AGRICULTURAL RESEARCH SERVICE • U. S. DEPARqFMENT.OF AGRICULTURE
THE AGAVE FAMILY

in

SONORA

Agriculture Handbook No. 399

U. S. DEPT. OF AGRICULTURE

NATIONAL AGRICULTURAL LIBRARY

MAY 1 1972

MTALOGINfi PREß

Agricultural Research Service

UNITED STATES DEPARTMENT OF AGRICULTURE
CONTENTS

Page

Environmental setting 1

Animal associates of agave 6

Agave 9

Yucca 12

Historical notes on authors and collectors of Agavaceae in Sonora 14

Some characters of agave 17

Concept of species 23

Agave flower measurements and their ideographs 28

Flower measurements 28

Floral ideographs 36

Annotation of species 40

Key to genera of agavaceae in Sonora 40

Agave L. 41

Key to agave species in Sonora 42

Subgenus Littaea 42

Subgenus Agave 43

Subgenus Littaea 46

Subgenus Agave 80

Manfreda Salisb. 150

Polianthes L. 153

Yucca L 154

Hesperaloe Engelm. 169

Dasylirion Zuce. 175

Nolina Michx. 177

Literature cited 184

Glossary of special terms 187

Index 191

Washington, D.C. Issued March 1972

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THE AGAVE FAMILY

IN SONORA

By HOWARD SCOTT GENTRY, research botanist, Plant Science Research

Division, Agricultural Research Service

ENVIRONMENTAL SETTING

In Sonora, agaves are scattered like gems in an arborescent

matrix. They grow mainly upon the rocky slopes of hills and

mountains and are generally lacking in the valleys and on the

plains. Hence, the distributional pattern is islandlike. Compared

with the massive populations of agaves in Baja California and

in the Coahuilan Desert, they are very sparse in Sonora. However,

they are distinctly characteristic of the succulent component in the

vegetation of our American deserts and arid regions (i2).'^ Sonora

is predominantly a country of open shrub and small tree growth

with intervening grasslands. The arborescent growth varies from

the widely spaced, drought-adapted shrubs of the northwestern

desert to the closely spaced, tropical short-tree forest of the southeastern barrancas and mountains. The whole constitutes a highly

varied community of plants, in which agaves are fortuitously

deployed.

Agaves occur from sea level to the tops of the higher mountains

at elevations of 7,000 to 8,000 feet. They thrive on limestone but

they appear to do as well on igneous rocks and other well-drained,

nonalkalinic land. One species. Agave aktites, is endemic to the

coastal sand dunes. Desert species exist with about 5 inches or

less annual precipitation and can endure rainless years ; montane

species receive 30 or more inches of annual precipitation. Nearly

all Sonoran agaves are subject to frosts. Very light frosts occur

rarely along the southern coast, while freezes are severe in the

northeastern mountains, where A, palmeri and A. huachucensis

may be covered with snow.

1 Italic numbers in parentheses refer to Literature Cited, page 184.
ù AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Agaves are heliophytic and xerophytic. They pioneer on hot rock

surfaces and the geologically transient rubble of talus and outwash

slopes. The crevices of rocks and sheltering thorn shrubs help

protect them during the delicate seedling stage w^hen life hangs

precariously betv^een recurrent droughts and the w^andering feet

and teeth of animals. In many desert localities, the survival of

agave populations appears dependent upon the nursemaid vegetation, much as Turner and associates found with saguaros (5i).

Seedling survival is enhanced during those unusually favorable

rainy periods that occur by decades rather than by yearly seasons.

Shreve commented on this v^ith other desert plants (^i). Once

established, the young agave has an inherent hardiness for the

rigors of local climate. Its developing armature of teeth and spines

and the bitter leaf juices repel animals.

The generative period for sexual progeny is long, 8 to 25 years,

and of uncertain issue. Genetic potential and environment appear

to have evolved the suckering plantlets, *'hijos,'' for species survival in arid lands. If the monocarpic parent does not leave progeny with its one burst of flowers and seeds, then the vegetative

sprouts it produces about the old dry base will have, in another

climatic cycle, another chance to leave its variable offspring along

the survival line. The immortal genes carry on. Sexual generations in such cases are two, three, or more times longer than the

monocarpic habit indicates and exceed the sexual generations of

many îrees. Because of the infrequency of gene assortment and

recombination, evolution in such lines may be very slow. Some

agave clusters, as in fairy rings, are perhaps hundreds of years

old and still v/ithout apparent seeded progeny. I have never seen

a wild seedling agave less than 1 year old although I have encountered 2- or 3-year-olds sporadically. This seedling scarcity is

not due to lack of seed because most wild agaves produce abundant

seed upon maturity. One spike of Agave chrysoglossa yielded about

750,000 seeds, as estimated by counting the capsules per inch of

the fruiting shaft. If the population near Bacanora is only just

maintaining itself, this indicates that only one seed in a half million to a million germinates and grows to maturity.

Many agaves are now known to have extensive ranges and populations, whereas others are relatively rare. Most of the Sonoran

species are among the latter. A, pelona and A. zebra are limited

to the rocky heights or slopes of a few northwestern mountains

within Sonora. A. parviflora and A. shrevei matapensis are known

only as a few dozen or a few hundred individuals. This apparent

rarity may be due to lack of exploration because their ecology indicates they may exist on many of the unbotanized mountains of
THE AGAVE FAMILY IN SONORA Ö

northeastern Sonora. A. subsimplex is presently rare, having been

greatly reduced by the Seri Indians who have eaten it. Because

of its palatability to the Warihio Indians who prefer it as a sweet

food to all the other agaves in their habitat, A. jaiboli has become

increasingly rarer during the 30 years I have known it. Man and

his animals, which are fond of the young flowering shoots, have

probably greatly reduced the native wild populations. See Castetter. Bell, and Grove on Indian uses of the Agaves (8),

The widely scattered populations of agave in Sonora consist

frequently of small colonies. A. pacifica in the coastal thorn forest

of the south is an exception. It extends over many miles and numbers thousands of individuals. Plants are not conspicuous unless

they are flowering because shrubs generally hide them from view.

A. pacifica grows on sandy soils and gravelly terraces in rather

regularly and widely spaced suckering clumps. This species is also

widely dispersed on rocky slopes through the short-tree forest

and is the most abundant agave of the Mexican west coast.

A. vilmoriniana^ A. pelona, A. ocahui, and A. multifilifera are

virtually limited to cliffs, where they are out of reach of most

larger animals. These cliffdwellers also are generally unarmed,

whereas most agaves in open accessible sites have some protection

furnished by spines and teeth. Some species contain bitter principles in addition to high sapogenin content that make them unpalatable to many animals and man. The escape habitat, the armature, and toxins all appear to be adaptations of some survival value

in the evolution of this genus.

Although the long sexual generation cycle in agave may have

slowed the evolution of species, their inherent predilection for

rocky sites has resulted in fragmented distributions. Evolution of

the colonies to species could have been fostered by their isolation.

As other studies have indicated (19, Si, 36, i5), mutations and

reassortment of genes in isolation have evolved new genotypes,

which in time become genetically distinct from former contemporaries, lose gametic compatibility, and evolve eventually as

species.

Mountains in Sonora have existed for a long stretch of geologic

time. The granitic pediments are precretaceous in age. Some of

the limestones are Pennsylvanian and Cretaceous (59), Volcanic

eruptives, which indicate faulting and differential land movem.ents,

are known from the Triassic, Miocene, and Pleistocene. Block

faulting and uplift, along with volcanism, created disjunct mountains out of uniform plains. The physiographic evolution involved

the Gulf of California (39), Several of the coastal mountains appear to have been islands in the Gulf for much of the upper
AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Tertiary, until they were grounded to the mainland during the

Pleistocene by the buildup of the coastal plain, as attrital materials

were carried coastward from the weathering Sierra Madre system.

Sierra Colorai, Sierra Seri, and the Guaymas monadnoch are examples of these coastal mountains. I have called such land bodies

postinsular (ü, pp. 85-88).

The disjunct pediments that appear to have been included in

the Gulf of California during much of the Upper Tertiary are

hachured on the map (fig. 1). Some of the more inland pediments

FIGURE 1.—Map of the state of Sonora with the principal plant localities referred to in this handbook.
THE AGAVE FAMILY IN SONORA 0

like Sierra Bojihuacame and the Bacatetes, if not islandic, most

probably were maritime headlands. Sierra Bojihuacame appears

but little degraded and rather more recent than the others. However, the caverns along its base appear to be wave cut, as in an

eustadial period, and marine shells occur on a Pleistocene surface

below Recent fills. In drawing the Miocene sea line, I have followed

Schuchert (39) in principle but not in detail, placing the Miocene

Sonoran sea line farther inland. This seems desirable to account

for the very large coastal plain buildup during the Pleistocene

and the generally emerging coastline. In any case, this shoreline

should not be considered as definitive but only to draw attention

to the geobotanical evolution indicated.

The yuccas, nolinas, and other genera of the Agavaceae do not

express as much geobotanical evolution in their distributions as

the agaves. Yucca arizonica, Hesperaloe nocturna, and Nolina

bigelovii are truly endemic ; but they are limited to physiographic

regions, not to certain isolated mountains. These are regional, not

insular, endemic plants. Among continental plants, the palms are

among the few groups that rival the agaves in their response to

physiographic fractionation. Oceanic islands show many insular

endemics among many genera and if we review Trelease's, "Agave

in the West Indies" (50), we find all of his species endemic to one

or a few adjacent islands. Even though we exercised a broader

species concept than Trelease, the Caribbean agaves would still

show a high degree of insular endemism. The genus appears to

have maintained a high rate of speciation under the aegis of

isolation.

The evolution of land forms was part of the environmental

change operating with climate and associated biota on the genetic

versatility of agave to create the rich assortment of agave forms

existing in Sonora today. Unfortunately, agaves are unknown as

fossils. Although their nondeciduous leaves are not suitable for

fossil preservation, their seeds or capsules could conceivably be

preserved in compressions, but their topographic situations on

rocky uplands provide small chance for internment in sedimentary

basins.

To what extent can land differentiation be correlated with

speciation? There are many more mountains to climb, more morphology to know, and genetic systems to trace before we can answer such questions. To understand agave relationships, to understand them as species, we must read them and their environments

back in time. Then perhaps we can mature a causal philosophy.
b AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

ANIMAL ASSOCIATES OF AGAVE

Many animals get food, drink, and shelter from the agaves.

Some animals are simply takers but others, including the pollinators, appear to pay for food with biological service. The follov^ing

notes are based upon my observations during visits to agave populations. While these notes reflect the cursorial nature of my visits,

they indicate that agaves are generally beneficial to v^ildlife and

in places v^here they are abundant, their presence may be critical

to the continuance of some animal populations. That copious flov^er

nectar should be a splendid refreshment for animals in the hot,

arid deserts. Man is the animal most injurious to the agaves. He

not only chops them up and eats them and encourages his domestic

animals to do likewise, but also hacks them needlessly and bulldozes them out of his way. His constructive uses of them are discussed in the section, ''Uses of Agave and Yucca," page 9.

Bighorn sheep and deer eat the young flowering shoots of

agaves. In addition to containing sugars, starches, and other

nutrients, these shoots contain abundant water and must be of

some importance to these animals, particularly when the flowering shoots appear during dry seasons when desert water tanks are

empty and fresh ephemeral vegetation is not available. The leaves

are rarely eaten. However, mummified stomachs of extinct ground

sloths showed that they fed on leaves of the Agavaceae {21).

Pack rats of the genus Neotoma frequently live among the

thickets of agave. These rodents make their characteristic piles

of sticks, thorns, and litter among the protective agave thorns.

Their runways may be chewed through the leaf bases and these

juicy organs probably supply them with food and moisture. Animal chewings on the fruits and flowers above Neotoma dwellings

are signs attributable to them also.

Ground squirrels of the genus Citellus have been observed eating and carrying off the flowers and green fruits of agaves. These

rodents forage out hundreds of feet from their burrows and they

may disperse seeds in this activity.

The desert spermophiles and the kangaroo rats of the genus

Dipodomys have runways around agave plants. Although it is not

certainly known if these rodents eat the seeds or other organs of

agaves, these armored plants afford them a measure of protection

when the rodents must elude preying foxes, coyotes, bobcats, and

birds of prey by night and day.

Pocket gophers of the genus Thomomys have proved very damaging in the agave garden. They tunnel up through the bases of

the plants and eat out the central meristematic tissue. They work
THE AGAVE FAMILY IN SONORA (

from below and within, out of surface sight, so their attacks may

be discovered too late to save the plants.

Bats have been photographed about agave flowers at night (fig.

2). The bats appear to be after the sweet nectar exuded in the

flower tube, and pollen is plainly evident on the head and whiskers,

indicating that the bats may be pollinators.

Several species of hummingbirds are frequent about the flowering agaves. During daylight hours, they buzz habitually about

the inflorescences, insert their bills in the selected flowers, and

perch on the lofty crossarms between forays. Flycatchers use these

tall perches, and insects are drawn to the strong-smelling flowers.

Doves, flickers, ravens, hawks, caracaras, vultures, and other large

birds also use the tall perches. The mutilated flowers, as well as

broken branches, appear to be due to bird activities. In Sinaloa,

parrots were observed eating agave flowers to such an extent that

many spikes had to be examined before undamaged flower speciPN-2107

FIGURE 2.—Bat at flowers of Agave schottii. (Photo courtesy of George Olin,

Naturalist. National Park Service.)
8 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

mens could be obtained. The capsules may be carried about by

strong birds and the seeds inadvertently widely scattered. Although this has not been observed, missing capsules in inflorescenes have been noted.

Agave flowers attract numerous insects like flies and beetles but

chief among these insects are Hymenoptera—the honey bees, wild

bees, and wasps. In Baja California honey bees were observed to

suck dry a massive bloom of Agave goldmaniana Trel. during the

early morning hours. Bumble bees were also observed visiting

the flowers of A. parviflora flexiflora, the structure of the flower

appearing to be specially modiñed for the bee's convenience. In

the summer of 1963, small, pollen-gathering bees were very active

on A. striata Zuce, and A. roezlii Baker near Tehuacan, Puebla,

divesting the anthers of all pollen before dehiscence. Wasps gorge

themselves on the sweet nectar. Hemiptera of the family Coreidae

regularly live with agaves and the pustulate scars dotting the

leaves and fruits are apparently due to the probings of these sapsucking insects. The gardening personnel at Huntington Botanical

Garden, San Marino, Calif., reported the loss of many small agaves

due to an unprecedented attack by small sap-sucking insects.

Agaves regularly host insects that live as borers in the stems,

leaves, and fruits during larval stages. These include butterflies

of the family Megathymidae and moths of the family Prodoxidae.

The host-parasite relation is conspecific in some cases. Carpenter

bees sometimes nest in the dry agave stalks, making deep borings

in the soft pithy tissues. A large snout beetle bores through the

stem and leaf base. In Mexico the larvae of one of the Megathymidae is eaten dried and roasted or is powdered and used as a

piquant flavoring. In Las Vigas, Veracruz, it was recommended

raw as a remedy for rheumatism, and my faithful assistant, Juan

Arguelles, who was suffering from sciatica at the time, valiantly

ate one, fresh and squirming, from a big maguey. Buprestids and

click beetles observed about agave plants may be predacious upon

the larval borers.

The fungi Diplodia theobromae and Colletotrichum agavae may

cause serious damage to agave leaves, for they have been reported

injurious to the fiber plantations in El Salvador. The climate of

El Salvador is relatively moist for fiber culture and probably explains why fungal diseases are not serious in drier regions where

agave plantations are generally found. On the whole, agaves are

remarkably free of plant diseases.

In summary, agaves operate with other elements of the vegetation to support a wide variety of animal life. The bats, hummingbirds, and some of the insects may serve as pollinators. In some
THE AGAVE FAMILY IN SONORA V

of the nonselfing agave species, this could be critical for their

survival. Conversely, the loss of agaves in some areas v^here they

form a dominant part of the vegetation could result in a decrease

of the animal life. We need to know much more about the biotic

relationships. For instance, the connection betv^een bats and

flowers may be a real mutualism and of evolutionary signiñcance.

However, whether this is of beneñt to either organism, we cannot

positively say. The sweet nectar might cause injurious dental

caries in the bats, or the cross-pollinations might genetically

'^swamp out'' a superiorly adapted variant among the agaves. We

can only assume that this mutualism may be of beneñt to the participants because they coexist in a functional relationship. The

agave fauna will appear to him who will sit quietly under a bush

and watch.

USES OF AGAVE AND YUCCA

Agave

Agaves have many uses. The fibers of sisal and henequén are

worldwide in commerce, whereas fibers of other species have more

limited uses. The coarse fibers of Agave lecheguilla Torr, and relatives are used for brushes. Many of the Sonoran agave fibers have

local uses in twine, rope, mats, saddle pads, packing, brushes, and

sacking. In preColumbian times, the Amerindians used fibers in

skirts and ornamental dress but these fibers are too coarse and

harsh to compete with cotton and modern fabrics for clothing.

Agaves still serve as food plants to many country peoples, including some in Sonora. One of my earliest experiences in the

local country cuisine was chewing on chunks of pit-baked agaves

prepared by the Mayo Indians along the Rio Cedros in 1933. The

white central tissue in the short stems or "cabezas'' is still eaten

as are the young tender inflorescence shoots, and many an agave

maturation is thwarted by cutting off the flowering shoot. Many

people believe that this decapitation also improves the sweetness

in the cabeza because the rosette goes on living for another year

or more. The starchy tissues contained in these cabezas are converted to sugars by cooking. The cooked tissue tastes like strong

molasses, frequently with a harsh, after-swallow feeling in the

throat. It may act as a purgative on those unaccustomed to this

food, to the amusement of the natives. Chewing frees the juice

from the fibers, which ball up in the mouth and are spit out, forming the ^'quids'' of archeology. The sugar in agaves and some other

members of the family was an important source of energy to the
10 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Indian peoples. However, not all species are edible and some of

the species reported as edible in publications are mistaken identities; for example, A. schottii was reported as eaten by the Tarahumara Indians in an area where the plant is not known to occur

(38).

In northeastern Mexico, agave leaves are fed to livestock. In San

Luis Potosí in 1963, I observed A. salmiana Otto ex Salm. being

carted daily to the dairy herds supplying milk to that city. The

fresh green leaves amounted to several thousand tons annually.

Along with opuntia pads, agave constitutes an important animal

food resource in that desert region.

Several of the Sonoran agaves are high in sapogenin content

and are used in washing clothes, animal hides, and hair. The basal

part of the leaf of A. vilmoriniana is used as a readymade cleansing brush, with a built-in soap, the water working into a bubbly

froth as the fibers are brushed upon the object being cleaned. Some

of the sapogenins, such as hecogenin and smilogenin, are used

in medicine as precursors for fabricating cortisone and the sex

hormones. The British and others are using hecogenin in pharmaceuticals as a by-product recovered from waste in sisal and henequén operations in Africa and Asia. The U.S. Department of

Agriculture in cooperation with the U.S. Department of Health,

Education, and Welfare conducted a survey for sapogenins between 1948 and 1956 (9) and the analytic results were published

(52-56), The analyses for the Sonoran Agavaceae are indicated brieñy below in the annotations for the respective species.

The Department of Health, Education, and Welfare is presently

(1971) conducting a survey for cancer-inhibiting constituents in

plants. Several agaves have inhibitive effects on pathological tissues, but the constituents have not been chemically identified and

the enquiry is in preliminary stages.

Such chemicals, some of which are already launched in industrial commerce, represent potential plant resources. It has been

estimated that the steroids in A. vilmoriniana would yield a gross

value of $6,000 to $7,000 per acre, with smilagenin at $10 per

pound (22). However, the development of smilagenin in the leaves

of the plant requires a full growth cycle of 7 or 8 years. The green

leaves of growing plants assayed only 1 percent or less of smilagenin, whereas the leaves on matured drying plants after flowering assayed 3 to 4.5 percent. The Warihio Indians knew this before we knew it and employed only the old dry leaves as soap.

Such prospects are long-term propositions such as tree farms,

which require large capital outlay. They naturally exclude the

adventurous small businessman who would pioneer on a short-
THE AGAVE FAMILY IN SONORA 11

term basis, and big business is inherently too conservative to pioneer. Such situations should merit government attention, but our

legislatures annually have many graver issues demanding their

attentions. Political or other conditions, however, could abruptly

change the steroid resource situation ; for instance, a shortage of

diosgenin from the wild Dioscorea plants, which currently supply

about 80 percent of our steroid imports. Agaves may eventually

be developed as a pharmaceutical resource, but none of the native

Sonoran species are presently (1971) so employed.

The Indians have reported agave leaves to be used as poultices

for itches, sores, bruises, and wounds. Pennington reported that

several agaves are used by the Tarahumara Indians as fish poisons

(S8), In addition to sapogenins, the leaves have been reported to

contain sterols, terpines, and vitamins (10, 53). The leaf chemistry

is complex and we know little about it.

There are many local special uses of agaves. I have observed

the dried flowering shafts used in fences and as props under fruiting branches in orchards. A modern innovation is the use of agaves

in Arizona as Christmas trees. In the United States their greatest

use is in ornamental plant culture and they are important as food,

shelter, and roosts to wildlife.

Castetter, Bell, and Grove presented an ethnic monograph of

agave uses in the Southwestern United States (8). They summarize American Indian uses : "In the Southwest, agave was used

for a considerable variety of purposes: food, alcoholic and nonalcoholic beverages, syrup, fiber, cordage, nets, bags, basketry,

mats, blankets, clothing (particularly aprons), sandals, pottery

rests, headrings, braids and other miscellaneous woven objects,

hair brushes, paint brushes, needle and thread, fish stringers,

armor, lances, fire hearths, musical instruments, paint, a gum-like

caulking material, soap, for smoking, medicine, and ceremonial

objects. Of these uses, food and fiber were most important and

most widespread."

The Aztec codices (17) and living Mexican peoples illustrate

numerous other basic and exotic uses of these singular American

plants. Many of the uses were never developed in Sonora, such as

drinking the sweet juice of the maguey agaves, either as ''agua

miel" or as ''pulque" made by fermenting this juice. The distilled

liquors of mescal and tequila did not appear until Hispanic occupation, for the Amerindian did not have distillation in his culture.

The Sonoran Indians, however, drank the fermented juices of

cooked agaves in ceremonies and cures. Those desiring more information on the uses of agaves would do well to consult the information retreaval compendium prepared by Ida Langman (31).
12 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Yucca

Yuccas were also important, useful plants to the native peoples

from the earliest times, as represented by archeological remains

found in prehistoric dwelling sites. As with agave, the strong

fibers of yucca were used for clothing and for cordage. The leaves

were used more in basketry and continue to be so used. The stems

were used in constructions for shelter. The young flowering shoots,

fruits, flowers, and inner stem tissues were used as food. The seeds

are oily but also contain sapogenin, and I find no reference to

them as food. In Sonora the sweet ripe fruits of the fleshy-fruited

species are still gathered and eaten either raw or roasted. Cruse

(10) reported fructose, fructosans, dilevans, and glucose present

in several yuccas. Wolf (60) found that the dried fruits of Yucca

brevifolia Engelm. compared favorably in food value with that of

dried orange peel.

Yucca flowers have long been eaten by the peoples indigenous

to yucca regions. The flowers are generally cooked by boiling and

eaten as a vegetable, or combined with other foods such as scrambled eggs. Miller and Hamilton (37) reported yucca flowers (probably Y. elephantipes Regel) to be rich in ascorbic acid (vitamin

C) as follows:

Ascorbic acid Moisture

Plant part Mg./g. %

Raw bud 474.8 82.4

Raw flower 371.3 85.4

Cooked bud 489.7 81.0

Cooked flower 360.0 84.2

In the more sapogenous species Y, arizonica and Y, grandiflora,

only the petals may be eaten, consumer informants having stated

that the gynaecium was too bitter. By the use of such wild foods,

the Amerindian could obtain his dietary necessities.

The indigenous uses of yucca have been thoroughly reported by

Bell and Castetter (3), who stated that 'Vucca ranked foremost

among the wild plants utilized by the inhabitants of the Southwest. It holds this place because of the great varieties of uses to

which it could be put and to the wide accessibility of the genus

within the Southwest."

More modern uses of the yuccas include Y. glauca Nutt. and Y.

elata as livestock feed. In times of range feed scarcity, the stems

are defoliated and chopped up for silage (57), Cruse (10) reported the chemical composition of such material on a dry-weight

basis as 4.4 percent protein, 6.7 percent ash, 1.5 percent fat, 3.2
THE AGAVE FAMILY IN SONORA 13

percent fiber, and 49.2 percent carbohydrates. Other chemical

assays show the leaves to contain up to 2 percent sapogenins, useful in the manufacture of steroid pharmaceuticals and also as

bases for soaps and cleansing powders like dental powders and

pastes. The U.S. Department of Agriculture found green fruits

and seeds of some species to contain up to 10 percent sapogenins.

The highest percentage was found in Y. brevifolia and Y. grandiflora. Samples of seeds of several species analyzed contained 17

to 32 percent oil and 12 to 22 percent protein.^

During World War II, the U.S. Department of the Navy set up

a pilot decorticating plant near Kingman, Ariz., for extracting

the fibers of Y. baccata and Y. schidigera Roez. Postwar trials

by private interests who took over the plant pretty well showed

that exploitation of the wild yucca stands could not compete economically with other sources of commercial cordage, although the

fibers were found suitable for rope, twine, and burlap. Because

of the splintering effect due to shorter fiber cells, the fibers of

yuccas are generally inferior to those of agaves. Weber reports

(57) that juice from yucca leaves may be more valuable than its

fibers. The juice reduces the surface tension of irrigation water

(thus increasing its penetration in heavy soils), assists in soil

flocculation, and serves as ''a carrying agent for the plant food

chemicals." Many other chemicals found in yuccas have been reported for a variety of potential uses. However, it is unlikely

that any of these potentials will develop in the United States until

such time as yucca is put into commercial plantings because the

dispersed nature of the wild stands over difficult topography makes

their collection uneconomical. The commercial cutting of the tree

yucca, y. fillfera Chab., as a source of paper pulp in Central Mexico

is a notable exception. The scarcity of a natural paper resource in

that country and the relatively weak competitive position of the

peso have favored the development of a local industry. Y. filifera

requires a century or more to grow, so it is unlikely that this enterprise will continue any longer than it will take to cut the existing wild stands convenient to roadways.

In the United States, yuccas are valued principally for their

ornamental and wildlife attractions. Many of them are being preserved in several public parks, such as the Joshua Tree National

Monument and Borego State Park, both in arid California. Yuccas

' WOLF, IVAN, and others. U.S. Dept. Agr. Rpts. Unpublished and interdepartmental, Northern Utilization Research and Development Division, Agr.

Res. Serv., Peoría, 111. 1960-65.
14 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

are increasingly used for landscaping private homes and public

grounds, particularly where water is scarce.

HISTORICAL NOTES ON AUTHORS AND

COLLECTORS OF AGAVACEAE IN SONORA

During the 19th century three agaves were reported from

Sonora: Agave parviflora, A. schottii (as A. geminiflora sonorae

Torr.) in 1859, and A, schlechtendalii Jacobi in 1864. Jacobi (25)

observed A. schlechtendalii in several north European gardens

and gave its origin as the Province of Sonora, Mexico. I have not

been able to recognize the plant from either Jacobi's descriptions

or Berger's later one, all drawn from immature specimens. Berger

placed it in the section Americanae.

In 1920 Trelease added A, subsimplex and A, yaquiana, the latter *'from rocky hillsides between Hermosillo and Ures, Trelease

391," which I am merging with his A. pacifica from Mazatlan. In

1924 I. M. Johnston described A. owenii, — A. pacifica Trel., from

"an islet in Guaymas Harbor, Sonora," and A. chrysoglossa from

San Pedro Nolasco Island off the Sonoran coast (29). In 1942

Gentry added five species, one of which is synonymized here, and

brought the total known agaves in Sonora to 10 species (13).

In 1964 Shreve and Wiggins ascribed 10 species to Sonora (i2).

Two of these, A. hartmanii S. Wats, and A. mescal Koch, are here

excluded as nomen confusum and nomen nudum, respectively, as

discussed below under A. polianthiflora and A. bovicornuta. Wiggins prefaced his taxonomic treatment with the following interesting commentary. "The treatment of the genus is dishearteningly inadequate. The range of variation within a species is often

extremely puzzling, and it is probable that hybridization occurs

frequently. These conditions make a satisfactory taxonomic solution exceedingly difficult. A reasonably acceptable classification

of the genus Agave can be presented only after extensive and

painstaking study in the field, prosecuted at all seasons of the year.

The keys and recognition of species here are strictly tentative."

The present treatment describes 27 species, 9 of which are new,

and 2 new subspecies. It seems unlikely that more wild species

will be added, although some of the coastal postinsular mountains

may hold some surprises. Trinomials are increasing in modern

taxonomy, so we can expect more of these as our knowledge of

relationships grows.

The botanical collection of Sonoran agaves can be briefly outlined, as based upon the herbarium and botanical garden specimens reviewed, as follows :
THE AGAVE FAMILY IN SONORA 15

Specimens

Year Collector Number

1855 Arthur Schott 2

1890 Edward Palmer 2

1899 E. A. Goldman 1

1899 E. W. Nelson 1

1900 William Trelease 1

1910 Rose, Standley & Russell 5

1921 I. M. Johnston 4

1923- Forest Shreve 4

1932-48 Ira L. Wiggins 4

1937-67 H. S. Gentry 110

1938-40 Steven White 4

1941 Wiggins & Rollins 1

1951-67 Gentry & Arguelles 40

1958-67 Richard S. Feiger 34

1966- Gentry & Weber 5

1966- Barclay & Arguelles 5

It might appear that more than 200 specimens of about 30 taxa

of agave in Sonora would provide a fair sample for taxonomic

revision. This is not the case. Many of the collections are fragmentary, consisting only of pieces of leaves or of capsules. Many

of my collection numbers are represented only by live plants sent

to botanic gardens, with the expectation of obtaining flowering

specimens in later years. As a result of neglect, loss of labels, lack

of a botanist at hand for collecting at the time of flowering, and

depredations by rodents, over half of such collections have been

lost. Only three species are represented by a series of flowering

specimens: Agave pacifica with 10, A. palmeri with 9, and A.

shrevei with 6. Most of the others are represented by 1 to 3 flowering specimens ; for one, flowers are unknown. Generally, the rarity

of flowering specimens is due to their highland inaccessibility, the

rarity and briefness of their flowering seasons, and the reluctance

of botanists to spend the time and effort necessary to collect the

spiney, wounding, and hard-to-dry succulent plants. The late Susan

Delano McKelvey left a note in the herbarium at the Arnold

Arboretum on collecting A. chrysantha Peebles near Sedona, Ariz.

