaps the terms should be 

designated “thamnics” and “thamnical descriptions” from the Ancient Greek “thamnos”, 

which means shrub. 

References

Abrams, L. 1940. Illustrated flora of the Pacific states. Vol. 1. Stanford University Press, 

Stanford, CA. 538 p. 

Abrams, L. 1944. Illustrated flora of the Pacific states. Vol. 2. Stanford University Press, 

Stanford, CA. 635 p. 

Abrams, L. 1951. Illustrated flora of the Pacific states. Vol. 3. Stanford University Press, 

Stanford, CA. 866 p. 

Abrams, L., and R.S. Ferris. 1960. Illustrated flora of the Pacific states. Vol. 4. Stanford 

University Press, Stanford, CA. 732 p. 

Alder, G.M., and W.K. Ostler. 1989. Native shrub propagation and nursery stock 

production. In: C.M. McKell, ed. The biology and utilization of shrubs. Academic Press, 

Inc., San Diego, CA. p. 535-552. 

Allaby, M. 1994. The concise Oxford Dictionary of ecology. Oxford University Press, 

Oxford, UK. 415 p. 

Benson, L. 1969. The cacti of Arizona. The University of Arizona Press, Tucson, AZ. 218 p. 
8 

Burns, R.M., and B.H. Honkala, eds. 1990a. Silvics of North America. Vol. 1, Conifers. 

U.S. Department of Agriculture, Forest Service, Agriculture Handbook 654. Washington, 

DC. 675 p. 

Burns, R.M., and B.H. Honkala, eds. 1990b. Silvics of North America. Vol. 2, Hardwoods. 

U.S. Department of Agriculture, Forest Service, Agriculture Handbook 654. Washington, 

DC. 877 p. 

Burns, R.M., M.S. Mosquera, and J.L. Whitmore, eds. 1998. Useful trees of the tropical 

region of North America. Publication 3. North American Forestry Commission, 

Washington, DC. 256 p. 

Correll, D.S., and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas 

Research Foundation, Renner, TX. 1881 p. 

Davis, R.J. 1952. Flora of Idaho. W.M.C. Brown Co., Dubuque, IA. 828 p. 

Degener, O. 1946. Flora Hawaiiensis. Books 1-4. Otto Degener, Waialua, Oahu, HI. [Not 

paged]. 

Degener, O. 1957. Flora Hawaiiensis. Book 5. Otto Degener, Waialua, Oahu, HI. [Not 

paged]. 

Degener, O. and I. Degener. 1963. Flora Hawaiiensis. Book 6. Otto Degener, Waialua, 

Oahu, HI. [Not paged]. 

Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs and cacti of South Texas. Texas Tech 

University Press. 213 p. 

Ford-Robertson, F.C., ed. 1971. Terminology of forest science, technology, practice, and 

products. Society of American Foresters, Washington, DC. 349 p. 
9

Francis, J.K. and H.A. Liogier. 1991. Naturalized exotic tree species in Puerto Rico. U.S. 

Department of Agriculture, Forest Service, General Technical Report SO-82. Southern 

Forest Experiment Station, New Orleans. 12 p. 

Francis, J.K. and C.A. Lowe., eds. 1999. Bioecología de especies natives y exóticas de 

Puerto Rico y las Indias Occidentales. U. S. Department of Agriculture, Forest Service, 

General Technical Report IITF-15. International Institute of Tropical Forestry. Río 

Piedras, PR. 583 p. 

Great Plains Flora Association. 1986. Flora of the Great Plains. University Press of Kansas. 

1391 p. 

Hansen, D. 1989. Reclamation and erosion control using shrubs. In: McKell, C. M., ed. The 

biology and utilization of shrubs. Academic Press, Inc, San Diego. 459-478. 

Jones, F.B. 1975. Flora of the Texas coastal bend. Mission Press, Corpus Christi, TX. 262 p. 

Lawrence, G.H.M. 1955. An introduction to plant taxonomy. The Macmillan Company, 

New York. 179 p. 

Liogier, H.A. 1985. Descriptive flora of Puerto Rico and adjacent islands. Vol. 1. Editorial 

de la Universidad de Puerto Rico, San Juan, PR. 352 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto Rico and adjacent islands. Vol. 2. Editorial 

de la Universidad de Puerto Rico, San Juan, PR. 481 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto Rico and adjacent islands. Vol. 3. Editorial 

de la Universidad de Puerto Rico, San Juan, PR. 461 p. 
10 

Liogier, H.A. 1995. Descriptive flora of Puerto Rico and adjacent islands. Vol. 4. Editorial 

de la Universidad de Puerto Rico, San Juan, PR. 617 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto Rico and adjacent islands. Vol. 5. Editorial 

de la Universidad de Puerto Rico, San Juan, PR. 436 p. 

Little, E.L., Jr. 1979. Checklist of United States trees (native and naturalized). U.S. 

Department of Agriculture, Forest Service, Agriculture 

 Handbook 541. Washington, DC. 375 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. U.S. Department of Agriculture, Forest Service, Agriculture 

Handbook 449. Washington, DC. 1,024 p. 

Little, E.L., Jr. and R. Skolmen. 1989. Common forest trees of Hawaii (native and 

introduced). U.S. Department of Agriculture, Forest Service, Agriculture Handbook 679. 

Washington, DC. 321 p. 

Long, R.W. and O. Lakela. 1971. A flora of tropical Florida. University of Miami Press, 

Coral Gables, FL. 962 p. 

Merriam-Webster Inc. 1984. Webster’s ninth new collegiate dictionary. Merriam-Webster 

Inc., Publishers. Springfield, MA. 1,563 p. 

McMinn, H. 1951. An illustrated manual of California shrubs. University of California 

Press, Berkeley, CA. 663 p. 

Morley, T. 1969. Spring flora of Minnesota. The University of Minnesota Press, 

Minneapolis, MN. 283 p. 
11

Natural Resource Conservation Service. 2003. Plants database. http://plants.usda.gov/. [not 

paged]. 

Nelson, Gil. 1996. The shrubs and woody vines of Florida. Pineapple Press, Inc., Sarasota, 

FL. 391 p. 

Orshan, G. 1989. Shrubs as a growth form. In: McKell, Cyrus M., ed. The biology and 

utilization of shrubs. Academic Press, Inc., San Diego, CA. p. 249-265. 

Rhoads, A.F., and W.M. Klein, Jr. 1993. The vascular flora of Pennsylvania, annotated 

checklist and atlas. American Philosophical Society, Philadelphia, PA. 636 p. 

Viereck, L.A., and E.L. Little Jr. 1972. Alaska trees and shrubs. Agriculture Handbook 410. 

U.S. Department of Agriculture, Forest Service, Washington, DC. 265 p. 

Young, J.A., J.D. Budy, and R.A. Evans. 1989. Use of shrubs for fuel. In: C.M. McKell, ed. 

The biology and utilization of shrubs. Academic Press, Inc, San Diego, CA. p. 479-492. 
12 

Abrus precatorius L. crab’s eye

FABACEAE 

Synonyms: Abrus abrus (L.) W.F. Wright 

 Glycine abrus L. 

John K. Francis 

General Description.—Crab’s eye, also known as 

jumbie bean and licorice plant in English, peronías 

and ojos de cangrejo in Spanish, and guen léglise 

in French (Howard 1988, Liogier 1988), is a 

slender woody vine that climbs shrubs and low 

trees. Its older stems are dark gray and the younger 

ones are green and very fine. The alternate, 

pinnately compound leaves are 5 to 10 cm long 

and have five to 20 pairs of leaflets. The racemes 

have tight clusters of white to purple flowers 

(Howard 1988, Liogier 1988). The most notable 

thing about this species is the 6-mm, spherical red 

and black seeds. Crab’s eye produces a tap and 

lateral root system with abundant fine roots. The 

larger roots are dark reddish brown. The lateral 

roots produce white nodules. 

Range.—Crab’s eye is possibly native of India 

(Acevedo-Rodríguez 1985) or of Guinea in Africa 

(Neal 1965), but today has naturalized throughout 

most of the tropics (Acevedo-Rodríguez 1985). 

 

Ecology.—In Puerto Rico, crab’s eye grows in 

subtropical moist (1000 to 2000 mm of 

precipitation), subtropical dry forest (below 1000 

mm of precipitation), and the lower extreme of the 

subtropical wet forest type on all drainage classes 

except very poorly drained and on soils of all 

textures and parent materials. All types of 

topography are colonized from near sea level to 

1,000 m in elevation (in India) (Parrotta 2001). 

Crab’s eye competes well with weeds and brush in 

abandoned farmland, disturbed areas, and early 

secondary forest. It requires disturbance to 

maintain itself in dense, closed stands. Although 

crab’s eye produces relatively few root nodules 

compared to other legumes, nitrogenase activity is 

comparatively high by nodule weight (Pokhriyal 

and others 1997). 

Reproduction.—A sample of crab’s eye seed 

from Puerto Rico averaged 0.1088 + 0.0091 

g/seed. These seeds, which were not treated in any 

way, germinated at a final rate of 61 percent 

between 11 and 182 days after sowing. 

Germination is epigeous. Seeds are produced in 

abundance. Seedlings are common in suitable 

habitat, but few gain access to sufficient sunlight 

to survive. Humans have been responsible for the 

long-distance transport that has resulted in the 

current pantropical distribution. Birds may move 

the seeds short distances either through curiosity 

or by being momentarily deceived into thinking 

that they are edible (personal communication with 

Joseph Wonderly, IITF, Río Piedras, Puerto Rico). 

Lateral extension of the vines also disperses the 

seeds short distances. These plants can be 

controlled by heavy grazing, hand removal, and 

with herbicides. 

 

Growth and Management.—Seedlings grow at a 

moderate rate. It takes about 6 months after 

germination for them to reach 30 cm in height. 

Mature plants may grow 2 m or more per year in 

each leader. They eventually reach 5 to 10 m into 

the canopy or a similar distance laterally 

(Acevedo-Rodríguez 1985). The longevity of 
13

crab’s eye is not reported, but certainly it lives 

longer than 3 years. New plants are easily grown 

in the nursery and probably can be established by 

direct seeding into prepared seed spots. Because 

the species is exotic in all of the U.S. Territories 

and because it becomes a weed in range and 

semimanaged pastures (Velez and von Overgeek 

1950), it is difficult to see a justification for the 

establishment of plantations. 

Benefits.—Crab’s eye seeds were used anciently 

as balance weights in the Eastern countries (Neal 

1965) and as rosary beads by the Buddhists 

(Bailey 1941) because they were so uniform in 

weight, as well as durable. Today, the seeds are 

used to make necklaces and other jewelry and for 

the sounding weights in maracas. The seeds 

contain a protein-based poison called abrin 

(Parrotta 2001). One seed contains enough poison 

to kill a human, but the seeds are very hard and 

must be chewed or ground to release the poison 

(Acevedo-Rodríguez 1985). However, cooking 

destroys the poison so that the seeds may be eaten 

(Neal 1965). The leaves and roots contain 

glycyrrhizin, the principal component of licorice. 

These tissues prepared in various ways are used to 

treat coughs and a number of other ailments 

(Parrotta 2001). Crab’s eye holds out promise in 

the treatment of Schistosoma haematobium

infections. Extracts of the plant were shown to be 

lethal to adult schistosomes in hamsters (Ndamba 

and others 1994). Crab’s eye vines are sometimes 

grown for vegetative screens (Bailey 1941). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. Gen. Tech. Rep. SO-58. 

U.S. Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 331 p. 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. Vol. 1. The MacMillan Co., New 

York. 1,200 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Vol. 4. Arnold Arboretum, 

Harvard University, Jamaica Plain, MA. 673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR 481 p. 

Ndamba, J., N. Nyazema, N. Makaza, C. 

Anderson, and K.C. Kaondera. 1994. Traditional 

herbal remedies used for the treatment of urinary 

schistosomiasis in Zimbabwe. Journal of 

Ethnopharmacology 42(2): 125-132. 

Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub. 

50. Bernice P. Bishop Museum Press, Honolulu, 

HI. 924 p. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CAB International, Wallingford, UK. 

944 p. 

Pokhriyal, T.C., S.P. Chaukiyal, and H.B. Naithani. 

1997. Nodule biomass and nitrogen fixation 

studies in some leguminous nitrogen fixing 

plants. Indian Forester 123(12): 1197-1198. 

Vélez, I. and J. von Overgeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Rìo Piedras, PR. 497 p. 
14 

Acacia farnesiana (L.) Willd. sweet acacia

FABACEAE

Synonyms: Acacia cavenia Bert. 

 Acacia leptophylla DC. 

 Vachellia farnesiana (L.) Wight & Arn. 

John A. Parrotta 

General Description.—Sweet acacia, also known 

as cassie, aroma, huisache, cambrón, espino 

blanco, and many other common names (Little and 

Wadsworth 1964), is a medium-sized shrub with 

many spreading branches and basal stems. The 

leaves are alternate, bipinnately compound with 

two to six pairs of pinnae, each with 10 to 25 pairs 

of narrow leaflets 3 to 5 mm in length. The slightly 

zigzag twigs are dark brown with light-colored 

dots (lenticels) and paired spines 3 to 20 mm in 

length at the nodes. The older bark is also dark 

brown and smooth. Its bright yellow or orange 

flowers, produced over a period of 2 to 4 months, 

depending on locality, are very fragrant and used 

in the perfume industry in France and elsewhere. 

 

Range.—Sweet acacia is believed to be native to 

the American tropics, although precise information 

is lacking about its range prior to the Spanish 

colonial era, during which it was introduced to 

numerous countries throughout the tropics and 

subtropics where it subsequently became 

naturalized. Today sweet acacia is found in the 

southern United States from California to Florida, 

throughout the West Indies, Mexico, Central 

America, in South America as far south as Chile 

and Argentina, and in many parts of the Old World 

tropics and subtropics. Present on all continents 

between 30° N. and 40° S. latitudes, it is the most 

widely distributed species of Acacia (Siegler and 

others 1986). 

Ecology.—Sweet acacia is a drought-hardy, fireresistant species that does not tolerate frost and 

grows well in areas receiving between 500 and 

750 mm of rainfall with a dry season of 4 to 6 

months (Webb and others 1980). Its best growth 

occurs on well-drained soils. It tolerates heavy 

clays to sands and a variety of soil conditions, 

including saline soils, at elevations up to 2,000 m. 

A light-demanding species, sweet acacia often 

forms dense thickets on disturbed sites and is 

associated with numerous other shrub and tree 

species in secondary thorn woodlands, shrublands, 

and dry forests in its tropical and subtropical 

American range (Rzendowski 1981). It is 

susceptible to attack by a number of insect 

species, leaf, stem, and root pathogens, though 

none appear to pose a serious threat to the species 

(Parrotta 1992). 

Reproduction.—Sweet acacia produces small (to 

5 mm in length) flowers that have functional male 

and female parts, borne in compact rounded heads 

0.6 to 1.3 cm across. The flowers are very fragrant 

and are pollinated by bees and other insects. The 

thick, slightly flattened pods, 4 to 9 cm in length 

and 0.5 to 1.3 cm broad, are produced in 

abundance after about 3 years. They mature 4 to 6 

months after flowering and contain a number of 

hard-coated, brown seeds embedded in a pulpy 

mesocarp. The species flowers and fruits between 

November and February in the Caribbean and 

between December and March in Central America 

(Hughes and Styles 1984, Little and Wadsworth 

1964). Natural reproduction is abundant, 

particularly on disturbed sites and in active 

pastures where cattle readily consume the pods. In 

nurseries, sweet acacia is usually propagated from 
15

seed, although branch cuttings can also be rooted 

(Webb and others 1980). A collection of seeds 

from Puerto Rico contained 7,600 seeds/kg, began 

germinating in 6 days, and achieved 57 percent 

germination after 13 days (Francis and Rodríguez 

1993). Although pregermination treatment is not 

necessary, cold- or hot-water soaking, chemical 

scarification, and, particularly, seed-coat 

scarification by abrasion with sandpaper greatly 

increase germination rates (Parrotta 1992). 

Growth and Management.—Early growth is 

relatively rapid. About 1 m of height growth can 

be obtained during the first year, although growth 

rates of 30 to 50 cm during the first year under 

semiarid field conditions are more typical 

(Foroughbakhch and others 1987). Depending on 

the environment, maximum heights of plants 

generally range from 3 to 5 m, with stem 

diameters up to 5 cm. Because sweet acacia is 

intolerant of shade and does not compete well with 

more aggressive woody vegetation such as 

Prosopis L., management activities for enhancing 

growth and natural regeneration in natural and 

plantation stands may include control of 

competing vegetation and periodic soil 

disturbance.

Benefits.—Sweet acacia is planted in many parts 

of its natural and introduced tropical and 

subtropical range for reforestation of degraded 

drylands, for fuelwood and small timber, livestock 

fodder, and for its flowers used in the perfume 

industry. In some areas, it is considered a pest due 

to its ability to colonize pastures and other 

disturbed habitats. The tannin-rich bark is used for 

tanning leather, the gum obtained from the cut 

bark is used as a substitute for gum arabic (from 

Acacia nilotica (L.) Delile), and a useful black dye 

is obtained from the pods. Various parts of the 

plant are used in traditional medicine (Liogier 

1990, Parrotta 2001). In Mexico, for example, the 

flowers are used to treat headache and indigestion, 

whereas a decoction of the green pods is used to 

treat dysentery and skin inflammations. In India, 

the bark, heartwood, and leaves are all used 

medicinally to treat a variety of ailments (Parrotta 

2001). 

References 

Fourouhbakhch, R., R. Penaloza, and H. Stienen. 

1987. Increasing productivity in the matorral of 

northeastern Mexico: domestication of ten 

native multipurpose tree species. In: Strategies 

for classification and management of native 

vegetation for food production in arid zones. 

General Technical Report RM-150. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Forest and Range Experiment 

Station, Fort Collins, CO. p. 90-98. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

Hughes, C.E. and B.T. Styles. 1984. Exploration 

and seed collection of multi-purpose dry zone 

trees in Central America. International Tree 

Crops Journal 3: 1-31. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

Parrotta, J.A. 1992. Acacia farnesiana (L.) Willd. 

–Aroma, huisache. Research Note SO-ITF-SM49. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans. 6 p. 

Parrotta, J.A. 2001. Healing plants of peninsular 

India. CAB International, New York. 944 p. 

Rzendowski, J. 1981. Vegetacíon de México. 

Editorial Limusa, Mexico City. 432 p. 

Siegler, D.S., S. Seilheimer, J. Keesy, and 

H.F.Huang. 1986. Tannins from four common 

Acacia species of Texas and northeastern 

Mexico. Economic Botany 40: 220-232. 

Webb, D.B., P.J. Wood, and J. Smith. 1980. A 

guide to species selection for tropical and 

subtropical plantations. Tropical Forestry Paper 

15. Overseas Development Administration, 

Commonwealth Forestry Institute, University of 

Oxford, London. 256 p. 
16 

Acacia greggii Gray catclaw acacia 

FABACEAE 

Synonyms: Senegalia greggii (Gray) Britt. & Rose. 

Juanita A. R. Ladyman 

 

Illustration source: USDA—Forest Service collection, 

Hunt Institute 

General Description.—A cacia greggii is 

commonly known as catclaw acacia, catclaw, 

Texas mimosa, tearblanket, uña de gato and waita-while (Epple 1995). Children sometimes call the 

plant “Frito” because the wide, flat fruits twist and 

curl like corn chips. Many of the common names 

are similar to those applied to Mimosa 

aculeaticarpa var. biuncifera which it superficially 

resembles. The genus name, Acacia, is derived 

from “akis” that means “thorn” in Greek (Gledhill 

1992). Catclaw acacia is a deciduous shrub, or 

small tree, with grayish-green bipinnately 

compound leaves that are up to 7.6 cm long. 

Catclaw acacia exhibits foliage dimorphism (Isely 

1973). The leaves clustered on spurs are small, 

very short petioled, and usually have one to two 

pairs of pinnae whereas the foliage on new growth 

is larger with petioles up to 1 cm long and with 

one to three pairs of pinnae. The short, sharp, 

curved spines on the branches are approximately 

0.64 cm long and resemble cats’ claws (Epple 

1995). Two varieties exist, var. greggii and var. 

wrightii (Kartesz 1994). Acacia greggii var. 

greggii is synonymous with A. greggii var. 

arizonica Isely; A. greggii var. wrightii (Benth.) 

Isely, is synonymous with A. wrightii Benth. 

(Kartesz 1994). Variety wrightii can be 

distinguished from var. greggii by its wider 

legume and larger leaflets. 

Range.—Catclaw acacia grows in New Mexico, 

Arizona, Nevada, southern Utah, southern 

Colorado, southern California, and Texas, and 

southwards into the Mexican states of Coahuila, 

Chihuahua, Sonora, and Baja California. Variety 

greggii is the most widespread. Variety wrightii

grows in parts of western Texas and New Mexico 

and in Sonora, Tamaupilas, and Nuevo León,

Mexico. 

Ecology.—Catclaw acacia is a drought resistant, 

deep-rooted plant that grows between 300 m and 

1,500 m in elevation in a variety of communities 

and soil types, from sandy or gravelly hills and 

slopes, to canyon bottoms and along washes and 

streams (Dayton 1931, McAuliffe 1995, 

Richardson 1995, Vines 1960). Although most 

commonly a 1- to 4-m tall shrub, it can grow to an 

8- to 10-m tall tree, especially in deep, rich soils 

(USDAFS 1988). It is a frequent member of desert 

grasslands and is often associated with other 

acacia species and a variety of other shrubs 

(Taylor and others 1997). It is not typically a 

dominant member of the community but may 

reach co-dominance in washes (Schmutz and 

others 1992, Uchytil 1990). Catclaw acacia is 

more cold tolerant than many acacia species 

(Bowers 1993). However, in the northern part of 

its range it generally grows no taller than 2 m 

whereas in its southern range it will grow to nearly 

7 m. Catclaw acacia is a winter-deciduous shrub 

that grows only when leafy, remaining dormant in 

the winter and early spring (McClaran 1995). 

Observations suggest that it is a common host to 

mistletoe. An ornithological study in Nevada noted 

that 67 percent of the catclaw acacia shrubs 

distributed along three 1-km transects were 

infested with large amounts of mistletoe, 

Phorodendron californicum Nutt. (Blake 1984). 

Reproduction.—Catclaw acacia flowers from 

April through October with the heaviest blooms in 

April and May (Epple 1995, Everitt and Drawe 

1993). Variety wrightii flowers as early as March 

(Vines 1960). Rain frequently prompts flowering. 

The creamy-yellow, fragrant flowers are insect 
17

pollinated (Bowers 1993). The fruit is a light 

brown to reddish legume often becoming 

constricted between the seeds. The circular seeds 

are 5- to 7-mm in diameter (Isely 1973). The waxcoated seeds delay germination for several years 

and need to be scarified to germinate (Bowers 

1993, Epple 1995). Regions where catclaw acacia 

grows are prone to flash floods and the tumbling 

action of floodwaters in sandy, gravelly washes 

accomplish both seed scarification and seed 

dispersal (Bowers 1993). 

Growth and Management.—Catclaw acacia is a 

thicket-forming shrub that has increased in 

abundance throughout the Southwestern United 

States since the 1890s (Bahre 1995). This increase 

is likely influenced if not directly caused by 

livestock overgrazing and fire suppression (Bahre 

1995). Overgrazing has reduced native perennial 

grasses that, when healthy and dense, can reduce 

woody shrub seedling establishment and also 

provide sufficient fuel to carry a fire. Catclaw is 

susceptible to fire when young, but mature plants 

re-sprout from the root crown following fire 

(Carmichael and others 1978, McPherson 1995). 

Catclaw is moderately resistant to phenoxy 

herbicides, and after only one application the 

shrubs tend to re-foliate or re-sprout from the 

crown (Hibbert and others 1974). Successive 

herbicide applications kill the plants. Apparently 

the invasive resinbrush, Euryops multifidus

(Thunb.) DC. out-competes, or is in some other 

way deleterious to, catclaw shrubs and may be a 

threat in some parts of its range (McAuliffe 1995). 

Benefits.—Catclaw acacia shrubs provide 

protective cover and shade for wildlife (Dayton 

1931). Pods, twigs and foliage, especially new 

growth, provides browse for a variety of wildlife, 

for example jack rabbit, white-tailed and mule 

deer, and the seeds are eaten by many species of 

small mammals and birds, for example, ground 

squirrels, western white-winged dove, scaled and 

bobwhite quail (Graham 1941, Everitt and Drawe 

1993, Taylor and others 1997). The leaves contain 

16 to 20 percent crude protein (Everitt and Drawe 

1993). Cattle browse the shrubs especially early in 

the spring before the new leaves appear, when the 

young green twigs provide some relatively 

succulent forage for a short period of time, and 

when other forage is limited (Graham 1941, 

USDAFS 1988). The mature foliage and pods are 

unpalatable to livestock (Graham 1941, Vines 

1960). The fruits were ground into a flour, or meal, 

called pinole by Native Americans (Epple 1995, 

Vines 1960). It is a food plant for both adult and 

larval butterflies (Taylor and others 1997) and an 

important “bee plant” (Dayton 1931). The lac 

insect, Tachardia lacca, feeds on the sap of 

catclaw acacia and produces a resinous substance 

that can be used in making varnish and shellac but 

probably not in commercial quantities (Powell 

1998). The wood has been used for fuel and tool 

handles (Epple 1995). It is also in the ornamental 

and landscape trade and is a good hedge plant 

(Taylor and others 1997). It has been used in 

projects to re-vegetate degraded land but is 

sensitive to some pollutants such as those found in 

copper mine wastes (Norem and others 1982). 

References 

Bahre, C.J. 1995. Human impacts on the 

Grasslands of southeastern Arizona. In: M.P. 

McClaran and T.R. Van Devender, eds. The 

Desert Grassland. The University of Arizona 

Press, Tucson, AZ. p. 230-264. 

Blake, J.G. 1984. A seasonal analysis of bird 

communities in southern Nevada. The 

Southwestern Naturalist. 29(4): 463-474. 

Bowers, J.E. 1993. Shrubs and trees of the 

Southwest Deserts. Southwest Parks and 

Monuments Assoc., Tucson, AZ. 140 p. 

Carmichael, R.S., O.D. Knipe, C.P. Pase, and W. 

W. Brady. 1978. Arizona chaparral: Plant 

associations and ecology. Research Paper RM202. U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Forest and Range 

Experiment Station, Fort Collins, CO. 17 p. 

Dayton, W.A. 1931. Important western browse 

plants. Miscellaneous Publication 101. U.S. 

Department of Agriculture, Washington, DC. 

214 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. LewAnn Publishing Company, Mesa, 

AZ. 347 p. 

Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs 

and cacti of South Texas. Texas Tech University 

Press, Lubbock, TX. 231 p. 

Gledhill, D. 1992. The names of plants. 

Cambridge University Press, Cambridge, UK. 

202 p. 
18 

Graham, E.H. 1941. Legumes for erosion control 

and wildlife. Miscellaneous Publication 412. 

U.S. Department of Agriculture, Washington, 

DC. 153 p. 

Hibbert, A.R., E.A Davis, and D.G. Scholl. 1974. 

Chaparral conversion potential in Arizona: Part 

I: water yield response and effects on other 

resources. Research Paper RM-126. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Forest and Range Experiment 

Station, Fort Collins, CO. 36 p. 

Isely, D. 1973. Leguminosae of the United States: 

I. Subfamily Mimosoideae. Memoirs of the New 

York Botanical Garden 25(1) 1-152. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada and 

Greenland. Vol. Checklist. Second Ed. Timber 

Press Portland, OR. 622 p. 

McAuliffe, J.R. 1995. Landscape evolution, soil 

formation, and Arizona’s desert grasslands. In:

The Desert Grassland. M.P. McClaran and T.R. 

Van Devender, eds. The University of Arizona 

Press, Tucson, AZ. p.100-129. 

McClaran, M.P. 1995. Desert grasslands and 

grasses. In: The Desert Grassland. M.P. 

McClaran and T.R. Van Devender, eds. The 

University of Arizona Press, Tucson, AZ. p. 1-

30. 

McPherson, G.R. 1995. The role of fire in desert 

grasslands. In: The Desert Grassland. M.P. 

McClaran and T.R. Van Devender, eds. The 

University of Arizona Press, Tucson, AZ. p. 

130-151. 

Norem, M.A., A.D. Day, and K.L. Ludeke. 1982. 

An evaluation of shrub and tree species used in 

revegetating copper mine wastes in the southwestern United States. Journal of Arid 

Environments 5: 299-304. 

Powell, A.M. 1998. Trees & shrubs of TransPecos and adjacent areas. University of Texas 

University Press, Austin, TX (First ed. copyright 

Big Bend Natural History Assoc.) 498 p. 

Richardson, A. 1995. Plants of the Rio Grande 

Delta. University of Texas Press, Austin, TX. 

426 p. 

Schmutz E.M., E.L Smith, P.R. Ogden, M.L. Cox, 

J.O. Klemmedson, J.J. Norris, and L.C. Fierro. 

1992. Desert grasslands. In: Ecosystems of the 

World. 8A. Natural Grasslands Introduction and 

Western Hemisphere. R.T. Coupland, ed. 

Elsevier, London, UK. p. 337-362. 

Taylor, R.B., J. Zrutledge, J.G. Herrera. 1997. A 

field guide to common south Texas shrubs. 

Texas Parks and Wildlife Press, Austin, TX. 

106 p. 

Uchytil, R.J. 1990. Acacia greggii. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/database/feis/ [not paged]. 

United States Department of Agriculture Forest 

Service. 1988. Range Plant Handbook. Dover 

Publications, Inc. New York. 837 p. 

Vines, R.A. 1960. Trees, shrubs, and woody vines 

of the Southwest. Sixth printing. University of 

Texas Press. Austin, TX. 1,104 p. 
19

Acacia retusa (Jacq.) Howard zarza brava 

FABACEAE 

Synonyms: Acacia westiana DC 

 Acacia riparia authors, not Kunth 

 Senegalia westiana (DC.) Britt. & Rose 

 Mimosa retusa Jacq. 

 Mimosa paniculata West 

 Acacia guadalupensis DC. 

 Acacia sarmentosa Griseb. 

John K. Francis 

General Description.—Zarza brava is also known 

as acacia zarza, amourette, and fleur du ben-aimé 

(Acevedo-Rodríguez 1985, Howard 1988, Liogier 

1988). The common name zarza brava is Spanish. 

The modifier “brava” means fierce, which aptly 

describes the way its thorns grab a person trying to 

walk through it. Usually, zarza brava is a climber, 

but when open-grown, it scrambles or grows 

upright. It is also reported to occasionally become a 

slender tree (Howard 1988). The stems are gray, 

squarish when young and angular or fluted when 

older. The old and young stems and even the leaf 

petioles and rachises are armed with recurved 

spines that protect the plant and facilitate climbing. 

The root system consists of many robust laterals 

without an apparent taproot. Nodules were not seen 

by the author. There may be single or multiple 

stems from the ground level, but few branches 

occur on the stem until it reaches increased light in 

the canopy. The leaves are bipinnately compound 

with four to 12 pinnae and 15 to 25 leaflet pairs per 

pinnae (Liogier 1988). The inflorescence is a 

panicle of small heads. The stamens are white. The 

legumes are flat, brown, and 6 to 10 cm long and 1 

to 2 cm wide. The seeds are oval, flat, and dark 

brown (Howard 1988). 

Range.—Zarza brava is native to Hispaniola, 

Puerto Rico, the Virgin Islands, the Lesser Antilles, 

Trinidad, and northern South America (Howard 

1988). It is not known to have been planted or 

naturalized elsewhere. 

Ecology.—Zarza brava grows in habitat that 

receives from as little as 700 mm to as much as 

2200 mm of annual rainfall in Puerto Rico, and is 

more common in the mid-range of this rainfall. 

Soils from all parent materials, including limestone 

and ultramafic rocks, are colonized. Soil textures 

from sand to clays and pH’s from at least 5.0 to 8.0 

are tolerated. Zarza brava can survive under a 

broken overstory. When mats are formed, only the 

upper surface stems produce leaves. The species is 

more common in disturbed habitat than in remnant 

old growth forests. During the abandonment of 

agricultural land, it invades during the brushy 

pasture stage and becomes very common in early 

secondary forest. After the canopy closes, zarza 

brava declines in importance, but does not 

disappear. 

Reproduction.—Zarza brava flowers in June and 
20 

December (Acevedo-Rodriguez 1985) and ripens 

fruit in January and July. Fruit production can be 

very heavy, and pods in one sample in Puerto Rico 

averaged 7.97 + 0.29 seeds/pod. The seeds 

averaged 0.041 + 0.001 g/seed or 24,000 seeds/kg. 

Some of the seeds had been attacked by an 

unknown species of beetle and a lepidopteran 

larva. When planted in commercial potting mix, the 

seeds began germinating in 4 days and by 11 days 

had reached their final 98 percent germination. 

Germination is epigeal. In the wild, seedlings are 

produced in abundance, but few survive beyond the 

first year. Zarza brava sprouts vigorously and 

grows rapidly after fires or cutting. 

Growth and Management.— Seeds are dispersed 

by lateral extension of the vines, by pods tumbling 

laterally through the air after becoming detached 

from vines, and by accidental dispersal by grazing 

animals and humans. New seedlings make 

relatively rapid early growth. Older plants also add 

length rapidly. The stems of zarza brava may 

eventually reach as much as 10 cm in diameter and 

as much as 12 m into tree canopies (AcevedoRodríguez 1985). No planting experience of zarza 

brava is known, but if establishment of plants is 

needed, doubtless the planting of potted nursery 

seedlings would succeed. The species is so 

aggressive that direct seeding into prepared seed 

spots in forest openings should also give good 

results. 

Benefits.—Zarza brava is eaten by goats, but 

seldom touched by cattle. In fact, it survives as 

scrambling patches in overgrazed cattle pasture. On 

the other hand, the species is a great nuisance to the 

cattleman, woods workers, and hikers. It is 

frequently necessary to cut a path through zarza 

brava thickets with a machete, in order to move 

through vegetation in the semiarid-zones within the 

zarza brava range. When one contacts the vines, 

the tips of the spines break off in the skin, causing 

pain and sometimes resulting in infection. No 

specific control recommendations have been 

developed. In the past, control was attempted with 

mixed success by repeated cutting. Control could 

probably be improved by treating the sprouts with 

herbicides after cutting. 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 
21

Acacia tortuosa (L.) Willd. twisted acacia 

FABACEAE 

Synonyms: Mimosa tortuosa L. 

 Poponax tortuosa (L.) Raf. 

 Acacia parvifolia Duss 

John K. Francis 

General Description.—Twisted acacia, also 

known as acacia-bush, wild-poponax, sweet-briar, 

cossie, aromo, cají torcido, and pompons jaunes, is 

a deciduous shrub or small tree to 6 m in height 

and 15 cm in trunk diameter. The plant may have 

single or multiple stems with gray-brown, finely 

fissured bark and a light brown inner bark with 

darker streaks. The wood is light brown and hard. 

The twigs are slender with a zig-zag profile, 

produce paired, straight spines at the nodes, and 

have whitish lenticels on the reddish-brown bark. 

Mature twisted acacia are flat topped and have a 

thin crown of feathery foliage. The bipinnately 

compound leaves are 2.5 to 9 cm long with two to 

eight pairs of pinnae and 10 to 20 pairs of leaflets 

per pinnae. Solitary or small clusters of heads of 

tiny, yellow-orange flowers are borne at the leaf 

axiles. Dark brown or black semiwoody, 8- to 14-

cm pods (legumes) are straight or slightly curved 

and somewhat constricted between the seeds. They 

do not open. The ovoid seeds are brown and hard 

(Howard 1988, Liogier 1988, Little and others 

1974). The chromosome number is 2n = 26 (Long 

and Lakela 1971). 

Range.—Twisted acacia is native to Jamaica, 

Hispaniola, Puerto Rico, the Virgin Islands, the 

Lesser Antilles, Trinidad, and in South America, 

Venezuela, Colombia, and Ecuador including the 

Galápagos Islands (Howard 1988, Instituto 

Humboldt 2002, Liogier 1988, Little and others 

1974). It is endangered in Florida where it may be 

native or introduced (Little and others 1974, 

Institute of Systematic Botany 2002). Although 

several reputable sources list Texas, Mexico, and 

Central America as being part of the A. tortuosa

range (e.g., Missouri Botanic Garden 2002, Nature 

Conservancy 2002), Little and others (1974) and 

Texas A & M Herbarium (2002) maintain that this 

western population is in fact A. schaffneri (S. 

Wats.) Herm. 

Ecology.—Twisted acacia is intolerant of shade 

and only moderately competitive. It finds 

conditions for establishment and growth to 

reproductive age in disturbed, dry areas. 

Disturbance since European colonization has been 

by heavy grazing, fires, excessive timber harvest, 

and clearing for farming or pasture improvement. 

One of the most common plants of disturbed areas 

in Curaçao is twisted acacia (Carambi Foundation 

2002). It is found on shell mounds and along 

roadsides in Southern Florida (Nelson 1996). The 

species grows in coastal areas of Puerto Rico and 

its offshore islands in areas that receive from 750 

to about 1100 mm of precipitation. The plant 

colonizes well-drained soils of the full range of 

textures derived from both sedimentary and 

igneous rocks. Most of these soils are neutral to 

mildly alkaline. Twisted acacia is not damaged by 

mild salt spray. It is usually ignored by cattle and 

tolerates browsing by goats. 

Reproduction.—Twisted acacia flowers 

intermittently throughout the year (Little and others 
22 

1974). Bees and other insects pollinate the flowers 

(Treeguide 2002). A collection of pods from 

Puerto Rico contained an average of 12.4 

seeds/pod and a range of eight to 15 seeds/pod. The 

air-dried seeds separated from them averaged 

0.0440 + 0.0004 g/seed or 22,700 seeds/kg. These 

were mechanically scarified (nicked) and incubated 

on moist blotter paper. Ninety-nine percent 

germinated between 5 and 21 days after sowing 

(author’s observation). Because of the hard, 

impermeable seed coat, germination without 

scarification is low and erratic. Any technique that 

perforates the seed coat will work. One source 

(Treeguide 2002) recommends placing the seeds in 

boiling water, immediately removing the heat 

source and letting the seeds soak for 24 hours. 

Germination is epigeal. The seeds are dispersed by 

herbivores, particularly ruminants that eat the pods. 

Seedlings often emerge in large numbers, but 

nearly all are consumed by grazing animals before 

they have a chance to become woody and thorny. 

Growth and Management.—Although acacias 

can be propagated with cuttings, the use of seeds is 

recommended (Bonsai Clubs International 2002). 

No management experience has been published; 

however, nursery production and field 

establishment will probably be as easy as it is for 

other acacias. Control of twisted acacia along with 

other secondary dry-forest species has been 

accomplished in several areas in Puerto Rico and 

the U.S. Virgin Islands by bulldozing. The use of 

heavy disks might be a better option where 

removal of woody debris is not required. 

Benefits.—Twisted acacia helps protect the soil in 

its disturbed habitat. It is listed as a nitrogen-fixing 

species (Winrock International 2002). The seeds 

are eaten by the threatened yellow-shouldered 

amazon parrot (Amazona barbadensis) in Bonaire 

and probably other parts of its range (Island 

Resource Foundation 2002). The species serves as 

cover for wildlife species. Cattle and other 

livestock eat the pods while goats and sheep 

browse the foliage, especially when other forage is 

scarce. The wood is used to a limited extent for 

fuel. Twisted acacia plants are used to create 

bonsai plants (Bonsai Clubs International 2002). 

The species has been grown and pruned for hedges 

(Little and others 1974). 

References 

Bonsai Clubs International. 2002. Acacia—Acacia

sp. http://www.bonsai-bci.com/species/acacia. 

html. 3 p. 

Carambi Foundation. 2002. Vegetation. http:// 

www.curacao.com/carambi/veget.htm. 1 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Institute of Systematic Botany. 2002. Atlas of 

Florida Plants: Acacia tortuosa. University of 

South Florida, Tampa, FL. 

http://plantatlas.usf.edu/main.asp?plantID=1545. 

1 p. 

Instituto Humboldt. 2002. Plantas vasculares: 

angiospermas y gimnospermas. http://www. 

humboldt.org.co/diqadeladiversidad/resultados/v

asculares.html. 6 p. 

Island Resource Foundation. 2002. Threatened and 

endangered birds of the Insular Caribbean. 

http://www.irf.org/bbarden.htm. 13 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Long, R.W. and O. Lakela. 1971. A flora of 

tropical Florida. University of Florida Press, 

Coral Gables, FL. 962 p. 

Missouri Botanical Garden. 2002. Current 

specimen list for Acacia tortuosa. 

http://mobot.mobot.org/cgi-bin/search_vast. [not 

paged]. 

Nature Conservancy. 2002. The history of the 

Plains and its biota. 

http://www.greatplains.org/resource/biodiver/bio

stat/ecoregio.htm. 13 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 
23

Texas A&M Herbarium. 2002. Checklist of the 

vascular plants of Texas. http://csdl.tamu.edu/ 

FLORA/cgi/msproj/shao/query.pl?query=Acacia

+tortuosa &rform=. 1 p. 

Treeguide. 2002. Huisachillo, Acacia tortuosa, 

Willd. Fabaceae. http://treeguide.com/nn/ 

Species.asp?SpeciesID=108&Region=NorthAme

rica. 2 p. 

Winrock International. 2002. Nitrogen fixing trees 

and shrubs. http://winrock.org/forestry/factpub/ 

nftlist.htm. 29 p. 
24 

Acer glabrum Torr. Rocky Mountain maple 

ACERACEAE 

Synonyms: Acer tripartitum Nutt. 

John K. Francis 

General Description.—Rocky Mountain maple, 

also known as mountain maple and Douglas 

maple, is an upright, deciduous shrub or small tree, 

that ranges in height from 2 to 10 m, depending on 

variety and habitat. It usually grows with multiple 

stems, suckering from the base, and narrow 

crowns, and may be spindly in closed stands. The 

stem bark is thin, smooth, and gray, brown, or 

reddish brown. Branches are opposite and sparse 

with reddish twigs. Roots are woody, extensive, 

and often deep. Leaves vary considerably, 

depending on variety. They are opposite, glabrous, 

and have three to five lobes and rarely three 

distinct leaflets, and range from 2 to 10 cm wide 

and long. The margins are serrate or doubly 

serrate. They may be papery or leathery, 

depending on location and variety. Rocky 

Mountain maple occurs as monoecious and 

dioecious trees that have small greenish flowers 

borne in loose, few-flowered terminal corymbose 

cymes. Fruits are light brown (when mature) 

double samaras that diverge at angles less than 45° 

with wings about 2 cm long. (Abrams 1951, 

Anderson 2001, Wasser 1982, Welsh and others 

1987). 

Range.—Rocky Mountain maple occurs from 

southern Alaska south into northern Mexico and 

east from the Pacific Coast to Alberta, South 

Dakota and Nebraska (Anderson 2001). There are 

currently six recognized varieties: var. diffusum

from the Pacific Southwest, var. douglasii from 

the Pacific Northwest, var. greenei from 

California, var. glabrum from the inland West, var. 

meomexicanum from the inland Southwest, and 

var. torreyi from Oregon, California, and Nevada 

(Natural Resources Conservation Service 2003). 

They differ at least in leaf shape, size, and 

thickness. 

Ecology.—Rocky Mountain maple is moderately 

shade tolerant. It can grow (slowly) in the 

understory and midstory of closed coniferous 

forest but grows in full sun as well. A pioneer in 

disturbed areas, it starts from sprouts and seeds in 

burned areas, clearcuts, avalanche fields, and road 

sides, cuts and fills. It may grow along streams in 

drier areas. Rocky Mountain maple is a member of 

a huge number of plant associations, and is 

frequently dominant during seral stages of 

succession (Anderson 2001, Johnson 1995). The 

species grows in soils derived from most rock 

types, in all soil textures, and in skeletal soils. 

These are moist, well-drained to somewhat poorly 

drained soils that are moderately acid to slightly 

alkaline. It may be found at elevations from 350 m 

in British Colombia to 3,900 m in Colorado and 

Utah. Rocky Mountain maple is slightly more 

common in southern than northern aspects. Some 

of its locations have mean annual precipitation as 

low as 230 mm and as high as 1,500 mm 

(Anderson 2001). It tolerates competition well, 

usually outgrowing and overtopping young trees 

and most shrub species. Dull red leaf spots are 

caused by an unknown pathogen and bright red 

eriophyid mite colonies develop on the underside 

of leaves in some areas. The effects of these and 

other diseases appear to be negligible to 

populations of the species. 

Reproduction.—Flowering occurs in early spring, 

depending on the elevation and latitude (Wasser 

1982). Fruits mature in late summer or early 
25

autumn. Good seed crops occur every 1 to 3 years. 

There are about 30,000 cleaned seeds/kg (Zasada 

and Strong 2003). Seeds disperse by whirling 

sideways as they descend, often carried by the 

wind. Seeds sometimes lie dormant for one or two 

growing seasons before germinating. Germination 

is epigeal. Natural seedlings can be common to 

abundant. 

Growth and Management.—Rocky Mountain 

maple seedlings grow about 30 cm/year (Hansen 

2003). On good sites sprouts may reach 1.3 m 

within 2 years and 3 m in 10 years. Maximum 

heights are reached in 30 to 40 years (Anderson 

2001). Fruits can be picked by hand from low 

shrubs or clipped or shaken from tall shrubs and 

small trees. Mechanical dewinging may be 

advantageous to reduce bulk and improve handling 

before planting. The seeds can be stored for up to 

3 years (Hansen 2003). Stratification of 180 days 

at 20 to 30 °C followed by 180 days at 3 to 5 °C is 

recommended before sowing. Germination of 

about 40 percent can be expected in about 30 days 

(Zasada and Strong 2003). Alternately, seeds are 

sown after collection and beds are maintained for 2 

years because a substantial amount of the 

germination will occur after the second winter. 

Seeds are sown 0.6 to 2.5 cm deep and mulched in 

shaded beds. Seedlings are produced as bare-root 

stock or containerized plants (Zasada and Strong 

2003). The species can be propagated by layering 

and rooted cuttings (Hansen 2003). 

Benefits.—Rocky Mountain maple is an important 

part of the forest understory and pioneer 

vegetation following fires and clearcuts. It helps 

protect the soil, adds beauty to the forest, and 

furnishes food and cover for wildlife. It is an 

important browse species for domestic livestock, 

especially sheep, and wild ungulates. Samples of 

summer growth contained about 8.7 percent crude 

protein, 3.1 percent fat, 34.0 percent crude fiber, 

51.0 percent N-free extract, and 3.2 percent ash. 

Samples of winter browse contained 5.9 percent 

crude protein, 2.4 percent fat, 33.3 percent crude 

fiber, and 54.2 percent N-free extract (Anderson 

2001). The seeds and vegetative parts are 

consumed by ruffed and blue grouse, grosbeaks, 

and small mammals (Anderson 2001). The wood is 

hard, close-grained, and flexible. It was used by 

Native Americans for arrow shafts, snowshoe 

frames, ceremonial equipment, hut frames, and 

drying and smoking racks. The inner bark 

furnished material for mats and ropes. Today, the 

wood is mainly used for firewood (Hansen 2003) 

and smoking meat (Johnson 1995). The young 

shoots may be cooked and eaten as a vegetable 

(Pratt and others 2003). Native Americans made 

infusions and decoctions of the twigs and wood to 

treat swelling and nausea, respectively (Moerman 

1986). Rocky Mountain maple is widely used in 

environmental restoration projects within its native 

range and is an important ornamental species 

valued for its petite size, tolerance to shade, and 

bright red, yellow, and orange autumn colors 

(Hansen 2003). 

References 

Abrams, L. 1951. Illustrated flora of the Pacific 

States. Vol. 3. Stanford University Press, 

Stanford, CA. 866 p. 

Anderson, M.D. 2001. Acer glabrum. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shrub/ 

acegla/all.html. 62 p. 

Hansen, W.W. 2003. Native plants of the 

Northwest: Acer glabrum (Douglas maple, 

Rocky Mountain maple, mountain maple). 

http://www.nwplants.com/plants/trees/aceraceae

/acer_glabrum/ 5 p. 

Johnson, F.D. 1995. Wild trees of Idaho. 

University of Idaho Press, Moscow, ID. 212 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports 19. The Museum of 

Anthropology, University of Michigan, Ann 

Arbor, MI. 534 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Acer glabrum Torr., Rocky 

Mountain maple. http://plants.usda.gov/ 

cgi_bin/plant_profile.chi?symbol=ACGL. 5 p. 

Pratt, M., J. Bowns, R. Banner, and A. Rasmussen. 

2003. Range plants of Utah: Rocky Mountain 

maple. Utah State University Extension. 

http://extension.usu.edu/coop/natres/range/ 

Woody/rockymtnmaple.htm. 4 p. 

 

Wasser, C.H. 1982. Ecology and culture of 

selected species useful in revegetating disturbed 

lands in the West. FWS/OBS-82/56. Fish and 

Wildlife Service, U.S. Department of Interior, 

Washington, DC. 347 p. 
26 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins (eds.). 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 

Zasada, J.C. and T.F. Strong. 2003. Acer L. In: 

F.T. Bonner, and R.G. Nisley, eds. Woody Plant 

Seed Manual. U.S. Department of Agriculture, 

Forest Service, Washington, DC. 24 p. 
27

Acer spicatum Lam. mountain maple 

ACERACEAE 

Synonyms: none 

Paula M. Pijut 

Illustration source: USDA—Forest Service collection, 

Hunt Institute 

General Description.—Mountain maple is a tall, 

deciduous shrub or clumped small tree that grows 

3 to 10 m tall and can spread to form dense 

thickets (Dirr 1998; Gleason and Cronquist 1991; 

van Gelderen and others 1994). In the northern 

part of its native range mountain maple reaches a 

maximum height of 6 m (Sullivan 1993). Leaves 

are opposite, simple, three-lobed or obscurely fivelobed, coarsely and irregularly serrate, dark 

yellowish-green, glabrous with impressed veins 

(adaxial), lighter green and pubescent (abaxial), 

truncate to cordate at the base, and 5 to 13 cm long 

and wide (Dirr 1998; Gleason and Cronquist 1991; 

van Gelderen and others 1994). The leaves turn 

yellow, orange, or most frequently brilliant red in 

the fall. Stems are grayish pubescent when young, 

developing a purplish-red color on the upper 

surface. Leaf scars are narrowly crescent shaped 

(Dirr 1998). Buds are approximately 0.6 cm long, 

stalked, pointed, red with minute, appressed, 

grayish hairs, and with two to four valvate scales 

(Dirr 1998; Gleason and Cronquist 1991). The 

flowers are small, greenish yellow, in fascicles of 

two to four, borne in erect 3 to 8 cm long panicles. 

Each flower is functionally either pistillate or 

staminate, although the non-functioning structures 

still exist in the flower, reduced to a greater or 

lesser extent. Staminate flowers drop soon after the 

pollen is shed (De Jong 1976). Each flower is 

borne on a slender stalk about 1.3 cm long and the 

panicles are found at the branchlet tips. The bark 

is thin, brown to reddish-brown, smooth, 

becoming furrowed or warty with age (Dirr 1998). 

The fruit is a 2.5 cm long, two-winged samara, 

bright red when mature, with divergent wings at an 

acute or right angle. The chromosome number of 

mountain maple is 2n = 26 (Gleason and Cronquist 

1991). 

 

Range.—Mountain maple occurs from 

Newfoundland to Saskatchewan, south to 

Connecticut, Pennsylvania, Ohio, and northeastern 

Iowa, and in the mountains to North Carolina and 

Tennessee (Gleason and Cronquist 1991). 

 

Ecology.—Mountain maple prefers rich, moist, 

acid, well-drained soils with low, diffuse light 

(Dirr 1998; Sullivan 1993). These sites include 

cool woods with humid climate and year-round 

precipitation, rocky slopes and flats, along 

streams, in ravines, and on moist hillsides. 

Mountain maple also grows well on podzol soils, 

talus slopes, and in forested bogs (Sullivan 1993). 

Mountain maple can form a canopy on cliff faces. 

Mountain maple is an understory or subcanopy 

shrub that is tolerant of deep shade, but also grows 

well in full sun, has medium moisture and fertility 

requirements, and colonizes the understory as 

pioneer tree species decline (Sullivan 1993; 

USDA, NRCS 2002). Mountain maple is not salt 

tolerant, and has low drought tolerance. Release 

of mountain maple by canopy tree harvest allows 

the shrub to dominate the site within 5 to 10 years, 

suppressing the growth of spruce and fir seedlings 

for at least 35 years (Sullivan 1993). In 

undisturbed, mature red pine-white pine sites in 

northeastern Minnesota, mountain maple forms a 

dense, high shrub layer with beaked hazel and 

American hazel that inhibits reproduction of later 

successional species, such as balsam fir and spruce 

(Sullivan 1993). Eradication of mountain maple is 

sometimes necessary for tree regeneration. The 

weevil, Phyllobius oblongus L., is especially 
28 

damaging to mountain maple (Johnson and Lyon 

1991). The larvae feed on the roots, mature, and 

overwinter in the soil. Adults are brown weevils 

that defoliate mountain maple. They are found 

throughout the northeast into Michigan and the 

border provinces of Canada (Johnson and Lyon 

1991). Kabatiella apocrypta (Ell. & Ev.) v. Arx 

(anthracnose pathogen) causes minor damage to 

mountain maple, although severe infections can 

led to premature defoliation (Sinclair and others 

1987). Mountain maple is also susceptible to 

attack by Verticillium dahilae Kleb. (verticillium 

wilt), Phyllactinia guttata (Wallr.:Fr.) Lev. 

(powdery mildew), Nectria sp. (nectria canker), 

Cristulariella depraedens (Cooke) Hohn. (leaf 

spot/blight), Pseudomonas syringae pv. syringae

van Hall (bacterial leaf spot and dieback), 

Phyllosticta minima (Berk. and M.A.Curtis) 

Underw. and Earle (leaf spot), Rhytisma sp. (tar 

spots), and Venturia acerina Plakidas ex Barr (leaf 

blotch) (Farr and others 1989, Jones and Benson 

2001, Sinclair and others 1987). 

Reproduction.—Mountain maple staminate 

flowers and pistillate flowers occur on the same 

plant. Some individuals may flower completely 

with staminate flowers. Mountain maple is 

predominately duodichogamous (producing 

flowers in three consecutive phases, male-femalemale, during anthesis), with a small part of the 

population protogynous (stigma receptive before 

the pollen is shed) (van Gelderen and others 1994). 

Low temperatures appear to favor development of 

female flowers and flowers with less reduction of 

the non-functioning parts. (van Gelderen and 

others 1994). The shrub blossoms in May to June 

after the leaves are fully developed; fruit ripens in 

September to October, with seed dispersal in 

October to December (Zasada and Strong 2000). 

Mountain maple is insect pollinated (Sullivan 

1983) and the seed is wind disseminated. Seeds of 

mountain maple need to be stratified for 90 to 120 

days at 5 oC for good germination (Dirr 1998). 

Mountain maple does not build up a seed-bank and 

seedling establishment is better on undisturbed 

soils (Sullivan 1993). Seedling reproduction of 

mountain maple does not result in dense, 

competitive stands, as can be the case with its 

vegetative reproduction. Mountain maples 

frequently layer, giving them the potential to 

develop relatively complex clones of varying size 

and morphology (Zasada and Strong 2000). 

Mountain maple can reproduce by sprouting from 

underground lateral stems, rarely from root 

suckers, and colonies usually develop following 

disturbances by browsing or cutting (Sullivan 

1993). Propagation of mountain maple by cuttings 

taken early in the summer produces strong plants 

(van Gelderen and others 1994). 

Growth and Management.—Mountain maple is a 

slow to medium growing shrub or small, short 

(crooked) trunked tree of bushy appearance (Dirr 

1998). It is hardy in zones three to seven at high 

elevations (USDA Plant Hardiness), and starts to 

decline at 40 to 50 years of age (Sullivan 1993). 

Mountain maple is girdled at the root collar by 

low, surface fires, and can re-sprout from 

underground stems (Sullivan 1993). A severe, hot 

fire is necessary in order to kill the roots. 

Mountain maple can withstand repeated and heavy 

browsing, producing new growth even when 80 

percent of the annual twig growth is removed each 

year (Sullivan 1993). Mountain maple is an 

understory or subcanopy component in a number 

of northeastern forest associations (Sullivan 1993). 

Cutting mountain maple near the ground level, 

clear-cutting, and spraying with 2,4-D in the 

spring will increase the availability of new growth 

for deer and moose browse (Sullivan 1993). 

Burning can be used to suppress mountain maple 

in order to promote reproduction of other, more 

desirable tree species. 

Benefits.—Mountain maple is a nutritious browse 

species for moose, white-tailed deer, cottontails, 

snowshoe hares, woodland caribou, beavers, and 

ruffed grouse (Sullivan 1993). Mountain maple is 

sometimes planted as an ornamental, but has little 

landscape value excluding its brief fall color 

display and colorful samaras. 

References

De Jong, P.C. 1976. Flowering and sex expression 

in Acer L. A biosystematic study. Mededelingen 

Landbouwhogeschool Wageningen 76 (2): 1-

202. 

 

Dirr, M.A. 1998. Manual of woody landscape 

plants, Their identification, ornamental 

characteristics, culture, propagation, and uses. 

Stipes Publishing, Champaign, IL. 1,187 p. 

Farr, D.F., G.F. Bills, G.P. Chamuris, and A.Y. 

Rossman. 1989. Fungi on plants and plant 

products in the Untied States. The American 

Phytopathological Society, St. Paul, MN. 

1,252 p. 

Gleason, H.A. and A. Cronquist. 1991. Manual of 

vascular plants of northeastern United States and 
29

adjacent Canada. The New York Botanical 

Garden, Bronx, NY. 910 p. 

Johnson, W.T. and H.H. Lyon. 1991. Insects that 

feed on trees and shrubs. Cornell University 

Press, Ithaca, NY. 560 p. 

Jones, R.K. and D.M. Benson. 2001. Diseases of 

woody ornamentals and trees in nurseries. The 

American Phytopathological Society, St. Paul, 

MN. 482 p. 

Sinclair, W.A., H.H. Lyon, and W.T. Johnson. 

1987. Diseases of trees and shrubs. Cornell 

University Press, Ithaca, NY. 575 p. 

Sullivan, J.R. 1983. Comparative reproductive 

biology of Acer pensylvanicum and A. spicatum 

(Aceraceae). American Journal of Botany 70(6): 

916-924. 

Sullivan, J. 1993. Acer spicatum. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/ 

database/feis/plants/shrub/acespi/index.html. 

14 p. 

USDA, NRCS. 2002. The PLANTS Database, 

Version 3.5 National Plant Data Center, Baton 

Rouge, LA. http://plants.usda.gov. [not paged]. 

van Gelderen, D.M., P.C. de Jong, and H.J. 

Oterdoom. 1994. Maples of the World. Timber 

Press, Portland, OR. 458 p. 

Zasada J.C. and T.F. Strong. 2000. Acer L. In: 

Woody plant seed manual. F.T. Bonner, tech. 

coord., R.G. Nisley, ed. U.S. Department of 

Agriculture, Forest Service, Washington, DC. 

http://www.wpsm.net/ Acer.pdf. [not paged]. 
30 

Acnistus arborescens (L.) Schlecht. gallinero 

SOLANACEAE 

Synonyms: Atropa arborescens L. 

 Cestrum cauliflorum Jacq. 

 Acnistus cauliflorus (Jacq.) Schott 

 Acnistus frutescens Bello 

 Cestrum macrostemon Sassé & Moç. 

 Dunalia arborescens (L.) Sleumer 

John K. Francis 

General Description.—Gallinero is also known as 

hollow heart, wild tobacco, siyou, bastard sirio, 

galán arbóreo, mata gallina, tabaco de monte, 

nigüito, marieneira, and tabak djab. It is an 

evergreen shrub or small tree to 6 m in height and 

15 cm in stem diameter. Multiple stems are usual 

for older plants. The trunk is supported by 

extensive lateral roots with “sinkers” and abundant 

fine roots. The roots are flexible, light tan, and 

have furrowed bark. Stem bark is light brown or 

gray and finely fissured. Stem wood is light brown 

and hard. Twigs of gallinero are stout, light brown 

or gray and finely hairy. The simple, alternate 

leaves are elliptical to lanciolate, 5 to 30 cm long 

by 3 to 14 cm broad, entire, and pointed at both 

ends. Gallinero bears tiny, fragrant, greenish-white 

flowers in clusters of 30 or more in axillary 

fascicles. The fruits (berries) are numerous in each 

cluster, yellow or orange, globose, 5 to 11 mm in 

diameter, and have a persistent calyx. The fruit 

pulp is juicy, almost tasteless or slightly bitter, and 

not edible to humans. Each fruit contains numerous 

yellow, flattened seeds 1.5 to 2 mm wide with a 

rolled embryo (Howard 1989, Liogier 1995, Little 

and others 1974, Stevens and others 2001). 

Range.—Gallinero is native to the Greater 

Antilles, the Lesser Antilles, Trinidad and Tobago, 

Mexico, Central America, and South America 

through Brazil and Peru (Howard 1989, Little and 

others 1974). 

Ecology.—Gallinero is widespread and rare to 

common on a wide range of soils over both igneous 

and sedimentary rocks. In Puerto Rico, it grows in 

areas that receive from about 1600 to 2500 mm of 

mean annual precipitation. The species is most 

common from 700 to 1,500 m in elevation in Costa 

Rica (Haber 2002) and from 450 to 900 m in 

Puerto Rico (Little and others 1974). The species is 

frequently found on roadsides, landslides, old 

pastures, and young secondary forests (Haber 

2002). Disturbance is necessary for establishment. 

Gallinero is intolerant of shade; at least broken 

sunlight is necessary for long-term survival and 

flowering. A number of insect species feed on the 

leaves and parasitize the fruits (Engriser 1995, 

Haber 2002, Simpson and others 2002). 

Reproduction.—Gallinero flowers in May and 

June and fruits from May to September in Costa 

Rica. Occasional individuals may be found in 

flower at any time during the year (Haber 2002). In 

Puerto Rico, the species flowers in spring and fruits 

in late spring and summer (Little and others 1974). 

The flowers are pollinated by bees, wasps, flies, 

butterflies, beetles, and occasionally hummingbirds 

(Haber 2002). A collection of fruits from Puerto 
31

Rico averaged 0.2025 + 0.0084 g/fruit. Seeds 

separated from them weighed an average of 

0.00074 + 0.00017 g/seed or 1,350,000 seeds/kg. 

Sown on commercial potting mix, 88 percent 

germinated between 11 and 35 days after sowing. 

Some of the seedlings were killed by “damping 

off” fungi (species unknown). A fruiting tree 

produces an enormous number of seeds. Birds and 

bats eat the fruits and disperse the seeds (Engriser 

1995). Layering (rooting) occurs whenever the 

trunk or branches come in contact with the ground; 

sprouts from prostrate trunks and branches 

sometimes become independent plants. 

Growth and Management.—Gallinero has a 

moderate growth rate and is relatively short-lived 

(10 to 20 years). Nursery and management 

experience has not been published. 

Benefits.—Gallinero is used occasionally as an 

ornamental and for hedges and living fence posts. 

Chickens eat the fruits (Little and others 1974). 

The wood is burned for fuel (Espinoza 1985). 

Practitioners of traditional herbal medicine use 

extracts of leaves to treat cancerous growths. An 

alcoholic extract was tested and found to inhibit 

cancerous cells in vitro and in vivo. The active 

ingredients were identified as Withaferin A and 

Withacnistin (Kupchan and others 1969). Extracts 

of gallinero were the most effective against 

hamster lung fibroblasts of 31 Columbian plants 

used in herbal medicine to treat cancer (Lopez de 

Cerain and others 1996). The species is used in 

Brazil as a diuretic, to treat liver infections, as well 

as an antitumor agent (Mingussi and Barata 2002). 

Gallinero helps reforest disturbed areas, protects 

the soil, and furnishes food and cover for wildlife. 

References 

Engriser, E.M. 1995. The effect of insect larvae 

infestation on fruit choice in phyllostomid fruit 

bats: an experimental study. Biotropica 27(4): 

523-525. 

Espinoza, P.L. 1985. Importance of the tree 

component in agroforestry-based coffee 

cultivation, with examples from Costa Rica. 

Thesis. Institut fur Waldbau der Tropen und 

Naturwaldforschung, University of Gottingen, 

Germany. 164 p. 

Haber, W.A. 2002. Acnistus arborescens (L.) 

Schltdl., Solanacear-Potatoe Family. 

http://www.cs.umb.edu/~whaber/Monte/Plant/ 

Sola/Acn-arb. htm. 2 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Kupchan, S.M., W.K. Anderson, P. Bollinger, 

R.W. Doskotch, R.M. Smith, J.A. SaenzRenauld, H.K. Schnoes, A.L. Burlingaure, and 

D.H. Smith. 1969. Tumor inhibitors. XXXIX. 

Active principles of Acnistus arborescens. 

Isolation and structural and spectral studies of 

Withaferin A and Withacnistin. Journal of 

Organic Chemistry 34(12): 3,858-3,866. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de 

la Universidad de Puerto Rico, San Juan, PR. 

617 p. 

 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Lopez de Cerain, A., R. Pinzón, J. Calle, A. Marín, 

and A. Monge. 1996. Cytotoxic activities of 

Colombian plant extracts on Chinese hamster 

lung fibroblasts. Phytotherapy Research 10(5): 

431-432. 

Minguzzi, S. and L.E.S. Barata. 2002 Vitanolídeos 

anti-câncer de Acnistus arborescens. 

http://www.sbq.org.br/ranteriores/ 

23/resumos/0398/ 2 p. 

Simpson, S.E., H.N. Nigg, and J.L. Knapp. 2002. 

Host plants of diaprepes root weevil and their 

implications to the regulatory process. 

http://www.fcprac.ifas.ufl.edu/citrustopics/pest

%20control/Diaprepes/Diaprepes%20Procee. 

18 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanical Garden, St. Louis, MO. 

p. 1,911-2,666. 
32 

Aegiphila martinicensis Jacq. Caribbean spiritweed 

VERBENACEAE 

Synonyms: Aegiphila glandulifera Moldenke 

 Aegiphila panamensis Moldenke 

John K. Francis 

General Description.—Caribbean spiritweed, 

also known as capaíllo, bois cabrit, bois de fer, 

bwa kabuit, sureau gros, and bois cabroit, is an 

upright to climbing, evergreen shrub or small tree 

up to 6 m in height and 7.5 cm in stem diameter. 

The older stems are gray and furrowed, branches 

are mostly smooth with lenticels, and twigs are 

four-angled. The ivory-colored wood is 

moderately hard and brittle, and the stems have a 

2-mm pith. Roots are tan colored and flexible. The 

opposite, thin leaves are oblong-lanceolate to 

oblong with entire edges, 8 to 16 cm long by 3 to 8 

cm broad, rounded or pointed at the base and longpointed at the tip. Inflorescences are axillary or 

terminal, many-flowered cymes. Flowers are white 

or pale yellow, tubular, and 2 to 6 mm long. From 

these develop globose to ovoid, yellow, orange, or 

red, 7- to 10-mm, fleshy fruits with persistent 

calyxes, a grainy texture, and little flavor. Fruits 

have one to four brown, elliptical seeds with 

longitudinal striations (author’s observations, 

Howard 1989, Liogier 1995, Little and others 

1974). 

Range.—Caribbean spiritweed is native to Cuba, 

Jamaica, Puerto Rico, the Virgin Islands, the 

Lesser Antilles, Trinidad, Mexico to Panama, 

Colombia, Venezuela, and French Guiana 

(Howard 1989, Liogier 1995, Little and others 

1974, Missouri Botanic Garden 2003). It is not 

known to have been planted or naturalized 

elsewhere. Stevens and others (2001) treat the 

Central American distribution as a separate species 

(A. panamensis Moldenke) based on the shape of 

the calyx. 

Ecology.—Caribbean spiritweed is rare in Puerto 

Rico (Little and others 1974) but is common in 

Nicaragua (Stevens and others 2001). It occurs in 

forests and brushy areas in foothills and lower 

mountains of Puerto Rico from 60 to 244 m in 

elevation in areas of limestone, other sedimentary, 

and igneous rocks (Little and others 1974). In 

Nicaragua, the species grows in forests and 

pastures in wet to dry areas from near sea level to 

900 m elevation (Stevens and others 2001). 

Caribbean spiritweed is moderately intolerant of 

shade and is able to flower and fruit in partial 

shade. 

Reproduction.—Caribbean spiritweed flowers 

and fruits irregularly throughout the year (Little 

and others 1974). The fruits ripen a few at a time 

in each infrutescence rather than all at once. A 

collection of fresh fruits from Puerto Rico weighed 

an average of 0.496 + 0.026 g/fruit. Air-dried seed 

separated from them averaged 0.0331 + 0.0007 

g/seed or 30,000 seeds/kg. Placed on moist blotter 

paper without pretreatment, 61 percent germinated 

between 1 and 4.5 months after sowing (author’s 

observation). Germination was hypogeal 

(cotyledons remain below ground). Seeds are 

probably dispersed by birds. Caribbean spiritweed 

sprouts when cut and may resprout to replace 

senescent stems. 

Growth and Management.—Caribbean 

spiritweed grows relatively rapidly, about 1 
33

m/year. Stems may live 5 years or more. No 

planting or management experience has been 

published. 

 

Benefits.—Carabbean spiritweed helps protect the 

soil and furnishes food and cover for wildlife. 

Larva of the hawk moth, Munduca hannibal 

hannibal, apparently feed exclusively on 

Caribbean spiritweed (Oehlke 2003). Infusions of 

the leaves and twigs have been used as a diuretic 

and as a pleasant tonic for treating asthma (NúñezMelédez 1982). In Costa Rica, the plant is 

considered an aphrodisiac (Duke 2003). 

References

Duke, J. 2003. Dr. Duke’s phytochemical and 

ethnobotanical databases: Tico ethnobotanical 

dictionary. http://www.ars-grin.gov/duke/ 

dictionary/tico/a.html. [not paged]. 

Howard, R.A. 1989. Flora of the Lesser Antilles. 

Vol. 6. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p.

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Missouri Botanic Garden. 2003. Flora of Panama 

checklist: Aegiphila martinicensis Jacq. http:// 

mobot.mobot.org/cgi-bin/search_pick. 1 p. 

Núñez-Melédez, E. 1982. Plantas medicinales de 

Puerto Rico. Editorial de la Universidad de 

Puerto Rico, Río Piedras, PR. 498 p. 

Oehlke, B. 2003. Manduca hannibal hannibal. 

http://www.silkmoths.bizland.com/mhannhan. 

htm. 4p. 

Stevens, R.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanic Garden, St. Louis, MO. 

p. 911-2,666. 
34 

Agave missionum Trel. corita 

AGAVACEAE 

Synonyms: Agave portoricensis Trel. 

John K. Francis 

General Description.—Corita, also known as 

maguey, is a succulent shrub with woody parts that 

is 1.0 to 1.5 m in height in its vegetative stage and 

3 to 6 m when in flower. The plant is considered a 

shrub because it is of shrub size and has fibrouswoody tissue at the base of the leaf whorl, woody 

thorns, and a woody flower stalk. The numerous 

fibrous roots are about 5 mm in diameter and the 

fibrous-woody base of the plant extends about 15 

cm below the surface. Lanciolate to broadly 

lanciolate, fleshy leaves are blue-green in color, 

contain many fine, high-quality fibers running their 

length, and thorns, 2 to 6 mm in length along their 

edges, and larger ones, 10 to 20 mm long, at the 

tips. The leaves die when the plant is about 

midway through flowering. The flower stalks 

(panicles) are about 10 cm in basal diameter with 

soft wood and a pithy center. The yellow or 

yellow-green perianth is six-parted and funnel 

shaped, 4.5 to 5.2 cm long. The capsules are 

elliptical or globose and 2.5 to 3.8 cm long and 

contains numerous seeds, 6 to 8.5 mm long and 5 

to 6 mm broad. If the flowers abort, numerous 

bulbils are formed on the pedicels (author’s 

observation, Britton and Wilson 1923). 

Range.—Corita is native to Puerto Rico and the 

offshore islands of Culebra and Vieques and St. 

Thomas, St. Croix, St. John, Tortola, Virgin Gorda 

and Anegada in the U.S. and British Virgin Islands 

(Breckon and García 2001, Britton and Wilson 

1923, University of the Virgin Islands 

Conservation Data Center 2002). Based on work 

by Trelease (1913), some sources (International 

Plant Name Index 2002, Missouri Botanical 

Garden 2002, Texas A & M University 

Bioinformatics Working Group 2002) separate the 

group into two species—A. missionum and A. 

portoricensis Trel. However, Britton and Wilson 

(1923) and Liogier and Martorell (2000) recognize 

just A. missionum. Because ecological and 

management differences between the two 

populations are probably minor, they will be 

treated here as a single species. 

Ecology.—Corita grow naturally in well-drained to 

excessively drained areas that receive between 750 

mm and about 1700 mm of annual precipitation. 

The species colonizes most soil types over both 

sedimentary and igneous rock formations. Corita is 

intolerant of shade and consequently does not grow 

well under a closed forest canopy and does not 

compete well in tall grass swards. It is often found 

in disturbed areas, cliffs, and rocky terrain on the 

coasts and inland. Corita plants are eaten by cattle 

when they are small but are generally avoided after 

the thorns harden. 

Reproduction.—Flowering of corita appears to be 

triggered more by size and vigor than by age. 

Probably, reserves of starch and other nutrients are 

important. Flowering plants are often about 1 m or 

more in height (tallest leaf), 2 m in diameter, and 5 

to 10 or more years old, depending on the quality 

of the habitat. Flowering proceeds from the bottom 

of the inflorescence to the top. Although honeybees 

and other insects visit the flowers in great numbers, 
35

most plants never produce seeds. After the flowers 

fall off, bulbils form at the point where the flowers 

were attached. Corita plants produce several 

hundred to several thousand bulbils each. A 

collection of 100 bulbils from Puerto Rico weighed 

(fresh) an average of 3.003 + 0.252 g/bulbil and 

ranged from 0.18 to 13.63 g. Placed with their 

bases slightly buried in moist potting mix, all of 

them rooted within 1 week. They grew rapidly 

under partial shade and were pricked into 

containers in about 3 weeks from sowing. After 2 

or 3 weeks they were moved to full sun and 

reached about 15 cm in height in 2 additional 

months and were dispersed to the public (author’s 

observation). Plants originating from bulbils can be 

numerous under fruiting plants. It is not known 

what specialized means of dispersal exist. 

However, new plants are frequently seen with no 

remains of flowering corita plants nearby. 

Normally a plant dies after completing flowering, 

but many plants produce basal sprouts in the year 

of flowering, or before, that continue for another 

generation. 

Growth and Management.—Corita grows at a 

steady, moderate rate from bulbils on well-watered 

soil and somewhat more slowly in poor, droughty 

ground. Growth and development of seedlings has 

not been reported. Transplanting is easy with little 

mortality. Because of low competition, disturbed, 

dry, and rocky areas are probably the best sites for 

planting. Corita is recommended for planting on 

shorelines and dunes in the Caribbean (United 

Nations Environment Program 1998). 

Benefits.—Corita contributes to the biodiversity 

and helps protect the soil in the sites where it 

grows. Because of its hardiness in almost any welldrained soil, moderate size, and pleasing dark bluegreen color, the species makes an excellent 

ornamental. Because exotic agaves were promoted, 

it was used sparingly in the past but is beginning to 

attract more attention. Corita appears to be an 

excellent honey source. The leaves contain strong 

fibers that probably were used for cordage in 

former times. 

References 

Breckon, G.J. and R.G. García. 2001. Vascular 

plants of Susúa Forest. http://www.uprm. 

edu/biology/profs.breckon/herbarium/florasusua.

htm. 52 p. 

Britton, N.L. and P. Wilson. 1923. Botany of Porto 

Rico and the Virgin Islands, descriptive floraSpermatophyta. Scientific Survey of Porto Rico 

and the Virgin Islands Vol. 5, Part 1. New York 

Academy of Sciences, New York. 626 p. 

International Plant Name Index. 2002. 

International Plant Name Index Query. 

http://www.ipni.org/ipni/query_ipni.html. 1 p. 

Liogier, H.A. and L.F. Martorell. 2000. Flora of 

Puerto Rico and adjacent islands, a systematic 

synopsis. 2nd ed. Editorial de La Universidad de 

Puerto Rico. 382 p. 

Missouri Botanical Garden. 2002. W3

 tropicos. 

http://mobot.mobot.org/cgi-bin/search_vast. 1 p. 

Texas A&M University Bioinformatics Working 

Group. 2002. Agavaceae (the century-plant 

family). http://www.csdl.tamu.edu/FLORA/ 

bonapfams/bonxxaga.htm. [not paged]. 

Trelease, W. 1913. Collections in Puerto Rico and 

St. Thomas. Memoirs of the National Academy 

of Sciences 11: 37. 

United Nations Environment Program. 1998. 

Manual for sand dune management in the wider 

Caribbean. http://www.cep.unep.org/issues/ 

sanddunes.pdf. 73 p. 

University of the Virgin Islands Conservation Data 

Center. 2002. U.S.V.I. vegetation classification 

system. http://cdc.uvi.edu/REAWEB/ 

vegbody.html. [not paged]. 
36 

Allamanda cathartica L. allamanda 

APOCYNACEAE 

Synonyms: Echites verticillata Sessé & Moç. 

 Orelia grandiflora Aublet 

 Allamanda grandiflora (Aublet) Poiret in Lam. 

 Allamanda hendersonii W. Bull ex Dombrain 

John K. Francis 

Illustration source: Missouri Botanical Garden 

General Description.—Allamanda, also known as 

angel’s trumpet, golden trumpet, yellow bell, 

canario, cautiva, dabel-da-dama, orelia, and liane à 

lait, is an evergreen, vine-like woody shrub. It may 

reach a free-standing height of 2 m and an 

extension of 5 m or more. The species also climbs 

a few meters into the crowns of tall brush and low 

trees. Older plants often have multiple stems from 

the root crown and long stems with relatively few 

branches. Bark of lower stems is brown and 

furrowed. Twigs are green or yellow green. Stems 

and twigs exude a milky sap when cut. The 

leathery, yellow-green to dark green leaves grow in 

whorls of three or four, or are sometimes opposite. 

Leaves are 6 to 16 cm long, obovate to oblonglanceolate, pointed at both ends and have entire 

margins and short petioles. Inflorescences are fewflowered, axillary cymes that grow near the ends of 

branches. The bright yellow flowers are 5 to 7.5 cm 

across. Flowers of cultivated varieties are often 

larger and may be colored white, cream, pink, or 

orange. Capsules, which rarely occur in cultivated 

varieties, are subglobose, 4 to 6 cm in diameter, 

and densely prickled. They contain many tan, 

flattened, winged seeds. There are 2n = 18 

chromosomes (Howard 1989, Liogier 1995, Long 

and Lakela 1976). 

Range.—Allamanda is apparently native to 

northern Brazil, Guyana, Surinam, and probably 

French Guiana (Liogier 1995, Pacific Island 

Ecosystems at Risk 2002, Tropilab Inc. 2002). The 

species has been planted and has become 

naturalized throughout the tropics (Howard 1989). 

Ecology.—Wild and naturalized allamanda grows 

on riverbanks in Suriname (Tropilab Inc. 2002), on 

disturbed sites in Florida (Long and Lakela 1976), 

along roads (Liogier 1995), and on abandoned 

farms, house places, and around clandestine dumps 

in Puerto Rico (author’s observation). The species 

is intolerant to intermediate in tolerance to shade. It 

can grow below brush and a thin forest canopy, 

usually without blooming, but blooms and makes 

its greatest growth in full sun. Allamanda grows 

best in well drained, moist, sandy soils rich in 

organic matter (Barcellos 2002). It does not 

tolerate salty soils, highly alkaline conditions, and 

is killed by temperatures of -1 °C (Floridata 2002, 

Tropilab Inc. 2002). In Nicaragua, the species 

occurs between 0 and 700 m in elevation (Stevens 

and others 2001). Allamanda has naturalized in 

Puerto Rico in areas that receive between about 

1000 and 2800 mm of mean annual precipitation. 

Reproduction.—Allamanda blooms all year in 

most habitats. Because capsules and seed are rarely 

produced by cultivated varieties, naturalization is 

usually by vegetative means. In Puerto Rico, the 

species has been planted widely, persists 

tenaciously, and spreads by layering as the vines 
37

extend. In addition, trimmings dumped in vacant 

lots and wildlands root readily and start new 

colonies. Plants coppice vigorously when cut. The 

species is invasive in Queensland, Australia 

(Pacific Island Ecosystems at Risk 2002). 

Growth and Management.—Established 

allamanda grows rapidly, adding 1 to 3 m to 

extended length per year. Pruning is necessary to 

keep it under control in most landscaping 

applications. There appears to be no reason to 

establish it in wildlands, but the shrub could easily 

be planted with cuttings placed directly in the soil. 

Getting rid of spot infestations can be difficult. 

Cutting is ineffective. For grubbing to work, 

removal of the plant stems and roots from the site 

must be complete. Control with herbicides may 

overcome some of these difficulties but is untested. 

Benefits and Detriments.—In wild stands, 

allamanda protects the soil and furnishes cover for 

wildlife. Year-long production of large, beautiful 

flowers endears allamanda to gardeners and 

landscapers throughout the tropics and subtropics. 

The species is particularly useful because it will 

grow in most areas with relatively little care. It is 

used as ground cover, for hedges and screens, and 

as upright shrubs. Because of its rapid growth, 

pruning is often necessary, which can expose 

gardeners to the toxic sap that causes dermatitis 

symptoms of rash, blisters, and itch. Although 

incidence is much less common, plant parts are 

also toxic if ingested. All parts contain the toxic 

iridoid lactone, allamandin (Ecology and 

Evolutionary Biology Conservatory 2002). In 

herbal medicine, teas prepared from leaves and 

roots are used as a strong purgative that must be 

used with caution (Liogier 1990). 

References 

Barcellos, D.C. 2002. Plantas ornamentais tóxicas: 

Allamanda cathartica. http://www. 

plantastoxicas.hpg.ig.com.br/toxicas/allcat.htm. 

2 p. 

Ecology and Evolutionary Biology Conservatory. 

2002. Allamanda cathartica L. University of 

Connecticut. http://florawww.eeb.uconn.edu/ 

acc_num/198500393.html. 3 p. 

Floridata. 2002. Allamanda cathartica. 

http://www.floridata.com/ref/a/alla_cat.cfm. 3 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Dicotyledoneae. 

Part 3. Vol. 6. Arnold Arboretum, Harvard 

University, Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc. San Juan, PR. 566 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p.

Long, R.W. and O. Lakela. 1976. A Flora of 

Tropical Florida. Banyan Books, Miami, FL. 

962 p. 

Pacific Island Ecosystems at Risk. 2002. 

Allamanda cathartica L., Apocynaceae. 

http://hear.org/pier_v3.3/alcat.htm. 2p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

1. Missouri Botanic Garden Press, St. Louis, 

MO. 943 p. 

Tropilab Inc. 2002. Allamanda cathartica L. 

http://www.tropilab.com/allamanda.thml. 2 p. 
38 

Amyris elemifera L. torchwood 

RUTACEAE 

Synonyms: Amyris maritima Jacq. 

 Amyris sylvatica Jacq. 

 Elemifera maritima (Jacq.) O. Kuntze 

John K. Francis 

General Description.—Torchwood, also known as 

candlewood, sea amyris, tea, cuabilla, bois 

chandelle, bois flambeau, and bois pini, is an 

evergreen shrub or small tree reaching maximum 

heights of 4 to 13 m, depending on the 

environment. It usually has a single stem emerging 

from the ground, unless damaged in the past but 

may begin branching near the ground when there is 

plenty of sunlight. The trunk is smooth and gray, 

becoming rough with furrows and plates when old. 

The plants usually have a weak taproot, strong, 

stiff laterals, and abundant fine roots. Torchwood 

has a vertical branching habit. It may have a dense, 

low crown in open-grown individuals. The twigs 

are fine and yellowish-gray, becoming gray with 

age. The foliage is pendulous and aromatic, palegreen or pale-green with blue-green highlights. The 

leaves are opposite or subopposite with three 

(sometimes five) leaflets on a 3-cm petiole. The 

blades are ovate, rhombic ovate, or lanciolate, with 

a pointed or long-pointed tip. The terminal or axial 

paniculate inflorescences contain many tiny 

flowers. The fruits are black globose drupes that 

are aromatic and contain one brown seed (Howard 

1988, Liogier 1988, Little and Wadsworth 1964). 

Range.—Torchwood is native to Florida, the 

Bahamas, the Greater Antilles, the Lesser Antilles, 

Trinidad, Guatemala, Belize, Honduras, and El 

Salvador (Liogier 1988, Little and Wadsworth 

1964). 

Ecology.—Torchwood is moderately intolerant of 

shade. Young plants compete well with shrubs and 

small trees, but do not survive in dense grass 

swards. In dry forests, understory plants linger for 

decades until openings in the overstory allow them 

to reach intermediate and codominant positions to 

flower and fruit. The species may be found in both 

remnant and secondary forests. In Puerto Rico, 

torchwood grows in areas receiving from 750 to 

over 2000 mm of mean annual precipitation. In the 

upper end of the rainfall range, the species is 

confined to excessively drained sites. Elevation 

ranges from near sea level to about 600 m. It 

tolerates a wide variety of soil textures, fertility 

levels, and pH’s. Torchwood may be found in areas 

underlain by sedimentary rocks including 

limestone, igneous, and metamorphic including 

ultramafic rocks, and coastal sands. In the 

Bahamas, torchwood grows in rocky and sandy 

coastal thickets (Britton and Millspaugh 1962); in 

Florida, it grows in coastal areas and on rich 

hammocks farther inland (4-H Youth Development 

2001). 

Reproduction.—In Puerto Rico, torchwood 

flowers irregularly through the year (Little and 

Wadsworth 1964). Flowers are reported to appear 

over several months in the fall in Florida (West and 

Arnold 1952). Nursery plants in pots in Puerto 

Rico bloomed at 2 years of age. Seeds cleaned 

from ripe fruits and sown without any pretreatment 

germinate normally. Birds are the principal seed 

disbursers. 

Growth and Management.—Nursery plants in 

pots reached 1.8 m in 2 years. However, older trees 
39

in the dry forests that torchwood inhabits grow 

slowly and may live for 50 years or more (author’s 

observation). A release cut or thinning should be 

effective if it were necessary to improve the 

position of torchwood in a stand. 

Benefits.—The wood of torchwood is used for 

fence posts today but was formerly used for fuel, 

furniture, and torches--hence the name torchwood. 

In a test of untreated service life of posts, 20 of 20 

posts were still sound after 13 years in the ground 

(Chudnoff and Goytia 1972). If the wood was more 

abundant and in larger pieces, it would be in 

demand for lumber because it is fragrant, finegrained, strong, durable, and repellent to dry-wood 

termites (Little and Wadsworth 1964). Torchwood 

is valued as a honey plant in the Dominican 

Republic (Marcano Fondeur 1973). Birds eat the 

fruits, but there usually are many other more 

attractive foods present (West and Arnold 1952). In 

Florida, torchwood is valued as a larval food for 

the rare and endangered Schaus’s swallowtail 

butterfly, Papilio aristodemus ponceanus (Hammer 

2001). Key deer (Odocollous virginanus clavium) 

will not eat the foliage (Schaus and others 2001). 

However, feral goats eat at least the bark, which 

has caused torchwood to decrease in abundance in 

Mona Island, Puerto Rico (Little and Wadsworth 

1964). Taxaline, an oxazole extracted from 

torchwood, had the most effective antibacterial 

activity of several bioactive chemicals from West 

Indian plants against Mycobacterium tuberculosis 

and two other Mycobacterium species (Rastogi and 

others 1998). Root and resin extracts from 

torchwood and others of the genus are used as an 

expectorant (Liogier 1990). 

References 

4-H Youth Development. 2001. Florida forest 

trees: torchwood (Amyris elemifera). 

http://www.sfrc.ufl.edu/4h/Torchwood/ 

torchwoo.htm. 2 p. 

Britton, N.L. and C.F. Millspaugh. 1962. The 

Bahama flora. The New York Botanical Garden, 

New York. 695 p. 

Chudnoff, M. and E. Goytia. 1972. Preservative 

treatments and service life of fence posts in 

Puerto Rico (1972 progress report). Research 

Paper ITF-12. U.S. Department of Agriculture, 

Forest Service, Institute of Tropical Forestry, 

Río Piedras, PR. 28 p. 

Hammer, R.L. 2001. Attracting birds to your 

garden. Fairchield Tropical Garden. 

http://www.ftg.org/horticulture/habitat 

plants.html. 4 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, part 1. Vol. 4. Arnold 

Arboretum, Harvard University. Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puetro 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

Marcano Fondeur, E. de J. 1973. La Flora Apícola 

de la República Dominicana. http.//marcano. 

Freeservers.com/nature/estudios/apicola/dicotsp.

html. 11 p. 

Rastogi, N, J. Abaul, G-K. Seng, A. Devallois, E. 

Philogene, P. Bourgeois, and K.S Goh. 1998. 

Antimycobacterial activity of chemically 

defined natural substances from the Caribbean 

flora of Guadeloupe. FEMS-Immunology and 

Medical Microbiology 20(4): 267-273. 

Schaus, C., S. Wade, and J. Dunan. 2001. Key 

deer and plants they won’t eat. 

http://monroe.ifas.ufl.edu/key_deer_plants.htm. 

3 p. 

West, E. and L.E. Arnold. 1952. The native trees 

of Florida. University of Florida Press, 

Gainesville, FL. 212 p. 
40 

Aralia spinosa L. devil’s walking stick 

ARALIACEAE 

 

Synonyms: Aralia leroana K. Koch. 

Kristina Connor 

Illustration source: U.S. Department of Agriculture [not 

dated] 

General Description.—Devil’s walking stick, also 

known as angelica tree, American angelica-tree, 

Hercules’ club, pigeon tree, pick tree, prickly ash, 

prickly elder, toothache bush, toothache tree, and 

shotbush, is a large, coarse-textured shrub or small 

tree, ranging from 6 to 10 m in height. The sturdy, 

ash gray to brown stems have dense, stout prickles, 

and diameters to 15 cm are not uncommon (U.S. 

Department of Agriculture [no date]). Young stems 

are mostly unbranched with leaves clustered near 

the top, producing a flat-topped, widely spreading 

crown. Stems tend to remain unbranched until the 

first terminal inflorescences are produced (Sullivan 

1992). Twigs are similar in color and armament to 

the stems and have a large pith (Synor and Cowen 

[no date]). Leaf scars are distinctive, lined with 

spines, and can extend half way around the twigs. 

Leaves are alternate, bi- or tri-pinnately compound, 

and 0.9 to 1.8 m in length. The rachis is prickly and 

swollen at nodes (Bailey and Bailey 1976). The 

rachis and rachilla may be the functional equivalent 

of branches in this species (Briand and others 

1998). The dark green, glaucous leaflets are sessile, 

5.1 to 10.2 cms long and 3.0 to 4.1 cm wide, 

mostly paired, ovate, serrate, with pointed apex and 

rounded base (Dirr 2002, Johnson and Hoagland 

1999, Krüssmann 1986). There are prickles on the 

veins and lower surfaces; they appear to be 

outgrowths of epidermal and parenchyma cells 

(Briand and Soros 2002). The wood is lightweight, 

brown but streaked with yellow (Synor and Cowen 

[no date]). It may be that the pithy nature of the 

trunk of Devil’s walking stick allows for rapid 

growth but ultimately restricts crown architecture 

and maximum size attainable (Briand and others 

1999). The species is deciduous and spreads by 

underground runners. 

Range.—Devil’s walking stick is found in the 

Eastern United States, from Pennsylvania south to 

Florida and west to Texas and southwestern Iowa. 

It has escaped from cultivation and thus can be 

found in New England, southern Ontario, 

Michigan, Wisconsin, Oregon, Washington, and 

western Europe (Sullivan 1992). It grows in 

regions where temperatures fall to –29 o

C; such 

severe freezes can kill stems back to ground level 

(Godfrey 1988, Frett and MacKenzie 1999, 

Sullivan 1992, Scheper 2002). 

Ecology.—This highly adaptable species grows 

best in well drained soils of low and moist 

woodlands but can be found growing in rocky, dry 

or clay soils and under a range of pH conditions 

(Frett and MacKenzie 1999). It grows luxuriantly 

on good sites, but plants may live longer and be 

sturdier on poorer sites (Scheper 2002). It has a 

fairly high heat tolerance. According to Russell 

(1997), it is shade tolerant, but Sullivan (1992) 

reports that it will decline if the overstory becomes 

thick, a common characteristic of moderately 

intolerant species. It has no serious pests, but 

margined blister beetles may defoliate plants early 

in the fall (Scheper 2002). 

Reproduction.—Devil’s walking stick spreads 

extensively by vegetative reproduction from 

underground rhizomes. The first terminal 

inflorescences usually occur when plants are, on 

the average, 3.5 years old (Sullivan 1992). The 
41

white to cream colored flowers are small, perfect, 

numerous, and occur in large clusters (umbels) in 

mid to late summer, depending on geographic area. 

Panicles of flowers may be 30 to 46 cm long. Each 

flower, composed of five sepals, five reflexed 

petals, and five stamens, is borne on a hairy stalk. 

The ovary is inferior, and there are usually five 

styles, united at the base (Brickell and Zuk 1996, 

U.S. Department of Agriculture [no date]). The 

purple-black fruits, maturing in late fall, are 

produced in large quantities on pink-red stems. 

They are juicy, ovoid drupes up to 6.4 mm long 

with three to five seed-like stones. The species can 

be propagated from seed sown outdoors in fall 

(Russell 1997) or from suckers taken in late winter 

and root cuttings taken in late fall and overwintered 

upside down in sand (Scheper 2002). Stored seeds 

require 3 to 5 months stratification, followed by 1 

to 4 months at 20 o

C for germination. 

Growth and Management.Devil’s walking stick 

can be maintained by mowing, cutting, and 

burning, and will commonly form dense thickets 

by resprouting after such disturbances. When the 

overstory becomes too dense, the species will 

decline. Defoliation by gypsy moth in 

Pennsylvania and Maryland has increased the 

presence of Devil’s walking stick by thinning the 

overstory (Sullivan 1992). The species can be 

controlled by injecting stems larger than 2 inches 

in diameter with herbicide (Loftis 1978). 

Benefits.—The flowers provide a pollen and nectar 

source to honeybees and a variety of other insects. 

The berries are edible to wildlife, a favorite of 

cedar waxwings and other birds, as well as other 

frugivores and omnivores, including the black bear. 

Deer browsing has been reported (Krüssmann 

1986, Sullivan 1992). The bark, roots, and berries 

have also been used for medicinal purposes by both 

Native Americans and early settlers. While various 

plant parts, extracted in alcohol or water, have been 

used to treat boils, fever, toothache, eye problems, 

skin conditions, and snakebite, the raw berries can 

be mildly toxic to humans if ingested, contact with 

the bark or roots can cause a brief skin irritation, 

and ingesting the fresh bark will cause vomiting 

(Felter and Lloyd 1898, Grieve 1971). The species 

is cultivated as an ornamental plant, the lacy 

appearance of the foliage, along with the prolific 

flowers and fruit making it an interesting 

landscape plant (Dirr 2002, Odenwald and others 

1996). It is also an excellent barrier plant (Frett 

and MacKenzie 1999, Michigan State University 

Extension 1999) and is good for urban use. 

However, it will send up shoots from its rhizomes 

and can be somewhat invasive. 

References 

Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: 

A Concise Dictionary of Plants Cultivated in the 

United States and Canada. McMillan Publishing 

Co., Inc. New York. 1,290 p. 

Briand, C.H., S.M. Campion, D.A. Dzambo, and 

K.A. Wilson. 1999. Biochemical properties of the 

trunk of the Devil’s walking stick (Aralia 

spinosa; Araliaceae) during the crown-building 

phase: Implications for tree architecture. 

American Journal of Botany 86(12): 1677-1682. 

Briand, C.H., A.D. Daniel, K.A. Wilson, and H.E. 

Woods. 1998. Allometry of axis length, diameter, 

and taper in the Devil’s walking stick (Aralia 

spinosa; Araliaceae). American Journal of Botany 

85(9): 1201-1206. 

Briand, C.H. and C. Soros. 2002. Prickle 

distribution in Aralia spinosa. 

http://henson1.salisbury.edu/~biology/ 

Briand/research/research.html. 5 p. 

Brickell, C. and J.D. Zuk. 1996. The American 

Horticultural Society A-Z Encyclopedia of 

Garden Plants. DK Publishing, Inc., New York. 

1,092 p. 

Dirr, M.A. 2002. Aralia spinosa – Devil’swalkingstick or Hercules-club. Noble Plants. 

Plant information webpage, 

http://www.nobleplants.com/Classnotes/ 

fall/fallprofiles/aralia.htm 

Felter, H.W. and J.U. Lloyd. 1898. Aralia 

spinosa.-Prickly Elder. King’s American 

Dispensatory. 18th ed., 3rd revision, 2 vol. Ohio 

Valley Co., Cincinnati, OH. 2,172 p. + index. 

Frett, J. and B. MacKenzie. 1999. Aralia spinosa. 

The University of Delaware Botanical Garden. 

Species descriptions. http://bluehen.ags.udel.edu. 

1 p. 

Godfrey, R.K. 1988. Trees, shrubs, and woody 

vines of northern Florida and adjacent Georgia 

and Alabama. The University of Georgia Press, 

Athens, GA. 734 p. 

Grieve, M. 1971. A Modern Herbal. The 

medicinal. culinary, cosmetic and economic 
42 

properties, collection and folk-lore of herbs, 

grasses, fungi, shrubs and trees with their modern 

scientific uses. Dover Publications Inc., New 

York. 902 p. 

Johnson, F.L. and B.W. Hoagland. 1999. Aralia 

spinosa. Catalog of the Woody Plants of 

Oklahoma, Oklahoma Biological Survey 

http://www. biosurvey.ou.edu. 1 p. 

Krüssmann, G. 1986. Manual of Cultivated BroadLeaved Trees and Shrubs. Volume I, A-D. 

Timber Press, Beaverton, OR. 448 p. 

Loftis, D.L. 1978. Preharvest herbicide control of 

undesirable vegetation in southern Appalachian 

hardwoods. Southern Journal of Applied Forestry 

2(2): 51-54. 

Michigan State University Extension. 1999. Aralia 

spinosa – Devil’s Walkingstick. Ornamental Plants 

plus Version 3.0. 

http://www.msue.msu.edu/msue/imp/ modzz/ 

00001941.html. 1 p. 

Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope. 

1996. Plants for American Landscapes. Louisiana 

State University Press, Baton Rouge. 266 p. 

Russell, A.B. 1997. Trees of the Maritime Forest. 

Department of Horticultural Science, North 

Carolina State University, 

http://www.ces.ncsu.edu/depts/hort/ 

consumer/factsheets/maritime. 2 p. 

Scheper, J. 2002. Aralia spinosa. Floridata. http:// 

www.floridata.com/ref/a/aral_spi.cfm. 4 p. 

Sullivan, J. 1992. Aralia spinosa. In: U.S. 

Department of Agriculture, Forest Service, Rocky 

Mountain Research Station Fire Sciences 

Laboratory (2002 February). Fire Effects 

Information System 

http://www.fs.fed.us/database/feis. 10 p. 

Synor, T.D. and W.F. Cowen. [No date]. Ohio 

Trees. Aralia. Department of Natural Resources, 

Ohio State University Extension Bulletin 700-00. 

http://www.ag.ohiostate.edu/~ohioline/b700/index. html. 2 p. 

U.S. Department of Agriculture. [No date]. 

Southern wetland flora: Field office guide to plant 

species. USDA Soil Conservation Service, South 

National Technical Center, Fort Worth, Texas. 

Jamestown, ND: Northern Prairie Wildlife 

Research Center Home Page, 

http://www.npwrc.usgs.gov/resource/1999 

/soutflor/soutflor.htm. [not paged]. 
43

Ardisia elliptica Thunb. shoebutton ardisia 

MYRSINACEAE 

Synonyms: Ardisia ketoensis Hayata 

Bladhia elliptica (Thunb.) Nakai 

John K. Francis 

 

General Description.—Shoebutton ardisia, also 

known as sea-shore ardisia, blackberry, 

mameyuelo, mata pelandok, penah periah, ati 

popa’a, and atiu (Corner 1952, Pacific Island 

Ecosystems at Risk 2002), is an invasive shrub or 

small tree to 6 m in height and 15 cm in basal 

diameter. The plants produce strong stems with 

gray bark. The stems are usually single, but 

additional sprouts may arise from the rootstalk, 

especially if the plant is injured. Shoebutton ardisia 

grows a strong taproot, much-branched laterals, 

and fine roots with rhizomorphic tips. Leaves have 

a rubbery or leathery texture and are pink when 

young, turning dark green later. They are glabrous 

and alternate with petioles about 1 cm long and 

blades 8 to 12 cm by about 3 cm. The 

inflorescences are axillary, and umbellate, 

corymbose, or racemose with numerous fivemerous, pink flowers. The drupes are globose 

drupes 6 to 11 mm in diameter, dark purple to dull 

black when ripe and contain one seed each 

(author’s observation, Corner 1952, Howard 1989, 

Pacific Island Ecosystems at Risk 2002). 

Range.—The taxonomy and range of shoebutton 

ardisia are uncertain. Native areas for Ardisia 

elliptica have been listed variously as Sri Lanka, 

India, China, Taiwan, Malaya, South East Asia, 

Indonesia, and the Philippines (Corner 1952, 

Tomlinson 1986, Yuen-Po 1999). Some authors 

unite A. polycephala Wall., A. solanacea Roxb., 

and A. humilis Vahl under A. elliptica (Center for 

Aquatic and Invasive Plants 2002, Neal 1965, 

Pacific Island Ecosystems at Risk 2002). These 

species are difficult to distinguish. Liogier (1995) 

erroneously cites A. solanacea instead of A. 

elliptica in Puerto Rico. Shoebutton ardisia is also 

naturalized and escaped in Hawaii (Pacific Island 

Ecosystems at Risk 2002), Florida (Center for 

Aquatic and Invasive Plants 2002), and many other 

areas. 

Ecology.—Shoebutton ardisia principally grows 

and is most aggressive in areas of wet soils. It 

forms dense stands in low areas and intermittent 

stream bottoms with clay soils in Puerto Rico. It is 

found in wet, lowland areas in Hawaii (Pacific 

Island Ecosystems at Risk 2002), in hammocks, 

disturbed wetlands, old fields, and tree islands in 

marshes in Florida (Center for Aquatic and 

Invasive Plants 2002), and in mangroves elsewhere 

(Tomlinson 1986). The annual precipitation of 

these areas in Puerto Rico and probably elsewhere 

is above 1500 mm/yr. The species is shade tolerant 

and usually grows under forest canopies. Seedlings 

appear in well-established forest understories—

disturbance is not required for establishment. 

Although significant insect herbivory does occur in 

Florida, it does not appear to inhibit the advance of 

the species into new habitats (Dominguez and 

others 2002). 

Reproduction.—Shoebutton ardisia blooms 

sporadically throughout the year (Miami-Dade 

County 2002). The species is insect pollinated and 

self-fertile (Pascarella 1997). A collection of fruits 

from Puerto Rico weighed an average of 0.321 +

0.005 g/fruit. Seeds separated from them weighed 

(air-dried) an average of 0.0463 + 0.001 g/seed. 

Both the fruits and seeds are uniform in weight 

(coefficients of variation 15 and 17, respectively). 

Sown without any pretreatment in commercial 

potting mix, 75 percent germinated between 42 and 

81 days after sowing. Germination is epigenous. 

Although seedlings are subject to damping-off 

mortality right after germination, older seedlings 

transplant with little mortality or loss of vigor. 
44 

Birds are the principal seed dispersers, although 

raccoons and opossums in Florida also eat the 

fruits and disperse seeds (Miami-Dade County 

2002). 

Growth and Management.—Although 

shoebutton ardisia may grow as much as 1 m the 

first year as a sprout, seedlings and established 

shrubs grow slowly, usually 0.25 to 0.5 m/year in 

height. No figures are known for longevity, but 10 

to 25 years or more are likely. It is often desirable 

to control shoebutton ardisia in natural areas. Small 

numbers of seedlings and shrubs can be eliminated 

by pulling and grubbing. A broadcast spray of 

glyphosate is effective in killing dense stands of 

seedlings but will also kill native plants. Larger 

plants can be killed by applying a basal spray of 

triclopyhr mixed with an oil diluent (Pacific Island 

Ecosystems at Risk 2002). 

Benefits and Detriments.—Shoebutton ardisia 

was once used heavily as an ornamental for accent 

and background plantings. Many plantings still 

remain, but the species is no longer cultivated 

commercially (Miami-Dade County 2002). The 

fruits are edible and taste slightly sour with a hint 

of starch but otherwise lack flavor. The species is 

useful for fuel and vegetable stakes. The air-dry 

specific gravity of stem wood in Puerto Rico was 

measured at 0.48 + 0.01 g/cm3

. Biomass (dry 

weight) of plants from 137 cm in height to plants 

with a 5-cm D.B.H. may be calculated by the 

model: Total Dry Weight in grams = 22.020(stem 

length in meters) (diameter at 30 cm)2

 (Francis 

2000). Shoebutton ardisia is extremely invasive. It 

forms dense stands under trees in wet sites in 

Florida, Hawaii, and Puerto Rico that suppress 

native understory plants (Center for Aquatic and 

Invasive Plants 2001). 

References 

Center for Aquatic and Invasive Plants. 2002. 

Ardisia elliptica Thumb., Myrsinaceae/Myrsine 

family. University of Florida, Gainesville, FL. 

www.aquat1.itas.ufl.edu. 2 p. 

Corner, E.J.H. 1952. Wayside trees of Malaya. 

Government Printing Office, Singapore, Malaya. 

772 p. 

Dominguez, J., A. Scott, T. Scott, G. Valdes, C. 

Glenn, and C. Moore. 2002. Herbivore damage 

on the invasive exotic Ardisia elliptica and the 

native A. escallonioides in Southeastern Florida. 

NSF Research in Ecology: Invasive Species. 

http://fig.cox.miami.edu/ys99/gladres.html. 3 p. 

Francis, J.K. 2000. Estimating biomass and carbon 

content of saplings in Puerto Rican secondary 

forests. Caribbean Journal of Science 36(3-4): 

346-350. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Miami-Dade County. 2002. Shoebutton ardisiaArdisia elliptica, Ardisia humilis. 

http://www.co.miami-dade.fl.us/derm/ 

environment/badplants/plant%20.../shoebutton_

ardisia.ht. 1 p. 

Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub. 

50. Bernice P. Bishop Museum Press, Honolulu, 

HI. 924 p. 

Pacific Island Ecosystems at Risk. 2002. Invasive 

plant species: Ardisia elliptica Thunberg, 

Myrsinaceae. http://www.hear.org/pier/arell. 

htm. 2 p. 

Pascarella, J.B. 1997. Breeding systems of Ardisia

Sw. (Myrsinaceae). Brittonia 49(1): 45-53. 

Tomlinson, P.B. 1986. The botany of mangroves. 

Cambridge University Press, London. 413 p. 

Yuen-Po, Y. 1999. An enumeration of 

Myrsinaceae of Taiwan. Botanical Bulletin of 

the Academy Sinensis 40: 39-47. 
45

Argythamnia candicans Sw. argythamnia 

EUPHORBIACEAE 

Synonyms: Tournesol candicans M. Gómez 

John K. Francis 

General Description.—No common name for A. 

candicans is listed in the literature. The convention 

among local natural resource workers, when a 

common name does not exist, is to use its genus or 

species name as a common name. Argythamnia is a 

common upright, low or half shrub that reaches 1.5 

m in height and 2 cm of ground-line diameter. 

Usual heights are from 0.5 to 1.0 m. It normally 

has a single stem, although branches may arise near 

the base. It is supported by a tap and lateral root 

system with fine tertiary roots. The branches and 

twigs are thin and gray-green or whitish-green in 

color. The leaves are green or gray-green above 

and reddish- or purplish-green and pubescent 

below. Petioles are 1.5 to 2.5 mm long. The blades 

are usually narrowly elliptic, 1.3 to 8.5 cm long 

and 0.9 to 3.6 cm wide. Argythamnia produces 

tiny, inconspicuous staminate and pistilate flowers. 

The latter develop into round, flattened capsules 5 

to 6 mm in diameter. The capsules have three 

chambers, each with a globose, dark brown seed 

1.5 to 2 mm in diameter (Howard 1989, Liogier 

1988). 

Range.—Howard (1989) lists the range of 

agrythamnia as the Bahamas, the Greater Antilles, 

Anguilla, St. Martin, St. Barts, Guadeloupe, and 

Martinique. However, Britton and Millspaugh 

(1962) state that the former listings for A. 

candicans in the Bahamas should be A. lucayana

Millsp., an endemic. 

Ecology.—Argythamnia inhabits areas of Puerto 

Rico that receive from 750 to 1800 mm of annual 

precipitation at elevations from near sea level to 

about 450 m, usually within a few kilometers of the 

sea. Soils of all textures derived from all local rock 

types appear to be suitable, if well drained. Most 

of the modern habitat is rocky, steep, or disturbed. 

These sites include primary and secondary forests, 

brushy natural pastures, and roadsides. The species 

occurs as scattered individual plants or 

occasionally in thickets. Argythamnia is 

moderately intolerant of shade. It requires at least 

good filtered sunlight to survive. Argythamnia 

plants withstand light grazing well but tend to 

disappear from areas that are overgrazed. It 

resprouts after fires but is eliminated from large 

areas by repeated fires and competition from the 

exotic grasses, Urochloa maximum (Jacq.) R.O. 

Webster and Bothriochloa pertusa (L.) A. Camus. 

Frosts do not occur in the native range. 

Reproduction.—Argythamnia flowers and fruits 

continuously. During the maturation and drying 

process, the capsules develop internal tension and 

eventually explode flinging seeds as much as 0.3 m 

away from the parent plant. A collection of 

capsules from Puerto Rico ranged in fresh weight 

from 0.0257 to 0.0381. Air-dry seeds collected 

from them averaged 0.0023 + 0.0000 g/seed or 

429,000 seeds/kg. On moist filter paper, 97 percent 

of these seeds germinated between 10 and 82 days 

after sowing. 

Growth and Management.—Argythamnia is a 

relatively short-lived species. Seed-bearing plants 

examined by the author all had two or three growth 

rings in the base of the stem. Establishment of 

seedlings could be encouraged by soil disturbance 
46 

near seed-bearing plants. No planting experience 

has been reported. 

Benefits.—Argythamnia is eaten by livestock and 

contributes to ground cover and environmental 

diversity. 

References 

Britton, N.L., and C.F. Millspaugh. 1962. The 

Bahama Flora. The New York Botanical 

Garden. New York. 695 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 
47

Artemisia arbuscula Nutt. low sagebrush 

ASTERACEAE 

Synonyms: Artemisia tridentata var. arbuscula (Nutt.) McMinn 

 Serphidium arbusculum (Nutt.) W.A. Weber 

E. Durant McArthur and J.R. Taylor 

 

General Description.—Low sagebrush is a short, 

spreading, irregularly-branched shrub up to 50 cm 

high. Twigs are slender and densely canescent 

(silvery white pubescence) but may become nearly 

glabrous, and thus darker green, in late summer. 

Leaves are broadly cuneate, 0.5 to 1.5 cm long, 

and 0.3 cm wide. The leaf apex is usually threetoothed or cleft, although leaves on the upper part 

of the flowering stalks may be entire. Flower 

heads are arranged in elongate, narrow racemous 

panicles. Each head usually contains five to 11 

disc flowers and is subtended by 10 to 15 

canescent involucral bracts. 

Taxonomy.—Three subspecies of Artemisia 

arbuscula are currently recognized by the 

International Plant Names Index (2003). These are 

arbuscula, longicaulis Winward & McArthur, and 

thermopola Beetle. Subspecies longicaulis, also 

known as Lahonton low sagebrush, is endemic to 

western Nevada (Winward and McArthur 1995). It 

is probably a hybrid between Wyoming big 

sagebrush (Artemisia tridentata ssp. wyomingensis 

Beetle and Young) and low sagebrush (Winward 

and McArthur 1995). Subspecies thermopola, also 

known as hotsprings sagebrush, is a dwarf form 

endemic to the Stanley Basin area of Idaho, 

Jackson Hole area of Wyoming and east-central 

Oregon. Beetle (1960) speculated that it is a hybrid 

derived from low sagebrush and threetip sagebrush 

(Artemisia tripartite Rydberg). Low sagebrush has 

a base chromosome number of x = 9 and can be 

diploid, tetraploid, or hexaploid depending on 

population and subspecies (McArthur and 

Sanderson 1999, Winward and McArthur 1995). 

Range.—The range of low sagebrush extends 

throughout Utah, Idaho, northern California, 

Nevada, Oregon, Washington, southern Colorado, 

and western Montana. It is usually found at 

elevations ranging from 700 to 3,500 m. Low 

sagebrush can grow well in mountains above 3,000 

m, particularly in arid regions such as 

southwestern Utah and Nevada. 

Ecology.—Low sagebrush is adapted to dry, 

sterile, often rocky and alkaline clay soils. Mean 

annual precipitation throughout its range can vary 

between 250 and 700 mm. Seedlings develop roots 

quickly to reduce the effects of soil surface 

drought stress. Typical low sagebrush habitat is 

generally drier and rockier than that of big 

sagebrush. It can, however, grow in mosaics with 

big sagebrush where each species is confined to a 

particular soil type. Low sagebrush rarely grows in 

mixed stands with black sagebrush (McArthur and 

others 1979), and when it does, they don’t share 
48 

the same chromosome number (McArthur and 

Sanderson 1999). In areas where the distributions 

of these two overlap, low sagebrush usually 

inhabits sites that have more moisture or are at 

higher elevations than black sagebrush (A. nova 

Nelson) (Tisdale and Hironaka 1981, Ward 1953, 

Zamora and Tueller 1973). Low sagebrush is 

usually associated with pinyon-juniper, mountain 

brush, and mountain big sagebrush communities. 

Reproduction.—Flowering occurs from August to 

September, depending upon strain and elevation. 

Seeds ripen in October and November (McArthur 

and others 1979). Seeds are usually wind dispersed 

during late fall or winter, and seedlings emerge 

during the following spring. Seedlings emerge 

rapidly and grow quickly, although root growth 

generally outpaces stem growth during early 

development. Plants can be established on adapted 

sites by direct seeding, broadcasting, and drilling. 

Seeding should be conducted in the fall on a firm 

seedbed and seeds should remain within 2 mm of 

the soil surface. There are approximately 1,800 

cleaned seeds per gram (Meyer 2003) 

Management.—Low sagebrush is a source of 

browse for livestock and big game during winter 

months (Kufeld and others 1973, McArthur and 

others 1979). Some forms are browsed more 

heavily than others. A gray-green form in Nevada, 

for example, may be heavily browsed, while the 

green form is only lightly browsed (Brunner 

1972). Stand maintenance is often crucial because 

shrubs provide forage and cover under conditions 

that are intolerable for most shrubs. 

Attempts to seed low sagebrush out of its 

ecological range have been unsuccessful. 

Seedlings planted “offsite” usually establish but 

fail to reproduce. Stands recover quickly after 

burning or other disturbance. Low sagebrush 

spreads vigorously and can invade perennial 

seeded grass stands. Perennial herb and grass 

understory production usually decreases as stands 

mature. Low sagebrush is not a good candidate for 

mechanical control because habitat terrain is often 

extreme. 

Benefits.—Low sagebrush is a native shrub that 

adds structural and biological richness to arid 

communities throughout the Intermountain West. 

Low sagebrush communities provide important 

habitat for a variety of domestic and wild animals 

(Dealy and others 1981). Its ability to grow on 

exposed sites is important for wintering animals 

and shrub cover helps reduce on-site soil erosion. 

References 

Beetle, A.A. 1960. A study of sagebrush, the 

section Tridentatae of Artemisia. Bulletin 368, 

University of Wyoming Agriculture Experiment 

Station, Laramie, WY. 83 p. 

Brunner, J.R. 1972. Observations on Artemisia in 

Nevada. Journal of Range Management 25: 205-

208. 

Dealy, J.E., DA. Leckenby, D.M. Concannon. 

1981. Wildlife habitats in managed 

rangelands—the Great Basin of southeastern 

Oregon. General Technical Report PNW-120, 

USDA Forest Service, Pacific Northwest Forest 

and Range Experiment Station, Portland, OR, 

66 p. 

International Plant Names Index. 2003. 

International plant names index. 

http://www.ipni.org. [not paged]. 

Kufeld, R.C., O.C. Wallmo, and C. Feddema. 

1973. Foods of the Rocky Mountain mule deer. 

Research Paper RM-111, USDA Forest Service, 

Rocky Mountain Forest and Range Experiment 

Station, Fort Collins, CO. 31 p. 

 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens. 1979. Characteristics and 

hybridization of important Intermountain shrubs. 

III. Sunflower family. Research Paper INT-220, 

USDA Forest Service, Intermountain Forest and 

Range Experiment Station, Ogden, UT. 82 p. 

McArthur, E.D. and S.C. Sanderson. 1999. 

Cytogeography and chromosome evolution in 

subgenus Tridentatae of Artemisia (Asteraceae). 

American Journal of Botany 86: 1,754-1,775. 

Meyer, S.E. 2003. Artemisia. In: T.F. Bonner and 

R.G. Nisley, eds. Woody plant seed manual. 

http://www.wpsm.net. 12 p. 

Tisdale, E.W. and M. Hironaka. 1981. The 

sagebrush—grass region: a review of the 

ecological literature. Bulletin 33. Forest, 

Wildlife, and Range Experiment Station, 

University of Idaho, Moscow, ID. 31 p. 

Ward, G.H. 1953. Artemisia, section Seriphidium, 

in North America, a cytotaxonomic study. 

Contributions from the Dudley Herbarium 4: 

155-205. 
49

Winward, A.H. and E.D. McArthur. 1995. 

Lahontan sagebrush (Artemisia arbuscula ssp. 

longicaulis): a new taxon. Great Basin 

Naturalist 55(2): 151-157. 

Zamora, B. and P.T. Tueller. 1973. Artemisia 

arbuscula, A. longiloba, and A. nova habitat 

types in northern Nevada. Great Basin Naturalist 

33: 225-242. 
50 

Artemisa bigelovii Gray Bigelow sagebrush 

ASTERACEAE 

Synonyms: Artemisia petrophila Wooton & Standley 

E. Durant McArthur and Jeffrey R. Taylor 

 

General Description.—Bigelow sagebrush, also 

known as flat sagebrush (Hall and Clements 1923), 

is a low shrub 20 to 40 cm high with numerous 

spreading branches. Leaves are narrowly cuneate, 

1 to 2 cm long, 0.2 to 0.5 cm wide, and normally 

tridentate, although tips may be variously shaped. 

The leaves on vegetative branches are similar to 

those of big sagebrush (Artemisia tridentata Nutt.), 

although the lobes of Bigelow sagebrush are 

shallower and more sharply dentate (McArthur and 

others 1979). The odor of crushed leaves is mild 

and pleasant. New growth is covered with a 

silvery-canescent pubescence. Flower stems are 

slender and erect and inflorescences form long 

narrow panicles with short, recurved branches. 

Flower heads normally bear one ray flower and 

two disc flowers. Occasionally, however, heads 

may include zero or two ray flowers and one or 

three disc flowers. The turbinate involucre consists 

of eight to 12 short, densely tomentose bracts 2 to 

4 mm long and 1.5 to 2.4 mm wide. 

Taxonomy.—This taxon is undivided, and 

ecotype variation has not been documented. 

Chromosome races from 2x to 8x, however, have 

been documented (McArthur and others 1981; 

McArthur and Sanderson 1999). Bigelow 

sagebrush occupies a taxonomic position between 

the true sagebrushes (subgenus Tridentatae) and 

other Artemisia species (subgenus Artemisia). We 

have chosen to treat Bigelow sagebrush as a 

member of subgenus Tridentatae because of its 

growth habit, wood anatomy, leaf form, 

chromosomal karyotype, RAPD molecular genetic 

markers, and ITS sequences in nuclear ribosomal 

DNA (Kornkven and others 1998, McArthur and 

others 1981, McArthur and others 1998). The 

confusion exists because flower heads may include 

one or two ray flowers in addition to the 

characteristic disc flowers of the Tridentatae. 

Range.—The distribution of Bigelow sagebrush is 

more southern than other sagebrushes. It covers 

approximately 88,000 km2

 through western Texas, 

southern Colorado, New Mexico, Arizona, Utah, 

Nevada, and California between 900 and 2,400 m 

elevation (Beetle 1960, Kearney and Peebles 1960, 

Ward 1953). 

Ecology.—Bigelow sagebrush is one of the most 

drought-tolerant sagebrushes. It typically grows in 

canyons, gravelly draws, and dry flats. It also 

grows on rocky soils of the southern portion of the 

pinyon-juniper woodlands (Hall and Clements 

1923). It is often found in mixed stands with big 

sagebrush, black sagebrush (Artemisia nova

Nelson), leafless green rabbitbrush 

[Chrysothamnus nauseosus Pallas ex Pursh ssp. 

junceus (Greene) Hall & Clements], shadscale 

[Artriplex confertifolia (Torr. & Frem.) Wats.], 
51

and especially broom snakeweed [Gutierrezia 

sarothrae (Pursh) Britt. & Rusby]. Bigelow 

sagebrush is usually resistant to the common rust 

diseases and insect galls common to other 

Tridentatae taxa (Beetle 1960). It is, however, 

susceptible to wilt disease (Nelson and Krebill 

1981). It grows in areas that usually receive 250 to 

400 mm of annual precipitation. 

Reproduction.—Flowering occurs from August to 

October and seeds mature during the following 

months. Seeds are wind dispersed in the fall and 

winter, and seedlings emerge in spring. Seeds are 

among the smallest of sagebrush seeds, about 5.5 

million cleaned seeds per kg (Meyer 2003). 

Seedlings grow rapidly, and young shrubs are 

robust and vigorous. Plants appear to have good 

drought tolerance, even as seedlings. 

 Direct seeding has been limited to 

experimental plantings. Seeded shrubs grow well 

and natural spread normally occurs around the 

parental plant, suggesting that Bigelow sagebrush 

can be successfully seeded. 

 

Management.—Bigelow sagebrush is palatable to 

livestock and game throughout its distribution. Its 

twigs are less woody, aromatic, and bitter than 

most species within the big sagebrush complex 

(Hall and Clements 1923). In addition, Bigelow 

sagebrush is more grazing tolerant than most other 

sagebrushes. These shrubs, however, are generally 

scattered, and extensive stands are found only near 

the Four Corners area of Utah, Arizona, New 

Mexico, and Colorado. 

Benefits.—Bigelow sagebrush adds biological 

diversity to shrub communities throughout the 

Southwestern United States. This species improves 

soil stability and reduces erosion. It also provides 

palatable forage for livestock and wildlife and adds 

structural diversity to the landscape. 

References 

Beetle, A.A. 1960. A study of sagebrush: the 

section Tridentatae of Artemisia. Bulletin 368. 

Agricultural Experiment Station, University of 

Wyoming, Laramie, WY. 83 p. 

Hall, H.M. and F.E. Clements. 1923. The 

phylogenetic method in taxonomy, the North 

American species of Artemisia, Chrysothamnus, 

and Atriplex. Publication 326. Carnegie 

Institution of Washington, Washington, DC. 

355 p. 

Kearney, T.H. and R.H. Peebles. 1960. Arizona 

flora, 2nd edition. University of California Press, 

Berkeley, CA. 1,065 p. 

Kornkven, A.B., L.E. Watson, and J.R. Estes. 

1998. Phylogenetic analysis of Artemisia section 

Tridentatae (Asteraceae) based on sequences 

from the internal transcribed spacers (ITS) of 

nuclear ribosomal DNA. American Journal of 

Botany 85: 1787-1795. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 82 p. 

McArthur, E.D., C.L. Pope, and D.C. Freeman. 

1981. Chromosomal studies of subgenus 

Tridentatae of Artemisia: evidence for 

autopolyploidy. American Journal of Botany 68: 

589-605. 

McArthur, E.D. and S.C. Sanderson. 1999. 

Cytogeography and chromosome evolution of 

subgenus Tridentatae of Artemisia (Asteraceae). 

American Journal of Botany 86: 1754-1775. 

McArthur, E.D., R. Van Buren, S.C. Sanderson, 

and K.T. Harper. 1998. Taxonomy of 

Sphaeromeria, Artemisia, and Tanacetum

(Compositatae, Anthemideae) based on 

randomly amplified polymorphic DNA (RAPD). 

Great Basin Naturalist 58: 1-11. 

Meyer, S.E. 2003. Artemisia. In: T.F. Bonner and 

R.G. Nisley, eds. Woody plant seed manual. 

http://www.wpsm.net 12 p. 

Nelson, D.L. and R.G. Krebill. 1981. A sagebrush 

wilt disease of unknown origin. Great Basin 

Naturalist 41: 184-191. 

Ward, G.H. 1953. Artemisia, section Seriphidium, 

in North America. A cytotaxonomic study. 

Contributions from the Dudley Herbarium 4: 

155-205. 
52 

Artemisia californica Less. California sagebrush

ASTERACEAE 

Synonyms: None 

Arlee M. Montalvo and Catherine E. Koehler 

General Description.—Mature California 

sagebrush are 0.6 to 2.5 m tall and abundantly 

branched from the base (Munz and Keck 1968, 

Hickman 1993). The common name refers to its 

strong, sage-like aroma and being endemic to 

California and Baja California. The stems are 

whitish with appressed hairs, long, slender, leafy, 

and flexible. The somewhat hairy gray-green 

leaves are soft, entire to divided into narrow linear 

segments, giving the entire shrub a wispy 

appearance. The many small flower heads are less 

than 5 mm wide, arranged in racemes, and have 15 

to 30 disk florets per flower head. Plants are 

diploid with n = 9 chromosomes (Hickman 1993). 

Range.—California sagebrush occurs in California 

from Contra Costa Co. south into Baja California, 

including the Channel Islands, at elevations of less 

than 800 m (Munz and Keck 1968, Hickman 

1993). Historically, it was the dominant shrub on 

north-facing slopes in coastal sage scrub but has 

declined to about a third of its past abundance in 

many areas of southern California, especially 

inland on north-facing slopes where it is being 

replaced by exotic annual grasses (Minnich and 

Dezzani 1998). 

Ecology.—California sagebrush is common in 

sage scrub and coastal strand on dry slopes and 

fans (Munz and Keck 1968, Hickman 1993). It is 

an indicator species of sage scrub (Kirkpatrick and 

Hutchinson 1977), occurs on virtually all soil types 

except serpentine (Westman 1981a), and its range 

is limited primarily by cold winter temperatures 

(Malanson and O'Leary 1995). Leaves are 

drought-deciduous and seasonally dimorphic 

(Westman 1981b, Gray 1982). In California, shoot 

and new leaf production begins with the winter 

rains, usually in December, and continues 

throughout winter and spring. Leaf drop and 

production of smaller leaves occurs during the 

summer drought (Westman 1981b). The shallow, 

branched roots (Harrison and others 1971) allow 

for rapid water absorption and growth response to 

shallow rains (Gray 1982). The thin leaf cuticle 

and numerous stomata allow a high photosynthetic 

rate in response to water availability (Poole and 

Miller 1975, Gray 1983). Consequently, there is a 

higher transpiration rate and longer period of water 

stress compared to species with thick leaves and 

deep roots (Poole and Miller 1975, Gray 1982). 

Roots form associations with arbuscular 

mycorrhizal (AM) fungi, but the beneficial nature 

of the interaction may be facultative and is 

potentially affected adversely by nitrogen 

deposition (Sigüenza 2000, Yoshida and Allen 

2001). 

Studies on chemistry of leaves, litter, and 

soil below shrubs documented the presence of both 

soluble compounds and volatile monoterpenes and 

sesquiterpenes that inhibit germination or growth 

of some plants (Halligan 1973, 1975, 1976). Small 

mammalian herbivores and seed eaters often 

restrict foraging to beneath shrub canopies and to 

within a short distance of protective cover 

(Halligan 1973, 1974) and may contribute to 

conspicuous bare zones on the edge of shrub 

stands and under canopies.

Reproduction.—Flowering of California 

sagebrush tends to peak in late fall, but in Baja 

California, summer rains may trigger flowering 

during summer (Minnich 1985). The reduced 

flowers are typical of other wind-pollinated 

species of Artemisia. The single-seeded fruits 
53

(achenes) ripen in December and January, with 

some variation among years and habitats, and are 

primarily wind-dispersed (Eliason and Allen 1997, 

DeSimone and Zedler 1999). Achenes are tiny 

(about 60 micrograms) and in bulk amount to 

about 14,300,000 seeds/kg. (personal 

communication with S&S Seeds, Carpenteria, 

California). Seeds germinate in canopy openings 

or in small grassland clearings generated by 

gophers (Thomomys spp.), but seedlings suffer 

high mortality from gopher activity (Eliason and 

Allen 1997, DeSimone and Zedler 1999, 2001). 

Seedlings emerge in the rainy season, and most 

growth occurs by May. 

Seed Germination.—Seeds will germinate when 

fresh, but stored seeds may need cold stratification 

to enhance germination (De Hart 1994). While 

testing the combined effects of light and fire 

components on germination, 73 percent of seeds 

exposed to light germinated on soil, but seeds in 

the dark did not germinate (Keeley 1987). These 

results are consistent with field data that show 

germination may be limited by reduced light 

(Elliason and Allen 1997, Montalvo and others 

2002). Certain components of fire also influence 

germination. In light, treatment with leachate from 

charred wood (charrate) alone or with heat 

increases germination by 5 to 14 percent, but 

germination is about a third lower when seeds are 

subjected to high temperatures (70 to 120 o

C). In 

contrast, in darkness both heat and charrate alone 

stimulate germination, although heat and charrate 

together generally decrease germination. Thus 

buried seeds may require some exposure to fire in 

order to germinate, but such exposure yields 

inferior germination compared to light alone 

(Keeley 1987). 

Genetics and Geographic Variation.—Near the 

coast, plants vary in pubescence, color, chemistry, 

and physiology (O'Brien 1980). Shrubs vary 

between a green form with sparse hairs on the 

leaves to a form with dense hairs that give the 

plants a gray hue. Populations differ in the 

distinctness of forms and in their relative 

frequency. Seedlings raised together generally 

retain the grayness of their parents but can become 

grayer with age (O'Brien 1980). Differences in 

pubescence also correspond to differences in water 

content, rate and timing of shoot elongation, leaf 

retention, and chemistry. The lower leaf-water 

content of gray plants negatively impacts larval 

growth and fecundity of the beetle, Trirhabda 

sericotrachyla (O'Brien 1980). 

Fire Effects.—California sagebrush resprouts 

after fire about 25 percent of the time (Keeley 

1998). Resprouting appears to be lower in burns 

through dense vegetation, where plants are older, 

or if fire intensity is high (Malanson and O'Leary 

1982, Keeley 1998, Minnich and Dezzani 1998). 

Seedling emergence after fire is variable and low 

(Zedler and others 1983, Keeley 1998). Seedlings 

tend to appear the second year from seeds of 

resprouts or seeds blown in from adjacent areas. 

Under high fire intensities or frequency, California 

sagebrush will likely be extirpated because of its 

poor resprouting ability and poor competitive 

ability of seedlings (Malanson and O'Leary 1982, 

Malanson and Westman 1991). Development, 

increased fire frequency, competition with exotic 

grasses, poor growth in grasslands, and air 

pollution are each detrimental to this shrub’s 

survival (Eliason and Allen 1997, Allen and others 

1998, Minnich and Dezzani 1998, Keeley 1998). 

Horticulture.—California sagebrush can be 

compact if kept pruned and subjected to occasional 

pinching. These aromatic plants remain attractive 

in the dry season with occasional water (Schmidt 

1980, Keator 1994). It prefers well-drained soils 

and a low organic matter content (Wasowski and 

Wasowski 1995). Naturally occurring prostrate or 

low-mounding varieties are in cultivation (Browse 

1987, Perry 1992, Wasowski and Wasowski 1995) 

and look good in dry borders or as a foreground to 

contrasting taller shrubs (Keator 1994). 

Benefits.—California sagebrush provides habitat 

for many plant and animal species and is an 

important component of critical habitat for the rare 

California gnatcatcher, Polioptila californica

(Weaver 1998). The bird forages preferentially in 

California sagebrush and the often co-dominant 

shrub Eriogonum fasciculatum Benth. (Beyers and 

Wirtz 1997), in part because they house a high 

number and diversity of arthropods. Osborne 

(1998) found high diversity and abundance of 

arthropods in both species and concluded that A. 

californica, together with other sage scrub shrub 

species, is important to the maintenance of high 

arthropod species richness and abundance in 

coastal sage scrub. California sagebrush is also 

involved in species-specific interactions. For 

example, it is the host for Trirhabda 

sericotrachyla, an herbivorous beetle whose 

developmental cycle is closely associated with the 

seasonal growth of its only known host plant 

(O'Brien 1980). 
54 

Historical Uses.—Leaves of California sagebrush 

have been used by Native Americans for smoking, 

in sweat-houses, and various other purposes (Bean 

and Saubel 1972). The Cahuilla used the plant to 

ensure proper maturation of girls into women. It is 

said to stimulate the uterine mucosa, ensuring 

rapid childbirth and, if regularly consumed as a 

decoction prior to the onset of each menstruation, 

to prevent menstrual cramps and alleviate 

menopausal trauma. Fresh or dried leaves were 

chewed to alleviate colds (Bean and Saubel 1972). 

Costanoan Indians applied leaves to wounds or 

aching teeth for pain reduction, and as a decoction 

to bathe patients with colds, coughs, rheumatism, 

or to be consumed and used as a poultice for 

treatment of asthma (Bocek 1984).

Growth and Management.—California 

sagebrush should be planted using shallow seeding 

methods such as hydroseeding or dry broadcasting 

followed by seed imprinting. Shallow methods are 

superior to planting with a range drill because the 

seeds need light for germination (Montalvo and 

others 2002). Attempts to use annual legumes as 

nurse plants can result in reduced seedling growth 

and survival (Marquez and Allen 1996), but 

seedling survival may be unaffected when sown 

with other native species at low density (Montalvo 

and others 2002). Clearing weeds before planting 

is critical. California sagebrush can return 

vigorously to areas where all vegetation has first 

been removed, even when sown with a variety of 

other species. Spring rains or artificial late fall and 

spring irrigation may enhance survival of seedlings 

(Williams and Hobbs 1989, Elliason and Allen 

1997, Padgett and others 2000). However, survival 

is not enhanced by late-season watering in wet 

years (DeSimone and Zedler 2001). 

References 

Allen, E.B., P.E. Padgett, A. Bytnerowicz, and R. 

Minnich. 1998. Nitrogen deposition effects on 

coastal sage vegetation of southern California. 

General Technical Report 166, U.S. Department 

of Agriculture, Forest Service. p. 131-139. 

Bean, J.L. and K.S. Saubel. 1972. Temalpakh: 

Cahuilla Indian knowledge and usage of plants. 

Malki Museum Press, Morongo Indian 

Reservation, CA. 225 p. 

Beyers, J.L. and W.O. Wirtz II. 1997. Vegetative 

characteristics of coastal sage scrub sites used by 

California gnatcatcher: implications for 

management in a fire-prone ecosystem. In: J. 

Greenlee, ed. Fire Effects on Rare and 

Endangered Species and Habitats. International 

Assoc. of Wildland Fire, Fairfield, WA. p. 81-89. 

Bocek, B. 1984. Ethnobotany of Costanoan Indians, 

California, based on collections by John P. 

Harrington. Economic Botany 38: 240-255. 

Browse, P.M. 1987. Artemisia californica 'canyon 

gray'. Pacific Horticulture 48: 56. 

De Hart, J. 1994. Propagation Secrets for California 

Native Plants. Jeanine De Hart, Encinitas, CA. 

28p. 

DeSimone, S.A. and P.H. Zedler. 1999. Shrub 

seedling recruitment in unburned Californian 

coastal sage scrub and adjacent grassland. 

Ecology 80: 2018-2032. 

DeSimone, S.A. and P.H. Zedler. 2001. Do shrub 

colonizers of southern Californian grassland fit 

generalities for other woody colonizers? 

Ecological Applications 11: 1,101-1,111. 

Eliason, S.A. and E.B. Allen. 1997. Exotic grass 

competition in suppressing native shrubland reestablishment. Restoration Ecology 5: 245-255. 

Gray, J.T. 1982. Community structure and 

productivity in Ceanothus chaparral and coastal 

sage scrub of southern California. Ecological 

Monographs 52: 415-435. 

Gray, J.T. 1983. Nutrient use by evergreen and 

deciduous shrubs in southern California. Journal 

of Ecology 71: 21-41. 

Halligan, J.P. 1973. Bare areas associated with 

shrub stands in grassland: the case of Artemisia

californica. BioScience 23: 429-432. 

Halligan, J.P. 1974. Relationship between animal 

activity and bare areas associated with California 

sagebrush in annual grassland. Journal of Range 

Management 27: 358-362. 

Halligan, J.P. 1975. Toxic terpenes from Artemisia

californica. Ecology 56: 999-1003. 

Halligan, J.P. 1976. Toxicity of Artemisia

californica to four associated herb species. The 

American Midland Naturalist 95: 406-421. 

Harrison, A.T., E. Small, and H.A. Mooney. 1971. 

Drought relationships and distribution of two 

Mediterranean-climate California plant 

communities. Ecology 52: 869-875. 

Hickman, J.C, ed. 1993. The Jepson Manual: 

Higher Plants of California. University of 

California Press, Ltd., Los Angeles, CA. 1,400 p. 

Keator, G. 1994. Complete Garden Guide to the 
55

Native Shrubs of California. Chronicle Books, 

San Francisco, CA. 314 p. 

Keeley, J.E. 1987. Role of fire in seed germination 

of woody taxa in California chaparral. Ecology 

68: 434-443. 

Keeley, J.E. 1998. Postfire ecosystem recovery and 

management: The October 1993 large fire episode 

in California. In: J. M. Moreno, ed. Large Forest 

Fires. Backbuys Publishers, Leiden, Netherlands. 

p. 69-90. 

Kirkpatrick, J.B. and C.F. Hutchinson. 1977. The 

community composition of California coastal sage 

scrub. Vegetatio 35: 21-33. 

Malanson, G.P. and J.F. O'Leary. 1982. Post-fire 

regeneration strategies of Californian coastal sage 

shrubs. Oecologia 53: 355-358. 

Malanson, G.P. and J.F. O'Leary. 1995. The coastal 

sage scrub-chaparral boundary and response to 

global climatic change. In: J. M. Moreno and W. 

C. Oechel, eds. Global change and Mediterraneantype ecosystems. Volume 117. Springer-Verlag 

New York, Inc., NY. p 203-224. 

Malanson, G.P. and W.E. Westman. 1991. 

Modeling interactive effects of climate change, air 

pollution, and fire on California shrubland. 

Climate Change 18: 363-376. 

Marquez, V.J. and E.B. Allen. 1996. Ineffectiveness 

of two annual legumes as nurse plants for 

establishment of Artemisia californica in coastal 

sage scrub. Restoration Ecology 4: 42-50. 

Minnich, R.A. 1985. Evolutionary convergence or 

phenotypic plasticity? Responses to summer rain 

by California chaparral. Physical Geography 6: 

272-287. 

Minnich, R.A. and R.J. Dezzani. 1998. Historical 

decline of coastal sage scrub in the RiversidePerris plane, California. Western Birds 29: 366-

391. 

Montalvo, A.M., P.A. McMillan, and E.B. Allen. 

2002. The relative importance of seeding method, 

soil ripping, and soil variables on seeding success. 

Restoration Ecology 10: 52-67. 

Munz, P.A. and D.D. Keck. 1968. A California 

Flora with Supplement. University of California 

Press, Berkeley, CA. 1,681 + 224 p. 

O'Brien, P.Y. 1980. Adaptive relations between a 

stenophagous herbivore, Trirhabda sericotrachyla

(Coleoptera: Chrysomelidae), and its host plant, 

Artemisia californica (Compositae). Ph.D. 

Dissertation. University of California, Irvine. 125 

p. 

Osborne, K.H. 1998. A description of arthropod 

community structure in southern California 

coastal sage scrub. M.S. Thesis. University of 

California, Riverside. 133 p. 

Padgett, P.E., S.N. Kee, and E.B. Allen. 2000. The 

Effects of irrigation of revegetation of semi-arid 

costal sage scrub in Southern California. 

Environmental Management 26: 427-435. 

Perry, B. 1992. Landscape plants for western 

regions: an illustrated guide to plants for water 

conservation. Land Design Publishing, Claremont, 

CA. 318 p. 

Poole, D.K. and P.C. Miller. 1975. Water relations 

of selected species of chaparral and coastal sage 

communities. Ecology 56: 1,118-1,128. 

Schmidt, M.G. 1980. Growing California Native 

Plants. University of California Press, Los 

Angeles, CA. 366 p. 

Sigüenza, C. 2000. Nitrogen deposition and soil 

microorganisms of Artemisia californica and 

exotic grasses in southern California. Ph.D. 

Dissertation. University of California, Riverside. 

153 p. 

Wasowski, S. and A. Wasowski. 1995. Native 

Gardens for Dry Climates, 1st Edition. Clarkson 

Potter/Publishers, New York. 176 p. 

Weaver, K.L. 1998. Coastal sage scrub variations of 

San Diego county and their influence on the 

distribution of the California gnatcatcher. Western 

Birds 29: 392-405. 

Westman, W.E. 1981a. Factors influencing the 

distribution of species of California coastal sage 

scrub. Ecology 62: 439-455. 

Westman, W.E. 1981b. Seasonal dimorphism of 

foliage in Californian coastal sage scrub. 

Oecologia 51: 385-388. 

Williams, K. and R.J. Hobbs. 1989. Control of 

shrub establishment by springtime soil water 

availability in an annual grassland. Oecologia 81: 

62-66. 

Yoshida, L.C. and E.B. Allen. 2001. Response to 

ammonium and nitrate by a mycorrhizal annual 

invasive grass and native shrub in southern 

California. American Journal of Botany 88: 1,430-

1,436. 

Zedler, P.H., C.R. Gautier, and G.S. McMaster. 

1983. Vegetation change in response to extreme 
56 

events: the effect of a short interval between fires 

in California chaparral and coastal scrub. Ecology 

64: 809-818. 
57

Artemisia cana Pursh silver sagebrush 

ASTERACEAE 

Synonyms: Seriphidium canum (Pursh) W.A. Weber 

E. Durant McArthur and Jeffrey R. Taylor 

General Description.—Silver sagebrush, also 

known as white sagebrush or hoary sagebrush, is 

an erect, freely branched, rounded shrub up to 1.5 

m tall. Older stems have a dark brown, fibrous 

bark while younger stems are covered with a dense 

white to yellowish-green tomentum. Branches 

often layer when in contact with the soil. Leaves 

on vegetative branches are 1 to 10 mm wide and 2 

to 8 mm long. Leaf shape is linear to linearoblanceolate, entire, or occasionally with one or 

two irregular lobes. Leaves are covered with a 

silvery-white pubescence that becomes slightly 

viscid with age. On flowering stalks, leaves are 

similar to those on vegetative branches but may be 

smaller, especially toward the distal end. The 

foliage emits a mild to pungent odor when 

crushed. Flower heads are arranged into dense, 

narrow, leafy panicles, sometimes reduced to a 

raceme or spike-like inflorescence. Each head 

contains from four to 20 disc flowers that are 

subtended by ovate involucral bracts. Silver 

sagebrush exhibits several levels of polyploidy 

based on x = 9, varying by subspecies and 

population. Diploid, tetraploid, and octaploid, 

populations have been documented (McArthur and 

others 1981, McArthur and Sanderson 1999, Ward 

1953). 

Taxonomy.—Silver sagebrush comprises three 

subspecies: A. cana ssp. cana, A. cana ssp. 

viscidula, and A. cana ssp. bolanderi. Ploidy levels 

are octaploid, diploid or tetraploid, and diploid, 

respectively (McArthur and others 1981, 

McArthur and Sanderson 1999, Ward 1953). 

Plains silver sagebrush (A. cana ssp. cana) and 

mountain silver sagebrush (A. cana ssp. viscidula) 

have a strong tendency to layer. Plains silver 

sagebrush is an erect, round, canescent, freely 

branched shrub up to 1.5 m tall. It has a more 

eastern distribution than the other subspecies. 

Putative natural hybrids have been found between 

plains silver sagebrush and big sagebrushes 

(Beetle 1960, Ward 1953). Plains silver sagebrush 

has the ability to spread rapidly after burning by 

resprouting and by rhizomes (Beetle 1960). 

Mountain silver sagebrush (A. cana ssp. 

viscidula) is an erect shrub usually not more than 

1.0 m tall. It is distinguished from plains silver 

sagebrush by its smaller, darker leaves, and more 

western distribution. Leaves on vegetative stems 

are 1 to 5 mm wide and up to 7 cm long. They are 

typically simple, entire, and grow in dark green 

clusters. This subspecies varies in appearance but 

is always darker than mountain big sagebrush, 

with which it is often growing (Beetle 1960). 

Bolander silver sagebrush (A. cana ssp. bolanderi) 

has narrow leaves like mountain silver sagebrush, 

but leaves are canescent like plains silver 

sagebrush. 

 

Range.—Aside from big sagebrush, silver 

sagebrush is the most widely distributed member 

of the genus. It occurs over approximately 140,000 

km2

, from British Columbia and Saskatchewan in 

the north, to Arizona and New Mexico in the 

south, and west to Oregon and California (Beetle 
58 

1960). Ranges for each subspecies, however, are 

smaller. Plains silver sagebrush is found mostly 

east of the Continental Divide, through Montana, 

the Dakotas, Wyoming, western Nebraska, and 

northern Colorado. Mountain silver sagebrush 

inhabits mountainous regions (around 1,800 m or 

higher) of Montana, south along the Continental 

Divide, and west to Arizona, Nevada, and Oregon. 

The range of Bolander silver sagebrush is 

restricted, generally to enclosed basins, in central 

Oregon and eastern California. 

Ecology.—Silver sagebrush grows on soils that 

are less mature, with less phosphorous, potassium, 

nitrogen, organic matter, and lower cation 

exchange capacity than soils supporting big 

sagebrush (Hazlett and Hoffman 1975). It inhabits 

various environments, each subspecies adapting to 

different soil conditions. Plains silver sagebrush 

grows particularly well on well-watered, deep soils 

of the northern Great Plains, especially along 

stream bottoms and drainageways (McArthur and 

others 1979, Walton and others 1986). Mountain 

silver sagebrush, on the other hand, grows in areas 

with a heavy, lingering snow pack (Beetle 1960, 

McArthur and others 1979, Tisdale and Hironaka 

1981, Winward 1980). Bolander silver sagebrush 

grows on poorly drained, alkaline soils (Beetle 

1960, McArthur and others 1979, Tisdale and 

Hironaka 1981, Winward 1980). Silver sagebrush 

usually grows in areas with precipitation between 

350 and 600 mm but it occurs in areas of water 

accumulation, such as near streams or in enclosed 

basins. 

Reproduction.—Flowers bloom between August 

and September and seed ripens in October and 

November. Seeds are wind dispersed during the 

winter months. Establishment by direct seeding 

has been successful (Kelsey 1986), especially 

when sown in the fall on the soil surface or at 

shallow depths (Wasser 1982). Seedlings emerge 

in spring and grow rapidly. Seeding silver 

sagebrush is best done in the late autumn or early 

spring to maintain high viability and high 

germination of seeds (Hou and Romo 1998, Romo 

and Grilz 2002, Romo and Young 2002). 

Seedlings are best established in lightly tilled soils 

so that stands of existing perennial grasses are 

maintained (Romo and Grilz 2002). 

Plants respond dramatically to moist 

conditions and may reach 35 to 50 cm in 1 year. 

Silver sagebrush can also spread asexually. 

Harvey (1981) successfully propagated plants 

through hardwood cuttings, and plants can also 

spread by rhizomes. There are approximately 

2,900 cleaned seeds per gram (Meyer 2003). 

Management.—Silver sagebrush can provide an 

important source of browse and is used quite 

extensively by livestock and big game, especially 

when other food sources are scarce (Kufeld and 

others 1973, Wasser 1982). In the western Great 

Plains area, silver sagebrush is an important 

survival food for antelope. Domestic sheep and 

mule deer found Bolander silver sagebrush among 

the most preferable sagebrushes during winter and 

fall feeding trials (Sheehy and Winward 1981). 

Sheep often browse silver sagebrush in the fall 

after forbs and grasses are dry. 

Silver sagebrush can occupy areas at high 

densities, and therefore is a candidate for plant 

control. It can be difficult to control because of its 

tendency to resprout from the crown and to spread 

by rhizomes. It is not as susceptible to fire as other 

species of sagebrush (Wright and others 1979, 

White and Currie 1983). White and Currie (1983) 

reported that silver sagebrush mortality after 

prescribed fall and spring burns was directly 

related to fire intensity. In general, fall burns are 

more effective at reducing silver sagebrush than 

spring burns. 

Benefits.—Silver sagebrush is an abundant native 

shrub that provides additional diversity to 

shrubland ecosystems. It is an important winter 

forage source for domestic sheep and game 

animals and adds structural variety to the 

landscape. Native Americans traditionally used a 

decoction from silver sagebrush to stop coughing 

(Stubbendieck and others 1997). In addition, EuroAmerican settlers and Native Americans used it 

for fuel (McArthur and others 1979). 

References 

Beetle, A.A. 1960. A study of sagebrush: the 

section Tridentatae of Artemisia. Bulletin 368, 

Agricultural Experiment Station, University of 

Wyoming, Laramie, WY. 83 p. 

Harvey, S.J. 1981. Life history and reproductive 

strategies in Artemisia. M.S. thesis, Montana 

State University, Bozeman, MT. 132 p. 

Hazlett, D.L. and G.R. Hoffman. 1975. Plant 

species distributional patterns in Artemisia 

tridentata—and Artemisia cana—dominatied 

vegetation in western North Dakota. Botanical 

Gazette 136: 72-77. 
59

Hou, J. and J.T. Romo. 1998. Cold-hardiness of 

silver sagebrush seedlings. Journal of range 

Management 51: 704-708. 

Kelsey, R.G. 1986. Emergence, seedling growth, 

and crude terpenoid concentration in a sagebrush 

garden. In: McArthur, E.D. and B.L Welch, 

comps. Proceedings—symposium on the biology 

of Artemisia and Chrysothamnus; 1984 July 9-

13, Provo, UT. General Technical Report INT200. U.S. Department of Agriculture, Forest 

Service, Intermountain Research Station, Ogden, 

UT. p. 358-365. 

Kufeld, R.C., O.C. Wallmo, and C. Feddema. 

1973. Foods of the Rocky Mountain mule deer. 

Research Paper RM-111. U.S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Forest and Range Experiment Station, Fort 

Collins, CO. 31 p. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service 

USDA Forest Service, Intermountain Forest and 

Range Experiment Station, Ogden, UT. 82 p. 

McArthur, E.D., C.L. Pope, and D.C. Freeman. 

1981. Chromosomal studies of subgenus 

Tridentatae of Artemisia: evidence for 

autopolyploidy. American Journal of Botany 68: 

589-605. 

McArthur, E.D. and S.C. Sanderson. 1999. 

Cytogeography and chromosome evolution of 

subgenus Tridentatae of Artemisia (Asteraceae). 

American Journal of Botany 86: 1,754-1,775. 

Meyer, S.E. 2003. Artemisia. In: F.T. Bonner, and 

R.G. Nisley, eds. Woody plant seed manual. 

http://www.wpsm.net 12 p. 

Romo, J.T. and R.W. Grilz. 2002. Establishment 

of silver sagebrush in the northern mixed prairie. 

Journal of Range Management 55: 217-221. 

Romo, J. T. and J.A. Young. 2002. Temperature 

profiles and the effects of field environments on 

germination of silver sagebrush. Native Plants 

Journal 3: 5-13. 

Sheehy, D.P. and A.H. Winward. 1981. Relative 

palatability of seven Artemisia taxa to mule deer 

and sheep. Journal of Range Management 34: 

397-399. 

Stubbendieck, J, Hatch, S.L., Butterfield, C.H. 

1997. North American Range Plants. University 

of Nebraska Press, Lincoln, Nebraska. 229 p. 

Tisdale, E.W. and M. Hironaka. 1981. The 

sagebrush—grass region: a review of the 

ecological literature. Bulletin 33, Forest, 

Wildlife, and Range Experiment Station, 

University of Idaho, Moscow, ID. 31 p. 

Walton, T.P., R.S. White, and C.L. Wambolt. 

1986. Artemisia rproductive strategies: a review 

with emphasis on silver sagebrush. In: E.D. 

McArthur and B.L Welch, comps. 

Proceedings—symposium on the biology of 

Artemisia and Chrysothamnus; 1984 July 9-13, 

Provo, UT. General Technical Report INT-200. 

U.S. Department of Agriculture, Forest Service, 

Intermountain Research Station, Ogden, UT: 67-

74. 

Ward, G.H. 1953. Artemisia, section Seriphidium, 

in North America. A cytotaxonomic study. 

Contributions from the Dudly Herbarium 4: 155-

205. 

Wasser, C.H. 1982. Ecology and culture of 

selected species useful in revegetating disturbed 

lands in the west. FSW/OBS-82/56. U.S. 

Department of the Interior Fish and Wildlife 

Service. Washington, DC. 347 p.

Winward, A.H. 1980. Taxonomy and ecology of 

sagebrush in Oregon. Station Bulletin 642. 

Argicultural Experiment Station, Oregon State 

University, Corvallis, OR. 15 p. 

White, R.S. and P.O. Currie. 1983. The effects of 

prescribed burning on silver sagebrush. Journal 

of Range Management 36: 611-613. 

Wright, H.A., L.F. Neuenschwander, and C.M 

Britton. 1979. The role and use of fire in 

sagebrush-grass and pinyon-juniper plant 

communites, a state-of-the-art review. General 

Technical Report INT-158. U.S. Department of 

Agriculture, Forest Service, Intermountain Forest 

and Range Experiment Station, Ogden, UT. 48 p.
60 

Artemisia filifolia Torr. sandsage 

ASTERACEAE 

Synonyms: Oligosporus filifolius (Torr.) W.A.Weber 

E.Durant McArthur and Jeffrey R. Taylor 

General Description.—Sandsage, also known as 

sand sagebrush and oldman sage or oldman 

sagebrush, is a freely branched, rounded shrub that 

grows up to 1.5 m tall. Young branches are 

covered with a canescent pubescence while the 

older stems are covered by a dark gray-green or 

blackish bark. Leaves are filiform, and covered 

with a canescent pubescence. Leaf length can 

range from 3 to 8 cm, and leaf width is less than 

0.5 cm. Leaves are entire or alternately divided 

into filiform divisions and are often fascicled. 

Flower heads are arranged into leafy narrow 

panicles. Each head usually contains two or three 

fertile, pistilate, ray flowers and one to six perfect, 

sterile, disc flowers. Five to nine canescent 

involucral bracts subtend each head. The 

receptacles and achenes are both glabrous. 

 

Taxonomy.—Sandsage does not have subspecific 

divisons. It has traditionally been assigned to the 

subgenus Drancunlus based on floral 

characteristics (McArthur 1979). Beetle (1979) 

suggested that it might have an affinity with true 

sagebrushes (subgenus Tridentatae). It does have 

some cytological, chloroplast DNA, and chemical 

similarities with Tridentatae (Kelsey and 

Shafizadeh 1979, Korkven and others 1998, 1999, 

McArthur and Pope 1979) although it differs from 

Tridentatae in floral characteristics, wood 

anatomy, and nuclear ribosomal DNA sequences 

(Korkven and others 1998, Moss 1940). The 

chromosome number for A. filifolia is 2n = 18; its 

karyotype is similar to that of the true sagebrushes 

(subgenus Tridentatae) (McArthur and Pope 

1979). 

Range.—Sandsage is common on sandy substrates 

from the southern Black Hills in South Dakota 

south to Texas and Chihuahua and west to Arizona 

and Nevada (Hall and Clements 1923, McArthur 

and others 1979, Rasmussen and Brotherson 

1986). It is particularly important in the central 

and southern Great Plains in bands along the sandy 

mantles that parallel stream drainages (Berg 1994) 

Ecology.—Sandsage grows almost exclusively on 

sand. It is probably the most widespread shrub on 

dunes and sandhills. In a study in southwestern 

Utah, Rasmussen and Brotherson (1986) found 

that sandsage communities were less diverse than 

adjacent communities, but plant densities were 

higher. They also reported that sandsage nutrient 

quality was not affected by the relative poor soil 

nutrient profile characteristic of some sandsage 

communities. Its accumulation of mineral nutrients 

above that of the soil is apparently an adaptation to 

sand habitats with inherently low soil fertility. It 
61

grows in areas with an annual precipitation range 

of 220 to 600 mm. 

Reproduction.—Blooming occurs during August 

and September, and seed ripens between October 

and December (McArthur and others 1979). Seeds 

are smaller than those of most sagebrushes, but 

seedlings grow well once established. There are 

approximately 3.2 million cleaned seeds per kg 

(Meyer 2003). Seedlings emerge in the spring and 

grow and mature quickly. Survival is reduced, 

however, if it is planted in habitats that extend 

beyond its natural range (Nelson and Krebill 1981). 

Sandsage has been seeded alone in most plantings 

but can be seeded with herbs adapted to arid sandy 

soils. It is easy to grow as bareroot stock and 

survives well when field planted. Wildings can be 

transplanted in the spring or fall, and plants can be 

easily reared from seed. 

Management.—The value of sandsage as a source 

of browse varies with community type. It is seldom 

eaten in grasslands where other forage is adequate 

but is consumed by cattle, domestic sheep, and big 

game in arid desert regions. It can be particularly 

important during dry years (Hall and Clements 

1923, McArthur and others 1979). It is important in 

the vegetation cover of harsh sandy sites, although 

limited water often reduces natural seed 

production. It is particularly useful in blackbrush 

and pinyon-juniper communities where sandy 

outcrops occur. 

 Sandsage resprouts vigorously after fire; 

fire may be useful in reducing canopy height and 

volume, thus allowing associated herbaceous 

species to express more vigor (Vermeire and 

others 2001. 

Benefits.—Sandsage can be an important, locally 

abundant, native shrub that provides habitat and 

forage for wildlife, including some small nongame 

birds. This species helps reduce wind erosion by 

stabilizing light sandy soils. People in Mexico 

have used a decoction of leaves to reduce the 

effects of intestinal worms and other stomach 

problems (Stubbendieck and others 1997). 

References 

Beetle, A.A. 1979. Autecology of selected 

sagebrush species. In: The sagebrush ecosystem: 

a symposium; 1978 April 27-28, Logan, UT. 

College of Natural Resources, Utah State 

University, Logan, UT. p. 23-26. 

Berg, W.A. 1994. Sand-sagebrush-mixed prairie, 

SRM 722. In: Shiflet, T. N., ed. Rangeland cover 

types of the United States. Society for Range 

Management, Denver, CO. p. 99. 

Hall, H.M. and F.E. Clements. 1923. The 

phylogenetic method in taxonomy, the North 

American species of Artemisia, Chrysothamnus, 

and Atriplex. Publication 326. Carnegie 

Institution of Washington, Washington, DC. 

355 p. 

Kelsey, R.G. and F. Shafizadeh. 1979. 

Sesquiterpene lactones and systematics of the 

genus Artemisia. Phytochemistry 18: 1,591-

1,611. 

Korkven, A.B., L.E. Watson, and J.R. Estes. 1998. 

Phylogenetic analysis of Artemisia section 

Tridentatae (Asteraceae) based on sequences 

from the internal transcribed spacers (ITS) of 

nuclear ribosomal DNA. American Journal of 

Botany 85: 1,787-1,795. 

Kornkven, A.B., L.E. Watson, and J.R. Estes. 

1999. Molecular phylogeny of Artemisia section 

Tridentatae (Asteraceae) based on chloroplast 

DNA restriction site variation. Systematic 

Botany 24: 69-84. 

McArthur, E.D. 1979. Sagebrush systematics and 

evolution. In: The sagebrush ecosystem: a 

symposium; 1978 April 27-28, Logan, UT. 

College of Natural Resources, Utah State 

University, Logan, UT. p. 14-22. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U. S. Department of Agriculture, Forest Service 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 82 p. 

McArthur, E.D. and C.L. Pope. 1979. Karyotypes 

of four Artemisia species: A. carruthii, A. 

filifolia, A. frigida, and A. spinescens. Great 

Basin Naturalist 39: 419-426. 

Meyer, S.E. 2003. Artemisia. In: T.F. Bonner and 

R.G. Nisley, eds. Woody plant seed manual. 

http://www.wpsu.net 12 p. 

Moss, E.H. 1940. Interxylary cork in Artemisia

with a reference to its taxonomic significance. 

American Journal of Botany 27: 762-768. 
62 

Nelson, D.L. and R.G. Krebill. 1981. A sagebrush 

wilt disease of unknown origin. Great Basin 

Naturalist 41: 184-191. 

Rasmussen, L.L. and J.D. Brotherson. 1986. 

Habitat relationships of sandsage (Artemisia 

filifolia) in southern Utah. In: McArthur, E.D. 

and B.L. Welch, comps. Proceedings—

symposium on the biology of Artemisia and 

Chrysothamnus; 1984 July 9-13, Provo, UT. 

U.S. Department of Agriculture, Forest Service, 

Intermountain Research Station, Ogden, UT. p. 

58-66. 

Stubbendieck, J., S.L. Hatch, and C.H. Butterfield 

1997. North American Range Plants. Fifth 

Edition. University of Nebraska Press, Lincoln, 

NB. 231 p. 

Vermeire, L.T., R.B. Mitchell, and S.C. 

Fuhlendorf. 2001. Sand sagebrush response to 

fall and spring prescribed burns. In: McArthur, 

E.D. and D.J. Fairbanks, comps. Shrubland 

ecosystem genetics and biodiversity: 

proceedings; 2000 June 13-15, Provo, UT. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Ogden, UT. 

p. 233-235. 
63

Artemisia frigida Willd. fringed sage 

ASTERACEAE 

Synonyms: Artemisia virgata Richardson 

E. Durant McArthur and Jeffrey R. Taylor 

General Description.—Fringed sage, known also 

as prairie sagewort, fringed sagewort and fringed 

sagebrush, is an aromatic, mat-forming, perennial 

shrub 20 to 60 cm tall (McArthur and others 1979, 

Stubbendieck and others 1997). The lower woody 

stems are spreading and often branched, while 

upper herbaceous stems are erect and leafy. 

Adventitous roots may occur when stems come in 

contact with soil. The entire plant is densely silvercanescent. Leaves are abundant, 6 to 12 mm long, 

and two or three times ternately (sub-pinnately) 

divided. Flowerheads are arranged into nodding 

racemes or open panicles. Small, densely hairy 

involucral bracts occur in series around each 

flowerhead. Each head contains 10 to 17 outer, 

seed-producing, pistillate, ray flowers and 

numerous (25 to 50) tubular, funnelform, perfect 

seed-producing disc flowers (McArthur and others 

1979, Stubbendieck and others 1997). This species 

has a deep perennial taproot with numerous 

extensive lateral roots as adaptations to withstand 

drought. 

Taxonomy.—Undoubtedly, fringed sage, with its 

broad geographical and altitudinal distribution, has 

considerable genetic variation. The taxon has three 

varieties: the typical variety, Artemisia frigida

Willd. var. frigida, Artemisia frigida var. 

gmelinianum Besser, and Artemisia frigida var. 

williamsae S.L. Welsh (International Plant Names 

Index 2003). Unlike several other widespread 

Artemisia species, fringed sage is known only at 

the diploid (2n = 2x = 18) chromosome level 

(McArthur and Pope 1979). Fringed sage is a 

member of the subgenus Artemisia (McArthur and 

Pope 1979). 

Range.—Fringed sage is probably the most widely 

distributed and abundant species of Artemisia L. 

Its range extends from Mexico in the south, 

through most of the Western United States, 

Western Canada, and Alaska in the north, and into 

Siberia, Mongolia, and Kazakhstan (Harvey 1981; 

USDA 1937). In the United States, it is most 

abundant in the eastern and northern parts of its 

range on the high plains along the eastern slope of 

the Rocky Mountains. The altitudinal gradient of 

fringed sage in the Intermountain area extends 

from 900 to 3,500 m; in other parts of its range 

(Great Plains and Asian Steppes) the lower 

elevational limits are somewhat less. 

Ecology.—Mean annual precipitation throughout 

the native range of fringed sage varies between 

210 and 550 mm. It inhabits a wide variety of sites 

but typically grows in full sunlight on dry, coarse, 

shallow soils. On winter livestock and game 

ranges in western Utah and eastern Nevada, 

fringed sagebrush may grow in dense stands along 

shallow depressions that collect moisture from 

summer rains. In such areas, it is frequently 

associated with winterfat [Ceratoides lanata

(Pursh) J. T. Howell], shadscale [Atriplex 

confertifolia (Torr. & Frem.) Wats.], and 

rabbitbrushes (Chrysothamnus Nutt. species). On 

plains, foothills, and mountain slopes, fringed sage 
64 

may grow in association with a variety of grasses 

and forbs as well as with various shrubs, including 

big sagebrush (Artemisia tridentata Nutt.), 

Bigelow sagebrush (Artemisia bigelovii Gray), 

sandsage (Artemisia filifolia Torr.), and especially 

in overgrazed areas, with broom snakeweed 

[Gutierrezia sarothrae (Pursh) Britt. & Rushby]. 

It is also a common understory shrub in ponderosa 

pine communities in several Western States 

(McArthur and others 1979). 

Reproduction.—Blooming occurs from June at 

high elevations and latitudes, to November at 

lower elevations and latitudes. Fringed sage is a 

prolific seed producer; each 2.5 cm length of 

inflorescence contains approximately 1,000 seeds 

(Harvey 1981). There are about 10 million cleaned 

seed per kg (Plummer and others 1968). Seed 

matures between September and December 

(McArthur and others 1979) and are wind 

dispersed in winter. Seedlings emerge in spring. 

Fringed sage can be established by seed, 

transplanting young plants, or from rooted 

segments of mature plants. Surface seeding on 

disturbed soils is recommended (Wasser 1982). 

Management.—The forage value of fringed sage 

varies considerably with location and season 

(Dietz 1972). On Western ranges, it is most 

valuable to big game and livestock in late fall, 

winter, and early spring (Cooperrider and Bailey 

1986, Kufeld 1973, USDA 1937, Wasser 1982). 

Nutritive quality is highest in the spring but 

remains adequate throughout much of the year 

(Cooperrider and Bailey 1986, Rauzi 1982). 

Individual plants are susceptible to overgrazing, 

but populations tend to increase under heavy 

grazing pressure (Cooperrider and Bailey 1986). 

In the northwestern Great Plains, fringed sage may 

be less palatable than other available forages and 

occasionally invades deteriorated grasslands. 

Pronghorn (Antilocapra Americana Ord) utilize 

fringed sage extensively during winter, and to a 

lesser extent, year round (McArthur and others 

1979). 

 Fringed sage is well suited to grass and 

broad-leafed herb communities. Seedlings are 

competitive and can establish in areas with 

herbaceous competition. It is adapted to mine 

spoils, perhaps better than any other species of 

Artemisia. It can be used as a biological control to 

reduce rapid weed expansion on large disturbed 

sites but can be controlled when it is too abundant 

(Alley 1972). 

Benefits.—Fringed sage is a good pioneer shrub 

for stabilizing disturbed sites. It is often used in 

seeding Western strip mines, especially coal areas. 

Its strong taproot and numerous lateral roots help 

stabilize gullies and reduce soil erosion. These 

rooting characteristics enable the shrub to resist 

considerable grazing and trampling. It also 

provides forage for livestock and forage and 

habitat for wildlife in both North America and 

Asia (Li and others 2002, USDA 1937). It is an 

important food for sage grouse (Wallestad and 

others 1975). Fringed sage has some value as a 

medicinal plant (Hall and Clements 1923). Native 

Americans traditionally used fringed sagebrush as 

toilet paper, to eliminate unpleasant odors from 

dried meat, as a chewed application for wounds 

prior to bandaging, and to make mats and fans 

(Stubbendieck and others 1997). 

References 

Alley, H. 1972. Range weed control. In: 

Proceedings 1970 Washington State Weed 

Control Conference, Yakima, WA. Washington 

State Weed Association. p. 39-45. 

Cooperrider, A.Y. and J.A. Bailey. 1986. Fringed 

sagebrush (Artemisia frigida)—a neglected 

forage species of Western ranges. In: McArthur, 

E.D. and B.L Welch, comps. Proceedings—

symposium on the biology of Artemisia and 

Chrysothamnus; 1984 July 9-13, Provo, UT. 

General Technical Report INT-200. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Research Station, Ogden, UT. p. 

46-54. 

Dietz, D.R. 1972. Nutritive value of shrubs. In: 

McKell, C.M., J.P. Blaisdell, and J.R. Goodin, 

tech. eds. Wildland shrubs—their biology and 

management; 1971 July, Logan, UT. General 

Technical Report INT-1. U.S. Department of 

Agriculture, Forest Service, Intermountain Forest 

and Range Experiment Station, Ogden, UT. p. 

289-302. 

Hall, H.M. and F.E. Clements. 1923. The 

phylogenetic method in taxonomy, the North 

American species of Artemisia, Chrysothamnus, 

and Atriplex. Publication 326. Carnegie 

Institution of Washington, Washington, DC. 

355 p. 

Harvey, S.J. 1981. Life history and reproductive 

strategies in Artemisia. M.S. thesis, Montana 

State University, Bozeman, MT. 132 p. 
65

International Plant Plant Names Index. 2003. 

http//www.ipni.org.) [not paged]. 

Kufeld, R.C. 1973. Foods eaten by the Rocky 

Mountain elk. Journal of Range Management 

26: 106-113. 

Li, F.R., A.f. Zhao, H.Y. Zhou, T.H. Zhang, and 

X. Zhao. 2002. Effects of simulated grazing on 

growth and persistence of Artemisia frigida in a 

semiarid sandy rangeland. Grass and Forage 

Science 57: 239-246. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 82 p. 

McArthur, E.D. and C.L. Pope. 1979. Karyotypes 

of four Artemisia species: A. carruthii, A. 

filifolia, A. frigida, and A. spinescens. Great 

Basin Naturalist 39: 419-426. 

Plummer, A.P., D.R. Christensen, and S.B. 

Monsen. 1968. Restoring big-game range in 

Utah. Publication 68-3. Utah Division of Fish 

and Game, Salt Lake City, UT. 183 p. 

Rauzi, F. 1982. Seasonal variations in protein and 

mineral content of fringed sagewort (Artemisia 

frigida). Journal of Range Management 35: 679-

680. 

Stubbendieck, J., S.L. Hatch, and C.H. 

Butterfield, 1997. North American Range 

Plants. Fifth Edition. University of Nebraska 

Press, Lincoln, NB. 233 p. 

U. S. Department of Agriculture, Forest Service. 

1937. Range plant handbook. United States 

Government Printing Office, Washington, DC. 

816 p. 

Wallestad, R., J.C. Peterson, and R.L. Eng. 1975. 

Foods of adult sage grouse in central Montana. 

Journal of Wildlife Management 39: 628-630. 

Wasser, C.H. 1982. Ecology and culture of 

selected species useful in revegetation disturbed 

lands in the west. FSW/OBS-82/56. U. S. 

Department of the Interior, Fish and Wildlife 

Service. Washington, DC. 347 p. 
66 

Artemisia longiloba (Osterhout) Beetle alkali sagebrush 

ASTERACEAE 

Synonyms: Artemisia spiciformis Osterhout var. longiloba Osterhout 

 Artemisia arbuscula Nutt. ssp. longiloba (Osterhout) L. Shultz 

E. Durant McArthur and Jeffrey R. Taylor 

General Description.—Alkali sagebrush, also 

known as early sagebrush and low sagebrush, is a 

small shrub up to 45 cm tall. It has lax, spreading 

stems that frequently layer. The bark is dark 

brown to black on older stems. The whole plant 

has a dark gray-green appearance (Beetle 1960). 

Leaves on vegetative stems are broadly cuneate, 

up to 2 cm long, and deeply three-lobed. Leaves 

on flowering stems are similar in shape but 

smaller. Its large heads and early blooming 

readily distinguishes alkali sagebrush from other 

small sagebrushes (Beetle 1959). Heads contain 

six to 11 disc flowers that are 3 to 5 mm wide. 

Disc flowers of similarly sized sagebrushes, such 

as low sagebrush (Artemisia arbuscula Nutt.) and 

black sagebrush (Artemisia nova A. Nelson), are 3 

mm wide or less. Beetle (1960) points out that this 

species has been confused with silver sagebrush 

(Artemisia cana Pursh) because of its large heads; 

with big sagebrush (Artemisia tridentata Nutt.) 

because of its broadly cuneate, three-lobed leaves; 

and with low sagebrush because of its diminutive 

size. 

Taxonomy.—The taxon includes no recognized 

subspecies or varieties. Although relegated to 

subspecific status by some, e. g., the U. S. 

Department of Agriculture, Natural Resource 

Conservation Service plant data base 2003, we 

prefer to maintain its specific status. Zamora and 

Tueller (1974) differentiated it from typical low 

sagebrush (Artemisia arbuscula Nutt.) in 

ecological habitat preference. Its large, scattered, 

distribution and its differential use by animals 

suggest that it harbors considerable genetic 

variation. Beetle (1960) suspected that it is a 

parent to the narrowly endemic coaltown 

sagebrush (Artemisia argillosa Beetle). Alkali 

sagebrush has both diploid (2n = 18) and tetraploid 

(2n = 36) populations (McArthur and others 1981). 

Range.—Alkali sagebrush is found along foothills 

and basins of the ranges that form the Continental 

Divide in southwestern Montana, south through 

Wyoming to northwestern Colorado, and scattered 

westward through northern Utah and Idaho to 

Nevada and Oregon (Beetle 1960, Winward 1980). 

Ecology.—Despite the common name of alkali 

sagebrush, the species also often grows well on 

neutral soils (Passey and Hugie 1962, Tisdale and 

Hironaka 1981). It is well adapted to xeric 

conditions (Robertson and others 1966, Zamora 

and Tueller 1973). Mean annual precipitation 

throughout its range can vary between 220 and 500 

mm. Alkali sagebrush typically grows in heavy, 

highly impermeable soils with dense “B” horizons 

(Passey and Hugie 1962, Tisdale and Hironaka 

1981). It does, however, grow on lighter soils as 

well (McArthur and others 1979). 
67

Reproduction.—Shrubs usually bloom in midJune to early August and seed ripens in August 

and September. Seeds are wind dispersed during 

late fall and winter, and seedlings emerge during 

the following spring. Seeds are about the size of 

low sagebrush seeds so we expect there would be 

about 2 million cleaned seeds per kg (Meyer 

2003). Natural seeding occurs rapidly following 

fires and other disturbances, yet mechanical tillage 

can limit establishment if seeds are sown too deep 

or if the seedbed is disrupted (Monsen and Shaw 

1986). 

Management.—Alkali sagebrush, depending on 

location and ecotype, can be a source of palatable 

browse for domestic sheep and provide habitat and 

forage for other animals (Dealy and others 1981, 

McArthur and others 1979). Stands of alkali 

sagebrush in good condition can support a diverse 

understory of perennial grasses and annual and 

perennial forbs. Such stands provide forage and 

cover for sage grouse, antelope, other wildlife, and 

domestic livestock (Dealy and others 1981, 

Monsen and Shaw 1986). Some stands, however, 

are much less diverse and provide little forage or 

habitat. Sites that lack a satisfactory understory 

remain closed unless shrubs are reduced by fire or 

other disturbance. Monsen and Shaw (1986) 

reported that decadent stands can be rejuvenated 

by mechanical means. 

This shrub has the potential to be a useful plant in 

rehabilitation, especially on heavy, seasonally 

dry soils. It has been observed invading 

roadcuts in Echo Canyon, Utah, and along 

roadsides near Kemmerer, Wyoming (McArthur 

and others 1979). 

Benefits.—Alkali sagebrush is a native shrub that 

provides habitat and forage for wildlife and 

livestock. In addition, it has characteristics that 

could make it useful in ecological restoration, 

rehabilitation, or soil conservation. It spreads well 

through natural seeding and tolerates disturbance. 

It also grows on argillic soils that may be too 

heavily textured for most other shrubs. 

References 

Beetle, A.A. 1959. New names within the section 

Tridentatae of Artemisia. Rhodora 61: 82-85. 

Beetle, A.A. 1960. A study of sagebrush, the 

section Tridentatae of Artemisia. Bulletin 368, 

University of Wyoming Agriculture Experiment 

Station, Laramie, WY. 83 p. 

Dealy, J.E., D.A. Leckenby, and D.M. Concannon. 

1981. Wildlife habitats in managed rangelands: 

The Great Basin of southeastern Oregon. Plant 

communities and their importance to wildlife. 

General Technical Report PNW-120. U.S. 

Department of Agriculture, Forest Service, 

Pacific Northwest Forest and Range Experiment 

Station, Portland, OR. 66 p. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 82 p. 

McArthur, E.D., C.L. Pope, and D.C. Freeman. 

1981. Chromosomal studies of subgenus 

Tridentatae of Artemisia: evidence for 

autopolyploidy. American Journal of Botany 68: 

589-605. 

Meyer, S.E. 2003. Artemisia. In: Bonner, F.T., and 

R. Nisley, eds. Woody plant seed manual. 

http://www.wpsm.net. [not paged]. 

Monsen, S.B. and N.L. Shaw. 1986. Response of 

an alkali sagebrush/fescue site to restoration 

treatments. In: McArthur, E.D. and B.L Welch, 

comps. Proceedings—symposium on the biology 

of Artemisia and Chrysothamnus; 1984 July 9-

13, Provo, UT. General Technical Report INT200. U.S. Department of Agriculture, Forest 

Service, Intermountain Research Station, Ogden, 

UT. p. 126-133. 

Passey, H.B. and V.K. Hugie. 1962. Sagebrush on 

relic ranges in the Snake River plains and 

northern Great Plains. Journal of Range 

Management. 15: 273-278. 

Robertson, D.R., J.L. Nielsen, and N.H. Bare. 

1966. Vegetation and soils of alkali sagebrush 

and adjacent big sagebrush ranges in North Park, 

Colorado. Journal of Range Management. 19: 

17-20. 

Tisdale, E.W. and M. Hironaka. 1981. The 

sagebrush—grass region: a review of the 

ecological literature. Bulletin No. 33, Forest, 

Wildlife, and Range Experiment Station, 

University of Idaho, Moscow, ID. 31 p. 
68 

U. S. Department of Agriculture, Natural 

Resources Conservation Service—

http//www.plants.usda.gov. Last visited August 

25, 3003. 

Winward, A.H. 1980. Taxonomy and ecology of 

sagebrush in Oregon. Bull. 642. Oregon State 

University, Oregon Agricultural Experiment 

Station, Corvallis, OR. 15 p. 

Zamora, B. and P.T. Tueller. 1973. Artemisia 

arbuscula, A. longiloba, and A. nova habitat 

types in northern Nevada. Great Basin Naturalist. 

33: 225-242. 
69

Artemisia ludoviciana Nutt. Lousiana sagewort 

ASTERACEAE 

Synonyms: Artemisia albula Wooton 

 Artemisia candicans Rydb. 

 Artemisia gnaphalodes Nutt. 

 Artemisia incompta Nutt. 

 Artemisia mexicana Willd. ex Spreng. 

Artemisia redolens A.Gray 

Artemisia sulcata Rydb. 

 Artemisia vulgaris var. americana Besser 

 Artemisia vulgaris ssp. ludoviciana H.M. Hall & Clem. 

 Artemisia vulgaris var. ludoviciana Kuntze 

Jeffrey R. Taylor and E. Durant McArthur 

 

Illustration credit: USDA—Forest Service collection, 

Hunt Institute 

General Description.—Louisana sagewort, also 

known as cudweed sagewort, Louisiana sagebrush, 

Louisiana wormwood, western mugwort, white 

sage, sage-weed, estafiate, and iztafiate, is an 

aromatic, morphologically variable, herbaceous 

forb or sub-shrub 0.3 to 1 m tall. Stems are erect, 

single to numerous, and woody near the base. 

Stems can be grayish-green, greenish-yellow, or 

green and are typically covered with a white to 

gray tomentum. Leaves may be entire, tridentate, 

variously-lobed oblanceolate, or linear. Lower 

leaves are grayish green, oblanceolate, lobed, or 

entire, 3 to 9 cm long and 0.5 to 2.5 cm long, while 

upper leaves are mostly entire, gray to green 

floccose to glabrous above, and white tomentose 

beneath (Hall and Clements 1923). The leaf apex 

is acute. Flower heads bear six to 12 pistilate ray 

flowers and five to 12 perfect disc flowers, 

arranged in elongate, leafy panicles 20 to 50 cm 

long (Stubbendieck and others 1997). Louisana 

sagewort has a strong tendency to spread through 

rhizomes. 

Taxonomy.—Artemisia ludoviciana is a highly 

polymorphic species. Hall and Clements (1923) 

chose to classify it within the Artemisia vulgaris 

L. complex. This system of classification 

consolidated 54 previously recognized species into 

one species and 15 subspecies. Keck (1946) 

suggested that the complex arose from one 

cenospecies, and subsequently revised the 

taxonomy. Keck’s revision classified A. 

ludoviciana on the specific, rather than subspecific 

level, with seven subspecies of its own. These 

subspecies include: typica, sulcata (Rydb.) Keck, 

albula (Woot.) Keck, candicans (Rydb.) Keck, 

incompta (Nutt.) Keck, mexicana (Wild.) Keck, 

and redolens (Gray) Keck. Cytogenetic studies 

show that the New World members within the 

vulgaris complex have a base chromosome 

number x = 9, as opposed to the Old World 

members, with a base chromosome number of x = 

8. Lousiana sagewort and its close relatives (A. 

ludoviciana species complex) can be diploid, 

tetraploid, or hexaploid (Estes 1969, Welsh and 
70 

others 1993). Estes (1969) suggested that 

autopolyploidy has been the driving evolutionary 

force within the complex. Molecular genetic 

studies have also shown New World and Old 

World members of the complex to be different 

from one another (Vallès and McArthur 2001). 

Range.—Lousiana sagewort is widely distributed 

throughout Western North America. Its range 

spans from Ontario and Michigan in the east, to 

British Columbia in the west, and south through 

Texas, Louisiana, California, and Mexico (USDA 

Forest Service 1937). Lousiana sagewort grows at 

various elevations from 250 to 3,050 m. 

Ecology.—Lousiana sagewort grows on rocky, 

sandy, or gravelly loams on open ridges, slopes, 

and mesas (USDA Forest Service 1937). It is also 

common on plains and prairies and is usually a late 

seral species that grows after grasses have declined 

(Hall and Clements 1923). Lousiana sagewort 

grows rapidly, is long lived, and is cold and 

drought tolerant (Shaw and Monsen 1982). It 

grows well in sunny sites even though it is shade 

tolerant. It is can be found in grasslands, 

sagebrush, ponderosa pine, and alpine 

communities (Shaw and Monsen 1982). Lousiana 

sagewort often associates with asters (Aster L.), 

goldenrods (Solidago L.), sunflowers (Helianthus 

L.), fringed sagebrush (Artemisia frigida Wild.), 

big sagebrush (Artemisia tridentata Nutt.), 

tarragon (Artemisia dracunculus L.), needlegrass 

(Stipa L.), blue grama [Bouteloua gracilis 

(H.B.K.) Lag. ex Steudel], muhly grasses 

(Muhlenbergia Schreber), wheatgrasses 

(Agropyron Gaertner), mountain dandelions 

[Agoseris glauca (Pursh) Raf.], and western 

yarrow (Achillea millefolium L.) (USDA Forest 

Service 1937, Hall and Clements 1923). Mean 

annual precipitation within its range varies 

between 400 and 1200 mm. 

Reproduction.—Blooming begins in August and 

ends in September (Stubbendieck and others 

1997). Seed matures in the fall and is dispersed by 

wind during late fall or winter. Seeds are 

relatively small, numbering about 8,360,000/kg 

(personal communication with S. Stranathan, 

Upper Colorado Environmental Plant Center, 

Meeker, CO). Seed viability decreases after 2 or 3 

years of storage (personal communication with 

K.R. Jorgensen, Great Basin Experimental Area 

Ephraim, UT). Seeding aerially, by drill, or 

broadcasting are all effective methods of planting 

and should be conducted during fall or winter 

(Shaw and Monsen 1982). In addition to seeds, 

Lousiana sagewort spreads asexually through 

rhizomes. 

Management.—Lousiana sagewort is used 

differentially by livestock and wildlife throughout 

its distribution. Populations in the southern 

portions of its range, such as in southern Utah, 

New Mexico, and southern Colorado, are more 

palatable than those in the north (USDA Forest 

Service 1937, Stubbendieck and others 1997). In 

these southern areas, it can provide valuable 

forage, especially on spring-fall and winter ranges. 

In more northern areas, it has a pungent taste and 

is generally not eaten until after the first frost (Hall 

and Clements 1923). Lousiana sagewort is 

generally considered poor forage, although it is 

utilized by domestic sheep, deer, and elk to some 

degree, and may be seasonally important (Shaw 

and Monsen 1982, Hall and Clements 1923, 

USDA Forest Service 1937). Lousiana sagewort 

has also been used to revegetate road, logging, and 

other disturbances (Shaw and Monsen 1982). In a 

revegetation experiment at Mesa Verde National 

Park, CO, it survived well when transplanted in 

roadcut sites (Paschke and others 1999). 

Benefits.—Lousiana sagewort is an extensive and 

genetically diverse native plant that is widespread 

throughout Western North America. It provides 

quick and excellent cover, reduces erosion, and 

serves as a nurse plant for other species (Shaw and 

Monsen 1982). It also has pharmaceutical 

potential. Zavala-Sanchez and others (2002) 

found that an essential oil found in Lousiana 

sagewort, nonanal, inhibited diarrhea in mice. In 

addition, Lousiana sagewort extracts inhibited 

reproduction in malarial parasites (Malagon and 

others 1997). The ssp. mexicana (estafiate) is used 

for gastrointestinal pain, as a verimfuge, and as a 

bitter stimulant (Heinrich 2002). It has also been 

used by Native Americans in ceremonial and 

purification rituals, to deodorize feet, as pillows, 

saddle pads, and to treat headaches, stomach 

ailments, and coughs (Stubbendieck and others 

1997). 

References

Estes, J.R. 1969. Evidence for autoploid 

evolution in the Artemisia ludoviciana complex 

of the Pacific Northwest. Brittonia 21: 29-43. 

Hall, H.M. and F.E. Clements. 1923. The 

phylogenetic method in taxonomy, the North 

American species of Artemisia, Chrysothamnus, 

and Atriplex. Publication no. 326, Carnegie 
71

Institution of Washington, Washington, DC. 

355 p. 

Heinrich, M. 2002. Ethnobotany, phytochemistry 

and biological/pharmacological activities of 

Artemisia ludoviciana ssp. mexicana (estafiate). 

In: Wright, C.W., ed. Artemisia, medicinal and 

aromatic plants Vol. 18. Taylor and Francis, 

London, UK. p. 107-117. 

Keck, D.D. 1946. A revision of the Artemisia 

vulgaris complex in North America. 

Proceedings of the California Academy of 

Sciences 25: 421-468. 

Malagon, F., J. Vasquez, and A. Ruiz. 1997. 

Antimalaric effect of an alchoholic extract of 

Artemisia ludoviciana mexicana in a rodent 

malaria model. Parassitologia 39: 3-7. 

Paschke, M.W., C. DeLeo, and E.F. Redente 1999. 

Revegetation of roadcut slopes in Mesa Verde 

National Park, U.S.A Restoration Ecology 8: 

263-268. 

Shaw, N. and S.B. Monsen. 1982. 

Nonleguminous forbs for rangeland sites. In: 

Monsen, S.B. and N. Shaw, comps. 

Proceedings: Managing Intermountain 

Rangelands—Improvement of range and 

wildlife habitats; September 15-17, 1981; Twin 

Falls, ID, June 22-24, 1982; Elko, NV. Gen. 

Tech. Rep. INT-157. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Research Station, Ogden, UT. p. 123-131. 

Stubbendieck, J., S.L. Hatch, and C.H Butterfield. 

1997. North American Range Plants. Fifth 

edition. University of Nebraska Press, Lincoln, 

NB. 235 p. 

U.S. Department of Agriculture, Forest Service. 

1937. Range plant handbook. United States 

Government Printing Office, Washington, DC. 

816 p. 

Vallès, J. and E.D. McArthur. 2001. Artemisia

systematics and phylogeny: cytogenetic and 

molecular insights. In: McArthur, E.D. and D.J. 

Fairbanks, comps. Shrubland ecosystem genetics 

and biodiversity: proceedings. 2000 June 13-15, 

Provo, UT. Proceedings RMRS-P-21. U.S. 

Department of Agrigulture, Forest Service, 

Rocky Mountain Research Station, Ogden, UT. 

p. 67-74. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1993. A Utah Flora. Second 

edition. Printing Services, Brigham Young 

University, Provo, UT. 986 p. 

Zavala-Sanchez, M.A., S. Perez-Gutierrez, C. 

Perez-Conzalez, and D. Sanchez-Saldivar. 

2002. Antidiarrhoeal activity of nonanal, an 

aldehyde isolated from Artemisia ludoviciana. 

Pharmaceutical Biology 40: 263-268. 
72 

Artemisia nova A. Nels. black sagebrush 

ASTERACEAE 

Synonyms: Artemisia tridentata ssp. nova (A. Nels.) H.&C. 

 Artemisia arbuscula ssp. nova (A. Nels.) McMinn 

 Artemisia arbuscula var. nova (A. Nels.) G. Ward 

 Serphidium novum (A. Nels.) W.A. Weber 

E. Durant McArthur and Jeffrey R. Taylor 

 

General Description.—Black sagebrush is a small 

aromatic shrub generally 15 to 20 cm tall, although 

it may occasionally exceed 75 cm. A dull grayishtomentose layer causes most populations of black 

sagebrush to appear darker than those of big 

sagebrush or low sagebrush. Branches are 

numerous and erect and arise from a spreading 

base. Leaf base is typically cuneate, and the leaf 

surface is covered with a viscid glandular 

pubescence. The leaf apex is three-toothed, 

although the uppermost leaves, particularly on 

flowering stems, may be entire. Leaf dimensions 

can vary between 0.5 to 2 cm long and 0.2 to 0.8 

cm wide. Flowering heads are grouped in narrow 

spike-like panicles that commonly extend above 

the foliage. Heads contain three to five disc 

flowers subtended by eight to 12 greenish-yellow 

involucral bracts. Leaf hairs, visible at 10x 

magnification, can be useful diagnostic clues to 

distinguish black sagebrush from other 

sagebrushes. 

Taxonomy.—Black sagebrush taxonomy is 

relatively simple. It includes one species and two 

varieties, A. nova var. duchesnicola and A. nova

var. nova (Welsh and Goodrich 1995). Beetle and 

Johnson (1982) have found forms of black 

sagebrush that are similar in appearance to 

Wyoming big sagebrush. This, in addition to other 

factors, has led some to suspect that black 

sagebrush may have been involved in the 

parentage of Wyoming big sagebrush (McArthur 

1983, Winward 1976). Glandular trichomes are a 

taxonomic feature that helps separate black 

sagebrush from big sagebrush (A. tridentata Nutt.) 

and low sagebrush (A. arbuscula Nutt.) (Kelsey 

1984). Black sagebrush has a base chromosome 

number of x = 9; both diploid (2n = 18) and 

tetraploid populations are known. About twothirds of the cytologically known populations are 

tetraploid (McArthur and Sanderson 1999). 

Range.—Black sagebrush is one of the most 

common shrubs in the Western United States. 

Populations can be found from California and 

Oregon in the west, to Colorado in the east, and 

from Montana in the north to northern Arizona and 

New Mexico in the south. Black sagebrush is most 

common at elevations ranging between 1,500 to 

2,400 m (McArthur and others 1979) 

Ecology.—Black sagebrush is well adapted to dry 

stony soils that are relatively shallow. Zamora and 

Tueller (1973) found root restricting layers 28 to 

69 cm deep in half of their black sagebrush study 

sites. Gravelly or sandy loam strata are also 

common subsurface soil profile characteristics in 

areas with black sagebrush. In addition, most soils 

supporting black sagebrush are calcareous. Black 

sagebrush is not restricted to adverse 

environments, however. If seeded, it can grow well 
73

on sites normally occupied by big sagebrush. 

Mean annual precipitation throughout the natural 

range of black sagebrush ordinarily is between 180 

and 320 mm. 

Most stands of black sagebrush do not 

burn because populations are relatively sparse 

(Tisdale and Hironaka 1981). Exceptions to this 

rule, however, have been observed in central Utah, 

where large stands of black sagebrush burned. In a 

comparative study, Nelson and Krebill (1981) 

found that black sagebrush is less susceptible to a 

wilt disease than other species of Artemisia grown 

under similar conditions. 

Reproduction.—Mature shrubs flower in late 

summer (usually August), and seed matures in 

September and October. Black sagebrush seed 

tends to be larger than those of other sagebrushes. 

There are about 2,000 seeds/g (Deitschman 1974). 

Seeds are wind dispersed in late fall or early 

winter. Seedlings emerge in the spring and often 

grow rapidly. Once established, young plants can 

persist well even under adverse conditions. With 

favorable moisture conditions, mature plants 

produce abundant seed, and natural spreading 

occurs quickly. 

Management.—Black sagebrush is usually 

considered a valuable browse species for wildlife 

and livestock, especially for deer, domestic sheep, 

and pronghorn (Clary 1986; McArthur and 

Plummer 1978, USDA Forest Service 1937). 

Black sagebrush has good winter nutritive value, 

but not as good as big sagebrush (Welch 1983). 

Palatability, however, can vary significantly 

between populations (Welch and others 1981). 

Deer and elk preference studies for black and big 

sagebrushes have yielded inconsistent results 

(Scholl and others 1977, Nagy and Regelin 1977, 

Smith 1950, Wambolt 1996). Welch and others 

have shown that some accessions are not eaten 

while others are highly preferred (Behan and 

Welch 1985, Welch and others 1981), which may 

partially explain differences in preference studies. 

Black sagebrush can also be a valuable 

conservation species for dry, shallow, stony soils 

because it produces abundant seed and seedlings 

establish readily. Due to its habitat, it is not 

usually a candidate for plant control. Black 

sagebrush can successfully be seeded in fall or 

early winter. Seeds should be covered with soil, 

but remain within 6 mm of the surface. It grows 

well with seeded herbs, but understory production 

tends to decrease as the stand matures. 

Benefits.—Black sagebrush is an important native 

shrub in many areas of the Western United States. 

It provides structural diversity within native plant 

communities and provides an important browse 

source for native and introduced ungulates. It can 

also reduce soil erosion on steep shallow slopes. 

‘Gordon Creek’ is a tested germplasm from 

Carbon County, Utah, that has been demonstrated 

to be high in nutritive quality and palatability to 

wildlife (Welch and others 1994). 

References 

Beetle, A.A. and K.L. Johnson. 1982. Sagebrush in 

Wyoming. Bulletin 779. Agricultural 

Experiment Station, University of Wyoming, 

Laramie, WY. 68 p. 

Behan, B. and B.L. Welch. 1985. Black sagebrush: 

mule deer winter preference and monoterpenoid 

content. Journal of Range Management 38: 278-

280. 

Clary, W.P. 1986. Black sagebrush response to 

grazing in the east-central Great Basin. In: E.D. 

McArthur and B.L. Welch, comps. Proceedings: 

symposium on the biology of Artemisia and 

Chrysothamnus; 1984 July 9-13; Provo, UT. 

General Technical Report INT-200. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Research Station, Ogden, UT: 

181-185 p. 

Deitschman, G.H. 1974. Artemisia. In: C.S. 

Schopmeyer, technical coordinator, Seeds of 

Woody Plants of the United States. Agriculture 

Handbook 450. U.S. Department of Agriculture, 

Forest Service, Washington, DC. 235-237 p. 

Kelsey, R.G. 1984. Glandular trichomes: a helpful 

taxonomic character for Artemisia nova (black 

sagebrush). Journal of Range Management 37: 

370-372. 

McArthur, E.D. 1983. Taxonomy, origin, and 

distribution of sagebrush (Artemisia tridentata) 

and allies (subgenus Tridentatae). In: K.L. 

Johnson, ed. Proceedings of the First Utah 

Shrub Ecology Workshop; 1981 September 9-

10; Ephraim, UT. College of Natural Resources, 

Utah State University, Logan, UT. 3-13 p. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 
74 

U.S. Department of Agriculture, Forest Service 

USDA Forest Service, Intermountain Forest and 

Range Experiment Station, Ogden, UT. 82 p. 

McArthur, E.D. and A.P. Plummer. 1978. 

Biogeography and management of native 

western North American shrubs: a case study, 

section Tridentatae of Artemisia. Great Basin 

Naturalist Memoirs 2: 229-241. 

McArthur, E.D. and S.C. Sanderson. 1999. 

Cytogeography and chromosome evolution of 

subgenus Tridentatae of Artemisia (Asteraceae). 

American Journal of Botany 86: 1,754-1,775. 

Nagy, J.G. and W.L. Regelin. 1977. Influence of 

plant volatile oils on food selection by animals. 

Congress of Game Biologists 13: 225-230. 

Nelson, D.L. and R.G. Krebill. 1981. A sagebrush 

wilt disease of unknown origin. Great Basin 

Naturalist 41: 184-191. 

Smith, A.D. 1950. Sagebrush as a winter feed for 

deer. Journal of Wildlife Management 14: 285-

289. 

Scholl, J.P., R.G. Kelsey, and F. Shafizadeh. 1977. 

Involvement of volatile compounds of Artemisia

in browse preference by mule deer. Biochemical 

Systematics and Ecology 5: 291-295. 

Tisdale, E.W. and M. Hironaka. 1981. The 

sagebrush—grass region: a review of the 

ecological literature. Bulletin 33, Forest, 

Wildlife, and Range Experiment Station, 

University of Idaho, Moscow, ID. 31 p. 

U.S. Department of Agriculture, Forest Service. 

1937. Range plant handbook. United States 

Government Printing Office, Washington, DC. 

816 p. 

Wambolt, C.L. 1996. Mule deer and elk foraging 

preference for 4 sagebrush taxa. Journal of 

Range Management 49: 499-503. 

Welch, B.L. 1983. Improving the nutritive value of 

winter range forage. In: S. B. Monsen and N. 

Shaw, comps. Managing Intermountain 

rangelands—improvement of range and wildlife 

habitats: proceedings of a symposium; 1981 

September 15-17; Twin Falls, ID; 1982 June 22-

24, Elko, NV. General Technical Report INT157, U.S. Department of Agriculture, Forest 

Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT: 158-164. 

Welch, B.L., E.D. McArthur, and J.N. Davis. 

1981. Differential preference of wintering mule 

deer for accessions of big sagebrush and black 

sagebrush. Journal of Range Management 34: 

409-411. 

Welch, B.L., E.D. Nelson, S.A. Young, A.R. 

Sands, F.J. Wagstaff, and D.L. Nelson. 1994. 

‘Gordon Creek’—a superior, tested germplasm 

of Wyoming big sagebrush. Research Paper INT461, U.S. Department of Agriculture, Forest 

Service, Intermountain Research Station. Ogden, 

UT. 7 p. 

Welsh, S.L. and S. Goodrich. 1995. Plant novelties 

in Lepidium (Cruciferae) and Artemisia

(Compositae) from the Uinta Basin, Utah. Great 

Basin Naturalist 55: 359-362. 

Winward, A.H. 1976. Evolutionary development 

of the Artemisia tridentata taxa. In: H. C. Stutz, 

ed. Wildland shrubs: proceedings, symposium, 

and workshop; 1975 November 4-6, Provo, UT. 

College of Biology and Agriculture, Brigham 

Young University, Provo, UT: 163. 

Zamora, B. and P.T. Tueller. 1973. Artemisia 

arbuscula, A. longiloba, and A. nova habitat 

types in northern Nevada. Great Basin Naturalist 

33: 225-242. 
75

Artemisia pygmaea Gray pygmy sagebrush 

ASTERACEAE 

Synonyms: Seriphidium pygmaeum (Gray) W.A.Weber 

E. Durant McArthur and Jeffrey R. Taylor 

General Description.—Pygmy sagebrush is a 

dwarf, depressed, evergreen, cushion-like shrub 

less than 20 cm tall. Bark on older stems becomes 

dark brown and fibrous with age. Bark on younger 

branches is nearly white to straw-colored and 

somewhat puberulent. Leaves on vegetative stems 

are green, nearly glabrous, viscidulous, and are 

pinnatified with three to 11 lobes, or sometimes 

may be toothed. Leaf dimensions range between 2 

to 4 mm wide and 2 to 8 mm long. Leaves on 

flowering branches are usually reduced and may 

be entire. Flower heads contain three to five disc 

flowers and are arranged into spike-like 

inflorescences. Ray flowers are lacking. Twelve 

to 18 greenish-yellow involucral bracts subtend 

each head (Cronquist 1994, McArthur and others 

1979). 

Taxonomy.—No subspecific entities have been 

described for pygmy sagebrush. Pygmy sagebrush 

is known only at the diploid, 2n = 18, chromosome 

level (McArthur and Sanderson 1999). 

Range.—The distribution of pygmy sagebrush is 

limited to approximately 54 km2

 in eastern Utah, 

western Nevada, and northern Arizona (Beetle 

1960; McArthur and Plummer 1978; Ward 1953). 

Ecology.—Pygmy sagebrush grows on calcareous 

desert soils. Mean annual precipitation within its 

range is approximately 200 to 300 mm. In 

Nevada, this species is often associated with 

halophytic threadleaf rubber rabbitbrush 

[Chrysothamnus nauseosus (Pallas ex Pursh) 

Britton ssp. consimilis (Greene) Hall & Clements]. 

In Utah, some relatively large stands are mixed 

with black sagebrush (Artemisia nova A. Nelson). 

Reproduction.—Blooming occurs between 

August and September. Seed matures in October. 

Seeds are relatively large compared to those of 

other subgenus Artemisia subgenus Tridentatae

species (McArthur and others 1979). Seeds are 

wind dispersed during late fall or winter. 

Seedlings emerge during spring. Shrubs in 

research plantings have spread naturally. It also 

establishes well by transplanting divided plants. 

Although it spreads through naturally dispersed 

seed, artificial seedings have been unsuccessful. 

Management.—Pygmy sagebrush provides little 

browse because of its scarcity and size. It is 

readily eaten, however, when available. It is a 

good candidate to plant in clay soils and may be of 

considerable value for revegetating mine spoils 

and roadway disturbances. 
76 

Benefits.—Although it is not abundant, pygmy 

sagebrush does provide important soil cover in 

dry, alkaline habitats where little else grows. It 

also adds to the overall biological diversity to the 

shrub communities where it is found. 

References 

Beetle, A.A. 1960. A study of sagebrush, the 

section Tridentatae of Artemisia. Bulletin 368, 

University of Wyoming Agriculture Experiment 

Station, Laramie, WY. 83 p. 

Cronquist, A. 1994. Asterales, Intermountain flora, 

volume 5. The New York Botanical Garden, 

Bronx, NY. 496 p. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens. 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service 

USDA Forest Service, Intermountain Forest and 

Range Experiment Station, Ogden, UT. 82 p. 

McArthur, E.D. and A.P. Plummer. 1978. 

Biogeography and management of native 

western shrubs: a case study, section Tridentatae

of Artemisia. Great Basin Naturalist Memoirs 2: 

229-243. 

McArthur, E.D. and S.C. Sanderson. 1999. 

Cytogeography and chromosome evolution of 

subgenus Tridentatae of Artemisia (Asteraceae). 

American Journal of Botany 86: 1,754-1,775. 

Ward, G.H. 1953. Artemisia, section Seriphicum, 

in North America. A cytotaxonomic study. 

Contributions from the Dudley Herbarium 4: 

155-205.
77

Artemisia rigida (Nutt.) Gray stiff sagebrush 

ASTERACEAE 

Synonyms: Artemisia trifida var. rigida Nutt. 

 Seriphidium rigidum W.A. Weber 

E. Durant McArthur and Jeffrey R. Taylor 

General Description.—Stiff sagebrush, also 

known as scabland sagebrush, is a low, pungently 

aromatic shrub with thick, rigid, somewhat brittle 

branches. The bush grows up to 40 cm tall. 

Leaves, mostly 1 to 4 cm long, are deciduous, 

silvery-canescent, spatulate, and usually deeply 

divided into three to five narrowly linear lobes. 

Occasionally, leaves are linear and entire 

(Cronquist 1994, McArthur and others 1979). 

Each head is composed of five to 16 perfect disc 

flowers. The campanulate involucre is 4 to 5 mm 

long with numerous canescent bracts. The 

inflorescence is a leafy spike with heads sessile or 

in small clusters in the axils of their subtending 

leaves, which are generally longer than the heads. 

This species somewhat resembles threetip 

sagebrush (Artemisia tripartita Rydb.) in size, 

silvery pubescence, and the deeply, narrowly lobed 

leaves, but is distinguishable by the spike-like 

inflorescence, large leafy bracts that subtend the 

heads, and the deciduous leaves (Cronquist 1994, 

McArthur and others 1979). 

Taxonomy.—Stiff sagebrush does not include 

subspecific taxa. However, both diploid, 2n = 18, 

and tetraploid, 2n = 36 populations are known 

(McArthur and others 1981, McArthur and 

Sanderson 1999, Ward 1953). 

Range.—Stiff sagebrush is found in the Columbia 

and Snake River basins and the northern end of the 

Great Basin between 910 to 1,500 m elevation. Its 

range includes portions of Idaho, central and 

eastern Oregon, and eastern Washington. The 

report that its range extends into Montana is 

apparently an error (Morris and others 1976; 

McArthur and others 1979). 

Ecology.—This shrub grows primarily, if not 

exclusively, on sites with rocky, basaltic parent 

material (Daubenmire 1982, Tisdale and Hironaka 

1981). Otherwise, its ecological niche is similar to 

that of low sagebrush (Artemisia arbuscula Nutt.). 

Stiff sagebrush grows in sites with annual 

precipitation between 250 and 350 mm. Stiff 

sagebrush is able to survive wildfires even though 

it is not known to resprout or layer, probably 

because it grows in scattered patches that carry 

fires poorly. Plants grow slowly and require 2 to 4 

years to attain a mature size (> 15 cm in height). It 

appears to have broader ecological amplitude than 
78 

its present range suggests (McArthur and others 

1979). 

Reproduction.—Blooming begins in late August 

and ends in early October, and seeds ripen in 

October and November. Seed is wind dispersed in 

late fall or winter, and seedlings emerge during the 

ensuing spring. Seed production and seed quality 

are low in comparison to other sagebrush species 

(McArthur and others 1979). Attempts to seed and 

transplant this species in arid sagebrush ranges 

near Boise, Idaho, have been unsuccessful. In 

such instances, plantings were made where 

cheatgrass (Bromus tectorum L.) and other annuals 

have invaded, and stiff sagebrush seedlings were 

unable to compete. Plantings have successfully 

established on prepared seedbeds, however. 

Management.—Due to scant foliage and stiff 

branches, few animals except domestic sheep 

browse stiff sagebrush (Hall and Clements 1923). 

It provides forage on dry sites for domestic sheep 

during midsummer when herbaceous plants are 

dry. 

Benefits.—Stiff sagebrush is a native shrub that 

adds biological diversity to Western ecosystems. 

It is an important plant that provides cover and 

reduces erosion on rocky sites. Its ecology 

suggests that it could be used in reclaiming harsh, 

disturbed sites. 

References 

Cronquist, A. 1994. Asterales, Intermountain flora, 

volume 5. The New York Botanical Garden, 

Bronx, NY. 496 p. 

Daubenmire, R. 1982. The distribution of 

Artemisia rigidia in Washington: a challenge to 

ecology and geology. Northwest Science 56: 

162-164. 

Hall, H.M. and F.E. Clements. 1923. The 

phylogenetic method in taxonomy; the North 

American species of Artemisia, Chrysothamnus, 

and Atriplex. Publication 326. Carnegie 

Institution of Washington, Washington, DC. 

355 p. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service 

USDA Forest Service, Intermountain Forest and 

Range Experiment Station, Ogden, UT. 82 p. 

McArthur, E.D., C.L. Pope, and D.C. Freeman. 

1981. Chromosomal studies of subgenus 

Tridentatae of Artemisia: evidence for 

autopolyploidy. American Journal of Botany 68: 

589-605 

McArthur, E.D. and S.C. Sanderson. 1999. 

Cytogeography and chromosome evolution of 

subgenus Tridentatae of Artemisia (Asteraceae). 

American Journal of Botany 86: 1754-1775. 

Morris, M.S., R.G. Kelsey, and D. Griggs. 1976. 

The geographic and ecological distribution of big 

sagebrush and other woody Artemisias in 

Montana. Proceedings Montana Academy of 

Sciences 36: 56-79. 

Tisdale, E.W. and M. Hironaka. 1981. The 

sagebrush-grass region: a review of the 

ecological literature. Bulletin no. 33. University 

of Idaho, College of Forestry, Wildlife and 

Range Sciences, Forest, Wildlife and Range 

Experiment Station, Moscow, ID. 31 p. 

Ward, G.H. 1953. Artemisia, section Seriphicum, 

in North America. A cytotaxonomic study. 

Contributions from the Dudley Herbarium 4: 

155-205.
79

Artemisia rothrockii Gray. rothrock sagebrush 

ASTERACEAE 

Synonyms: Artemisia tridentata ssp. rothrockii Hall & Clements 

 Artemisia tridentata var. rothrockii McMinn 

Sally A. Reynolds and Eric L. Berlow 

 

General Description.—Rothrock sagebrush, also 

known as timberline sagebrush, is a low, aromatic, 

evergreen shrub, spreading to ascending from a 

narrow trunk, 20 to 60 cm tall. Bark is gray to dark 

grayish brown, becoming fibrous with age. Young 

twigs are densely tomentose. Leaves are dark 

green, narrowly cuneate, ranging from 0.5 to 2 cm 

long and 0.2 to 1.5 cm wide. Vegetative leaves are 

three-toothed but, leaves on flowering stalks are 

generally entire. Young leaves are canescent, 

becoming glabrous with age (Shultz 1993). 

Range.—Rothrock sagebrush is native and 

endemic to California. It occurs at high elevations 

in the Sierra Nevada and the White and San 

Bernardino Mountains. This species is commonly 

found in the southern Sierra Nevada on the Kern 

Plateau, at Monache Meadows, Big Meadows, 

Ramshaw Meadows, Templeton Meadows, and 

Mulkey Meadow. It is also found at Mono Pass, 

Tuolumne Pass and Mount Dana in Tuolumne 

County, Angora Peak, and near Fallen Leaf Lake 

in El Dorado County (McMinn 1951). Its 

northernmost extent is Placer County (Sawyer and 

Keeler-Wolf 1995). 

Ecology. –Rothrock sagebrush generally vegetates 

well-drained, gravelly soils on the margins of 

montane meadows from 2,000 to 3,500 m. In the 

southern Sierra Nevada, evidence suggests 

rothrock sagebrush was historically limited to dry 

soils in the areas between the meadows and 

surrounding slopes of lodgepole or red fir forest 

but has expanded into mesic meadows since the 

onset of livestock grazing in the 1850’s (Ratliff 

1985, Odion and others 1988, Dull 1995). Grazing 

and trampling along stream banks are proposed to 

have caused channel incision and an associated 

lowering of the water table in the adjacent meadow 

(Kattelmann and Embury 1996, Kirchner and 

others 1998, Knapp and Mathews 1996, USDA 

Forest Service 1998, 2000). While more often 

found on drier soils, rothrock sagebrush appears to 

be tolerant of very high soil moisture as 

individuals survive periods of standing water 

following spring snowmelt (Berlow and others 

2002). Additionally, in a pot experiment, rothrock 

sagebrush seedlings showed 100 percent survival 

after 2 weeks and over 60 percent survival after 4 

weeks of complete immersion in flowing water 

(Swartz, Berlow and D’Antonio unpublished data). 

The highest potential germination and seedling 

growth rates were observed in mesic, rather than 

xeric, meadow habitat. However, in these areas 

micro-site availability and seed dispersal limit 

establishment, with seedlings preferentially 

associated with small soil disturbances within 0.5 

m of a seed source (Berlow and others 2002). 

Reproduction.—Flowers develop on narrow 

inflorescences. Heads are sessile to shortpeduncled, 3 to 5 mm in diameter. Phyllaries 

(involucre bracts) occur in two series. Both series 

are shiny with wide margins. The outer phyllaries 

are canescent with acute tips, whereas the inner 

phyllaries are elliptic and only sparsely tomentose. 

Discoid flowers (ray florets are absent) have 

between eight and 20 fertile florets. Seeds are 

small and resinous, ranging from 0.8 to 2 mm, 

with pappus present on outer seeds (Shultz 1993). 

Inflorescences begin development in late spring. 

Flowering takes place at the end of the growing 

season in early to late fall, and seed dispersal 

begins soon after. Seeds are primarily wind 

dispersed and do not appear to travel far, as 

seedlings tend to be aggregated within 0.5 m of 

adult shrubs (Berlow and others 2002). 

Germination occurs immediately following spring 
80 

snow-melt. Rothrock sagebrush does not form a 

large soil seedbank (Reynolds unpublished data). 

An average sized adult plant can produce over 

5000 seeds (Reynolds unpublished data). 

However, field germination rates are low (< 1 

percent), and seed viability appears to decline 

rapidly after 1 year (Berlow and others 2002, and 

unpublished data). 

Growth and Management.—The growing season 

for rothrock sagebrush in montane habitats is 

relatively short, beginning in early spring and 

continuing until the soil moisture is depleted in 

late summer. Under mesic conditions, rothrock 

sagebrush can begin flowering within 5 years of 

establishment (Berlow and others 2002). While 

age to senescence is unknown, individuals up to 40 

years old have been documented by counting 

growth rings (Bauer and others 2002). Rates of 

increase in stem diameter have been observed to 

vary between 0.10 to 0.14 mm/year. Faster growth 

rates were found in microhabitats characterized by 

moist surface soil and a shallow water table, 

suggesting that this woody species can use similar 

water sources as herbaceous species (Bauer and 

others 2002, Berlow and others 2003). Rothrock 

sagebrush seedlings are not palatable to livestock 

or native grazers, which may contribute to their 

encroachment into intensively grazed meadows. 

While large areas of sagebrush expansion are 

associated with stream incision, rothrock 

sagebrush also has the demonstrated potential to 

invade un-incised, mesic meadow habitat given the 

presence of exposed soil and a nearby seed supply 

(Berlow and others 2002). Thus, on a large scale, 

preventing further stream incision is critical to 

slowing sagebrush invasion. Within mesic 

meadow habitat, removal of scattered individual 

shrubs that serve as local seed sources may be the 

most effective way to prevent rapid expansion into 

un-incised areas. 

Benefits.—Rothrock sagebrush contributes to the 

diversity of California’s montane plant 

communities. It stabilizes sandy soil on the slopes 

of meadow fringes and provides habitat for a 

variety of birds and small mammals. 

References 

Bauer, K., E.L. Berlow, and C.M. D’Antonio. 

2002. Shrub expansion in montane meadows: 

The relationship between climate and Rothrock 

sagebrush colonization patterns. Journal of 

Range Management. 55: 620-625. 

Berlow, E.L., C.M. D’Antonio, and S.A. 

Reynolds. 2002. Shrub expansion in montane 

meadows: the interaction of local-scale 

disturbance and site aridity. Ecological 

Applications 12: 1103-1118. 

Berlow, E.L., C.M. D’Antonio, and H. Swartz. 

2003. Response of herbs to shrub removal across 

natural and experimental variation in soil 

moisture: Implications for restoration. Ecological 

Applications. (in press). 

Dull, R.A. 1995. Palynological evidence for 19th

century grazing-induced vegetation change in 

the southern Sierra Nevada, California. Journal 

of Biogeography 26: 899-912. 

 Kattelmann, R. and M. Embury. 1996. Riparian 

areas and wetlands. In Sierra Nevada 

Ecosystem Project. University of California, 

Centers for Water and Wildland Resources, 

Davis, CA. p. 34-68. 

Kirchner, J.W., L. Micheli, and J.D. Farrington. 

1998. Effects of herbaceous riparian vegetation 

on streambank stability. Technical Completion 

Report, Project #W-872, University of 

California Berkeley Water Resources Center. 

43 p. 

Knapp, R.A. and K.R. Matthews. 1996. 

Livestock grazing, Golden Trout, and streams 

in the Golden Trout Wilderness, California: 

impacts and management implications. North 

American Journal of Fisheries Management 16: 

805-820. 

McMinn, H.E. 1951. An Illustrated Manual of 

California Shrubs. University of California 

Press, Berkeley, CA. p. 608-609. 

Odion, D.C., T.L. Dudley, and C.M. D'Antonio. 

1988. Cattle grazing in southeastern Sierran 

meadows: Ecosystem change and prospects for 

recovery. In: C.A. Hall and V. Doyle-Jones, 

editors. Plant Biology of Eastern California, 

Mary Decker Symposium. White Mountain 

Research Station, Los Angeles, CA. p. 277-292. 

Ratliff, R.D. 1985. Meadows in the Sierra Nevada 

of California: state of knowledge. General 

Technical Report PSW-84. U.S. Department of 

Agriculture, Forest Service, Pacific Southwest 

Forest and Range Experiment Station. 

Berkeley, CA. 52 p. 
81

Sawyer, J.O. and T. Keeler-Wolf. 1995. A Manual 

of California Vegetation. California Native Plant 

Society, Sacramento, CA. p. 188. 

 

Shultz, L.M. 1993. Artemisia. In: J.C. Hickman, 

ed. The Jepson Manual: Higher Plants of 

California. University of California Press, 

Berkeley, CA. p. 202-205. 

USDA Forest Service. 1998. Kern Plateau 

Ecosystem Analysis. U.S. Department of 

Agriculture, Forest Service, Inyo National 

Forest, Bishop, CA. 215 p. 

USDA Forest Service. 2000. Templeton and 

Whitney Grazing Allotments Environmental 

Assessment. U.S. Department of Agriculture, 

Forest Service, Inyo National Forest, Bishop, 

CA. 111 p. 
82 

Artemisia spinescens Eaton budsage 

ASTERACEAE 

Syononyms: Picrothamnus desertorum Nutt. 

E. Durant McArthur and Jeffrey R. Taylor 

General Description.—Budsage, also known as 

spring sage and bud sagebrush, is a low, 

spinescent, pungently aromatic, rounded shrub 10 

to 50 cm high and profusely branched at the base. 

Young branches and leaves are covered with a 

white-tomentose pubescence that becomes stiff 

and gray with age. Leaves are small and normally 

2 cm or less in length. Leaves are three- to fivepalmately parted, again divided into three linearspatulate lobes. Leaves are crowded on short 

stems, with those near the apex being smaller and 

more entire. Unlike most members of the genus, 

budsage is deciduous, losing most leaves by 

midsummer. Flowerheads are small and arranged 

in glomerate racemes of one to three heads in leaf 

axils on flower branches. Each head contains two 

to six fertile, pistilate ray flowers and five to 13 

perfect, but sterile, disc flowers. Four to eight 

rounded involucral bracts subtend each flowerhead 

(Cronquist 1994, McArthur and others 1979). 

Taxonomy.—Budsage is distinct from closely 

related Artemisia species and is sometimes 

considered a separate monotypic genus although 

(Cronquist 1994) states that “despite this proposed 

segregation, its affinity with Artemisia is not in 

question”. The inclusion or exclusion of segregate 

genera from Artemisia including Picrothamnus has 

not been confirmed by molecular genetic studies 

as of yet (Vallès and McArthur 2001). Polyploidy 

within the budsage has been documented; both 

diploid, 2n = 18, and tetraploid, 2n = 36, 

populations occur (McArthur and Pope 1979). 

Range.—The range of budsage extends from 

Wyoming to western Montana, southern Idaho, 

and easten Oregon south to northwestern New 

Mexico, northern Arizona, and southeastern 

California (Cronquist 1994, McArthur and others 

1979). 

Ecology.—Budsage is well adapted to xeric 

conditions. It is quite common in semiarid valley 

bottoms, benches, and foothills throughout the 

Interior Western United States (Hall and Clements 

1923, Wood and Brotherson 1986). Mean annual 

precipitation throughout its habitat varies between 

150 and 250 mm. It has an extensive root system 

that grows primarily within the top 15 to 56 cm of 

the soil. Interxylary cork is formed over the 

previous year’s wood that restricts the upward 

movement of water to the narrow zone of wood 

formed by the current year’s growth. The corky 

tissue develops during the early summer and 

reduces water loss during dormancy (Wood 1966). 

 Summer dormancy is an adaptation that 

reduces the effects of drought stress. Early in the 

spring, when budsage first shows signs of breaking 

dormancy and prior to bud elongation, the 

previous season’s bark growth can be removed 

easily. This condition is known as “slipping” and 

is used as a sign to tell when the plant is palatable 

to sheep (Wood and Brotherson 1986). Shrubs 

subsequently produce new, bright-green leaves as 

soon as February or March. Terminal and lateral 

buds generally expand and begin to elongate in 

late March and early April. Although budsage 

ordinarily begins to grow in early spring and 

becomes dormant by midsummer, it occasionally 

may break summer dormancy after late summer 

storms. 
83

 

Reproduction.—Blooming normally begins 

during the last week in April and continues through 

the last week in May. Exceptions, however, have 

been documented where blooming began as early 

as late March or has extended as late as mid-June 

(Wood 1966). Seeds normally mature in early 

June. There are about 3 million cleaned seeds per 

kg (Meyer 2003). Freezing temperatures in early 

spring frequently kill developing embryos. Intact 

heads regularly fall from the plant without breaking 

apart to release seeds, although some seeds are 

usually dispersed independently. Abundant natural 

reproduction occurs in years when seed production 

is plentiful and moisture conditions are favorable. 

Management.—Budsage is a palatable, nutritious 

winter forage for upland birds, small game, big 

game, and domestic sheep. It is particularly 

preferred just after dormancy. Budsage is high in 

calcium, magnesium, phosphorous, and protein 

(Wood and Brotherson 1986). It is usually more 

palatable in late winter than early winter 

(Holmgren and Hutchings 1972). Palatability 

again decreases once twigs have elongated and 

volatile oil content increases (Cook and others 

1954). Budsage is especially important to grazing 

animals during late winter in areas where there is 

an abundance of dry grass. Unfortunately, it does 

not tolerate grazing well, and continual heavy 

grazing may eliminate it from certain areas 

(Holmgren and Hutchings 1972). In order to 

maintain the species in plant communities, 

periodic rest from grazing is needed. This allows 

seed and seedlings to develop. 

Benefits.—Budsage is an important native shrub 

because of its forage value to various species of 

wildlife and domestic livestock. Wildlife species 

that utilize budsage include mule deer (Odocoileus 

heminonus Rafinesque), pronghorn (Antilocapra 

Americana Ord), mountain sheep (Ovis canadensis 

Shaw), black-tailed jackrabbits (Lepus californicus 

Gray), and chucker partridge (Alectoris graeca 

Meisner) (Wood and Brotherson 1986). In areas 

of local abundance, budsage can be an important 

source of protein during late winter months. 

References 

Cook, C.W., L.A, Stoddart, and L.E. Harris. 1954. 

The nutritive value of winter range plants in the 

Great Basin as determined with digestion trials 

with sheep. Bulletin 372. Utah State University, 

Agricultural Experiment Station, Logan, UT. 

56 p. 

Cronquist, A. 1994. Asterales, Intermountain flora, 

Vol. 5. The New York Botanical Garden, Bronx, 

NY. 496 p. 

Hall, H.M., and F.E. Clements. 1923. The 

phylogenetic method in taxonomy; the North 

American species of Artemisia, Chrysothamnus, 

and Atriplex. Publication 326. Carnegie 

Institution of Washington, Washington, DC. 

355 p. 

Holmgren, R.C. and S.S. Hutchings. 1972. Salt 

desert shrub response to grazing use. In: McKell, 

C.M., J.P. Blaisdell, and J.R. Goodin, tech. eds. 

Proceedings-symposium on wildland 

shrubs--their biology and utilization, 1971 July; 

Logan, UT. Gen. Tech. Rep. INT-1. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. p. 153-164. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 82 p. 

McArthur, E.D. and C.L. Pope. 1979. Karyotypes 

of four Artemisia species: A. carruthii, A. 

filifolia, A. frigida, and A. spinescens. Great 

Basin Naturalist 39: 419-426. 

Meyer, S. E. 2003. Artemisia. In: Woody plant 

seed manual. http://www.wpsu.net [not paged]. 

Vallès, J. and E.D. McArthur. 2001. Artemisia 

systematics and phylogeny: cytogenetic and 

molecular insights. In: E.D. McArthur and D.J. 

Fairbanks, comps. Shrubland ecosystem genetics 

and biodiversity: proceedings, 2000 June 13-15; 

Provo, UT. Proceedings RMRS-P-21. U.S 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Ogden, UT. 

p. 67-74. 

Wood, B. W. 1966. An ecological life history of 

budsage in western Utah. M.S. thesis. Brigham 

Young University, Provo, UT. 85 p. 

Wood, B.W. and J.D. Brotherson. 1986. 

Ecological adaption and grazing response of 

budsage (Artemisia spinescens) in southwestern 

Utah. In: McArthur, E.D. and B.L. Welch, 
84 

comps. Proceedings--symposium on the biology 

of Artemisia and Chrysothamnus, 1984 July 

9-13; Provo, UT. General Technical Report 

INT-200. U.S. Department of Agriculture, Forest 

Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. p. 75-92. 
85

Artemisia tridentata Nutt. big sagebrush 

COMPOSITAE 

Synonyms: Artemisia tridentata var. angustifolia Gray 

 Artemisia angusta Rydb. 

 Artemisia tridentata ssp. typica Hall & Clements 

Bruce L. Welch 

General Description.—Big sagebrush is an erect, 

aromatic, evergreen species, that lacks winter 

buds. It has numerous ecotypes that range in size 

from 0.4 to 4.5 m tall (Beetle 1960, Beetle and 

Young 1965, Diettert 1938, McArthur and others 

1979). Growth forms are of two types; 

multistemmed and single trunks and are correlated 

with subspecific taxonomy. This species produces 

two types of branches, vegetative and flowering 

(Diettert 1938). The bark on older vegetative 

branches is stringy and black or dark brown in 

color. Bark on younger vegetative and 

inflorescences is heavily covered with trichomes 

that give the branches a silvery green to gray color. 

The leaves are also silvery green or gray. Principal 

leaves are narrowly cuneate or oblanceolate, 1.0 to 

6.5 cm long and 2 mm to 2 cm wide, and usually 

are three-toothed at the apex (McArthur and others 

1979). Leaf arrangement is spiral with a 2/5 

divergence (Diettert 1938). Internodes are often so 

short that the leaves form dense, rosette-like 

clusters at the shoot tips (Diettert 1938). The 

haploid chromosome number of big sagebrush is 

nine (Diettert 1938, McArthur and Sanderson 

1999). Two levels of ploidy have been reported 

among the three subspecies of big sagebrush–

diploid (2n = 2x = 18) and tetraploid (2n = 4x = 

36) (Diettert 1938, McArthur and Sanderson 

1999). All three subspecies of big sagebrush 

contain polyploid populations (McArthur and 

Sanderson 1999). 

Taxonomy.—There are three widely recognized 

subspecies of big sagebrush (Beetle 1960, Beetle 

and Young 1965). These are Wyoming big 

sagebrush (A. t. ssp. wyomingensis), basin big 

sagebrush (A. t. ssp. tridentata), and mountain big 

sagebrush (A. t. ssp. vaseyana). Subspecies can be 

separated on morphological, chemical, and 

ecological characteristics (McArthur and others 

1979, Welch 2002). Wyoming big sagebrush 

occupies the drier sites that range from 20 to 32 

cm of precipitation, 31 to 149 cm for mountain big 

sagebrush with basin big sagebrush somewhat 

between the other two subspecies (Welch 2002). 

Some authorities have divided mountain big 

sagebrush into three forms, varieties, or subspecies 

depending on the authority being quoted. One is 

called subalpine big sagebrush, A. t. ssp. 

spiciformis, (McArthur and Goodrich 1986) or A. 

t. ssp. vaseyana f. spiciformis (Beetle and Johnson 

1982). The second is referred to as “X” big 

sagebrush, or xeric big sagebrush or A. t. ssp. 

xericensis (Welch 2002). Lastly, a variety of 

mountain big sagebrush is called A. t. ssp. 

vaseyana var. pauciflora (Welch 2002). 

Range.—Beetle (1960) describes the range or 

distribution of big sagebrush in these terms: 

“occupying a great variety of sites at scattered 

localities in southern Montana and central to 
86 

western Wyoming; southern-central British 

Columbia, south through central Washington, to 

the Columbia River; from the south side of the 

Blue Mountains in northeastern Oregon, 

throughout central and southeastern Oregon, 

southwestern and central Idaho to northern Utah, 

Nevada, and northwestern California; from the 

northern great basin areas of California, Nevada, 

and Utah, extending southward at elevations from 

5,000 to 7,000 feet and extending into western 

Colorado, northwestern New Mexico, northern 

Arizona; at much lower elevations in southern 

California and northern Lower California, 

Mexico.” It also occupies areas of the northern 

Great Plains in the States of North and South 

Dakota, Montana, Wyoming, and Nebraska 

(Welch 2002). 

Ecology.—Big sagebrush grows in a variety of 

soils throughout its range from arid plains, valleys, 

foothills, to mountain slopes, from 500 to 3,400 m 

elevation (McArthur and others 1979, Welch 

2002). It has been found growing on five soil 

order: Alfisols, Aridisols, Entisols, Inceptisols, and 

Mollisols. It can be found growing on all 12 soil 

textural classes, but it is most often found on 

loams or sandy loams (Welch 2002). Chemical 

properties of big sagebrush soils are highly 

variable (pH 5.9 to 10.0 and organic carbon 0.62 to 

4.14 percent) but two characteristics are common: 

they are well-drained and contained low 

concentrations of salts (Welch 2002). Depending 

on subspecies, as many as 40 plant species 

(grasses and forbs) can grow with big sagebrush 

(Welch 2002). Big sagebrush is not fire tolerant 

and can be killed by a number of pathogenic fungi, 

insects, and environmental conditions such as 

winter kill or winter induced drought (Welch 

2002). 

Reproduction.—The growth of inflorescence 

starts in late spring and is complete by late 

summer to early fall (Welch 2002). Flowering 

occurs during early to late fall with fruit 

development commencing almost simultaneously 

with flowering. Thus reproductive growth occurs 

when water supplies and temperatures are not 

particularly favorable; in fact, during this period 

big sagebrush plants have started shedding 

ephemeral leaves and neighboring plants species 

are largely dormant (Welch 2002). However, 

inflorescences do generate positive net 

photosynthesis at a time when whole plant is in a 

water conservation mode (Welch 2002). Big 

sagebrush achenes or seeds are small, weighing 

from 0.00018 to 0.00025 g (Welch 2002). Time of 

seed dispersal is highly variable among stands and 

somewhat dependent on elevation; some starting in 

mid-fall other not until early winter. Most seeds 

germinate immediately after snow melt or after 

late-winter or early spring storms (Welch 2002). 

The seeds possess no special adaptations for wind 

dispersal, but are wind dispersed to a maximum 

distance of 30 m from the mother plant. Hence 

seedlings placement is heavily depended on wind 

direction (Welch 2002). Big sagebrush forms a 

weak soil seed bank (Welch 2002). A single 

mature big sagebrush plant can produced upward 

of 500,000 seeds, although, excessive browsing 

can reduce inflorescences production by a factor of 

20 (Welch 2002). The seeds must lie on the soil 

surface or be buried not more than 5 mm for 

successful emergence (Welch 2002). 

Growth and Management.—Vegetative growth 

starts in early spring and declines when soil 

moisture becomes deficient. In general, Wyoming 

big sagebrush, the smallest of the subspecies, has 

the slowest vegetative growth rate and produces 

the least amount of biomass. Basin big sagebrush 

has the fastest growth rate and is the largest of the 

subspecies (McArthur and others 1979, Welch 

2002). In the past 50 years or more, management 

has erroneously perceived big sagebrush to be a 

highly competitive, dominating, and suppressive 

species of grasses and forbs (Welch 2002). Thus 

a variety of techniques have been developed to kill 

big sagebrush (Welch 2002). As more studies 

show the importance of big sagebrush in 

maintaining it’s ecosystem, techniques have been 

and continue to be developed to restore big 

sagebrush to its native sites (Welch 2002). 

 

Benefits.—Big sagebrush is the keystone species 

of a large and highly variable ecosystem of 

Western United States (Welch 2002). It is literally 

a “nursing mother” to a host of organisms ranging 

from microscopic to large mammals and is what 

supports life in Bailey’s (1896) description: “One 

never recovers from his surprise that there should 

be so much life where apparently there is so little 

to support it.” Species that depend directly or 

indirectly on big sagebrush are many: 31 species 

of fungi, 52 species of aphids, 10 species of insects 

that feed on aphids, 42 species of midges and fruit 

flies that induce galls, 20 species of insects that 

parasitize the gall inducers, 6 species of insects 

that hibernate in big sagebrush galls, 18 species of 

beetles, 13 species of grasshoppers, 13 species of 

shield-back katydids, 16 species of thrips, 74 

species of spiders, 24 species of lichens, 16 species 

of paintbrushes, 7 species of owl-clovers, 5 species 
87

of bird’s beaks, 3 species of broom rapes, and a 

host of large and small mammals, birds, and 

reptiles (Welch 2002). Soil fertility is higher 

under big sagebrush canopy cover than outside the 

canopy (Welch 2002). Roots of other plants can 

tap into these islands of fertility (Krannitz and 

Caldwell 1995). Also, by a process known as 

“hydraulic lift or hydraulic redistribution” 

associated plants receive part of their needs from 

water pumped by big sagebrush to the soil surface 

from greater soil depths (Welch 2002). Lastly, big 

sagebrush provides a safe haven for grasses and 

forbs growing under its canopy from grazing 

domestic livestock (Welch 2002). 

References 

Bailey, W.W. 1896. The sage brush. American 

Naturalist 3:356-360. 

Beetle, A.A. 1960. A study of sagebrush–The 

section Tridentatae of Artemisia. Bulletin 368. 

University of Wyoming, Agricultural 

Experiment Station. Laramie. 83 p. 

Beetle, A.A. and K.L. Johnson. 1982. 

Sagebrush in Wyoming. B-779, University of 

Wyoming, Agricultural Experiment Station, 

Laramie. 68p. 

Beetle, A.A. and A. Young. 1965. A third 

subspecies in the Artemisia tridentata complex. 

Rhodora 67:405-406. 

Diettert, R.A. 1938. The morphology of Artemisia 

tridentata Nutt. Lloydia 1:3-74. 

Krannitz, P.G. and M.M. Caldwell. 1995. Root 

growth responses of three Great Basin 

perennials to intra-and inter specific contact with 

other roots. Flora. 190:161-167. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens. 1979. Characteristics and 

hybridization of important Intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 82 p. 

McArthur, E.D. and S.K. Goodrich. 1986. 

Artemisia tridentate ssp. spiciformis: 

distribution and taxonomic placement. In: E.D. 

McArthur and B.L. Welch, comps. Proceedings: 

Symposium on the biology of Artemisia and 

Chrysothamnus; July 9-13, 1984; Provo, UT. 

Gen. Tech. Rep. INT-200. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Research Station, Ogden, UT. 55-57. 

McArthur, E.D. and S.C. Sanderson. 1999. 

Cytogeography and chromosome evolution of 

subgenus Tridentatae of Artemisia (Asteraceae). 

American Journal of Botany. 86:1,754-1,775. 

Welch, B.L. [in press]. Big sagebrush: A sea 

fragmented into lakes, puddles, and ponds. 

General Technical Report. U.S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Research Station, Fort Collins, CO. 
88 

Artemisia tripartita Rydb. threetip sagebrush 

ASTERACEAE 

Synonyms: Artemisia trifida Nutt. 

 Seriphidium tripartitum (Rydb.) W.A.Weber 

E. Durant McArthur and Jeffrey R. Taylor 

General Description.—Threetip sagebrush is a 

round, evergreen shrub up to 1.8 m tall. It may 

have a simple, trunk-like stem or several, lesser 

stems arising from the base. The bark on young 

branches is canescent but becomes shredded and 

grayish, light brown to dark brown or black, on 

older stems. This species can layer and resprout 

after fire or herbicide treatment (Beetle 1960, 

Pechanec and others 1965, Schlatterer 1973, 

Winward 1980). Leaves on vegetative branches are 

canescent, 0.5 to 4 cm long, and typically deeply 

divided into three linear or narrowly linearlanceolate lobes, which in turn may be three-cleft. 

Upper leaves are often entire. Crushed foliage 

emits a pungent odor. Flowerheads contain three to 

11 disc flowers typically arranged into panicles. 

Ray flowers are lacking. Eight to 12 canescent 

involucral bracts subtend each head. Achenes are 

resinous-granuliferous (Cronquist 1994, McArthur 

and others 1979). 

Taxonomy.—The taxon includes two subspecies: 

Artemisia tripartita ssp. tripartita and Artemisia 

tripartita ssp. rubicola Beetle. This species, as is 

quite common for other species in the subgenus 

Tridentatae, includes populations that are diploid, 

2n = 18 and tetraploid, 2n = 36 (McArthur and 

Sanderson 1999). 

Subspecies tripartita, also known as tall 

threetip sagebrush, is a freely-branching shrub up 

to 1.8 m high. It can layer when conditions are 

conducive, but it is seldom found layering in the 

field. It can stump-sprout after fire (Beetle 1960). 

Leaves on vegetative branches are 1.5 to 4 cm long 

and deeply divided into three linear lobes, each 

less than 1 mm wide, that may be further divided 

(Beetle 1960). Flower heads bear four to eight disc 

flowers arranged into panicles that may be reduced 

to a spicate form. 

Subspecies rubicola, also known as 

Wyoming threetip sagebrush, is a dwarf shrub 

usually less than 15 cm tall with decumbent 

branches. It layers frequently and may have a 

crown diameter of 30 to 50 cm (Beetle 1960). 

Leaves on vegetative branches are often 3 cm long 

and deeply divided into linear lobes, each at least 1 

mm wide (Beetle 1959, 1960). Flower heads bear 

three to 11 disc flowers arranged in leafy, 

narrowly racemose panicles. 
89

Range.—Threetip sagebrush covers approximately 

34,000 km2 in the Northern Rocky Mountains and 

Great Basin States. It usually grows at elevations 

between 910 to 2,700 m. It extends from British 

Columbia south through Montana and Wyoming to 

Colorado, and west to Washington, Oregon, 

northern Nevada, and northern Utah (Beetle 1960). 

Subspecies rubicola, however, is limited to areas 

of central and southeastern Wyoming and southern 

Oregon (Brunner 1972). 

Ecology.—Threetip sagebrush grows well on 

moderate to deep, well-drained, loamy and sandy 

loam soils (Winward 1980). In some places, 

particularly in Idaho, this species grows between 

the lower, hot, dry sites dominated by Wyoming 

big sagebrush (Artemisia tridentata Nutt. ssp. 

wyomingensis Beetle & Young) and the higher, 

cooler sites dominated by mountain big sagebrush 

[Artemisia tridentata ssp. vaseyana (Rydb) Beetle] 

(Schlatterer 1973). It often grows intermixed, but 

as separate stands, with mountain big sagebrush, 

Wyoming big sagebrush, or alkali sagebrush 

[Artemisia longiloba ([Osterh.) Beetle] in areas 

with 300 to 600 mm of annual precipitation. 

Wyoming threetip sagebrush grows on rocky 

knolls at high elevations (2,100 to 2,700 m) 

adjacent to mountain big sagebrush sites. 

Reproduction.—Flowers bloom between July and 

September and seed matures in October. Seeds are 

wind dispersed and seedlings emerge during the 

ensuing spring. Plants grow quickly and attain 

maturity in 3 to 5 years. The seeds are smaller than 

many Artemisia species; there are approximately 5 

million cleaned seeds/kg (Meyer 2003). 

Management.—This species is a vigorous seeder. 

Some forms are more palatable than others. 

Threetip sagebrush establishes well by direct 

seeding, rearing, or transplanting. Seedlings are 

able to persist with both a dense perennial and 

annual understory, although it is no more effective 

at excluding invasive annuals than other 

sagebrushes. It frequently grows with a number of 

other shrubs and herbs. It is compatible with many 

introduced and native grasses including mountain 

brome (Bromus carinatus H. & A.), slender 

wheatgrass [Elymus trachycaulus (Link) Gould ex 

Shinn], intermediate wheatgrass [Elymus hispidus 

(Opiz) Meld], and orchard grass (Dactylis 

glomerata L.). 

Attempts to establish threetip sagebrush 

on arid sites dominated by Wyoming big 

sagebrush have been unsuccessful. Some ecotypes, 

however, can extend into drier sites. Threetip 

sagebrush generally recovers well after 

disturbance, but hot summer fires can kill large 

stands, inhibit resprouting, and significantly delay 

natural seedling recovery. 

Benefits.—Three-tip sagebrush adds structural and 

biological diversity to the landscape and reduces 

the effects of wind and soil erosion. Although 

Beetle (1960) reported it was not palatable to 

livestock or game, Kufeld (1973) reported that elk 

used it, and Kufeld and others (1973) also reported 

that deer use it. A form near Salmon, Idaho, is 

palatable to deer and may have introgressed into a 

nearby population of Wyoming big sagebrush 

(McArthur and others 1979). 

 

References 

Beetle, A.A. 1959. New names within the section 

Tridentatae of Artemisia. Rhodora 61: 82-85. 

Beetle, A.A. 1960. A study of sagebrush. The 

section Tridentatae of Artemisia. Bulletin 368. 

University of Wyoming, Agricultural 

Experiment Station, Laramie, WY. 83 p. 

Brunner, J.R. 1972. Observations on Artemisia in 

Nevada. Journal of Range Management 25: 205-

208. 

Cronquist, A. 1994. Asterales, Intermountain flora, 

Vol. 5. The New York Botanical Garden, Bronx, 

NY. 496 p. 

Kufeld, R.C. 1973. Foods eaten by the Rocky 

Mountain elk. Journal of Range Management. 

26: 106-113. 

Kufeld, R.C., O.C. Wallmo, and C. Feddema. 

1973. Foods of the Rocky Mountain mule deer. 

Research Paper RM-11. U.S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Forest and Range Experiment Station, Fort 

Collins, CO. 31 p. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 82 p. 

McArthur, E.D. and S.C. Sanderson. 1999. 

Cytogeography and chromosome evolution of 
90 

subgenus Tridentatae of Artemisia (Asteraceae). 

American Journal of Botany 86: 1,754-1,775. 

Meyer, S.E. 2003. Artemisia. In: F.T. Bonner, and 

R.G. Nisley, eds. Woody plant seed manual. 

http://www.wpsu.net [not paged]. 

Pechanec, J. F., A.P. Plummer, J.H. Robertson, 

and A.C. Hull, Jr. 1965. Sagebrush control on 

rangelands. Agriculture. Handbook 277. U.S. 

Department of Agriculture, Washington, DC. 

40 p. 

Schlatterer, E.F. 1973. Sagebrush species and 

subspecies. U.S. Department of Agriculture, 

Forest Service, R-4 Range Improvement Notes 

18(2): 1-11. 

Winward, A.H. 1980. Taxonomy and ecology of 

sagebrush in Oregon. Bullitin 642. Oregon State 

University, Oregon Agricultural Experiment 

Station, Corvallis, OR. 15 p. 
91

Arundinaria gigantea (Walt.) Muhl. giant cane 

POACEAE 

Synonyms: Arundinaria macrosperma Michx. 

 Arundinaria tecta (Walter) Muhl. 

 Arundinaria tecta var. decidua Beadle 

 Arundo gigantea Walter 

 Arundo tecta Walter 

Kristina Connor 

Illustration source: Grelen and Duvall 1966 

General Description.—Giant cane, also known as 

cane or switchcane, is a perennial monocot, a 

woody grass, and one of only two native bamboos. 

With its stem-like rhizomes and hard, ‘woody’ 

stems, giant cane can grow to a height of 8 to 9 m 

but is typically less. Lower sheaths are about half 

as long as the culm internodes, while the upper six 

to 10 sheaths culminate in 10 to 12 bristles that are 

5 to 9 mm long (Hitchcock 1971). Leaves range 

from 10 to 30 cm in length and from 2 to 3 cm 

wide. They have rounded base, are lance shaped, 

and have finely toothed margins (Bailey and 

Bailey 1976). Leaves are typically pubescent 

underneath and are either pubescent or glabrous 

above. Leaves on stems 2 years and older grow on 

side branches. Branches are short, less than 30 cm, 

also with loose papery sheaths. 

Range.—Giant cane is found at elevations ranging 

from sea level in southern floodplains to 610 m 

elevation in the Appalachian Mountains. It can 

grow in dense thickets in the Mississippi Delta and 

in other southern swamplands. Its range extends 

from southern Maryland west into Ohio, Indiana, 

and Illinois, south to the Gulf Coast and west to 

Texas, Oklahoma, and Arkansas (Bailey and 

Bailey 1976, Brickell and Zuk 1996, Walkup 

1991). A recognized variety is A. gigantea subsp. 

tecta (Walter) McClure. 

Ecology.—Giant cane formerly occupied large 

areas (canebrakes) in floodplains of southern 

rivers; now these thickets are usually found only in 

the Mississippi Delta where they form in lowlying, shady moist areas. Elsewhere, giant cane is 

usually intermixed with shrubs. It is fire dependent 

and resprouts from rhizomes (Walkup 1991). It has 

a broad tolerance for weather and can withstand 

temperatures ranging from -23 to 41 o

C. It grows 

in a variety of soil types (muck lands to mountain 

slopes and rich alluvial soils) and is rugged, coldhardy and adaptable. 
92 

Reproduction.—Giant cane primarily spreads by 

rapid vegetative reproduction from large rhizomes. 

It flowers infrequently, and at irregular intervals, 

in early spring, forming simple panicles on the 

branches. The panicles consist of a few racemes of 

five to 15 large, stalked spikelets, which may be 

purple (Bailey and Bailey 1976, Brickell and Zuk 

1996). The inflorescence may be a single axis or 

have branches. Seed production from these flowers 

is sparse and unpredictable. 

Growth and Management.—Giant cane is a 

short-lived, evergreen monocot. Individual stems 

survive less than 10 years, but the species sprouts 

prolifically from its rhizomes. In the presence of 

fire and on the best-suited sites, it forms dense 

thickets called canebrakes. While it can withstand 

flooding, drought, and intense surface fires, giant 

cane is sensitive to overgrazing (Grelen and 

Duvall 1966, Hitchcock 1971). One source reports 

rapid growth (Brickell and Zuk 1996), especially 

after a fire. Fuel accumulates rapidly in a 

canebrake, and, to eliminate the hazard, a short 

burning cycle is advised (Hughes 1966, Walkup 

1991). 

Benefits.—The dense thickets of giant cane 

provide cover for nesting birds and small 

mammals. In addition, the young shoots are edible, 

sometimes used as a potherb (Hitchcock 1971), 

and of good nutritional quality. The leaves are a 

preferred food for southern pearly eye butterfly 

caterpillars (University of Florida 2001). The 

leaves extend well above the ground, plants can be 

completely defoliated by cattle and are also 

uprooted by swine (Hitchcock 1971, Walkup 

1991). Thus, while grazing capacity of cane is 

high, careful management is required to prevent 

deterioration of the plants (Grelen and Duvall 

1966). 

References

Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: 

A Concise Dictionary of Plants Cultivated in the 

United States and Canada. McMillan Publishing 

Co., Inc. New York. 1,290 p. 

Brickell, C. and J.D.. Zuk. 1996. The American 

Horticultural Society A-Z Encyclopedia of 

Garden Plants. DK Publishing, Inc., New York. 

1,092 p. 

Grelen, H.E. and V.L. Duvall. 1966. Common 

plants of longleaf pine-bluestem range. Res. Pap. 

S0-23, U.S. Department of Agriculture Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 96 p. 

Hitchcock, A.S. 1971. Manual of Grasses of the 

United States. Vol. 1, 2nd Ed. Revised by A. 

Chase. Dover Publications Inc., New York. 569 

p. 

Hughes, R.H. 1966. Fire ecology of canebrakes. 

In: Proceedings 5th Annual Tall Timbers Fire 

Ecology Conference, Tallahassee, FL. Tall 

Timbers Research Station. p. 148-158. 

University of Florida. 2001. Website, Center for 

Aquatic and Invasive Plants. http://aquat1.ifas. 

ufl.edu/arugig.html. 1 p. 

USDA, NRCS. 2001. The PLANTS database, 

National Plant Data Center, Baton Rouge, LA. 

http://plants.usda.gov. [not paged]. 

Walkup, C. 1991. Arundinaria gigantea. In: U.S. 

Department of Agriculture, Forest Service, Rocky 

Mountain Research Station Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed. us/database/feis. 9 p. 
93

Atriplex canescens (Pursh) Nutt. fourwing saltbush 

CHENOPODIACEAE 

Synonyms: Calligonum canescens Pursh 

E. Durant McArthur, Stewart C. Sanderson, and Jeffrey R. Taylor 

Illustration source: USDA—Forest Service collection, 

Hunt Institute 

General Description.—Fourwing saltbush, also 

known as costilla de vaca and chamizo, is an 

upright shrub between 10 and 200 cm tall. Stature 

varies with site conditions and genotype. Branches 

arise freely from the base and are brittle and hard at 

maturity. Young branches and leaves are covered 

with inflated or collapsed bladder hairs that create 

a scurfy, grey-green appearance. Leaves are 

evergreen, linear to oblanceolate or spatutulate, 

usually 1 to 4 cm long and 0.1 to 1.0 cm wide. The 

species is usually dioecious, although some are 

monoecious or trioecious. Pistillate (female) 

flowers are small and inconspicuous. A pair of 

small bracts united along their edges encloses each 

pistil. Each of these bracts has two ridges down the 

face that expand into “wings,” giving the utricle 

(fruit) four “wings” at maturity. Staminate (male) 

flowers are small, yellow to red to brown, and 

borne in glomerules 2 to 3 mm wide (Blauer and 

others 1976). Sexual expression varies between 

populations and with ploidy level. Diploids are 

usually monoecious, with a few exceptions. At 

higher ploidy levels (tetraploids and hexaploids), 

fourwing saltbush is trioecious (three sexual 

states). In the trioecious condition, some plants are 

consistently female and others are consistently 

male and some are able to switch during stressful 

conditions from the less physiologically taxing 

female state to the male state (McArthur 1977, 

McArthur and others 1992). Gender differences 

appear to be adaptive and allow for greater species 

radiation (Freeman and others 1993). 

Taxonomy.—Fourwing saltbush is composed of a 

number of polyploid races with a base chromosome 

number of x = 9 (Sanderson and Stutz 1994, 2001). 

Many have been named as varieties. It appears that 

many of these are autopolyploids (Stutz 1978). The 

common form of fourwing saltbush in the 

Intermountain region (race Occidentalis) is 

tetraploid (Sanderson and Stutz 2001). Of the 

several distinct diploids the most widespread are 

race Angustifolia of the Chihuahuan Desert and 

race Linearis of the Sonoran Desert. Hexaploids 

are also generally southern in distribution (Nevada, 

New Mexico and Mexico) (Sanderson and Stutz 

2001). Polyploids higher than 6x (up to 20x) are 

usually restricted endemics (Sanderson and Stutz 

2001). 

The North American perennial saltbushes 

form a group of related species. Fourwing saltbush 

apparently hybridizes, at least to some extent, with 

all of these species (Blauer and others 1976, Stutz 

1978). 

Range.—Fourwing saltbush is a widespread shrub 

throughout much of Western North America. It 

grows from the Great Plains to the Pacific Coast 

and from Canada to Mexico at elevations ranging 

from below sea level to 2,400 m. Fourwing 

saltbush has been planted in various parts of 

Eurasia and Australia but naturalization has not 

been documented. 

Ecology.—Fourwing saltbush is adapted to desert 

climates. Mean annual precipitation within its 
94 

distribution usually ranges between 150 and 400 

mm. It grows on deep, usually halophytic, welldrained sandy soils, gravelly washes, mesas, 

ridges, and slopes. Soil salts enhance growth of 

fourwing saltbush by increasing days to wilting, 

organic matter production, water use efficiency, 

and ability to extract water by means of osmotic 

adjustments (Glenn and Brown 1998). It often 

grows intermixed with a wide variety of shrubs, 

grasses and forbs. 

Reproduction.—Flowering generally occurs 

between May and September (Blauer and others 

1976). This period can vary, however, with 

genotype and location. Flowers are wind 

pollinated, and seed matures in October. Most 

wildland stands of fourwing saltbush produce 

abundant seed 3 out of every 5 years. 

Seed fill in high quality lots may only 

reach 50 percent, although rates of 40 percent or 

lower are generally considered substandard (Meyer 

2003). Like many species of saltbush, Atriplex 

canescens exhibits high levels of dormancy 

controlled by multiple and complex mechanisms 

(Meyer 2003). Woody bracts inhibit germination 

mechanically and also contain water-soluble 

substances, salt and perhaps saponin, which inhibit 

germination (Clor and others 1989). Seed normally 

requires 20 to 30 days of stratification to assure 

uniform germination. There are about 120,000 

cleaned, dewinged seeds (utricles)/kg (Blauer and 

others 1976). 

Artificial seeding should take place during 

the fall or winter unless spring frosts threaten 

seedling survival. Fall and winter seeding promotes 

seed stratification and overall germinability. When 

sown, seeds should remain within the top 1.25 cm 

of soil. Broadcast seeding followed by chaining 

has produced excellent stands. Shrubs can also be 

established using drills alone or in separate rows 

among other species. Care should be taken to keep 

herbs from displacing fourwing saltbush seedlings. 

Seedlings grow vigorously and quickly produce a 

dense canopy although interspecific competition 

can reduce stand densities. 

Shrubs can also be grown from nursery 

stock, container stock, or by stem cuttings (Shaw 

and Monsen 1984). Rooted stem cuttings have 

been used successfully to establish shrub seed 

orchards with desirable traits (McArthur and others 

1978, 1992). Some populations reproduce 

asexually through root sprouting (Barrow 1997). 

Management.—Fourwing saltbush is a valuable 

forage shrub because it is abundant, palatable, 

provides large quantities of forage, is nutritious, 

and grows rapidly (Cibils 1998, McArthur and 

others 1983, Peterson and others 1987). Leaves, 

stems, and fruits provide browse throughout the 

year. Winter protein content has been shown to 

vary between accessions (Welch and Monsen 

1981). Relative saponin concentrations may affect 

palatability. In mixed species communities, 

herbage production of fourwing saltbush and forbs 

remains quite high. Fourwing saltbush is a 

facultative selenium absorber, making it mildly 

poisonous to browsing animals in selenium-rich 

soils (Davis 1972). 

 When propagating fourwing saltbush, it is 

important to select ecotypes that are adapted to the 

site. If possible, seed should be from a source with 

a similar climate unless the strain being planted is 

known to have a broad ecological range. Climatic 

and biological factors may affect plant persistence. 

Plants established from southern seed sources have 

not persisted in colder northern areas. However, 

seed collected from colder areas have done well at 

warmer sites (McArthur and others 1983, Plummer 

and others 1966). 

Benefits.—Fourwing saltbush is a valuable plant 

that provides livestock and wildlife habitat and 

food. It has been widely and successfully used in 

revegetation and restoration projects including soil 

stabilization (Blauer and others 1976, Plummer and 

others 1966). Its ability to adapt and to hybridize 

with other woody Atriplex can facilitate in situ

selection during revegetation (Stutz 1982). 

References 

Barrow, J.R. 1997. Natural asexual reproduction 

in saltbush Atriplex canescens (Pursh) Nutt. 

Journal of Arid Environments 36: 267-270. 

Blauer, A.C., A.P. Plummer, E.D. McArthur, R. 

Stevens, and B.C. Giunta. 1976. Characteristics 

and hybridization of important Intermountain 

shrubs. II. Chenopod family. Research Paper 

INT-177. U.S. Department of Agriculture, Forest 

Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. 42 p. 

Cibils, A.F., D.M. Swift, and E.D. McArthur. 

1998. Plant-herbivore interactions in Atriplex: 

current state of knowledge. General Technical 

Report RMRS-GTR-14. U.S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Research Station, Ogden, UT. 31 p. 

Clor, M.A., F.M.R. Al-Charachafchi, and N. 

Mahmood. 1989. Seed dormancy and 
95

germination inhibition of Atriplex canescens. 

Journal of Agriculture and Water Resources 

Research Plant Production 8: 55-64. 

Davis, A.M. 1972. Selenium accumulation in a 

collection of Atriplex species. Agronomy 

Journal. 64: 823-824.

Freeman, D.C., E.D. McArthur, S.C. Sanderson, 

and A.R. Tiedemann. 1993. The influence of 

topography on male and female fitness 

components of Atriplex canescens. Oecologia 

93: 538-547. 

Glenn, E.P. and J.J. Brown. 1998. Effects of salt 

levels on the growth and water use efficiency of 

Atriplex canescens (Chenopodicaceae) varieties 

in drying soil. American Journal of Botany 85: 

10-16. 

McArthur, E.D. 1977. Environmentally induced 

changes of sex expression in Atriplex canescens. 

Heredity 38: 97-193. 

McArthur, E.D., D.C. Freeman, L.S. Luckinbill, 

S.C. Sanderson, and G.L. Noller. 1992. Are 

triocy and sexual morphs of Atriplex canescens

genetically based: evidence from clonal studies. 

Evolution 46: 1708-1721. 

McArthur, E.D., A.P. Plummer, G.A. Van Epps, 

D.C. Freeman, and K.R. Jorgensen. 1978. 

Producing fourwing saltbush seed in seed 

orchards. In: Hyder, D. N., ed. Proceedings of 

the First International Rangeland Congress; 

1978 Aug. 14-18; Denver, CO. Denver, CO: 

Society for Range Management: 406-410. 

McArthur, E.D., R. Stevens, and A.C. Blauer. 

1983. Growth performance comparisons among 

18 accessions of fourwing saltbush (Atriplex 

canescens) at two sites in central Utah. Journal 

of Range Management. 36: 78-81. 

Meyer, S.E. 2003. Atriplex L. In: Woody Plant 

Seed Manual, http://www.wpsm.net [not paged]. 

Peterson, J.L., D.N. Ueckert, R.L. Potter, and J.E. 

Huston. 1987. Ecotype variation in western 

Texas fourwing saltbush. Journal of Range 

Management. 40: 361-366. 

Plummer, A.P., S.B. Monsen, and D.R. 

Christensen. 1966. Fourwing saltbush, a shrub 

for future game ranges. Publication 66-4. Salt 

Lake City, UT: Utah State Department of Fish 

and Game. 12 p. 

Sanderson, S.C. and H.C. Stutz. 1994. 

Chromosome numbers in Mojave and Sonoran 

Desert Atriplex canescens (Chenopodiaceae). 

American Journal of Botany 81: 1045-1053. 

Sanderson, S.C. and H.C. Stutz. 2001. 

Chromosome races of fourwing saltbush 

(Atriplex canescens), Chenopodiaceae. In: E.D. 

McArthur and D.J. Fairbanks, comps. 

Proceedings—shrubland ecosystem genetics and 

biodiversity; 2000 June 13-15; Provo, UT. Proc. 

RMRS-P-21. U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research 

Station, Ogden, UT. p. 75-88. 

Shaw, N. and S.B. Monsen. 1984. Nursery 

propagation and outplanting of bareroot 

chenopod seedlings. In: A.R. Tiedemann, E.D. 

McArthur, H.C. Stutz, R Stevens, and K.L. 

Johnson, K., comps. Proceedings-symposium on 

the biology of Atriplex and related chenopods, 

1983 May 2-6; Provo, UT. Gen. Tech. Rep. 

INT-172. U.S. Department of Agriculture, Forest 

Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. p. 251-260. 

Stutz, H.C. 1982. Broad gene pools required for 

disturbed lands. In: E.F. Aldon and W. Oaks W. 

eds. Reclamation of mined lands in the 

southwest: Proceedings of the symposium; 1982 

Oct. 20-22; Albuquerque, NM. Soil 

Conservation Society of America, New Mexico 

Chapter, Albuquerque, NM. p.113-118. 

Stutz, H.C. 1978. Explosive evolution of perennial 

Atriplex in western America. Great Basin 

Naturalist Memoirs. 2: 161-168. 

Welch, B.L. and S.B. Monsen. 1981. Winter crude 

protein among accessions of fourwing saltbush 

grown in a uniform garden. Great Basin 

Naturalist. 41: 343-346.
96 

Atriplex confertifolia (Torr. & Frém.) S. Wats. shadscale 

CHENOPODIACEAE 

Synonyms: Obione confertifolia Torr. & Fremont 

 Atriplex collina Woot. & Standl. 

Stewart C. Sanderson, E. Durant McArthur, and Jeffrey R. Taylor

Illustration source: USDA—Forest Service collection, 

Hunt Institute 

General Description.—Shadscale is a compact 

spinescent shrub that is typically 15 to 120 cm tall 

and 30 to 170 cm wide. Branches are scruffy when 

young but become rigid, brittle, and spiny with 

age. Leaves are nearly circular to elliptic, oval, or 

oblong, 9 to 25 mm long and 4 to 20 mm wide 

(Blauer and others 1976). Flowers are imperfect. 

Staminate (male) flowers are arranged in 

glomerate inflorescences borne in leaf axils. 

Pistilate (female) flowers are enclosed by foliose 

bracts, 5 to 12 mm long, broadly ovate to almost 

round, united at the base, and have entire, free, 

somewhat spreading margins. Staminate flowers 

are small and inconspicuous. Populations may 

have biased sex ratios and can include monocieous 

as well as dioecious individuals (Freeman and 

McArthur 1984). 

Taxonomy.—Shadscale includes several 

chromosome races ranging from diploid (2n = 2x 

= 18) to decaploid (2n = 10x = 90). Plant 

similarities and the paucity of close relatives 

suggest that these races are largely autoploid 

(Stutz and Sanderson 1983). Of all the races, 

tetraploids are most common (Sanderson and 

others 1990). Generally, polyploid populations 

occur in uniform stands at lower elevations and on 

valley floors while diploid forms occur on uplands 

in more diverse communities and are usually 

larger in stature (Sanderson and others 1990, Stutz 

and Sanderson 1983). 

Range.—Shadscale is present throughout much of 

Western North America, from Montana in the 

north to Texas and Chihuahua in the south, and 

from North Dakota in the east to California and 

Oregon in the west (Stutz and Sanderson 1983). It 

usually grows at elevations from 460 to 2,100 m. 

Among the woody chenopods, it ranks next to 

fourwing saltbush [Atriplex canescens (Pursh) 

Nutt.] and winterfat [Ceratoides lantata (Pursh) J. 

T. Howell] in frequency and distribution. 

Ecology.—Shadscale usually grows on finetextured alkaline soils, although it can be found on 

coarser, sandier soils (Blaisdell and Holmgren 

1984, McArthur and others 1978). It can occur in 

nearly pure stands, but also grows intermixed with 

many other shrubs, especially of the chenopod 

and sunflower families, and with junipers 

(Juniperus L.) and grasses. It tolerates alkali soils 

better than most of these associates (McArthur and 

others 1978). Relationships between microhabitat, 

synecology, and physiology have explained 

differences in distribution patterns within 

intermixed stands of shadscale and winterfat 

(Caldwell and others 1977, West and Gasto 1978). 

Shadscale grows slowly, has a relatively short life 

span and can be killed by drought or high water 

(Blaisdell and Holmgren 1984, McArthur and 

others 1978). High water can kill plants directly, 

lead to anaerobiosis, induce abiotic disease, or 

predispose plants to biotic pathogens, all of which 

may have contributed to die-offs over the past 

decades (Wallace and Nelson 1990). Shadscale 
97

typically grows in arid climates where mean 

annual precipitation can range between 180 and 

360 mm. 

Reproduction.—Shadscale usually blooms during 

April and May. In isolated locations populations 

may bloom as early as January and as late as June. 

Utricles mature, in general about mid-October. 

Shadscale seeds are largely dormant after 

maturation but respond to cold treatment (chilling) 

through dry after-ripening (Meyer and others 

1998). Chilling and leaching soluble inhibitors 

from seeds has been shown to increase 

germination (Garvin and others 1996). 

Interestingly, seeds from warm desert populations 

responded to chilling more and after-ripened 

quicker than those from cold deserts (Meyer and 

others 1998). Ecotypic germination variation is 

common in shadscale. 

 Shadscale spreads naturally through seed 

but is difficult to establish by artificial seeding 

(McArthur and others 1978, Monsen and 

Richardson 1984). Low seed fill often compounds 

seed dormancy and germination problems. Fall 

seeding is recommended because it reduces the 

effects of dormancy, and utricles are more likely to 

break down before spring. Seeds should remain 

between 6 and 13 mm of the soil surface to 

improve the likelihood of germination and 

establishment. Soil type and seedbed preparations 

should be considered prior to seeding. Seed should 

be planted at rates of 2.2 to 4.5 kg of pure live 

seed/ha and in separate rows from other seeded 

species. Seedlings grow slowly and compete 

poorly with herbs and aggressive grasses, 

particularly within the first 2 years. Diploid races 

from pinyon-juniper woodlands and big sagebrush 

communities establish more readily in biologically 

diverse communities than other races. 

Shadscale has also been successfully 

transplanted when moisture conditions are right 

during spring or fall (Luke and Monsen 1984). 

Transplants can be from container or bare-root 

stock. Survival is usually poor unless care is taken 

during handling and planting. Young shrubs are 

particularly susceptible to overwatering. 

Transplanting is most successful if plants are 

dormant at the time of planting. 

Management.—Despite its spiny nature, 

shadscale is grazed by livestock and game. Forage 

value is directly related to leaf persistence, which 

is a characteristic that varies by habitat. Many 

species prefer the nutritious seeds (USDA 1937). 

Livestock and wildlife tend to seek seed and leaf 

accumulations under parent shrubs after leaves and 

fruits have fallen in autumn. Grazing pressure can 

reduce more palatable shrub species, such as 

budsage and blacksage, resulting in increased 

shadscale populations (Blaisdell and Holmgren 

1984). Shadscale may be more effective in 

restoring and stabilizing salty areas than fourwing 

saltbush or other saltbushes because it is more salt 

tolerant (McArthur and others 1978). In large 

stands, shadscale should be managed rather than 

mechanically controlled. Managing for shadscale 

is easier than reducing shrub densities and seeding 

in new species because limited soil moisture and 

precipitation make it difficult to manipulate these 

plant communities (Bleak and others 1965). 

Aggressive annuals, such as cheatgrass, usually 

invade such areas if they are disturbed. 

Benefits.—Shadscale is a genetically and 

morphologically diverse native shrub that occupies 

a considerable range throughout the West. It 

provides soil cover, habitat for birds, and forage 

for domestic livestock and wildlife. Some small 

passerine birds, such as horned lark (Eremophila 

alpestris), Brewer’s sparrow (Spizella breweri), 

and sage thrashers (Oreoscoptes montanus) nest in 

shadscale stands (Medin 1990). Healthy stands 

also reduce the proliferation of exotic, undesirable 

annuals. In addition, seeds were historically 

ground into flour by Native-Americans for bread 

(Stubbendieck and others 1997). 

References 

Blaisdell, J.P. and RC. Holmgren. 1984. Managing 

Intermountain rangelands-salt-desert shrub 

ranges. General Technical Report INT-163. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 52 p. 

Blauer, A.C., A.P. Plummer, E.D. McArthur, R. 

Stevens, and B.C. Giunta. 1976. Characteristics 

and hybridization of important Intermountain 

shrubs. II. Chenopod family. Research Paper 

INT-177. U.S. Deptment of Agriculture, Forest 

Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. 42 p. 

Bleak, A.T., N.C. Frischknecht, A.P. Plummer, 

and R.E. Eckert, Jr. 1965. Problems in artificial 

and natural revegetation of the arid shadscale 

vegetation zone of Utah and Nevada. Journal of 

Range Management. 18: 59-65. 

Caldwell, M.M., R.S. White, R.T. Moore, and L.B. 

Camp. 1977. Carbon balance, productivity, and 
98 

water use by cold desert shrub communities 

dominated by C3 and C4 species. Oecologia. 29: 

275-300. 

Freeman, D. C. and E.D. McArthur. 1984. The 

relative influences of mortality nonflowering, 

and sex change on the sex ratios of six Atriplex 

species. Botanical Gazette 145: 385-394. 

Garvin, S.C., S. E. Meyer, and S. L. Carlson. 1996. 

Seed germination studies in Atriplex confertifolia

(Torr. & Frem.) Wats. In: J.R. Barrow, E.D. 

McArthur, R.E. Sosebee, and R.J. Tausch, 

comps. Proceedings: shrubland ecosystem 

dynamics in a changing environment. 1995 May 

23-25; Las Cruces, NM. General Technical 

Report INT-GTR-338. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Research Station, Ogden, UT. p. 165-169. 

Luke, F. and S.B. Monsen. 1984. Methods and 

costs for establishing shrubs on mined lands in 

southwestern Wyoming. In: A.R. Tiedemann, 

E.D. McArthur, H.C. Stutz, R. Stevens, and K.L. 

Johnson, comps. Proceedings-symposium on the 

biology of Atriplex and related chenopods, 1983 

May 2-6; Provo, UT. Gen. Tech. Rep. INT-172. 

U.S. Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. p. 286-292. 

McArthur, E.D., A.P. Plummer, and J.N. Davis. 

1978. Rehabilitation of game range in the salt 

desert. In: K.L Johnson, ed. Wyoming 

shrublands: proceedings 7th Wyoming shrub 

ecology workshop; 1978 May 31-June 1; Rock 

Springs, WY. University of Wyoming, Division 

of Range Management, Laramie, WY. p. 23-50. 

Medin, D.E. 1990. Birds of a shadscale (Atriplex 

confertifolia) habitat in east central Nevada. 

Great Basin Naturalist 50: 295-298. 

 

Meyer, S.E., S.L. Carlson, and S.C. Garvin. 1998. 

Seed germination regulation and field seed bank 

carryover in shadscale (Atriplex confertifolia: 

Chenopodiaceae). Journal of Arid Environments 

38: 255-267. 

Monsen, S.B. and B.Z. Richardson. 1984, Seeding 

shrubs with herbs on a semiarid mine site with 

and without topsoil. In: A.R. Tiedemann, E.D. 

McArthur, H.C. Stutz, R. Stevens, and K.L. 

Johnson, comps. Proceedings: Symposium on 

the biology of Atriplex and related chenopods, 

1983 May 2-6; Provo, UT. Gen. Tech. Rep. 

INT-172. U.S. Department of Agriculture, 

Forest Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. p. 298-305. 

Sanderson, S.C., H.C. Stutz, and E.D. McArthur 

1990. Geographic differentiation in Atriplex 

confertifolia. American Journal of Botany 77: 

490-498. 

Stubbendieck, J., S.L Hatch, and C.H. Butterfield 

1997. North American Range Plants. Fifth 

Edition. University of Nebraska Press, Lincoln, 

NB. 305 p. 

Stutz, H.C. and S.C. Sanderson. 1983. 

Evolutionary studies of Atriplex: chromosome 

races of A. confertifolia (shadscale). American 

Journal of Botany 70: 1536-1547. 

U.S. Department of Agriculture, Forest Service. 

1937. Range Plant Handbook. U.S. Government 

Printing Office, Washington, DC. 816 p. 

Wallace, A. and D.L. Nelson. 1990. Wildland 

shrub dieoffs following excessively wet periods: 

a synthesis. In: E.D. McArthur, E.M. Romney, 

S.D. Smith, and P.T. Tueller, comps. 

Proceedings: Symposium on cheatgrass 

invasion, shrub die-off, and other aspects of 

shrub biology and management; 1989 April 5-7; 

Las Vegas NV. General Techical Report INT276. Ogden, UT. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Research Station. p. 84-90. 

West, N.E. and J. Gasto. 1978. Phenology of the 

aerial portions of shadscale and winterfat in 

Curlew Valley, Utah. Journal of Range 

Management. 31: 43-45. 
99

Atriplex corrugata S. Wats. mat saltbush 

CHENOPODIACEAE 

Synonyms: Atriplex nuttallii var. corrugata A. Nelson 

 Obione corrugata Ulbr. 

E. Durant McArthur, Stewart C. Sanderson, and Jeffrey R. Taylor 

General Description.—Mat saltbush, also known 

as matscale, is a low shrub that forms dense, 

prostrate (15 cm tall) and nearly white mats five to 

20 times wider than they are tall. Prostrate 

branches often form adventitious roots when in 

contact with the soil. The bark is soft, spongy, and 

white. Leaves are sessile, evergreen, oblong, 

linear-oblancelate, or spatulate, rounded at the 

apex, opposite on the lower portions of stems, and 

alternate above. Leaf blades are densely scurfy and 

7 to 18 mm wide (Blauer and others 1976, 

McArthur and others 1978, Welsh and others 

1993). Individual shrubs are commonly dioecious 

but may be monoecious. Staminate (male) flowers 

are yellow to light-brown, 3 to 6 mm wide, and 

borne in glomerulate spikes. Pistilate (female) 

flowers are enclosed by sessile or subsessile 

fruiting bracts. Fruiting bracts are 3 to 5 mm long, 

4 to 6 mm wide, and united along two-thirds of 

their length and densely tuberculate on the lower 

one-third (Blauer and others 1976, McArthur and 

others 1978, Welsh and others 1993). 

Taxonomy.—This species will hybridize with 

other saltbushes (Blauer and others 1976, Stutz 

1978, 1984). Mat saltbush forms intermediates 

with Atriplex confertifolia Torr. & Fremont and 

Atriplex cuneata A. Nelson (Welsh and others 

1993). Plants can be either diploid (2n = 18) or 

tetraploid (2n = 36) (Sanderson and Stutz, 

unpublished data). 

Range.—Mat saltbush is an endemic of the 

Colorado Plateau and Uintah Basin, found mainly 

on Mancos shale formations in eastern Utah, 

western Colorado, and northwestern New Mexico 

between 1,200 and 2,100 m (Hall and Clements 

1923; Hanson 1962). 

Ecology.—Mat saltbrush grows on fine textured, 

often saline, soils. It tolerates up to 13,000 ppm 

soluble salts and is often the only perennial plant 

present in such high-saline environments (Hanson 

1962). Although it is probably the most halophytic 

member of the genus, in less saline habitats, 

patches of mat saltbush may alternate with patches 

of winterfat [Ceratoides lanata (Pursh) 

J.T.Howell], cuneate saltbush (Atriplex cuneata A. 

Nelson), shadscale (Atriplex confertifolia Torr. & 

Fremont), black greasewood (Sarcobatus 

vermiculatus Torr.), budsage (Artemisia 

spinescens Eaton), and greenmolly summercypress 

(Kochia americana S. Wats.). It grows in areas 

with 150 to 280 mm of annual precipitation. 

Reproduction.—Blooming occurs between March 

and May and fruits (utricles) ripen 6 to 10 weeks 

later. Seed production is usually modest, although 

some seed is usually produced even during 

drought. 

 Because germination is near zero without 

cold treatment, fall is normally the recommended 

season for artificial seeding. Where possible, 

midwinter planting is also successful. Preparatory 

measures are usually unnecessary if the site 

previously supported mat saltbush and is not 

infested with weeds. Recommended seeding rates 

range between 2.2 and 4.5 kg of pure live seed/ha. 

Seed should be sown in separate rows and covered 
100 

with a drag or harrow. Seeds should be planted 

near the soil surface and no more than 1.3 cm 

deep. Seedlings emerge rapidly and grow 

vigorously during the first year. Growth continues 

until soil moisture becomes limiting (usually by 

mid-summer). There are about 220,000 cleaned 

seed per kg (Blauer and others 1976). 

Transplants from bare root stock, 

wildings, or containers have all established 

successfully. Wildings and bare root stock tolerate 

considerable distress. Transplants survive 

surprisingly well, even when planted on seemingly 

infertile soils. Young plants have been 

successfully transplanted into former black 

greasewood communities in the Great Basin 

(Blauer and others 1976). 

Management.—Mat saltbush is valuable winter 

forage for wildlife and domestic livestock. It often 

grows interspersed among other plant communities 

on shale outcrops at medium elevations. Shrubs 

often grow on windblown slopes and provide 

important, accessible winter forage. Big game 

have been known to congregate on these areas 

during periods of deep snow accumulation. Shrubs 

tolerate grazing and trampling and maintain vigor 

even after serious abuse. 

 Mat saltbush has been seeded with 

selected grasses including Russian wildrye 

(Elymus junceus Fisch.), tall wheatgrass 

[Agropyron elongatum (Host) P.Beauv.], and 

crested wheatgrass [Agropyron cristatum (L.) 

Gaertn.]. Although these species may increase 

forage production, they do not usually persist in 

dense stands. Grasses and forbs will suppress 

shrub establishment, but grasses interseeded into 

established shrub stands do not reduce shrub 

density or vigor. 

 This species is useful in revegetation and 

restoration projects and can help reclaim disturbed 

areas with heavy textured soils such as road 

construction sites or mine disturbances. 

Benefits.—Mat saltbush is a native shrub that 

occupies a specialized niche in Western 

ecosystems. Its ability to grow on fine textured, 

often saline soils helps reduce soil erosion on sites 

that are inhospitable to many shrubs. It also 

provides important forage for wildlife during 

winter when other sources of forage may be 

inaccessible. 

References 

Blauer, A.C., A.P. Plummer, E.D. McArthur, R. 

Stevens, and B.C. Giunta. 1976. Characteristics 

and hybridization of important Intermountain 

shrubs. II. Chenopod family. Research Paper 

INT-177. U.S. Department of Agriculture, Forest 

Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. 42 p. 

Hall, H.M. and F.E. Clements. 1923. The 

phylogenetic method in taxonomy, the North 

American species of Artemisia, Chrysothamnus, 

and Atriplex. Publication 326. Carnegie 

Institution of Washington, Washington, DC. 355 

p. 

Hanson, C.A. 1962. Perennial Atriplex of Utah and 

the northern deserts. M.S. thesis. Brigham 

Young University, Provo, UT. 133 p. 

McArthur, E.D., A.P. Plummer, and J.N. Davis. 

1978. Rehabilitation of game range in the salt 

desert. In: K.L. Johnson, ed. Wyoming 

shrublands: proceedings of the 7th Wyoming 

shrub ecology workshop; 1978 May 31-June1; 

Rock Springs, WY. University of Wyoming, 

Division of Range Management, Laramie, WY. 

p. 23-50. 

Sanderson, S.C., and H.C. Stutz. 2003. 

Unpublished data. U.S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Research Station, Provo, UT 

Stutz, H.C. 1978. Explosive evolution of perennial 

Atriplex in western America. Great Basin 

Naturalist Memoirs. 2: 161-168. 

Stutz, H.C. 1984. Atriplex hybridization in western 

North America. In: A.R. Tiedemann, E.D. 

McArthur, H.C. Stutz, R. Stevens, and K.L. 

Johnson, comps. Proceedings: Symposium on 

the biology of Atriplex and related chenopods, 

1983 May 2-6; Provo, UT. General. Technical 

Report INT-172. U.S. Department of 

Agriculture, Forest Service, Intermountain Forest 

and Range Experiment Station, Ogden, UT. 

p. 25-27. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1993. A Utah flora. 2nd Ed., 

revised. Brigham Young University Print 

Services, Provo, UT. 986 p. 
101

Atriplex cuneata A. Nels. Castle Valley clover saltbush 

CHENOPODIACEAE 

Synonyms: Atriplex gardneri var. cuneata (A. Nels.) S.L.Welsh 

 Atriplex acanthocarpa var. cuneata Jones 

 Atriplex nuttalli subsp. cuneata H.M. Hall & Clem. 

E. Durant McArthur, Stewart C. Sanderson, and Jeffrey R. Taylor 

General Description.—Castle Valley clover 

saltbush, also known as Castle Valley saltbush and 

cuneate saltbush, is a low shrub 10 to 45 cm tall 

with a more or less prostrate, woody, muchbranched base with erect branches. Leaves are 

evergreen, light-grey green, spatulate to broadly 

elliptic, 2 to 6 cm long and 0.5 to 2.5 cm wide 

(Hall and Clements 1923). Staminate (male) 

flowers are dark, almost black, and are borne in 

glomerules arranged in panicles. Pistilate (female) 

flowers are borne in axillary clusters and consist of 

pistils enclosed by wingless bracts. At maturity, 

bracts are 5 to 9 mm wide, irregularly toothed 

along their margins, and have numerous, crest-like 

tubercles on their side. Sex expression in Castle 

Valley clover saltbush is trioecious (Freeman and 

McArthur 1984). In this system, some individuals 

are consistently male, some are consistently 

female, and others can switch between the less 

energistically expensive male state, and the more 

taxing female state, depending on climatic 

conditions. 

Taxonomy.—Castle Valley clover saltbush is one 

of several related saltbush species. It hybridizes 

with a number of other saltbushes, particularly 

with fourwing saltbush (Atriplex canescens

(Pursh) Nutt.). Atriplex cuneata is tetraploid (2n = 

36). Some taxonomists consider it to be a variety 

or subspecies of other woody saltbush species. Its 

exact taxonomic distinction remains somewhat 

unclear. Hanson (1962) described one subspecies 

for Atriplex cuneata, ssp. introgressa, found to be 

diploid (2n = 18) (Sanderson and Stutz, 

unpublished data). It is now considered to be a 

variant of A. welshii Hanson, also diploid. Atriplex 

cuneata hybridizes with other perennial Atriplex, 

especially with A. confertifolia Torr. & Freemont, 

a hybrid that has been named as A. X 

neomexicana. Populations of this hybrid and its 

derivatives may expand enormously during 

favorable years but are eradicated during drought 

(Sanderson and Stutz, unpublished data). 

Range.—The distribution of Castle Valley clover 

saltbush includes portions of eastern Utah, 

southwestern Colorado, and northern New 

Mexico. It usually grows at elevations between 

1,220 and 2,170 m (Welsh and others 1993). 

Ecology.—This species often grows on fine 

textured soils, typically clay or soils derived from 

shale, that are moderately to highly alkaline. It 

grows on variably saline soils and may be 

associated with many other shrubby chenopod 

species and several of the sagebrush (Artemisia L.) 

species. It is often the dominant or codominant 

shrub with shadscale (Atriplex confertifolia Torr. 

& Frem.) or mat saltbush (Atriplex corrugata S. 

Wats.). It grows in areas with 160 to 310 mm of 

precipitation. 

Reproduction.—Castle Valley clover saltbush 

blooms in April and May, or sometimes later 

depending upon rainfall. Fruits (utricles) ripen 
102 

about 7 weeks after blooming (Hanson 1962). 

Annual seed production is often erratic and only 

about 50 percent of utricles contain viable seed. 

Germination rates are also erratic and vary among 

ecotypes. Fruits are small and generally number 

about 180,000/kg (Blauer and others 1976). 

 This species should not be seeded in areas 

beyond its natural distribution. It is poorly adapted 

to big sagebrush (Artemisia tridentata Nutt.) and 

other upland habitats but can be seeded into most 

salt desert shrublands. Young plants grow 

vigorously and are competitive but can be 

suppressed by perennial grasses and annual weeds. 

It should not be seeded directly with herbaceous 

species but competes well when seeded with 

fourwing saltbush [Atriplex canescens (Pursh) 

Nutt.], green ephedra (Ephedra viridis Coville), 

and spiny hopsage [Grayia spinosa (Hook.) Moq.]. 

Management.—Castle Valley clover saltbush is 

preferred by livestock throughout the year. It 

remains green and succulent, even through winter. 

It is an important source of protein in winter 

months for both livestock and game when most 

other forages are dry (McArthur and others 1978). 

A stabilized hybrid between fourwing saltbush and 

Castle Valley clover saltbush has great potential to 

improve the forage quality on winter ranges. Such 

a hybrid would be taller than Castle Valley clover 

saltbush and retain more green foliage than 

fourwing saltbush. 

This species shows particular promise in 

winter game range restoration and on disturbed 

alkaline soils. Like other woody saltbushes, it 

grows on freshly disturbed soils, such as exposed 

substrata created by mining or road construction 

activities (Blauer and others 1976, McArthur and 

others 1978). 

Benefits.—Castle valley clover saltbush is a native 

shrub that is adapted to fine textured soils that are 

inhospitable to many other shrubs. It adds 

biological and structural diversity to the plant 

communities on these otherwise difficult sites. It 

helps stabilize the soil, can be useful in wildland 

restoration projects, and provides valuable and 

nutritious forage for wildlife and livestock on 

Western shrublands. 

References 

Blauer, A.C., A.P. Plummer, E.D. McArthur, R. 

Stevens, and B.C. Giunta. 1976. Characteristics 

and hybridization of important Intermountain 

shrubs. II. Chenopod family. Research Paper 

INT-177. U.S. Department of Agriculture, Forest 

Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. 42 p. 

Freeman, D.C. and E.D. McArthur. 1984. The 

relative influences of mortality, non-flowering, 

and sex change on the sex ratios of six Atriplex 

species. Botanical Gazette 145: 385-394. 

Hall, H.M. and F.E. Clements. 1923. The 

phylogenetic method in taxonomy, the North 

American species of Artemisia, Chrysothamnus, 

and Atriplex. Publication 326. Carnegie 

Institution of Washington, Washington, DC. 

355 p. 

Hanson, C.A. 1962. Perennial Atriplex of Utah and 

the northern deserts. M.S. thesis. Brigham 

Young University, Provo, UT. 133 p. 

McArthur, E.D., A.P. Plummer, and J.N. Davis. 

1978. Rehabilitation of game range in the salt 

desert. In: K.L. Johnson, ed. Wyoming 

shrublands: proceedings of the 7th Wyoming 

shrub ecology workshop; 1978 May 31-June1; 

Rock Springs, WY. University of Wyoming, 

Division of Range Management, Laramie, WY. 

p. 23-50. 

Sanderson, S.C. and H.C. Stutz. 2003. 

Unpublished data. U.S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Research Station, Provo, UT 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1993. A Utah Flora. Second 

edition, revised. Brigham Young University 

Print Services, Provo, UT. 986 p. 
103

Baccharis pilularis DC. coyote brush 

ASTERACEAE 

Synonyms: None

Christopher Ross

General Description—Coyote brush, also known 

as chaparral broom, is a shrub to 3 m tall, with 

glabrous, sticky, oblanceolate to obovate leaves 8 

to 55 mm long, with toothed or entire margins. It 

may either be spreading or ascending in form 

(Sundberg 1993). 

Range.—Coyote brush occurs from Oregon to 

northern Mexico. In California it ranges from the 

northwestern coast south, including the peninsular 

and transverse ranges and the Channel Islands, and 

eastward to the Sierra Nevada foothills (Sundberg 

1993). 

Ecology.—Coyote brush is found on coastal bluffs 

and oak woodlands from sea level to 1500 m. It 

sometimes occurs on serpentine soils (Sundberg 

1993). Coyote brush may invade stabilized dunes 

in northern California, especially after yellow bush 

lupine has established (Pickart and Sawyer 1998). 

It has been very invasive in grassland areas near 

Oakland, California, apparently as a result of 

decreased grazing and fire, and appears to favor 

areas of bare soil (McBride and Heady 1968). It 

may be a seral stage leading to oak and bay 

woodlands in the San Francisco Bay area (McBride 

1974). The annual grass Bromus mollis L. has been 

found to interfere competitively with Baccharis

seedlings, especially in dry conditions (Da Silva 

and Bartolome 1984). In coastal regions of central 

California, a cecidomyiid midge, Rhopalomyia 

californica Felt forms terminal galls on coyote 

brush, and the midges are in turn preyed upon by 

various parasites (Latto and Briggs 1995). 

Reproduction.—Flowers are headed in leafy 

panicles, with staminate heads 3.2 to 5 mm long, 

and pistallate heads 3.5 to 5 mm long. Phyllaries 

(involucre bracts) are in five to six series, linearlanceolate, hairy, and glandular, with convex 

receptacle. Staminate flowers are 19 to 26, with 4 

to 7 mm corollas and 3.5 to 4.5 mm pappus. 

Pistillate flowers number 19 to 43, with 2.5 to 3.5 

mm corollas. Seeds are glabrous, 1 to 2 mm long 

with 8 to 10 ribs and a 5.5 to 9 mm pappus 

(Sundberg 1993). Seed dispersal begins in 

November, with germination in winter and early 

spring (McBride and Heady 1968). Coyote brush is 

sometimes propagated horticulturally from tip 

cuttings. 

Fire Effects.—Coyote brush has been observed to 

burn readily in coastal California, despite the fact 

that it is sometimes marketed as a fire-resistant 

species. Coyote brush seedlings and young plants 

are susceptible to fire damage, but older plants 

quickly resprout from their base after fire (McBride 

and Heady 1968).

Growth and Management.—Coyote brush 

seedlings are adversely affected by grazing and 

trampling by livestock, although grazing animals 

may also foster its spread by decreasing grass 

cover (McBride and Heady 1968). However, 

Williams and others (1987) found no proximal link 

between decreased grazing and fire and coyote 

brush invasion. Mowing of specimens used as 
104 

ground cover is effective in controlling height 

without long-term detrimental effects (Hodel and 

Pittenger 1994). Coyote brush may be removed by 

burning and then pulling out the roots. 

 Benefits.—Although several cultivated varieties 

are often sold as a landscaping shrub or ground 

cover, some growers caution that it is neither an 

especially attractive plant, nor as fire-proof as 

sometimes stated. It is of low value as livestock 

forage (McBride and Heady 1968). Coyote brush 

was used by Native Americans as a tea for use on 

poison oak rash (Timbrook 1987). It is considered 

problematic in recreation areas due to its 

impenetrability and frequent association with 

poison oak (Rhus diversiloba T. & G.) (McBride 

and Heady 1968). 

References 

Da Silva, P.G. and J. W. Bartolome. 1984. 

Interaction between a shrub, Baccharis pilularis 

subsp. consanguinea (Asteraceae) and an annual 

grass, Bromus mollis (Poaceae), in coastal 

California. Madrono 31: 93-101. 

Hodel, D.R. and D.R. Pittenger. 1994. Responses 

of eight groundcover species to renovation by 

mowing. Journal of Environmental Horticulture 

12: 4-7. 

Latto, J. and C.J. Briggs. 1995. Factors affecting 

distribution of the gall forming midge 

Rhopalomyia californica (Diptera: 

Cecidomyiidae). Environmental Entomology 

24(3): 679-686. 

McBride, J. 1974. Plant Succession in the 

Berkeley Hills, California. Madrono 22: 317-

380. 

McBride, J. and H.F. Heady. 1968. Invasion of 

Grassland by Baccharis pilularis DC. Journal 

of Range Management 21: 106-108. 

Pickart, A.J. and J. O. Sawyer. 1998. Ecology and 

Restoration of Northern California Coastal Dunes. 

California Native Plant Society. Sacramento, CA. 

152 p. 

Sundberg, S. 1993. Baccharis. In J.C. Hickman, 

ed. The Jepson Manual: higher plants of 

California. University of California Press, 

Berkeley and Los Angeles. 1,400 p. 

Timbrook, J. 1987. Virtuous herbs: plants in 

Chumash medicine. Journal of Ethnobiology 

Winter 1987: 171-180. 

Williams, K., R.J. Hobbs, and S.P. Hamburg. 

1987. Invasion of an annual grassland in 

Northern California by Baccharis pilularis ssp. 

consanguinea. Oecologia 72:461-465.
105

Barleria prionitis L. porcupine flower 

ACANTHACEAE 

Synonyms: none 

John K. Francis 

General Description.—Porcupine flower, also 

called espinosa amarilla, and picanier jaune, is a 

flowering, spiny invader that reaches 1.5 m in 

height and 2.5 cm in basal diameter. It has a tap 

and lateral root system. Plants commonly have a 

single stem but may have multiple stems or 

branches from near the ground. The stems and 

branches are stiff, round, and light tan or light gray. 

The shrub is armed with 5- to 20-mm long spines 

in the leaf axils. Leaves are elliptic to oblong, 3 to 

10 cm long and 1.5 to 4 cm broad. The yellow, 

tubular flowers, 3 to 4 cm long and broad, are 

sessile in the leaf axils or in terminal branched 

spikes. The two-celled capsule is ovate-lanceolate 

and 1.3 to 2 cm long, with a sharp-pointed beak 

about 6 mm long. The seeds are 8 by 5 mm, 

flattened and covered with matted hairs (Howard 

1989, Liogier 1997). 

Range.—Porcupine flower is a native of Tropical 

East Africa and Asia (Burkill 1985). It is widely 

cultivated as an ornamental and has escaped in 

many tropical countries including Hawaii (Neal 

1965) and Puerto Rico (Liogier 1997). 

Ecology.—Porcupine flower grows in a wide 

variety of well-drained soils derived from igneous, 

metamorphic (including ultramafics) and 

sedimentary (including limestone) rocks. In the 

Puerto Rican naturalized range, it grows in areas 

receiving from about 750 to 900 mm of mean 

annual precipitation (author’s observation). Within 

the native range, it probably tolerates both higher 

and lower precipitation, but no data are available. 

Porcupine flower is found throughout the hotter 

parts of India and defoliates annually during the 

dry season (Parrotta 2001). The species is 

moderately intolerant of shade, growing in both 

full sunlight and under light forest canopies. 

Porcupine flower may grow as single plants or in 

large, tangled thickets. Dense stands eliminate 

most other ground vegetation. However, the 

species does not compete well in dense stands of 

tall grass. It is most often found on roadsides, 

bluffs and bars above streams, overgrazed range, 

disturbed areas, and farmsteads. 

Reproduction.—In the deciduous tropical forest 

zone of central India, these shrubs flower from 

September to December and fruit from January to 

April (Parrotta 2001). The flowering and fruiting 

seasons are about the same in Puerto Rico. Sixty 

fruits collected in Puerto Rico averaged 0.0998 +

0.0044 g/fruit. Air-dried seeds separated from the 

same collection averaged 0.0305 + 0.0005 g/seed 

or 33,000 seeds/kg. Forty-four percent of the seeds 

sown on commercial potting mix germinated 

between 13 and 77 days following sowing. Natural 

regeneration generally occurs within 1 or 2 m of 

the parent plant in Puerto Rico, perhaps because 

animal vectors for seed dispersal are lacking. 

Seedlings can be abundant near fruiting plants. 

Growth and Management.—Seedlings of 

porcupine flower grow slowly at first. However, 

established plants add up to 0.5 m per year to their 

height and nursery seedlings reach 0.6 m in 6

months. In Puerto Rico, these shrubs live about 4 
106 

years. Because it is spiny and generally ignored by 

cattle, the species is considered a weed in much of 

its range. Mowing, followed by spraying of the 

sprouts with broadleaf herbicide, is a general 

control measure for low shrubs and would probably 

be effective against porcupine flower. 

Benefits.—Porcupine flower furnishes cover for 

wildlife and protects the soil against erosion. It is 

widely planted as an ornamental and cultivated in 

Asia as a hedge plant (Burkill 1985). Whole-plant 

extracts of porcupine flower contain iridoid 

glycosides, barlerin, and verbascoside, which have 

shown potent activity against respiratory syncytial 

virus in vitro and may account for the plant’s use in 

treating fever and several respiratory diseases in 

herbal medicine (Balick and others 1998). Extracts 

of the plant have also been shown to effectively 

suppress the fungi Trichophyton mentagrophytes in 

vitro (Panwar and others 1979). A mouthwash 

made from root tissue is used to relieve toothache 

and treat bleeding gums (Burkill 1985). The whole 

plant, leaves, and roots are used for a variety of 

purposes in traditional Indian medicine. For 

example, the leaves are used to promote healing of 

wounds and to relieve joint pains and toothache 

(Parrotta 2001). Because of its antiseptic 

properties, extracts of the plant are incorporated 

into herbal cosmetics and hair products to promote 

skin and scalp health (Prakruti 2002, Probiotics 

New Zealand 2002, Vaipani 2002). 

References 

Balick, M., P. Blanc, M. Morgan, J.L. Chen, R. 

Cooper, M.R. Kernan, W. Nanakorn, N. 

Parkinson, E.J. Rozhon, C.A. Stoddard, and Z.J. 

Yee. 1998. New iridoids from the medicinal 

plant Barleria prionitis with potent activity 

against respiratory syncytial virus. Journal of 

Natural Products 61(10): 1,295-1,297. 

 Burkill, H.M. 1985. The useful plants of West 

Tropical Africa. Vol. 1. Royal Botanic Garden, 

Kew, UK. 960 p. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H. A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Museum 

Press, Honolulu, HI. 924 p. 

Panwar, H.S., M.M. Nauriyal, and H.C. Joshi. 

1979. In vitro screening of certain indigenous 

plants for their antimycotic activity. Veterinary 

Research Bulletin 2(2): 164-167. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CABI Publishing. Wellington, UK & New 

York. 917 p. 

Prakruti. 2002. Suddh Bhangra (maka) oil. http:// 

www.prakrutiherbals.com/hairoil.htm 2 p. 

Probiotics New Zealand. 2002. Probiotics for 

life—improving your health and quality of life 

through good bacteria. http://www.probiotics. 

co.nz/prdcts_efml.asp?ProductID=5. 3 p. 

Vaipani. 2002. Vaipani Herbal Ayurvedic 

Partisthan—A herbal ayurvedic cosmetics 

company. http://www.vaipani.com/skincare/ 

skin%20Care-Page3.html. 2 p. 
107

Batis maritima L. saltwort 

BATACEAE 

Synonyms: none 

John K. Francis 

General Description.—Saltwort, also known as 

turtleweed, pickleweed, barilla, planta de sal, 

camphire, herbe-à-crâbes, and akulikuli-kai, is a 

low, yellow-green shrub with succulent leaves. It 

is prostrate or upright and occasionally reaches 1 

m in height, 2 m in lateral extent, and 5 cm in 

basal diameter. Stems are usually multiple as 

sprouts from the root crown, and as they become 

tall and heavy, lie down and root along the stems 

forming loose mats. Weak roots with light-tan, 

corky bark form tap and lateral root systems. The 

bark is grayish white; stem wood is weak and 

brittle. The glabrous leaves are succulent, linear or 

narrowly oblanceolate and round or three- to fourangled in cross section. Tiny, white male and 

female flowers occur on different plants. Fruits are 

axillary drupaceous, yellow-green syncarps 1 to 2 

cm long (Liogier 1985, Stevens and others 2001, 

Wiggins and Porter 1971). 

Range.—Saltwort is native to coastal areas of 

southern United States including California, 

Mexico, Central America, South America to Brazil 

and Peru, the West Indies, and the Galapagos 

Islands (Howard 1988, Wiggins and Porter 1971). 

It has been planted in a number of areas in the 

tropics and has naturalized in at least Hawaii (Neal 

1965). 

Ecology.—Saltwort is uncommon to abundant in 

low-laying areas near seashores. It grows in salt 

marshes, at the upper edge of tidal flats, at the 

edge of mangrove stands, and between scattered 

mangroves. It is recognized as a major colonizer 

after mangroves are destroyed by hurricanes. 

Although it is not a water plant, it can endure brief 

flooding and long periods of waterlogged soils 

(Nelson 1996). Saltwort grows slowly in soils 

with high salt concentrations and areas with 

seawater overwash where it suffers little 

competition from other plants. The species 

manages salts by sequestering them in cell 

vacuoles and eventually shedding the leaves 

(Barbuda Turf Company 2002). It also grows in 

soils without salt but is vulnerable to competition 

from nonhalophytes. The soils are usually sandy, 

marly, or gravely. Deposits of wrack (dead plant 

material) by high tides have been shown to be 

beneficial to the species (Pennings and Richards 

1998). Saltwort is intolerant of shade. The species 

is not seriously affected by insects, disease, or 

grazing. 