''Reduced to crutches (all way back east) after running a terminal

spine of this plant into my ankle; never retrieved by Dr." Some

spines appear to carry a toxin for they cause swelling and pain

beyond what a mere puncture would produce.

Collections of other genera in the Agavaceae are even scarcer

than those of Agave, The specimens reviewed and cited for Sonora

in this work total only 52, as follows : Yucca 25, Nolina 10, Dasylirion 8, Manfreda 5, Hesperaloe 4. These specimens are all I found
16 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

in the following herbaria : U.S. National Arboretum, Smithsonian

Institution, Arnold Arboretum and Gray at Harvard University,

University of Arizona, and Gentry. Doubtless a fev^ other specimens exist in herbaria I did not review. Altogether, the specimens

are insufficient for an adequate account of the family in Sonora.

Our knowledge of Sonoran plants is primarily due to three

works : The vegetation and floristic treatises of Shreve and Wiggins, which cover the desertic region of the northwestern part of

the state H2) ; Stephen White's account of the flora on the mountainous area comprised by the Loop of the Rio Bavispe in the

northeastern part {58) \ and Gentry's survey of the Rio Mayo

basin in the tropical southeastern part {13), Most of the specimens cited here (agaves excepted) were part of the general floristic sampling on these projects and on the unpublished work of

Richard Feiger. There is still a great deal to be discovered about

the plants of Sonora, even unknown plants. It is a rich field for

botanical research for anyone who has the capacity to engage it.

The writer is grateful to the many people who have helped him

in so many ways in his study of the agaves. The Sonoran people

are first among these because from the beginning, they were always ready to give information about the stands of wild plants,

the trails and roads for reaching them, the names of plants, and

their uses. These people included businessmen, ranchers, miners,

and the specialists in making ''mescal," the ''mescaleros." Sonora

has an obscure maze of side roads and trails. Where routes are

too long or complex to follow by directions, local people will frequently serve as guides. Many will refuse pay, but tips or gifts

are acceptable, particularly to children. The native Sonoran people

are among the most polite and hospitable in the world and can

contribute to any receptive traveler's education. The Sonoran

character is epitomized in Juan Arguelles of San Bernardo, my

assistant for many years, who accompanied me on my agave

forays.

Among the institutions that made their collections and facilities

available are: The Huntington Botanical Garden, San Marino,

Calif.; Desert Botanical Garden, near Phoenix, Ariz.; and the

Herbarium of the Botany Department and the Ecology Herbarium

of the Zoology Department of the University of Arizona. The loan

of Agavaceae collections of Richard S. Feiger of the University of

Arizona has been particularly helpful. His collection labels carry

the names of several student assistants : R. L. Bezy, Oda Kleine,

Joe T. Marshall, Peter Marshall, Becky Moser, Cathy Moser, Alexander Rüssel, Robert Rüssel, Dirk D. Stronak, and Robin Thomas.
THE AGAVE FAMILY IN SONORA 17

These were all students of Charles Lowe of the Department of

Zoology.

The materials in the Harvard University Herbaria and their

extensive botanical libraries were made available. The historical

literature there and the agave collections made in Southwestern

United States by Susan Delano McKelvey were of paramount value

for comparative studies.

The National Herbarium of the Smithsonian Institution in

Washington, D.C., has the most complete agave collection in the

world and includes the classical specimens made by Alwin Berger

at the Hanbury Gardens (La Mórtola) near Ventimiglia, Italy.

The Smithsonian collection and my own form the core of evidence

accumulated in this work. All type specimens of new species described here are deposited in the Smithsonian Herbarium, along

with duplicates of most other numbers I collected.

The drawings of Regina 0. Hughes, Plant Science Research

Division, ARS (retired), speak eloquently of her careful labor.

I am grateful to Frederick Hermann of the U.S. Forest Service

Herbarium in Washington for editing my Latin diagnoses.

SOME CHARACTERS OF AGAVE

Generally, agaves are succulent plants with spirally arranged

leaves forming a rosette, some with definite trunks, but more

often nearly stemless. Agaves are like annuals in that the rosettes

are monocarpic (in Sonora) and ñower but once and die. They

are like perennials in regard to the time needed to reach ñowering maturity. However, perennial as a term does not suit them

because perennials flower repeatedly. I have therefore called

agaves multiannuals, meaning plants that require several to many

years to mature and flower once. Many of the palms have this

same habit. However, contingent perennials may characterize the

agaves better because their floral maturation appears to depend

not only upon a number of years of growth, but also upon favorable coordination of seasonal temperatures and rainfall. Rosette

or growth maturity may be reached several years ahead of floral

maturity. This phenomenon is more apparent in more arid environments and reflects the holding capacity of agaves as adaptation to desert conditions.

Ehrler (11) has recently discovered that unlike most plants,

agave stomates close by day and open by night. This prevents the

loss of water through transpiration during hot daylight hours,

although it results in a hot leaf surface, hotter than most plants

tolerate. Damage to the leaf by high temperature is apparently
18 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

prevented by the thick cuticle overlying the epidermis, v^hich is

SO evident in desert species of agaves. This circadian mechanism

and specialized structure are adaptive characters evolved at the

environmental edge of life where water is critical. The agave leaf

forms a greenhouse over itself, insulated for protection against

sun and wind, and translucent to admit the vital light to the photosynthetic cells. The Mixe Indians of Oaxaca use this peelable

waxy layer of the agave leaf for wrapping tortilla sandwiches,

as we use our modern plastic bags. One may well wonder how this

leaf structure came about. The countless gene complexes that were

formed were either rejected or accepted by environmental stresses

until gradually and finally the genetic combinations balanced and

structures were synchronized with the increasing deserts. This is

a most interesting area about which little is known, and original

research, not only in genetics and evolution but also in ecology and

physiology, could find principles applicable to more than agaves.

It is out of such particulars that man forms his cogent universals.

The sizes and outlines of the mature rosettes are characteristic

and expressive in many agaves. Some radiate the leaves out stiffly,

forming an armed hemisphere ; others may be urn-shaped or balled

by the incurving leaves. The larger leaves sometimes collapse and

drop in severe droughts, spoiling the healthy posture. Many species

tend to develop a fixed number of leaves, 60 to 80 being median ;

others may develop as many as 300. However, this number is not

fixed because individual conditions or cultivation in foreign sites

may delay flowering for years while leaves go on developing. We

know little about the factors that finally trigger the maturity of

inflorescence, but maturity appears to be a coordinate timing of

rainfall and seasonal temperature gradients. Most species are

armed with spine-tipped leaves, and many have strong marginal

teeth as well.

The relative sizes and shapes of leaves are important. Leaves

vary greatly according to age, and their ontogeny is a study in

itself. The taxonomist does better to compare and use only the

mature leaves. The stifit sword leaf is always the stiff sword leaf

(unless it is in shade) and other forms may be just as characteristic of other groups and species. The tendency to be narrowed

just above the spreading, clasping leaf base may be fairly constant throughout species populations. This is a helpful character

in separating the broad-leaved A. shrevei from the related and

relatively narrow-leaved A. palmeri, because the latter has little

or no narrowing below the midblade. Leaves may be thick or thin,

hard or soft, according to the species. Basal thickness is correlative with the length and weight of leaf as a structural necessity
THE AGAVE FAMILY IN SONORA 19

for support, so in itself adds little to diagnosing a leaf when the

dimensions are given.

The margins of leaves show many characters. On leaves of A.

palmeri the margins are nearly straight. Other species have leaves

with crenate or undulate margins due to the fleshy prominences,

teats, under the teeth. This crenate character is variably present

in A, shrevei and A. bovicormita, while it appears always well

developed in others, for example, A. colorata, A hard, corneous

margin may be constantly present in some species, but either

present or lacking in others. Frequently, this margin is limited

to the apex of the leaf as a deccurency of the spine base. As far

as I have observed, the corneous margin is invariably present in

the section that takes its name from this structure, the Marginatae, which is not known in Sonora.

Leaf color, though overlaid and lightened by a pruinose or

glaucous cuticle, is characteristic in some species, as the dark

green of A. wocomahi, the yellow of A. cerulata, and the light gray

of A. deserti. In other species leaf color is variable. Light glaucous

forms are common in many species and this character can be used

safely only in conjunction with other characters. The center pale

stripe, which has been used to characterize species, is a form that

occurs homologously in many species.

Teeth or marginal prickles occur in many species in both subgenera, but many species in Littaea never have them. In subgenus

Agave they are more nearly universal, although there are a few

species in which teeth occur only weakly or sporadically, as in

A. desmettiana and A. sisalana Perrine. In other cases, toothless

forms appear in species that are generally well stocked with teeth,

as A. deserti and A. parryi Engelm. Other characters in leaf and

flower identify these as merely forms. Teeth have many forms,

varying on a single leaf from small towards the leaf base to large

in the midblade, flexed upward or downward, curved or straight.

There is intraspecific variation, like subtle composition, and it

serves to characterize some species; e.g., the down-flexed teeth

along the basal half of leaves in A. asperrima Jacobi. An unusual

and persistent tooth type helps mark some species, as the round

tooth form, in cross section, in A. applanata Koch and the fishtailed or biscuspid form in A. goldmaniana. Study of teeth variations in populations teaches one how to read them out for species

orientation.

Terminal spine forms are less bizarre than teeth but they also

vary within species. In some cases it is difficult to determine the

specific dimensions of their variability, while in others there is a

more recognizable definite tooth form. Spines are sometimes dis-
20 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

torted in the bud by the exigencies of early growth and emerge

distorted or flexed sharply downward. A still unrecognized species

in Puebla has a spine sometimes centrally decurrent down along

the ventral keel of the leaf and bearing small teeth, incongruously,

along the midleaf line. For self-perpetuation through armor, evolution is producing another innovation.

Some characters, as central leaf stripe, bicuspid or fishtailed

teeth, polycarpy, and toothless leaves, appear in unrelated species

more or less at random through the genus or even in the genera

Manfreda and Furcraea. Such characters recurring in separate

genetic lines are to be regarded as homologous variations and have

been discussed by Vavilov and by Darwin as convergent or parallel

evolution. This is briefly discussed in ^^Concept of Species,''

page 25.

Flower structure has been the most important guide to plant

classification since before Linnaeus. Because of the general lack

of flowering material, floral morphology has been neglected by

agave taxonomists. Berger employed some floral characters in

setting up his sections, but the flowers of many species he and

others described were like their origins, unknown. My incomplete

study of flowers finds them indispensable for an adequate classification and my revision of the genus is still concerned with obtaining flowering specimens of many presumed species.

Agave flowers take many forms although there is a basic likeness throughout their development. The ovary is inferior in all

taxa but not quite completely so in what I consider a primitive

section, Striatae (not known in Sonora). A definite tube is usually

present but is virtually lacking in A. bracteosa S. Wats., and A.

guiengola Gentry (also not known in Sonora). Besides being either

deep or shallow, the tube varies in its structural components, in

the nectarius inner lining, in ribbing, in form, and its relation

with the tepal lobes and filaments. The tepal lobes have been modified in various ways. In the putative primitive form, the six tepals

are essentially equal and all alike. More specialized forms develop

larger outer tepals, such as those that characterize my section

Ditepalae. A 2-ranked perianth has tended to develop with biserial

modifications in form and structure, such as in the conspicuous

keel on the inner tepal, which is lacking on the outer. The tepals

may be open and reflexing or erect to incurved, sometimes involute; some wilt early, others wilt late or not at all. The outer

tepals are often hooded and apparently always glandular papillate

at the tip in varying degrees or patterns. In some flowers as in

the section Marginatae, the tepals fold about and support the long

filaments.
THE AGAVE FAMILY IN SONORA 21

The insertion of the filament in the tube is a valuable character

on the sectional level. The filament may be inserted in the base of

the tube or on the sides of the tube and, at its upper extreme, on

the base of the tepals, as in the Deserticolae. In species v^ith a

thick glandular or nectariferous tube lining, the base of the filaments grows as out of an inner rim, forming a six-point star

shape, as shown by Regina Hughes in figure 22, page 77. Length of

filament is variable by growth and not all filaments of the same

flower are equal, but in species with exceptionally long or short

filaments, the character becomes significant, as with the long

stamens of A. americana L. Anthers are much alike in gross morphology. Size is generally correlative with flower size, but when

this ratio is exceptional, size of the anthers may be of diagnostic

value. Anther fixation is always versatile, centric or eccentric,

but it appears irregular within species and I find it an unreliable

character.

Flower colors range from the common greens and yellows to

various shades of purple and red. The strong, dark red found in

A. pelona is exceptional, as is the nearly white bloom of A. braeteosa. Flower color varies within species, with yellow, lavender,

and pink forms being found like Mendelian segregates in A.

striata. The range of variability of flower color or its constancy

in populations may be of taxonomic value, but generalized statements of flower color may be misleading. Different flower parts

frequently show color differences, as green ovaries and yellow

tepals, or yellow perianths with pink filaments. Anthers are commonly yellow, but pinks and reds appear and their color changes

with dehiscence.

Capsules and seeds have not been thoroughly studied. The

former vary considerably in size and shape in a single infloresence and cannot be used to separate species, as Trelease used them.

There are gross differences in stipitation, beaking, sizes, and ribbings between sections. Seeds also show significant size differences

between sections and it is quite probable that microscopic studies

would reveal further taxonomic criteria.

The inflorescence shaft with its branches is a massive structure,

generally distinct as the spicate form of subgenus Littaea and the

paniculate form of subgenus Agave. However, these two forms

are not always just one or the other because there are intermediate

forms, as of A. utahensis Engelm. and A. potatorum Zuce, where

the lateral flower umbels may be upon spreading lateral branches

or sessile on the shaft. Diagnostic features of value have been

noted in the ratio of peduncle length to inflorescence length and

the general outline of inflorescence and individual flower clusters,
22 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

whether broad, deep, or narrow. The series of bracts generally

decrease in size from the base to the tips of the inflorescence in

a graduated sequence. On the whole, they express little specific

difference, but valuable exceptions do occur, as the fleshy imbricated bracts in the section Salmianae, the short bracteoles of A.

pacifica, and the long acicular bracteoles of A. lophantha Schiede

contrasting with the short ones of A. lecheguilla.

Flower size varies from the lO-mm.-long flowers found in A.

parviflora to the over lOO-mm.-long flowers in A. salmiana Otto

ex. Salm. The size ratios between floral organs have been found

useful in making diagnostic alinements of sections and species.

The diagrammatic drawing in figure 3 shows the various measurements of flower parts. These measurements are useful for analytic

comparisons and the floragraph is a kind of taxonomic shorthand

based on the ratio of tube and tepal lengths, further qualified by

the position of filament insertion.

^-^ a = 24

> f = 35

s= 19 a 17-á

ovory

^* 0 = 25

^'ji^^. a Hiu^Uu

pedicel

brocteole

o-ovo..oa,,en,.. „ - „ecK o, ovo.. .„.H. t-,o.e ,en.., ...,o.en, .se.on

(measured to bottom of t-tube) s tonm i *K r
THE AGAVE FAMILY IN SONORA 23

Gross morphology is indeed gross in agaves, because all essential organs are large scale. Variability is expressed profusely and

smites the naked eye impressively. I have generally limited my

taxonomic criteria to the macroscopic characters rather than belabor the microscopic for the following reasons :

(1) Except for certain floral structures, there appear to be

ample characters for classification in gross morphology.

(2) Much of the use of this handbook will probably be made

by people who will be looking at living plants, either in the wild

or in cultivation.

(3) Brief descriptions of principal characters are better than

long, detailed ones when ample illustrations are provided.

It is hoped that users of this handbook will respond to the deficiencies they find because this is also a preliminary trial for a

general revision of the genus. Critiques can help to improve the

next one to be written.

Other Sonoran genera of the Agavaceae have been included here

to orient the genus Agave more completely and to make known

other obscure Sonoran plants of the family. Like the agaves, the

yuccas of Sonora have scarcely been collected. I have known several undescribed species for years. It is time they were formally

registered. There is hybridization in both Yucca and Agave, creating puzzling problems. To unravel them will take more time

than this one life; for such a task a man would be fortunate to

extend himself by offsets, like an agave. As it is, one will have

to let the next botanical generation do it.

The best single taxonomic contrivance for identification of the

Agave taxa is their separation on a geographic basis. This permits

the use of some obvious leaf and rosette characters, which are

not serviceable when treating large numbers of taxa in the genus

as a whole. There are too many homologous characters in unrelated species or too little leaf difference in some related species

to separate them without inflorescence characters. Inflorescence

characters are frequently unavailable to those who wish to identify the living wild and cultivated representatives. Hence, whenever possible, for separating taxa, I have used both vegetative

and inflorescence characters, supplemented by distribution data.

CONCEPT OF SPECIES

The species in agave are frequently what may be called hard

cases. This is partly because many species grade morphologically

and variously towards other species and partly because traditional

taxonomy has erected many specific names on poorly understood
24 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

leaf variation. Jacobi (24-27) named many species on the trivial

differences exhibited by cultivated plants, many of them immature

and grov^ing unnaturally in pots and greenhouses and of unknov^n

origin. Berger (5) used floral morphology mainly for his sections,

but regarded Jacobi as an authority and conservatively follov^ed

close splitting in recognizing species. Baker (2), just before

Berger, was also unav^are of the natural variability of populations

in their native states and maintained narrow specific limits. Engelmann was first to evaluate characters in agave and broadened

the concept of species, but his studies were brief (12). His successor, Trelease, was astute in recognizing differences, and, as

though he saw more variability than he could conveniently handle,

came to place undue weight in a single character—the terminal

spine (47), It is very useful for separating clones when growth

factors are considered, but agave populations demonstrate that

like other organs the spine too can have a wild variability. Trelease, however, was barely acquainted with wild agave populations. Most of the agaves he observed on his Latin American

forays were cultivated clones in fields and hedgerows.

The namers of the agaves cultivated in Europe were horticulturists concerned with naming their novelties. The names satisfied

their needs for communication, but the practice gave taxonomy

a poor start. It set a precedent that conservatism and ignorance

follow and that is inadvertently supported by the priority of

names. When you name a thing, you have it, in a limited sense,

but a label does not always tell what is inside. Every agave is

a one-million-year-old package still ticking out genes and, to this

practicing taxonomist, they sometimes appear like challenging

fiends. Our pages will grow to many before our taxonomic formulas are very truthful.

Evolutionary biology has emphasized that species are composed

of populations. This concept is very effective because it leads us

to evaluate variability as a collective whole. The reproductions

of variability patterns become taxonomic criteria that the systematist can apply to taxa as he judges them on the generic, specific, and subspecific levels. However, if botanists accept zoologist

Mayr's (35) bological species concept entirely, ''groups of actually

or potentially interbreeding natural populations which are reproductively isolated from other such groups,'' we are soon in trouble.

Hybrids are much more common among plants than animals. They

occur not only among species but also between genera (U)^ Mayr's

phrase ''potentially interbreeding'' is particularly unsuitable for

the definitions of plant species. We do not know what species are

potentially crossable, and considering the hundreds of thousands
THE AGAVE FAMILY IN SONORA 25

of plant species, it would take us forever to find them all out. For

instance, species are known, that are isolated genetically and geographically, but may be bridged by a third species carrying genes

between the isolates. (See Harlan and De Wet compilospecies

{20),)

If one were to apply Mayr's species definition to the agaves, not

only would many species be abandoned, but subgenera would be

in danger also {16). We might eventually end up with one species.

Agave americana, which would satisfy nobody with an interest in

plants. It is not surprising that animal species are basically different from plant species. There are many different kinds of plant

species, as we have come progressively to realize ever since the

arrival of Camp and Gilly's article on this subject nearly 30 years

ago (7). There is no universal model for species, no adequate

definition for all plant species ; and there are about as many concepts of plant species as there are plant taxonomists.

Species are interfertile groups of individuals composing one to

many populations with a recognizable unison of characters. This

statement appears to express the working concept I have followed

in this enumeration of Sonoran species. Regardless of how species

are defined, the practical purpose of a taxonomic work is to show

both close and more distant relationships.

There are inbreeding and outbreeding agaves and agaves that

do not breed at all. There are diploids and polyploids {18), and

clones that change their karyotypes have been reported (^0).

Variations are phenotypically expressed in cells, organs, habits,

and physiologies in baffling sequences throughout the genus.

Homologous variation presents special problems for the taxonomist. Certain characters appear irregularly among species,

among genera, and even among families. I have listed some of

them above that have been observed in agaves—leaf stripe, bicuspid teeth, toothless leaves, and polycarpy. These characters are

misleading for separating species and especially mischievious in

keys. The presence or absence is doubtless controlled by genetic

factors, but the mechanism is not understood. It appears to be

genetic segregation operative on a long-term basis, through hundreds of generations rather than through the few generations

that geneticists have documented by experiments with short-term

segregating characters. New analytic approaches must be developed to study the geologic-long segregates of homologous variation. The comparisons of distantly related DNA slurries of mice

and men, as reported by Hoyer, McCarthy, and Bolton {23) demonstrate the compatibility in portions of distantly related chromosomes. Such experiments indicate the great age of some genes,
26 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

common origins for higher categories, the great truth of evolution, and the immortality of the generative cell. When our young

science of genetics has matured sufficiently to show the causal

relationships of homologous variation, we can then orient it more

properly with other biological characteristics. At the extremes of

the genus, one finds taxa that could be separated generically were

the continuum only a bit more broken. Natural breaks occur with

distinct species, but more often than not the breaks are expressed

by a lack of concordance in the patterns of variation. Study seeks

to find the limitation of these variations, which appear as subtle

patterns that become familiar with repeated observations and

comparisons. To solve these problems, I have sought to study the

wild agave populations whenever and wherever the various botanical assignments of the last two decades have allowed me to

do so.

In general, the characters of the several major organs require

inspection and study to determine the species outline. Taxa may

appear very distinct when only one or two individuals are seen,

but lose much of their separateness when seen in their variable

hundreds. Hybrids are very confusing unless they can be recognized as such. Usually a complex of character variables must be

recognized and correlated with distributions and habits. A kind

of unison of characters emerges correlative to time and place. Few

are those single characters that serve to distinguish a species at a

glance, but a few species do have them. Floral morphology is a

unifying guide, showing the common relationships of variables;

it broadens species and reduces surplus names. However, the

flower alone provides no universal frame for every species. Leaf

and rosette characters may or may not display significant differences. Frequently it is the breaks in variation patterns of leaves

that decide species limits. The supporting evidence in distribution,

breeding habits, and biotic relationships provides valuable supplementary evidence to morphology.

My concept of species has been and continues to be an evolving

one. My judgments are based primarily upon certain morphological characters that appear to link variable populations or a single

population together as hereditary self-perpetuating units. I view

species as aggregate lines of descent in the Darwinian philosophy,

rather than as discrete typological units in the Linnaean sense.

I have drawn some specific lines arbitrarily and tentatively because of insuflîcient knowledge of inflorescences and respective

biologies. Two disjunct populations of sectional alinement that

show floral and leaf differences are generally treated as separate

species, but unknown intervening populations or future breeding
THE AGAVE FAMILY IN SONORA 27

work might reduce those disjunct populations to subspecific ranks.

For example, A. inaequidens Koch of the central transverse Mexican ranges and A. bovicornuta Gentry of the northern Sierra

Madre Occidental are separable by flower and leaf variation pattern differences. Since the former is fall blooming and the latter

spring blooming, it seems advisable to treat them as separate

species. However, between these two taxa are several related complexes in the mountains of Durango, Nayarit, and Jalisco. Additional specimens and studies are needed for adequate definitions.

I have tried to maintain a fairly narrow interpretation of species because there is a great convenience in the binomial system

as compared with the polynomial, which in effect is a retreat to

pre-Linnaean taxonomy where descriptive phrases represented the

basic taxonomic unit. I am also reluctant to make synonyms out

of previous students' species unless I have good evidence to the

contrary. I have, therefore, maintained some species that appear

doubtful to me, but for which I have insufficient evidence, morphologically or otherwise, to pass judgment.

Trinomials appear indispensable to account for those numerous

intergrades that are not clearly one species or the other. Trinomials are inevitable with evolving organisms and usually occupy

definite geographic areas in relation to their source species. I call

these subspecies, distinct from the varieties, which in agave are

mainly horticultural varieties based on leaf forms. I find no taxonomic merit whatever in the use of subvarieties and subformas

because they only burden nomenclature and minds with trivial

variations, which are better left as conversational subjects between hobby enthusiasts.

The circumstances controlling my investigations of agaves and

their long generative period have prevented the practice of genetic

experiments or experimental taxonomy to determine relationships.

Our larger herbaria can now provide systematists with sufficient

specimens among many genera of plants for detailed studies in

species variation and limitations. This is not the case with the

Agavaceae because collections have been very few and fragmentary. Most of my specimens for this study were collected from

Arizona to Costa Rica. I have attempted to emphasize floral morphology, so basic to plant relationships, because preceding accounts of the genus have neglected it. This has meant repeated

trips to distant mountain populations. Relationships not apparent

to earlier students are appearing with the growth of the inflorescence collections.

With a feeling of reluctant inadequacy I am attempting to formalize my taxonomic studies in print. There is too much that I
28 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

do not know and I hope this essay will not be regarded as authoritarian. The Sonoran agaves are but a small segment of a widespread, large, native American genus. I think my attention was

first irremedially called when the late great Paul S. Standley in

1934 refused to identify my first collections of Sonoran agaves;

they comprised only two or three specimens of the many hundreds

of other plants he identified. Now, like an agave, I am an old being,

who must make a show after years of rain and light catching.

Among modern men it has become almost mandatory that his

genes be given an outing or an inning via the cerebellum. I hope

this account will serve to acquaint others with the obscure and

interesting Agavaceae in Sonora. Many of them have been markedly depleted by the exploits of man and to be conserved need men

to protect them from other men.

AGAVE FLOWER MEASUREMENTS AND

THEIR IDEOGRAPHS

Flower Measurements

Measurements of organs and parts have been widely employed

in the classification of both plants and animals. Measurements

provide basic data for direct and analytical comparisons between

species and other taxonomic categories or comparisons of individual specimens. Table 1 gives a nearly complete record of flower

measurements made of Sonoran agaves. Figure 3 shows the organs

these measurements represent. However, the experience of making

these measurements teaches that to be reliably comparable, the

measurements must be made in a standardized way subject to

certain qualifying conditions imposed by the developing flower.

The flowers used should be normal, healthy, and precisely in the

stage of anther dehiscence.
TABLE 1.—Agave flower measurements (in mm.): *, measurements from dried flowers; **, measurements

from dried flowers, relaxed by boiling; ***, measurements from fresh or pickled flowers

Taxon and

localityOvary

and neck

Subgenus Littaea

multifilifera

Sierra Charuco _

polianthiflora

Sierra Charuco

parviflora

Í

Pajarito Mountains \

p, flexiflora

Guasabas {

Bacadéhuachi

Matapé

schottii

Pajarito Mountains

Huntington Bot. Gard.

Tube Tepals

Filament insertion Total Collection

and length Anther length No.

1f> 5 5 XIO 17X5 4-5 25 17

14 5 5 XIO 16X5 4-5 32 16

16 5 5 XIO 17X5 4-5 37 17

8 1 30X5 3 6 35 6

8 ?, 30X6 3 X3 4 34 13

8 1 24X5 4 X4&3 4 37 11

8 1 28X5 4 X4 5 37 12

4 ?, 5 X4 2.5 1 9 5.5

5 2 5.5 2 1 10 5

4.5 2 5 2.5 1 10 5

.4.5 2 5 2 1 — —

5 1 4 4-5 0 __

5.5 1 4 3.5 0 11 6

6 8 3-4 4-3.5 0 12

5 2 3-4 4 0 13 6

5.5 2 4 3.5 0 11 6

6 2 3.5 4-5 0 15 5

6 1 3 4 0 13 5

5.5 1 3 4.5 0 15 b

'8 4 10X8 7 6 17 __

8 4 10 7 6 18

8 2 11X7 9 X5 8-9 18 9

¡10 6 10X7 11X5 8 20 10

\^ 5 11X8 11X6 9 20 11

8 4 10X8 15 X 5.5 10 20 15

43*** 8167

40*** 8167

42*** 8167

40* 8013 > 40** 8013

37** 8013 >

40** 8013 <

13*** 17752 >

14*** 17752 g

13*** 17752 f

14*** 17752 H-1

13** 16638 m

14** 16638 O

17** 16643 o

14** 16643 íi

14** 16643

14*** 17744

13*** 17744

14*** 17744

30*** 17751

29*** 17751

30*** 17751

36*** 19858

36*** 19858 IN

36*** s.n. C^
TABLE 1.—Agave flower measurements (in mm,): *, measurements from dried flowers; **, measurements

from dried flowers, relaxed by boiling; ***, measurements from fresh or pickled flowers—Continued

Taxon and Ovary Filament insertion Total Collection

locality and neck Tube Tepals and length Anther length No.

(8 5 8 X8 12X5 5-6 16 16 32*** 21988

Rancho Primavera {9 4 11X8 16X5 8 22 17 40*** 21988

(10 7 10X8 17X5 7 25 17 44*** 21988

Q. ., ^. , 18 5 8 X7 10X4 6 12 11 31*** Noel

bierrita l^icu |g ^ g ^6 11X4 6-7 16 11 31*** Noel

feigeri

n2 2 3 12 2.5 _- __ 30** 11366

n^n\.c.nc. 19 3 2 11 1 20 __ 25** 17882 líacnaca jg 42 12 1 18 __ 25** 17882

19 2 4 12 2 18 __ 26** 17882

Bahia San Carlos __ 3 10 2.5 __ __ __ 11343

vilmoriniana

Q. ^^^ p,^^„_ Í17 3 4X8 16X4&15 4 35 16 40*** 10232

öierra unaruco ^^ 3 4X8 15 X 5 & 15 4 32 16 39*** 10232

(12 3 3 X8 17X4&16 3-3.5 38 15 35*** 2016

Sierra Tecurahui < 14 3 3.5X8 16X4.5 3.5 38 16 37*** 2016

(15 4 4 X8 17X4.5 4 40 16 39*** 2016

TTiir.fir.o.f^r. "R^f Po^^ Í16 4 4 X10 15X4 4 35 16 40*** 19677 Huntington Bot. Gard. 1 ^g ^ ^ ^ 10 17X4 4 34 16 41*** 19677

chrysoglossa

(13 5 4.5X9 15X6 4-4.5 40 15 37*** 19882

Bahia San Carlos { 12 5 4 X9 16X6 4 40 15 37*** 19882

(13 5 4 X9 15X6 4&3 42 15 37*** 19882

Bacanora 13 2 4 13 4 25 __ 33* 16623 ßacanora ^^ 2 4 14 4 26 13 35** 16623

ocahui

Guasabas Í14 1 4 15 4 40 __ 35** 16637

uuasaoas ^2 2 4 13 4 __ __ 31* 16637

CO

o

>

o

o

f

a

>

Ö

W

O

O

o

c!

tn

Ö

O

>

S

a
pelona

Cerro Quituni.

Subgenus Agave

a. expansa

Amado, Ariz.j

bovicornuta

Sierra Saguariboj

Sierra Tecurahuij

Sierra Surotato, Sinaloa

jaiboli

Sierra de la Ventana<

desmettiana

Guasave, Sinaloaj

Brownsville, Tex.j

murpheyi

Huntington Bot. Gard. |

palmeri

Northwest of Nogales{

17

3

9 X17 18X8

9 38 15 47*** 19898

18

38 18X7

8 20 17 47*** 19898

18

3 8.5 18X8 8.5 35 15 46*** 19898

336 14X19 31X9&29 8-9 68 12&16 85*** 21983

32

4 13X19 30X9 8-9 70 28 80*** 21983

20

27 X12 21X4 4-5 40-45 12&8 50** 3672

222

8 19X4&18 3-4 42-45 50** 3672

2967 X13 20

X5& 19 4-5 45 10&14 62*** 2017

28

6

6 X12 19X5&18 4-5 45 23 58*** 2017

>

18

49 X14 15

X 6-7

4 30 18 46*** 18379

O

1948 16

4 34 20 45*** 18379

<

1634 X8 15X4&14 3-4 35 13 36** 21177

244 4.5X8 16X4&15 3-4 35 13 47** 21177

>

244

4 X8 16X5 3-4 36 14 47** 21177

12

1 10X14 14X5&13

6 28-30 15 37*** 11569

Kî

111 10 14

6 25 15 37*** 11569 "Z

14

1 11X14 15

X5& 14

7 35 13 41*** 18411 Ul o

14

1 10 15

7 35 13 40*** 18411

o

28

4 17X15 18X7&17 10

&8 50 11&14 68***

>

31

4 19X19 18X7&17 11-10 45 12&14 71***

354 16 15 10

&9

—— 74***

22

4 13X10 12

X5& 10 7&6 46 17 51*** 17750

26

2 14 12

X5& 10 8&6 42 17 52** 17750

282 12 10 &8 6&5 40 17 49*** 17749

26

2 12 10 &8 6&5 40 9&7 50*** 17749

CO
CO

TABLE 1.—Agave flower measurements (in mm.): *, measurements from dried flowers; **, measurements

from dried flowers, relaxed by boiling; ***, measurements from fresh or pickled flowers—Continued

Taxon and

localityOvary

and neck Tube Tepals

Filament insertion

and length Anther

Total

length

Collection

No.

>

O

2

o

a

a

w

>

Ö

td

O

o

o

Ö

O

>

O

t—t

O

f

a

r23

21

)22 Sierra Jojoba (21

26

131

Sierra Baviso j 2]^

wocomahi

(27

Huicorichi, Chihuahua ) 29

(32

Sierra Charuco { 36

(32

(28

29

fp/?

25

27

Guajaráy Scarp j 25

Sierra Potrero j 09

Sierra Saguaribo { OQ

17X11

15X11

16X11

17X11

15X13

14X15

18X12

17X12

18

17

22 X 15

22X15

22 X 15

18

18

23

21

20 X 11

19X11

19X12

20X12

18

18

21X12

22X12

15X5&13

12 X 5 & 9

15 X 5 & 12

15 X 5 & 13

12X6&11

13X7&11

12 X 6.5

13 X 6.5

13 &11

19X5&17

23 X 5 & 18

23X5&21

22 X 5 & 20

15X5&13

14 X 5 & 13

10X6&8

10X6&8

13X7&11

13X7&11

11X7&10

12 X 7 & 10

12 X 5 & 10

11

13X7&11

12X7&11

7-8

7

8-9

8

7-8

7-8

8-9

7-8

11 & 9

11

13&10

13&10

14&11

ll&lO

10 & 8

14&10

13 & 9

12&10

11 & 9

9&8

10 & 9

10 & 8

10 & 8

10 & 8

11 & 9

46

42

50

50

43

43

40

50

55

55

62

60

60

65

63

50

45

58

55

50

40

50

50

40

40

25

20

23

25

21

23

21

21

22

13&14

21&12

20&14

21&13

25

26

14 & 8

13 & 9

26

25

16 (fell

16&11

14 & 9

16

15&12

59***

50***

55***

58***

g2***

52***

53***

62*

g4***

rrg***

65**

64**

62**

65***

g4***

62***

62***

65*

67***

Y][***

21987

21987

21987

21987

21987

21987

21989

21989

1989

1989

10252

10252

10252

6340

6340

2028

202,8

8080

8080

11582

11582

11378

11378

11633

11633
s. matapensis

Matapé

colorata

Aquibiquichij

Sierra Bojihuacame

huachucensis

Mount Huachuca{

Sonoita<

fortiflora

Sierra Jojoba

zebra

Sierra Viejo

subsimplex

Puerto Libertad

'20

2 15X11 12

X7& 10 9&8 36 11

&9 59*** 16600

,203 16X11 11X7&9 10

&8 45 20 51*** 16600

'20

2 15X11 11X7&10 10

&8 43 20 48*** 16600

|39

2 20X14 16X7&14 10

&8 52 30 77*** 11607

382 20X14 15

X7& 13 10

&9 56 28 75*** 11607

.31

3 19X14 16

X7& 14 10

&7 55 28 68*** 11607

24

5 16X14 16X5&14 8&6 48 26 61** 10270 18 4 16X14 17X5&15 9&7 42 16&10 54** 10270

23

4 13X14 13X7&11 6&4 50 22 51*** 11641 22 4 15X14 12 X 7 & 9 7&5 53 21 52*** 11641

24

3 15X14 12

X7& 10 7&5 54 21 53*** 11641

35

9

8 X16 26

X6& 24 5-7 45 27 78*** 22580

37 10

9 X17 27X5& 25 7-8 58 27 81*** 22580

31

6

8 X14 22X5&21 7-6 52 24 68*** 22581

35

6

8 X15 23

X5& 22

7 58 25 71*** 22581

33

8

9 X14 19X5&17

7 50 20 68*** 22582

33

78 X14 19X4&17

7 50 20 66*** 22582

34 11 11X18 20

X7& 19 8&6 60 20 75*** 11630 31 11 11X18 20X7&18 7&5 58 20 ^72*** 11630

33 12 12X18 19X7&18 8&6 60 20 75*** 11630

25

5

6 XIO 14

X 5.5& 133 35 17 50*** 21984

21

4

5 XIO 11X5.5

3 30 12 40*** 21984

20

4

6 X13 14

X5& 134 35 16 43*** 21984

28

4

6 X12 15

X5& 14 3-4 38 18 53*** 21984

19

54 X9 11X4 2-3 24

9 40* 4486

20

5

4 XIO 13X6

3 20 13 41** 4486

—

5

3 XIO 12X5

3 20 13 ** 4486

>><>OO>

CO

CO
TABLE 1.—Agave flower measurements (in mm,): *, measurements from dried flowers; **, measurements

CO

>

o

s

o

a

ir*

from dried flowers, relaxed by boiling; ***, measurements from fresh or pickled flowers—Continued

Taxon and Ovary Filament insertion Total Collection

> locality and neck Tube Tepals and length Anther length No.

deserti

Ö

C18 5 4 X15 14X6 5 31 13 40*** 19741 OQ

18 4 4 X15 14X6 5 31 13 40*** 19741 W

)l6 5 4 X15 13X6 5 30 13 38*** 19741 I2; San Felipe, Calif <

21 6 5 X15 16X8 5.5 23 15 47*** 19940 0

20 6 5 X15 16X8 6 28 15 48*** 19940 CO

V21 6 5 X15 16X8 6 26 14 53*** 19940 to

fourcroydes h

w

Í30

30

5 16X15 17X3 8 55 20 66* 6/8/09

Mórtola (Berger) 6 15X15 17 60 70* 6/8/09

pacifica

(20 2 6 X12 2C '; 4 5-3 45 20 48*** 19885

Felis Gomez \ 22 1 7 X12 1 X ^ 4 45 20 49*** 19885 0

(25 1 8 X12 18X4 4-5 45 52*** 19885 >

(27 3 11X14 19 X 5 & 18 5-6 40 13&11 61*** 19875 0

Navojoa 23 4 10X14 18X5 5-6 40 13&10 56*** 19875

(24 4 9 XIO 18X4 5-6 38 21 54*** 19875 0

f21 4 11X10 19X3 6 35 14&10 57*** 19359 tr"

I 21 4 12X10 20 X 4 & 18 6 35 24 58*** 19359 ci

l22 4 11X10 20 6 45 14&10 57*** 19359 w Sierra Tecurahui \25 3 14X13 24 X 5 & 22 10 45 17&13 67*** 19368 H

/25 3 15X13 24X5 10 45 16&13 67*** 19368

126 3 14X13 25X5 10 46 16&14 68*** 19368
Sonora-Sinaloa border I in

(2,5

Cerros del Fuerte l^^

rhodacantha

r20

Mazatan, Nayarit <|g

(28

9 Xll

9 XIO

8 XIO

13X10

11X10

14X10

10X9

9 X9

8 XIO

8 XIO

22X4

22X4

24X4

18X4

22 X 4 & 20

18X4

16X4

16X4

22X4

23X4

5-6

5

5

10

8

10

6-7

37

42

40

40

42

50

40

33

46

44

16&12

16&12

16&12

13&11

15&10

13&13

8&5

22

28

52***

52***

ß2***

63***

65***

54**

54**

56*

62*

19656

19656

19656

19655

19655

19655

10783

10783

10704

10806

w

>

>

<

t—I

O

o

>

00

en
36 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

The flower bud grows slowly but opens and expands very rapidly

during anthesis (expansion of the flower) within 1 day. Therefore, measurements for comparative purposes must be taken at

about the time of anther dehiscence, which occurs during a few

daylight hours. However, flowering begins at the base of the inflorescence and continues to move upwards in the inflorescence

for 2 to 4 weeks, depending upon individual plants and species.

One can usually find a series of flowers in late bud stage, at anther

dehiscence, and after dehiscence, all in one flower cluster. I prefer

to have measurements of flowers just before, during, and right

after stamen dehiscence so maturation is bracketed. For this purpose, I collect and immerse the flowers in a pickling solution of

1 part formaldehyde, 3 parts 95 percent alcohol, and 6 parts water.

Pint-size canning jars in cartons of one dozen are suitable for

field work and accommodate all except a few of the very largest

flowers. For permanent storage, they must be transferred to laboratory jars with noncorrosive resinoid tops. Labels are put inside

the jars.

The pistil continues to grow after anther dehiscence, usually

for 2 or 3 days, when the stigmas open and are receptive. Because

of this later growth, measurements of pistils at anther dehiscence

have no comparative value and have not been made.

Floral Ideographs

The measurements can be analytically applied to form the symbolic floral ideograph. The various forms of ideographs and their

applications were very ably described and applied by the late

Edgar Anderson (1), The form ideographs may take is more or

less determined by the organs, which the taxonomist finds best

representative of the variability that expresses speciation, introgression, or developmental features of a related group of organisms. The ideograph here selected shows the tube/tepal ratio,

together with the insertion of the filament within or on the rim

of the tube and, more rarely, on the base of the tepal. The ideograph is a kind of taxonomic shorthand that throws into perspective the selected characters, so strikingly apparent in figure 4,

A and B,

These floral ideographs are also employed on the maps to show

the known distributions of the respective agave species (figs. 13,

24, 31, 39, and 55). The ideographs are reduced in size and still
THE AGAVE FAMILY IN SONORA 37

function as meaningful symbols. Ideographs with a collection number below indicate that the ideograph is based upon a flower collection at that locality. Others without a number are typical or

mean species ideographs drawn from the measurements in table 1

or selected from the ideographs shown in figure 4, ß. The ideographs serve as specific representatives of documented localities.

Old dried flowers are shrunken and distorted by the growing

ovary and measurements of them are unreliable. They are little

improved by relaxing. Berger at La Mortola Botanical Garden

pressed split flowers under great pressure and, so-treated, their

proportions remain fairly reliable.

The junction of tube and tepals is a particularly critical level,

as the length of both organs derive from this. In measuring the

length or depth of tube, one must note the various tepal sinuses

because some may be deeper than others in the same flower. The

upper limit of tube is taken at the median level of the sinuses.

Where there is marked overlap of tepal bases in the sinuses, the

median point of the overlap is taken as the upper limit of tube.

There is, thus, a built-in indefiniteness about many tubes that may

amount to a 1-mm. variance or a margin of about 10-percent inexactness for tube measurement. Also, one must watch for damage

by insects and birds, which sometimes split the sinuses. With

measurements of three or four flowers, made after splitting the

flower lengthwise, these difficulties are largely overcome and the

mean or medium measurement is suitable for comparisons with

other collections.

Diameter of tube is its widest extent, usually at about the level

of filament insertion.

Width of outer tepal is the median width just above the basal

widening, or about Vs to i/^ tepal length above the tepal sinuses.

This is bound to be a general-impression kind of measurement

as it depends on degree of wilting, or degree of inrolling, and

cannot, therefore, always be precise.

Where there is marked difference in length of outer and inner

tepals, the length of the outer is given first. Width of the inner

tepal is not given, but its form, shape, or other qualifying characteristics are indicated in the description of species.

Where two measurements appear under anther, these indicate

the parts on each side of filament attachment. Total anther length

is equal to the sum of the two figures. Anther length is given in

this way only to indicate the degree of eccentric fixation. Dried

anthers swell more than any other organ when boiled in water.
38 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

SÜIEEIIUS tun

piliantliiflori

1013

sibsiinplti

j 44B6

FIGURE 4.—A, Long sections of agave flowers showing general floral organization and diversification. Scale is not uniform; A. parviflora and A. deserti only slightly reduced, others reduced from % to % natural size. The

diversity in agave flowers is symbolized in the ideographs on the opposite

page (fig. 4,5).
vilmorín.

ji19677

ocahui

16637

pelona

19898

multífl.

8167

17822

B

THE AGAVE FAMILY IN SONORA

bovicorn.

schottü

El

17751

toumey.

22307

parvifl.

D17752

flexífl.

u16638

políanth.

8013

palmerí

3672 17750

shrevei

8080

shrevei

11569

pacífica

11633

shrevei

17710 11378

shrevei

expansa

11582

m ata pe.

colorata

ñ 11641

wocomahi

snreveí .

^^^ murpheyi

desmett. ^^H H H

ü FI El H.B.G.

huachuc

19823

fortifl.

19808

zebra

39

pacifica

^19882 ñ21988í21177 ñ8080 U10254 Ù19875

rhodacan.

u10704

fourcroy. ú

Berger

0 1 I deserti

19741

subsimplex

21983 16600 21984 4486

B, Ideographs of agave flowers, which depict relative proportions of

organs derived by measurements, ratio of tube (black) to outer tepal

(white column), and insertion of ñlament in tube (black square). Measurements are listed in table 1. See also figures 3 and 4, A.
40 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

ANNOTATION OF SPECIES

Key to Genera of Agavaceae in Sonora

Ovary inferior ; perianth segments united to form a tube ; anthers

exserted or, rarely, included. (Agaveae)

Succulent multiannuals^ or, rarely, perennials, without fleshy

underground stems and succulent roots; leaves enduring for

many years, large, succulent, with conspicuous terminal spines ;

inflorescence a dense spike, raceme, or panicle; anthers exserted Aö^at;e

Perennial herbs with fleshy underground stems and succulent

roots; leaves enduring only 1 season or short-lived, relatively

small, weak, and without terminal spines; inflorescence of lax

spikes or racemes ; anthers exserted or included.

Filaments exserted ; flowers actinomorphic, the tube not amplíate above Manfreda

Filaments included; flowers zygomorphic, the tube amplíate

above Polianthes

Ovary superior; perianth segments free or nearly so; anthers

included.

Leaves with terminal spine ; flowers large (2 cm. wide or more),

usually succulent, perfect, campanulate ; capsules large, woody ;

seeds black, usually discoid, numerous in a cell. (Yuccae)

Perianth segments spreading, not connivent, broad, white;

filaments pubescent Yucca

Perianth segments narrow, connivent forming a tubelike

flower, variously colored; filaments glabrous . .. .Hesperaloe

Leaves without terminal spine; flowers small (less than 1 cm.

wide), not succulent, often unisexual, stellate; capsules small,

papery; seeds not black, never discoid, only 1-3 in a cell.

(Nolineae)

Leaf margins armed with sharp, curved spines; plants completely dioecious; capsules 1-celled, winged; seeds angled;

flowers in dense catkinlike spikes Dasylirion

Leaf margins entire or serrulate; plants incompletely dioecious, some flowers perfect; capsules 3-celled, 3-lobed; seeds

globose; flowers in diffuse panicles Nolina

^ Multiannuals are plants that flower once and die, but require several to

many years to mature.
THE AGAVE FAMILY IN SONORA 41

AGAVE L., SP. PL. 323. 1753.

Succulent rosettes, monocarpic or polycarpic, perennials or

multiannuals with long-lived leaves, frequently suckering at base

and occasionally bulbiferous in the inflorescence; roots hard

fibrous, radiately and shallov^ly deployed ; stems thick, very short,

usually shorter than the terminal bud, simple or branched ; leaves

large, generally succulent, spine-tipped, margin armed or unarmed

with teeth; inflorescence tall, bracteate, scapose, spikate, racemose, or paniculate with flowers in umbellate clusters; flowers

mostly large, generally proterandrous ; perianth tubular to shallowly funnelform, the 6 segments erect to variously curved, similar

or dimorphic, imbricate in the bud ; stamens 6 exserted ; filaments

long, inserted in tube or on tepal bases ; anthers versatile ; ovary

inferior, 3-celled, succulent, thick-walled with numerous axile

ovules in 2 rows ; pistil elongate, filiform, tubular ; stigma 3-lobate,

papillate glandular; fruit a dehiscent, loculicidal capsule; seeds

flattened, black.
42 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Key to Agave Species in Sonora

Inflorescence spicate or racemose Subgenus Littaea

Inflorescence paniculate, flowers in umbellate clusters on lateral

branches Subgenus Agave

Subgenus Littaea

1- Leaves filiferous, small or narrow, usually with white bud

printing on the surface; flower tube deep, narrow 2

1- Leaves not filiferous, the margins smooth, larger ; flower tube

very short, the tepals several times longer than the tube (except in pelona) 6

2- Rosettes large, (40-60 cm. tall), with numerous leaves (150

or more) ; leaves 40-50 cm. long; tepals greenish (section

Filiferae) multifiUfera, p. 46.

2- Rosettes small (20-30 cm. tall), with relatively few leaves

40-70) ; leaves 10-40 cm. long; flowers yellow or pink (section Parviflorae) 3

3- Flowers red or pink, long-tubular, the tube 25-30 mm. long,

7-10 times longer than the segments polianthiflora, p. 51.

3- Flowers yellow or yellowish green, the tubes much shorter,

3-10 mm. long, little longer or about equaling the segments __4

4- Flowers very small, 11-15 mm. long; leaves 10-20 cm. long,

linear-lanceolate parviflora, p. 54.

4- Flowers larger, 25-40 mm. long; leaves 20-40 cm. long,

linear 5

5- Flower tube narrow and deep, about equaling or exceeding

tepals. Mountain highlands schottii, p. 57.

5- Flower tube shallow, open, about l^ the length of the tepals.

Coastal lowlands ^^^felgeri, p. 60.

6- Leaves large, 1-2 m. long 7

6- Leaves small, 20-40 cm. long 8

7- Leaves deeply guttered, arching, 1.2-2 m. long; inflorescence

frequently bulbiferous; tepals linear-lanceolate, 4-6 mm.

wide vilmoriniana, p. 62.

7- Leaves plane, about 1 m. long, straight ; inflorescence not

bulbiferous; tepals ovate-lanceolate, the inner 8 mm. wide

chrysoglossa, p. 69.

8- Leaves rather pliant, green, with fine brown margins ; flowers

yellowish, short-pedicellate (5-10 mm. long) ocahui, p. 72.

8- Leaves rigid, purple-tinged and with conspicuous white margins; flowers red with a large tube, long-pedicellate (15-25

mm. long) pelona, p. 76.
THE AGAVE FAMILY IN SONORA 43

Subgenus Agave

1- Leaves sword-shaped, 10-25 times longer than wide, mostly

9-20 dm. long, straight, stiff, and always armed with teeth;

tepals wilting with anthesis, early drying sharply reflexed;

inflorescence frequently bulbiferous (section Ensiformae)

15

1- Leaves rarely sword-shaped, 3-10 times longer than wide,

variously curved, if stiff, then short and broad, sometimes

toothless ; tepals not normally wilting at anthesis and not drying sharply reflexed; inflorescence rarely bulbiferous except

in two species 2

2- Rosettes large, suckering prolifically; mature leaves 120-170

cm. long, broadly lanceolate; flower umbells diffuse and

broadly spreading (section Agavae, 1 cultivar in Sonora)

__ americana, p. 80.

2- Rosettes small to medium, usually suckering sparsely or not

at all; mature leaves less than 120 cm. long; flower umbells

compact 3

3- Rosettes not suckering; leaves green to yellowish green,

lanceolate to spatulate, frequently with interstitial teeth between prominent teats; panicles usually narrow, elongate,

with 25-40 laterals; tepals about twice as long as tube (section Crenatae) 4

3- Rosettes suckering, soon or late; leaves mostly glaucous gray

but sometimes green, linear to broadly lanceolate, if interstitial teeth present then not between prominent teats;

panicles less narrow and shorter, with mostly 10-20 laterals

(A. huachucensis up to 25-30) ; tepals shorter or much longer

than tube 5

4- Leaves broadly lanceolate to spatulate, about 4 times as long

as wide, strongly narrowed above base, margin markedly

crenate bovicornuta, p. 85.

4- Leaves narrowly lanceolate, about 6-10 times as long as wide,

only slightly narrowed above base, margin merely undulate

jaiboK, p. 89.

5- Leaves without teeth or teeth small and irregularly occurring,

tender and arching ; ovary about 1/2 ^s long as perianth ; tube

with prominent ridges within on each side of filaments;

tepals thin (section Sisalanae, 1 cultivar in Sonora).

desmettiana, p. 94.

5- Leaves usually armed with teeth (some species have toothless

forms), generally tough and stiffer than above; ovary and

perianth about equal in length ; tube without prominent ridges

within; tepals thick 6
44 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

6- Rosettes medium size; leaves lanceolate, mostly broadly so

and short acuminate, 30-90 cm. long; flowers with a welldeveloped tube ; tepals less than 3 times as long as tube ; ñlaments usually inserted well below orifice of tube 7

6- Rosettes small; leaves triangular-acuminate, 20-50 cm. long;

flowers with short open tube; tepals 3-5 times as long as

tube ; filaments inserted at orifice or on base of tepals 14

7- Tepals unequal, the outer longer and overlapping the inner,

about equaling or shorter than the tube, persistently erect;

filaments inserted unequally in midtube, those below outer

tepals conspicuously higher (section Ditepalae) 8

7- Tepals about equal, longer than the tube, not drying erect;

filaments inserted about equally in tube 12

8- Leaves linear or long-lanceolate, 5-10 times longer than wide,

widest at or below the middle ; margin straight or nearly so ;

teeth small to medium 9

8- Leaves ovate to lanceolate, 3-6 times longer than wide, widest

at or above the middle; margin undulate or prominently

teated under teeth; teeth frequently large and flexuous 10

9- Leaves linear; spine conical, short, up to 2 cm. long; flowers

about 70 mm. long ; inflorescence bulbiferous ^murpheyi, p. 99.

9- Leaves lanceolate; spine long acicular, 2.5-6 cm. long; flowers

40-60 mm. long; inflorescence rarely bulbiferous.

palmeri, p. 101.

10- Rosettes green, medium to large, not suckering; tepals brownish at tips, about equaling tube, sinuses open ; panicles large,

5-7 m. tall wocomahi, p. 105.

10- Rosettes glaucous gray, small to medium, suckering at base;

tepals red or brownish, usually shorter than the tube, sinuses

closed; panicles relatively small or short, usually less than

5 m. tall 11

11- Leaves uniformly gray; marginal teats lacking or moderate

in size; panicles tall, small, open with few laterals in upper

1/3 of shaft; tepals shorter than the deep tube ^shrevei, p. 111.

11- Leaves conspicuously cross-zoned and sometimes pink-tinged;

teats prominent; panicles deep, relatively dense with many

umbells in upper % of shaft; tepals about equaling the

tube colórala, p. 117.
THE AGAVE FAMILY IN SONORA 45

12- Rosettes compact with numerous (80-110), short, stiff, incurving, nearly plane leaves, not cross-zoned, smooth.

huachucensis, p. 119.

12- Rosettes open with few (40-70), ascending or arching, guttered leaves conspicuously cross-zoned and scabrous 13

13- Leaves straightly ascending, slightly guttered, relatively narrow, 5-8 times longer than wide; margins straight or with

small teats; flowers 70-80 mm. long, long-lasting; ñlaments

flattened, 4-5 mm. wide fortiflora, p. 122.

13- Leaves arching low, spreading, deeply guttered, relatively

broad, 3-5 times longer than wide; margins deeply crenate

with prominent teats and teeth; flowers about 50 mm. long;

filaments slender, oval in cross section, 2 mm. wide.

zebra, p. 126.

14- Plants 20-30 cm. tall; filaments inserted just below base of

tepals in orifice of tube ; tepals elliptic, broadest at the middle.

subsimplex, p. 131.

14- Plants 30-50 cm. tall; filaments inserted on base of tepals

above tube; tepals lanceolate, broadest at the base.

deserti, p. 134.

15- Rosettes developing a trunk 1-2 m. tall ; leaves uniform, 8-10

cm. wide ; teeth regular, mostly 2-4 cm. apart, small, less than

5 mm. long; panicles sterile (cultivar) fourcroydes, p. 138.

15- Rosettes acaulescent or with a short trunk; leaves and teeth

variable ; teeth generally longer than above, the larger along

midblades over 5 mm. long ; panicles bearing seeds ; capsules

large, ovate to oblong 16

16- Leaves mostly less than 150 cm. long; teeth proximal or remote; inflorescence 3-5 m. tall with fewer than 25 laterals;

capsules 3.5-5 cm. long, short-stipitate (less than 7 mm.) 17

16- Leaves mostly 150-200 cm. long; teeth proximal, strong; inflorescence 7-9 m. tall with 30-45 laterals; capsules 6-8 cm.

long, long-stipitate (10-15 mm.) rhodacantha,-p. 141.

17- Stems elongate; leaves variable, mostly 70-150 X 4-7 cm.,

yellowish green to light green, rarely glaucous; teeth proximal ; spine rounded to the base, gradually subulate.

pacífica, p. 143.

17- Stems globose; leaves regular, 40-60 X 2-3 cm. bluish

glaucous; teeth remote; spine broad and flattened above at

base, abruptly subulate aktites, p. 148.
46 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Subgenus Littaea

Agave miiltifilifera Gentry, sp. nov. Figure 5

Single, monocarpic, nonsurculose, green rosettes, about 1 m. tall ;

1.5 m. broad, with a short trunk at maturity and 200 or more

leaves; leaves 50-80 X 1.2-3.5 cm., linear-lanceolate, broadest

at base, erectly ascending to declined, long-filiferous, plane above,

slightly convex below, firm but pliant, smooth, light green, spine

1-1.5 cm. long, subulate, flattened above, castaneous to aging

gray; spike to 5 m. tall, 18-21 cm. thick with flowers, densely

flowered from level of upright upper leaves ; lower flowering bracts

about twice as long as flowers, grayish, subulate, the upper bracts

much shorter ; buds green with lavender luster ; flowers pale waxy

green with pink tinge on tepals, 40-43 mm. long; ovary 20-21

mm. long with constricted, faintly grooved neck 5 mm. long ; tube

5 mm. long, 10 mm. broad at apex, short-funnelform, the nectarious innerliner about filling the tube; tepals 16-17 mm. long,

subequal, narrowly elliptic, some erect, some recurving at anthesis,

apiculate, the papillate area broad and decurrent on the inner

tepals, the outer tepals narrower, 5-5.5 mm. wide, and also with

a low keel ; filaments 35-40 mm. long, slender, red, elliptic in cross

section, anthers yellow, 16-17 mm. long; pistil slender, whitish;

capsule 20-25 X 10-12 mm., slender ovate, apiculate; seeds small,

3.5-4.5 X 2.5-3 mm., the curved margin with a wrinkled or erose

winglike edge, hilum notch abrupt.