Reproduction.—In Florida, saltwort flowers in 

the spring and fruits in the summer (Long and 

Lakela 1976). Flowering and fruiting occurs yearround in Puerto Rico. Little is known about seed 

production or germination. Most effective 

reproduction of the species appears to be 

vegetative. Sprouting from the root crown occurs 

with and without disturbance. Layering is a 

constant process of prostrate stems. New plants 

can be started by cuttings and probably broken 

pieces of plants are carried to new habitat by water 

and machinery. It is speculated that whole plants 

washed to sea during torrential rains in South 
108 

America were carried by currents to establish the 

species in the Galapagos Islands (Trillmich 2002). 

Growth and Management.—Saltwart plants 

established in experiments from cuttings and 

irrigated with fresh and brackish water grew 65 cm 

and reached plant dry weights of 120 g in 13 

weeks. Plants receiving saltier water grew much 

less but still added height and weight with water 

almost twice as salty as sea water. Tissue ash 

concentrations did not differ significantly between 

treatments (Miyamoto and others 1996). Wild 

plants in stands in Puerto Rico appear to extend 

their stems about 1 m/year. Leaf succulence 

increases as much as 1.5 times during the dry 

season to maintain a near static solute (chiefly Na+

, 

Cl-

, and S04

--) concentration (Lüttge and others 

1989). Probably all planting for landscaping and 

environmental restoration is by rooted cuttings or 

uprooted wild plants. Except in Hawaii, where it is 

being suppressed as an exotic plant (Big Island 

Invasive Species Committee 2002), there is little 

reason to control stands of saltwort. It grows in 

disturbed areas where few other plants can survive. 

Benefits.—The principal benefit of saltwort is that 

it grows in, covers, and protects salty low-laying 

areas where few other species will grow. It is used 

as an ornamental and ground cover on similar 

sites. Salt tolerance of the species allows it to be 

irrigated with brackish and sea water, as well as 

fresh water (Miyamoto and others 1996). Ashes of 

the plant were once used in the manufacture of 

glass and soap. The leaves are sometimes eaten as 

a salad (Neal 1965). However, it is toxic in large 

quantities (Austin 1998). Saltwort serves as the 

larval and adult hosts for the great Southern white 

and Eastern pigmy blue butterflies (Florida 

Cooperative Extension Service 2002) and is an 

important food source for the Galapagos marine 

iguana (Amblyrhynchus cristatus) (Wrege 2002). 

The species has applications in herbal medicine to 

treat eczema, psoriasis, and other skin conditions, 

rheumatism, gout, blood and vein disorders 

(Liogier 1990). 

References

Austin, D.F. 1998. Poisonous plants of southern 

Florida. http://www.fau.edu/divdept/science/ 

envsci/poison-pl.html. 9 p. 

Barbuda Turf Company. 2002. Halophytes in 

deep. Barbuda Turf Company, Codrington 

Village, Barbuda, Antigua-Barbuda, West 

Indies. http://www.geocities.com/barbudaturf/ 

hallophites%20in%20deep.htm. 3 p. 

Big Island Invasive Species Committee. 2002. Pest 

prevention/suppression: eradication of incipient 

forest pest plants by the Big Island Species 

Committee. http://www.hear.org/ 

operationmicronia/2002biiscannual.pdf. 9 p. 

Florida Cooperative Extension Service. 2002. 

Butterfly gardening in Florida. University of 

Florida, Gainesville, FL. http://edis.ifas.ufl.edu/ 

BODY_UW057. 24 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain, 

MA: Arnold Arboretum, Harvard University. 

673 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Lüttge, U., M. Popp, E. Medina, W.J. Cram, M. 

Diaz, H. Griffiths, H.S.J. Lee, C. Schäfer, J.A. 

Smith, and K.H. Stimmel. 1989. Ecophysiology 

of xerophytic and halophytic vegetation of a 

coastal alluvial plain in Northern Venezuela. 

New Phytologist 111(2): 283-291. 

Miyamoto, S., E.P. Glenn, and M.W. Olsen. 1996. 

Growth, water use and salt uptake of four 

halophytes irrigated with highly saline water. 

Journal of Arid Environments 32(2): 141-159. 

Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub. 

50. Bernice P. Bishop Museum Press, Honolulu, 

HI. 924 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

 391 p. 

Pennings, S.C. and C.L. Richards. 1998. Effects of 

wrack burial in salt-stressed habitats: Batis 
109

maritima in a southwest Atlantic salt marsh. 

Ecography 21(6): 630-638. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.H. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs of Systematic Botany Vol. 85, No. 

1. Missouri Botanical Garden Press. 943 p. 

Trillmich, F. 2002. El Niño in the Galapagos 

Islands: a natural experiment. International 

Council of Scientific Unions. http://www.icsuscope.org/downloadpubs/scope45/ch01-1.3.1. 

html. [not paged]. 

Wiggins, I.L. and D.M. Porter. 1971. Flora of the 

Galápagos Islands. Stanford University Press, 

Stanford, CA. 998 p. 

Wrege, P.H. 2002. Bee-eater research: Galapagos! 

Department of Ecology and Evolutionary 

Biology, Cornell University, Ithaca, New York. 

http://www.eeb.cornell.edu/wrege/pages/iggires.

html. 4 p. 
110 

Bidens menziesii (Gray) Sherff ko'oko'olau

ASTERACEAE 

Synonymns: Coreopsis menziesii A. Gray 

 Bidens menziesii subsp. filiformis (Sherff) Ganders & Nagata

 Bidens m. var. filiformis Sherff 

 Bidens lepida Degener & Sherff 

Sarah A. Taylor and Randy S. Senock 

 

General Description.―Bidens menziesii, known 

as ko'oko'olau, is a sparingly branched shrub with 

erect stems reaching 1 to 4 m in height. Leaves are 

4 to 26 cm long, including petiole, bipinnately 

divided into long linear divisions and may be 

glaborous or pubescent. Margins are entire. 

Densely branched compound cymes terminate the 

main stem and lateral branches. Flowers measure 5 

to 7 cm in diameter. Ray florets are 4 to 5 per 

head, disk florets 6 to 8 per head. Corollas are 

yellow. Fruits are indehiscent, dry, grayish black, 

straight, wingless or very rarely winged, barbless 

and 9 to 12 mm long (Wagner 1990). 

Range.―Ko'oko'olau can be found on Molokai 

and West Maui scattered on arid, leeward slopes 

and cliffs in shrubland vegetation, generally 

between the elevations of 200 to 750 m. On 

leeward sides of Mauna Loa and Mauna Kea of 

Hawaii, this species is common on slopes of cinder 

cones in montane a'ali'i shrublands, and in 

subalpine forest, from elavations of 750 to 2,200 

m. The species is one of 19 Bidens unique to the 

Hawaiian islands (Wagner 1990). 

Ecology.―Ko'oko‘olau is an important dryland 

shrub and forest plant of ecosystems seriously 

under threat by agriculture and development. On 

the island of Hawaii, the species is a minor 

component of the Ohi'a forest. In montane dry 

shrublands of leeward Mauna Kea and Mauna Loa, 

ko'oko'olau becomes structurally codominant with 

a'alii. Due to its tall growth habitat, it appears 

dominant over other species when it is in flower. 

Ko'oko'olau appears intollerant of wildfire and is 

conspicuously absent from burned areas on 

leeward Hawaii. Ko'oko'olau does not resprout 

following fire (Sherry and others. 1999). The plant 

is worth preserving because, like the 

honeycreepers, Loxops spp. and Hemignathus spp., 

ko'oko'olau serves as an example of the adaptive 

radiation that occurred in Hawaii, with all 19 

species of Bidens having a single ancestor. 

Reproduction.―Seed production is fairly prolific 

as the inflorescence is a large terminal panicle 

composed of 8 to 100 disc florets. Fruits are 

achenes. Chromosome number has been identified 

as 2n = 72 (Wagner 1990). Interestingly, the 

species hybridizes readily and for this reason 

individuals unique to a particular local should not 

be introduced to a different zone, in order to 

protect the integrity of endemic species. 

Growth and Management.―Reasonably fast 

growing, ko'oko'olau may not be long-lived. 

Ko'oko'olau can be grown easily from seeds or 

from terminal cuttings using a root hormone. It 

prefers light to moderate watering and full sun 

(Krauss 1998). 

Benefits.―Ko'oko‘olau was widely used by 

Hawaiians prior to European arrival and is still 

sold as tea, although the introduced pantropical B. 

pilosa is usually the species incorrectly labeled 

and sold as the traditional Hawaiian tea. 

References 

Carlquist, S. 1970. Hawaii--a natural history. The 

Natural History Press, Garden City, NY. 484 p. 
111

Krauss, B. 1998. How to plant a native Hawaiian 

garden. http://www.hawaii.gov/health/oeqc/ 

garden/index.html. [not paged]. 

Sherry, K., J.M Castillo, and R.B Shaw. 1999. 

Effects of wildfire on vegetation and rare plants 

in arid montane shrublands, Pohakaloa training 

area, Hawaii. 1999 Hawaiian Conservation 

Conference, Honolulu, HI. 

Wagner, W.L., D.R. Herbst, and S.H. Sohmer. 

1990. Manual of the Flowering Plants of 

Hawaii. University of Hawaii Press, Honolulu. 

1,854 p. 
112 

Bixa orellana L. annatto 

BIXACEAE 

Synonym: Bixa katagensis Delpierre 

John K. Francis 

General Description.—Annatto, which grows as 

a shrub or a small tree, is known by many other 

common names including lipstick plant, roucou, 

achiote, bija, urucú, and shambu (Little and others 

1974). Its single or multiple stems are light brown. 

The bark is more or less smooth with many warty 

lenticels, but may become fissured in old 

individuals. If given ample space, annatto 

generally branches several times near the ground 

and develops a dense, spreading crown. The plant 

roots firmly with a thick taproot and finer laterals. 

The alternate leaves have long petioles, thin ovate 

blades with long-pointed tips. Panicles at the 

branch tips have few to many pink or white 

flowers. The fruits are spiny capsules that dry and 

split open in two parts to expose red seeds on their 

inner surfaces (Howard 1989, Liogier 1995). 

Range.—Although annatto is native to continental 

tropical America, it is unclear where the original 

native stands of the species were located because 

the shrub has been cultivated since ancient times. 

It now grows in cultivation and naturalized from 

Mexico to Argentina and throughout the Caribbean 

Islands. The plant is also widely cultivated and 

naturalized in tropical and subtropical regions 

throughout the rest of the world. 

 

Ecology.—Annatto is shade-intolerant and must 

have disturbance or a broken forest canopy to 

become established. All the naturally growing 

annatto shrubs in Puerto Rico are found on 

neglected or abandoned farmland, some of which 

has grown up to early secondary forest. Rainfall 

ranges from 1000 mm to 3000 mm per year in 

areas where the species grows naturally or under 

cultivation. Soils with textures from sands to clays 

are colonized. The species tolerates relatively low 

base saturation and moderate compaction. Anatto 

is vulnerable to overtopping and smothering by 

trees, shrubs, vines, and grass. Plants that have 

become overtopped and shaded cease to flower 

and bear fruit. The species is frost sensitive (von 

Carlowitz 1991). 

 

Reproduction.—Annatto plants flower and fruit 

heavily and almost continuously in favorable 

habitat. Cuttings taken from flowering plants will 

produce flowers and fruits and a smaller shrub than 

plants of seed origin (Bailey 1941). Belfort and 

others (1992) report that seeds dried to moisture 

contents of 10 to 15 percent germinated at from 8 

to 58 percent compared to fresh seeds (65 percent 

moisture) that gave 96 percent germination. 

Scarification was reported to improve germination 

of fresh seeds in another study. Mechanical 

scarification proved superior to acid or hot water 

treatments (Amaral and others 1995). Air-dry fruits 

collected in Puerto Rico weighed an average of 

1.701 + 0.078 g. Seeds from that collection 

averaged 0.029 + 0.008 g/seed or 35,000 seeds/kg. 

Of these seeds that received no pregermination 

treatment, 60 percent germinated between 11 and 

110 days after sowing. The resulting plants were 

ready to prick out (10 cm in height) about 3 months 

after germination. Success in air layering of shoots 

ranged from 93 to 100 percent except when 

sawdust was used as substrate (7 percent) (Barbosa 

e Silva and others 1993). Stem cuttings rooted (up 

to 60 percent) when treated with IAA or IBA. 

Untreated cuttings did not root (Thirunavoukkarasu 

and Saxena 1997). Annatto can be grafted by 

several techniques. The best method is budding (70 

percent success in tests) (Bruckner and others 

1991). 

Growth and Management.—Annatto shrubs will 

bear fruit when 2 years old in Hawaii (Neal 1965).
113

Under good management, plants will fruit within 1 

year of planting (Nepstad and others 1991). An 

Indian plantation yielded 529 kg/ha of seed at 2 

years old and 2,483 kg/ha of seed at 3 years old 

(Kanjilal and Singh 1995). Annatto seldom 

reaches more than 5 m in height and 10 cm in stem 

diameter (Little and others 1974). Pruning of 

ornamentals is recommended to shape and thicken 

the crowns (Warren 1997). 

 

Benefits.—Annatto, obtained from the oily arils of 

the seeds is the world’s second most important 

(after caramel) natural colorant (Mercadante and 

Pfander 1998), yielding yellow to red colors. The 

colors are produced by several apocarotenoides 

and may reach up to 7 percent of the seed’s dry 

mass (Katzer 1999). World production of annatto 

seed, both for commercial and home use, was 

estimated in 1990 at 10,000 tons per year (Arkcoll 

1990). Brazil is the world’s largest exporter 

(Katzer 1999). Not only was the dye used 

anciently to color food, but also to dye cloth and 

paint the skin (which is still done today). The 

species is also planted as an ornamental, 

particularly the varieties with bright pink pods. 

Branches with the dry pods are used in dry floral 

arrangements (Warren 1997). Bees collect nectar 

from its flowers to make honey. The wood is lightweight (specific gravity 0.4), weak, and not 

durable. It was used in former times to start fires 

by friction. Ropes and twine were made from the 

fibrous bark (Little and others 1974). The pulp 

surrounding the seeds is widely used in herbal 

medicine to treat burns, bleeding, dysentery, 

gonorrhea, constipation, and fever (Parrotta 2001). 

Extracts of leaves, bark, and roots are reported to 

be antidotes for poisoning from Manihot esculenta

Crantz, Jatropha curcas L., and Hura crepitans L. 

(Liogier 1990). 

References

Amaral, L.I.V., M. de F.A. Pereira, and A.L. 

Cortelazzo. 1995. Dormancy breaking in seeds 

of Bixa orellana. Revista Brasiliera de 

Fisiologia Vegetal 7(2): 151-157. 

Arkcoll, D. 1990. New crops from Brazil. In: J. 

Janick and J.E. Simon, eds. Advances in new 

crops. Timber press, Portland, OR. p. 367-371. 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. Vol. 1. The MacMillan Co. New 

York, NY. 1,200 p. 

Barbosa e Silva, K.M., F.C.G. Almeida, F.A.G. 

Almeida, P.S. Lima e Silva, J.J.L. de 

Albuquerque, and J.J.L. de Albuquerque. 1993. 

Effect of substrate on rooting of air-layered 

shoots of annatto. Pesquisa Agropecuaria 

Brasileira 28(1): 101-106. 

Belfort, A.J.L., O.R. Kato, and M. do S.A. Kato. 

1992. Practical method of drying annatto seeds 

for seedling production. Circular Tecnica 67. 

Centro de Pesquisa Agropecuaria do Tropico 

Umido, EMBRAPA-CPATU, Belem, Brazil. 

14 p. 

Bruckner, C.H., S.S. Khouri, and A.V. Melgaco. 

1991. Propagation of annatto (Bixa orellana L.) 

by five grafting methods. Revista Ceres 38: 218, 

340-344. 

Howard, R.A. 1989. Flora of the Lesser Antilles. 

Vol. 6. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 658 p. 

Kanjilal, P.B. and R.S. Singh. 1995. Agronomic 

evaluation of annatto (Bixa orellana L.). Journal 

of Herbs, Spices, and Medicinal Plants 3(3): 13-

17. 

Katzer, G. 1999. Annatto (Bixa orellana L.). 

http://www-ang.kfunigraz.ac.at/~katzer/engl/ 

Bixa_ore.html. 3 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 563 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial 

de la Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Mercadante, A.Z. and H. Pfander. 1998. 

Carotenoids from annatto: a review. Recent 

Research Developments in Agriculture and Food 

Chemistry 2(1): 79-91. 

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Museum 

Press, Honolulu, HI. 924 p. 
114 

Nepstad, D.C., C. Uhl, and E.A.S. Serrao. 1991. 

Recuperation of a degraded Amazonian 

landscape: forest recovery and agricultural 

restoration. Ambio 20(6): 248-255. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CAB International, Wallingford, UK. 

944 p. 

Thirunavoukkarasu, M. and H.O. Saxena. 1997. A 

short note on the effect of auxins (IAA, IBA) 

on rooting of Bixa orellana L. stem cuttings. 

Orissa Journal of Horticulture 25(1): 84-86. 

 

von Carlowitz, P.G. 1991. Multipurpose trees and 

shrubs, sources of seeds and inoculants. 

International Council for Research in 

Agroforestry, Nairobi, Kenya. 328 p. 

Warren, W. 1997. Tropical plants for home and 

garden. Thames and Hudson, New York. 240 p. 
115

Bocconia frutescens L. pan cimarrón 

PAPAVERACEAE 

Synonyms: Bocconia frutescens var. cernua Moc. & Sessé ex DC. 

 Bocconia glauca Salisb. 

 Bocconia pearcei Hutch. 

 Bocconia quercifolia Moench 

 Bocconia sinuatifolia Stokes 

 Bocconia subtomentosa L’Hér. ex Stahl 

John K. Francis 

General Description.—Pan cimarrón is also 

known as parrot weed, celadonia, plume-poppy, 

palo de toro, panilla, llorasangre, curarador, palo 

amarillo, saúco, yagrumito, chocolate blanco, 

gordolobo, guacamayo, trompeto, tabaquillo, 

grande chelidoine, and bois codine. It is a shrub or 

small tree, usually 2 to 3 m in height and 2 to 3 cm 

in stem diameter. Branching near the ground 

normally results in multiple stems. The plant is 

supported by a tap and lateral system of orange-red 

roots that are fleshy outside and fibrous within. 

There are relatively few branches. The outer bark is 

light brown, smooth to lightly fissured; the inner 

bark is orange-red and bitter. An orange or yellow 

sap exudes from wounds. The deeply lobed and 

toothed leaves are alternate but usually clustered at 

the ends of older branches. Petioles are stout, 1.5 to 

3 cm long, and blades are 14 to 50 cm long. 

Panicles up to 50 cm long have lax branches. The 

numerous small flowers are yellow-cream in color. 

The fruits are ellipsoidal capsules each containing a 

black seed with a red, fleshy aril running along one 

side (Howard 1988, Liogier 1985, Little and others 

1974). 

Range.—Pan cimarrón is reported to be native 

from the middle of Mexico through Central 

America to Argentina and Bolivia in South 

America, and throughout the Caribbean (Howard 

1988, Liogier 1985, Stevens and others 2001, New 

York Botanical Garden 2002). It has naturalized in 

forests in the islands of Maui and Hawaii in the 

State of Hawaii (University of Hawaii Botany 

2002). 

Ecology.—Pan cimarrón grows in a wide variety 

of soil types at middle elevations up to 2,150 m in 

Mexico (Secretaría de Madio Ambiente y Recursos 

Naturales 2002). In Puerto Rico, it grows in 

subtropical dry forest along streams (750 to 1000 

mm/year precipitation), in subtropical moist forest 

(1000 to 2000 mm/year precipitation), and in 

subtropical wet forests (2000 to 3000 mm/year 

precipitation). In Nicaragua, the species grows 

even in the cloud forests (Stevens and others 2001). 

In Puerto Rico, pan cimarrón is most common 

along streams, road cuts, and landslides. The 

species also grows widely scattered in brush lands, 

which develop from abandoned fields and pastures, 

and in secondary forest. Pan cimarrón is intolerant 

of shade. It competes vigorously with herbs and 

brush after establishment. The shrub does not form 

continuous stands in Puerto Rico, but occurs as 

scattered individuals or small patches. However, as 

an invader in Hawaii, it does form large, dense 

stands in dry and mesic habitat (University of 

Hawaii Botany 2002). 
116 

Reproduction.—Pan cimarrón flowers and fruits 

throughout the year (Little and others 1974, 

Stevens and others 2001). The infrutescences may 

contain tens to hundreds of fruits. Fruits in the 

infrutescences mature a few at a time beginning at 

the top and proceeding downward. A collection of 

seeds from Puerto Rico weighed (air-dried) an 

average of 0.0146 + 0.0002 g/seed (author’s 

observation). Germination begins in 25 to 55 days 

with 40 percent germinating. Germination is 

epigeal (Ricardi and others 1977). The seeds are 

dispersed by birds (Environmental Protection 

Agency 2002). Damaged plants usually resprout. 

Growth and Management.—Once established, 

pan cimarrón has relatively rapid growth (Jardín 

Botánico de Bogotá José Mutis 2002). Only in 

Hawaii is the species mentioned as a nuisance 

where it displaces native vegetation and threatens 

the endangered Blackburn’s sphinx moth 

(Environmental Protection Agency 2002). In the 

absence of tests of control measures, such standard 

practices as grubbing and spraying with broadleaf 

herbicides are recommended. 

Benefits.—Pan cimarrón contributes to 

biodiversity, helps protect the soil, and furnishes 

food and cover for wildlife. The sap has been used 

as a dye (Little and others 1974). The orange wood 

is soft and brittle with a thick pith, and is of little 

use. Extracts of various tissues of pan cimarrón are 

employed in herbal medicine to control mange, 

lice, and intestinal worms, to treat ulcers of the 

eyes, to treat wounds, and to treat edema and 

jaundice (Pérez-Arbelaez 1978). A dose of 15 

drops of the sap causes a strong purgative effect 

(Guzmán 1975). The sap will cure warts and if 

injected under the skin acts as a local anesthetic 

although the injection itself causes considerable 

pain. The bark contains the alkaloids, 

bocconietrine, cocconichlorine, bocconiyoidine, 

and bocconixatine (Secretaría de Madio Ambiente 

y Recursos Naturales 2002). Pan cimarrón is being 

evaluated as a species for restoration of disturbed 

sites (Jardín Botánico de Bogotá José Mutis 2002). 

References 

Environmental Protection Agency. 2002. Proposed 

endangered status for Blankburn’s sphinx moth. 

Federal Register Vol. 62, No. 63. http:// 

www.epa.gov/fedrgstr/EPA-SPECIES/1997/ 

April/Day-02/e8550/htm. 12 p. 

Guzmán, D.J. 1975. Especies útiles de la flora 

salvadoreña. Ministerio de Educación, Dirección 

de Publicaciones. San Salvador, El Salvador. 

703 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Jardín Botánico de Bogotá José Mutis. 2002. 

Conservación ex situ. http://www.jbb.gov.co/ 

conservacion/ex_situ/semillas/body_semillas. 

html. 4 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 

New York Botanical Garden. 2002. Specimens 

search results: Bocconia frutescens. 

http://scisun.nybg.org.8890/searchdb/owa/www

catalog.search_list. [not paged]. 

Pérez-Arbelaez, E. 1978. Plantas útiles de 

Colombia. Litografía Arco, Bogota, Colombia. 

831 p. 

Ricardi, M., F. Torres, C. Hernandez, and R. 

Quintero. 1977. Morphology of seedlings of 

Venezuelan trees. Revista Forestal Venezuelana 

27: 15-56. 

Secretaría de Madio Ambiente y Recursos 

Naturales. 2002. Especies con usos no 

maderables en bosques tropicales y 

subtropicales: Bocconia frutescens L. 

http://www.semarnat.gob.mx/pfnm2/fichas/bocc

onia_frutescens.htm. 2 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany, Vol. 85, No. 

3. Missouri Botanical Garden, St. Louis, MO. 

p. 1,911-2,666. 

University of Hawaii Botany. 2002. Bocconia 

frutescens, bocconia, Papaveraceae. 

http://www.botany. hawaii.edu/faculty/ 

cw_smith/boc_fru.htm. 1p. 
117

Bontia daphnoides L. white alling 

MYOPORACEAE 

Synonyms: Bontia daphnifolia Salisb. 

 Bontia daphnoides var. menor (Gaertn. f.) A. DC. 

 Bontia menor Gaertn. f. 

John K. Francis 

General Description.—White alling, also known 

as wild olive, mangle bobo, aceituna Americana, 

olivier bard, and oliba, is a shrub or small tree 

usually 3 or 4 m in height, occasionally reaching 9 

m and 15 cm in stem diameter. Although seedlings 

have a tap and lateral root system, older plants rely 

on shallow lateral roots that are stiff and brittle 

with a brown, corky bark. Trunks of white alling 

are usually covered with a thick, light brown, 

furrowed bark. The heartwood is light gray-brown, 

hard, heavy, and somewhat brittle; the sapwood is 

tan in color. Multiple stems commonly diverge 

from the base of the principal stem. The crowns are 

vertical and narrow with a thin complement of 

foliage. Leaves are oblong to linear-lanceolate, 3 to 

11 cm long by 1 to 2 cm broad, and pointed at both 

ends. The usually solitary, axillary tubular flowers 

are yellow blotched with purple and 2 cm long. 

Drupes are ovoid, tapering to a point with a 

permanently attached style. Their corky-textured 

flesh is yellowish-green to yellow at maturity, has a 

slightly bitter taste, and contains a single hardshelled stone (Howard 1989, Liogier 1997, Little 

and others 1974). 

Range.—White alling is native to the Bahamas, the 

Greater and Lesser Antilles, Trinidad, Venezuela, 

and Guyana (Howard 1989, Little and others 

1974). It is cultivated as an ornamental in Hawaii 

(Hawaiian Ecosystems at Risk 2001) and Spain 

(Sánchez 2001) and has naturalized in Florida 

(Nelson 1996). 

Ecology.—White alling is most common in coastal 

thickets between the tidal mangroves and the 

upland forests. These areas are flooded during 

storm surges and receive a moderate amount of salt 

spray. The soils vary from sandy to clayey, are 

usually high in organic matter, and range in pH 

from 7.0 to 8.5. The water table is usually within 2 

m of the surface. In Puerto Rico, the species grows 

in areas that receive from 750 to 1800 mm of mean 

annual precipitation. White alling plants are 

moderately intolerant of shade, generally starting in 

openings. Seedlings and saplings survive in 

relatively sunny understories. Adults flower and 

fruit in intermediate crown positions. The species 

is rare in uplands, probably because of competition. 

It has been cultivated successfully at elevations up 

to 1,500 m (Little and others 1974). White alling 

grows as rare to common components of stands but 

does not form pure or nearly pure stands. 

Reproduction.—White alling flowers and fruits 

throughout the year (Little and others 1974). Fruits 

collected in Puerto Rico averaged 0.311 + 0.015 

g/fruit. Air-dried seeds cleaned from the above 

collection averaged 0.0898 + 0.0033 g/seed or 

11,000 seeds/kg. Sown in commercial potting mix, 

44 percent of the seeds germinated between 12 and 

68 days after sowing (author’s observation). Seeds 

are dispersed by water and presumably by birds 

and mammals that eat the fruits. Fruit production is 

usually good and seedlings are common in small 

openings near seed sources. Although white alling 

are often tipped in hurricanes, they sprout from the 

trunks and reform vertical crowns. 
118 

Growth and Management.—A small group of 

white alling seedlings grown in the nursery in 

Puerto Rico ranged in height from 21 to 86 cm, 8 

months after being pricked into containers. Sapling 

and sprout growth rate is moderate (about 0.5 

m/year) and individual plants live 10 to 30 years. 

Although no management experience has been 

published, the species can probably be established 

with nursery seedlings and be managed like other 

short-statured species. 

Benefits.—White alling helps protect the soil and 

provides food and cover for wildlife. The wood is 

useful for fuel and stakes. Ethanol extracts of white 

alling showed promising insecticidal activity 

against Boophilus microplus, Cylas formicarius, 

and Tribolium confusum (Mansingh and Williams 

1998, Williams and Caleb-Williams 1997, 

Williams and Mansingh 1993). White alling is 

frequently used to control diabetes in Trinidad 

(Mahabir and Gullifor 1997). Extracts of the plant 

are used to control intestinal worms, treat herpes, 

treat inflammation, insect bites, scarring, ulcers, 

and wounds (Liogier 1990). 

References 

Hawaiian Ecosystems at Risk 2001. Mystery plant 

# 2, Bontia daphnoides Linn. (Myoporaceae). 

http://hear.org/mysteryplants/autogendhtml/ 

mysteryplant2.htm. 3 p. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Mahabir, D. and M.C. Gullifor. 1997. Use of 

medicinal plants for diabetes in Trinidad and 

Tobago. Revista Panamericana de Salud Pública 

1(3): 174-179. 

Mansingh, A. and L.A.D. Williams. 1998. 

Pesticidal potential of tropical plants II. 

Acaricidal activity of crude extracts of several 

Jamaican plants. Insect Science and its 

Application. 18(2): 149-155. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Sánchez deL.C., J. 2001. Plantas de la flora cubana 

cultivadas en España. http://floraguide.es/ 

arboles/plantasdecuba.htm. 8 p. 

Williams, L.A.D. and L. Caleb-Williams. 1997. 

Insecticidally active sesquiterpene furan from 

Bontia daphnoides L. Philippine Journal of 

Science 126(2): 155-162. 

Williams, L.A.D. and A. Mansingh. 1993. 

Pesticidal potential of tropical plants I. 

Insecticidal activity in leaf extracts of sixty 

plants. Insect Science and its Application. 14(5): 

697-700.
119

Borrichia arborescens (L.) DC. sea ox-eye 

ASTERACEAE 

Synonyms: Buphthalmum arborescens L. 

 Borrichia argentea DC. 

 Borrichia glabrata Small. 

John K. Francis 

General Description.—Sea ox-eye, also known as 

tree oxeye, silver sea ox-eye, oxeye daisy, sea 

daisy, seaside tansy, gull feed, clavelón de playa, 

and fleur-sorleil bord de mar, is an evergreen, low 

shrub usually 1 m or less in height with stem 

diameters of 1 cm, which forms mounds and mats. 

Older plants have numerous stems. The plants are 

supported by short rhizomes and a moderate 

number of lateral roots. The stems are gray with a 

white, brittle wood and a 3-mm pith. Twigs bear 

deep leaf scars. The foliage tends to be crowded at 

ends of twigs. The simple, opposite, sessile leaves 

are fleshy and yellow-green, light-green, or graygreen, oblanceolate to spatulate, entire, and 3 to 8 

cm long. The foliage is resinous and aromatic. Sea 

ox-eye flowers are usually solitary, terminal heads 

about 2.5 cm across on peduncles 2 to 5 cm long. 

The corolla and the ray florets are bright yellow 

and the disk florets are orange-yellow. The black 

achenes are 3 to 4 mm long with a pappus in the 

form of a dentate cup less than 1 mm long. The 

chromosome number is 2n = 28 (Howard 1989, 

Liogier 1997, Long and Lakela 1976, Nelson 1996) 

Range.—Sea ox-eye is native to southern Florida, 

the Bahamas, Bermuda, the West Indies, Mexico 

and Guatemala (Liogier 1997). Although it is 

planted as an ornamental, it is not known to have 

naturalized outside the native range. Borrichia x 

cubana of southern Florida is believed to be a 

natural hybrid of sea ox-eye and Borrichia 

frutescens (L.) DC. (Nelson 1996). 

Ecology.—Natural sea ox-eye grows near the 

seashore, usually within the influence of salt spray. 

Because the species is low in stature, grows slowly, 

and is intolerant of shade, it must occupy areas 

with low competition from other vegetation. It 

finds these conditions on headlands, seaside rocks, 

dunes, beach strands, low hammocks, and the 

edges of mangroves and brackish marshes. Sea oxeye will grow in more favorable conditions if 

artificially protected from competition. Almost any 

soil will do, including both acid and alkaline soils. 

Sea ox-eye tolerates both excessively drained and 

poorly drained conditions. Mean annual rainfall 

may range from 900 to 1500 mm. It is drought 

resistant (Gillman 1999). 

Reproduction.—Sea ox-eye blooms in spring and 

summer in Florida (Long and Lakela 1976) and 

throughout the year in Puerto Rico (author’s 

observation). It is pollinated by insects. Seeds 

collected in Puerto Rico averaged 0.00115 g/seed 

or 870,000 seeds/kg. Only 16 percent of this 

collection germinated when placed in moist potting 

mix. Workman (1980) recommends propagating 

sea ox-eye by sowing in pots. Also, the plant can 

be propagated by cuttings (Gillman 1999). Seed 

dispersal appears to be by water (storm surges) and 

strong winds. Once established, plants widen by 

constant layering of the semiprostrate stems. New 

clumps probably are established by the rooting of 

broken stems moved by storms. 

Growth and Management.—Growth of sea oxeye is slow. Branch extension is about 10 to 30 

cm/year. Although, stems and branches die, as long 

as conditions remain favorable, clonal plants may 

live almost indefinitely. Newly established plants 

should be protected from overtopping by grass, 

brush, and trees. It is important not to regularly 
120 

irrigate sea ox-eye plants used in landscaping 

(Gillman 1999). 

Benefits.—Sea ox-eye is available commercially 

and is widely planted as an ornamental. It is 

recommended for foundation plantings and as a 

border shrub (Workman 1980), particularly where 

salt and salt spray are prevalent and in xeric 

(unwatered) gardens (Knox 2002). It also attracts 

butterflies (Gilman 2002). A tea is prepared from 

branch tips and leaves to relieve colds, coughs, and 

fish poisoning (Garland 2002). 

References 

Garland, K. 2002. Borrichia arborescens (Sea Oxeye). University of the Virgin Islands. 

http://rps.uvi.edu/VIMAS/seaoxeye.htm. 1 p. 

Gillman, E.F. 1999. Borrichia arborescens. Fact 

Sheet FPS-68. Cooperative Extension Service, 

University of Florida, Gainesville, FL. 3 p. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Knox, G.W. 2002. Drought tolerant plants for 

North and Central Florida. Pinellas County 

Extension. http://coop.co.pinellas.fl.us/fyn/ 

publications/fyn_drought_tolerant.html. 12 p. 

 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Long, R.W., and O. Lakela. 1976. A flora of 

tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Workman, R.W. 1980. Growing native. The 

Sanibel-Captiva Conservation Foundation, Inc., 

Sanibel, FL. 137 p. 
121

Bourreria virgata (Sw.) G. Don roble de guayo 

BORAGINACEAE 

Synonyms: Ehretia virgata Sw. 

John K. Francis 

General Description.—Roble de guayo, also 

known as palo de vaca, hoja menuda, guazumilla, 

raspalengua, and cafecillo, is an evergreen shrub or 

small tree sometimes reaching 9 m in height and 

15 cm in stem diameter at breast height. It is 

somewhat branchy and may have single or 

multiple stems emerging from the ground. The 

stem bark is gray and smoothish to slightly 

fissured. The wood is light brown, hard, heavy, 

and relatively brittle. Roble de guayo is deeply and 

solidly rooted with a tap and lateral root system of 

brown, somewhat stiff roots. Twigs are slender, 

hairless to densely hairy, green when young, 

becoming gray. The coriaceous, shiny, green 

leaves are alternate but may be crowded near twig 

ends. Blades are elliptic, oblong, or obovate, 1.5 to 

7 cm long by 0.7 to 2.5 cm broad, rounded to 

slightly notched at the apex, and have a short 

petiole. The terminal inflorescence is a corymb or 

a cyme with two to 15 tubular, white flowers with 

five corolla lobes. Fruits are orange or red-orange, 

subglobose, 5 to 8 mm in diameter, often threelobed, drupes with persistant calyx and style. Each 

fruit contains four or fewer light brown, hard 

nutlets. The nutlets are a quarter sphere with the 

outer surface deeply grooved (author’s 

observations, Liogier 1995, Little and others 

1974). 

Range.—Roble de guayo is native to Cuba, 

Hispaniola, and Puerto Rico, including the Island 

of Vieques (Little and others 1974). It is not 

known to have been planted or naturalized 

elsewhere. 

Ecology.—Roble de guayo grows in areas that 

receive about 700 to 2000 mm of mean annual 

precipitation. Elevations of habitat in Puerto Rico 

range from 30 to 300 m (Little and others 1974). It 

colonizes well- to excessively well-drained soils 

with a wide range of textures and pH’s from about 

5.5 to 8.0. Most of the hillside, ridge, and hilltop 

soils are shallow and rocky. In Puerto Rico, roble 

de guayo is locally common in the moist and dry 

limestone hills and in dry foot hills of igneous and 

metamorphic (including ultramafic) rocks 

(Breckon and García 2001, Little and others 1974). 

The species is an important component of the 

cactus scrub vegetation type in Cuba (World 

Wildlife Fund 2001). It is moderately intolerant of 

shade, growing in the understory of low basal area 

forests but needs overhead sun to produce good 

fruit and seed crops. The species survives best in 

harsh environments where competition is minimal. 

Reproduction.—Roble de guayo flowers and 

fruits throughout the year (Little and others 1974). 

Production of fruits and seeds is moderate. Two 

collections of fresh fruits from Puerto Rico 

weighed an average of 0.453 + 0.093 g/fruit 

(Francis and Rodríguez 1993) and 0.423 + 0.012 

g/fruit. Air-dried seeds separated from the 

respective groups of fruits averaged 43,500 

seeds/kg and 51,500 seeds/kg. Sown without 

pretreatment, the first sample of seed failed to 

germinate and the second gave only 2 percent 

germination (author’s observation). Germination is 

epigeal. The seeds are undoubtedly dispersed by 

birds that eat the fruits of this as well as other 

species of the genus. Seedlings are relatively 

common. 

Growth and Management.—Both seedlings and 

adult plants grow slowly. Apparently, roble de 
122 

guayo lives for several decades. No management 

experience has been published. The species is not 

weedy in agricultural land or forest plantations and 

will rarely need control. 

Benefits.—Roble de guayo helps protect the soil 

and furnishes food and cover for wildlife. The 

stems are too small for use as lumber but should 

make excellent fuel. The species would probably 

be a good ornamental for landscaping xeric 

gardens. 

References

Breckon, G.J. and R.G. García. 2001. Vascular 

plants of Susúa Forest. University of Puerto 

Rico, Mayaguez, PR. http://www.uprm.edu/ 

biology/profs/breckon/herbarium/florasusua. 

htm. 52 p. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de 

la Universidad de Puerto Rico, San Juan, PR. 

617 p. 

 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

World Wildlife Fund. 2001. Cuban cactus scrub 

(NT1306). http://www.worldwildlife.org/ 

wildworld/profiles/terrestrial/nt/nt1306_full. 

html. 8p. 
123

Brickellia californica (Torr. & Gray) Gray California brickellbush 

ASTERACEAE 

Synonyms: Bulbostylis californica Torr. & Gray 

 Brickellia wrightii A. Gray 

 Coleosanthus californicus Kuntze 

 Coleosanthus albicaulis Rydb. 

 Brickellia tenera A. Gray 

 

John K. Francis 

General Description.—California brickellbush is 

a suffruticose shrub 0.5 to 1.2 m in height. Stem 

bases reach a maximum diameter of about 2 cm, 

and clumps may reach a diameter of 1 m. Plants 

usually have multiple stems arising from the root 

crown or short, near-surface horizontal stems. The 

branches are slender, upright, with even finer 

twigs. Current-year branches, which annually die 

back to a few cm from the ground-line, have a 

white to silver-gray bark, and older branches are 

fine-furrowed gray or gray-brown. The wood is 

yellowish white, medium density, and brittle with 

annual rings. California brickellbush shrubs are 

supported by tap and lateral roots that are covered 

by corky brown, finely-furrowed bark. The 

alternate aromatic, gray-green leaves are deltoidovate with a rounded or cordate base and a shortacute tip and a crenate-dentate margin. They are 1 

to 5 cm long and almost as wide with petioles 4 to 

20 mm long. Inflorescences are heads in terminal 

clusters on small lateral branches of leafy panicles. 

The fragrant heads are 7 to 10 mm long and 

contain eight to 18 greenish-cream florets. The 

olive green achenes are 3 mm long tipped with a 

white to brownish pappus. There is a base number 

of n = 9 chromosomes (author’s observation, 

Abrams and Ferris 1960, Harrington 1964, Munz 

1974). 

Range.—California brickellbush occurs in two 

varieties. Variety californica grows in Oregon, 

Idaho, California (including Santa Catalina, 

Anacapa, and Santa Cruz Islands), Nevada, Utah, 

Colorado, Arizona, New Mexico, Oklahoma, 

Texas, and at least Chihuahua State in Mexico. 

Variety jepsonii B.L. Robins is found only in 

California (Abrams and Ferris 1960, Munz 1974, 

Natural Resources Conservation Service 2003). 

Although the species is planted as an ornamental, 

it has not been reported to have naturalized outside 

its native range. 

Ecology.—California brickellbush is intolerant of 

shade and does not grow under tree or closed 

shrub canopies. Stands observed by the author 

were along trails, intermittent streams, and steep, 

north-facing colluvial slopes. Abrams and Ferris 

(1960) say that California brickellbush grows in 

dry canyons and streambeds. It grows in soils 

derived from a wide range of parent materials, 

through the whole range of textures, and with pH’s 

of 5 to 8. California brickellbush grows in habitat 

that receives annual rainfall amounts ranging from 

220 to 640 cm (Las Pilitas Nursery 2003). The 

species grows at between 915 and 2,290 m in 

elevation in Arizona (Dittman 2003). It grows in 
124 

coastal and island vegetation types in California 

(Munz 1974) and therefore at low elevations. 

Disturbance is probably required for 

establishment. 

Reproduction.—California brickellbush flowers 

August through October (Tarweed Native Plants 

2003). The species flowers abundantly and 

potentially produces vast numbers of seeds. Airdried seeds collected in Utah averaged 3.525 

million/kg. Hand-sorted seeds placed without any 

pretreatment on moist blotter paper germinated at 

71 percent beginning between 5 and 14 days after 

sowing. Germination is epigeal (author’s 

observation). Seed will have to be harvested by 

hand because wild plants are scattered. Stems layer 

(root) when covered by soil or colluvial material. 

The seeds are wind-dispersed. Seedlings probably 

appear and survive only when conditions are 

nearly ideal. 

Growth and Management.—Save for a few 

centimeters of caudex, California brickellbush 

adds and loses its entire height (0.5 to 1.0 m) each 

growing season. The aerial portion of the caudex 

also dies back about every 3 to 5 years while 

others are produced each year. Plant may live for 

one to several decades, renewing themselves with 

suckers. The species is rarely abundant and 

probably does not require control. Management 

experience has not been published. 

Benefits.—California brickellbush helps protect 

the soil and furnishes food and cover for wildlife. 

It has been reported to be a larval food plant for 

the tiger moth, Dysschema howardi Hy. Edwards 

(Walsh 2003). The species has been planted as an 

ornamental, particularly for the floral fragrance, in 

natural landscaping (Oklahoma Biological Survey 

2003). Native Americans used infusions of the 

leaves as a ceremonial emetic, as a lotion to treat 

infant skin sores, and for coughs and fevers 

(Moerman 1986). 

References

Abrams, L., and R.S. Ferris. 1960. Illustrated flora 

of the Pacific States. Stanford University Press, 

Stanford, CA. 732 p. 

Dittman, L. 2003. Asteraceae: aster, daisy or 

composite family. http://www.nazflora.org/ 

asteraceae.htm. [not paged]. 

Harrington, H.D. 1964. Manual of the plants of 

Colorado. Sage Books, Denver, CO. 666 p. 

Las Pilitas Nursery. 2003. Brickellia californica. 

http://www.laspilitas.com/plants/123.htm. 3 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports 19. University of 

Michigan Museum of Anthropology, Ann 

Arbor, MI. 534 p. 

Munz, P.A. 1974. A flora of Southern California. 

University of California Press, Berkeley, CA. 

1,086 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Brickellia Ell., brickellbush. 

http://plants.usda.gov/cgi_bin/plant_profile.cgi?

symbol=BRICK. [not paged]. 

Oklahoma Biological Survey. 2003. Brikellia 

californica (Torr. & Gray) Gray. 

http://www.biosurvey.ou.edu/shrub/brca3.htm. 

3 p. 

Tarweed Native Plants. 2003. Current plant 

availability list. http://www.tarweednativeplants. 

com/botanical.html. 5 p. 

Walsh, B. 2003. Moths of southeastern Arizona: 

Arctiidae (tiger moths). http://nitro.biosci. 

Arizona.du/zeeb/butterflies/artic.html. 8 p. 
125

Bromelia pinguin L. maya 

BROMELIACEAE 

Synonyms: none 

John K. Francis 

General Description.—Maya is a pineapple-like 

plant with large, sword-shaped, dark-green leaves 

that have alternate curved spines about 5 mm long 

on their edges. The name used is Spanish 

meaning “net” which derives from its use as a 

barrier or possibly because its fibers may have 

been used for making nets. Other common names 

are karatas, pingouin, bayonette, and pinguin 

(Howard 1979). Although it is mostly fleshy, 

maya is classed as a shrub because it is perennial, 

of shrub size, and has a woody core at its base 

and fibrous leaves. The plants are formed of a 

large basal rosette and rarely develop a 

discernible stem. The roots are shallow, relatively 

fine, all of a similar size, and radiate in all 

directions. At the start of a plant’s last year, it 

grows a stout scape inflorescence with many 

wooly red-orange flowers. The new leaves 

surrounding the inflorescence are also intensely 

red-orange. A few months later, 3.5-cm-long 

elliptical yellow berries ripen. After the fruits 

have withered, about 1 year after the start of 

fruiting, the plant dies. 

Range.—The native range of maya extends from 

Mexico through tropical South America and the 

Caribbean islands (Howard 1979). In Puerto Rico 

nearly all the stands are on or near abandoned 

farmland, which may indicate a relatively recent 

introduction. It has been planted and naturalized 

in Hawaii (Neal 1965) and many other tropical 

areas. 

Ecology.—Maya is intermediate in shade 

tolerance. Although it sometimes grows in open 

areas, the most vigorous stands are found under 

forest stands with moderate basal areas. In Puerto 

Rico, natural stands occur in areas with from 850 

to 2000 mm of rainfall and from near sea level to 

600 m of elevation. All types of soils except very 

poorly drained and saline soils are colonized. 

Maya is sensitive to fire. Although many plants in 

a colony will recover from a burn, they do so 

slowly. Usually flowering is synchronized, 

although an occasional plant flowers out of phase, 

especially in moist habitat. 

Reproduction.—Maya reproduces vegetatively 

and by seeds. The fruits, whose fresh weight 

averaged 12.26 + 0.35 g in a Puerto Rican 

collection, contain 0 to over 100 seeds, depending 

on their size. The fruits have a tough, fibrous rind. 

In Puerto Rico, most of the fruits are eaten by 

fine-toothed animals (probably rats, mice, or fruit 

bats) and it is assumed that the seeds are dispersed 

by these small mammals. The black, teardropshaped seeds averaged 0.245 + 0.006 g (air dry). 

Seventy-five percent of these seeds germinated 

between 133 and 175 days after sowing. The 

plants are very fragilely rooted at first and develop 

at a moderate rate. After maya plants have reached 

their full size and before flowering, most healthy 

individuals produce one or sometimes two stiff 

horizontal stolons about 0.5 m long. A new plant 

forms at the terminus. The new plants grow 

rapidly and reach roughly half the parent plant’s 

height and diameter and become independent in 

about a year. Consequently, (with the exception of 

those planted by humans) new colonies are started 

as seedlings from dispersed seeds and most plants 

within colonies arise vegetatively. Maya has been 

propagated with apical bud explants with a high 

degree of success (Mesa and Lajonchere 1996) 

Growth and Management.—Maya plants grow 

to full size (1 to 2 m in height and 2 to 3 m in 

diameter) in 2 or 3 years. Mature stands can be 

dense with interlacing crowns and little clear 

space. Five stands inventoried in Puerto Rico 

ranged from 723 to 1,948 plants/ha. Four whole 
126 

plants averaged 81.6 percent water and ranged 

from 0.3 kg to 1.6 kg dry weight. The maya 

portion of total stand dry biomass would run from 

about 500 to 2,000 kg/ha. For the most part, maya 

is undesirable in forests and pastures because it 

takes up valuable space, restricts access, inhibits 

grazing, and provides nesting habitat and cover for 

rats (Vélez and van Overbeek 1950). Maya stands 

may be controlled by uprooting and piling the 

plants. If management objectives so dictate, maya 

can be established by transplanting top-trimmed 

wildlings. 

Benefits.—Historically, and to some extent 

today, maya was used as a hedge or living fence 

to inhibit entry into fields and homesteads (Crane 

1945). These hedges were never more than 

marginally effective—cattle and many species of 

animals pass relatively easily and people can 

ignore the spines or cut a path with a machete. 

The fruit, which has a white flesh, is bitingly acid, 

and tastes like pineapple, is eaten occasionally out 

of hand and used to make a refreshing drink (Neal 

1965). The long, parallel leaf fibers were found to 

be of good quality and proposed for commercial 

production (Guzman 1975). Maya offers food and 

cover for wildlife, such as small mammals and 

birds. 

References 

Crane, J.C. 1945. Living fence posts in Cuba. 

Agriculture in America 5(2): 34-35, 38. 

Guzman, D.J. 1975. Especies útiles de la flora 

Salvadoreña. Ministerio de Educación, Dirección 

de Publicaciones. San Salvador, El Salvador. 

703 p. 

Howard, R.A. 1979. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 3. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 586 p. 

Mesa, A.R. and G. Lajonchere. 1996. 

Micropropagation of Bromelia pinguin Lindl. 

Pastos y Forrajes 19(3): 217-223. 

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Museum 

Press, Honolulu, HI. 924 p. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitario, Río Piedras, PR. 497 p. 
127

Brunfelsia lactea Krug & Urban jasmín del monte 

SOLANACEAE 

Synonyms: none 

John K. Francis 

General Description.—Jasmín del monte (a 

Puerto Rican name), also known as aguacero, vega 

blanca, dama de noche, is an evergreen shrub or 

occasionally a small tree. It has been recorded to 7 

m in height and 10 cm in stem diameter but 

usually measures only 1 to 3 m in height. Older 

plants tend to have multiple stems and many, often 

intertwining branches. The bark is light brown, 

slightly furrowed, and usually covered with 

mosses and liverworts. The inner bark is white. 

The wood is light brown, hard, and brittle. Plants 

are supported by a tap and lateral root system of 

light brown, somewhat brittle roots. Roots 

sometimes arise from the lower trunk. The leathery 

foliage is moderately abundant. Dark green leaves 

are elliptic, oval, or obovate, 5 to 15 cm long and 

2.5 to 6.5 cm broad, and have petioles 5 to 15 mm 

long. The flowers are solitary at the terminus of 

twigs. They are strikingly fragrant, particularly at 

night. White or pale yellow, later tinged with 

purple, the corolla is trumpet shaped, 5 to 7 cm 

long and 5 to 6 cm across, with five rounded lobes. 

Purple to light brown, fleshy fruits (berries) are 

globose, 2 to 2.5 cm in diameter and contain many 

4-mm brown, elliptic seeds (Liogier 1995, Little 

and others 1974). 

Range.—Jasmín del monte is endemic to highelevation forests in eastern Puerto Rico (Liogier 

1995, Little and others 1974) and is confined to 

only a few locations. The species has been grown 

widely in greenhouses and gardens, but there is no 

record of it naturalizing. 

Ecology.—Dwarf forests at the summits of 

mountains are the habitat of jasmín del monte. 

These areas lie between 980 and 1,100 m in 

elevation (Little and others 1974) and receive more 

than 3000 mm of mean annual precipitation. The 

soils are clayey over igneous rocks with pH’s 

between 5 and 6 and are saturated in their subsoils 

most of the time. As an ornamental, jasmín del 

monte blooms in partial shade (Gehrke 2000). In 

natural habitat, the species can survive under forest 

canopies but requires broken or full sunlight to 

flower and fruit. Hurricane-force winds impact the 

mountain tops where the species grows every one 

to few decades, uprooting trees and defoliating the 

forest. This disturbance, plus occasional 

landslides, probably allows adult plants of jasmín 

del monte to flower and fruit and seedlings to 

establish themselves. In recent years, the 

disturbance of road cuts and other construction 

have created additional habitat for the species. The 

minimum overwintering temperature for jasmín 

del monte as an ornamental is 10 °C 

(Kuipplantenkwekerij Gommer 2003). 

Reproduction.—Jasmín del monte flowers 

throughout the year (Little and others 1974). There 

are good fruit and seed production. A collection of 

fresh fruits weighed from 2 to 8 g each. Air-dried 

seed separated from those fruits averaged 0.0062 +

0.0002 g/seed. Sown without pretreatment in 

commercial potting mix, 13 percent germinated 

between 1 and 3 months after sowing. Germination 

is epigeal. Seedlings are relatively common in 

disturbed patches near seed-bearing shrubs but rare 

elsewhere. Stems layer (root) whenever they come 

in contact with the ground. Cut roots and stems 

produce sprouts. 
128 

Growth and Management.—Jasmín del monte 

grows slowly in its natural habitat and appears to 

be long-lived. The author knows of no wildland 

plantings or intentional management. Because the 

species reproduces after disturbance, soil 

unavoidably exposed during construction and road 

work should be left in good tilled condition to 

encourage seedling establishment. 

Benefits.—Jasmín del monte helps hold the soil, 

provides cover and probably food for wildlife, and 

adds to the aesthetics of the forest. The species is 

planted as an ornamental in gardens and 

greenhouses and is available for purchase from a 

number of commercial nurseries. 

References 

Gehrke, P. 2000. Sale plants 2000. Tropical 

Flowering Tree Society, Miami, FL. 

http://www.tfts.org/Plant_Sale_May_2000.htm. 

12 p. 

Kuipplantenkwekerij Gommer. 2003. Brunfelsia 

lactea. Kuipplantenkwekerij Gommer, Dalen, 

Netherlands. http://www.kwekerijgommer.com/ 

fotos/kuipplanten/Brunfelsia%20Lactea.htm. 

1 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial 

de la Universidad de Puerto Rico, Río Piedras, 

PR. 617 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 
129

Caesalpinia bonduc (L.) Roxb. gray nicker bean 

FABACEAE 

Synonyms: Guilandina bonduc L. 

 Guilandina bonducella L. 

 Caesalpinia bonducella (L.) Fleming 

 Caesalpinia crista of Small and other authors 

 Guilandina crista (L.) Small 

John K. Francis 

General Description.—Gray nicker bean, also 

known as nicker nut, wait-a-bit, hold back, fever 

nut, hava de San Antonio, mato de playa, z’yeux à 

chatte, canique grise, and uri, is a spiny, 

scrambling shrub to 1.5 m in height (unsupported) 

and 6 m or more in extension, and has stems up to 

5 cm in diameter or more. Plants usually have a 

single stem arising from the ground but often 

branch low on the stem. Seedlings form taproots 

and may retain them later; lateral roots are 

extensive. The stems grow stiffly upright for 0.75 

to 1 m before curving into a more horizontal 

direction. The stems, twigs, and leaf rachises are 

covered with straight or curved prickles. The 25- to 

80-cm-long leaves are bipinnately compound with 

four to nine pairs of pinnae, each with four to eight 

pairs of oblong to elliptic leaflets. Racemes of 

yellow flowers are lateral or terminal. Prickley, 

inflated legumes are flattened oval shaped, 5 to 10 

cm long, and reddish-brown when dry. Within each 

pod are one to three (usually two) smooth, hard, 2-

cm long seeds that are olive drab in the pods and 

remain so until exposure to the sun bleaches them 

to a light gray color. The pods open partially upon 

drying and eventually release the seeds (author’s 

observation, Howard 1988, Liogier 1988, Nelson 

1996). There are 2n = 24 chromosomes (Long and 

Lakela 1976). 

Range.—Gray nicker bean is established in nearly 

every tropical shore worldwide, including Florida, 

Louisiana, Puerto Rico, the U.S. Virgin Islands, 

Hawaii, Guam, and American Samoa (Howard 

1988, Liogier 1988, Pacific Island Ecosystems at 

Risk 2002, Vermillion 2001). It has apparently 

arrived at these locations by natural means, 

although unrecorded accidental or intentional 

introductions cannot be ruled out. 

Ecology—Gray nicker bean is intolerant of shade 

and is usually open-grown or at least grows in 

broken sunlight. It tolerates salt spray, salty soils, 

and occasional flooding with seawater. The species 

grows in all textures of mildly acid to alkaline soil. 

Annual rainfall in the areas where gray nicker bean 

grows in Puerto Rico ranges from 750 mm to 1800 

mm. The species grows from sea level to 850 m in 

elevation in India (Parrotta 2001). It grows most 

frequently on the beach strand, on coastal dunes, 

and at the better-drained edge of mangroves. Gray 

nicker bean also grows inland in disturbed areas. It 

competes well with grass and herbs and may 

ascend into the crowns of low trees. 

Reproduction.—Gray nicker bean flowers and 

fruits year-round in Florida (Nelson 1996). A 

collection of seeds from Puerto Rico weighed an 

average of 2.444 + 0.034 g/seed or 409 seeds/kg. 

Twenty-five seeds each of weathered-scarified (by 

nicking), weathered-unscarified, fresh-scarified, 

and fresh-unscarified treatments were incubated in 

plastic bags of moist potting mix. Most of the 

scarified seed germinated within a few days and 

had all germinated within 1 year. By 16 months, 23 

of the fresh, unscarified seeds had germinated and 

just one of the weathered, unscarified seeds had 

germinated. Germination is hypogeal (author’s 

observation). Gray nicker bean seeds are 

apparently carried to sea by floods and storm 

surges where they float until deposited on shore. 

The seeds are found on beaches in northern 

Scotland and are known to be able to float in sea 
130 

water for as long as 19 years (Markland 2002). The 

scarifying action of sand, weathering, insects, or 

rodents eventually allows water to enter the seeds 

and they germinate and establish themselves in the 

new environments. A seed collected at Macquarie 

Island, Antarctica, a drift of not less than 1 year, 

was scarified and grown successfully (Costin 

1965). The stems layer (root) whenever they come 

in contact with the ground. Stems sprout when cut 

and lateral roots sometimes sucker. Clonal plants 

tend to remain attached to one another. 

Growth and Management.—Growth of gray 

nicker beans is fairly rapid at all stages. Seedlings 

reached 26 cm in 40 days after sowing. Older 

plants grow a meter or more per year. Individual 

stems live at least 4 years. The author knows of no 

published management experience. Plantations 

undoubtedly could be established with potted 

seedlings or by direct seeding scarified seeds. 

Control could probably be obtained by lopping 

with a machete followed by herbicide treatment of 

the sprouts. 

Benefits.—Gray nicker bean contributes to the 

biodiversity of the forests where it grows, helps 

protect the soil, and furnishes cover for wildlife. 

The seeds have been used for centuries and are still 

used as jewelry, prayer beads, good luck charms, 

and worry stones (Rancho Leona 2002, Workman 

1980). They were anciently used as standards of 

weight in India (Vijayanagara Coins 2002). The 

ancient African game of mancala traditionally 

employs gray nicker beans as game pieces 

(Driedges 1972). The species is sometimes planted 

as a hedge to prevent undesired entry into property 

(Nelson 1996) and could be planted for dune 

stabilization. Preparations of the seeds and other 

plant parts are used to treat a large range of 

ailments (Burkill 1995, Parrotta 2001). 

Triterpenoids, fatty acid triglicerides, and sterols 

isolated from seeds may possibly explain some of 

the activity (Ali and others 1997, Rostogi and 

others 1996). Seed extracts have been shown to 

lower blood sugar in laboratory animals (Biswas 

and others 1997, Sharma and others 1997) and 

effectively suppress or cure infections of several 

species of round worms (Amarsinghe and others 

1993, Rastogi and others 1996). The fat content of 

gray nicker nut seeds was measured at 34 + 0.83 

percent. The semidrying oil extracted would be 

useful for manufacture of high-quality alkyd resins, 

polishes, and paint (Ajiwe and others 1996). 

References 

Ajiwe, V.I.E., C.A. Okeke, H.U. Agbo, G.A. 

Ogunleye, and S.C. Ekwuozor. 1996. Extraction, 

characterization and industrial uses of velvettamarind, physic-nut and nicker-nut seed oils. 

Bioresource Technology 57(3): 297-299. 

Ali, M.S., S. Shameel, V.U. Ahmad, and K. 

Usmanghanim. 1997. Chemical constituents of 

Caesalpinia bonduc. Pakistan Journal of 

Scientific and Industrial Research 40(1-4): 20-

22. 

Amarsinghe, A.P.G., R.D. Sharma, C. Chaturvedi, 

and D.K. Agarwal. 1993. Anthelmintic effect of 

Ayurvedic recipe Kuberakshadi yoga in 

intestinal worms among children.Journal of 

Research and Education in Indian Medicine 

12(1): 27-31. 

Biswas, T.K., S. Bandyopadhyay, BiswapatiMukherjee, Bhaswar-Mukherjee, B.R. Sangupta, 

and B. Mukherjee. 1997. Oral hypoglycemic 

effect of Caesalpinia bonducella. International 

Journal of Pharmacognosy 35(4): 261-264. 

Burkill, H.M. 1995. The useful plants of West 

Tropical Africa. Vol. 3. Royal Botanic Gardens, 

Kew, UK. 857 p. 

Costin, A.B. 1965. Long-distance seed dispersal to 

Macquarie Island. Nature 206: 317. 

Driedges, W. 1972. The game of Boa, or Mankala, 

in East Africa. Mila 3(1): 7-19. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Markland, J. 2002. Drift seeds. In: West Word, 

Community paper for Glenfinnan, Lochailort, 

Glenuig, Arisaig, Morar, Mallaig, Knoydart, and 

the Small Isles. March 2002. http://road-to-the-
131

isles.org.uk/westword/march2002.html. p. 6-7. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

 391 p. 

Pacific Island Ecosystems at Rick. 2002. 

Caesalpinia bonduc (L.) Roxb., Fabaceae. 

http://www.hear.org/pier3/cabon.htm. 2 p. 

Parrotta, J.A. 2001. Healing plants of Pininsular 

India. CABI Publishing, Wellingford, UK and 

New York. 917 p. 

Rancho Leona. 2002. Rancho Leona jewelry 

samples: rainforest seed chart. 

http://rancholeona.com/seed.html. 2 p. 

Rastogi, S., A.K Shaw, and D.K. Kulshreshtha. 

1996. Characterization of fatty acids of 

antifilarial triglyceride fraction from Caesalpinia 

bonduc. Fitoterapia 67(1): 63-64. 

Sharma, S.R., S.K. Dwivedi, and D. Swarup. 1997. 

hypoglycaemic, antihyperglycaemic and 

hypolipidemic activities of Caesalpinia 

bonducella seeds in rats. Journal of 

Ethnopharmacology. 58(1): 39-44. 

Vermillion, W.G. 2001. Caesalpinia bonduc

(Fabaceae) new to Louisiana. Sida 19(4): 1,181-

1,182. 

Vijayanagara Coins. 2002. Coinage. http://www. 

vijayanagaracoins.com/htm/coinage.htm. 7 p. 

Workman, R.W. 1980. Growing native. The 

Sanibal-Captiva Conservation Foundation, Inc., 

Sanibal, FL. 137 p 

.
132 

Cajanus cajan (L.) Millsp. pigeon pea 

FABACEAE 

Synonyms: Cystisus cajan L. 

 Cajanus bicolor DC. 

 Cajanus flavus DC. 

 Cajanus indicus Spreng. 

 Cajanus luteus Bello 

John K. Francis 

General Description.—Pigeon pea, also known as 

red gram, Congo pea, gungo pea, no eye pea, dhal, 

gandul, gandure, frijol de árbol, and pois cajan, 

occurs in several varieties. The old varieties in 

cultivation and semicultivation in the West Indies 

treated here are semideciduous, short-lived shrubs 

usually 1 to 4 m in height and 1 to 4 cm in basal 

stem diameter. They are usually single stemmed, 

freely branching, and become woody after a few 

months. The wood is moderately hard and brittle. 

They have a deep taproot (to 3 m) with lateral roots 

and nodulated fine roots. The branches and fine 

twigs support abundant light-green or yellow-green 

foliage. Spirally arranged, silky-pubescent, 

trifoliolate leaves have narrowly elliptic, lanceolate 

or oblong leaflets, 2.5 to 9 cm long, the center 

leaflet being slightly longer than the laterals. The 

five- to 12-flowered racemes are axillary. Flowers 

are about 2 cm long, yellow, the standard often 

being orange to purple outside. The legumes, 

which are flattened, somewhat constricted between 

seeds, and 4 to 8 cm long, are mottled bronzepurple when immature, drying to brown. They 

contain two to nine mottled brown (white, red, 

brown, gray, or black in improved varieties) seeds, 

7 to 8 mm long by 6 mm broad. There are 2n = 22, 

44, or 66 chromosomes (Liogier 1988, Long and 

Lakela 1976). 

Range.—Pigeon pea probably originated in India, 

but may have come from Africa. Both are centers 

of diversity for the genus Cajanus. It is clear that 

the species has been under cultivation for a long 

time and was spread by traders thousands of years 

ago. The wild progenitor may be Cajanus 

cajanifolius (Haines) van der Maesen of India and 

Myanmar (van der Maesen 1990). Today, pigeon 

pea is cultivated throughout the tropics and has 

naturalized in many areas including Florida, Puerto 

Rico, and the U.S. Virgin Islands (Liogier 1988, 

Long and Lakela 1976). 

Ecology.—Pigeon pea grows on a broad range of 

well-drained soils, from sands to clays over 

sedimentary, igneous, and metamorphic parent 

materials. It tolerates pH’s of from 4.5 to 8.4 and 

some varieties tolerate 6 to 12 mmhos/cm of 

salinity. However, the species is sensitive to 

waterlogging. Pigeon pea will grow in areas that 

receive from 530 to 4000 mm of mean annual 

precipitation (Van Den Beldt 1988). Pigeon pea 

grows best in temperatures of 18 to 30 °C. It is 

subject to frost damage (Skerman and others 1988). 

The species is grown under cultivation from near 

sea level to 3,000 m in elevation (Van Den Beldt 

1988). Although a large number of insects and 

diseases affect it (American Phytopathological 

Society 2002), the actual incidence of attack is low 

(Center for New Crops and Plants Products 2002). 

Pigeon pea is intolerant of shade and tolerates only 

moderate competition. It does best in full sun on 

bare ground but can grow with side shade or 

broken shade from trees and a low cover of grass 
133

and forbs. Growth is moderately slow during the 

first 2 to 3 months of life during which time 

seedlings are not competitive with grass and 

weeds; afterwards pigeon pea competes well with 

vegetation equal or lower in height (van der 

Maesen 1990). In Puerto Rico, the species persists 

for a generation or two after cultivation and grows 

along roadsides and in waste places where seeds 

have fallen, but the populations are not sustained 

indefinitely. Although seldom devastating, a large 

number of insects and diseases affect pigeon pea 

and can reduce yield or shorten a plant’s life 

(American Phytopathological Society 2002). 

Reproduction.—Pigeon pea flowers nearly 

throughout the year (Red de Grupos de Agricultura 

de Cobertura 2002). The flowers are selfcompatible and usually self-pollinated (Smartt 

1976). Insects visit and pollinate the flowers (5 to 

40 percent cross pollination), but it is not known 

whether they increase seed yield (McGregor 2002). 

Upon drying, the pods spring and coil to release the 

seeds. Pigeon pea seeds weigh about 0.1g and 

germinate in about 2 weeks (Center for New Crops 

and Plants Products 2002). 

Growth and Management.—Pigeon pea is 

normally sown directly into prepared ground. 

Seeding rates for pure stands are 12 to 25 kg of 

seed/ha (Smartt 1976). Seeding depths of 2.5 to 5 

cm are recommended (Center for New Crops and 

Plants Products 2002). No pregermination 

treatment of the seed is needed. Although some 

varieties mature seed in 5 to 6 months, longerlived, tall varieties including those that are more 

competitive in the wild take 10 to 12 months to 

mature seed. These plants live about 5 years 

(Smartt 1976). Experimental yields of 50 dry 

t/ha/year have been demonstrated; yields of 3 to 8 

dry t/ha/year are obtained under normal 

management (Van Den Beldt 1988). 

Benefits.—Pigeon peas are an important food in 

developing tropical countries. An excellent source 

of protein, the seeds (and sometimes the pods) are 

eaten as a vegetable, as a flour additive to other 

foods, in soups, and with rice (Center for New 

Crops and Plants Products 2002). Although they 

vary slightly, typical nutritional values for seeds 

are: moisture, 10.1 percent, protein 19.2 percent, 

fat, 1.5 percent, carbohydrates, 57.3 percent, fiber 

8.1 percent, and ash, 3.8 percent (Smartt 1976). 

About 3.4 million ha were under cultivation in 

1978-1988 period, 88 percent of it in India (Nene 

and Sheila 1990). Although pigeon pea makes 

excellent forage, because of the brittleness of its 

stems, the plant is damaged by browsing, 

especially by cattle (Department of Primary 

Industries 2002). The species is planted as a green 

manure crop, nurse crop, cover crop, a windbreak 

hedge, as a host for lac insects, and as food for silk 

worms. The stalks are used for fuel, thatch, and 

basketry (Center for New Crops and Plants 

Products 2002). Pigeon pea forms root nodules in 

association with Rhizobium sp. bacteria and is 

capable of fixing 41 to 280 kg/ha of nitrogen (Red 

de Grupos de Agricultura de Cobertura 2002). 

Preparations of the leaves are used to treat 

jaundice, inflammation, and sores of the mouth 

(Parrotta 2001). 

References 

American Phytopathological Society. 2002. 

Common names of plant diseases: Diseases of 

pigeonpea [Cajanus cajan (L.) Millsp.]. 

http://www.apsnet.org/online/common/names/ 

pigeon.asp. 5 p. 

Center for New Crops and Plants Products. 2002. 

Cajanus cajan (L.) Millsp. Purdue University. 

http://www.hort.purdue.edu/newcrop/duke_ 

energy/Cajanus_cajun.html. 6 p. 

Department of Primary Industries. 2002. Legumes 

for the tropics: pigeon pea (Cajanus cajan). 

Queensland Government. http://www.dpi.qld. 

gov.au/pastures/4571.html. 2 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyan Books, Miami, FL. 

962 p. 

McGregor, S.E. 2002. Insect pollination of 

cultivated crop plants. Agriculture Research 

Service. http://gears.tucson.ars.ag.gov/book/ 

chap4/pig.html. [not paged]. 

Nene, Y.L. and V.K. Sheila. 1990. Pigeonpea: 

geography and importance. In: Y.L. Nene, S.H. 

Hall, and V.K. Sheila. The pigeonpea. CAB 

International, Wellingford, UK. p. 1-14. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CABI Publishing, Wallingford, UK and 

New York. 917 p. 
134 

Red de Grupos de Agricultura de Cobertura. 2002. 

Base de información sobre especies con 

potential de abonos verdes y cultivos de 

cobertura. Rockefeller Foundation. http://www. 

rockfound.org.mx/cajanusbiesp.html. [not 

paged]. 

Skerman, P.J., D.G. Cameron, and F. Riveros. 

1988. Tropical forage legumes. FAO Plant 

Production and Protection Series 2. Food and 

Agriculture Organization of the United Nations. 

692 p. 

Smartt, J. 1976. Tropical pulses. Longman Group 

Limited, London. 348 p. 

Van Den Beldt, R.J. 1988. Cajanus cajan: it’s 

more than just a pulse crop. Nitrogen Fixing 

Tree association. NFT Highlights 88-06. 5 p. 

van der Maesen, L.J.G. 1990. Pigeonpea: origin, 

history, evolution, and taxonomy. In: Y.L. 

Nene, S.H. Hill, and V.K. Sheila. The 

pigeonpea. CAB International. Wellingford, 

UK. p. 15-46. 
135

Callicarpa americana L. American beautyberry 

VERBENACEAE (also placed in LAMIACEAE alt. Labiatae) 

 

Synonyms: None 

Kristina Connor 

Illustration source: UDA [no date] 

General Description.—American beautyberry, 

also known as beauty-berry, Bermuda mulberry, 

French mulberry, sourbush, sow-berry, or Spanish 

mulberry, is a shrub that can reach 2.4 to 2.7 m in 

height but is typically shorter, averaging 0.9 to 1.8 

m. The irregular, spreading bush has an open 

growth habit and grows as a single plant or in 

colonies. Bark is brown and raised lenticels are 

found on older stems. Branches are scurfy or 

tomentose, gray to reddish brown in color. The 

opposite, short-petioled, simple, deciduous leaves 

are ovate to elliptic in shape. They are pointed at 

both the base and the apex, and the margins are 

coarsely serrate. Leaves are glabrous or slightly 

pubescent above and pubescent with stellate 

trichomes below (Radford and others 1968, Martin 

and Mott 1997). Leaf size is variable, ranging from 

7.6 to 20.3 cm in length and 5 to 12.7 cm in width. 

Range.—American beautyberry ranges through the 

Southeastern United States from Maryland and 

Virginia south to Florida and west to Arkansas, 

Oklahoma, and Texas. It also grows in the 

Bahamas, Bermuda, Cuba, northern Mexico, and in 

the West Indies (Bailey and Bailey 1976, Brickell 

and Zuk 1996). 

Ecology.—American beautyberry is a droughttolerant perennial shrub. Classified as a pioneer 

species (Odenwald and Turner 1988), it is 

common along forest edges and fence rows, and is 

also a common understory plant in open pine 

plantations in the southeastern United States. It is 

found in openings after clear-cutting (Odenwald 

and others 1996) and in moist thickets and 

bordering swamps (Krüssman 1976). American 

beautyberry is very tolerant of fire and resprouts 

easily. However, it is intolerant of deep shade and 

is only found in the better drained areas of 

bottomland hardwood stands. It will grow in a 

variety of soil textures and in a wide pH range 

(USDA-NRCS 2001). Tolerant of temperatures 

ranging from –17 o

C to over 37.8 oC (Martin and 

Mott 1997), it is cultivated for its fall display of 

purple to violet fruits. 

 

Reproduction.—The perfect, bluish white to 

white axillary flowers first appear in early spring. 

The bloom period can continue through late 

summer. It is not uncommon to see both flowers 

and fruits on the same plant (Grelen and Duvall 

1966). The flowers are borne in dichotomous 

cymes 8 to 36 mm long (Bonner in press). The 3- 

to 6-mm fruits are violet to purple berry-like 

drupes, formed in clusters that encircle the stem. 

Each drupe contains four small seeds that 

germinate without scarification or stratification, 

although germination can be slow. If seeds are 

sown in the fall, germination the following spring 

may be excellent (Dirr and Heuser 1987). Bonner 

(in press) reported that seeds can be easily cleaned 

using any type of macerator, even a kitchen 

blender for small lots. Seeds average 600 per 

gram. Longevity of seeds in storage is unknown, 

but they survive for at least 1 year in the soil seed 

bank and are thought to be orthodox. Orthodox 

seeds can be dried to a moisture content of less 

than 12 percent and stored under refrigeration for 

long periods (Roberts 1973). These seeds are 

transported by animals and birds. American 

beautyberry may also be propagated from 

softwood stem cuttings taken in the summer and 
136 

fall that are treated with indol-3-butyric acid (IBA; 

1000 ppm) and put in a mist bed (Dirr and Heuser 

1987). 

Growth and Management.—American 

beautyberry is a short-lived shrub that is easy to 

grow and widely distributed in the Southeastern 

United States. It can be used as a landscape border 

and makes a good mass planting. However, regular 

pruning is required if it is to maintain its form and 

produce masses of the fruits so attractive in the fall 

of the year. The fruits occur only on new growth, 

and plants should be heavily pruned in early spring 

(Martin and Mott 1997). It will not do well in 

areas that experience flooding but can be used to 

restore surface-mined sites (Martin and Sick 

1995). It will resprout after fire (Martin and Mott 

1997) and can be easily transplanted. 

Benefits.—American beautyberry is a good source 

of food for deer and the fruits attract birds. It can 

be used in reclamation work and for erosion 

control (Brown 1945). It easily reseeds in nature. 

References 

Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: 

A Concise Dictionary of Plants Cultivated in the 

United States and Canada. McMillan Publishing 

Co., Inc., New York. 1,290 p. 

Bonner, F.T. (In press). Callicarpa americana L. 

American beautyberry. In: Seeds of Woody 

Plants in the United States. Revised edition. 

Brickell, C. and J.D.. Zuk. 1996. The American 

Horticultural Society A-Z Encyclopedia of 

Garden Plants. DK Publishing, Inc., New York. 

1,092 p. 

Brown, C.A. 1945. Louisiana trees and shrubs. 

Bulletin 1, Louisiana Forestry Commission, 

Baton Rouge, LA. 262 p. 

Dirr, M.A. and C.W. Heuser, Jr. 1987. The 

reference manual of woody plant propagation: 

from seed to tissue culture. Varsity Press, Athens, 

GA. 239 p. 

Grelen, H.E. and V.L. Duvall. 1966. Common 

plants of longleaf pine-bluestem range. Research 

Paper SO-23. U.S. Department of Agriculture 

Forest Service, Southern Forest Experiment 

Station, New Orleans, LA. 96 p. 

Krüssmann, G. 1976. Manual of Cultivated BroadLeaved Trees and Shrubs. Volume I, A-D. 

Timber Press, Beaverton, OR. 448 p. 

Martin, C.O. and S.P. Mott. 1997. American 

Beautyberry (Callicarpa americana). Section 

7.5.8, U.S. Army Corps of Engineers Wildlife 

Resources Management Manual. Ecosystem 

Management and Restoration Research Program 

Technical Report EL- 97.15. Conservation 

Communications, Vicksburg, MS. 17 p. 

Martin, H. and G. Sick. 1995. American 

beautyberry for borrow pit reclamation in South 

Carolina Restoration and Management Notes 

13(1): 90-96. 

Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope. 

1996. Plants for American Landscapes. Louisiana 

State Univ. Press, Baton Rouge, LA. 266 p. 

Odenwald, N.G. and J.R. Turner. 1988. Plants for 

the South: A guide for landscaping design. 

Claitor’s Publication Division, Baton Rouge, LA. 

565 p. 

Radford, A.E., H.E. Ahles, and C.R. Bell. 1968. 

Manual of the Vascular Flora of the Carolinas. 

University of North Carolina Press, Chapel Hill. 

 1,183 p. 

Roberts, E.H. 1973. Predicting the storage life of 

seeds. Seed Science and Technology 1: 499-514. 

USDA, NRCS. 2001. The PLANTS database. 

National Plant Data Center, Baton Rouge, 

http://plants.usda.gov. LA [not paged]. 

USDA. [no date]. Southern wetland floral field 

office guide to plant species. U.S. Department 

of Agriculture Soil Conservation Service, South 

National Technical Center, Fort Worth, TX. 

http://www.npwrc.usgs.gov. [not paged] 

.
137

Calotropis procera (Ait.) Ait. f. giant milkweed 

ASCLEPIADACEAE 

Synonym: Asclepias procera Ait. 

John K. Francis 

General Description.—Giant milkweed is also 

known as sodom apple, calotrope, French cotton, 

small crown flower (English), algodón de seda, 

bomba (Spanish), cotton-france, arbre de soie, and 

bois canon (French) (Howard 1989, Liogier 1995, 

Neal 1965, Parrotta 2001). 

This plant is a soft-wooded, evergreen, 

perennial shrub. It has one or a few stems, few 

branches, and relatively few leaves, mostly 

concentrated near the growing tip. The bark is 

corky, furrowed, and light gray. A copious white 

sap flows whenever stems or leaves are cut. Giant 

milkweed has a very deep, stout taproot with few 

or no near-surface lateral roots. Giant milkweed 

roots were found to have few branches and reach 

depths of 1.7 to 3.0 m in Indian sandy desert soils 

(Sharma 1968). The opposite leaves are oblongobovate to nearly orbicular, short-pointed to blunt 

at the apex and have very short petioles below a 

nearly clasping, heart-shaped base. The leaf blades 

are light to dark green with nearly white veins. 

They are 7 to 18 cm long and 5 to 13 cm broad, 

slightly leathery, and have a fine coat of soft hairs 

that rub off. The flower clusters are umbelliform 

cymes that grow at or near the ends of twigs. The 

flowers are shallowly campanulate with five sepals 

that are 4 to 5 mm long, fleshy and variable in 

color from white to pink, often spotted or tinged 

with purple. The fruits are inflated, obliquely ovoid 

follicles that split and invert when mature to 

release flat, brown seeds with a tuft of white hairs 

at one end (Howard 1989, Liogier 1995, Little and 

others 1974). 

Range.—Giant milkweed is native to West Africa 

as far south as Angola, North and East Africa, 

Madagascar, the Arabian Peninsula, southern Asia, 

and Indochina to Malaysia (Rahman and Wilcock 

1991). The species is now naturalized in Australia, 

many Pacific islands, Mexico, Central and South 

America, and the Caribbean islands. 

Ecology.—Giant milkweed favors open habitat 

with little competition. This condition is most 

completely met in overgrazed pastures and 

rangeland. Other common habitats are beachfront 

dunes, roadsides, and disturbed urban lots. The 

species grows in dry habitat (150 to 1000 mm 

precipitation) and sometimes in excessively 

drained soils in areas with as much as 2000 mm of 

annual precipitation. Giant milkweed may be found 

in areas up to 1,000 m in elevation in India 

(Parrotta 2001). It roots very deeply and rarely 

grows in soils that are shallow over unfractured 

rock. Soils of all textures and derived from most 

parent materials are tolerated, as well as soils with 

high sodium saturation. Beachfront salt spray is not 

detrimental. Competition with tall weeds, brush, 

and especially grass weakens existing plants, and 

being overtopped and shaded by trees soon 

eliminates them. During droughts in Puerto Rico, 

giant milkweed is attacked by the orange aphid, 

Aphis nerii Boyer de Fonscolombe, which causes 

defoliation, death of branches, and aborted fruits 

(Little and others 1974). 

Reproduction.—Flowering and fruiting takes 

place throughout the year (Little and others 1974). 

Hundreds to thousands of seeds may be produced 

per plant each year. The seeds in a Puerto Rican 

collection averaged 0.0095 + 0.0027 g/seed or 
138 

about 100,000 seeds/kg (author’s observation). 

Eighty-nine percent germination took place in 

potting mix between 7 and 64 days after sowing. 

Half the seed weight was found in the wing (silk). 

The seeds are dispersed by wind and may fly for 

several hundred yards in gentle breezes. Seedlings 

may arise in abundance after rainy periods, but 

only a few survive the first season. Using the 

reserves in its large taproot, giant milkweed can 

resprout year after year when burned or cut. 

Growth and Management.—Giant milkweed 

usually reaches heights of about 2 m, but may 

occasionally reach 5 m in height and stem 

diameters of 25 cm (Little and others 1974). 

Growth is about 1 m the first year after sprouting 

in Puerto Rico. Senescence of individual stems 

takes place in about 5 years, but plants often 

resprout afterwards. Giant milkweed does not form 

dense stands, normally occurring as scattered 

individuals. It can be a serious weed in pastures, 

overgrazed rangelands, and poorly managed hay 

fields. Although it probably is not possible to 

eliminate existing stands through management 

without ceasing grazing and harvest, it may be 

possible to prevent their establishment. Some form 

of chemical control would seem to be the only 

practical option for eliminating existing stands but 

no specific guidelines are available. Establishing 

giant milkweed has been advocated for 

environmental protection and as a nurse crop for 

more valuable species (Campolucci and Paolini 

1990). This can be done easily by planting 

containerized seedlings or rooted cuttings. 

Benefits.—Giant milkweed tissues, especially the 

root bark, are used to treat a variety of illness 

including leprosy, fever, menorrhagia, malaria, 

and snake bite (Parrotta 2001). The latex is toxic 

and can cause blisters and rash in sensitive 

persons. The plant is occasionally grown as an 

ornamental in dry or coastal areas because it is 

handsome, of a convenient size, and is easy to 

propagate and manage. It is recommended as a 

host plant for butterflies (Mikula 2001). In the 

past, the silky hairs were used to stuff pillows 

(Little and others 1974). Giant milkweed was 

tested as a host for sandalwood, Santalum album 

L., a partial root parasite. It resulted in greater 

growth of sandalwood than all other species tested 

(Shinde and others 1993). Extracts, chopped 

leaves, and latex have shown great promise as 

nematicides, in vitro and in vivo (Anver and Alam 

1992, Charu and Trivedi 1997). Sheep, goats, and 

camels will eat the leaves of giant milkweed 

during droughts, but consumption is low. If the 

leaves are chopped and mixed with other feed, 

consumption greatly increases with no ill effects. 

(Abbas and others 1992, Nehra and others 1987). 

Shade-dried giant milkweed leaves contained 94.6 

percent dry matter, 20.9 percent ash, 19.6 percent 

crude protein, 2.2 percent fat, 43.6 percent acid 

detergent fiber, and 19.5 percent neutral detergent 

fiber (Abbas and others 1992). Although it is 

lightweight, the wood is used in impoverished 

desert areas for a cooking fuel (Varshney and Bhoi 

1988). 

References 

Abbas, B., A.E. El Tayeb, and Y.R. Sulleiman. 

1992. Calotropis procera: feed potential for arid 

zones. Veterinary Record 131(6): 132. 

Anver, S. and M.M. Alam. 1992. Effect of latex 

seed dressing on interacting root-knot and 

reniform nematodes. Afro-Asian Journal of 

Nematology 2: 1-2, 17-20. 

Campolucci, P. and C. Paolini. 1990. 

Desertification control in the Sahel regions—

low-cost large-scale afforestation techniques. 

Note Tecniche 10. Centro di Sperimentazione 

Agricola e Forestale di Sperimentazione per la 

Pioppicoltura. 24 p. 

Charu-Jain and P.C. Trivedi. 1997. Nematicidal 

activity of certain plants against root-knot 

nematode, Meloidogyne incognita, infecting 

chickpea, Cicer arietinum. Annals of Plant 

Protection Sciences 5(2): 171-174. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Dicotyledoneae. 

Part 3. Vol. 6. Arnold Arboretum, Harvard 

University, Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial 

de la Universidad de Puerto Rico, San Juan, PR. 

617 p.

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 

Mikula, R. 2001. Butterfly plants for your garden. 

www.butterflybreeders.com/pages/bflygdning/bu

tterflyplants.html. 5 p. 
139

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Museum 

Press, Honolulu, HI. 924 p. 

Nehra, O.P., M.C. Oswal, and A.S. Faroda. 1987. 

Management of fodder trees in Haryana. Indian 

Farming 37(3): 31, 33. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CAB International, Wallingford, UK and 

New York. 944 p. 

Rahman, M.A. and C.C. Wilcock. 1991. A 

taxonomic revision of Calotropis

(Asclepiadaceae). Nordic Journal of Botany 

11(3): 301-308. 

Sharma, B.M. 1968. Root systems of some desert 

plants in Churu, Rajasthan. Indian Forester 

94(3): 240-246. 

Shinde, S.R., R.D. Ghatge, and S.S. Mehetre. 1993. 

Comparative studies on the growth and 

development of sandalwood tree in association 

with different hosts. Indian Journal of Forestry 

16(2): 165-166. 

Varshney, A.C., and K.L. Bhoi. 1988. Cloth from 

bast fibre of the Calotropis procera (aak) plant. 

Biological wastes 26(3): 229-232.
140 

Capparis indica (L.) Druce sapo prieto 

CAPPARACEAE 

Synonyms: Capparis indica (L.) Rawc. & Rendle 

 Breynia indica L. 

 Capparis breynia L. 

 Capparis amygdalifolia Jacq. 

 Capparis amygdalina Lam. 

 Linnaeobreynia indica (L.) Hutch. 

John K. Francis 

General Description.—Sapo prieto, a name used 

in Puerto Rico that means black toad in Spanish, is 

also known as caper, linguam, burro, white willow, 

colorín, vara prieta, palo zapo, taiche, endurece 

maíz, curumo, guacoco, naranjuelo, pachaca, olivo 

macho, olivo, bois de mêche, bois-puant, bois-noir, 

pois à mabou, paaloe pretae, and raba stokki (Little 

and others 1974, Zamora 1989). It is a shrub or a 

small tree, which under favorable circumstances 

may reach 8 m in height and 15 cm in diameter at 

breast height. Sapo prieto usually has a single stem 

unless it has been damaged, but the species is very 

limby and may have major branches emerging low 

on the trunk. The stem and branch bark is gray or 

light brown and smooth with a bitter, yellow or 

reddish inner bark. The twigs, petioles, and 

undersides of leaves are covered with silvery, gray, 

or golden scales. The alternate, simple leaves are 

supported by 6- to 10-mm petioles and have 

narrowly elliptic, leathery blades 4 to 11 cm long 

and 2 to 4.5 cm broad. The inflorescences 

(corymbs) are near the ends of branches and 

contain a few small, white flowers with long white 

stamens and short yellow anthers. The fruits are 

silvery-brown legume-like pods 4 to 26 cm long 

that split along one side at maturity to expose a 

bright-red interior and pulp-covered seeds. The 

seeds are black, elliptic, and 5 to 7 mm in diameter 

(Howard 1988, Liogier 1985, Little and others 

1974, Zamora 1989). 

Range.—The range of sapo prieto includes 

Jamaica, Hispaniola, Puerto Rico, the Virgin 

Islands, the Lesser Antilles, the Dutch Antilles, 

Trinidad, Venezuela, Southern Mexico to Costa 

Rica and possibly Panama (Liogier 1985, Little and 

others 1974, Zamora 1989). 

Ecology.—Sapo prieto grows in a wide range of 

well-drained soils derived from both sedimentary 

(including limestone), igneous, and metamorphic 

rocks. It may be found from near sea level to about 

600 m on steep as well as level slopes, and in areas 

receiving from 750 to about 1700 mm of 

precipitation in Puerto Rico. In Costa Rica, sapo 

prieto grows from sea level to 300 m in elevation 

in very rocky soils (Zamora 1989). The species is 

reported to root deeply (Zoológico Virtual de 

Barranquilla 2002). Sapo prieto may be found 

growing in secondary and remnant forests. It is not 

a pioneer and will rarely be found in abandoned 

fields or areas where all vegetation has been 

removed. It is moderately intolerant of shade and 

consequently does not grow under dense forest 

canopies. It most frequently grows in an 

intermediate crown position in relatively open dry 

forests or as a codominant in clumps of shrubs and 

low trees in disturbed secondary forests. 

Reproduction.—Little and others (1974) state that 
141

sapo prieto flowers and fruits intermittently. 

Flowers were observed in Costa Rico in May and 

September and fruits were present from May to 

September (Zamora 1989). A collection of sapo 

prieto fruits from Puerto Rico weighed an average 

of 6.948 + 0.773 g/fruit. The air-dry seeds collected 

from these fruits weighed an average of 0.0838 +

0.0021 g/seed or 12,000 seeds/kg. Sown on 

commercial potting mix without any pre-treatment, 

the seedlings began to emerge in 7 days and 

reached a maximum germination of 49 percent in 

64 days (author’s observation). The seeds are 

covered with a thin, pasty, scarlet pulp that is 

presumably eaten by birds that transport the seeds. 

Growth and Management.—Sapo prieto has been 

referred to as slow growing (Zoológico Virtual de 

Barranquilla 2002). Weaver (1990) measured one 

tree in St. John, U.S. Virgin Islands, as a part of a 

larger study and found a 0.4 mm annual diameter 

increment over a 5-year period. Increases in sapo 

prieto density can probably be encouraged by 

treatments to dry and moist forests near seed 

sources to eliminate dense shade and create brush 

patches and small openings. No wildland planting 

experience has been reported. 

Benefits.—Sapo prieto makes a good ornamental 

that is evergreen, with attractive bicolor leaves. It 

has showy flowers and fruits, and a crown that can 

be shaped by pruning. However, the species is not 

commonly used because it grows very slowly 

(Virgin Islands Wetlands Reserve 2002, Zoológico 

Virtual de Barranquilla 2002). The wood is heavy 

and hard and useful for fuel, stakes, and smalldiameter fence posts. Sapo prieto contributes to the 

biodiversity of the forests it inhabits, helps protect 

those sites from erosion, and furnishes minor 

amounts of food and cover for wildlife. The 

flowers are nectar sources for butterflies (Debrot 

and others 1999). 

References 

Debrot, A.O., J.Y. Miller, L.D. Miller, and B.T. 

Leysner. 1999. The butterfly fauna of Curaçao, 

West Indies: 1996 status and long-term species 

turnover. Caribbean Journal of Science 35(3-4): 

184-194. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p.

Virgin Islands Wetlands Reserve. 2002. A list of 

plants tagged along the trail at the reserve and 

some of their uses. Virgin Islands Wetlands 

Reserve, University of the Virgin Islands, St. 

Croix, U.S. Virgin Islands. 

http://rps.uvi.edu/VIMAS/plantlist.htm. 5 p. 

Weaver, P.L. 1990. Tree diameter growth rates in 

Cinnamon Bay Watershed, St. John, U.S. Virgin 

Islands. Caribbean Journal of Science 26(1-2): 

1-6. 

Zamora, N. 1989. Flora arborescente de Costa 

Rica. Editorial Techologica de Costa Rica, San 

Jose, Costa Rica. 262 p. 

Zoológico Virtual de Barranquilla. 2002. 

Resultados de busca por: Olivo macho. 

http://zoobaq.org/formas/consultaveg2.php3?co

mun=Olivo. 1 p. 
142 

Capraria biflora L. goat-weed 

SCROPHULARIACEAE 

Synonyms: Capraria lanceolata Vahl 

John K. Francis 

General Description.—Goatweed, also known as 

wild tea, savadilla, té del país, thé du pays, ditay 

paye, and balsaminha, is an evergreen (or 

sometimes suffruticose) shrub to 1.5 m in height 

and 6 mm in basal stem diameter. The plant is 

supported by a well-developed taproot with laterals 

through its length. The shrub usually has multiple 

stems and many fine branches and twigs. The 

degree of hairiness is variable. Most plants have 

foliage the full length of their stems. The leaves are 

subsessile, narrowly lanceolate to obovate, 2 to 12 

cm long, with entire to serrate margin. The white, 

five-lobed, campanulate, 5 mm-long flowers are 

borne in groups of one to three in the leaf axils. 

Ovoid, 5-mm capsules contain many tiny, yellow 

seeds about 0.4 mm long (Howard 1989, Liogier 

1995, Stevens and others 2001). 

Range.—Goatweed is native to Florida and Texas, 

the Bahamas, the West Indies, Trinidad, Mexico 

through Central America, South America as far 

south as Bolivia, and the Galapagos Islands (Flora 

of Texas Consortium 2002, Grisebach 1963, 

Pezzatti and others 1998). The species has 

naturalized in Ghana, the Cape Verde Islands, and 

Mauritius (Burkill 2000). 

Ecology.—Goatweed is intolerant of shade. It is 

usually open-grown but may be found in stands of 

brush and in the understory of open forest. It will 

die as the shade becomes dense. In Puerto Rico, it 

grows in pastures, occasionally burned grasslands, 

dry, early secondary forest, rocky coasts, and sandy 

areas behind beaches. It is reported to grow on 

beaches, sandy soils, and disturbed areas in Florida 

(Long and Lakela 1976). The species grows on 

soils derived from both sedimentary and igneous 

rocks in a wide range of soil textures and fertility 

but requires well-drained conditions. In Puerto 

Rico, goatweed occurs in areas that receive from 

750 to about 1700 mm of precipitation. It grows at 

elevations from near sea level to 1,000 m in 

Nicaragua (Stevens and others 2001). Disturbance 

is probably required for establishment. Once 

established, goatweed competes relatively well 

with grass, weeds, and low brush. It is little 

browsed by cattle. 

Reproduction.—Goatweed blooms and fruits all 

year (Long and Lakela 1976, Stevens and others 

2001). A collection of seeds from Puerto Rico 

averaged 42,300 seeds/g. Placed on moist filter 

paper, 21 percent germinated between 10 and 35 

days after sowing. Germination is epigeal. The tiny 

seeds are apparently dispersed by wind and water, 

or by fortuitously sticking to animals or machinery. 

Natural seedlings are not abundant. Plants readily 

sprout from the lower stem or root when cut or 

burned. 

Growth and Management.—Individual goatweed 

stems live 2 or 3 years or more and are replaced by 

other sprouts. The plant may live for many years. 

Sprouts grow about 0.5 m in their first year, much 

less in succeeding years. Large numbers of seeds 

can be collected by picking unopened capsules, 

drying them in a paper bag, and separating with a 

fine screen. Nursery procedures have not been 
143

published. Goatweed is considered weedy in 

pastures. In the absence of tested procedures for its 

control, standard techniques of grubbing or spot 

spraying with broadleaf herbicides is 

recommended. Mowing has little lasting effect on 

stands. 

Benefits.—Goatweed helps revegetate disturbed 

areas, protects the soil, and furnishes cover for 

wildlife. It is a nectar source for butterflies and is 

recommended in natural landscaping (University of 

Florida Extension 2002). Goatweed has many uses 

in herbal medicine. A tea prepared from the leaves 

is used as an eyewash (Burkill 2000), to soothe 

skin itch (Secritaría de Medio Ambiente y 

Recursos Naturales 2002), and as a general tonic. 

However, overindulgence can result in stupor, 

disorientation, and paralysis (Guzmán 1975). The 

infusion is also used to treat fever, flu, vomiting, 

childbirth recovery, diarrhea, hemorrhoids, 

rheumatism, and swelling (Scofield 2002). The 

alkaloid, biflorine, present in the leaves has 

antibiotic properties (Burkill 2000), and 

sesquiterpenoids, caprariolides A and B from aerial 

parts, have demonstrated strong insecticidal 

activity against adult sweet potato weevils, Cylas 

formicarius elegantulus (Collins and others 2000). 

References 

Burkill, H.M. 2000. The useful plants of West 

Tropical Africa. Vol. 5. Royal Botanical Garden, 

Kew, UK. 686 p. 

Collins, D.O., W.A. Gillimore, W.R. Reynolds, 

L.A.D. Williams, and P.B. Reese. 2000. New 

skeletal sesquiterpenoids, caprariolides A-D, 

from Capraria biflora and their insecticidal 

activity. Journal of Natural Products 63: 1,515-

1,518. 

Flora of Texas Consortium. 2002. Texas vascular 

plant checklist: Scrophulariaceae. 

http://csdl/tamu.edu/FLORA/ftc/dft/ftc_scr.htm. 

5 p. 

Grisebach, A.H.R. 1963. Flora of the British West 

Indian Islands. J. Cramer-Weinheim. New York. 

789 p. 

Guzmán, D.J. 1975. Especies útiles de la flora 

Salvadoreña. Ministerio de Educación, 

Dirección de Publicaciones, San Salvador, El 

Salvador. 703 p. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Pezzatti, B., T. Irzan, and D. Cherix. 1998. Ants 

(Hymenoptera, Formicide): Lost paradise. 

Darwin Foundation, Galapagos, Equador. 

http://www.darwinfoundation.org/articles/n5900

049802.html. 10 p. 

Scofield, D. 2002. Medicinal usage: Capraria 

biflora L. (Scrophulariaceae). http://www. 

cassiakeyensis.com/sofl_plants/med_caprariabifl

ora.html. 1 p. 

Secritaría de Medio Ambiente y Recursos 

Naturales. 2002. Especies con usos do 

maderables en bosques de encino, pino, y pinoencino. http://www/semarnat.gob.mx/pfnm/ 

CaprariaBiflora.html. 2 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanical Garden, St. Louis, MO. p. 

1,911-2,666. 

University of Florida Extension. 2002. Ask a 

master gardener: List of native Florida 

specimens for butterfly gardening. http:// 

volusia.org/extension_service/butterfly.htm. 2 p.
144 

Capsicum annuum L. bird pepper 

SOLANACEAE 

Synonyms: Capsicum indicum microcarpum var. aviculare Dierb. 

 Capsicum bacatum of authors, not of L. 

John K. Francis 

General Description.—Bird pepper is also known 

as red pepper, wild pepper, wild chili, pimiento, 

and ají de gallina (Liogier 1995). The current 

growth is herbaceous, but it later hardens and 

becomes woody. The wood is brittle. Older plants 

are multistemmed and very branchy, with a thick, 

leafy crown. The leaves are dark green, ovate to 

lanciolate, 4 to 15 cm long, with petioles 0.5 to 3 

cm long. The twigs are stiff and straight between 

nodes. The flowers are mostly solitary or in pairs. 

The corolla is greenish-white or yellowish-white, 1 

to 2 cm broad, with blue, violate, or yellow 

anthers. The fruits, which are ellipsoidal berries 

with thin flesh, are 0.4 to 3 cm long (usually about 

0.8 cm in wild plants) and red or orange-red. The 

seeds are cream to yellow in color (Howard 1989, 

Liogier 1995). The variety growing as a shrub in 

wildland vegetation in Puerto Rico, Florida, and 

elsewhere is C. annuum var. glabriusculum

(Dunal) H. Pickersgill (Liogier 1995). 

 

Range.—Bird pepper ranges today from the 

Southern United States to Argentina and 

throughout the West Indies (Liogier 1995). The 

species has been widely spread and cultivated by 

Native Americans and European settlers. Escapes 

from cultivation have masked the original range. 

Bird pepper may be found competing with native 

vegetation in Puerto Rico (author’s observations), 

Florida (Nelson 1996), Texas (Everitt and Drawe 

1993), and Arizona (Tewksbury and other 1999). It 

is felt that the species was introduced into Puerto 

Rico by the aboriginal inhabitants (Barrett 1925). 

Bird pepper is also widely cultivated and escaped 

in the tropical and subtropical Eastern Hemisphere 

(Bailey 1941). It is the wild ancestor of the 

pimento, the bell pepper, and some of the hot 

peppers (Bailey 1941). The domestication of these 

pepper varieties from bird pepper is reported to 

have taken place in Mexico or Central America 

(Hawkens 1991). 

Ecology.—In Puerto Rico, bird pepper inhabits the 

subtropical moist forest and the wetter portion of 

the subtropical dry forest (about 800 to 2000 mm 

of annual precipitation). It is cold sensitive, and 

hot, dry weather is desirable for fruit ripening 

(California Antilles Trading Consortium 2001). In 

favorable habitat, bird pepper can grow under a 

broken canopy or in disturbed areas without tree 

cover. However, in Arizona, the plants were found 

exclusively under partially shading (nurse) plants 

(Tewksbury and other 1999). Under conditions of 

low fertility, moisture, and light, bird pepper plants 

do not become large and probably do not live more 

than 1 year. The species favors a near-neutral soil 

reaction and a generous supply of bases. Welldrained soil with a sandy loam or silt loam texture 

is best (California Antilles Trading Consortium 

2001). After it is well established, bird pepper can 

survive dry seasons of 2 to 4 months. During this 

period, it will partially defoliate and take on a 

wilted appearance, but will refoliate and grow 

vigorously after the rains return. 

Reproduction.—When conditions are favorable, 

bird pepper begins flowering at about 3 months of 

age (California Antilles Trading Consortium 2001) 

and flowers and fruits throughout the year. A 

sample of fruits collected in Puerto Rico weighed 

an average of 0.0926 + 0.0057 g/fruit. Seeds from 

the sample weighed an average of 0.0039 + 0.0001 

g/seed or 260,000 seeds/kg. Bird pepper plants are 

prolific seed producers. Large plants may produce 

hundreds of fruits per year. Fruits in the above 

sample contained an average of 8.2 seeds/fruit. 
145

Twenty-eight percent of the seeds in this collection 

germinated between 17 and 118 days after being 

sown in potting mix. Seedlings may be established 

by sowing in prepared seed spots in the field or 

grown in the nursery and transplanted. The seeds 

are dispersed by birds that are immune to the 

capsaicin the fruits contain. Rodents that would 

otherwise chew up and destroy the seeds will not 

eat the fruits (UniSci 2001). Seedlings and plants 

are common but scattered in early secondary forest 

in Puerto Rico. However, bird pepper may 

occasionally form small thickets. 

Growth and Management.—Because the species 

name “annuum” means annual and because 

cultivated forms are, or are treated as, annuals, bird 

pepper has been assumed to be an annual (see 

Liogier 1995). In its wild state, it is, in fact, a 

short-lived perennial (Floridata 2001), living 3 or 4 

years, if conditions are favorable. Plants may 

occasionally reach 5 m in height (Howard 1989). 

Heights in Puerto Rico are commonly 2 to 3 m. 

The stems are slender, about 1 cm in diameter, in 

larger plants (author’s observation). 

Benefits.—Bird pepper has been cultivated for 

thousands of years and is the wild ancestor for 

hundreds of named varieties of cultivated peppers. 

However, the wild fruits are still harvested and 

used today. Bird pepper fruits are “hot” to very 

“hot” with a slight musky flavor. They are used to 

flavor food and make sauces. Capsicum 

derivatives are used in a wide variety of medicinal 

applications, mostly related to pain relief (eg: sore 

muscles, toothache, phantom limb pain). 

Capsaicin causes the brain to release endorphins 

that promote a sense of well-being and deadens 

pain receptors (Floridata 2001). Concentrated 

pepper sprays are used for riot control and 

personal defense. Gardeners also use pepper 

extracts to protect crops from insects and animals. 

Extracts of leaves and fruits of bird pepper have 

shown insecticidal and fungicidal properties in 

laboratory tests (Hongo and Karel 1986, Patil and 

Joi 1992, Williams and Mansingh 1993). 

However, to sensitive individuals, exposure to C. 

annuum fruits can be dangerous. This species 

ranked fourth in plant poisoning incidents reported 

to American poison information centers 

(Krenzelok and Provost 1995). In Texas and 

Arizona, several species of birds including the Rio 

Grande turkey and the curve-billed thrasher eat the 

fruits (Everitt and Drawe 1993, UniSci 2001). 

References 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. Vol. 1. The MacMillan Co. New 

York, NY. 1,200 p. 

Barrett, O.W. 1925. The food plants of Puerto 

Rico. The Journal of the Department of 

Agriculture of Puerto Rico 9(2): 168. 

California Antilles Trading Consortium. 2001. 

Capsicum peppers. www.calantilles.com/ 

capsicum_peppers.htm. 9 p. 

Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs 

and cacti of South Texas. Texas Tech University 

Press. 213 p. 

Floridata. 2001. Capsicum spp. 

www.floridata.com/ref/c/caps_spp.cfm. 6 p. 

Hawkens, J.G. 1991. The centers of plant diversity 

in Latin America. Diversity 7(1/2): 7-9 

Hongo, H., and A.K. Karel. 1986. Effects of plant 

extracts on insect pests of common beans. 

Journal of Applied Entomology 102(2): 164-

169. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Krenzelok, E.P., and F.J. Provost. 1995. The ten 

most common plant exposures reported to 

poison information centers in the United States. 

Journal of Natural Toxins 4(2): 195-202. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Nelson, Gil. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc. Sarasota, FL. 

391 p. 

Patil, B.H. and M.B. Joi. 1992. Inhibition of 

tomato spotted wilt virus with leaf extracts of 

some plants. Journal of Maharashtra 

Agricultural University 17(2): 340-341. 
146 

Tewksbury, J.J., G.P. Nabhan, D. Norman, H. 

Suzan, J. Tuxill, and J. Donovan. 1999. In situ 

conservation of wild chilis and their biotic 

associates. Conservation Biology 13(1): 98-107. 

UniSci 2001. Zing in the flavor helps chili pepper 

plants survive. Daily University Science News. 

http://unisci.com/stories/20013/0726015.htm. 

3 p. 

Williams, L.A.D. and A. Mansingh. 1993. 

Pesticidal potential of tropical plants—I. 

Insecticidal activity of leaf extracts of sixty 

plants. Insect Science and its Application 14(5): 

697-700.
147

Capsicum frutescens L. red pepper

SOLANACEAE 

Synonyms: Capsicum fastigiatum Blume 

 Capsicum annuum L. var. frutescens (L.) Kuntze 

John K. Francis 

General Description.—Red pepper, also known as 

bird pepper, chili pepper, Cayenne pepper, Guinea 

pepper, ají, and ají picante, is a short-lived 

evergreen shrub usually 1 to 1.5 m in height and 1 

to 3 cm in basal stem diameter. The shrub is 

supported by a short to long taproot (depending on 

soil conditions), many spreading lateral roots, and 

moderately abundant fibrous roots. The stem and 

larger branches of mature plants are woody but 

moderately soft and weak. Bark of stems and older 

branches is light gray. The form is upright, the 

abundant branching is often dichotomous, and the 

branches and twigs are slender. The ovate to ovatelanceolate leaves vary in size. The larger of them 

are 4 to 12 cm long and 1 to 4.5 cm broad. 

Greenish-white to yellowish-white flowers with 

blue, violet, or yellow anthers occur in groups of 

two or more at the nodes. The berries are red or 

red-orange at maturity, elongated with a pointed or 

rounded tip, 1.5 to 3.5 cm long and 0.5 to 1.2 cm 

thick. The fruits are somewhat dry and contain few 

to many (depending on fruit size) cream to yellow 

lenticular seeds about 3 mm in diameter. The fruits, 

especially the seeds and placenta, have a biting, 

pungent taste. The species has 2n = 24 

chromosomes (Bailey 1941, Bentley and Trimer 

1880, Bosland and Votava 2000, Liogier 1995). 

Range.—The original range of red pepper is 

unknown, but it is believed to have been 

domesticated in Central America, possibly Panama, 

thousands of years ago. It spread throughout the 

Neotropics before Columbus and has since become 

almost pan-tropically cultivated and naturalized 

(Bosland and Votava 2000). Unlike C. annuum L., 

which has many widely varying varieties, 

domesticated red pepper has relatively few 

varieties with a minimum of variation (Bentley and 

Trimer 1880) and is not greatly different from the 

wild type. 

Ecology.—Red pepper grows on soils of all 

textures in a wide range of fertilities. Moist, welldrained conditions and loose structure is best for 

rapid growth. Soil pH’s of 4.3 to 9.7 are tolerated 

(Center for New Crops and Plant Products 2002). 

The species is intolerant of shade: it will grow with 

broken overhead shade and moderate competition 

from grass and forbs, but fruits best in full sun. Red 

peppers can be cultivated in areas that receive from 

30 to 430 cm of annual precipitation at elevations 

from near sea level to more than 2,000 m. The 

species is not frost tolerant and does not grow in 

temperatures below 7 °C (Center for New Crops 

and Plant Products 2002). Wild red pepper grows 

as individuals or groups in gardens, fields, vacant 

lots, river flood plains, abandoned fields, roadsides, 

and early secondary forest. Although damping-off 

fungi, various species of insects, and nematodes 

sometimes damage or kill individual plants, serious 

effects are rarely widespread. 

Reproduction.—After about 3 months of growth, 

red pepper plants flower (Center for New Crops 

and Plant Products 2002) and fruit continuously as 
148 

long as they live. The flowers are insect pollinated 

(Bosland and Votava 2000). Production of fruits 

and seeds can be abundant. Fresh fruits from 

several plants in a stand in Puerto Rico averaged 

0.3668 + 0.0242 g/fruit. The variability was high 

(CV = 39.5 percent). They ranged from 2 to 35 

seeds/fruit and averaged 16.6 + 2.9 seeds/fruit. Airdried seeds averaged 0.0041 + 0.0001 g/seed or 

244,000 seeds/kg. Placed in moist potting mix, 72 

percent of the seeds germinated between 13 and 34 

days after sowing. Germination is epigeous. The 

seeds can be safely stored under refrigeration after 

air-drying and need no treatment before planting. 

The seeds are dispersed by birds that eat the fruits 

and are immune to the pungent chemicals they 

contain. Young plants coppice when cut or broken. 

Rooting of cuttings and tissues has been 

demonstrated for Capsicum (Bosland and Votava 

2000). 

Growth and Management.—Under continually 

favorable conditions, red peppers live about 2 

years. They grow rapidly during the first year, then 

much more slowly, and finally dwindle and die. 

Red peppers grown commercially are managed as 

annuals, much as other Capsicum peppers but for a 

longer season. Plants that arise spontaneously in 

gardens and vacant land are usually allowed to 

grow for occasional future harvest. 

Benefits.—Red pepper is used and loved the world 

over as a condiment, added to food fresh, dried, 

refined, and ground (for Cayenne pepper and 

curry), and as the principal or incidental ingredient 

in sauces. The source of the popular biting 

sensation are the capsaicinoids, principally 

capsaicin, which ranges from 600 to 13,000 ppm in 

the fruits (Center for New Crops and Plant 

Products. 2002). The fruits are an excellent source 

of vitamins A and C (Bosland and Botava 2000). A 

few thousand hectares are grown commercially, 

and probably even more are grown in gardens or 

harvested from the wild in rural areas of the 

tropics. Coatings and powders are used with 

varying degrees of success to deter browsing 

animals and insects, and break children of thumb 

sucking and nail biting. Pepper spray, whose active 

ingredient is capsaicin, is used widely for personal 

protection, law enforcement, and defense. Acetone 

and petroleum ether extracts caused complete 

mortality of rice weevil (Sitophillus oryzae) in 15 

days. Fruit powder was much less effective (ElLakwah and others 1997). Red pepper has many 

medicinal applications. Some of the most widely 

used and reliable are as a salve to relieve muscle, 

joint, and toothache pain, to treat cough, asthma, 

and sore throat, as a stimulant, and to treat stomach 

ache, seasickness, and flatulence. Anciently, it was 

even used as an instrument of torture (Bentley and 

Trimer 1880, Bosland and Votava 2000, Center for 

New Crops and Plant Products 2002, 

Gardenguides.com 2002). 

References 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. Vol. 1. The MacMillan Co. New 

York, NY. 1,200 p. 

Bentley, R. and H. Trimer. 1880. Medicinal plants. 

Vol. 3. J.&A. Churchill, London. p. 147-227. 

Bosland, P.W. and E.J. Votava. 2000. Peppers: 

Vegetable and spice capsicums. CABI 

Publishing, Oxon, UK and New York. 204 p. 

Center for New Crops and Plant Products. 2002. 

Capsicum pepper. Purdue University. http:// 

hort.purdue.edu/newcrop/med-aro/factsheets/ 

CAPSICUM_PEPPER.html. 2 p.

El-Lakwah, F.A., O.M. Khaled, M.M. Khattab, 

and T.A. Abdel-Rahman. 1997. Toxic effects of 

extracts and powders of certain plants against 

the rice weevil (Sitophilus oryzae). Annals of 

Agricultural Science (Moshtohor) 35(1): 553-

566. 

Gardenguides.com. 2002. Cayenne. http://www. 

gardenguides.com/herbs/cayenne.htm. 4 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p.
149

Casearia decandra Jacq. tostado 

FLACOURTIACEAE 

Synonyms: Casearia parvifolia Jacq. 

 Casearia parvifolia Will. 

 Casearia albicaulis Risby 

Samyda lancifolia Sessé & Moç. 

John K. Francis 

 

General Description.—Tostado (a name used in 

Puerto Rico) is also known by wild honey-tree, 

wild-cherry, pipewood, biscuitwood, jumbie-apple, 

palo blanco, caracolillo, cerezo, corcho blanco, 

cotorrerillo, duro-duro, guía mansa, machacomo, 

tapaculo, limoncaspi, bois jaune, coco-ravet, jaune 

d’oeuf, and fortuga caspi. It is a shrub or small tree 

usually 2 to 6 m in height and 2 to 8 cm in diameter 

at breast height (d.b.h.). Tostado is supported by a 

strong taproot, somewhat finer lateral roots, and 

abundant near-surface fine roots. The roots are stiff 

and ivory-colored. The species usually has a single 

stem, unless disturbed, and smooth gray bark. The 

stemwood is moderately hard and moderately 

heavy. The branches and twigs are numerous, 

slender, and form a relatively dense crown. The 

alternate leaves have petioles 2 to 5 mm long, 

elliptic blades 3 to 9 cm long, with a finely sawtoothed margin. The small white, cream, or 

greenish-white flowers are born in nearly sessile 

clusters at the defoliate leaf nodes. The globose 

fruits are capsules about 8 to 10 mm in diameter, 

opening on three valves. The fruits are ripe when 

they turn from greenish-white to cream with a 

salmon-colored blush or light brown. The fleshy 

part of the fruit is orange to red. There are one to 

four, 4- to 5-mm seeds per fruit, depending on fruit 

size (Howard 1989, Liogier 1994, Little and 

Wadsworth 1964). 

Range.—The native range of tostado covers the 

islands of the West Indies from Hispaniola south 

and extends from Honduras through Panama and 

into South America as far south as Paraguay, 

Bolivia, and Northern Argentina (Howard 1989, 

Liogier 1994, Little and Wadsworth 1964). 

Ecology.—Tostado grows on well-drained soils 

ranging from coastal sands to upland clays, and on 

soils derived from sedimentary (including 

limestone), igneous, and metamorphic rocks. 

Annual precipitation in habitat in Puerto Rico 

ranges from about 900 mm to about 2200 mm. The 

species grows to altitudes of at least 1,120 m in 

Brazil (Silva and others 2002). In Bolivia it occurs 

from 230 to 650 m in elevation (Killeen and others 

1993), and in Puerto Rico, it grows from near sea 

level to over 600 m. Tostado is moderately 

intolerant of shade. It grows in disturbed areas, 

forest openings, very rocky sites, and the 

understories of lower density forests. In southern 

Brazil, it occurs in the third canopy layer of 

Araucaria forests (Silva and others 2002), and in 

Bolivia in Amazon forests and savanna woodlands 

(Killeen and others 1993). In Venezuela, the 

species functions as a gap-filling species in bush 

island savannas (San José and others 1991). 

Reproduction.—Tostado flowers and fruits 

irregularly throughout the year in Puerto Rico 

(Little and Wadsworth 1964), only a minority of 

plants being in flower or fruit at any time. It is 
150 

reported to flower from August to October in 

Bolivia (Killeen and others 1993). Plants 1 m or 

more in height in partial or full sunlight bear fruits. 

Shrubs and small trees bear from hundreds to 

thousands of fruits each year. A collection of fruits 

from Puerto Rico weighed an average of 0.340 +

0.031 g/fruit. They are extremely variable (CV = 

69.9 percent), even within the same plant. Seeds 

separated from the above collection weighed (airdried) an average of 0.0406 + 0.0005 g/seed or 

24,600 seeds/kg. Without pre-treatment, these 

seeds were sown on commercial potting mix and 

91 percent germinated 7 days later (author’s 

observation). Tostado sprouts when cut or burned. 

Growth and Management.—Although tostado is 

usually a shrub, if it survives several decades in 

fertile sites with sufficient sunlight, it may 

occasionally become a tree 12 to 18 m in height 

(Howard 1989, Liogier 1994). Weaver (1990) 

reported an average 5-year annual diameter 

increment of only 0.05 cm for plants 4 to 9 cm 

d.b.h. in St. John, U.S. Virgin Islands. The airdried specific gravity of stemwood was measured 

at 0.630 + 0.031. The above-ground average 

carbon content was measured at 0.508. Total 

above-ground dry weight in plants up to 5 cm 

d.b.h. can be estimated by the model: Wt = 

34.356(D2

S), where Wt is weight in grams, D is 

diameter in cm at 30 cm above the ground-line, S is 

total stem length in meters, and adjusted r-squared 

equals 0.977 (Francis 2000). 

Benefits.—The stems of tostado are sometimes 

used as fuel and fenceposts. This is an important 

honey plant. The small fruits are edible, but almost 

tasteless according to Little and Wadsworth (1964). 

However, the author found them sweet and 

pleasantly flavored. The species furnishes food and 

cover for wildlife, contributes to biodiversity, and 

helps stabilize the soil. 

References 

Francis, J.K. 2000. Estimating biomass and carbon 

content of saplings in Puerto Rican secondary 

forests. Caribbean Journal of Science 36(3-4): 

346-350. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Killeen, T., E. García E., and S.G. Beck. 1993. 

Guía de árboles de Bolivia. Herbario Nacional 

de Bolivia and the Missouri Botanical Garden. 

St. Louis, MO. 958 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

San José, J.J., M.R. Farinas, and J. Rosales. 1991. 

Spatial patterns of trees and structuring factors 

in a Trachypogon savanna of the Orinoco 

Llanos. Biotropica 23(2): 114-123. 

Silva, D.W., E. Seitz, M. Burgo, S.M. Silva, and 

J.J. Soares. 2002. Levantamento de espécies 

arbóreas no parque municipal das araucarias, 

Guapapuava, PR. Sociedad Botanica de São 

Paulo, Brazil. http://www.ib.usp.br/sbsp/ 

congresso/ec.htm. 1 p. 

Weaver, P.L. 1990. Tree diameter growth rates in 

Cinnamon Bay Watershed, St. John, U.S. Virgin 

Islands. Caribbean Journal of Science 26(1-2): 

1-6.
151

Casearia sylvestris Sw. wild-coffee 

FLACOURTIACEAE 

Synonyms: Samyda parviflora L. 

 Casearia parviflora L. 

 Anavinga samyda Gaertn. f. 

John K. Francis 

 

General Description.—Wild-coffee, also known 

as crack-open, cafeíllo, café silvestre, sarnilla, 

corta-lengua, guayabillo, guassatonga, and 

papelite, is a shrub or small tree usually 2 or 3 m in 

height, but occasionally reaching 10 m or more. On 

clay soils, the plant relies for absorption and 

support on extensive lateral roots that are white, 

moderately stiff, and have a corky bark. The plant 

may have a single or multiple stems with nearly 

smooth, thin gray bark and a moderately branchy 

habit. The sapwood is light brown and the 

heartwood is dark-brown, fine-textured, hard, 

heavy, and strong. The twigs are long and slender, 

often horizontal or drooping. The previous year’s 

growth often has tufts in the leaf axils of what were 

the fruit stalks. The alternate leaves have petioles 2 

to 8 mm long and ovate to lanciolate blades 4 to 13 

cm long by 2 to 5 cm broad with a wavy edge and a 

long-pointed tip. Tiny white, cream, or greenish 

flowers are crowded (20 to 50) on short stalks in 

the leaf axils. The flowers smell like a mixture of 

honey and urine. The fruits are globose or slightly 

flattened, 3 to 4 mm in diameter, splitting open on 

three valves to reveal usually three brown seeds 

covered with a red or red-orange aril (author’s 

observations, Croat 1978, Howard 1989, Liogier 

1994, Little and Wadsworth 1964). 

Range.—Wild-coffee is native to Cuba, Jamaica, 

Hispaniola, Puerto Rico, the Virgin Islands, the 

Lesser Antilles, Trinidad and Tobago, Mexico 

through Central America, and South America as far 

south as northern Argentina, Uruguay, and Bolivia 

(Howard 1989, Instituto de Botánica Darwinion 

2002, Killeen and others 1993, Little and 

Wadsworth 1964). 

Ecology.—Wild-coffee grows on soils of all 

textures derived from nearly all parent materials. 

However, it does not grow on swampy ground and 

usually not on excessively drained sites. In Puerto 

Rico, wild-coffee grows in areas receiving from 

about 900 to over 3000 mm of annual precipitation 

(author’s observation). However, in southern 

Brazil, wild-coffee grows even in annually flooded 

areas called várzeas (Universidad do Extremo Sul 

Catarinense 1999). In Nicaragua, the species grows 

from near sea level to 1,300 m in elevation 

(Stevens and others 2001). Wild-coffee 

demonstrates an intermediate tolerance to shade, 

being able to survive and grow slowly under a 

forest canopy that is not too dense but requiring 

partial sunlight to reproduce. Plants growing in 

pine plantations and natural secondary forests in 

Puerto Rico held similar concentrations of N, P, 

and ash in their leaves, but K levels were nearly 

twice as high in natural forest (Lugo 1992). Wildcoffee is common in brushy pastures, early 

secondary forests, roadsides, fence rows, and 

gallery forests of stream bottoms (author’s 

observation, Little and Wadsworth 1964). It is one 

of the principal arborescent species of the Llanos 

(plains) of the Orinoco area of Venezuela (van Os 

2000). 

Reproduction.—Under favorable conditions, wildcoffee is able to flower and fruit at a young age, 

about the second year as a sprout or at about 1 m in 
152 

height. Although individual plants do not flower 

continuously, members of the population may be 

seen flowering throughout the year (Little and 

Wadsworth 1964, Stevens and others 2001). In 

Panama, the species flowers principally in August 

to December with sporadic flowering during the 

dry season. The fruits mature in about 1 month 

(Croat 1978). A collection of fresh fruits (seed + 

aril) from Puerto Rico weighed an average of 

0.0286 + 0.0021 g/fruit. They vary considerably in 

size (coefficient of variation = 60.5). Air-dry seeds 

from the same collection averaged 0.0011 + 0.0001 

g/seed or 909,000 seeds/kg. Sown on peat, these 

seeds germinated at 31 percent between 20 and 103 

days after sowing (author’s observation). In 

southern Brazil, 84,000 seeds/kg were reported and 

a low rate of germination (Universidad do Extremo 

Sul Catarinense 1999). Germination is epigeal. Just 

one or two fruits per inflorescence are ripe at any 

time. The only method of seed collection known to 

the author is to hand-pick the tiny ripe fruits 

(capsules split exposing the red aril) and wet sieve 

to separate the seeds. Birds disburse the seeds. 

Natural seedlings are rare to common. Plants past 

the seedling stage have a high survival rate. 

Damaged plants coppice readily. 

Growth and Management.—Wild-coffee has a 

moderate growth rate and lives for at least 20 

years. Artificial reproduction is usually by seeds, 

but cuttings can also be rooted. Recommended 

spacing for plantings in Brazil is 4 by 4 m 

(Universidad do Extremo Sul Catarinense 1999). 

Because it becomes established as part of the early 

secondary forest or in disturbed openings of more 

advanced forest, management to promote 

regeneration of the species might include the use of 

clearcuts or group selection in closed forest and 

protection of open land from fires. 

Benefits.—The wood of wild-coffee is used for or 

is suitable for fuel, fence posts, stakes, small poles, 

rustic carpentry, and tool handles (Little and 

Wadsworth 1964). The species is recommended for 

urban planting as a food source for birds (Rio 

Grande Energia 2002). It is a honey plant (Little 

and Wadsworth 1964). Leaves of wild-coffee 

contains 2.5 percent essential oils with a pleasant 

aroma. They also contain caprionic acid, saponins, 

alkaloids, flavonoids, and a host of other 

chemicals. Extracts (particularly of the leaves) are 

used in traditional herbal medicine to treat a large 

number of different ailments (Universidad do 

Extremo Sul Catarinense 1999). Laboratory 

experiments have confirmed antiseptic, antitumor, 

antiulser, and abortive activity (Basile and others 

1990, Carvalho, and others 1999, Itokawa and 

others 1990, Silva and others 1988). A commercial 

perfume and a weight-loss product contain extracts 

of wild-coffee (Fontovit 2002, Multinível do Brasil 

S.A. 2002). 

References 

Basile, A.C., J.A.A. Sertie, S. Panizza, T.T. 

Oshiro, and C.A. Azzolini. 1990. 

Pharmacological assey of Casearia sylvestris. I. 

Preventive anti-ulsur activity and toxicity of the 

leaf crude extract. Journal of 

Ethnopharmacology 30(2): 185-197. 

Carvalho, J.C.T., V.V. Vignoli, G.H.G. de Souza, 

K. Ujikawa, J.J. Neto, and G.H.B. de Souza. 

1999. Antimicrobial activity of essential oils 

from plants used in Brazilian popular medicine. 

Acta Horticulturae 501: 77-81. 

Croat, T.B. 1978. Flora of Barro Colorado Island. 

Stanford University Press, Stanford, CA. 943 p. 

Fontovit. 2002. Porangaba Fontovit. 

http://fontovit.com.br/produtos/porangaba.htm. 

1 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Instituto de Botánica Darwinion. 2002. Catálogo 

de las plantas vasculares de la Argentina: 

Flacourtiaceae. http://www.darwin.edu.ar/ 

Catalogo/Flacourtiaceae.pdf. 3 p. 

Itokawa, H., N. Totsuka, H. Morita, K. Takeya, Y. 

Iitaka, E.P. Schenkel, and M. Motidome. 1990. 

New antitumor principles, casearines A-F, for 

Casearia sylvestris Sw. (Flacourtiaceae). 

Chemical and Pharmaceutical Bulletin 38(12): 

3,384-3,388. 

Killeen, T.J., E. García E., and S.G. Beck. 1993. 

Guía de árboles de Bolivia. Herberio Nacional 

de Bolivia and Missouri Botanical Garden, St. 

Louis, MO. 958 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 
153

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

Lugo, A.E. 1992. Comparison of tropical tree 

plantations with secondary forests of similar 

age. Ecological Monographs 62: 1-41. 

Multinível do Brasil S.A. 2002. Uma nova geração 

de colônias. http://www.multiniveldobrasil. 

com.br/bio_collection.html. 2 p. 

van Os, M. 2000. The Llanos de Orinoco. 

Wageningen Agricultural University, 

Wageningen, Netherlands. http://www.wau.nl/ 

rpv/ond_proj/dirjongm/Llanos/teksten/objectives

/chapter03.html. 8 p. 

Río Grande Energia. 2002. Manual de arborização 

e poda: espécies recomendadas. http://www.rgers.com.br/manual_poda/esp_recomendadas.asp. 

5 p. 

Silva, F.A., A.L.M. Baisch, B. Oliveira, A.M. 

Battastini, F. Torres, G. Rocoski, E.S. Silva, 

M.F. Alam, J.C.G. Apolinario, and A.J. Lapa. 

1988. Preliminary pharmacological studies on 

extracts from Casearia sylvestris Swartz. Acta 

Amazonica Suplemento 18(1-2): 219-229. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora of Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden Press. St. Louis, 

MO. p. 945-1,910. 

Universidad do Extremo Sul Catarinense. 1999. 

Jornada Catarinense de plantas medicinais: 

Guaçatonga. http://www.unesc.rct-sc.br/ 

plantas_medicinais/guaca.htm. 4 p. 
154 

Cassine xylocarpa Vent. marble tree 

CELASTRACEAE 

Synonyms: Elaeodendron xylocarpum (Vent.) DC. 

 Freziera dioica Macfad. 

 Elaeodendron dioicum (Macfad.) Griseb. 

 Elaeodendron attenuatum A. Rich. 

 Elaeodendron xylocarpum (Vent.) DC. 

 Cassine rotundata (DC.) Kuntze 

John K. Francis 

General Description.—Marble tree, also known as 

spoon tree, poison tree, wild nutmeg, coscorrón, 

aceituno, guayarote, mate prieto, pinipinche de 

sabana, bois-tan, bord-de-mer, and grosse-peau, is 

an evergreen shrub or small tree commonly 2 to 4 

m in height and 5 to 8 cm in diameter at breast 

height. The plant is supported by an extensive 

shallow and deep lateral root system with sinkers 

and sometimes a discernable taproot. The roots are 

orange and stiff. Marble tree often has several 

stems emanating from the root crown or low on the 

trunk. The crowns are fairly dense (especially 

when open-grown), branchy, and often contorted. 

The bark is gray, smooth to finely fissured, with 

inner bark that is purplish or pinkish. The twigs are 

slender and green, becoming gray with age. Leaves 

are variable, whitish-green to yellow-green, mostly 

opposite, but sometimes alternate, thick and stiff. 

They are entire or with a few teeth, elliptic to 

obovate, 2.5 to 12.5 cm long, with a rounded to 

spine-tipped apex. The inflorescences are tinyflowered, branched axillary cymes. The fruit is a 

yellow (at maturity), rounded drupe, 1.5 to 3 cm 

long with a thin, hard-fleshy exocarp and hard, 

thick-walled stone with one to three cells and 

elliptic seeds (Howard 1989, Liogier 1994, Little 

and others 1974). 

Range.—The range of marble tree consists of the 

Bahamas, the Greater and Lesser Antilles, Mexico, 

Belize, Costa Rica, Nicaragua, Panama, and 

Venezuela (Howard 1989, Instituto Nacional de 

Biodiversidad. 2000, Liogier 1994, Missouri 

Botanical Garden 2002a, 2002b). 

Ecology.—Marble tree is moderately intolerant of 

shade. It can persist and grow slowly under low 

forest but needs to be open-grown or at least have 

broken sunlight to flower and fruit. The species is 

drought-tolerant and usually grows on excessively 

well-drained situations—beaches, bluffs, rocky 

headlands, and rocky ridges. Marble tree often 

grows out of cracks in rocks. It is most often seen 

in areas of limestone and ultramafic (serpentine) 

rocks. The species is reported to be a nickel 

accumulator (Medina and others 1994). Colonized 

areas in Puerto Rico range from 750 to about 1800 

mm of mean annual rainfall and at elevations from 

near sea level to 300 m or more. The species may 

be found in remnant and late secondary forests. 

Reproduction.—Marble tree flowers and fruits 

intermittently throughout the year (Little and 

others 1974). A collection of air-dried seeds in 

Puerto Rico averaged 3.19 + 0.076 g/seed or 313 

seeds/kg. Sown in commercial potting mix, they 

began germinating in 26 days and completed 82 

percent germination. The species can also be 

propagated by asexual means. Fifty-six and 21 
155

percent, respectively, of air layers and misted 

cuttings treated with IBA rooted. The newly 

formed roots are fragile and easily broken off 

during transplanting (author’s observation). The 

seeds are probably dispersed by fruit bats. 

Seedlings and saplings are relatively common in 

and around Puerto Rican stands of the species. 

Growth and Management.—Marble tree grows 

slowly. A 1-m tall sapling with a basal diameter of 

1.5 cm growing in the understory of low basal area 

moist forest had 37 annual rings (author’s 

observation). Individuals as large as 20 cm in 

diameter at breast height (Little and others 1974) 

must be several decades old. Management of the 

species should include protection from fire and 

from commercial development of habitat. Existing 

understory seedlings and saplings could be 

released from competition by thinning or 

eliminating the overstory in patches. 

Benefits.—Marble tree is pretty enough to make a 

good ornamental but has rarely been used as such. 

It is an important component of the plant 

community in the harsh environment where it is 

most common and furnishes food and cover for 

animals. The fruits of marble tree are listed as a 

food item for the endangered Anegada iguana 

(Cyclura pinguis) (Zoological Society of San 

Diego 2002). The wood is light-brown, hard, 

heavy, fine-textured, strong, and durable (Little 

and others 1974). It is certainly useful for fuel and 

fence posts and possibly carving and turnery. The 

fruits are reported to be edible, and extracts of the 

plant are a stimulant (Liogier 1990). 

References

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Instituto Nacional de Biodiversidad. 2000. Leaps 

and bounds. The Cutting Edge 7(2): 1-2, 

http://inbio.ac.cr/papers/manual_plantas?apr00le

a.html. 2 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Edicones, 

Inc. San Juan, PR. 566 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

461 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 

Medina, E., E. Cuevas, J. Figueroa, and A.E. 

Lugo. 1994. Mineral content of leaves from 

trees growing on serpentine soils under 

contrasting rainfall regimes in Puerto Rico. Plant 

and Soil 158(1): 13-21. 

Missouri Botanical Garden. 2002a. Tropicos-Flora 

of Panama Checklist. http://mobot.org/cgibin/search_pick. 1 p. 

Missouri Botanical Garden. 2002b. W3

-Specimen 

data base. http://mobot.org/cgi-bin/search_vast. 

1 p. 

Zoological Society of San Diego. 2002. 

Unraveling the relationships between Caribbean 

iguanas and their ecosystems. http://www. 

sandiegozoo.org/conservation/fieldproject_ 

anegada_plant.html. 2 p. 
156 

Castela erecta Turp. cockspur 

SIMAROUBACEAE 

Synonyms: Castela nicholsoni Hook. 

 Castelaria micholsoni (Hook.) Small 

John K. Francis 

General Description.—Cockspur, also known as 

goat-bush, retama, and urupagüita, is an evergreen, 

spiny shrub 1 to 4 m in height and up to 10 cm on 

stem diameter. The plant is multi-stemmed and 

branchy. The twigs are stiff, sometimes zig-zag, 

whitish from fine hairs, and end in spines. There 

are also short spines at the leaf bases. The foliage 

is sometimes dense, composed of alternate simple 

oblong to elliptic, almost sessile leaves, 0.6 to 2.5 

cm long by 0.3 to 1.2 cm broad, dark green and 

glabrous above, and hairy below. The foliage and 

twigs are bitter. Flowers are tiny, whitish to red 

and tightly clustered in the leaf axils. The fruits are 

6- to 10-mm, red, fleshy drupes, one to four 

developing from a flower. Each fruit contains one 

hard seed (Howard 1988, Liogier 1988, Little and 

others 1974). 

Range.—Cockspur is native to Puerto Rico, the 

Virgin Islands, Antigua, Barbuda, Anguilla, 

Curacao, Aruba, northern Venezuela and northern 

Colombia (Howard 1988, Little and others 1974). 

It is not known to have been planted or naturalized 

elsewhere. 

Ecology.—Cockspur is a coastal species. It grows 

in beach strand vegetation, in sandy soils behind it, 

and on rocky escarpments and hills somewhat 

inland (Fundación La Salle de Ciencias Naturales 

2002), a dominant to minor part of local xeric 

scrub communities (Locklin 2002). It occurs to an 

elevation of about 100 m in Puerto Rico (Little and 

others 1974). These areas receive annual rainfall 

totals of about 700 to 900 mm. Soil type does not 

appear to be critical except that because of the 

species’ short stature and slow growth, it tends to 

be more successful in poor sites. Although 

cockspur tolerates salt spray and mild salt in the 

soil, it does not grow in very salty soils. 

Reproduction.—In Puerto Rico, cockspur flowers 

and fruits in the spring. The species bears male and 

female flowers on different plants (dioecious). 

Fleshy, red fruits are probably eaten and the seeds 

dispersed by birds. 

Growth and Management.—Cockspur is a slowgrowing species. No management experience has 

been published. 

Benefits.—Cockspur helps protect the soil and 

furnishes food and cover for wildlife. The sister 

species C. texana (T.&G.) Rose, once considered a 

part of cockspur as C. erectas subsp. texana (T. & 

G.) Cronq., is considered an important browse 

species (Secretaría de Medio Ambiente y Recursos 

Naturales. 2002). The common name, goat-bush, 

suggests that it is browsed by goats. 

 

References

Fundación La Salle de Ciencias Naturales. 2002. 

Playas de Aragua para uso turístico sustentable. 

http://www.fundacite.arg.gov.ve/papelesf/docs/ 

playas.pdf. 24 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 
157

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 

Locklin, C. 2002. Guajira-Barranquilla xeric scrub 

(NT1308). World Wildlife Fund, Washington, 

DC. http://www.worldwildlife.org/wildworld/ 

profiles/terrestrial/nt/nt1308_full.html. 7 p. 

Secretaría de Medio Ambiente y Recursos 

Naturales. 2002. Especies forestales no 

maderables y maderables no tradicionales de 

zonas áricas y semiáridas: Castela texana

(T.&G.) Rose. http://www.semarnat.gob.mx/ 

pfnm3/fichas/castela_texana.htm. 2 p.
158 

Ceanothus velutinus Dougl. ex Hook. snowbush ceanothus 

RHAMNACEAE 

Synonyms: none 

John K. Francis 

Drawing source: Britton and Brown 1913 

General Description.—Snowbush ceanothus is 

also known as shiny-leaf ceanothus, varnish-leaf 

ceanothus, sticky laurel, tobacco brush, mountain 

balm, buckbrush, deerbush, and snowbrush. It is 

an evergreen, upright to semiprostrate shrub 1 to 2 

m (rarely to 4 m) in height. The shrub has one to 

many stems with reddish-brown bark. The twigs 

are rigid, slightly flattened, and olive green. The 

plant is supported by a taproot and a deep, 

spreading root system. Simple, alternate leaves are 

shiny dark green above, and pale beneath leathery, 

and covered with a sticky resin. The 2- to 8-cm 

blades are broadly elliptical to ovate with rounded 

tips and rounded or subcordate bases. They have a 

fine serrate margin and three prominent veins 

proceeding from the base. The foliage is strongly 

aromatic, especially when crushed. Some have 

described it as being similar to cinnamon or 

balsam. Inflorescences, which are borne on short 

lateral branches on the previous year’s growth, 

contain many small white, five-maris, perfect 

flowers. Fruits are three-lobbed capsules 3 to 6 

mm long. The seeds are shiny and tan to dark 

brown. There are 2n = 24 chromosomes (Anderson 

2001, Banner 2002, Brayshaw 1996, Harrington 

1964, Nobs 1963, Pollock 2003, Welsh and others 

1987). 

Range.—The native range of snowbush ceanothus 

includes British Columbia and Alberta south to 

California and east to South Dakota and Colorado 

(Abrams 1951, Anderson 2001, Natural Resources 

Conservation Service 2003). There are two 

varieties: hookeri and velutinus. The former, of the 

Coast and Cascade ranges, in glaberous beneath 

the leaves. The latter, occurring farther inland, is 

finely hairy on the underside of the leaves 

(Brayshaw 1996). The species is not known to 

have naturalized outside its native range. 

Ecology.—Snowbush ceanothus is a member of a 

large number of plant associations (Anderson 

2001). It favors well-drained soils and is 

susceptible to root rot in more poorly drained 

conditions (Hansen 2002). These are moderately 

acid to neutral, often rocky or gravely soils, poor 

in nutrients, and of granitic parent material. The 

taproot penetrates 1.8 to 2.4 m deep (Pollock 

2003), and with the moisture-conserving leaf 

structure, impart the plant a great deal of drought 

resistance. Snowbush ceanothus grows in 

relatively low coastal habitat and in mountain sites 

up to 2,900 m in elevation (Welsh and others 

1987). Although found on all aspects, it is more 

likely to occur on southerly exposures (Anderson 

2001). Precipitation and length of growing season 

vary a great deal because of the wide elevational 

and latitudinal range of the species. Snowbush 

ceanothus is a pioneer species. Large numbers of 

seedlings often appear after logging and especially 

after fires. The species generally grows in small to 

large thickets, usually as the dominate species. It 

develops best in full sun but can persist in open 

forest stands. It grows more rapidly than conifer 

seedlings and will suppress them for a time. 

However, after 10 to 75 years, conifers overtop it, 

and its vigor and abundance declines (Pollock 

2003). 

Reproduction.—Snowbush ceanothus flowers 

from May through July and the fruits mature in 
159

August and September (Banner 2002). The flowers 

are pollinated by at least bees (Clark 1976). There 

is an average of 207,000 seeds/kg. When seeds 

mature, beginning in August, they are ejected from 

the pod and dispersed a short distance (Anderson 

2001). Good seed production begins when plants 

are about 8 years old. As many as 1,250,000 

seeds/ha/year can be produced. However, as much 

as 99 percent of the annual seed crop can be 

consumed by ants, birds, and rodents. (Pollock 

2003). Seeds still accumulate in large numbers in 

the soil seed bank and can apparently remain 

viable for 200 to 300 years until conditions 

become favorable. Germination is epigeal (Reed 

1974). Heat, as from forest fires, appears to scarify 

the seeds. Burned or damaged plants sprout from 

the root crown. Layering also occurs when 

branches come in contact with the soil (Anderson 

2001). 

Growth and Management.—The natural life 

span of snowbush ceanothus is longer than 25 

years, and plants sometimes reach 50 years old. 

However, stands sometimes begin to deteriorate 

after 15 years (Anderson 2001). A 17-year-old 

stand in the Cascade Mountains in Oregon 

contained over 4,000 stems/ha and averaged 

34,000 kg/ha of above-ground biomass (Pollock 

2003). Seeds may be sown in the late summer and 

will germinate in the spring. Pretreatments are 

necessary for good germination of spring-planted 

seeds. A hot water soak (90 °C until the water 

cools) followed by cold stratification at 1 to 5 °C 

for 30 to 90 days. Seeds should be sown in flats at 

depths of twice the diameter of the seed in 

amended mineral soil. Germination may be as high 

as 82 percent. Seedlings should be pricked into 

pots when they have gained several sets of true 

leaves (Reed 1974). Plants can be propagated with 

stem cuttings. Survival of transplanted seedlings is 

low (about 9 percent). Small seedlings survive 

better than large ones (Pollock 2003). Snowbush 

ceanothus stands are difficult to control. Broadleaf 

herbicides such as 2,4-D tend to top-kill rather 

than eliminate it. Opening the brush canopy also 

allows seeds to germinate and grow. Glyphosate 

and hand slashing result in greater densities of the 

species in the long run (Anderson 2001). 

Benefits.—Snowbush ceanothus adds to the 

beauty of the forest, helps protect the soil, 

furnishes food and cover for wildlife, and is a 

useful ornamental and medicinal plant. The 

species is little used by cattle and horses. Sheep 

browse it only when other forage is unavailable. 

Deer and elk browse snowbush ceanothus yearround, although it makes up only a small portion 

of their diet. It is important fall and winter moose 

browse in some areas. It has an in vitro dry matter 

digestibility of about 57 percent and a crude 

protein content of 7 to 18 percent (Anderson 

2001). Snowbush ceanothus is important resting 

and escape cover for many types of wildlife 

(Pollock 2003). Nitrogen is fixed by actinomycetes 

of the genus Frankia in root nodules (Dalton 

1997). Snowbush ceanothus makes a beautiful and 

hardy ornamental that can be pruned and shaped 

(Hansen 2002). Native Americans used the leaves 

as a deodorant, smoke of the plant to kill bedbugs, 

and decoctions to treat flu, pain, and gonorrhea 

(Moerman 1986). They also used the leaves as a 

tobacco substitute (Pollock 2003). 

References

Abrams, L. 1951. Illustrated flora of the Pacific 

States. Vol. 3. Stanford University Press, 

Stanford, CA. 866 p. 

Anderson, M.D. 2001. Ceanothus velutinus. In: 

U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shrub/ 

ceavel/all.html. 73 p. 

Banner, R. 2002. Deerbush. Utah State University, 

Logan, UT. http://extension.usu.edu/coop/ 

natres/range/Woody/deerbush.htm. 4 p. 

Brayshaw, T.C. 1996. Trees and shrubs of British 

Columbia. UBC Press, Vancouver, British 

Colunbia, Canada. 373 p. 

Britton, N.L. and A. Brown. 1913. Illustrated flora 

of the northern states, Canada, and the British 

possessions. Vol. 2, 2nd ed. Scribner, New York. 

735 p. 

Clark, L.J. 1976. Wild flowers of the Pacific 

Northwest. Gray’s Publishing Limited, Sidney, 

British Columbia, Canada. 604 p. 

Dalton, D. 1997. Nitrogen fixation: range of 

organisms that can fix nitrogen. Reed College, 

Portland, OR. http://academic.reed.edu/ 

biology/Nitrogen/Nfix1.html. 7 p. 

Hansen, W. 2002. Ceanothus. http://www. 

nwplants.com/plants/shrubs/ceanothus/. 6 p. 
160 

Harrington, H.D. 1964. Manual of the plants of 

Colorado. Sage Books, Denver, CO. 666 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports 19. University of 

Michigan, Museum of Anthropology, Ann 

Arbor, MI. 524 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Ceanothus velutinus Dougl. ex 

Hook. http://plants.usda.gov/cgi_bin/ 

plant_profile.cgi?symbol=CEVE. 4 p. 

Nobs, M.A. 1963. Experimental studies on species 

relationships in Ceanothus. Publication 623. 

Carnegie Institution of Washington, 

Washington, DC. 94 p. 

Pollock, T. 2003. Ceanothus velutinus. University 

of University of Saskatchewan, Saskatoon, 

Saskatchewan, Canada. http: //www.usask.ca/ 

agriculture/plantsci/classes/range/ceanothus/ 

html. 5 p. 

Reed, M.J. 1974. Ceanothus L., ceanothus. In: 

C.S. Schopmeyer, tech. coord. Seeds of woody 

plants in the United States. Agriculture 

Handbook 450. U.S. Department of Agriculture, 

Forest Service, Washington, DC. p. 284-290. 

Welsh, S.L., N.D. Atwood, S. Gooddrich, and L.C. 

Higgins. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo UT. 894 p. 

 
161

Celastrus orbiculatus Thunb. oriental bittersweet 

CELASTRACEAE 

Synonyms: Celastrus articulatus Thunb.

 Celastrus insularis Koidz. 

 Celastrus jeholensis Nakai 

 Celastrus lancifolius Nakai 

 Celastrus stephanotiifolius Makino 

 Celastrus strigulosus Nakai 

 Celastrus tatarinowii Rupr. 

 Celastrus versicolor Nakai 

Jinshuang Ma and Gerry Moore 

Photo credit: Steven Clemants 

General Description.—Oriental bittersweet is the 

name used most often in North America to refer to 

this nonnative vine. Other common names include 

Asian or Asiatic bittersweet and round-leaved 

bittersweet. The woody roots can be up to 2 cm 

thick and are yellow to orange. Oriental 

bittersweet is multistemmed with brown, round 

twigs; the pith is solid and white. It can behave 

like a shrub but it is usually a climbing woody vine 

that can reach up to 20 m in height. Oriental 

bittersweet is able to climb and effectively cover 

other woody vegetation. The plant lacks tendrils or 

aerial roots. The leaves of oriental bittersweet are 

deciduous, alternate, spiral, stipulate, and measure 

(including the petiole) 2.8 to 13.7 by 2.9 to 7.6 cm, 

with the blades suborbicular to broadly obovate in 

outline with crenulate to serrulate margins. The 

leaf blades can have white hairs along the midvein 

on the underside but are otherwise glabrous; the 

lateral veins are ascending. The leaf scars are flush 

with the stem, not raised. The small, greenishwhite, five-parted flowers are in axillary cymes of 

three or four functionally unisexual flowers, with 

the partially fused sepals 3 to 7 by 1 to 2 mm and 

the petals 2.6 to 5.0 by 0.9 to 2.0 mm. The 

functional male flowers have five stamens about as 

long as the petals and a vestigial pistil. The 

functional female flowers have a syncarpous pistil 

with a three-lobed stigma, short columnar style, 

and well-developed superior ovary. The fruit of 

Oriental bittersweet is a bright orange, threevalved, globose capsule, with each valve covering 

one or two seeds that are enclosed in a fleshy, 

crimson aril. The seeds (sans aril) are smooth, light 

orange, oblong, 2.5 to 2.6 mm by 1.5 to 1.6 mm 

(Clemants 2003). The genus name is derived from 

the ancient Greek name Celastros (Kelastros), 

which in Greek was applied to a different 

Mediteranean evergreen tree probably in the genus 

now known as Phillyrea (Rehder 1940). 

Range.—Oriental bittersweet is native to east Asia 

in Japan, Korea, and China, its southern limit 

occurring along the Yangtze River watershed in 

China. In its native range it occurs in areas with 

elevation ranges from 450 to 2200 m (Cheng and 

Huang 1999). The plant has become widely 

established in the Eastern United States, occurring 

in 25 states (Shetler and Orli 2000, USDA 2003). 

It is also known in Canada from Ontario and 

Quebec (Scoggan 1978). 

Systematic Botany.—Oriental bittersweet is 

similar to American bittersweet (Celastrus 

scandens L.), and can be distinguished from it on 

the basis of its leaves and inflorescences (Hou 

1955). The leaves of oriental bittersweet are 

suborbicular to obovate and are usually less than 

twice as long as wide, whereas the leaves of 

American bittersweet are elliptic to oblong and are 

usually twice as long as wide. The flowers (and 

fruits) of oriental bittersweet are in short axillary 

clusters; the flowers (and fruits) of American 
162 

bittersweet are in long terminal clusters. Both 

species have the same chromosome number (n = 

23; Bowden 1945) and they can be hybridized 

(White and Bowden 1947, Pooler and others 

2002). The presence of some intermediate material 

in the wild with both axillary and terminal 

inflorescences has led to the speculation that the 

two species may hybridize in the wild (Dreyer and 

others 1987, Mehrhoff in Pooler and others 2002). 

Some spell the specific epithet “orbiculata” instead 

of “orbiculatus” used here. This is because 

botanists disagree as to whether the genus name 

Celastrus L. should be treated as feminine 

(“orbiculata”) or masculine (“orbiculatus”) (Paclt 

1998). In classical Greek, the generic name was 

treated as feminine; however, Linnaeus treated the 

genus as masculine. The matter is now being 

reviewed by a formal nomenclatural committee 

(Brummitt 2000). 

Ecology.—Oriental bittersweet grows in a wide 

variety of habitats, including, dune thickets, 

fencerows, forests, forest edges, and roadsides. It 

seems to do particularly well in disturbed habitats 

(Dreyer 1994, Steward and others in press). This 

species was perhaps first introduced in the New 

World during the 1860s and was later popularized 

for ornamental plantings (Patterson 1973, Rehder 

1940). Oriental bittersweet is an aggressive 

invader that can dominate all vegetation levels of 

forested and open areas (Bergmann 2003). It 

grows over other vegetation, completely covering 

it. It can kill other plants by preventing them from 

photosynthesizing and through girdling and 

uprooting. Based on its native range and habitat 

types, Oriental bittersweet can be expected to 

spread to additional areas in the United States and 

Canada (J. Ma, personal observation). Oriental 

bittersweet is a shade tolerant species, able to 

acclimate to low light levels under heavy shading 

and grow explosively if shading is reduced. Once 

large clones are established, it is difficult to 

eliminate since it has developed such an extensive 

root system. While oriental bittersweet is 

spreading and increasing in abundance, the native 

American bittersweet is concurrently declining 

(Dreyer and others 1987, Forman and Stark 2001, 

Steward and others in press). Pooler and others 

(2002) have hypothesized that the spread of 

Oriental bittersweet and its hybridization with 

American bittersweet may be threatening the 

genetic identity of the American bittersweet. The 

very properties–faster growth (Bailey 1922, 

Rehder 1940), greater fecundity (Clemant and 

others 1991, Dreyer and others 1987, Hart 1928), 

greater tolerance of environmental heterogeneity 

(Baker 1974, Newsome and Noble 1986, Sakai 

and others 2001)–that made early horticulturists 

prefer oriental bittersweet over American 

bittersweet (C. scandens L.) are the same 

characteristics that enable oriental bittersweet to be 

an extremely successful invasive plant. 

Reproduction.—Oriental bittersweet blooms in 

the spring and is pollinated by hymenopterous 

insects, especially bees, although wind may also 

be involved (Brizicky 1964). The fruit ripens in 

the fall. The seeds are dispersed by birds and small 

mammals (Dreyer 1994). Humans are also 

important dispersal agents since the fruiting stems 

are commonly used in dried flower arrangements 

and subsequently discarded in compost and brush 

piles (Dreyer 1994). Oriental bittersweet can also 

reproduce asexually through root suckering. 

Growth and Management.—Due to the plant’s 

highly invasive nature, it is not recommended that 

this plant be grown. Small infestations can be 

controlled by cutting the stems and removing the 

roots. Larger infestations require the application of 

glycophosphate herbicides (Miller 2002). 

References 

Bailey, L.H. 1922. The standard cyclopedia of 

horticulture. vol 2. Macmillan Co., New York. 

1,200 p 

Baker, H.G. 1974. The evolution of weeds. Annual 

Review of Ecology and Systematics 5: 1-24. 

Bergmann, C. 2003. Oriental Bittersweet Celastrus 

orbiculatus Thunb. http://www.nps. 

gov/plants/alien/fact/ceor1.htm. [not paged]. 

Bowden, W.M. 1945. A list of chromosome 

numbers in higher plants I. Acanthaceae to 

Myrtaceae. American Journal of Botany 32: 81-

92. 

Brizicky, G.K. 1964. The genera of Celastrales in 

the southeastern United States. Journal of the 

Arnold Arboretum 45: 206-235. 

Brummitt, R.K. 2000. Report of the Committee for 

Spermatophyta: 49. Taxon 49: 261-278. 

Cheng, C.Y., and P.H. Huang. 1999. Celastraceae. 

Flora Reipublicae Popularis Sinicae 45: 1-218 

(in Chinese). 
163

Clemant, C., R. Warren, G. Dreyer, and P. Barnes. 

1991. Photosynthesis, water relations, and 

fecundity in the woody vines American and 

Oriental bittersweet (Celastrus scandens and 

Celastrus orbiculatus). American Journal of 

Botany 78 (Suppl.): 134-135. 

Clemants, S.E. 2003. Celastrus orbiculatus. New 

York Metropolitan Flora Project. Brooklyn 

Botanic Garden http://www.bbg.org/sci/nymf/ 

encyclopedia/cel/cel0020b.htm#description. [not 

paged]. 

Dreyer, G.D. 1994. Element stewardship abstract 

for Celastrus orbiculata Asiatic bittersweet. The 

Nature Conservancy, Arlington, Virginia. 

http://tncweeds.ucdavis.edu/esadocs/documnts/ 

celaorb.html.11p. 

Dreyer, G.D., L.M. Baird, and C. Fickler. 1987. 

Celastrus scandens and Celastrus orbiculatus: 

comparisons of reproductive potential between a 

native and an introduced woody vine. Bulletin of 

the Torrey Botanical Club 114: 260-264. 

Forman, J., and J. Stark. 2001. The introduction of 

non-native plants into Massachusetts. 

http://omega.cc.umb.edu/~conne/jennjim/ 

celastrus.html. [not paged]. 

Hart, H.T. 1928. Delayed germination in seedsof 

Peltandra virginica and Celastrus scandens. 

Publications of the Puget Sound Biological 

Station of the University of Washington 6: 255-

261. 

Hou, D. 1955. A revision of the genus Celastrus. 

Annals of the Missouri Botanical Garden 42: 

215-302. 

Miller, J.H. 2002. Exotic pest plants and their 

control. The Bugwood Network. 

http://www.bugwood.org/weeds/forestexotics.ht

ml. Accessed 30 June 2003. [not paged]. 

Newsome, A.E. and I.R. Noble. 1986. Ecological 

and physiological characters of invading species. 

In: R.H. Groves and J.J. Burdon, eds. Ecology of 

biological invasions. Cambridge University 

Press. p. 1-20. 

Paclt, J. 1998. Proposal to conserve the name 

Celastrus (Celastraceae) as being of feminine 

gender. Taxon 47: 879-880. 

Patterson D. 1973 Distribution of Oriental 

bittersweet in the United States. Journal of the 

Elisha Mitchell Science Society 89: 245. 

Pooler, M.R., R.L. Dix, and J. Feely. 2002. 

Interspecific hybridization between the native 

bittersweet, Celastrus scandens and the 

introduced invasive species, C. orbiculatus. 

Southeastern Naturalist 1: 69-76. 

Rehder, A. 1940. Manual of cultivated trees and 

shrubs. Macmillan Publishing Co., New York. 

996 p. 

Saikai, A.K., F.W. Allendorf, J.S. Holt, D.M. 

Lodge, J. Molofsky, K.A. With, S. Baughman, 

R.J. Cabin, J.E. Cohen, N.C. Ellstrand, D.E. 

McCauley, P.O’Neil, I.M. Parker, J.N. 

Thompson, and S.G. Weller. 2001. The 

population biology of invasive species. Annual 

Review of Ecology and Systematics 32: 305-

332. 

Scoggan, H.J. 1978. The Flora of Canada Part 3 – 

Dicotyledoneae (Saururaceae to Violaceae). 

National Museum of Natural Sciences, National 

Museums of Canada, Ottawa. 568 p. 

Shetler, S.G., and S.S. Orli. 2000. Annotated 

checklist of the vascular plants of the 

Washington-Baltimore area. Part. 1. Ferns, fern 

allies, gymnosperms, dicotyledons. Department 

of Systematic Biology – Botany. National 

Museum of Natural History, Smithsonian 

Institution, Washington, D.C. 186 p. 

Steward, A.M., S.E. Clemants, and G. Moore 

2003. The concurrent decline of the native 

Celastrus scandens and spread of the non- native 

Celastrus scandens in the New York City 

metropolitan area. Journal of the Torrey 

Botanical Society 130: 143-146. 

USDA, NRCS. 2003. The PLANTS Database. 

National Plant Data Center, Baton Rouge, 

LA. http:plants.usda.gov. [not paged]. 

White, O.E. and W.M. Bowden 1947. Oriental and 

American bittersweet hybrids. Journal of 

Heredity 38: 125-127. 
164 

Celastrus scandens L. American bittersweet 

CELASTRACEAE 

Synonyms: Celastrus bullata L. 

 Euonymoides scandens (L.) Medic. 

Jinshuang Ma and Gerry Moore 

Illustration source: Britton and Brown 1897 

General Description.—American bittersweet is 

the name used most often in North America to 

refer to this native vine. Other common names 

include climbing orange root, false bittersweet, 

fever twig, staff-tree, waxwork, and yellow-root 

(Dillingham 1907). The woody roots can be up to 

2 cm thick and are yellow to orange. American 

bittersweet is multistemmed with brown, round 

twigs; the pith is solid and white. It can behave 

like a shrub but it is usually a climbing woody vine 

that can reach up to 10 m in height. The plant lacks 

tendrils or aerial roots. The leaves of American 

bittersweet are deciduous, glabrous, alternate, 

spiral, stipulate, and measure (including the 

petiole) 4.5 to 14.8 by 2.7 to 5.5 cm, with the 

blades elliptic to ovate in outline with serrulate 

margins. The lateral veins are ascending. The leaf 

scars are flush with the stem. The small, greenishwhite, five-parted flowers are in terminal racemes 

of 14 to 50 functionally unisexual flowers, with 

the partially fused sepals 3 to 7 by 1 to 2 mm and 

the petals 1.8 to 4.0 by 0.9 to 1.2 mm. The 

functional male flowers have five stamens about as 

long as the petals and a vestigial pistil. The 

functional female flowers have a syncarpous pistil 

with a three-lobed stigma, short columnar style, 

and well-developed superior ovary. The fruit of the 

American bittersweet is a bright orange, threevalved, globose to ellipsoid capsule, with each 

valve covering one or two seeds that are enclosed 

in a fleshy, crimson aril. The seeds (sans aril) are 

smooth, light orange, ellipsoid, 3.5 to 4.5 by 1.8 to 

2.0 mm (Clemants 2003). The genus name is 

derived from the ancient Greek name Celastros 

(Kelastros), which in Greek was applied to a 

Mediterranean evergreen tree probably in the 

genus now known as Phillyrea (Rehder 1940). 

Range.—American bittersweet has been reported 

throughout most of the United States, except 

Florida and the Western States of Arizona, 

California, Colorado, Idaho, Nevada, New 

Mexico, Oregon, Utah and Washington (USDA. 

2003). It is also known in Canada from Manitoba, 

New Brunswick and Quebec (Scoggan 1978). 

Reports (Bailey 1949, Britton and Brown 1897) of 

this plant from New Mexico (then a territory with 

the same area as the present state) are probably in 

error. No voucher specimens exist for these reports 

and there has been no other report of American 

bittersweet from New Mexico. 

Systematic Botany.—American bittersweet is 

similar to the Oriental bittersweet (Celastrus 

orbiculatus Thunb.) and can be distinguished from 

it on the basis of its leaves and inflorescences 

(Hou 1955). The leaves of American bittersweet 

are elliptic to oblong and are usually twice as long 

as wide, whereas the leaves of Oriental bittersweet 

are suborbicular to obovate and are usually less 

than twice as long as wide. The flowers (and 

fruits) of American bittersweet are in long terminal 

clusters; the flowers (and fruits) of Oriental 

bittersweet are in short axillary clusters. Both 

species have the same chromosome number (n = 
165

23, Bowden 1945) and they have been hybridized 

(White and Bowden 1947, Pooler and others 

2002). The presence of some intermediate material 

in the wild with both lateral and terminal 

inflorescences has led to the speculation that the 

two species may hybridize in the wild (Dreyer and 

others 1987, Mehrhoff in Pooler and others 2002). 

Ecology.—American bittersweet grows in a wide 

variety of habitats, including dune thickets, 

fencerows, forests, forest edges, and roadsides. 

The native American bittersweet is declining, 

while the nonnative Oriental bittersweet (Celastrus 

orbiculatus Thunb.) is spreading and increasing in 

abundance (Dreyer and others 1987, Forman and 

Stark 2001, Steward and others in press). Pooler 

and others (2002) have hypothesized that the 

spread of Oriental bittersweet and its hybridization 

with American bittersweet may be threatening the 

genetic identity of the American bittersweet. The 

very properties--faster growth (Bailey 1922, 

Rehder 1940), greater fecundity (Clemant and 

others 1991, Dreyer and others 1987, Hart 1928), 

greater tolerance of environmental heterogeneity 

(Baker 1974, Newsome and Noble 1986, Sakai 

and others 2001)--that made early horticulturists 

prefer Oriental bittersweet over American 

bittersweet are the same characteristics that enable 

Oriental bittersweet to outcompete American 

bittersweet. 

Reproduction.—American bittersweet blooms in 

the spring and is pollinated by hymenopterous 

insects, especially bees, although wind may also 

be involved (Brizicky 1964). The fruit ripens in 

the fall. The seeds are dispersed by birds and small 

mammals (Dreyer 1994). American bittersweet 

can also reproduce asexually through root 

suckering. 

Growth and Management.—American 

bittersweet does well in many conditions. It is a 

rapid grower, adding up to several meters in one 

growing season. Adequate sun is important for 

fruit production. It can be planted in Hardiness 

Zones 3 to 8, in soils with pH ranges from 3.7 to 

6.5. It cannot tolerate saturated soils. Propagation 

can be effected either through layering or seed 

(Chittenden and Synge 1956). Seeds should be 

stratified in the dark at about 4 °C for 3 months 

(Young and Young 1992) 

Benefits.—The branches of American bittersweet 

were used by Native Americans, including the 

Chippewa, as a last resort food source (Dillingham 

1907, Palmer 1871). The leaves have been 

reported as being poisonous to cattle (Brizicky 

1964). Fruiting stems of the species are often used 

in dried arrangements. 

References 

Bailey, L.H. 1922. The standard cyclopedia of 

horticulture Vol 2. Macmillan Co., New York. 

1,200 p. 

Bailey, L.H. 1949. Manual of cultivated 

plants.Macmillan Co., New York. 1,116 p. 

Baker, H.G. 1974. The evolution of weeds. Annual 

Review of Ecology and Systematics 5: 1-24. 

Bowden, W.M. 1945. A list of chromosome 

numbers in higher plants I. Acanthaceae to 

Myrtaceae. American Journal of Botany 32: 

 81-92. 

Britton, N. L. and A. Brown 1897. An illustrated 

flora of the northern United States, Canada, and 

the British possessions Vol 2. Charles Scribner’s 

Sons, New York. 643 p. 

Brizicky, G.K. 1964. The genera of Celastrales in 

the southeastern United States. Journal of the 

Arnold Arboretum 45: 206-235. 

Chittenden, F.J. and P.M. Synge. 1956. Dictionary 

of gardening. Vol. 1, 2nd Ed. Clarendon Press, 

Oxford, UK. 512 p. 

Clemant, C., R. Warren, G. Dreyer, and P. Barnes. 

1991. Photosynthesis, water relations, and 

fecundity in the woody vines American and 

Oriental bittersweet (Celastrus scandens and 

Celastrus orbiculatus). American Journal of 

Botany 78 (Suppl.): 134-135. 

Clemants, S.E. 2003. Celastrus orbiculatus. New 

York Metropolitan Flora Project. Brooklyn 

Botanic Garden http://www.bbg.org/sci/nymf/ 

encyclopedia/cel/cel0030b.htm#description. [not 

paged]. 

Dillingham, F.T. 1907. The staff-tree, Celastrus 

scandens, as a former food supply of starving 

Indians. American Naturalist 41: 391-393. 

Dreyer, G.D. 1994. Element stewardship abstract 

for Celastrus orbiculata Asiatic bittersweet. The 

Nature Conservancy, Arlington, Virginia. 

http://tncweeds.ucdavis.edu/esadocs/documnts/c

elaorb.html. 11p. 
166 

Dreyer, G.D., L.M. Baird, and C. Fickler, 1987. 

Celastrus scandens and Celastrus orbiculatus: 

comparisons of reproductive potential between a 

native and an introduced woody vine. Bulletin of 

the Torrey Botanical Club 114: 260-264. 

Forman, J. and J. Stark. 2001. The introduction of 

non-native plants into Massachusetts. http:// 

omega.cc.umb.edu/~conne/jennjim/celastrus. 

html. [not paged]. 

Hart, H.T. 1928. Delayed germination in seeds of 

Peltandra virginica and Celastrus scandens. 

publications of the Puget Sound Biological 

Station of the University of Washington 6: 255-

261. 

Hou, D. 1955. A revision of the genus Celastrus. 

Annals of the Missouri Botanical Garden 42: 

215-302. 

Newsome, A.E. and I.R. Noble. 1986. Ecological 

and physiological characters of invading species. 

In: R.H. Groves and J.J. Burdon, eds. Ecology of 

biological invasions. Cambridge University 

Press, Cambridge, MS. p. 1-20. 

Palmer, E. 1871. Food products of the North 

American Indians. In H. Capron, Report of the 

Commissioner of Agriculture for the Year 1870, 

Washinton, D.C. p. 404-428 

Pooler, M.R., R.L. Dix, and J. Feely. 2002. 

Interspecific hybridization between the native 

bittersweet, Celastrus scandens and the 

introduced invasive species, C. orbiculatus. 

Southeastern Naturalist 1: 69-76. 

Rehder, A. 1940. Manual of cultivated trees and 

shrubs. Macmillan Publishing Co., New York. 

996 p. 

Saikai, A.K., F.W. Allendorf, J.S. Holt, D.M. 

Lodge, J. Molofsky, K.A. With, S. Baughman, 

R.J. Cabin, J.E. Cohen, N.C. Ellstrand, D.E. 

McCauley, P.O’Neil, I.M. Parker, J.N. 

Thompson, and S.G. Weller. 2001. The 

population biology of invasive species. Annual 

Review of Ecology and Systematics 32: 305-

332. 

Scoggan, H.J. 1978. The Flora of Canada Part 3 – 

Dicotyledoneae (Saururaceae to Violaceae). 

National Museum of Natural Sciences, National 

Museums of Canada, Ottawa. 568 p. 

Steward, A.M., S.E. Clemants, and G. Moore 

2003. The concurrent decline of the native 

Celastrus scandens and spread of the non- native 

Celastrus scandens in the New York City 

Metropolitan area. Journal of the Torrey 

Botanical Society 130: 143-146. 

USDA, NRCS. 2003. The PLANTS Database, 

Baton Rouge, LA. http:plants.usda.gov. [not 

paged]. 

White, O.E. and W.M. Bowden 1947. Oriental and 

American bittersweet hybrids. Journal of 

Heredity 38: 125-127. 

Young, J.A. and C.G. Young. 1992. Seeds of the 

woody plants of North America. Dioscorides 

Press, Portland, OR. 407 p. 
167

Celtis reticulata Torr. netleaf hackberry 

ULMACEAE 

Synonyms: Celtis douglasii Planch.

C. occidentalis L. var. reticulata (Torr.) Sarg.

C. laevigata Willd. var. reticulata (Torr.) L. Benson 

Ann M. DeBolt 

Illustration credit: J.R. Janish in Hitchcock and 

others 1964. 

General Description.—Netleaf hackberry is also 

known as western or Douglas hackberry. It is a 

deciduous shrub to small tree from 3 to 20 m tall. 

Near the northern edge of its range in Idaho, its 

maximum height is 12 m (DeBolt and McCune 

1995). This species’ growth form varies from a 

singular, reasonably straight-stemmed tree to a 

gnarled shrub with multiple stems. The bark is 

gray, hard, and moderately thick, forming 

protruding corky ridges with age (Elias 1980). 

Leaves are 3 to 9 cm long, alternate, simple, ovate 

to ovate lanceolate, and somewhat leathery with 

scabrous surfaces and entire to serrate margins. 

The specific epithet reticulata comes from the 

reticulate or netlike pattern of prominent venation 

on the leaf underside (Lanner 1983). Twigs and 

leaves are frequently infested by insect galls, but 

vigor is not seriously affected (Hayward 1948). 

The small, green, apetalous, polygamomonoecious flowers number from one to several in 

the leaf axils. Pea-sized fruits are fleshy, reddishorange to purple globose drupes, each with a 

single large seed. Some authorities consider the 

highly variable netleaf hackberry to be a variety of 

sugarberry (C. laevigata Willd.) (Natural 

Resources Conservation Service 2003, Stephens 

1973). Others recognize many intergrading forms, 

and it is possible that future distribution maps of 

C. reticulata, if the name is retained, will not 

resemble those available today. 

Range.—Netleaf hackberry is found east of the 

Cascade and Sierra Nevada Mountains from 

central Washington south to northern Mexico and 

east to central Kansas, Texas, and Oklahoma 

(Lanner 1983, Little 1976). 

Ecology.—At elevations ranging from 100 to 

2,000 m, netleaf hackberry grows in a variety of 

habitats including mountain shrub, deciduous 

riparian woodlands, live oak-mixed shrub and 

wash scrub communities, rocky ravines, and as 

scattered individuals in semi-desert grasslands, 

pinyon-juniper, and Joshua tree woodlands (Albee 

and others 1988, Brown 1982, Carmichael and 

others 1978, Plummer 1977). Populations can be 

small or highly localized, particularly at the 

northern latitudes (Daubenmire 1970), or they can 

comprise the dominant vegetation matrix. On the 

Edwards Plateau of Texas, netleaf hackberry 

grows as an overstory codominant in many 

riparian woodlands (Van Auken and others 1979). 

A hackberry association was presented by DickPeddie and Hubbard (1977) for the mixed 

deciduous series of their riparian classification 

scheme for New Mexico, and several ecologists 

have recognized it as a vegetation type on the 

lower Snake and Salmon Rivers in Idaho and 

Oregon (Huschle 1975, Tisdale 1986), where 

warm canyons are adequate for its existence. At 
168 

northern latitudes stands are often associated with 

rock (DeBolt and McCune 1995), which provides 

extra moisture and some protection from wildfire. 

Reproduction.—Wind-pollinated, nonshowy 

flowers appear in spring, just before or as the 

leaves unfold, depending on latitude. Fruits ripen 

from September to November and persist on the 

trees until midwinter. They are dispersed by birds, 

rodents, and other small mammals. Netleaf 

hackberry exhibits low germination percentages 

(37 percent) and high seed dormancy (Bonner 

1974). Seeds are hard and thick-walled, probably 

the major impediment to rapid germination. 

Germination is epigeal. Dormancy can be 

overcome with stratification at 5 o

C in moist sand 

or other suitable media for 120 days (Bonner 

1974). Mechanical scarification enhances 

germination after 45 days (DeBolt 1992). Netleaf 

hackberry can reproduce vegetatively by sprouting 

from the root crown, stem base, or lateral roots. It 

can also be propagated by cuttings (Bonner 1974). 

Growth and Management.—Seedlings establish 

from animal-, gravity-, or water-dispersed 

propagules in sites free of herbaceous competition, 

often in rocky areas or in the duff of an existing 

stand. Plummer (1977) found netleaf hackberry 

useful in stabilizing disturbed areas in pinyonjuniper and mountain brush types of Utah. In 

Texas and Arizona, it is increasingly being used 

for riparian restoration and wildlife plantings. It 

currently receives limited use in artificial plantings 

in the northern portion of its range, though once 

established, young plants are hardy and persistent. 

Netleaf hackberry is generally slow-growing, but 

plants may live for 300 to 400 years (DeBolt and 

McCune 1995). 

Benefits.—Netleaf hackberry provides cover and 

food for a host of wildlife species. Its fruits are 

particularly important for birds during winter 

(Hayward 1948, Lanner 1983), and its leaves and 

twigs are browsed by bighorn sheep, mule deer, 

whitetail deer, elk, and occasionally by livestock. 

It could be used more widely for domestic 

landscaping as it requires little maintenance and is 

drought tolerant. Its hard wood is still used today 

on a limited basis for tool handles and firewood. 

References 

Albee, B.J., L.M. Schultz, and S. Goodrich. 1988. 

Atlas of the vascular plants of Utah. Occasional 

Publication 7. Utah Museum of Natural History, 

Salt Lake City, UT. 670 p. 

Bonner, F.T. 1974. Celtis L. Hackberry. In: C.S. 

Shopmeyer, tech. coord. Seeds of woody plants 

in the United States. Handbook 450. U.S. 

Department of Agriculture, Agriculture 

Washington, DC. p. 298-300. 

Brown, D.E. 1982. Great Basin montane 

scrubland. In: Biotic communities of the 

American Southwest–United States and Mexico. 

Desert Plants 4: 83-84. 

Carmichael, R.S., O.D. Knipe, C.P. Pase, and 

W.W. Brady. 1978. Arizona chaparral: plant 

associations and ecology. Agriculture Research 

Paper RM-202. U.S. Department of Agriculture, 

Ft. Collins, CO. 16 p. 

Daubenmire, R. 1970. Steppe vegetation of 

Washington. Technical Bulletin 62. Agriculture 

Experiment Station, Pullman, WA. 131 p. 

DeBolt, A.M. 1992. The ecology of netleaf 

hackberry in Idaho. Master’s thesis, Oregon 

State University, Corvallis, OR. 167 p. 

DeBolt, A.M. and B. McCune. 1995. Ecology of 

Celtis reticulata in Idaho. Great Basin Naturalist 

55: 237-248. 

Dick-Peddie, W.A. and J.P. Hubbard. 1977. 

Classification of riparian vegetation. Pages 85-

90 In: R.R. Johnson and A. Dale, eds. 

Importance, preservation, and management of 

riparian habitat. General Technical Report RM43. U.S. Department of Agriculture, Ft. Collins, 

CO. 

Elias, T.S. 1980. The complete trees of North 

America. Times Mirror Magazines, Inc., New 

York. 565 p. 

Hayward, C.L. 1948. Biotic communities of the 

Wasatch chaparral, Utah. Ecological 

Monographs 18: 473-506. 

Hitchcock, J.C., A. Cronquist, M. Ownbey, and 

J.W. Thompson. 1964. Vascular plants of the 

Pacific Northwest. Part 2: Salicaceae to 

Saxifragaceae. University of Washington Press, 

Seattle, WA. 597 p. 

Huschle, G. 1975. Analysis of the vegetation along 

the middle and lower Snake River. M.S. thesis, 

University of Idaho, Moscow. 271 p. 
169

Lanner, R.M. 1983. Trees of the Great Basin. 

University of Nevada Press, Reno, NV. 215 p. 

Little, E.L., Jr. 1976. Atlas of United States trees. 

Volume 3-Minor western hardwoods. 

Miscellaneous Publication 1,314. U.S. 

Department of Agriculture, Washington, DC. 

215 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: http://plants.usda.gov/ [not 

paged]. 

Plummer, A.P. 1977. Revegetation of disturbed 

intermountain area sites. In: J.C. Thames, ed. 

Reclamation and use of disturbed lands of the 

Southwest. University of Arizona Press, Tucson. 

p. 302-337. 

Stephens, H.A. 1973. Woody plants of the North 

Central Plains. University Press of Kansas, 

Lawrence, KS. 530 p. 

Tisdale, E.W. 1986. Canyon grasslands and 

associated shrublands of west-central Idaho and 

adjacent areas. Bulletin 40. Forestry, Wildlife, 

and Range Experiment Station, University of 

Idaho, Moscow. 42 p. 

Van Auken, W.W., A.L. Ford, and A. Stein. 1979. 

A comparison of some woody upland and 

riparian plant communities of the southern 

Edwards Plateau. Southwestern Naturalist 24: 

165-180.
170 

Cephalanthus occidentalis L. buttonbush 

RUBIACEAE 

 

Synonyms: Cephalanthus berlandieri Wernh. 

Kristina Connor 

General Description.—Buttonbush, also known 

as buttonball, button willow, common buttonbush, 

honey-bells, globe-flower, and riverbush, is a 

deciduous, wetland shrub or small tree that can 

reach 6 m in height but generally averages 1 to 3 

m tall. The trunk base is often swollen (Snyder 

1991). Branches are generally green when young 

but darken upon maturity and have conspicuous, 

raised lenticels (Brown and Kirkman 1990). The 

short-petioled glossy green leaves are elliptic or 

lanceolate-oblong; they are mostly opposite but, 

on the same plant, can occur in whorls of three or 

four. Leaves range in size from 7 to 18 cm in 

length and are 4 to 10 cm wide. 

Range.—Buttonbush is common along stream and 

pond borders, in swamps, floodplains and other 

riparian areas throughout the eastern half of the 

United States. It occurs naturally in southern Nova 

Scotia, New Brunswick, Quebec, and Ontario as 

well as through the eastern half of the Great Plains 

States; scattered populations and varieties are 

found in Arizona, New Mexico, southern 

California, and Texas. It also grows in Mexico, 

Cuba, Central America, and in the West Indies 

(Little 1979). Recognized varieties are C. 

occidentalis L. var. californicus Benth., C. 

occidentalis var. pubescens Raf., and C. 

occidentalis var. angustifolius Dippel. 

Ecology.—Buttonbush is a wetland species that 

cannot tolerate drought. It commonly grows in 

thickets in areas that have intermittent flooding. 

The open, rangy plant is not particularly attractive 

and is seldom found in cultivation today, although 

Van Dersal (1938) and Vines (1960) reported that 

it was cultivated as early as 1735 as a honey plant. 

The creamy white summer flowers of buttonbush 

attract butterflies, honeybees, and hummingbirds. 

While it is common in natural environments, its 

habitat is threatened in California and it grows 

poorly along manmade waterways (Holstein 

1984). Faber-Langendoen and Maycock (1989) 

state that buttonbush abundance increases with 

increasing depth of water and light levels, while 

Holstein (1984) suggests distribution may be 

limited by mean July temperatures of 20 o

C. It is 

classified as a pioneer species and grows best in 

wet areas that receive full sun. It is able to tolerate 

some salinity that might result from hurricane 

storm surges but will not survive long-term 

exposure to salt water (McCarron and others 

1998). 

Reproduction.—The perfect, creamy white 

terminal and axillary flowers of buttonbush occur 

in dense spherical heads 2 to 3.5 cm in diameter. 

The flower balls stand on stalks that are 5 cm long. 

The fragrant individual flowers have corollas 6 to 

10 mm long, with pubescent lobes, and four 

stamens that extend 5 to 8 mm beyond the 

corollas. Flowers are produced over a long period, 

from late spring, throughout the summer months, 

and into early autumn. Flowers are thought to be 

self-incompatible (Imbert and Richards 1993). The 

long-stemmed fruits are clusters of achenes. The 

hard nutlets are 4 to 7 mm long and turn reddishbrown when mature (August to November). Fruit 

balls may persist throughout the winter. Bonner 

(1974a) reported an average of 295,000 seeds/kg. 

Seeds are high in carbohydrates (Bonner 1974b). 
171

Longevity of seeds in storage is unknown but they 

are thought to be orthodox. Thus, if seeds are dried 

to a moisture content of 12 percent or less, they 

can be stored under refrigeration for long periods 

(Roberts 1973). Germination is epigeal, cotyledons 

forced above ground, and no pretreatment is 

required. DuBerry (1963) and Bonner (1974a) 

reported germinations from 78 to 86 percent while 

Vines (1960) reported a low germination 

percentage. Seeds may be transported by animals, 

birds, or by water. Buttonbush may also be 

propagated from tip cuttings in the spring or 

mature-wood cuttings in the winter. 

Growth and Management.—Buttonbush is a fast 

growing but short-lived shrub. It can be used as a 

landscape shrub border but if not planted in moist 

soil, it must be watered frequently. It must also be 

pruned to maintain a good form. In nature, 

buttonbush occurs in dense thickets. It will 

resprout after fire (Vogl 1973, Wade and others 

1980). It has no reported pest problems but is 

moderately sensitive to herbicides. Thickets can be 

reduced by cutting. 

Benefits.—Buttonbush seeds are an important 

food for water birds but can be toxic to other 

animals (Snyder 1991). Dense buttonbush thickets 

provide a safe nesting ground for many wetland 

birds. Buttonbush also serves as a source of honey 

for butterflies, bees, and hummingbirds. A 

decoction of the inner bark was used by Native 

Americans as an emetic. The bark was also used as 

a substitute for quinine. The wilted leaves, which 

contain bitter glycosides, cephalin and 

cephalanthin (ACES 2001), are reportedly toxic to 

some animals, especially cattle that eat them when 

other foliage is scarce. However, buttonbush 

leaves are eaten by deer (Bramble and Goddard 

1943), apparently with no ill effects. Other plant 

parts are less toxic. 

References

ACES, Alabama Cooperative Extension System. 

2001. Poisonous Plants of the Southeastern 

United States.ANR-975 http://www.aces.edu/ 

dept/extcomm/ publications. 56 p. 

Bonner, F.T. 1974a. Cephalanthus occidentalis L., 

common buttonbush. In: C.S Schopmeyere, 

tech. coord. Seeds of woody plants in the United 

States. United States Department of Agriculture 

Forest Service Agriculture Handbook 450, 

Washington, D.C. 883 p. 

Bonner, F.T. 1974b. Chemical components of 

some southern fruits and seeds. United States 

Department of Agriculture Forest Service Res. 

Note SO-183, New Orleans. 3 p. 

Bramble, W.C. and M.K. Goddard. 1943. Seasonal 

browsing of woody plants by white-tailed deer 

in the bear oak forest type. Journal of Forestry 

41(7): 471-475. 

Brown, C.L. and L.K. Kirkman. 1990. Trees of 

Georgia and adjacent states. Timber Press, 

Portland, OR. 292 p. 

DuBerry, A.P., Jr. 1963. Germination of 

bottomland tree seed while immersed in water. 

Journal of Forestry 61: 225- 226. 

Faber-Langendoen, D. and P.F. Maycock. 1989. 

Community patterns and environmental 

gradients of buttonbush, Cephalanthus 

occidentalis, ponds in lowland forests of 

southern Ontario. Canadian Field-Naturalist 

103(4): 479-485. 

Holstein, G. 1984. California riparian forests: 

deciduous islands in an evergreen sea. In: R.E. 

Warner and K.M. Hendris, eds. California 

riparian systems: Ecology, conservation, and 

productive management Proceedings of a 

conference; 1981 Sept. 17-19, Davis CA. 

University of California Press, Berkeley, CA. 

p. 2-22. 

Imbert, F.M. and J.H. Richards. 1993. Protandry, 

incompatibility, and secondary pollen 

presentation in Cephalanthus occidentalis

(Rubiaceae). American Journal of Botany 80: 

395-404. 

Little, E.L., Jr. 1979. Checklist of United States 

trees (native and naturalized). Agriculture 

Handbook 541. United States Department of 

Agriculture, Forest Service, Washington, D.C. 

375 p. 

McCarron, J.K., K.W. McLeod, and W.H. Conner. 

1998. Flood and salinity stress of wetland 

woody species buttonbush (Cephalanthus 

occidentalis) and swamp tupelo (Nyssa sylvatica

var. biflora). Wetlands 18: 165-175. 

Roberts, E.H. 1973. Predicting the storage life of 

seeds. Seed Science and Technology 1: 499-514. 
172 

Snyder, S.A. 1991. Cephalanthus occidentalis. In: 

United States Department of Agriculture, Forest 

Service, Rocky Mountain Research Station Fire 

Sciences Laboratory. Fire Effects Information 

System, http://www.fs.fed.us/database/feis. 10 p. 

Van Dersal, W.R. 1938. Native woody plants of 

the United States: their erosion-control and 

wildlife values. United States Department of 

Agriculture Miscellaneous Publication 393, 

Washington, D.C. 362 p. 

Vines, R.A. 1960. Trees, shrubs, and woody vines 

of the Southwest. University of Texas Press, 

Austin, TX, 1,104 p. 

Vogl, R.J. 1973. Effects of fires on the plants and 

animals of a Florida wetland. American Midland 

Naturalist 89: 334-347. 

Wade, D., J. Ewel, and R. Hofstetter. 1980. Fire in 

South Florida ecosystems. United States 

Department of Agriculture Forest Service 

General Technical Report SE-17. Asheville, NC. 

125 p. 
173

Cercocarpus intricatus Wats. littleleaf mountain-mahogany 

ROSACEAE 

Synonyms: Cercocarpus ledifolius var. intricatus (Wats.) Jones 

 Cercocarpus arizonicus Jones 

Stanley G. Kitchen 

 

General Description.—Littleleaf mountainmahogany, also known as little leaf cercocarpus 

and dwarf mountain-mahogany, is a long-lived, 

unarmed, intricately branched and occasionally 

tree-like, evergreen shrub. Height of mature plants 

varies from 0.5 to 2.5 m. Mature bark is smooth 

and light gray. Dark brown heartwood is dense and 

extremely hard. Persistent leaves are 3 to 18 mm 

long, 0.8 to 1.4 mm wide (oblong to linear) and 

leathery with strongly revolute margins (Welsh 

and others 1987). Leaf pubescence varies from 

glabrous to villous. Perfect, apetalous flowers, 3 to 

9 mm long, occur singularly or in small clusters 

arising primarily from short spur branchlets on 

second-year growth. Hard, light brown fruits are 

cylindrical achenes, approximately 1 mm wide and 

4 to 10 mm long with a persistent twisted or 

spiraling tail (style) 1 to 3 cm long (Welsh and 

others 1987). Fruit body and especially the tail are 

covered with short (1 mm) hairs that extend from 

the axis when dry and facilitate wind dispersal. 

Taxonomy.—The evolutionary process by which 

Littleleaf mountain-mahogany segregated from 

curlleaf mountain-mahogany (C. ledifolius Nutt. 

var. ledifolius) appears to still be in progress with 

intermediates between the species widespread 

(Stutz 1990). The shorter and more compact 

littleleaf mountain-mahogany is more drought 

tolerant, has fewer stamens and shorter tails on 

fruits than does curlleaf mountain-mahogany 

(Stutz 1990). Although hybridization occurs 

between the two species, littleleaf mountainmahogany appears to exhibit much greater 

reproductive isolation from true mountain 

mahogany (C. montanus Raf.) than does its 

ancestral species (Stutz 1990). Walker and Turley 

(1999) cited examples of hybrids between littleleaf 

and true mountain-mahoganies, suggesting that a 

lack of hybridization may be due, at least in part, 

to limited interspecies contact. 

Range.—Littleleaf mountain-mahogany is found 

throughout most of Nevada and Utah and in parts 

of eastern California, northern Arizona, and 

extreme western Colorado at elevation of 900 to 

3,000 m (Davis 1990). Limited populations have 

also been reported for New Mexico and Wyoming 

(NatureServe Explorer 2002). 

Ecology.—Within desert shrub, pinyon-juniper, 

and mountain brush communities, littleleaf 

mountain mahogany is typically found rooted in 

cracks and crevices of exposed sedimentary and 

metamorphic rocks; commonly limestone, 

dolomite, and quartzite in the Great Basin and 

sandstone in the Colorado Plateau (Davis 1990). It 

is also found in shallow, poorly developed soils 

derived from these substrates. It grows best in full 
174 

sun. Soil pH of littleleaf mountain-mahogany 

habitat is neutral to slightly basic, (Fairchild and 

Brotherson 1980). Annual precipitation is 200 to 

500 mm. Although undocumented, it is believed 

that this species possesses nitrogen fixing root 

nodules similar to those reported for close relatives 

(Davis 1990). Littleleaf mountain-mahogany is 

browsed heavily by wild ungulates and domestic 

sheep when it can be reached. 

Reproduction.—Flowering occurs from mid-May 

to late June (Walker and Turley 1999). Fruits ripen 

and disperse in July and August. Years of 

abundant seed production are rare. A high 

percentage of empty fruits is common. Plummer 

and others (1968) reported 112,000 cleaned seeds 

(fruits with tails removed) per kg. Seed dispersal is 

by wind. 

Growth and Management.—Although 

information on seed germination for this species is 

lacking in the literature, primary dormancy at seed 

dispersal is probably similar to that of curlleaf 

mountain-mahogany. Kitchen and Meyer (1990) 

observed almost no germination for this species 

without prechill (1 o

C). They reported germination 

of six seed collections after 6 weeks of prechill 

ranged from 53 to 96 percent of viable. Seedling 

growth rate is slow, and plants require 10 to 30 

years to reach maturity. Management alternatives 

for littleleaf mountain-mahogany are limited due 

to the nature of the sites it occupies.

Benefits.—Littleleaf mountain-mahogany 

provides cover and year-round forage on harsh 

landscapes. It is of particular value to native 

ungulates that depend on rugged escape terrain 

such as mule deer (Ococolius hemionus), bighorn 

sheep (Ovis Canadensis), and mountain goat 

(Oreamnos americanus). 

References.

Davis, J.N. 1990. General ecology, wildlife use, 

and management of the mountain mahoganies in 

the Intermountain West. In: K.L. Johnson, ed. 

Proceedings of the Fifth Utah Shrub Ecology 

Workshop: The Genus Cercocarpus; 1988 July 

13-14; Logan, UT. College of Natural 

Resources, Utah State University, Logan, UT. 

p. 1-13. 

 

Fairchild, J.A. and J.D. Brotherson. 1980. Microhabitat relationships of six major shrubs in 

Navajo National Monument, Arizona. Journal of 

Range Management 33: 150-156. 

Kitchen, S.G. and S.E. Meyer. 1990. Seed 

dormancy in two species of mountain-mahogany 

(Cercocarpus ledifolius and Cercocarpus 

montanus). In: K.L. Johnson, ed. Proceedings of 

the Fifth Utah Shrub Ecology Workshop: The 

Genus Cercocarpus; 1988 July 13-14; Logan, 

UT. College of Natural Resources, Utah State 

University, Logan, UT. p. 27-42. 

NatureServe Explorer. 2002. An online 

encyclopedia of life [web application]. Version 

1.6. NatureServe. Arlington, VA, USA. 

http://www.natureserve.org/explorer. [not 

paged]. 

Plummer, A.P., D.R. Christensen, and S.B. 

Monsen. 1968. Restoring big-game range in 

Utah. Publ. No. 68-3. Utah division of Fish and 

Game, Ephraim, UT. 183 p. 

Stutz, H.C. 1990. Taxonomy and evolution of 

Cercocarpus in the western United States. In: 

K.L. Johnson, ed. Proceedings of the Fifth Utah 

Shrub Ecology Workshop: The Genus 

Cercocarpus, 1988 July 13-14, Logan, UT. 

College of Natural Resources, Utah State 

University, Logan, UT. p. 15-25. 

Walker, S.C. and D. Turley. 1999. Characteristics 

of mountain mahogany (Cercocarpus) species 

and hybrids in Utah hybrid zone. In: E.D. 

McArthur, W.K. Ostler, and C.L. Wambolt, 

comps. Proceedings: Shrubland Ecotones; 1998 

August 12-14; Ephraim, UT. Proceedings 

RMRS-P-11. U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research 

Station, Ogden, UT. p. 32-34. 

Welsh, S.L., N.D. Atwood, L.C. Higgins, and S. 

Goodrich. 1987. A Utah Flora. Brigham Young 

University Print Services, Provo, UT. 894 p.
175

Cercocarpus ledifolius Nutt. curl-leaf mountain mahogany 

ROSACEAE 

Synonyms: None 

Christopher Ross 

General Description.—Curl-leaf mountain 

mahogany is a large-sized shrub to small or 

medium tree. Young stems are smooth and gray, 

becoming rough and brown in mature specimens, 

and often scaly in older trees. The evergreen leaves 

are narrowly lanceolate, often revolute, about 1 to 

1.5 cm long with a 1- to 3-mm petiole. They are 

densely white-wooly underneath. (Lis 1993). The 

trees are extremely long lived in the absence of 

external sources of mortality and are often by far 

the oldest members of the communities in which 

they occur (Ross 1999). 

Range.—Curl-leaf ountain mahogany is widely 

distributed in North America, from Montana to 

Baja California and from near the Pacific Coast in 

southwestern Oregon to the Bighorn Mountains in 

Wyoming and near Chadron, Nebraska. It occurs at 

elevations of 610 to 1,372 m in the north, and to 

3,000 m or higher in the south (USDA USFS 1937, 

Martin 1950). 

Ecology.—Curl-leaf mountain mahogany occurs in 

a variety of plant associations, including sagebrush, 

pinyon/juniper, scrub oak/mountain brush, aspen, 

ponderosa, jeffrey and other pines, and spruce/fir 

(Martin 1950, Ross 1999). It occurs on a wide 

variety of soils (Davis and Brotherson 1991), 

including decomposed granite and limestone, 

altered andesites (Billings 1976) and others. Curlleaf mountain mahogany often occurs in rock 

outcrops and talus slopes with no apparent soil 

development. In pine and spruce/fir forests it may 

eventually be overshadowed by tall trees and 

shaded out. It often forms dense, closed canopy 

stands that permit little understory or interspecific 

competition. Sapsuckers and other woodpeckers 

may attack intermediate age class trees, girdling 

the stems and causing extensive stand mortality in 

some cases (Ross 1999). Other predators include 

the bark beetle Renoci’s heterodoxus (Kraft 1960) 

and the leaf defoliater Anacamtodes clivinavia 

profanata (Furniss and Barr 1969). A variety of 

other insects also attack the species at times. 

Reproduction.—Flowers are one to three per 

inflorescence, axillary, and more or less sessile, 

with a straight stigma and 15 to 25 stamens. Fruits 

are unusual for a member of the rose family, 4 to 8 

mm long, hard, with a 2.4 to 3.5 cm plume style 

(Lis 1993). Seed production is episodic but may be 

very heavy at times, with the seeds forming drifts 

beneath trees. Seed predation by insects in fall may 

be nearly complete at times (Dealy 1975). 

Reproduction is sporadic, occurring usually on bare 

mineral soil, including fresh road cuts, but very 

uncommon in established plant communities. The 

increase in cheatgrass (Bromus tectorum L.) and 

other annuals over its range has apparently reduced 

reproduction in many areas (Ross 1999). Under 

established stands, germination is common but the 

seedlings usually damp off and die (Ross 1999). 

Episodic regeneration and mortality, especially 

from fire and sapsuckers, may create even-aged 

stands with almost no diversity of age class. 

Fire Effects.—Although curl-leaf mountain 

mahogany is sometimes referred to as a weak 

resprouter after fire, this is very uncommon. In the 

western Great Basin, it is invariably killed by fire 

regardless of intensity, and never resprouts. Even 
176 

very light burns that do no apparent damage to 

mature trees result in full mortality within 1 year. 

Closed mature stands may have insufficient 

understory to carry fire, so that the fire and 

resulting mortality are confined to the edges of the 

stands. Regeneration by seed may occur after fire 

if the soil is not colonized by other species. In 

some cases, the resulting stands occupy essentially 

the same area as the burned stand. Historic fire 

lines and burns are often clearly demarcated by the 

sharp boundaries of living stands. The wood is 

rarely consumed by fire and often persists for 

decades after, offering precise information about 

past distribution and age classes. With changes in 

fire intensity and return interval, usually related to 

increases in exotic annuals, curl-leaf mountain 

mahogany in many areas is decreasing and is 

increasingly restricted to higher elevations, steeper 

slopes, and cooler aspects (Ross 1999).

Growth and Management.—Although curl-leaf 

mountain mahogany is very long lived and 

eventually assumes a tree form, young specimens 

grow quickly as multistemmed shrubs. Stands are 

typically composed of a single age class, perhaps 

with younger specimens on the edges. It may be 

readily grown from seed for transplanting, although 

young transplants are attractive to rodents, 

lagomorphs, and large game. Seed collection is 

easily done, although the hairy styles are a strong 

respiratory and skin irritant. Management has 

focused on increasing the forage production of old, 

browsed stands via clipping and pruning (Davis 

and Brotherson 1991), although this may result in 

tree mortality (Garrison 1953, Thompson 1970). 

Benefits.—Curl-leaf mountain mahogany has 

found extensive use in the past as a fuel and 

charcoal source. The very hard, beautiful wood 

has been used for lathe work and parts for musical 

instruments (Ross 1999). Living stands provide 

significant blue grouse nesting habitat (Stauffer 

and Petersen 1986). It is one of the few species that 

meet protein requirements for wintering deer 

(Welch and McArthur 1979) and is heavily favored 

by bighorn sheep in summer (Rominger et al. 

1988). Where present in even small amounts in 

sagebrush, curl-leaf mountain mahogany allows 

greater wildlife diversity and abundance than 

would otherwise occur (Furniss et al. 1988). It is 

also used increasingly as a commercial, residential, 

and highway landscaping species. 

References 

Billings, W. D. 1976. Islands of sierran plants on 

the arid slopes of Peavine Mountain. Mentzelia 

6: 32-39. 

Davis, J.N. and J. D. Brotherson. 1991. Ecological 

relationships of curlleaf mountain mahogany 

(Cercocarpus ledifolius Nutt.) communities in 

Utah and implications for management. Great 

Basin Naturalist 51: 153-66. 

Dealy, J.E. 1975. Ecology of curlleaf mountain 

mahogany (Cercocarpus ledifolius Nutt.) in 

eastern Oregon and adjacent areas. PhD. 

dissertation. Oregon State University, Corvallis, 

OR. 

Furniss, M.M. and W. F. Barr. 1969. Bionomics of 

Anacamprodes clivinaria profanata on mountain 

mahogany in Idaho. Agricultural Experiment 

Station Research Bulletin 73. University of 

Idaho, Moscow, ID. 24 p. 

Furniss M.M., D.C. Ferguson, K.W. Boget, J.W. 

Burkhardt, A.R. Tiedemann, and J.L. 

Oledmeyer. 1988. Taxonomy, life history, and 

ecology of a mountain mahogany defoliator, 

Stamnodes animata Pearsall in Nevada. Fish 

and Wildlife Research 3, USFWS, Washington, 

D.C. 

Garrison, G.A. 1953. Effects of clipping on some 

range shrubs. Journal of Range Management 6: 

309-317. 

Kraft, G.F. 1960. Insects affecting bitterbrush and 

other range plants. Office Report. Entomology 

Dept., Oregon State University, Corvallis, OR. 

55 p. 

Lis, R. 1993. Cercocarpus Mountain Mahogany. 

In: J.C Hickman, ed. 1993. The Jepson Manual: 

higher plants of California. U.C. Press, Berkeley 

and Los Angeles, CA. 

Martin, F.L. 1950. A revision of Cercocarpus. 

Brittonia 7: 91-111. 

Rominger, E.M., A.R. Dale, and J.A. Bailey. 1988. 

Shrubs in the summer diet of Rocky Mountain 

bighorn sheep. Journal of Wildlife Management 

52: 47-50. 

Ross, C. 1999. Population dynamics and changes 

in curlleaf mountain mahogany (Cercocarpus 
177

ledifolius Nutt.) in two adjacent Sierran and 

Great Basin mountain ranges. PhD. Dissertation, 

University of Nevada, Reno. 

Stauffer, D.F. and S.R. Petersen. 1986. Seasonal 

microhabitat relationships of blue grouse in 

southeastern Idaho. Great Basin Naturalist 46: 

117-22. 

Thompson, R.M. 1970. Experimental top pruning 

of curlleaf mountain mahogany on the South 

Horn Mountain, Ferron Ranger District, MantiLaSal National Forest, USDA Forest Service 

Range Improvement Notes, Vol. 15, No. 3. 

USDA USFS. 1937. Range Plant Handbook. 

Washington D.C. U.S. Govt. Printing Office. 

p. 132–133. 

Welch, B.L. and E.D. McArthur. 1979. Nutritive 

value of big sagebrush and other shrubs. In L.H. 

Stelter and others., eds. Proceedings, Shrub 

establishment on disturbed arid and semi-arid 

lands, Wyoming Fish and Game Dept. p. 9-22.
178 

Cercocarpus montanus Raf. true mountain-mahogany 

ROSACEAE 

Synonyms: Cercocarpus betuloides Nutt. in T. & G. 

 Cercocarpus betulifolius Nutt. ex Hook. 

 Cercocarpus parvifolius Nutt. 

 Cercocarpus flabellifolius Rydb. 

Stanley G. Kitchen 

General Description.—True mountainmahogany, also known as mountain cercocarpus, 

birchleaf cercocarpus, birchleaf mountainmahogany, alderleaf mountain-mahogany, 

blackbrush, deerbush, tallowbrush, and lintisco 

(Spanish), is a long-lived, unarmed, mostly 

deciduous shrub, with few to many ascending 

stems generally 1 to 4 m tall and 0.5 to 5 cm in 

diameter at the base. Mature bark is smooth and 

light gray. Stout tap and lateral roots penetrate 

deep into available soil and rock crevices. Leaves 

are simple and alternate. Leaf length varies from 6 

to 44 mm. Leaf blade shape is oblanceolate to 

roundish with shallow forward-pointing teeth on 

the terminal margin. The upper surface is glabrous 

while the lower surface is generally pubescent 

(Welsh and others 1987). Perfect, apetalous 

flowers, 10 to 17 mm long, occur singularly or in 

small clusters arising from axiles or on short spur 

branchlets along upper portions of 2-year-old 

stems. Hard, pale green to light brown fruits are 

pubescent, cylindrical achenes, approximately 1 to 

2 mm wide and 8 to 15 mm long with a persistent 

twisted or spiraled style or tail 3 to 10 cm in 

length. The numerous 1 to 2 mm long hairs present 

on this appendage extend almost perpendicular to 

the style axis when dry and facilitate wind 

dispersal. Because of the reflective nature of these 

hairs, abundant seed crops give plants a frosted 

look in sunlight. 

Taxonomy.—Nine varieties of true mountain 

mahogany are recognized of which var. montanus 

Raf. is most widespread (Marshall 1995, 

NatureServe Explorer 2002). Varieties with 

southwestern United States distributions include 

vars. argenteus (Rydb) F.L. Martin, glaber (S. 

Wats.) F.L. Martin, and the evergreen 

paucidentatus (S. Wats) F.L. Martin. Closely 

related species of the Southwest and Mexico 

include: C. mexicanus Hendrickson, C. rzedowski

Hendrickson, and C. fothergilloides Kunth, 

suggesting a southern North American origin 

(Stutz 1990). Coastal California has varieties with 

restricted distributions including the critically 

imperiled (S1) Santa Catalina mountain-mahogany 

or C. montanus traskiae (Eastw.) F.L. Martin, 

which is sometimes treated as a separate species, 

and the vulnerable (S3) Island mountainmahogany or C. montanus blancheae (Schneid.) 

F.L. Martin (NatureServe Explorer 2002). True 

mountain-mahogany is known to hybridize with 

other mountain-mahogany species where 

populations are in contact (Stutz 1990, Walker and 

Turley 1999). Hybrids derived from crosses with 

the more tree-like and evergreen curlleaf mountain 

mahogany (Cercocarpus ledifolius var. 

intermontanus N. Holmgren) are evergreen, 

intermediate in stature and leaf shape, and usually 
179

sterile. In contrast, hybrids formed with C. 

ledifolius Nutt. spp. ledifolius, are quite fertile and 

backcross with parent populations (Stutz 1990). 

Range.—True mountain-mahogany, like other 

members of the genus, is endemic to dry coastal 

and interior foothills and mountains of the Western 

United States and Mexico (Marshall 1995, Stutz 

1990). In the United States, it is centrally located 

in Colorado, Utah, and Wyoming and to lesser 

degrees in South Dakota, Nebraska, Kansas, 

Oklahoma, Texas, New Mexico, Arizona, Nevada, 

California, Oregon, Idaho and Montana (Davis 

1990, NatureServe Explorer 2002). Within its 

range it grows at elevations of 1,000 and 3,000 m 

(Davis 1990). 

Ecology.—True mountain-mahogany grows in 

solid, sometimes extensive stands, and in mixtures 

in sagebrush and mountain shrublands and pinyonjuniper woodlands. This species is also found in 

openings of ponderosa pine, mixed conifer, and 

aspen forests. It does best in full sun. Annual 

precipitation ranges from 250 to 600 mm. Soils are 

well drained, near-neutral to slightly basic, and 

deep to absent (rock crevices). Nitrogen fixation in 

root nodules and ectomycorrhizal facilitation of 

phosphorus uptake (Hoeppel and Wollum 1971) 

are probably critical for the success of this species 

on infertile soils. Plants often appear hedged from 

heavy winter use by wildlife. Regeneration by 

crown sprouting is best after early season fires 

(Marshall 1995). Ferguson (1983) suggested that 

the ability to sprout following fire might be 

ecotype dependent. Stand regeneration by seed is 

slow. 

Reproduction.—Anthesis for this wind pollinated 

species occurs between April and early July 

depending on elevation, latitude, and aspect. Fruits 

ripen and disperse from July to October. A unique 

pattern of resource partitioning is commonly 

observed in which branches with few seeds and 

long leader growth occur concurrently with 

branches sporting many fruits and little leader 

development (Walker and Turley 1999). Abundant 

fruit production may occur at intervals of 1 to 10 

years (Plummer and others 1968). A high 

percentage of nonviable (empty) fruit is common 

for this species and the genus Cercocarpus in 

general. Means of 90,000 and 130,000 cleaned 

seeds per kg (fruits with tails removed) have been 

reported in the literature (Kitchen and others 1989, 

Deitschman and others 1974). Primary seed 

dispersal is by wind. However, seeds are also 

gathered and likely cached by rodents. The 

possible significance of rodents as secondary 

dispersers of mountain-mahogany seeds is 

unknown. 

Growth and Management.—Although withinplant flowering and fruit maturation are well 

synchronized, some within-population asynchrony 

is expected. Ripened fruits are easily dislodged 

and collected by striking or shaking branches over 

hand-held hoppers. Optimal season for harvest 

varies from June to September. During harvest and 

handling, fruit hairs dislodge and can cause 

considerable discomfort to eyes and skin, 

revealing the motivation for the cowboy moniker, 

“hell feathers” (Plummer and others 1968). 

Commercially available seed is typically cleaned 

to 95 percent purity and 85 percent viability. Seeds 

have been warehouse-stored for 7 years without 

significant loss in viability (Stevens and others 

1981). Although primary dormancy is apparently 

absent for some collections of true mountainmahogany seed (Deitschman and others 1974), 

moist chilling (stratification) treatments of 2 to 12 

weeks are typically required to break dormancy 

(Kitchen and Meyer 1990). In one study, seeds 

germinated in chilling (2 o

C) between 7 and 15 

weeks of treatment (Kitchen and Meyer 1990). A 

germination inhibitor may play a role in 

germination regulation (Moore 1963). Laboratory 

viability evaluations using standard TZ 

(tetrazolium chloride) tests can be problematic due 

to difficulty in extracting undamaged embryos 

from fruits (Kitchen and others 1989). Seeds are 

used in restoration plantings of highly disturbed 

sites (e.g. surfacing mines) and for wildlife habitat 

improvement. Seedling vigor is fair and growth 

rate is slow to moderate. Plants reach maturity in 5 

to 15 years. Plants are tolerant of moderate to 

heavy use by browsers (Turley 2000) and shoots of 

true mountain-mahogany provide adequate 

nutrition as winter forage for wildlife. Productivity 

decreases when plants are protected from 

herbivory over extended periods (Waugh 1990). 

Decadent stands can be rejuvenated using light to 

moderate top removal techniques such as one-way 

chaining (Davis 1990). Both containerized and 

bare-root stock (1 and 2 year old) is available from 

select nurseries. 

Benefits.—True mountain mahogany is an 

important, often dominant shrub in many parts of 

the Western United States. It provides valuable 

winter forage for wildlife. It also provides 

protection for, and increases the fertility of, 

naturally occurring and human-caused (e.g. mine 

spoils) infertile soils. 
180 

References

Davis, J.N. 1990. General ecology, wildlife use, 

and management of the mountain mahoganies in 

the Intermountain West. In: K.L. Johnson, ed. 

Proceedings of the Fifth Utah Shrub Ecology 

Workshop: The Genus Cercocarpus; 1988 July 

13-14; Logan, UT. College of Natural 

Resources, Utah State University, Logan, UT. 

p. 1-13. 

Deitschman, G.H., K.R. Jorgensen, and A.P. 

Plummer. 1974. Cercocarpus. In: C.S. 

Schopmeyer, tech. coord.. Seeds of Woody 

Plants of the United States. Agricultural 

Handbook 450. U.S. Department of Agriculture, 

Forest Service, Washington, DC. p. 309-312. 

Ferguson, R.B. 1983. Use of wildland shrubs for 

wildland plantings in the Intermountain West. 

In: S.B. Monsen and N. Shaw, comps. Managing 

Intermountain Rangelands—Improvement of 

Range and Wildlife Habitats; Proceedings of 

Symposia; 1981 September 15-17; Twin Falls, 

ID and 1982 June 22-24; Elko NV. General 

Technical Report INT-157. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Research Station, Ogden, UT. p. 136-147. 

Hoeppel R.E. and A.G. Wollum. 1971. 

Histological studies of ectomycorrhizae and root 

nodules from Cercocarpus montanus and 

Cercocarpus paucidentatus. Canadian Journal of 

Botany 49: 1315-1318. 

Kitchen, S.G. and S.E. Meyer. 1990. Seed 

dormancy in two species of mountain-mahogany 

(Cercocarpus ledifolius and Cercarpus 

montanus). In: K.L. Johnson, ed. Proceedings of 

the Fifth Utah Shrub Ecology Workshop: The 

Genus Cercocarpus; 1988 July 13-14; Logan, 

UT. College of Natural Resources, Utah State 

University, Logan, UT. p. 27-42. 

Kitchen, S.G., S.E. Meyer, G.R. Wilson, and R. 

Stevens. 1989. Addition of Cercocarpus 

montanus—true mountain-mahogany to the 

Rules. Association of Official Seed Analysts 

Newsletter 63: 28-30. 

Marshall, K.A. 1995. Cercocarpus montanus. In: 

U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System, 

http://www.fs.fed.us/database/feis/ 17 p. 

Moore, T.C. 1963. A germination inhibitor in 

achenes of Cercocarpus montanus. Ecol. 44: 

406-409. 

NatureServe Explorer. 2002. An online 

encyclopedia of life. Version 1.6. NatureServe, 

Arlington, VA, USA. http://www.natureserve. 

org/explorer [not paged]. 

Plummer, A.P., D.R. Christensen, and S.B. 

Monsen. 1968. Restoring big-game range in 

Utah. Publication 68-3. Utah Division of Fish 

and Game, Ephraim, UT. 183 p. 

Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981. 

Viability of seed from thirty-two shrub and forb 

species through fifteen years of warehouse 

storage. Great Basin Naturalist 4: 274-277. 

Stutz, H.C. 1990. Taxonomy and evolution of 

Cercocarpus in the western United States. In: 

K.L. Johnson, ed. Proceedings of the Fifth Utah 

Shrub Ecology Workshop: The Genus 

Cercocarpus; 1988 July 13-14; Logan, UT. 

College of Natural Resources, Utah State 

University, Logan, UT. p. 15-25. 

Turley, D. 2000. Effects of browsing on true 

mountain mahogany (Cercocarpus montanus

Raf.) MS Thesis. Brigham Young University. 

Provo, UT. 115 p. 

Walker, S.C. and D. Turley. 1999. Characteristics 

of mountain mahogany (Cercocarpus) species 

and hybrids in Utah hybrid zone. In: E.D. 

McArthur, W.K. Ostler, and C.L. Wambolt, 

comps. Proceedings: Shrubland Ecotones; 1998 

August 12-14; Ephraim, UT. Proceedings 

RMRS-P-11. U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research 

Station, Ogden, UT. p. 32-34. 

Waugh, W.J. 1990. Stagnation and decadence of 

Cercocarpus montanus in a southeastern 

Wyoming big game exclosure. In: K.L. Johnson, 

ed. Proceedings of the Fifth Utah Shrub Ecology 

Workshop: The Genus Cercocarpus; 1988 July 

13-14; Logan, UT. College of Natural 

Resources, Utah State University, Logan, UT. p. 

89-95. 

Welsh, S.L., N.D. Atwood, L.C. Higgins, and S. 

Goodrich. 1987. A Utah Flora. Brigham Young 

University Print Services, Provo, UT. 894 p.
181

Cereus greggii Engelmann night-blooming cereus 

CACTACEAE 

Synonyms: Peniocereus greggii (Engelm.) Britt. & Rose 

 Cereus greggii Engelmann var. roseiflorus Kuntze 

Cereus pottsii Salm-Dyck 

Juanita A. R. Ladyman 

Illustration credit: Tonia Masaood

General Description.—The common names for 

Cereus greggii are night-blooming cereus, Texas 

night-blooming cereus, Arizona queen-of-thenight, reina-de-la-noche, sweet-potato cactus, and 

deer horn cactus. It is a perennial, succulent shrub 

(Vines 1960). Night-blooming cereus is a Federal 

Species of Concern and is listed as endangered by 

the State of New Mexico (Sivinski and Lightfoot 

1995, NMRPTC 2002). It is threatened by illegal 

collection throughout its range. Although this plant 

is cryptic and may be more common than verified 

documentation suggests, many occurrences of 

night-blooming cereus are known to have been 

extirpated due to collection pressures, and its 

range-wide abundance is believed to be declining. 

Night-blooming cereus has erect or sprawling 

stems that may grow to 2.4 m in length but are 

typically less than 1.5 m. The gray-brown stems 

are four to five ribbed and approximately 1.25 cm 

in diameter. There are 11 to 13 dark spines that 

tend to be swollen at the base, per areole. The 

lower spines may have some white coloration. The 

flower is particularly remarkable, having waxy 

white, pointed “petals” with numerous white to 

yellow-tipped stamens. It is fragrant and up to 5 

cm in diameter and 15 cm long. The orange-red, 

oblong fruit is up to 7.6 cm long and 3.8 cm in 

diameter with short spines. The root is a fleshy 

taproot and typically weighs 2.3 to 6.8 kg 

(Weniger 1991, Earle 1963, Vines 1960). In old 

plants it may be up to 0.6 m in diameter and weigh 

125 lbs (Weniger 1991). In 1919 a specimen with 

purplish flowers was collected in Organ, New 

Mexico, but since then only plants with white 

flowers have been recorded (Weniger 1969). There 

are two varieties of night-blooming cereus, Cereus 

greggii var. greggii and Cereus greggii var. 

transmontanus. Variety transmontanus differs 

from the typical variety in that the flowers are 

larger, approximately 7.5 cm in diameter, and the 

hypanthium is particularly spiny, and the spines 

generally 1.5 to 3.0 cm long (Benson 1982, Martin 

and Hutchins 1980). The hypanthium is the cuplike enlargement of the floral axis below the calyx 

that surrounds the ovary. The hypanthium of 

variety greggii is covered by minute, 

inconspicuous spines and the flowers are smaller. 

These varieties are in need of further study 

(NMNPPAC 1984). Some sources (Natural 

Resources Conservation Service 2003) maintain 

that the synonym, Peniocereus greggii (Engelm.) 

Britt. & Rose is the correct name. Night-blooming 

cereus provides an interesting evolutionary link. It 

appears to possess anatomical characteristics that 

are relictual from its nonsucculent, woody, 

Pereskia-like ancestor (Mauseth and others 1998). 

Range.—Night-blooming cereus grows in 

southern New Mexico, southeastern Arizona, 

western Texas, eastern Chihuahua, northeastern 

Durango, northern Zacatecas, and Coahuila, 

Mexico (Kearney and others 1960, Powell 1998, 
182 

Vines 1960). Benson (1982) defined the range of 

variety transmontanus as the Arizona and Sonoran 

Desert that possibly also includes Hildalgo County 

in New Mexico, and that of variety greggii as the 

Chihuahuan Desert of New Mexico, western Texas 

and Mexico. 

Ecology.—Night-blooming Cereus grows in dry 

alluvial soils at elevations between 370 and 1,500 

m. Again, the varieties are distinguished by variety 

transmontanus growing at the lower end of the 

elevation range (300 to 1,050 m) and variety 

greggii growing above 1200 m (Benson 1982). 

Night-blooming cereus generally grows in slightly 

broken to level terrain in desert grassland or 

Chihuahuan desert scrub. Typically it grows in 

sandy to silty gravelly soils on upper to mid 

bajadas among, depending upon the desert region, 

creosotebush (Larrea tridentata), mesquite 

(Prosopsis glandulosa), paloverde (Cercidium

species), and knife-leaf condalia (Condalia

species). It is also commonly associated with 

Ocotillo (Fouquieria splendens), tarbush 

(Flourensia cernua) and dropseed (Sporobolus

species). Habitat alteration such as that caused by 

the high impact trampling of heavy grazing is 

suspected to reduce population size (NMNPPAC 

1984). 

Reproduction.—As the common name suggests, 

night-blooming cereus blooms only at night. Just 

after dark the flowers open in a series of “jerks” 

(Benson 1982). Individual flowers only bloom for 

one night. The total flowering period of plants 

within a population is also brief and generally all 

flowers bloom within a 2-day period. However, in 

unusual circumstances, flowers on different 

individuals within a population may bloom 

periodically over a period of almost one month. 

Flowering occurs in late May into July depending 

on geographic area (Earle 1963, Epple 1995, 

Kearney and others 1960). The fragrance of the 

flowers is pleasant, and humans can detect the 

perfume of the flowers up to 30 m away from the 

plant (Benson 1982, Epple 1995, Vines 1960). 

Flowers are pollinated principally by hawk moths 

(ASDM 2002) and possibly other night feeding 

insects (Epple 1995) that search for nectar. The 

flowers are not self-fertile and must be crosspollinated. Therefore the hawk moths (Sphingidae) 

must fly hundreds of yards between the sparsely 

distributed plants (ASDM 2002). In areas where 

pesticides are heavily used for agriculture, the 

hawkmoth populations are devastated, and most of 

the flowers on plants in adjacent natural habitat 

fail to fruit (ASDM 2002). 

Growth and Management.—The stems generally 

grow up through, and later become supported by, 

shrubs. It is likely that the shrubs provide 

protection and a suitable microclimate for 

germination and subsequent plant development. 

Any activity that reduces shrub cover is a threat to 

night-blooming cereus. Therefore, fire is a 

potential hazard, although the direct effect of fire 

on night-blooming cereus is not well documented. 

The substantial tuberous root may provide the 

plant the means to recover if the above-ground 

stems are killed, and therefore, it is likely that 

individual plants can tolerate light fires that do not 

decimate the shrubs with which they are associated 

(Thomas 1991). More severe fires that scorch and 

heat the soil are likely to be more detrimental. 

Herbivores such as white-throated woodrats 

(packrats) and cactus borers (Cactobrosis 

fernaldialis) eat the stems, but new stems soon 

sprout from the tuberous root (ASDM 2002). 

Many populations throughout its range have been 

extirpated due to private and commercial 

collection, and the time required for population reestablishment is not known. However, although 

slow growing, it is easily propagated by short stem 

cuttings and from seed. Seeds of many cacti 

species, including Cereus species, are subject to 

physiological dormancy. For example, seeds of 

Cereus griseus were dormant at maturity but 

germinated after 8 weeks in a dry environment 

(Baskin and Baskin 2001). Cereus griseus seeds 

have an absolute light requirement for germination 

(Baskin and Baskin 2001). If the plant is dug up 

for transplantation it is actually difficult to keep 

alive (Weniger 1969). Apparently the root is 

susceptible to fungus. Damp soil has to be 

avoided, but even if root rot is prevented, the plant 

tends to decline over a few years if kept in a pot or 

in a greenhouse (Weniger 1969). 

Benefits.—Night-blooming Cereus plants have 

commercial value especially among cactus 

collectors (Epple 1995). Not only is it an 

aesthetically desirable cactus, but it has 

considerable medicinal value. The root is generally 

believed to be the organ that has medicinal 

properties (Moerman 1998, Moore 1989, Powell 

1998). However, the stems also have the same 

properties although “more feebly” (Moerman 

1998, Moore 1989, Powell 1998). Active 

ingredients include penicerol, viperidone, desoxy 

viperidone, viperidinone, β-sitosterol and, most 

likely, caffeine (Moore 1989). It is likely that 

night-blooming cereus is not unique in its 

medicinal value as other species of Cereus contain 

similar active ingredients (Ecdybase 2002). The 
183

commercial toll on this species has not been 

established, but wild-harvested material is 

available on the Internet (Pacific West Botanicals 

2002). It is called “pain in the heart” by the Death 

Valley Shoshones (Moore 1989). This tribe 

presumably uses it in a manner similar to Native 

Americans of Nevada who ingest an infusion of 

the roots as a cardiac stimulant (Moerman 1998). 

Other Native Americans have used a decoction of 

the roots for diabetes, the seedpods mixed with 

deer fat as a salve for sores, and the cut slices of 

root as an externally applied cure for chest colds 

(Moerman 1998, Vines 1960). The fruits, flowers, 

young stalks, and roots have been eaten for food 

(Moerman 1998). This use of the root may account 

for the common name “sweet potato cactus.” 

Chewing the raw root has been reported to quench 

thirst. Birds eat the seed and contribute to seed 

dispersal (Vines 1960). 

References 

Arizona Sonora Desert Museum. 2002. 

Peniocereus greggii. Internet web site: 

http://www.desert 

museum.org/books/nhsd_peniocereus.html#top. 

Baskin, C.C. and J.M. Baskin. 2001. Seeds, 

ecology, biogeography, and evolution of 

dormancy and germination. Academic Press, 

New York. 666 p. 

Benson, L. 1982. The cacti of the United States 

and Canada. Stanford University Press, 

Stanford, CA. p. 588-593. 

Earle, W.H. 1963. Cacti of the Southwest. Desert 

Botanical Garden, Phoenix, AZ. 210 p. 

Ecdybase. 2002. Internet Web Page: http:// 

ecdybase.org/index.php?row=308&action= 

browse. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. Falcon Press Publishing Co., Helena, 

MT. 347 p. 

Kearney, T.H., R.H. Peebles, J.T. Howell, and E. 

McClintock. 1960. Arizona flora. 2d ed. 

University of California Press. Berkeley, CA. 

1,085 p. 

Martin, W.C. and C.R. Hutchins. 1980. A Flora of 

New Mexico. Strauss & Cramer, Hirschberg, 

Germany. 2,592 p. 

Mauseth, J.D., T. Terrazas, and S. Loza-Cornejo. 

1998. Anatomy of relictual members of 

subfamily Cactoideae, IOS Group 1a 

(Cactaceae). Bradleya 16: 31-43. 

Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. 927 p. 

Moore, M. 1989. Medicinal plants of the desert 

and canyon west. Museum of New Mexico 

Press, Santa Fe, NM. 184 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Peniocereus greggii (Engelm.) 

Britt. & Rose. http://plants.usda.gov/ [not 

paged]. 

New Mexico Native Plants Protection Advisory 

Committee. 1984. A handbook of rare and 

endemic plants of New Mexico. University of 

New Mexico Press, Albuquerque, NM. 291p. 

New Mexico Rare Plant Technical Council. 2002. 

NM Rare Plants Web Page: http://nmrareplants. 

unm.edu/reports/ pengre.htm. 

Pacific West Botanicals. 2002. Internet web site: 

http://www.pacificwestbotanicals.com/ 

Night_Blooming_Cereus.html. 

Powell, A. M. 1998. Trees & shrubs of TransPecos and adjacent areas. University of Texas 

University Press, Austin, TX. 498 p. 

Sivinski, R. and K. Lightfoot 1995. Inventory of 

rare and endangered plants of New Mexico. 

Miscellaneous Publication 4. New Mexico 

Forestry and Resources Conservation Division, 

Energy, Mineral and Natural Resources 

Department, Santa Fe, NM. 

Thomas, P.A. 1991. Response of succulents to 

fire: A review. International Journal of Wildland 

Fire 1(1): 11-22. 

Vines, R.A. 1960. Trees, shrubs, and woody vines 

of the Southwest. Sixth printing 1986. 

University of Texas Press. Austin, TX. 1104 p. 

Weniger, D. 1969. Cacti of the Southwest. 

University of Texas Press, Austin, TX. 249 p. 

Weniger, D. 1991. Cacti of Texas and neighboring 

states. University of Texas Press, Austin, TX. 

356 p. 
184 

Cestrum diurnum L. day jasmine 

SOLANACEAE 

Synonyms: Cestrum fastigiatum Jacq. 

 Cestrum diurnum L. var. fastigiatum (Jacq.) Stehlé in Fournet 

 Cestrum diurnum portoricense O.E. Schulz in Urban 

John K. Francis 

General Description.—Day jasmine is a single or 

multistemmed shrub or rarely a small tree that is 

also known as day cestrum, wild jasmine, ink-bush, 

Chinese inkberry (English), dama de día, rufiana, 

galán de día, and saúco tintóreo (Spanish) (Little 

and others 1974). The bark of day jasmine is nearly 

smooth and gray. The inner bark is yellow-brown 

and slightly bitter. The twigs are slender, greenish 

gray, and sometimes drooping (Liogier 1995, Little 

and others 1974). The leaves of day jasmine are 

oblong or oblong-elliptic, and shiny green to 

yellow green, and somewhat membranous. They 

are about 15 by 6.5 cm and have petioles about 2.5 

cm long. The small, white flowers are grouped in 

panicles attached at the bases of the last leaves in 

each branch. These develop into elliptical, 

purplish-black berries about 6.5 mm long (Howard 

1989, Liogier 1995, Little and others 1974). 

Range.—Day jasmine is native to the Bahamas, 

Cuba, Jamaica, Hispaniola, and the Cayman 

Islands (Little and others 1974). It has been 

introduced as an ornamental into most of tropical 

and subtropical America. The species has escaped 

from cultivation in at least Florida, southern Texas, 

Hawaii, Puerto Rico, Guam, and American Samoa 

(Florida Exotic Plants Council 2001, Institute of 

Pacific Islands Forestry 2001). 

Ecology.—Day jasmine grows in gardens, 

roadsides, fencerows, pastures, vacant lots, and 

abandoned farmland in Puerto Rico (Liogier 1995). 

The species is particularly associated with pasture 

fencerows because its seeds are deposited there by 

birds, vegetative competition is controlled by 

grazing, and day jasmine is rarely eaten by cattle. 

There is no mention in the literature of the types of 

natural forest stands where day jasmine grows 

within its native range. However, because the 

species is intolerant of heavy shade and quickly 

disappears when overtopped by forest, it can be 

inferred that day jasmine is an opportunist that 

invades disturbed areas and quickly completes its 

life cycle ahead of encroaching forest. The species 

may grow as individual plants or in thickets. Day 

jasmine will grow in areas receiving from about 

800 to 3000 mm of precipitation, but most 

aggressively colonizes areas that receive from 

about 1400 to 2400 mm of precipitation. It tolerates 

soils of all textures, apparently from all parent 

materials, but is most common in limestone areas 

(Florida Exotic Plants Council 2001). Day jasmine 

does not prosper on exposed subsoil nor grow on 

swampy ground. The plant is salt tolerant if not 

exposed to heavy salt spray or overwash from 

storms (Florida Exotic Plants Council 2001). 

Reproduction.—Day jasmine flowers and fruits 

year-round (Florida Exotic Plants Council 2001). 

The fruits from a Puerto Rican collection were 

sweet and slightly bitter with little other flavor and 

weighed (fresh) an average of 0.208 + 0.010 

g/fruit. The seeds weighed an average of 0.0021+

0.0005 g/seed (air dry). The fruits are much more 

variable in size (CV = 49) than the seeds (CV = 

25). There are from four to 14 seeds/fruit (Florida 

Exotic Plants Council 2001). Day jasmine seeds 

are dispersed by frugiverous birds (Neal 1965, 

Little and others 1974). Seventy-nine percent of the 

seeds from the Puerto Rican collection cited above 
185

germinated between 7 and 30 days after sowing. 

Growth and Management.—Day jasmine rarely 

reaches more than 4.5 m in height and 8 cm in 

trunk diameter (Little and others 1974). Early 

growth is relatively fast and life spans are 

relatively short (5 to 20 years). Although seeds are 

still sold by regional seed houses and ornamental 

seedlings are offered by nurseries, nothing is 

published about propagation and management. 

With a plant so aggressive, few nursery and 

planting problems seem likely. Because the species 

is a weed in many situations, control may be 

desirable. In the absence of control guidelines, 

cutting followed by herbicide treatment of the 

sprouts is suggested. 

Benefits.—Day jasmine has long been planted as 

an ornamental for its pleasing appearance, 

moderate size, ease of establishment, and fragrant 

flowers. The plant has the drawback of being 

poisonous to livestock. The leaves contain a 

calcinogenic glycoside called 1,25-

dihydroxycholecalciferol that leads to a vitamin D 

toxicity that results in elevated serum calcium and 

deposition of calcium in soft tissues. Fifteen to 30 

percent of day jasmine leaves in an animal’s diet is 

sufficient to cause symptoms (Animal Science at 

Cornell University 2001). However, the fruits of 

day jasmine are one of the three foods that make up 

the bulk of the diet of the endangered plain pigeon 

(Columba inornata) in Puerto Rico (Island 

Resource Foundation 2001). 

References 

Animal Science at Cornell University. 2001. 

Calcinogenic glycosides. http://www.ansci. 

edu/plants/toxicagents/calglyco.htm. 2 p. 

Florida Exotic Plants Council. 2001. Cestrum 

diurnum L., Solanaceae/nightshade family. 

www.fleppc.org/pdf/Cestrum%20diurnum.pdf. 

2 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Dicotyledoneae. 

Part 3. Vol. 6. Arnold Arboretum, Harvard 

University, Jamaica Plain, MA. 658 p. 

Institute of Pacific Islands Forestry. 2001. Plant 

threats to Pacific Ecosystems: Cestrum diurnum

L., Solanaceae. www.hear.org/pier/cediu.htm. 

2 p. 

Island Resource Foundation. 2001. Threatened and 

endangered birds of the insular Caribbean: plain 

pigeon. www.irf.org/binornat.htm. 14 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de 

la Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Museum 

Press, Honolulu, HI. 924 p. 
186 

Cestrum laurifolium L’Hér. dama de noche 

SOLANACEAE 

Synonyms: Cestrum citrifolium Ritz in Hoffm. 

 Cestrum alaternoides Poir. in Lam. 

 Cestrum lambertii Dunal in DC. 

John K. Francis 

General Description.—Dama de noche, also 

known as candlewood, torch, galán de monte, boisflambeau, and citronnier, is an evergreen shrub 

usually 1 to 2 m in height but sometimes reaching 

4 m. Dama de noche is a Puerto Rican name 

meaning lady of the night. It has one or a small 

number of stems arising from the root crown or 

low on the main stem. The plant tends to root 

shallowly with stout lateral roots that are hard and 

woody with a corky or fleshy bark. The stems and 

branches are slender with gray bark and grow 

vertically for 1 to 2 m and then curve horizontally. 

The alternate, glabrous leaves are shiny green on 

the upper surface, narrowly elliptic to obovate, 

with a short petiole and blades that are 4 to 15 cm 

long. The inflorescences are short axillary racemes 

with three to five yellow or greenish-yellow, fivelobed, tubular flowers 10 to 20 mm long. The fruits 

are purple to black, ovoid to subglobose berries, 6 

to 10 mm long, with a persistent calyx. There are 

one to three or more black seeds per fruit with an 

ivory spot on the ventral side (author’s observation, 

Howard 1989, Liogier 1995). 

Range.—Dama de noche is native to Cuba, 

Hispaniola, Puerto Rico, the Virgin Islands, the 

Lesser Antilles, and Trinidad (Grisebach 1963, 

Howard 1989, Liogier 1995). 

Ecology.—Dama de noche is most common in 

areas underlain by limestone rocks but also grows 

in areas of igneous rocks. The soils are well 

drained, often rocky with a wide range of textures. 

In Puerto Rico, it may be found from a little above 

sea level to about 500 m in elevation and in areas 

that receive from about 900 to 2200 mm of annual 

precipitation. Dama de noche has an intermediate 

tolerance of shade. It usually grows in the 

understory of medium- to low-density forest. The 

species grows and fruits best when small openings 

allow broken overhead sunlight to enter. Dama de 

noche does not invade disturbed areas quickly and 

so is mainly found in middle and late secondary 

forests and remnant forests. It can be quite 

common but does not form dense stands. 

Reproduction.—Dama de noche is a good fruit 

and seed producer and may yield a few to 100 or 

more fruits per season. Flowering and fruiting 

seem to be synchronized, at least within a given 

area. Fruits were observed ripening in a forest in 

the north of Puerto Rico during the dry season 

(March). A collection of fruits from that area 

averaged 0.290 + 0.012 g/fruit. Air-dried seeds 

separated from them averaged 0.0169 + 0.0004 

g/seed or 59,200 seeds/kg. Sown on commercial 

potting mix, 93 percent germinated between 14 and 

60 days after sowing. Seeds are apparently 

dispersed by birds, and seedlings are well scattered 

and not abundant. Plants resprout when cut or 

damaged. 

Growth and Management.—Dama de noche 

appears to have a moderate growth rate, probably 

not exceeding 1 m of height growth per year. No 
187

nursery, planting, or management experience has 

been published. Protection of forests from fires and 

development is recommended to encourage the 

formation and development of natural stands. 

Benefits.—Dama de noche contributes to the 

biodiversity of the forests where it grows, helps 

protect the soil, and furnishes food and cover for 

wildlife. The stems are generally too small to use 

for fuel. 

References 

Grisebach, A.H.R. 1963. Flora of the British West 

Indies. J. Cramer-Weinheim, New York. 789 p. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 
188 

Chamaebatiaria millefolium (Torr.) Maxim. fern bush 

ROSACEAE 

Synonyms: Spiraea millefolium Torr.

Sorbaria millefolium Focke

Basilima millefolium Greene

Chamaebatiaria glutinosa Rydb.

Spiraea glutinosa Fedde 

Nancy L. Shaw 

 

Drawing source: Hitchcock and others 1961 

General Description.—Fern bush or desert sweet, 

the single species in its genus, is an upright, 

multistemmed, sweetly aromatic shrub 0.3 to 2 m 

tall. Bark of young branches is brown, becoming 

smooth and gray with age. Leaves are leathery, 

alternate, simple, twice pinnately divided, stipulate, 

and clustered near the branch tips. Foliage and 

young branches are viscid and pubescent with 

simple and stellate, pointed and glandular-capitate 

hairs. Southern populations are evergreen (Phillips 

1949), while northern populations are largely 

deciduous, retaining a few leaves though winter 

and initiating new leaf growth in early spring. The 

profuse, terminal, leafy-bracteate panicles are 5 to 

20 cm long and produce abundant flowers that are 

showy, creamy white, and insect-pollinated. 

Flowers are complete, regular, and five-merous. 

Follicles are pubescent and coriaceous. Seeds are 

yellowish to brownish and linear to narrowly 

fusiform. Germination is epigeal (Hickmann 1993, 

Hitchcock and others 1961, Welsh and others 

1987). Chromosome number is x = n = 9 

(McArthur and Sanderson 1985). Variability 

within the species appears low and hybridization 

with other species has not been reported. 

Range.—Fern bush is endemic to the Great Basin, 

Colorado Plateau, and surrounding areas. It occurs 

east of the Cascade and Sierra Nevada Mountains 

from central Oregon eastward into Wyoming and 

south to southeastern California, Nevada, Utah, 

northern Arizona, and northern New Mexico 

(Hitchcock and others 1961, Phillips 1949, Welsh 

and others 1987). 

Ecology.—Fern bush grows in fissures in rock 

outcrops and cliffs and on well-drained soils of 

dry, rocky, gravelly canyons and mountain slopes 

at elevations ranging from 900 to 3,400 m 

(Hickman 1993). It is often an early successional 

species on cinder cones and basalt lava flows, but 

it also grows on soils derived from limestone and 

granite (Eggler 1941, Everett 1957, Merkle 1952). 

Fern bush occurs in isolated populations or as an 

associated species in Artemisia spp., Pinus spp.-

Juniperus spp., northern Juniperus, mountain 

brush, Populus tremuloides Michx., Pinus flexilis

James, P. ponderosa Dougl., Picea spp. A.--Abies 

spp., Pinus longaveva D. Bailey, and chaparrel 

communities (Hickman 1993, Welsh and others 

1987). 

Reproduction.—Plants flower from June to 

September (Hitchcock and others 1961, Phillips 

1949) with irrigation prolonging the flowering 

season (Shaw and Hurd 2001). Seeds ripen in 

August to October. The follicles are hand 

collected. They open as they dry, and seeds are 

extracted by screening. Shaw and Hurd (2001) 

found an average of about 3.7 million seeds/kg for 

two seed lots grown under irrigation. Fresh seeds 

are nondormant while stored seeds require a 1- to 
189

3-month prechill (Phillips 1949, Shaw and Hurd 

2001). The optimum temperature for germination 

of southwestern populations is 18 to 26 °C 

(Phillips 1949). 

Growth and Management.—Seedlings establish 

from shallow seeding on well-drained soils where 

vegetative competition is limited (Shaw and Hurd 

2001). Fall seeding or artificial prechilling is 

required for seeds of northern populations. 

Seedlings develop rapidly if adequate water is 

available. Bareroot seedlings can be lifted 

following one growing season. Irrigated plants 

begin flowering during the second year (Shaw and 

Hurd 2001). Unlike its namesake genus, 

Chamaebatia Benth., fern bush is not nodulated by 

nitrogen-fixing actinomycetes (McArthur and 

Sanderson 1985). Fern bush is a rare host of 

Phoradendron juniperinum A. Gray, the juniper 

mistletoe (Hawksworth and Mathiasen 1978). 

Benefits.—Fern bush provides cover for many 

organisms. It is sometimes browsed by mule deer 

(Odocoilus hemionus Rafinesque), sheep (Ovis 

spp.), and goats (Capra hircus), but it receives 

little use by cattle (Bos spp.). First cultivated in 

1878 (Rehder 1940), fern bush is a valued 

ornamental due to its showy flowers, long 

flowering period, and fernlike foliage (Hitchcock 

and others 1961, Phillips 1949). Mass plantings, 

xeriscapes, screens, hedges, and specimen plants 

thrive in full sun (Phillips 1949). Native 

Americans used a tea made from its leaves for 

treatment of stomachaches (Mozingo 1987). Plants 

produce cyanic acids in their tissue (Fikensher and 

others 1981). 

References 

Eggler, W.A. 1941. Primary succession on 

volcanic deposits in southern Idaho. Ecological 

Monographs 3: 277-298. 

Everett, P.C. 1957. A summary of the culture of 

California plants at the Rancho Santa Ana 

Botanic Garden. Rancho Santa Ana Botanic 

Garden, Claremont, CA. 233 p. 

Fikenscher, L.H., R. Hegnauer, and H.W.L. 

Ruijgrok. 1981. Distribution of hydrocyanic acid 

in Cormophyta 15. New observations on 

cyanogenesis in Rosaceae. Planta Medica 41: 

313-327. 

Hawksworth, F.G. and R.L. Mathiasen. 1978. 

Hosts of juniper mistletoe at Walnut Canyon 

National Monument, Arizona, USA. Great Basin 

Naturalist 38: 89. 

Hickman, J.C., ed. 1993. The Jepson manual of 

higher plants of California. University of 

California Press, Berkeley, CA. 1,400 p. 

Hitchcock, C.L., A. Cronquist, M. Ownbey, and 

J.W. Thompson. 1961. Vascular plants of the 

Pacific Northwest. Part 3: Saxifragaceae to 

Ericaceae. University of Washington Press, 

Seattle, WA. 614 p. 

McArthur, E.D. and S.C. Sanderson 1985. A 

cytotaxonomic contribution to the western North 

American rosaceous flora. Madroño 32: 24-28. 

Merkle, J. 1952. An analysis of a pinyon-juniper 

community at Grand Canyon, Arizona. Ecology 

33:385-384. 

Mozingo, H.N. 1987. Shrubs of the Great Basin. 

University of Nevada Press, Reno, NV. 342 p. 

Phillips, J. 1949. Southwestern landscaping with 

native plants. Museum of New Mexico Press, 

Santa Fe, NM. 140 p. 

Rehder, A. 1940. Manual of cultivated trees and 

shrubs. Macmillan Publishing Company, Inc., 

New York. 996 p. 

Shaw, N.L. and E.G. Hurd. 2001 Chamaebatiaria 

millefolium (Porter) Maxim. fern bush, desert 

sweet. In: F.T. Bonner and R.G. Nisley, eds. 

Woody plant seed manual. Agriculture 

Handbook. U.S. Department of Agriculture, 

Forest Service, Washington, D.C. 

http://wpsm.net/index.html. 6 p. 

Welsh, S.L., N.D. Atwood, L.C. Higgins, and S. 

Goodrich, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 
190 

Chenopodium oahuense (Meyen) Aellen. 'aheahea 

CHENOPODIACEAE 

 

Synonyms: Atriplex oahuensis Meyen, 

Chenopodium oahuense var discosperma Fosb., 

Chenopodium pekeloi Degener, I. Degener & Aellen, 

Chenopodium sandwicheum Moq. 

Randy S. Senock and Sarah A. Taylor 

 

General Description.―Chenopodium oahuense, 

known most commonly as 'aheahea or 'aweoweo, 

is a nonscented to lightly scented shrub that 

sometimes takes on the form of a small tree. Plants 

may be erect, ascending or prostrate, branched, 

and total height 0.5 to 2 m. Older branches turn 

gray and woody with age, while younger tissue 

remains mealy pubescent, thickened and somewhat 

fleshy. Leaves are rhombic to broadly deltate, 2 to 

4.2 cm long, and 1.3 to 2.5 cm wide, with both 

surfaces pubescent, the upper surface less so and 

greener. Leaves three-lobed, teeth rounded to 

obtuse, base truncate to cuneate, petioles 1.3 to 2.5 

cm long (Wagner and others 1990). The leaves are 

strongly scented when crushed (Lamb 1981). 

Flowers in small, dense clusters grouped into 

terminal, almost leafless panicles. A calyx nearly 1 

mm long, entirely surrounding the fruit at 

maturity. Seeds are light to dark brown and may be 

horizontal or sometimes vertical, about 0.8 mm in 

diameter, the surface covered with short, blunt, 

rounded projections (Wagner and others 1990). 

Range.―While the genus is pantropical in 

distribution, this species is indigenous to Hawaii. 

‘Aheahea occurs on Laysan, French Frigate 

Shoals, Necker, Nihoa, Lisianski, and all of the 

main Hawaiian Islands except Kaho‘olawe 

(Wagner and others 1990). 

Ecology.―‘Aheahea is dispersed across the full 

range of dryland habitats in Hawaii, from coastal 

to subalpine. It appears primarily as a shrub but 

has been found on Mauna Kea on the island of 

Hawaii 3.5 to 4.5 m tall (Lamb 1981). ‘Aheahea 

behaves as a colonizer on old lava flows following 

site disturbance. In many dry upper elevation 

shrublands on the island of Hawaii, ‘aheahea 

dominates woody plant density and may be found 

sharing habitat with native naio, Myoporum 

sandwicense Gray, and mamani, Sophora 

chrysophylla (Salisb.) Seem, trees (Lamb 1981). 

The US Army has documented 'aheahea as being 

the most widespread woody plant species across 

their 108,000 acre Pohakuloa Training Area. 

Following wildfire in that same area, ‘aheahea 

reestablishment was delayed for approximately 1 

year. Within 4 years, however, it had exceeded its 

preburn density approximately 15-fold (Sherry and 

others 1999). 'Aheahea appears well adapted to 

tolerate drought by dying-back during periods of 

low moisture and then rapidly growing during 

periods of higher available moisture. 

Reproduction.―Seed production is extremely 

high for this species. The inflorescence is a large 

terminal panicle composed of many small, dense 

glomerules of flowers. ‘Aheahea peak 

reproductive events appear correlated with 

favorable environmental conditions such as 

prolonged periods of available moisture. However, 

plants can be found in flower nearly any time of 

year. Chromosome number 2n = 36 (Wagner and 

others 1990). 

Growth and Management.―‘Aheahea grows 

rapidly and remains fleshy during the first several 

months of growth. In later stages, branches are 

brittle and easily damaged. However larger, older 

individuals become single-stemmed and develop 

thickened woody branches. ‘Aheahea has been 

used sparingly as an ornamental in native and 

xeriscape gardens. The parent plant produces 

hundreds of seeds, which are easily propagated. 
191

'Aheahea prefers partial to full sun with light 

watering. The species may have potential in native 

habitat restoration as a nurse species or to enhance 

microclimatic understory conditions in favor of 

woody plant establishment. 

Benefits.―The leaves and plant tips are used for 

greens, and may be eaten like spinach, wrapped in 

ti [Cordyline fruticosa (L.) A. Chev.] leaves and 

cooked on hot coals. Hawaiians used the plant to 

make a poultice for minor wounds during battle. 

Leaves were pounded or crushed and applied like 

rubbing alcohol, the healing effect due to the 

presence of chlorophyll in the leaf (Ching 2002). 

Hawaiians also used the dark red bark pounded in 

conjunction with other ingredients to create a 

concoction that would enhance and beautify the 

skin of a newborn baby when ingested by a 

pregnant or nursing mother (Anonymous 1996). 

Two limbs of the shrub wood were connected to 

create a "makua mano," or shark hook, and in this 

fashion, 'aheahea was used to catch sharks (Ching 

2002). The native shrub is also of great importance 

to a certain rare beetle, Rhyncogonus biformis, that 

lives on Necker Island. The beetle is nocturnal and 

emerges at night where it feeds soley on the leaves 

of the 'aheahea (Kawaharada 2002). 

Detrimental Effects.―The flower heads are host 

plants for Nysius nemorivagus, a white Lygaeid 

bug that can become a nuisance pest for certain 

agricultural crops such as cabbage, cucumbers, 

potatoes and all types of squash (Kessing and 

others 1993). 

References 

Anonymous. 1996. Hawaiian use of native plants. 

http://www.k12.hi.us/~waianaeh. 

hawaiianstudies/nplants6.html 

Ching, F. 2002. Ulu Ka Hoi. http://kms. 

kapalama.ksbe.edu/projects/yellow/default. 

html. [not paged]. 

Kawaharada, D. 2002. Visiting Mokumanamana. 

http://explorers.bishopmuseum.org. [not paged]. 

Kessing, J., L. Martin, and R.FL Mau. 1993. 

http://www.extento.hawaii.edu/kbase/crop/ 

Type/Nysius.htm 

Lamb, S.H. 1981. Native trees and shrubs of the 

Hawaiian Islands. The Sunstone press, Santa Fe, 

New Mexico. 158 p. 

Sherry, K., J.M. Castillo, and R.B Shaw. 1999. 

Effects of wildfire on vegetation and rare plants 

in arid montane shrublands. Pohakaloa training 

area, Hawaii. 1999 Hawaii Conservation 

Conference, Honolulu, HI. 

Wagner, W.L., D.R. Herbst, and S.H. Sohmer. 

1990. Manual of the Flowering Plants of 

Hawaii. University of Hawaii Press, Honolulu, 

HI. 1,854 p. 
192 

Chiococca alba (L.) A.S. Hitchc. West Indian snow-berry

RUBIACEAE 

Synonyms: Lonicera alba L. 

 Chiococca racemosa L. 

John K. Francis

General Description.—West Indian snow-berry, 

is also known as David’s root, bejuco de berac, 

buenda, liane des sorciers and several other 

common names (Howard 1989). It is one of the 

most common woody plants in Puerto Rican moist 

forests. The species grows as a scrambling shrub 

or woody vine that often climbs taller vegetation 

and may reach 6 m of extension (AcevedoRodríguez and Woodbury 1985). The slender, 

green stems and branches are four-angled to round 

in cross section. Opposite branching occurs at the 

nodes where one or two shoots develop. A vertical 

shoot at each node commonly ascends from 

prostrate stems. The root system consists of 

shallow laterals and abundant fine roots. West 

Indian snow-berry has opposite, thin to coriaceous, 

dark green leaves that are ovate or oblong and 

pointed at the tip and pointed to rounded at the 

base. The petioles are 2 to 10 mm long and tiny 

lobbed stipules are present (Liogier 1997). 

Range.—West Indian snow-berry grows naturally 

in Florida, the Bahamas, through the Greater and 

Lesser Antilles, in Mexico, Central America, and 

northern South America. The species is not known 

to have naturalized outside its native range. 

Ecology.—West Indian snow-berry grows in moist 

and dry forests from near sea level to 700 m or 

more in all types of topography. It will grow on 

soils of all textures derived from both sedimentary 

and igneous parent materials including ultramafic 

rocks (serpentine). Very poorly drained and highly 

saline conditions are not tolerated. The species may 

be found in remnant forests in both natural 

openings and the understory. It invades brushy 

pastures and roadsides and appears early in the 

reforestation process. It is intermediate in tolerance 

to shade. West Indian snow-berry competes well 

with weeds and brush, but does not prosper in 

dense grass swards. The species does not appear to 

be seriously affected by any insect or disease. 

Reproduction.—West Indian snow-berry flowers 

in Puerto Rico from June through October and 

yields ripe fruits from June through December 

(Acevedo-Rodriguez and Woodbury 1985). The 

small white, cream, or yellow flowers are borne in 

racemes that arise from the leaf axils. The globose 

but slightly flattened fruits are white and vary in 

size (4 to 8 mm) and weight (by a factor of 3) in 

the same inflorescence. The fruits are sweet with 

little other flavor. The average fruit weight from a 

Puerto Rico collection was 0.117 + 0.003 g. The 

dark brown seeds, which averaged about two per 

fruit, weighed 0.004 + 0.000 g. Germination is 

epigeal. For the above sample, germination was 48 

percent and occurred between 33 and 59 days after 

sowing. Stems root when they come in contact with 

the soil, which results in interconnected patches 

and many independent plants. Asexual 

reproduction by air layers and rooted cuttings 

should be easy. Long distance dispersal appears to 

be principally done by birds. 

Growth and Management.—West Indian snowberry plants under moderate canopy openings grow 

up to 0.5 m per year in each of its many stems. 
193

Plants develop slowly in the nursery, requiring 3 

months to grow large enough to prick into pots and 

about an additional year to reach outplanting size. 

Seedlings and sections of older plants can be dug 

up in the forest and transplanted successfully 

(Workman 1980). Once established, West Indian 

snow-berry needs little maintenance. 

Benefits.—West Indian snow-berry is used to a 

limited extent as an ornamental in naturalistic 

landscaping (Workman 1980) and is sometimes 

cultivated as a flowering vine in greenhouses 

(Bailey 1941). Being very common and 

widespread, the species contributes to the 

biodiversity of many Neotropical forests. It is 

grazed by cattle, goats and certainly wild ruminants 

and produces fruits that are eaten by birds and 

other animals. The roots of West Indian snowberry are used as a purgative, diuretic, vomitive, 

and antidiarrhetic in herbal medicine (Liogier 

1990). 

References 

Acevedo-Rodriguez, P. and R.O. Woodbury. 1985. 

Los bejucos de Puerto Rico. Vol. 1. General 

Technical Report SO-58. U.S. Department of 

Agriculture, Forest Service, Southern Forest 

Experiment Station, New Orleans, LA. 331 p. 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. Vol. 1. The MacMillan Company, 

New York, NY. 1,200 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Dicotyledoneae. 

Part 3. Vol. 6. Arnold Arboretum, Harvard 

University, Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p.

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Workman, R.W. 1980. Growing native. The 

Sanibel-Captiva Conservation Foundation, Inc., 

Sanibel, FL. 137 p. 
194 

Chromolaena geraniifolia (Urban) King & H.E. Robins. geranium-leaf eupatorium 

ASTERACEAE 

Synonyms: Osmia geraniifolia (Urban) Britt. & Wilson 

 Eupatorium geraniifolium Urban 

John K. Francis 

General Description.—Geranium-leaf eupatorium, a name assigned by the author in the absence 

of a local common name, is an evergreen shrub up 

to 2 m in height and 1 cm in stem diameter. The 

plant usually has several stems arising from the 

root crown. The root system is composed of fibrous 

lateral and fine roots. Geranium-leaf eupatorium 

produces relatively few branches. The foliage, 

which has a chrysanthemum-like odor, tends to be 

concentrated near the branch ends. The opposite 

leaves, which have a short petiole, are densely 

pubescent, 2 to 6 cm long by 1.5 to 5 cm broad, 

and deeply lobed or coarsely toothed. 

Inflorescences are small terminal corymbs of 

oblong-cylindrical heads. The corolla of the small 

flowers is blue. The fruits are dark-colored 

achenes, 4 to 5 mm long with a pappus 4 mm long 

(author’s observation, Liogier 1997). 

Range.—Geranium-leaf eupatorium is endemic to 

Puerto Rico. It occurs mainly in mountainous areas 

of the central part of the island (Liogier 1997). It is 

not known to have been planted or naturalized 

elsewhere. 

Ecology.—Geranium-leaf eupatorium can be 

locally common in scattered populations. It may be 

found on a wide variety of soils but most 

frequently is found on well-drained, mediumtextured, often rocky soils that develop over 

igneous, sedimentary, and metamorphic (including 

ultramafic) rocks. The species is most common in 

areas of volcanic rock. Geranium-leaf eupatorium 

grows on sites that receive from about 1200 to 

about 3000 mm of mean annual precipitation at 

elevations of 400 to 1,000 m (Liogier 1997). The 

species is moderately intolerant of shade; it grows 

well in openings and under the canopy of low 

basal-area forest. Geranium-leaf eupatorium occurs 

in remnant and middle- to late-secondary forests, 

old road cuts and fills, bluffs, and unstable slopes. 

It grows as scattered individual plants and in small 

clumps under favorable conditions. 

Reproduction.—Geranium-leaf eupatorium was 

observed in bloom in March by the author. It 

flowers and fruits abundantly. Seeds collected near 

Cayey, Puerto Rico, averaged 0.00033 g/seed or 31 

million seeds/kg. These seeds were sown on moist 

filter paper and yielded 67 percent germination 

between 7 and 24 days after sowing. The seeds are 

dispersed by the wind. Seedlings and successful 

natural reproduction are not common. 

Growth and Management.—Individual stems of 

geranium-leaf eupatorium live about 5 years. Plants 

live much longer because of sprouting that replaces 

senescent stems. No management guidelines have 

been published. Planting is probably the best way 

of establishing new populations, and natural 

reproduction probably can be encouraged by 

creating openings with scattered patches of bare 

soil in forests near seed sources. 

Benefits.—Geranium-leaf eupatorium is a pretty 

plant and contributes to the aesthetic appeal of the 

forest. It helps stabilize the soil and furnishes cover 

for wildlife. 
195

References 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 
196 

Chromolaena odorata (L.) King & H.E. Robins. Christmas bush 

ASTERACEAE 

Synonyms: Eupatorium odoratum L. 

 Eupatorium conyzoides Vahl 

 Eupatorium brachiatum Sw. ex Wikstr. 

 Eupatorium atriplicifolium Vahl 

 Osmia odorata (L.) Schultz-Bip. 

John K. Francis 

General Description.—Christmas bush, also 

known as bitter bush, Siam weed, baby tea, 

cariaquillo, Santa María, and fleurit-Noël, is a 

scrambling shrub (Howard 1989, Liogier 1997). It 

may reach 1 m or more as a free standing shrub and 

4 m or more when climbing into trees or shrubs. 

Stems reach 2 cm in diameter. The plants are 

maintained by a system of abundant, yellowish, 

fine lateral roots. Multiple sprouts arise from the 

root crown and lower stems. The individual 

branches are long with relatively few branches. 

Foliage occurs only on recent growth. The 

opposite, three-nerved leaves are deltoid to ovatelanciolate, usually with a dentate margin and a long 

pointed tip. The leaves are aromatic when crushed. 

The inflorescences are corymbs of cylindrical 

heads located on the terminals of lateral branches. 

There are 15 to 25 tubular florets per head, white, 

lavender, pink, or blue in color. The seeds are a 

brownish gray to black achene that is 4 mm long 

with a pale brown pappus 5 or 6 mm long (Howard 

1989, Liogier 1997). 

Range.—Christmas bush is native from Florida 

through the West Indies and from Texas through 

Central and South America to Argentina (Howard 

1989, Liogier 1997). It has been accidentally or 

deliberately introduced and has naturalized 

throughout much of the tropics, including Guam 

and Hawaii (Pacific Island Ecosystems at Risk 

2001). 

Ecology.—Christmas bush grows from near sea 

level to over 1,000 m in elevation (Binggeli 1999). 

It thrives on all types of well-drained soil and can 

grow on soils relatively low in fertility. 

Disturbance is required before a site can be 

colonized (Pacific Island Ecosystems at Risk 

2001). Once established, Christmas bush competes 

aggressively with herbs, grass, and shrubs in open 

areas. In its native range, it is frequently seen on 

roadsides, riverbanks, vacant lots, abandoned 

farmland, and neglected pastures. Christmas bush 

has found a particular niche in the slash-and-burn 

agriculture cycle. In Borneo, Christmas bush and 

other perennial grasses and shrubs invade within 3 

years of abandonment and are gradually replaced 

by trees (Ohtsuka 1999). The species is not shadetolerant and will not grow under a closed forest 

stand. It is also intolerant of frost (Binggeli 1999) 

and is limited by drought (below about 900 mm of 

mean annual precipitation). 

Reproduction.—Christmas bush blooms annually 

and is an abundant producer of seeds. Flowering 

and fruiting begins after plants are 1 year old 

(Binggeli 1999). The flowers are pollinated by 

insects. The small fruits mature in about a month 

(Binggeli 1999). One collection of seeds in Puerto 

Rico averaged 2,670,000 seeds/kg but did not 

germinate. A second collection averaged 1,560,000 

seeds/kg and gave 11 percent germination between 

3 and 120 days after sowing. Germination is 

epigeal. The seeds are wind-dispersed, and 

transport by animals is possible because of small 

hooks on the seeds. In India, it was observed that 

only about 1.4 percent of the first-year seedlings 

survived into the second year (Binggeli 1999). 

Stems root whenever they come in contact with the 

ground. 

Growth and Management.—Individual stems last 

about 2 years and die back to or near the base and 

are replaced by new sprouts. Plants easily survive 

cutting and fire. The best current control method is 
197

mechanical or hand cutting followed by herbicide 

treatment. Partial control can be obtained through 

the use of aggressive cover crops. Relatively good 

biological control has been obtained with 

Pareuchaetes pseudooinsulata Rego Barros 

(Lepidoptera) in Guam and several other Pacific 

islands (Pacific Island Ecosystems at Risk 2001). 

Detriments and Benefits.—Invasion of Christmas 

bush has been disastrous by seriously suppressing 

native species in disturbed forests and pastures in 

the tropics outside its native range. The shrub is 

reported to be highly allelopathic to nearby 

vegetation (Muniappan 1994), a fact that has been 

demonstrated in controlled studies (Sahid and 

Sugau 1993). Christmas bush reduces the diameter 

growth of teak in infested plantations (Daryono and 

Hamzah 1979). It was thought to be useful in the 

control of Imperata grass and for this reason was 

deliberately introduced into the Ivory Coast, but 

the results were disappointing (Binggeli 1999). 

Because of the abundance of dead leaves and dry 

shoots, Christmas bush stands are a fire hazard 

(Muniappan 1994). Cattle do not eat Christmas 

bush; however, it is browsed by white-tailed deer 

(Meyer and others 1984). In herbal medicine, leaf 

extracts with salt are used as a gargle for sore 

throats and colds. It is also used to scent aromatic 

baths (Liogier 1990). Extracts of Christmas bush 

have been shown to inhibit or kill Neisseria 

gonorrhoeae (the organism that causes gonorrhoea) 

in vitro (Caceres and others 1995) and to accelerate 

blood clotting (Triratana and others 1991). A 

satisfactory medium-density particleboard was 

prepared from Christmas bush stems (Kaleta and 

others 1999). During fallows between cultivation, 

Christmas bush adds copious amounts of organic 

matter to the soil and may reduce the populations 

of nematodes (M’Boob, 1991). It is also useful as 

mulch for row crops (Swennen and Wilson 1984). 

References 

Binggeli, P. 1999. Chromolaena odorata (L.) King 

& Robinson (Asteraceae). http://members.tripod. 

co.uk/WoodyPlantEcology/docs/web-sp4.htm. 

4 p. 

Caceres, A., H. Menendez, E. Mendez, E. 

Cohobon, B.E. Samayoa, E. Jauregui, E. Peralta, 

and G. Carrillo. 1995. Antigonorrheal activity of 

plants used in Guatemala for the treatment of 

sexually transmitted diseases. Journal 

Ethnopharmacol 48(2): 85-88. 

Daryono, H. and Z. Hamzah. 1979. A study of 

Eupatorium odoratum as a weed in teak 

(Tectona grandis) forest. Lembaga Penelitian 

Hasil hutan 312. Lopora, Indonesia. 25 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Kaleta, D., S.R. Ghosh, and C.N. Saikia. 1999. 

Medium density particle board from weeds. 

Journal of Scientific and Industrial Research 

58(9): 705-710. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

M’Boob, S.S. 1991. Preliminary results of a 

survey and assessment of Chromolaena odorata

(Siam weed) in Africa. Biotropica Special Pub. 

44: 51-55. 

Meyer, M.W., R.D. Brown, and M.W. Graham. 

1984. Protein and energy content of white-tailed 

deer diets in the Texas coastal bend. Journal of 

Wildlife Management 48(2): 527-534. 

Muniappan, R. 1994. Chromolaena odorata (L.) 

R.M. King and H. Robinson. In: R. Labrada, 

J.C. Caseley, and C. Parker, eds. Weed 

management for developing countries. Plant 

Production and Protection Paper 120. Food and 

Agriculture Organization of the United Nations, 

Rome. 93-94. 

Ohtsuka, T. 1999. Early stages of secondary 

succession on abandoned cropland in north-east 

Borneo Island. Ecological Research 14(3): 281-

290. 

Pacific Island Ecosystems at Risk. 2001. Invasive 

plant species: Chromolaena odorata (L.) King 

& Robinson, Asteraceae. 

http://www.hear.org/pier/chodo.htm. 3 p. 

Sahid, I.B. and J.B. Sugau. 1993. Allelopathic 

effects of lantana (Lantana camara) and siam 

weed (Chromolaena odorata) on selected crops. 

Weed Science 41(2): 303-308. 
198 

Swennen, R. and G.F. Wilson. 1984. In-situ mulch 

production for plantain. Banana Newsletter 7: 

20-22. 

Triratana, T., R. Suwannuraks, and W. 

Naengchomnong. 1991. Effect of Eupatorium 

odoratum on blood coagulation. Journal of 

Medical Association of Tailand 74(5): 283-287.
199

Chrysobalanus icaco L. coco-plum 

CHRYSOBALANACEAE 

Synonyms: Chrysobalanus pellocarpus G.F.W. Meyer 

 C. icaco L. var. genuinus Stahlé & Quentin 

 C. icaco L. var. pillocarpa (G.F.W. Meyer) C. Martius 

 John K. Francis 

 

General Description.—Coco-plum, also known as 

icaco, icaque ponne, pork-fat-apple, zicate, and 

many other common names, is a medium-sized 

coastal shrub or, rarely, a small tree. It generally 

has multiple brown or gray stems that are smooth 

to scaly. The twigs are green and hairless when 

young and turn reddish brown with raised lenticels. 

The shrub’s habit may be creeping or erect. The 

branches support many shiny, dark-green, leathery, 

round, or elliptic leaves 3.8 to 8.2 cm in length and 

2.5 to 5.7 cm in width. The under surfaces of the 

leaves are light green. The petiole is about 3 mm 

in length. The simple leaves are alternate in two 

rows, turned upward (Little and others 1974). 

Range.—Coco-plum is native to coastal areas of 

southern Florida and the Bahamas through the 

Caribbean. It is also found along the coasts of 

Mexico, through Central America and South 

America, to Ecuador and northern Brazil. The 

range has been extended inland in those areas by 

disturbance and planting (Little and others 1974). 

Chyrysobalanus orbicularis Schum., C. ellipticus

Soland. ex Sabine, and C. atacarensis A. Chev. 

from Africa were all once considered to be 

subspecies of coco-plum (Paradis 1987). Probably 

all the African references to C. icacos are really 

one of these species. There is disagreement among 

taxonomists as to whether varieties exist among 

coco-plumb (Howard 1988, Liogier 1985). At 

most, variation among populations is minor. 

Ecology.—Coco-plum is a coastal species. It 

commonly grows as single plants or thickets on 

dunes and rocky headlands. It may also be found 

on shallow soils in moist areas up to a 450-m 

elevation in Puerto Rico (Little and others 1974). 

Although the species can survive a great deal of 

stress from storms, salt spray, and flooding, it is 

low in stature, relatively intolerant of shade, and 

only persists where competing vegetation is short. 

Scale insects and caterpillars sometimes damage 

natural and ornamental plants in Florida (VargasSimon and others 1997). 

Reproduction.—The flower clusters (cymes) are 

shorter than the leaves and borne at the bases of the 

leaves near the ends of the branches. Several 

greenish-white flowers less than 1 cm in diameter 

compose the clusters. Coco-plum flowers and fruits 

nearly throughout the year. The white to purple 

fruits are drupes that resemble plumbs. Their thin 

flesh is spongy, whitish, and slightly sweet to 

almost tasteless when ripe. Immature fruits are 

astringent (Little and others 1974). The fleshiness 

of the mesocarp of fruits varies considerably across 

its range (Howard 1988). One large (2.5 m tall) 

coco-plum plant growing under a Pinus caribaea 

Morelet plantation in Puerto Rico yielded 760 ripe 

fruits in a single picking. Ripe fruits from another 

Puerto Rican collection weighed 4.36 + 1.17 g 

(mean and standard deviation), and seeds averaged 

1,790/kg in one Puerto Rican collection. This seed 

lot gave 89 percent germination beginning 34 days 

after sowing sample (Francis and Rodríguez 1993). 

No scarification or other seed treatment is needed. 
200 

Seed dispersion is presumed to be by gravity, 

water, birds, bats, domestic animals, and humans. 

Natural reproduction may be sparse to abundant. 

Artificial reproduction is usually by seeds with 

plants grown as potted seedlings. Apical and basal 

semiwoody, leafy cuttings treated with hormones 

will root in 6 to 8 weeks in mist bed conditions 

(Vargas-Simon and others 1997). The best 

treatment tested was Indol Acidic Acid (IAA) at 

5,000 ppm applied to apical stem cuttings. 

Adventitious roots arose from the vascular 

cambium, and lateral roots arose from the 

paricycle. 

Growth and Management.—Growth is slow in 

the nursery and the field, so new plantings of cocoplum must be protected from both herbaceous and 

woody competition. 

Benefits.—The fruits are edible raw and can be 

made into preserves. Coco-plum seeds, which have 

a high oil content, are also edible. The wood is 

light brown, hard, and heavy (specific gravity 0.8) 

and is used for fuel and rustic construction (Little 

and others 1974). Various parts of the plant have 

been used in folk medicine. The species is known 

to have hypoglycemic effects (Costa 1977). It is a 

honey plant and furnishes food for wildlife. Cocoplum is used in the Eastern and Western 

Hemespheres as an ornamental. Another important 

benefit from the species is for dune and soil 

stabilization. 

References 

Costa, O. de A. 1977. Brazilian plants with 

hypoglycemic effects. Leandra 6-7: 7, 63-75. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station. New 

Orleans, LA. 5 p. 

Howard, R.A. 1988. Flora of Lesser Antilles. Vol. 

4. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 673 p. 

Liogier, H.A. 1985. Discriptive flora of Puerto 

Rico and adjacent islands. Vol. 1. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 352 

p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Paradis, G. 1987. Concerning the division of the 

genus Chrysobalanus into species and 

subspecies in West Africa (Chrysobalanaceae). 

A Sciences Naturelles 45(3,4): 246-254. 

Vargas-Simon, G., G. Arellano-Ostoa, and E. 

García-Villanueva. 1997. Propagación por 

estacas con hojas de icaco (Chrysobalanus icaco

L.) y anatomía del enraizamiento. Proceedings of 

the International Society for Tropical 

Horticulture 41: 264-269. 
201

Chrysophyllum oliviforme L. satinleaf 

SAPOTACEAE 

Synonyms: Chrysophyllum microphyllum Jacq. 

 Chrysophyllum monopyrenum Sw. 

 Cynodendron oliviforme (L.) Baehni 

John K. Francis 

General Description.—Satinleaf, also known as 

wild star-apple, damson plum, saffron-tree, 

caimitillo, caimitillo de perro, camitillo cimarró, 

teta de burra, macanabo, and caïmite marron, is a 

beautiful, evergreen shrub or small tree usually 3 to 

5 m in height. Under favorable conditions, older 

individuals occasionally reach 10 m or more in 

height and 30 cm in diameter at breast height. The 

single or multiple stems have gray-brown bark with 

fissures and plates. Twigs are slender and rusty or 

reddish-brown. The alternate leaves are elliptic, 

ovate or oblong, 3 to 11 cm long and 2 to 5 cm 

broad, with shiny, dark-green, hairless upper 

surfaces and coppery under-surfaces owing to a 

dense covering of short hairs. The bell-shaped 

flowers are one to many on short pedicels in the 

leaf axils. The 3- to 5-mm, five-lobbed corolla is 

yellowish- or greenish-white. The fleshy fruits are 

dark-purple, blue, or black, elliptical, one-seeded 

berries 1 to 2 cm long. They have whitish flesh 

surrounded by a gummy or rubbery skin that 

secretes a white milky sap when cut. The fruits are 

sweet and edible. Satinleaf has 2n = 52 

chromosomes (Gilman and Watson 1993, Liogier 

1995, Little and Wadsworth 1964, Long and 

Lakela 1976, Nelson 1996). 

Range.—Satinleaf is native to Florida, the 

Bahamas, the greater Antilles, and Belize (Liogier 

1995). The species is planted in many tropical 

countries as an ornamental and has become 

naturalized in Hawaii and French Polynesia 

(Pacific Island Ecosystems at Risk 2002). It has 

been declared endangered in the wild in Florida 

(Crowley 2002). 

Ecology.—Satinleaf grows on a wide variety of 

soils in all textures and in pH’s from about 5 to 8, 

and has a moderate tolerance to salt (Institute of 

Food and Agricultural Sciences 2002). It grows on 

coastal sands and shallow clays over limestone in 

Puerto Rico, in pinelands and hammocks of the 

everglades and keys of Florida, and in lowelevation moist, secondary forests and Eucalyptus

plantations in Florida (Little and Wadsworth 1964, 

Pacific Island Ecosystems at Risk 2002). Satinleaf 

occurs in remnant and middle secondary forests. 

When young, it demonstrates an intermediate 

tolerance to shade and can grow and develop in the 

understory of relatively low-density forests. To 

bear fruit, it must have increased light, as in 

intermediate and codominant crown positions. 

Satinleaf plants are severely damaged by 

temperatures of 0 °C and below. They are resistant 

to drought and storm damage (Gilman and Watson 

1993). 

Reproduction.—Puerto Rican satinleaf plants 

flower July to October and mature fruits in 

February (Little and Wadsworth 1964). They are 

reported to flower and fruit all year in Florida 

(Long and Lakela 1976). The flowers are insect 

pollinated. Seeds are dispersed by frugivorous 

birds (Pacific Island Ecosystems at Risk 2002). 

Seedlings are relatively common in suitable habitat 

in Puerto Rico. Seed weights and treatment 
202 

information is not available. Misted hardwood 

cuttings and air layers, both treated with 0.3 

percent IBA (indol-butyric-acid), were tested as 

means of propagating the species vegetatively. 

Thirty-five of 69 cuttings rooted in 7 months and 

one of 39 air layers rooted in 6 months (author’s 

observation). Ornamental production is by seed and 

semihardwood cuttings under mist (Gilman and 

Watson 1993). 

Growth and Management.—Growth is relatively 

slow at all stages, and the plants are relatively 

long-lived. Natural reproduction can probably be 

encouraged by disturbance under low-density 

secondary forests near seed sources. 

Benefits.—Satinleaf contributes to the aesthetics of 

the forests where it grows, helps protect the soil, 

and furnishes food and cover for wildlife. It was 

listed as an important source of nectar for 

honeybees in the Dominican Republic (Marcano 

Fondeur 1973). The foliage is probably unpalatable 

to ungulates; at least the key deer (Odocoilleus 

virginianus ssp. clavium) will not eat it (Schaus 

and others 2002). The fruits are edible and 

sometimes used to make jelly (Little and 

Wadsworth 1964). In areas of infestation, satinleaf 

is usually heavily attacked by Mediterranean fruit 

fly (Ceratitis capitata (Wiedemann) (Department 

of Entomology and Nematology 2002). The wood, 

which has a specific gravity of 0.9, is hard, heavy, 

and strong. It is used for construction in Cuba 

(Little and Wadsworth 1964). Satinleaf has been 

extensively, but not intensively, planted as an 

ornamental in Florida, Hawaii, and elsewhere. It 

makes a pretty addition to natural landscaping, a 

good foundation plant, and an attractive lawn, 

street, and parking lot tree. Pruning may be 

necessary to maintain a single stem and to remove 

drooping branches (Gilman and Watson 1993). 

References 

Crowley, M. 2002. Florida’s native plants—

protected. Native Florida Heritage Web Ring. 

http://nsis.org/garden/pr-plants-c.html. 5 p. 

Department of Entomology and Nematology. 

2002. Mediterranean fruit fly—Ceratitis 

capitata (Wiedemann). University of Florida. 

http://creatures.ifas.ufl.edu/fruit/Mediterranean_

fruit-fly.htm. 19 p. 

Gilman, E.F. and D.G. Watson. 1993. 

Chrysophyllum oliviforme, Satinleaf. Fact Sheet 

ST-166. Environmental Horticulture 

Department, Florida Cooperative Extension 

Service, University of Florida. 2 p. 

Institute of Food and Agricultural Sciences. 2002. 

Flowering and shade trees (A-E). University of 

Florida Extension http://miami-dade.ifas.ufl. 

edu/programs/fyn/publications/dtplfloweringtree

s-a-e.htm. 7 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agric. Handb. 249. U.S. Department of 

Agriculture, Washington, DC. 548 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Marcano Fondeur, E. de J. 1973. La flora apícola 

de la República Dominicana. http.// 

marcano.freeservers.com/nature/estudios/apicola

/dicotsp.htm. 11 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Pacific Island Ecosystems at Risk. 2002. 

Chrysophyllum oliviforme L., Sapotaceae. 

http://www.hear.org/pier3/choli.htm. 2 p. 

Schaus, C., S. Wade and J. Dunan. 2002. Key deer 

and plants they won’t eat. Monroe County 

Extension Service, University of Florida. 

http://monroe.ifas.ufl.edu/key_deer_plants.htm. 

4 p. 
203

Chrysothamnus nauseosus (Pallas ex Pursh) Britt. rubber rabbitbrush 

ASTERACEAE 

Synonyms: Bigelovia nauseosa Gray 

Chrysocoma nauseosa Pallas ex Pursh 

 Ericameria nauseosa (Pallas ex Pursh) Nesom & Baird 

E. Durant McArthur and Jeffrey R. Taylor 

General Description.—Rubber rabbitbrush also 

know as common rabbitbrush is a shrub typically 

0.3 to 2.0 m in height. Some types, however, are 

smaller while others exceed heights of 3.0 m. 

Usually several erect stems arise from the base and 

branch to create a rounded form. Branches are 

covered with a felt-like tomentum, permeated by a 

resinous gum, which can vary in color from green, 

yellow-green, gray-green, to white. Leaf shape can 

vary from filiform to broadly linear, and leaf 

length can range from 18 to 63 mm. Like 

branches, leaves are typically covered with 

tomentum. The flower heads are generally 

arranged into a cymose infloresence. Involucral 

bracts in each head may vary from glabrous to 

highly tomentose and number between 20 and 25. 

Chrysothamnus nauseosus is uniformly diploid, x 

= 9, 2n = 18 (Anderson 1966). 

Taxonomy.—The International Plant Names 

Index (IPNI) recognizes 30 subspecies of C. 

nauseosus (IPNI. 2002). Some sytematists 

promote the transfer of rubber rabbitbrush to the 

genus Ericameria (Anderson 1995). Subspecies 

may vary by morphology, color, and secondary 

metabolic compounds. Each subspecies is likely 

the result of reproductive isolation inbreeding, 

although subspecific sympatry has been observed. 

Some of the more common subspecies include ssp. 

albicaulis, hololeucus, consimilis, graveolens, and 

salicifolius (McArthur and others 1979). White 

stem rubber rabbitbrush (C. n. ssp. albicaulis and 

hololeucus) vary from other subspecies by their 

dense white to grayish tomentum, the size and 

arrangement of involucral bracts (five distinct 

vertical ranks, 8 to 10 mm in length), and the 

corolla lobe length (1 to 2 mm in ssp. albicaulis, 

0.5 to 1 mm in ssp. hololeucus). These two 

subspecies are segregated by elevation. Threadleaf 

rubber rabbitbrush (C. n. consimilis) is a tall form, 

up to 3.0 m, with green to yellow-green tomentum 

and thread-like leaves. Ecologically, spp. 

consimilis is more likely to be found in alkaline 

valley bottoms. Green rubber rabbitbrush (C. n.

ssp. graveolens) has yellow-green to green stems 

and is less pubescent than other subspecies. 

Mountain rubber rabbitbrush (C. n. ssp. 

salicifolius) has larger leaves (4 to 8 cm long and 3 

to 10 mm wide), less uniform involucaral bracts, 

and is one of the more palatable subspecies to 

livestock and big game. 

Range.—Rubber rabbitbrush is common 

throughout the Intermountain West extending from 

British Colombia to Saskatchewan and south to 

western Texas, Sonora, Baja California, and 

eastern California (Anderson 1986). It inhabits 

plains, valley bottoms, and foothills and is often 

associated with sagebrush, pinyon-juniper, and 

ponderosa pine communities. 
204 

Ecology.—Rubber rabbitbrush establishes readily 

on disturbed areas and can often become the 

dominant vegetation on lands that have been 

disturbed by fire, insects, vehicles, or heavy 

grazing. The species spans a broad elevation 

gradient, extending from 150 to 2,700 m. with 

annual precipitation generally in the 250 to 350 

mm range. This shrub grows well in sandy, 

gravelly, or clay-alkaline soils, depending on the 

subspecies or ecotype. Some subspecies have 

broader ecological amplitudes than others, 

allowing them to establish in a variety of habitats 

(McArthur and others 1979). Although it grows 

vigorously, it does not exclude or inhibit 

herbaceous species. On the contrary, production of 

herbaceous cover was greater where rubber 

rabbitbrush was present than when it was not 

present (Frischknecht 1963). 

Reproduction.—Rubber rabbitbrush blooms from 

August to October. Seeds mature beginning in 

mid-October until late fall. In most conditions, 

rubber rabbitbrush is a heavy seed producer. Seed 

germination in rubber rabbitbrush is quite high, 

and seed viability remains relatively high (65 

percent) for up to 3 year’s storage under ordinary 

warehouse storage conditions (Stevens and others 

1981). When exposed to cool night (4.7 o

C) and 

warm day temperatures, seed germinates in about 

2 days (Weber and others 1985). Under 

standardized single cool temperature conditions, 

however, seeds from lower elevations and latitudes 

germinate more quickly (2 weeks as compared to 

4) than those from higher elevations and latitudes 

(McArthur and others 1987, Meyer and others 

1989). Seed (achene) size in rubber rabbitbrush 

appears to be correlated to habitat; subspecies 

adapted to sandy sites have larger achenes (Meyer 

1997). There are about 1,530 cleaned seeds per 

gram (McArthur and others 1979). 

Growth and Management.—Once established, 

wind-disseminated achenes allow rubber 

rabbitbrush to spread easily and quickly (Meyer 

1997, Young and others 1984). It is fast growing, 

reaching maturity in 2 to 4 years but relatively 

short-lived shrub, usually 5 to 20 years (personal 

observation). It can be aerially seeded. Rubber 

rabbitbrush can be troublesome on rangelands 

when it invades and occupies areas at high 

densities. It is a difficult to control because it is 

resistant to herbicides and has the ability to 

resprout from the crown (Cook and others 1965). 

Benefits.—Rubber rabbitbrush can be an 

important structural and ecological component in 

stable and disturbed plant communities. Its ability 

to stabilize soil and reduce wind and water erosion 

preserves habitat for other shrubs, forbs, and 

grasses. This shrub is an excellent plant for erosion 

control because it has deep roots, produces heavy 

litter, and is able to establish on severe sites 

(Aldon and Pase 1981, McArthur and others 1995, 

USDA 1937). Rubber rabbitbrush is browsed by 

deer, pronghorn, and by domestic animals, but 

palatability varies greatly between subspecies and 

ecotypes. The white to grayish subspecies 

(albicaulis, hololoeucus, and salicifolius) are more 

palatable than the green subspecies (gravelens and 

consimilis) (Goodrich and others 1999 Hanks and 

others 1975). Unlike some forage shrubs, however, 

rubber rabbitbrush is quite resilient to browsing. 

In addition to food, C. nauseosus also provides 

cover for small mammals, birds, and newly born 

pronghorn (Yoakum 1986). In addition to its 

ecological importance, people have also 

recognized the chemical properties of C. 

nauseosus. Rubber rabbitbrush has been 

investigated as a possible source of rubber and 

other chemicals (Hall and Goodspeed 1919; 

Hegerhorst and others 1987). Some accessions, for 

example, produce up to 6 percent stem rubber and 

20 percent resin. Finally, some forms have 

ornamental shrub potential (McArthur and others 

1979, Weber and others 1985). 

 

References 

Aldon, E.F. and C.P. Pase. 1981. Plant species 

adaptability on mine spoils, in the Southwest: a 

case study. Research Note RM-398. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Forest and Range Experiment 

Station, Fort Collins, CO. 3 p. 

Anderson, L.C. 1966. Cytotaxonomic studies in 

Chrysothamnus (Asterae, Compositae). 

American Journal of Botany 53: 204-212. 

Anderson, L.C. 1986. Sympatric subspecies in 

Chrysothamnus nauseosus. In: E. D. McArthur 

and B.L Welch, comps. Proceedings—

symposium on the biology of Artemisia and 

Chrysothamnus; 1984 July 9-13, Provo, UT. 

General Technical Report INT-200. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Research Station, Ogden, UT: 98-

103. 

Anderson, L.C. 1995. The ChrysothamnusEricameria connection (Asteraceae). Great 

Basin Naturalist 55: 84-88. 
205

Cook, C.W., P.D. Leonard, and C.D. Bonham. 

1965. Rabbitbrush competition and control on 

Utah rangelands. Bulletin 454. Utah 

Agricultural Experiment Station., Logan, UT. 

 28 p. 

Frischknecht, N.C. 1963. Contrasting effects of big 

sagebrush and rubber rabbitbrush on production 

of crested wheatgrass. Journal of Range 

Management 16: 70-74. 

Goodrich, S., E.D. McArthur, A. Huber, and J.E. 

Ott. 1999. Ungulate browsing of two 

populations of rubber rabbitbrush. In: E.D. 

McArthur, W.K. Ostler, and C.L. Wambolt, 

comps. Proceedings: shrubland ecotones; 1998 

August 12-14; Ephraim, UT. Proc RMRS-P-11. 

Ogden, UT: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research 

Station: 196-199. 

Hall, H.M. and T.H. Goodspeed. 1919. A rubber 

plant survey of Western North America. II. 

Chrysil, a new rubber from Chrysothamnus 

nauseosus. University of California Publications 

in Botany 7: 183-264. 

Hanks, D.L., E.D. McArthur, A.P. Plummer, B.C. 

Giunta, and A.C. Blauer. 1975. 

Chromatographic recognition of some palatable 

and unpalatable subspecies of rubber rabbitbrush 

in and around Utah. Journal of Range 

Management 28: 144-148. 

Hegerhorst, D., D.J. Weber, and E.D. McArthur. 

1987. Resin and rubber content in 

Chrysothamnus. Southwestern Naturalist 32: 

475-482. 

International Plant Names Index. 2002. 

International Plant Names Index. 

xhttp//www.ipni.org. [not paged]. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service 

USDA Forest Service, Intermountain Forest and 

Range Experiment Station, Ogden, UT. 82 p. 

McArthur, E.D., S.E. Meyer, and D.J. Weber. 

1987. Germination rate at low temperature: 

rubber rabbitbrush population differences. 

Journal of Range Management 40: 530-533. 

McArthur, E.D., R. Stevens, and S.B. Monsen 

1995. Adaptation and success of big sagebrush 

and rubber rabbitbrush on disturbed sites. In: 

G.E. Schuman and G.F. Vance, eds. Decades 

later: a time for reassessment, proceedings of the 

12th Annual Meeting of the American Society 

for Surface Mining and Reclamation; 1995 June 

3 -8, Gillette, WY. American Society for 

Surface Mining and Reclamation, Princeton, 

WV. p. 811-823. 

Meyer, S.E. 1997. Ecological correlates of achene 

mass variation in Chrysothamnus nauseosus

(Asteraceae). American Journal of Botany. 84: 

471-477. 

Meyer, S.E., E.D. McArthur, and G.L. Jorgensen. 

1989. Variation in germination response to 

temperature in rubber rabbitbrush 

(Chrysothamnus nauseosus:Asteraceae) and its 

ecological implications. American Journal of 

Botany 76: 981-991. 

Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981. 

Viability of seed from thirty-two shrub and forb 

species through fifteen years of warehouse 

storage. Great Basin Naturalist 41: 274-277. 

U.S. Department of Agriculture, Forest Service. 

1937. Range plant handbook. United States 

Government Printing Office, Washington, D.C. 

816 p. 

Weber, D.J., T. Davis, E.D. McArthur, and N. 

Sankhla. 1985. Chrysothamnus nauseosus

(rubber rabbitbrush): mutiple-use shrub of the 

desert. Desert Plants 7: 172-209. 

Yoakum, J. 1986. Use of Aremisia and 

Chrysothamnus by pronghorns. In: McArthur, E. 

D. and B. L Welch, comps. Proceedings—

symposium on the biology of Artemisia and 

Chrysothamnus; 1984 July 9-13, Provo, UT. 

General Technical Report INT-200. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Research Station, Ogden, UT. 

p. 176-180. 

Young, J.A., R.A. Evans, and B.L. Kay. 1984. 

Persistence and colonizing ability of rabbitbrush 

collections in a common garden. Journal of 

Range Management 37: 373-377. 
206 

Chrysothamnus parryi (Gray) Greene Parry rabbitbrush 

ASTERACEAE 

Synonyms: Bigelovia parryi Gray 

 Ericameria parryi (Gray) G.L. Nesom & G.I. Baird 

 Linosyris parryi Gray 

E. Durant McArthur and Jeffrey R. Taylor 

 

General Description.—Parry rabbitbrush is a 

shrub, intermediate in height, growth habit, and 

stem and leaf tomentum between rubber 

rabbitbrush [Chrysothamnus nauseosus (Pallas ex 

Push) Britton] and low rabbitbrush 

(Chrysothamnus viscidiflorus Nutt.). It is usually 

short, being 20 to 60 cm tall, with numerous 

spreading to erect flexible branches. Branches are 

covered with a felt-like white to green tomentum. 

Leaves are narrowly linear to elliptic. Leaf 

dimensions range from 0.5 to 8 mm wide and 1 to 

8 cm long. The leaf surface is somewhat viscid and 

leaf pubescence can vary from glabrous to 

tomentose. Flower heads are composed of four to 

twenty yellow disc flowers; heads are usually 

arranged in terminal racemes that sometimes form 

panicles. Corollas are tubular to funnelform and 8 

to 11 mm long. Involucral bracts can vary in 

length (9 to 14 mm) and shape (acuminate to very 

attenuate) according to subspecies. Parry 

rabbitbrush is uniformly diploid n = 9, 2x = 18 

(Anderson 1966). 

Taxonomy.—Parry rabbitbrush is a diverse group 

with 12 subspecies (Anderson 1986, McArthur and 

Meyer 1987). Subspecies vary one from another 

by numerous characteristics including stature, 

growth habit, leaf shape, leaf size, involucral bract 

shape, number of flowers per head, pubescence, 

and habitat. Neesom and Baird (1993) recently 

suggested that Parry rabbitbrush would better be 

placed in Ericameria based on molecular genetic 

data. Anderson (1995) was not comfortable with 

this change. Notable subspecies include asper, 

attenuatus, howardii, monocephalus, nevadensis, 

and parryi. 

 Subspecies asper is a low shrub 15 cm or 

more high and inhabits mountainsides that border 

the deserts in western Nevada and eastern 

California (Anderson 1986, McArthur and others 

1979). Leaves are covered with short-stalked resin 

glands and flower heads contain five to ten disc 

flowers. 

Chrysothamnus parryi ssp. attenuatus has 

small leaves up to 4 mm in length. Stems are 

mostly erect and shrubs can grow as high as 60 

cm. Flower heads contain five to seven disc 

flowers and bracts are ranked into five vertical 

rows. This subspecies is found in Utah, 

southwestern Colorado, and northwestern New 

Mexico (Anderson 1986, McArthur and others 

1979). 

Subspecies howardii is similar in size to 

subspecies attenuatus. The uppermost leaves 

usually extend beyond the uppermost heads of the 

inflorescence. Flower heads have five to seven 

pale yellow disc flowers and involucral bracts are 

vertically ranked with spreading tips. This 

subspecies is endemic to Utah, southern Wyoming, 

Colorado, New Mexico, and Nebraska (Anderson 

1986, McArthur and others 1979). 
207

In subspecies monocephalus, leaf shape 

can vary from linear-oblanceolate to spatulate. 

Like subspecies howardii, the upper leaves usually 

extend beyond the inflorescence. One unique 

characteristic is that flower heads occur singly or 

in pairs rather than forming racemes. Flower heads 

contain five or six disc flowers subtended by 

obscurely ranked involucral bracts. This 

subspecies occurs in high mountains between 790 

and 3,400 m in eastern California and western 

Nevada. 

Subspecies nevadensis is comprised of 

small shrubs up to 60 cm in height. Leaves are 

linear to linear-oblanceolate and covered with 

resin or gray tomentum. The uppermost leaves 

rarely extend beyond the inflorescence. Flower 

heads contain four to six yellow disk flowers and 

are subtended by ranked involucral bracts with 

slender recurved tips. Subspecies nevadensis

occurs on dry mountainsides from eastern 

California to eastern Nevada, southwestern Utah, 

and Northern Arizona, being most common on the 

eastern slopes of the Sierra Nevada (Anderson 

1986, McArthur and others 1979). 

Subspecies C. parryi ssp. parryi consists 

of low shrubs with erect branches and can be as 

tall as 76 cm. The uppermost leaves usually extend 

beyond the inflorescence. Flower heads include 10 

to 20 disc flowers and are subtended by obscurely 

ranked bracts with attenuate tips. This subspecies 

grows on dry hillsides and plains in central 

Nevada, southern Utah, south-central Wyoming, 

western Colorado, and northern New Mexico 

(Anderson 1986, McArthur and others 1979). 

Anderson (1970) made a case for this taxon being 

the most primitive of all Chrysothamnus. 

Parry rabbitbrush in the form of C. parryi 

ssp. attenuatus has been implicated in the 

parentage of the putative stabilized hybrid species 

C. nauseosus ssp. uintahensis Anderson 1984). 

Range.—The range of Parry rabbitbrush extends 

from California in the west, to western Nebraska 

in the east, and from Wyoming in the north, to 

New Mexico and Arizona in the south. Subspecies, 

however, may be more geographically restricted. 

Within its range, Parry rabbitbrush may be found 

at elevations between 790 and 3,400 m. 

Ecology.—Parry rabbitbrush grows on dry, open 

foothills and mountains. Precipitation throughout 

its range generally exceeds 380 mm. Parry 

rabbitbrush tends to increase on overgrazed or 

otherwise disturbed areas. Populations are usually 

smaller and more scattered than those of more 

common species, such as rubber and low 

rabbitbrushes. 

Reproduction.—Parry rabbitbrush flowers from 

July to September and seed matures in October 

and November. Achenes are 5 to 6 mm long and 

covered with long, shaggy, oppressed hairs. We 

presume there are about 1,600 cleaned achenes per 

gram and that they have good fertility as is the 

case for other Chrysothamnus species 

(Deitschmann and others 1974). Achenes are wind 

disseminated in late fall or winter. In addition to 

sexual reproduction, Parry rabbitbrush has been 

observed to spread through underground roots 

(Paulsen and Miller 1968, McArthur and others 

1979). We are unaware of seed germination data. 

Growth and Management.—Seed germinates in 

the early spring and seedlings become established 

during the following spring and summer. Plants 

generally mature within 5 years and typically live 

15 to 20 years. Paulsen and Miller (1968) reported 

that, when necessary, Parry rabbitbrush can be 

controlled by the herbicide Tordon®.1

Benefits.—Although Parry rabbitbrush is a 

component of western uplands, it is spotty in 

distribution and usually sparse in population 

density. It has some importance as a browse source 

in those areas where it is locally abundant. Some 

subspecies of Parry rabbibrush, including C. parryi

ssp. asper and C. parryi ssp. monocephalus, are 

particularly attractive plants and have horticultural 

potential as ornamentals. 

 

References 

Anderson, L.C. 1966. Cytotaxonomic studies in 

Chrysothamnus (Asterae, Compositae). American 

Journal of Botany 53: 204-212. 

Anderson, L.C. 1970. The karyotype of 

Chrysothamnus parryi ssp. parryi and its 

implication. Transactions of the Kansas 

Academy of Science 72: 399-401. 

Anderson, L.C. 1984. Chrysothamnus nauseosus

ssp. uintahensis: a stabilized hybrid. Great Basin 

Naturalist 44:416-420. 

Anderson, L.C. 1986. An overview of the genus 

Chrysothamnus. In: E.D. McArthur and B.L 

Welch, comps. Proceedings—symposium on the 

 

1

 Use of trade names does not imply endorsement 

of the U. S. Department of Agriculture 
208 

biology of Artemisia and Chrysothamnus; 1984 

July 9-13, Provo, UT. General Technical Report 

INT-200. U.S. Department of Agriculture, Forest 

Service, Intermountain Research Station, Ogden, 

UT: 29-45. 

Anderson, L.C. 1995. The ChrysothamnusEricameria connection (Asteraceae). Great Basin 

Naturalist 55: 84-88. 

Deitschman, G.H., K.R. Jorgensen, and A.P. 

Plummer. 1974. Chrysothamnus. In: C.S. 

Schopmeyer, tech. coord., Seeds of Woody 

Plants of the United States. Agriculture 

Handbook 450. Washington, DC, U.S. 

Department of Agriculture, Forest Service. 

p. 326-328. 

McArthur, E.D., A.C. Blauer, A.P. Plummer, and 

R. Stevens. 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U.S. Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 82 p. 

McArthur, E.D. and S.E. Meyer. 1987. A review 

of the taxonomy and distribution of 

Chrysothamnus. In: K.L. Johnson, ed. 

Proceedings of the Fourth Utah Ecology 

Workshop; 1986 September 17-18, Cedar City, 

UT. College of Natural Resources, Utah State 

University, Logan, UT. p. 9-18. 

Nesom, G.L. and G.I. Baird. 1993. Completion of 

Ericameria (Asteraceae: Asterae), diminution of 

Chrysothamnus. Phytologia 75: 74-93. 

Paulsen, H.A., Jr. and J.C. Miller. 1968. Control of 

Parry rabbitbrush on mountain grasslands of 

Western Colorado. Journal of Range 

Management 21: 165-177 

. 
209

Chrysothamnus viscidiflorus Nutt. low rabbitbrush 

ASTERACEAE 

Synonyms: Bigelovia douglasi Gray 

 Chrysothamnus douglasi Clements & Clements 

 Crinitaria viscidiflora Hook. 

 Ericameria viscidiflora (Hook) L. C. Anderson 

E. Durant McArthur and Jeffrey R. Taylor 

General Description.—Low rabbitbrush, also 

known as yellowbrush, green rabbitbrush, little 

rabbitbrush, or Douglas rabbitbrush, is an erect 

shrub, with many branches stemming from a 

simple base. It is typically 0.3 to 1.1 m tall. 

Brittle, erect branches can be glaborous or 

puberulent with pale green or white bark. Leaves 

are often twisted. Leaf shape can vary between 

narrowly linear to oblong or lanceolate, with leaf 

length ranging from 1 to 6 cm. Leaves may be 

glaborous or pubescent, the degree of which may 

vary by subspecies. Flowerheads include five 

perfect, fertile disc flowers, which are arranged in 

compact terminal cymes. Each head includes 

approximately 15 involucral bracts that are 

arranged in poorly to well-defined vertical ranks. 

The bracts of some subspecies have a greenish to 

brownish thickened spot near their apex 

(McArthur and others 1979). Low rabbitbrush is 

the only species within the genus with frequent 

polyploidy; diploids, tetraploids, and hexaploids 

have been documented (x = 9, 2n = 18, 36, and 54, 

respectively) (Anderson 1966). In addition, 

occasional triploids and pentaploids are known 

(Anderson 1971). 

Taxonomy.—This species includes five 

subspecies and several ecotypes within subspecies 

(Anderson 1980, McArthur and Meyer 1987). The 

most common subspecies include two glabrous 

subspecies, viscidiflorus and axillaris, and two 

pubescent subspecies lanceolatus and puberlus. 

Abrams and Ferris (1960) describe low rabbitbrush 

as a highly polymorphic species with several 

freely integrating subspecies with overlapping 

distributions. Stickyleaf low rabbitbrush or green 

rabbitbrush (ssp. viscidiflorus) is the largest 

subspecies of low rabbitbrush. Mature shrubs are 

generally more than 50 cm tall whereas mature 

shrubs of other subspecies are generally less than 

50 cm tall. In addition, the leaves, stems, and 

inflorescences of ssp. viscidiflorus are viscid 

(sticky). Both mountain low rabbitbrush or 

yellowbrush (ssp. lanceolatus) and hairy low 

rabbitbrush (ssp. puberlus) are shorter at maturity 

than ssp. viscidiflorus, and both have finely 

pubescent stems. Mountain low rabbitbrush has 

gray or straw-colored stems, whereas hairy low 

rabbitbrush has yellow to green stems. All four of 

these subspecies are fairly widespread throughout 

the Great Basin. Some previously recognized 

subspecies, such as ssp. elegans and pumulis, have 

been consolidated; chromatographic work 

substantiates the consolidations (Anderson 1980; 

McArthur and others 1978). There is a difference 

of opinion on the taxonomic affinity of C. 
210 

viscidiflorus with C. nauseosus and C. parryi 

based on molecular genetic techniques (Nesom 

and Baird 1993; Anderson 1995) 

Range.—Low rabbitbrush is one of the most 

widely distributed shrubs in Western North 

America (McArthur and others 1979). One reason 

for its broad distribution is the species’ great 

ecological amplitude. It is found growing on dry, 

open areas from British Columbia and Montana, 

south to New Mexico, Arizona, and eastern 

California. 

Ecology.—Due to its adaptability, low rabbitbrush 

grows in a variety of habitats. Plants generally, 

however, inhabit drier sites, such as foothills, 

mountains, dry plains, and valleys. Annual 

precipitation within its range varies between 250 

and 535 mm, with notable exceptions at higher 

elevations. Anderson (1986) has shown that 

polyploid races, in subspecies where polypoidy 

occurs (ssp. lanceolatus, puberulus, and 

viscidiflorus), are adapted to lower and drier sites 

than their diploid counterparts. Interestingly, 

Anderson has also observed a correlation within 

certain subspecies between plant stature and leaf 

size with altitude and precipitation (personal 

communication). In addition to its ability to 

tolerate xeric conditions, some variants of low 

rabbitbrush can grow well in areas with poor or 

disturbed soils. Low rabbitbrush can be found at 

elevations ranging from 790 to 3,400 m. It is 

commonly associated with big sagebrush 

(Artemisia tridentata), broom snakeweed 

(Gutierrezia sarothrae), and other rabbitbrushes 

Chrysothamnus sp.), although it can be associated 

with shadscale (Atriplex confertifolia), winterfat 

(Ceratoides lanata), fourwing saltbush (Atriplex 

canescens), greasewood (Sarcobatus 

vermiculatus), and occasionally pinyon (Pinus sp.) 

and juniper (Juniperus sp.). Goodrich and others 

(2001) note that C. viscidiflorus does not 

commonly overtop associated herbaceous 

vegetation. Elevation tends to partition the 

subspecies; mountain low rabbitbrush (ssp. 

lanceolatus) especially and stickyleaf low 

rabbitbrush (ssp. viscidiflorus) grow well at higher 

elevations, while hairy low rabbitbrush (ssp. 

puberlus) does best on lower desert foothills. 

Reproduction.—Shrubs flower from August 

through October; seed matures from October until 

December. Like other members of the genus, low 

rabbitbrush seeds (or achenes) are wind 

disseminated. Meyer (1996) found that 

germination rates at 15 o

C for ssp. lanceolatus and 

viscidiflorus were 37 and 58 percent, respectively. 

Seeds from plants adapted to higher elevations 

(where seeds ripen earlier in the year) are more 

dormant than those collected from lower 

elevations and may be more chill responsive 

(Meyer 1996). There are about 1,720 cleaned seeds 

per gram (Deitschman and others 1974). 

Growth and Management.—Seedlings emerge in 

spring and grow quickly. After a period of growth 

and establishment, shrubs often produce seed after 

their second growing season (Meyer 1996). Low 

rabbitbrush can dominate rangelands that are 

grazed heavily or cleared. This species has a strong 

tendency to sprout from the base (Wasser 1982), 

which may make it more difficult to remove and 

difficult to control with herbicides (Cook and 

others 1965). Fall is the preferred time to seed; 

seed can be drilled or broadcast and should remain 

within the top 3 mm of the soil surface. Seeds can 

be harvested by hand and by vacuum harvesting 

techniques. 

Benefits.—Low rabbitbrush is an important native 

component of Western North American 

shrublands. It can provide important browse to 

both game and livestock, especially during late fall 

and winter months after more desirable forage has 

been consumed. As a forage shrub, the palatability 

of low rabbitbrush varies significantly among 

subspecies, varieties, locality, and season. Some 

subspecies, e.g., ssp. lanceolatus, may be used 

heavily by domestic livestock and wildlife, 

whereas others are consumed little, if at all (USDA 

1937, Goodrich and others 2001). In addition to its 

value as a browse species, low rabbitbrush has 

successfully been used to revegetate depleted 

rangelands and other disturbed sites (Plummer 

1977). 

References 

Abrams, R. and R.S. Ferris. 1960. Illustrated flora 

of the Pacific States, Vol. 4, Bignoniaceae to 

Compositae. Stanford University Press, 

Stanford, CA. 732 p. 

Anderson, L.C. 1966. Cytotaxonomic studies in 

Chrysothamnus (Asterae, Compositae). 

American Journal of Botany 53: 204-212. 

Anderson, L.C. 1971. Additional chromosome 

numbers in Chrysothamnus (Asteraceae). 

Bulletin of the Torrey Botanical Club 98: 222-

225. 
211

Anderson, L.C. 1980. Identity of narrow-leaved 

Chrysothamnus viscidiflorus (Asteraceae). 

Great Basin Naturalist 40: 117-120. 

Anderson, L.C. 1986. Cytogeography of 

Chrysothamnus viscidiflorus. In: E.D. McArthur 

and B.L. Welch, comps. Proceedings—

symposium on the biology of Artemisia and 

Chrysothamnus; 1984 July 9-13, Provo, UT. 

General Technical Report INT-200. Ogden, UT: 

U.S. Department of Agriculture, Forest Service, 

Intermountain Research Station. p. 93-97. 

Anderson, L.C. 1995. The ChrysothamnusEricameria connection (Asteraceae). Great 

Basin Naturalist 55: 84-88. 

Cook, C.W., P.D. Leonard, and C.D. Bonham. 

1965. Rabbitbrush competition and control on 

Utah rangelands. Bulletin 454. Utah 

Agricultural Experiment Station., Logan, UT. 

28 p. 

Deitschman, G.H., K.R. Jorgensen, and A.P. 

Plummer. 1974. Chrysothamnus. In: C.S. 

Schopmeyer, tech. coord., Seeds of Woody 

Plants of the United States. Agriculture 

Handbook 450. U.S. Department of Agriculture, 

Forest Service. p. 326-328. 

Goodrich, S., R.M. Thompson, and A. Huber. 

2001. A yellowbrush/grass community type 

from the Uinta Mountains and Utah plateaus. 

In: E.D. McArthur and D.J. Fairbanks, comps. 

Shrubland ecosystem genetics and biodiversity: 

proceedings; 2000 June 13-15, Provo, UT. 

Proceedings RMRS-P-21. U. S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Research Station, Ogden, UT. p. 344-346. 

McArthur, E.D., A.C. Blauer, A. P. Plummer, and 

R. Stevens. 1979. Characteristics and 

hybridization of important intermountain shrubs. 

III. Sunflower family. Research Paper INT-220. 

U. S. Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. 82 p. 

McArthur, E.D., D.L. Hanks, A.P. Plummer, and 

A.C. Blauer. 1978. Contributions to the 

taxonomy of Chrysothamnus visicidiflorus

(Astereae, Compositae) and other 

Chrysothamnus species using paper 

chromatography. Journal of Range Management 

31: 216-223. 

McArthur, E.D. and S.E. Meyer. 1987. A review 

of the taxonomy and distribution of 

Chrysothamnus. In: K.L. Johnson, ed. 

Proceedings of the Fourth Utah Ecology 

Workshop; 1986 September 17-18, Cedar City, 

UT. College of Natural Resources, Utah State 

University, Logan, UT. p. 9-18. 

Meyer, S.E. 1996. Chrysothamnus Nutt. In:

Woody Plant Seed Manual, 

http://www.wpsm.net/Chrysothamnus.pdf. 11 p. 

 Nesom, G.L., and G.I. Baird. 1993. Completion of 

Ericameria (Asteraceae: Asterae), diminution of 

Chrysothamnus. Phytologia 75: 74-93. 

Plummer, A.P. 1977. Revegetation of disturbed 

intermountain area sites. In: J. L. Thames, 

editor, Reclamation and use of disturbed land in 

the Southwest. University of Arizona Press, 

Tucson, AZ: 302-339. 

U.S. Department of Agriculture, Forest Service. 

1937. Range plant handbook. United States 

Government Printing Office, Washington, DC. 

816 p. 

Wasser, C.H. 1982. Ecology and culture of 

selected species useful in revegetating disturbed 

lands in the West. U.S. Department of the 

Interior, Fish and Wildlife Service, Washington, 

DC. 347 p. 
212 

Cissampelos pareira L. velvet leaf 

MENISPERMACEAE 

Synonyms: Cissampelos caapeba L. 

 Cissampelos microcarpa DC. 

John K. Francis 

General Description.—Velvet leaf is also known 

as abuta, pareira, alcotá, bejuco de mona, oreja de 

raton, hierba de peso, curarina, liane amère, lianecordé, and liane molle. It is a shrubby climber (or 

liana) that reaches 3 to 6 m along the ground or 

into the crowns of trees. The stem is woody, 

flexible, and slender (to 1 cm), and twines for 

support. The pith and rays of the stem cross section 

resemble a wagon wheel with spokes; annual rings 

are not visible. The root system consists of flexible, 

light-brown lateral roots with sinkers and 

moderately abundant fine roots. There are 

relatively few branches. Alternate leaves are 

usually softly pubescent on both surfaces. The 

petioles are 3 to 7 cm long. Venation is palmate in 

widely oval or nearly round 4- to 10-cm blades. 

The small staminate and pistillate inflorescences 

(cymes) are borne in leaf axils. The fruits are juicy 

red, red-orange, or yellow, hairy drupes 4 to 5 mm 

in diameter. Each fruit is partially covered by a 

rounded bract. The seeds have a croissant shape 

(Acevedo-Rodríguez 1985, Howard 1988, Liogier 

1985, Stevens and others 2001). The species’ 

chromosome number is 2n = 24 (Long and Lakela 

1976). 

Range.—Velvet leaf is native from Mexico to 

Argentina and Peru on the New World mainland 

and in the West Indies (Instituto Botánico 

Darwinian 2002, Secretaría de Medio Ambiente y 

Recursos Naturales 2002, Stevens and others 

2001). It is native to Florida, although rare or 

possibly locally extinct (Nelson 1996). It is also 

found throughout tropical Asia and Africa (Long 

and Lakela 1976, Parrotta 2001), although it is not 

clear whether it is native or naturalized there. 

Ecology.—Velvet leaf is a “good site” species. It 

usually does not grow on exposed clay subsoils, 

compacted soils, excessively drained, or very 

poorly drained soils. The species occurs on a wide 

variety of soil textures, pH levels, and on soils 

derived from most parent materials including 

limestone and ultramafic rocks (serpentine). In 

Puerto Rico, velvet leaf grows in areas that receive 

from 750 to about 2400 mm of annual precipitation 

at elevations from near sea level to about 1,500 m 

(Stevens and others 2001). Velvet leaf may be 

found on roadsides, fencerows, river banks, 

hammocks, brushy pastures, and secondary and 

remnant forests. It is moderately intolerant of shade 

and does not grow under the closed canopy of high 

forest. Openings, and therefore disturbance, appear 

necessary for the maintenance of the species. 

Reproduction.—Velvet leaf flowers and fruits all 

year in the Americas (Acevedo-Rodríguez 1985, 

Stevens and others 2001). In India it flowers 

between July and October and fruits from October 

to December (Parrotta 2001). Fruits collected in 

Puerto Rico weighed an average of 0.1925 +

0.0008 g/fruit. Seeds from them averaged 0.0109 

g/fruit (air-dried). When planted in commercial 

potting mix without pretreatment, 26 percent of the 

seeds germinated between 28 and 61 days after 

sowing (author’s observation). Birds presumably 

disperse the seeds. Fruit and seed production are 

generally moderate, and seedlings are rarely 

abundant. Stems layer (root) wherever they contact 

the soil. 

Growth and Management.—No information is 
213

available on the growth rate of seedlings. Growth 

of sprouts is rapid (at least 3 m in the first year). 

Individual stems are not long-lived, but by 

sprouting and layering, plants or clones may last 

many years. The creation of small forest openings 

and protection against heavy grazing are probably 

good strategies to promote natural establishment of 

new plants. 

Benefits.—Velvet leaf adds to biodiversity and 

biomass, helps stabilize the soil, and furnishes food 

and cover for wildlife. A major interest in the 

species arises from the natural medicinal benefits 

of the plant’s chemical contents. Known as the 

“midwife’s herb,” it has been used for centuries by 

native peoples of South America to treat menstrual 

cramps, prevent threatened miscarriage, control 

uterine hemorrhages, and ease childbirth and 

postpartum pain (Rain-tree 2002). The list of other 

natural medicinal applications to which the herb is 

applied is large: urinary infections, kidney stones, 

arthritis, snakebite, cough, dysentery, piles, ulcers, 

pain, indigestion, colic, skin irritations, stings, 

intestinal worms, and wounds (Parrotta 2001, Raintree 2002). Ground tissues and preparations of 

velvet leaf are sold throughout the world in 

markets, shops, and mail-order companies. The 

physiological effects are apparently derived from a 

number of alkaloids found in the tissues of the 

plant (International BioPark Foundation 2002, 

Morita and others 1993a, Morita and others 

1993b). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Instituto de Botánica Darwinian. 2002. Catálogo 

de las plantas vasculares de la Argentina. 

http://www.darwin.edu.ar/Catalogo/indicevascul

ares.htm. [not paged]. 

International BioPark Foundation. 2002. Abuta, 

family Menispermaceae. http://www.biopark. 

org/peru/abuta.html. 3 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

tropical Florida. Banyan Books. Miami, FL. 

962 p. 

Morita, H., K. Matsumoto, K. Takeya, and H. 

Itokawa. 1993a. Azafluoranthene alkaloids from 

Cissampelos pareira. Chemical and 

Pharmaceutical Bulletin 41(7): 1,307-1,308. 

Morita, H., K. Matsumoto, K. Takeya, H. Itokawa, 

and Y. Iitaka. 1993b. Structures and solid state 

tautomeric forms of two novel antileukemic 

tropoloisoquinoline alkaloids, pareirubrines A 

and B, from Cissampelos pareira. Chemical and 

Pharmaceutical Bulletin 41(8): 1,418-1,422. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CABI Publishing, Wallingford, UK and 

New York. 917 p. 

Rain-tree. 2002. Abuta. http://www.raintree.com/abuta.htm. 7 p. 

Secretaría de Medio Ambiente y Recursos 

Naturales. 2002. Especies con usos no 

maderables in bosques tropicales y 

subtropicales: Cissampelos pareira L. 

http://www.semarnat.gob.mx.pfnm2/fichas/cissa

mpelos_pareira.htm. 2 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel (eds.). 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden, St. Louis, MO. 

p. 945-1,910. 
214 

Citrus aurantiifolia (Christm.) Swingle Key lime 

RUTACEAE 

Synonyms: Limonia aurantifolia Christm. 

 Limon spinosum Mill. 

 Citrus limonia Osbeck 

 Citrus lima Luman 

 Citrus spinosissima G.F.W. Meyer 

 Citrus acida Roxb. 

John K. Francis 

General Description.—Key lime is the name used 

most often to refer to a primitive race of Citrus 

arantiifolia cultivated and naturalized in the West 

Indies. It is also referred to as Mexican lime, West 

Indian lime, lima, limón criollo, limón agria, limón 

boba, and citron (Little and Wadsworth 1964). 

Key lime is an evergreen, spiny shrub or small tree 

to 6 m in height. The plant has single or multiple 

stems and irregular branches covered with 

smoothish brown to gray bark. The twigs are 

quadrangular (when young), green, and bare sharp 

axillary spines 3 to 17 mm long. The leaves are 

yellow-green to dark green, with 5- to 28-mm 

winged petioles and elliptic to oval leathery 4- to 

13-cm long blades with edges that have minute 

rounded teeth. The crushed foliage has a strong, 

distinct, spicy (citrus) odor and taste. The four- to 

five-petaled white flowers occur in few-flowered 

axillary clusters. The fruits (hesperidiums) are 

ellipsoidal, 3 to 5 cm in diameter, have juicy, 

greenish-yellow flesh, and are yellow at maturity. 

They contain a few white, pointed seeds about 1 

cm long (Liogier 1988, Little and Wadsworth 

1964). 

Range.—Key lime is believed to be native of 

eastern Malaysia. It was introduced to the Asian 

mainland early in historical times and carried by 

Arab traders to the Middle East and eventually 

came to Europe during the Crusades (Burkill 

1997). The species was introduced to the West 

Indies by Columbus during his second voyage 

(Ehler 2002). Key lime has been planted 

throughout the tropics and has naturalized in at 

least Puerto Rico, the Virgin Islands (Little and 

Wadsworth 1964), and the Florida Keys (Nelson 

1996). 

Ecology.—Key lime is most competitive in areas 

that receive from about 700 to 1000 mm of mean 

annual precipitation. It tolerates drought better than 

any of the other citrus fruit species (Morton 1987). 

If planted, it will grow but becomes increasingly 

susceptible to disease in areas that receive up to 

about 2000 mm of annual precipitation. Most welldrained soils are suitable, particularly those rich in 

calcium (Morton 1987). Elevations below 900 m 

are best (Secritaría del Medio Ambiente y Recursos 

Naturales 2002). Key lime is intolerant of shade 

and will not survive long under a closed forest 

canopy. Cattle grazing encourages it somewhat by 

eliminating some of the competition without 

damaging it. Plants are top-killed by fires and hard 

frosts, but will sprout and survive if the disturbance 

is not repeated frequently. Naturally reproduced 

trees may be seen on roadsides, fencerows, 

abandoned farms, secondary forests, and coastal 

hammocks. 
215

Reproduction.—Key lime flowers in late spring 

and fruits in fall to spring in Florida (Nelson 1996). 

In Puerto Rico, it flowers in spring to early summer 

and fruits in summer and fall (Little and 

Wadsworth 1964). The fruits ripen and fall from 

the trees 5 to 6 months after flowering (Morton 

1987). Fruits in one collection in Puerto Rico 

ranged from 3.0 to 4.5 cm in diameter and weighed 

from 19 to 51 g. Air-dried seeds separated from 

them averaged 0.0702 + 0.0037 g/seed or 14,000 

seeds/kg. Placed on moist blotter paper, 84 percent 

germinated between 15 and 49 day after sowing. 

Germination is epygeal. Many of the seeds are 

polyembryonic. Seedlings begin to fruit in 3 to 6 

years (Morton 1987). In the West Indies, the seeds 

are mainly dispersed by gravity and humans. There 

are undoubtedly animal vectors in the original 

native habitat. Key lime can be air-layered using 

indol buteric acid (IBA) with a high degree of 

success (Morton 1987). 

Growth and Management.—Growth of Key lime 

is slow, less than 0.5 m/year. Plants may live for 25 

years or more. Although the species is not managed 

in the wild, existing trees are usually protected by 

local inhabitants, whenever possible. Plantations 

are established using potted material that is set at 

7.5- by 7.5-m spacing. Pruning is usually not 

necessary (Morton 1987). 

Benefits.—Key lime has been under cultivation or 

semicultivation for thousands of years. Improved 

horticultural varieties, which have bigger, generally 

seedless fruits and thornless plants, now dominate 

the local and international fruit markets. The 

unimproved type is still managed commercially 

and is the basis for a juice and lime oil industry 

(Morton 1987). It is also planted for a dooryard 

fruit tree, and fruits are harvested from the wild. 

The principal use is still for food, refreshing drinks, 

tasty disserts, and for seasoning meats, vegetables, 

salads, sauces, and casseroles (Ehler 2002, Katzer 

2002). The wild type is superior in flavor to the 

improved varieties because it has a stronger flavor 

and a higher acid content (Ehler 2002). The 

pericarp (rind) contains 7 percent essential oil with 

principal constituants, citral, limonene, and 

fenchon as well as terpineol, bisabolene, and other 

terpenoids (Katzer 2002). Key lime is used to treat 

a huge number of ailments (Burkill 1997, Liogier 

1990). The author recommends a tea prepared from 

juice, fruit rind, or leaves as an expectorant and to 

relieve catarrh brought on by colds and flu. The 

fresh fruits and bottled juice are an excellent source 

of vitamin C and were once relied upon to prevent 

scurvy. Essential oils of Key lime and some other 

citrus fruits cause phytophotodermatosis in 

sensitive individuals (Bruneton 1999). The wood is 

hard and heavy, but is used for little other than 

fuel. It is a good honey plant. With pruning, the 

plant can be used for a living fence post (Little and 

Wadsworth 1964) and can be formed into a hedge 

(Burkill 1997). 

References 

Bruneton, J. 1999. Toxic plants dangerous to 

humans and animals. Lavoisier Publishing, 

Paris. 545 p. 

Burkill, H.M. 1997. The useful plants of West 

Tropical Africa. Vol. 4. Royal Botanic Gardens, 

Kew, UK. 969 p. 

Ehler, J.T. 2002. Key lime (Citrus aurantifolia). 

Food Reference, Key West, FL. 

http://foodreference.com/html/artkeylimes.html. 

2 p. 

Katzer, G. 2002. Lime [Citrus aurantifolia

(Christm. & Panz.) Swengle]. Gernet Katzer’s 

spice pages. http://www-ang.kfunigraz. 

ac.at/~katzer/engl/Citr_aur.html. 4 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR 566 p. 

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

Morton, J.F. 1987. Citrus aurantifolia Swingle. 

from: Fruits of warm climates. 

http://www.hort.perdue.edu/newcrop/morton/me

xican_lime.html. 8 p. 

Nelson, Gil. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc. Sarasota, FL. 

 391 p. 
216 

Secritaría del Medio Ambiente y Recursos 

Naturales. 2002. Especies con usos no 

maderables en bosques tropicales y 

subtropicales: Lantana involucrata L. 

http://www.semarnat.gob.mx.pfnm2.fichas/ 

citrus_aurantifolia.htm. 2 p. 
217

Clematis orientalis L. oriental virginsbower 

RANUNCULACEAE 

Synonyms: Clematis aurea A. Nels. & J.F. Macbr. 

 Viticella orientalis (L.) W.A. Weber 

John K. Francis 

General Description.—Oriental virginsbower, 

also known as orange peel or orange peel clematis, 

is a deciduous, sometimes suffruticose vine or 

scrambling shrub. The multiple stems may reach 8 

m in length and as much as 7 cm in thickness. 

Stems are covered with longitudinally fissured 

gray-brown bark. Inner bark is green. Vines cling 

to rocks and other plants by twining as they climb. 

The roots are weak and flexible. Fine roots are tan 

or orange-tan. The leaves are pinnately compound 

with three to seven coarsely-toothed to entire, 

ovate to linear-lanceolate leaflets 1 to 5 cm long. 

The foliage has a gray-green color. Axillary 

cymes, which may have one to many flowers, are 

borne on the current season’s growth. Four yellow, 

yellow-orange, or yellowish-green sepals give the 

flowers their color. Flowers have a delicate 

fragrance. The feather-like style, 3 to 10 cm long, 

remains attached to the achene and functions as a 

wing. There are 2n = 16, 32 chromosomes (Plants 

for a Future 2003, Welsh and others 1987, Wu and 

others 2001). 

Range.—Oriental virginsbower is native from 

Turkey through Asia to the Korean Peninsula and 

south to Iran and Northwest India (Griffiths 1994). 

The species has been widely planted as an 

ornamental and has naturalized and escaped in 

Idaho, Nevada, Utah, Colorado, and New Mexico 

(Natural Resources Conservation Service 2003) 

and probably other places in the world. There are 

two varieties, orientalis and robusta, separated on 

the basis of pedicel length and thickness (Wu and 

others 2001). 

Ecology.—Oriental virginsbower grows in 

shrublands, riverbanks, gullies, sand depressions, 

and riparian forests in hot, dry valleys and desert 

and semidesert areas. In Utah, oriental 

virginsbower is mostly found along rivers, creeks, 

and intermittent streams at the mouth of canyons. 

It is not present in cooler sites farther up the 

canyons. It requires well-drained soils, but is not 

particular about soil texture, and tolerates acid and 

alkaline soils from a wide range of parent 

materials. It does well on chalky soils (Plants for a 

Future 2003). The species inhabits sites with 

elevations between 400 and 3,800 m (Wu and 

others 2001). It is hardy to about –15 °C (Plants 

For a Future 2003) and presumably survives from 

roots after top damage. Oriental virginsbower will 

climb taller vegetation, fences, and rocks, but 

forms mounds and mats when objects to climb are 

not available. 

Reproduction.—Oriental virginsbower flowers 

between August and October. Both flowers and 

fruits are often present on the plant at the same 

time. The flowers are pollinated by bees and flies 

(Plants for a Future 2003). A group of air-dried 

seeds collected in Utah averaged 911 seeds/g. 

Oriental virginsbower seeds are dispersed by the 

wind. Seedlings are not abundant. It reproduces 

vegetatively by sprouting from the root crown and 

by layering. 

Growth and Management.—Growth of oriental 

virginsbower is rapid, at least 1 m/year from 

sprouts or existing stems. Although the risk of 
218 

environmental damage is not severe, because it is 

an alien, the species should not be used in 

environmental restoration plantings. No specific 

recommendations for control are available, but the 

general procedure of cutting plants and spot 

spraying the sprouts with broadleaf herbicide 

would probably be effective. The use of potted 

seedlings or rooted cuttings for ornamentals is 

recommended. They should be outplanted in the 

spring after the frost danger has passed. Pruning 

should take place in the spring (Plants for a Future 

2003). 

Benefits.—Oriental virginsbower is planted as an 

ornamental and ground cover. Plants escaped into 

the wild help protect the soil, furnish cover for 

wildlife, and add beauty to the landscape, 

especially during the fall when it produces flower 

and seed heads. Oriental virginsbower is used in 

landscaping as a flowering ground cover. Infusions 

are used as a gargle for sore and ulcerated throats 

and to treat dog bites (Plants for a future 2003). 

References

Griffiths, M. 1994. Index of garden plants. Timber 

Press, Portland, OR. 1,234 p. 

Natural Resources Conservation Service. 2003. 

Plants profiles: Clematis orientalis L., oriental 

virginsbower. http://plants.gov/cgi_bin/ 

plant_profile.cgi?symbol=CLOR. 5 p 

Plants For a Future. 2003. Database search results: 

Clematis orientalis. http://www.scs.leeds.ac.uk/ 

cgi-bin/pfaf/arr_html?Clematis+orientalis. 7 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Hinnins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 

Wu Z., P.H. Raven, and Hong D. (eds.). 2001. 

Flora of China. Vol. 6. Science Press, Beijing 

and Missouri Botanical Garden Press, St. Louis, 

Mo. 512 p. 
219

Clibadium erosum (Sw.) DC. carruzo 

ASTERACEAE 

Synonyms: Trixis erosa Sw. 

John K. Francis 

General Description.—Carruzo, also known as 

jackass breadnut, cachimbo, tuchima, turma de 

toro, bois énivrant, and medsinnyé benní, is a 

shrub or small tree up to 6 m in height and 10 cm 

in basal stem diameter. These plants have a single 

stem from the ground, unless damaged, and are 

much branched in the crown. The trunk bark is 

gray to greenish brown, smooth to slightly 

fissured. The white wood is soft to medium in 

hardness, brittle, without annual rings, and has a 3-

mm pith. The plant often breaks at forked stems. 

Carruzo plants are supported by a root system of 

tan, flexible laterals and may or may not retain a 

taproot. The twigs are ringed at the nodes by 

opposite, three-lobbed leaf scars and densely 

pubescent. The leaves are broadly ovate, with 

long, 2- to 8.5-cm petioles, coarsely and finely 

incised-dentate, and have an elongated tip. The 

inflorescences are many-branched clusters 

(corymbs) containing many heads with white to 

rose-purple flowers. Several black, 2- by 1.5-mm, 

seed-like fruits (akenes) are produced per flower 

(Howard 1989, Liogier 1997, Little and 

Wadsworth 1964). 

Range.—Carruzo is native to Puerto Rico, St. 

Martin, Saba, St. Kitts, Nevis, Montserrat, 

Guadeloupe, Dominica, Martinique, St. Lucia, St. 

Vincent, Costa Rica, and Panama (Howard 1989, 

Missouri Botanical Garden 2003). The species is 

not known to have been planted or naturalized 

elsewhere. 

Ecology.—Carruzo is a species of moist and wet 

mountains. It grows at elevations above about 400 

m in Puerto Rico and it is reported from 800 and 

1,300 to 1,500 m in Costa Rica (Instituto Nacional 

de Biodiversidad 1997, Missouri Botanic Garden 

2003). Mean annual rainfall of its habitat in Puerto 

Rico ranges from 2000 to over 4000 mm/year. The 

soils are mostly clayey, derived from a variety of 

rock types, anaerobic in their subsoils, and have 

pH’s between 5 and 6. Because the species is lightdemanding, it grows mainly in disturbed areas 

(Little and Wadsworth 1964) such as roadsides, 

cuts and fills, landslides, tree-tip mounds, logged 

areas, plantations, fields, and abandoned 

construction sites. Once established, carruzo 

competes aggressively with grass, herbs, and 

shrubs of equal or lesser height. Overtopped, it 

soon dies. 

Reproduction.—Carruzo flowers and fruits nearly 

throughout the year, at least in Puerto Rico (Little 

and Wadsworth 1964). Fruits are produced in great 

numbers. A group of akenes harvested in Puerto 

Rico averaged 0.001867 g/fruit or 536,000 

fruits/kg. The author could not germinate the hard 

akenes on moist blotter paper. Some sort of 

pretreatment is probably necessary. Fruits can be 

collected in quantity by hand. Means of dispersal 

is unknown. Seedlings are common on disturbed 

ground near seed sources. Carruzo shrubs will 

sprout if disturbed and will layer if intact stems 

come in contact with the ground. 

Growth and Management.—Carruzo grows 

rapidly, adding 1 to 1.5 m/year to its height. It 

begins flowering and fruiting after about 1 year 

and probably lives 2 to 5 years. There is no 

published record of planting or propagation 

techniques. Because it grows rapidly and thickly in 

disturbed areas, carruzo sometimes becomes 

weedy in fields, plantations, and along roads. 
220 

Stands along roads are routinely controlled by 

mowing. 

Benefits.—Carruzo helps protect the soil in 

disturbed areas, serves as a nurse species for later 

successional species, and provides cover and 

possibly food for wildlife. 

References

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Dicotyledoneae. 

Vol. 6. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 658 p. 

Instituto Nacional de Biodiversidad. 1997. Lista de 

especímenes de Clibadium erosum. 

http://www.inbio.ac.cr/bins/k03/p13/c045/o0144

/f01362/g007091/s020661.htm. 1 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

Missouri Botanical Garden. 2003. Flora 

Mesoamericana: lista anotada—03 Feb 2003. 

http://mobot.org/cgi-bin/search_vast. [not 

paged]. 
221

Clidemia cymosa (Wendl. ex Spreng.) Alain camasey peludo 

MELASTOMATACEAE 

Synonyms: Heterotrichum cymosum (Wendl.) Urban 

Melastoma cymosa Wendl. in Spring. 

 Heterotrichum eggersii Cogn. 

John K. Francis 

General Description.—Camasey peludo, which 

means hairy melastome, is also known locally as 

camasey colorado, camasey de paloma, pelúa, and 

terciopelo. It is a medium to large shrub to 4.5 m in 

height and 7.5 cm in trunk diameter. Shrubs 1.5 to 

2.5 m in height are typical. The stems may be 

single or multiple. The bark is thin, gray-brown, 

and smooth or slightly fissured. There are 

relatively few branches. The twigs are red or 

orange near the tips due to a thick covering of 

reddish hairs, greenish below, becoming brown as 

they grow older. The young leaves are bright red or 

orange. As the leaves approach full size, the color 

fades to green on the upper surface with reddish 

accents at the edges and other areas where hairs are 

concentrated and yellow-green on the lower 

surface. The leaves are ovate with a cordate base 

and an elongated tip and have seven veins radiating 

from the base. The petioles are 4 to 9 cm long and 

the blades are 8 to 18 cm long and 7 to 12 cm 

broad. The 1.9-cm, white or pinkish flowers are 

grouped in terminal panicles. The purple berries 

that develop are about 12 mm in diameter. They 

are juicy and edible with a delightful flavor 

somewhat like blueberries. Each berry contains up 

to hundreds of tiny, light brown seeds (Liogier 

1995, Little and Wadsworth 1964). 

Range.—Camasey peludo is endemic to Puerto 

Rico (Liogier 1995). It grows mainly in the 

Luquillo Mountains, the Cordillera Central, and 

moist foothills in the eastern, northern, and western 

districts (Little and Wadsworth 1964). 

Ecology.—Camasey peludo is common and 

widespread but never abundant. It is confined to 

disturbed, open areas, such as roadsides, landslides, 

recently abandoned fields, and large tree-fall gaps. 

Camasey peludo grows in areas receiving from 

about 1600 to over 3000 mm of annual 

precipitation on soils derived from igneous and 

sedimentary rocks. These upland soils are mostly 

highly weathered clays with slow internal drainage. 

The species grows from a few meters above sea 

level to nearly 1,000 m in elevation. Camasey 

peludo are occasionally attacked (sometimes 

lethally) by white flies (Aleyrodidae) and mealy 

bugs (Pseudocoddidae, author’s observation). 

Cyrtopeltis modesta Distant (Hemiptera), 

Paurocephala heterotrichii Caldwell (Homoptera), 

and Bleparomastix ebulealis Guenée (Lepidoptera) 

have also been observed attacking camasey paludo 

(Martorell 1975). 

Reproduction.—Camasey peludo flowers and 

fruits nearly throughout the year (Little and 

Wadsworth 1964). A collection of fruits averaged 

1.298 + 0.046 g/fruit and ranged from 0.650 to 

1.998 g/fruit. Seeds separated from them averaged 

98 million seeds/kg (air dry). Sown on moist filter 

paper, 85 percent germinated between 11 and 49 

days after sowing. No data on nursery management 

of seedlings is available. Forty-four branches of 

camasey peludo were air layered by girdling and 

treating with 0.3 percent IBA in talcum, then 

covered with commercial potting mix and wrapped 

with aluminum foil. Eighty-nine percent rooted. 

However, IBA-treated cuttings placed in a mist bed 

entirely failed to root and grow. Natural layering is 

common. 
222 

Growth and Management.—Camasey peludo 

plants are relatively short-lived, lasting about 3 to 6 

years. Resprouting and layering may enable them 

to persist somewhat longer. There is no reported 

planting or management experience for this 

species. Plants cloned by layering should be used 

for planting until methods of growing the plants 

from seed are developed. Natural reproduction in 

areas of abundant rainfall may be encouraged by 

cultivation or scalping to expose a bare seedbed 

during wet portions of the year. 

Benefits.—Camasey peludo is an attractive 

roadside plant and should be encouraged whenever 

possible. Its beauty also suggests real potential as 

an ornamental, although its life is short and it has a 

tendency to unsightly decline in its last year. The 

fruit, currently almost unknown to the public, is 

very tasty and deserves attention. The fruits are a 

food source for birds. The light-brown, moderately 

hard wood has a specific gravity of 0.6 but is not 

used because of the poor form and small size of the 

plants (Little and Wadsworth 1964). 

References 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

Martorell, L.F. 1975. Annotated food plant catalog 

of the insects of Puerto Rico. Department of 

Entomology, Agricultural Experiment Station, 

University of Puerto Rico, Río Piedras, PR. 

303 p. 
223

Clidemia hirta (L.) D. Don Koster’s curse 

MELASTOMATACEAE 

Synonyms: Melastoma hirta L. 

 Clidemia crenata DC. 

 Melastoma elegans Aublet 

 Clidemia elegans (Aublet) D. Don 

John K. Francis 

General Description.—Koster’s curse (a 

Hawaiian name) is also known as clidemia, soap 

bush, camasey, camasey peludo, nigua, kak mél, 

bon bon mél, canot-macaque, and bonbon bleu. It 

is a weak upright or scrambling shrub that varies in 

mature size from less than 1 m to 5 m in height, 

depending on habitat. Koster’s curse is supported 

by shallow lateral roots and abundant fine roots. 

There are usually multiple stems arising as sprouts 

from the rootstalk and base of the principal stem, 

especially in older and disturbed plants, resulting in 

dense shrubs. The branches and twigs are slender. 

The 5- to 18-cm, opposite leaves have a short 

petiole, five to seven principal nerves radiating 

from the base, an ovate blade with a serrated to 

entire edge, usually subcordate base and pointed 

tip. The whole plant is covered with brown hairs. 

Inflorescences are branched axillary cymes with 

five-merous white flowers. The fruits are 

ellipsoidal, purple-to-black berries 6 to 8 mm long 

with a flavor similar to blueberries, but weaker. 

Each fruit contains over 100, 0.5- to 0.7-mm-long, 

coffee-colored seeds (author’s observation, 

Howard 1989, Liogier 1995, Smith 1992, Stevens 

and others 2001). 

Range.—Koster’s curse is native to the Greater 

and Lesser Antilles, southern Mexico, Central 

America, and South America as far south as 

Northern Argentina and Bolivia (Howard 1989, 

Liogier 1995, Stevens and others 2001). It has 

naturalized in Hawaii, American Samoa, many of 

the Pacific and Indian Ocean islands, Australia, 

Southern Asia, Sri Lanka, India, and East Africa 

(Binggeli 1997, Pacific Island Ecosystems at Risk 

2002). 

Ecology.—In its native range, Koster’s curse 

colonizes disturbed habitat—landslides, river 

banks, burned areas, tree tip mounds, pastures, old 

fields, plantations, fence rows, and roadsides. It 

occurs as scattered plants, occasionally as thickets, 

which flourish for a few years and succumb to 

competition or disease and insects. It tolerates a 

wide range of soil properties as long as moisture is 

adequate. Annual rainfall ranges from about 1200 

to 4000 mm. Elevation may vary from near sea 

level to 1,500 m. The species grows well in full sun 

but is moderately tolerant of shade. In some nonnative habitat in the absence of the usual insects, 

fungi, and fast-growing competitors, Koster’s curse 

forms dense and tall, monotypic thickets under 

forest canopies that shade out virtually all 

vegetation below them (Binggeli 1997, Smith 

1992). 

Reproduction.—Koster’s curse flowers and fruits 

year-round in continuously moist habitat. After dry 

seasons or droughts, in which it may defoliate or 

die back, it recovers and begins flowering again. A 

large plant may produce more than 500 fruits/year 

(Smith 1992). A collection of fresh fruits from 

Puerto Rico weighed an average of 0.2171 +

0.0081 g/fruit. Air-dried seeds separated from them 

averaged 0.00383 g/seed or 260,000 seeds/kg 

(author’s observation). The seeds are dispersed by 
224 

birds, feral pigs, other animals, and humans. The 

seeds can remain viable in the soil for up to 4 

years. The stems root on contact with moist soil; 

even detached leaves have been observed to root 

(Smith 1992). 

Growth and Management.—Koster’s curse 

grows slowly in the first months and faster 

thereafter. It often dominates at the end of the first 

year (Binggeli 1997). Individual stems live 2 or 3 

years in Puerto Rico, although plants may live 

longer by sprouting. There is little motivation to 

encourage Koster’s curse and often a need to 

control it. Single-pass mowing is usually 

ineffective. Spraying with broadleaf herbicides is 

the most widely used control method, 

supplemented by hand pulling (Rubber Research 

Institute of Malaya 1973). At best, such measures 

are temporary. Some success has been achieved 

with biocontrol by the introduction of Liothrips 

urichi, a thrips (Binggeli 1997). A number of other 

natural controls are being studied (State of Hawaii 

Department of Agriculture 2002). 

Benefits and Detriments.—Desired or not, 

Koster’s curse helps revegetate disturbed areas and 

provides a reliable food source for wildlife. The 

species is used in Brazil to treat Leishmania 

braziliensis skin infections (Franca and others 

1996). On the other hand, Koster’s curse is a 

serious threat to understory plant species in fragile 

tropical island ecosystems (Pacific Island 

Ecosystems at Risk 2002). It is a serious weed in 

tropical plantations such as rubber (Hevea 

brasiliensis Muell. Arg.). Sheep have been shown 

to control most weeds in plantations but will not 

eat Koster’s curse (Chee and Faiz 2002). Goats 

suffer toxicity from hydrolysable tannin when fed 

the plant (Murdiati and others 1990).

References 

Binggeli, P. 1997. Clidemia hirta (L.) D. Don 

(Melastomataceae). http://members.lycos.co.uk/ 

WoodyPlantEcology/docs/web-sp5.htm. 4 p. 

Chee, Y.K. and A. Faiz. 2002. Sheep grazing 

reduces chemical weed control in rubber. 

Australian Centre for International Agriculture 

Research. http://www.aciar.gov.au/publications/ 

proceedings/32/paper26.pdf. 4 p. 

Franca, F., E.L. Lago, and P.D. Marsden. 1996. 

Plants used in the treatment of leishmanial 

ulcers due to Leishmania (Vannia) braziliensis

in an endemic area of Bahia, Brazil. Revista da 

Sociedade Brasileira de Medicina Tropical 

29(3): 229-232. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Murdiati, T.B., C.S. McSweeney, R.S.F. 

Campbell, and D.S. Stoltz. 1990. Prevention of 

hydrolysable tannin toxicity in goats fed 

Clidemia hirta by calcium hydroxide 

supplementation. Journal of Applied Toxicology 

10(5): 325-331. 

Pacific Island Ecosystems at Risk. 2002. Clidemia 

hirta (L.) D. Don, Melosomataceae. http://www. 

hear.org/pier3clhir.htm. 5 p. 

Rubber Research Institute of Malaya. 1973. 

Clidemia hirta in South Johore. Planter’s 

Bulletin 128: 140-144. 

Smith, C.W. 1992. Distribution, status, phenology, 

rate of spread, and management of clidemia in 

Hawai’i. In: C.P. Stone, C.W. Smith, and J.T. 

Tunison, eds. Alien plant invasions in native 

ecosystems of Hawaii, management and 

research. University of Hawaii Cooperative 

National Park Resources Studies Unit, 

Honolulu, HI. p. 241-253. 

State of Hawaii Department of Agriculture. 2002. 

Plant pest control, persisting biological control 

problems. http://www.hawaiiag.org/hdoa/ 

pi_ppc_bioprob.htm. 6 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. A Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden Press, St. Louis, 

MO. p. 945-1,910. 
225

Coccoloba krugii Lindau wild-grape 

POLYGONACEAE 

Synonyms: none 

John K. Francis 

General Description.—Wild-grape, also known 

as white wood, crabwood, and bow-pigeon, is an 

evergreen shrub or small tree 1.5 to 6 m in height 

and stem diameters up to 10 cm. The plant is 

supported by an extensive lateral root system of 

black roots. Older plants usually have multiple 

stems arising from the root crown. The smooth or 

slightly fissured bark is gray. A dense, branchy 

crown is formed of trident or bifurcate branches 

and slender twigs. Leaves have a reddish petiole 5 

to 6 mm-long, and reddish midvein. The blades are 

coriaceous, entire, ovate to suborbicular, and 2 to 6 

cm long. This species is dioecious (male and 

female flowers on different trees). The flower 

clusters are terminal spikes. Flowers are tiny, 

greenish-white, five-merous, and nearly stalkless. 

The fruits are fleshy and black, ovoid and 

somewhat triangular in cross section, 4 to 5 mm 

long, and have a mildly sweet flavor somewhat 

like grapes. The fruit juice stains skin and clothing. 

Each fruit contains one dark brown akene (Howard 

1988, Liogier 1985, Little and others 1974). 

Range.—Wild-grape is native to the Bahamas, 

Jamaica, Hispaniola, Puerto Rico, the Virgin 

Islands, St. Martin, Anguilla, Barbuda, and 

Antigua. A natural hybrid, Coccoloba krugii x 

uvifera, with features intermediate between the two 

parents is found in Puerto Rico, the Virgin Islands, 

Anguilla, and Barbuda (Howard 1988, Liogier 

1985, Little and others 1974). 

Ecology.—Wild-grape grows on well-drained to 

excessively drained, mostly slightly acid to mildly 

alkaline soils in the whole range of soil textures. It 

is found in Puerto Rico and the Virgin Islands from 

near sea level to 125 m in elevation (Little and 

others 1974) in areas that receive from 750 to 950 

mm of annual precipitation. Wild-grape is 

intolerant of shade. Because it grows slowly, in 

order to compete it must grow on difficult sites 

such as beach strands, dry coastal forests, shallow 

rocky headlands, hillsides, and ridges. The species 

is mostly found in remnant forests and sometimes 

highly disturbed galleries. Wild-grape may occur 

as individual plants, in groups, or in thickets. 

Reproduction.—Wild-grape flowers in the fall 

and bears fruits during the winter (Little and others 

1974). A collection of fruits from Puerto Rico 

averaged 0.0985 + 0.0021 g/fruit. Air-dried seeds 

separated from them weighed an average of 0.0148 

+ 0.003 g/seed or 68,000 seeds/kg. However, 

sowing them in commercial potting mix with no 

pretreatment resulted in only 5 percent 

germination. The seeds are dispersed by birds, 

iguanas, and possibly mammals. Seedlings are not 

common. Plants sprout when cut or burned. 

Growth and Management.—Growth of wildgrape is slow, perhaps 10 to 20 cm of height per 

year. Plants are relatively long-lived and may 

renew themselves by coppicing and continue 

clonally through more than one generation of 

crowns. No management experience has been 

published. Planting any slow-growing species in 

wildlands is a challenge; protection of remnant 

forests is probably the best approach to promoting 

wild-grape. 

Benefits.—Wild-grape wood is heavy and hard and 
226 

useful for at least fuel. The fruits are edible and 

tasty, though small. The fruits are counted among 

the food sources of the endangered Anegada iguana 

(Cyclura pinguis) (International Reptile 

Conservation Foundation 2002). 

References 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

International Reptile Conservation Foundation. 

2002. Effects of introduced ungulates on the 

iguana on Anegada. http://cyclura.com/ 

article/articleview/248/1/6/. 6 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Little, E.L., Jr. R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agric. Handb. 449. U.S. 

Department of Agriculture, Washington, DC. 

1,024 p. 
227

Coccoloba microstachya Willd. pockhout 

POLYGONACEAE 

Synonyms: Coccoloba parvifolia Poir. in Lam. 

 Coccoloba klotzschiana Meisn. in DC. 

 Coccoloba microstachya var. ovalifolia Miesn. in DC. 

 Coccoloba microstachya var. lanceolata Meins. in DC. 

 Coccoloba microstachya var. rotundifolia Urb. ex Lindau 

 Coccoloba microstachya var. obtusifolia Lindau 

John K. Francis 

General Description.—Pockhout, a name used in 

the Virgin Islands, is known as uvillo, uverillo, 

and negra loca in Puerto Rico and Hispaniola. It is 

an evergreen or semideciduous shrub or small tree 

usually 2 to 6 m in height and 2 to 7 cm in 

diameter at breast height. It is limby and twiggy 

with a contorted form and often develops multiple 

stems from suckers on the lower stem. The wood 

is light brown and hard. The plants are supported 

by a weak taproot and many, extensive, reddishbrown, tough and flexible lateral roots. The fine 

roots have clearly visible mycorrhizal sheaths. The 

stem bark is gray, smooth, becoming deeply 

furrowed into plates in old individuals. The twigs 

are brown to gray, smooth with lenticels. The 

light-green, alternate leaves have short petioles, 3 

to 6 mm long, have broadly elliptic to ovatelanceolate blades 3.5 to 7 cm long, entire edges 

and generally rounded at both ends. Inflorescences 

are terminal spikes. Male and female flowers are 

borne on different shrubs (dioecious). The flowers 

are greenish white and tiny. The egg-shaped fruits 

are sessile, 6 mm long, nearly black or dark brown 

when ripe, and contain one blackish seed (Liogier 

1985, Little and others 1974). 

Range.—Pockhout is native to Hispaniola, Puerto 

Rico and its offshore islands, and the Virgin 

Islands (Liogier 1985). Little and others (1974) 

also list Anguilla as having a population. The 

species in not known to have been planted or 

naturalized elsewhere. 

Ecology.—Pockhout grows on a wide variety of 

well- to excessively-drained soils over 

sedimentary (especially limestone), igneous, and 

metamorphic (especially ultramaphic) rocks. The 

sites are usually hill tops, ridges, or upper slopes 

with shallow and rocky soils. In Puerto Rico, this 

habitat varies from 750 to about 1700 mm of mean 

annual precipitation at elevations from near sea 

level to 760 m. The species is normally dispersed 

throughout the secondary forest and does not form 

pure thickets (Weaver and others 1999). Pockhout 

is moderately intolerant of shade. It grows in 

intermediate and codominant positions in open, 

low forests as well as dominating in more difficult 

sites. 

Reproduction.—Pockhout flowers during the 

summer and fruits during the winter (Little and 

others 1974). Fruit and seed production can be 

relatively heavy in open-grown trees. Fresh fruits 

collected in Puerto Rico averaged 0.0517 + 0.0075 

g/fruit. Air-dried seeds separated from them 

averaged 70,400 seeds/kg. Sown without 

pretreatment on moist filter paper, 98 percent 

germinated beginning 16 days after sowing 

(Francis and Rodríguez 1993). Seedlings are 
228 

uncommon. 

Growth and Management.—Pockhout is slow 

growing and long lived. No published planting or 

management experience is known to the author. 

Benefits.—Pockhout contributes to the 

biodiversity of secondary and remnant forests, 

helps protect the soil, and furnishes food and cover 

for wildlife. The wood is good for fuel. 

References 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Res. Note SO-374. Southern Forest 

Experiment Station, U.S. Department of 

Agriculture, Forest Service, New Orleans, LA. 

5 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Weaver, P.L., J.L. Ramirez, and J.L. Coll-Rivera. 

1999. Las Cabezas de San Juan Nature Reserve 

(El Faro). General Technical Report IITF-5. 

International Institute of Tropical Forestry, U.S. 

Department of Agriculture, Forest Service, Río 

Piedras, PR. 58 p. 
229

Coccoloba uvifera (L.) L. seagrape

POLYGONACEAE

Synonyms: Coccolobis uvifera Jacq. 

 Polygonum uvifera L. 

John A. Parrotta 

General Description.—Seagrape, also known as 

seaside grape, bay grape, uva de playa, uvero, 

raisin de la mer, and several other common names, 

is a large, low-branching shrub or small tree with 

thick, smooth branches and a stout trunk. It is a 

coastal, tropical species easily recognized by its 

large, thick, nearly circular leaves and long, 

grapelike clusters of edible, purple fruits. 

 

Range.—Seagrape is widely distributed along the 

Atlantic, Caribbean and Pacific coasts of the 

American tropics and subtropics between 25° N. 

and 10° S. latitudes (Parrotta 1994). It is believed 

to be native to southern Florida, the Bahamas, and 

the West Indies, and is currently naturalized on 

both coasts of Mexico from Sinaloa and Veracruz 

southward through Central America and South 

America to northern Peru and northeastern Brazil. 

It has been introduced as an ornamental and 

seashore windbreak species in Hawaii and several 

tropical countries in the Asia-Pacific region 

(National Research Council 1983).

Ecology.—Seagrape is one of the first species to 

colonize sandy and rocky shores within its natural 

range, and is rarely found at any great distance 

from sea coasts (Little and Wadsworth 1964, 

Adams 1972). It is a drought-hardy, salt-tolerant 

species that grows in areas receiving between 500 

and 1500 mm of rainfall with a dry season of up to 

8 months (Von Carlowitz 1986). Its best growth 

occurs on well-drained loamy sands that are 

slightly to moderately alkaline in reaction. It can, 

however, tolerate rocky, calcareous soils and soils 

derived from igneous rocks, provided they are 

freely drained (FAO 1982). A light-demanding 

species, seagrape often forms dense thickets on 

sand dunes, and is typically associated with a 

number of other shrub, vine and tree species. 

While most commonly found in dune shrub 

formations, it does occur farther inland in coastal 

woodland formations up to 150 m in elevation in 

the Caribbean and elsewhere in its tropical and 

subtropical American range (Stoffers, 1956, Craig 

1984, Gooding 1974). Throughout its range, it is 

susceptible to attack by a number of defoliating 

insect species, as well as leaf, stem, and root 

pathogens, though none appear to pose a serious 

threat to the species (USDA 1960, Spaulding 

1961, Chellman 1978, Martorell 1975, National 

Research Council 1983, Parrotta 1994). 

Reproduction.— Flowering and fruiting in 

seagrape usually begins when trees are 6 to 8 years 

old. Its small (5 mm across), fragrant, greenishwhite, male and female flowers, borne on separate 

trees, form terminal and lateral clusters 10 to 23 

cm in length. Flowering and fruiting may occur 

throughout the year, although flowering is 

generally concentrated between January and 

August and fruiting between March and October in 

its native Caribbean range (Little and Wadsworth 

1964, Parrotta 1994). The flowers are pollinated 

by bees and other insects. The elliptic or eggshaped fruits, measuring about 2 cm in diameter, 

are grouped in clusters resembling bunches of 

grapes. The fruits ripen approximately 2 months 
230 

after flowering. The ripe, purple fruits contain a 

single, hard, elliptic seed about 1 cm long 

surrounded by an edible, tartly sweet pulp and a 

thin, fleshy covering. Fresh seed weights range 

from 1,000 to 1,400 seeds/kg, based on collections 

in Puerto Rico (Parrotta 1994). The fruits are 

consumed by a variety of birds that help to 

disperse the seeds. Natural reproduction is 

generally good on open coastal sites, although 

seedlings do not tolerate shading or competition 

from grasses and herbs. In the nursery, seagrape 

can be propagated from seed in light-textured, 

well-drained soils; no pretreatment is required to 

break seed dormancy. Seeds should be sown on or 

near the soil surface. Germination takes place 

between 18 and 50 days after sowing, and 

seedlings reach plantable size (15 cm tall) 4 to 5 

months after sowing. It can also be propagated 

vegetatively by air-layering, ground-layering, and 

grafting, though mature wood cuttings are most 

commonly used to produce rooted cuttings for 

mass propagation of selected cultivars (FAO 1982, 

Parrotta 1994). 

Growth and Management.—Growth rates in 

seagrape are strongly influenced by wind exposure; 

on exposed sites, heights typically increase from 

the windward to leeward edges of stands. On a less 

exposed coastal site with sandy soils receiving an 

annual rainfall of approximately 1500 mm in 

northeastern Puerto Rico, average heights of 0.9 

and 2.9 m were recorded for 1- and 3-year-old 

plantings (Parrotta 1994). No information is 

available on growth rates of older trees. Under 

optimal conditions seagrape can attain a height of 8 

to 18 m and a stem diameter of 45 to 60 cm 

(Record and Hess 1943). Being shade-intolerant 

and susceptible to competition from grasses, herbs 

and other woody species, planted stands should be 

regularly weeded until they are taller than 

competing vegetation. 

Benefits.—Throughout its range, the wood is used 

as firewood and for making charcoal. The reddishto dark-brown heartwood, distinguished from the 

light-brown sapwood in larger trees, is hard, closegrained, and used in turning and occasionally for 

furniture, inlay work and cabinetry (Gooding 

1974). The edible fruits can be eaten raw, used in 

preserves, or fermented like grapes to make wine. 

The flowers yield abundant nectar, and the 

resulting honey is of good quality, light amber in 

color, with a spicy flavor (National Research 

Council 1983). The astringent red sap exuded from 

the cut bark, known commercially as West Indian 

or Jamaican kino, was formerly used for tanning 

and dyeing (Little and Wadsworth 1964, Uphof 

1968). The astingent roots and bark have been 

used in traditional medicine in Puerto Rico and 

elsewhere in the Caribbean (Liogier 1990). 

Seagrape’s tolerance of saline soils and sea spray 

has made it an excellent and popular choice for 

ornamental plantings and coastal windbreaks. It 

prunes well and makes an attractive hedge. 

References 

Adams, C.D. 1972. Flowering plants of Jamaica. 

University of the West Indies, Mona, Jamaica. 

848 p. 

Chellman, C.W. 1978. Pests and problems of 

South Florida trees and palms. Florida 

Department of Agriculture and Consumer 

Services, Division of Forestry, Talahassee, FL. 

103 p. 

Craig, R.M. 1984. Plants for coastal dunes of the 

Gulf and South Atlantic coasts and Puerto Rico. 

Information Bulletin 460. U.S. Department of 

Agriculture, Soil Conservation Service, 

Washington, DC. 41 p. 

FAO. 1982. Fruit-bearing forest trees: technical 

notes. FAO For. Pap. 34. Food and Agriculture 

Organization of the United Nations, Rome. 

177 p. 

Gooding, E.G.B. 1974. The plant communities of 

Barbados. Government Printing Offic,e 

Bridgetown, Barbados. 177 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

Martorell, L.F. 1975. Annotated food plant catalog 

of the insects of Puerto Rico. Agricultural 

Experiment Station, University of Puerto Rico, 

Río Piedras, PR. 303 p. 

National Research Council. 1983. Firewood crops: 

shrub and tree species for energy production, 
231

Vol. 2. BOSTID Report 40. National Academy 

of Sciences, Washington, DC. 92 p. 

Parrotta, J.A. 1994. Coccoloba uvifera (L.) L. – 

Sea grape, uva de playa. Research Note SO-ITFSM-74. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 5 p. 

Record, S.J. and R.W. Hess. 1943. Timbers of the 

New World. Yale University Press, New Haven, 

CT. 640 p. 

Spaulding, P. 1961. Foreign diseases of forest trees 

of the world. Agriculture Handbook 191. U.S. 

Department of Agriculture, Washington, DC. 

361 p. 

Stoffers, A.L. 1956. The vegetation of the 

Netherlands Antilles. 15: Studies on the flora of 

Curaçao and other Caribbean islands. Vol. 

1.Nartinus Nijhoff, The Hague. 142 p. 

Uphof, J.T.C. 1968. Dictionary of economic 

plants. Verlag von J. Cramer, New York. 

591 p. 

USDA. 1960. Index of plant diseases in the United 

States. Agriculture Handbook 165. U.S. 

Department of Agriculture, Washington, DC. 

531 p. 

Von Carlowitz, P.G. 1986. Multipurpose tree and 

shrub seed directory. International Council for 

Research on Agroforestry, Nairobi. 265 p. 
232 

Coffea arabica L. coffee 

RUBIACEAE 

Synonyms: Coffea vulgaris Moench 

 Coffea laurifolis Salisb. 

 Coffea moka Hort. ex Heynh. 

John K. Francis 

General Description.—Coffee, also known as 

café, is an upright, evergreen shrub or small tree up 

to 5 m in height and 7 cm in diameter at breast 

height. The plant may grow with a single stem, but 

often develops multiple stems by branching at the 

base or on the lower stem. The bark is light gray, 

thin, and becomes fissured and rough when old. 

The wood is light-colored, hard, heavy, and tough. 

The root system consists of a short, stout central 

root, secondary roots radiating at all angles, and 

abundant fine “feeder” roots. The glabrous, shiny, 

dark-green, opposite leaves have petioles 4 to 12 

mm long and ovate to elliptic blades 7 to 20 cm 

long, with entire edges, and pointed at both ends. 

The fragrant, white flowers are in axillary clusters 

of two to nine. The 1.0- to 1.8-cm drupes are 

ovoid, fleshy, green turning red and finally blueblack. The fruits usually contain two greenish 

seeds, 8 to 12 mm long, that are rounded and 

flattened on one site with a medial groove. Coffee 

is an allotetraploid with 2n = 44 chromosomes 

(Bailey 1941, Howard 1989, Liogier 1997, Little 

and Wadsworth 1964, Wrigley 1988). 

Range.—The original native population of coffee 

was in the highlands of Ethiopia with possible 

disjunct populations in nearby highland areas of 

Sudan and Kenya. All of those areas have been 

altered and under semicultivation for many years 

(Charrier and Berthaud 1985). Coffee was first 

cultivated by Arabs during the 14th century and 

introduced into the New World and much of the 

rest of the tropics during the 17th century (Smith 

1985, Wrigley 1988). Today it is cultivated 

throughout the moist subtropics and high-altitude, 

moist tropics and has naturalized in many of these 

areas including Puerto Rico, the Virgin Islands 

(Little and Wadsworth 1964), Guam, and Samoa 

(Pacific Island Ecosystems at Risk 2002). 

Ecology.—The native range of coffee lies at 1,370 

to 1,830 m in elevation (Wrigley 1988). In higher 

latitudes, altitude becomes less critical. Optimum 

temperatures range from 15 to 24 °C. Growth is 

impaired above 25 °C. Frosts destroy both leaves 

and fruits (Willson 1985). The species requires a 

minimum of 1200 to 1500 mm of annual 

precipitation. Precipitation in excess of 2500 to 

3000 mm begins to be detrimental (Wrigley 1988). 

Although the species tolerates soil pH’s from 4 to 

8, pH’s of 5.2 to 6.2 are preferred. Good drainage 

is essential, and soil textures lighter than clays are 

best (Willson 1985). Coffee is tolerant of shade and 

usually grows in the forest understory. This 

environment is simulated when coffee is grown 

under Inga vera Willd. and other shade species. 

Also, it will grow well and is often cultivated in 

full sun. There are a number of serious pests and 

diseases of coffee in plantations; the effects appear 

to be negligible in dispersed naturalized 

populations. 

Reproduction.—Coffee usually flowers and fruits 

once per season, but in some areas it flowers twice. 

In Puerto Rico, flowering occurs January to April 

(Barrett 1925). It is reported to flower from 

February through May in Nicaragua (Stevens and 
233

others 2001). The flowers are insect pollinated 

(Wrigley 1988). The fruits take from 61⁄2 to 7 

months to mature (Bailey 1941). Seeds of the 

varieties naturalized in Puerto Rico vary from 

0.139 to 0.147 g/seed (Barrett 1925). Seeds 8 

weeks old germinate (95 percent in 32 days) better 

than fresh seed (Wrigley 1988). However, coffee 

seeds stored more than 21 weeks at ambient 

temperature begin to rapidly lose their viability 

(Harrer 1963). The seeds are dispersed by birds, 

and seedlings can be abundant. Artificial 

propagation is usually by seed, but the species can 

be reproduced by budding and cuttings (Center for 

New Crops and Plants Products 1996). 

Growth and Management.—Coffee has a 

moderate growth rate. Plants begin bearing in 3 to 

4 years and are in full fruit production in 6 to 8 

years. Coffee plants may live over 100 years 

(Center for New Crops and Plants Products 1996). 

Nursery plants are maintained in containers under 

shade for 6 to 12 months before outplanting 

(Wrigley 1988). Also, plantations are established 

by direct seeding into prepared seed spots (Center 

for New Crops and Plants Products 1996). Weed 

control is necessary in the first years after 

plantation establishment. Naturalized and escaped 

coffee is usually not common enough or does not 

grow fast enough to warrant control. 

Benefits.—As an exotic species in New World 

forests, coffee has had a gentle impact on 

biodiversity and contributes to wildlife food and 

cover, and soil stability. The wood is used mainly 

for fuel in the New World but is turned into chairs 

and other types of furniture in Africa (Cheney 

1925). Coffee is a good honey plant and yields a 

light-colored honey (Little and Wadsworth 1964). 

Coffee berries, edible and slightly sweet, are eaten 

occasionally by children and field workers. The 

fruit pulp, which is removed during processing, is 

sometimes fed to livestock but more often is 

composted for fertilizer and mulch (Center for New 

Crops and Plants Products 1996). Coffee seeds 

have been chewed as a stimulant in East Africa 

from ancient times (Center for New Crops and 

Plants Products 1996). The hot drink “coffee” is 

brewed from the roasted and ground seeds (or 

“beans”) and is one of the world’s most popular 

beverages. It is used to flavor candies, liquors, and 

pastries. Probably the principal reason for its 

popularity is the addictive stimulant alkaloid, 

caffeine (1,3,7-trimethylxanthine), 1.1 to 1.3 

percent in the beans, but varying greatly in the 

beverage due to different brewing practices. The 

alkaloid is present in the leaf at 0.30 percent, twig, 

0.04 percent, stem 0.01 percent, and central root, 

0.01 percent. Other alkaloids, xanthine, guanine, 

and trigonelline, which have stimulant and diuretic 

properties, are also present (Burkill 1997). Caffeine 

protects vegetative plant parts from insect and 

fungal attack and inhibits the growth of plants and 

bacteria near germinating seeds (Steiman 1997). 

Purified caffeine is widely sold as a medicinal 

stimulant, dietary aid, and headache remedy. Leaf 

poultices are used to treat sores in Trinidad, and 

root sap or root infusions are drunk to relieve 

scorpion stings (Burkill 1997). Coffee is also 

employed in folk medicine to treat asthma, flu, 

headache, jaundice, nephrosis, malaria, sores, and 

vertigo (Center for New Crops and Plants Products 

1996). 

References 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. MacMillan, New York. 3,639 p. 

Barrett, O.W. 1925. The food plants of Puerto 

Rico. Journal of the Department of Agriculture 

of Puerto Rico 9(2): 61-208. 

Burkill, H.M. 1997. The useful plants of West 

Tropical Africa. Vol. 4. Royal Botanic Gardens, 

Kew, UK. 969 p. 

Center for New Crops and Plants Products. 1996. 

Coffea arabica L. Perdue University. http://hort. 

perdue.edu/newcrop/duke_energy/Coffea_ 

arabica.html. 7 p. 

Charrier, A. and J. Berthaud. 1985. Botanical 

classification of coffee. In: M.N. Clifford and 

K.C. Willson, eds. Coffee: botany, biochemistry, 

and production of beans and beverage. The AVI 

Publishing Company, Inc., Westport, CN. p. 13-

47. 

Cheney, R.H. 1925. Coffee. The New York 

University Press, New York. 244 p. 

Harrer, A.E. 1963. Coffee growing. Oxford 

University Press, London. 127 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Dicotyledoneae. 

Part 3. Vol. 6. Arnold Arboretum, Harvard 

University, Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 
234 

436 p.

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

Pacific Island Ecosystems at Risk. 2002. Coffea 

arabica L., Rubiaceae. http://www.hear.org/ 

pier_v3.3/coara.htm. 3 p. 

Smith, R.F. 1985. History of coffee. In: M.N. 

Clifford and K.C. Willson, eds. Coffee: botany, 

biochemistry, and production of beans and 

beverage. The AVI Publishing Company, Inc., 

Westport, CN. p. 1-12. 

Steiman, S. 1997. The effects of caffeine in Coffea 

arabica L. http://www.grayskies.net/honeybear/ 

arabica.htm. 2 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and 

O.M. Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, 

No. 3. Missouri Botanic Garden Press, St. 

Louis, MO. p. 1,911-2,666. 

Willson, K.C. 1985. Climate and soil. In: M.N. 

Clifford and K.C. Willson, eds. Coffee: botany, 

biochemistry, and production of beans and 

beverage. The AVI Publishing Company, Inc., 

Westport, CN. p. 97-107. 

Wrigley, G. 1988. Coffee. Longman Scientific 

Technical and John Wiley & Sons, Inc. New 

York. 639 p. 
235

Colubrina arborescens (P. Mill.) Sarg. coffee colubrina 

RHAMNACEAE 

Synonyms: Colubrina colubrina (Jacq.) Millsp. 

 Colubrina ferruginosa Brongn. 

 Rhamnus colubrinus Jacq. 

 Ceanothus arborescens Mill. 

 Ceanothus colubrina (Jacq.) Lam. 

John K. Francis 

General Description.—Coffee colubrina, also 

known as greenheart, wild coffee, snakebark, 

blackbead tree, abeyelo, corazón de paloma, 

bijáguara, cascalata, sonzonate, and bois de fer, is 

an evergreen or semideciduous shrub or small tree. 

It grows as a shrub with multiple stems 3 to 6 m in 

height in dry areas on beach fronts, deep sands, 

and porous limestone with little soil, and as a tree 

to 25 m in height on moist sites and fertile soils. 

Bark on trunks is gray or brown, smoothish, 

fissured, or platy. Inner bark is pink, brown, or 

reddish-brown. The roots are dark brown with 

reddish inner bark and are somewhat brittle. The 

twigs, flower clusters, young leaves, petioles, and 

underside of older leaves are covered with a rusty 

pubescence. Leaves, which grow alternately in two 

rows on twigs, are papery to leathery, ovate to 

elliptic, 5 to 18 cm long, pointed at the tip and 

rounded at the base, and have prominent, curved 

veins. The tiny yellow or yellow-green flowers 

grow in short-stalked cymes in the leaf axils. 

Fruits are globose capsules 6 to 10 mm in diameter 

that split into three parts and release globose, 3- to 

3.5-mm, hard, black seeds (Liogier 1990, Little 

and Wadsworth 1964, Sargent 1923). 

Range.—Coffee colubrina is native to southern 

Florida, the Bahamas, the West Indies, and from 

southern Mexico to Panama (Liogier 1994, Little 

and Wadsworth 1964, Stevens and others 2001). 

Although the species is planted as an ornamental, 

it is not known to have become naturalized outside 

its native range. 

Ecology.—Coffee colubrina grows in a wide 

variety of well-drained soils in its native range, 

including soils of all textures, pHs ranging from 

about 5.0 to 8.0, and soils derived from 

sedimentary (including limestone), igneous, and 

metamorphic (including ultramaphic) rocks. The 

species grows in areas of Puerto Rico that receive 

from about 750 to 2500 mm of mean annual 

precipitation at altitudes from near sea level to 

about 600 m. Although the species is evergreen in 

moist sites, in dry areas, especially in excessively 

drained sites, it defoliates in response to prolonged 

drought. Coffee colubrina grows on hammocks in 

Florida (Nelson 1996) that have sandy soils 

(Gilman 1999), and in beach strand vegetation, 

coastal sands, dry to moist foothills and mountains 

in remnant and secondary forests in the West 

Indies (author’s observation). The species tolerates 

salt spray and moderate amounts of salt in the soil 

(Gilman 1999). Coffee colubrina is moderately 

intolerant; it grows in full sun and partial shade of 

low forest or broken high forest. 

Reproduction.—Coffee colubrina is reported to 

bloom from spring to fall in Puerto Rico (Little 

and Wadsworth 1964) and throughout the year in 

Florida (Long and Lakela 1976). The flowers are 

insect pollinated. Seeds collected by the author in 

Puerto Rico averaged 70,400 seeds/kg (air-dried). 

Besides minor movement by gravity, wind, and 

water, the fruits pop open when dry and fling the 

seeds a short distance (Gilman 1999). 

Growth and Management.—Growth is slow in 

dry forest areas and generally moderate in moist, 
236 

fertile soils. Four experimental plantings of coffee 

colubrina were made in Puerto Rico. Complete 

mortality occurred in the two drier, poorer sites, 

but the trees grew 1 m/year in height for the first 

few years on the better sites (Francis 1998). 

Mortality probably occurred on the drier sites 

because they were overcome by faster-growing 

competition. To ensure quick establishment, 

planting of containerized stock for ornamental and 

wildland plantings is recommended. Protection 

from weeds, vines, and faster-growing trees for 2 

or more years is imperative. Ornamental plants can 

be forced into tree shape by continual pruning of 

the lower branches (Gillman 1999). 

Benefits.—Coffee colubrina helps protect the soil, 

contributes to the aesthetics of the forest, and 

furnishes food and cover for wildlife. The plant 

attracts abundant insects including bees, wasps, 

butterflies, and diurnal moths, which in turn 

provide food for warblers (Dendroica spp.), 

gnatchtchers (Polioptila spp.), kingbirds (Tyrannus

spp.), and vireos (Vireo spp.) (Florida Fish and 

Wildlife Conservation Commission 2003). The 

species is considered a honey plant, furnishing 

both nectar and pollen to honeybees (Apis 

mellifera L.) May through September in the 

Dominican Republic (Rivas-Laureano 2003). 

Coffee colubrina is planted as a shade tree and 

ornamental in Florida, Guatemala, Nicaragua, and 

El Salvador (Little and Wadsworth 1964, Stevens 

and others 2001). It is recommended for planting 

in parking lot islands, in large containers, and 

small lawns, and for borders, screens, and hedges, 

as well as for reclamation plantings (Gilman 

1999). A tea made from the leaves and the wood is 

used as a remedy for rheumatism. Extracts are also 

used for antiseptic baths (Liogier 1990). The shiny 

black seeds are used in necklaces in Jamaica. The 

wood is hard and heavy with a specific gravity of 

0.7. The sapwood is ivory or light brown, and the 

heartwood is yellowish brown. Because it is 

resistant to decay, it is used for fence posts and 

was formerly used for marine pilings (Little and 

Wadsworth 1964). 

References 

Florida Fish and Wildlife Conservation 

Commission. 2003. Native plants for backyard 

Florida habitats. http://www.floridaconservation. 

org/viewing/inyourbackyard/nativeplants.htm. 

11 p. 

Francis, J.K. 1998. Tree species for planting in 

forest, rural, and urban areas of Puerto Rico. 

General Technical Report IITF-3. U.S. 

Department of Agriculture, Forest Service, 

International Institute of Tropical Forestry, Río 

Piedras, PR. 82 p. 

Gilman, E.F. 1999. Colubrina arborescens. Fact 

Sheet FPS-137. University of Florida, 

Cooperative Estension Service, Gainesville, FL. 

3 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc. 566 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Rivas-Laureano, S. 2003. Flora apícola 

Dominicana. http://www.apinetla.com.ar/rdom/ 

editorial-rdo/editorial1.htm. 2 p. 

Sargent, C.S. 1923. Manual of the trees of North 

America (exclusive of Mexico). Houghton 

Mifflin, Boston, MA. 910 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua, 

Angiospermas. Monographs in Systematic 

Botany Vol. 85, No. 3. Missouri Botanical 

Garden Press, St. Louis, MO. p. 1,911-2,666. 
237

Comptonia peregrina (L.) Coult. sweet fern 

MYRICACEAE 

Synonyms: Comptonia asplenifolia L. 

 Myrica asplenifolia (L.) Ait. 

Paula M. Pijut 

General Description.—Sweet fern is a densely 

branched, deciduous, rhizomatous shrub that 

grows 0.5 to 1.5 m tall and 1.2 to 2.5 m wide or 

more (Dirr 1998, Gleason and Cronquist 1991). 

Sweet fern is a dioecious or seldom monoecious 

(Gleason and Cronquist 1991), actinorhizal 

nitrogen-fixing shrub (Del Tredici 1996, Ziegler 

and Huser 1963) with fern-like leaves and stems 

that are aromatic when crushed. Leaves are 

alternate, oblong-linear, deeply pinnately lobed, 

stipulate, dark green, pubescent, 5 to 12 cm long, 1 

to 1.5 cm wide, with petioles 0.3 to 0.6 cm long 

(Dirr 1998, Gleason and Cronquist 1991, Hall and 

others 1976). Stems are green, yellowish, or 

reddish-brown when young, turning reddish-purple 

or coppery-brown with age. Buds are globular, 

minute, solitary, and sessile, with two to four 

exposed scales (Dirr 1998). The flowers are 

inconspicuous. Staminate catkins are olive green, 

cylindrical, clustered (usually three to four) on the 

tip of the previous year’s wood, 1.5 to 4 cm long, 

and drooping (Gleason and Cronquist 1991, Hall 

and others 1976). Pistillate catkins are dark red at 

the beginning of anthesis, subglobose at maturity, 

up to 0.7 cm long, with the ovary surrounded by 

eight persistent bracts (Gleason and Cronquist 

1991, Hall and others 1976). The fruit is an ovoid 

nut, olive-brown, 3 to 5 mm long, enclosed in a 

burr-like cluster of bracts (Dirr 1998, Gleason and 

Cronquist 1991). The chromosome number of 

sweet fern is 2n = 32 (Gleason and Cronquist 

1991). Two varieties are recognized: var. 

asplenifolia (L.) Fernald and var. tomentosa

A.Chev. (Krüssmann 1984, Missouri Botanical 

Garden 2002). 

Range.—Sweet fern occurs from Nova Scotia to 

North Carolina, western South Carolina, and 

northern Georgia, and west to Saskatchewan, 

Minnesota, Illinois, and Tennessee (Gleason and 

Cronquist 1991). Comptonia peregrina var. 

asplenifolia occurs only from Long Island, New 

York, to Virginia (Hall and others 1976). In 

Canada, sweet fern is found in all three Maritime 

Provinces, Ontario, and certain parts of Quebec 

(Hall and others 1976). 

Ecology.—Sweet fern prefers well-drained, dry, 

acid, sandy or gravely soils with full exposure to 

the sun (Del Tredici 1996, Schwintzer 1989, 

Snyder 1993). These sites include dry piney 

woods, exposed mountain slopes, abandoned 

pastures, barrens, highway embankments, gravel 

pits, weathered mine tailings, and cut-over forested 

land (Del Tredici 1996). Sweet fern is shade 

intolerant, adaptable to infertile soils, fixes 

nitrogen in a symbiotic association with Frankia, a 

filamentous bacterium (actinomycete) (Callaham 

and others 1978), has cluster roots that aid in 

phosphorus uptake (Hurd and Schwintzer 1997), 

and is a pioneer colonizer of sites disturbed by fire, 

logging, clear-cutting, and road construction (Del 

Tredici 1996, Lynham and others 1998, Snyder 

1993). Sweet fern is drought and salt tolerant. 

Sweet fern colonizes areas via lateral roots that 

form endogenous root buds from which shoot 

sprouts are readily formed (Louis and Torrey 

1991). The shrub is a serious weed problem in the 

commercial lowbush blueberry fields in Canada 

and New England (Hall and others 1976, Snyder 
238 

1993). The nymphs of the Saratoga spittlebug 

(Aphrophora saratogensis Fitch.) feed on sweet 

fern but do not cause serious damage (Johnson and 

Lyon 1991). Sweet fern is susceptible to attack by 

Botryosphaeria dothidea (Moug.:Fr.) Ces. & 

DeNot. (causing dieback) and Gymnosporangium 

ellisii (Berk.) Ellis (alternate host for aecia stage) 

(Sinclair and others 1987). Sweet fern is the 

alternate host of the fungus, Cronartium 

comptoniae Arth., that causes sweet fern blister 

rust on hard pines (Farr and others 1989, Sinclair 

and others 1987). The fungus is not seriously 

harmful to sweet fern, but it can reduce the growth 

and even cause death of susceptible pines, such as 

jack (Pinus banksiana Lamb.), pitch (P. rigida

Mill.), shortleaf (P. echinata Mill.), and loblolly 

(P. taeda L.) (Del Tredici 1996, Snyder 1993). 

Reproduction.—Sweet fern flowers are small, 

inconspicuous, wind-pollinated, and shed large 

amounts of pollen (Hall and others 1976). In the 

United States the shrub blossoms in April or early 

May with fruit maturation by fall. In Canada, 

sweet fern blossoms mid-May to mid-June with 

fruit maturation from July through September 

(Hall and others 1976). Hall and others (1976) 

determined the mean number of staminate catkins 

per shoot based on 42 shoots to be 6.43 + 0.48 

(SE) and for the pistillate catkins 0.60 + 0.01 (SE). 

The fruit of sweet fern is borne in a bur-like cluster 

of bracts containing on average 5.52 + 0.43 (SE) 

nuts per bur (Hall and others 1976). The pericarp 

is composed of a thin, fleshy outer layer and a 

hard, thick inner layer (Del Tredici and Torrey 

1976). Seeds do not germinate readily but can 

remain viable (for as long as 70 years) buried in 

the soil (Del Tredici 1977). Sweet fern seeds are in 

deep primary dormancy when they are shed from 

the plant and become incorporated into the soil 

(Dow and Schwintzer 1999). They enter seasonal 

cycles of secondary dormancy over time in which 

dormancy is induced during the summer. The 

seeds are then released from this dormancy by 

chilling during the winter. However, most buried 

seeds still require exposure to strong daily 

temperature fluctuations before they germinate 

(Dow and Schwintzer 1999). Strong temperature 

fluctuations indicate that shade cover has been 

removed, which stimulates the seeds to germinate 

in an open situation favorable for seedling 

establishment. Fresh seed will germinate (80 

percent) after scarification and treatment with 500 

mg/l gibberellic acid (GA3) for 24 hours (Del 

Tredici and Torrey 1976). Sweet fern propagates 

itself vegetatively via lateral roots that form 

endogenous root buds from which shoot sprouts 

are readily formed (Louis and Torrey 1991). These 

sprouts can spread extensively over large areas 

forming thickets. Juvenile stems (7.6 cm or less in 

length) root when treated with 3 g/l indole-3-

butyric acid and placed under mist (Dirr 1998). 

Root pieces (10 cm long x 0.2 cm wide, or 5 cm x 

1.3 cm) dug in late winter or early spring, placed 

horizontally at a depth of 1.3 cm in a mixture of 

fine sand and sphagnum peat, will develop shoots 

and additional roots (Dirr 1998). Sweet fern can be 

grown in containers and successfully transplanted 

(Dirr 1998). Micropropagation of sweet fern has 

been achieved by the induction of root buds in 

excised root culture (Louis and Torrey 1991). 

Growth and Management.—Sweet fern is a slow 

to medium growing shrub that develops a broad, 

flat-topped to rounded habit as it spreads and 

colonizes (Dirr 1998). It is hardy in zones two to 

six (USDA Plant Hardiness) and is sexually 

mature in 2 to 3 years (Snyder 1993). Burning and 

disking following logging will stimulate the 

growth of sweet fern (Snyder 1993). Sweet fern is 

used for erosion control, low maintenance 

plantings along highway embankments, and as 

naturalistic plantings under power company rightsof-way (Del Tredici 1996, Snyder 1993). Sweet 

fern can become weedy in pastures, old fields, and 

open woods, but can be controlled with herbicides, 

such as dicamba, 2,4-D, and glyphosate (Hall and 

others 1976, Heyd and others 1987, Snyder 1993). 

Benefits.—Sweet fern is planted as a landscape 

plant because of its fern-like, aromatic foliage and 

stems. Because it fixes nitrogen, sweet fern is 

useful for rehabilitation of disturbed sites. Sweet 

fern has limited use as a food source and cover for 

wildlife. The fruits are consumed by flickers, 

moose and deer browse sweet fern, and prairie 

chickens and sharp-tailed grouse use it for nesting 

cover (Snyder 1993). The leaves can be used for 

potpourri and tea (Stokes 1981). The chemical 

betulin occurs in the leaves, root, and stem of 

sweet fern, and its biological activities have been 

described (Duke 1996). 

References

Callaham, D., P. Del Tredici, and J.G. Torrey. 

1978. Isolation and cultivation in vitro of the 

actinomycete causing root nodulation in 

Comptonia. Science 199: 899-902. 

Del Tredici, P. 1977. The buried seeds of 

Comptonia peregrina, the sweet fern. Bulletin of 

the Torrey Botanical Club 104: 270-275. 
239

Del Tredici, P. 1996. Bulldozers and bacteria: the 

ecology of sweet fern. Arnoldia 56: 2-11. 

Del Tredici, P. and J.G. Torrey. 1976. On the 

germination of seeds of Comptonia peregrina, 

the sweet fern. Botanical Gazette 137: 262-268. 

Dirr, M.A. 1998. Manual of woody landscape 

plants, Their identification, ornamental 

characteristics, culture, propagation, and uses. 

Stipes Publishing, Champaign, IL. 1,187 p. 

Dow, M.A. and C.R. Schwintzer. 1999. Seed 

germination, seedling emergence, and seed bank 

ecology of sweet fern (Comptonia peregrina

(L.) Coult.). Canadian Journal of Botany 77: 

1,378-1,386. 

Duke, J.A. 1996. Phytochemical Database, U.S. 

Department of Agriculture, Agricultural 

Research Service, Beltsville, Maryland. 

http://www.ars-grin.gov/cgibin/duke/farmacy2.pl. 

Farr, D.F., G.F. Bills, G.P. Chamuris, and A.Y. 

Rossman. 1989. Fungi on plants and plant 

products in the United States. The American 

Phytopathological Society, St. Paul, MN. 1,252 

p. 

Gleason, H.A. and A. Cronquist. 1991. Manual of 

vascular plants of northeastern United States and 

adjacent Canada. 2nd Edition. The New York 

Botanical Garden, Bronx, NY. 910 p. 

Hall, I.V., L.E. Aalders, and C.F. Everett. 1976. 

The biology of Canadian weeds. 16. Comptonia 

peregrina (L.) Coult. Canadian Journal of Plant 

Science 56: 147-156. 

Heyd, R.L., R.L. Murray, and L.F. Wilson. 1987. 

Managing Saratoga spittlebug in pine 

plantations by suppressing sweet fern. Northern 

Journal of Applied Forestry 4: 16-17. 

Hurd, T.M. and C.R. Schwintzer. 1997. Formation 

of cluster roots and mycorrhizal status of 

Comptonia peregrina and Myrica pensylvanica

in Maine, USA. Physiologia Plantarum 99: 680-

689. 

Johnson, W.T. and H.H. Lyon. 1991. Insects that 

feed on trees and shrubs. Cornell University 

Press, Ithaca, NY. 560 p. 

 

Krüssmann, G. 1984. Manual of cultivated broadleaved trees and shrubs, Volume I, A-D. Timber 

Press, Beaverton, OR. 448 p. 

Louis, I. and J.G. Torrey. 1991. In vitro clonal 

multiplication of the actinorhizal plant 

Comptonia peregrina. Plant Cell, Tissue and 

Organ Culture 26: 89-96. 

Lynham, T.J., G.M. Wickware, J.A. Mason, and 

H. Krause. 1998. Soil chemical changes and 

plant succession following experimental burning 

in immature jack pine. Canadian Journal of Soil 

Science 78: 93-104. 

Missouri Botanical Garden. 2002. w3

TROPICOS, 

Nomenclatural database. http://mobot.mobot. 

org/W3T/Search/vast.html. [not paged]. 

Schwintzer, C.R. 1989. All field-collected 

actinorhizae examined on Comptonia peregrina

and Myrica pensylvanica in Maine are spore 

negative. Canadian Journal of Botany 67: 1,460-

1,464. 

Sinclair, W.A., H.H. Lyon, and W.T. Johnson. 

1987. Diseases of trees and shrubs. Cornell 

University Press, Ithaca, NY. 575 p. 

Snyder, S.A. 1993. Comptonia peregrina. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shrub/c

omper/index.html. 

Stokes, D.W. 1981. The natural history of wild 

shrubs and vines, eastern and central North 

America. Harper and Row Publishers, New 

York. 246 p. 

Ziegler, H. and R. Huser. 1963. Fixation of 

atmospheric nitrogen by root nodules of 

Comptonia peregrina. Nature 199: 508. 
240 

Conocarpus erectus L. button mangrove 

COMBRETACEAE 

Synonyms: Conocarpus procumbens L. 

 Conocarpus erectus var. procumbens (L.) DC. 

 Conocarpus erectus var. sericeus DC. 

 Conocarpus sericeus Jiménez 

 Conocarpus supinus Crantz 

 Terminalia erecta (L.) Baill. 

John K. Francis 

General Description.—Button mangrove is also 

known as buttonwood, sea mulberry, botoncillo, 

mangle prieto, mangle negro, palétuvier, and 

mangue. The species is usually a shrub 1.5 to 4 m 

in height but can become a tree up to 20 m or more 

in height. The root system consists mainly of 

laterals and fine roots that are dark brown, weak 

and brittle, and have a corky bark. The plant 

usually has an erect trunk or multiple trunks, but it 

may assume a prostrate form and have limbs that 

layer and become new individuals. The bark is gray 

or brown, furrowed, fibrous, and moderately thin 

(about 8 mm). The inner bark is dark cream in 

color. Stemwood (specific gravity of 1.0) is hard, 

heavy, and strong. Branches are brittle. The twigs 

are slender, yellow-green, angled, flattened, or 

winged. The spirally arranged, elliptic to lanceolate 

leaves are cartaceous to somewhat fleshy, 2 to 10 

cm long, with petioles 3 to 9 mm long. 

Inflorescences are terminal or axillary panicles of 

tiny greenish-white flowers grouped in spheroidal 

heads 3 to 5 mm in diameter. The thin, dry, 5- to 

15-mm, two-winged seeds are densely packed into 

globose clusters (Howard 1989, Liogier 1994, 

Little and Wadsworth 1964, Nelson 1996, 

Pennington and Sarukhan 1968, Stevens and others 

2001). 

Range.—Button mangrove is native to Bermuda, 

both coasts of southern Florida, the Bahamas, the 

West Indies, both coasts of Mexico, Central 

America, South America (through Ecuador and 

Brazil), and the Galapagos Islands. It is also native 

to coastal areas of Tropical West Africa (Howard 

1989, Little and Wadsworth 1964). The species has 

been planted widely as an ornamental and has 

naturalized in at least Hawaii (Pacific Island 

Ecosystems at Risk 2002). 

Ecology.—Button mangrove is intolerant of shade. 

It is almost always open-grown or in codominant 

stands. It will tolerate competition by plants of 

similar size but will not grow under the canopy of 

taller trees. The species grows above the high tide 

line, along beaches and just landward of 

Laguncularia racemosa (L.) Gaertn. and other 

mangroves. The soils in these situations are usually 

sandy or marly but sometimes loamy or clayey. 

Button mangrove is particularly adaptable as an 

ornamental because it tolerates compacted soil, air 

pollution, poor drainage, and drought (Gilman and 

Watson 1993). It also tolerates salt water overwash 

from storm surges and heavy salt spray. The 

species grows, although much less commonly, on 

the lower floodplains of rivers and in upland 

forests. Herbarium samples have been collected at 

elevations of 745 m in Costa Rica (Instituto 

Nacional de Biodiversidad 2002). 

Reproduction.—Button mangrove blooms 
241

throughout the year in México (Pennington and 

Sarukhan 1968) and from March to September in 

Florida (Nelson 1996). Tomlinson (1986) 

maintains that this species is dioecious. It is a 

consistent and abundant seed producer throughout 

most of its range. The fruit heads may contain from 

35 to 56 fruits each (Masís and others 1999). Seeds 

collected in Puerto Rico averaged 250,000 

seeds/kg. Seeds from this collection began 

germinating on moist filter paper in 9 days but 

were only 12 percent viable (Francis and 

Rodríguez 1993). The seeds are dispersed by water 

(Pacific Island Ecosystems at Risk 2002). Button 

mangrove can be propagated by partially burying 

large stakes in moist ground for living fence posts 

(Little and Wadsworth 1964). 

Growth and Management.—Button mangrove 

has a medium growth rate (Gilman and Watson 

1993) and may live for several decades. 

Information is not available on wildland plantation 

establishment. The planting of potted or 

containerized seedlings seems the safest approach. 

The species is not aggressive or invasive and only 

requires control when land is converted to other 

uses. 

Benefits.—Button mangrove provides food and 

cover for wildlife (various species of crabs and 

insects). It protects the soil during storm surges and 

helps “fix” dunes (Popp and others 1989). Both 

button mangrove and silver buttonwood (C. erectus

var. sericeus Griseb.) are widely planted as 

ornamentals in yards, parking lots, streets and 

parks (Gilman and Watson 1993, Nelson 1996). It 

can be trained to form hedges. Potted plants are 

used to form bonsai (Gilman and Watson 1993). 

Button mangrove wood is reported to be ideal for 

smoking fish and meat (Gilman and Watson 1993). 

The wood is durable but susceptible to dry-wood 

termites [Cryptotermes brevis (Walker)] and is 

used to make railroad ties, posts, for turnery, boat 

building, fuel, and charcoal. The bark and leaves 

have been used in tannery (Little and Wadsworth 

1964). Bark from Belize contains 18 percent tannin 

(Burkill 1985). Extracts of the bark are used to 

treat bleeding gums, vaginal bleeding, colic, and 

skin ulcers (Liogier 1990). 

References 

Burkill, H.M. 1985. The useful plants of West 

Tropical Africa. Ed. 2. Vol. 1. Royal Botanic 

Gardens, Kew, UK. 960 p. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. Southern 

Forest Experiment Station, U.S. Department of 

Agriculture, Forest Service, New Orleans, LA. 

5 p. 

Gilman, E.F. and D.G. Watson. 1993. Conocarpus 

erectus, buttonwood. Fact Sheet ST-179. U.S. 

Forest Service and Southern Group of State 

Foresters, Gainesville, FL. 3 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Instituto Nacional de Biodiversidad. 2002. Lista de 

especímenes de Conocarpus erectus. http:// 

www.inbio.ac.cr/bims/k03/p13/c045/o0264/f016

17/g007434/s021856.htm. 3 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR 566 p. 

Liogier, H. A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

Masís, A, D. Pérez, F. Chavarría, R. Espinoza, A. 

Guadamuz. 1999. Species page de Conocarpus 

erectus (Combretaceae), 14 April 1999. Species 

Home Pages. Area de Conservación Guanacaste, 

Costa Rica. http://www.acguanacaste.ac.cr/ 

paginas_especie/plantae_online/magnoliophyta/

combretaceae/conocarpus_erectus/c_erectus_14’

... 6 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Pacific Island Ecosystems at Risk. 2002. 

Conocarpus erectus L., Combretaceae. 

http://www.hear.org/pier_v3.3/coere.htm. 2 p. 

Pennington, T.D. and J. Sarukhan. 1968. Arboles 

tropicales de México. Instituto Nacional de 

Investigaciones Forestales, Secretaría de 
242 

Agricultura y Ganadaría, México D.F., México. 

413 p. 

Popp, M., U. Lüttge, W.J. Cram, M. Díaz, H. 

Griffiths, H.J.S. Lee, E. Medina, C. Schäfer, K.-

H. Stimmel, and B. Thonke 1989. Water 

relations and gas exchange of mangroves. New 

Phytologist 111: 293-307. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.H. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs of Systematic Botany Vol. 85, No. 

1. Missouri Botanical Garden Press. p. 1-943. 

Tomlinson, P.B. 1986. The botany of mangroves. 

Cambridge University Press, Cambridge, UK. 

419 p. 
243

Corchorus hirsutus L. jack-switch 

TILIACEAE 

Synonyms: none 

John K. Francis 

General Description.—Jack-switch, also known 

as wooly corchorus, mallet, cadillo, and 

malvavisco, is an upright woody to semiprostrate 

shrub usually about 1 m in height but sometimes 

reaching 2 m and 1.5 cm in basal diameter. It 

usually has a single main stem and a somewhat 

sparse, open crown. Plants examined by the author 

had a deep taproot and fine lateral roots at all 

levels. All parts except older branches and stems 

are densely pale scruffy tomentulose with stellate 

hairs. The alternate leaves are ovate to oblonglanceolate, 2 to 6 cm long, have crenate to serrate 

margins and obtuse to acute tips, and have petioles 

2 to 7 mm long. The leaves, which have a graygreen color, tend to wilt rather than defoliate 

during dry seasons. Yellow flowers occur in groups 

of two to eight on axillary peduncles opposite the 

leaves. The four-chambered capsules are 

ellipsoidal, about 12 mm long, and contain a 

number of 1.5- to 2-mm black seeds (Britton and 

Millspaugh 1962, Howard 1989, Liogier 1994). 

Range.—Jack-switch is native throughout the 

West Indies and has been reported from Mexico 

through Central and South America and in 

northeastern Africa (Britton and Millspaugh 1962, 

Howard 1989, Liogier 1994). Herbarium 

specimens exist for Bolivia and Paraguay 

(Missouri Botanical Garden 2002). The species is 

present as an exotic in the southern tip of Florida 

(Institute of Systematic Botany 2002). 

Ecology.—Jack-switch is intolerant of shade and 

grows in open areas or openings in low forest. It 

cannot endure severe competition. The species 

usually grows near the coast or inland on dry, often 

excessively-drained sites. These are usually coastal 

sands or rocky ridges and hillsides over limestone, 

igneous, and metamorphic (including ultramaphic) 

rocks. Jack-switch occurs as scattered plants or as 

open stands with other species of similar size. 

Because cattle do not eat or rarely eat the foliage, 

the species benefits from over-grazing. It also 

invades eroded and physically disturbed soils. 

Reproduction.—Jack-switch blooms 

continuously, except during periods of drought. 

The species is insect pollinated (Marcano-Fondeur 

1973). Capsules collected in Puerto Rico averaged 

28.0 seeds/capsule. Air-dried seeds from that 

collection averaged 1.109 million seeds/kg. Sown 

without pretreatment on the surface of wet peat, 10 

percent germinated over a 4-month period 

(author’s observation). Dispersal undoubtedly 

occurs by wind and water; specialized means of 

seed dispersal are unknown. Seedlings in wildland 

sites in Puerto Rico vary from common to rare. 

Plants apparently do not renew themselves by 

resprouting after stem death. Reaction to fire is 

unknown. 

Growth and Management.—Jack-switch plants 

have a moderate to slow growth rate and live about 

5 years. Mechanical soil disturbance near seed 

sources and heavy grazing would probably lead to 

natural establishment of the shrubs. Jack-switch is 

seldom common enough to warrant control, but if 

needed, mowing may be sufficient. 

Benefits.—Jack-switch furnishes cover for wildlife 

and helps protect the soil. The plant is attractive 

enough to be used as an ornamental in natural 

landscaping, although that use is not yet reported. 

It is a honey plant (Marcano-Fondeur 1973). The 

leaves are cooked and eaten, and made into tea in 

St. Croix, U.S. Virgin Islands. Branches were 

formerly used to make brooms (Garland 2002). 
244 

References 

Britton, N.L. and C.F. Millspaugh. 1962. The 

Bahama flora. The New York Botanical Garden, 

New York. 695 p. 

Garland, K. 2002. UVI Wetland Reserve plants: 

Corchorus hirsutus (jack-switch). 

http://rps/uvi/edu/VIMAS/jackswitch.htm. 1 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Institute of Systematic Botany. 2002. Atlas of 

Florida Vascular Plants: Corchorus hirsutus. 

University of South Florida, Tampa, FL. 

http://www.plantatlas.usf.edu/main.asp?plantID

=4142. [not paged]. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Marcano-Fondeur, E.J. 1973. La flora apícola de la 

Republica Dominicana. http://marcano. 

freeservers.com/nature/studios/apicola/dicotsp. 

html. 11 p. 

Missouri Botanical Garden. 2002. W3

-Specimen 

Data Base: Current specimen list for Corchorus 

hirsutus. http://www.mobot.mobot.org/cgibin/search_vast. [not paged]. 
245

Cordia lima (Desv.) Roemer & J.A. Schultes lija 

BORAGINACEAE 

Synonyms: Varronia lima Desv. 

 Cordia radula Spreng. 

 Cordia asperrima DC. 

John K. Francis 

General Description.—Lija, which means 

sandpaper in Spanish, is an evergreen, vine-like, 

arching shrub 1 to 2.5 m in height and 1 to 2 cm in 

basal diameter. Lija plants are supported by weak 

taproots, slender lateral roots, and abundant fine 

roots. The roots are dark brown. Older plants 

usually have a number of stems that grow 

vertically for 1 to 2 m and then arch horizontally. 

Supported on other vegetation without twining or 

tendrils, they sometimes extend 4 m or more 

laterally. The relatively few branches and twigs are 

slender. The leaves are 2 to 6 cm long, stiff and 

almost woody, ovate or elliptic, with toothed edges 

and a rough texture imparted by short, stiff hairs. 

The inflorescences are terminal or lateral 

glomerules of small white subsessile flowers. 

These produce globose one-seeded fruits (drupes) 

that are red when ripe and 4 to 5 mm in diameter. 

The fruits have a mild, pleasantly sweet flavor 

(author’s observation, Grisebach 1963, Liogier 

1995). 

Range.—Lija is native to Hispaniola, Puerto Rico, 

and the Bahamas (Grisebach 1963, Liogier 1995). 

It is not known to have been planted or naturalized 

elsewhere. 

Ecology.—Lija grows in dry to moist habitat 

(about 700 to 1800 mm of mean annual 

precipitation) at elevations from near sea level to at 

least 600 m. Colonized soils are variable in texture 

with pH’s from about 5.0 to 7.5, well drained, and 

usually rocky, on ridges, hillsides, and in draws. 

Although the species grows over other rock types, 

it is most common in limestone and ultramafic 

(serpentine) areas. Lija is moderately intolerant of 

shade and competes well with herbs and brush. It 

grows under and at the edges of remnant forests 

and middle to late secondary low forests with 

relatively low basal areas. The species varies from 

uncommon to relatively common in Puerto Rico. 

Individual plants are usually widely scattered. 

Reproduction.—A collection of fresh fruits from 

Puerto Rico averaged 0.261 + 0.010 g/fruit. Airdried seeds separated from them averaged 0.0515 +

0.0013 g/seed or 19,000 seeds/kg. Placed to 

germinate on moist filter paper, 16 percent 

germinated within 6 months. Other germination 

tests are needed to determine whether this is low 

for the species. Pretreatments do not appear 

necessary. Apparently, the seeds are dispersed by 

birds. Seedlings in natural forests are uncommon. 

Established plants sprout when disturbed. 

Growth and Management.— Lija is a slowgrowing shrub. Adult plants add only 10 to 30 cm 

to their stem length annually. Individual stems 

appear to live 1 or 2 decades while plants, by 

sprouting, may live much longer. No planting or 

management experience has been published for the 

species. Protection from fire and development are 

probably all that is required in relatively 

undisturbed sites. 

Benefits.—Lija contributes to the biodiversity in 

forests where it lives, helps protect the soil, and 

furnishes food and cover for wildlife. The fruits are 

tasty but too small and too infrequently found to 

justify collection. 
246 

References 

Grisebach, A.H.R. 1963. Flora of the British West 

Indian Islands. J. Cramer-Weinheim, New York. 

789 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 
247

Cordia polycephala (Lam.) I.M. Johnston black sage 

BORAGINACEAE 

Synonyms: Varronia polycephala Lam. 

 Varronia paniculata Wakström 

 Cordia wickstroemii Steudel 

 Cordia sulfurata Krause 

 Cordia ulmifolia Juss. ex Dumont 

 Lantana corymbosa L. 

John K. Francis 

General Description.—Black sage, also known as 

basora, palo de perico, saraguero, saragüero, Santa 

María (Spanish), petit mahot, and mahot fin 

(French), is a scrambling shrub or woody vine 

(Acevedo-Rodríguez 1985, Howard 1989, Liogier 

1995). The single or multiple stems are gray, 

cylindrical, and strong with relatively few, slender 

branches. There may or may not be a weak taproot; 

the plants rely on shallow lateral root systems with 

abundant fine roots. Alternate scabrous leaves are 

ovate to lanciolate and 8 to 15 cm long. The leaves 

have a serrate margin and are dark green above and 

yellow green below. Whitish-green flowers are 

grouped in tightly packed cymes arising near the 

branch ends. The fruits (drupes) are red, globose, 

2.5 to 5 cm in diameter, and have little flavor. The 

seed has an irregular surface (Acevedo-Rodríguez 

1985, Howard 1989, Liogier 1995). 

Range.—Black sage is native to Hispaniola, Puerto 

Rico, Vieques Island (Puerto Rico), the Virgin 

Islands, St. Barts, St. Kitts, Guadeloupe, Dominica, 

Martinique, St. Lucia, St. Vincent, Granada, and 

South America as far south as Bahia, Brazil 

(Centro Nordestino de Informações Sobre Plantas 

2002, Howard 1989). It is not known to have been 

planted or naturalized elsewhere. 

Ecology.—Black sage most frequently grows in 

abandoned fields and pastures, along roads, in early 

secondary forest, in open forest on difficult terrain, 

and in disturbed openings in old-growth forest. It 

occurs in areas receiving from 1000 to 2500 mm of 

annual precipitation, in soils of all textures and 

parent materials at elevations from near sea level to 

at least 600 m. The species does not occur in very 

poorly drained soils. Black sage is not among the 

early pioneers to colonize disturbed sites but enters 

with brush that follows the weed and grass stage. It 

competes well by scrambling across low vegetation 

and climbing into the crowns of other shrubs and 

low trees. It is moderately intolerant to shade and is 

able to grow under the canopy of low basal-area 

forest. 

Reproduction.—As a population, black sage 

blooms and fruits all year, particularly during the 

summer (Acevedo-Rodríguez 1985). However, 

individual plants usually flower at one time and 

then rest for several months before flowering 

again. The fresh fruits in one Puerto Rican 

collection averaged 0.12 g/fruit. The fruits are 
248 

consumed by birds that disperse the seeds. 

Seedlings are common but rarely abundant. 

Growth and Management.—Black sage achieves 

1 m or more of stem extension per year. Stems may 

grow to 1.5 cm of diameter and reach 5 m in height 

or extension from the point of rooting (AcevedoRodríguez 1985). One 1.5-cm stem was observed 

to have five growth rings. Black sage can live at 

least 5 years and perhaps much longer by 

sprouting, which it readily does when injured. No 

planting, management, or control experience has 

been published. In the absence of other control 

information, grubbing out individual plants or spot 

treating with broad-leaf herbicides is 

recommended. 

Benefits.—Black sage helps protect the soil and 

furnishes food and cover for wildlife. The fruits are 

edible. It is a minor weed in pastures (Vélez and 

van Overbeek 1950). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Centro Nordestino de Informações Sobre Plantas. 

2002. Base de dados CNIP, Checklist das 

plantas do Nordeste-Nomes provisórios-Versão 

12: Cordia polycephala (Lam.) Johnst. 

Universidade Federal de Pernambuco. 

http:150.161.125.13/pnechk/taxa/1057.shtml. 

2 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles. 

Vol. 6. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Río Piedras, PR. 497 p. 
249

Cornus sericea L. redosier dogwood 

CORNACEAE 

Synonyms: Cornus stolonifera Michx. 

Paula M. Pijut 

Illustration source: USDA—Forest Service Collection, 

Hunt Institute 

General Description.—Redosier dogwood is a 

multi-stemmed, deciduous, stoloniferous shrub 

that grows 1 to 3 m tall and 3 m wide or more, 

often forming dense thickets. Leaves are opposite, 

simple, ovate to oblong-lanceolate, acuminate, 

rounded at the base, 5 to 12 cm long, 2.5 to 6.4 cm 

wide, with petioles 1.3 to 2.5 cm long (Dirr 1998, 

Gleason and Cronquist 1991). Leaves are medium 

to dark green in summer, purplish to reddish in the 

fall, distinctly glaucous beneath, with five to seven 

lateral vein pairs. Stems are slender, upright, bright 

red to dark blood red, appressed pubescence on 

younger stems, with prominent lenticels, and a 

white pith (Dirr 1998). Buds are valvate, 

appressed, and elongated. The inflorescence is a 

cyme (3.8 to 6.4 cm in diameter) with small (2 to 3 

mm), white to cream-colored flowers (Gleason and 

Cronquist 1991). The fruit is a globose, white 

drupe (stone), 7 to 9 mm in diameter (Gleason and 

Cronquist 1991). The chromosome number of 

redosier dogwood is 2n = 22 (Gleason and 

Cronquist 1991). Two forms of C. sericea are 

recognized: f. baileyi (J.M. Coult and W.H. Evans) 

Fosberg and f. stolonifera (Michx.) Fosberg 

(Missouri Botanical Garden 2002). 

Range.— Redosier dogwood occurs from Alaska 

to Newfoundland, south to Virginia in the east, to 

Kansas in the Great Plains, to northern Mexico, in 

the Rocky Mountains, and through California on 

the West Coast (Crane 1989). 

Ecology.—Redosier dogwood is extremely 

adaptable to a wide range of soil and climatic 

conditions. It prefers rich, moist, poorly drained 

soils with high levels of mineral nutrients, and 

moderate to full sunlight (Crane 1989). These sites 

include swamps, low meadows, riparian zones, 

wetlands, floodplains, forest openings and 

understories. Redosier dogwood is suppressed in 

shade, a dominant understory shrub in the early 

successional willow and balsam poplar 

communities, not a primary invader of wet prairies 

and meadows (enters the stand later in succession), 

and enters a stand after fire (Crane 1989). It is 

tolerant of fluctuating water tables and extremely 

cold temperatures. Redosier dogwood can spread 

and form dense thickets (by layering) when stems 

touch the ground and roots form at the nodes. The 

larvae (instars) of the dogwood sawfly 

[Macremphytus tarsatus (Say)] feed on redosier 

dogwood causing defoliation (Johnson and Lyon 

1991). Pandemis pyrusana Kearfott (leafroller) 

attacks redosier dogwood in British Columbia and 

California (Johnson and Lyon 1991). Redosier 

dogwood is susceptible to attack by Discula 

destructiva Redlin. sp. nov. (dogwood 

anthracnose), Botryosphaeria dothidea

(Moug.:Fr.) Ces. & De Not. (canker and dieback), 

Septoria cornicola Desmaz. (leaf spot), and 

Phyllactinia guttata (Wallr.:Fr.) Lev. (powdery 

mildew) (Brown and others 1996, Jones and 

Benson 2001, Sinclair and others 1987). Tobacco 

ringspot virus has been detected in naturally 

infected redosier dogwood (Sinclair and others 

1987). Mycoplasma-like organisms have been 

detected in witches-brooms of redosier dogwood 

(Sinclair and others 1987). 

Reproduction.—Redosier dogwood flowers are 

small, self-sterile with obligate outcrossing, and 
250 

are insect-pollinated (Crane 1989). In the United 

States the shrub blossoms from late May to June 

(June to August in the Northern United States) 

with fruit maturation by fall. The fruit is a drupe 

containing a single two-celled and usually twoseeded stone, but only one seed is fully developed 

(Brinkman 1974). Redosier dogwood seeds have a 

hard seed coat and dormant embryos (Crane 1989). 

Seeds do not germinate readily (can remain viable 

in seedbanks) and require cold stratification and 

sometimes scarification in order for germination to 

occur (Crane 1989). Fresh seed can be fall sown or 

stratified for 60 to 90 days at 5 oC (Brinkman 

1974). Acharya and others (1992) found that 

native (Alberta, Canada) redosier dogwood seed 

germination was strongly influenced by the 

population from which seeds were collected, and 

the yearly precipitation in those areas. Redosier 

dogwood propagates itself vegetatively via stolons 

(stems just on or under the ground) and by 

layering forming dense thickets. Plants are also 

produced from the roots, the bases of dying 

branches, and injured stems (Crane 1989). 

Cuttings (with leaves) root readily when treated 

with 1 g/l indole-3-butyric acid (Dirr 1998). 

Hardwood cuttings root (90 to 100 percent) when 

placed in the field in late winter without a rooting 

treatment (Dirr 1998). Redosier dogwood can be 

grown as bare-root, containerized, or balled-andburlapped plants (Dirr 1998). 

Growth and Management.—Redosier dogwood 

is a fast (vigorous) growing shrub that develops a 

loose, broad-spreading, rounded habit with 

horizontal branches at the base (Dirr 1998). It is 

hardy in zones 2 to 7 (USDA Plant Hardiness) and 

is sexually mature in 3 to 4 years (Crane 1989). 

Redosier dogwood generally increases following 

fire (Crane 1989). Coppicing stimulates new, 

vigorous stem growth. Pruning and burning 

redosier dogwood helps produce long straight 

stems. Redosier dogwood is used for soil 

stabilization, reclamation sites, residential 

landscape plantings, mass plantings along highway 

banks, parks, and golf courses (Crane 1989, Dirr 

1998, Renault and others 2001). Redosier 

dogwood can be controlled by spraying with 

mixtures of 2, 4-D and dicamba (Crane 1989). 

Benefits.—Redosier dogwood is planted as a 

landscape plant because of its bright red stems, 

which are especially beautiful contrasting against a 

snowy landscape. Because it is easy to establish 

and grows rapidly, redosier dogwood is useful for 

rehabilitation of disturbed sites. It is an excellent 

soil stabilizer along stream banks because of its 

thick, extensive root system (Crane 1989). 

Redosier dogwood is relatively tolerant to high 

salinity tailings waters produced as a result of oil 

and sand mining, making it useful for reclamation 

of sites (Renault and others 1999, 2001). The bark, 

stems, leaves, roots, and fruit of redosier dogwood 

were used by Native Americans for a variety of 

uses including basket weaving, dyes, and tonics 

(Stevens and Dozier 2002). Redosier dogwood 

thickets provide cover for birds and other small 

wildlife. It is an important food source and winter 

browse for deer, elk, moose, bear, cottontail 

rabbits, snowshoe hares, grouse, quail, and other 

small birds and mammals (Crane 1989). 

References

Acharya, S.N., C.B. Chu, R. Hermesh, and G.B. 

Schaalje. 1992. Factors affecting red-osier 

dogwood seed germination. Canadian Journal of 

Botany 70: 1,012-1,016. 

Brinkman, K.A. 1974. Cornus L. dogwood. In: 

C.S. Schopmeyer, tech. coord. Seeds of woody 

plants in the United States. Agriculture 

Handbook 450, U.S. Department of Agriculture, 

Forest Service, Washington, D.C. p. 336-342. 

Brown, D.A., M.T. Windham, and R.N. Trigiano. 

1996. Resistance to dogwood anthracnose 

among Cornus species. Journal of Arboriculture 

22(2): 83-86. 

Crane, M.F. 1989. Cornus sericea. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shrub/c

omper/index.html. 

Dirr, M.A. 1998. Manual of woody landscape 

plants, their identification, ornamental 

characteristics, culture, propagation, and uses. 

Stipes Publishing, Champaign, IL. 1,187 p. 

Gleason, H.A. and A. Cronquist. 1991. Manual of 

vascular plants of Northeastern United States 

and adjacent Canada. 2nd Edition. The New York 

Botanical Garden, Bronx, NY. 910 p. 

Johnson, W.T. and H.H. Lyon. 1991. Insects that 

feed on trees and shrubs. Cornell University 

Press, Ithaca, NY. 560 p. 

Jones, R.K. and D.M. Benson. 2001. Diseases of 

woody ornamentals and trees in nurseries. The 
251

American Phytopathological Society, St. Paul, 

MN. 482 p. 

Missouri Botanical Garden. 2002. w3

TROPICOS, 

Nomenclatural database. http://mobot. 

mobot.org/W3T/Search/vast.html 

Renault, S., C. Croser, J.A. Franklin, and J.J. 

Zwiazek. 2001. Effects of NaCl and Na2SO4 on 

red-osier dogwood (Cornus stolonifera Michx) 

seedlings. Plant and Soil 233: 261-268. 

Renault, S., E. Paton, G. Nilsson, J.J. Zwiazek, and 

M.D. MacKinnon. 1999. Responses of boreal 

plants to high salinity oil sands tailings. Journal 

of Environmental Quality 28: 1957-1962. 

Sinclair, W.A., H.H. Lyon, and W.T. Johnson. 

1987. Diseases of trees and shrubs. Cornell 

University Press, Ithaca, NY. 575 p. 

Stevens, M. and I. Dozier. 2002. Redosier 

dogwood. In: U.S. Department of Agriculture, 

Natural Resources Conservation Service, The 

PLANTS Database, National Plant Data Center, 

Baton Rouge, LA. http://plants.usda.gov. [not 

paged]. 
252 

Coursetia caribaea (Jacq.) Lavin añil falso 

FABACEAE 

 

Synonyms: Cracca caribaea (Jacq.) Lavin 

 Galega caribaea Jacq. 

 Tephrosia caribaea (Jacq.) DC. 

 Benthamantha caribaea (Jacq.) Kuntze 

Brittonamra caribaea (Jacq.) Kuntze 

Tephrosia aniloides Bello 

John K. Francis 

 

General Description.—Añil falso is a low shrub, 

usually about 0.6 m in height, but sometimes 

reaching 2 m in height and 1 cm or more in basal 

diameter. The common name is Spanish from 

Puerto Rico and means false indigo. Añil falso is 

supported by a tap and lateral root system with a 

moderate amount of fine roots. The species 

produces round to elongated root nodules (Dubey 

and others 1972). The plant has a slender single 

stem that usually resprouts with a single stem 

when the plant is damaged. Plants grazed by cattle 

may be stunted or contorted. The current year’s 

mature growth is tender, but the stem is woody and 

normally without branches. The leaves are 4 to 12 

cm long, have short petioles, and 9 to 18 opposite, 

elliptic leaflets, 0.6 to 3.0 cm long. The foliage is 

yellow-green. The racemes contain one to six pealike flowers that are white or yellowish, sometimes 

lined with pink or purple. The pods are linear, 4 to 

6 cm long and somewhat compressed between 

seeds. The 10 to 25 seeds are about 2 mm long and 

have squared ends (Howard 1988, Liogier 1988). 

The diversity of the species across its range has 

resulted in the description of nine varieties 

(International Plant Index 2001). 

Range.—Añil falso is native to northern Mexico 

through Central America and South America to 

Paraguay, and to Jamaica, Hispaniola, Puerto Rico, 

the Virgin Islands, the Lesser Antilles, and 

Trinidad in the Caribbean (Liogier and Martorell 

2000). 

Ecology.—Añil falso grows at elevations from 

near sea level to over 400 m in Puerto Rico. These 

areas receive from about 750 mm to about 1200 

mm of annual precipitation. The species grows in a 

wide variety of well-drained soils that originate 

from igneous and sedimentary (including 

limestone) rocks. Añil falso may be found in brush 

lands, secondary forests, and remnant forests. It is 

moderately intolerant of shade. While it normally 

grows in the understory, it requires plenty of 

filtered light. It can tolerate moderate competition 

from shrubs, herbs, and grass, but cannot survive in 

heavy grass swards. The leaves are shed during the 

dry season. In relatively undisturbed habitat in 

Puerto Rico, plants may grow as densely as one per 

m2

. Añil falso disappears from areas that are 

overgrazed. 

Reproduction.—Of a collection made in Puerto 

Rico, air-dry pods of añil falso weighed an average 

of 0.073 + 0.005 g/pod. Eighteen pods were 

threshed and yielded an average of 12.1 seeds/pod 

with a minimum of two and a maximum of 19. 

These seeds weighed an average of 0.0028 +
253

0.0000 g/seed. They were sown without 

pretreatment on filter paper and yielded only 10 

percent germination in 206 days. After mechanical 

scarification, 98 percent of the remainder of the 

seeds germinated within 12 days. Grazing animals 

are the principal seed vectors. Seedlings are 

common and well-disbursed. Large plants are 

prolific seed producers. Even small and suppressed 

plants produce a few seeds. 

Growth and Management.—Añil falso grows up 

to about 0.5 m per year in height. Plants live from 2 

to several years. Wise management of the species 

must preclude all but light grazing. Plantations for 

range improvement or environmental restoration 

could probably be established by sowing scarified 

seed into cultivated understory sites before or at the 

beginning of the rainy season. 

Benefits.—Añil falso, when available, is grazed 

heavily by cattle. The species improves the soil by 

fixing nitrogen. 

References 

Dubey, H.D., R. Woodbury, and R.L. Rodríguez. 

1972. New records of tropical legume 

nodulation. Botany Gazette 133(1): 35-38. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University. Jamaica Plain, 

MA. 673 p. 

International Plant Index. 2001. International plant 

names index query: Coursetia caribaea

www.ipni.org/ipni/query-ipni.html. 17 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. and L.F. Martorell. 2000. Flora of 

Puerto Rico and adjacent islands. Second ed., 

Rev. Editorial fe la Universidad de Puerto Rico, 

Río Piedras, PR. 382 p. 
254 

Crataegus douglasii Lindl. black hawthorn 

ROSACEAE 

Synonyms: Crataegus brevispina Doug. ex Steud. 

 Crataegus brockwayae Sarg. 

 

John K. Francis 

General Description.—Black hawthorn, also 

known as Douglas hawthorn, river hawthorn, 

western thorn apple, and Douglas thorntree, is a 

thorny deciduous shrub or small tree 1.5 to 9 m in 

height. It may have multiple stems from the base 

or a single stem that begins branching just above 

the ground. The bark of older stems is smooth to 

scaly and gray to brown; twigs are hairless, shiny, 

slender, and reddish, and support stout, slightly 

curved spines up to 3 cm long. Leaves are 

alternate, thin, lanceolate to obovate, serrate to 

doubly serrate, and 1.5 to 9.5 cm long and almost 

as wide. Showy, five-petaled, white flowers about 

12 mm wide with 10 pink stamens are borne in 

terminal corymbs. The flowers emit a somewhat 

fishy odor. Fruits are small (12 mm in diameter), 

apple-like black pomes with thick, light-yellow 

pulp and three to five nutlets (Borialforest.com 

2003, Hansen 2003, Sargent 1923, Viereck and 

Little 1972, Welsh 1974). 

Range.—Black hawthorn is native to southern 

Alaska, British Colombia, Alberta, Saskatchewan, 

Ontario, Washington, Oregon, California, Nevada, 

Utah, Idaho, Wyoming, Montana, North Dakota, 

South Dakota, Minnesota, Wisconsin, and 

Michigan (Borialforest.org 2003, Natural 

Resources Conservation Service 2003, Soper and 

Heimburger 1982). There are two recognized 

varieties. Crataegus douglasii var. douglasii

occupies nearly all the range (Natural Resources 

Conservation Service 2003) and var. duchenensis

is found in a small area in Utah (Welsh and others 

1987). Former varieties rivularis and suksdorfii are 

now recognized as separate species (Natural 

Resources Conservation Service 2003). 

Ecology.—Black hawthorn grows mostly in forest 

understories but also occurs in the open, often in 

pure thickets. It is both a riparian species and 

grows in upland sites where adequate moisture is 

available. Slopes may be level to steep. Elevations 

vary from 670 to 1,646 m. Soils vary greatly but 

typically are deep and fine textured. The species 

usually does not occupy recently disturbed sites. 

One exception is that it is common in avalanche 

tracks in Glacier National Park. Although the 

species grows along streams, it seldom grows in 

flood-prone areas. Black hawthorn burns readily 

but is somewhat fire tolerant due to sprouting and 

suckering. However, the extent of its stands and 

thickets in eastern Washington has been limited by 

frequent fires (Habeck 1991). 

 

Reproduction.—Black hawthorn flowers open in 

May (Sargent 1923) or June (Soper and 

Heimburger 1982). The flowers are insect 

pollinated (Hansen 2003). The fruits ripen and fall 

in August through September (Sargent 1923). 

Fruits in southeastern Washington averaged 4.78 

seeds/fruit (Habeck 1991). Seeds make up about 

15 percent of the weight of ripe fruits. There are 

from 47,000 to 52,000 cleaned seeds/kg 

(Brinkman 1974). The species is difficult to 

reproduce from cuttings (Hansen 2003). Black 

hawthorn sprouts readily when cut and may sucker 

from the roots (Habeck 1991). 
255

Growth and Management.—Growth of newly 

established seedlings is slow (Habeck 1991). 

Particulars on natural stand management are 

lacking. Fruits are ripe when they are black and 

lustrous (Habeck 1991) and must be picked by 

hand or clipped with a pruning pole. Pulp should 

be removed from the seeds by maceration and then 

seeds should be air dried before sowing or storage. 

Seeds should be scarified with H2SO4 and 

stratified at 1 °C for 84 to 112 days before sowing. 

Brinkman (1974) found that germination reached 

about 30 percent in 35 to 45 days. Another source 

cites 50 to 80 percent germination (Habeck 1991). 

Dumroese and others (1997) report germination 

from 20 to 80 percent for lots treated identically. 

Seedlings of most hawthorn species develop long 

taproots and should not be kept in the nursery for 

more than 1 year (Habeck 1991). Pruning should 

be avoided and final target height of seedlings 

should be 25 cm (Dumroese and others 1997). 

Benefits.—Black hawthorn contributes to the 

beauty of the riparian vegetation, helps protect the 

soil from streambank and overflow erosion, and 

offers benefits to fauna. Although not as “showy” 

as some other hawthorns, the species is hardy and 

has been used to a limited extent as an ornamental. 

It makes excellent hedges and natural barriers, and 

is planted as a wildlife cover and food plant, and 

for stream bank stabilization (Hansen 2003). 

Livestock readily eat black hawthorn foliage but 

the thorny structure of shrubs and small trees 

makes it difficult for them to utilize it well. They 

prefer shrubs less than 1 m in height which they 

may hedge. Mule deer also browse it. With 3.7 

percent protein, 3.8 percent fat, 19.3 percent fiber, 

and 4.0 percent ash, the species is moderately 

palatable to browsing animals. Deer, upland game 

birds, song birds, and rodents eat the fruits. Black 

hawthorn makes excellent escape and nesting 

cover for wildlife (Habeck 1991). The fruits are 

made into pies, jellies, and jams (Hansen 2003). 

Native Americans ate the fruits, sometimes dried 

and mixed with fat and other ingredients. The 

wood is hard and heavy and was used for tools and 

weapons (Washington State Department of 

Transportation 2003). Native Americans used the 

thorns for piercing ears, lancing boils, and making 

fish hooks (Hansen 2003). They prepared a 

poultice of chewed leaves to relieve swelling and 

took decoctions of sap, bark, wood, and root for 

stomach problems (Moerman 1986). 

References

Borialforest.com 2003. Shrub species of the 

World’s boreal forests: Crataegus douglasii, 

black hawthorn. http://www.borealforest.org/ 

world/herbs_shrubs/black_hawthorn.htm. 2 p. 

Brinkman, K.A. 1974. Crataegus L., hawthorn. In: 

C.S. Schopmeyer, tech. coord. Seeds of woody 

plants in the United States. Agriculture 

Handbook 450. U.S. Department of Agriculture, 

Forest Service, Washington, DC. p. 356-360. 

Dumroese, R.K., K.M. Hutton, and D.L. Wenny. 

1997. Propagating woody riparian plants in 

nurseries. In: T.D. Landis and J.R. Thompson, 

tech. coords. National proceedings, Forest and 

Conservation Nursery Associations. Gen. Tech. 

Rep. PNW-GTR-419. Pacific Northwest 

Research Station, U.S. Department of 

Agriculture, Forest Service, Portland, OR. p. 71-

76. 

Habeck, R.J. 1991. Crataegus douglasii. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shrub/ 

cradou/all.html. 15 p. 

Hansen, W.W. 2003. Native plants of the 

Northwest: black hawthorn, Douglas hawthorn, 

western thorn apple, and Douglas thorntree 

(Crataegus douglasii). http://www.nwplants. 

com/plants/trees/rosaceae/crataegus_douglasii/ 

index. html. 5 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports 19. University of 

Michigan Museum of Anthropology, Ann 

Arbor, MI. 534 p. 

Natural Resources Conservation Service. 2003. 

Plant profile: Crataegus douglasii Lindl., black 

hawthorn. http://plants.usda.gov/cgi_bin/ 

 plant_profile.cgi?symbol=CRDO2. 5 p. 

Sargent, C.S. 1923. Manual of the trees of North 

America (exclusive of Mexico). Houghton 

Mifflin, Boston, MA. 910 p. 

Soper, J.H. and M.L. Heimburger. 1982. Shrubs of 

Ontario. Royal Ontario Museum, Toronto, 

Ontario, Canada. 495 p. 
256 

Viereck, L.A. and E.L. Little, Jr. 1972. Alaska 

trees and shrubs. Agriculture Handbook 410. 

U.S. Department of Agriculture, Forest Service, 

Washington, DC. 265 p. 

Washington State Department of Transportation. 

2003. Crataegus douglasii, black hawthorne. 

http://www.wsdot.wa.gov/environment/eao/ 

culres/ethbot/a-c/Crataegus.htm. 2 p. 

Welsh, S.L. 1974. Anderson’s flora of Alaska. 

Brigham Young University Press, Provo, Utah. 

724 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University Press, Provo, UT. 894 p. 
257

Critonia portoricense (Urban) Britt. & Wilson guerrero 

ASTERACEAE 

Synonyms: Eupatorium portoricense Urban 

John K. Francis 

General Description.—Guerrero, which means 

warrior in Spanish, is a large shrub or small tree to 

6 m in height and 15 cm in diameter at breast 

height. Older plants usually have multiple stems, 

arising from the root crown or lower stem. The 

wood is brittle and has a central pith 2 or 3 mm 

thick. Plants may or may not have well-defined 

taproots. Lateral roots are robust and send down 

sinker roots. The roots are tan-colored and stiff. 

Stem bark is thin, gray, and nearly smooth. The 

inner bark of guerrero is light green and bitter. 

Branches and twigs are not numerous, resulting in 

thin crowns. The hairless leaves are opposite, 7 to 

16 cm long with short petioles, ovate-elliptic 

blades with serrated or wavy-toothed edges, and 

long-pointed tips. The foliage is fragrant. The 

inflorescences are heads within large, terminal 

corymbs with tiny white flowers that produce, at 

maturity, tiny (3 mm long) achenes tipped with 3-

mm white hairs (pappus) (Liogier 1997, Little and 

others 1974). 

Range.—Guerrero is found only in Puerto Rico 

and the adjacent island of Vieques (Liogier 1997, 

Little and others 1974). It is not known to have 

been planted or naturalized elsewhere. 

Ecology.—Guerrero grows on nearly all welldrained soil types derived from sedimentary and 

igneous rocks (including limestone and 

ultramafics). Rooting is deep when soil permits, 

but the typical sites where the species grows are 

shallow over bedrock. The species is also found on 

poor and eroded soils. It grows from near sea level 

to 900 m in elevation (Little and others 1974). 

Mean annual rainfall ranges from about 850 to 

2200 mm. Although present in openings and rock 

outcrops in late secondary and remnant forests, 

guerrero is frequently common in early and middle 

secondary forests. It invades neglected pastures. 

Minor soil disturbance such as trampling by cows 

appears necessary for establishment. 

Reproduction.—Guerrero mainly flowers from 

November to February and matures fruits from 

December to March (Little and others 1974). Plants 

receiving full or nearly full sun are heavy seed 

producers. The seeds are wind-disbursed and may 

travel considerable distances. A collection of seeds 

averaged 0.00025 g/seed or about 4,000,000 

seeds/kg. Seedlings are rare to common, but never 

abundant. Resprouting occurs after adult plants are 

burned or cut. 

Growth and Management.—Guerrero grows at a 

similar rate as associated species in the difficult 

habitat where it occurs. Sprouts add about 0.5 m of 

height per year for the first 4 or 5 years. Life span 

appears to be 2 to 3 decades. No planting 

experience has been published. 

Benefits.—Guerrero is browsed by goats but 

mostly ignored by cattle. The species is sometimes 

planted as an ornamental for its fragrant foliage 

(Little and others 1974). Three pyrrolizidine 

alkaloids were isolated from the aerial parts of the 

shrub (Wiedenfeld and others 1995). As a 

medicinal plant, extracts from its leaves are said to 

relieve arthritis, and to relieve pain when applied as 
258 

a topical alcoholic rub (Liogier 1990). Guerrero 

contributes to biodiversity of the forests in which it 

grows, furnishes cover and nesting sites for birds, 

and helps protect and stabilize soil in disturbed 

areas. 

References 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Wiedenfeld, H., R. Guerrero, and E. Roeder. 1995. 

Pyrrolizidine alkaloids from Eupatorium 

portoricense. Planta Medica 61(4): 380-381. 
259

Crossopetalum rhacoma Crantz maidenberry 

CELASTRACEAE 

Synonyms: Rhacoma crossopetalum L. 

 Myginda rhacoma Sw. 

 Myginda rotundata Lam. 

 Myginda pallens J.E. Sm. in Rees 

 Crossopetalum pallena (J.E. Sm.) Kuntze 

 Rhacoma pallens (J.E. Sm.) G. Maza 

 Myginda latifolia M. Vahl 

John K. Francis 

General Description.—Maidenberry is also 

known as Florida crossopetalum, rhacoma, poisoncherry, coral, manto, maravedí, palo de paloma, 

pico de paloma, limonejo, membrillo, ti bonbon, 

bonbon rouge, and placa chiquitu. It is an 

evergreen shrub sometimes becoming a small tree. 

It commonly measures 1 to 3 m in height but may 

reach 8 m. The shrub is supported by a major 

taproot and significant lateral roots with yellow 

bark. Maidenberry usually has multiple stems from 

the root crown and just above, and many branches. 

Three shoots form at each node, one of which 

usually dies. The opposite or ternate leaves have a 

petiole about 1 mm long, lanceolate to obovate 

leathery blades usually with wavy-toothed edges. 

The tiny greenish-red flowers are borne in axillary 

cymes. The fruits that follow are fleshy, eggshaped drupes that ripen to a bright red. Each 

contains a stone and one or two seeds (Howard 

1989, Liogier 1994, Little and others 1974). 

Range.—Maidenberry is native to Southern 

Florida, the West Indies, and Venezuela (Howard 

1989, Liogier 1994, Little and others 1974, 

Alarcón 2001). Howard (1989) and Liogier (1994) 

state that is found in Mexico and Central America, 

but the species is not listed in the current check 

lists for countries in the region. 

Ecology.—Maidenberry is usually found in rocky 

and dry or excessively drained sites. It tolerates a 

wide range of well-drained soil types derived from 

sedimentary (including limestone), igneous, and 

metamorphic (including ultramafic) rocks. In 

Puerto Rico, maidenberry grows from near sea 

level to 400 m in elevation in areas that receive 

from 750 to about 1600 mm of precipitation. It is 

moderately intolerant of shade and withstands only 

moderate competition from grasses and shrubs. The 

species will survive but not prosper under low 

basal-area dry forest. The roots support vasculararbuscular mycorrhizae (TreeGuide Inc. 2002). 

Maidenberry is listed as endangered in Florida 

(Florida Fish and Wildlife Conservation 

Commission 1997); however, it is relatively 

common in Puerto Rico and several West Indian 

islands (Little and others 1974). In Florida, it is 

found in pinelands and occasionally hammocks 

(Nelson 1996). In the West Indies, maidenberry 

grows in remnant and secondary forests, most often 

on rocky ridges and hillsides. The species’ 

continued presence in overgrazed areas suggests 

that it is not palatable to cattle. 

Reproduction.—Maidenberry flowers and fruits 

year-round as a process associated with new stem 

growth (Nelson 1996). Birds disperse the seeds. 

Despite almost constant fruit production, seedlings 
260 

are not common. Plants readily sprout when cut or 

burned. 

Growth and Management.—Maidenberry grows 

slowly and may live for several decades. No 

management experience has been published. 

However, cattle grazing to reduce competition 

from grass, herbs, and vines will probably 

encourage reproduction and survival. 

Benefits.—Maidenberry furnishes food and cover 

for wildlife and protects the soil. It is listed as one 

of the food plants for the endangered Anegada 

iguana, Cyclura pinguis (International Reptile 

Conservation Foundation 2002). The wood of 

maidenberry is light brown and hard, but is used 

only for fuel. All the plant tissues, especially the 

roots, are diuretic and are used to promote 

urination and to treat kidney and bladder infections 

(Liogier 1990). 

References 

Alarcón, C. 2001. Paraguana xeric scrub 

(NT1313). World Wildlife Fund. 

http://worldwildlife.org/wildworld/profiles/terre

strial/nt1313_full.html. 14 p. 

Florida Fish and Wildlife Conservation 

Commission. 1997. Florida’s endangered 

species, threatened species and species of 

special concern. http://floridaconservation.org/ 

pubs/endanger.html. [not paged]. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

International Reptile Conservation Foundation. 

2002. Effects of introduced ungulates on the 

iguana of Anegada. http://www.cyclura.com/ 

article/articleview/248/1/6/. 12 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR 566 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. 461 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

TreeGuide Inc. 2002. Florida crossopetalum, 

Crossopetalum rhacoma Crantz, Celestraceae. 

http://www.treeguide.com/nn/Species.asp? 

SpeciesID=345&Region=NorthAmerican. 2 p. 
261

Crotalaria lotifolia L. cascabelillo axilar 

FABACEAE 

Synonyms: Crotalaria lotifolia L. var. eggersii Senn 

 Crotalaria lotifolia L. var. grandiflora Urban 

 Crotalaria lotifolia L. var. grandifolia Urban 

John K. Francis 

General Description.—Cascabelillo axilar is a 

suffruticose or relatively short-lived shrub usually 

1 m in height and 6 mm in basal diameter but 

sometimes reaching 2 m in height and more than 1 

cm of basal diameter. It usually has a single stem, 

stiff and straight in its lower portions, which 

branches sparingly. The wood is moderately hard 

and tough. The branches are slender. Stem bark is 

yellowish-green and striated. The plant is 

supported by a tap and lateral root system with 

moderately stiff roots with yellow bark. Foliage is 

concentrated near the top of the plant. Leaves are 

trifoliate on long (2 to 6 cm) petioles. Leaflets are 

light green or yellow-green, oblong to elliptic, and 

1 to 5 cm long. The inflorescences are fewflowered, axillary racemes whose unequal flowers 

have a yellow corolla 5 to 7 mm long. The legumes 

(pods) are 2 to 3 cm long and 6 mm wide and 

contain a few tan seeds that rattle when dry 

suggesting the Spanish name cascabelillo, which 

means little rattlesnake (author’s observations, 

Howard 1988, Liogier 1988). 

Range.—Cascabelillo axilar is native to the 

Bahamas, the Greater Antilles, St. Barts, Dominica, 

Martinique, the Grenadines, Barbados, Mexico, 

and Honduras (Howard 1988, Liogier 1988). 

Ecology.—In Puerto Rico, cascabelillo axilar 

grows from a few meters above sea level to about 

500 m elevation on the southern slopes of the 

Sierra Central. These areas receive from 750 to 

about 2200 mm of mean annual precipitation. It 

grows on a variety of soils derived from igneous, 

sedimentary (including limestone), and 

metamorphic (including ultramaphic) rocks 

(Breckon 2002). Cascabelillo axilar is moderately 

intolerant of shade. It grows in openings and in the 

understory of low basal-area forest, often in rocky 

and excessively drained areas. The forests are 

remnants and middle to late secondary. The species 

does not tolerate competition from tall grass swards 

and thick herb and brush growth. 

Reproduction.—Cascabelillo axilar probably 

flowers near the end of the wet season. Plants with 

mature pods were observed by the author in Puerto 

Rico in March. Seed production can be abundant. 

Eighteen pods collected in Puerto Rico averaged 

6.7 seeds per intact pod. An unknown seed insect 

had attacked some of the pods. Seeds collected 

from the same area averaged 0.0086 + 0.0008 

g/seed or 117,000 seeds/kg. Sown on moist blotter 

paper without pretreatment, 35 percent germinated 

in 5 months. Seeds from the same batch, 

mechanically scarified, all germinated in 12 days. 

Germination is epigeal. Seed pods spring open 

upon drying flinging the seeds a short distance. 

Grazing animals probably also move seeds. 

Seedlings are relatively common on sites where the 

species is present. 
262 

Growth and Management.—Once established, 

cascabelillo axilar adds about 0.5 m in height each 

year and loses most of it to die-back the following 

dry season. Individual plants probably live 1 to 3 

years or more. No management experience has 

been published. Scarified seed sown and 

incorporated into prepared ground or seed spots at 

the start of the wet season may offer a good chance 

of success. 

Benefits.—Cascabelillo axilar contributes in a 

minor way to standing biomass, cover for wildlife, 

the aesthetics of the forest understory, and it helps 

protect the soil. The species is browsed by cattle 

and consequently is more common in protected 

areas. Cascabelillo axilar is present in moderately 

grazed areas but absent from seriously overgrazed 

areas. 

References 

Breckon, G.J. 2002. Preliminary checklist for 

Sierra Bermeja. http://www.uprm.edu/biology/ 

profs/breckon/herbarium/FLORABERMEJA. 

htm. [not paged]. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University. Jamaica Plain, 

MA. 673 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 
263

Crotalaria spectabilis Roth showy rattlebox 

FABACEAE 

Synonyms: Crotalaria leschenaultii DC. 

 Crotalaria macrophylla Weinm. 

 Crotalaria retzii Hitchc. 

 Crotalaria sericea Retz. 

John K. Francis 

General Description.—Showy rattlebox, also 

known as showy crotalaria and rattlebox, is an 

annual semiwoody herb in temperate areas, and a 

short-lived shrub in its frost-free subtropical and 

tropical range. In Puerto Rico, it grows up to 2 m in 

height and 3.5 cm in basal diameter. It normally 

has a single stem. The lower stems are covered 

with whorled, short, shelf-like old leaf bases and 

stipule remnants. The leaves and fruiting branches 

are shed progressively upward after they mature so 

that foliage and branches are only found on the 

upper portion of the stem. The mid- and lower stem 

has a white, brittle, medium-hard wood with a 1.5-

mm pith. The upper stem, branches, and foliage are 

green to yellow-green. The plant is supported by a 

tap and lateral system of stiff, tan roots. The 

simple, whorled leaves are oblanceolate to elliptic, 

5 to 17 cm long, entire, and have a 2-mm petiole 

and broad, triangular stipules. The inflorescences 

are terminal or subterminal racemes with 20 to 25 

flowers with linear-triangular bracts. The bright 

yellow unequal flowers are 1.5 to 2 cm long. 

Inflated brown to black legumes are 3 to 5 cm long 

and 1.8 to 2 cm thick. They contain several hard, 

shiny brown to black seeds 4.5 mm long (author’s 

observation, Damron and Jacob 2001. Howard 

1988, Liogier 1988, Stevens and others 2001). 

Range.—Showy rattlebox is native to the IndoMalaysia area (Parrotta 2001). It has been planted 

widely and has naturalized in many tropical 

countries including the Southern United States, 

Hawaii, and Puerto Rico (International Legume 

Database and Information Service 2002). 

Ecology.—Not much is known about showy 

rattlebox in its natural habitat. It grows on 

roadsides and disturbed areas in Florida (Long and 

Lakela 1976), and in Puerto Rican coastal sands 

and limestone rubble in areas receiving about 1650 

mm of annual precipitation (author’s observation) 

and at the edge of mangroves (Liogier 1988). 

Parrotta (2001) reports that it is common along 

stream banks in deciduous forests in India. The 

species is a weed of agricultural crops in the 

Southern United States (Bradley and Hagood 

2002). It is intolerant of shade and does not grow 

under a forest canopy. The species seems to favor 

sandy soils (McGregor 1976), grows well in 

alkaline soils, and at least tolerates moderate soil 

salinity. Plants are often parasitized by dodder 

(Cuscuta spp.) in Puerto Rico. Showy rattlebox is 

well nodulated with nitrogen-fixing bacteria 

(McGregor 1976). 

Reproduction.—Showy rattlebox blooms all year 

in Florida (Long and Lakela 1976) but is reported 

to flower in April and May and fruit in October in 

Nicaragua (Stevens and others 2001) and flower 

between November and January and fruit from 

December to February in India (Parrotta 2001). 
264 

The flowers are visited by honey bees and other 

insects (McGregor 1976). A collection of seeds 

from Puerto Rico averaged 0.0154 + 0.0000 or 

65,000 seeds/kg. Only 3 percent of unscarified seed 

placed on moist blotter paper germinated in 6 

months. However, 99 percent of scarified seed 

germinated between 4 and 7 days after sowing. 

Germination is epigeal. When sufficiently dry, 

pods spring open, flinging the seeds a short 

distance. 

Growth and Management.—Showy rattlebox has 

a moderate to fast growth rate and may reach a 

meter of height by 1 year old. Plants appear to live 

2 or 3 years in Puerto Rico. Parrotta (2001) refers 

to the species as biennial. Plantations are 

established by sowing scarified seed into prepared 

ground. 

Benefits and Detriments.—Showy rattlebox is a 

beautiful plant and contributes to the aesthetics of 

wildlands and gardens where it is planted. It serves 

as a nurse species during early reforestation and 

helps protect the soil. Showy rattlebox is used as a 

green manure crop, particularly where rotation for 

control of nematodes is important (Zago and others 

2002). It has also been grown as a fodder plant. 

This has largely been abandoned because of the 

toxic alkaloids (principally monocrotaline) it 

contains. Concentrations are reported as: leaves 

0.008 percent, seedpods 0.366 percent, and seeds 

1.958 percent (Burkill 1995). Chickens, turkeys, 

and quail are very sensitive. Concentrations of 

crotalaria alkaloids between 0.01 and 0.1 percent of 

the diet have adverse effects and 0.3 percent is fatal 

(Damron and Jacob 2001). Horses, cattle, and 

swine are also sensitive. Sheep, goats, and dogs are 

more resistant. Effects can be both acute and 

chronic and include bloody diarrhea, anemia, 

jaundice, hair loss, and unthriftiness. There is no 

known treatment (Alabama Cooperative Extension 

Service 2002). In herbal medicine, extracts of the 

whole plant are used to treat impetigo and scabies 

(Jain and De Filipps 1991), as an antiseptic for 

cuts, and to treat intestinal worms (Parrotta 2001). 

References 

Alabama Cooperative Extension Service. 2002. 

Poisonous plants of the Southeastern United 

States: Showy crotalaria Crotalaria spectabilis. 

http://www.aces.edu/dept/extcomm/publications

/anr/anr-975/Showy%20crotalaria.pdf. 1 p. 

Bradley, K. and S. Hagood. 2002. Virginia Tech 

weed identification guide: Showy crotalaria: 

Crotalaria spectabilis. http://ppws.vt.edu/scott/ 

weed_id/cvtsp.htm. 2 p. 

Burkill, H.M. 1995. The useful plants of West 

Tropical Africa. Vol. 3. Royal Botanic Garden, 

Kew, UK. 857 p. 

Damron, B.L. and J.P. Jacob. 2001. Toxicity to 

poultry of common weed seeds. University of 

Florida, Cooperative Extension Service. 

http://edis.ifas.ufl.edu/BODY_PS052. 7 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University. Jamaica Plain, 

MA. 673 p. 

International Legume Database and Information 

Service. 2002. ILDIS Legumes of the world: 

Crotalaria spectabilis Roth. http://ildis.org/ 

LegumeWeb/6.00/taxa/4077.shtml. 4 p.

Jain, S.K. and R.A. De Filipps. 1991. Medical 

plants of India. Vol. 1. Reference Publications, 

Inc., Algonac, MI. 408 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyan Books, Miami, FL. 

962 p. 

McGregor, S.E. 1976. Insect pollination of 

cultivated crop plants. Internet printing by the 

USDA Agriculture Research Service, http:// 

gears.tucson.ars.ag.gov/book/chap9/crotalaria. 

html. [not paged]. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CABI Publishing, Oxon, UK and New 

York. 917 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

1. Missouri Botanical Garden Press, St. Louis, 

MO. p. 945-1,910. 
265

Zago, C., V.P. de Melo and W.R. Maluf. 2002. A 

aplicação da adubação verde em hortaliças. 

Boletim Técnico de Hortaliças 16. 

Departamento de Agricultura, Universidade 

Federal de Lavras. http://www2.ufla.br/ 

~wrmaluf/bth016/bth016.html. 5 p. 
266 

Cuphea hyssopifolia Kunth false heather 

LYTHRACEAE 

Synonyms: Parsonia petiolata (L.) Rusby 

 Cuphia petiolata Koehne 

John K. Francis 

General Description.—False heather is sometimes 

called cuphia, clammy cuphia, tarweed, elfin herb, 

or simply, heather (Griffiths 1992, Pope 1968). It is 

a woody, much branched shrub, although 

diminutive in size. While Howard (1989) states 

that the species may reach 70 cm in height and 

Turner and Wasson cite 60 cm as maximum height, 

25 to 30 cm are more typical (Liogier 1994). A 

definite leader is maintained, but many of the 

lateral branches grow along the ground surface. 

The bark is reddish brown and stringy. The root 

system consists of a short taproot and many laterals 

of equal thickness. The tertiary roots are fibrous. 

The leaves are subsessile, 1 to 3 cm long and 

linear, lanceolate, or oblanceolate. Young stems 

and the underside of leaves are pubescent. The 

small white, pink, violate, or reddish purple 

flowers appear in the axils of new leaves (Turner 

and Wasson 1997). There 5 to 8 ovules in each 

flower, and generally six 1-mm seeds per fruit 

(Howard 1989). 

Range.—False heather is native from Mexico to 

Panama (Howard 1989, Liogier 1994). It has been 

planted widely and has naturalized and escaped in 

at least Hawaii (Pope 1968), Puerto Rico (Liogier 

1994), Montserrat, and St. Lucia (Howard 1989). 

Ecology.—A moderate amount of disturbance is 

required for false heather to establish and maintain 

itself. Mowing, animal and vehicular traffic, and 

light cultivation are typical enabling disturbances. 

A moist climate (above 1400 mm of annual 

precipitation) is required for natural stands. It does 

not tolerate frost or salt (Watkins and Sheehan 

1975). Partial shade from trees and light 

competition from grass and weeds are tolerated, 

but tall grass and heavy weeds will eliminate it. In 

Hawaii, false heather is most abundant in moist and 

partially shaded places above 240 m elevation 

(Pope 1968). Clayey and medium textured soils 

with good drainage are best. Japanese beetles 

(Popillia japonica Newman) sometimes defoliate it 

in Hawaii (Pope 1968). 

Reproduction.—Flowering and seed production 

takes place constantly. The seeds are tiny (1.3 

million per kg) and difficult to collect. The period 

between flowering and seed formation is short, and 

the seeds are expelled from the fruits at maturity. A 

germination test of 94 seeds on filter paper resulted 

in only two seeds germinating, over a period of 14 

days. Seedlings are not uncommon on moist, 

disturbed soil. False heather also reproduces by 

layering whenever horizontal branches touch the 

ground. Both seedlings and layers are important in 

the maintenance of natural stands. Obtaining new 

plants from seed is difficult. Nursery plants are 

reproduced by cuttings; tips will root any time of 

year (Watkins and Sheehan 1975). Artificial 

reproduction by air and ground layers is also 

possible. 

Growth and Management.—False heather is a 

short-lived plant except when layering prolongs its 

life. After about 1 year, most plants begin a decline 

in vigor and eventually die. Renewal of decadent 
267

older plants can be promoted by mowing at about 5 

cm height, which cuts away the old plant structure 

from the newly layered branches. Nursery plants 

should be watered during dry periods or as needed 

and given frequent light fertilizer applications. 

False heather is susceptible to nematodes. Use of 

nematode-free media is recommended (Watkins 

and Sheehan 1975). 

Benefits.—False heather has been cultivated 

widely as an ornamental. It is used as a potted 

plant, for low borders, and as ground cover in small 

beds. False heather may become a weed in warm 

climates (Turner and Wasson 1997) and has 

become a serious weed in some parts of Hawaii 

(Pope 1968). 

References 

Griffiths, M. 1992. Index of garden plants. Timber 

Press, Portland, OR. 1,234 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Pope, W.T. 1968. Manual of wayside plants of 

Hawaii. Charles E. Tuttle Co, Publishers, 

Rutland, VT. 289 p. 

Turner, R.G., Jr. and Wasson, E. 1997. Botanica. 

Barns and Noble, Inc. New York. 1,020 p. 

Watkins, J.V. and T.J. Sheehan. 1975. Florida 

landscape plants, native and exotic. The 

University Press of Florida, Gainsville, FL. 

420 p. 
268 

Curatella americana L. chaparro 

DILLENIACEAE 

Synonyms: none 

Ariel E. Lugo and Ernesto Medina 

General Description.—Chaparro in English is 

known as the rough-leaf tree and it is also known 

as careicillo, peralejo macho, vacabuey, aperalejo, 

curatela, peralejo, peralejo de sabana, pomme 

torche, raspa viejo, tlachicón, hoja man, yaha, 

lengua de vaca, malcajaco, hoja chigüe, raspa, 

raspa guacal, chumico de palo, hoja chigüe, yahal, 

saha, chumico, azufre, curata, paricá, chaparro de 

sabana, chaparro sabanero, chaparro colorado, 

parica, feuille rude, curatahie, carne de fiambre, 

caimbe, cajueiro bravo, sambaiba, caimbahiba, 

cambaiba, sobro, sabaibinha, and chaáco (Liogier 

1983, 1990, Standley and Record 1936, Salas 

Estrada 1993, Siegel and Paguaga 1991, Vargas 

Ulate 2001, Standley 1920, 1937, Pérez Arbelaez 

1978, Record and Mell 1924, Bastos 1984, Kileen 

and others 1993). Curatélla means “plants being 

used or worked”, for example, for polishing 

weapons and metal (Bailey 1941). It is an 

evergreen woody shrub less than 50 cm tall to a 

tree 6 to 10 m in height (Lundell 1937, Liogier 

1990, Killeen and others 1993). Its most frequent 

habit is that of a gnarled shrub or small tree 

(Sarmiento and Monasterio 1983). 

In a family dominated by lianas, chaparro 

is a twisted, thick-barked shrub/tree with 

pubescent young branches (Schnee 1960). Leaves 

are simple, alternate, oval to elliptic oval, 5 to 12 

cm wide and 10 to 15 cm long, rounded to 

emarginated, pinnately veined, hard, pubescent 

(when young), and with very scabrous surfaces 

and sinuous margins (Pittier 1926, Gentry 1993, 

Killeen and others 1993). Leaf borders are dentate. 

The apex is retuse and the base is decurrent on a 

short petiole (Schnee 1960). Stipules are 

deciduous and form small scars on the stem. 

Leaf morphology contributes to reduction 

of heat loads and prevents overheating (Medina 

1982). Stomata are slightly elevated and the leaf 

epidermis is heavily cutinized (Mérida and Medina 

1967). Leaves of chaparro are scleromorphic, with 

low specific leaf area (<100cm2

/g) and low 

nutrient concentrations (Mérida and Medina 1967, 

Medina 1977, 1982, Sarmiento and Monasterio 

1983). 

Throughout its range, chaparro is most 

commonly associated with the following species: 

trees such as Bowdichia virgilioides and 

Byrsonima crassifolia; and grasses and sedges 

such as Leptocoryphium lanatum and 

Trachypogon plumosus; and species of 

Andropogon, Axonopus, Bulbostylis, Paspalum, 

and Rhynchospora (Sarmiento 1983). In Belize, 

chaparro grows in pine forests (Standley and 

Record 1936). Pollen from chaparro appeared in 

Suriname and Colombia in strata dated at 45,000 

years before present at the beginning of the 

Pleistocene and during glacial times (Van der 

Hammen 1983). Chaparro pollen was associated 

with grass pollen and dry periods and has shown 

cycles of abundance and disappearance in response 

to climate change. The presence of chaparro pollen 

in lake sediments established the evolution of 

savanna vegetation in the Neotropics before the 

presence of humans (Sarmiento 1983). Chaparro 

pollen also established the boundary between rain 

forests and savannas in Bolivia and the southern 

expansion of the rain forest boundary over the past 

3,000 years in response to climate change (Mayle 

and others 2000), and vegetation changes in 

southeastern Brazil and Colombia since the 

Holocene and late Quaternary (Behling 1995, 

Behling and Hooghiemstra 1999). 

Range.—Chaparro has a large geographic range 

that includes Cuba (Grisebach 1963); Hispaniola; 

the states of Tepic, Tabasco, Chiapas, Veracruz, 

Campeche, Oaxaca, and Guerrero of México 

(Standley 1920); Honduras; Belize (Standley and 

Record 1936); Nicaragua; Costa Rica; Panamá 

(Woodson and Schery 1965); Guatemala; El 

Salvador; French Guiana; Suriname; Guyana; 

Trinidad; Venezuela; Colombia (Pérez Arbelaez 

1978); Perú (Macbride 1956); Brazil; and Bolivia 

(Beard 1953, Sarmiento 1983, Liogier 1990, Salas 

Estrada 1993). 
269

Genetics.—Comparative flavonoid chemistry of 

chaparro and other members of the Dilleniaceae 

family resulted in reticulate distribution of the 

substances identified and did not permit 

recognition of taxa beyond the level of genus and 

family (Gurni and Kubitzki 1981). Woodson and 

Schery (1965) reported pentagyna as a variety of 

chaparro in Panamá. 

Ecology.—Chaparro is a savanna plant with a 

uniform spatial distribution (San Jose and others 

1991). It is adapted to fire, high air temperature, 

and harsh soil conditions (Pittier 1926, Lundell 

1937). Chaparro is among the first invaders after a 

fire (Salas Estrada 1993). Medina and Silva (1990) 

describe the savannas where chaparro grows as 

“steady state (s) regulated by water-fire 

interactions on a background of low nutrient 

availability.” However, during the early phases of 

growth, chaparro is susceptible to fire disturbance 

(San José and Medina 1975). A low allocation of 

N to leaves--even young ones--reduces 

photosynthetic capacity, but it is adaptive to fire as 

only a small amount of N is lost when leaves burn 

(Medina and Francisco 1994). 

Chaparro grows up to 500 m elevation in 

Bolivia (Killeen and others 1993), and Venezuela 

(Sarmiento 1983), and it has extended its range in 

Colombia to up to 500 to 1,500 m due to 

savannization of forest lands (Cavalier and others 

1998). Frost appears to limit its northern and 

southern distribution, and flooding or soil 

saturation also limits its distribution (Sarmiento 

1983). 

High acidity, high clay content, low 

mineral nutrient concentration, high exchangeable 

Al, poor drainage, and poor aeration in the deeper 

soil horizons favor chaparro (Lundell 1937, 

Gottsberger and Morawetz 1986). Chaparro will 

grow in depressions. In some sites, chaparro 

dominates the community and forms dense groves. 

For example, within a matrix of flooded savannas 

in Bolivia, chaparro forms groves on ant and 

termite hills and on hummocks (Killeen and others 

1993). Sahales is the name used to describe groves 

of chaparro in northern Petén (Lundell 1937). 

Chaparro grows in a wide range of moisture 

conditions from dry to wet savannas. In the 

Venezuelan Llanos, San José and Medina (1975) 

report a mean annual rainfall of 1335 mm and a 

range of 580 to 1990 mm. A “normal” year has a 4 

to 6 month dry season. Beard (1953), Sarmiento 

and Monasterio (1975), Sarmiento (1983), Medina 

and Silva (1990), and Medina and Huber (1992) 

report details of the climatic and edaphic 

conditions of tropical American savannas where 

chaparro is one of the most common woody 

species. 

Chaparro grows in many soil types. The 

most favorable are well-drained soils. Examples of 

soils are red sands, red sandy loams, yellow sands, 

well drained latosols, hard laterite crusts, illdrained latosols, deeply weathered quartz gravel 

and sands, serpentine-derived soils, entisols on 

well drained sites, entisols on young alluvial 

surfaces to oxisols on more ancient ones, alfisols, 

and vertisols (Sarmiento 1983). Vargas Ulate 

(2001) attributed the presence of chaparro in 

Guanacaste, Costa Rica, to acid and infertile soils. 

In Colombia, the loss of the A and B horizons of 

granitic soils in sierra Nevada have led to 

savannization and the establishment of chaparro in 

lands where it normally does not grow (Cavalier 

and others 1998). 

Chaparro has a deep root system reaching 

the water table in its adult stage. The species is a 

phreatophyte, so it grows new leaves during the 

dry season (Sarmiento and Monasterio 1983, 

Medina and Francisco 1994). 

Competition between chaparro and other 

plants in the savanna occurs in two stages: 

establishment and subsequent growth. During 

establishment, grasses may hinder the growth of 

saplings and constitute a risk because of the high 

fire frequency. Chaparro roots face water shortages 

as they grow through the grass root layer, so 

establishment is favored by sequences of wet years 

when fire frequency diminishes and soil moisture 

increases. Eventually, chaparro will out-grow 

grasses and shade them out (for example in tree 

groves). 

The chlorophyll content of chaparro is 

low and correlated with low N and high ash 

concentrations (Medina and Francisco 1994). Leaf 

photosynthesis rate reached maximum values in 

mature leaves during the humid season and 

declined in younger and older leaves and during 

the dry season (Sobrado 1996b). Young leaves are 

more efficient in water use than mature leaves. 

Stomatal conductance, and water loss, 

decline during the dry season. Chaparro leaves do 

not show photoinhibition. Stomatal conductance 

differences between dry and wet season appear to 

be responsible for maintaining a homeostatic 

balance of leaf water and thus maintain xylem 

integrity in spite of the species vulnerability to 

embolisms (Sobrado 1996a). Predawn water 

potential, minimum leaf water potential, and 

hydraulic resistance of chaparro, did not show 

significant differences throughout the year. In spite 

of the scleromorphic appearance of leaves, 

chaparro appears adapted only to short-term 
270 

diurnal deficits (Meinzer and others 1983). The 

transpiration curves for chaparro during the dry 

season are always bimodal (Medina 1977, 1982) as 

plants close their stomata at noontime, 

independently of water availability (Medina 1967, 

1983). Transpiration rates are also low. The 

dependency of leaf dark respiration on air 

temperature changes seasonally (Medina 1982). 

The steepness of the relationship decreases from 

December, to February, to March, and to 

September through November, when the 

relationship is flat. 

The fungus Pseudocercospora curatellae

was isolated from leaves of chaparro (Furlanetto 

and Dianese 1999). The earthworm Pontoscolex 

roraimensis was associated with chaparro (Guerra 

Torquemada 1994). Epiphytes are not abundant on 

chaparro bark, probably due to its flaky deciduous 

nature (Gottsberger and Morawetz 1993). Leaf 

herbivory on chaparro was 4.6 percent near 

Cuiaba, Mato Grosso, Brazil (Nascimento and 

others 1990). 

Reproduction.—The white flowers have a 

“disagreeable smell” (Grisebach 1963), occur in 

lateral panicles on defoliated old branches, and are 

rarely terminal. Flowers are actinomorphic and 

hermaphroditic. Four to five petals are white or 

pinkish, 5 to 6 mm, free, and imbricate (Liogier 

1990). Four to five green sepals are rounded, 5 

mm long, persistent, and externally pilose (Schnee 

1960). Stamens are numerous, free, and persistent. 

The gynoecium has two hirsute carpels of 6 to 7 

mm (Liogier 1990) with longitudinal dehiscence 

(Schnee 1960). They are coned at the base and 

covered by long and abundant hairs. Follicles are 

globose, hispid, coned in the lower half, with two 

seeds (Killeen and others 1993). Flowers and fruits 

of chaparro are fasciculate and ramiflorous 

(Gentry 1993). Chaparro flowers between July and 

November in Bolivia (Killeen and others 1993) 

and December to February in Venezuela (Montes 

and Medina 1977). Seeds are oblong, bright, 

chestnut brown to black, 3 mm long, with a small 

membranous hilum (Schnee 1960, Liogier 1983). 

Seeds are long-lived and remain buried until the 

proper conditions for germination occur (Medina 

and Francisco 1994). Fire induces germination and 

sprouting of stems that are too thick to sprout 

under normal conditions (Foldats and Rutkis 

1969). Seeds are dispersed by birds (Salas Estrada 

1993), including parrots and parakeets who eat 

chaparro fruits (Pérez Arbelaez 1978). 

Growth and Management.—Chaparro 

assimilates carbon all year, but its photosynthetic 

capacity is reduced due to high mesophyll 

resistance to CO2 transfer (Medina 1982). Growth 

is seasonal and its flowering is tardy (Sarmiento 

and Monasterio 1983). It flowers and produces 

new leaves during the dry season (Medina 1982). 

Leaf development takes 45 days (Nascimento and 

others 1990), and leaf longevity is about 11 

months (Montes and Medina 1977). However, 

young, mature, and old leaves overlap in an 

individual (Medina and Francisco 1994). Fruiting 

and seed dispersal may continue for a short time 

during the rainy season. Fire does not interrupt this 

cycle as burned leaves fall and within days there is 

a new flush. 

Two hundred randomly selected trees of 

chaparro growing in the seasonal savannas of 

western Venezuela had a mean stem diameter at 

ground level of 8.3 cm. The largest recorded 

diameter was 39.8 cm (Sarmiento and Monasterio 

1983). Chaparro wood has no growth rings and its 

sapwood is not clearly defined (Record and Mell 

1924). 

Under a regime of annual burning, 

chaparro will experience die back of all its annual 

growth shoots, and growth will start the next 

season from adventitious buds that develop from 

vascular cambium of older branches (Sarmiento 

and Monasterio 1983). As a result of this 

response, “the tree behaves as a mechanical 

structure supporting a crop of annual branches” 

(Sarmiento and Monasterio 1983). The vascular 

cambium assumes the role of apical meristem as 

the main replacement tissue. Chaparro can 

maintain itself as a hemixyle--with annual shoots 

and woody underground structures--and even 

complete the reproductive process in this growth 

form (Sarmiento and Monasterio 1983). Chaparro 

can develop new individuals from injured roots 

(Foldats and Rutkis 1975). Usually, saplings that 

develop from injured roots have higher water 

potentials than normal saplings because they are 

already connected to the water table. Injury to 

roots will stimulate vegetative regrowth, which in 

turn also facilitates formation of groves (San Jose 

and others 1991). 

The mineralization of nitrogen, soil 

organic matter, soil exchangeable Ca, Mg, and K, 

and total soil nitrogen were higher in soils under 

chaparro trees than soils under savanna grasses. 

For soil organic matter, K, and cation exchange 

capacity, chaparro exceeded values measured 

under deciduous trees in savannas (García 

Miragaya and others 1994, Sánchez and others 

1997). Although chaparro is not an aluminum 

accumulating plant, it exhibits seasonal variation 

in its nutrient concentration (Villela and de 
271

Lacerda 1992). Potassium, Fe, Cu, and Zn were 

highest in August, Ca and Mg in November, and 

Al and Mn in May. The Al and Mn concentration 

of mature chaparro leaves is 171 and 105 parts per 

million, respectively (Medina 1977). 

Benefits.—Chaparro produces good wood with a 

reddish hue and specific gravity of 0.85. Record 

and Mell (1924) and de Matos Araujo and de 

Matos Filho (1977) contain wood anatomy 

information for chaparro. Small stem sizes limit 

wood utilization of shrubs and trees. However, the 

wood is durable and used for charcoal, fuelwood, 

posts, small cabinetwork, and for turning (Record 

and Mell 1924, Standley and Record 1936, Killeen 

and others 1993, Salas Estrada 1993). Termites do 

not attack chaparro wood (Pérez Arbelaez 1978). 

Wood utilization increases where trees or shrubs 

become locally abundant and grow in groves 

(Pittier 1926). People use dry leaves as substitutes 

for sand paper and for cleaning kitchen utensils or 

polishing metals (Killeen and others 1993). 

Chaparro bark produces tannins that provide a 

gray color when used for dying pelts (Killeen and 

others 1993). Ground seeds mixed with chocolate 

flavors the drink in Oaxaca, México (Standley 

1920). Chaparro was one of the most abundant 

pollens in honey samples during the months of 

January, February, and March from an apiary 

north of Roraima State, Brazil (da Silva and Absy 

2000). Siegel and Paguaga (1991) suggested using 

chaparro twigs and pods for prospecting for gold 

in Costa Rica. In Venezuela, Clamens and others 

(1999) evaluated chaparro for industrial gum 

production and found the species promising. 

Pharmaceuticals.—Chaparro is a medicinal plant 

used extensively in folk medicine throughout its 

range. Infusions from leaves and stems are used 

for arthritis, diabetes, and to lower blood pressure 

(Killeen and others 1993, Liogier 1990). Guerrero 

and others (2002) found that a dose of 20 mg/kg of 

ethanolic extracts of chaparro showed significant 

antihypertensive activity in rats. In Brazil, 

chaparro is used for treatment of inflammation and 

ulcer. Anti-inflammatory and analgesic tests with 

hydroalcoholic extract of bark were successful in 

mice (Alexandre Moreira and others 1999). 

Cooked leaves are used to mitigate skin eruptions, 

for dressing wounds, and the water for purifying 

blood (Liogier 1990). 

References 

Alexandre Moreira, M.S., M.R. Piuvezam, A.C. 

Araujo, and G. Thomas. 1999. Studies on the 

anti-inflammatory and analgesic activity of 

Curatella americana L. Journal of 

Ethnopharmacology 67:171-177. 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. The Macmillan Co. New York. 

Bastos, M.D.N.D.C. 1984. Floristic survey of 

fields of the state of Para Brazil. 1. The field of 

Joannes Marajo Island. Boletim do Museu 

Paraense Emilio Goeldi Serie Botanica 1:67-86. 

Beard, J.S. 1953. The savanna vegetation of 

northern tropical America. Ecological 

Monographs 23:149-215. 

Behling, H. 1995. A high-resolution Holocene 

pollen record from Lago do Pires, SE Brazilvegetation, climate and fire history. Journal of 

Paleolimnology 14:253-268. 

Behling, H. and H. Hooghiemstra. 1999. 

Environmental history of the Colombian 

savannas of the llanos Orientales since the last 

glacial maximum from lake records El Pinal and 

Carimagua. Journal of Paleolimnology 21:461-

476. 

Cavalier, J., T.M. Aide, C. Santos, A.M. Eusse, 

and J.M. Dupy. 1998. The savannization of 

moist forests in the Sierra Nevada de Santa 

Marta, Colombia. Journal of Biogeography 

25:901-912. 

Clamens, C., Leon de Pinto G., F. Rincon, V. 

Antonio, and O. Beltran. 1999. The behavior of 

Samanea saman and Curatella americana as 

gum producers. Boletín del Centro de 

Investigaciones Biológicas Universidad del Zulia 

32(2): 67-78. 

da Silva, S.J.R. and M.L. Absy. 2000. Analyses of 

pollen found in honey samples of Apis mellifera

L. (Hymenoptera, Apidae) in a savanna area in 

Roraima, Brazil. Acta Amazonica 30:579-588. 

Foldats, E. and E. Rutkis. 1969. Sprouting of 

Curatella americana as affected by fire. XI 

International Botanical Congress Abstracts, 

p. 61. 

Foldats, E. and E. Rutkis. 1975. Ecological studies 

of chaparro (Curatella americana) and manteco 

(Byrsonima crassifolia) in Venezuela. Journal of 

Biogeography 2:159-178. 
272 

Furlanetto, C. and J.C. Dianese. 1999. Some 

Pseudocercospora species and a new Prathigada 

species from the Brazilian cerrado. Mycological 

Research 103: 1,203-1,209. 

García Miragaya, J., S. Flores, and N. Chacón. 

1994. Soil chemical properties under individual 

evergreen and deciduous trees in a protected 

Venezuelan savanna. Acta 

Oecologica/International Journal of Ecology 15: 

477-484. 

Gentry, A.H. 1993. A field guide to the families 

and genera of woody plants of northwest South 

America (Colombia, Ecuador, Perú). 

Conservation International Washington, DC. 

895 p. 

Gottsberger, G. and W. Morawetz. 1986. Floristic 

structural and phytogeographical analysis of the 

savannas of Humaita, Amazonas, Brazil. Flora 

Gena 178:41-71. 

Gottsberger, G. and W. Morawetz. 1993. 

Development and distribution of the epiphytic 

flora in an Amazonian savanna in Brazil. Flora 

188:145-151. 

Grisebach, A.H.R. 1963. Flora of the British West 

Indian Islands. Wheldon & Wesley LTD and 

Hafner Publishing Co. New York. 

Guerra Torquemada, R. 1994. Earthworm activity 

in forest and savanna soils near Boa Vista, 

Roraima, Brazil. Acta Amazonica 24: 303-307. 

Guerrero, M.F., P. Puebla, R. Carrón, M.L. Martin, 

L. Arteaga, and L San Román. 2002. 

Assessment of the antihypertensive and 

vasodilator effects of ethanolic extracts of some 

Colombian medicinal plants. Journal of 

Ethnopharmacology 80:37-42. 

Gurni, A.A. and K. Kubitzki. 1981. Flavonoid 

chemistry and Systematics of the Dilleniaceae. 

Biochemical Systematics and Ecology 9: 109-

114. 

Killeen, T.J., E. García, and S.G. Beck. 1993. Guía 

de árboles de Bolivia. Missouri Botanical 

Gardens and Herbario Nacional de Bolivia. La 

Paz, Bolivia. 598 p. 

Liogier, A.H. 1983. La flora de la Española. II. 

Universidad Central del Este Volumen 44 Serie 

Científica XV. San Pedro de Macorís, República 

Dominicana. 

Liogier, H.A. 1990. Las Plantas medicinales de 

Puerto Rico y del Caribe. Iberoamericana de 

Ediciones. San Juan PR. 

Lundell, C.L. 1937. The vegetation of Petén. 

Carnegie Institute of Washington, Washington 

D.C. 244 p. 

Macbride, J.F. 1956. Flora of Perú. Botanical 

Series Field Museum of Natural History 

Chicago IL. 13(3a)(2): 677. 

Mayle, F.E., R. Burbridge, and T.J. Killeen. 2000. 

Millennial-scale dynamics of southern 

Amazonian rain forests. Science 290: 2,291-

2,294. 

Medina, E. 1967. Intercambio gaseoso de árboles 

de las sabanas de Trachypogon en Venezuela. 

Boletín No. 111 Sociedad Venezolana de 

Ciencias Naturales 27: 56-69. 

Medina, E. 1977. Introducción a la ecofisiología 

vegetal. Monografía 16 Serie de biología. 

Programa regional de desarrollo científico y 

tecnológico. Departamento de Asuntos 

Técnicos, Secretaría General de los Estados 

Americanos. Washington, D. C. 102 p. 

Medina, E. 1982. Physiological ecology of 

neotropical savanna plants. In: B.J. Huntley and 

B.H. Walker, eds. Ecology of tropical savannas. 

Springer Verlag, Berlin. p. 308-335 

Medina, E. 1983. Adaptations of tropical trees to 

moisture stress. In: F.B. Golley, ed. Tropical 

rain forest ecosystems, A. structure and 

function. Elsevier, Amsterdam, The 

Netherlands. p. 225-237 

Medina, E. and M. Francisco. 1994. 

Photosynthesis and water relations of savanna 

tree species differing in leaf phenology. Tree 

Physiology 14:1367-1381. 

Medina, E. and O. Huber. 1992. The role of 

biodiversity in the functioning of savanna 

ecosystems. In: O.T. Solbrig O.H.M. van Emden, 

and P.G.W.J. van Oordt, eds. Biodiversity and 

global change. Monograph 8. International 

Union of Biological Sciences, Paris. p. 139-158 
273

Medina, E. and J.F. Silva. 1990. Savannas of 

northern South America: a steady state regulated 

by water-fire interactions on a background of 

low nutrients availability. Journal of 

Biogeography 17: 403-413. 

Meinzer, F., V. Seymour, and G. Goldstein. 1983. 

Water balance in developing leaves of 4 tropical 

savanna woody species. Oecologia 60: 237-243. 

Mérida, T. and E. Medina. 1967. Anatomía y 

composición foliar de árboles de las sabanas de 

Trachypogon en Venezuela. Boletín 111 

Sociedad de Venezuela de Ciencias Naturales 

27:45-55. 

Montes, R. and E. Medina. 1977. Seasonal 

changes in nutrient content of leaves of savanna 

trees with different ecological behavior. Geo 

Eco Trop 4: 295-307. 

Nascimento, M.T., Villela, D.M., and L.D. de 

Lacerda. 1990. Foliar growth longevity and 

herbivory in two cerrado species near Cuiaba 

Mato Grosso Brazil. Revista Brasileira de 

Botanica 13(1):27-32. 

Pérez Arbelaez, E. 1978. Plantas útiles de 

Colombia. Litografía Arco, Bogota, Colombia. 

Pittier, H. 1926. Manual de las plantas usuales de 

Venezuela. Litografía del Comercio. Caracas, 

Venezuela. 458 p. 

Record, S.J. and C.D. Mell. 1924. Timbers of 

tropical America. Yale University Press, New 

Haven, CN.

Salas Estrada, J.B. 1993. Árboles de Nicaragua. 

Instituto Nicaragüense de Recursos Naturales y 

del Ambiente. Managua, Nicaragua. 388 p. 

Sánchez, L.F., J. García Miragaya, and N. Chacón. 

1997. Nitrogen mineralization in soils under 

grasses and under trees in a protected 

Venezuelan savanna. Acta 

Oecologica/International Journal of Ecology 18: 

27-37. 

San José, J.J. and E. Medina. 1975. Effect of fire 

on organic matter production and water balance 

in a tropical savanna. In: F.B. Golley and E. 

Medina, eds. Tropical ecological system. 

Springer Verlag, NY. p. 251-264 

San Jose, J.J., M.R. Fariñas, and J. Rosales. 1991. 

Spatial patterns of trees and structuring factors in 

a Trachypogon savanna of the Orinoco Llanos. 

Biotropica 23: 114-123. 

Sarmiento, G. 1983. The savannas of tropical 

America. In: F. Bourlière, ed. Tropical savannas. 

Elsevier, Amsterdam, The Netherlands. p. 245-

288 

Sarmiento, G. and M. Monasterio. 1975. A critical 

consideration of the environmental conditions 

associated with the occurrence of savanna 

ecosystems in tropical America. In: F.B. Golley 

and E. Medina, eds. Tropical ecological system. 

Springer Verlag, NY. p. 223-250 

Sarmiento, G. and M. Monasterio. 1983. Life 

forms and phenology. In: F. Bourlière, ed. 

Tropical savannas. Elsevier, Amsterdam, The 

Netherlands. p. 79-108. 

Schnee, L. 1960. Plantas comunes de Venezuela. 

Revista de la Facultad de Agronomía de la 

Universidad Central de Venezuela, October: 

230. 

Siegel, F.R. and A.S. Paguaga. 1991. Curatella 

americana L.--a biogeochemical sample medium 

for the Tilaran-Montes del Aguacate gold belt, 

Costa Rica. Journal of Geochemical Exploration 

41: 169-180. 

Sobrado, M.A. 1996a. Embolism vulnerability of 

an evergreen tree. Biologia Plantarum 38: 297-

301. 

Sobrado, M.A. 1996b. Leaf photosynthesis and 

water loss as influenced by leaf age and seasonal 

drought in an evergreen tree. Photosynthetica 

32:563-568. 

Standley, P.C. 1920. Trees and shrubs of México. 

Contributions from the United States National 

Herbarium 23(1):819. 

Standley, P.C. 1937. Flora of Costa Rica, part II. 

Field Museum of Natural History, Chicago IL. 

p. 690. 

Standley, P.C. and S.J. Record. 1936. The forests 

and flora of British Honduras. Field Museum of 

natural History Botanical Series 12: 254. 

Van der Hammen, T. 1983. The palaeoecology and 

palaeogeography of savannas. In: F. Bourlière, 
274 

ed. Tropical savannas. Elsevier, Amsterdam, The 

Netherlands. p. 19-35 

Vargas Ulate, G. 2001. Phytogeography of dry 

ecosystems in the ignimbrite plateau of 

Guanacaste, Costa Rica. Revista de Biología 

Tropical 49: 227-238. 

Villela, D.M. and D. de Lacerda. 1992. Leaf 

mineral element dynamics in two “cerrado” tree 

species. Revista Brasileira de Biología 52:151-

160. 

Woodson, R.E. and R.W. Schery. 1965. Flora of 

Panamá. Annals Missouri Botanical Garden 52: 

587-588. 
275

Cytisus scoparius (L.) Link Scotch broom 

FABACEAE 

Synonyms: Sarothamnus scoparius (L.) Wimmer ex Koch 

 Genista scoparius Lam. 

 Spartium scoparium L. 

John K. Francis 

Drawing source: Britton and Brown 1913. 

General Description.—Scotch broom is also 

known as broom, common broom, Scot’s broom, 

English broom, green broom, broom tops, Irish 

tops, witch’s broom, basam, escoba negra, and 

retama de escobas. It is a shrub, usually 1 to 2, 

occasionally 3 m in height, which sheds its leaves 

early and remains leafless for most of the year. 

The green twigs are the principal photosynthetic 

organs. Branches are upright to arching. The 

angular, flexible twigs often occur in moderately 

long sprays. Old stems become gray-brown. The 

plant grows a deep root system with a taproot. 

Leaves are trifoliate low on the twigs and simple 

and sessile near the tips. The obovate to lanceolate 

leaflets are 4 to 8 mm long, and petioles are 2 to 

10 mm long. It’s perfect flowers, which are 

solitary in leaf axils or in clusters of two or three, 

are about 2 cm long, brilliant yellow, and have the 

typical form of flowers in the pea family. Orange, 

red, pink, purple, and lilac flowers are available in 

ornamental varieties. The fruits are dark brown, 

linear pods 4 to 5 cm long that are ciliate at the 

margins and contain five or six seeds each. The 

seeds are 3 to 4 mm long, greenish brown, and in 

the form of a flattened oval. There are 2n = 46 or 

48 chromosomes (Abrams 1944, Meyer 2003, 

New England Wild Flower Society 2003 

University of Connecticut 2003, Welsh and others 

1987). 

Range.—Scotch broom is native to Western and 

Eastern Europe, including the Mediterranean 

islands, the British Isles, and western Scandinavia. 

It has been planted and naturalized in Eastern and 

Western United States, Hawaii, Canada, the 

Azores, the Canary Islands, Australia, New 

Zealand, South Africa, India, China, and parts of 

southern South America (International Legume 

Database and Information Service 2003, Natural 

Resources Conservation Service 2003). Two 

varieties of Scotch broom have naturalized in the 

United States. Variety andreanus occurs only in 

California and var. scoparius accounts for the rest 

of the naturalized range (Natural Resources 

Conservation Service 2003). 

Ecology.—Scotch broom grows as scattered 

plants, small clumps, and pure stands, and 

competes fiercely with forbs, grasses, and tree 

seedlings. It requires disturbance, especially 

disturbance resulting in bare mineral soil to 

become established. It prefers sandy soils, but will 

grow in a wide variety of well-drained soils 

(Prasad 2003). Scotch broom is drought tolerant. 

The species is intolerant of shade and cannot 

survive under a closed forest stand. Nitrogen 

fixation occurs in root nodules, and fixed nitrogen 

is released to the environment in the litter (Parker 

and others 1994). Top injury from subfreezing 

temperatures may occur in cold climates, but 
276 

plants usually survive (Michigan State University 

Extension 1998). Scotch broom is commonly seen 

along roads, in vacant lots and inactive 

construction sites, in old logging areas, in 

neglected pastures and orchards, and along fencerows. 

Reproduction.—Scotch broom flowers in May or 

June (New England Wild Flower Society 2003). 

The flowers are pollinated by honey bees and 

bumblebees (Meyer 2003). Seed production starts 

when plants are 2 years old in Tasmania with 

around 300 seeds/year and increases to 12,000 to 

30,000 seeds/shrub/year. The seeds are spread by 

explosive release from the pods, which can throw 

the seeds 1 to 3 m (Alchemy-works 2003), as well 

as by water, soil movement, and machinery 

(National Haritage Trust 2003). Ants also disperse 

seeds. Seeds average 125 seeds/g. (Meyer 2003). 

Seeds can remain viable in the soil seed bank for 

as long as 80 years (National Haritage Trust 2003). 

The species can also be propagated with cuttings 

(University of Connecticut 2003). 

Growth and Management.—Scotch broom 

grows from 30 to 46 cm/year (Michigan State 

University Extension 1998) and usually lives from 

10 to 15 years (Prasad 2003). Fruits can be picked 

by hand after they are mature but before they burst 

open. They should be spread to dry, threshed, and 

screened. These seeds are orthodox and can be 

stored for many years. Because they are hardcoated, scarification is needed to ensure adequate 

and uniform germination. This can be done by 

nicking or sanding the seed coat or by applying 

dry heat for 1 minute at 130 °C or 15 minutes at 70 

°C. Viability is normally about 80 percent. 

Because roots are delicate, the use of containerized 

stock is recommended for planting rather than 

bare-root stock (Meyer 2003). However, because 

the species is invasive, planting for conservation or 

ornamental purposes is usually discouraged. In 

fact, the species has been declared a noxious weed 

in five Western States where planting is prohibited 

(Natural Resources Conservation Service 2003). 

Scotch broom is not a problem in cultivated land 

or in land fully timbered, only in pastures, 

roadsides, sites normally occupied by low 

vegetation, and in regenerating timberlands. 

Control of Scotch broom is difficult. Grubbing 

large plants and pulling seedlings, or power 

cultivation, effectively kills the plants, but new 

seedlings continually appear from the soil seed 

bank. Mowing only encourages branching near the 

ground. Stands of Scotch broom do not carry a fire 

readily (Parker and others 1994), and burned 

stands reproduce prolifically. The application of 

herbicides has sometimes been effective (Prasad 

2003). A number of insect enemies of Scotch 

broom have arrived accidentally or have been 

introduced into the United States. They may 

already be having a minor impact on the 

competitive ability of the species (Syrett and 

others 1999). Research into biological control 

continues, particularly in Australia and New 

Zealand (CSIRO 2003). 

Benefits.—Scotch broom, which produces a 

spectacular annual floral display and possesses a 

unique form, has been planted throughout the 

temperate regions of the world as border and 

accent plants. It is particularly useful in this regard 

because it succeeds on poor soils with little care. It 

has also been used successfully to protect exposed 

soils and stabilize dunes. Scotch broom furnishes 

food (mainly from the seeds) and cover for 

wildlife. The leaves and twigs are generally 

unpalatable to domestic livestock and wild 

mammals. The tissues contain the alkaloids 

cytisin, sparteine, and isosparteine, and the 

glycoside scoparin. However, large amounts must 

be ingested before illness or death results (Everest 

and others 2003). The stems and twigs have been 

used from ancient times to make brooms and were 

used for ritual brooms called besoms. The species 

was also used to make baskets, thatching, wattle 

fencing, and a green dye, and to flavor beer before 

the introduction of hops (Alchemy-works 2003). 

Decoctions of the twigs and leaves were and are 

still occasionally used to treat gout, jaundice, 

edema, and other conditions (Grieve 2003). 

References

Abrams, L. 1944. Illustrated flora of the Pacific 

States. Vol. 2. Stanford University Press, 

Stanford, CA. 635 p. 

Alchemy-works. 2003. Cytisus scoparius-Scotch 

broom. http://www.alchemy-works.com/ 

cytisus_scoparius.html. 1 p. 

Britton, N.L. and A. Brown. 1913. Illustrated flora 

of the northern states, Canada, and the British 

possessions. Vol. 2, 2nd Ed. Scribner, New York. 

735 p. 

CSIRO-European Laboratory. 2003. Our research: 

Scotch broom-Cytisus scoparius Link, Fabaceae. 

http://www.csiro-europe.org/cytisus.html. 2 p. 
277

Everest, J.W., T.A. Powe, Jr., and J.D. Freeman. 

2003. Poisonous plants of the southwestern 

United States. Alabama Cooperative Extension 

System. http://www.aces.edu/dept/extcomm/ 

publications/anr/anr-975.pdf. 56 p. 

Grieve, M. 2003. Broom. http://www.botanical. 

com/botanical/mgmh/b/broom-70.html. 9 p. 

International Legume Database and Information 

Service. 2003. Cytisus scoparius (L.) Link. http: 

//www.ildis.org/LegumeWeb/6.00/taxa/6046. 

shtml. 4 p. 

Meyer, S.E. 2003. Cytisus scoparius (L.) Link. In: 

F.T. Bonner and R.G. Nisley, eds. Woody plant 

seed manual. U.S. Department of Agriculture, 

Forest Service, Washington, DC. 

http://wpsm.net/Cytisus.pdf. 4 p. 

Michigan State University Extension. 1998. 

Cytisus scoparius—Scotch broom. http://www. 

msue.msu.edu/msue/imp/modop/00000485.html. 

2 p. 

National Heritage Trust. 2003. English broom 

(Cytisus scoparius). http://www.bushcare. 

tas.gov.au/ToAdd/Data/weeds/English%20%20 

broom.pdf. 2 p. 

Natural Resources Conservation Service. 2003. 

Plant profile: Cytisus scoparius (L.) Link. 

http://plants.usda.gov/chi_bin/plant_profile.cgi?

symbol=CYSC4. 7 p. 

New England Wild Flower Society. 2003. Invasive 

plant atlas of New England: Cytisus scoparius

(Scotch broom). http://webapps.lib.uconn.edu/ 

ipane/browsing.cfm?descriptionid=48. 6 p. 

Parker, B., G. Miller, L.C. Burrell. 1994. Weeds: 

Scotch broom. PNW 130. Pacific Northwest 

Extension, Oregon State University, Corvallis, 

OR. 10 p. 

Prasad, R. 2003. Scotch broom, Cytisus scoparius

L. in British Columbia. Canadian Forest Service, 

Ottawa, Ontario, Canada. http://www.pfc. 

forestry.ca/biodiversity/broom_e.html. 9 p. 

Syrett, P., S.V. Fowler, E.M. Coombs, J.R. 

Hosking, G.P. Markin, Q.E. Paynter, and A.W. 

Sheppard. 1999. The potential for biological 

control of Scotch broom (Cytisus scoparius) 

(Fabaceae) and related weedy species. 

Biocontrol News and Information 20: 17N-34N. 

University of Connecticut. 2003. University of 

Connecticut plant database: Cytisus scoparius. 

http://www.hort.uconn.edu/plants/c/cytsco/ 

cytsco3.html. 4 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1987. A flora of Utah. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 
278 

Dalbergia ecastaphyllum (L.) Taubert coin vine 

FABACEAE 

Synonyms: Hedysarum ecastaphyllum L. 

 Pterocarpus ecastaphyllum L. 

 Ecastophyllum ecastophyllum (L.) Britton 

 Amerimnon ecastophyllum (L.) Standl. 

John K. Francis 

General Description.—Coin vine is a scrambling 

and climbing shrub that sometimes becomes a 

small tree with stems that extend as much as 10 m 

(Howard 1988). It is also known by the common 

names maraimaray, palo de pollo, mangle médaille, 

popian, and a host of other common names (Burkill 

1995, Liogier 1988). Coin vine develops a tap and 

lateral root system. The tan colored roots support 

many nodules that attach directly to the tap and 

lateral roots. The older stems, which may reach 7 

cm or more in diameter, are gray and extend into 

the crowns of low trees and scramble over low 

obstacles. There are many long, vine-like branches 

that bear leaves only on the current year’s growth. 

The simple leaves have petioles 5 to 12 mm long 

and elliptical or ovate blades 2.5 to 14 cm long and 

2 to 8 cm broad. They are leathery, glossy green, 

rounded at the base, pointed at the tip, and 

pubescent on the under side. The small white 

flowers are grouped in panicles in the leaf axils. 

These develop into small groups of copper-colored 

to gray-brown, elongated lenticular fruits about 1.5 

to 3 cm long that contains one flattened, brown 

seed (Holdridge and Poveda 1975, Howard 1988, 

Liogier 1988, Nelson 1996). 

Range.—Coin vine is native to Florida, the 

Bahamas, the Greater and Lesser Antilles, 

Trinidad, the east coasts of Mexico, Central 

America, and South America to southern Brazil, 

and the coast of tropical West Africa (Liogier 

1988, Burkill 1995). 

Ecology.—Coin vine inhabits coastal sand dunes, 

open coastal forests and brush fields, lagoon sides, 

landward sides of mangrove swamps, shell mounds 

within mangroves, coastal hammocks, and 

estuaries (Burkill 1995, Nelson 1996). It also 

encroaches on dense grass swards. The species is 

most common on sands or sandy soils, but also 

grows on heavier soils. Coin vine inhabits areas in 

Puerto Rico that receive from about 1200 to 2200 

mm of annual precipitation. It also survives 

droughts well (Workman 1980). The species 

frequently grows on sites where there is constant 

salt spray and sometimes in moderately salty soils. 

Old plants tend to form dense thickets. 

Reproduction.—A collection of mature fruits 

from Puerto Rico weighed an average of 0.290 + 

0.010 g/fruit. Seeds extracted from 24 of the pods 

constituted 69 percent of the pod weight. The pods 

are tough and the seeds are fragile. Extraction is 

difficult and unnecessary because seeds germinate 

through the exocarp without difficulty. Seventythree percent of 100 fruits sown in commercial 

potting mix germinated between 31 and 181 days 

after sowing. Germination is hypogeal. Seeds 

disburse by lateral branch extension and by water. 

Seed may be collected after they wash up on the 

shores (Workman 1980), directly from the shrubs, 

and from the ground under them. Natural seedlings 
279

are abundant near coin vine plants. However, few 

survive more than a few weeks. Branches layer 

whenever they come in contact with the ground. 

Growth and Management.—Early height growth 

is moderate but steady. A small group of nursery 

seedlings reached about 15 cm in 5 months. Older 

plants grow about 2 m per year. By resprouting and 

layering, plants (clones) can survive almost 

indefinitely. 

Benefits.—The extensive horizontal branches and 

their roots cover, bind, and stabilize coastal sand 

dunes (Burkill 1995). The stems are brittle and not 

suitable for wooden implements or basketry. The 

crushed roots and bark contain a chemical that was 

used by Native Americans to stupefy and catch fish 

(Workman 1980). Coin vine is listed among the 

honey plants of the Dominican Republic (Marcano 

Fondeur 2001). In Senegal, the leaves are put into 

inhalations and baths to treat various debilities 

(Burkill 1995). Various extracts are used in herbal 

medicine as a diuretic, an emetic, and a vermicide. 

Care must be taken, because some of the tissues are 

toxic (Liogier 1990). 

References 

Burkill, H.M. 1995. The useful plants of West 

Tropical Africa. Vol. 3. Royal Botanci Gardens, 

Kew, United Kingdom. 857 p. 

Holdridge, L.R. and L.J. Poveda A. 1975. Arboles 

de Costa Rica. Vol. 1. Centro Cientifico 

Tropical, San Jose, Costa Rica. 546 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University. Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc. San Juan, Puerto Rico. 566 p. 

Marcano Fondeur, E. de J. 2001. La flora apícola 

de la Republica Dominicana. http://marcano. 

Freeservers.com/nature/estudios/apicola/dicotsp.

html. 11 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Workman, R.W. 1980. Growing native. The 

Sanibal-Captiva Conservation Foundation, Inc., 

Sanibal, FL. 137 p. 
280 

Dasiphora floribunda (Pursh) Kartesz, comb. nov. ined. shrubby cinquefoil

ROSACEAE 

Synonyms: Potentilla fruticosa L. 

 Dasiphora fruticosa auct. non. (L.) Rydh. 

 Potentilla floribunda Pursh. 

 Potentilla fruticosa ssp. floribunda (Pursh) Elkington

 Potentilla fruticosa var. tenuifolia Lehm. 

Juanita A.R. Ladyman 

Illustration source: USDA—Forest Service collection, 

Hunt Institute 

General Description.—Pentaphylloides 

floribunda is commonly known as shrubby or bush 

cinquefoil, but local names also include golden 

hardback, prairie-weed, fivefinger, buckbrush, 

ninebark, widdy, and yellow rose. It is also 

frequently referred to by the most common 

synonym, Potentilla fruticosa (Kartesz 1994). This 

is a distinctive shrub and easily separable from the 

other Potentilla species that are all herbaceous 

(Cronquist and others 1997). Recently it has been 

reported as Dasiphora floribunda (Pursh) Kartesz 

comb. nov. ined (ITIS 2002). The genus 

Dasiphora includes approximately 8 or 9 species 

that are all exclusively in Asia except for shrubby 

cinquefoil. The genus name “Pentaphylloides” is 

derived from the Greek and refers to the pinnate 

five-foliate leaves. “Penta-” means “five-“and 

“phylloides” means “leaved.” “Fruticosa” is a 

Latin adjective that means “shrubby” or “bushy.” 

This shrub is usually from 10 cm to 1 m tall but 

may grow to a little over 1.5 m. The herbage of the 

current year is covered by soft hairs, but the stem 

becomes smooth on aging with, eventually, a redbrown shreddy bark. The leaves comprise usually 

five, but sometimes three to seven, elliptic-shaped 

leaflets. The flowers are solitary in the leaf axils 

along the stem but form three- to seven-flowered 

clusters (cymes) at the stem tips. Petal color is 

usually yellow, although it may be white, or even 

pink and red, in cultivated varieties, and there are 

15 to 25 stamens per flower (Carter 1997, 

Cronquist and others 1997, Hitchcock and 

Cronquist 1973, Vines 1986). This is a wideranging species, but apparently chromosome 

number differs between new and old world 

material. North American plants have a 

chromosome number of 2n = 14 while Asian and 

European plants have 2n = 28 (Cronquist and 

others 1997). 

Range.—Shrubby cinquefoil is a common, wideranging circumboreal shrub ranging from 

Greenland and Labrador, to Alaska and south to 

the mountains of California, Nevada, Arizona, 

Colorado, New Mexico, and through the northern 

states into Iowa, Indiana, Illinois, Ohio, 

Pennsylvania and New Jersey in North America. It 

also occurs in Europe, for example England, 

Ireland, Scandinavia and Bulgaria, and in Asia 

(Stace 1997, Gibbons and Davies 1994, Polunin 

1997). 
281

Ecology.—Shrubby cinquefoil grows in moist 

soils in mountain regions from lower foothills to 

sub alpine. The elevation tends to depend 

somewhat upon latitude, but generally plants are 

found between approximately 1,800 m and 3,400 

m. It grows in open woods, wet mountain 

meadows, calcareous bogs, limestone pavements, 

and along streams (Cronquist and other 1997, 

Gibbons and Davies 1994, Vines 1986, Welsh and 

others 1993). It has also been reported from 

relatively xeric habitats such as mountain shrub 

lands and rocky mountain slopes (McGregor and 

others 1986, Stace 1997, Thilenius 1972). The 

degree of permanent soil moisture required is 

likely to depend on average precipitation, 

temperature, and localized water collection sites or 

drainage channels. It is a frequent member of 

willow (Salix)-dominated communities, and in 

some parts of its range, for example New Mexico, 

it is a member of the Krummholz communities 

found at the lower tundra edge (Dick-Peddie 

1993). Shrubby cinquefoil may be the dominant 

shrub in many situations and can also grow in 

fairly pure stands. For example, it may be the 

dominant shrub in aspen stands or other forest 

clearings where snow accumulation prevents tree 

establishment (Knight 1994, Thilenius 1972). In 

addition, it frequently attains dominance in 

limestone-derived soils (Thilenius 1972). 

Reproduction.—Shrubby cinquefoil generally 

flowers in June through August, although flowers 

may be found until the first frost (Epple 1995, 

McGregor and others 1986, USDA FS 1988). The 

achenes mature in summer and fall. The bright 

yellow flowers are insect pollinated. The fruit is a 

densely hairy achene. 

Growth and Management.—Shrubby cinquefoil 

increases in response to overgrazing and other 

disturbance. In the Western United States it does 

not appear to be an invasive species, but in New 

England it can be an aggressive invader of 

agricultural land and may be difficult to eradicate 

once established (USDA FS 1988). It has been 

used as an indicator species of past disturbance 

(Dick-Peddie 1993). Shrubby cinquefoil is widely 

used as an ornamental and in wild land seeding. It 

does best in full sun as it tends to flower poorly in 

shade conditions. Seed apparently does not require 

cold stratification for germination and will 

germinate immediately after harvesting (Bonde 

1965, Tykač 1990). The highest germination rate 

was achieved at 18 o

C (Bonde 1965). However, it 

has been proposed that cold stratification may 

reduce the temperature requirements for seed 

germination (Baskin and Baskin 2001). A cold, 

moist stratification period of 2 months has been 

generally recommended for Potentilla species, 

including Potentilla fructicosa. (Phillips 1987). 

Shrubby cinquefoil can also be propagated from 

softwood cuttings in the spring or from suckers 

(Tykač 1990). Container grown plants, that are 

very popular within the landscape business, can be 

planted in spring and fall. Even old plants are 

reported to tolerate transplanting (Tykač 1990). 

Benefits.—Shrubby cinquefoil leaves are bitter 

tasting, and the forage and browse value varies 

depending upon the part of the world in which it 

grows. It is generally on summer range, although at 

lower elevations it may be valuable for fall 

browsing (Dayton 1931). It is likely that just as the 

shrubs’ morphology varies, the levels of secondary 

plant products that impart the bitter taste differ 

between habitats and geographic range. Generally, 

it is considered inferior forage for livestock, but it 

is used by cattle in Arizona and Montana (Epple 

1995, Vines 1986). It is also often an important 

browse for sheep and goats especially in the 

Southwestern United States and in southwestern 

Montana and contiguous portions of southeastern 

Idaho (Dayton 1931). In Montana it is rated 18th

among the most important browse species. 

Similarly, it is reported unpalatable to deer 

(Thilenius 1972) but does provide browse for deer 

and elk in some regions (Epple 1995, Vines 1986). 

On some overgrazed range it is often grazed quite 

closely, and the shrubs assume a hedge-like 

appearance (USDA FS 1988). Such use indicates 

overstocking and a critical reduction, if not 

elimination, of other palatable plant species. It 

provides effective protection against soil erosion 

(Epple 1995, Vines 1986). Native Americans have 

used various parts of the plant for an assortment of 

purposes including as a medicine and in 

ceremonials (Moerman 1998). The Blackfoot 

peoples have used the leaves to fill pillows and the 

bark as tinder, while the Eskimos of Alaska and the 

Arctic have used dried leaves for tea (Moerman 

1998). Leaves were also used for tea in Russia. The 

name “potentilla” is derived from the Latin 

“potens,” which means powerful and may refer to 

the medicinal value of some species. A preparation 

made from the leaves was used by the Cheyenne 

for protection and was also considered a deadly 

arrow poison, only to be administered by holy 

people (Carter 1997, Moerman 1998). Shrubby 

cinquefoil has commercial value in the landscape 

and horticultural trade. It is a popular ornamental 

shrub, and many cultivars exist that have been 

chosen for the degree of leaf pubescence 
282 

(hairiness), flower color, and growth form. A list of 

some of the recognized cultivars available can be 

found in Bailey and others (1976) and Tykač

(1990).

References. 

Bailey, L.H., E.Z. Bailey, and the staff of the 

Liberty Hyde Bailey Hortorium. 1976. Hortus 

Third – a concise dictionary of plants cultivated 

in the United States and Canada. Macmillian 

Publishing Company, New York. 1,290 p. 

Baskin, C.C. and J.M. Baskin. 2001. Seeds, 

ecology, biogeography, and evolution of 

dormancy and germination. Academic Press, 

New York. 666 p. 

Bonde, E.K. 1965. Further studies on the 

germination of seeds of Colorado alpine plants. 

University of Colorado Studies 18. 

Carter, J.L. 1997. Trees and shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Cronquist, A., N.H. Holmgren, and P.K. 

Holmgren. 1997. Intermountain flora, vascular 

plants of the Intermountain West, U.S.A. Vol. 3, 

Part A. New York Botanical Garden, Bronx, 

NY. 446 p. 

Dayton, W.A. 1931. Important western browse 

plants. Miscellaneous Publication 101. U.S. 

Department of Agriculture, Washington DC. 

214 p 

Dick-Peddie, W.A. 1993. New Mexico vegetation 

– past, present, and future. University of New 

Mexico Press, Albuquerque, NM. 244 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. Falcon Press Publishing Co., Helena, 

MT. 347 p. 

Gibbons, B. and P. Davies. 1994. Field guide to 

wild flowers of Britain and Northern Europe. 

The Crowood Press, Marlborough, Wiltshire, 

UK. 320 p. 

Hitchcock, C.L. and A. Cronquist. 1973. Flora of 

the Pacific Northwest – an illustrated manual. 

University of Washington Press, Seattle, WA. 

730 p. 

ITIS, the Integrated Taxonomic Information 

System. 1996. Dasiphora floribunda (Pursh) 

Kartesz comb. nov. ined. Taxonomic Serial No.: 

565123 and Plants, database (version 5.1.1) 

http://www.itis.usda.gov/ [not paged]. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada and 

Greenland. Vol. 1 - Checklist. 2nd Ed. Timber 

Press, Portland, OR. 622 p. 

Knight, D.H. 1994. Mountains and plains–the 

ecology of Wyoming landscapes. Yale 

University Press, New Haven, CN. 338 p. 

McGregor, R.L., T.M. Barkley, R.E. Brooks, E.K. 

Schofield. 1986. Flora of the Great Plains. 

University Press of Kansas, Lawrence, KN. 

1,402p. 

Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. 927 p. 

Phillips, J. 1987. Southwestern landscaping with 

native plants. University of New Mexico Press, 

Albuquerque, NM. 140p. 

Polunin, O. 1997. Flowers of Greece and the 

Balkans–a field guide. Oxford University Press, 

New York. 567p. 

Stace, C. 1997. New Flora of the British Isles. 2nd

edition. Cambridge University Press, New York. 

1,130 p. 

Thilenius, J.F. 1972. Classification of deer habitat 

in the ponderosa pine forest of the Black Hills, 

South Dakota. Research Paper RM-91. USDA 

Forest Service Rocky Mountain Forest and 

Range Experiment Station, Fort Collins, CO. 28 

p. 

United States Department of Agriculture Forest 

Service. 1988. Range Plant Handbook. Dover 

Publications, Inc. New York. 838 p. 

Vines, R.A. 1986. Trees, shrubs, and woody vines 

of the Southwest. University of Texas Press. 

Austin, TX. 1,104 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, L.C. 

Higgins, eds. 1993. A Utah flora. Second 

Edition, revised. Brigham Young University, 

Provo, UT. 986 p. 
283

Dasylirion wheeleri S. Wats. Wheeler sotol

AGAVACEAE 

Synonyms: None. 

Juanita A. R. Ladyman 

Illustration credit: Robert De Witt Ivey 

General Description.—The common names for 

Dasylirion wheeleri are Wheeler sotol, sotol, 

desert spoon, spoonplant and spoonleaf. The latter 

names refer to the enlarged, spoon-like bases of 

the long serrated leaves. Dasylirion wheeleri

grows from a thick, woody, mostly subterranean 

caudex with a large rosette of rigid leaves that are 

armed with sharp curved spines, or teeth, around 

the margins (Kearney and others 1960). The long 

flowering stalk arises out of the center of the 

rosette and is topped by a slender compound 

panicle. The genus name, Dasylirion, means thick, 

or hairy, white lily from “dasy-“ and “lirio” 

(Gledhill 1992). Dasylirion wheeleri is 

distinguished from Dasylirion leiophyllum

Engelm. ex Trel. by having the leaf margin spines 

directed apically rather than basally (Carter 1997). 

There are two recognized varieties of Dasylirion 

wheeleri, var. wheeleri and var. wislizenii (Vines 

1960). 

Range.—Wheeler sotol grows on well-drained 

rocky and gravelly hillsides or slopes in Western 

Texas, southern to central New Mexico, southern 

Arizona in the U.S.A. and in Chihuahua in 

northern Mexico (Kearney and others 1960, 

Powell 1998, Tesky 1993, Vines 1960). 

Ecology.—Wheeler sotol occurs in chaparral, 

southwestern oak (Quercus) woodland, desert and 

semi-desert grasslands communities at 914 to 

1,830 m in elevation (Carter 1997). It occurs in 

seral, climax, and postclimax communities. In the 

desert plains grasslands it is often sub-dominate in 

the beargrass-scrub oak postclimax community 

and the curly mesquite (Hilaria belangeri)-blue 

grama climax community. It has been described 

as a diagnostic climax member of desert 

grasslands (Dick-Peddie 1993). In Trans-Pecos, 

Texas, it is commonly found growing on limestone 

and granite (Powell 1998, Schmutz and others 

1992). Wheeler sotol usually grows on rocky soils 

in desert grasslands that are often described as too 

moist for open desert scrub but too dry for oak or 

juniper woodland (Burgess 1995, Dick-Peddie 

1993, Schmutz and others 1992). These are areas 

where subsurface water is generally not available. 

Wheeler sotol can store relatively large amounts of 

water internally that act as a buffer against the 

rapid onset of drought stress. It is classed as a 

water storer, or “succulent” plant (Burgess 1995, 

Thomas 1991). The species can also tolerate 

several degrees of frost (Vines 1960). 

Reproduction.—The flowering stalks elongate to 

up to 5 m in late spring and bloom from May to 

August (Bowers 1993). Wheeler sotol is dioecious, 

the male and female flowers being borne on 

separate plants. Female flowers frequently flower 

later than the male plants (Warnock 1974). The 

female flowers are wind pollinated, and wind also 

disperses the seed that are contained within a 

papery, one-celled, three-winged capsule. Wheeler 

sotol grows readily from seed. 

Growth and Management.—Wheeler sotol 

occurs in desert grasslands that are being 

increasingly managed with fire (Thomas and 

Goodson 1992). In these areas, fire is primarily 

used to stimulate grass production and reverse 

dense scrub invasion. The apical meristem of sotol 

is usually protected from fire by a cluster of green 

leaves, and this species can also resprout from the 
284 

caudex that is the region at the base of the stem 

(McPherson 1995, Thomas 1991). However, 

frequent fires and stable soil moisture regimes tend 

to exclude wheeler sotol (Burgess 1995). The 

response of sotol to fire tends to vary depending 

upon conditions. Kittams (1973) observed that 

mature sotol (in this case Dasylirion leiophyllum) 

were usually killed by fire. White (1969) randomly 

selected 50 individuals after a fire in desert 

grassland in Arizona and classified them with 

respect to the level of the damage (severe, 

moderate, or light) experienced. All of the lightly 

or moderately damaged plants survived, but only 3 

percent of the severely damaged plants survived. 

Sprouting did not occur except as refoliation from 

the terminal bud (White 1969). Ahlstrand (1982) 

investigated the response of Dasylirion 

leiophyllum to fire at sites in New Mexico and 

western Texas. He reported losses of 50 percent or 

more on burned sites, but surviving top-killed 

plants regained much of their pre-burn coverage 

after three growing seasons. This was similar to 

the observations after a semidesert grassland fire 

in southern Arizona where 47 percent of the 

Wheeler sotol plants burned to death (Thomas and 

Goodson 1992). Of the 71 Wheeler sotol plants 

that survived, all had apical regrowth, but despite 

the fact that sotol may occasionally produce new 

shoots from below ground, no offsets or seedlings 

were found 11 months after the fire (Thomas and 

Goodson 1992). In this study no individuals 

evaded the fire in refugia. 

Benefits.—The head-like trunks and leaf bases of 

Wheeler sotol are edible and American Indians 

roasted and ate the young flower stalks (Moerman 

1998, Powell 1998). The basal trunks contain 

starch and are also roasted, fermented and distilled 

to make alcohol (Powell 1998). Currently an 

alcoholic beverage, generally known as sotol, is 

made in parts of Mexico (Bowers 1993). Bighorn 

sheep and javelinas browse sotols (Epple 1995, 

Kearney and others 1960, Starr 1999). In times of 

stress Wheeler sotol has been used as feed for 

livestock in a variety of ways. The trunks and leaf 

bases have been used during drought, and the 

plants have also been machine-cut and shredded 

for emergency feed (Dayton 1931, Powell 1998). 

In addition, ranchers have burned off the leaves 

and split the heads (trunks) for livestock (Vines 

1960). The leaves have been used to make mats, 

baskets, thatch, and paper (Moerman 1998, Powell 

1998). The long flowering stalks have been used 

extensively within its range for corral, porch and 

house roofs, and other structures (Powell 1998). 

They have also been used for ceremonial purposes 

by Native Americans (Moerman 1998). Wheeler 

sotol plants are commercially used as ornamentals 

and described as “one of the most striking accent 

plants available” (Starr 1999). The leaves were 

once sold for use in floral arrangements, but this 

practice has been discouraged due to conservation 

considerations (Epple 1995). 

References 

Ahlstrand, G.M. 1982. Response of Chihuahuan 

Desert Mountain Shrub Vegetation to Burning. 

Journal of Range Management 35 (1): 62-65. 

Bowers, J.E. 1993. Shrubs and trees of the 

southwest deserts. Southwest Parks and 

Monumens Association, Tucson, AZ. 140 p. 

Burgess, T.L. 1995. The dilemma of coexisting 

growth forms. In: The desert grassland. M.P. 

McClaran and T.R. Van Devender, eds. The 

University of Arizona Press, Tucson, AZ. p. 31-

67. 

Carter, J.L. 1997. Trees and shrubs of New 

Mexico. Johnson Books, Boulder, CO. 533 p. 

Dayton, W. A. 1931. Important western browse 

plants. Miscellaneous Publication 101. U.S. 

Department of Agriculture, Washington, DC. 

214 p. 

Dick-Peddie, W.A. 1993. New Mexico vegetation: 

past, present, and future. University of New 

Mexico press, Albuquerque NM. 244 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. Falcon Press Publishing Co., Helena, 

Montana. 347 p. 

Gledhill, D. 1992. The names of plants. 2nd ed. 

Cambridge University Press, Cambridge, 

England. 202 p. 

Ivey, R.D. 1995. Flowering plants of New Mexico. 

3rd ed. Rio Rancho Printing, Albuquerque, NM. 

504 p. 

Kearney, T.H., R.H. Peebles, J.T. Howell, and E. 

McClintock. 1960. Arizona flora. 2d ed. 

University of California Press. Berkeley, CA. 

1,085 p. 

Kittams, W.H. 1972. Effect of fire on vegetation of 

the Chihuahuan Desert region. Proc. Tall 

Timbers Fire Ecology Conference 12: 427-444 
285

McPherson, G.R. 1995. The role of fire in desert 

grasslands. In: The desert grassland. M.P. 

McClaran and T.R. Van Devender, eds. The 

University of Arizona Press, Tucson, AZ. p. 130-

151. 

Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. 927 p. 

Powell, A. M. 1998. Trees & shrubs of TransPecos and adjacent areas. University of Texas 

University Press, Austin, TX . 498 p. 

Schmutz, E.M., E.L. Smith, P.R. Ogden, M.L. 

Cox, J.O. Klemmedson, J.J. Norris, and L.C. 

Fierro. 1992. Desert grassland. In: R.T. 

Coupland, ed. Natural grasslands: Introduction 

and Western Hemisphere. Elsevier, New York. 

p. 337-362. 

Starr, G. 1999. Dasylirion – The shaggy lilies. 

Desert Plants 15(1): 29-31. 

Tesky, J.L. 1993. Dasylirion wheeleri. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire 

Sciences Laboratory. Fire Effects Information 

System. http://www.fs.fed.us/database/feis/. 8 

p. 

Thomas, P.A. 1991. Response of succulents to 

fire: A review. International Journal of Wildland 

Fire 1(1): 11-22. 

Thomas, P.A. and P. Goodson. 1992. Conservation 

of succulents in desert grasslands managed by 

fire. Biological Conservation. 60(2): 91-100. 

Vines, R.A. 1960. Trees, shrubs, and woody vines 

of the Southwest. Sixth printing 1986. 

University of Texas Press. Austin, TX. 

 1,104 p. 

Warnock, B.H. 1974. Wildflowers of the 

Guadalupe Mountains and the Sand Dune 

Country, Texas. Sul Ross State University, 

Alpine, TX. 176 p. 

White, L.D. 1969. Effects of a wildfire on several 

desert grassland shrub species. Journal of Range 

Management 22: 284-285. 
286 

Datura inoxia P. Mill. devil’s trumpet 

SOLANACEAE 

Synonyms: Datura metel auct. non L. 

Datura fastuosa sensu Griseb. non L. 

John K. Francis 

Drawing source: Missouri Botanic Garden 2003 

General Description.—Devil’s trumpet is also 

known as angel’s trumpet, thorn apple, Indian 

apple, purple datura, garden datura, horn-of-plenty, 

David bush, chaico blanco, buenas tardes, 

concombre zombi, cornicopio, trompette du 

jugement, and pomme épineuse. It is a shrub or 

woody herb to 2 m in height that is often grown as 

an annual in temperate zones. The stems are 

semiwoody and suffruticose in the moist tropics. In 

drier environments, it dies to the ground annually. 

The alternate leaves have petioles 3 to 7 cm long 

and ovate to elliptic blades 6 to 15 cm long with 

sinuate to irregularly toothed edges. The tubular 

flowers are axillary and usually solitary. They are 

erect or nodding, have a five-toothed calyx 5 to 7 

cm long and a white, purple, or yellow corolla 8 to 

20 cm long, often double in horticultural varieties. 

The ovoid capsule is nodding, about 3 cm in 

diameter and covered with stout, soft prickles 2 to 

4 mm long. The capsules remain on the plant for a 

long period. The yellowish-brown seeds are flat, 

kidney-shaped, about 5 mm long, and have a small 

fleshy aril. Datura species normally have 12 pairs 

of chromosomes (Bonde 2001, Burkill 2000, 

Howard 1989, Liogier 1995, Stevens and others 

2001). 

Range.—The native range of devil’s trumpet 

appears to have been Mexico and the U.S. 

Southwest (Schultes and Hofmann 1992), the 

Caribbean Islands (Howard 1989), India, and 

China (Schultes and Hofmann 1992). There are 

related species in all these areas and ancient 

traditions of its use (Bonde 2001, Schultes and 

Hofmann 1992). The species apparently 

naturalized in Africa hundreds of years ago 

(Burkill 2000). Today, devil’s trumpet grows 

throughout the Tropics. 

Ecology.—Devil’s trumpet grows naturally in 

disturbed areas such as eroded sites, old fields, 

vacant lots, overgrazed pastures and rangeland, 

roadsides and abandoned roadbeds, and fencerows. 

Apparently, disturbance and reduced competition 

are required for the plant to become established 

and grow. A wide variety of well-drained soils on 

both igneous and sedimentary parent materials are 

suitable. In Puerto Rico, the species grows 

naturally in areas that receive from 750 to about 

1000 mm of mean annual precipitation from near 

sea level to about 400 m. Devil’s trumpet grows on 

sites up to 900 m in elevation in Nicaragua 

(Stevens and others 2001) 

Reproduction.—Devil’s trumpet flowers and 

fruits throughout the year in some environments, 

but in India principally from July to September 

(Parrotta 2001), and from September through 

November in Nicaragua (Stevens and others 2001). 

Hummingbirds sometimes visit the flowers, but are 

affected by the alkaloids in the nectar and must 

limit their consumption. Honeybees are apparently 

unaffected. The flowers have an intense night 
287

fragrance (Annie’s Annuals 2002), which perhaps 

helps attract night-flying moths. The seeds are 

distributed by ants and some species of birds that 

are resistant to the chemicals they contain (Bonde 

2001). Seeds germinate in less than 2 weeks at soil 

temperatures of 27 °C (Hardy Plants 2002). Devil’s 

trumpet is normally propagated with seeds. 

Growth and Management.—In dry environments, 

devil’s trumpet reaches a maximum height of about 

0.5 m in one growing season. In moist areas, it can 

reach two to four times this height although it is 

not clear whether it does so in a single season. New 

stands can be established by cultivation and 

planting. Although control information has not 

been published, weedy plants can probably be 

reduced or eliminated without difficulty by 

cultivation or spraying with broadleaf herbicides. 

Benefits and Detriments.—Devil’s trumpet is 

grown in all but the coldest climates as a flowering 

ornamental. Beautiful white, purple, and yellow 

varieties with large, single and double blossoms are 

available. From ancient times continuing to the 

present, the taking of Datura spp. tissues, 

particularly the seeds, was used in shamanistic 

rituals as a path to enlightenment (Schultes and 

Hofmann 1992). Today, people frequently 

experiment with it for the hallucinogenic effect, but 

the results are so unpleasant (dark visions, 

disorientation, amnesia, blurred vision, dry mouth, 

and incontinence) that they seldom recommend the 

experience (Erowid 2002). Overdoses can result in 

death. The plant has been used to treat impotence, 

asthma, diarrhea, as an analgesic, to control fever, 

kill parasites, and as a drug for criminal purposes 

(Bonde 2001, Parrotta 2001). Devil’s trumpet 

contains a host of phytoactive chemicals including 

atropine, hyoscyamine, hyoscine, scopolamine, 

norscopolamine, meteloidine, hydroxy-6-

hyoscyamine, tiglic esters of dihydroxytropine, and 

a number of withanolides (Gupta and others 1991, 

Janthangvith and others 2000). Alkaloid 

concentration in tissues is around 0.5 percent 

(Niber and others 1992) but varies according to 

environmental circumstances (Burkill 2000). 

Devil’s trumpet is a minor weed in tropical and 

some subtropical countries. It causes erratic 

behavior and even death of livestock that have 

eaten it, but it is seldom a problem for pastured 

animals because they carefully avoid consuming it. 

References 

Annie’s Annuals. 2002. Datura metel, ‘Belle 

Blanche.’ http://www.anniesannuals.com/signs/ 

D%20-%20G/Datura_metel_’Belle_Blanche’. 

htm. 1 p. 

Bonde, K. 2001. The genus Datura: From research 

subject to powerful hallucinogen. 

http://leda.lycaeum.org/Documents/The_Genus_

Datura:_From_Research_Subject_to_Power. 

9 p. 

Burkill, H.M. 2000. The useful plants of West 

Tropical Africa. Vol. 5. 2nd ed. Royal Botanical 

Gardens, Kew, UK. 686 p. 

Erowid. 2002. Erowid experience vaults. 

http://www.erowid.org/experiences/subs/ex_Dat

ura.shtml#retrospective_/_Summary. [not 

paged]. 

Gupta, M., A. Bagchi, and A.B. Ray. 1991. 

Additional withanolides of Datura metel. 

Journal of Natural Products. 54(2): 599-602. 

Hardy Plants. 2002. Specialty perennials. 

http://hardyplants.com/D.htm. [not paged]. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Janthangvith, J., P. Chumsri, K. Kraisintu, and A. 

Pongpan. 2000. Comparison of tropane alkaloid 

production by Datura innoxia and Datura metel

varieties of white, soil, hydroponic, in vitro plant 

culture and tranformed root culture. Mahidol 

University annual Research Abstracts. 

http://www.hahidol.ac.th/abstracts/annual1999/0

355.htm. 2 p. 

 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Missouri Botanic Garden. 2003. Missouri Botanic 

Garden. Missouri Botanic Garden, St. Louis, 

MO. http://www.mobot.org/. [not paged]. 

Niber, B.T., J. Helenius, and A.L. Varis. 1992. 

Toxicity of plant extracts to three storage beetles 

(Coleoptera). Journal of Applied Entomology 

113(2): 202-208. 

Parrotta, J.A. 2001. Healing plants of India. CABI 

Publishing, Wallingford, UK and New York. 

917 p. 
288 

Schultes, R.E. and A. Hofmann. 1992. Plants of 

the gods. Healing Plants Press, Rochester, NY. 

192 p. 

Stevens, R.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanic Garden, St. Louis, MO. pp. 

911-2,666. 
289

Desmanthus virgatus (L.) Willd. wild-tantan 

FABACEAE 

Synonyms: Mimosa virgata L. 

 Acuan virgatum (L.) Medikus 

 Acuan guadeloupense Britton & Rose 

 Desmanthus depressus Humb. & Willd. 

John K. Francis 

General Description.—Wild-tantan, also called 

prostrate bundleflower, dwarf koa, desmanthus, 

desmanto, acacia courant, acacia savane, and 

pompon blank, is a suffruticose shrub or perennial 

woody herb that reaches 0.5 to 2 m in height and 3 

to 10 mm in basal diameter. The plant is supported 

by a deep woody taproot and descending laterals. 

Each dry season, wild-tantan dies back to the root 

crown or lower stems and grows anew when the 

rains return. The stems and branches are semiwoody to woody, pithy in the center, angular, 

slender, green turning brown, and erect or 

semiprostrate. There may be as many as 50 stems, 

depending on the age, vigor, and disturbance 

history of the plant. The 2- to 8-cm leaves have a 

short petiole, less than seven pinnae, 10 to 25 pairs 

of leaflets per pinnae that are linear-oblong and 4 

to 9 mm long. The flowers occur in axillary heads 

with peduncles 2 to 7.5 cm long. The petals are 

white or creamy yellow. The pod, a legume, is 

linear, 50 to 90 mm long, 3 to 4 mm broad, 

reddish-brown, and contains 20 to 30 seeds 

(Howard 1988, Liogier 1988, Skerman and others 

1988). There are 2n = 28 chromosomes (Long and 

Lakela 1976). 

Range.—Wild-tantan is native to the Greater and 

Lesser Antilles, Trinidad and Tobago, from Texas 

to Argentina, and the Galapagos Islands (Liogier 

1988, Howard 1988, Skerman and others 1988). It 

has naturalized in Florida and Hawaii (Institute of 

Systematic Botany 2002, Neal 1965). The species 

has been planted throughout the tropics and 

subtropics and has probably naturalized in many 

other areas. 

Ecology.—Wild-tantan grows well in deep, wellto moderately well-drained silts, clays, loams, and 

sands with pH’s from about 5 to about 8, formed 

over both igneous and sedimentary rocks. It is 

seldom seen in shallow, rocky sites. The species is 

recommended for planting in Australia in areas 

receiving from 550 to 750 mm of mean annual 

precipitation (Department of Primary Industries 

2002) and grows naturally in Puerto Rico in areas 

that receive from 750 to about 1600 mm of mean 

annual precipitation. Wild tantan tolerates seasonal 

drought without difficulty. The species grows from 

near sea level up to 300 m in elevation (Skerman 

and others 1988). Wild-tantan is intolerant of 

shade. It tolerates severe competition from grass 

and herbs but disappears when overtopped by trees. 

The species is usually more or less evenly 

dispersed in stands of grass and herbs, often at 1 or 

2 per square meter. Although plants are killed to 

the ground by heavy frost, they regrow from the 

root crown (Forlin and others 2000). It withstands 

heavy grazing pressure and recovers well from fire. 
290 

Reproduction.—Wild-tantan blooms at the end of 

the wet season, and fruits mature about 1 month 

later. Seeds collected in Puerto Rico weighed an 

average of 0.0041 + 0.0005 g/seed or 244,000 

seeds/kg. Placed to germinate on moist filter paper, 

only 28 percent germinated in 1 year. The 

remaining seeds were scarified (by nicking) and 

100 percent germinated starting in 2 days (author’s 

observation). In nature, scarification occurs by 

weathering of the seed coat in the soil or while 

passing through an herbivore. Artificial 

scarification by treatment with concentrated 

sulfuric acid for 8 minutes is recommended 

(Skerman and others 1988). The seeds are 

dispersed by ruminant animals. Many seedlings 

spring up under favorable conditions in natural 

stand, but few survive. The species can be 

propagated by rooting hormone-treated leaves 

(Forlin and others 2000). 

Growth and Management.—Wild-tantan 

normally grows from 0.5 to 1.5 m/year in height. 

After establishment, the root crown gradually 

increases in size until it reaches 15 cm across in 3 

years. Plantations are established by sowing 2 kg of 

scarified seed per hectare into well-prepared 

seedbeds at a depth of 1 to 1.5 cm. Plantations can 

be cut for feed 4 times/year and yield 35 

tons/ha/year of dry matter. There is little mortality 

up to at least the fourth year (Skerman and others 

1988). 

Benefits.—Wild-tantan is planted throughout the 

tropics as a forage and hay crop and is appreciated 

as a forage in range land. The forage is nontoxic 

and palatable to cattle and other ruminants. It is a 

favorite of white-tailed deer in southern Texas 

(Soltero-Gardea and others 1994). Crude protein 

content of whole plants from plantations ranges 

from 10.5 to 15.5 percent, the leaves having 22.4 

percent and the stems 7.1 percent (Skerman and 

others 1988). Wild-tantan is planted as a hedgerow plant in the alley cropping system on steep 

land in the Philippines (Labios and others 1994). It 

has been shown to be an efficient nursery-stage 

host for sandalwood (Santalum album L.) (Fox and 

others 1996). The species can become a serious 

weed in sugar cane fields (Neal 1965). 

References 

Department of Primary Industries. 2002. Legumes 

for the tropics and subtropics: Desmanthus 

(Desmanthus virgatus). Queensland 

Government, Brisbane, Australia. http://dpi.qld. 

gov.au/pastures/4490.html. 2 p. 

Forlin, S.M., H.Y. Rey, and L.A. Mroginski. 2000. 

Cultivo de tejidos de Desmanthus virgatus: 

atención de plantas a partir de hojas. 

Universidad Nacional del Nordeste, Corrientes, 

Argentina. Comunicaciones Científicas y 

Tecnológicas 2000. 3 p. 

Fox, J.E.D., A.I. Doronila, D.R. Barrett, and I.K. 

Surata. 1996. Desmanthus virgatus (L.) Willd. 

An efficient intermediate host for the parasitic 

species Santalum album L. in Timor, Indonesia. 

Journal of Sustainable Forestry 3(4): 13-23. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Institute of Systematic Botany. 2002. Atlas of 

Florida plants: Desmanthus virgatus. University 

of Florida, Gainesville, FL. http://www. 

 plantatlas.usf.edu/maps.asp?plantID=1474. 1 p. 

Labios, R.V., J.G. Montesur, and R.O. Retales. 

1994. Alley cropping in sloping upland rice 

areas of Cavite, Philippines. Philippine Journal 

of Crop Science 19(1): 33-37. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Río Piedras, PR: Editorial de la Universidad 

de Puerto Rico. 481 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Museum 

Press, Honolulu, HI. 924 p. 

Skerman, P.J., D.G. Cameron, and F. Riveros. 

1988. Tropical forage legumes. 2nd. Ed. Food 

and Agriculture Organization of the United 

Nations, Rome. 692 p. 

Soltero-Gardea, S., I.M. Ortega, and F.C. Bryant. 

1994. Nutrient content of important deer forage 

plants in the Texas costal bend. Texas Journal of 

Science 46(2): 133-142. 
291

Distictis lactiflora (Vahl) DC. pega palo 

BIGNONIACEAE 

Synonyms: Bignonia lactiflora Vahl 

 Bignonia rigescens Jacq. 

 Bignonia odorata Bello 

 Distictis rigescens (Jacq.) DC. 

 Macrodiscus lactiflorus (Vahl) Bureau ex Baill. 

John K. Francis 

General Description.—Pega palo, also known as 

liana fragante and viuda, is a climbing woody vine 

that clings to objects by means of sticky disks and 

may extend 6 m or more into the crowns of trees or 

laterally along rocks and fences (AcevedoRodríguez 1985). Older plants may have multiple 

stems arising from ground level. Like many other 

vines in the Bignoniaceae, pega palo has a 

characteristic pattern in the stem cross-section, 

shamrock-shaped in this case. The brown stems are 

slender, furrowed, and fissured. Older plants have 

no taproot, only lateral roots with sinkers and a 

moderate amount of fine roots. The larger roots 

have the same general appearance as the stems. 

There are few branches until the crown of the plant 

is reached. Branches usually arise in three’s at the 

nodes (author’s observation). The opposite, 

compound leaves have two leaflets with the tendril 

in place of a third leaflet. The leaf blades are ovate, 

3 to 8 cm long, and obtuse, rounded, or acute at the 

tip. Terminal panicles support a few white tubular 

flowers with a yellow throat, 3 to 7 cm long. A few 

of the flowers develop into narrowly ellipsoid, 

somewhat flattened capsules 7 to 12 cm long and 

brown at maturity. They split open by a medial 

suture and release brown, winged seeds (Liogier 

1995, Acevedo-Rodríguez 1985). Pega palo occurs 

in the same areas and is similar in appearance to 

Macfadyena unguis-cati (L.) A. Gentry. They are 

distinguished by the tripartite tendril, yellow 

flower, and long pod of M. unguis-cati. 

Range.—Pega palo is native to Cuba, Hispaniola, 

Puerto Rico, and the Virgin Islands. It has been 

introduced into Guadeloupe (Liogier 1995). 

Ecology.—Pega palo grows in areas that receive 

from about 750 to 2000 mm of annual 

precipitation. It grows on most types of welldrained to excessively drained soils derived from 

sedimentary (including limestone), igneous, and 

metamorphic (including ultramafic) rocks. Pega 

palo grows from near sea level to over 600 m in 

elevation. The species is moderately intolerant, 

growing in low basal-area forests, forest edges and 

openings, brushy pastures, fencerows, and 

disturbed forests. Pega palo is not eaten by cattle 

and is often common in overgrazed range. 

Although the usual habit of the species is to ascend 

into trees or over rocks, it forms mounds and 

scrambles over the ground when trees are absent. 

Pega palo resprouts after fires. An unknown insect 

in Puerto Rico often destroys all the seeds in 

immature capsules. 

Reproduction.—Pega palo flowers from 

December to July (Acevedo-Rodríguez 1985). 

Because the species requires nearly full sunlight to 

bloom and fruit and because it does not climb to 

the tops of most of the trees it invades, fruiting 

plants are not abundant. A collection of fruits from 

Puerto Rico contained seeds of which only about 

one in four contained endosperm. Visually culled, 

filled seed weighed an average of 0.0153 g or 

65,000 seeds/kg. Sown without any pretreatment 

on peat, 55 percent germinated within 44 days. 

Germination is hypogenous. Seedlings, whose first 

two leaves have an unusual dark-green color tinged 

with purple and a whitish net-like vein pattern, are 

common, but few survive the first year (author’s 
292 

observation). Vines layer (root) when they come in 

contact with the ground. 

Growth and Management.—Pega palo grows at 

least a meter per year from sprouts. Individual 

vines live several years, and strong plants develop 

new stems periodically. Grazing of dry and moist 

forest areas probably promotes the species. 

Benefits.—Pega palo contributes to the 

biodiversity of moist and dry forests and helps 

protect the soil. Its foliage and white flowers add to 

the beauty of the forest. The species has been used 

as an ornamental to a limited extent. 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. Gen. Tech. Rep. SO-58. 

U.S. Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 331 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 
293

Dodonaea viscosa (L.) Jacq. ‘a‘ali‘i 

SAPINDACEAE 

Synonymns: Dodonaea angustifolia L. 

Dodonaea eriocarpa Sm. 

Dodonaea sandwicensis Sherff 

Dodonaea scottsbergii (Sherff) Degener & I. Degener 

Dodonaea spathulata DC. 

Amanda J. Stevens, Sarah A. Taylor, and Randy S. Senock 

General Description.—‘A’ali’i is also known as 

a’ali’i-ku ma kua, a’a’li’i ku makani, lampuaye, 

hop bush, hopseed bush, togovao, and akeake. It is 

usually a many-branched shrub or small tree 

reaching a height of 4.5 to 9.0 m, and 7.5 cm in 

diameter (Little and Skolmen 1989). The bark is 

reddish brown to gray or blackish gray, finely 

furrowed, black heartwood. Leaves are simple, 

alternate, and membranous to chartaceous, 3 to 10 

cm long and 1 to 3 cm wide and vary in shape 

from elliptic to spatulate, oblanceolate or 

lanceolate, entire margins, finely hairy when 

young, yellow green, sticky coating, slightly shiny 

above, and paler below. Petioles are 0.1 to 1.1 cm 

long and slightly winged (Wagner and others 

1990, Little and Skolemen 1989, Lamb 1981). 

Range.—‘A’ali’i has a pantropical distribution 

occuring in temperate regions of Australia, Africa, 

Mexico, New Zealand, India, Samoa, Guam, 

Northern Mariana Islands, Virgin Islands, Puerto 

Rico, Florida, Arizona, South America, Hawaii 

and elsewhere (West and Noble 1984). ‘A‘ali‘i 

occurs on all the main Hawaiian islands except 

Kaholawe. 

Ecology.―‘A’ali’i grows on most types of soils 

and substrates in Hawaii, from old deeply leached 

soils, to fertile soils high in organic materials, to 

barren ‘a‘a and pahoehoe lava, and ash and cinder. 

A’ali’i seedlings are considered fairly shade 

intolerant and usually grow in open sites (Williams 

and Buxton 1989). The ‘a‘ali‘i has a broad 

ecological amplitude and can occur from near sea 

level up into the sub-alpine zone, and from wet 

forests to arid shrub and grasslands. It resists 

droughts well and is generally one of the heartiest 

plants in the arid shrublands of leeward Hawaii. In 

response to wildfire ‘a’ali’i resprout from both the 

root crowns and from seed. Following wildfire in 

diverse ‘a‘ali‘i-dominated montane shrublands, it 

doubled its preburn density within 1 year. Its 

density, however, declined over subsequent years 

but did regain preburn cover values 4 years 

following the burns (Sherry and others 1999). In 

Hawaii, a’ali’i can be affected by a yellowing 

disease distinguished by chlorotic witches’-

brooms. Infected areas of the plants fail to produce 

flowers, while flowering is normal on the 

nonafflicted parts of the same plants. The terminal 

leaves of the portion of the afflicted branches 

become chlorotic, crinkled, and reduced in size. 

This disease can affect all stages of maturity, and 

plants will eventually die. A relatively small 

portion of these plants are infected in Hawaii, but 

this disease is spreading and can be found on all 

the major islands (Borth and others 1995). 

Reproduction.—The flowers are unisexual, 

sometimes perfect, in paniculate inflorescences 

usually 1 to 5 cm long, clustered at the ends of the 

twigs (Wagner and others 1990). The flowers are 

sticky with fine hairs, three to five sepals, and 

without petals. Female flowers have four stigmas 

and male flowers have ten stamens (Little and 

Skolmen 1989). Male and female flowers are born 

on different trees. The fruit is a capsule straw-
294 

colored to dark reddish purple, exceedingly 

variable, turgid to inflated bladder or compressed. 

Seeds are compressed-ovoid, about 3 mm long. 

The two to four-winged, papery capsules are either 

dispersed by wind or adhere to plumage of birds 

(Burrows 1995). The seed coats of most of the 

‘a’ali’i seeds are impermeable to water and there 

also appears to be a micropylar plug present. The 

hard seed coat and the micropylar plug may play a 

role in delayed germination. Some evidence 

suggests that higher temperatures, which may 

enhance the breakdown of the plug, as well as 

scarification of the seed coat, may be necessary for 

germination (Burrows 1995). ‘A‘ali‘i is able to 

reproduce through sprouting following 

disturbance. 

Growth and management.—‘A‘ali‘i is popular as 

an ornamental in xeriscape gardens. Seedlings 

should be planted from containers after last frost in 

any kind of soil. Water infrequently once 

established and prune if a more formal appearance 

is desired. Plant emits strong odor during pollen 

season. Plant is frost damaged at approximately –

6.6° C (Anonymous 1999). 

Benefits.—‘A‘ali‘i has enormous potential as a 

landscape species. ‘A’ali’i also possesses many 

medicinal properties and has been used by native 

peoples from all regions where it is found. Its 

leaves can be used to combat rheumatism, skin 

infections, fevers, swellings, aches, and 

“gastrointestinal disorders” including diarrhea, and 

can be used as a antispasmodic agent (Rojas and 

others 1996). The Australian aborigines used the 

leaves and roots as a painkiller to soothe 

toothaches and headaches (Crib and Crib 1981). 

The flowers (or hops) of ‘a’ali’i are used as a 

“home-brew” substitute to bestow a bitter flavor, 

and also used as a tonic. A red dye is extracted 

from the fruit, and the capsules are also commonly 

used in leis (Little and Skolmen 1989). In India the 

seeds are used as fish poison (Wagner and others 

1987). The wood from ‘a’ali’i is extremely hard 

and has been used for tools, spears and weapons 

(Lamb 1981). In India ‘a’ali’i is an important 

source of fuelwood (Jain 1998). 

References

Anonymous. 1999. Dodonaea viscosa: Hopbush. 

www.plantadviser.com/plants/dodovisc.shtml 

Borth, W.B., J.S. Hu, B.C. Kirkpatrick, D.E. 

Gardner, and T.L. German. 1995. Occurrence of 

phytoplasmas in Hawaii. Plant Disease. 79(11): 

1,094-1,097. 

Burrows, C.J. 1995. Germination behavior of the 

seeds of six New Zealand woody plant species. 

New Zealand Journal of Botany 33: 365-377. 

Cribb, A.B. and Cribb, J.W. 1981. Wild Medicine 

in Australia. Collins, Sydney. 228 p. 

Jain, R.K. 1998. Fuelwood characteristics of 

medium tree and shrub species of India. In: 

Bioresource technology. Volume 68, Issue 3. 

Elsevier Science Limited, Essex, UK. p. 305-

308. 

Lamb, S.H. 1981. Native trees and shrubs of the 

Hawaiian Islands. The Sunstone press, Santa 

Fe, NM. 160 p. 

Little, E.L. and R.G. Skolmen. 1989. Common 

forest trees of Hawaii (native and introduced). 

Agriculture Handbook. 679. U.S. Department of 

Agriculture, Washington, DC. 321 p. 

Pengelly, A. Dodonaea viscosa. www.nhaa.org.au/ 

forum/text/dodvis2.html. 

Rojas, A., S. Cruz, H. Ponce-Monter, and R. Mata. 

1996. Smooth muscle relaxing compounds from 

Dodonaea viscosa. Planta Medica. 62:154-159. 

Sherry, K., J.M. Castillo, and R.B Shaw. 1999. 

Effects of wildfire on vegetation and rare plants 

in arid montane shrublands. Pohakaloa training 

area, Hawaii. 1999 Hawaii Conservation 

Conference, Honolulu, HI. 

Wagner, H., C. Ludwig, L. Grotjahn, and M.S.Y. 

Khan. 1987. Phytochemistry 26(3): 697-701. 

Wagner, W.L., D.R. Herbst, and S.H. Sohmer. 

1990. Manual of the Flowering Plants of 

Hawaii. University of Hawaii Press, Honolulu, 

HI. 1,854 p. 

Williams, P.A. and R.P. Buxton. 1989. Response 

to reduced irradiance of 15 species of native and 

adventive shrub and tree seedlings from eastern 

Canterbury. New Zealand Journal of Ecology. 

12: 95-101. 

West, J.G. and I.R. Noble. 1984. Analyses of 

digitized leaf images of the Dodonaea viscosa

complex in Australia. Taxon 33(4):595-613.
295

Dryas octopetala L. white mountain avens

ROSACEAE 

Synonyms: Dradetum octopetalae Keiner 

Juanita A.R. Ladyman 

Drawing source: USDA—Forest Service collection, 

Hunt Institute 

General Description.—Dryas octopetala is 

commonly known as alpine dryad, mountain 

avens, or white mountain avens in areas where 

more than one Dryas species occur (Stace 1997, 

Kershaw and others 1998, Carter 1988). The 

epithet “dryad” and “dryas” is derived from the 

mythical oak-nymph, Dryas, because the shape of 

its leaves resembles that of an oak species 

(Gledhill 1992, Zwinger and Willard 1996). This 

prostrate, trailing perennial shrub is usually less 

than 30 cm tall, but can grow up to 50 cm, and 

commonly forms mats. The oval to oblong leaves 

are 5 to 25 cm long and the margins are coarsely 

round-toothed (crenate) and somewhat rolled 

under (Carter 1988, Kershaw and others 1998, 

Stace 1997). Typically the shrub is described as 

“evergreen” but has also been observed as “semideciduous,” because it loses some of its foliage in 

winter (Williams and others 1986). The leaves are 

dark green, wrinkled and hairless on the upper side 

but densely white-hairy on the lower surface 

(Kershaw and others 1998). The stalk, midrib, and 

veins have yellowish-brown glands on the lower 

surface. Each flower has eight (sometimes nine or 

10) egg-shaped white, rarely yellowish, petals and 

lance-shaped sepals. The flowers are 2 to 3 cm 

across and are solitary on hairy stalks that are 1 to 

20 cm long. The seeds (achenes) are 2.5 to 4 mm 

long with a feathery plume (style) at the tip 

(Komarov 1941). The chromosome number is 18 

(Stace 1997, Welsh and others 1993). The species 

tends to be heterogeneous throughout its range. 

The stature and organs of white mountain avens 

plants in North America and a part of the Western 

Arctic appear to have smaller dimensions than 

those in Russia and alpine Western Europe 

(Komarov 1941). At least four subspecies and 

several varieties of Dryas octopetala are 

recognized (Hitchcock and Cronquist 2001, 

Kartesz 1994): Dryas octopetala L. ssp. 

alaskensis (Porsild) Hultén, ssp. hookeriana (Juz.) 

Breit., ssp. punctata (Juz.) Hultén, and ssp. 

octopetala of which there are at least three 

varieties; namely var. angustifolia Hitch., var. 

kamtschatica (Juz.) Hultén, and var. luteola

Hultén. Some other varieties are described in 

Kartesz (1994) and Komarov (1941). 

Range.—White mountain avens is a circumpolar 

species occurring in arctic and alpine zones in 

Europe, Asia, and North America. In North 

America, it grows in sub-alpine and alpine tundra 

zones from Greenland, Labrador, and Alaska to 

the Cascade Mountains in Washington and Oregon 

and through the Rocky Mountains to Montana, 

Wyoming, and Colorado and west to high 

elevation sites in Utah and Idaho (Britton 1901, 

Hitchcock and Cronquist 2001, Welsh and others 

1993, Kershaw and others 1998). It is found in 

Scandinavia, the British Isles, Ireland, and the 

Alps, Pyrenees, Carpathians and Dinarid 

Mountains of Europe (Praprotnik 1997, Ellenberg 

1988). It also dominates the tundras of Novaya 

Zemlya, Wrangel Island, the Urals and mountin 

slopes in Siberia (Komárková 1979, Botch and 

Masing 1983). 

Ecology.—Mountain avens is often the dominant 

or co-dominant species in areas where it occurs 

throughout its range. The elevation at which it 
296 

grows is associated with latitude. Plants are found 

at 3,500 m to 3,965 m in Utah whereas they may 

occur at 100 m or less in northern parts of its 

range, such as Greenland (Borge Wills 1996, 

Welsh and others 1993). It typically grows above 

the timberline in sites with low snow cover on 

calcareous or basic soils (Dorn 2001, Dorn 1984, 

Komárková 1979, Ellenberg 1988, Stace 1997, 

Weber and Wittman 2001). It is not an indicator of 

calcareous sites per se as it is common on granitic 

substrates in the Colorado Rocky Mountains 

(Weber 1960). However, on this substrate calcium 

is likely available to the plants as a leachate 

(Weber and Wittman 2001). Mountain avens 

grows in dry meadows, rocky tundra, fellfields, 

and in relatively moist sites in fen lands. In Russia, 

it has been reported as a co-dominant in “frost 

mound bogs” that are essentially permanently 

frozen ridges, or mounds, that alternate with wet 

hollows (Botch and Masing 1983). In contrast, 

mountain avens and lichens are of primary 

importance on the dry upland meadows in the 

Alaskan Arctic and on the windswept fellfield 

slopes of the Rocky Mountains of Colorado U.S.A.

(Komárková 1979, Langenheim 1962). These 

shrubs have an trellis-like growth habit clinging 

closely to rocks and stones and anchoring its roots 

in cracks or between stones. The radiation is so 

intense at high elevations that the temperature can 

rise to more than 65 o

C where the ground is dark 

colored. In these locations the prostrate seedlings 

and young plants can be heat damaged (Ellenberg 

1988). A mutualistic association exists between 

mountain avens and ectomycorrhizal fungi 

(Monson and others 2001). The most dominant 

fungus is Cenococcum geophilum, an ascomycete. 

Reproduction.—White mountain avens reproduce 

by seed. Flowering is from June into August. The 

flowers always face the sun on calm, sunny days, 

and their parabolic shape directs the sun’s heat on 

to the stigma, which can then become up to 3.5 o

C 

warmer than the surrounding air temperature 

(Kershaw and others 1998). The warmer 

temperatures can attract insects and also speed 

pollen germination. Hybridization occurs between 

sympatric Dryas species (Komarov 1941). Seed is, 

at least in part, distributed by wind. 

Growth and Management.—White mountain 

avens seeds have physiological dormancy but do 

not appear to require stratification for germination 

(Baskin and Baskin 2001). Forty-eight percent of 

freshly matured seeds germinated after 4 days in 

the light at 25 o

C (Baskin and Baskin 2001). 

However, in England seeds failed to germinate 

when sown outdoors in the autumn while seeds did 

geminate the following spring, suggesting that 

cold stratification lowers the temperature 

requirement for germination (Elkington 1971). The 

stems root easily and contribute to the rapid spread 

of plants. As part of studies on the consequences 

of global climate change, experiments have been 

made on the response of white mountain avens to 

elevated temperatures in the Colorado Rocky 

Mountains (Welker and others 2001). The results 

suggested that warmer, wetter conditions would 

initially lead to greater productivity (increased leaf 

mass and higher numbers of seed per ramet) 

although decomposition may be retarded, leading 

to a lower nutrient availability in subsequent years 

(Welker and others 2001). Under longer term 

conditions, a switch in the source-sink activity of 

the community occurred. Under current conditions 

the mountain avens community is a carbon sink of 

about 8 g CO2

-C m-2. However, after 4 years of 

experimental warming, the community switched to 

a carbon source of almost 10 g CO2

-C m-2 per 

summer due primarily to an increase in the rate of 

respiration without a concomitant increase in 

photosynthesis (Welker and others 2001). This 

switch in source sink activity has been reported for 

Dryas and other communities in the Arctic 

(Welker and others 2001, Oechel and others 1993). 

This finding may be significant when considering 

the repercussions of increasing industrial 

“greenhouse gas” emissions 

Benefits.—White mountain avens is a pioneer 

species and is important in stabilizing the thin soils 

on mountain slopes (Zwinger and Willard 1996, 

Ellenberg 1988). It spreads relatively rapidly and 

is popular in landscape rock gardens (Williams 

and others 1986). Several cultivated varieties, 

including hybrids, are available (Kershaw and 

others 1998). One hybrid, xSuendermannii

Kellerer, between Dryas octopetala and D. 

drummondii is morphologically similar to D. 

octopetala, but the flowers are yellow in bud 

becoming white and nodding upon opening 

(Bailey and Bailey 1976). White mountain avens is 

an important food source for ptarmigian (Nelson 

1992) and pikas (Martin and others 1951). 

References. 

Bailey, L.H., E.Z. Bailey, and the staff of the 

Liberty Hyde Bailey Hortorium. 1976. Hortus 

Third. Macmillian Publishing Company, New 

York, NY. 1,290 p. 
297

Baskin, C.C. and J.M. Baskin. 2001. Seeds, 

ecology, biogeography, and evolution of 

dormancy and germination. Academic Press, 

New York, NY. 666 p. 

Borge Wills, L. 1996. Sanderling review. Marine 

and Coastal Species Information System, 

Conservation Management Inst., Roanoke, VA. 

http://fwie. fw.vt.edu [not paged]. 

Botch, M. and V. Masing. 1983. Mire ecosystems 

in the U.S.S.R. In: A.J.P. Gore, ed. Ecosystems 

of the World, Vol. 4B. Elsevier Company, New 

York. p. 95-152. 

Britton, N.L. 1901. Manual of the flora of the 

northern states and Canada. Henry Holt and 

Company, New York, NY. 1,080 p. 

Carter, J.L. 1988. Trees and shrubs of Colorado. 

Johnson Books, Boulder, CO. 165 p. 

Dorn, R. D. 1984. Vascular plants of Montana. 

Mountain West Publishing Cheyenne, WY. 412 

p. 

Dorn, R. D. 2001. Vascular plants of Wyoming. 

Mountain West Publishing Cheyenne, WY. 412 

p. 

Elkington, T.T. 1971. Biological flora of the 

British Isles. Dryas octopetala. Journal of 

Ecology 59: 887-905. 

Ellenberg, H. 1988. Vegetation ecology of central 

Europe. 4th edition. Cambridge University Press, 

New York, NY. 731 p. 

Gledhill, D. 1992. The names of plants. 

Cambridge University Press, Cambridge, 

England. 202 p. 

Hitchcock, C.L. and A. Cronquist. 2001. Flora of 

the Pacific Northwest. University of Washington 

Press, Seattle, WA. 730 p. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada and 

Greenland. Vol. 1 - Checklist. Timber Press, 

Portland, OR. 622 p. 

Kershaw, L., A. MacKinnon, and J. Pojar. 1998. 

Plants of the Rocky Mountains. Lone Pine 

Publishing, Edmonton, Canada. 383 p. 

Komárková, V. 1979. Alpine vegetation of the 

Indian Peaks area, Front Range, Colorado Rocky 

Mountains. J. Cramer, Vaduz, Germany. 

Komarov, V.L. 1941. Flora of the U.S.S.R. 

Volume 10. Izdatel’stvo Akademii Nauk SSSR. 

Translated by N. Landau 1970. The Smithsonian 

Institution, Washington, D.C. p. 272-273. 

Langenheim, J.H. 1962. Vegetation and 

environmental patterns in the Crested Butte area. 

Ecological Monographs 32(3): 249-285. 

Martin, A.C., H.S. Zim, and A.L. Nelson. 1951. 

American wildlife and plants. McGraw-Hill 

Book Company, New York, NY. 500 p. 

Monson, R.K., R. Mullen, and W.D. Bowman. 

Plant nutrient relations. 2001. In: W.D. Bowman 

and TR. Seastedt, eds. Structure and function of 

an alpine ecosystem. Oxford University Press, 

New York, NY. 337 p. 

Nelson, R.A. 1992. Handbook of Rocky Mountain 

plants. Robert Rinehart Publishers, Niwot, CO. 

444 p. 

Oechel, W.C., S.J. Hastings, G.L. Vourlitis, M.A. 

Jenkins, G. Riechers, and N. Grulke. 1993. 

Recent changes of arctic tundra ecosystems from 

a carbon sink to a source. Nature 361: 520-523. 

Praprotnik, N. 1997. Alpski botanicni vrt Juliana v 

Trenti. Prirodoslovni muzej Slovenije, 

Ljubljana, Slovenia. 126 p. 

Stace, C. 1997. New Flora of the British Isles. 2nd

edition. Cambridge University Press, New York, 

NY. 1,130 p. 

Weber, W.A. 1960. Some features of the 

distribution of Arctic relicts at their Austral 

limits. Twelfth International Botanical Congress. 

Recent Advances in Botany. p. 912-914 

Weber, W.A., and R.C. Wittman 2001. Colorado 

flora, western slope. University Press of 

Colorado, Boulder, CO. 488 p. 

Welker, J.M., W.D. Bowman, and T.R. Seastedt. 

2001. Environmental change and future 

directions in alpine research. In: W.D. Bowman 

and T.R. Seastedt, eds. Structure and function of 

an alpine ecosystem. Niwot Range, Colorado. 

Oxford University Press, New York. 337 p. 
298 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1993. A Utah flora. Second 

Edition, Brigham Young University, Provo, UT. 

986 p. 

Williams, J., R. Radebaugh, D. Hall, G. Keladis, P. 

Keladis, and P. A. Pachuta. 1986. Rocky 

Mountain Alpines. Timber Press, Portland, OR. 

333 p. 

Zwinger, A.H. and B.E. Willard. 1996. Land above 

the trees, a guide to American alpine tundra. 

Johnson Printing, Boulder, CO. 425 p. 
299

Duranta erecta L. golden dewdrop 

VERBENACEAE 

Synonyms: Duranta repens L. 

 Duranta ellisia Jacq. 

 Duranta plumieri Jacq. 

John K. Francis 

General Description.—Golden dewdrop, also 

known as skyflower, pigeon berry, angels-whisper, 

duranta, cuentas de oro, azota-caballo, fruita de 

paloma, lila, lluvia, and grão de galo, is an upright 

to drooping shrub that sometimes takes the form of 

a scrambling shrub or rarely a small tree (Liogier 

1995, Little and others 1974). It usually matures 

with a height 1 to 3 m. The species develops tap 

and lateral root systems with abundant fine roots. 

There are normally several stems arising from 

below and just above the ground line. Crowns of 

golden dewdrop are replete with fine branches and 

twigs that are often thorny. The bark is light gray, 

becoming rough and fissured when old. Lightgreen, opposite leaves are elliptic to ovate and 1.5 

to 8 cm long. Light-blue, lavender, or white, 

tubular, five-lobed flowers are borne on terminal or 

axillary racemes. The yellow or yellow-orange 

fleshy fruits are ellipsoidal with five lobes and 

grow in hanging clusters. These fruits may contain 

up to eight nutlets (Hardin and Arena 1969, 

Howard 1989, Liogier 1995, Little and others 

1974). 

Range.—Golden dewdrop is a native of Mexico, 

Central America, South America to Argentina, 

southern Florida (possibly naturalized), Bermuda, 

the Bahamas, and the West Indies (Howard 1989, 

Liogier 1995, Little and others 1974). The species 

is widely cultivated and escaped in the tropics and 

subtropics including Hawaii, American Samoa, and 

Guam (Pacific Islands Ecosystems at Risk 2002). 

Ecology.—Golden dewdrop grows wild mostly in 

dry coastal areas (750 to 900 mm of annual 

precipitation in Puerto Rico) from near sea level to 

over 100 m in elevation (Little and others 1974). It 

also grows in disturbed areas in moister habitat, 

especially along roads (Pacific Island Ecosystems 

at Risk 2002). Because it is moderately intolerant 

of shade and does not compete well with taller 

vegetation, golden dewdrop is usually found in 

rocky or sandy areas with low shrubs and sparse 

grass and herbs. Although it is more common on 

limestone, the shrub also grows in areas with 

igneous rocks. The species tolerates light to 

moderate salt spray. In natural stands, golden 

dewdrop may be found in small tangled stands or 

as occasional plants. Plants under droughty and 

infertile natural conditions tend to be short (about 1 

m in height). Where cultivated in deep, moist, 

fertile soils, golden dewdrop grows faster and to 

larger sizes. As an ornamental, it can be grown in 

moderate shade at elevations up to 1,300 m 

(Bruggeman 1974). Plants in the northerly extent 

of the naturalized range die to the ground after 

frosts but resprout from the roots in the spring 

(Floridata 1999). The species is subject of attack by 

scale, mealy bugs, caterpillars, and nematodes 

(Watkins and Sheehan 1975, Woman’s Club of 

Havana 1958). 

Reproduction.—Natural stands of golden dewdrop 

in Puerto Rico flower and fruit in spring and 

summer (Little and others 1974). However, 

ornamental plants in Puerto Rico flower and fruit 
300 

throughout the year (author’s observation), and in 

Florida, both flowers and fruit are present on the 

shrubs at the same time (Nelson 1996). A 

collection of golden dewdrop fruits from Puerto 

Rico averaged 0.515 + .009 g/fruit. Air-dried seeds 

separated from them averaged 0.0346 + 0.0005 

g/seed or 28,900 seeds/kg. Set to germinate on 

moist filter paper without any pretreatment, 29 

percent of the seeds germinated beginning 128 

days after sowing (Francis and Rodíguez 1993). 

Birds disseminate the seeds (Watkins and Sheehan 

1975). However, seedlings are seldom common. 

After establishment, plants thicken and spread 

laterally a few decimeters vegetatively. Golden 

dewdrop are produced commercially from seeds, 

cuttings, and by layering (Floridata 1999). 

Growth and Management.—Golden dewdrop 

plants live at least 15 years. The species has a 

moderate growth rate, usually about 0.5 m/year for 

the first few years. Ornamental plants need regular 

thinning and pruning to keep them under control 

and to remove dead branches (Floridata 1999). 

Although they withstand poor soil and drought 

well, plants grow best if watered and fertilized 

(Woman’s Club of Havana 1958). 

Benefits and Detriments.—Golden dewdrop 

forms a part of the coastal scrub community and 

contributes to soil and ecosystem stability. It is a 

popular ornamental used for accent plants and 

hedges in tropical and subtropical parts of the 

world because of its profuse displays of flowers 

and fruits (Floridata 1999). Golden dewdrop 

flowers attract butterflies and hummingbirds 

(Floridata 1999). It is sometimes grown in 

greenhouses in areas too cold for natural plants. 

The wood is light brown and hard and useful for 

stakes and fuel. Golden dewdrop is a poisonous 

plant and has caused deaths of children. Saponins 

in the fruits and foliage cause gastroenteric 

irritation, drowsiness, fever, nausea, vomiting, and 

convulsions. Dermatitis sometimes occurs from 

handling the plants (Hardin and Arena 1969, 

Westbrooks and Preacher 1986). However, birds 

feed on the fruits without difficulty (Nelson 1996). 

Ethyl acetate and aqueous extracts of leaves 

showed significant antimalarial activity when 

administered to mice (Castro and others 1996). In 

small quantities, fruits are used to treat intestinal 

worms (Whistler 2000). 

 

References 

Bruggeman, L. 1964. Tropical plants and their 

cultivation. Thames and Hudson, London. 228 p. 

Castro, O., M. Barrios, M. Chinchilla, and O. 

Guerrero. 1996. Chemical and biological 

evaluation of the effects of plant extracts against 

Plasmodium berghei. Revista de Biologia 

Tropical 44(2A): 361-367. 

Floridata. 1999. Duranta erecta. http:// 

floridata.com/ref.d.dra_ere.cfm. 3 p. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

Hardin, J.W. and J.M. Arena. 1969. Human 

poisoning from native and cultivated plants. 

Duke University Press, Durham, NC. 167 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles. 

Vol. 6. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p.

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Pacific Island Ecosystems at Risk. 2002. Duranta 

erecta L., Verbenaceae. http://www.hear. 

org/pier3/duere.htm. 2 p. 

Watkins, J.V. and T.J. Sheehan. 1975. Florida 

landscaping plants, native and exotic. The 

University Presses of Florida, Gainesville, FL. 

420 p. 

Westbrooks, R.G. and J.W. Preacher. 1986. 

Poisonous plants of Eastern North America. 

University of South Carolina Press, Columbia, 

SC. 226 p. 

Whistler, W.A. 2000. Tropical ornamentals, a 

guide. Timber Press, Inc., Portland, OR. 542 p. 
301

Woman’s Club of Havana. 1958. Flowering plants 

from Cuban gardens. Criterion Books, New 

York. 365 p. 
302 

Encelia farinosa Gray ex Torr. brittlebush 

ASTERACEAE 

Synonyms: none 

Catherine E. Koehler and Arlee M. Montalvo 

General Description.—Brittlebush, a dry 

environment-adapted shrub with weak branches, is 

also known as incienso because it has fragrant 

resin, and as desert encelia because it occurs in 

desert and semidesert habitats. The specific epithet 

farinosa refers to the mealy pubescence of the 

leaves, which is composed of appressed wooly 

hairs. These shrubs have dense rounded canopies 

with alternate whitish to greenish-gray leaves 

clustered near stem tips. Adults are 0.3 to 1.5 m 

tall, shallowly rooted (40 cm), with one to several 

many-branched stems. Leaves are simple, 3 to 8 

cm long, ovate to rhombic in outline, with wavy 

margins, and short petioles. Heads are arranged in 

loose, naked panicles and have both disk and ray 

flowers subtended by green, glandular bracts that 

embrace the flat, obovate, 4.5 mm long achenes. 

Involucre bracts are imbricate in three to four 

series. Disk florets are yellow or rich purple and 

are surrounded by 8 to 12 mm long yellow ray 

florets (Munz and Keck 1968, Hickman 1993). 

Munz and Keck (1968) recognize two varieties: E. 

f. var. radians Bdg. ex Blake from southeastern 

California with glabrate leaves and involucre and 

yellow disk flowers, and E. f. var. phenocodonta

(Blake) Jtn. from the southern Colorado desert 

with purple disk flowers. Plants have n = 18 

chromosomes (Hickman 1993). Brittlebush

hybridizes readily with E. fructescens (A. Gray) A. 

Gray, E. californica Nutt. and E. actoni Elmer

(Ehleringer and Clark 1988, Hickman 1993). 

Intergeneric hybrids with the annual Gerea 

canescens T. & G. have also been reported (Kyhos 

1967). 

Range.—Brittlebush occurs within sage scrub of 

the inland valleys of southern California, eastward 

and southward into arid habitats of the Sonoran 

and Mojave deserts and into Arizona, northwestern 

Mexico, southern Nevada, and southwestern Utah 

at under 1,000 m elevation (Munz and Keck 1968, 

Hickman 1993, Sandquist and Ehleringer 1997). In 

California, it has been extending westward and 

northward due to roadside and other plantings. 

Geographic Variation.—Geographic variation 

and adaptation to local environments have been 

documented for many brittlebush traits. In 

particular, adaptation to water availability has been 

well demonstrated, including genetic 

differentiation in water use efficiency (Sandquist 

and Ehleringer 2003). Also, individuals with 

brown-purple disk florets (var. phenicodonta) 

occur in areas with higher levels of soil moisture 

and are replaced by the yellow-disked form (var. 

farinosa) in drier sites. This pattern may involve 

natural selection in response to water availability 

(Kyhos 1971) but may also be linked to the lower 

frost tolerance of var. phenicodonta (Sandquist 

and Ehleringer 1996). Similarly, Monson and 

others (1992) found localized physiological and 

genotypic differences in water use between plants 

at the base and the top of a slope that coincided 

with a moisture stress gradient. 

The degree of leaf pubescence varies across 

regions with different mean annual rainfall, and 

variation in number of leaf hairs is both a plastic 

response and genetically determined. Leaves of 

plants growing in arid regions are more pubescent, 

thereby having greater control over leaf 

temperature and water loss, but they have lower 

photosynthetic capacity due to higher reflectance 

of light than plants from more mesic regions. 

Differences are maintained when offspring are 

planted together in common gardens (Sandquist 

and Ehleringer 1997, Housman and others 2002). 

However, an additional plastic response allows
303

plants in more mesic gardens to grow lesspubescent leaves than sibling plants in desert 

gardens, a response which allows plants to take 

advantage of higher water availability and 

increased photosynthetic ability through leaf-hair 

reduction (Ehleringer and Clark 1988). The 

inverse relationship for leaf pubescence and 

photosynthetic activity may involve pleiotropy or 

linkage to other genetically determined 

morphological differences between populations or 

from environments with different moisture 

regimes (Housman and others 2002). 

There is also clinal variation in production of 

chemical compounds that provide defense against 

herbivores. From north to south in Baja California 

and east to west from the Sonoran desert to coastal 

regions of California, plants produce progressively 

less of a sequiterpene and more of a chromene 

toxin, which may influence local resistance to 

herbivores (Wisdom 1985, Kunze and others 

1995). Seasonal production of these compounds 

may also influence herbivores (Wisdom and 

Rodriquez 1983). There are higher concentrations 

of these chemicals and nitrogen in young tissues. 

In addition, populations differ in the relative 

amount of different compounds. The specialist 

beetle Trirhabda geminata Horn experienced 

lowered larval growth rates when fed higher levels 

of the secondary compounds. 

Ecology.—In sage scrub, brittlebush occurs on 

flats and slopes, primarily on weathered sandstone, 

granite, diorite, and alluvial deposits; in the desert 

it occurs on rocky slopes, flats, and in washes 

(Munz and Keck 1968, Hickman 1993). Studies on 

the adaptive significance of variation in leaf form 

and pubescence both within and among 

populations is reviewed by Housman and others 

(2002) and by Sandquist and Ehleringer (1997, 

1998). Plants react to seasonal increase in water 

stress at the end of the rainy season by replacing 

the larger, less hairy leaves produced earlier in the 

growing season with more pubescent leaves that 

are smaller and thicker. This reduces water loss 

and regulates leaf temperature, but it also 

decreases photosynthetic capacity. Prolonged 

drought leads to dormancy and leaf drop. 

Brittlebush shows variation among 

individuals and populations in water-use efficiency 

(the ratio of photosynthesis to transpiration, or 

delta value). Individuals with a high delta have a 

higher growth response if water stress is decreased 

but perform poorly in response to drought stress, 

while those with a low delta show lower growth 

response under low water stress and a greater 

capacity to survive drought conditions (Ehleringer 

1993). In general, however, brittlebush is drought 

adapted and responds quickly to water addition 

through rapid CO2 uptake, leaf production, and 

stem growth (Nobel and others 1998). 

Reproduction.—Brittlebush flowers from March 

through May (Munz and Keck 1968) and 

individuals must be cross-pollinated in order to 

produce seed (Ehleringer and Clark 1988). 

Flowers are visited by various insects including 

butterflies, moths, flies, bees, wasps, and beetles 

(Kyhos 1971, Moldenke 1976). In one study, a 

beetle (Malachiidae) was found to be 10 times 

more common than all other insect species 

combined (Kyhos 1971). These potential 

pollinators do not discriminate between plants with 

different disk-flower color (Kyhos 1971). Achenes 

are collected from May to July (Mirov and Kraebel 

1939) with about 770,000 bulk seeds/kg (personal 

communication with S&S seeds, Carpenteria, CA). 

Seed viability varies among years, with as few as 

35 percent of seeds viable (personal 

communication with M. Wall, Rancho Santa Ana 

Botanical Garden, Claremont, CA). Seed 

production is influenced by water stress 

heightened by competition. In a desert study in 

which nearby neighbors were removed, shrubs 

experienced lower water stress, had higher 

survival, grew to nearly twice the mass, and 

produced 53 percent more flower heads per twig 

and 220 percent more achenes per head than 

shrubs with brittlebush neighbors within 2 m 

(Ehleringer 1984). 

Establishment and Growth.—Brittlebush 

becomes established rapidly following 

disturbance. Seedlings emerge and become 

established in open areas in the winter rainy 

season. Plants become dormant and drop many 

leaves during the dry season and then sprout new 

leaves with the onset of winter rains. Plants can 

reach maturity within 2 years and often live for 10 

to 15 years. In sage scrub vegetation, resprouting 

success of shrubs from the base is inversely related 

to fire intensity (Westman and others 1981, Martin 

1984). In one study, 2 to 30 percent of brittlebush

resprouted on slopes previously dominated by the 

shrub, and resprouts and seedlings surpassed 

prefire densities within 2 years (Martin 1984). For 

coastal sage scrub in general, both resprouting and 

seedling emergence from a soil seed bank are 

negatively correlated with fire intensity (Keeley 

1998). 

Seed germination.—Some authors report seed 

germination without pretreatment (Mirov and 
304 

Kraebel 1939, Emery 1988), but pretreatment can 

increase otherwise low germination rates. Padgett 

and others (1999) found that seed stored for 6 

months at room temperature had 2 to 4 percent 

germination while seed stored at 5 to 10 o

C in a 

refrigerator had 10 to 12 percent germination. In 

addition, treatment with gibberelic acid or 

Ca(NO3)2 increased germination of both warmand cold-stored seeds approximately two-fold, and 

leaching with water for several days increased 

germination by about 50 percent. 

Growth and Management.—Maximum growth 

of roots occurs in the winter and early spring 

(Drennan and Nobel 1996), so plants will establish 

best if sown in late fall. Planting brittlebush seeds 

can yield vegetative cover and visual appeal 

relatively quickly. However, seed mixtures should 

be balanced carefully because overuse can retard 

establishment of other species (Went 1942, Gray 

and Bonner 1948). Ample evidence for local 

adaptation in this species underscores the need to 

collect seed material for wildland restoration from 

within the same ecological zone and vegetation 

type as the targeted planting site so as to maximize 

success of planting projects. Because of potential 

competition and hybridization, it is also important 

that correct native species are specified and used. 

Mistaken plantings of E. californica instead of 

brittlebush, or vice versa, abound (authors’ 

observation), and hybrids between species have 

been found in such locations (personal 

communication with A. Sanders, University of 

California, Riverside). Improper seed choices can 

compromise the success of restoration efforts and 

the genetic integrity of wild populations. 

Benefits.—Brittlebush was used by native tribes 

for medicinal and other purposes. The resinous 

gum, heated or made into a salve, was applied to 

the chest to relieve pain and loosen bronchial 

mucous. A decoction of boiled blossoms, leaves, 

and stems was held in the mouth to relieve gum 

and tooth ache (Bean and Saubel 1972, Moore 

1989). In addition, tea made from the gum has a 

numbing effect and was used to relieve arthritic 

pain (Moore 1989). The resin was also burnt as 

incense or melted and used as a varnish (Moore 

1989, Hickman 1993). 

Brittlebush feeds numerous pollinators and 

herbivores. It is an important nectar and pollen 

source of the bee, Calliopsis pugionis Cockerell, 

which is the host of the rare cleptoparasitic bee, 

Holcopasites ruthae Cooper in Riverside County, 

CA (Visscher and Danforth 1993). Mountain 

sheep eat brittlebush, but it is only found in fecal 

pellets in spring, summer, and fall in trace amounts 

(Perry and others 1987). The dominant herbivores 

on brittlebush leaves are the larvae and adults of 

the beetle, Trirhabda geminata (Wisdom 1985). 

The fly Neotephritis finalis Loew lays its eggs 

between the florets, and the larvae feed on the 

achenes (Goeden and others 1987). 

Horticulture.—Brittlebush does best with full sun 

and good drainage (Hickman 1993, Keator 1994). 

The rounded form with striking yellow flowers is 

attractive near the back of borders or rock gardens, 

and it is especially suitable on dry slopes (Perry 

1992, Keator 1994). Plants can be established 

quickly from seed or containers (Perry 1992). 

Provision of occasional summer water allows 

plants to remain attractive throughout the year 

(Keator 1994). 

References 

Bean, J.L. and K.S. Saubel. 1972. Temalpakh: 

Cahuilla Indian Knowledge and Usage of Plants. 

Malki Museum Press, Morongo Indian 

Reservation, CA. 225 p. 

Drennan, P.M. and P.S. Nobel. 1996. Temperature 

influences on root growth for Encelia farinosa

(Asteraceae), Pleuraphis rigida (Poaceae), and 

Agave deserti (Agavaceae) under current and 

doubled CO2 concentrations. American Journal of 

Botany 83: 133-139. 

Ehleringer, J.R. 1984. Intraspecific competitive 

effects on water relations, growth and 

reproduction in Encelia farinosa. Oecologia 63: 

153-158. 

Ehleringer, J.R. 1993. Variation in leaf carbon 

isotope discrimination in Encelia farinosa: 

implications for growth, competition, and drought 

survival. Oecologia 95: 340-346. 

Ehleringer, J.R. and C. Clark. 1988. Evolution and 

adaptation in Encelia (Asteraceae). In: L. 

Gottlieb, L. Jain and S. Jain, eds. Plant 

Evolutionary Biology. Chapman and Hall, 

London. p. 221-248. 

Emery, D.E. 1988. Seed propagation of native 

California Plants. Santa Barbara Botanical 

Garden, Santa Barbara, CA. 23 p. 

Goeden, R.D., T.D. Cadatal, and G.A. Cavender. 

1987. Life history of Neotephritis finalis (Loew) 

on native Asteraceae in southern California 
305

(Diptera: Tephritidae). Proceedings of the 

Entomological Society of Washington 89: 552-

558. 

Gray, R. and J. Bonner. 1948. An inhibitor of plant 

growth from the leaves of Encelia farinosa. 

American Journal of Botany 35: 52-57. 

Hickman, J.C., ed. 1993. The Jepson Manual: 

Higher Plants of California. University of 

California Press, Ltd., Los Angeles, CA. 1,400 p. 

Housman, D.C., M.V. Price, and R.A. Redak. 

2002. Architecture of coastal and desert Encelia 

farinosa (Asteraceae): consequences of plastic 

and heritable variation in leaf characters. 

American Journal of Botany 89: 1,303-1,310. 

Keator, G. 1994. Complete garden guide to the 

native shrubs of California. Chronicle Books, San 

Francisco, CA. 314 p. 

Keeley, J.E. 1998. Postfire ecosystem recovery and 

management: The October 1993 large fire 

episode in California. In: J.M. Moreno, ed. Large 

Forest Fires. Backbuys Publishers, Leiden, 

Netherlands. p. 69-90. 

Kunze, A., C. Muller, and P. Proksch. 1995. 

Chemical variation and defense of Encelia

farinosa. Biochemical Systematics and Ecology 

23: 355-363. 

Kyhos, D.W. 1967. Natural hybridization between 

Encelia and Geraea (Compositae) and some 

related experimental investigations. Madroño 19: 

33-43. 

Kyhos, D.W. 1971. Evidence of different 

adaptations of flower color variants of Encelia

farinosa (Compositae). Madroño 21: 49-61. 

Martin, B.D. 1984. Influence of slope aspect on 

postfire reproduction of Encelia farinosa

(Asteraceae). Madroño 31: 187-189. 

Mirov, N.T. and C.J. Kraebel. 1939. Collecting and 

handling seeds of wild plants. Forestry 

Publication 5. Civilian Conservation Corps, 

United States Government Printing Office, 

Washington, D.C. 

Moldenke, A.R. 1976. California pollination 

ecology and vegetation types. Phytologia 34: 305-

361. 

Monson, R.K., S.D. Smith, J.L. Gehring, W.D. 

Bowman, and S.R. Szarek. 1992. Physiological 

differentiation within an Encelia farinosa

population along a short topographic gradient in 

the Sonoran Desert. Functional Ecology 6: 751-

759. 

Moore, M. 1989. Medicinal Plants of the Desert 

and Canyon West. Museum of New Mexico 

Press, Santa Fe, NM. 184 p. 

Munz, P.A. and D.D. Keck. 1968. A California 

Flora with Supplement. University of California 

Press, Berkeley, CA. 1,681 + 224 p. 

Nobel, P.S., H. Zhang, R. Sharifi, M. Castañeda, 

and B. Greenhouse. 1998. Leaf expansion, net 

CO2 uptake, rubisco activity, and efficiency of 

long-term biomass gain for the common desert 

subshrub Encelia farinosa. Photosynthesis 

Research 56: 67-73. 

Padgett, P.E., L. Vázquez, and E.B. Allen. 1999. 

Seed viability and germination behavior of the 

desert shrub Encelia farinosa Torrey & A. Grey 

(Compositae). Madroño 46: 126-133. 

Perry, B. 1992. Landscape Plants for Western 

Regions: an Illustrated Guide to Plants for Water 

Conservation. Land Design Publishing, 

Claremont, CA. 318 p. 

Perry, W.M., J.W. Dole, and S.A. Holl. 1987. 

Analysis of the diets of mountain sheep from the 

San Gabriel Mountains, California. California 

Fish and Game 73: 156-162. 

Sandquist, D.R. and J.R. Ehleringer. 1996. Potential 

adaptability and constraints of response to the 

changing climates for Encelia farinosa var. 

phenicondonta from southern Baja California, 

Mexico. Madroño 43: 465-478. 

Sandquist, D.R. and J.R. Ehleringer. 1997. 

Intraspecific variation of leaf pubescence and 

drought response in Encelia farinosa associated 

with contrasting desert environments. New 

Phytologist 135: 635-644. 

Sandquist, D.R. and J.R. Ehleringer. 1998. 

Intraspecific variation of drought adaptation in 

brittlebush: Leaf pubescense and timing of leaf 

loss vary with rainfall. Oecologia 113: 162-169. 

Sandquist, D.R. and J.R. Ehleringer. 2003. Carbon 

isotope discrimination differences within and 
306 

between contrasting populations of Encelia 

farinosa raised under common-environmental 

conditions. Oecologia 134: 463-470. 

Visscher, P.K. and B.N. Danforth. 1993. Biology of 

Calliopsis pugionis (Hymenoptera: Andrenidae): 

nesting, foraging, and investment sex ratio. 

Annals of the Entomological Society of America 

86: 822-832. 

Went, F.W. 1942. The dependence of certain annual 

plants on shrubs in southern California deserts. 

Bulletin of the Torrey Botanical Club 69: 100-

114. 

Westman, W.E., J.F. O'Leary, and G.P. Malanson. 

1981. The effects of fire intensity, aspect and 

substrate on post-fire growth of California coastal 

sage scrub. In: N.S. Margaris and H.A. Mooney, 

eds. Components of Productivity of 

Mediterranean-Climate Regions: Basic and 

Applied Aspects. W. Junk Publishers, Boston, 

MA. p. 151-179. 

Wisdom, C.S. 1985. Use of chemical variation and 

predation as plant defenses by Encelia farinosa

against a specialist herbivore. Journal of 

Chemical Ecology 11: 1,553-1,565. 

Wisdom, C.S. and E. Rodriguez. 1983. Seasonal 

age-specific measurements of the sesquiterpene 

lactones and chromenes of Encelia farinosa. 

Biochemical Systematics and Ecology 11: 345-

352. 
307

Ephedra nevadensis S. Wats. Nevada ephedra 

EPHEDRACEAE 

Synonyms: Ephedra aspera Engelm. ex S. Wats 

 Ephedra reedii Cory 

 Ephedra antisyphilitica sensu S. Watts in King 

Stanley G. Kitchen 

General Description.—Nevada ephedra, also 

known as Nevada jointfir, gray ephedra, or 

mormon tea, is an erect, 0.3 to 1.5 m tall, 

coniferous shrub with opposite spreading 

branches. Roots are deep and fibrous extending 

from an expanded root crown. Stems are jointed 

and grooved. Internodes are 1.5 to 6 cm long 

(Cronquist and others 1972). The combination of 

small cones, ring porous wood, wide multiseriate 

rays, and vessels in older stems separate Ephedra

from other gymnosperms (Stevenson 1993). Bark 

on older branches is gray and fissured. The erect, 

photosynthetic branchlets (1 to 5 mm wide) are 

pale green to gray, glaucous and smooth. Leaves 

are reduced to 2 to 5 mm long, deciduous scales, 

positioned in opposite pairs at the nodes. A 

thickened gray base remains after leaf-fall. One to 

several oval-shaped male cones, 4 to 8 mm long, 

are subtended by several pairs of membranous 

bracts attached at the nodes (Cronquist and others 

1972). Egg-shaped female cones, 5 to 11 mm long 

and light-brown to yellowish-green, are attached 

on short stalks (peduncles) and partially enclosed 

by 8 to 10 bracts at young nodes. Fruits are smooth 

brown nutlets, 4 to 9 mm long, 2 to 4 mm wide, 

and occur individually or in pairs (Cronquist and 

others 1972). The inner cone scales enclose the 

seed mimicking the angiosperm pericarp. The 

embryo has two cotyledons. 

Taxonomy.—Ephedra is monotypic to 

Ephedraceae with a worldwide distribution (except 

for Australia) and is represented by approximately 

60 species (Stevenson 1993). Two varieties, of 

Nevada ephedra are recognized; E. nevadensis var. 

nevadensis and var. aspera (Engelm. ex S. Wats) 

L. Benson (Kartesz 1994). Chromosome number is 

2n = 14 or 28 (Stevensen 1993). 

Range.—Nevada ephedra is endemic to arid and 

semiarid parts of the Great Basin, Colorado 

Plateau, and desert Southwest of western North 

America. Its range extends from Oregon and 

California on the north and west, into Texas in the 

east, and Mexico in the south (Tirmenstein 1990). 

In Utah it grows from 1,220 to 1,830 m in 

elevation (Dittberner and Olson 1983). 

Ecology.—Nevada ephedra grows in well drained, 

sandy to rocky soils on arid hills, alluvial fans, and 

plains, and on ridges and canyons partially to 

completely devoid of trees (Stanton 1974). It is 

moderately salt-tolerant (Stanton 1974, Plummer 

1977). It occurs in large pure stands and in mixed 

pinyon-juniper woodlands, salt-desert, sagebrush, 

and hot-desert transitional shrublands, and 

mountain and desert grasslands (Tirmenstein 

1990). Mean annual precipitation varies from 150 

to 400 mm. Plants sprout from the woody crown 

and from roots following fire (Young and Evan 

1978, West and Hassan 1985). Fire seasonality, 

intensity, and severity affect plant capacity to 
308 

recover after burning (Stanton 1974, Wright 1980, 

McLaughlin and Bowers 1982). A lack of fuel 

continuity in many Nevada ephedra communities 

results in fire being a rare event. Plants spread 

vegetatively through underground stolons in 

response to burial. Natural recruitment from seeds 

is often from rodent caches (Tirmenstein 1990). 

Of North American ephedra, Nevada ephedra is 

considered to be the most important forage species 

(Tirmenstein 1990). It is used extensively by mule 

deer (Ococolius hemionus), pronghorn 

(Antilocapra americana), and bighorn (Ovis 

canadensis) and domestic sheep during winter and 

in periods of drought (Stanton 1974, Smith and 

Beale 1980). Nutritive value and palatability of 

young branchlets is considered to be fair to good 

(Tirmenstein 1990). 

Reproduction.—Nevada ephedra is dioecious 

(occasional monoecious) with female and male 

cones occurring on separate plants. Male plants are 

more common on dry slopes and female plants 

dominate more favorable locations, maximizing 

reproductive output (Freeman and others 1976). 

Plants are wind-pollinated. The timing of cone 

maturation, or coning, and pollination is weatherdependent and generally occurs over a period of 1 

to 2 weeks between early March and early June 

(Stanton 1974, Turner and Randall 1987). Seeds 

ripen between May and August. Turner and 

Randall (1987) observed considerable inter-annual 

variation in timing of phonological development. 

Cones are not produced every year, and good seed 

crops are typically several years apart for any 

given population (Plummer and others 1968). 

Years of good cone production are apparently 

synchronized among most plants of a population 

by an interaction between years of above-average 

precipitation and time since last major seed crop. 

There are approximately 45,000 seeds (nutlets) per 

kg (Kitchen and others 1999). 

Growth and Management.—Once ripe, seeds are 

easily dislodged. Ripened seeds can be collected 

by gently striking or shaking seed-bearing 

branches over tarps or hand-held hoppers. In good 

years, large quantities of seeds can be harvested in 

relatively short time if harvests are well timed. 

After drying, seeds are cleaned to remove light, 

unfilled seeds, stem fragments, cone scales, and 

other inert material. Purity and viability levels of 

commercially available seed are generally high. 

Seed germination is rapid across a wide range of 

temperatures (5 to 20 o

C) with little or no 

dormancy (Young and others 1977, Kitchen and 

others 1999). Seed viability has remained high for 

up to 5 years (Stevens and others 1981). Although 

seedlings tolerate drought well, they are slow 

growing and may take 5 to 10 years to reach 60 cm 

in height (Plummer and others 1968). Plants 

establish well from direct seeding when more 

competitive species are controlled. Successes with 

transplants and cuttings to revegetate minelands 

and other degraded shrublands have also been 

reported (Luke and Monsen 1984, Shaw and 

Monsen 1984).

Benefits.—Nevada ephedra provides forage and 

cover for numerous species of wildlife, especially 

during winter and periods of drought (Stanton 

1974, Smith and Beale 1980, Dittberner and Olsen 

1983). It is also used as forage by domestic sheep 

and cattle but is considered poor forage for horses. 

The species is used effectively in restoration 

planting on harsh sites (Plummer and others 1968, 

Luke and Monsen 1984, Shaw and Monsen 1984). 

Indigenous humans used roasted seeds in foods 

and various concoctions from seeds and stems to 

treat a variety of symptoms including coughs, 

headaches, cold, fever, and kidney ailments 

(Steward 1938, Tirmenstein 1990). Mormon 

pioneers also made teas from stems. All members 

of the genus produce unique secondary compounds 

with potential medicinal applications (Caveney 

and others 2001). Nevada ephedra, and other 

North American members of the genus, lack the 

nueroactivating ephedrine alkaloids found in some 

Old World species. However, a class of 

compounds with antimicrobial potential, 

kynurenates, are found in Nevada ephedra 

(Caveney and others 2001). It is occasionally used 

as an ornamental (Cronquist and others 1972). 

References 

Caveney, S., D.A. Charlet, H. Freitag, M. MaierStolte, and A.N. Starratt. 2001. New 

observations on the secondary chemistry of 

world Ephedra (Ephedraceae). American 

Journal of Botany 88: 1,199-1,208. 

Cronquist, A., A.H. Holmgren, N.H. Holmgren, 

and J.L. Reveal. 1972. Intermountain flora: 

Vascular plants of the Intermountain West, 

U.S.A. Vol. 1. Hafner Publishing Company, 

Inc., New York. 270 p. 

Dittberner, P.L. and M.R. Olson. 1983. The plant 

information network (PIN) data base: Colorado, 

Montana, North Dakota, Utah, and Wyoming. 

FWS/OSB-83/86. U.S. Department of the 

Interior, Fish and Wildlife Service, Washington 
309

DC. 786 p. 

Freeman, D.C., L.G. Klikoff, and K.T. Harper. 

1976. Differential resource utilization by the 

sexes of dioecious plants. Science 193: 597-599. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular plants of the United States, Canada, and 

Greenland. Vol. II—thesaurus . 2nd ed. Portland, 

OR: Timber Press. 816 p. 

Kitchen, S.G., S. Walker, and G.R. Wilson. 1999. 

Addition of Ephedra nevadensis—Nevada 

ephedra to the Rules. The Seed Technologist 

Newsletter 73: proposal #9. 

Luke, F. and S.B. Monsen. 1984. Methods and 

costs for establishing shrubs on mined lands in 

southwestern Wyoming. In: A.R. Tiedemann, 

E.D. McArthur, H.C. Stutz, R. Stevens, and K.L. 

Johnson, comps. Proceedings—Symposium on 

the biology of Atriplex and related chenopods; 

1983 May 2-6; Provo, UT. General Technical 

Repeport INT-172. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Forest and Range Experiment Station, Ogden, 

UT. p. 286-291. 

McLaughlin, S.P. and J.E. Bowers. 1982. Effects 

of wildfire on a Sonoran Desert plant 

community. Ecology 63: 246-248. 

Plummer, A.P. 1977. Revegetation of disturbed 

Intermountain area sites. In: J.C. Thames, ed. 

Reclamation and use of disturbed lands of the 

Southwest. University of Arizona Press, Tucson, 

AZ. p. 302-337. 

Plummer, A.P., D.R. Christensen, and S.B. 

Monsen. 1968. Restoring big-game range in 

Utah. Publication 68-3. Utah Division of Fish 

and Game, Ephraim, UT. 183 p. 

Shaw, N. and S.B. Monsen. 1984. Nursery 

propagation and outplanting of bareroot 

chenopod seedlings. In: A.R. Tiedemann, E.D. 

McArthur, H.C. Stutz, R. Stevens, and K.L. 

Johnson, comps. Proceedings—Symposium on 

the biology of Atriplex and related chenopods; 

1983 May 2-6; Provo, UT. General Technical 

Report INT-172. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Forest and Range Experiment Station, Ogden, 

UT. p. 251-260. 

Smith, A.D. and D.M. Beale. 1980. Pronghorn 

antelope in Utah: some research and 

observations. Publication 80-13. Utah Division 

of Wildlife Resources, Salt Lake City, UT. 88 p. 

Stanton, F. 1974. Wildlife guidelines for range fire 

rehabilitation. Technical Note 6712. U.S. 

Department of the Interior, Bureau of Land 

Management, Denver, CO. 90 p. 

Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981. 

Viability of seed from thirty-two shrub and forb 

species through fifteen years of warehouse 

storage. Great Basin Naturalist 41: 274-277. 

Stevensen, D.W. 1993. Ephedraceae. In: Flora of 

North America. http://flora.huh.harvard.edu/ 

[not paged]. 

Steward, J.H. 1938. Basin-Plateau Aboriginal 

Sociopolitical Groups. Bulletin 120. 

Smithsonian Institution Bureau of American 

Ethnology, Washington, DC. [1997 reprinted by 

the University of Utah Press, Salt Lake City, 

UT]. 346 p. 

Tirmenstein, D. 1990. Ephedra nevadensis. In: 

U.S. Department of Agriculture, Forest Service, 

Fire Sciences Laboratory. Fire Effects 

Information System, http://www.fs.fed. 

us/database/feis/ [not paged]. 

Turner, F.B. and D.C. Randall. 1987. The 

phenology of dessert shrubs in southern Nevada. 

Journal of Arid Environments 13: 119-128. 

West, N.E. and M.A. Hassan. 1985. Recovery of 

sagebrush-grass vegetation following wildfire. 

Journal of Range Management 38: 131-134. 

Wright, H.A. 1980. The role and use of fire in the 

semidesert grass-shrub type. General Technical 

Report INT-85. U.S. Department of Agriculture, 

Forest Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT: 24 p. 

Young, J.A. and R.A. Evans. 1978. Population 

dynamics after wildfires in sagebrush 

grasslands. Journal of Range Management 31: 

283-289. 

Young, J.A., R.A. Evans, and B.L. Kay. 1977. 

Ephedra seed germination. Agronomy Journal 

69: 209-211. 
310 

Ephedra viridis Coville green ephedra 

EPHEDRACEAE 

Synonyms: Ephedra nevadensis subvar. pluribracteata Palmer ex Stapf. 

 Ephedra nevadensis var. viridis (Colville) M.E. Jones 

Stanley G. Kitchen 

General Description.—Green ephedra, also 

known as Mormon tea, or jointfir, is a spreading to 

erect, coniferous shrub, 0.5 to 1.5 m tall and with a 

crown 1 to 3 m wide at maturity (Cronquist and 

others 1972, USDA Forest Service 1976). A 

fibrous, somewhat fragile root system originating 

from several, almost parallel, taproots descends 

deep into the soil profile. Stems are jointed and 

grooved. Internodes are 1 to 4.5 cm long 

(Cronquist and others 1972). Lower bark is gray 

and irregularly fissured (Anderson 2001). 

Photosynthetic upper branches are thin (1 to 3 

mm), bright green to yellow green and broom-like, 

with opposite or falsely whorled attachment 

(Welsh and others 1987). Leaves are reduced to 

1.5 to 4 mm, deciduous scales, positioned in 

opposite pairs at nodes. A thickened brown base 

remains after leaf-shed. Egg-shaped cones are 

borne at nodes, singly or whirled. Two or more 

male cones, 5 to 7 mm long, are subtended by 

opposite membranous bracts, 2 to 4 mm long, 

while female cones, 6 to 10 mm long, are partially 

enclosed by thickened bracts, 4 to 7 mm long 

(Cronquist and others 1972). Brown, three-sided 

seeds are smooth and 5 to 8 mm long and 2 to 4 

mm wide. The combination of small cones, ring 

porous wood, wide multiseriate rays, and vessels 

in older stems separate Ephedra from other 

gymnosperms (Stevenson 1993). 

Taxonomy.—Two varieties of green ephedra are 

recognized. Variety viscida (Cutler) L. Benson 

differs from the more widespread var. viridis by 

the presence of short stalks (peduncles) on female 

cones and the often sticky nature of stems (Welsh 

and others 1987). Chromosome number is 2n = 28 

(Stevensen 1993). 

Range.—Green ephedra is closely associated with 

the Great Basin and Colorado Plateau provinces of 

western North America (Anderson 2001). Its range 

extends from California and Oregon east and south 

to western Texas and Chihuahua, Mexico. In Utah 

and California its elevational range varies from 

900 to 3,000 m (Welsh and others 1987, Anderson 

2001). 

Ecology.—Green ephedra is found on 

sedimentary, volcanic, and metamorphic substrates 

in sandy, gravelly, or rocky, well-drained, 

underdeveloped soils (Anderson 2001). 

Populations are found on a variety of landforms 

including: plains, alluvial fans, mesa tops, washes, 

and canyon walls on all aspects. Salt tolerance is 

limited. Plants tolerate partial shade but do best in 

full sun. Although rarely considered a dominant, 

green ephedra is associated with a wide variety of 

plant community types including: big sagebrush-
311

bunchgrass, black sagebrush, blackbrush, desert 

grassland, pinyon-juniper, mountain mahogany, 

oak woodland, mountain shrub, mixed conifer, and 

ponderosa pine (Anderson 2001). It is present in 

early, mid, and late successional stages (Young 

and Evans 1973, Koniak 1985). Mean annual 

precipitation varies from 200 to 430 mm 

(Anderson 2001). Plants sprout from crown or 

roots following top-kill by fire (USDA Forest 

Service 1976, Young and Evans 1978, Koniak 

1985, Everett 1987). Reported survival following 

fire is variable (Anderson 2001). Sprouting 

response following fire may vary by season of 

burn, fire severity, or among population variation 

in fire tolerance. Green ephedra is found in 

communities for which a wide range of fire 

frequencies is known. It also establishes on 

disturbed sites from seed (Koniak 1985). Green 

ephedra is used as winter forage by big game and 

livestock but receives only limited use during 

other seasons (Jameson and others 1962, Kufeld 

and others 1973). Palatability and nutritive quality 

of young stems are generally ranked as fair to 

good for most wildlife species and classes of 

domestic livestock (Dittberner and Olson 1983, 

Anderson 2001). Seeds are eaten by birds and 

rodents and appear to be moderately prefered 

(Everett and others 1978). Rodents probably play 

an important role in seed dispersal. 

Reproduction.—Green ephedra is dioecious. 

Spatial segregation of the sexes results from high 

male plant representation on dry ridges and slopes 

and high female plant representation on microsites 

with relatively favorable soil moisture conditions 

(Freeman and others 1976, Brunt and others 1988) 

providing for greater reproductive efficiency. 

Plants are wind-pollinated. Timing of cone 

maturation, or coning, and seed maturation is 

dependent upon variation in weather, elevation, 

aspect, and geography. Coning and pollination 

occur for approximately 2 weeks, usually between 

April and June. Seeds mature between June and 

September. Cones are not produced each year, and 

good seed crops may be several years apart 

(Plummer and others 1968). There are 

approximately 45,000 seeds per kg (Meyer and 

others 1988). 

Growth and Management.—In years of good 

seed crops, significant quantities are easily 

harvested by hand (Plummer and others 1968). 

Ripened cones are easily dislodged by gently 

shaking or striking cone-bearing branches over 

tarps or hand-held hoppers. Timing of harvest is 

critical as thunder storms can cause considerable 

shattering after seeds ripen. After drying, seeds are 

cleaned to high purity and viability levels using 

conventional milling equipment. Freshly-harvested 

seed may be partially dormant (Meyer and others 

1988). Dormancy can be broken with a 4-week 

moist prechill or stratification (Meyer and others 

1988). Germination for after-ripened or prechilled 

seed is relatively rapid. Germination of nondormant seeds occurs over a range of cool 

temperatures (Young and others 1977). Seed 

viability remains high for at least 5, and perhaps as 

long as 15 years (Stevens and others 1981). 

Seedling growth rate is slow. Seedlings may 

require 5 to 10 years to reach 60 cm in height 

(Anderson 2001). In restoration plantings, 

seedlings are susceptible to competition from 

weeds and faster-growing grasses and forbs in the 

seed mix (Ferguson and Frischknecht 1981). 

Plants establish well from direct seeding and 

transplants (USDA Forest Service 1976, Plummer 

1977, Everett 1980, Luke and Monsen 1984). Stem 

cutting are capable of root development in 

greenhouse conditions (Wieland and others 1971). 

Green ephedra was toxic to domestic cattle and 

sheep during pregnancy in a controlled diet study 

(Keeler 1989). No detrimental effects were 

observed for developing calves and lambs. 

Conflicting reports have been published on the 

possible presence of ephedrine alkaloids in green 

ephedra. While Anderson (2001) suggests the 

presence of this group of compounds in this 

species, Caveney and others (2001) failed to find 

these compounds common to Old World ephedras 

in any New World species. Tannin content is high. 

Benefits.—Green ephedra is valuable for restoring 

vegetation structure and diversity and for erosion 

control on disturbed lands, including roadsides, 

minelands, and degraded rangelands (USDA 

Forest Service 1976, Everett 1980, Ferguson and 

Frischknecht 1981, Ferguson and Frischknecht 

1985, Herbel 1986), in areas in which it is adapted. 

When competition is controlled, it establishes well 

from seed and persists after wildfire. It provides 

important winter forage for wildlife and domestic 

livestock (USDA Forest Service 1976). Green 

ephedra has had limited use as an ornamental 

(Anderson 2001). Native Americans made both 

medicinal and nonmedicinal teas brewed from 

stems and used seeds and roasted seeds (nutlets) 

for food (Steward 1938, USDA Forest Service 

1976). Mormon settlers were also known to use it 

in the production of a beverage, a cause for the 

frequently used moniker, ‘mormon tea’. 

Undetermined medicinal applications for a group 

of unique secondary compounds associated with 
312 

members of the genus have not been fully 

determined, including possible antimicrobial 

applications for kynurenates (Caveney and others 

2001). 

References

Anderson, M.D. 2001. Ephedra viridis. In: U.S. 

Department of Agriculture, Forest Service, Fire 

Sciences Laboratory, Fire Effects Information 

System, http://www.fs.fed.us/database/feis/ 

26 p. 

Brunt, J.W., M.R. Conley, and G.L. Cunningham. 

1988. Sex in Ephedra trifurca (Ephedraceae) 

with relation to Chihuahuan Desert habitats. The 

American Midland Naturalist. 119: 137-142. 

Caveney, S., D.A. Charlet, H. Freitag, M. MaierStolte, and A.N. Starratt. 2001. New 

observations on the secondary chemistry of 

world Ephedra (Ephedraceae). American 

Journal of Botany 88: 1199-1208. 