Type—Gentry 8167, north rim of Arroyo Hondo, Sierra Charuco. Chihuahua, live plants collected April 1948, flowered at Murrieta, Calif., June 10-July 18, 1966, deposited in U.S. Nati. Herbarium No. 2558493 and No. 2558494. Also at type locality, Gentry 102Í7, March 7, 1951, (live pi.) ; Sierra Guicorichi, Chihuahua,

Aug. 25, 1951, Gentry & Arguelles 11S99 (Iv. & live pi.) ; 20 km.

north of Puente de Basihuare (Rio Urique) along road from Creel

to La Bufa, Chihuahua, June 9, 1963, Feiger & Russell 8135 (Ivs.),

deciduous oaks. Its habitat is the rocky ledges and cliffs in the

Planta singulo brevicaulis; foliis viridibus lineari-lanceolatis, 50-80 cm.

longis 1.2-3 cm. latis, supra plana infra convexa, margine longo-griseo-filiferis; spina terminali parva 1 cm. longa, castanea vel grisea; scapo inflorescentia inclusa 5 m. alto, bracteis ad basim inflorescentibus 8-10 cm. longis

subulatis; floribus viridis pallidoroseis, 40-43 mm. longis; ovario 20-21 mm.

longo ad apicem constricto; tubo 4.5-5 mm. longo 9-10 mm. lato infundibuliformis; segmentis subequalibus 16-17 mm. longis 5-7 mm. latis, anguste

elliptis apiculatis; filamentis 35-40 mm. longis roséis ad apicem tubo insertis;

antheris luteis, 16-17 mm. longis; capsula parva 20-25 X 10-12 mm., angusti

ovata, apiculata; seminibus parvi 3.5-4.5 X 2.5-3 mm.
THE AGAVE FAMILY IN SONORA 47

FIGURE 5,—Agave multifilifera: Leaf, flower cluster, and capsules less than X

1/2; flower section and spine tip slightly enlarged; seeds X 2. Drawn from

Gentry 8167.
48 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

pine and oak forests elevations between 4,500 and 6,500 feet. The

species is known only from the northern Sierra Madre, the southern record being live plants in the Jardin Nacional, Mexico, collected in Durango. The photograph in figure 6 shows several plants

on brecciated lava in the scattered population on Sierra Charuco,

where the soil is too thin to support trees. This is very near the

Sonora-Chihuahua border.

The habit, filiferous leaves (fig. 6), and flower form place this

species in the section Filiferae, of which it is the largest known

member. The stems in cultivation reached about 2 feet in height.

Like other species of the Filiferae, the flower has a regular funnelform tube and recurving subequal tepals, and the filaments are

inserted on the rim or orifice of the tube. However, the tube of

A, multifilifera is considerably shorter in reference to the tepals

than that of other relatives. The short tube is like that of A. filifera, while the thinner flatter, leaves are more like those of A,

schidigera. Some of the size differences may be compared in the

following way :

Tube Tepals Leaves

Mm. Mm.

A. multifilifera 5 16-17 Thin, elongate.

A.filifera 5-8 14 Thick, short.

A, schidigera 8-10 13-16 Thin, short.

The flower proportions with the larger broader tepals, the green

flower color, the elongate leaves, and the large size of the rosettes

together with minor differences appear suflScient to separate this

northwestern group as a species. The bud printing is less distinct

than in most other filiferous agaves.

The name ''chahuiqui'' was applied to the plant on Sierra Guicorichi and is obviously of Indian origin, but I have no notes on

local uses. The normal lifespan of this species appears to be 18 to

25 years, as the young plants, assumed to be 3 to 5 years old when

collected in 1948, did not flower until 18 and 21 years later. In

1966 two plants at the Huntington Botanic Garden also flowered,

but starting in May, about 1 month earlier than my plant at Murrieta (fig. 7). These were probably from my 1951 collection. It

forms a handsome plant in the garden, the copious white threads

among the leaves being curiously attractive, and because of the

larger size would be more suitable than other Filiferae for landscaping grounds. It should be more cold hardy than other members

of the Filiferae, as it grows in sites of the higher Chihuahua

mountains where snow falls in winter. It showed no frost damage

during the 15 years in Murrieta where winter lows reached 24° F.
THE AGAVE FAMILY IN SONORA 49

.'r f -

V ?^';^-*f#^w . y* . ^

ji ' <-•••,ä'Hl .S!*2Sii t"«, du»,.*

: .Y*'*^í^t!^'r^i»5a¿^

PN-2108, PN-2109

FIGURE 6.—Agave multifilifera : Upper, on the bracciated lava of Sierra Charuco, the type locality, with one plant of A. vilmoriniana; lower, copious

threads in the rosette.
50 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

PN-2110

FIGURE 7.—Agave multifilifera in flower at Murrieta, Calif., July 1966; 18

years from a small transplant.
THE AGAVE FAMILY IN SONORA 51

Agave polianthiflora Gentry, sp. nov. Figure 8

Small, single or cespitóse rosettes 10-20 cm. high and 20-30 cm.

broad; leaves 10-20 X 1-1.3 cm., linear lanceolate, voidest in the

middle, plane above, convex below, green, v^hite bud-printed above

and below, the margin conspicuously white-filiferous and minutely

toothed near the base; spine 7-10 mm. long, grayish, weak; spike

12-20 dm. tall, stalk red, bearing flowers above the middle ; bracts

1-2 cm. long, lanceolate, subulate, chartaceous; flowers 40-45

mm. long, short-pedicellate, usually in pairs, or 1 or 3, at a node,

pruinose pink, long-tubular, subtended by scarious, triangular

bractlets 3-6 mm. long; ovary 9-12 mm. long, red, ovoid; tube

28- 32 mm. long, 7-8 mm. wide at the top, very narrow and curved

below; tepals reduced to triangular lobes of the tube, subequal,

3-4 mm. long; ñlaments, inserted 5-6 mm. above base of tube;

stamens short, reddish before anthesis, yellow afterwards, shortly

exserted ; pistils exserted before anthesis, slender ; capsules round

to ovoid, 10-15 mm. long, apiculate, pruinose, rugose; seeds black,

lucid, thick, 3.5-4 X 2.5-3 mm.

Type—Gentry 8013, high rocky rims around Arroyo Hondo,

Sierra Charuco, Chihuahua, Apr. 17-25, 1948, and cultivated at

Murrieta, Calif., flowering there in June 1957, deposited in U.S.

Nati. Herbarium No. 2549705. SONORA : Sierra Saguaribo, Dec.

26, 1951, Gentry 11S8Í (live pi. only) ; Canyon de la Escalera

(loop of the Bavispe River), June 23, 1940, White 2820 (If. & fl.) ;

Sierra Yecora, Feiger (live pi.). CHIHUAHUA : San Jose de Piñal,

Sept. 22, 1936, Gentry 2850 (capsules) ; Arroyo Hondo, Sierra

Charuco, Mar. 6-7, 1951, Gentry 10229; Majalca, alt. 6,000 feet,

June 24, 1936, LeSueur 572 (If. & fl.). It occurs infrequently in

small clones on rocky outcrops in the pine and oak forests at elevations between 4,000 and 6,500 feet from the Rio Bavispe south

on both sides of the Sonora-Chihuahua border. (Fig. 9, left,)

Planta parva, caule singulo vel caespitosulo ; foliis viridibus, diagonaliter

striati maculatis, albo-filiferis, lineari-lanceolatis, 8-18 cm. longis, 1-1.3 cm.

latis, parvi dentatis ad basim; spina terminali parva, grísea, 6-8 mm. longa;

scapo inflorescentia inclusa 12-20 dm. alto, vlrgato; bracteis l'-2 cm. longis

lanceolatis, subulatis, chartaceis; bracteolis minutis, 3-6 mm. longis, triangulari-acutis; floribus puniceis roséis, longi-tubiformibus, 40-45 mm. longis;

ovario brevi, ovato, 9-12 mm. longo; tubo 28-32 mm. longo, 7-8 mm.^diam., ad

basim aliquantulum curvato ; segmentis subaequalibus, 3-4 mm. longis, 3 mm.

latis, imbricatis; filamentis 30-32 mm. longis, roséis, intra tubo profunde insertis; antheris 6-8 mm. longis, post dehiscentiam luteis; capsulis ovatis, 10-

15 mm. longis, 8-11 mm. latis, apiculatis, rugosis; seminibus nigris, lucidis

3.5-4 X 2.5-3 mm. crassis.
52 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

FIGURE 8.—Agave polianthiflora : Spike and leaves X Vs; spine, flower cluster,

and flower section slightly enlarged. Drawn from Gentry 8013.

The long, tubular, red or pink flower with its very short tepal

lobes distinguishes this small agave from all others. The red pigment is made dusky with a pruinose overlay. The flowering shaft

is also reddish. The flowers closely resemble those of the genus

Polianthes, whence the specific name. Were it not for the agavoid

leaves and the exserted filaments, this plant could be placed in
THE AGAVE FAMILY IN SONORA 53

Polianthes. It is also unlike any other Agave I have observed in

not having a proterandrous flower, as the pistil in this plant is

exserted before anther dehiscence.

The leaves on the smaller rosettes of A. polianthiflora are

scarcely separable from those of A. parviflora (fig. 10). In moist

or more fertile situations and if watered frequently in cultivation,

the leaves become much larger than in its natural habitat. The

Feiger collection from Yecora has unusually broad coarse leaves.

In "Rio Mayo Plants" (13), A. polianthiflora was reported doubtfully as A. hartmanii S. Wats. Watson described that taxon from

a growing plant in the now extinct Cambridge Botanical Garden,

collected originally by Hartmen and Lloyd from an unspecified

locality in the northern Sierra Madre Occidental of Sonora or

PN-2111, PN-2112

FIGURE 9.—Left, Agave polianthiflora in flower at Murrieta, Calif.; right, A.

parviflora near type locality in Arizona.
54 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Chihuahua on the Lumholtz expedition of the 1890's. Because no

herbarium specimen was prepared and the disposition of the garden plant is unknown, it has not been possible to orient Watson's

species. Watson's brief description could apply to A. parviflora,

to A, polianthiflora, or to another small filiferous rosette related

to A. filifera Salm., all growing in the same region. These taxa

can be identified certainly only by their flowers. Wiggin's citation

of A. hartmanii {Í2), based on a collection near Matapé, is referrable to A, pariflora flexiflora. It, therefore, seems appropriate

to drop Watson's name as a nomen confitsum.

The Warihio Indian name for this plant is ''taiehcholi." They

reported it as a good source of sweet food when pit baked, and

the flowering stalks were formerly used as arrow shafts. The Rio

Mayo Mexicans refer to the small plants as ''mescalitos." The

species appears limited naturally to the higher elevations of the

mountains, 4,000 to 6,500 feet, growing on rocky sites in the open

pine and oak forests. It is not common, being found in small and

distant colonies. It makes a distinctive pot ornamental, but should

be watered very sparingly and given full sunlight, or the leaves

become etiolated, even floppy, and the rosette loses its diminutive

and attractive compactness. The filiferous leaves are green with

white diagonal brushlike markings formed by bud printing of the

white, waxy exúdate secreted in the growing bud.

Jacobsen (28) gives an illustration of this plant as A. parviflora.

Agave parviflora Terr. ssp. parviflora Figure 10

Agave parviflora Torr., United States and Mexico Boundary Bot. 214. 1859.

Very small, single or cespitóse rosettes 10-15 cm. high and 15-

20 cm. broad; leaves 6-10 X 0.8-1 cm., oblong-linear, widest at

or above the middle, plane above, convex below, green, white budprinted above and below, the margin conspicuously white-filiferous

and minutely toothed near the base; spine weak-subulate, 5-8

mm. long, brown to grayish white; spike 10-18 dm. tall, laxly

flowered through upper half of shaft, which is frequently reddish ;

bracts with lowest flowers 1-3 cm. long, much smaller above,

scarious, long-subulate from a broad deltoid base, ephemeral;

flowers pale yellow, in 2's, 3's or 4's, 13-15 mm. long; ovary proper

4-5 mm. long ; neck 2 mm. long ; tube 5 mm. long, urceolate ; tepals

2-3 mm. long, the outer slightly longer than the inner, the latter

much broader than long, 1.5 X 3-4 mm. and abruptly narrowed
THE AGAVE FAMILY IN SONOEA 55

PARVIFLORA

16638

FIGURE 10.—The similar leaves of three agaves, v^hich are certainly distinguishable as three taxa only by their flowers. A. parviflora flexiflora has

two leaf forms.

below in the tepal sinus, all erect to incurved ; filaments erect 10-

12 mm. long, inserted at base of tube; anthers 5-6 mm. long;

capsules orbicular to oblong, 6-10 mm. in diameter, sessile to short

pedicellate, shortly beaked ; seeds half-round, black, wedge-shaped,
56 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

3 mm. long on the thin flat edge, 2.5 mm. in diameter, much thickened on the curved side.

Type—Collected by Schott in the 'Tajarito Mountains, Sonora,"

which was located by Trelease {J^S) as near the international

boundary marker No. 129, between Sonora and Arizona, 10-15

miles west of Nogales. Locally, these mountains are now referred

to as the 'Teña Blanca Mountains" and on some roadmaps are

shown as the ''Atasco Mountains." Peebles IIUUU and Gentry

10022 and 17752, about 11 miles southeast of Ruby, Ariz., are on

this same mountain and topotypic (ñg. 9, right). More recently

A. parviflora was found 16 miles northeast of Nogales along Arroyo Sonoita, Arizona, Gentry and Kaiser 19900, Apr. 10, 1963,

alt. about 4,000 feet on rocky open slope in oak woodland. SONORA : 18.3 miles west of Moctezuma along road to Mazochahui,

alt. about 3,000 feet, July 18, 1960 Feiger 3Í92 (If. & fl.) ; "lower

edge of oaks on N-facing slopes, inflo, ca. 6-10 ft. high." Inconspicuous, the plant has been overlooked by collectors and probably

has a much denser distribution than indicated by the limited

collections.

^ It flowers in June at the height of the spring dry season. Occasionally the inflorescence becomes diffusely racemose with numerous, strictly ascending, slender, lateral branches 5-10 cm. long.

With A, pumila Baker, it is among the smallest of agaves.

Agave parviflora subsp. flexiflora Gentry, ssp. nov.

Small, mostly single rosettes with filiferous, dimorphic, white

bud-printed leaves; leaves linear or lanceolate, 6-10 X 1 cm. or

15-18 X 1.2 cm., plane above, convex below, short-acuminate, the

margin minutely toothed near the base ; spine small, weak, whitish, 5-8 mm. long; spikes slender, 15-25 dm. tall, laxly flowered,

with small, triangular, scarious, caducous bracts; flowers 1-3 at

a node, mostly geminate, short-pedicellate, yellow, saccate, 14-17

mm. long, the corolla with anthers and pistil flexed downward at

anthesis; ovary 6-8 mm. long, fusiform; tube 3-4 mm. long; tepals

3.5-5 mm. long, broadly lanceolate, obtuse the inner sharply

^ Planta parva, caule singulo vel caespitoso; foliis viridibus, striati-maculatis, albo-filiferis, lineari-lanceolatis, 10-18 cm. longis, 1-1.2 cm. latis, paucidentatis ad basim; spina terminali parva, grisea, 5-8 mm. longa; scapo

inflorescentia inclusa 15-25 dm. alto, virgato; bracteis chartaceis, caducis,

brevi-triangulatis; floribus reflexi-saccatis, luteis; ovario 6-8 mm. longo,

luteo-viridi, fusiformo; tubo 3-4 mm. longo; segmentis subaequalibus, luteis,

3.5-5 mm. longis, ovati-lanceolatis, obtusis; filamentis 14-16 mm. longis, ad

basim tubi insertis; antheris luteis, 5-6 mm. longis; capsula et semina non

vidi.
THE AGAVE FAMILY IN SONORA 57

keeled and with broad hyaline margins, slightly shorter than the

outer; filaments 14-16 mm. long, inserted in base of tube; anthers

5-6 mm. long, yellow ; capsules and seeds not seen.

Type—Gentry 16638, collected 15 miles east of Guasabas along

road to Huachinera, Sonora, Mexico, May 20, 1957, on volcanic

rocks in oak woodland, alt. about 4,000 feet, deposited in U.S.

Nati. Herbarium, No. 2549710. SONORA: 6-8 miles north of

Bacadéhuachi, May 21, 1957, Gentry 166Í3 (ñ.) ; Ures of Moctezuma, Gentry 16655 (live pi.) ; 3-4 miles northeast of Matapé in

savanilla, alt. 2,000-2,500 feet. May 13, 1957, Gentry 16601 (buds)

and July 2, 1959, Gentry 177ii (fl.) ; western slope of Sierra

Batuc, 5 miles north of Matapé, Sept. 8, 1941, Wiggins & Rollins

il3 (If. & cap.) ; 9.3 miles northwest of Nacori Chico, alt. about

3,450 feet, June 6, 1960, Feiger & Marshall 3305 (fl. & If.), rocky

low hills, grassland. The habitat is the short, sparse, open, grama

grasslands with gravelly clay soils derived from volcanics at elevations between 2,000 and 4,500 feet.

A, parviflora flexiflora differs from A. parviflora parviflora in

the downward flexed flowers, in the longer tepals, and in the tendency to develop longer leaves (fig. 10). This last character is conspicuous in the collections east of Guasabas, but is seldom apparent

in the collections west of the Bavispe and Moctezuma Rivers, in

which the leaves are not separable from typical A, parviflora. The

downward flexing of the flower develops with anthesis, becoming

marked with the downward flexure of the growing pistil. Carpenter bees enter the flower from below the perianth.

Local names for these diminutive agaves were reported as **tauta,"

''tautilla," and "sóbari'' or "sobali.'' They were said to be among

the sweetest and most edible of their kind when properly cooked,

but were rarely used because of their small size. All forms of the

species are attractive as pot ornamentals. Carpenter bees were

working the flowers assiduously in the populations east of Guasabas at the time of specimen collections.

Agave schottii Engelm., Acad. Sei. St. Louis Trans. 3: 307. 1875.

Figure 11

Densely cespitóse, small, yellowish green to green rosettes;

leaves narrowly linear, 0.7-1.2 X 25-40 cm., widest at the base,

straight, incurved, or falcate, pliant, flat or somewhat convex

above, deeply convex below, smooth above and below, margins

with a narrow brown border and sparse brittle threads; spine

8-12 mm. long, grayish, fine, rather weak and brittle; spikes 1.8-

2.5 m. tall, slender, frequently crooked, flowering in upper Vs to
58 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

14 of shaft ; bracts straw-colored, filiform, acicular, 2-4 cm. long ;

flowers yellow 1, 2, or 3 on stout pedicels 3-5 mm. long with setaceous bracteoles 10-15 mm. long on lower spike but shorter above;

ovary greenish yellow, 8-10 mm. long with neck 4-6 mm. long;

tube deeply funnelform, 9-12 mm. long, very narrow below; tepals

10-16 mm. long, yellow, unequally spreading at anthesis, the outer

without a keel; filaments 15-20 mm. long, inserted high in tube

at 6-9 mm. above base of tube ; anthers light yellow, 10-15 mm.

long; capsule 10-20 mm. long, rounded to apiculate; seeds 3-3.5

mm. along the straighter thinner edge, with a notch at one corner,

variably thickened.

Type—Arthur Schott in 1855, Sierra del Pajarito in southern

Arizona, flowering in August. Also collected on the same mountain

is Gentry 17751, about 12 miles west of Nogales along road to

Ruby, flowering in July 1959. SONORA: Western foothills of

Sierra de la Caballera, about 10 miles east of Colonia Morelos,

alt. 3,400 feet, Sept. 22, 1941, White U66 (If. & fr.). Between

Santa Rosa canyon and Bavispe, July 20, 1938, White 628 (If. &

fl.) ; about 16 miles southeast of Magdalena along road to Cucurpe

by Sierra Baviso, May 22, 1960, Feiger 3276 (If. & fl.) ; Rancho

Primavera, west of Sierra Jojoba, July 24, 1966, Gentry & Weher

21988 (fl.). The following leaf collections are tentatively assigned

to this species. SONORA: Picú Pass, 18-20 miles northeast of

Libertad, MacDougal & Shreve H; Wiggins 6097; Port Libertad,

MacDougal & Shreve s,n. In northeastern Sonora, colonies of this

species were observed on the lower slopes of Sierra Huachinera

near the Chihuahua border. The distribution in Chihuahua is unknown. It is probably common in the oak and grama grassland

highlands of northern Sonora at elevations between 4,000 and

6,000 feet, as in adjacent Arizona and New Mexico. It is less common about the desert mountains of northwestern Sonora, where

it has been observed along arroyos and outwash bajadas at elevations between 1,500 and 2,500 feet. The western limit appears

to be in the low Picú Mountains some 15 to 20 miles east of Libertad. (Fig. 11.)

The slender, deep tube, together with the narrow, ovarian neck,

gives the flower a characteristically long, tubular appearance. The

ratio of tube length to tepal length, however, is variable, but the

tube usually equals and sometimes exceeds the tepals in length.

In gardens, A. schottii is easily confused with narrow-leaved forms

of A, falcata and A. felgeri, but all three are readily separated by

their respective flowers.
THE AGAVE FAMILY IN SONORA 59

PN-2113, PN-2114

FIGURE 11.—Upper, Agave schottii with Yucca peninsularis and Fouquieria

splendens in pass by Sierra Baviso, 16 miles southeast of Magdalena;

lower, A. schottii at road fork north of El Datil.
60 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Among Spanish-speaking people these small, narrow-leaved,

nondescript agaves are knov^n as "amole" or ''amoliyo'' and they

are employed locally for v^ashing clothes. They are not eaten because of the insignificant size of the ''heads'' (stems) or because

of bitter constitutents. The leaves v^ere found to contain 0.5-1.2

percent sapogenins, mainly gitogenin and chlorogenin. The plants

have little if any value as ornamentals. They are so abundant on

some cattle ranges in southeastern Arizona that cattlemen regard

them v^ith disfavor. However, they contribute a great deal towards

building and holding soil on the steep, rocky slopes they inhabit.

Agave felgeri Gentry, sp. nov. Figure 12

Small, green to yellow-green, surculose rosettes with rather few

leaves forming rather closely cespitóse clones; leaves 25-35 X

.7-1.5 cm., linear to narrowly lanceolate, straight or.falcate, plane

above, convex below, widest at the base, faintly bud-printed and

frequently with a pale median stripe, epidermis rugose or scabrous

above, the margin with a narrow brown border weekly filiferous,

smooth; spine small, weak, 8-15 mm. long, gray; spikes 1.5-2.5 m.

tall, flowering in upper 14 of shaft ; bracts chartaceous, the lower

ones 2-3 cm. long, very narrow; pedicels strong, single or bifurcate, 2-5 mm. long; old dry flowers (relaxed) 25-30 mm. long;

ovary 12-14 mm. long; tube 2-4 mm. deep; tepals 10-12 mm. long,

about equal, linear; filaments 20-25 mm. long inserted in orifice

of tube ; capsules oblong or obovoid, narrowed towards the base,

apiculate, 15-20 X 9-12 mm. ; seeds irregular, 4-5 X 3 mm., thick,

angulate, wrinkled, the hilar notch narrow and deep.

Type—Geniri/ 118J^S Bahia San Carlos, District Guaymas, Sonora, Dec. 13, 1951 (If. & cap.) with coastal shrub on volcanic

agglomerate (fig. 12), deposited in U.S. Nati. Herbarium No.

2540143 and No. 2540144. Same locality: Nov. 7, 1954, Feiger

650 (If. & cap.) and at later dates. Feiger 2Jf92; Feiger, Kleine,

& Russell 1U56; Felger, Thomas, & Russell 12121, SONORA:

Planta rhizomati-caespitosa, parva, 20-30 cm. alta, 30-40 cm. lata; foliis

paucis, pallido-viridibus, saepe medio pallido-lutis-striatibus, linearibus, 25-35

X .7-1.5 cm; asperibus sparse brunneofiliferis, planus supra convexus infra;

spina terminali parva, 8-15 mm. longa, grísea; scapo inflorescentia inclusa

1.5-2.5 m. alto, virgato; bracteis inferioris 2-3 cm. longis, chartaceis; bracteolis 5-8 mm. longis; floris 25-30 mm. longis; ovario 12-14 mm. longo; tubo

2-4 mm. longo; segmentis aequalibus 10-12 mm. longis, linearis; filimentis

20-25 mm. longis, ad apicem tubi insertis; capsulis oblongis vel obovatis,

apiculatis, 15-20 X 9-12 mm.; seminibus 4-5 X 3 mm. crassis rugosis angulatis.
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PN-2115, PN-2116

FIGURE 12.—Broad-leaved form of Agave felgeri at San Carlos Bay. Spikes (right) were carried to beach for photographing. 05
62 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Rancho Bachaca, 15-18 miles east of Navojoa, Dec. 19, 1951, Gentry 11366 (fl. & cap.) ; Sept. 16, 1959, Gentry 17822 (If. & dry fl.).

Collections from near Libertad in northern Sonora, D. T, McDougal in 1923 and Wiggins 6097, Oct. 26, 1932, may be assignable to this species, and if so, indicate a narrow coastal range

through middle Sonora. I have seen A. felgeri for sure only in

the thin parched rocky soil on volcanic ledges near the coast.

In habit, size, and leaf form this proposed species resembles A.

schottii, especially in the forms having narrow leaves, as those

at the Rancho Bachaca. However, the short tube with the relatively long tepals, 3-4 times as long as the tube, indicate a basic

difference between the highland A. schottii and the coastal A. felgeri. Fresh flowers are needed to judge the relationships. Its flowering season appears to be June and July, but the San Carlos Bay

population certainly does not flower every year, as several trips

to that easily accessible locality have shown. (Fig. 12.)

At Rancho Bachaca (Mayo, meaning bad water) the people

called this plant ''mescalito" and said they did not use it. I found

it there in one isolated colony upon one of the bald volcanic domes

that characterize the place. Analyses of the rosettes yielded about

1 percent sapogenins, that from Rancho Bachaca consisting wholly

of chlorogenin, while that from the Bahia San Carlos material

contained 67 percent chlorogenin and 33 percent tigogenin.

It is a pleasure to commemorate the name of Richard Feiger

by giving his name to this species, in recognition of his sustained

interest in this Agave and of his years of labor with the Sonoran

biota as a graduate student of the University of Arizona.

For distribution of Agave multifilifera and the Parviflorae

group, see figure 13.

Agave vilmoriniana Berger, in Fedde's, Repert. Spec. Nov. Regni

Veg. 12: 503.1913. Figure 14

Agave mayoensis Gentry, Rio Mayo Plants, Carnegie Inst. Wash. Pub. 527,

p. 94. 1942.

Short-caulescent, Hght green to yellowish green, single rosettes,

1 m. tall X 2 m. broad with toothless, arching, deeply guttered,

pliant leaves, 90-180 X 7-10 cm., broadest at base, linear-lanceolate, long acuminate, heavily thickened towards the base, concave

to conduplicate above; margin unarmed with a fine brown continuous border about 1 mm. wide, scaly in age; epidermis smooth;

spine acicular, brown to grayish brown, 1-2 cm. long ; spike 3-5

m. tall, densely flowered from 1-2 m. above base, bulbiferous or
THE AGAVE FAMILY IN SONORA 63

FIGURE 13.—Distribution of Agave multifilifera and the Parviflorae group in

Sonora, represented by their ideographs. Ideographs with numbers indicate flower collection at the locality occupied. (See p. 37.)

nonbulbiferous ; lower bracts 10-20 cm. long, scarious, acicular,

brownish towards the brittle tip; upper bracts 1-2 cm. long; bracteoles of the pedicels deltoid, 2-3 mm. long; pedicels 8-20 mm.

long, 1-2 bifurcate ; flowers yellow, 35-40 mm. long with a shallow

open tube; ovary 15-20 mm. long including neck (3-4 mm.) ; tube

4 mm. long x 8-9 mm. wide; tepals 14r-17 X 4-5 mm., nearly

plane, ascending to spreading at anthesis, clasping the filaments
64 AGRICULTURE HANDBOOK NO. 399. U.S. DEPT. OF AGRICULTURE

FIGURE 14.—Agave vilmoriniana: Lower and upper thirds of leaf (spine tip

enlarged); flower cluster from lower part of spike about X %; flower

section slightly enlarged. Drawn from Gentry 10232. Upper right, A,

chrysoglossa: Flower section slightly enlarged. Drawn from Gentry

19882.
THE AGAVE FAMILY IN SONORA 65

in drying, equal ; filaments 30-40 mm. long, inserted at upper edge

of tube ; anthers 16 mm. long.

Type—None cited. Berger reported it was collected in Mexico

by Diguet. In an undated letter to J. N. Rose, fide copy in U.S.

Nati. Herbarium, Berger wrote that the plant was collected by

"Mr. Diguet for the Jardin des Plantes at Paris and which I named

Agave vilmoriniana in 1913 in honour of M. Maurice de Vilmorin,

an excellent man, in whose garden at Verreves I found this Agave

for the first time. . . . The fruits in your photo resemble greatly

those of A. ellemeetiana etc. of the same group *Anoplagave.' It

was not in cultivation at La Mortola and of course if I v/ere still

there I should be delighted to receive your plants." Rose had found

the plant on the rim of the barranca near Guadalajara, Jalisco,

July 9, 1899, Rose & Hough ^833, Hough having risked his life

to obtain the specimen. As Diguet is known to have collected in

the vicinity of Guadalajara, it is probable that this barranca is

the type locality. There are two spines and a pencil sketch by

Berger in the U.S. Nati. Herbarium. The flowers are described

and shown here for the first time (figs. 14 and 15). SONORA:

Arroyo Gochico, Rio Mayo, canyon cliffs in barranca forest, Apr.

5, 1938, Gentry 3673 (fl.) ; also in Arroyo Gochico, 5 miles above

San Bernardo, May 1, 1957, Gentry 16531 (with bulbils & fl.) ;

Rancho San Antonio, Sierra Tecurahui, May 20, 1966, Barclay &

Arguelles 2016 (fl. & If.). ''Abundant on bare rocky cliffs; pineoak zone.'' CHIHUAHUA: High rocky rims and cliffs around

Arroyo Hondo, Sierra Charuco, alt. 5,000-5,500 feet, March 6-7,

1951, Gentry 10232 (If. only). DURANGO: 8-10 miles south of

Boca de Mezquital, along road from Durango to Mezquital, June

11, 1951, Gentry & Gilly 10587 (If.). (Figs. 16 and 17.)

A cliffdweller, the typical habitat of this handsome plant is the

volcanic brecciated cliffs of the barrancas from southern Sonora

south through Sinaloa and Durango to Jalisco and Aguascalientes,

at altitudes between 2,000 and 5,500 feet. In the larger, deeper

canyons it forms extensive vertical colonies, which, when viewed

from a distance, resemble giant spiders on a wall. Individuals

may be found upon gentler adjacent slopes in open light. The cliff

situations are virtually frostless, but the open heights above 4,000

feet have sharp frosts. In southern California the plants have

resisted overnight freezes as low as 24° F. They withstand protracted drought easily and will take in water during the chilling

winter rains in California. Many of the cliff plants, for lack of

soil, remain stunted through their lives, their maturation inhibited,

and the inflorescence is feeble when at last it does appear.
66 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

PN-2117, PN-2118

FiGXniE 15.—Left, Agave vilmoriniana: Bulbils growing out of

flowering axils; right, A. chrysoglossa: seeding spike.
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fruiting spikes.

Dried flowers hang, bleached and

PN-2119

beardlike, on

OS
68 AGRICULTURE HANDBOOK NO. 399. U.S. DEPT. OF AGRICULTURE

PN-2120, PN-2121

FIGURE 17.—Agave vilmormiana : Upper, plants 1 year old from bulbils at

Huntington Botanical Garden; lower, maturing plant on Sierra Charuco,

Sonora. Compare these deeply guttered leaves with those of A. chrysoglossa (fig. 18).
THE AGAVE FAMILY IN SONORA 69

This species is distinguished by its unarmed, large, gracefully

arching, deeply guttered leaves (fig. 17, lower). In Sonora and

elsewhere it is known as ''amole," and, like other plants of the

name, is used in washing clothes and other fabrics. For this use,

the old dried leaf bases on dead flowered plants are cut off about

6 or 8 inches above the base. The fibers in the lower butt are beaten

and loosened by pounding with a rock, forming an effective brush

with a built-in soap or detergent, which foams and cleans in water

as one brushes. These brushes have been noted for sale in local

markets of the Mexican west coast. The cleaning agent is principally the sapogenin, smilagenin, which by analyses was found to

form 3-4.5 percent of the dry leaf, among the highest known for

any agave leaf. Younger green leaves contain much less. For remarks on the commercial potential, see ''Uses of Agave and

Yucca,'' p. 9.

As an ornamental A. vilmoriniana lends itself well to design

plantings and propagates readily from bulbils. The clone in the

Huntington Botanical Garden in San Marino, Calif., forms mature

flowering plants in 7 or 8 years from bulbils (fig. 17, upper).

However, young plants collected on Sierra Charuco, Gentry 10232,

and planted at Murrieta in 1951 did not flower until 1966—15

years later. The spike emerged soon after April 1 and reached

about its full height in early May. The more rapid maturation in

the Huntington Botanical Garden is apparently due to better culturing there and, perhaps, also to genetic early maturation factors. Capsules and seeds are also produced, hence, a given population may consist of both apomictic and sexual generations. This

was noted in the Arroyo Gochico population (fig. 16). No rhizomatous offsets have ever been observed.

Agave chrysoglossa IM. Jtn., Calif. Acad. Sei. Proc, ser. 4, 12:

998. 1924.

Single, short-stemmed, openly spreading, few-leaved, green

rosettes, 1-1.3 X 2-2.4 m. ; leaves 70-120 X 4-7 cm., wider at the

base, straight or slightly curved, flat above, convex below, deflexed

at maturity, light green, smooth, linear-lanceolate, toothless, with

a thin brown fragile margin 1 mm. wide; spine 2-4 cm. long,

acicular, brown, aging grayish, with a short fine groove at base

above; spike mostly 2-4 m. tall, densely flowered in upper %;

bracts dry, chartaceous, the lower ones exceeding the flowers;

pedicels bifurcate, 10-15 mm. long, their bracteoles small, whitepapery, curling; flowers yellow, 35-40 mm. long, gemminate;

ovary 16-18 mm. long including neck 3-5 mm. long, slender,
70 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

scarcely grooved; tube shallow, 4-4.5 mm. deep, 9 mm. wide; tepals

ovate, plane, outcurving at anthesis, clasping filaments in drying,

thin, the outer 14-16 X 6 mm., keeled, the inner wider (8 mm.)

and unkeeled; filaments 30-40 mm. long, inserted on the rim of

tube at base of tepals; anthers 15 mm. long; capsules mostly oblong, 2x1 cm., short-apiculate, parallel-veined and slightly crossridged; seeds lunate, mostly 4-4.3 X 2.5-2.8 mm., 1-several wrinkles on faces, marginal wing low, firm, the hilar notch sharply

angled {Gentry 16611),

Type—/. M, Johnston 3123, found flowering Apr. 17, 1921, on

the rocky, granitic slopes of San Pedro Nolasco Island, Gulf of

California; Gentry 113Í9, same locality, Dec. 16, 1951, without

flowers. The following collections document it for the mainland.

SONORA : 26 miles northwest of Sahuaripa, along the Rio Yaqui,

May 24, 1957, Gentry 16611, (cap.) ; 14 miles south of Bacanora

along road to Tonichi, May 16, 1957, Gentry 16623, (fl.) ; cerro

4-5 miles north of Bahia San Carlos, March 29, 1963, Gentry

19882, (fl.) ; San Pedro Bay, July 7, 1921, Johnston Í338, ^^common

on rocky mountain side'' ; about 2 miles northwest of Bahia San

Carlos along road to Bahia Algodones, riparian canyon with desert

shrub and thorn scrub, Feb. 9, 1960, Feiger & Stronck 3113.

A few plants once observed in a lower canyon of Sierra de Alamos

may belong to this species.

A, chrysoglossa is closely related to A. vilmoriniana, but the

straight, narrow, plane leaves and nonbulbiferous inflorescence

are sufficient to distinguish A. chrysoglossa, (fig. 18). A. chrysoglossa is the coastal or lowland relative of this alliance in Sonora,

more xerophytic and at home upon the hot, frequently bare rock

surfaces. It flowers from mid-March to mid-May during the spring

dry season when air temperatures during the day usually rise to

100-110° F. I found the inland population about Bacanora in May

to have prolific sets of maturing black seeds (fig. 15, right). Estimates based on partial seed and capsule counts indicated as many

as 500,000 to 750,000 seeds per plant. It appears to be a freeseeding, cross-pollinated species, with the liquid nectar attracting

animals during the thirsty season. Numerous insects, hummingbirds, and other kinds of birds were observed about the flowers.

Johnston wrote of the island plants (loc. cit.), *When found it

was enlivening the rocky mid-slopes of the island with spectacular

tongues of color. The plants grew singly and produced dense elegant spicate floral clusters 1-2 m. long and 8-10 cm. broad which,

due to their weight, almost invariably bent over with their tips

nearly touching the ground." All mainland plants observed had

erect spikes.
THE AGAVE FAMILY IN SONORA

PN-2122, PN-2123

FIGURE 18.—Agave chrysoglossa: Sections of flowering spike and a typical

plant, showing the flat upper surface of leaves, at San Carlos Bay, Sonora.
72 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

This Agave, also known as '^amole," is almost as sapogenous as

A, vilmoriniana and has the same local use for washing clothes.

Small plants were transplanted from Bancanora to my Murrieta

garden, where they have done well, but in 10 years they have not

flowered.

Agave ocahui Gentry, sp. nav. Figure 19

Short-caulescent, single, dense, green, yuccoid rosettes, 30-50

cm. tall, 50-100 cm. broad ; leaves numerous, 25-70 cm. long, 1.5-2.5

cm. broad, widest at the base, linear-lanceolate, plane above, convex below, mostly stiff, erect to ascending, some older declined

or falcate, the margin straight and lined with a very narrow,

reddish-brown, firm border detachable on dried leaves, surface

smooth, minutely, densely punctulate in fine lines; teeth none;

spine weak, rather brittle, 1-2 cm. long, pruinose gray over brown ;

spike about 3 m. tall, slender, densely flowered from 1-1.5 m. above

the base ; flowers 30-35 mm. long, yellow, on short bifurcate pedicels; ovary 14-15 mm. long with short constricted neck; tube about

4 mm. deep, openly spreading; tepals subequal, 14-15 mm. long,

5 mm. wide, oblong, the tips outcurving; filaments 40 mm. long,

yellow, inserted on base of tepals; anthers 14-15 mm. long, yellow

(measurements from relaxed dried flowers) ; capsules small, 12-15

mm. long, 7-8 mm. broad, ovoid, apiculate, yellowish tan to reddish, tardily dehiscent; seeds small, 2.5-3 mm. long, 1.5-1.7 mm.

wide, rather thick, half-ovate.

Type—Gentry & Arguelles 16637, deep volcanic canyon above

(east of) Guasabas, Sonora, May 21, 1957, alt. about 4,000 feet,

deposited in U.S. Nati. Herbarium No. 2549703 and No. 2549704.

SONORA : Sierra Baviso, 16 miles southeast of Magdalena, near

Canyon de Palmas, March 31, 1963, Gentry & Arguelles 19886

(fr.); Ahuaje de Pescado, 8 miles east of Matapé, Gentry &

Arguelles 16603 (If. for analysis), on rock cliff, the leaves green.

Planta brevicaulis, yuccoidea, 30-50 cm. alta, 50-100 cm. lata; foliis multis,

25-70 cm. longis, 1.5-2.5 cm. latis, lineari-lanceolatis, firmis, aliquando curvatis aut falcatis, supra plana, infra convexa; margine inermi, firmo, angusto,

castaneo, continuato; spina terminali 1-2 cm. larga, grisea, fragili; scapo

inflorescentia inclusa ca. 3 m. alto; pedicellis 5-8 mm. longis, bracteolis chartaceis, 5-20 mm. longis, filiformibus; floribus luteis, 30-35 mm. longis, densis;

ovario 14-15 mm. longo, viridi, cylindrico; tubo ca. 4 mm. longo; segmentis

subequalibus, 14-15 mm. longis, 5 mm. latis, oblongis, recurvatis; filamentis

40 mm. longis, ad basim segmenti insertis; antheris luteis, 14-15 mm. longis;

capsula parva, 12-15 mm. longa, 7-8 mm. lata, ovata, apiculata; seminibus

parvi, 2.5-3 mm. longis, 1.5-1.7 mm. latis, crassis, nigris, margine curvato

angusto-alatis.
THE AGAVE FAMILY IN SONORA 73

FIGURE 19.—Agave ocahui: Leaves and spike section X Vs; flower pair X 1;

flower section slightly enlarged; cuticular pattern X 20. Drawn from

Gentry 16637.
74 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

recurved and weeping; 6-8 miles northeast of Matapé, Sierra

Batuc, Gentry & Arguelles 16607 (live pi. & seed) ; about 4 miles

w^est of Sahuaripa, alt. about 1,500-1,800 feet on volcanic rocks,

Gentry & Arguelles 16612 (live pi.) ; Sierra Murrieta, 15 miles

v^est of Bacanora, in Oak Woodland, alt. about 4,500 feet. May 17,

1957, Gentry & Arguelles 16626; southern rim of Canyon Cruz de

Peñasco, 7 miles east of Rio Bavispe along road to Bacadéhuachi,

alt. about 3,650 feet, June 5, 1960, Feiger 3301 (If., fl., & cap.),

''Lvs. green v^ith white margin; fls. yellow; inflo, ca. 8 ft. tall."

A. ocahui is known only in northeastern Sonora, where it occurs

on cliffs and other outcrops of volcanic rocks at elevations between

1,500 and 4,500 feet. Also noted as ''scattered on limestone crest

of the Murrieta Mountain'' (Gentry notes). Such sites commonly

have dry, poor soils inhibiting growth. Plants found on deeper

soils and in partial tree shade develop longer, more pliant leaves.

(Figs. 20 and 21.)

The smooth, narrow leaves with fine, brown, unarmed margins,

the prolifically flowering spike with its small, slender, yellow

flowers with shallow tube, and the high insertion of filaments, all

relate this species with the other Sonoran amóles, A. vilmoriniana

and A. chrysoglossa. The leaves and flowers of A. ocahui were

found to contain 0.5-1.0 percent smilagenin, which also relates

it to the amóles. It is easily distinguished from the others by its

small rosette with more numerous, but smaller leaves and by its

smaller flowers with small stamens on long filaments. Also, in

contrast to the other Sonoran amóles that have fine, weak leaf

fibers, the leaf fiber of A. ocahui is coarse, abundant, strong, and

employed locally for cordage and brushes.

Neither the chemistry nor the fibers of A. ocahui were assayed

sufficiently to assess the prospects of this plant as a potential resource. Considering the fiber-sapogenin industry of the cultivated,

but frost-susceptible fiber agaves, A. ocahui, because of its apparent cold hardiness, its superior fibers, and good genin content,

merits investigation for cultivation in the United States. The

length of generation is not known, nor are its cultural requirements except as generally apply to agaves. It seeds abundantly

and all the plants observed in the several populations visited were

nonsurculose and showed relatively little variation. The considerable variation in size of rosette and leaf length appeared correlative to site situations. The small plants in the Murrieta collection

have grown very slowly since they were planted in 1957. The generative period is probably very long. The flowering season is not

definitely known. Although the type plant was flowering in May,
THE AGAVE FAMILY IN SONORA 75

PN-2124, PN-2126

FIGURE 20.—Agave ocahui: On tuffaceous rocks on sun slope of Sierra Baviso,

Sonora.
76 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

^ #0 » ' •

^^^Txmf}

PN-2126

FIGURE 21.—Agave ocahui: Section of flowering spike on volcanic rocks above

Guasabas, Sonora.

repeated visits to other populations during May, June, September,

and December found no flowering. (Fig. 20.)

"Ocahui," "ojajui," and "amoliyo" are local names for this

agave. The first two terms are of Indian origin and signify fiber

or cordage. A related term "majahui" carries the same meaning

and is used by country people in Sinaloa and southeastward, bu1

it is applied to such bast fibers as those from the Malvaceae anc

Sterculiaceae and to Ceiba.

Agave pelona Gentry, sp. nov. Figure 22

Acaulescent, single, dark green to purplish, shiny, multileaved,

toothless, compact rosettes 40-50 cm. tall, 50-80 cm. broad ; leaves

Planta monocarpica, acaulescens, 40-50 cm. alta, 50-80 cm. lata; foliis

35-50 X 3-4.5 cm., lineari-lanceolatis, longi-acuminatis, rigidis, margine

inermo, albo, firmo, 2 mm. lato; spina terminali valida, alba, 4-7 cm. longa,

supra plana, vel canaliculata infra angulata; inflorescentia racemosa, scapo

incluso 2-3 m. alta, 12-15 cm. lata; bracteis filamentosis; floribus rubidis,

45-50 mm. longis, longi-pedicellatis ; ovario cylindrico, viridi, apice incluso ca.

20 mm. longo; tubo 8-9 mm. longo, 17 mm. lato, infundibuliformi; segmentis

subaequalibus, 18 mm. longis, 8-10 mm. latis, relexis, apiculatis; filamentis

35-40 mm. longis, rubris ad apicem tubi insertis; antheris rubris, 15-17 mm.

longis, capsulis 25-30 mm. longis, 12-15 mm. latis, oblongis, valvulis tenuibus;

seminibus 4x4 vel 5x5 mm., rugosis, nigris.
THE AGAVE FAMILY IN SONORA 77

FIGURE 22.—Agave pelona: Leaf with spine offset of ventral side, flower pair,

and capsules X V2 ; flower section slightly enlarged with apex of central

tepal raised; section of leaf cuticle and margin greatly enlarged. Drawn

from Gentry 19898.
78 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

35-50 X 3-4.5 cm., linear-lanceolate, long acuminate, sometimes

slightly narrowed towards the base, nearly plane above, rounded

below, thick, stiff, erect to ascending, shiny dark green, turning

reddish to purplish with drought or maturity; epidermis smooth,

waxy, minutely circle-punctate ; margin toothless, with a narrow,

smooth, white, firm border becoming brittle and detachable on

dried specimens ; spine strong, sharp, 4-7 cm. long, white to reddish, sharply angled below, grooved or plane above, decurrent as

a white border down the leaf; inflorescence racemose, 2-3 m. tall,

flowering through upper half of shaft; flowers 45-50 mm. long,

dark red, campanulate, on bifurcate pedicels 30-40 mm. long;

ovary including neck about 20 mm. long, cylindric, slender, light

greeji, faintly striate; tube 8-9 mm. long, 17 mm. wide, openly

funnelform, lined with a nectariferous disk; tepals 18 mm. long,

8-10 mm. wide, linear-ovate, plane and unkeeled, the inner widest,

8-9 nerved, recurved at the tip; tip apiculate, white-pappillate;

filaments 35-40 mm. long, red, inserted on rim of tube; anthers

red, 15-17 mm. long, centrally affixed ; pistil small ; capsules 25-30

mm. long, 12-15 mm. broad, oblong, light pruinose, thin-walled,

rounded and short-apiculate ; seeds irregular in size and shape,

mostly 4 X 4 to 5 X 5 mm., thick, rugose, sooty black, numerous.

Type—Gentry & Arguelles 19898, Cerro Quituni, about 26 miles

south of Caborca, Sonora, April 7, 1963, on arid limestone mountain, alt. 1,500-3,000 feet, deposited in U.S. Nati. Herbarium, No.

2549712 and No. 2549713. Also on Cerro Quituni, May 20, 1966,

Barclay & Arguelles 2023 (If.). SONORA: Cerro del Viejo, about

25 miles south of Caborca, alt. 3,500-4,000 feet, April 5, 1963,

Gentry & Arguelles 19896 (moth-eaten buds) and Feb. 25, 195l'

Gentry 10202 (If.) ; Sierrita de Lopez, about 35 miles northwest

of Hermosillo, Feb. 19, 1952, Gentry 11615 (If.). It was reported

by miners to grow on Sierra del Humo, but this is doubtful. The

largest population known grows on the precipitous limestone

heights of the 25-mile-long Sierra (or Cerro) del Viejo. (Fig. 23.)

This beautiful and distinct, endemic xerophyte has no close relative. The rosettes cling firmly to the rough limestone rocks and

cliffs. The rigid, dark-green leaves, often tinged brightly with red

and purple, are strikingly outlined by the white, smooth margins

and the long, strong, white spine. The long-pedicellate, dark-red

flowers with spreading recurved tepals are distinct in the genus

The funnelform tube with its thickened, nectareous inner lining

with 6-pointed apex of filaments is like the section Filiferae, but

the leaf is more like agaves of the amole group or the section

Margmatae. However, both leaf and inflorescence of A. pelona
THE AGAVE FAMILY IN SONORA

i

PN-2127, PN-2128

FIGURE 23.—Agave pelona: Upper, old dry plant with seeded spike; lower,

live growing rosette in natural habitat on Sierra del Viejo.
80 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

show some similarities to A, utahensis. One tooth on one leaf was

observed on one plant.

Flowering appears to be normally during April, but in dry years

flowering is sparce, or not at all. In the dry spring of 1963, all of

the several inflorescences found on Sierra del Viejo had the buds

completely gutted by an insect, assumed to be a Lepidopteran

larva. The flowers on the one flowering plant found on Cerro

Quituni the same season were being eaten by a rodent. Several of

the buds were collected and placed in a small water jar. These

opened the following day and it is from these that my flower

measurements and description and Regina Hughes' drawings are

taken.

I was first guided to this agave by a ''mescalero'' encountered

along the streets of Caborca, who called this plant ''mescal pelon,''

in reference to the bald or toothless leaf margins. He stated that

it was a sweet kind and suitable for making the ''moonshine"

called "mescal,'' except its "cabezas" (heads) were too small for

the labor involved in cutting and trimming. For this reason, he

preferred the larger "mescal lechuguilla" in the same region, described on page 126 as A. zebra. The mescaleros of Mexico have

frequently been very helpful in my investigations of agaves. Presumably, A, pelona was formerly eaten and its strong fibers were

used by the local Amerindians, such as the Papagos and Seris who

still inhabit adjacent areas, but there are no reports of uses other

than those I have already given. Analyses of the leaves for steroids showed 0.06 percent sapogenin from Sierra del Viejo and 0.3

percent smilagenin from Sierrita de Lopez.

For the dry, rocky, succulent gardens of the Southwest, A.

pelona should make an attractive addition, but should be placed

where children cannot fall against its strong, stiff spines. Both

leaves and flowers are strongly colored. The lifespan is not known,

but must be many years.

For distribution of the Amolae group of agaves, see figure 24.

Subgenus Agave

Agave americana L., var. expansa (Jacobi) Gentry, slat. nov.

Agave expansa Jacobi, Nachtrage I in Abh. Schles. Ges. Vaterl. Cult., Abth.

Naturwiss. 1868: 151. 1868.

Large, light-gray rosettes, 1.5-2 m. tall, 2 m. broad, with a

short trunk in age, suckering abundantly from early age ; leaves

12-15 dm. long, 18-24 cm. wide, lanceolate to spatulate, rather

abruptly acuminate, narrowed towards the base, deeply rounded
THE AGAVE FAMILY IN SONORA 81

X%

"iß"'!. S Pajarilo ÏÏ

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w,,# SOIOBA

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—28° T j^ 1 "C \ ^£/"^ Jl^AYMA^g- %% ^/ \ \ r-|^X_^^

1^5 S Bojihuacame \ .asLnTs Choruco

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IM37 19898 110°

FIGURE 24.—Distribution of the Amolae group of agaves in Sonora, represented by their ideographs. Ideographs with numbers indicate flower collection at the locality occupied. (See p. 37.)

below, the upper % of leaf more plane, firm, thick, erect or ascending, frequently cross-zoned, sometimes valleculate, the margin

nearly straight, usually with several sharply angled, low teats in

the midblade ; the larger teeth 5-8 mm. high, 1-4 cm. apart, shortly

cuspidate from low broad bases, brown becoming gray; spine

short, conical, 2-3 cm. long, narrowly grooved above, brown to

gray; panicle large, 7-9 m. tall, deeply oval in outline, with 20-
82 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

30 long, horizontal laterals; bracts of the peduncle 10-15 cm. long,

triangular, chartaceous, reflexed; flowers thick and fleshy, 70-85

mm. long ; ovary green, short, tapering from unconstricted neck ;

tube green, 13-14 mm. deep, 12-20 mm. broad, scarcely bulging,

but deeply grooved and thick-walled ; tepals pale yellow, erect,

thick, 26-32 mm. long, 7-9 mm. broad, linear, involute, cuculate,

the outer 2-3 mm. longer, the inner with thick keel, 2 strong

costae within, and thin involute margin; filaments stout 65-70

mm. long, flattened on adaxial side and bowed towards pistil, inserted 8-9 mm. above tube base ; anthers yellow, 28-32 mm. long,

excentric ; capsules and seeds not seen.

This variety is known only as a cultivar, introduced in western

Europe, where it was described by Jacobi as growing at ''St. Germain en Laye bei Paris,'' France. What is taken for this taxon

has been observed from southwestern United States to Jalisco,

where it probably originated. It was collected at Amado, Ariz.,

along the route to Nogales, July 23, 1966, Gentry & Weher 21988,

from which the present flower description is drawn (fig. 25),

Tumacacori, Ariz., Nov. 6, 1950, Gentry & Ogden 99W (If.). SONORA : Rancho Monte Carlo, about 30 miles west of Hermosillo,

Gentry 11631 (If. for analysis). It has been observed around the

Sonoran towns. Nogales and Hermosillo.

It has taken botanists a century since this plant was first described to collect and describe the flowers, a rather shocking delinquency because the plant flowers in both the Mediterranean

and the Southwestern United States regions almost every year.

Wiggins H2) treats A. expansa as a synonym of A. americana.

Indeed, the large, glaucous gray, prolifically suckering rosettes

and tall, diffuse panicle resemble the type of the species and genus

in its Mediterranean form. Trelease (43) regarded the Arizona

form as a separate species, distinguishable by its short, heavy,

conic spine and unreflexed leaves vs. the long subulate spine and

reflexed leaves of A. americana. Until 1963 only a few, incomplete

herbarium specimens of A. americana were available in American

herbaria and only two leaf specimens and one poor, withered

flower collection of A. expansa. The recent flower collection (Gentry & Weber 21983) confirms the close relationship of A. expansa

to A. americana. The flowers of A. americana, both in typical form

and in the numerous variants found wild and cultivated in northeastern Mexico, are characterized by: a short, tapering ovary,

shorter than the perianth; large, thick-fleshy, elongate outer

tepals ; smaller inner tepals with thick keels, thin involute margins,

and paired prominent inner costae; and long, strong filaments
THE AGAVE FAMILY IN SONOEA 83

PN-2129

FIGURE 25.—Agave americana expansa at Amado, Ariz., as ornamental hedge,

well developed with irrigation. Source of Gentry & Weber 21983.

divergent above the tepals. The species forms a highly varied

complex, both in flov^ers and leaves, as represented in w^ild and

cultivated populations of Mexico. The western relative is treated

here as a cultivated variety.

The origin of A. americana L. has never been known, Linnaeus

listing only "America calidiore." (So. PI. 1: 323, 1753.) A form

very close to the Mediterranean form Linnaeus had in mind grows

along the lower Rio Grande around Laredo. South of there, in
84 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

eastern Mexico, are several variant forms in cultivation. I suspect

that some port like the Rio Soto la Marina in northern Tamaulipas,

which v^as among the earliest sheltered harbors along the eastern

Agave coast, may v^ell have been the source area of A. americana.

The Portuguese carried A, americana to the Azores in the 16th

century. From the Azores it was carried to Europe (Linnaeus).

Although they are not of the European or Laredo clone, the collection of several flowering variants in eastern Mexico during the

summer of 1963 greatly expand the dimensions of this species.

The polymorphic populations from southeastern Texas to Hidalgo

enjoin complex relationships awaiting detailed studies. The maguey or metl-culturing Amerindians apparently collected the fortuitous and attractive larger variants from local wild populations.

These they propagated by their sucker offsets and several of these

clones have diffused widely during the thousands of years of man's

cultivation.

The east Mexican A. americana americana appear separable

from the west Mexican A. americana expansa on the basis of leaf

characters, as follows :

Leaves long-acuminate, recurved to reflexed; larger teeth

longer than (or as long as) their corneous bases are wide,

not on sharply raised teats; spines slender, 3-5 cm. long—

var. americana

Leaves short-acuminate, straight, erectly ascending; larger

teeth shorter than their broad, low, corneous bases are wide,

raised on teats, some of which are abruptly angled with leaf

edge ; spines short, thick, conical, 2-3 cm. long—var. expansa

Some of the varieties of A. americana in Jalisco are used for

pulque and as fence plants; one of these, A. abrupta Trel., is a

form with a thick, short spine on a valleculate short-acuminate

leaf. In Sonora, A. americana expansa has only desultory use as

an ornamental. The indigenous peoples there did not drink pulque.

It's sporadic occurrence in and about the modern towns and its

scarcity in older Indian villages indicate that A. americana expansa was brought northward to Sonora, Arizona, and California

in modern times, probably after 1850. It is very common now in

southern Arizona and California, where it is used as an ornamental, as a hedge plant, and for holding soil on banks. However,

its prolific suckering makes it weedy and the untended bunches

become untidy and troublesome in time. If the suckers are controlled, it makes a good and hardy ornamental in our Southwestern States.
THE AGAVE FAMILY IN SONORA 85

Agave bovicornuta Gentry, Carnegie Inst. Wash. Pub. 527, p. 92.

1942. Figure 26

Medium to large, single, nonsuckering, light green rosettes, 8-10

dm. tall, 15-20 dm. wide; leaves 60-100 X 14-20 cm., lanceolate

FIGURE 26.—Agave bovicomuta: Leaf and flowers drawn from Barclay and

Arguelles 2017, the flower section about natural size; capsules X Vs and

seeds X 1.5 from Gentry 21175.
86 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

to spatulate, widest at or above the middle, much narrowed towards the base, yellowish green to green, the younger leaves

frequently satiny glaucous, smooth, bud-printed, the margins crenate with prominent teats under the larger teeth ; teeth dimorphic,

the larger mostly 8-12 mm. long, flexuous and slender above a

broad base, mostly 2-4 cm. apart, the smaller mostly 2-5 mm. long,

1-several between the larger, all castaneous or dark brown to grayish brown in age ; panicles 5-7 m. tall, narrow and deep with 20-

30 short laterals in upper half of shaft ; bracts below the laterals

20-30 cm. long, triangular-acuminate, soon drying reflexed, those

above smaller; flowers small in compact umbels, greenish yellow,

55-65 mm. long, on pedicels 5-10 mm. long; ovary pale green,

30-35 mm. long including neck 4-6 mm. long ; tube 6-8 mm. deep,

12-14 mm. wide; tepals yellow, 18-21 mm. long, 4-6 mm. wide,

linear-lanceolate, ascending-spreading, conduplicate, involute,

broadly overlapping sinuses, the inner wider and 2-costate within ;

filaments yellow, 40-45 mm. long, inserted 4-5 mm. above base of

tube; anthers 20-23 mm. long, yellow; capsule stipitate, 40-50

mm. long, 15-20 mm. in diameter, oblong, the valves thin; seeds

7x5 mm., finely punctate, the curved side with a flange or wing,

the hylar notch shallow.

Type—Curohui, Sierra Saguaribo, Rio Mayo, Apr. 4, 1938, Gentry 3672. SONORA : Sierra Tecurahui, Dist. de Alamos, November 1961, Gentry, Barclay, & Arguelles (photos); Sierra de

Alamos, Mar. 14, 1910, Rose, Standley, & Russell 12781 (If., fl., &

cap.). La Tinaja, north side of Sierra de Alamos, riparian canyon,

Feb. 20, 1954, Feiger 376 (If. & fl.) ^^common on rock walls of

canyon, flowers bright yellow, only plant found in flower, inflo,

to 8 feet"; vicinity of Rancho San Antonio, Sierra Tecurahui,

May 20, 1966, Barclay & Arguelles 2017 (If. & fl.), "abundant on

slopes in pine-oak woods." CHIHUAHUA: High rocky rims

around Arroyo Hondo, Sierra Charuco, Mar. 6-7, 1951, Gentry

10236. SINALOA: Sierra Surotato, Dec. 13, 1959, Gentry &

Arguelles 18379 (fl. only). Generally it occurs at elevations between 3,000 and 6,000 feet on the rocky open slopes in oak woodland and the pine-oak forest zones. Although common and widespread in the northern Sierra Madre Occidental, the populations

are usually local with limited numbers of individuals, and one may

travel for many mountain miles without seeing any. (Fig. 27.)