Cronquist, A., A.H. Holmgren, N.H. Holmgren, 

and J.L. Reveal. 1972. Intermountain flora: 

Vascular plants of the Intermountain West, 

U.S.A. Vol. 1. Hafner Publishing Company, 

Inc., New York. 270 p. 

Dittberner, P.L. and M.R. Olson. 1983. The plant 

information network (PIN) data base: Colorado, 

Montana, North Dakota, Utah, and Wyoming. 

FWS/OSB-83/86. U.S. Department of the 

Interior, Fish and Wildlife Service, Washington 

DC. 786 p. 

Everett, R.L. 1980. Use of containerized shrubs for 

revegetating arid roadcuts. Reclamation Review 

3: 33-40. 

Everett, R.L. 1987. Plant response to fire in the 

pinyon-juniper zone. In: Everett, R.L., comp. 

Proceedings—pinyon-juniper conference; 1986 

January 13-16; Reno, NV. General Technical 

Report INT-215. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Research Station, Ogden, UT. p. 152-157. 

Everett, R.L., R.O. Meeuwig, and R Stevens. 

1978. Deer mouse preference for seed of 

commonly planted species, indigenous weed 

seed, and sacrifice foods. Journal of Range 

Management. 31: 70-73. 

Ferguson, R.B. and N. Frischknecht. 1981. Shrub 

establishment on reconstructed soils in semiarid 

areas. In: Shrub establishment on disturbed arid 

and semi-arid lands: Proceedings of the 

symposium; 1980 December 2-3; Laramie, WY. 

Wyoming Game and Fish Department. p. 57-63. 

Ferguson, R.B. and N. Frischknecht. 1985. 

Reclamation on Utah’s Emery and Alton coal 

fields: techniques and plant materials. Research 

Paper INT-335. U.S. Department of Agriculture, 

Forest Service, Intermountain Research Station, 

Ogden, UT. p. 78. 

Freeman, D.C., L.G. Klikoff, and K.T. Harper. 

1976. Differential resource utilization by the 

sexes of dioecious plants. Science 193: 597-599. 

Herbel, C.H. 1986. Seeding shrubs in the field. 

Reclamation and Revegetation Research 5: 377-

385. 

Jameson, D.A., J.A. Williams, and E.W. Wilton. 

1962. Vegetation and soils of Fishtail Mesa, 

Arizona. Ecology 43: 403-410. 

Keeler, R.F. 1989. Investigation of maternal and 

embryo/fetal toxicity of Ephedra viridis and 

Ephedra nevadensis in sheep and cattle. Journal 

of Range Management 42: 31-35. 

Koniak, S. 1985. Succession in pinyon-juniper 

woodlands following wildfire in the Great 

Basin. The Great Basin Naturalist 45: 556-566. 

Kufeld, R.C., O.C. Wallmo, and C. Feddema. 

1973. Foods of the Rocky Mountain mule deer. 

Research Paper RM-111. U.S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Forest and Range Experiment Station, Fort 

Collins, CO. 31 p. 

Luke, F. and S.B. Monsen. 1984. Methods and 

costs for establishing shrubs on mined lands in 

southwestern Wyoming. In: A.R. Tiedemann, 

E.D. McArthur, H.C. Stutz, R. Stevens, and K.L. 

Johnson, comps. Proceedings—Symposium on 

the biology of Atriplex and related chenopods; 

1983 May 2-6; Provo, UT. Gen. Tech. Rep. 

INT-172. U.S. Department of Agriculture, 

Forest Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. p. 286-291. 

Meyer, S.E., S.G. Kitchen, G.R. Wilson, and R. 

Stevens. 1988. Addition of Ephedra viridis

green mormon tea to the Rules. AOSA 

Newsletter 62: 18-19. 
313

Plummer, A.P. 1977. Revegetation of disturbed 

Intermountain area sites. In: J.C. Thames, (ed). 

Reclamation and use of disturbed lands of the 

Southwest. University of Arizona Press, Tucson, 

AZ. p. 302-337. 

Plummer, A.P., D.R. Christensen, and S.B. 

Monsen. 1968. Restoring big-game range in 

Utah. Publ. No. 68-3. Utah Division of Fish and 

Game, Ephraim, UT. p. 183. 

Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981. 

Viability of seed from thirty-two shrub and forb 

species through fifteen years of warehouse 

storage. Great Basin Naturalist 41: 274-277. 

Stevensen, D.W. 1993. Ephedraceae. In: Flora of 

North America. http://flora.huh.harvard.edu/ 

[not paged]. 

Steward, J.H. 1938. Basin-Plateau Aboriginal 

Sociopolitical Groups. Bulletin 120. 

Smithsonian Institution Bureau of American 

Ethnology, U.S. Government Printing Office, 

Washington, DC. [1997 reprinted by the 

University of Utah Press, Salt Lake City, UT]. 

346 p. 

USDA Forest Service. 1976. Some important 

native shrubs of the west. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Forest and Range Experiment Station. Ogden, 

UT. 16 p. 

Welsh, S.L., N.D. Atwood, L.C. Higgins, and S. 

Goodrich. 1987. A Utah Flora. Brigham Young 

University Print Services, Provo, UT. 894 p. 

Wieland, P.A.T., E.F. Frolich, and A. Wallace. 

1971. Vegetative propagation of woody shrub 

species from the northern Mojave and southern 

Great Basin deserts. Madrono 21: 149-152. 

Young, J.A. and R.A. Evans. 1973. Downy bromeintruder in the plant succession of big sagebrush 

communities in the Great Basin. Journal of 

Range Management 26: 410-415. 

Young, J.A. and R.A. Evans. 1978. Population 

dynamics after wildfires in sagebrush 

grasslands. Journal of Range Management 31: 

283-289. 

Young, J.A., R.A. Evans, and B.L. Kay. 1977. 

Ephedra seed germination. Agronomy Journal 

69: 209-211. 
314 

Eriogonum fasciculatum Benth. California buckwheat

POLYGONACEAE 

Synonyms: Eriogonum rosmarinifolium Nutt. 

Eriogonum fasciculatum var. maritimum Parish 

Eriogonum fasciculatum var. oleifolium Gand. 

Eriogonum aspalathoides Gand. 

Eriogonum fasciculatum ssp. aspalathoides S. Stokes 

Arlee M. Montalvo 

General Description.—California buckwheat is a 

widespread and morphologically variable shrub 

and is one of the most important components of 

sagebrush scrub, desert scrub, and especially 

coastal sage scrub (Kirkpatrick and Hutchinson 

1977, Westman 1981). Although the four 

taxonomic varieties partly overlap in geographic 

distribution each is associated with a different 

climatic zone and combination of traits, especially 

color, chromosome number, leaf shape, degree of 

revolute leaf margins, and pubescence (Munz and 

Keck 1968, Hickman 1993). All forms have 

alternate cauline, linear to oblanceolate leaves that 

are white wooley with appressed hairs beneath, 

and darker, generally less hairy to glabrous above. 

Leaves are 1 to 2 cm long and cluster tightly at 

nodes on short axillary shoots, giving the leaves a 

fascicled appearance. Flowers are small (about 3 

mm long) with six white to pinkish sepals and are 

born in heads of tightly to loosely packed 

involucres with few flowers per involucre. The 

heads (or dense cymes) are born singly or in 

umbels subtended by long, naked peduncles (often 

> 5 cm). Branching of the umbels varies from one 

to four levels. Umbels are rounded to flat-topped 

depending on the extent of branching. Flowers 

produce small (about 1.5 to 2 mm long), single 

seeded achenes that are partially enclosed in a dry, 

persistent calyx. Below is a summary of diagnostic 

differences among varieties following Munz and 

Keck (1968), Reveal (1989), and Hickman (1993). 

Range and Taxonomy.—E. f. var. fasciculatum

(see insert) has 2n = 40 chromosomes (Munz and 

Keck 1968). Shrubs are often low and spreading 

and 0.6 to 1.2 m tall. Leaves are linear to linearoblanceolate, green and glabrate above, sparsely 

pubescent beneath, with leaf margins strongly 

rolled under (revolute). Involucres and flowers are 

usually glabrous. Plants occur from nearly sea 

level to just over 300 m in coastal scrub and sage 

scrub along coastal bluffs and cliffs, and in 

canyons and dry slopes of the foothills near the 

coast from San Luis Obispo Co. south to central 

Baja California. 

E. f. var. foliolosum (Nutt.) S. Stokes ex 

Jones (Munz and Keck 1968), the interior flattopped buckwheat, is tetraploid with 2n = 80 

chromosomes (Stebbins 1942). Shrubs are erect to 

spreading, 0.5-1.5 m tall and occur on interior 

slopes and mesas below 1000 m (seldom on coast) 

in chaparral and sage scrub from Monterey Co., 

California south to northern Baja California. 

Leaves are often linear, dark to gray-green and 

pubescent above, and densely pubescent below. 

Involucres and perianth are usually pubescent. 

Synonyms are E. rosmarinifolium var. foliolosum 

Nutt., E. f. ssp. foliolosum S. Stokes, and E. f. var. 

obtusiflorum S. Stokes. 
315

Eriogonum f. var. polifolium (Benth.) Torrey 

& A. Gray (flowering shoot in photograph) has 2n 

= 40 chromosomes (Stebbins 1942). These 

greyish, pubescent, rounded shrubs occur at 600 to 

2300 m on dry slopes and washes. They are 

widespread across inland mountains and deserts of 

California, from the inner coast ranges of San Luis 

Obispo and Fresno Cos., south to San Diego Co., 

eastward into the Mojave Desert to southern 

Nevada, Utah, and Arizona, and south into 

northwestern Senora, Mexico and central Baja 

California. In inland sage scrub, plants overlap in 

distribution with var. foliolosum, especially in 

Riverside Co., but occupy drier sites. Leaves are 

oblanceolate to obovate, densely pubescent 

beneath, and usually tomentose above with leaf 

margins sometimes barely revolute. Involucres and 

sepals are pubescent. Heads tend to be born in 

simple to once-compound umbels. Reveal (1989) 

reports hybridization between var. polifolium and 

E. cinereum Benth in areas were E. cinereum has 

been introduced within the range of var. 

polifolium. Synonyms are E. f. ssp. p. S. Stokes 

and E. p. Benth. 

Eriogonum f. var. flavoviride Munz & 

Johnson has n = 20 chromosomes (Reveal 1989) 

and is 2 to 3 dm tall. The rounded shrubs occur on 

dry rocky slopes and washes from 200 to 1300 m 

in creosote bush scrub of the southern Mojave 

Desert, San Bernardino Co., south through eastern 

Riverside Co. to far eastern San Diego Co., and 

south to central Baja California. Leaves are light 

green to yellowish-green, linear, strongly revolute, 

and glabrous above. Peduncles are glabrous; 

involucres and sepals are subglabrous. Synonym is 

E. f. ssp. flavoviride S. Stokes. 

Ecology.—California buckwheat occurs on dry 

slopes and often granitic substrate (Westman 

1981). The woody, branched roots penetrate to 

under 1.5 m (Kummerow and others 1977). Plants 

lose some leaves in the dry season and can shed 

many leaves in severe drought. Plants are 

vulnerable to hot fires so resprout success is low 

and most regeneration is from seeds (Keeley 

1998). Frequent fires deplete the seed bank, 

making populations vulnerable to local extinction. 

In Riverside Co., this shrub has decreased 

dramatically since the 1940’s, with the largest 

decreases on alluvial soils (Minnich and Dezzani 

1998). Decreases are attributed to a combination of 

competition with exotic annual grasses, too 

frequent a fire interval, that is exacerbated by 

weeds which carry fire, and by atmospheric 

deposition of nitrogen, which encourages growth 

of competing weeds. California buckwheat does 

well on rocky road cuts and in shallow soils that 

are inhospitable to annual grasses. 

Reproduction.—Plants flower from early spring 

through summer. The four taxa have slightly 

shifted flowering times, with var. flavoviridae

having the earliest season (Munz and Keck 1968) 

and var. polifolium flowering before the other two. 

In all taxa, stamens and stigmas are exerted and 

open to generalist pollinators, including honey 

bees and many species of small native bees, flies, 

wasps, and beetles. Achenes mature from June to 

September and disperse when dried inflorescences 

shatter in autumn. The dry calyx provides 

buoyancy to detached achenes and assists dispersal 

by wind and water. Harvester ants also disperse 

achenes (DeSimone and Zedler 1999). Seedlings 

emerge in the rainy season in mid to late winter 

within light gaps and open areas. Survival is 

unusually high as shown by studies in a variety of 

sites and years, often ranging from 80 to 90 

percent (Wright and Howe 1987, Miriti and others 

1998, Montalvo and others 2002). Most shoot 

elongation and leaf growth occurs before 

flowering from late winter to mid spring (Cole 

1967). Plants can reach maturity in 1 to 2 years. 

Seed Germination.—Light improves germination 

(Cole 1967, Keeley 1987). Atwater (1980) 

suggests that light increases permeability of seed 

coats, water adsorption, and possibly leaching of 

inhibitors, allowing seeds to germinate. Lightinduced seed coat permeability would allow some 

buried seeds to remain viable and dormant in the 

seed bank, a bet-hedging strategy. Cole (1967) 

found coastal populations germinated at 15 to 25 

o

C while inland populations germinated over a 

larger range of 5 to 35 o

C. Keeley (1987) 

examined the effects of light, dark, heat, and 

leachate through charred wood (charrate) on 

germination. Seeds germinated equally well on 

soil vs. filter paper, and light controls had a twofold increase in germination compared to dark 

controls (83 vs. 45 percent germination). Charrate 

did not affect germination. In light, seeds exposed 

to 120 o

C for 5 min suffered a large decrease in 

germination under all conditions relative to seeds 

heated to 70 or 100 o

C, suggesting fire may kill 

many seeds in the upper centimeter of the 

seedbank. Heat sensitivity and a light requirement 

for germination encourage colonization of 

disturbed or open sites. 

Horticulture.—Plants can be propagated from 

seeds or rooted from cuttings. Seeds and container 

plants should be planted in the fall to early winter 
316 

to take advantage of the moist cool season for 

growth. In one study, seeds planted by seed 

imprinting and hydroseeding, shallow methods 

that allow light to reach planted seeds, had at least 

twice the emergence success as drilling that 

covered seeds with 0.5 to 1 cm of soil (Montalvo 

and others 2002). Seeds germinate within 2 weeks 

if soil is moist and over about 15 o

C. Plants require 

full sun and good drainage. No irrigation is 

required if planted before seasonal rains. 

The grey-green var. polifolium provides a 

nice contrast to the other green varieties in the 

garden, but all do well in dry landscaping. The 

flowers last far into the summer and become an 

attractive rusty-brown in fruit. Some prostrate 

cultivars of California exist, including “Theodore 

Payne,” “Prostrata,” “Dana Point,” and 

“Wildwood” (Perry 1992). 

Greenhouse studies indicate that California 

buckwheat forms facultative associations with 

arbuscular mycorrhizal fungi (Egerton-Warburton, 

Montalvo, and Allen, unpublished report). The 

plant’s facultative dependence on mycorrhizal 

fungi may be instrumental to successful 

colonization of barren sites. Montalvo and others 

(2002) found that plants grew quickly to maturity 

on a graded site with low organic matter and 

nutrients, but establishment, growth, and flowering 

all decreased significantly with decreasing soil 

nitrogen (NO3-N mean = 7.89 mg/g, range = 0.8-

69 mg/g in the study plot). 

Benefits.—California buckwheat provides habitat 

and food for numerous animals. Osborne (1998) 

found high abundance and diversity of arthropod 

species on California buckwheat. Some are 

species-specific feeders such as the larvae of the 

moth, Hemileuca electra (Wright) (Rubinoff 

1998). Numerous species of butterfly larvae feed 

on California buckwheat and specialize on 

different plant parts and taxa (Howe 1975), 

sometimes according to timing of growth. For 

example, the larvae of the butterfly, Apodemia 

mormo (Felder & Felder), has three biotypes in the 

Mojave Desert. The spring morph feeds on 

California buckwheat while the others feed on 

species with different flowering times. E. 

parvifolium Sm. in Rees. is the host of the rare El 

Segundo blue butterfly (Euphilotes bernardino 

allyni Shields). When California buckwheat was 

planted instead of later flowering E. parvifolium, 

competing insect species used California 

buckwheat, built up populations, and then 

competed with the rare butterfly for E. parvifolium

flowers (Longcore and others 2000). 

 Vertebrates also use California 

buckwheat. The rare orange-throated whiptail is 

associated with open vegetation with this shrub 

(Brattstrom 2000). Rare California gnatcatchers 

(Polioptila californica californica) forage and nest 

in the shrub. California buckwheat and Artemisia 

californica Less. are dominant shrubs used by 

gnatcatchers (Mock and Bolger 1992). The rare 

mountain sheep (Ovis canadensis nelsoni) browses 

on California buckwheat in the San Gabriel 

Mountains of southern California (Perry and others 

1987), and deer eat the inflorescences 

(Schopmeyer 1974). 

Genetics, Geographic Variation, Hybridization, 

and Fitness.—Stebbins (1942) hypothesized that 

tetraploid populations of var. foliolosum arose 

from hybridization between diploid var. 

fasiculatum and var. polifolium (= E. f. subsp. 

foliolosum, E. f. subsp. typicum, and E. f. subsp. 

polifolium, respectively, in the paper). He 

examined the distribution of taxa in relation to 

climatic factors and concluded that the 

morphology and ecological tolerances of var. 

foliolosum combined the morphology and 

tolerances of the two diploid taxa. The variety 

polifolium was in areas with some snow, frost, and 

only 10 to 13 cm of rainfall, fasciculatum was in 

the areas with no frost, often frequent fog, and 22 

to 37 cm of rainfall, and foliolosum often occurred 

in habitats intermediate to the two diploids. Cole 

(1967) tested assumptions about the distribution of 

morphology and ecological tolerances of vars. 

fasciculatum and foliolosum. Cole compared their 

physiologies over an ecological gradient from the 

cool coast across the mountains to the hot inland 

valleys and found that coastal var. fasciculatum

were the least hairy, and populations of var. 

foliolosum became more hairy inland. There was a 

clear morphological cline from the coast to the 

interior among and within ecotypes that correlated 

with habitat. Photosynthesis vs respiration ratios 

and photosynthetic rates and respiration rates all 

varied in a clinal manner that correlated with 

morphological characters. The variety

fasciculatum forms a distinctly adaptive coastal 

ecotype, while var. foliolosum forms an inland 

ecotype. 

Analysis of eight allozyme loci for eight 

populations of var. foliolosum and four 

fasciculatum revealed ample genetic variation 

(Montalvo, Ellstrand, and Clegg, unpublished 

data). All loci were polymorphic and there was an 

average of 5.7 alleles/locus. Expected 

heterozygosity was also high at 0.42. Nei’s genetic 

distances among populations ranged from 0 to 
317

0.074, with the largest distance between varieties. 

Overall, the proportion of total variation explained 

by differences among populations (GST) was 

0.025. By itself, the tetraploid var. foliolosum had 

no significant structure (GST = 0.005), while GST = 

0.028 for var. fasciculatum. These values suggest 

high levels of gene flow and outbreeding.

Seed Collection and Processing.—Seeds are 

collected after heads have turned rusty brown. The 

chaff can be separated from seeds by pushing 

seeds through a screen and then separating seeds 

from chaff with an air separator or fan (author’s 

observation). Commercial seed companies usually 

skip the screening and just break up the chaff, air 

separate the seeds from crude chaff, and sell seeds 

with the persistent calyx intact. A recommended 

target for minimum quality of purchased seeds is 

15 percent purity and 65 percent germination. 

Here, a bulk seed pound with about 990,000 

seeds/kg, would have about 96,800 pure live seeds 

(personal communication, S & S Seeds, 

Carpenteria, CA). 

Growth and Management.—Seeds have been 

widely planted for restoration, roadside erosion 

control, slope stabilization, landscaping, and 

apiary (Schopmeyer 1974, Perry 1992). Despite 

the clear geographic pattern and ecological 

affinities of the four taxonomic varieties, most 

researchers and consultants fail to designate 

taxonomic variety on publications, plant lists, or 

plant palettes. They also commonly fail to specify 

appropriate ecological zone for seeds used in 

wildland plantings. This has resulted in extensive 

plantings outside natural ranges and habitats 

(Reveal 1989). Projects should use seeds of 

appropriate taxa to maximize project success. 

References 

Atwater, B.R. 1980. Germination, dormancy and 

morphology of the seeds of herbaceous 

ornamental plants. Seed Science and Technology 

8: 523-573. 

Brattstrom, B.H. 2000. The range, habitat 

requirements, and abundance of the orangethroated whiptail, Cnemidophorus hyperythrus

beldingi. Bulletin of the Southern California 

Academy of Sciences 99: 1-24. 

Cole, N.H. A. 1967. Comparative physiological 

ecology of the genus Eriogonum in the Santa 

Monica Mountains, southern California. 

Ecological Monographs 37: 1-24. 

DeSimone, S.A., and P.H. Zedler. 1999. Shrub 

seedling recruitment in unburned Californian 

coastal sage scrub and adjacent grassland. 

Ecology 80: 2018-2032. 

Hickman, J.C., ed. 1993. The Jepson Manual: 

Higher Plants of California. University of 

California Press, Ltd., Los Angeles, CA. 1,400 p. 

Howe, W.H., ed. 1975. The butterflies of North 

America. Doubleday and Company, Inc., Garden 

City, NY. 633 p. 

Keeley, J.E. 1987. Role of fire in seed germination 

of woody taxa in California chaparral. Ecology 

68: 434-443. 

Keeley, J.E. 1998. Postfire ecosystem recovery and 

management: The October 1993 large fire 

episode in California. In: J. M. Moreno (ed.). 

Large Forest Fires. Backbuys Publishers, Leiden, 

Netherlands. p. 69-90. 

Kirkpatrick, J.B., and C.F. Hutchinson. 1977. The 

community composition of California coastal 

sage scrub. Vegetatio 35: 21-33. 

Kummerow, J., D. Krause, and W. Jow. 1977. Root 

systems of chaparral shrubs. Oecologia 29: 163-

177. 

Longcore, T., R. Mattoni, G. Pratt, and C. Rich. 

2000. On the perils of ecological restoration: 

lessons from the El Segundo blue butterfly. In: J. 

E. Keeley, M. Baer-Keeley and C. J. 

Fotheringham, eds. 2nd Interface Between 

Ecology and Land Development in California: 

U.S. Geological Survey Open-File Report 00-62. 

Vol. 62. Sacramento, CA. p. 281-286. 

Minnich, R.A., and R.J. Dezzani. 1998. Historical 

decline of coastal sage scrub in the RiversidePerris plane, California. Western Birds 29: 366-

391. 

Miriti, M.N., H.F. Howe, and S.J. Wright. 1998. 

Spatial patterns of mortality in a Colorado desert 

plant community. Plant Ecology 136: 41-51. 

Mock, P.J., and D. Bolger. 1992. Ecology of the 

California gnatcatcher at Rancho San Diego. 

Ogden Environmental and Energy Services Co., 

Inc.; Project No. 110970000. 50 p. 

Montalvo, A.M., P.A. McMillan, and E.B. Allen. 

2002. The relative importance of seeding method, 
318 

soil ripping, and soil variables on seeding success. 

Restoration Ecology 10: 52-67. 

Munz, P.A., and D.D. Keck. 1968. A California 

Flora with Supplement. University of California 

Press, Berkeley, CA. 1681 + 224 p. 

Osborne, K.H. 1998. A description of arthropod 

community structure in southern California 

coastal sage scrub. M.S. Thesis. University of 

California, Riverside, CA. 133 p. 

Perry, B. 1992. Landscape plants for western 

regions: an illustrated guide to plants for water 

conservation. Land Design Publishing, 

Claremont, CA. 318 p. 

Perry, W.M., J.W. Dole, and S.A. Holl. 1987. 

Analysis of the diets of mountain sheep from the 

San Gabriel Mountains, California. California 

Fish and Game 73: 156-162. 

Reveal, J.L. 1989. The eriogonoid flora of 

California (Polygonaceae: Eriogonoideae). 

Phytologia 66: 295-414. 

Rubinoff, D. 1998. Field observations on mating 

behavior and predation of Hemileuca electra

(Saturniidae). Journal of the Lepidopterists' 

Society 52: 212-214. 

Schopmeyer, C.S., ed. 1974. Seeds of Woody Plants 

in the United States. Vol. 450 Agricultural 

Handbook. USDA Forest Service, Washington, 

D.C. 883 p. 

Stebbins, G.L. 1942. Polyploid complexes in 

relation to ecology and the history of floras. The 

American Naturalist 76: 36-45. 

Westman, W.E. 1981. Factors influencing the 

distribution of species of California coastal sage 

scrub. Ecology 62: 439-455. 

Wright, S.J., and H.F. Howe. 1987. Pattern and 

mortality on Colorado Desert plants. Oecologia 

73:543-552. 
319

Erithalis fruticosa L. black torch 

RUBIACEAE 

Synonyms: Erithalis revoluta Urban 

 Erithalis angustifolia DC. 

 Erithalis elliptica Raf. 

 Erithalis inodora Jacq. 

 Erithalis odorata Pers. 

 Erithalis parviflora Griseb. 

John K. Francis 

 

General Description.—Black torch, also known as 

candlewood, tea, parrot apple, cubra prieta, 

jayajabico, manglillo, tarro de chivo, bois 

flambeau, and bois chandelle, is an evergreen shrub 

that typically is 1 or 2 m in height but occasionally 

reaches 8 m in height. Usually the species has 

multiple stems arising near the ground line and 

many branches that are stiff and sometimes 

gnarled. The bark is gray and smoothish to warty. 

The resinous wood is brown, hard, fine-textured, 

heavy, and has visible growth rings. Black torch 

plants are supported by brown roots that are often 

contorted as they grow over rocks and into cracks. 

The foliage may be dense or diffuse depending on 

whether the shrubs grow in full sun or partial 

shade. The opposite leaves have 3 to 20 mm 

petioles, orbicular to oblanciolate blades 2 to 12 cm 

long with entire edges, and dark-green upper 

surfaces. The inflorescences are lateral or terminal 

panicles containing many small, white flowers. The 

black fruits are globose or flattened drupes 3 to 4 

mm in diameter that have a bitter-sweet flavor and 

contain five to 10 nutlets (Howard 1989, Liogier 

1997, Little and others 1974, Nelson 1996). It is 

morphologically a highly variable species (Long 

and Lakela 1976). 

Range.—Black torch is native to Florida, the 

Bahamas, the Greater and Lesser Antilles, 

Trinidad, Margarita and other Venezuelan islands, 

Quintana Roo (Mexico), and Belize (Howard 1989, 

Little and others 1974). 

Ecology.—Black torch grows from near sea level 

to 120 m in elevation in Puerto Rico (Little and 

others 1974) in areas that receive from 750 mm to 

about 1800 mm of annual precipitation. The 

species is drought-hardy and moderately intolerant 

of shade and can grow under the canopy of lowdensity forest. Because of slow growth, it is 

restricted to areas with reduced competition. Black 

torch is most common near beaches and on rock 

outcroppings and bluffs near the shoreline, 

especially in moist limestone areas, and on 

limestone hills in the interior. In Florida, black 

torch is found in beach strand vegetation, on sand 

dunes, and coastal hammocks (Nelson 1996). 

Reproduction.—Black torch flowers and fruits 

throughout the year (Little and others 1974, Nelson 

1996). Open-grown individuals produce fruits and 

seeds in abundance. Fruits collected in Puerto Rico 

averaged 0.1087 + 0.0071 g/fruit. Air-dried seeds 

separated from them averaged 0.00092 g/seed or 

1,080,000 seeds/kg. Presumably, the seeds are 

dispersed by birds that eat the fruits. Seedlings and 

saplings are relatively common in Puerto Rico. The 

stems layer (root) when they come in contact with 

the ground. 

Growth and Management.—Black torch is slow 
320 

growing, especially when situated on rock 

outcrops. Management should mainly consist of 

protection from fires and clearing of sites for 

development. 

Benefits.—Black torch contributes to biodiversity 

in the forests where it grows, helps protect the soil, 

and furnishes food and cover for wildlife. It would 

probably make an acceptable ornamental for green 

areas and nonirrigated gardens. The wood is good 

for turned articles but available only in small sizes. 

It is resistant to rot and has been used for piles and 

posts. Formerly, it was split and used for torches 

(Little and others 1974) but tended to be smoky 

(University of the Virgin Islands 2002). The bark, 

fruits, and the resin have diuretic and astringent 

properties and are used to treat inflammation of the 

kidney and bladder, and blennorrhoea (Liogier 

1990). The leaves are parched and ground and used 

to treat skin sores (University of the Virgin Islands 

2002). 

References 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

 Liogier, H.A. 1990. Plantas medicinales de 

Puerto Rico y del Caribe. Iberoamericana de 

Edicones, Inc., San Juan, PR. 566 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyan Books, Miami, FL. 

962 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

390 p. 

University of the Virgin Islands. 2002. Erithalis 

fruticoas (blacktorch). http://rps.uvi.edu/ 

VIMAS/blacktorch.htm. 1 p.
321

Ernodea littoralis Sw. golden creeper 

RUBIACEAE 

Synonyms: Ernodea angusta Small 

John K. Francis 

General Description.—Golden creeper, also 

known as cough bush, beach creeper, wild 

pomegranate, ernodia de playa, wild pamgramma, 

and liane-séche, is an evergreen prostrate or 

arching shrub to about 1.5 m in height. Heavy 

lateral roots support the plants. The stems are fourangled and orange or reddish brown. Golden 

creeper leaves have a short petiole (1 to 2 mm), 

lanceolate blades, and a spiny, pointed tip. In some 

environments, leaves are somewhat fleshy; in 

others they are hard or coriaceous. White-to-pink 

tubular flowers are solitary, axillary, and sessile. 

The fleshy fruits are golden orange, in the shape of 

tiny pomegranates, and have an apple-like flavor. 

The seeds, borne one per fruit, are brown with a 

longitudinal furrow (Howard 1989, Liogier 1997, 

Nelson 1996). 

Range.—Golden creeper is native to Southern 

Florida, the Bahamas, the Greater Antilles, the 

Lesser Antilles as far south as Marie Galante, 

Mexico, Belize, and Honduras (Howard 1989, 

Liogier 1997). It is planted as an ornamental but is 

not known to have naturalized into any new range 

as a result. 

Ecology.—Golden creeper will grow in acidic to 

mildly alkaline soils with the whole range of 

textures (Gilman 1999) and over a variety of 

parent materials including serpentine. In Puerto 

Rico, it grows in areas receiving from 750 to about 

1400 mm of mean annual precipitation and at 

elevations from near sea level to about 400 m. 

Golden creeper is tolerant of salt spray and mild 

salt concentration in the soil. The species is 

drought-tolerant and usually grows in excessively 

drained sites. However, it withstands short-term 

flooding (Carney-Kulig and others 2002). In 

Florida, it is common in coastal dunes (Nelson 

1996). The variety angusta grows in pinelands in 

Florida (Workman 1980). In Puerto Rico, golden 

creeper also grows on coastal rocks (usually 

limestone) and on rocky ultramafic (serpentine) 

slopes. It is a sun-loving plant that usually grows 

in the open. When the shrubs grow under low trees 

in open forest, they do not become thick and 

robust. Golden creeper benefits from mild 

disturbance that eliminates shade and taller 

competition. 

Reproduction.—Golden creeper blooms and 

fruits all year (Gilman 1999, Nelson 1996). Fruits 

collected in Puerto Rico weighed an average of 

0.0467 + 0.0014 g/fruit. Air-dried seeds separated 

from them weighed an average of 0.0078 + 0.0002 

g/seed or 128,000 seeds/kg. Sown without 

pretreatment on commercial potting mix, 24 

percent germinated between 35 and 146 days of 

sowing (author’s observation). The fruits are 

presumably eaten and the seeds dispersed by birds. 

Once established, plants thicken and spread by 

root suckering and layering. 

Growth and Management.—The growth rate of 

golden creeper is reported to be moderate (Gilman 

1999). Individual stems appear to live about 2 to 5 

years, but the plants, being clonal, can go on 

indefinitely. Golden creeper is produced and sold 

in commercial nurseries. Plants in commercial 

production are produced by ground layering and 

rooted cuttings. Field planting of hormone-treated 

cuttings is also recommended (Centro Ecologico 

Sian Ka’an 2002). Wildlings (seedlings and plants 

dug up from natural stands) transplant well 

(Workman 1980). Recommended spacing for 

planting is 0.9 to 1.5 m (Gilman 1999). Once 

planted, golden creeper needs little attention 

(Workman 1980); in fact, it often dies if overirrigated (Gilman 1999). 
322 

Benefits.—Golden creeper contributes to the 

biodiversity of the sites where it grows, stabilizes 

the soils in those fragile environments, and 

provides food and cover for wildlife. It is planted 

as an ornamental for landscaping, usually as 

ground cover. Golden creeper is also used for 

beach dune stabilization. It is particularly 

promoted for water-restricted landscaping 

(Workman 1980). A tea is made from the leafy 

shoots as a treatment for coughs (Centro Ecologico 

Sian Ka’an 2002). 

References 

Carney-Kulig, S., M. Jolly, T. Weyant, S. Buehler, 

and G. Davis. 2002. Planting the right plant in 

the right place. Monroe County Extension 

Service, University of Florida. http:// 

monroe.ifas.ufl.edu/gardeningguide_5.htm. 6 p. 

Centro Ecologico Sian Ka’an. 2002. Flora. 

http://cesiak.org/flora1.htm. 3 p. 

Gilman, E.F. 1999. Ernodea littoralis. Fact Sheet 

FPS-196. Cooperative Extension Service, 

University of Florida. http:// 

hort.ifas.ufl.edu/shrubs/ERNLITA.pdf. 3 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Dicotyledoneae. 

Part 3. Vol. 6. Arnold Arboretum, Harvard 

University, Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Workman, R.W. 1980. Growing native. The 

Sanibel-Captiva Conservation Foundation, Inc., 

Sanibel, FL. 137 p. 
323

Erythroxylum areolatum L. false cocaine 

ERYTHROXYLACEAE 

Synonyms: none (genus also spelled Erythroxylon) 

John K. Francis 

General Description.—False cocaine, also known 

as redwood, swamp redwood, thin-leafed 

erythroxylon, indio, palo de hierro, arabo 

carbonero, limoncillo, huesito, cocaina falsa, and 

poirier, is a deciduous shrub or small tree 2 to 7 m 

in height and 5 to 15 cm in stem diameter. The 

plant may have a single stem or multiple stems 

branching from near the ground and is supported 

by a tap and lateral root system. The roots are dark 

brown, stiff, and woody. The bark is gray and scaly 

and the inner bark is red. Moderately robust 

branches and many twigs form a “branchy” crown, 

often with a dense display of foliage. Alternate 

leaves are tightly grouped on short side branches or 

spread out on new twigs. The leaves are light green 

to dark green, elliptic to oblong, 3 to 13 cm long, 

with entire margins and petioles 5 to 7 mm long. 

Tiny, fragrant, five-merous white flowers are borne 

in clusters in the leaf axils. At maturity, the thinfleshed, 6- to 9-mm drupes are bright red. Each 

fruit contains one oblong, bony seed (Britton and 

Millspaugh 1962, Liogier 1988, Little and 

Wadsworth 1964, Stevens and others 2001). 

Range.—False cocaine is native to the Bahamas, 

the Greater Antilles, the Cayman Islands, southern 

Mexico, and Central America (National Trust for 

the Cayman Islands 2002, Stevens and others 

2001). It is not known to have been planted or 

naturalized elsewhere. 

Ecology.—False cocaine grows in areas of Puerto 

Rico that receive from about 750 to 900 mm of 

mean annual precipitation at elevations of a few 

meters above sea level to about 300 m. It grows in 

gallery forests in Nicaragua from 40 to 380 m 

elevation, frequently associated with limestone 

rocks (Stevens and others 2001). False cocaine is 

common in limited areas but uncommon in most of 

its range, growing in remnant and middle to late 

secondary forests. False cocaine grows on deep, 

medium-textured soil and sandy beach-strand soils 

(Vásquez and Kolterman 1998). The species is 

most frequent on limestone parent material, as 

skeletal rock or porous solid rocks but grows in 

areas with igneous and metamorphic (including 

ultramaphic) rocks. It has an intermediate tolerance 

to shade and will grow in openings or in the 

understory of medium to low basal area forests. 

Reproduction.—False cocaine has been observed 

flowering from October to June in Puerto Rico 

(Little and Wadsworth 1964). Fresh fruits collected 

in Puerto Rico weighed an average of 0.119 +

0.013 g/fruit. Seeds cleaned from them weighed 

(air-dried) an average of 0.0493 g/seed or 20,300 

seeds/kg. Sown on moist filter paper without 

pretreatment, they germinated at 35 percent, 

beginning 12 days after sowing (Francis and 

Rodríguez 1993). Apparently, birds are the 

principal dispersers of seeds. Seed production can 

be abundant in some years. Seedlings and saplings 

vary from rare to common. 

Growth and Management.—False cocaine grows 

slowly, about 1 to 3 mm in diameter per year and is 

capable of living several decades. Published 

planting or management experience is not known 

to the author. Protecting forests and stands 

containing the species is probably the best 

management approach until research can be 
324 

performed. 

Benefits.—False cocaine protects the soil and 

furnishes food and cover for wildlife. The wood is 

heavy, hard, fine-grained, durable, and strong. The 

sapwood is light brown and the heartwood is 

reddish or chocolate brown (Little and Wadsworth 

1964). Available only in small sizes, it is useful for 

stakes, fuel, carving, and turnery. The foliage of 

this species does not contain useful amounts of 

cocaine, which is extracted from a sister species (E. 

coca Lam.) from the Andean Highlands (Little and 

Wadsworth 1964). 

References 

Britton, N.L. and C.F. Millspaugh. 1962. The 

Bahama flora. The New York Botanical Garden, 

New York. 695 p. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: Second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

National Trust for the Cayman Islands. 2002. 

Virtual herbarium of the National Trust for the 

Cayman Islands: Erythroxylaceae, Erythroxylum 

areolatum L. http://www.virtualherbarium.org/ 

vh/CAYM/403.htm. [not paged]. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany, Vol. 85, No. 

1. Missouri Botanical Garden, St. Louis, MO. 

943 p. 

Vásquez, O.J. and D.A. Kolterman. 1998. Floristic 

composition and vegetation types of the Punta 

Guaniquilla Natural Reserve—Cabo Rojo, 

Puerto Rico. Caribbean Journal of Science 

43(304): 265-279. 
325

Erythroxylum brevipes DC. brisselet 

ERYTHROXYLACEAE 

Synonyms: Erythroxylum brevipedatum St. Lager 

 Erythroxylum rhamnoides Peyr. ex Schulz 

John K. Francis 

General Description.—Brisselet is a shrub, 

usually 1 or 3 m in height, that sometimes 

becomes a small tree to 8 m in height. It is also 

known by the common names, rat-wood, jibá, 

ratón, yaría de costa and rocío in Spanish and bois 

vinette and brésillette in French (Liogier 1988, 

Little and others 1974). Older plants often have 

multiple stems from below or just above the 

ground level. They are supported by a tap and 

lateral root system consisting of dark-brown, 

tough, and brittle roots. The shrub’s crowns are 

cluttered with slender branches and twigs. The 

bark of brisselet is gray or light brown. Foliage on 

older branches is borne on short spurs. The 1- to 3-

cm obovate leaves have prominent light-colored 

central nerves and are sometimes notched at the 

tips. The small, white, five-petaled flowers are 

clustered one to four at the leaf axils. Brisselet 

fruits are red, ellipsoidal, fleshy drupes 5 to 9 mm 

long (Howard 1988, Little 1988, Little and others 

1974). 

Range.—Brisselet is native to Hispaniola, Puerto 

Rico, the Virgin Islands, and St. Barts in the West 

Indies (Howard 1988). It has been much confused 

in the literature with E. rotundifolium Luman that 

also occurs in Puerto Rico (Liogier and Martorell 

2000), southern Mexico through Central America 

as far south as Costa Rica, and in the Bahamas, 

Cuba and Jamaica (Stevens and others 2001). 

Ecology.—Brisselet grows in most types of soil, 

except poorly drained or salty soils. Plants are 

more frequent in eroded areas and rocky terrain 

where competition is weaker. Annual rainfall in the 

Puerto Rican range varies from about 750 to about 

2000 mm. Elevation where it grows ranges from 

near sea level to about 760 m (Little and others 

1974). Brisselet is moderately intolerant of shade. 

It invades abandoned pastures, fields, and disturbed 

areas, usually as scattered plants. It is common in 

early and mid-secondary forest and is an important 

plant of the understory of remnant and late 

secondary forests that are not too dense. Grazing 

cattle benefits the species, probably by reducing its 

competition. Cut stems will resprout. No 

observations are published on sprouting after single 

fires, but repeated fires will eliminate it from an 

area. 

Reproduction.—Brisselet flowers during the 

summer months and fruits in late summer or early 

fall (Little and others 1974). Fruit production can 

be heavy in open-grown plants but is rare in shaded 

understory plants. A collection of fruits from 

Puerto Rico weighed an average of 0.0936 +

0.0035 g/fruit. Air-dried seeds separated from them 

weighed an average of 0.0401 + 0.0007 g/seed or 

24,900 seeds/kg. Sown in commercial potting mix 

without any pretreatment, these seeds began 

germinating in 19 days and completed 99 percent 

germination in 47 days. Germination is epigenous. 

A thin, deep taproot is quickly produced. The 

seedlings are woody, stiff, and easy to lift and 

transplant. Birds disperse the seeds. Seedlings are 

relatively common and widespread. 

Growth and Management.—Both height and 

diameter growth of brisselet is slow. Weaver 

(1990) measured the mean annual diameter growth 
326 

of 33 brisselet stems 4 to 14 cm in diameter in St. 

John, U.S. Virgin Islands at 0.5 mm/year. 

Apparently, plants live for several decades. No 

management experience has been published. 

Benefits.—Brisselet is an important component of 

dry and moist forests, protects the soil, and 

furnishes cover for wildlife. Although seasonal, the 

fruits are a consistent source of food for birds. Bees 

are attracted to the fragrant flowers (Little and 

others 1974). The sapwood is light tan while the 

heartwood is dark brown, hard, and flexible. 

Because of its small diameters and lengths, the 

wood is useful for little other than fuel. Brisselet 

tissues do not contain cocaine, as obtained from its 

more famous relative Erythroxylum coca Lam. 

References 

Howard, R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. and L.F. Martorell. 2000. Flora of 

Puerto Rico and adjacent islands, a systematic 

synopsis. 2nd Ed. Editorial de la Universidad de 

Puerto Rico, Río Piedras, PR. 382 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.H. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs of Systematic Botany Vol. 85, No. 

1. Missouri Botanical Garden Press. p. 1-943. 

Weaver, P.L. 1990. Tree diameter growth rates in 

Cinnamon Bay Watershed, St. John, U.S. Virgin 

Islands. Caribbean Journal of Science 26(1-2): 

1-6.
327

Eugenia foetida Pers. boxleaf stopper 

MYRTACEAE 

Synonyms: Eugenia buxifolia (Sw.) Willd. 

 Eugenia myrtoides Poir. 

 Myrtus axillaries Poiret in Lam. 

 Myrtus poiretii Sprengl 

 Eugenia triplinervia O. Berg. 

 Eugenia lateriflora sensu Griseb. 

John K. Francis 

General Description.—Boxleaf stopper is also 

known as Spanish stopper, gurgeon stopper, 

stopper bush, white watting, anguila, hoja menuda, 

escobón, pico de paloma, guairaje, bois petites 

feuilles. It is an upright, evergreen shrub or small 

tree sometimes reaching 10 m in height and 12.5 

cm in diameter at breast height. There is usually a 

single stem from the ground, but it may branch 

near the ground. The stems and branches are 

slender with gray, smooth to lightly fissured or 

platy bark. The inner bark is pink and slightly 

bitter. The sapwood is light brown, and the 

heartwood is dark reddish brown. The wood is 

hard, heavy, and fine-grained. This deep-rooted 

species has a tap and lateral root system of slender, 

flexible to moderately stiff brown roots. The 

paired, leathery, green to dark green leaves are 

obovate or oblanciolate 2 to 5 cm long and have a 

1.5 to 3 mm petiole and usually have a rounded 

tip. The foliage, especially when crushed, emits a 

mild, offensive, skunk-like odor. Small, fourpetaled, white flowers are grouped in fewflowered, axillary racemes. Fruits are black 

globose berries 4 to 7 mm in diameter with one or 

two small seeds (Howard 1989, Liogier 1994, 

Little and others 1974). 

Range.—Boxleaf stopper is native to Florida, 

Puerto Rico, the U.S. Virgin Islands, Jamaica, 

Hispaniola, Cuba, Anguilla, St. Martin, St. Barts, 

Netherlands Antillies, Yucatan in Mexico, Belize, 

and Guatemala (Howard 1989, Liogier 1994, Little 

and others 1974). The species has not been 

reported to have naturalized outside its native 

range. 

Ecology.—Boxleaf stopper most frequently grows 

in areas underlain by limestone rock, usually on 

ridges and hill sides. It grows on clay and coursertextured soils, rocky rubble, and often directly into 

cracks in fractured rock. The species also occurs 

on sandy flats behind beaches. In Florida, it grows 

on hammocks near the coast and on pinelands in 

the Lower Keys (Nelson 1996). Elevations vary 

from near sea level to about 100 m. Mean annual 

rainfall of the native range in Puerto Rico ranges 

from about 750 and 1700 mm (author’s 

observation, Little and others 1974). The species is 

drought tolerant (Gilman 1999). Although boxleaf 

stopper eventually reaches the stature of a small 

tree on favorable sites, much of its habitat is so 

difficult that the plant completes its life cycle in 

sizes (2 to 5 m in height) usually associated with 

shrubs. The species is moderately intolerant of 

shade, growing in the understory, as an 

intermediate or codominant in low-basal area, 

short-statured, dry forests. 
328 

Reproduction.—Flowering of boxleaf stopper in 

Puerto Rico is reported to occur in spring and 

summer (Little and others 1974). It flowers in the 

summer in Florida (Gilman 1999). Fresh fruits 

collected in Puerto Rico averaged 0.181 + 0.032. 

Air-dried seeds separated from them averaged 

14,900 seeds/kg. Placed in moist potting mix, 51 

percent germinated beginning 69 days after 

sowing. The new seedlings grow slowly. In the 

wild, seeds are dispersed by birds. Seedlings and 

saplings are widely scattered to relatively 

common. 

Growth and Management.—The growth rate of 

boxleaf stopper is moderate to slow, depending on 

habitat. Apparently, it is relatively long lived (a 

few decades). Potted stock is available from 

nurseries for planting. Although expensive for 

wildland plantings, this may be the best way to 

establish them in order to get them above 

competing weeds and grass. Ornamental plants 

may be pruned into hedges or forced to singlestemmed trees (Gilman 1999). Because they are 

relatively slow growing and not aggressive 

invaders, control of the species in natural stands 

should not be necessary. 

Benefits.—Boxleaf stopper protects the soil, 

furnishes cover for wildlife, and contributes to the 

aesthetics of the forest. Aromatic compounds 

released from the leaves of this species apparently 

are responsible for the slight musky smell present 

in forests in Puerto Rico where it is common. 

Foliage of the species is not eaten by Key deer 

(Odocolleus virginanus clavium) (Schaus and 

others 2003) and probably is avoided by other 

ruminant herbivores. Although small, the fruits are 

edible to humans, and birds consume them when 

available. Boxleaf stopper is recommended and 

used for screens, hedges, parking lot, yard, and 

street trees in Florida (Gilman 1999). 

References

Gilman, E.F. 1999. Eugenia foetida. Fact Sheet 

FPS-200. Cooperative Extension Service, 

University of Florida, Gainesville, FL. 3 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Schaus, C., S. Wade and J. Dunan. 2003. Key deer 

and plants they won’t eat. http://monroe. 

ifas.ufl.edu/key_deer_plants.htm. 4 p. 
329

Eugenia pseudopsidium Jacq. wild guava 

MYRTACEAE 

Synonyms: Eugenia megalocarpa Urban 

 Eugenia portoricensis DC. 

 Eugenia thomasiana Berg 

 Myrtus willdenovii Spreng. 

Stenocalyx portoricensis Berg 

John K. Francis 

General Description.—Wild guava, which is also 

known as bastard guava, Christmas cherry, 

guayaba silvestre, quiebrahacha, goyavier de 

montague, bois plié, and goyavier bâtard, is a 

shrub or a small tree reaching a maximum of 20 m 

in height and 30 cm in trunk diameter (Howard 

1989, Liogier 1994, Little and others 1974). 

Mature plants are usually 3 to 5 m in height and 4 

to 8 cm in stem diameter. Wild guava has a tough, 

woody root system with tap and lateral roots. The 

stems are light gray, slightly fissured, and scaly. 

The wood is light to reddish brown, hard and very 

heavy (specific gravity = 1.3). Twigs are green, 

turning to brown (Little and others 1974). The 

crowns branch to the third order and branches tend 

to be slender. The opposite leaves have short 

petioles, are generally elliptic, rounded or short 

pointed at the base and long pointed at the apex. 

They are dark green above and light green 

beneath. Small white flowers arise in the leaf 

axils, one to three per axil. The globose fruits 

retain the calyx of the flower. They are orange, 

turning to bright red, and astringent with little 

flavor. Each fruit contains one rounded seed. 

Range.—The native range of wild guava includes 

Hispaniola, Puerto Rico, the Virgin Islands, 

Guadalupe, Dominica, Martinique, and St. Lucia. 

The same plant or a very similar one also occurs in 

Venezuela, the Guyanas, and Amapá, Brazil 

(Howard 1989). That it is wild guava is supported 

by its listing in the Smithsonian (2001) checklist 

of plants for Guyana. 

Ecology.—Wild guava grows inland a few meters 

above sea level to elevations of 670 m in Puerto 

Rico (Little and others 1974). It occurs in areas 

that receive from about 1400 to 2200 mm of 

annual precipitation. Medium- to heavy-textured 

soils derived from sedimentary, igneous, and 

metamorphic (including ultramafic) rocks are 

colonized. It does not appear sensitive to 

topographic position but rarely if ever grows on 

excessively or very poorly drained soils. Wild 

guava is shade tolerant and normally is confined to 

the understories of remnant old growth and 

advanced secondary forests. 

Reproduction.—Wild guava flowers and fruits 

irregularly throughout the year (Little and others 

1974). Understory plants produce fruits in small 

numbers; plants receiving increased sunlight in 

gaps or thinned canopies produce several times 

more fruits. Wild guava fruits collected in Puerto 

Rico weighed an average of 1.284 + 0.043 g. 

Seeds extracted from these fruits weighed an 

average of 0.565 + 0.010 g (air dry) or 1,800 

seeds/kg. Eighty-one percent of the seeds from 

this collection germinated between 59 and 143 

days after sowing in commercial potting mix. 

When the new seedlings are damaged above 

ground by fungi or insects, they sprout from the 
330 

ground level, usually with more than one shoot 

(author’s observation). Birds disperse the seeds. 

Seedlings and saplings are well-scattered and 

relatively common. Wild guava sprouts readily 

after cutting or damage. 

Growth and Management.—The growth rate of 

wild guava appears to be relatively good for an 

understory shrub. A 2.5-m tall sapling, 2.5 cm 

d.b.h., under partial shade in Puerto Rico, had five 

growth rings in its trunk near the ground. Wild 

guava germinates, survives, and transplants well in 

the nursery, but growth is somewhat slower than 

most commercial nursery plants. Unfortunately, 

no plantation or ornamental use has been reported. 

Benefits.—Wild guava is used to a limited extent 

for firewood and fence posts. With its dark green 

leaves, bright red fruits, and clean form, the 

species has a pleasing appearance and probably 

would make a fine ornamental, especially in shady 

locations. It provides food and cover for wildlife 

and contributes to the scenic beauty and 

biodiversity of the forests where it grows. 

References 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. U.S. Department of 

Agriculture, Agriculture Handbook 449. 

Washington, DC. 1,024 p. 

Smithsonian. 2001. Regional plant species lists for 

Guyana. Mason.gmu.edu/~ckelloff/region2.pdf. 

6 p. 
331

Euonymus alatus (Thunb.) Sieb. winged burning bush 

CELASTRACEAE 

Synonyms: Celastrus alatus Thunb. 

 Celastrus striatus Thunb. 

 Euonymus arakianus Koidz. 

 Euonymus sacrosanctus Koidz. 

 Euonymus striatus (Thunb.) Loes. 

 Euonymus subtriflorus Blume 

 Euonymus thunbergianus Blume 

Jinshuang Ma and Gerry Moore 

Illustration soruce: Ma, J.S. 2001

General Description.—This species is commonly 

known as winged burning bush or winged spindle 

tree because of its stems, which have two to four 

conspicuous brown, corky ridges or wings, 2 to 3.5 

mm wide and 1.0 to 1.2 mm thick. It is the only 

species of Euonymus L. in North America to have 

such prominent wings on its twigs. It is also 

occasionally called Asian spindle tree because it is 

native to Asia. Winged burning bush is a lowgrowing shrub, usually up to 2 m tall, never more 

than 4 m. The plant does spread well horizontally, 

sending out stems in many directions. Its buds are 

axillary (two per axil), ovoid or obovate, scattered 

along the stem. The leaves are opposite, two per 

node, and are spaced somewhat evenly along the 

stem. The leaf petioles are short, round in cross 

section, glabrous, 2 to 3.5 mm long (rarely shorter 

than 2 mm., the leaves then appearing subsessile). 

The leaf blades are glabrous, lanceolate to obovate, 

4.5 to 7.3 (rarely to 10) by 1.2 to 2.3 (rarely to 4.5) 

cm, leathery to papery, with cuneate or attenuate 

bases, crenate to serrate margins and acuminate to 

cuspidate apices. The five to seven pairs of 

ascending lateral veins disappear before reaching 

the leaf margin. The flowers develop on the 

current season’s growth and are are bisexual 

(rarely functionally unisexual). They are in small 

axillary clusters (cymes), the clusters on slender 

peduncles, 2.0-2.5 cm long, usually with one 

dichotomous branch and three or fewer flowers. 

Each flower is on a pedicel 4 to 7 mm long. The 

flowers are about 9 mm in diameter with the parts 

in fours, and the sepals and petals readily 

distinguishable from one another. The petals are 

yellowish green. The fruits are ovoid, four-lobed 

capsules, 8 to 14 by 10 to 16 mm. At maturity the 

capsules are red to brown-purple and they split 

open along the lobes (often one or more lobe is 

abortive), exposing the one to four seeds, each 

enclosed in a fleshy orange aril. The seeds (sans 

aril) are glabrous, light brown to black, ellipsoid, 6 

to 12 by 4.0 to 5.5 mm. Some spell the specific 

epithet “alata” instead of “alatus” used here. This 

is because botanists disagree as to whether the 

genus name Euonymus should be treated as 

feminine (“alata”) or masculine (“alatus”) (Paclt 

1998). In classical Greek, the generic name was 

treated as feminine; however, Linnaeus treated the 

genus as masculine. The spelling “Euonymus” has 

formally been conserved over Linnaeus’s 

originally spelling “Evonymus,” which has also 

occasionally been used (Gilstra 1991). 

Range.—Winged burning bush is native to east 

Asia being distributed in far eastern Russia (Far 

East), Japan, Korea, and China. In its native range, 

it occurs in areas with annual precipitation of 400 
332 

mm or more and in elevations ranging from 40 to 

2,700 m (Ma 2001). It was first introduced into the 

United States about 1860 (Rehder 1940), as an 

ornamental plant, and it eventually became 

naturalized in the Eastern United States (Ebinger 

1996, Gleason and Cronquist 1991, J. Ma in 

personal observation). Currently winged burning 

bush is known from 21 Eastern and Mid-Western 

States (Connecticut, Illinois, Indiana, Iowa, 

Kentucky, Maryland, Massachusetts, Michigan, 

Minnesota, Missouri, North Carolina, New 

Hampshire, New Jersey, New York, Ohio, 

Pennsylvania, Rhode Island, South Carolina, 

Virginia, Vermont, West Virginia), as well as 

Montana (USDA 2003). It can be expected to be 

found elsewhere with the continued spread of 

naturalized populations and additional material 

escaping from cultivation. 

Ecology.—It is not as aggressive an invader as the 

related Oriental bittersweet (Celastrus orbiculatus

Thunb). It is found in disturbed habitats, such as 

roadsides, forest margins and woodlands that are 

in early stages of succession (Ma, personal 

observation). 

Reproduction.—Hymenopterous and dipterous 

insects, especially bees and flies, appear to be the 

main pollinators, and the attractive arillate seeds 

are widely dispersed by birds and small mammals 

(Brizicky 1964). The flowers bloom in the spring 

and the fruit mature in the fall. The species can be 

asexually propagated through stem cuttings and 

root separation. 

Growth and Management.—Winged burning 

bush grows in a variety of soil conditions. It is not 

a fast growing plant, reaching perhaps up to 1 m of 

growth in a single growing season. Propagation 

can be effected by layering or seed (Chittenden 

and Synge 1956). However, there is concern about 

the continued use of this plant in ornamental 

plantings, since it can be invasive in some areas 

(Martin 2002) 

Benefit.—The foliage of winged burning bush 

turns bright scarlet in autumn, giving the plant its 

common name burning bush. The purplish 

capsules and seeds with orange arils add to the 

“burning” effect of the plant late in the year. This 

effect of the plant has made it a popular 

ornamental and horticultural species. Some 

specimens can be shaped into bonsai (Smith 2003). 

References 

Brizicky, G.K. 1964. The genera of Celastrales in 

the southeastern United States. Journal of the 

Arnold Arboretum. 45: 210-215. 

Chittenden, F.J. and P.M. Synge. 1956. Dictionary 

of gardening. Vol. 2, 2nd Ed. Clarendon Press, 

Oxford, UK. 583 p. 

Ebinger, J.E. 1996. Euonymus alatus. In: J.M. 

Randall, J.M. and J. Marinelli, eds. Invasive 

plants–weeds of the global garden. 21st Century 

Gardening Series. Brooklyn Botanic Garden, 

Brooklyn, NY. p 55. 

Gilstra, Z. 1991. Proposal to conserve the spelling 

of 4618 Euonymus (Celastraceae). Taxon 40: 

137-139. 

Gleason, H.A. and A. Cronquist 1991. Manual of 

Vascular Plants of northeastern United States 

and adjacent Canada. New York Botanical 

Garden, Bronx, NY. 910 p. 

Ma, J.S. 2001. A revision of Euonymus 

(Celastraceae). Thaiszia Journal of Botany 11: 

1-264. 

Martin, T. 2002. Weed alert! Euonymus alata 

(Thunb.) Siebold. http://tncweeds.ucdavis. 

edu/alert/alrteuon.html. [not paged]. 

Paclt, J. 1998. Proposal to amend the gender of 

Euonymus, nom. cons. (Celastraceae) to 

feminine. Taxon 47: 473-474. 

Rehder, A. 1940. Manual of cultivated trees and 

shrubs. Macmillan Publishing Co., New York. 

996 p. 

Smith, B. 2003. The bonsai guide 

http://www.thebonsaiguide.com/BPG/BS_BPBT

E.HTM. [not paged]. 

USDA, NRCS. 2003. The PLANTS Database. 

Baton Rouge, LA. http:plants.usda.gov. [not 

paged].
333

Euonymus americanus L. strawberry bush 

CELASTRACEAE 

Synonyms: Euonymus alternifolius Moench. 

 Euonymus americanus var. augustifolius (Pursh) Wood 

 Euonymus americanus var. sarmentosus Nutt. 

 Euonymus augustifolius Pursh 

 Euonymus heterophylus Raf.

 Euonymus muricatus Raf. 

 Euonymus sarmentosus G. Don 

 Euonymus sempervirens Marsh. 

Jinshuang Ma and Gerry Moore 

Illustration source: Britton and Brown 1913 

General Description.—Strawberry bush is also 

known as bursting heart, hearts–a-bursting, or 

American burning bush. It is a low-growing, 

deciduous shrub, up to 2 m high. Its stems are stiff, 

green, glabrous, straight, round, 2 to 5 mm in 

diameter, and are occasionally four-angled. The 

pith is white, round, and continuous. The leaves of 

the strawberry bush are opposite (two per node), 

spaced somewhat evenly along stem. The petioles 

are short, less than 3 mm and sometimes are nearly 

absent, the leaves then appearing sessile or 

subsessile. The leaf blades are slightly leathery to 

papery, oblong to elliptic, 4.0 to 10 by 2.0 to 3.5 

cm wide, with cuneate or attenuate bases, crenulate 

margins, and acuminate apices. The five to seven 

pairs of ascending lateral leaf veins disappear 

before reaching the margin. The flowers occur on 

the current season’s growth in small axillary 

clusters (cymes), each cluster usually consisting of 

two peduncles, each with one to three flowers 

(often one or two of the flowers will aborte, 

leaving only one functional flower per peduncle). 

The peduncles are slender, 1.5 to 2.2 cm long. 

When there is more than one flower per peduncle 

each flower is subtended by a short pedicel, 1 to 

3.5 mm long. The flowers are bisexual, about 10 

mm in diameter, with the sepals and petals readily 

distinguishable from one another. The calyx 

comprises five yellow to green sepals, 1.0 to 1.5 

by 1.6 to 1.8 mm. The corolla comprises of five 

greenish to reddish petals up to 4 mm by 4 mm 

wide. There are five stamens, 1.5 to 2.0 mm long. 

The five carpels are fused. A sticky nectarous disk 

is present, 3 to 4 mm in diameter. The color and 

shape of the fruit gives it its common names. The 

fruit is a plump, red globose dry, five-lobed 

capsule up to 1.5 cm in diameter prior to opening. 

The surface of the capsule is densely covered with 

minute spines or warts on the surface. This species 

is the only species of Euonymus L. in North 

America to have a capsule with a spiny or warty 

surface. The capsule opens along the lobes, 

exposing the seeds that are covered by a fleshy, 

scarlet aril. The seeds (sans aril) are brown, 

glabrous, ellipsoid, 7 to 9 by 4.0 to 5.5 mm. Some 

spell the specific epithet “americana” instead of 

“americanus” used here. This is because botanists 

disagree as to whether the genus name Euonymus 

should be treated as feminine (“americana”) or 

masculine (“americanus”) (Paclt 1998). In 
334 

classical Greek, the generic name was treated as 

feminine; however, Linnaeus treated the genus as 

masculine. The spelling “Euonymus” has formally 

been conserved over Linnaeus’s originally spelling 

“Evonymus,” which has also occasionally been 

used (Gilstra 1991) 

Range.—Strawberry bush occurs throughout 

much of the Eastern United States (Alabama, 

Arkansas, Delaware, District of Columbia, Florida, 

Georgia, Illinois, Indiana, Kentucky, Louisiana, 

Maryland, Mississippi, Missouri, New Jersey, 

New York, North Carolina, Ohio, Oklahoma, 

Pennsylvania, South Carolina, Tennessee, Texas, 

Virginia and West Virginia), except New England. 

(Shetler and Orli 2000, USDA 2003) It is also 

known from Ontario (Gleason and Cronquist 1991, 

Ma 2001, Scoggan 1978). 

Ecology.—Strawberry bush grows well in many 

soils types, up to 1 m in a growing season. It is 

usually found in forests and thickets, including 

lowland areas and edges of streams. It occurs from 

sea level up to 400 m in elevation (Ma 2001). 

Reproduction.—Hymenopterous insects, 

especially bees, appear to be the main pollinators, 

although wind may also be involved (Brizicky 

1964). The seeds of strawberry bush are dispersed 

by birds and other small mammals (Brizicky 

1964). Strawberry bush blooms in late spring and 

early summer, and the fruits mature in autumn. 

Growth and Management.—Strawberry bush 

prefers a humus-rich, slightly acidic soil. It does 

well in shady situations, tolerating even full shade. 

It can be planted in USDA Zones 5 to 9. 

Propagation can be effected through stem cuttings, 

layering and seed (Chittenden and Synge 1956) 

Seeds require approximately 3 months of 

prechilling and stratification. 

Benefits.—Strawberry bush is an excellent 

ornamental species. The leaves turn a dark red in 

autumn. The red capsules and scarlet arillate seeds 

add to the “burning effect” and contributes to some 

of its common names. Strawberry bush is best 

used in naturalistic settings, in the shade of larger 

shrubs and trees. Large specimens can have 

hundreds of the “bursting” red capsules. In the 

winter, the bright green twigs are also handsome. 

References 

Britton, N.L. and A. Brown. 1913. Illustrated flora 

of the northern states, Canada, and the British 

possessions. Vol. 2, 2nd Ed. Scribner, New York. 

735 p. 

Brizicky, G.K. 1964. The genera of Celastralesin 

the Southeastern United States. Journal of the 

Arnold Arboretum. 45: 206-235. 

Chittenden, F.J. and P.M. Synge. 1956. Dictionary 

of gardening. Vol. 2, 2nd ed. Clarendon Press, 

Oxford, UK. 583 p. 

Gilstra, Z. 1991. Proposal to conserve the spelling 

of 4618 Euonymus (Celastraceae). Taxon 40: 

137-139. 

Gleason, H.A., and A. Cronquist 1991. Manual of 

vascular plants of northeastern United States and 

adjacent Canada. 2nd Ed. New York Botanical 

Garden, Bronx, NY. 910 p. 

Ma, J.S. 2001. A Revision of Euonymus 

(Celastraceae). Thaiszia, Journal of Botany 11: 

1-264. 

Paclt, J. 1998. Proposal to amend the gender of 

Euonymus, nom. cons. (Celastraceae) to 

feminine. Taxon 47: 473-474. 

Scoggan, H.J. 1978. The Flora of Canada Part 3 – 

Dicotyledoneae (Saururaceae to Violaceae). 

National Museum of Natural Sciences, National 

Museums of Canada, Ottawa. 568 p. 

Shetler, S.G. and S.S. Orli. 2000. Annotated 

checklist of the vascular plants of the 

Washington-Baltimore area. Part. 1. Ferns, fern 

allies, gymnosperms, dicotyledons. Smithsonian 

Institution, Washington, D.C. 186 p. 

USDA, NRCS. 2003. The PLANTS Database. 

Baton Rouge, LA. http:plants.usda.gov. [not 

paged].
335

Euonymus atropurpureus Jacq. eastern wahoo 

CELASTRACEAE 

Synonyms: Euonymus carolinensis Marsh. 

 Euonymus latifolius Marsh. 

Jinshuang Ma and Gerry Moore 

Illustration source: Britton and Brown 1913 

General Description.—Eastern wahoo, also 

known as burning bush, is a shrub or small tree, up 

to 8 m high, with the main stem up to 10 cm in 

diameter at breast height. The stems are round, 

smooth, and green. The pith is white, round, 

continuous, and somewhat spongy. Its leaves are 

opposite (two per node) and spaced somewhat 

evenly along stem. The petioles can be up to 14 

mm long. The leaf blades are elliptic to oblong 

elliptic, 8.5 to 11.3 by 3.2.to 5.5 cm papery, with 

cuneate to attenuate bases, crenulate margins, and 

acuminate apices. The blades have nine to 10 pairs 

of ascending lateral veins that disappear before 

reaching the margin. The flowers are formed on 

the current season’s growth in small axillary 

clusters (cymes). Each peduncle is 2 to 3 cm long 

and can have several flowers, each one on a 

pedicel 5 to 7 mm long. The flowers are bisexual, 

10 to 12 mm in diameter, with four greenish sepals 

and four brown-purple petals. There are four 

stamens, each 1.5 to 2.0 mm long. The four carpels 

are fused and a nectarous disk is present. The fruit 

of the eastern wahoo is a smooth reddish to pink 

four-lobed (sometimes one or more of the lobes 

aborte) obrhombic capsule, up to 1.7 cm in 

diameter. The capsule opens along the lobes, 

exposing the seeds that are covered by a bright red 

fleshy aril. The seeds (sans aril) are orange, 

glabrous, ellipsoid, 7.0 to 11.0 by 4.0 to 5.5 mm, 

glabrous. Some botanists spell the specific epithet 

“atropurpurea” instead of “atropurpureus” used 

here. This is because they disagree as to whether 

the genus name Euonymus L. should be treated as 

feminine (“atropurpurea”) or masculine 

(“atropurpureus”) (Paclt 1998). In classical Greek, 

the generic name was treated as feminine; 

however, Linnaeus treated the genus as masculine. 

The spelling “Euonymus” has formally been 

conserved over Linnaeus’s originally spelling 

“Evonymus,” which has also occasionally been 

used (Gilstra 1991). 

Range.—Eastern wahoo is widely distributed 

throughout the Eastern United States (no records 

are known for Vermont) westward to the states of 

Kansas, Montana, North Dakota, Oklahoma, South 

Dakota, and Texas (Ma 2001, Shetler and Orli 

2000, USDA 2003). It also is known from Ontario 

(Gleason and Cronquist 1991, Scoggan 1978). 

Ecology.—Eastern wahoo can be found in 

thickets, forests, often in lowlands or near creek 

bottoms, riversides and along the margins of 

marshes. It can also be found along mountain 

slopes. It usually occurs in elevations from sea 

level to 360 m. 

Reproduction.—Hymenopterous insects, 

especially bees, seem to be the main pollinators, 

although wind may also be involved (Brizicky 

1964). The fruits are dispersed by birds and small 

mammals (Brizicky 1964). Eastern wahoo blooms 

in late spring and early summer and the fruits ripen 

in autumn. The plant reproduces asexually through 

rhizomes, which allows it to form large colonies. 

Growth and Management.—Eastern wahoo 

refers well-drained loamy soils. It is a moderately 

fast growing but short-lived shrub or tree. It can be 

planted in Hardiness Zones 3 to 7. Seed requires 8 
336 

to 16 weeks cold stratification prior to sowing. The 

plant can also be propagated with cuttings. 

Benefits.—The common name “burning bush” 

comes from the plant’s bright red fall foliage. The 

red fruit and arillate seeds add to the “burning” 

effect. This makes the eastern wahoo an excellent 

shrub for horticultural use (Rehder 1940). The 

seeds and the vegetative organs of eastern wahoo 

are said to be poisonous (Brizicky 1964). 

References 

Britton, N.L. and A. Brown. 1913. Illustrated flora 

of the northern states, Canada, and the British 

possessions. Vol. 2, 2nd Ed. Scribner, New York. 

735 p. 

Brizicky, G.K. 1964. The genera of Celastrales in 

the Southeastern United States. Journal of the 

Arnold Arboretum. 45: 206-235. 

Gilstra, Z. 1991. Proposal to conserve the spelling 

of 4618 Euonymus (Celastraceae). Taxon 40: 

137-139. 

Gleason, H.A. and A. Cronquist 1991. Manual of 

vascular plants of northeastern United States and 

adjacent Canada. 2nd Ed. New York Botanical 

Garden, Bronx, NY. 910 p. 

Ma, J.S. 2001. A Revision of Euonymus 

(Celastraceae). Thaiszia, Journal of Botany 11: 

1-264. 

Paclt, J. 1998. Proposal to amend the gender of 

Euonymus, nom. cons. (Celastraceae) to 

feminine. Taxon 47: 473-474. 

Rehder, A. 1940. Manual of cultivated trees and 

shrubs, Macmillan Publishing Co, New York. 

996 p. 

Scoggan, H.J. 1978. The Flora of Canada Part 3 – 

Dicotyledoneae (Saururaceae to Violaceae). 

National Museum of Natural Sciences, National 

Museums of Canada, Ottawa. 568 p. 

Shetler, S.G. and S.S. Orli. 2000. Annotated 

checklist of the vascular plants of the 

Washington-Baltimore area. Part. 1. Ferns, fern 

allies, gymnosperms, dicotyledons. Smithsonian 

Institution, Washington, D.C. 186 p. 

USDA, NRCS. 2003. The PLANTS Database. 

Baton Rouge, LA. http:plants.usda.gov. [not 

paged].
337

Fallugia paradoxa (D. Don) Endl. ex Torr. Apache-plume

ROSACEAE 

Synonyms: Sieversia paradoxa D. Don. 

 Geum cercocarpoides DC. ex Ser. 

Juanita A. R. Ladyman 

Drawing source: USDA-Forest Service collection, Hunt 

Institute 

General Description.—Fallugia paradoxa, 

commonly called “Apache-plume,” is an evergreen 

shrub that grows up to 2.5 m tall (Stubbendieck 

and others 1993, Cronquist and others 1997). It is 

often referred to as “semi-evergreen.” In the north 

part of its range, seedlings are deciduous (Meyer 

1974). However, the term “semi-evergreen” may 

be applied because the leaves take on a bronze 

color during some times of the year. The leaves are 

alternate, somewhat fascicled, 0.8 to 1.5 cm long 

and have three to seven deep, narrow lobes. The 

bark on young twigs is white, or light-colored, and 

hairy (villous). The bark becomes darker, hairless 

(glabrous), and shreddy, exfoliating in flakes, as it 

gets older. The showy, flowers have five white 

petals. There are numerous fruits (achenes) in each 

head. Each achene is tipped by a 2.5 to 5 cm long 

feathery, densely hairy, tail-like style that makes 

the plants appear as if they have pom-poms on the 

end of each stalk during the fruiting period. The 

length of the flowering period varies depending 

upon geographic location and elevation. It flowers 

in June through August in all parts of its range but 

flowering is extended from May to December in 

Texas (Powell 1998) and from April to October in 

Arizona and New Mexico (Epple 1995, Martin and 

Hutchins 1980). The common name for Fallugia 

paradoxa, Apache-plume, was coined because the 

feathery seed clusters look like Indian feather 

headdresses (Elmore 1976). Other common names 

include poñil, fallugie (Dayton 1931), feather rose, 

and feather duster bush (Epple 1995). Fallugia is a 

monotypic genus, that is, a genus of a single 

species (Cronquist and others 1997). In addition, 

Apache-plume shows minimal genetic variation 

throughout its wide range (Jones and Johnson, 

1998). Hybrids between Apache-plume and 

cliffrose (Purshia mexicana (D. Don) Henrickson) 

have been reported (Blauer and others 1975, 

McArthur and others 1977). 

Range.—Apache-plume occurs in western Texas, 

Oklahoma, New Mexico, Arizona, southern 

Colorado, south and central Utah, Nevada, and 

southern California in the U.S.A. (USDA 

PLANTS database 2002, Baldwin and others 

2002, McGregor and other 1986, Vines 1986). In 

Mexico it occurs in Coahuila, Chihuahua, and 

Durango (Powell 1998). 

Ecology.—Apache-plume generally grows at 

elevations between 915 and 2,430 m, but in 

northern New Mexico it has been found as high as 

2,700 m (Dick-Peddie 1993). It grows on mesas 

and rocky hillsides in montane shrubland, 

grassland, pinyon-juniper woodland, and yellow 

pine in a variety of alluvial and gravel soils 

(Tidestrom and Kittell 1941, Dick-Peddie 1993, 

Powell 1998, Baldwin and others 2002). It is 

especially abundant along dry margins of watercourses and arroyos (author personal observation, 

Powell 1998). Dick-Peddie (1993) described it as a 

“semi-riparian” species because it tends to 

colonize microhabitats, such as run-off catchments 

and seep areas, which contain more available 
338 

moisture than the surrounding area. Apache-plume 

is relatively drought tolerant and requires 20 to 50 

cm of annual precipitation (Hayward 1990). It is 

tolerant of weakly saline and neutral to moderately 

basic soils (McWilliams 2000). Endomycorrhizae 

were associated with Apache-plume in New 

Mexico (Williams and Aldon 1976). 

Reproduction.—Apache-plume reproduces 

sexually by seed and vegetatively by sprouts 

(suckers) from the roots. The feather-tailed seeds 

are probably primarily wind dispersed. Apacheplume seeds apparently have no dormant period 

(Baskin and Baskin 2001). They are small and 

average approximately 925,932 per kg (Vines 

1986). 

Growth and Management.—Seed can be sown 

by broadcasting it on a prepared bed and covered 

by approximately 5 mm of fine loam or sand. 

Germination rate is from 19 to 65 percent (Vines 

1986). In pots, the germination rate is 30 to 40 

percent, but seedlings are reported to damp off 

easily and require adequate air circulation 

(Hayward 1990). A 4-litre plant is fully established 

and ready for sale in 18 months for the landscape 

market (Hayward 1990). Plants can also be grown 

fairly easily from root cuttings. It is closely grazed 

on overstocked range and, although tolerant of 

grazing with “excellent recuperative powers,” it is 

also reported to decrease under grazing pressure 

(Dayton 1931, McGregor and others 1986). 

Availability of adequate moisture may be an 

important factor in the degree of recovery. 

Apache-plume sprouts after fire (Shaw and 

Monson 1983, McWilliams 2000). In a pinyon 

juniper community it was reported to tolerate fire 

well (Aro 1971). 

Benefits.—Apache-plume is valuable for erosion 

control (Dayton 1931, Stubbendieck and others 

1993, Vines 1986). Its long roots provide effective 

soil stabilization especially in arid situations 

(McWilliams 2000). Apache-plume is important 

winter range for livestock (Dayton 1931, 

McGregor and others 1986, Stubbendieck and 

others 1993, USDAFS 1988). It is reported to have 

only a fair palatability for sheep, goats and cattle 

but closely cropped, stunted plants have been cited 

as evidence of high palatability (Dayton 1931, 

McGregor and others 1986, Stubbendieck and 

others 1993, Warnock 1974). It also furnishes 

important browse for some species of wildlife 

(Dayton 1931, Stubbendieck and others 1993) and 

makes up 1 percent to 5 percent of desert mule 

deer diet (Krausman and others 1997). Time of 

year, environmental conditions, and availability of 

alternate browse appears to influence palatability 

and use. It provides cover for a variety of small 

mammals and birds (Haywood 1990, McWilliams 

2000). It was used for arrow shafts and brooms by 

many Native American peoples (Dunmire and 

Tierney 1995, Moerman 1998). Apache-plume 

brooms are specifically kept inside some Sandia 

Pueblo houses because there is a belief that the 

brooms exert a positive spiritual effect on the 

household (Dunmire and Tierney 1995). The long 

roots were used as cord to tie fencing and make 

ramadas (Dunmire and Tierney 1995, Moerman 

1998). An infusion of leaves was used as a 

shampoo and hair growth stimulant by some 

Native American tribes, and petals are reported to 

prevent stomach gas (Dunmire and Tierney 1995, 

Moerman 1998). One tribe used it in witchcraft to 

cause “insanity” (Moerman 1998). Shrubs are 

attractive as ornamentals and are popularly used 

for landscaping (Hayward 1990, Phillips 1987, 

Morrow 1995). 

References 

Aro, R.S. 1971. Evaluation of pinyon-juniper 

conversion to grassland. Journal of Range 

Management. 24(3): 188-197 

Baldwin B.G., S. Boyd, B.J. Ertter, R.W. 

Patterson, T.J. Rosatti, and D.H. Wilken. 2002. 

The Jepson Desert Manual: Vascular plants of 

southeastern California. University of California 

Press, Berkeley, CA. 624 p. 

Baskin, C.C. and J.M. Baskin. 2001. Seeds. 

Ecology, biogeography, and evolution of 

dormancy and germination. Academic Press, 

New York, NY. 666 p. 

Blauer, A. C., A.P. Plummer, and E.D. McArthur. 

1975. Characteristics and hybridization of 

important Intermountain shrubs. I. Rose family. 

Research Paper INT-169. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Forest and Range Experiment Station, Ogden, 

UT. 36 p. 

Cronquist, A., N.H. Holmgren, and P.K. 

Holmgren. 1997. Intermountain Flora – vascular 

plants of the Intermountain west, U.S.A. Vol. 3 

part A. Subclass Rosidae. The New York 

Botanical Garden, Bronx, NY. 446 p. 

Dayton, W.A. 1931. Important western browse 

plants. Miscellaneous Publication 101. U.S. 
339

Department of Agriculture, Washington, DC. 

214 p. 

Dick-Peddie, W.A. 1993. New Mexico Vegetation 

– past, present, and future. University of New 

Mexico Press, Albuquerque, NM. 244 p. 

Dunmire, W.W. and G.D. Tierney. 1995. Wild 

plants of the Pueblo Province. Museum of New 

Mexico Press, Santa Fe, NM. 290 p. 

Elmore. F.H. 1987. Shrubs and Trees of the 

Southwest Uplands. Southwest Parks and 

Monuments Assoc. Tucson, AZ. 214 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. Falcon Press Publishing Co., Helena, 

MT. 347 p. 

Hayward, P. 1990. Fallugia paradoxa. American 

Nurseryman. 172(7): 126 

Jones, T.A. and D A. Johnson. 1998. Integrating 

Genetic Concepts into Planning Rangeland 

Seedings. Journal of Range Management 

51(6): 594-606. 

Krausman, P.R., A.J. Kuenzi, R.C. Etchberger, 

K.R. Rautenstrauch, L.L. Ordway, and J.J. 

Hervert. 1997. Diets of mule deer. Journal of 

Range Management. 50(5): 513-522. 

Martin, W.C. and C.R. Hutchins. 1980. A Flora of 

New Mexico. Strauss & Cramer, Hirschberg, 

Germany. 2,592 p. 

McArthur, E.D., B.C. Giunta, and A.P. Plummer. 

1977. Shrubs for restoration of depleted range 

and disturbed areas. Utah Science. 35: 28-33. 

McGregor, R.L. and the Great Plains Flora 

Association. 1986. Flora of the Great Plains. 

University Press of Kansas, Lawrence, KS. 

1,402 p. 

McWilliams, J. 2000. Fallugia paradoxa. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/database/feis/ 19 p. 

Meyer, S. 1974. Fallugia paradoxa (D. Don) Endl. 

ex Torr. Apache-plume. In: . F.T. Bonner and 

R.G. Nisley, eds. USDA Forest Service Woody 

Plant Seed Manual Internet site: http:// wpsm.net 

4 p. 

Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. 927 p. 

Morrow, B.H. 1995. Best Plants for New Mexico 

Gardens and Landscapes. University of New 

Mexico Press, Albuquerque, NM. 267 p. 

Phillips, J. 1987. Southwestern landscaping with 

native plants. Museum of New Mexico Press, 

Santa Fe, NM. 141 p. 

Powell, A.M. 1998. Trees and shrubs of TransPecos and adjacent areas. University of Texas 

University Press, Austin, TX . 498 p. 

Shaw, N.L. and S.B. Monson. 1983. Phenology 

and growth habits of nine antelope bitterbrush, 

desert bitterbrush, stansbury cliffrose, and 

apache-plume accessions. In: A.R. Tiedemann 

and K.L. Johnson, comps. Proceedings: 

Research and Management of bitterbrush and 

cliffrose in Western North America: 1982, April 

13-15, Salt Lake City, UT. General technical 

Report: INT-152, USDA Forest Service, 

Intermountain Forest and Range Station, Ogden, 

UT. 

Stubbendieck, J., S.L. Hatch and C.H. Butterfield. 

1993. North American range plants. 4th edition. 

University of Nebraska Press, Lincoln, NE. 

493 p. 

Tidestrom, I. and T. Kittell. 1941. A flora of 

Arizona and New Mexico. The Catholic 

University of America Press, Washington, DC. 

897 p. 

USDA Forest Service. 1988. Range Plant 

Handbook. Dover Publications, Inc., New York, 

NY. 842 p. 

USDA PLANTS database. 2002. Internet site: 

http:// plants.usda.gov/cgi_bin/plant_profile. 

cgi? symbol=FAPA#. [not paged]. 

Vines, R.A. 1986. Trees, shrubs, and woody vines 

of the Southwest. Sixth printing. University of 

Texas Press. Austin, TX. 1,104 p. 

Warnock, B.H. 1974. Wildflowers of the 

Guadalupe Mountains and the sand dune 

country, Texas. Sul Ross university, Alpine, TX. 

176 p. 

Williams, S.E. and E.F. Aldon. 1976. 

Endomycorrhizal (vesicular arbuscular) 
340 

associations of some arid zone shrubs. The 

Southwestern Naturalist. 20(4): 437-444. 
341

Faramea occidentalis (L.) A. Rich. cafeíllo

RUBIACEAE 

Synonyms: Coffea occidentalis Jacq. 

 Faramea odoratissima DC. 

 Pavetta coffeoides Ham. 

 Tetramerium odoratissimum (Gaertn. f.) DC. 

 Ixora occidentalis L. 

John K. Francis 

General Description.—Cafeíllo, also known as 

false coffee, café cimarrón, palo de toro, hueso, 

jasmín de estrella, and bois flèche, is a tall shrub in 

the Caribbean islands and a small understory tree 

in continental tropical America. It has dark-green, 

opposite lanciolate leaves and straight, usually 

single, gray or dark-brown stems. The twigs are 

slender and green with rings at the nodes (Little 

and Wadsworth 1964). The stems are straight and 

often without branches for a meter or more. 

Range.—The native range extends from Cuba 

through Trinidad and from central Mexico through 

Central America to Ecuador, Peru, Brazil, and the 

Guyanas (Little and Wadsworth 1964). The 

species is not known to have naturalized 

elsewhere. 

Ecology.—Cafeíllo is tolerant and grows and 

reproduces in the understories and mid-stories of 

secondary forests. The species is most common in 

mid- to late-secondary forests. Cafeíllo is found in 

two of the Life Zones in Puerto Rico: moist 

subtropical forest and much of the wet subtropical 

forest from about 900 to 2800 mm of precipitation 

(Holdridge 1967). Alluvial soils and residual soils 

derived from sedimentary and volcanic rocks are 

suitable habitat. Excessively drained and very 

poorly drained soils are usually not colonized. 

Although not forming pure or dense stands, 

cafeíllo is common, at least in Puerto Rico and 

Panama. 

Reproduction.—Small, white, fragrant flowers 

have a short tube tipped by four pointed lobes. 

Cafeíllo flowers generally occur in threes. The 

fruits are blue-black or dark-purple drupes borne 

singly or in pairs from the leaf axils near the twig 

ends. The fruits measure 8 to 10 mm in diameter 

(Howard 1989). Each fruit contains a single seed. 

A Puerto Rican collection averaged 0.825 + 0.123 

g/fruit. The seeds from the same collection 

numbered 3,980 seeds/kg. Ninety-six percent 

germinated beginning 57 days after sowing 

(Francis and Rodríguez 1993). In Central America, 

the seeds are disbursed by birds and monkeys. 

Rodents and a number of other animal species eat 

and destroy the seeds after fruit-fall (Schupp 

1990). A rust, Aecidium farameae Arthur, which 

attacks the ovaries and pedicels of the flowers, 

results, in some years, in the destruction of 32 to 

83 percent of the fruits of individual plants 

(Travers 1998). A high recruitment of seedlings 

(compared to other species) results in high 

population densities in the forest understorey in 

Panama (Schupp 1990). 

Growth and Management.—Cafeíllo 

occasionally exceeds 20 m in height and 20 cm in 

diameter at breast height in Panama (O’Brien and 

others 1995). However, in Puerto Rico, mature 

shrubs usually reach 4 m in height and sometimes 

attain as much as 10 m in height (Liogier 1997). 

Total above-ground, ovendried weight of cafeíllo 

plants in Puerto Rico that ranged between 1.37 m 

in height (with diameter greater than 0) and 5 cm 

in d.b.h. may be predicted by the model: 

 
342 

TDW = 30.045D2L (adjusted R-squared = 0.962), 

where TDW is total dry weight in grams, D is 

diameter in cm at 30 cm above the ground, and L 

is stem length in meters. Lower stem density of 

Puerto Rican shrubs averaged 0.62 + 0.04 g/cm3 

(Francis 2000). Stem density of saplings from 

Panama was 0.57 g/cm3

 (King 1990). Allometric 

relationships for cafeíllo in Panama may be found 

in O’Brien and others (1995) and King (1990). As 

an understory tree, cafeíllo is slow-growing and 

long lived. Based on recent growth rates, the 

largest individuals in a Panamanian stand were 

estimated to exceed 200 years in age. 

There is no record of plantations or 

planted ornamental cafeíllo. The species is able to 

reproduce by sprouting after injury. Thirty-six 

percent of the individuals in a Panamanian stand 

were of sprout origin (Putz and Brokaw 1989).

Benefits.—Cafeíllo wood is heavy, hard, and 

tough. However, uses are mainly limited to fence 

posts, vegetable stakes, tool handles, and 

fuelwood. In herbal medicine, cafeíllo is 

reportedly used as an antiseptic, astringent, and for 

diarrhea and anemia (Liogier 1990). Birds 

consume the fruits. The use of cafeíllo as an 

ornamental or in forest restoration efforts has not 

been reported. Cafeíllo is a fine looking plant and 

could probably be used as a back-ground plant or 

screen under shade in yards, estates, and parks. 

References 

Francis, J.K. 2000. Estimating biomass and carbon 

content of saplings in Puerto Rican secondary 

forests. Caribbean Journal of Science 36(3-4): 

346-350. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

Holdridge, L.R. 1967. Life zone ecology. Tropical 

Science Center, San José, Costa Rica. 206 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

King, D.A. 1990. Allometry of saplings and 

understory trees of a Panamanian forest. 

Ecology 4(1): 27-32. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

O’Brien, S.T., S.P. Hubbell, P. Spiro, R. Condit, 

and R.B. Foster. 1995. Diameter, height, crown, 

and age relationships in eight neotropical tree 

species. Ecology 76(6): 1926-1939. 

Putz, F.E. and N.V.L. Brokaw. 1989. Sprouting of 

broken trees on Barro Colorado Island, Panama. 

Ecology 70(2): 508-512. 

Schupp, E.W. 1990. Annual variation in seedfall, 

postdispersal predation, and recruitment of a 

neotropical tree. Ecology 71(2): 504-515. 

Travers, S.E. 1998. The effect of rust infection on 

reproduction in a tropical tree (Faramea 

occidentalis). Biotropica 30(3): 438-443. 
343

Forestiera acuminata (Michx.) Poir. swamp privet

OLEACEAE 

Synonyms: Adelia acuminata Michx. 

 Borya acuminata (Michx.) Willd. 

Kristina Connor 

Drawing credit: USDA—Forest Service collection, Hunt 

Institute 

General Description.—Swamp privet, also called 

eastern swamp privet, is a deciduous shrub or 

small, open-crowned tree, occasionally reaching 

10 m in height but more often averaging 1.5 to 2.5 

m (Krüssmann 1986, Johnson and Hoagland 

1999). It is commonly multitrunked (Mackay and 

Finical 1999, U.S. Army Corps of Engineers 

1997). The bark ranges from gray to dark brown in 

color and is either smooth or ridged (Johnson and 

Hoagland 1999, USGS 2002). Twigs are light 

brown, glabrous, and have conspicuous lenticels 

(Johnson and Hoagland 1999). Leaves are simple, 

opposite, and dull, yellowish green. They range in 

size from 3 to 11 cm in length, and 2.5 to 5 cm in 

width. Leaves have long, pointed tips (acuminate) 

and petioles that are slender and slightly winged at 

the base. While leaf margins are usually smooth, it 

is not uncommon for the apex half to be finely 

serrated. 

Range.—Swamp privet is found in the wet 

woodlands and swamps of the Southeastern United 

States, from South Carolina south to Florida and 

westward into Texas, Oklahoma, and Kansas. It 

ranges as far north as Illinois, Indiana, and 

Missouri (Krüssmann 1986, USGS 2002). 

Ecology.—Swamp privet grows primarily in wet 

areas, in swamps and bottomland forests, along 

streams, in wet woods and sloughs, and around 

ponds and lakes (Johnson and Hoagland 1999, 

USGS 2002). It will grow in either full or partial 

sun and is highly adaptable to various soils 

(MacKay and Finical 1999). The species 

reportedly thrives in a variety of soil types, sand, 

loam or clay, and at pH’s ranging from acidic to 

basic. Swamp privet does, however, require that 

the soils be moist (Plants for a Future 2001). It 

frequently produces new stems from the base of 

senescing individuals, especially when soils are 

flooded (Mackay and Finical 1999, U.S. Army 

Corps of Engineers 1997). 

Reproduction.—Swamp privet plants are 

monoecious; flowers emerge in the spring before 

the leaves unfold. They are yellow, fragrant and 

small, the males sessile or almost so, female 

flowers in small panicles or fascicles (Bailey and 

Bailey 1976, Botanical Institute 2002, Johnson and 

Hoagland 1999, Krüssmann 1986). They have no 

petals and minute or absent sepals. Male flowers 

have one to four stamens, female flowers have 

simple or two-lobed stigmas (Johnson and 

Hoagland 1999, Botanical Institute 2002, USGS 

2002). Fruits are dark purple or black oblong 

drupes, 8 to 12 mm long with one, rarely two, 

seeds. They appear in summer and are promptly 

shed (Johnson and Hoagland 1999). Although no 

data are provided to support this, the Plants for a 

Future website (2001) suggests that seeds be sown 

as soon as they ripen and that cuttings can be taken 

of half-ripe wood in summer or mature wood in 

winter. 

Growth and Management.—Swamp privet, 

while adaptable to many soil types, must be 

located in moist areas. It is moderately tolerant of 

heat and can tolerate full or partial sunlight 

(Mackay and Finical 1999). It has been reported in 

cultivation (Krüssmann 1986). 

Benefits.—The fruits of swamp privet are 

considered good food for wildlife (Johnson and 
344 

Hoagland 1999). A decoction of the roots and bark 

has been used as a health beverage, and the wood 

has been used for turning (Bailey and Bailey 

1976), although reports on its durability vary 

(Plants for a Future 2001). 

 

References

Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: 

A Concise Dictionary of Plants Cultivated in the 

United States and Canada. McMillan Publishing 

Co., Inc., New York. 1,312 p. 

Botanical Institute. 2002. Forestiera Poiret. 

Goteborg University. http://www.systbot. 

gu.se/staff/evawal/ oleaceae/forestiera.html. 2 p. 

Johnson, F.L. and B.W. Hoagland. 1999. 

Forestiera acuminata (Michx.) Poir. Catalog of 

the Woody Plants of Oklahoma, Oklahoma 

Biological Survey http://www.biosurvey.ou.edu. 

1 p. 

Krüssmann, G. 1986. Manual of Cultivated BroadLeaved Trees and Shrubs. Volume II, E-Pro. 

Timber Press, Beaverton, OR. 445 p. 

Mackay, W. and L. Finical. 1999. A special 

project of the Texas Agricultural Experiment 

Station. Benny Simpson’s Texas Native Trees. 

Texas A&M University, College Station. 

http://aggie-horticulture.tamu.edu/natives. 1 p. 

Plants for a Future (PFAF). 2001. Resource and 

Information Centre for Edible and Other Useful 

Plants. Forestiera acuminata. 

http://www.scs.leeds. ac.uk/ pfaf/index.html. 4 

p. 

U.S. Army Corps of Engineers. 1997. Wetlands 

Delineation Manual. Appendix C: Vegetation. 

The Wetlands Regulation Center 

http://www.wetlands. com/coe/87manapc.htm. 

6 p. 

USGS. 2002. Swamp privet (Forestiera 

acuminata). Northern Prairie Wildlife Research 

Center. Directory of Resource 

http://www.npwrc.usgs.gov/resource/ 

othrdata/plntguid/species. 2 p. 
345

Forestiera segregata (Jacq.) Krug & Urban Florida privet 

OLEACEAE 

Synonyms: Myrica segregata Jacq. 

 Adelia porulosa Michx. 

 Forestiera porulosa (Michx.) Poir. 

 Adelia segregata (Jacq.) O. Kuntze 

 Adelia pinetorum Small 

 Forestiera cassinoides Poir. 

John K. Francis 

General Description.—Florida privet, also known 

as Florida swamp privet, wild olive, and ink-bush, 

is a semideciduous shrub or small tree to 7 m in 

height. The plant is supported by a weak taproot 

and more robust, tan-colored lateral roots. The 

branches are slender. The opposite leaves have 

short petioles, 1 to 6 mm long, and narrowly 

elliptic to obovate blades, 1 to 7 cm long, with 

entire margins. Tiny, greenish-yellow flowers are 

borne in clusters at the leaf axils. The fruits are 

slightly curved, ovoid, black drupes, 6 to 10 mm 

long, that have a bitter, unpleasant flavor and stain 

cloth and skin. Each contains one greenish-yellow 

seed (Howard 1989, Liogier 1995, Long and 

Lakela 1976, Nelson 1996). 

Range.—Florida privet is native to Florida, 

Georgia, South Carolina, the Bahamas, the Greater 

Antilles, the Cayman Islands and Anguilla, 

Barbuda, Antigua, La Désirade, and Marie Galante 

in the Lesser Antilles (Howard 1989, Liogier 1995, 

National Trust for the Cayman Islands 2002, 

Natural Resources Conservation Service 2002). 

Long and Lakela (1976) identify two varieties: var. 

segregata with glabrous stems and leaf blades 

more than 3 cm long in Florida and Georgia, and 

var. pinatorum, which has puberulent stems and 

leaf blades 1 to 3 cm long in South Florida. 

Ecology.—Florida privet grows best in partial 

shade (Horticopia 2002). It is tolerant of drought, 

salt, and alkaline soils (University of Florida 

Cooperative Extension Service 2002). Florida 

privet competes well with shrubs and herbs and 

grows in hammocks and pinelands in Florida (Long 

and Lakela 1976). In Puerto Rico and other 

Caribbean Islands, it is most frequently seen in dry 

forests in limestone areas near the coast. The 

species is rare in the United States and has become 

a candidate for Federal listing (Everglades 

Mitigation Bank 2002). 

Reproduction.—In Florida, Florida privet flowers 

in the spring (Nelson 1996). It is insect pollinated. 

A collection of fruits from Puerto Rico weighed an 

average of 0.1209 + 0.0035 g/fruit. Air-dried seeds 

cleaned from them weighed an average of 0.0359 +

0.0006 g/seed or 27,900 seeds/kg. Sown in moist 

potting mix, they began germinating in 7 days and 

completed 89 percent germination in 49 days 

(author’s observation). The seeds are dispersed by 

birds that eat the fruits. Seedlings do not occur in 

large numbers. 

Growth and Management.—Florida privet grows 

slowly in the early seedling stage but later 

accelerates so that seedlings are ready to transplant 

at the end of 1 year. Nursery production is 

normally from seed. Wildlings of all sizes can be 
346 

successfully transplanted (Workman 1980). 

Beyond the seedling stage, it has a moderate 

growth rate (University of Florida Cooperative 

Extension Service 2002). The largest Florida privet 

recorded measures 20.2 cm diameter at breast 

height and 5.5 m in height (American Forests 

2002). 

Benefits.—Florida privet is recommended for 

xeriscape (unwatered) landscaping (South Florida 

Water Management District 2001) and is available 

through a number of commercial nurseries. It is 

useful for hedges and foundation and specimen 

plantings. Warblers, bluebirds, mockingbirds, and 

vireos feed upon the fruits in South Florida 

(Broschat and Verkade 2002, Cowley 2002). 

Honeybees, butterflies, and other insects are 

attracted to the flowers for the nectar (Workman 

1980). 

References 

American Forests. 2002. National register of big 

trees: Florida-privet, Forestiera segregata. 

http://www.americanforests.org/resources/bigtre

es/register.php?details=2000. 1 p. 

Broschat, T.K. and S.V. Verkade. 2002. 

Landscaping to attract birds in South Florida. 

http://www.floridaplants.com/landscape/birds. 

htm. 3 p. 

Cowley, M. 2002. Florida native plants: olive 

family. Native Florida Web Ring. 

http://nsis.org/garden/family/olive.html. 2 p. 

Everglades Mitigation Bank. 2002. Endangered 

and threatened plants at the EMB. 

http://www.fpl.com/environment/emb/contents/e

ndangered_plants.shtml. 1 p. 

Hortocopia. 2002. Forestiera segregata, wild olive. 

http://hortpix.com/pc1848.htm. 1 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Vol. 6. Arnold Arboretum, 

Harvard University, Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

tropical Florida. Banyon Books, Miami, FL. 

962 p. 

National Trust for the Cayman Islands. 2002. 

Virtual herberium: Oleaceae, Forestiera 

segregata (Jacq.) Krug & Urban. http://www. 

virtualherbarium.org/vh/CAYM/1002.html. 2 p. 

Natural Resources Conservation Service. 2002. 

Plants profile: Forestiera Poir. swampprivet. 

http://plants.usda.gov/cgi_bin/plant_profile.cgi?

symbol=FORES. 3 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391. p. 

South Florida Water Management District. 2001. 

Quick facts on xeriscape: how to save water 

through water-smart landscaping. http://www. 

sfwmd.gov/images/pdfs/splash/splxeris.pdf. 2 p. 

University of Florida Cooperative Extension 

Service. 2002. Native shrubs of South Florida. 

http://edis.ifas.ufl.edu/BODY_EH159. 9 p. 

Workman, R.W. 1980. Growing native. The 

Sanibel-Captiva Conservation Foundation, Inc., 

Sanibel, FL. 137 p. 
347

Fouquieria splendens Engelm. ocotillo

FOUQUIERIACEAE 

Synonyms: none 

Juanita A.R. Ladyman 

Illustration credit: Tonia Masaood 

General Description.—The common name for 

Fouquieria splendens is Ocotillo. This is a relative 

of the renowned boojum tree (Fouquieria 

columnaris) of Baja California. Ocotillo is an erect 

shrub with several spiny ascending stems that 

grow up to 3 m tall. A mature individual may have 

as many as 75 stems (Epple 1995). The leaves are 

oblanceolate to oblong-obovate and only 10 to 25 

mm long. The bright scarlet flowers are showy, 

and an individual may flower several times during 

the year. The flowers are arranged in an 8 to 25-cm 

-long panicle. Each flower has a tubular corolla, 

five petals, and 10 to 15 exerted stamens. The seed 

capsule is ovoid and incompletely three-celled. 

The flat seeds are white with a fringe of hair-like 

filaments. Flowering is from March to June 

depending upon latitude (Martin and Hutchins 

1980). 

Range.—Ocotillo occurs in Arizona, southern 

California, New Mexico, Nevada, and Texas in the 

USA. In Mexico it occurs in Sonora, Coahuila, 

Chihuahua, Durango, Zacatecas, San Luis Potosí, 

Hildalgo, and Baja California (Mason and Mason 

1987). 

Ecology.—Ocotillo grows in shallow soils on 

mesas, outwash plains, and rocky slopes from sea 

level to as high as 2,050 m elevation in the 

Guadalupe and Del Norte mountains of the TransPecos region of Texas. Soils are generally rocky, 

shallow, well-drained, and of limestone or granitic 

origin and are often underlain by caliche (Mathews 

1994). Although not restricted to calcareous soils, 

ocotillo tends to be characteristic of a calcareous 

community (Schmutz and others 1992). It has a 

relatively shallow root system (Burgess 1995). It is 

often found on southern facing slopes. In Mexico 

it also occurs in both the Sonoran and Chihuahuan 

deserts. Communities in which it commonly 

occurs are mesquite-grasslands (Mason and Mason 

1987), high desert bunch-grass (Schmutz and 

others 1992), desert scrub (Van Devender 1995) 

and mesquital (mesquite), motorral desértico 

microfilo (fine-leaf desert shrubland), and pastizal 

(grassland) defined by Flores-Mata and others 

(1971). 

Reproduction.—Ocotillo may self-pollinate to a 

limited extent, but flowers are generally crosspollinated by hummingbirds and bees. In southern 

Arizona flowering coincides with the northward 

migration of hummingbirds (Mathews 1994). Seed 

set per flower is greater for plants flowering during 

migration than for plants flowering after 

hummingbirds have migrated (Rathcke and Lacey 

1985, Waser 1979). The winged seeds of ocotillo 

are probably wind dispersed. 

Growth and Management.—Ocotillo is a longlived stable element of desert vegetation but with 

apparently specialized requirements for seed 

germination that limit its capacity to exploit 

opportunities for population expansion (Zedler 

1981). There seems to be no significant change in 

the distribution and abundance of this species 

within the recent past. Evidence suggests that 

ocotillo habitat has increased as grasslands have 

declined (Van Devender 1995). However, 

considering the amount of urban development 

within octotillo habitat, the actual area occupied by 

the species may not have changed significantly. It 

may be significant to overall species sustainability 

that seedlings are rarely found on disturbed soils 

(Yeaton and others 1977) but are found under the 

canopy of mature plants (McAuliffe 1988, Young 
348 

and Young 1986). It is a slow-growing species, 

and estimates of its life span range from less than 

60 to greater than 72 years (Mathews 1994). It is 

likely that life span is related to latitude and 

substrate. The leaves are winter- and droughtdeciduous (Schmutz and others 1992). The leaves 

appear after rains and are a reliable indicator of 

recent rainfall (Powell 1998). They tend to mature 

and fall within a few weeks indicating that 

available photosynthate is limited and that growth 

is dependent upon precipitation. Ocotillo is a 

“water storer” in that the stems store relatively 

large amounts of water that buffer against drought 

stress (Burgess 1995). Wildfires may be 

detrimental to ocotillo populations. It rarely 

sprouts from the root crown following damage 

from fire, and its sprouting ability is probably 

dependent on fire severity (Ahlstrand 1982, 

McPherson 1995, White 1969). Mortality rates of 

40 percent to 50 percent were reported after light 

to moderate fire in Arizona (Thomas 1991). 

Seedlings are not known to establish in recently 

burned areas (Mathews 1994), and its seeds are 

unlikely to survive for long in the soil (Zedler 

1981). In fact, burning has been suggested as a 

control method for woody species, such as 

Fouquieria splendens, in desert grasslands 

(Mathews 1994). 

Benefits.—The endangered Lucifer hummingbird's habitat in New Mexico centers on slopes and 

adjacent canyons in arid montane areas dominated 

by ocotillo and agave species (New Mexico 

Department of Game and Fish 1991). Ocotillo has 

no recorded forage value, presumably because its 

leaves are so short-lived (Dayton 1931). Ocotillo 

plants are popularly used for landscaping. Plants 

sold in reputable nurseries are obtained from 

private land under permit. There are typically two 

“types” of product. Smaller plants are sold in 1 

gallon pots or larger specimens are sold “balled 

and burlapped.” Although the plants transplant 

well if there are sufficient roots attached, many 

that are dug are not treated well and two thirds or 

more of those dug up are likely not to survive. 

Propagation has been reported using cuttings 

although conversations with horticulturalists 

indicate that this may be less easy than the 

literature indicates. The plant can also be grown 

from seed but is a relatively slow grower. Seeds 

apparently collected from Mexico can be 

purchased in the United States (Tradewinds 

Import-Export Company 2002). This trade 

between regions where the plant is native does 

raise the potential for genetic modification, or 

“dilution,” of locally adapted genotypes by 

cultivated genotypes originally adapted to another 

region. This may pose a threat although, because 

of the slow growth and low turnover of 

individuals, the effects would be a long time in 

becoming apparent. For example, the cultivated 

genotypes may have particularly high fecundity 

and may be physiologically robust but do not have 

the genetic composition to be resistant to 

environmental or biological (e.g. disease or insect 

infestation) events that are periodic (perhaps 

decades apart) in the local area. Hedges or fences 

of living stems are common, especially in Mexico. 

It does not appear to be grown for its medicinal 

value. Historically the Mahuna Native Americans 

used it as a “blood medicine” (Moerman 1998). 

The Apache Indians used powdered roots to treat 

wounds and painful swellings and they also bathed 

in an ocotillo root mixture to relieve fatigue 

(Krochmal and others 1954, Powell 1998, Vines 

1960). A beverage made from ocotillo flowers was 

used for cough medicine (Vines 1960). The 

Cahuilla, Papago and Yavapai tribes used the 

flowers in beverages, the fruits for a “candy,” and 

the stalks as building material (Moerman 1998). 

The thorns were used for ear piercing by the 

Papago. The resin and wax from ocotillo bark was 

used for conditioning leather (Krochmal and others 

1954, Powell 1988, Vines 1960). It seems that all 

uses are localized, personal, or historic in the 

Southwestern United States. It is not known if 

ocotillo is more widely used today in Mexico. 

References 

Ahlstrand, G.M. 1982. Response of Chihuahuan 

Desert Mountain Shrub Vegetation to Burning. 

Journal of Range Management 35 (1): 62-65. 

Burgess, T.L. 1995. The dilemma of coexisting 

growth forms. In: The desert grassland. M.P. 

McClaran and T.R. Van Devender, eds. The 

University of Arizona Press, Tucson, AZ. p. 31-

67. 

Dayton, W.A. 1931. Important western browse 

plants. Miscellaneous Publication 101. U.S. 

Department of Agriculture, Washington, DC. 

214 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. Falcon Press Publishing Co., Helena, 

MT. 347 p. 

Flores-Mata, G.F. J.J. Lopez, X.M. Sánchez, F.M. 

Ruiz, and F.T. Takaki. 1971. Memoria del Mapa 

Tipos de Vegetación de la República Mexicana. 
349

Secretaria de Recursos Hidráulicos, Dirección 

de Agrología, Mexico, DF. 59 p. 

Krochmal, A., S. Paur, and P. Duisberg. 1954. 

Useful native plants in the American 

Southwestern deserts. Economic Botany. 8: 3-

20. 

Martin, W.C. and C.R. Hutchins. 1980. A Flora of 

New Mexico. Strauss & Cramer, Hirschberg, 

Germany. 2,592 p. 

Mason, C.T. and P.B. Mason. 1987. A handbook 

of Mexican roadside flora. The University of 

Arizona Press, Tucson, AZ. 380 p. 

Matthews, R.F. 1994. Fouquieria splendens. The 

Fire Effects Information System Data base. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Research Station, Intermountain 

Fire Sciences Laboratory, Missoula, MT. http:// 

www.fs.fed.us/database/feis/plants/shrub/fouspl/ 

all.html. 15 p. 

McAuliffe, J.R. 1988. Markovian dynamics of 

simple and complex desert plant communities. 

American Naturalist. 131(4): 459-490. 

McPherson, G.R. 1995. The role of fire in desert 

grasslands. In: The desert grassland. M.P. 

McClaran and T.R. Van Devender, eds. The 

University of Arizona Press, Tucson, AZ. p. 

130-151. 

Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. 927 p. 

New Mexico Department of Game and Fish. 1991. 

Handbook of species endangered in New 

Mexico. Dept. of Game and Fish., Santa Fe, 

NM. 185 p. 

Powell, A.M. 1988. Trees & shrubs of TransPecos Texas including Big Bend and Guadalupe 

Mountains National Parks. Big Bend Natural 

History Association, Big Bend National Park, 

TX. 536 p. 

Powell, A.M. 1998. Trees & shrubs of TransPecos and adjacent areas. University of Texas 

University Press, Austin, TX . 498 p. 

Rathcke, B. and E.P. Lacey, 1985. Phenological 

patterns of terrestrial plants. Annual Reviews of 

Ecological Systems. 16: 179-214. 

Schmutz, E.M., E.L. Smith, P.R. Ogden, M.L. 

Cox, J.O. Klemmedson, J.J. Norris, and L.C. 

Fierro. 1992. Desert grassland. In: Natural 

grasslands: Introduction and Western 

Hemisphere. R.T. Coupland, ed. Elsevier, New 

York. p. 337-362. 

Thomas, P.A. 1991. Response of succulents to 

fire: A review. International Journal of Wildland 

Fire 1(1): 11-22. 

Tradewinds Import-Export Company. 2002. Seeds 

Department – USA. P.O. Box 218650, Houston, 

Texas 77218 USA. http://www.abonline. 

com/products/seeds_f.htm#f. [not paged]. 

Van Devender, T.R. 1995. Desert grassland 

history: changing climates, evolution, 

biogeography, and community dynamics. In:

The desert grassland. M.P. McClaran and T.R. 

Van Devender, eds. The University of Arizona 

Press, Tucson, AZ. p. 68-99. 

Vines, R.A. 1986. Trees, shrubs, and woody vines 

of the Southwest. Sixth printing. University of 

Texas Press. Austin, TX. 1,104 p. 

Waser, N.M. 1979. Pollinator availability as a 

determinant of flowering time in ocotillo 

(Fouquieria splendens). Oecologia. 39(1): 107-

121. 

White, L.D. 1969. Effects of a wildfire on several 

desert grassland shrub species. Journal of Range 

Management 22: 284-285. 

Yeaton, R.I., J. Travis, and E. Gilinsky. 1977. 

Competition and spacing in plant communities: the 

Arizona upland association. Journal of Ecology. 65: 

587-595 

Young, J.A. and C.G.Young. 1986. Collecting, 

processing, and germinating seeds of wildland 

plants. Timber Press. Portland, OR. 236 p. 

Zedler, P.H. 1981. Vegetation change in chaparral 

and desert communities in San Diego County, 

California. In: D.C. West, H.H. Shugart, D. B. 

Botkin, eds. Forest succession: Concepts and 

application. Springer-Verlag, New York. p. 

406-430
350 

Furcraea foetida (L.) Haw. Mauritius hemp 

AGAVACEAE 

Synonyms: Furcraea gigantea Vent. 

 Agave gigantea D. Dietr. 

John K. Francis 

General Description.—Mauritius hemp, also 

known as green aloe, female karata, maguey, 

mayuey criollo, cocuisa, giant cabuya, and aloes 

vert, is a robust shrub with a basal rosette about 

2.5 to 3.5 m in diameter and flowering stalks 5 to 

10 m in height. The plant can be called a shrub 

because it is perennial, because it is shrub size, and 

because it has a fibrous-woody sheath surrounding 

the core of the short stem (20 to 30 cm long) 

within the rosette, and a woody flowering stalk. 

Mauritius hemp has no taproot, relatively fine 

lateral roots, and many fine roots. Its green to 

yellow-green leaves are linear-lanceolate to 

oblanceolate, pointed at the tip, and are fleshy with 

thread-like parallel fibers. They are up to 25 cm 

wide, have marginal spines but lack a terminal 

spine. The inflorescences (panicles) are terminal 

and contain many pendulous, fragrant, white, 

greenish-white, yellowish-green, or pale bluegreen, 2.5 to 3.3 long by 1.0 by 1.8 cm wide 

flowers. Flowers open a few at a time for several 

weeks. Seed development has not been observed 

by the author in Puerto Rico and is apparently rare 

elsewhere. Bulbils 1 to 16 cm long develop 

abundantly on the peduncles after flower 

dehiscence. There are 2n = 60 chromosomes 

(author’s observation, Bailey 1941, Howard 1979, 

Pacific Island Ecosystems at Risk 2003, Schlegel 

2003). 

Range.—Mauritius hemp is native to the Greater 

Antilles, and from Guadaloupe south through 

northern South America to Brazil (Grisebach 

1963, Howard 1979). The species has been widely 

planted. It has naturalized in Florida, Hawaii, 

Marquesas Islands, French Polynesia, Tonga, and 

is present and probably naturalized in many other 

places (Pacific Island Ecosystems at Risk 2003). 

Taxonomy.—Mauritius hemp in Puerto Rico was 

known previously as Furcraea tuberose (Miller) 

Ait. f. (Liogier and Martorell 2000). The current 

assignment of the species to F. foetida is based on 

the statement by Howard (1979) that F. foetida 

occurred throughout the Antilles and that F. 

tuberosea is endemic to the Lesser Antilles, 

specifically from St. Barts southward. 

Characteristics given to separate the two species 

are unreliable by Howard’s own admission. 

Mature plants examined by the author had 

maximum leaf widths of 18 to 20 cm, leaf lengths 

of 1.5 to 2 m, oblanceolate leaves, weak fetid 

smell of leaf juices, and bulbiferous reproduction. 

Reliable identification of the Puerto Rican and 

Caribbean Furcrea species will have to wait for 

further study. 

Ecology.—In Puerto Rico, Mauritius hemp grows 

in moist areas that receive from about 1200 to 

2500 mm of mean annual precipitation from near 

sea level to 1,000 m in elevation. The species 

grows in all types of well-drained soils, including 

poor and eroded soils, and frequently grows on 

rocks, cliffs, and rarely in crotches of trees. It is 

resistant to short-term drought and salt spray and 

can survive temperatures as low as -7 to -4 °C. 

Growth is best in full sun, which most often leads 

to successful flowering. Mauritius hemp also 

grows in partial but not heavy shade. When plants 

are young, competition with tall grass, weeds, 

shrubs, and trees is a critical limiting factor. Cattle 

graze and often completely uproot small plants. 

They are ignored by cattle once they become large 

(author’s observation, Desert-Tropicals Nursery 

2003, Pacific Island Ecosystems at Risk 2003). 

This species, as others of the genus, has 
351

crassulacean acid metabolism by which it fixes 

CO2 during the night and incorporates it into 

carbohydrates during the day (personal 

communication with Ernesto Medina, plant 

physiologist, Centro de Ecología, Instituto de 

Investigaciones, Caracas, Venezuela). 

Reproduction.—Flowering, which may occur at 

any time of the year, apparently begins when 

plants attain sufficient size and vigor to support the 

large flower stalk. Plants die about 1 year after the 

onset of flowering. The flowers are visited by 

honey bees (Apis mellifera L.). There is no record 

in the literature of seed weights. Fresh bulbils (n = 

100) collected by the author in Puerto Rico 

weighed an average of 2.72 + 0.31 g/bulbil with a 

range of 0.18 to 16.27 g. Placed in moist potting 

mix, 100 percent of them rooted within 1 week. 

Bulbils are formed by the thousands on single 

plants and can form dense thickets. Gravity is the 

only known natural means of dispersal. The 

presence of scattered individuals indicates other 

transport, perhaps fruit bats. 

Growth and Management.—The best way to 

artificially propagate Mauritius hemp is to place 

bulbils in pots of potting mix. A group of bulbils 

planted in Puerto Rico grow rapidly with no 

mortality, reached an average of 17.8 + 0.4 cm in 

height in 1 month, and were ready to outplant in 1 

to 2 months. New plants need protection from 

weeds for a few months, but afterwards the plants 

need little care. Individual plants probably live 

from 5 to 20 years depending on growing 

conditions. One owner reported that an ornamental 

plant died after flowering at 15 years old (author’s 

observation). Mauritius hemp is rarely common 

enough in its native range to need control. When 

control is needed, particularly in invaded areas, 

individual plants can be killed by grubbing them 

out or by spray or drizzle application of 2,4-D or 

triclopyr in oil (Pacific Island Ecosystems at Risk 

2003). 

Benefits.—Mauritius hemp helps hold the soil, 

furnishes cover for wildlife, and adds to the 

aesthetics of wildlands. The species is widely, 

although not heavily, used as a landscaping plant 

for accent and curiosity. A variegated form is 

available (Desert-Tropicals Nursery 2003). 

Mauritius hemp was once widely cultivated for 

fiber, hence the common name. Extracts of the 

roots are used as ingredients in tonics for purifying 

the blood, and the dried leaves are used to control 

swelling and to aid in wound healing (NúñezMeléndez 1982). 

References

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. Vol. 2. The MacMillan Company, 

New York. p. 1,201-2,422. 

Desert-Tropicals Nursery. 2003. Mauritius hemp, 

sisal, maguey. http://www.desert-tropicals.com/ 

Plants/Agavaceae/Furcraea_foetida.html. 2 p. 

Grisebach, A.H.R. 1963. Flora of the British West 

Indian Islands. J. Cramer, Weinheim, Germany. 

789 p. 

Howard, R.A. 1979. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 3. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 586 p. 

Liogier, H.A. and L.F. Martorell. 2000. Flora of 

Puerto Rico and adjacent islands: a systematic 

synopsis. 2nd ed. Editorial de la Universidad de 

Puerto Rico, Río Piedras, PR. 382 p. 

Núñez-Meléndez, E. 1982. Plantas medicinales de 

Puerto Rico. Editorial de la Universidad de 

Puerto Rico, Río Piedras, PR. 498 p. 

 

Pacific Island Ecosystems at Risk. 2003. Furcraea 

foetida (L.) Haw., Agavaceae. http;//www.hear. 

org/pier_v3.3/fufoe.htm. 3 p. 

Schlegel, R. 2003. Plant breeding update: crop 

plants II. http://www.desicca.de/plant_breeding/ 

Crop_plants/Crops_II/hauptteil_crops_ii.html. 

[not paged.]. 
352 

Gesneria pedunculosa (DC.) Fritsch arbol de navidad 

GESNERIACEAE 

Synonyms: Conradia pedunculosa A.P. DC. 

 Gesneria exogenia Sassé & Moc. ex A.P. DC. 

 Pentarhaphia pedunculosa (A.P. DC.) Decne. 

 Codonoraphia albiflora (Decne.) Oersted 

John K. Francis 

 

General Description.—Arbol de navidad is an 

upright, evergreen shrub or occasionally a small 

tree 3 to 5 m in height and 5 to 10 cm in basal 

diameter. On rocks, the shrub sometimes takes a 

natural bonsai form. The root system, which may 

or may not have a taproot, takes whatever form 

necessary to anchor itself in the rocky ground 

where it grows. The numerous fine roots have 

clearly visible, white mycorrhizal growth. There 

are usually multiple stems from the base of the 

trunk. Arbol de navidad bark is gray or brown, 

furrowed and rough. The inner bark is light brown 

and fibrous. Twigs are slender, greenish gray, and 

form a relatively dense crown. The leaves, which 

are crowded near the ends of branches, have 

elliptic to obovate blades, 4 to 11 cm long, with 

smooth to fine serrate margins. The inflorescences 

usually have 2 to 4 flowers borne at the end of 

glandular-resinous peduncles 8 to 16.5 cm long. 

The five-lobbed tubular corolla is yellow-green to 

pinkish-white and 1.1 to 1.9 cm long. The fruit that 

forms is a four-valved, 6 to 9 mm capsule with five 

long, narrow calyx lobes that release numerous tiny 

reddish-brown or brown seeds (Liogier 1995, Little 

and others 1974). 

Range.—Arbol de navidad is endemic to Puerto 

Rico (Liogier 1995, Little and others 1974). It has 

been planted elsewhere in botanic gardens and 

arboretums but is not reported to have naturalized. 

A hybrid between arbol de navidad and G. 

pedicellaris Alain, dubbed “golden oldie,” has 

been developed for ornamental purposes 

(American Gloxinia and Gesneriad Society 2002). 

Ecology.—Arbol de navidad is locally common 

and usually grows on steep slopes and shallow, 

rocky soils, sometimes in fractured rock. The 

parent materials of the soils may be sedimentary 

(including limestone), igneous, and metamorphic 

(including ultramafic) rocks. The species is found 

as scattered individuals or in nearly pure thickets. 

Plants are less common on south slopes than on 

north-, east-, or west-facing slopes in areas that 

receive from about 1000 to 3000 mm of 

precipitation from about 50 to 800 m above sea 

level. Arbol de navidad is moderately shade 

tolerant. It grows and flowers in openings and 

below forest canopies, if they are not too dense. 

The species does not appear to compete in dense 

stands of grass. It grows in remnant forest stands 

and advanced secondary forests, some of which are 

old road cuts, abandoned pastures, and fields. 

Reproduction.—Arbol de navidad flowers and 

fruits nearly throughout the year. The flowers are 

probably pollinated by bats (Little and others 

1974). Seed production is high and consistent from 

year to year. Samples of seed collected in Cayey 

and Susúa, Puerto Rico, in different years weighed 

an average of 1.78 x 10-5g and 1.75 x 10-5g, 

respectively, or 56 million seeds/kg. Both 

collections failed to germinate when placed on 

moist filter paper. The seeds are dispersed by the 
353

wind. Seedlings are relatively common on bare 

ground near fruiting adult plants. Adult and sapling 

plants sprout when cut or damaged. 

Growth and Management.—Arbol de navidad is 

slow growing at all stages. It probably lives several 

decades. No plantations are known, and 

recommended management practices have not been 

published. Disturbance of the soil surface in steep, 

rocky terrain in moist areas near seed sources 

would likely result in seedling establishment. 

Benefits.—Arbol de navidad is a pretty plant that 

adds to the aesthetics of the forest. It helps protect 

the soil in fragile areas and furnishes cover for 

wildlife. Because of the small diameters and 

quantities in which it is available, the wood, which 

is light brown and hard, is useful mainly for fuel. 

References 

American Gloxinia and Gesneriad Society. 2002. 

Registered gesneriads. American Gloxinia and 

Gesneriad Society, Salsbury, CT. http:// 

www.aggs.org/ir_ges/. 7 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 
354 

Gonzalagunia hirsuta (Jacq.) K. Schum. rabo de ratón 

RUBIACEAE 

Synonyms: Barleria hirsuta (Jacq.) Jacq. 

Coccocypselum spicatum (Lam.) HBK 

 Duggena hirsuta (Jacq.) Britton & Wilson 

 Duggena richardii West 

 Duggena spicata (Lam.) Gómez 

 Gonzalagunia spicata (Lam.) DC. 

 Gonzalagunia coccocypselum C. & S. 

 Gonzalia spicata (Lam.) Standl. 

Heydyotis secunda Spreng. ex Schultes 

 Justicia hirsuta Jacq. 

Lygistum spicatum Lam. 

John K. Francis 

General Description.—Rabo de ratón, also 

known as mata de mariposa, palo pelado, yerba 

pelada, bios-foufou, and bois colibri (Howard 

1989, Liogier 1997), is a suberect, medium-sized 

shrub. It has membranous, opposite, ovate to 

lanceolate leaves, 6.5 to 19.0 cm in length and 2.0 

to 8.5 cm in width (Howard 1989) with 

pronounced, sunken veins. Plants usually have 

single straight stems but begin branching near the 

ground. The branches and twigs are slender and 

sometimes extend a meter horizontally. 

Range.—The natural range of rabo de ratón 

includes Hispaniola, Puerto Rico, the Lesser 

Antilles, Trinidad, Venezuela, the Guayanas, and 

northern Brazil (Howard 1989). The species is not 

known to have naturalized elsewhere. 

Ecology.—Rabo de ratón is intolerant and requires 

disturbance to grow and reproduce. It is most 

frequently found in brushy pastures, in early 

secondary forests after abandonment of 

cultivation, in tree-fall gaps and other disturbed 

areas in advanced secondary forests, and along 

roads. In a Puerto Rican study (Myster and Walker 

1997), rabo de ratón was the fifth most common 

species colonizing recent landslides. Dense 

thickets sometimes form. The species grows in 

areas receiving from about 1100 to 3400 mm of 

annual rainfall and at elevations ranging from near 

sea level to more than 600 m. Apparently, all types 

of soils are colonized, if not excessively well 

drained or very poorly drained. The root system is 

shallow, much branched, and extensive. 

Reproduction.—Rabo de rató begins flowering 

near the end of its first year. The inflorescences, 

which arise from leaf axils, are spiciformpaniculate and up 56 cm in length. The flowers are 

white with four or five lobes (Liogier 1997). 

Flowering and fruiting begin at the base and 

progress toward the tip. The flowers are pollinated 

by butterflies and small insects. When the 

inflorescence is laden with fruit progressing in size 

from base to tip, it resembles a mouse’s tail—

hence the name, rabo de ratón (Vélez and von 

Overbeek 1950). An inflorescence may contain 

100 or more fruits. The fruits are two-seeded 

fleshy drupes that are white, turning pink, violet, 

or blue (Howard 1989) and are available most of 

the year. The fruits in a Puerto Rican collection 

averaged 0.0885 + 0.0419 g and contained seeds 

numbering 182,000/kg. Seeds in this collection 

failed to germinate (Francis and Rodríguez 1993). 

Another collection from Puerto Rico had between 

55 and 68 percent germination in a field test. Of 

those failing to germinate, about 5 percent were 

lost to pathogens. There was no significant 
355

predation (Myster 1997). The seeds are dispersed 

by birds (Parrotta 1995). 

Growth and Management.—Rabo de ratón is 

fast growing and may reach 4 m in height and 3 

cm in lower stem diameter. Individual plants 

usually live 1 to 2 years (Vélez and von Overbeek 

1950). The species can probably be established by 

traditional methods (planted seedlings and direct 

seeding), but no plantations have been reported. 

Natural reproduction may be encouraged by 

cultivation near seed sources. 

Benefits.—Rabo de rató is beneficial to the 

environment in that it readily invades and helps to 

stabilize disturbed areas. Because the shrub forms 

a thin canopy, it does not inhibit the invasion of 

later successional vegetation. While probably not 

suitable for formal gardens, rabo de rató would 

contribute to wildflower gardens, forest parks, and 

greenbelts. The species is an abundant producer of 

fruits that are eaten by birds.

References 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

 Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Dicotyledoneae. 

Part 3. Vol. 6. Arnold Arboretum, Harvard 

University, Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p.

Myster, R.W. 1997. Seed predation, disease and 

germination on landslides in neotropical lower 

montane wet forest. Journal of Vegetation 

Science 8: 55-64. 

Myster, R.W., and L.R. Walker. 1997. Plant 

successional pathways on Puerto Rican 

landslides. Journal of Tropical Ecology 13: 165-

173. 

Parrotta, J.A. 1995. Influence of overstory 

composition on understory colonization by 

native species in plantations on a degraded 

tropical site. Journal of Vegetation Science 6: 

627-636. 

Vélez, I., and J. von Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Río Piedras, PR. 497 p. 
356 

Gossypium hirsutum L. wild cotton 

MALVACEAE 

Synonyms: Gossypium religiosum L. 

 Gossypium latifolium Murry 

 Gossypium punctatum Schum. 

 Gossypium taitense Parl. 

 Gossypium mexicanum Tod. 

 Gossypium marie-galante Watt 

John K. Francis 

General Description.—Wild cotton, also known 

as upland cotton, in wild populations is a shrub or 

occasionally a small tree to 5 m in height and 15 

cm in stem diameter. The plant is deeply rooted 

with a taproot and laterals. Wild cotton usually has 

a single trunk, unless disturbed, but may have 

branches arising just above the ground. The bark is 

gray and smooth, but tough and stringy. The stem 

wood of older plants is of medium density and 

brittle. The twigs are relatively stout. The leaves on 

young plants are usually without lobes; older plants 

usually have three lobes or occasionally are deeply 

incised with five lobes. The light green leaves have 

petioles 3 to 8 cm long and blades 10 to 15 by 12 to 

15 cm. Bracteoles are pointed and may have one to 

seven or more teeth. The flowers of wild cotton are 

large and showy with cream to pale yellow petals, 

sometimes with a red spot at the base of the petals. 

The capsules have three to five valves with a 

smooth surface and many black gland dots and 

yield white or brown lint with seeds embedded. 

The seeds vary from black and smooth to green 

with tightly adhering fuzz (Howard 1989, Liogier 

1994, Watt 1907). In certain instances (Little and 

others 1974), wild cotton has been misidentified as 

Gossypium barbadense L., which occurs today as a 

residual from cultivation and as a true wild plant 

only on the coast of Ecuador (Izuko Museums of 

Cape Town 2000). Wild cotton occurs in at least 

three botanical varieties. The most commonly seen 

in the West Indies is G. hirsutum var. mariegalante (Watt) J.B. Hutchinson (Liogier 1994). 

Doubtless a legacy of agricultural cultivation, wild 

populations in Puerto Rico have a wide range of 

leaf and bract shapes, seed characteristics, and lint 

length and color (author’s observation). In fact, 

wild cotton is a tetraploid. There is a theory that 

wild cotton is the result of natural hybridization 

between Old World diploid cottons, G. arborium

L. or G. herbacium L. and G. raimondii Ulbrich. 

(Bajaj 1998). 

Range.—The native range of wild cotton includes 

Tropical Florida, the West Indies, Northern South 

America, Mexico, and Central America (Howard 

1989, Liogier 1994, Long and Lakela 1976) and 

apparently several Pacific Islands including Samoa 

(Kohel and Lewis 1984). Because of eradication 

intended to control the pink bole worm, the species 

has become very rare and has been listed as 

endangered in Florida (Nelson 1996). Gossypium 

hirsutum is the principal cultivated cotton 

throughout the world (Vijayakumar 2002). It has 

naturalized in many locations, especially in island 

habitats, including Hawaii (Neal 1965). 

Ecology.—Wild cotton is mainly a plant of the 

coastal strand and lower coastal plains. The species 

has a particular affinity for small islands. It also 

grows in disturbed places, particularly along roads 

and on river overflow areas, well inland. Wild 

cotton can grow in almost all types of well-drained 

soils. Optimum pH’s are said to be from 5.2 to 7.0 
357

(Center for New Crops and Plants Products 2002). 

Soil disturbance is usually necessary for its 

establishment. It tolerates moderate amounts of 

salts in the soil and salt spray. Wild cotton 

competes well with weeds and grasses but does not 

tolerate shade and cannot grow under tree cover. In 

Puerto Rico, wild cotton is most competitive at low 

rainfalls (750 to 900 mm of annual precipitation) 

but grows in well-drained areas that receive up to 

1700 mm of precipitation at from near sea level to 

about 300 m elevation. Wild cotton does not 

tolerate frost. 

Reproduction.—Wild cotton is dormant during 

the winter dry season and becomes vegetative as 

the rains begin again in early summer. At the end 

of the summer, vegetative growth ceases and plants 

begin to flower and fruit (Kohel and Lewis 1984). 

Three collections of wild cotton seeds of Puerto 

Rico from three locations averaged 0.088 + 0.001, 

0.075 + 0.001, and 0.081 + 0.001 g/seed or about 

12,000 seeds/kg. Germination is epigeal. Seeds 

from two of the collections were sown without 

pretreatment in commercial potting mix. One 

collection germinated at 100 percent starting in 17 

days and ending in 41 days. The second group 

germinated at 67 percent starting in 22 days and 

ending in 140 days. The wind disburses the seeds 

over short distances by blowing the lint until it 

catches on vegetation. Birds also move seeds when 

they use cotton lint for nest building materials. 

Under greenhouse conditions, seedlings may grow 

more than 1 m in 6 months. In wild stands, 

seedlings can be abundant but they naturally thin 

themselves quickly. 

Growth and Management.—Although wild 

cotton in its cultivated form is managed as an 

annual, it is a true perennial, and if allowed to do 

so under favorable conditions, will live for several 

years (Kohel and Lewes 1984) and become tall and 

woody. Wild cotton in Puerto Rico grows 1 to 1.5 

m in its first year. The wild variety requires 15 or 

more nodes to be formed along the main axis 

before it can flower. Consequently, the plant is 

rarely able to bloom during its first year (Kohel and 

Lewis 1984). Cotton can be transplanted using 

containerized nursery plants. However, direct 

seeding following removal of overhead 

competition and cultivation of the soil is 

recommended. Soil temperatures must be at least 

18 °C before planting (Center for New Crops and 

Plant Products 2002). 

Benefits.—About 87 percent of the cotton grown 

commercially is G. hirsutum (Vijayakumar 2002). 

It has been selected, bred, and, in some cases, 

hybridized to give the manageable and productive 

varieties planted today. Evidence from Mexico 

shows that wild cotton has been used by man for 

making string and cloth since at least 200 BC 

(Iziko Museums of Cape Town 2000). Today, 

cotton is the world’s most widely used natural 

fiber. The seeds of cultivated cotton form the basis 

for an important oil and animal food industry. The 

seeds contain 7.3 g water, 23.1 g protein, 22.9 g 

fat, 43.2 g total carbohydrate (16.9 g fiber), and 3.5 

g ash per 100 g (Center for New Crops and Plant 

Products 2002). Gossypol, a sesquiterpinoid, is 

present in vegetative parts of wild cotton as well as 

the cultivated varieties. It is a natural deterrent to 

insects but is toxic to man and nonruminant 

animals. It has been shown to have strong 

antifertility, antitumor, antiparasite, and anti-HIV 

properties (Bajaj 1989). Gossypol is widely used in 

China as a male contraceptive (Center for New 

Crops and Plant Products 2002). Raw cottonseed 

oil (containing gossypol) used alone showed 

promise as an insecticide (Tadas and others 1994). 

In herbal medicine, cottonseed and roots have been 

used to treat nasal polyps, asthma, diarrhea, 

hemorrhoids, dysentery, uterine fibroids and 

certain cancers, to induce abortions, and as a 

diuretic. (Center for New Crops and Plant Products 

2002). Wild cotton is occasionally planted as an 

ornamental. The species contributes to biodiversity, 

soil stability, wildlife habitat, and scenic beauty in 

its beach strand and lowland habitat. 

References 

Bijaj, Y.P.S. 1998. Cotton. In: Y.P.S. Bijaj, ed. 

Biotechnology in Agriculture and Forestry 42. 

Springer-Verlag, New York. p. 3-36. 

Center for New Crops and Plant Products. 2002. 

Gossypium hirsutum L. Purdue University. 

http://www.hort.perdue.edu/newcrop/duke_ener

gy/Gossypium_hirsutum.html. 6 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Iziko Museums of Cape Town. 2000. Gossypium

(cotton). Biodiversity explorer. http://www. 

museums.org.za/bio/plants/malvaceae/gossypiu

m.htm. 2 p. 

Kohel, R.J. and C.F. Lewis, eds. 1984. Cotton. 

Agronomy 24. American Society of Agronomy, 
358 

Inc., Crop Science Society of America, Inc., and 

Soil Science Society of America, Inc., Madison, 

WI. 605 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books. Miami, FL. 

962 p. 

Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub. 

50. Bernice P. Bishop Museum Press, Honolulu, 

HI. 924 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Sarasota, FL. 391 p. 

Tadas, P.L., H.K. Kene, and S.D. Deshmukh. 

1994. Effect of raw cottonseed oil against 

sucking pests of cotton. PKV Research Journal 

18(1): 142-143. 

Vijayakumar, T. 2002. Cotton. http://www. 

geocities.com/vijayakumar777/cotton1.html. 

14 p. 

Watt, Sir G. 1907. The wild and cultivated cotton 

plants of the world. Longmans, Green, and Co., 

London. 406 p. 
359

Gouania lupuloides (L.) Urban chew-stick 

RHAMNACEAE 

Synonyms: Banisteria lupuloides L. 

 Rhamnus domingensis Jacq. 

 Gouania domingensis (Jacq.) L. 

 Gouania glabra Jacq. 

 Lupulus lupuloides Kuntze var. domingensis (Jacq.) Kuntze 

John K. Francis 

General Description.—Chew-stick, also known as 

toothbrush tree, bejuco de sopla, bejuco de indio, 

mascapalo, bejuco de fuego, liane-savon, and 

lyenn-savon, is a scrambling and climbing woody 

vine that may reach 7 to 12 m of extension. The 

plant is supported by an extensive system of 

medium and fine, white, lateral roots. The stem and 

branches are relatively slender and support 

themselves by means of coiled tendrils. The 

alternate leaves have petioles 0.5 to 2 cm long and 

thin, ovate to elliptic blades 4 to 10 cm long with a 

serrate or wavy-toothed margin and a pointed tip. 

Tiny white, yellow, or greenish flowers are borne 

on slender axillary or terminal racemes. Few-tomany, 6- to 13-mm, gray, three-winged fruits 

(schizocarps splitting into three mericarps) develop 

later in the season. Each mericarp contains a 3- to 

4-mm dark-brown seed (Acevedo-Rodríguez 1985, 

Croat 1978, Howard 1989, Liogier 1994, Stevens 

and others 2001). 

Range.—The native range of chew-stick includes 

Florida, the West Indies, Mexico, Central America, 

and South America into Brazil and Colombia 

(Howard 1989, New York Botanical Garden 2002, 

Pérez-Arbelaez 1978, Secritaria de Medio 

Ambiente y Recursos Naturals 2002). It is not 

known to have been planted or naturalized 

elsewhere. 

Ecology.—In Puerto Rico, Chew-stick occurs in 

forests that receive from 750 to over 2000 mm of 

precipitation. It grows on soils with the entire range 

of textures, both poor and rich in nutrients, 

excessively well drained to somewhat poorly 

drained over sedimentary (including limestone), 

igneous, and metamorphic (including ultramafic) 

rocks. The species is found from near sea level to 

elevations of 1,300 m (Acevedo-Rodríguez 1985, 

Stevens and others 2001). Chew-stick has an 

intermediate tolerance of shade. It can live and 

grow slowly in all but dark forest understories but 

requires increased sunlight to flower and fruit. The 

species grows on roadsides, in brushy pastures, and 

in secondary and remnant forests. In Florida, chewstick grows in mangroves and coastal hammocks 

(Nelson 1996). 

Reproduction.—In Puerto Rico, chew-stick 

blooms from August to March, and fruit matures 

from November to March (Acevedo-Rodríguez 

1985). In Panama, it flowers in the early dry season 

(November to March), and the seeds are dispersed 

from February to May (Croat 1978). Fruits 

collected in Puerto Rico averaged 0.0374 + 0.0005 

g/fruit. Seeds separated from them averaged 0.0052 

+ 0.0001 g/seed or 192,000 seeds/kg. Seed should 

be collected by hand from fruiting vines. Hand 

separation is practical for small lots; larger lots 

could be pulverized and winnowed. Fifty-seven 

percent of untreated seed from the previously 

mentioned collection germinated between 16 and 

75 days after sowing on commercial potting mix 

(author’s observation). Seed is dispersed by its 

lateral tumbling action in still air or wind and by 

lateral extension on the vines. Seedlings and young 

plants are relatively common in Puerto Rico. The 

plant sprouts after fire and cutting. 

Growth and Management.—Once established, 
360 

chew-stick grows relatively rapidly. In the presence 

of seed sources, reproduction is usually adequate. 

Thinning the overstory would probably allow 

existing suppressed plants to grow to larger sizes 

and fruit. 

Benefits.—Chew-stick contributes to biodiversity 

and biomass accumulation, helps protect the soil, 

and furnishes wildlife cover in forests where it 

grows. It is an important honey plant (Townsend 

1984). In Jamaica and the West Indies, stem 

cuttings are used to clean teeth and gums and 

extracts are used to manufacture a dentifrice 

(Secretaría de Medio Ambiente y Recursos 

Naturales 2002). Triterpenoid saponins may 

contribute to the beneficial activity (Kennelly and 

others 1993). In Colombia, the plant is used for a 

bitter flavoring in alcoholic beverages and to treat 

water retention and stomach problems (PérezArbelaez 1978). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Croat, T.B. 1978. Flora of Barro Colorado Island. 

Stanford University Press, Stanford, CA. 943 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Kennelly, E.J., W.H. Lewis, R.E.K. Winter, S. 

Johnson, M. Elvin-Lewis, and J. Gossling. 1993. 

Triterpenoid saponins from Gouania lupuloides. 

Journal of Natural Products 56(3): 402-410. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

 391 p. 

New York Botanical Garden. 2002. Flora and 

mycota of Acre, Brazil. http://nybg.org/ 

bsci/acre/rhamnaceae.html. [not paged]. 

Pérez-Arbelaez, E. 1978. Plantas útiles de 

Colombia. Litografía Arco, Bogotá, Colombia. 

831 p. 

Secretaría de Medio Ambiente y Recursos 

Naturales. 2002. Especies con usos no 

maderables en bosques tropicales y 

subtropicales. http://www.semarnat.gob.mx/ 

pfnm2/fichas/gouania_lupuloides.htm. 2 p. 

Stevens, W.D., C. Ulloa-Ulloa, A. Pool, O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanic Garden Press, St. Louis, 

MO. 2,666 p. 

Townsend, G.F. 1984. Multi-purpose trees for 

honey production. Echo technical note. 

http://www.echonet.org/tropicalag/technotes/Mu

ltiPur.pdf. 3 p. 
361

Grayia spinosa (Hook.) Moq. spiny hopsage 

CHENOPODIACEAE 

Synonyms: Atriplex spinosa Collotzi 

 Atriplex grayia Collotzi 

Grayia polygaloides Hook. & Arn. 

Nancy L. Shaw, Marshall R. Haferkamp, and Emerenciana G. Hurd 

 

General Description.—Alternative common 

names for spiny hopsage are grayia, Gray’s 

saltbush, saltbrush, spiny-sage, horsebrush, and 

applebush (Shaw and others 2001, Welsh and 

others 1987). Shrubs are 0.3 to 1.2 (1.5) m tall, 

summer deciduous, and erect to rounded with 

divergent, thorn-tipped branches and gray to 

brownish exfoliating bark. Young twigs and 

herbage are scurfy to pubescent, becoming 

glabrous with age. Alternate, entire, gray-green, 

fleshy leaves, 5 to 30 (40) mm long and 2 to 13 

mm wide, develop from prominent, globose, 

axillary buds. Blades are linear-oblanceolate with 

sessile or short-petiolate bases. The species is 

tetraploid (4x = 36) (McArthur and Sanderson 

1984). Inflorescences develop on floral shoots that 

die back following fruit dispersal. Flowers are 

inconspicuous. Staminate flowers develop in 

glomerate spikes and consist of a four or five 

lobed perianth and four or five stamens. Pistillate 

flowers are produced on dense terminal spicate 

inflorescences. Some flowers are vestigial. Each 

flower consists of a pistil enclosed in two 

obcompressed chordate bracteoles united along 

their margins except for a minute apical opening. 

Bracteoles enlarge at maturity to form a papery 

sac, 9 to 15 mm in diameter with a dorsally 

winged margin. Fruits are utricles with the papery 

pericarp free from the seed. Seeds are 1 to 2 mm in 

diameter. The testa is comprised of a thin, dark 

brown outer layer and an elastic inner layer. A 

well-developed embryo with an inferior radicle 

encircles the perisperm (Hitchcock and others 

1964, Shaw and others 1996, 2001). 

Range.—Spiny hopsage is widely distributed in 

the interior Western United States. It occurs east of 

the Cascade and Sierra Nevada Mountains from 

central Washington to southern California and 

eastward from southern Montana to western 

Colorado and northern Arizona (Hitchcock and 

others 1964, Welsh and others 1987). 

Ecology.—Spiny hopsage occurs at elevations 

from 160 to 2,130 m in Wyoming big sagebrush 

(Artemisia tridentata Nutt. ssp. wyomingensis

Beetle & A. Young), salt desert shrub, Mohave 

Desert, and pinyon-juniper communities receiving 

125 to 300 mm of annual precipitation (Welsh and 

others 1987). It grows on soils that are silty to 

sandy, frequently high in calcium, and neutral to 

strongly basic. It also occurs on sand dunes. Spiny 

hopsage generally occurs in small patches or as 

scattered plants. Extensive stands are rare. Plants 

initiate growth in early spring and rapidly 

complete their vegetative and reproductive cycles 

prior to entering summer dormancy. They are most 

tolerant of summer wildfires after summer leaf fall 

and can resprout from surviving root crowns 

(Daubenmire 1970). Regeneration is episodic. 

Germination occurs in early spring. Most seedlings 

establish beneath canopies of spiny hopsage or 

other shrubs. Growth of cheatgrass (Bromus 

tectorum L.), a highly competitive annual, 

however, is favored by nutrient accumulation 

beneath spiny hopsage canopies. This can interfere 
362 

with natural regeneration of native vegetation and 

result in increased fine fuel accumulation. 

 

Reproduction.—Most plants are dioecious, but 

the proportion varies among populations. Flowers 

appear in February to May and are wind 

pollinated. Fruits ripen in March to June (Shaw 

and others 2001) and are dispersed by wind, 

gravity, and insects. Herbage, flower, and fruit 

production are highly variable among years 

(Rickard and Warren 1981); many plants fail to 

produce flowers in dry years (McArthur and 

Sanderson 1984). During a drought, spiny hopsage 

in a southern Idaho wildland planting began 

flowering 4 years after seeding. Incubating 

debracted utricles at 15 and 5 o

C (8 hrs, 16 hrs) or 

at 15 o

C for 14 days provides an estimate of 

germination. A 30 to 60 day prechill at 3 to 5 oC is 

required to enhance germination of utricles from 

northern populations. Shaw and others (2001) 

reported an average of 1,030,600 debracted 

utricles/kg. Germination is epigeal. 

Growth and Management.—Seedlings develop 

rapidly if adequate water is available. Mature 

plants are drought tolerant. Spiny hopsage is 

considered one of the most palatable shrubs in salt 

desert shrub communities in spring and early 

summer. Its use by wildlife and livestock declines 

rapidly as plants enter summer dormancy. 

Although fairly tolerant of grazing, some 

populations receiving heavy use have been 

replaced by less palatable species (Blaisdell and 

Holmgren 1984). 

Benefits.—Spiny hopsage contributes to diversity 

on native sites that generally support few other 

woody species. It provides cover for birds and 

other small animals, spring and early summer 

forage for big game and livestock, and soil 

stabilization on moderate slopes. Litter rich in 

potassium and other accumulated cations 

enhances growth and nutrient content of plants 

growing beneath its canopy (Rickard and Keough 

1968). Native Americans ground parched seed of 

spiny hopsage to prepare pinole flour 

(Stubbendieck and others 1986). 

References 

Blaisdell, J.P. and R.C. Holmgren. 1984. 

Managing Intermountian rangelands—salt desert 

shrub ranges. General Technical Report INT163. U.S. Department of Agriculture, Forest 

Service, Intermountain Research Station, Ogden, 

UT. 52 p. 

Daubenmire, R. 1970. Steppe vegetation of 

Washington. Technical Bulletin 62. Washington 

State University, Pullman, WA. 131 p. 

Hitchcock, C.L., A. Cronquist, M. Ownbey, and 

J.W. Thompson. 1964. Vascular plants of the 

Pacific Northwest. Part 2. Flora of the Pacific 

Northwest. University of Washington Press, 

Seattle, WA. 730 p. 

McArthur, E.D. and S.C. Sanderson. 1984. 

Distribution, systematics, and evolution of the 

Chenopodiaceae, In: Proceedings—symposium 

on the biology of Atriplex and related 

chenopods. General Technical Report INT-172. 

U.S. Department of Agriculture, Forest Service, 

Intermountain Research Station, Ogden, UT. p. 

14-24. 

Rickard, W.H. and R.F. Keough. 1968. Soil-plant 

relationships of two steppe desert shrubs. Plant 

and Soil. 19:205-212. 

Rickard, W.H. and J.L. Warren. 1981. Response of 

steppe shrubs to the 1977 drought. Northwest 

Science. 55:108-112. 

Shaw, N.L., M.R. Haferkamp, and E.G. Hurd. 

2001. Grayia Hook. & Arn. hopsage. In: F.T. 

Bonner, and R.G. Nisley, eds. Woody Plant 

Seed Manual. U.S. Department of Agriculture, 

Forest Service, Washington, D.C. 

http://wpsm.net/ 12 p. 

Shaw, N.L., E.G. Hurd, and M.R. Haferkamp. 

1996. Spiny hopsage fruit and seed morphology. 

Journal of Range Management. 49:551-553. 

Stubbendieck, J., S.L. Hatch, and K. Hirsch. 1986. 

North American range plants. University of 

Nebraska Press, Lincoln. 465 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 
363

Guettarda elliptica Sw. velvetseed 

RUBIACEAE 

Synonyms: Guettarda bladgettii Shuttlew. ex Chapm. 

 Matthiola elliptica (Sw.) Kuntze 

John K. Francis 

General Description.—Velvetseed, also known as 

Everglade velvetseed, pickle-wood, cucubano liso, 

cigüillo, cuero de sabana crucillo, guayabillo 

prieto, cabrito, and punteral, is a shrub or small tree 

to 6 m in height and 10 cm in stem diameter. It 

usually has multiple stems from below or just 

above the ground level. The stem bark is gray or 

gray-brown, smooth with light colored dots or 

lines. The stem wood is light brown, hard, finetextured, and heavy with a specific gravity of 0.83 

(Little and others 1974). The plant is supported by 

hard, brittle, dark-brown roots. Velvetseed tends to 

form branchy crowns of opposite, moderately fine 

branches and twigs. The opposite leaves have 

short, hairy petioles and oval to elliptic blades 3 by 

7.5 cm long. Axillary, few-flowered cymes bear 

small tubular yellowish-white flowers. Three to 4 

months later, globose, 4- to 8-mm drupes with a 

persistent calyx develop. They turn red to black at 

maturity and contain two to four seeds enclosed in 

a bony shell (Liogier 1997, Little and others 1974, 

Nelson 1996, West and Arnold 1952). The fruits 

have a slightly sweet, slightly sour, and slightly 

astringent flavor (author’s observation). 

Range.—The native range of velvetseed extends 

from southern Florida through the West Indies, 

from Mexico through Central America, and 

Venezuela and Guyana in South America (Little 

and others 1974). 

Ecology.—Velvetseed grows in pinelands and 

hammocks in Florida (Nelson 1996). In Puerto 

Rico, it is most common in limestone areas and 

occasionally in areas of igneous rock, on rocky 

ridge sites too difficult for tall vegetation, and in 

coastal thickets and rocky hills (Liogier 1997, 

Little and others 1974). Annual rainfall varies from 

750 mm to 2200 mm, and elevations range form 

near sea level to about 400 m. Vegetation sites 

include relatively pristine to highly disturbed 

remnant forests and late secondary forests. 

Velvetseed is moderately intolerant of shade. It 

competes well with slow-growing vegetation but 

not with tall grass or overtopping trees. Opengrown plants assume a symmetrical shape, but in 

thickets, the stems are usually crooked and the 

crowns asymmetrical. 

Reproduction.—Velvetseed blooms in summer in 

Florida (West and Arnold 1952) and in the spring 

in Puerto Rico (Little and others 1974). Flowers 

require pollination to develop seeds and are 

ordinarily pollinated by insects (Koptur and 

Richards 1996). A collection of fruits in Puerto 

Rico averaged 0.350 + 0.008 g/fruit. Seeds 

separated from them weighed (air-dried) an 

average of 0.0626 + 0.0015 g/seed or 16,000 

seeds/kg. Sown on commercial potting mix, 68 

percent of the seeds germinated between 3 and 7 

months after sowing (author’s observation). 

Although there is usually good fruit and seed 

production, seedlings are relatively uncommon. 

Birds disperse the seeds. Velvetseed sprouts readily 

when cut. 

Growth and Management.—The Florida 

champion velvetseed is 7 m in height and 29 cm in 

diameter at breast height (Champion Tree Project 

2002). The species grows slowly, at least in the 
364 

difficult habitat where most occur. Individuals 

probably live from two to several decades. No 

management experience has been published. 

However, long-term protection of velvetseed sites 

is recommended. 

Benefits.—Velvetseed contributes to biodiversity 

and wildland biomass while it helps protects the 

soil from erosion and provides food and cover for 

wildlife. The species is listed as the larval food 

plant for the moths, Calidota strigosa Walker in 

Florida and the West Indies (Barns 2002) and 

Hylesia lineata Druce in Mexico (Pescador 1995). 

The wood is useful for fuel and stakes. Velvetseed 

is a pretty shrub and would make a fine foundation 

plant in seminatural landscaping. 

References 

Barns, M.J.C. 2002. Moths of Jamaica, Calidota 

strigosa. http://members.fortunecity.co.uk/ 

jamaicamoth/images/calstr.htm. 1 p. 

Champion Tree Project. 2002. National Champion 

Trees, Florida. http://www.championtrees.org/ 

database/championsFL.htm. 8 p. 

Koptur, S. and J.H. Richards. 1996. Comparative 

floral biology and breeding systems of 

smooth- and rough-leaved velvetseed, 

Guettarda elliptica Sw. and G. scabra in the 

Everglades. American Journal of Botany 

83(supplement): 70. 

 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de 

la Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

390 p. 

Pescador, A.R. 1995. Distribution and abundance 

of Hylesia lineata egg masses in a tropical dry 

forest in western Mexico. Southwestern 

Entomologist 20(3): 367-375. 

West, E. and L.E. Arnold. 1952. The native trees 

of Florida. University of Florida Press, 

Gainesville, FL. 212 p. 
365

Gundlachia corymbosa (Urban) Britt. ex Boldingh yambush 

ASTERACEAE 

Synonyms: Baccharis dioica sensu Duss 

 Gundlachia domingensis var. corymbosa Urban 

John K. Francis 

General Description.—Yambush, also known as 

horsebush, broombush, soldier’s-bush, Jamaican 

trash, sereno, and bois-Guillaume, is an evergreen 

shrub to 1.5 m in height. Stems are typically 

slender, numerous, branched near the ground, and 

rooted at intervals where they touch the ground. 

The plant is supported by brown lateral and fine 

roots that are flexible and weak. The leaves are 

mostly clustered near the twig ends. The subsessile 

leaves are fleshy, sticky, oblanceolate to oblonglanceolate, mostly rounded at the apex and 3 to 8 

cm long by 0.5 to 2 cm wide. The terminal 

inflorescences are dense corymbs of heads on 

short peduncles. The involucre is about 5 mm long 

and surrounds five to nine white florets. Fruits are 

oblong, angular achenes with tawny pappus 

(Britton and Millspaugh 1962, Howard 1989, 

Liogier 1997). 

Range.—Yambush is native to the Bahamas, 

Cuba, Hispaniola, Puerto Rico, the Virgin Islands, 

Barbuda, Saba, Monserrat, Guadeloupe, La 

Désirade (Guadaloupe), Curaçao, Aruba, and 

Bonaire (Howard 1989, Liogier 1997). It is not 

known to have been planted or naturalized 

elsewhere. 

Ecology.—Seaside and small island habitat is 

normal for the species. Yambush mostly grows on 

shoreline rocks and cliffs in Puerto Rico. It was 

collected between rocks and gravel deposited at 40 

to 50 m elevation in La Désirade (New York 

Botanical Garden 2003) and on calcareous silt 

with emergent limestone on Turks and Caicos 

(Darwin Institute 2002), and on rocky shores on 

San Salvador, Bahamas (Gerace and others 2003). 

Britton and Millspaugh (1962) indicate that it 

grows on sand dunes, pine lands, and borders of 

saline marshes. Yambush can grow in a wide 

variety of soils, gravel, and even in cracks in 

rocks. It is tolerant of salty soils, heavy salt spray, 

sea water overwash, and constant, sometimes high 

winds. Mean annual precipitation in its natural 

habitat varies from about 700 mm to 1,700 mm. It 

grows from near sea level to over 50 m in 

elevation. The species can grow with light 

overhead shade but generally grows where few 

other species can survive to avoid competition. 

Reproduction.—Yambush probably blooms 

throughout the year. The seeds are small, produced 

in great numbers, and dispersed by the wind. 

Seedlings are uncommon, but once established in 

favorable environments, they layer profusely and 

form interconnected mats. 

Growth and Management.—Yambush grows 

slowly. In seaside environments in Puerto Rico, it 

adds about 20 cm per year to its multiple stems. 

Individual plants and clumps apparently live for 

many years. The author knows of no reports of 

planting or management of the species. Because it 

is uncommon in much of its range, not aggressive, 

and grows in areas not normally used for 

agriculture or forestry, control of stands of the 

species should rarely be necessary. 

Benefits.—Yambush adds to the aesthetics of sites 

where it grows, helps protect the soil, and 
366 

furnishes cover for wildlife. Although there is no 

record of it having been used for ornamental 

purposes, yambush seems an excellent choice for 

groundcovers and borders in seaside and xeric 

gardens. Poultices of crushed leaves relieve and 

help heal allergic dermatitis (Wood 2003). 

References

Britton, N.L. and C.F. Millspaugh. 1962. The 

Bahama flora. The New York Botanical Garden, 

New York. 695 p. 

Darwin Institute. 2002. Plan for biodiversity 

management and sustainable development 

around Turks and Caicos Ramsar site: 

introduction. http://www.ukotcf.org/pdf/ 

TCIRamMP/Parts%20One%20and%Two.pdf. 

48 p. 

Gerace, D.T., G.K. Ostrander, and G.W. Smith. 

2003. San Salvador, Bahamas. Coastal region 

and small island papers 3. UNESCO, Rome. 

http://www.unesco.org/csi/pub/papers/gerace. 

htm. 20 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

New York Botanical Garden. 2003. Specimens 

detailed results: Gundlachia domingensis A. 

Gray var. corymbosa Urb. (syntype). 

http://scisun.nybg.org:8890/searchdb/owa/www

catalog.detail_list. 1 p. 

Wood, K.M. 2003. Medicinal plants for what ails 

you in Paradise. http://www.timespub.tc/ 

Natural%20History/Archive/Winter%20200102/

medicinal.htm. 2 p. 
367

Gutierrezia sarothrae (Pursh) Britt. & Rusby broom snakeweed

ASTERACEAE 

Synonyms: Xanthocephalum sarothrae (Pursh) Shinners 

 Solidago sarothrae Pursh 

Gutierrezia diversifolia Greene 

Gutierrezia linearis Rydb. 

Juanita A. R. Ladyman 

 

Illustration source: USDA-Forest Service collection, 

Hunt Institute 

General Description.—Gutierrezia sarothrae is a 

woody perennial named for Pedro Gutierrez, a 

Spanish nobleman. The epithet “sarothrae” is a 

derivative of a Latin word meaning “broom.” Its 

most popular common name is broom snakeweed, 

or snakeweed, but is also referred to as common 

matchbrush, broomweed, brownweed, yellow-top, 

yellow weed, resin-weed, turpentine-weed and, 

principally in Mexico, yerba de vibora (Dayton 

1931). It has also many botanical synonyms and 

for a complete list see Kartesz (1994). The Navajo 

Native Americans call it Ch’il diilyesiitoh, “big 

dodge weed” (Mayes and Lacy 1994). Broom 

snakeweed is generally 20 to 60 cm tall with a 

dense crown. The young, greenish stems turn tan 

and then brown with age. It has linear green leaves 

up to 3.8 cm long and numerous small, brightyellow flowers with two to seven rays that are 2 to 

3 mm long. In its northern range it is deciduous 

and dies back, but within its southern range it 

commonly only dies back part way in winter 

(Cronquist 1994, Whitson et al. 1991). Snakeweed 

contains saponin, a toxic secondary product 

(Whitson et al. 1991). Details of the plant may be 

viewed on the Texas A&M University “Virtual 

Herbarium” Website (2000). 

Range.—Broom snakeweed is native to the 

Americas. It is a very common shrub throughout 

Western North America, from Saskatchewan and 

Alberta, Canada south through western Nebraska 

and Kansas into Texas and central Mexico, and 

west to Washington, Oregon, California, and Baja 

California. Broom snakeweed has been described 

as a diagnostic climax member of desert 

grasslands (Dick-Peddie 1993). It is a member of 

both Chihuahuan and Sonoran desert grasslands 

(Schmutz et al. 1992). It has been estimated that it 

covers more acres than any other perennial plant 

on Navajo Tribal lands in the Southwestern United 

States (Mayes and Lacy 1994). 

Ecology.—Broom snakeweed grows in dry, open 

places from 732 m to 2,898 m (Tirmenstein 1999). 

As its wide range suggests, it grows in a variety of 

soil conditions, from sandy and well-drained loam 

to heavy clay soils and in various plant 

communities, including Douglas-fir and ponderosa 

pine forests, pinyon-juniper woodlands, sagebrush 

shrublands, and desert grasslands. Broom 

snakeweed root systems are fairly shallow, which 

is unusual for a shrub adapted to semi arid and arid 

environments (Schmutz and others 1992). They 

occupy the same soil layers as those of perennial 

grasses and sometimes appear to compete with 

them (Burgess 1995). Broom snakeweed tends not 

to resprout after fire and frequently dies during 
368 

drought (Schmutz et al. 1992). It suffers from 

drought to about the same extent as black grama 

grass, but when insect pressure is present the 

snakeweed mortality is much higher (Dick-Peddie 

1993). However, by virtue of its high seed 

production and rapid seedling establishment 

(Osman and Pieper 1988) it often increases, or 

“comes in”, after prolonged drought and other 

disturbances (Dayton 1931). 

Reproduction.—Snakeweed flowers from July to 

December depending upon the latitude and water 

availability. The plant is insect pollinated 

(Mayeux and Leotta 1981). It is a prodigious 

producer of seed that can remain viable in the soil 

for a considerable length of time (Osman and 

Pieper 1988). A single plant is capable of 

producing 9,000 to 10,000 seeds annually 

(Tirmenstein 1999). The oval fruit (seeds) are 

achenes and covered with stiff hairs or nearly 

glabrous. Chaffy scales, called the pappus, are on 

one end of the achene and aid seed dispersal by 

wind (Cronquist, 1994). Seeds are dispersed by 

animals as well as wind, although it has been 

reported that they also accumulate under the shrub 

canopy (Mayeux and Leotta 1981). Light enhances 

germination, which suggests that the best position 

for seeds is on or near the soil surface (Mayeux 

and Leotta 1981). 

Growth and Management.—Snakeweed is not 

palatable and can be poisonous to livestock and 

increases under grazing pressure (Cable 1975; 

Carter 1988; Cronquist 1994; Pase and Pond 

1964). Most losses in sheep and cattle are due to 

abortions (Whitson et al. 1991), but the plant also 

apparently affects the kidneys of both sheep and 

horses (Dayton 1931). Data suggest that the 

presence of jackrabbits may reduce cattle losses 

from poisonous plants because, under moderate 

stocking rates, jackrabbits consume increased 

amounts of poisonous plants and shrubs such as 

snakeweed but have minimal forage competition 

with cattle (Daniel et al. 1993). When abundant, 

snakeweed is generally considered an indicator of 

overgrazing (Dayton 1931; Hatch and Pluhar 

1993; Powell, 1998). Pastures are also not 

properly utilized when broom snakeweed is 

present amongst the grasses (Whitson et al. 1991). 

Snakeweed is killed by certain borers and other 

insects. Species from the genera, Crossidius, 

Diplotaxis, and Mecas have been proposed as 

possible biological controls (Dayton 1931). Fire 

can be used to reduce snakeweed (Humphrey 

1949; Tirmenstein 1999). However, the success of 

controlled burns depends upon fuel quantity and 

dryness, and an absence of grazing for at least one 

season prior to burning has been recommended 

(Tirmenstein 1999). In many cases, fire may 

prove an ineffectual control. For example, 5 years 

after a wildfire snakeweed had gone from 0 

percent canopy cover to over 40 percent canopy 

cover on monitored plots in Arizona (Pase and 

Pond 1964). In west-central Utah broom 

snakeweed was found on eight out of the nine ageclasses of burns, being absent on the 71-year-old 

burns and most abundant on the 22-year-old burns 

(Barney and Frischknecht 1974). Broom 

snakeweed is susceptible to some herbicides 

(Whitson et al. 1991). 

Benefits.—Broom snakeweed provides some 

stability to the soil against both wind and water 

erosion (Tirmenstein 1999, USDAFS 1988). It is 

fair browse for wildlife but is poisonous to 

livestock due to the saponin content, which is 

variable and depends upon both growth phase and 

environmental conditions (Hatch and Pluhar 

1993). The vegetation is most toxic during leaf 

formation (Whitson and others 1991). At times 

when there has been no alternative forage, 

snakeweed has been used by livestock with no ill 

effects (USDAFS 1988). Presumably saponin 

levels have been low at these times. It has several 

medicinal uses among the Navajo (Mayes and 

Lacy 1994). It is used to heal cuts and bites in 

humans and animals. A snakeweed medicine is 

given in childbirth and is used to treat 

stomachaches and other internal disorders. It is 

reported to be a powerful diuretic (Powell 1998). It 

is also used in various Navajo ceremonies 

including as an emetic and fumigent (Mayes and 

Lacy 1994). A yellow dye is made from the 

flowers, and the stems can be used to make fire by 

friction. Birds, for example quail, eat the seeds 

(Epple 1995). Broom snakeweed also provides 

cover and resting sites for species of small 

mammals, for example jackrabbits (Daniel and 

others 1993) and birds, for example lesser prairie 

chicken (USDAFS 1988) and Columbia basin 

burrowing owl (Green and Anthony 1989). 

References 

Barney, M.A. and N.C. Frischknecht. 1974. 

Vegetation changes following fire in the pinyonjuniper type of west-central Utah. Journal of 

Range Management 27(2): 91-96. 

Burgess, T.L. 1995. The dilemma of coexisting 

growth forms. In: M.P. McClaran and T.R. Van 
369

Devender, eds. The desert grassland. The 

University of Arizona Press, Tucson, AZ. 346 p. 

Cable, D.R. 1975. Range management in the 

chaparral type and its ecological basis: the status 

of our knowledge. Research Paper RM-155. 

U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Forest and Range Experiment 

Station. Fort Collins, CO. 30 p. 

Carter, J. L. 1988. Trees and Shrubs of Colorado. 

Johnson Books, Boulder, CO. 165 p.

Cronquist, A. 1994. Intermountain Flora, Vascular 

Plants of the Intermountain West, USA. Vol. 5. 

Asterales. New York Botanical Garden, Bronx, 

NY. 496 p. 

Daniel, A., J. L. Holechek, R. Valdez, A. Tem, L. 

Saiwana, M Rusco, and M. Carden. 1993. Range 

condition influences on Chihuahuan Desert 

cattle and jackrabbit diets. Journal of Range 

Management 46(4): 296-301. 

 Dayton, W. A. 1931. Important western browse 

plants. Miscellaneous Publication 101. U.S. 

Department of Agriculture, Washington, DC. 

214 p. 

Dick-Peddie, W.A. 1993. New Mexico Vegetation 

– past, present, and future. University of New 

Mexico Press, Albuquerque, NM. 244 p. 

Epple, A.O. 1995. A Field Guide to the Plants of 

Arizona. Falcon Press Publishing Co., Helena, 

Montana. 347 p. 

Green, G.A. and R.G Anthony. 1989. Nesting 

success and habitat relationships of burrowing 

owls in the Columbia Basin, Oregon. The 

Condor. 91: 347-354. 

Hatch, S.L. and J. Pluhar. 1993. Texas range 

plants. The W.L. Moody Jr Natural History 

Series 123. Texas A&M University Press, 

College Station, TX. 326 p. 

Humphrey, R.R. 1949. Fire as a means of 

controlling velvet mesquite, burroweed, and 

cholla on southern Arizona ranges. Journal of 

Range Management. 2: 175-182. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada, and 

Greenland. Vol. 1 – Checklist. 2nd ed. Timber 

Press, Portland, OR. 622 p. 

Mayes, V.O. and B.B. Lacy, 1994. Nanise’ Navajo 

herbal. Navajo Community College Press, 

Tsaile, AZ. 153 p. 

Mayeux, H.S., Jr. and L. Leotta. 1981. Germination 

of broom snakeweed (Gutierrezia sarothrae) and 

threadleaf snakeweed (G. microcephalum) seed. 

Weed Science. 29: 530-534. 

Osman, A. and Pieper, R.D. 1988. Growth of 

Gutierrezia sarothrae seedlings in the field. 

Journal of Range Management 41(1): 92-93. 

Pase C.P. and F.W. Pond. 1964. Vegetation 

changes following the Mingus Mountain burn. 

Note RM-18. U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research 

Station, Fort Collins, CO. 8 p. 

Powell, A. M. 1998. Trees & shrubs of TransPecos and adjacent areas. Big Bend Natural 

History Association. (Reprinted by University of 

Texas University Press, Austin, TX.) 498 p. 

Schmutz E.M., E.L Smith, P.R. Ogden, M.L. Cox, 

J.O. Klemmedson, J.J. Norris, and L.C. Fierro. 

1992. Desert Grasslands. In: R.T. Coupland, ed. 

Ecosystems of the World, 8A; Natural 

Grasslands Introduction and Western 

Hemisphere. Elsevier, London, NY. p. 337-362 

Texas A&M University “Virtual Herbarium”. 

Uvalde Research and Extension Center. 2000. 

Copyright Texas A&M University System. 

http://uvalde.tamu.edu/herbarium/gusa.htm [not 

paged]. 

Tirmenstein, D. 1999. USDA Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System, 

http://www.fs.fed.us/database/ [not paged]. 

Whitson, T.D., ed., L.C. Burrill, S.A. Dewey, D.W. 

Cudney, B.E. Nelson, R.D. Lee, and R. Parker. 

1991. Weeds of the West. The Western Society 

of Weed Science, Western U.S. Land Grant 

Universities Cooperative Extension Services and 

the University of Wyoming, Pioneer of Jackson 

Hole, Jackson WY. p 132-133. 

United States Department of Agriculture, Forest 

Service. 1988. Range Plant Handbook. Dover 

Publications, Inc. New York. 837 p. 
370 

Gynerium sagittatum (Aubl.) Beauv. wild cane 

POACEAE 

Synonyms: Saccharum sagittatum Aubl. 

 Arundo saccharoides Poir. 

 Airo gigantea Steud. 

John K. Francis 

General Description.—Wild cane, also known as 

bitter cane, uva grass, and caña brava, is a tall 

shrub with a grass-like habit. Its culms are usually 

5 or 6 m in height and 2 or 3 cm in diameter but 

may reach 10 m in height and 4 cm in diameter in 

Puerto Rico. The species varies from 5 to 14 m in 

height in the western Amazon Basin (Kalliola and 

others 1992). The culms arise from underground 

rhizomes which also produce weak and flexible 

lateral roots, mostly 1 mm or less in diameter. The 

culms have closely imbricated woody sheaths 

around a hard, woody exterior, and a fibrous 

interior. They are usually unbranched and taper 

little except near the top. The older leaves are shed, 

leaving a plainer, fan-like group near the apex. The 

leaf blades are 1 to 2 m long and have sharp 

serrulate margins. The clonal groups of plants are 

dioecious. The grayish-white plume-like terminal 

panicles are large, up to 2 m long. The male and 

female inflorescences are similar in appearance, 

but pistillate plants have a slightly fuzzy 

appearance because of hairy lemmas. The fruits are 

brown and about 1 mm long (Croat 1978, Howard 

1979, Pohl 1983, Stevens and others 2001). 

Range.—Wild cane is native to the West Indies 

except the Bahamas, and from Mexico through 

Central America and South America to Paraguay 

(Howard 1979). It is not known to have naturalized 

elsewhere. Two types coexist in the western 

Amazon Basin: a “small” and a “large” type that 

differ considerably in physical form and mode of 

reproduction (Kalliola and others 1992). The 

distinction is not reported elsewhere. 

Ecology.—Wild cane grows on sites with moist 

soils, usually high in organic matter, often with the 

water table near the surface. These sites are 

seasonally flooded areas such as lake shores, 

swamps, river flood planes, or sand bars. The 

species grows at elevations from 10 to 1,600 m 

above sea level in Costa Rica (Instituto Nacional 

de Biodiversidad 2002). Wild cane resists damage 

from moderate flooding and sprouts after being 

covered with sediment. “Large type” stands in the 

western Amazon region vary in density from 0.6 to 

2.6 culms/m2

. Forest edges “shade out” portions of 

wild cane stands, and occasional trees grow up 

through stands and eventually suppress culms 

growing under their crowns. The species affects the 

course of forest succession (Kalliola and others 

1991). Apparently, disturbance that creates bare, 

wet soil is necessary for seedling establishment. 

Reproduction.—In some environments flowering 

occurs throughout the year (Croat 1978); in others 

it occurs near the end of the low water period 

(Kalliola and others 1992). The species is 

apparently wind pollinated (Pohl 1983). There are 

1.67 million seeds/kg, and they can be expected to 

germinate between 3 and 7 days following sowing 

at temperatures between 20 and 30 °C (Association 

of Official Seed Analysis 2002). Almost all the 

seeds of the “short” type from the Amazon Basin 

germinated within 3 weeks, and 0 to 2 percent of 

the “large” type germinated (Kalliola and others 

1992). Seeds are dispersed by wind and water 

(Kalliola and others 1992). Vegetative propagation 

is also important, both for expanding colonies and 

establishing new ones. Horizontal runners or 

rhizomes, surface or underground, are constantly 

active and establish new plants or clumps as far as 
371

20 m from the parent plants (Pohl 1983). Segments 

of culm or rhizome, carried by floodwaters and 

covered with soil or debris, sprout and start new 

colonies. 

Growth and Management.—Growth of wild cane 

is rapid. Nursery seedlings reached 20, 30, and 50 

cm after 1, 2, and 4 months (Kalliola and others 

1992). How long seedlings take to reach maturity 

and how rapidly suckers grow is unknown. 

Theoretically, baring catastrophes and invasion and 

shading by trees, clones can endure indefinitely. 

Culms of Amazon Basin plants produced close to 

200 leaves during their lifetimes, having from 19 to 

28 living leaves at a time (Kalliola and others 

1992). Unbranched culms die after flowering, but 

only the branches of branched culms die. If not 

controlled, wild cane slowly invades wet 

bottomland pastures and eliminates forage plants. 

Periodic mowing appears to be adequate for control 

of advancing clumps. 

Benefits.—Wild cane provides cover for wildlife 

and protects stream banks from erosion (Kalliola 

and others 1992). Its culms lack the strength and 

toughness of hardwoods and bamboo but still are 

used in rude construction, drying racks, vegetable 

stakes and fruit props, and for weaving mats, 

baskets, and hats (Kalliola and others 1992). In the 

Amazon area, arrow shafts are made from the dried 

culms (Russo 2002). Plumes are used for dry floral 

arrangements. 

References 

Association of Official Seed Analysis. 2002. 

Reference base. http://www.aosaseed.com/ 

species/speciesG.pdf. [not paged]. 

Croat, T.B. 1978. Flora of Barro Colorado Island. 

Stanford University Press, Stanford, CA. 943 p. 

Howard, R.A. 1979. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 3. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 586 p. 

Instituto Nacional de Biodiversidad. 2002. 

Jerarquía taxonómica: Lista de especímenes de 

Gynerium sagittatum. http://www.inbio.ac.cr/ 

bims/k03/p13/c046/00159/f01382/g008647/s027

308.htm. 2 p. 

Kalliola, R., M. Puhakka, and J. Salo. 1992. 

Interspecific variation, and the distribution and 

ecology of Gynerium sagittatum (Poaceae) in the 

western Amazon. Flora 186(3-4): 153-167. 

Kalliola, R., J. Salo, M. Puhakka, and M. Rajasilta. 

1991. New site formation and colonizing 

vegetation in primary succession on the western 

Amazon floodplains. Journal of Ecology 79(4): 

877-901. 

Pohl, R.W. 1983. Gynerium sagittatum (caña 

brava, cane). In: D.H. Janzen, ed. Costa Rican 

natural history. University of Chicago Press, 

Chicago and London. p. 248-249. 

Russo, E.B. 2002. Village life of the Machiguenga. 

http://montana.com/manu/village.html. 4 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanic Garden Press, St. Louis, 

MO. p. 1,911-2,666. 
372 

Hamelia patens Jacq. firebush 

RUBIACEAE 

Synonyms: Hamelia erecta Jacq. 

 Hamelia pedicellata Wernh. 

 Hamelia latifolia Reichb. ex DC. 

John K. Francis 

 

General Description.—Firebush, which is also 

known as hummingbird bush, scarletbush, 

bálsamo, busunuvo, pata de pájaro, Doña Julia, 

fleur-corail, and many other names, is a mediumsized shrub. This shrub commonly ranges from 1 

to 3 m in height, but sometimes reaches 7 m in 

height (Howard 1989). It grows as a tree in the 

Atlantic tropical lowland of Costa Rica (Paciorek 

and others 1995). Firebush has a tap and lateral 

root system with abundant fine roots. The roots are 

red-brown. Stem bark is gray and smooth and the 

inner bark is light green (Little and others 1974). 

Plants may have single or multiple stems. The 

twigs are orange to purple. Leaves are opposite or 

grouped in threes or fours, and finely hairy to 

glabrous. The leaves have petioles 1 to 3.5 cm long 

and blades that are mostly ovate-elliptic to 

obovate-elliptic with an acute or acuminate tip. The 

lateral and especially the midveins are red or pink. 

In temperate areas, as the temperatures turn cool in 

the fall, the foliage turns to a brilliant red—hence 

the common name. The inflorescence is terminal, a 

modified dichasium with flowers that are tubular, 

12 to 22 mm long, and orange to red in color. The 

fruit is a berry, spherical to elliptical, 7 to 10 mm 

long, turning red and then black at maturity. The 

seeds are orange-brown, 0.6 to 0.9 mm long 

(Howard 1989, Liogier 1997). 

Range.—The native range of firebush extends 

from southern Florida and Bermuda, through the 

Bahamas, the Greater and Lesser Antilles, Trinidad 

and Tobego, and from Mexico through Central 

America and South America to Paraguay and 

Argentina (Little and others 1974). The species is 

also cultivated throughout the moist tropics and 

subtropics but is not reported to have naturalized 

outside its native range. 

Ecology.—Firebush grows in deforested areas, in 

thickets with other brushy species, in forest 

openings, or in the understory of low basal-area 

forest stands. The species is found in moist and wet 

areas that receive from about 1600 to about 3000 

mm of precipitation. Firebush prefers loamy or 

clayey soil. It grows on soils derived from volcanic 

and sedimentary parent materials and is most 

common in areas with limestone rocks. 

Reproduction.—Firebush flowers throughout the 

year. It is a hummingbird-pollinated plant 

(Cunningham 1994). The flowers are also visited 

by butterflies (Desert-Tropicals 2001). There were 

970,000 seeds/kg in a collection of seeds from 

Puerto Rico. Sown on wet peat, germination started 

after 20 days and ended at 40 days with 46 percent 

germinated. Although the species is a good 

producer of fruits and seeds in Puerto Rico, 

seedlings are not common. The fruits are eaten by 

birds (Floridata 2001), which disburse the seeds. 

Layering occurs whenever stems come in contact 

with the ground. The species can be propagated 

from seeds, but most commercial ornamentals are 

produced from cuttings. In one test, cuttings 15 cm 

long with four leaves, which had been taken from 

the shoot tip, treated with 4000 ppm of indolbuteric acid (IBA) and planted in sand, gave the 

best rooting (Bhattacharjee and Balakrishna 1992). 
373

Another experiment using an auto-humid chamber 

obtained 100 percent rooting of softwood cuttings 

treated with hormones and 40 percent without 

hormone treatment (Maiti 1974). 

Growth and Management.—In its native and 

other frost-free habitat, firebush grows as a typical 

shrub, but cultivated in warm temperate areas with 

hard frosts, it will freeze to the ground and sprout 

again in the spring. Firebush is also planted as an 

annual bedding plant in temperate climates. 

Ornamental plants need frequent watering during 

the establishment phase, but afterward, they are 

drought tolerant (Floridata 2001). Pruning of 

ornamental plants is recommended to keep them 

from becoming tall and spindly (Desert-Tropical 

2001). However, pruning inhibits blooming, which 

occurs at the terminals. 

Benefits.—Firebush is loved and planted as an 

ornamental almost worldwide in warm, moist 

areas. The fruit is edible (Little and others 1974). A 

phytochemical screening found firebush tissues 

rich in phytochemicals including alkaloids and 

flavonoids (Raintree Nutrition 2001). Firebush is 

used in herbal medicine to treat athlete’s foot, skin 

lesions and rash, insect bites, nervous shock, 

inflammation, rheumatism, headache, asthma, and 

dysentery (Liogier 1990, Mast Arboretum 2001, 

Raintree Nutrition 2001). Firebush is one of the 

woody species foraged by sheep and goats in 

brushy pastures in Costa Rica. These animals 

consumed 1.3 percent of their body weight of this 

species in a free-choice test (Benavides 2001). 

Firebush contains 17.5 percent crude protein and 

has an in vitro digestibility of 61.6 percent 

(Benavides 2001). 

References 

Benavides, J.E. 2001. Arboles y arbustos 

forrajeros: una alternativa agroforestal para la 

ganadería. Conferencia electrónica de la FAO 

sobre la producción animal en Latinoamerica. 

http://lead.virtualcentre.org/es/ele/conferencia1/

bnvdes23.htm. 22 p. 

Bhattacharjee, S.K. and M. Balakrishna. 1992. 

Studies on propagation of Hamelia patens Jacq. 

and Ixora singaporensis Hort. from stem 

cuttings. Progressive Horticulture 24(3/4): 157-

164. 

Cunningham, S.A. 1994. Measuring the 

relationships between floral duration and fruit 

set for Hamelia patens (Rubiaceae). Biotropica 

26(2): 227-229. 

Desert-Tropicals. 2001. Texas firecracker bush, 

scarlet bush. http://www.deserttropicals.com/Plants/Rubiaceae/Hamelia_patens.

html. 2 p. 

Floridata. 2001. Hamelia patens. 

http://www.floridata.com/ref/h/hame_pat.cfm. 

3 p. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p.

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc. San Juan, PR. 566 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Maiti, R.G. 1974. Regeneration of terminal 

greenwood cuttings of some ornamental shrubs 

under auto-humid chamber. Plant Science 6: 

104-105. 

Mast Arboretum. 2001. Hamelia patens, firebush. 

Steven F. Austin State University. 

http://www.sfasu.edu/ag/arboretum/plants/hamel

iapatens/hamelia%20patens.htm. 3 p. 

Paciorek, C.J., B.R. Moyer, R.A. Levin, and S.L. 

Halpern. 1995. Pollen consumption by the 

hummingbird flower mite, Proctolaelaps 

kirmsei, and possible fitness effects on Hamelia 

patens. Biotropica 27(2): 258-262. 

Raintree Nutrition. 2001. Scarlet bush. Raintree 

Nutrition, Inc. http://www.raintree.com/scarletbush.htm. 3 p. 
374 

Helicteres jamaicensis Jacq. cowbush 

STERCULIACEAE 

Synonym: Helicteres spiralis Northop 

John K. Francis 

General Description.—Cowbush is a shrub or 

sometimes a small tree 7 m in height and up to 20 

cm in basal diameter. It is also known as cowitch, 

blind-eye-tree, salzbush, wild salve, screwtree, 

jeucon, cottonrat, cuernecillo, gato soga, huevo de 

gato, gato, majagüilla de costa, and tapaculo (Little 

and others 1974). The shrub has a dense covering 

of soft hairs throughout. The trunk is gray with 

conspicuous warts. A somewhat swollen taproot 

with abundant laterals and fines form the root 

system. The tan-colored roots are somewhat stiff 

and woody. There is usually a single stem unless 

the plant has been damaged. It develops relatively 

few long, stout, spreading or drooping branches. 

The alternate leaves have short petioles and ovate 

to ovate-lanciolate blades with toothed edges. 

Flowers are few and form in groups of one to three 

at leaf bases. They have five white petals. The fruit 

consists of five gray-green hairy carpals that twist 

upon themselves to form an ellipse about 2 to 4.5 

cm long. The seeds are blackish and about 1 mm in 

diameter (Liogier 1994, Little and others 1974). 

Range.—About 40 tree or shrub species of 

Helicteres are found in tropical America and Asia 

(Griffiths 1994). Cowbush is native to the 

Bahamas, the Greater Antilles, the Virgin Islands, 

and St. Martin (Little and others 1974). Planting or 

naturalization in other areas has not been reported. 

Ecology.—Cowbush grows on soils of all textures 

that are derived from sedimentary, igneous, and 

metamorphic rocks, including ultramafics. In 

Puerto Rico, it grows between sea level and 400 m 

elevation and in areas that receive between about 

700 and 1700 mm of annual precipitation. Welldrained soils are required. The species competes 

well with herbs and shrubs in early and middle 

secondary forest. It grows on roadsides, in 

openings, and in low and open forest canopies, but 

not under closed canopies. Some degree of 

disturbance appears to be necessary to perpetuate 

the species. It is common in many partially forested 

rangelands, but disappears under severe 

overgrazing or frequent fires. 

Reproduction.—Cowbush flowers irregularly 

through the year. The fruits in a Puerto Rican 

collection averaged 1.436 g/fruit and contained an 

average of 152 seeds/fruit. The seeds weighed an 

average of 0.0032 g/seed or 312,000 seeds/kg. The 

carpels of the fruits open at the ends to gradually 

release the seeds as the fruits shake in the wind. To 

collect small quantities, tear the fruits apart and 

pick out the seeds or cut off the ends of the fruits 

and tap them to release most of the seeds. Larger 

quantities can be obtained by pulverizing the fruits 

and winnowing the seed. Seeds from the Puerto 

Rican collection were sown in commercial potting 

mix and germinated (99 percent) between 11 and 

22 days after sowing. Germination is epigeal. 

Natural seedlings are usually common wherever a 

seed source is present. However, few progress past 

the small seedling stage. 

Growth and Management.—Growth is relatively 

rapid after the early seedling stage, but the shrubs 

are not long-lived (5 to 15 years). The use of 

containerized seedlings for plantations is 

recommended. 

Benefits.—Cowbush is occasionally planted as an 

ornamental. It is attractive as a large foliage or 

background plant and makes an interesting 

curiosity plant. It has been planted in revegetation 

projects (Reforesta 2001). The wood of the species 

is yellow, hard and heavy, but not available in sizes 

sufficient to be used for anything other than fuel. 
375

The fibrous bark is strong and was once used for 

making rope (Little and others 1974). Cowbush is 

reported to have several uses in herbal medicine. A 

tea made from the wood and leaves is used to treat 

bilious conditions and as an emetic. A decoction of 

those same tissues is good for fever and cough, and 

the cooked root is able to stop bleeding (Liogier 

1990). The hairs of the species are reported to 

cause serious irritation (Jamaicans 2001), but this 

is not reported by Little and others (1974) and was 

found to be untrue by the author, at least for Puerto 

Rican plants. 

References

Griffiths, M. 1994. Index of garden plants. Timber 

Press, Portland, OR. 1,234 p. 

Jamaicans. 2001. Jamaica: Bites and Stings, etc. 

http://jamaicans.com/tourist/bites.htm. 3 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. 461 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR 566 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 

Reforesta. 2001. Restauración ribereña. 

http://www.reforesta.com/projects/Espanol 

 /riparian_restoration_esp.htm. 2 p. 
376 

Hibiscus pernambucensis Arruda seaside mahoe

MALVACEAE 

Synonyms: Hibiscus tiliaceus var. pernambucensis (Bertol.) I.M. Johnston 

 Hibiscus tiliaceus ssp. pernambucensis (Arruda) Castell 

 Hibiscus arboreus Ham. 

 Paritium pernambucense (Bertol.) Don 

John K. Francis 

General Description.—Seaside mahoe is also 

known as sea hibiscus and rope mangrove in 

English, emajagua and majagua in Spanish, and 

mahoe doux, mahaut, and bois flot in French 

(Howard 1989, Liogier 1994). It is a large shrub or 

occasionally a small tree. The species normally 

grows in tangled thickets formed by stems that sag 

until they touch the ground and form roots. 

Simultaneously, sprouts arise from the horizontal 

stems and grow upward until they in turn become 

prostrate. Occasionally, especially on dry ground, 

open-grown individuals grow in a tree-like habit. 

Adult plants have an aggressive lateral root system 

with abundant fine roots. The bark of the limbs 

and trunks is gray and smooth, but can become 

furrowed and scaly in old stems. The leaves are 

nearly round (10 to 15 cm in diameter) except for a 

deeply cordate base and a pointed tip. The petioles 

are long (6 to 9 cm). Seaside mahoe produces large 

yellow flowers that change to orange or red as they 

age. Soon after, 2-cm long, hairy, cone-shaped 

capsules develop, and upon drying, split open 

along five lines to release their seeds. 

Range.—Seaside mahoe is native to coastal areas 

of Bermuda, southern Florida, the Bahamas, the 

Greater and Lesser Antilles, and from Mexico 

through Brazil on the Atlantic Coast and in Pacific 

coastal areas from Mexico to Peru (Little and 

Wadsworth 1964). Seaside mahoe is similar to and 

once was classified as a variety of Hibiscus 

tiliaceus L. All the New World natives are now 

thought to be H. pernambucensis (Howard 1989). 

Ecology.—Seaside mahoe usually grows just 

inland of mangroves around brackish lakes and 

along rivers in the lower reaches of their 

floodplains. Thickets have formed from planted 

individuals in abandoned subsistence farms in 

rainy (usually above 2000 mm of precipitation) 

upland sites with poorly drained soils. It also 

grows as a beachfront tree in association with 

Terminalia catappa L. (Malavasi and Malavasi 

1999). Seaside mahoe grows on hummocks and 

sand dunes in southern Florida (West and Arnold 

1952). The sandy habitat probably has a water 

table within reach of the roots. Seaside mahoe 

suffers when there are droughts and appears to 

require poorly drained sites to form the 

characteristic thickets. Although the species grows 

near brackish lakes, it is definitely less tolerant of 

saline conditions than the mangroves. All textures 

of soil with a wide range of mineral fertility are 

represented in seaside mahoe habitat. The northern 

extent of the range is limited by the frost line. 

 Seaside mahoe grows in nearly pure 

stands. Even artificially established stands in 

upland areas maintain themselves for many 

decades. This can probably be attributed to its 

rapid growth, profuse sprouting, and the dense 

crowns that are formed. Seedlings of other species 

rarely survive for long. A number of insects have 

been collected from seaside mahoe (Martorell 

1975). Although no insect seriously threatens the 

species, aphids (Aphidoidea), white flies 
377

(Aleyrodidae), and mealy bugs (Pseudococcideae) 

sometimes cause chlorosis and slower growth. 

Reproduction.—The principal means of 

reproduction is by the layering and sprouting of 

prostrate stems. The old prostrate stems eventually 

die and rot, leaving the sprouts as new, isolated 

individuals. Seeds are produced in relative 

abundance and occasionally result in seedlings that 

can eventually establish new clones. Streams and 

floodwaters are the only known effective vector of 

seed dispersal. 

 In wetter habitat, seaside mahoe flowers 

continuously throughout the year (Little and 

Wadsworth 1964). The species flowers seasonally 

in habitat with drought stress during part of the 

year. The flowers are pollinated by insects, and 

yield mature seeds about 1 month later. Fruits can 

potentially produce at least 40 seeds per fruit. In 

one collection from upland habitat in Puerto Rico, 

fruits averaged only 7.7 filled seeds per fruit. In 

two other Puerto Rican collections from upland 

habitat, the seeds averaged 0.018 + 0.006 and 

0.024 + 0.003 g/seed. These seed collections 

yielded 41 and 53 percent germination, which 

occurred between 15 and 220 days after sowing. 

Germination is epigeal. Seedlings are easily raised 

in the nursery from seed using traditional 

techniques. Plants can be established from cuttings 

and air layers (Workman 1980). In one small 

planting in Puerto Rico, 40 of 140 untreated 

cuttings (45 cm long and 2.5 cm minimum 

diameter) placed directly in the planting site rooted 

and grew aggressively. Plants can also be started 

from very large cuttings as living fence posts 

(Little and Wadsworth 1964). 

Growth and Management.—Seaside mahoe 

grows rapidly from sprouts and can extend its 

leader 3 m or more per year. Seedlings grow 

slowly at first. A group of 6-month-old seedlings 

averaged only 17 cm in height. Probably, 1 year is 

required before nursery seedlings are ready for 

outplanting. Seaside mahoe may occasionally 

reach 12 to 15 m in height (Workman 1980), but 

more typically reaches only 6 m in height and 15 

cm in stem diameter. Because the species is 

mainly valued in natural stands for environmental 

protection and because it aggressively maintains 

nearly pure stands, little or no management is 

required once it is established. One unwanted 

upland stand was converted to timber species by 

slashing the seaside mahoe, piling the slash 

between the planting spots, and planting seedlings 

of Eucalyptus grandis Hill ex Maiden, Terminalia 

ivorensis L., and Hibiscus elatus Sw. in the small 

cleared planting spots with no other treatment. The 

fast-growing species that were planted overtopped 

the sprouting seaside mahoe and eventually 

suppressed them. 

Benefits.—The fibrous inner bark was formerly 

used to make ropes, cords, fish nets, floats, cloth, 

and mats. It is still used for emergency ties and 

lashings. The wood is used for fuel in 

underdeveloped areas (National Academy of 

Sciences 1983). The flowers make a colorful and 

tasty addition to salads and can be cooked as a 

vegetable (Workman 1980). The young leaves, 

bark, and roots were reportedly used for food 

during famines in former times (Little and 

Wadsworth 1964). Bees make honey from the 

nectar. The plant is used as an ornamental and to 

stabilize stream banks (Workman 1980). 

References 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

Malavasi, U.C. and M.M. Malavasi. 1999. Tree 

shade preferred by recreational users of a 

Brazilian beach sidewalk. Journal of 

Arboriculture 25(2): 85-87. 

Martorell, L.F. 1975. Annotated food plant catalog 

of the insects of Puerto Rico. Agricultural 

Experiment Station, University of Puerto Rico, 

Río Piedras, PR. 303 p. 

National Academy of Sciences. 1983. Firewood 

crops: shrub and tree species for energy 

production. BOSTID Report 40. National 

Academy of Sciences. Washington, DC. 92 p. 

West, E. and L.E. Arnold. 1952. The native trees 

of Florida. University of Florida Press, 

Gainsville, FL. 212 p. 
378 

Workman, R.W. 1980. Growing native. The 

Sanibel-Captiva Conservation Foundation, Inc. 

Sanibal, FL. 137 p. 
379

Holodiscus discolor (Pursh) Maxim. creambush oceanspray 

ROSACEAE 

Synonyms: Spiraea discolor Pursh 

Spiraea ariaefolia Smith in Rees 

Schizonotus discolor Raf. 

Sericotheca discolor Rydb. 

Nancy L. Shaw 

Illustration credits: Hitchcock and others 1961 and E.G. 

Hurd

General Description.—Creambush oceanspray, 

also known as arrowwood, hardhack, rock-spirea, 

and mountain spray, is a usually erect, deciduous 

shrub with multiple arching branches. Plants range 

from 1 to 3 (6) m in height. Large tree-like forms 

grow in coastal areas. Young twigs are finely 

hairy; bark on older stems is gray to deep grayish 

red. Leaves are alternate, 2 to 10 cm long, and 

ovate to ovate-lanceolate with lobed to doubly 

toothed edges and prominent veins. Upper surfaces 

are hirsute; lower surfaces are lanate and 

sometimes glandular. The common name derives 

from the showy, diffuse, drooping panicles that are 

10 to 17 (30) cm long with numerous, tiny, white 

to pink flowers. Dried flowers and panicle 

branches turn brown and persist into winter 

(Hitchcock and others 1961). 

Range.—Creambush oceanspray grows at 

elevations from sea level to 2,150 m. It is most 

abundant in coastal forests from British Columbia to 

southwestern California. It occurs eastward to 

Montana in drier conifer types of the interior Pacific 

Northwest. It is also common in riparian areas and 

on rocky talus slopes with shallow sandy to clay 

loam soils. Remnant stands are found on higher 

peaks of Great Basin mountain ranges (Hitchcock 

and others 1961, Stark 1966, USDA Forest Service 

1937). 

Ecology.—Creambush oceanspray is a prolific root 

sprouter, capable of recovering from fire, grazing, 

or mechanical damage. Regeneration from seed 

following burns is often limited, and seedlings 

develop slowly (Wright and others 1979). However, 

regeneration from seed may be important following 

extremely hot fires (Morgen and Neuenschwander 

1988). The species is a host for vesicular-arbuscular 

mycorrhizae (Berch and others 1988) and 

Orobanche pinorum Geyer ex Hook. (Harrington 

and others 1999). It is shade tolerant and disease 

resistant. 

Reproduction.—Flowers appear from late spring 

to mid summer and are insect-pollinated. Fruits 

ripen in late summer (Hitchcock and others 1961); 

seed is wind dispersed. Fruits are hand harvested 

and conditioned by air drying and screening larger 

material with a fanning mill. Number of cleaned 

seed/kg exceeds 11 million (King 1947, Link 1993), 

but few seeds are generally sound. Seed supplies are 

rare and costly. Moist prechilling at 1 to 2 o

C for 

120 days relieves dormancy (Marchant and 

Sherlock 1984). Natural germination may be 

enhanced by a heat pretreatment or microsite 

conditions provided by exposed mineral soils 

(Morgan and Neuenschwander 1988). Seed may be 

broadcast over a rough seedbed and covered by 

natural soil sloughing. It should be planted 

separately or included in seed mixes of other native 

species, particularly if broadcast seeded. It should 

not be seeded with competitive introduced species 

(Shaw and others 2001). Bareroot and container 

stock are easily propagated (Everett 1957). Achenes 

should be fall sown or artificially prechilled and 

spring sown in the bareroot nursery. Creambush 

oceanspray wildings and layers have also been used 

successfully (Kruckeberg 1982). Plants can be 
380 

grown from cuttings, but rooting varies widely 

among clones, cutting type, and propagation 

technique (Antieau 1987, Link 1993). 

Growth and Management.—Seedlings develop 

slowly, producing spreading, fibrous root systems 

(Wright and others 1979). Growth rate is moderate 

and protection from introduced grass and forb 

competition is required for one or two growing 

seasons. Excessive browsing is rarely a threat to 

developing seedlings, but they may be girdled by 

rodents or trampled by big game or livestock. Brush 

fields may be rejuvenated by prescribed burning at 

10 to 15 year intervals (Orme and Leege 1980). 

Benefits.—Creambush oceanspray provides habitat 

for a wide variety of organisms. Its palatability and 

forage value vary geographically but are generally 

low for big game and domestic livestock. Use may 

be greater in winter when other forage is limited. 

Palatability may be improved following fire. 

Creambush oceanspray offers considerable potential 

for revegetating disturbed areas. Populations 

capable of growing on dry, rocky, well-drained sites 

or exhibiting low palatability may be particularly 

valuable. It has been recommended for use in 

highway plantings, riparian areas, windbreaks, 

erosion control, wildlife habitat improvement, 

conservation plantings, and low maintenance 

landscaping (Antieau 1987). Native Americans ate 

the seeds and made digging sticks and arrow shafts 

from the hard, straight branches. 

References 

Antieau, C.J. 1987. Field notes: Holodiscus

discolor. American Nurseryman 166: 110. 

Berch, S.M., S. Gamiet, and E. Deom. 1988. 

Mycorrhizal status of some plants of 

southwestern British Columbia. Canadian 

Journal of Botany 66: 1,924-1,928. 

Everett, P.C. 1957. A summary of the culture of 

California plants at the Rancho Santa Ana 

Botanic Garden. The Rancho Santa Ana Botanic 

Garden, Claremont, CA. 223 p. 

Harrington, C.A., J.M. McGrath, and J.M. Kraft. 

1999. Propagating native species: experience at 

the Wind River Nursery. Western Journal of 

Applied Forestry 14: 61-64. 

Hitchcock, C.L., A. Cronquist, M. Ownbey, and 

J.W. Thompson. 1961. Vascular plants of the 

Pacific Northwest. Part 3: Saxifragaceae to 

Ericaceae. University of Washington Press, 

Seattle, WA. 614 p. 

King, J.E. 1947. The effect of various treatments 

to induce germination of seeds of some plants 

valuable for soil conservation and wildlife. 

Master’s thesis. University of Idaho, Moscow, 

ID. 97 p. 

 

Kruckeberg, A.R. 1982. Gardening with native 

plants of the Pacific Northwest. University of 

Washington Press, Seattle, WA. 252 p. 

Link, E. 1993. Native plant propagation techniques 

for National Parks interim guide. Rose Lake 

Plant Materials Center, East Lansing, MI. 240 p. 

Marchant, C. and J. Sherlock. 1984. A guide to 

selection and propagation of some native woody 

species for land rehabilitation in British 

Columbia. Forest Research Report RR84007-

HQ. Ministry of Forests, Victoria, BC. 117 p. 

Morgan, P. and L.F. Neuenschwander. 1988. Seedbank contributions to regeneration of shrub 

species after clear-cutting and burning. Canadian 

Journal of Botany 66: 169-172. 

Orme, M.L. and T.A. Leege. 1980. Phenology of 

shrubs on a north Idaho elk range. Northwest 

Science 54: 187-198. 

Shaw, N.L., E.G. Hurd, and P.F. Stickney. 

Holodiscus (K. Koch) Maxim. 2001. Oceanspray. In: F.T. Bonner and R.G. Nisley, eds. 

Woody plant seed manual. U.S. Department of 

Agriculture, Forest Service, Agriculture 

Handbook. Washington, D.C. 

http://wpsm.net/index.html. 7 p. 

Stark. N. 1966. Review of highway planting 

information appropriate to Nevada. College of 

Agriculture Bulletin B-7. University of Nevada, 

Desert Research Institute. Reno, NV. 209 p. 

USDA Forest Service. 1937. Range plant 

handbook. U.S. Government Printing Office, 

Washington, D.C. 512 p. 
381

Wright, H.A., L.F. Neuenschwander, and C.M. 

Britton. 1979. The role and use of fire in 

sagebrush-grass and pinyon-juniper plant 

communities: A state of the art review. General 

Technical Report INT-58. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Forest and Range Experiment Station, Ogden, 

UT. 48 p. 
382 

Holodiscus dumosus (Nutt. ex Hook.) Heller rockspirea 

ROSACEAE 

Synonyms: Spiraea dumosa (Nutt.) T. & G.

 Schizonotus dumosus Koehne. 

 Holodiscus microphyllus Rydb. 

 Sericotheca dumosa Rydb. 

Nancy L. Shaw 

 Illustration credits: Hitchcock and others 1961 and 

E.G. Hurd 

General Description.—Rockspirea, also 

known as gland oceanspray, bush oceanspray, 

mountain-spray, and creambush, is a compact, 

rounded, intricately branched deciduous shrub 

0.1 to 4 m tall with a crown diameter of 1.5 to 3 

m (Hitchcock and others 1961, Welsh and 

others 1987). Leaf blades are oval and 

shallowly or coarsely lobed or toothed, but 

generally without secondary teeth. Petioles are 

decurrent along the stems. Panicles are showy 

and terminal, overtopping the shrub. Flowers 

are numerous, small, creamy-white, perfect, 

perigynous, and insect pollinated. Fruits are 

tiny hirsute achenes. Roots are densely fibrous 

and spreading. The species has n = 18 

chromosomes (McArthur and Sanderson 

1986). 

Range.—Rockspirea grows east of the Cascade and 

Sierra Nevada Mountains from north central 

Oregon, east to Wyoming, and south to Chihuahua, 

Mexico at elevations ranging from 760 to 3,660 m 

(Harrington 1964, Hopkins and Kovalchek 1983). 

Ecology.—Rockspirea occurs in a wide array of 

plant communities including Artemisia spp., 

Pinus spp.--Juniperus spp. L., Cercocarpus 

ledifolius Nutt., chaparral, Populus tremuloides

Michx.--Pinus contorta Dougl. ex Loud., Picea

spp.--Abies spp., Pseudotsuga menziesii (Mirbel) 

Franco, Abies concolor (Gord. & Glend.) Lindl. ex 

Hildebr., and Pinus ponderosa Dougl. (Sutton and 

Johnson 1974, USDA 1937). It grows as a pioneer 

species in cracks and fissures on rock surfaces, as 

a seral species in brush fields of forested areas, and 

as a climax species in self-replacing, monospecific 

stands. Within its range it commonly occurs on 

well-drained, dry to moderately dry sandy or 

gravelly soils that may be somewhat alkaline, but 

it is also found on finer-textured soils. Rockspirea 

often grows on dry, rocky ridges, talus slopes, and 

basalt outcrops (Hitchcock and others 1961, Welsh 

and others 1987). It is a prolific root sprouter, 

capable of recovering from fire, grazing, or 

mechanical damage. 

Reproduction.—Panicles and floral buds develop 

in early spring, but flowering is delayed until 

summer or early fall depending upon elevation. 

Fruits are wind and gravity dispersed through late 

fall (Hitchcock and others 1961). King (1947) 

determined that although there are about 

12,000,000 seed/kg, only about 7 percent were 

sound. These required moist prechilling at 5 o

C for 

18 weeks to release dormancy. Heat generated by 

wildfires may also stimulate germination. Seed can 

be broadcast seeded on a rough seedbed. Planting 

stock can be grown by fall seeding in the bareroot 

nursery or planting moist prechilled seed in the 

greenhouse. 

 

Growth and Management.—Excessive browsing 

rarely occurs as the species is present primarily in 
383

rocky, inaccessible areas and on summer ranges 

where other species receive preferential use. Based 

on results of clipping studies in Oregon, Garrison 

(1953) recommended a maximum of 50 to 60 

percent use for sustained production. On some 

western juniper/big sagebrush habitat types of 

eastern Oregon, the species tends to increase when 

other species receive excessive use (Ferguson 

1983, Hopkins and Kovalchik 1983). 

 

Benefits.—Rockspirea is a potentially valuable 

species for revegetation as it is drought tolerant 

and adapted to sites with dry, rocky, unstable 

surface conditions in a variety of forested and 

nonforested communities (Stark 1966). Its use has 

been limited by a lack of high quality seed 

supplies. Palatability and forage value of 

rockspirea are low (USDA 1937), but it does 

provide food and cover for small animals. Kufeld 

and others (1973) reported its use by mule deer 

(Odocoilus hemionus) was moderate in fall and 

light during the remainder of the year. Rockspirea 

is used in summer by bighorn sheep (Ovis 

canadensis) and rabbits (Sutton and Johnson 1974, 

Todd 1975). It is an attractive, low water use 

ornamental. Native Americans made digging sticks 

and arrow shafts from the hard, straight branches 

(Anderson and Holmgren 1969, Daubenmire 1970, 

Hopkins and Kovalchik 1983). Native Americans of 

the Great Basin ate the seeds, and pioneers made 

nails from its wood. 

 

References 

Anderson, B.A. and A.H. Holmgren. 1969. 

Mountain plants of northeastern Utah. Circular 

319. Utah State University, Logan, UT. 148 p. 

Daubenmire, R. 1970. Steppe vegetation of 

Washington. Technical Bulletin 62. Washington 

State University, Pullman, WA. 131 p. 

Ferguson, R.B. 1983. Use of rosaceous shrubs for 

wildland plantings in the intermountain west. In: 

S.B. Monsen, and N. Shaw, comps. Managing 

Intermountain rangelands--Improvement of 

range and wildlife habitats: Proceedings. 

General Technical Report INT-157. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. p. 136-139. 

Garrison, G.A. 1953. Effects of clipping on some 

range shrubs. Journal of Range Management 6: 

309-317. 

Harrington, H.D. 1954. Manual of the plants of 

Colorado. Sage Books, Denver, CO. 666 p. 

Hitchcock, C.L., A. Cronquist, M. Ownbey, and 

J.W. Thompson. 1961. Vascular plants of the 

Pacific Northwest. Part 3: Saxifragaceae to 

Ericaceae. University of Washington Press, 

Seattle. 614 p. 

Hopkins, W.E. and B.L. Kovalchik. 1983. Plant 

associations of the Crooked River National 

Grassland. R6 Ecology 133-1983. U.S. 

Department of Agriculture, Forest Service, 

Pacific Northwest Region, Portland, OR. 98 p. 

King, J.E. 1947. The effect of various treatments 

to induce germination of seeds of some plants 

valuable for soil conservation and wildlife. . 

Master’s thesis. University of Idaho, Moscow, 

ID. 97 p. 

Kufeld, R.C. 1973. Foods eaten by the Rocky 

Mountain elk. Journal of Range Management 

26: 106-113. 

McArthur, E.D. and S.C. Sanderson. 1985. A 

cytotaxonomic contribution to the western North 

American rosaceous flora. Madroño 32: 24-28. 

Stark. N. 1966. Review of highway planting 

information appropriate to Nevada. College of 

Agriculture Bulletin B-7. University of Nevada, 

Desert Research Institute, Reno. 209 p. 

Sutton, R. and C.W. Johnson. 1974. Landscape 

plants from Utah’s mountains. EC-368. Utah 

State University, Logan, UT. 137 p. 

Todd, J.W. 1975. Foods of Rocky Mountain 

bighorn sheep in southern Colorado. Journal of 

Wildlife Management 39: 108-111. 

USDA Forest Service. 1937. Range plant 

handbook. U.S. Government Printing Office, 

Washington, D.C. 512 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, L. 

Higgins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoir 9. Brigham Young 

University, Provo, UT. 894 p. 
384 

Hyperbaena laurifolia (Poir.) Urban limestone snakevine 

MENISPERMACEAE 

Synonyms: Cissampelos laurifolia Poir. in Lam. 

 Anelasma laurifolia (Poir.) Meirs 

 Cocculus laurifolia Eggers 

John K. Francis 

General Description.—Limestone snakevine is an 

uncommon scrambling shrub or vine-like small 

tree. It is also known as hyperbaena, following the 

convention among natural resources workers to use 

the Latin genus or species name as a common 

name. The vine develops a tap and lateral root 

system. Plants of the species have single or 

multiple smooth, gray stems with a moderate 

number of branches and stiff, slender, hairless 

twigs. Limestone snakevine has alternate, oblong 

to lanciolate, hairless, leathery leaves with a short 

petiole and blades from 7 to 16 cm long and 2.5 to 

6 cm broad. The leaves are shiny, dark green above 

and dull, lighter green below, have smooth edges, 

and may be rounded or pointed at either end. Male 

and female flowers grow on different plants 

(dioecious). The individual flowers are tiny, 

greenish-yellow, and grow in clusters at the leaf 

bases. The fruits (drupes) that develop are rounded 

and somewhat flattened, bright red when ripe, 2 to 

3 cm broad with a plum-like skin and flesh about 5 

mm thick and contain a single seed. The large 

yellow stone within is grooved and shaped like a 

crescent curved back upon itself (Liogier 1985, 

Little and others 1974, Mathias and Theobald 

1981). 

Range.—Limestone snakevine is native to Puerto 

Rico, St. Thomas in the U.S. Virgin Islands 

(Liogier 1985), and Haiti (Mathias and Theobold 

1981). It had been reported in Montserrat (Little 

and others 1974), but apparently this was in error 

(Howard 1988). 

Ecology.—Little and others (1974) and Mathias 

and Theobald (1981) reported that this species is 

confined to limestone areas. However, the author 

knows of two stands near Salinas, Puerto Rico that 

are growing on soil residual from igneous rocks. 

Limestone snakevine grows from near sea level to 

1,200 m in elevation (presumably in Haiti) 

(Mathias and Theobald 1981). In Puerto Rico, it 

grows to elevations of 600 m (Little and others 

1974), and in areas that receive from about 800 to 

about 2000 mm of mean annual precipitation. 

Limestone snakevine grows in a variety of soils, 

most of which are very rocky in the currently 

available habitat. The species is moderately shade 

tolerant and grows and reproduces mainly under 

the crowns of trees in remnant and advanced 

secondary forests. 

Reproduction.—Limestone snakevine flowers in 

the spring and bears fruits during the spring and 

summer (Little and others 1974). The fruits are 

easily detached and presumably dispersed by fruit 

bats. Forty fruits in a Puerto Rican collection 

weighed an average of 9.31 + 0.27 g each. Seeds 

from that collection averaged 4.18 + 0.12 g/seed or 

240 seeds/kg. Ninety-eight percent of the seeds 

sown in potting mix germinated between 45 and 72 

days after sowing. Two of the stones produced two 

seedlings each. Germination is hypogeal, a stiff, 

well-branched root developing considerably before 

the top emerges. The tops reach 10 to 15 cm in 

height from the reserves in the seeds. The seedlings 

are ready to transplant within a month of 

emergence. They are hardy and survive well when 

transplanted from nursery beds to pots. Seedlings 

are not common in the wild. 

Growth and Management.—Limestone 

snakevine may reach 3 m in height, 8 m in lateral 

extension, and 10 cm of basal stem diameter. 

Established plants appear to have a moderate 
385

growth rate, and plants known to the author appear 

to be several decades old. However, no data on 

growth rate and longevity or experience on 

management have been published. 

Benefits.—The fruits of limestone snakevine are 

slightly acid, bitter, and unpleasant to the taste and 

smell. The fruit pulp immediately stains skin and 

cloth a lasting reddish-brown and might be useful 

as a natural dye in crafts. The wood is heavy and 

hard and would certainly be good for smalldiameter fuel. The species contributes to the 

biodiversity of local forests and in a minor way, 

furnishes wildlife food and cover, and protects 

against erosion. 

References 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 

Mathias, M.E. and W.L. Theobald. 1981. A 

revision of the genus Hyperbaena

(Menispermaceae). Brittonia 33(1): 81-104. 
386 

Indigofera suffruticosa P. Mill. wild indigo 

FABACEAE 

 

Synonyms: Indigofera anil L. 

 Indigofera truxillensis HBK. 

Indigofera divericata Jacq. 

Anila tinctoria vera Kuntze 

John K. Francis 

General Description.—Wild indigo, also known 

as indigo, Guatamala indigo, añil, añil de pasto, 

and ti cafe, is a short-lived shrub that reaches 1 to 2 

m in height and 1 to 2 cm in stem diameter. The 

shrub may have multiple stems, especially if it has 

been disturbed by grazing or fire. The stems are 

gray-brown, pubescent, and more or less straight. 

The light green leaves are pinnately compound 

with 9 to 17 narrowly oblong, pubescent leaflets 

that are 1.5 to 2.5 cm long and about 9 mm wide. 

There are 6- to 8-mm lanciolate stipules at the base 

of the leaves. Crimson to rust-red flowers grow in 

short, many-flowered racemes. The curved 

legumes are short (1.1 to 2.5 cm) and contain three 

to seven seeds that are 1.5 mm wide and 1 mm 

thick (Howard 1988, Liogier 1988). 

Range.—Wild indigo is native to Southern United 

States through tropical and subtropical South 

America as well as the Caribbean Islands (Howard 

1988). The species has naturalized in Hawaii (Neal 

1965) and is present in American Samoa, Guam, 

and a large number of the Pacific Island groups 

(Pacific Island Ecosystems at Risk 2001). Wild 

indigo has been introduced into tropical regions of 

the Old World (Liogier 1988). 

Ecology.—In Hawaii, wild indigo has naturalized 

in dry, highly disturbed areas from near sea level to 

1,160 m in elevation (Pacific Island Ecosystems at 

Risk 2001). In Puerto Rico, the species grows from 

near sea level to about 700 m in elevation in areas 

that receive from about 750 to 1800 mm of 

precipitation annually. Soils derived from volcanic 

and sedimentary rocks in a wide range of soil 

textures are colonized. Good drainage is required, 

but not high fertility. Wild indigo is not shade 

tolerant and will not grow under a closed forest 

stand. 

Reproduction.—In Texas, wild indigo flowers 

from July through November (Everitt and Drawe 

1993). In Mexico, fruiting occurs between October 

and February (Moreno-Casasola and others 1994). 

The pods remain open on the branches, exposing 

but not expelling the seeds. Mechanisms of 

transport have not been reported. The seeds in a 

collection of fruits in Puerto Rico had been 

partially consumed by an unknown insect and 

averaged only 0.9 undamaged seeds per pod. These 

seeds averaged 0.0039 g/seed or about 257,000 

seeds/kg. Of these, 17 percent germinated in 7 to 

24 days after sowing. Seed from a Mexican 

collection gave more than 90 percent germination 

(in the field) when a mechanical scarification was 

used and nearly 100 percent when wet-dry cycles 

were combined with scarification (MorenoCasasola and others 1994). 

Growth and Management.—Neal (1965) reports 

that in Hawaii, 20,400 kg of wild indigo can be 
387

produced per hectare from which 227 kg of indigo 

paste can be separated. Wild indigo tested for 

revegetation of soil-lignite overburden varied in 

biomass yield from 162 to 2,432 kg/ha and 

maintained from 8 to 30 percent cover over a 3-

year period (Skousen and Call 1987). Surface 

sowing at the start of the rainy season on freshly 

tilled ground is recommended. Plants usually live 2 

or 3 years in Puerto Rico. 

Benefits.—Although indigo obtained from other 

species of Indigofera was used in the Old World, 

the use of wild indigo by pre-Columbian natives of 

Mexico to dye cloth and paint in various shades of 

blue was passed down to the Spanish colonists 

(Haude 1997). Indigofera species became 

important commercial crops in various tropical and 

subtropical areas. The blue dye was produced by 

fermentation of the leaves, usually with caustic 

soda or sodium hydrosulfite, and the exudates 

processed into dry cake. The blue color developed 

as the cake was exposed to the air (Simon and 

others 1984). Indigo was used as a bluing to 

counter the yellowing in clothes from washing with 

soap (Vélez and van Overbeek 1950). In the last 

few decades, natural indigo has been almost wholly 

replaced by synthetic dyes. Poultices and extracts 

of wild indigo leaves, alone or in combination with 

other ingredients, are used in herbal medicine to 

treat fever, headaches, hemorrhages, convulsions, 

acute cough, skin parasites, and boils (HealthLink 

2001, Liogier 1990). Domestic and wild ruminant 

animals browse wild indigo plants. It and another 

Indigofera species are reported to be the sole hosts 

of the false dusky wing butterfly [Gesta gesta

(Herrich-Schäffer)] (Opler and Malikul 1992). 

Wild indigo is useful as a natural cover plant in 

disturbed areas. The species may become weedy, 

but is seldom aggressive or common enough to 

cause difficulties in croplands. 

References 

Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs 

and cacti of South Texas. Texas Tech University 

Press. 213 p. 

Haude, M.E. 1997. Identification and classification 

of colorants used during Mexico’s early colonial 

period. Book and Paper Group Annual Vol. 16. 

The American Institute of Conservation. 

http://aic.stanford.edu/conspec/bpg/annual/v16//

bp16-05.html. 26 p. 

HealthLink. 2001. Monograph: indigo naturalis. 

http://www.healthlink.com.au/nat_lib/htmdata/htm-herb/bhp1016.htm. 3 p. 

Howard, R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Moreira-Casasola, P., J.P. Grime, and M.L. 

Martinez. 1994. A comparative study of the 

effects of fluctuations in temperature and 

moisture supply on hard coat dormancy in seeds 

of coastal tropical legumes in Mexico. Journal of 

Tropical Ecology 10(1): 67-86. 

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Museum 

Press, Honolulu, HI. 924 p. 

Opler, P.A. and V. Malikul. 1992. A field guide to 

eastern butterflies. Peterson field guide 4. 

Houghton-Mifflin Co., Boston, MA. 396 p. 

Pacific Island Ecosystems at Risk. 2001. Invasive 

plant species: Indigofera suffruticosa Miller, 

Fabaceae. http://www.hear.org/pier/insuf.htm. 

 2 p. 

Simon, J.E., A.F. Chadwick, and L.E. Craker. 

1984. Herbs: An indexed bibliography. 1971-

1980. Scientific literature on selected herbs, and 

aromatic and medical plants of the Temperate 

Zone. Archon Books, Hamden, CT. 770 p. 

Skousen, J.G. and C.A. Call. 1987. Grass and forb 

species for revegetation of mixed soil-lignite 

overburden in East Central Texas. Journal of 

Soil and Water Conservation 42(6): 438-441. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Univeresitaria, Río Piedras, PR. 497 p. 
388 

Isocoma tenuisecta Greene burro weed 

ASTERACEAE 

Synonyms: Haplopappus tenuisectus (Greene) Blake 

 Aplopappus tenuisectus (Greene) Blake 

 

Juanita A. R. Ladyman 

Illustration credit: Robert DeWitt Ivey 

General Description.—Burro weed is a compact, 

rounded shrub, sometimes referred to as a subshrub, growing to approximately 1 m high and 

approximately 1 m wide. Its grayish-green, 

glandular leaves are pinnately cleft (deeply lobed) 

into four to eight linear acute lobes. The yellow 

flowers are discoid, having no ray flowers 

(“petals”), and are arranged in dense terminal 

clusters. The flowers dry and turn a light brown 

color while remaining on the stems (PCC 2001). 

The bracts around the flowers are also glandular 

(Martin and Hutchins 1980). Other members of the 

same genus are called golden weeds or yellow bush 

(Dayton 1931). Historically, Haplopappus was a 

large and variable genus that was a depository for 

loosely related “yellow daisies” that shared a 

limited number of characteristics (Taylor 1998). 

However, with the aid of molecular (DNA) data 

and more powerful computer analyses, taxonomists 

have placed members of Haplopappus into 

different genera, such as Isocoma, Ericameria, 

Stenotus and Machaeranthera (Nesom 1991, Lane 

and Hartman 1996). 

Range.—Burro weed occurs in southwestern 

Texas, southern New Mexico, southern Arizona 

and northern Mexico (Kearney and others 1960, 

Martin and Hutchins 1980, Ivey 1995). 

Ecology.—Burro weed grows on dry slopes and 

mesas between 600 and 2,000 m in elevation. It is 

a member of various vegetation communities 

including desert scrub, desert grassland, and pineoak woodland (Bowers and McLaughlin 1987). 

Although burro weed can be moderately long 

lived, up to 20 years, long-term monitoring studies 

indicate as few as 2 percent survive more than 7 

years (Goldberg and Turner 1986). Burro weed has 

a deep (3 to 6 m long), relatively unbranched, root 

system but is susceptible to drought and often dies 

during drought periods (Schmutz and others 1992). 

However, following droughts burro weed shrubs 

often increase in number (Schmutz and others 

1992). Similar to other species within the Isocoma

genus, fire easily kills burro weed, but burned sites 

will eventually become reoccupied (White 1969, 

Cable 1967). For example, fire reduced a pre-burn 

shrub density of 3,762 plants per acre to 327 plants 

per acre that was reduced even further to 87 plants 

per acre in the following year. Those plants that 

survived were in unburned or light burned patches 

within the burn area (Cable 1967). However, the 

numbers of shrubs gradually increased in 

subsequent years to number 7,175 shrubs per acre 

13 years after the burn. The spatial association 

between different three-awn grass species and 

burro weed is an interesting example of the 

tradeoffs between protection and liability of a 

potentially advantageous microclimate and refuge. 

Santa Rita three-awn (Aristada glabrata (Vasey) 

Hitchc.) grows between burro weed individuals, 

whereas some other three-awn species, such as 

Aristida hamulosa Henr. and Aristida ternipes

Cav., grow within the burro weed crowns. The 

latter species are subjected to greater heat in the 

smoldering, resinous crowns than in the open areas 

and, in combination with the shallow subsurface 

buds that characterize all three-awns, suffer a 
389

higher mortality rate than the Santa Rita three-awn 

(Blydenstein 1966, Cable 1967, McPherson 1995). 

Reproduction.—Burro weed flowers from August 

into October (Martin and Hutchins 1980). The 

seeds (achenes) are partially covered by silky hairs 

(Martin and Hutchins 1980). The long hairs that 

comprise the pappus at one end of the seed aid 

dispersal (Parker 1990). 

Growth and Management.—Burro weed is a 

resilient species that can flourish on highway 

shoulders, parking lots, dirt roads, vacant lots and 

other disturbed areas. It tends to be sporadically 

scattered on range covered by perennial bunch 

grasses. However, on disturbed ranges, with even 

a good stand of annual grasses, it can become one 

of the principal plant species (USDA FS 1988). 

Burro weed foliage is poisonous to livestock 

(AVDL 1998, Bowers 1993). Horses are most 

sensitive while sheep are least sensitive. If 

livestock eat large amounts of the vegetation they 

suffer from “trembles,” “alkali sickness,” or “milksickness” and humans can become ill after 

drinking the milk from affected cows (Dayton 

1931, Epple 1995). The toxic substance 

transmitted by the milk is called tremetol (AVDL 

1998, Kingsbury 1964). Many other species of 

Isocoma contain the same toxic constituent 

(AVDL 1998, Kingsbury 1964). As a consequence 

of its toxic properties, livestock avoid it unless 

there is nothing else to eat. In overgrazed land 

burro weed behaves as an invasive weed 

(Whitfield and Anderson 1938, Parker 1990). 

Benefits.—Burro weed provides cover and 

stabilizes soils that are denuded by overgrazing or 

other disturbance. It adds to the biodiversity of 

communities of which it is part and may be 

important for pollinators and other arthropods in 

areas that are depauperate in native species. 

References 

Arizona Veterinary Diagnostic Laboratory 

Quarterly. 1998. Equine 3 (4): 2. 

Blydenstein, J. 1966. Root systems of four desert 

grassland species on grazed and protected sites. 

Journal of Range Management 19: 93-95. 

Bowers, J.E. 1993. Shrubs and trees of the 

southwest deserts. Southwest Parks and 

Monuments Assoc., Tucson, AZ. 140 p. 

Bowers, J.E. and S.P. McLaughlin. 1987. Flora 

and vegetation of the Rincon Mountains, Pima 

County, Arizona. Desert Plants 8(2): 51-94. 

Cable, D.R. 1967. Fire effects on semidesert 

grasses and shrubs. Journal of Range 

Management 20: 170-176. 

Dayton, W.A. 1931. Important western browse 

plants. Misc. Publ. 101. U.S. Department of 

Agriculture. Washington, DC. 214 p. 

Elmore, F.H. 1987. Shrubs and Trees of the 

Southwest Uplands. Southwest Parks and 

Monuments Assoc. Tucson, AZ. 214 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. LewAnn Publishing Company, Mesa, 

AZ. 347 p. 

Goldberg, D.E. and R.M. Turner. 1986. Vegetation 

change and plant demography in permanent 

plots in the Sonoran Desert. Ecology 67(3): 695-

712. 

Ivey, R.D. 1995. Flowering plants of New 

Mexico. 3rd Ed. Rio Rancho Printing, 

Albuquerque, NM. 504 p. 

Kearney, T.H., R.H. Peebles, and collaborators. 

1960. Arizona flora. 2nd Ed. University of 

California Press, Berkeley, CA. 1,085 p. 

Kingsbury, J.M. 1964. Poisonous Plants of the 

United States and Canada, Prentice-Hall, Inc., 

Englewood Cliffs, NJ. 626 p. 

Lane, M.A. and R.L. Hartman. 1996. 

Reclassification of North American 

Haplopappus (Compositae: Asteraceae) 

Completed: Rayjacksonia gen. nov. American 

Journal of Botany 83(3): 356-370. 

Martin, W.C. and C.R. Hutchins. 1980. A Flora of 

New Mexico. Strauss & Cramer, Hirschberg, 

Germany. 2,590 p. 

McPherson, G.R. 1995. The role of fire in desert 

grasslands. In: M.P. McClaran and T.R. Van 

Devender, eds. The desert grassland. The 

University of Arizona Press, Tucson, AZ. p. 

130-151. 

Neson, G.L. 1991. Taxonomy of Isocoma 

(Compositae: Asteraceae). Phytologia 70: 69-

114. 
390 

Parker, K.F. 1990. An illustrated guide to Arizona 

weeds. The University of Arizona Press, 

Tucson, AZ. 338 p. 

Pima Community College. 2001. Internet site. 

http://wc.pima.edu/Bfiero/tucsonecology/plants/

shrubs_bw.htm [not paged]. 

Schmutz E.M., E.L Smith, P.R. Ogden, M.L. Cox, 

J.O. Klemmedson, J.J. Norris, and L.C. Fierro. 

1992. Desert grasslands. In: R.T. Coupland, ed. 

Ecosystems of the World, 8A; Natural 

grasslands: Introduction and Western 

Hemisphere. Elsevier, London, UK. p. 337-362. 

Taylor, R.J. 1998. Desert wildflowers of North 

America. Mountain Press Publishing Co., 

Missoula, MT. 349 p. 

United States Department of Agriculture, Forest 

Service. 1988. Range Plant Handbook. Dover 

Publications, Inc. New York. 837 p. 

White, L.D. 1969. Effects of a wildfire on several 

desert grassland shrub species. Journal of Range 

Management 22: 284-285. 

Whitfield, C.J and H.L. Anderson. 1938. 

Secondary succession in the desert plains 

grassland. Ecology 19 (2): 171-180. 
391

Itea virginica L. Virginia sweetspire 

SAXIFRAGACEAE 

Synonyms: None 

Kristina Connor 

Illustration source: U.S. Department of Agriculture [No 

date] 

General Description.—Virginia sweetspire, also 

known as sweetspire, tassel-white, Virginia tea, or 

Virginia willow, is an upright, multistemmed, 

deciduous or semievergreen shrub with arching 

branches (Bailey and Bailey 1976, Krüssmann 

1984, Odenwald and others 1996). The branches 

are light green or brown, pubescent when young, 

while old stems are glabrous, gray and striated. 

Twigs are slender and may be red when young 

(Krüssmann 1984) or in the fall (Seiler and 

Peterson 2001). The medium to dark green, 

simple, alternate leaves are elliptic to oblong in 

shape, glabrous above and finely pubescent 

beneath, four to 10 cm long, and have finely 

toothed margins. The plants are noted for their 

racemes of white flowers in early summer and 

their brilliant red to reddish purple foliage in 

autumn. Leaves can remain on the plant in a mild 

winter or in the more southern hardiness zones. 

Virginia sweetspire can occasionally attain the 

form of a small tree but is generally multistemmed 

and reaches a height between 0.9 and 2.4 m. This 

species has also been placed in the Grossulariaceae 

and Iteaceae (USDA-ARS-GRIN 2001, USDANRCS 2001). 

Range.—Virginia sweetspire is found throughout 

the Southeastern United States, from the Atlantic 

to eastern Texas and Oklahoma, although it is 

uncommon in the latter state (Johnson and 

Hoagland 1999). It can be found in wetlands as far 

north as Maryland, Delaware, and New Jersey and 

as far south as Florida. It is also found in southern 

Illinois and throughout the Mississippi River 

Valley. 

Ecology.—Virginia sweetspire grows primarily in 

wet areas, in swamps, along stream beds and lake 

edges, and in floodplains where it can form 

thickets. However, it can thrive in a variety of soil 

types and pHs and also shows tolerance to 

droughty conditions (Bailey and Bailey 1976, 

Odenwald and others 1996, Ohio State University 

2002). The species is tolerant of various light 

regimes and will grow well in both the sun and the 

shade; however, it can become leggy and sparse in 

the shade. It flowers more prolifically and has 

optimal fall coloration when grown in full sun. 

The species is highly adaptable and tough. 

Reproduction.—Virginia sweetspire is named for 

its drooping to upright 5 to 15 cm-long pubescent, 

terminal racemes of fragrant, white flowers that 

occur from early spring to early summer. 

Individual flowers have five petals and five sepals 

and are small, only 6 to 9 mm across (Johnson and 

Hoagland 1999). Flowering period is fairly long, 

lasting from May to early July (Krüssmann 1984). 

Brown seeds are produced in small elongate 

capsules with cranesbill-like projections (The Ohio 

State University 2002). Capsules can remain on 

the plant long after leaf fall. Seeds are quick to 

germinate and reportedly require no pretreatment 

for germination to occur. Virginia sweetspire can 

be propagated by rooting cuttings, from seed, or 

from root suckers (Bailey and Bailey 1976). 
392 

Growth and Management.—Virginia sweetspire 

is semievergreen and fairly long lived. With its 

long flowering period and attractive fall foliage, 

the species can do well in landscape plantings. Its 

adaptability to both sun and shade, and to wet and 

dry areas, gives it a versatility lacking in some 

other shrub species. However, it can grow beyond 

desired boundaries because it spreads by root 

suckers, and if not pruned it will form thickets. It 

grows fairly rapidly and easily from basal shoots 

and is relatively free from insect pests and diseases 

(The Ohio State University 2002). 

Benefits.—The dense thickets of Virginia 

sweetspire provide cover for wildlife (Silberhorn 

1996). Although the dry fruit capsules and small 

seeds are not a preferred food, they are eaten by 

birds. The species can be used for erosion control 

on wet or sloped sites. The fragrant flowers are 

attractive to butterflies. 

References

Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: A 

Concise Dictionary of Plants Cultivated in the 

United States and Canada. McMillan Publishing 

Co., Inc., New York. 1,312 p. 

Johnson, F.L. and B.W. Hoagland. 1999. Itea 

virginica. Catalog of the Woody Plants of 

Oklahoma, Oklahoma Biological Survey. 

http://www.biosurvey.ou.edu. 1 p. 

Krüssmann, G. 1976. Manual of Cultivated BroadLeaved Trees and Shrubs. Volume I, A-D. Timber 

Press, Beaverton, OR. 448 p. 

Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope. 

1996. Plants for American Landscapes. Louisiana 

State Univ. Press, Baton Rouge, LA. 266 p. 

Ohio State University. 2002. Horticulture and Crop 

Science in Virtual Perspective. 

http://www.hcs.ohio-state.edu/hcs/TMI/ 

Plantlist/it_inica.html. 4 p. 

Seiler, J.R. and J.A. Peterson. 2001. Sweetspire. 

Virginia Tech. Dendrology Webpage 

http://www.fw.vt.edu/ 

dendro/dendrology/syllabus/ivirginica.htm. 1 p. 

Silberhorn, G. 1996. Virginia willow (Itea 

virginica). Technical Report. Wetlands Flora. No. 

96-5. Virginia Institute of Science, School of 

Marine Science, College of William and Mary, 

Gloucester Point, VA. 2 p. 

U.S. Department of Agriculture. [no date]. 

Southern wetland flora: Field office guide to 

plant species. U. S. Department of Agriculture, 

Soil Conservation Service, South National 

Technical Center, Fort Worth, TX. Jamestown, 

ND: Northern Prairie Wildlife Research Center 

Home Page. http://www.npwrc.usgs.gov. [not 

paged]. 

USDA-ARS-GRIN. 2001. Germplasm Resources 

Information Network, National Germplasm 

Resources Laboratory. Plant Sciences Institute, 

Beltsville, Agricultural Research Center, 

Agricultural Research Service, U.S. Department of 

Agriculture. http://www.ars-grin.gov. [not paged]. 

USDA, NRCS. 2001. The PLANTS database, 

Version 3.1. National Plant Data Center, Baton 

Rouge, LA. http://plants.usda.gov. [not paged]. 
393

Ixora coccinea L. jungle-geranium

RUBIACEAE 

Synonyms: Ixora grandiflora Bot. 

 Ixora bandhuca Roxbg. 

John K. Francis 

General Description.—The common name of this 

plant, jungle-geranium, seems to be in slightly 

more common use than others: flame of the woods, 

flame of the forest, jungle flame, burning love, red 

ixora, ixora, amor ardiente, cruz de Malta, bola de 

coral, equisósea, and rajana (Griffiths 1994, 

Liogier 1997). Jungle-geranium is a densely 

branched shrub to 3 m in height and 3 or 4 cm in 

basal diameter. The stems and branches are gray. 

The plant produces a strong taproot with fine 

laterals. Mycorrhizal bifurcation is visible on the 

tips of fine roots. The dark green, oblong leaves are 

sessile or subsessile and hairless or nearly so 

(Liogier 1997). The inflorescences are terminal, 

dense corymbs that contain from 15 to 50 flowers 

(Whistler 2000). The flowers are tubular with four 

or five calyx lobes. The “wild” flower color is red 

or red-orange, but ornamental varieties may have 

white, yellow, salmon, or pink flowers. There are 

also dwarf varieties. The fruits are fleshy, globose 

berries that ripen to a dark red or purplish-black 

(Liogier 1997). When fully developed, there are 

two seeds per fruit. However, usually one 

develops and one aborts in Puerto Rican plants. 

Range.—Jungle-geranium is native to India and 

Sri Lanka (Griffiths 1994). It is planted worldwide 

in tropical and subtropical climates and has 

naturalized in at least Puerto Rico (author’s 

observation, Liogier 1997). 

Ecology.—Jungle-geranium grows well in full sun 

and light to moderate shade. It competes well with 

shrubs and herbs but not with dense grass. Little is 

known about environmental conditions in its native 

range, but it grows naturally in Puerto Rico in areas 

that receive around 1800 mm of annual 

precipitation. Jungle-geranium grows well in all 

textured soils if moisture is adequate and drainage 

is good. The species prefers slightly acid soils with 

good fertility and becomes chlorotic under alkaline 

conditions. It tolerates mild salt spray (Watkins and 

Sheeham 1975). Jungle geranium is cold sensitive 

and will partially defoliate after chilling (Michaelia 

and others 1999). Frosts will kill it to the ground, 

but it recovers afterward (Tropilab 2001). Junglegeranium in Florida is damaged by nematodes, root 

rot, leaf spots, and scale insects (Florida Gardener 

2001). 

Reproduction.—Jungle-geranium flowers 

throughout the year in full sunlight and light shade 

(Whistler 2000). Shrubs in heavy shade survive but 

rarely flower. A collection of fruits in Puerto Rico 

averaged 0.618 + 0.015 g/fruit. Air-dry seeds from 

these fruits averaged 0.067 + 0.001 g or 15,000 

seeds/kg. Planted in commercial potting mix, 70 

percent of these seeds germinated between 30 and 

72 days following sowing. Jungle-geranium 

coppices when cut or burned and prostrate plants 

send up numerous vertical shoots. Frequent 

suckering occurs from lateral roots out to a 

distance of 30 cm or more from the parent plant. It 

also layers whenever a branch touches the ground. 

Jungle-geranium can be easily propagated from 

untreated cuttings with four leaves placed in a 

sand-peat mixture with bottom heat (Bailey 1941). 

Jungle-geranium is spreading slowly in Puerto 

Rico, presumably dispersed by birds. 

Growth and Management.—Suckers and sprouts 

may grow as much as 1 m the first year, after 

which growth slows. Individual stems live at least 

15 years; by coppicing and suckering, a plant may 

live almost indefinitely. Maintenance of junglegeranium in hedges requires frequent pruning. The 

shrub is difficult to kill by mechanical means. 

Benefits.—Jungle-geranium is one of the world’s 
394 

most popular tropical flowering shrubs. It is used 

for hedges and borders, for accent plants, in 

planters and as an indoor potted plant. The species 

may also be shaped into bonsais (Frommer 2001). 

In laboratory tests, extracts of jungle-geranium 

have shown antibacterial (Kumer and others 1997) 

and antitumor (Latha and Panikkar 2000, Serrame 

and Lim-Sylianco 1995) activity. From a methanol 

extract of the flowers, 13 chemicals were 

identified, including ursolic acid, which has known 

antitumor and antiviral activity (Monteath and 

others 2001). In the traditional medicine of India, 

infusions of the leaves and juice from the roots are 

used to treat a wide variety of ailments including 

dysentery, ulcers, and gonorrhea (Parrotta 2001). 

The flowers are visited by butterflies (Collins 

2001), and birds eat the fruits. 

References 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. The MacMillan Company, New 

York. 3,639 p. 

Collins, M. 2001. Marge Collins flying flowers: 

Marge’s story. http://www.ffgc.org/horticulture 

/flgardens/collins/garden_collins.htm. 4 p. 

Florida Gardener. 2001. Plant of the month: Ixora 

coccinea. http://www.floridagardener.com 

/ixora.htm. 2 p. 

Frommer A. 2001. Growing an indoor bonsai. 

Arthur Frommer’s Budget Travel Newsletter. 

http://www.dummies.com/Lifestyle/ 

Home_and_Garden/Healthy.../0-7645-5102-

7_0019.htm. 4 p. 

Griffiths, M. 1994. Index of garden plants. Timber 

Press, Portland, OR. 1,234 p. 

Kumer, T.K., C.E. Rajini, V.K. Sasidharan, and T. 

Krishna-Kumar. 1997. Antibacterial and 

antifungal activity of secondary metabolites 

from some medicinal and other common plant 

species. Journal of Life Science 2: 14-19. 

Latha, P.G. and K.R. Panikkar. 2000. Inhibition of 

chemical carcinogenesis in mice by Ixora 

coccinea flowers. Pharmaceutical Biology 38(2): 

152-156. 

Liogier, H. A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Michaelia, R., S. Philosoph-Hadasa, J. Riovb, and 

S. Meira. 1999. Chilling-induced leaf abscission 

of Ixora coccinea plants. I. Induction by 

oxidative stress via increased sensitivity to 

ethylene. Physiologia Plantarum 107(2): 166-

173. 

Monteath, S.A.F.A., V.V. Veiga, A.C. Pinto, A. 

Echevarria, and M.A.M. Maciel. 2001. 

Constituentes químicos das flores de Ixora 

coccinea. Universidade Federal Rural do Rio de 

Janeiro. http://www.sbq.org.br/ranteriores/23/ 

resumos /1147. 3 p. 

Parrotta, J.A. 2001. Healing plants of peninsular 

India. CAB International, Wallingford, UK and 

New York. 944 p. 

Serrame, E. and C.Y. Lim-Sylianco. 1995. Antitumor promoting activity of decoctions and 

expressed juices from Philippine medicinal 

plants. Philippine Journal of Science 124(3): 

275-281. 

Tropilab. 2001. Ixora coccinea L. Tropilab Inc. 

http://www.tropilab.com/jungle-ger.html. 1 p. 

Watkins, J.V. and T.J. Sheeham. 1975. Florida 

landscape plants, native and exotic. The 

University Presses of Florida, Gainesville, FL. 

420 p. 

Whistler, W.A. 2000. Tropical ornamentals, a 

guide. Timber Press, Portland, OR. 542 p. 
395

Jacquinia arborea Vahl barbasco 

THEOPHRASTACEAE 

Synonyms: Jacquinia armillaris Jacq. 

 Jacquinia barbasco Mez in Engl. 

 Chrysophyllum barbasco Loefl. 

John K. Francis 

General Description.—Barbasco, also known as 

torchwood, azúcares, and bizcocho, is an evergreen 

shrub or small tree to 4.5 m in height and 15 cm in 

stem diameter. It has a frequently thickened 

rootstalk and one or a few stems. The bark is dark 

brown and finely fissured, and the inner bark is 

light brown and bitter. The wood is light brown or 

yellowish and hard. The crowns are compact and 

often wind-hedged in their sea-side habitat. The 

thick and stiff, dull yellow-green leaves are 

spatulate or obovate, alternate or in threes, and 

clustered at the ends of the twigs. Terminal 

racemes are several to many-flowered. The small, 

white or yellowish, five-lobbed flowers are 

fragrant. The fruit (berries) are globose 8 to 12 mm 

in diameter, orange-red, and contain one to four 

rounded, brown seeds (Howard 1989, Liogier 

1995, Little and others 1974). 

Range.—There has been confusion about the 

botanic identity of barbasco. Stahl (1992) 

concluded that three species should be recognized 

from the J. armillaris group: J. keyensis Mez from 

the Northern West Indies (Florida and the 

Bahamas), J. arborea in the Central Caribbean, and 

J. armillaris Jacq. in the Lesser Antilles, northern 

South America, and eastern Brazil. Stearn (1992) 

gives the range of barbasco as the Cayman Islands, 

Jamaica, Hispaniola, Puerto Rico, the Virgin 

Islands, the Lesser Antilles, Trinidad, and Curaçao. 

He has J. arborea and J. armillaris coexisting in 

the Lesser Antilles. The Missouri Botanical Garden 

Herbarium lists specimens for Jamaica, Puerto 

Rico, the Virgin Islands, Quintana Roo, Mexico, 

and Honduras (Missouri Botanical Garden 2002). 

Barbasco is reported to have naturalized on Key 

Largo, Florida (Nelson 1996). 

Ecology.—Barbasco is relatively common along 

undisturbed coasts. Because the species is 

intolerant of shade and not very competitive, it 

usually grows on rocks and rocky headlands, 

exposed ridges and slopes, and dry coastal forests. 

In Puerto Rico, barbasco grows in areas that 

receive from 750 to 1700 mm of annual 

precipitation at elevations from near sea level to 

100 m. It is tolerant of heavy salt spray and 

moderately salty soils. Barbasco grows on welldrained, mildly acid to mildly alkaline soils of all 

textures derived from both sedimentary and 

igneous rocks. It is frequently seen growing out of 

cracks in limestone rocks. The species appears to 

have no major insect or disease problems. 

Reproduction.—Barbasco flowers from winter to 

summer and matures fruits from spring to fall 

(Little and others 1974). Fruits collected in Puerto 

Rico averaged 0.254 + 0.005 g/fruit. Air-dried 

seeds cleaned from those fruits averaged 0.0363 +

0.0007 g/seed or 27,500 seeds/kg. Sown without 

any pre-treatment in commercial potting mix, 100 

percent of the seeds germinated between 40 and 60 

days of sowing. Seeds are easily collected by hand 

and can be cleaned by maceration and wet sieving. 

The seeds are probably dispersed by birds. 

Seedlings are not common. 
396 

Growth and Management.—Growth of barbasco 

seedlings in the nursery is very slow. A group of 

seedlings averaged only 6.2 cm 1 year after 

pricking into containers. However, survival was 

relatively good (82 percent). Growth of nativegrown plants of all ages is also believed to be very 

slow. They seem to be long-lived. Probably the 

best strategy for management of native stands is 

strict protection from fire, harvesting, and 

development. 

Benefits.—Barbasco helps hold the soil in 

topography where there is little soil to lose and 

contributes significantly to the aesthetics of island 

coastlines. The fruits were once used to poison or 

stupefy fish and because of that are suspected to be 

poisonous to humans (Little and others 1974). The 

sister species J. keyensis is used as an ornamental 

(Nelson 1996), and probably barbasco has been 

used as well. It is a pretty and well-shaped plant, 

suited for borders and low backgrounds. 

References 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Missouri Botanical Garden. 2002. Current 

specimen list for Jacquinia arborea. 

http://mobot.org/cgi-bin/search_vast. 1 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

 391 p. 

Stahl, B. 1992. On the identity of Jaquinia 

armillaris (Theophrastaceae) and related 

species. Brittonia 44 (1): 54-60. 

Stearn, W.T. 1992. The genus Jacquinia

(Theophrastaceae) in Jamaica. Nordic Journal of 

Botany 12(2): 231-238. 
397

Jasminum fluminense Vell. Brazilian jasmine 

OLEACEAE 

Synonyms: Jasminum azoricum auct., not L. 

 Jasminum bahiense DC. 

John K. Francis 

General Description.—This plant became known 

as Brazilian jasmine because it was first described 

from Brazil. It is of African origin, having been 

introduced earlier by Portuguese settlers (Florida 

Exotic Plant Council 2001). The species is also 

known as jasmine, Gold Coast jasmine, jasmín 

oloroso, jasmín de canario, jasmín de trapo, jasmin 

à bouguet, and jasmin blanc (Acevedo-Rodríguez 

1985, Howard 1989, Pacific Islands Ecosystems at 

Risk 2001). Brazilian jasmine is a woody 

evergreen scrambling and climbing shrub (vine) 

that may extend as much as 12 m along the ground 

or into the crowns of trees. The cylindrical stems, 

which are commonly about 1 cm in diameter but 

may grow to 5 cm, have gray, fine-furrowed bark 

and are flexible and strong. There are often 

multiple stems from a root crown. Foliage is borne 

only on the current year’s growth. The opposite, 

compound leaves have three leaflets, the central 

one being larger than the lateral two. The 

inflorescences are cymes that arise from leaf axils. 

The tubular white flowers are fragrant, especially 

at night. The fruits, which form in groups of two, 

are globose, 8 mm in diameter, and dark purple or 

dark blue to almost black when ripe (AcevedoRodríguez 1985, Howard 1989, Liogier 1995). The 

fruits have a bitter, disagreeable flavor. Each fruit 

contains one spherical gray seed. 

Range.—The African and Middle Eastern native 

range of Brazilian jasmine includes Mauritius, the 

Seychelles, Arabia, Ethiopia, southern Zimbabwe, 

Malawi, Mozambique, Angola, Nigeria, South 

Africa (Miami-Dade County 2001), the Azores, 

and the Canary Islands (Acevedo-Rodríguez 1985). 

The species has naturalized and escaped 

throughout much of the tropics and subtropics 

including southern Florida (Florida Exotic Plant 

Council 2001), Puerto Rico (Liogier 1995), 

Hawaii, and Guam (Pacific Island Ecosystems at 

Risk 2001). 

Ecology.—Brazilian jasmine occurs in Puerto Rico 

in areas that receive from about 750 to 1800 mm of 

annual rainfall. It grows from near sea level to 

more than 600 m in elevation. Soils of all textures 

and parent material are colonized. However, it does 

not tolerate poorly drained soils. The species is 

restricted to areas with minimum temperatures 

above 1.7 °C (Florida Exotic Plant Council 2001). 

It will grow on the coast in areas that do not 

receive salt spray (Florida Exotic Plant Council 

2001). Brazilian jasmine will grow in partial shade 

and climb upward to better light. It can survive but 

is not aggressive in the denser shade of unbroken 

forest canopies. Most plants grow in natural and 

artificial openings in the forest such as fencerows, 

river banks, roadsides, brushy pastures, and logged 

or burnt-over forest. 

Reproduction.—Brazilian jasmine blooms 

throughout the year (Miami-Dade County 2001). 

Under favorable conditions, fruits are produced in 

abundance by open-grown plants. Shaded plants 

produce few or no fruits. A collection of fruits of 

Brazilian jasmine from Puerto Rico weighed an 

average of 0.413 ± 0.066 g each. Air-dried seeds 

from those fruits averaged 0.164 + 0.002 g/seed or 

6,000 seeds/kg. Sown on commercial potting mix, 

100 percent of the seeds collected from this sample 

germinated between 18 and 52 days after sowing. 

Brazilian jasmine roots (layers) whenever stems 

come in contact with the ground. The seeds are 

dispersed by birds and mammals (Miami-Dade 

County 2001) and by the lateral extension of the 

stems. Seedlings are common in suitable habitat in 

Puerto Rico. Ornamental plants are propagated by 
398 

means of cuttings (Florida Exotic Plant Council 

2001). 

Growth and Management.—The stems of 

established plants may extend as much as 2 or 3 m 

in 1 year. Seedlings grow much more slowly. 

Brazilian jasmine can engulf shrubs, small trees, 

and fences and can ascend power poles. Control of 

thickets and mats is often needed. Young plants 

can be pulled up by hand. Older plants should be 

cut at the ground level and the stumps treated with 

herbicide. Follow-up treatments will probably be 

required (Pacific Island Ecosystems at Risk 2001).

Benefits.—Brazilian jasmine has been widely 

planted as an ornamental, both in greenhouses and 

outdoors (Bailey 1941, Liogier 1995). Because of 

its invasive nature, the species should not be 

planted where it might escape to the wild (Nelson 

1996). It is a food source for birds and mammals. 

The fruits are heavily consumed by raccoons in 

Florida (Miami-Dade County 2001). Goats browse 

the foliage. 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. The MacMillan Company, New 

York, NY. 3,639 p. 

Florida Exotic Plant Council. 2001. Jasminum 

fluminense Vell. www.fleppc. 

org/pdf/Jasminum%20fluminense.pdf. 2 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Dicotyledoneae. 

Vol. 6. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 617 

p. 

Miami-Dade County. 2001. Brazilian jasmineJasminum fluminense. www.co.miamidade.fl.us/derm/environment/badplants/plant%2

0.../brazilian_jasmine.ht. 1 p. 

Nelson, Gil. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc. Sarasota, FL. 

 391 p. 

Pacific Island Ecosystems at Rick. 2001. Invasive 

plant species: Jasminum fluminense Vell., 

Oleaceae. www.hear.org/pier/jaflu.htm. 2 p. 
399

Jatropha gossypiifolia L. red physic nut 

EUPHORBIACEAE 

Synonyms: Adenoropium gossypifolium Pohl 

 Jatropha elegans (Pohl) Klotzsch in Seemann 

John K. Francis 

General Description.—Red physic nut, also 

known as belly-ache bush, figus nut, wild cassava, 

higuereta cimarrona, tautuba, túatúa, purga del 

fraile, médicinier noir, médicinier bâtard, 

médicinier rouge, is a short-lived perennial shrub 

0.5 to 2 m in height and 1 to 3 cm in basal 

diameter. It may be evergreen or deciduous, 

depending on climate. The plant usually has a 

single stem and is woody at the base. The small 

number of branches are stout, green, and semiwoody. A cloudy, yellow sap collects when stems 

and branches are cut. Red physic nut has a short 

taproot, robust laterals, and many fine tertiary roots 

and is usually not deeply rooted. The opposite 

leaves are three- or five-lobed, 3 to 13 by 3 to 19 

cm, reddish or purplish tinged when young and 

green when fully developed. Small red or purple 

flowers are borne in terminal corymbose cymes. 

The fruits that follow are oblong, three-lobed 

capsules that are green, turning tan at maturity. 

Within are three mottled, gray-brown or grayishred seeds (Burkill 1994, Howard 1989, Liogier 

1988, Parrotta 2001). 

Range.—Red physic nut is a native of the New 

World (Burkill 1994). The original range is not 

known, but today it grows wild from Florida 

through the West Indies and from Mexico through 

tropical South America (Howard 1989). It has 

naturalized in most tropical areas of the world 

(Burkill 1994). 

Ecology.—Red physic nut grows on nearly all 

types of soils within its range. In Puerto Rico, it 

may be found in areas receiving from 750 to about 

2000 mm of annual precipitation. The species is 

more common in soils with high base saturation, 

such as dry areas, sites near the ocean, and soils 

derived from limestone. Red physic nut usually 

requires disturbance to establish itself and does not 

compete well with tall grass or heavy weed growth. 

It is intolerant of shade. Although plants may 

survive for a season in moderate shade, they need 

full or nearly full sun for longer-term survival and 

fruiting. Cattle and horses generally avoid red 

physic nut, except in extreme shortages of forage, 

so that the species in often abundant in overgrazed 

range and poorly managed pastures. It is also 

common in waste lands, roadsides, poorly tended 

agricultural fields, and river overflow areas. 

Reproduction.—Flowering in India occurs from 

February through July (Parrotta 2001). Sometimes 

both flowers and fruits will be present at the same 

time on plants. Upon drying, the capsule valves 

spring open propelling the seeds a few centimeters. 

It is not known if there are other seed dispersal 

mechanisms. Seeds collected in Puerto Rico 

averaged 0.017 ± 0.001 g/seed or 59,000 seeds/kg. 

The seeds of this collection were highly variable in 

weight (C.V.= 53.7 percent) and just 4 percent 

germinated. Collecting seed in quantity is difficult 

because capsules are mature for only a short period 

before discharging their seeds, and there are only a 

few capsules per large plant at any time. 

Growth and Management.—Red physic nut 

usually grows without branches during its first year 

and adds adventitious branches in the year or two 

following. Growth is about 0.5 m per year and 

plants usually live 2 or 3 years. The species is 
400 

considered an agricultural weed but can be 

suppressed by common weed-control practices. 

Detriments and Benefits.—Red physic nut 

contains toxins in the seeds, sap and other tissues, 

capable of killing humans (Marcano-Fondeur 

2002). Apparently, poisoning cases are rare. 

Extracts of the plant are used as a purgative and 

emetic, and to treat headache, diarrhea, venereal 

disease, skin sores, mouth sores, and cancer 

(Burkill 1994, Parrotta 2001). The use of the seeds 

in herbal medicine is advised against because of 

their high toxicity (Liogier 1990). Phytoactive 

chemicals found in tissues of red physic nut 

include the protease curcain, the alkaloid 

jatrophine, the cyclic heptapeptide, cyclogossine A, 

saponins, and numerous fatty acids (Burkill 1994, 

Horsten and others 1996, Marcano-Fondeur 1992, 

Ogbobe and Akano 1993). The seeds contain 36 

percent oil (Ogbobe and Akano 1993) and are used 

as an illuminant in Africa (Burkill 1994). Swaths of 

plants are frequently established around villages in 

Africa as fire barriers. The species is also attributed 

magical powers that protect against lightning, 

snakes, and violence (Burkill 1994). The wood is 

soft and weak and of little use. A red-leafed 

variety is used as an ornamental (Howard 1989). 

References 

Burkill, H.M. 1994. The useful plants of West 

Tropical Africa. Vol. 2. Royal Botanic Gardens, 

Kew, UK. 636 p. 

Horsten, S.F., A.J. van den Berg, J.J. Kettenes-van 

den Bosch, B.R. Leeflang, and R.P. Lababie. 

1996. Cyclogossine A: a novel cyclic 

heptapeptide isolated from the latex of Jatropha 

gossypifolia. Planta Med 62(1): 46-50. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Marcano-Fondeur, E. de J. 1992. Las plantas 

venenosas en la medicina popular. Naturaleza 

Dominicana. http://marcano.freeservers.com/ 

nature/conference/venen.html. 6 p. 

Ogbobe, O. and V. Akano. 1993. The physicochemical properties of the seed and seed oil of 

Jatropha gossipifolia. Plant Foods for Human 

Nutrition 43(3): 197-200. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CABI Publishing, Wallingford, UK and 

New York. 917 p. 
401

Juniperus communis L. common juniper 

CUPRESSACEAE 

Synonyms: Juniperus canadensis Lodd. ex Burgsd. 

Juniperus alpina (Sm.) S.F. Gray 

Juniperus sibirica Burgsd. 

Juanita A. R. Ladyman 

Illustration source: USDA-Forest Service collection, 

Hunt Institute 

General Description.—Common juniper is 

known by a variety of common names: dwarf, low, 

mountain, alpine, prostrate or Siberian juniper, 

ground cedar, genevrier commun, and enebro 

(Elmore 1987, FNA 1993). There are several subspecies or varieties, each having several 

synonyms. For a list of the synonyms consult 

Kartesz (1994). Common juniper is a low shrub, 

with spreading or ascending branches, that 

generally grows no more than 1 m high. Larger 

individuals, up to 10 m, have been observed. It 

typically forms dense mats 1 to 4 m in diameter. 

The brown fibrous bark is smooth on small 

branches (<1 cm diameter) but exfoliates into thin 

strips on larger branches. The leaves are awlshaped, in groups of three and tend to be “prickly” 

standing out from the stems. Common juniper is 

listed as extremely rare or critically imperiled in 

many Southeastern States, including Georgia 

(Georgia Natural Heritage Program 2001) and 

Alabama (Alabama Natural Heritage Program 

1996), or of likely rarity in South Carolina (South 

Carolina Heritage Trust 2000). 

Range.—The range of common juniper is 

extensive. It is the only circumpolar conifer in the 

northern hemisphere (Pojar and Mackinnon 1994). 

It is found across the U.S.A. and Canada to 

Greenland, through Europe, across Siberia and 

Asia. However, within its range there are at least 

five sub-species or varieties (FNA 1993). For 

example, var. depressa occurs throughout much of 

Canada and the United States where it occurs in 

the Southwestern States of Arizona, New Mexico, 

Colorado, and Utah and extends west to California 

and east to North Carolina through Arkansas; var. 

megistocarpa occurs in Nova Scotia, 

Newfoundland, and Quebec; var. montana occurs 

in Greenland, British Columbia, and California, 

Oregon and Washington States (FNA 1993); var. 

communis and var. nana occur in Britain and 

Ireland (Stace 1997). Larger specimens of var. 

depressa in the United States have frequently been 

misidentified as var. communis (FNA 1993). 

Ecology.—Common juniper is a hardy shrub that 

grows in a wide range of ecological conditions. 

Typically it grows on dry, open, rocky slopes and 

mountainsides but may be found in stressed and 

acid environments where competition with other 

plants is almost non-existent (Ward 1982). It also 

often grows in partial shade. Depending upon the 

latitude it can be found from lowland bogs at sealevel to sub-alpine ridges and alpine tundra at over 

3,600 m (FNA 1993; Lanzara and Pizzetti 1977; 

Pojar and Mackinnon 1994). It is also a common 

shrub in abandoned lowland fields in Northern 

United States (Steele 1982). 

Reproduction.—Common juniper has berry-like 

female cones that mature in 2 years in 

Mediterranean climates and every 3 years in cooler 
402 

climates (Lanzara and Pizzetti 1977). Male and 

female cones are on separate plants. The dark blue 

seed (female) cones are globose to ovoid in shape, 

6 to 13 mm in diameter and are resinous to 

obscurely woody with two to three triangularshaped seeds. From pale green the female cones 

ripen to dark blue in late summer and then tend to 

whiten with a waxy “bloom.” The staminate cones 

are 3 to 6 mm in diameter (Cronquist and others 

1972). Although the cones are generally terminal 

in other species of juniper, in J. communis the 

cones are axillary (FNA 1993). The fruits are of 

relatively low nutritional quality and on average 

number 80,500 seeds/kg (Stiles 1980). The seed is 

dispersed by birds, at least in North America, and 

commonly require a long maturation and 

germination period (Crane and Fischer 1986; Stiles 

1980). Because seeds do not readily germinate, 

the establishment of seedlings, for example for revegetation projects, can be a problem (Dietz and 

others 1980, Stiles 1980). As well as poor 

germination, poor seed dispersal has been blamed 

for the relative rarity of juniper in areas where the 

majority of potential habitat is not occupied 

(Diotte and Bergeron 1989). Plant age also affects 

reproduction; 40 to 60 percent of old individuals 

are sterile and up to 94.8 percent of seeds of seedproducing old plants are non-viable. In contrast, 80 

percent of seeds produced by young plants are 

viable (Diotte and Bergeron 1989). 

Growth and Management.—Common juniper is 

often killed by fire (Crane and Fischer 1986). It 

has been described as having minimal “firesurviving regeneration properties," and resprouting 

after fire is rare (Mallik 1995). The foliage is 

resinous and flammable (Diotte and Bergeron 

1989). Mortality depends upon fire intensity. For 

example, in eastern Canada, older common juniper 

shrubs will survive fires of low severity, and some 

fire regimes allow common juniper to survive 

several fires; individuals of more that 170 years 

old have been observed to survive in areas exposed 

to fire (Diotte and Bergeron 1989). However, it 

should be noted that the patchy nature of fires 

leads to refugia for juniper shrubs, and so 

particular individuals can escape from harm and 

provide seed for re-vegetation of the burnt areas 

(Diotte and Bergeron 1989). Re-establishment of 

common juniper as a browse species after 

disturbance and fire has had varying success 

(Dietz and others 1980). In the Black Hills, South 

Dakota, the only successful means for establishing 

common juniper was from nursery stock, and seed 

germination was particularly poor. 

Benefits.—Common juniper has low value for 

short-term re-vegetation projects but moderate to 

high value for long-term rehabilitation projects 

and is useful in preventing soil erosion 

(Tirmenstein 1999). Common juniper provides 

important cover and browse for wildlife such as 

mule deer (Dusek 1975). The cones are eaten by 

several species of birds and are important food 

sources for wild turkeys (Decker and others 1991), 

Cedar and Bohemian waxwings (Catling and 

Brownell 1998), and others. Domestic livestock 

rarely utilize common juniper (Tirmenstein 1999). 

The seed cones of common juniper are used to 

flavor gin, and the word for this alcoholic 

beverage was derived from the Latin “iuiperus” 

through the Old French and Dutch words 

“genevre” and “genever” respectively (Elmore 

1987). The seed cones are also used as a flavoring 

for foods. Many Native American tribes used 

common juniper for medicinal and ceremonial 

purposes (Moerman 1998). It has been used 

medicinally for childbirth and to treat urinary 

infections (Pojar and Mackinnon 1994). However, 

it can also cause miscarriages. The different 

varieties of common juniper make excellent, 

vigorous landscaping shrubs, which are readily 

propagated by cuttings in the horticultural trade 

(Cronquist and others 1972). 

References 

Alabama Natural Heritage Program. 1996. 

NatureServe, Member Program--Plants of 

Alabama. Website Last Update: 8/1/96:http:// 

www.abi.org/nhp/us/al/plants.html 1 p. 

Catling, P.M. and V.R. Brownell. 1998. 

Importance of fire in Alvar EcosystemsEvidence from the Burnt Lands, Eastern 

Ontario. Canadian Field Naturalist 112(4): 661-

667. 

Crane M.F. and W.C. Fischer. 1986. Fire Ecology 

of the Forest habitat Types of Central Idaho. 

General Technical Report INT-218. USDA 

Forest Service Intermountain Research Station, 

Ogden, Utah. 85 p. 

Cronquist, A., A.H. Holmgren, N. H. Holmgren, 

and J.L. Reveal. 1972. Intermountain Flora. 

Volume 1. Vascular Plants of the Intermountain 

West, U.S.A. New York Botanical Garden. 

Hafner Publishing Co. New York. p. 238-240. 

Decker, S. R., P. J. Pekins, W. W. Mautz. 1991. 

Nutritional evaluation of winter foods of wild 
403

turkeys. Canadian Journal of Zoology. 69 (8): 

2,128-2,132. 

Dietz, R.D., D.W. Uresk, H.E. Messner, and L.C. 

McEwen. 1980. Establishment, Survival, and 

Growth of Selected Browse Species in a 

Ponderosa Pine Forest. General Technical 

Report RM-219. USDA Forest Service Rocky 

Mountain Forest and Range Experiment Station. 

Fort Collins, CO. 12 p 

Diotte, M. and Y. Bergeron. 1989. Fire and the 

distribution of Juniperus communis L. in the 

boreal forest of Quebec, Canada. Journal of 

Biogeography 16: 91-96. 

Dusek, L. 1975. Range relations of Mule Deer and 

cattle in prairie habitat. Journal of Wildlife 

Management. 39(3): 605-616. 

Elmore. F.H. 1987. Shrubs and Trees of the 

Southwest Uplands. Southwest Parks and 

Monuments Assoc. Tucson, AZ. 214 p. 

Flora of North America Editorial Committee. 

1993. Flora of North America. Vol. 2. Oxford 

University Press, New York. 475 p. 

Georgia Natural Heritage Program. 2001. 

http://www.dnr.state.ga.us/dnr/wild/natural/sppl

_t.htm. 1 p 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada, and 

Greenland. Vol.1 – Checklist. 2nd Ed. Timber 

Press, Portland, OR. 622 p. 

Lanzara, P. and M. Pizzetti. 1977. Simon and 

Schuster’s Guide to Trees. Simon and Schuster 

Inc., New York. 316 p. 

Mallik, A. U. 1995. Conversion of temperate 

forests into heaths: role of ecosystem 

disturbance and ericaceous plants. 

Environmental Management. 19(5): 675-684. 

Moerman, D.E. 1998. Native American 

Ethnobotany. Timber Press, Inc, Portland, OR. 

927 p. 

Pojar, J. and A. Mackinnon. 1994. Plants of the 

Pacific Northwest Coast. British Columbia 

Ministry of Forests and Lone Pine Publishing, 

Vancouver, Canada. 527 p. 

Stace, C. 1997. New Flora of the British Isles. 

Second ed. Cambride University Press, 

Cambridge, England. 1,130 p. 

Steele, F.L. 1982. At Timberline: A Nature Guide 

to the Mountains of the Northeast. Appalachian 

Mountain Club, Boston, MA. 285 p. 

South Carolina Heritage Trust. 2000. Website. 

http://www.abi.org/nhp/us/sc/speclist.htm. 1 p. 

Stiles, E.W. 1980. Patterns of fruit presentation and 

seed dispersal in bird-disseminated woody plants 

in the eastern deciduous forest. The American 

Naturalist. 116(5): 670-688. 

Tirmenstein 1999. Juniperus communis. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System, 

 http://www.fs.fed.us/database/feis/ [not paged]. 

Ward, L.K. 1982. The Conservation of Juniper. 

Journal of Applied Ecology. 19: 165-188. 
404 

Juniperus osteosperma (Torr.) Little Utah juniper 

CYPRESSACEAE 

Synonyms: Juniperus utahensis (Engelm.) Lemmon 

 Juniperus californica Carr. ssp. osteosperma (Torr.) E. Murr. 

 Juniperus californica Carr. var. utahensis Engelm. 

 Juniperus knightii A. Nels. 

 Juniperus megalosperma Sudworth 

 Juniperus occidentalis Hook. var. utahensis (Engelm.) Kent 

 Sabina osteosperma (Torr.) Antoine 

John K. Francis 

General Description.—Utah juniper, also known 

as cedar, desert cedar, sabina, and sabina morena, 

has a habit between that of shrub and tree. 

Although slow-growing and long-lived, rare 

individual plants attain large diameters (record 

tree: 217 cm diameter breast height and height 

11.6 m, American Forests 1998). The species is 

short statured, (mainly 2 to 4 m tall), (Welsh and 

others 1987), usually has multiple stems (limbs 

arising near the ground that become codominant 

with the original central stem), and becomes 

reproductively mature at small sizes. The bark is 

gray-brown or gray, thin, fibrous, and shredding. 

Sapwood is white and the heartwood is yellowbrown with narrow annual rings. It is relatively 

soft, brittle, and has a dry-weight specific gravity 

of 0.553 g/cm3

 (Paettie 1953). The wood and all 

other parts are aromatic. Crowns tend to be 

branchy, rounded to flat-topped, and sometimes 

dense; old trees often have bleached, dead limbs. 

Utah junipers have deep taproots and lateral roots 

that may extend as much as 30 m, a few cm below 

the soil surface. The roots are colonized by 

vasicular-arbuscular mycorrhizae. The 

photosynthetic organs are yellow-green branchlets 

with opposite (or whorled), reduced scale-like 

leaves. The species is monoecious with small, 

separate male and female flowers. Fruits are 

subglobose, 6- to 12-mm diameter, leathery cones 

(“berries”), brownish to blue-purple at maturity. 

They have a strong, resiny flavor and contain one 

(sometimes two) hard seed (Abrams 1940, Welsh 

and others 1987, Zlatnik 1999). 

Range.—Utah juniper occurs as large populations 

and scattered disjuncts in southern Idaho, southern 

Montana, Wyoming, western Colorado, Utah, 

Nevada, Arizona, eastern and southern California, 

northeastern New Mexico, and Chihuahua and 

Durango in Mexico (Natural Resources 

Conservation Service 2003, Secretaría de Medio 

Ambiente y Recursos Naturales 2003, Tree Guide 

2003). The species is not known to have 

naturalized outside its native range. It hybridizes 

with J. occidentalis Hook. where the species grow 

together (Terry and others 2000). 

Ecology.—Utah juniper grows on a wide variety 

of well-drained soils. It accommodates all the soil 

textures, pH’s of 6.5 to 8.2, high free calcium 

carbonate, but does not tolerate salty soils (Natural 

Resources Conservation Service 2003). It is 
405

commonly found on rocky, gravely, and sandy 

sites, usually originating from igneous (Stuart and 

Sawyer 2001) and sedimentary (sandstone and 

limestone) rocks, at 900 to 2,500 m above sea 

level (Zlatnik 1999). Mean annual precipitation 

varies from 300 to 510 mm, but the species can 

withstand severe droughts (Natural Resources 

Conservation Service 2003) and is a fierce 

competitor for the little available moisture in its 

habitat. It has a specific leaf area of 18 cm2

/g 

(Grier and others 1992). Utah juniper also tolerates 

cold and heat well. However, the species is 

intolerant of shade and does not survive under the 

canopy of taller vegetation. It is alleged to exert an 

allelopathic effect on some understory grass 

species. Utah juniper is a climax species in many 

vegetation types with species such as Artimisia 

tridentata Nutt., Pinus edulis Engelm., and Pinus 

monophylla Torr. & Frém. The range of Utah 

juniper is known to have expanded since European 

colonization, and stands have changed from open, 

savanna types to dense stands probably because of 

reduction of fine fuels by livestock grazing, fire 

suppression, and changes toward warmer and drier 

climates (Tausch 1999). Utah juniper is easily 

killed by fire, both by scorching the top or heatgirdling through the thin bark. However, the 

habitat seldom has sufficient fine fuels to sustain a 

fire. Stands most at risk are young stands with 

scattered trees in brush and grass, and continuous 

stands of large, decadent trees during high winds 

(Zlatnik 1999). Utah juniper is sometimes heavily 

infested by the mistletoes, Phoradendron 

juniperinum Engelm. ex Gray and P. bolleanum

(Seem.) Eichl. (Zlatnik 1999). The larvae of 

Eurytoma juniperina Marcovitch attack the seeds 

(Bonner 2003). 

Reproduction.—Utah juniper flowers from 

January to April (depending on site) and is wind 

pollinated. Fruits ripen in the second year and 

germinate the following spring or the second 

spring after dispersal (Zlatnik 1999). Fruits may be 

collected by hand from the shrubs or low trees or 

picked up by hand after they have fallen. Seed 

should be extracted by maceration with water 

before the fruits dry out. Cleaned seed from 

Arizona ranged from 7,900 to 15,700 seeds/kg. 

Seed should be dried to about 10 percent moisture 

and can be stored at 5 to -18 °C for several years. 

Moist stratification at 3 to 5 °C for 30 to 180 days 

is necessary for good germination. Drilled or 

broadcast in a nursery bed, seeds should be 

covered with 6 mm of fine soil or sand. 

Germination ranges from 50 to 64 percent (Bonner 

2003). 

Growth and Management.—Utah juniper grows 

slowly, usually only 0.13 cm/year in diameter. An 

86-year-old stand in Utah had 17.2 percent canopy 

cover and a basal area of 7.7 m2

/ha (Zlatnik 1999). 

Poor site conditions may result in 15 cm-tall plants 

over 50 years old. Individuals of the species may 

live 650 years or longer. Closing Utah juniper 

stands eliminate most forage plants; killing trees 

can result in a doubling of herbaceous cover 

(Zlatnik 1999). Controlling Utah juniper has been 

attempted with fire, but fires on these sites are 

difficult to start under humid, still conditions and 

hard to control under dry, windy conditions. 

Controlled burns can be done with the greatest 

precision in late winter or early spring when the 

wood is low in water content (personal 

communication with Stanley Kitchen, Shrub 

Sciences Laboratory, USDA Forest Service, 

Provo, UT). The shrubs and trees can be killed 

with chainsaws or herbicides, but the preferred 

method is “chaining,” dragging sections of anchor 

chain between two bulldozers. Many young trees 

are not killed, but the treatment does open stands 

and allows greater shrub and herbaceous growth 

(Plummer and others 1968, Stevens 1999). 

Benefits.—Utah juniper is a major constituent of 

semidry ecosystems of the West. It protects the 

soil from wind erosion but in some instances may 

aggravate sheet erosion from rains by suppressing 

understory grasses, forbs, and shrubs. The species 

furnishes cover and reproductive habitat for many 

species and is a major source of food for wildlife, 

especially during winter. Many birds, rodents, as 

well as jackrabbits (Lepus californicus) and 

coyotes (Canis latrans) eat the fruits and mule 

deer (Odocoileus hemionus), and elk (Cervus 

elaphus) browse the foliage when better food is 

not available. It is not eaten by cattle and is 

browsed to a limited extent by domestic sheep 

(Stuart and Sawyer 2001, Zlatnik 1999). The 

forage is of low quality because of low 

digestibility (44 percent) and low protein content 

(6 to 8 percent) (Zlatnik 1999). The wood is used 

for firewood, charcoal, and novelties (Secretaría de 

Medio Ambiente y Recursos Naturales 2003). The 

bark is used as tinder and formerly was used as a 

“slow match” to maintain and transport fire (Plants 

For a Future 2003). Native Americans used the 

bark to make sandals, mats, bedding, baskets, 

thatch, and cordage, and they made meal from the 

fruits (Peattie 1953). The wood is highly decay 

resistant, which makes it desirable for fence posts 

and rude construction despite its poor form 

(Zlatnik 1999). Tissues of Utah juniper are not 

high enough in cedarwood oil to be commercially 
406 

valuable. Heartwood contains 1.19 percent oil, 

principally thujopsene (40 percent) and cedrol 

(13.2 percent) (Adams 1987). The species was 

used by Native Americans to treat bladder and 

kidney troubles, wounds, and mouth sores (Plants 

for a Future 2003). Utah juniper has been used to a 

limited extent in revegetation projects and 

xeriscape gardening. 

References

Abrams, L. 1940. Illustrated flora of the Pacific 

States. Vol. 1. Stanford University Press, 

Stanford, CA. 538 p. 

Adams, R.P. 1987. Investigation of Juniperus

species of the United States for new sources of 

cedarwood oil. Economic Botany 41(1): 48-54. 

American Forests. 1998. National register of big 

trees 1998-1999. American Forests 104(1): 24-

46. 

Bonner, F.T. 2003. Juniperus L., juniper. In: F.T. 

Bonner and R.G. Nisley, eds. Woody Plant Seed 

Manual. U.S. Department of Agriculture, Forest 

Service, Washington, DC. 15 p. 

Grier, C.C., K.J. Elliot, and D.G. McCullough. 

1992. Biomass distribution and productivity of 

Pinus edulis-Juniperus monosperma woodlands 

of north-central Arizona. Forest Ecology and 

Management 5093-4): 331-350. 

Natural Resources Conservation Service. 2003. 

Plants profile: Juniperus osteosperma (Torr.) 

Little, Utah juniper. http://plants.usda.gov/ 

cgi_bin/plant_search.cgi?mode=Scientific+Nave

&keywordquery=Juniperus+osteosperma&earl=

plant_sea. 5 p. 

Peattie, D.C. 1953. A natural history of Western 

trees. Houghton Mifflin Company, Boston, MS. 

749 p. 

Plants For a Future. 2003. Plants for a future: 

database search results: Juniperus osteosperma. 

Plants for a Future, Cornwall, UK. http://www. 

ibiblio.org/pfaf/cgi-bin/arr_html?Juniperus+ 

osteosperma&CAN=LATIND. 8 p. 

Plummer, A.P., D.R. Christensen, and S.B. 

Monson. 1968. Restoring big-game range in 

Utah. Publication 68-3. Utah Division of Fish 

and Game, Salt Lake City, UT. 183 p. 

Secretaría de Medio Ambiente y Recursos 

Naturales. 2003. Especies con usos no 

maderables en bosques de Encino, pino y pinoencino: Juniperus osteosperma (Torr.) Little. 

http://www.semarnat.gob.mx/pfnm/Juniperus 

Osteosperma.html. 2 p. 

Stevens, R. 1999. Mechanical chaining and 

seeding. In: S.B. Monson and R. Stevens, 

comps. Proceedings: ecology and management 

of pinyon-juniper communities within the 

Interior West; Sep. 15-18, 1997. Provo, UT. 

PMRS-P-9. U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research 

Station, Ogden, UT. p. 281-284. 

Stuart, J.D., and J.O. Sawyer. 2001. Trees and 

shrubs of California. University of California 

Press, Berkeley, CA. 467 p. 

Tausch, R.J. 1999. Historic pinyon and juniper 

woodland development. In: S.B. Monson and R. 

Stevens, comps. Ecology and management of 

pinyon-juniper communities within the Interior 

West. Proc. RMRS-P-9. U.S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Research Station, Ogden, UT. p. 12-19. 

Terry, R.G., R.S. Nowark, and R.J. Tausch. 2000. 

Genetic variation in chloroplast and nuclear 

ribosomal DNA in Utah juniper (Juniperus 

osteosperma, Cupressaceae): evidence of 

interspecific gene flow. American Journal of 

Botany 87: 250-258. 

Tree Guide. 2003. Treeguide species details: Utah 

juniper. http://www.treeguide.com/Species. 

asp?SpeciesID=544. 3 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1987. A Utah flora. Brigham 

Young University, Provo, UT. 894 p. 

Zlatnik, E. 1999. Juniperus osteosperma. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/tree/ 

junost/all.html. 33 p. 
407

Kalmia latifolia L. mountain laurel 

ERICACEAE 

Synonyms: Kalmia latifolia var. laevipes Fern. 

W. Henry McNab and Barton D. Clinton 

 

General Description.―Mountain laurel, known 

also as calico-bush, ivy, ivybush, kalmia, laurel, 

and spoonwood, is a medium to large (2 to 9 m in 

height) evergreen shrub, often with contorted stem 

and branching habit. The thick, flat, leathery 

leaves are shiny, dark-green above and pale-green 

below; margins are entire. Leaf length ranges from 

5 to 12 cm and width from 1.5 to 5 cm. The 

yellowish green petioles range from 0.7 to 3 cm 

long. The thin, reddish-brown bark shreds and 

subdivides into ridges. In the absence of 

disturbance (usually fire), the species is long-lived: 

40 to 60 years (Chapman 1950). Botanical 

characteristics of the genus Kalmia have been 

studied intensively (Ebinger 1974). Kalmia is the 

state flower of Connecticut and Pennsylvania. 

Range.―Mountain laurel is primarily a shrub of 

the Appalachian Mountain region in the eastern 

U.S. It also occurs in the Atlantic Coastal Plain 

from Maine to Virginia, throughout the 

Appalachian Piedmont, southwest into Mississippi, 

and west into much of Tennessee, Kentucky, and 

southern Indiana and Ohio. Reports of native 

mountain laurel in Canada have not been 

confirmed and are likely ornamentals or largeleaved specimens of sheep laurel [Kalmia 

angustifolia (Small) Fernald] (Jaynes 1997). 

Kalmia has been planted widely as an ornamental 

in the western United States, throughout Europe, 

and in New Zealand and Australia. 

Ecology.―Mountain laurel occurs most 

commonly on dry sites of acid soils, although it 

occasionally occurs on well-drained floodplain 

soils, and it will tolerate clay soils that are not 

waterlogged. Mountain laurel forms dense, almost 

impenetrable patches known as "laurel hells" or 

"ivy thickets," particularly on upper slopes and 

ridges where the tree canopy is sparse or lacking. 

In the southern Appalachians, mountain laurel 

occurs at elevations to 1500 m and the probability 

of its occurrence increases on sites with a thin soil 

A horizon (McNab et al. 1999), and likely low 

available water capacity. Lipscomb and Nilsen 

(1990) found that mountain laurel has high wateruse efficiency, which could give it a strong 

competitive advantage on relatively hot and dry 

southwest slopes. Mountain laurel adapts to a 

range of light regimes, from little to moderately 

dense shade, typically beneath an oak (Quercus

spp.) or pine (Pinus spp.) canopy. The ratio of 

foliage chlorophylls a to b is lower (1.5) in Kalmia

than for most understory species and may benefit 

this adaptation (Al-Hamdani and others 2002). 

Rosebay rhododendron (Rhododendron maximum

L.) is a common associate of mountain laurel on 

subxeric/submesic ecotones in the southern 

Appalachians. A mycorrhizal association has been 

demonstrated for seedlings of mountain laurel 

(Flemer 1949). Natural stands of mountain laurel 

are little affected by insects, although at epidemic 

levels caterpillars of gypsy moths (Lymantria 

dispar L.) and elm spanworms (Ennomos 

subsignaria Hubner) will feed on the foliage. The 

most serious disease is phytophthora root rot 

(Phytophthora cinnamomi Rands), which can 

infect damaged plants (Jaynes 1997). Foliage of 

mountain laurel near roadsides is particularly 

sensitive to sodium chloride (NaCl) used for deicing roads in New England (Bryson and Baker 

2002). Although not shown in mountain laurel, 

other Kalmia species are allelopathic. Waterman 

and others (1995) found that Kalmia thickets do 

not exclude tree seedlings but inhibit their growth 
408 

by reducing light and increasing competition for 

water. 

Reproduction.―The showy, white to pink 2- to 

3- cm saucer-shaped flowers form convex clusters 

at the ends of branches and appear after the 

foliage, from March to July, depending on latitude 

and altitude (Olson and Barnes 1974). Purple spots 

mark 10 anther pockets in the petals. The method 

of pollen dispersal in Kalmia is unusual in that the 

anthers are under tension as the flower matures, 

and they spring forward when disturbed by an 

insect, forcefully and effectively transferring 

grains of pollen to the insect (Jaynes 1997). The 

flowers are pollinated by more than a dozen 

insects, primarily bumblebees, but generally not 

honeybees, likely because the flowers produce 

little nectar (Jaynes 1997). Glandular hairs on the 

flower stalk, calyx, and corolla may deter 

nonflying insects. Flowers are most abundant on 

plants in full sunlight and occur rarely on plants in 

dense shade (Kurmes 1961). Fruit ripens in late 

summer and consists of a brown, erect, five-celled, 

globular capsule, 4 to 7 mm wide. Each capsule 

contains about 600 seeds, which are 0.8 to 1 mm 

long and average about 27 million per kilogram 

(Olson and Barnes 1974). The nonwinged seeds 

are dispersed by wind, but only for distances of 

less than 15 m (Robinette 1974). Seeds remain 

viable for more than a year under field conditions. 

Light, cold stratification, and moisture enhance 

germination (Olson and Barnes 1974). Seeds 

germinate on mineral soil and on moss (Robinette 

1974). Kalmia may also regenerate by layering, 

sprouts, or suckers (Robinette 1974). Techniques 

for commercial production of mountain laurel 

plants have been well developed (Jaynes 1997). 

Growth and Management.―Growth rate of 

mountain laurel is relatively slow; young plants (< 

15 years) add about 12 cm in height and 9 cm in 

width annually, depending on site moisture 

relations (Monk and others 1985). Older mountain 

laurel stems may attain heights up to 9 m and 

diameter at ground level of 15 cm (Jaynes 1997). 

Mountain laurel stem density can range from 

sparse on mesic sites to over 26,000 per ha in 

thickets on upper, southwest slopes; basal area at 3 

cm above ground level can exceed 25 m2

/ha. 

Herbicides may be used to control mountain laurel 

on sites where establishment of tree seedlings is 

desired (Neary and others 1984). It is sensitive to 

juglone, a substance produced by black walnut 

(Juglans nigra L.) trees. Mountain laurel generally 

does not present a problem to timber management 

activities on submesic to mesic sites. 

Benefits and Detriments.―Mountain laurel has 

long been cultivated as an ornamental--since about 

1740 when American naturalist John Bartram sent 

plants to England (Jaynes 1997). Cultivars of 

mountain laurel are particularly valuable as 

ornamentals and are propagated commercially by 

many nurseries. Nearly 100 horticultural varieties 

of have been selected for variation in flower color, 

leaf variegation, and growth habit (Jaynes 1997). 

Mountain laurel burls have been used for making 

pipes for smoking tobacco, and early European 

settlers in America used the fine-grained wood for 

making eating utensils and weaver's shuttles, and 

produced a yellow dye from its foliage. Tinctures 

of fresh Kalmia leaves have been used for home 

medical remedies of skin disorders. An 

unidentified anticancer compound has been 

extracted from its sap (Jaynes 1997). Mountain 

laurel provides cover and stability to thin soils, 

however its presence can reduce water yields 

(Johnson and Kovner 1956). Foliage of mountain 

laurel is toxic to domestic cattle and sheep but 

provides subsistence winter food for ruffed grouse 

(Bonasa umbellus L.) and white-tailed deer 

(Odocoileus virginianus Boddaert) (Robinette 

1974). Foliage from new sprouts of burned 

mountain laurel provides greater potential benefit 

to wildlife diets compared to unburned foliage 

(Thackston and others 1982). 

 

Fire.―Mountain laurel contributes to intensity of 

fires in mountainous terrain and is classified as 

"extreme" in rate of spread when it occurs in dense 

thickets beneath hardwood stands (Jemison and 

Keetch 1942). Flame length can exceed 30 m in 

fires burning in Table Mountain pine (Pinus 

pungens Lambert) stands with mountain laurel 

(Waldrop and Brose 1999). Moisture of Kalmia

foliage is lower than other species and is little 

affected by seasonal changes of soil moisture 

(Reifsnyder 1961). The high flammability of 

Mountain laurel thickets results from low foliate 

moisture (Richards 1940) and a high proportion of 

total biomass in small (<5 mm) dead twigs. 

Caloric content of foliage is estimated at about 

5,000 cal/g based on similar vegetation (Hough 

1969). Allometric equations for estimating dry 

biomass of 6- to 8-year old open-grown mountain 

laurel stems were developed by Boring and Swank 

(1986), which can be used to predict fuel loadings 

of leaf (log10Y = -0.754 + 1.882 log10X), branch 

(log10Y = -1.222 + 2.359 log10X), and bole (log10Y 

= -0.636 + 1.948 log10X) components of individual 

plants, where Y is dry weight (g) and X is stem 

diameter (mm) at 3 cm from the ground. Kalmia is 

killed more readily by headfires than by backfires 
409

(Hooper 1969), but almost all topkilled stems 

recover by producing basal sprouts (Clinton and 

others 1993, Johnson and Kovner 1956). Equations 

for estimating stem, foliage, and total biomass of 

young mountain laurel sprouts on recently burned 

areas were developed by Elliott and Clinton 

(1993). 

References 

Al-Hamdani, S.H., P.B. Nichols, and G.R. Cline. 

2002. Seasonal changes in the spectral 

properties of mountain laurel (Kalmia latifolia

L., Ericaceae) in north east Alabama. Castanea. 

67(1): 25-32. 

Boring, L.R. and W.T. Swank. 1986. Hardwood 

biomass and net primary production following 

clearcutting in the Coweeta basin. In: Brooks, 

R.T., Jr. ed. Proceedings of the 1986 southern 

forest biomass workshop. Tennessee Valley 

Authority, Knoxville, TN: 43-50. 

Bryson, G.M. and A.V. Barker. 2002. Sodium 

accumulation in soils and plants along 

Massachusetts roadsides. Communications in 

Soil Science and Plant Analysis. 33:67-78. 

Chapman, G.L. 1950. The influence of mountain 

laurel undergrowth on environmental conditions 

and oak reproduction. Doctoral dissertation. 

Yale University, New Haven, CT. 

Clinton, B.D., J.M Vose, and W.T. Swank. 1993. 

Site preparation burning to improve southern 

Appalachian pine-hardwood stands: vegetation 

composition and diversity of 13-year-old stands. 

Canadian Journal Forest Research. 23: 2,271-

2,277. 

Ebinger, J.E. 1974. A systematic study of the 

genus Kalmia (Ericaceae). Rhodora. 76: 315-

398. 

Elliott, K.J. and B.D. Clinton. 1993. Equations for 

estimating biomass of herbaceous and woody 

vegetation in early-succession Southern 

Appalachian pine-hardwood forests. Research 

Note SE-365. U.S. Department of Agriculture, 

Forest Service, Southeastern Forest Experiment 

Station, Asheville, NC. 7 p. 

Flemer,, W., III 1949. The propagation of Kalmia 

latifolia from seed. Bulletin of the Torrey 

Botanical Club. 76: 12-16. 

Hooper, R.M. 1969. Prescribed burning for laurel 

and rhododentrol control in the southern 

Appalachians. Research Note SE-116. U.S. 

Department of Agriculture, Forest Service, 

Southeastern Forest Experiment Station, 

Asheville, NC. 6 p. 

Hough, W.A. 1969. Caloric value of some forest 

fuels of the southern United States. Research 

Note SE-120. U.S. Department of Agriculture, 

Forest Service, Southeastern Forest Experiment 

Station, Asheville, NC. 6 p. 

Jaynes, R.A. 1997. Kalmia--Mountain laurel and 

related species. Timber Press, Portland, OR. 

295 p. 

Jemison, G.M. and J.J. Keetch. 1942. Rate of 

spread of fire and its resistance to control in the 

fuel types of eastern mountain forests. Technical 

Note 2. U.S. Department of Agriculture, Forest 

Service, Appalachian Forest Experiment Station, 

Asheville, NC. 15 p. 

Johnson, E.A. and J.L. Kovner. 1956. Effect on 

streamflow of cutting a forest understory. Forest 

Science. 2: 82-91. 

Kurmes, E.A. 1961. The ecology of mountain 

laurel in southern New England. Yale 

University, New Haven, CT. 85 p. Doctoral 

Dissertation. 

Lipscomb, M.V. and Nilsen, E.T. 1990. 

Environmental and physiological factors 

influencing the natural distribution of evergreen 

and deciduous ericaceous shrubs on northeast 

and southwest slopes of the southern 

Appalachian Mountains. II. Water relations. 

American Journal Botany. 77(4): 517-526. 

McNab, W.H., S.A. Browning, S.A. Simon, P.E. 

Fouts. 1999. An unconventional approach to 

ecosystem unit classification in western North 

Carolina, USA. Forest Ecology and 

Management. 114: 405-420. 

Monk, C.D., D.T. McGinty, and F.P. Day, Jr. 

1985. The ecological importance of Kalmia 

latifolia and Rhododendron maximum in the 

deciduous forest of the Southern Appalachians. 

Bulletin Torrey Botanical Club. 112(2): 187-

193. 

Neary, D.G., J.E. Douglass, J.L. Ruehle, and W. 

Fox. 1984. Converting rhododendron-laurel 
410 

thickets to white pine with picloram and 

mycorrhizae-inoculated seedlings. Southern 

Journal of Applied Forestry. 8(3): 163-168. 

Olson, D.F., Jr. and R.L. Barnes. 1974. Kalmia 

latifolia L., mountain laurel. In: C.S 

Schopmeyer, ed. Seeds of woody plants in the 

United States. Agriculture Handbook 450. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. p. 470-471. 

Robinette, S.L. 1974. Mountain laurel-Kalmia 

latifolia. In: J.D. Gill and W.M. Healy, eds. 

Shrubs and vines for northeastern wildlife. 

General Technical Report NE-9. U.S. 

Department Agriculture, Forest Service, 

Northeastern Forest Experiment Station, Radnor, 

PA. p. 102-105. 

Reifsnyder, W.E. 1961. Seasonal variation in the 

moisture content of the green leaves of mountain 

laurel. Forest Science. 7: 16-23. 

Thackston, R.E., P.E. Hale, A.S. Johnson, and 

M.J. Harris. 1982. Chemical composition of 

mountain-laurel leaves from burned and 

unburned sites. Journal Wildlife Management. 

46(2): 492-496. 

Waldrop, T.A. and P.H. Brose. 1999. A 

comparison of fire intensity levels for stand 

replacement of table mountain pine (Pinus 

pungens Lamb.). Forest Ecology and 

Management. 113: 155-166. 

Waterman, J.R., A.R. Gillespie, J.M. Vose, and 

W.T. Swank. 1995. The influence of mountain 

laurel on regeneration in pitch pine canopy gaps 

of the Coweeta Basin, North Carolina, U.S.A. 

Canadian Journal Forest Research. 25: 1,756-

1,762. 
411

Koanophyllon polyodon (Urban) King & H.E. Robins. polyodon 

ASTERACEAE 

Synonyms: Eupatorium polyodon Urban 

John K. Francis 

General Description.—The name polyodon is 

assigned according to the convention that if no 

common name exists, the species (or sometimes 

the genus) Latin name is used as a common name. 

Polyodon is a somewhat spindly, upright or arching 

shrub usually 1.5 to 3 m (occasionally to 6 m) in 

height and 2 to 3 cm (sometimes 5 cm) in basal 

diameter. The shrub may have a single or multiple 

stems, straight lower sections 0.5 to 1 m long, then 

curving and branchy with fine twigs above. The 

plant is supported by an extensive lateral and fine 

root system. The opposite leaves are papery and 

rough with short petioles and three main veins 

arising from or near the base. The leaf blades are 

ovate with a heart-shaped or nearly heart-shaped 

base, pointed tip, and serrate edges. The 

inflorescences are terminal corymbs of white or 

rose compound flowers (heads). Each develops 

several, five-ribbed, 3-mm achenes tipped with 

numerous, 3 to 4 mm hairs (pappus) (author’s 

observation, Liogier 1997). 

Range.—Polyodon is endemic to Puerto Rico 

(Liogier 1997) and is not known to have been 

planted or naturalized elsewhere. 

Ecology.—Polyodon grows on moderately and 

well-drained but rarely excessively drained soils of 

loamy and clayey textures with pH’s ranging from 

about 5 to 7. Both sedimentary (including 

limestone), igneous, and metamorphic (including 

ultramafic) parent materials are colonized. The 

species is found in areas that receive from about 

1000 to 2500 mm of mean annual precipitation at 

elevations ranging from near sea level to 900 m. 

Polyodon grows as scattered individuals, diffuse 

stands (under forest canopies), or in thickets. The 

species has an intermediate tolerance for shade and 

is able to grow and reproduce under shade if not 

too dense. It depends on taller vegetation for 

support in order to reach heights of more than 

about 1.5 m. The species is occasional to common 

in open forest, brushlands, abandoned pastures, and 

old road cuts and banks. 

Reproduction.—Polyodon flowers near the end of 

the wet season (November-January) and fruits 

during the dry season (March-May). It is insect 

pollinated. A collection of seeds averaged 0.00019 

g/seed or 5.2 million seeds/kg. Placed on moist 

filter paper without any pretreatment, 67 percent 

germinated between 12 and 68 days after sowing. 

Germination is epigeal. The fruits are dispersed by 

the wind. Seeds are produced in abundance, but 

seedlings are scattered. Plants will resprout when 

damaged. 

Growth and Management.—Individual stems 

may live for 6 years or more. Plants can survive 

longer by producing sprouts. Sprouts grow at least 

1 m in their first year. No management experience 

has been published. However, maintaining areas in 

early and middle secondary forest would probably 

encourage populations of polyodon. 

Benefits.—Polyodon contributes to biodiversity in 

its wildland habitat, helps protect the soil from 

erosion, and furnishes wildlife cover. It is browsed 

to some extent by cattle and goats. 

References 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 
412 

Krameria ixine L. abrojo colorado 

EUPHORBIACEAE 

Synonyms: Krameria cuspidata C. Presl. 

John K. Francis 

General Description.—Abrojo colorado, 

sometimes known as retama falsa, is a shrub or 

sufruticose shrub to 1 m in height and 1 cm in stem 

basal diameter. The stems are stiff and woody, the 

twigs flexible and less woody. Older plants have 

lateral roots with a small or no taproot and many 

fine roots. The roots are brittle, dark brown outside, 

with red inner bark. There are multiple stems from 

the root crown. The branches and leaves are 

covered with white hairs, giving the whole plant a 

gray-green color. The leaves are narrowly oblong 

to oblong-lanciolate, 1 to 2.5 cm long, with a 

spiny-acuminate tip. The inflorescences are 

compact lateral or terminal racemes bearing several 

flowers. The pink- to purplish-red flowers are 

about 1.2 cm in diameter, from which develop 

globose 5 to 6 mm, woody capsules (burrs) covered 

with orange or red, hooked spines. The seeds are 

light tan and 4 mm in diameter (Howard 1988, 

Liogier 1988, Stevens and others 2001). 

Range.—Abrojo colorado is native to Hispaniola, 

Puerto Rico, the Lesser Antilles, Mexico through 

Costa Rica, Colombia, Venezuela, and Guyana 

(Liogier 1988, Missouri Botanical Garden 2002, 

New York Botanical Garden 2002, Stevens and 

others 2001). It is not known to have naturalized 

outside its native range. 

Ecology.—In Puerto Rico, abrojo colorado grows 

in areas that receive from 750 to about 1000 mm of 

mean annual precipitation and at altitudes from 

near sea level to about 400 m above sea level. 

Because the shrub is intolerant of shade and low in 

stature, it does not withstand serious competition. It 

will grow in almost any well-drained, nonsaline 

soil. It is most frequently seen on rocky ridges and 

south slopes, disturbed sites, and overgrazed range. 

It is rarely browsed by cattle or horses. 

Reproduction.—Abrojo colorado flowers and 

fruits throughout the year in Costa Rica where both 

flowers and fruit may be present (Instituto 

Nacional de Biodiversidad 2002). Both flowers and 

fruits are often present in Puerto Rican plants 

(author’s observation). Fruit and seed production is 

moderate to abundant. A collection of air-dried 

fruits in Puerto Rico weighed an average of 0.0516 

± 0.0009 g. The seeds are transported as they cling 

to passing animals or humans. After drying 

sufficiently, the capsules open, releasing the seeds. 

After disturbance, the plants regenerate by 

sprouting. 

Growth and Management.—Individual stems of 

abrojo colorada grow about 0.5 m/year in the first 

year from sprouts and live 3 or 4 years. By 

continual sprouting, established plants live much 

longer. Although no management experience has 

been published, it probably is safe to say that 

overgrazing and disturbance encourages the 

species, and that reestablishment of tree, tall shrub, 

or heavy grass cover discriminates against or 

eliminates it. 

Benefits.—Abrojo colorado contributes ground 

cover, soil stability, and scenic beauty in rocky and 

often disturbed terrain where it grows. On the other 

hand, the burrs can be a nuisance to both humans 

and animals. 
413

References 

Howard, R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Instituto Nacional ce Biodiversidad. 2002. Lista de 

especímenes de Krameria ixine. 

http://www.inbio.ac.cr/bims/k03/p13/c045/o013

2/f01646/g008002/s024181.htm. 2 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Missouri Botanical Garden. 2002. Manual de la 

Flora de Costa Rica. http://www.mobot.org/ 

manual.plantas/019706/S019706.html. 1 p. 

New York Botanical Garden. 20023. Specimens 

search results: Krameria ixine Loefl. 

http://scis.../wwwspecimen.search_list?taxon= 

Krameria+ixine+Loefl.+++++&projcode=VAS. 

1 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden, St. Louis, MO. 

p. 945-1,910. 
414 

Lantana camara L. lantana

VERBENACEAE 

Synonyms: Lantana aculeata L. 

Lantana antillana Raf.

 Lantana camara L.var.aculeata (L.) Mold. 

 Lantana scabrida Soland. 

John K. Francis 

General Description.—Lantana has many 

common names including cariaquillo, filigrana, mille fleurs, sauge, red sage, yellow sage, prickly 

sage, and lakana. It is a medium-sized aromatic 

shrub with quadrangular stems, sometimes having 

prickles. The posture may be suberect, scrambling, 

or occasionally clambering (ascending into shrubs 

or low trees, clinging to points of contact by means 

of prickles, branches, and leaves). Frequently, 

multiple stems arise from ground level. The leaves 

are generally oval or broadly lance-shaped, 2 to 12 

cm in length, and 2 to 6 cm broad, having a rough 

surface and a yellow-green to green color. The 

flat-topped inflorescence may be yellow, orange, 

white, pale violet, pink, or red (Howard 1989, 

Liogier 1995). The small individual flowers of 

wild Puerto Rican shrubs darken with age so that 

inflorescences are yellow in the center and orange 

toward the perimeter. Fruits form in clusters 

similar in appearance to a blackberry. 

Range.—The native range of lantana extends from 

Bermuda, the Bahamas, the Greater Antilles, the 

Lesser Antilles, through Trinidad and Aruba. On 

the mainland, it is native to coastal areas of the 

United States from Georgia through Texas and 

from northern Mexico to South America including 

Brazil and Peru and probably Bolivia, Paraguay 

and northern Argentina. Lantana has naturalized in 

most suitable habitats in tropical and subtropical 

Africa, Asia, and Australia. Most of the world’s 

tropical and subtropical islands including the 

Hawaiian Islands, Guam, Pitcairn Island, 

Madagascar, the Juan Fernandez archipelago, 

Reunion, and the Galapagos have naturalized 

populations. This colonization has occurred largely 

during the last century. 

Ecology.—Lantana grows on all types of welldrained soil in areas that receive from about 250 

mm to 2900 mm of rainfall. It resists droughts very 

well and tolerates salt spray. Aerial portions of the 

plant are killed by temperatures of –2 °C, but 

quickly grow back (Anonymous 2000). Large and 

vigorous plants survive fires and cutting well, 

although less vigorous plants are often killed. 

Lantana is an intolerant pioneer that colonizes 

disturbed areas. It will grow under an open forest 

canopy but quickly disappears when the shade 

becomes heavy. Many pests and diseases lightly 

and incidentally affect the species across its broad 

range. 

Reproduction.—The inflorescence is a capitate, 

many-flowered head. The corolla may vary widely 

in color depending on the variety but 

characteristically changes colors between the 

center flowers and older, outer flowers. Lantana 

blooms almost continuously under favorable 

conditions. Somatic chromosome numbers of 33, 

44, and 55 were recorded in India, the latter 

tetraploid being the most common (Sinha and 

others 1995). Insects, especially butterflies, 

pollinate the flowers. Clusters of drupes are 

produced abundantly. The fruits are blueblack 
415

when ripe and contain one seed each. They are 

eaten by birds and are widely scattered. If not 

eaten, they dry and remain on the shrub for weeks.

A sample of seed collected in Puerto Rico 

contained 100,700 seeds/kg of which 75 percent 

germinated within 7 weeks of sowing. Early 

growth is rapid. Also, lantana can be propagated 

with cuttings and air layers. 

Growth and Management.—Ornamental plants 

are established as potted seedlings. This is 

probably also the best method to establish plants 

for environmental protection projects such as 

reclamation of mine spoils. Lantana may reach 3 

m in height within 3 or 4 years. In naturalized 

habitat, the species often forms dense thickets. 

These brush stands may be eliminated at 

considerable expense from pastures and tree 

plantations by cutting and herbicide application. 

The preferred methods are to cut the clumps and 

drench the stumps with herbicide or to later spray 

the tender regrowth with postemergence herbicide. 

Once established, tree plantations with closed 

canopies will keep lantana from reestablishing 

itself. 

Benefits.—Lantana, in many horticultural 

varieties, is planted the world over as a flowering 

ornamental. It is grown as an annual bedding plant 

in temperate areas. Lantana oil, an aromatic 

mixture that varies by local plant variety, is 

exported from at least Brazil (Weyerstahl and 

others 1999). In herbal medicine, infusions of the 

leaves and other plant parts are used as an 

antiinflammatory (Oyedapo and others 1999), a 

tonic and expectorant, and added to baths as an 

antirhumatic. Lantana extracts have also been 

shown to be a powerful febrifuge (Liogier 1990). 

Because the leaves and some other parts of lantana 

are poisonous, care must be taken when it is used 

medicinally. The ripe fruit is benign and heavily 

consumed by birds and frequently eaten by 

humans in some countries (Herzog and others 

1994). Extracts of lantana leaves have shown 

strong insecticidal and antimicrobial activity in 

numerous experiments. Storing potatoes with 

lantana leaves nearly eliminates damage by 

Phthorimaea operculella Zeller, the potato tuber 

moth (Lal 1987). Stems and leaves are used as 

mulch. Although of inferior quality because of size 

and form, lantana stems are widely used as fuel in 

less developed countries. 

Detrimental Effects.—Lantana has become a 

weedy invader of disturbed forest land and 

neglected pasture in much of its naturalized range. 

In some areas, competition by lantana results in a 

reduction of biodiversity (Kumar and Rohatgi 

1999). Despite the establishment of a number of 

natural enemies of lantana into exotic populations, 

control of lantana populations has been usually 

limited or a failure (Day and others 1999, Hill 

1999). In thick stands, the shrub increases costs in 

forest management by inhibiting access in stands 

for thinning and felling, competes with 

reproduction, and increases fire hazards (Graaff 

1986). Lantana leaves contain poisonous 

triterpines and lantadenes A and B that cause death 

of horses, cattle, sheep, goats, and rabbits by 

failure of the liver and other organs (Morton 1994, 

Munyua and others 1990). However, most animals 

carefully avoid eating this plant when given a 

choice. Green fruits also contain the poisons and 

have caused illness and death in children (Morton 

1994). Lantana leaves and their leachates exert 

alelopathic effects in vetro and to a lesser extent in 

soil on seed germination, root elongation, and 

plant growth of many species (Casado 1995, Sahid 

and Sugau 1993). 

References

Anonymous. 2000. Floridata, Lantana camara. 

www.streetside.com/plants/floridata/ref/l/lant_c.

ht. 1p. 

Casado, C.M. 1995. Allelopathic effects of 

Lantana camara (Vergenaceae) on morning 

glory (Ipomoea tricolor). Rhodora 97: 264-274. 

Day, M.D., R.H. Holtkamp, and P. Blackmore. 

1999. The status of biological control of 

Lantana camara in Australia. In: Practical weed 

management: protecting agriculture and the 

environment. 10th Biennial Noxious Weeds 

Conference, 10-22 July 1999, Ballina, Australia. 

p. 257-260. New South Wales Agriculture, 

Armidale, Australia. 

Graaff, J.L. 1986. Lantana camara, the plant and 

some methods for its control. South African 

Forestry Journal 136: 26-30. 

Hill, M.P. 1999. Past and present initiatives on 

the biological control of Lantana camara 

(Verbenaceae) in South Africa. In: J. R. Baars, 

S. Neser, and T. Olckers, eds. Biological control 

of weed in South Africa 1990-1998. African 

Entomology Memoir 1. 21-33. 
416 

Herzog, F., Z. Farah, and R. Amado. 1994. 

Composition and consumption of gathered wild 

fruits in the V-Baoule, Cote d’Ivoire. Ecology of 

Food and Nutrition 32(3-4): 181-196. 

Howard, R.A. 1989. Flora of the Lesser Antilles. 

Vol. 6. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 658 p. 

Kumar, S. and N. Rohatgi. 1999. The role of 

invasive weeds in changing floristic diversity. 

Annals of Forestry 7:(1): 147-150. 

Lal, L. 1987. Studies on natural repellents against 

potato tuber moth (Phthorimaea operculella

Zeller) in country stores. Potato Research 30(2): 

329-334. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 563 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p.

Morton, J.F. 1994. Lantana, or red sage (Lantana 

camara L. Vergenaceae), notorious weed and 

popular garden flower; some cases of poisoning 

in Florida. Economic Botany 48(3): 259-270. 

Munyua, S.J.M., M.J. Nienga, T.P. Karitu, T.P. 

Kimoro, J.E. Kiptoon, and I.B.J. Buoro. 1990. A 

note on clinical-pathological findings and serum 

enzyme activity in sheep, goats and Friesian 

calves with acute Lantana camara poisoning. 

Bulletin of Animal Health and Production in 

Africa 38(3): 275-279. 

Oyedapo, O.O., F. C. Sab, and J.A. Olagunju. 

1999. Bioactivity of fresh leaves of Lantana

camara. Biomedical Letters 59: 179-183. 

Sahid, I.B. and J. B. Sugau. 1993. Allelopathic 

effects of lantana (Lantana camara) and siam 

weed (Chromolaena odorata) on selected crops. 

Weed Science 41(2): 303-308. 

Sinha, S., B. Sinha, and A. Sharma. 1995. 

Chromosome composition of Lantana camara

L.: karyotype, basic number and DNA diversity. 

Nucleus Calcutta 38(1-2): 16-22. 

Weyerstahl, P., H. Marschall, A. Eckhardt, and C. 

Christiansen. 1999. Constituents of commercial 

Brazilian lantana oil. Flavour and Fragranced 

Journal 14(1): 15-28. 
417

Lantana involucrata L. wild sage 

VERBENACEAE 

Synonyms: Lantana odorata L. 

John K. Francis 

 

General Description.—Wild sage, also known as 

button sage, common sage, sea sage, white sage, 

orégano, terete, peonía colorada, contite, and 

frutilla, is a shrub commonly 1 to 2 m in height 

with showy flowers. The plant is supported by 

many stiff lateral roots and abundant fine roots. 

The roots are light brown with a corky bark. Older 

plants, often with a basal diameter of 5 cm or more, 

have several main branches arising at or near the 

ground line and a branchy crown. The bark is 

yellowish and the wood is brittle. Petioles are 3 to 

5 mm long and the leaf blades are 1 to 4 by 1 to 2.5 

cm wide and broad, rotund to narrowly ellipticoblong, rough and stiff with a crenate or toothed 

margin. The leaves are aromatic with a spicy, sagelike smell. The small tubular flowers form compact 

terminal heads. The corolla may be white, pink to 

pinkish purple, or pale blue. The infrutescences are 

clusters of purple to blue-black drupes 3 to 4 mm 

in diameter containing one seed each. The 

chromosome number is 2n = 36 (author’s 

observation, Howard 1989, Liogier 1995, Long and 

Lakela 1971, Nelson 1996). 

Range.—The native range of wild sage includes 

southern Florida, the West Indian islands, Mexico 

through northern South America bordering the 

Caribbean, and the Galápagos Islands (Howard 

1989, Liogier 1995). It is naturalized or possibly 

native in Bermuda (Britton 1918). Wild sage is 

now widely planted as an ornamental and probably 

has naturalized in additional areas. 

Ecology.—Wild sage grows on most well-drained 

soil types derived from both sedimentary 

(including limestone), igneous, metamorphic 

(including ultramafic) rock types. Mean annual 

precipitation in areas where it grows in Puerto Rico 

varies from 750 to about 1700 mm. The species 

grows from near sea level to 600 m or more in 

Puerto Rico. In Mexico, wild sage is reported to 

occur from 1,000 to 2,000 m above sea level 

(Secritaría del Medio Ambiente y Recursos 

Naturales 2002). It requires disturbance to become 

established and full or nearly full sun to grow and 

reproduce. Wild sage is especially common in 

over-grazed range and poorly managed pastures, 

and may be found in abandoned farmland and early 

secondary forest as well as road cuts, cliffs, and 

rocky sites. It grows in pinelands, hammocks, and 

shell mounds in Florida (Long and Lakela 1971, 

Nelson 1996). 

Reproduction.—Wild sage blooms all year 

(Nelson 1996) and also is a good fruit and seed 

producer. The seeds number about 110,000 

seeds/kg (air-dried). They may be cleaned by 

maceration and wet sieving. No pretreatment is 

necessary, and germination begins in about 15 

days. Wild sage sprouts when cut. Seedlings are 

fairly common in favorable habitat near seed 

sources. 
418 

Growth and Management.—The growth rate is 

moderate and plants live for 10 years or more. 

Propagation may be by seed, by rooting of 

softwood cuttings, or by transplanting wildlings. 

Wildlings transplant well if the roots are not badly 

damaged and about one-third of the top is pruned 

away. Periodic pruning of ornamental plants is 

necessary to prevent them from becoming lank 

(Workman 1980). 

Benefits.—Wild sage contributes to biodiversity, 

helps protect the soil, and furnishes wildlife food 

and cover. The wood is useful for fuel, but it is 

generally too small for anything but campfires. The 

species is used as a foundation plant and to form 

low hedges (Workman 1980). It is listed as one of 

the best butterfly nectar plants (Malone 2002). The 

leaves are used as a condiment in cooking, and the 

essential oil is used in cosmetics and liquors 

(Secritaría del Medio Ambiente y Recursos 

Naturales 2002). Leaves are added to baths to ease 

heat rashes and mild insect bites (McNary-Wood 

2002). Extracts of the plant are also said to have a 

sedative effect (Beattie and others 2002). Teas and 

decoctions of leaves and twigs are used to control 

colic, vomit, cough, fever, and congestion 

(Secritaría del Medio Ambiente y Recursos 

Naturales 2002). It is reportedly not as poisonous 

as the more widespread L. camara L. (Workman 

1980). 

References 

Beattie, L., M. Martin, M. Shaposhnik, and K. 

Vaga. 2002. Stress treatments. http://members. 

tripod.com/~Moiraine/treatment.html. 7 p. 

Britton, N.L. 1918. Flora of Bermuda. Charles 

Schribner’s Sons, New York. 585 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles. 

Vol. 6. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de 

la Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Long, R.W. and O. Lakela. 1971. A flora of 

tropical Florida. University of Miami Press, 

Coral Gables, FL. 962 p. 

Malone, K. 2002. Palm Beach County 

(Southeastern Florida) top butterfly nectar 

flowers. North American Butterfly Association. 

http://www.naba.org/ftp/pbco.pdf. 4 p. 

McNary-Wood, K. 2002. Medicinal plants for 

what ails you in paradise. http://www. 

timespub.tc/Natural%History/Archive/Winter%

20200102/medicinal.htm. 2 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Secritaría del Medio Ambiente y Recursos 

Naturales. 2002. Especies con usos no 

maderables en bosques tropicales y 

subtropicales: Lantana involucrata L. 

http://www.semarnat.gob.mx.pfnm2.fichas/ 

lantana_involucrata.htm. 1 p. 

Workman, R.W. 1980. Growing native. The 

Sanibel-Captiva Conservation Foundation, Inc., 

Sanibel, FL. 137 p. 
419

Larrea tridentata (Sesse' and Moc. ex DC.) Coville creosote bush 

ZYGOPHYLLACEAE 

Synonyms: Covillea tridentata (Sesse' & Moc. ex DC.) Vail 

 Covillea glutinosa (Engelm.) Rydb. 

 Larrea divaricata Cav. 

 Larrea glutinosa Engelm. 

 Larrea mexicana Moric. 

James E. Nellessen 

General Description.—Creosote bush is also 

know by the names gobernadora, hediondilla 

(Spanish for little stinker), and guamis. It is 

sometimes erroneously referred to as greasewood. 

It is an evergreen resinous shrub of warm deserts. 

It is a well branched shrub with no defined trunk, 

typically reaching heights of 1 to 1.5 m, but can 

grow as high as 3 m. It is strongly scented, 

especially noticeable after a rainstorm. The stems 

are gray with black, somewhat swollen bands at 

the nodes; the twigs and young branches are 

relatively flexible. The leaves are opposite, 

bifoliately compound, each leaflet joined at the 

base, divergent, but the tips often point toward 

each other. Leaflets generally are olive green in 

color, to 1 cm in length, 3 to 4 mm wide, 

asymmetrical, oblong to obovate in shape, and 

broader away from the base (Carter 1997, Correll 

and Johnston 1970, Hickman 1993, Kearney and 

others 1951, Martin and Hutchins 1980-81). The 

leaves are often glossy with a thick resinous 

coating. Stipules are persistent. The South 

American L. divaricata is so similar to L. 

tridentata that some researchers have considered 

them the same species (Mabry and others 1977). 

These two species have been demonstrated to 

hybridize in experimental gardens. South America 

is the likely origin for the genus, with creosote 

bush ancestors arriving in North America 8.4 to 

4.2 million years before present (Lia and others 

2001) with current distribution occurring since the 

last glaciation. Creosote bush is believed to be a 

migrant through possible bird dispersal of the L. 

divaricata populations (Mabry and others 1977). 

There is a ploidy gradient from east to west (from 

moister to dryer deserts) across the Southern 

United States: Chihuahuan Desert populations are 

diploid, Sonoran tetraploid, and Mojave hexaploid. 

From a study of plant remains in packrat middens, 

the plants appear to have spread slowly from lower 

Rio Grande refugia in Texas north and west, 

reaching the Southern Mojave about 8,500 years 

ago, the Northern Sonoran about 6,400 years ago, 

and the Northern Chihuahuan about 4,000 years 

ago (Hunter and others 2001). Larrea divaricata

and diploid L. tridentata have homologous 

genomes (Mabry and others 1977). Two varieties 

have been recognized by one author: L. tridentata

var. tridentata and L. tridentata var. arenaria L. 

Benson (Kartesz 1994). 

Range.—Creosote bush occurs from Southern and 

Western Texas through Southern New Mexico 

(with a band up the Rio Grande valley to 

Albuquerque), into Arizona, and Southern 

California, as far north as Southern Nevada and 

Southwestern Utah, and south well into Mexico. 

Its elevational range is from below sea level 

(Death Valley, CA) to about 1,700 m. 

Ecology.—Creosote bush is a characteristic 

dominant evergreen shrub of the Southwestern 

warm Chihuahuan, Sonoran, and Mojave deserts. 

On the northern fringes of the Mojave its range 

just touches the Great Basin Desert (Mabry and 

others 1977). It grows best in gravelly to sandy, 

well drained soils of desert plains, benches, mesas, 

alluvial fans, and rocky slopes. It does not occur 

on alkali flats or pans. It roots deeply, to 3 m or 

more, but most roots occur in the upper soil layers 

(but below about 10 cm), and it will tolerate a 
420 

caliche layer. The shallow roots are able to utilize 

small rainfall events. Creosote roots inhibited 

further elongation of the roots of both adjacent 

creosote bushes and the shrub Ambrosia dumosa

(A. Gray) Payne (Mahall and Callaway 1992). 

Stem angle facilitates water stemflow to the roots, 

which is maximal with stems greater than 65° and 

almost none less than 45°. Creosote bushes in the 

driest part of their range, for example, Death 

Valley, generally have steeper angles, while there 

is more variability in stem angle in the Chihuahuan 

Desert. Shrubs with shallower stem angles take on 

a more hemispherical shape as opposed to an 

inverted cone shape. Shrub canopy architecture 

also affects soil characteristics creating "islands of 

fertility." Shrubs with hemispherical shapes collect 

much more litter underneath, hence soil nitrogen is 

greater under hemispherical shrubs than under 

inverted cone shaped shrubs (Whitford 2002). 

Creosote bushes often require a nurse shrub such 

as the small shrub A. dumosa for seedling/sapling 

establishment, and creosote bush itself will serve 

as a nurse plant for certain other species (Whitford 

2002). Creosote bush often forms monotypic 

stands that extend for miles, although its density 

varies greatly from site to site, with the greatest 

densities occurring in the Chihuahuan Desert (up 

to 8,000 plants/ha), and lesser densities in the 

Sonoran (up to 4,000 per ha) and the Mojave (up 

to 1,800 per ha) (Mabry and others 1977). Aboveground standing crop biomass tends to reflect 

these density trends. Chihuahuan Desert 

populations tend to show a greater size or age class 

structural diversity than those in other deserts 

(Mabry and others 1977). The resinous coating on 

the leaves helps retard water loss. During extreme 

droughts creosote bush will produce smaller, 

tougher leaves, and will also shed many of its 

older leaves. It quickly produces new shoot 

growth after substantial summer rains. Creosote 

bush has the C3 photosynthetic pathway, with 

peak photosynthetic activity (Pmax) in the spring 

and during summer monsoon seasons especially 

when pre-dawn water potentials are high 

(Whitford 2002). Pmax is as high as that found in 

C4 plants (Fitter and Hay 1987), and creosote 

shrubland has a similar community energy balance 

as the C4 desert grasslands they have invaded 

(Dugas and others 1996). Creosote bush has 

demonstrated a threshold for stomatal closure at -

5.8 MPa, has maintained full turgor at -3.0 MPa, 

with positive net photosynthesis occurring at -8 to 

-16 MPa (Fitter and Hay 1987, Mabry and others 

1977). It can tolerate leaf water potentials below -

50 MPa. It is able to maintain a relatively stable 

photosynthetic rate all year in Death Valley, with 

increases associated with spring temperatures and 

moisture availability (Mabry and others 1977). It 

has high water use efficiency and ability to resorb 

some nutrients, especially phosphorus, 72 to 86 

percent, while nitrogen resorption is within the 

general range for other evergreen shrubs, 47 to 57 

percent. Creosote bush stem and leaf orientations 

maximize light collection during cooler and 

moister early morning periods. Creosote bush has 

a greater ability to withstand xylem cavitation 

during droughts than riparian species (Pockman 

and Sperry 2000). An Arizona population of 

creosote bush experiences freezing-induced xylem 

vessel cavitation and consequent disruption of 

water transport at temperatures from -11 to -20 °C 

(Pockman and Sperry 1997). These temperatures 

correspond to 20+ year minimum isotherms and 

also correspond with the northern distribution in 

both the Sonoran and Mojave Deserts. A few 

studies have examined the effects of 

environmental pollutants on creosote bush. Foliage 

demonstrates leaf injury when exposed to acute 

levels of sulfur dioxide (SO2) (Olyszyk and others 

1987) and elevated CO2 concentrations may 

ameliorate the effects of heat stress (Hamerlynck 

and others 2000). Creosote foliage may 

accumulate heavy metals from industrial pollution 

as demonstrated in a 1980 to 1995 study from El 

Paso, TX, and Ciudad Juarez, Mexico (Mackay 

and others 1998). 

Reproduction.—Creosote bush blooms 

periodically during the growing season, blooming 

correlated first with seasonality, secondarily with 

the timing of moisture availability and other local 

environmental factors. Peak blooming periods 

occur in spring (February to May) and again in late 

summer into the fall (August to December). A 

single mature shrub may produce more than 2,000 

flowers (or about 50 g of flowers) during the 

growing season. Numerous insects visit the 

flowers, but bees are the most effective pollinators. 

Many species of bees are known to visit the 

flowers; some are generalists but some are 

specialists and forage solely on creosote bush 

(Minckley and others 2000). It is an outcrossing 

species but is always self-compatible (Mabry and 

others 1977) with localized populations having 

higher levels of inbreeding in the Chihuahuan 

Desert. The flowers are borne solitary in the axils, 

are 2 to 3 cm in diameter, are perfect, with five 

yellow clawed petals, obovate to spatulate, 6 to 10 

mm long, and 5 mm wide. There are five 

yellowish green, unequal, deciduous sepals, 5 to 8 

mm long. There are 10 stamens with winged 

filaments, and the ovary has five locules. The 
421

petals twist at a 90 degree angle after pollination. 

The fruit is a roundish capsule, about 5 to 7 mm in 

diameter, covered with a dense concentration of 

white to reddish hairs, and separates into five 

indehiscent one-seeded carpels at maturity (Carter 

1997, Correll and Johnston 1970, Kearney and 

others 1951, Martin and Hutchins 1980-81). Both 

fruits and flowers are commonly found on the 

plant at the same time. Flowering may begin on 

plants as young as 4 to 6 years, although heavy 

flowering and fruiting generally occurs after 8 to 

13 years. Greenhouse-raised seedlings may flower 

at 2 years of age. Fruit production ranges from 39 

to 278 fruits per 100 g of branches or 120 to 1,710 

per plant (Young and Young 1992). There are 

about 370 seeds/g. The seeds have mechanical 

dormancy and scarification aids germination 

(Young and Young 1992). Germination success 

varies by population, latitudinal and local 

environmental conditions with reported success 

rates of 2 to 70 percent. Salt (NaCl) at 10,000 ppm 

(-0.78 MPa) resulted in 0 percent germination, and 

seed germinated equally well at pH's from 7 to 10 

(Barbour 1968), although some studies have 

shown some variation from these pH's (reviewed 

in Baskin and Baskin 2001). Leachates from fruit 

coats inhibit germination of some plant species 

[Bouteloua eriopoda (Torr.) Torr. but not 

Muhlenbergia porteri Scribn. ex Beal], but not of 

creosote bush seeds (Baskin and Baskin 2001). 

Seeds stored dry in paper bags at room 

temperature for as long as 8 years showed some 

germinability (Barbour 1968). 

Growth and Management.—Growth response 

and productivity can be rapid after rain, but there 

is not a strong pattern of productivity and rainfall, 

while nitrogen has been shown as an important 

factor in creosote bush productivity (Whitford 

2002). Maximum leaf longevity is 16 months. 

Maximal root growth occurs in the fall. Branches 

will root if covered with soil. Ring counts of stems 

have indicated that creosote bush is long-lived, at 

least 30 to 50 years, but ring counts may not be 

reliable. Individual shrubs may sometimes 

reproduce by fragmentation or separation of the 

root crowns along with new basal sprouting, 

causing the plant to spread out and away from the 

center. Such fragmentation may contribute to longterm survival estimated at more than 100 years to 

several thousand years. Creosote bush shrubland 

has been expanding over the past 100 years or 

more due to overgrazing of desert grasslands by 

livestock, although climate trends have also been 

implicated. One experiment to test this involved 

transplanting seedling creosote bushes into 

fertilized, irrigated, or unmodified ungrazed black 

grama (B. eriopoda) grassland, a creosote 

shrubland, or a heavily grazed grassland. The only 

site in which seedlings survived was in the heavily 

grazed grassland (Whitford 2002). Prior to heavy 

livestock grazing creosote bush was generally 

restricted to well-drained gravelly or sandy soils of 

steep slopes. Creosote bush and mesquite 

(Prosopis glandulosa Torrey) invasion likely 

occurred in episodes in conjunction with both 

reduced grass cover and drought (Grover and 

Musick 1990). Localized decreases have also been 

reported. In an area of Big Bend National Park, 

previously overgrazed by livestock, the dry period 

from 1960 to 1967 showed increased creosote 

bush cover, while the wetter period from 1967 to 

1981 showed a decrease in creosote bush cover 

and increased cover of perennial grasses, forbs, 

and most other shrubs (Wondzell and Ludwig 

1995). At a site in southern New Mexico creosote 

shrub size has decreased over a 10-year period, 

possibly due to longer term climate change (Miller 

and Huenneke 1996). Certain creosote bush 

populations in the Sonoran Desert of Mexico 

declined by 50 to 90 percent, perhaps due to 

prolonged droughts from 1936 to 1964 (Turner 

1990). Creosote bush shrublands have been 

experimentally treated with a variety of herbicides 

for shrub management. Several liquid-applied 

herbicides (2,4-D, dicamba, picloram, 2,3,6-TBA) 

either singly or in combination with other 

herbicides have demonstrated greater than 50 

percent root kill (Herbel and Gould 1995). But 

individual shrub treatment with dry herbicides 

(bromacil, fenuron) or aerially applied tebuthiruon 

pellets have given greater control, 80 to 90 

percent. Perennial grass and annual forb 

production increased substantially on tebuthiurontreated plots. 

Benefits and Disadvantages.—Creoste bush is 

unpalatable as livestock forage and is usually 

toxic, sometimes causing death (Gay and Dwyer 

1998, Kearney and others 1951, Stubbendieck and 

others 1997, USDA 1937). Jackrabbits are one of 

the few mammals that will consume the plant to a 

certain degree, largely based on the water content 

of a particular plant (Whitford 2002). Some 

woodrat populations will also consume creosote 

bush, demonstrating differentially developed 

tolerances to the resin (Mangione and others 

2000). Woodrats consume creosote leaves and 

stems in the Mojave Desert when necessary to 

maintain water balance but suffer ill effects such 

as loss of body mass (Karasov 1989). Both rabbits 

and woodrats will peel open the stems to get water. 
422 

Once a particular shrub is browsed by a jackrabbit, 

that same shrub is much more likely to be browsed 

again than another shrub that has never been 

browsed (Ernest 1994). Younger leaves have a 

higher concentration of resins that affect relative 

leaf palatability and digestibility. Resin content 

may be as high as 26 percent in young leaves, 

dropping to 11 to 16 percent in older leaves, with 

insects showing preference for the older leaves 

(Mabry and others 1977). A phenoloxidase system 

in the leaves appears to make digestibility of the 

resinous leaves even more difficult. Chemical 

treatment of creosote foliage to remove the resins 

results in nutritious edible forage, but due to 

economics this practice is not often implemented 

(Mabry and others 1977). Numerous species of 

arthropods set up habitation in creosote bush. In 

particular males of the grasshopper species, 

Ligurotettix coquilletti McNeil, select a particular 

shrub as their territory and defend it against other 

male grasshoppers (Mabry and others 1977). The 

mounds created by kangaroo rats (Dipodomys

spp.) appear to have a positive effect on creoste 

bush survival, flowering, and fruiting, while 

creosote bush itself has a negative effect on 

kangaroo rat populations (Chew and Whitford 

1992). Creosote bush exposure to 0.1 to 0.2 ppm 

ozone consistently reduced the toxic 

nordihydroguaiaretic acid content of leaves, which 

could result in potentially greater levels of 

herbivory (Gonzalez and others 1988). Powdered 

creosote leaves added to stored pinto beans made 

them significantly less attractive to the bean 

weevil (Zabrotes subfasciatus Boheman) (CortezRocha and others 1993). 

Pharmaceutical Chemistry.—Although a toxic 

plant for livestock and many wildlife, it has value 

due to this same toxicity because of antiseptic and 

other potential medicinal uses. Native Americans 

of the Southwestern deserts have used this plant in 

teas, tinctures, and salves, as a poultice to retard 

bacterial growth, as an emetic, expectorant, and 

diuretic to treat venereal disease, tuberculosis, 

bowel cramps, and rheumatism (Kearney and 

others 1951, Mabry and others 1977). It has been 

used for stiff limbs, sores, skin ailments, 

snakebites, menstrual cramps, and chicken pox 

(Bowers and Wignall 1993, Mabry and others 

1977). Creosote bush may cause dermatitis in 

certain people with allergic skin sensitivity. 

Secretions and contents of the lac scale insect 

found on creosote bush have been used for 

repairing pottery, fastening arrow points, water 

proofing baskets, and making mosaics (Kearney 

and others 1951, Warnock 1974). One of the 

principal compounds in creosote bush resin is the 

anti-oxidant nordihydroguaiaretic acid (NDGA). 

This is a phenolic aglycone and comprises 5 to 10 

percent of leaf dry weight, 80 percent of all 

phenolics in the resin, and once was used as an 

anti-oxidant in foods, pharmaceuticals, and 

industrial materials (Mabry and others 1977). 

NDGA has demonstrated an ability to inhibit 

tumors and cancers but does have toxic side effects 

to both animals and humans. Its use as a 

chemoprevention for skin cancer and protection 

against the effects of ultaviolet light has been 

investigated (Gonzales and Bowden 2002). The 

botanical dietary supplement "chapparal" has been 

associated with nonviral toxic hepatitis and 

contains lignans similar to estrogens (Obermeyer 

and others 1995) and may cause acute liver 

damage (Sheikh and others 1997). NDGA has 

shown potential to inhibit the human 

immunodeficiency virus (HIV) (Gnabre and others 

1995). NDGA (masoprocol) lowered glucose and 

triglyceride levels in rats and may have potential in 

Type II diabetes treatment (Reed and others 1999). 

Numerous flavanoid aglycones, flavanoid 

glycosides, wax esters, saponins, and volatile 

constituents such as terpenes, have been identified 

from creosote bush. 

References

Barbour, M.G. 1968. Germination requirements of 

the desert shrub Larrea divaricata. Ecology 49: 

915-923. 

Baskin, C.C. and J.M. Baskin. 2001. Seeds: 

Ecology, Biogeography, and Evolution of 

Dormancy, and Germination. Academic Press, 

San Diego, CA. 666 p. 

Bowers, J.E. and B. Wignall. 1993. Shrubs and 

Trees of the Southwest Deserts. Southwest Parks 

and Monuments Association. Tucson, AZ. 140 

p. 

Carter, J.L. 1997. Trees and Shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Chew, R.M. and W.G. Whitford. 1992. A longterm positive effect of kangaroo rats, Dipodomys 

spectabilis, on creosote bushes Larrea tridentata. 

Journal of Arid Environments 22(4): 375-386. 

Correll, D.S. and M.C. Johnston. 1970. Manual of 

the Vascular Plants of Texas. Texas Research 

Foundation, Renner, TX. 1,881 p. 
423

Cortez-Rocha, M.O., R.I. Sanchez-Martinez, G. 

Garcia-Sanchez, M.I. Villaescusa-Moreno, and 

F.J. Cinco-Moroyoqui. 1993. Plant powders as 

stored grain protectants against Zabrotes 

subfasciatus (Boheman). Southwestern 

Entomologist 18(1): 73-75. 

Dugas, W.A., R.A. Hicks, and R.P. Gibbens. 

1996. Structure and function of C-3 and C-4 

Chihuahuan Desert plant communities: energy 

balance components. Journal of Arid 

Environments 34(1): 63-79. 

Ernest, K.A. 1994. Resistance of creosote bush to 

mammalian herbivory: temporal consistency and 

browsing induced changes. Ecology 75(6): 

1,684-1,692. 

Fitter, A.H. and R.K.M. Hay. 1987. 

Environmental Physiology of Plants, 2nd Ed. 

Academic Press, London. 423 p. 

Gay, C.W., Jr. and D.D. Dwyer. 1998. Reprint. 

New Mexico Range Plants. Cooperative 

Extension Service Circular 374. Revisions by: 

C. Allison, S. Hatch, and J. Schickedanz. New 

Mexico State University, Las Cruces, NM. 84 

p. 

Gnabre, J.N., J.N. Brady, D.J. Clanton, Y. Ito, J. 

Dittmer, R.B. Bates, and C.C. Huang-Ru. 1995. 

Inhibition of human immunodeficiency virus 

type 1 transcription and replication by DNA 

sequence-selective plant lignans. Proceedings of 

the National Academy of Sciences of the United 

States of America. 92(24): 11,239-11,243. 

Gonzales, M. and G.T. Bowden. 2002. 

Nordihydroguaiaretic acid-mediated inhibition 

of ultraviolet B-induced activator protein-1 

activation in human keratinocytes. Molecular 

Carcinogenesis 34(2): 102-111. 

Gonzalez, C.A., C.S. Wisdom, and P.W. Rundel. 

1988. Ozone impact on the antioxidant 

nordihydroguaiaretic acid content in the external 

leaf resin of Larrea tridentata. Biochemical 

Systematics and Ecology 16(1): 59-64. 

Grover, H.D. and H.B. Musick. 1990. Shrubland 

encroachment in southern New Mexico, USA, 

an analysis of desertification processes in the 

American Southwest. Climatic Change 17(2-3): 

305-330. 

Hamerlynck, E.P., T.E. Huxman, M.E. Loik, and 

S.D. Smith. 2000. Effects of extreme high 

temperature, drought and elevated CO2 on 

photosynthesis of the Mojave Desert evergreen 

shrub, Larrea tridentata. Plant Ecology 148(2): 

183-193. 

Herbel, C.H. and W.L. Gould. 1995. Management 

of Mesquite, Creosotebush, and Tarbush with 

Herbicides in the Northern Chihuahuan Desert. 

New Mexico State University and United States 

Department of Agriculture, Agricultural 

Research Service Experiment Station Bulletin 

775. Las Cruces, New Mexico. 53 p. 

Hickman, J.C., ed. 1993. The Jepson Manual: 

Higher Plants of California. University of 

California Press, Berkeley, CA. 1,400 p. 

Hunter, K.L., J.L. Betancourt, B.R. Riddle, T.R. 

Van Devender, K.L. Cole, and W.G. Spaulding. 

2001. Ploidy race distributions since the last 

glacial maximum in the North American desert 

shrub, Larrea tridentata. Global Ecology and 

Biogeography 10(5): 521-533. 

Karasov, W.H. 1989. Nutritional bottleneck in a 

herbivore, the desert wood rat, Neotoma lepida. 

Physiological Zoology 62(6): 1,351-1,382. 

Kartesz, J.T. 1994. A Synonymized Checklist of 

the Vascular Flora of the United States, Canada, 

and Greenland, Vol. 1, 2nd Ed. Biota of North 

America Program of the North Carolina 

Botanical Garden. Timber Press, Portland, OR. 

622 p. 

Kearney, T.H., R. Peebles, and Collaborators. 

1951 with 1960 supplement. Arizona Flora. 

University of California Press, Berkeley, CA. 

1,085 p. 

Lia, V.V., V.A. Confalonieri, C.I. Comas, and J.H. 

Hunzikert. 2001. Molecular phylogeny of 

Larrea and its allies (Zygophyllaceae): reticulate 

evolution and the probable time of creosote bush 

arrival in North America. Molecular 

Phylogenetics and Evolution 21(2): 309-320. 

Mabry, T.J., J.H. Hunziker, and D.R. DeFeo, Jr., 

eds. 1977. Creosote Bush: Biology and 

Chemistry of Larrea in New World Deserts. 

United States/International Biological Program 

Synthesis Series 6. Dowden, Hutchinson, and 

Ross, Inc., Stroudsburg, PA. 284 p. 
424 

Mackay, W.P., R. Mena, N.E. Pingitore, Jr., K. 

Redetzke, C.E. Freeman, H. Newman, J. 

Gardea, and H. Navarro. 1998. Seasonal 

changes in concentration and distribution of 

heavy metals in creosotebush, Larrea tridentata

(Zygophyllaceae), tissues in the El Paso, 

TX/Ciudad Juarez, Mexico area. SidaContributions to Botany 18(1): 287-296. 

Mahall, B.E. and R.M. Callaway. 1992. Root 

communication mechanisms and 

intracommunity distribution of two Mojave 

Desert shrubs. Ecology 73(6): 2,145-2,151. 

Mangione, A.M., M.D. Dearing, and W.H. 

Karasov. 2000. Interpopulation differences in 

tolerance to creosote bush resin in desert 

woodrats (Neotoma lepida). Ecology 81(8): 

2,067-2,076. 

Martin, W.C. and C.E. Hutchins. 1980-1981 

(reprinted 2001). A Flora of New Mexico, Vol. 

1. Bishen Singh Mahendra Pal Singh, India and 

Koeltz Scientific Books, Germany. 1,276 p. 

Miller, R.E. and L.F. Huenneke. 1996. Size 

decline in Larrea tridentata (creosotebush). 

Southwestern Naturalist 41(3): 248-250. 

Minckley, R.L., J.H. Cane, and L. Kervin. 2000. 

Origins and ecological consequences of pollen 

specialization among desert bees. Proceedings of 

the Royal Society of London Series B Biological 

Sciences 267(1440): 265-271. 

Obermeyer, W.R., S.M. Musser, J.M. Betz, R.E. 

Casey, A.E. Pohland, and S.W. Page. 1995. 

Chemical studies of phytoestrogens and related 

compounds in dietary supplements: flax and 

chaparral. Proceedings of the Society for 

Experimental Biology and Medicine 208(1): 6-

12. 

Olszyk, D.M, A. Bytnerowicz, C.A. Fox, G. Kats, 

P.J. Dawson, and J. Wolf. 1987. Injury and 

physiological responses of Larrea tridentata DC 

Coville exposed in-situ to sulfur dioxide. 

Environmental Pollution 48(3): 197-212. 

Pockman, W.T. and J.S. Sperry. 1997. Freezinginduced xylem cavitation and the northern limit 

of Larrea tridentata. Oecologia 109(1): 19-27. 

Pockman, W.T. and J.S. Sperry. 2000. 

Vulnerability to xylem cavitation and the 

distribution of Sonoran Desert vegetation. 

American Journal of Botany 87(9): 1,287-1,299. 

Reed, M.J., K. Meszaros, L.J. Entes, M.D. 

Claypool, J.G. Pinkett, D. Brignetti, J. Luo, A. 

Khandwala, and G.M. Reaven. 1999. Effect of 

masoprocol on carbohydrate and lipid 

metabolism in a rat model of Type II diabetes. 

Diabetologia 42(1): 102-106. 

Sheikh, N.M., R.M. Philen, and L.A. Love. 1997. 

Chaparral-associated hepatotoxicity. Archives of 

Internal Medicine 157(8): 913-919. 

Stubbendieck, J., S.L. Hatch, and C.H. Butterfield. 

1997. North American Range Plants, 5th Edition. 

University of Nebraska Press, Lincoln, NE. 501 

p. 

Turner, R.M. 1990. Long-term vegetation change 

at a fully protected Sonoran Mexico desert site. 

Ecology 71(2): 464-477. 

U.S. Department of Agriculture Forest Service. 

1937 (1988 Dover edition). Range Plant 

Handbook. Dover Publications Inc., NY. 816 p. 

Warnock, B.H. 1974. Wildflowers of the 

Guadalupe Mountains and the Sand Dune 

Country, Texas. Sul Ross State University, 

Alpine, TX. 176 p. 

Whitford, Walter G. 2002. Ecology of Desert 

Systems. Academic Press, San Diego, CA. 343 

p. 

Wondzell, S. and J.A. Ludwig. 1995. Community 

dynamics of desert grasslands: influence of 

climate, landforms, and soils. Journal of 

Vegetation Science 6(3): 377-390. 

Young, J. A. and C. G. Young. 1992. Seeds of 

Woody Plants in North America. Dioscorides 

Press, Portland, Oregon. 407 p. 
425

Lasiacis divaricata (L.) A.S. Hitchc. wild bamboo 

POACEAE 

Synonyms: Panicum bambusoides Desv. ex Ham. 

 Panicum chauvinii Steud. 

 Lasiacis harrisii of Britton and Wilson 

 Lasiacis sloanei of Britton and Wilson 

Panicum divericatum L. var. glabrum Kuntze 

 Panicum divericatum L. var. stenostachyum Griseb. 

John K. Francis 

General Description.—Wild bamboo, also known 

as smallcane, pitillo de monte, tibisí, and chico, is 

an evergreen, slender, woody, climbing or 

scrambling shrub that sometimes reaches 5 m in 

length and is commonly about 6 mm in basal stem 

(culm) diameter. There are usually several stems 

from the root crown. The plant develops an 

extensive and abundant system of fibrous roots. 

The whole pant is more or less glabrous. The culms 

are hollow between nodes, thin-walled, brittle, and 

tough, so that they have low bending strength but 

high tensile strength. The lower internodes are 

greenish yellow and smooth. There are few or no 

branches on the culms until near the ends. 

Relatively few leaves are located on recently 

grown branches. The alternate leaves are narrowly 

lanceolate, 5 to 20 cm long and 5 to 15 cm wide. 

Inflorescences are terminal panicles with few 

branches. The fruits are ovoid, about 4 mm long, 

and covered (at maturity) with a thin, black 

pericarp (Hitchcock 1935, Nelson 1996, Stevens 

and others 2001). There are 2n = 36 chromosomes 

(CromoPar 2002). 

Range.—Wild bamboo is native to southern 

Florida, the West Indies, and from Central Mexico 

to northern Argentina (Stevens and others 2001). 

Stevens and others (2001) report two varieties, 

divaricata and leptostachya (Hitchc.) Davidse, in 

Nicaragua. The species is not reported to have 

naturalized outside its native range. 

Ecology.—Wild bamboo is intermediate in its 

tolerance to shade. It normally grows in the 

understory of medium to low-density forest, in 

small openings, and in brushy areas. It is common 

along shady forest trails and roads, and sometimes 

invades brushy pastures. In Puerto Rico, it grows 

from near sea level to about 800 m in elevation in 

areas that receive from about 1000 to 2400 mm of 

mean annual precipitation. Wild bamboo grows up 

to an elevation of 1300 m in Nicaragua (Stevens 

and others 2001). The species colonizes soils with 

the full gamut of textures and pH’s from about 5.5 

to 7.5 in areas of sedimentary (including 

limestone), igneous, and metamorphic (including 

ultramaphic) rocks. The soils are usually well 

drained. 

Reproduction.—Wild bamboo flowers throughout 

the year except during periods of high drought 

stress. The flowers are assumed to be wind 

pollinated. Collections of air-dried seeds from two 

areas in Puerto Rico in different years yielded 

averages of 0.0064 ± 0.0001 and 0.0156 + 0.0001 

g/seed or roughly 90,000 seeds/kg. Some 

pretreatment is probably needed: one lot yielded 0 
426 

percent germination and the other 11 percent. The 

seeds are somewhat delicate and probably damaged 

by scarification. The pericarp can be removed by 

rubbing with great care with unknown effect on 

germination. It is assumed that birds and possibly 

rodents are seed dispersers. Seedlings are not 

common. Culms layer (root) whenever they come 

in contact with the ground and are covered with 

litter. Wild bamboo sprouts when cut or damaged 

to renew damaged culms. 

Growth and Management.—Growth of wild 

bamboo from sprouts is rapid. Sprouts from mature 

plants may reach most of their eventual length 

during the first year. Thereafter, branches are 

added for 1 or 2 years. After old culms die, new 

ones grow from the root crown. It is not known 

how fast seedlings grow. Wild cane seldom forms 

thickets and mostly grows in forests and brush 

lands so that control does not seem warranted. If it 

should be needed, grubbing out the root crowns or 

cutting and spot-spraying the resulting sprouts with 

glyphosate would probably be effective. 

Benefits.—Wild bamboo furnishes food and cover 

for wildlife and helps protect the soil. The leafy 

and more succulent portions are browsed by horses 

and cattle. The species seems resistant to light and 

moderate browsing. The tough stems, which often 

run across low brush, make it difficult to walk 

through the forest. 

References 

CromoPar. 2002. Recuentos cromosómicos 

foráneos de plantas presentes en Paraguay. 

Universidad de Barcelona, Spain. http:// 

www.ub.es/botanica.cromopar.cro-out/pdf. 24 p. 

Hitchcok, A.S. 1935. Manual of the grasses of the 

United States. Miscellaneous Publication 200. 

U.S. Department of Agriculture, Washington, 

DC. 1,040 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

390 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanic Garden Press, St. Louis, 

MO. p. 1,911-2,666. 
427

Leandra krugii (Cogn.) Judd & Skean leandra 

MELASTOMATACEAE 

Synonyms: Calycogonium krugii Cogn. 

John K. Francis 

General Description.—Leandra is an evergreen 

shrub usually 1 to 1.5 m in height and 1 to 3 cm in 

basal diameter, but rarely reaching 3 m in height 

and 6 cm in basal diameter. Multiple stems from 

basal sprouts are the rule and stems are covered 

with finely furrowed brown bark. Young plants 

have deep taproots; older plants have deep and 

extensive root systems. Branches and twigs are 

stiff, slender, and tend to grow vertically. 

Trichotomous branching is common. The green to 

dark-green leaves are opposite, usually oval to 

elliptic, rough, and coriaceous. They are 2 to 5.5 

cm long and have a 6-mm petiole. Edges are 

serrulate. Five nerves are raised on the underside, 

the three principal ones radiating from a point 

above the base of the leaf. The six-parted white 

flower is about 3 cm across. Fruits (berries) are 

globose, bright red, and about 12 mm in diameter. 

They are edible and contain many minute seeds 

(author’s observations, Liogier 1995). 

Range.—Leandra is endemic to central and 

western Puerto Rico (Liogier 1995). It is not 

known to have been planted or naturalized outside 

Puerto Rico. 

Ecology.—Leandra is intermediate in tolerance. As 

single plants and in thickets, it grows naturally in 

small openings and in the understory of low to 

moderate basal-area remnant and secondary 

forests, and in plantations. It occurs at elevations of 

about 600 to 1000 m in areas that receive from 

about 1800 to 3000 mm of mean annual 

precipitation. The soils it grows on are usually 

moderately acid, loamy or clayey, deep or rocky, 

well drained to somewhat poorly drained Ultisols, 

Alphisols, and Inceptisols. These soils develop 

from sedimentary, igneous, and metamorphic 

(ultramaphic) rocks. Leandra is most common on 

side slopes and ridge tops, in forest plantations, 

secondary forest, and primary remnants (author’s 

observations). 

Reproduction.—Leandra blooms during the late 

spring and early summer (April through June) and 

fruits during late summer and fall (August through 

October). The flowers appear to be insect 

pollinated. There are several million seeds per 

kilogram. They germinated profusely when 

smeared on the surface of potting mix and placed 

in a mist bed. The minute seedlings did not survive, 

however. The plant can be propagated by air 

layering. Thirty-nine of 46 air layers treated with 

0.3 percent IBA on wild plants rooted in 6 months. 

Wild plants layer whenever branches come in 

contact with the ground. Woody cuttings treated 

with IBA (indol-butyric acid) and misted failed to 

successfully root. Apparently, birds are the 

principal dispersers of seeds. Reproduction can be 

abundant in shaded areas that have been disturbed 

(author’s observations). 

Growth and Management.—Leandra grows 

slowly as a seedling and at a moderate rate from 

sprouts. Plants appear to be relatively long-lived. 

Management experience is lacking or has not been 

published. Disturbance of the understory of forests 

in suitable habitat with low to moderate basal area 

in which mineral soil is exposed should result in 

the establishment of seedlings. Stands could be 

thickened by ground-layering existing plants. 

Benefits.—Leandra adds to the aesthetics of the 

forests where it grows, helps protect the soil, and 

furnishes food and cover for wildlife. The 

convenient size, pretty flowers, and foliage 

(W3

Tropicos 2002), as well as the attractive fruits 

suggest leandra as an ornamental. 
428 

References 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

W3

Tropicos. 2002. Calycogonium krugii Cogn., 

flowering shrub.http://digitalis.mobot.org/mrsid/ 

bin/mosid/mosid.pl?client=203&image=MOA06799_001.sid&title=Calycogonium+krugii+ Cog... 

1 p. 
429

Ledum groenlandicum Oeder Labrador tea 

ERICACEAE 

Synonyms: Ledum palustre L. ssp. groenlandicum (Oeder) Hulten 

 Ledum palustre L. var. latifolium (Jacq.) Michx. 

 Ledum latifolium Ait. 

 Rhododendron groenlandicum (Oeder) Kron & Judd 

James E. Nellessen 

General Description.—Labrador tea is an 

evergreen, well branched, spreading shrub up to 

about 1 m in height. The twigs are densely hairy 

and the buds scaly. The leaves are simple, 

alternate, entire (not toothed), lanceolate, 

somewhat narrowly elliptic to oblong, 1.5 to 5 cm 

long, about 0.7 to 2 cm wide, thick, leathery and 

evergreen with rolled margins, dark green above, 

densely off-white to rusty hairy beneath, and 

fragrant when crushed (Gleason and Cronquist 

1963, Britton and Brown 1913). Several 

synonyms, as given above, may be found for this 

species (Kartesz 1994, Britton and Brown 1913). 

The genus Ledum appears to be monophyletic, and 

cladistic analysis places it as most closely related 

to genus Rhododendron subsection Edgeworthia, 

hence its synonym placing it in the genus 

Rhododendon (Kron and Judd 1990). 

Range.—This shrub occurs from Greenland, 

across Canada from Labrador to British Columbia, 

north to Alaska, south to New Jersey, 

Pennsylvania, Massachusetts, Michigan, 

Wisconsin, Minnesota, and Washington, with 

isolated sites or occurrences in Ohio and South 

Dakota (Gleason and Cronquist 1963, Britton and 

Brown 1913, Great Plains Flora Association 1986, 

McCance and Burns 1984). Labrador tea has been 

classified as an endangered species in Ohio, where 

one population occurs in a bog habitat in the 

northeastern corner of the state (McCance and 

Burns 1984). One isolated collection has occurred 

in the Black Hills of South Dakota (Great Plains 

Flora Association 1986). 

Ecology.—This species grows in acidic soils of 

bogs, especially sphagnum (Sphagnum spp.) bogs, 

and other swampy and wet shoreline habitats in 

association with Picea mariana (Mill.) BSP (black 

spruce), Chamaedaphne calyculata (L.) Moench. 

(leatherleaf), and Kalmia spp. (laurels). Labrador 

tea will also grow on some drier upland soils with 

a reasonable moisture regime in association with 

Pinus spp. (pines) and Vaccinium spp. 

(blueberries). Stands of Labrador tea appear to 

retard black spruce growth and regeneration 

(Inderjit 1996). Labrador tea has been reported as 

an undergrowth species on serpentine substrates on 

mountain tops in eastern townships of Quebec, 

Canada. It has also been observed in shrub and 

sedge dominated serpentine ecotonal communities 

of Newfoundland with a soil pH of 6.27, and 

exchangeable soil mineral concentrations: calcium, 

13 μg/ml, magnesium 237 μg/ml, and nickel 0.72 

μg/ml (reviewed in Brooks 1987). 

Ecology in Human Impacted Environments.—

Exposure to sulfur dioxide has been shown to 

cause visible foliar symptoms and significantly 

reduce photosynthetic net assimilation rates, but 

the rate of reduction was significantly slower than 

that in deciduous species (Addison and others 

1984). In a radionuclide study, lead-210 and 

polonium-210 tended to accumulate in lichens and 

mosses, while radium-226 tended to accumulate in 

shrubs such as Labrador tea (Sheard 1986). 

Labrador tea may be an indicator of lead 

contamination in the environment in the vicinity of 

lead/zinc mines (Pugh and others 2002). Labrador 

tea was among four ericaceous shrub species 

naturally revegetating an abandoned vacuum-
430 

mined peatland, despite the absence of new 

sphagnum moss colonization (Berube and Lavoie 

2000). 

Reproduction.—Labrador tea flowers are 

relatively small, about 1 cm wide and 2 cm long, 

white, with five to seven stamens, and are grouped 

in terminal clusters. It blooms from May to July, 

sometimes as late as August. The fruit is a slender 

capsule, oval in shape, about 5 to 8 mm long and 2 

to 3 mm wide, with a persistent style (Gleason and 

Cronquist 1963, Britton and Brown 1913). The 

seeds are elongated, numerous and small. For 

germination, they should be placed onto a good 

moisture-supplying substrate and covered with a 

clear plastic film (Young and Young 1992). Seeds 

germinate best after a 30-day cold stratification 

period, with optimal germination temperatures of 

20 to 25 °C, and light is required (Calmes and 

Zasada 1982, Baskin and Baskin 2001). The seeds 

are considered to have non-deep physiological 

dormancy because of the relatively shorter cold 

periods needed and that some seeds can germinate 

shortly after ripening. The cold stratification 

period can decrease the germination temperature 

requirement. Full sunlight and high soil moisture 

seem to be requirements, including a low soil pH, 

e.g. 5.5 (Karlin and Bliss 1983, Baskin and Baskin 

2001). Apparently winter-time cuttings will root 

well (Dirr and Heuser 1987, Young and Young 

1992). 

Growth and Management.—This shrub is 

generally considered a slow growing, late 

successional, pre-climax or climax species. The 

low shrub stratum consisting of Labrador tea and 

leatherleaf was estimated to have an aboveground 

productivity of 0.4 and 2.0 t/ha/yr on perched and 

raised bogs, respectively, in northern Minnesota 

(Grigal and others 1985). Rust fungi of the genus 

Chrysomyxa alternate between spruce and 

members of the Ericaceae such as Labrador tea. 

Chrysomyxa reticulata sp. nov. may spread from 

Ledum spp. to cultivated rhododendrons (Crane 

2001). 

Benefits.—As the name implies, the dried leaves 

have been used for tea. Native Americans, such as 

the Chippewa, have used this shrub for various 

purposes. The root has been used for medicinal 

purposes to treat ulcers and the leaves for making 

tea (Densmore 1928). As a tea this plant has been 

used for treating asthma, colds, stomach aches, 

kidney problems, scurvy, and fevers (Foster and 

Duke 1990). Externally it has been applied as a 

wash for burns, ulcers, and stings. It has been used 

as a folk remedy for lung ailments, dysentery, 

indigestion, and to kill lice and treat leprosy. 

Successful traditional use for gout treatment may 

be due to the presence of phenolics and tannins 

within the plant (Owen and Johns 1999). This 

species, as well as others in the genus, has 

generally been regarded as having low palatability, 

being unpalatable, or even slightly poisonous 

(USDA 1937). Since this species often grows in 

wet boggy habitats, it is generally not accessible to 

most livestock. The plant may serve as reindeer 

forage in Alaska. Its unpalatability may also be 

due to essential oils of the monoterpene family 

(sabinene and limonene) and the sesquiterpene 

family (alpha- and beta-selinene and germacrone) 

(Belleau and Collin 1993). Germacrone has been 

shown to be a feeding deterrent for snowshoe 

hares (Lepus americanus Erxleben), although they 

do eat the plant to some extent (Reichardt and 

others 1990, MacCracken and others 1988). 

References

Addison, P.A., S.S. Malhotra, and A.A. Khan. 

1984. Effect of sulfur dioxide on woody boreal 

forest species grown on native soils and tailings. 

Journal of Environmental Quality 13(3): 333-

336. 

Baskin, C.C. and J.M. Baskin. 2001. Seeds: 

ecology, biogeography, and evolution of 

dormancy, and germination. Academic Press, 

San Diego, CA. 666 p. 

Belleau, F. and G. Collin. 1993. Composition of 

the essential oil of Ledum groenlandicum. 

Phytochemistry (Oxford) 33(1): 117-121. 

Berube, M.E. and C. Lavoie. 2000. The natural 

revegetation of a vacuum-mined peatland: eight 

years of monitoring. Canadian Field Naturalist 

114(2): 279-286. 

Britton, N.L. and A. Brown. 1913 (1970 Dover 

edition). An illustrated flora of the northern U.S. 

and Canada, Vol. 2. Dover Publications Inc., 

NY. 735 p. 

Brooks, R.R. 1987. Serpentine and its vegetation: 

A multidisciplinary approach. Dioscorides 

Press, Portland, OR. 454 p. 

Calmes, M.A. and J.C. Zasada. 1982. Some 

reproductive traits of four shrub species in the 

black spruce forest type of Alaska. Canadian 

Field Naturalist 96: 35-40. 
431

Crane, P.E. 2001. Morphology, taxonomy, and 

nomenclature of the Chrysomyxa ledi complex 

and related rust fungi on spruce and Ericaceae in 

North America and Europe. Canadian Journal of 

Botany 79(8): 957-982. 

Densmore, F. 1928 (1974 Dover edition). How 

Indians use wild plants for food, medicine and 

crafts. Dover Publications, New York. 397 p. 

Dirr, M.A. and C.W. Heuser. 1987. The reference 

manual of woody plant propagation. Varsity 

Press, Athens, GA. 

Foster, S.A. and J.A. Duke. 1990. A field guide to 

medicinal plants (eastern/central). Peterson Field 

Guide Series, Houghton Mifflin Co., Boston, 

MA. 366 p. 

Gleason, H.A. and A.R. Cronquist. 1963. Manual 

of the vascular plants of northeastern U.S. and 

adjacent Canada. D. Van Nostrand Co., New 

York. 810 p. 

Great Plains Flora Association. 1986. Flora of the 

Great Plains. University Press of Kansas, KS. 

1,392 p. 

Grigal, D.F., C.G. Buttleman, and L.K. Kernik. 

1985. Biomass and productivity of the woody 

strata of forested bogs in northern Minnesota. 

Canadian Journal of Botany 63(12): 2416-2424. 

Inderjit, A.U.M. 1996. Growth and physiological 

responses of black spruce (Picea mariana) to 

sites dominated by Ledum groenlandicum. 

Journal of Chemical Ecology 22(3): 575-585. 

Karlin, E.F. and L.C. Bliss. 1983. Germination 

ecology of Ledum groenlandicum and Ledum 

palustre ssp. decumbens. Arctic Alpine 

Research 15: 397-404. 

Kartesz, J.T. 1994. A synonymized checklist of 

the vascular flora of the United States, Canada, 

and Greenland, Vol. 1, 2nd Edition. Biota of 

North America Program of the North Carolina 

Botanical Garden. Timber Press, Portland, OR. 

622 p. 

Kron, K.A. and W.S. Judd. 1990. Phylogenetic 

relationships within the Rhodoreae Ericaceae 

with specific comments on the placement of 

Ledum. Systematic Botany 15(1): 57-68. 

MacCracken, J.G., W.D. Steigers, Jr., and P.V. 

Mayer. 1988. Winter and early spring habitat 

use by snowshoe hares, Lepus americanus, in 

south-central Alaska, USA. Canadian Field 

Naturalist 102(1): 25-30. 

McCance, R.M., Jr. and J.F. Burns, eds. 1984. 

Ohio endangered and threatened vascular plants: 

Abstracts of state listed taxa. Department of 

Natural Resources, Columbus, OH. 635 p. 

Owen, P.L. and T. Johns. 1999. Xanthine oxidase 

inhibitory activity of northeastern North 

American plant remedies used for gout. Journal 

of Ethnopharmacology 64(2): 149-160. 

Pugh, R.E., D.G. Dick, and A.L. Fredeen. 2002. 

Heavy metal (Pb, Zn, Cd, Fe, and Cu) contents 

of plant foliage near the Anvil Range lead/zinc 

mine, Faro, Yukon Territory. Ecotoxicology and 

Environmental Safety 52(3): 273-279. 

Reichardt, P.B., J.P. Bryant, B.J. Anderson, D. 

Phillips, T.P. Clausen, M. Meyer, and K. Frisby. 

1990. Germacrone defends Labrador tea from 

browsing by snowshoe hares. Journal of 

Chemical Ecology 16(6): 1961-1970. 

Sheard, J.W. 1986. Distribution of uranium series 

radionuclides in upland vegetation of northern 

Saskatchewan, Canada I: Plant and soil 

concentrations. Canadian Journal of Botany 

64(11): 2446-2452. 

U.S. Department of Agriculture Forest Service. 

1937 (1988 Dover edition). Range plant 

handbook. Dover Publications Inc., New York. 

816 p. 

Young, J. A. and C. G. Young. 1992. Seeds of 

woody plants in North America. Dioscorides 

Press, Portland, OR. 407 p. 
432 

Lepianthes peltata (L.) Raf. monkey’s hand 

PIPERACEAE 

Synonyms: Lepianthes umbellata (L.) Raf. 

 Piper peltatum L. 

 Heckeria peltata (L.) Kunth in L. 

 Pothomorphe peltata (L.) Miq. 

 Piper umbellatum L. 

 Pothomorphe dussii Trel. in Stehlé 

John K. Francis 

 

General Description.—Common names for 

monkey’s hand include cowheel bush, cow-foot 

leaf, baquiña, chapeau g’leau, bois-anisette, fèy a 

kè, and many others. It is a weak-stemmed shrub 

up to 2 m in height and 3 cm in basal diameter. 

This plant usually has a single stem, but may 

produce multiple sprouts from the root collar if the 

top is damaged. The stems have a vertical habit 

with no branches until the stems reach 0.5 to 1.0 m 

in height and then form only second order 

branches. A taproot does not develop in clayey 

soil, but the plant develops many robust, semifleshy lateral roots. Adventitious roots may form at 

the lower nodes. The stems may be glaberous or 

pubescent. The leaves have petioles 8 to 20 cm 

long. The leaf blades may be sub-orbicular to 

ovoid, peltate or not, and about 15 to 35 cm long 

and broad. They have a cordate base and an acute 

tip, with 11 to 15 palmate veins. Spikes of minute 

flowers in small groups are born on peduncles 1 to 

7 cm long. In these “fruits,” after development, 

have numerous 0.5 mm-long druplets embedded 

(Howard 1988, Liogier 1985). The stems, leaves, 

and young fruits have a pine-like aromatic smell. 

However, overripe spikes, have a smell reminiscent 

of oysters or cow’s breath. 

Range.—Monkey’s hand is native to Mexico, 

Central America, tropical South America, the 

Greater and Lesser Antilles, and Trinidad in the 

Americas, and Guinea, Sierra Leone, Ivory Coast, 

Ghana, and Nigeria in Africa (Burkill 1997, 

Howard 1988, Liogier 1985, Liogier and Martorell 

2000). The species is also reported as naturalized in 

Florida (Langeland and Stocker 2001). 

Ecology.—Monkey’s hand grows in full sunlight 

and in light to moderate shade. Continually moist 

soil is imperative. Rainfall may vary from about 

1500 to over 3000 mm of annual precipitation. The 

plant usually grows on loamy or clayey soils with 

relatively good fertility. In Puerto Rico, monkey’s 

hand grows from near sea level to over 1,000 m in 

elevation. Plants may grow singly or in clumps 

along streams and roads, in old fields and other 

disturbed areas, in forest tree-fall gaps, and on 

nursery benches. Disturbance and bare soil are 

necessary for successful establishment. 

Reproduction.—Monkey’s hand shrubs bloom 

and fruit continuously after reaching about 0.5 m in 

height. A collection of fresh fruits from Puerto 

Rico weighed an average of 0.95 ± 0.11 g. Seeds 

separated from these fruits weighed an average of 

0.000061 g/seed or 16,400,000 seeds/kg (air-dried). 

For unknown reasons, germination on moist filter 

paper failed entirely (author’s observations). Seeds 

are produced in large numbers and disbursed by 

fruit bats and birds. Apparently, only a small 

percent of the seeds germinate and develop, 

because seedlings are scattered and rarely 
433

abundant. Monkey’s hand plants are easy to 

propagate from cuttings. A small group of stem 

cuttings were treated with IBA and placed in a mist 

bed. After about 5 weeks, 85 percent had rooted. 

Because rooting takes place at the nodes, a node 

should be included in the buried portion of the 

cutting. 

Growth and Management.—Monkey’s hand 

plants live about 2 years but may perpetuate 

themselves by sprouting from the root collar. 

Plants also root at the nodes when nodes are buried 

with soil or leaf mold, or if laid prostrate. The plant 

is a common weed in plantations of cacao and oil 

palm in Africa (Burkill 1997). Monkey’s hand may 

be controlled by hand pulling. It is important to 

remove the pulled material from the site to prevent 

it from rerooting. If hand pulling is not possible, 

plants may be killed by spraying with a 20 percent 

solution of Garlon1 applied as a basal bark spray or 

by cutting the plants and applying a 50 percent 

solution to the cut stumps (Langeland and Stocker 

2001). 

Benefits and Medicinal Use.—Monkey’s hand 

plants are browsed by cattle and other ruminants. 

The species is pretty enough to be used as a border 

or background plant in landscaping, but there are 

no current reports of its use in landscaping. In 

several parts of Africa, the leaves and stems are 

cooked and eaten as a vegetable with meat and 

fish. It is also used as a condiment and as an 

ingredient in sauces. The fruits and leaves are used 

as bait in trapping fish and small birds (Burkill 

1997). In Haiti, monkey’s hand leaves are used in 

combination with other natural products in a 

treatment for hypertension (The Temple of Yehwe 

2001). In Africa, it is used in various ways in 

magical rites to promote good things and ward off 

the influences of evil fetishes. Leaves, root, and 

fruits of the plant are also used to control migraine 

and general pain, as an antiseptic, to control 

tapeworms, gonorrhea, urinary problems, and 

rheumatism (Burkill 1997). Monkey’s hand is used 

in many of the same ways in the West Indies 

(Liogier 1990).

 

1

Garlon is named here for identification purposes 

only. The Forest Service does not endorse any 

commercial product. 

References 

Burkill, H.M. 1997. The useful plants of West 

Tropical Africa. Vol. 4. Royal Botanic Gardens, 

Kew, UK. 969 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Langeland, K.A. and R.K. Stocker. 2001. Control 

of non-native plants in natural areas of Florida. 

University of Florida, Cooperative Extension 

Service. http://edis.ifas.ufl.edu/BODY_WG209. 

56 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands. Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Liogier, H.A. and L.F. Martorell. 2000. Flora of 

Puerto Rico and adjacent islands. 2nd Ed. 

Revised. Editorial de la Universidad de Puerto 

Rico, Río Piedras, PR. 382 p. 

The Temple of Yehwe. 2001. Treatments. 

http://www/vodou.org/treatmen.htm. 3 p. 
434 

Lepidospartum burgessii B.L. Turner gypsum scalebroom 

ASTERACEAE 

Synonyms: none

Juanita A. R. Ladyman 

General Description.—The common name for 

Lepidospartum burgessii is gypsum scalebroom or 

Burgess’s broomshrub. The botanical name comes 

from the Greek, “lepidotus” meaning scaly and 

“spartium” meaning broom (Gledhill 1992). There 

are two other species in the genus Lepidospartum

but neither is sympatric with gypsum scalebroom 

(Baldwin and others 2002). The range of 

Lepidospartum latisquamum S. Wats. extends from 

southern California to Utah and that of L. 

squamatum (Gray) Gray ranges from central 

California to Baja California (Baldwin and others 

2002). Gypsum scalebroom was relatively recently 

described (Turner 1977). Prior to that time, it is 

likely to have frequently been mistaken for 

rabbitbrush, Chrysothamnus nauseosus (Pallas ex

Pursh) Britt. Gypsum scalebroom is a woody shrub 

with numerous stems growing up to 1.2 m tall. 

The stems have multiple branches and are covered 

with silvery, matted, felt-like hairs out of which 

protrude numerous small oil blisters. The leaves 

are needlelike, alternate, and 5 to 12 mm long. 

There are three or four terminal flower heads on 

stems with three, rarely four, bright yellow flowers 

per head. The achenes are covered by dense white 

hairs and topped by a pappus of many slender 

bristles. Although gypsum scalebroom has been 

mistaken for rabbitbrush, the three rounded bracts 

of the involucre on the flower heads clearly 

distinguish it from any species in that genus. 

Range.—Gypsum scalebroom is both a geographic 

and substrate endemic being restricted to certain 

gypsum soils in an approximate 1,000 square km 

area in northern Culberson County, Texas, and 

southern Otero County, New Mexico. 

Ecology.—Gypsum scalebroom is found only on 

the gypsum flats at the fringe of the northern 

Chihuahuan Desert. Dick-Peddie (1993) designated 

the region Chihuahuan desert scrub whereas some 

authorities have regarded the whole area as part of 

the desert plains grassland (Humphrey 1958). The 

latter designation may be more appropriate for the 

relatively small areas where gypsum scalebroom

occurs because creosote bush, which is the 

dominant shrub of the Chihuahuan desert scrub 

vegetation type, does not occur with gypsum 

scalebroom. Shrubs and sub-shrubs associated 

with Gypsum scalebroom include Atriplex 

canescens (Pursh) Nutt., Opuntia leptocaulis

DeCandolle, and Yucca elata (Engelm.) Engelm. as 

well as Tiquilia hispidissima (Torr. & Gray) A. 

Richards. Poliomintha incana (Torr.) Gray is a 

common associate, and Allenrolfia occidentalis (S. 

Wats.) Kuntze is also observed to a lesser extent in 

Texas. Associated grasses include Bouteloua 

breviseta Vasey, Sporobolus nealleyi Vasey, and 

Sporobolus airoides (Torr.) Torr. Gallardia 

multiceps Greene, Senecio warnockii Shinners and 

species of Mentzelia and Isocoma are common 

forbs associated with gypsum scalebroom,

although their contribution to the ground cover is 

low. Plants colonize two habitats, semistabilized 

gypsum dunes and relatively compacted gypsum 

soils at the edge of dry alkaline lakes (Powell 

1998). In New Mexico the plants generally grow 

on stabilized, microbiotic-covered, gypsum soils 

with approximately 5 percent basal vegetation/litter 

cover. In Texas and at one site in New Mexico, 

shrubs grow on more mobile gypsum dunes with 

an average of 20 percent basal vegetation/litter 

cover (Ladyman and Gegick 2001). The plants 

occur singly or in scattered colonies. The climate in 

this area is relatively extreme with summer 

temperature highs between 34 o

C and 41 oC and 
435

winter temperature ranging from lows of –2 oC and 

highs of 14 oC. Freezing is common during this 

time. Precipitation comes primarily during the 

summer (ranging from 13.1 to 46.8 cm). However, 

substantial spring rains have been reported, and 

there is a high degree of year-to-year variability 

characteristic of most arid and semiarid climates 

(Noy-Mier 1973). 

Reproduction.—The peak of flowering is during 

late July through early September, but within a 

population, flowering can extend from late April to 

early October. The number of flowers per shrub is 

highly variable; from one flower to literally 

hundreds of flower heads on any given shrub. 

There does not appear to be a close relationship 

between the size of the plant and the number of 

flowers because the largest shrubs are often in a 

decadent condition, that is, with more than 50 

percent dead wood. In 1991/1992 and 1997 plants 

were counted in seven colonies in New Mexico 

(Huenneke 1991, Ladyman and others 1999). 

Approximately 15 percent of the individuals 

counted within those colonies in 1991/1992 were 

dead in 1997. The number of juvenile plants had 

also declined. Recruitment is only by clonal 

propagation. No seeds have been observed despite 

abundant yellow flowers and apparently adequate 

pollinators (Ladyman and others 1999). The results 

of studies on pollen germination on stigmas 

indicate an incompatibility system is operating 

(Ladyman and others 1999). The prematurely 

stunted pollen tube growth observed by florescent 

microscopy is indicative of sporophytic 

incompatibility, which is typical of members of the 

Asteraceae and other species with trinucleate 

pollen (Ladyman and others 1999, de Nettancourt 

1977). 

Growth and Management.—Gypsum scalebroom 

responds to irrigation and fertilization and can 

grow to over 1.5 m with luxurious silvery foliage 

in cultivation (author’s personal observation). 

Transplantation is possible, but the success rate is 

not known. In 1998, two potential disease problems 

were identified. Corythuca marmorata, a known 

pathogen of some Asteraceae species, caused leaf 

loss and stem necrosis. A potential pathogenic 

fungus, Alternaria alternata, was observed within 

some flowers (Rotem 1994, Ladyman and Gegick 

2001). Genetic diversity appears to exist between 

the Texas and New Mexico populations. The 

involucre length of plants growing in Texas was 

significantly longer than from those in New 

Mexico (Ladyman and Gegick 2001). It is 

unknown if individuals between these two 

populations are reproductively compatible. Some 

protective measures, including erecting exclosures, 

have been made for some colonies growing in an 

Area of Critical Concern and managed by the 

Bureau of Land Management (BLM 1997). This 

species appears to be facing relatively imminent 

extinction. From a simple model that assumed the 

rate of death was linear, extinction is predicted 

within 35 years (Ladyman and others 1999). 

Benefits.—Gypsum scalebroom is visited by a 

wide variety of arthropods and possibly provides 

cover and food for small wildlife species. It is 

unlikely to be palatable to livestock, but they do 

appear to brush against it even if they do not use it 

as food (anecdotal information from ranchers in 

area and author’s personal observation). This is 

likely due to the absence of other large shrubs in 

the region. It is a beautiful silvery ornamental 

shrub. It has been suggested that one way to slow 

its eventual extinction is to introduce it into the 

horticultural and landscape trade. 

References 

Baldwin, B.G., S. Boyd, B.J. Ertter, R.W. 

Patterson, T.J. Rosatti, and D.H. Wilken. 2002. 

The Jepson Desert Manual, vascular plants of 

southeastern California. University of California 

Press, Berkeley, CA. 624 p. 

Bureau of Land Management. 1997. Areas of 

Critical Environmental Concern Resource 

Management Plan Amendment, August 1997. 

Bureau of Land Management, Las Cruces 

District, Caballo Resource Area, Otero County, 

Las Cruces, NM [not paged]. 

De Nettancourt, D. 1977. Incompatibility in 

Angiosperms. Springer Verlag, Berlin, 

Heidelberg, and New York. 230 p. 

Dick-Peddie, W.A. 1993. New Mexico Vegetation 

– past, present, and future. University of New 

Mexico Press, Albuquerque, NM. 244 p. 

Gledhill, D. 1992. The names of plants. Second ed. 

Cambridge University Press, Cambridge UK. 

202 p. 

Huenneke, L.F. 1991. Biological studies of 

Lepidospartum burgessii, a rare endemic shrub 

of southern New Mexico – First Progress 

Report. Bureau of Land Management, New 

Mexico State Office, Santa Fe, NM [not paged]. 
436 

Humphrey, R.R. 1958. The desert grassland, a 

history of vegetational change and an analysis of 

causes. Botanical Review 24:193-252. 

Ladyman, J.A.R., L. DeLay, P. Gegick, M. Bogan. 

1999. Status and Reproductive Biology of 

Lepidospartum burgessii (Burgess broomshrub 

or gypsum broomscale). Unpublished report 

submitted to the USGS Biological Survey, 

Albuquerque, NM. 108 p. 

Ladyman, J.A.R. and Patricia Gegick. 2001. 

Status of Lepidospartum Burgessii (Burgess 

broomshrub or gypsum broomscale). In: 

Southwestern Rare and Endangered Plants. 

Proceedings of Third Conference. RMRS-P-23. 

USDA Forest Service, Rocky Mountain 

Research Station, Fort Collins, CO. p. 116-127. 

Noy-Mier, I. 1973. Desert ecosystems: 

environment and producers. In: Annual Reviews 

of Ecology and Systematics IV: 25-51. 

Powell, A. Michael. 1998. Trees & shrubs of 

Trans-Pecos and adjacent areas. University of 

Texas University Press, Austin, TX (First Ed. 

copyright Big Bend Natural History Assoc.) 498 

p. 

Rotem, J. 1994. The Genus Alternaria – Biology, 

Epidemiology, and Pathogenicity. APS, St. Paul, 

MN. 325 p. 

Turner, B.L. 1977. Lepidospartum burgessii

(Asteraceae, Senecioneae), a remarkable new 

gypsophilic species from Trans-Pecos Texas. 

Wrightia 5: 354-355. 
437

Leucaena leucocephala (Lam.) de Wit leadtree 

FABACEAE 

 

Synonyms: Leucaena glauca Benth. 

 Mimosa leucocephala Lam. 

 Leucaena latisiliqua (L.) Benth. 

 Mimosa glauca L. 

 Acacia glauca Willd. 

John K. Francis 

General Description.—Leadtree occurs in three 

forms or types known as the “Hawaiian” or 

common type, the “Salvador” or giant type, and the 

“Peru” type. The common type is widespread and 

shrubby and will be dealt with in this document. 

Leadtree is also known as white leadtree, leucaena, 

tantan, wild tamarind, koa haole, ipil-ipil, zarcilla, 

guaje, monval, macata, and a host of other names. 

It commonly reaches 2 to 5 m in height and 5 to 10 

cm in stem diameter at breast height. Undisturbed, 

closed stands usually are of single-stemmed plants; 

disturbed plants, especially dispersed ones, almost 

always develop multiple stems. Leadtrees are 

supported by tap and lateral root systems. They 

root deeply and extensively and form nodules when 

appropriate strains of Rhizobium bacteria are 

available. The stem bark is gray or brownish gray, 

more or less smooth with many lenticels. The 

sapwood is light yellow, and the heartwood is 

brown, hard, heavy (specific gravity 0.7), and has 

visible annual rings. The branches and twigs are 

grey-green to gray-brown and somewhat brittle in 

older shrubs. Bipinnate leaves 9 to 25 cm long are 

borne alternately. There are three to 10 pairs of 

pinnae each with 9 to 20 pairs of leaflets that are 

linear-oblong to lanceolate and 8 to 15 mm long. 

Solitary or paired inflorescences are 12- to 20-mm 

diameter, axillary heads on 2- to 3-cm-long 

peduncles. The heads contain numerous fragrant, 

greenish-white flowers. Usually four to 10 

flattened, brown legumes, 10 to 15 cm long, 

develop per inflorescence. Each legume contains 

15 to 20 hard, shiny, brown seeds that are flattened 

and tear-drop shaped. Leadtree is a polyploid with 

2n = 104 chromosomes (Brewbaker 1997, Howard 

1988, Liogier 1988, Little and Wadsworth 1964, 

Pennington and Sarukhan 1968, Stevens and others 

2001). 

Range.—Leadtree is native to southern Texas and 

the lowlands of Mexico from about 20° N. south to 

Nicaragua at about 12° N (Parrotta 2000). The 

Mexican state of Oaxaca derives its name from the 

Native American word “Uaxia,” which means the 

“place where Leucaena grows” (National Academy 

of Science 1977). Most authors treat the range in 

vague terms or state that it is uncertain. The species 

is established so completely in the West Indies and 

the tropical lowlands of Central and South America 

that apparently all adapted sites are occupied and 

the population supports many natural enemies. 

While it is assumed that leadtree was spread to 

these areas through early Spanish commercial 

activity (Parrotta 2000), earlier introductions by 

humans or animals cannot be ruled out without 

archeological evidence. From the mid-1500’s 

onward, leadtree has been spread intentionally and 

unintentionally throughout the Tropics and the 

frost-free and lightly frosted subtropics including 

Florida, Georgia, Arizona, Hawaii, and the U.S. 

Territories in the Caribbean and Pacific (Natural 

Resources Conservation Service 2002, Pacific 

Islands Ecosystems at Risk 2002). 

Ecology.—Leadtree can grow on most soils 

derived from most parent materials in a wide 

variety of site conditions. It grows fairly well on 
438 

disturbed, eroded, and partially compacted soils. 

However, the species does not grow well in poorly 

drained soils or periodically flooded soils, in soils 

with high salinity or those below pH 5.0 or above 

pH 8.0, at temperatures below 10 °C, and in areas 

with mean annual rainfall below 750 mm or above 

2500 mm (Skerman and others 1988). Leadtree is 

intolerant of shade. It does best in the open, but 

competes aggressively in low stands of weeds and 

grass and will grow in low forests with low basal 

areas. It often forms thickets that briefly eliminate 

nearly everything underneath but soon self-thin to 

fairly open stands with normal understory 

vegetation. Leadtree is drought tolerant. It folds its 

leaves under water stress preventing water loss 

and, under severe conditions, will defoliate until 

the rains return (National Academy of Sciences 

1977). 

Reproduction.—Leadtree blooms throughout the 

year (Pennington and Sarukhan 1968) and may 

begin blooming as young as 4 to 6 months of age 

(National Academy of Sciences 1977). The flowers 

are largely self-compatible and self-fertilized. It 

takes about 4 months after flowering for pods to 

mature and liberate seeds (Binggeli 1997). From 

10,000 to 21,000 seeds/kg are reported although it 

is sometimes unclear from what variety they were 

sampled (Brewbaker 1997, Parrotta 2000). A 

sample of seeds from shrubby plants from the north 

coast of Puerto Rico averaged 0.0392 ± 0.0006 

g/seed or 25,500 seeds/kg (author’s observation). 

Leadtree seeds have a hard coat that requires 

scarification for reliable germination. This can be 

done by abrasion, or treatments with hot water or 

concentrated acid. Germination of scarified seed is 

usually between 50 and 98 percent in 6 to 10 days 

after sowing. Seed can be stored for up to 5 years 

at temperatures of 2 to 6 °C (Parrotta 2000). Pods 

open while still attached to liberate their seeds and 

are dispersed by granivorous and herbivorous 

animals (Pacific Island Ecosystems at Risk 2002) 

and machinery. Detached pods are blown some 

distance by wind and occasionally retain attached 

seeds. Reproduction is common in disturbed sites 

near seed sources. Plants aggressively coppice 

when burned, cut, or broken off. The plant can be 

propagated vegetatively from cuttings (Parrotta 

2000). 

Growth and Management.—Leadtree grows 

moderately rapidly (not as fast as the giant variety 

for which most of the data are available). Seedlings 

may add about 1 m of height in the first year and 

0.5 to 0.75 m for 2 or 3 years thereafter. Diameter 

growth may vary from 2 to 10 mm per year. 

Individual shrubs live 10 to 20 years or more. 

Plantations are relatively easy to establish with 

containerized seedlings or by direct seeding 

scarified seed into prepared seedspots at the 

beginning of the wet season. Because of its 

aggressive nature, it is not advisable to introduce 

leadtree into areas where it does not already occur. 

Within the native or naturalized range, disturbed 

wildlands are usually quickly colonized by leadtree 

from the soil seedbank or from nearby seed trees. 

However, where seed is not present, planting the 

species may be an excellent way of rehabilitating 

damaged sites. Grass swards should be cultivated 

or burned or sprayed followed by cultivation before 

planting. About 5,000 to 10,000 plants/ha should 

be established for erosion control and sward 

conversion. Dense stands of leadtree needing 

control may be slashed and grazed or sprayed with 

broadleaf herbicide. 

Benefits.—Leadtree quickly invades disturbed 

areas and prevents further erosion while allowing 

succession to secondary forest cover. It is one of 

only a few species that can invade and rehabilitate 

dense tall grass swards such as Panicum maximum 

Jacq. and Imperata cylindrica (L.) Raeusch. 

(author’s observation, National Academy of 

Sciences 1977). Leadtree furnishes food and cover 

for wildlife, especially browsing animals. The 

species has been planted widely in agroforestry 

applications. It is useful for forming erosion 

barriers, windbreaks, for shade and support, green 

manure, and cut fodder. It is known to fix useful 

quantities of nitrogen (Parrotta 2000). Plantations 

are established for pasture and cut for hay. The 

foliage is highly digestible (60 to 70 percent) 

(Brewbaker 1997). Young leaves are reported to 

contain 68 calories/100 g and 80 percent moisture 

(Duke 1983). By dry weight, leaves contain 21 

percent crude protein, 18 percent crude fiber, 8 

percent ash, 6 percent fat, and 46 percent total 

digestible nutrients (FAO 2002). Cattle readily eat 

the forage but suffer symptoms of mimosine 

toxicity if given a pure diet of the species. Pigs and 

chickens must not be given more than 5 to 10 

percent in their diet, but goats have no difficulty 

with the forage, and sheep can become accustomed 

to diets rich in leadtree foliage (National Academy 

of Sciences 1977). The wood is a preferred fuel 

that burns slowly with little smoke or ash and 

makes an excellent charcoal (Brewbaker 1997). 

The stems are used for stakes and tool handles. The 

seeds are widely employed for making jewelry and 

placemats (Howard 1988). The young pods and 

green seeds are cooked as a vegetable, and the 

mature seeds are used as a coffee substitute and 
439

parched for snacks (Duke 1983). The flowers are a 

pollen source for honeybees (Little and Wadsworth 

1964). Extracts of the roots and bark are a powerful 

emetic and have been used as an abortifacient 

(Liogier 1990). 

References 

Binggeli, P. 1997. Leucaena leucocephala (Lam.) 

de Wit (Mimosaceae). http://members.lycos.co. 

uk/WoodyPlantEcology/docs.web-sp7.htm. 5 p. 

Brewbaker, J.L. 1997. Leucaena leucocephala--a 

versatile nitrogen fixing tree. Fact Sheet 97-06. 

Winrock International, Morrilton, AK. 

http://www.winrock.org/forestry/factpub/factsh/l

eucaena.htm. 5 p. 

Duke, J.A. 1983. Handbook of energy crops: 

Leucaena leucocephala (Lam.) de Wit. Center 

for New Crops & Plants Products, Purdue 

University, West Lafayette, IN. http;/www. 

hort.purdue.edu/newcrop/duke_energy/Laucaena

_leucocephala.html. 5 p. 

FAO. 2002. Sistema de información de los 

recursos del pienso: B81 Leucaena leucocephala

(Lam.) de Wit [L. glauca (L.) Benth.]. Food and 

Agriculture Organization of the United Nations, 

Rome. http://www.fao.org/livestock/agap/frg/ 

afris/espanol/document/tfeed8/Data/417.htm. 

3 p. 

Howard, R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain, 

MA: Arnold Arboretum, Harvard University. 

673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc. San Juan, PR. 566 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

National Academy of Sciences. 1977. Leucaena: 

promising forage and tree crop for the Tropics. 

National Academy of Sciences, Washington, 

DC. 115 p. 

Natural Resources Conservation Service. 2002. 

Plants profile: Leucaena leucocephala (Lam.) de 

Wit white leadtree. http://plants.usda.gov/ 

cgi_bin/plant_profile.chi?symbol=LELE10. 5 p. 

Pacific Island Ecosystems at Risk. 2002. Leucaena 

leucocephala (Lam.) de Wit, Mimosaceae. 

http://www.hear.org/pier_v3.3/leleu.htm. 3 p. 

Parrotta, J.A. 2000. Leucaena leucocephala (Lam.) 

de Wit Leucaena, tantan. In: J.K. Francis and 

C.A. Lowe, eds. Bioecología de árboles nativos 

y exóticos de Puerto Rico y las Indias 

Occidentales. General Technical Report IITF15. U.S. Department of Agriculture, Forest 

Service, International Institute of Tropical 

Forestry, Río Piedras, PR. p. 308-316. 

Pennington, T.D. and J. Sarukhan. 1968. Arboles 

tropicales de México. Instituto Nacional de 

Investigaciones Forestales, Secretaría de 

Agricultura y Ganadaría. México D.F., México. 

413 p. 

Skerman, P.J., D.G. Cameron, and F. Riveros. 

1988. Tropical forage legumes. 2nd Ed. FAO 

Plant Production and Protection Series 2. Food 

and Agriculture Organization of the United 

Nations, Rome. 692 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany, Vol. 85, No. 

2. Missouri Botanical Garden, St. Louis, MO. 

p. 945-1,910. 
440 

Lindera melissifolia (Walt.) Blume pondberry 

LAURACEAE 

Synonyms: Lindera melissaefolium (Walt.) Blume 

 Benzoin melissaefolium (Walt.) Nees 

 Laurus melissaefolia Walt. 

Margaret S. Devall and Nathan M. Schiff 

General Description.—Pondberry, also known as 

southern spicebush (Morgan 1983), a member of 

the Lauraceae, is a stoloniferous, deciduous, 

aromatic shrub up to 2 m tall. Pondberry usually 

occurs in clones of numerous stems with erect or 

ascending shoots and few branches. The alternate 

drooping leaves are subcordate with prominent 

venation and pubescence on the lower surface 

(Klomps 1980). Lindera benzoin var. pubescens 

also is pubescent beneath. The species is 

dioecious, with small yellow flowers that bloom in 

spring before leaf-out. The fruit is a red drupe 

about 1 cm long that matures in late summer or 

fall. Female clones are often smaller than male 

clones and are sometimes absent from stands 

(Wright 1989, 1990). As in many clonal species, 

seedlings are rarely observed (Wright 1990, Devall 

and others 2001). 

Range.—Pondberry occurs in the Southeastern 

United States. At present there are populations in 

Arkansas, Georgia, Mississippi, Missouri, North 

Carolina and South Carolina; it has apparently 

been extirpated from Alabama and Louisiana and 

possibly Florida. 

Ecology.—Pondberry has probably always been a 

rare species (Radford and others 1968, Kral 1983), 

and knowledge of its ecology is limited. The 

species occurs in lowland habitats with hydric 

soils (Morgan 1983), in areas that are usually 

flooded in winter. In Mississippi pondberry occurs 

in bottomland hardwood forests. In northeastern 

Arkansas and southeastern Missouri pondberry is 

found on the bottoms and edges of shallow 

seasonal ponds in old dune fields, but in 

southeastern Arkansas it occurs in low habitat 

along a river. In South Carolina the species occurs 

in areas with karst topography, around the edges of 

sinkholes, and in Georgia it occurs along the 

borders of sphagnum bogs. Ambient light at the 

different sites ranges from deep shade to almost 

full sun. It appears that pondberry can occupy 

different habitats as long as its requirements for 

water are met (Devall and others 2001). The 

distribution and abundance of pondberry have 

been affected by habitat destruction and alteration, 

such as timber cutting and clearing of land. The 

species was listed as endangered by the U.S. Fish 

and Wildlife Service in 1986 (U.S. Fish and 

Wildlife Service 1986). Many of the existing 

colonies are small, and occupy only a portion of 

the apparently suitable habitat. Although the 

pondberry recovery plan states that there are 36 

extant populations (U.S. Fish and Wildlife Service 

1993), new colonies have been discovered since 

1993, some in new locations and some near known 

populations. Some populations that were thought 

to be separated by enough distance to preclude 

interbreeding (as on the Delta National Forest, 

MS) may be linked by recently discovered 

colonies. 

Reproduction.—Stems flower in the second to 

fourth year of growth (U.S. Fish and Wildlife 

Service 1993). The flowers are pollinated by 

several species of small bees and are visited by 
441

numerous insect species. Flower and fruit 

production are highly variable (Morgan 1983, 

Tucker 1984). Some years the flowers are 

damaged by late frosts, while other years flower 

and fruit production may be heavy, with up to 100 

fruits per plant. 

Growth and Management.—Seedlings are rarely 

observed (Devall and others 2001). Stems usually 

live 6 or 7 years, and when a stem dies it is almost 

always replaced by a new stem that grows from 

the base of the plant. Clones expand vegetatively, 

and a mature colony often includes numerous leafy 

stems along with some dead stems (U.S. Fish and 

Wildlife Service 1993). Most pondberry colonies 

occur in light shade, but a few grow in almost full 

sun. In unshaded conditions, competition may be a 

problem. More knowledge is needed about 

management, and studies are currently being 

conducted on the Delta National Forest in 

Mississippi and at other locations. The U.S. Fish 

and Wildlife Service (1993) states that pondberry 

should be protected from forestry and agricultural 

management actions and protected from grazing 

and browsing animals. 

Benefits.—The fruits are eaten by hermit thrushes 

(Catharus guttatus), cardinals (Cardinalis 

cardinalis) and other birds (Smith and others 

2003). Steyermark (1949) reported that the fruits 

are used as ammunition in pop guns, tubular 

contrivances constructed from elderberry 

[Sambucus nigra L. ssp. canadensis (L.) R. Bolli] 

twigs. 

References 

Devall, M., N. Schiff, and D. Boyette. 2001. 

Ecology and reproductive biology of the 

endangered pondberry, (Lindera melissifolia

(Walt) Blume. Natural Areas Journal 21: 250-

258.

Klomps, V.L. 1980. The status of Lindera 

melissifolia (Walt.) Blume, pondberry, in 

Missouri. Transactions of the Missouri Academy 

of Science 14: 61-66. 

Kral, R. 1983. A report on some rare, threatened, 

or endangered forest-related vascular plants of 

the South: Volume 1, Isoetaceae through 

Euphorbiaceae. Technical Publication R8-TP 2. 

U.S. Department of Agriculture, Forest Service, 

Southern Region, Atlanta, GA. 718 p. 

Morgan, S. 1983. Lindera melissifolium: A rare 

southeastern shrub. Natural Areas Journal 3: 62-

67. 

Radford, A.E., H.E. Ahles, and C.R. Bell. 1968. 

Manual of the Vascular Flora of the Carolinas. 

University of North Carolina, Chapel Hill, NC. 

1,183 p. 

 

Smith, C.G. III, P.B. Hamel, M.S. Devall and 

N.M. Schiff. 2003. Hermit thrush is the first 

observed dispersal agent for pondberry. 

Castanea, in press. 

Steyermark, J.A. 1949. Lindera melissifolia.

Rhodora 51: 153-162. 

Tucker, G.E. 1984. Status report on Lindera 

melissifolia (Walt) Blume. Provided under 

contract to the U.S. Fish and Wildlife Service, 

Southeast Region, Atlanta, GA. 41 p. 

U.S. Fish and Wildlife Service. 1986. Endangered 

and threatened wildlife and plants; 

determination of endangered status for Lindera 

melissifolia. Federal Register 51: 27,495-27,500. 

U.S. Fish and Wildlife Service. 1993. Recovery 

plan for pondberry (Lindera melissifolia). U.S. 

Fish and Wildlife Service. Atlanta, GA. 56 p. 

Wright, R.D. 1989. Reproduction of Lindera 

melissifolia in Arkansas. Proceedings of the 

Arkansas Academy of Science 43: 69-70. 

Wright, R.D. 1990. Photosynthetic competence of 

an endangered shrub, Lindera melissifolia. 

Proceedings of the Arkansas Academy of 

Science 44: 118-120. 
442 

Lonicera japonica Thunb. Japanese honeysuckle 

CAPRIFOLIACEAE 

Synonyms: Nintooa japonica (Thunb.) Sweet 

John K. Francis 

General Description.—Japanese honeysuckle, 

also known as woodbine and Chinese 

honeysuckle, is a semi-evergreen woody vine that 

forms mounds and mats on open ground and by 

twining, climbs shrubs, low trees, windfalls, and 

fences. The ropy stems, which have gray-brown, 

fissured to shredding bark, can extend 5.5 m or 

more and reach 10 cm in diameter in old plants. 

The wood is white, soft, and fine-grained. Young 

stems are reddish brown to light brown and about 

3 mm in diameter with hollow pith. Roots are 

extensive, reaching 2.5 m laterally and 15 to 100 

cm in depth. Opposite, light-green leaves are 2.5 to 

12 cm long, ovate to oblong, entire to lobbed, with 

short petioles. The leaves are evergreen in tropical 

and subtropical settings and become late deciduous 

to deciduous in cooler temperate climates. The 

paired axillary flowers are 2 to 3 cm long, white 

(sometimes tinged with pink or purple) turning 

yellow, tubular with five lobes, fused in two 

unequal lips. The fruits are black, glossy, globose 

berries 4 to 6 mm in diameter containing two to 10 

black seeds (Miller 1999, Munger 2002, Stephens 

1973). Japanese honeysuckle is a diploid plant 

with 2n = 18 chromosomes (Schierenbeck and 

others 1995). 

Range.—Japanese honeysuckle is native to Japan, 

Korea, China, and Taiwan (Bravo 2003, Pacific 

Island Ecosystems at Risk 2003). The species has 

naturalized in much of the United States except 

Alaska and a few states in the Northwest, northern 

Midwest, and Vermont (Natural Resources 

Conservation Service 2003). It has also naturalized 

in Puerto Rico (Liogier and Martorell 1982), 

Ontario, Canada (Munger 2002), Juan Fernandez 

archipelago, Chile (Sharma and others 1997), 

Australia, New Zealand, Christmas Island (Pacific 

Island Ecosystems at Risk 2003), England, 

Portugal, Corsica, Brazil, and Argentina (Nuzzo 

2003). There are two generally accepted varieties, 

chinensis and hilliana. The latter is the variety 

commonly encountered in naturalized populations 

(Nuzzo 2003). 

Ecology.—Japanese honeysuckle grows well in a 

wide variety of soils with a few exceptions such as 

coarse sands and poor peat soils (Munger 2002). It 

invades fencerows, roadsides, vacant lots, forest 

openings, understories of open forest stands, and 

abandoned pastures and fields. The species is 

present in nearly all the forest associations in the 

Southeastern U.S., the region where it is most 

common. Japanese honeysuckle is adapted to 

climbing small-diameter stems and sometimes 

smothers or strangles small trees. It cannot ascend 

trees with trunks larger than 15 cm in diameter 

(Nuzzo 2003). It grows best under full sunlight but 

can still survive with 25 percent of sunlight in a 

forest understory. Retaining leaves and 

photosynthesizing through all or part of the winter 

after canopy forest trees have defoliated helps 

plants survive through the summer in low-light 

conditions (Nuzzo 2003). Northern distribution of 

naturalized Japanese honeysuckle is limited by 

short growing seasons and late spring frosts, and is 

generally south of an isotherm where mean 

January temperature is -1 °C. It has not generally 

naturalized west of the 1000 mm mean annual 
443

precipitation limit (except where exogenous water 

is available), roughly corresponding with native 

hardwood forests. Japanese honeysuckle survives 

fire by sprouting from stem tissue buried under 

litter and mineral soil. It generally disappears from 

communities (such as longleaf pine) where fires 

are frequent (Munger 2002). 

Reproduction.—Japanese honeysuckle in the U.S. 

flowers from April through July (sometimes 

through October) varying by local climate (Nuzzo 

2003). The flowers open in late afternoon and 

remain open for about 3 days (Nuzzo 2003). The 

flowers are pollinated by hummingbirds and 

insects (nocturnal hawkmoths and diurnal bees). 

The species requires pollen from genetically 

distinct individuals for successful pollination and 

as a result is pollinator limited. Naturallypollinated flowers produced seeds in 17.4 percent 

of the resulting fruits while hand-pollinated 

flowers produced seeds in 78.7 percent of the 

fruits (Larson 2001). The seeds are dispersed 

primarily by birds. Germination of most seeds 

appears to occur in the spring following dispersal 

and occurs at similar rates in mineral soil and 

under leaf litter (Munger 2002). Japanese 

honeysuckle regenerates vegetatively by sprouting 

from underground stems and by the layering of 

above-ground stems that come in contact with the 

soil. Discarded prunings will also take root. 

Growth and Management.—Seedlings grow 

slowly for the first 2 years but afterwards may 

extend their stems 1.5 m/year (Nuzzo 2003). The 

species begins bearing fruit at age 3 in full sun and 

at age 5 when growing in the shade. Fruit 

production generally peaks between ages 4 and 6 

(Munger 2002). By fertilization, researchers were 

able to increase browse produced by Japanese 

honeysuckle by over 2,000 kg/ha and increase 

crude protein as well (Dyess and others 1994). The 

species is easy to propagate with cuttings and most 

ornamental plants are multiplied in this fashion. 

The greater challenge is to eliminate infestations 

of Japanese honeysuckle. Pulling and grubbing 

individual plants and small infestations can be 

effective but will require repeat visits to eliminate 

missed sprouts and new seedlings. Tethered goats 

can provide effective control for spot infestations 

(Bravo 2003). Over-the-top spraying with 

glyphosate, Dichlorprep + 2,4-D, picloram + 2,4-

D, tebuthiuron, sulfometuron, or triclopyr will kill 

it (Nuzzo 2003) but also tend to eliminate most 

other plant species as well. A more selective 

method is to cut stems and apply the above 

mentioned herbicides to the cut surfaces (Bravo 

2003). Mowing is ineffective (Nuzzo 2003). 

Benefits and Detriments.—In spite of the 

invasive and weedy tendencies of the species, 

Japanese honeysuckle is still widely used as an 

ornamental. It is especially desirable in arid areas 

where it has little tendency to spread. Managed for 

screens and ground cover, Japanese honeysuckle is 

beautiful in and out of flower and especially 

captivating because of the strong and pleasant 

aroma of its blooms. Some gardeners favor it 

because it attracts hummingbirds. The fruits are 

toxic to humans if large quantities are eaten and 

result in vomiting, diarrhea, rapid heartbeat, 

respiratory failure, and coma. However, children 

suck the nectar from flowers without ill effects 

(Russell 1997). The species was formerly planted 

for erosion control and wildlife benefits. Planting 

is currently discouraged or prohibited because the 

species has dominated large areas and threatens 

sensitive native species (Nuzzo 2003). Japanese 

honeysuckle is important wildlife cover and an 

important, sometimes the most important, browse 

species for white tailed deer in Eastern and 

Southern United States. It is also eaten by 

cottontail rabbits, cattle, sheep, and goats. The 

leaves yield 8 to 12 percent crude protein and have 

an in vivo dry matter digestibility of 35 to 70 

percent. Wild turkeys, northern bobwhite, and a 

number of songbirds eat the fruits (Munger 2002). 

Japanese honeysuckle tissues contain anticomplementary polysaccharides and polyphenolic 

compounds. Used in herbal medicine, extracts are 

reported to protect against cellular injury, help 

maintain vascular health, and prevent infections 

(Nuzzo 2003) 

References

Bravo, M.A. 2003. Japanese honeysuckle, 

Lonicera japonica Thunb. Alien Plant Working 

Group, National Park Service, Washington, DC. 

http://www.nps.gov/plants/alien/fact/loja1.htm. 

6 p. 

Dyess, J.G., M.K. Causey, and H.L. Stribling. 

1994. Effects of fertilization on production and 

quality of Japanese honeysuckle. Southern 

Journal of Applied Forestry 18(2): 68-71. 

Larson, K.C. 2001. Lack of pollinators limits fruit 

set in the exotic Lonicera japonica. Botanical 

Society of America, Botany 2001 Abstracts. 

p. 39. 
444 

Liogier, H.A. and L.F. Martorell. 1982. Flora of 

Puerto Rico and adjacent islands: a systematic 

synopsis. Editorial de la Universidad de Puerto 

Rico, Río Piedras, PR. 342 p. 

Miller, J.H. 1999. Japanese honeysuckle, Lonicera 

japonica. Protection Report RX-PR 036. U.S. 

Department of Agriculture, Forest Service, 

Region 8, Atlanta, GA. 2 p. 

Munger, G.T. 2002. Lonicera japonica. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/vine/ 

lonjap/all.html. 43 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Lonicera japonica Thunb. 

http://plants.usda.gov/cgi_bin/plant_search.cgi?

mode=Scientific+Name&keywordquery+Lonice

ra+japonica&earl=plant_search.cgi. 6 p. 

Nuzzo, V. 2003. Element stewardship abstract for 

Lonicera japonica, Japanese honeysuckle. The 

Nature Conservancy, Arlington, VA. 

http://tncweeds.ucdavis.edu/esadocs/documnts/ 

lonijap.html. 23 p. 

Pacific Island Ecosystems at Risk. 2003. Lonicera 

japonica Thunb., Caprifoliaceae. http://www. 

hear.org/pier/lojap.htm. 3 p. 

Russell, A.B. 1997. Poisonous plants of North 

Carolina, scientific name: Lonicera japonica.

North Carolina State University, Charlotte, NC. 

http://www.ces.ncsu.edu/depts/hort/consumer/ 

poison/Lonicja.htm. 2 p. 

Schierenbeck, K.A., J.L. Hamricks, and R.N. 

Mark. 1995. Comparison of allozyme variability 

in a native and an introduced species of 

Lonicera. Heredity 75(1): 1-9. 

Sharma, A., A. Murayama, T. Osaki, K. Ooi, T. 

Yahara, M. Ikegami, Silva-O., M. 1997. Plant 

conservation in the Juan Fernandez archipelago, 

Chile. Aliso 16(2): 89-101. 

Stephens, H.A. 1973. Woody plants of the North 

Central Plains. The University Press of Kansas, 

Laurence, KS. 530 p. 
445

Lotus scoparius (Nutt.) Ottley California broom 

FABACEAE 

Synonyms: Hosackia scoparia Nutt. in T. & G. 

 Syrmatium galbrum Vogel 

 Hosackia glaber Greene 

 Hosackia crassifolia Nutt., not Benth 

 Lotus glaber Greene, not Mill. 

Arlee M. Montalvo 

General Description.—California broom is a 

diverse and widely distributed subshrub (Munz 

and Keck 1968, Isely 1981, Hickman 1993). The 

Latin name “scoparius” refers to the broom-like 

form which is also evident in the common name 

“California broom.” The other common name, 

“deerweed,” refers to its use as browse by deer. Of 

five previously recognized subspecific taxa (Ottley 

1923, Munz and Keck 1968), the three from the 

Channel Islands were reassigned to L. dendroideus

(Greene) Greene by Isley (1981). His treatment of 

the genus recognizes two varieties of Lotus 

scoparius (L. s. var. scoparius and L. s. var. 

brevialatus Ottley) and has been adopted in 

Hickman (1993). In the former, the keel of the 

corolla is about as long as the wings and the calyx 

has broad triangular teeth (figure: top left and 

bottom left). In the latter, the keel is longer than 

the shortened wings and the calyx has narrow teeth 

(figure: top center and bottom right). The erect 

suffrutescent shrubs grow from 0.4 to 1.5 m tall 

but prostrate forms often occur along the coastal 

strand. A spreading, prostrate ecotype occurs on 

coastal sand dunes at Monterey Bay. The many 

flexible green branches arise from a woody base, 

and are mostly glabrous except toward tips. The 

soft, alternate leaves are pinnately compound with 

three (occasionally four to five) oblong to 

oblanceolate leaflets, usually 4 to 10 mm long with 

short, sparse, appressed hairs, and are subtended 

by small, gland-like stipules. The sessile, usually 

two- to seven-flowered, umbellate inflorescences 

are in the axils of leaves. The yellow pea-like 

flowers are 7 to 10 (occasionally to 15) mm long, 

have short pedicels, and sometimes bear a red 

splotch of varying intensity on the back of the 

banner. Depending on the plant, flowers fade to 

dark orange or red following pollination. Plants 

bear hundreds of inflorescences and are an 

attractive flag for native bees. The small 

indehiscent pods (figure: top right) bear one to two 

(rarely three) seeds that vary from brown to 

greenish brown with or without dark brown 

mottling. Seeds of var. scoparius are narrow beanshaped, 1.2 - 2mm long. Seeds of var. brevialatus

are narrower (ca. 3 x longer than wide). Both 

varieties have 2n = 14 chromosomes (Munz and 

Keck 1968, Grant 1995). 

Range.—California broom occurs primarily after 

fire or in open areas of coastal sage scrub, 

chaparral, desert scrub, washes, coastal strand, or 

along roadsides. The var. scoparius is distributed 

in cismontane California below 1,500 m from 

Humboldt and Plumas Counties, south into Baja 

California, primarily in shrublands and associated 

open areas, and primarily in the coastal regions of 

California and north of Los Angeles. In addition to 

the distinct differences in floral form between the 

two varieties, there is variation in floral form 

within var. scoparius that is associated with 

coastal verses inland habitats from the Transverse 

Range of California, northward (author’s 
446 

observation). Variety brevialatus occurs from Los 

Angeles Co., south into Baja California primarily 

in the hotter and drier interior regions of Riverside, 

Los Angeles, western San Bernardino, and eastern 

San Diego Counties. Although the two varieties 

have somewhat different geographic distributions 

and are easily distinguished by floral characters 

(Isely 1981, Steppan 1991), they overlap and 

hybridize in some areas of Los Angeles, Riverside, 

and San Diego Counties. Putative hybrids have 

been observed in contact zones between the two 

varieties (Isely 1981, Steppan 1991, Montalvo and 

Ellstrand 2001). Flowers of synthetic F1 hybrids 

are somewhat intermediate, but have wings as long 

as the keel. Many areas of overlap may be due to 

natural secondary contact, but some are clearly due 

to seeding projects along highways and utility 

corridors. The various floral forms are maintained 

when plants are grown from seed in a common 

environment (Montalvo and Ellstrand 2000, 2001). 

Ecology.—In nature, seeds typically germinate 

after scarification by fire but some germinate in 

open, disturbed sites (Munz and Keck 1968, 

Keeley and Keeley 1984). Between 2 to 3 years 

after fire in sage scrub vegetation, following a 

burst of herbaceous species, California broom can 

become the dominant canopy species, eventually 

becoming replaced by long-lived shrub species. Its 

abundance gradually decreases in 5 to 10 years 

after fire. The branched tap root is relatively 

shallow (< 1 m), and roots form symbiotic 

associations with nitrogen fixing bacteria and 

arbuscular mycorrhizal fungi (author’s 

observation). Plants are facultatively droughtdeciduous, a trait commonly associated with 

shallow-rooted shrubs of coastal sage scrub 

vegetation in California. Seasonality of leaf 

production, nutrient accumulation, and leaf drop in 

response to summer drought has been studied 

extensively in L. scoparius var. scoparius (Nilsen 

and Muller 1980, 1981a,b, 1982, Nilsen and 

Schlesinger 1981, Nilsen 1982). Other researchers 

have examined response to photoperiod 

(Comstock and Ehleringer 1986). 

Reproduction.—Seeds of California broom 

germinate in mid to late winter during the rainy 

season. Plants establish quickly with normal 

rainfall and typically reach flowering size the 

second year. Flowering occurs primarily from 

March to June but may start as early as January in 

warm, wet winters, and last much longer in more 

moist, coastal areas and in years with long-lasting 

soil moisture. Flowers are self-compatible and 

insect-pollinated (Moldenke 1976, Hickman 1993, 

Jones and Cruzan 1999) primarily by native bees 

in the genera Bombus, Hoplitus, Anthophora, 

Habropoda, Osmia, and Anthidium, but flowers 

are also visited by butterflies and non-native 

honeybees (author’s observation, Jones and 

Cruzan 1999). The indehiscent pods ripen in about 

4 to 6 weeks and are primarily passively dispersed 

short distances. The hard seeds require heat or 

mechanical scarification to break dormancy. Of 

several treatments including control, soil heated to 

100 o

C for 1 hr, ash /chemical fertilizer, and heat 

plus fertilizer, Christensen and Muller (1975) 

found that heat treatment yielded the highest 

germination. In addition, Keeley (1987) found that 

heating seeds in their pods to 120 o

C for 5 minutes 

increased germination over that of unheated 

controls. 

Genetics.—There are significant genetic 

differences among populations of the two varieties 

of California broom (Montalvo and Ellstrand 

2000, 2001). An analysis of genetic marker data 

(13 allozyme loci) from three populations of var. 

brevialatus and nine populations of var. scoparius

showed significant population substructure due 

primarily to differences among populations of the 

two varieties. In an analysis of all populations, 18 

percent of the variation was due to differences 

among populations, while analysis of just var. 

brevialatus or var. scoparius populations showed 

only 1 and 8 percent of the variation due to 

differences among populations. Thus, populations 

within a variety are substantially more genetically 

similar to each other than to populations of the 

other variety. Inbreeding coefficients were low for 

all populations, a pattern consistent with 

substantial cross pollination, severe loss of inbred 

progeny, or both (mean f = 0.09, range = 0.00 – 

0.18) (Montalvo, Clegg, and Ellstrand, manuscript 

in preparation). 

There is also genetically based geographic 

variation in floral form in this species. In San 

Diego Co., Steppan (1991) detected distinct 

discontinuities in floral morphology between 

varieties and moderate correlations among 

environmental variables and floral traits of wild 

populations. Montalvo and Weaver (unpublished 

data) measured floral traits on 12 populations of 

plants raised in a common environment and made 

pairwise comparisons of genetic, floral, 

geographic, and environmental distances of source 

populations. Floral morphological distance 

correlated with environmental distance, genetic 

distance correlated with floral distance, but genetic 

distance did not correlate with geographic distance 
447

until var. brevialatus was removed from the 

analysis. Results were consistent with floral form 

having a genetic basis and with environmental 

factors playing a role in the evolution of floral 

divergence. 

The substantial genetic differentiation of 

populations has implications for translocation of 

California broom for restoration. Common garden 

experiments testing plants originating from seed 

collected from 12 source populations (both 

varieties represented) demonstrated a significant 

home site advantage (Montalvo and Ellstrand 

2000). Furthermore, Montalvo and Ellstrand 

(2001) directly tested the potential for 

“outbreeding depression,” a loss of fitness upon 

crossing genetically differentiated populations, by

crossing individuals from six populations of the 

two varieties in every combination and testing the 

progeny in two common gardens at wild sites. 

Seeds per flower and seedling emergence 

decreased significantly with an increase in genetic 

distance of the crossed parental populations. 

Among variety crosses were only 70 percent as fit 

as within variety crosses by the time seedlings 

emerged, and further fitness differences 

accumulated after seedlings were outplanted into 

field plots. In these common gardens, success of 

progeny decreased with increasing differences 

between parental environments and the transplant 

location. Results indicate that careful attention 

should be given to the similarity of taxonomic 

position and environment of source populations 

relative to planting location, especially when 

choosing source populations for restoration, 

mitigation, and roadside landscaping. 

Interspecific Hybridization.—Putative hybrids 

have been reported between California broom and 

L. junceus (Benth.) Greene and L. benthamii Green 

in central and northern California, as well as 

between other members of the species complex 

(Isely 1981), but there have not been any genetic 

studies confirming introgression. Liston and others 

(1990), however, did genetic studies on San 

Clemente Is. and documented hybridization 

between the rare L. dendroideus (Greene) Greene 

var. traskiae (Nodden) Isely, formally L. scoparius

ssp. traskiae (Noddin) Raven, and the more 

widespread L. argophyllus (A. Gray) E. Greene 

var. ornithopus (E. Greene) Ottley. They 

concluded that genetic assimilation of the rare 

species by the widespread species is possible. 

Growth and Management.—The small pods of 

California broom can be collected from May to 

July depending on location. Pods or cleaned seeds 

can be covered with boiling water and left to soak 

to break dormancy (Atwater 1980, Young and 

Young 1986, Emery 1988, Montalvo and Ellstrand 

2000, 2001), or exposed to dry heat (Keeley 1987). 

Whole pods can be dry or wet broadcast in the fall 

for revegetation and restoration. Seedlings plugs 

can also be used for small projects. In greenhouse 

plantings, seeds germinate best when planted less 

than 2 mm deep (author’s observation). Despite 

the strong geographic and morphological 

differences, the two varieties have been used 

indiscriminately in many seeding projects. Due to 

the adaptive differences and the observed 

outbreeding depression following hybridization, 

land managers should use the variety native to the 

planting location. 

Benefits.—California broom is an important, fast 

growing, early successional species used 

extensively in erosion control, post-fire mitigation, 

and habitat restoration in California, especially in 

the rapidly disappearing coastal sage scrub. The 

flowers and seeds are an important food resource 

for a variety of insects and seed foraging rodents 

and birds (Duncan 1968). The plants provide 

valuable forage for deer, especially in drought 

years when growth of herbaceous vegetation is 

sparse (Dale 2000). 

References 

Atwater, B.R. 1980. Germination, dormancy and 

morphology of the seeds of herbaceous 

ornamental plants. Seed Science and Technology 

8: 523-573. 

Christensen, N.L. and C.H. Muller. 1975. Relative 

importance of factors controlling germination and 

seedling survival in Adenostoma chaparral. The 

American Midland Naturalist 93: 71-78. 

Comstock, J. and J.R. Ehleringer. 1986. 

Photoperiod and photosynthetic capacity in Lotus

scoparius. Plant, Cell and Environment 9: 609-

612. 

Dale, N. 2000. Flowering Plants: The Santa 

Monica Mountains, Coastal and Chaparral 

Regions of Southern California. California Native 

Plant Society, Sacramento, CA. 240 p. 

Duncan, D.A. 1968. Food of California quail on 

burned and unburned central California foothill 

rangeland. California Fish and Game 54: 123-127. 

Emery, D.E. 1988. Seed propagation of native 

California Plants. Santa Barbara Botanical 

Garden, Santa Barbara, CA. 23 p. 
448 

Grant, W.F. 1995. A chromosome atlas and 

interspecific-intergenic index for Lotus and 

Tetragonolobus (Fabaceae). Canadian Journal of 

Botany 73: 1,787-1,809. 

Hickman, J.C., ed. 1993. The Jepson Manual: 

Higher Plants of California. University of 

California Press, Ltd., Berkeley, CA. 1,400 p. 

Isely, D. 1981. Leguminosae of the United States III 

subfamily Papilionoidae: tribes Sophoreae, 

Podalyrieae, Loteae. In: Memoirs of the New 

York Botanical Garden. Volume 25. The New 

York Botanical Garden, Bronx, NY. p. 127-191. 

Jones, C.E. and M.B. Cruzan. 1999. Floral 

morphological changes and reproductive success 

in deer weed (Lotus scoparius; Fabaceae). 

American Journal of Botany 86: 273-277. 

Keeley, J.E. 1987. Role of fire in seed germination 

of woody taxa in California chaparral. Ecology 

68:434-443. 

Keeley, J.E. and S.C. Keeley. 1984. Postfire 

recovery of California coastal sage scrub. The 

American Midland Naturalist 111: 105-117. 

Liston, A., L.H. Rieseberg, and O. Mistretta. 1990. 

Ribosomal DNA evidence for hybridization 

between island endemic species of Lotus. 

Biochemical Systematics and Ecology 18: 239-

244. 

Moldenke, A.R. 1976. California pollination 

ecology and vegetation types. Phytologia 34: 305-

361. 

Montalvo, A.M. and N.C. Ellstrand. 2000. 

Transplantation of the subshrub Lotus scoparius: 

testing the home site advantage hypothesis. 

Conservation Biology 14: 1034-1045. 

Montalvo, A.M. and N.C. Ellstrand. 2001. Nonlocal transplantation and outbreeding depression 

in the subshrub Lotus scoparius (Fabaceae). 

American Journal of Botany 88: 258-259.

Munz, P.A. and D.D. Keck. 1968. A California 

Flora with Supplement. University of California 

Press, Berkeley, CA. 1,681 + 224 p. 

Nilsen, E.T. 1982. Productivity and nutrient cycling 

in the early postburn chaparral species Lotus

scoparius. In: C. E. Conrad and W. C. Oechel, 

eds. General Technical Report PSW-58. Pacific 

Southwest Forest and Range Experiment Station, 

USDA Forest Service, Berkeley, CA. p. 291-296. 

Nilsen, E.T. and W.H. Muller. 1980. An evaluation 

of summer deciduosness in Lotus scoparius

(Nutt.) T.& G. The American Midland Naturalist 

103: 88-95. 

Nilsen, E.T. and W.H. Muller. 1981a. The influence 

of low plant water potential on the growth and 

nitrogen metabolism of the native California fire 

response shrub Lotus scoparius. American Journal 

of Botany 68: 402-407. 

Nilsen, E.T. and W.H. Muller. 1981b. Phenology of 

the drought-deciduous shrub Lotus scoparius: 

climatic controls and adaptive significance. 

Ecological Monographs 51: 323-341. 

Nilsen, E.T. and W.H. Muller. 1982. The influence 

of photoperiod on drought induction of dormancy 

in Lotus scoparius. Oecologia 53: 79-83. 

Nilsen, E.T. and W.H. Schlesinger. 1981. 

Phenology, productivity, and nutrient 

accumulation in the post-fire chaparral shrub 

Lotus scoparius. Oecologia 50: 217-224. 

Ottley, A.M. 1923. A revision of the Californian 

Species of Lotus. University of California 

Publication in Botany, Berkeley, CA. 10: 189-

305. 

Steppan, S.J. 1991. Geographic distribution of 

flower morphological traits in subspecies of Lotus

scoparius. Journal of Biogeography 18: 321-331. 

Young, J.A. and C.G. Young. 1986. Collecting, 

Processing, and Germinating Seeds of Wildland 

Plants. Timber Press, Portland, OR. 236 p. 
449

Ludwigia octovalvis (Jacq.) Raven primrose willow 

ONAGRACEAE 

Synonyms: Oenothea octovalvis Jacq. 

Jussiaea suffruticosa L. 

Jussiaea pubescens L. 

Jussiaea octovalvis (Jacq.) Sw. 

Jussiaea angustifolia Lam. 

(many others Howard 1989, Liogier 1995) 

John K. Francis 

 

General Description.—Primrose willow, also 

known as Jamaica loostrife, wild clove, many-seed, 

kamole, cangá, yerba de clavo, girofle-ma, grand 

giroflé, and manger mouton, is a suffruticose shrub 

or perennial woody herb to 2 m in height and 1 cm 

in stem diameter. The stems of older plants are 

woody through most of their height; branches 

become increasingly succulent toward the tips. 

Only the lower parts of young plants are woody. 

Primrose willow plants are supported by “sinker” 

roots, one of which may be the taproot, and long 

white, lateral roots that grow just under the soil 

surface. These roots have a hard central core and 

are surrounded by a cottony mass of root hairs. In 

the first year, the stem produces a number of fine 

lateral branches. In the next one or more years, 

thelateral branches each produce similar secondary 

or tertiary branches. The alternate leaves are linear 

to narrowly ovate with blades 1.3 to 16.2 cm long. 

The species is highly variable in leaf size, shape, 

and pubescence. Primrose willow flowers are 

solitary in the upper leaf axils. Four sepals form a 

peg-like hypanthium, 5 to 12 mm long, from which 

emerge four yellow petals. The twigs, leaf veins, 

and hypanthium are often tinged in red. The 

capsules, which have a clove-like shape, are 2.5 to 

5 cm long and contain many rounded, black seeds. 

Chromosome number for the species is 2n = 32, 48 

(Howard 1989, Jones 1975, Liogier 1995, Long 

and Lakela 1976). 

Range.—The original range of primrose willow is 

unknown. It grows today from Southern United 

States through the West Indies, and from Mexico 

through Central and South America to at least 

Bolivia (Hodgdon Herberium 2002, Natural 

Resources Conservation Service 2002). It has 

naturalized in Hawaii (Fish and Wildlife 

Information Exchange 2002) and grows in tropical 

and subtropical regions almost worldwide (Burkill 

1997, Howard 1989, Liogier 1995). 

Ecology.—Primrose willow grows in wet soils 

and shallow water, such as drainage ditches, 

borrow pits, sloughs, marshes, ponds, seasonally 

flooded bottoms, river banks, and rice paddies. 

Soil type within those sites is apparently not very 

important. Primrose willow grows from near sea 

level to at least 800 m in elevation (Missouri 

Botanical Garden 2002). It prefers full sun and 

tolerates no more than side shade. The species 

grows singly or in clumps. 

Reproduction.—After primrose willow plants in 

the tropics reach about 0.4 m, blooming and 

fruiting is continuous (author’s observation, Long 
450 

and Lakela 1976). In Texas, it is reported to bloom 

from July through October (Correll and Johnston 

1970). Seeds collected in Puerto Rico averaged 54 

million per kg. Placed on moist filter paper, 28 

percent of them germinated within 14 days, most 

on the seventh day (author’s observation). Local 

dispersal is by wind and water. Long-distance 

dispersal is probably accidental on birds, 

machinery, or materials being shipped. 

Growth and Management.—Primrose willow 

grows up to 1 m/year. Under favorable conditions, 

plants live at least 3 years and possibly more. 

There seems to be little reason to plant the species. 

Control may be necessary in drainage ditches to 

maintain flow and in rice paddies. Control can be 

achieved by cultivation, hand pulling, and by 

applying commercial weed killers. 

Benefits and Detrements.—Primrose willow 

grows in wet areas where few other plants will 

grow and as such it aids in the successional 

process. It furnishes cover for wildlife. Its yellow 

flowers add a splash of color to areas often devoid 

of colorfully flowering plants. In herbal medicines, 

preparations of the leaves are used to treat diarrhea, 

dysentery, as a laxative and a vermifuge, and to 

relieve headache, chest pains, and rheumatoid pain 

(Burkill 1997, Liogier 1990, Parrotta 2001). A test 

of a local crude drug from Taiwan made from 

primrose willow effectively inhibited the 

cariogenic bacteria Streptococcua mutans (Chen 

and others 1989). The plant is a major weed in rice 

in the tropics and sometimes chokes drainage 

ditches and irrigation canals (Burkill 1997). 

References 

Burkill, H.M. 1997. The useful plants of West 

Tropical Africa. Vol. 4. Royal Botanic Gardens, 

Kew, UK. 969 p. 

Chen, C.P., C.C. Lin, and T. Namba. 1989. 

Screening of Taiwanese crude drugs for 

antibacterial activity against Streptococcus 

mutans. Journal of Ethnopharmacology 27(3): 

285-295. 

Correll, D.S. and M.C. Johnston. 1970. Manual of 

the vascular plants of Texas. Texas Research 

Foundation, Renner, TX. 1,881 p. 

Fish and Wildlife Information Exchange. 2002. 

Species moorhen, common, Hawaiian. 

http://fwie.vt.edu/WWW/esis/lists/e101014.htm. 

17 p. 

Hodgdon Herbarium. 2002. Dicotyledons 

associated with Bolivian wetlands. 

http://unh.edu/herbarium/bolivia/dicots.htm. 5 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Jones, F.B. 1975. Flora of the Texas Costal Bend. 

Mission Press, Corpus Christi, TX. 262 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de 

la Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Liogier, H.A. 1990. Plantas mediciales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Missouri Botanical Garden. 2002. Manual de la 

flora de Costa Rica. http://www.mobot.org/ 

manual.plantas/028593/S028700.html. 3 p. 

Natural Resources Conservation Service. 2002. 

Plant profile for Ludwigia octovalvis (Jacq.) 

Raven ssp. octovalvis. http://plants.usda.gov/ 

cgi_bin/plant_profile.cgi?symbol=LUOCO. 2 p. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CABI Publishing, Wallingford, UK and 

New York. 917 p. 
451

Lupinus arboreus Sims yellow bush lupine 

FABACEAE 

Synonyms: None 

Christopher Ross

General Description.—Yellow bush lupine is a 

small erect shrub, less than 20 dm tall, with 

cauline leaves on 2- to 3- cm petioles, and 

composed of five to 12 leaflets, on 8 to 12 mm 

stipules. Leaf colors range from green and smooth 

to silver and hairy. In northern California it grades 

into and coexists with L. rivularis Doug. ex Lindl., 

distinguished by blue flowers (Sholars 1993). 

Range.—Yellow bush lupine is native on coastal 

bluffs and dunes, or less than 5 km inland 

(Davidson and Barbour 1977), in central 

California from Ventura to Sonoma Counties, and 

is widely naturalized on the northern California 

coast (Sholars 1993) and as far north as 

Vancouver, Canada (Hitchcock and Cronquist 

1973). There has been considerable debate over 

the native range, although it probably encompasses 

only the southern part of the current range. 

Ecology.—Yellow bush lupine is an aggressive 

colonizer of dunes, where it outcompetes native 

dune mat species. It encourages other non native 

invaders by fixing nitrogen (Pickart and others 

1998) and creating shade. Outside its former 

range, it also hybridizes with native lupines, 

especially L. rivularis (National Park Service 

2001). The California Native Plant Society places 

it on the “A” List of invasive weeds in Humboldt 

County; however, it is apparently only a serious 

invader of dune communities (Anderson et al. 

1999). Most lupines are poisonous to large 

herbivores, but I found no data on this species. 

Yellow bush lupine is a strong nitrogen fixer, and 

is used in New Zealand as a planted precursor to 

tree planting in unstable, nitrogen-deficient dune 

environments, where it has been estimated to 

contribute 220 kg of nitrogen/ha in 4 1⁄2 years of 

growth (Gadgil 1971). In California coastal 

prairies it is a facilitator of exotic weed and annual 

grass invasions, which establish after the dense 

canopies of the short-lived lupines are opened by 

fire or insects (Maron and Connors 1996). A 

variety of insects prey upon it, sometimes killing 

extensive stands. Among the predators are the 

subterranean ghost moth Hephialus californicus

and the tussock moth Orgyia vetusta. The nitrogen 

rich litter that results from such mortality makes 

establishment of native vegetation uncertain and a 

long-term process (Maron and Jefferies 1999). 

Reproduction.—Inflorescences are 10 to 30 cm, 

with flowers sometimes arranged in whorls. 

Flowers are 14 to 18 mm with upper lip 5 to 9 mm 

and lower lips 5 to 7 mm. Petals are yellow, or 

north of central California lilac to purple. Pods are 

hairy, brown or black, 4 to 7 cm long. Eight to 12 

seeds per pod are black, tan, or striped, and are 4 

to 5 mm long (Sholars 1993). Flowers are both 

self-compatible and outcrossing and have 

generated considerable interest from genetic 

investigators. Self-pollination requires facilitation, 

usually from insects, but is responsible for only 

about 25 percent of seed production (Kittelson and 

Maron 2000). 

Fire Effects.—Although fire may cause 

widespread mortality in dense lupine stands, and 

may be followed by invasion of annual weeds and 

grasses (author’s observation), the long-lived 

lupine seed banks may result in reestablishment of 

lupine (Maron and Connors 1996). The annual 

weeds and grasses that follow the nitrogen 
452 

enrichment by lupine potentially alter fire regimes, 

increasing frequency and intensity of burns. 

Growth and Management.—Yellow bush lupine 

is fast growing and short lived (maximum 7 years) 

(Davidson and Barbour 1977). Stands may grow 

rapidly and die off in as little as 3 years (Strong et 

al. 1995). Restoration of lupine-invaded sites 

should include litter and duff removal in addition 

to direct removal of lupine, because it enriches soil 

to the detriment of native competitors (Pickart and 

others 1998). In New Zealand, where it is used as a 

soil developer, it is controlled prior to tree planting 

by crushing and herbicides (Gadgil 1971). Coastal 

dune restoration in northern California has focused 

heavily on removal of yellow bush lupine. Manual 

removal is accomplished by pulling up small 

plants and chopping larger ones, which rarely 

crown sprout. Heavy equipment has also been used 

successfully, but all mechanical methods are labor 

intensive and expensive (Pickart and Sawyer 

1998). Although others have noted a long-lived 

seed bank (see Maron and Connors 1996 and 

above), Pickart and Sawyer (1998) found no 

recruitment by 3 years after removal of established 

stands. 

Benefits.—Yellow bush lupine has been widely 

cultivated as a sand stabilizer and ornamental in 

coastal California. It is a strikingly beautiful plant, 

both in and out of bloom, although its short life 

and messy litter probably limit its value as an 

ornamental. Elsewhere it has been used to prepare 

coastal and dune soils for planting of 

commercially useful trees (Gadgil 1971). It 

provides habitat for birds, reptiles, and rodents. 

References 

Anderson, L.W.J., J. DiTomaso, G.F. Hruss, and 

M. Rejmanek. 1999. The CalEPPC List: Exotic 

Pest Plants of Greatest Ecological Concern in 

California. California Exotic Pest Plant 

Council. Petaluma, CA. 12 p. 

Davidson, E.D. and M.G. Barbour. 1977. 

Germination, establishment, and demography 

of coastal bush lupine (Lupinus arboreus) at 

Bodega Head, California. Ecology 58:592-600. 

Gadgil, R.L. 1971. The nutritional role of Lupinus 

arboreus in coastal sand dune forestry. Plant 

and Soil 35:113-126. 

Hitchcock, C.L. and A. Cronquist. 1973. Flora of 

the Pacific Northwest: an illustrated manual. 

University of Washington Press. Seattle, WA. 

730 p. 

Kittelson, P.M. and J.L. Maron. 2000. 

Outcrossing rate and inbreeding depression in 

the perennial yellow bush lupine, Lupinus 

arboreus (Fabaceae). American Journal of 

Botany 87: 652-660. 

Maron, J.L. and P.G. Connors. 1996. A native 

nitrogen-fixing shrub facilitates weed invasion. 

Oecologia 105:302-312. 

Maron, J.L. and R.L. Jefferies. 1999. Bush lupine 

mortality: Altered resource availability and 

alternative vegetation states. Ecology 80: 443-

454. 

Pickart, A.J., L.M. Miller, and T.E. Duebendorfer. 

1998. Yellow bush lupine invasion in northern 

California coastal dunes. Ecological impacts 

and manual restoration techniques. Restoration 

Ecology 6(1): 59-68. 

Pickart, A.J. and J.O. Sawyer. 1998. Ecology and 

Restoration of Northern California Coastal 

Dunes. California Native Plant Society. 

Sacramento, California. 152 p. 

Sholars, T. 1993. Lupinus Lupine in Hickman, 

J.C., ed. 1993. The Jepson Manual: higher 

plants of California. University of California 

Press, Berkeley and Los Angeles. 1,400 p. 

Strong, D.R., J.L. Maron, P.G. Connors, A. 

Whipple, S. Harrison, and R.L. Jefferies. 1995. 

High mortality, fluctuation in numbers, and 

heavy subterranean insect herbivory in bush 

lupine, Lupinus arboreus. Oecologia 104: 85-

92. 

U.S.D.I. National Park Service, 2001. What is 

yellow bush lupine? http://www.nps. 

gov/redw/y-lupine. 
453

Lycium pallidum Miers pale wolfberry

SOLANACEAE 

Synonyms: None 

Juanita A.R. Ladyman 

 

Illustration credit: Tonia Masaood 

General Description.—Lycium pallidum is 

commonly known as pale wolfberry, pale lycium, 

rabbit thorn (Epple 1995), box thorn (Derig and 

Fuller 2001), chico, cambronera, and tomatillo 

(Elmore 1976). The latter name can be confusing 

because “tomatillo” also commonly refers to 

species of Physalis (Tull 1987). The epithet 

“pallidum” refers to the pale colored leaves. Pale 

wolfberry are thorny, intricately-branched shrubs 

that form loose thickets. Unlike other Lycium

species it is seldom drought deciduous (Bowers 

1993, Kearney and others 1960). The pale green, 

elongated-oblong leaves are adjacent to short, 

sharp thorns or spines. The flowers are greenish 

cream, sometimes tinged with purple, and 

funnelform in shape. The corolla tube is longer 

than 12 mm. The stamens are usually exerted 

(Vines 1986, Welsh and others 1993). The oval 

fruits are red colored and approximately 1 cm or 

less in diameter (Vines 1986). Typically the plants 

are 1 m high shrubs but may grow to 2 m (Elmore 

1976, Carter 1997). Two varieties of Lycium

pallidum are recognized: var. oligospermum C.L. 

Hitchc., and var. pallidum (Kartesz 1994). Variety 

oligospermum differs from var. pallidum in having 

smaller flowers (corolla tube less than 12mm), 

smaller anthers (1 to 1.5mm versus 1.8 to 3 mm) 

and fewer, only approximately seven seeds, per 

berry (Cronquist and others 1984, Baldwin and 

others 2002). 

Range.—Pale wolfberry is a common shrub in 

Arizona, southern Colorado, southern California, 

Nevada, New Mexico, western Oklahoma, western 

Texas, and Utah in the U.S.A., and in Sonora, 

Chihuahua, Zacatecas, and San Luis Potosí in 

Mexico (McGregor and others 1986, Vines 1986, 

Carter 1988, Welsh and others 1993). Variety 

oligosperma occurs only in Nevada and California 

(Integrated Taxonomic Information System 2002). 

Ecology.—Pale wolfberry grows at elevations of 

910 m to 2,134 m in a variety of habitats and soils, 

including those derived from limestone and 

igneous formations. It often grows in sandy soils 

and is tolerant of saline soils (Kearney and others 

1960). It is frequently a member, and may be a codominant, in blackbrush, sagebrush, four-wing 

saltbush, mesquite, greasewood, mountain brush, 

and pinyon-juniper communities (Dick-Peddie 

1993, MacMahon 1988, Welsh and others 1993). 

However, it has never been reported as a dominant 

species. Pale wolfberry grows throughout the 

Sonoran, Chihuahuan, and Mohave Deserts. At 

higher elevations it grows on dry slopes and 

plains, but in the deserts it tends to grow in washes 

or other areas where run-off supplements rainfall 

(Bowers and McLaughlin 1987, Bowers 1993). 

Wolfberry species have tough and fibrous roots 

and the root systems are relatively extensive 

extending 25 to 30 feet (7.5 to 9.0 m) from the 

plant (Matthews 1994) that suggests they may be 

important in retarding soil erosion. 

Reproduction.—Pale wolfberry generally flowers 

in April to June but in some regions may flower as 

early as February (Powell 1998, Epple 1995). The 

fruits mature in the late spring and summer 

(Bowers 1993, Vines 1986). The fragrant, creamyyellow to yellowish-green flowers are insectpollinated. The bright, shiny red berries contain 20 

to 50 seeds. 
454 

Growth and Management.—Pale wolfberry 

sprouts readily when cut or broken (Van Dersal 

1938). Once established, shrub longevity may be 

substantial. In one Arizona study 69 percent of 

individual shrubs of a related species, Lycium 

berlandii, survived for at least 50 years (Goldberg 

and Turner 1986). Seeds are likely dispersed by 

birds and other animals that feed on the berries 

(Bowers 1993). It can be propagated from 

hardwood cuttings, suckers, layers, and seeds. 

Benefits.— Pale wolfberry is an important source 

of food for wildlife, especially birds and rodents 

(Elmore 1987, Bowers 1993, Powell 1998). In 

many areas of degraded habitat in the 

Southwestern United States it frequently provides 

a source of berries and browse, in an otherwise 

depauperate landscape (authors personal 

observation). It has been reported to be locally 

important and valuable as browse for livestock 

especially on winter range (Van Dersal 1938). The 

berries also are eaten by humans, especially Native 

Americans, who have also used other parts of the 

plant as medicine and in ceremonials (Moerman 

1998, Stevenson 1915). In times of famine the 

Native Americans of northern Arizona ate the 

dried berries mixed with saline clay (Kearney and 

others 1960). In this form they were called “food 

clay” by the Navajo or “potato clay” by the Hopi 

(Dunmire and Tierney 1997). As well as being an 

extender, the clay may have been used to reduce 

bitterness. With the recent interest in using native 

plants, more palatable recipes for sauces and jams 

using pale wolfberry berries are available (Derig 

and Fuller 2001). Medicinal uses include applying 

the soaked leaves to cuts and using the ground-up 

roots for toothaches. These hardy shrubs have been 

used as ornamentals since 1878 but on a limited 

scale (Vines 1986, Powell 1998). Stands of pale 

wolfberry are frequently associated with ancient 

Anasazi ruins in the Four Corners area of the 

Southwestern United States (Dunmire and Tierney 

1995). Some stands have been speculated to be 

deliberate plantings and others the result of 

inadvertent seed dispersal. After a pueblo is 

abandoned, moisture tends to collect over 

decomposing subsurface floors and plazas and 

these conditions may make suitable habitat for 

seeds that were unintentionally dropped by the 

inhabitants of the ancient Pueblos (Dunmire and 

Tierney 1997). 

References 

Baldwin, B.G., S. Boyd, B.J. Ertter, R.W. 

Patterson, T.J. Rosatti, D.H. Wilken, M. 

Wetherwax. 2002. The Jepson desert manual. 

University of California Press, Berkeley, CA. 

626 p. 

Bowers, J.E. 1993. Shrubs and trees of the 

Southwest Deserts. Southwest Parks and 

Monuments Association Tucson, AZ. 140 p. 

Bowers, J. E. and S.P. McLaughlin. 1987. Flora 

and vegetation of the Rincon Mountains, Pima 

County, Arizona. Desert Plants. 8(2): 50-94. 

Carter, J.L. 1988 Trees and shrubs of Colorado. 

Johnson Books, Boulder, CO. 165 p. 

Carter, J.L. 1997 Trees and shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Cronquist, A., A. H. Holmgren, N.H. Holmgren, J. 

L. Reveal and P. K. Holmgren. 1984. 

Intermountain Flora, Vascular Plants of the 

Intermountain West, U.S.A. Vol. 4, New York 

Botanical Garden, Bronx, NY. 279 p. 

Derig, B. B. and M. C. Fuller. 2001. Wild berries 

of the west. Mountain Press Publishing 

Company, Missoula, Montana. 235 p. 

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation–past, present, and future. University 

of New Mexico Press, Albuquerque, NM. 244 p. 

Dunmire, W.W. and G.D. Tierney. 1995. Wild 

plants of the Pueblo Province. Museum of New 

Mexico Press, Santa Fe, NM. 290 p. 

Dunmire, W.W. and G.D. Tierney. 1997. Wild 

plants and native peoples of the Four Corners. 

Museum of New Mexico Press, Santa Fe, NM. 

312 p. 

Elmore. F.H. 1976. Shrubs and Trees of the 

Southwest Uplands. Southwest Parks and 

Monuments Assoc. Tucson, AZ. 214 p. 

Epple, A.O. 1995. A Field Guide to the Plants of 

Arizona. Falcon Press Publishing Co., Helena, 

MT. 347 p. 

Goldberg, D.E. and R.M. Turner. 1986. Vegetation 

change and plant demography in permanent 
455

plots in the Sonoran Desert. Ecology 67(3): 695-

712. 

Integrated Taxonomic Information System. 2002. 

Lycium pallidum. Plants, database 

http://www.itis.usda.gov/ [not paged]. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada and 

Greenland. Vol.1 - Checklist. 2nd ed. Timber 

Press, Portland, OR. 622 p. 

Kearney, T.H., R.H. Peebles, and collaborators. 

1960. Arizona flora. 2nd Ed. University of 

California Press, Berkeley, CA. 1,085 p. 

MacMahon, J.A. 1988. Warm deserts. In: M.G. 

Barbour, Billings and W. Dwight, eds. North 

American terrestrial vegetation. Cambridge 

University Press, New York. p 231-264. 

Matthews, R. F. 1994. Lycium pallidum. U.S.D.A. 

Forest Service, Rocky Mountain Research 

Station, Fire Sciences Laboratory. Fire Effects 

Information System. http://www.fs.fed. 

us/databse/feis/plants. 

McGregor, R. L., T.M. Barkley, RE Brooks, and 

EK Schofield. 1986. Flora of the Great Plains. 

University Press of Kansas, Lawrence KS, 

U.S.A. 1,402 p. 

Moerman, D.E. 1998. Native American Ethnobotany. Timber Press, Portland, OR. 927 p. 

Powell, A. M. 1998. Trees & shrubs of TransPecos and adjacent areas. University of Texas 

Press, Austin, TX. 498 p. 

Stevenson, M. C. 1915. Ethnobotany of the Zuni 

Indians. Annual Report 30. Smithsonian 

Institution-Bureau of American Ethology. 102 p. 

Tull, D. 1987. Edible and Useful Plants of Texas 

and the Southwest. University of Texas Press. 

Austin, TX. 518 p. 

Van Dersal, W.R. 1938. Native woody plants of 

the United States, their erosion-control and 

wildlife values. Miscellaneous Publication 303. 

U.S. Deptment of Agriculture. Washington D.C. 

Vines, R.A. 1986. Trees, shrubs, and woody vines 

of the Southwest. University of Texas Press. 

Austin, TX. 1,104 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1993. A Utah Flora. Second Edition, 

revised. Brigham Young University, Provo, 

Utah. 986 p. 
456 

Macfadyena unguis-cati (L.) A.H. Gentry cat’s claw 

BIGNONIACEAE 

Synonyms: Bignonia unguis L. emend. DC. 

 Bignonia unguis-cati L. 

 Batocydia unguis (L. emend. DC.) Mart. ex DC. 

 Doxantha unguis (L. emend. DC.) Miers 

 Doxantha praesignis Miers 

 Doxantha serrulata Miers 

John K. Francis 

 

 

General Description.—Cat’s claw is a woody 

vine or occasionally a scrambling shrub. The 

name, and its equivalent in Spanish, uña de gato, 

comes from the tripartite, hooked tendrils 

resembling an animal’s claws that enable the vine 

to adhere to tree bark and other surfaces 

(Acevedo-Rodríguez 1985). Old plants are more 

or less free hanging. It is also known by the 

common names bejuco de gato, paz y justicia, and 

griffe à chatte (Howard 1989, Liogier 1995). Cat’s 

claw has a strong and flexible, cylindrical stem 

that is brown in color with many lenticels. The 

stems produce adventitious roots to anchor them 

tightly to vertical surfaces. Cat’s claw may exceed 

8 cm in stem diameter and extend more than 20 m 

into the crowns of trees. A long main root extends 

laterally near the soil surface with finer roots 

scattered over its length. Tubers are produced by 

both young and old plants (Stockard 2001). The 

plant grows and maintains few branches until the 

growing tip reaches increased light. The 

compound leaves have two leaflets with the 

clawed tendril between them. The leaves are 

generally ovate to lanceolate in shape but quite 

variable in size. Depending on both age and 

environment, petioles may vary from 0.25 to 2.5 

cm in length, and blades may vary from 1 to 16 

cm in length. They are dark green above and 

lighter below. Inflorescences are axillary with one 

to three flowers. The 4.5 to 10 cm-long tubular 

flowers have five lobes and are bright yellow with 

red-orange lines in the throat. From them develops 

a long (up to 75 cm), narrow (1.0 to 1.5 cm), 

flattened capsule that produces brown, flattened, 

winged seeds. The species has both diploid (2n = 

40) and tetraploid (2n = 80) populations 

(Acevedo-Rodríguez 1985, Gentry 1983, Howard 

1989, Liogier 1995). 

Range.—Cat’s claw is native to the Greater and 

Lesser Antilles, Mexico, Central America, and 

South America to Argentina (Howard 1989). The 

species is reported to have naturalized and escaped 

in Florida, Texas, Hawaii (Aquatic Plant Control 

Operations Support Center 2001), New Caledonia 

(Pacific Island Ecosystems at Risk 2001), 

Australia (Stockard 2001), South Africa (Pest 

Cabweb 2001). 

Ecology.—In Puerto Rico, cat’s claw grows from 

near sea level to over 600 m in elevation and in 

sites that receive mean annual rainfalls from about 

750 to about 2400 mm. It will tolerate most soils, 

except very poorly drained and salty soils. The 

species can tolerate a few degrees of frost, being 

killed to the ground but resprouting afterward 

(Watkins and Sheehan 1975). Cat’s claw is 
457

moderately shade tolerant as a young plant and 

grows in both full sun and under forest canopies. It 

may be less shade tolerant as an adult. The species 

is common in savannas, secondary forest, and 

remnant high forest. It can survive grazing and fire 

but is eliminated by deep grass swards. 

Reproduction.—In Hawaii, flowering takes place 

during the spring (Rauch and Weissich 2000) 

whereas there are actually two periods of 

flowering in Puerto Rico, both during dry seasons 

(Acevedo-Rodríguez 1985). In Costa Rica, nearly 

all the plants flower at once and flowering lasts 

just a short time (Gentry 1983). The capsules 

mature about 6 months after flowering. The 

winged seeds are dispersed by the wind. Seedlings 

are common and widespread in suitable habitat. 

Eleven capsules collected in Puerto Rico had a 

maximum of 212 seeds and a minimum of 106 

seeds. These seeds (air dry) averaged 0.0224 ± 

0.0005 g/seed or 45,000 seeds/kg. One hundred 

percent of the seeds tested germinated between 49 

and 95 days after sowing in commercial potting 

mix. Young plants sprout when damaged and layer 

(root) whenever stems touch the ground. Cat’s 

claw, for ornamental uses, have been propagated 

by seed, cuttings, and layering (Turner and 

Wasson 1999). 

Growth and Management.—Stem extension is 

moderately rapid, especially in sprouts. Diameter 

growth of cat’s claw stems is slow. The vines are 

long-lived, nearly as long as the trees that they 

claim for support. There are relatively few 

problems in establishing cat’s claw. Insect and 

disease problems are rare. The biggest problem is 

controlling ornamental plants. Yearly pruning 

after the flowering season is essential to 

maintaining them within the bounds desired. In its 

native range cat’s claw vines have only a minor 

effect on the trees they parasitize. However, the 

species is beginning to seriously suppress native 

vegetation in parts of Florida (Florida Exotic Plant 

Council 2001) and Australia (Stockard 2001). The 

only means of control recommended at present is 

to cut the vines and paint the cut ends with 

glyphosate herbicide (Stockard 2001). However, 

Vélez and van Overbeek (1950) report that older 

plants are killed by simply cutting the stems. A 

more practical long-term solution is hoped for in 

the ongoing biological control introductions in 

several countries (Pest Cabweb 2001). 

Benefits.—Despite the propensity to naturalize 

and compete with native vegetation, cat’s claw is 

still a handsome ornamental. It is often used to 

cover fences or screen unsightly buildings. Cat’s 

claw is recommended for areas where there is little 

chance of invasion and especially in desert areas 

because it requires relatively little water for 

maintenance (Desert-Tropicals 2001). It was 

reported that a preparation made from cat’s claw 

was used to treat dermatitis from Hippomane 

mancinella L. (Michell and Rook 2001). A 

number of other uses in herbal medicine from 

diverse locations are cited by Liogier (1990). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Aquatic Plant Control Operations Support Center. 

2001. Macfadyena unguis-cati—catclaw vine. 

www.saj.usace.army.mil/conops/apc/newtt/cat1

maps/macfadyenaunguis-cati.htm. 1 p. 

Desert-Tropicals. 2001. Philippe and Sura’s 

Phoenix Tropical Gardens. http://www.deserttropicals.com. 3 p. 

Florida Exotic Plant Council. 2001. Macfadyena 

unguis-cati (L.) A. Gentry. http://www. 

fleppc.org/pdf/Macfadyena%20unguis-cati.pdf. 

2 p. 

Gentry, A.H. 1983. Macfadyena unguis-cati (uña 

de gato, cat-claw, bignone). In: D.H. Janzen, ed. 

Costa Rican natural history. University of 

Chicago Press, Chicago and London. p. 272-

273. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Vol. 6. Arnold Arboretum, 

Harvard University, Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Edicones, 

Inc. San Juan, PR. 566 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Michell, J. and A. Rook. 2001. Botanical 
458 

Dermatology Database. http://bodd.cf.ac.UK/ 

BotDermFolder/BotDermB/BIGN.html. 3 p. 

Pacific Island Ecosystems at Risk. 2001. 

Macfadyena unguis-cati (L.) Gentry, 

Bigniniaceae. http://www.hear.org/pier/maung. 

htm. 2 p. 

Pest Cabweb. 2001. Spotlight on biological control 

of weeds. http://pest.cabweb.org/Features/ 

Spotlight/spot00-2.htm. 21 p. 

Rauch, F.D. and P.R. Weissich. 2000. Plants for 

tropical landscapes, a gardener’s guide. University 

of Hawaii Press, Honolulu, HI. 139 p. 

Stockard, J. 2001. The regeneration of Wingham 

Brush, NSW. AABR lecture series. http:// 

www.zip.com.au/~aabr/info/seminars/seminar0.

4.html. 5 p. 

Turner, R.J. and E. Wasson. 1999. Botanica. 

Barnes and Noble, Inc., New York. 1020 p. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Río Piedras, PR. 497 p. 

Watkins, J.V. and T.J. Sheehan. 1975. Florida 

landscape plants, native and exotic. The 

University Presses of Florida, Gainesville, FL. 

420 p. 
459

Machaonia portoricensis Baill. alfilerillo 

RUBIACEAE 

Synonyms: none 

John K. Francis 

General Description.—Alfilerillo, also known as 

roseta, is a spiny shrub or small tree that 

occasionally reaches 6 m in height and 7.5 cm in 

stem diameter. It is supported by an extensive 

lateral root system. Alfilerillo may have single or 

multiple stems arising just above the ground level, 

but it is always branchy and thorny. The stem bark 

is gray and smooth becoming fissured in old 

plants. The branches and twigs are brown, slender 

and stiff and support paired spines 1 to 2.5 cm 

long at the nodes and ends of twigs. The plant 

tends to have a slender, vertical profile. Opposite 

or whorled, almost sessile leaves are orbicular to 

elliptic with entire edges and blades 7 to 16 mm 

long and 5 to 10 cm broad. Tiny, white flowers are 

grouped in many-flowered terminal panicles. The 

capsules are turbinate (inverse conical), brown, 

quadrangular, 4 to 5 mm long, crowned with four 

calyx lobes and contain two seeds (Liogier 1997, 

Little and others 1974). 

Range.—Alfilerillo is native to Puerto Rico 

(Liogier 1997) from Salinas to Cabo Rojo. A 

closely-related, recently-separated species, 

Machaonia woodburyana Acev.-Rodr., occurs on 

St. John in the U.S. Virgin Islands (New York 

Botanical Garden 2002) where it is endangered 

(University of the Virgin Islands 2002). Alfilerillo 

is not known to have been planted or naturalized 

elsewhere. 

Ecology.—Alfilerillo grows on a wide variety of 

well-drained soils derived from a variety of rock 

types including limestone and ultramafics 

(serpentine). These are usually rocky, sandy, or 

moderately salty so that competition is somewhat 

reduced. It is moderately intolerant of shade and 

can inhabit forests with low basal areas. The 

species grows in dry habitat that receives from 

about 750 to about 950 mm of mean annual 

precipitation at elevations ranging from near sea 

level to about 400 m. Alfilerillo is relatively 

tolerant of fire, being killed to the ground, but 

readily sprouting. It seems to be resistant to 

grazing by cattle probably owing to the spines. 

The species may be found in remnant forests on 

ridges and bluffs, in overgrazed brush lands and 

savannas, near mangroves and salt ponds, and in 

beach strand vegetation (author’s observation, 

New York Botanical Garden 2002, Vázquez and 

Kolterman 1998). 

Reproduction.—Alfilerillo flowers and fruits 

irregularly through the year (Little and others 

1974). Seeds collected from trees in western 

Puerto Rico weighed an average of 0.00182 g/seed 

or 548,000 seeds/kg. Sown on moist filter paper, 

24 percent germinated within 54 days. 

Germination is epigeal. No specialized means of 

seed dispersal is apparent aside from a weak 

tendency to cling to clothing. Seedlings are 

uncommon in spite of relatively good seed 

production. Plants sprout when disturbed. 

Growth and Management.—Growth of 

alfilerillo is relatively slow. A 3-cm thick stem cut 

by the author had 11 growth rings. No planting or 

management experience has been published. 

Benefits.—Afilerillo helps protect the soil in the 

fragile environments where it grows and furnishes 
460 

cover for wildlife. The wood, which is lightbrown, hard, and brittle, is good for fuel. 

References 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

New York Botanical Garden. 2002. West Indian 

vascular plant types catalog. http:// 

scisun.nybg.org:8890/searchdb/owa/wwwcatalo

g.detail_list. [not paged]. 

University of the Virgin Islands. 2002. Endangered 

and threatened plants and animals in the U.S. 

Virgin Islands. http://rps.uvi.edu/CES/ 

endangered.html. 4 p. 

Vázquez, O.J. and D.A. Kolterman. 1998. Floristic 

composition and vegetation types of the Punta 

Guaniquilla Natural Reserve—Cabo Rojo, 

Puerto Rico. Caribbean Journal of Science 34(3-

4): 265-279. 
461

Mahonia aquifolium (Pursh) Nutt. Oregon grape 

BERBERIDACEAE 

Synonyms: Berberis aquifolium Pursh 

 Berberis piperiana (Abrams) McMinn 

 Mahonia piperiana Abrams 

 Odostemon aquifolium (Pursh) Rydb. 

John K. Francis 

General Description.—Oregon grape, also called 

tall mahonia, hollyleaved barberry, mountain 

grape, Oregon grape-holly, and Oregon hollygrape, is an upright evergreen shrub from 0.6 to 3 

m in height. The stems are slender, stiff, usually 

upright, and usually without branches. The wood 

is whitish and brittle and has a significant pith. 

Bark of young stems is green; older bark is rough 

and gray-brown with yellow inner bark. The root 

systems produce horizontal rhizomes up to 1.8 m 

long from which suckers arise. The foliage 

attaches directly to the stems. Alternate, stiff and 

leathery, pinnately compound leaves have five to 

13 leaflets with a form like holly (coarse dentate 

with prickles). Leaves are dark green above, 

except in winter, when exposed leaves take on a 

purplish color. The species is monoecious with 

perfect, bright-yellow flowers in terminal 

(sometimes lateral) racemes. Flowers have a 

honey-like fragrance. Fruits (berries), 6 to 10 mm 

long, ripen from green to blue or blue-black in late 

summer (Abrams 1950, Center for Applied 

Nursery Research. 2003, Dirr 1983). Oregon grape 

is the state flower of Oregon (Paghat.com 2003). 

Range.—Oregon grape is native to British 

Colombia, Washington, Oregon, northern 

California, and northern Idaho (Abrams 1950, 

Natural Resources Conservation Service 2003). 

The species has been planted widely as an 

ornamental and medicinal plant and has 

naturalized in Michigan, Ohio, Kentucky, New 

York, New Jersey, United Kingdom, Germany and 

probably other areas (Auge and Brandl 1997, 

Natural Resources Conservation Service 2003, 

Plants for a Future 2003). Oregon grape hybridizes 

naturally with Mahonia repens (Lindl.) G. Don 

(many ornamentals are actually this hybrid) and 

Mahonia pinnata (Lag.) Fedde (Plants For a 

Future 2003). At least three artificial intergeneric 

crosses between Oregon grape and Berberis

species have been made (Dirr 1983). 

Ecology.—Oregon grape is tolerant of shade and 

normally grows in the forest understory. Natural 

stands contain scattered individual plants arising 

from seeds and clumps of the species arising both 

from seed and suckers. The species grows well in 

soils with a wide variety of properties. However, 

ornamentals do not do well in dry, wind-swept 

sites, or soils with high pH’s (above 8.0). The tops 

are severely injured at –29 °C (Dirr 1983). It 

grows in areas with natural precipitation ranging 

from 950 to 5450 mm annually (Las Pilitas 

Nursery 2003). A number of diseases and insects 

attack the leaves and fruits but usually do not 

cause serious damage. 

Reproduction.—Oregon grape blooms from April 

to June (Clark and Trelawny 1976). Flowers are 

self-fertile and pollinated by insects (Plants for a 

Future 2003). The fruits mature in late July 

through September and remain on the plants into 

December (Center for Applied Nursery Research 

2003). Fruits collected by the author from 

ornamental plants in Utah averaged 0.312 ± 0.013 

g/fruit. Air-dried seeds separated from them 

averaged 0.0067 + 0.0002 g/seed or 149,000 

seeds/kg. The number of seeds/fruit varied from 

zero to nine. Fruits can be harvested in quantity by 

hand and can be cleaned by gentile maceration, 
462 

wet sieving, and screening. Seeds should not be 

allowed to dry out completely and should be 

stratified at 5 °C before sowing. Taken in 

November and treated with 0.8 to 1.0 percent of 

IBA (Indolbuteric acid), cuttings root well. 

Division of established plants is an alternate 

method of vegetative propagation (Dirr 1983). In 

nature, mostly birds disperse the seeds 

(Paghat.com 2003) and both sexual and asexual 

reproduction are effective (Auge and Brandl 

1997). 

Growth and Management.—Oregon grape is 

moderately slow growing. Individual stems reach a 

60- to 90-cm height in 3 to 4 years (Dirr 1983). 

While individual stems probably have a life of not 

more than 10 years, clones can last much longer. 

Plants are grown for the nursery trade in containers 

and are usually 1 year old when sold. The species 

is planted both in sunny and shady locations. 

Ornamental plants tolerate pruning well (Plants for 

a Future 2003). Data on management of the 

species in natural stands are lacking. 

Benefits.—Oregon grape shrubs contribute to the 

diversity and beauty of understory vegetation in 

native habitats, help protect the soil, and furnish 

food and cover for wildlife. The species is planted 

in temperate areas throughout the world as an 

ornamental, for hedges, and foliage, and leafy 

branches with their fall colors are sometimes used 

for Christmas decorations. The fruits, edible but 

sour, are made into jellies and juice drinks (Clark 

and Trelawny 1976). Flowers are eaten in salads, 

cooked in tempuras, and used to make a lemonadelike drink (Paghat.com 2003). Yellow, green, 

violet, and purplish-blue dyes are made from 

various tissues for arts and crafts (Plants For a 

Future 2003). 

Herbal Medicine.—Infusions of Oregon grape 

root and doses of isolated alkaloid have been used 

effectively in the past to relieve many of the 

symptoms of syphilis (Felter 1922). Ointments 

made from the plant and isolated alkaloids are 

currently enjoying tremendous popularity in 

treating psoriasis and similar skin conditions. In 

clinical trials, symptoms improved or disappeared 

in 81 percent of 443 patients suffering from 

subacute and chronic psoriasis (Gieler and others 

1995). Of the constituent compounds tested, the 

benzylisoquinoline alkaloids berbamine and 

oxyacanthine were principally responsible for 

inhibition of abnormal cell growth, with berberine 

showing lesser activity (Muller and others 1995). 

The fruit serves as a safe and gentle laxative 

(Plants for a Future 2003). 

References 

Abrams, L. 1950. Illustrated flora of the Pacific 

States. Vol. 2. Stanford University Press, 

Stanford, CA. 635 p. 

Auge, H. and R. Brandl. 1997. Seedling 

recruitment in the invasive clonal shrub, 

Mahonia aquifolium (Pursh) Nutt. Oecologia 

Berlin 110(2): 205-211. 

Center for Applied Nursery Research. 2003. 

Mahonia aquifolium—Oregongrapeholly, 

Oregon Grapeholly, Oregon Hollygrape, Oregon 

grape. http://www.nobleplants.com/classnotes/ 

spring/springprofiles/evergreen/mahoniaaquifoli

um.htm. 2 p. 

Clark, L.J. and J.G. Trelawny. 1976. Wild flowers 

of the Pacific Northwest. Gray’s Publishing 

Limited, Sidney, B.C., Canada. 604 p. 

Dirr, M.A. 1983. Manual of woody landscape 

plants: their identification, ornamental 

characteristics, culture, propagation and uses. 

Stipes Publishing Company, Champaign, IL. 

826 p. 

Felter, H.W. 1922. The eclectic material medica, 

pharmacology and therapeutics: Berberis

(Mahonia) aquifolium. published as a database 

in: http://www.ibiblio.org/herbmed/eclectic/ 

felter/hahonia-aqui.html. 2 p. 

Gieler, U., A. Von der Weth, and M. Heger. 1995. 

Mahonia aquifolium—a new type of topical 

treatment for psoriasis. Journal of 

Dermatological Treatment 6(1): 31-34. 

Las Pilitas Nursery. 2003. Mahonia aquifolium. 

Las pilitas Nursery, Escondido, CA. 

http://www.laspilitas.com/plants/420.htm. 2 p. 

Muller, K., K. Ziereis, and I. Gawlik. 1995. The 

anipsoriatic Mahonia aquifolium and its active 

constituents: II. Antiproliferative activity against 

cell growth of human keratinocytes. Planta 

Medica 61(1): 74-75. 

Natural Resources Conservation Service. 2003. 

Plants profile: Mahonia aquifolium (Pursh) Nutt. 

http://plants.usda.gov/cgi_bin/topics.cgi. 4 p. 
463

Paghat.com. 2003. Paghat’s garden: Mahonia 

aquifolium. http://www.paghat.com/ 

oregongrape.html. 4 p. 

Plants For a Future. 2003. Plants for a future: 

database search results: Mahonia aquifolium. 

http://www.ibiblio.org/pfaf/cgi-bin/arr_html 

?Mahonia+aquifolium. 12 p. 
464 

Mahonia repens (Lindl.) G. Don creeping barberry 

BARBERIDACEAE 

Synonyms: Berberis amplectens (Eastw.) L.C. Wheeler 

 Berberis aquifolium Pursh var. repens (Lindl.) Scoggan 

 Berberis pumila Greene 

 Berberis sonnei (Abrams) McMinn 

Mahonia amplectens Eastw. (additional synonyms: Natural Resources 

Conservation Service 2003) 

John K. Francis 

General Description.—Creeping barberry, also 

known as Oregon grape, creeping Oregon grape, 

creeping mahonia, creeping hollygrape, ash 

barberry, and dwarf Oregon grape, is an upright to 

recumbent, evergreen shrub 12 to 40 cm in height 

and 3 to 9 mm in basal diameter. The stems are 

solitary, usually unbranched but occur in clonal 

groups arising from rhizomes. The main root is 

stiff and deep, and the lateral roots are short and 

fine. Stem bark is slightly rough and gray brown, 

and the inner bark is bright yellow, especially the 

inner root bark. The wood and pith are yellow. The 

alternate leaves, which have petioles 2 to 3 cm 

long, are pinnately compound with three to seven 

leathery, sessile, ovate leaflets that are dull on both 

surfaces, and have spiny-toothed margins. There 

are usually two to eight leaves per plant. Several 

flowers with six petals and six sepals, both yellow, 

are clustered in terminal racemes 3 to 7 cm long. 

Fruits are glaucous, blue berries containing one to 

four seeds. The seeds are 6 to 10 mm long. There 

are 2n = 28 chromosomes (Auger and other 2002, 

Harrison 2003, Lesica 2002, Rehder 1951, Welsh 

and others 1987). Creeping barberry readily 

hybridizes with M. aquifolium (Pursh) Nutt. and 

M. pinnata (Lag.) Fedde when growing near them 

(Pagahat.com 2003). 

Range.—Creeping barberry is native to the 

Western United States as far east as Texas and 

Minnesota, and to British Colombia and Alberta in 

Canada. It is present (probably naturalized) in 

Indiana and Pennsylvania (Natural Resources 

Conservation Service 2003, Walkup 1991). The 

species has been planted widely as an ornamental 

and has probably naturalized in many areas outside 

its native range. 

Ecology.—Low in stature and moderately tolerant 

of shade, creeping barberry is not highly 

competitive. In Utah, it will not grow in heavy 

grass and forb stands. It is most frequently seen in 

rocky or gravely areas with low vegetative cover 

or under open conifer or hardwood stands with 

sparse understory vegetation. However, the species 

is reported to be the dominant understory shrub in 

closed-canopy Douglas fir in central Idaho 

(Walkup 1991). Creeping barberry apparently 

grows well under conifers that produce a thick 

layer of decomposing needles. However, it does 

not flower when growing in shade (Pahgat.com 

2003). Natural populations are affected little by 

treatments such as prescribed fire, clearcutting, 

and scarification. It usually survives (by sprouting) 

even moderate and high-intensity fires (Walkup 

1991). Creeping barberry is an invader of 

disturbed sites (Harrison 2003). It will grow in 

most well-drained soils and is especially common 

in rocky or gravelly soils. The species is much 

more common over limestone or other alkaline 

parent materials than granite or quartzite (Walkup 
465

1991). It tolerates acidic soils but grows poorly 

when concentrations (less than 2.5 mg/L) of boron 

are low (Harrison 2003). Creeping barberry grows 

at elevations of 1,125 to 3,300 m in the inland 

West (Harrison 2003, Walkup 1991). Although it 

is most common on north-facing slopes, it grows 

on south-facing slopes and ridge tops at higher 

elevations. 

Reproduction.—Creeping barberry flowers in 

May through June and matures fruit in June 

through July (Rudolf 1974). The flowers are 

pollinated by bees and butterflies (Paghat.com 

2003). Flowers not cross pollinated with other 

plants will self pollinate, but produce much fewer 

seeds (Walkup 1991). Ratio weight of fresh fruits 

to air-dried seeds is 12.5:1 (Auger and others 

2002). There are 119,000 to 157,000 seeds/kg 

(Rudolf 1974). Cleaned seed can be sealed and 

stored for 5 years (Harrison 2003). Rudolf (1974) 

recommends stratification of 30 days at 1 °C 

followed by 60 days at 21 °C, and then 196 days at 

1 °C. Treated thus, 74 percent of test seeds 

germinated. Alternately, a hot water treatment 

followed by cold stratification for 60 days is 

recommended (Kjelgren 2003). Seeds can be sown 

in fall without pretreatment. In nature, seeds are 

dispersed by birds and to a lesser extent by 

mammals including black bears and rodents 

(Auger and others 2002). Once established, plants 

spread through sprouts from rhizomes (Walkup 

1991). 

Growth and Management.—Growth is slow. 

Individual stems may live 10 years or more. It is 

difficult to imagine the need to control this 

unobtrusive species in its native habitat. On the 

other hand, it should frequently be planted as part 

of environmental restoration projects. The species 

can be artificially propagated by seed, divisions, 

cuttings, and layers (Walkup 1991). Direct seeding 

in the fall is the most practical method to introduce 

it to disturbed sites. Repellants or poisons may be 

necessary because the seeds are readily consumed 

by mice (Kjelgren 2003). 

Benefits.—Creeping barberry adds beauty to the 

forest, helps protect the soil, provides food for 

wildlife, and makes a useful ornamental. It is 

planted as ground cover, especially under light tree 

canopy cover or on poor, gravelly ground. In the 

summer, the leaves are dark green; in winter they 

are green, mottled with bright red, or a bonzepurple. The species is particularly recommended 

because it requires little or no irrigation (Kansas 

State University Research and Extension 1996). It 

is included in reclamation plantings (Welsh and 

others 1987) and has been used especially for 

revegetation of mine spoils (Harrison 2003). Mule, 

white-tailed deer, and sometimes elk browse 

creeping barberry during fall and winter. The 

protein content ranges from 4.7 to 5.5 percent, and 

the species contains high contents of carotene in 

July. Creeping barberry foliage is mildly toxic and 

domestic animals normally make little use of it for 

forage except when other food is scarce (Walkup 

1991). In one case, browsing of creeping barberry 

killed cattle in Arizona (Harrison 2003). Many 

mammals (including black bears) and birds eat the 

fruits (Walkup 1991). The fruits become edible to 

humans after they have passed through at least one 

frost. They are used to make jelly, wine, and a 

juice drink (Harrison 2003). Native Americans 

used the berries and the roots for lavender and 

yellow dyes, respectively, and made teas from the 

roots to treat a range of afflictions (Lesica 2002, 

Paghat.com 2003). Creeping barberry contains the 

alkaloids berberine and oxyacanthin (Harrison 

2003). The species is an alternate host for black 

stem rust of cereal grains (Walkup 1991). 

References 

Auger, J., S.E. Meyer, and H.L. Black. 2002. Are 

American black bears (Ursus americanus) 

legitimate seed dispersers for fleshy-fruited 

shrubs? American Midland Naturalist 147: 352-

367. 

Harrison, S. 2003. Mahonia repens. Crop 

Development Centre, University of 

Saskatchewan, Saskatoon, Saskatchewan, 

Canada. http://www.usask.ca/agriculture/ 

plantsci/classes/range/berberis.html. 4 p. 

Kansas State University Research and Extension. 

1996. Low water use plants for Kansas 

landscapes. MF2067. Kansas State University, 

Manhattan, KS. 4 p. 

Kjelgren, R. 2003. Mahonia repens. Utah State 

University, Logan, UT. http://www.hort.usu. 

edu/natives/shrubs/repens.htm. 1 p. 

Lesica, P. 2002. A flora of Glacier National Park, 

Montana. Oregon State University Press, 

Corvallis, OR. 512 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Mahonia repens (Lindl.) G. Don, 
466 

creeping barberry. http://plants.usda.gov/ 

cgi_bin/plant_search.chi?mode=Scientific+Nam

e&keywordquerry=Mahonia+repens&earl=plant

_search.cgi. 5 p. 

Paghat.com. 2003. Creeping mahonia, aka 

creeping barberry, aka creeping hollygrape, aka 

ash barbarry, aka small or dwarf Oregon grape. 

http://www.paghat.com/oregongrape2.html. 4 p. 

Rehder, A. 1951. Manual of cultivated trees and 

shrubs. The MacMillan Company, New York. 

996 p. 

Rudolf, P.O. 1974. Berbers L., barberry, mahonia. 

In: C.S. Schopmeyer. tech. coord. Seeds of 

woody plants of the United States. Agric. 

Handb. 450. U.S. Department of Agriculture, 

Forest Service, Washington, DC. p. 247-251. 

Walkup, C.J. 1991. Mahonia repens. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shrub/ 

mahrep/all.html. 10 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University Press, Provo, UT. 894 p. 
467

Malvastrum americanum (L.) Torr. Indian Valley false mallow 

MALVACEAE 

Synonyms: Malva americana L. 

 Malva spicata auct. non L. 

 Malvastrum spicatum auct. non (L.) Gray 

 Malva blumeana Steud. 

 Malva gangetica L. 

 Malva borbonica Willd. (and others, Institute of Systematic Botany 2003) 

John K. Francis 

General Description.—Indian Valley false 

mallow, also known as spiked malvastrum, false 

mallow, malva loca, malva silvestre, and mauve 

d’Amérique, is a shrub, subshrub, or woody herb 

that often reaches 1.5 m in height and 1 cm in 

basal diameter. The semideciduous plant generally 

has a single stem that is upright and sparsely 

branched. The wood is light colored, moderately 

hard, and moderately strong. Plants are supported 

by a tap and lateral root system of ivory colored 

roots. The twigs and leaves are stellate-pubescent. 

Gray-green leaves have a 1- to 6-cm petiole, 

usually ovate blades 2 to 8 cm long with serrate 

margins. Inflorescences are dense spikes, terminal 

on stems and branches. The flowers are sessile, 

subtended by an involucel of three linear bracts. 

The petals are pale yellow to yellow-orange and 8 

to 9 mm long. The fruits are 5 to 6 mm in diameter 

and contain 10 to 15 mericarps without spines with 

seeds 1.5 mm long (Howard 1989, Liogier 1994, 

Stevens and others 2001). There are 2n = 24 

chromosomes (Lavia and others 2000). 

Range.—Indian Valley false mallow occurs in 

Florida and throughout the Caribbean including 

Puerto Rico and the U.S. Virgin Islands and from 

Texas through Central and South America to 

Argentina (Howard 1989, Stevens and others 

2001). The species occurs as a naturalized exotic 

in Hawaii (Natural Resources Conservation 

Service 2003), Australia, India, and other locations 

in the Old World tropics and subtropics (Howard 

1989). 

Ecology.—Plants observed by the author appear to 

be short-lived shrubs that grow yearly without 

dieback. The species is reported also growing as 

an herb (annual) and subshrub (dieing back to 

permanent woody parts each year) (Howard 1989, 

Stevens and others 2001). Indian Valley false 

mallow is usually not eaten by cattle or other 

herbivores. Moderate grazing coupled with fire 

tends to make the species more abundant. It also 

grows in areas disturbed by river overflows, 

farming, road building, and land clearing. The 

species is intolerant of shade and severe 

competition. It disappears under closed tree 

canopies or in dense tall grass swards. However, 

Indian Valley false mallow does grow under a 

broken tree canopy. It grows on most types of 

well-drained soil. Mean annual rainfall in Puerto 

Rican habitat ranges from about 750 to 1000 mm 

(author’s observation). It is known from near sea 
468 

level to 1000 m in elevation in Nicaragua (Stevens 

and others 2001). 

Reproduction.—Indian Valley false mallow is 

reported to flower and fruit throughout the year in 

Nicaragua (Stevens and others 2001). In Puerto 

Rico, plants begin flowering at about 15 cm in 

height and continue through life except for pauses 

during dry seasons. Air-dried seeds collected in 

Puerto Rico averaged 1.19 million seeds/kg. 

Placed on moist filter paper without pretreatment, 

15 percent germinated over a period of 6 months. 

No specialized means of dispersal is apparent; the 

seeds are small and undoubtedly are moved to 

some extent by wind, water, and animals. 

Seedlings are relatively common in favorable sites. 

The dry fruits can easily be collected in quantity 

by hand but cleaning the tiny seeds is difficult. 

Growth and Management.—Growth rates are 

moderate (0.3 to 0.5 m/year) and plants live 1 to 5 

years, depending on competition and growing 

conditions. Although control is occasionally 

needed in rangelands and disturbed areas, no 

specific recommended treatments are known to the 

author. 

Benefits and Detriments.—Indian Valley false 

mallow helps protect the soil, furnishes cover for 

wildlife, and because of its blooms contributes 

moderately to the aesthetics of wildlands where it 

grows. It is a host for the cotton bollworm, 

Helicoverpa armigera (Hubner) (Mensah and 

others 2003). The species also is the larval foodplant for the Laviana white-skipper butterfly, 

Heliopetes laviana (Hewitson) of Texas (Quinn 

2003) and a Cuban butterfly, Strymon columella 

cybira (Hewitson) (Fernández 2001). 

References

Fernández, D.M. 2001. New oviposition and larval 

hostplants for twenty-three Cuban butterflies, 

with observations of the biology and distribution 

of some species. Caribbean Journal of Science 

37(1-2): 122-125. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Institute of Systematic Botany. 2003. Atlas of 

Florida vascular plants: Malvastrum 

americanum. http://www.plantatlas.usf.edu/ 

main.asp?plantID=3662. 2 p. 

Lavia, G., G. Seijo, A. Fernández, and A. 

Krapovickas. 2000. Los cariotipos de algunas 

especies de Malvastrum A. Gray (Malvaceae). 

Universidad Nacional del Nordeste, Corrientes, 

Argentina. http://www.unne.edu.ar/cyt/2000/ 

6_biologicas/6_pdf/6_034.pdf. 3 p. 

Liogier, H. A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Mensah, R., M. Dillon, M. Kahn, C. Tann, and L. 

Wilson. 2003. Cotton insect pests and their weed 

hosts. Australian Cotton Cooperative Research 

Centre. http://www.cotton.pi.csiro.au/Assets/ 

PDFFiles/IPMGL99/IPMSO4.pdf. 2 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Malvastrum americanum (L.) 

Torr., Indian Valley false mallow. http:// 

plants.usda.gov/cgi_bin/plant_search.cgi?mode=

Scientific+Name&keywordquery=Malvastrum+

americanum&earl=plant. 3 p. 

Quinn, M. 2003. Caterpillar food plants for the 

lower Rio Grande Valley. http://www.naba. 

org/ftp/plants_info.pdf. 8 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. A Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden Press, St. Louis, 

MO. p. 945-1,910. 
469

Mecranium latifolium (Cogn.) Skean camasey almendro 

MELASTOMATACEAE 

Synonyms: Mecranium amygdalinum (Desr.) C. Wright ex Sauvalle 

 Melastoma amygdalinum Desr. 

John K. Francis 

General Description.—Camasey almendro is a 

shrub or small tree sometimes reaching 7.5 m in 

height and 13 cm in stem diameter. The plant 

frequently has multiple stems arising near the base. 

Horizontal stems and branches produce vertical 

sprouts. Stems, which are covered with graybrown, thin, slightly furrowed bark, tend to be 

straight with relatively few branches. Twigs are 

light green turning brown, slightly four-angled, 

and marked with half-round leaf scars. The wood 

is light tan, weak and brittle, and has a 2-mm pith. 

Plants are supported by a shallow system of dark 

tan, somewhat brittle roots. Opposite, light green 

leaves are elliptic to ovate, pointed at both ends, 

with the margin mostly finely serrate, and 5.4 to 

15.4 cm long. There are three main veins, the two 

lateral ones arising 0.3 to 1.3 cm above the base of 

the blade. Small inflorescences (panicles) are 

axillary and bear several-to-many small, white, 

four-petaled flowers. The juicy fruits (berries) are 

globose, 5.5 mm in diameter, purplish black when 

mature, and have a slightly sweet and slightly 

bitter flavor. Within are many seeds that are light 

brown, tear-drop shaped, and about 0.5 mm long 

by 0.25 mm thick (author’s observation, Liogier 

1995, Little and others 1974). 

Range.—Camasey almendro is native to the 

Luquillo Mountains and the Cordillera Central of 

Puerto Rico and to St. Thomas in the U.S. Virgin 

Islands (Liogier 1995). DNA evidence indicates a 

likelihood that this species arose from M. 

multiflorum (Desr.) Triana or M. septentrionala

Skean now found in the adjacent island of 

Hispaniola (McKenny 2002). The species is not 

known to have been planted or naturalized outside 

the native range. 

Ecology.—Camasey almendro is widely 

distributed throughout the mountain forests in 

Puerto Rico from 300 m elevation upward (Little 

and others 1974) in areas that receive more than 

about 1800 mm of precipitation. Soils are usually 

clayey or loamy, developed from igneous and 

sedimentary rocks with pH’s from about 5.0 to 6.0. 

Camasey almendro flowers and fruits in full sun 

and partial shade. It grows well in moderate shade 

but eventually succumbs to the heavy shade of a 

fully closed forest canopy. Apparently, at least 

mild disturbance is necessary for establishment. 

Such disturbance results naturally from hurricanes 

and other winds that cause tree-fall gaps, rarely 

from landslides, and more recently from road cuts 

and other construction. However, invasion is slow 

and never with large populations. Thirty years 

after a plane crash removed the elfin forest 

vegetation at 1,000 m elevation in the Luquillo 

Mountains of Puerto Rico, 0.1 percent of the stems 

of the resurgent vegetation was camasey almendro 

(Weaver 2000). 

Reproduction.—Camasey almendro flowers and 

fruits throughout the year (Little and others 1974). 

The small fruits are produced in relatively large 

numbers, and each fruit contains a large number of 

seeds. A collection of fruits from Puerto Rico 
470 

averaged 0.138 ± 0.005 g/fruit. The seeds are tiny; 

an air-dried sample averaged 25.4 million/kg. The 

seeds can be cleaned by crushing, wet sieving with 

a fine mesh screen, filtering and drying on the 

paper filter, and rubbing the seed off the paper. 

Without pretreatment seeds took 4 months to begin 

germinating and completed 11 percent germination 

in 11 months. Germination is epigeal, and 

seedlings are about 1 mm across the two cotyledon 

leaves. It is assumed that birds eat the fruits and 

disperse the seeds. Seedlings are scattered to 

relatively common in suitable, disturbed habitat. 

Layering (rooting) occurs whenever horizontal 

stems come in contact with the ground. Camasey 

almendro sprouts when stems are broken or cut. 

Growth and Management.—Camasey almendro 

grows moderately slowly and probably lives at 

least 2 or 3 decades. No planting or management 

experience has been published. Removal of the 

forest canopy and soil disturbance near seed 

sources would likely result in reproduction. 

Benefits.—Camasey almendro helps protect the 

soil and furnishes food and cover for wildlife. It is 

an attractive plant and adds to the aesthetics of the 

forest. The species might be suitable as an 

ornamental in natural landscaping if it proves 

adaptable at lower elevations where the island 

urban centers are located. 

References

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

McKenny, S. 2002. DNA evidence for the origin 

of Mecranium latifolium (Melastomataceae), a 

Puerto Rican endemic. Abstract in: Elkin Isaac 

Research Symposium, Apr. 18, 2002, Albion 

College, Albion, MI. 1 p. 

Weaver, P.L. 2000. Elfin woodland recovery 30 

years after a plane wreck in Puerto Rico’s 

Luquillo Mountains. Caribbean Journal of 

Science 36(1-2): 1-9. 
471

Melochia nodiflora Sw. malva colorada 

STERCULIACEAE 

Synonyms: Mougeotia nodiflora (Sw.) HBK. 

 Riedlea nodiflora (Sw.) DC. 

 Visenia nodiflora (Sw.) Spreng. in L. 

John K. Francis 

General Description.—Malva colorada, also 

known as britónica prieta and mauve, is a 

medium-sized shrub to 3 m in height and 5 cm in 

stem basal diameter. The species forms a fairly 

robust tap and lateral root system. The species is 

herbaceous when small, becoming woody before 

the end of the first year. Malva colorada may have 

single or multiple stems that arise from the root 

crown and lateral roots and large lateral branches 

may form low on the stem. Malva colorada often 

develops only primary and secondary branches. 

The bark of older stems and branches is reddish 

brown. The leaves are alternate with petioles 0.5 

to 3 cm long and ovate or ovate-lanciolate blades 

are 2.5 to 10 cm long and 1.2 to 6 cm broad. The 

tip is pointed, the margin serrate, and the veins 

prominent. The inflorescences are axillary in 

dense globose clusters in which are situated tiny, 

dark pink, pink, or white flowers. The 

inflorescences mature into clusters of reddishbrown capsules each of which contain, when 

completely filled, five 2-mm brown seeds 

(Grisebach 1864, Howard 1989, Liogier 1994). 

Range.—Malva colorada is native to the 

Bahamas, the West Indies, and from Mexico 

through Brazil (Grisebach 1864, Howard 1989, 

Liogier 1994). The species is not known to have 

been planted or naturalized outside its natural 

range. 

Ecology.—Malva colorada grows on all types of 

well drained soils with medium or high fertility 

(Sánchez, and Uranga 1993) derived from 

sedimentary (including limestone), igneous, and 

metamorphic (including ultramafic) rocks. In 

Puerto Rico, it grows in areas that receive from 

750 to about 2000 mm of mean annual 

precipitation. In dry areas, the species is confined 

to riparian zones. Malva colorada grows in Puerto 

Rico from near sea level to over 600 m in 

elevation. The species does not tolerate heavy 

shade and seems to require at least minor soil 

disturbance to become established. Malva 

colorada grows in old fields and neglected 

pastures, on roadsides and fencerows, along river 

flood planes and bars, and on neglected 

construction sites and vacant lots. It competes well 

with low grass and broadleaf weeds but does not 

grow under closed tree canopies. 

Reproduction.—Malva colorada flowers over a 2 

or 3 month period. After flowering, capsules take 

about 6 months to mature seeds. The species 

produces large quantities of seeds that are released 

during the dry season. They are small and are 

carried by wind, water, animals and agricultural 

equipment (Sánchez and Uranga 1993); no 

specialized means of dispersal is known. Seeds of 

malva colorada collected in Puerto Rico weighed 

an average of 0.00167 g/seed or 597,000 seeds/kg. 

Seeds with no pretreatment were placed to 

germinate on moist filter paper, and 36 percent 

germinated over an 11-month period. Germination 

is epigenous. 

Growth and Management.—Individual stems 

live from 2 to about 4 years. Through suckers and 

coppices, plants can live several times longer. 

Malva colorada is sometimes a weed and is 

primarily a problem in perennial plantations such 

as citrus, pasture, coffee, and sugar cane (Sánchez 
472 

and Uranga 1993). The species is controlled by 

mowing, cutting with machete, and by spraying 

with broadleaf weed killers. 

Benefits.—Malva colorada is browsed by goats 

and to a limited extent by cattle. It protects against 

erosion in recently disturbed areas. 

References 

Grisebach, A.H.R. 1864. Flora of the British West 

Indian Islands. J. Cramer-Weinheim, New York. 

789 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Sánchez, P. and H. Uranga. 1993. Plantas 

indeseables de importancia economica en los 

cultivos tropicales. Editorial Científico-Técnisa. 

La Habana, Cuba. 166 p. 
473

Melochia tomentosa L. pyramid bush 

STERCULIACEAE 

Synonyms: Melochia frutescens Jacq. 

 Moluchia frutescens (Jacq.) Medikus 

 Melochia turpiniana Kunth 

 Moluchia tomentosa (L.) Britton 

John K. Francis 

General Description.—Pyramid bush, also 

known as tea bush, raichie, broom wood, broom 

weed, black widow, balsam, bretónica afelpada, 

and bois-champignon, is an erect shrub 0.5 to 4 m 

in height and up to 10 cm in basal diameter. 

Pyramid bush is supported by a tap and lateral root 

system. Usually, a single stem emerges from the 

ground, but it may branch near the base. The 

branches farther up the stem are slender. The bark 

is slate-gray with many lenticels. Stemwood is tan 

colored, moderately hard, and tough. The leaves 

are covered with a short wooly hair giving the 

plant a gray flannel look. Ovate to lanciolate 

leaves are attached by short petioles and have 

blades 1.5 to 10.5 cm long and 0.9 to 8.5 cm broad 

with serrate margins. Its inflorescences are 

axillary umbelliform cymes of perfect flowers 

with five pink, purple, or blue petals that are 8 to 

13.5 mm long. The fruits that develop are 

pyramidal capsules 6 to 9 mm across. The seeds 

are 2 mm long and reddish brown (Howard 1989, 

Liogier 1994). 

Range.—The native range of pyramid bush 

extends from southern Florida and southern Texas 

through the West Indies and Central America into 

Brazil and Colombia (Howard 1989, Liogier 1994, 

Instituto Nacional de Biodiversidad 2002, 

Recursos Hidricos 2002). No planting or escapes 

have been reported outside its natural range. 

Ecology.—Pyramid bush grows in dry areas 

(areas receiving from 750 to 1000 mm of annual 

precipitation in Puerto Rico), usually within a few 

km of the coast. It grows from near sea level to 

altitudes of about 400 m in Puerto Rico. The 

species tolerates a wide variety of well-drained 

soils derived from sedimentary and igneous rocks 

and grows on coastal sands. Because there is 

relatively little competition, pyramid bush is most 

common on sites that are rocky, eroded, or 

excessively drained. In Texas, the species is found 

on sandy or rocky soil in mesquite thickets, palm 

groves, and dry streambeds (San Antonio 

Botanical Gardens 2002). The species inhabits 

pinelands in southern Florida (Long and Lakela 

1976). In Brazil, it is part of the pioneer 

community on the sandy soils of the coastal plain 

(Recursos Hidricos 2002). The species does not 

tolerate heavy shade, and overtopping by trees will 

eliminate it. It is not eaten by cattle and tends to be 

common on overgrazed rangeland--probably 

because competition from grasses and other plants 

is reduced. 

Reproduction.—Pyramid bush flowers at the end 

of the wet season (November and December) and 

ripens fruits about 2 months later. It flowers 

during the summer and early fall in Texas (Everitt 

and Drawe 1993). Air-dried capsules of pyramid 

bush collected in Puerto Rico weighed an average 

of 0.0135 g. Seeds separated from them weighed 

an average of 0.00104 g/seed or 963,000 seeds/kg. 
474 

Sown without pretreatment on moist filter paper, 

79 percent germinated within a short period 6 

months later. The species can be propagated with 

cuttings (San Antonio Botanical Gardens 2002). 

There appear to be no specialized means of seed 

transport; the small seeds are probably disbursed 

incidentally by wind, water, machinery, and cattle. 

Growth and Management.—Nursery-grown 

pyramid bush seedlings in Texas reached 60 cm in 

just 2 months after outplanting (San Antonio 

Botanical Gardens 2002). However, adult plants in 

Puerto Rico add only about 30 cm to their height 

each year. Based on ring counts in Puerto Rico, 

plants appear to live about 10 years. Overgrazing 

to reduce competition may be the best way of 

promoting this species. Pyramid bush is often 

considered a weed in rangeland (Liogier 1990). 

Mowing may be sufficient to control it. 

Benefits.—Pyramid bush appears to have great 

potential as a flowering ornamental for xeric 

landscaping (San Antonio Botanical Gardens 

2002). In natural habitat, it contributes to 

biodiversity, soil stability, and wildlife cover. 

Although cattle avoid it, sheep and goats in Brazil 

browse it in mixture with other native species 

(Charles and others 1983). A tea is made from the 

foliage to treat colds and as an eye wash (Liogier 

1990). 

References 

Charles, T.N.P., A.M. Maia, C. Guimaraes Filho, 

L.M.C. Salviano, and E.A.P. de Figueiredo. 

1983. Effect of roughage supplementation and 

mineral mixture plus deworming on sheep and 

goat production. II. Lamb and kid growth rates. 

Boletim de Pesquisa No. 20. Centro de Pesquisa 

Agropecuaria do Tropico Semi-Arido. 28 p. 

Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs, 

and cacti of South Texas. Texas Tech University 

Press. Lubbock, TX. 213 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Instituto Nacional de Biodiversidad. 2002. The 

cutting edge. Vol. 7, No. 3. http://www. 

inbio.ac.cr/papers/manual_plantas/jul00/jul00lea

.html. 1 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Recursos Hidricos. 2002. Plano diretor de recursos 

hídricos das bacias hidrográficas dos tributaries 

estaduaia dos rios vazas barris e real. Vol. 3, part 

4.1. http://hidricosw.mg.gov.br/vazareal/vol3/ 

41.htm. 4 p. 

San Antonio Botanical Gardens. 2002. Friendly 

natives for Texas. http://www.sabot.org/ 

natives/native.html. 3 p. 
475

Miconia impetiolaris (Sw.) D. Don ex DC. camasey de costilla 

MELASOMATACEAE 

Synonyms: Melastoma impetiolaris (Sw.) D. Don 

 Miconia wydleriana DC. 

 Tamonea impetiolaris (Sw.) Cook & Collins 

John K. Francis 

General Description.—Camasey de costilla, also 

known as camasey colorado, cordobán, oreja de 

mula, dos caras, auguey, hoja de pasmo, punta de 

sarvia, maya, and trios côtes, is a shrub or small 

tree 2 to 8 m in height. The plant is supported by a 

deep taproot and strong, lateral roots. These roots 

are light gray and have fissured bark. Camasey de 

costilla forms multiple stems if cut or damaged. 

Stem bark is brown and finely fissured. Young 

branches are stout, quadrangular, and covered with 

brown hairs; lower leaf surfaces, petioles, and 

flower stalks of camasey de costilla are also 

covered with brown pubescence. Leaf petioles are 

2 to 7 mm long and mostly obscured by the 

cordate leaf base. The leaves are opposite and 

usually have five main nerves. The blades are 

ovate, minutely serrate at the edges, long pointed, 

12 to 40 cm long, and roughly half as wide. Large 

terminal panicles bear many tiny, stalkless white 

flowers. The fruits are berries 4 to 7 mm in 

diameter progressing from red to purple or blueblack as they ripen. Each fruit contains numerous 

tiny brown seeds (Croat 1978, Howard 1989, 

Liogier 1995, Little and others 1974, Stevens and 

others 2001). 

Range.—Camasey de costilla is native to Mexico, 

Central America, South America as far south as 

Bolivia, and to the Greater and Lesser Antilles 

(Liogier 1995, Little and others 1974, Stevens and 

others 2001). Three varieties have been described 

(Stevens and others 2001). The species is not 

known to have been planted or naturalized outside 

its native range. 

Ecology.—Camasey de costilla grows from a few 

meters above sea level to about 600 m in elevation 

(Little and others 1974, Stevens and others 2001). 

It requires moist to wet habitat where it rains from 

about 1500 to about 3000 mm/year. Although a 

wide variety of soils are tolerated, the species is 

most frequent on well drained to somewhat poorly 

drained, neutral to strongly acid clays and clay 

loams, often with the A horizons absent or 

partially gone because of erosion. Excessively 

drained soils are not colonized. Camasey de 

costilla is moderately intolerant of shade. It 

succumbs in dense shade, grows but does not fruit 

under a forest canopy, and fruits in broken or full 

sun. The species requires disturbance to become 

established (Stevens and others 2001). It may be 

found in early and middle secondary forests, 

plantations, fencerows, pastures, and roadsides. 

The species varies from an occasional plant to 

common as dispersed plants or in clumps. 

Reproduction.—Camasey de costilla flowers 

from spring until fall in Puerto Rico (Little and 

others 1974). It reportedly flowers during the dry 

season (February and March) and fruits during the 

wet season (April and May) in Panama (Croat 

1978), and flowers and fruits throughout the year 

in Nicaragua (Stevens and others 2001). The seeds 

are disbursed by birds and primates (Molano and 

others 2002). A collection of fruits from Puerto 

Rico averaged 0.242 + 0.008 g/fruit. Air-dried 

seeds from these fruits averaged 0.00044 g/seed or 
476 

2.3 million seeds/kg. Sown on wet peat, 56 

percent of the seeds germinated between 13 and 

23 days after sowing. The new seedlings are tiny 

and develop slowly (author’s observation). 

Growth and Management.—Camasey de costilla 

is relatively slow growing. About 0.25 to 0.5 m of 

height is added annually. Plants live at least 10 

years and probably much longer. No known 

attempts have been made to plant or manage 

camasey de costilla. Probably management 

practices that maintain forests interspersed with 

pastures, brushlands, and roadsides will ensure at 

least a presence of camasey de costilla. 

Benefits.—Camasey de costilla is employed as a 

living fencepost in Colombia when it springs up 

naturally along fence rows (Molano and others 

2002). The wood is light colored and hard (Little 

and others 1974) and used to a limited extent for 

stakes and fuel. Camasey de costilla benefits 

wildlife by furnishing food (fruits) and cover. It 

also contributes to biodiversity and protects the 

soil from erosion. In herbal medicine, leaf extracts 

are used for aromatic baths, to arrest bleeding, 

promote healing, and to treat mouth sores (Liogier 

1990). The species is pretty enough to be 

employed as an ornamental, but perhaps because 

of the difficulty of propagation, it has not been 

used so far. 

References 

Croat, T.B. 1978. Flora of Barro Colorado Island. 

Stanford University Press, Stanford, CA. 943 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de 

la Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Edicones, 

Inc. San Juan, PR. 566 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Molano, J.G., M.P. Quicero, and C. Roa. 2002. El 

papel de las cercas vivas en un sistema de 

produccion agropecuaria en el Pidemonte 

Llanero. http://lead.virtualcentre.org/es/ele/ 

conferencia2/vbconfe3.htm. 12 p. 

Stevens, W.D., C. Ulloa-U., and O.M. Motiel, eds. 

2001. Flora of Nicaragua, Angiospermas. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden Press, St. Louis, 

MO. p. 945-1,910. 
477

Miconia prasina (Sw.) DC. camasey blanco 

MELASTOMATACEAE 

Synonyms: Melastoma prasina Sw. 

 Miconia collina DC.

 Acinodendrom prasinum (Sw.) Krasser in Engl. & Prantl. 

 Melastoma parviflora Aublet 

 Melastoma pendulifolia Rich. 

 Miconia parviflora (Aublet) Cogn. in A. DC. 

John K. Francis 

General Description.—Camasey blanco, also 

known as sardine, granadillo bobo, cenizoso, 

mullaca colorado, waraia, santo selele bélétére, 

jacatirão, and mondururu preto, is a shrub or small 

tree usually 2 to 6 m in height. It is an evergreen 

with single or multiple stems arising at or near the 

ground level. Camasey blanco tends to be 

branchy. The sapwood is light brown and the 

heartwood is grayish brown. It is a hard and heavy 

wood with a specific gravity of 0.7. The bark is 

smooth, gray, and thin. The plant is supported by a 

lateral root system with sinkers and a moderate 

amount of fine roots. The leaves frequently have 

winged petioles and leaf blades that are elliptic to 

lanciolate, 10 to 30 cm long, entire or with wavytoothed edges, three- to five-nerved, and pointed 

at both ends. The terminal panicles contain many 

tiny whitish flowers that develop into berries 3 to 

8 mm in diameter and blue to black at maturity. 

The fruits are juicy, slightly sour but mostly 

tasteless. Each fruit contains many minute brown 

seeds (Croat 1978, Howard 1989, Liogier 1995, 

Little and Wadsworth 1964, Stevens and others 

2001). 

Range.—Camasey blanco is native to the Greater 

and Lesser Antilles, Trinidad, Mexico, Central 

America, South America to Bolivia, Brazil, and 

Paraguay (Howard 1989, Killeen and others 1993, 

Liogier 1995, Little and Wadsworth 1964). The 

species is not known to have been planted or 

naturalized elsewhere. 

Ecology.—Camasey blanco grows in secondary 

and remnant forests, brushy pastures, fencerows, 

creek-bottom galleries, coastal thickets, and 

disturbed areas. The sites where it grows are moist 

or wet and sometimes swampy, receiving from 

1600 to 3500 mm of annual precipitation. The 

species occupies elevations from near sea level to 

650 m in Costa Rica (Missouri Botanical Garden 

2002) and up to 1,500 m in Bolivia (Killeen and 

others 1993). Camasey blanco grows in loamy and 

clayey soils derived from both sedimentary and 

igneous rocks. Most of these soils have anaerobic 

subsoils. Leaves collected under a Pinus caribaea

Morelet plantation had 2.34 percent N, 0.13 

percent P, 0.90 percent K, and 10.29 percent ash; 

were higher in N, P, and K, but lower in ash than 

these leaves of plants from nearby natural forest of 

similar age (Lugo 1992). Camasey blanco is a 

shade intolerant pioneer species. It was the second 

most abundant species 7 years following 

destruction of forests in Nicaragua by Hurricane 

Juana (Granzow-de la Cerdal and others 2002). In 

Puerto Rico, the species appears to invade active 

pastures, taking advantage of disturbance by cattle 
478 

and persists as small individuals until the pastures 

are abandoned, after which they grow rapidly and 

dominate for 10 to 20 years (Pascarella 2002). It 

usually grows as dispersed individual shrubs or 

small clumps. Camasey blanco disappears in a few 

seasons after being overtopped by a forest canopy. 

Reproduction.—Camasey blanco flowers from 

February to June and fruits from April to 

September in Nicaragua (Stevens and others 

2001). The species is reported to flower and fruit 

twice each year in Panama (Croat 1978). In Puerto 

Rico, it is reported to flower and fruit almost 

throughout the year (Little and Wadsworth 1964). 

Fruits collected in Puerto Rico averaged 0.137 ± 

0.003 g/fruit. Seeds separated from them averaged 

10.6 million seeds/kg. Sown on wet peat, 57 

percent of the seeds germinated between 20 and 

63 days after sowing. The seedlings are tiny and 

difficult to manage. In nature, birds are the 

principal dispersers of seeds. Camasey blanco 

sprouts readily when disturbed. 

Growth and Management.—The largest 

camasey blanco recorded in Puerto Rico had a 

diameter at breast height of 16.3 cm and a height 

of 11 m (Puerto Rico Champion Tree List on file 

at the International Institute of Tropical Forestry, 

Río Piedras, PR). The species has a moderate 

growth rate and is relatively short lived. None of a 

group of 24 camasey blanco stems in a survey in 

Puerto Rico were present 24 years later at a 

second visit (Weaver 1979). However, by 

suckering and ground layering, plants can 

perpetuate themselves somewhat longer than 

individual stems. Nursery propagation of the 

species has not been reported. 

Benefits.—Camasey blanco wood is not durable 

in the ground and therefore not suitable as fence 

posts (Little and Wadsworth 1964), unless treated. 

Penetration of preservative was about average for 

52 Puerto Rican species tested (Englerth and 

Goytía-Olmedo 1960). The wood is used for fuel 

and occasionally as tool handles. Wood specific 

gravity of 30 plants sampled in Puerto Rico was 

measured at 0.630 + 0.021. A weighted average of 

carbon content for above-ground parts was 50.0 

percent. Total above-ground biomass in grams 

may be predicted by multiplying the stem 

diameter in cm squared at 30 cm above the soil 

times stem length in meters by 38.344 (R2

 = 

0.981) (Francis 2000). The fruits are edible but 

seldom eaten by people. Camasey blanco is 

browsed sparingly by cattle and more heavily by 

goats. The fruits are eaten by the Puerto Rican 

birds, Mimocichla ardosiaceae portoricensis

Bryant, Vircosylva calidris calidris L., Spindalis 

portoricensis Bryant (Whetmore 1916) and 

certainly many other species across the range. 

Camasey blanco serves as a transitional 

(successional) species between pasture, farmland, 

or disturbed forest and secondary forest. 

References 

Croat, T.B. 1978. Flora of Barro Colorado Island. 

Stanford University Press, Stanford, CA. 943 p. 

Englerth, G.H. and E. Goytía-Olmedo. 1960. 

Preservation of Puerto Rican fence posts treated 

by cold soaking and hot-and-cold bath method. 

Tropical Forest Notes 2. U.S. Department of 

Agriculture, Forest Service, Tropical Forest

Recearch Center, Río Piedras, PR. 4 p.

Francis, J.K. 2000. Estimating biomass and carbon 

content of saplings in Puerto Rican secondary 

forest. Caribbean Journal of Science 36(3-4): 

346-350. 

Granzow-de la Cerdal, I., N. Zamora, J. 

Vandermeer, and D. Boucher. 2002. Diversidad 

de especies arbóreas en el bosque tropical 

húmedo del Caribe nicaragüense siete años 

después del huracán Juana. http://www.rbt. 

ac.cr/revistas/45-4/granzow.htm. 15 p. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Killeen, T.J., E. García-E., and D.G. Beck. 1993. 

Guía de árboles de Bolivia. Herbario Nacional 

de Bolivia and Missouri Botanical Garden, La 

Paz and St. Louis, MO. 958 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 617 

p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 
479

Lugo, A.E. 1992. Comparison of tropical tree 

plantations with secondary forests of similar 

age. Ecological Monographs 62: 1-41. 

Missouri Botanical Garden. 2002. Flora of Costa 

Rica: Miconia prasina. http://www.mobot.org/ 

manual.plantas/023023/S024494.html. 2 p. 

Pascarella, J.B. 2002. Tropical forest succession in 

Puerto Rico. http://chiron.valdosta.edu/ 

jbpascar/Research/Prico/puerto_rico.htm. 6 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany, Vol. 85, No. 

2. Missouri Botanical Garden, St. Louis, MO. 

pp. 945-1,910. 

Weaver, P.L. 1979. Tree growth in several tropical 

forests of Puerto Rico. Research Paper SO-152. 

U.S. Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 15 p. 

Whetmore, A. 1916. Birds of Porto Rico. Bulletin 

326. U.S. Department of Agriculture. 

Washington, DC. 140 p. 
480 

Miconia racemosa (Aubl.) DC. camasey de felpa 

MELASTOMATACEAE 

Synonyms: Melastoma racemosa Aubl. 

 Melastoma decussata Vahl 

 Miconia brachypoda DC. 

 Tamonea racemosa (Aubl.) Cook & Collins 

John K. Francis 

General Description.—Camasey de felpa 

(meaning “felt” Miconia in Spanish), also known 

as terciopelo and camasey racimoso, is an 

evergreen shrub or small tree 2 or 3 m in height 

and 3 to 5 cm in stem diameter. The stems are 

smooth and gray with an inner bark of light gray. 

The plants are supported by an ample lateral root 

system. Multiple stems arising from the root 

crown and lower trunk are common. The wood is 

light brown and hard. Twigs are light green and 

somewhat four-angled and have a ring of hairs at 

the nodes. The opposite elliptic leaves have 

petioles 8 to 45 mm long, five main veins from the 

base, a saw-toothed, hairy edge, and are pointed at 

both ends. The inflorescences are panicles with 

many tiny white, pink, or purple flowers. The 

fruits are berries, 3 to 5 mm in diameter, whitish 

when immature, dark blue, purple, or black at 

maturity, and contain numerous tiny brown seeds 

(Howard 1989, Liogier 1995, Little and others 

1974). 

Range.—Camasey de felpa is native to 

Hispaniola, Puerto Rico, Dominica, St. Lucia, 

Granada, Trinidad and Tobago, Venezuela, the 

Guianas, and Brazil to the Atlantic forest in the 

South (Centro Nordestino de Informaçõas Sobre 

Plantas 2002, Fundaçao Andre Tosello 2002, 

Howard 1989, Liogier 1995, Little and others 

1974). 

Ecology.—Camasey de felpa inhabits areas that 

receive 1600 to 3000 mm of annual precipitation. 

Common habitat is on moderately well drained, 

somewhat poorly, and poorly drained, clayey, 

weathered soils, particularly ultisols. Areas of both 

sedimentary and igneous rocks are colonized at 

elevations from near sea level to 900 m in 

elevation. The species is shade intolerant, 

requiring partial sunlight to flower and fruit. 

Disturbance favors reproduction, but this 

requirement does not appear to be absolute. 

Seedlings are frequently found under plantations 

in mid-rotation or trees that have colonized old 

fields. Camasey de felpa may be found in old 

fields, tree plantations, secondary forests, 

roadsides, and landslides (author’s observation, 

Little and others 1974). 

Reproduction.—Camasey de felpa flowers and 

fruits throughout the year (Little and others 1974). 

A collection of fruits from Puerto Rico weighed 

0.1315 ± 0.0041 g/fruit. The seeds are tiny 

amounting to several million per kg. Placed on 

moist filter paper at ambient temperature, 33 

percent germinated within 18 days. The seeds are 

principally dispersed by birds (Devoe 1989). A 

study in Puerto Rico showed predispersal losses of 

seeds to vertebrates and insects accounted for 3 

percent of the seeds, fungi destroyed 9 percent, 

and postdispersal loss to ants was 15 percent 

(Myster 1997). Although seedlings are rarely 
481

abundant, the species is common at least in Puerto 

Rico, growing in small clumps or as scattered 

individuals. Camasey de felpa coppices readily 

when cut. 

Growth and Management.—Camasey de felpa 

has a moderate growth rate and probably lives 10 

to 20 years. No management experience has been 

published. Natural reproduction could be 

encouraged by disturbance of moist and wet forest 

and unwanted plants eliminated by grubbing or 

spraying with broadleaf herbicides. 

Benefits.—Camasey de felpa contributes to the 

aesthetics of forests where it grows, helps 

revegetate disturbed sites and protect the soil, and 

provides food and cover for birds and other 

wildlife. The fruits are juicy and edible but small 

and almost tasteless (Little and others 1974). The 

wood is useful for fuel, small fence posts, and 

stakes. 

References 

Centro Nordestino de Informaçõas Sobre Plantas. 

2002. Checklist das plantas do Nordeste. 

Universidade Federal de Pernambuco. 

http://umbuzeiro.cnip.org.br/db/pnechk/fam/ 

f104.shtml. 6 p. 

Devoe, N.N. 1989. Differential seeding and 

regeneration in openings and beneath closed 

canopy in sub-tropical wet forest. Ph.D. 

dissertation, Yale University, New Haven, CN. 

307 p. 

Fundaçao Andre Tosello. 2002. Base de dados 

tropicles: Espécies arbóreas da Mata Atlântica. 

http://www.bdt.fat.org.br/mata.atlantica/flora/ 

local?SC. 5 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Myster, R.W. 1997. Seed predation, disease and 

germination on landslides in neotropical lower 

montane wet forest. Journal of Vegetation 

Science 8(1): 55-64. 
482 

Mimosa aculeaticarpa Ortega wait-a-bit

FABACEAE 

Synonyms: Mimosa biuncifera Benth.

 Mimosa lindheimeri Gray 

 Mimosa warnockii B.L. Turner 

 Mimosopsis biuncifera (Benth.) Britt. & Rose 

Juanita A. R. Ladyman 

Illustration source: USDA-Forest Service collection, 

Hunt Institute 

General Description.—Mimosa aculeaticarpa is 

a straggling, thicket-forming shrub typically armed 

with curved prickles that are responsible for the 

common names, wait-a-bit, Catclaw mimosa, Una 

de Gato and Gatura. The origin of the genus name 

is likely from the Spanish word Mimoso meaning 

“sensitive”, which refers to the sensitive, 

thigmotactic leaf movement (Barneby 1991). An 

equally appropriate derivation may be from the 

Greek mimos that means “to mimic” and refers to 

the animal-like response of the leaf movements 

(Gledhill 1992, Vines 1986). Although plants may 

have straight prickles or be unarmed, they most 

commonly have two curved spines per node and 

spines on the leaves and pods (Barneby 1991, 

Bowers 1993, Carter 1997, Powell 1998). Shrubs 

are generally 1 to 3m tall with deciduous, divided 

leaves that may have as many as 300 tiny leaflets. 

The species has variable phenotypic and habitat 

characteristics and has been described as 

genetically and ecologically confusing (Carter 

1997). Barneby (1991) recognized four variants of 

Mimosa aculeaticarpa of which var. biuncifera 

(Benth.) Barneby is a particularly common one. 

However, even within each variety he emphasizes 

that substantial variability exists. Varieties 

glabrescens and lindheimeri are synonymous with 

var. biuncifera (Kartesz 1994) and should not be 

confused with the variants described by Barneby 

(1991). 

Range.—Catclaw mimosa occurs in central and 

southern Arizona, southern New Mexico, western 

and central Texas, and northern Mexico in Nuevo 

León, Tamaulipas, Durango, Coahuila, Puebla, 

San Luis Potosí, and Chihuahua (Carter 1997, 

Kearney and others 1960, Martin and Hutchins 

1981, Vines 1986). 

Ecology.—Catclaw mimosa grows at lower 

elevations, between 600 to 1,525 m (2,000 to 

5,000 ft), frequently in limestone and igneous rock 

habitats (Powell 1998). It often becomes abundant 

on dry, rocky, open slopes (Pase and Brown 1982) 

although it can be equally profuse in washes where 

water availability is periodically copious. Catclaw 

mimosa is a member of various communities: 

Desert shrub, Texas savanna, Southwestern shrub 

steppe, Chaparral - mountain shrub, Pinyon – 

juniper woodlands, Plains, and Desert grasslands 

(Dick-Peddie 1993, Cable 1975, Uchytil 1990). 

Many authors have mentioned the importance and 

frequency of low shrubs, such as Catclaw mimosa, 

in a perennial-grass dominated matrix on rocky 

boulder-ridden terrain in Arizona (Shreve 1915, 

Whittaker and Niering 1965), Mexico (Muller 

1939, 1947), New Mexico (Dick-Peddie and Moir 

1970), and Texas (Gehlbach 1967). 

Reproduction.—Flowering dates are variable 

(generally late April to September), and available 

moisture is the limiting factor for flowering 

(Carter 1997). The individual flowers are small 

(<4 mm) and they are massed in a pale-pink to 

whitish globose head. The flowers are fragrant and 
483

are pollinated by insects. Only a few flowers in 

each inflorescence produce pods (Bowers 1993). 

The fruit is an elongated flattened legume (pod), 

the margins separate from the valves at 

dehiscence. The valves often break transversely 

into one-seeded sections. The seeds exhibit high 

germination rates over a relatively wide range of 

temperatures (Jordan and Haferkamp 1989). 

Livestock and wildlife are likely to disperse the 

seeds (Uchytil 1990). 

Growth and Management.—Catclaw mimosa 

forms true brier patches that may become almost 

impenetrable to humans (author’s personal 

observation). Plants can profusely re-sprout from 

protected buds following fire-induced mortality of 

aboveground tissue (Cable 1975, Carmichael and 

others 1978, McPherson 1995). However, in 

general, plants must be several years old before 

they have the ability to re-sprout. Re-sprouting 

decreases as soil-moisture decreases and is less 

common after fires that occur during the growing 

season compared with dormant-season fires 

(McPherson 1995). Stands can recover from 

burning in 5 years (Ahlstrand 1982, Kittams 

1972). However, at least in Arizona, wildfire does 

not appear to stimulate sprouting (Pase and Pond 

1964). Catclaw mimosa is moderately resistant to 

phenoxy herbicides and after only one application 

tend to re-foliate or re-sprout from the crown 

(Hibbert and others 1974). Successive applications 

kill the plants. 

Benefits.—Catclaw mimosa thickets provide 

cover and food for a variety of small wildlife 

species, especially quail (Graham 1941, Powell 

1998). The flowers are a good source of nectar for 

honey bees (Powell 1998, Vines 1986). Deer and 

pronghorn browse Catclaw mimosa and its 

palatability is rated good (Nichol 1938, Buechner 

1950). Its livestock palatability is low (Dayton 

1931). However, livestock use it when other 

forage is scarce, and by virtue of its abundance in 

New Mexico and Arizona it has been described as 

important cattle browse (Dayton 1931, Vines 

1986). Both livestock and wildlife eat the pods 

(Dayton 1931, Cable 1975, Uchytil 1990). 

Because plants have a tendency to form thickets 

that effectively bind soil, Catclaw mimosa shows 

potential for erosion control (Kearney and others 

1960, Vines 1986). In a comparison with 100 other 

plant species studied for the production of fuels 

and chemicals, it yielded substantial amounts of 

oils, polyphenols, and hydrocarbons (Carr and 

others 1986). 

References 

Ahlstrand, G.M. 1982. Response of Chihuahuan 

Desert Mountain Shrub Vegetation to Burning. 

Journal of Range Management. 35 (1): 62-65. 

Barneby, R.C. 1991. Sensitivae Censitae. Memoirs 

of the New York Botanical Garden Vol. 65. 

The New York Botanical Garden, Bronx, NY. 

Bowers, J.E. 1993. Shrubs and Trees of the 

Southwest Deserts. Southwest Parks and 

Monumens Association, Tucson, AZ. 140 p. 

Buechner, H.K. 1950. Life history, ecology, and 

range use of the pronghorn antelope in TransPecos Texas. American Midland Naturalist. 

43(2): 257-354. 

Cable, D.R. 1975. Range management in the 

chaparral type and its ecological basis: the status 

of our knowledge. Research Paper RM-155. 

U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Forest and Range Experiment 

Station, Fort Collins, CO. 30 p. 

Carmichael, R.S., Knipe, O.D., Pase, C.P. and 

Brady, W.W. 1978. Arizona chaparral: plant 

associations and ecology. Research Paper RM202. U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Forest and Range 

Experiment Station, Fort Collins, CO. 16 p. 

Carr, M.E., Mason, C.T., Jr., and Bagby, M.O. 

1986. Renewable resources from Arizona trees 

and shrubs. Forest Ecology and Management.16: 

155-167. 

Carter, J.L. 1997. Trees and Shrubs of New 

Mexico. Johnson Books, Boulder, CO. 533 p. 

Dayton, W.A. 1931. Important western browse 

plants. Miscellaneous Publication 101. U.S. 

Department of Agriculture, Washington, DC. 

214 p. 

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation--past, present, and future. 

University of New Mexico Press, Albuquerque, 

NM. 244 p. 

Dick-Peddie, W.A. and W.H. Moir. 1970. 

Vegetation of the Organ Mountains, New 

Mexico. Range Science Department Science 
484 

Series 4. Colorado State University, Ft. Collins, 

CO. 

Gehlbach, F.R. 1967. Vegetation of the Guadalupe 

Escarpment, New Mexico-Texas. Ecology 48: 

793-806. 

Gledhill, D. 1992. The names of plants. Second 

Ed. Cambridge University Press, Cambridge 

England. 202 p. 

Graham, E.H. 1941. Legumes for erosion control 

and wildlife. Miscellaneous Publication 412. 

U.S. Department of Agriculture, Washington, 

DC. 153 p. 

Hibbert, A.R., Davis, E.A., and Scholl, D.G. 1974. 

Chaparral conversion potential in Arizona: Part 

I: water yield response and effects on other 

resources. Research Paper RM-126. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Forest and Range Experiment 

Station, Fort Collins, CO. 36 p. 

Jordan, G.L. and M.R Haferkamp. 1989. 

Temperature responses and calculated heat units 

for germination of several range grasses and 

shrubs. Journal of Range Management. 42(1): 

41-45. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada and 

Greenland. Vol.1-Checklist. Second ed. Timber 

Press Portland, OR. 622 p. 

Kearney, T.H., Peebles, R.H. and collaborators. 

1960. Arizona flora. 2nd ed. University of 

California Press, Berkeley, CA. 1,085 p. 

Kittams, W.H. 1972. Effect of fire on vegetation of 

the Chihuahuan Desert region. Proceedings: 

Tall Timbers Fire Ecology Conf. A Quest for 

Ecological Understanding 12: 427-444. 

Martin, W. C. and C.R. Hutchins. 1981. A Flora of 

New Mexico. Volume 2. J. Cramer, Vaduz, 

Germany. p. 1,277-2,591. 

McPherson, G.R. 1995. The role of fire in desert 

grasslands. In: M. P. McClaran and T. R. Van 

Devender, eds. The Desert Grassland. The 

University of Arizona Press, Tucson, AZ. 

Muller, C.H. 1939. Relation of the vegetation and 

climatic types in Nuevo Leon, Mexico. 

American Midland Naturalist 21: 687-729. 

Muller, C.H. 1947. Vegetation and Climate of 

Coahuila, Mexico. Madrono. 9: 33-57. 

Nichol, A.A. 1938. Experimental feeding of deer. 

Technical Bulletin 75. Arizona Agriculture 

Experiment Station. 39 p. 

Pase C.P. and F.W. Pond. 1964. Vegetation 

changes following the Mingus Mountain burn. 

Note RM-18. U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Station, Fort 

Collins, CO. 8 p. 

Pase, C.P. and D.E. Brown. 1982. Interior 

chaparral. In: D.E. Brown, ed. Biotic 

Communities of the American Southwest—

United States and Mexico. Desert Plants. 4 (1-

4): 95-99. 

Powell, A.M. 1998. Trees & shrubs of TransPecos and adjacent areas. (First Ed. copyright 

Big Bend Natural History Assoc.) University of 

Texas University Press, Austin, TX 498 p. 

Shreve, F. 1915. The vegetation of a desert 

mountain range as conditioned by climatic 

factors. Publication 529. Carnegie Institute, 

Washington, DC. p. 1-45. 

Uchytil, R.J. 1990. Mimosa biuncifera. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/database/feis/. 

Vines, R.A. 1986. Trees, shrubs, and woody vines 

of the Southwest. University of Texas Press, 

Austin, TX. 1,104 p. 

Whittaker, R.H. and W.A. Niering. 1965. 

Vegetation of the Santa Catalina Mountains, 

AZ: a gradient analysis of the south slope. 

Ecology 46: 429-452. 
485

Mimosa arenosa (Willd.) Poir. tepehuiste 

FABACEAE 

 

Synonyms: Acacia arenosa Willd. 

 Acacia malacocentra Mart. 

 Mimosa caudero L. 

 Mimosa leiocarpa DC. 

 Mimosa malacocentra (Mart.) Benth. 

 Mimosa xantholasia Benth. 

John K. Francis 

General Description.—Tepehuiste is a thorny 

shrub or tree usually 3 to 5 m (up to 12 m) in 

height and stem diameters to 15 cm or more. 

Young and undisturbed plants have a single stem 

that may branch several times near the ground. As 

trees become older and heavier, they tend to lie 

down and produce new, vertical sprouts. 

Sometimes, the mid section of the trunk is thicker 

than the base. The hard, somewhat flexible wood 

has light tan sapwood and light brown or brown 

heartwood. The plant is supported by a tap and 

lateral root system of brown, strong, and flexible 

roots. Stems are covered by dark brown, shallowly 

furrowed bark. The inner bark is green. Twigs are 

greenish brown with 3 to 4 mm, curved spines. 

The alternate, compound leaves commonly have 

four to 22 pinnae each with 15 to 35 pairs of 

leaflets. Inflorescences are 6-cm spikes with 

paniculiform branching bearing many tiny, white 

flowers. The fruits, which are borne in clusters, 

are linear-oblong, flat, brown legumes, 4 to 5 cm 

long by 5 to 6 mm broad. The seeds are yellow, 

flattened, and about 4.5 by 5 mm wide and long 

(author’s observations, Liogier 1988, Stevens and 

others 2001). 

Range.—Tepehuiste is native to South America 

from Colombia and Venezuela through Brazil and 

lowland Bolivia. It is possibly native to Mexico 

(recorded in Jalisco State) and Nicaragua. 

Populations in Puerto Rico and the Dominican 

Republic were introduced and have become 

naturalized (Liogier 1988, Missouri Botanical 

Garden 2002, New York Botanical Garden 2002, 

Stevens and others 2001). 

Ecology.—Tepehuiste is intolerant of shade. It 

competes rigorously with grass, herbs, and low 

shrubs but cannot survive overtopping by tall 

trees. The Puerto Rican populations occur in areas 

that receive from 1200 to 1600 mm of mean 

annual precipitation on well-drained to somewhat 

poorly-drained sites. Tepehuiste does not appear 

to compete or survive on rocky, excessively welldrained sites. The species grows at elevations from 

near sea level to 500 m in Nicaragua. Tapehuiste 

grows in catinga (scrub) forest in Brazil, savannas 

in Nicaragua, and vacant lots, roadsides, 

fencerows, and neglected pastures in Puerto Rico 

(author’s observations, Nunez de Medeiros 2002, 

Stevens and others 2001). 

Reproduction.—Tepehuiste blooms June through 

September and fruits September and October in 

Nicaragua (Stevens and others 2001). The author 

observed stands in Puerto Rico in bloom in March 

and other stands with heavy seed crops during that 
486 

same period. Insects pollinate the flowers. Fruits 

collected in Puerto Rico averaged 7.0 + 0.2 

seeds/pod. Seeds separated from them weighed an 

average of 0.0054 ± 0.0006 g/seed or 186,000 

seeds/kg. One percent of unscarified seed 

germinated in 6 months. but a lot of mechanically 

scarified seed germinated at 99 percent between 3 

and 5 days after sowing on moist filter paper 

(author’s observation). Germination is epigeal 

(Parra 1984). Plants sprout when cut or burned. 

Growth and Management.—Tepehuiste grows 

rapidly. Because the species is thorny, forms 

almost impenetrable thickets, and aggressively 

invades pastures and rangeland, it is a highly 

undesirable species in exotic habitats. Every effort 

should be made to prevent it from invading new 

areas. Control techniques have not been published. 

However, the same measures used against Mimosa 

pigra L. probably would be effective. 

Benefits.—Tepehuiste helps sites move through 

the grass stage to secondary forest, protects the 

soil, and provides cover for wildlife. It is an 

important forage plant for goats in northeastern 

Brazil (Nunez de Medeiros 2002). The wood is 

useful for fuel. 

References 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Missouri Botanical Garden. 2002. VAST 

nomenclatural database. http://mobot.mobot. 

org/cgi-bin/search_vast?name=Mimosa 

+arenosa. [not paged]. 

New York Botanical Garden. 2002. Neotropical 

flora mycota catalog. http://scisun.nybg.org: 

8890/searchdb/owa/wwwspecimin.search_list? 

taxon=Mimosa+arenosa&projcode=NETR. [not 

paged]. 

Nunez de Medeiros, A. 2002. Caprinocultura de 

Corte no Nordeste Brasileiro. Departamento de 

Agropecuária, Bananeiras, PB, Brazil. 

http://www.capritio.com.br/art18.htm. [not 

paged]. 

Parra-G., P. 1984. Estudio de la morfologia 

externa de plantulas de Calliandra gracilis, 

Mimosa albida, Mimosa arenosa, Mimosa 

camporum y Mimosa tenuiflora. Revista de la 

Facultad de Agronomía (Maracay) 8(1-4): 311-

350. 

Stevens, W.D., C. Ulloa-U., A. Pool, O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in systematic botany Vol. 85, No. 

2. Missouri Botanic Garden Press, St. Louis, 

MO. p. 945-1,910. 
487

Mimosa ceratonia L. zarsa 

FABACEAE 

Synonyms: Lomoplis ceratonia (L.) Raf. 

John K. Francis 

General Description.—Zarza, also known as 

climbing mimosa, black ambret, amourette-grandbois, grate-jambe, and croc-chien, is a multistemmed scrambling shrub. It supports itself on 

other vegetation by means of curved spines and 

may reach 4 to 6 m into the crowns of trees and as 

far laterally. The plant has a weak taproot, a 

moderate amount of lateral roots, and abundant 

fine roots. Lower stems are gray, scaly, and 

cylindrical. The branches tend to be long, almost 

straight or evenly curved, with few divisions. The 

twigs are green, sometimes reddish with many 

curved 2-mm spines. Lateral twigs usually die 

back after 1 year. The leaves are bipinnately 

compound with three to five pairs of pinnae, each 

with three to five pairs of leaflets. The leaflet 

blades are obliquely obovate, three-nerved, 1.0 to 

2.5 cm long and 6 to 10 mm broad, rounded at the 

apex and asymmetrically acute at the base. The 

inflorescences are terminal racemes of globose 

heads of flowers that are white or cream-colored. 

The pods, which are borne in clusters, are straight 

or slightly curved, oblong, 4 to 6 cm long and 

armed at the edges. The flattened seeds are brown 

or black (Acevedo-Rodríguez 1985, Howard 1988, 

Liogier 1988). 

Range.—The range of zarza includes Hispaniola, 

Puerto Rico, the Virgin Islands, Antigua, 

Guadeloupe, Martinique, St. Lucia, and Barbados 

(Howard 1988, Liogier 1988). Also, populations 

of zarza have been identified in Brazil including 

the varieties M. ceratonia L. var. pseudo-obovata

(Taubert) Barneby and M. ceratonia L. var. 

interior Barneby (Barneby 1985, Paganucci de 

Queiroz 2001). 

Ecology.—Zarza is moderately intolerant of 

shade. It grows in fencerows, roadsides, wooded 

drains, brushy pastures, forest edges and openings, 

and in low-density secondary forest. The species 

often forms compact, nearly impenetrable clumps. 

Diffuse stands mixed with tall grass, weeds, and 

shrubs form in areas that are mowed or burned on 

one- to several-year cycles. Zarza is most common 

on soils derived from limestone and serpentine 

(Vélez and van Overbeek 1950), but also grows on 

soils overlying other sedimentary and igneous 

rocks. It tolerates all textures of well-drained soils 

and a wide range of fertility levels. In Puerto Rico, 

the species may be found in areas receiving 

rainfalls from 750 mm per year to over 2000 mm 

per year. Zarza grows from near sea level to at 

least 600 m in elevation. Frosts do not occur 

within its natural range. 

Reproduction.—Zarsa flowers between June and 

December and fruits December through January in 

Puerto Rico (Acevedo-Rodríguez 1985). A 

collection of zarza pods from Puerto Rico weighed 

an average of 0.1371 ± 0.0005 g (air dry). They 

averaged 4.18 + 0.12 seeds/pod and ranged from 

three to six seeds per pod. The seeds weighed an 

average of 0.0201 + 0.004 g/seed or 50,000 

seeds/kg. Sown on commercial potting mix, 71 

percent germinated between 7 and 165 days after 

sowing. Germination is epigeal. Stems layer (root) 

if covered by soil or organic debris. Stems sprout 
488 

when cut and lateral roots sometimes sucker when 

damaged. The seeds are disbursed by lateral 

extension of the vines and by the pods clinging to 

clothing or to the fur of animals. Seedlings are 

common, but only a few survive for more than a 

few weeks. 

Growth and Management.—Both seedlings and 

sprouts grow rapidly, up to 2 or 3 m per year. 

Zarza plants may endure for many years. Because 

of the thorny nature of this species, property 

owners often choose to eliminate it from pastures 

and plantations. Historically, farmers controlled it 

by repeated cutting. This method could probably 

be improved by treating the resulting sprouts with 

broadleaf weed killers. 

Detriments and Benefits.—Thickets and tangles 

of zarza are almost impenetrable to humans. Paths 

may be cut to allow passage, but they soon grow 

over. Because of the thorns, cattle will not eat 

zarza (Vélez and van Overbeek 1950), but it is 

browsed by goats. The species is a host for the 

diaprepes root weevil (Diaprepes abbreviatus), 

which is of grave concern for growers of citrus 

and other agricultural crops (Simpson and others 

2001). Zarza is a valuable honey plant (Marcano 

1973), protects the soil, and serves as wildlife 

cover. 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Barneby, R.C. 1985. The identity and synonymy 

of Acacia guiladinae, Mimosa obovata, M. 

pseudo-obovata and M. laticifera (Mimosaceae). 

Brittonia 37(1): 85-87. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Marcano F., E. de J. 1973. La flora apícola de la 

República Dominicana. http://naturalista. 

virtualave.net/estudios/apicola/dicotse.html. 8 p. 

Paganucci de Queiroz, L. 2001. Leguminosas da 

caatinga da Bahia com potencial forrageiro. 

http://umbuzeiro.cnip.org.br/db/forrag/fam/ 

f19.html. 6 p. 

Simpson, S.E., H.N. Nigg, and J.L. Knapp. 2001. 

Host plants of diaprepes root weevil and their 

implications to the regulatory process. Florida 

Department of Agriculture and Consumer 

Services. http://www.fcprac.ifas.ufl.edu/ 

citrustopics/pest%20control/Diaprepes/.../simps

on.hostplant.ht. 16 p. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria. Río Piedras, PR. 497 p.
489

Mimosa pigra L. black mimosa 

FABACEAE 

Synonyms: Mimosa asperata L 

John K. Francis 

General Description.—Black mimosa is also 

known as catclaw mimosa, thorny sensitive plant, 

giant sensitive plant, bashful plant, giant mimosa, 

dormilona, pigra, agüiste, amouretto, amouretterivière, and banglin. It is a thorny upright to 

scrambling shrub 2 to 4 m in height and up to 12 

cm in basal diameter. The plant usually has a 

single stem from the ground but may branch near 

the ground and form an inverse cone-shaped 

crown. There are relatively few branches. The 

shrub is supported by extensive lateral roots with 

numerous fine roots that have occasional nodules. 

Stems, limbs, twigs, and leaves bear curved thorns 

up to 7 mm long. Black mimosa produces a thin 

crown of touch-sensitive bipinnately compound 

leaves. Each leaf has four to 14 pairs of pinnae 

and 20 to 40 pairs of linear-oblong leaflets per 

pinna. The inflorescences are tight, subglobose 

mauve or pink heads with about 100 flowers, 

grouped one to three in the upper axils. The 

clustered brown legumes are densely bristled, 4 to 

12 cm long, and breaking transversally into 14 to 

26 segments, each containing one seed. The seeds 

are flattened, about 6 by 2.5 mm, and brown to 

olive green (Howard 1988, Pacific Island 

Ecosystems at Risk 2002, Stevens and others 

2001). 

Range.—The original range of black mimosa is 

uncertain. The contention that the native range 

extends from Mexico to northern Argentina 

(Stevens and others 2001) is supported by the fact 

that 440 species of insects and several fungi 

species keep it under control there (CSIRO 2002). 

On the other hand, a rich native tradition supports 

Equatorial Africa as being native range (Burkhill 

1995). Perhaps it crossed the South Atlantic in 

prehistoric times or early in the colonization of the 

New World. More recently, the species has 

invaded most humid tropical and subtropical areas 

including Florida, Texas, and Puerto Rico (Natural 

Resources Conservation Service 2002). There are 

two varieties: var. berlandieri (Gray ex Torr.) B.L. 

Turner that occurs from the Texas to Costa Rica, 

and var. pigra that grows from Mexico to 

Argentina, Africa, and in the invaded ranges in the 

New and Old Worlds (Stevens and others 2001). 

Ecology.—Presettlement populations of black 

mimosa in Costa Rica were found mainly on the 

edges of marshes and on large river floodplains. 

Today, they are found along roads and in marshy 

spots in pastures (Janzen 1983). The species is 

highly intolerant of shade. It requires full sunlight 

to flower and fruit and nearly full sunlight to 

survive. The two principal ingredients for 

successful establishment are moist or wet soils and 

disturbance that allows full sunlight. It is a fierce 

competitor with low vegetation and one of the few 

shrubs capable of succeeding in dense, tall grass 

swards. Black mimosa tolerates a wide variety of 

soil conditions, short-term flooding, seasonal 

drought, and grows at elevations from near sea 

level to 700 m (Janzen 1983). Although it grows 

mixed with other vegetation in its native habitat, 

the species forms pure stands with little understory 

in many of its exotic habitats (Pacific Island 

Ecosystems at Risk 2002). 

Reproduction.—Black mimosa blooms 
490 

continuously in moist habitat, intermittently in 

seasonally dry habitat. Bees pollinate the flowers 

and pods mature about 1 month after pollination 

(Binggeli 1997). Black mimosa grows rapidly and 

can begin producing seed in as little as 3 months 

after emergence (Department of Natural Resources 

and Mines 2002). One average plant can produce 

more than 9,000 seeds annually (Marko 2002). 

The pod segments disperse by floating on water 

and clinging to clothing and possibly animal fur 

and feathers. A sample of black mimosa pods in 

Puerto Rico contained an average of 15.0 

seeds/pod. The seeds weighed an average of 

0.0166 g/seed. Sown on commercial potting mix 

between 6 and 20 days, 81 percent of the seed 

germinated. Germination is epigeal (author’s 

observation). Scarification is required for quick 

and uniform germination. In an experiment with 

several treatments of temperature and storage, 

germination percentages of 75 to 94 percent were 

observed (Creager 1992). Seeds can remain viable 

in dry soil for more than 2 years. Plants sprout 

freely following fires (Binggeli 1997). Layering 

occurs when stems become prostrate and covered 

with litter. 

Growth and Management.—Black mimosa 

grows rapidly. Seedlings grown in a greenhouse 

reached 25.8 ± 0.5 cm in 30 days, 97.1 + 16.2 cm 

in 90 days, and 335.1 ± 15.9 cm in 360 days 

(Swarbrick and Mercado 1987). Plants in Puerto 

Rico in tall, dense grass swards and disturbed 

upland sites grow about 1.0 to 1.5 m the first year, 

1 m in the second year, and usually live 2 to 3 

years. Larger plants tend to lie down due to weight 

and often do not become taller than 3 m (author’s 

observation). Maximum age is about 5 years 

(Binggeli 1997). In an effort to control the 

aggressively spreading infestation, nine insect 

enemies and two pathogenic fungi have been 

released in Australia. Most have become 

established but so far have had little effect on 

black mimosa populations (Marko 2002). 

However, a recent release of Malacorhinus 

irregularis, a defoliater beetle from Mexico shows 

early promise (CSIRO 2002). Mowing is 

ineffective (Creager 1992). Grubbing of adult 

plants achieved only about 82 percent mortality 

(Cross and Wiedemann 1997). Until effective 

biological control can be achieved, chemical 

control is probably the best approach for spot 

infestations (Marko 2002). 

Benefits and Detriments.—Black mimosa 

invades dense grass swards and in some 

environments helps them move on to the brush and 

secondary forest stages. It also protects the soil in 

disturbed sites. Although the cover and diversity it 

furnishes is probably beneficial for wildlife in 

artificially maintained exotic grass swards, the 

species can reduce wildlife populations in natural 

swards such as river floodplains in Australia 

(Marko 2001). The plant has been used as a green 

manure, a cover crop, for fuel wood, and as 

beanpoles. Although cattle and horses generally do 

not eat black mimosa because of the thorns 

(Binggeli 1997), with 22 percent foliar protein, it 

was found to be perfectly acceptable to rabbits 

(Lebas and others 1986). Black mimosa can be 

weedy in row crops and plantations, and a serious 

problem in pastures and hay fields. The thorny 

branches make it difficult for people to walk 

through fields and along trails where it grows. In 

Africa, it is planted as a barrier around fields and 

cattle enclosures to discourage human intruders 

and animal predators. The species is used in a 

number of herbal remedies and magic rites in 

Africa (Burkill 1995). 

References 

Binggeli, P. 1997. Mimosa pigra L. (Mimosaceae). 

http://members.lycos.co.uk/WoodyPlantEcology

/docs/web-sp11.htm. 5 p. 

Burkill, H.M. 1995. The useful plants of West 

Tropical Africa. Vol. 3. Royal Botanic Gardens, 

Kew, UK. 857 p. 

CSIRO. 2002. Mimosa under attack. Global Net 

News.com. http://ausissues.com/ausissues/ 

AINews.nsf/0/DE44CB72D21850A6CA256C10

00169466?OpenDocument. 4 p. 

Creager, R.A. 1992. Seed germination, physical 

and chemical control of catclaw mimosa 

(Mimosa pigra var. pigra) Weed Technology 

6(4): 884-891. 

Cross, B.T., and H.T. Wiedemann. 1997. Control 

of catclaw acacia and mimosa by grubbing. 

Applied engineering in Agriculture 13(2): 291-

293. 

Department of Natural Resources and Mines. 

2002. NR & M acts on Mimosa pigra outbreak 

at Peter Faust Dam, Proserpine. Department of 

Natural Resources and Mines, Queensland, 

Australia. http://www.nrm.qld.gov.au/about/ 

media/apr/mimosa.html. 4 p. 
491

 Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University. Jamaica 

Plain, MA. 673 p. 

Janzen, D.H. 1983. Mimosa pigra (zarza, 

dormilona). In: D.H. Janzen, ed. Costa Rican 

natural history. University of Chicago Press, 

Chicago and London. p. 277-278. 

Lebas, F., P. Coudert, R. Rouvier, and H. de 

Rochambeau. 1986. The rabbit: husbandry, 

health and production. Food and Agriculture 

Organization of the United Nations, Rome. 

http://fao.org/docrep/x5082e/X5082E06.htm. 

[not paged]. 

Marko, M. 2002. Controlling invasion of the 

exotic shrub (Mimosa pigra) in tropical 

Australia wetlands. University of Minnesota 

Department of Horticultural Science. 

http://www.hort.agri.umn.edu/h5015/99papers/

marko.htm. 10 p. 

Natural Resources Conservation Service. 2002. 

Plants profile: Mimosa pigra L. black mimosa. 

http://plants.usda.gov/cgi_bin/plant_profile.cgi?

symbol=MIPI. [not paged]. 

Pacific Island Ecosystems at Risk. 2002. Mimosa 

pigra L., Fabaceae. http;//www.hear. 

org/pier_v3.3/mipig.htm. 3 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden Press, St. Louis, 

MO. p. 945-1,910. 

Swarbrick, J.T. and B.L. Mercado. 1987. Weed 

science and weed control in Southeast Asia. 

FAO Plant Production and Protection Paper 81. 

Food and Agriculture Organization of the United 

Nations, Rome. 203 p. 
492 

Mimosa pudica L. sensitive plant 

FABACEAE 

Synonyms: Mimosa tetrandra Humb. & Bonpl. ex Willd. 

 Mimosa pudica L. var. tetrandra (Willd.) DC. 

 Mimosa unijuga Duch. & Walp. 

 Mimosa pudica L. var. unijuga (Duch. & Walp.) Griseb. 

John K. Francis 

General Description.—Sensitive plant is a small, 

prostrate or ascending, short-lived shrub. Some 

authors consider it a woody herb. It may reach 1 m 

in height when supported on other vegetation and 

more than 2 m in horizontal extension. The 

reddish-brown, woody stems are sparsely or 

densely armed with curved prickles. The root 

system consists of a taproot and extensive fibrous 

roots with nodules. The twigs are fine and flexible 

and support leaves with one or two pairs of pinnae 

and 15 to 25 pairs of oblong leaflets 3 to 12 mm 

long. The flowers are pink and clustered in 

globose heads. The legume (pod) is linear-oblong, 

1.0 to 1.5 cm long and 3 mm broad, with bristles 

on the margins. The pods are born in groups and 

contain two to four brown seeds (Howard 1988, 

Liogier 1988, Pacific Island Ecosystems at Risk 

2001). Sensitive plant is also known as dorme 

dorme, dormidera, humble plant, marie-honte, 

mayhont, morivivi, honteuse, sleeping grass, ti 

mawi, touch-me-not, and many other names 

(Holm and others 1977, Howard 1988, Liogier 

1988). The great curiosity of sensitive plant and 

the source of most of its names is that when 

touched, it quickly folds its leaflets and pinnae and 

droops downward at the petiole attachment. The 

leaves also droop at night, and when exposed to 

rain or excessive heat. This response may be 

defenses against herbivorous insects, leaching loss 

of nutrients, or desiccation. 

Range.—Sensitive plant was first described from 

Brazil (Pacific Island Ecosystems at Risk 2001) 

and is perhaps native to much or all of the New 

World Tropics (Liogier 1988). Today, it is 

pantropical in its distribution (Howard 1988). 

Ecology.—Sensitive plant grows on most welldrained soils, even scalped or eroded subsoils and 

soils with low nutrient concentrations. It requires 

disturbed soils to establish itself. Repeated 

burning may encourage its spread in pastures 

(Siregar and others 1990). Sensitive plant is shade 

intolerant and does not compete with tall 

vegetation or grow under forest canopies. The 

species’ roots produce carbon disulfide, which 

selectively inhibits colonization of the rhizosphere 

by mycorrhizal and pathogenic fungi (Feng and 

others 1998). This plant occurs in croplands, 

orchards, pastures, mowed areas, roadsides, and 

areas disturbed by construction. It may grow as a 

single plant or in tangled thickets. Sensitive plant 

grows from near sea level up to 1,300 m in 

elevation (Holm and others 1977) and in areas 

with annual precipitations from about 1000 to over 

2000 mm. The species is frost-sensitive. 

Reproduction.—In the Philippines, sensitive 

plant flowers all year and may produce as many as 

675 seeds per plant per year (Holm and others 

1977). The species is both wind (Chieng and 

Huang 1998) and bee-pollinated (Payawal and 

others 1991). Air-dry seeds from Puerto Rico 

weighed an average of 0.0065 ± 0.0002 g/seed. 

With no pretreatment, seeds from this collection 

began germinating 7 days after sowing and 

reached a maximum germination of 17 percent by 

94 days (author’s observation). In another test, 80 

percent germination was obtained in 4 weeks with 

alternating temperatures of 20 and 40 °C (Holm 

and others 1977). Bui (2001) recommends a 

pretreatment with hot water followed by overnight 
493

soaking. Germination is epigeal. Seeds are 

transported by means of the bristles on the edges 

of their pods that cling to clothing or to the fur of 

mammals. Most nursery and home propagation is 

done using seeds, but summer cuttings may also 

be used (Bui 2001). 

Growth and Management.—In Puerto Rico, 

sensitive plants live 1 to 2 years. Seedlings grow 

slowly for 2 or 3 months and then accelerate, 

reaching 0.5 to 2 m of extension at the end of the 

first year. Growth of plants that survive into the 

second year is much slower. Potted and fieldgrown individuals are sensitive to overwatering 

(Bui 2001). This species has been successfully 

tested and recommended for erosion control 

plantings using potted material at a spacing of 60 

x 60 cm (Coimbra and Magnanini 1953). 

Benefits and Detriments.—Sensitive plant has 

become a serious weed in fields of corn, soybeans, 

tomatoes, upland rice, cotton, bananas, sugarcane, 

coffee, oil palms, papayas, coconuts, and rubber in 

many tropical areas. It is particularly troublesome 

where hand pulling of weeds is practiced. The 

species may be controlled by a number of 

commercial broad-leaf herbicides (Bui 2001). On 

the other hand, it is tolerated or valued as a forage 

plant in pastures (Holm and others 1977, Turbet 

and Thuraisingham 1948). In fact, sheep grazing 

is reported to control sensitive plant in pastures 

and plantations (Simonnet 1990). The root nodules 

have been shown to fix nitrogen (Pokhriyal and 

others 1990). Thickets of sensitive plant may be a 

fire hazard when dry (Pacific Island Ecosystems at 

Risk 2001). The seeds and other plant parts of 

sensitive plant contain mimonsine, an amino acid 

that is known to cause hair loss and depressed 

growth in mammals (Arora 1983). An unlikely 

large dose is necessary to cause problems, 

however. The pollen is important to honeybees in 

the Philippines (Payawal and others 1991). 

Extracts of the plant have been shown in scientific 

trials to be a moderate diuretic, depress duodenal 

contractions similar to atropine sulphone, promote 

regeneration of nerves, and reduce menorrhagia 

(Modern-natural 2001). Anitdepressant activity 

has been demonstrated in humans (Martínez and 

others 1996). Root extracts are reported to be a 

strong emetic (Guzmán 1975). 

References 

Arora, S.K., ed. 1983. Chemistry and biochemistry 

of legumes. Edward Arnold (Publishers) 

Limited, London. 358 p. 

Bui, L-D. 2001. Mimosa pudica. http://bio. 

maimi.edu/mimosa/mimosa.html. 7 p. 

Chieng, H-T. and T.C. Huang. 1998. Aeropollen 

of the Pingtung area, South Taiwan. Taiwania 

43(2): 73-100. 

Coimbra, A.F. and A. Magnanini. 1953. 

Considerations sobre Mimosa pudica no 

combate a arosao superficial. Anucio Brazilero 

Economico da Floresta 6. Instituto Nacional, 

Pinho, Brazil. 131-136. 

Feng, Z., P.G. Hartel, R.W. Roncadori, S.J.S. 

Sung, and J.E. Box. 1998. Inhibition of fungal 

colonization on the rhizoplane of the CS2-

production plant, Mimosa pudica L. In: Plant 

and Soil Sciences 82. Kluwer Academic 

Publishers, Dordrecht, Netherlands. 115-126. 

Guzmán, D.J. 1975. Especies utiles de la flora 

salvadoreña. Ministerio de Educación, Dirección 

de Publicaciones. San Salvador, El Salvador. 

703 p. 

Holm, L.G., D.L. Plucknett, J.V. Paucho, and J.P. 

Herberger. 1977. The world’s worst weeds. 

East-West Center, University of Hawaii, 

Honolulu, HI. 609 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain, 

MA: Arnold Arboretum, Harvard University. 

673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Martínez R., G., F.F. Rodríguez L., C.M. 

Contreras, and M. Molina H. 1996. Estudio 

preliminary de las possibles acciones 

antidepresivas de Mimosa pudica L. In: 

Resumen de ponencias del Primer Congreso 

Nacional de Plantas Medicinales de Mexico. 

p. 69. July 24-30, 1996. Tlaxcala, Tlaxcala, 

Mexico. 

Modern-natural. 2001. Mimosa pudica Linn. 

http://modern-natural.com/mimosa_pudica.htm. 

4 p. 
494 

Pacific Island Ecosystems at Risk. 2001. Invasive 

plant species: Mimosa pudica Mart. ex Colla, 

Fabaceae. http://www.hear.org/pier/mipud.htm. 

2 p. 

Payawal, P.C., A.C. Tilde, and A.L. Manimtim. 

1991. Year round pollen sources of Italian honey 

bees (Apis mellifera L.) in the Philippines. III. 

Selected areas. Philippine Agriculturist 74(4): 

503-509. 

Pokhriyal, T.C., H.C.S. Bhandari, D.S. Negi, S.P. 

Chaukiyal, and B.B. Gupta. 1990. Identification 

of some fast growing leguminous tree species 

for nitrogen fixation studies. Indian Forester 

116(6): 504-507. 

Simonnet, P. 1990. Sheep flock management in a 

tropical environment under coconut. 

Oleagineux Paris 45(10): 451-456. 

Siregar, M.E., B. Haryanto, and S. Tjitrosemito. 

1990. A review of weed management in 

Indonesian pastures. BIOTROP Special 

Publication 38. A symposium on weed 

mangement. 7-9 June 1989. Bogor, Indonesia. p. 

229-235. 

Turbet, C.R. and K. Thuraisingham. 1948. Feeding 

trials with the sensitive plant Mimosa pudica. 

Tropical Agriculturist, Ceylon 104(2): 81-86. 
495

Mimulus aurantiacus W. Curtis bush monkeyflower

PHRYMACEAE (formally in SCROPHULARIACEAE) 

Synonyms: Diplacus aurantiacus Jeps. 

 Mimulus glutinosus Wendl. 

 Diplacus latifolius

 Diplacus leptanthus Nutt. 

Arlee M. Montalvo and Paul A. McMillan 

General Description.—Bush monkeyflower, or 

sticky monkeyflower, is a beautiful, erect to 

sprawling subshrub, that is named for its woody 

habit and typical “monkey-faced” flowers. Bush 

monkeyflower represents a complex assemblage of 

hybridizing forms with intergrading floral colors 

and morphologies. For this reason, the most recent 

taxonomic treatment lumps together all the species 

of woody monkeyflowers from California, 

excepting M. clevelandii Brandegee, under M. 

aurantiacus (Thompson 1993). The various taxa, 

sensu Munz and Keck (1968), are distinguished by 

stature, flower size and color, and range, but share 

similar leaf morphology. Given the taxonomic 

uncertainty, we describe M. aurantiacus sensu 

Munz and Keck. Then under “Range and 

Taxonomy,” we provide a brief description of 

diagnostic differences and distributions of their 

other taxa that were treated as a single species 

complex by Thompson (1993). 

Shrubs are erect, 6 to 12 dm tall, and the 

specific epithet “aurantiacus” refers to the often 

deep orange corollas (sometimes yellowish) that 

are 3.5 to 4.5 cm long. The simple, opposite, 

leaves are elliptic to narrowly-elliptic, with entire 

to serrulate margins that are often rolled under. 

Leaves are 2 to 8 cm long and dark green, and 

resinous above. Smaller leaves are clustered on 

shortened shoots in the axils of larger leaves. The 

lighter underside of leaves and the 2 to 2.5 cm 

long green, pleated calyx are covered with sticky 

glandular and nonglandular hairs. Flowers have a 

long floral tube, spreading limbs, and a large, twolobed stigma. Plants are often sticky with 

glandular secretions. 

Range and Taxonomy.—Recent phylogenetic 

work favors placement of woody monkeyflowers 

(section Diplacus) in Mimulus and transfers the 

genus from family Scropulariaceae to the 

Phrymaceae (Beardsley and Olmstead 2002). Bush 

monkeyflower (broad sense) ranges from the coast 

to coastal ranges to interior foothills from 

northwestern Baja California, northward to 

Mendocino County, California below 1,600 m and 

along the western foothills of the Sierra Nevada 

range. The underlying topography of coastal hills, 

interior valleys, and foothills is complex, as are the 

west to east and north to south gradients in 

temperature and rainfall. The correspondingly 

complex pattern of variation in bush 

monkeyflower may reflect once isolated and 

differentiated populations that have come into 

secondary contact and hybridized. The resulting 

patterns of genetic diversity do not match the 

patterns of morphological diversity (Beardsley and 

others 2003). The taxonomy is necessarily 

complex and may be resolved with detailed 

phylogeographic analysis. See Munz and Keck 

(1968) and Tulig (2000) for details of the 

geographic distribution of the following color 

forms (narrow sense). 

The small, orange-flowered M. aurantiacus 

occurs in the foothills and coastal ranges north of 

Santa Barbara County. M. a. ssp. australis

(McMinn) Munz has orange-yellow to buff or 

white corollas and occurs in interior San Diego 

County to Baja California. M. aridus (Abrams) 

A.L. Grant, the lowbush monkeyflower, has a 

yellow corolla, low, spreading habit to 4 dm tall, 

and occurs farther east in San Diego County to 

Baja Calfornia. M. bifidus Pennel has a larger 
496 

orange corolla with larger, deeply-notched lobes, a 

somewhat spreading habit to 1 m tall, and occurs 

from coastal San Luis Obispo to Monterey County 

and along the western base of the Sierra Nevada 

from Placer into Butte and Plumas Counties. M. 

flemingii Munz has red corollas with limbs about 

half as long as other forms, is low growing from 1 

to 6 dm tall, and is restricted to the Channel 

Islands. M. longiflorus (Nutt.) A. L. Grant, the 

southern bush monkeyflower (see figure), has an 

orange-yellow to salmon corolla, a long and 

pubescent calyx (to 3.5 cm), is 3 to 12 dm tall, and 

occurs from central San Diego County through 

Riverside to San Bernardino County. A shorter, 

pale-yellow form [M. l. ssp. calycinus (Eastw.) 

Munz] occurs primarily farther inland in Riverside 

and western San Bernardino Counties. A redcorolla form (M. l. var. rutilus Grant) occurs 

mostly in interior Los Angeles County. Finally, M. 

puniceus (Nutt.) Steudel, the red bush 

monkeyflower, has a red corolla, short calyx (to 

2.5 cm), and erect habit from 5 to 15 dm tall. It 

occurs on Catalina Island, coastal Orange County, 

and inland through the Santa Ana Mountains to 

southwestern Riverside County south into Baja 

California. 

Hybridization.—All bush monkeyflowers have n 

= 10 chromosomes and crosses among them 

produce viable progeny. Natural, putative hybrids 

between forms have been hypothesized based on 

morphology (Munz and Keck 1968, Wells 1980, 

Grant 1993, Tulig 2000). Preliminary phylogenetic 

work suggests widespread introgression among 

forms (personal communication with Paul 

Beardsley, Colorado College, CO). Grant (1993) 

examined pollinators in relation to several floral 

forms (equals species of Diplacus in his paper) and 

postulated that the populations with different floral 

forms would be partially reproductively isolated. 

He hypothesized that the red-colored D. puniceus

Nutt. and salmon D. longiflorus Nutt. were 

primarily hummingbird-pollinated, and that D. 

calycinus Eastw. (M. l. var. calycinus) with pale 

yellow flowers and long tube) were hawkmothpollinated. He found that habitats interdigitated 

and resulted in D. puniceus and D. calycinus

populations overlapping in distribution, with D. 

longiflorus habitat forming a connecting link 

between the different forms. He hypothesized that 

the intermediate nature of D. longiflorus suggested 

past hybridization of D. puniceus and D. calycinus. 

Tulig (2000) analyzed floral and vegetative 

variation throughout the geographic range of bush 

monkeyflowers and found statistical evidence for 

hybrid populations that were intermediate between 

parental forms. For example, in San Diego County, 

putative hybrid populations were intermediate 

between M. puniceus (coastal form) and M. 

aurantiacus ssp. australis (inland form), consistent 

with analysis of Waayers (1996). In the region 

between the San Gabriel and San Bernardino 

Mountains, putative hybrids were intermediate 

between M. longiflorus ssp. calycinus (lemon 

yellow corolla) and M. longiflorus (salmon 

corolla). 

Ecology.—Bush monkeyflowers occur in areas 

with cool moist winters and warm to hot, dry 

summers. They grow on rocky hillsides and cliffs, 

usually on the border of chaparral or sage scrub, or 

in open foothill-woodland forest. The roots are 

fibrous, less than 2 m deep, with feeder roots 

concentrated within the first 8 cm of soil (Hellmers 

and others 1955). There is evidence for 

differentiation and local adaptation in 

morphological and physiological characters 

relating to water use. For example, differences in 

wood anatomy among forms and among 

populations are correlated with habitat (Michener 

1983). Plants from mesic habitats have relatively 

few, large vessel-elements, while plants from xeric 

habitats have many, small vessel-elements. Under 

humid, low-stress conditions, both mesic-adapted 

and xeric-adapted populations had high water-use 

efficiency and maintained a high rate of 

photosynthesis relative to transpiration (Mooney 

and Chu 1983). Plants from xeric populations had 

high water-use efficiency under low relative 

humidity, while the efficiency of coastal plants 

diminished rapidly with decreasing relative 

humidity.

The primary herbivore of bush monkeyflower 

is a specialist butterfly Euphydryas chalcedona

Doubleday and Hewitson. The timing of growth of 

both plants and the butterfly is linked to water 

availability. Emergence of larvae from diapause is 

synchronized with the initiation of leaf growth. 

The leaves produce a resin that inhibits the growth 

of the larvae. During the growing season the 

youngest leaves have the highest nitrogen (N) 

content, the highest carbon gain, and the highest 

resin content. Growth rate of the larvae increases 

with increasing N but decreases with increasing 

resin content (Lincoln and others 1982). Larvae 

feed initially on young leaves with high nutritional 

value, then switch to older leaves with lower resin 

content. During flowering, N is translocated from 

the leaves, and larvae stop feeding (Mooney and 

others 1981). 
497

Such studies demonstrate physiological 

trade-offs in resource allocation to growth, 

reproduction, or defense. Han and Lincoln (1994) 

found significant heritability and maternal effects 

for resin content within one population, and they 

found negative genetic and phenotypic correlations 

between resin production and growth rate. This 

indicates potential trade-offs between traits during 

selection. Leaf resin content varies among 

populations, but little is known about geographic 

differentiation in such trade-offs. 

Hare (2002) found that nearly 30 percent of 

the dry weight of leaves is resin. In a common 

garden experiment, he examined the chemistry and 

relative quantities of important resin components 

for six populations of bush monkeyflower from a 

range of environments and that differed in attack 

by E. chalcedona. The populations differed in 

chemical components of the resins but not in the 

pattern of insect attack. This study followed 

Thompson’s (1993) classification and did not 

mention if there were differences in floral 

morphology among the populations. The collection 

locations could have represented at least three 

species or subspecific taxa following Munz and 

Keck (1969) or Tulig (2000). 

Emergence and Growth.—Seedlings emerge in 

the cool rainy season, and most growth is in winter 

and spring. Plants can reach reproductive maturity 

in a single growing season, with flowering 

primarily from mid spring to early summer. The 

capsules mature in summer, and the many, tiny 

seeds (about 1 mm long, half as wide and nearly 

flat) disperse by gravity and wind. Plants become 

dormant in the summer dry season and shed many 

leaves by fall. Leaf production begins after the 

first fall rain and ends near the beginning of July 

(Mooney and others 1981). A large and relatively 

constant proportion of carbon and nitrogen are 

allocated to reproduction, and reproductive 

structures supply some of their own carbon (Alpert 

and others 1985). Both growth and reproduction 

are primarily water limited. 

Reproduction.—Bush monkeyflowers are 

pollinated by diverse flower visitors, and the 

dominant pollinators vary with color form and 

location. This may reflect local differentiation, or 

the relative abundances of pollinators. Fetscher 

and Kohn (1999) reported Anna’s hummingbird as 

the primary pollinator of a red flower form in San 

Diego County, California, with black-chinned, 

Costa’s, and rufous hummingbirds as occasional 

visitors. Bromer and others (1990) reported that M. 

longiflorus was “generally but not exclusively” 

pollinated by Black-chinned and Anna's 

hummingbirds. Six populations from the Santa 

Monica Mountains varied in floral form, sucrose 

content of nectar, and anther-stigma separation. 

Populations with high pollinator diversity but low 

visitation rates had shorter corolla tubes and lower 

sucrose:hexose ratios in nectar than populations 

with less diverse pollinators. In a different 

population of M. longiflorus, Eckert (1970) found 

that almost all pollination was done by solitary 

bees and that Anna's hummingbirds were only 

occasional visitors. Most pollination was effected 

by bees in the genus Osmia and carpenter bees in 

Ceratina. Both visited multiple flowers per plant, 

and self-pollination appeared common. Eckert also 

found that selfed flowers produced fewer capsules 

per plant and had more aborted ovules than 

outcrossed flowers. 

Horticulture.—The bush monkeyflowers and 

their horticultural hybrids have striking flower 

colors, produce masses of flowers, and are popular 

ornamental plants (Schmidt 1980, Perry 1992, 

Keator 1994). Plants can be grown from seeds or 

cuttings, but plants grown from seed tend to live 

longer (personal communication with Steve 

Morgan, University of California Riverside 

Botanic Garden, CA). The shrubs grow best in full 

sun to light shade. Flowering and leaf production 

may be extended by summer watering in more arid 

regions. However, Atkinson and others (1988) 

found that watering may increase susceptibility to 

atmospheric pollution in the dry season. In 

summer, unwatered plants drop most leaves, are 

not photosynthetically active, and take up little 

SO2, while watered plants retain their leaves, are 

photosynthetically active, and take up SO2. 

Growth and Management.—Bush 

monkeyflowers have been planted along highways 

corridors and are used in habitat restoration. There 

are about 145 million seeds/kg (Mirov and Kraebel 

1939). Seeds will germinate without pretreatment 

(Mirov and Kraebel 1939, Schmidt 1980), but 

seeds need to be exposed to light and leachate 

from charred wood may improve germination 

slightly (Keeley 1987). For pot culture, the tiny 

seeds should be sown on the surface of the soil and 

kept moist until seedlings emerge. Outdoors, they 

should be planted with shallow broadcasting 

methods. The phylogenetic relationships among 

populations and the fitness effects of translocations 

and hybridization are not yet known, and there are 

no published studies on population genetic 
498 

patterns. However, there is a complex pattern of 

geographic variation, genetic differentiation of 

floral form and resin chemistry, local adaptation to 

moisture environments, and extensive ability to 

hybridize. Consequently, attention should be made 

of flower color and habitat matching when 

obtaining seeds for large planting projects. Use of 

regionally local seed sources for planting projects 

can mitigate unknown fitness consequences of out 

of range plantings. A careful analysis of hybrid 

zones and the fitness of hybrids is needed to 

understand if hybridization following 

translocations will cause decreases in fitness of 

adjacent wild populations. 

Benefits.—Bush monkeyflowers are important 

plants for water-wise landscaping and revegetation 

projects. They provide nectar to hummingbirds 

and large bees and are an important larval host of 

the checkerspot butterfly. 

References 

Alpert, P., E.A. Newell, C. Chu, J. Glyphis, S.L. 

Gulmon, D.Y. Hollinger, N.D. Johnson, H.A. 

Mooney, and G. Puttick. 1985. Allocation to 

reproduction in the chaparral shrub, Diplacus

aurantiacus. Oecologia 66: 309-316. 

Atkinson, C.J., W.E. Winner, and H.A. Mooney. 

1988. Gas exchange and SO2 fumigation studies 

with irrigated and unirrigated field grown 

Diplacus aurantiacus and Heteromeles

arbutifolia. Oecologia 75: 386-393. 

Beardsley, P.M. and R.G. Olmstead. 2002. 

Redefining Phrymaceae: the placement of 

Mimulus, tribe Mimuleae, and Phryma. 

American Journal of Botany 89: 1093-1102. 

Beardsley, P.M., S.E. Schoenig, J.B. Whittall, and 

R.G. Olmstead. 2003. The radiation of Mimulus

(Phrymaceae) in western North America. 

American Journal of Botany. In Press. 

Bromer, W., J. Barnette, J. Lee, D. Green, and V. 

Ervin. 1990. Genetic variation within and among 

populations of Mimulus longiflorus in the Santa 

Monica Mountains, CA: consequences of 

breeding system and pollination. Bulletin of the 

Ecological Society of America 71: 103. 

Eckert, J.R. 1970. Pollination studies in Mimulus

longiflorus (Nutt.) Grant. Master’s thesis. 

California State University, Los Angeles. 88 p. 

Fetscher, A.E. and J.R. Kohn. 1999. Stigma 

behavior in Mimulus aurantiacus

(Scrophulariaceae). American Journal of 

Botany 86: 1,130-1,135. 

Grant, V. 1993. Origin of floral isolation between 

ornithophilous and sphingophilous plant species. 

Proceedings of the National Academy of 

Sciences of the United States of America 90: 

7729-7733. 

Han, K. and D.E. Lincoln. 1994. The evolution of 

carbon allocation to plant secondary 

metabolites: a genetic analysis of cost in 

Diplacus aurantiacus. Evolution 48: 1,550-

1,563. 

Hare, J.D. 2002. Geographic and genetic variation 

in the leaf surface resin components of Mimulus 

aurantiacus from southern California. 

Biochemical Systematics and Ecology 30: 281-

296. 

Hellmers, H., J.S. Horton, G. Juhren, and J. 

O'Keefe. 1955. Root systems of some chaparral 

plants in southern California. Ecology 36: 667-

678. 

Keator, G. 1994. Complete Garden Guide to the 

Native Shrubs of California. Chronicle Books, 

San Francisco, CA. 314 p. 

Keeley, J.E. 1987. Role of fire in seed germination 

of woody taxa in California chaparral. Ecology 

68: 434-443. 

Lincoln, D.E., T.S. Newton, P.R. Ehrlich, and K.S. 

Williams. 1982. Coevolution of the checkerspot 

butterfly Euphydryas chalcedona and its larval 

food plant Diplacus aurantiacus: larval response 

to protein and leaf resin. Oecologia 52: 216-223. 

Michener, D.C. 1983. Systematic and ecological 

wood anatomy of Californian Scrophulariaceae. 

I. Antirrhinum, Castilleja, Galvezia, and 

Mimulus sect. Diplacus. Aliso 10: 471-487. 

Mirov, N.T. and C.J. Kraebel. 1939. Collecting 

and handling seeds of wild plants. Civilian 

Conservation Corps, Forestry Publication 5. 

United States Government Printing Office, 

Washington, D.C. 

Mooney, H.A., and C. Chu. 1983. Stomatal 

responses to humidity of coastal and interior 
499

populations of a Californian shrub. Oecologia 

57: 148-150. 

Mooney, H.A., K.S. Williams, D.E. Lincoln, and 

P.R. Ehrlich. 1981. Temporal and spatial 

variability in the interaction between the 

checkerspot butterfly, Euphydryas chalcedona

and its principal food source, the Californian 

shrub, Diplacus aurantiacus. Oecologia 50: 195-

198. 

Munz, P.A. and D.D. Keck. 1968. A California 

Flora with Supplement. University of California 

Press, Berkeley, CA. 1,681 + 224 p. 

Perry, B. 1992. Landscape Plants for Western 

Regions: an Illustrated Guide to Plants for Water 

Conservation. Land Design Publishing, 

Claremont, CA. 318 p. 

Schmidt, M.G. 1980. Growing California Native 

Plants. University of California Press, Los 

Angeles. 

Thompson, D.M. 1993. Mimulus. In: J.C. 

Hickman, ed. The Jepson Manual: Higher Plants 

of California. University of California Press, 

Ltd., Los Angeles, CA. p. 1,037-1,046. 

Tulig, M. 2000. Morphological variation in 

Mimulus section Diplacus (Scrophulariaceae). 

Masters Thesis, Department of Biological 

Sciences, California State Polytechnic 

University, Pomona, CA. 82 p. 

Waayers. 1996. Hybridization, introgression, and 

selection in Mimulus aurantiacus ssp. australis

and M. puniceus. Master’s thesis. San Diego 

State University, San Diego, CA. 77 p. 

Wells, H. 1980. A distance coefficient as a 

hybridization index: an example using Mimulus

longiflorus and M. flemingii (Scrophulariaceae) 

from Santa Cruz Island, California. Taxon 29: 

53-65.
500 

Mitracarpus portoricensis (Urban) Urban Puerto Rico girdlepod 

RUBIACEAE 

Synonyms: Mitracarpus frigidus portoricensis Urban 

John K. Francis 

General Description.—Puerto Rico girdlepod (a 

name assigned by the Natural Resources 

Conservation Service) is a low, 20- to 60-cm tall 

evergreen shrub with basal diameters of 0.5 to 1.0 

cm. It may have one or more gray stems from the 

root crown and a compact, rounded crown with 

bifurcate or trifurcate branching. The stems, 

branches, and roots are woody but flexible. Plants 

are supported by a tap and lateral root system of 

yellow or tan roots. The glabrous branches and 

twigs are four-striate and four-angled. The leaves, 

which are crowded on recent twig growth, are 

sessile, nearly linear, 30 to 60 mm long and 1 to 3 

mm broad. The 2-cm broad, many-flowered heads 

are lateral or terminal on long peduncles. Tubular 

flowers are bright white, about 5 mm long. 

Capsules disperse their seeds when the tops come 

off releasing 1-mm long, ovate, brown seeds 

(Liogier 1997). 

Range.—Puerto Rico girdlepod is endemic to 

Puerto Rico, particularly to the northern, western, 

and southwestern portions of the Island (Liogier 

and Martorell 1982). The species is not known to 

have been planted or naturalized anywhere else. 

Ecology.—Puero Rico girdlepod is a low-statured, 

slow-growing plant that requires full or nearly full 

sunlight. It can compete with low herbs and 

grasses but cannot grow under a forest or shrub 

canopy. Consequently it must grow in areas with 

low competition, usually disturbed areas and areas 

of low fertility. It finds these conditions in eroded 

areas, dirt roads and road cuts, old fields, rocky 

hillsides, and coastal sands. Puerto Rico girdlepod 

may be found in areas of sedimentary (including 

limestone), igneous, and metamorphic (including 

ultramaphic) rocks. It tolerates salt spray, low soil 

fertility, compacted soils, and moderate physical 

abuse. Elevations range from a little above sea 

level to about 400 m in areas with rainfall from 

about 900 to about 1800 mm of mean annual 

precipitation. 

Reproduction.—Puerto Rico girdlepod blooms 

and produces seed throughout the year. A 

collection of seeds from the Susúa State Forest 

contained an average of 2,400 seeds/g. Placed on 

moist blotter paper, 29 percent of the seeds 

germinated between 4 and 90 days after sowing 

(author’s observation). Seeds are dispersed by 

wind and water. Opening of the forest canopy and 

probably bare soil are necessary for establishment. 

Seedlings are not common. 

Growth and Management.—Growth of Puerto 

Rico girdlepod is slow. Plants observed by the 

author appeared to have grown about 10 to 25 

cm/year. Life span of the shrubs appears to range 

from about 2 to 6 years. No planting or 

management experience has been reported. The 

species is not common enough or aggressive 

enough to warrant control. 

Benefits.—Puerto Rico girdlepod helps protect the 

soil, revegetates disturbed sites, adds to the 

aesthetics of wildland areas (because of its white 

flowers), and furnishes cover for wildlife. 
501

References 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Liogier, H.A. and L.F. Martorell. 1982. Flora of 

Puerto Rico and adjascent islands: a systematic 

synopsis. Editorial de la Universidad de Puerto 

Rico, Río Piedras, PR. 342 p. 
502 

Morella caroliniensis (P. Mill.) Small bayberry 

MYRICACEAE 

Synonyms: Myrica caroliniensis P. Mill. 

 Myrica curtissii A. Chev. 

 Myrica heterophylla Raf. 

 Myrica pensylvanica Mirb. 

Gerry Moore 

Illustration credit: Britton and Brown 1913 

General Description.—Bayberry is the name used 

most often to refer to this common eastern shrub. 

Other common names include candleberry, 

tallowshrub, and waxberry. Bayberry is a 

deciduous (northern part of range) to evergreen 

(southern portion of range) aromatic shrub or 

small tree to 4 m. The plant is single or 

multistemmed. The branchlets and twigs are terete 

and vary in color from brown-gray-blackish; they 

are dotted with yellow aromatic glands and vary 

from being glabrous to pubescent. The leaves are 

resinous, glandular, aromatic, with short pubescent 

petioles (to 7 mm) and elliptic to oblanceolate, 

membranaceous-leathery 3.0 to 7.0 cm by 1.5 to 

2.5 cm blades with margins (sometimes slightly 

revolute) entire or more usually toothed from the 

middle to the apex. The flowers are in unisexual 

catkins (aments), with the staminate (male) and 

pistillate (female) catkins appearing on different

plants. The staminate catkins range from 0.4 to 1.8 

cm; the pitsillate catkins range from 0.3 to 1.4 cm. 

The fruit of the bayberry is technically not a berry 

but a globose 3.0- to 6.0-mm drupe (one-seeded 

stone fruit) that at maturity is covered by a thick 

coat of gray-white wax. The fruits persist 

throughout the winter. The globose seed in each 

fruit measures slightly less in diameter than the 

fruit (author, personal observation). 

Range.—Bayberry occurs in eastern Canada (New 

Brunswick, Newfoundland, Nova Scotia, Ontario, 

Prince Edward Island, and Quebec) and the United 

States, from Maine south to Florida and west to 

Texas and Arkansas. Inland populations are known 

from western New York, Ohio and southern 

Ontario (Gleason and Cronquist 1991). 

Systematic Botany.—There are taxonomic and 

nomenclatural difficulties associated with the 

bayberry. Taxonomists disagree regarding 

bayberry’s generic placement, with some 

(Bornstein 1997, Fernald 1950, Gleason and 

Cronquist 1991) placing the bayberry and sweet 

gale (Myrica gale L.) in the same genus, Myrica, 

and others choosing to place the bayberry in its 

own genus, Morella (Wilbur 1994, 2002). Some 

taxonomists also choose to recognize two species 

of bayberry, a northern bayberry Morella 

pensylvanica and a southern bayberry, Morella 

caroliniensis (synonym: Myrica heterophylla 

Raf.). The two supposedly can be distinguished

based on whether the leaves are evergreen 

(southern) or deciduous (northern), the fruit wall is 

hairy (northern) or not (southern), and the twigs 

possess black hairs (southern) or not (northern). 

Further complicating matters, bayberry and the 

wax myrtle [Morella cerifera (L.) Small] can 

hybridize when growing together (Bornstein 

1997). Some authors also treat the name Morella 

caroliniensis (Miller) Small as a synonym of the 

wax myrtle (M. cerifera). This is due to different 

interpretations as to which plant–bayberry or wax 

myrtle–Miller (1768) was actually referring to in 
503

his original description. The recent work of Wilbur 

(1994, 2002) is followed here; therefore only one 

species is recognized under the name M. 

carolininesis. 

Ecology.—Bayberry can tolerate full sun to dense 

shade and can grow in dry to wet conditions. 

Habitats where it can be found include bogs, 

swamps, dunes, fields, heathlands, oak forests, 

pine forests, and the margins of streams and lakes. 

The roots of bayberry contain nitrogen fixing 

nodules, and this allows it to be particularly 

tolerant of nitrogen-poor, acidic soils (Morris and 

others 1974). Results are mixed as to whether 

bayberry’s nitrogen-fixing properties may assist 

the growth of other plant species growing nearby 

(Tiffney and Barrera 1979, Dudley and others 

1996). Leachate from the leaf litter of bayberry 

may have alleopathic properties and affect some 

other plants (Collins and Quinn 1982). 

Reproduction.—Bayberry is wind pollinated and 

blooms in the spring. The pollen is allergenic 

(Lewis 1977). The seeds of bayberry are dispersed 

by birds that eat the fruits (McClanahan and Wolfe 

1993, Place and Stiles, 1992, Ridley 1930). 

Growth and Management.—Bayberry is often 

cultivated on dry, sandy, sterile soils where few 

other species can grow (Bailey 1922, Rehder 

1940). It thrives best in acid peaty soils; it can not 

tolerate liming. Propagation can be by seed or 

layering (Chittenden and Synge 1956). 

Propagation can also be done with cuttings taken 

in late summer or fall. The cuttings should be 

placed in a bed of peat moss and sand (Everett 

1960). 

Benefits.—Settlers used the bark of bayberry in 

dentifrice mixtures (Lewis 1977) and the fruits in 

candle making (Bornstein 1997). Bayberry candles 

remain popular today; it can take up to 8 kg of 

bayberry fruit to yield 1 kg of wax. The fruits are 

also often used in decorative arrangements. Native 

Americans used the plant for various medicinal 

purposes (Moerman 1986). 

References 

Bailey, L.H. 1922. Standard cyclopedia of 

 horticulture. Vol. 4. MacMillan Co., London. 

 660 p. 

Bornstein, A.J. 1997. Myricaceae. In: Flora of 

North America Editorial Committee, eds. Flora 

of North America. Vol. 3. Oxford University 

Press, New York. p. 430-434. 

Britton, N.L. and A. Brown. 1913. Illustrated flora 

of the northern states, Canada, and the British 

possessions. Vol. 2, 2nd Ed. Scribner, New York. 

735 p. 

Chittenden, F.J. and P.M. Synge. 1956. Dictionary 

of Gardening. 2nd Ed. Clarendon Press, Oxford, 

UK. 623 p. 

Collins, B.S. and J.A. Quinn. 1982. Displacement 

of Andropogon scoparius on the New Jersey 

Piedmont by the successional shrub Myrica 

pensylvanica. American Journal of Botany 69: 

680-689. 

Dudley, J.L., B. Michener, and K. Lajtha. 1996. 

The contributions of nitrogen-fixing symbioses 

to coastal heathland succession. American 

Midland Naturalist 135: 334-342. 

Everett, T.H. 1960. New illustrated encyclopedia 

of gardening. Vol. 3. Greystone Press, New 

York. 479 p. 

Fernald, M.L. 1950. Gray’s manual of botany. 

American Book Co., New York. 1,632 p. 

Gleason, H.A. and A. Cronquist. 1991. Manual of 

vascular plants of the northeastern United States 

and adjacent Canada. 2nd Ed. New York 

Botanical Garden, Bronx. 910 p. 

Lewis, W.H. 1977. Medical botany. John Wiley & 

Sons, New York. 515 p. 

McClanahan, T.R. and R.W. Wolfe. 1993. 

Accelerating forest succession in a fragmented 

landscape: the role of birds and perches. 

Conservation Biology 7: 279-288. 

Miller, P. 1768. The gardeners dictionary. 8th Ed. 

Printed for the author, London. 1,329 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. 2 Vols. Technical Report 19. 

University of Michigan Museum of 

Anthropology, Ann Arbor, MI. 910 p. 

Morris, M., D.E. Eveleigh, S.C. Riggs, and W.N. 

Tiffney, Jr. 1974. Nitrogen fixing in the 

bayberry (Myrica pensylvanica) and its role in 
504 

coastal succession. American Journal of Botany 

61: 867-870. 

Place, A.R. and E.W. Stiles. 1992. Living off the 

wax of the land: bayberries and yellow-rumped 

warblers. Auk 190: 334-345. 

Rehder, R. 1940. Manual of cultivated trees and 

shrubs, Macmillan Publishing Co, New York. 

996 p. 

Ridley, H.N. 1930. The dispersal of plants 

throughout the world. L. Reeve & Co., Ashford, 

Kent, UK. 744 p. 

Tiffney, W.N., Jr. and J.F. Barrera. 1979. 

Comparative growth of pitch and Japanese black 

pine in clumps of the N2 –fixing shrub, bayberry. 

Botanical Gazette 140 (Suppl.): S108-S109. 

Wilbur, R.L. 1994. The Myricaceae of the United 

States and Canada: genera, subgenera, and 

series. Sida 16: 93-107. 

Wilbur, R.L. 2002. The identity and history of 

Myrica caroliniensis (Myricaceae). Rhodora 

104: 31-41. 
505

Morinda citrifolia L. noni 

RUBIACEAE 

Synonyms: none 

John K. Francis 

General Description.—Noni, is also known as 

Indian mulberry, great morinda, cheezefruit, 

morinda, mouse’s pineapple, yellow root, jumbie 

breadfruit, hog apple, pain killer, mengkudu, 

nono, feyukke friudem rhubarbe caraïbe, bilimbi, 

pomme-macaque, and pomme de singe. It is a 

large evergreen shrub or small tree to 6 m or more 

in height and 13 cm or more in stem diameter. 

Sapwood is yellow-brown and soft. The bark is 

gray or brown, smoothish to slightly rough. Twigs 

are light green and four-angled. The opposite 

leaves are attached by stout petioles 1 to 2 cm 

long. The blades are dark green and shiny, ovate 

or elliptic, 14 to 30 cm long by 8 to 18 cm broad, 

and have prominent veins. The white tubular 

flowers are grouped in globose heads at the leaf 

axils. The five-lobbed flower tubes are about 6 

mm long. The greenish-white to pale-yellow, 

fleshy fruits are ovoid or globose syncarps 5 to 7 

cm long. They have an unpleasant odor 

resembling cheese. They contain a number of 

seeds about 4 mm long (Howard 1989, Liogier 

1997, Little and Wadsworth 1964, Nelson 1996). 

Range.—Noni is probably native to maritime 

forests of Northern Australia, the Western Pacific, 

and Indian Oceans (Stevens and others 2001). It 

was widely spread by native peoples beginning 

2000 years ago (Smith 2002) and more recently by 

Europeans so that it is naturalized in most of the 

tropical and tropical coastal forests of the World 

(Howard 1989) including Florida, Puerto Rico, 

and the U.S. Virgin Islands (Little and Wadsworth 

1964).

Ecology.—Noni is most competitive on sands and 

loamy sands but will grow on soils of all other 

textures if planted and protected from competition. 

It also grows well on very rocky soils. Although it 

grows naturally only a few meters above sea level, 

noni can be cultivated on sites up to 800 m or 

more above sea level (Stevens and others 2001). 

The species tolerates salty soils and salt spray. It is 

intermediate in shade tolerance, growing under the 

canopy of forests as well as in the open. Noni 

grows naturally on the edges of mangroves, in 

coastal forests, and on the landward side of beach 

strand vegetation. The species is grown in 

plantations, as an ornamental, and as a garden 

medicinal tree. A symbiotic association in the 

native range with weaver ants (Oecophylla 

smaragdina) provides the ants with food and 

leaves for nesting in exchange for protection from 

insect predators (Tan 2001). 

Reproduction.—Noni flowers and fruits nearly 

throughout the year (Little and Wadsworth 1964). 

Seeds (air dried) in Puerto Rico weighed an 

average of 0.0259 g/seed or 38,600 seeds/kg. 

Without any pretreatment, 24 percent germinated 

in commercial potting mix beginning 70 days after 

sowing (Francis and Rodríguez 1993). In its native 

habitat, the seeds of noni are probably dispersed 

by fruit bats and other mammals. The plant may 

be propagated from both seeds and cuttings 

(Association of Societies for Growing Australian 

Plants 2000). 

Growth and Management.—Published 
506 

information is lacking on growth and 

management. 

Benefits.—Although the fruits of noni are 

somewhat tasteless and have an unpleasant smell, 

they are eaten as famine food in Indonesia, 

Australia, and the Pacific Islands. The young 

leaves are also eaten as a vegetable and contain 4 

to 6 percent protein. Noni is planted as a 

windbreak, for support of pepper vines, and shade 

for coffee bushes (Tan 2001). The species is 

useful for coastal erosion control (Association of 

Societies for Growing Australian Plants 2000), 

and it is planted as an ornamental in some areas. 

The bark has been the source of a red dye (Little 

and Wadsworth 1964). Roots are used to produce 

an orange dye (Nelson 1996). Harvested from both 

plantations and from the wild, noni is one of the 

most important botanical remedies and food 

supplements traded on the international market. It 

is usually the fruit juice that is sold, fresh or dried. 

Leaves, bark, and roots are also used for a great 

many maladies. Pain, arthritis, diabetes, high 

blood pressure, skin and stomach ulcers, 

depression, senility, diarrhea, arteriosclerosis, 

cancer, AIDS, skin parasites, and bad breath are 

all treated (Association of Societies for Growing 

Australian Plants 2000, Cambie and Ash 1994, 

Tan 2001). A number of physiologically active 

chemicals, anthraquinones, alkaloids, 

asperuloside, caproic, caprylic, and ursolic acids, 

β-sitosterol, and asperuloside may account for 

some of the effects (Cambie and Ash 1994). 

Antitumor activity expressed in enhanced survival 

of tumor-bearing mice has been demonstrated 

after treating with juice extracts (Hirazumi and 

Furusawa 1999). Aqueous extracts of roots were 

shown to have an analgesic effect on mice without 

any sign of toxicity, and a sedative effect at high 

doses (Younos and others 1990). Octanoic acid, 

which is present in the ripe fruits, effectively 

poisons a fruit fly (Drosophila melanogaster), 

honey bee (Apis mellifera), an ant (Lasius sp.), 

and cockroaches (Periplaneta americana and 

Blattella germanica) (Legal and Plawecki 1995). 

Noni is the principal larval host of the hawk moth, 

Macroglossum hirundo vitiensis in Fiji 

(Stampsfiji.com 2002). 

References 

Association of Societies for Growing Australian 

Plants. 2000. Morinda citrifolia. 

http:/farrer.csu.edu/ASGAP/m-cit.html. 2 p. 

Cambie, R.C. and J. Ash. 1994. Fijian medicinal 

plants. CSIRO, Canberra, Australia. 365 p. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

 

Hirazumi, A. and E. Furusawa. 1999. An 

immunomodulatory polysaccharide-rich 

substance from the fruit juice of Morinda 

citrifolia (noni) with antitumour activity. 

Phytotherapy Research 13(5): 380-387. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Part 3. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Legal, L. and M. Plawecki. 1995. Comparative 

sensitivity of various insects to toxic compounds 

from Morinda citrifolia L. Entomological 

Problems 26(2): 155-159. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Sarasota, FL. 391 p. 

Smith, R.S. 2002. History and tradition of 

Morinda citrifolia. http://rsscomp.freeyellow. 

com/morindacitrifoliastory.htm. 2 p. 

Stampsfiji.com. 2002. The Sphingid (hawk) moths 

of Fiji stamp issue. http://www.stampsfiji. 

com/stamps/moths/ 4 p. 

Stevens, W.D., C. Ulloa U., A. Pool, and O. M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanic Garden, St. Louis, MO. 

p. 1,911-2,664. 
507

Tan, R. 2001. Mangrove and wetland wildlife at 

Sungei Buloh Nature Park: Great morinda. 

http://www.naturia.per.sg/buloh/plants/morinda.

htm. 3 p. 

Younos, C., A. Rolland, J. Fleurentin, M.C. 

Lanhers, R. Misslin, and F. Mortier. 1990. 

Analgesic and behavioral effects of Morinda 

citrifolia. Planta Medica 56(5): 430-434. 
508 

Murraya exotica L. orange jasmine 

RUTACEAE 

Synonyms: Murraya paniculata (L.) Jack 

 Chalcas paniculata L. 

 Chalcas exotica (L.) Millsp. 

John K. Francis 

General Description.—Orange jasmine is also 

known as mock orange, satin wood, honey bush, 

China-box, café de la India, mirto, azahar, naranjo 

jazmín, limonaria, and bun (Little and others 

1974). It is an evergreen shrub or occasionally a 

small tree, usually 2 to 3 m in height but reaching 

7.5 m and 13 cm in stem diameter. Older orange 

jasmine normally have multiple stems from the 

ground level. The stems are supported by taproots 

with lateral roots and abundant fine roots. Stem 

bark is gray, becoming fissured and rough. Orange 

jasmine branches and twigs are slender and 

abundant at all heights. The alternate leaves are 

pinnately compound with three to nine leaflets 

alternating on the rachis. The 1- to 5-cm, leaflets 

are dark-green, stiff, ovate, and smell of citrus 

when crushed. The shrub produces fragrant, fivepetaled, white flowers borne in small clusters near 

the branch ends. Later, shiny, red elliptic fruits 

about 1 cm long develop. One or two light green 

seeds are embedded in the bitter, watery pulp 

(Liogier 1988, Little and others 1974). The seeds 

are tear-drop shaped, rounded or flattened on one 

side depending on whether there are one or two 

seeds per fruit. 

Range.—The native range of orange jasmine 

includes China, India, Sri Lanka, the Andaman 

Islands, Myanmar, Thailand, Kampuchea, Viet 

Nam, Malaysia, northeastern Australia, New 

Caledonia, and Taiwan (Parrotta 2001). In 

addition, the shrub has been planted throughout 

the Tropics and has naturalized in many locations 

including Puerto Rico (author’s observation). 

Some authors consider M. exotica and M. 

paniculata to be separate species (Howard 1988, 

Pacific Island Ecosystems at Risk 2002). 

Ecology.—Orange jasmine is adapted to a wide 

range of conditions. Naturalized plants in Puerto 

Rico grow in areas receiving from about 750 mm 

to 1900 mm of annual precipitation. The species 

grows from nearly sea level to elevations of 1,300 

m (Neal 1965). It grows on most well-drained 

soils derived from both sedimentary and igneous 

rocks, although it is said to favor limestone areas 

(Pacific Island Ecosystems at Risk 2002). Plants 

survive temperatures to about –4 °C (DesertTropicals 2002). Ornamental plants are attacked at 

times by white flies, scale insects, nematodes, and 

sooty mold (Sheehan 1975). It was noted that 

powdery mildew attacked the species in pruned 

hedges but not in free-growing unpruned plants 

(Pathak and others 1992). Orange jasmine is 

moderately intolerant of shade. Although growing 

well under partial shade, it produces few flowers 

or fruits. Escaped plants in Puerto Rico are most 

often seen at the edges and in the understory of 

dry and moist secondary forests. Seedlings do not 

compete well with grass or herbaceous vegetation. 

Reproduction.—Orange jasmine flowers 

irregularly throughout the year (Little and others 

1974). In Hawaii, it flowers from June to 

September (Neal 1965). Fruits collected from 

plants in Puerto Rico averaged 0.370 + 0.012 

g/fruit. Air-dried seeds from these fruits averaged 

0.0568 + 0.0012 g/seed or 17,600 seeds/kg. Sown 
509

on commercial potting mix, 65 percent germinated 

between 25 and 60 days of sowing. Germination is 

hypogenous. Seedlings quickly develop deep root 

systems and grow at a moderate rate. Plants 

coppice vigorously after disturbance. Nursery 

production is by seed, cuttings (Little and others 

1974), and air-layers (Woman’s Club of Havana 

1952). 

Growth and Management.—Orange jasmine 

plants can live at least 15 years (author’s 

observation). Once established as an ornamental, 

they need little care. The species has not yet been 

reported to be a weed in any area. Orange jasmine 

can be killed (with moderate success) with 

herbicides recommended for broad-leaf weeds, by 

girdling (Negreros-Castillo and Hall 1994), or by 

grubbing out the plants. The species is the 

preferred host of the citrus psyllid, Diaphorina 

citri, the vector for “citrus greening” disease 

(Pacific Island Ecosystems at Risk 2002), and is a 

host of Mediterranean fruit flies (Diptera: 

Tephritidae) (Harris and Lee 1986). 

Benefits.—Orange jasmine is a popular hedge 

plant in the tropics. It can be used as a background 

plant or an accent plant. It can be pruned into 

animal and other shapes (topiary), pruned into tree 

form, grown as a potted plant (Whistler 2000), or 

cultivated as a bonsai (Gonzalez 2002). The 

sapwood is light yellow, and the heartwood is 

light brown and heavy, hard, fine-textured and 

good for small turned articles (Little and others 

1974). Branches or stems 2 to 5 cm in diameter in 

the Indian Himalayan region were evaluated for 

fuelwood suitability. Results were as follows: 

oven-dry density 0.72, dry-weight caloric value 

20.2 kj/g, ash 0.6 percent, moisture 54.7 percent, 

and nitrogen 0.42 percent (Negi and Todaria 

1993). In the wild, orange jasmine contributes to 

the biodiversity, protects the soil, and furnishes 

food and cover for wildlife. The leaves and other 

tissues have both stimulant and astringent 

properties and are used to treat diarrhea, 

dysentery, cuts, joint pain, body aches (Parrotta 

2001), venereal disease (Kinoshita and Firman 

1996), and as an abortive (Xiao and Wang 1991). 

In addition to essential oils, tissues of orange 

jasmine contain the indole alkaloid yuehchukene 

(Xiao and Wang 1991) and at least eight highly 

oxygenated flavones (Kinoshita and Firman 1996). 

References 

Desert-Tropicals. 2002. Orange jasamine. 

http://www.desert-tropicals.com/Plants/ 

Rutaceae/Murraya_paniculata.html. 2 p. 

Gonzalez, G. 2002. Bonsai gallery photos of 

Murraya paniculata bonsi. Tropical 

Bonsai.com. http://www.Tropicalbonzai.com/ 

MurrayaPaniculata.htm. 1 p. 

Harris, E.J. and C.Y.L. Lee. 1986. Seasonal and 

annual occurrence of Mediterranean fruit flies 

(Diptera: Tephritidae) in Makaha and Waianae 

valleys, Oahu, Hawaii. Environmental 

Entomology 15(3): 507-512. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University. Jamaica Plain, 

MA. 673 p. 

Kinoshita, T. and K. Firman. 1996. Highly 

oxygenated flavonoids from Murraya 

paniculata. Phytochemistry 42(4): 1207-1210. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Museum 

Press, Honolulu, HI. 924 p. 

Negi, A.K. and N.P. Todaria. 1993. Fuelwood 

evaluation of some Himalayan trees and shrubs. 

Energy 18(8): 799-801. 

Negreros-Castillo, P. and R.B. Hall. 1994. Four 

methods for partial overstory removal in tropical 

forests of Mexico. Journal of Environmental 

Management 41(3): 237-243. 

Pacific Island Ecosystems at Risk. 2002. Invasive 

plant species: Murraya paniculata (L.) Jack, 

Rutaceae. http://www.hear.org/pier3/mupan. 

htm. 2 p. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CABI Publishing, Wallingford, UK and 
510 

New York. 917 p. 

Pathak, R.K., Madhulika-Mahajan, and S.N. 

Sachan. 1992. Powdery mildew and cultural 

practices. Indian Journal of Forestry 15(1): 73. 

Sheehan, M.R. 1975. Florida landscape plants, 

native and exotic. The University Presses of 

Florida, Gainesville, FL. 420 p. 

Whistler, W.A. 2000. Tropical ornamentals, a 

guide. Timber Press, Portland, OR. 542 p. 

Woman’s Club of Havana. 1952. Flowering plants 

from Cuban gardens. Criterion Books, New 

York. 365 p. 

Xiao, P.G. and N.G. Wang. 1991. Can 

ethnopharmacology contribute to the 

development of anti-fertility drugs? Journal of 

Ethnopharmacology 32(1-3): 167-177. 
511

Myrica gale L. sweet gale 

MYRICACEAE 

Synonyms: Gale palustris A. Chev. 

 Myrica gale var. subglabra (A. Chev.) Fernald 

 Myrica gale var. subarctica J. Rousseau. 