A. bovicornuta is quite clearly of the polymorphic group represented by A. inaequidens Koch and several other later names in

Berger's sections Scolymoides and Americanae. Trelease and

others have used Koch's name, A. mescal (1865), for this complex.
THE AGAVE FAMILY IN SONORA 87

JßK^

PN-2130, PN-2131

FIGURE 27.—Agave bovicornuta: Upper, on volcanic rocky open slope in the

oak woodland on Sierra Tecurahui, Sonora; lower, detail of teeth and

bud printing on leaf backs.
88 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

However, Berger dropped the taxon and I find Koch never diagnosed it, remarking about it only as a handsome plant observed

in tv^o or three gardens. Koch's A. inaequidens (1860) appears

as the oldest diagnosed name applicable to this group. A. mescal

should be dropped as a nomen nudum. These plants are recognizable by their nonsuckering habit, their yellov^ish to glaucous green,

lanceolate, smooth leaves with crenate margins, their hig teeth

on large teats with smaller interstitial teeth, and their tall, narrow

panicles with numerous short, lateral, compact umbels of greenishyellow flowers. Some individual plants may not have the small

interstitial teeth, but all populations observed exhibit them in the

majority of individuals. Some teeth arrangements are bizarre,

as seen in figure 27, lower. The section Crenatae occurs throughout the Sierra Madre Occidental south of the Rio Yaqui watershed

and across the trans-Mexican range from Colima to Morelos and

if A. seemanniana is placed with them, on into Central America.

Materials collected to date indicate a large polymorphic group,

which with more collections and study may be best systematized

as geographic subspecies. For the time being, A. bovicornuta as

a species is separable from its southern congeners by its small

flowers, 45-55 mm. vs. 60-90 mm., and its winter-spring flowering vs. fall flowering.

In Sonora the only species with which A. bovicornuta may be

confused is A. wocomahi, which is also nonsuckering and may

show occasional interstitial teeth. However, the leaves of A. wocomahi are darker green, not yellowish green, and the teeth are

longer and deflexed downward from the mid-blade below. On the

basis of flower morphology, I place the two species in separate

sections; the tepals of A. bovicornuta (section Crenatae) are twice

as long as the relatively short tube, whereas the tepals of A. wocomahi (section Ditepalae) are about equal to the deep tube in larger

flowers, 70-85 mm. long ; compare respective ideographs ; fig. 4, B,

page 39. Although flowers are seldom available for field identifications, they are very helpful in separating the species when they

are available. These species live sympatrically with A. shrevei,

and no morphological intergrades have been observed that indicate

hybridizing.

Some Rio Mayo people may call both A. bovicornuta and A.

wocomahi "lechuguilla verde," but the mescal makers and the

Indians differentiate between the two and call the latter '*wocomahi." A. wocomahi is employed for distilling the drink mescal

and is also eaten, whereas A. bovicornuta is usually employed for

neither. The latter is probably one of the agaves reported by Ben-
THE AGAVE FAMILY IN SONORA 89

nett and Zingg H) and again by Pennington {38) as used for

poisoning fish. The juice is caustic and produces a temporary

dermatitis on tender skin, with a burning sensation, inñammation,

and white welts or blisters. Some people are more susceptible than

others. Washing with soap only increases the burning. It is better

to wipe dry or wash with clear water only. A. bovicornuta was

among many samples of section Crenatae analyzed for sapogenins ;

like the others, it contained little or no sapogenin. The constituents

toxic to fish and skin have not been identified chemically. On the

Sierra Surotato, Sinaloa, I was given the nam.e "noriba'' for A,

bovicornuta,

A. bovicornuta forms a handsome plant in the succulent garden,

especially the more glaucous individuals, where a satiny sheen

appears on the new leaves along with crenulated bud printing.

Because they do not sucker or form bulbils and apparently require

cross-pollination, garden stocks are exhausted in one generation,

12-18 years, unless new field collections are made. The fertile black

seeds germinate readily and can easily be cultured in flats or pots

during early stages.

Agave jaiboli, sp. nov. Figure 28

Medium-size, single, nonsuckering, green to yellowish green,

usually open rosettes, 6-10 dm. tall, 14-20 dm. broad ; leaves 60-

100 X 8-12 cm. linear to lanceolate, widest at or above the middle,

gradually narrowed below, usually straightly ascending to spreading, long acuminate, sometimes incurved, plane to conduplicate,

the margins noncorneous or narrowly corneous with the decurrent

spine for less than its length; the larger teeth mostly 2-3 cm.

apart, on small regular teats, 5-8 mm. long, flexed downwards or

upwards, reddish brown, the smaller interstitial teeth 1-several,

1-4 mm. long; terminal spine 3-4 cm. long, subulate, terete, reddish brown, shiny, openly or narrowly grooved in lower 1/2 to %

Planta non-surculosa 6-10 dm. alta, 14-20 dm. lata; foliis viridibus, 60-100

cm. longis, 8-12 cm. latis, lanceolatibus, rectis-adscendentibus ; dentis

castaneis, dimorphis, majoris 5-8 mm. longis, flexuosis, in mammae 2-3 cm.

distantis, minoris 1-4 mm, longis; spina terminalis 3-4 cm. longa, castanea,

subulata, supre canaliculata, teres, decurrenta; inflorescentia paniculata, 6-8

m. alta, diffusa, cum ca. 15 ramii lateralibus; floris parvis, 35-50 mm. longis,

g-racilis; ovario parvo, fusiformo, 20-30 mm. longo, 8 mm. lato; tubo 4-5 mm.

longo, 8 mm. lato; segmentis 15-17 mm. longis, 4-5 mm. latis, linearis, involutis, ad apicem rotundis, galeatis, filamentis 35-40 mm. longis ad médius

tubo insertis; antheris luteis, 13-14 mm. longis; capsulis oblongis, 40-50 X

18-20 mm., stipitatis; semini 5x7 mm., negris, luteis, marginis alatis.
90 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

FIGURE 28.—Agave jaiboli: Leaf drawn from Arguelles 78; flower cluster and

capsules X Vs, flower section X 1, and seeds X IV2 from Gentry 21177.

The flower cluster was reconstructed from old withered flowers persisting

on the thickened and still growing ovaries or young fruits.
THE AGAVE FAMILY IN SONORA 91

of length, smooth; panicle 6-8 m. tall, the laterals about 15 in

upper 1/3 or 1/2 of shaft, ascending and rather sigmoid, with small

diffuse clusters of flowers on long (1-2 cm.) bifurcate pedicels

with discoid apices in fruit; flowers small, slender, 35-50 mm.

long; ovary fusiform, 20-30 mm. long including 3-4 mm. neck;

tube shallow, 4-5 mm. deep, 8 mm. broad, bulging at filament insertions; tepals yellow, 15-17 mm. long, 4-5 mm. wide, subequal,

linear, rounded at tip and glandular floccose, involute, ascending,

persisting erect, the outer broader, the inner strongly keeled on

back and with 2 prominent ridges within that unite in apex ; filaments yellow, 35-40 mm. long; anthers yellow, short 13-14 mm.

long (flower description from relaxed dried flowers from inflorescence tip broken off by animal) ; capsules mostly 40-50 X 18-20

mm., long-stipitate, shortly beaked, light brown ; seeds 5x7 mm.,

shiny black, very finely punctate, the straight edge as well as the

rounded ones, with a wavy wing.

Type—Gentry 21177, Sierra de la Ventana, Rio Mayo, Sonora,

Mexico, May 27, 1965, near Sonora-Chihuahua border, in oaknolina grassland at about 3,500-foot altitude, deposited in U.S.

Nati. Herbarium No. 2549714 and No. 2549715. Same locality.

Gentry 21186, May 28, 1965 (fig. 29). SONORA: 2 miles northwest of Conejos along Arroyo Guajaráy, north of San Bernardo,

Dec. 22-25, 1951, Gentry 1137Í (If. only) ; near San Bernardo, Rio

Mayo, Mar. 8, 1951, Gentry 10256 (If. only) ; vicinity of San Bernardo, May 1959, Arguelles 78 (If. only). The species occurs

sparsely as widely scattered individuals between 1,500- and 3,500-

foot elevations on the more open slopes of the short-tree forest and

the grasslands of the lower parts of oak woodland. The open,

rocky, grassy slopes in the Guajaráy country appear to be the

center of area and habitat (fig. 30). Only one to a dozen or so

individuals have been observed in one locality, all as single, nonsuckering rosettes. (Figs. 29 and 30.)

The flower structure and dimorphic teeth on the soft, fleshy

leaves place A. jaiboli in section Crenatae. The narrow, almost

ensiform leaves and the small flowers with shallow tubes distinguish this species from its relatives. It is further distinguished

from its Sonoran relative A, bovicornuta by the unnarrowed haft

of the leaves so characteristic of the latter. The natives have no

difliculty in separating ^'jaiboli'' or ^'temeshi,*' Warihio Indian

names, from the unpalatable A. bovicornuta, Jaiboli is much

esteemed for its sweet edible qualities, so the two species have a

quite different chemical makeup. Chemical analyses showed neither

of the species to contain appreciable quantities of sapogenins.
PN-2132, PN-2133

FiGXJRE 29.—Agave jaiboli on Sierra de la Ventana: Left, tip of panicle was broken off (by a large bird?), caught on oak

branches at fall, and provided dried flowers for Gentry 21177; riffht, unusual plant with incurving leaves.

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THE AGAVE FAMILY IN SONORA 93

PN-2134

FIGURE 30.—Agave jaiboli with budding inflorescence in December near Conejos along Arroyo Guajaráy. Leaf form here is more typical of species

than that shown in ñgure 29.
94 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Both the young flowering shoots and the headlike stem of jaiboli

are cooked and eaten by the peoples of the Rio Mayo country;

the succulent flowering shoots are usually boiled, the heads pit

baked over hot stones and coals. The roving eye of the man with

a ready machete is quick to discover a new shoot and it takes

but a moment to cut and lay it upon the shoulder to carry home.

The species appears to become ever rarer, as it reproduces only

by seeds, and my search for flowering specimens has been frustrated during the many years I have known the plant. The nonIndian people, "gente de razón,'' living on Sierra de la Ventana

frequently decapitate the flowering shoots, so the rosette will remain green and mature until such time that it can be conveniently

collected for making mescal. This is a common practice in many

parts of Mexico.

The young plants are easily cultured and have grown well in

my Murrieta collection, showing no ill effects of the 5 to 8 degrees

of frost occurring there. However, rabbits and gophers are fond

of the plants and completely destroyed three of the plants introduced there. The cottontail rabbit, Sylvilagus, ate the leaves of

half-grown plants clear down to the base. The food qualities of

this plant appear to merit detailed investigation as a potential

resource.

For distribution of seven species of the subgenus Agave, see

figure 31.

Agave desmettiana Jacobi, Hamburger Garten-BIumenzeitung

22:217.1866. Figure 32

Agave regeliana Jacobi, Hamburger Garten-BIumenzeitung 22: 214. 1866.

Agave miradorensis Jacobi, Abh. Schles. Ges. Vaterl. Cult., Abth. Naturwiss.

1868: 147. 1868.

Medium to small, dark green to glaucous green, symmetrical

rosettes 7X9 dm., surculose in early years, with graceful arching leaves; leaves 50-60 X 10-12 cm. to 70-80 X 7-9 cm., linearlanceolate, openly ascending, turgidly brittle, abruptly or gradually narrowed towards the base, the base thicker than wide in

cross section, finely and sparsely fibrous, unarmed with smooth

unmargined edges or armed with small teeth ; teeth 1-2 mm. high,

regular, chestnut brown, 1-2 cm. apart or few and irregularly

spaced; spine 2-3 cm. long, subulate, dark brown to fuscous,

shortly and broadly grooved above ; inflorescence paniculate, rather

deep and narrow in upper 1/2 to % of shaft, 2.5-3 m. tall; bracts
THE AGAVE FAMILY IN SONORA 95

FIGURE 31.—Distribution of 7 species of the subgenus Agave in Sonora, represented by their ideographs. Ideographs with numbers indicate flower collection at the locality occupied. (See p. 37.)

short, deltoid ; laterals 20-25 with congested umbels of pale yellow

flowers about 40 mm. long, on short petiolules subtended by short

scarious bracteoles ; ovary short-stipitate, small, 12-14 mm. long,

green, 6-grooved above, with a very short unconstricted neck;

tube 10-12 mm. long, 14 mm. broad, prominently flanged within

below inner tepals, deeply grooved and ridged outside; tepals

13-15 mm. long, green in the bud, yellow at anthesis, drying fer-
96 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

FIGURE 32.—Agave desmettiana: Leaf and flower clusters X Vs; flower section slightly enlarged. Drawn from Gentry 11569.

ruginous, the outer 1 mm. longer than the inner and broadly overlapping at base, the inner sharply involute ; filaments 30-35 mm.

long, inserted 6-8 mm. above the tube base, yellow ; anthers 13-15

mm. long, attached in the middle ; capsule and seeds not seen.
THE AGAVE FAMILY IN SONORA 97

Type—Not specified and origin unknown. Syntype—Gentry

11569, Guasave, Sinaloa, Mexico, Feb. 8, 1952, cultivated in yard

of house (If. & ñ.) (fig. 33). SONORA: Guaymas, Dec. 17, 1951,

Gentry 11359 (live pi. only). TEXAS: Ringold Park, Brownsville,

Mar. 14, 1960, Gentry & Barclay 18U1 (If. & fl.). It has been

observed frequently as a cultivated ornamental along the west

coast of Mexico from Hermosillo to Mazatlan. Inquiries from

several owners along the west coast route in 1952 established the

story that it was carried by a peddler 7 or 8 years previously,

who sold small plants to housewives. It was subsequently observed

as houseplants along the east coast from Tampico northward. Its

natural habitat remains unknown; but judging from its tropical,

frost-sensitive, delicate leaves, it should be expected in the more

humid mountains of Central Mexico. Trelease suggested the Huatusco region of Veracruz as the place of origin {J^3, p. 121. 1920).

This species is distinguished by its smooth, unarmed, arching

leaves, short, compact, paniculate inflorescence, and small flowers

with very short ovary and broad, bulging, deeply furrowed tube.

The deep exterior furrows are complemented within by conspicuous costae that border the filaments along the inner tepals. These

provide structural strength to the thin perianth above the thickened nectariferous tissue of the lower half of the tube. This costate

structure is well developed in the section Sisalanae. The present

taxon differs from all other members of the section reviewed by

its smaller, more graceful rosettes, small dense panicles (vs. tall,

diffuse), and smaller flowers (35-40 mm. vs. 60-90 mm.) with

a very short, nearly neckless ovary. Like A. sisalana and some

other members of the section, A. desmettiana may have either perfectly smooth leaf margins, or some leaves with small, irregularly

spaced teeth, or others with a regular set of small teeth, as in the

Ringold Park collection in Brownsville. The nearer relatives include A, miradorensis and A. regeliana, both described by Jacobi

in European collections but which apparently originated from the

environs of Mirador in Veracruz, Mexico.

There are no tangible characters in Jacobi's descriptions or

in his sketches (reviewed in the U.S. Nati. Herbarium) that distinguish these taxa as species. Berger (5) in a footnote under

A. miradorensis states that it is the same as A. desmettiana, which

is the priority name. The inflorescence of A. regeliana is still unknown, and unless recorded clonal descendants are still available

in Europe, it will never be known. The other sectional relatives,

including A. franceschiana Trel., A. weberi Gels, and A. neglecta

Small, are larger, coarser plants with tall, diffuse panicles and

larger flowers, and are all more xerophytic than A. desmettiana.
98 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

W- »»,

PN-2135

FIGURE 33.—Agave desmettiana flowering panicle cut from a cultivated plant

at Guasave, Sinaloa.
THE AGAVE FAMILY IN SONORA 99

There appear to be more names than species in this group, but

these will be dealt with more comprehensively in a later, more

appropriate work. The Desert Nursery near Riverside, Calif., has

a fine growing series of this section, with some individuals that

appear to be recent introgressive hybrids from local cultivations.

A, desmettiana with its soft, pliant, arching, and frequently

glaucous leaves is an attractive ornamental. The glaucous quality

and color varies with growing conditions and among individuals.

It is, however, sensitive to frost and young plants were killed at

Murrieta, Calif. In the Mexican gardens, it grows rapidly when

given plenty of water, throws vigorous offsets early in life, and

blooms in the early spring within 8 to 10 years. The fiber is fine

and weak. The leaves were found to contain mostly negligible

amounts of sapogenin, up to 0.3 percent. Except as an ornamental,

no local uses were observed or reported for the plant. (Fig. 33.)

Agave murpheyi F. Gibson, Boyce Thompson Inst. Contrib. 7: 83,

fig. 1.1935.

Medium-size, freely suckering by rhizomes, light green, compact

rosettes, 60-80 cm. tall, about 1 m. broad, with numerous leaves,

spring-blooming but fall-starting; leaves 50-65 X 6-8 cm., linear,

short-acuminate, light glaucous green to yellowish green, frequently lightly cross-zoned, firm, straight, bud-printing clear, the

margin undulate with small regular teeth 3-4 mm. long, mostly

1-2 cm. apart, bases brown, cusps graying; spines short, 12-20

mm. long, conic, very shortly grooved or flattened above, dark

brown becoming grayish; panicle bulbiferous, 3-4 m. tall on a

thick shaft with scarious, triangular bracts 10-15 cm. long and

10-15 short, compact laterals in upper 14 of shaft; flowers pale

waxy green with purplish to brownish tips, 65-75 mm. long ; ovary

thick, rounded, 32-40 mm. long, scarcely narrowed at neck ; tube

deep, urceolate, 16-20 mm. deep, 15-19 mm. wide, 6-grooved from

the tepal sinuses; tepals unequal, the outer longer, 15-19 mm.

long, 7 mm. wide in the middle, overlapping the inner, linear,

rounded on back, strictly erect, thick, the tip thickly hooded,

brownish, the inner strongly keeled, pale yellow ; filaments 45-50

mm. long, unequally inserted 8-12 mm. above bottom of tube,

yellow; anthers 25-26 mm. long, yellow, slightly excentric: capsules obovate to oblong or ovate, 2-2.5 X 5-6 cm., stipitate,

beaked; seeds "dull black, 6X9 mm., thin'' (loc. cit.).

Type—from near Boyce Thompson Southwestern Arboretum,

Superior, Ariz., deposited there and the type clone cultivated. The

species is reported to grow near Roosevelt Dam, Tonto Basin, and
100 AGRICULTURE HANDBOOK NO. 399. US. DEPT. OF AGRICULTURE

Ü

PN-2136, PN-2137

FIGURE 34.—Agave murpheyi at San Luisito, Sonora. Small off-sets clutter

around the base of the flowering plant. The panicle is typically full of

bulbils.
THE AGAVE FAMILY IN SONORA 101

Paradise Valley, and along the base of Superstition Mountains,

all in central Arizona. The writer also saw it cultivated at Sells,

Ariz., and a Papago Indian reported the same had been brought

in from the neighboring mountains in view to the southwest of

Sells. SONORA: San Luisito, along road to Quitovac from Caborca, Sept. 4, 1965, Gentry & Arguelles 2120Jf, Cultivated in a

yard, whose owner stated that his plants had been brought from

the nearby hills (1-2 miles), where it is wild. He dissuaded me

from going there by saying that all the larger plants had been

cut for eating or for making mescal. (Fig. 34.)

The flower structure clearly alines this agave with other members of the section Ditepalae. In addition to the separating characters given in the key to the species, I may add here its proliñcally bulbiferous inflorescence, which is short, compact, and

limited to the top 1/4 or Vs of the shaft. It blooms much earlier

than other agaves in its area. Its habit of starting the flowering

shoot in the fall means that the succulent, turgid tissues are resistant to the winter freezes regularly occurring in its habitat.

Gibson wrote (loc. cit.) : ''Inflorescence very strongly bulbiferous,

lasting into the second or third year. The bulbs form in the axils

of small scaly bracts on the pedicels, from adventitious buds, or

from the base of other bulbs.'' All plants I have seen show relatively little variation and once the species is observed, it is readily

recognized again. The plants cultivated at San Luisito have longer

leaves with more closely set teeth than the type clone, but the

aspect and essential characters of the species are plainly there,

including the prolific reproduction with bulbils and rhizomatous

offsets. (Fig. 34.)

Agave palmeri Engelm., Acad. Sei. St. Louis Trans. 3: 313.1875.

Rosettes at first single, later suckering, 5-12 dm. tall, 10-12

dm. broad, rather open about the conal bud ; leaves mostly 35-75

X 7-10 cm. usually narrowed above the base, lanceolate, longacuminate, rather rigid, somewhat guttered, thick at base, convex below, pale green to light glaucous green or reddish-tinged,

the margins nearly straight or undulate with small teatlike bases

under closely set, rather regular, slender teeth, which are variously flexed, smaller teeth sometimes between the larger; spine

strong, 3-6 cm. long, shortly and openly grooved above the base,

castaneous or brown to aging gray, acicular ; panicle deep, broad,

open, 3-5 m. tall, with triangular bracts and 8-12 horizontal

laterals in upper 1/3 of shaft; flowers 45-55 mm. long, narrow,

pale greenish yellow to waxy white, reddish in bud ; ovary shiny
102 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

green, 25-30 mm. long including short neck ; tube 12-14 mm. deep,

10 mm. wide; tepals persistently erect, leathery, yellow to pink

below, conspicuously red to brownish on calloused tip, dimorphic,

the outer 10-13 mm. long, longer than and overlapping the inner,

the inner 2 mm. shorter and narrowly keeled on back; filaments

40-45 mm. long, reddish or pale yellow, inserted midway in tube

with those opposite the outer tepals inserted higher than the

others; anthers 15-18 mm. long, yellow; capsule oblong to pyriform, 3.5-6 X 1.8-2 cm., the dried walls thick and strong, short

to long apiculate; seeds sooty black, 5-7 mm. along the straight

edge, 4-5 mm. broad, thin, flat.

Type—Not specified; from Schott, Palmer, & Rothrock who

made scattered collections in southeastern Arizona. SONORA:

Nogales, June 4, 1897, Rose 1200 (If.) ; Arroyo de la Galera (loop

of the Bavispe River), July 27, 1940, White 30Í9 (If. & fl.) ; Santa

Rosa canyon near Bavispe, July 15, 1938, White 517 (If. & fl.) ; 16

miles southeast of Magdalena along road to Cucurpe, April 1963,

Gentry 19888; Sierra de Pajaritos, about 12 miles east of Ures,

Mar. 11, 1951, Gentry 1027Í (If., cap., & photo) ; mountain above

east of Guasabas on volcanics in oak woodland, alt. 4,000 feet.

May 21, 1957, Gentry 166Jfl (live pi.) ; 6.3 miles northwest of

Nacori Chico, June 10, 1960, Feiger & Marshall 33^3 (If. & buds)

'Vith lower oaks ca. 3,400 feet alt., common, non-suckering'' ; El

Coyote northeast of Moctezuma on north-facing slopes with lower

oaks, alt. about 3,400 feet, July 19, 1960, Feiger & Beasly 3525,

''now in full flower^—flowers white with callous tip, ovary green'' ;

16.1 kms. north of Nacozari, oak-grassland, July 22, 1960, Feiger

3595 (If. & fl.) ; 15.4 miles southeast of Magdalena along road to

Cucurpe, alt. about 3,400 feet, July 16, 1960, Feiger SU57 (If. & fl.)

and at 16 miles southeast of Magdalena, Barclay & Arguelles

2013; 8.3 miles north of Magdalena, alt. about 3,100 feet, Aug.

19, 1960, Feiger 3781 (If. & fl.) ; 15 miles north of Fronteras, alt.

about 4,100 feet, Sept. 7, 1960, Feiger Í026 (If. & fl.) ; Sierra del

Humo, Dist. Caborca, near Rancho Cimarrón, Sept. 3, 1965, Gentry 21200 (If., cap., & live pi.). CHIHUAHUA: Near Sierra en

Media, plains, Sept. 24-26, 1899, Nelson 6Í69 (If. & fl.), the only

collection I have seen from Chihuahua. (Fig. 35, lower,)

A. palmeri is a widely scattered but characteristic plant of the

oak woodland and grama grassland communities at altitudes between 3,000 and 6,000 feet. It shows good development on the

limestone slopes at altitudes between 4,000 and 5,000 feet on the

Patagonia Hills and along highway 86 where it passes through

the granite of Texas Canyon in Arizona. A smaller form occurs

in northern Sonora in somewhat lower elevations on the rocky
THE AGAVE FAMILY IN SONORA 103

PN-2138, PN-2139

FIGURE 35.—Agave palmeri: Upper, glaucous form on the lower slope of

Sierra Baviso ; lower, small form on Sierra del Humo.
104 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

brush slopes and comprises the western segment of the species.

(Fig. 35.)

It flowers mainly in June and July. Hummingbirds, other wild

fowl, and many insects visit the flowers. The night visitors are

unknown to me. The capsules from the Sierra de Pajaritos show

large pustulate scars, probably made by the large, sap-sucking

bugs of the family Coreidae observed on the plants. Good black

seeds have been collected from wild plants, and progeny are also

produced by rhizomatous offsets.

Characteristic of the species are the long lanceolate leaf, typically narrow in its northern range, its close-set, slender teeth,

sometimes with smaller ones between, and the reddish tepals about

equaling the tube. About the margins of its considerable area it

shows what is taken for introgressive variation ; on the north in

the Ricon Mountains with A. chrysantha Peebles, on the south

in east central Sonora with A. shrevei. I have separated the southern representatives of A. palmeri from A. shrevei on leaf and

flower characters; i.e., the broader leaves and the much deeper

tube and relatively shorter tepals of the latter (fig. 4, A and B).

The broad, glaucous green leaves in populations in the Rio Bavispe

vvatershed resemble those of A. shrevei, but flowers collected by

Feiger—3595, 3525, 33Í3—show the smaller flowers with shallower tubes of A. palmeri and led me to cite these collections,

along with Gentry 166Jfl, under this species. An introgressive

aspect in these eastern Sonoran populations extends as far south

as the upper Guajaray country.

Peebles {30) has given good key characters for separating A.

chrysantha from A. palmeri. Little {32) reduced A. chrysantha

to a variety of A. palmeri on the grounds of intergrading characters, but hybridization is much better understood now, and no

longer in itself constitutes grounds for combining species. In fact,

it can be a measure of their separateness. The scattered population along the main highway in Sonora between Nogales and

Magdalena consists of small plants, reflecting the drier and lower

elevations. Similar small plants were encountered on the Sierra

del Humo (flg. 35) and on the northern slopes of the Baboquivari

Mountains, along the grade up Kitt Peak. These populations are

apparently A. palmeri and perhaps their small apartness should

be nominated subspecifically. However, there is insuflScient evidence for doing this. They are a small part of a large polymorphic

group, including, besides the species mentioned, A. colorata, A.

flexispina, and other nondesignated forms ranging from central

Arizona and New Mexico into Durango, Mexico. There are very
THE AGAVE FAMILY IN SONORA 105

few specimens and far fewer flowers available for studying the

systematic problems posed by this group.

In northeastern Sonora, the people called A, palmeri ''lechuguilla'' and said it was collected for eating and for making mescal.

It was much used for food, fiber, and beverage by the Indian tribes

inhabiting its region. Castetter, Bell and Grove (8) report its

uses by the Papago, Pima, and Apache Indians. The young, tender,

emerging flowering shoots are still eaten by the Mexicans. They

are also eaten by cattle, rodents, and other animals. It is among

the sweeter kinds with little or no sapogenin reported in assays

(52) ; 0.5 percent hecogenin was found in the leaves of Gentry &

Ogden 9915, Pyramid Mountains, New Mexico. Several other

assays from plants in other localities were negative.

Agave wocomahi Gentry, Carnegie Inst. Wash. Pub. 527, p. 96.

1942. Figure 36

Rosettes medium to large, 0.8-1.3 m. tall, 1.5-2 m. broad, nonsuckering, eventually depressed and open at maturity; leaves 30

X 9 cm. to 90 X 25 cm., mostly lanceolate to linear-lanceolate,

rarely ovate, somewhat narrowed towards the base, plane, thickfleshy, rather rigid, ascending to depressed in age, dark green

to glaucous green, smooth, the margins straight to undulate with

large teeth, 1-2 cm. long, dark brown to glaucous brown, variously flexed, these below the midblade frequently down-flexed,

and smaller interstitial teeth irregularly occurring; spine stout,

3-6 cm. long, usually sinuous, flattened or hollowed in a broad

groove above, short or long-decurrent ; panicle 3-5 m. tall, open,

with 8-15 laterals in upper 1/3 of shaft; bracts in the midshaft

15-20 cm. long, scarious, appressed or reflexed; flowers yellow,

in small umbels, 65-85 mm. long, erect; ovary cylindrical, 34-40

mm. long including the neck 2-5 mm. long, light green ; tube deeply

funnelform, 18-22 mm. deep, 15 mm. broad, light yellow, narrowly

grooved below the tepal sinuses; tepals dimorphic, 15-23 mm.

long, 5 mm. wide, yellow, erect, thick, linear, conduplicate, rounded

and deeply hooded at tip, the outer with papillate pubescence well

below apex and sometimes red-tipped, the inner shorter and prominently keeled ; fllaments 60-65 mm. long, inserted unequally 9-14

mm. above base of tube; anthers yellow, large, 26-34 mm. long,

eccentric; pistil with a large stigmatic head; capsules shortstipitate, oblong, 50-60 mm. long, 15 mm. or more in diameter;

seeds 7 X 4.5-5 mm., shiny black, oblique, flattened on the end

opposite the hylar notch, finely punctulate, wavy corrugate, with

a narrow margin partly upcurved.
106 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

FIGURE 36.—Agave wocomahi: Leaf drawn from type, Gentry 1989; flowers

from Gentry 10254, cluster X %, flower section slightly enlarged; capsule

X 'V2 and seeds X 2 from Gentry 17666.
THE AGAVE FAMILY IN SONORA 107

Type—Huicorichi, Sierra Huicorichi, Rio Mayo, Chihuahua,

Oct. 7, 1935, Gentry 1989 (fl. & If.). CHIHUAHUA: same locality

as type specimens. Gentry & Arguelles 11397; high rocky rims

around Arroyo Hondo, Sierra Charuco, Mar. 6-7, 1951, Gentry &

Arguelles 10252, 1025J, (flowered at Murrieta, Calif., July 25-

Aug. 10,1962 (ñg. 37, upper) ) ; Sierra Chinatu, Dist. Guadelupe y

Calvo, Oct. 9, 1959, Gentry, Correll, & Arguelles 17988 (If.). SONORA: Sierra Charuco, Rio Mayo, Mar. 6, 1951, Gentry 10225 in

part (If.) ; Guajaráy Scarp, north of Curopaco and Arroyo Guajaráy, Feb. 13, 1952, Gentry 11586, 11588 (If.) ; Sierra de Yecora,

Nov. 15, 1958, Gentry & Arguelles 17666; 6 miles east of Reparo

along road from Nuri to Yecora, Dec. 11, 1959, Feiger & Stronack

3052 (If. & cap.) ''Upper edge of Oak Forest—common, leaves,

green." SINALOA: Ocurahui, Sierra-Surotato, Sept. 1-10, 1941,

Gentry 68iO (fl. & If.). Native to the rocky, calcareous, open

mountain slopes through the pine-oak forests at elevations between 4,500 and 7,500 feet (fig. 38). Because of the montane elevations it occupies the most humid habitat of all the paniculate

Sonoran agaves. Although I have observed the species from northern Durango and northern Sinaloa to Sierra de Yecora, Sonora,

it is an uncommon plant, with relatively few individuals in small,

widely scattered colonies. (Figs. 37 and 38.)