 Myrica gale ssp. tomentosa (C. DC) E. Murray 

 Myrica gale var. tomentosa C. DC 

Gerry Moore 

Illustration source: Britton and Brown 1913 

General Description.—Sweet gale is the name 

used most often to refer to this shrub species. It is 

also known as bog myrtle, Dutch myrtle English 

myrtle, and meadow fern, although this species is 

neither a true fern nor myrtle. Sweet gale is an 

aromatic, deciduous, shrub up to 2.5 m tall. The 

plant is single or multistemmed, with strongly 

ascending branches. The twigs are terete, brown 

with scattered yellow glands. The leaves are 2 to 6 

cm long (including the short petiole), oblanceolate 

(i.e., widest toward apex) with cuneate bases, 

toothed only toward the apex, glabrous and shining 

above but usually pubescent on the blade’s 

underside. Like the twigs, the blades on both sides 

are often beset with yellow glands. The flowers are 

in unisexual, unbranched catkins (aments), with 

the staminate (male) and pistillate (female) catkins 

appearing on different plants. Individual plants 

have been known to change sex from year to year 

(Burges 1993). The catkins are stalkless and borne 

on the upper portions of the preceding year’s 

branchlets. These branchlets die after flowering. 

The staminate catkins range from 6 to 10 mm long, 

are crowded on the branchlet, with the individual 

flowers overtopped by the broad, shining, 

subtending, brown bracts. The pistillate catkins 

range from 7 to 10 mm long, each flower 

subtended by two wing-like bracts that remain 

fused to the fruit. The fruit of the sweet gale is a 

three-pointed, compressed, ovoid nutlet that at 

maturity is dotted with shining red to yellow resin 

glands. 

Range.—Sweet gale is found in the New and Old 

World temperate regions. It occurs in the 

Northeastern (Connecticut, Maine, Massachusetts, 

New Hampshire, New Jersey, Michigan, 

Minnesota, New York, North Carolina, 

Pennsylvania, Rhode Island, Vermont, and 

Wisconsin) and the Northwestern (Oregon and 

Washington) United States (USDA 2003). The 

species is also known throughout much of Canada 

(Alberta, British Columbia, Manitoba, New 

Brunswick, Newfoundland, Northwest Territory, 

Nova Scotia, Ontario, Prince Edward Island, 

Quebec, Saskatchewan, and Yukon) and Alaska 

(Bornstein 1997). In Eurasia it is known from 

northwest Europe to Portugal, Poland and 

northwestern Russia (Belgium, Britain, Denmark, 

Finland, France, Germany, Ireland, Netherlands, 

Portugal, Norway, Poland, Russia, Spain, and 

Sweden). 

Systematic Botany.—Taxonomists disagree as to 

whether the sweet gale and the waxy-fruited 

bayberry [Morella caroliniensis (Mill.) Small] and 

wax myrtle [Morella cerifera (L.) Small] should 

be placed in the same genus (Bornstein 1997, 

Fernald 1950, Gleason and Cronquist 1991) or in 

separate genera (Wilbur 1994, 2002). Sweet gale 
512 

can be readily distinguished from the waxy fruited 

species (bayberry, wax myrtle) on the bases of 

flowers and fruits. If sweet gale is to be recognized 

as generically distinct from bayberry and wax 

myrtle, as is done here, the generic name Myrica is 

to be retained for sweet gale, with the bayberry 

and wax myrtle being placed in the genus Morella 

(Wilbur 1994, 2002). Various varieties and 

subspecies [e.g., M. gale var. subglabra Fernald, 

M. gale var. tomentosa C. DC., M. gale ssp. 

tomentosa (C. DC) E. Murray] have been 

recognized within sweet gale based on the degree 

of pubescence present on the leaves. 

Ecology.—Sweet gale can tolerate full sun to 

dense shade and is usually found in moist to wet 

areas. Habitats where it can be found include bogs, 

fens, swamps, and the margins of streams and 

lakes. The roots of sweet gale contain nitrogenfixing nodules, which allow it to be particularly 

tolerant of nitrogen-poor, acidic soils (Baker and 

Parsons 1997). Nitrogen fixation is accomplished 

by the actinomycetous fungal genus, Frankia. 

Reproduction.—Sweet gale is wind pollinated 

and blooms in the spring. The pollen is allergenic 

(Lewis 1977). The fruits of sweet gale mature in 

the fall and are dispersed by water (Ridley 1930, 

Wilbur 1994). 

Growth and Management.—Sweet gale is often 

cultivated on moist acidic soils where few other 

species can be grown (Bailey 1922, Rehder 1940). 

It thrives best in peaty soils and cannot tolerate 

liming. Propagation can be by seed, layering, or 

root suckers (Chittenden and Synge 1956). 

Propagation can also be effected through cuttings 

taken in late summer or fall. The cuttings should 

be placed in a bed of peat moss and sand (Everett 

1960). 

Benefits.—The branches and bark have been used 

in making a gale beer and in tanning; a decoction 

has been used for an insecticide and to kill vermin 

(Rook 1998). The branches of sweet gale were 

used by the Native Americans, Bella Coola, to 

prepare decoctions to be used as a diuretic or as a 

treatment for gonorrhea (Moerman 1986). 

References 

Bailey, L.H. 1922. Standard cyclopedia of 

horticulture Vol. 4. MacMillan Co., London. 

660 p. 

Baker, A. and R. Parsons. 1997. Rapid 

assimilation of recently fixed N2 in root nodules 

of Myrica gale. Physiologia Plantarum 99: 640-

647. 

Britton, N.L. and A. Brown. 1913. Illustrated flora 

of the northern states, Canada, and the British 

possessions. Vol. 2, 2nd Ed. Scribner, New York. 

735 p. 

Bornstein, A.J. 1997. Myricaceae. In: Flora of 

North America Editorial Committee, eds. Flora 

of North America Vol. 3. Oxford University 

Press, New York. p. 430-434. 

Burges, N.A. 1993. Myrica. In: T.G. Tutin, N.A. 

Burges, A.O. Chater, J.R. Edmondson, V.H. 

Heywood, D.M. Moore, D.H. Valentine, S.M. 

Walters, and D.A. Webb, eds. Flora Europaea. 

2nd Ed.. Vol. 1. Cambridge University Press, 

Cambridge, UK. p. 66-67. 

Chittenden, F.J. and P.M. Synge. 1956. Dictionary 

of Gardening 2nd Ed. Vol. 1. Clarendon Press, 

Oxford, UK. 623 p. 

Everett, T.H. 1960. New illustrated encyclopedia 

of gardening. Vol. 3. Greystone Press, New 

York. 479 p. 

Fernald, M.L. 1950. Gray’s manual of botany. 

American Book Co., New York. 1,632 p. 

Gleason, H.A. and A. Cronquist. 1991. Manual of 

vascular plants of the northeastern United States 

and adjacent Canada 2nd Ed. New York 

Botanical Garden, Bronx, NY. 910 p. 

Lewis, W.H. 1977. Medical botany. John Wiley & 

Sons, New York. 515 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. 2 Vols. Technical Report 19. 

University of Michigan Museum of 

Anthropology, Ann Arbor, MI. 910 p. 

Rehder, A. 1940. Manual of cultivated trees and 

shrubs, Macmillan Publishing Co, New York. 

996 p. 

Ridley, H.N. 1930. The dispersal of plants 

throughout the world. L. Reeve & Co., Ashford, 

Kent, UK. 744 p. 
513

Rook, J.S. 1998. A boundary waters compendium. 

http://www.rook.org/earl/bwca/nature/shrubs/ 

myricagale.html [notpaged]. 

USDA, NRCS. 2003. The PLANTS Database, 

Version 3.5 Baton Rouge, LA. 

http:plants.usda.gov. [not paged]. 

Wilbur, R.L. 1994. The Myricaceae of the United 

States and Canada: genera, subgenera, and 

series. Sida 16: 93-107. 

Wilbur, R.L. 2002. The identity and history of 

Myrica caroliniensis (Myricaceae). Rhodora 

104: 31-41. 
514 

Neea buxifolia (Hook. f.) Heimerl nia 

NYCTAGINACEAE 

Synonyms: Eggersia buxifolia Hook. f. 

John K. Francis 

General Description.—Nia, also known as 

saltwood, is a fine-leafed, upright, evergreen shrub 

from 2 to 7 m in height and 2 to 8 cm in basal stem 

diameter. There is usually a single stem emerging 

from the ground with multiple branches low on the 

stem. Nia plants have a weak taproot with lateral 

and fine roots, all brownish orange in color. The 

few slender branches form a diffuse crown. Bark is 

smooth and gray. The inner bark is bitter. The 

wood is whitish, soft, and does not have 

discernable annual rings. Oblong to oblanceolate, 

entire leaves are opposite or in whorl-like groups 

along the twigs. The 9- to 24-mm long and 3- to 9-

mm broad blades are supported by 1- or 2-mm 

petioles. Small light yellow flowers are borne on 

separate plants (dioecious). The elliptic, red, 

slightly fleshy fruits are 4 to 5 mm long and 

contain one seed each (Liogier 1985, Little and 

others 1974). 

Range.—Nia is native to Puerto Rico, its offshore 

island of Culebra, St. Thomas and St. John of the 

U.S. Virgin Islands, and Virgin Gorda of the 

British Virgin Islands (Clubbe 2000, Liogier 1985, 

Little and others 1974). It has been widely used as 

a bonsai plant and planted somewhat as an 

ornamental but is not known to have naturalized 

anywhere. 

Ecology.—Nia has an intermediate tolerance for 

shade. It usually grows in the understory of low to 

medium-density remnant and late secondary 

forests. The species sometimes grows in semiexposed positions with other shrubs and low trees 

on the tops of ridges. These are moist and dry 

forests over sedimentary (especially limestone), 

igneous, and metamorphic rocks. It is rare to 

infrequent in most habitats except the moist 

limestone hills in northern Puerto Rico where it is 

locally common. Nia is sometimes found in coastal 

thickets. A wide variety of well-drained soils are 

colonized. Mean annual precipitation ranges from 

750 to about 2200 mm at elevations from a little 

above sea level to 500 m. The minimum 

temperature for potted plants is reported to be 4.5 

°C (Zane 2002). 

Reproduction.—Nia flowers in the spring and 

early summer and matures fruits in mid- to late 

summer. The fruits, which are produced in sparing 

quantities, ripen a few at a time over a period of a 

few weeks. A group of fruits collected in Puerto 

Rico averaged 0.0291 ± 0.0011 g/fruit. However, 

the seeds from this collection that averaged 0.0021 

± 0.0001 g/seed (air dried) failed to germinate. 

Seeds of this species are probably dispersed by 

birds. Natural seedlings are rare to scattered. 

According to Zane (2002), nia can be propagated 

by seed, branch cuttings taken in March through 

June, and by air layering. 

Growth and Management.—Nia is relatively 

slow growing. Open-grown ornamentals reached 2 

m in height in about 8 years. Understory saplings 

add just a few centimeters of height per year. Life 

span in forests may reach several decades. 

Although methods of propagation and culture in 

pots as bonsais is well understood, no wildland 

plantings have been documented. Probably the 

best management is protection of the stands where 

the species occurs. 

Benefits.—Nia contributes to the diversity and 

aesthetics of the forests where it occurs, helps 

protect the soil, and furnishes food and cover for 

wildlife. It is one of the best broadleaf tropical 
515

(room temperature) bonsai species available and 

offers promise as an ornamental, especially for 

hedges and other shaped ornamentals. 

References

Clubbe, C. 2000. British Virgin Islands: 

conservation and training. Kew Scientist 17:5. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Zane, T.L. 2002. Neea buxifolia—Neea buxifolia. 

http://www.bonsai-bci.com/species/neea.html. 

2 p. 
516 

Odontonema cuspidatum (Nees) Kuntze cardinal’s guard 

ACANTHACEAE 

Synonyms: Odontonema tubiforme (Bertol.) Kuntze 

Odontonema strictum (Nees) Kuntze 

 Thrysacenthus cuspidatus Nees in DC. 

 Thyrsacanthus strictus Nees in DC. 

John K. Francis 

General Description.—Cardinal’s guard, also 

known as firespike, is an evergreen shrub 1 to 2 m 

in height and 1 to 2.5 cm in basal stem diameter. It 

grows in patches and clumps that generate from 

root suckers. The smooth, green stems, which are 

woody below and semiwoody above, are 

supported by stiff lateral roots. The stems develop 

few branches. The opposite leaves are dark green, 

glabrous, elliptic, 10 to 30 cm long, and acuminate 

at the tip. The inflorescences are terminal racemes, 

sometimes branched, composed of tubular flowers 

of scarlet or other colors including pink, white, 

and lavender. Fruits are clavate (club-shaped) 

capsules 1 to 2 cm long that contain flattened 

semiorbicular seeds (Liogier 1997). 

Range.—Cardinal’s guard is native from Mexico 

to Panama (Stevens and others 2001). The species 

has naturalized and escaped in at least Puerto 

Rico, Hispaniola, and Cuba (Liogier 1997), and is 

widely grown as an ornamental in Florida 

(Watkins 1975). Some authors propose separating 

O. cuspidatum and O. tubiforme into individual 

species (Whistler 2000) or using only the name O. 

tubiforme (Stevens and others 2001). 

Ecology.—Cardinal’s guard prospers on fertile 

and moderately fertile soils that are continually 

moist. It is moderately tolerant of shade and will 

bloom in full sun, broken sun, and moderate shade 

(author’s observation, Blomber 2000). In Central 

America, it grows from near sea level to 1,400 m 

in elevation (Stevens and others 2001). Cardinal’s 

guard is not salt tolerant (Watkins 1975) and dies 

to the ground if subjected to frost. In areas subject 

to frost, it is grown as an annual or herbaceous 

perennial (Blomber 2000, Watkins 1975). The 

plants are mostly free of pests and disease but may 

occasionally be attacked by mealy bugs 

(Pseudococcidae) (Woman’s Club of Havana 

1952). 

Reproduction.—In tropical countries, cardinal’s 

guard blooms throughout the year (Whistler 

2002), but in warm temperate areas, it blooms in 

the fall (Watkins 1975). Although the species is 

common in Puerto Rico, the plants produce few 

viable seeds. Capsules fall off before drying and 

liberating seeds. Fresh capsules collected in Puerto 

Rico weighed an average of 0.0095 + 0.0002 

g/capsule. One hundred of these capsules were 

sown on moist potting mix and only a single 

seedling emerged. Cardinal’s guard is most 

common in disturbed areas (Stevens and others 

2001), along streams and drains, and in extramoist places in secondary forests. Most of the 

stands of cardinal’s guard in Puerto Rico have 

originated from abandoned gardens or errant 

pieces of stem or root that have been transported 

by streams or dumped in the woods with garden 

prunings. Once established, plants spread by root 

suckers. The stems also layer (root) readily when 

they become prostrate. 

Growth and Management.—Cardinal’s guard 

grows rapidly from suckers and must be pruned 

frequently when used as hedges (Woman’s Club 

of Havana 1952). Normally, little care is needed 

once the plant is established (Whistler 2000). 

Benefits.—In natural stands, cardinal’s guard adds 

to biodiversity and protects against soil erosion. 
517

The species is widely cultivated as an ornamental 

in tropical and subtropical areas for its striking red 

flowers. It is used as an accent and background 

plant and sometimes employed to form hedges 

(Watkins 1975, Whistler 2000, Woman’s Club of 

Havana 1952). It is also popular in gardens 

because it attracts butterflies and hummingbirds 

that feed on the nectar (Blomber 2000, Watkins 

1975). 

References 

Blomber, M. 2000. In the garden: tropical plants 

can add diversity to gardens. 

http://gainesvillesun.com/news/marina/02-26-

00marina.shtml. 4 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in systematic botany Vol. 85, No. 1. 

Missouri Botanical Garden Press. 943 p. 

Watkins, J.V. 1975. Florida landscape plants, 

native and exotic. The University Presses of 

Florida, Gainesville, FL. 420 p. 

Whistler, W.A. 2000. Tropical ornamentals, a 

guide. Timber Press, Inc., Portland, OR. 542 p. 

Woman’s Club of Havana. 1952. Flowering plants 

from Cuban gardens. Criterion Books, New 

York. 365 p. 
518 

Olyra latifolia L. carricillo 

POACEAE 

Synonym: Olyra paniculata Sw. 

 Olyra cordifolia Kunth. 

John K. Francis 

General Description.—Carricillo (or carrucillo), 

also known as lintentwa, cortadora, lambedora, 

and sonadora, is a slender, arching bamboo-like 

grass. Unsupported culms reach about 1.5 m 

before sagging to horizontal. Climbing into the 

crowns of shrubs and low trees, plants may reach 3 

to 7 m in height and sometimes as much laterally. 

The culms (stems) are thin-walled, brittle but 

tough, and can reach 1.5 cm in diameter. Each 

plant usually has several culms arising from a tight 

root crown composed of brief rhizomes that 

support abundant fine roots. The culms are green, 

sometimes mottled with purple, throughout their 

length and are enveloped with up to 8-cm sheaths 

arising from the nodes. The culms are lineal except 

for vertical sprouts on reclining stems and 

“brooms” or diffuse multiple branches that form 

near the ends of older culms. The leaves are 

ribbon-like, 15 to 25 cm long and 3 to 6 cm broad 

with long pointed tips and one edge of the leaf 

base rounded and the other straight, forming a 50 ° 

angle with respect to the leaf axis for about 2 cm. 

Inflorescences are panicles 10 to 15 cm long, 

terminal or upper axillary, and contain few to 

many flowers. The 5-mm fruits are smooth, shiny, 

ivory-colored, and hard (author’s observation, 

Croat 1978, Howard 1979, Stevens and others 

2001). 

Range.—Carricillo is native to southern Mexico, 

Central America, South America to northern 

Argentina, the Antilles, moist Tropical Africa, and 

Sri Lanka (Burkill 1994, Howard 1979, Judziewicz 

and others 1999). There is speculation that the 

species may be exotic in the Old World 

(Judziewicz and others 1999). 

Ecology.—Carricillo is moderately intolerant to 

intermediate in tolerance to shade. It is widespread 

and relatively common in the understory of thin 

canopy forests, brushy forests, small openings, and 

edges of high forest. These are most often medium 

to late-secondary forests but can be primary forests 

and remnants. Carricillo grows on a wide variety 

of well-drained to somewhat poorly drained soils 

with pH’s from near neutral to about 5.5 over 

sedimentary (including limestone), igneous, and 

metamorphic (including ultramafic) rocks. 

Elevation may vary from near sea level to 1,100 m 

in areas that receive from about 1000 to more than 

3000 mm of mean annual precipitation (author’s 

observation, Croat 1978, Shaka and others 1997, 

Stevens and others 2001). 

Reproduction.—Carricillo flowers and fruits 

primarily during the rainy season in Panama 

(Croat 1978) and throughout the year in Nicaragua 

(Stevens and others 2001). Seed production is not 

abundant in Puerto Rico, but may be in Costa Rica 

(Missouri Botanical Garden 2002). Seeds in two 

collections made in Puerto Rico on the same site 

on different years averaged 0.0210 + 0.0003 and 

0.0225 + 0.0003 g/seed or about 46,000 seeds/kg. 

Placed in moist potting mix, only 1 percent had 

germinated after 18 months. Scarification, 

alternate wetting and drying, and heat treatment all 
519

failed to stimulate germination (author’s 

observation). Birds disperse the seeds (Burkill 

1994, Judziewicz and others 1999). New plants are 

uncommon. Attempts to cultivate the species in 

temperate greenhouses have failed (Judziewicz and 

others 1999). 

Growth and Management.—Carricillo is a 

relatively fast growing plant. Individual culms 

grow 2 m or more during their first year and 1 m 

or so thereafter and live 2 to 4 years. By continual 

sprouting, plants may live for many years. 

Artificial vegetative propagation has not been 

explored. Until vegetative methods are developed 

or the extended dormancy problem of seeds is 

solved, planting is not advised. Eradication can be 

done by grubbing out individual plants or probably 

by cutting and spraying the sprouts with 

glyphosate or other grass herbicides. 

Benefits.—Carricillo contributes to the diversity 

of the forest, helps protect the soil, and furnishes 

food and cover for wildlife. Cattle eat the leaves 

and fine twigs. Birds eat the fertile florets and 

fruits, although there is speculation that they may 

not be able to digest the hardened seeds (Burkill 

1994). The species is the food plant for larva of the 

moth Eryphanis reevesii (Doubleday) (North 

Carolina Botanical Garden 2002). The hollow 

culms are used for flutes (Judziewicz and others 

1999) and drinking straws. Leaves, culms, roots, 

and seeds are used in a number of applications in 

herbal medicine (Burkill 1994). Blades made from 

the culms are used by the Cuiba tribe in Colombia 

to cut the umbilical cords of newborns (Judziewicz 

and others 1999). 

References

Burkill, H.M. 1994. The useful plants of West 

Tropical Africa. Vol. 2. Royal Botanic Gardens, 

Kew, UK. 636 p. 

Croat, T.B. 1978. Flora of Barro Colorado Island. 

Stanford University Press, Stanford, CA. 943 p. 

Howard, R.A. 1979. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 3. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 586 p. 

Judziewicz, E.J., J.G. Clark, X. Londoño, and M.J. 

Stern. 1999. American bamboos. Smithsonian 

Institution Press, Washington, DC. 

http://email.uwsp.edu/publicanon/Course%20Inf

ormation/200120%-%20Spring%202002/ 

Biology/PF-BIOL345_xF8FF_545.1_200120/ 

American%20Bamboos%20book.doc?Cmd= 

open. 175 p. 

Missouri Botanical Garden. 2002. Manual de la 

flora de Costa Rica. Missouri Botanical Garden, 

St. Louis, MO. http://www.mobot.org/manual. 

plantas/050531/S050880.html. 7 p. 

North Carolina Botanical Garden. 2002. The biota 

of North America program: Olyra. North 

Carolina Botanical Garden, Chapel Hill, NC. 

http://www.funet.fi/pub/sci/bio/life/plants/magno

liophyta/magnoliophytina/liliopsida/poaceae/ 

olyra/ 2p. 

Shaka, J.M., W. Kabushemera, and A. Msangi. 

1997. Soils and vegetation of Semdoe proposed 

forest reserve Bombwera Division, Muheza 

District, Tanga. Tech. Paper 36. Ministry of 

Agriculture, National Soil Service, Agricultural 

Research Institute, Mlingano, Tanganyika. 14 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanic Garden Press, St. Louis, 

MO. p. 1,911-2,666. 
520 

Opuntia leptocaulis DC. desert Christmas cactus

CACTACEAE 

Synonyms: Cylindropuntia leptocaulis (DC.) D.M. Knuth 

 Opuntia ramulifera Salm Dyck 

 Cylindropuntia leptocaulis var. stipata Coulter 

 Cylindropuntia leptocaulis var. longispina Berger 

Juanita A. R. Ladyman 

 

General Description.—The common name for 

Opuntia leptocaulis is desert Christmas cactus. 

This name is likely to have originated because the 

green -colored fruits turn a very festive red color in 

December just in time for Christmas. Other 

common names include slender-stem cactus, 

turkey pear, tasajillo, aguijilla and garrambulla 

(Taylor and others 1997, Weniger 1991). In the 

indigenous language of Nahuatl it is called 

tzazahuistli (Crook and Mottram 1999). A widely 

applied name that includes all the long jointed 

cylindropuntia cacti, of which desert Christmas 

cactus is one, is cholla. The genus name, Opuntia, 

is taken from the name given to succulent plants 

that were originally from the old Greek town of 

Opus. The species name, leptocaulis means 

slender stemmed in Greek (Crook and Mottram 

1999). Desert Christmas cactus is an erect shrub 

that is usually less than 1.5 m tall but can grow up 

to 2.8 m in shady, protected environments such as 

under trees (Earle 1990). Although relatively 

compact, it is extensively branched from a main 

trunk that is covered by scaly bark in old age. It 

has pencil-like woody joints that are 12 to 40 cm 

long and 5 mm in diameter and can be easily 

detached from the plant. The tubercules, usually 

less than 1 cm long, may be indistinct and the 

joints smooth. The areoles, which are the small, 

clearly defined raised areas on the surface, are oval 

or almost diamond-shaped with short white wool. 

The areoles may be spineless or have from one to 

three spines that are from 19 to 51 mm long. There 

are few, usually short, yellowish-brown glochids, 

or minute bristles, in one to three small clusters in 

the upper part of the areole. The leaves are green, 

less than 1.3 cm long, and are early deciduous. The 

greenish-yellow flowers are approximately 1.3 to 

1.9 cm in diameter and up to 2.5 cm tall. The fruits 

are up to 2.5 cm long, and smooth with brown 

glochids in the areoles. There are usually less than 

12 seeds per fruit (Earle 1990, Weniger 1991, 

Vines 1986). Desert Christmas cactus is a diploid 

species where n = 11 (McGregor and others 1986). 

Range.—Desert Christmas cactus occurs in 

Arizona, New Mexico, Oklahoma, and Texas in 

the U.S.A. and in Sonora and Chihuahua in 

northern Mexico (Bowers 1993, McGregor and 

others 1986, Powell 1998). 

Ecology.—Desert Christmas cactus is a water 

storer and employs a type of photosynthesis called 

crassulacean acid metabolism that allows them to 

make efficient use of available water (Burgess 

1995, Nobel 1988). The species is found at 

elevations below 1,524 m. It grows in most soil 

types but appears to be most successful in 

gravelly, sandy, and heavier bottomland soils 

(Taylor and others 1997, Epple 1995, Rondeau and 

others 1996, Bowers and McLaughlin 1987, 

Kearney and others 1960). It is commonly in the 

same habitat with creosotebush, Larrea tridentata
521

(DC) Colville, with which it has a cyclical 

relationship in the Chihuahuan desert (Yeaton 

1978). Creosotebush colonizes sites and modifies 

microhabitat and edaphic conditions for desert 

Christmas cactus establishment and is eventually 

replaced by it. However, a combination of 

burrowing rodents and soil erosion increases the 

mortality rate of desert Christmas cactus while 

having little effect on creosotebush. The openings 

left by the dead cactus are then re-colonized by 

creosotebush. 

Reproduction.—Desert Christmas cactus 

reproduces both sexually by seed and vegetatively. 

Flowers generally appear in May to July 

depending upon regional and environmental 

conditions. In response to wet weather, they can 

flower as early as April and again in August. The 

flowers open wide in late evening (Taylor and 

others 1997). The fruit ripens in December and 

persists on the plant until approximately February. 

Seeds may sprout while in the fruit and have a 

shoot of up to 7 cm long while still on the plant 

(Weniger 1991). Detached joints root and grow 

into plants. The terminal stem segments detach 

particularly easily and readily take root near the 

parent plant so that natural populations that appear 

to be dense monocultures of clonal individuals 

develop (Rebman and Pinkava 2001). Cuttings 

taken from woody stems do not root easily (Earle 

1990). Hybrids between cholla species are 

common in nature (Rondeau and other 1996). An 

example herbarium specimen of such a hybrid, O. 

leptocaulis x O. spinosior (cane cholla), can be 

seen on the Internet (Pinkava 2001). 

Growth and Management.—Desert Christmas 

cactus is frequently abundant on overgrazed range 

(Powell 1998). It tends to form thickets that can 

become almost impenetrable to humans and 

livestock. Individual plants can be long-lived and 

may be more than 53 years old (Goldberg and 

Turner 1986). In a study over a 72-year period, 40 

percent of plants were determined to survive at 

least 7 years (Goldberg and Turner 1986). 

Survival after fire is dependent upon the intensity 

and frequency of the fire (Thomas 1991, Cave and 

Patten 1994). Plants can re-sprout after the fire 

kills the above ground parts (Bunting and others 

1980). However, desert Christmas cactus appears 

more susceptible to direct damage from fire than 

some other Opuntia species (Bunting and others 

1980). After a fire in a mixed-grass prairie in 

Texas, mortality was 65 percent in the first year 

and reached a cumulative value of 80 percent by 

the fourth year (Bunting and others 1980). Desert 

Christmas cactus recovery appeared particularly 

sensitive to the amount of precipitation. 

Precipitation was required before plants would resprout. Plants tolerate minimum temperatures of 

7.3 o

C in cultivation (Innes and Glass 1991). In 

cultivation, desert Christmas cactus is susceptible 

to the cochineal bug and periodic oil spray is 

recommended (Earle 1990). 

Benefits.—Weniger (1991) describes it as a major 

pest and “probably one of the most hated cacti in 

our area [Texas].” This is an unfortunate sentiment 

because, although the spines can impale people 

and livestock that venture too close, it provides 

significant food, protected nesting sites, and cover 

for a variety of wildlife (Earle 1990, Dayton 

1931). Most birds, including the bobwhite quail 

and wild turkey, as well as small mammals, favor 

the fruit (Powell 1998, Taylor and others 1997). 

Some animals, such as white-tailed dear, consume 

the joints as well as the fruit (Everitt and Drawe 

1993). There is apparently no seasonal variation 

in the crude protein content of the stems, which is 

similar to the fruits; both are 8 percent (Taylor and 

others 1997). The larger shrubs provide cover and 

nesting sites, especially for the cactus wren. In 

addition, there is now a commercial value to 

maintaining these shrubs. Land management for 

wildlife has become increasingly important to 

livestock producers in the Western U.S.A. because 

of fluctuations in livestock prices. In some parts of 

Texas, income generated from consumptive (e.g., 

hunting) and non-consumptive (e.g., bird 

watching) wildlife activities already exceed 

revenue obtained through livestock operations 

(Hernandez and others 1999). Under these 

circumstances upland gamebirds, such as northern 

bobwhites and Rio Grande wild turkeys, have 

become particularly important. Recommendations 

have been made that low, thorny brush composed 

of species such as lotebush [Ziziphus obtusifolia 

(Hook. ex Torr. & Gray) Gray] and desert 

Christmas cactus should be protected in order to 

enhance nesting cover (Hernandez and others 

1999) and provide a food source. Desert Christmas 

cactus also provides refugia to native plant species. 

A clear example of this on a landscape scale is in 

Arizona where clumps of tobosa grass are almost 

completely restricted to refugia that are most 

commonly under the desert Christmas cactus 

canopy where it is inaccessible to livestock 

(McAuliffe 1995). Native Americans gathered the 

fruits and ate them raw or cooked them into a jam 

(Earle 1990). The Apache tribes mixed the 
522 

crushed fruits with a beverage to produce narcotic 

effects (Moerman 1998). 

References 

Bowers, J.E. 1993. Shrubs and trees of the 

Southwest Deserts. Southwest Parks and 

Monuments Assoc. Tucson, AZ. 140 p. 

Bowers, J.E. and S.P. McLaughlin. 1987. Flora 

and vegetation of the Rincon Mountains, Pima 

County, Arizona. Desert Plants 8(2): 51-94 

Bunting, S.C., H.A. Wright, and L.F. Neuenschwander. 1980. Long-term effects of fire on 

cactus in southern mixed prairie of Texas. 

Journal of Range Management 33(2): 85-88 

Burgess, T.L. 1995. The dilemma of coexisting 

growth forms. In: M.P. McClaran and T.R. Van 

Devender, eds. The desert grassland. University 

of Arizona Press, Tucson, AZ. p. 31-67 

Cave, G.H. and D.T. Patten. 1994. Short-term 

vegetation responses to fire in the upper Sonoran 

desert. Journal of Range Management 37(6): 

491-496 

Crook, R. and R. Mottram, 1999. Opuntia Index – 

Part 5: Nomenclatural note and I-L. Bradleya 

17: 109-131 

Dayton, W.A. 1931. Important western browse 

plants. Misc. Publication 101. U.S. Department 

of Agriculture, Washington, DC. 214 p. 

Earle, W.H. 1990. Cacti of the Southwest. Desert 

Botanical Garden, Phoenix, AZ. 210 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. Falcon Press Publishing Co., Helena, 

MT. 347 p. 

Everitt, J.H. and D.L. Drawe. 1993. Trees, Shrubs 

and Cacti of South Texas. Texas Tech 

University Press, Lubbock, TX. 213 p. 

Goldberg, D.E. and R.M. Turner. 1986. 

Vegetation change and plant demography in 

permanent plots in the Sonoran Desert. Ecology 

67(3): 695-712 

Hernandez, F., C. Kleberg, and A. Radomski. 

1999. Northern bobwhite and Rio Grande turkey 

management in relation to livestock grazing in 

semiarid environments. Proceedings from 

Northern Texas and South Texas Wildlife 

Conservation and Management Workshop, 

Saltillo, Mexico. August 19-20. Internet site: 

http://cnrit.tamu.edu. 

Innes, C. and C. Glass. 1991. Cacti. Portland 

House, New York. 320 p. 

Kearney, T.H., R.H. Peebles, and collaborators. 

1960. Arizona flora. 2nd Ed. University of 

California Press, Berkeley, CA. 1,085 p. 

McAuliffe, J.R. 1995. Landscape evolution, soil 

formation, and Arizon’s desert grassland. In: 

M.P. McClaran and T.R. Van Devender, eds. 

The desert grassland. The University of Arizona 

Press, Tucson, AZ. p. 100-129. 

McGregor, R.L., T.M. Barkley, RE Brooks, EK 

Schofield. 1986. Flora of the Great Plains. 

University Press of Kansas, Lawrence, KN. 

1,402p. 

Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. 927 p. 

Nobel, P.S. 1988. Environmental biology of 

agaves and cacti. Cambridge University Press, 

New York. 288 p. 

Pinkava, D. 2001. Biologist profile, specimens at 

Arizona State University herbarium. 

http://lsvl.la.asu.edu/askabiologist/profiles/ 

pinkava/pink.html. 

Powell, A.M. 1998. Trees & shrubs of TransPecos and adjacent areas. University of Texas 

University Press, Austin, TX . 498 p. 

Rebman, J.P., and D. J. Pinkava. 2001. Opuntia 

cacti of North America–An overview. Florida 

Entomologist 84 (4): 474-483 

Rondeau, R., T.R. Van Devender, C.D. Bertelsen, 

P. Jenkins, R.K. Wilson, and M.A. Dimmitt. 

1996. Annotated flora and vegetation of the 

Tucson Mountains, Pima County, Arizona. 

Desert Plants 12(2): 3-46 

Taylor, R.B., J. Zrutledge, and J.G. Herrera. 1997. 

A field guide to common south Texas shrubs. 

Texas Parks and Wildlife Press, Austin, TX. 

p. 106. 
523

Thomas, P.A. 1991. Response of succulents to 

fire: A review. International Journal of Wildland 

Fire 1(1): 11-22. 

Vines, R.A. 1986. Trees, shrubs, and woody vines 

of the Southwest. Sixth printing. University of 

Texas Press. Austin, TX. 1,104 p. 

Weniger, D. 1991. Cacti of Texas and neighboring 

states. University of Texas Press, Austin, TX. 

356 p. 

Yeaton, R.I. 1978. A cyclical relationship between 

Larrea tridentata and Opuntia leptocaulis in the 

northern Chihuahuan desert. Journal of Ecology. 

66: 651-656. 
524 

Palicourea crocea (Sw.) J.A. Schultes cachimbo 

RUBIACEAE 

Synonyms: Palicourea coccinea Poiteau ex DC. 

 Psychotria crocea Sw. 

 Palicourea reparia Benth. 

 Palicourea croceoides Ham. 

 Palicourea crocea DC. var. riparia (Benth.) Griseb. 

 Palicourea brevithyrsa Britt. & Standl. 

John K. Francis 

General Description.—Cachimbo is a medium to 

large shrub also known as red palicourea, yellow 

palicourea, yellow-cedar, tapa camino, ponasí, 

bois de l’encore, bois cabrit, and bois fou-fou 

(Little and others 1974). There is some ambiguity 

in the classification of this species. Liogier (1997) 

separates P. crocea and P. croceoides on the basis 

of flower color; Howard (1989) treats them as a 

single species under P. crocea. We will follow 

Howard for the purposes of this description. The 

stems of cachimbo are green when young, 

changing to gray. The inner bark remains green. 

The wood is moderately hard and brittle. There are 

usually several sprouts from the base and long 

straight branches arising from major stems. 

Cachimbo develops a robust, shallow lateral root 

system with abundant fine roots. Foliage is 

relatively sparse and concentrated near the 

growing tips. The plant is glabrous or nearly so. 

The leaves are elliptic, 7.0 to 14.5 cm long and 3.5 

to 5.5 cm broad, and pointed at both ends with a 

petiole 0.5 to 1.0 cm long. The tubular flowers are 

grouped in cymes. Flower color ranges from 

yellow to red. The fruits are ovoid to globose, 4 to 

6 mm in diameter and dark red, purple, or black. 

They are faintly sweet with a grassy flavor. There 

are two seeds per fruit when perfectly developed. 

The seeds are black or grayish with a hard, bony 

shell, and hemispherical, with a groove on the flat 

side and three ridges on the rounded side. 

Range.—The natural range of cachimbo extends 

from southern Mexico through Central America 

and South America to southern Brazil and 

Paraguay on the mainland and includes the 

Greater and Lesser Antilles in the Caribbean 

(Howard 1989, Liogier 1997) 

Ecology.—Cachimbo grows in areas receiving 

precipitations ranging from 1500 to over 4000 

mm/yr. Soils where it grows are usually loams to 

clays derived from volcanic or sedimentary parent 

material. Elevations may be a few meters above 

sea level up to 1,000 m (Little and others 1974). 

Cachimbo requires disturbance to establish itself. 

It grows well in tree-fall gaps, landslides, artificial 

openings, and under thin forest canopies. It 

competes aggressively with forbs, other shrubs, 

and young trees. When the canopies close and 

shade becomes dense, existing plants decline and 

eventually disappear. Labrón (1977) calls the 

species a “gap opportunist.” Cachimbo was a 

principal species in abandoned pastures in the 

Luquillo Mountains, Puerto Rico, 60 years ago, 

but not in abandoned shade coffee plantations 

(Zimmerman and others 1995). The species grows 

slowly on nutrient-poor substrate (as in landslide 

areas), but responds to N and P fertilization under 

those conditions (Fetcher and others 1996). 
525

Reproduction.—Cachimbo flowers and fruits 

irregularly throughout the year (Little and others 

1974). The fruits in one Puerto Rican sample 

weighed an average of 0.1488 + 0.0055 g/fruit. 

Seeds cleaned from the sample averaged 0.0314 +

0.0048 g/seed or 32,000 seeds/kg. Thirty-five 

percent of these seeds germinated between 39 and 

84 days after sowing in commercial potting mix. 

Palicourea flowers are pollinated by humming 

birds and the fruits are dispersed by frugiverous 

birds (Taylor 1996). Bees were also observed to 

visit the flowers (Labrón 1977). Cachimbo suffers 

relatively little seed loss to pathogens and 

predators and germinates at a higher percent after 

natural dispersal than without it (Myster 1997). 

Seedlings are common and widespread in suitable 

habitat. There is no significant difference between 

seasons in seedling appearance, but regardless of 

season, seed germination is always higher in the 

open than in closed canopy areas (Labrón 1977). 

Transplanted wildlings survive and grow poorly 

(personal communication with F.N. Scatena, IITF, 

Río Piedras, PR) 

Growth and Management.—Cachimbo grows 

slowly (a few cm) the first year and rapidly (1 m 

or more per year) for about 2 years in openings 

(Labrón 1977). After about 3 years, seedlings 

sown under an open canopy had accumulated 

about 20 times more biomass than seedlings sown 

under closed canopy conditions. The aboveground to below-ground ratio of the open-canopy 

plants was 4.8 (Labrón 1977). Cachimbo may 

reach 6 m in height and 7.5 cm in stem diameter 

(Howard 1989, Little and others 1974). No annual 

rings are visible. However, single stems last about 

10 years, and plants can continue to regenerate 

with new sprouts. Plants cut during plantation 

cleaning and trail clearing quickly regain their 

former height (author’s observation). Scatena and 

others (1993) present an equation to predict ovendry above-ground biomass (To) from diameter at 

breast height (D) and height (H): To = exp(0.752 

ln (D2

H) – 2.362) where R2 = 0.861. 

Benefits.—Cachimbo is a food source for several 

species of song birds (Devoe 1989). It is higher in 

the major nutrients in its tissues than most of the 

trees and shrubs with which it is associated 

(Scatena and others 1993) and is probably a good 

recycler of nutrients and a good forage plant. 

People in some areas of Puerto Rico use crushed 

cachimbo leaves to stop bleeding (Labrón 1977). 

References 

Devoe, N.N. 1989. Differential seeding and 

regeneration in openings and beneath closed 

canopy in sub-tropical wet forest. Ph.D. 

dissertation. Yale University, New Haven, CN. 

307 p. 

Fetcher, N., B.L. Haines, R.A. Cordero, D.J. 

Lodge, L.R. Walker, D.S. Fernandez, and W.T. 

Lawrence. 1996. Responses of tropical plants to 

nutrients and light on a landslide in Puerto Rico. 

Journal of Ecology 84(3): 331-341. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Labrón, M. L. 1977. An autoecological study of 

Palicourea riparia Bentham (RUBIACEAE): an 

ecologically important species in the recovery of 

a disturbed tropical rain forest in Puerto Rico. 

Ph.D. Dissertation. University of North 

Carolina, Chapel Hill, NC. 238 p. 

Liogier, H. A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Myster, R.W. 1997. Seed predation, disease and 

germination on landslides in neotropical lower 

montane wet forest. Journal of Vegetation 

Science 8(1): 55-64. 

Scatena, F.N., W. Silver, T. Siccama, A. Johnson, 

and M.J. Sánchez. 1993. Biomass and nutrient 

content of the Bisley Experimental Watershed, 

Luquillo Experimental Forest, Puerto Rico, 

before and after Hurricane Hugo, 1989. 

Biotropica 25(1): 15-27. 

Taylor, C.M. 1996. Palicourea (Rubiaceae). 

www.mobot.org/MOBOT/Staff/Research/taylor 

//palihome.html. 9 p. 
526 

Zimmerman, J.K., T.M. Aide, M. Rosario, M. 

Serrano, and L. Herrera. 1995. Effects of land 

management and a recent hurricane on forest 

structure and composition in the Luquillo 

Experimental Forest, Puerto Rico. Forest 

Ecology and Management 77(1-3): 65-76. 
527

Parathesis crenulata (Vent.) Hook. f. rascagarganta 

MYRSINACEAE 

Synonyms: Ardisia crenulata Vent. 

John K. Francis 

General Description.—Rascagarganta, also 

known as secagarganta, jalapón, and raisin 

marron, is a shrub or small tree 1 to 3 m 

(occasionally reaching 6 m) in height and 1 to 4 

cm (occasionally to 7.5 cm) in trunk diameter. The 

common names in Spanish imply that it will make 

one’s throat raspy or dry. The species, unless 

disturbed, has a single stem and is supported by a 

taproot and lateral roots that are orange-brown in 

color. Most of the branches arise from the main 

stem, often in whorls of three’s or four’s. The 

alternate, dark-green, 7- to 23-cm leaves are 

narrowly elliptic, pointed at both ends, and often 

have wavy-toothed edges. The small pink flowers 

are borne in large numbers in terminal panicles. 

The 6- to 8-mm fleshy fruits progress from green 

to pink to red to black as they ripen. Each fruit 

contains one seed (Howard 1989, Liogier 1995, 

Little and others 1974). 

Range.—Rascagarganta is native to Hispaniola, 

Puerto Rico, and Martinique (Howard 1989, 

Liogier 1995, Little and others 1974). Some 

taxonomists suggest that P. serrulata (Sw.) Mez, 

of Cuba and Hispaniola is synonymous with P. 

crenulata (Howard 1989). 

Ecology.—Rascagarganta grows in moist forests 

that receive from about 1500 mm to 2800 mm of 

annual precipitation. In the dry end of the 

precipitation range, it only occurs near streams 

and extra moist hollows. The species is found 

mainly on slightly- to moderately-acid soils with 

loamy to clayey texture. Although areas with both 

igneous and sedimentary rocks are colonized, it is 

most abundant in valleys in limestone formations. 

Rascagarganta grows from a little above sea level 

to 760 m in elevation (Little and others 1974). The 

species is shade tolerant and prefers to grow under 

closed forests with moderate to light understory 

vegetation. It sometimes forms its own understory 

canopy under which little vegetation grows. There 

appear to be few insect or disease problems in 

natural stands. 

Reproduction.—Rascagarganta is reported to 

flower from May to July and fruits from July to 

December (Little and others 1974). Fruits are 

often present in quantity and are easy to pick. 

They may be cleaned by maceration and wet 

sieving. A collection of fruits from Puerto Rico 

weighed an average of 0.407 g/fruit. Air-dried 

seeds cleaned from that collection weighed an 

average of 0.054 g/seed or 18,000 seeds/kg. Sown 

in peat, 45 percent germinated between 48 and 

104 days after sowing. Many seedlings died from 

damping-off; therefore, a looser germination 

medium than peat is recommended. Seedlings 

grew slowly in the nursery, reaching less than 10 

cm in 6 months (author’s observation). Birds 

disperse the seeds. Seedlings are relatively 

common and widespread near fruit-bearing stands. 

Growth and Management.—Rascagarganta 

grows slowly and appears to live several decades. 

No management experience has been published. 

However, the best strategy for establishment of 

natural stands may be to maintain plantations or 

natural forest stands in moist areas for long 

periods with closed canopies and relatively clear 

understories to allow rascagarganta to build up. 

Benefits.—Rascagarganta contributes to 

biodiversity, cover for wildlife, and protects the 

soil from erosion in the forests where it grows. 

Although not usually heavily used by wildlife, the 

fruits are produced in large quantity and are 

available after hurricanes when overstory fruits are 
528 

scarce. The wood is hard, but due to its small size 

is useful mainly for stakes and fuel. 

References 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

617 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 
529

Parkinsonia aculeata L. Jerusalem-thorn 

FABACEAE 

Synonyms: none 

John K. Francis 

General Description.—Jerusalem-thorn occupies 

the transition zone between shrub and tree lifeforms. In difficult sites and portions of its range, it 

is very shrubby, but in moist and fertile areas, it 

becomes a small tree up to 10 m in height. 

Jerusalem-thorn often has multiple stems, 

especially if burnt or cut, and usually has a 

branchy, low, and diffuse crown. The branches 

and foliage are often somewhat pendulous. It has 

a taproot, carrot-like in young plants, and slender 

laterals radiating out from it at all depths. Old 

trunks are brown and fissured or scaly. The bark 

of smaller stems and twigs is thin, smooth, and 

yellow-green or blue-green. There are paired 

spines at the nodes and a larger spine (1 to 2 cm 

long) at the end of the leaf axis. The leaves are 

alternate, bipinnately compound (20 to 40 cm 

long) with a flattened rachis and many tiny (2 to 4 

mm long) leaflets that are shed during the winter 

or the dry season. The yellow or yellow and 

orange fragrant flowers are grouped in racemes 

and have five sepals and five petals. The fruits are 

legumes 3 to 10 cm long, slightly flattened with 

constrictions between each seed. The seeds are 

one to several in number, ellipsoid, gray, and 

about 4 by 8 mm. The species has a haploid 

chromosome number of 14 (Pantulu 1942). 

Jerusalem-thorn is also known as horse bean, palo 

verde, retama, palo de rayo, espinillo, sulfato, 

cina-cina, mataburro, madam naiz, and arrêteboeuf (Correll and Johnston 1970, Howard 1988, 

Liogier 1988, Little and Wadsworth 1964). 

Range.—Jerusalem-thorn is native to Texas, 

Arizona, Mexico, Central America, and South 

America as far south as the north of Uruguay 

(Holdridge and Poveda 1975, Little and 

Wadsworth 1964, Piaggio 2001). It has been 

planted and has naturalized in many of the islands 

of the Caribbean, including Puerto Rico and the 

U.S. Virgin Islands (Liogier 1988). The species 

has been introduced into the Old World and has 

become widespread in Australia. Jerusalem-thorn 

is planted as an ornamental in Florida, and the 

Southern and Western United States. 

Ecology.—Jerusalem-thorn will grow on most 

types of soils, including sand dunes, clay soils, 

strongly alkaline, chalky, and mildly salty soils. It 

withstands a great deal of heat and can survive in 

areas that receive less than 30 cm of annual 

rainfall. Jerusalem-thorn is damaged by 

temperatures below –8 °C (Floridata 2001). 

Individual plants are attacked and damaged by 

termites, scales, and a number of other insect 

species, but populations are generally relatively 

free of pests. It requires full or nearly full sunlight 

to survive and reproduce. In most environments, 

Jerusalem-thorn needs some kind of disturbance to 

become established. 

Reproduction.—Jerusalem-thorn flowers and 

fruits throughout the year in Puerto Rico (Little 

and Wadsworth 1964). It often flowers and fruits 

at 2 m in height or less. The flowers are pollinated 

by bees that are attracted to and orient themselves 

to the banner petal, which absorbs UV radiation 

(Jones and Buchmann 1974). Seeds collected in 

Puerto Rico by the author numbered 13,300 

seeds/kg. Little and Wadsworth (1964) report 

12,300 seeds/kg. Seeds from the former collection 
530 

were scarified and germinated at 59 percent, 

beginning 2 days after sowing (Francis and 

Rodríguez 1993). Jerusalem-thorn produces two 

kinds of seeds. About 25 percent of them have thin 

testae and will germinate readily without 

pretreatment; the rest have hard seed coats and 

must be scarified before they will germinate 

(Floridata 2001). Mechanical scarification was 

used by the author; soaking for 45 minutes in 

concentrated sulfuric acid worked equally well in 

another test (Everitt 1983). In nature, seeds are 

transported by water, birds, and animals (Pacific 

Island Ecosystems at Risk 2001). Artificial 

propagation is routinely done with seeds followed 

by ordinary nursery culture in containers. In vitro 

propagation using nodal explants has been 

demonstrated (Jaideep and others 1992). 

Jerusalem-thorn will continue to resprout after 

many cycles of annual disturbance. 

Growth and Management.—The growth rate is 

moderate, from 0.5 to 1 m per year in early years. 

Jerusalem-thorn plants live about 30 years (Pima 

Community College 2001). As an exotic, 

Jerusalem-thorn can be problematic in rangeland, 

as it has become in Australia. Biological control 

attempts are ongoing, but have not yet been 

successful. Jerusalem-thorn can be controlled with 

tractors by pulling or dozing the large plants out, if 

the seedlings are also controlled. Good control has 

been achieved with several herbicides as well 

(Pacific Island Ecosystems at Risk 2001). 

Benefits.—The principal use of Jerusalem-thorn 

today is for landscaping. Its abundant yellow floral 

display alone justifies its use. Perhaps its best use 

is in xeric gardens, where it contrasts well with 

succulents and other desert plants (Floridata 

2001). It is also useful in hedges and for living 

fenceposts (Hoekstra and others 2001). Jerusalemthorn is also planted for environmental restoration 

in desert areas. Besides being hardy, it has the 

added benefit of fixing nitrogen (Harris 1982). 

The wood is moderately hard, heavy (specific 

gravity 0.6), and brittle. It is used principally for 

fuel. Native Americans harvested the seeds, which 

they parched before eating (Floridata 2001). 

Various extracts of leaves, flowers, fruits, and 

bark are used in herbal medicine to treat arthritis 

and fever, and as a nerve stimulant and abortive 

(Liogier 1990). Livestock eat the foliage and fruits 

(Little and Wadsworth 1964). Jerusalem-thorn is 

occasionally browsed by white-tailed deer and the 

seeds are eaten by bobwhite quail (Everitt and 

Drawe 1993). 

References 

Correll, D.S., and M.C. Johnston. 1970. Manuel of 

the vascular plants of Texas. Texas Research 

Foundation, Renner, TX. 1,881 p. 

Everitt, J.H. 1983. Seed germination 

characteristics of two woody legumes (retama 

and twisted Acacia) from South Texas. Journal 

of Range Management 36(4): 411-414. 

Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs 

and cacti of South Texas. Texas Tech University 

Press. 213 p. 

Floridata. 2001. Plant profile: Parkinsonia 

aculeata. http://www.floridata.com/ref/p/ 

park_acu.cfm. 3 p. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station. New 

Orleans, LA. 5 p. 

Harris, S.C. 1982. Nitrogen fixation by tropical 

woody legumes: potential source of soil 

enrichment. In: J.P. Roskoski, J. Monano, C. van 

Kessel, G. Castilleja, and P.H. Graham, eds. 

Biological nitrogen fixation technology for 

tropical agriculture. Centro International de 

Agricultura Tropical. Cali, Colomabia. p. 447-

454. 

Hoekstra, D.A., P.J. Wood, and F. Anap Sang. 

2001. Proposed second phase dryland 

agroforestry research project. 

http://www.idrc.ca/library/document/031091/ 

31 p. 

Holdridge, L.R. and L.J. Poveda A. 1975. Arboles 

de Costa Rica. Vol. 1. Centro Científico 

Tropical, San José, Costa Rica. 546 p. 

Howard, R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain, 

MA: Arnold Arboretum, Harvard University. 

673 p. 

Jaideep. M., S. Mukunthakumar, and J. Mathur. 

1992. Micropropagation of Bauhinia variegata

and Parkinsonia aculeata from nodal explants of 

mature trees. Plant Cell, Tissue and Organ 
531

Culture 28(1): 119-121. 

Jones, C.E. and S.L. Buchmann. 1974. Ultraviolet 

floral patterns as functional orientation cues in 

hymonopterous pollination systems. Animal 

Behavior 22(2): 481-485. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamricana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr., and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

Pacific Island Ecosystems at Risk. 2001. Invasive 

plant species: Parkinsonia aculeata L. Fabaceae. 

http://www.hear.org/pier/paacu.htm. 2 p. 

Pantulu, J.V. 1942. Chromosome numbers of some 

Caesalpiniaceae. Current Science 11: 152-153. 

Piaggio, M. 2001. Plantas natives del Uruguay. 

Red Academica de Uruguay. 

http://www.rau.edu.uy/uruguay/flora/Uy.flora3.

htm. 8 p. 

Pima Community Collage. 2001. Desert ecology 

of Tucson: mexican paloverde (Parkinsonia 

aculeata). http://wc.pima.edu/Bfiero/ 

tucsonecology/plants/trees_mpv.htm. 1 p. 
532 

Parthenocissus quinquefolia (L.) Planch. Virginia creeper 

VITACEAE 

Synonyms: Amphelopsis latifolia Tausch 

 Amphelopsis quinquefolia (L.) Michx. 

 Hedera quinquefolia L. 

 Vitis quinquefolia (L.) Lam. 

 Parthenocissus hirsute (Pursh) Graebn. 

Parthenocissus inserta (Kerner) Fritsch (and other synonyms: Natural 

Resources Conservation Service 2003) 

John K. Francis 

General Description.—Virginia creeper, 

occasionally referred to as woodbine, thicket 

creeper, American ivy, and five-leaved ivy, is a 

deciduous woody vine that may reach 15 m or 

more into the crowns of trees or up masonry walls. 

Single or multiple stems are slender, round, light 

brown, and have prominent lenticels. Young 

growth is reddish. The vines adhere to surfaces by 

means of five- to eight-branched tendrils ending in 

cup-like, adhesive tips. The alternate leaves are 

palmately compound with five (sometimes three or 

four) leaflets. The leaflets are lanceolate, 

oblanceolate or obovate, 3 to 15 cm long, and 1.6 

to 8 cm broad with singly or doubly serrate 

margins. Petioles are 15 to 20 cm long, and leaflets 

have distinct petiolules. Small, greenish-white to 

yellowish-green flowers are borne in terminal 

cymes or panicles positioned under the foliage. 

Fruits are berries, somewhat flattened at the apex, 

4 to 6 mm in diameter, bluish-black with a definite 

bloom when ripe. Fruits usually contain two to 

three seeds (Dirr 2003, Nelson 1996, Rehder 1951, 

Welsh and others 1987). 

Range.—Virginia creeper is native to all the 

conterminous States except Washington, Oregon, 

California, Nevada, Arizona, New Mexico, Utah, 

Idaho, Montana, Wyoming, and North Dakota 

(Natural Resources Conservation Service 2003). It 

is also native to northern Mexico and southeastern 

Canada from Nova Scotia to Ontario (Coladoanto 

1991, Nelson 1996). It has naturalized in at least 

Utah, England, and France (Compton 2003, 

Services Culture Editions Ressources 2003, Welsh 

and others 1987). The species is cultivated as an 

ornamental in many moist temperate areas of the 

world. 

Ecology.—Virginia creeper is found in most of the 

vegetation associations in the Eastern and 

Midwestern United States (Coladoanto 1991). It 

grows in most moist soils types. Virginia creeper 

grows well in full sun but also tolerates shade. 

When growing under a forest canopy, it attempts 

to ascend the tree trunks or obstacles it encounters. 

It is a mid- to late-seral species. It is sensitive to 

fire and is top-killed by most fires. Root crowns 

and occasional surviving stems sprout and partially 

renew burned stands (Coladoanto 1991). 

Reproduction.—Virginia creeper flowers from 

June to August, matures fruits from August to 

October and drops fruits (or they are eaten by 

birds) from September to February. Good fruit and 

seed crops are frequent. There are 21,600 to 

58,000 air-dried seeds/kg of which 44 to 99 
533

percent are sound (Gill and Pogge 1974). The 

seeds are dispersed by birds (Dirr 2003). Seeds 

germinate the first or second spring after dispersal; 

germination is epigeal (Gill and Pogge 1974). It 

layers readily at the nodes when stems come in 

contact with the soil (Crandall and Crandall 1995). 

Growth and Management.—Once well 

established, Virginia creeper grows rapidly. It 

often must be pruned to prevent it from getting out 

of control in ornamental settings. It can seed into 

nearby flower beds and wildlands. Virginia 

creeper should not be planted to climb up 

buildings with wooden siding because it is difficult 

to remove and because it increases the humidity 

and hastens rot (Gilman 1999). The species 

occasionally interferes with reproduction of pines 

and hardwoods and may be controlled with 

herbicides. Fire can also be an effective control 

agent (Coladoanto 1991). Fruits can usually be 

picked in quantity by hand. Seeds are separated 

from fruit tissue by maceration followed by 

floating off the pulp and empty seeds with water. 

However, maceration must be gentle because seed 

coats are soft and easily damaged. There are 16 to 

27 g of seeds/100 g of fruits. The seeds may be 

fall-sown 1 cm deep in soil and mulched. 

Alternatively, seeds may be stratified in moist peat 

or sand at 5 or 6 °C for 60 days and sown in the 

spring. Forty-one to 94 percent of the seeds will 

germinate after 30 days. Seedlings are ready for 

outplanting when they reach 15 cm in height after 

1 or 2 years in the nursery. The species can also be 

propagated with cuttings and layers (Gill and 

Pogge 1974). 

Benefits.—Virginia creeper is important to the 

aesthetics of forests, helps protect the soil, and 

furnishes food and cover for wildlife. Cattle and 

white-tailed deer sometimes browse the foliage. 

Songbirds are the principal consumers of the fruits. 

The foliage, both as a ground cover and climber, 

provides cover for small mammals and birds 

(Coladoanto 1991). It has been planted for 

watershed protection and erosion control (Gill and 

Pogge 1974). The principal attraction of Virginia 

creeper is its fall color. The leaves turn fiery 

shades of purple, red, and scarlet after the first 

frost (Dirr 2003). The blue-black fruits and their 

stems (which turn red also) add color and interest 

after the leaves have fallen. The species is planted 

to cover building walls, run up tree trunks, and 

form trellises and arbors. It also makes a fine 

ground cover (Crandall and Crandall 1995). Native 

Americans took decoctions of the roots for 

diarrhea and difficult urination, infusions for 

jaundice, and applied decoctions and poultices 

topically for swelling and lockjaw (Moerman 

1986). The fruits are reported to be poisonous by 

reason of the oxalic acid they contain (Russell and 

others 1997). However, consuming more than a 

small amount is unlikely due to their bad flavor 

(author, personal observation). 

References

Coladoanto, M. 1991. Parthenocissus 

quinquefolia. In: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain 

Research Station, Fire Sciences Laboratory, Fire 

Effects Information System. http://www.fs.fed. 

us/database/feis/plants/vine/parqui/all.html. 

10 p. 

Compton, G. 2003. Cambridgeshire flora records 

since 1538. http://mnlg.com/gc/species2pz/ 

p/par_qui.html. [not paged]. 

Crandall, C. and B. Crandall. 1995. Flowering, 

fruiting, and foliage vines. Sterling Publishing 

Co., Inc., New York. 192 p. 

Dirr, M.A. 2003. Parthenocissus quinquefolia—

Virginia creeper, also called woodbine. 

http://www.nobleplants.com/classnotes/spring/ 

springprofiles/vines/parthenoquinque.htm. 1 p. 

Gill, J.D. and F.L. Pogge. 1974. Parthenocissus 

Planch., creeper. In: C.S. Schopmeyer, tech. 

coord. Seeds of woody plants in the United 

States. U.S. Department of Agriculture, Forest 

Service, Washington, DC. p. 568-571. 

Gilman, E.F. 1999. Parthenocissus quinquefolia. 

Fact Sheet FPS-454. Cooperative Extension 

Service, University of Florida, Gainesville, FL. 

3 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports. 19. University of 

Michigan Museum of Anthropology, Ann 

Arbor, MI. 534 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Parthenocissus quinquefolia (L.) 

Planch. http://plants.udsa.gov/cgi_bin/ 

plant_search.cgi?mode=Common+Name&keyw

ordquery=Vi.... 4 p. 
534 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Rehder, A. 1951. Manual of cultivated trees and 

shrubs hardy in North America. The MacMillan 

Company, New York. 996 p. 

Russell, A.B., J.W. Hardin, L. Grand, and A. 

Fraser. 1997. Poisonous plants: Parthenocissus 

quinquefolia. North Carolina State University, 

Raleigh, NC. http://www.ces.ncsu.edu/depts/ 

 hort/consumer/poison/Parthqu.htm. 2 p. 

Services Culture Editions Ressources. 2003. 

Parthenocissus quinquefolia (Linné) Planchon. 

http://crdp.ac-besancon.fr/ressourc/flore/flore/ 

especes/parth_quinq.htm. 1 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 
535

Passiflora edulis Sims passionfruit 

PASSIFLORACEAE 

Synonyms: none 

John K. Francis 

General Description.—Passionfruit is also 

known as parcha, granadilla, maracuyá, ceibey, 

lilikoi, and linmangkon (Morton 1987). The 

English prefix “passion” derives from the passion 

of Christ suggested by the prominent fourbranched style that appears in a few of the flowers. 

Passionfruit is a woody, perennial vine that bears a 

delicious fruit and occurs in purple- and yellowfruited forms (Passiflora edulis Sims f. edulis and 

P. edulis f. flavicarpa) known as purple and 

yellow passionfruits. The plants have a weak 

taproot and extensive ivory-colored lateral roots. 

The stem is usually solitary, up to 7 cm in basal 

diameter, extends 5 to 10 m or more into the 

crowns of trees, and is covered by a thin, flaky, 

light brown bark. The stemwood is light and 

brittle. The twigs are yellow-green, turning brown, 

and support themselves on vegetation by means of 

tendrils that arise at the leaf axils. The leaves are 

alternate, green to yellow-green, three-lobed (on 

mature plants) with serrate edges. The petioles are 

3 to 6 cm long and the blades are 5 to 11 cm long 

by 4 to 10 cm broad. Solitary flowers arise at the 

leaf axils. The flowers measure 5 to 7 cm across 

with five greenish-white sepals and five white 

petals topped with a fringe-like corona of straight 

purple and white rays. There are five stamens with 

large anthers and a triple-branched style. The fruit 

is globose or ovoid, purple or yellow and 4 to 7 

cm in diameter. Inside a thick rind are many darkbrown to black seeds enveloped in small sacs 

filled with aromatic yellow or orange juice. The 

fruits of the purple passion fruit are smaller but 

more aromatic than those of the yellow form 

(Acevedo-Rodríguez 1985, Liogier 1994, Morton 

1987). 

Range.—The purple passionfruit is a native of 

southern Brazil through Paraguay to northern 

Argentina. The yellow form of passionfruit is 

probably native of the Amazon region of Brazil 

(Morton 1987). Passionfruit is cultivated today 

throughout the tropics and subtropics and has 

naturalized and escaped in many areas including 

Florida, Puerto Rico, U.S. Virgin Islands, Hawaii, 

Guam, and Western Samoa (Acevedo-Rodríguez 

1985, Morton 1987, Pacific Island Ecosystems at 

Risk 2002). 

Ecology.—Altitude and latitude do not appear to be 

a constraint other than through the temperatures 

associated with them. Purple passionfruit grows 

best in a subtropical climate, and the yellow 

passionfruit prefers a tropical climate with fullseason warm days and nights. However, even 

yellow passionfruit can survive temperatures down 

to – 5 °C (Sentelhas and other 1996). Generally, 

annual rainfall should be at least 900 mm. Rainfall 

in Indian areas that grow passionfruit ranges from 

1000 to 2500 mm/year (Morton 1987). The species 

is shallow-rooted but withstands drought by 

defoliating. Passionfruit tolerates a wide variety of 

soils and grows best on well-drained sandy loams 

with pH’s of 6.5 to 7.5 (Morton 1987). Passionfruit 

is moderately intolerant of shade, requires trees, 

brush, or fences for support, and benefits from but 

does not require soil disturbance for reproduction. 

Wild plants are found in broken forests, streambottom galleries, fencerows, abandoned farms, and 

neglected city lots. Young plants are eaten by 

livestock, so passionfruit is almost never found in 

moderate to heavily grazed areas. A large number 

of insects, nematodes, fungi, and viruses attack the 

species (Morton 1987). 

Reproduction.—The purple passionfruit blooms 

in spring and early summer and again for a shorter 

period in fall and early winter (Morton 1987). 
536 

Yellow passionfruit in Puerto Rico flower from 

April to September and yield fruits from June to 

October (Acevedo-Rodríguez 1985). In some 

areas, plants fruit twice each year (Popenoe 1920). 

Plants usually begin blooming and fruiting in their 

second year. Yellow passionfruit flowers have 

both male and female parts but are self-sterile. 

They rely mainly on carpenter bees (Xylocopa

spp.) for pollination. Other insects and 

hummingbirds also visit the flowers. The flowers 

of purple passionfruit can self-pollinate (Morton 

1987). Fruits of the naturalized yellow-fruited 

form range from about 45 to 120 g in Puerto Rico. 

There is a large variation between plants in size 

and shape of fruits. Small fruits are sometimes 

completely devoid of seeds, and large fruits may 

have over 200 seeds. A collection of seeds from 

naturalized plants in Puerto Rico averaged 0.0251 

+ 0.0004 g/seed or 40,000 seeds/kg. Passionfruit is 

usually propagated from seeds but can be started 

from cuttings, layers, and grafts (Morton 1987). 

Seeds germinate best if allowed to ferment for a 

few days in the fruit pulp before cleaning and are 

lightly scarified by clipping or sandpapering 

(Morton 1987). A group of seeds in Puerto Rico 

were sown without pretreatment in commercial 

potting mix and began germinating in 14 days and 

completed germination in 24 days with 61 percent 

germinated. Plants are grown in beds or pots and 

transplanted when they reach about 25 cm in 

height. The seedlings are heavily watered after 

outplanting (Morton 1987). Seeds are disbursed in 

the wild by humans, animals especially pigs, and 

birds, and by vine extension. 

Growth and Management.—Plants in fertile soil 

extend their stems about 3 m per year. Each year 

during the annual dry season, the leaves fall off 

and the twigs die, leaving the main stem and a few 

important branches alive to rebuild the crown after 

the rains begin. Because fruiting takes place on 

new wood, light pruning does not reduce yield. 

Plants live from 3 to 8 years and do not resprout. 

Commercial stands are managed in vineyards, 

somewhat like grapes. They are planted in trellised 

rows 4.5 m apart and spaced 4.5 m apart within 

rows. The orchards are replanted every 4 to 6 

years (Bailey 1941). Alternately, vineyards may 

be established using small trees or bamboo as 

standards (Heenkenda and Punchikumarihami 

1991). Fruiting plants can even be grown in pots 

under glass (Anonymous. 2001). Fruits fall after 

ripening on the vines and are picked up from the 

ground at least twice per week. The roots normally 

form mycorrhizal associations and benefit from 

inoculation with superior strains of fungi 

(Cavalcante and others 2001). Wild plants are 

usually scattered and attempts at management 

have not been reported. 

Benefits.—By far the greatest benefit of 

passionfruit to humankind is its fruit and the 

delicious juice made from it. In addition to being 

collected by local people in the forests, the fruit is 

now grown in vineyards in dozens of countries. It 

is condensed, frozen, and shipped worldwide. The 

fruit pulp contains 2.2 percent protein, 0.7 percent 

fat, and 21.2 percent carbohydrates. In addition, 

the seeds contain 23 percent oil similar to 

sunflower or soybean oil, and the rind residue is 

used for cattle feed (Morton 1987). The fruits of 

native and naturalized stands furnish food for 

numerous species of wild mammals and birds. The 

whole plant, especially the leaves, contains 

alkaloids and a number of other phytoactive 

chemicals. Among these is passiflorine, a known 

sedative and tranquilizer (Morton 1987). Extracts 

of the leaves have been used for centuries as 

sedatives by native Brazilians. They prepare a 

drink from the flower to treat asthma, bronchitis, 

and whooping cough. The plant is also used as a 

diuretic to treat urinary infections (Rain-tree 

2002). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Anonymous. 2001. Exotic ornamental. The Hindu. 

http://www.hinduonnet.com/thehindu/2001/04/ 

22/stories/1322045j.htm. 3 p. 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. Vol. 3. The MacMillan Co. New 

York. p. 2,423-3,639. 

Cavalcante, U.M.T., L.C. Maia, C.M.C. Costa, and 

V.F. Santos. 2001. Mycorrhizal dependency of 

passion fruit (Passiflora edulis f. flavicarpa). 

Fruits 56: 317-324. 

Heenkenda, H.M.S. and S.M.A. 

Punchikumarihami. 1991. Alternate trellising 

materials for passion fruit (Passiflora edulis

Sims) vines. Tropical Agriculturist 147: 11-18. 
537

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Morton, J.F. 1987. Fruits of warm climates. 

Creative Resources Systems, Inc. Winterville, 

NC. 505 p. 

Pacific Island Ecosystems at Risk. 2002. Invasive 

plant species: Passiflora edulis Sims, 

Passifloraceae. http://hear.org/pier/paedu/htm. 

 2 p. 

Popenoe, W. 1920. Manual of tropical and 

subtropical fruits. The MacMillan Company. 

New York. 474 p. 

 

Rain-tree. 2002. Maracuja, “passion fruit”. 

http://rain-tree.com/passionf.htm. 3 p. 

Sentelhas, P.C., C.T.P. Junior, J.M.M Sigristi, R. 

Kavati, and M.T. Parodi. 1996. Freezing points 

of various tropical fruits. Bragantina 55(2): 231-

235. 
538 

Paullinia pinnata L. bejuco de costilla

SAPINDACEAE 

Synonyms: none 

John K. Francis 

General Description.—Bejuco de costilla is also 

known in Spanish-speaking areas as bejuco de 

paloma and bejuco de guajanilla and in Englishspeaking areas as supple jack. It is a stout and 

strong woody vine that forms mats in forest 

openings and ascends rocks and trees by means of 

forked tendrils to access full sunlight. It may reach 

10 m of extension (Vélez and Overbeek 1950) and 

12 cm or more of stem diameter. The name bejuco 

de costilla (rib vine) arises from the appearance of 

young stems. The young, green stems are 

angularly striated; the large, lower stems have a 

rounded triangular cross section. The plant is 

easily recognized from the leaves that have five 

serrated leaflets with prominent veins and a 

winged rachis and petiole. The fruits are red or 

dark pink when ripe (Acevedo-Rodriguez 1985, 

Howard 1989, Liogier 1994). 

Range.—Bejuco de costilla is native to Cuba, 

Hispaniola, Puerto Rico, the Lesser Antilles, 

Trinadad, Central America, South America, and 

tropical Africa (Howard 1989, Liogier 1994). 

Ecology.—Bejuco de costilla is found primarily in 

the moist (1000 to 2000 mm of annual rainfall) 

forests. It is also reported to occur in subtropical 

dry (less than 1000 mm precipitation) forest in 

Puerto Rico (Acevedo-Rodriguez 1985). Soils 

derived from limestone and other sedimentary 

rocks, and various igneous rocks at low and 

moderate elevations (below 400 m) are colonized. 

The species appears to be rare or absent from 

serpentine. It is present in both primary remnant 

and secondary forest stands. 

Reproduction.—In Puerto Rico, the species 

blooms from June to November and fruits from 

July to December (Acevedo-Rodriguez 1985). The 

fruits are capsules that split in three valves when 

ripe to expose one to three black seeds partially 

covered by a white aril. A collection of fruits in 

Puerto Rico weighed 1.674 + 0.0564 g. The fresh 

seeds recovered from those fruits weighed 0.150 +

0.008 g and gave 5 percent germination. The seeds 

are recalcitrant (drying kills them). A later 

collection carefully protected from drying gave 

100 percent germination. Germination is 

hypogenous. The seedlings grew rapidly and 

reached a maximum of 18 cm in 18 days after 

emergence. The seedlings quickly develop a deep 

and extensive tap and lateral root system. 

Moderate amounts of seed can be obtained by 

clipping the fruit clusters with a pruning pole and 

shelling out the seeds by hand. Seeds should not be 

picked up from the ground. Seed should be planted 

immediately or stored refrigerated in a closed 

plastic bag containing a moistened paper towel. 

Natural reproduction in forest stands is common to 

rare. Good survival and long life enable the 

species to maintain itself. More than one stem may 

arise from a horizontal root segment. Stems layer 

when they come in contact with the ground. 

Growth and Management.—If needed for 

establishment during environmental restoration 

projects, containerized plants can be easily and 

quickly grown from seed. However, no planting 

projects are known. In forest management activity, 

it is more often desirable to eliminate vines than 

establish them. An effective method in closed 

forest stands is to cut the stems of vines near the 

ground a year before harvest. Although the vines 

resprout, most of the species, being shade 
539

intolerant, die or are seriously weakened and do 

not pose a serious threat to the reproduction stand. 

Herbicides can also be applied to the sprouts to 

make the method more effective, especially in 

more open stands. Bejuco de costilla is one of the 

alternate hosts of the tsetse flies (Glossina palpalis

and G. tachnioides) and has become the target of 

eradication measures in some parts of tropical 

Africa (Morris 1944). 

Benefits.—The stems of bejuco de costilla are 

used to make baskets. Ground leaves and seeds are 

used to stupefy fish and in preparing herbal 

remedies (Acevedo-Rodriguez 1985). The seeds 

and leaves of this species are poisonous to humans 

and are reported to have been used in arrow 

poisons (Liogier 1990). Aqueous extracts of 

bejuco de costilla from Africa demonstrated 

inhibitory effects against several important 

infectious organisms (Souza and others 1993). The 

white aril associated with the seed is reported to be 

edible (Vélez and Overbeek 1950). 

References 

Acevedo-Rodriguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Morris, K.R.S. 1944. A large-scale experiment in 

the eradication of tsetse (Glossina palpalis and 

G. tachninoides). Farm and Forest 5: 149-156. 

Souza, C. de, K.K. Amegavi, K. Koumaglo, M. 

Gbeassor, and C. De Souza. 1993. Study of the 

antimicrobial activity of the total aqueous 

extracts of ten medicinal plants. Revue de 

Medecines et Pharmacopees Africaines 7: 109-

115. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Río Piedras, PR. 497 p. 
540 

Pavonia spinifex (L.) Cav. cadillo espinoso 

MALVACEAE 

Synonyms: Hibiscus spinifex L. 

 Malache spinifex (L.) Kuntze 

John K. Francis 

General Description.—Cadillo espinoso, also 

known as spiny abutilon, abutilon espinoso, mahot 

jaune, and coquelicot, is a suffruticose shrub 

usually 1 to 1.5 m in height, but occasionally 

reaching 5 m. The plant is supported by a shallow 

root system consisting of a weak taproot, lateral 

roots, and abundant fine roots. Older plants have 

multiple stems arising from the root crown and the 

lower 15 cm of stem. The plant dies back to the 

main stem or sometimes to the root crown during 

the dry season. The branches are slender and 

relatively few in number. The alternate leaves are 

ovate with rounded or cordate bases and pointed 

tips. The leaf blades are 4 to 12 cm long and have 

serrate edges. Bright, 2- to 3-cm yellow flowers 

are usually axillary and solitary and develop into 

fruits that are schizocarps, dividing into five 

mericarps. Each mericarp has three barbed spines, 

the central spine, the largest, being about 1 cm 

long (Howard 1989, Liogier 1994). 

Range.—There is considerable confusion about 

the extent of the native range of cadillo espinoso. 

It is reported to be native from Bermuda through 

the West Indies and from Mexico to Peru and 

Argentina (Griffiths 1994, New York Botanical 

Garden 2002). Howard (1989) states that 

Southeastern United States and the West Indies 

are the only range; other claims are based on 

unsubstantiated reports or misidentifications of 

other species. Finally, Nelson (1996) refers to the 

populations in Florida as escapes from cultivation. 

Ecology.—Cadillo espinoso grows in dry areas. It 

is found in Puerto Rico in habitat that receives 

from 750 to about 1100 mm of precipitation and at 

elevations from near sea level to about 600 m. The 

species grows in well-drained soils with a wide 

variety of textures and fertility levels. Cadillo 

espinoso is more common in areas of igneous 

rocks but does occur over limestone and other 

sedimentary rocks. Cadillo espinoso is moderately 

intolerant of shade. It grows in open areas but 

more frequently under low, open stands of small 

trees. The species competes well with tall grass, 

herbs, and brush, except on the most fertile sites 

where forest stands are dense. Probably because of 

competition, the plant is rarely found in bottoms 

and north slopes. Most of the areas where the 

species is common are recovering from past 

disturbance such as fire, severe overgrazing, or 

clearing. Cadillo espinoso tolerates all but the 

most severe grazing. It sprouts after fires. The 

species occurs as occasional plants or in mixture 

with other species, not as pure or dense stands. 

Reproduction.—Cadillo espinoso flowers during 

the summer (July and August). Fruits mature at 

the start of the dry season (January-February) and 

are dispersed by sticking to the clothing or fur of 

animals. In addition to attaching directly from the 

shrub, fruits rest on the ground so that the central 

spine points up, so they may be picked up by the 

feet of passing animals. Seed production is reliable 

and moderately abundant. Air-dried fruits 

collected in Puerto Rico averaged 0.0258 + 0.0005 

g. Incubated in a plastic bag with moist peat, only 

4 percent germinated in 10 months. Germination is 

epigeal. 

Growth and Management.—Annual growth is 
541

rapid (about 1 m) from sprouts arising from 

established suffruticose plants. Growth begins 

during the early summer rains and continues 

through the fall rainy season. The leaves and 

branches dry out and die during the late winterearly spring dry season (in Puerto Rico). 

Individual plants probably live at least a decade. 

Except for the spiny fruits that cling to clothing, 

there seems to be little need to control cadillo 

espinosa in range or forest land. When necessary, 

the species probably could be controlled by spot 

spraying with broadleaf weed killer to eliminate 

individual plants. The development of a closed 

canopy forest eventually eliminates the shrub. 

Benefits.—Cadillo espinoso is planted to a limited 

extent as an ornamental for its handsome yellow 

flowers. It contributes to the recovery of disturbed 

areas, furnishes cover for wildlife, and is eaten by 

grazing animals. In herbal medicine, infusions of 

the flowers are used to treat sore throats and skin 

problems, and infusions of leaves and twigs are 

used to treat stomach problems, gall stones, and 

liver pain (Liogier 1990).

References 

Griffiths, M. 1994. Index of garden plants. Timber 

Press, Portland, OR. 1,234 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Nelson, G. 1996. The shrub and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

New York Botanical Garden. 2002. NYBG species 

search results. http://s.../wwwspecimen. 

search_list?taxon=Pavonia+spinifex+(L.)+Cav.+

+++&projcode=SEB. 1 p. 
542 

Pennisetum purpureum Schumacher elephant grass 

POACEAE 

Synonyms: none 

John K. Francis 

General Description.—Elephant grass is also 

known as napier grass, marker grass, Uganda 

grass, yerba elefante, capim elefante, herbe à 

elephants, fausse-cane à sucre, and a host of native 

African names (Burkill 1994, Holm and others 

1977). The species is a robust grass with 

perennial stems. The plants produce short, 

creeping rhizomes 15 to 25 cm long with fine 

roots at the nodes and culms that are from 2 to 8 m 

in height, up to 2.5 cm in diameter at the base, and 

have a solid center. Older culms may branch 

several times. Leaf blades are 50 to 90 cm long 

and 1 to 3 cm wide, flat, and have a white midrib. 

Leaves of new, vigorous growth have wide, robust 

leaves; older culms have finer, narrow leaves. 

Leaf margins are rough (fine-toothed). The 

inflorescence is a compact, erect, bristly tawny or 

purplish spike 8 to 30 cm long and 1.5 to 3 cm 

wide. Spikelets are arranged around a hairy axis, 

and fall at maturity. The chromosome number is 

2n = 27, 28, or 56 (Burkill 1994, Holm and others 

1977, Long and Lakela 1971, Skerman and 

Riveros 1990). 

Range.—Elephant grass is native throughout 

humid, tropical mainland Africa and the island of 

Bioko (Burkill 1994). It has been planted for 

forage and has naturalized in many tropical areas 

in Asia, the America’s, and Oceania. It grows wild 

in the U.S. territories of Florida, Texas, California, 

Hawaii, Guam, American Samoa, Puerto Rico, 

and U.S. Virgin Islands (Natural Resource 

Conservation Service 2002, Pacific Island 

Ecosystems at Risk 2002). 

Ecology.—Elephant grass will grow on poorly 

drained clay soils through the gamut of soil types 

to excessively drained sandy soils. Growth is best 

on rich, moist, well-drained medium-textured 

soils. Soil reaction should range from pH 4.5 to 

8.2 (Center for New Crops and Plant Products 

2002). Rainfall should be in excess of 1500 mm 

per year and temperatures for optimum growth 

should be from 25 to 40 °C (Skerman and Riveros 

1990). In its native range, the species is a fire subclimax to broadleaf tropical forest (Center for 

New Crops and Plant Products 2002). Elephant 

grass will grow in light shade but is does not 

survive under a closed tree canopy. In turn, it will 

suppress most grasses, herbs, and tree seedlings. A 

frost will kill the above-ground parts, but the soil 

must be frozen to kill the rhizomes (Center for 

New Crops and Plant Products 2002). 

Reproduction.—Flowering takes place mainly in 

the fall and winter (Long and Lakela 1971, Holm 

and others 1977). Because of asynchrony of male 

and female flower parts, the plant relies on crosspollination by wind. Elephant grass is an 

inconsistent seed producer and rarely develops 

seeds in some habitats. When seeds are produced, 

they are often of low viability (Holm and others 

1977). About 3.8 million seeds per kg were 

reported for a United States source (Skerman and 

Riveros 1990). A collection of seeds from Puerto 

Rico numbered 1.78 million per kg (author’s 

observation). The seeds are wind-dispersed. 
543

Colonization of new habitats is slow (Skerman 

and Riveros 1990). 

Growth and Management.—The growth of 

culms is vertical continually, but their weight 

bends them in the middle and lower part, causing a 

j-shaped habit. As the lower stem makes contact 

with moist soil, it roots at every node. Individual 

culms live more than 1 year, and by layering, they 

can continue until disturbed or a barrier is reached. 

New plantations are established by planting stem 

pieces with at least three nodes or with root 

cuttings (Skerman and Riveros 1990). In an 

agricultural research plot in Puerto Rico, elephant 

grass reached the height of 4 m in 3 months 

(Barrett 1925). The record yield for heavily 

fertilized elephant grass is 84,800 kg/ha/yr 

(Skerman and Riveros 1990). Not-withstanding its 

value as forage, elephant grass has become one of 

the worst weeds in the tropics because of the 

difficulty of controlling it in croplands and fallow 

areas. Cultivation alone is usually insufficient to 

control it in croplands (Skerman and Riveros 

1990). Frequent mowing will cause it to be 

replaced by other grasses. The herbicide 

glyphosate provides acceptable control in at least 

aquatic sites (McCann and others 1996). 

Benefits.—Elephant grass makes up the bulk of 

the diet of forest elephants in West Cameroon 

(Tchamba and Seme 1993). The species is an 

important forage and pasture grass in its native 

Africa and throughout the Tropics, especially for 

cattle. It is also cut for hay and fermented for 

silage. A number of forage samples of different 

ages of grass from several countries varied from 4 

to 15 percent in crude protein, from 28 to 40 

percent in crude fiber, from 10 to 16 percent in 

ash, from 0.9 to 3.8 percent in fat, and from 39 to 

49 percent in nitrogen-free extract (Skerman and 

Riveros 1990). The species has been employed 

successfully using sown cuttings to replace 

Imperata swards in the Phillippines (Agus and 

others 1996, Skerman and Riveros 1990). It is 

planted as hedgerows for erosion protection and 

forage production in the alley cropping system of 

agroforestry (Magcale-Macandog and others 1998, 

Menz and others 1999). The plant is also effective 

as a windbreak for agricultural crops (Karschon 

and Heth 1958). Lines of plants are used to mark 

boundaries between plots and properties. Elephant 

grass is planted on riverbanks to prevent erosion. 

In Africa, the plant is used for thatch, and the thick 

culms are made into fences, screens, and 

reinforcement for mud huts. The young leaves and 

young shoots are eaten in soups and stews (Burkill 

1994). Elephant grass is used for mulch in East 

Africa where a 25-cm depth of mulch is needed 

for good weed control (Nishimoto 1994). Extracts 

of the plant are strongly diuretic and are used for 

that purpose in Africa. It is also used in a number 

of other herbal remedies (Burkill 1994). The seeds 

are eaten by many bird species. However, because 

of the aggressive spread of the species, it is a 

menace to native vegetation in the Galapagos 

Islands (Mauchamp 1997) and at the margins of 

swamps and streams in Florida (Miami-Dade 

County 2002). 

References 

Agus, F., D.K. Cassel, and D.P. Garrity. 1996. 

Soil-water and soil physical properties under 

contour hedgerow systems on sloping Oxisols. 

Soil and Tillage Research 40(3-4): 185-199. 

Barrett, O.W. 1925. The food plants of Puerto 

Rico. Journal of the Department of Agriculture 

of Puerto Rico 9(2): 193. 

Burkill, H.M. 1994. The useful plants of West 

Tropical Africa. Royal Botanic Gardens. Kew, 

UK. 636 p. 

Center for New Crops and Plant Products. 2002. 

Pennisetum purpureum K. Schumach. Perdue 

University. http://hort.purdue.edu/newcrop/ 

duke_energy/Pennisetum_purpurium.html. 4 p. 

Holm, L.G., D.L. Plucknett, J.V. Pancho, and J.P. 

Herberger. 1977. The World’s worst weeds. 

East-West Center. Honolulu, HI. 609 p. 

Karschon, R. and D. Heth. 1958. Wind speed, 

wind-borne salt and agricultural crops as 

affected by windbreaks. La Yaaran 8(3/4): 8-13, 

32-38. 

Long, R.W. and O. Lakela. 1971. A flora of 

Tropical Florida. University of Miami Press, 

Coral Gables, FL. 962 p. 

Magcale-Macandog, D.B., C.D. Predo, K.M. 

Menz, and A.D. Calub. 1998. Napier grass strips 

and livestock: a bioeconomic analysis. 

Agroforestry Systems 40(1): 41-58. 

Mauchamp, A. 1997. Threats from alien plant 

species in the Galapagos Islands. Conservation 

Biology 11(1): 260-263. 
544 

McCann, J.A., L.N. Arkin, and J.D. Williams. 

1996. Noniddigenous aquatic and semi-aquatic 

plants in freshwater systems. University of 

Florida, Center for Aquatic Plants. 

http://aquat1.ifas.ufl.edu/mctitle.html. [Not 

paged]. 

Menz, K.M., D. Magcale-Macandog, and I.W. 

Rusastra, eds. 1999. Improving smallholder 

farming systems in Imperata areas of Southeast 

Asia: alternatives to shifting cultivation. 

Australian Centre for International Agricultural 

Research. Canberra, Australia. 280 p. 

Miami-Dade County. 2002. Napier grassPennisetum purpureum. http://co.miamidade.fl.us/derm/environment/badplants/plant%2

0desc.../napier_grass.ht. 1 p. 

Natural Resources Conservation Service. 2002. 

Plants profile: Pennisetum purpureum

Schumacher. Washington, DC. http://plants. 

usda.gov/cgi_bin/plant_profile.cgi?symbol=PEP

U2. 3 p. 

Nishimoto, R.K. 1994. Weed control in coffee 

plantations. In: R. Labrada, J.C. Caseley, and C. 

Parker, eds. Weed management for developing 

countries. FAO Plant Production and Protection 

Paper 120. Food and Agriculture Organization 

of the United Nations. Rome. p. 354-359. 

Pacific Island Ecosystems at Risk. 2002. Invasive 

plant species: Pennisetum purpureum

Schumacher, Poaceae. http://www.hear.org/ 

pier3/pepur.htm. 2 p. 

Skerman, P.J. and F. Riveros. 1990. Tropical 

grasses. FAO Plant Production and Protection 

Series 23. Food and Agriculture Organization of 

the United Nations. Rome. 832 p. 

Tchamba, M.N. and P.M. Seme. 1993. Diet and 

feeding behavior of the forest elephant in the 

Santchou Reserve, Cameroon. African Journal 

of Ecology 31(2): 165-171. 
545

Penstemon ambiguus Torr. sand penstemon 

SCROPHULARIACEAE 

Synonym: Leiostemon ambiguus (Torr.) Greene 

James E. Nellessen 

General Description.—Sand penstemon is also 

known by the names bush penstemon, prairie or 

pink plains penstemon, moth penstemon, gilia 

penstemon, phlox penstemon, and cow-tobacco. 

This species occurs in sandy soils of plains, and 

mesas, and somewhat hilly terrain. The plant 

generally ranges from 20 to 60 cm in height, 

sometimes to 1 m, is highly branched, the stems 

are slightly hairy, and become distinctly woody 

well above the bases. Leaves are simple, linear, 

opposite, 6 to 30 mm long, enrolled, smooth to 

slightly hairy, with slightly rough edges (Carter 

1997, Epple 1995, Great Plains Flora Association 

1986, Kearney and others 1951, Martin and 

Hutchins 1981). There are two recognized 

varieties: P. ambiguus var. ambiguus and P. 

ambiguus var. laevissimus (Keck) N. Holmgren 

(Kartesz 1994, Allred 2002). Both varieties have 

also been classified as subspecies. Variety 

laevissimus differs from the typical in that it has 

smooth stems, with the leaf edges smooth or very 

slightly rough. The chromosome number is 

reported as n = 8, which is common to many 

members of this genus (Nisbet and Jackson 1960). 

Range.—This species occurs from eastern 

Colorado, western Kansas, Oklahoma, and Texas, 

through eastern and central New Mexico, then 

west into Arizona, Utah, and Nevada, at elevations 

from about 850 m to 2,000 m. Variety ambiguus

occurs in the northern and eastern portions of this 

range while variety laevissimus occurs in the 

southern and western portions of this range. 

Ecology.—Sand penstemon occurs on alluvial 

plains, valleys, and in sandy hill country. It is 

considered one of the major shrub species 

comprising Plains-Mesa Sand Scrub vegetation 

that occurs along drainages of the Rio Grande, 

Pecos River, San Juan River, and areas within the 

Llano Estacado ("staked" plains) of western Texas 

and eastern New Mexico (Dick-Peddie 1993). It is 

common in the Painted Desert of Arizona 

(Kearney and others 1951). A couple of hybrid 

swarms between P. ambiguus and P. thurberi Torr. 

(P. ambiguus x thurberi Nisbet and Jackson) are 

known from: 1) an area near Magdalena, Socorro 

County, and 2) another near Carrizozo, Lincoln 

County, both in New Mexico (Nisbet and Jackson 

1960). P. thurberi has a similar shrubby habit and 

occurs in sandy soils. Hybrids have shown a wide 

range of characteristics between the two species. 

Although generally considered a separate species, 

P. thurberi has also been classified as a variety of 

P. ambiguus (P. ambiguus Torr. var. thurberi

Gray). 

Biochemical Ecology.—Several iridoid glucosides 

have been isolated and identified from sand 

penstemon: catalpol, speciocide, nemoroside, 

ambiguuside; as well as two phenylpropanoid 

glycosides: verbascoside and martynoside 

(Arslanian and others 1990). Iridoids are a group 

of bitter tasting terpenoid toxins also referred to as 

monoterpene lactones (Harborne 1988). These 

compounds may be toxic to birds and insects, 

hence functioning as a chemical defense against 

herbivory. 

Reproduction.—Each plant has numerous manyflowered inflorescences, and each inflorescence is 

narrow, with one to two flowers per peduncle, the 

flowers spreading. The calyx is 2 to 3 mm long, 

the corolla 15 to 25 mm long, pale to deep pink, 

the five corolla lobes whitish, the upper ones 

reflexed at an oblique angle to the tube, the lower 

one projecting forward, making it distinctive from 

most other penstemons. The opening of the tube is 

bearded with short hairs, in two prominent 

guidelines at the base of the throat (Nisbet and 

Jackson 1960, Martin and Hutchins 1981, Carter 
546 

1997). The sterile stamen (staminode) is beardless 

(hairless), a feature different from most 

penstemons (beardtongues), hence the alternate 

genus name Leiostemon, leio meaning smooth and 

stemon for stamen (Kirkpatrick 1992, Weber and 

Wittman 2001). It blooms from May to as late as 

October, depending on local environmental 

conditions, although mass blooming generally 

occurs from June through July. The hybrid, P. 

ambiguus x thurberi has smaller corollas, 12 to 15 

mm long, corolla lobes white to blue-purple, the 

throat and tube pale pink to deep reddish purple. 

The fruit is a capsule from 5 to 10 mm long that 

contains many very small (1.2 to 2 mm) seeds. 

Seed germination characteristics (for example, the 

amount of cold stratification) vary as a function of 

the elevation of the source population (Meyer and 

others 1995). Sand penstemon seeds have a 

physiological dormancy requiring 8 to 15 weeks of 

cold stratification before germination will occur 

(Kitchen and Meyer 1991, Baskin and Baskin 

2001). 

Growth and Management.—Seedlings take a 

few years becoming established before this longlived plant becomes larger and more conspicuous 

in the landscape. Only a few flowers may be 

produced the first year of blooming, before the 

plant gains enough mass and root reserves for the 

large floral displays that make this plant so 

characteristic. 

Benefits.—Since this species’ typical habitat is in 

sandy, unstable type soils, it acts as a soil 

stabilizer. Because of its conspicuous and colorful 

floral displays, it is a welcome addition to habitats 

that might otherwise appear drab. This is an 

attractive plant for native plant landscaping 

(Morrow 1995, Warnock 1974), and in full bloom 

with the numerous stems and flowers per stem, it 

makes for an attractive display of pink bouquets. 

References

Allred, K.W. 2002. A working index of New 

Mexico vascular plant names. Available on a 

New Mexico State University web page. 

http://web.nmsu.edu/~kallred/herbweb. [not 

paged]. 

Arslanian, R.L., T. Anderson, and F.R. Stermitz. 

1990. Iridoid glucosides of Penstemon 

ambiguus. Journal of Natural Products--Lloydia 

53(6): 1,485-1,489. 

Baskin, C.C. and J.M. Baskin. 2001. Seeds: 

ecology, biogeography, and evolution of 

dormancy, and germination. Academic Press, 

San Diego, CA. 666 p. 

Carter, J.L. 1997. Trees and shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation: Past, Present and Future. University 

of New Mexico Press, Albuquerque, NM. 244 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. Falcon Publishing Inc., Helena, MT. 

347 p. 

Great Plains Flora Association. 1986. Flora of the 

Great Plains. University Press of Kansas, KS. 

1,392 p. 

Harborne, J. B. 1988. Introduction to ecological 

biochemistry, 3rd Ed. Academic Press, London. 

356 p. 

Haflin, J. 1997. Penstemons: the beautiful 

breadtongues of New Mexico. Jackrabbit Press, 

Albuquerque, NM. 50 p. 

Kartesz, J.T. 1994. A synonymized checklist of 

the vascular flora of the United States, Canada, 

and Greenland. Vol. 1, 2nd Edition. Biota of 

North America Program of the North Carolina 

Botanical Garden. Timber Press, Portland, OR. 

622 p. 

Kearney, T.H., R. Peebles, and Collaborators. 

1951 with 1960 supplement. Arizona flora. 

University of California Press, Berkeley. 

1,085 p. 

Kirkpatrick, Z.M. 1992. Wildflowers of the 

Western Plains. University of Texas Press. 

Austin, TX. 240 p. 

Kitchen, S.G. and S.E. Meyer. 1991. Seed 

germination of intermountain Penstemons as 

influenced by stratification and GA3 treatments. 

Journal of Environmental Horticulture 9: 51-56. 

Martin, W.C. and C.E. Hutchins. 1981. (reprinted 

2001). A flora of New Mexico. Vol. 2. Bishen 

Singh Mahendra Pal Singh, India and Koeltz 

Scientific Books, Germany) p. 1,277-2,591. 
547

Meyer, S.E., S.G. Kitchen, and S.L. Carlson. 1995. 

Seed germination timing patterns in 

intermountain Penstemon (Scrophulariaceae). 

American Journal of Botany 82: 377-389. 

Morrow, B.H. 1995. Best plants for New Mexico 

gardens and landscapes. University of New 

Mexico Press, Albuquerque, NM. 267 p. 

 

Nisbet, G.T. and R.C. Jackson. 1960. The genus 

Penstemon in New Mexico. The University of 

Kansas Science Bulletin 41(5): 691-759. 

Warnock, B.H. 1974. Wildflowers of the 

Guadalupe Mountains and the Sand Dune 

Country, Texas. Sul Ross State University, 

Alpine, TX. 176 p. 

Weber, W.A. and R.C. Wittman. 2001. Colorado 

flora: Eastern Slope. University Press of 

Colorado, Boulder CO. 521 p. 
548 

Philadelphus lewisii Pursh Lewis mock orange 

HYDRANGEACEAE 

 

Synonyms: Philadelphus gordonianus Lindl. 

Philadelphus cordatus Petz. & Kirchn. 

Philadelphus grahami Petz. & Kirchn. 

Philadelphus columbianus Koehne 

 

Nancy L. Shaw

Illustration credits: E.G. Hurd, and Hitchcock and others 

1961 

General Description.—Lewis mock orange, also 

known as Indian arrowwood, Lewis syringa, mock 

orange, syringa, or western syringa, was named for 

Captain Meriwether Lewis, the famous Western 

explorer, who made the first collection of the 

species in 1806. The state flower of Idaho, Lewis 

mock orange is a long-lived and highly variable 

deciduous species, ranging from densely branched 

and rounded to erect, open shrubs 1.5 to 2.5 m in 

height. Clusters of arching stems develop on older 

specimens. Bark on young branches is red to 

chestnut brown, turning gray and exfoliating with 

age. Leaves are 2 to 8 cm long and 1 to 4 cm wide, 

opposite, and ovate to oblong with acute to 

acuminate tips. Terminal cymes of showy, perfect, 

four-merous, fragrant white flowers develop on 

lateral branches. The fruit is a woody, four-celled 

capsule containing numerous tiny, fusiform seeds 

(Hitchcock and others 1961, Welsh and others 

1987). 

Range.—Lewis mock orange is distributed from 

British Columbia south to northern California and 

east to southwestern Alberta and western Montana. 

It is most common at mid-elevations but grows 

from near sea level to 2,100 m (USDA Forest 

Service 1937). Lewis mock orange exhibits wide 

ecological amplitude, growing in communities 

ranging from Pseudotsuga menziesii (Mirbel) 

Franco and Sequoia sempervirens (Lamb. ex D. 

Don) Endl. to Artemisia L., chaparral, and Pinus 

contorta Dougl. ex Loud and Pinus ponderosa P. 

& C. Lawson. 

Ecology.—Lewis mock orange grows on soils 

ranging from dry, rocky, gravelly loams on open 

hillsides, to well-drained deep, rich alluvial loams 

near riparian zones (USDA Forest Service 1937). 

It is found in early to late seral as well as in climax 

communities. It occurs as small thickets, isolated 

plants, or in association with other shrubs on talus 

slopes, cliffs, canyons, or rocky hillsides and in 

transition zones of riparian areas (Hitchcock and 

others 1961, Hopkins and Kovalchik 1983). It is 

most common on northern and eastern exposures 

(USDA Forest Service 1937). The species is 

classified as fire resistant because it resprouts from 

the root crown following burning. It also 

regenerates from seeds that accumulate in the soil 

seed bank (Kramer and Johnson 1987). Lewis 

mock orange exhibits moderate drought and shade 

tolerance. 

Reproduction.—Lewis mock orange flowers in 

late spring to early summer. Capsules mature in 

late summer and dehisce in September or October. 

Seeds are dispersed by wind and gravity. They are 

harvested by hand-stripping the capsules before 

the valves begin to open. Dried capsules are 

crushed to release the seeds, and trash is removed 

with an aspirator or fanning mill. There are 

7,716,000 to 17,637,000 seeds/kg of clean seed 

(Stickney and others 2001), but seed fill is often 

low. Seed may be stored in airtight containers for 

up to 1 year. Wet prechilling for 8 weeks at 5 °C is 

required to release embryo dormancy (Stickney 

and others 2001). Intermittent exposure to light 

may be required for germination. Seeds may be 
549

broadcast seeded on a rough seedbed and covered 

using a Brillion seeder or similar device. They 

may also be spot seeded in selected, prepared areas 

that are well-drained and free of herbaceous 

competition. Seeds may be mixed with other shrub 

seeds that require shallow or surface planting. 

Bareroot stock may be produced by fall seeding or 

by seeding moist, prechilled seeds in spring. 

Seedlings develop rapidly and can be transplanted 

as 1-year-old stock (Stickney and others 2001). 

Container stock is grown from seed or propagated 

from hardwood or softwood cuttings (Marchant 

and Sherlock 1984). Rooted suckers and crown 

divisions are also used. 

Growth and Management.—Lewis mock orange 

seedlings establish well on a wide variety of soils 

but should be protected from competition and 

browsing. Plants grow at a moderate rate, flower 

reliably, and are generally free of insect and 

disease problems. Seedlings, however, are 

sometimes susceptible to damping off. Plants 

grown from seed may begin flowering in the 

second or third year (Everett 1957). Flowers are 

produced on twigs of the previous year; hence, 

landscape plants should be pruned after flowering. 

Benefits.—Seedlings or larger planting stock of 

Lewis mock orange are transplanted into steep, 

rocky, unstable slopes where they provide soil 

stabilization and vegetative cover. The species is 

also useful in transitional areas of degraded 

riparian zones. Lewis mock orange is usually not 

grazed heavily by livestock, but in some areas it 

does receive fair amounts of use by cattle (Bos

spp. L.) and sheep (Ovis spp. L.) (Leege 1968, 

USDA Forest Service 1937). It frequently occurs 

with other species that are more palatable to big 

game, and consequently, it may receive little use, 

except under severe conditions. However, it can 

provide good browse for deer (Odocoileus spp. 

Rafinesque) and elk (Cervus canadensis

Erxleben), especially on winter ranges (Kufeld 

1973, Leege 1968, Marchant and Sherlock 1984, 

USDA Forest Service 1937). New growth is 

generally highly palatable to big game (Leege 

1968). The species provides food and cover for 

birds and other small animals. Lewis mock orange 

is a valued landscape species because of its showy, 

white flowers. It is used in borders, screens, 

hedges, and as specimen plants (Marchant and 

Sherlock 1984). Several commercial cultivars of 

Lewis mock orange have been developed. The 

flowers are used in preparing perfumes and teas 

(Taylor 1972). Native Americans made arrow 

shafts from the long woody shoots (USDA Forest 

Service 1937). 

References 

Everett, P.C. 1957. A summary of the culture of 

California plants at the Rancho Santa Ana 

Botanic Garden. The Rancho Santa Ana 

Botanic Garden, Claremont, CA. 223 p. 

Hitchcock, C.L., A. Cronquist, M. Ownbey, and 

J.W. Thompson. 1961. Vascular plants of the 

Pacific Northwest. Part 3: Saxifragaceae to 

Ericaceae. University of Washington Press, 

Seattle, WA. 614 p. 

Hopkins, W.E. and B.L. Kovalchik. 1983. Plant 

associations of the Crooked River National 

Grassland. R6 Ecol. 133-1983. U.S. 

Department of Agriculture, Forest Service, 

Pacific Northwest Region, Portland, OR. 98 p. 

Kramer, N.B. and F.D. Johnson. 1987. Mature 

forest seed banks of three habitat types in 

central Idaho. Canadian Journal of Botany 65: 

1961-1966. 

Kufeld, R.C. 1973. Foods eaten by the Rocky 

Mountain elk. Journal of Range Management 

26: 106-113. 

Leege, T.A. 1968. Prescribed burning for elk in 

northern Idaho. Proceedings: Annual tall 

timbers fire ecology conference. 8: 235-253. 

Marchant, C. and J. Sherlock. 1984. A guide to 

selection and propagation of some native 

woody species for land rehabilitation in British 

Columbia. Forest Research Report RR84007-

HQ. Ministry of Forests, Victoria, BC. 117 p. 

Stickney, P.F., N.L. Shaw, and E.G. Hurd. 2001. 

Philadelphus L. Mockorange. In: F.T. Bonner 

and R.G. Nisley, eds. Woody plant seed 

manual. Agriculture Handbook. U.S. 

Department of Agriculture, Forest Service, 

Washington, D.C. http://wpsm.net/index.html. 

5 p. 

Taylor, S. 1972. Philadelphus lewisii Pursh ‘mock 

orange’ or ‘syringa’. Davidsonia 3: 4-7. 

USDA Forest Service. 1937. Range plant 

handbook. U.S. Government Printing Office, 

Washington, D.C. 512 p. 
550 

Welsh, S.L., N.D. Atwood, L.C. Higgins, and S. 

Goodrich. 1987. A Utah flora. Great Basin 

Naturalist 9. Brigham Young University, Provo, 

UT. 894 p. 
551

Philadelphus microphyllus Gray littleleaf mock orange 

HYDRANGEACEAE 

Synonyms: Philadelphus occidentalis A. Nels. 

 Philadelphus argenteus Rydb.

 Philadelphus minutus Rydb. 

 Philadelphus nitidis A. Nels. 

 Philadelphus stramineus Rydb. 

Nancy L. Shaw 

 

Illustration source: Cronquist and others 1997 

General Description.—Littleleaf mock orange, 

also known as desert mock orange, is a small, 

rounded, multistemmed shrub 0.9 to 2.1 m tall 

with a crown diameter of about 1.2 m. New woody 

growth is strigose, while bark of mature stems is 

gray to yellowish and exfoliating. Some stems may 

be subspinose. Leaves are opposite, entire, short 

petiolate, 8 to 25 mm long, 2 to 13 mm wide, 

elliptic to lanceolate or linear, and sometimes 

revolute. Flowers are solitary or in clusters of three 

at the ends of branches produced the previous 

year. They are showy, white or creamy, fragrant, 

and four-merous with numerous stamens. Fruits 

are four-chambered loculicidal capsules containing 

numerous black, short-caudate seeds. The 

chromosome number is 2n = 26 (Cronquist and 

others 1997, Hickman 1993, Munz and Keck 1973, 

Welsh and others 1987). 

Range.—Littleleaf mock orange is distributed 

from southern California east into southern and 

central Nevada; most of Utah; southwestern 

Wyoming; western, central, and southern 

California; Arizona; New Mexico; central Texas; 

and central Mexico (Cronquist and others 1997). 

The species exhibits considerable morphological 

variability and has been subdivided into a number 

of species or subspecific taxa by various authors. 

Ecology.—Littleleaf mock orange grows in Pinus

spp. L.-Juniperus spp. L., Quercus spp. L., 

mountain brush, Populus tremuloides Michx., 

Pinus contorta Dougl. ex Loud., Pseudotsuga

menziesii (Mirb.) Franco, and Abies concolor

Lindl. communities at elevations from 1,200 to 

3,100 m (Cronquist and others 1997, Hitchcock 

1943, Welsh and others 1987). A drought-tolerant 

species, it is often found growing in cracks or 

fissures in rocky surfaces and in other dry, sandy, 

gravelly, or rocky areas. 

Reproduction.—Plants flower in May to 

September depending upon location. The capsules 

mature in late summer. Mature capsules are 

harvested by hand just after they have begun to 

dehisce. Dried capsules are crushed with a barley 

debearder to permit seed extraction. Seed is 

separated from debris using an aspirator or airscreen machine. Littleleaf mock orange is readily 

propagated from seed (Sutton and Johnson 1974, 

Swingle 1939). Germination of a New Mexico 

seed collection was 12 times greater when 

incubated at 15 compared to 20/10 o

C (8 hrs/16 

hrs) for 28 days (Stickney and others 2001). 

Germination was improved by a 28-day moist 
552 

prechill at 3 to 5 oC when seeds were subsequently 

incubated at 20/10 oC, but not at 15 oC. 

Germination is epigeal. 

Growth and Management.—Little data are 

available on the propagation of littleleaf mock 

orange, but species of the genus Philadelphus are 

generally easily propagated from softwood or 

hardwood cuttings, rooted suckers, divisions, or 

layers (Hartmann and others 1990, Macdonald 

1986). Littleleaf mock orange is easily 

transplanted (Sutton and Johnson 1974). Seeds of 

Philadelphus spp. are tiny and may be broadcast 

seeded on a rough seedbed and covered lightly or 

spot seeded on prepared seedbeds (Stevens and 

others, in press). Seeds may also be surfaceplanted using a Brillion or similar seeder. Best 

results are obtained if seeds are planted in welldrained sites free of herbaceous competition. 

Seeds may be mixed with other shrub seeds that 

require surface or shallow planting. Growth of 

littleleaf mock orange is moderate to rapid (Sutton 

and Johnson 1974). 

Benefits.—Littleleaf mock orange provides good 

cover for small animals. It is used to some extent 

by mule deer (Odocoileus hemionus Rafinesque) 

(Patton and Ertl 1982). Although the species 

receives little use in revegetation, it has potential 

for planting on disturbances on steep, rocky, 

unstable slopes within its native range (Stevens 

and others, in press). Seedlings or larger stock are 

recommended for such use. The species might 

also be used to advantage in drier areas of 

degraded riparian zones. Littleleaf mock orange is 

an attractive ornamental because of its showy 

flowers and fall coloration. It was first cultivated 

in 1883 (Rehder 1940). It can be used in borders, 

screens, hedges, or as isolated specimens in sunny 

areas. It can also be used for low maintenance 

landscaping and in recreational area plantings. The 

small seeds were eaten by Native Americans 

(Cronquist and others 1997). 

References 

Cronquist, A., N.H. Holmgren, and P.K. 

Holmgren. 1997. Intermountain flora. Volume 3, 

Part A. New York Botanical Garden, Bronx, 

NY. 446 p. 

Hartmann, H.T., D.E. Kester, and F.T. Davies, Jr. 

1990. Plant propagation principles and practices. 

5th Ed. Prentice-Hall, Inc., Englewood Cliffs, 

NJ. 657 p. 

Hickman, J.C. 1993. The Jepson manual: higher 

plants of California. University of California 

Press, Berkeley, CA. 1,400 p. 

Hitchcock, C.L. 1943. The xerophyllous species of 

Philadelphus in southwestern North America. 

Madroño 17: 35-56. 

Macdonald, B. 1986. Practical woody plant 

propagation for nursery growers. Vol. I. Timber 

Press, Portland, OR. 669 p. 

Munz, P.A. and D.D. Keck. 1973. A California 

flora and supplement. University of California 

Press, Berkeley, CA. 1,681 p. 

Patton, D.R. and M.G. Ertl. 1982. Run wild 

wildlife/habitat relationships. Wildlife Unit 

Technical Series. U.S. Government Printing 

Office, Washington, D.C. 49 p. 

 

Rehder, A. 1940. Manual of cultivated trees and 

shrubs. 2nd Ed. The MacMillan Co., New York. 

996 p. 

Stevens, R., S.B. Monsen, and N.L. Shaw. [in 

press]. Shrubs of other families. In: S.B. Monsen 

and R. Stevens, comps. Restoring western 

ranges and wildlands. INT-GTR. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Ogden, UT. 

Stickney, P.F., N.L. Shaw, and E.G. Hurd. 2001. 

Philadelphus L. Mockorange. In: F.T. Bonner 

and R.G. Nisley, eds. Woody plant seed manual. 

U.S. Department of Agriculture, Forest Service, 

Agriculture Handbook. Washington, D.C. 

http://wpsm.net/ index.html. 5 p. 

Sutton, R. and C.W. Johnson. 1974. Landscape 

plants from Utah’s mountains. EC-368. Utah 

State University, Logan, UT. 137 p. 

Swingle, C.F. 1939. Seed propagation of trees, 

shrubs and forbs for conservation planting. SCSTP-27. Washington, DC: U.S. Department of 

Agriculture, Soil Conservation Service. 198 p. 

Welsh, S.L., N.D. Atwood, L.C. Higgins, and S. 

Goodrich. 1987. A Utah flora. Great Basin 

Naturalist 9. Brigham Young University, Provo, 

UT. 894 p. 
553

Phoradendron quadrangulare (Kunth) Griseb. guacimilla de canario 

VISCACEAE (sometimes placed in LORANTHACEAE) 

Synonyms: Phoradendron randiae (Bello) Britt. 

 Loranthus quadrangularis Kunth in Humb. 

 Viscum quadrangulare (Kunth) DC. 

 Phoradendron rubrum sensu Griseb. 

 Viscum trigonum D. Dietr. 

John K. Francis 

General Description.—Guacimilla de canario is 

an evergreen, epiparasitic shrub that obtains water, 

minerals, and support from its tree hosts and 

produces stems up to 1 m long and 1 cm thick. 

The woody stems are much branched, brown, and 

lightly furrowed in their lower parts with a 

swelling at the point of attachment to the host. The 

bark of branches and twigs is green. The young 

branches especially are square or rhombic in cross 

section. The stem wood is light-colored and 

brittle. The dull green leaves have a leatheryfleshy texture. They are linear, lanciolate, or 

oblanciolate with a tapered base and acute to 

obtuse apex. Inflorescences are small spikes, one 

to three in number at the leaf axils, and support 

small flowers. The yellow, globose fruits are 3 to 

4 mm in diameter (Howard 1988, Liogier 1985). 

However, fruits in Costa Rica range from pale 

yellow to red-orange (Missouri Botanical Garden 

2002). Within the pigmented skin is a layer of 

sticky jelly and a flat, green seed with longitudinal 

striations (author’s observations). 

Range.—Guacimilla de canario is native to 

Central America, South America, and the West 

Indies (Howard 1988, Liogier 1985).

Ecology.—Guacimilla de canario colonizes a 

number of tree species, especially Guazuma 

ulmifolia Lam. in Puerto Rico (author’s 

observation). The former species, as well as 

Amphitecna latifolia (Mill.) A.H. Gentry, Cordia 

stellifera I.M. Johst., Erythrina sp., Malvaviscus 

arboreus Cav., and Persea Americana Miller, are 

mentioned as hosts in a Costa Rican herbarium 

collection (Missouri Botanical Garden 2002). 

Most of the guacimilla de canario plants occur in 

the mid-crown of the affected trees. After the 

parasite becomes general throughout a tree crown 

(a process that takes many years), trees are 

weakened and sometimes die as a result. More 

often, parasitized trees die of other causes or old 

age before they succumb to the parasite. Trees in 

forest, rangeland, farmland, roadside, and cities 

are attacked. Guacimilla de canario grows at lower 

and middle elevations in Puerto Rico (Liogier 

1985), that is, from sea level to about 600 m. 

These areas range from 750 to about 2400 mm of 

annual precipitation. In Ecuador, the plant may be 

found from near sea level to 1,000 m in elevation 

(Jorgensen and León-Yánez 1999). 

Reproduction.—Fruits collected in Puerto Rico 

weighed an average of 0.0294 + 0.0004 g/fruit. 

Seeds cleaned from those fruits weighed (airdried) an average of 0.0017 + 0.0000 g/seed or 

600,000 seeds/kg. Sown without any pretreatment 

on moist filter paper, 59 percent germinated within 

35 days of sowing. After 4 days, mesoscopic roots 

emerged followed at 17 days by a shoot that 

extended hypogenally (author’s observation). 

Birds transport the seeds (the fruits are sticky and 

adhere to their feet or beaks) to new trees or 

stands. After fruits stuck to the bark of a suitable 

host tree, they require a rainy period long enough 

to allow the roots of the germinating seed to 

penetrate the phloem of the host, or the seed or 

plantlet will desiccate. Once established in a new 

tree, the guacimilla de canario spreads by seeds 
554 

within the crown of the host tree. It does not 

spread vegetatively. 

Growth and Management.—The growth rate of 

guacimilla de canario is not known. Although it is 

a parasite and does harm its hosts, the plant takes 

many years to become sufficiently abundant to 

seriously weaken a host tree. Guazuma ulmifolia

and most of the other hosts are not used as 

ornamentals and are not commercially valuable. If 

control is desired, probably the best method is to 

clip the guacimilla de canario plants from the 

limbs each year or until they die. This will prevent 

it from bearing fruit and spreading within the tree. 

Alternately, one may prune out the infected limbs 

of the parasitized tree. 

Benefits and Detriments.—Guacimilla de 

canario contributes in a minor way to biodiversity, 

to tree canopy density, and to biomass production. 

It furnishes fruits for wildlife food. On the other 

hand, a few trees/km2

 are attacked, some are 

weakened, and an occasional tree dies as a result. 

Parasitized trees are often unsightly. 

References 

Howard, R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Jorgensen, P.M., and S. León-Yánez. 1999. 

Catalogue of the vascular plants of Ecuador. 

Missouri Botanical Garden, St. Louis, MO. 

1,181 p. 

Liogier, H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Missouri Botanical Garden. 2002. Manual de la 

flora de Costa Rica. http://www.mobot. 

org/manual.plantas/040567/S040679.html. 4 p. 
555

Phragmites australis (Cav.) Trin. ex Steud. common reed 

POACEAE 

Synonyms: Phragmites communis Trin. 

 Phragmites phragmites (L.) Karst. 

John K. Francis 

Drawing source: Britton and Brown 1913 

General Description.—Common reed is also 

known as giant reed, giant reedgrass, roseau, 

roseau cane, yellow cane, and cane (Uchytil 1992). 

It is a shrub or perennial woody graminoid herb 

that grows annually from woody rhizomes to 

heights of 2 to 4 m (occasionally to 6 m). The 

normally unbranched, hollow, jointed stems 

(culms) reach nearly 2.5 cm in diameter but 

usually measure 1.0 to 1.5 cm in diameter. In 

addition to rhizomes, plants occasionally produce 

stolons. Alternate leaves are glabrous except for 

being strongly scabrous at the margins. They are 

flat, 1 to 5 cm wide, and up to 50 cm long. 

Terminal plume-like panicles are 15 to 50 cm long 

with branches ascending to nodding and densely 

flowered. The panicles are initially grayish purple, 

becoming whitish, and finally weathering to light 

brown. There are 2n = 36, 48, and 54 

chromosomes (Duke 1998, Gould 1975, 

Harrington 1964, Uchytil 1992, Welsh and others 

1987). 

Range.—Common reed occurs in every state in 

the Continental United States except Arkansas 

(probably present but undocumented), in Puerto 

Rico (Natural Resources Conservation Service 

2003), southern Canada, and Mexico. In fact it is 

found on all continents except Antarctica 

(Harrington 1964). It is native to the United States 

and is known from the archeological record. 

However, genetic analysis has shown that more 

aggressive European genotypes have replaced the 

native genotype in much of the former American 

range and have invaded new areas (Blossey 2002). 

Ecology.—Common reed is a warm-season grass 

that starts its growth after the last frost has 

occurred but remains green until frost in temperate 

areas (Uchytil 1992). Wide climatic tolerance is 

indicated by its range that stretches from the 

tropics to cold temperate areas and habitat from 

sea level to elevations of at least 1,980 m 

(Harrington 1964, Welsh and others 1987). 

Common reed typically forms pure stands that 

contain up to 205 stems/m2

 (Uchytil 1992). In Gulf 

Coast marshes, it is often codominant with 

Spartina cynosuroides (L.) Roth. (Duke 1998). 

The species grows on the shores of streams and 

lakes, and in marshes, both fresh and brackish. 

Although common reed grows best in clay, it 

occurs in a wide variety of soils. The species 

tolerates flooding, frost, high pH, salt, and weeds 

(Duke 1998). Stands in the Nile River Delta 

tolerate soil pH’s of 7.0 to 9.3 (Serag 1996). 

Standing dead material often totals twice as much 

biomass as current growth, allowing stands to burn 

even during the growing season. New stands arise 

from rhizomes that are under the soil or water. The 

species also tolerates late season drought (author, 

personal observation). Moderately deep rooting (to 

1 m, Uchytil 1992) undoubtedly imparts drought 
556 

resistance. A large number of fungal species and at 

least one nematode have been isolated from the 

species, none of which cause extensive damage 

(Duke 1998). 

Reproduction.—Common reed blooms mostly 

from July through November (Diggs and others 

1999). It apparently blooms and fruits throughout 

the year in frost-free areas (Gould 1975). Under 

favorable artificial conditions, germination begins 

1 day after sowing and may reach nearly 100 

percent (Ekstam and Forseby 1999). Although 

common reed is a relatively good seed producer 

and seeds are added annually to the soil seed bank, 

successful seedling establishment is rare (Uchytil 

1992). New stands can also be started by floodborne pieces of rhizome. Once a new plant is 

established, it spreads by vegetative means. 

Growth and Management.—Because stems are 

renewed each growing season, annual height 

growth is from 2 to 4 m per year. Growth can 

reach 4 cm/day. In southern Manitoba, stems reach 

their full height by the end of July (Uchytil 1992). 

During periods of falling water, colonies can 

increase in width by as much as 15 m in a single 

season (Fewless 2003). Annual productivity ranges 

from 7.5 to 63 metric tons/ha (Duke 1998). If 

harvested during the dormant season, productivity 

is not affected, except through compaction. New 

stands are established by planting rhizome 

segments (Duke 1998). This is normally not 

recommended because of the invasive nature of the 

species. Eliminating stands is difficult. The best 

approaches are heavy grazing by cattle, and 

spraying with amatrol, dalapon, or glyphosate with 

follow-up treatments. Stands can be degraded by 

cultivation or summer mowing (Uchytil 1992). 

Benefits and Detriments.—Common reed 

aggressively colonizes large areas of shoreline and 

shallow marsh, and with its thick sod and heavy 

stand of stems effectively prevents wave- and 

current-caused erosion (Uchytil 1992). It also traps 

sediment and pollutants. In fact the species has 

been used to aid in settling and drying sewage 

sludge (Graduate College of Marine Studies 2003). 

The dense single-species stands completely 

displace other native marsh communities (Fewless 

2003) and many of the fauna they support 

(Weinstein and Balletto 1999). On the other hand, 

strong declines in populations in many European 

countries have occurred during the last two 

decades (Clevering 1998). Native Americans used 

the sap from green stems to loosen phlegm and 

sooth lung pain, especially during pneumonia, and 

to treat stomach problems and diarrhea (Moerman 

1986). There are numerous other applications in 

herbal medicine throughout the world. The stems 

were used as arrow shafts by Native Americans 

(Duke 1998). Native people of Tasmania used the 

hollow stems to make rafts, jewelry, baskets, and 

light spears (Australian National Botanic Gardens 

2003). Common reeds are harvested today in 

Britain for thatching (Tyler-Walters 2002). They 

were once harvested for fuel where better sources 

were unavailable and may hold promise as an 

energy crop in the future. Stands yield 415 

calories/100g (Duke 1998). Early in the growing 

season, common reed is high-quality forage for 

cattle and horses and may be cut for hay. It 

contains 11.4 percent protein, 2.3 percent fat, 42.1 

percent carbohydrates, 31.1 percent crude fiber, 

and 10.8 percent ash (Duke 1998). It becomes 

woody and unpalatable later in the season. The 

seeds are eaten by waterfowl and the roots are 

eaten by muskrats. The edges are occasionally 

used for nesting by ducks and the interior of the 

stands are nesting habitat for snowy egrets, blackcrowned night herons, and yellow-headed 

blackbirds. A few species such as deer use the 

stands for escape cover (Uchytl 1992). It is 

important habitat for wildlife in Britain and 

supports such species as the bittern, the reed 

bunting, and the marsh harrier (Tyler-Walters 

2002). Throughout the world the stems have been 

used for building materials, lattice work, baskets, 

and mats. The young shoots and rhizomes are 

sometimes eaten, and the rhizomes processed for 

starch. The stems are processed into pulp, paper, 

and fiberboards. Stalks contain over 50 percent 

cellulose and have fibers 0.8 to 3.0 mm long by 

5.0 to 30.5 μm in diameter. The dried plumes are 

used for decorations and a variegated variety is 

employed as an ornamental (Duke 1998). 

References

Australian National Botanic Gardens. 2003. 

Aboriginal plant use in South-eastern Australia: 

Phragmites australis. http://www.anbg.gov. 

au/aborig.s.e.aust/phragmites-australis.html. 2 p. 

Blossey, B. 2002. Native to North America or 

introduced (or both)? Ag Web, Cornell 

University, Ithaca, NY. http://www. 

invasiveplants.net/phragmites/phrag/natint.htm. 

3 p. 
557

Britton, N.L. and A. Brown. 1913. Illustrated flora 

of the northern states, Canada, and the British 

possessions. Vol. 2, 2nd Ed. Scribner, New York. 

735 p. 

Clevering, O.A. 1998. An investigation into the 

effects of nitrogen on growth and morphology of 

stable and die-back populations of Phragmites 

australes. Aquatic Botany 60(1): 11-25. 

Diggs, G.M., Jr., B.L. Lipscomb, and R.J. 

O’Kennon. 1999. Thinner’s and Mahler’s 

Illustrated flora of North Central Texas. 

Botanical Research Institute of Texas, Fort 

Worth, TX. 1,622 p. 

Duke, J.A. 1998. Handbook of energy crops: 

Phragmites australis (Cav.) Trin. ex Steud. 

Center for New Crops and Plants Products, 

Purdue University, West Lafayette, IN. http:// 

 www.hort.purdue.edu/newcrop/duke_energy/ 

phragmites_australis.html. 5 p. 

 

Ekstam, B. and A. Forseby. 1999. Germination 

response of Phragmites australis and Typha 

latifolia to diurnal fluctions in temperature. Seed 

Science Research 9(2): 157-163. 

Fewless, G. 2003. Invasive plants of Wisconsin: 

Phragmites australis: common reed. http:// 

www.uwgb.edu/biodiversity/herbarium/invasive

_species/phraus01.htm. 2 p. 

Gould, F.W. 1975. The grasses of Texas. Texas 

A&M University Press, College Station, Texas. 

653 p. 

Graduate College of Marine Studies. 2003. 

Optimized reed (Phragmites australis) function 

in sludge drying beds: producing a more 

effective genotype. University of Delaware, 

Newark, DL. http://www.ocean.udel.edu/ 

level1/facultystaff/faculty/dseliskar/sludgebeds.

html. 3 p. 

Harrington, H.D. 1964. Manual of the plants of 

Colorado. Sage Books, Denver, CO. 666 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports. 19. University of 

Michigan Museum of Anthropology, Ann 

Arbor, MI. 534 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Phragmetes australis (Cav.) Trin. 

ex Steud. http://plants.usda.gov/cgo_bom/ 

plant_search.cgi?mode=Scientific+Nave&keyw

ordquery=Phragmites+australis&earl=plant_s... 

6 p. 

Serag, M.S. 1996. Ecology and biomass of 

Phragmites australes (Cav.) Trin. ex Steud. In 

the north-eastern region of the Nile Delta, 

Egypt. Ecoscience 3(4): 473-482. 

Tyler-Walters, H. 2002. Phragmites australis, 

common reed. Marine Life Information 

Network, Plymouth, UK. http://www.marlin. 

ac.uk/species/Phraus.htm. 4 p. 

Uchytil, R.J. 1992. Phragmites australis. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/gramin

oid/phraus/all.html. 19 p. 

Weinstein, M.P. and J.H. Balletto. 1999. Does the 

common reed, Phragmites australis, affect 

essential fish habitat? Estuaries 22(3B): 793-

802. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 
558 

Physocarpus malvaceus (Greene) Kuntze mallow ninebark 

ROSACEAE 

Synonyms: Spiraea oputifolia var. pauciflora Torr. & Gray 

 Neillia malvaceae Greene 

 Opulaster malvaceus Kuntze 

 Physocarpus pauciflorus Piper 

John K. Francis 

General Description.—Mallow ninebark, also 

known as ninebark, mallow-leaved ninebark, 

mountain ninebark, few-flowered ninebark, and 

shallow ninebark, is a multistemmed, deciduous 

shrub 1 to 2 m (rarely 3 m) in height. The stems 

and branches are slender and covered with a brown 

bark that exfoliates in papery strips on older 

branches. The 3- to 5-cm, alternate leaves are 

round to ovate, three to five lobed, and doubly 

serrate or crenate. The petioles are 1 to 2 cm long. 

Inflorescences are terminal, corymbose with five 

to 30 flowers. The hypanthium of the flowers is 

stellate-pubescent; twigs, petioles, and leaf blades 

vary from pubescent to glabrous. The flowers have 

five sepals and five white, rounded petals 3.5 to 

6.5 mm long. Each produces two inflated, brown 

follicles united for two-thirds their length. Each 

fruit may contain one to several shiny, pairshaped, straw-colored seeds (Abrams 1950, Welsh 

and others 1987). 

Range.—Mallow ninebark is native to British 

Columbia, Alberta, Washington, Oregon, Idaho, 

Montana, Wyoming, Colorado, Utah, and Nevada 

(Abrams 1950, Natural Resources Conservation 

Service 2003). It is not known to have naturalized 

outside its native range. 

Ecology.—Mallow ninebark grows on a wide 

variety of soils derived from igneous, 

metamorphic, and calcareous and noncalcareous 

sedimentary rocks. Textures range from sandy 

loams to silty clay loams and surface pH ranges 

from 5.5 to 7.1. In Idaho and Montana, the average 

duff layer thickness was 6.5 cm with an effective 

rooting depth of 38 cm. In northern areas, mallow 

ninebark is found mainly in areas with no exposed 

rock on mountain slopes, ridges, and streambanks 

(Habeck 2003); in Utah, it often grows in skeletal 

soils and rubble in steep canyons and 

mountainsides. It occurs at elevations of 1,600 to 

3,300 m at the southern end of its Utah range 

(Welsh and others 1987) and at somewhat lower 

elevations farther north. The species is associated 

with Abies lasiocarpa (Hook.) Nutt., Abies grandis 

(Dougl. ex D. Don) Lindl., and Picea engelmannii

Perry ex Engelm. in cool/moist habitat and with 

Pseudotsuga menziesii (Marbel) Franco and Pinus 

ponderosa P. & C. Lawson in warmer, moist sites 

(Habeck 2003). While associated with these trees, 

mallow ninebark usually grows in disturbancecaused openings or diffuse stands. It also grows on 

nonforested shrubland sites. Some common shrub 

associates are Holodiscus discolor (Pursh) 

Maxim., Symphoricarpos albus (L.) Blake, 

Symphoricarpos oreophilis Gray, Spirea butilifolia 

Pallas, Amelanchier alnifolia (Nutt.) Nutt. ex M. 

Roemer, and Mahonia repens (Lindl.) G. Don 

(Habeck 2003). Mallow ninebark is fire resistant, 

sprouting after topkill from surviving root crowns 

or horizontal rhizomes (Palouse Prairie Foundation 

2003). Having 36 to 99 percent of its roots buried 

in mineral soil enables it to escape the heat of 

surface fires (Habeck 2003). Mallow ninebark 
559

stands often owe their existence to logging or fires. 

The species is an obligate pioneer that increases 

and grows rapidly following disturbance, but as 

competition, particularly overhead shade, 

increased, it decreases in mean height and 

gradually disappears (Habeck 2003). The species 

is generally free of insect and disease problems, 

although it is occasionally attacked by foliageeating and seed-eating insects (Youngblood and 

others 2003). 

Reproduction.—Flowering takes place between 

late May and late July, depending on altitude, and 

fruits ripen between late August and late 

September (Habeck 2003). Seed weights are 

reported (geographic source unspecified) at 

165,000 seeds/kg (Youngblood and others 2003). 

Seeds collected by the author in Utah averaged 

158,000 seeds/kg. Viability is usually less than 50 

percent (Youngblood and others 2003). The fruits 

split open at maturity but most of the seeds do not 

immediately fall. Dispersal is by gravity, wind, 

and probably by browsing animals. Seeds in the 

soil seedbank are reported to be 11 percent viable 

and will germinate when the surface is scarified 

(Habeck 2003). Ripe fruits can be picked by hand 

or beaten off shrubs onto a tarp. Fruits may be 

threshed by rubbing for small quantities or passed 

through a hammer mill for larger quantities and 

screened and blown to clean them. They may be 

stored cool and dry for at least 5 years. Seeds may 

be sown in the fall, or in the spring after 30 days of 

prechilling. Seeds should be sown to a depth of 3 

mm and mulched with another 6 mm of sawdust 

(Youngblood and others 2003). Mallow ninebark 

can easily be vegetatively reproduced with misted 

softwood cuttings or by planting hardwood 

cuttings directly in the field (Youngblood and 

others 2003). 

Growth and Management.—Mallow ninebark 

shrubs in northern Idaho reached their maximum 

height by the fourth growing season after a fire 

(Habeck 2003). Although stems are added and lost 

continuously, shrubs do not grow taller. Individual 

plants may live for many years if growing 

conditions permit. Mallow ninebark is sometimes 

planted for soil stabilization and other 

conservation objectives. Killing the shrubs to 

make room for more desirable species is 

sometimes attempted. Several herbicides can be 

effective if sprayed early in the season from late 

foliar development to flowering or in early fall 

before leaves change color. Sprouts will 

sometimes appear later and will have to be sprayed 

again (Habeck 2003). 

Benefits.—Mallow ninebark helps protect the soil 

during secondary succession until the shrub is 

replaced by taller vegetation. It adds beauty to 

forest openings and shrublands when its leaves 

turn russet-red in the fall (Clark 1976). The species 

provides cover for both large and small animals. It 

is one of the principal nesting sites of the dusky 

flycatcher (Empidonax oberholseri) (Sedgwick 

1993). The shrub is not browsed by cattle and is 

utilized only to a limited extent by deer, bighorn 

sheep, and domestic sheep. The upper third of the 

annual growth of plants from a burned area 

contained 1.3 percent N, 0.3 percent P, and 1.2 

percent Ca. Plants in an unburned site contained 

even lower levels (Habeck 2003). Early summer 

sheep grazing increases the nutritional quality of 

mallow ninebark for fall and winter browse (Alpe 

and others 1999). 

References 

Abrams, L. 1950. Illustrated flora of the Pacific 

States. Vol. 2. Stanford University Press, 

Stanford, CA. 635 p. 

Alpe M.J., J.L. Kingery, and J.C. Mosley. 1999. 

Effects of summer sheep grazing on browse 

nutritive quality in autumn and winter. Journal 

of Wildlife Management 63(1): 346-354. 

Clark, L.J. 1976. Wild flowers of the Pacific 

Northwest. Gray’s Publishing Limited, Sidney, 

BC, Canada. 604 p. 

Habeck, R.J. 2003. Physocarpus malvaceus. In: 

U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shrub/ 

phymal/all.html. 17 p. 

Natural Resources Conservation Service. 2003. 

Plant profile: Physocarpus malvaceus (Greene) 

Kuntze. Plants database. http://plants.usda.gov/ 

cgi_bin/plant_profile.chi?symbol=PHMA5. 3 p. 

Palouse Prairie Foundation. 2003. Palouse Prairie 

Foundation native species search results: 

Physocarpus malvaceus. http://www. 

palouseprairie.org/readppf.pl?entry=PHMA5. 

3 p. 
560 

Sedgwick, J.A. 1993. Reproductive ecology of 

dusky flycatchers in western Montana. Wilson 

Bulletin 105(1): 84-92. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 

Youngblood, A.Y., J.D. Gill, and F.L. Pogge. 

2003. Physocarpus (Camb.) Raf. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. http://wpsm.net/Physocarpus. 

pdf. 5 p. 
561

Picramnia pentandra Sw. Florida bitterbush 

SIMAROUBACEAE 

 

Synonyms: Tariri pentandra (Sw.) Baill. 

 Picramnia antidesmoides Griseb. 

 Picramnia cubensis Turcz. 

 Picramnia micrantha Tulasne 

 Picramnia oblongifolia Turcz. 

John K. Francis 

General Description.—Florida bitterbush is also 

known as bitterbush, doctor-bar, wild-coffee, 

guarema, hueso, aguedita, palo de peje, palo de 

paz, roble agalla, quina del país, bois poison, 

vaillant garcon, and bois montagne. It is an 

evergreen shrub or small tree usually less than 6 m 

in height and 10 cm in stem diameter. The plant 

usually has a single slender stem with smooth, 

gray bark and relatively few, slender branches. The 

inner bark is brown. The wood is light colored, 

hard, and heavy. The inner bark, roots, leaves, 

fruits, and seeds have a strongly bitter taste. The 

alternate, compound leaves are 10 to 30 cm long 

and contain five to nine, usually seven, 5- to 10-

cm leaflets. The elliptic to ovate, papery leaflets 

have short stalks, smooth edges, and long points. 

The side leaflets are subopposite. The leaflets are 

green tinged with red, have a net-like structure of 

fine veins when viewed against the light, and are 

shiny when dried. Male and female flowers occur 

on different trees (dioecious) in branched terminal 

clusters. The tiny flowers are five-parted, green 

with a reddish tinge. Fruits (berries) are scarlet to 

almost black when ripe, oblong to obovoid, 9 to 15 

mm long, and contain one to three shiny brown 

seeds 6 to 10 mm long (Howard 1988, Liogier 

1988, Little and Wadsworth 1964, Long and 

Lakela 1976). 

Range.—Florida bitterbush is native to southern 

Florida, the Bahamas, the West Indian Islands, and 

Colombia and Venezuela in northern South 

America (Howard 1988, Little and Wadsworth 

1964). It is not known to have naturalized outside 

its native range. 

Ecology.—Florida bitterbush occurs in coastal 

hammocks in southern Florida and the Keys (Long 

and Lakela 1976). It grows in both remnant forests 

and secondary forests (logged, pastured, and 

abandoned fields) in Puerto Rico. During old-field 

or pasture succession, Florida bitterbush invades 

during the brushy stage. The species adapts to a 

wide variety of well-drained soils derived from 

most parent materials. It usually does not grow on 

the most fertile sites (where competition is severe) 

or on the worst, such as eroded, rocky ridge tops. 

Elevations vary from near sea level to 600 m or 

more, and mean annual precipitation ranges from 

750 mm to about 2400 mm. Florida bitterbush is 

moderately intolerant to shade, being able to grow 

under the canopy of low basal-area forests as well 

as in openings. Seedlings are apparently more 

tolerant than adults and are able to progress from 

the understory into the canopy of low forests 

(Pascarella 1996). 

Reproduction.—Florida bitterbush flowers and 

fruits nearly throughout the year in Puerto Rico 

(Little and Wadsworth 1964). It blooms March 

through July in Florida. Flowers of both sexes 

attract small, generalist insect pollinators with 

scent, pollen, and small amounts of nectar. Peak 
562 

ripening occurs in November and December 

(Pascarella 1996). Fruit and seed production is 

moderately abundant and consistent from year to 

year. Fresh fruits collected in Puerto Rico weighed 

an average of 0.231 + 0.043 g/fruit. Air-dried 

seeds separated from them averaged 11,900 

seeds/kg. Placed on moist filter paper without 

pretreatment, 100 percent germinated, beginning 

14 days after sowing (Francis and Rodríguez 

1993). Pascarella (1996) noted fruits collected in 

Florida weighed an average of 0.483 + 0.034 g and 

that 85 to 90 percent of the fruits contained just 

one seed. Most seeds fall under the parent trees but 

some are dispersed by birds. Seedlings are 

relatively common in forests where the species 

grows. 

Growth and Management.—Growth rate of 

Florida bitterbush is slow to moderate. Weaver 

(1979) followed the growth of two plants in a 

forest in Puerto Rico with characteristic slow 

growth and found diameter growth rates over the 

24-year period of about 0.2 cm/year. Florida 

bitterbush in Puerto Rico live about 30 to 60 years 

(author’s observation). Plants begin flowering and 

fruiting when they are about 1.5 to 2 m in height if 

sufficient light is available. Fruits are easily 

collected in quantity by hand and with short 

pruning poles. They may be cleaned by macerating 

or by working them against a screen under running 

water. Production of seedlings, at least to the 

pricking-out stage, is straightforward. Where a 

seed source is available in favorable habitat, 

natural reproduction is usually adequate. Florida 

bitterbush has not been reported to cause any 

problems in forests and therefore control measures 

are probably not needed. 

Benefits.—Florida bitterbush helps protect the soil 

and contributes to the biodiversity and aesthetics 

of forest stands. A number of bird and animal 

species eat the fruits. It furnishes larval food for 

the bush sulphur butterfly, Urema dina helios

(Institute of Food and Agricultural Sciences 2003). 

Florida bitterbush is a honey plant (Little and 

Wadsworth 1964). The wood is little used, except 

occasionally for fuel. Florida bitterbush is planted 

in Florida and Cuba as an ornamental (Little and 

Wadsworth 1964). A bitter tonic is made from 

tissues of the species in the Caribbean to treat 

fevers, anorexia, and stomach problems and as an 

enema to treat for worms (Liogier 1990). 

Qualitative tests indicated the presence of 

antioxidants, phenols, terpenoids, and alkaloids in 

ethanol extracts of tissues. A number of bacteria 

and fungi were treated with ethanol extracts of 

various tissues. Growth reductions of Bacillus 

cereus, Bacillus subtilis, and Pseudomonas 

aeruginosa were observed which was enhanced by 

exposure to long-wave ultraviolet radiation 

(O’Neal and others 2002). 

References 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Institute of Food and Agricultural Sciences. 2003. 

Native landscape plants for South Florida: 

shrubs and small trees (P-Z). http://miamidade.ifas.ufl.edu/programs/fyn/publications/nati

veplantlist/npl-shrubs-small-trees-p-z.htm. 5 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. University of Miami Press, 

Coral Gables, FL. 962 p. 

O’Neal, C.B., J. Salazar, and E. Rodríguez. 2002. 

A preliminary analysis of the anti-microbial and 

phytochemical properties of Picramnia 

pentandra (Simaroubaceae) and its potential 

relationship to gastrointestinal health. 

Emanations 4: 57. 
563

Pascarella, J.B. 1996. Reproductive ecology of 

Picramnia pentandra (Picramniaceae) in South 

Florida. Caribbean Journal of Science 32(1): 99-

104 

Weaver, P.L. 1979. Tree growth in several tropical 

forests of Puerto Rico. Research Paper SO-152. 

U.S. Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 15 p. 
564 

Piper aduncum L. bamboo piper 

PIPERACEAE 

Synonyms: Artanthe adunca Miq. 

 Piper hebecarpum C. DC. in Urban 

 Piper martinicense C. DC. in Briq. 

 Piper stehleorum Trel. in Stehlé 

 Piper subrectinerve C. DC. in Urban 

John K. Francis 

General Description.—Bamboo piper is also 

known as jointwood, spiked pepper, false kava, 

cow’s foot, higuillo, higuillo de hoja menuda, 

cordoncillo, anisillo, guayayo, Santa María negra, 

aperta ruão, and matico. It is a multistemmed 

evergreen shrub or small tree 1 to 6 m in height 

and up to 10 cm or more in stem diameter. The 

wood is whitish, hard, and brittle with a thick, 

solid pith, and no annual rings. The stems have 

swollen nodes and bark that is smooth and gray or 

green. Plants are supported by abundant lateral 

roots and, at times, proproots arising from nodes 

near the base. The green twigs are not abundant. 

The alternate leaves have short petioles and 

elliptic to lanciolate blades 12 to 20 cm long. All 

plant parts have a peppery taste and odor. The 

cordlike spikes are borne opposite leaves and 

contain many, minute flowers that soon develop 

into numerous tiny, imbedded drupes with brown 

or black seeds (Howard 1988, Liogier 1985, Little 

and Wadsworth 1964, Stevens and others 2001). 

Range.—Bamboo piper is native to the Greater 

and Lesser Antilles and on the mainland from 

Mexico to northern Argentina. The species has 

been widely planted as an ornamental and has 

spread by the movement of equipment between 

land masses. It has escaped and established itself 

in Florida, Southeast Asia, and a number of Pacific 

Islands (Instituto de Botánica Darwinion 2002, 

Howard 1988, Pacific Island Ecosystems at Risk 

2002, Plant Protection Service 2001, Stevens and 

others 2001). 

Ecology.—Bamboo piper grows in areas that 

receive from about 1500 to over 4000 mm of 

mean annual rainfall. It grows in Puerto Rico from 

near sea level to over 400 m in elevation. Most 

soils are colonized including fill dirt, but not salty 

soils and excessively well-drained soils at the dry 

end of the range. Disturbance, which is required 

for establishment, includes removal of the forest 

canopy to allow light to enter and the creation of a 

bare soil surface, although plants occasionally 

establish themselves on rotting logs and stumps. 

Bamboo piper is moderately intolerant of shade. It 

can survive and grow slowly under a moderate 

overstory but requires at least partial exposure to 

grow large and flower. The species commonly 

grows in clearcuts, tree-fall gaps, shade-grown 

coffee and other tree plantations, brushy pastures, 

roadsides, and landslides. Bamboo piper grows as 

individual plants or in thickets. As many as 2,632 

plants/ha were reported in areas invaded in the 

Philippines (Philippine Council of Agriculture 

2002). 

Reproduction.—Bamboo piper flowers and fruits 

throughout the year (Little and Wadsworth 1964). 

Air-dried seeds from a Puerto Rican source 

averaged 0.000237 g/seed or 4.2 million/kg. Only 

7 percent germinated when placed on moist filter 
565

paper (author’s observation). The seeds are 

dispersed by bats, birds, and possibly arboreal 

rodents (Plant Protection Service 2001). Seedlings 

are fairly common on disturbed ground. 

Established plants thicken into clumps by suckers 

arising from the root crown (Pacific Island 

Ecosystems at Risk 2002). Plants are established 

for agroforestry purposes in Papua New Guinea by 

shoving cuttings into moist soil (Bourke 1997). 

Growth and Management.—The growth of 

bamboo piper is moderately rapid after the early 

seedling stage. Sprouts and suckers grow more 

than a meter in their first year. Producing 

seedlings by means of seed is difficult. 

Propagating plants with cuttings, which need no 

hormonal treatment, in the nursery or directly in 

the field, is recommended. Individual stems live 2 

to several years; by sprouting, plants live much 

longer. Infestations of bamboo piper can be 

controlled by uprooting young plants and spraying 

older plants with broadleaf weed killers such as 

2,4-D. Treated areas should be checked in a few 

weeks and treated again if regrowth has occurred 

(Plant Protection Service 2001). 

Benefits.—Bamboo piper helps revegetate 

disturbed areas and contributes to the biodiversity 

and biomass of forests. It also is a source of food 

and cover for wildlife. Bamboo piper established 

in contour rows for soil erosion brakes helps 

facilitate agroforestry on steep land in Papua New 

Guinea (Bourke 1997). The wood is useful for 

fuel, stakes, fences, and rude construction 

(Philippine Council of Agriculture 2002). 

However, exposed wood rots quickly (Vélez and 

van Overbeek 1950). The species is planted as an 

ornamental. The peppery fruits have been used to 

season food (Little and Wadsworth 1964). 

Essential oil content of bamboo piper tissue 

(leaves and twigs), of which dillapiole is the major 

component, ranged from 1.2 to 3.4 percent (Maia 

and others 1998). Teas and other extracts of the 

leaves and roots of bamboo piper are used in 

herbal medicine as a tonic to ease diarrhea, 

dysentery, vomiting, ulcers, and to control 

bleeding (Liogier 1990). The chemical 2’, 6’-

dihydroxy-4’-methoxychalcone isolated from 

bamboo piper inhibited 98 percent of the growth 

of Leishmania amazonensis parasites in vitro with 

low host cell toxicity (Torres-Santos and others 

1996). In addition, the essential oils have shown 

strong insecticidal, molluscicidal, and antibacterial 

effects (Gómez and others 1997, Ibrahim and 

others 1996, Orjala and others 1992). 

References 

Bourke, R.M. 1997. Management of fallow species 

composition with tree planting in Papua New 

Guinea. http://coombs.anu.edu.au/Depts/ 

RSPAS/RMAP/bourke.htm. 8 p. 

Gómez, P., D. Cubillo, G.A. Mora, and L. Hilje. 

1997. Evaluation of possible repellents for 

Bemisi tabaci. II. Botanical substances. Manejo 

Integrado de Plagas 46: 17-25. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Ibrahim, J. A. Abu-Said, A. Abdul-Rashih, M.A. 

Nor-Azah, M.Z. Zaridah, A.K. Azizol, NorharaHussein, P.S. Bacon, and K.C. Khoo. 1996. 

Essential oils of selected Malaysian plants and 

their potential uses. Forestry and Forest Products 

Research, Proceedings of the Third Conference, 

Oct. 3-4, 1995; Kuala Lumpur, Malaysia. p. 97-

103. 

Instituto de Botánica Darwinion. 2002. Catálogo 

de las plantas vasculares de la Argentina. 

http://www.darwin.edu.ar/Catalogo/indicevascul

ares.htm. [n.p.]. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

Maia, J.G.S., M.G.B. Zohhbi, E.H.A. Andrade, 

A.S. Santos, M.H.L. da Silva, A.I.R. Luz, C.N. 

Bastos, and M.H.L. da Silva. 1998. Constituents 

of the essential oil of Piper aduncum L. growing 

wild in the Amazon region. Flavour and 

Fragrance Journal 13(4): 269-272. 

Orjala, J., A.D. Wright, T. Rali, and O. Sticher. 
566 

1992. Three new prenylated benzoic acid 

derivatives and molluscicidal sesquiterpenoids 

from Piper aduncum leaves. Medica 58(7): 714. 

Pacific Island Ecosystems at Risk. 2002. Invasive 

plant specie, Piper aduncum L. Rutaceae. 

http://www.hear.org/pier3/ piadu.htm. 2 p. 

Philippine Council of Agriculture. 2002. Research 

highlights 1997. http://www.pcarrd.dost.gov.ph/ 

division/FERD/new/highlights97.html. 18 p. 

Plant Protection Service. 2001. Pest alert: false 

kava. Pest Alert No. 19. http://www.spc.org.nc/ 

pps/PestAlerts/PestAlertNo19-False_kave.pdf. 

2 p. 

Stevens, W.D., C. Uloa-U., A. Pool, O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanical Garden Press. p. 1911-

2666. 

Torres-Santos, E.C., V.C. Moura, G.M. Sperandio, 

D.L. Moreira, M.A. Kaplan, and B. RossiBergmann. 1996. Anti-leishmanial effect of a 

pure chalcone isolated from Piper aduncum

(Piperaceae). Memórias Vol. 91 (Supplement). 

Instituto Oswaldo Cruz, Rio de Janeiro, RJ, 

Brazil. 18 p. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Río Piedras, PR. 497 p. 
567

Piper amalago L. higuillo de limón 

PIPERACEAE 

Synonyms: Enckea amalago (L.) Griseb. 

 Enckea seiberi Miq. 

 Piper medium Jacq. 

 Piper seiberi (Meq.) DC. 

John K. Francis 

General Description.—Higuillo de limón is a 

Spanish name meaning lemon pepper. Although 

the species is a pepper with the characteristic 

peppery taste of foliage and seeds, the pungent, 

disagreeable odor of crushed leaves, stems, and 

fruits hardly suggests lemon. Other common 

names include joint wood, soot-soot, guayuyo, 

cordoncillo, anisillo, alcotán, anisetto, and 

malimbé (Little and others 1974). 

Higuillo de limón is a shrub that 

sometimes becomes a small tree. It is recognized 

by its enlarged, ringed nodes on smooth, green 

stems with many brown lenticels. Stems on older 

plants become gray. Higuillo de limón often has 

multiple stems from the base, but usually does not 

branch further until the crown is reached. The 

crowns have relatively few branches and may be 

flattened in older individuals. The wood is of 

moderate density and brittle. A taproot forms in 

seedlings, but secondary roots that fan out and 

down later match it in size, and even become 

thickened in older plants. The alternate, dark 

green, hairless leaves are 4 to 14 cm long and 2 to 

7 cm broad. They are pointed at the tip, ovate to 

elliptic, and have palmate venation with five main 

veins. Tiny flowers are borne in compact, cordlike, gray-green spikes 6 to 12 cm long. At 

maturity, the small (1.5 mm) fruits (drupes) are 

closely packed along the spike (Liogier 1985, 

Little and others 1974). 

Range.—Higuillo de limón is native to tropical 

Mexico, Central America, South America, and the 

Greater and Lesser Antilles (Howard 1988). 

Ecology.—Higuillo de limón is primarily an 

understory species of old growth and secondary 

forests. It is also found in openings and along 

roads. The species will endure all but the densest 

shade under stands, but needs openings or at least 

intermediate crown positions in open forest or 

brush thickets to reproduce. However, higuillo de 

limón is rarely among the pioneers that colonize 

disturbed sites, tending instead to invade after a 

low stand of trees or brush has become 

established. It endures competition with herbs, 

trees, and shrubs, but does not survive in deep 

grass swards. The species will grow in soils with 

textures from sand to clay and in all drainage 

classes except excessively drained and very poorly 

drained. It does not appear to be sensitive to 

differences in soil parent material. In Puerto Rico, 

higuillo de limón grows in areas receiving from 

about 1200 to 2500 mm of annual precipitation 

and from near sea level to more than 600 m in 

elevation. 

Reproduction.—Higuillo de limón flowers and 

fruits throughout the year (Little and others 1974). 

In Costa Rica, frugivorous bats are the most and 

perhaps the only important dispersers of higuillo 

de limón seeds (Elizondo 2000, Fleming and 

others 1977). One sample of fruits from Puerto 

Rico yielded 890,000 seeds/kg. On moist filter 

paper, these seeds gave 77 percent germination 

between 8 and 37 days after sowing.
568 

Growth and Management.—In Puerto Rico, 

higuillo de limón reaches 5.5 m in height and 8 cm 

in diameter (Little and others 1974). The species 

may become a tree to 15 m in high forests in 

Mexico, although it remains a shrub or small tree 

in successional forests and gaps (Chazdon and 

others 1988). Its stems have no visible annual 

rings and are therefore difficult to age. Judging 

age by stem internodal segments, individual stems 

in Puerto Rico appear to reach 5 to 10 years old. 

Because resprouting occurs from the roots, 

individual plants may live much longer. Natural 

regeneration of higuillo de limón occurs readily in 

early secondary forest and in open stands of older 

forest when seed sources are present. Stands can 

be easily manipulated to encourage it. Small 

quantities of seed have been separated by hand 

with tweezers after mashing the fruit. Larger 

quantities probably can be obtained by gentle 

maceration and wet sieving. Germination on the 

surface of wet soil or peat is recommended. 

Plantations could be established using 

containerized seedlings, although no plantings 

have been reported.

Benefits.—Higuillo de limón is eaten by cattle 

after the more desirable grasses and forbs are 

gone. Infusions of the leaves are said to alleviate 

colic and intestinal gas. The roots are used as a 

diuretic and to treat water retention (Liogier 

1990). The fully ripe fruits and their seeds have 

the same taste as black pepper, Piper nigrum L., 

and are sometimes used as a substitute for it, 

especially in Jamaica. The fruits are usually dried 

whole and ground as needed (Grieve 2001).

References 

Chazdon, R.L., K. Williams, and C.B. Field. 1988. 

Interactions between crown structure and light 

environment in five rain forest Piper species. 

American Journal of Botany 75(10): 1,459-

1,471. 

Elizondo C., L.H. 2000. Artibeus toltecus

Saussure. http://www.Inbio.ac.cr/bims/ubi/ 

mamiferos/ubiespejo/ubiid=1557&-find.html. 

 5 p. 

Fleming, T.H., E.R. Heithaus, and W.B. Sawyer. 

1977. An experimental analysis of the food 

location behavior of frugivorous bats. Ecology 

58(3): 619-627. 

Grieve, Mrs. M. 2001. A modern herbal: pepper. 

http;//www.botanical.com/botanical/mgmh/p/ 

pepper24.html. 5 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., Río Piedras, PR. 566 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 
569

Piper swartzianum (Miq.) C. DC. higuillo 

PIPERACEAE 

Synonyms: Ariantha swartziana Miq. 

John K. Francis 

General Description.—This shrub, known by its 

Spanish name higuillo in Puerto Rico (Little and 

others 1974), has been assigned the name Spanish 

elder in English (Natural Resources Conservation 

Service 2002). The species is an evergreen shrub 

that may take on vine-like or tree-like form. 

Higuillo usually has heights of 1.5 to 3 m but may 

reach 4.5 m in height and 7.5 cm in stem diameter 

(Little and others 1974). Plants are supported by 

relatively abundant lateral and fine roots. White 

mycorrhizal fungi associated with the fine roots 

are clearly visible. The shrubs usually have 

multiple stems that have arisen as sprouts from the 

root crown. The stems are gray with many raised 

lenticels and have enlarged nodes and a thick pith. 

The wood is light brown and brittle. The twigs are 

light green turning gray and often have a zig-zag 

form. The inner bark, twigs, and leaves have a 

pleasant spicy odor when crushed, and a peppery 

taste. The leaves are dull green to dark green, 

alternate in two rows, with minute gland dots. The 

leaf blades are oblong to lanceoate, 11 to 24 cm 

long by 3 to 7 cm broad, long-pointed at the apex 

and unequally rounded at the base. The flowers 

and fruits are grouped in 5- to 11-cm cord-like 

spikes, 4 mm thick borne singly opposite leaf 

attachments near the branch tips. The spikes are 

white during flowering, turning light green or 

yellow-green as the fruits mature. The individual 

fruits are imbedded within the spike in spirals 

(Liogier 1985, Little and others 1974). 

Range.—Higuillo is native to Puerto Rico and 

Hispaniola (Liogier 1985). It is relatively common 

to rare in the moist and wet portions of Puerto 

Rico, specifically the public forests of Guajataca, 

Maricao, and Río Abajo (Little and others 1974), 

and private lands nearby. Higuillo was 

misidentified as P. tuberculatum Jacq. in Little 

and others (1974) (Liogier 1985).

Ecology.—Higuillo inhabits middle and late 

secondary and remnant forests. It is shade tolerant 

and is usually found in the understory, often in 

fairly deep shade. Higuillo does not compete well 

in tall, dense thickets of vigorous shrubs and 

young trees. At the lower end of the rainfall range 

(about 1200 mm/yr), the species is confined to 

extra-humid situations such as deep ravines. It is 

most common in forests that receive from 1600 to 

3000 mm of precipitation. Higuillo grows on sites 

from a few meters above sea level to more than 

1,000 m in elevation on a wide variety of soils 

derived from all types of parent material including 

ultramafics and limestone. 

Reproduction.—In order to flower and fruit, 

mature higuillo plants need breaks in the forest 

canopy that admit over-head sun for part of the 

day. Vigorous plants flower and fruit continuously 

(Little and others 1974). Air-dried seeds weighed 

an average of 0.00156 g/seed or 641,000 seeds/kg 

(author’s observation). The seeds are probably 

disbursed by fruit bats that eat the fruits. Seedlings 

are widely scattered and uncommon. 

Growth and Management.—New seedlings 

grow slowly. Well-established plants and sprouts 

grow about 0.5 m/year in height. Individual stems 

live 3 or 4 years and are replaced by other sprouts 

so that established plants can live indefinitely. No 
570 

management experience has been published. 

Natural reproduction could probably be 

encouraged by disturbance that exposes mineral 

soil under a moderately shady tree canopy. 

Benefits.—Higuillo contributes to the 

biodiversity, scenic beauty, and stability of the soil 

of the forests where it grows. Like other pipers, its 

fruits are food for fruit bats. 

References 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Natural Resources Conservation Service. 2002. 

Plants profile: Piper blattarum Spreng. 

http://plants.usda.gov/cgi_bin/checklist.cgi. 2 p. 
571

Piptocoma antillana Urban Antilles velvetshrub 

ASTERACEAE 

Synonyms: Piptocoma rufescens auct. non Cass.

John K. Francis 

General Description.—Antilles velvetshrub is an 

evergreen, narrow-crowned, upright shrub usually 

1 to 2 m in height but sometimes reaching 4 m in 

height and 5 cm in basal diameter. Mature plants 

often have several stems arising from the root 

crown or low on the stem. Lower stems are 

furrowed and gray; branches are relatively few, 

slender, straight, and covered with gray bark. The 

pale yellow wood is of moderate density, 

moderately strong, and has a dark brown pith at 

the center of the stem. Plants are supported by 

flexible, weak roots covered by corky, brown bark. 

The species is easily recognized by the narrow 

leaves that are dark green above and bright white 

beneath. The narrowly elliptic to lanceolate leaves 

are 3 to 8 cm long and 0.5 to 2.5 cm broad and 

have petioles up to 5 mm long. Inflorescences are 

corymbs of numerous campanulate heads. The 

seeds are straw-colored achenes about 2 mm long, 

tipped with a pappus about 4.5 mm long (author’s 

observation, Liogier 1997). 

Range.—Antilles velvetshrub is native to Puerto 

Rico and the Virgin Islands (Liogier 1997). 

Previous reports (Liogier and Martorell 1982) that 

it occurred in Hispaniola were based on 

misidentification of P. rufescens Cass. Antilles 

velvetshrub is not known to have been planted or 

naturalized outside its native range. 

Ecology.—Antilles velvetshrub grows in coastal 

and upland sites from near sea level to 500 m or 

more in elevation. Mean annual rainfall varies 

from about 750 mm to 2000 mm. It is most 

common on ridges and hillsides of areas underlain 

with ultramafic rocks and limestone. Antilles 

velvetshrub is sometimes seen on road cuts and the 

center between the tracks of abandoned roads. 

Soils are excessively well or well drained, have 

pH’s from about 5.5 to 7.7, and vary widely in 

texture and quantity of stones in the profile. 

Antilles velvetshrub is drought tolerant--leaves 

wilt but the plants usually do not defoliate. The 

species tolerates competition from shrubs, herbs, 

and a moderate amount of grass. It grows in 

openings or under partial shade of low basal-area 

forests and may be found in remnant stands and 

mid- to late-secondary forests. The species 

probably requires bare soil or small-scale 

disturbance to establish itself. Antilles velvetshrub 

occurs as scattered to occasional plants. 

Reproduction.—Antilles velvetshrub blooms 

annually, apparently near the end of the wet 

season. Seeds were collected by the author in 

Puerto Rico in March (dry season). The air-dried 

seeds averaged 0.00046 g/seed or 2,150,000 

seeds/kg. Placed on moist filter paper without 

pretreatment, 8 percent germinated between 11 and 

14 days after sowing. Germination is epigeal. 

Seeds are dispersed by wind. Seed crops can be 

moderately abundant, but seedlings are 

uncommon. 
572 

Growth and Management.—Antilles velvetshrub 

grows 0.5 to 1.0 m per year. The stems do not live 

long--perhaps 2 to 5 years--but plants can renew 

themselves and live longer by sprouting from the 

root crown. Planting and management experience 

have not been published. Because the species is 

not abundant or aggressive, control should not be 

necessary. 

Benefits.—Antilles velvetshrub helps protect the 

soil, furnishes cover for wildlife, and adds to the 

aesthetics of the forest. If means of propagation 

can be developed, the species would be valuable 

for background and accent in naturalistic 

landscaping. 

References

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Liogier, H.A. and L.F. Martorell. 1982. Flora of 

Puerto Rico and adjacent islands, a systematic 

synopsis. Editorial de la Universidad de Puerto 

Rico, Río Piedras, PR. 342 p. 
573

Pisonia aculeata L. pull-back-and-hold 

NYCTAGINACEAE 

Synonyms: Pisonia helleri Standley 

Pisonia yaguapinda D. Parodi 

John K. Francis 

General Description.—Pull-back-and-hold (a 

name used in Florida) is a species known by a 

large number of common names worldwide. Other 

names used in U.S. territories are devil’s claw 

(Florida), wild bougainvillea (Texas), escambrón 

(Puerto Rico), and prickly mampoo (U.S. Virgin 

Islands). Pull-back-and-hold is a large, scrambling 

and climbing woody shrub (vine) that rarely 

becomes a small tree. It is reported as a medium to 

large tree in Costa Rica (Enquist and Sullivan 

2001). Its stems and branches are smooth, dark 

gray, and usually support opposite, stout, hooked 

and branched spines. The spines aid in 

clinging to tree crowns. Pull-back-and-hold may 

have one to several branches arising near the base, 

and then the stems are relatively free of branches 

until they reach increased light, usually in the 

canopy of trees where they branch profusely. 

Seedlings have a taproot and finer laterals. In older 

plants the lateral roots become extensive and 

diffuse. The fine roots are tough and fibrous. The 

species may be evergreen or deciduous depending 

on the climate. The leaves are variable, ovate to 

elliptical and 2.5 to 15 cm long by 1.5 to 6 cm 

broad. They are light to dark green, leathery to 

somewhat fleshy. The flowers are small axillary 

cymes that are succeeded a few months later by 11 

to 15 mm long fruits (anthocarps) with five single 

or double length-wise rows of sticky glands 

(Acevedo-Rodríguez 1985, Howard 1988, Liogier 

1985). 

Range.—Pull-back-and-hold is native to most of 

the humid tropical and subtropical areas of the 

world including five continents, most of the 

islands of the Caribbean and many of the Pacific 

islands. Its range includes southern Florida and the 

Rio Grande Valley of Texas (Florida 4-H 

Foundation 2001, Friend 2001, Howard 1988).

Ecology.—Pull-back-and-hold grows from near 

sea level in coastal areas to as much as 1,000 m in 

elevation in continental areas (Múlgura 2000). In 

Puerto Rico, the species grows in sites receiving 

from 750 mm to over 2000 mm of precipitation. 

These shrubs grow well in soils with textures from 

sand to clay, with drainage from somewhat poorly 

drained to excessively well drained in limestone, 

igneous, and alluvial areas. In Florida, the species 

grows on hummocks and pinelands (Florida 4-H 

Foundation 2001). Seedlings and young plants 

survive in broken sunlight, but to reproduce, a 

plant must be relatively large and must have 

reached nearly full sunlight. This usually means 

growing in the crowns of trees or sometimes on 

rocks or fences 

 

Reproduction.—In Puerto Rico, pull-back-andhold flowers from December to May (the dry 

season) and fruits in August (just before the wet 

season) (Acevedo-Rodríguez 1985) although 

fruiting of occasional individuals may occur at 

almost any time of the year. Plants in Costa Rica 

fruit late in the wet season (Enquist and Sullivan 

2001). A large plant can produce thousands of 

seeds during a fruiting episode. Fruits from a 

Puerto Rican collection averaged 0.077 + 0.028 
574 

g/fruit. Seeds extracted by hand, which have the 

appearance of grains of wild rice, averaged 0.041 

+ 0.011 g/seed. It is not necessary to extract seed 

because they germinate satisfactorily within the 

fruit. Seventy-five percent of seeds from the above 

Puerto Rican collection, sown as fruits in 

commercial potting mix, germinated between 21 

and 49 days after sowing. Pull-back-and-hold is 

susceptible to damping-off as seedlings in 

germination beds. Seedlings are ready to prick out 

into containers about 1 month after germination. 

The sticky seeds can cling to birds while on the 

trees; then, after they fall, still attached to the 

infrutescences, stick to the clothing or fur of 

passing humans and animals. 

Growth and Management.—Although pull-backand-hold has only a moderate growth rate, it can 

live for several decades and become quite large. 

Plants may reach 20 m of extension (AcevedoRodríguez 1985) and 20 cm of stem diameter in 

Puerto Rico. It is sometimes desirable to remove 

pull-back-and-hold from forest stands. This 

increases safety during timber harvest and helps 

ensure that the regenerating stand will not be 

suffocated by resurgent vines. A time-honored 

method is to cut the vines hanging from trees 1 

year before harvest and then to cut or spray the 

few surviving sprouts just before harvest. 

Benefits.—Pull-back-and-hold may be used as a 

hedge or a foundation plant in landscaping, but it 

requires regular pruning to keep it under control 

(Friend 2001) An occasional plant occurs entirely 

free of spines. Perhaps these could be propagated 

vegetatively for ornamental use. The wood of wild 

pull-back-and-hold is sometimes used for fuel. It 

is hard, heavy, and often available in good 

diameters and considerable lengths. Root extracts 

have been used in herbal medicine as a purgative, 

and extracts from leaves and bark are used to treat 

arthritis (Acevedo-Rodríguez 1985). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Enquist, B.J. and J.J. Sullivan. 2001. Vegetative 

key and descriptions of tree species of the 

tropical dry forests of upland Sector Santa Rosa, 

Area de Conservación, Guanacaste, Costa Rica. 

http://www.acguanacaste.ac.cr/paginas_ especie/ 

plantae_online/EnquistSullivanTreeKey.pdf. 

68 p.

Florida 4-H Foundation. 2001. Florida forest 

plants: Devil’s claws (Pisonia aculeata). 

http://www.sfrc.ufl.edu/4h/Devils_claw/ 

deviclaw.htm. 2 p. 

Friend, W.H. 2001. Plants of ornamental value for 

the Rio Grande Valley of Texas. http:// 

www.aggiehorticulture.tamu.edu/plantanswers/ 

publications/riograndeornamentals.htm. 16 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier, H.A. 1985. Discriptive flora of Puerto 

Rico and adjacent islands. Vol. 1. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

352 p. 

Múlgura, M.E. 2000. Catálogo de las plantas 

vasculares de la República Argentina. 

http://www.Darwin.edu.ar/Catologo/ 

nyctaginaceae.pdf. 4 p. 
575

Pithecellobium unguis-cati (L.) Benth. bread-and-cheese 

FABACEAE 

Synonyms: Mimosa unguis-cati L. 

 Zygia unguis-cati (L.) Sudw. 

John K. Francis 

General Description.—Bread-and-cheese is the 

common name used in the Virgin Islands and the 

English-speaking Lesser Antilles. The species is 

also know as cat’s claw and black bead in Florida 

and doctor long, groven eye, uña de gato, rolón, 

bois crabbe, griffe-chatte, collier-diable, tendra à 

caulilou-rivière, and bébèl elsewhere (Howard 

1988, Liogier 1988, Little and others 1974). 

Bread-and-cheese is a shrub or small tree usually 3 

to 9 m in height and 4 to 13 cm in stem diameter 

(Little and others 1974). However, it occasionally 

grows to impressive proportions. The national 

record tree, found in Camino Real, Florida, 

measures 22 m in height and 121 cm in diameter 

(Champion Tree Project 2001). The plant usually 

has multiple stems arising at or below ground 

level. The stems and branches are gray and nearly 

smooth with lenticels and rings at the nodes. The 

paired spines are persistent. Bread-and-cheese 

produces a taproot and abundant lateral and fine 

roots. Alternate hairless leaves have petioles 2 to 5 

cm long and leaf blades 2 to 6 cm long. There are 

four obliquely obovate or oblong leaflets per leaf. 

The greenish-yellow, yellow, or pink flowers form 

terminal or axillary racemes or panicles in heads. 

The legume, 5 to 10 cm long, is coiled or curved 

and splits open to reveal 4- to 6-mm shiny, black 

seeds surrounded by white to red fleshy arils 

(Howard 1988, Liogier 1988, Little and others 

1974). Bread-and-cheese is easily confused with P. 

dulce (Roxb.) Benth., which grows as an exotic in 

many Caribbean islands. The latter is a tree with 

somewhat smaller leaves, coarser twigs, larger 

fruits, and less pronounced rings at the nodes. 

Range.—Bread-and-cheese ranges from Florida 

through the Greater and Lesser Antilles, Trinidad 

and Tobago, Bonaire, Curacao, Aruba, Guyana, 

and Venezuela (Liogier 1988, Little and other 

1974). Planting has not been reported outside its 

native range. 

Ecology.—Bread-and-cheese grows in Puerto 

Rico from near sea level to about 450 m in 

elevation on soils derived from both igneous and 

sedimentary rocks (author’s observation). The 

species prefers well-drained soils, but all soil 

textures appear to be tolerated. With few 

exceptions, it grows in areas receiving less than 

1000 mm of annual precipitation. Because of 

reduced competition from trees, the species tends 

to grow on sand dunes, coastal strands and keys, 

and on shallow rocky soils, sometimes forming 

thickets (West and Arnold 1952). It does survive 

and grow in the understory of open, dry forests, 

but maintains thin crowns and seldom flowers in 

these situations. Bread-and-cheese appears to 

tolerate salt spray and salty groundwater. 

Reproduction.—Bread-and-cheese flowers and 

fruits irregularly through the year (Little and 

others 1974). Seed yield can be heavy but 

inconsistent from year to year. A sample of seed 

collected in Puerto Rico contained 14,300 

seeds/kg and germinated at 31 percent, beginning 

3 days after sowing (Francis and Rodríguez 1993). 

Artificial propagation is by seed; it is one of the 

easiest shrubs to grow (Workman 1980). Natural 

seedlings are seldom abundant. The shrubs 

resprout readily after cutting or fire. 
576 

Growth and Management.—Plants started from 

seed grow relatively slowly at first and somewhat 

faster after deep rooting. Supplemental watering 

and fertilization are unnecessary. As a landscaping 

plant, it can be easily pruned and shaped 

(Workman 1980). In the poor habitat where the 

species usually occurs in the wild, growth is very 

slow and plants live for several decades. 

Benefits.—Although it is difficult to harvest 

because of the spines, the wood of bread-andcheese is used for fuel. It is heavy and hard and 

has an oven-dry heat of combustion of 19.05 

megajoules/kg (Timyan 1996). The fleshy aril is 

edible, although not highly desirable because of 

the lingering flavor. The seeds of bread-andcheese are used to make necklaces. The species is 

planted to form impenetrable hedges (UVI 

Wetlands Reserve 2001). Bread and cheese is 

listed as a nitrogen fixing species (Winrock 

International 2001). The fruits are one of the food 

plants of the endangered yellow-shouldered 

amazon parrot (Amazona barbadensis) in 

Venezuela, and the islands of Margarita, La 

Blanquilla, and Bonaire (Island Resource 

Foundation 2001). The large orange sulfur 

(Phoebis agarithe Boisduval) and the Miami blue 

(Hemiargus thomasi Clench) butterflies both use 

bread-and-cheese as rearing plants for their larva 

(Association of Florida Native Nurseries 2001). A 

small treehopper insect (Homoptera) has 

developed an extended thorax that mimics a 

bread-and-cheese thorn and serves as protective 

camouflage (Workman 1980). 

References 

Association of Florida Native Nurseries. 2001. 

Florida gardening with native plants—Legume. 

http://www.nsis.org/garden/family/legume.html. 

4 p. 

The Champion Tree Project. 2001. National 

champion trees, Florida. 

http://www.championtrees.org/database/champi

onsFL.htm. 8 p. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. Southern 

Forest Experiment Station, U.S. Department of 

Agriculture, Forest Service, New Orleans, LA. 

5 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Island Resource Foundation. 2001. Threatened and 

endangered birds of the insular Caribbean: 

Yellow-shouldered amazon, Amazona 

barbadensis. http://www.irf.org/bbarden.htm. 

13 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1024 p. 

Timyan, J. 1996. Bwa yo: important trees of Haiti. 

South-East Consortium for International 

Development, Washington, DC. 418 p. 

UVI Wetlands Reserve. 2001. A list of plants 

tagged along the trail at the Reserve, and some 

of their uses. http://rps.uvi.edu/ 

VIMAS/plantlist.htm. 5 p. 

West, E. and L.E. Arnold. 1952. The native trees 

of Florida. University of Florida Press, 

Gainesville, FL. 212 p. 

Winrock International. 2001. Nitrogen fixing trees 

and shrubs. http.//www.winrock.org 

/forestry/factpub/nftlist /htm. 25 p. 

Workman, R.W. 1980. Growing Native. The 

Sanibel-Captiva Conservation Foundation, Inc. 

137 p. 
577

Pluchea carolinensis (Jacq.) G. Don cure-for-all 

ASTERACEAE 

Synonyms: Conyza symphytofolia sensu Gillis 

 Pluchea odorata authors, non P. odorata (L.) Cass. 

Conyza carolinensis Jacq. 

 Conyza coresii Kunth 

 Pluchea cortesii (Kunth) DC. 

John K. Francis 

General Description.—Cure-for-all, also known 

as sourbush, sweetscent, wild tabacco, cattle 

tongue, salvia, sauge rouge, guerit-tout, tabak 

djab, zówèy mouten, is a shrub 1 to 3 m in height 

and up to 6 cm in basal stem diameter. Stem wood 

is moderately soft and brittle. Older plants are 

supported by many flexible, lateral roots and may 

be sparsely branched to somewhat branchy. The 

twigs are stout. The foliage is concentrated on the 

branch ends. Leaves have petioles 1 to 3 cm long, 

and ovate to elliptic blades 6 to 20 cm long with 

rounded bases, pointed ends, usually entire edges, 

upper surfaces glabrate to densely pubescent, and 

lower surfaces velvet-pubescent. The terminal 

inflorescences are broadly rounded corymbose 

panicles of heads of over 500 flowers each. The 

corollas are usually pink, but vary from light 

purple to white. The achenes are brown, 0.6 to 0.8 

mm long, and have a pappus of 10 to 15 

yellowish-white bristles. Chromosome number is 

2n = 20 (author’s observation, Howard 1989, 

Liogier 1997, Long and Lakela 1976, Stevens and 

others 2001) 

Range.—Cure-for-all is native to Florida, 

Bermuda, the Bahamas, the West Indies, Mexico, 

Central America, and Colombia, Venezuela, and 

Ecuador in South America (Stevens and others 

2001, Natural Resources Conservation Service 

2002). The species has naturalized in Hawaii, 

Guam and other Pacific Islands, Taiwan, West 

Africa, and probably in many other places in the 

tropics (Peng and others 1998, Pacific Islands 

Ecosystems at Risk 2002, Stevens and others 

2001). 

Ecology.—Cure-for-all is adapted to a wide 

variety of soils and sites. It tolerates excessivelywell to poorly-drained soils, the full range of soil 

textures, acid and alkaline reactions, salt and salt 

spray, and compaction. A minimum of about 

1000 mm of mean annual precipitation is required 

in upland sites, but it grows in much drier climates 

along streams and near mangroves and marshes. 

Cure-for-all may be found from near sea level to 

1,000 m in elevation in Hawaii (University of 

Hawaii Botany 2002). The species is intolerant 

and cannot endure overhead shade or severe 

competition from brush or grass. It is common in 

disturbed areas such as construction sites, 

riverbanks, the margins of hammocks, road cuts 

and fill, vacant lots, eroded sites, landslides, 

burned areas, and abandoned fields. In Hawaii, 

cure-for-all quickly invades burned areas, but 

being early successional, is soon replaced by other 

species (Smith and Tunison 1992). Plants sprout 

after fires if they are not too intense (University of 

Hawaii Botany 2002). 

Reproduction.—Cure-for-all blooms in spring 
578 

and summer in Florida (Long and Lakela 1976) 

and produces seeds prolifically. A collection of 

seeds from Puerto Rico weighed an average of 

0.000025 g/seed or 40 million seeds/kg. Because 

the seeds failed to germinate on moist filter paper, 

it is not known whether this represents a 

reasonable estimate for the species (author’s 

observation). The seeds are wind-dispersed and 

probably require wet, bare soil to germinate and 

establish themselves. 

Growth and Management.—In Puerto Rico, 

most cure-for-all live for 2 to 4 years before dying 

or dying back to the root and resprouting. Sprouts 

grow about 1.5 m in the first year. Establishment 

of new plants can probably be assured by 

scarifying the soil before the wet season in the 

presence of a seed source. Where it is necessary to 

eliminate cure-for-all, grubbing out the plants or 

spraying with broadleaf herbicides is 

recommended until tested treatments are available. 

A seed insect, Acinia picturata (Diptera: 

Tephritidae), was introduced in Hawaii and 

although now well established, has had no 

significant effect on the shrub (Alyokhin and 

others 2001). 

Benefits.—Cure-for-all has an excellent ability to 

colonize and stabilize disturbed areas and act as a 

nurse crop for later-successional species. Although 

not specifically reported, as other members of the 

family, it probably provides a source of nectar and 

pollen for honeybees and other insects. It has 

several herbal applications including aromatic 

baths, control of fever, treatment of uterine 

fibroids, relief of sore throat and stomach pain, 

poultices for wounds and skin ulcers, as an 

analgesic, and for the treatment of malaria (Balick 

and others 2000, Liogier 1990, Vélez and van 

Overbeek 1950). Analgesic and antiinflammatory 

effects have been demonstrated in laboratory trials 

with rats (Gavilán-Yodú and Hechavarría 2002). 

References 

Alyokhin, A.V., R.H. Messing, and J.J. Duan. 

2001. Utilization of the exotic weed Pluchea 

odorata (Asteraceae) and related plants by the 

introduced biological control agent Acinia 

picturata (Diptera: Tephritidae) in Hawaii. 

Biocontrol Science and Technology 11: 711-

718. 

Balick, M.J., F. Kronenberg, A.L. Ososki, M. 

Reiff, A. Fugh-Berman, B. O’Connor, M. Roble, 

P. Lohr, and D. Atha. 2000. Medical plants used 

by Latino healers for woman’s health conditions 

in New York City. Economic Botany 54(3): 

344-357. 

Gavilán-Yodú, R. and P.N. Hechavarría. 2002. 

Efecto analgésico y antiinflamatorio de la tintura 

de salvia (Pluchea carolinensis) al 30 % en ratas 

Wistar. Instituto Superior de Ciencias Médicas, 

La Habana, Cuba. http://fcmfajardo.sld.cu/ 

jornada/trabajos/salvia/resumen/htm. 5 p. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyan Books, Miami, FL. 

962 p. 

Natural Resources Conservation Service. 2002. 

Plant profile: Pluchea carolinensis (Jacq.) G. 

Don. http://plants.usda.gov/cgi_bin/plant_profile 

.cgi?symbol=PLCA10. 3 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Pacific Island Ecosystems at Risk. 2002. Invasive 

plant species: Pluchea carolinensis (Jacq.) G. 

Don, Asteraceae. http://hear.org/pier3/plcar.htm. 

2 p. 

Peng, C.-I., C.-H. Chen, W.-P. Leu, and H.-F. 

Yen. 1998. Pluchea Cass. (Asteraceae: Inuleae) 

in Taiwan. Botanical Bulletin of Academia 

Sinica 39(4): 287-297. 

Smith, C.W. and J.T. Tunison. 1992. Fire and alien 

plants in Hawai’i: Research and management for 

native ecosystems. In: C.P. Stone, C.W. Smith, 

and J.T. Tunison, eds. Alien plant invasions in 

native ecosystems of Hawai’i: Management and 

Research. University of Hawaii Cooperative 
579

National Park Resources Studies Unit, 

Honolulu, HI. p. 394-409. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany. Vol. 85, No. 

1. Missouri Botanical Garden Press, St. Louis, 

MO. 943 p. 

University of Hawaii Botany. 2002. Alien plants 

of Hawaii: Pluchea symphytifolia (Mill.) Gillis. 

http://www.botany.hawaii.edu/faculty/cw_smith

/plu_sym.htm. 1 p. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Río Piedras, PR. 497 p. 
580 

Prosopis glandulosa Torr. honey mesquite

FABACEAE 

Synonyms: Prosopis chilensis (Molina) Stuntz 

Prosopis odorata Torr. & Frém. 

Juanita A.R. Ladyman 

Illustration source: USDA-Forest Service collection, 

Hunt Institute 

General Description.—Prosopis glandulosa is 

commonly known as honey mesquite, mesquite, 

glandular mesquite, prairie mesquite, algarobo, 

and la péchita. The Spanish conquistadores named 

it “algaroba” as it reminded them of the algarroba, 

or carob tree of Spain (USDAFS 1988). Mesquite 

are deciduous shrubs, or small trees, armed with 

one or two, rarely absent, straight, stout, stipular 

spines and pinnately twice-compound leaves. 

Flowers are in fragrant, globose heads and the 

fruits are straight, slightly curved, long 

(approximately 20 cm) pods with partitioned and 

embedded seeds (Barneby 1989, Vines 1986). 

Typically the plants are 1- to 2-m high shrubs but 

may develop into 10 m tall trees (Bowers 1993). 

An unusually large individual in Texas was 

reported to be 12.8 m tall with a 15.8 m crown 

spread and 2.4 m girth (Powell 1998). In some 

habitats the plants remain less than 0.5 m tall 

(USDAFS 1988). Three varieties of P. glandulosa

are recognized: var. torreyana, var. prostrata, and 

var. glandulosa (Kartesz 1994). Prosopis 

glandulosa has been known erroneously as P. 

juliflora (Kartesz 1994) that grows in the West 

Indies. Inter- and intraspecific hybridization within 

mesquites is common and identification of the 

intermediate forms may be difficult (Uchytil 

1990). The mesquite introduced into Hawaii is also 

called “algarroba” but is a different species 

(Sohmer and Gustafson 1987). 

Range.—Honey mesquite is a common shrub in 

Oklahoma, southern Kansas, Texas, New Mexico, 

extreme southwestern Utah, Arizona, southeastern 

California, and southern Nevada, and in Nuevo 

Leòn, Tamaulipas, and Coahuila in Mexico 

(Barneby 1989, Powell 1998). 

Ecology.—Honey mesquite grows at elevations 

below 1,800 m in a variety of soils and habitats. It 

is an aggressive plant and will occupy disturbed 

soils. It has a very deep (to 20 m) and wide root 

system that confers drought tolerance (Dayton 

1931, Kearney and others 1960). It grows in areas 

of Death Valley that receive only 5 cm annual 

rainfall (Bainbridge and others 1990). It is 

moderately frost tolerant. It has always had a wide 

geographical range but has been reported as being 

historically restricted to moist, rather narrow 

habitats such as arroyos, floodplains, and canyons 

(Dick-Peddie 1993, Powell 1998). However, 

contrary to these reports mesquite was not totally 

restricted to these more moist habitats but was a 

component of upland communities prior to 

European settlement and may have been overlooked by early explorers as it tended to be a low, 

small shrub obscured in thick grasslands (Smiens 

and others 1992). This breadth of habitat is 

consistent with its hardiness to adverse 

environmental conditions. 

Reproduction.—Honey mesquite flowers in 

March to May, and the fruits mature before the 

summer rains in June through September (Bowers 

1993, Vines 1986). However, depending upon 

environmental conditions, flowering may extend 

into September (Everitt and Drawe 1993, Vines 

1986). The fragrant, creamy-yellow to yellowishgreen flowers are insect pollinated (Everitt and 

Drawe 1993, Uchytil 1990). The fruit is a manyseeded, brown, shiny, dehiscent pod (legume). 
581

Scarified seeds germinate readily and can remain 

undigested and viable as they pass through animal 

digestive tracts (Wright 1982). The germination 

rate typically ranges from 75 to 90 percent (Vines 

1986). Seeds frequently contain weevil larvae 

(Vines 1986). The pollen is reported responsible 

for some hay fever (Kearney and others 1960) 

implying that it is also disseminated by wind. 

Growth and Management.—Since livestock 

grazing was introduced in the Southwestern United 

States and Northern Mexico, honey mesquite has 

increased to such an extent as to be considered a 

pest. However, mesquite is a valuable soil binder 

and slows erosion of otherwise denuded land. The 

increase in honey mesquite has been attributed to a 

variety of factors. The two most often cited are a 

reduction in perennial grasses, which when healthy 

and dense reduces mesquite seedling 

establishment, and the reduced incidence of fire 

due to livestock over-grazing and fire suppression 

(Bahre 1995, Humphrey 1958, Fisher 1977). Other 

reasons include: reduced effectiveness of 

invertebrate seed predators, increased 

dissemination of seed by livestock and/or 

kangaroo rats, land clearing and cultivation, soil 

compaction by livestock that hinders grass 

establishment, decreased activity of jackrabbits 

and wood rats, and climate change (Bahre 1995, 

Glendening 1952). Depending upon the situation, 

herbicide and mechanical treatments are often both 

economical and effective in controlling scattered 

and dense stands of mesquite (Schmutz and others 

1992). Chaining and burning the trees destroy 

above-round tissue, but old stumps of honey 

mesquite readily resprout. Burning is effective 

only on small mesquite plants and where fuel is 

adequate to carry a fire (Schmutz and others 1992). 

Benefits.—Honey mesquite plants bind soils, and 

plantings are recommended for erosion control 

(Graham 1941, Rorabaugh 1995). Mesquite, 

particularly var. torreyana, is tolerant of saline 

soils and is a particularly useful cover in degraded 

areas (Rorabaugh 1995, Schmutz and others 1992). 

It has been used, with limited success, in 

revegetating copper mine wastes (Norem and 

others 1982). Mesquite has been used in riparian 

restoration projects and provides excellent habitat 

for many species of wildlife and birds (Rorabaugh 

1995). In one study, the number of quail using an 

area decreased as a consequence of mesquite 

removal (Goodwin and Hungerford 1977). Honey 

mesquite also provides shelter for the germination 

and development of forbs, grasses, and cacti such 

as Peniocereus greggii Britt. and Rose (Tull 1987, 

author’s personal observation). It is not clear 

whether this is due to a favorable microclimate or 

a response to grazing pressures. Seeds of mesquite 

are nutritionally rich and are important food for a 

large number of wildlife species (Graham 1941, 

Tull 1987). Honey mesquite seeds may constitute 

5 to 37 percent of scaled quails' diets year round 

(Davis and others 1975). Deer, javelina, and 

smaller wildlife species such as jackrabbits, feed 

on both pods and vegetation. It is important 

livestock feed, especially in times of drought. 

However, if livestock eat too large amounts, 

rumen stasis, impaction, and death may result 

(Stubbendieck and others 1993). It is an important 

“honey plant” and bees that forage its flowers 

produce excellent quality honey (Dayton 1931). It 

provides a good source of nectar and food for 

butterfly adults and larvae (Taylor and others 

1997). The wood is resistant to termites and decay, 

and is used for building. It is also prized as bar-bcue fuel as it is a hard, close-grained wood that 

burns hot and slowly (Barneby 1989, CWAR 

2002). Honey mesquite was essential to Native 

Americans who made breads from the pods, fuel 

and utensils from the wood, medicines from the 

stems and leaves, rope from the roots, and dye 

from the black pitch of the trunks (Moerman 1998, 

Tull 1987). The gum exudates of the bark may be 

a substitute for gum arabic and have been used to 

mend pottery (Powell 1998). An intoxicating 

beverage is made from fermented meal (Bowers 

1993). 

References 

Bahre, C.J. 1995. Human impacts on the 

grasslands of southeastern Arizona In: M.P. 

McClaran and T.R. Van Devender, eds. The 

Desert Grassland. The University of Arizona 

Press, Tucson, AZ. p. 230-264. 

Bainbridge, D.A., R.A. Virginia, and W.M. Jarrell. 

1990. Honey Mesquite. Forest, Farm, and 

Community Tree Network (FACT Net) 

http://www.winrock.org/forestry/factnet.htm. 

Barneby, R.C. 1989. Fabales. Intermountain Flora, 

Vascular Plants of the Intermountain West, 

U.S.A. Vol. 3, Part B. New York Botanical 

Garden, Bronx, NY. 279 p. 

Bowers, J.E. 1993. Shrubs and trees of the 

Southwest Deserts. Southwest Parks and 

Monuments Assoc. Tucson, AZ. 140 p. 
582 

Center for Wood Anatomy Research. 2002. U.S. 

Department of Agriculture. Forest Service. 

Tech. Fact Sheet. Prosopis. http://www2.fpl. 

fs.fed.us/TechSheets/HardwoodNA/htmlDocs/ 

prosop1.html. 

Davis, C.A., R.C. Barkley, and W.C. Haussamen. 

1975. Scaled quail foods in southeastern New 

Mexico. Journal of Wildlife Management 

39(3): 496-502. 

Dayton, W.A. 1931. Important western browse 

plants. U.S. Department of Agriculture. Misc. 

Publ. 101. Washington, DC. 214 p. 

Dick-Peddie, W.A. 1993. New Mexico Vegetation 

– past, present, and future. University of New 

Mexico Press, Albuquerque, NM. 244 p. 

Everitt, J.H. and D.L. Drawe. 1993. Trees, Shrubs 

and Cacti of South Texas. Texas Tech 

University Press, Lubbock, TX. 213 p. 

Fisher, CE. 1977. Mesquite and modern man in 

southwestern North America. In: B.B. Simpson, 

ed. Mesquite, its biology in two desert shrub 

ecosystems. Downden, Hutchinson, and Ross, 

Stroudsburg, PA. p 17-88. 

Glendening, G.E. 1952. Some quantitative data on 

the increase of mesquite and cactus on a desert 

grassland range in southern Arizona. Ecology. 

33 (3): 319-328. 

Goodwin, J.G. and C.R. Hungerford. 1977. Habitat 

use by native Gambel’s and scaled quail and 

released masked bobwhite quail in southern 

Arizona. Research Paper RM-197. USDA 

Forest Service, Rocky Mountain Research 

Station, Fort Collins, CO. 8 p. 

Graham, E.H. 1941. Legumes for erosion control 

and wildlife. U.S. Department of Agriculture 

Misc. Pub. 412. Washington D.C. 153 p. 

Humphrey, R.R. 1958. The desert grassland: a 

history of vegetational changes and an analysis 

of causes. Botanical Review 24: 193-252. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada and 

Greenland. Vol.1 - Checklist. 2nd ed. Timber 

Press, Portland, OR. 622 p. 

Kearney, T.H., R.H. Peebles, and collaborators. 

1960. Arizona flora. 2nd ed. University of 

California Press, Berkeley, CA. 1085 p. 

Moerman, D.E. 1998. Native American Ethnobotany. Timber Press, Portland, OR. 927 p. 

Norem, M.A., A.D. Day, and KL. Ludeke. 1982. 

An evaluation of shrub and tree species used in 

revegetating copper mine wastes in the United 

States. Journal of Arid Environments 5: 299-

304. 

Powell, A.M. 1998. Trees and shrubs of TransPecos and adjacent areas. University of Texas 

Press, Austin, TX. 498 p. 

Rorabaugh, J.C. 1995. A superior accession of 

western honey mesquite (var. torreyana) for 

riparian restoration projects. Desert Plants. 

11(4) 32-40 

Schmutz, E.M., E.L. Smith, P.R. Ogden, M.L. 

Cox, J.O. Klemmedson, J.J. Norris, and L.C. 

Fierro. 1992. Desert Grassland. In: R.T. 

Coupland ed. Ecosystems of the World. Vol. 

8A. Elsevier, London and New York. 469 p. 

Smiens, F.E., D.D. Diamond, and C.W. Hanselka. 

1992. Coastal Prairie. In: R.T. Coupland, ed. 

Ecosystems of the World. Vol. 8AElsevier, 

London and New York. 469 p. 

Sohmer, S.H. and R. Gustafson. 1987. Plants and 

flowers of Hawaii. University of Hawaii Press, 

Honolulu, HI. 160 p. 

Stubbendieck, J., S.L. Hatch, and C.H. Butterfield. 

1993. North American Range Plants. 4th ed. 

Univ. of Nebraska Press, Lincoln, NB. 493 p. 

Taylor, R.B., J. Zrutledge, and J.G. Herrera. 1997. 

A field guide to common south Texas shrubs. 

Texas Parks and Wildlife Press, Austin, TX. 

p. 106. 

Tull, D. 1987. Edible and Useful Plants of Texas 

and the Southwest. University of Texas Press. 

Austin, TX. 518 p. 

Uchytil, R.J. 1990. Prosopis glandulosa var. 

glandulosa. U.S.D.A. Forest Service, Rocky 

Mountain Research Station, Fire Sciences 

Laboratory Fire Effects Information System, 

http:// www.fs.fed. us/databse/feis/. 
583

United States Department of Agriculture Forest 

Service. 1988. Range Plant Handbook. Dover 

Publications, Inc. New York. 837 p. 

Vines, R.A. 1986. Trees, shrubs, and woody vines 

of the Southwest. University of Texas Press. 

Austin, TX. 1,104 p. 

Wright, R.A. 1982. Aspects of desertification in 

Prosopis dunelands of southern New Mexico. 

Journal of Arid Environments 5: 277-284. 
584 

Proustia vanillosma C. Wright sweet yellowcrown 

ASTERACEAE 

Synonyms: Proustia krugiana Urban 

 Perezia vanillosma Molinet & Gómez 

John K. Francis 

General Description.—Sweet yellowcrown, a 

name assigned by the Natural Resources 

Conservation Service (2003), is a clambering 

woody shrub reaching 1 to 3 m in height and 5 m 

of extension. The species usually has multiple 

stems that sometimes reach 2 cm in diameter. 

They are covered with a lightly striated, graybrown bark and a green inner bark, and have a 

brittle, greenish-white wood with a 2-mm pith. 

The stems layer (root) whenever they come in 

contact with the soil. The roots are brown and 

flexible. Wood of branches is flexible, strong, and 

hard. The alternate, light-green leaves are thin to 

leathery with a rough surface, elliptic to narrowly 

ovate, pointed to rounded at the tips, have margins 

rough to dentate, and are up to 9 cm long. The 

fragrant inflorescences are leafy, terminal panicles 

of heads that have three to seven florets. The 

corolla is yellow, 8 to 10 mm long. The seeds have 

an 8-mm straw-colored pappus (Liogier 1997). 

Turner (1993) proposed a new genus, 

Berylsimpsonia, to replace Proustia in the Greater 

Antilles. 

Range.—Sweet yellowcrown is native to Puerto 

Rico, Hispaniola, and Cuba (Liogier 1997). It is 

not known to have been planted or naturalized 

elsewhere. 

Ecology.—The species occurs in dry and moist 

areas of Puerto Rico that receive from about 750 to 

1800 mm of mean annual precipitation from a few 

meters above sea level to elevations of about 550 

m. Sweet yellowcrown grows on a wide variety of 

well-drained soils derived from sedimentary 

(including limestone), igneous, and metamorphic 

(including ultramaphic) rocks. It is moderately 

tolerant to shade, growing under open to 

moderately dense forest stands, as well as in 

openings and at forest edges. The species is most 

common in remnant primary stands and secondary 

forests that have never been completely converted 

to agriculture. Sweet yellowcrown slowly invades 

abandoned fields and pastures after they become 

forested. Although occasionally common and 

forming small thickets, plants are usually scattered 

(author’s observations). 

Reproduction.—Sweet yellowcrown in Puerto 

Rico flowers near the end of the wet season 

(November and December) and matures seed at the 

beginning of the dry season (January and 

February). A group of seeds collected in Puerto 

Rico averaged 0.000654 g/seed or 1,529 seeds/g. 

Sown on moist blotter paper, 88 percent of the 

seeds germinated within 13 days. Germination is 

epigeal. Seeds are dispersed by wind. Seedlings 

occur infrequently in forests. Once established, 

plants spread locally by layering (author’s 

observations). 

Growth and Management.—Sweet yellowcrown 

has a moderate growth rate (about 0.5 m of stem 

extension/year) and can live for at least a decade. 

Apparently plants (clones) survive longer than 

individual stems. No planting or natural stand 

management experience is published or known to 

the author. 

Benefits.—Sweet yellowcrown helps protect the 

soil, furnishes cover for wildlife, and adds to the 

aesthetics of the forest. 

References

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 
585

Universidad de Puerto Rico, San Juan, PR. 

436 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Proustia vanillosma C. Wright, 

sweet yellowcrown. http://plants.usda.gov/ 

cgi_bin/plant_profile.cgi?symbol=PRVA. [not 

paged]. 

Turner, B.L. 1993. Berylsimpsonia (Compositae: 

Mutiseae), a new genus for the Greater Antilles. 

Phytologia 74: 349-355. 
586 

Prunus americana Marsh. American plum 

ROSACEAE 

Synonyms: Prunus americana Marsh. var. floridana Sarg. 

 Prunus domestica L. var. americana (Marsh.) Castigl. 

 

John K. Francis 

General Description.—American plum, also 

known as Pottawattami plum, wild plum, yellow 

plum, red plum, river plum, goose plum, hog 

plum, and ciruela, is a shrub or small gnarled tree 

generally 1 to 8 m in height (rarely to 11 m) with 

stem diameters up to 30 cm. Maximum size 

depends on the environment. It usually produces a 

single stem and branches low to the ground but 

forms thickets by suckering from the roots. Stem 

bark is about 1.25 cm thick, gray or dark brown 

tinged with red, with the outer layer separating 

into thin, persistent plates. The twigs are green at 

first, becoming orange-brown and darker as they 

age. The plants become branchy and somewhat 

thorny, especially in older individuals. The wood 

is hard, heavy, and close-grained, with reddishbrown heartwood and light tan sapwood. Leaves 

are dark green, elliptic to obovate, 4 to 10 cm long, 

with an elongated point at the tip and a rounded 

base, margin serrate or doubly serrate, and having 

usually glandless petioles 1.5 to 2 cm long. 

Flowers are white, five-petaled, about 2.5 cm 

across, growing in two- to five-flowered lateral 

umbels. They are unpleasantly aromatic. Fruits 

(globose to oblong drupes) are 1.8 to 2.5 cm long 

and red to yellow (usually orange-red) when ripe. 

The fruits have a thick skin and sweet-tart, yellow 

flesh. The brown stone forms a flattened oval, 

ridged on one edge and grooved on the other 

(Britton and Shafer 1908, Sargent 1923). 

Range.—American plum is native to southeastern 

Canada and the conterminous states of the United 

States except Texas, Washington, Idaho, Oregon, 

Nevada, and California where it is planted and 

probably naturalized (Natural Resources 

Conservation Service 2003, Sargent 1923). It has 

been suggested that the current broad range is, in 

part, the result of planting by Native Americans 

and white settlers (Treeguide 2003). The species is 

widely planted in temperate areas of the World and 

probably has naturalized in some of them. 

Ecology.—Across its vast range, American plum 

grows in a wide variety of sites. In the eastern and 

mid portions of the continent, it grows on 

roadsides, old fields, vacant lots, fencerows, 

gullies, clearings, edges of woodlots, and open 

prairies. In the West, it grows in riparian areas and 

ditch banks, at the edges of fields, and in moist 

foothills. American plum grows at elevations up to 

2,300 m in Utah. The species grows on sand 

through clay soils on moist to somewhat dry sites. 

It tolerates a moderate amount of salt in the soil 

(Forest Service 2003). American plum is intolerant 

of shade and will not grow in the understory of 

forest or persist if overtopped by trees (Treeguide 

2003). Other members of the genus and 

presumably American plum recover rapidly from 

fires by sprouting (Forest Service 2003). 

Reproduction.—Flowering occurs from March 

through May and fruit ripens from late June 

through October, depending on habitat (Forest 

Service 2003, Oklahoma Biological Survey 1999). 

The flowers are pollinated principally by 

honeybees (Apis mellifera L.). Good fruit and seed 

crops are borne every 1 to 2 years (Grisez and 

others 2003). Fresh fruits collected by the author 

in Utah averaged 6.572 + 0.139 g/fruit. Air-dried 

seeds separated from them averaged 0.5336 +

0.0095 g/seed or 1,874 seeds/kg. Grisez (1974) 
587

reported 1,918 cleaned seeds/kg, and that 60 

percent germinated after cold stratification. 

Germination is hypogeal (Young and Young 

1992). Seeds are dispersed by mammals and birds. 

Animals that ingest the seeds, such as black bear 

(Ursus americanus), are more effective dispersers 

than those that do not (Forest Service 2003). Seeds 

ingested by coyotes (Canis latrans) had a 

significantly lower germination rate than uneaten 

seeds (Cypher and Cypher 1999). Seeds may 

remain in the soil seed bank for many years until 

disturbance creates conditions for growth 

(Treeguide 2003). After dispersed plants become 

well established, they begin forming clonal 

thickets. Suckers may appear as much as 3 m away 

from the parent plants (Colorado Springs Utilities 

2003). 

Growth and Management.—American plum is 

only capable of a moderate growth rate, usually 

less than 30 cm in height per year (Michigan State 

University Extension 2003). Minimum seedbearing age is 4 years (Grisez and others 2003). 

Individual stems rarely live longer than 20 years 

(Treeguide 2003). However, clones may last much 

longer. American plum fruits should be collected 

when fully mature for best quality seed. This can 

be done by hand stripping or shaking or beating 

onto a tarp spread under the shrub or tree. The 

seeds should be cleaned of all pulp by macerating 

and washing. The seeds are usually 96 to 100 

percent filled. Seeds of this species can be stored 

at room temperature for up to 30 months without 

loss of viability. Seeds to be stored for longer 

should be surface dried and placed in a sealed 

container at 1 to 5 °C. Stratification of 90 to 150 

days at 2 to 5 °C before planting is recommended 

for after-ripening of the seed. Alternately, seed 

may be sown in nursery beds 2.5 to 5 cm deep in 

the fall. Seedlings are ready for lifting (bare-root) 

at the end of one growing season (Grisez and 

others 2003). Plantings may be made with bareroot stock and potted nursery seedlings. Direct 

seeding in the fall with unstratified seed or spring 

with stratified seed in prepared seedspots will 

normally yield new seedlings. The species can be 

grafted or used as root stock, and it has been 

successfully propagated from stem cuttings (Forest 

Service 2003). 

Benefits.—American plum is an early spring 

bloomer and with its white flowers beautifies the 

forests and prairies for a week or more each year. 

The species also helps protect the soil and provides 

benefits to wildlife and humans. The browse value 

of the plants to domestic ruminants and wild game 

animals varies from good to poor depending on the 

animal species and the location (Forest Service 

2003). The cover provided by American plum 

thickets is important to many wild animals. The 

fruits are eaten by a number of species including 

bluejays (Cyanocitta cristata), brown thrashers 

(Toxostoma rufum), mockingbirds (Mimus 

polyglottos), red-headed woodpeckers 

(Melanerpes erythrocephalus), bobwhite quail 

(Colinus virginianus), white-tailed deer 

(Odocoileus virginianus), raccoons (Procyon 

lotor), squirrels (Sciurus spp.), and coyotes (Canis 

latrans) (Cowley 2003, Cypher and Cypher 1999, 

Kaiser 2001). The flowers furnish nectar food for 

great purple hairstreak [Atlides halesus (Cramer)] 

and Sweadner’s Jupiter hairstreak [Callophrys 

gryreus (Hübner)] butterflies (Cowley 2003). 

Fruits are eaten raw or cooked, made into jams and 

jellies, and were dried for winter food in former 

times by Native Americans and pioneers. Several 

horticultural varieties of plum have been derived 

from American plum. It has also been used in 

hybrid crosses with other species of plums 

(Bircher and Bircher 2000). The wood, which has 

a specific gravity of 0.73 (Britton and Shafer 

1908), makes excellent firewood except for being 

crooked and limby. American plum is used in 

amenity planting for wildlife and in revegetation 

projects. 

References 

Bircher, A.B. and W.H. Bircher. 2000. 

Encyclopedia of fruit trees and edible flowering 

plants in Egypt and the Subtropics. The 

American University in Cairo Press, Cairo, 

Egypt. 568 p. 

Britton, N.L. and J.A. Shafer. 1908. North 

American trees. Henry Holt and Company, New 

York. 964 p. 

Colorado Springs Utilities. 2003. Plant details. 

http://www.csu.org.cgi-bin/xeri/Xeriinclude? 

Xeridetail?PIS-pra. 2 p. 

Cowley, M. 2003. Rose family (Rosaceae). 

http://www.nsis.org/garden/family/rose.html. 

4 p. 

Cypher, B.L. and E.A. Cypher. 1999. Germination 

rates of tree seeds ingested by coyotes and 

raccoons. American Midland Naturalist 142(1): 

71-76. 
588 

Forest Service. 2003. Fire effects information 

system: species: Prunus americana. http://fs. 

fed.us/database/feis/plants/tree/pruame/all.html. 

14 p. 

Grisez, T.J. 1974. Prunus L. cherry, peach, and 

plum. In: C.S. Schopmeyer, tech. coord. Seeds 

of woody plants in the United States. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. p. 658-673. 

Grisez, T.J., J.R. Barbour, and R.P. Karrfalt. 2003. 

Prunus L. cherry, peach, and plum. In: F.T. 

Bonner and R.G. Nisley, eds. Woody plant seed 

manual. http://wpsm.net/Prunus.pdf. [not 

paged]. 

Kaiser, J. 2001. 2001 featured plant, American 

plum, Prunus americana. In: Plants for 

conservation, Vol. 4, No. 1. Elsberry Plant 

Materials Center, Elsberry, MO. p. 1. 

Michigan State University Extension. 2003. 

Prunus americana—American plum. http:// 

www.msue.msu.edu/msue/imp/modzz/00002025

.html. 1 p. 

Natural Resources Conservation Service. 2003. 

Prunus americana Marsh. http://plants.usda. 

gov/cgi_bin/plant_profile.chi?symbol=PRAM. 

4 p. 

Oklahoma Biological Survey. 1999. Prunus 

americana Marsh. http://www.biosurvey.ou. 

edu/shrub/prun-ame.htm. 2 p. 

Sargent, C.S. 1923. Manual of the trees of North 

America (exclusive of Mexico). Houghton 

Mifflin, Boston, MA. 910 p. 

Treeguide. 2003. American plum, Prunus 

americana Marsh. http://www.treeguide.com/ 

Species.asp?SpeciesID=770. 3 p. 

Young, J.A. and C.G. Young. 1992. Seeds of 

woody plants in North America. Dioscorides 

Press, Portland, OR. 407 p. 
589

Prunus emarginata (Dougl. ex Hook.) D. Dietr. bitter cherry 

ROSACEAE 

Synonyms: None 

Christopher Ross 

 

General Description.—Bitter cherry ranges from 

a shrub to a tree of 10 m. It often forms dense 

thickets. Clustered, deciduous elliptic to obovate, 

crenate-serrate leaves 20 to 65 mm in length are on 

3 to 12 mm petioles (Wilken 1993, Mozingo 

1987). The leaves, stems, and flowers all have a 

very strong cyanide odor (bitter almond) when 

crushed. 

Range.—Bitter cherry is found in cooler parts of 

Arizona, California, far western Nevada, Idaho, 

Utah, New Mexico and Montana and north to 

British Columbia (Mozingo 1987, Esser 1995). 

Ecology.—Bitter cherry site dominance may not 

peak until three decades after disturbance, 

suggesting that it is not an obligate seed bank 

species (Oakley and Franklin 2001). However, in 

some cases following fire it may quickly dominate 

a site (personal. observation.). However, others 

(Morgan and Neuenschwander 1988) have 

considered it an obligate seed bank species. It may 

achieve major dominance of disturbed sites, which 

persists for a long time but does not occur until 

late in the post-disturbance seral progression. 

Seed dispersal is by animals (Halperin 1989, 

personal observation). 

Reproduction.—Three to 12 five-petaled flowers 

on 3- to 12-mm pedicels occur in sometimes flattopped racemes. The sepals and hypanthium are 

glabrous to puberulent. White petals are 4 to 8 mm 

long. Fruits are obvoid to round about 7 to 14 mm 

long, smooth, with a red to purple fleshy pulp 

(Wilken 1993). Seeds may at times undergo heavy 

predation and mortality while green, apparently 

from sawflies of the genus Hoplocampa (Weaks 

2001). Rodents, including chipmunks, eat the 

seeds (John Francis and Maurice Beck, personal 

communication). 

Fire Effects.—Although bitter cherry is a prolific 

sprouter after fire, Leege (1979) found that 

repeated fires eventually result in heavy plant 

mortality. However, in those cases, seedlings 

developed to replace mature plants lost to fire. 

Growth and Management.—Bitter cherry has 

been effectively controlled on conifer planting 

sites by application of the herbicide 2,4,5-T (now 

banned in the United States) (Bock and others 

1978). It may be heavily attacked by tent 

caterpillars and other insects at times. 

Benefits.—Blue grouse and other birds are fond of 

the fruits of bitter cherry. They are also a preferred 

summer and fall food of black bears (Unsworth 

and others 1989). Leaves are browsed by sheep 

and cattle, despite their cyanogenic properties, 

which have caused reported stock poisonings 

(Mozingo 1987). In the Pacific Northwest, bitter 

cherry is eaten by deer and elk (Crouch 1968), 

although it is not a preferred forage (Leege 1979, 

Klebenow 1965). Native Americans used long 

fibrous strips of bark for twine and basketry, and 

used the roots for a variety of medicinal purposes 

but made little use of the bitter fruit (Turner and 

Bell 1973). Bitter cherry essence is marked by 

herbalists, who claim that it aids patience and 

short attention spans. Bitter cherry has also been 
590 

used for mine and highway reclamation (Everett 

and others 1980 and personal observation). 

References 

Bock, J.H., M. Raphael, and C.E. Bock. 1978. A 

comparison of planting and natural succession 

after a forest fire in the northern Sierra Nevada. 

Journal of Applied Ecology 15:597-602. 

Crouch, G.L. 1968. Forage availability in relation 

to browsing of Douglas-fir seedlings by blacktailed deer. Journal of Wildlife Management 32: 

542-553. 

Esser, L.L. 1995. U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research 

Station, Fire Sciences Laboratory (2001, 

October). Fire Effects Information System, 

http://www.fs.fed.ss/database/feis/. [not paged]. 

Everett, R.L., R.O. Meeuwig, and R. I. Butterfield. 

1980. Revegetation of untreated acid spoils at 

Leviathan mine, Alpine County, California. 

California Geology 32: 8-10. 

Halperin, C.B. 1989. Early successional patterns of 

forest species: Interactions of life history traits 

and disturbance. Ecology 70: 704-720. 

Klebenow, D.A. 1965. A montane forest winter 

deer habitat in western Montana. Journal of 

Wildlife Management 29: 27-33. 

Leege, T.A. 1979. Effects of repeated prescribed 

burns on Northern Idaho elk browse. Northwest 

Science 53:107-113. 

Morgan, P. and L.F. Neuenschwander. 1988. 

Seed-bank contributions to regeneration of shrub 

species after clear-cutting and burning. 

Canadian Journal of Botany 66:169-72. 

Mozingo, H. N. 1987. Shrubs of the Great Basin. 

University of Nevada Press, Reno Nevada. 342 

p. 

Oakley, B.B. and J. F. Franklin. 2001. Bitter 

cherry (Prunus emarginata) distribution, 

successional dynamics, and implications for the 

role of the seed bank. 

http://www.cisti.nrc.ca/cisti/journals/cjb/b98-

162.html. p. 1725-1732. 

Turner, N.C. and M.A.M. Bell. 1973. The 

Ethnobotany of the Southern Kwakiutl Indians 

of British Columbia. Economic Botany 27: 257-

310. 

Unsworth, J.W., J.J. Beecham, and L.R. Irby. 

1989. Female black bear habitat use in Westcentral Idaho. Journal of Wildlife Management 

53: 668-673. 

Weaks, A. 2001. Bitter cherry sawfly. 

http://tardigrade.org/natives/cherrysawfly.html. 

3 p. 

Wilken, D.H. 1993. Prunus In: Hickman, J.C., ed. 

1993. The Jepson Manual: higher plants of 

California. University of California Press, 

Berkeley and Los Angeles. 1,400 p. 
591

Prunus pumila L. sand cherry 

ROSACEAE 

Synonyms: Cerasus pumila (L.) Michx. 

 Prunus susquehanae Willd. 

 Prunus cuneata Raf. 

 Prunus depressa Pursh. 

 Prunus besseyi Bailey. 

Don C. Bragg 

 var. pumila 

 var. cuneata

Illustration source: (used with permission) Cranbrook 

Institute of Science 

General Description.—Sand cherry, also called 

beach plum, “cerise de sable” (Fernald 1923), or 

dwarf American cherry, is a diffusely-branched, 

low growing (0.5 to 3 m tall, depending on variety 

and habitat) to sometimes decumbent or prostrate 

shrub (Fernald 1923, Gleason 1952, LamsonScribner 1891). Older stems develop a grayish, 

glabrous bark, while younger twigs are often 

tannish- to reddish-brown or brown. Its alternate 

leaves have glandular petioles up to half the blade 

length. While Fernald (1923) listed three different 

species of sand cherry (P. pumila, P. susquehanae, 

and P. depressa), Gleason (1952) recognized four 

varieties of P. pumila:P. pumila L. var. pumila, the 

Great Lakes sand cherry, has narrow (10 to 18 mm 

wide) oblanceolate leaves, narrowly cuneate at 

their base, acute or acuminate at their apex, 

lustrous on their top surface and pale below. Stems 

are erect and diffusely branched, with some 

decumbent branches on active dunes.

 P. pumila L. var. cuneata (Raf.) Bailey, 

the Appalachian sand cherry, has oblong to 

oblong-obovate leaves with acute bases, often 20 

to 30 mm wide at maturity, pale green above and 

glaucous below. Stems are erect or diffusely 

branched. This variety is most common on dry or 

rocky sites. 

 P. pumila L. var. depressa (Pursh) 

Gleason, the flat sand cherry, has leaves that are 

narrow (10 to 20 mm wide), oblanceolate, often 

with obtuse, long-tapering bases, pale green above 

and whitish below. Stems are prostrate, forming 

low mats up to 2 m wide. New shoots are reddish, 

highly lustrous, and often freely rooted. 

 P. pumila L. var. besseyi (Bailey) 

Gleason, the western sand cherry, has leaves that 

are somewhat glaucous underneath, oblanceolate, 

up to 18 mm wide, with acute to acuminate tips 

and long-cuneate bases. 

Range.—Sand cherry is widely distributed in the 

northern half of the United States and eastern 

Canada, from New Brunswick down the Atlantic 

Seaboard to North Carolina, westward to Utah, 

Wyoming, and Montana, and as far north as 

northern Ontario and Quebec (BONAP 1999, 

Cusik 1985). The varieties of sand cherry are 

generally geographically distinct (BONAP 1999, 

Gleason 1952). Prunus pumila var. pumila is most 

common in the Great Lakes region, especially in 

Michigan, Minnesota, and Wisconsin. Prunus 

pumila var. depressa has been reported from New 

Brunswick to Pennsylvania and New Jersey, with 

isolated populations in Wisconsin, Tennessee, and 

possibly Kentucky. Prunus pumila var. cuneata

ranges from Maine to Minnesota (and possibly 

Arkansas), southeast to Indiana and North 

Carolina. Prunus pumila var. besseyi is primarily a 

western subspecies, extending from Minnesota to 

Kansas westward to Utah and Montana, with 

isolated subpopulations noted as far east as 
592 

Michigan. The widespread distribution of sand 

cherry has helped to conserve the species, although 

it is considered locally threatened by some States 

(e.g., Arkansas, Ohio) because of the rarity of the 

sometimes specialized habitats it occupies 

(Arkansas Department of Planning 1974, Cusik 

1985, Emmitt and Cusik 1983). 

Ecology.—As its name suggests, sand cherry is 

abundant in sandy areas, although var. cuneata is 

most common on rocky sites (Billington 1943, 

Gleason 1952). In the Great Lakes region, sand 

cherry is often found on deep, excessively drained 

glacial sand plains or sand dunes bordering major 

bodies of water (Cowles 1899, McAtee 1920, 

Walp 1935). Sand cherry also grows along gravel 

bars or shorelines, cliff faces, rocky slopes, or 

even on calcareous, saline, or serpentine soils 

(Fernald 1923, Gleason 1952). The habitat 

preferences of sand cherry have restricted its 

abundance in many parts of its range. For example, 

the sand cherry in Arkansas appears to have been 

limited to two locations on remnants of the Grand 

Prairie in Prairie County (Smith 1988). Sand 

cherry is opportunistic in its distribution, 

frequently colonizing road cuts, gravel pits, or 

railroad beds (Cusik 1985, Fernald 1923, Stevens 

1961). Preferred habitat is typically open, with few 

trees, other shrubs, or herbs to compete for light, 

nutrients, and water on the harsh sites it occupies. 

However, some have reported sand cherry as 

abundant in closed forests (Gysel 1966). Human 

activities and habitat alteration from changing 

natural disturbance regimes have been blamed for 

localized extinctions of sand cherry (Arkansas 

Department of Planning 1974, Cusick 1985, 

Drayton and Primack 1996). 

Reproduction.—Depending on geographic 

location, from April to June a sand cherry may 

produce two to four white insect-pollinated 

flowers in umbels scattered amongst its leaves. 

Voss (1954) reported that butterflies pollinated 

sand cherry flowers in northern Michigan. Sand 

cherry fruits ripen by late July or August and are 

typically reddish- or purplish-black to nearly 

black, without bloom, subglobose to globose, and 

10 to 15 mm in diameter (Billington 1943, Fernald 

1950, Gleason 1952). Prunus pumila var. pumila is 

virtually inedible for humans (Rehder 1958), 

though Fernald (1923, 1950) mentions the 

palatability of var. cuneata and var. besseyi. Given 

their size, fleshy and edible fruit, and large stone, 

seed dispersal is primarily through birds and small 

mammals. Sand cherry can also vegetatively 

propagate (Olson 1958). 

Growth and Management.—Sand cherry growth 

is best under open canopy conditions. As with 

many other species that specialize on poor sites, 

rates of growth are usually better on higher quality 

locations (i.e., those with abundant moisture and 

nutrients). However, these conditions also promote 

the growth of competitors that can exclude this 

low-growing, shade-intolerant species. Sand 

cherry’s inconspicuous stature and spreading root 

system leave it vulnerable to overshading, 

trampling, soil compaction, erosion, and other 

surface disturbances (Cusick 1985, Emmitt and 

Cusick 1983). Road grading, for example, may 

have destroyed the two known pockets of sand 

cherry in Arkansas (Arkansas Department of 

Planning 1974). Habitat protection coupled with 

the restoration of openings should help conserve 

this species. 

Benefits.—The small stature of sand cherry has 

minimized its economic benefits, as it does not 

produce merchantable wood and fruit production is 

generally limited. At least one cultivated hybrid is 

commercially available. Popular for its colorful 

foliage, the purpleleaf sand cherry [P. x cistena

(N.E. Hansen) Koehne] is a cross between P. 

pumila and P. cerasifera Ehrh. Fernald (1923) 

fondly described the fruit of sand cherry along the 

rivers of New England and southeastern Canada: 

“...its juicy black ‘plums’ are highly prized either 

raw, cooked or as the source of a rich syrup-like 

jelly,” suggesting that specialty food markets may 

be possible. While the ecological benefits of sand 

cherry are poorly understood, the vanishing 

habitats occupied by this species often have 

considerable value. For example, var. cuneata was 

one of the primary floristic components of the 

Albany Pine Bush in northern New York, which is 

an important local refuge for numerous 

amphibians and reptiles (Stewart and Rossi 1981). 

Sand cherry may also play a critical pioneering 

role in the ecosystems where it is abundant. Its 

deep root network and dense thickets help to 

stabilize shifting sand, allowing for the invasion of 

other plant species and colonization by important 

invertebrates such as ants (Olson 1958, Talbot 

1934). Soil stabilization and organic matter 

production by sand cherry also contribute to 

nitrogen biogeochemistry in dune habitats 

(Robertson and Vitousek 1981). 
593

References

Arkansas Department of Planning. 1974. Arkansas 

natural areas plan. Arkansas Department of 

Planning, Little Rock, AR. 248 p. 

Billington, C. 1943. Shrubs of Michigan. Bulletin 

20. Cranbrook Institute of Science, Bloomfield 

Hills, MI. 249 p. 

BONAP. 1999. Synthesis of the North American 

flora, version 1.0 (CD). North Carolina 

Botanical Garden, University of North 

Carolina, Chapel Hill, NC. 

Cowles, H.C. 1899. The ecological relations of the 

vegetation on the sand dunes of Lake 

Michigan. Botanical Gazette 27: 167-202. 

Cusick, A.W. 1985. Prunus pumila var. pumila L. 

(Great Lakes Sand Cherry). Abstract. Ohio 

Dept. of Natural Resources Division of Natural 

Areas and Preserves. http://www.dnr. 

state.oh.us/ODNR/dnap/Abstracts/P/prupumi. 

htm. 2 p. 

Drayton, B. and R.B. Primack. 1996. Plant species 

lost in an isolated conservation area in 

metropolitan Boston from 1894 to 1993. 

Conservation Biology 10: 30-39. 

Emmitt, D.P. and A.W. Cusick. 1983. Prunus 

pumila var. cuneata Willd. (Sand Cherry). 

Abstract. Ohio Dept. of Natural Resources 

Division of Natural Areas and Preserves. 

http://www.dnr.state.oh.us/ODNR/dnap/ 

Abstracts/P/prupumi2.htm. 2 p. 

Fernald, M.L. 1923. The identities of the sand 

cherries of eastern America. Rhodora 25: 69-

74. 

Fernald, M.L. 1950. Gray’s manual of botany, 8th

ed. American Book Co., New York. 1,632 p. 

Gleason, H.A. 1952. Illustrated flora of the 

Northeastern United States and adjacent 

Canada, Vol. 2. Lancaster Press, Inc., 

Lancaster, PA. 655 p. 

Gysel, L.W. 1966. Ecology of a red pine (Pinus 

resinosa) plantation in Michigan. Ecology 47: 

465-472. 

Lamson-Scribner, F. 1891. A sketch of the flora of 

Orono, ME. Botanical Gazette 16: 228-234. 

McAtee, W.L. 1920. Notes on the jack pine plains 

of Michigan. Bulletin of the Torrey Botanical 

Club 47: 187-190. 

Olson, J.S. 1958. Rates of succession and soil 

changes on southern Lake Michigan sand 

dunes. Botanical Gazette 119: 125-170. 

Rehder, A. 1958. Manual of cultivated trees and 

shrubs, 2nd ed. The Macmillan Co., New York. 

996 p. 

Robertson, G.P. and P.M. Vitousek. 1981. 

Nitrification potentials in primary and 

secondary succession. Ecology 62: 376-386. 

Smith, E.B. 1988. An atlas and annotated list of 

the vascular plants of Arkansas, 2nd ed. 

Department of Botany and Microbiology, 

University of Arkansas, Fayetteville, AR. 

489 p. 

Stevens, O.A. 1961. Plants of Fargo, North 

Dakota. American Midland Naturalist 66: 171-

177. 

Stewart, M.M. and J. Rossi. 1981. The Albany 

Pine Bush: a northern outpost for southern 

species of amphibians and reptiles in New 

York. American Midland Naturalist 106: 282-

292. 

Talbot, M. 1934. Distribution of ant species in the 

Chicago region with reference to ecological 

factors and physiological toleration. Ecology 

15: 416-439. 

Voss, E.G. 1954. The butterflies of Emmet and 

Cheboygan Counties, Michigan with other 

notes on northern Michigan butterflies. 

American Midland Naturalist 51: 87-104. 

Walp, R.L. 1935. Shrubs of Cheboygan and 

Emmet Counties, Michigan. American 

Midland Naturalist 16: 230-247. 
594 

Prunus virginiana L. chokecherry 

ROSACEAE 

Synonyms: Cerasus demissa Nutt. 

 Cerasus virginiana L. 

 Padus demissa Nutt. 

 Padus melanocarpa A. Nelson 

 Padus virginiana L. 

 Prunus demissa Nutt. 

 

Bruce L. Welch 

General Description.—Chokecherry is a 

thornless, deciduous shrub or small tree that grows 

to 8 m tall (Cronquist and others 1997). Young 

stems are finely puberulent, greenish at first then 

become glabrous and reddish in color. Older stems 

are ashy-gray with a reddish-brown undertone 

(Cronquist and others 1997, Welsh and other 

1987). The alternate leaves are simple. A pair of 

reddish glands appears on the petioles near the 

base of the leaves. Leaves are 2 to 10 cm in length 

and 1.5 to 7 cm in width. Leaves are elliptic to 

oblong-ovate, finely serrated and abruptly 

acuminate apically. At the base, the leaves are 

acute to rounded. Leaves are dark green above and 

pale beneath. The midrib is impressed above and 

prominent beneath. (Cronquist and others 1997, 

Welsh and others 1987). Inflorescence is a raceme 

that is 4 to 20 cm long with leafy peduncles 2 to 8 

cm long. The white perfect flowers are 10 to 20 

mm wide and numerous with 4 to 17 mm long 

pedicels. Petals are white and are 4 to 6 mm long 

and suborbicular. Sepals are fringed and glabrous. 

Hypanthium and pedicels are also glabrous. The 

mature fruit 6 to 8 mm thick, known as a drupe, is 

dark red to black in color (Cronquist and others 

1997, Welsh and other 1987). The diploid 

chromosome number of chokecherry is 16, 26, or 

32 (Cronquist and others 1997). 

 

Taxonomy.—Two varieties of chokecherry are 

recognized: var. melanocarpa and var. demissa 

(Cronquist and others 1997). They are in the words 

of Cronquist and others 1997: “weakly separated 

as follows: 1. Leaves glabrous or rarely with a 

few tufts of hairs in the axils of the lateral leaf 

veins beneath; drupe blackish.........var. 

Melanocarpa (A. Nelson) Sarg. 2. Leaves 

pubescent; drupe dark red; California, in White 

Mountains and Sierra Nevada, westward 

......var. demissa (Nutt.) Torr.” 

Range.—Chokecherry occupies a variety of sites 

across Canada and the United States (Little 1976). 

It grows in 20 ecosystems and occurs in 47 of 

Kuchler’s plant associations (Pacific Southwest 

Experiment Station. 2002). The Society of 

American Foresters list chokecherry as occurring 

in 51 of its cover types, and the Society for Range 

Management finds chokecherry growing in 40 of 

its rangeland cover types (Pacific Southwest 

Experiment Station 2002). It ranges from 

Newfoundland in eastern Canada to British 

Columbia in the west (Little 1976). Scatted stands 

can be found as far north as the southern portion of 

the Northwest Territories of Canada and as far 

south as western Texas, Southwest, and southern 

California (Little 1976). In the United States 

chokecherry occurs in the Northeast, Midwest, 

Central Plains, Pacific Northwest, and 

Intermountain regions (Little 1976). Isolated 

patches occur in Virginia, West Virginia, 

Tennessee, Kentucky, North Carolina and 

Oklahoma (Little 1976). 
595

Ecology.—Usually chokecherry can be found 

growing in canyon bottoms, sheltered slopes, 

along streams and roads. It grows in a wide range 

of soils, ranging from Entisols to Mollisols that 

have textures ranging from sandy loams to clays. It 

can be found from 177 (Michigan) to 3,100 (Utah) 

m in elevation where the combinations of soil and 

topography permit greater than average 

accumulation of moisture (Pacific Southwest 

Experiment Station. 2002). Weakly salty soils are 

tolerated by chokecherry, but it cannot tolerant 

soils that are poorly drained or suffer from 

prolonged flooding. It grows in soil pH ranges 

from 3.5 to 7.6. It is found in numerous habitat and 

plant associations that range from post disturbance 

invaders to early successional to climax or stable 

(Pacific Southwest Experiment Station 2002). 

Chokecherry is intolerant to intermediate in 

tolerance of shade and resprouts from root crowns 

and rhizomes readily, thus giving it the ability to 

persists under open or forest canopies of moderate 

densities. It is well adapted to disturbance by fire. 

Chokecherry is a primary host of the eastern tent 

caterpillar and a fungus Plowrightia stansburiana,

which causes black knot-like tumors on stems 

(Pacific Southwest Experiment Station 2002). 

Western X virus can kill entire stands of 

chokecherry. 

Reproduction.—Chokecherry is among the first 

of the deciduous woody shrubs to leaf out and to 

flower in the spring. It can reproduce sexually 

from seeds and asexually from root crowns and 

rhizomes. Seeds are encased in a stony endocarp 

and have an after-ripening requirement for 

germination. Heat treatment improves 

germination. Seeds weigh about 0.095 grams 

(Grisez 1974). Passing through digestive systems 

of mammals and birds may enhance germination 

(Auger and others 2002). Also a number of birds 

and mammals may be responsible for long 

distance dispersal (Auger and others 2002). 

Chokecherry seeds persists in the soil seedbank . 

Growth and Management.—Heights of this 

species vary greatly as to variety and site quality. 

Chokecherry may grow as high as 12 m with a 

truck diameter 20 cm on some sites in the Great 

Basin. It has a deep root system that can grow to 

depths exceeding 1.8 m with lateral roots more 

than 10.6 m in length. Rhizomes ranging from 1 to 

2 cm in diameter are produced by this species. 

Chokecherry rhizomes sprout at a faster rate with a 

higher percentage of sprouts than Gambel oak. It is 

moderately tolerant of browsing. However, 

excessive livestock grazing has damaged some 

populations in many areas of the northern Great 

Plains. Hydrogen glycoside prunasin is a toxin 

produced by chokecherry that could be poisonous 

to grazing livestock. Highest levels (5 percent) are 

found in new stems and leaves. These levels 

diminish over the growing season to 1.2 to 2.2 

percent. 

Benefits.—Chokecherry adds to the biodiversity 

of a multitude of ecosystems. It provides habitat 

and food for a number of wildlife species, and 

watershed protection. Fruits, leaves, or twigs are 

eaten by bears, moose, bighorn sheep, pronghorn, 

elk and deer. A number of small of mammals also 

consume chokecherry including coyotes, 

snowshoe hares, red foxes, bobcats, raccoons, and 

porcupines. Chokecherry fruits are eaten by many 

birds including robins, western, eastern, and 

mountain bluebirds, European starlings, and 

Columbian sharp-tailed grouse. Livestock also eat 

chokecherry. Winter twigs contain 38.9 percent 

total digestible nutrients, which is lower than a 

number of winter shrubs (Dietz 1972, Welch 

1981.) Winter protein, calcium, and phosphorus 

levels are above average. Fruits are harvested to 

make wines, syrups, jellies, and jams. Chokecherry 

plants are planted as ornamentals, for enhancing 

backyard wildlife habitats, and as windbreaks. 

 

References 

Auger, J., S.E. Meyer, and H.L. Black. 2002. Are 

American black bears (Ursus americanus) 

legitimate seed dispersers of fleshy-fruited 

shrubs?American Midland Naturalist. 147:352-

367. 

 

Cronquist, A., N.H. Holmgren, and P.K. Holmgren 

1997. Intermountain flora: vascular plants of the 

Intermountain west, U.S.A. Vol. 3. Part A; 

Subclass Rosidae (except Fabales). The New 

York Botanical Garden, New York. 446 p. 

Dietz, D.R. 1972. Nutritive value of shrubs. In: 

C. M. McKell, J. P. Blaisdell, J. R. Goodin, tech. 

eds. Wildland shrubs–their biology 

andutilization, an international symposium; 

Proceedings; 1971 July; Logan, UT. General 

Technical Report INT-1. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Forest and Range Experiment Station, Ogden, 

UT. p. 289-302. 
596 

Grisez, T.J. 1974. Prunus L. Cherry, peach, and 

plum. In: C.S. Schopmeyer, tech. coord. Seeds 

of woody plants in the United States. 

Agriculture Handbook 654. U.S. Department of 

Agriculture, Forest Service, Washington, DC. 

p. 658-673. 

Little, E.L., Jr. 1976. Atlas of United States trees. 

Vol. 3. Minor Western hardwoods. 

Miscellaneous Publication 1,314. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 210 p. 

Pacific Southwest Experiment Station. 2002. Fire 

effects information system. http://www. 

fs.fed.us/ database/feis/plant. 33 p. 

Welsh, S. L., N. D. Atwood, S. Goodrich, L. 

C.Higgins. 1987. A Brigham Young 

University, Utah flora. Great Basin Naturalist 

Memoirs 9. Provo, UT. 894 p. 

 
597

Psidium guajava L. guava 

MYRTACEAE 

Synonyms: Psidium pumilum Vahl 

 Psidium guajava L. var. pumilum (Vahl) Griseb. 

 Psidium guajava L. var. cujavillum King & Urban 

John K. Francis 

 

General Description.—Guava, also known as 

guayaba, guayabo, arazá-puitá, goyavier, and 

gobiabiera, is an evergreen shrub or small tree 2 to 

8 m in height and up to 40 cm in diameter at breast 

height. Plants may have a single stem, especially if 

crowded in secondary forest, but individuals 

receiving ample light usually develop secondary 

stems arising from the main stem near the ground. 

The branches and stems are usually crooked and 

have a smoothish, cream to reddish-brown bark 

between thin, irregular scales that peel off. Guava 

bark is 5 to 8 mm thick (Pennington and Sarukhan 

1968). The sapwood is light brown and the 

heartwood is reddish brown, hard, heavy (specific 

gravity of 0.8), and strong (Little and Wadsworth 

1964). The roots are slender. The young twigs are 

four-angled, slightly winged, and green, turning 

brown with age. The leathery and light green or 

yellow green opposite leaves have petioles 4 to 7 

mm long, and elliptic or oblong blades, 8 to 14 cm 

long that are short or round pointed at both ends. 

The foliage is aromatic when crushed. The axillary 

flowers are usually solitary with four or five white 

petals. Fruits (berries) are globose or pear-shaped, 

with a prominent, persistent calyx. They are at 

first hard and green, becoming softer, and yellow 

at maturity and 3 to 5 cm long. Inside the thin rind 

is a sweet-tart flavored pink or yellow pulp 

containing many hard, angular, yellow seeds about 

2 mm long. The species has 2n = 21, 22, 30, or 33 

chromosomes (Howard 1989, Liogier 1994, Long 

and Lakela 1976, Pennington and Sarukhan 1968, 

Stevens and others 2001). 

Range.—Guava originated somewhere in the 

Neotropics and was spread nearly to the extent of 

its adapted climate before the arrival of Europeans 

(Morton 1987). Seeds found in Peruvian 

archeological sites seem to indicate that it 

originated in that area (Rain-tree 2002). Guava has 

been planted in nearly every tropical and frost-free 

subtropical country and has naturalized in most of 

them (Howard 1989). 

Ecology.—Guava grows on soils of all textures 

derived from most parent materials. Well-drained 

and poorly drained soils, soils with pH’s from 4.5 

to 9.4, mildly salty soils, and soils both rich and 

poor in basic cations are tolerated. Guava grows at 

near sea level in coastal environments up to 2,300 

m in elevation in Ecuador (Morton 1987). It grows 

naturally in areas of Puerto Rico that receive from 

about 1000 to 3000 mm of mean annual 

precipitation. Guava withstands drought very well. 

The species is moderately intolerant of shade. It 

develops a broad, low crown if open grown, grows 

a more vertical crown with side shade, and 

becomes tall and spindly in intermediate crown 

positions. Saplings can endure a few years in the 

understory of low basal-area secondary forests. 

Guava survives the competition of weeds, grass, 

and brush well. Growth is benefited by root 

association with arbuscular mycorrhizal fungi 

(Samarao and Martins 1999). Although they rarely 

kill the plants, a number of insects and diseases 

affect the species. As many as 80 percent of 

unprotected fruits may be attacked by 
598 

Mediterranean fruit flies (Ceratitis capitata Wied.) 

(Popenoe 1948). This thin-barked species is easily 

top-killed by fire and is sensitive to frost (von 

Carlowitz 1991). 

Reproduction.—Guava flowers and fruits nearly 

throughout the year (Little and Wadsworth 1964). 

Many individual plants bear just once per year but 

not necessarily synchronized with other guava 

plants in the area. Honey bees (Apis mellifera) are 

the chief pollinators (Morton 1987). Time from 

flowers to ripe fruits ranges from 102 to 124 days 

(Samson 1986). Thirty-one ripe fruits collected 

from wild plants in Puerto Rico averaged 32.8 g 

and ranged from 13.5 to 61.8 g. Air-dried seeds 

averaged 0.0079 + 0.0001 g/seed or 127,000 

seeds/kg. Sown on moist peat, 63 percent 

germinated between 15 and 60 days of sowing. 

Germination was epigeal (author’s observation). 

Birds and mammals disperse the seeds (Invasive 

Species Specialist Group 2002). Root cuttings are 

often used for commercial propagation of 

improved varieties (Morton 1987). Branches and 

stems layer (root) when they come in contact with 

moist soil (author’s observation), and suckers also 

arise from roots near the trunk. Established plants 

coppice readily and withstand repeated cutting 

(Advisory Committee on Technology Innovation 

1983). 

Growth and Management.—Guava seedlings 

grow at a moderate rate, but older plants grow 

more slowly. Under good conditions, guava begins 

bearing fruits in 3 or 4 years. Plants live 40 years 

or more (Popenoe 1948). It is probably unwise and 

unnecessary to plant wild guava because the 

species is tough and aggressive and often invades 

agricultural (especially cattle pasture) and forest 

lands. Control is sometimes attempted, especially 

in cattle pastures and plantations. Sheep and goats 

graze the leaves and strip the bark and have been 

used to control it. Several herbicides are effective 

in controlling infestations (Pacific Island 

Ecosystems at Risk 2002). Repeated heavy 

plowing or repeated burning are also effective 

measures (Mune and Parham 1956). 

Benefits.—Guava helps protect the soil and can be 

a major participant in reforestation of disturbed 

areas and abandoned pastures. It furnishes food 

and cover for wildlife. Domestic animals eagerly 

consume the fruits. Guava is one of the most 

important fruits in the tropics and is exported to 

temperate areas. It is consumed fresh and made 

into juice, jams and jellies, and paste or “cheese.” 

The fruit rinds are candied or stewed in syrup. 

Nearly all the commercial production comes from 

improved varieties with large fruits and few seeds. 

The ripe fruits contain the following components: 

water 84 percent, ash 0.7 percent, protein 0.8 

percent, fiber 5.6 percent, total sugars 5.4 percent, 

starch 2.5 percent, and fat 1.0 percent (Popenoe 

1948). The fruit pulp is rich in vitamins A and C 

(Morton 1987). Guava wood is used for tool 

handles, carving, and fuel (Advisory Committee 

on Technology Innovation 1983). In folk 

medicine, extracts of roots, bark, and leaves are 

used to treat gastroenteritis, vomiting, diarrhea, 

dysentery, wounds, ulcers, toothache, coughs, sore 

throat, inflamed gums, and a number of other 

conditions (Morton 1987). Guava leaf tea is 

widely used to control blood sugar of diabetics in 

Japan and elsewhere. It has been shown to be 

effective in vitro, in mice, and in human 

volunteers (Deguchi and others 1998). The basis 

for herbal treatment of diarrhea was established by 

demonstrating inhibition of eight bacteria species 

and amoebas, and antispasmodic activity (Tona 

and others 1999). 

References 

Advisory Committee on Technology Innovation 

1983). Firewood crops. Vol. 2. National 

Academy of Sciences, National Academy Press, 

Washington, D.C. 92 p. 

Deguchi, Y., K. Osada, K. Uchida, H. Kimura, M. 

Yoshikawa, T. Kudo, H. Yasui, and M. 

Watanuki. 1998. Effects of extract of guava 

leaves on the development of diabetes in the 

db/db mouse and on the postprandial blood 

glucose of human subjects. Nippon 

Nogeikagaku Kaishi 72(8): 923-931. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Invasive Species Specialist Group. 2002. Notes 

from Jim Space on his survey of invasive plant 

species in Tonga. The World Conservation 

Union. http://www.issg.org/features/ 

invasives_on_tonga.html. 4 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 
599

Little, E.L., Jr. and F.L. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Washington, DC. 

548 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Morton, J. 1987. Guava. In: J.F. Morton. Fruits of 

warm climates. Julia F. Morton, Maimi, FL. 

p. 356-363. 

Mune, T.L. and J.W. Parham. 1956. Weed control: 

Guava and its control in Fiji. Agricultural 

Journal, Fiji 27(3/4): 103-108. 

Pacific Island Ecosystems at Risk. 2002. Psidium 

guajava L., Myrtaceae. http://hear.org/pier_v3.3/ 

psgua.htm. 4 p. 

Pennington, T.D. and J. Sarukhan. 1968. Arboles 

tropicales de México. Instituto Nacional de 

Investigación Forestales, Secretaría de 

Agricultura y Ganadaría. Ciudad de México, 

México. 413 p. 

Popenoe, W. 1948. Manual of tropical and 

subtropical fruits. Hafner Press, New York. 

474 p. 

Rain-tree. 2002. Guava. Raintree Nutrition, Inc., 

Austin, Texas. http://www.rain-tree.com/guava. 

htm. 5 p. 

Samarao, S.S. and M.A. Martins. 1999. Influence 

of arbuscular mycorrhizal fungi, associated with 

addition of rutin, on the growth of guava 

(Psidium guajava L.). Revista Brasileira de 

Fruticultura 21(2): 196-199. 

Samson, J.A. 1986. Tropical fruits. 2nd ed. 

Longman Scientific & Technical, Harlow, UK. 

336 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden, St. Louis, MO. 

p. 945-1,910. 

Tona, L., K. Kambu, K. Mesia, K. Cimanga, S. 

Apers, T. de Bruyne, L. Pieters, J. Totte, A.J. 

Vlietinck, and T. de Bruyne. 1999. Biological 

screening of traditional preparations from some 

medicinal plants used as antidiarrhoeal in 

Kinshasa, Congo. Phytomedicine 6(1); 59-66. 

von Carlowitz, P.G. 1991. Multipurpose trees and 

shrubs: Sources of seeds and inoculants. 

International Council for Research in 

Agroforestry, Nairobi, Kenya. 328 p. 
600 

Psorothamnus scoparius (Gray) Rydb. broom dalea 

FABACEAE 

Synonyms: Dalea scoparia Gray 

 Parosela scoparia (A. Gray) Heller 

James E. Nellessen 

General Description.—Broom dalea is also 

known as broom indigobush and purple sage 

(Dick-Peddie 1993, Ivey 1995). This is a highly 

branched shrub with slender twigs and small 

leaves. Its overall growth form gives it a broomlike appearance. It may reach heights of 1.5 m or 

more, but is typically about 1 m when mature. 

Horizontal spread is likewise about 1 m but in 

older plants may be as much as 2 m or more. 

Leaves are alternate, linear to linear-spatulate, 

from 1 to 2 mm in width and 0.5 to 1.9 cm in 

length, sometimes trifoliate, and gland-dotted 

(Carter 1997, Martin and Hutchins 1980-81). 

Populations in New Mexico are actually much 

more frequently trifoliate than some floras would 

indicate. Flowers are pealike, dark blue, grouped 

in semi-spherical clusters, and fragrant. 

Range.—Broom dalea occurs from far western 

Texas through central and southern New Mexico 

into eastern Arizona and northern Mexico 

(Kearney and others 1951, Martin and Hutchins 

1980-81, Warnock 1974). It occurs at elevations 

from about 915 m to 1,830 m in upland desert 

scrub habitat, usually in sandy soils. 

Ecology.—This species is common in Plains-Mesa 

Sand Scrub habitats and can be one of the 

dominant shrubs in this vegetation type. It is a 

deep-sand tolerant or deep-sand adapted shrub. It 

may occur on gypsum sands as well as quartz 

sands. More specifically it occurs in a mixed shrub 

series in association with Artemisia filifolia Torr., 

Atriplex canescens (Pursh) Nutt., Oryzopsis 

hymenoides (Roem. & Schult.) Ricker ex Piper, 

Sporobolus spp., and other mixed grasses and 

forbs (Dick-Peddie 1993). 

Reproduction.—Broom dalea often appears 

relatively dormant, with little new growth, until 

summer monsoon rains begin, whereupon it 

rapidly enters the blooming stage. The dark blue 

flowers are grouped in semi-spherical spikes, of a 

few to several flowers, the clusters 1 to 2 cm in 

diameter (Carter 1997, Martin and Hutchins 1980-

81). The fragrant blooms attract numerous 

pollinating bees. Fruits are short, egg shaped pods 

with hairs and glands. Under proper moisture 

conditions seeds may germinate the following 

year. New plants may reach reproductive status in 

as little as 3 to 6 years depending on local 

environmental conditions. 

Pollination.—Numerous species of bees are 

attracted to the blue flowers and collect pollen 

from broom dalea. The plant emits a noticeable 

sweet, aromatic scent when in bloom. During the 

peak of summer blooming, these shrubs are often 

focal points for bee foraging activity. Some bees 

visiting broom dalea include crepuscular (twilight 

active) bees of the family Colletidae (yellow-faced 

and plasterer bees), subfamily Diphaglossinae, 

species Caupolicana ocellata Michener; and 

family Anthophoridae (cuckoo, digger, and 

carpenter bees), subfamily Anthophorinae, Tribe 

Eucerini, species Martinapis luteicornis

(Cockerell) (Rozen and Rozen 1986). These bee 

species were observed in southeastern Arizona. 

Both groups of bees nest in the ground, in crevices 

in rocks or walls, in plant stalks or other cavities. 

Both groups use the pollen, or pollen mixed with 

nectar as a food source. 
601

Growth and Management.—This species grows 

relatively quickly and can reach 1 m in height or 

diameter within 4 to 7 years depending on local 

environmental conditions. The sandy Plains-Mesa 

Sand Scrub habitats in which this shrub may be a 

dominant are by nature poor rangelands for 

livestock. 

Benefits.—Broom dalea is an important natural 

stabilizer of dune and other sandy soils. Its 

relatively rapid growth rate allows for reasonably 

quick establishment in situations of shifting and 

disturbed soils. Like many members of the legume 

family, it is a nitrogen fixer and forms rhizobial 

root nodules (Allen and Allen 1981). In fact, this 

species has been documented to transmit the 

rhizobial bacteria to another genus within the 

legume family, Crotalaria (Wilson 1939). Broom 

dalea is not an important range or browse plant for 

livestock and may increase under grazing. It is not 

substantially browsed, likely due to oils contained 

within the glands (Warnock 1974). Very little, 

until recently, has been known about the volatile 

oil composition of this species. Sixty-four volatile 

compounds have been identified (Lucero and 

others 2002). The three most abundant compounds 

were gamma-terpinene (22.3 percent), p-cymene 

(14.0 percent), and alpha-pinene (9.0 percent). 

Some of these are known deterrents to herbivore 

browsing in other plant species and may serve the 

same function in broom dalea. 

References

Allen, O.N. and E.K. Allen. 1981. The 

Leguminosae: A Source Book of 

Characteristics, Uses, and Nodulation. 

University of Wisconsin Press, WI. 812 p. 

Carter, J.L. 1997. Trees and Shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation: Past, Present and Future. University 

of New Mexico Press, Albuquerque, NM. 244 p. 

Ivey, R.D. 1995. Flowering Plants of New Mexico, 

3rd Ed. Published by the author, NM. 504 p. 

Kearney, T.H., R. Peebles, and Collaborators. 

1951 with 1960 supplement. Arizona Flora. 

University of California Press, Berkeley, CA. 

1,085 p. 

Lucero, M.E., R.E. Estell, and E.L. Frederickson. 

2002. The essential oil composition of

Psorothamnus scoparius (A. Gray) Rydb. 

United States Department of Agriculture, 

Agricultural Research Service, Jornada 

Experimental Range http://www.nal. 

usda.gov/ttic/tektran. In press, Journal of 

Essential Oils. 

Martin, W.C. and C.E. Hutchins. 1980-1981 

(reprinted 2001). A Flora of New Mexico. Vol. 

1. Bishen Singh Mahendra Pal Singh, India and 

Koeltz Scientific Books, Germany. p. 1-1,276. 

Rozen, Jr., J.G. and Rozen B.L. 1986. Bionomics 

of crepuscular bees associated with the plant 

Psorothamnus scoparius: Hymenoptera, 

Apoidea. Journal of the New York 

Entomological Society. 94(4): 472-479. 

Warnock, B.H. 1974. Wildflowers of the 

Guadalupe Mountains and the Sand Dune 

Country, Texas. Sul Ross State University, 

Alpine, TX. 176 p. 

Wilson, J.K. 1939. Symbiotic promiscuity of two 

species of Crotalaria. Journal of the American 

Society of Agronomy 31: 934-939. 
602 

Psychotria brachiata Sw. palo de cachimbo 

RUBIACEAE 

Synonyms: Psychotria neurotricha DC. 

 Cephaelis polycephala Schltdl. 

 Palicourea caerulea (Ruiz & Pav.) Roem. & Schult. 

 

John K. Francis 

General Description.—Palo de cachimbo is an 

evergreen shrub occasionally reaching 5 m in 

height but usually 2 to 3 m in height and 3 to 6 cm 

in basal diameter. The shrub usually has several 

stems formed by suckers from the lateral roots a 

few cm out from the central stem and by branches 

low on the principal stems. The wood has a 

moderately low density, is moderately strong, and 

has faint annual rings. Palo de cachimbo plants are 

supported by relatively shallow lateral root 

systems, at least in soils that have poorly aerated 

subsoil. The roots are tan colored and flexible. The 

branches and twigs are green, slender, and usually 

paired. Shiny, dark-green leaves are ovate to 

lanceolate, entire and pointed at both ends, 7 to 20 

cm long and 3 to 8 cm broad, with a 1- to 3-cm 

petiole. Small yellow to greenish-white flowers 

are tightly clustered in open terminal panicles. The 

4-mm ellipsoidal berries are tightly clustered. 

They are dark purple or dark blue at maturity and 

usually contain two hemispherical seeds having 

the inside face concave and the outside face with 

five ridges (Croat 1978, Liogier 1997, Stevens and 

others 2001). 

Range.—Palo de cachimbo is native to the 

Greater Antilles, Trinidad, and Guatemala to Peru 

(Liogier 1997). The Missouri Botanical Garden 

(2002) lists some 73 herbarium specimens from 

South America. However, some researchers (Croat 

1978, Stevens and others 2001) contend that the 

South American range is based on 

missidentification of the similar P. caerullea Ruiz 

& Pav. Palo de cachimbo is not known to have 

been planted or naturalized elsewhere. 

Ecology.—Palo de cachimbo grows mostly in wet 

forest areas that receive from 2000 to 3000 mm of 

mean annual precipitation. It occurs at elevations 

from near sea level to 800 m in Nicaragua 

(Stevens and others 2001). Palo de cachimbo is 

intolerant of shade. Plants compete well with 

herbs and other shrubs in openings but do not 

survive long after a forest canopy closes over 

them. The species is common to uncommon in 

roadsides, secondary forest, and openings in 

primary forest. Plants sampled in secondary forest 

in Puerto Rico had the following nutrient levels in 

their leaves: 2.36 + 0.03 percent N, 0.12 + 0.00 

percent P, 2.08 + 0.25 percent K, and 12.66 + 0.71 

percent ash. Lower levels of those nutrients were 

reported for twigs and stem (Lugo 1992). 

Reproduction.—Palo de cachimbo flowers from 

February to August and fruits from July to April in 

Nicaragua (Stevens and others 2001). The species 

is a good producer of fruits and seeds. A collection 

of fresh fruits from Puerto Rico weighed an 

average of 0.1569 + 0.0125 g/fruit. Air-dried seeds 

separated from them averaged 0.0048 + 0.0002 

g/seed or 208,000 seeds/kg. Placed in commercial 

potting mix, 48 percent germinated between 34 

and 57 days following sowing. Seeds are 

presumably dispersed by birds. Seedlings are 

common in the vicinity of fruit-producing plants. 

Growth and Management.—Palo de cachimbo 

grows slowly in the early seedling stage. It grows 
603

0.5 to 0.8 m/year for 2 or 3 years from sprouts. 

Individual stems probably live from 10 to 20 

years, but plants may persist longer by sprouting. 

Natural regeneration probably can be encouraged 

by disturbance designed to create forest openings. 

Planting and management experience have not 

been published. 

Benefits.—Palo de cachimbo contributes to the 

biodiversity of forests, helps protect the soil, and 

furnishes food and cover for wildlife. 

References 

Croat, T.B. 1978. Flora of Barro Colorado Island. 

Stanford University Press, Stanford, CA. 943 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Lugo, A.E. 1992. Comparison of tropical tree 

plantations with secondary forests of similar 

age. Ecological Monographs 62: 1-41. 

Missouri Botancial Garden. 2002. W3

Tropicos: 

nomenclature data base. http://mobot.mobot. 

org/cgi-bin/search_vast#samer. [not paged]. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanical Garden, St. Louis, MO. 

p. 1,911-2,666. 
604 

Psychotria deflexa DC. garricillo

RUBIACEAE 

Synonyms: Psychotria patens of authors, not of Sw. 

 Psychotria flexulosa of authors, not of Willd. 

 Palicourea patens (Sw.) Urban 

John K. Francis 

General Description.—Garricillo (a Panamanian 

name), sometimes referred to by the general name 

“cachimbo,” is an evergreen shrub usually 1.5 to 

2.0 m and 1.0 to 2.0 cm in basal diameter, 

sometimes reaching 3 m in height. Multiple stems 

are common, but not thick clumps. The wood is 

moderately hard with faint annual rings. The 

angular twigs and the foliage are glabrous. 

Persistent, bilobed, long-pointed stipules are 

characteristic of the species. Paired, dark-green, 

elliptic to oblong lanceolate leaves are 5 to 18 cm 

long with 3- to 10-mm petioles. Inflorescences are 

few-flowered terminal panicles. The small, sessile 

flowers are white. Fruits are subglobose, 3 to 5 

mm in diameter, white or pale blue. Normally, 

there are two hemispherical seeds per fruit that are 

striated on the convex surface and have a medial 

groove on the opposite, flat or slightly concave 

surface (Croat 1978, Liogier 1997, Stevens and 

other 2001). 

Range.—Garricillo is native to the Greater 

Antilles, Trinidad, southern Mexico, Central 

America, and South America to Bolivia (Croat 

1978, Liogier 1997, Stevens and others 2001). It is 

not known to have naturalized outside the native 

range. 

Ecology.—Garricillo is a shrub of moist and wet 

forests, areas receiving from about 1600 mm to 

over 3000 mm of mean annual precipitation. 

Although the species may occur from a few meters 

above sea level to over 1,000 m in elevation 

(Instituto Nacional de Biodiversidad 2002), it 

generally grows at medium to high elevations 

(Liogier 1997) because there is greater available 

moisture. The species grows on poorly drained to 

well drained soils generally with loamy to clayey 

texture and pH’s from about 4.5 to 6.5. Garricillo 

is intermediate in tolerance to shade and grows in 

the forest understory (if not too dark), at forest 

edges, in overgrown fencerows (Molano and 

others 2002), and in openings. Most fruit and seed 

production takes place in openings or situations 

with increased to full sunlight. The species 

competes well with herbs and low brush. 

Reproduction.—Garricillo flowers from May to 

July and matures fruits mostly from August to 

December in Panama (Croat 1978). The flowers 

are both insect-pollinated and self-pollinated 

(Faivre 2002). Fresh fruits collected in Puerto 

Rico weighed an average of 0.0758 + 0.0006 

g/fruit. Air-dried seeds separated from them 

weighed an average of 0.0057 + 0.0001 g/seed, or 

175,000 seeds/kg. There was an average of 1.7 

seeds/fruit in this sample. Sown without 

pretreatment in moist potting mix, 69 percent 

germinated between 68 and 117 days after sowing. 

Fruit and seed production is good and consistent. 

Birds disperse the seeds (Molano and others 

2002). Seedlings are common but suffer high 

mortality. Damaged plants resprout. Prostrate 
605

stems of young plants layer (root) readily when in 

contact with the soil. 

Growth and Management.—Seedlings of 

garricillo grow slowly at first. Older plants and 

sprouts add from 20 to 40 cm to their height each 

year. Individual stems live about 10 years; plants 

may last longer by producing new sprouts. 

Although the species is more or less common, it 

does not appear to cause any difficulties for forest 

users. 

Benefits.—Garricillo contributes to the 

biodiversity and biomass of forest communities, 

helps protect the soil, and furnishes food and 

cover for wildlife. 

References 

Croat, T.B. 1978. Flora of Barro Colorado Island. 

Stanford University Press, Stanford, CA. 943 p. 

Faivre, A.E. 2002. Reproductive biology of seven 

understory Psychotria species on Barro 

Colorado Island, Panama. Cedar Crest College, 

Allentown, PA. http://botany2002.org.section3/ 

abstracts/21.shtml. 1 p. 

Instituto Nacional de Biodiversidad. 2002. Lista de 

especímenes de Psychotria deflexa. Costa Rica. 

http://www.inbio.ac.cr/bims/k03/p13/c045/o014

2/f01359/g008913/s028252.htm. 4 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Molano, J.G., M.P. Quiceno, and C. Roa. 2002. El 

papel de las cercas vivas en un sistema de 

producción agropecuaria en el Piédemonte 

Llanero. http://www.lead.virtualcentre.org/ 

es/ele/conferencia2/vbconfe3.htm. 18 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol.85, No. 

3. Missouri Botanical Garden Press, St. Louis, 

MO. p. 1,911-2,666. 
606 

Psychotria microdon (DC.) Urban thicket wild coffee 

RUBIACEAE 

Synonyms: Psychotria pinularis Sessé & Moc. 

 Rondeletia microdon DC. 

John K. Francis 

General Description.—Thicket wild coffee, also 

known as café bâtard, and café marron, is a 

semideciduous arching or scrambling shrub 1 to 3 

m in height with a basal diameter of up to 4 cm. 

There is usually a single stem from the ground in 

undisturbed plants with multiple branches low on 

the stem. The arching stems sometimes extend 2 or 

3 m horizontally and tend to be very branchy. The 

wood is of medium density, moderately weak and 

brittle. Stem and branch bark is smooth and tan 

outside and green within. The plant is supported 

by a tap-and-lateral root system of dark tan and 

flexible roots. The foliar display is medium to thin. 

Membranous, glabrous, obovate to oblanceolate, 

light-green leaves are 4 to 11 cm long with 10- to 

18-mm petioles. The inflorescences are terminal 

corymbiform cymes with three to five rays. The 

flowers have a five-toothed white corolla tube 5 to 

10 mm long. Fruits are red or red-orange 

ellipsoidal berries about 7 mm long. They usually 

contain two tan seeds that are flattened on one side 

and rounded on the other (Howard 1989, Liogier 

1997, Stevens and others 2001). 

Range.—Thicket wild coffee is native to the 

Greater Antilles (except Jamaica), the Lesser 

Antilles, from Central Mexico south through 

Central America to Bolivia and eastwards to 

Guiana (Howard 1989, Liogier 1997, Stevens and 

others 2001). It is not known to have been planted 

or naturalized elsewhere. 

Ecology.—Thicket wild coffee is occasional to 

relatively common in secondary and remnant 

forests from near sea level to 840 m (Stevens and 

others 2001). Mean annual precipitation for the 

species in Puerto Rico ranges from 750 to about 

1700 mm. It grows on a variety of well-drained 

soils that developed from alluvium, and igneous 

and sedimentary (including limestone) rocks. 

Thicket wild coffee is moderately intolerant to 

intermediate in tolerance to shade. It will grow and 

fruit in openings and low basal-area stands and 

survives in forest stands with moderate basal areas. 

The species endures light grazing but disappears 

under heavy grazing pressure. 

Reproduction.—In Nicaragua, thicket wild coffee 

flowers May through September and fruits August 

through September (Stevens and others 2001). 

Fruit and seed production is moderate. A 

collection of fresh fruits from Puerto Rico weighed 

an average of 0.259 + 0.005 g/fruit. Air-dried 

seeds separated from them averaged 0.0187 +

0.0004 g/seed or 53,000 seeds/kg. Sown without 

pretreatment on moist blotter paper, 78 percent 

germinated between 65 and 217 days after sowing. 

Germination is epigeal. The seeds are apparently 

dispersed primarily by birds. Tapirs eat the fruits, 

but the seeds are destroyed in the process (Olmos 

1997). Seedlings are common near seed sources in 

Puerto Rico. Survival to adulthood is relatively 

rare, however. 

Growth and Management.—Thicket wild coffee 

is slow growing as a new seedling and grows at a 

moderate rate as a sapling and adult. Plants survive 

at least 5 years and perhaps much longer. Planting 

and management experience has not been 

published. The species is not common enough or 

aggressive enough to warrant control. 

Benefits.—Thicket wild coffee contributes to the 

biodiversity of forests, helps protect the soil, and 

furnishes food and cover for wildlife. The fruits 

are slightly sweet and edible but not highly 
607

desirable. The species is a larval host for the 

Xylophanes pluto (Fabricius) moth in Puerto Rico 

(Torres-Bauzá 2000). 

References

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Part 3. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p.

Olmos, F. 1997. Tapirs as seed dispersers and 

predators. IUCN/SSC Tapir Specialist Group. 

http://www.tapirback.com/tapirgal/iucn-ssc/tsg/ 

 action97/ap97-05.htm. 12 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua, 

Angiospermas. Monographs in Systematic 

Botany Vol. 85, No. 3. Missouri Botanical 

Garden Press, St. Louis, MO. p. 1,911-2,666. 

Torres-Bauzá, J.A. 2000. Ciclo de vida y aspectos 

de la biología de Xylophanes pluto (Fabricius) 

en Puerto Rico (Lepidoptera: Sphingidae). 

Caribbean Journal of Science 36(3-4): 227-232. 
608 

Psychotria nervosa Sw. shiny-leafed wild coffee 

RUBIACEAE 

Synonyms: Psychotria undata Jacq. 

 Psychotria chimarrhoides DC. 

 Psychotria lanceolata Nutt. 

 Psychotria hirsuta Spreng. 

 Psychotria oligotricha DC. 

 Psychotria portoricensis DC. 

John K. Francis 

General Description.—Shiny-leafed wild coffee, 

also known as wild coffee, Bahaman wild coffee, 

wood balsam, Seminole balsamo, café marrón, 

palo moro, ti café marron, ti kafé mawon, and 

remaroogu, is a 1- to 3-m erect shrub. Stem-like 

low branches are common, and multiple stems are 

formed in response to damage to the trunk. The 

stems, sometimes up to 7 cm in diameter, are 

covered with a bone-colored, smooth bark that is 

hairy in some variants. The plant is supported by 

stiff tap and lateral roots and abundant fine roots. 

Relatively few branches and twigs are formed 

with foliage maintained only near the tips. The 

leaves are shiny light or dark green, opposite, 

simple, glabrous or nearly so, elliptic to 

lanceolate, with sunken veins and elongated tips. 

A large, flattened bud is almost always visible 

between the youngest pair of leaves. The flower 

clusters, which are terminal or axillary, are nearly 

globose and contain a number of small, white 

flowers. The clusters become more diffuse as the 

fruits mature. The fruits (drupes) are bright red, 

elliptic or globose at maturity and 6 to 7 mm long 

(Howard 1989, Liogier 1997, Nelson 1996, 

Stevens and other 2001). They are fleshy and juicy 

with little flavor. Each contains two seeds that 

have a rounded side with shallow striations and a 

flattened side with a division down the middle like 

a coffee bean (author’s observation). 

Range.—Shiny-leafed wild coffee is native to 

Florida, the West Indies, Southern Mexico, 

Central America, and Colombia, Ecuador, and 

Venezuela in South America (Liogier 1997, 

Stevens and others 2001). 

Ecology.—Shiny-leafed wild coffee grows in dry 

and moist forests that receive from about 700 to 

2000 mm of mean annual precipitation. At the 

moist end of the precipitation range, it tends to be 

found only on excessively drained sites. It is coldsensitive, tolerating temperatures only to about 

freezing (Dolan 2002). It grows well above the 

frost line in Florida; presumably shiny-leafed wild 

coffee freezes to the ground and resprouts later. 

Well-drained soils with the range of textures and 

both igneous and sedimentary (especially 

limestone) parent materials are colonized. Wild 

coffee is moderately tolerant of shade, usually 

growing in the understories of low basal area 

forests. The shrub may also be found in natural 

and artificial openings. In Florida, it is usually 

found in shell ridges, hammocks, and pinelands 

(School of Forest Resources and Conservation 

2002). Shiny-leafed wild coffee is occasionally 

found in pure clumps, but more often it grows as 

well-distributed plants. 
609

Reproduction.—Shiny-leafed wild coffee blooms 

in spring and summer in Florida (Nelson 1996) 

and irregularly through the year in Nicaragua 

(Stevens and others 2001). The flowers are visited 

by honeybees and butterflies (Dave’s Garden 

Network 2002) that presumably pollinate them. 

A collection of fruits from Puerto Rico weighed an 

average of 0.256 + 0.009 g/fruit. Air-dried seeds 

from those fruits weighed an average of 0.0175 +

0.0004 g/seed or 57,000 seeds/kg. About 85 

percent of fruits from the collection contained two 

seeds; the reminder contained one filled seed and 

one aborted seed. Sown in commercial potting 

mix, 95 percent of the filled seeds germinated 

between 64 and 164 days after sowing (author’s 

observation). 

Growth and Management.—Shiny-leafed wild 

coffee has a moderate growth rate. Small wildlings 

and pruned larger plants transplant well. They also 

can be grown easily from seed. Once established 

in a garden or natural area, they will reproduce 

and eventually fill the area (Workman 1980). 

Where a seed source is available, probably the 

best strategy to promoting natural establishment is 

to manipulate the stand for an open understory. 

Benefits.—Shiny-leafed wild coffee is a 

widespread and important understory plant in dry 

and moist forest and as such contributes to the 

biodiversity and biomass accumulation, and 

protects against soil erosion. It is frequently 

planted in Florida as a native landscape plant in 

both formal and natural gardens (Workman 1980). 

It is appreciated for its attractive foliage and bright 

red berries and because it attracts birds and 

butterflies (Dave’s Garden Network 2002). The 

fruits are food for tapirs (Olmos 1997), a number 

of bird species (School of Natural Resources and 

Conservation 2002), and probably many other 

animals. Shiny-leafed wild coffee contains the 

alkaloid dimethyltryptamine, which is 

hallucinogenic (Ted 2002) and may eventually 

prove to have medicinal applications. The seeds 

were once used as a coffee substitute (Dave’s 

Garden Network 2002). 

References 

Dave’s Garden Network. 2002. Wild coffee 

(Psychotria nervosa). The Plants Database. 

http://plantsdatabase.com/go/2326.html. 2 p. 

Dolan, S. 2002. Native shrubs. 

http://www.indialantic.com/contents/ecology/ 

shurbs.htm. 3 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Part 3. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Olmos, F. 1997. Tapirs as seed dispersers and 

predators. IUCN/SSC Tapir Specialist Group. 

http://www.tapirback.com/tapirgal/iucn-ssc/tsg/ 

 action97/ap97-05.htm. 12 p. 

School of Forest Resources and Conservation. 

2002. Florida forest plants: wild coffee 

(Psychotria nervosa). University of Florida, 

Gainesville, FL. http://www.plantatlas.usf.edu/ 

maps.asp?plantID=2811. 2 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua, 

Angiospermas. Monographs in Systematic 

Botany Vol. 85, No. 3. Missouri Botanical 

Garden Press, St. Louis, MO. p. 1,911-2,666. 

Ted, M. 2002. Smokable dimethyltryptamine from 

organic sources. http://www.deoxy.org/ 

smokedmt.htm. 11 p. 

Workman, R.W. 1980. Growing native. The 

Sanibel-Captiva Conservation Foundation, Inc., 

Sanibel, FL. 137 p. 
610 

Purshia tridentata (Pursh) DC. antelope bitterbrush 

ROSACEAE 

Synonyms: Kunzia tridentata Spreng.

Tigarea tridentata Pursh 

 

Nancy L. Shaw and Stephen B. Monsen 

 

Illustration source: USDA Forest Service 1937 

General Description.—Antelope bitterbrush, also 

known as antelope brush, antelope buckbrush, 

deerbrush, or quininebrush, is an intricately 

branched, deciduous shrub varying widely in 

growth habit from low, decumbent, spreading 

forms to upright arborescent plants over 4 m in 

height (Blauer and others 1975, Welsh and others 

1987). Leaves are alternate, simple, pinnatifid or 

apically three-toothed and sometimes glandular. 

Flowers are insect-pollinated and borne on short 

spurs of previous year’s growth. They are 

numerous and white to yellow with petals 5 to 9 

mm long (Welsh and others 1987). The fruit is a 

cartilaginous achene with a persistent tapering 

style and a dark pinkish gray pyriform seed 

(Blauer and others 1975). Purshia DC. ex Poir. 

likely derived from Cowania (D. Don) (Jabbes 

2000, McArthur and others 1983) and hybrids 

between Cowania and Purshia are common. 

Purshia glandulosa Curran is considered a 

stabilized hybrid between Cowania and Purshia 

tridentata. 

Range.—Antelope bitterbrush is widely 

distributed from British Columbia southward on 

the east side of the Cascade and Sierra Nevada 

Mountains to California and northern Arizona and 

eastward into western Montana, the Black Hills of 

South Dakota, and northwestern New Mexico 

(Cronquist and others 1997, Welsh and others 

1987). 

Ecology.—Antelope bitterbrush grows on welldrained, slightly basic to slightly acidic soils at 

elevations from 60 to 3,510 m (Nord 1965). It 

often grows as the dominant shrub with bluebunch 

wheatgrass and a variety of forbs. It is also a 

common associated species in many big sagebrush 

(Artemisia tridentata L.), mountain brush, pinyon–

juniper (Pinyon L.-Juniperus L.), ponderosa pine 

(Pinus ponderosa P. & C. Lawson), and 

occasionally lodgepole pine (Pinus contorta

Dougl. ex Loud.) communities (McArthur and 

others 1983, Nord 1965). It is a pioneering species 

on steep, rocky, unstable disturbances. Erect forms 

generally do not survive wildfires, but layering 

forms may resprout. Antelope bitterbrush is 

sometimes associated with the nitrogen-fixing 

actinomycete Frankia, but root nodulation varies 

with factors such as moisture availability and soil 

chemistry (Righetti and others 1983). Emerging 

seedlings are susceptible to a number of dampingoff organisms, while seedlings and mature plants 

may be damaged by grasshoppers, Great Basin tent 

caterpillars (Malacosoma fragile Stetch), and other 

insects (Nord 1965, Shaw and Monsen 1983). 

 

Reproduction.—Plants flower in April to June, 

and fruits ripen in late June to August depending 

upon elevation and latitude (Nord 1965). Seed 

production is related to precipitation and leader 

growth in the previous year, but yields are often 

drastically reduced by insect predators (Shaw and 

Monsen 1983). Seeds dehisce rapidly at maturity. 

High percentages of the achenes are cached by 

rodents, and most successfully establishing natural 

recruitment is from these caches (Evans and others 

1983, Nord 1965, Van der Wall 1994). Large 

quantities of seed are hand harvested from 

wildland stands each year. Dry seed may be stored 

in a warehouse for up to 15 years (Stevens and 

others 1981). A 28-day moist prechill is required 

to release seed dormancy (AOSA 1993, Booth 
611

1999, Meyer 1989). Site-adapted seed sources 

should be selected for plantings. Antelope 

bitterbrush should be drill seeded in late fall or 

winter to provide overwinter prechilling. It may be 

seeded with other native species that are not highly 

competitive. Seeding failures resulting from rodent 

predation of seed can be avoided by seeding in late 

fall on fairly large disturbances that do not provide 

cover for the rodents (Evans and others 1983). 

Containerized or 1-0 bareroot seedlings are easily 

grown, but fungicide treatments may be required 

to reduce losses to damping-off organisms. 

Nursery stock can be planted on steep, rocky 

slopes that are inaccessible to seeding equipment. 

It establishes best if planted in early spring when 

antelope bitterbrush in the surrounding area is yet 

dormant. Herbaceous vegetation must be cleared 

around each seedling to reduce competition. 

Growth and Management.—Antelope 

bitterbrush seedings, plantations, and wild stands 

are vulnerable to heavy use by wild or domestic 

ungulates and predation by gophers, mice, birds, 

and insects. Seedlings are not competitive with 

weedy annuals or most introduced perennial 

grasses. Many mature stands are damaged by 

continuous and close browsing by deer and 

livestock. 

Benefits.—Next to sagebrush, antelope bitterbrush 

is probably one of the most widespread shrubs in 

the Great Basin region, although its prevalence has 

been reduced by range fires and exotic plant 

introductions (Mozingo 1987). It is important for a 

host of wildlife and provides nutritious browse and 

essential cover on fall, winter, and summer mule 

deer ranges (Plummer and others 1968). Protein 

content varies among populations depending upon 

over-winter leaf retention. New growth may also 

be heavily used in spring and summer. Because of 

the varied growth forms, attractive foliage, and 

showy flowers, antelope bitterbrush has potential 

as an ornamental in low-maintenance landscapes. 

Populations of antelope bitterbrush with distinctive 

attributes have been recognized and are 

commercially harvested and sold. Native 

Americans reportedly used bitterbrush as a remedy 

for smallpox, measles, tuberculosis, and 

pneumonia, and as an antiseptic for rashes and 

insect bites (Mozingo 1987). 

References 

AOSA [Association of Official Seed Analysts]. 

1993. Rules for testing seeds. Journal of Seed 

Technology 16: 1–113. 

Blauer, A.C., A.P. Plummer, E.D. McArthur, R. 

Stevens, and B.C. Giunta. 1975. Characteristics 

and hybridization of important Intermountain 

shrubs. I. Rose family. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Forest and Range Research Station, Ogden, UT. 

35 p. 

Booth, D.T. 1999. Imbibition temperatures affect 

bitterbrush seed dormancy and seedling vigor. 

Journal of Arid Environments 48: 35-39. 

Cronquist, A., N.H. Holmgren, and P.K. 

Holmgren. 1997. Intermountain flora: vascular 

plants of the Intermountain West, U.S.A. Vol. 3, 

part 1. Subclass Rosidae: (except Fabales). The 

New York Botanical Garden, The Bronx, NY. 

446 p. 

Evans, R.A., J.A. Young, G.J. Cluff, and J.K. 

McAdoo. 1983. Dynamics of antelope 

bitterbrush seed caches. In: A.R. Tiedemann and 

K.L. Johnson, comps. Proceedings: Research 

and management of bitterbrush and cliffrose in 

western North America. GTR-INT-152. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. p. 195–202. 

Jabbes, M. 2000. Hybridization and its 

evolutionary consequences in Purshia and 

Cowania. Ph.D. dissertation. University of 

Idaho, Moscow, ID. 185 p. 

McArthur, E.D., H.C. Stutz, and S.C. Sanderson. 

1983. Taxonomy, distribution, and cytogenetics 

of Purshia, Cowania, and Fallugia (Rosoideae, 

Rosaceae). In: A.R. Tiedemann and K.L. 

Johnson, comps. Proceedings, Research and 

management of bitterbrush and cliffrose in 

western North America. GTR-INT-152. U.S. 

Department of Agriculture, Intermountain Forest 

and Range Experiment Station. p. 4–24. 

Meyer, S.E. 1989. Warm pretreatment effects on 

antelope bitterbrush (Purshia tridentata) 

germination response to chilling. Northwest 

Science 63: 146–153. 

Mozingo, H.N. 1987. Shrubs of the Great Basin: a 

natural history. University of Nevada Press, 

Reno, NV. 342 p. 
612 

Nord, E.C. 1965. Autecology of bitterbrush in 

California. Ecological Monographs 35: 307–

334. 

Plummer, A.P., D.R. Christensen, and S.B. 

Monsen. 1968. Restoring big game range in 

Utah. Publication 68-3. Utah Division of Fish 

and Game, Salt Lake City, UT. 183 p. 

Righetti, T.L., C.H. Chard, and D.N. Munns. 1983. 

Opportunities and approaches for enhancing 

nitrogen fixation in Purshia, Cowania and 

Fallugia. In: A.R. Tiedemann and K.L. Johnson, 

comps. Proceedings: Research and management 

of bitterbrush and cliffrose in western North 

America. GTR-INT-152. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Forest and Range Experiment Station, Ogden, 

UT. p. 214–224. 

Shaw, N.L. and S.B. Monsen. 1983. Phenology 

and growth habits of nine antelope bitterbrush, 

desert bitterbrush, Stansbury cliffrose, and 

Apache plume accessions. In: A.R. Tiedemann 

and K.L. Johnson, comps. Proceedings: 

Research and management of bitterbrush and 

cliffrose in western North America. GTR-INT152. U.S. Department of Agriculture, Forest 

Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. p. 55–69. 

Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981. 

Viability of seed from thirty-two shrub and forb 

species through fifteen years of warehouse 

storage. Great Basin Naturalist 41: 274–277. 

USDA Forest Service. 1937. Range plant 

handbook. U.S. Government Printing 

Office, Washington, D.C. 512 p. 

Van der Wall, S.B. 1994. Seed fate pathways of 

antelope bitterbrush: dispersal by seed-caching 

yellow pine chipmunks. Ecology 75: 1911-1926. 

Welsh, S.L., N.D. Atwood, L.C. Higgins, and S. 

Goodrich. 1987. A Utah flora. Great Basin 

Nturalist Memoirs 9. Brigham Young University 

Press. 894 p. 
613

Quercus havardii Rydb. Havard shin oak 

FAGACEAE 

Synonyms: None 

James E. Nellessen 

General Description.—Havard shin oak is also 

known as just Havard oak, shin oak or shinnery 

oak, but the names sand shinnery oak, sand scrub 

oak, panhandle oak, and midget oak have also 

been applied to it (McCoy 1981). Peterson and 

Boyd (1998) supply a fairly comprehensive review 

of this species. Havard shin oak grows to 1 m tall 

at maturity, but hybrids with other white oak 

species may attain greater height. It forms dense 

thickets because of an extensive underground 

system of horizontal rhizomes. The rhizomes are 

usually 3 to 10 cm in diameter but may be as thick 

as 15 cm. Most of the rhizomes and roots occur in 

the upper 60 cm of soil, but the plant is known to 

root to depths of 9 m. The bark is gray becoming 

papery scaly to rough with age. The twigs are 

brown to grayish, smooth to densely hairy with 

short yellowish or grayish hairs. The buds are dark 

red-brown, semi-globose, and lightly hairy. The 

leaves are simple, alternate, leathery, somewhat 

variable in shape, oblong, ovate, or lanceolate in 

outline, with rounded shallow lobes, although 

sometimes they may be more coarsely toothed 

than lobed (the teeth are not bristle-tipped), 

lustrous green and often smooth above, sometimes 

with hairy patches, densely yellowish to brownish 

green felty-hairy below, the veins prominent. The 

leaves range in length from 1.5 to 10 cm, and in 

width from 1.2 to 5.5 cm, with occasionally curled 

or wavy margins (Carter 1997, Correll and 

Johnston 1970, Flora of North America 1997, 

Great Plains Flora Association 1986, Martin and 

Hutchins 1980). Havard shin oak is a member of 

the white oak group, (i.e. it does not bear bristle 

tips on the leaves and the fruits mature in the first 

year). There are two varieties: Q. havardii var. 

havardii and Q. havardii var. tuckeri Welsh 

(Kartesz 1994). 

Range.—This species occurs from western 

Oklahoma, and western Texas, into the 

southeastern portion of New Mexico, with 

outlying populations elsewhere in these states and 

as far west as Utah and Arizona (Great Plains 

Flora Association 1986, Peterson and Boyd 1998). 

Havard shin oak has recently been documented 

from Kansas for the first time (Freeman 2000). 

The elevational range of habitats in which shin oak 

may occur is from 500 m to 1,525 m. Variety 

tuckeri occurs in the Four Corners area (Utah and 

Arizona populations); these populations appear to 

be hybrids between shin oak and Q. turbinella

Greene (scrub live oak) and Q. gambelii Nutt. 

(Gambel oak) (Flora of North America 1997). 

Community Ecology.—Havard shin oak 

historically covered an estimated 5 to 7 million 

acres within the southern Great Plains (Peterson 

and Boyd 1998). Of this, 1 million acres occurred 

in Oklahoma, 1.5 million acres in New Mexico, 

and 3.5 million acres in Texas. About 30 percent 

of the acreage in Texas has been converted to 

cropland and grassland, and about 10 percent has 

been converted in Oklahoma and New Mexico. 

Shin oak occurs on deep sandy soils and plains, 

including sand dunes, and is the major shrub 

species comprising Plains-Mesa Sand Scrub 

vegetation in southeastern New Mexico and 

similar sandy soil habitats in western Texas 

including portions of the Llano Estacado ("staked" 

plains) (Dick-Peddie 1993). The shin oak or 

shinnery vegetation series variously places Havard 

shin oak in dominance with Schizachyrium 

scoparium (Michx.) Nash (little bluestem), 

Artemisia filifolia Torr. (threadleaf sagebrush), 

Andropogon hallii Hack. (sand bluestem), Aristida 

purpurea Nutt. (purple three-awn), Bothriochloa 

saccharoides (Sw.) Rydb. (silver beardgrass), 

Yucca campestris McKelvey (plains yucca), 
614 

Atriplex canescens (Pursh) Nutt. (four-wing 

saltbush), Sporobolus airoides (Torr.) Torr. (alkali 

sacaton), and Eriogonum annuum Nutt. (annual 

buckwheat). But there are a wide variety of other 

shrubs, forbs and grasses comprising numerous 

other vegetation series within the greater shinnery 

community. The tall grasses such as sand 

bluestem, big bluestem (Andropogon gerardii

Vitmann), and Indiangrass [Sorghastrum nutans

(L.) Nash] have been reduced in abundance 

through overgrazing of Havard shin oak 

communities. Havard shin oak itself may remain 

stable or also decrease with overgrazing, the cover 

of grass and Havard shin oak often being 

positively correlated. Stands of Havard shin oak on 

sites with shallow water tables, such as in Chavez 

County New Mexico, may contain cottonwoods. 

Hybrids with other oak species, such as Q. stellata

Wang (post oak), occur and may be found within 

typical Havard shin oak stands, the hybrids often 

attaining greater height (2 to 3 m), and stand out as 

readily visible clumps from the surrounding 

shorter plants. These hybrids tend to have 

intermediate characteristics such as larger leaves 

and smaller acorns. These hybrids tend to be more 

common within the eastern portion of Havard shin 

oak’s range (Correll and Johnston 1970, Flora of 

North America 1997, Peterson and Boyd 1998). 

Havard shin oak also hybridizes with Q. mohriana

Buckley ex Rydb. (Mohr shin oak) that occurs 

within the rocky “breaks” area of the Texas 

panhandle. Hybrids are intermediate in 

characteristics and habitat location (Muller 1952). 

Autoecology.—The plant primarily spreads by its 

underground rhizomes to form large clonal clumps 

that may range in size from as small as 2 to 15 m 

in diameter to as large as 0.8 ha. Isozyme 

electrophoresis identified 37 clones/ha, larger 

clones being free of interclonal growth, and the 

sample population was outcrossing based on 

Hardy-Weinberg expected genotype ratios (Mayes 

and others 1998). There are no data on horizontal 

growth rates but they are believed to be slow. The 

plants are long-lived, estimated at hundreds to 

thousands of years (Peterson and Boyd 1998). The 

rootstocks are massive, comprising the bulk of the 

plant mass (73 percent); the above-ground shoots 

are merely small side shoots to the below ground 

parts. Above-ground biomass has been measured 

at 1,821 kg/ha (Sears and others 1986) and annual 

production has been estimated to be anywhere 

from 550 to 4,500 kg/ha depending on the site 

(reviewed in Peterson and Boyd 1998). The roots 

(which are mycorrhizal) and rhizomes absorb 

considerable amounts of water during wet weather 

with up to 50 percent of their mass being water. 

This gives the plant a great ability to survive 

drought. During spring drought it may not leaf out 

or may defoliate and leaf out later once water 

becomes available. It generates more negative leaf 

water potentials, -25 to -32 bars, than many other 

species in the surrounding community. Soils under 

shin oak, in addition to being sandy, are low in 

nitrogen (0.017 percent), low in organic matter 

(0.36 percent) neutral to slightly alkaline in pH, 

and low in clay and carbonates (a caliche layer is 

inhibitory). Havard shin oak has been reported to 

have allelopathic effects on other plant species. 

Reproduction.—Shin oak flowers in April to 

May. As with other oaks, male flowers are born in 

pendulous catkins, the female flowers solitary or a 

few clustered within leaf axils. The male catkins 

are 2 to 3 cm long and densely flowered. The 

female catkins are 3 to 7 mm long with one to five 

flowers. For a small shrubby oak, the acorns are 

quite large, 12 to 25 mm long, up to about 20 mm 

wide, brown, ovoid to oblong in shape. They are 

born solitary or in pairs and the cup, 10 to 12 mm 

deep, covers about a third to a half of the fruit. The 

fruits mature from mid-July into September during 

the first year of development on the current 

season’s branches (Carter 1997, Correll and 

Johnston 1970, Flora of North America 1997, 

Great Plains Flora Association 1986, Martin and 

Hutchins 1980). Acorns of the white oak group, 

show little or no dormancy, and will germinate 

immediately (Young and Young 1992). Acorn 

moisture content for this group cannot drop below 

30 to 50 percent otherwise seed viability will be 

lost. Within a particular clone, acorn crops 

generally only occur every 3 to 10 years. Within a 

larger landscape scale area containing many 

different clones, at least one or a few clones will 

have acorns. Most acorns are attacked by weevils 

and will decompose quickly. There is little 

reproduction by seed because of the short lifespan 

of the acorn, the need for good environmental 

conditions for germination and establishment at 

the time of acorn drop, attack by weevils, and 

decomposers. 

Growth and Management.—For purposes of 

agriculture and livestock, shin oak has generally 

been managed as an undesirable species. Havard 

shin oak has sometimes been classified as an 

increaser under livestock grazing, but reports do 

not substantiate this (reviewed in Peterson and 

Boyd 1998). Its poor reproduction by seed restricts 
615

its spread primarily to the slow growing rhizome 

system. An estimated 100,000 acres of land in 

New Mexico have been treated with the herbicide 

tebuthiuron for brush control. Tebuthiuron 

gradually kills the plant over a 3 to 4 year period. 

Other chemicals that have historically been used 

on Havard shin oak are 2,4-D, benzoic acids, 

2,4,5-T, and picloram. Tebuthiruron should not be 

used in drought years, and the treated land should 

not be summer grazed for 2 years. Various 

techniques of rotational and continuous grazing in 

shin oak communities have been tested with mixed 

results for livestock productivity. But rotational 

grazing is best for overall ecosystem management 

including maintenance of proper native grass 

covers, and management of the lesser prairie 

chicken and perhaps other wildlife. Goats have 

been used to control shin oak with reasonable 

success, although goats eating it continuously will 

lose weight. In addition to livestock use and direct 

agricultural conversion of shin oak habitat, another 

reason for shin oak management concerns the boll 

weevil (Anthonomus grandis Boheman). This 

cotton pest is known to overwinter in oak leaf litter 

and to reinfest cotton fields for up to a half mile 

away (reviewed in Peterson and Boyd 1998). 

Above-ground shoots may live 11 to 15 years. 

Shoot regeneration is rapid, 30 to 60 cm within the 

first year, when above ground tissues are removed 

mechanically or through fire. The response to fire 

is variable and dependent on timing of the fire. 

Burning reduced shin oak cover by at least 25 

percent, with the greatest reduction, 50 percent, 

following spring burns (Boyd and Bidwell 2002). 

Winter and annual fires increased rhizomatous 

grass cover, and although oak cover was reduced, 

regrowth resulted in greater stem densities (e.g. 

increasing from 20 stems/ha to 40 stems/ha). 

Benefits.—The sandy soils that Havard shin oak 

grows in are susceptible to wind erosion. The 

extensive underground rootstock system and dense 

above ground shoots make it an important soil 

stabilizer. Oak leaves can be good forage for sheep 

and goats, but are less utilized by cattle. 

Consumption of too much, especially young spring 

shoots, can be toxic and result in illness or even 

death for cattle and lambs (Gay and Dwyer 1998, 

Warnock 1974). As with many oak species, 

toxicity is due to phenolics and tannins. Later in 

the growing season cattle will utilize it in small 

and varying degrees. The crude protein content of 

foliage is about 9 percent. The plants are browsed 

by deer, pronghorn antelope, and southern plains 

woodrats. Phenolic content (15 to 20 percent) of 

catkins and buds were higher in livestock grazed 

plots than ungrazed plots. Crude protein and other 

nutritional qualities did not differ between 

treatments (Boyd and others 2001). The acorns are 

eaten by a variety of wildlife and are a staple food 

for lesser prairie chickens (Tympanuchus 

pallidicinctus Ridgeway) in the fall and winter 

seasons. Havard shin oak communities are 

important breeding and nesting habitats for the 

lesser prairie chicken. Prescribed fire in shin oak 

habitats can improve grass and forb cover and 

associated insect abundance for prairie chickens 

but at least a 2 to 3-year recovery period, no 

burning in spring, and burning in patches are 

necessary (Boyd and Bidwell 2001). The rare sand 

dune lizard (Sceloporus arenicolus Degenhardt 

and Jones), a threatened species in New Mexico, is 

restricted to Havard shin oak communities. Both 

Native Americans and early Spanish colonists 

would eat the acorns raw or baked. A flour can be 

made from the large acorns by first boiling them, 

then roasting, chopping into pieces, drying for 30 

minutes, then passing them through a grinder a 

couple of times (Warnock 1974). 

References

Boyd, C.S. and T.G. Bidwell. 2002. Effects of 

prescribed fire on shinnery oak (Quercus 

havardii) plant communities in western 

Oklahoma. Restoration Ecology 10(2): 324-

333. 

Boyd, C.S. and T.G. Bidwell. 2001. Influence of 

prescribed fire on lesser prairie chicken habitat 

in shinnery oak communities in western 

Oklahoma. Wildlife Society Bulletin 29(3): 

938-947. 

Boyd, C.S., L.T. Vermeire, T.G. Bidwell, and R.L. 

Lochmiller. 2001. Nutritional quality of 

shinnery oak buds and catkins in response to 

burning or herbivory. Southwestern Naturalist 

46(3): 295-301. 

Carter, J.L. 1997. Trees and Shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Correll, D.S. and M.C. Johnston. 1970. Manual 

of the Vascular Plants of Texas. Texas Research 

Foundation, Renner, TX. 1,881 p. 

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation: Past, Present and Future. 
616 

University of New Mexico Press, Albuquerque, 

NM. 244 p. 

Flora of North America Editorial Committee. 

1997. Flora of North America, Vol 3. 

Magnoliophyta: Magnoliidae and Hamamelidae. 

Flora of North America Editorial Committee, 

Oxford University Press, New York. 590 p. 

Freeman, C.C. 2000. Vascular plants new to three 

states in the central United States. Transactions 

of the Kansas Academy of Science 103(1-2): 51-

54. 

Gay, C.W., Jr. and D.D. Dwyer. 1998 Reprint. 

New Mexico Range Plants. Cooperative 

Extension Service Circular 374. Revisions by: 

C. Allison, S. Hatch, and J. Schickedanz. New 

Mexico State University, Las Cruces, NM. 

84 p. 

Great Plains Flora Association. 1986. Flora of the 

Great Plains. University Press of Kansas, KS. 

1,392 p. 

Kartesz, J.T. 1994. A Synonymized Checklist of 

the Vascular Flora of the United States, Canada, 

and Greenland, Vol. 1, 2nd Edition. Biota of 

North America Program of the North Carolina 

Botanical Garden. Timber Press, Portland, OR. 

622 p. 

Martin, W.C. and C.E. Hutchins. 1980 (reprinted 

2001). A Flora of New Mexico, Vol. 1. Bishen 

Singh Mahendra Pal Singh, India and Koeltz 

Scientific Books, Germany. 1,276 p. 

Mayes, S.G., M.A. McGinley, and C.R. Werth. 

1998. Clonal population structure and genetic 

variation in sand-shinnery oak, Quercus 

havardii (Fagaceae). American Journal of 

Botany 85(11): 1609-1617. 

McCoy, D. 1981. Roadside Trees and Shrubs of 

Oklahoma. University of Oklahoma Press, 

Norman, OK. 116 p. 

Muller, C.H. 1952. Ecological control of 

hybridization in Quercus; A factor in the 

mechanism of evolution. Evolution 6: 147-161. 

Peterson, R.S. and C.S. Boyd. 1998. Ecology and 

Management of Sand Shinnery Communities: A 

Literature Review. General Technical Report 

RMRS-GTR-16. United States Department of 

Agriculture Forest Service, Rocky Mountain 

Research Station, Fort Collins, CO. 44 p. 

Sears, W.E., C.M. Britton, D.B. Wester, and R.D. 

Pettit. 1986. Herbicide conversion of a sand 

shinnery oak (Quercus havardii) community: 

effects on biomass. Journal of Range 

Management 39: 399-403. 

Warnock, B.H. 1974. Wildflowers of the 

Guadalupe Mountains and the Sand Dune 

Country, Texas. Sul Ross State University, 

Alpine, TX. 176 p. 

Young, J.A. and C.G. Young. 1992. Seeds of 

Woody Plants in North America. Dioscorides 

Press, Portland, OR. 407 p. 
617

Randia aculeata L. white indigo berry

RUBIACEAE 

Synonyms: Randia mitis L. 

 Gardenia randia Sw. 

 Randia latifolia Lam. 

 Randia aculeata L. var. mitis (L.) Griseb. 

John K. Francis 

 

General Description.—White indigo berry is a 

shrub or small tree from 2 to 6 m in height. It is 

spiny and may have one to several stems. White 

indigo berry has smooth to slightly fissured gray 

bark, opposite, often horizontal, branches, a thin 

crown, and rough appearance. The plant is 

supported by an extensive tap and lateral root 

system. The roots have a corky, ivory colored bark 

and are stiff and woody. The spines are paired and 

the foliage is crowded on the ends of short lateral 

twigs. The leaves are almost stalkless and entire, 

ovate, obovate, or orbicular. They are 1 to 6 cm 

long and 0.5 to 3 cm broad, slightly thickened, 

shiny green above and light green below. The 

solitary flowers are small, white, funnel-shaped, 

with a five-lobed corolla, and located on very 

short lateral spurs. The 6- to 13-mm, globose, 

white fruits (or ivory with a salmon blush), with a 

brittle shell-like skin, have a blue or black pulp 

and five to 10 round, flattened seeds (Howard 

1989, Liogier 1997, Little and others 1974). There 

are populations and transition populations with 

and without spines, with large leaves and small 

leaves, and with large and small fruits. These gave 

rise to variety designations that are no longer 

recognized (Howard 1989). White indigo berry is 

also known as box-brier, inkberry, prickle bush, 

fishing rod, five fingers, goat horn, tintillo, tintero, 

palo de cotorra, árbol de navidad, sota-caballo, 

resuelesuele, café cimmarrón, espino cruz, 

crucete, crucito, maíz tostado, raboe, peetschkitam, cabai nache, croc-à-chien, bois-lance, ti 

coco, and petit coco (Howard 1989, Little and 

others 1974). 

Range.—White indigo berry is native to southern 

Florida, Bermuda, the Bahamas, the Caribbean 

Islands, and from Mexico through Central 

America and South America as far south as 

Colombia (Little and others 1974, Smithsonian 

Institution 2001). It is not known to have 

naturalized outside its native range. 

Ecology.—White indigo berry grows in most 

types of soils including excessively drained and 

well drained, but not poorly drained soils. It 

inhabits areas that receive from about 700 to 2200 

mm of annual precipitation from near sea level to 

over 600 m of elevation in Puerto Rico. In Florida, 

white indigo berry grows in unburned pinelands 

and along the margins of coastal hammocks 

(Nelson 1996). In Puerto Rico, the species grows 

in dry and moist forests, more commonly over 

limestone rocks, but also over igneous rocks and 

ultramafics (serpentine). Because white indigo 

berry is moderately intolerant of shade, it must 

grow in openings or under low-density forest. The 

species invades abandoned land, but not quickly, 

and because of slow growth, is not common in 

early secondary forest following cultivation. 

However, white indigo berry does hang on 
618 

tenaciously after disturbance and is prominent in 

secondary forests arising after logging or partial 

clearing for pasture. Plants are usually well 

disbursed and seldom, if ever, form thickets. 

Reproduction.—White indigo berry blooms and 

fruits irregularly throughout the year. Fruits (n = 

73) collected in Puerto Rico weighed an average 

of 0.472 + 0.014 g/fruit. Fourteen of the fruits 

contained from three to 14 seeds and averaged 8.2 

seeds. The air dry seeds from these fruits averaged 

0.0264 + 0.0003 g/seed or 38,000 seeds/kg. Sown 

in commercial potting mix, 25 percent germinated 

between 46 and 168 days after sowing. 

Growth and Management.—Weaver (1990) 

measured 12 plants over a 5-year period in closed 

low forest in St. John, U.S. Virgin Islands and 

found the vary slow diameter growth rate of 0.06 

+ 0.02 cm/yr. A 1-m tall sapling in a moist area of 

Puerto Rico with a basal diameter of 2 cm had 10 

growth rings (author’s observation). Ornamental 

plants also grow slowly (Workman 1980). Seeds 

for nursery production should be cleaned by 

maceration and wet sieving before sowing. 

Production of seedlings in containers is 

recommended. Because growth is slow, a year or 

more in the nursery is required followed by 

protection from weeds in the field. 

Benefits.—White indigo berry has been used as an 

ornamental to a limited extent in Florida and, 

because it is native and resists drought and 

hurricanes, it is currently being recommended for 

planting (Dade County 2001). It is available as 

nursery-grown plants, and plants from the wild 

can be successfully dug up and transplanted. The 

fruits are edible although of poor flavor 

(Workman 1980). White indigo berry provides 

food and nesting sites for birds (Vélez and van 

Overbeek 1950) and nectar for butterflies (Dade 

County 2001). The wood is hard and heavy and 

useful for fuel. The shrub is sometimes decorated 

for a Christmas tree, hence the Spanish name, 

árbol de navidad. Other names (tintillo and 

inkberry) arose from the former use of the berries 

for dye and for ink (Little and others 1974). The 

plant is also frequently formed into bonsai plants. 

The fruits are used in herbal medicine to control 

dysentery. An unspecified part of the plant is used 

to control fever, and the latex is reported to 

effectively stop bleeding (Liogier 1990). 

References 

Dade County. 2001. Some trees and shrubs native 

to South Florida. http://www.fnps.org/dade/ 

pubs/SomeTrees.html. 6 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Part 3. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p.

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. U.S. Department of 

Agriculture, Agriculture Handbook 449. 

Washington, DC. 1,024 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc. Sarasota, FL. 

391 p. 

Smithsonian Institution. 2001. Centers of plant 

diversity: Columbian Central Massif. 

http://www.nmnh.si.edu/botany/projects/cpd/sa/

sa29.htm. 14 p. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Río Piedras, PR. 497 p. 

Weaver, P.L. 1990. Tree diameter growth rates in 

Cinnamon Bay Watershed, St. John, U.S. Virgin 

Islands. Caribbean Journal of Science 26(1-2): 

1-6. 

Workman, R.W. 1980. Growing native. The 

Sanibel-Captiva Conservation Foundation, Inc., 

Sanibel, FL. 137 p. 
619

Rauvolfia viridis Willd. ex Roemer & J.A. Schultes bitter bush 

APOCYNACEAE 

Synonyms: Rauvolfia lamarckii A. DC. in DC. 

 Rauvolfia latifolia A. DC in DC. 

 Rauvolfia lanceolata sensu Griseb. 

John K. Francis 

General Description.—Bitter bush, also known 

as bellyache bush, balsam, sassafras, milk tree, 

and ti bois-lait, is a branchy shrub to 2.5 m in 

height (Howard 1989, Liogier 1995). The species 

rarely becomes a small tree to 4 m in height 

(Liogier 1995) and 7 cm in diameter at breast 

height (Weaver 1990). Usually, there are multiple 

stems, as suckers arising from lateral roots, 

especially after fire or other disturbance. The root 

system is dominated by lateral roots, which 

support sinker roots and a moderate amount of 

fine roots. The ovate or ovate-elliptic leaves are 5 

to 15 cm long by 3 to 5 cm broad, and grow in 

whorls of four or rarely three leaves. The severalto many-flowered inflorescences are usually 

terminal. The tiny flowers are tubular and white. 

The fruits are spherical or slightly flattened and 5 

to 8 mm in diameter. Each contains two seeds 

(Howard 1989, Liogier 1995). 

Range.—According to Howard (1989) and 

Liogier (1995) bitter bush is native to Hispaniola, 

Puerto Rico, the Virgin Islands, the Lesser 

Antilles, Trinidad, Margarita, and Venezuela. 

There are also records for Cuba (Martínez-Pérez 

and others 1997), Navassa Island (New York 

Botanical Garden 2001), Costa Rica 

(Departamento de Botánica 1999), and Colombia 

(Missouri Botanical Garden 2001). 

Ecology.—In Puerto Rico, bitter bush grows in 

dry and moist forests that receive from 750 to 

about 1700 mm of annual precipitation. Usually 

rocky or otherwise poor sites are colonized in 

areas of both igneous and sedimentary (including 

limestone) rocks. The species grows from near sea 

level to about 400 m in elevation. Bitter bush 

requires full or partial sunlight. It grows in open, 

disturbed areas and low basal area, secondary and 

remnant forest. It withstands competition from 

herbs and shrubs well but eventually succumbs to 

competition from tall grass swards and closed 

forests. Although never abundant, the species 

becomes more common in overgrazed areas. 

Reproduction.—A sample of bitter bush fruits 

collected in Puerto Rico weighed an average of 

0.124 + 0.005 g/fruit. Air-dried seeds separated 

from these fruits averaged 0.0247 + 0.0004 g/seed 

or 40,500 seeds/kg. Sown in commercial potting 

mix without any pretreatment, 5 percent 

germinated after 11 months. Birds that eat the 

fruits disburse the seeds. 

Growth and Management.—Growth of bitter 

bush is slow. The author estimated height growth 

of young shrubs from sprouts at 0.5 m/year. 

Weaver (1990) measured the diameter growth rate 

of two old small trees and found an average of 

only 0.01 cm/yr. No planting or management 

experience has been reported. 

Benefits.—The roots of bitter bush were analyzed 

for alkaloids. Ajmalidine, a-yohimina, reserpinine, 

quebrachidine, and yohimbina were isolated. 

Reserpine, a sedative present in other Rauvolfias, 

was not found (Martínez Pérez and others 1997). 

Bitter bush furnishes minor food and cover for 

birds and other wildlife. With other dry forest 

vegetation, it protects the soil from erosion. 
620 

References 

Departamento de Botánica. 1999. Especies de 

plantas nuevas para Costa Rica. Departamento 

de Botánica, Inventario Biótico. 

http://www.minae.go.cr/estrategia/estudio/planta

s1.htm. 4 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Part 3. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

617 p. 

Martínez Pérez, J., A. Bello Alarcío, O. Montero 

Tápanes, and V. Fuente Fiallo. 1997. Bases 

débiles minoritarias de la raíz de Rauvolfia 

viridis Roem. et Schult. Revista Cubana Farm 

31(3): 188-191. 

Missouri Botanical Garden. 2001. Current 

specimen list for Rauvolfia viridis. 

http://mobot.mobbot.org/cgi-bin/search_vast. 

1 p. 

The New York Botanical Garden. 2001. Flora of 

Navassa Island: vascular plant checklist. 

http://www.nybg.org/bsci/hcol/navassa/checklist

vasc.html. 9 p. 

Weaver, P.L. 1990. Tree diameter growth rates in 

Cinnamon Bay Watershed, St. John, U.S. Virgin 

Islands. Caribbean Journal of Science 26(1-2): 

1-6. 
621

Rhododendron macrophyllum D. Don ex G. Don Pacific rhododendron

ERICACEAE 

Synonyms: Hymenanthyes macrophyllum (D. Don ex G. Don) H.F. Copel 

Rhododendron californicum Hook. 

Christopher Ross 

General Description.—Pacific rhododendron, 

also known as coast rhododendron and California 

rosebay, is a coarse-branched evergreen shrub to 4 

m tall. The alternate leathery ovate to obovate 

leaves are 7 to 15 cm long and 3 to 6 cm wide. 

Flowers are showy and bell shaped, about 4 cm 

long, with rose pink to purple corolla, five petals, 

five sepals, and 10 stamens (Wallace 1993).

Range.—Pacific rhododendron occurs near the 

Pacific coast from British Columbia south to 

Monterey, California, and in the coastal and 

Cascade mountain ranges of Oregon, Washington, 

and California (Hitchcock and others 1959, 

Wallace 1993). It occurs in redwood, Douglas fir, 

yellow pine, and mixed evergreen forests fron sea 

level to over 1,000 m elevation (CalFlora 2000). 

Ecology.—Pacific rhododendron grows in acidic 

(pH 5.5) to neutral fine textured soils (USDA 

2002). Pacific rhododendron, like many others of 

the genus, is highly toxic to humans and animals. 

The poison is a neurotoxin, andromedotoxin, or 

grayanotoxin, which has occasionally caused large 

livestock losses (Kingsbury 1964). About 30 

diterpine, polyhydroxylated cyclic hydrocarbons 

are found in rhododendrons and related species 

(Harborne and Baxter 1993). Sucking nectar from 

flowers has caused illness in humans, as has 

consumption of rhododendron honey (Alberta 

Government 2002). The species has recently been 

identified as a host of the algae Phytophthora 

ramorum responsible for sudden oak death in 

California, but the effects are not yet known 

(University of California Cooperative Extension 

2002). A variety of insect and fungi may attack 

rhododendrons, including several caterpillars such 

as cutworms, loopers, and leafrollers (Foss and 

Antonelli 1999).

 

Reproduction.—Pacific rhododendron flowers are 

insect pollinated. Fruits are oblong capsules that 

split along the sides to release minute seeds (4400 

to 12,500/g) after ripening. Germination occurs 

with no pretreatment, but requires light (Young 

and Young 1992). Vegetative reproduction by 

crown sprouting is common after fire, logging, and 

other disturbance. Propagation is by seeds or 

cuttings.

Fire Effects.—Rhododendron has been classed as 

a residual shrub (rather than a colonizer) after fire 

(Dyrness 1973). It recovers well but slowly after 

fire, primarily from vegetative sprouting from root 

crowns and stem bases. Fire dramatically reduces 

it initially, and peak abundance is not reached until 

late in the fire recovery process (Halpern 1989). It 

is scarce after severe burns (Dyrness 1973).

Growth and Management.—Pacific 

rhododendron typically occupies drier sites within 

its range (Dyrness 1973). Like some other 

rhododendrons, it may have a mycorrhizal 

relationship that enables it to thrive on lownutrient soils (USDA USFS 2002). The tendency 

to resprout from stem crowns has caused difficulty 

for conifer reforestation (Halverson and others 

1986), so that spring spraying with triclopyr has 

been used for control in some areas (Burrill and 

others 1989). At California’s Kruse Rhododendron 
622 

State Reserve, Pacific rhododendron is considered 

to be an early seral stage after fire or logging, and 

before tanoak, which is being thinned in order to 

set back the succession process and maintain the 

rhododendrons (California State Parks 2002). 

Benefits.—Pacific rhododendron is widely 

available from commercial nurseries and is valued 

for native landscaping purposes. It has served as a 

source of genetic material for many ornamental 

cultural selections. It is also used in erosion 

control (Olson 1974). It has no other common uses 

(USDA 2002). 

References 

Alberta Government. 2002. Rhododendron 

toxicity. http://www.Agric.gov.ab.ca/crops. 

[not paged]. 

Burrill, L.C., W.S. Braunworth Jr., and R.D. 

William, comps. 1989. Pacific Northwest weed 

control handbook. Oregon State University, 

Extension Service, Agricultural 

Communications. Corvallis, OR. 276 p. 

CalFlora. 2000. Information on California plants 

for education, research and conservation. 

Berkeley, California: The CalFlora Database. 

http://www. Calflora.org/ [not paged]. 

California State Parks–Kruse Rhododendron State 

Reserve. 2002. http://cal-parks.ca.gov. 

Dyrness, C.T. 1973. Early stages of plant 

succession following logging and burning in the 

western Cascades of Oregon. Ecology 54: 57-

69. 

Foss, C. and A. Antonelli. 1999. Hortsense. 

http://pep.wsu.edu/hortsense. [not paged]. 

Halpern, C.B. 1989. Early successional patterns of 

forest species: Interactions of life history traits 

and disturbance. Ecology 70: 704-720. 

Halverson, N.M., C. Topik, and R. Van Vickle. 

1986. Plant association and management guide 

for the western hemlock zone: Mt. Hood 

National Forest. R6-ECOL-232A. U.S. 

Department of Agriculture, Forest Service, 

Pacific Northwest Region, Portland, OR. 

 111 p. 

Harborne, J.B. and H. Baxter. 1993. 

Phytochemical Dictionary. Taylor and Francis, 

London and Washington, D.C. 791 p. 

Hitchcock, C.L., R. Cronquist, and M. Ownbey. 

1959. Vascular plants of the Pacific northwest. 

Part 4: Ericaceae through Campanulaceae. 

University of Washington Press. Seattle, WA. 

510 p. 

Kingsbury, J.M. 1964. Poisonous plants of the 

United States and Canada. Prentice Hall, Inc. 

Englewood, NJ. 626 p. 

Olson, D.F., Jr. 1974. Rhododendron L. 

rhododendron. In: C.S. Schopmeyer, ed. Seeds 

of woody plants in the United States. 

Agricultural Handbook 450. U.S. Department 

of Agriculture, Forest Service, Washington, 

D.C. p. 709-712. 

University of California Cooperative Extension. 

2002. Sudden oak death. http://cemarin. 

ucdavis.edu. 2 p. 

USDA NRCS. 2002. Plants Source and Reference. 

http://plants.usda.gov. [not paged]. 

U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire 

Sciences Laboratory. 2002. Fire Effects 

Information System, http://www.fs.fed.us/ [not 

paged]. 

Wallace, G. 1993. Rhododendron. In: J.C. 

Hickman, ed. 1993. The Jepson Manual: 

higher plants of California. U.C. Press, 

Berkeley and Los Angeles, CA. 1,400 p. 

Young, J.A. and C.G. Young. 1992. Seeds of 

woody plants of North America. Disceroides 

Press, Portland, OR. 407 p. 
623

Rhododendron maximum L. rosebay rhododendron 

ERICACEAE 

Synonyms: none 

Barton D. Clinton 

General Description.—Rosebay rhododendron, 

sometimes known as ‘great laurel’, is an 

understory evergreen shrub. It ranges in height 

from 2 to 8 m, and can occasionally reach 

diameters of 20 to 25 cm. Leaves are 

schlerophyllous, simple, alternate, and oblong (10 

to 30 cm long, 5 to 8 cm wide). It retains its waxy, 

deep-green leaves for up to 8 years, but once shed 

are slow to decompose. It produces large, showy, 

white to purple flowers each June. 

Range.—Worldwide, there are over 800 species in 

the genus Rhododendron. Most of the 26 North 

American species occur in the East along the 

mountain chain stretching from Nova Scotia in the 

north to Georgia and Alabama in the south, but are 

concentrated in the southern Appalachians (Dobbs 

1998). Rosebay rhododendron is the most 

frequently occurring and dominant species of this 

genus in the southern Appalachian region 

(Swanson 1994), and occurs occasionally on mesic 

hill-slopes throughout the upper Piedmont 

Crescent of the Southeastern United States. 

Ecology.—Approximately 1.2 million ha in the 

southern Appalachians are occupied by this 

species (Dobbs 1995) where it dominates the 

understory. This species has historically been 

confined to riparian areas and other mesic sites but 

takes advantage of disturbed areas where it is 

present to advance onto sub-mesic sites. It prefers 

deep well-drained acid soils high in organic matter 

where it produces a thick, peat-like humus. It 

prefers low to medium light conditions for 

optimum carbon gain, and has a tremendous 

capacity for avoiding cavitation during freeze-thaw 

cycles (Lipp and Nilsen 1997). Where extensive 

overstory mortality has eliminated most of the 

overstory, this species forms a thick and 

continuous subcanopy known locally as ‘laurel 

slicks’ or ‘laurel hells’. Rosebay rhododendron is 

an important structural and functional component 

of southern Appalachian forest ecosystems. What 

isn’t clear is whether or not we are in a period of 

advancement or retreat for this species. For 

example, on poorly drained sites on ridge or upper 

slope positions, large areas of rosebay 

rhododendron, particularly at the high elevations,

have recently died out presumably due to the 

phytophthora fungus, or due to recent prolonged 

periods of below-average precipitation. Yet, 

rosebay rhododendron now occupies sites that 

historically were free of evergreen understory. 

There are still important questions to be answered 

regarding this species to completely understand its 

role in forest understories. 

Reproduction.—Rosebay rhododendron is clonal. 

It is capable, however, of reproducing both 

vegetatively and sexually. It reproduces 

vegetatively through a process called ‘layering’ 

where it produces roots from aboveground woody 

parts when in contact with the forest floor. The 

fruit is produced from showy flowers from March 

to August. The fruit is an oblong capsule that 

ripens in the fall, and splits along the sides soon 

after ripening to release large numbers of minute 

seed (approx. 400 per capsule) (Schopmeyer 

1974). Microsite requirements for seed 

germination are relatively specific (e.g., high in 

organic matter such as rotting logs); hence, the 

majority of reproduction is vegetative resulting in 

a clonal distribution. 

Growth and Management.—Seeds from rosebay 

rhododendron are minute and it is estimated that 

approximately 11 million are contained in 1 kg. 

Commercial seed production is generally from 

cultivated hybrids. Seeds from wild sources are not 

commonly sold commercially. Rosebay 

rhododendron is a slow-growing shrub and has a 

very high sprout potential. If mechanical removal 

is attempted in the case of forest management, 
624 

extremely high densities are attained by this 

species in a manner of a few years. Prescribed fire 

has also been used to control this species but with 

limited success (Clinton and Vose 2000). 

Benefits.—Rosebay rhododendron is a striking 

and aesthetically pleasing feature of mesic 

southern Appalachian forests. It is one of the 

largest and hardiest rhododendrons grown 

commercially. Several cultivars with white to 

purple flowers have been selected for the 

horticultural trade (Brown and Kirkman 1990). 

Where it occurs naturally, it produces a showy, 

white, pink, or light purple flower primarily in 

June, but occurs from March into August. Rosebay 

rhododendron maintains deep-green foliage year 

round. This species affords protection to steep 

watersheds and shelter for wildlife. The wood is 

very hard and is occasionally used for specialty 

wood products. 

 

Detrimental Effects.—For all its prized qualities 

as a naturally occurring component of the 

landscape or as plantings in residential and 

commercial landscaping, rosebay rhododendron 

can have an inhibitory effect on regeneration of 

other plant species. There is some evidence to 

suggest that due to fire suppression and the 

absence of other cultural activities (i.e., mountainland grazing), this species has advanced beyond 

the mesic forest sites into sub-mesic understories 

(Dobbs 1998). The significance of this movement 

onto previously unoccupied sites centers around 

the impacts of rosebay rhododendron on plant 

succession (Clinton and Vose 1996) and resource 

availability (Nilsen and others 2001). Rosebay 

rhododendron is associated with reduced woody 

and herbaceous seedling abundance throughout its 

range, and hence poses a serious impediment to 

the production of wood products. The 

mechanism(s) by which rosebay rhododendron 

reduces seedling survival has been the subject of 

much debate. Possible sources of inhibition 

include allelopathy, competition for resources 

including light, physical and chemical attributes of 

the forest floor and soil, and interactions between 

some or all sources (Nilsen and others 1999, 

Nilsen and others 2001). 

 

References 

Brown, C.L. and L.K. Kirkman. 1990. Trees of 

Georgia and Adjacent States. Timber Press, 

Portland, OR. 292 p. 

Clinton. B.D. and J.M. Vose. 1996. Effects of 

Rhododendron maximum L. on Acer rubrum L. 

seedling establishment. Castanea 61(1):38-45. 

Clinton, B.D. and J.M. Vose. 2000. Plant 

succession and community restoration 

following felling and burning in the southern 

Appalachian Mountains. In: W. K. Moser and 

C.F. Moser, eds. Fire and forest ecology: 

innovative silviculture and vegetation 

management. Tall Timbers Fire Ecology 

Conference Proceedings, No. 21. Tall Timbers 

Research Station, Tallahassee, FL. p . 22-29. 

Dobbs, M.M. 1995. Spatial and temporal 

distribution of the evergreen understory in the 

southern Appalachians. Master’s Thesis, 

University of Georgia, Athens, GA. 100 p. 

Dobbs, M.M. 1998. Dynamics of the evergreen 

understory at Coweeta Hydrologic Laboratory, 

North Carolina. PhD Dissertation, University 

of Georgia, Athens GA. 179 p. 

Lipp, C.C. and E.T. Nilsen. 1997. The impact of 

subcanopy light environment on the hydraulic 

vulnerability of Rhododendron maximum to 

freeze-thaw cycles and drought. Plant, Cell 

and Environment 20:1,264-1,272. 

Nilsen, E.T., B.D. Clinton, T.T. Lei, O.K. Miller, 

S.W. Semones, and J.F. Walker. 2001. Does 

Rhododendron maximum L. (Ericaceae) reduce 

the availability of resources above and 

belowground for canopy tree seedlings? 

American Midland Naturalist 145:325-343. 

Nilsen, E.T., J.F. Walker, O.K. Miller, S.W. 

Semones, T.T. Lei, and B.D. Clinton. 1999. 

Inhibition of seedling survival under 

Rhododendron maximum (Ericaceae): could 

allelopathy be a cause? American Journal of 

Botany 86(11):1,597-1,605. 

Schopmeyer, C.S. 1974. Seeds of woody plants in 

the United States. Agricultural Handbook 450. 

U.S. Department of Agriculture, Forest 

Service, Washington, DC. 883 p. 

Swanson, R.E. 1994. A Field Guide to the Trees 

and Shrubs of the Southern Appalachians. The 

Johns Hopkins University Press, Baltimore, 

MD. 399 p. 
625

Rhus copallinum L. shining sumac 

ANACARDIACEAE 

Synonyms: None 

Kristina Connor 

Illustration source: Grelen and Duvall 1966 

General Description.—Shining sumac, also 

known as black sumac, dwarf sumac, flameleaf 

sumac, mountain sumac, shiny sumac, and winged 

sumac, is an upright, deciduous shrub or (rarely) 

small tree from 3 to 6 m tall (Bailey and Bailey 

1976, Odenwald and others 1996). Bark ranges in 

color from light brown to gray to reddish-brown. 

Shoots and twigs are hairy and reddish in color. 

Twigs have conspicuous lenticels. The sparsely 

branched, flat crown is composed of alternate, 

pinnately compound leaves approximately 15 to 30 

cm long, with wings between each of the 7 to 27 

glossy, dark green leaflets (Brickell and Zuk 1996, 

Odenwald and others 

1996, Seiler and Peterson 2001). Leaflets are 

paired, olbong-lanceolate in shape, and from 3 to 

10 cm long (Bailey and Bailey 1976, Krussmann 

1976, Johnson and Hoagland 1999). Buds are 

glabrous, and terminal buds are absent (Synor and 

Cowen [no date]). Sap is sticky and yellow 

(Johnson and Hoagland 1999). 

Range.—Shining sumac grows from southern 

Ontario south along the Coastal Plain to Florida. It 

extends westward into eastern Texas, and inland 

from central Michigan and Wisconsin to Kansas 

and Oklahoma (USDA-ARS [no date], Coladonato 

1992). It is also found in Cuba. Recognized 

varieties are R. copallinum var. latifolia Engl., R. 

copallinum var. lanceolata Gray, and R. 

copallinum var. leucantha (Jacq.) DC (USDAARS [no date], USDA-NRCS 2001). 

Ecology.—Shining sumac is a deciduous, clonal 

shrub. It is an early pioneer species and grows best 

on well drained soils in full sunlight (Coladonato 

1992, Odenwald and others 1996, Seiler and 

Peterson 2001). It can form thickets in abandoned 

fields, along roadsides, in glades and open woods, 

and in waste areas (Tenaglia 2002, Oplin 2001, 

Arborquest 2001). It is difficult to get rid of, once 

established, due to its ability to spread from 

rhizomes (Arborquest 2001). This characteristic 

also makes it well adapted to fire (Coladonato 

1992). Shining sumac will tolerate compacted soil, 

drought, pollution, heavy pruning, and 

transplanting, but the shallow roots make it 

susceptible to uprooting and the stems can break in 

strong wind storms (Arborquest 2001). It can be a 

serious competitor with young pines and 

hardwoods (Coladonato 1992). 

Reproduction.—Shining sumac is a clonal species 

which sprouts from roots and the root crown. It 

also regenerates sexually (Coladonato 1992). From 

May to August, plants produce panicles of small 

yellow-green dioecious flowers that have five 

petals and five sepals (Tenaglia 2002). The 
626 

panicles are pubescent, about 12 to 15 cm long, 

and can be either terminal or axillary (Brickell and 

Zuk 1996, Johnson and Hoagland 1999, 

Krussmann 1976). Brown and Kirkman (1990) 

report the presence of polygamous flowers but 

state that shining sumac is functionally dioecious 

since the pistils of the polygamous flowers abort. 

Thus, both male and female plants are needed for 

good fruit set to occur (Arborquest 2001) The 3- to 

5- mm pubescent fruits are small drupes that 

contain a single nutlet (Coladonato 1992). Fruits 

ripen from August to October. They turn dark red 

when ripe and often remain on the plant 

throughout the winter. The hard-coated seeds 

germinate poorly without pretreatment, but a 1 hr 

soak in sulfuric acid at room temperature can 

result in a 75 percent germinative capacity 

(Brinkman 1974). 

Growth and Management.—Shining sumac is a 

fast growing but short lived clonal shrub, first 

cultivated in 1688 (Brinkman 1974). It is planted 

as an ornamental and used in mass plantings 

because of its brilliant red fall foliage but it can 

spread well beyond desired borders because of its 

ability to sprout from the roots. 

Benefits.—Shining sumac is nonpoisonous to 

humans (USDA-ARS [no date], Seiler and 

Peterson 2001, Synor and Cowen [no date]), and 

the fresh fruits can be used to make a lemontasting beverage. It was also used by Native 

Americans to treat dysentery and mouth sores 

(Tenaglia 2002). Dense thickets of shining sumac 

serve as cover for birds and mammals. The seeds 

are eaten by a variety of birds, while the flowers 

attract butterflies. Deer and rabbits commonly 

browse the twigs in winter; rabbits also eat the 

bark (Grelen and Duvall 1966, Oplin 2001, 

Arborquest 2001, Coladonato 1992). It is not, 

however, a preferred food and is considered a poor 

to moderately important browse (Coladonato 

1992). However, the seeds, bark and leaves, while 

low in nutritive value, have a high tannin content 

and have been used by the leather industry (Bailey 

and Bailey 1976, Brown and Kirkman 1990, 

Coladonato 1992). Its ability to sprout from the 

roots, coupled with its rapid growth rate, make 

shining sumac a good species to plant for erosion 

control (Arborquest 2001). 

References

Arborquest. 2001. Rhus copallina. 

http://www.arborquest.com. 2 p. 

Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: 

A Concise Dictionary of Plants Cultivated in the 

United States and Canada. McMillan Publishing 

Co., Inc., New York. 1,312 p. 

Brickell, C. and J.D.. Zuk. 1996. The American 

Horticultural Society A-Z Encyclopedia of 

Garden Plants. DK Publishing, Inc., New York. 

1,092 p. 

Brinkman, K.A. 1974. Rhus L. Sumac. In: C.S. 

Schopmeyere, tech. coord. Seeds of woody 

plants in the United States. Agriculture 

Handbkook 450. U.S. Department of 

Agriculture, Forest Service Washington, D.C. 

883 p. 

Brown, C.L. and L.K. Kirkman. 1990. Trees of 

Georgia and adjacent states. Timber Press, 

Portland, OR. 292 p. 

Coladonato, M. 1992. Rhus copallinum. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station Fire Sciences 

Laboratory, Fire Effects Information System 

http://www.fs.fed.us/database/feis. 9 p. 

Grelen, H.E. and V.L. Duvall. 1966. Common 

plants of longleaf pine-bluestem range. Research 

Paper SO-23. U.S. Department of Agriculture, 

Forest Service, Southern Forest Experiment 

Station, New Orleans, LA. 96 p. 

Johnson, F.L. and B.W. Hoagland. 1999. Rhus 

copallinum L. Catalog of the Woody Plants of 

Oklahoma, Oklahoma Biological Survey. 

http://www.biosurvey.ou.edu. 1 p. 

Krüssmann, G. 1976. Manual of Cultivated BroadLeaved Trees and Shrubs. Volume I, A-D. 

Timber Press, Beaverton, OR. 448 p. 

Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope. 

1996. Plants for American Landscapes. 

Louisiana State University Press, Baton Rouge, 

LA. 266 p. 

Oplin. 2001. Dwarf Sumac--Rhus copallina. Ohio 

Public Library Information Network (OPLIN) 

and the Ohio Historical Society (OHS). 

http://www. oplin.lib.oh.us/products/tree. 2 p. 

Seiler, J.R. and J.A. Peterson. 2001.Shining 

sumac. Virginia Tech. Dendrology Webpage 
627

http://www.fw.vt.edu/dendro/dendrology/syllab

us/ivirginica.htm. 1 p. 

Synor, T.D. and W.F. Cowen. [no date]. Ohio 

trees. Rhus--Sumac. The Ohio State University 

College of Food, Agricultural, and 

Environmental Sciences, Extension Bulletin 

700-00. http://www.ag.ohiostate.edu/~ohioline/b700. 3 p. 

Tenaglia, D. 2002. Rhus copallinum. The Missouri 

Flora Website. http://www.missouriplants.com. 

4 p. 

USDA, ARS National Genetic Resources Program. 

Germplasm Resources Information Network- 

(GRIN). [no date]. Online Database. National 

Germplasm Resources Laboratory, Beltsville, 

MD. http://www.ars-grin.gov/npgs/ 

searchgrin.html. 

USDA, NRCS. 2001. The PLANTS database. 

National Plant Data Center, Baton Rouge, LA. 

http://plants.usda.gov. [not paged]. 
628 

Rhus glabra L. smooth sumac 

ANACARDIACEAE 

Synonyms: Rhus borealis Greene 

 Rhus calophylla Greene 

 Rhus canadensis Mill. 

 Rhus carolineana Mill. 

 Rhus cismontana Greene 

Rhus occidentalis (Torr.) Blank. (additional synonyms: Institute of 

Systematic Botany 2003). 

John K. Francis 

Illustration source: USDA-Forest Service collection, 

Hunt Institute 

General Description.—Smooth sumac, also 

known as common sumac, Rocky Mountain 

sumac, red sumac, scarlet sumac, lemonade sumac, 

western sumac, and white sumac, is a deciduous 

shrub or a small tree from 0.5 to 6 m in height and 

1 to 18 cm in stem diameter. It usually has a single 

gray-brown stem but often occurs in clonal 

thickets formed by suckers from horizontal roots. 

The relatively few branches and twigs are thick, 

have a soft pith, and large, horseshoe-shaped leaf 

scars. Stems and branches ooze a milky-resinous 

sap when cut. The alternate leaves are uneven, 

pinnately compound, up to 46 cm long, and 

support seven to 29 leaflets. Leaflets are almost 

stalkless and have serrate edges. (Clark 1973). The 

foliage turns a brilliant red or scarlet during the 

fall. Small, greenish-yellow flowers are grouped in 

tight, 15- to 25-cm terminal panicles. The fruits are 

dry, hairy, red to scarlet drupes that contain one 

hard ovoid, dark-gray nutlet. The fruits have a 

tangy, sour taste (Johnson 1995). Smooth sumac is 

dioecious (male and female plants) (Wyman 

1969). Total above-ground dry biomass may be 

predicted by the equation: dry weight = 1.5130 + 

0.6292 D2

H, where D = diameter at 1 cm above 

groundline, H = total height and r2 = 0.974 

(Reeves and Lenhart 1988). 

Range.—Smooth sumac is native to every state of 

the United States (except Alaska and Hawaii) and 

from Nova Scotia to British Colombia and south 

into northern Mexico (Abrams 1951, Fire Sciences 

Laboratory 2003, Natural Resources Conservation 

Service 2003). However, it is most common in the 

eastern half of United States north of Florida 

(Hutchison 2003). It has been planted widely as an 

ornamental but has not been reported to have 

naturalized outside its native range. 

Ecology.—Smooth sumac prefers rich, welldrained soil but will grow on excessively drained, 

sandy soils and eroded sites without topsoil. In 

much of the Eastern range, it grows as a seral 

species on disturbed sites such as logged areas, 

abandoned fields, neglected pastures, old 

construction sites, strip mines, stream overflow 

areas, roadsides, and fence rows (Hutchison 2003). 
629

In Western areas it grows in stable communities on 

mountains, hillsides, and canyons. The species is 

used as a climax indicator in a number of shrubgrassland communities (Fire Sciences Laboratory 

2003). Smooth sumac usually grows with full sun 

but tolerates partial shade. It grows from low 

elevations in many areas and up to 2,100 m in 

Arizona (Kearney and Peebles 1951). 

Reproduction.—Smooth sumac flowers from 

May to late July, depending on location. Fruits 

ripen in September and October and persist 

through the fall and winter (Fire Sciences 

Laboratory 2003). The flowers are pollinated by 

bees (Rowe and Blazich 2003). Smooth sumac 

produces at least some fruits and seeds nearly 

every year (Fire Sciences Laboratory 2003). There 

are about 2,000 seeds per well-formed panicle 

(Browse-Shrub and Forb Committee 1985), much 

less in some areas and during dry years. Rowe and 

Blazich (2003) report 50,600 to 105,600 fruits/kg 

and 52,800 to 277,200 seeds/kg. Plummer and 

others (1968) found 137,700 seeds/kg in samples 

from Utah. Air-dried fruits collected by the author 

in Utah averaged 0.0202 + 0.0004 g/fruit. Seeds 

separated from them averaged 0.0103 + 0.0002 

g/seed or 97,000 seeds/kg. The seeds are dispersed 

principally by birds and form a soil seed bank 

(Fire Sciences Laboratory 2003). Once individual 

plants are established, horizontal roots sucker to 

form clonal thickets. 

 Nursery plants are usually produced from 

seed, although production from cuttings is also 

done to preserve favorable traits of select strains 

(such as assuring only female plants). Rootstalks 

pulled up from wild stands in the early spring 

when the ground is wet will survive almost 100 

percent if planted properly (Plummer and others 

1968). Seed can be collected in quantity by hand in 

fall or early winter. Seeds may be separated from 

the fruits by rubbing or beating followed by 

screening (Rowe and Blazich 2003) and can be 

stored for as long as 5 years without loss of 

viability (Fire Sciences Laboratory 2003). 

Scarification to break physical (endocarp) 

dormancy may be done by a 3 to 4 hr soak in 

concentrated sulfuric acid (Rowe and Blazich 

2003) or emersion for less than 1 min. in boiling 

water (Li and others 1999). 

Growth and Management.—Smooth sumac has a 

moderate growth rate of 30 to 46 cm in height per 

year (Michigan State University Extension 1999). 

Although clones may last much longer, individual 

stems are not long-lived, perhaps 2 to 10 years. It 

requires more than ordinary maintenance to keep 

smooth sumac from spreading vegetatively. One 

method of keeping plants in check is to establish 

them in soil space limited by rock or concrete 

barriers (Clark 1973). The tops are killed by fires, 

but underground rhizomes (7.5 to 30 cm deep) are 

not damaged and sprout vigorously, increasing the 

density and extent of smooth sumac stands. Seeds 

from the soil seed bank are also stimulated to 

germinate. Spring fires are more beneficial than 

late summer fires (Fire Sciences Laboratory 2003). 

When it is necessary to control the plant, it is 

recommended that they be lopped during or 

shortly after flowering and the sprouts spotsprayed with glyphosate (Hutchison 2003). 

Benefits and Detriments.—Smooth sumac helps 

protect the soil in disturbed areas, adds to the 

aesthetics of wildlands, and furnishes food and 

cover for wildlife. The fruits are eaten by a wide 

variety of birds (Johnson 1995). Smooth sumac 

foriage is rated poor in both energy and protein 

value (Fire Sciences Laboratory 2003). White tail 

deer (Odocoileus virginianus) browse it during fall 

and winter but do not weaken the plants (Strauss 

1991). It is also browsed by mule deer (Odocoileus 

hemionus) (Fire Sciences Laboratory 2003). 

Smooth sumac is valued as an ornamental for its 

brilliant red and scarlet fall colors and because it 

can be planted almost anywhere in the Continental 

United States. It is particularly useful for 

vegetating sandy banks and for low screens (Clark 

1973) and has been recommended for establishing 

“living snow fences.” The species has been planted 

to revegetate roadsides and cuts, landfills, and strip 

mines (Fire Sciences Laboratory 2003, Plummer 

and others 1968). In addition to using dried leaves 

as an adulterant for tobacco and making drinks 

from the fruits, Native Americans employed 

smooth sumac to treat a large number of ailments, 

particularly mouth and throat sores, burns, to 

control diarrhea, and to promote urination (Fire 

Sciences Laboratory 2003, Giese 1996, Moerman 

1986). Basis for medicinal uses may be found in 

the fact that methanol extracts of ground branches 

were effective (although not as effective as 

commercial antibiotics) against total of 11 species 

of bacteria. The antibacterial compounds were 

identified as methyl gallic acid, gallic acid, and the 

methyl ester of 3,4,5-trihydroxybenzoic acid 

(Saxena and others 1994). Some people develop 

allergic dermatitis after contacting the sap, but the 

species is not as dangerous as many other Rhus

species (Clark 1973). Thickets of smooth sumac 

can prevent the development of tree seedlings 
630 

through shade and root competition and arrest 

succession (Putz and Canham 1992). 

References 

Abrams, L. 1951. Illustrated flora of the Pacific 

States. Vol. 3. Stanford University Press, 

Stanford, CA. 866 p. 

Browse-Shrub and Forb Committee. 1985. 

Handbook on seeds of browse-shrubs and forbs. 

Tech. Pub. R8-TP8. Association of Official Seed 

Analysts and U.S. Department of Agriculture, 

Forest Service, Southern Region, Atlanta, GA. 

246 p. 

Clark, L.J. 1973. Wild flowers of British 

Columbia. Gray’s Publishing Limited, Sidney, 

BC, Canada. 591 p. 

Fire Sciences Laboratory. 2003. Fire effects 

information system: Rhus glabra. U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Missoula, MT. http://www.fs. 

fed.us/database/feis/plants/shrub/rhugla/all.html. 

28 p. 

Giese, P. 1996. Red sumac: multi-tribal use. 

http://www.kstrom.net/isk/food/rhususes.html. 

4 p. 

Hutchison, M. 2003. Vegetation management 

guideline: smooth sumac (Rhus glabra). Illinois 

nature Preserves Commission, Belknap IL. 

http://www.inhs.uiuc.edu/VMG/smsumac.html. 

5 p. 

Institute of Systematic Botany. 2003. Atlas of 

Florida plants: Rhus glabra. http://www. 

plantatlas.usf.edu/main.asp?plantID=3941. 2 p. 

Johnson, F.D. 1995. Wild trees of Idaho. 

University of Idaho Press, Moscow, ID. 212 p. 

 

Kearney, T.H. and R.H. Peebles. 1951. Arizona 

flora. University of California Press, Berkeley 

and Los Angeles, CA. 1,032 P. 

Li, X., J.M. Baskin, and C.C. Baskin. 1999. 

Anatomy of two mechanisms of breaking 

physical dormancy by experimental treatments 

in seeds of two North American Rhus species 

(Anacardiaceae). American Journal of Botany 

86(11): 1,505-1,511. 

Michigan State University Extension. 1999. Rhus 

glabra—smooth sumac. http://www.msue.msu. 

edu/msue/imp/modzz/00001245.html. 1 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Research Reports in Ethnobotany Vol. 

1. University of Michigan Museum of 

Anthropology, Ann Arbor, MI. 534 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Rhus glabra L., smooth sumac. 

http://plants.usda.gov/cgi_bin/plant_profile.cgi?

symbol=RHGL. 5 p. 

Plummer, A.P., D.R. Christensen, and S.B. 

Monsen. 1968. Restoring big-game range in 

Utah. Pub. 68-3. Utah Division of Fish and 

Game, Salt Lake City, UT. 183 p. 

Putz, F.E. and C.D. Canham. 1992. Mechanism of 

arrested succession in shrublands: root and shoot 

competition between shrubs and tree seedlings. 

Forest Ecology and Management 49(3-4): 267-

275. 

Reeves, H.C. and J.D. Lenhart. 1988. Fuel weight 

prediction equations for understory woody 

plants in eastern Texas. Texas Journal of 

Science 40(1): 49-53. 

Rowe, D.B. and F.A. Blazich. 2003. Rhus L., 

sumac. In: F.T. Bonner and R.G. Nisley, eds. 

Woody plant seed manual. U.S. Department of 

Agriculture, Forest Service, Washington, DC. 

http://wpsm.net.Rhus.pdf. 16 p. 

Saxena, G., A.R. McCutcheon, S. Farmer, G.H.N. 

Towers, and R.E.W. Hancock. 1994. 

Antimicrobial constituents of Rhus glabra. 

Journal of Ethnopharmacology 42(2): 95-99. 

Strauss, S.Y. 1991. Direct, indirect, and 

cumulative effects of three native herbivores on 

a shared host plant. Ecology 72(2): 543-558. 

Wyman, D. 1969. Shrubs and vines for American 

gardens. The Macmillan Company, London. 

613 p. 
631

Rhus microphylla Engelm. ex Gray little-leaf sumac 

ANACARDIACEAE 

Synonym: Rhoeidium microphyllum (Engelm.) Greene 

James E. Nellessen

General Description.—Little-leaf sumac is 

sometimes referred to as desert sumac because of 

its occurrence in habitats that are drier than the 

habitats of other sumacs. Another name is 

correosa. This highly branched shrub may reach 

heights of up to 4 to 5 m and is frequently as broad 

as it is tall. Some twigs may be somewhat spinytipped. Leaves are odd-pinnately compound, 

alternate, with five to nine elliptic to oblong 

leaflets, which are sessile, 8 to 18 mm long, 2 to 6 

mm wide, and generally not leathery or shiny. 

Leaflet margins are entire and may be somewhat 

revolute, and the rachis is winged (Carter 1997, 

Correll and Johnston 1970, Epple 1995, Great 

Plains Flora Association 1986, Kearney and others 

1951, Martin and Hutchins 1980-81). Leaves 

persist on the plant until the first hard frost. 

Range.—Little-leaf sumac occurs from western 

Texas, southwestern Oklahoma, through the 

southern half of New Mexico into southeastern 

Arizona, and south into Mexico. It occurs at 

elevations from about 1,060 m to 1,830 m. 

Ecology.—This species is a common member of 

Chihuahuan Desert grassland vegetation although 

its distribution is not uniform. It may occur in 

canyons, along river banks, arroyos, rocky 

hillsides, even alkali flats. It grows on both 

limestone and sandy soils and can be a dominant 

in some sandy-dune soils forming extensive 

coppice conditions. Littleleaf sumac will often 

grow adjacent to arroyos but is considered a 

facultative or semiriparian species (Dick-Peddie 

1993, Whitford 2002). It demonstrated a 46 

percent nitrogen and phosphorus resorption 

efficiency from leaves prior to leaf drop in the 

autumn (Whitford 2002). This was lower than the 

obligate riparian shrubs, desert willow [Chilopsis 

linearis (Cavanilles) Sweet] and cutleaf brickellia 

(Brickellia laciniata A. Gray), but higher than the 

nitrogen fixing mesquite (Prosopis glandulosa

Torrey). Hence, the greater resorption efficiencies 

of these shrubs may be due to their biology and 

ecology as it relates to plant nitrogen demand and 

nitrogen availability within their local habitats 

(Killingbeck and Whitford 2001). Littleleaf sumac 

as well as Torrey yucca (Yucca torreyi Shafer) and 

javelina bush (Condalia warnockii MC Johnston) 

were the most frequently used species for nesting 

birds in a Chihuahuan Desert location in southcentral New Mexico (Kozma and Mathews 1997). 

This observation was made despite the fact that 

little-leaf sumac was lower in density (209 

stems/ha in arroyo sites and six stems/ha in upland 

sites) than other shrub species. Twenty-seven bird 

species were monitored from 1993-1995. Large 

junipers may function as nurse plants, aiding the 

establishment of certain shrub species, including 

little-leaf sumac, in western Texas (McPherson 

and others 1988). 

Reproduction.—Flowers are small, greenish 

white with five petals and five stamens. Flowers 

appear before the leaves in both axillary and 

terminal clusters. Blooming may occur from 

March through May depending on local 

environmental conditions and availability of 

moisture. Fruits are red to orange drupes, from 4.5 

to 6 mm in diameter, with a covering of glandular 

hairs, and maturing in July to August. 

Seed Dormancy.—Seed dormancy appears to be 

caused by a water impermeable endocarp (Li and 

others 1999). After incubation on moist substrate 

for 4 weeks, 29 to 34 percent of all seeds had 

imbibed water, and after 1 year imbibition had 

climbed to 93 to 100 percent among all of the 

seeds. This type of physical dormancy occurs in 

other members of the genus Rhus (Young and 

Young 1992), and some members of the genus 
632 

also have physiological dormancy (Baskin and 

Baskin 2001). There is a high proportion of 

species with physical dormancy in savannas, hot 

deserts, and similar climates with distinct dry and 

wet seasons, the wet season being the season 

favorable for germination. The impermeable 

endocarp in the genus Rhus may have evolved as 

long ago as the Middle Eocene epoch (Baskin and 

Baskin 2001). 

Growth and Management.—Little-leaf sumac is 

not one of the faster growing shrubs. It produces 

numerous stems and resprouts readily. 

Tebuthiuron, an herbicide often used for brush 

control on rangelands, proved effective in littleleaf sumac control (Emmerich and others 1984). 

Tebuthiuron was applied at 0.84 kg/ha to 

rangeland in both pelleted and dissolved forms. 

After 21 months there was a 100 percent kill of 

little-leaf sumac, tarbush (Flourensia cernua DC.), 

and four-wing saltbush [Atriplex canescens

(Pursh) Nutt.], all dominant species on the test site. 

Tebuthiuron moved to a soil depth of 15 cm within 

8 months, and after 21 months 38 percent of the 

application was remaining, while only 0.47 percent 

was detected in runoff water. 

Benefits.—The fruits are readily eaten by many 

birds. Little-leaf sumacs are also important nesting 

sites for birds. The fruits may be steeped in warm 

or cold water to make soothing tea-like drinks that 

can sooth sore throats (Bowers and Wignall 1993, 

Warnock 1974). 

References

Baskin, C.C. and J.M. Baskin. 2001. Seeds: 

Ecology, Biogeography, and Evolution of 

Dormancy, and Germination. Academic Press, 

San Diego. 666 p. 

Bowers, J.E. and B. Wignall. 1993. Shrubs and 

Trees of the Southwest Deserts. Southwest 

Parks and Monuments Association, Tucson, AZ. 

140 p. 

Carter, J.L. 1997. Trees and Shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Correll, D.S. and M.C. Johnston. 1970. Manual of 

the Vascular Plants of Texas. Texas Research 

Foundation, Renner, TX. 1,881 p. 

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation: Past, Present and Future. 

University of New Mexico Press, Albuquerque. 

244 p. 

Emmerich, W.E., J.D. Helmer, K.G. Renard, and 

L.J. Lane. 1984. Fate and effectiveness of 

tebuthiuron applied to a rangeland watershed. J. 

of Environmental Quality 13(3): 382-386 

Epple, A.O. 1995. A Field Guide to the Plants of 

Arizona. Falcon Publishing Inc., Helena, MT. 

347 p. 

Great Plains Flora Association. 1986. Flora of the 

Great Plains. University Press of Kansas. 

1,392 p. 

Kearney, T.H., R. Peebles, and Collaborators. 

1951 with 1960 supplement. Arizona Flora. 

University of California Press, Berkeley, CA. 

1,085 p. 

Killingbeck, K.T. and W.G. Whitford. 2001. 

Nutrient resorption in shrubs growing by design, 

and by default in Chihuahuan Desert arroyos. 

Oecologia, Berlin 128(3): 351-359. 

Kozma, J.M. and N.E. Mathews 1997. Breeding 

bird communities and nest plant selection in 

Chihuahuan desert habitats in south-central New 

Mexico. Wilson Bulletin 109(3): 424-436. 

Li, Xiaojie, J.M. Baskin, and C.C. Baskin. 1999. 

Seed morphology and physical dormancy of 

several North American Rhus species 

(Anacardiaceae). Seed Science Research 9(3): 

247-258. 

Martin, W.C. and C.E. Hutchins. 1980-1981 

(reprinted 2001). A Flora of New Mexico. 2 

volumes. Bishen Singh Mahendra Pal Singh, 

India and Koeltz Scientific Books, Germany. 

2,591 p. 

McPherson, G.R., H.A. Wright, and D.B. Wester. 

1988. Patterns of shrub invasion in semiarid 

Texas USA grasslands. American Midland 

Naturalist 120(2): 391-397. 

Warnock, B.H. 1974. Wildflowers of the 

Guadalupe Mountains and the Sand Dune 

Country, Texas. Sul Ross State University, 

Alpine, TX. 176 p. 
633

Whitford, Walter G. 2002. Ecology of Desert 

Systems. Academic Press, San Diego, CA. 

343 p. 

Young, J.A. and C.G. Young. 1992. Seeds of 

Woody Plants in North America. Dioscorides 

Press, T. R. Dudley, general editor, Portland, 

OR. 407 p. 
634 

Rhus trilobata Nutt. oakleaf sumac 

ANACARDIACEAE 

Synonyms: Rhus aromatica Ait. var. trilobata (Nutt.) Gray 

Toxicodendron trilobatum Kuntze 

Schmaltzia oxyacanthoides Greene 

Schmaltzia pubescens Osterh. 

 

Nancy L. Shaw and Ann M. DeBolt 

 

Illustration credit: E.G. Hurd 

 

General Description.—Also known as skunkbush 

sumac or lemonade sumac, oakleaf sumac is a 

morphologically variable deciduous species with a 

spreading, mound-like, or erect growth form. It 

frequently forms clumps or thickets 0.6 to 2.1 m in 

height (Hitchcock and others 1961, Thornburg 

1982, Welch and others 1987). Young branches 

are brownish and puberulent, turning gray with 

age. The alternate, trifoliate leaves are glossy, dark 

green, and 3 to 7 cm long. Leaflet lobes are 

shallowly to deeply incised and puberulent below. 

The terminal leaflet is fan-shaped; the lower two 

are about half as large and shallowly crenate-lobed 

(Hitchcock and others 1961). Plants are polygamodioecious (nearly dioecious, but with some 

male/female flowers). Inflorescences consist of 

clusters of spikes located near branch tips 

(Goodrich and Neese 1986). Flowers are 

inconspicuous, pale-yellow and mostly imperfect, 

developing before the leaves. Fruits are sticky, 

subglobose, orange-red drupes 6 to 8 mm in 

diameter with a lemony flavor when fresh. This 

species closely resembles R. aromatica Ait. and is 

often treated as a variety of that more eastern 

species. 

Range.—Oakleaf sumac is a widespread and 

highly variable taxon, ranging from Alberta to 

Iowa and south to California, Mexico, and Texas 

(Harrington 1964). It grows at elevations from 

1,050 to 2,400 m on sites that receive 250 to 500 

mm of precipitation (Hitchcock and others 1961, 

USDA Forest Service 1937, Wasser 1982). 

Ecology.—In the Intermountain Region, oakleaf 

sumac is common in mountain brush and pinyonjuniper communities (Hitchcock and others 1961, 

Welch and others 1987) and also grows in 

openings in communities of oakbrush, ponderosa 

pine, and other conifers (Vories 1981). At lower 

elevations it may be found in the blackbrush, basin 

big sagebrush, or Wyoming big sagebrush zones. 

In these areas it occurs on streambanks and 

terraces, floodplains, seep, and spring margins, 

mesic slopes, talus slopes, swales, drainageways, 

and on other sites receiving extra moisture (Welsh 

and others 1987). It is frequently associated with 

riparian communities, growing intermixed with 

other shrubs or as scattered thickets in 

communities dominated by herbaceous species. 

Adapted to a wide range of soil conditions, oakleaf 

sumac grows on basic to acidic soils with textures 

ranging from clayey to sandy. 

Reproduction.—Flowers appear in early spring, 

usually March or April (Brinkman 1974). Fruits 

ripen from June to September (Plummer and 

others 1968, Shaw 1984, van Dersal 1938) but 

many remain on the shrub well into winter. Seeds 

are dispersed primarily by birds but also by 

rodents and other animals (Brinkman 1974). There 

are approximately 44,000 cleaned seeds/kg 

(Brinkman 1974, Plummer and others 1968). 

Germination is restricted by the presence of a hard 

seedcoat and embryo dormancy, with the degree of 

dormancy varying considerably among 

populations. Seedcoat permeability of most 
635

populations is improved by a 0.3 to 2 hour 

sulphuric acid scarification (Babb 1959, Brinkman 

1974, Glazebrook 1941, Heit 1968), while 30 to 

120 days of moist prechilling is required to release 

embryo dormancy (Babb 1959, Swingle 1939). 

Mechanically rupturing the seedcoat improves 

germination (McKeever 1938), but techniques 

have not been developed for treating large seed 

lots. Following fire, oakleaf sumac generally 

sprouts vigorously from the root crown and 

rhizomes (Wasser 1982). 

Growth and Management.—Local seed sources 

should be selected for range seedings or 

transplanting projects as the range of adaptability 

for most populations is not known (Swenson 

1957). Scarified seed may be fall planted to 

provide overwinter moist prechilling. Seed should 

be planted approximately 1.3 cm deep or slightly 

deeper in dry, coarse textured soils. Seedlings are 

considered to be drought and cold hardy but may 

succumb to competition from herbaceous species 

when young (Clark and Depuit 1981, Monsen 

1987). Range or wildland seedings normally 

require 2 to 5 years for establishment (Plummer 

and others 1968). 

Benefits.—Oakleaf sumac can be used to stabilize 

disturbed riparian areas. It provides excellent 

ground cover, and the spreading root system 

reduces soil erosion. It is particularly useful for 

restoration of meadows and other riparian sites 

where extensive erosion and gully cutting has 

lowered water tables. Oakleaf sumac thickets 

provide excellent cover for many small birds and 

mammals, and winter browse where shrubs are 

limited. It is increasingly being used by 

homeowners seeking drought-tolerant native 

species since it is commercially available, easy to 

grow, long-lived, responds well to pruning, and 

has attractive red, orange to yellow fall color. 

Native Americans have used oakleaf sumac for 

food, tobacco substitute, basketry, and as a 

mordant in dyeing (LHBH 1976). 

References 

Babb, M.F. 1959. Propagation of woody plants by 

seed. Bulletin 26. Alaska Agricultural 

Experiment Station. [Place of publication 

unknown]. 12 p. 

Brinkman, K.A. 1974. Rhus L. Sumac. In: C.S. 

Schopmeyer, tech. coord. Seeds of woody plants 

in the United States. Agriculture Handbook 450. 

U.S. Department of Agriculture, Forest Service, 

Washington, DC. p. 715-719. 

Clark, J.W. and J.E. Depuit. 1981. Analysis of 

direct seeding methods for establishment of 

selected native shrub species on minesoils in 

southeastern Montana. In: L.H Stelter, E.J. 

DePuit, and S.A. Mikol, tech. coords. Shrub 

establishment on disturbed arid and semi-arid 

lands. Wyoming Game and Fish Department, 

Cheyenne, WY. p. 89-103. 

Glazebrook, T.B. 1941. Overcoming delayed 

germination in the seed of plants valuable for 

erosion control and wildlife utilization. Master’s 

thesis, University of Idaho, Moscow, ID. 97 p. 

Goodrich, S. and E. Neese. 1986. Uinta Basin 

flora. U.S. Department of Agriculture, Forest 

Service, Intermountain Region, Ogden, UT. 319 

p. 

Harrington, H.D. 1964. Manual of the plants of 

Colorado. The Swallow Press, Inc., Chicago, IL. 

666 p. 

Heit, C.E. 1968. Thirty-five years’ testing of tree 

and shrub seed. Journal of Forestry 66: 632-634. 

Hitchcock, C.L., A. Cronquist, M. Ownbey, and 

J.W. Thomson. 1961. Vascular plants of the 

Pacific Northwest. Part 3: Saxifragaceae to 

Ericaceae. University of Washington Press, 

Seattle, WA. 614 p. 

LHBH [Liberty Hyde Bailey Hortorium]. 1976. 

Hortus third: a concise dictionary of plants 

cultivated in the United States and Canada. 

Third Edition, Macmillan Publishing, New 

York. 1,290 p. 

McKeever, D.G. 1938. The effects of various 

methods of treatment on germination of seeds of 

some plants valuable for game and erosion 

purposes. Master’s thesis, University of Idaho, 

Moscow, ID. 128 p. 

Monsen, S.B. 1987. Shrub selections for pinyonjuniper plantings. In: R.L. Everett (comp.). 

Pinyon-juniper conference. INT-GTR-215. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Research Station, Ogden, UT. p. 

316-334. 
636 

Plummer, A.P., D.R. Christensen, and S.B. 

Monsen. 1968. Restoring big game range in 

Utah. Publication 68-3. Utah Division of Fish 

and Game, Salt Lake City, UT. 183 p. 

Shaw, N. 1984. Producing bareroot seedlings of 

native shrubs. In: P.M. Murphy, comp. The 

challenge of producing native plants for the 

Intermountain area. INT-GTR-168. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Forest and Range Experiment 

Station, Ogden, UT. p. 6-15. 

Swenson, W.S. 1957. Squawbush in windbreaks in 

eastern Colorado. Journal of Soil and Water 

Conservation 12: 184-185. 

Swingle, C.F. 1939. Seed propagation of trees, 

shrubs, and forbs for conservation planting. 

SCS-TP-12. U.S. Department of Agriculture, 

Soil Conservation Service, Washington, DC. 

198 p. 

Thornburg, A.A. 1982. Plant materials for use on 

surface mined land in arid regions. T.P. 157, 

EPA 600-7-79-133. U.S. Department of 

Agriculture, Soil Conservation Service, 

Washington, DC. 88 p. 

USDA Forest Service. 1937. Range plant 

Handbook. U.S. Department of Agriculture, 

Forest Service, Washington, DC. Republished 

by Dover Publications, London. 816 p. 

van Dersal, W.R. 1938. Native woody plants of the 

United States: Their erosion-control and wildlife 

values. Miscellaneous Publication 303. U.S. 

Department of Agriculture, Washington, DC. 

302 p. 

Vories, K.C. 1981. Growing Colorado plants from 

seed: a state of the art. Vol. 1: Shrubs. INTGTR-103. U.S. Department of Agriculture, 

Forest Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. 80 p. 

Wasser, C.H. 1982. Ecology and culture of 

selected species useful in revegetating disturbed 

lands in the West. FWS/OBS-82/56. U.S. 

Department of the Interior, Fish and Wildlife 

Service, Office of Biological Services, Western 

Energy and Land Use Team, Washington, DC. 

347 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoir 9. Brigham Young 

University, Provo, UT. 894 p. 
637

Ribes aureum Pursh golden currant

GROSSULARIACEAE 

Synonyms: Chrysobotrya aurea (Pursh.) Rydb. 

Juanita A.R. Ladyman 

 Illustration credit: J. Shoemaker in Wasser 1982 

General Description.—Ribes aureum Pursh. is 

commonly known as golden currant, but local 

names also include buffalo, flowering, fragrant, 

and clove currant (Elmore 1976). The species 

name “aureum” is derived from the Latin word for 

“golden-yellow” and refers to the rich-yellow 

colored flowers. In some areas flowers may also 

be a cream or reddish color (Derig and Fuller 

2001). The derivation of “Ribes” is apparently 

debatable. Gledhill (1992) suggests it is derived 

from the Persian word for “acid-tasting” whereas 

Vines (1986) indicates “Ribes” may be derived 

from an ancient Danish name. Another likely 

derivation is a link to historical cultural use. In the 

Middle East, a popular syrup was made from an 

Oriental species of rhubarb (Rheum ribes) called 

“Ribas” (USDA FS 1988, ISTA 1968). In Western 

Europe, currants were substituted for the rhubarb 

in this prized juice, and apparently this use 

influenced Linnaeus to adopt “Ribes” as the 

generic name for currants (USDA FS 1988). 

Golden currant is a deciduous, unarmed (spineless) 

shrub. It is usually from 60 cm to 3 m tall but may 

grow to a little over 4 m. The light green, 

semiglossy leaves are usually three-lobed. The 

flowers are large (up to 9 mm long) and each has 

five petals. They are solitary or in erect clusters, or 

racemes, of up to 15 flowers. The smooth, hairless, 

spherical fruit is a juicy several-seeded berry 

approximately 0.5 to 1.5 cm across and variously 

colored from yellow, red, brown to black. This 

description is after Carter 1997, Cronquist and 

others 1997, Hitchcock and Cronquist 1973, and 

Vines 1986. Kartesz (1994) and the U.S. 

Department of Agriculture (2003) database 

PLANTS recognize three varieties: the type 

variety aureum, variety gracillimum (Coville and 

Britt.) Jepson that is synonymous with Ribes 

gracillimum, and variety villosum DC that is 

synonymous with Ribes odoratum H. Wendl. 

Despite some confusing references to the contrary, 

Ribes aureum var. aureum is not a synonym of 

Ribes odoratum. The likely source of the 

taxonomic confusion is that Ribes odoratum H. 

Wendl is synonymous with Ribes aureum Gray, as 

opposed to the more familiar and taxonomically 

accepted, Ribes aureum Pursh. 

Range.—Golden currant, Ribes aureum var. 

aureum, is a wide-ranging species across Western 

North America (Cronquist and others 1997). It is 

common from Alberta, Canada, to western Texas 

(Kershaw and others 1998, Powell 1998). Ribes 

aureum var, odoratum is native to mid-America, 

west of the Great Lakes (Cronquist and others 

1997). Both varieties have become naturalized in 

Europe and possibly in other parts of the world 

(Stace 1997, Cronquist and others 1997, Komarov 

1939). 

Ecology.—Golden currant grows along water 

courses and in moist meadows in a variety of 

vegetation communities from grasslands, shrub 

lands, piñon juniper woodland, western hardwood, 

pine, and fir-spruce forests (Marshall 1995, DickPeddie 1992, Welsh and others 1993). Common 

associates include rabbitbrush (Chrysothamnus

species), maples (Acer species), serviceberry 

(Amelanchier species), chokecherry (Prunus 

virginiana L.), juniper (Juniperus species), 

ponderosa pine (Pinus ponderosa P. & C. 

Lawson), willows (Salix species), and 

cottonwoods (Populus species). The elevation at 

which it occurs tends to depend somewhat upon 

latitude, but generally plants are found between 
638 

approximately 400 to 2,800 m (Cronquist and 

others 1997, Carter 1997, Powell 1998). Golden 

currant grows in full sunlight but also does well in 

partial shade (Tykač 1990). It is tolerant to a range 

of weakly acid (pH 6) to weakly basic soils 

(Wasser 1982). Bacterial diseases have been noted 

to infect canes of some genotypes, but resistant 

varieties are available (Wasser 1982). Ribes

species are intermittent hosts of the white pine 

blister rust (Cronartium ribicola), a fungal 

pathogen that can be ruinous to most native white 

pine species. Because of this, Ribes species have 

been subject to specific eradication programs in 

many white pine (Pinus monticola Dougl. ex D. 

Don) regions (Martin and others 1951, Van 

Arsdel and others 1998). However, the 

significance of specifically golden currant in the 

life cycle and pathology of this rust is uncertain. 

Throughout the Sacramento Mountains of New 

Mexico, R. pinetorum Greene was a major 

contributor to the spread of rust to nearby pines, 

but blister rust was not found on golden currant 

that generally grew below the elevation of the 

pines (Van Arsdel and others 1998). 

Reproduction.—Golden currant generally flowers 

in June through August, although flowers may be 

found from March until the first frost (Epple 1995, 

McGregor and others 1986, USDA FS 1988, Vines 

1986). The fragrant flowers are insect pollinated. 

Some insects have adapted to the long narrow 

flower tube by boring a hole at its base to “steal” 

the nectar (Derig and Fuller 2001). Golden currant 

readily crosses with other Ribes species (Wasser 

1982). The fruits mature in summer and fall. The 

seed is a densely hairy achene. The cleaned seed 

averages approximately 478,398 to 513,672 

individuals per kg with a germination rate of 60 to 

98 percent (Vines 1986, Wasser 1982). The seeds 

undergo a physiological dormancy that is broken 

by 60 days of cold stratification (Baskin and 

Baskin 2001). Germination is epigeal. The plant 

also spreads by growing shoots from rhizomes 

(Wasser 1982). 

Growth and Management.—Golden currant is 

easily propagated from hardwood cuttings in June 

or September and bare rooted plants are best 

planted when leafless in the spring or fall (Tykač

1990). When starting plants from seed, fall 

seeding is preferred, but stratified seed can be 

planted in the spring (Wasser 1982). Golden 

currant is used as a rootstock for grafting on other 

species of currant that root poorly (Tykač 1990). 

Cutting back old shoots stimulates new growth, 

and plants can sprout after fire (Tykač 1990, 

Marshal 1995). Tolerance to fire in the dormant 

state is good (Wasser 1982). However, the 

severity, or heat, of the fire is likely to be 

important to survival, and very hot fires can kill 

shrubs completely (Marshall 1995). 

Benefits.—Golden currant rates from “poor” and 

“fair” to “medium” palatability for cattle (Dayton 

1931, USDA FS 1988). It has good palatability for 

sheep (Dayton 1931). In Montana, twigs and 

foliage can make up to 2 to 5 percent of a sheep’s 

diet (Martin and others 1951). The fruits are an 

important food source for a variety of wildlife 

such as songbirds, chipmunks, mice, and ground 

squirrels (Martin and others 1951, Marshall 1995, 

Tykač 1990). The fruits may contribute up to 10 

percent of a chipmunk and ground squirrels’ diet 

(Martin and others 1951). Both fruit and foliage 

are used by a range of species that include grouse, 

coyote, and beaver (Martin and others 1951). Elk 

have been reported to eat the foliage in Colorado, 

and one report indicated that mule deer browsed it 

infrequently, but cattle not at all, during a 1-year 

study in Montana (Dusek 1975). Golden currant 

has been planted for erosion control and in 

restoration projects (Vines 1986, Wasser 1982). It 

is particularly used for restoration of game habitat 

(Wasser 1982). When golden currant is used in 

vegetation restoration projects it is usually a small 

component of the seed mix and used at 

approximately 461 to 922 grams per hectare 

(Wasser 1982). The berries, both fresh, dried, and 

made into jams and cakes, have been used for food 

by many Native American tribes as well as by 

European settlers (Elmore 1976, Moerman 1998, 

Welsh and others 1993). It has also been used as a 

pharmaceutical by some of the tribes of the Great 

Plains. The dried, pulverized inner bark was 

sprinkled on sores, used with other ingredients in a 

poultice, and a decoction was taken for leg 

swelling (Moerman 1998). The flowers are edible, 

apparently flavorful, and eaten by gourmets 

(Elmore 1976). Golden currant is a popular 

ornamental plant in North America, Europe, and 

parts of the former Soviet Union (Bailey and 

others 1976, Komarov 1939, Tykač 1990).

References 

Bailey, L.H., E.Z. Bailey, and the staff of the 

Liberty Hyde Bailey Hortorium. 1976. Hortus 

Third–a concise dictionary of plants cultivated 

in the United States and Canada. Macmillian 

Publishing Company, New York. 1,290 p. 
639

Baskin, C.C. and J.M. Baskin. 2001. Seeds, 

ecology, biogeography, and evolution of 

dormancy and germination. Academic Press, 

New York. 666 p. 

Carter, J.L. 1997. Trees and shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Cronquist, A., N.H. Holmgren, and P.K. 

Holmgren. 1997. Intermountain flora, vascular 

plants of the Intermountain West, U.S.A. Vol. 

3, Part A. New York Botanical Garden, Bronx, 

NY. 446 p. 

Dayton, W. A. 1931. Important western browse 

plants. Miscellaneous Publication 101. U.S. 

Department of Agriculture, Washington DC. 

214 p 

Derig, B.B. and M.C. Fuller. 2001. Wild berries of 

the West. Mountain Press Publishing Company, 

Missoula, MT. 235 p. 

Dick-Peddie, W.A. 1992. New Mexico 

Vegetation– past, present, and future. 

University of New Mexico Press, Albuquerque, 

NM. 244 p. 

Dusek, G.L. 1975. Range relations of mule deer 

and cattle in prairie habitat. Journal of Range 

Management 39(3): 605-615. 

Elmore. F.H. 1976. Shrubs and Trees of the 

Southwest Uplands. Southwest Parks and 

Monuments Assoc. Tucson, AZ. 214 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. Falcon Press Publishing Co., Helena, 

MT. 347 p. 

Gledhill, D. 1992. The names of plants. 2nd

edition, reprinted 1992. Cambridge University 

Press, Cambridge, UK. 202 p. 

Hitchcock, C.L. and A. Cronquist. 1973. Flora of 

the Pacific Northwest. University of 

Washington Press, Seattle, WA. 730 p. 

International Seed Testing Association (ISTA). 

1968. A multilingual glossary of common 

plant-names 1. Field crops, grasses and 

vegetables. International Seed Testing 

Association, Wageningen, The Netherlands. 

371 p. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada and 

Greenland. Vol.1 - Checklist. 2nd Ed. Timber 

Press, Portland, OR. 622 p. 

Kershaw, L., A. MacKinnon, and J. Pojar. 1998. 

Plants of the Rocky Mountains. Lone Pine 

Publishing, Edmonton, Canada. 383 p. 

Marshall, K.A. 1995. Ribes aureum. In: 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire 

Sciences Laboratory. Fire Effects Information 

System, http://www.fs.fed.us/ database/feis/. 

13 p. 

Martin, A C., Zim, H. S., and Nelson, A.L. 1951. 

American wildlife and plants. 2nd printing. 

McGraw-Hill Book Company, Inc., New York. 

500 p. 

McGregor, R. L., T.M. Barkley, R.E. Brooks, and 

E.K. Schofield. 1986. Flora of the Great Plains. 

University Press of Kansas, Lawrence, KS. 

1,402 p. 

Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. 927 p. 

Powell, A.M. 1998. Trees and shrubs of TransPecos and adjacent areas. University of Texas 

University Press, Austin, TX. 498 p. 

Stace, C. 1997. New Flora of the British Isles. 2nd

Ed. Cambridge University Press, New York. 

1,130 p. 

Komarov. B.K. 1939. Flora SSSR. Vol. 9. Flora of 

the USSR translated 1964 by the Israel Program 

for Scientific Translations, Jerusalem, Israel. p. 

205 

Tykač, J. 1990. The illustrated guide to ornamental 

shrubs. Treasure Press, London, UK. 224 p. 

U.S. Department of Agriculture. 2003. Plants 

database. http://plants.usda.gov/cgi _bin/plant_ 

profile.cgi?symbol=FAPA# [not paged]. 

United States Department of Agriculture Forest 

Service (USDA FS). 1988. Range Plant 

Handbook. Dover Publications, Inc. New York. 

838 p. 
640 

Van Arsdel, E.P., D.A. Conklin, J.B. Popp, and 

B.W. Geils. 1998. The distribution of white 

pine blister rust in the Sacramento Mountains of 

New Mexico. Finnish Forest Research Institute 

Research Papers 712: 275-283. 

Vines, R.A. 1986. Trees, shrubs, and woody vines 

of the Southwest. University of Texas Press. 

Austin, TX. 1,104 p. 

Wasser, C.H. 1982. Ecology and culture of 

selected species useful in revegetating disturbed 

lands in the West. FWS?OBS-82/56. Fish and 

Wildlife Service, U.S. Department of the 

Interior, Washington, DC. 347 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1993. A Utah flora. 2nd Edition, 

Revised. Brigham Young University, Provo, 

UT. 986 p. 
641

Ribes cereum Dougl. wax current 

GROSSULARIACEAE 

Synonyms: Ribes inebrians Lindl. 

 Ribes cereum Dougl. var. inebrians (Lindl.) C.L. Hitchc. 

John K. Francis 

Drawing source: Britton and Brown 1913 

General Description.—Wax current, also known 

as squaw current, is a deciduous shrub 0.5 to 1.5 m 

in height. Plants support one to several stems and 

are unarmed, dark gray to brown, sticky, and hairy 

when young, becoming glabrous with age. The 

yellow-green to grayish-green, simple leaves are 

circular to kidney-shaped with three to seven 

shallow lobes and a fine toothed or roundedtoothed margin. Petioles are 0.4 to 2.2 cm long and 

blades are 0.5 to 2.5 long and 0.7 to 3.0 cm broad. 

Inflorescences are two- to four-flowered racemes 

that are shorter than the leaves. The 4- to 7-mm 

long, glandular, tubular, greenish-white to pale 

pink flowers have five tiny petals. Fruits (berries) 

are red, shiny, and 6 to 8 mm in diameter. Each 

fruit contains several rough-surfaced, elliptical 

seeds about 2.5 mm long. There are 2n = 16 

chromosomes (Abrams 1944, Brayshaw 1996, 

Mesler and Sawyer 2003, Pfister 1974, Welsh and 

others 1987). 

Range.—Wax current is native to the states of 

Washington, Oregon, California, Nevada, Utah, 

Arizona, New Mexico, Oklahoma, Idaho, 

Montana, Wyoming, North Dakota, South Dakota, 

and Nebraska, as well as the provinces of British 

Columbia and Alberta. There are three recognized 

varieties. Variety cerium is apparently found in all 

the range except Oklahoma. Variety colubrinum

C.L. Hitchc. occurs in Washington, Oregon, Idaho, 

(and probably British Columbia). Variety 

pedicellare Brewer & S. Wats., known as whisky 

current, occupies the range east and south of 

Washington and Oregon (Natural Resources 

Conservation Service 2003). 

Ecology.—Wax current is most common and 

grows best in the open but it can grow under open 

conifer forest canopies. It therefore classifies as 

intolerant of shade. New seedlings appear after 

fires (that may scarify seeds) and when the soil is 

disturbed and opened to sunlight (Marshall 1995). 

The species grows on well-drained soils of the full 

range of textures that have developed from a wide 

variety of parent materials. These soils are often 

rocky or gravelly. Wax current grows at elevations 

of 1,500 to 4,000 m (Marshall 1995) in areas with 

warm, dry to moist summers and cool to cold 

winters. Minimum annual precipitation is around 

330 mm/year. Wax current occurs in many 

vegetation types including open and broken 

coniferous forests, exposed subalpine slopes and 

ridges, and sagebrush (Artemisia spp.) steps 

(author’s observation). 

Reproduction.—Wax current flowers between 

April and June and ripens fruit in August (Pfister 

1974). Flowers are pollinated by insects (Plants 

For a Future 2003). There was an average of 

553,000 seeds/kg from fruits collected in 

California (Pfister 1974). Seeds are dispersed by 

birds and small mammals, as well as by gravity 

(Marshall 1995). At 21 °C, air-dried seeds retained 

some viability for 27 years (Pfister 1974). 

Reproduction is mainly by seed; the species has 

only a “weak” ability to sprout from the root 

crown (Marshall 1995). 
642 

Growth and Management.—Fruits must 

generally be picked by hand. Seeds are cleaned of 

fruit by maceration, and air dried. They can be 

stored in sealed containers at ambient temperatures 

or refrigerated (Pfister 1974). Seeds require 4 to 5 

months of cold stratification (at temperatures near 

freezing) after which they can be planted in the 

spring (Marshall 1995). Alternately, they can be 

sown in the fall and allowed to stratify naturally. 

Plants can also be started from cuttings of semihardened wood (Plants For a Future 2003). Adult 

plants can be eliminated from stands by fire, 

grubbing, and probably herbicides. However, in at 

least one study, because many new seedlings 

appeared, the number of plants increased in sites 

disturbed by treatments (Marshall 1995). Because 

wax current does poorly in shade, tree planting is 

suggested as a long-term solution. 

Benefits.—Wax current contributes to the beauty 

of forest lands where it grows and helps protect the 

soil from erosion. Shade from their crowns shelters 

conifer seedlings and thus aids forest succession 

(Marshall 1995). The species is used to a limited 

extent as a foundation plant in formal landscaping 

(Wildland Nursery 2003). Wax current is fair to 

poor browse for domestic livestock and wild 

ungulates and is mainly eaten when little else is 

available. New annual growth contains about 5.6 

percent protein. Several species of birds and 

rodents consume the fruits (Marshall 1995). The 

larva of the Rocky Mountain Agepema moth 

(Agapema homogena Dyar) feed on the foliage 

(Oehlke 2003). Wax current berries are edible but 

have little flavor. They are picked for making 

jams, jellies, and pies and were formerly used by 

Native Americans for making pemmican (Marshall 

1995). People of the Secwepemc tribe of British 

Columbia ate them to relieve diarrhea (Secwepemc 

Nation 2003). An infusion of inner bark was used 

as a wash for sore eyes (Plants For a Future 2003). 

Wax current and other Ribes species are alternate 

hosts for white pine blister rust (Cronartium 

ribicola), which can be lethal to five-needle pines. 

Efforts to eradicate Ribes from several forest areas 

failed and did not decrease the incidence of blister 

rust (Marshall 1995). 

References

Abrams, L. 1944. Illustrated flora of the Pacific 

States. Vol. 2. Stanford University Press, 

Stanford, CA. 635 p. 

Brayshaw, T.C. 1996. Trees and shrubs of British 

Columbia. UBC Press, Vancouver, B.C., 

Canada. 373 p. 

Britton, N.L., and A. Brown. 1913. Illustrated flora 

of the northern states, Canada, and the British 

possessions. Vol. 2, 2nd Ed. Scribner, New York. 

735 p. 

Marshall, K.A. 1995. Ribes cereum. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shurb/ 

ribcer/all.html. 14 p. 

Mesler, M.R. and J.O. Sawyer, Jr. 2003. 

Grossulariaceae, gooseberry family. http:// 

ucjeps.berkeley.edu/chi-bin/get_JM_treatment. 

pl?4451,4452,4463. 3 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Ribes cereum Doul. 

http://plants.usda.gov/cgi_bin/plant_profile.chi?

symbol=RICE. 3 p. 

Oehlke, B. 2003. Agapema homogena moth, Dyar, 

1908. http://www3.islandtelecom.com/ 

~oehlkew/sahomoge.htm. 4 p. 

Pfister, R.D. 1974. Ribes L., current, gooseberry. 

In: C.S. Schopmeyer, tech. coord. Seeds of 

woody plants in the United States. Agriculture 

Handbook 450. U.S. Department of Agriculture, 

Forest Service, Washington, DC. p. 720-727. 

Plants For a Future. 2003. Database search results: 

Ribes cereum. http://www.ibiblio.org/pfaf/cgibin/arr_html?Ribes+cereum&CAN=LATIND. 

5 p. 

Secwepemc Nation. 2003. Secwepemc 

ethnobotanical garden: other plants. Sccwepemc 

Nation, Kamloops, BC, Canada. 4 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1987. A flora of Utah. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University Press, Provo, UT. 894 p. 

Wildland Nursery. 2003. Wildland nursery: 

shrubs. Wildland Nursery, Joseph, UT. 

http://www.wildlandnursery.com/shrubs/shrubs

M-Z.html. 6 p. 
643

Ricinus communis L. castor bean 

EUPHORBIACEAE 

Synonyms: none 

John K. Francis 

General Description.The castor bean, also 

known as the castor-oil plant, higüerito, higüerillo, 

palma Christi, carrapateira, ricin commún, and 

many other common names, is grown as an annual 

herb in temperate countries but grows to treelike 

proportions on fertile ground in tropical climates. 

The plant usually has only one stem arising from 

the soil but develops a number of low robust 

branches when open grown. Castor bean is 

herbaceous when young but becomes woody with 

age. The wood is soft and light with a thick central 

pith. Occasionally, irregular brown heartwood 

develops (Kadambi and Dabral 1955). The bark is 

light brown, smooth, and exhibits rings at the 

nodes and raised lenticels. There are a moderate 

number of large, star-shaped leaves with 7 to 9 

long pointed lobes (Little and others 1974). Castor 

bean has been cultivated in Africa and Asia from 

ancient times and used medicinally (CISR 1972). 

Range.—Castor bean is a native of tropical Africa 

but has naturalized in moist tropical and 

subtropical regions throughout the world. It may be 

found growing naturally and in cultivation as an 

ornamental throughout the tropical and subtropical 

areas of the United States and its territories. 

Ecology.—Disturbance is required for successful 

natural stands of castor bean. The species is an 

intolerant pioneer. If disturbance is not repeated, it 

will be succeeded in a few years by grass, vines, or 

trees. Castor bean is competitive and most 

frequently seen in flood zones, on neglected 

farmland, and roadsides. In Puerto Rico, wild 

stands rarely succeed on exposed subsoils and 

highly weathered soils such as Ultisols. In 

cultivation, where excessive vegetative growth is a 

detriment, castor beans are often grown on 

marginal agricultural soils (CSIR 1972). Castor 

bean is not frost hardy. Many pests and diseases 

are known worldwide, but they rarely present a 

serious threat to cultivated or wild stands. 

Reproduction.—Castor bean flower clusters grow 

erect at the ends of twigs. Male flowers form on the 

lower part and female flowers on the upper part of 

each raceme. The seed capsules are covered with 

weak spines (spineless types exist). The capsules 

split into three parts, each containing one seed. The 

seeds are shiny brown with darker streaks or spots 

and resemble a bloated tick (Little and others 

1974). Two seed sizes are reported in India: 1,164 

seeds/kg and 2,610 seeds/kg (Kadambi and Dabral 

1955). Stored seeds should be kept cool. Seeds 

stored at ambient temperature lose 75 percent of 

their viability within 3 months (CISR 1972). 

Soaking the seeds for 12 hours prior to sowing is 

reported to improve germination (CISR 1972). 

Germination starts in 7 to 10 days and is complete 

in 25 to 45 days (Kadambi and Dabral 1955). 

There are hundreds of natural forms and cultivated 

varieties of the species, including annual and 

perennial types. 

Growth and Management.—Castor beans are 

planted in rows spaced from 1 to 2 m apart with 

spacing within the rows of about 0.5 m. When 

grown as an annual crop, it takes 5 to 9 months 

from planting to harvest (CSIR 1972). Castor beans 

are reported to survive for 8 to 10 years in India 
644 

and may reach 6 m in height (Kadambi and Dabral 

1955). Annual varieties reach 1 to 2 m in height. 

Seed yields under cultivation vary from 200 to 

1,700 kg/ha, depending on variety and site quality 

(CISR 1972). 

Benefits.—Castor beans contain from 31 to 61 

percent oil (CISR 1972). After decorticating, they 

are subject to a series of hot or cold presses 

followed by solvent extraction, each step yielding a 

different grade of oil (Kirschenbauer 1960). The 

largest producers are Brazil and India. The 

principal consumer is the United States 

(Encyclopedia Brittanica 2000), but an annual 

demand for 100,000 tons of castor oil is reported 

for Europe (NF-2000 Database 2000). The seed 

cake remaining after extracting the oil is used as 

fertilizer or cooked to destroy the toxin and 

incorporated into animal feeds. Not as popular as it 

once was, castor oil is still widely used in 

traditional and herbal medicine, especially in less 

developed countries. Its principal use in medicine 

is as a purgative and laxative. Castor oil is also 

used as a lubricant, lamp fuel, a component of 

cosmetics, and in the manufacture of soaps, 

printer’s ink, plastics, fibers, hydraulic fluid, break 

fluid, varnishes, paints, embalming fluid, textile 

dyes, leather finishes, adhesives, waxes, and 

fungicides (Encyclopedia Brittanica 2000, CISR 

1972). It is gradually being replaced as a raw 

material for some of the uses by petroleum-based 

products. In India, the leaves are used as food for 

eri silk worms (Kadambi and Dabral 1955). The 

stalks from fields are burned for fuel in India and 

have been shown to be suitable for short-fiber pulp 

(Kadambi and Dabral 1955). The species has been 

planted for dune stabilization (CISR 1972). Castor 

bean is widely planted as an ornamental. Its large, 

star-shaped leaves make it a bold foliage plant. 

Some varieties have red- or purplish-colored leaves 

and stems. 

Detrimental Effects.—Castor bean may become a 

weed in neglected cropland and pasture. It is not 

difficult to control through cultivation and mowing. 

Of greater concern than its weedy potential is the 

high toxicity of its seeds, which contain ricin, a 

water-soluble protein. Even a small amount of 

masticated seed is likely to cause death. Humans 

and horses are especially vulnerable. Fatal doses 

are from 2.5 to 6 seeds for humans and about 6 

seeds for horses (CISR 1972). Symptoms are 

stomach irritation, diarrhea, abdominal pain, 

increased heart rate, profuse sweating, collapse, 

and convulsions. Broken seeds can cause skin 

irritation. The foliage is only slightly toxic 

(Anonymous 2000). It is advisable to completely 

eliminate castor bean from pastures, especially 

horse pastures, and pinch off flowers of ornamental 

plants to prevent possible poisoning of children. 

References 

Anonymous. 2000. Castorbean, castor oil plant. 

http://www.vet.perdue.edu/depts/addl/toxic/ 

plant11. htm. 1 p. 

 

CSIR. 1972. The wealth of India. Raw materials. 

Vol. 9. Publications & Information Directorate, 

Council for Scientific and Industrial Research, 

New Delhi. 472 p. 

Encyclopedia Britanica. 2000. Castor oil. 

http://www.Britanica.com/bcom/eb/article/4/0,57

16,2105+1+20724,00htm?query=castoroil%20 

oil. [not paged]. 

Kadambi, K. and S.N. Dabral. 1955. The 

silviculture of Ricinus communis Linn. Indian 

Forester 81(1): 53-58. 

Kirschenbauer, H.G. 1960. Fats and oils, an 

outline of their chemistry and technology. 

Reinhold Publishing Corporation, New York. 

240 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook. 

449. U.S. Department of Agriculture, Forest 

Service, Washington, DC. 1,024 p. 

NF-2000 Online Database Information Items. 

2000. Crops: castor oil (Ricinus communis). 

http://www.nf-2000.org/secure/Crops/S600.htm 

[not paged]. 
645

Rondeletia inermis (Spreng.) Krug & Urban cordobancillo 

RUBIACEAE 

Synonyms: Catesbaea inermis Spreng. 

 Rondeletia inermis angustifolia Krug & Urban 

 Rondeletia inermis oblongifolia Krug & Urban 

 Rondeletia inermis intermedia Krug & Urban 

 Rondeletia inermis latifolia Krug & Urban 

John K. Francis 

General Description.—Cordobancillo is an 

upright evergreen shrub or occasionally a small 

tree sometimes reaching 4.5 m in height and 7.5 

cm in stem diameter. The plant occasionally takes 

a vine-like form. Older plants usually have 

multiple stems arising at or near ground level. 

Stems and branches are slender, mostly straight. 

The bark is gray or dark brown and nearly smooth. 

Inner bark is green. The wood is hard or 

moderately hard and stiff; sapwood is yellowish 

and heartwood is light brown. Cordobancillo 

plants are supported by a tap and lateral root 

system of stiff, brown roots. Foliage is mostly 

grouped at the ends of twigs. The opposite leaves 

are variable: with or without hairs, cartaceous to 

coriaceous, linear to elliptic, and 1 to 10 cm long. 

There are deltoid stipules about 2 mm long and 

petioles 3 to 6 mm long. Groups of one to five 

small, yellow (or white to purple or brown) 

flowers with four corolla lobes are borne on 

peduncles 0.5 to 3.0 cm long. The fruit is a twocelled, 3- to 4-mm, globose capsule containing 

tiny, wrinkled, brown seeds (author’s observations, 

Liogier 1997, Little and others 1974). 

Range.—Cordobancillo is endemic to Puerto Rico 

and its offshore island, Muertos (Little and others 

1974). There are several races differing in size and 

shape of leaves and amount of pubescence (Liogier 

1997). The species is not known to have been 

planted or naturalized elsewhere. 

Ecology.—The species is intermediate in tolerance 

to shade. It usually grows in the understory of 

open to moderately dense forests, or in small 

openings or edges. Cordobancillo may be found in 

both primary remnants and secondary forests. It 

invades disturbed areas but takes many years to do 

so. Cordobancillo grows in a wide range of welldrained, neutral to mildly acidic soils derived from 

sedimentary (including limestone), igneous, and 

metamorphic (including ultramafic) rocks. It 

grows from a few meters above sea level to 

elevations of 1,000 m (Little and others 1974) in 

areas that receive from about 750 to 2500 mm of 

mean annual precipitation. 

Reproduction.—Cordobancillo flowers and fruits 

throughout the year. A collection of air-dried seeds 

from the moist limestone hills near Bayamon, 

Puerto Rico averaged 1,863,000 seeds/kg. Seed 

dispersal is probably by gravity and wind. 

Collecting seeds is difficult because the capsules 

ripen, dry, and open a few at a time and 

immediately release their tiny seeds. 

Growth and Management.—Cordobancillo 

appears to have a slow to moderate growth rate 

and be relatively long-lived. By sprouting from the 

base and root crown, plants live longer than 

individual stems. There is no published experience 
646 

on plantation establishment or management of 

natural stands. 

Benefits.—Cordobancillo helps protect the soil, 

furnishes cover for wildlife, and adds to the 

aesthetics of Puerto Rican forests. Too small for 

lumber or poles, the wood is occasionally used for 

fuel. 

References

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 
647

Rondeletia pilosa Sw. cordobancillo peludo 

RUBIACEAE 

Synonyms: Rondeletia triflora Vahl 

 Oldenlandia longiflora Lam. 

 Hedyotis longiflora (Lam.) Spreng. 

John K. Francis 

General Description.—Cordobancillo peludo, 

also known as cachimbo peludo and lechecillo, is 

an evergreen shrub or small tree commonly 1.5 to 

2 m in height and 2 cm in basal diameter but 

sometimes reaching 5 m in height. It may have 

several stems from the base. The form is usually 

upright but occasionally vine-like. Stem bark is 

gray and smooth. The plants examined by the 

author had a tan colored, robust, shallow lateral 

root system with sparse secondary and tertiary 

roots. The twigs are long, slender, forking, and 

ringed at the nodes. Soft hairs cover the twigs and 

lower surfaces of the leaves. The leaves are 

opposite, with entire, elliptic blades 4 to 10 cm 

long and 2 to 3.5 cm broad, and pointed at both 

ends, and have a petiole 3 to 6 mm long. 

Inflorescences are axillary and usually have three 

flowers with peduncles up to 7 cm long. The 

flowers have four-lobbed hairy calyxes and 

corollas. The corollas are white with red-tipped 

lobes. The two-celled capsules are rounded, 3 mm 

in diameter, with calyx lobes persistent. The 

capsules produce many minute brown seeds 

(Liogier 1997, Little and others 1974). 

Range.—Cordobancillo peludo is native to Puerto 

Rico, the Virgin Islands, and Monsarrat (Liogier 

1997). It is not known to have been planted or 

naturalized elsewhere. 

Ecology.—Cordobancillo peludo occurs in areas 

that receive from about 750 to about 1000 mm of 

mean annual precipitation from near sea level to 

about 450 m in elevation. Soil texture does not 

appear to be a limiting factor. All are well- to 

excessively-well drained. Soil pH’s range from 

about 5.5 to 8.0. Parent materials may be 

sedimentary (including limestone), igneous, and 

metamorphic (including ultramaphic) rocks. It is 

moderately intolerant and grows in open areas as 

well as under low basal-area low forest. Because 

the species is not highly competitive, 

cordobancillo peludo is more likely to survive 

where growth otherwise is limited—such as very 

rocky terrain. It is scattered to rare in coastal 

thickets, and remnant and middle to late secondary 

forests in upland areas. 

Reproduction.—Cordobancillo peludo appears to 

flower and fruit throughout the year. In terms of 

numbers, production of seeds is huge. The seeds 

are dispersed by the wind. However, seedlings are 

uncommon. The seeds number a few million/kg. 

Of a group of seeds sown on moist filter paper, 69 

percent germinated between 15 and 70 days after 

sowing. Germination is epigeal. 

Growth and Management.—Growth of 

cordobancillo peludo is slow. Basal sprouts in one 

stand grew about 30 cm/year. One plant with a 

2.5-cm basal diameter had 10 growth rings. 

Individual stems probably last 10 to 20 years, and 

plants can prolong their lives by coppicing. No 

planting or management experience is known to 

the author. Probably the best management strategy 

is to protect remnant stands and late-secondary 

forests containing the species. 
648 

Benefits.—Cordobancillo peludo contributes to 

the aesthetics of the forest, it helps protect the soil, 

and furnishes cover for wildlife. The wood is 

useful for small-diameter fuel. 

References 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p.

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 
649

Rosa woodsii Lindl. Wood’s rose 

ROSACEAE 

Synonyms: Rosa fendleri Crep 

Rosa californica Watson. 

Rosa macounii Greene 

Rosa chrysocarpa Rydb. 

Bruce L. Welch 

General Description.—Wood’s rose is a prostrate 

to upright shrub from 0.5 to 3.0 m tall (Blauer and 

others 1975). It is a strongly armed to nearly 

unarmed shrub with well-developed, straight, or 

less often, somewhat curved infrastipular prickles, 

and often with other stout or weak prickles 

(Cronquist and others 1997). The opposite 

compound leaves are composed of five to 11 

leaflets, which are oval, elliptic, or obovate in 

shape, with serrate margins (Blauer and others 

1975). Leaflets are 2 to 5 cm long and 1 to 2.5 cm 

wide; the teeth are not gland-tipped except in var. 

woodsii. Flowers are light-pink to deep-rose color 

and are borne on lateral shoots from old wood, 

which are usually over 10 cm long (Blauer and 

others 1975). Flowers are solitary or two to several 

in small corymbiform cymes that terminate the 

lateral branches of the season and are relatively 

small (Cronquist and others 1997). The 

hypanthium is glabrous and 3 to 5 mm thick at 

anthesis (Cronquist and others 1997). Pedicels are 

glabrous (Cronquist and others 1997). The five 

persistent sepals are from 1 to 2 cm long and 2 to 

3.5 mm wide at the base. The tip of each sepal is 

usually a little expanded above the median 

constriction (Cronquist and others 1997). The 

sepals are erect or spreading in the fruit (Blauer 

and others 1975). Petals, five in all, are 1 to 2.5 cm 

long (Cronquist and others 1997, Welsh and others 

1987). Hips (fruits) are red to purple or nearly 

black in color and ellipsoid to subglobose in shape 

(Cronquist and others 1997, Welsh and others 

1987). Hip size varies from 6 to 15 mm wide and 

up to 2.0 cm in length and contains from 15 to 30 

achenes (Blauer and others 1975). Each achene is 

3 to 4 mm long with stiff hairs along one side. 

Achenes weigh from 0.007 to 0.013 g (Gill and 

Pogge 1974). The diploid chromosome number of 

Wood’s rose is 14 (Blauer and others 1975).

Taxonomy.—Two varieties of Wood’s rose are 

recognized: var. ultramontana and var. woodsii 

(Blauer and others 1975, Cronquist and other 

1997, Welsh and others 1987). Variety 

ultramontana is the most prevalent in the 

Intermountain area of Western United States. 

Woodsii is found on the prairies and plains of 

central North America and is smaller and stiffer 

than var. utlramontana with smaller leaflets that 

often have gland-tipped teeth (Blauer and others 

1975, Cronquist and others 1997, Welsh and 

others 1987). 

Range.—Wood’s rose occurs from Minnesota 

west and northwest to Alaska and British 

Columbia, south to Arizona, northern Mexico and 

western Texas and north to western Kansas and 

North Dakota. It is the most widespread native 

rose in Alberta (Blauer and others 1975, Cronquist 

and others 1997, Pacific Southwest Experiment 

Station 2002). It grows in 24 ecosystems and 

occurs in 51 of Kuchler’s plant associations 

(Pacific Southwest Experiment Station. 2002). 

The Society of American Foresters list Wood’s 

rose as occurring in 55 of its cover types (Pacific 

Southwest Experiment Station 2002). 

Ecology.—Wood’s rose can grow at elevations 

reaching 3,549 m (Pacific Southwest Experiment 

Station 2002). It occurs on bluffs, dry grassy 
650 

slopes, and on sandhills throughout the prairies 

and on riverbanks and clearings in boreal and 

subalpine forests. It can be found on the eastern 

slopes of the Rocky Mountains and throughout the 

Great Basin. It is found along roadsides and 

south-facing cutbanks (Pacific Southwest 

Experiment Station. 2002). In the Great Basin, 

Wood’s rose prefers moist sites along streams, in 

seepage areas along fences, irrigation canals, 

marsh lands, lake shores, and hillsides in 

palustrine and lacustrine habitats (Welsh and 

others 1987). When conditions are favorable, 

nearly impenetrable thickets of Wood’s rose are 

formed along some mountain streams (Pacific 

Southwest Experiment Station 2002). It flourishes 

in moderate shade to full sunlight and so makes 

good growth on all aspects (Blauer and other 

1975). Wood’s rose is adapted to a wide range of 

soil types and textures (Pacific Southwest 

Experiment Station 2002). Growth is generally 

best on moderately fertile, well-drained clay loam, 

sandy loam, or sandy soils. It is tolerant of 

moderately acid to weakly basic soils. (Pacific 

Southwest Experiment Station 2002). It is seldom 

found where the average annual precipitation is 

less than 260 mm (Blauer and others 1975). 

Reproduction.—Seeds are produced on 2 to 5 

year old plants. Seeds of Wood’s rose are ingested 

with the hip and dispersed in the dropping of birds 

and mammals. The seeds have a seed coat 

dormancy and require warm or cold stratification. 

Seed viability ranged from 44 to 64 percent for 

seed collected in northern Alberta. The seeds 

remain viable for 2 to 5 years. Seeds will 

germinate within 30 to 40 days (Blauer and others 

1975, Gill and Pogge 1974, Pacific Southwest 

Experiment Station 2002). Wood’s rose spreads 

vegetatively through underground rhizomes, 

sprouting from the root crown, and by layering 

(Blauer and other 1975, Pacific Southwest 

Experiment Station 2002). 

Management.—Wood’s rose has a fairly high 

tolerance to browsing (Pacific Southwest 

Experiment Station 2002). It is susceptible to 

various leaf spots, as well as to leaf rusts, gray 

mold, powdery mildew, common gall, and stem 

cankers (Pacific Southwest Experiment Station 

2002). The principal forest insect pests are tent 

caterpillar, rose leaf hopper, and a subspecies of 

western tussock moth (Pacific Southwest 

Experiment Station 2002). 

Benefits.—Wood’s rose adds to the biodiversity 

of a multitude of ecosystems. It provides habitat 

and food for a number of wildlife species, and 

watershed protection. Fruits, leaves, or twigs are 

eaten by a number of birds and mammals. These 

include pronghorn, elk, mule deer, white-tailed 

deer, porcupines, beavers, squirrels, coyotes, bear, 

small nongame birds and upland game birds 

(Pacific Southwest Experiment Station 2002). 

Livestock also eat Wood’s rose. Crude protein 

content of Wood’s rose leaves vary from 5.7 (fall) 

to 16.4 (spring) percent of dry matter and stems 

from 5.4 (winter) to 12.0 (spring) percent (Dietz 

1972). Phosphorus content of leaves varied from 

0.29 (fall) to 0.48 (spring) percent of dry matter 

with stems varying from 0.12 (winter) to 0.32 

(spring) percent (Dietz 1972). In vitro digestible 

for Wood’s rose was 65.9 percent of dry matter 

digested for spring tissues and 54.5 percent for 

winter tissues (Dietz 1972). These in vitro 

digestible values would rank Wood’s rose as a 

highly digestible spring and winter forage (Welch 

1989, Welch and Andrus 1977). The dense 

thickets formed by Wood’s rose are used for 

nesting and escape cover by many birds and small 

mammals (Pacific Southwest Experiment Station 

2002). The extensive rhizomes and good 

survivability and revegetation characteristics 

makes this species an effective tool in erosion 

control (Pacific Southwest Experiment Station 

2002). Good results have been obtained with 

rooting of hardwood cuttings, direct seeding, and 

transplanting trials (Pacific Southwest Experiment 

Station 2002). Native Americans made extensive 

use of Wood’s rose roots, stems, leaves, flowers, 

and hips for foods and therapeutic purposes. The 

hips are a source of vitamin C and are dried for 

use in flavoring teas, jellies, fruitcakes, and 

puddings. Wood’s rose is used as an ornamental 

near homes to attract birds and other wildlife 

(Pacific Southwest Experiment Station 2002). 

References 

Blauer, A.C., A.P. Plummer, E.D. McArthur, R. 

Stevens, and B.C. Giunta. 1975. Characteristics 

and hybridization of important intermountain 

shrubs. I. Rose family. Research Paper INT169. U.S. Department of Agriculture, Forest 

Service, Intermountain Forest and Range 

Experiment Station, Ogden, UT. 36 p. 

Cronquist, A., N.H. Holmgren, and P.K. 

Holmgren. 1997. Intermountain flora: vascular 

plants of the Intermountain west, U.S.A. Vol. 3. 

Part A; Subclass Rosidae (except Fabales). The 

New York Botanical Garden., New York. 446 p. 
651

Dietz, D. R. 1972. Nutritive value of shrubs. In: 

C. M. McKell, J. P. Blaisdell, and J. R. Goodin, 

tech. eds. Wildland shrubs–their biology and 

utilization, An international symposium; 

Proceedings; 1971 July; Logan, UT. General 

Technical Report INT-1. U.S. Department of 

Agriculture, Forest Service, Intermountain 

Forest and Range Experiment Station, Ogden, 

UT. p. 289-302. 

Gill, J.D. and F.L. Pogge. 1974. Rosa L. Rose In: 

C.S. Schopmeyer, tech. coord. Seeds of woody 

plants in the United States. Agric. Handbook 

654. U.S. Department of Agriculture, Forest 

Service, Washington, DC. p. 732-737. 

Pacific Southwest Experiment Station. 2002. Fire 

effects information system. 

www.fs.fed.us/database/feis.plant. 33p. 

Welch, B.L. 1989. Nutritive value of shrubs. In: C. 

M. McKell, ed. The biology and utilization of 

shrubs. Academic Press, Inc., San Diego, CA. 

p. 405-424. 

Welch, B.L. and D. Andrus. 1977. Rose hips-a 

possible high-energy food for wintering mule 

deer? Research Note INT-221. Intermountain 

Forest Experiment Station, U.S. Department of 

Agriculture, Forest Service, Ogden, UT. 6 p. 

Welsh, S.L, N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1987. A Utah flora. Great Basin 

Naturalist Memoirs. No. 9. Brigham Young 

University, Provo, UT. 894 p. 
652 

Rubus discolor Weihe & Nees Himalayan blackberry 

ROSACEAE 

Synonyms: Rubus procerus auct. non P. Muell. ex Genev 

 Rubus fruticosus L. 

 Rubus macrostemon (Focke) Sampaio 

 Rubus thyrsanthus (Focke) Foerster. 

 Rubus praecox Bertol. 

 Rubus grabowskii Weihe ex Gunther et al. 

John K. Francis 

General Description.—Himalayan blackberry, 

also known as Armenian blackberry or just 

blackberry, is a deciduous to evergreen, spiny, 

arching woody shrub that may reach heights of 4 

m with stem (cane) lengths up to 10 m. The 

multiple stems are fluted with stout, red, straight or 

recurved prickles along ridges. The canes grow 

straight upward at first but arch as they become 

long. The root crown may reach 20 cm in 

diameter; the root may penetrate up to 1.5 m into 

the soil, and lateral roots extend 30 to 60 cm. The 

bark is green to purplish-red. The 7- to 20-cm-long 

leaves are usually penta-foliate except on fruiting 

branches where they can be trifoliate or unifoliate. 

The petioles, rachises, and lower surfaces of the 

midveins are also armed. The blades are 3 to 12 

cm long, ovate to orbicular, dark green, and have 

biserrate margins. The white to pink, five-peteled 

flowers 2- to 2.5-cm broad are borne three to 20 in 

terminal panicles. Fruits are aggregates 12 to 20 

mm in diameter of black, juicy, sweet druplets on 

a torus (central core). Each drupe contains one 

hard, flattened seed that is covered with a fine 

network of pits and ridges (Brayshaw 1996, 

Bruzzese 1998, Starr and others 2003, Welsh and 

others 1987). Luther Burbank introduced the fruit 

cultivar he called Himalayan blackberry in 1885. 

After its release to gardeners, it quickly spread to 

wildlands on the West Coast and later to other 

parts of the United States (Jacobson 2001). 

Taxonomists disagree on its identity. Some 

maintain that it is in fact Rubus armeniacus Focke 

(Ceska 1999). Himalayan blackberry hybridizes 

with R. thyrsiger Banning & Focke, R. calvatus

Lees ex Bloxam, and R. schlechtendalii Weihe ex 

Link (Tirmenstein 1989). 

Range.—Himalayan blackberry originated in the 

Old World, probably Europe. It has been under 

semicultivation for a long time and has escaped 

into the wild in Western and East-Central United 

States, Hawaii, parts of Europe, Australia, New 

Zealand, South Africa, and probably southern 

South America (Ceska 1999, Natural Resources 

Conservation Service 2003, Starr and others 2003). 

In many populations, it is not always clear from 

the literature whether it is Himalayan blackberry or 

other closely related taxa. 

Ecology.—Himalayan blackberry grows in vacant 

lots, logging sites, burned areas, along rivers, 

roads, fences, and railroad tracks. It grows best in 

full sunlight but does well under a light forest 

cover. However, it will not grow under a dense 

canopy. Himalayan blackberry colonizes most soil 

types in moist areas but is confined to riparian 

areas and the edges of irrigated fields in interior 

dry climates. It grows at elevations from near sea 

level on the West Coast to 1,500 m in the inland 

West (Welsh and others 1987). Himalayan 

blackberry is confined at the northern extent of its 

range to mild coastal areas and low-elevation, 

sheltered inland sites. It tolerates periodic flooding 

of fresh or even brackish water. The species 

vigorously sprouts from root systems after fires 

(Tirmenstein 1989). 
653

Reproduction.—Himalayan blackberry 

commonly flowers from June to August. The 

flowers are intensely visited by honey bees, but 

insect pollination is not necessary for seed 

production. In fact, sexual reproduction may be 

rare; 17 samples from Himalayan blackberry 

plants collected throughout Australia proved to 

have no genetic variation (Evans and others 1998). 

Good seed crops are produced nearly every year 

(Tirmenstein 1989). There are 324,000 cleaned 

seeks/kg. The seeds are dispersed by birds and 

mammals as well as by gravity. Passing through 

birds and mammals is reported to increase 

germination by 30 percent in the first year 

(Bruzzese 1998). Seeds in the soil seed bank 

remain viable for several years (Brinkman 1974). 

Stands in Victoria, Australia have been estimated 

to produce 7,000 to 13,000 seeds/m2

/year 

(Bruzzese 1998). Himalayan blackberry seedlings 

commonly appear after fires or disturbance that 

exposes the soil and allows sunlight to reach the 

surface. Once seedlings become established, most 

subsequent reproduction is vegetative. Plants 

reproduce by sprouts from rhizomes and by 

layering (rooting) at the nodes when stems come in 

contact with the ground. 

Growth and Management.—Canes do not bloom 

during their first year in which they make most of 

their growth (2 to 8 m of elongation). Canes bloom 

and fruit in their second and sometimes third years 

and die at the end of their second or third years. 

Individual root crowns live a maximum of 7.5 

years in Australia (Bruzzese 1998). By suckering 

from rhizomes and layering, plants (clones) can 

endure almost indefinitely. Live biomass of a stand 

in Victoria, Australia totaled 3.1 tons/ha (59 

percent above-ground and 41 percent belowground) and 27 tons of dead canes and litter 

(Bruzzese 1998). Scarification is required for 

prompt germination. Although several methods 

will work, soaking in concentrated H2SO4 for 50 to 

60 minutes followed by 90 days of cold 

stratification is recommended. About 33 percent 

germination in about 70 days can be expected. 

Sowing in late summer for spring germination can 

substitute for artificial cold stratification. Sowing 

depth is 3 to 9 mm. (Brinkman 1974). The species 

can be propagated by digging up and replanting 

suckers. However, many more plants can be 

started by using root cuttings. Root pieces between 

3 and 13 mm in thickness and 10 to 18 cm long are 

recommended. They are grown for 1 year in the 

nursery bed before outplanting (Shoemaker 1978). 

Himalayan blackberry can be troublesome in many 

habitats and is a potential threat to native 

ecosystems in places such as Hawaii (Starr and 

others 2003). Plants can be eliminated by grubbing 

with follow-up treatment a year later, repeated 

mowing, and by pasturing with goats (Cox 2003). 

Several herbicides are effective using foliar spray, 

stem injection, treatment of cut stumps, and basal 

spray methods (Starr and others 2003, Tirmenstein 

1989). Dead canes accumulate in older stands and 

are as much a nuisance to remove as the plants 

themselves. Because of dormant seeds in the soil, 

checking and retreatment for many years may be 

required to completely eliminate the species. 

Benefits.—Himalayan blackberry provides food 

and cover for many mammals and birds. It is little 

used by domestic livestock except for goats. 

However, deer, elk, rabbits, porcupines, beavers, 

and mountain beavers consume leaves, buds, 

twigs, and cambium, especially during the winter 

months. A large number of species consume the 

fruits and a number rely on the thickets of stems 

for escape and reproductive cover (Tirmenstein 

1989). Himalayan blackberry fruits are among the 

most delicious of wild fruits but are difficult to 

pick because of the spines. They are eaten fresh, 

canned, used to flavor ice cream, and made into 

pies, jams, jellies, juices, and wines. Some people 

object to the seeds getting caught in one’s teeth 

when eaten whole, but it is a minor inconvenience 

that can be avoided by juicing the fruit. In spring, 

the succulent canes can be pealed and eaten, fresh 

or cooked (Jacobson 2001). The brambles are 

allowed to grow over fences and trellises to create 

barriers almost impenetrable to people and 

livestock. While the species does protect the soil 

from erosion and helps revegetate disturbed sites, 

the thickets and mounds of brambles restrict 

movement of hikers and woods workers, and 

suppress other vegetation and slows succession. 

References 

Brayshaw, T.C. 1996. Trees and shrubs of British 

Columbia. UBC Press, Vancouver, BC, Canada. 

373 p. 

Brinkman, K.A. 1974. Rubus L., blackberry, 

raspberry. In: C.S. Schopmeyer, tech. coord. 

Seeds of woody plants in the United States. 

Agriculture Handbook 450. U.S. Department of 

Agriculture, Forest Service, Washington, DC. 

p. 738-743. 

Bruzzese, E. 1998. The biology of blackberry in 

south-eastern Australia. Plant Protection 

Quarterly 13(4): 160-162. 
654 

Ceska, A. 1999. Rubus armeniacus—a correct 

name for Himalayan blackberries. Botanical 

Electronic News ISSN 1188-603X. http://www. 

ou.edu.cas/botany-micro/ben/ben230.html. 3 p. 

Cox, C. 2003. Nonchemical methods for removing 

unwanted blackberry plants. Journal of Pesticide 

Reform 23(1): 10-11. 

Evans, K.J., D.E. Symon, and R.T. Roush. 1998. 

Taxonomy and genotypes of the Rubus 

fruticosus L. aggregate in Australia. Plant 

Protection Quarterly 13(4): 152-156. 

Jacobson, A.L. 2001. Himalaya blackberry: Rubus 

armeniacus Focke. http://www.arthurleej.com/ 

a-himalayabb.html. 4 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Rubus discolor Weihe & Nees, 

Himalayan blackberry. http://plants.usda.gov/ 

cgi_bin/plant_search.cgi?mode=Scientific+Nam

e&keywordquery=Rubus+discolor&earl=plant_

search.cgi. 5 p. 

Shoemaker, J.S. 1978. Small fruit culture. The 

AVT Publishing Company, Inc., Westport, CN. 

357 p. 

Starr, F., K. Starr, and L. Loope. 2003. Plants of 

Hawai’i: Rubus discolor. http://www/jear/ 

prg/starr/hiplants/reports/html/rubus_discolor.ht

m. 6 p. 

Tirmenstein, D. 1989. Rubus discolor. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shurb/ 

rubdis/all.html. 15 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University Press, Provo, UT. 894 p. 
655

Rubus parviflorus Nutt. thimbleberry 

ROSACEAE 

Synonyms: Rubus nutkanus Moc. ex Ser. 

 Rubus lacer Kuntze 

 Rubacer parviflorum Rydb. 

 Bossekia parviflora Greene 

 Rubus nutkanus var. parviflorus Focke 

John K. Francis 

Illustration source: USDA-Forest Service collection, 

Hunt Institute 

General Description.—Thimbleberry, also known 

as western thimbleberry, salmonberry, mountain 

sorrel, and western thimble raspberry, is a 

spineless, deciduous shrub 0.5 to 2.5 m in height. 

Twigs and new stems are green and glandular; old 

stems are covered with tan, papery, shredding 

bark. Stems support relatively few branches. 

Clonal groups of plants are usually represented by 

several to many somewhat dispersed stems that 

have arisen from rhizomes (underground stems). 

Leaves, which have a maple-leaf appearance, are 

simple, alternate, and palmately veined with three 

to seven lobes and have entire to serrate margins. 

The yellow-green to dark-green blades are 4.5 to 

15 cm long and 5.5 to 20 cm broad and are 

supported by long petioles. Perfect, white flowers 

2 to 4 cm broad are grouped in terminal corymbs 

of 2 to 7 flowers. The fruits, which are aggregates 

of tiny drupes, are hemispherical, thin, somewhat 

dry, fragile, and dark pink to bright red, and detach 

easily from the central core (torus), which has five 

sepals at its base. The hard seeds are 1.5 to 2.5 mm 

long (Abrams 1944, Rook 1998, Welsh 1974) 

Range.—Thimbleberry occurs from Alaska to 

western Ontario, from Michigan westward, and 

south through California, Arizona, New Mexico, 

and northern Mexico (Natural Resources 

Conservation Service 2003, Welsh 1974, Welsh 

and others 1987). It is also reported in 

Massachusetts, probably as a naturalized 

population. There are two recognized varieties of 

thimbleburry, var. parviflorus, accounting for 

virtually all of the range, and var. velutinus (Hook. 

& Arn.) Greene, which is densely hairy on the 

underside of leaves, found in California (Abrams 

1944, Natural Resources Conservation Service 

2003). 

Ecology.—Thimbleberry grows on most types of 

moist soils, including skeletal soils. The species is 

most common in cool, moist sites, particularly on 

north slopes. Thimbleberry also occurs on dry, 

exposed ridges in high-elevation areas. It grows 

from near sea level to 2,900 m in elevation in 
656 

Wyoming (Tirmenstein 1989). Thimbleberry is 

moderately shade tolerant. Although it is most 

abundant and grows and fruits best in 60 to 100 

percent sunlight, the species persists in the 

understory of closed stands (Tirmenstein 1989). It 

occurs both as scattered individual plants and as 

large continuous stands. Habitat includes burned 

sites, logged areas, avalanche tracks, roadsides, 

natural shrublands, hardwood forest understories, 

understories of low to moderate basal area conifer 

forests, and river overflow terraces and shorelines. 

It is a part of a large number of different plant 

communities. Thimbleberry is resistant to fire, 

readily resprouting from rhizomes. In fact, the 

species generally benefits from all kinds of 

disturbance. It usually appears during the first year 

after disturbance and often dominates the 

understories of logged areas within 5 years. 

Moderate to light thimbleberry stands can serve as 

a good protective cover for conifer seedlings. In 

recreational sites, the species showed relatively 

low resistance to trampling (Tirmenstein 1989). 

Reproduction.—Thimbleberry generally blooms 

in June and July. Its fruits mature in July and 

August (Borialforest.org 2003). However, at the 

extremes of its range, it may bloom anywhere 

from May to September and fruit between late 

June through September (Tirmenstein 1989). The 

flowers are insect pollinated. Good fruit and seed 

crops are usually produced every year except in 

high-elevation areas where production may be 

unreliable (Tirmenstein 1989). The seeds are 

dispersed by birds and mammals that eat the fruits, 

and by gravity. The seeds accumulate in the soil 

and duff to form a seed bank and germinate in 

great numbers following fire and other 

disturbances that remove the forest canopy (Rook 

1998). After establishment by seeds, stands 

thicken through sprouts from rhizomes (Rook 

1998). 

Growth and Management.—The stems (or 

canes) live for 2 or 3 years. In the first year, canes 

grow in height and develop only leaves. During 

the second and third years height growth continues 

and flowers and fruits are produced. The canes 

then die and are replaced by others that sprout 

from rhizomes. Maximum height of current canes 

is reached within 10 years after establishment 

(Tirmenstein 1989). Seeds must be collected by 

hand and should be cleaned of adhering fruit. 

Seeds may be sown in the fall for spring 

germination or may be warm (20 to 30 °C for 90 

days) and cold stratified (2 to 5 °C for 90 days) for 

spring sowing. Also, scarifying with H2SO4 prior 

to the cold stratification may enhance germination. 

Tests have given maximum germination estimates 

of 62 percent (Tirmenstein 1989). Hardwood 

cuttings can be easily rooted. New plants can also 

be started from rhizome cuttings and plant 

divisions. Wildlings survive well but are slow to 

become established (Washington State University 

Cooperative Extension 2003). Occasionally, it is 

desirable to release conifer seedlings in and under 

thimbleberry thickets or to make way for other, 

more palatable forage species. The species is 

moderately susceptible to herbicides. Picloram, 

2,4-D, and glyphosate have been effective in 

appropriate applications (Tirmenstein 1989). 

Benefits.—Thimbleberry is an important shrub of 

the understory and forest openings. It helps protect 

the soil and adds to the beauty of the forest. The 

species is occasionally planted as an ornamental 

for its fragrant flowers and brilliant orange to 

maroon fall foliage (Borialforest.org 2003, Rook 

1998). It is also occasionally planted in 

conservation plantings in disturbed areas. 

Thimbleberry is relatively low in energy and 

protein (4 to 8 percent), and is little used by cattle 

and horses and is only fair forage for sheep. It is 

sometimes important for deer, elk, and other wild 

ungulates during the summer while the leaves are 

still present. Rodents consume bark, buds, and 

foliage to a limited extent. On the other hand, the 

fruits are an important food item for numerous 

wild mammals and birds. Thickets of thimbleberry 

are also important escape, resting, and 

reproductive cover for many species of wildlife 

(Tirmenstein 1989). Fruits are certainly edible to 

humans but reports vary greatly on their 

palatability (Clark 1976, Welsh 1974). The author 

has eaten them for years and finds them inferior to 

cultivated raspberries (Rubus idaeus L.) but well 

worth the trouble of picking while hiking in the 

forest. The fruits are made into an excellent jelly 

and were once dried for later use by Native 

Americans. The tender young shoots are juicy and 

sweet and can be boiled or eaten fresh. The leaves 

are sometimes made into herb teas 

(Borialforest.org 2003, Washington State 

Department of Transportation 2003). Native 

Americans applied poultices of leaves to burns and 

wounds and took decoctions of roots as a tonic, for 

vomiting, and certain internal disorders (Moerman 

1986). 
657

References

Abrams, L. 1944. Illustrated flora of the Pacific 

States. Vol. 2. Stanford University Press, 

Stanford, CA. 635 p. 

Borialforest.org. 2003. Rubus parviflorus, 

thimbleberry. http://www.borialforest.org/ 

shrubs/shrub41.htm. 2 p. 

Clark, L.J. 1976. Wild flowers of the Pacific 

Northwest. Gray’s Publishing Limited, Sidney, 

BC, Canada. 604 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

Americans. Technical Report 19. Museum of 

Anthropology, University of Michigan, Ann 

Arbor, MI. 534 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Rubus parviflorus Nutt. 

http://plants.usda.gov/cgi_bin/topics.cgi. 4 p. 

Rook, E.J.S. 1998. Rubus parviflorus, 

thimbleberry. http://www.rook.org.earl/bwca/ 

hature/shurbs/rubuspar.html. 7 p. 

Tirmenstein, D. 1989. Rubus parviflorus. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shurb/ 

rubpar/all.html. 37 p. 

Washington State Department of Transportation. 

2003. Environmental affairs: Rubus parviflorus. 

http://www.wsdot.wa.gov/environment/eao/ 

culres/ethbot/q-s/RubusPar.htm. 2 p. 

Washington State University Cooperative 

Extension 2003. Native plants: Rubus 

parviflorus var. parviflorus. http://cahedb. 

wsu.edu/nativePlant/scripts/webDisplayText.asp

?ID=nv044. 1 p. 

 

Welsh, S.L. 1974. Anderson’s flora of Alaska. 

Brigham Young University Press, Provo, UT. 

724 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1987. A flora of Utah. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University Press, Provo, UT. 894 p. 
658 

Rubus rosifolius Sm. roseleaf raspberry 

ROSACEAE 

Synonyms: Rubus coronaries (Sims) Sweet 

John K. Francis 

General Description.—Roseleaf raspberry, also 

known as wild raspberry, Mauritius raspberry, 

Queensland raspberry, fresa de montaña, 

framboisier, and akalakala, is an evergreen 

scrambling shrub, freestanding to 0.75 m, taller 

when supported on other vegetation, and extending 

laterally 2 m or more. Individual plants usually 

have several arching stems (canes) with basal 

diameters of 0.5 to 1 cm and have a 3-mm pith. 

Stems are green with reddish coloration at the 

nodes and lower portions. Wood of the lower 

stems is moderately hard and brittle. The stems, 

twigs, and leaf petioles and rachises are armed 

with short, straight or curved prickles. Adult plants 

have a fibrous lateral root system. Leaves are oddpennate with mostly seven ovate to linear, doubly 

serrate leaflets. The terminal or axile 

inflorescences contain one to four, 1.5 to 3 cm 

diameter flowers with five white petals. The 

globose to narrowly oblong aggregate fruit is 

bright red and juicy when ripe. The pitted 

(foveolate), yellow seeds (pyrenes) are about 1.5 

mm long (author’s observation, Howard 1988, 

Liogier 1985, Lu and Boufford 2002). 

Range.—Roseleaf raspberry is apparently native 

from China and Taiwan south through Indochina 

and Indonesia along the eastern coast of Australia, 

Mauritius, New Caledonia, the Solomon Islands, 

and Vanuatu, and eastward as far as northeastern 

India. However, it is difficult to tell the native 

from naturalized range that extends from Japan to 

Africa and through the Caribbean (including 

Puerto Rico) and Brazil and to many of the islands 

of the Pacific (including Hawaii) (Liogier 1985, 

Lu and Boufford 2002, Pacific Island Ecosystems 

at Risk 2002, Peixoto 2002). 

Ecology.—Although roseleaf raspberry invades 

forest understories in some habitats (Pacific Island 

Ecosystems at Risk 2002), it is moderately shade 

intolerant, requiring partial sun to flower and fruit; 

it does best in full sun. Preferred habitats include 

natural and artificial openings in primary and 

secondary forest, such as tree-fall gaps, landslides, 

abandoned fields, neglected pastures, and 

roadsides. Roseleaf raspberry has a high moisture 

requirement, which usually means more than 1800 

mm of mean annual precipitation and no severe 

annual droughts. The species usually grows at 

moderate to high elevations. It is reported at 2,200 

m in Tahiti and 1,730 m in Hawaii (Pacific Island 

Ecosystems at Risk 2002). A variety of welldrained to poorly-drained soils are colonized. 

Reproduction.—In China, roseleaf raspberry 

flowers in March through May and fruits in June 

and July (Lu and Boufford 2002). The species 

blooms and fruits throughout the year in 

continually moist areas of Puerto Rico and 

Equatorial Africa (Burkill 1997). Rubus species 

are pollinated by insects, particularly honey bees 

(Apis mellifera L.) (Dalaplane and Mayer 2000). 

Fresh fruits collected in Puerto Rico averaged 

1.516 + 0.102 g/fruit. Air-dried seeds separated 

from them averaged 0.00045 g/seed or 2.2 million 

seeds/kg. The seeds are dispersed by birds and 

rodents (Pacific Island Ecosystems at Risk 2002). 

Stems root (layer) whenever they come in contact 

with the soil, which helps it fill in newly colonized 

habitat. 

Growth and Management.—Stems (canes) of 

roseleaf raspberry originating from sprouts grow 1 

m or more per year and live about 1.5 years. New 

sprouts then arise to replace them and the plant 

lives on (potentially) for several years. In Puerto 

Rico, clumps and thickets of roseleaf raspberry 

usually disappear in 2 or 3 years due to mounting 

competition and encroaching shade from trees, 
659

shrubs, and herbs. The species is occasionally 

planted from wildlings and rooted layers for 

personal use in gardens. It is controlled along with 

other vegetation by slashing or mowing, and by 

tillage to prepare ground for crops. 

Benefits.—Roseleaf raspberry helps return 

disturbed areas to forest vegetation, protects the 

soil, and furnishes food and cover for wildlife. 

Leaves and tender shoot are browsed by livestock. 

The berries are eaten out of hand and made into 

drinks and preserves throughout its range. Fresh 

fruits are sold nearly year-round in Ugandan 

markets (Burkill 1997). The fruits contain 

relatively high (1.58 and 12.93 mg/100g fresh 

weight respectively) of vitamins E and C (Wei and 

Payne 2002). Roseleaf raspberry has been grown 

as a ground cover in Cameroon plantations 

(Burkill 1997). A double petaled variety (R. 

rosifolius var. coronaries (Sims) Focke) is grown 

as an ornamental (Garden Plant Conservation 

2002). Infusions of flower petals are used to 

control diarrhea, vomiting, and other flu 

symptoms, and as a tonic (Liogier 1990). An 

infusion of the leaves was brewed in former times 

to relieve menstrual cramps, morning sickness, and 

labor pains (Nortan 2002). 

References

Burkill, H.M. 1997. The useful plants of West 

Tropical Africa. Ed. 2. Vol. 4. Royal Botanical 

Garden, Kew, UK. 960 p. 

Dalaplane, K.S. and D.F. Mayer. 2000. Crop 

pollination by bees. CABI Publishing, 

Wallinford, UK. 352 p. 

Garden Plant Conservation. 2002. Rubus rosifolius

‘Coronarius.’ http://www.manntaylor.com/ 

plantweek9c.html. 7 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc. 566 p. 

Lu, L. and D.E. Boufford. 2002. Rubus Linnaeus 

[Draft]. In: A.R. Brach and Hong S., eds. Flora 

of China, Harvard University Herbaria, 

Cambridge, MA. http://hua.huh.harvard.edu/ 

china/mss/volume09/Rosaceae-AGH_Rubus_ 

coauthoring.htm. 200 p. 

Nortan, A. 2002. Bush tucker: Roseleaf raspberry 

(Rubus rosifolius). Rumbalara Environmental 

Ecucation Centre, Sydney, NSW, Australia. 

http://rumbalara-e.schools.nsw.edu.au/ 

bushtucker/ Rubus_rosifolius.htm. [not paged]. 

Pacific Island Ecosystems at Risk. 2002. Rubus 

rosifolius Sm., Rosaceae. http://hear.org/ 

pier_v3.3/ruros.htm. 2 p. 

Peixoto, M. 2002. Plantas Brasileiras: Rosaceae, 

Rubus rosifolius. http://sites.uol.com.br/ 

mpeixoto/outras/rosifolius.html. 2 p. 

Wei, C. and T.J. Payne. 2002. Raspberry (Rubus 

L.). Washington Red Raspberry Commission, 

Lyndon, WA. http://www.red-raspberry.org/ 

china.pdf. 7 p. 
660 

Salix arctica Pallas arctic willow

SALICACEAE 

Synonyms: Salix anglorum Cham. 

Salix brownie (Anderss.) Bebb 

Salix caespitosa Kennedy 

Salix pallasii Anderss. 

Salix petrophila Rydb. 

Juanita A.R. Ladyman 

Illustration source: USDA-Forest Service collection, 

Hunt Institute 

General Description.—Salix arctica is commonly 

known as arctic willow. This deciduous, prostrate, 

trailing shrub is usually less than 10 cm tall and 

commonly forms mats. However, its growth 

habitat is variable, and in the Pacific Northwest it 

grows up to 50 cm tall (Pojar and MacKinnon 

1994). The broadly oval, alternate leaves are 2 to 6 

cm long and sparsely hairy when mature. They are 

a glossy green on the upper side and pale greengrey below (Kershaw and others 1998, Fertig and 

Markow 2001). Plants are dioecious, possessing 

male and female flowers on separate, “unisexual” 

plants. The flowers of both sexes are arranged in 

dense, erect clusters or catkins. The female catkins 

are densely hairy, 1 to 8 cm (usually 2 to 4 cm) 

long and are borne on lateral, sometimes appearing 

terminal, branchlets (Kershaw and others 1998, 

Fertig and Markow 2001). The male, staminate, 

catkins are up to 4 cm long. Flowers appear with 

the leaves. There are several more synonyms than 

those listed above and may be referred to in 

Kartesz (1994), Argus and others (1999), and 

Dallwitz (1980). Identification may be difficult. 

Environment can affect the morphology to such an 

extent it appears a unique species is present (Argus 

and others 1999). Although many subspecies have 

been described, they have generally been discarded 

in favor of the type species. In addition, hybrids 

are reported, and introgression with other species, 

such as S. cascadensis in Utah and Salix reticulata

in Russia, is well documented (Hultén 1968, 

Komarov 1936, Pojar and MacKinnon 1994, 

Welsh and others 1993). 

Range.—Arctic willow is a circumpolar species 

occurring in North America, Europe, and Asia 

(Britton 1901, Fertig and Markow 2001, Hultén 

1968, Komarov 1936). In North America it grows 

in the sub-alpine and alpine tundra zones from 

Quebec to Alaska and south, through the Rocky 

Mountains, into New Mexico (Kershaw and others 

1998). Occurrences are also reported from Utah 

(Welsh and others 1993) and the Ruby Mountains 

of Nevada (Charlet 1994). It has disjunct 

occurrences on the west coast of the U.S.A., 

occurring in California and in the northern 

Cascade Mountains in Washington and in the 

Wallowa Mountains in Oregon (Hitchcock and 

Cronquist 1973). 

Ecology.—Arctic willow grows in dry meadows, 

rocky tundra, fellfields, and in relatively moist 

sites in fen lands and at the edge of snowfields. 

Habitat tends to be related to latitude. In the 

mountains of New Mexico it is an obligate wetland 

species, whereas at higher latitudes it grows in 

both dry and moist habitats (Cooper and others 

1997, Dick-Peddie 1993, Fertig and Markow 2001, 

Hultén 1968). It may be locally dominant in some 

cushion plant communities and co-occur with 

other species of dwarf willow such as Salix 

reticulata L. (snow willow). In New Mexico 

tundra, arctic willow and snow willow are often 

found associated with lichens and other vascular 

plants in snowbank and rivulet communities 

(Dick-Peddie 1993). Similarly in Montana and 
661

elsewhere in the Rocky Mountains, a common S. 

arctica/Polygonum bistortoides Pursh community 

was identified in “water-receiving positions” 

(Cooper and others 1997). However, the latter sites 

were also reported to have a good probability of 

experiencing wind scouring, and some of the 

associated vegetation suggested the area could be 

relatively dry (Cooper and others 1997). The 

elevation at which it occurs is also associated with 

latitude. Plants are only found above 3,350 m in 

New Mexico, whereas they occur at sea level and 

up to 700 m in northern parts of its range, such as 

Greenland (Carter 1997, Argus and others 1999). 

It grows in a range of soils, from acidic to 

calcareous (Argus and others 1999, Cooper and 

others 1997, Pojar and MacKinnon 1994). A 

mutualistic association exists between arctic 

willow and ectomycorrhizal fungi that are 

primarily agarics, or gilled mushrooms (Cripps 

and Horak 2002). 

Reproduction.—Arctic willow reproduces by 

seed and also vegetatively by rooting at the nodes 

of stems (Fertig and Markow 2001). Flowering is 

from June into August. On calm, sunny days the 

female catkins can be up to 8.5 o

C warmer and the 

male catkins up to 7 oC warmer than the 

surrounding air temperature (Kershaw and others 

1998). The warmer temperatures speed pollen and 

seed development and also attract insects 

(Kershaw and others 1998). 

Growth and Management.—Seeds have a 

physiological dormancy and require a 30-day 

stratification period for germination (Baskin and 

Baskin 2001). Seeds germinate at 25 o

C although 

minimum temperatures for germination are not 

defined (Densmore and Zasada 1983). Thompson 

(1992) reported that seeds of Salix species are 

“normally absent” from the seed bank in alpine 

and arctic communities. Individual shrubs live for 

60 to 85 years although one specimen was reported 

to be 180 and another 236 years old (Savile 1979, 

Raup 1959). Habitat appears to influence the sex 

that is most abundant. Dawson (1987) reported 

female plants were significantly more numerous 

than male plants in mesic-wet, more fertile, low 

soil-temperature sites, whereas male plants were 

most prevalent in drier, less fertile sites. In 

addition, both habitat and sex may influence a 

plant’s response to environmental conditions such 

as temperature. In a 3-year gas exchange field 

study, male and female willows from dry and wet 

habitats were subjected to passively enhanced 

summer temperature using small open-top 

chambers (Jones and others 1999). Overall net 

assimilation was higher in the dry habitat than in 

the wet habitat, and higher in females than in 

males, although there appeared to also be some 

habitat-sex interactions (Jones and others 1999). 

Elevated temperature enhanced development and 

growth of both male and female catkins, but the 

response of the leaves to elevated temperatures 

was more variable particularly among male 

individuals (Jones and others 1999). These 

findings indicate the importance of defining both 

sex and habitat conditions when making 

observations on this dioecious species, and they 

may also have significance in predicting the 

response of Arctic willow to global climate 

change. 

Benefits.—The majority of Salix species provide 

important browse for wildlife and livestock within 

their range (Dayton 1931). Arctic willow, 

particularly the young leaves stems and buds, 

provide valuable browse for ptarmigan and other 

arctic animals including muskox and reindeer 

(Kershaw and others 1998, Komarov 1936, Tolven 

and others 2001). Arctic willow is tolerant of 

varied conditions, and cuttings have been rooted 

and used in restoration projects (Bittman 1997). 

Willow leaves and bark are commonly used for 

medicinal purposes and the roots, twigs, and bark 

are used for a multitude of fiber work, such as 

basketry and clothing by Native American peoples 

(Moerman 1998). Arctic willow has been used for 

similar purposes by peoples native to Asian Arctic 

regions (Moerman 1998). The Yakuts have used 

an infusion as a substitute for tea called “chaitalak” (Komarov 1936). It is used as fuel in some 

regions where it is the only source of wood 

(Kershaw and others 1998). Arctic willow is 

cultivated, and it is especially popular for rock 

gardens (Bailey and others 1976, Pojar and 

MacKinnon 1994).

References. 

Argus, G.W., C.L. McJannet, and M.J. Dallwitz. 

1999 onwards. ‘Salicaceae of the Canadian 

Arctic Archipelago: Descriptions, Illustrations, 

Identification, and Information. http://www. 

mun.ca/biology /delta/arcticf/. 

Bailey, L.H., E.Z. Bailey, and the staff of the 

Liberty Hyde Bailey Hortorium. 1976. Hortus 

Third. Macmillian Publishing Company, New 

York. 1,290 p. 
662 

Baskin, C.C. and J.M. Baskin. 2001. Seeds, 

ecology, biogeography, and evolution of 

dormancy and germination. Academic Press, 

New York. 666 p. 

Bittman, K.K. 1997. High elevation native species 

island model for mine reclamation. Quintette 

Operating Corporation, Tumbler Ridge, BC. 

http://www.teckcominco.com/presentations/kbhighelv-1997.pdf. 

Britton, N.L. 1901. Manual of the Flora of the 

northern states and Canada. Henry Holt and 

Company, New York. 1,080 p. 

Carter, J.L. 1997. Trees and shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Charlet, D. 1994. Plant Species in the Ruby 

Mountains, Nevada. 

http://www.brrc.unr.edu/data/plants/ ruby.html. 

Cooper, S.V., P. Lesica, and D.Page-Dumrose. 

1997. Plant community classification for alpine 

vegetation on the Beaverhead National Forest, 

Montana. General Technical Report INT-362. 

USDA Forest Service Intermountain Research 

Station, Ogden, UT. 61 p. 

Cripps, C.L., and E. Horak. 2002. A survey 

of alpine agaricales in the Rocky Mountains. 

http://plantsciences.montana.edu/alpinemushroo

ms/ rocky_mountain_ alpine_field_sites.htm 

Dallwitz, M.J. 1980. A general system for coding 

taxonomic descriptions. Taxon, 29: 41–46. 

Dawson, T. E. 1987. Comparative 

ecophysiological adaptations in arctic and alpine 

populations of a dioecious shrub, Salix arctica

Pall. Ph.D. dissertation, University of 

Washington, Seattle, WA. 208 p. 

Dayton, W. A. 1931. Important western browse 

plants. Miscellaneous Publication 101. U.S. 

Department of Agriculture, Washington DC. 

214 p. 

Densmore, R. and Zasada, J.C. 1983. Seed 

dispersal and dormancy patterns in northern 

willows: Ecological and evolutionary 

significance. Canadian Journal of Botany 61: 

3207-3216 

Dick-Peddie, W.A. 1993. New Mexico 

vegetation. University of New Mexico Press, 

Albuquerque, NM. 244 p. 

Fertig, W. and S. Markow. 2001. Guide to the 

willows of Shoshone National Forest. General 

Technical Report RMRS-GTR-83. U.S. 

Department of Agriculture, Forest Service 

Rocky Mountain Research Station, Fort Collins, 

CO. 80 p. 

Hitchcock, C.L., and A. Cronquist. 1973. Flora of 

the Pacific Northwest. University of Washington 

Press, Seattle, WA. 730 p. 

Hultén, E. 1968. Flora of Alaska and neighboring 

territories Stanford University Press, Stanford, 

CA. p. 340-341. 

Jones M.H., S.E. Macdonald, and G.H.R. Henry. 

1999. Sex- and habitat-specific responses of a 

high arctic willow, Salix arctica, to experimental 

climate change. Oikos 87(1): 129-138. 

Kartesz, J.T. 1994. A synonymized checklist of the 

vascular flora of the United States, Canada and 

Greenland. Vol.1 - Checklist. 2nd Ed. Timber 

Press, Portland, OR. 622 p. 

Kershaw, L., A. MacKinnon, and J. Pojar. 1998. 

Plants of the Rocky Mountains. Lone Pine 

Publishing, Edmonton, Canada. 383 p. 

Komarov, V.L. 1936. Flora of the U.S.S.R. 

Volume V. Izdatel’stvo Akademii Nauk SSSR, 

Moscow-Leningrad, U.S.S.R. Translated from 

Russian by N. Landau 1970. Israel Program for 

scientific translations, Jerusalem, Israel and The 

Smithsonian Institution Washington, D.C. p. 

36-37. 

Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. 927 p. 

Pojar, J. and A. MacKinnon. 1994. Plants of the 

Pacific Northwest coast–Washington, Oregon, 

British Columbia and Alaska. Lone Pine 

Publishing, Edmonton, Canada. 526 p. 

Raup, H.M. 1959. The willows of boreal western 

America. Contributions from the Gray 

Herbarium of Harvard University 185: 3–95. 
663

Savile, D.B.O. 1979. Ring counts in Salix arctica 

from northern Ellesmere Island. Canadian FieldNaturalist 93: 81-82. 

Thompson, K. 1992. The functional ecology of 

seed banks. In: M. Fenner, ed., Seeds, the 

ecology of regeneration in plant communities. 

CAB International, Wallingford, Oxon, U.K. 

373 p. 

Tolven, A. J., Schroderus, and G.H.R. Henry. 

2001. Age- and stage-based bud demography of 

Salix arctica under contrasting muskox grazing 

pressure in the High Arctic. Evolutionary 

Ecology 15: 4-6 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1993. A Utah flora. Second Edition, 

revised. Brigham Young University, Provo, UT. 

986 p. 
664 

Salix exigua Nutt. coyote willow 

SALICACEAE 

Synonyms: Salix interior Rowlee 

James E. Nellessen 

General Description.—Coyote willow is also 

known as sandbar willow, basket willow, 

narrowleaf willow, slender willow, riverbank 

willow, acequia willow, long-leaved willow, and 

gray willow (Elmore 1976). It generally ranges in 

height from 0.5 to 3 m, is occasionally a small tree 

to 6 m and trunk diameter to 13 cm, with one 

author (Dorn 1998) reporting up to 17 m in height. 

Coyote willow is a clonal species that spreads by 

creeping rootstocks that generate new shoots 

forming multistemmed dense thickets. Coyote 

willow is one of few willow species that reproduce 

this way. As with other willows the species is 

dioecious. The bark is greenish to reddish on 

young twigs and ashy gray to grayish brown on 

older stems. Lenticels are elliptic and raised. The 

buds are reddish brown. The leaves are alternate, 

simple, long and narrow, linear to lanceolate in 

shape, tapering at both ends, generally six or more 

times longer than wide, 2 to 16 cm long, 4 to 15 

mm wide, the margins are smooth or irregularly 

toothed, with one to five teeth per cm, and the 

petioles are short, 2 to 7 mm long (Argus 1986, 

Britton and Brown 1913, Carter 1997, Gleason and 

Cronquist 1963, Great Plains Flora Association 

1986, Martin and Hutchins 1980-81, Newsholme 

1992). Upon emergence the young twigs and 

leaves are densely silvery hairy, but with time and 

aging, the upper leaf surface reduces to a 

moderately gray hairy or smooth surface, 

sometimes slightly shiny. Plants in moist sites at 

higher elevations in the mountains may be 

noticeably less hairy (Mozingo 1987). The lower 

leaf surface generally remains whitish and much 

hairier. Coyote willow is one of few willow 

species with stomata on both leaf surfaces (Argus 

1986). Stipules are small and sometimes absent. 

The chromosome number is 2n = 38 (Dorn 1998). 

Taxonomy.—Coyote willow has a complicated 

taxonomy. It is a member of Salix section 

Longifoliae. Several botanists now consider S. 

interior as a much less hairier to smooth form of S. 

exigua that occurs in the Great Plains and Eastern 

United States, while variety exigua is the hairier 

Western form (Argus 1986, Carter 1988, Dorn 

1998, Gleason and Cronquist 1963, Great Plains 

Flora Association 1986, Weber and Wittman 

2001a, b). The Eastern interior is more distinctly 

toothed, leaf veins are more prominent, and the 

fruits (capsules) slightly longer. But characteristics 

are highly variable across the entire range of this 

plant. Under this taxonomic organization and 

using the Dorn (1998) treatment, there would be 

two subspecies: S. exigua subsp. exigua (coyote 

willow) and S. exigua subsp. interior (Rowlee) 

Cronq. (sandbar willow). The Western subspecies 

is divided into four varieties: var. exigua, var. 

hindsiana (Benth.) Dorn, var. sessilifolia (Nutt.) 

Dorn, and var. columbiana Dorn. The Eastern 

subspecies, interior, contains no varieties under 

Dorn’s treatment. There are various synonyms 

(Argus 1986, Britton and Brown 1913, Dorn 1998, 

Gleason and Cronquist 1963, Great Plains Flora 

Association 1986, Kearney and others 1951, 

Newsholme 1992). A close to complete list of 34 

synonyms can be found in Kartesz (1994). 

Hybridization is likely an important factor 
665

complicating the taxonomy for this species, the 

entire section, likely a syngameon undergoing 

geographic speciation (Brunsfield and others 

1992). 

Range.—This is a widely distributed species of 

plains and lower elevations to lower montane 

habitats in the Western United States, as well as 

throughout Eastern deciduous forests if S. interior

is considered a subspecies. The Western form 

(exigua) occurs from Saskatchewan to Eastern 

British Columbia in Canada, south through the 

United States to Eastern and Southern California, 

Arizona, New Mexico, and into Northern Mexico 

and the Western Great Plains. The Eastern form or 

subspecies (interior) occurs from New Brunswick 

in Eastern Canada, westward throughout the Great 

Plains, as far west as Idaho, north to Alaska, south 

to Eastern New Mexico, and extends into the 

Southeast along a line from Northern Virginia to 

Tennessee and Louisiana. The two subspecies 

overlap in the Western Great Plains. In the 

Western mountains, coyote willow can be found as 

high as 2,900 m in elevation. 

Ecology.—Coyote willow commonly forms 

thickets along streams and rivers and is an 

important riparian species stabilizing streambanks. 

It will commonly colonize sand bars along rivers, 

hence the frequently used alternate common name 

sandbar willow. It also occurs in ditches, and on 

the edges of swamps, lakeshores, and other 

wetland habitats. This species will grow anywhere 

groundwater is close to the surface. This includes 

sites where human activities result in surface water 

availability on a reasonably regular and reliable 

basis (exclusive of saline conditions). Coyote 

willow is also one of the few willow species that 

can be found in lower elevation, hot desert areas in 

the Western United States. But because if its wide 

elevational range it is both a floodplain-plains and 

montane riparian species (Dick-Peddie 1993). Due 

to the extensive rootstock and clonal nature of the 

species, cut stands of coyote willow will 

regenerate relatively quickly. The presence of an 

aerenchyma-containing wetland plant, Typha 

latifolia L., may facilitate a nonaerenchyma 

bearing species like coyote willow to survive in 

low oxygen wetland soils (Callaway and King 

1996). Leaf water potentials as low as -2.7 MPa 

have been measured for high elevation populations 

in Wyoming (Foster and Smith 1991). Coyote 

willow leaves within the Colorado River (Arizona) 

decomposed substantially more slowly than those 

of Populus fremontii S. Watson and salt cedar 

(Tamarix ramosissima Ledebour), with 52 percent 

of the leaf mass remaining after 142 days 

(Pomeroy and others 2000). A number of insects 

attack coyote willow, including leaf mining 

beetles, lepidopterans, sawflies (Euura spp.), and 

galling insects. A bud galling midge (Rabdophaga

sp.) affects both terminal and lateral shoots, 

arresting growth of the immediate shoot (Declerck 

and Price 1994). Coyote willow responds by 

producing new lateral buds and shoots, but many 

do not survive. 

Reproduction.—The male flowers are in densely 

flowered catkins from 1 to 6 cm in length, each 

flower with two stamens, the filaments hairy. The 

female catkins are loosely flowered, from 2 to 10 

cm long, the ovaries are smooth to slightly silky 

with a short or almost nonexistent style. Each 

ovary bears nine to 15 ovules (Argus 1986). 

Catkin scales are yellow and deciduous. The 

catkins appear with the leaves or shortly after the 

leaves in spring (March to May) on short leafy 

stalks. The flowers are bee pollinated. There may 

even be a second blooming period from June to 

August (Argus 1986). The fruits are smooth 

capsules from 4 to 8 mm long, green, turning 

yellow to reddish brown upon ripening. The fruits 

open by two valves releasing tiny seeds (22,000/g 

[Brinkman 1974]) with long hairs that aid in wind 

dispersal. The seeds are short-lived, do not require 

stratification, must fall on a moist substrate, and 

can germinate within 12 to 24 hours under proper 

conditions (Young and Young 1992). Successful 

germination is reported at 22 C for the Western 

subspecies exigua (Brinkman 1974) and 5 to 25 

C for the Eastern subspecies interior (Densmore 

and Zasada 1983, Baskin and Baskin 2001). 

Flower-and-fruit-bearing shoots have been 

observed to sprout new leafy shoots and become 

new side branches (Argus 1986). 

Growth and Management.—Along the Rio 

Grande in New Mexico, clearing riparian areas of 

the invasive, nonnative salt cedar prior to peak 

river flows in the spring can facilitate the 

reestablishment of natives such as coyote willow 

(Taylor and others 1999). In Alberta, Canada, 

successful seedling establishment of several 

riparian species, including coyote willow, occurred 

best on deposits 80 to 120 cm above the late 

summer stream flow (Rood and Mahoney 2000). 

In Manitoba, Canada, average annual height 

increases were 30 cm, stem diameter increases 

were 2.6 mm, flowering began in 2 to 3 year old 

stems, stem mortality was greatest in 3 to 6 year 
666 

old clones, entire plant senescence first occurred at 

12 years, and the oldest plant recorded was 31 

years on a point bar in the Assiniboine River 

(Ottenbreit and Staniforth 1992). 

Benefits.—Because coyote willow forms dense 

thickets and spreads clonally, it is an important 

stabilizer of streambanks. It is frequently and 

successfully used in riparian and wetland 

revegetation and restoration. It is an important 

food for wildlife such as deer and beaver. It is 

considered good browse for sheep and fair for 

cattle, but cattle generally find it more palatable 

later in the growing season (Mozingo 1987, 

Stubbendieck and others 1997, USDA 1937). 

Coyote willow is used in intensive culture coppice 

plantations (Aravanopoulos and others 1999), was 

one of 32 woody plants tested for biomass 

gasification, and was one of three species that 

exceeded 0.30 liter methane per gram of volatile 

solids (Turick and others 1991). Coyote willow 

has been successfully grown in tissue culture with 

roots developing from shoots longer than 1 cm 

(Stoehr and others 1989). Native Americans have 

used the slender stems in making baskets and have 

made a tea from the bark to treat fever and 

headache. Willow bark, as with most willows, is 

bitter and acts as an astringent for diarrhea, has 

been used for fevers, pain, arthritis, rheumatism, 

and a poultice for cuts, ulcers, and rashes. Salicylic 

acid, commonly used in asperin as acetylsalicylic 

acid, was originally derived from salicin contained 

in the bark of willows (Foster and Duke 1990). 

Salicin is a member of the chemical class of 

phenols and affords some protection against 

browsing mammals such as rabbits. It tends to 

attain a greater concentration in low growing 

willows but becomes lost in larger, taller willows 

(Harborne 1988). Oestriol (an oestrogen), a human 

sex hormone, has also been found in willow 

flowers. 

References

Aravanopoulos, F.A., K.H. Kim, and L. Zsuffa. 

1999. Genetic diversity of superior Salix clones 

selected for intensive forestry plantations. 

Biomass and Bioenergy 16(4): 249-255. 

Argus, G.W. 1986. The Genus Salix (Salicaceae) 

in the Southeastern United States. Systematic 

Botany Monographs, Vol. 9. The American 

Society of Plant Taxonomists, USA. 170 p. 

Baskin, C.C. and J.M. Baskin. 2001. Seeds: 

Ecology, Biogeography, and Evolution of 

Dormancy, and Germination. Academic Press, 

San Diego, CA. 666 p. 

Brinkman, K.A. 1974. Salix L. Willow. In: C.S. 

Schopmeyer, tech. coord. Seeds of Woody 

Plants in the United States. U.S. Department of 

Agriculture Forest Service, Handbook 450. 

Washington, DC. p. 746-750. 

Britton, N.L. and A. Brown. 1913 (1970 Dover 

edition). An Illustrated Flora of the Northern 

U.S. and Canada, Vol. 1. Dover Publications 

Inc., New York. 680 p. 

Brunsfield, S.J., D.E. Soltis, and P.S. Soltis. 1992. 

Evolutionary patterns and processes in Salix: 

evidence from chloroplast DNA. Systematic 

Botany 17(2): 239-256. 

Callaway, R.M. and L. King. 1996. Temperature 

driven variation in substrate oxygenation and the 

balance of competition and facilitation. Ecology 

77(4): 1,189-1,195. 

Carter, J.L. 1988. Trees and Shrubs of Colorado. 

Johnson Books, Boulder, CO. 165 p. 

Carter, J.L. 1997. Trees and Shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Declerck, F.R. and P.W. Price. 1994. Impact of a 

bud-galling midge on bud populations of Salix 

exigua. Oikos 70(2): 253-260. 

Densmore, R. and J.C. Zasada. 1983. Seed 

dispersal and dormancy patterns in northern 

willows: ecological and evolutionary 

significance. Canadian Journal of Botany 61: 

3,207-3,216. 

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation: Past, Present and Future. University 

of New Mexico Press, Albuquerque, NM. 244 p. 

Dorn, R.D. 1998. A taxonomic study of Salix

section Longifoliae (Salicaceae). Brittonia 

50(2): 193-210. 

Elmore, F.H. 1976. Shrubs and Trees of the 

Southwest Uplands. Southwest Parks and 

Monuments Association, Tucson, AZ. 214 p. 
667

Foster, S.A. and J.A. Duke. 1990. A Field Guide to 

Medicinal Plants (Eastern/Central). Peterson 

Field Guide Series, Houghton Mifflin Co., 

Boston, MA. 366 p. 

Foster, J.R. and W.K. Smith. 1991. Stomatal 

conductance patterns and environment in high 

elevation phreatophytes of Wyoming, USA. 

Canadian Journal of Botany 69(3): 647-655. 

Gleason, H.A. and A.R. Cronquist. 1963. Manual 

of the Vascular Plants of Northeastern U.S. and 

Adjacent Canada. D. Van Nostrand Co., New 

York. 810 p. 

Great Plains Flora Association. 1986. Flora of the 

Great Plains. University Press of Kansas, KS. 

1,392 p. 

Harborne, J.B. 1988. Introduction to Ecological 

Biochemistry, 3rd Ed. Academic Press, London. 

356 p. 

Kartesz, J.T. 1994. A Synonymized Checklist of 

the Vascular Flora of the United States, Canada, 

and Greenland, Vol. 1, 2nd Edition. Biota of 

North America Program of the North Carolina 

Botanical Garden. Timber Press, Portland, OR. 

622 p. 

Kearney, T.H., R. Peebles, and Collaborators. 

1951 with 1960 supplement. Arizona Flora. 

University of California Press, Berkeley, CA. 

1,085 p. 

Martin, W.C. and C.E. Hutchins. 1980-1981 

(reprinted 2001). A Flora of New Mexico, Vol. 

1. Bishen Singh Mahendra Pal Singh (India) and 

Koeltz Scientific Books (Germany). 1,276 p. 

Mozingo, H.N. 1987. Shrubs of the Great Basin: 

A Natural History. University of Nevada Press, 

Reno, NV. 342 p. 

Newsholme, C. 1992. Willows: The Genus Salix. 

Timber Press, Portland, OR. 224 p. 

Ottenbreit, K.A. and R.J. Staniforth. 1992. Life 

cycle and age structure of ramets in an 

expanding population of Salix exigua (sandbar 

willow). Canadian Journal of Botany 70(6): 

1,141-1,146. 

Pomeroy, K.E., J.P. Shannon, and D.W. Blinn. 

2000. Leaf breakdown in a regulated desert 

river: Colorado River, Arizona, U.S.A. 

Hydrobiologia 434: 193-199. 

Rood, S.B. and J.M. Mahoney. 2000. Revised 

instream flow regulation enables cottonwood 

recruitment along the St. Mary River, Alberta, 

Canada. Rivers 7(2): 109-125. 

Stoehr, M.U., M. Cai, and L. Zsuffa. 1989. In-vitro 

plant regeneration via callus culture of mature 

Salix exigua. Canadian Journal of Forest 

Research 19(12): 1,634-1,638. 

Stubbendieck, J., S.L. Hatch, and C.H. Butterfield. 

1997. North American Range Plants, 5th Edition. 

University of Nebraska Press, Lincoln, NE. 

501 p. 

Taylor, J.P., D.B. Wester, and L.M. Smith. 1999. 

Soil disturbance, flood management, and 

riparian woody plant establishment in the Rio 

Grande floodplain. Wetlands 19(2): 372-382. 

Turick, C.E., M.W. Peck, D.P. Chynoweth, D.E. 

Jerger, E.H. White, L. Zsuffa, and W.A. 

Kenney. 1991. Methane fermentation of woody 

biomass. Bioresource Technology 37(2): 141-

147. 

U.S. Department of Agriculture Forest Service. 

1937 (1988 Dover edition). Range Plant 

Handbook. Dover Publications Inc., New York. 

816 p. 

Weber, W.A. and R.C. Wittman. 2001a. Colorado 

Flora: Eastern Slope. University Press of 

Colorado, Boulder CO. 521 p. 

Weber, W.A. and R.C. Wittman. 2001b. Colorado 

Flora: Western Slope. University Press of 

Colorado, Boulder CO. 488 p. 

Young, J.A. and C.G. Young. 1992. Seeds of 

Woody Plants in North America. Dioscorides 

Press, Portland, Oregon. 407 p. 
668 

Salix scouleriana Barratt ex Hook. Scouler’s willow 

SALICACEAE 

Synonyms: Salix brachystachys Benth. 

 Salix capreoides Anderss. 

 Salix flavescens Nutt. 

 Salix nuttallii Sarg. 

 Salix stagnalis Nutt. 

John K. Francis 

Illustration source: USDA-Forest Service Collection, 

Hunt Institute 

General Description.—Scouler’s willow, also 

known as fire willow, Nuttall willow, mountain 

willow, and black willow, is a deciduous shrub or 

small tree, depending on the environment. It 

usually has multiple stems that reach 2 to 7 m in 

height in dry, cold, high elevation, and other 

difficult environments, and 10 to 20 m in favorable 

sites. The stems are straight and support few 

branches generally resulting in narrow crowns. 

The root system is fibrous, deep, and widespread. 

The thick sapwood is nearly white, and heartwood 

is light brown tinged with red. Stem bark is thin, 

gray or dark brown with broad, flat ridges. Twigs 

are stout and whitish green. Leaves are 

oblanceolate to elliptic, 5 to 12.5 cm long, mostly 

short-pointed at the apex and tapered toward the 

base with entire to sparsely wavy-toothed margins. 

The leaves are dark-green and nearly hairless 

above and white- or grayish-hairy below. 

Scouler’s willow is dioecious, having male and 

female flowers on different trees. Tiny flowers are 

grouped in “pussy willow”-like catkins. The 

anthers, two per flower, are yellow, sometimes 

tipped with red; pistols are red. Fruits are light 

reddish-brown, long-pointed capsules about 0.75 

cm long. At maturity, they open to release a white 

fluff with imbedded tiny seeds. The species has 2n 

= 76 or 114 chromosomes (Anonymous 2003, 

Department of Ecology 2003, Harrington 1964, 

Peattie 1953, Sargent 1923, Viereck and Little 

1972). 

Range.—Scouler’s willow occurs from south 

central Alaska to central Manitoba, in the Black 

Hills of South Dakota, in the Rocky Mountains 

south to New Mexico and Arizona, and along the 

coast through British Colombia, Washington, 

Oregon, and California (Natural Resources 

Conservation Service 2003, Peattie 1953, 

Treeguide.com 2003). 

Ecology.—Scouler’s willow is the most common 

upland willow through most of its range. It invades 

quickly and abundantly after fires and logging 

(Viereck and Little 1972). Mineral soil seedbeds 

are required for seedling establishment (Forest 

Practices Branch 1997). In northern areas, 

Scouler’s willow occurs in muskegs, willow 

thickets, disturbed areas, and forests (Welsh 1974). 

At lower latitudes, the species grows in former 

clearcuts, burned areas, thinned forests, and areas 

of natural disturbance such as avalanche areas and 

river flood zones. These are all moist, well-drained 

to poorly drained sites. Although this willow 

tolerates drier conditions than most other willows, 

it does not tolerate xeric conditions. Scouler’s 

willow is a component in a large number of 

vegetation types throughout its range (Anderson 

2001). With few exceptions, it is the only willow 

found growing with other trees in upland Western 

forests (Johnson 1995). Soils of all textures, 

including skeletal soils and soils derived from 

most parent materials are colonized. Sites may 
669

vary from near sea level to about 3,000 m in 

elevation (Peattie 1953). Scouler’s willow is topkilled by all but gentle fires, but usually greater 

than 65 percent of the plants sprout quickly 

afterwards (Anderson 2001). The species is 

intolerant of shade, and when overtopped by 

conifers and other hardwoods, it begins to decline. 

Reproduction.—Scouler’s willow flowers from 

April through June, flowers appearing before 

leaves, often while snow is still on the ground, and 

fruiting occurs from May through July, depending 

on area. The flowers are insect pollinated. There 

are about 14,300 cleaned seeds/g. Germination, 

which is epigeal, begins to occur in 12 to 24 hours 

after seeds alight on wet ground. Germination 

usually reaches 95 percent in 1 or 2 days 

(Brinkman 1974). The seeds are dispersed by the 

wind. Plants sprout from the root collar when cut 

or top-killed. Pieces of stem and root will root and 

grow if partially buried in moist soil (Forest 

Practices Branch 1997). 

Growth and Management.—Annual height 

growth of sprouts from cut stems varies from 1 to 

3 m/year. Up to 60 sprouts are produced per stem 

(Forest Practices Branch 1997). Maximum height 

at 20 years is about 9 m. At higher elevations, 

shrubs reach 4 to 5 m in 15 years after which 

growth slows until a maximum height of 10 m is 

reached (Natural Resources Conservation Service 

2003). Fruits should be collected by hand or with 

pruning poles as soon as they turn from green to 

yellow. The capsules are air-dried until opening. 

Generally, the seeds should be sown as soon as 

possible because they remain viable for only a few 

days. Seed can be stored in sealed containers under 

refrigeration for 4 to 6 weeks, but germination 

begins to drop rapidly after 10 days. Seeds are 

broadcast on well-prepared beds that are kept 

continually moist until germination and seedling 

emergence. Light is required for successful 

germination (Brinkman 1974). Recommended 

spacing using rooted cuttings for erosion control is 

1.8 m by 1.8 m; for unrooted whips or shorter 

cuttings, 0.6 m. Rooted cuttings can be grown to 3 

m tall in containers. Cuttings should be 45 to 60 

cm long, and whips (not recommended) should be 

1.2 m long (Department of Ecology 2003). 

Benefits.—Scouler’s willow protects the soil and 

helps return sites to forest cover following 

disturbance. When growing along streams, it helps 

protect the stream banks from erosion and shade 

the watercourse, thus maintaining cooler water 

temperatures. It is an important browse species for 

domestic livestock and wild animals. Cattle, sheep, 

and goats all like it as browse. It is sometimes the 

most preferred food species for white tailed, black 

tailed and mule deer, elk, moose, and bighorn 

sheep. Small mammals, bears, upland game birds, 

and waterfowl feed to a lesser extent on leaves, 

buds, and seeds. Fresh browse (twigs and leaves) 

contain 41 percent dry matter, 4 percent protein, 2 

percent fat, 20.8 percent nitrogen-free extract, 11.2 

percent crude fiber, and good quantities of mineral 

nutrients (Anderson 2001). The cover provided by 

Scouler’s willow is important for mammals and 

birds. The flowers provide pollen and nectar to 

honey bees in early spring (Anderson 2001). The 

wood, which is soft and close-grained, is not sawn 

into lumber but is used to a limited extent for 

firewood and wood carving (Viereck and Little 

1972). The Secwepemc people of British 

Colombia used Scouler’s willow wood for 

smoking fish, drying meat, and constructing 

fishing weirs, the inner bark for lashing, sowing, 

cordage, and headbands, and decoctions of twigs 

for treating pimples, body odor, and diaper rash 

(Secwepemc Cultural and Education Society 

2003). 

References

Anderson, M.D. 2001. Salix scouleriana. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Missoula, MT. http://www.fs. 

fed.us/database/feis/plants/tree/salsco/all.html. 

57 p. 

Anonymous. 2003. Treatment from the Jepson 

manual: Salicaceae. University of California, 

Berkeley, CA. http://ucjeps.berkeley.edu/chibin/get_JM_treatment.pl?7043,7050,7079. 3 p. 

Brinkman, K.A. 1974. Salix L., willow. In: C.S. 

Schopmeyer, tech. coord. Seeds of woody plants 

in the United States. Agriculture Handbook 450. 

U.S. Department of Agriculture, Forest Service, 

Washington, DC. p. 746-750. 

Department of Ecology. 2003. Plant selection 

guide. State of Washington, Department of 

Ecology, Olympia, WA. http://www.ecy.wa. 

gov/programs/sea/pubs/93-30/table3.html. 6 p. 

Forest Practices Branch. 1997. Operational 

summary for vegetation management: willow 

complex. ISBN 0-7726-3166-2. Forest Practices 
670 

Branch, Ministry of Forests, Victoria, British 

Colombia, Canada. 11 p. 

Harrington, H.D. 1964. Manual of the plants of 

Colorado. Sage Books, Denver, CO. 666 p. 

Johnson, F.D. 1995. Wild trees of Idaho. 

University of Idaho Press, Moscow, ID. 212 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Salix scouleriana Barratt ex 

Hook. http://plants.usda.gov/chi_bin/ 

plant_search.cgi?mode=Scientific+Name&keyw

ordquery=Salix+scouleriana&earl=plant_search. 

4 p. 

Peattie, D.C. 1953. A natural history of western 

trees. Houghton Mifflin Company, Boston, MA. 

751 p. 

Sargent, C.S. 1923. Manual of the trees of North 

America (exclusive of Mexico). Houghton 

Mifflin, Boston, MA. 910 p. 

Secwepemc Cultural and Education Society. 2003. 

Flood plain garden. Secwepemc Nation, 

Kamloops, BC, Canada. http://www.secwepemc. 

org/SECethnogardens2/flood_plain_garden.htm. 

4 p. 

Treeguide.com. 2003. Scouler willow, Salix 

scouleriana Barratt ex Hook. http://www. 

treeguide.com/Species.asp?Region=NorthAmeri

can&SpeciesID=996. 2 p. 

Viereck, L.A. and E.L. Little, Jr. 1972. Alaska 

trees and shrubs. Agriculture Handbook 410. 

Forest Service, U.S. Department of Agriculture, 

Washington, DC. 265 p. 

Welsh, S.L. 1974. Anderson’s flora of Alaska and 

adjacent parts of Canada. Brigham Young 

University Press, Provo, UT. 724 p. 
 

671

Salvia apiana Jepson white sage

LAMIACEAE 

Synonyms: Audibertia polystachya Benth. 

 Ramona polystachya Briq. 

 Audibertiella polystachya Briq. 

 Salvia californica Jepson not Bdg. 

Arlee M. Montalvo

General Description.—White sage is a rounded, 1 

to 2.5 m tall shrub of low elevations that has 

highly aromatic, 3 to 9 cm long, whitish to pale 

gray-green leaves that persist throughout the year. 

The thick, petioled, lance-oblong leaves have 

crenulate margins and are covered with short 

appressed hairs and oil glands that give the leaves 

a silvery sheen. The attractive white to pale 

lavender flowers with short greenish calyx are 

clustered in compact cymose branches of a tall 

graceful, often pink-tinged inflorescence (thyrsoid 

panicle) that extends from 0.5 to 1.5 m above the 

foliage (Munz and Keck 1968). The lower lip of 

the two-lipped corolla is long and ruffled and 

obstructs the corolla tube, whereas the upper lip is 

reduced to a small lobe. Anthers release most 

pollen prior to maturity of stigmas. The two 

stamens are exerted to about 15 mm, diverging up 

and outward from their insertion on the lower lip. 

The exerted stigma protrudes forward about 15 

mm beyond the floral tube. There is a north to 

south and east to west geographic gradient in floral 

morphology (Meyn and Emboden 1987). For 

example, longer lips occur toward the coast. On 

the desert edge, there is a form with more compact 

inflorescences called S. apiana var. compacta

Munz (Munz and Keck 1968). Plants have n = 15 

chromosomes (Epling and others 1962). The four 

nutlets are about 2 to 3 mm long, somewhat 

rectangular, narrow in cross-section, keeled on one 

side, convex on the other side, and often greybrown to light brown.

Range.—White sage ranges from Santa Barbara 

County, California south to the middle of Baja 

California and east to the western edge of the 

Colorado desert and is an important component of 

inland and coastal sage scrub vegetation, with peak 

abundance in Riverside, Orange, and San Diego 

Counties, California. It is found on dry slopes and 

benches in many plant communities below 1500 m 

(occasionally higher), including chaparral, coastal 

sage scrub, yellow-pine forest, and the upper edges 

of desert scrub (Munz and Keck 1968). 

Ecology.—Plants often thrive in droughty soils 

and in habitats with high summer temperatures. 

Westman (1981a) reports occurrence on coarse to 

loamy soils derived from sandstone, conglomerate 

and sandstone, shale, granitic/dioritic rocks, and 

volcanic rocks. In western Riverside County, 

plants also occur on serpentine outcrops (author’s 

observation). White sage tends to occur with 

Artemisia californica Less. and Eriogonum 

fasciculatum Benth. Plants are notably drought 

tolerant even though roots are thought to be less 

than 1.5 m deep (Hellmers and others 1955). The 

crowded, whitish leaves are often held vertically 

and become folded during drought, likely 

adaptations for keeping leaves cool. In addition 

many leaves dehisce late in the dry season. In a 

desert wash in Baja California during summer 

drought, leaf conductance, transpiration rates, and 

turgor potentials were high, and plants did not 

show signs of drought stress (Schmitt and others 

1993). In chaparral, white sage had higher water 

potentials throughout the day well into the summer 

drought than did most of the evergreen shrubs 

measured (Poole and Miller 1975). Water 

potentials decreased late in the dry season and 

recovered after November at the end of the 

seasonal drought. The roots of white sage form 

associations with arbuscular mycorrhizal fungi that 

may also assist plants in low nutrient and droughty 
 

672 

habitats (personal communication, James Beaver, 

Indiana University, Bloomington). 

Fire is a normal disturbance factor of most 

plant communities within which white sage 

occurs. Following fire, this shrub frequently 

sprouts from distinct basal burls (Keeley 1998), 

but year-old seedlings are killed by fire, and 

repeated burning kills mature plants (Zedler and 

others 1983). Seedlings can emerge from the 

seedbank in the first season after fire, but reported 

densities are low (Zedler and others 1983, Keeley 

1998), and high intensity fires can kill seeds 

(Keeley and Fotheringham 1998). 

Reproduction.—Inflorescences arise from the 

upper nodes in spring and produce flowers from 

late May to July. Based on the floral morphology, 

only some of the larger bees are capable of 

transferring pollen to the highly exerted stigmas. 

Grant and Grant (1964) report three species of 

Xylocopa and a species of Bombus as effective 

pollinators. Hummingbirds, honeybees, bombyliid 

flies, and small species of native bees (primarily 

Anthophora, Diadasia, and Osmia) sometimes 

visit flowers (Grant and Grant 1964) but are 

thought to be ineffective as pollinators. The 

protandrous flowers and presence of interspecific 

hybrids are consistent with an outcrossing 

breeding system. Inflorescences are large and 

many-flowered, making some self-pollination 

likely. The small nutlets fall from the dried calyx 

during mid to late summer but are secondarily 

dispersed by harvester ants and seed caching 

rodents. Seed traps revealed that seeds dispersed 

up to 3 m away and that there was a higher 

secondary removal rate of seeds in coastal sage 

scrub (80 percent) than in grassland (0 percent) or 

the ecotone of the two habitats (about 40 percent) 

(DeSimone and Zedler 2001). 

Establishment and Growth.—Seedlings emerge 

in the cool wet season in late winter to early spring 

and can take 2 years or more to reach maturity, 

depending on rain patterns and growing 

conditions. Seedling establishment in shrublands is 

associated with gaps in mature vegetation and 

within grasslands in areas where herbs have been 

removed by gophers (DeSimone and Zedler 1999, 

2001). Seedlings show a relatively high growth 

rate, little herbivore damage and they fold their 

leaves in response to drought (DeSimone and 

Zedler 2001). In plots that were hand weeded and 

seeded in a twice-normal rainfall year, about a 

quarter of the seedlings survived to the second 

year (Storms 1999). Seedling survival is expected 

to be much lower in dry years and weedy areas. 

Most growth occurs after winter rains begin and 

continues until the soil becomes dry in summer. 

Leaf growth over the season results in a seasonal 

dimorphism in leaf size (Westman 1981b). Smaller 

leaves are produced on short shoots that develop 

from leaf axils along the main elongated stems and 

are often retained when the older, longer leaves 

dehisce during drought. With the return of 

seasonal rains, the short shoots elongate into long 

shoots, and the retained short leaves continue to 

expand and subtend the next generation of short 

shoots and leaves. 

Hybridization.—Epling (1938) reports that white 

sage hybridizes with other Salvias with the same 

chromosome number, including S. mellifera

Greene, S. munzii Epl., S. leucophylla Greene, S. 

clevelandii (Gray) Greene, S. eremostachya Jeps., 

S. pachyphylla Epl. Ex Munz., and S. vaseyi

(Porter) Parish. Hybrid zones between S. mellifera

(black sage) and white sage are especially 

common. Their hybrids are fully viable but, on 

average, suffer reduced fertility relative to parental 

species (Epling 1947a, Meyn and Emboden 1987), 

and there are fewer seeds/flower in F1 backcrosses 

to both parental species (Grant and Grant 1964). 

Hybrids exhibit a range of intermediate floral and 

leaf traits (Epling 1947a, b, Anderson and 

Anderson 1954) and differ in leaf anatomy (Webb 

and Carlquist 1964). Most hybrids are thought to 

be F1 crosses or backcrosses to black sage and 

they tend to grow close to the parental types. 

Although black and white sage overlap in range, 

they have different habitat affinities. Their ranges 

overlap in most of coastal southern California, but 

white sage is not found north of Santa Barbara 

County and it ranges farther south into Baja 

California and eastward into the edge of the desert 

than does black sage. Both species can occur 

intermingled, but black sage tends to be found in 

flatter and wetter areas, while white sage is usually 

on drier slopes (Epling 1947a, Anderson and 

Anderson 1954, Grant and Grant 1964, Gill and 

Hanlon 1998). As such, most sites contain a single 

species, but hybrids are present at most sites where 

the species co-occur (Meyn and Emboden 1987). 

Authors have speculated about factors that may 

limit the success of hybrids and maintain black and 

white sage as distinct species. Epling (1947a) first 

suggested that hybrids were less adapted to wet 

sites than black sage and less adapted to dry sites 

than white sage, but that they may be better 
 

673

adapted to recently disturbed conditions. Hybrids 

usually occur on recently disturbed areas, 

suggesting that hybrids are better adapted to 

intermediate habitats (Anderson and Anderson 

1954, Meyn and Emboden 1987). Within a natural 

hybrid zone, Gill and Hanlon (1998) found that 

xylem pressure potential was significantly higher 

in white than in black sage, that hybrids were 

intermediate to the two parental species, and that 

white sage drops fewer leaves during drought. 

These data support that white sage is more drought 

adapted than black sage. It is possible that 

selection will maintain the integrity of the hybrid 

zones if the intermediate hybrids do better in 

intermediate habitats relative to their performance 

in the main habitats of the parental species. 

The potential for reproductive isolation of the 

species within areas of overlap has been examined 

most thoroughly by Grant and Grant (1964), 

augmenting earlier observations (Epling 1947a, 

Anderson and Anderson 1954). Black sage 

generally flowers earlier than white sage and the 

short overlap in flowering time of one to several 

weeks limits hybridization. In addition, white sage 

has large, highly modified flowers that are 

pollinated almost exclusively by large carpenter 

and bumblebees (Xylocopa and Bombus), while 

black sage has smaller, unmodified flowers that 

are pollinated primarily by honeybees and small 

native solitary bees. 

Seed germination.—Keeley (1987) showed that 

exposure to 70 o

C for 1 hour followed by 

incubation in light substantially increased 

germination over controls, that heat shock for 5 

minutes at 100 o

C had no effect, and that exposure 

for 5 min at 120 o

C killed seeds. Leachate from 

charred wood did not increase germination 

(Keeley 1987), but cool smoke treatment increased 

germination (Keeley and Fotheringham 1998). 

Liquid smoke can also be used to significantly 

increase germination (author’s observation). 

Horticulture.—The generally rounded form with 

attractive whitish leaves and long graceful, pinktinged inflorescence stalks make this an attractive 

shrub for background areas of dry landscape 

gardens (Perry 1992, Keator 1994, Clebsch 1997). 

Plants require full sun, well-drained soil, do not 

tolerate constantly damp conditions, and are 

sensitive to conditions below –7 o

C (Keator 1994, 

Clebsch 1997), although populations in desert 

scrub are likely to be more cold tolerant than those 

from coastal locations. Plants should be kept dry 

all summer long. Clebsch (1997) recommends 

trimming back the dried inflorescence stalks to 

encourage compact growth. Plants can be easily 

started from seeds in the fall or planted from 

containers. 

Growth and Management.—Because of its 

historical abundance, fast growth, attractive nature, 

and importance to the biodiversity of sensitive 

sage scrub habitat, white sage is used in many 

restoration and roadside revegetation projects, 

especially in endangered sage scrub vegetation. 

Both container plants and seeds are used 

successfully in dry-land restoration in southern 

California. Seeds are collected from the dry, 

fruiting inflorescences, usually in July and August, 

depending on site, and processed to remove chaff 

and insects. S&S Seeds (Carpenteria, CA) and 

Mirov and Krabel (1939) report 715,000 to 

750,200 bulk seeds/kg, with often 70 percent 

purity and 50 percent germination. Seeds should 

be planted before the onset of the cool rainy 

season. If seed germination tests show low 

germination but high seed viability, seeds can be 

treated with dry or liquid smoke before sowing. If 

liquid smoke is used, seeds must be air-dried prior 

to dry broadcasting. Dormant seeds can result in a 

beneficial seed bank and establishment over 

several years. Hydroseeding can be successful on 

steep slopes and road cuts. Shallow planting 

methods such as hydroseeding and imprinting are 

likely to produce higher germination than drilling 

because seed germination is improved with 

exposure to light. Based on seedling survival data 

(Storms 1999, DeSimone and Zedler 2001), even 

under well-weeded and rainy conditions, fewer 

than 25 percent of seedlings should be expected to 

survive through their first year. This species has 

decreased dramatically in percent cover in western 

Riverside County since the early 1930’s (Minnich 

and Dezzani 1998). Successful restoration will 

depend on controlling the factors associated with 

its decrease, especially invasive plants and fire 

frequency, so that shrubs can establish, mature, 

and buildup a seed bank. 

Benefits.—White sage leaves are an important 

browse of mountain sheep in the winter and spring 

(Perry and others 1987). Leaves contain essential 

oils and a variety of diterpines and triterpenes, 

including carnosic acid, oleaolic acid, and ursolic 

acid, some of which have been linked to the 

medicinal use of white sage by native Tribes. 

Some may also deter many herbivores (Dentali 

and Hoffman 1992). A tincture of the leaves has 
 

674 

been used internally as a diaphoretic and diuretic, 

and externally to wash skin. The tincture may have 

antimicrobial properties. Antibacterial and 

antifungal activity of several compounds have 

been verified in vitro (Dentali and Hoffman 1992). 

In addition to medicinal use, the dried leaves have 

been used as a smudge and crushed as a soapless 

shampoo (Bean and Saubel 1972, Moore 1989). 

Seeds have been used as food and leaves for 

flavoring. 

References 

Anderson, E. and B.R. Anderson. 1954. 

Introgression of Salvia apiana and Salvia

mellifera. Annals of the Missouri Botanical 

Garden 41: 329-338. 

Bean, J.L. and K.S. Saubel. 1972. Temalpakh: 

Cahuilla Indian Knowledge and Usage of Plants. 

Malki Museum Press, Morongo Indian 

Reservation, CA. 225 p. 

Clebsch, B. 1997. A Book of Salvias: Sages for 

Every Garden. Timber Press Inc., Portland, OR. 

221 p. 

Dentali, S.J. and J.J. Hoffmann. 1992. Potential 

antiinfective agents from Eriodictyon

angustifolium and Salvia apiana. International 

Journal of Pharmacognosy 30: 223-231. 

DeSimone, S.A. and P.H. Zedler. 1999. Shrub 

seedling recruitment in unburned Californian 

coastal sage scrub and adjacent grassland. 

Ecology 80: 2,018-2,032. 

DeSimone, S.A. and P.H. Zedler. 2001. Do shrub 

colonizers of southern Californian grassland fit 

generalities for other woody colonizers? 

Ecological Applications 11: 1,101-1,111. 

Epling, C. 1938. The California salvias. Annals of 

the Missouri Botanical Garden 25: 95-188. 

Epling, C. 1947a. The genetic aspects of natural 

populations: actual and potential gene flow in 

natural populations. The American Naturalist 81: 

104-113. 

Epling, C. 1947b. Natural hybridization of Salvia

apiana and S. mellifera. Evolution 1: 69-78. 

Epling, C., H. Lewis, and P.H. Raven. 1962. 

Chromosomes of Salvia: section Audibertia. 

Aliso 5: 217-221. 

Gill, D.S. and B.J. Hanlon. 1998. Water potentials 

of Salvia apiana, S. mellifera (Lamiaceae), and 

their hybrids in the coastal sage scrub of southern 

California. Madroño 45: 141-145. 

Grant, K.A. and V. Grant. 1964. Mechanical 

isolation of Salvia apiana and Salvia mellifera

(Labiatae). Evolution 18: 196-212. 

Hellmers, H., J.S. Horton, G. Juhren, and J. 

O'Keefe. 1955. Root systems of some chaparral 

plants in southern California. Ecology 36: 667-

678. 

Keator, G. 1994. Complete Garden Guide to the 

Native Shrubs of California. Chronicle Books, 

San Francisco, CA. 314 p. 

Keeley, J.E. 1987. Role of fire in seed germination 

of woody taxa in California chaparral. Ecology 

68: 434-443. 

Keeley, J.E. 1998. Postfire ecosystem recovery and 

management: The October 1993 large fire 

episode in California. In: J. M. Moreno (ed.) 

Large Forest Fires. Backbuys Publishers, Leiden, 

Netherlands. p. 69-90. 

Keeley, J.E., and C.J. Fotheringham. 1998. Smokeinduced seed germination in California 

chaparral. Ecology 79: 2,320-2,336. 

Meyn, O. and W.A. Emboden. 1987. Parameters 

and consequences of introgression in Salvia

apiana X S. mellifera (Lamiaceae). Systematic 

Botany 12: 390-399. 

Minnich, R.A. and R.J. Dezzani. 1998. Historical 

decline of coastal sage scrub in the RiversidePerris plane, California. Western Birds 29: 366-

391. 

Mirov, N.T. and C. J. Kraebel. 1939. Collecting and 

handling seeds of wild plants. Civilian 

Conservation Corps, Forestry Publication No. 5, 

United States Government Printing Office, 

Washington, D.C. 42 p. 
 

675

Moore, M. 1989. Medicinal Plants of the Desert 

and Canyon West. Museum of New Mexico 

Press, Santa Fe, NM. 184 p. 

Munz, P.A. and D.D. Keck. 1968. A California 

Flora with Supplement. University of California 

Press, Berkeley, CA. 1681 + 224 p. 

Perry, B. 1992. Landscape Plants for Western 

Regions: an Illustrated Guide to Plants for Water 

Conservation. Land Design Publishing, 

Claremont, CA. 318 p. 

Perry, W.M., J.W. Dole, and S.A. Holl. 1987. 

Analysis of the diets of mountain sheep from the 

San Gabriel Mountains, California. California 

Fish and Game 73: 156-162. 

Poole, D.K. and P.C. Miller. 1975. Water relations 

of selected species of chaparral and coastal sage 

communities. Ecology 56: 1,118-1,128. 

Schmitt, A.K., C.E. Martin, V.S. Loeschen, and A. 

Schmitt. 1993. Mid-summer gas exchange and 

water relations of seven C3 species in a desert 

wash in Baja California, Mexico. Journal of Arid 

Environments 24: 155-164. 

Storms, N. 1999. Restoration of a Native Shrubland 

in an Area of Frequent Disturbance and High 

Nitrogen Deposition. M.S. Thesis. University of 

California, Riverside. 60 p. 

Webb, A.-A. and S. Carlquist. 1964. Leaf anatomy 

as an indicator of Salvia apiana-mellifera

introgression. Aliso 5: 437-449. 

Westman, W.E. 1981a. Factors influencing the 

distribution of species of California coastal sage 

scrub. Ecology 62: 439-455. 

Westman, W.E. 1981b. Seasonal dimorphism of 

foliage in Californian coastal sage scrub. 

Oecologia 51: 385-388. 

Zedler, P.H., C.R. Gautier, and G.S. McMaster. 

1983. Vegetation change in response to extreme 

events: the effect of a short interval between fires 

in California chaparral and coastal scrub. 

Ecology 64: 809-818. 
 

676 

Salvia mellifera Greene black sage 

LAMIACEAE 

Synonyms: Audibertia stachyoides Benth. 

 Ramona stachyoides Briq. 

 Audibertiella stachyoides Briq. 

Arlee M. Montalvo. and Paul A. McMillan 

General Description.—Black sage has a dark 

appearance, especially during drought. Salvia is 

derived from the Latin word for safe and refers to 

the medicinal properties of sages. The specific 

epithet “mellifera” means “honey producing” and 

refers to its use by nectar foraging bees. Shrubs are 

open, 1 to 2 m tall with twisted, furrowed woody 

stems at the base, and herbaceous greenish branch 

tips that are square in cross section. Twigs and 

leaves are glandular, highly aromatic, and 

opposite. The oblong-elliptical blades are 2 to 6 

cm long, dark green and rugulose above (convex 

between obvious veins) with crenulate (finely 

scalloped) margins, and taper to a few to 12 mm 

long petiole (Munz and Keck 1968). Many small, 

6- to 12-mm long, two-lipped flowers occur in 

compact clusters subtended by green bracts and are 

spaced 2 to 6 cm apart along the often branched 

inflorescence. Corollas vary from pure white to 

pale blue or lavender. The style and stamens 

extend just beyond the corolla tube. Flowers bear 

four oblong nutlets (single–seeded fruits) within a 

persistent calyx. Nutlets vary from mottled gray to 

dark brown and are about 1 mm wide and 2 mm 

long. 

Range.—Black sage occurs on dry slopes and 

alluvial fans within interior and coastal sage scrub 

and lower montane chaparral from sea level to 

1,200 m, ranging from Contra Costa County, CA 

(near San Francisco) south to northern Baja 

California (Munz and Keck 1968, Hickman 1993). 

On leeward slopes of the Coast Ranges it extends 

into desert scrub (Keeley 1986). It is one of the 

dominant shrubs of sage scrub (Westman 1983), 

and can form dense, nearly monospecific stands on 

steep slopes. 

Ecology.—Black sage occurs on a variety of soils 

derived from sandstone, shale, granite, and 

especially serpentinite, and gabbro basalt 

(Westman 1981). Plants produce numerous 

volatile compounds, including camphor, cineole, 

terpenes, and sesquiterpenes (Neisess and others 

1987, Gonzalez and others 1992, Arey and others 

1995), which may be important in herbivore or 

pathogen defense. 

Black sage is semideciduous, depending on 

site and severity of drought, and shallow rooted. 

Plants are drought tolerant by leaf curling rather 

than drought avoiding through leaf drop (Gill and 

Hanlon 1998). Leaf drop is drought induced 

possibly due to embolisms that occur in xylem 

tissues (Kolb and Davis 1994). At Stanford, CA, 

leaves began to senesce and drop in October in 

both irrigated and unirrigated treatments, leaf 

stomata did not control transpiration when xylem 

water potentials were low, and rates of 

photosynthesis did not decline until after leaf drop 

(Gigon 1979). In Santa Barbara County, CA, most 

leaves senesced between May and July, 32 percent 

persisted into the following growing season, leaf 

curling and uncurling was related to water stress, 

and leaf drop was more related to photoperiod 

than to low xylem potentials (Gill and Mahal 

1986). 
 

677

Reproduction.—The timing of growth and 

flowering is closely adapted to the Mediterranean 

climate of southern California, with cool wet 

winters and hot dry summers (Grant and Grant 

1964, Gill and Mahall 1986). Common garden 

studies with 12 populations revealed that date of 

first flowering varies with source population and 

ranged from early February to early May 

(Montalvo, unpublished data). Also, shifts in 

flowering time occur in years with contrasting 

rainfall patterns (Meyn and Emboden 1987). 

The self-compatible flowers are 

hermaphroditic with stamens dispersing pollen 

before stigmas are receptive. Flowers are 

pollinated by small to meduim-sized solitary bees 

in families Andrenidae, Anthophoridae, 

Halictidae, Megachilidae, and Xylocopinae as well 

as introduced honey bees (Grant and Grant 1964). 

Rarer visitors include large-bodied Bombus and 

Xylocopa bees, syrphid and bombyliid flies, and 

Anna’s hummingbirds. Each flower produces up to 

four seeds that are gravity dispersed in June and 

July from the dry, persistent calyces, and can be 

secondarily dispersed by ants. Seeds accumulate in 

a dormant seed bank (Keeley 1986). 

Establishment and Growth.—Black sage occurs 

in plant communities adapted to fire and drought. 

Seedlings emerge in clearings between adult 

shrubs, but most emerge in the first 2 years 

following fire, primarily from February to April 

(Keeley 1986, Westman and O’Leary 1986). 

Plants take two or more growing seasons to reach 

maturity. After fire, plants sometimes resprout 

from the base (Went and others 1952, Keeley 

1986, 1998). However, where fire intensity is high, 

shrubs tend to be killed, and recruitment is 

primarily by seed. Across a range of conditions, 

resprouting success averaged 15.8 percent, and 

shrubs that resprouted had significantly smaller 

basal diameters, suggesting resprouting decreases 

with age (Keeley 1998). The seasonal pattern of 

growth and dormancy varies both geographically 

and between years. Generally, seasonal growth 

begins after the start of fall rains, most new leaves 

are produced by mid March, and flowering occurs 

from late winter to late spring. 

Roots are colonized by arbuscular 

mycorrhizal (AM) fungi. After 4 months of growth 

in a greenhouse, root and shoot mass was higher in 

seedlings inoculated with AM fungi than in 

controls, but in younger plants the opposite 

occurred (E. Allen, L. Egerton-Warburton, A. 

Montalvo, unpublished report). 

Seed germination.—Seeds of black sage often 

have low germination rates unless exposed to light 

or components of fire such as charred wood, 

smoke, or KNO3 (Keeley 1986, Thanos and 

Rundel 1995, Keeley and Fotheringham 1998). 

Exposure to dry cool smoke for 5 min (Keeley and 

Fotheringham 1998) yields higher germination 

than charred wood. A 12 to 15 hour soak in a 1:25 

dilution of Regen 2000 Smokemaster, a liquid 

smoke product, also breaks dormancy (author’s 

observation). Heat of fire may reduce germination 

of seeds from sage scrub populations (Keeley and 

Fotheringham 1998). Oddly, seeds from desert 

populations, where fires are less frequent, were 

stimulated to germinate by heat and charred wood, 

but not by light (Keeley 1986). 

Genetics, Geographic Variation, and Fitness.—

Black sage is genetically variable over its 

geographic range. In a survey of 12 populations 

(Montalvo, Clegg, and Ellstrand unpublished, 14 

allozyme loci), expected heterozygosity was high 

(He = 0.23) and alleles averaged 2.81 per locus. 

Within sites, a low inbreeding coefficient (f = 

0.072) suggests the protandrous flowers and 

pollinators combine to promote outcrossing. There 

is significant but low structure both among and 

within populations, and results are consistent with 

a relatively high level of historical gene flow 

(Theta-p (~FST, sites relative to total population) = 

0.041; Theta-s (plots relative to sites) = 0.057; and 

F (FIT) = 0.125). 

Populations cluster by genetic similarity, and 

geographic floristic associations and this is 

somewhat mirrored by distribution patterns of 

morphological variation and timing of flowering. 

Flowers from populations in the Santa Monica 

Mountains and Simi Hills can be strikingly bluish. 

In the drier, more interior hills and valleys of 

Riverside Co., flowers tend to be white to pale 

blue or lavender, and plants flower earlier. These 

differences are retained in common gardens. 

Differences in timing of flowering could affect 

seed production of translocated populations if 

pollinator activity, seed predation, or important 

physical environmental factors differ from home 

sites. Plants from contrasting populations also 

smell different, likely due to differences in 

composition of volatile chemicals. Such 

differences among populations may be adaptive 
 

678 

and contribute to their success under particular 

environments. Montalvo and Ellstrand 

(unpublished) evaluated seedling survival, growth 

and reproduction of 12 populations in a common 

garden. A combination of climatic data and soil 

traits was used to calculate environmental 

similarity among garden and source sites. After 

three growing seasons, populations from sources 

most similar to the garden site were the most 

successful (measured as survival x flower 

production). This effect was small (r2

 = 0.18) but 

is consistent with the presence of weak adaptive 

differences among source populations. 

Hybridization.—Black sage hybridizes with the 

shrubs S. apiana Jeps. (n = 15), S. leucophylla

Greene (2n = 30), and S. clevelandii (Gray) Greene

(2n = 30), and rarely with annuals S. columbariae 

Benth (n = 13) and S. carduacea Benth (n = 16)

(Epling 1938, 1947a, Epling and others 1962, 

Munz and Keck 1968). Most hybridization is with 

S. apiana. These hybrids are fully viable but suffer 

reduced pollen fertility (Epling 1947a, Meyn and 

Emboden 1987) and numbers of seeds/flower in 

F1 back crosses to parental species (Grant and 

Grant 1964). Hybrids differ from both parental 

species by a range of intermediate floral, leaf, and 

anatomical traits (Epling 1947a, b, Anderson and 

Anderson 1954, Webb and Carlquist 1964). Most 

are thought to be F1s or backcrosses to S. mellifera

and tend to grow near parental types. Hybrid zones 

can be stable in position but may shift in 

morphology (Meyn and Emboden 1985). This 

suggests differences in the relative success of 

hybrids and backcrossed progeny over time. The 

influence of gene exchange on floral form of black 

sage does not appear to extend beyond hybrid 

zones (Epling 1947a, b). 

Several authors have speculated on the 

factors promoting hybridization while also 

maintaining black and white sage as distinct 

species. While ranges overlap in most of southern 

California, S. apiana is not found north of Santa 

Barbara County and it ranges farther south into 

Baja California and eastward. In areas of contact, 

S. mellifera tends to occur in flatter and wetter 

microsites, while S. apiana occurs on drier slopes 

(Epling 1947a, Anderson and Anderson 1954, 

Grant and Grant 1964, Meyn and Emboden 1987, 

Gill and Hanlon 1998). Hybrids also occur in 

recently disturbed areas, suggesting that hybrids 

are adapted to intermediate habitats (Anderson and 

Anderson 1954, Meyn and Emboden 1987). 

Within a hybrid zone, Gill and Hanlon (1998) 

found that xylem pressure potential was 

significantly higher in white than in black sage and 

that putative hybrids were intermediate to the two 

parental species. These data support that S. apiana

is more drought adapted than S. mellifera, and that 

the hybrids are intermediate. In addition, 

reproductive isolation is favored by flowering 

differences. S. apiana flowers later than black 

sage, limiting opportunities to hybridize (Grant 

and Grant 1964). In addition, white sage has 

larger, highly modified flowers that are pollinated 

almost exclusively by large bees in Xylocopa and 

Bombus. Black sage has smaller, two-lipped 

flowers that are rarely visited by these largebodied bees. 

Growth and Management.—Seeds are collected 

from June-August and should be cleaned to 

remove seed predators. Air separation can be used 

to remove lighter unfilled seeds that are sometimes 

abundant. There are about 1,375,000 seeds/bulk kg 

(personal communication with Victor Schaff, S&S 

Seeds, Carpenteria, CA). Treatment of seeds with 

dry or liquid smoke increases germination 

substantially. Seeds should be air-dried before 

handling. Plants can be sown in flats for 

subsequent transplanting or directly sown into the 

ground in the fall. Plants can be started from 

cuttings, but this should be avoided for restoration 

because it reduces genetic variation. Prostrate 

cultivars (e.g., Clebsch 1997) should not be used 

in restoration. Black sage grows in well-drained 

soil and can tolerate some summer water in a 

garden. 

Benefits.—Black sage is important for restoration, 

erosion control, and native landscape gardening in 

California. Plants provide cover and seeds for 

wildlife, and in western Riverside Co. the listed 

California gnatcather frequently nests in 

vegetation containing black sage (Weaver 1998). 

The seeds were eaten and the leaves used for 

flavoring by native California tribes (Bean and 

Saubel 1972). In addition, the leaves have a long 

history of medicinal use by native tribes (Bocek 

1984). Their diterpene compounds have 

antimicrobial activity against gram (+) bacteria 

(Moujir and others 1996). 

Decreasing Populations.—Black sage 

populations are declining. Increased ignition 

sources and invasion of shrublands by annual 

grasses have increased fire frequency, jeopardizing 
 

679

recovery of seed banks (Westman and O’Leary 

1986, Haidinger and Keeley 1993, Minnich and 

Dezzani 1998). Black sage is also inhibited by 

exotic annual grasses, black mustard (Went and 

others 1952, Storms 1999), and air pollution 

(Westman 1985, Preston 1988). 

References 

Anderson, E. and B. R. Anderson. 1954. 

Introgression of Salvia apiana and Salvia

mellifera. Annals of the Missouri Botanical 

Garden 41: 329-338. 

Arey, J., D.E. Crowley, M. Crowley, M. Resketo, 

and J. Lester. 1995. Hydrocarbon emissions 

from natural vegetation in California's south 

coast air basin. Atmospheric Environment 29: 

2,977-2,988. 

Bean, J. and K. Saubel. 1972. Temalpakh: Cahuilla 

Indian knowledge and usage of plants. Malki 

Museum Press, Morongo Indian Reservation, 

CA. 225 p. 

Bocek, B. 1984. Ethnobotany of Costanoan 

Indians, California, based on collections by 

John P. Harrington. Economic Botany 38: 240-

255. 

Clebsch, B. 1997. A Book of Salvias: Sages for 

Every Garden. Timber Press, Portland, OR. 221 

p. 

Epling, C. 1938. The California salvias. Annals of 

the Missouri Botanical Garden 25: 95-188. 

Epling, C. 1947a. The genetic aspects of natural 

populations: actual and potential gene flow in 

natural populations. The American Naturalist 

81: 104-113. 

Epling, C. 1947b. Natural hybridization of Salvia

apiana and S. mellifera. Evolution 1: 69-78. 

Epling, C.,H. Lewis, and P.H. Raven. 1962. 

Chromosomes of Salvia: section Audibertia. 

Aliso 5: 217-221. 

Gigon, A. 1979. CO2-gas exchange, water 

relations and convergence of Mediterranean 

shrub-types from California and Chile. 

Oecologia Plantarum 14: 129-150. 

Gill, D.S. and B.J. Hanlon. 1998. Water potentials 

of Salvia apiana, S. mellifera (Lamiaceae), and 

their hybrids in the coastal sage scrub of 

southern California. Madroño 45: 141-145. 

Gill, D.S. and B.E. Mahall. 1986. Quantitative 

phenology and water relations of an evergreen 

and deciduous chaparral shrub. Ecological 

Monographs 56: 127-143. 

González, A.G., L.S. Andrés, Z.E. Aguiar, and 

J.G. Luis. 1992. Diterpenes from Salvia

mellifera and their biogenetic significance. 

Phytochemistry 31(4): 1,297-1,305. 

Grant, K.A. and V. Grant. 1964. Mechanical 

isolation of Salvia apiana and Salvia mellifera

(Labiatae). Evolution 18: 196-212. 

Haidinger, T.L. and J.E. Keeley. 1993. Role of 

high fire frequency in destruction of mixed 

chaparral. Madroño 40: 141-147. 

Hickman, J.C., editor. 1993. The Jepson Manual: 

Higher Plants of California. University of 

California Press, Ltd., Los Angeles, CA. 1,400 

p. 

Keeley, J.E. 1986. Seed germination patterns of 

Salvia mellifera in fire-prone environments. 

Oecologia 71: 1-5. 

Keeley, J.E. 1998. Postfire ecosystem recovery 

and management: The October 1993 large fire 

episode in California. In: J. M. Moreno, ed. 

Large Forest Fires. Backbuys, Leiden, The 

Netherlands. p. 69-90. 

Keeley, J.E. and C.J. Fotheringham. 1998. Smokeinduced seed germination in California 

chaparral. Ecology 79: 2,320-2,336. 

Kolb, K.J. and S.D. Davis. 1994. Drought 

tolerance and xylem embolism in co-occurring 

species of coastal sage and chaparral. Ecology 

75: 648-659. 

Meyn, O. and W.A. Emboden. 1987. Parameters 

and consequences of introgression in Salvia

apiana X S. mellifera (Lamiaceae). Systematic 

Botany 12: 390-399. 

Minnich, R.A. and R.J. Dezzani. 1998. Historical 

decline of coastal sage scrub in the RiversidePerris plane, California. Western Birds 29: 366-

391. 
 

680 

Moujir, L., A.M. Gutiérrez-Navarro, L. San 

Andres, and J.G. Luis. 1996. Bioactive 

diterpenoids isolated from Salvia mellifera. 

Phytotherapy Research 10: 172-174. 

Munz, P.A. and D.D. Keck. 1968. A California 

Flora with Supplement. University of California 

Press, Berkeley, CA. 1,681 + 224 p. 

Neisess, K.R., R.W. Scora, and J. Kumamoto. 

1987. Volatile leaf oils of California Salvias. 

Journal of Natural Products 50: 515-517. 

Preston, K.P. 1988. Effects of sulphur dioxide 

pollution on a Californian coastal sage scrub 

community. Environmental Pollution 51: 179-

195. 

Storms, N. 1999. Restoration of a native shrubland 

in an area of frequent disturbance and high 

nitrogen deposition. Master of Science in 

Botany Thesis. University of California, 

Riverside. 60 p. 

Thanos, C.A. and P.W. Rundel. 1995. Firefollowers in chaparral: nitrogenous compounds 

trigger seed germination. Journal of Ecology 

83: 207-216. 

Weaver, K.L. 1998. Coastal sage scrub variations 

of San Diego County and their influence on the 

distribution of the California gnatcatcher. 

Western Birds 29: 392-405. 

Webb, A.-A. and S. Carlquist. 1964. Leaf anatomy 

as an indicator of Salvia apiana-mellifera

introgression. Aliso 5: 437-449. 

Went, F.W., G. Juhren, and M.C. Juhren. 1952. 

Fire and biotic factors affecting germination. 

Ecology 33: 351-364. 

Westman, W.E. 1981. Factors influencing the 

distribution of species of California coastal sage 

scrub. Ecology 62: 439-455. 

Westman, W.E. 1983. Xeric Mediterranean-type 

shrubland associations of Alta and Baja 

California and the community/continuum 

debate. Vegetatio 52: 3-19. 

Westman, W.E. 1985. Air pollution injury to 

coastal sage scrub in the Santa Monica 

Mountains, southern California. Water, Air, and 

Soil Pollution 26: 19-41. 

Westman, W.E. and J.F. O'Leary. 1986. Measures 

of resilience: the response of coastal sage scrub 

to fire. Vegetatio 65: 179-189. 
 

681

Sassafras albidum (Nutt.) Nees sassafras 

LAURACEAE 

Synonyms: Sassafras sassafras (L.) Karst. 

 Laurus albidus Nutt. 

 Laurus sassafras L. 

 Sassafras officinale Nees & Eberm. 

 Sassafras variifolium (Salsb.) Kuntze 

 Sassafras albidum (Nutt.) Nees var. molle (Raf.) Fern. 

John K. Francis 

General Description.—Sassafras is also known as 

white sassafras, common sassafras, ague tree, 

cinnamon wood, saloop, smelling stick, gumbo 

file, and mitten tree. It is a shrub or small tree up 

to 15 m in height and 35 cm in diameter. The 

species grows as a medium to large shrub in 

Florida, in the northern part of its range, and on 

sand or poor, rocky sites. It may have single or 

multiple stems. Older stems have deeply furrowed, 

red-brown bark. Slender, brittle, twigs with thick, 

white, mucilaginous pith are covered by green 

bark. The species depends primarily on a shallow, 

lateral root system. Long lateral roots develop 

suckers that in turn develop their own root system. 

The alternate dark green leaves occur in three 

forms: ovate or elliptical leaves, leaves with one 

side lobe (“mitten”), and trilobate leaves with the 

center lobe being symmetrical and somewhat 

larger than the side lobes. Leaves are 8 to 15 cm 

long. Small yellowish-green flowers grow in 

axillary racemes. The fruit is a fleshy, oblong, dark 

blue or black drupe partially set in a red or orange 

thickened base (pedicel and calyx). The seeds are 

oblong, pointed, and brown (Griggs 1990, Sargent 

1923, Vines 1982). 

Range.—Sassafras is native to all of the States 

east of a line running between Lake Superior and 

western Texas, and in extreme southern Ontario 

(Natural Resources Conservation Service 2003, 

Sullivan 1993). It is planted as an ornamental in at 

least the United Kingdom, Germany, and Spain 

(Oberle 2003, Plants for a Future 2003, Sanchez de 

Lorenzo 2003). There are no reports of it 

naturalizing outside its native range. The genus 

contains two other species, natives of China and 

Taiwan (Center for Wood Anatomy Research 

2003) 

Ecology.—Sassafras is intolerant of shade, and 

reproduction from seed is rare under closedcanopy forest. However, it is a pioneer in old 

fields and an invader of disturbed and early 

secondary forests. It grows as scattered individual 

plants or as dense thickets, often of sucker origin. 

Removal of the forest overstory by almost any 

means (i.e. gypsy moth, logging, fire) results in 

increased density of the species. On the other 

hand, as competition increases and forest canopies 

close, sassafras decreases in importance. Although 

uncommon, it persists in old-growth stands. The 

species grows from a few meters above sea level 

to 1,220 m (Sullivan 1993). Mean annual 

precipitation ranges from 760 to 1400 mm/year 

and the average frost-free period ranges from 160 

to 300 days. Average January temperatures range 

from -7 to 13 °C and average July temperatures 
 

682 

range from 21 to 27 °C. (Griggs 1990). Sassafras 

grows in a wide variety of soil types and grows 

best on well-drained sandy loams with a pH 

between 6.0 and 7.0 (Griggs 1990). Seedlings and 

saplings are easily top-killed by fire but readily 

sprout. Thick-barked trees are injured by fire 

(Sullivan 1993). Sassafras is sometimes attacked 

by leaf blight caused by Actinopelte dryina, by leaf 

spots caused by Mycosphaerella sassafras, and by 

a Nectria canker. A number of insects also attack 

the species (Griggs 1990). 

Reproduction.—Flowers open in early spring 

with the first leaves. Male and female flowers 

usually occur on different plants (Sargent 1923). 

Fruits mature in August or September. Seed 

production begins when plants are about 10 years 

old (Griggs 1990). There are about 12,800 

seeds/kg (Bonner and Maisenhelder 1974) or 

according to Griggs (1990), 8,800 to 13,200 

seeds/kg. About 35 percent of the seeds are sound. 

Birds are the principal dispersers of seeds, with 

gravity, water, and mammals playing minor roles. 

Germination, which occurs in the spring, is 

hypogeal. Seeds may remain viable in the soil seed 

bank for up to 6 years (Griggs 1990). Sassafras 

sprouts prolifically from stumps and suckers from 

lateral roots. 

Growth and Management.—Growth of sassafras 

is moderately fast. Sprouts can reach 1.2 m in the 

first year and 4.5 m in 4 years (Floridata 2003). 

Rare individual trees reaching the forest canopy 

may live as long as 300 years (TreeGuide 2003). 

Fruits are ripe and can be harvested when they turn 

from green to dark blue and can be picked by hand 

or by flailing them from the trees or shrubs onto a 

tarp. The seeds can be cleaned of fruit tissue by 

rubbing against hardware cloth and washing away 

the pulp. One kg of fruits will yield about 310 g of 

seeds. Seeds should be placed in sealed containers 

and stored at 2 to 5 °C. Before planting, seeds 

should be moistened and held at the above 

temperatures for 120 days. Drilling in nursery beds 

in 20- to 30-cm-wide rows and covering with 6 to 

12 mm of soil is recommended (Bonner and 

Maisenhelder 1974). Sassafras can also be 

reproduced from root cuttings but not from stem 

cuttings (Griggs 1990). Wildlings are difficult to 

transplant. Sassafras shrubs and trees can be killed 

by injecting 2,4-D, picloram, or glyphosate 

(Sullivan 1993). 

Benefits.—Sassafras adds beauty in both summer 

and winter to forest lands where it grows. It has 

been used to help restore depleted soils of old 

fields and is considered superior to black locust 

(Robinia pseudoacacia L.) and pines in this 

application (Grigg 1990). Sassafras is browsed 

year-round by white tailed deer. The plants are 

also utilized to some extent by woodchucks, 

rabbits, beavers, and bears. Many species of birds 

and some small mammals consume the fruits. 

Crude protein ranges from 21.0 percent (leaves in 

April) to 6.1 percent (twigs in January). The fruits 

have a high lipid and energy value (Sullivan 

1993). Sassafras makes an attractive ornamental 

small tree and is widely, although not heavily 

used. The fall foliage colors of yellow, orange, and 

red are particularly beautiful. Sassafras heartwood 

is pale brown to orange brown; sapwood is 

yellowish-white. The wood is ring-porous, coarsebut straight-grained, brittle, soft, and has a spicy 

aromatic odor. It is resistant to decay making it 

useful for fence posts and other items exposed to 

moisture. It is also useful for lumber, millwork, 

furniture, and small boats. Technical details of the 

wood have been published (Center for Wood 

Anatomy Research 2003). The use of sassafras in 

herbal medicine has a long history beginning with 

Native Americans who used it to treat colds, high 

blood pressure, heart troubles, swelling, as a tonic, 

to treat for worms, and to control fever (Moerman 

1986). Sassafras tea is still widely drunk both for 

pleasure and as a tonic. Extracts and infusions of 

the plant are used to treat a wide range of physical 

complaints. Dry, powdered leaves, called file, are 

used to thicken gumbo and other Cajun dishes 

(Katzer 1999). Oil of sassafras, extracted from the 

root bark, was the original flavoring for root beer 

and was used to flavor candy, chewing gum, and 

medicines. Because it contains safrole, a 

carcinogen, it is no longer permitted in food, but 

may be used to add scent to cosmetics and soap 

(TreeGuide 2003). 

References

Bonner, F.T. and L.C. Maisenhelder. 1974. 

Sassafrass albidum (Nutt.) Nees sassafras. In: 

C.S. Schopmeyer, tech. coord. Seeds of woody 

plants in the United States. Agriculture 

Handbook 450. U.S. Department of Agriculture, 

Forest Service, Washington, DC. p. 761-762. 

Center for Wood Anatomy Research. 2003. 

Sassafras albidum. Technical Transsfer Fact 
 

683

Sheet. U.S. Department of Agriculture, Forest 

Service, Forest Products Laboratory, Madison, 

Wisconsin. http://www2fpl.fs.fed.us/ 

TechSheets/HardwoodNA/htmlDocs/sassafrasal

bi.html. 3 p. 

Floridata. 2003. Sassafras albidum. http:// 

www.floridata.com/ref/s/sass_alb.cfm. 4 p. 

Griggs, M.M. 1990. Sassafras albidum (Nutt.) 

Nees, sassafras. In: R.M. Burns, and B.H. 

Honkala. Silvics of North America. Vol. 2., 

Hardwoods. Ag. Handb. 654. U.S. Department 

of Agriculture, Forest Service, Washington, DC. 

p. 773-777. 

Katzer, G. 1999. Sassafras [Sassafras albidum

(Nutt.) Nees]. http://www-ang.kfunigraz. 

ac.at/~katzer/engl/Sass_alb.html. 4 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports 19. University of 

Michigan Museum of Anthropology, Ann 

Arbor, MI. 534 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Sassafras albidum (Nutt.) Nees. 

http://plants.usda.gov/cgi_bin/plant_profile.cgi?

symbol=SAAL5. 6 p. 

Oberle, R. 2003. Sassafras albidum (Nutt.) Nees. 

http://www.biologie.uni-ulm.de/systax/ 

dendrologie/sasalblv.htm. 1 p. 

Plants For a Future. 2003. Plant portrait-Sassafras 

albidum. http://www.scs.leeds.ac.uk/pfaf/ 

sassafr.html. 3 p. 

Sanchez de Lorenzo-C., J.M. 2003. Árboles 

ornamentals: Sassafras albidum (Nutt.) Nees. 

http://www.arbolesornamentales.com/Sassafrasa

lbidum.htm. 1 p. 

Sargent, C.S. 1923. Manual of the trees of North 

America (exclusive of Mexico). Houghton 

Mifflin, Boston, MA. 910 p. 

Sullivan, J. 1993. Sassafras albidum. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/tree/ 

sasalb/all.html. 17 p. 

TreeGuide. 2003. Sassafras. http://www. 

treeguide.com/Species.asp?SpeciesID=953. 3 p. 

Vines, R.A. 1982. Trees of North Texas. 

University of Texas Press, Austin, TX. 466 p. 
 

684 

Schaefferia frutescens Jacq. Florida-boxwood

CELASTRACEAE 

Synonyms: Schaefferia completa Sw. 

 Schaefferia frutescens Jacq. var. buxifolia DC. 

 Schaefferia buxifolia (DC.) Nutt. 

John K. Francis 

General Description.—Florida-boxwood is a 

shrub or occasionally a small tree. Floridaboxwood is also known as yellowwood, jiga, 

cafeíllo, cabra cimarrona, amansa, guapo, jasmín, 

limoncillo, fruta de paloma, petit bois blanc, bois 

capable, petit garcon, and merisier (Little and 

others 1974). It has multiple stems, branching near 

the base, or a single stem, and usually reaches 

heights of 4 to 5 m. However, the Florida record 

tree measures 8 m in height and 18 cm diameter at 

breast height (Champion Tree Project 2001). The 

bark is light gray and nearly smooth, becoming 

increasingly furrowed in older shrubs and trees. 

The inner bark is light yellow. The roots are bright 

orange. There is a weak taproot supporting robust 

and stiff lateral roots and fine 3rd and 4th order 

roots. Twigs of Florida-boxwood are slender, 

slightly angled, hairless and pale yellow-green 

when young, turning light gray with age. The 

branch habit is upright to partially drooping. The 

leaves are yellow-green, alternate, simple, 

leathery, and elliptic to oval shaped with pointed 

ends. They measure 4 to 7 cm in length and 1.2 to 

2.5 cm in width. The flowers are small, pale green 

or greenish-yellow or greenish-white, and borne in 

small clusters at the leaf axiles. The fruits that 

arise from them are spherical to ovoid, 4 to 8 mm 

long, and red or orange-red. Each fruit contains 

two light-brown seeds (Howard 1989, Little and 

others 1974, Liogier 1994, Nelson 1996). 

Range.—Florida-boxwood is native to southern 

Florida, the Bahamas, the West Indies, Mexico 

through Central America, Colombia, Venezuela, 

and Ecuador (Liogier 1994). It is not known to 

have naturalized outside its native range. 

Ecology.—Florida-boxwood grows in soils 

ranging from sandy to clayey and from mildly 

alkaline to moderately acid derived from both 

sedimentary and igneous rocks. In Puerto Rico, 

the species inhabits areas receiving from 750 to 

about 1700 mm of mean annual precipitation and 

from near sea level to 600 m in elevation. In the 

upper range of rainfall, it grows only on 

excessively drained sites. Florida-boxwood is 

moderately shade tolerant. It is usually found in 

the understory of low-density forest where it 

grows well and reproduces. After the forest 

overstory is removed, Florida-boxwood grows 

well in full sun. In Puerto Rico, the species grows 

in remnant dry and moist forests and middle and 

late secondary forests. In Florida, it grows in 

tidewater areas and hammocks. The species is 

drought tolerant and somewhat resistant to soil 

salinity (Gilman 1999). Although the species is 

common in Puerto Rico and in much of its range, 

it is considered endangered in natural 

environments in Florida (Institute of Systematic 

Botany 2001). 

Reproduction.—Florida-boxwood flowers in 

March through May in Florida (Nelson 1996). 

Fruits collected in Puerto Rico averaged 0.142 +

0.002 g/fruit. Seeds separated from them averaged 

0.011 + 0.000 g/seed or 98,000 seeds/kg. Twentyone percent of these seeds germinated between 62 
 

685

and 93 days of sowing in moist peat. Germination 

is epigeal. The fruits are apparently eaten and the 

seeds dispersed by birds. 

Growth and Management.—Florida-boxwood is 

reported to have a high growth rate, withstands 

heavy pruning, and can be formed into hedges and 

shaped into small trees (Gilman 1999). 

Benefits.—Florida-boxwood makes superior 

quality hedges, screens, and accent plantings. It is 

also used for reclamation plantings (Gilman 

1999). The wood is light brown to yellow, hard 

and moderately heavy. It has been used in the past 

for wood engraving (Little and others 1974). It is 

also used, when available, for carving, for boxes 

(Gilman 1999), and, to a limited extent, for fuel. 

The leaves have been used in the past as 

emergency soap for the body and cloths (Liogier 

1990). Almost nothing is known about its forage 

value except that Key deer (Odocolleus virginanus 

clavium) will not eat it (Schaus and others 2001). 

Decoctions are used in herbal medicine for colds, 

flu, and chronic cough, and crushed leaves are 

used as a soothing bath for skin irritations (Liogier 

1990). 

References 

The Champion Tree Project. 2001. National 

champion trees: Florida. http://www. 

 championtrees.org/database/championsFL.htm. 

8 p. 

Gilman, E.F. 1999. Schaefferia frutencens. Fact 

Sheet FPS-540. University of Florida, 

Cooperative Extension Service. 3 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Institute of Systematic Botany. 2001. Atlas of 

Florida Plants: Schaefferia frutescens. 

University of South Florida. http:// 

plantatlas.usf.edu/main.asp?plantID==2227. 1 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico. Río Piedras, PR. 

461 p. 

 Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Schaus, C., S. Wade, and J. Dunan. 2001. Key 

deer and plants they won’t eat. 

http://monroe.ifas.ufl.edu/key_deer_plants.htm. 

3 p. 
 

686 

Schinus terebinthifolius Raddi Brazilian pepper tree 

ANACARDIACEAE 

Synonyms: Sarcotheca bahiensis Turcz. 

 Schinus antiarthriticus Mart. ex Marchand 

 Schinus mellisii Engl. 

 Schinus mucronulatus Mart. 

 Rhus terebinthifolia Schlecht. & Cham. 

John K. Francis 

General Description.—Brazilian pepper tree, also 

known as Christmas berry, Florida holly, pimienta 

de Brazil, copal, chichita, and aroeira pimenteira, 

is a multiple-stemmed evergreen shrub or small 

tree, usually 2 to 6 m in height and 3 to 12 cm in 

stem diameter. A few individuals of the species 

develop large central stems and a dense growth of 

low limbs and basal sprouts. The bark is smooth 

and gray. Wounds in the stem and twigs exude a 

resinous sap that turns black on exposure to air. 

The alternate, compound leaves have a rachis 3 to 

14 cm long, often winged, with five to nine 

leaflets. The leaflets are 1.5 to 7.5 cm long, 

lanceolate to elliptic, pointed at each end, with 

entire to serrate edges. The midribs, rachis, and 

petiole are often reddish, especially when young. 

Crushed foliage smells like turpentine. The 

inflorescences (panicles) are mostly born in the 

leaf axils near the twig ends and contain many 

small white flowers. Male and female flowers are 

borne on different plants (dioecious). The fruits 

are bright-red, fleshy drupes 4 to 6.5 mm in 

diameter with an aromatic brown pulp and an 

elliptic light-brown stone (Liogier 1988, Long and 

Lakela 1976, National Parks Service 2002). 

Range.—Brazilian pepper tree is a native of 

Brazil, Argentina, and Paraguay (Liogier 1988). It 

was introduced into Florida in 1898 and has since 

been distributed widely. The species has escaped 

cultivation and occupies huge areas in natural 

stands (Nelson 1996). It has also naturalized in 

Arizona, California, Hawaii, Puerto Rico, the U.S. 

Virgin Islands, and many tropical and subtropical 

countries (Little and others 1974, National Parks 

Service 2002). 

Ecology.—Brazilian pepper tree is an aggressive 

pioneer species that quickly colonizes disturbed 

areas. The species has an intermediate tolerance of 

shade and can survive and grow slowly under 

forest canopies until disturbance releases it. 

Although seedlings are quickly killed by 

inundation, large plants can withstand up to 6 

months of flooding. It is very drought resistant. 

Brazilian pepper tree also survives fire well and 

can withstand high winds without significant 

damage. It is apparently at home in tropical, 

Mediterranean, and desert climates (National 

Parks Service 2002). In Puerto Rico, the species is 

most common in low-elevation, moist limestone 

areas and nearby coastal plains. In Florida, it 

grows in mangrove associations, hammocks, 

pinelands, old fields, and disturbed areas (Long 

and Lakela 1976). 

Reproduction.—Brazilian pepper tree flowers 

most prolifically in the spring in Florida (Nelson 

1996). It flowers and fruits intermittently 

throughout the year in Puerto Rico (Little and 

others 1974). In Florida, the flowers are pollinated 

by a native syrphid fly (Fire Sciences Laboratory 
 

687

2002). Seeds from Brazil were reported to average 

80,600 seeds/kg (Instituto de Pesquisas e Estudos 

Florestais 2002). Germination of intact fruits 

(without the exocarp digested or manually 

removed) is minimal, and seeds do not remain 

viable in the soil bank more than 9 months 

(Panetta and Mckee 1997). Birds, particularly 

robins, mockingbirds, and cedar waxwings, are the 

chief dispersers of seeds in Florida (Fire Sciences 

Laboratory 2002). Brazilian pepper tree is 

propagated from both seeds and cuttings (Little 

and others 1974). It readily sprouts when cut. 

Growth and Management.—Brazilian pepper 

tree is fast growing and in certain circumstances is 

capable of growing to large diameters. The U.S. 

Champion tree in Florida measures 134 cm in 

diameter at breast height and 10.7 m in height 

(American Forests 2002). Brazilian pepper tree 

has shown large growth responses to inoculation 

with mycorrhizal fungi and superphosphate 

(Carneiro and others 1996). Hand pulling, 

bulldozing, prescribed fire, and spraying with 

herbicides have all been used with varying degrees 

of success to control Brazilian pepper tree. So far, 

biological control methods have failed (National 

Parks Service 2002). 

Benefits and Detriments.—Widely introduced as 

an ornamental, Brazilian pepper tree was popular 

for its red berries and bright green foliage. It is 

still used for Christmas decorations. Because it 

and the species growing in its understory do not 

burn readily (Fire Sciences Laboratory 2002), 

Brazilian pepper tree has been recommended for 

planting as fire resistant barriers (Castronovo 

1997). Goats browse on the foliage with no ill 

effects (National Parks Service 2002). The species 

is a honey plant (Little and others 1974) and the 

wood has been used for fuel, lumber, stakes, posts, 

and railway sleepers (Fire Sciences Laboratory 

2002). The dried fruits are marketed in Brazil as a 

substitute for black pepper (Piper nigrum) 

(personal communication with J.A. Parrotta, 

USDA Forest service, Washington, DC). A 

decoction of bark is used in baths to relieve 

rhumatisum and back pain (Liogier 1990). The 

essential oils from leaves and flowers are made up 

of a mixture of a large number of chemicals, the 

most important of which are alpha-pinene (24 

percent), limonene (12 percent), and p-cymene (14 

percent) (Singh and others 1998). Extracts of 

Brazilian pepper tree were shown to be the most 

effective of a number aromatic and medicinal 

plant species in suppressing several important 

pathogenic bacteria (Martínez and others 1996, 

Siddiqui and others 1995). Despite these benefits, 

Brazilian pepper tree is often undesirable outside 

its native range. It is so well adapted and 

aggressive in Florida and Hawaii that it suppresses 

and replaces native vegetation, including 

endangered species. For this reason it is banned 

from cultivation there (Fire Sciences Laboratory 

2002, Nelson 1996). Brazilian pepper tree is 

reported to have an allelopathic effect on 

competing plants (National Parks Service 2002). 

The plant causes skin irritation similar to poison 

ivy (Toxicodendron radicans Linn.) and 

respiratory difficulties in some people when it is in 

bloom (Nelson 1996). It is a host of the black twig 

borer of coffee, Xylosandrus compactus (Eichoff) 

(College of Tropical Agriculture and Human 

Resources 2002). The fruits are eaten by mammals 

and birds, but excessive feeding has been blamed 

for massive bird kills in Florida (National Parks 

Service 2002), and the unripe fruits can be fatal to 

horses (Fire Sciences Laboratory 2002). 

References 

American Forests. 2002. National register of big 

trees. American Forests, Washington, DC. http:// 

www.americanforests.org/resources/bigtrees/ 

regester.php?details=2297. 1 p. 

Carneiro, M.A.C., J.O. Siqueira, A.C. Davide, L.J. 

Gomes, N. Curi, and F.R. do Vale. 1996. 

Mycorrhizal fungi and superphosphate and the 

growth of tropical woody species. Scientia 

Forestalis 50: 21-36. 

Castronovo, T. 1997. Fire resistive landscaping 

can save your house and your life. 

http://www.themastergardenershow.com/ 

fire_resistive_landscaping_.htm. 5 p. 

College of Tropical Agriculture and Human 

Resources. 2002. Crop knowledge master: 

Xylosandrus compactus (Eichoff). University of 

Hawaii. http://extento.hawaii.edu/kbase/Type/ 

xylosand.htm. 4 p. 

Fire Sciences Laboratory. 2002. Fire effects 

information database: Schinus terebinthifolius. 

Rocky Mountain Research Station, Missoula, 

MT. http://www.fs.fed.us/database/feis/plants/ 

tree/ schter/all.html. 9 p. 
 

688 

Instituto de Pesquisas e Estudos Florestais. 2002. 

Tabla de preços de sementes de espécies nativas 

e exóticas. http://www.ipef.br/especies/ 

natexoticas.asp. 2 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. University of Miami Press, 

Coral Gables, FL. 962 p. 

Martínez, M.J., J. Betancourt., N. A. González., 

and A. Jauregui. 1996. Screening of some Cuban 

medicinal plants for antimicrobial activity. 

Journal of Ethnopharmacology 52(3): 171-174. 

National Parks Service. 2002. Exotic weeds I. 

http://www.nature.nps.gov/wv/ipm/exweeds1.ht

m. 9 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Panetta, F.D. and J. McKee. 1997. Recruitment of 

the invasive ornamental, Schinus 

terebinthifolius, is dependent upon frugivores. 

Australian Journal of Ecology 22(4): 432-438. 

Siddiqui, R.R., Uzma-Zafar, S.S. Chaudhry, and 

Hamid-Ahamad. 1995. Antimicrobial activity of 

essential oils from Schinus terebinthifolius, 

Cypress sempervirens, Citrus limon, Ferula 

assafoetida. Part 1. Pakistan Journal of 

Scientific and Industrial Research. 38(9-10): 

358-361. 

Singh, A.K., J. Singh, K.C. Gupta, and J.J. 

Brophy. 1998. Essential oil of leaves and 

inflorescence of Schinus terebinthifolius: an 

exotic plant of India. Journal of Essential Oil 

Research 10(6): 697-699. 
 

689

Securidaca virgata Sw. bejuco de sopla 

POLYGALACEAE 

Synonyms: Elsota virgata (Sw.) Kuntze 

John K. Francis 

General Description.—Bejuco de sopla, also 

known as jaboncillo and maravedí, is a woody 

vine-like shrub that ascends trees and other 

vegetation and can reach 9 m in length and 3 cm 

in basal stem diameter. The plants have a tap and 

lateral root system of tough and flexible roots with 

a moderate amount of fine roots. The stems give 

rise to few branches until they reach full or nearly 

full light after which there are abundant secondary 

and tertiary branches. The greenish, cylindrical 

woody stems have pronounced annual rings. The 

fine branches are green and produce relatively few 

leaves, which they lose during dry seasons. 

Foliage is found only on the current year’s growth. 

The alternate leaves are oval, have a 1 mm petiole, 

and are 10 to 20 mm long by 8 to 15 mm broad. 

Small, 8 mm-long, pink to violate, fragrant 

flowers are borne near the ends of the branches in 

paniculate racemes. The fruit, a reticulate-veined 

samara 2 to 4 cm long and about 1 cm broad, turns 

from green to yellow to light tan as it ripens, and 

contains a single seed (Acevedo-Rodrígiez 1985, 

author’s observations, Grisebach 1963, Liogier 

1988). 

Range.—Bejuco de sopla is native to Puerto Rico, 

its offshore island, Vieques, as well as Jamaica, 

and Cuba (Acevedo-Rodrígiez 1985, Grisebach 

1963, Liogier 1988). 

Ecology.—Bejuco de sopla grows on a wide 

variety of soils over both sedimentary and igneous 

rocks at lower and middle elevations. Annual 

rainfall in its native areas ranges from about 900 

to 2500 mm/year. It sustains itself on other 

vegetation by twining around stems and branches 

(Vélez and van Overbeek 1950). Although it 

depends on trees and shrubs for support, it does 

not smother them, tending to hang down from side 

branches. Bejuco de sopla is intolerant of shade 

and does not grow under closed, shady forest 

canopies. It grows in brushy pastures, old fields, 

fencerows, roadsides, canyons, gallery creek 

bottoms, secondary forest, and low-density 

remnant forests. The species does not form pure 

stands or dense thickets, although is may grow in 

mixture with other vines and shrubs in thickets. 

Reproduction.—Bejuco de sopla flowers and 

fruits sporadically throughout the year (AcevedoRodrígiez 1985, Vélez and van Overbeek 1950). A 

collection of fruits from Puerto Rico weighed an 

average (air-dried) of 0.0728 + 0.0011 g/fruit. The 

acorn-like seeds separated from the wing weighed 

an average of 0.0455 + 0.0012 g/seed or 22,000 

seeds/kg. Sown in moist potting mix, 22 percent of 

these seeds germinated in 32 days by which time 

some of the seedlings began to die from 

“damping-off” (fungal species unknown). The 

survivors were 10 cm tall 2 months after sowing 

and ready to prick out. Fruits of bejuco de sopla 

are mature and ready to harvest when they are dry 

on the tips or have a yellow color. They should 

then be air-dried before storage or planting. If the 

seeds are separated, it must be done by hand, a 

laborious process that probably is not necessary. 

Bejuco de sopla vines layer (root) whenever they 

come in contact with the ground. Although 

untested, this probably indicates that the species 

can be propagated by air layering and from 

cuttings. The seeds are dispersed as they spiral 

sideways in the wind or still air. Seedlings are 

usually well scattered. 

Growth and Management.—Bejuco de sopla has 

a moderate growth rate. Height growth of 

established plants is 1 m or more per year and 

diameter growth is about 3 mm/year. Individual 
 

690 

stems live 5 to 10 years or more; plants may last 

longer by resprouting. Bejuco de sopla is not 

abundant, aggressive, or weedy. Should it be 

necessary to eliminate individual plants, grubbing 

them out will probably be effective. 

Benefits.—Bejuco de sopla contributes to 

biodiversity, helps protect the soil, and furnishes 

wildlife cover. It is a honey plant (Marcano-F. 

1973). The roots, principally, are used to treat flu, 

laryngitis, edema, urinary infections, and as an 

expectorant and a diuretic (Liogier 1990). The 

plant is prescribed by traditional Latin healers in 

New York City to treat uterine fibroids (Balick 

and others 2000). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Balick, M.J., F. Kronenberg, A.L. Ososki, M. 

Reiff, A. Fugh-Berman, B. O’Connor, M. Roble, 

P. Lohr, and D. Atha. 2000. Medicinal plants 

used by Latino healers for woman’s health 

conditions in New York City. Economic Botany 

54(3): 344-357. 

Grisebach, A.H.R. 1963. Flora of the British West 

Indian Islands. J. Cramer-Weinheim, New York. 

789 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR 481 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Marcano-F., E. de J. 1973. La flora apícola de la 

República Dominicana. http://marcano. 

freeservers.com/nature/estudios/apicola/floraap1

.html. 7 p. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Río Piedras, PR. 497 p. 
 

691

Senecio flaccidus Less. threadleaf groundsel 

ASTERACEAE 

Synonyms: Senecio douglasii DC. 

James E. Nellessen 

 

General Description.—Threadleaf groundsel is 

also known as threadleaf butterweed, Douglas 

groundsel or ragwort, felty groundsel, sand wash 

groundsel, comb butterweed, old man, yerba cana, 

squawweed, creek senecio, and cenicillo (Elmore 

1976, Epple 1995, Stubbendieck and others 1997). 

This species commonly reaches heights of 80 to 

130 cm and under ideal conditions may reach 2 m. 

It is probably best classified as a subshrub. It 

forms woody stems to a meter or so but often dies 

back to less than 50 cm from the ground-line. It 

tends to become somewhat straggly as branches 

may lean over in different directions, some 

becoming almost decumbent. The leaves are 

alternate, pinnately compound (sometimes not 

divided), narrow and thread-like, up to 12 cm in 

length, and usually covered with whitish hairs 

giving the plant an obvious grayish color (Carter 

1997, Ivey 1995, Kearney and others 1951, Martin 

and Hutchins 1980-81). It may have some 

tendencies to be evergreen in the southern portion 

of its range. 

Taxonomy.—There has been a profusion of 

taxonomic names and varietal subclassifications 

attached to this species, making for a complicated 

taxonomy. The oldest specific name applied to this 

species complex is now known to be Senecio 

flaccidus Lessing (Turner and Barkley 1990). 

Three varieties are recognized with the following 

synonymies (Allred 2002, Kartesz 1994). Variety 

1: S. flaccidus Less. var. flaccidus, with four 

synonyms; S. douglasii DC var. longilobus 

(Benth.) L. Benson, S. filifolius Nutt., S. flaccidus

Nutt. var. jamesii Torr. & Gray, and S. longilobus

Benth. This variety tends to be more uniformly 

white hairy with the column of the flower head 

narrower (8 to 10 mm wide at summit). Variety 2: 

S. flaccidus Less. var. douglasii (DC) B. Turner & 

T. Barkley, with two synonyms; S. douglasii DC 

var. douglasii, and S. warnockii Shinners. This 

variety is not white hairy throughout and the heads 

are wider (10 to 15 mm). Variety 3: S. flaccidus

Less. var. monoensis (Greene) B. Turner & T. 

Barkley, with two synonyms; S. douglasii DC var. 

monoensis (Greene) Jeps., and S. monoensis 

Greene. This variety is generally hairless 

throughout, more herbaceous, and has a more 

comb-like leaf, the linear lobes curling upward. 

Range.—Threadleaf groundsel occurs from 

western Texas, west through New Mexico, 

Arizona, to California, north to Wyoming and 

Nebraska, Utah, and Nevada, and south into 

northern Mexico. It is a wide ranging species 

occurring in a variety of soil types and elevations 

from about 760 to 2,440 m. 

 

Ecology.—This species grows in short grass 

prairies of the southwestern Great Plains and a 

wide variety of scrubland and grassland habitats in 

the Southwest. It has been considered a diagnostic 

or climax species for desert grassland (DickPeddie 1993). It tends to become more abundant in 

disturbed habitats and overgrazed rangelands and 

often grows in washes. It is a taprooted plant that 

grows relatively quickly and tends to be a shortlived perennial shrub to subshrub. Oxygen flux, as 

it relates to photosynthesis, from leaves and twigs 

was constant with increasing drought stress, up 

until leaf drop occurred (Ehleringer and Cooper 

1992). Twigs may serve to maintain some 

photosynthesis through drought periods when new 

foliage cannot easily be produced, such as during 

dry springs and early summers. Threadleaf 

groundsel was one of several shrub and subshrub 

species, studied in Mojave and Sonoran desert 

areas that generated substantial net carbon gains 
 

692 

from twig photosynthesis (Comstock and others 

1988). 

Reproduction.—Threadleaf groundsel generally 

blooms in the late summer and fall but will bloom 

earlier in the summer if sufficient moisture is 

available. It produces numerous yellow flower 

heads with eight to 17 ray florets, the rays 10 to 15 

mm long (Carter 1997, Martin and Hutchins 1980-

81). Seeds (fruits) are wind-borne with a white 

fluffy pappus. 

Growth and Management.—Threadleaf 

groundsel is a fast growing, short lived perennial. 

Plants may live 3 to 6 years or more. It increases in 

rangelands as a result of overgrazing pressure. Its 

abundance may be controlled through proper 

grazing management and careful use of herbicides. 

Spraying with 2,4-D has been used to control this 

species. 

Benefits and Disadvantages.—Because this 

species will colonize open disturbed areas, it helps 

achieve a quick ground cover in a natural 

succession process. The perennial nature of the 

plant further helps stabilize soil for longer-lived 

perennials to eventually become established. 

Direct human uses are minimal, but it was once 

used medicinally by Native Americans. It is a poor 

forage plant for cattle and horses and is known to 

be toxic (Gay and Dwyer 1998, Stubbendieck and 

others 1997, Warnock 1974) and causes liver 

damage when consumed in large quantities 

(Kirkpatrick 1992). All parts of the plant are 

poisonous, younger leaves more so, and poisoning 

is more of a problem on drier range conditions. 

Symptoms in cattle may include a dry scaly nose, a 

rough coat, and continuous walking with no 

direction (Phillips Petroleum Company 1963). 

Because it has low palatability, it tends to be 

avoided except during long droughts or in heavily 

overgrazed areas where it may increase in 

abundance (USDA 1937). Toxicity of threadleaf 

groundsel may be attributed to the pyrrolizidine 

alkaloids (PA). About 50 percent of all identified 

PA are toxic, and some are carcinogenic in rodents 

(Stegelmeier and others 1999). Threadleaf 

groundsel contains the pyrrolizidine alkaloids: 

senecionine, seneciphylline, florosenine, 

otonecine-based florosenine, and retrorsine 

(Cooper and others 1996). As with many toxic 

plants, concentration and preparation are critical 

factors. For example, the Navajo would boil entire 

plants and drink the mixture to assist the voice and 

singing during some ceremonial songs (Elmore 

1976). The Navajo would also use the flower 

heads to clean cactus fruits of their spines. 

References

Allred, K.W. 2002. A Working Index of New 

Mexico Vascular Plant Names. New Mexico 

State University. http://web.nmsu.edu/~kallred/ 

herbweb/ [not paged]. 

Carter, J.L. 1997. Trees and Shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Comstock, J.P., T.A. Cooper, and J.R. Ehleringer. 

1988. Seasonal patterns of canopy development 

and carbon gain in nineteen warm desert shrub 

species. Oecologia, Berlin 75(3): 327-335. 

Cooper, R.A., R.J. Bowers, C.A. Beckham, and 

R.J. Huxtable. 1996. Preparative separation of 

pyrrolizidine alkaloids by high-speed counter 

current chromatography. Journal of 

Chromatography 732(1): 43-50. 

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation: Past, Present and Future. University 

of New Mexico Press, Albuquerque, NM. 244 p. 

Ehleringer, J.R. and T.A. Cooper. 1992. On the 

role of orientation in reducing photoinhibitory 

damage in photosynthetic-twig desert shrubs. 

Plant Cell and Environment 15(3): 301-306. 

Elmore, F.H. 1976. Shrubs and Trees of the 

Southwest Uplands. Southwest Parks and 

Monuments Association, Tucson, AZ. 214 p. 

Epple, A.O. 1995. A Field Guide to the Plants of 

Arizona. Falcon Publishing Inc., Helena, MT. 

347 p. 

Gay, C. W., Jr. and D. D. Dwyer. 1998 Reprint. 

New Mexico Range Plants. Cooperative 

Extension Service Circular 374. Revisions by: 

C. Allison, S. Hatch, and J. Schickedanz. New 

Mexico State University, Las Cruces, NM. 84 p. 

Ivey, R.D. 1995. Flowering Plants of New Mexico, 

3rd Edition. Published by the author, NM. 

504 p. 

Kartesz, J.T. 1994. A Synonymized Checklist of 

the Vascular Flora of the United States, Canada, 
 

693

and Greenland, Vol. 1, 2nd Edition. Biota of 

North America Program of the North Carolina 

Botanical Garden. Timber Press, Portland, OR. 

622 p. 

Kearney, T.H., R. Peebles, and Collaborators. 

1951 with 1960 supplement. Arizona Flora. 

University of California Press, Berkeley, CA. 

1,085 p. 

Kirkpatrick, Z. M. 1992. Wildflowers of the 

Western Plains. University of Texas Press. 

Austin, TX. 240 p. 

Martin, W.C. and C.E. Hutchins. 1980-1981 

(reprinted 2001). A Flora of New Mexico. Vol. 

2. Bishen Singh Mahendra Pal Singh (India) and 

Koeltz Scientific Books (Germany). p. 1,277-

2,591. 

Phillips Petroleum Company. 1963. Pasture and 

Range Plants. Phillips Petroleum Co., 

Bartlesville, OK. 176 p. 

Stegelmeier, B.L., J.A. Edgar, S.M. Colegate, D.R. 

Gardner, T.K. Schoch, R.A. Coulombe, and R.J. 

Molyneux. 1999. Pyrrolizidine alkaloid plants, 

metabolism and toxicity. Journal of Natural 

Toxins 8(1): 95-116. 

Stubbendieck, J., S.L. Hatch, and C.H. Butterfield. 

1997. North American Range Plants, 5th Edition. 

University of Nebraska Press, Lincoln, NE. 

501 p. 

Turner, B.L. and T.M. Barkley. 1990. Taxonomic 

overview of the Senecio flaccidus complex in 

North America, including Senecio douglasii. 

Phytologia 69(1): 51-55. 

U.S. Department of Agriculture, Forest Service. 

1937 (1988 Dover edition). Range Plant 

Handbook. Dover Publications Inc., New York. 

816 p. 

Warnock, B.H. 1974. Wildflowers of the 

Guadalupe Mountains and the Sand Dune 

Country, Texas. Sul Ross State University, 

Alpine, TX. 176 p. 
 

694 

Senecio spartioides Torr. & Gray many-headed groundsel 

ASTERACEAE 

Synonym: Senecio multicapitatus Greenm. ex Rydb. 

James E. Nellessen 

General Description.—Many-headed groundsel is 

also known as broom groundsel, broom 

butterweed, or grass-leaved ragwort (Elmore 1976, 

Epple 1995). This plant produces many stems from 

a large woody crown, in a tight to spreading 

cluster, giving it a broom-like appearance. Plants 

range in height from 20 cm to slightly more than 1 

m. In large plants, some stems may lean over, 

becoming almost decumbent. The leaves are bright 

green, both lobed (pinnately) or nonlobed, 

alternate, linear, 4 to 10 cm long, and to 5 mm 

wide, generally hairless or slightly pubescent 

(Carter 1997, Great Plains Flora Association 1986, 

Ivey 1995, Martin and Hutchins 1980-81, Weber 

and Wittman 2001a and 2001b). This plant is best 

classified as a subshrub because even though the 

stems do become woody, especially the bases, it 

often dies back during the winter to less than 20 

cm above the ground-line. 

Taxonomy.—Two varieties are recognized: the 

typical variety, S. spartioides var. spartioides, and 

S. spartioides var. multicapitatus (Greenm. ex 

Rydb.) S.L. Welch (Allred 2002). The leaves of 

variety spartioides are generally simple, or 

infrequently with small lobes near the base, and 

the plant is hairless. The variety multicapitatus

tends to have leaves irregularly pinnately divided, 

more hairs, and smaller, more numerous flower 

heads. The latter variety is also synonymized with 

the full species designation: S. multicapitatus

Greenm. ex Rydb. (Kartesz 1994). 

Range.—This is a wide-ranging species occurring 

from western Texas, west through New Mexico, 

northern Mexico, into Arizona, Utah, and 

California, north to Wyoming, western Nebraska, 

and southwestern South Dakota. Variety 

spartioides tends to occur further north and west 

within this range, while variety multicapitatus

tends to be more southern in its distribution 

(Kearney and others 1951, Great Plains Flora 

Association 1986). Within its range it occurs at 

elevations from about 1,070 m to 2,745 m. 

Ecology.—Many-headed groundsel occurs on 

plains, open slopes, valleys, arroyos, and semistabilized dunes in piñon-juniper woodlands, 

ponderosa pine forests, and desert areas. It tends to 

be an early colonizer of disturbed soil situations, is 

a prolific seed producer because of the numerous 

flower heads, and is generally a short-lived (2 to 4 

years) perennial. 

Reproduction.—This species produces numerous 

yellow-flowered heads, the ray flowers numbering 

from four to eight per head (over 18 flowers with 

the disk flowers included). The heads are 7 to 12 

mm high, 3 to 7 mm wide, subcylindric to 

cylindric with about eight to 12 phyllaries, and the 

rays are from 7 to 12 mm long. It generally blooms 

in the late summer and fall but will bloom early in 

the summer given sufficient moisture. Hence, the 

full range of potential blooming runs from May 

into early November, depending on local 

environmental conditions. A bright white fluffy 

pappus enables easy dispersal of seeds by the 

wind. 

Growth and Management.—Seeds will 

germinate under moist conditions in the fall, the 

seedlings over-wintering. Growth continues slowly 

in the spring but is rapid once summer rains begin. 

Many-headed groundsel is a short-lived perennial. 

Most plants appear to survive only 2 to 4 years, 

some perhaps slightly longer. This species, 

although toxic to livestock, is not palatable and is 

generally not highly abundant or dominant in most 

habitats to be a significant management problem. 
 

695

Benefits.—Many-headed groundsel will colonize 

open disturbed areas and provide quick ground 

cover in a natural succession process. Although 

relatively short-lived the perennial nature of the 

plant further helps stabilize soil for longer-lived 

perennials to eventually become established. 

Many-headed groundsel is toxic to cattle, but it is 

infrequently eaten (USDA 1937, Kearney and 

others 1951, Warnock 1974). This species should 

be utilized more in urban and native plant 

landscaping. When in bloom it bears, as the name 

implies, numerous yellow flower heads, adding 

considerable color to a desert backyard. It is a 

prolific seed producer and will easily regenerate 

itself. This may pose a problem for some 

gardeners, managing the abundance of seedlings, 

although many of the seedlings will not survive to 

maturity within a natural landscape anyway. 

References

Allred, K.W. 2002. A Working Index of New 

Mexico Vascular Plant Names. Available on a 

New Mexico State University web page, 

http://web.nmsu.edu/~kallred/herbweb/ [not 

paged]. 

Carter, J.L. 1997. Trees and Shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Elmore, F.H. 1976. Shrubs and Trees of the 

Southwest Uplands. Southwest Parks and 

Monuments Association, Tucson, AZ. 214 p. 

Epple, A.O. 1995. A Field Guide to the Plants of 

Arizona. Falcon Publishing Inc., Helena, MT. 

347 p. 

Great Plains Flora Association. 1986. Flora of the 

Great Plains. University Press of Kansas, KS. 

1,392 p. 

Ivey, R.D. 1995. Flowering Plants of New Mexico, 

3rd Edition. Published by the author, NM. 

504 p. 

Kartesz, J.T. 1994. A Synonymized Checklist of 

the Vascular Flora of the United States, Canada, 

and Greenland, Vol. 1, 2nd Ed. Biota of North 

America Program of the North Carolina 

Botanical Garden. Timber Press, Portland, OR. 

622 p. 

Kearney, T.H., R. Peebles, and Collaborators. 

1951 with 1960 supplement. Arizona Flora. 

University of California Press, Berkeley, CA. 

1,085 p. 

Martin, W.C. and C.E. Hutchins. 1980-1981 

(reprinted 2001). A Flora of New Mexico. 

Bishen Singh Mahendra Pal Singh, India and 

Koeltz Scientific Books, Germany. Vol. 2, p. 

1,277-2,591. 

U.S. Department of Agriculture Forest Service. 

1937 (1988 Dover edition). Range Plant 

Handbook. Dover Publications Inc., New York. 

816 p. 

Warnock, B. H. 1974. Wildflowers of the 

Guadalupe Mountains and the Sand Dune 

Country, Texas. Sul Ross State University, 

Alpine, TX. 176 p. 

Weber, W.A. and R.C. Wittman. 2001a. Colorado 

Flora: Eastern Slope. University Press of 

Colorado, Boulder CO. 521 p. 

Weber, W.A. and R.C. Wittman. 2001b. Colorado 

Flora: Western Slope. University Press of 

Colorado, Boulder CO. 488 p. 
 

696 

Senna atomaria (L.) Irwin & Barneby flor de San José 

FABACEAE 

Synonyms: Cassia atomaria L. 

 Cassia emarginata L. 

 Cassia arborescens Miller 

 Cassia triflora Vahl 

Isandrina emarginata (L.) Britt. & Rose (and others: International 

Legume Database and Information Service 2002) 

John K. Francis 

General Description.—Flor de San José is also 

known as yellow candlewood, senna-tree, palo de 

chivo, palo de burro, cañafístola cimarrona, 

carbonera, alcaparro, chile perro, palo de zorrillo, 

arguchoco, vainillo, chivato, carángano, platanillo, 

bois cabrite, casse marron, and petite casse. It is a 

flowering, drought-deciduous shrub or small tree 2 

to 12 m in height that occasionally exceeds 20 cm 

in stem diameter. The plant may or may not have 

multiple stems but almost always supports stout 

branches relatively low to the ground. The 

branches are brittle. It is supported by a tap and 

extensive lateral root system that has slender fine 

roots. The roots are woody and flexible with a 

dark gray bark over a pale yellow interior. The 

crown may be described as spreading and branchy 

but not dense. The bark is light gray. Leaves, 

twigs, and other vegetative parts have an 

unpleasant odor and are covered with soft hairs. 

The alternate, pinnate leaves have petioles 2 to 5 

cm long and two to five pairs of ovate to obovate 

entire leaflets, 2 to 11 cm long. The flower 

clusters are corymbose racemes that bear many 

flowers and arise from the bases of the upper 

leaves. The flowers have five unequal sepals and 

five unequal yellow petals 8 to 15 mm long. The 

fruits are flattened, linear, red-brown to black, 

legumes 12 to 35 cm long and 8 to 12 mm broad, 

and contain numerous brown seeds 4 to 5 mm 

long (Howard 1988, Liogier 1988, Little and 

others 1974, Secretaría de Medio Ambiente, 

Recursos Naturales y Pesca 2002, Stevens and 

others 2001). 

Range.—Flor de San José is native to the 

Bahamas, the West Indies, Mexico, Central 

America, Colombia, Venezuela, and Guyana 

(Howard 1988, Little and others 1974). It has been 

planted in Hawaii and probably other areas outside 

the natural range as an ornamental (Neal 1965). 

Ecology.—Flor de San José is common to rare in 

coastal and upland dry forests at elevations from 

near sea level to 1,100 m that receive annual 

rainfalls of about 600 to 1,000 mm. Soils, often 

rocky and excessively drained, developed from 

both igneous and sedimentary parent materials 

with a wide range of textures and with pH’s that 

are mildly acid to mildly alkaline. Flor de San José 

is moderately intolerant of shade; it grows in the 

open, as a codominant in low forest, and in the 

understory of open forests. The species may be 

found in remnant and late secondary forests. 

Reproduction.—Flor de San José flowers and 

fruits throughout the year in Nicaragua (Stevens 

and others 2001). Although flowers and fruits 

have been observed at every season of the year in 

Puerto Rico, individual trees normally flower and 

fruit once per year. Because of extended 

flowering, flowers and fruits may be present on 

the same tree. Seeds collected by the author from 

Puerto Rico weighed an average of 0.0238 +

0.0004 g/seed. Sown on moist filter paper, 96 

percent germinated between 3 and 15 days after 

sowing. Germination is epigeal. A test of percent 

germination of sowings throughout the year in 

Mexico resulted in emergence varying from 1.2 to 
 

697

44 percent over 45 day periods depending on 

ambient temperature and precipitation (Vera and 

Sánchez 1995). Reproduction may be common in 

areas overgrazed by cattle. Seeds germinate after 

pods rot on the ground under mother trees 

(Stevens and others 2001) and may be dispersed to 

a limited degree by ruminants. 

Growth and Management.—Flor de San José 

has a moderate growth rate. Although planted as 

an ornamental, the author is not aware of any 

reports of wildland plantings. Management and 

protection of existing stands are recommended 

(Secretaría de Medio Ambiente, Recursos 

Naturales y Pesca 2002). 

Benefits.—Flor de San José is planted widely but 

not abundantly as an ornamental for its beautiful 

floral display and moderate size. The sapwood is 

yellow and hard, and the heartwood is brown and 

hard with a density of 0.57 to 0.85 g/cm3

 (Timyan 

1996). It is used for fuel and rude construction 

(Holdridge and Poveda 1975, Secretaría de Medio 

Ambiente, Recursos Naturales y Pesca 2002). 

Cows eat the pods but do not browse the leaves. 

Some of the common names imply that goats 

browse the species. Meal made from seed-free 

pods of flor de San José was eagerly consumed by 

lambs in an acceptance test (Palma and Román 

2002b); the meal contained 6.5 percent crude 

protein, 6.8 percent fat, 4.5 percent ash, and 32.1 

percent crude fiber (Palma and Román 2002a). It 

has been demonstrated that Baird’s tapir (Tapirus 

bairdii) kills 100 percent of the flor de San José 

seeds when it consumes the pods, apparently 

because they germinate within the gut (Olmos 

1997). Flor de San José is the principal or a key 

larval host for the yellow angled-sulphur butterfly, 

Anteos maerula (Fabricius), and the large moth, 

Sphingicampa montana (Packard) (Northern 

Prairie Wildlife Research Center 2002, Savela 

2002). A tea made from the leaves is used as a 

purgative (Secretaría de Medio Ambiente, 

Recursos Naturales y Pesca 2002). 

References 

Holdridge, L.R. and L.J. Poveda A. 1975. Arboles 

de Costa Rica. Centro Cientifico Tropical, San 

José, Costa Rica. 546 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

International Legume Database and Information 

Service. 2002. Senna atomaria (L.) H. Irwin & 

Barneby. http://www.ildis.org/LegumeWeb/ 

6.00/taxa/15514.shtml. 2 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Press, 

Honolulu, HI. 924 p. 

Northern Prairie Wildlife Research Center. 2002. 

Butterflies of North America: Yellow angledsulphur (Anteos maerula). U.S. Geological 

Survey. http://www.npwrc.usgs.gov.resource. 

distr/lepid/bflyusa/usa/723.htm. 3 p. 

Olmos, F. 1997. Tapirs as seed dispersers and 

predators. UNCN/SSC Tapir Specialist Group. 

http://www.tapirback.com/tapirgal/iucn-ssc/tsg/ 

action97/ap97-05.htm. 14 p. 

Palma, J.M. and L. Román. 2002a. Frutos de 

especies arbóreas leguminosas y no 

leguminosas para la alimentación de rumiantes. 

Centro Universitario de Investigación y 

Desarrollo Agropecuario. http://lead. 

virtualcentre.org/es/ele/conferencia2/vbconfe15

.htm. 15 p. 

Palma, J.M. and L. Román. 2002b. Prueba de 

selectividad con ovinos de pelo de harinas de 

frutos de especies arbóreas. Centro 

Universitario de Investigación y Desarrollo 

Agropecuario. http://www.cipav.org.co/ 

redagrofor/memoirias99/PalmaJM.htm. 4 p. 

Savela, M. 2002. Sphingicampa Walsh, 1864. 

http://www.funet.fi/pub/sci/bio/life/insecta/lepi

doptera/ditrysia/bombycoidea/saturniidae/ 

ceratocampinae/sphingicampa/. 3 p. 
 

698 

Secretaría de Medio Ambiente, Recursos Naturales 

y Pesca. 2002. Especies forestales no 

maderables y maderables no tradicionales de 

zonas áridas y semiáridas: Senna atomeria (L.) 

Irwin et Barneby. http://www.semarnat.gob.mx/ 

pfnm3/fichas/senna_atomaria.htm. 2 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

1. Missouri Botanic Garden Press, St. Louis, 

MO. 943 p. 

Timyan, J. 1996. Bwa yo: important trees of Haiti. 

South-East Consortium for International 

Development, Washington, DC. 418 p. 

Vera-S., F. and M. Sánchez-R. 1995. Epoca de 

siembra de doce especies forestales de Baja 

California Sur. Revista de Ciencia Forestal en 

México 20(78): 59-78. 
 

699

Senna occidentalis (L.) Link coffee senna 

FABACEAE 

Synonyms: Cassia occidentalis L. 

 Ditremexa occidentalis (L.) Britt. & Rose 

John K. Francis 

General Description.—Coffee senna is also 

known as stinking weed, miki palaoa, hedionda, 

bucho, pico de pájaro, furrusca, bois puante, 

fedegoso, and many other names. The species 

varies from a semi-woody annual herb in warm 

temperate areas to a woody annual shrub or 

sometimes a short-lived perennial shrub in frostfree areas (Haselwood and Motter 1966, Henty 

and Pritchard 1975, Holm and others 1997). It 

usually matures at from 0.5 to 2.0 m in height. In 

Brazil it is reported to reach 5 to 8 m in height 

(Raintree 2002). Coffee senna produces a hard, 

woody tap root with relatively few laterals. It 

usually has a single purplish stem and sparse 

branching. Young stems are four-angled, 

becoming rounded with age. The crushed foliage 

has an unpleasant odor. Compound alternate 

leaves have four to six pairs of glabrous leaflets 

and a gland near the base of the petiole. Leaflets 

are ovate to ovate-lanceolate, pointed at the tip 

and rounded at the base. Inflorescences are fewflowered axillary racemes with yellow-petaled 

flowers about 2 cm across. The legumes (pods) are 

brown, flat, slightly curved and 5 to 12 cm long. 

They contain 40 or more brown to dark-olive, 

ovoid seeds about 4 mm long. The species has 2n 

= 26, 28 chromosomes (Henty and Pritchard 1975, 

Liogier 1988, Long and Lakela 1976, Stevens and 

others 2001). 

Range.—Coffee senna grows throughout the 

tropics and subtropics (Liogier 1988, Stevens and 

others 2001) including the United States from 

Texas to Iowa eastward, Hawaii, the Pacific Island 

Territories, Puerto Rico, and the U.S. Virgin 

Islands (Natural Resources Conservation Service 

2002). It appears to be of South American or New 

World origin (Haselwood and Motter 1966, Henty 

and Pritchard 1975, Raintree 2002). 

Ecology.—In Cuba, coffee senna is said to be 

most common in fertile, cultivated areas (Sánchez 

and Uranga 1993) and in Nicaragua it also grows 

in disturbed sites (particularly fertile ones), river 

overflow areas, meadows, and forests from near 

sea level to 900 m in elevation (Stevens and others 

2001). The species occurs on pinelands in Florida 

and grows better on near-neutral soils than acid 

(pH 4.7) soils (Long and Lakela 1976). Coffee 

senna grows in moist areas (mean annual rainfall 

from about 1000 to 2200 mm) in Puerto Rico. 

Reproduction.—Coffee senna flowers and fruits 

throughout the year or seasonally, depending on 

rainfall and cold seasons. Seeds collected in 

Puerto Rico averaged 0.0158 + 0.0002 g/seed or 

63,000 seeds/kg. Some 95 percent of scarified 

seeds sown in potting mix germinated between 5 

and 36 days after sowing (author’s observation). 

Scarification is necessary for good seed 

germination (Institute of Food and Agricultural 

Sciences 2002). Mechanical scarification, acid 

treatment, and immersing in boiling water all 

worked well, giving 82 to 100 percent germination 

(Demel and Teketay 1996). The seeds are 
 

700 

dispersed by grazing animals (Sánchez and 

Uranga 1993). 

Growth and Management.—In seasonally cold 

or dry climates, the life cycle of coffee senna is 

complete in 6 to 9 months, but in warm, 

continually moist areas plants may last a full year 

or grow through the second year. Perhaps in Brazil 

where extraordinary heights are reached, the 

species may live a third or fourth year. Growth is 

moderately rapid. Plants add 0.5 to 1.5 m during 

the first season. Although coffee senna is planted 

to yield medicinal materials (Barbadine 2002), 

because of its short life and weedy potential, it is 

not advisable to plant it in wildlands. The species 

can be controlled with broadleaf herbicides (Henty 

and Prichard 1975). 

Benefits and Detriments.—As the name implies, 

the seeds of coffee senna are roasted and used as a 

coffee substitute (Guzmán 1975). The plant’s 

tissues contain a host of phytoactive chemicals 

that may support its numerous applications in folk 

medicine. Extracts or powdered leaves are used as 

an analgesic, antibacterial, anti-hepatotoxic, 

antifungal, anti-inflammatory, antiseptic, 

antispasmodic, antiparasitic, antiviral, 

carminative, diaphoretic, emmenagogue, 

febrifuge, insecticidal, immunostimulant, laxative, 

purgative, sudorific, and vermifuge. Several of 

these effects have been demonstrated in laboratory 

and clinical tests (Raintree 2002). While coffee 

senna can be a weed of cultivated fields and 

plantations, it is principally a problem because it 

accumulates in heavily-grazed pastures (Henty 

and Pritchard 1975). The foliage is poisonous and 

generally avoided by livestock (Stevens and others 

2001). Ingestion of large amounts of seeds has 

been implicated in deaths of cows, horses, and 

goats (Raintree 2002). Poisoning of pigs fed 

coffee senna seeds resulted in muscle necrosis 

(Timm and Riet-Correa 1997). 

References 

Barbadine. 2002. Senna occidentalis (L.). 

Barbadine, Reunion Island, France. http://www. 

barbadine.com/pages/senna_occ_lien.htm. 2 p. 

Demel, T. and D. Teketay. 1996. The effect of 

different pre-sowing seed treatments, 

temperature, and light on the germination of five 

Senna species from Ethiopia. New Forests 

11(2): 155-171. 

Guzmán, D.J. 1975. Especies útiles de la flora 

salvadoreña. Ministerio de Educación, Dirección 

de Publicaciones, San Salvador, El Salvador. 

703 p. 

Haselwood, E.L. and G.G. Motter, eds. 1966. 

Handbook of Hawaiian weeds. Experiment 

Station, Hawaiian Sugar Planters’ Association, 

Honolulu, HI. 479 p. 

Henty, E.E. and G.H. Pritchard. 1975. Weeds of 

New Guinea and their control. Botany Bulletin 

7. Division of Botany, Department of Forests, 

Lae, Papua New Guinea. 189 p. 

Holm, L., J. Doll, E. Holm, J. Pancho, and J. 

Herberger. 1997. World weeds. John Wiley & 

Sons, New York. 1,129 p. 

Institute of Food and Agricultural Sciences. 2002. 

Coffee senna, Cassia occidentalis L. Institute of 

Food and Agricultural Sciences, Cooperative 

Extension Service, University of Florida, 

Gainesville, FL. http://edis.ifas.ufl.edu/ 

BOFY_FW008. 3 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Natural Resources Conservation Service. 2002. 

Plant profile: Senna P. Mill, senna. U.S. 

Department of Agriculture, Natural Resources 

Conservation Service, Washington, DC. 

http://plants.usda.gov/cgi_bin/plant_profile.chi?

symbol=SENNA. 6 p. 

Rain-tree. 2002. Fedegoso. http://rain-tree.com/ 

fedegosa.htm. 7 p. 

Sánchez, P. and H. Uranga. 1993. Plantas 

indeseables de importancia economica en los 

cultivos tropicales. Editorial Científico-Técnica, 

Havana, Cuba. 166 p. 
 

701

Stevens, W.D., C. Ulloa-U., A. Pool, and O.H. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs of Systematic Botany Vol. 85, No. 

1. Missouri Botanical Garden Press, St. Louis, 

MO. p. 1-943. 

Timm, C.D. and F. Riet-Correa. 1997. Plants toxic 

to pigs. Ciencia Rural 27(3): 521-528. 
 

702 

Senna polyphylla (Jacq.) Irwin & Barneby desert cassia 

FABACEAE 

Synonyms: Cassia polyphylla Jacq. 

 Peiranisia polyphylla (Jacq.) Britt. & Rose 

John K. Francis 

General Description.—Desert cassia, also known 

in Spanish as hediondilla and retama prieta, is a 

shrub or occasionally a small tree of dry and moist 

forests of the middle Caribbean, now cultivated as 

an ornamental. It is usually 2 or 3 m in height, but 

rarely reaches 5 m in height and 12 cm in trunk 

diameter. The bark of older stems is dark gray and 

furrowed with short scaly plates. The inner bark is 

light brown. Multiple stems sprouting from the 

root crown are common, even in small plants. The 

stems are mostly clean, long and wand-like. 

Desert cassia develops a root system with a strong 

taproot. The twigs are slender, warty, and light 

green, maturing to brown. The leaves are alternate 

or clustered three to five at the nodes. These 

pinnately compound leaves have three to 15 pairs 

of leaflets that are 4 to 10 mm long. The fivepetaled flowers are yellow and about 3.8 cm 

across, grouped in axillary racemes of mostly two 

flowers. The legume is linear, 8 to 15 cm long, 

slightly contorted, flattened between the seeds, 

and dark brown at maturity. The seeds are round, 

flattened, and dark brown (Howard 1988, Liogier 

1988, Little and others 1974). 

Range.—Senna polyphylla var. polyphylla Irwin 

& Barneby is native to Puerto Rico and the Virgin 

Islands and has been recorded under cultivation in 

Florida, Grenada, Guyana, Surinam, Brazil, and 

probably elsewhere. The variety montis-christi

Irwin & Barneby is an endemic of Hispaniola, and 

the variety neglecta Irwin & Barneby is found 

only on the island of Anegada, British Virgin 

Islands (Liogier 1988). 

Ecology.—In Puerto Rico, desert cassia grows on 

a wide variety of well-drained soils that have 

developed over igneous, metamorphic (including 

ultramafics), and sedimentary (including 

limestone) rocks. It is tolerant of salt spray 

(Botanics Wholesale 2001). The species occurs at 

elevations from near sea level to about 300 m 

above sea level (Little and others 1974). Annual 

rainfall ranges from about 750 mm to about 1200 

mm. Because desert cassia is intolerant of shade 

and grows slowly, it is usually found in areas 

where competition is minimal—dry, very rocky, 

excessively drained, overgrazed, and sites subject 

to occasional fires. Bare ground is probably 

necessary for reproduction. 

Reproduction.—Desert cassia flowers and fruits 

throughout the year (Little and others 1974). The 

seeds from a Puerto Rican collection averaged 

20,400 seeds/kg. Placed on moist filter paper 

without any pre-treatment, they began germinating 

in 2 days and gave 59 percent germination 

(Francis and Rodríguez 1993). The principal mode 

of dispersal today is by livestock. Seedlings in the 

wild are scattered and not abundant. 

Growth and Management.—Growth is slow. 

About 30 cm of annual height increase is normal 

in Puerto Rico. Diameter growth ranges between 

about 1 and 4 mm/yr. Although no planting 

experience is published, plantations could 

probably be established with containerized 

seedlings followed by several years of weeding 

until the seedlings grew above herb and grass 

competition. 
 

703

Benefits.—Desert cassia is grown by commercial 

nurseries and widely planted as a flowering 

ornamental in the New World Tropics. It is 

employed as an accent plant or trained into a small 

background tree. The wood is light brown and 

hard and used occasionally for fence posts and 

fuel. Cows apparently do not readily eat the 

foliage of desert cassia—the species is more 

common in heavily grazed rangeland than 

elsewhere. 

References 

Botanics Wholesale. 2001. http://www.botanics. 

com/Products/botanics_details.asp?NameAssoc

+153.htm. 1 p. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University. Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 
 

704 

Serenoa repens (Bartr.) Small saw palmetto 

PALMAE 

Synonyms: None 

Deborah K. Kennard, G.W. Andrews, Kenneth Outcalt, and Mary Carrington 

General Description.—Saw palmetto, also 

known as palmetto, is a low-growing palm 

endemic to the Southeastern United States. Saw 

palmetto stems typically lie prostrate at the soil 

surface but can grow upright and reach heights of 

5 to 7 m (Tanner and others 1996). Multiple, 

persistent, palmate leaves up to 1 m wide emerge 

from the stem’s terminal buds. Short recurved 

spines line the petioles (Small 1933). Flowers are 

perfect and borne on paniculate inflorescences that 

emerge from the leaf bases. The fruit is a oneseeded ellipsoid or subglobose drupe with a fleshy 

mesocarp, 16 to 25 mm long and 12 to 19 mm 

wide. Fruit color turns from green, to yellow, to 

orange, and then to bluish-black when fully ripe 

(Hilmon 1968).

 

Range.—Saw palmetto is endemic to the Coastal 

Plain of the Southeastern United States. The 

northern limits of its range extend from 

southeastern Louisiana through Tifton Georgia to 

Charleston County, South Carolina (Hilmon 1968, 

McNab and Edwards 1980). 

Ecology.—Saw palmetto occurs as a major 

understory plant in seasonally wet pine flatwoods, 

well-drained scrubby flatwoods, and on sandy 

berms and dunes along rivers and the coast 

(Tanner and others 1996). Within these habitats, it 

can grow in a variety of conditions from shade to 

full sun (Small 1926). Common associates include: 

gallberry [Ilex glabra (L.) Gray], wax myrtle 

(Myrica cerifera L.), staggerbush [Lyonia 

ferruginea (Walter) Nuttall], oaks (Quercus spp. 

L.), wiregrass (Aristida beyrichiana Trin. & 

Rupr.), several bluestem species (Andropogon spp. 

L.), and lop-sided Indian grass [Sorgastrum 

secundum (Ell.) Nash.] (Tanner and others 1996). 

Saw palmetto occurs on a wide range of soil types 

but most commonly is found on seasonally 

flooded, sandy, acidic spodosols typical of 

flatwoods ecosystems in the Lower Coastal Plain. 

Typical soil types are Leon fine sand, Myakka fine 

sand, and Immokalee fine sand. Saw palmetto also 

grows on deep, sandy entisols, on calcareous 

sandy soils near coasts, and on limestone in 

southern Florida. Annual rainfall varies from 114 

cm in the northern part of its range to over 150 cm 

along the southeastern coast of Florida. Rainfall 

also becomes more seasonal going from north to 

south, with increasingly more rainfall occurring 

during summer. In southern Florida, the 

southernmost part of its range, 64 percent of 

average annual rainfall occurs from June to 

September (Hilmon 1968). 

Reproduction.—Saw palmettos primarily 

reproduce vegetatively through suckers from the 

main stem (Fisher and Tomlinson 1973). Each 

sucker has the capacity to become a ramet, 

forming extensive clumps of genetically identical 

clones. Saw palmettos must be at least 0.6 m in 

height to flower (Carrington and others 2000) after 

which probability of flowering increases with plant 

height. Inflorescences emerge from buds at the 

bases of previous season’s leaves in February to 

April, and flowering occurs from April to June 

(Hilmon 1968). Flowers are insect pollinated 

(Tanner and others 1996). While 34 insect species 

are known to pollinate saw palmetto flowers, the 

European honeybee (Apis mellifera L.) is the 

primary pollinator (Carrington and others in 

review). Emerging saw palmetto inflorescences are 

subject to attack by cabbage palm caterpillars 

(Litoprosopus futilis G. & R.), and green fruits are 

subject to anthracnose [Colletotrichum

gloeosporioides (Penz.) Penz. & Sacc. in Penz.] 
 

705

(Carrington and others 2000). Fruits ripen in 

August to November. Saw palmettos flower 

heavily about every 2 to 4 years. During these 

years, individual ramets commonly produce two to 

three inflorescences. Due to high flower density 

(several thousand flowers/inflorescence), typically 

0.4 to 0.5 kg of fruit are produced on each 

inflorescence. In some cases, individual 

inflorescences can produce up to 12 kg of fruit. 

Average fruit yield for a site is approximately 200 

kg/ha; however, yields can vary from less than 100 

kg/ha to more than 1,500 kg/ha (Carrington and 

others 1997). Fruits are bird and mammal 

dispersed (Tanner and others 1996). Seed 

germination ranges from 20 percent after 15 

months in field conditions to 55 percent after 6 

months under lab conditions (Hilmon 1968). Seeds 

can remain viable for up to 1 year, albeit with 

reduced germination rates (Carrington and others 

2000). Seed germination may be enhanced by 

passing through animal digestive systems (Tanner 

and others 1996). 

Growth and Management.—Growth of saw 

palmetto is slow, ranging between 0.6 and 2.2 

cm/year in stem elongation (Abrahamson 1995, 

Hilmon 1968). Based on these estimates, some 

saw palmettos may be 500 to 700 years old 

(Abrahamson 1995, Tanner and others 1996). In 

some cases, land managers may want to reduce the 

density and cover of saw palmetto in order to 

improve cattle forage and/or timber production or 

to reduce fire hazard. Because saw palmetto is 

fire-tolerant, mechanical or chemical treatments 

are often needed for control. Roller-chopping 

reduces abundance of saw palmetto (Hilmon and 

others 1963, Lewis 1970) by crushing vegetative 

parts and removing terminal buds. This treatment 

is particularly effective during periods of low soil 

moisture (Moore 1974) or when followed by 

prescribed burning. Among herbicide treatments, 

repeated cycles of 2,4,5-T followed by burning 

will reduce palmetto cover (Altobellis and Hough 

1968), but this chemical is no longer available. 

Metsulfuron, often sold as Escort, is the most 

effective herbicide currently available for control 

of saw palmetto (personal communication J.L. 

Michael, Southern Research Station, Auburn, AL). 

In other cases, land managers may want to 

enhance saw palmetto flowering and fruiting. The 

most cost-efficient practice to increase fruit 

production is prescribed burning (Carrington and 

others 2000). Optimal burning frequency is every 

5 to 8 years. Soil fertilization has been used to 

increase coverage of saw palmetto (Carrington and 

others 2000). 

Fire.—Saw palmetto naturally occurs in plant 

communities adapted to periodic burning. Saw 

palmetto itself is highly flammable due to the 

accumulation of dead fronds that remain on the 

plants for several years (Carrington and others 

2000) and volatile waxes that cover green leaves. 

Fires commonly consume all above-ground foliage 

of saw palmettos, and therefore, prescribed fire is 

used to reduce leaf coverage for short periods. 

However, leaf growth usually begins a few days 

after fires, and saw palmettos can regain 80 

percent of crown coverage the first year after 

burning (Hilmon 1968). Fire also benefits fruit 

production of saw palmetto, especially within the 

first year following burns (Carrington and others 

2000). However, flowering may be reduced for 

several years after burns (Abrahamson 1999) as 

palmettos recover from both burning and heavy 

fruit production. Frequent burning (every 1 to 4 

years) curtails flowering and fruiting by keeping 

carbohydrate reserves low (Hilmon 1968). 

Detriments and Benefits.—Saw palmetto was 

viewed by early settlers in the Southeastern United 

States as an obstacle to establishing agricultural 

fields, cattle pastures, and home sites. Ranchers 

continue to regard it as a competitor of native 

forage grasses, and foresters cite that it inhibits 

pine regeneration. Despite these detriments, saw 

palmetto is viewed as a significant pharmaceutical 

resource in Florida and southern Georgia due to 

the effectiveness of certain fruit compounds (such 

as free fatty acids, phytosterols) in treating benign 

prostatic hyperplasia (Tasca 1985, Braeckman 

1994, Wilt and others 1998), or swelling of the 

prostate gland. In 1995, saw palmetto fruits 

collected for pharmaceuticals sold for over $6/kg. 

Total estimated value of fruits sold in 1996 was 

approximately $5 million (Carrington and others 

2000). In addition to its medicinal value, saw 

palmetto can be a significant source of honey 

production (Bennett and Hicklin 1998). Saw 

palmetto is also a good native plant for 

enviroscaping due to its natural drought- and 

insect-resistance, and low requirements for 

fertilization (Tanner and others 1996). Saw 

palmettos serve as nesting and denning habitat for 

over 100 animal species, including: the 

endangered Florida grasshopper sparrow 

(Ammodramus savannarum Gmein), Florida 

panther (Felis concolor Linnaeus), Florida 
 

706 

woodrat (Neotoma floridana Ord), wild turkey 

(Meleagris gallopavo Linnaeus), and white-tailed 

deer (Odocoileus virginianus Zimmermann) 

(Tanner and others 1996). Fruits are eaten by black 

bears (Ursus americanus Pallas), white-tailed deer 

(Odocoileus virginianus), raccoons (Procyon lotor 

Linnaeus), turkeys (Meleagris gallopavo), bobwhite quail (Colinus virginianus L.), gray foxes 

(Urocyon cinereoargenteus Schreber), opossum 

(Dasypus novemcinctus Linnaeus), and gopher 

tortoises (Gopherus polyphemus Daudin) (Maehr 

and Layne 1996). 

References 

Abrahamson, W.G. 1995. Habitat distribution and 

competitive neighborhoods of two Florida 

palmettos. Bulletin of the Torrey Botanical 

Club 122: 1-14. 

Abrahamson, W.G. 1999. Episodic reproduction in 

two fire-prone palms, Serenoa repens and 

Sabal etonia (Palmae). Ecology 80:100-115. 

Altobellis, A.T. and W.A. Hough. 1968. 

Controlling palmetto with fire and herbicides. 

Georgia Forestry Research Paper 52. Georgia 

Forest Research Council, Macon, Ga. 

Bennett, B.C. and J.R. Hicklin. 1998. Uses of saw 

palmetto (Serenoa repens, Arecaceae) in 

Florida Economic Botany 52: 381-393. 

Braeckman, J. 1994. The extract of Serenoa repens

in the treatment of benign prostatic 

hyperplasia: A multicenter open study. 

Current Therapy Research 55: 776-785. 

Carrington, M.E., T.D. Gottfried, and J.J. 

Mullahey. (In review) Pollination biology of 

saw palmetto (Serenoa repens: Palmae) in 

southwestern Florida. Palms. 

Carrington, M.E., J.J. Mullahey, G. Krewer, B. 

Boland, and J. Affolter. 2000. Saw palmetto 

(Serenoa repens): an emerging forest resource 

in the southeastern United States. Southern 

Journal of Applied Forestry 24(3): 129-134. 

Carrington, M.E., J.J. Mullahey, and F. Roka. 

1997. Saw palmetto: A fountain of youth. 

Proceedings American Forage Grassland 

Council 6: 233-237. 

Fisher, J.B. and P.B. Tomlinson. 1973. Branch 

and inflorescence production in saw palmetto 

(Serenoa repens). Principes 17: 10-19. 

Hilmon, J.B. 1968. Autecology of saw palmetto 

[Serenoa repens (Bartr.) Small]. Ph.D. 

dissertation. Duke University, Durham, N.C. 

191 p. 

Hilmon, J.B., C.E. Lewis, and J.E. Bethune. 1963. 

Highlights of recent results of range research 

in Southern Florida. Society of American 

Foresters Proceedings 1962: 73-76. 

Lewis C.E. 1970. Responses to chopping and rock 

phosphate on south Florida ranges. Journal of 

Range Management 23: 276-282. 

Maehr, D.S. and J.N. Layne. 1996. Florida’s allpurpose plant the saw palmetto. Palmetto 

(Fall): 16 –10, 15, 21. 

McNab, W.H. and M.B. Edwards. 1980. Climatic 

factors related to the range of saw-palmetto 

[Serenoa repens (Bartr.) Small]. American 

Midland Naturalist 103: 205-208. 

Moore, W. H. 1974. Some effects of chopping 

saw-palmetto pineland threeawn range in 

south Florida. Journal of Range Management 

27(2): 101-104. 

Small, J.K. 1926. The saw-palmetto–Serenoa 

repens. Journal of the New York Botanical 

Garden 27: 193-202. 

Small, J.K. 1933. Manual of the southeastern 

flora. John Kunkel Small, New York. 

1,554 p. 

Tanner, G.W., J.J. Mullahey, and D. Maehr. 1996. 

Saw-palmetto: an ecologically and 

economically important native palm. Circular 

WEC-109, Florida Cooperative Extension 

Service, Institute of Food and Agricultural 

Sciences, University of Florida, Gainesville, 

FL. 7 p. 

Tasca, A. 1985. Treatment of obstructive 

symptomatology in prostatic adenoma with an 

extract of Serenoa repens. Minerva Urologica 

e Nefrologica 37: 87-91. 
 

707

Wilt, T., A. Ishani, and C. Mulrow. 1998. Saw 

palmetto extracts for treatment of benign 

prostatic hyperplasia: a systemtatic review. 

Journal of the American Medical Association 

280: 1,604-1,609. 
 

708 

Serjania polyphylla (L.) Radlk. bejuco de corrales 

SAPINDACEAE 

Synonyms: Paullinia polyphylla L. 

 Paullinia triternata Jacq. 

 Serjania lucida Schum. 

 Serjania triternata (Jacq.) Willd. 

 Serjania dubia Spreng. 

John K. Francis 

General Description.—Bejuco de corrales, also 

known as bejuco de canastas, bejuco de costillas, 

basket wood, and blackwith, is a woody vine that 

ascends as much as 20 m into the crowns of trees. 

Bejuco de corrales is a Spanish name meaning 

corral vine. The older stems are brown, rough, 

with three main striations and up to 5 cm in 

diameter. The stem cross section has a 

characteristic pattern reminiscent of a string of 

beads around a central core. The vines have a 

taproot, extensive lateral, and abundant fine roots, 

and all have a dark brown color. Periodic 

sprouting from the root crown replaces dying 

stems and gives rise to multiple stems, usually less 

than five. The branches are slender and flexible 

with few bifurcations. Tendrils, which support the 

vine on trees and other structures, grow at the leaf 

axils. The alternate compound leaves divide by 

threes. Leaflets, which number three to 19 or 

more, are dentate from the middle to the tip, 1.5 to 

6.0 cm long and 1.0 to 3.5 cm broad. 

Inflorescences are axillary panicles and contain 

many white flowers. The brown or red-brown 

fruits are samaras, 1.5 to 2.5 cm long, that grow in 

groups of three. Each samara contains a single 

seed (Acevedo-Rodrígues 1985, Liogier 1994). 

Range.—Bejuco de corrales is native to Puerto 

Rico and offshore islands, the Virgin Islands, and 

Hispaniola (Liogier 1994). It is not known to have 

been planted elsewhere. 

Ecology.—Bejuco de corrales grows in remnant 

and secondary forests, on roadsides, fence rows, 

river banks, and brushy pastures. It is usually 

common but rarely abundant. The species is 

moderately intolerant of shade, requiring full 

sunlight to flower and fruit; yet it can survive in 

the understory of low basal-area forest. Bejuco de 

corrales grows on soils with textures from sand to 

clay that are derived from igneous and 

sedimentary (including limestone) rocks. In Puerto 

Rico, the species grows from near sea level to over 

600 m in elevation and in areas that receive from 

750 to over 2000 mm of annual rainfall. In the 

areas with the highest rainfall, it is confined to 

excessively drained sites. Bejuco de corrales 

tolerates drought and at least mild salt spray. 

Frosts are not known within the native range. 

Reproduction.—Bejuco de corrales flowers and 

fruits from November through June (AcevedoRodrígues 1985). An infrutescence may contain 

25 or more fruits, and a plant may produce several 

infrutescences per year. A collection of bejuco de 

corrales samaras averaged 0.0248 g/fruit or 40,000 

fruits/kg. When the clusters break up, the samaras 
 

709

spin and fly sideways as they descend and may 

travel a considerable distance before reaching the 

ground. 

Growth and Management.—Seedlings grow 

about 0.5 m in their first year. Older plants and 

sprouts grow 2 or 3 m each year. Individual stems 

can live for several years; by sprouting, plants 

may survive for at least several decades. No 

planting experience for the species has been 

reported. In the past, farmers controlled bejuco de 

corrales by repeated cutting with a machete. These 

vines could probably be killed faster by spraying 

the sprouts that arise after cutting with broadleaf 

weed killer. 

Benefits.—The strong and flexible stems of 

bejuco de corrales are suitable for basketry, and 

some of the common names indicate that it was 

used for this purpose. Bejuco de corrales is 

considered to be a honey plant in the Dominican 

Republic (Marcano Fondeur 1973). Livestock eat 

young plants and the foliage of older ones that are 

within their reach. Bejuco de corrales is reputed to 

be a diuretic and purifier of the blood in herbal 

medicine (Liogier 1990). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc. San Juan, PR. 566 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Marcano Fondeur, E. de J. 1973. La flora apícola 

de la República Dominicana. http:// 

marcano.freeservers.com/nature/estudios/apicola

/dicotsp.htm. 11 p. 
 

710 

Sesbania sericea (Willd.) Link silky sesban 

FABACEAE 

Synonyms: Coronilla sericea Willd. 

John K. Francis 

General Description.—Silky sesban, also known 

as papagayo in Spanish, is a short-lived woody 

shrub of frequently disturbed areas. Its mature 

height may vary from 1 to 6 m with basal 

diameters reaching as much as 10 cm under the 

most favorable conditions. A single stem is 

usually developed unless the plant sprouts after 

damage. Robust low side branches develop on 

open-grown individuals. Secondary branches on 

stems are usually unbranched. The species is 

unarmed or may occasionally have weak prickles. 

Stems are green when young, turning gray. There 

may or may not be a well-developed taproot, 

depending on the aerated rooting depth. The white 

lateral roots are relatively thick (3 to 5 mm) and 

support many nodules with reddish centers and 

abundant fine roots. The pinnately compound 

leaves are 5 to 25 cm long and support 10 to 20 

leaflet pairs. The leaflets are oblong, rounded at 

both ends, mucronate at the apex, with a silky 

pubescence below. The racemes have two to eight 

yellow, greenish-yellow, or orange flowers. The 

brown pods that develop after flowering are 10 to 

20 cm long, 3 mm broad, and contain 20 to 30 

seeds (Howard 1988, Liogier 1988, Nelson 1996). 

The species has 2n = 24 chromosomes (Long and 

Lakela 1976). 

Range.—There is confusion about the native 

range of silky sesban. Howard (1988) states that it 

is a native of Sri Lanka. Others refer to tropical 

Asia (Long and Lakela 1976) or Africa and the 

Caribbean (Evans and Rotar 1987) as the native 

ranges. It is clear that the species has been widely 

introduced. The current New World range covers 

southern Florida, the Bahamas, the West Indies, 

Trinidad, Surinam, the Guianas, Venezuela, and 

Central America (Liogier 1988). 

Ecology.—Silky sesban is highly intolerant of 

shade and must have disturbed soil in which to 

germinate. Once established, it competes 

aggressively with grass and herbs, but it must 

dominate all its life or die. Stands of silky sesban 

vigorously self-thin. The species develops best on 

moist alluvial soils. It will grow on soils with low 

fertility, particularly when poorly or very poorly 

drained. Modest amounts of salts in the soil are 

tolerated. Apparently, the northern extent of the 

range is limited by cool weather and frost. 

Reproduction.—A collection of 45 air-dry pods 

of silky sesban weighed an average of 0.378 +

0.0008 g/pod. The seeds separated from them 

averaged 0.0053 + 0.0001 g each or 189,000 

seeds/kg. Sown without any pretreatment on 

commercial potting mix, these seeds germinated at 

99 percent between 3 and 24 days after sowing. 

Germination is epigeal. Although the original 

mode of seed dispersal has not been documented, 

in the Caribbean it seems to be opportunistic, 

disbursing by wind, water, grazing animals, and 

farm machinery. Seedlings may be abundant in the 

presence of a seed source on moist bare soil. 
 

711

However, relatively few of the seedlings progress 

past the early seedling stage. 

Growth and Management.—Plants of silky 

sesban grow rapidly. Seedlings are about 10 cm 

tall at the end of 1 month and reach 2 to 6 m in 

height in about 6 months. Silky sesban was 

reported to yield 26.8 Mg/ha (fresh weight) in a 

84-day growing period (Evans and Rotar 1987). 

After flowering, height growth of silky sesban 

ceases. Plants live from about 8 months to a little 

over a year, depending on conditions. When their 

seeds mature, the plants die. The species may 

grow and die in an annual cycle timed with wet 

and dry seasons, or it may simply grow in 

response to favorable soil conditions as they 

occur. If land managers find it necessary to 

establish silky sesban, sowing into a moist, 

prepared seedbed should be sufficient. 

Benefits.—The species of Sesbania, including 

silky sesban, are used as green manure in India. 

Silky sesban is readily consumed by cattle in 

pastures. The closely related S. sesban (L.) Merr. 

has been shown to have moderately good 

nutritional values. Some of the species of Sesbania

have toxic saponins and canavanine in their seeds. 

So far, the seeds of silky sesban have not been 

reported to be toxic. Crushed seeds of this species 

were fed to chicks at 1 percent of body weight 

each day for 3 days without any signs of toxicity 

(Evans and Rotar 1987). Silky sesban can be 

weedy but usually causes few problems. It was 

experimentally intercropped with maize and 

resulted in no depression in maize yield. In 

Guyana, the species grows in upland rice fields 

but does not occur in flooded rice paddies (Evans 

and Rotar 1987). The stemwood of silky sesban is 

brash, has a relatively low specific gravity, and 

offers few prospects for commercial use. 

References 

Evans, D.O. and P.P. Rotar. 1987. Sesbania in 

agriculture. Tropical Agriculture Series 8. 

Westview Press, Boulder, CO. 192 p. 

Howard, R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University. Jamaica Plain, 

MA. 673 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR 481 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc. Sarasota, FL. 

391 p. 
 

712 

Sida fallax Walp 'ilima

MALVACEAE 

Synonyms: Anoda ovata Meyen 

 Sida cordifolia L. 

 Sida dielli Gray, 

 Sida fallax var. kauaiensis Hochr. 

 Sida ledyardii St. John 

Sarah A. Taylor and Randy S. Senock 

General Description.―Sida fallax, known as 

'ilima, is an erect to prostrate perennial herb or 

subshrub that grows to 0.2 to 1.5 m tall. The shrub 

varies from smooth to densely covered in woolly 

hairs, the hairs usually stellate. 'Ilima has simple 

leaves, the blades from 1 to 12 cm long, 

lanceolate-ovate to oblong or elliptic. The upper 

surfaces are typically bright green and smooth, or 

sparsely to densely pubescent, the lower surface 

densely pubescent. Flowers arise from an axil, 

solitary or two to seven per node born in the 

pedicel, at the end branches or in the leaves. 

Corolla is usually yellow to orange-yellow, 

sometimes dark maroon at the base. Fruits split 

into two or more carpels at maturity. Extensive 

variation in the morphology of this species exists 

in leaf size and shape, stature, pubescence, and 

arrangement of its inflorescence (Wagner and 

others 1990). 

 

Range.―Sida is a genus of 125 to 150 species of 

tropical and subtropical regions of the world, from 

the Pacific Islands to Hawaii. 'Ilima has a series of 

similar forms common to all islands but also 

exhibits morphological variation from island to 

island. The shrub exists on all the main islands of 

Hawaii, including the atoll Midway and the 

pinnacle Nihoa from coastal to subalpine 

environments. 

Ecology.―'Ilima displays numerous ecological 

types, occurring from sea level to altitudes of 600 

m. Beach forms are low growing, dense, spreading 

mats, while upland types may grow erect and 

shrub-like (Krauss 1998). Lowland types are 

typically densely pubescent, while upland types 

are glabrous. 'Ilima occurs on a variety of 

substrate, from limestone reefs, dry lava fields, and 

along sandy and rocky shores. Often 'ilima 

represents the only ground cover, aside from 

nonnative species, on arid, rocky coastlines. 'Ilima 

appears intolerant of wildfire as adult plants 

typically do not resprout within burned areas. In 

montane dry shrublands or leeward Hawaii, 'ilima 

re-established from seed within 4 years of burning 

(Sherry and others 1999). 

Reproduction.―The flower parts are typically in 

fours or fives and are true flowers with stamen and 

pistils. The ovules are covered by ovaries. Seeds 

are brown or black, obovoid, with three angles, 1.7 

to 2.2 mm long, minutely pubescent around the 

raphe. Somatic chromosome number of 2n = 28 

has been identified (Wagner and others 1990). It is 

probably wind pollinated, though the native yellow 

faced bee species Hylaeus has been observed 

visiting the flowers of ‘ilima (Anonymous 2002). 

Growth and Management.―Easily grown from 

seeds, 'ilima makes an excellent xeriscape plant. 

Seedlings germinate and grow quickly, while 

cuttings do less well. It requires full sunlight and 

light to moderate watering. Upland varieties can be 

susceptible to nematodes, while lowland types are 

resistant. Leaves may become yellow spotted and 

chlorotic, and will respond well to fertilizer with 

micronutrients (Krauss 1998). It does well in 

sandy or rocky soils. 

Benefits.―'Ilima is an important plant to Hawaii 

and its people. Four separate types of 'ilima were 
 

713

recognized. These included 'ilima-ku-kala, ilimalei, 'ilima-ku-kahakai and 'ilima-koli-kukui 

(Stephens 2000). 'Ilima-ku-kala is the wild tall 

form found in the mountains. 'Ilima-ku-kahakai is 

the creeping form found along the coastline. 

'Ilima-lei were cultivated by the Hawaiians for use 

in lei making. 'Ilima-koli-kukui was another 

cultivar with reddish brown flowers (Stephens 

2000). 'Ilima is considered a special flower of the 

island of Oahu. 'Ilima leis are among the most 

treasured leis and are difficult to make, requiring 

over a thousand blossoms. At one time, only 

royalty were allowed to wear an 'ilima lei 

(Anonymous 2001). 'Ilima was also used 

medicinally. The juice squeezed from the flowers 

was used as a mild laxative for children, and the 

root bark mixed with flowers was used to treat 

asthma (Krauss 2000). Buds were chewed to 

quench thirst on hot, dry days. Stems were used to 

make baskets. 'Ilima is said to be one of the forms 

that Laka, the goddess of hula, could take at will 

(Quensell 2000). 

Detrimental Effects.―‘Ilima is known to be host 

to nonnative rust, Puccinia heterospora, which is 

now becoming established on native plants. 

Infection may be heavy and conspicuous. 

References 

Anonymous. 2001. Hawaii's Island symbols. 

http://www.geobop.com/World/NA/US/HI/Islan

d.htm [not paged]. 

Anonymous. 2002. ttp://www.anglefire.com/hi4/ 

nhps/news122 k. html [not paged]. 

Krauss, B. 1998. How to plant a native Hawaiian 

garden. http://hawaii.gov/health/ 

oeqc/garden/index.html 

Krauss, B. 2000. Native plants used as medicine in 

Hawaii. http://library.kcc.hawaii.edu/~ 

soma/krauss/ilima.html. [not paged] 

Sherry, K., J.M. Castillo, and R.B Shaw. 1999. 

Effects of wildfire on vegetation and rare plants 

in arid montane shrubland. Pohakaloa Training 

Area, Hawaii. 1999 Hawaii Conservation 

Conference, Honolulu, HI. 

Stephens, M. 2000. The comparative ecophsiology 

of mountain and coastal populations of Sida 

fallax Walp. (Malvaceae). M.S. thesis. 

University of Hawaii, Hilo, HI. 

Quensell, N. 2000. Native Hawaiian plants. 

http://www.kcc.hawaii.edu/campus/tour/plants/p

ilima.htm [not paged] 

Wagner, W.L., D.R. Herbst, and S.H. Sohmer. 

1990. Manual of the Flowering Plants of 

Hawaii. University of Hawaii Press, Honolulu, 

HI. p. 897-898. 
 

714 

Sida rhombifolia L. arrowleaf sida 

MALVACEAE 

Synonyms: Sida hondensis Kunth in Humb. 

 Sida ruderata Macfad. 

John K. Francis 

General Description.—Arrowleaf sida is known 

by many common names: broom weed, paddy’s 

lucerne, Cuba jute, common sida, escoba dulce, 

escoba blanca, escoba dura, malva de cochino, 

huang hau mu, rhomboid ilima, antibala, idem, 

afata, guanxuma, and basbasot (Holm and others 

1997). It is a short-lived perennial subshrub 

(woody stem and herbaceous branches) commonly 

growing to 60 cm, but sometimes reaching 1.5 m 

in height. (Holm and others 1997, Howard 1989, 

Liogier 1994). Arrowleaf sida develops a taproot 

and many lateral and fine roots. The plant usually 

has a single stem unless disturbed, but may branch 

near the ground. The stems are woody, flexible, 

and tough, and may reach 1 cm or more in basal 

diameter. The twigs are slender, green, and 

semiwoody. The alternate leaves are variable in 

both shape and size. They have a 3- to 8-mm 

petiole with broadly ovate to lanciolate (often 

rhomboidal) blades 2 to 6 cm long and serrate at 

the margins, especially from the middle to the tip. 

The axillary flowers are solitary on slender 

pedicels 2 to 3 cm long. The corolla contains five 

pale yellow to yellow-orange petals. The fruits are 

round, flattened schizocarps, 4 to 5 mm in 

diameter and contain 10 to 14 small dark-brown 

seeds. Both diploid (2n = 14) and tetraploid (2n = 

28) forms exist (Holm and others 1997, Howard 

1989, Liogier 1994). 

Range.—Arrowleaf sida grows today in over 70 

countries throughout the tropical, subtropical, and 

warm temperate regions (Holm and others 1997, 

Howard 1989). Its original range is not known, but 

the presence of multiple subspecies and varieties 

seems to indicate that it was from the Old World. 

The species was introduced into the United States 

in the late 1800’s as a promising fiber crop (Holm 

and others 1997). 

Ecology.—Arrowleaf sida grows only in disturbed 

areas and in situations where overhead 

competition is controlled or naturally light. It is 

common in cultivated fields, pastures, abandoned 

farmland, roadsides, vacant lots, construction 

sites, landslides, and river overflow areas. It does 

not survive in tall grass swards, tall brushlands, 

and closed forests. Arrowleaf sida grows on both 

fertile and degraded soils of all textures and 

derived from most parent materials. It may be 

found growing from near sea level to 2,000 m in 

elevation (Holm and others 1997). In Puerto Rico, 

the species grows in areas receiving from 900 to 

3000 mm of precipitation. 

Reproduction.—Arrowleaf sida flowers and 

fruits continuously starting at 3 or 4 months of age 

in Puerto Rico. In Central India, plants flower 

from September to December and fruit from 

October to January (Parrotta 2001). A single plant 

can produce as many as 11,600 seeds. Individual 

seeds have been reported to weigh between 0.0012 

and 0.0015 g (Holm and others 1997). Seeds 

collected in Puerto Rico averaged 0.0016 g/seed or 

625,000 seeds/kg. Sown on moist filter paper 

without any pretreatment, 33 percent germinated 

within 13 months (author’s observation). Fresh 

seed is dormant, and dry storage, freezing, various 

light regimes, and several chemical treatments 
 

715

failed to induce germination. However, acid 

scarification, heating, and cold storage succeeded 

in breaking dormancy. Some 80 percent 

germination was obtained at the best soil depth for 

germination, which was 0.5 to 2 cm (Holm and 

others 1997). Arrowleaf sida plants sprout 

vigorously if cut. The seeds are dispersed by 

water, farm machinery, ruminant animals, and in 

impure agricultural seed. Ants disperse the seeds 

in Africa (Holm and other 1997). 

Growth and Management.—Arrowleaf sida 

plants may reach 0.5 m or more in their first year. 

Growth is most rapid in warm conditions (days of 

30 °C and nights of 25 °C). Growth nearly ceases 

below 20 °C. Plants survive frosts and winters as 

far north as Tennessee (Holm and others 1997). 

Although the species is able to perpetuate itself as 

an annual in difficult climates and under cultivated 

agriculture, the aerial portions of individual plants 

in favorable climates may live 3 years, and 

possibly more by resprouting. Control, but not 

elimination of arrowleaf sida in plantations and 

pastures may be achieved by hand weeding and 

herbicide spraying. Hand pulling and mowing are 

only partially effective because arrowleaf sida is 

difficult to pull and quickly sprouts after cutting. 

The weed problem from arrowleaf sida becomes 

more severe in reduced tillage agriculture (Holm 

and others 1997). Mowing or chain slashing is 

recommended in pastures to suppress arrowleaf 

sida and other unpalatable Sida species and allow 

pasture forage plants to grow (Food and 

Agriculture Organization 2002). Effective 

biological control has been obtained in Australia 

and Papua New Guinea by introducing the leafeating beetle Calligrapha pantherina, which feeds 

exclusively on three species of weedy Sida’s 

(Kuniata and Rapp 2001). 

Benefits.—Arrowleaf sida stems are used as 

rough cordage, sacking, and for making brooms. 

The stems have a high quality fiber and were once 

exported from India and elsewhere as “hemp” 

(Guzmán 1975, Holm and others 1997). Chemical 

analysis revealed that the leaves contain 

respectable amounts of nutrients: 74,000 to 

347,000 ppm protein, 94,000 to 475,000 ppm 

carbohydrates, 33,000 to 167,000 ppm fiber, 

14,000 to 71,000 ppm fat, and 16,000 to 81,000 

ppm ash. However, it was reported that the root 

contained 450 ppm alkaloids and the presence of 

ephedrine and saponin (Southwest School of 

Botanical Medicine 2002). Another source reports 

an alkaloid content in the root of 0.1 percent and 

the presence of choline, pseudoephedrine, betaphenethylamine, vascin, hipaphorine and related 

indole alkaloids (Shaman Australis 

Ethnobotanicals 2002). Perhaps because of these 

chemicals, arrowleaf sida is unpalatable to cattle 

(Kuniata and Rapp 2001). Arrowleaf sida has 

significant medicinal applications for which it is 

cultivated throughout India. The pounded leaves 

are used to relieve swelling, the fruits are used to 

relieve headache, the mucilage is used as an 

emollient, and the root is used to treat rheumatism 

(Parrotta 2001). Australian Aborigines use the 

herb to treat diarrhea. Leaves are smoked in 

Mexico and a tea is prepared in India for the 

stimulation it provides (Shaman Australis 

Ethnobotanicals 2002). 