On the basis of floral morphology A. wocomahi belongs in the

section Ditepalae, as indicated in the key to the species, page 42. It

sometimes grows with A. shrevei, from which it is usually easily

distinguished by its dark green color, larger size, or larger and

more numerous distal teeth. It is more easily confused with A.

bovicornuta, as discussed under that species (See p. 88) ; compare

also respective ideographs in figure 4, B,

An unusual variant with very short, broad leaves is represented

by Gentry 11588, found along the Guajaráy Scarp, a striking topographic feature described in ''Rio Mayo Plants'' {13). I have noted

no intergrading of A. wocomahi with its neighbors.

The Warihio Indians and their Mexican neighbors called this

plant ''wocomahi," stating that it was sweet and suitable for eating and for making mescal. Both these peoples and the mountaineers of Sinaloa stated that the flowers were cooked and eaten

like squash. The fiber of the leaves is used locally for cord, rope,

and pack saddle pads. A single plant that flowered in my Murrieta

garden in 1962 did not set viable seed, indicating that it requires

cross-pollination. This maturation required 11 years from the

small plant collected, Gentry 10252, in 1951. No vegetative reproduction has been observed in this species and it appears to be a

confirmed seeder and outbreeder.
108 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

PN-2140, PN-2141

FIGURE 37.—Agave wocomahi: Upper, with budding inflorescence in garden

at Murrieta, Calif. ; lower, form with more uniform teeth among volcanic

rocks on Guajaráy Scarp in southeastern Sonora.
THE AGAVE FAMILY IN SONORA 109

PN-2142, PN-2143

FIGURE 38.—Upper, Agave wocomahi in flower at Murrieta, Calif., Gentry

10254, summer of 1962 ; lower, Agave shrevei in flower at Murrieta, summer of 1965.
lio AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

For distribution of the Ditepalae group of agave, see figure 39.

FIGURE 39.—Distribution of Ditepalae group of agave in Sonora, represented

by their ideographs. Ideographs with numbers indicate flower collection

at the locality occupied. (See p. 37.)
THE AGAVE FAMILY IN SONORA 111

Agave shrevei Gentry spp. shrevei Figure 40

Agave shrevei Gentry, Carnegie Inst. Wash. Pub. 527, p. 95.1942.

Small to medium, light gray, glaucous rosettes, closely or widely

suckering with maturity; leaves ovate, short-acuminate, 20-35

FIGURE 40.—Agave shrevei: Smaller leaf drawn from type. Gentry 2028;

larger leaf from Gentry 11580; flower cluster X Va from Gentry 19865;

flower section slightly enlarged from Gentry 8080; capsules X % and

seeds x 1% from Gentry 21211.
112 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

X 8-10 cm., plane to conduplicate, lanceolate, acuminate, 50-60

X 12-18 cm., generally narrowed above the base, firm, thick,

straight or outcurving near the tip; teeth variable, on small to

prominent teats, the larger teeth in midblade 5-10 mm. long,

straight or flexed upward or downward, dark brown to gray;

spine stout, mostly 25-50 mm. long, brown, acicular, with a narrow or open groove from base to above the middle; panicle 2.5-5

m. tall with 8-16 distal, ascending laterals in upper 1/3 ; bracts

deltoid, long-acuminate, drying early ; umbels small ; flowers persisting erect, slender, leathery, 60-70 mm. long, light green to

pale yellow, buds red-tipped; ovary 25-35 mm. long including

constricted neck; tube cylindric or urceolate, 18-23 mm. deep,

10-12 mm. wide ; tepals strictly erect, leathery, pea green to light

yellow, unequal, the outer 10-12 mm. long with red to purplish

calloused tips, plane, widely overlapping the shorter, keeled inner ;

filaments 40-50 mm. long, yellow or red, inserted unequally in

midtube, flattened; anthers large, 22-26 mm. long, yellow; capsules oblong, 4.5-7 X 1.5-2.5 cm., short-stipitate and shortly

beaked; seeds 6-7 X 4.5-5 mm., hilar notch small, the marginal

wing prominent.

Ty^Q—Gentry 2028, Sierra Canelo, Chihuahua, Oct. 8, 1935

(old dried flowers) deposited in Dudley Herbarium, Stanford

Univ. SONORA : Sierra Tecurahui, Nov. 1961, Gentry, Barclay, &

Arguelles s,n. (fig. 41, lower) ; Sierra Potrero, west of Arroyo

Guajaráy, Gentry 11378 (flowered at Murrieta, Calif., August

1957) ; Guajaráy Scarp, north of Curopaco, Gentry 11580

(flowered at Huntington Bot. Gard., August 1962), 11582, Feb. 13,

1952; Rancho Santa Barbara del Agua Blanca, northwest of

Curopaco, Gentry 11598, Feb. 14, 1952 (If.). CHIHUAHUA:

Arroyo Hondo, Sierra Charuco, April 1948, Gentry 8080 (pi.),

flowered at Murrieta, summer 1965. Common and widely scattered

from middle Sonora southward on both sides of the ChihuahuaSonora boundary, from 3,000- to 6,000-foot altitudes of the

northern Sierra Madre Occidental, in open, rocky, calcareous

slopes of the oak woodland and pine-oak forests. Flowers in

summer. (Figs. 38, 40-42, 44.)

Known locally as "lecheguilla ceniza" or *'totosali'' by the

Warihio Indians, who pit bake the heads for eating.

Good distinguishing features of this agave are its light glaucous

gray, broad leaves with teated margins under well-developed

brown teeth and its leathery perianth with deep tube, short,

persistently erect tepals, much shorter than the tube. Some of

the flower forms are quite striking, as the red filaments in a
THE AGAVE FAMILY IN SONORA 113

7"^

PN-2144, PN-2145

FIGURE 41.—Agave shrevei: Upper, flowers with visiting bee; lower, colony

of plants on Sierra Tecurahui, southeastern Sonora.
114 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

PN-2146, PN-2147

FIGURE 42.—Agave shrevei: Upper, large-leaved form along Guajaráy Scarp;

lower, cultivated as food plant in Chijucu, a Mayo Indian suburb pf

Navojoa.
THE AGAVE FAMILY IN SONORA 115

pea-green perianth. The typical or smaller leaved A. shrevei show^s

leaf resemblance to A. flexispina Trel., but the latter's flowers

appear closer to A, palmeri, the tube and tepals being about equal.

The Guajaráy population with its larger leaf and variable forms

shows some A. palmeri characters and there appears to be some

introgression. The obscure relative Gentry 11605, cultivated in

Navojoa, was reported as from Tepahue, which is adjacent to the

Guajaráy country (fig. 42, lower). The Mayo Indian growing the

plants in his garden in the ''bario'' known as Chijuca also stated

that he grew them to eat, as the pit-baked heads were very sweet.

Agave shrevei subsp. matapensis Gentry, ssp. nov. Figure 43

Small to medium, light gray glaucous, rather open rosettes,

suckering late and sparsely ; leaves short-lanceolate, 30-45 X 8-14

cm., mostly plane to conduplicate towards apex, the larger teeth

in the midblade down-flexed, the margin nearly straight to undulate; panicles 4-5 m. tall with 8-14 laterals in upper half of

shaft ; flowers waxy yellow-green with tips of outer tepals reddish ;

filaments greenish to pink, anthers yellow ; ovary 22-40 mm. long

including the short unconstricted neck 2-3 mm. long; tube

cylindric, 15-20 mm. long, 11-14 mm. broad; outer tepals 11-16 X

7 mm., rounded on back, thick ; filaments inserted 8-10 mm. above

base of tube ; filaments 45-55 mm. long, slightly flattened ; anthers

20-28 mm. long; capsules (Gentry 17623) short, 35-40 X 20 mm.,

short-oblong, apiculate, the valves strong and woody ; seeds smokey

black, mostly 5-7 mm. along straight edge, 5-6 mm. broad, rugose,

scarcely winged and unfluted along margin.

Type—Gentry 11607, Sierra Batuc, North of Matapé, Sonora,

Feb. 16, 1952, deposited in U.S. Nati. Herbarium. No. 2540344.

SONORA: 2 miles east of Nacori, Feb. 16, 1952, Gentry 11612

(If.) ; between Nacori and Matapé, May 12, 1957, Gentry 16600

(live pi., flowered at Murrieta, Cahf., in 1962). Mountain pass 6-8

miles northeast of Matapé, Nov. 4, 1958, Gentry 17622 and 17628

Planta caule singulo vel caespitosula ; foliis pallidis, glauco-viridibus,

lanceolatis, 30-45 cm. longis, 8-14 cm. latis ; dentibus validis, 4-14 mm, longis,

subulatis, deflexis, in mammae proximis, castaneis; spine terminali 2-4 cm.

longa, valida, castanea, supra canaliculata, decurrente; inflorescentia 4-5 m.

alta, paniculata, cum 8-15 ramis lateralibus; floribus luteo-viridibus, ca. 50

mm. longis, gracilibus; ovario 20-40 mm. longo, ápice crasso incluso; tubo

15-20 mm. longo, urceolato; segmentis inaequalibus, exterioribus 11-16 mm.

longis, 7 mm. latis, apice rubro; filamentis 45-55 mm. longis, luteis, medio

tubi insertis; antheris luteis, 20-28 mm. longis; capsulis oblongis 3.5-4 cm.

longis, 2 cm. latis; seminibus 5-6 X 6-7 mm., nigris, enervis, apice emarginatis.
116 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

FIGURE 43.—Agave shrevei matapensis: Leaf drawn from Gentry 11607;

flowers from Gentry 16600; cluster X %, section X 1; capsules X Vs and

seeds X IV2 from Gentry 17623.
THE AGAVE FAMILY IN SONORA 117

(cap. & seed). Widely scattered in sparse colonies in the savanillas

of lysiloma-oak-nolina woodland on granitics and deformed limestones, alt. 2,000 to 3,000 feet. The largest colony observed was in

and about the mountain pass 6-8 miles northeast of Matapé along

the road to Sahuaripa.

These small to medium, light-gray glaucous rosettes resemble

some forms of A. shrevei, from which it is distinguished by its

deflexed teeth, smaller flowers with shorter ovary unconstricted

in the very short neck, the broader and longer tepals in proportion

to the shorter tube, and the short, broad capsules. It produces

offsets late and seeds abundantly. The variability observed about

Matapé indicates an outbreeding population. The flower description is drawn mainly from pickled flowers from plants that

flowered in my Murrieta planting.

Agave colorata Gentry, Carnegie Inst. Wash. Pub. 527, p. 95.1942.

Small to medium, light-gray, compact, few-leaved rosettes on

short stems, suckering while young; leaves 25-60 X 12-18 cm.,

ovate, short-acuminate, thick, firm, rounded below, plane to shallowly concave above, frequently cross-zoned and red-tinted, glaucous, the margins prominently crenate, asperous; teeth mostly

4-8 mm. long, smaller below, 1-3 cm. apart, mildly flexed to

straight, brown to grayish; spine subulate; straight or flexuous,

25-50 mm. long, narrowly grooved above in lower half; panicle

2-3 m. tall, narrow, with densely flowered laterals in upper 1/2

or % of red shaft; lower bracts leaflike, the others soon drying

appressed at base, reflexed, reddish, 15-10 cm. long; flowers 50-65

mm. long ; ovary including neck 24-30 mm. long, yellowish green ;

tube 14-16 mm. long, 14 mm. wide, yellowish; tepals 12-16 X 5

mm., coppery red to yellow, leathery, strictly erect, unequal ; filaments yellow or pink, 40-50 mm. long, inserted unequally 6-9 mm.

above base of tube ; anthers 20-26 mm. long, eccentrally attached,

yellow; fruits large, oblong, green; capsules and seeds not seen.

Type—SONORA : Peñasco Blanco, near Aquibiquichi, February

1937, Gentry 3050 in Dudley Herbarium, Stanford Univ.; same

locality, March 9, 1951, Gentry 10270 (fl.) ; Rancho Pachaca, about

20 miles east of Navojoa, Dec. 19, 1951, Gentry 11368 (live pi.) ;

Sierra Bojihuacame, east of Ciudad, Obregon, March 1952, Gentry

& Arguelles 11641 (live pi.). SINALOA: Los Pucheros, Sierra

Surotato, Mar. 17-24, 1945, open rocky slope in pine-oak forest,

Gentry 7237. Mainly on open rocky sites in the volcanic foothills

of the "tierra caliente'' at elevations between 200 and 2,000 feet.
118 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

¿%6

PN-2148, PN-2149

FIGURE 44.—Upper, Agave colórala flowering at Murrieta, Calif., in June

1965, grown from Gentry 11368. Flowering shoots of A. shrevei adjacent

are later (see opposite page) ; lower, rosette details of A. colorata.
THE AGAVE FAMILY IN SONORA 119

With its dimorphic tepals and large, deep tube, A. colórala fits

nicely into the section Ditepalae. Its nearest relative, morphologically and geographically, is A. shrevei, from which it differs

in the broader leaf with deeply crenate to undulate margins, the

more densely ñowered panicle with more numerous and more

closely set laterals, and the shorter, wider tube and thicker ovary.

A. shrevei grows in the higher altitudes of the Sierra Madre,

while A. colórala in Sonora is restricted to the foothills on open

rocky sites in the thorn forest.

The small live plants collected in 1951 flowered at Murrieta in

June 1965, indicating a sexual generation period of about 15 years

(fig. 44). The first flowers along the shaft did not set fruits, but

fruits developed with ovules on the last terminal laterals. It was

during the anthesis of these later flowers that a nearby specimen

of A. shrevei came into flower. Hummingbirds visited the flowers

of the two plants. From this one may infer that A. colórala is

perhaps self-sterile and that it can be pollinated with A. shrevei.

However, ground squirrels chewed off the capsules before seed

development could be verified.

A. colórala is uncommon in nature and on my very intensive

travels through its area I do not recall it from any localities except those listed above. It was reported by the natives as suitable

for pit baking and was referred to as ''mescal ceniza.'' It was

perhaps formerly decimated as a sugar source by the Mayo and

Yaqui Indians who surrounded its area rather densely in precolonial times. Because of its compact size and bright, glaucous

leaves with pinkish cross-zones, it is an attractive ornamental,

especially those plants from the type locality. It is now (1971)

found in southwestern gardens (6),

Agave huachucensis Baker, Handb. of the Amaryllideae, p. 172.

1888.

Agave applanata var. huachucensis (Baker) Mulford, Mo. Bot. Gard. Ann.

Rpt. 7: 85. 1896.

A. parryi var. huachucensis (Baker) Little ex Benson, Amer. Jour. Bot. 30:

235. 1943.

Compact, light gray, multileaved, short, broad, widely suckering rosettes 5-7 dm. tall, 8-12 dm. broad ; leaves 25-40 X 10-15

cm., incurving, nearly plane, ovate-lanceolate, short-acuminate,

slightly narrowed below the midblade, light gray glaucous, thick,

rigid; teeth small or larger, the larger along the upper part of

blade 2-3 mm. long and nearly straight or 5-7 mm. long and
120 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

somewhat flexuous, the margin slightly undulate with or without

a narrow corneous margin ; spine 2-3.5 cm. long, brown to aging

grayish, acute-triangular and flattened or hollowed above, decurrent; panicle 3.5-4.5 m. tall, deep, broad, with 24-30 horizontal

laterals in upper half of shaft, peduncular bracts 10-20 cm. long,

broad, drying early, recurving; flowers bright lemon yellow, 55-65

mm. long, closely clustered ; ovary 30-35 mm. long, 3-angled and

grooved, virtually neckless, tapering from base to tube ; tube 8-10

mm. deep, 16-18 mm. wide, funnelform ; tepals 19-21 mm. long,

5-6 mm. wide, lanceolate, appressed, incurving after anthesis,

with open sinuses; filaments 45-65 mm. long, flattened on inner

side, rounded on outer side, inserted 7-8 mm. above bottom of

tube at base of tepals, with nectary rimming the tube; anthers

20 mm. long, eccentrically affixed, yellow (from Gentry 19829) ;

capsules '^broadly oblong, 20-25 X 55-75 mm., stipitate and

beaked ; seeds 6x8 mm.'' (Trel. loc. cit.).

Type—Collected by Pringle in 1884 in Huachuca Mountains,

Arizona, at 5,000 to 8,000 feet elevation. SONORA : Sierra Huachinera, south of Bavispe, alt. about 7,500 feet in pine forest.

May 21, 1957, Gentry & Arguelles 166Í2 (photo & live pi.). Northern Sierra Madre including southeastern Arizona, where it occurs

as widely scattered clones in small colonies in oak woodland and

pine forests, alt. 4,500 to 7,500 feet. In Sonora it is restricted to

the northeast quarter of the state. (Fig. 45.)

The Mexican populations of this taxon are not well known.

Flowering specimens have not been collected from Sonora. The

highlands of both Sonora and Chihuahua have scattered populations with rosette and leaf characters strongly resembling the

Arizona plants around Sonoita and in the Huachuca Mountains.

Some extensive stands occur west of Chihuahua city, which Trelease called A. chihuahuana. Without more materials for study

I am reserving judgment about taxonomic changes and am reporting it under Baker's original binomial. If this group is to be

considered a part of the species, A. parryi Engelm., then a subspecific status can be established when the morphological and

geographical limits are known.
THE AGAVE FAMILY IN SONORA 121

PN-2150

FIGURE 45.—Agave huachucensis near Sonoita in southeastern Arizona, June

1968.
122 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Agave fortiflora Gentry, sp. nov. Figure 46

Light-gray, open rosettes up to 1 X 1.8 m., mostly single but

occasionally suckering; leaves 50 X 8 to 100 X 12 cm., straightly

ascending or outcurving and conduplicate, light gray glaucous,

usually cross-zoned, widest in the middle, gradually narrowed

above the dilated base, long-acuminate, the margin straight or

small-teated under the midteeth ; teeth in the midblade 5-10 mm.

long, 1-3 cm. apart, generally flexed downward but also erect,

smaller and variously flexed above and below the middle, with

smaller interstitial teeth irregularly occurring; spine 3-5 cm.

long, subulate, chestnut to light gray, rounded below, narrowly

grooved above, decurrent with a narrow corneous margin to the

upper teeth; surface of leaf, spine, and teeth finely tuberculate

rugose; panicle 4-6 m. tall with 12-18 laterals, open and deeply

ovoid in outline; bracts short, triangular, chartaceous, 5-6 cm.

long at the lower laterals; pedicels 4-20 mm. long, bracteolate;

flowers in dense yellow umbels, long-persisting, erect, 72-82 mm.

long; ovary pale green, 45-50 mm. long, conspicuously angled below outer tepals; tube pale yellow, broadly bulging, 11-13 mm.

deep, 18-20 mm. wide, the sinuses overlapping and deeply grooved

below; tepals erect, yellow, 20-23 mm. long, 7-8 mm. wide, the

outer rounded, linear-lanceolate, with an involute margin, obtuse,

the inner with a high narrow keel and involute hyaline margins,

broadly hooded at tip; filaments strong, persisting erect after

anthesis, 60 mm. long, elliptic in cross section; anthers 20-22

mm. long, yellow, centrally aflîxed; pistil stout, triquetrously

knobed at the stigma ; capsules and seeds not seen.

Type—Gentry 19808, Huntington Botanical Garden, San

Marino, Calif., July 2, 1962, deposited in U. S. Nati. Herbarium,

No. 2549708 and No. 2549709 believed to have been collected

Planta acaulis, sing-ularis vel surculosa, grisea, 6-10 dm. alta, 10-20 dm.

lata; foliis 50-100 X 8-12 cm., lanceolatis, acuminatis, adscendentibus, planis

vel conduplicatis, palidis vel glauco-viridibus, dentibus lateralis plerumque

5-10 mm. longis, 1-3 cm. separatim, plerumque retrorso-flexuosis ; spina

terminal! recta, valida, subulata, 3-5 cm. longa, castanea vel maturitate

g-risea;^ scapo inflorescentia inclusa 4-6 m. alto, paniculate, 12-18 ramis

lateralibus; bracteis 5-6 cm. longis deltoideis chartaceis; pedicellis 4-20 mm.

longis, bracteolatis; floribus luteis, persistenti-rectis, congestis, 72-82 mm.

longis, crassis, ovario crasso, 3-angulato, apice incluso 45-50 mm. longo,

viridi; tubo 11-13 mm. longo apice 18 mm. diámetro, 6-angulato, 6-furcato,

infundibuliformi; segmentis rectis, 20-23 mm. longis, 7-8 mm. latis, linearilanceolatis, luteis, involutis, apice obtuso-galeatis ; filamentis validus, ca. 60

mm. longis, ligulatis, base 4-5 mm. latis, 6-8 mm. supra basim tubi insertis;

antheris 20-22 mm. longis, luteis; fructum et semina non vidi.
THE AGAVE FAMILY IN SONORA 123

FIGURE 46.—Agave fortiflora: Leaf and flower cluster X % ; flower section

slightly reduced; cuticular pattern X 18. Drawn from Gentry 19808.
124 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

originally on Sierra Jojoba, 15 miles northeast of El Datil, Dist.

Altar, Sonora, Mar. 23, 1952, Gentry & Fox 11630 (If. & pi.).

The Huntington Botanical Garden tag was lost. Other numbers

consisting of leaf specimens only, assigned here are all from

SONORA : On valley plain, 3 miles northeast of Sierrita de Lopez,

February 1952, Gentry 11618; cerro 4-5 miles north of Bahia

San Carlos, Mar. 29, 1963, Gentry and Arguelles 19881; rocky

hillslopes at north end of Ensenada Grande (=Bahia San Carlos)

Nov. 13, 1964, Feiger & Russell 11U55 '^leaves bluish'^ Cerro de

la Vigia, Guaymas, south side of mountain at l,125--feet alt., Jan.

9, 1965, Feiger & Thomas 11765, ^^generally non-suckering.''

Widely and sparsely scattered on the bajadas and lov^er slopes

of the desert coastal mountains, elevations between 50 and 2,500

feet, in northwestern Sonora. It is a rare plant, the observed

colonies being limited in area and members. The old Papago Indian who guided me to the clone of Gentry 11618 stated that it

was the only one he had ever seen in that valley.

There persists a nagging doubt in my mind regarding the exact

origin of the type plant, which was grown from a small plant

that I collected and sent to the Huntington Botanical Garden.

The identifying tag was lost during the decade of its culture, but

the late Joseph Schneider, a head gardner, believed I had collected

it in Baja California. However, I remember no such plant in Baja

California and it matches none of my collections from the peninsula. The leaves are closely matched, however, by my 1952 Sonoran

number 11630, Unfortunately, despite our recent efforts, neither

Feiger nor I have been able to find flowering plants in Sonora.

This light-gray, glaucous xerophyte is distinct in floral morphology; the large flowers are strong (whence the name, fortiflora) and long lasting, persisting erect in close-set umbels long

after anthesis. The ovary is thick, roundly 3-angulate, with the

grooves from the tepal sinuses furrowing the thick neck. The tube

is broad and bulging at the attachment of the filaments and 3-

angulate, with the rounded corners descending from the outer

tepal bulges. These characters aline it with Trelease's section

Umbelliflorae of Baja California. Its relatives are A. shawii

Engelm. and A. goldmaniana Trel., from which A. fortiflora is

easily distinguished by its simple habit (i.e., not branching from

the leaf axils), its light-gray color, its less open tube, and its erect

rather than spreading tepals. The flattening of the filaments is

extreme in A. fortiflora, making them almost strap shaped and

quite the broadest observed in the genus.

The coastal population of A. fortiflora in the mountains north

of San Carlos Bay consists of a long-leaved, boldly crossed-zone
THE AGAVE FAMILY IN SONORA 125

Vf \'L~ t

PN-2151, PN-2162

FIGURE 47.—Agave fortiflora: Upper, long-leaved form in the mountains

north of Bahia San Carlos; lower, source of Gentry 11618 in valley near

Sierrita de Lopez, western Sonora.
126 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

form, withered and collapsing with protracted drought when

observed in the spring of 1963 (fig. 47, upper). The leaf margins

are unbordered and generally straight to undulate, but the isolated clone or colony found on the plain near Sierrita de Lopez

has small, sharply rising teats under the midteeth (fig. 47, lower).

The flowering season appears to be in summer, since the Huntington Botanical Garden specimen flowered during June and July.

The rosettes observed produced offsets sparingly and old or extensive clonal colonies are rare. The widely scattered members

indicate a freely seeding sexual species. The single plant that

Johnston found at San Pedro Bay (29, p. IfiOi) may have belonged to this species. Locally, the plant is called "lechuguilla^'

and the Papago Indian guide said it was good for pit baking.

Agave zebra Gentry, sp. nov. Figure 48

Rosettes mostly single, medium-size, low spreading, rather open,

light gray, xerophytic, strongly armed, 4-6 X 10-16 dm. leaves

generally 50-80 X 12-17 cm. lanceolate, broadest near the middle,

narrowed above the base, deeply guttered, arcuate, thick, rigid,

light gray glaucous, scabrous, conspicuously cross-zoned, with

strongly undulate margins ; spine 3.5-7.5 cm. long, acicular, mostly

very narrowly grooved above, scabrously decurrent for 5-10 cm.

to upper teeth, yellowish brown to light gray ; teeth large, 25-40

on a side, on prominent teats, those on the midblade mostly 10-20

mm. long, with broad, low, scabrous bases, 1-3 cm. apart, flattened, variously curved and flexed, gray with castaneous tips;

panicles to 6-8 m. tall, narrow, with large bracts towards the shaft

base and 7-14 small laterals in upper 14 to 1/5 of shaft; flowers

small, yellow, 40-55 mm. long; ovary 25-32 mm. long, slender,

cylindro-angulate, straight from angled very short pedicels, neck

slightly constricted and 6-sulcate ; tube funnelform, 6-7 mm. deep,

bulging, deeply sulcate ; tepals about equal, 12-15 mm. long, 5-6

Planta plerumque caule singulo vel caespitosula, 40-60 cm. alta, 100-160

cm. lata; foliis paucis, 30-40, lanceolatis, 50-80 cm. longis, 12-17 cm. latis,

glauco-zonatis, pallidi-viridibus, conduplicatis, reflexis, valde scabrosis;'

margine valde crenata, dentibus supra mammas plerumque 10-20 mm. longis',

reflexis, pallido-brunneis vel albis; spina terminali robusta, 3.5-7.5 cm. longai

leviter flexuosa, albocastanea, supra anguste canaliculata; inflorescentia paniculata, scapo incluso 6-8 m. alta 7-14 ramis remotis; bracteis ad basim

grandis, siccis, triangulis; floribus luteis, ca. 50 mm. longis; ovario cylindricoangulato apice incluso ca. 30 mm. longo; tubo infundibuliformi, 6-7 mm.

longo ;^ segmentis ca. 12-15 mm. longis, aequalibus; filamentis medio tubi

insertis; capsulis parvis, ovatis vel oblongis, 4-5 cm. longis, 1.2-1.5 cm. latis,

stipitatis, brevi-apiculatis; semenibus 4.5-5 X 4-4.5 mm., ovatis, rugosis.
THE AGAVE FAMILY IN SONORA 127

FIGURE 4S.—Agave zebra: Flower cluster X Vs; flower section slightly enlarged; capsules X V2 ; seeds X 2. Drawn from Gentry 21984.
128 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

mm. wide, erect to ascending, lanceolate, revolute, cucullate, the

inner prominently keeled; filaments broad, inserted in midtube,

35-40 mm. long; anthers small, 15-18 mm. long; capsules small,

mostly 4-5 X 1.2-1.5 cm., oblong to obovate, the walls strong and

conspicuously lined, stipitate, apex round, apiculate; seeds 4.5-5

X 4-4.5 mm. obliquely rounded, concave on one or both faces,

the marginal wing very low, hilar notch shallow.

Type—Gentry 2198i, Puerto Blanco, Sierra del Viejo, about 30

miles south of Caborca, Sonora, July 24, 1966, open rocky slopes

of limestone mountain, alt. 2,000 to 2,500 feet, deposited in U.S.

Nati. Herbarium No. 2549706 and No. 2549707. Also from the

same mountain on the northern end, alt. 2,500 to 4,000 feet, Feb.

25, 1951, Gentry 10205 and Apr. 5, 1963, Gentry & Arguelles

19897 (If. & live pi.) ; Puerto Blanco, alt. 1,500 feet, September

6, 1965, Gentry & Arguelles 21207 (fruiting). SONORA: Cerro

Quituni, about 26 miles south of Caborca, Apr. 7, 1963, Gentry

& Arguelles 19899 (If.). (Figs. 49 and 50.)

A, zebra is certainly known from only the two above-named

limestone mountains with a desert climate. Sierra del Viejo, a

north-south trending range about 20 miles long, has a large population of this Agave, Many young flowering shoots were observed

in April 1963, but were still without buds ; July appears to be the

principal flowering month. On Feb. 11, 1894, Mearns collected

Meams 305 in the Tule Mountains along the Sonora-Arizona

boundary. The specimen in the U.S. Nati. Herbarium consists of

two leaves; one is A, deserti, the other suggests A, zebra, having

large, down-flexed teeth on prominent teats. Both leaves were

cut above the base.