References 

Food and Agriculture Organization. 2002. Cultural 

practices. http://www.fao.org/ag/AGP/AGPC/ 

doc/Publicat/FAOGUL2/B204.htm. 2 p. 

Guzmán, D.J. 1975. Especies útiles de la flora 

Salvadoreña. Ministerio de Educación, 

Dirección de Publicaciones. San Salvador, El 

Salvador. 703 p. 

Holm, L., J. Doll, E. Holm, J. Pancho, and J. 

Herberger. 1997. World weeds. John Wiley and 

Sons, Inc. New York. 1,129 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Kuniata, L. and G. Rapp. 2001. Biocontrol of Sida 

rhombifolia in Papua New Guinea. http://w 

ww.nt.gov.au/dpif/pubcat/agnotes/ 542.htm. 4 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Parrotta, J.A. 2001. Healing plants of Peninsular 

India. CABI Publishing, Wallingford, UK and 

New York. 917 p. 

Shaman Australis Ethnobotanicals. 2002. Sida 

rhombifolia, common sida. http://www.shamanaustralis.com/Website/Sidarhombifolia.htm. 1 p. 
 

716 

Southwest School of Botanical Medicine. 2002. 

Constituents. http://www.rt66.com/hrbmoore/ 

Constituents/Sida_rhombifolia.txt. 1 p. 
 

717

Smilax domingensis Willd. bejuco de membrillo 

SMILACACEAE 

Synonyms: Smilax caudata Lundell 

 Smilax microscola (B.L. Rob.) Killip & C.V. Morton 

John K. Francis 

General Description.—Bejuco de membrillo, 

also called bejuco de sarzaparilla and zarzaparilla, 

is an evergreen woody vine or scrambling shrub 

that reaches 5 to 20 m of extension and 5 cm of 

basal diameter and supports itself by means of 

tendrils. The roots produce rhizomes with 

occasional tuber-like structures. The plant has a 

round, smooth stem, gray in older parts and 

greenish-yellow in younger portions. The stem 

wood does not have annual rings. Some specimens 

have small spines or prickles in their lower stems. 

Bejuco de membrillo has few branches until it 

reaches increased sunlight and forms a crown. The 

alternate leaves are oval to lanceolate, glabrous, 

and leathery. Leaves have short petioles and entire 

blades 8 to 12 cm long. The inflorescences are 

axillary cymes of minute, greenish, fragrant 

flowers. The fruits are 7 mm in diameter, globose, 

juicy, maturing to dark purple or black, and 

contain one seed each (Acevedo-Rodríguez 1985, 

Stevens and others 2001). The fruits taste slightly 

sour and slightly sweet, somewhat like apples. The 

seeds are shiny tan or light brown and flattened on 

one side (author’s observation). 

Range.—Bejuco de membrillo is native to the 

Greater Antilles, and Mexico to Panama 

(Acevedo-Rodríguez 1985, Stevens and others 

2001). Many herbarium specimens also exist from 

Colombia, Ecuador, and Peru (Missouri Botanical 

Garden 2003). At least one major authority 

(Natural Resources Conservation Service 2003) 

has bejuco de membrillo as conspecific under S. 

smallii Morong that ranges across the Southern 

United States. Time will tell whether this union is 

reliable or whether further taxonomic adjustments 

are necessary. It is not known to have been planted 

or naturalized elsewhere. 

Ecology.—Bejuco de membrillo has been 

collected from near sea level to over 2,000 m 

elevation in Costa Rica (Missouri Botanical 

Garden 2002). It grows on most soils over nearly 

all types of parent material in areas that receive 

from about 1400 to over 3200 mm of 

precipitation. Bejuco de membrillo is moderately 

intolerant of shade and cannot sustain itself 

indefinitely under a closed forest canopy. It is 

most often found in natural openings of secondary 

and remnant forests. It may also grow in brushy 

pastures, fencerows, and roadsides. 

Reproduction.—In Puerto Rico, bejuco de 

membrillo blooms during July and August and 

fruits in January and February (AcevedoRodríguez 1985). It is reported to flower and fruit 

throughout the year in Nicaragua (Stevens and 

others 2001). A collection of fresh fruits in Puerto 

Rico weighed an average of 0.486 + 0.018 g/fruit. 

Seeds cleaned from these fruits averaged 0.105 +

0.002 g/seed or 9,500 seeds/kg. Planted on 

commercial potting mix, 50 percent of the seeds 

germinated between 65 and 94 days after sowing 

(author’s observation). The seeds are dispersed by 

vine extension and presumably by birds and 

animals. The plant suckers from rhizomes and 

sprouts heavily when cut. 

Growth and Management.—The growth of 

established bejuco de membrillo sprouts is rapid, 

more than 3 m/year. Individual stems live several 

years, and plants, by suckering, live almost 

indefinitely. New plants can probably be 

established by planting the thickened rhizomes in 

tree-fall gaps and other forest openings. Creating 

the essential gaps could also promote the natural 
 

718 

establishment of the species. The seeds must be 

collected by hand and can be cleaned by 

maceration and wet sieving. 

Benefits.—Bjuco de membrillo adds to 

biodiversity, helps to protect the soil, and furnishes 

food and cover for wildlife. Extracts of the plant 

are used as a tonic, a sudorific, and a purifier, and 

to treat skin conditions, rheumatism, and venereal 

disease (Liogier 1990) 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR 566 p. 

Missouri Botanical Garden. 2002. Flora de Costa 

Rica: Smilax domingensis Willd. http://mobot. 

org/manual.plantas/051162/S051173.html. 10 p. 

Missouri Botanical Garden. 3002. W3

 specimen 

data base--28 Aug. 2003. http://mobot. 

mobot.org/cgi_bin/search_vast. [not paged]. 

Natural Resources Conservation Service. 2003. 

Plant profile: Smilax smallii Morong. 

http://plants.usda.gov/ [not paged]. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanical Garden, St. Louis, MO. 

p. 1,911-2,666. 
 

719

Solanum drymophilum O.E. Schulz erubia 

SOLANACEAE 

Synonyms: none 

John K. Francis 

General Description.—Erubia is a rare shrub that 

reaches 3 to 6 m and 7.5 cm or more in basal 

diameter. The plant may have a single stem or 

multiple branches from the base or near the base. 

It is supported primarily by semiflexible lateral 

roots. The foliage is concentrated near the ends of 

twigs. Alternate lanceolate to oblong leaves, 8 to 

20 cm long and 2 to 4 cm wide, are long-pointed 

at the tip and obtuse or subcordate at the base with 

petioles up to 1 cm long. There are sharp, stiff, 

yellow spines up to 1 cm long on the mid-vein of 

the leaves and sometimes on the twigs and stems. 

Five-lobbed white flowers about 2 cm across with 

yellow anthers are grouped in lateral or 

subterminal racemes. The fruits are spherical, 

shiny black berries about 6 mm in diameter that 

contain many tiny seeds (Liogier 1995, Little and 

others 1974, U.S. Fish and Wildlife Service 2002). 

Range.—Erubia is endemic to Puerto Rico. In 

1974, it was known only from the upper Cordillera 

forest of the eastern and central mountains (Little 

and others 1974). Since then, the range has 

become constricted and fragmented until it is 

currently known from one natural site and one 

plantation (Miner Solá 1999) and has been 

declared an endangered species (U.S. Fish and 

Wildlife Service 2002). Deforestation and 

misguided weed control have contributed to the 

species’ scarcity. 

Ecology.—The environment of only the extant 

stands are known. Erubia may actually have a 

much wider ecological amplitude. The existing 

natural stand is situated at 840 m on clayey soil 

derived from volcanic rock. The area receives 

about 1800 mm of mean annual rainfall. The 

plantation is located on clay soil over porous 

limestone in an area that receives about 2200 mm 

of mean annual precipitation. The plants in the 

plantation are healthy but do not appear to be 

reproducing on the site. The species appears to be 

intolerant of shade and requires disturbance to 

become established. Before colonization, it 

probably relied on landslides, tree tips, and other 

natural disturbances to become established. The 

known wild stand is confined to a brushy pasture. 

Reproduction.—Erubia appears to flower and 

fruit throughout the year (U.S. Fish and Wildlife 

service 2002). Flower and fruit production can be 

abundant. The flowers are insect pollinated. 

Twenty-eight fruits collected from a plantation in 

Río Abajo, Puerto Rico weighed (fresh) an 

average of 0.0504 + 0.0050 g. The variability in 

weight is high (coefficient of variation = 52 

percent). Air-dried seeds separated from them 

averaged 0.00092 g/seed or 1.1 million seeds/kg. 

Sown on moist potting mix without pretreatment, 

89 percent of the seeds germinated between 12 

and 50 days after sowing. Birds apparently 

disperse the seeds. Plants that are disturbed 

resprout. 

Growth and Management.—Erubia has a 

moderate growth rate and probably lives at least 

10 years. Nursery production appears to be easy, 

but field planting must be accompanied by at least 

2 years of weed control. 

Benefits.—Erubia helps reforest disturbed sites 

such as abandoned pastures and helps protect the 

soil. It contributes cover for wildlife, and 

presumably, the fruits are eaten by birds. As most 

other members of the Solanaceae family, its 

tissues probably contain physiologically active 

chemicals that might be useful in herbal medicine 

and control of pathogens. 
 

720 

References 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

617 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. U.S. Department of 

Agriculture, Forest Service, Agriculture 

Handbook. 449. Washington, DC. 1,024 p. 

Miner Solá, E. 1999. Arboles y plantas en peligro 

de extinción en Puerto Rico. Puerto Rico 

Ecológico Vol. 3. First Book Publishing of 

Puerto Rico, San Juan, PR. 91 p. 

U.S Fish and Wildlife Service. 2002. Species 

accounts: Erubia (Solanum drymorphilum). 

http://endangered.fws.gov/i/q/saq50.html. 2 p. 
 

721

Solanum dulcamara L. bittersweet nightshade 

SOLANACEAE 

Synonyms: none 

John K. Francis 

General Description.—Bittersweet nightshade is 

also known as bitter nightshade, climbing 

nightshade, woody nightshade, scarlet berry, 

fellenwort, poison bittersweet, and dulcamara. The 

name bittersweet comes from the report that stems 

first taste bitter, then sweet when chewed. It is a 

vine or slender scrambling shrub up to 7 m in 

extension and 2 or 3 cm in basal diameter. The 

base of the plant is woody, and the upper 

herbaceous branches die back to the woody 

portion each year. The slender branches are green 

becoming gray-brown and finely furrowed in the 

lower stem. The wood, which has annual rings, is 

creamy-white, brittle, and of light to medium 

density. The main root grows horizontally just 

below the surface and suckers frequently. Its bark 

is tan, corky, and knotty. The crushed bark and 

foliage have an unpleasant smell. The alternate, 

dark-green leaves are ovate to lanceolate, entire, 

simple or with small, opposite lobes or leaflets 

near the base. The blades are 5 to 12 cm long and 

the petioles 1 to 4 cm long. Inflorescences are 

drooping lateral cymes of usually more than 10 

flowers. The corolla is deeply five-lobbed (star 

shaped), 12 to 16 mm in diameter, blue to violate 

(occasionally white), with exerted yellow anthers 

about 5 mm long. The ripe fruits (berries) are 

bright red, ovoid to globose, 8 to 12 mm long, and 

contain numerous flattened, pale-yellow seeds. 

Chromosome number of the species is 2n = 24 

(Abrams 1951, Stance 1997). 

Range.—Bittersweet nightshade is native to 

temperate Eurasia and northern Africa (Crossley 

1974, Steyermark 1963). It is generally considered 

to be naturalized well into Canada and the United 

States including all the conterminous states except 

Arizona, New Mexico, Texas, Oklahoma, 

Arkansas, Louisiana, Mississippi, Alabama, and 

South Carolina (Natural Resources Conservation 

Service 2003). Its observation by early botanists 

and its presence in native habit led one author to 

conclude that it is native to North America (Dean 

1940). The species has been widely planted for 

ornamental and medicinal purposes. 

Ecology.—Bittersweet nightshade often grows in 

wet sites or sites close to water, but also grows in 

moist upland sites: river banks, weedy ditches, sea 

shores, at the edges of lakes, bogs, and fens, 

roadsides, fencerows, orchards, and open forests. 

A wide variety of soils are colonized. The species 

tolerates dormant-season temperatures well below 

freezing but for unknown reasons has not invaded 

tropical climates. Continuous soil moisture is 

required. Seedlings are tolerant enough to grow 

under a moderate forest canopy (Samodien and 

others 2003). However, mortality of seedlings 

tends to be high (89 to 94 percent) (Kollmann and 

Grubb 1999). Plants in full sun or light shade 

flower and fruit much more heavily than those in 

moderate to medium shade. Bittersweet nightshade 

climbs into small trees, shrubs, weeds, and fences 

or remains prostrate as opportunities avail 

themselves. Plants of the species grow singly or in 

small patches. 
 

722 

Reproduction.—Bittersweet nightshade flowers 

from May to September (Horticulture Purdue 

1998). Fruit and seed production can be abundant. 

Fresh fruits collected by the author in Utah (n = 

50) averaged 0.441 + 0.013 g/fruit. They contained 

an average of 30.9 + 1.6 seeds/fruit, and the airdried seeds averaged 0.00149 + 0.00002 g/seed or 

670,000 seeds/kg. Crossley (1974) reports 938,000 

seeds/kg. The fruits can be collected by hand, 

cleaned by macerating and wet screening, and 

stored at 6 percent moisture for periods of less 

than 1 year. When seeds are germinated under 

light, stratification is not necessary and total 

germination percentages of 61 to 98 percent may 

be reached. Stems and branches layer (root) 

whenever they come in contact with the ground. 

Plants coppice when they are cut or damaged. 

Nursery propagation is usually by seeds, but the 

species can be propagated asexually by root or 

stem cuttings (Crossley 1974). Seeds of wild 

plants are dispersed by birds (Samodien and others 

2003). 

Growth and Management.—Bittersweet 

nightshade branches grow and die back 1 to 3 m or 

more each year. Larger stems examined by the 

author had two or three growth rings. Plants 

probably live much longer by resprouting, 

suckering, and layering. Bittersweet nightshade 

usually is not abundant or aggressive enough to 

need control. Occasional plants or patches in 

gardens, orchards, or landscaped areas can be 

eliminated by grubbing or spot spraying with 

broadleaf or broad-spectrum herbicides. 

Benefits and Detriments.—Bittersweet 

nightshade contributes to the aesthetics of 

wildlands where it grows and furnishes food and 

cover for wildlife. It is a pretty plant, cultivated as 

an ornamental for its colorful flowers and fruits 

and dark green foliage. Several species of moths 

use it for larval food (Savela 2003). The fruits are 

eaten by birds, including pheasants (Phasianus 

colchicus). Although all parts are poisonous to 

cattle, horses, and sheep, which normally will not 

eat it, the stems are eaten by muskrats (Ondatra 

zibethica). The berries have been used to poison 

rabbits and dogs (Steyermark 1963). Although the 

fruits have an attractive appearance, the flavor is 

so disagreeable that it is doubtful that anyone 

would mistakenly eat enough to be poisoned. Also, 

poisonous principals are low in ripe fruits (Lambo 

Seeds 2003). Bittersweet nightshade contains the 

alkaloid solanine (the poisonous principal) and the 

glycoside dulcamarin (responsible for the bittersweet taste). Decoctions, usually of the twigs, were 

once used to treat a variety of ailments. These have 

been largely discontinued because of severe side 

effects. The herb is used today to treat scaly skin 

afflictions and deficient capillary circulation in the 

skin (Herbdata New Zealand 2003). 

References

Abrams, L. 1951. Illustrated flora of the Pacific 

States. Vol. 3. Stanford University Press, 

Stanford, CA. 866 p. 

Crossley, J.A. 1974. Solanum dulcamara L. In: 

C.S. Schopmeyer, tech. coord. Seeds of woody 

plants of the United States. Agriculture 

Handbook 450. U.S. Department of Agriculture, 

Forest Service, Washington, DC. p. 777-778. 

Dean, C.C. 1940. Flora of Indiana. Department of 

Conservation, Division of Forestry, 

Indianapolis, IN. 1,236 p. 

Herbdata New Zealand. 2003. Solanum 

dulcamara, Linn. http://www.herbdatanz.com/ 

Bittersweet.html. 4 p. 

Horticulture Purdue. 1998. Bitter nightshade. 

Purdue University, West Lafayette, IN. http:// 

www.hort.purdue.edu/newcrop/herbhunters/ 

nightshade.html. 1 p. 

Kollmann, J. and P.J. Grubb. 1999. Recruitment of 

fleshy-fruited species under different shrub 

species: control by under-canopy environment. 

Ecological Research. http://dogwood.ag. 

utk.edu/literature/1999/99_kollmann.htm. 1 p. 

Lambo Seeds. 2003. Genus: Solanum. http://www. 

aros.net/~lambo/dulcamara/dulcamara01.htm. 

8 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Solanum dulcamara L., climbing 

nightshade. http://plants.usda.gov.cgi_bin/ 

plant_profile.cgi?symbol=SODU. 6 p. 

Samodien, J., M. Wood, K. Epps, and M. Thandy. 

2003. An introduction to common exotic species 

to British Columbia’s ecosystems. University 

College of Fraser Valley, Abbotsford, BC, 

Canada. http://www.ucfv.bc.ca/biology/Biol210/ 

1999/Exotic/Exotic_plant.htm. 9 p. 
 

723

Savela, M. 2003. Solanum L. http://www. 

funet.fi/pub/sci/bio/life/plants/magnoliophyta/ 

magnoliophytina/magnoliopsida/solanaceae/ 

solanum/. 5 p. 

Stance, C. 1997. New flora of the British Isles. 2nd. 

Ed. Cambridge University Press, Cambridge, 

UK. 1,130 p. 

 

Steyermark, J.A. 1963. Flora of Missouri. The 

Iowa State University Press, Ames, IA. 1,725 p. 
 

724 

Solanum persicifolium Dunal berenjena de playa 

SOLANACEAE 

Synonyms: Solanum ignaeum parvifolium Vahl 

 Solanum persicifolium angustifolium Dunal in DC. 

 Solanum persicifolium belloni O.E. Schulz in Urban 

 Solanum persicifolium parvifolium (Vahl) O.E. Schulz 

John K. Francis 

General Description.—Berenjena de playa, 

which means beach eggplant, is a semideciduous 

shrub usually about 2 m in height with a basal 

diameter of 1.5 cm. The species is also known as 

rubia. It may have a single or multiple stems and 

is supported by a stout taproot and many lateral 

roots. The most striking feature of the species is 

the sharp yellow to orange prickles on the stem, 

branches and midveins of the leaves. The prickles 

may reach as much as 13 mm long. However, the 

density of prickles is variable and some plants 

have none. The branches and twigs are slender and 

form a thin crown. The alternate leaves, which 

have 4- to 7-mm petioles, are thin, oblong to 

linear-oblong, 5 to 13.5 cm long and 0.6 to 3 cm 

wide with pointed ends and edges rolled under 

(revolute). The leaves take on a wilted appearance 

during dry periods. The raceme-like inflorescences 

are lateral and contain up to 15 small, blue, violet, 

or rarely white flowers. The fruits are globose, red 

berries about 5 mm in diameter with the calyx still 

attached. Each contains several flattened, yellow 

seeds (author’s observation, Liogier 1995). 

Range.—Berenjena de playa is native to 

Hispaniola, Puerto Rico, and the Virgin Islands 

(Liogier 1995). It is not known to have been 

planted or naturalized elsewhere. 

Ecology.—Berenjena de playa is most common 

near the sea shore but does occur several km 

inland. It grows on the whole range of soil textures 

on well-drained soils. In Puerto Rico, the species 

colonizes areas that receive from 750 mm to about 

1700 mm of mean annual precipitation from near 

sea level to over 400 m of elevation. Berenjena de 

playa is moderately intolerant of shade and grows 

within low basal-area forest as well as in 

openings. The plants are apparently avoided by 

cattle and appear to benefit by overgrazing. Some 

degree of disturbance appears to be necessary for 

establishment. Vázquez and Kolterman (1998) 

report it as “occasional” in pasture, shrubland, and 

woodland. It may also be found on roadsides, 

trails, fencerows, bluffs, and beach strands. 

Reproduction.—Berenjena de playa flowers and 

fruits more or less continuously in moist habitat 

but seasonally in dry habitat. A group of fruits 

collected in Puerto Rico averaged 0.0708 + 0.0035 

g/fruit. They contained from three to 20 seeds 

each, which averaged (air-dried) 0.00137 + 0.0003 

g/seed or 730,000 seeds/kg. The seeds are 

apparently dispersed by birds that eat the fruits. 

Plants sprout when they are broken or cut. 

Growth and Management.—Sprouts of 

berenjena de playa grows about 0.5 to 1.0 m in the 

first year and slower thereafter. Seedling growth is 

undoubtedly slower. Stems apparently live about 3 

to 10 years, but the rootstalks resprout and may be 

older than the top. No management experience has 
 

725

been published. Small areas of disturbance in 

suitable habitat would probably encourage natural 

establishment. The species is rarely common 

enough to control as a weed in pastures or 

plantations. 

Benefits.—Berenjena de playa contributes to the 

biodiversity of habitat where it grows, helps to 

protect the soil, and furnishes food and cover for 

wildlife, particularly birds. 

References 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

617 p. 

Vázquez, O.J. and D.A. Kolterman. 1998. Floristic 

composition and vegetation types of the Punta 

Guaniquilla Natural Reserve—Cabo Rojo, 

Puerto Rico. Caribbean Journal of Science 34(3-

4): 265-279. 
 

726 

Solanum torvum Sw. turkey berry 

SOLANACEAE 

Synonyms: Solanum ficifolium Ortega 

 Solanum daturifolium Dunal 

 Solanum torvum var. daturifolium (Dunal) Schulz 

 Solanum maccai L.C. Rich. ex Spreng. 

John K. Francis 

General Description.—Turkey berry is an erect 

spiny shrub that is also known by prickley 

solanum, shoo-shoo bush, devil’s fig, wild egg 

plant, boo, terongan, berenjena cimarrona, 

berenjena de gallina, berenjena silvestre, tabacón, 

pendejera, tomatillo, bâtard balengène, zamorette, 

friega-platos, and many other names (Howard 

1989, Little and others 1974, Pacific Island 

Ecosystems at Risk 2001). It is usually 2 or 3 m in 

height and 2 cm in basal diameter, but may reach 

5 m in height and 8 cm in basal diameter. The 

shrub usually has a single stem at ground level, 

but it may branch on the lower stem. The stem 

bark is gray and nearly smooth with raised 

lenticels. The inner bark has a green layer over an 

ivory color (Little and others 1974). The plants 

examined by the author, growing on firm soil, had 

weak taproots and well developed laterals. The 

roots are white. Foliage is confined to the growing 

twigs. The twigs are gray-green and covered with 

star-shaped hairs. The spines are short and slightly 

curved and vary from thick throughout the plant, 

including the leaf midrib, to entirely absent. The 

leaves are opposite or one per node, broadly ovate 

with the boarder entire or deeply lobed. The 

petioles are 1 to 6 cm long and the blades are 7 to 

23 by 5 to 18 cm and covered with short hairs. The 

flowers are white, tubular with 5 pointed lobes, 

and grouped in corymbiform cymes. They are 

shed soon after opening. The fruits are berries that 

are yellow when fully ripe. They are thin-fleshed 

and contain numerous flat, round, brown seeds 

(Howard 1989, Liogier 1995, Little and others 

1974). 

Range.—Turkey berry apparently is native from 

Florida and southern Alabama through the West 

Indies and from Mexico through Central America 

and South America through Brazil (Little and 

others 1974). Because of its rapid spread as a 

weed in disturbed lands, it is difficult to tell which 

populations are native and which are introduced. 

Turkey berry has been introduced and naturalized 

throughout tropical Africa, Asia, Australia, and 

the Pacific Islands including Hawaii, Guam, and 

American Samoa (Pacific Island Ecosystems at 

Risk 2001). 

Ecology.—In Puerto Rico, turkey berry grows in 

upland sites that receive from about 1000 to 4000 

mm of annual precipitation. It also grows in 

riparian zones in drier areas. Turkey berry grows 

on all types of moist, fertile soil at elevations from 

near sea level to almost 1,000 m in Puerto Rico 

(Little and others 1974) and 2,000 m in Papua 

New Guinea (Pacific Island Ecosystems at Risk 

2001). Given an equal start after disturbance, 

turkey berry quickly overtops most herbs, grasses, 

and other shrubs. It grows best in full sunlight and 

does well in light shade or shade for part of the 

day, but cannot survive under a closed forest 

canopy. Turkey berry single plants, groups, and 

thickets are most frequently seen on roadsides, 

vacant lots, brushy pastures, recently abandoned 

farmland, landslides, and river banks. 

Reproduction.—Flowering and fruiting is 

continuous after the shrubs reach about 1 to 1.5 m 

in height. Ripe fruits collected in Puerto Rico 

averaged 1.308 + 0.052 g. Air dry seeds from 

these fruits weighed an average of 0.00935 g or 

1,070,000 seeds/kg. These seeds were sown on 

commercial potting mix and 60 percent 

germinated between 13 and 106 days following 

sowing. The seedlings are common in recently 
 

727

disturbed ground. Frugivorous birds eat the fruits 

and spread the seeds (Pacific Island Ecosystems at 

Risk 2001). Turkey berry can be propagated 

vegetatively by placing branch cuttings, with or 

without leaves, in a mist chamber for one month 

(Badola and others 1993). 

Growth and Management.—Turkey berry grows 

about 0.75 to 1.5 m in height per year. The species 

is not long-lived; most plants live about 2 years. 

Physical control of the shrub may be done by 

grubbing out the plants; lopping will not kill them. 

They can be killed by translocated herbicides 

applied to the leaves or the cut stumps (Pacific 

Island Ecosystems at Risk 2001). 

Benefits.—The wood is soft and light and of little 

use except for emergency fuel. The fruits are 

edible and used in Thai cuisine (Royal 

Horticultural Society 2001) and incorporated into 

soups and sauces in the Ivory Coast (Herzog and 

Gautier-Béguin 2001). Turkey berry contains a 

number of potentially pharmacologically active 

chemicals including the sapogenin steroid, 

chlorogenin (Badola and others 1993). Aqueous 

extracts of turkey berry are lethal to mice or 

depress the erythrocytes, leukocytes and platelets 

in their blood (Tapia and others 1996). A related 

chemical, cholecalciferol, is the active ingredient 

in a number of commercial rodentacides 

(American Board of Veterinary Toxicology 2001). 

Extracts of the plant are reported to be useful in 

the treatment of hyperactivity (Null 2001), colds 

and cough (CPR Environmental Education Centre 

2001), pimples, skin diseases, and leprosy (Liogier 

1990). Turkey berry is being crossed with 

eggplant in an attempt to incorporate genes for 

resistance to Verticillium wilt into the vegetable 

(Bletsos and others 2001). 

References 

American Board of Veterinary Toxicology. 2001. 

An overview of cholecalciferol toxicosis. 

http://www.abvt.org/tow.htm. 6 p. 

Badola, K.C., P. Mohinder; H.C.S. Bhanderi, and 

M. Pal. 1993. Vegetative propagation of 

ranbaigan (Solanum torvum Sw.) by rooting 

branch cuttings. Indian Forester 119(12): 1,027-

1,028. 

Bletsos, F.A., D.G. Roupakias, and C.C. 

Thanassoulopoulos. 2001. Morphological traits 

of the interspecific eggplant hybrid Solanum

melongena x Solanum torvum. 

http://www.actahort.org/books/467/467_17.htm. 

1 p. 

CPR Environmental Education Centre. 2001. 

Medicinal plants. http://cpreec.org/edu/ 

 medi-pln.htm. 4 p. 

Horzog, F. and D. Gautier-Béguin. 2001. 

Uncultivated plants for human nutrition in Côte 

d’Ivorie. http://www.fao.org/docrep/W3735e/ 

w3735e10.htm. 12 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Part 3. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

617 p. 

Liogier, H.A. 1990. Plantas mediciales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Null, G. 2001. The biochemical activity of plants. 

http://www.garynull.com/Documents/phytoche

micals/Phytochemicals5.htm. 15 p. 

Pacific Island Ecosystems at Risk. 2001. Invasive 

plant species: Solanum torvum Sw., Solanaceae. 

http://www.hear.org/pier/sotor.htm. 2 p. 

Royal Horticultural Society. 2001. Fruits and 

vegetables: felty leaves. 

http://www.rhs.org.uk/about/mn_pubs_journals_

garden_0898_aubergine_b.asp. 1 p. 
 

728 

Tapia A., R., A. Astudillo V., R. Uribe H., 1996. 

Resultados preliminaries del efetco de Solanum

torvum y Plantago major sobre la proliferación 

de células hematopoyéticas in vivo e in vitro. 

Resumen de Ponencias de Primer Congreso 

Nacional de Plantas Medicinales de México. 24-

30 June 1996. Tlaxcala, Tlax., Mexico. 102-

104. 

 
 

729

Sorbus scopulina Greene Greene’s mountain-ash 

ROSACEAE 

Synonyms: Sorbus sambucifolia non Roem. 

 Sorbus cascadensis G.N. Jones 

John K. Francis 

General Description.—Greene’s mountain-ash, 

also known as western mountain-ash, is a 

deciduous shrub or small tree ranging in height 

from 1 to 6 m and up to 10 cm in stem diameter. It 

usually has multiple stems with smooth yellowish 

to grayish-red bark and slender light brown twigs 

that are white-hairy when young. Winter buds are 

glutinous and glossy. The alternate leaves are 10 to 

20 cm long, odd-pinnately compound with seven 

to 15 lanceolate leaflets that are nearly sessile, 

pointed, and serrate on the margins. They are thin, 

shiny-green above and paler beneath. 

Inflorescences are much-branched corymbs, 6 to 

12 cm broad, that contain many 10-mm broad, 

white to cream, five-petaled flowers. Fruits, which 

grow in clusters, are shiny, orange to red, berrylike, 5- to 10-mm-long, globose pomes with an 

attached calyx at the apex. Each contains up to 

eight flattened, brown or red-brown seeds 3 to 4 

mm long (Davis 1952, Viereck and Little 1972, 

Welsh 1974). 

Range.—Greene’s mountain-ash is native to broad 

areas and disjuncts in the southern half of Alaska, 

disjuncts and broad areas of the Yukon, Northwest 

Territories, British Columbia, Alberta, 

Saskatchewan, Montana, Idaho, Washington, and 

scattered areas in Oregon, California, Wyoming, 

North Dakota, South Dakota, Utah, Colorado, 

Nevada, and New Mexico (Natural Resources 

Conservation Service 2003, Treeguide 2003). 

There are two varieties. The typical variety 

occupies the entire inland range and var. 

cascadensis (G.N. Jones) C.L. Hitchc. occupies 

coastal habitat from British Columbia into 

California (Natural Resources Conservation 

Service 2003). In addition, an intergeneric hybrid 

named Amelasorbus jackii Rehder between 

Amelanchier alnifolia Nutt. and Greene’s 

mountain-ash has been observed in Oregon and 

Idaho (Love 2003). Greene’s mountain-ash has 

been widely planted, but no reports of it 

naturalizing outside its range are known. 

Ecology.—Greene’s mountain-ash colonizes welldrained soils on rocky hillsides, canyons, wooded 

slopes, forest clearings, avalanche chutes, and 

along streams. It grows as dispersed individuals or 

in nearly pure (usually small) clumps. The sites are 

moist; the species is not drought resistant. 

Elevations range from near sea level in the 

northern portions of its range to 3,000 m in the 

Southern Rocky Mountains. Greene’s mountainash is moderately intolerant of shade; it sometimes 

grows in the understory of open stands but usually 

grows in openings. Low-vigor plants are also 

common under closed, but not dense, conifer 

stands (Alaska Department of Fish and Game 

2001, Ibiblio 2003, Luna and Wick 2001, Umatilla 

National Forest 2003, Utah State University 

Horticulture 2003). 

Reproduction.—Flowering of Greene’s 

mountain-ash occurs between May and July 

depending on location (Stein 2003). The flowers 

are pollinated by insects (Ibiblio 2003). The fruits 

ripen in July through September and remain on the 

shrubs into the winter (Alberta Agriculture, Food 

and Rural Development 2003). Fruits can be 

collected in quantity by hand after the fruits turn 

orange or red and the seeds are brown. The seeds 

are extracted by maceration, washing, and 

screening. The weight of seeds has been reported 

at 100 seeds/g (Luna and Wick 2001) and 180 

seeds/g (Browse-Shrub and Forb Committee of the 

Association of Official Seed Analysis 1985). In a 
 

730 

collection made by the author in Utah, fresh fruits 

weighed an average of 0.333 + 0.009 g/fruit, and 

air-dried seeds weighed an average of 0.00386 +

0.0001 g/seed (259 seeds/g). There were 24 to 117 

fruits/cluster and an average of 2.4 seeds/fruit. 

Seed will remain viable in sealed, refrigerated 

containers at low relative humidity for 5 years. 

Recommended pretreatment is a 3:1 

water/hydrogen peroxide soak for 10 minutes 

followed by a 25-hour water soak and a cold 

stratification in peat at 3 °C for 90 to 120 days. 

Germination varies from 90 to 100 percent (Luna 

and Wick 2001). A test of viability can be made by 

incubating imbibed, excised embryos at 20 °C for 

6 days. Viable embryos either retain their freshly 

excised appearance or become deep green; 

nonviable embryos deteriorate or turn pale yellow 

green (Umatilla National Forest 2003). In the wild, 

seeds are dispersed mainly by birds. Greene’s 

mountain-ash may be asexually reproduced using 

cuttings. Late-summer semihardwood stem 

cuttings 15 to 25 cm long and 0.3 to 1.3 cm in 

diameter are recommended. After hormone 

treatment, cuttings placed in a bottom-heated 

mistbed for 6 weeks yielded 47 percent rooted 

cuttings (Luna and Wick 2001). 

Growth and Management.—This species grows 

slowly as seedlings and only at an intermediate 

rate later. Plants are able to flower and fruit after 3 

to 5 years (Luna and Wick 2001). Greene’s 

mountain-ash is usually grown for revegetation 

projects from seed in nurseries as container or 

bare-root stock. Potted seedlings can be expected 

to reach 15 cm tall with a 2.0 cm caliper by the 

end of the first year (Luna and Wick 2001). They 

are outplanted as either 1- or 2-year-old stock. 

Greene’s mountain-ash planted in seven sites in 

Alberta averaged 1.2 to 2.3 m in height after 4 to 8 

years of growth (Alberta Agriculture, Food and 

Rural Development 2003). Direct seeding is also 

possible. Drilling seed 2 mm deep in fertile, welldrained soil is recommended. Many seeds will not 

germinate until the second or third year (Umatilla 

National Forest 2003). Although the species is 

adapted to many soil conditions, it is sensitive to 

high pH in the nursery and the field and is 

damaged by the pear slug [Caliroa cerasi (L.)] 

(Alberta Food and Rural Development 2003). 

Browsing keeps the plants compact in many 

locations (Clark 1976). The author knows of no 

published data on management of natural stands. 

Benefits.—Greene’s mountain-ash is an important 

component of the Western shrub community and 

furnishes a number of benefits. The species helps 

protect the soil, adds to the aesthetics of wildland 

sites, especially with its yellow to orange-red fall 

foliage and red-orange berries, and furnishes cover 

for wildlife. The foliage and twigs are browsed by 

deer (Odocoileus spp.), elk (Cervus canadensis), 

moose (Alces americana), and to a lesser extent, 

cattle. The fruits are eaten by black bears 

(Euarctos americanus), martin (Martes 

americana), fisher (Martes pennanti), rodents, 

American robin (Turdus migratorius), hermit 

(Catharus guttatus), gray-cheeked (Catharus 

minimus) and Swainson’s (Catharus ustulatus) 

thrushes, pine grosbeaks (Pinicola enucleator), 

Bohemian waxwings (Bombycilla garrulus), 

northern flickers (Colaptes auratus), Steller’s jays 

(Cyanocitta stelleri), and blue grouse 

(Dendragapus obscurus) (Alaska Department of 

Fish and Game 2001, Gullion 1964, Stein, 2003, 

Wier 2003). The fruits are edible to humans, fresh, 

cooked, and dried, but it is necessary to wait until 

the bitterness disappears after multiple frosts 

(Moser 2003). A wine is also made from the 

berries (Keller 2003). In herbal medicine, 

infusions of bark have been used to reduce fevers 

and as a tonic, and infusions of branches have been 

given to children with bed-wetting problems 

(Ibiblio 2003). Greene’s mountain-ash is planted 

to a limited extent as an ornamental, especially in 

naturalistic landscape settings. The wood is soft 

and has a specific gravity of 0.59 g/cm3

 (Ibiblio 

2003). It is probably useful for firewood, but other 

uses are not known. 

References

Alaska Department of Fish and Game. 2001. 

Native Alaskan and exotic plants used by 

wildlife. http://www.state.ak.us/adfg/wildlife/ 

geninfo/birds.htm. 9 p. 

Alberta Agriculture, Food and Rural Development. 

2003. Evaluating woody plants for hardiness and 

landscaping quality in Alberta: Sorbus sp. 

(mountain ash). http://www.agric.gov.ab.ca/ 

crops/trees/rwptp/ sorbus.html. 3 p. 

Browse-Shrub and Forb Committee of the 

Association of Official Seed Analysis. 1985. 

Handbook on seeds of browse-shrubs and forbs. 

Technical Publication R8-TP8. U.S. Department 

of Agriculture, Forest Service, Southern Region, 

Atlanta, GA. 246 p. 

Clark, L.J. 1976. Wild flowers of the Pacific 

Northwest. Gray’s Publishing, Limited. Sidney, 

BC, Canada. 604 p. 
 

731

Davis, R.J. 1952. Flora of Idaho. Brigham Young 

University Press, Provo, UT. 836 p. 

Gullion, G.W. 1964. Wildlife uses of Nevada 

flora. Contr. Toward a Flora of Nevada 49. U.S. 

National Arboretum, Washington, DC. 170 p. 

Ibiblio. 2003. Plants for a future: database search 

results: Sorbus scopulina. University of North 

Carolina, Chapel Hill, NC. http://www.ibiblio. 

org/pfaf.cgi-bin/arr_html?Sorbus+scopulina& 

CAN =LATIND. 6 p. 

Keller, J. 2003. The winemaking home page: 

mountain ash. http://winemaking.jackkeller.net/ 

mtnash.asp. 2 p. 

Love, R. 2003. Amelasorbus: An intergeneric 

hybred and a new taxon for the Oregon 

checklist. http://www.oregonflora.org/ofn/v4n2/ 

amelasorbus.html. 3 p. 

Luna, T. and D. Wick. 2001. Propagation protocol 

for production of container Sorbus scopulina

Greene var. scopulina Greene plants (172 ml 

containers), Glacier National Park, West 

Glacier, Montana. http://www. 

nativeplantnetwork.org. 8 p. 

Moser, R. 2003. Edible Sierra Nevada plants: 

Rosaceae-rose family, mountain ash. 

Backcountry rangers, Fresno, CA. http://www. 

backcountryrangers.com/edibles/SORBUS.html. 

2 p. 

Natural Resources Conservation Service. 2003. 

Sorbus scopulina Greene, Greene’s mountain 

ash. http://plants.usda.gov/cgi_bin/plant_profile. 

cgi?symbol=SOSC2. 4 p. 

Stein, W.I. 2003. Sorbus L, mountain-ash. In: F.T. 

Bonner, and R.G. Nisley, eds. Woody plant seed 

manual. U.S. Department of Agriculture, Forest 

Service, Washington, DC. http://wpsm.net/ 

Sorbus.pdf. 12 p. 

Treeguide. 2003. Greene mountain-ash, Sorbus 

scopulina, Greene, Rosaceae. 

http://www.treeguide.com/Species.asp?Region=

NorthAmerican&SpeciesID=1021. 2 p. 

Umatilla National Forest. 2003. Umatilla National 

Forest native plant species: Sorbus scopulina. 

U.S. Department of Agriculture, Forest Service, 

Umatilla National Forest, Pendleton, OR. http:// 

www.fs.fed.us/r6/uma/native/ts100.htm. 8 p. 

Utah State University Horticulture. 2003. Sorbus 

scopulina. http://www.usu.edu/natives/shrubs/ 

scopulina.htm. 1 p. 

Viereck, L.A. and E.L. Little, Jr. 1972. Alaska 

trees and shrubs. Agriculture Handbook 410. 

U.S. Department of Agriculture, Forest Service, 

Washington, DC. 265 p. 

Welsh, S.L. 1974. Anderson’s flora of Alaska. 

Brigham Young University Press, Provo, UT. 

724 p. 

Wier, S.K. 2003. Greene’s mountain-ash of the 

southern Rocky Mountains. http://home. 

earthlink.net/~swier/MtnAsh.html. 1 p. 
 

732 

Spermacoce verticillata L. botón blanco

RUBIACEAE 

Synonyms: Borreria verticillata (L.) Meyer 

 Borreria podocephala DC. 

 Borreria stricta DC. 

John K. Francis 

General Description.—Botón blanco, which 

means white button in Spanish (Puerto Rico), is 

also known as shrubby false buttonwood, 

vassourinha, cardio de frade, poaia, borrerie 

verticillée, éribun, and many other names (Burkill 

2000, Instituto de Biociências 2002, Natural 

Resources Conservation Service 2002). It is a finestemmed scrambling shrub that may reach a few 

meters of lateral extension and 1.2 m in height as a 

free-standing plant. The square stems are 

herbaceous to semiwoody in their first year, 

becoming woody and more rounded in the 

following year. The brown stems reach a 

maximum diameter of about 8 mm, have a solid 

pith, and lack visible annual rings. Botón blanco 

produces a weak taproot, many important laterals 

that are pale yellow and flexible, and a moderate 

amount of fine roots. Branching is bifurcate or 

ternate. The leaves are opposite but appearing with 

two or a cluster of smaller leaves in whorls at the 

nodes. The leaves are sessile or nearly so, linear or 

linear-lanceolate, 2 to 6 cm long, and pointed at 

both ends. The tiny white flowers grow in heads or 

glomerules in terminal or lateral positions. The 

terminals continue to grow through the center of 

the inflorescence so that the fruits develop at 

nodes in mid-stem. The capsules are oblong or 

subglobose with two carpels, each with one seed. 

The seeds are ellipsoidal, brown, and about 1 mm 

long (Correll and Johnston 1970, Howard 1989, 

Liogier 1997).

Range.—Botón blanco appears to be a native of 

the New World and possibly Africa, but the 

original range is uncertain. It grows as a native or 

naturalized species from Florida through the West 

Indies, and Texas through Central and South 

America to Argentina, and throughout the moist 

portions of Tropical Africa and Madagascar 

(Burkill 2000, Howard 1989, Instituto Botánico 

Darwin 2002, Liogier 1997, Natural Resources 

Conservation Service 2002). It has also been 

reported from India (Kudremukh Wildlife 

Foundation 2002). 

Ecology.—Botón blanco grows on moist soils, 

both acid and alkaline, of all textures derived from 

nearly all types of rocks. The species grows in 

areas that receive from about 750 to 3000 mm of 

annual rainfall from near sea level to 600 m or 

more in elevation in Puerto Rico. It grows on sand 

and caliche in prairies and openings in Texas 

(Jones 1975). In Puerto Rico, it grows on 

roadsides, construction sites, old fields, and 

pastures. Botón blanco competes with cultivated 

crops and plantations in Brazil and Africa (Holm 

and others 1997). It is one of the major invaders of 

abandoned pastures and slash-and-burn fields 

(Ministério de Ciencia e Tecnología 2002). The 

species requires disturbance to establish itself and 

must have full or good partial sunlight to survive. 

It competes well with disbursed grass and weeds, 
 

733

but is overcome by dense, tall grass, brush, and 

trees. Because of grazing, mowing, and 

cultivation, most plants do not progress beyond 

the herbaceous stage. If allowed to grow, they will 

form dense clumps and mats.

Reproduction.—In Brazil botón blanco blooms 

from February through August (Instituto de 

Biociêcias 2002). In Texas, it flowers from March 

through May (Correll and Johnston 1970). 

Flowering is almost continuous in moist portions 

of Puerto Rico. Plants begin blooming in the 

nursery at about 9 months. The flowers are 

pollinated by several species of bees (Instituto de 

Biociências 2002). Seeds collected in Puerto Rico 

averaged 0.00016 g each or 6,250,000 seeds/kg. 

Sown on peat without pretreatment, these seeds 

germinated at 49 percent beginning in 13 days and 

ending at 74 days. The seeds are disbursed by 

grazing animals and farm equipment. Established 

plants root readily at the nodes when covered by 

soil or rotting plant material.

Growth and Management.—Botón blanco 

seedlings grow slowly at first but begin rapid 

growth after about 6 months. Twenty-three 9-

month-old nursery plants averaged 64 cm in 

height with a maximum of 109 cm. Botón blanco 

shrubs appears to live at least 4 years and probably 

much longer in Puerto Rico. Botón blanco is 

controlled in crops and pasture by cultivation, 

mowing, and spraying with broadleaf herbicides. 

The importance value of botón blanco in a 

Colombian pasture was reduced by 39 percent by 

simply fertilizing with potassium and sulfur to 

increase the vigor of the pasture grasses (Tejos 

1981). 

Benefits.—Botón blanco is a forage plant, but not 

one highly favored by livestock. African material 

was found to contain 0.2 percent alkaloid 

including borreine and borreverine (Burkill 2000). 

At least part of the alkaloids are beta-carbolines 

and would represent a poisoning hazard if they 

were present in higher concentrations (Animal 

Science at Cornell University 2002). Botón blanco 

has a number of uses in herbal medicine, most 

frequently for skin conditions. In Africa, leaf 

extracts are used to treat leprous conditions, 

furuncles, ulcers, and gonorrheal sores (Burkill 

2000, Environnement et Développement du Tiersmonde 2002). A lotion is prepared to relieve skin 

itches (Liogier 1990). Other preparations are used 

internally to treat diarrhea, as a diuretic in the 

treatment of schistosomiasis, and as an abortive. 

An essential oil extracted from the leaves has been 

shown to inhibit Escherichia coli and 

Staphylococcus aureus (Burkill 2000).

References 

Animal Science at Cornell University. 2002. Mind 

bending beta-carbolines. http://www.ansci. 

cornell.edu/plants/toxicagents/betacarbolines/ 

bcarb.html. 5 p. 

Burkill, H.M. 2000. The useful plants of West 

Tropical Africa. Royal Botanic Gardens, Kew, 

UK. 686 p. 

Correll, D.S. and M.C. Johnston. 1970. Manual of 

the vascular plants of Texas. Texas Research 

Foundation, Renner, TX. 1,881 p. 

Environnement et Développement du Tiersmonde. 2002. Reconnaisssez, Protégez et 

utilisez: Borreria vericillata L. Rubiaceae. 

http://www.enda.sn/plantesmed/borreria.html. 

2 p. 

Holm, L., J. Doll, E. Holm, J. Pancho, and J. 

Herberger. 1997. World weeds. John Wiley & 

Sons, Inc. New York. 1,129 p. 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Instituto de Biociências. 2002. Borreria 

verticillata. Instituto de Biociêcias, 

Universidade de São Paulo, São Paulo, Brazil. 

http://ib.usp.br/beeplant/bove.htm. 1 p. 

Instituto Botánico Darwin. 2002. Catálogo de las 

plantas vasculares de la Argentina. http:// 

www.darwin.edu.ar/Catalogo/rubiaceae.pdf. 

22 p. 

Jones, F.B. 1975. Flora of the Texas Costal Bend. 

Mission Press, Corpus Christi, TX. 262 p. 

Kudremukh Wildlife Foundation. 2002. Checklist 

of non-woody plants reported from Kudremukh 

National Park. http://kudremukh.org/ 

kudremukh/nonwoody.html. 4 p. 
 

734 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Ministério de Ciencia e Tecnología. 2002. 

Recuperação de areas de pastagems 

abandonadas e degradadas através de sistemas 

agroflorestais na Amazônia Ocidental. 

http://www.mct.gov.br/prog/ppg7/projetos/ 

proj991.pdf. 16 p. 

Natural Resources Conservation Service. 2002. 

Plants profile: Spermacoce verticillata L., 

shrubby false buttonweed. http://plants.usda. 

gov/cgi_bin/plant_profile.cgi?symbol=SPVE2. 

2 p. 

Tejos, R. 1981. Efecto del potasio y azufre sobre el 

pastizal nativo de una sabana. 1. Producción 

contenido de material seca y composicion 

botánica. Agronomía Tropical 29(6): 503-516. 
 

735

Strumpfia maritima Jacq. pride-of-Big-Pine 

RUBIACEAE 

Synonyms: none 

John K. Francis 

General Description.—Pride-of-Big-Pine, also 

known as snowbank, rosemary, strumpfia, lirio, 

and rosarin-bord-de-mer, is an evergreen shrub 

usually 1 m or less in height but sometimes 

reaching 2 m. Most plants have a profusion of gray 

stems and branches. The twigs are slender (3 to 4 

mm in diameter), and prominently ringed from 

persistent stipule bases. The whorled dark-green 

foliage is crowded at the ends of branches. Leaves 

are linear and leathery, 1 to 3 cm long with 

petioles about 1 mm long, with an entire and 

revolute margin. Inflorescences are axillary, fewflowered racemes. The individual flowers are 

white to pink, campanulate with a tube about 1 

mm long and corolla 6 mm across with five lobes. 

The fruit, a white, fleshy globose drupe, is 3 to 6 

mm in diameter with a persistent calyx. It contains 

one or two seeds (Howard 1989, Liogier 1997, 

Nelson 1996). 

Range.—Pride-of-Big-Pine is native to the Florida 

Keys, the Bahamas, the Greater and Lesser 

Antilles, Grand Cayman, several islands north of 

Venezuela, and Yucatan, Mexico (Howard 1989, 

Liogier 1997, Nelson 1996). The species is listed 

as endangered in Florida (Florida Fish and 

Wildlife Conservation Commission 1997). It is not 

known to have naturalized outside its native range. 

Ecology.—Pride-of-Big-Pine is confined to 

coastal areas. Types of habitat include rocky 

headlands, coastal cliffs, beach strands, and coastal 

flats from near sea level to a few meters in 

elevation. It frequently grows in cracks in rocks or 

rocky rubble. The soils are salty, and the plant 

tolerates sea water overwash and heavy salt spray. 

Mean annual precipitation ranges from about 600 

to 900 mm. Pride-of-Big-Pine grows singly, 

scattered, or in small thickets. When growing in 

severe habitats, it sometimes forms natural 

bonsais. 

Reproduction.—Pride-of-Big-Pine blooms and 

fruits throughout the year (Nelson 1996). The 

flowers are probably pollinated by insects. The 

fresh fruits weigh about 0.02 g each. A collection 

of air-dried seeds (n = 99) from Puerto Rico 

averaged 0.0052 + 0.0001 g/seed or 192,000 

seeds/kg. The seeds were placed to germinate on 

the surface of moist peat, and 6 percent germinated 

between 4 and 6 months after sowing. Germination 

is epigeous. Plants coppice when broken or cut 

(author’s observation). The seeds are probably 

dispersed by birds. 

Growth and Management.—Pride-of-Big-Pine 

grows slowly. Planting experience for this species 

has not been published. Because it is rare or 

uncommon and never aggressive, control is not 

necessary. Probably the best management is to 

totally protect existing stands and suitable habitat 

from disturbance. 

Benefits.—Pride-of-Big-Pine helps protect the soil 

in coastal sites and provides cover for wildlife. 

The Exuma Island iguana (Cyclura cychlura 

figginsi), an endangered species from the 

Bahamas, feeds on fruits, flowers, buds, and leaves 

of Pride-of-Big-Pine (Knapp 2002). Probably 

many other species eat the fruits. It is a pretty plant 

and adds to the aesthetics of coastlines. If 

propagation and cultural techniques can be worked 

out, it would make a valuable ornamental. 

Infusions of the leaves are reported to be a 

stimulant useful in treating poisonous bites, fever, 

and stupor and weakness caused by fever (Liogier 

1990). 
 

736 

References

Florida Fish and Wildlife Conservation 

Commission. 1997. Florida’s endangered 

species, threatened species, and species of 

special concern. http://floridaconservation.org/ 

pubs/endanger.html. 18 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Part 3. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Knapp, C. 2002. Exuma Island iguana, Cyclura 

cychlura figginsi. Iguana Specialist Group home 

page. http://www.iucn-isg.org/actionplan/ch2/ 

exumaisland.php. 5 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 
 

737

Styrax americanus Lam. American snowbell 

STYRACACEAE 

 

Synonyms: Styrax pulverulentus Michx. 

Kristina Connor 

Illustration source: USDA—Soil Conservation Service, 

Southern Technical Center

General Description.—American snowbell, also 

known as mock orange or storax, is a deciduous 

shrub or small tree with a widely branched crown. 

It reaches 3 to 5 m in height, and the stems can 

reach 7.5 cm in diameter (Johnson and Hoagland 

1999, Plant Identification Resource 2001). While 

the bark on the stems is smooth and dark grey to 

brown, branches range in color from green to grey 

to red-brown. Young stems are pubescent, 

becoming glabrous with age (Johnson and 

Hoagland 1999). The alternate, simple leaves are 

2.6 to 9 cm long, ovate to elliptic, entire to finely 

serrate, glabrous above, and pubescent beneath 

(Bailey and Bailey 1976, Krüssmann 1986). The 

leaf apex is acute, venation is pinnate. The showy, 

fragrant white flowers for which the plant is noted 

form in racemes from April to June (Iverson and 

others 1999, Krüssmann 1986). American 

snowbell is distinguishable from bigleaf snowbell 

(S. grandifolius) by its smaller leaves and shorter 

flowering racemes with fewer flowers per raceme 

(Brown and Kirkman 1990). 

Range.—American snowbell is found in the 

Southeastern United States, from Virginia south to 

Florida and west to Louisiana (Bailey and Bailey 

1976) and eastern Texas. It is also recorded in 

McCurtain County in Oklahoma (Johnson and 

Hoagland 1999). It will grow as far north as 

southern Missouri, southern Illinois, Indiana, and 

southern Ohio but is rare, endangered or possibly 

extirpated in Ohio and Illinois (Ohio Department 

of Natural Resources 2002, Iverson and others 

1999). 

Ecology.—American snowbell grows primarily in 

rich, poorly-drained acidic soils along swamp 

margins, stream beds and lake edges, oxbow lakes, 

and in floodplains. (Johnson and Hoagland 1999). 

It grows well in the shade but will tolerate full sun. 

It is associated with trees commonly found in wet 

areas, such as baldcypress and overcup oak. Plants 

are particularly tender when young (Krüssmann 

1986). 

Reproduction.—American snowbell is named for 

its showy, fragrant, white flowers. Flowers are 

complete and their placement hypogynous 

(Iverson and others 1999). They occur in 2.5 to 12 

cm long racemes, one to four flowers per raceme. 

Raceme stalks are pubescent. The flowers have 

five petals, and the white style extends beyond the 

10 stamens. The calyx is shallowly five-lobed and 

has tiny, triangular teeth. The 6 to 8 mm 

subglobose fruits are one-seeded drupes that 

mature from July to October (Krüssmann 1986, 

Johnson and Hoagland 1999). 

Growth and Management.—This species is often 

planted and grows well in cultivation. Since it 

requires moist soils and prefers shade, it can be 

seriously threatened by drainage and removal of 

the forest canopy. Its major pollinators are bees 

(Iverson and others 1999), and the species relies 

on seed dispersal for regeneration. It is, however, 

easily propagated from softwood cuttings (Johnson 

and Hoagland 1999). 
 

738 

Benefits.—American snowbell is an attractive 

landscape plant. Its leaves are also a preferred food 

of the caterpillar of Callosamia promethea, the 

promethea moth (Oehlke [no date]). 

References

Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: 

A Concise Dictionary of Plants Cultivated in the 

United States and Canada. McMillan Publishing 

Co., Inc., New York. 1,312 p. 

Brown, C.L. and L.K. Kirkman. 1990. Trees of 

Georgia and adjacent states. Timber Press, 

Portland, OR. 292 p. 

Iverson, L.R., D. Ketzner and J. Karnes. 1999. 

Styrax americanus. Illinois Plant Information 

Network. Database at 

http://www.fs.fed.us/ne/delaware/lpin. html. 

Illinois Natural History Survey and USDA Forest 

Service. 3 p. 

Johnson, F.L. and B.W. Hoagland. 1999. Styrax 

americanus. Catalog of the woody plants of 

Oklahoma, Oklahoma Biological Survey Home 

Page, http://www.biosurvey.ou.edu. 1 p. 

Krüssmann, G. 1986. Manual of cultivated broadleaved trees and shrubs. Volume III, Pro-Z. 

Timber Press, Beaverton, OR. 510 p. 

Oehlke, B. [no date]. The Promethea Moth 

Callosamia promethes (Drury, 1773). 

http://www3. 

islandtelecom.com/~oehlkew/scpromet.htm. 5 p. 

Ohio Department of Natural Resources. 2002. 

Styrax americanus. Ohio Division of Natural 

Areas and Preserves. Rare Native Ohio Plant 

2000-2001 Status List. 2 p. 

Plant Identification Resource. 2001. Styrax 

americanus. Auburn University College of 

Agriculture, Landscape/Horticulture webpage, 

http://www.ag.auburn.edu/landscape/386.html. 

3 p. 
 

739

Suaeda suffrutescens S. Wats. desert seepweed 

CHENOPODIACEAE 

Synonyms: Dondia suffrutescens (Wats.) Heller 

 Suaeda nigrescens I.M. Johnst. 

James E. Nellessen 

 

General Description.—Desert seepweed is also 

known as inkweed, iodinebush, and quelite salado. 

Desert seepweed has stout stems and branches, 

woodiness greatest toward the base, side branches 

generally short, although the plant is highly 

branched. It could be classified as a subshrub. The 

plant is erect and ascending, or spreading. Desert 

seepweed ranges in height from about 50 to 120 

cm, often forming mounds as broad as 150 cm. 

Young stems and leaves are soft and felty with 

numerous hairs. Leaves are simple, alternate, 

numerous, fleshy, linear to linear-lanceolate, 5 to 

20 mm long, narrower at the base, semirounded to 

somewhat flat (Correll and Johnston 1970, Great 

Plains Flora Association 1986, Ivey 1995, Kearney 

and others 1951, Martin and Hutchins 1980). Two 

varieties have been recognized: S. suffrutescens

var. suffrutescens, and S. suffrutescens var. 

detonsa I.M. Johnston. The latter variety (detonsa) 

has three synonyms: (1) S. duripes I.M. Johnst., 

(2) S. nigrescens I.M. Johnst., and (3) S. 

nigrescens var. glabra I.M. Johnst. (Allred 2002, 

Great Plains Flora Association 1986, Kartesz 

1994). The latter variety, S. suffrutescens var. 

detonsa (S. nigrescens var. glabra) is distinguished 

by having smooth, or relatively smooth (hairless) 

stems and leaves. This variety is more common in 

Mexico, but extends into southern Texas. 

Range.—Desert seepweed occurs from western 

Oklahoma and Texas across central and southern 

New Mexico into Arizona and California, and 

south into Mexico. It occurs at elevations from 

about 750 m to 1,830 m. 

Ecology.—Desert seepweed is a perennial shrub 

of saline or alkaline soils in plains and valleys. 

These habitats include the margins of ephemeral 

lake beds, salt flats, and flood plains of rivers and 

streams. Some moisture needs to be relatively near 

the surface for part of the growing season. This 

species is classified as a halophyte, or salt tolerant 

plant. Desert seepweed may dominate alkali sink 

habitats in association with other salt tolerant 

species, including other species of Suaeda

(seepweeds), Salicornia spp. (glasswort or 

saltwort), Distichlis spicata (L.) Green var. stricta

(Torr.) Beetle (inland saltgrass), Allenrolfea 

occidentalis (Wats.) Kuntze (burro weed or 

quinine bush), and Heliotropium curassavicum L. 

(salt heliotrope), to name a few (Hendrickson 

1974, Dick-Peddie 1993). 

Halophyte Ecology and Physiology.—Although 

little if anything is reported about desert seepweed, 

much is known about the common salt tolerant 

characteristics of the genus Suaeda. Chapman 

(1942) used the term halophyte in describing 

species that could grow in an environment with 

greater than 0.5 percent sodium chloride (NaCl). 

Halophytes can be divided into two groups: (1) 

those requiring salt for survival and best growth, 

and (2) those tolerating or resisting salty 

environments (Waisel 1972). The second group, 

the salt tolerators, have adapted to saline habitats 

and survive there because salt-sensitive plants 
 

740 

(glycophytes) cannot survive there. Halophytes 

accumulate salts obtained from the soil solution 

within their tissues during the transpiration 

process. Many members of the genus Suaeda

readily translocate salts to upper portions of the 

plant. This second group of halophytes can be 

subdivided into three more categories of tolerance. 

Salt enduing species simply tolerate the presence 

of salt in their cells. Salt excluding species may 

excrete salt from shoots, including those with 

specialized hairs on the plant, or transport them 

back to the roots. Salt evading species have 

mechanisms to prevent transport to the leaves or 

prevent absorption from the soil solution (Waisel 

1972). Some members of the genus Suaeda fall 

into the first group requiring salt for survival, 

while others are salt enduring halophytes. Salt 

(NaCl) and other ions (including sulfate and 

potassium) may be sequestered into cell vacuoles 

where they do not interfere with normal cell 

biochemistry. These plants will generally have 

higher water potential gradients between the soil 

and the leaves, which may add to succulence. 

Waisel (1972) proposed a new order to the 

classification of halophytic plant communities: 

Suaedetalia--persistent communities in dry desert 

saline habitats, dominated by annual and perennial 

members of the family Chenopodiaceae (as well as 

other families). Zahran (1982) likewise includes a 

Suaeda community classification. Some shrubby 

members of the genus (S. fruticosa Forsk. and S. 

monoica Forsk.) are reported to accumulate large 

amounts of salt annually when in dense stands, as 

much as 2 to 4 tons/ha (Waisel 1972). In this 

function they may help to desalinate some saline 

habitats. 

Reproduction.—Flowers are small and greenish, 

and occur in spiked clusters of three to nine 

flowers in the leaf axils. The corolla is absent. The 

five-lobed calyx has all parts equal, may be hood 

shaped, and lacks appendages, that is, it is not 

winged. Seeds are small, black, and 0.7 to 1 mm 

wide. Desert seepweed blooms from March to 

September, depending on local environmental 

conditions (Correll and Johnston 1970, Great 

Plains Flora Association 1986, Kearney and others 

1951, Martin and Hutchins 1980). Many of the salt 

tolerant members of the Chenopodiaceae have 

physiological seed dormancy (Baskin and Baskin 

2001). Although no seed germination reports 

regarding this particular species were located, 

numerous studies have been performed on other 

members of the Suaeda genus. Three of four 

members of this genus required prior cold 

stratification of the seeds with optimal daytime 

germination temperatures of 25 to 27 °C 

(summarized in Baskin and Baskin 2001). NaCl 

concentrations ranging from 0.17 to 0.90 molar 

have depressed maximum germination of 75 to 

100 percent down to 10 percent in four different 

species of Suaeda (summarized in Baskin and 

Baskin 2001). 

Growth and Management.—Desert seepweed 

can grow relatively quickly. It has been observed 

to resprout readily in managed and mowed areas 

adjacent to levees along the lower Rio Grande 

floodplain in New Mexico. This species is 

restricted to saline and riparian habitats and is not 

highly browsed. Consequently, it does not pose as 

a major management concern on most rangelands. 

It does not typically form dense stands. 

Benefits.—Desert seepweed leaves and young 

shoots have been added to vegetable greens or 

cacti fruits by Native Americans during cooking to 

add flavor. It has also been added between layers 

of cacti fruit and sotol butts (stems of Dasylirion 

sp.) during the 24 to 48 hour cooking process in 

mescal pits (Warnock 1974). Pima Indians would 

eat the plant for greens and the roasted seeds, 

called pinole. Coahuila Indians extracted a black 

dye from the plant used for artwork (Dodge 1985). 

This species is not browsed much by livestock but 

may be utilized when other vegetation is in short 

supply (Warnock 1974, Dodge 1985). 

References

Allred, K.W. 2002. A Working Index of New 

Mexico Vascular Plant Names. http://web. 

nmsu.edu/~kallred/herbweb. [not paged]. 

Baskin, C.C. and J.M. Baskin. 2001. Seeds: 

Ecology, Biogeography, and Evolution of 

Dormancy, and Germination. Academic Press, 

Sand Diego, CA. 666 p. 

Chapman, V.J. 1942. The new perspective in the 

halophytes. Quarterly Review of Biology 17: 

291-311. 

Correll, D.S. and M.C. Johnston. 1970. Manual of 

the Vascular Plants of Texas. Texas Research 

Foundation, Renner, TX. 1,881 p. 
 

741

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation: Past, Present and Future. 

University of New Mexico Press, Albuquerque, 

NM. 244 p. 

 Dodge, N.N. 1985. Flowers of the Southwest 

Deserts. Southwest Parks and Monuments 

Association, Tucson, AZ. 136 p. 

Great Plains Flora Association. 1986. Flora of the 

Great Plains. University Press of Kansas, KS. 

1,392 p. 

Hendrickson, J. 1974. Saline habitats and 

halophytic vegetation in the Chihuahuan Desert 

region. In: R.H. Wauer and D.H. Riskind, eds. 

Transactions of the Symposium of the 

Biological Resources of the Chihuahuan Desert 

Region, United States and Mexico. U.S. 

Department of Interior, National Park Service 

Transactions and Proceedings Series, No. 3, 

U.S. Goverment Printing Office, Washington 

DC. p. 289-314. 

Ivey, R.D. 1995. Flowering Plants of New 

Mexico, 3rd Edition. Published by the author, 

NM. 504 p. 

Kartesz, J.T. 1994. A Synonymized Checklist of 

the Vascular Flora of the United States, Canada, 

and Greenland, Vol. 1, 2nd Ed. Biota of North 

America Program of the North Carolina 

Botanical Garden. Timber Press, Portland, OR. 

622 p. 

Kearney, T.H., R. Peebles, and Collaborators. 

1951 with 1960 supplement. Arizona Flora. 

University of California Press, Berkeley, CA. 

1,085 p. 

Martin, W.C. and C.E. Hutchins. 1980 (reprinted 

2001). A Flora of New Mexico, Vol. 1. Bishen 

Singh Mahendra Pal Singh (India) and Koeltz 

Scientific Books (Germany). 1,276 p. 

 

Waisel, Y. 1972. Biology of Halophytes. 

Academic Press, NY. 395 p. 

Warnock, B.H. 1974. Wildflowers of the 

Guadalupe Mountains and the Sand Dune 

Country, Texas. Sul Ross State University, 

Alpine, TX. 176 p. 

Zahran, M.A. 1982. Ecology of halophytic 

vegetation of Egypt. In: D.N. Sen and K.S. 

Rajpurohit, eds. Contributions to the Ecology of 

Halophytes.. Dr. W. Junk Publishers, The 

Hague, Netherlands. p. 3-20. 
 

742 

Suriana maritima L. bay-cedar 

SURIANACEAE (formerly in SIMAROUBACEAE) 

Synonyms: none 

John K. Francis 

General Description.—Bay-cedar, is also known 

as tassel-plant, guitarán, temporana, jovero, 

cuabilla, pantsil, cucharo, palo corra, oseille bord 

de mer, romarin noir, and crises marine. It is an 

evergreen shrub usually 1 to 2 m in height but 

sometimes reaching 6 m. Multistemmed and 

much-branched, it has a prostrate to ovoid crown. 

Stems have a dark brown, rough and flakey bark. 

Heartwood is dark red or reddish brown; the 

sapwood being somewhat lighter. The wood is 

hard, heavy, fine-textured, strong, and durable. 

The gray-green, succulent, downy foliage has a 

salty taste and a cedar-like fragrance when 

crushed. Leaves are crowded at the ends of the 

branches, sessile, narrowly obovate, and 1 to 4 cm 

long. Inflorescences, solitary or few-flowered 

cymes, are almost hidden among the leaves. The 

small, five-merous, bisexual flowers are yellow 

and develop into clusters of five hard, 3- to 4-mm, 

dry drupes surrounded by five gray sepals. 

(Gilman 1999, Howard 1988, Liogier 1988, Little 

and others 1974). 

Range.—Bay-cedar is native to the shores of 

southern Florida, Bermuda, the Bahamas, the West 

Indies, the eastern side of Mexico through Brazil, 

East Africa, tropical Asia, Australia, and many of 

the islands of the Indian and Pacific Oceans (not 

Hawaii) (Howard 1988, Stevens and others 2001). 

The species has been planted widely as an 

ornamental. It is listed as endangered in the wild in 

Florida (Smithsonian Institution 2002). 

Ecology.—Bay-cedar is strictly costal. It grows on 

beaches, dunes, sandy thickets (Nelson 1996), and 

rocky headlands. It tolerates moderately salty soils, 

storm-surge overwash, heavy salt spray, blowing 

sand, high surface heat, drought, and strong winds 

(Wildflower Nervana 2002). If planted and given 

protection from competition, it will grow well on 

inland sites (Nelson 1996) on a wide range of 

well-drained soils. The species requires full sun to 

grow and flower. Bay-cedar is rare to uncommon 

in much of its habitat, but occasionally forms 

thickets. 

Reproduction.—Bay-cedar flowers and fruits 

intermittently throughout the year (Little and 

others 1974). The fruits are buoyant and remain 

viable for long periods in sea water enabling 

colonization of tropical beaches throughout the 

world (Nelson 1996). 

Growth and Management.—Growth rate of baycedar is described as moderate. The species is 

currently being propagated by seeds to produce 

nursery plants. The recommended spacing for 

ornamentals is 90 to 150 cm. Irrigation is 

recommended during establishment but afterwards 

is not needed (Gilman 1999). 

Benefits.—Bay-cedar helps stabilize beaches and 

costal dunes, and furnishes food and cover for 

wildlife. It is widely planted as an ornamental for 

hedges, screens, borders, accents, seminatural 

covers, and potted plants. The fruits attract birds 

that feed on them (Gilman 1999). The foliage 

furnishes food for blue and hairstreak butterfly 

larva (Schaefer and others 2002). Key deer 

(Odocolleus virginanus clavium) and probably 

other ruminants will not eat the foliage of baycedar (Schaus and others 2002). Herbalists employ 
 

743

extracts of the leaves and bark to treat rheumatism 

and skin ulcers and to stop bleeding (Liogier 

1990). 

References

Gilman, E.F. 1999. Suriana maritima. Cooperative 

Extension Service, University of Florida, 

Gainesville, FL. 3 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A. 1990. Plantas mediciales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washingtion, DC. 1,024 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Schaefer, J., C.N. Huegel, and F.J. Mazzotti. 2002. 

Butterfly gardening in Florida. Cooperative 

Extension Service, University of Florida, 

Gainesville, FL. http://edis.ifas.ufl.edu/ 

BODY_UW057. 24 p. 

Schaus, C, S. Wade, and J. Dunan. 2002. Key deer 

and plants they won’t eat. Monroe County 

Extension Service, Key West, FL. 

http://monroe.ifas.ufl.edu/key_deer_plants.htm. 

4 p. 

Smithsonian Institution. 2002. Special status 

species in the Indian River Lagoon. Smithsonian 

Marine Station at Fort Pierce, Smithsonian 

Institution, Washington, DC. http://www. 

sms.si.edu/IRLSpec/ListedSpec.htm. 3 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora of Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

3. Missouri Botanical Garden Press, St. Louis, 

MO. p. 1,911-2,666. 

Wildflower Nervana. 2002. Suriana maritima—

guitarán or temporana (bay-cedar). http:// 

www.wfnirvana.com/pr/surimari/html. 3 p. 
 

744 

Symphoricarpos albus (L.) Blake common snowberry 

CAPRIFOLIACEAE 

Synonyms: Symphoricarpos racemosus Michx. 

 Symphoricarpos rivularis Suksdorf 

 Viburnum album L. 

John K. Francis 

General Description.—Common snowberry, also 

known as snowberry, waxberry, and white 

coralberry, is a deciduous shrub. The slender, 

branchy, multiple stems are usually low (0.5 to 1 

m) and arching or straight, but sometimes reaching 

as much as 2 m in height. The stems rarely exceed 

2.5 cm in thickness. The bark is yellow-gray in 

young stems, tan to grayish-brown in older ones, 

becoming scaly or splitting lengthwise in larger 

stems. Stem wood is soft and greenish-white. The 

opposite-branching twigs are slender and flexible. 

Thin, papery leaves are opposite, simple, shortstalked, and round to oblong-elliptic with margins 

entire, lobbed, or wavy-toothed. Petioles are 3 to 4 

mm long. The perfect flowers are solitary to 

several in terminal or lateral spikes. The fivelobbed corolla is pink and white, tubular or bellshaped, and about 6 mm long. Fruits are white, 

spongy, berry-like drupes 6 to 12 mm in diameter. 

Each contains two white, flattened, elliptical 

nutlets with one seed per nutlet. Chromosome 

number is 2n = 54 or 72 (Gilbert 1995, Grimm 

1966, McWilliams 2000, Soper and Heimburger 

1982, Stephens 1973, Welsh and others 1987). 

Range.—Common snowberry is native from Nova 

Scotia to southern Alaska, throughout the northern 

half of the United States, and in California, Utah, 

Colorado, New Mexico, Tennessee, and North 

Carolina (McWilliams 2000, Natural Resources 

Conservation Service 2003). It does not occur in 

Nevada and Kansas, and is reported to be only 

naturalized in Utah (Welsh and others 1987). The 

species was introduced to Great Britain in 1817 

and has naturalized widely in the United Kingdom, 

Ireland, and northern Europe (Binggeli 1998, 

Gilbert 1995). There are two varieties, albus and 

laevigatus (Fern.) Blake. Variety laevigatus, a 

coarser and taller (up to 2 m) plant, and has a 

Western distribution. Variety albus occurs 

intermittently across the Continent (Natural 

Resources Conservation Service 2003). Both are 

cultivated and naturalized in many places, making 

determination of distribution somewhat 

complicated. 

Ecology.—Common snowberry grows best in full 

sunlight or light shade on well-drained, moist, 

fertile soils. It does survive and grow slowly in 

medium shade. The species grows in many types 

of open forests, forest edges, shrublands, riparian 

vegetation, and fencerows. It does not grow in 

swampy sites, deserts, and mountain tops above 

the tree line. Common snowberry grows in soils 

with mildly acidic pH (6.0) and mildly alkaline 

conditions (pH’s of 7.8) and does very well on 

soils derived from limestone; less aggressively on 

soils derived from granite. It tolerates poor fertility 

and has been used to revegetate disturbed sites. 

Common snowberry in interior Western United 

States grows at elevations from 800 to 2,800 m. It 

grows at much lower elevations in other areas of 

North America. Because it sprouts readily from 

rhizomes, common snowberry is resistant to fire 

and browsing. It may increase dramatically in 

density after logging opens the forest canopy 

(Gilbert 1995, McWilliams 2000, Natural 

Resources Conservation Service 2003). 
 

745

Reproduction.—Common snowberry blooms 

between early May and late July depending on 

location and matures fruits in August to October 

(Grimm 1966). The flowers are pollinated by a 

range of bees, wasps, and syrphids (Gilbert 1995). 

Fruit and seed production can be abundant, 

especially when plants grow in full sun on fertile 

sites. Normally a good seed crop is produced every 

year. Fruits remain on stems until midwinter or 

early spring, if not eaten. Numbers of seeds per 

gram vary from 119 to 250 for var. albus and from 

86 to 144 for var. laevigatus. Germination may be 

as high as 87 percent if exacting conditions of 

pretreatment are met. Germination is epigeal 

(Walker 2003). Seeds are dispersed by birds and 

mammals. Seedlings are relatively uncommon. 

However, vegetative reproduction is common by 

sprouting of underground stems that run 

horizontally at a depth of 2 to 5 cm for up to 60 cm 

before turning up to form new tops. Stems layer 

when they come in contact with the ground, and 

severed stems lying on the ground will also take 

root (Gilbert 1995). 

Growth and Management.—Planted common 

snowberry reached 30 cm in height during the first 

year (Center for Urban Horticulture 2002). Shoot 

extension of plants in England is reported at 0.6 m 

per year. Individual stems have been aged up to 34 

years old in England (Gilbert 1995). By suckering, 

plants may survive indefinitely. Fruits may be 

collected by hand picking or flailing the plants 

with a tarp spread underneath. The fruit is then 

macerated and water is used to float off the pulp 

and empty seeds. The residue is then dried and 

cleaned. The air-dried seeds can be stored at 5 °C 

for up to 5 years. Pretreatments are necessary for 

acceptable germination. Warm stratification for 3 

to 4 months at 22 to 30 °C followed by cold 

stratification at 5 °C for 4 to 6 months is 

recommended. Sowing in the late fall can be used 

instead of the cold treatment. Sown seeds should 

be covered with about 6 mm of soil and 2 cm of 

mulch. Seedlings for planting can be grown 

successfully from cuttings, wildlings, and pieces of 

stem with roots collected in the spring. Transplant 

establishment success of bareroot and 

containerized nursery stock is as high as 90 

percent when done properly (Walker 2003). 

Benefits.—Common snowberry helps protect the 

soil, provides food and cover for wildlife, browse 

for livestock, and adds beauty to the landscape, 

especially in fall and winter when the white fruits 

are on the shrubs. Containing 4 to 13 percent 

protein, depending on tissue and season, it 

provides important early-season browse for cattle 

and domestic sheep. It is also browsed by most 

wild ungulates. It is generally unpalatable to 

horses and moose. The fruits are consumed by 

both black and grizzly bears and are eaten by a 

number of birds and small mammals (McWilliams 

2000). Some sources refer to the fruits as being 

toxic to humans (Gilbert 1995, Moerman 1986). 

The toxic agent appears to be the isoquinoline 

alkaloid chelidonine and causes mild symptoms of 

vomiting, dizziness, and sedation in children 

(Canadian Biodiversity Information Facility 2003). 

The fruits were reported to have been eaten but not 

favored by Native Americans (McWilliams 2000). 

Various tissues were used by Native Americans as 

a diuretic, to treat gonorrhea, and for sore eyes 

(Moerman 1986). Common snowberry is planted 

as an ornamental and in conservation plantings for 

wildlife amenity, soil stabilization, and strip mine 

reclamation in temperate areas around the world 

(Gilbert 1995). 

References

Binggeli, P. 1998. Plants of the Pacific Northwest 

in Western Europe. Botanical Electronic News 

ISSN 1188-603X. http://www.ou.edu/cas/ 

botany-micro/ben/ben195.html. p. 1-4 

Canadian Biodiversity Information Facility. 2003. 

Notes on poisoning: Symphoricarpos albus. 

http://sas.agr.gc.ca/pls/pp/ppack.info?p_psn=122

&p_type=all&p_sci=sci&p_x=px. 5 p. 

Center for Urban Horticulture. 2002. Does mulch 

improve plant survival and growth in restoration 

sites? Fact sheet 38. University of Washington, 

College of Forest Resources. http://www.cfr. 

washington.edu/Research/fact_sheets/38-mulch. 

pdf. 2 p. 

Gilbert, O.L. 1995. Symphoricarpos albus (L.) 

S.F. Blake (S. rivularis Suksd., S. racemosus

Michaux). Journal of Ecology 83(1): 159-166. 

 

Grimm, W.C. 1966. Recognizing native shrubs. 

The Starkpole Company, Harrisburg, PA. 319 p. 

McWilliams, J. 2000. Symphoricarpos albus. In: 

U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory. Fire Effects Information System. 
 

746 

http://www.fs.fed.us/database/feis/plants/shrub/ 

symalb/all.html. 32 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports 19. University of 

Michigan Museum of Anthropology, Ann 

Arbor, MI. 534 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Symphoricarpos albus (L.) Blake 

http://plants.usda.gov/cgi_bin/plant_profile. 

cgi?symbol =SYAL. [not paged]. 

Soper, J.H. and M.L. Heimburger. 1982. Shrubs of 

Ontario. Royal Ontario Museum, Toronto, 

Ontario, Canada. 495 p. 

Stephens, H.A. 1973. Woody plants of the North 

Central Plains. The University Press of Kansas, 

Lawrence, KS. 530 p. 

Walker, S.C. 2003. Symphoricarpos Duham., 

snowberry. U.S. Department of Agriculture, 

Forest Service, Washington, DC. http://wpsm. 

net/Symphoricarpos.pdf. 10 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 
 

747

Tabebuia haemantha (Bertol. ex Spreng.) DC. roble cimarrón 

BIGNONIACEAE 

Synonyms: Bignonia haemantha Bert. ex Spreng. 

 Tecoma haemantha (Bert.) Griseb. 

 Spathodea portoricensis Bello 

John K. Francis 

General Description.—Roble cimarrón, also 

known as roble colorado and roble bobo, is a 

shrub or a small tree up to 8 m in height and 15 

cm in basal diameter. Commonly, plants are 3 to 5 

m in height and 4 to 8 cm in diameter. Young 

roble cimarrón usually have a single stem, until 

they have been damaged mechanically or by fire. 

Older plants often develop multiple stems by 

spontaneously sprouting just above the ground 

level. The bark is gray, smooth except for a slight 

fissuring. The inner bark is nearly white and 

slightly bitter. There are relatively few branches. 

The twigs are light gray and slightly flattened 

below the nodes. The species develops a tap and 

lateral root system. The roots are brown and 

flexible. Roble cimarrón is evergreen with 

palmately compound leaves. There are three or 

five stiff, leathery leaflets on a stout 2.5- to 5.0-cm 

petiole. The 3- to 15-cm long leaf blades are 

elliptic or ovate with entire edges and rounded to 

pointed at the tip. The inflorescences are panicles 

of several to many flowers on short branches. The 

flowers have a crimson, tubular corolla 3 to 5 cm 

long with five irregular lobes. The capsules are 6 

to 11 cm long and contain many membranous twowinged seeds 1.9 cm long (author’s observations, 

Liogier 1995, Little and Wadsworth 1964). 

Range.—Roble cimarrón is endemic to the island 

of Puerto Rico. Specifically, the species is found 

in the dry forests in the southern and eastern parts 

of the island and the moist slopes above them, 

including serpentine areas in western Puerto Rico, 

and on the limestone hills in the northern coast 

and inland (Little and Wadsworth 1964, Vélez and 

van Overbeek 1950). 

Ecology.—Roble cimarrón grows on loamy and 

clayey soils with pH’s from about 5.5 to 7.5. It 

often occurs in sites that have been eroded and 

partially compacted by grazing animals. The soils 

where it grows today are usually steep and rocky. 

The species formerly inhabited all types of 

topography from near sea level to about 400 m in 

elevation. These sites receive from about 700 to 

2100 mm of precipitation. In areas that receive the 

higher rainfall of the range, roble cimarrón tends 

to grow on excessively drained sites. The species 

is frequently common in overgrazed pastures 

indicating that it is unpalatable to cattle. Roble 

cimarrón is moderately intolerant of shade. It 

grows well in the open and in low basal area forest 

stands. Adult plants are usually found in 

codominant and intermediate crown positions in 

dry forests. It is top-killed by fires but sprouts 

vigorously afterwards. 

Reproduction.—Flowering occurs almost 

throughout the year according to Little and 

Wadsworth (1964). However, most shrubs in 

particular areas appear to flower synchronously 

(author’s observation). Hummingbirds, other bird 

species, and insects pollinate the flowers. Capsules 

form and mature about 6 weeks after flowering. 

They are green, turning tan at maturity, after 

which they quickly split open. The seeds adhere to 

a flat central placenta but detach and become 

airborne in the slightest breeze (Vélez and van 

Overbeek 1950). A sample of 17 capsules 
 

748 

collected in Salinas, Puerto Rico contained an 

average of 59.9 ± 2.01 seeds/capsule. The air-dry 

seeds averaged 0.0148 + 0.0003 g/seed or 68,000 

seeds/kg. Sown on peat moss, these seeds 

germinated at 90 percent between 7 and 31 days 

after sowing. The species grows and survives well 

in the nursery, similar to other Tabebuia seedlings. 

In the wild, seedlings are well scattered and 

uncommon. 

Growth and Management.—Roble cimarrón 

grows about 0.5 m/yr in early years after it is well 

established. It does not appear to be a long-lived 

species. Unfortunately, no plantations are known 

and no management experience has been reported. 

Benefits.—Roble cimarrón would probably be 

suitable as an ornamental for background 

plantings in moist and dry frost-free areas. It 

would have the advantage of being drought-hardy 

and tolerant of infertile and partially compacted 

soils and should be a good species to plant in 

environmental restoration plantings within its 

native range. The light brown, hard wood is 

suitable for firewood and charcoal but is little used 

because of the plant’s small size and relative 

scarcity. 

References 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

617 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service. 

Washington, DC. 548 p. 

Vélez, I. and J. van Overbeek. 1950. Plantas 

indeseables en los cultivos tropicales. Editorial 

Universitaria, Río Piedras, PR. 497 p. 
 

749

Tamarix chinensis Lour. tamarisk

TAMARICACEAE 

Synonyms: Tamarix pendandra Pallas 

 Tamarix ramosissima Ledebour 

 Tamarix gallica L. var. micrantha Ledebour 

 Tamarix pentandra Pallas ssp. tigrensis (Bge.) Hand.-Mazz. 

Juanita A.R. Ladyman 

General Description.—Tamarix chinensis

Loureiro is commonly known as tamarisk or 

saltcedar. Occasionally it grows with a defined 

trunk as a tree but generally it grows as a multiplestemmed shrub up to 6 m tall. The bark of larger 

trunks, or stems, is reddish-brown to dark brown 

or blackish. The leaves, that are only 1.5 to 3.5 

mm long and appear scale-like, are deciduous. The 

tiny flowers are five-merous, having five sepals, 

five petals, five stamens, and a five carpellate 

pistil, and are arranged on a flowering stalk that is 

2 to 7 cm long. The petals are 1 to 2.3 mm long, 

and persistent in fruit. They are frequently a shade 

of pink but may be white or red. The sepals are 0.5 

mm to 1.3 mm long. The scale-like leaves and 

shape of the shrub are reminiscent of a juniper or 

cedar and this accounts for the common name 

“saltcedar.” This shrub has been the subject of 

considerable taxonomic confusion. McClintock 

(1951) stated “there is probably not another genus 

of plants as well known as the tamarisks in which 

the species is so poorly understood or separated on 

more obscure characters.” For most species, the 

tiny floral organs and the use of a microscope, or 

at least hand lens, are essential for identification 

(Allred 2002). Tamarix pentandra var, pentandra

Pallus has been mistakenly applied to this species 

within North America. Actually, T. pentandra

Pallus is synonymous with T. hispida Willd. that is 

distributed throughout central Asia (Baum 1978). 

Other confusion has resulted from this species 

being referred to as a sub-species of T. gallica L., 

which is a unique taxon originally from Europe, 

and now also naturalized within the United States 

(Allred 2002, Baum 1978). Although Baum’s 

treatment (1978) described T. chinensis Loureiro 

and T. ramosissima Ledebour as unique taxa 

placing them in two different sections, recent 

chloroplast and nuclear DNA studies suggest the 

two entities should be merged (Allred 2002). 

These two names are synonymized in the Utah 

Flora (Welsh and others 1993). Tamarix chinensis

has a chromosome number of 2n = 24 (Welsh and 

others 1993). 

Range.—Tamarisk is a common, wide-ranging 

species that has been collected across Asia from 

western Turkey through Mongolia, Afghanistan, 

and China to Korea and Japan. It was introduced 

into the Western United States and has become 

naturalized. 

Ecology.—Tamarisk grows in generally moist 

sites at seeps, streams, and along river banks and 

roadsides (Dick-Peddie 1993). It tolerates saline 

soils and also contributes to salinization of the 

soils in which it grows. It forms equally dense 

stands in both the Western United States and in 

central China where it is an apparent native 

(author’s personal observation). It has an extensive 

root system and uses such large volumes of water 

that streams and ponds become dry. Studies in the 

Southwestern United States show that tamarisk 

stands have far less biodiversity than stands of 

native riparian vegetation. Miles of riverbank can 

be almost monotypic with dense tamarisk stands 
 

750 

that have 50 percent fewer small mammal species 

as well as fewer reptile and amphibian species than 

native riparian stands (Olson 1999). Although bees 

visit the flowers (Epple 1995) the diversity of 

insects is also low, and nestling birds rely on 

insects brought to the nests by their parents. 

Although some birds can nest in the shrubs the 

trunks are not large enough to provide habitat for 

animals such as wood-peckers, owls, chickadees, 

and squirrels. The tiny, hairy seeds provide poor 

nutrition and are largely indigestible. 

Reproduction.—Tamarisk generally flowers in 

April through August. The pink, fragrant flowers 

are insect pollinated. The seeds, which are small, 

mature in summer and fall and are dispersed by 

birds and by water. However, the seed may have 

limited longevity under humid conditions. Seeds 

retain high viability if stored in sealed containers 

with desiccant at temperatures between 3 to 27 oC 

but lose viability if stored at “moderate humidity” 

(Wilgus and Hamilton 1962, Baskin and Baskin 

2001). Germination temperature appears to be 

flexible. Freshly matured seeds germinated from 

87 to 98 percent when kept at constant 

temperatures between 19 to 43 o

C. Percent 

germination did not appear related to temperature; 

96 percent germinated at 19 o

C and 91 percent at 

43 oC (Wilgus and Hamilton 1962, Baskin and 

Baskin 2001). Vegetative reproduction by suckers 

and rooting branches is copious. 

Growth and Management.—Tamarisk is an 

aggressive, invasive species. Its invasive behavior 

is illustrated by its colonization of the Pecos River 

Valley in New Mexico (Allred 2002). In 1912, a 

few seedlings were observed at Lake McMillan. 

By 1915 the shrub covered 600 acres and extended 

up and down the riverbanks. By 1926 it covered 

12,300 acres and extended to 57,000 acres by 1960 

(Allred 2002). It out-competes many native 

species and is difficult to eradicate once 

established (USDA Forest Service 1988). The 

extremely deleterious impact of this species on the 

riparian and wetland areas of the Southwest has 

led to intense study on how to control and reverse 

its spread. There are four general methods--

biological control, physical removal by hand or by 

machine, herbicides, and fire--that can be used to 

control tamarisk, and each has shortcomings. The 

most appropriate method, or combination of 

methods, depends upon the particular site. 

Biological controls for tamarisk are not yet 

commercially available, although a variety of 

insects have been investigated--from grasshoppers 

to leaf hoppers. Two species specific to tamarisk, a 

mealybug (Trabutina mannipara) from Israel and 

a leafbeetle (Diorhaba elongata) from China, have 

received advance testing (DeLoach and others 

1996, DeLoach, 1994). Physical cutting and 

removal of tamarisk will only be successful if the 

roots are removed or killed. By itself, cutting 

tamarisk by hand (that is by chain saw, weed eater, 

or axe) is not often an effective treatment. Cutting 

tamarisk tends to just cause more shoots to grow. 

However, there was a report of successful control 

where large tree stumps (20 to 25 cm diameter) 

had been cut back to the soil surface and then 

covered by a black, liner material (Cinnamon 

1990). Purely mechanical removal is also 

sometimes successful if there is enough standing 

vegetation to substantially shade the resprouting 

tamarisk. For example, in Nevada (Busch and 

Smith, 1995) when tamarisk had been completely 

cleared above ground in a stand of willows, there 

was essentially no grow back during the 4 years 

follow up. Tamarisk is sensitive to numerous 

herbicides (Kunzman and Bennett 1990, Sisneros 

1991). At the present time, the most successful 

approach to control tamarisk is by cut-stump 

herbicide or cut-stump/frill herbicide treatments 

(Neill 1990, Hughes 1996). This approach is quite 

costly but it is very controlled, does not affect nontarget organisms, and causes the least 

environmental impact. The tamarisk shrubs should 

be cut to within 2.5 cm of the ground surface 

(Cinnamon 1990) and a systemic herbicide applied 

to the stump within a few minutes after cutting. 

The time between cutting and herbicide treatment 

should be as short as possible. Leaving the cut 

stump for 2 to 5 days before herbicide treatment 

reduces success rate from greater than 

approximately 88 percent tamarisk death to less 

than 10 percent (Hays and Mitchell 1990). Fire is 

often ineffective. The high water and salt content 

of tamarisk makes it difficult to burn. Too light 

(low temperature) a fire will permit, or even 

encourage, tamarisk to re-sprout and become even 

denser. If the fire burn is so hot it kills the 

tamarisk, it is likely that the fire will leave the soil 

bare and in such a poor condition that it cannot 

support grasses and desirable vegetation. In the 

horticultural trade, tamarisk is propagated from 

hardwood cuttings in the spring or softwood 

cuttings in the summer (Tykač 1990). Plants 

grown for a long time in containers often do not 

establish after transplanting (Tykač 1990). 
 

751

Benefits.—Tamarisk were once planted for 

erosion control and as wind breaks (Baum 1978, 

Allred 2002). It is not significantly grazed or 

browsed. The tannin substances in the vegetation 

are likely to make it unpalatable. Tamarisk has 

commercial value in the landscape and 

horticultural trade. It is a popular ornamental shrub 

although its use in the United States is now 

generally discouraged. Many cultivars exist that 

have been chosen for flower color and growth 

form (Bailey and others 1976, Tykač 1990). 

Tamarix species, although not specifically T. 

chinensis, have been used for fuel and building 

materials by Native American tribes in the 

Western United States (Moerman 1998).

References. 

Allred, K. 2002. Identification and taxonomy of 

Tamarix (Tamaricaceae) in New Mexico. 

Desert plants 18(2): 26-31 

Bailey, L.H., E.Z. Bailey, and the staff of the 

Liberty Hyde Bailey Hortorium. 1976. Hortus 

Third. Macmillian Publishing Company, New 

York. 1,290 p. 

Baskin, C.C. and J.M. Baskin. 2001. Seeds, 

ecology, biogeography, and evolution of 

dormancy and germination. Academic Press, 

New York. 666 p. 

Baum, B.R. 1978. The genus Tamarix. The Israel 

Academy of sciences and humanities, 

Jerusalem, Israel. 209 p. 

Busch, D.E. and S.D. Smith. 1995. Mechanisms 

associated with decline of woody species in 

riparian ecosystems of the Southwestern U.S.A. 

Ecological Monographs. 65(3) 347-370. 

Cinnamon, S.K. 1990. Wupatki National 

Monument tamarisk and camelthorn eradication 

program 1983-1988. In: Tamarisk control in 

Southwestern United States. Special Report 9. 

Cooperative National Park Resources Studies 

Unit. p. 20-24. 

DeLoach, C.J. 1994. Petition to release into the 

field the leaf beetle Diorhaba elongata from 

China for biological control of saltcedar, 

Tamarix ramosissima, a weed of riparian areas 

of the Western United States and Northern 

Mexico. Submitted to the Technical Advisory 

Group for the Introduction of Biological 

Control Agents of Weeds, USDA-APHIS. 21 

March. 

DeLoach, C.J., D. Gerling, L. Fornasari, R. 

Sobhain, S. Myartseva, I.D. Mityaev, Q.G. Lu, 

J.L. Tracy, R. Wang, J.F. Wang, A. Kirk, J.W. 

Pemberton, V. Chikatunov, R.V. Jashenko, J.E. 

Johnson, H. Zheng, S. L. Jiang, M.T. Liu, P.A. 

Liu, and J. Cisneros. 1996. Biological control 

program against salt cedar (Tamarix spp.) in the 

U.S.A. In: V. C. Moran and J.H. Hoffman, eds. 

Proc. of the IX International symposium on the 

control of weeds, Cape Town, South Africa. p. 

253-260. 

Dick-Peddie, W.A. 1993. New Mexico vegetation 

– past, present, and future. University of New 

Mexico Press, Albuquerque, NM. 244 p. 

Epple, A.O. 1995. A field guide to the plants of 

Arizona. Falcon Press Publishing Co., Helena, 

MT. 347 p. 

Hays, F. and J. Mitchell. 1990. Tamarisk 

eradication in Zion national Park. In: Tamarisk 

control in Southwestern United States. Special 

Report 9. Cooperative National Park Resources 

Studies Unit. p. 36-38. 

Hughes, D. 1996. Restoring native riparian 

vegetation. In: Desired future conditions for 

South western riparian ecosystems: Bringing 

interests and concerns together. General 

Technical Report RM-GTR-272. U.S. 

Department of Agriculture, Forest Service, Fort 

Collins, CO. 

Kunzman, M.R., and P.S. Bennett. 1990. Arsenal 

as a control agent for Saltcedar (Tamarix). In:

Tamarisk control in Southwestern United 

States. Special Report 9. Cooperative National 

Park Resources Studies Unit. p. 82-90. 

McClintock, E. 1951. Studies in California 

ornamental plants. 3. The tamarisks. California 

Horticulture Society Journal 12: 76-83 

Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. 927 p. 

Neill, W.M. 1990. Control of Tamarisk by cutstump herbicide treatment. In: Tamarisk 

control in Southwestern United States. Special 
 

752 

Report 9. Cooperative National Park Resources 

Studies Unit. p. 91-98. 

Olson, BE. 1999. Impacts of noxious weeds on 

ecologic and economic systems. In: R.L Sheley 

and J.K. Petroff, eds. Biology and Management 

of noxious rangeland weed. Oregon State 

University Press, Corvallis, OR. 528 p. 

Sisneros, D. 1991. Herbicide analysis: Lower 

Colorado River saltcedar vegetation 

management study. Publication R-91-06. 

Bureau of Reclamation, Denver, CO. 165 p. 

Tykač, J. 1990. The illustrated guide to ornamental 

shrubs. Treasure Press, London. 224 p. 

United States Department of Agriculture, Forest 

Service. 1988. Range Plant Handbook. Dover 

Publications, Inc. New York. 838 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins. 1993. A Utah flora. Brigham Young 

University, Provo, UT. 986 p. 

Wilgus, F. and K.C. Hamilton. 1962. Germination 

of salt cedar seed. Weeds 10: 332-333. 
 

753

Tamonea boxiana (Moldenke) Howard crow broom 

VERBENACEAE 

Synonyms: Ghinia boxiana Moldenke 

 Tomonea spicata sensu Urban 

 Ghinia spinosa sensu Britt. & Wils. 

John K. Francis 

 

General Description.—Crow broom, also known 

as coast broom and cardero, is a branchy shrub 40 

cm to 100 cm in height. The stems are slender and 

woody with many bifurcating branches. Crow 

broom plants are supported by robust taproots 

with laterals through their lengths. The shortpetioled lower leaves are oblong, 0.8 to 1 cm long 

and may have three to five lobes; the upper leaves 

are linear and up to 2.5 cm long. Leaves and twigs 

are yellow-green. The plant is deciduous or 

semideciduous during the dry season. Tiny purple 

or whitish flowers are grouped in terminal 

racemes. The fruits are dry and shiny with four 

horns and contain a four-celled central cavity with 

1-mm oblong seeds (Howard 1989, Liogier 1995). 

Range.—Crow broom is native to Puerto Rico, 

the Virgin Islands, Barbuda, and Antigua (Howard 

1989, Liogier 1995). It is not known to have been 

planted or naturalized elsewhere. 

Ecology.—Crow broom grows in dry habitats that 

receive from about 700 to 900 mm of annual 

precipitation. It is normally found within a few km 

of the coast at elevations from near sea level to 

about 450 m. Soils are variable in texture, pH, and 

parent material, but are always well drained. Crow 

broom is intolerant of shade and heavy 

competition and apparently requires disturbance to 

become established. It is usually open-grown but 

also occurs in savannas. Crow broom is not eaten 

or is browsed little by cattle and appears to benefit 

from control of competition in overgrazed 

pastures. Vasquez and Kolterman (1998) list it as 

“occasional” in pasture and shrublands in 

Southeastern Puerto Rico. It is also found on 

roadsides, disturbed areas, and on rocky ridges 

and hill sides. 

Reproduction.—Crow broom blooms during the 

dry season (March and April). It is insect 

pollinated. The fruits mature during the wet season 

and are released at the end of the wet season 

(December through February). The seed-like fruits 

are dispersed by sticking to animals or clothing, 

and after being released, germinate (presumably 

during the wet season) and grow. A collection of 

air-dried fruits from Puerto Rico weighed an 

average of 0.0108 + 0.0003 g/seed or 93,000 

fruits/kg. 

Growth and Management.—Crow broom plants 

grow 20 to 30 cm/year and live about 4 or 5 years. 

No management experience has been reported. 

There appears to be no reason to encourage the 

species and possibly motivation in rangeland to 

control it. However, it is seldom common enough 

to warrant control measures. 
 

754 

Benefits and Detriments.—Crow broom 

contributes to biodiversity in the habitat where it 

grows, helps protect the soil, and furnishes cover 

for wildlife. It may become a minor weed in 

rangelands and the clinging seeds can be a 

nuisance. 

References 

Howard, R.A. 1989. Flora of the Lesser Antilles. 

Vol. 6. Arnold Arboretum, Harvard University, 

Jamaica Plain, MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de 

la Universidad de Puerto Rico, Río Piedras, PR. 

617 p. 

Vasquez, O.J. and D.A. Kolterman. 1998. Floristic 

composition and vegetation types of the Punta 

Guaniquilla Natural Reserve—Cabo Rojo, 

Puerto Rico. Caribbean Journal of Science 34(3-

4): 265-279. 
 

755

Taxus brevifolia Nutt. Pacific yew 

TAXACEAE 

Synonyms: Taxus baccata L. ssp. brevifolia (Nutt.) Pilger 

 Taxus baccata var. canadensis Benth. 

 Taxus boursieri Carrière 

 Taxus lindleyana A. Murray 

John K. Francis 

General Description.—Pacific yew, also known 

as western yew and mountain mahogany, is an 

evergreen large shrub or small tree. In good sites 

in coastal areas and a few inland areas, old 

specimens sometimes reach 60 cm in diameter at 

breast height and 15 m in height. In high elevation 

and poor sites, Pacific yew usually grows to only a 

few meters in height and a few centimeters in 

diameter, sometimes forming low mats. The bark 

of stems is thin (6 mm) and has a multihued 

purplish scale pattern. Bark on young twigs is rose 

colored and on older ones, reddish-brown. The 

sapwood is yellow and the heartwood red to 

reddish brown. Roots are deep and widespreading. The green leaves are linear, 8 to 35 mm 

long and 1 to 3 mm broad, acute at the tip, with a 

median ridge on the upper surface and two yellowgreen bands on the lower surface. They are 

whorled but bent to give a two-ranked appearance. 

Inconspicuous male and female strobili are borne 

on the underside of branches and occur on 

different trees (dioecious). The fruit (seed) is twoto four-angled, ovoid, 5 to 6.5 mm long, enclosed 

in a cup-like, fleshy red aril about 10 mm in 

diameter. Seeds have a large oily endosperm and a 

small embryo (Abrams 1940, Bolsinger and 

Jaramillo 1990, Earle 2002, Rudolf 1974, Viereck 

and Little 1972). 

Range.—Pacific yew is native to the coastal 

forests from southern Alaska through northern 

California and a moist interior corridor from 

southern British Columbia and Alberta to 

northwestern Montana, north-central Idaho, and a 

little of eastern Washington and Oregon (Bolsinger 

and Jaramillo 1990). Although the species has 

been planted to a limited extent in Europe, it is not 

known to have naturalized outside its native range. 

Ecology.—Pacific yew is very tolerant according 

to the five-step tolerance rating. Seedlings grow, 

develop, and eventually fruit under heavy conifer 

forest canopies. The species is most often found in 

deep ravines, along streams, and under coniferous 

stands (Sargent 1923). It grows as an understory 

dominant in most of the forest types where it 

occurs. Under closed stands, the limbs are often as 

long as the trees or shrubs are tall (Bolsinger and 

Jaramillo 1990, Termenstein 1990). Leaves persist 

on the twigs 4 to 5 years (Abrams 1940). Although 

initially shocked, it is able to adapt to full sunlight 

after overstory removal by logging, and it 

sometimes grows in the open in avalanche chutes, 

and on ridges and rocky slopes at high elevations 

(Bolsinger and Jaramillo 1990, Termenstein 1990). 

Plants growing in the interior of the continent are 

more often shrubby than those near the coast 

(Sargent 1923). It typically grows as an understory 

tree 3 to 5 m tall near the coast and as a shrub 

about 1 m tall east of the Cascade Mountains. It 

grows from near sea level to about 2,200 m in 

elevation (Earle 2002). 

Reproduction.—Pacific yew flowers mostly in 

June, matures fruits August to October, and 
 

756 

releases seeds in October. The pollen is dispersed 

by the wind. There are 32,000 to 36,000 seeds/kg 

(Washington source). About 50 to 99 percent will 

eventually germinate under proper conditions. The 

seeds are dispersed primarily by birds. Natural 

germination usually takes place in the second 

spring after dispersal (Rudolf 1974). Development 

of seedlings is slow. Pacific yew sprouts from 

stumps and rootstalks after being cut or top-killed. 

It commonly layers when branches or stems are 

pressed to the ground by snow or falling debris 

(Bolsinger and Jaramillo 1990). 

Growth and Management.—Growth of Pacific 

yew is slow. Trees in Idaho took 25 years to reach 

a stem diameter (15 cm above the ground) of 2.5 

cm and 100 years to reach 15 cm. Height growth is 

correspondingly slow (Bolsinger and Jaramillo 

1990). Even in moist, rich soils, few plants live 

long enough to reach tree sizes. Fruits should be 

picked by hand as soon as they are ripe. The fruit 

pulp is removed by maceration and floating off the 

pulp, or by soaking in water at room temperature 

for 4 or 5 days, rubbing the fruits on screens, and 

washing away the pulp. The seeds should be airdried for 1 to 2 weeks and should be sown or 

stored as soon as drying is complete. Air-dried 

seeds can be stored for 5 to 6 years in sealed 

containers at 1 to 2 °C. Seeds sown in nursery beds 

should be covered by about 1 cm of mineral soil 

and mulched. Germination primarily occurs in the 

second spring afterwards. To obtain germination a 

shorter time after sowing, seeds should be 

stratified for 90 to 210 days at 16 °C followed by 

60 to 120 days at 3 to 6 °C (Rudolf 1974). 

Benefits.—Pacific yew contributes to the 

aesthetics of the forest, helps protect the soil, and 

provides food and cover for wildlife. It is a 

preferred moose browse. Although eaten in all 

seasons, during the winter, moose will eat all the 

available leaves and twigs, and even strip the bark. 

Deer, elk, and rabbits also browse the species. 

Livestock make limited use of it during the winter 

and when other food is lacking. The fruit is sweet 

and eaten by many species of birds (Tirmenstein 

1990). Plants growing along streams help protect 

against streambank erosion and shade the water 

and help maintain cool stream temperatures 

(Termenstein 1990). Pacific yew wood is finegrained, heavy, hard, elastic, and strong. It has 

been used and still is used to a limited extent (due 

to scarcity) for archery bows, harpoon shafts, 

canoe paddles, household utensils, tool handles, 

mauls, splitting wedges, gunstocks, boat decking, 

fence posts, musical instruments, carved figurines, 

novelty items, furniture, oriental ceremonial 

“Toko” poles, turnery, cabinetry, and firewood 

(Bolsinger and Jaramillo 1990, Termenstein 1990). 

Infusions, decoctions, and poultices of leaves, 

twigs, and bark were used by Native Americans to 

treat lung problems, stomachache, wounds, and 

pain (Moerman 1986). Pactitaxel (often called by 

the trade name Taxol) was discovered during a 

massive anticancer activity screening program by 

the National Cancer Institute during the 1960’s 

(21cecPharm 2003). Paclitaxel is a white 

crystalline powder extractable from any of the 

Taxus species. It is a cytotoxic anticancer drug 

used to treat ovarian, breast, and lung cancers, and 

Kaposi’s sarcoma (SFT Enterprise 2003). 

Harvesting for peclitaxel extraction threatened to 

decimate the species’ populations until it was 

discovered that other Taxus species were better 

sources and that the drug could be produced by 

semi-synthesis and by cell cultures (21cecPharm 

2003). Pacific yew is used to a limited extent as an 

ornamental shade tree, foundation plant, hedge, 

and topiary plant (Bolsinger and Jaramillo 1990). 

References 

21cecPharm. 2003. Paclitaxel story. http://www. 

21cecpharm.com/px/story.htm. 4 p. 

Abrams, L. 1940. Illustrated flora of the Pacific 

States. Stanford University Press, Stanford, CA. 

538 p. 

Bolsinger, C.L. and A.E. Jaramillo. 1990. Taxus 

brevifolia Nutt., Pacific yew. In: R.M. Burns, 

and B.H. Honkala, eds. Silvics of North 

America. Agriculture Handbook 654. Forest 

Service, U.S. Department of Agriculture, 

Washington, DC. p. 573-579. 

Earle, C.J. 2002. Taxus brevifolia Nuttall 1849. 

http://www.botanik.uni-bonn.de/conifers/ta/ta/ 

brevifolia.htm. 3 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports 19. University of 

Michigan Museum of Anthropology, Ann 

Arbor, MI. 534 p. 

Rudolf, P.O. 1974. Taxus L., yew. In: C.S. 

Schopmeyer, tech. coord. Seeds of woody plants 

in the United States. Agriculture Handbook 450. 
 

757

U.S. Department of Agriculture, Forest Service, 

Washington, DC. p 799-802. 

Sargent, C.S. 1923. Manual of the trees of North 

America (exclusive of Mexico). Houghton 

Mifflin, Boston, MA. 910 p. 

SFT Enterprise. 2003. Natural pactitaxel, full 

information. http://www.21cep.com/sft/ 

pxsft.htm. 9 p. 

Tirmenstein, D.A. 1990. Taxus brevifolia. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Missoula, MT. Fire Effects 

Information System. http://www.fs.fed.us/ 

database/feis/plants/tree/ taxbre/all.html. 19 p. 

Viereck, L.A. and E.L. Little, Jr. 1972. Alaska 

trees and shrubs. Agriculture Handbook 410. 

U.S. Department of Agriculture, Forest Service, 

Washington, DC. 265 p. 
 

758 

Taxus canadensis Marsh. Canada yew 

TAXACEAE 

Synonyms: Taxus minor (Michaux) Britt. 

 Taxus procumbens Loddiges 

 Taxus baccata L. var. canadensis Gray 

 Taxus baccata L. var. minor Michx. 

 Taxus baccata L. var. procumbens Loud 

Gerry Moore and John K. Francis 

Drawing source: Britton and Brown 1913 

General Description.—The common name most 

often used for Taxus canadensis is Canada yew. It 

is also known as American yew and ground 

hemlock; a common French name used in Quebec 

for this species is buis de sapin. This evergreen 

species is usually a low-growing, straggling nonaromatic shrub, with ascending branches up to 2 m 

tall. Branches often extend two-thirds of their 

length laterally before curving upwards. The bark 

is thin, scaly and red to reddish-brown. Twigs are 

green and alternately arranged, and winter buds are 

composed of imbricate (overlapping), lanceolate, 

keeled scales. The leaves are needle-like, on short 

stalks, and persist for several years. The leaf 

blades are spirally arranged, flattened, often in two 

ranks, linear, abruptly narrowed into a fine point, 

1.2 to 2.2 cm long, with pale green bands on the 

under surface. The midrib is slightly elevated on 

the top surface. The leaves lack resin ducts. In 

winter, the leaves can be slightly reddish. Being a 

gymnosperm, Canada yew lacks true flowers or 

fruits. It does produce pollen and seeds, on 

separate plants (dioecious) or on the same plant 

(monoecious) in axillary cones. The pollen is 

produced by the male cones, which appear as 

stalked heads and mature in the first season. Each 

male cone is a compound of four to 16 reduced, 

stalked, specialized leaves called sporophylls. The 

stalk is attached to the center of the sporophyll (i.e. 

they are peltate). Each sprophyll produces two to 

nine pollen sacs (sporangia) where the pollen is 

produced. The pollen is spherical and lacks wings. 

The female cones are reduced, subtended by a 

series of small, inconspicuous bracts. Each cone 

produces one ovule that develops into a single 

seed, maturing in one season. The seeds are 

brown, flattened, slightly broader than long, 4 to 5 

mm long. Each seed is surrounded by a fleshy, 

scarlet, cup-shaped aril that is open at the top, 

exposing the seed. Inside the aril is a clear, 

mucilaginous liquid called albumen (Fernald 1950, 

Pilger 1903, Soper and Heimburger 1982). 

Range.—Canada yew is native to Eastern Canada 

(Manitoba, New Brunswick, Newfoundland, Nova 

Scotia, Ontario, Prince Edward Island, Quebec) 

and the Northeastern United States (Connecticut, 

Illinois, Indiana, Iowa, Kentucky, Maine, 

Maryland, Massachusetts, Michigan, New 

Hampshire, New Jersey, New York, North 

Carolina, Ohio, Pennsylvania, Rhode Island, 

Tennessee, Vermont, Virginia, West Virginia, 

Wisconsin) (Hils 1993, Natural Resources 

Conservation Service 2003). It becomes rare above 

50 °N (Soper and Heimburger 1982). 

Systematic Botany.—The genus name Taxus is 

from the Greek word, taxos, for yew. 

Approximately eight species of Taxus are 

recognized, and all of them are similar 

morphologically, even though they are wellseparated geographically (Hils 1993). Canada yew 
 

759

is the only native yew throughout its range. 

However, it can be confused with two other yew 

species--English yew (T. baccata L.) and Japanese 

yew (T. cuspidata Sieb. & Zucc.)--and a hybrid 

between them, (T. media Rehd.) that occasionally 

escapes from cultivation. Canada yew can be 

distinguished from these by its low-growing, 

straggling nature. Immature saplings are 

unidentifiable to species. Canada yew has 2n = 24 

chromosomes (Hils 1993). 

Ecology.—Canada yew, a slow-growing species, 

is tolerant of shade, which gives it a competitive 

advantage over other species with which it must 

compete. It grows best in at least partial shade 

(Sullivan 1993). Canada yew is found in rich soil 

in a variety of habitats including bogs, swamps, 

and rocky banks at elevations up to 1,500 m. It 

prefers soils that are well-drained and that have 

pH’s between 5.0 and 7.5. The climate is humid 

and cool. Exposure to winds from the ocean is not 

tolerated. The species grows in many forest 

associations and is taken as an indicator of oldgrowth forest. It does not occur in seral 

communities. Canada yew is apparently easily 

killed by fire. Wildfires and other influences that 

open the forest canopy reduce the competitive 

ability of the species (Sullivan 1993). 

Reproduction.—Canada yew flowers in April and 

May and is wind pollinated. Individual plants are 

not self-fertile (Plants For a Future 2003). It 

produces seed almost every year (Sullivan 1993). 

The seeds mature in late summer and autumn. 

There are 33,000 to 62,400 seeds/kg (Rudolf 

1974). The seeds are dispersed by birds (Sullivan 

1993). Canada yew spreads vegetatively by 

layering (Rook 1998). The connections between 

plants of a clonal group usually rot away (Sullivan 

1993). Small plants tend to be males and browsing 

and other stresses on a population tends to increase 

the proportion of male plants in the population 

(Sullivan 1993). 

Growth and Management.—Canada yew is slow 

growing (Public Forest Council 2003). It is the 

hardiest of all yew species; however, it is the least 

attractive yew species used as an ornamental due 

to its low-growing, straggling habit (Rehder, 

1940). The species may be propagated with seeds 

and by cuttings of half-hardened or hardened 

terminal shoots that root with good success (Plants 

For a Future 2003). Warm stratification for 4 

months followed by 4 months of cold stratification 

are suggested to promote more rapid germination 

of seeds (Dirr and others 1987). Seeds may be 

sown in nursery beds in the fall and covered by 10 

to 12 mm of mineral soil topped by mulch. 

However, fall-sown seeds usually germinate the 

second spring (Rudolf 1974). Seedlings should be 

transplanted while still small. However, they are 

slow growing and they must be maintained for 2 

years in nursery beds or pots at a minimum. 

Seedlings should be outplanted in late spring or 

early summer after the last frost has occurred. 

Established plants will tolerate pruning (Brand 

2003). 

Benefits.—Canada yew is an important 

component of the forest understory in Northeastern 

forests, helping to protect the soil and imparting 

beauty to forest landscapes. The species is used as 

an ornamental for ground cover and mass 

plantings, and for parental stock in yew breeding 

(Sullivan 1993). The toxic compound that is 

present in the plant is the alkaloid taxine 

(Kingsbury 1964). The seeds and the dried foliage 

have been fatal to livestock, but the fresh foliage is 

browsed by deer (Fernald 1950, Hils 1993). Moose 

also browse the species and have seriously reduced 

Canada yew abundance on Isle Royale, Michigan, 

since their introduction. The fleshy aril is eaten by 

numerous species of birds (Sullivan 1993). The 

pulp of the arils is sweet and edible to humans 

(Fernald 1950) but has a slimy texture (Soper and 

Heimburger 1982). A number of Native American 

groups made decoctions of leaves and twigs to 

treat rheumatism. Infusions and decoctions were 

also used to treat numbness of fingers and legs, 

colds, gonorrhea, and as a diuretic (Moerman 

1986). New foliage and green stems are 

commercially harvested for the extraction of 

paclitaxel (Public Forest Council 2003). Paclitaxel 

is a white crystalline powder extractable from any 

of the Taxus species. It is a cytotoxic anticancer 

drug used to treat ovarian, breast, and lung 

cancers, and Kaposi’s sarcoma (SFT Enterprise 

2003). 

References 

Brand, M.H. 2003. Plant UConn database of trees, 

shrubs and vines: Taxus canadensis. 

http://www.hort.uconn.edu/plants/t/taxcan/ 

taxcan3/html. 2 p. 

Britton, N.L. and A. Brown. 1913. Illustrated flora 

of the northern states, Canada, and the British 
 

760 

possessions. Vol. 2, 2nd Ed. Scribner, New York. 

735 p. 

Dirr, M.A., M.W. Heuser, Jr., and B.L. Dirr. 1987. 

The reference manual of woody plant 

propagation. Varsity Press, Athens, GA. 239 p. 

Fernald, M.L. 1950. Gray’s manual of botany. 

American Book Co., New York. 1,632 p. 

Hils, H.J. 1993. Taxaceae. In: Flora of North 

America Editorial Committee, eds. Flora of 

North America. Vol. 2. Oxford University Press, 

New York. p. 423-427. 

Kingsbury, J.M. 1964. Poisonous Plants of the 

United States and Canada. Prentice-Hall, Inc., 

Englewood Cliffs, NJ. 626 p. 

Moerman, D.E. 1986. Medicinal plants of Native 

America. Technical Reports 19. University of 

Michigan Museum of Anthropology, Ann 

Arbor, MI. 534 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Taxus canadensis Marsh. http:// 

plants.usda.gov/cgi_bin/plant_search.cgi?mode=

Scientific+Name&keywordquery=Ta... 

 [not paged]. 

Pilger, R.K.F. 1903. Taxaceae. In: H.G.A. Engler, 

ed. Das Pflanzenreich. V. 18(IV,5). 

p. 1-124. 

Plants For a Future. 2003. Taxus canadensis. 

http://www.ibiblio.org/pfaf/chi-bin/arr_html? 

Taxus+canadensis. 7 p. 

Public Forest Council. 2003. Ground hemlock 

(Taxus canadensis). Public Forest Council Fact 

Sheet. Charlottetown, Prince Edward Island, 

Canada http://www.gov.pe.ca/af/agweb/ 

library/factsheets/ground_hemlock.pdf. 2 p. 

Rehder, A. 1940. Manual of cultivated trees and 

shrubs. Macmillan Publishing Co., New York. 

996 p. 

Rook, J.S. 1998. Taxus canadensis, Canada yew. 

http://www.rook.org/earl/bwca/nature/shrubs/ 

taxuscan.html. 3 p. 

Rudolf, P.O. 1974. Taxus L., yew. In: C.S. 

Schopmeyer, tech. coord. Seeds of woody plants 

in the United States. Agriculture Handbook 450. 

U.S. Department of Agriculture, Forest Service, 

Washington, DC. p. 799-802. 

SFT Enterprise. 2003. Natural pactitaxel, full 

information. http://www.21cep.com/sft/ 

pxsft.htm. 9 p. 

Soper, J.H. and M.L. Heimburger. 1982. Shrubs of 

Ontario. Royal Ontario Museum, Toronto, 

Ontario, Canada. 495 p. 

Sullivan J. 1993. Taxus canadensis. In: U.S. 

Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/shrub/ 

taxcan/all.html. 10 p. 
 

761

Tecoma stans (L.) Juss. ex Kunth ginger-thomas 

BIGNONIACEAE 

Synonyms: Bignonia stans L. 

Gelseminum stans (L.) Kuntze 

Stenolobium stans (L.) Seem. 

John K. Francis 

 Illustration source: USDA-Forest Service collection, 

Hunt Institute 

General Description.—Ginger-thomas, also 

known as roble amarillo, saíco amarillo, bois 

caraîbe, trumpet flower, yellow-elder, and many 

other common names (Little and Wadsworth 

1964), is a medium-sized shrub with many 

branches and basal stems. The leaves are opposite, 

pinnately compound with 5 to 13 saw-toothed 

leaflets. The twigs are green, turning brown, and 

the older bark is light gray and very furrowed. A 

profusion of bright yellow flowers has made the 

species much loved throughout the tropics. Gingerthomas is the official flower of the U.S. Virgin 

Islands. 

 

Range.—The native range extends from southern 

Texas, New Mexico, and Arizona to Bolivia and 

northern Argentina and from Florida and the 

Bahamas to Trinidad in the Caribbean. It has 

naturalized in much of tropical and subtropical 

Africa, Asia, the Pacific islands, and Australia. 

Ecology.—Ginger-thomas does not tolerate heavy 

frost. However, it will grow in most well-drained 

soils, including calcareous fill, infertile sands, 

acidic Ultisols, and volcanic regolith in areas 

receiving from 700 to 1800 mm of rainfall. The 

species is described as a water spender that is able 

to convert to a water saver (Tipton 1994). It 

requires minimal competition and nearly full 

sunlight to survive (shade intolerant). Roadsides 

and disturbed areas are the most common habitats. 

Given a start in secondary forest, it is usually 

present for only 10 to 20 years following 

disturbance. Many species of insects feed on 

ginger-thomas, and it is attacked by the genera of 

parasitic plants, Cuscuta, and a number of disease 

organisms; but, nothing seems to pose a serious 

threat to the species. 

Reproduction.—Ginger-thomas produces flowers 

that have functional male and female parts. The 

tubular flowers are fragrant, 4 to 5 cm in length, 

and pollinated by bees and other insects and 

hummingbirds. The 10- to 25-cm pods (capsules) 

develop in about 1 month and liberate large 

numbers of papery-winged seeds. The species may 

flower and fruit nearly throughout the year (Little 

and Wadsworth 1964) in climates without strong 

seasonal change, or flower heavily in autumn in 

seasonal climates such as southern Florida. 

Developing pollen becomes sterile when 

temperatures rise above 34 °C. This leads to seed 

failures during summer months in many areas 

(Kumar and Singh 1988). Natural reproduction 

may vary from dense to scattered. In nurseries, 

ginger-thomas is usually propagated from seed, 

although greenwood cuttings can also be rooted 

(Baily 1941). A collection of seeds from Puerto 

Rico averaged 208,000 seeds/kg. Germinating 

began in 3 days and finished with 97 percent 
 

762 

germinated (Francis and Rodríguez 1993). No 

pregermination treatments are necessary. 

Growth and Management.—Early growth is 

relatively rapid. About 1 m of height growth can be 

obtained during the first year. Depending on the 

environment, maximum height of plants may vary 

from 1.5 m to 7 m, and diameters of 8 cm are 

sometimes achieved. Establishment activities may 

include removal of competition and disturbance of 

the soil to encourage natural regeneration, planting 

(more reliable for smaller numbers of plants), and 

control of competing vegetation. When control of 

ginger-thomas is needed, cutting, grubbing out 

stumps, or poisoning is recommended. In Brazil, 

tebuthiuron was found to effectively kill plants, but 

picloram and 2, 4-D were not (Passini and Kranz 

1997). 

Benefits.—Ginger-thomas is planted as an 

ornamental throughout the topics and subtropics. It 

is especially prized as a flowering hedge plant. The 

shrub is planted and managed to enhance the 

beauty of green belts and natural forests used for 

recreation. Used in combination with trees, gingerthomas contributes to effective windbreaks and 

sound breaks. Growing in thick patches that shade 

out grass, the species has become a serious weed in 

Brazilian pastures (Kranz and Passini 1997). 

Browsing cattle and goats in Mexico consume up 

to 20 percent of the leaves and 100 percent of the 

available flowers (Susano Hernandez 1981). The 

foliage in one study in India contained 17 percent 

crude protein, 6 percent ash, 18 percent fat, 25 

percent fiber, and 14 percent total polyphenols 

(Nag and others 1994). Ginger-thomas leaves, 

bark, and roots contain many biologically active 

chemicals, and extracts from those tissues have 

been used in traditional folk medicine to treat many 

diseases and conditions (Liogier 1990). Perhaps the 

most promising compounds are monoterpine 

alkaloids, which have been shown to effectively 

reduce the symptoms of diabetes mellitus in rats, 

dogs, and mice (Aguilar and others 1993, LozoyaMeckes and Mellado-Campos 1985, Perez and 

others 1984) 

References 

Aguilar, L.C., S. Macias, A. Chagoya, A. 

Cardenas, P. Díaz, and J M. Cantu. 1993. 

Antidiabetic activity of Tecoma stans in rats. 

Fitoterapia 64(4): 304-305. 

Bailey, L.H. 1941. The standard cyclopedia of 

horticulture. Vol. 3. The MacMillan Company, 

New York. p. 2,423-3,639. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

Kranz, W.M. and T. Passini. 1997. Amarelinho:

biologia e control. Informe da Pesquisa 

No.121.Instituto Agronomico do Parana. 

Londrina, PR, Brazil. 19 p. 

Kumar, R. and G. Singh. 1988. Investigations into 

the cause of sterility. Tecoma stans L. Letters 

Botaniques (France) 135(2): 131-135. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the 

VirginIslands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service. 

Washington, DC. 548 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Lozoya-Meckes, M. and V. Mellado-Campos. 

1985. Is the Tecoma stans infusion an antidiabetic remedy? Journal of Ethnopharmacology 

14(1): 1-9. 

Neg Datta, A., A.K. Lehiri, S. Matai, and L. Si. 

1994. Proximate composition and polyphenol 

content of some tree leaves. Indian Forester 

120(12): 1,122-1,125. 

Passini, T. and W.M. Kranz. 1997. Eficacia de 

herbicidas no controle de amarelinho (Tecoma

stans) em pastagem. Planta Daninha 15(2): 190-

197. 

Perez G., R.M., A. Ocegueda Z., J.L. Muñoz L., 

J.G. Avila A., and W.W. Morrow. 1984. A study 

of the hypoglycemic effect of some Mexican 

plants. Journal of Ethnopharmacology 12(3): 

253-262. 

Susano Hernandez, R. 1981. Especies arboreas 

forestales susceptibles de aprovecharse como 

forraje. Ciencia Forestal 6(29): 31-39. 
 

763

Tipton, J.L. 1994. Relative drought resistance 

among selected southwestern landscape plants. 

Journal of Agriculture 20(3): 150-155. 
 

764 

Tetrapterys inaequalis Cav. bejuco de paralejo 

MALPIGHIACEAE 

Synonyms: Tetrapteris citrifolia (Sw.) Pers. 

 Banisteria bracteata DC. 

 Triopteris citrifolia Sw. 

 Tetrapteris paniculata Bello 

John K. Francis 

General Description.—Bejuco de paralejo, also 

known as brjuco de sopla and aile à ravet, is a 

woody vine or climbing shrub that may extend 15 

m laterally and reach 10 m into the crowns of 

trees, with stems as thick as 8 cm. The root system 

is superficial, especially roots arising from layered 

stems. The stems are cylindrical, smooth, and have 

rings at the nodes. The wood is moderately hard 

and has growth rings. The stems tend to be long 

and little branched except near the extremities. 

The simple, opposite ovate or elliptical leaves 

have petioles 1 to 1.5 cm long, and blades 5 to 17 

cm long and 5 to 8 cm broad, rounded to cordate 

at the base and pointed at the tip. Inflorescences 

are panicles of four-flowered cymes flanked by 

leaves of reduced size that often shrivel as the 

fruits mature. The five-merous, yellow-petaled 

flowers are about 2 cm in diameter. The fruits 

consist of four samaras united at the base. Each 

has an upper lateral wing 17 to 28 mm by 6 to 10 

mm and a lower lateral wing about half as large 

and a medial ridge protruding about 1 mm 

(Acevedo-Rodríguez 1985, Howard 1988, Liogier 

1988). 

Range.—The range of bejuco de paralejo includes 

Puerto Rico and the offshore island of Vieques, St. 

Thomas, and St. Croix in the U.S. Virgin Islands, 

Antigua, Guadaloupe, Martinique, and St. Vincent 

(Howard 1988). This differs from Liogier (1988) 

and Acevedo-Rodríguez (1985) who use the 

synonym T. citrifolia and add Jamaica, Hispaniola, 

and Tobago to the range. Howard states: “The 

flowering specimen from Tobago that Niedenzu 

called T. citrifolia is almost certainly T. discolor

(G. Meyer) DC. ...Niedenzu considered [T. 

inaequalis] a synonym of Tetrapterys citrifolia

(Sw.) Pers. They are certainly closely related, both 

belonging to the difficult group called Section 

Lophogynixa by Niedenzu. However, T. citrifolia

from Jamaica, the type locality, has a welldeveloped dorsal wing on the samara and young 

stems that are tightly sericeous with straight 

appressed hairs. These differences lead me to 

consider T. citrifolia endemic to Jamaica and apply 

the later name T. inaequalis to the plant of Puerto 

Rico and the Lesser Antilles. ...The species of the 

Lesser Antilles seems not to occur on 

Hispaniola...” In their latest checklist, Liogier and 

Martorell (2000) now concur with Howard. 

Ecology.—Bejuco de paralejo colonizes a wide 

range of soils derived from both sedimentary and 

igneous rocks. It requires mean annual rainfall of 

from about 1200 to about 2500 mm. The species 

grows at elevations between 110 and 500 m in the 

Lesser Antilles (Howard 1988) and from a few 

meters above sea level to 600 m or more in Puerto 

Rico. Bejuco de paralejo does not tolerate shade 

well. It will survive and grow slowly in light shade 

but requires full or nearly full sunlight to flower 

and fruit. The species may be found on roadsides, 

along rivers, in brushy pastures, early secondary 

forests, and clearings in secondary and remnant 

forests. 

Reproduction.—Bejuco de paralejo usually 

flowers in Puerto Rico in October and November 

and fruits in July to October (Acevedo-Rodríguez 

1985). Howard (1988) notes collections in the 

Lesser Antilles with flowers and fruits in every 

month except May and June. Hundreds of seeds 

can be produced by a large plant. Air-dried 
 

765

samaras from Puerto Rico collected by the author 

in April averaged 0.0953 + 0.0014 g/fruit. Planted 

on commercial potting mix without any 

pretreatment, 79 percent germinated between 34 

and 82 days after sowing. Samaras spiral sideways 

a few meters from fruiting adult plants. While 

young plants are relatively common in many 

areas; fruiting plants are relatively rare. Layering 

(rooting in contact with the ground) is common 

and appears to be an important means of 

perpetuating established plants. 

Growth and Management.—Bejuco de paralejo 

vines extend rapidly from sprouts. A large vine 

(7.7 cm in diameter) cut by the author had 27 

growth rings. Although large plants are capable of 

smothering small trees, the species causes few 

problems. No planting or management experience 

has been published. 

Benefits.—Bejuco de paralejo contributes to the 

biodiversity of forests, helps protect the soil, and 

furnishes cover for wildlife. 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Liogier, H.A, and L.F. Martorell. 2000. Flora of 

Puerto Rico and adjacent islands, a systematic 

synopsis. 2nd ed. Editorial de la Universidad de 

Puerto Rico, Río Piedras, PR. 382 p. 
 

766 

Thrinax morrisii H. Wendl. brittle thatch palm 

ARECACEAE 

Synonyms: Sampsonia microcarpa (Sarg.) O.F. Cook 

 Thrinax keyensis Sarg. 

 Thrinax microcarpa Sarg. 

 Thrinax ponceana O.F. Cook 

 Thrinax praeceps O.F. Cook 

John K. Francis 

General Description.—Brittle thatch palm is also 

known as Key thatch palm, small-fruited thatch 

palm, broom palm, buffalo-tip, palma de escoba, 

yaray, pandereta, palma de petate, palma de 

cogollo, guano de sierra, miraguano, and palmita. 

It is an evergreen, single-stemmed shrub or small 

tree 1 to 6 m in height. Although most plants in 

natural stands do not live long enough to develop a 

discernable stem, when formed, it is brown or 

gray, 7 to 13 cm in diameter at breast height, with 

many vertical cracks and prominent leaf scars in 

the clean portion, covered with old leaf bases near 

the top, and thick with rootlets at the base. Brittle 

thatch palms maintain about 20 fan-shaped, pale 

blue-green or yellow-green leaves, spirally 

arranged on the trunk. The petioles are 27 to 84 cm 

long, split at the base. There are deeply incised leaf 

segments 33 to 75 cm long and 2.3 to 4.8 cm 

broad. The inflorescences (spadixes or panicles) 

are 55 to 100 cm long, arching or straight and 

usually extend beyond the leaves. Small, white 

flowers are borne in large numbers. The fruits are 

white (turning yellow at maturity) drupes, 3.5 to 8 

mm in diameter, containing one hard, spherical 

seed 2.7 to 4.2 mm in diameter. Chromosome 

number is 2n = 36 (Gilman and Watson 1994, 

Howard 1979, Little and others 1974, Long and 

Lakela 1976, Nelson 1996) 

Range.—Brittle thatch palm is native to southern 

Florida, the Bahamas, Cuba, Hispaniola, Puerto 

Rico, the Virgin Islands, Anguilla, and Barbuda 

(Howard 1979, Little and others 1974). It is 

considered endangered in Florida (Institute of 

Systematic Botany 2002). The species is planted as 

an ornamental within and outside its natural range 

(Hoyos-F. and Braun 2001, Little and others 1974) 

but has not been reported to have naturalized. 

Ecology.—Brittle thatch palm grows in sites 

where it suffers a minimum of competition. It 

grows along the edges of hammocks and in 

pinelands in the Florida Keys (Nelson 1996). In 

Puerto Rico, it grows primarily on cliffs, and 

ridges of dry and moist limestone and ultramafic 

rocks at elevations from near sea level to 300 m 

(Little and others 1974). The soils are sands, or are 

loams or clays shallow over fractured rocks. Brittle 

thatch palm is moderately intolerant of shade. It 

occurs singly, in small groups or in nearly pure 

thickets in the open, in small openings, or under 

open stands, usually of low trees. The species can 

withstand temperatures as low as -4 °C (Deserttropicals 2002), is tolerant of drought and salt 

spray, and resists breakage in storms (Gilman and 

Watson 1994). 

Reproduction.—Brittle thatch palm in natural 

stands flowers seasonally or irregularly in response 
 

767

to favorable conditions in Puerto Rico (Little and 

others 1974) and flowers in the spring in Florida 

(Gilman and Watson 1994). The flowers are insect 

pollinated. A collection of seeds in Puerto Rico 

averaged 0.0396 + 0.0007 g/seed or 25,000 

seeds/kg. Sown without pretreatment on moist 

blotter paper, 85 percent germinated in 17 months 

(author’s observation). The seeds are reported to 

begin germination 50 to 70 days after sowing 

(Hoyos-F. and Braun 2001). The seeds can stand 

storage at -40 °C and 66 °C for at least 1 week 

without loss of viability (Broschat and Meerew 

2000). Because seeds have such a long dormant 

period, they can be collected by hand from plants 

or from the ground at almost any time of the year. 

The seeds are dispersed by animals. Surviving 

seedlings are not usually common, but the plants 

are long-lived and populations can build to high 

levels. 

Growth and Management.—Brittle thatch palm 

is slow growing. A specimen in the Fairchild 

Tropical Garden, Florida was 42 years old and 

1.16 m in height (Zona and Maidman 2000). Plants 

mature and begin flowering and fruiting with only 

a rosette of basal leaves, long before they develop 

a discernable stem. Plants with long stems are 

uncommon and are probably more than 50 years 

old. Wildland plantings are not documented but 

presumably could be done with container stock. 

Either they must be established in poor sites where 

few other species will grow, or weed protection 

must be given almost in perpetuity. Management 

of existing stands should consist mainly of 

protection from fire, development, or heavy 

grazing. 

Benefits.—Brittle thatch palm helps protect the 

soil, contributes to the aesthetics of natural stands, 

and furnishes food and cover for wildlife. The 

species is grown and sold as an ornamental. It is 

used as a container plant and for buffer strips in 

parking lots and medians on highways as well as 

background and specimen plantings (Gilman and 

Watson 1994). Leaves are still used to make 

brooms, thatch, and mats (called petate) (Osvaldo 

2002), although now for ornamentation rather than 

necessity. The fruit pulp contains 4,083 μg/g of 

oxalate—capable of causing a burning sensation 

on the skin of sensitive people—that may reduce 

its palatability to some species of wildlife 

(Broschat and Meerew 2000). However, it is eaten 

in Florida by the introduced Mexican red-bellied 

squirrel (Sciurus aureogester (Cowley 2002) and 

the eastern box turtle (Terrapene carolina) (Liu 

and others 2002), and in the Bahamas by the 

Bahamas amazon parrot (Amazona leucocephala 

bahamensis) (Attril 2002). 

References

Attril, R. 2002. Bahamas wildlife pages: The status 

and conservation of the Bahamas amazon. 

http://www.bahamaswildlife.fsnet.co.uk/ 

parrotconservation.htm. 8 p. 

Broschat, T.K. and A.W. Meerew. 2000. 

Ornamental palm horticulture. University Press 

of Florida, Gainesville, FL. 255 p. 

Cowley, M. 2002. Florida native plants: palm 

family. http://www.nsis.org/garden/family/palm. 

html. 3 p. 

Desert-tropicals. 2002. Broom palm. http://deserttropicals.com/Palm/Thrinax_morrisii.html. 3 p. 

Gilman, E.F. and D.G. Watson. 1994. Thrinax 

morrisii, Key thatch palm. Fact Sheet ST-628. 

U.S. Department of Agriculture and Southern 

Group of State Foresters. Winder, GA. 3 p. 

Howard, R.A. 1979. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 3. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 586 p. 

Hoyos-F., J. and A. Braun. 2001. Palmas en 

Venezuela. Monograph 47. Sociedad de 

Ciencias Naturales La Salle, Caracas, 

Venezuela. 424 p. 

Institute of Systematic Botany. 2002. Thrinax 

morrisii. Institute of Systematic Botany, 

University of South Florida, Tampa, FL. 

http://www.plantatlas.usf.edu/main.asp?plantID

=555. 2 p. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1024 p. 

Liu, H., S.G. Platt, and C.K. Borg. 2002. Seed 

dispersal by the eastern box turtle (Terrapene 

carolina) in subtropical pine rockland of the 
 

768 

Lower Florida Keys. Botany 2002 Conference, 

August 2-4, Madison, WS. Abstract. 

http://www.botany2002.org/section3/abstracts/3

6.shtml. 1 p. 

 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyan Books, Miami, FL. 

962 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc. Sarasota, FL. 

391 p. 

Osvaldo, J. 2002. El petate. http://joseosvaldo. 

freeyellow.com/Petate/html. 12 p. 

Zona, S. and K. Maidman. 2000. Growth rates of 

palms in Fairchild Tropical Garden. Palms 

45(3): 151-154. 
 

769

Toxicodendron radicans (L.) Kuntze eastern poison ivy 

ANACARDIACEAE 

Synonyms: Rhus radicans L. 

 Rhus littoralis Mearns 

 Rhus blodgettii Kearney 

 Philostemon radicans Raf. 

 Toxicodendron blodgettii (Kearney) Greene 

 Rhus scandens Salisb. (and others: Institute of Systematic Botany 2003) 

John K. Francis 

General Description.—Eastern poison ivy, also 

known as climbing poison ivy, common poison 

ivy, poison ivy, poison vine, and bemberecua, is a 

deciduous vine or shrub capable of ascending into 

tree crowns 20 m or more and along the ground for 

as much as 45 m. The stems sometimes reach 15 

cm in diameter. The older stems are brown and 

covered with brown adventitious roots by which 

they cling to tree trunks and other surfaces and 

layer (root) whenever they come in contact with 

the ground. The small stems are lighter colored 

and covered with dark brown dots. The alternate 

leaves have long (to 15 cm) petioles and three 

ovate to lanceolate leaflets 4 to 20 cm long with 

entire, coarsely toothed, or lobed margins. Small 

white or greenish white flowers grow in axillary 

panicles. Clusters of white or greenish-white 

drupes 3 to 6 mm in diameter develop later and 

remain on the vine well after the leaves fall. The 

leaves are dark green during the summer, turning 

yellow to red in the fall. The species has 2n = 30 

chromosomes (Long and Lakela 1976, Pavek 

1992, Secretaría de Madio Ambiente y Recursos 

Naturales 2003). 

Range.—Eastern poison ivy is native to Prince 

Edwards Island and from Nova Scotia to southern 

Ontario of Canada, all of the Eastern and 

Midwestern States of the United States (except for 

North Dakota), Arizona, Mexico, Guatemala, the 

Bahamas, Bermuda, China, Taiwan, and Japan 

(Brooks 2003, Griffiths 1994, Natural Resources 

Conservation Service 2003, Pavek 1992). The 

species has been planted as an ornamental and 

medicinal plant and presumably naturalized in 

Great Britain, Europe, and Australia (Schwartz 

1999). 

 There are at least seven subspecies of 

eastern poison ivy. Subspecies divericatum occurs 

in Arizona, ssp. eximium ranges from Texas to 

Guatemala, ssp. negundo grows in the Midwestern 

and Northeastern sections of North America, 

pubens is native to the Southwestern States, ssp. 

radicans ranges across the Southern and Eastern 

portions of U.S., ssp. verrucosm occurs in Texas, 

Oklahoma, and Missouri, and ssp. barkleyi

inhabits the Far East portion of the range (Griffiths 

1994, Natural Resources Conservation Service 

2003). 

Ecology.—Eastern poison ivy grows in soils of all 

textures from nearly all parent materials. Although 

it grows best in fertile, near-neutral, moist but well 

drained soils, it tolerates a wide range of fertility, 

pH, and other conditions. Mean annual 

precipitation ranges from 391 mm to 1,450 mm in 

the United States and Canada. It grows from sea 

level on the Eastern and Gulf Coasts to 2,158 m in 

elevation in New Mexico. In Florida swamps, it 

survives inundation by fluctuations in water level 

and tolerates mildly saline water. An average 

growing season of 150 days with a mean annual 

temperature of 5.3 °C occurs at the northern extent 
 

770 

of its range. There is an average growing season of 

240 days in southern Florida, and a mean annual 

temperature of 19 °C in Arizona (Pavek 1992). 

The mean annual temperatures are probably 

higher, precipitation greater, and growing seasons 

may approach 365 days in Central America. 

Eastern poison ivy is moderately tolerant of shade. 

It is often the most important understory plant, 

climbs nearly every tree in many Southern river 

floodplain forests, and is a component of most 

forest and vegetation types in its ample range. It 

also grows in full sunlight in forest openings, 

fencerows, bluffs, railroad tracks, and vacant lots. 

Eastern poison ivy grows in old-growth forests in 

some portions of its range and is an early invader 

during succession in many environments. It is 

tolerant of fire, browsing, insects, and disease, and 

competes aggressively with weeds and brush. 

Reproduction.—Eastern poison ivy blooms from 

May through June in Virginia (Von Essen 2000) 

and from March through May in Mexico 

(Secretaría de Madio Ambiente y Recursos 

Naturales 2003). The start of flowering 

corresponds with leaves being about half formed 

in the spring (Pavek 1992). The flowers are 

pollinated by insects, including honey bees (Apis 

mellifera L.) (Pollen Lab 2003). Good crops of 

fruits and seeds are produced annually. Seeds are 

dispersed by birds and mammals and sometimes 

by water. Seeds that have been regurgitated from 

the crop or passed through the digestive tract of 

birds demonstrate excellent germination (Pavek 

1992). Eastern poison ivy spreads extensively by 

means of scandent stems that root along their 

whole lengths and below-ground stems (rhizomes) 

that send up shoots frequently. 

Growth and Management.—Eastern poison ivy 

grows rapidly after establishment and assumes 

several forms. It scrambles along the ground and 

sometimes forms thick mats. When given the 

opportunity, it ascends trees as a woody vine. 

Plants take 3 years to reach the flowering stage 

from seed (Pavek 1992). It often forms diffuse 

stands of short upright stems. It can be partially 

controlled by grazing, but there is a resurgence of 

Eastern poison ivy after secession of grazing 

because of the reduction in competition. Spot 

spraying or wiping with the herbicide glyphosate 

during the active growing season can effectively 

control it (Pavek 1992). Grubbing and mowing 

offers some degree of control but puts the worker 

at risk of contracting dermatitis. 

Poison Effects.—The pale yellow oil called 

urushiol, present in all plant parts, is the source of 

all the allergenic pain and suffering associated 

with this species. Even in minute amounts, if it 

stays in contact with human skin for more than 5 

minutes, it will be absorbed and metabolized. The 

metabolites bind with proteins, forming 

compounds that the immune system recognizes as 

“foreign” and attacks. The immune response 

results in rash, blisters, and an intense itch 

(Borialforest.org 2003) that lasts for 1 to 3 weeks. 

Sensitivity can vary over the life of a person. A 

few people have life-threatening reactions to the 

allergen (Schwartz 1999). The chemical is the 

same and the effect (called rhus dermatitis) is the 

same for all Toxicodendrons. Ten to 15 percent of 

the U.S. population is completely immune to rhus 

dermatitis, 25 to 35 percent react only to high 

doses, and 50 percent have a consistent reaction to 

relatively small doses of the allergen (Family 

Practice Notebook.com 2003). The best defense is 

to avoid contact with the plant, especially with the 

sap or smoke from burning Toxicodendron

material. The next line of defense is to quickly 

wash the exposed skin with soap and copious 

amounts of water, or if not available, with mud 

and water or an alkaline material such as baking 

soda or wood ashes and water. After a rash 

develops, the sufferer is advised to apply 

hydrocortisone cream or any of several creams 

sold over-the-counter for the treatment of rhus 

dermatitis. A folk remedy mentioned in the 

literature (Felter 1922) and found by the author to 

almost instantly and completely (but temporarily) 

stop the itch and irritation of rhus dermatitis is to 

rub the mashed juicy stems of jewel weed 

(Impatiens spp.) on the affected area. 

Benefits.—Eastern poison ivy, for its dark green 

foliage during the summer and bright fall colors, 

adds considerably to the beauty of forests and 

shrublands where it grows. It has been used to a 

limited extent as an ornamental and for 

conservation plantings. Eastern poison ivy is an 

important browse species. It is one of the seven 

most important food species for white-tailed deer 

in Indiana (Pavek 1992). Goats are said to like 

eastern poison ivy and have been used to control it 

(Schwartz 1999). The fruits are an important fall, 

winter, and early spring food for upland game 

birds and many species of song birds (Pavek 

1992). Rabbits and rodents also eat the leaves and 

fruits. In herbal medicine, extracts of the leaves 

have been and still are used to treat herpetic 
 

771

eruptions, palsy, and rheumatism. Small internal 

doses will act as a sedative but must be used with 

care (Felter 1992, Grieve 2003). 

References

Borialforest.org. 2003. Toxicodendron radicans, 

poison ivy. http://www.lakeheadu.ca/~borfor/ 

world/herbs_shrubs/poison_ivy.htm. 2 p. 

Brooks, B. 2003. The Toxicodendrons: poison ivy, 

poison oak and poison sumac. http://nac. 

tamu.edu/x075bb/caddo/frameidx.html. 7 p. 

Family Practice Notebook.com. 2003. Rhus 

dermatitis. http://www.fpnotebook.com/ 

Der10.htm. 4 p. 

Felter, H.W. 1922. The eclectic material medica, 

pharmacology and therapeutics: Rhus 

toxicodendron. http://www.ibiblio.org/herbmed/ 

eclectic/felter/toxicodendron-radi.html. 6 p. 

Grieve, M. 2003. A modern herbal: ivy, poison. 

http://www.botanical.com/botanical/mgmh/i/ 

ivypoi17.html. 3 p. 

Griffiths, M. 1994. Index of garden plants. Timber 

Press, Portland, OR. 1,234 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. Banyan Books, Miami, FL. 

962 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Toxicodendron radicans (L.) 

Kuntze. http://plants.usda.gov/cgi_profile. 

cgi?symbol=TORA2. 6 p. 

Pavek, D.S. 1992. Toxicodendron radicans. In: 

U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/vine/ 

toxrad/all.html. 29 p. 

Pollen Lab. 2003. Pollen descriptions: 

Toxicodendraon radicans. http://pollen.usda. 

gov/Pollen_descriptions/Toxicodendron_radican

s.htm. 1 p. 

Schwartz, D. 1999. Poison ivy (Toxicodendron 

radicans (Rhus radicans). http://www.cloudnet. 

com/~djeans/FlwPlant/PoisonIvy.htm. 4 p. 

Secretaría de Madio Ambiente y Recursos 

Naturales. 2003. Especies con usos no 

maderables en bosques tropicales y 

subtropicales: Toxicodendron radicans (L.) 

Kuntze http://www.semarnat.gob.mx/pfnm2/ 

fichas/toxicodendron_radicans.htm. 2 p. 

Von Essen, T. 2000. Poison Ivy, Toxicodendron 

radicans. Wildwood Park, Radford, VA. 

http://www.radford.va.us/Community/wildwood

/today/Plant_articles/Poison_ivy.htm. 2 p. 
 

772 

Toxicodendron rydbergii (Small ex Rydb.) Greene western poison ivy 

ANACARDIACEAE 

Synonyms: Rhus radicans L. var. rydbergii (Small ex Rydb.) Rehd. 

 Rhus radicans L. var. vulgaris (Small ex Rydb.) Rehd. 

 Rhus toxicodendron L. var. vulgaris Michx. 

 Toxicodendron desertorum Lundell 

John K. Francis 

General Description.—Western poison ivy, also 

called poison oak, poison ivy, and Ryberg’s poison 

ivy, is a deciduous, normally upright shrub, 

occasionally a scrambling vine, usually 20 to 100 

cm in height with stems 4 to 10 mm in diameter. 

The woody stems, which arise from underground 

stems that extend up to 2 m from the parent plant, 

are sparingly branched and have smoothish, brown 

to gray bark, with dark brown lenticels, and wooly, 

brown, winter buds. The alternate, bright-green 

leaves are trifoliate with long petioles. Leaflets are 

ovate to rhombic, 3 to 15 cm long, the lateral 

leaflets usually somewhat oblique and shorter than 

the central leaflet. Leaflet margins are lobbed, 

rounded-dentate, or entire. Small yellowish 

flowers are grouped in dense axillary panicles. The 

fruits, which remain attached throughout the 

winter, are ivory-colored, globose drupes 4 to 6 

mm in diameter and contain one seed each. There 

are 2n = 30 chromosomes (Lasica 2002, St. John 

1963, Welsh and others 1987). Western poison ivy 

is similar in appearance to eastern poison ivy [T. 

radicans (L.) Kuntze] except that it is a shrub and 

rarely and weakly climbs. It is known to hybridize 

with eastern poison ivy (Rook 1998). 

Range.—Western poison ivy is native across 

southern Canada from Nova Scotia to British 

Colombia and throughout the United States except 

California and New Jersey and from Missouri, 

Kentucky, and South Carolina southward 

(McMurray 1988, Natural Resources Conservation 

Service 2003). Although it has been planted 

outside its natural range (botanic gardens, etc.), 

there are not reports of it escaping. 

Ecology.—Western poison ivy grows on a wide 

range of sites--on river bottoms, streambanks, 

disturbed upland sites such as roadsides, 

fencerows, sand dunes, logged areas, orchards, 

talus slopes in subalpine areas, and around seeps 

and on shaded canyon sides in cold desert areas. It 

grows from 600 to 2,500 m elevation in the 

interior West and probably near sea level in coastal 

areas. It tolerates short-term flooding. The species 

grows in both full sun and under forest canopies. It 

colonizes all soil textural types derived from most 

parent materials with pH’s from mildly acid to 

mildly alkaline. Western poison ivy often grows 

on skeletal soils, rocky slopes, and cliff sides. The 

species invades quickly after disturbance. Bare 

mineral soil is the best surface for seed 

germination and seedling establishment (Moye 

1999). It often forms clonal thickets. Western 

poison ivy tolerates fire well by sprouting from 

underground stems but does not increase in density 

as a result (McMurry 1988, Welsh and others 

1987). 

Reproduction.—Flowering occurs in late spring 

through early summer. Plants on the Great Plains 

sometimes flower a second time in August or 

September (McMurry 1988). Good fruit and seed 

crops occurs almost every year. The seeds are 

dispersed by birds, mammals, and gravity. 
 

773

Seedlings are necessary to establish new colonies 

but vegetative (by sprouts from underground 

stems) reproduction is more important for local 

spread. 

Growth and Management.—Western poison ivy 

grows at a moderate rate. Clones are relatively 

stable over time. Although it is not as invasive as 

eastern poison ivy, it is often desirable to eliminate 

it from areas of heavy human use. Spot spraying or 

wiping with the herbicide glyphosate during the 

active growing season can effectively control 

eastern poison ivy (Pavek 1992) and probably will 

be effective for western poison ivy. Grubbing and 

mowing offers some degree of control but puts the 

worker at risk of contracting dermatitis so that 

protective clothing should be worn during the 

operation. Branches and roots should be piled to 

rot in an out-of-the-way place or bagged and sent 

to the landfill, never burned. 

Poison Effects.—The pale yellow oil called 

urushiol, present in all plant parts, is the source of 

the allergenic pain and suffering associated with 

this species. If even a minute amount (nanograms) 

the oil stays in contact with human skin for more 

than 5 minutes, it will be absorbed and 

metabolized. The metabolites bind with proteins, 

forming compounds that the immune system 

recognizes as “foreign” and attacks. The immune 

response results in rash, blisters, and an intense 

itch (Borialforest.org 2003) that lasts for 1 to 3 

weeks. Sensitivity can vary over the life of a 

person. A few people have life-threatening 

reactions to the allergen (Schwartz 1999). The 

chemical is the same and the effect (called rhus 

dermatitis) is the same for all Toxicodendrons. Ten 

to 15 percent of the U.S. population is completely 

immune to rhus dermatitis, 25 to 35 percent react 

only to high doses, and 50 percent have a 

consistent reaction to relatively small doses of the 

allergen (Family Practice Notebook.com. 2003). 

The best defense is to avoid contact with the plant, 

especially with the sap or smoke from burning 

Toxicodendron material. The next line of defense 

is to quickly wash the exposed skin with soap and 

copious amounts of water, or if not available, with 

mud and water or an alkaline material such as 

baking soda or wood ashes and water. After a rash 

develops, the sufferer is advised to apply 

hydrocortisone cream or any of several creams 

sold over-the-counter for the treatment of rhus 

dermatitis. A folk remedy mentioned in the 

literature (Felter 1922) and found by the author to 

almost instantly and completely (but temporarily) 

stop the itch and irritation of rhus dermatitis is to 

rub the mashed juicy stems of jewel weed 

(Impatiens spp.) on the affected area. 

Benefits.—Western poison ivy helps protect the 

soil and adds beauty in the summer with its dark 

green leaves and in autumn with bright colors to 

forest and shrublands. Although it causes browsing 

animals no ill effects, western poison ivy is low in 

protein and energy and consequently is only 

occasionally grazed by domestic animals and wild 

ungulates. However, the fruits are eaten by quail, 

wild turkeys, and some songbirds. It does provide 

cover for small mammals and birds (McMurray 

1988). In herbal medicine, extracts of the leaves of 

this and other Toxicodendron species have been 

and still are used to treat herpetic eruptions, palsy, 

and rheumatism. Small internal doses will act as a 

sedative but must be used with care (Grieve 2003). 

References 

Borialforest.org. 2003. Toxicodendron radicans, 

poison ivy. http://www.lakeheadu.ca/~borfor/ 

world/herbs_shrubs/poison_ivy.htm. 2 p. 

Family Practice Notebook.com. 2003. Rhus 

dermatitis. http://www.fpnotebook.com/ 

Der10.htm. 4 p. 

Felter, H.W. 1922. The eclectic material medica, 

pharmacology and therapeutics: Rhus 

toxicodendron. http://www.ibiblio.org/herbmed/ 

eclectic/felter/toxicodendron-radi.html. 6 p. 

Grieve, M. 2003. A modern herbal: ivy, poison. 

http://www.botanical.com/botanical/mgmh/i/ 

ivypoi17.html. 3 p. 

Lasica, P. 2002. A flora of Glacier National Park, 

Montana. Oregon State University Press, 

Corvallis, OR. 512 p. 

McMurray, N.E. 1988. Toxicodendron rydbergii. 

In: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Research Station, Fire 

Sciences Laboratory, Fire Effects Information 

System. http://www.fs.fed.us/database/feis/ 

plants/shrub/ toxryd/all.html. 15 p. 

Moye, H. 1999. The Toxicodendrons: poison ivy, 

poison oak, and poison sumac. 
 

774 

http://chiron.valdosta.edu/rgoddard/biol4900/mo

ye/moye.html. 7 p. 

Natural Resources Conservation Service. 2003. 

Plants profile: Toxicodendron rydbergii (Small 

ex Rydb. Greene. http://plants.usda.gov/ 

cgi_profile.cgi?symbol=TORA2. 5 p. 

Pavek, D.S. 1992. Toxicodendron radicans. In: 

U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station, Fire Sciences 

Laboratory, Fire Effects Information System. 

http://www.fs.fed.us/database/feis/plants/vine/ 

toxrad/all.html. 29 p. 

Rook, E.J.S. 1998. Toxicodendron rydbergii, 

poison ivy. http://www.rook.org/earl/ 

bwca/nature/shrubs/toxicodendron.html. 4 p. 

Schwartz, D. 1999. Poison ivy (Toxicodendron 

radicans (Rhus radicans). http://www.cloudnet. 

com/~djeans/FlwPlant/PoisonIvy.htm. 4 p. 

St. John, H. 1963. Flora of Southeastern 

Washington and adjacent Idaho. Outdoor 

Pictures, Escondido, CA. 583 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 
 

775

Trema lamarckianum (J.A. Schultes) Blume West Indian trema 

ULMACEAE 

Synonyms: Celtis lamarckiana Roem. & Schult. 

 Sponia lamarckiana (Roem. & Schult.) Decne. 

 Celtis lima Lam. 

 Trema lima authors, not Blume 

John K. Francis 

General Description.—West Indian trema, also 

known as Lamarck trema, pain-in-back, cabrilla, 

memizo de majagua, capulí cimarrón, and orme 

petites feuilles, is an evergreen shrub or small tree 

to 8 m in height and 20 cm in diameter at breast 

height. The plant is supported by a system of tancolored, stiff, but flexible, tap, lateral, and fine 

roots. There is usually a single main stem. It is 

covered by smoothish, light brown to gray bark 

with many tiny warty lenticels over a light brown 

to pinkish, fibrous inner bark. The species has a 

monopodial branching pattern, in which the main 

stem produces a continuous progression of fine 

lateral branches (Massey and Murphy 1996) some 

of which may thicken and become major branches 

in older plants. The alternate leaves, which are 

attached to twigs by petioles 8 to 10 mm long, 

have ovate-lantiolate, green to gray-green blades 

that are rough on both surfaces and have finetoothed edges. The 2-mm diameter, greenish 

flowers are clustered at the leaf axils. The fruits 

are rounded, fleshy, 1- to 3-mm pink drupes. The 

fruits have little flavor and each contain one 

brown seed (Howard 1988, Liogier 1985, Little 

and Wadsworth 1964). 

Range.—West Indian trema is native to Florida, 

Bermuda, the Bahamas, Grand Cayman, and the 

Greater and Lesser Antilles south to St. Vincent 

(Liogier 1985, Little and Wadsworth 1964). 

Ecology.—West Indian trema is a pioneer species. 

It colonizes disturbed sites on a wide variety of soil 

types over both sedimentary and igneous rocks. It is 

especially common in ultramafic (serpentine) areas. 

Areas in Puerto Rico with West Indian trema 

populations receive annual rainfall ranging from 

about 1200 to about 3000 mm at elevations ranging 

from a few meters above sea level to about 900 m. 

The species is intolerant of shade and usually grows 

in areas with a sparse to moderate cover of herbs, 

shrubs and trees. It does not grow under a closed 

forest canopy. Common sites are road cuts and fill, 

abandoned roads, unstable slopes, landslides, 

mechanically disturbed sites, and hammocks 

(author’s observation, Long and Lakela 1976). 

Reproduction.— After West Indian trema reaches 

about 1 m, it flowers and fruits continuously 

throughout its life. During periods of favorable 

moisture, plants produce fruits and seeds in large 

numbers. A collection of fruits from Puerto Rico 

weighed an average of 0.0077 + 0.0001 g/fruit. 

Air-dried seeds cleaned from them averaged 

0.0024 + 0.0000 g/seed or 416,000 seeds/kg. 

Placed in commercial potting mix, 38 percent of 

the seeds germinated between 60 and 120 days 

after sowing. Birds disperse the seeds. Seedlings 

are common on disturbed ground near seedbearing plants. Young plants sprout when cut. 

Growth and Management.—Early seedling 

growth is slow. Later, growth reaches about 1 

m/year, slowing as the plants get old. West Indian 
 

776 

trema lives about 10 to 20 years. The species has 

not been reported as weedy. Reproduction 

probably can be promoted by scarifying the soil 

near seed-bearing plants before the start of 

seasonal rains. 

Benefits.—West Indian trema is an important 

colonizer of disturbed areas and helps protect the 

soil from erosion. It has not been used for, but 

might be useful for, site stabilization plantings. 

West Indian trema is listed as a nitrogen-fixing 

species (Winrock International 2002). The wood is 

soft and seldom used. The fruits are one of the 

most important foods of the endangered Puerto 

Rican plain pigeon, Columba inornata wetmorei

(Division of Endangered Species 2002). 

References 

Division of Endangered Species. 2002. Puerto 

Rican plain pigeon, Columba inornata 

wetmorei. U.S. Fish and Wildlife Service. 

http://endangered. Fws.gov/i/b/sab40.html. 3 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

Tropical Florida. University of Miami Press, 

Coral Gables, FL. 962 p. 

 

Massey, J.R. and J.C. Murphy. 1996. Vascular 

plant systematics. Section B. General characters 

and character states: 9. Patterns. http://www. 

ibiblio.org/botnet/glossary/b_ix.html. 2 p. 

Winrock International. 2002. Nitrogen fixing trees 

and shrubs. http://www.winrock.org/forestry/ 

factpub/nftlist.htm. 29 p. 
 

777

Trichilia hirta L. broomstick 

MELIACEAE 

Synonyms: Trichilia spondioides Jacq. 

 Trichilia wawrana antillana C. DC. 

John K. Francis 

General Description.—Broomstick is also known 

as broomwood, cabo de hacha, garbancillo, guaita, 

jojobán, jabillo, molinillo, palo de anastasio, 

retamo, canalete, conejo colorado, mata piojo, 

carrapeta, and mombin bâtard. It is an evergreen to 

semideciduous shrub or small tree ranging in 

mature height from 3 to 15 m in different parts of 

its range. The shrubs usually grow with a single 

stem in closed stands but often have multiple 

stems when open-grown or when they have been 

damaged. The bark is brown or gray, rough and 

lightly furrowed or scaly. Sapwood is yellowish 

white, and the heartwood is reddish brown with 

darker streaks. Twigs are green, becoming brown 

with age. The plants are supported by a tap and 

extensive lateral root system of flexible, somewhat 

brittle roots with thin, reddish-brown bark. The 

alternate leaves are pinnately compound, 20 to 35 

cm in length with nine to 21 ovate to lanceolate 

leaflets. The inflorescences are unisexual axillary 

racemes. However, these plants are dioecious. The 

flowers are small and greenish white. The two- to 

four-chambered capsules split open to expose one 

or two seeds per chamber. The 6- to 8-mm ovular 

seeds are covered by a thin, scarlet to orange aril 

(Howard 1988, Liogier 1988, Little and 

Wadsworth 1964, Stevens and others 2001). 

Range.—Broomstick is native to the West Indies 

and from Mexico through Central America to 

Brazil and Bolivia in South America (Killeen and 

others 1993, Little and Wadsworth 1964). It has 

been planted in southern Florida (Little and 

Wadsworth 1964). 

Ecology.—Broomstick is relatively common and 

widely scattered through most of its range. It does 

not form stands but tends to be scattered in open 

forests, roadsides, brushy pastures, and disturbed 

areas. Although the species is found in primary 

remnants, it is more common in secondary forests, 

usually with low basal areas. Broomstick may be 

found in Puerto Rican forests that receive from 

750 to about 1700 mm of mean annual 

precipitation at elevations from near sea level to 

600 m. It grows in Nicaragua in deciduous to 

moist forests from 0 to 1,000 m elevation (Stevens 

and others 2001). Broomstick is reported in 

mountain rainforests up to 3,000 m in elevation in 

Bolivia (Killeen and others 1993). Although the 

species grows on a wide range of soil types, it is 

more often seen on rocky and infertile sites where 

competition is less severe. Broomstick can grow in 

partial shade but requires good overhead sunlight 

to flower and fruit. 

Reproduction.—Broomstick flowers and fruits 

throughout most of the year (Little and Wadsworth 

1964). The species is reported to flower from 

March through May and fruit from January 

through April in Nicaragua (Stevens and others 

2001) and flowers from May through July in 

Panama. The flowers are insect pollinated (Center 

for Tropical Forest Science 2002). Plants receiving 

ample sunlight can produce several hundred seeds 

per year. A collection of fresh fruits from Puerto 

Rico weighed an average of 0.129 + 0.0019 g/fruit. 

Surface-dry seeds separated from them weighed an 

average of 0.0891 + 0.0020 g/seed or 11,000 

seeds/kg. Sown without pre-treatment on moist 
 

778 

blotter paper, 100 percent of them germinated 

between 17 to 232 days after sowing. Germination 

is hypogeal, and they appear to be recalcitrant 

(lose viability if dried beyond a certain point) 

(author’s observation). In Panama, the seeds are 

animal-dispersed (Center for Tropical Forest 

Science 2002). In Puerto Rico, seedlings are 

relatively common but only an occasional 

individual lasts more than a few weeks or months. 

Growth and Management.—Broomstick in 

Puerto Rico appears to have a moderate growth 

rate and to live about 10 to 30 years. Management 

information is not available. 

Benefits.—Broomstick adds to secondary forest 

biodiversity, helps protect the soil, and furnishes 

food and cover for wildlife. The species is used to 

a limited extent for landscaping and is considered 

a honey plant. The wood is of medium density 

(specific gravity 0.5), moderately hard and 

somewhat brash, and is resistant to attack by drywood termites (Criptotermes brevis (Walker)). 

Another source lists the wood density as 0.48 to 

0.96 at 12 to 15 percent moisture content (World 

Agroforestry Centre 2002). It has been used for 

oars, broom handles, and rustic furniture; today the 

wood is mainly used for fuel, stakes, and fence 

posts (Little and Wadsworth 1964). An infusion of 

leaves, flowers, and roots has been used as a 

treatment for skin ulcers and as an emetic that has 

the unfortunate side effect of causing abortions 

and bleeding. Young branches were strewn on 

floors to control fleas and lice (Núñez Meléndes 

1982). 

References

Center for Tropical Forest Science. 2002. Arboles 

del area de Canal de Panama. Smithsonian 

Institution, Washington, DC. http://ctfs.si.edu/ 

webatlas/spanish/tri2hi.html. 2 p. 

Howard, R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Killeen, T.J., E. García-E., and S.G. Beck, eds. 

1993. Guía de árboles de Bolivia. Herbario 

Nacional de Bolivia and the Missouri Botanical 

Garden, La Paz, Bolivia and St. Louis, MO. 

958 p. 

Liogier, H.A. 1988. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

2. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 481 p. 

Little, E.L., Jr. and F.H. Wadsworth. 1964. 

Common trees of Puerto Rico and the Virgin 

Islands. Agriculture Handbook 249. U.S. 

Department of Agriculture, Forest Service, 

Washington, DC. 548 p. 

Núñez-Meléndez, E. 1982. Plantas medicinales de 

Puerto Rico. Editorial de la Universidad de 

Puerto Rico, Río Piedras, PR. 498 p. 

 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden Press, St. Louis, 

MO. p. 945-1,910. 

World Agroforestry Centre. 2002. Trichilia hirta. 

http://www.worldagroforestrycentre.org/sea/ 

AgroModels/Dbases/WD/asps/DisplayDetail.asp

?SpecID=3487. 1 p. 
 

779

Trichostigma octandrum (L.) H. Walt. hoop vine 

PHYTOLACCACEAE 

Synonyms: Rivinia octandra L. 

 Rivinia humilis var. scandens L. 

 Rivinia scandens (L.) Miller 

 Trichostigma rivinoides A. Rich. in Sagra 

 Villamilla octandra (L.) Hook. f. in Benth. & Hook. 

John K. Francis 

General Description.—Hoop vine, which is also 

known as basket wiss, bejuco de paloma, bejuco 

de nasa, bejuco de palma, murette, liane à 

barriques, and liane à terre, is a scrambling shrub 

that may extend 10 m laterally or 6 m up into the 

crowns of trees and develops stems with diameters 

of 5 to 15 cm. These plants produce weak 

taproots, extensive lateral roots and many fine 

roots. The stems are gray, cylindrical, smooth, and 

relatively stiff. Older plants often have several 

branches near the base and then few branches until 

near the growing tips. Young branches of hoop 

vine are cylindrical, slender, and covered with 

elongated lenticels. The leaves are alternate, 

elliptic to lanceolate, 5 to 15 cm long and 1.5 to 6 

cm broad. The petioles and young leaves are 

reddish. The inflorescence is a many-flowered 

raceme about 15 cm long, often terminal on a 

leafless stem. The flowers are about 10 mm 

across, greenish-white, becoming red as the fruits 

develop. Hoop vine fruits are elliptic, about 5 to 6 

mm in diameter, and purple to black when fully 

ripe. They have little flavor but leave a slightly 

unpleasant aftertaste. The seeds are 4 to 5 mm in 

diameter, black, and shiny (Acevedo-Rodríguez 

1985, Howard 1988, Liogier 1985). 

Range.—Hoop vine is native to southern Florida, 

throughout the West Indies, and from Mexico to 

northern Argentina (Anonymous 2002, Howard 

1988, Liogier 1985). Although relatively common 

in most other areas, the species is listed as 

critically endangered in Florida (Gann and others 

2001). 

Ecology.—Although hoop vine has no specialized 

climbing structures, it often ascends into the lower 

and mid-crowns of trees. The shrub also covers 

windfalls, fences, rocks, and forms mounds of 

tangled stems to 2 m high in open areas. Hoop 

vine grows on soils of all textures in a wide range 

of pH’s derived from both igneous, metamorphic 

(including ultramafics) and sedimentary (including 

limestone) rocks. In Puerto Rico, the species 

occurs in areas receiving from 900 mm to 2400 

mm of precipitation. In Costa Rica, herbarium 

specimens have been collected from near sea level 

to 1,100 m in elevation (Anonymous 2002). Hoop 

vine is moderately intolerant of shade. It invades 

early successional forest and continues at least 

into the late secondary forest stage, and also 

occurs in remnant forests, especially in tree-fall 

gaps and on the edges of natural openings. In 

farms and urban areas, the species grows on 

roadsides, fencerows, woodlots, brushy pastures, 

brushy vacant lots, and stream bottom galleries. 

Reproduction.—In Puerto Rico, hoop vine plants 

flower from April to August and again in October 

(Acevedo-Rodríguez 1985). In Costa Rica, 

herbarium specimens were collected with fruit and 

flowers from November through June (Instituto 

Nacional de Biodiversidad 2002). Fruits collected 

in Puerto Rico weighed an average of 0.178 +

0.007 g/fruit. Air-dried seeds separated from them 

averaged 0.0324 + 0.0004 g/seed or 31,000 
 

780 

seeds/kg. Sown in commercial potting mix, 32 

percent of these seeds germinated between 10 and 

71 days after sowing. Birds apparently disperse 

the seeds. Natural seedlings are not common. 

Stems layer (root) whenever they are covered with 

soil or organic material. 

Growth and Management.—Growth of hoop 

vine is rapid from sprouts, as much as 3 m/year. 

Established plants appear to live for several 

decades, about as long as the trees they use for 

support. No management experience has been 

published. 

Benefits.—The young leaves of hoop vine are 

cooked and eaten as a vegetable after the cooking 

water is discarded to remove the bitter flavor 

(Educational Concerns for Hunger 2002). Both the 

split stems and the bast fibers of hoop vine are 

used for making baskets. The stems are also used 

for making bent furniture and craft items (Ruiz 

2002). Juice from the fruits leaves a purple stain 

on whatever it contacts and may have potential as 

a natural stain for cloth and crafts. The species 

contributes to biodiversity and soil stability, and 

provides food and cover for wildlife. In herbal 

medicine, the vegetative parts are used as a 

treatment for heart palpitations, and a powder of 

the root bark is applied topically to treat colds and 

water retention (Liogier 1990). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Anonymous. 2002. Contribución a la flora polínica 

de Noreste Argentina. http://www.unne.edu.ar/ 

cyt/2001/6-Biologicas/B-048.pdf. [not paged]. 

Educational Concerns for Hunger. 2002. Tropical 

vegetables. http://www.echonet.org/eln&herbs/ 

eln_catalog/veggies.htm. 4 p. 

Gann, G.D., K.A. Bradley, and S.W. 

Woodmansee. 2001. Status of South Florida 

flora. Institute of Regional Conservation. 

http://www.Regionalconservation.org/CH2.htm. 

10 p. 

Howard R.A. 1988. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae, Part 1. Vol. 4. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 673 p. 

Instituto Nacional de Biodiversidad. 2002. Lista de 

especimenes de Trichostigma octandrum. http:// 

www.inbio.ac.cr.bims/k03/p13/c045/o0245/f015

56/g008574/s026880.htm. 2 p. 

Liogier H.A. 1985. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vol. 

1. Editorial de la Universidad de Puerto Rico, 

Río Piedras, PR. 352 p. 

Liogier, H.A. 1990. Plantas medicinales de Puerto 

Rico y del Caribe. Iberoamericana de Ediciones, 

Inc., San Juan, PR. 566 p. 

Ruiz, D.L. 2002. Cuba. Food and Agriculture 

Organization of the United Nations. http:// 

www.fao.org/docrep/t2354s/t2354s0r.htm. 4 p. 
 

781

Triumfetta semitriloba Jacq. burweed 

TILIACEAE 

Synonyms: Triumfetta hirta Vahl 

 Triumfetta ovata DC. 

 Triumfetta tiliifolia Vahl 

 Heliocarpus hirta (Vahl) R.O. Williams & Sandw. 

John K. Francis 

General Description.—Burweed is also known as 

Sacramento bur, black bush, cadillo de perro, 

cousin-petit, petit mahot-cousin, mahot-cousin 

rouge, tête à nègre, and nedyah (Howard 1989, 

Liogier 1994, Pacific Island Ecosystems at Risk 

2001). It is a single-stemmed, short-lived shrub 

that reaches 1 to 2 m in height and 3 cm in stem 

diameter. The stems have gray, smooth bark and a 

woody center. Burweed plants are supported by a 

weak taproot and many lateral roots. The roots are 

tan colored, flexible, and have a fleshy outer layer 

over a woody center. The leaves have petioles 

from 0.8 to 9.5 cm long, growing progressively 

shorter toward the plant apex. The leaves, 3 to 8 

cm long, are broadly ovate to rhombic-ovate, and 

sometimes three-lobed, nearly hairless above, but 

densely stellate-pubescent below. The flowers, 

which are located at the leaf axils, have yellow 

petals about 5 mm long. The fruits, round burs 6 to 

8 mm in diameter with numerous hooked spines, 

are borne in groups of two or three. Each fruit has 

three compartments and contains three seeds, if 

complete (Howard 1989, Liogier 1994, Long and 

Lakela 1976). 

Range.—Burweed is native to southern Florida, 

Bermuda, the West Indies, Mexico, Central 

America, and South America (Howard 1989, 

Liogier 1994, Long and Lakela 1976). It was 

thought that burweed had been recently introduced 

to Rapa Nui (Easter Island), but fossil pollen 

indicates that it has been present for at least 

35,000 years (National Geographic Society 2001). 

The species has naturalized in Hawaii, Guam, 

Western Samoa and a number of other Pacific 

Islands, and in fact is reported to be a pantropical 

weed (Pacific Island Ecosystems at Risk 2001). 

Ecology.—Burweed grows in pinelands and 

hammocks in Florida (Long and Lakela 1976) and 

in roadsides, neglected pastures, and wastelands in 

Puerto Rico. It grows in sand to clay soils with pH 

values from about 5.5 to 8.0. In Puerto Rico, 

burweed grows from near sea level to about 700 m 

in elevation. This diverse habitat receives annual 

precipitation from about 900 to 2200 mm. In 

Hawaii, it grows from sea level to 1,067 m in 

elevation in areas with annual rainfalls of 760 to 

1500 mm (Haselwood and Motter 1966). The 

species needs disturbance for establishment. If 

given an equal start, it competes well with other 

herbs, grasses, and shrubs in full sun or partial 

shade. Burweed may be found growing in small 

patches and as single, dispersed plants. 

Reproduction.—Burweed blooms and fruits 

continuously (Long and Lakela 1976), beginning 

at about 6 months of age. Seed production is 

guaranteed by self-pollination, but crosspollination increases seed set and probably quality 

(Collevatti and others 1997a). The species has a 
 

782 

chromosome count of 2n = 32 (Long and Lakela 

1976). In Brazil, solitary and social bees visit and 

pollinate burweed (Collevatti and others 1997b). It 

is an abundant seed producer. Fruits collected in 

Puerto Rico averaged 0.0040 + 0.0016 g/fruit. 

Seeds separated from those fruits averaged 0.0039 

+ 0.0001 g/seed or 256,000 seeds/kg. Placed on 

filter paper, 41 percent germinated between 10 and 

119 days from sowing. Germination is epigeal. 

Seeds are dispersed when they cling to passing 

animals. 

Growth and Management.—Burweed lives 

between 1 and 3 years. In seasonally dry habitat, 

this species behaves as an annual. The plants 

grow about 1 m the first year, with growth slowing 

in subsequent years. Because of its weedy nature, 

planting burweed is unlikely, and ill-advised 

outside its native range. Indeed, control is often 

needed in croplands and pastures. Treatments 

follow normal practice for coarse broadleaf weeds: 

mechanical cultivation or grubbing and spraying 

with herbicides such as 2,4-D. 

Benefits.—The stem wood has moderate strength 

and hardness, but because of the plant’s small size 

the wood is of little value. Fibers from the stem 

bark were once used by the inhabitants of Rapa 

Nui to make fishing lines (Moanalua Gardens 

Foundation 1999). Burweed is reported to have 

little or no forage value for domestic livestock 

(Haselwood and Motter 1966). 

References 

Collevatti, R.G., M.E.C. Amaral, and F.S. Lopes. 

1997a. Role of pollinators in seed set and a test 

of pollen limitation hypothesis in the tropical 

weed Triumfetta semitriloba (Tiliaceae). Revista 

de Biologia Tropical 45(4): 1,401-1,407. 

Collevatti, R.G., L.A.O. Campos, and J.H. 

Schoereder. 1997b. Foraging behaviour of bee 

pollinators on the tropical weed Triumfetta 

semitriloba: departure rules from flower patches. 

Insectes Sociaux 44(4): 345-352. 

Haselwood, E.L. and G.G. Motter, eds. 1966. 

Handbook of Hawaiian weeds. Hawaiian Sugar 

Planters Association. Honolulu, HI. 479 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Long, R.W. and O. Lakela. 1976. A flora of 

tropical Florida. Banyan Books, Maimi, FL. 

962 p. 

Moanalua Gardens Foundation. 1999. The mystery 

of Rapa Nui. Moanalua Gardens Foundation, 

Honolulu, HI. http://gamma.mhpcc.edu/voyage/ 

moanalua/6_rapa.pdf. 20 p. 

National Geographic Society. 2001. Rapa Nui and 

Sala-y-Gomez subtropical broadleaf forests. 

National Geographic Society, Washington, DC. 

http://nationalgeographic.com/wildworld/ 

profiles/terrestrial/oc/oc0111.html. 3 p. 

Pacific Island Ecosystems at Risk. 2001. Invasive 

plant species: Triumfetta semitriloba Jacaquin, 

Tiliaceae. http://www.hear.org/pier/trsem.htm. 

2 p. 
 

783

Turbina corymbosa (L.) Raf. Christmas vine 

CONVOLVULACEAE 

Synonyms: Rivea corymbosa (L.) Hall.f. 

 Convolvulus corymbosus L. 

 Convolvulus sidaefolius HBK. 

 Ipomoea sidaefolia (HBK.) Choisy 

 Ipomaea corymbosa (L.) Roth. 

John K. Francis 

General Description.—Christmas vine, also 

known as Christmas wreath, aguinaldo blanco, 

corona de novia, ololiuqui, and badoh, is a woody 

vine that may extend 5 m or more laterally and 

into the crowns of trees and shrubs. Older stems 

reach about 2.5 cm in diameter. The gray, threesided stems have many lenticels and may be 

grooved. The young stems are cylindrical and 

tough. The foliage is concentrated on current 

year’s growth. The leaves, with slender petioles, 

have cordate blades 5 to 8 cm long with an 

elongated point. The inflorescences are corymbose 

cymes that arise from leaf axils. The 2.5- to 3-cm 

corolla is trumpet shaped, white with a red or 

purple throat and green or greenish gray radiating 

stripes. The ellipsoidal capsule has three long and 

three short sepals that cause it to spin and glide 

laterally when released. Each capsule contains one 

brown, pubescent seed (Acevedo-Rodríguez and 

Woodbury 1985, Howard 1989, Liogier 1995). 

Range.—Christmas vine is native to the West 

Indies, Mexico through Central America, and the 

tropical portion of South America (Liogier 1995). 

It has naturalized in Florida (Correll and Johnston 

1970), Hawaii, a few other Pacific Islands, 

Australia (Pacific Island Ecosystems at Risk 

2002), and some parts of the Old World Tropics 

(Howard 1989). 

Ecology.—Christmas vine grows as single plants 

or matted patches of vines in secondary forest 

openings, old fields, neglected pastures, road 

sides, stream banks, and vacant lots. In Florida, it 

may be found in hammocks and brushy areas 

(Long and Lakela 1971). It demands full or nearly 

full sunlight and will not grow under a closed 

forest canopy. Christmas vine will grow on most 

well-drained soils. It is reported to bloom more 

profusely on “red” and limey soils (Woman’s 

Club of Havana 1952). The species has been 

observed by the author growing in areas of Puerto 

Rico that receive from 1000 to 2000 mm of annual 

precipitation. 

Reproduction.—A collection of fruits from 

Puerto Rico weighed an average of 0.0389 +

0.0004 g/fruit. Seeds separated from those fruits 

weighed an average of 0.0265 + 0.0002 g/seed or 

38,000 seeds/kg. Sown in commercial potting mix, 

95 percent of the seeds germinated between 33 

and 48 days after sowing (author’s observation). 

Well-established plants root whenever vines touch 

the soil. Natural seedlings tend to be widely 

scattered. Besides spreading by wind, water, and 

lateral vine extension, seeds of the species are now 

widely sold, exchanged, and grown for the 

narcotic the seeds contain. 

Growth and Management.—Seedlings of 

Christmas vine grow slowly at first and develop a 

strong tap and lateral root system. Leaders of wellestablished plants may extend 2 m or more per 

year. The plants can be pruned back to the woody 

stems or thinned in the spring after blooming is 

complete (Woman’s Club of Havana 1952). No 

specific recommendations are given for 

controlling the species when it grows as a weed in 

agricultural settings. Cutting the vines near the 
 

784 

ground, and spraying the resulting sprouts with 

broad-leaf weed killers, should be effective. 

Benefits.—Christmas vine is sometimes grown as 

an ornamental for the clusters of white, scented 

flowers it produces during the early winter. The 

nectar gathered from its flowers makes one of the 

finest honeys (Woman’s Club of Havana 1952). 

Extracts from the seeds are used as an analgesic in 

herbal medicine (Schultes and Hoffmann 1992). 

Narcotic Properties.—The seeds of Christmas 

vine were valued as a sacred hallucinogen by 

Chinantec, Mazatec, Mixtec, Zapotec, and other 

groups in Southern Mexico in Pre-Columbian 

times and are still cultivated and used today as 

aids in divination and witchcraft. It was 

administered by grinding about 13 seeds, adding 

water, filtering, and drinking the filtrate in a quiet, 

secluded place. Hallucinations follow that last 

about 3 hours, sometimes with aftereffects. The 

active ingredients are the ergoline alkaloids, 

lysergic acid amide, and lysergic acid 

hydroxyethylamide that are closely related to LSD 

(Schultes and Hoffmann 1992). 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Correll, D.S. and M.C. Johnston. 1970. Manuel of 

the vascular plants of Texas. Texas Research 

Foundation. Renner, TX. 1,881 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. 

Dicotyledoneae. Part 3. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1995. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 4. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

617 p. 

Long, R.W. and O. Lakela. 1971. A flora of 

Tropical Florida. University of Miami Press, 

Coral Gables, FL. 962 p. 

Pacific Island Ecosystems at Risk. 2002. Invasive 

plant species: Ardisia elliptica Thunberg, 

Myrsinaceae. http://www.hear.org/pier/ 

arell.htm. 2 p. 

Shultes, R.E. and A. Hoffmann. 1992. Plants of the 

gods. Healing Arts Press, Rochester, VT. 192 p. 

Woman’s Club of Havana. 1952. Flowering plants 

from Cuban Gardens. Criterion Books, New 

York. 365 p. 
 

785

Urena lobata L. Caesar weed 

MALVACEAE 

Synonyms: Urena americana L. f. 

 Urena grandiflora DC. 

 Urena trilobata Vell. 

 Urena lobata L. 

 Urena diversifolia Schumach. 

 (and others: Institute of Systematic Botany 2003). 

John K. Francis 

General Description.—Caesar weed is also 

known as hibiscus bur, aramina, pink Chinese 

burr, bur mallow, grand cousin, cadillo, carrapicho 

do mata, malva, mahot cousin, cousin petit, cousin 

rouge, jut africain, cooze mahot, dadangsi, and 

mautofu. It is a subshrub 0.6 to 3 m in height and 

up to 7 cm in basal diameter. The species usually 

has a single stem emerging from the ground but 

normally produces several stems or major 

branches low on the stem and many branches 

throughout. The nearly smooth bark is tough and 

fibrous, brown on the outside and green within. Its 

pale yellow wood is of medium density. Plants are 

supported by a tap and lateral root system of tan or 

ivory-colored, tough and flexible roots. Grayishgreen, discolorous, alternate leaves are stellatepubescent both above and below, ovate, angulate 

or shallowly lobbed, 1 to 12 by 1 to 12 cm, and 

have serrate margins. Axillary flowers are usually 

solitary and have five pink petals. The flowers are 

about 1 cm broad. The fruits are 8- to 10-mm 

globose capsules that break into five fine-barbed 

(glochidate) mericarps. (Howard 1989, Liogier 

1994, Stevens and others 2001) 

Range.—The original range of Caesar weed is 

probably Asiatic. Today it grows throughout moist 

tropic and subtropical regions including American 

Samoa, Florida, Guam, Hawaii, Louisiana, Puerto 

Rico, and the U.S. Virgin Islands (Natural 

Resources Conservation Service 2003, Pacific 

Island Ecosystems at Risk 2003). 

Ecology.—Caesar weed readily invades disturbed 

areas, especially poorly managed pastures, 

scarified and eroded areas, and perennial crop 

plantations but is much less of a problem in annual 

crops. The species does not compete well in tall 

grass swards and brushlands and does not grow 

under forest canopies. Ceasar weed grows on a 

wide variety of soils of varying fertility derived 

from most parent materials. It tolerates salt spray 

and a moderate amount of salt in the soil but does 

not grow in saturated soils. The species ranges to 

elevations of 1,500 m (Pacific Island Ecosystems 

at Risk 2003). In Puerto Rico, it occurs in areas 

that receive from about 1400 to 3000 mm of mean 

annual precipitation, forming thickets in favorable 

sites. 

Reproduction.—Caesar weed flowers and fruits 

throughout the year (Stevens and others 2001). A 

collection of seeds from Puerto Rico averaged 

0.0239 + 0.0003 or 41,800 seeds/kg. Placed on 

moist blotter paper without pretreatment, just 3 
 

786 

percent germinated in 9 months. Germination is 

epigeal (author’s observation). Sulfuric acid 

scarification was tested as a means of breaking 

dormancy of seed from Sierra Leone. The best 

treatment (18 molar solution for 90 minutes) gave 

96 percent germination as compared with distilled 

water that gave 4 percent germination (Harris 

1986). Seeds are dispersed by clinging to fur and 

clothing. Seedlings are common in disturbed sites 

near seed sources. 

Growth and Management.—Caesar weed grows 

rapidly and can reach 0.5 to 2 m by the end of the 

first year. In Puerto Rico, it can live 2 years, 

usually dying back to midheight after the first 

growing season. It is not known whether the 

shrubs sprout from the roots multiple times. A 

fiber crop yielding 1,800 kg/ha is ready to harvest 

after 6 to 7 months and seed crops of 300 to 500 

kg/ha are ready after 7 or 8 months (Fagundes 

2003). Fiber crops are established with seed. 

Because of the aggressive nature of the species, 

wildland plantings are not recommended. The 

author knows of no published specific control 

measures. 

Benefits and Detriments.—Caesar weed 

colonizes disturbed areas and helps to protect the 

soil while furnishing cover for wildlife. It has 

attractive flowers and contributes to aesthetics of 

areas it has colonized. Tens of thousands of tons of 

a jute-like fiber from Caesar weed called aramina 

fiber and Congo jute are produced in Brazil 

(Fagundes 2003) and Africa (Câmara de Comércio 

e Indústria Portugal-Angola 2003). Various 

extracts of leaves and roots are used in herbal 

medicine to treat such diverse ailments as colic, 

malaria, gonorrhea, fever, wounds, toothache, and 

rheumatism (Forest Research Institute of Malaysia 

2003). A semipurified glycoside obtained from 

Caesar weed leaves was 86 percent as effective an 

anti-inflammatory as aspirin in rats (Bautista 

2000). The leaves and flowers are eaten as famine 

food in Africa (Freedman 1998). Raw leaves are 

reported to contain 81.8 percent moisture, 54 cal, 

3.2 g of protein, 0.1 g fat, 12.8 g carbohydrates, 

1.8 g fiber, and 2.1 g ash, 558 mg calcium, and 67 

mg of phosphorous per 100 g (FAO 2003). 

However, the plant is little browsed by cattle and 

can become a severe weed in pastures and 

plantations. Burs that collect on clothing and in 

animal fur are a nuisance. 

References 

Bautista, L.M.A. 2000. Inquiry into the antiinflammatory activity of the syrup from the 

glycosides of the leaves of kulutkulutan (Urena 

lobata, Linn., family Malvaceae). Centro 

Escolar Universitario, Mandiola, Philippines. 

http:// www.ceu.edu.ph/research1.htm. 3 p. 

Câmara de Comércio e Indústria Portugal-Angola. 

2003. Caracterização económico social de 

Angola. http://www.cciportugal-angola.pt/ 

caract.htm. [not paged]. 

Fagundes, M.H. 2003. Sementes de juta e malva: 

algumas observações. Companhia Nacional de 

Abastecimento, Brazil. http://www.conab.gov. 

br/politica_agricola/Conjunturasemanal/Especiai

s/SEMENTES%20JUTA%MALVA.doc. 10 p. 

FAO. 2003. Food composition table for use in 

Africa. Food and Agriculture Organization of 

the United Nations, Rome. http://www.fao. 

org/docrep/003/x6877e/X687705.htm#ch5. [not 

paged]. 

Forest Research Institute of Malaysia. 2003. Plants 

information: Urena lobata Griff., pulut-pulut, 

Malvaceae. Forest Research Institute of 

Malaysia, Kuala Lumpur, Malaysia. http://www. 

frim.gov.my/tu/Urena.htm. 1 p. 

Freedman, R. 1998. Famine foods: Malvaceae. 

Perdue University, Perdue, IN. http://www.hort. 

perdue.edu/newcrop/faminefoods/ff_families/M

ALVACEAE.html. 3 p. 

Harris, P.J.C. 1986. Dormancy of Urena lobata L. 

seeds. I. Development of sulphuric acid 

scarification techniques. Ghana Journal of 

Agriculatural Science 14-19: 79-84. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Institute of Systematic Botany. 2003. Atlas of 

Florida vascular plants. University of South 

Florida, Tampa, FL. http://plantatlas.usf. 

edu/synonyms.asp?plantID=1364&genus=Urena

&species=lobata. [not paged]. 
 

787

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Natural Resources Conservation Service. 2003. 

Plant profile: Urena lobata L., Caesarweed. 

http://plants.usda.gov/cgi_bin/plant_profile.cgi?

symbol=URLO. 5 p. 

Pacific Island Ecosystems at Risk. 2003. Urena 

lobata L., Malvaceae. http://www.hear.org/ 

Pier/urlob.htm. 3 p. 

Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. 

Montiel, eds. 2001. Flora de Nicaragua. 

Monographs in Systematic Botany Vol. 85, No. 

2. Missouri Botanical Garden, St. Louis, MO. 

p. 945-1,910. 
 

788 

Urera baccifera (L.) Gaud. ortiga brava 

URTICACEAE 

Synonyms: none 

John K. Francis 

General Description.—Ortiga brava means “bad 

nettle” in Spanish. The species is known by a host 

of other common names, among which are: 

stinging nettle, ortiga, pringamoza, mala mujer, 

chichicaste, nigua, guaritoto, ishanga, manman 

guêpes, and urtiga bronca (Little and others 1974). 

Ortiga brava is a weak-stemmed woody shrub that 

sometimes becomes a small tree. Individual plants 

may have one or several stems, branching near the 

ground. Probably the most recognizable feature of 

the plant is the presence of many sharp, stinging 

spines on the leaves, branches, and stem. The 

alternate leaves are 12.5 to 25 cm in length and 7.5 

to 12.5 cm in width, coarse-toothed with thin 

blades and stout petioles. The stem bark is smooth, 

light gray with green-over-white inner bark, and 

thick greenish pith. There are raised leaf scars and 

lenticels on the stems and branches. 

Range.—Ortiga brava ranges throughout the 

Greater and Lesser Antilles, Trinidad, and Tobago 

and on the mainland from Mexico to Brazil, 

Argentina, Bolivia, and Peru (Little and others 

1974). The species is not known to have 

naturalized outside its native range. 

Ecology.—Ortiga brava grows in the higher 

rainfall portion of the moist forest (>1600 mm 

precipitation), especially along drainages, and 

throughout the wet and rain forests (2000 to 4000 

mm of precipitation). Alluvial, coluvial, and 

residual soils from acidic and calcareous rocks 

make suitable habitat. Well-drained and somewhat 

poorly drained soils of all textures are colonized. 

Being moderately shade intolerant, disturbance is 

generally required for the species to get started, 

and it disappears after the forest understory 

becomes dark. A high importance value of ortiga 

brava in a Brazilian forest and an unbalanced 

diameter distribution was used to infer past 

disturbance (Bertoni and others 1988). 

Reproduction.—Minute male and female flowers 

are borne in pink or purple, branched clusters on 

the stem on different plants (dioecious). The fruits 

are white or pinkish, spongy and watery, with one 

greenish-black seed per fruit (Little and others 

1974). A Puerto Rican collection of seeds 

averaged 4.36 + 1.17 g each (Francis and 

Rodríguez 1993) or 500,000 seeds/kg. They began 

germinating in 26 days with a final germination of 

49 percent (Francis and Rodríguez 1993). Ortiga 

brava built up to high populations in a firedisturbed forest in the Atlantic forest in southern 

Brazil through a high recruitment rate and low 

mortality (Nascimento and others 1999). The seeds 

are disbursed by birds. 

Growth and Management.—Individual stems of 

ortiga brava may grow for 5 years or more and 

reach 5 m in height. Heights of 1.5 to 2 m are more 

typical. Ortiga brava stems cut during site 

management activities quickly sprout and regain 

their former height. Whether sprouting from the 

rootstalk occurs after senescence and death of 

individual stems is not known.

Benefits and Detriments.—In Central America, 

ortiga brava has been planted in hedges and fences 

to make an almost impenetrable barrier (Little and 

others 1974). Although not generally sought after 

for food under normal conditions, ortiga brava is 

one of the first plants to fruit after hurricanes, and, 

as such, is an important factor in the survival of 

frugiverous birds (personal communication with 

Joseph Wunderle, IITF, Río Piedras, PR). The 

stems of ortiga brava were used by Aztec and 

Otomi Indians in Mexico for making paper (Hagen 

1943). The plant is considered a weed in shaded 

coffee plantations. (Francis and Rodríguiz 1993). 
 

789

All parts of the plant are covered with the spinelike hairs that cause considerable pain when they 

penetrate the skin. Normal clothing does not 

protect against the stinging effect. However, the 

pain is usually short-lived, lasting from a few 

seconds to a few minutes. Although no permanent 

injury results, some people may experience rash or 

blisters; swelling, fever, and ulcers can occur in 

extreme cases (Allen 1943). The plant has long 

been used in folk medicine. Recent laboratory tests 

using rats have shown anti-inflamatory and 

analgesic activity of aqueous extracts of ortiga 

brava (Badilla and others 1999). Amerindians in 

Costa Rica chastised themselves with branches of 

ortiga brava in a toughening ritual (Badilla and 

others 1999).

References 

Allen, P.H. 1943. Poisonous and injurious plants 

of Panama. Journal of Tropical Medicine 

23(suppl.): 3-76. 

Badilla, B., G. Mora, A.J. Lapa, and J.A. Silva 

Emim. 1999. Anti-inflammatory activity of 

Urera baccifera (Urticaceae) in Sprague-Dawley 

rats. http://www.ots.duke.edu/tropibiojnl/claris/ 

47-2/badilla.html. 

Bertoni, J.E. de A., F.R. Martins, J.L. de Moraes, 

and G.J. Shepherd. 1988. Composicão floristica 

e estrutura fitossociologica do parque de 

Vacununga, Santa Rita do Passa Quatro, Sao 

Paulo, gleba Praxides. Boletim Tecnico do 

Instituto Florestal, Sao Paulo 42: 149-170. 

Francis, J.K. and A. Rodríguez. 1993. Seeds of 

Puerto Rican trees and shrubs: second 

installment. Research Note SO-374. U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, New 

Orleans, LA. 5 p. 

von Hagen, V.W. 1943. Mexican paper-making 

plants. Journal of the New York Botanical 

Garden 44: 1-10. 

Little, E.L., Jr., R.O. Woodbury, and F.H. 

Wadsworth. 1974. Trees of Puerto Rico and the 

Virgin Islands. Vol. 2. Agriculture Handbook 

449. U.S. Department of Agriculture, 

Washington, DC. 1,024 p. 

Nascimento, H.E.M., A. da S. Dias, A.A.J. 

Tabanez, and V.M. Viana. 1999. Tree 

population and dynamics of a semidecidous 

seasonal forest fragment in the Piracicaba 

region, Sao Paulo State, Brazil. Revista 

Brasileira de Biologia 59(2): 329-342. 
 

790 

Vernonia albicaulis Pers. Santa María 

ASTERACEAE 

Synonyms: Vernonia vhliana Less. 

 Vernonia thomae Benth. 

 Cacalia thomae (Benth.) Kuntze 

 Vernonia longifolia vahliana (Less.) Urban 

 Vernonia longifolia sintenisii Urban 

 Vernonia sintenisii (Urban) Gleason 

 Lepidaploa glabra (Willd.) H. Rob. 

John K. Francis 

General Description.—Santa María, also known 

as tabac à jacot, is a shrub to 2 m tall and stem 

basal diameter to 1 cm. The ribbed, woody, glanddotted, hairy stems are upright or arching. Several 

stems may arise from the root crown. The lateral 

root system has brown bark and a moderate 

amount of fine roots. The alternate leaves are 

evenly dispersed on the branches and have 

petioles 1 to 11 mm long. The blades are oblong to 

elliptic, rounded at the apex and pointed at the 

base, and have entire to weakly sinuate margins. 

Inflorescences are cymes of heads of 10 to 23 

florets having lavender to white corollas 5 to 10 

mm long. The fruits (achenes) are 3 mm long with 

pale brown to white pappus with bristles 0.5 to 1 

mm long (Howard 1989, Liogier 1997). 

Range.—Santa María is native to Puerto Rico, the 

Virgin Islands, Anguilla, St. Martin, St. Barts, 

Antigua, Saba, St. Eustatius, St. Kitts, Monserrat, 

Guadeloupe, La Désidade, and Dominica (Howard 

1989). A second subspecies, longistylis Keeley, a 

taller and more robust plant, has been described on 

Guadeloupe (Howard 1989). Santa María is not 

known to have been planted or naturalized 

elsewhere. 

Ecology.—Santa María is moderately intolerant to 

intermediate in tolerance of shade. It grows at 

forest edges, small openings, under low basal-area 

stands, and on difficult terrain with few trees. This 

may be in secondary or remnant forests. The 

species is most common in limestone areas from 

near sea level to 600 m elevation (Liogier 1997) 

but also occurs in terrain underlain by igneous 

rocks. Mean annual precipitation varies from 

about 900 to 2400 mm. The soils are well-drained, 

usually clayey or loamy, and have pH’s from 

about 5.0 to 8.0. 

Reproduction.—Seed production is abundant. A 

sample of seeds from Puerto Rico averaged 3.9 

million per kg. They were sown on the surface of 

wet peat but failed to germinate. The seeds are 

dispersed by the wind. Seedlings are common to 

widely scattered.

Growth and Management.—Santa María has a 

moderate growth rate. Individual stems live about 

2 to 4 years, and plants may live much longer by 

resprouting from the roots. No planting or 

management experience has been published. 

Because Santa María appears to benefit from 

disturbance to existing forest, thinnings and other 

silvicultural activities may help maintain it. 

Benefits.— Santa María contributes to the 

diversity and beauty of the forest, helps protect the 

soil, and furnishes cover for wildlife. 
 

791

References 

Howard, R. A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 6. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 658 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, San Juan, PR. 

436 p. 
 

792 

Vernonia borinquensis Urban Puerto Rican ironweed 

ASTERACEAE 

Synonyms: Vernonia borinquensis var. resinosa Gleason 

 Vernonia borinquensis var. hirsuta Gleason 

John K. Francis 

General Description.—No local name is known 

for this species in Puerto Rico. The Natural 

Resources and Conservation Service (2002) has 

assigned it the name Puerto Rico ironweed. The 

plant is a vine-like scrambling shrub reaching 5 to 

10 m of lateral extension and sometimes as much 

as 3 m into the crowns of trees and shrubs. 

Younger plants often have a single stem, but older 

plants may form clumps and mats. The root system 

consists mainly of slender lateral roots and 

abundant, fleshy, fine roots. The principal 

branches are cylindrical, about 0.5 cm thick, and 

covered with gray bark. Fine branches are densely 

hairy, striated, and have a zig-zag form. Leaves, 

concentrated near the branch ends, are somewhat 

stiff, have a short petiole, and blades in an ovate to 

lanceolate form. They have a rounded base and a 

pointed tip. Leaves remain on the twigs 1 year and 

twigs remain green for an additional year. 

Inflorescences are leafy cymes of sessile 

campanulate heads, 5.5 to 8 mm long, of 13 to 22 

florets. The corolla is white or pale lavender. At 

maturity, the fruits (achenes) are 1 to 2 mm long, 

ribbed, and have a brown pappus 5 to 6.5 mm long 

(Acevedo-Rodríguez 1985, Liogier 1997). 

Range.—Puerto Rico ironweed is endemic to 

Puerto Rico and mainly found in the Cordillera 

Central (Acevedo-Rodríguez 1985). The species is 

not known to have been planted or naturalized 

elsewhere. 

Ecology.—Puerto Rico ironweed grows on welldrained soils with a wide range of textures derived 

from most parent materials. Fertility is usually 

moderate to good, and pH’s range from about 5 to 

7. Exposed subsoils are not colonized. Mean 

annual precipitation ranges from about 900 to 

2400 mm and at elevations from a few to about 

800 m above sea level. Puerto Rico ironweed has 

an intermediate tolerance to shade. It grows in 

openings but is often found under low to medium 

basal-area forest canopies. It is resistant to grazing 

and/or avoided by cattle enough to survive in 

light- to moderately-grazed semiforested range 

and brushy pastures. The species may be found in 

both remnant and secondary forests. 

Reproduction.—Puerto Rico ironweed flowers 

from November to July and fruits from February 

to September (Acevedo-Rodríguez 1985). 

Presumably, flowers are insect pollinated. In a 

sample of seeds collected near Calley, Puerto 

Rico, about 3 percent of the seeds were filled. 

Sorted by examination under a microscope, 

apparently filled seeds weighed an average of 

0.00098 g/seed or about 1 million seeds/kg. After 

being placed on moist filter paper, 76 percent 

germinated within 24 days. The seeds are winddispersed. Although seed production can be heavy, 

established seedlings are relatively uncommon. 
 

793

Disturbance is probably necessary for seedling 

establishment. The plants layer (root) whenever 

stems come in contact with the soil. 

Growth and Management.—Puerto Rico 

ironweed grows at a moderate rate. Established 

plants extend stems about 0.5 m/year. Although 

individual stems probably do not live more than 2 

to 4 years, plants may live much longer by 

sprouting and layering. No management 

experience has been published; thinning the forest 

overstory with accompanied soil disturbance is 

recommended to encourage regeneration. 

Benefits.—Puerto Rico ironweed helps protect the 

soil and furnishes cover for wildlife. It adds a 

pretty aspect to trailsides and forest edges but 

gives the understory a “closed” appearance and 

makes it somewhat more difficult to walk through. 

References 

Acevedo-Rodríguez, P. 1985. Los bejucos de 

Puerto Rico. Vol. 1. General Technical Report 

SO-58. U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station, 

New Orleans, LA. 331 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Natural Resources Conservation Service. 2002. 

Plants database. http://plants.usda.gov/cgi_ 

bin?topics.chi?earl=dl_all.html. [not paged]. 
 

794 

Vernonia proctori Urbatsch Proctor’s vernonia 

ASTERACEAE 

Synonyms: none 

John K. Francis 

General Description.—Proctor’s vernonia (a 

name assigned by the author) is an upright graygreen shrub to 1.5 m in height and 1 cm in basal 

stem diameter. Older plants may have as many as 

50 stems arising from the root crown. The plants 

have many lateral and abundant fine roots. 

Branches are greenish gray, turning brown or gray 

with age. The foliage is crowded on new twig 

growth. Alternate leaves have short petioles and 

blades that are broadly elliptic to suborbicular, 1.2 

to 3.5 cm long, and rounded at the tip. They are 

dark green above and white beneath and have a 

densely ciliate margin. The inflorescences are 

terminal, sessile clusters of usually three heads 

with 16 to 22 florets. The corollas are purple or 

blue. At maturity, the heads contain 2- to 3-mm 

long achenes with a white pappus (Liogier 1997, 

Miner-Solá 1999, U.S. Fish and Wildlife Service 

2002). 

Range.—Proctor’s vernonia is endemic to Puerto 

Rico and found only at the summit of Cerro 

Mariquita in the range of hills known as Sierra 

Bermeja in southwestern Puerto Rico. The species 

is endangered and has an estimated population of 

950 individuals in an area of a few hectares (U.S. 

Fish and Wildlife Service 2002). 

Ecology.—Proctor’s vernonia grows in exposed 

rock crevices and in shallow cherty clay loam soil 

up to 30 cm deep that has developed over 

weathered siliceous rock. Mean annual 

precipitation is about 900 mm/year with a fall wet 

season and spring dry season. Humidity is high 

and breezes are almost continuous. Elevations 

where the population grows ranges from 270 to 

300 m above sea level (U.S. Fish and Wildlife 

Service 2002). Proctor’s vernonia is intolerant of 

shade. It grows in the open or in broken stands of 

low trees, but not under closed-canopy forest. 

Bare ground and 3 to 4 weeks of almost daily 

precipitation is probably necessary for 

establishment. The endemic area was once 

severely grazed and cut over for fuelwood, but has 

been much less disturbed in recent years. 

Reproduction.—Proctor’s vernonia flowers and 

fruits during April and May (U.S. Fish and 

Wildlife Service 2002). The plant produces seed in 

abundance. A small quantity of seeds collected by 

the author averaged 0.00038 g/seed or about 2.6 

million seeds/kg. Placed on moist filter paper, 13 

percent germinated within 15 days. The seeds are 

dispersed by the wind. Seedlings are infrequent. 

Growth and Management.—Proctor’s vernonia 

appears to grow slowly as a seedling and take 

many years to attain large plant size. Individual 

stems arising from sprouts grow about 20 cm/year. 

Although individual stems last only a few years, 

the plant, renewing itself with new sprouts, 

appears to live for many years. Propagation 

techniques have not been published. Complete 

protection of the Cerro Mariquita site from 

development, grazing, and fire is critical. 

Benefits.—Proctor’s vernonia is a pretty plant and 

certainly contributes to the aesthetics of the area 

where it grows. It protects the soil and furnishes 

cover for wildlife. 
 

795

References 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Miner-Solá, E. 1999. Arboles y plantas en peligro 

de extinción en Puerto Rico. Puerto Rico 

Ecológico Vol. 3. First Book Publishing of 

Puerto Rico, San Juan, PR. 91 p. 

U.S. Fish and Wildlife Service. 2002. Species 

accounts: Vernonia proctorii. http://endangered. 

fws.gov/i/q/saqa9.html. 2 p. 
 

796 

Vernonia sericea L.C. Rich. long-shoot 

ASTERACEAE 

Synonyms: Lepidaploa phyllostachya Cass. 

 Vernonia berteriana DC. 

 Conya portoricensis Bert. ex DC. 

 Vernonia arborescens lessingiana Griseb. 

 Cacalia sericea (L.C. Rich.) Kuntze 

 Cacalia arborescens lessingiana (Griseb.) Kuntze 

John K. Francis 

General Description.—Long-shoot, also known 

as escobilla, huye-que-te-coge, tapa caminos, and 

yerba socialista, is an upright shrub usually 1 to 

1.5 m (occasionally 2 m) in height and 1 cm in 

basal stem diameter. It usually has a single main 

stem, although older and previously damaged 

plants sometimes have multiple stems branching 

from the root collar or from near the base. The 

wood of the stems is stiff and hard. The plant is 

supported by a tap and lateral root system of stiff, 

orange to tan roots. Smaller roots are tough and 

flexible. Branches and twigs are slender. The 

leaves are linear to elliptic, 2.5 to 10 cm long, with 

an entire edge, and pointed at the tip and 

somewhat rounded at the base. The inflorescences 

are elongated scorpioid cymes with leafy bracts 

and contain lavender to white, sessile flower heads 

about 6 mm long with 11 to 18 florets. They 

produce achenes 2 mm long tipped with a brown 

pappus about 4 mm long (author’s observation, 

Liogier 1997). 

Range.—Long-shoot is native to Cuba, 

Hispaniola, Puerto Rico, and the Virgin Islands 

(Liogier 1997). It is not known to have been 

planted or naturalized elsewhere. 

Ecology.—Long-shoot grows on a variety of soil 

types, mostly of medium to heavy texture with 

pH’s from 5 to 7 that are derived from 

sedimentary (including limestone), igneous, and 

metamorphic parent materials. These are well 

drained to somewhat poorly drained soils in areas 

that receive from about 850 to 2200 mm of mean 

annual precipitation at elevations from near sea 

level to about 700 m. Long-shoot is moderately 

intolerant of shade and grows in openings and 

under the canopy of low basal-area forest. It 

competes well with forbs and grasses if the 

vegetation is not too heavy. It is browsed by cattle, 

but not preferentially, so that it can survive and 

reproduce in lightly and moderately grazed 

pastures and range. Long-shoot grows on 

abandoned agricultural land, secondary forest, 

stream banks, open remnant forests, roadsides, and 

fence rows. 

Reproduction.—Long-shoot blooms seasonally, 

probably at the latter part of the rainy season 

(November through January). The abundant seeds 

produced are dispersed by wind during the dry 

season (March and April). A collection of seeds 

from Puerto Rico weighed an average of 0.000309 
 

797

g/seed or 3.2 million seeds/kg. Placed on moist 

filter paper, 69 percent germinated over the course 

of 2 months, beginning 7 days after sowing. 

Growth and Management.—Long-shoot grows 

between 0.5 and 1 m in the first year from sprouts. 

The growth rate of seedlings is undoubtedly much 

slower. Individual stems live about 3 years, but 

plants sprout and live through a second, perhaps 

several, cycles of tops. No propagation or 

management experience has been published. 

Presumably, conditions for healthy populations 

can be created by maintaining broken or low 

basal-area forest stands. 

Benefits.—Long-shoot helps protect the soil, adds 

to the aesthetic beauty of forests, and furnishes 

cover for wildlife. The species is one of the nectar 

sources for the butterfly, Eurema leuce antillarum

Hall (Torres Bauzá 1999). 

References 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 

Torres Bauzá, J.A. 1999. Ciclo de vida de Eurema 

leuce antillarum Hall (Lepidoptera: Pieridae) en 

Puerto Rico. Caribbean Journal of Science 35(3-

4): 195-200. 
 

798 

Waltheria indica L. sleepy morning 

STERCULIACEAE 

Synonyms: Waltheria americana L. 

 Waltheria elliptica Cav. 

John K. Francis 

General Description.—Sleepy morning is also 

known as velvet leaf, marsh-mallow, monkey 

bush, boater bush, leather coat, buff coat, basora 

prieta, malvavisco, hierba de soldado, guimauve, 

mauve-gris, moto-branco, hioloa, fulutafu, kafaki, 

and many other names (Haselwood and Motter 

1966, Howard 1989, Liogier 1994, Burkill 2000). 

It is a short-lived shrub or subshrub sometimes 

reaching 2 m in height and 2 cm in stem diameter. 

Sleepy morning develops a weak taproot, robust 

lateral roots, and abundant fine roots. The roots 

are brown and flexible. This shrub usually has a 

single, strong stem emerging from the ground, but 

frequently branches near the ground. Sleepy 

morning usually has an upright and somewhat 

branchy form. However, in some environments, it 

may grow in a semiprostrate habit. The young 

stems and leaves are covered with a gray, velvety 

pubescence. The alternate leaves are narrowly 

ovate or oblong with a rounded to subcordate 

base, irregularly serrate edges, and a rounded to 

acute tip. The petioles are 0.5 to 3.3 cm long and 

the blades are 2 to 12 cm long and 1 to 7 cm 

broad. Axillary inflorescences are usually dense 

glomerules that contain fragrant, yellow to orange 

flowers. Each 2-mm capsule holds one tiny, black, 

obovoid seed (Howard 1989, Liogier 1994, 

Tropilab, Inc. 2001). 

Range.—Sleepy morning now grows throughout 

the tropics and warmer subtropics (Liogier 1994). 

It apparently naturalized in Hawaii soon after the 

arrival of nonnative colonists (Haselwood and 

Motter 1966). Howard (1989) indicates that the 

species is native to the New World where it occurs 

from Florida and Texas to Brazil (Dick 2001, 

Nelson 1996, Texas A & M University 2001).

Ecology.—Sleepy morning grows in disturbed dry 

and well drained, moist habitat. In Puerto Rico, 

the species occupies areas that receive from 750 to 

about 1800 mm of annual precipitation from near 

sea level to more than 400 m of elevation. The 

species grows on sites up to 1,220 m in elevation 

in Hawaii (University of Hawaii 2001). It 

colonized a wide variety of soils in areas with 

igneous, metamorphic (including ultramafics) and 

sedimentary (including limestone) rocks. The 

species may be found on old fields, construction 

sites, roadsides, burned forest and grasslands, and 

stream overflow areas in Puerto Rico. In Florida, it 

grows in open pinelands, hammocks, and 

disturbed sites (Long and Lakela 1976). It is 

intolerant of shade and will not survive under a 

closed tree canopy and cannot compete with grass 

in dense swards. It withstands drought, salt spray, 

and mildly salty soils. 

Reproduction.—Sleepy morning plants begin 

flowering at about 6 months of age and bloom 

more or less continuously for the rest of their 

lives. A collection of seeds made in Puerto Rico 

averaged 0.0013 g/seed or 764,000 seeds/kg. 
 

799

These seeds were sown without any pretreatment 

on filter paper and after 16 weeks yielded 13 

percent germination. Reproduction is by seeds. 

The seeds are disbursed by water, agricultural 

equipment (Sánchez and Uranga 1993), and 

grazing animals. Seedlings are relatively common 

in disturbed habitat. 

Growth and Management.—Sleepy morning 

plants in Puerto Rico live for 1 or 2 and 

occasionally 3 years. Death usually occurs during 

the dry season. Perennial growth is more likely in 

continuously moist habitat. Stands of sleepy 

morning in agricultural plantations can be 

controlled by cultivation and probably by 

broadleaf herbicides. 

Benefits and Detriments.—In the Turks and 

Caicos Islands, sleepy morning is used to make an 

herb tea (Wood 2001). The plant produces a fiber 

that was formerly used for making cords, sacking, 

padding, and sandals (Guzmán 1975). Durawhite, 

an extract of sleepy morning, is used in a 

commercial cosmetic for its ability to inhibit 

melanin synthesis and whiten the skin (Janssen 

Cosmeceutical Care 2001). The plant contains 

steroid derivatives and alkaloids of the adouetine 

group that perhaps make it physiologically active. 

Various extracts are used as standard febrifugal, 

purgative, emollient, tonic, analgesic, and 

astringent herbal medicines in Africa (Burkill 

2000). In Hawaii, the root is chewed to relieve 

sore throat (Neal 1965). Stems are used as a chew 

stick, and extracts of the plant are used as an eye 

bath and a remedy for hemoptysis in Panama 

(Agricultural Research Service 2001). Seeds are 

sold commercially, and the species is cultivated in 

gardens as a medicinal plant. The plant is browsed 

by all types of livestock, especially when young 

(Burkill 2000). Sleepy morning forage in a 

Mozambique valley during the rainy season 

contained 6.4 percent crude protein, 0.12 percent 

phosphorus, and 0.51 percent calcium (Faftine and 

others 2001). Sleepy morning is considered a 

weed in much of its range, but it is seldom 

aggressive enough to be a major problem 

(Sánchez and Uranga 1993). It is host for a 

number of insects harmful to agricultural crops 

(Centro de Desarrollo de Agronegocios 2000, 

Lastres 2000, Sánchez and Uranga 1993). 

References 

Agricultural Research Service. 2001. Tico 

ethnobotanical dictionary. http://www.arsgrin.gov/duke/dictionary/tico/w.html. 4 p. 

Burkill, H.M. 2000. The useful plants of West 

Tropical Africa. Royal Botanic Gardens, Kew, 

UK. 686 p. 

Centro de Desarrollo de Agronegocios. 2000. Guia 

prictica de manejo de plagas y enfermedades del 

chile. CDA Boletín Técnico de Producción 13. 

http://www.hondurasag.org/fintrac-cda/pubs/ 

Prod_13_Esp.pdf. 4 p. 

Dick, C. 2001. Amazon plant list. Ecotour 

Expeditions, Inc. http://www.naturetours.com/ 

Am_lispl.html. 8 p. 

Faftine, O., A. Alage, and J.P. Muir. 2001. 

Characterization of forage selected by cattle on 

communal range in Manhição, Mozambique. 

http://stephenville.tamu.edu/~jmuir/faftine.htm. 

6 p. 

Guzmán, D.J. 1975. Especies útiles de la flora 

Salvadoreña. Ministerio de Educación, 

Dirección de Publiciones, San Salvador, El 

Salvador. 703 p. 

Haselwood, E.L. and G.G. Motter. 1966. 

Handbook of Hawaiian weeds. Experiment 

Station/Hawaiian Sugar Planter’s Association. 

Honolulu, HI. 479 p. 

Howard, R.A. 1989. Flora of the Lesser Antilles, 

Leeward and Windward Islands. Vol. 5. Arnold 

Arboretum, Harvard University, Jamaica Plain, 

MA. 604 p. 

Janssen Cosmeceutical Care. 2001. Supreme 

secrets. http://www.janssen-beauty.com/ 

supreme.htm. 11 p. 

Lastres, L.. 2000. Guia practica de manejo de 

plagas y enfermedades del chile. CDA Technical 

Production Bulletin 13. Centro de Desarrollo de 

Agronegocios. http://www/jpmdirasag/org/ 

fintrac-cda/pubs/Prod_13_Esp.pdf. 4 p. 

Liogier, H.A. 1994. Descriptive flora of Puerto 

Rico and adjacent Islands. Vol. 3. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

461 p. 

Long, R.W. and O. Lakela. 1976. A flora of 
 

800 

Tropical Florida. Banyon Books, Miami, FL. 

962 p. 

Neal, M.C. 1965. In gardens of Hawaii. Special 

Publication 50. Bernice P. Bishop Museum 

Press, Honolulu, HI. 924 p. 

Nelson, G. 1996. The shrubs and woody vines of 

Florida. Pineapple Press, Inc., Sarasota, FL. 

391 p. 

Sánchez, P. and H. Uranga. 1993. Plantas 

indeseables de importancia economica el los 

cultivos tropicales. Editorisal CientíficoTécnica. La Habana, Cuba. 166 p. 

Texas A & M University. 2001. Texas vascular 

plant checklist: Sterculiaceae. Texas A & M 

University, Department of Botany. 

http://www.csdl.tamu.edu/FLORA/ftc/dft/ 

ftc_str.htm. 1 p. 

Tropilab, Inc. 2001. Waltheria indica L. 

http://www.tropilab.com/sleepy_waltheria.html. 

2 p. 

University of Hawaii. 2001. Plants of Hawaii 

Volcanoes National Park. 

http://www.botany.edu/b308/bigisland/species/ 

waltheria/waltheria.htm. 3 p. 

Wood, K.M. 2001. Roadside wildflowers. 

http://www.timespub.tc/Natural%History/ 

Archive/Summer2001/wildflowers.htm. 2 p. 
 

801

Wedelia reticulata DC. manzanilla de monte 

ASTERACEAE 

Synonyms: Seruneum reticulatum (DC.) Kuntze 

 Stemmodontia reticulata (DC.) Cook & Collins 

John K. Francis 

General Description.—Manzanilla de monte is 

an evergreen shrub to 1.5 m in height and 1.5 cm 

in basal diameter. Adult plants have few-to-many, 

slender, brown, brittle, woody stems arising from 

the root crown. The stems have a 1-mm pith and 

discernible annual rings. The root system is 

dominated by stiff and brittle, slender, dark tanover-green, wide spreading laterals. Also, 

adventitious roots grow from the bases of 

branches. Branching is often bifurcate or 

trifurcate, sometimes forming a tangled structure 

in older plants. However, foliage tends to 

concentrate at the periphery of the crown. Leaves 

are opposite, narrowly oblong ovate to oblong 

lanceolate, short-pointed at the tip and rounded to 

subcordate at the base, 3 to 10.5 cm long, with a 

0.5- to 1.0-cm petiole and fine serrate edges. The 

leaves are green to yellow-green. Compound 

flowers are terminal, solitary, or occasionally in 

two’s or three’s. The yellow involucral bracts 

(“petals”) are 4 to 7 mm in length and numerous 

florets. The fruits are four-sided achenes 3 mm 

long (author’s observations, Liogier 1997). 

Range.—Manzanilla de monte is native to Puerto 

Rico and the island of Hispaniola (Liogier 1997). 

It is not known to have been planted or naturalized 

elsewhere. 

Ecology.—Manzanilla de monte grows on soils of 

all textures, pH’s from mildly acid to mildly 

alkaline, over parent materials of igneous, 

sedimentary (including limestone), and 

metamorphic (including ultramafic) rocks. Soils 

are usually rocky or shallow over bedrock. The 

species grows from near sea level to elevations of 

600 m or more in Puerto Rico (Breckon 2001, 

Francis and others 1998). Annual rainfall ranges 

from about 900 to 2000 mm. Manzanilla de monte 

tolerates salt spray and mild salinity in the soil. It 

is drought tolerant. During drought, the leaves wilt 

but usually do not fall off. Manzanilla de monte is 

intolerant of shade. It is open-grown or grows in 

small openings in low forest. The species is hardy 

and competes well with grass and herbs of similar 

height. Well-established stands become clumps 

and thickets that tend to shade out plants under 

them. 

Reproduction.—Manzanilla de monte blooms 

and fruits continually. Seed production can be 

abundant although unknown insects sometimes 

destroy most of the seeds. Other than shortdistance transport by wind, gravity, and water, 

specific means of dispersal are unknown. 

Seedlings are relatively uncommon. Suckers at the 

edge of clumps are common, and plants 

occasionally layer. 

Growth and Management.—Manzanilla de 

monte is slow growing in natural environments. 

Sprouts grow 10 to 30 cm/year. Individual stems 

persist about 4 to 10 years. Through continually 

suckering and layering, plants can last almost 

indefinitely.

Benefits.—Manzanilla de monte contributes to the 

aesthetics of natural areas where it grows, helps 
 

802 

protect the soil, and furnishes cover and possibly 

food for wildlife. Although somewhat rough in 

appearance during dry periods, it has pretty 

flowers and should make a fine ornamental for 

natural landscaping, especially in coastal areas and 

rocky terrain. 

References 

Breckon, G.J. 2001. Preliminary check list for 

Maricao Forest. http://www.uprm. 

edu/biology/profs/breckon/herbarium/FLORAM

ARICAO.htm. [not paged]. 

Francis, J.K., S. Alemañy, H.A. Liogier, and G.R. 

Proctor. 1998. The flora of Cañón de San 

Cristóbal, Puerto Rico. General Technical 

Report IITF-4. U.S. Department of Agriculture, 

Forest Service, International Institute of Tropical 

Forestry, Río Piedras, PR. 37 p. 

Liogier, H.A. 1997. Descriptive flora of Puerto 

Rico and adjacent islands. Vol. 5. Editorial de la 

Universidad de Puerto Rico, Río Piedras, PR. 

436 p. 
 

803

Ziziphus obtusifolia (Hook. ex Torr. & Gray) Gray graythorn 

RHAMNACEAE 

Synonyms: Condalia obtusifolia (Hook.) Weberbauer 

 Condalia lycioides (Gray) Weberbauer 

 Zizyphus lycioides Gray 

James E. Nellessen 

General Description.—Graythorn is also known 

as lote bush, southwestern condalia, prairie bush, 

or thornbush. This shrub reaches heights of 2 to 3 

m but is often shorter. Stems are grayish green due 

to a whitish or grayish wax-like bloom, which 

helps to distinguish Ziziphus from Condalia. The 

stems are robust, grooved, highly branched, 

although terminal branches are short, and bear 

stout spine-tips. The stem branching pattern gives 

this shrub an irregular shape. Side branches tend to 

come out at right angles from the main axis. The 

leaves are alternate, simple, generally less than 2.5 

cm long, range from 0.4 to 0.7 cm in width, are 

ovate, oblong, or elliptic with three main veins. 

Leaf shape is variable which has lead to some 

confusion in identification. Leaves are dropped 

during dry periods (Carter 1997, Correll and 

Johnston 1979, Great Plains Flora Association 

1986, Kearney and others 1951, Martin and 

Hutchins 1980-81). 

Taxonomy.—This shrub has a somewhat 

complicated proliferation of taxonomic names. 

Two varieties are recognized: Ziziphus obtusifolia

var. canescens (Gray) M.C. Johnston; and the 

typical variety Z. obtusifolia var. obtusifolia. 

Synonyms for variety canescens are: Condalia 

lycioides (Gray) Weberb. var. canescens (Gray) 

Trel.; Zizyphus lycioides Gray var. canescens

Gray; and Condaliopsis lycioides var. canescens

(Gray) Suess. Synonyms for variety obtusifolia

are: Condalia lycioides (Gray) Weberb., including 

var. lycioides; Condalia obtusifolia (Hook. ex 

Torr. & Gray) Weberb.; Zizyphus lycioides Gray; 

and Condaliopsis lycioides (Gray) Suess. (Allred 

2002, Kartesz 1994). 

Range.—This species occurs from just a few 

counties in southwestern Oklahoma (McCoy 

1981), portions of central and western Texas, 

through southern New Mexico to northwestern 

Arizona and southeastern California and south 

well into Mexico. It occurs at a wide elevational 

range from about 300 m to 1,525 m. Populations 

in the more western portions of its range are 

variety canescens (Kearney and others 1951). 

Ecology.—Graythorn will grow in a variety of 

situations from silty and sandy floodplains and 

washes to gravelly slopes and plains in desert 

grasslands and scrublands. This species tends to be 

diffusely distributed within its local habitat, but 

may also form thickets. This species is considered 

a characteristic component of Chihuahuan desert 

scrub and a diagnostic climax indicator species of 

desert grassland (Dick-Peddie 1993). In a northern 

Texas study, although graythorn had a minimal 

influence on overall grass cover, there appeared to 

be differences in species distributions around 

graythorn bushes (Foster and others 1984). 

Buffalo grass [Buchloe dactyloides (Nutt.) 

Engelm.] was more abundant near graythorn than 

in shrub free zones, while Texas winter- or 

needlegrass (Stipa leucotricha Trin. & Rupr.) 

cover and standing crop was greater away from 

graythorn. A study conducted in a savanna 

parkland habitat in southern Texas showed that 

graythorn is a deciduous shrub with an average 

leaf lifespan measured at 66 days (Nelson and 

others 2002). Supplemental watering had no 

detectable effect on leaf longevity. Root infection 

with symbiotic strains of Frankia and Rhizobium 

did not occur on four species of Rhamnaceae from 

southern Texas, including graythorn, when 

planted in pots in the laboratory, in a 1:1 native 

soil (with symbionts) to vermiculite substrate 

(Zitzer and others 1996). 
 

804 

Reproduction.—Flowers of graythorn are small, 

inconspicuous, and occur in axillary cymes of two 

to seven flowers. The five-petalled flowers are 

greenish yellow. Graythorn may bloom at various 

times during the growing season, from May 

through September, depending on the availability 

of moisture. The fruit is a dark blue to black 

globose drupe, 6 to 10 mm long, covered with a 

whitish bloom, and matures from July into early 

October. Seeds are nondormant, but the seed coats 

are impermeable to water, hence are classified as 

physically dormant (Baskin and Baskin 2001). 

Physically dormant seeds need some type of 

scarification of the seed coat to initiate water 

penetration and germination. Optimal germination 

occurs at 25 °C. Both light and prechilling 

maximized the percent germination (Speer and 

Wright 1981, Young and Young 1992). Graythorn 

seeds from the lower Rio Grande valley in 

southern Texas did not need pretreatment for 

germination (Vora 1989). 

Growth and Management.—Graythorn stem 

regeneration generally occurs just below the cut or 

removed portion. When stems are removed to the 

ground line, regeneration will occur from the root 

crown, and stems can be regenerated from roots, 

but generally only when the root crown itself is 

removed (Flinn and others 1992). Graythorn may 

demonstrate rapid growth when resprouting from 

cut stumps or during periods of good moisture but 

is generally a slow growing shrub. Although this 

species may on occasion become locally dense and 

form thickets, it is generally a species that is 

diffusely distributed within its habitat, such that 

management should not be a problem. 

Benefits.—The fruits are readily eaten by birds 

such as Gambel's quail and white-winged doves. 

Graythorn was among several shrubs in a northern 

Mexico study testing the preferred browse 

composition and digestibility of foliage. Graythorn 

was grouped with species having a high crude 

protein content, lower cell wall content, and 

smoother or waxy leaf surfaces with the absence of 

trichomes. Range goats and white-tailed deer 

included graythorn as an important part of their 

diets (Ramirez and others 1997). In another 

nutrient study in northern Mexico (Ramirez and 

others 2001) graythorn was among several shrubs 

studied for seasonal mineral levels. General 

mineral concentrations were higher in the spring 

and summer, but only calcium, magnesium, 

potassium, and iron were at levels to meet adult 

goat requirements during all seasons. Phosphorus 

was low in all seasons except the spring. Native 

Americans have used the plant for medicinal 

purposes. Pima Indians have steeped the roots in 

water and applied them to sore eyes. Seri Indians 

have powdered the roots for application to skin 

and scalp sores. Solutions made from roots also 

served as a soap substitute (Bowers and Wignall 

1993, Epple 1995, Kearney and others 1951). 

References

Allred, K.W. 2002. A Working Index of New 

Mexico Vascular Plant Names. Available on a 

New Mexico State University web page: 

http://web.nmsu.edu/~kallred/herbweb. [not 

paged]. 

Baskin, C.C. and J. M. Baskin. 2001. Seeds: 

Ecology, Biogeography, and Evolution of 

Dormancy, and Germination. Academic Press, 

San Diego, CA. 666 p. 

Bowers, J.E. and B. Wignall. 1993. Shrubs and 

Trees of the Southwest Deserts. Southwest 

Parks and Monuments Association. Tucson, 

AZ. 140 p. 

Carter, J.L. 1997. Trees and Shrubs of New 

Mexico. Johnson Books, Boulder, CO. 534 p. 

Correll, D.S. and M.C. Johnston. 1970. Manual 

of the Vascular Plants of Texas. Texas Research 

Foundation, Renner, TX. 1,881 p. 

Dick-Peddie, W.A. 1993. New Mexico 

Vegetation: Past, Present and Future. 

University of New Mexico Press, Albuquerque, 

NM. 244 p. 

Epple, A.O. 1995. A Field Guide to the Plants of 

Arizona. Falcon Publishing Inc., Helena, MT. 

347 p. 

Flinn, R.C., C.J. Scifres, and S.R. Archer. 1992. 

Variation in basal sprouting in co-occurring 

shrubs: implications for stand dynamics. 

Journal of Vegetation Science 3(1): 125-128. 

Foster, M.A., C.J. Scifres, and P.W. Jacoby, Jr. 

1984. Herbaceous vegetation - lotebush 

(Ziziphus obtusifolia var. obtusifolia) 
 

805

interactions in north Texas USA. Journal of 

Range Management 37(4): 317-320. 

Great Plains Flora Association. 1986. Flora of the 

Great Plains. University Press of Kansas, KS. 

1,392 p. 

Kartesz, J.T. 1994. A Synonymized Checklist of 

the Vascular Flora of the United States, Canada, 

and Greenland. Vol. 1, 2nd Edition. Biota of 

North America Program of the North Carolina 

Botanical Garden. Timber Press, Portland, OR. 

622 p. 

Kearney, T.H., R. Peebles, and Collaborators. 

1951 with 1960 supplement. Arizona Flora. 

University of California Press, Berkeley, CA. 

1,085 p. 

Martin, W.C. and C.E. Hutchins. 1980-1981 

(reprinted 2001). A Flora of New Mexico. Vol. 

1. Bishen Singh Mahendra Pal Singh, India and 

Koeltz Scientific Books, Germany. 1,276 p. 

McCoy, D. 1981. Roadside Trees and Shrubs of 

Oklahoma. University of Oklahoma Press, 

Norman, OK. 116 p. 

Nelson, J.A., P.W. Barnes, and S. Archer. 2002. 

Leaf demography and growth responses to 

altered resource availability in woody plants of 

contrasting leaf habit in a subtropical savanna. 

Plant Ecology 160(2): 193-205. 

Ramirez, R.G., G.F.W. Haenlein, and M.A. 

Nunez-Gonzalez. 2001. Seasonal variation of 

macro and trace mineral contents in 14 browse 

species that grow in northeastern Mexico. Small 

Ruminant Research 39(2): 153-159. 

Ramirez, R. G., J.L. Pineiro-Hernandez, and R.K. 

Maiti. 1997. Nutritional profile and leaf surface 

structure of some native shrubs consumed by 

small ruminants in semiarid regions of 

northeastern Mexico. Journal of Applied 

Animal Research 11(2): 145-156. 

Speer, E.R. and H.A. Wright. 1981. Germination 

requirements of lotebush (Ziziphus obtusifolia

var. obtusifolia). Journal of Range Management 

34: 365-368. 

Vora, R.S. 1989. Seed germination characteristics 

of selected native plants of the lower Rio 

Grande valley Texas, USA. Journal of Range 

Management 42(1): 36-40. 

Young, J.A. and C.G. Young. 1992. Seeds of 

Woody Plants in North America. Dioscorides 

Press, Portland, OR. 407 p. 

Zitzer, S.F., S.R. Archer, and T.W. Boutton. 1996. 

Spatial variability in the potential for symbiotic 

N-2 fixation by woody plants in a subtropical 

savanna ecosystem. Journal of Applied Ecology 

33(5): 1125-1136. 
 

806 

Zuckia brandegei (Gray) Welsh & Stutz ex Welsh siltbush 

CHENOPODIACEAE 

Synonyms: Grayia brandegei Gray var. brandegei

 Grayia brandegei Gray var. plummeri Stutz & Sanderson 

 Zuckia arizonica Standley 

Nancy L. Shaw, Rosemary L. Pendleton, and Emerenciana G. Hurd 

 

General Description.—Siltbush or spineless 

hopsage is a deciduous shrub or subshrub ranging 

from 0.1 to 0.8 m in height (Goodrich and Neese 

1986). Stems of the current year are thornless and 

ascending to erect, branching from a persistent, 

woody base. Leaves are alternate, gray-scurfy and 

entire to lobed. Overwintering leaf buds are 

prominent, axillary, and globose (Welsh and others 

1987). There are three varieties (Welsh 2000). 

Diploid (2x = 18) G. brandegei (Gray) Welsh & 

Stutz var. brandegei is small with narrow, linear 

leaves. Tetraploid (4x = 36) G. b. (Gray) Welsh & 

Stutz var. plummeri (Stutz & Sanderson) Dorn 

(Plummer’s siltbush) is larger with ovate to 

lanceolate leaves. Utricles of both varieties are 

obcompressed and vertical with two-winged 

samara-like bracts (see illustration). Utricles of the 

third variety, G. b. (Gray) Welsh & Stutz var. 

arizonica (Standley) Welsh (Arizona siltbush), are 

dorsiventrally compressed and mostly horizontal 

with six-keeled bracts. Flowers of all three 

varieties develop in small clusters in bract axils. 

Staminate flowers consist of four or five stamens 

and a four- or five-lobed perianth. Pistillate 

flowers contain a single pistil enveloped by two 

united bracts. Embryos are well developed. 

Range.—Siltbush is a narrowly distributed 

edaphic endemic, largely restricted to the Colorado 

River drainage. G. b. var. brandegei occurs 

primarily in south-central Utah and northeastern 

Arizona. G. b. var. plummeri grows in isolated 

populations in northeastern Utah, south-central 

Wyoming, western Colorado, and northwestern 

New Mexico. G. b. var. arizonica occurs in 

scattered populations from northern Arizona to 

northeastern Utah (Goodrich and Neese 1986, 

Shaw and others 2001, Welsh and others 1987). 

Ecology.—Siltbush grows in isolated monotypic 

populations on shale outcrops in desert shrub and 

lower juniper communities at elevations of 1,280 

to 2,240 m (Goodrich and Neese 1986). Soils on 

these outcrops are poorly developed, fine textured, 

and characterized by moderate levels of soluble 

salts, and high levels of exchangeable sodium and 

potassium. Micronutrient levels are generally high. 

Siltbush tissues accumulate sodium and 

magnesium, but potassium levels and the calcium 

and magnesium ratio are low (Pendleton and 

others 1996). 

Reproduction.—All siltbush varieties are 

monoecious and heterodichogamous (Pendleton 

and others 1988) with individual plants either 

protogynous (producing pistillate, then staminate 

flowers) or protandrous (producing staminate, then 

pistillate flowers). Flowering occurs in late spring 

or summer and fruits ripen in mid to late summer 

or fall (Pendleton and others 1988). Protogynous 

plants generally produce more seed, but 

protandrous plants may be equally productive in 

years when precipitation is high and seed 
 

807

predation is low (Pendleton and others 2000). 

Wind and gravity disperse the fruits over time; 

some fruits generally remain on the plant until late 

winter. There are about 625,000 seeds/kg 

(Pendleton and others 2000). Germination 

experiments conducted with seeds of Z. b. var. 

brandegei and Z. b. var. plummeri showed that 

seeds of warm winter populations germinated over 

a wide range of constant temperatures (15 to 30 

o

C) (Meyer and Pendleton 1990). Seeds of coldwinter populations were dormant at fall and winter 

temperatures and required exposure to overwinter 

chilling to promote germination. Germination 

generally increased with duration of wet 

prechilling at 1 o

C for up to 8 weeks, dry 

afterripening for up to 14 months, or bract removal 

(Meyer and Pendleton 1990). Germination is 

epigeal. 

Growth and Management.—Few data are 

available. Siltbush is a stress-tolerant species 

capable of establishing on steep, eroded sites 

where the combination of salinity, aridity, and fine 

soils preclude establishment of other species 

(Pendleton and others 1996). Seedling growth rate 

and palatability of mature plants are moderate 

(personal communication with Steve Monsen, 

USDA-FS-RMRS, Provo, UT, and Howard Stutz, 

Brigham Young University, Provo, UT). Plants 

may attract rodents, other small animals, and deer. 

Benefits.—Siltbush is a potential revegetation and 

soil stabilization species for disturbances resulting 

from mining and other human activities on sites 

where it is native. Few other species will grow on 

these outcrops (Pendleton and others 1996). 

 

References 

Goodrich, S. and E. Neese. 1986. Uinta Basin 

flora. U.S. Department of Agriculture, Forest 

Service, Intermountain Region, Ashley National 

Forest and USDI Bureau of Land Management, 

Vernal District, Ogden, UT. 320 p. 

Meyer, S.E. and R.L. Pendleton. 1990. Seed 

germination biology of spineless hopsage: 

Between population differences in dormancy 

and response to temperature. In: Proceedings, 

Symposium on cheatgrass invasion, shrub dieoff, and other aspects of shrub biology and 

management. U.S. Department of Agriculture, 

Forest Service, General Technical Report INT276. Intermountain Research Station, Ogden, 

UT. p. 187-292. 

Pendleton, R.L., D.C. Freeman, E.D. McArthur 

and S.C. Sanderson. 2000. Gender specialization 

in heterodichogamous Grayia brandegei.

American Journal of Botany. 87:508-516. 

Pendleton, R.L., E.D. McArthur, D.C. Freeman, 

and A.C. Blauer. 1988. Heterodichogamy in 

Grayia brandegei (Chenopodiaceae): Report 

from a new family. American Journal of Botany. 

75: 267-274. 

Pendleton, R.L., S.D. Nelson, and R.L. Rodriguez. 

1996. Do soil factors determine the distribution 

of spineless hopsage (Grayia brandegei)? In:

Proceedings, Symposium on shrubland 

ecosystem dynamics in a changing climate. 

General Technical Report INT-GTR-338. U.S. 

Department of Agriculture, Forest Service, 

Intermountain Research Station, Ogden, UT. p. 

205-209. 

Shaw, N.L., R.L. Pendleton, and E.G. Hurd. 2001. 

Zuckia brandegei (Gray) Welsh & Stutz ex

Welsh. In: F.T. Bonner and R.G. Nisley, eds. 

Woody Plant Seed Manual. U.S. Department of 

Agriculture, Forest Service, Washington, D.C. 

http://wpsm.net/index. html. 8 p. 

Welsh, S.L. 2000. Zuckia. Flora of North America. 

http://hua.huh.harvard.edu/FNA/zuckia.ed2.html 

3 p. 

Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. 

Higgins, eds. 1987. A Utah flora. Great Basin 

Naturalist Memoirs 9. Brigham Young 

University, Provo, UT. 894 p. 
808

Glossary of Technical Terms Used in this Volume

Achene—an indehiscent one-seeded fruit type common in Asteraceae. 

Alfisols—order of moderately leached soils, developed under forests, with relatively high native fertility, 

and having well developed subsurface horizons with clay accumulation. 

Allele—one of an array of genes possible at a certain position (locus) on a given chromosome. 

Allelopathy—the negative influence of plants on each other generally understood to arise from released 

metabolic products. 

Alpine—sites in mountains above the timberline or vegetation characteristic of those areas. 

Anthesis—the time at which a flower comes into full bloom. 

Arborescent—having a treelike habit. 

Aridisols—order of soils of arid regions that contain free CaCO3 and have at least some subsurface horizon 

development. 

Aril—an appendage or outer covering of some seeds, often fleshy or pulpy. 

Armed—having spines or thorns. 

Awn—bristle-like appendage of a flower and fruit usually seen in grasses. 

Axillary—borne in the axils (or inside the angles of attachment) of leaves. 

Beach strand—narrow strip of vegetation between a beach and inland formations. 

Berry—fleshy fruit containing one or usually more seeds. 

Biomass—the dry weight of the plants and animals (usually just plants in vegetation studies) in a unit area. 

Bonsai—a tree or shrub grown in miniature form through cultural practices; the term is sometimes used for 

naturally miniature trees or shrubs. 

Bract—modified leaves associated with flowers. 

Calyx—the whorl of green leaf- or bract-like sepals. 

Campanulate—bell shaped. 

Canescent—hoary, gray-colored pubescence. 

Canopy—the more or less continuous cover of tree crowns in a forest. 

Capsule—a fruit arising from a multicarpellate ovary that usually dries and splits open. 

Cartaceous—papery texture (usually used to describe leaves). 

Catkin—a drooping elongated inflorescence of unisexual flowers typical of willows. 

Cespitose shrubs—very short shrubs with a cushion-like habit. 

Chaining—dragging a heavy chain between two bulldozers to kill unwanted brush and small trees and to 

scarify the soil. 

Clambering—habit of growing over obstacles or into low vegetation without tendrils or other specialized 

climbing structures. 

Claviculate—club shaped. 

Clearcut—silvicultural harvesting method in which all the trees are removed at one time and usually 

unwanted material is felled as well. 

Climax—terminal stage of ecological succession in which the vegetation association remains stable over a 

relatively long period. 

Codominant—a crown class in a tree or shrub canopy in which a plant shares the main canopy level with 

many other individuals of similar height. 

Colluvium (colluvial soil)—fractured rock and soil accumulated at the foot of a slope or the soil developed 

from it. 

Coppice—reproduce by sprouts after being cut or damaged. 

Cordate—heart shaped. 

Coriaceous—thickened and leathery. 

Corolla—the whorl or aggregate of the petals of a flower. 

Corymb—a flat-topped inflorescence in which the outer flowers open first. 

Crenate—margin with rounded, broad teeth. 

Cyme—a more or less flat-topped inflorescence in which the central flowers open first 

d.b.h.—stem diameter at breast (1.37 m above the ground) height. 

Decumbent—branching habit that is flattened, growing along the ground with ends turned up. 

Dioecious—having staminate (male) and pistillate (female) flowers on different plants. 

Diploid—having two sets of chromosomes in a somatic nuclei. 
809

Disjunct population—a population isolated from the main part of the range. 

Diurnal—occurring (blooming) during the day. 

Dominant—a crown class in forest or shrub canopy in which the individual is taller than the general 

canopy level; may also refer to species with a preponderance of numbers or biomass. 

Drupe—fleshy, one-seeded fruit whose seed is encased in a stony endocarp. 

Ecotone—a zone of transition between adjacent plant communities. 

Ecotype—a population within a species that is genetically adapted to a certain habitat distinct from that 

which is general for the species. 

Edaphic—pertaining to the soil and its ecological relationships. 

Endosperm—starchy or oily tissue in many seeds. 

Entire—without teeth or indentations on the margin. 

Epigeal—germination and emergence in which the cotyledons raise above the soil surface. 

Escapes—introduced populations now established and competing in the wild and spreading beyond the 

original point of introduction. 

Excessively drained soil—water runs off, infiltrates, or evaporates rapidly so that the soil tends to be 

droughty even with ample rainfall. 

Floret—individual, usually minute, flower of a compound flower. 

Fruticose—being woody or shrub-like. 

Galleries—small patches of vegetation (usually trees or shrubs) in protected locations distinct from 

surrounding vegetation. 

Germinative capacity—the percentage of seeds that germinate during the normal period of germination. 

Glabrous—not hairy; hairless or nearly so. 

Glaucous—with a waxy bloom or whitish substance that rubs off. 

Habit—the growth form or appearance of a plant. 

Head—a compact, multiple inflorescence, often disk-shaped. 

Hammock—a fertile area in Florida and Southeastern United States that is elevated above the surrounding 

terrain. 

Hypanthium—a floral tube formed by the fusion of the basal portions of the sepals, petals, and stamens 

from which the rest of the floral parts arise. 

Hypogeal—germination and emergence in which the cotyledons remain below the soil surface. 

Inceptisols—order of soils that are young, frequently rocky, and have a poorly developed profile. 

Intermediate—a crown class of tree or shrub canopies in which the individual plant crown is within and 

does not quite rise to the level of the top of the canopy. 

Lanate—soft, woolly pubescence. 

Layering—rooting at a point on a stem where the stem comes in contact with the soil or with artificial 

treatment called air layering. 

Legume—one to multiple-seeded fruit (pod) common in the Fabaceae family. 

Lenticels—lenticular or diamond-shaped dots, pits, or corky protrusions on stems. 

Litter—organic debris on the surface of the soil or the uppermost layer of the forest floor organic horizon. 

Maceration—the process of soaking and gently grinding fruits to remove the pulp from seeds. 

Mesic—moist soil or environment; neither excessively wet nor dry. 

Mine spoil—dirt and rock left after a mining operation. 

Monoecious—having staminate (male) and pistillate (female) flowers on the same plant. 

Naturalized—naturally reproduced progeny of horticultural plants or wildland plantings outside their 

native range. 

Nerve—vein, usually used in relation to leaves. 

Nut—a one-seeded fruit enclosed in an involucre (diminuative=nutlet). 

Old growth forest—characterized by old, usually large, and often decadent trees. 

Open forest—a forest characterized by low basal area, a large amount of space between crowns, and a lot 

of sunlight reaching the forest floor. 

Open-grown—growing without significant competition from plants of similar size or taller. 

Panicle—an inflorescence with indeterminate branching, the primary axis bearing branched secondary axes 

and pedicellate flowers. 

Pappus—hairs attached to small seeds that together function as a wing. 

Perfect flower—a flower with both stamens (male parts) and carpels (female parts). 

Pericarp—the wall of a ripened ovary (fruit). 
810

pH—a measure of soil acidity or alkalinity; values above 7.0 are alkaline and values below are acid. 

Phenotype—a plant as it exists; the product of the interaction of genetics and environment. 

Pioneer—shrub or other plant that establishes on or has the ability to colonize newly disturbed habitat. 

Pith—spongy central cylinder of parenchyma tissue in stems. 

Ploidy—the degree of repetition of the basic chromosome number. 

Poorly drained soil—water runs off or evaporates so slowly that the soil remains wet for a large part of the 

year. 

Provenance—the original geographic source of a plant’s genetic material. 

Puberulent—covered with soft, very fine hairs. 

Raceme—an inflorescence with indeterminate single axis and pedicellate flowers. 

Rachis—the axis of a pinnately compound leaf. 

Remnant (forest)—lightly to moderately disturbed pieces or stands of the original forest. 

Reticulate—with a net-like structure. 

Rhizome—modified underground stem. 

Riparian vegetation—plant associations and stand types growing along streams. 

Samara—a one-seeded, winged fruit. 

Savannah—tropical or subtropical grassland with scattered trees; usually maintained by periodic fires. 

Scabrous—covered with scattered, short, stiff hairs. 

Scandent—vine-like habit. 

Scarify—to perforate or make a seed coat permeable by mechanical or chemical means. 

Scrub—vegetation consisting of shrubs and sometimes trees stunted because of poor site conditions. 

Secondary forest—forest that has grown up after clearing or lesser disturbance. 

Selfing—pollination of a plant with its own pollen. 

Seral—transitional as a site or vegetation association moves toward a climax state. 

Serrulate—finely serrate. 

Sessile—connected directly to the stem or other structure without a stalk. 

Shade intolerant—requiring full or nearly full sunlight to survive, grow and reproduce. 

Shade tolerant—the ability to survive, grow, and reproduce in the shade of taller vegetation, as under a 

tree canopy. 

Skeletal soil—substrate in which stones form the structure of the soil with sand, silt, and clay filling the 

voids between. 

Slash—the cut stems and limbs of trees and brush, or to cut unwanted trees and brush leaving it in place. 

Soil texture—classes defining the proportion of sand, silt, and clay making up a soil. 

Spike—indeterminate inflorescence of sessile flowers arranged linearly along a central axis. 

Stamen—male or pollen-producing part of the flower. 

Stipule—a small leaf-like structure found at the base of some leaf petioles, usually present in pairs. 

Stolon—a stem or shoot that bends to the ground and takes root after contact. 

Stratify—breaking the physiological dormancy of certain seeds with a period of cold or warm incubation. 

Succulent—class of plants with fleshy leaves and usually stems that store water in their tissues and usually 

have special metabolic pathways (crassulan acid metabolism). 

Sucker—root sprouts or to reproduce by root sprouts; sometimes used to mean an epichormic branch. 

Suffruticose—perennial plant with stems that are woody near their base. 

Suppress—inhibit the growth (and often eventually kill) by shading and competition for water and 

nutrients. 

Synonym—an equivalent but superseded species name. 

Testa—the outer coat of a seed. 

Ultisols—a family of highly weathered soils found in subtropical and tropical environments. 

Ultramafic rocks—metamorphic rocks (including serpentine) characterized by low percentages of silica 

and elevated percentages of iron and magnesium. 

Variety—a subdivision of a species that has one or more inherited morphological characteristics. 

Well-drained soil—water runs off, infiltrates, or evaporates readily but not rapidly. 

Xeric garden—a garden planted with drought-resistant vegetation that is not irrigated. 
811

References Consulted 

Burns, R.M. and B.H. Honkala, eds. 1990. Silvics of North America. Vol. 2. Agriculture Handbook 654. 

U.S. Department of Agriculture, Forest Service, Washington, DC. 877 p. 

Editorial staff. 1984. Webster’s ninth new collegiate dictionary. Marriam-Webster Inc., Publishers, 

Springfield, MA. 1,563 p. 

Lawrence, G.H. 1955. An introduction to plant taxonomy. The Macmillan Company, New York. 179 p. 

Soil Survey Staff. 1951. Soil survey manual. Agriculture Handbook 18. Agriculture Research 

Administration, U.S. Department of Agriculture, Washington, DC. 503 p. 
812

Common Name Index 

Common name page 

‘a’ali’i 293 

‘a’ali’i ku makani 293 

‘a’ali’i-ku ma kua 293 

abeyelo 235 

abrojo Colorado 412 

abuta 212 

abutilon espinoso 540 

acacia bush 21 

acacia courant 289 

acacia savane 289 

acacia zarza 19 

aceituna Americana 117 

aceituno 154 

acequia willow 664 

achiote 112 

afata 714 

aguacero 127 

aguedita 561 

ague tree 681 

aguinaldo blanco 783 

agüiste 489 

‘aheahea 190 

aile à ravet 764 

aji 147 

aji de gallina 144 

aji picante 147 

akalakala 658 

akeake 293 

akulikuli-kai 107 

alcaparro 696 

alcotá 212 

alcotán 567 

alderleaf mountain-mahogany 178 

alfilerillo 459 

algodón de seda 137 

algorobo 580 

alkali sagebrush 66 

allamanda 36 

aloes vert 350 

alpine dryad 295 

alpine juniper 401 

amansa 684 

American angelica tree 40 

American beautyberry 135 

American bittersweet 164 

American burning bush 333 

American ivy 532 

American plum 586 

American snowball 737 

American yew 758 

amor ardiente 393 

amourette 19 

Common name page

amourette-grand-bois 487 

amourette-rivière 489 

amouretto 489 

angelica tree 40 

angel’s trumpet 36, 286 

angel’s whisper 299 

anguila 327 

añil 386 

añil de pasto 386 

añil falso 252 

anisette 567 

anisillo 564, 567 

annatto 112 

antelope bitterbrush 610 

antelope brush 610 

antelope buckbrush 610 

antibala 714 

Antilles velvetshrub 570 

Apache-plume 337 

aperta ruão 564 

applebush 361 

aramina 785 

arabo carbonero 323 

arazá-puitá 597 

árbol de navidad 352, 617 

arbre de soie 136 

arctic willow 660 

arête-boeuf 529 

argythamnia 45 

Arizona queen-of-the-night 181 

Armenian blackberry 652 

Aroeira pimenteira 686 

aroma 14 

aromo 21 

arrowleaf sida 714 

arrowwood 379 

ash barberry 464 

Asian bittersweet 161 

ati popa’a 43 

atiu 43 

auguey 475 

‘aweoweo 190 

azahar 508 

azota-caballo 299 

azúcares 395 

baby tea 196 

badoh 783 

Bahaman wild coffee 608 

balsam 473, 619 

balsaminha 142 

bálsamo 372 

basbasot 714 
813

Common Name Index 

bamboo piper 564 

banglin 489 

baquiña 432 

barbasco 395 

basam 275 

bashful plant 489 

basket willow 664 

basket wiss 779 

basket wood 708 

basora 247 

basora prieta 798 

bastard guava 329 

bastard sirio 30 

bâtard balengène 726 

bayberry 502 

bay-cedar 742 

bay grape 229 

bayonette 125 

beach creeper 321 

beach plum 591 

beautyberry 135 

bébèl 575 

bejuco de berac 192 

bejuco de canastas 708 

bejuco de corrales 708 

bejuco de costilla 538, 708 

bejuco de fuego 359 

bejuco de gato 456 

bejuco de guajanilla 538 

bejuco de indio 359 

bejuco de membrillo 717 

bejuco de mona 213 

bejuco de nasa 779 

bejuco de palma 779 

bejuco de paloma 538, 779 

bejuco de paralejo 764 

bejuco de sarsaparilla 717 

bejuco de sopla 359, 689, 764 

belly-ache bush 399, 619 

bemberecua 769 

berenjena cimmarona 726 

berenjena de gallina 726 

berenjena de playa 724 

berenjena silvestre 726 

Bermuda mulberry 135 

Bigelow sagebrush 50 

big sagebrush 85 

bija 112 

bilimbi 505 

birchleaf cercocarpus 178 

birchleaf mountain-mahogany 178 

bird pepper 144, 147 

biscuitwood 149 

bitter bush 196, 561, 619 

bitter cane 370 

bitter cherry 589 

bitter nightshade 721 

bittersweet nightshade 721 

bizcocho 395 

black ambret 487 

blackbead 575 

blackbead tree 235 

blackberry 43, 652 

black bush 781 

black hawthorn 254 

black mimosa 489 

black sage 247, 676 

black sagebrush 72 

black sumac 625 

black torch 319 

black widow 473 

black willow 668 

blackwith 708 

blind-eye-tree 374 

boater bush 798 

bog myrtle 511 

bois-anisette 432 

bois bolibri 354 

bois cabrit 32, 524, 696 

bois cabroit 32 

bois candelle 38, 319 

bois canon 137 

bois capable 684 

bois caraibe 761 

bois-champignon 473 

bois codine 115 

bois crabbe 575 

bois de fer 32, 235 

bois de l’encore 524 

bois énivrant 219 

bois flambeau 38, 186, 319 

bois fleche 341 

bois flot 376 

bois-foufou 354, 524 

bois-guillaume 365 

bois jaune 149 

bois mêche 140 

bois montagne 561 

bois-noir 140 

bois petites feuilles 327 

bois poison 561 

bois-puant 140 

bois pini 38 

bois puante 699 

bois tan 154 

bola de coral 393 

bomba 137 

bonbon bleu 223 
814

Common Name Index 

bon bon mél 223 

bonbon rouge 259 

boo 726 

bord-de-mer 154 

borrerie verticillée 732 

botón blanco 732 

botoncillo 240 

bow-pigeon 225 

box-brier 617 

boxleaf stopper 327 

box thorn 453 

Brazilian pepper tree 686 

Brazilian jasmine 397 

bread-and-cheese 575 

brilla 107 

brisselet 325 

britónica afelpada 473 

britónica prieta 471 

brittlebush 302 

brittle thatch palm 766 

broom 275 

broombush 365 

broom butterweed 694 

broom dalea 600 

broom groundsel 694 

broom indigobush 600 

broom palm 766 

broom snakeweed 367 

broomstick 777 

broom tops 275 

broomweed 367, 473, 714 

broomwood 473, 777 

brownweed 367 

bucho 699 

buckbrush 158, 280 

budsage 82 

buenas tardes 286 

buenda 192 

buffalo current 637 

buffalo-tip 766 

buff coat 798 

buis de sapin 758 

bun 508 

bur mallow 785 

burning bush 335 

burning love 393 

bursting heart 333 

burro 140 

burro weed 388 

burweed 781 

bush cinquefoil 280 

bush monkey flower 495 

bush penstemon 545 

busunuvo 372 

buttonball 170 

buttonbush 170 

button mangrove 240 

button sage 417 

button willow 170 

buttonwood 240 

bwa kabuit 32 

cabai nache 617 

cabo de hacha 777 

cabra cimarrona 684 

cabrilla 775 

cabrito 363 

cachimbo 219, 524, 604 

cachimbo peludo 647 

cadillo 243, 785 

cadillo espinoso 540 

cadillo de perro 781 

Caesar weed 785 

café 232 

cafecillo 121, 151, 341 

café cimarrón 341 

café bâtard 606 

café de la India 508 

cafeillo 684 

café marron 606, 608 

café silvestre 151 

caïmite marron 201 

caimitillo 201 

caimitillo cimarrón 201 

caimitillo de perro 201 

cají torcido 21 

calico bush 407 

California brickellbush 123 

California broom 445 

California buckwheat 314 

California rosebay 621 

California sagebrush 52 

calotrope 137 

camasey 223 

camasey almendro 469 

camasey blanco 477 

camasey colorado 219, 475 

camasey de costilla 475 

camasey de felpa 480 

camasey de paloma 219 

camasey peludo 221, 223 

camasey racimoso 481 

cambrón 14 

cambronera 453 

cammy cuphia 266 

camphire 107 

caña brava 370 

Canada yew 758 

cañafistola cimarrona 696 
815

Common Name Index 

canalete 777 

canario 36 

candleberry 502 

candlewood 38, 186, 319 

cane 91, 555 

cangá 449 

canique grise 129 

canote-macaque 223 

capaillo 32 

caper 140 

capim elefante 542 

capulí cimarrón 775 

caracolillo 149 

carbonera 696 

cardero 753 

cardinal’s guard 516 

cardio de frade 732 

cariaquillo 196 

Caribbean spiritweed 32 

carrapata 777 

carrapateira 643 

carrapicho do mata 785 

carricillo 518 

carruzo 219 

cascabelillo axiliar 261 

cascalata 235 

casie 14 

casse marron 696 

Castle Valley clover saltbush 101 

Castle Valley saltbush 101 

castor bean 643 

castor-oil plant 643 

catclaw 16 

catclaw acacia 16 

catclaw mimosa 482, 489 

cat’s claw 456, 575 

cattle tongue 577 

cautiva 36 

Cayenne pepper 147 

cedar 404 

ceibey 535 

celadonia 115 

cenicillo 691 

cenizoso 477 

cerezo 149 

cerise de sable 591 

chaico blanco 286 

chamizo 93 

chaparral broom 103 

chaparro 268 

chapeau g’leau 432 

cheezefruit 505 

chew-stick 359 

chichicaste 788 

chichita 686 

chico 425, 453 

chile perro 696 

chili pepper 147 

China box 508 

Chinese honeysuckle 442 

Chinese inkberry 184 

chocolate blanco 115 

chokecherry 594 

cholla 520 

Christmas berry 686 

Christmas bush 196 

Christmas cherry 329 

Christmas vine 783 

Christmas wreath 783 

cigüillo 363 

cina-cina 529 

cinnamon wood 681 

ciruela 586 

citron 214 

citronnier 186 

clavelón de playa 119 

clidemia 223 

climbing mimosa 487 

climbing nightshade 721 

climbing orange root 164 

climbing poison ivy 769 

clove current 637 

coast broom 753 

coast rhododendron 621 

cocaine falsa 323 

cockspur 156 

coco-plum 199 

coco-ravet 149 

cocuisa 350 

coffee 232 

coffee colubrine 235 

coffee senna 699 

coin vine 278 

collier-diable 575 

colorín 140 

comb butterweed 691 

common broom 275 

common buttonbush 170 

common juniper 401 

common poison ivy 769 

common rabbitbrush 203 

common reed 555 

common sage 417 

common sassafras 681 

common sida 715 

common snowberry 744 

common sumac 628 

conchombre 286 
816

Common Name Index 

conejo colorado 777 

Congo pea 132 

contite 417 

cooze mahot 785 

copal 686 

coquelicot 540 

coral 259 

corazón de paloma 235 

corcho blanco 149 

cordobán 475 

cordobancillo 645 

cordobancillo peludo 647 

cordoncillo 564, 567 

corita 34 

cornicopio 286 

corona de novia 783 

correosa 631 

cortaduro 518 

corta-lengua 151 

coscorrón 154 

cossie 21 

costilla de vaca 93 

cotorrerillo 149 

cotton-france 137 

cottonrat 374 

cough bush 321 

cousin-petit 781, 785 

cousin rouge 785 

cowbush 374 

cowfoot leaf 432 

cowheel bush 432 

cowitch 374 

cow’s foot 564 

cow-tobacco 545 

coyote bush 103 

coyote willow 664 

crab’s eye 12 --- 

crabwood 225 

crack-open 151 

creambush oceanspray 379 

creek senecio 691 

creeping barberry 464 

creeping mahonia 464 

creeping Oregon grape 464 

creosote bush 419 

criaquillo 414 

crises marine 742 

croc-chien 487, 617 

crow broom 753 

cruz de Malta 393 

cuabilla 38, 742 

Cuba jute 714 

cubra prieta 319 

cucharo 742 

cucubano liso 363 

cudweed sagewort 69 

cuentas de oro 299 

cuernecillo 374 

cuero de sabana crucillo 363 

cuneate saltbush 101 

cuphia 266 

curarador 115 

curarina 212 

cure-for-all 577 

curl-leaf mountain mahogany 175 

curumo 140 

dabel-da-dama 36 

dadangsi 785 

dama de día 184 

dama de noche 127, 186 

damson plum 201 

David bush 286 

David’s root 192 

day cestrum 184 

day jasmine 184 

deerbush 158, 178, 610 

deer horn cactus 181 

deerweed 445 

desert cassia 702 

desert cedar 404 

desert Christmas cactus 520 

desert mock orange 551 

desert seepwood 739 

desert spoon 283 

desert sumac 631 

desert sweet 188 

desmanthus 289 

desmanto 289 

devil’s claw 573 

devil’s fig 726 

devil’s trumpet 286 

devil’s walking stick 40 

dhal 132 

ditay paye 142 

doctor-bar 561 

doctor long 575 

Doña Julia 372 

dorme dorme 492 

dormidera 492 

dormilona 489 

dos caras 475 

Douglas groundsel 691 

Douglas hackberry 167 

Douglas hawthorn 254 

Douglas maple 24 

Douglas rabbitbrush 209 

Douglas ragwort 691 

Douglas thorntree 254 
817

Common Name Index 

dulcamara 721 

duranta 299 

duro-duro 149 

Dutch myrtle 508 

dwarf American cherry 591 

dwarf juniper 401 

dwarf koa 289 

dwarf mountain-mahogany 173 

dwarf Oregon grape 464 

dwarf sumac 625 

early sagebrush 66 

eastern poison ivy 769 

eastern swamp privet 343 

eastern wahoo 335 

elephant grass 542 

elfin herb 266 

emajagua 376 

endurece maiz 140 

enegro 401 

English broom 275 

English myrtle 511 

equisósea 393 

éribun 733 

ernodia de playa 321 

erubia 719 

escambrón 573 

escoba blanca 714 

escoba dulce 714 

escoba dura 714 

escoba negra 275 

escobilla 796 

escobón 327 

espinillo 529 

espino blanco 14 

espinosa amarilla 105 

estafiante 69 

Everglade velvetseed 363 

false bittersweet 164 

false cocaine 323 

false coffee 341 

false heather 266 

false kava 564 

false mallow 467 

fausse-cane à sucre 542 

fedegoso 699 

fellenwort 721 

felty groundsel 691 

female karata 350 

fern bush 188 

fever nut 128 

fever twig 164 

few-flowered ninebark 558 

fèy a kèy 432 

feyukke friudem 505 

figus nut 399 

filigrana 414 

firebush 372 

firespike 516 

fire willow 668 

fishing rod 617 

fivefinger 280 

five fingers 617 

five-leaved ivy 532 

flameleaf sumac 625 

flame of the woods 393 

flat sagebrush 50 

fleur-corail 372 

fleur de ben-aimé 19 

fleuret-Noël 196 

fleur-sorleil bord de mar 119 

flor de San José 696 

Florida bitterbush 561 

Florida boxwood 684 

Florida crossopetalum 259 

Florida holly 686 

Florida privet 345 

Florida swamp privet 345 

flowering current 637 

fortuga caspi 149 

fourwing saltbush 93 

fragrant current 637 

framboisier 658 

French cotton 137 

French mulberry 135 

fresa de montaña 658 

friega-platos 726 

frijol de árbol 132 

fringed sage 63 

fringed sagebush 63 

fringed sagewort 63 

fruta de paloma 299, 684 

frutilla 417 

fulutufa 798 

furrusca 699 

galán arbóreo 30 

galán de día 184 

galán de monte 186 

gallinero 30 

gandul 132 

gandure 132 

garbancillo 777 

garden datura 286 

garricillo 604 

gato 374 

gato soga 374 

gatura 482 

genevrier commun 401 

geranium-leaf eupatorium 194 
818

Common Name Index 

giant cabuya 350 

giant cane 91 

giant laurel 623 

giant milkweed 136 

giant mimosa 489 

giant reed 555 

giant reedgrass 555 

giant sensitive plant 489 

gilia penstemon 545 

ginger-thomas 761 

giroflé 449 

girofle-ma 449 

glandular mesquite 580 

globe-flower 170 

goat-bush 156 

goat horn 617 

goat-weed 142 

gobernadora 419 

gobiabiera 597 

Gold Coast jasmine 397 

golden creeper 321 

golden current 637 

golden dewdrop 299 

golden hardback 280 

golden trumpet 36 

goose plum 586 

gordolobo 115 

goyavier 598 

goyavier bâtard 329 

goyavier de montague 329 

granadilla 535 

granadillo bobo 477 

grand cousin 785 

grande chelidoine 115 

grand giroflé 449 

grão de galo 299 

grass-leaved ragwort 694 

grate-jambe 487 

great morinda 505 

gray ephedra 307 

grayia 361 

gray nicker bean 129 

Gray’s saltbush 361 

graythorn 803 

gray willow 664 

green aloe 350 

green broon 275 

green ephedra 310 

greenheart 235 

green rabbitbrush 209 

Green’s mountain-ash 729 

griffe á chatte 456, 575 

grosse-peau 154 

ground cedar 401 

ground hemlock 758 

groven eye 575 

guacamayo 115 

guacimilla de canario 553 

guacoco 140 

guairaje 327 

guaita 777 

guaje 437 

guamis 419 

guano de sierra 766 

guano pea 132 

guanxuma 714 

guapo 684 

guarema 561 

guaritoto 788 

guassatonga 151 

guatama indigo 386 

guava 597 

guayaba 597 

guayaba silvestre 329 

guayabilla 151 

guayabillo prieto 363 

guayabo 597 

guayarote 154 

guayayo 564, 567 

guazumilla 121 

guen léglise 12

guerit-tout 577 

guerrero 257 

guía mansa 149 

guimauve 798 

Guinea pepper 147 

guitará 742 

gull feed 119 

gumbo file 681 

gurgeon stopper 327 

gypsum scalebroom 434 

hardhack 379 

hava de San Antonio 129 

Havard oak 613 

Havard shin oak 613 

hearts-a-bursting 333 

heather 266 

hedionda 699 

hediondilla 419, 702 

herbe-à-crâbes 107 

herbe à elephants 542 

Hercules’ club 40 

hibiscus bur 785 

hierba de peso 212 

hierba de soldado 798 

higuereta cimarrona 399 

higüerillo 643 

higüerito 643 
819

Common Name Index 

higuillo 564, 569 

higuillo de hoja menuda 564 

higuillo de limo 567 

Himalayan blackberry 652 

Hioloa 798 

hoary sagebrush 57 

hog apple 505 

hog plum 586 

hoja de pasmo 475 

hoja menuda 121, 327 

hold back 129 

hollyleaved barberry 461 

hollow heart 30 

honey bush 508 

honey mesquite 580 

honteuse 492 

hoop vine 779 

hop bush 293 

hopseed bush 293 

horn-of-plenty 286 

horse bean 529 

horsebrush 361, 365 

huang hau mu 714 

hueso 561 

huesito 323 

hueso 342 

huevo de gato 374 

huisache 14 

humble plant 492 

hummingbird bush 370 

huye-que-te-coge 796 

icaco 199 

icaque ponne 199 

idem 714 

‘ilima 712 

Indian apple 286 

Indian arrowwood 548 

Indian mulberry 505 

Indian Valley false mallow 467 

indigo 386 

indio 323 

inkberry 617 

ink-bush 184, 345 

inkweed 739 

iodine bush 739 

ipil-ipil 437 

Irish tops 275 

ishanga 788 

ivy 407 

ivybush 407 

ixora 393 

jabillo 777 

jaboncillo 689 

jacatirão 477 

jackass breadnut 219 

jackswitch 243 

jalapón 527 

Jamaica loostrife 449 

Jamaican trash 365 

Japanese honeysuckle 442 

jasmin à bouguet 397 

jasmin blanc 397 

jasmín de canario 397 

jasmín de estrella 341 

jasmín del monte 127 

jasmín de trapo 397 

jaune d’oeuf 149 

jasmine 397, 684 

jasmín oloroso 397 

jayajabico 319 

Jerusalem-thorn 529 

jeucon 374 

jigá 325, 684 

jointfir 310 

jointwood 564, 567 

jojobán 777 

jovero 742 

jumbie apple 149 

jumbie bean 12

jumbie breatfruit 505 

jungle flame 393 

jungle-geranium 393 

jut africain 785 

kafaki 798 

kak mé 223 

kalmia 497 

kamole 449 

karatas 125 

Key lime 214 

Key thatch palm 766 

koa haole 437 

ko’oko’olau 110 

Koster’s curse 223 

Labrador tea 429 

Lakana 414 

Lamark trema 775 

lambedora 518 

lampuaye 293 

lantana 414 

la péchita 580 

laurel 407 

leadtree 437 

leandra 427 

leather coat 798 

lechecillo 647 

lemonade sumac 628, 634 

leucaena 437 

Lewis mock orange 548 
820

Common Name Index 

Lewis syringe 548 

liana fragante 291 

liane á barriques 779 

liane à lait 36 

liane amère 212 

liane á terre 779 

liane cordé 212 

liane des sorciers 192 

liane molle 213 

liane-savon 359 

liane-séche 321 

licorice plant 12 --- 

lija 245 

lila 299 

lilikoi 535 

lima 215 

limestone snakevine 384 

limón agria 214 

limón boba 214 

limoncaspi 149 

limón criollo 214 

limoncillo 323, 684 

limoneria 508 

limonejo 259 

linguam 140 

linmangkon 535 

lintentwa 518 

lintisco 178 

lipstick plant 112 

lirio 735 

little leaf cercocarpus 173 

littleleaf mock orange 551 

littleleaf mountain-mahogany 173 

little-leaf sumac 631 

little rabbitbrush 209 

llorasangre 115 

lluvia 299 

long-leaved willow 664 

long-shoot 796 

lote bush 803 

Lousiana sagewort 69 

Louisiana wormwood 69 

low juniper 401 

low rabbitbrush 209 

low sagebrush 45, 66 

lyenn-savon 359 

macanabo 201 

macata 437 

machacomo 149 

madam naiz 529 

maguey 34, 350 

maguey criollo 350 

mahot-cousin 781, 785 

mahot-cousin rouge 781 

mahot fin 247 

mahot jaune 540 

maidenberry 259 

majagua 376 

majagüilla de costa 374 

mahoe daux 376 

mala mujer 788 

malimbé 567 

mallet 243 

mallow-leaved ninebark 558 

mallow ninebark 558 

malva 785 

malva colorada 471 

malva de cochino 714 

malva loca 467 

malva silvestre 467 

malvavisco 243, 798 

mameyuelo 43 

mangle bobo 117 

mangle médaille 178 

mangle negro 240 

mangle prieto 240 

manglillo 319 

mangue 240 

manman guêpes 788 

manto 259 

many-headed groundsel 694 

many-seed 449 

manzanilla de monte 801 

maracuyá 535 

maraimaray 278 

maravedí 259, 689 

marble tree 154 

marie-honte 492 

marieniera 30 

marker grass 542 

marsh-mallow 798 

mascapalo 359 

mataburro 529 

mata de mariposa 354 

mata gallina 30 

mata pelandok 43 

mata piojo 777 

matchbrush 367 

mate prieto 154 

matico 564 

mato de playa 129 

mat saltbush 99 

matscale 99 

Mauritius hemp 350 

Mauritius raspberry 658 

Mautofu 785 

mauve 471 

mauve d’Amérique 467 
821

Common Name Index 

mauve-gris 798 

maya 125, 475 

mayhont 492 

meadow fern 511 

médicinier bâtard 399 

médicinier noir 399 

médicinier rouge 399 

medsinnyé benni 219 

membrillo 259 

memizo de majagua 775 

mengkudu 505 

merisier 684 

mesquite 580 

Mexican lime 214 

miki palaoa 699 

milk tree 619 

mille fleurs 414 

miraguano 766 

mitten tree 681 

mock orange 508, 548, 737 

molinillo 777 

mombin bâtard 777 

mondururu preto 477 

monkey bush 798 

monkey’s hand 432 

monval 437 

morinda 505 

morito 508 

morivivi 492 

mormon tea 307, 310 

moth penstemon 545 

moto-branco 798 

mountain avens 295 

mountain balm 157 

mountain cercocarpus 178 

mountain grape 461 

mountain juniper 401 

mountain laurel 407 

mountain mahogany 755 

mountain maple 27 

mountain ninebark 558 

mountain sorrel 655 

mountain spray 379 

mountain sumac 625 

mountain willow 668 

mouse’s pineapple 505 

mullaca colorado 477 

murette 779 

napier grass 542 

naranjo jasmine 508 

naranjuelo 140 

narrowleaf willow 664 

nedyah 781 

negra loca 227 

netleaf hackberry 167 

Nevada ephedra 307 

Nevada jointfir 307 

Nia 514 

night-blooming cereus 181 

nigua 223, 788 

nigüto 30 

ninebark 280, 558 

no-eye pea 132 

noni 505 

nono 505 

Nuttall willow 668 

oakleaf sumac 634 

ocotillo 347 

ojos de cangrejo 12 

old man 691 

oldman sage 60 

oldman sagebrush 60 

oliba 117 

olivier bard 117 

olivo 140 

olivo macho 140 

ololiuqui 783 

orange jasmine 508 

orange peel 217 

orange peel clematis 217 

oregano 417 

Oregon grape 461, 464 

Oregon grape-holly 461 

Oregon holly-grape 461 

oreja de mula 475 

oreja de rató 212 

orelia 36 

Oriental bittersweet 161 

Oriental virginsbower 217 

orme petites feuilles 775 

ortiga 788 

ortiga brava 788 

oseille bord de mer 742 

oxeye daisy 119 

paaloe pretae 140 

pachaca 140 

Pacific rhododendron 621 

Pacific yew 755 

paddy’s lucerne 714 

pain-in-back 775 

pain killer 505 

pale lyceum 453 

palétuvier 240 

pale wolfberry 453 

palma Christi 643 

palma de cogollo 766 

palma de escoba 766 

palma de petate 766 
822

Common Name Index 

palmetto 704 

palmita 766 

palo amarillo 115 

palo blanco 149 

palo corra 742 

palo de anastasio 777 

palo de burro 696 

palo de cachimbo 602 

palo de chivo 696 

palo de cotorra 617 

palo de hierro 323 

palo de paloma 259 

palo de paz 561 

palo de peje 561 

palo de perico 247 

palo de pollo 278 

palo de rayo 529 

palo de toro 115, 341 

palo de vaca 121 

palo de zorrillo 696 

palo moro 608 

palo pelado 354 

palo verde 529 

palo zapo 140 

pan cimarrón 115 

pandereta 766 

panilla 115 

pantsil 742 

papagayo 710 

papelite 151 

parcha 535 

pareira 212 

Parry rabbitbrush 206 

parrot apple 319 

parrot weed 115 

passionfruit 535 

pata de pájaro 372 

paz y justicia 456 

peetsch-kitam 617 

pega palo 291 

pelúa 219 

penah heriah 43 

peronías 12 --- 

petit bois blanc 684 

petit casse 696 

petit coco 617 

petit garcon 684 

petit mahot 247 

phlox penstemon 545 

picanier jaune 105 

pickleweed 107 

picklewood 363 

pick tree 40 

pico de pájaro 699 

pico de paloma 259, 327 

pigeonberry 299 

pigeon pea 132 

pigeon tree 40 

pigra 489 

pimiento de Brazil 686 

pink Chinese bur 785 

pink plains penstemon 545 

pionia colorada 417 

pitillo de monte 425 

placa chiquitu 259 

planta de sal 107 

plume poppy 115 

pimiento 144 

pingouin 125 

pinguin 125 

pinipinche de sabana 154 

pipewood 149 

poaia 732 

pockhout 227 

poirier 323 

pois à mabou 140 

pois cajan 132 

poison bittersweet 721 

poison cherry 259 

poison ivy 769, 772 

poison oak 772 

poison tree 154 

poison vine 769 

polyodon 411 

pomme de singe 505 

pomme épineuse 286 

pomme-macaque 505 

pompon blank 289 

pompons jaunes 21 

ponasi 524 

pondberry 440 

popian 278 

porcupine flower 105 

pork-fat-apple 199 

Pottawattami plum 586 

prairie bush 803 

prairie mesquite 580 

prairie penstemon 545 

prairie sagewort 63 

prairie-weed 280 

prickle bush 617 

prickly ash 40 

prickly elder 40 

prickly mampoo 573 

prickly sage 414 

prickly solanum 726 

pride of Big Pine 735 

primrose willow 449 
823

Common Name Index 

pringamoza 788 

Proctor’s vernonia 794 

prostrate bundleflower 289 

prostrate juniper 401 

Puerto Rican ironweed 792 

Puerto Rico girdlepod 500 

pull-back-and-hold 573 

punta de sarvia 475 

punteral 363 

purga del fraile 399 

purple datura 286 

purple sage 600 

pygmy sagebrush 75 

pyramid bush 473 

Queensland raspberry 658 

Quelite salado 739 

quiebrahacha 329 

quina del país 561 

quininebrush 610 

raba stokki 140 

rabbit thorn 453 

rabo de ratón 354 

raboe 617 

raichie 473 

raisin de la mar 229 

raisin marron 527 

rajana 393 

rascagarganta 527 

raspalengua 121 

ratón 325 

rattlebox 263 

rat-wood 325 

red gram 132 

red ixora 393 

redosier dogwood 249 

red palicourea 524 

red pepper 144, 147 

red physic nut 399 

red plum 586 

red sage 414 

red sumac 628 

redwood 323 

reina-de-la-noche 181 

remaroogu 608 

resin-weed 367 

retama 156, 529 

retama de escobas 275 

retama falsa 412 

retama prieta 702 

retamo 777 

rhacoma 259 

rhomboid ilima 714 

rhubarbe caraïbe 505 

ricin commún 643 

riverbank willow 664 

riverbush 170 

river hawthorn 254 

river plum 586 

roble agalla 561 

roble amarillo 761 

roble bobo 747 

roble de guayo 121 

roble cimarrón 747 

roble colorado 747 

rocío 325 

rockspirea 379, 382 

Rocky Mountain maple 24 

Rocky Mountain sumac 628 

rolón 575 

romarin noir 742 

rope mangrove 376 

rosarin-bord-de-mer 735 

roseau 555 

roseau cane 555 

rosebay rhododendron 623 

roseleaf raspberry 658 

rosemary 735 

roseta 459 

rothrock sagebrush 78 

roucou 112 

rough-leaf tree 278 

round-leaved bittersweet 161 

rubber rabbitbrush 203 

rufiana 184 

Ryberg’s poison ivy 772 

sabina 404 

sabina morena 404 

sacagarganta 527 

Sacramento bur 781 

saffron-tree 201 

sageweed 69 

saico Amarillo 761 

salmonberry 655 

saloop 681 

saltbrush 361 

saltcedar 749 

saltwood 514 

saltwort 107 

salvadilla 142 

salvia 577 

salzbush 374 

sandbar willow 664 

sand cherry 591 

sand penstemon 545 

sandsage 60 

sand sagebrush 60 

sand wash groundsel 691 

Santa María 196, 247, 790 
824

Common Name Index 

Santa María negra 564 

santo selele bélétére 477 

sapo prieto 140 

saragüero 247 

saraguero 247 

sardine 477 

sarnilla 151 

sassafras 619, 681 

satinleaf 201 

satinwood 508 

saúco 115 

saúco tintóreo 184 

sauge 414 

saw palmetto 705 

scabland sagebrush 77 

scarlet berry 721 

scarletbush 372 

scarlet sumac 628 

Schouler’s willow 668 

Scotch broom 275 

screwtree 374 

sea amyris 38 

sea daisy 119 

seagrape 229 

sea hibiscus 376 

sea mulberry 240 

sea ox-eye 119 

sea sage 417 

sea-shore ardisia 43 

seaside grape 229 

seaside mahoe 376 

seaside tansy 119 

Seminole balsamo 608 

Senna-tree 696 

sensitive plant 492 

sereno 365 

shadscale 96 

shallow ninebark 558 

shambu 112 

shining sumac 625 

shinnery oak 613 

shin oak 614 

shiny-leaf ceanothus 158 

shiny-leaf wild coffee 608 

shiny sumac 625 

shoebutton ardisia 43 

shoo-shoo bush 726 

shot bush 40 

showy crotalaria 263 

showy rattlebox 263 

shrubby cinquefoil 280 

shrubby false buttonwood 732 

Siam weed 196 

Siberian juniper 401 

silky sesban 710 

siltbush 806 

silver sagebrush 57 

silver sea oxeye 119 

siyou 30 

skunkbush 634 

skyflower 299 

sleeping grass 492 

sleepy morning 798 

slender willow 664 

smallcain 425 

small crown flower 137 

small-fruited thatch palm 766 

smelling stick 682 

smooth sumac 628 

snakebark 237 

snowbank 735 

snowberry 744 

snowbush 158 

snowbush ceanothus 158 

soap bush 223 

Sodom apple 137 

soldier’s-bush 365 

Sonadora 518 

sonzonte 235 

soot-soot 567 

souge rouge 577 

sourbush 135, 577 

southern spicebush 440 

southwestern condalia 803 

sow-berry 135 

Spanish elder 569 

Spanish mulberry 135 

Spanish stopper 327 

spiked malvastrum 467 

spiked pepper 564 

spineless hopsage 806 

spiny abutilon 540 

spiny hopsage 361 

spiny sage 361 

spoonleaf 283 

spoon plant 283 

spoon tree 154 

spoonwood 407 

springsage 82 

squaw current 641 

squawweed 691 

staff-tree 164 

sticky laurel 158 

sticky monkeyflower 495 

stiff sagebrush 77 

stinging nettle 788 

stinking weed 699 

stool 283 
825

Common Name Index 

stopper bush 327 

storax 737 

strawberry bush 333 

strumpfia 735 

sulfato 529 

supple jack 538 

sureau gros 32 

swamp privet 343 

swamp redwood 323 

sweet acacia 14 

sweet-briar 21 

sweet gale 511 

sweet fern 237 

sweet potatoe cactus 181 

sweetscent 577 

sweetspire 391 

sweet yellowcrown 584 

switchcane 91 

syringa 548 

tabac à jacot 790 

tabac djab 30, 577 

tabaco del monte 30 

tabacón 726 

tabaquillo 115 

taiche 140 

tall mahonia 461 

tallowbush 178 

tallow shrub 502 

tamarisk 749 

tantan 437 

tapacamino 524, 796 

tapaculo 374 

tapaculop 149 

tapehuiste 485 

tarro de chivo 319 

tarweed 266 

tassel-plant 742 

tassel-white 391 

tautuba 399 

tea 38, 319 

tea bush 473 

tearblanket 16 

te bois-lait 619 

te café 386 

té del pais 142 

te kafé mawon 608 

temporana 742 

tendra à caulilou-rivière 575 

terciopelo 480 

terete 417 

terongan 726 

teta de burro 201 

tête à nègre 781 

Texas mimosa 16 

Texas night-blooming cereus 181 

thé du pays 142 

thicket creeper 532 

thicket wild coffee 606 

thimbleberry 655 

thin-leafed erythroxylon 323 

thornapple 286 

thornbush 803 

thorny sensitive plant 489 

threadleaf butterweed 691 

threadleaf groundsel 691 

threetip sagebrush 88 

tibisi 425 

ti bonbon 259 

ti café marron 608 

ti coco 617 

ti mawi 492 

timberline sagebrush 79 

tintero 617 

tintillo 617 

tobacco bush 158 

togovao 293 

tomatillo 453, 726 

torchwood 395 

torciopelo 219 

toothache bush 40 

toothbrush tree 359 

torch 186 

torchwood 38 

tostado 149 

touch-me-not 492 

tree oxeye 119 

trios côtes 475 

trombeto 115 

trompete du jugement 286 

true mountain-mahogany 178 

trumpet flower 761 

túatúa 399 

tuchima 219 

turkey berry 726 

turma de toro 219 

turpentine-weed 367 

turtleweed 107 

twisted acacia 21 

tzazahuistli 520 

Uganda grass 542 

uña de gato 16, 456, 484, 575 

upland cotton 356 

uri 129 

urtiga branca 788 

urucú 112 

urupagüita 156 

Utah juniper 404 

uva de playa 229 
826

Common Name Index 

uva grass 370 

uverillo 227 

uvero 229 

uvillo 227 

vaillant garcon 561 

vara prieta 140 

varnish-leaf ceanothus 158 

vassourinha 732 

vega blanca 127 

velvet leaf 212, 798 

velvetseed 363 

Virginia creeper 532 

Virginia sweetspire 391 

Virginia tea 391 

Virginia willow 391 

viuda 291 

wait-a-bit 129, 482 

wait-a-while 16 

waraia 477 

waxberry 502, 744 

wax current 641 

waxwork 164 

western hackberry 167 

western mountain-ash 729 

western mugwort 69 

western poison ivy 772 

western sumac 628 

western syringa 548 

western thimbleberry 655 

western thimble raspberry 655 

western thorn apple 254 

western yew 755 

West Indian lime 214 

West Indian snow-berry 192 

West Indian trema 775 

Wheeler stool 283 

whisky current 631 

white alling 117 

white coalberry 744 

white indigo berry 617 

white leadtree 437 

white mountain avens 295 

white sage 69, 417, 671 

white sagebrush 57 

white sassafras 681 

white sumac 628 

white watting 327 

white willow 140 

white wood 225 

widdy 280 

wild bamboo 425 

wild bougainvillea 573 

wild cane 370 

wild cassava 399 

wild cherry 149 

wild chili 144 

wild clove 449 

wild-coffee 151, 235, 561, 608 

wild cotton 356 

wild egg plant 726 

wild grape 225 

wild guava 329 

wild honey-tree 149 

wild indigo 386 

wild jasmine 184 

wild nutmeg 154 

wild olive 117, 345 

wild pepper 144 

wild plum 586 

wild pomegranate 321 

wild poponax 21 

wild raspberry 658 

wild salve 374 

wild sage 417 

wild star-apple 201 

wild tamarind 437 

wild tantan 289 

wild tea 142 

wild tobacco 30, 577 

winged burning bush 331 

winged spindletree 331 

winged sumac 625 

witch’s broom 275 

woodbine 442, 532 

wood balsam 608 

Wood’s rose 649 

woody nightshade 721 

wooly corchorus 243 

yagrumito 115 

yaray 765 

yaría de costa 325 

yambush 365 

yellow bell 36 

yellowbrush 209 

yellow bush lupine 451 

yellow candlewood 696 

yellow cane 555 

yellow cedar 524 

yellow-elder 761 

yellow palicourea 524 

yellow plum 586 

yellow-root 164, 505 

yellow rose 280 

yellow sage 414 

yellow-top 367 

yellow weed 367 

yellowwood 684 

yerba cana 691 
827

Common Name Index 

yerba de clavo 449 

yerba de vibora 367 

yerba elephante 542 

yerba pelada 354 

yerba socialista 796 

zamorette 726 

zarca 487 

zarcilla 437 

zarsa brava 19 

zarzaparilla 717 

zicate 199 

zombie 286 

zówèy mouten 577 

z’yeux à chatte 129 
828

Authors of Thamnic Descriptions

G.W. Andrews, USDA Forest Service, Forestry Sciences Laboratory, 520 Devall Drive, Auburn, AL 

36830 

Eric L. Berlow, Research Scientist, University of California San Diego, White Mt. Research Station, 3000 

E. Line St., Bishop, CA 93514 

Don C. Bragg, Research Forester, USDA Forest Service, Southern Research Station, P.O. Box 3516 UAM, 

Monticello, AR 71656 

Mary Carrington, Assistant Professor, Science Division, Governors State University, 1 University 

Parkway, University Park, IL 60466 

Barton D. Clinton, Research Ecologist, USDA Forest Service, Southern Research Station, Asheville, NC 

28802 

Kristina Connor, Plant Physiologist, USDA Forest Service, Center for Bottomland Hardwoods Research, Box 

9681, Mississippi State, MS 39762 

Ann M. DeBolt, Botanist, USD Forest Service, Rocky Mountain Research Station, Boise, ID 83702 

Margaret S. Devall, Research Ecologist, USDA Forest Service, Center for Bottomland Hardwoods 

Research, Stoneville, MS 38776 

John K. Francis, formerly Research Forester, U.S. Department of Agriculture, Forest Service, International 

Institute of Tropical Forestry, Jardín Botánico Sur, 1201 Calle Ceiba, San Juan PR 00926-1119, in 

cooperation with the University of Puerto Rico, Río Piedras, PR 00936-4984, now retired and volunteering 

with the USDA Forest Service, Shrub Sciences Laboratory, 735 N 500 E, Provo, UT 84606 

Marshall R. Haferkamp, Rangeland Scientist, Fort Keogh Livestock and Range Research Laboratory, 

USDA Agricultural Research Service, Miles City, MT 59301 

Emerenciana G. Hurd, Botanist, Rocky Mountain Research Station, U.S. Department of Agriculture, 

Forest Service, Boise, ID 83702 

Deborah K. Kennard, Research Ecologist, USDA Forest Service, Forestry Sciences Laboratory, 320 

Breen Street, Athens, GA 30602 

Stanley G. Kitchen, Research Botanist, USDA Forest Service, Rocky Mountain Research Station, Shrub 

Sciences Laboratory, 735 N. 500 E. Provo, UT 84606 

Catherine E. Koehler, Reserve Manager, University of California Davis-McLaughlin Reserve, 26775 

Morgan Valley Rd., Lower Lake, CA 95457-9411 

Juanita A. R. Ladyman, Botanist, JnJ Associates, 6760 S. Kit Carson Cir. E., Centennial, CO 80122 

Ariel E. Lugo, Director, International Institute of Tropical Forestry, U.S. Department of Agriculture, 

Forest Service, Ceiba 1201, Jardín Botanico Sur, Río Piedras, PR 00926 

Jinshuang Ma, Research Taxonomist, Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, 

NY 11225 

E. Durant McArthur, Research Geneticist, USDA Forest Service, Rocky Mountain Research Station, Shrub 

Sciences Laboratory, Provo, UT 84606 
829

Paul A. McMillan, Coordinator, Biology Department, Capilano College, 2055 Purcell Way, North Vancouver, 

British Columbia, Canada V7J 3H5 

W. Henry McNab, Research Forester USDA Forest Service, Southern Research Station, Asheville, NC 

28802 

Ernesto Medina, Investigador Titular, Centro de Ecología, Instituto de Investigaciones Científicas, 

Caracas, Venezuela and Adjunct Scientist, International Institute of Tropical Forestry, USDA Forest 

Service, Ceiba 1201, Jardín Botanico Sur, Río Piedras, PR 00926 

Stephen B. Monsen, retired and volunteering with the USDA Forest Service, Shrub Sciences Laboratory, 

735 N 500 E, Provo, UT 84606 

Arlee M. Montalvo, Plant Population Biologist, Department of Botany and Plant Sciences, University of 

California, Riverside, CA 92521-0124 

Gerry Moore, Research Taxonomist, Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, NY 

11225 

James E. Nellessen, Botanist, Biologist and Environmental Scientist, Taschek Environmental Consulting, 

8901 Adams St., NE, Albuquerque, NM 87113 

Kenneth Outcalt, Research Ecologist, USDA Forest Service, 320 Green Street, Athens, GA 30602 

Paula M. Pijut, Plant Physiologist, USDA Forest Service, North Central Research Station, Hardwood Tree 

Improvement and Regeneration Center, 195 Marsteller Street, West Lafayette, IN 47907 

John A. Parrotta, Biological Scientist, Research & Development, USDA Forest Service, Washington, DC 

20090-6090 

Rosemary L. Pendleton, Research Ecologist, USDA Forest Service, Rocky Mountain Research Station, 

Albuquerque, NM 87102 

Sally A. Reynolds, Botanist, U.S. Fish and Wildlife Service, San Francisco Bay National Wildlife Refuge 

Complex, P.O. Box 524, Newark, CA 94560 

Christopher Ross, Natural Resource Specialist, Bureau of Land Management, Nevada State Office, P.O. Box 

12000, Reno, NV 89520

Stewart C. Sanderson, Research Geneticists, USD Forest Service, Rocky Mountain Research Station, 

Shrub Sciences Laboratory, 735 N. 500 E., Provo, UT 84606 

Nathan M. Schiff, Research Entomologist, USDA Forest Service, Center for Bottomland Hardwoods 

Research, Stoneville, MS 38776 

Randy S. Senock, Assistant Professor of Tropical Forestry, College of Agriculture, Forestry and Natural 

Resource Management, University of Hawaii at Hilo, 200 West Kawili St., Hilo, HI 96720 

Nancy L. Shaw, Research Botanist, USDA Forest Service, Rocky Mountain Research Station, Boise, ID 

83702 

Amanda J. Stevens, student, College of Agriculture, Forestry and Natural Resource Management, 

University of Hawaii at Hilo, 200 West Kawili St., Hilo, HI 96720 
830

Jeffrey R. Taylor, formerly Biological Technician, USDA Forest Service, Rocky Mountain Research 

Station, Shrub Sciences Laboratory, 735 N. 500 E., Provo, UT 84606, now student, University of Utah, Salt 

Lake City, UT 

Sarah A. Taylor Student, College of Agriculture, Forestry and Natural Resource Management, University 

of Hawaii at Hilo, 200 West Kawili St., Hilo, HI 96720 

Bruce L. Welch, Plant Physiologist, USDA Forest Service, Rocky Mountain Research Station, Shrub 

Sciences Laboratory, 735 N. 500 E., Provo, Utah 84606 



Federal Recycling Program Printed on Recycled Paper

The U.S. Department of Agriculture (USDA) prohibits discrimination in all its programs and activities on the

basis of race, color, national origin, sex, religion, age, disability, political beliefs, sexual orientation, or marital

or family status. (Not all prohibited bases apply to all programs.) Persons with disabilities who require

alternative means for communication of program information (Braille, large print, audiotape, etc.) should

contact USDA’s TARGET Center at (202) 720-2600 (voice and TDD).

To file a complaint of discrimination, write USDA, Director, Office of Civil Rights, Room 326-W, Whitten

Building, 1400 Independence Avenue, SW, Washington, DC 20250-9410 or call (202) 720-5964 (voice or

TDD). USDA is an equal opportunity provider and employer.