The aspects of the leaves (fig. 49) are like those of A. marmorata Roezl of Puebla and Oaxaca, and the lax narrow panicle is

like A. deserti. The flowers collected from four plants of A. zebra

show considerable variation, mainly in size, the proportions of

the main organs being relatively about the same, an exception

being the neck. All the ovaries show a moderate neck constriction,

but when sectioned longitudinally, the carpel cells may reach the

tube bottom level as they taper into the base of the style, as in the

flower drawn by Regina Hughes (fig. 48). In other flowers, the

carpel cells terminate 3-4 mm. below the tube bottom level. The

flower structure shows relationship to those of A. fortiflora, but

are much smaller, are without the long neck, and lack the latter's

long-lasting qualities. The leaves are quite different from those

of A. fortiflora.
THE AGAVE FAMILY IN SONORA 129

PN-21B3, PN-2154

FIGURE 49.—Agave zebra on limestone rubble on Sierra del Viejo, Sonora.
130 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

A. zebra was used for a short while for making mescal, but no

such activity has been observed since 1950. The leaves have a

rather weird aspect; marbled crossbanding on deeply guttered,

outcurving leaves, flagrantly armed with prominent teeth. For

desert gardens it could play the beast to accompany the beauty,

A. pelona, already naturally sympatric but not crossing at all.

i

'¿•i

^-i^^:4*"'^

Vi»

PN-216B

FIGURE 50.—Agave zebra inflorescence with capsules on Sierra del Viejo, in

September.
THE AGAVE FAMILY IN SONORA 131

Agave subsimplex Trel., Mo. Bot. Gard. Ann. Rpt. 22: 60. 1911.

Figure 51

Small, single or cespitese, glaucous, colorful, low-spreading

rosettes, 20-35 cm. tall, 50-70 cm. broad; leaves variable, 12-35

X 3-5 cm., lanceolate to ovate, long-acuminate to short-acuminate,

but little narrowed towards the base, thick, rigid, rounded on

FIGURE 61.—Agave subsimplex: Larger leaf drawn from Gentry 10217;

smaller leaf, capsules X Vs, and seeds X 1% from Gentry 10221; flowers

from Feiger & Bezy 14171, section X 1, cluster X %.
132 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

back, hollowed in inner face, gray glaucous or light yellow green,

or sometimes purple-tinged, the margin nearly straight or strongly

teated ; teeth variable, friable, the larger 3-15 mm. long, straight

or variously flexed, rarely bicuspid, brown or more often yellowish

gray; spine subulate, 2-4 cm. long, not or but little decurrent,

frequently sinuous, shallowly grooved above, glaucous gray ; panicle slender, narrow, 2-3.5 m. tall, with 5-8 short laterals ; flowers

in small umbels, yellow to pink, 40-45 mm. long; ovary about

25 mm. long with a unconstricted long (5 mm.) neck; tube shallow, spreading, 3-4 mm. deep, 10 mm. wide ; tepals 12-15 X 6-7

mm., equal, ascending, elliptic, plane, widest at the middle, apiculate and scarcely hooded; filaments 25-28 mm. long, round in

cross section, inserted below base of tepals 3 mm. above bottom

of tube; anthers 13-15 mm. long, centrically attached; capsules

40 X 15 mm. to 35 X 10 mm., variable, oblong, sometimes narrowly so, light glaucous {Gentry 10221), bluntly apiculate, narrowly or broadly stipitate, the valves thick and striate-nerved ;

seeds (10221) mostly 4.5 X 3 mm., sooty black, roughly lunate,

hylum notch narrow and the opposite corner frequently apiculate,

edges rimmed with a sharp, winglike flange.

Type—SONORA: Seal Island, just off Tiburón Island, Apr. 13,

1911, Rose 16811; Puerto Libertad, May 22, 1939, Gentry U86

(flg.) ; north end of Sierra Colorai, Feb. 28, 1951, Gentry 10217

(If., cap., & pi.) ; Cerro Punta Tepopa, Mar. 1, 1939, Gentry 10221,

10222, 10223 (If., cap., & pi.) ; Kino Point, Nov. 17, 1923, MacDougal & Shreve s,n, ; Punto Cirio, about 7 miles south of Puerto

Libertad, Jan. 3, 1963, Hastings, McClanahan, & Turner 63-8

(If. & cap.)-small leaf with big teats; Tiburón Island, vicinity of

Tecomate, June 14-22, 1951, Whiting 90Í7 (If.) '^Abundant on

rocky mountain slopes on west side of island. This one from NW

corner" ; foothills at north side of Cerro Tepopa, 4 miles by road

west of 9 miles by road south of Desemboque, May 15, 1966, Feiger & Bezy lJfl71 (fl.) ; 6.3 miles south of Desemboque San Ignacio, middle bajada with sandy soil. May 14, 1966, Feiger & Bezy

1U82 (fl.) 'Tew scattered colonies, Ivs. faintly cross-handed,

tepals white to pinkish." Isla Cholludo (Roco Fuca, Seal Island),

Oct. 30, 1958, Feiger 2721 (If. & fl.), ''common & widespread,

suckering, leaves bluish. Inflo, branched, only one plant in fl. ;

anthers yellow, filaments & tepals dustry rose-colored; base of

tube pinkish white." Known only from coastal Sonora, where it

it thinly scattered in small colonies on the outwash slopes of the

granitic and volcanic mountains and the adjacent islands. (Figs.

52 and 55.)
THE AGAVE FAMILY IN SONORA 133

t."

PN-2166, PN-21B7

FIGURE 52.—Agave subsimplex along arid Sonoran coast. Scattered clones

show individual variation.
134 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

This small, xerophytic, dusky agave is closely related to A.

deserti and A. cerulata Trel., as expressed by their common characters of small variable leaves, the nearly tubeless flowers, and

the narrov^ small panicles, v^hich characterize the section Deserticolae. I think the characters outlined in the key to species (p.

45) should be sufficient to distinguish A. subsimplex from

A. deserti, although depauperate plants of the latter in the Lechuguilla Desert of Arizona resemble A. subsimplex. The tv^o species

are not known to cohabit, but it is possible they may be found

together on some unexplored mountains in northwestern Sonora,

such as the coastal Sierra del Alamo. A. subsimplex may be more

closely related to the polymorphic A. cerulata of Baja California,

as indicated by the small, variable leaves and the narrow, oblong,

waxy capsules. However, the Sonoran plants have more spreading

rosettes, gray not yellow, with thicker, wider, less acuminate

leaves, and the filaments are inserted below the base of the tepals.

In A. subsimplex the nectarious interliner of the tube, from the

rim of which the filaments ascend, does not completely fill the

tube, as it does in A. cerulata. The flowers of A. subsimplex are

smaller than those of A. cerulata, 40-45 mm. vs. 45-60 mm. The

whole section Deserticolae around the Gulf of California needs

intensive field study.

The Seri Indians, a maritime hunting and gathering people of

the Sonoran coast, called this agave ^^ahmmo," as nearly as I could

render the sound, and stated that it was gathered for cooking

and eating. Whiting noted the Seri name as "den;kl." Small

amounts of sapogenin (0.07-0.14 percent) were found in the

leaves. OflFsets that were collected for the Huntington Botanical

Garden and my own live collection did not survive.

Agave deserti Engelm., Acad. Sei. St. Louis Trans. 3: 310, 370.

1875.

Agave consociata Trel., Mo. Bot. Gard. Ann. Rpt. 22: 53. 1911.

Medium-size, light-gray, freely suckering rosettes mostly 80-40

cm. tall, 40-50 cm. in diam ; leaves variable, mostly 25-40 X 6-8

cm., lanceolate to linear-lanceolate, scarcely narrowed above the

broad clasping base, acuminate, gray glaucous to bluish glaucous,

often cro3s-zoned, thick, rigid, concave above, convex below, usually regularly armed with slender-cusped teeth, from small, the

larger 2-3 mm. long, to large, the larger 6-8 mm. long, gray,

loosely attached, mostly 15-30 mm. apart; spine strong, generally

2-4 cm. long, light brown to grayish, openly grooved above, decur-
THE AGAVE FAMILY IN SONORA 135

rent to the first or second tooth above; panicles generally 2.5-4 m.

tall, on slender shafts with scarious triangular bracts 8-15 cm.

long and 6-12 short laterals with small umbels in upper 14 to

% of shaft; flowers yellow, 40-53 mm. long; ovary 22-35 mm. long

with a slightly narrowed neck 4-6 mm. long, greenish ; tube shallow, spreading, 4-5 mm. deep, 12-15 mm. wide, lined with a thick

nectiferous disk; tepals equal, 14-19 mm. long, 6-8 mm. wide,

light yellow, broadly linear, spreading at anthesis, rounded and

abruptly hooked inward at apex; filaments 25-35 mm. long, inserted at base of tepals ; anthers 13-18 mm. long, yellow ; capsules

ovoid to oblong or obovoid, mostly 3.5-5 cm. long, 1.5-1.8 cm.

wide, thick-walled, short-stipitate ; seeds black, 4X5 mm.

Type—Based on Emory in 18Í6 and Hitchcock & Palmer in

1875 collected on Rancho San Felipe, San Diego County, California. SONORA: At km. 2,509 in mountain pass southeast of

Quitovac, alt. 1,800 to 2,000 feet, on granitic slope, Sept. 4, 1965,

Gentry & Arguelles 21203 (fruiting) (fig. 53). This is the only

Sonoran collection observed. It has been collected in Arizona near

the Sonoran border ; Cabeza Prieta Mountains, Gentry 20590, and

in the Tule Mountains, Mearns 305, The latter is about 100 miles

from the collection southeast of Quitovac, where the plants are

sparsely but widely scattered. Other populations of the species

are to be expected on the rocky pediments of the intervening

mountains in Sonora. (Fig. 54, upper.)

The Lechuguilla Desert, one of the most arid regions in North

America, takes its name from the small pauperized A. deserti

found thinly scattered over the rocky mountain slopes of this part

of southwestern Arizona. The leaves commonly are only 20-25 cm.

long. Other than the small size, the characters of the plants are

quite like those of the species; rather broad, gray, crossbanded,

thick, rigid leaves and slender, small panicles. As with other

Arizona populations, the rosettes sucker more sparingly than the

rosettes of California colonies, and the large cespitóse Californian

clones are absent. Although the extensive population of A. deserti

at the type locality along the arroyo San Felipe is morphologically

homogeneous, there are other localities showing wide variability

in leaf forms. One of these localities is Piñón Flats by the northwestern slopes of Santa Rosa Mountain along route 74 in southern

California, and another, described by Ivan Johnson (29) is on

Angel de la Guarda Island. Some of the leaf variants to be found

at Piñón Flats are shown in figure 54, upper. The sources of such

genetic turbulence may be due to hybridization, but if so, the

parents involved have not been detected. A consociata Trel. is only
136 AGRICULTURE HANDBOOK NO. 399, U.S. DEFT. OF AGRICULTURE

PN-2158, PN-2159

FIGURE 53.—Agave deserti in granitic hills southeast of Quitovac, Sonora.

Organ pipe cactus and acacia tree in baclîground.
THE AGAVE FAMILY IN SONORA 137

PN-2160, PN-2161

FIGURE 54.—Upper, Agave deserti leaves from population around Pinon Flats,

Southern California. Not an unusual range of variability for an Agave

species; lower, Agave fourcroydes in Obregon plantation southeast of

Ciudad Obregon, Sonora.
138 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

one of the variant forms and has no value as a specific name.

Cave reports (Madroño 17: 166. 1964) a collection from Baja

California, Hutchinson 710, with chromosomes n=59, indicating

polyploidy in this species. As far as chromosomes have been reported for the genus, 30 is the basic number, typified by a karyotype of 5 long and 25 short chromosomes.

A, deserti is known to have been eaten by the Arizona Indians,

e.g., the Yumans and Papagos (<^, p. J^9). It may still be eaten by

the Papagos in northwestern Sonora. San Francisquito is an important Papago village within the probable range of the species.

The plant contains little or no sapogenins and I find no report of

medicinal uses. It is important to wildlife ; birds visit the flowering stalks for nectar and insects. Small rodents live around the

plants and are protected by the armed rosettes. The pack rats,

Neotoma, make nests among the clumps of rosettes, gnaw through

the leaves, and eat the flowers and perhaps the seeds. Wild bighorn sheep, like cattle, eat the flowering shoots. As a wildlife resource growing in our most arid of deserts, A. deserti deserves

conservation and perhaps should be established by planting on

desert mountains where it is now lacking.

Agave fourcroydes Lern., 111. Hort. 11: 65.1864.

Agave sullivani TreL, U.S. Nati. Herbarium Contrib. 23: 119. 1920.

Large, sword-leaved rosettes developing a thick stem 1-1.7 m.

tall; leaves straight, rigid, 1-1.8 m. long, 8-12 cm. wide, linear,

thickly rounded at base, guttered, acuminate, the margins straight

with regularly spaced, slender, dark brown teeth 3-6 mm. long;

spine stout, conical, mostly 2-3 cm. long, dark brown, openly

short-grooved above at the base; influorescence an open panicle

5-6 m. tall with 10-18 laterals in upper half; flowers greenish

yellow, 60-70 mm. long; ovary 35 mm. long; tube 15-16 mm. deep,

15 mm. wide; segments subequal, 17-16 X 3 mm., wilting and

twisting before or at anthesis, sharply reflexed when dry; filaments 55-60 mm. long, inserted 8 mm. above bottom of tube ; an-
THE AGAVE FAMILY IN SONORA 139

thers 20 mm. long. (Measurements from dried flowers of Berger

6-VIII-09, U.S. Nati. Herbarium.)

The typical plantation form of this fiber resource, ^'henequén/'

is still (1971) represented in Sonora by the Obregon plantation

12 miles southeast of Cuidad Obregon, Gentry 11361, The planting

stock was reported to have been obtained from the Pericos plantation, Sinaloa, which in turn was reported by the owner-manager

of the Pericos plantation to have been brought from Yucatan in

1890. In 1951, the Obregon planting was reported to be about 25

years old and occupied several hundred acres (figure 54, lower).

However, it had never been harvested except in a desultory handlabor way by one or two resident families. The 15-18 inches of

annual rainfall was apparently not sufficient to produce a large

enough leaf for first-quality fiber.

On several occasions I have searched the panicles for capsules

with seeds but without success. One would expect that the related

A. pacifica Trel., which is in the same section of the genus and

which occurs in the surrounding coastal thorn forest, would crosspollinate occasionally, but no instance was found. The Obregon and

similar plantings probably represent a sterile clone, perhaps of

hybrid origin, which has been transported for fiber planting

around the world.

Propagation is made by transplanting the rhizomatous offsets

produced during the early years of growth. They are preferred

over the bulbils of the inflorescence because they grow more vigorously and require fewer years to reach the commercial production

stage. Fiber as a cottage industry from the Obregon plantation

goes into local trade.

The leaves contain less than 1 percent sapogenins, which as a side

product from the bagasse of fiber operations in eastern Mexico and

elsewhere has been utilized for synthesizing cortisone and some of

the sex hormones. The plant occurs widely as a backyard cultivate

or fence plant. It was found at El Triumfo, Cape District, Baja

California, Gentry 11268, and I find it unseparable from A.

sullivani Trel. described from La Paz in the same area.

For distribution of the Deserticolae and the Ensiformae sections

of agave, see figure 55.
140 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OP AGRICULTURE

FIGURE 65.—Distribution of Agave in Sonora, sections Deserticolae and Ensiformae, represented by their ideographs. Ideographs with numbers indicate flower collection at the locality occupied. (See p. 37.)
THE AGAVE FAMILY IN SONORA 141

Agave rhodacantha Trel., U.S. NatL Herbarium Contrib. 23: 117.

1920. Figure 56

Large, single or cespitóse, multileaved, radial rosettes 2-3 m.

tall, 3-5 m. in diameter, truncate, 5-9 dm., or acaulescent; leaves

FIGURE 56.—Agave rhodacantha: Leaf and flower umbel X ^/4 ; flower sections

slightly enlarged. Drawn from Gentry and Gilly 10704.
142 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

1.4-2.5 m. X 8-15 cm. linear, hard-fibrous, rigid, straight, much

thickened and scarcely narrowed at base, green to faintly glaucous

green, smooth, the margin straight to undulate; teeth regular,

mostly 4-8 mm. long, firm, slender, very sharp, ñexed upwards,

dark brown, mostly 1-3 cm. apart; spine 1-2.5 cm. long, conical

but frequently with a subulate tip, dark brown, and with a short

open groove above; panicle 7-9 m. tall, deep, broad, with 30-45

large remote laterals; flowers green with tepals yellowing at

anthesis, 55-65 mm. long (dried & relaxed) ; ovary 25-35 mm.

long, including short neck, fusiform; tube urceolate, 8-10 mm.

long, 9-10 mm. wide; tepals subequal, 16-23 mm. long, 4 mm.

wide, linear, strongly involute, rounded and strongly hooded at

apex, wilting at anthesis and soon reflexing; filaments 40-45 mm.

long, inserted 5-7 mm. above bottom of tube ; anthers 14-28 mm.

long; capsules 7-8 cm. long, 2.5-3 cm. wide, oblong, shortly

beaked long-stipitate, the stipe 10-15 mm. long; seeds large,

oblique, 8 X 10 mm., the curved margin not winglike, the hylum

notch shallow.

Type—Lundstrom in 1909, Mocorito, Sinaloa, in Mo. Bot. Gard.

Herbarium, St. Louis (reported as lost). SONORA: 5 miles northwest of Conejos, along Arroyo Guajaráy north of San Bernardo,

Dec. 22-25, 1951, Gentry 11376. Cultivated in Tupeyeca, Rio Mayo,

originally from Sierra Charuco, Gentry, Barclay & Arguelles

20Í79.

Trelease's description is very brief. The above description is

based primarily upon 3 flowering specimens collected in Nayarit :

1-2 miles southwest of Jalisco, on lower slopes of Cerro San Juan,

July 9, 1951, Gentry & Gilly 10806; 1 mile east of Mazatan along

road to Compostela, July 6, 1951, Gentry & Gilly 10786; 7-8 miles

west of Compostela, along road to Mazatan, June 27, 1951, Gentry

& Gilly 1070J^ (fig. 56). Widely scattered along the more moist

mountain slopes at elevations from 2,000 to 3,500 feet from southern Sonora at least to about Bahia Banderas, Nayarit. It is rare in

Sonora.

The very long, rigid leaves and large inflorescences with large,

long-stipitate capsules distinguish this agave from its near relative, A. pacifica. However, leaf size and armature of A. rhodacantha can become quite variable and some of the forms with small

distal teeth are inseparable in themselves from A. pacifica, like

Gentry & Gilly 10756, 1 mile east of Mazatan along road to

Compostela, Nayarit, but this collection has the long-stipitate

capsules of A. rhodacantha. It is not known if the two species

hybridize, but some of the variants suggest they do. Some young
THE AGAVE FAMILY IN SONORA 143

plants received from the Tres Marias Islands, Gentry 11565,

11567, appeared to represent A. rhodacantha and are to be identified with the "giant Agave'' referred to by Nelson (North Amer.

Fauna 14: 7-13. 1899); Maria Madre Island, May 3-25, 1897,

Nelson i26i (cap. & If.). During the 1940's and 1950's it was

being exploited on Maria Madre Island for its long fibers with

labor of the convicts incarcerated there.

This western long-leaved population of the section Ensiformae

eventually may be found cospecific with eastern Mexican taxa,

including A, gutierreziana described by Trelease from Oaxaca, a

name of the same date as A. rhodacantha. Other long-leaved taxa,

A. elongata Jacobi, A. candalahrum Tod., and A. spectabilis Tod.

appear closely related, but they are light glaucous and appear to

belong with A. angustifolia Haw., an older name.

I once made a search for A. rhodacantha at Mocorito, the type

locality, but found nothing. Lundstrum quite possibly brought

the plant in from a nearby mountain or perhaps the planting, if

such existed, has been destroyed.

Agave pacifica Trel., U.S. Nail. Herbarium Contrib. 23: 118.1920.

Figure 57

Agave yaquiana Trel., op. cit.: 120.

A. owenii I.M. Jtn., Calif. Acad. Sei. Proe., ser. 4, 12: 999. 1924.

Short-stemmed, multileaved, freely suckering, sword-leaved,

radial rosettes, 1-1.5 m. tall, 1.5-2 m. diameter; leaves mostly

50-120 X 4-8 cm., linear to linear-lanceolate, rigid, straight to

falcate, ascending, guttered, green or glaucous green to yellowing

green, margins nearly straight; teeth regular, mostly 3-6 mm.

high, 15-30 mm. apart, dark brownish to black, the cusps flexed

upward ; spines 15-20 mm. long, dark brown, flattened above at the

base, otherwise conical, the tip short or subulate ; panicle 3-6 m.

tall with 6-20 horizontal to ascending, short, trifúrcate laterals in

upper 1/3 to 14 of shaft; bracts triangular, 5-12 cm. long,

appressed, scarious ; flowers greenish yellow, glaucous lavender to

brownish red or pale green in the bud, 45-65 mm. long, the tepals

wilting before or at anthesis, reflexing; ovary 20-30 mm. long including the short neck, fusiform, 6-grooved at apex, pale green ;

tube 6-12 mm. deep, 10-13 mm. in diameter, 6-grooved from the

tepal sinuses, pale green; tepals subequal, 15-24 mm. long, 4-5

mm. wide, appearing narrower with involution, linear, withered

and sharply reflexed in post-anthesis ; filaments 35-45 mm. long,

inserted in midtube ; anthers 18-30 mm. long, eccentrically afiixed ;

capsules 35-50 X 20-26 mm., ovoid to oblong, short-stipitate.
144 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

FIGURE 57.—Agave pacifica: Leaf drawn from Gentry 19875; leaf section

from Gentry 10280; flowers from Gentry 19875, cluster X %, section X

1 ; capsules X Vs and seeds X 2 from Gentry 10278.
THE AGAVE FAMILY IN SONORA 145

short-beaked at apex; seeds large, 8-10 X 5-8 mm., shiny black,

oblique, with a broad outcurving wing or margin, hylum notch

very shallow or lacking.

Type—Trelease in 1904, from Crestón Island, Mazatlan,

Sinaloa, Mexico, in Mo. Bot. Gard. Herbarium, St. Louis.

SONORA: Between "Ures & Hermosillo, 8/18/00,'' Trelease 391

(type of A, yaqidana) ; islet in Guaymas Harbor, Apr. 14, 1921,

Johnston 3085 (type of A. owenii) ; Gentry 113Í6; 4 miles north

of Rancho Tinaja, Alter District, Feb. 27, 1951, Gentry 10212

(in bud) ; 8 miles east of Ures, Mar. 11, 1951, Gentry 10280; 6-8

miles northwest of Bacadéhuachi, May 21, 1957, Gentry 166H;

near Navojoa, Mar. 28, 1963, Gentry & Arguelles 19875 (fl.) ; San

Bernardo, Rio Mayo, Gentry & Arguelles 10258; Sierra Tecurahui,

District de Alamos, Oct. 26-28, Gentry, Barclay, & Arguelles 19359

(fl.) ; Sonora-Sinaloa border along highway. Gentry, Barclay, &

Arguelles 19656,19657 (fl. only) ; near Alamos, March 1910, Rose,

Standley, & Rüssel 128^7, 1295i; vicinity of Empalme, Rose,

Standley, & Rüssel 12260 (If. & cap.) ; 7 miles north of La Colorado

Mina San Fernando Road, May 5, 1948, Wiggins 116W (If. & fl.) ;

Peon, between Rio Yaqui and Empalme, Wiggins 61^68; vicinity of

Hermosillo, Rose, Standley, & Rüssel 12JfS0; Camoa, Rio Mayo,

Jan. 25, 1899, Goldman 302 (If. & fl.) ; Tonichi, hills above Rio

Yaqui, Sept. 8, 1941, Spicer s.n, ; between Nacori Grande and

Reveico, 10 miles from Nacori Grande, District Sahuaripa, alt.

about 2,350 feet, Nov. 14, 1953, Feiger, Soto & Lenks 229 (If. &

fl.) ; 40 miles south of Navojoa and 15 miles north of Sinaloa

borders, coastal plain, Mar. 9, 1960, Feiger & Stronck 3139 (If. &

fl. Tepals are unusually long, 28 mm., tube 12 mm., dried fl.)

'Terianth chartruse, stamens yellow-green ; inflo. 10-12 feet talF' ;

near Sabino, 10 miles southeast of Alamos, short-tree forest, Jan.

22, 1961, Feiger 5050 (If., fl., & cap.) ^^Inflo. ca. 15 feet high; fls.

greenish, Ivs. bluish green'' ; 9 miles south of Torres, Oct. 22, 1965,

alt. 825 feet, Hasting & Turner 65-18J, (If.). SINALOA: Capadero. Sierra Tacuichamona, alt. 3,000 feet, rocky, sunny hill slope

in oak woodland, Feb. 13, 1940, Gentry 5630.

This agave is characteristic of the coastal thorn forest of Sinaloa

and southern Sonora and is also widely scattered in the shorttree forest of the Sierra Madrean foothills below the oak forests ;

see distribution map, figure 55. It is thinly scattered in the

arborescent desert of northwestern Sonora. It occupies the sandy,

gravelly, and rocky soils of plains and mountains from sea level

to 3,500 feet. It is the most common agave of both Sonora and

Sinaloa (fig. 58). The only agave in this region likely to be con-
146 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

PN-2162, PN-2163

FIGURE 58.—Agave pacifica: Upper, at its northern limits near Rancho Dátil;

lower, at Bacanora still, wild plants transplanted show variability of

seeding population.
THE AGAVE FAMILY IN SONORA 147

fused with A. pacifica is A, aktites a smaller, bluish-leaved, bigheaded, member of the section Ensiformae (Rigidae of Berger),

growing on the maritime sands of Isla Lechuguilla, Sinaloa, and

the adjacent mainland. A. pacifica is related to the A. angustifolia

complex of eastern Mexico and Central America. A, pacifica^ A.

collina Greenm., A. elongata Jacobi, A. kirchneriana Berger, A.

bergeri Trel., and others too numerous to mention here, form a

most difficult area of agave taxonomy yet to be systematically

resolved.

There is a rather even spread of variability through the extensive populations of A. pacifica. The coastal population have shorter

leaves with a wide midspread, while the more interior populations

have longer, narrower leaves and may become more lax in forest

shade, with a concomitant lengthening of stem. The northern

Sonora form is more xerophytic with narrow, stiff, heavily armed

leaves (A, yaquiana) but the dark-brown, upward-flexed, slendercusped teeth, although variable, are characteristic throughout.

A, owenii is a local form with low, broad, blackish distal teeth and

a short, conic spine. The teeth on Johnston's type are at most 5-6

mm. long, not *'20-35 mm.'' as was printed (loc. cit.). The flower

varies in size and ratio of tube to tepals, but there is no correlation

of floral variability with leaf difference. Flower color varies during

development and between individual plants.

Large, black, viable-looking seed is regularly produced, bulbils

are not uncommon, and rhizomatous suckers are widely deployed

around the older rosettes. Individual sexual variability, therefore,

occurs between the widely scattered thickets, which are composed genetically of identical individuals. The flowering season

starts in November, especially in the south, and continues until

March. The inflorescences in the north are frequently delayed by

the lower winter temperatures. The nectariferous flowers attract

many insects during the spring dry season, and judging by the

many birds observed, the high inflorescence is also an attractive,

sustaining perch. The firm, sharp teeth and spines are woundingly

repellent to man and other large mammals, but protective to

rodents and rabbits. Altogether, the plant is a wildlife asset.

The leaves have a thick, strong fiber, comparable in tensile

strength to henequén, and are used locally for cordage. The famous

"mescal bacanora" is made from the population growing wild

about the settlement, Bacanora, near Sahuaripa in eastern Sonora

(fig. 58). This mescal has a distinctive flavor not matched by any

others even when it is made from other populations of the same

species farther south. Whether this flavor is due to inherent

genetic constitution, to local soil or other écologie factors, or to
148 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

the kind of bacteria or method used in the primitive still has not

been determined. Mescal bacanora continues to be an epicurean

product among the northwestern mescal drinkers. Were it properly aged before being sold, it should find a distinctive place among

the fine tequilas of Mexico.

Analyses of several samples showed the leaves to contain little

or no sapogenins, the highest found having 0.2 percent of

tecogenin, hecogenin, and chlorogenin; Gentry 11337 from

Cieneguita, 40 miles north of Guaymas, Sonora. The heads have

been pit baked and eaten but are inferior to lechuguilla ceniza and

jaiboli, according to local informants. Following are some notes

concerning this agave, taken during a journey up the Arroyo

Guajaray in December 1951.

A. pacifica was observed frequently all along the trail from

Chorijoa to Conejos in the short-tree forest; commonly as scattered individuals or in groups of 2 to 4 or 5, rarely crowded

clumps of a dozen. Inflorescences in the bud were common and a

few flowers were opening beyond Conejos on the higher slopes;

flower buds brownish red to dull green. Flowers are greenish

yellow at anthesis. In this area with elevations from 2,000 to 3,500

feet the species is abundant. Common names are mague and

mescal.

Agave aktites Gentry, sp. nov. Figure 59

Small, glaucous, surculose rosettes with broadly globose stems,

4-7 X 6-11 dm.; leaves 40-60 X 2-3 cm., linear, straight, patulous, unequal, nearly flat above, concave below, broadly clasping

at the base, bluish glaucous gray, sometimes cross-zoned, smooth

or asperous; spine short, 12-20 mm. long, broad and flattened

above at the base, abruptly subulate, dark brown to grayish ; teeth

3-5 mm. long, mostly 3-5 cm. apart, dark brown, generally upcurved with subulate flexuous tips; panicle 3-4 m. tall, narrow,

with 10-14 short laterals in upper third of shaft. Flowers and

capsules unknown.

Type—Gentry 11Í70, Isla Lechuguilla, north of Topolobampo,

Sinaloa, Mexico, Jan. 7, 1952, on sand dunes with coastal thorn

forest, deposited in U.S. Nati. Herbarium No. 2539974. Also from

Sinaloa: 8 miles southeast of Los Mochis in coastal thorn forest

Planta caulis globosis surculosa, 4-7 dm. alta, 6-11 dm. lata, species

foliorum anguste glaucedine insignis ; foliis 40-60 X 2-3 cm. minerais patulis

inaequalibus, plerumque trans-zonatis ; dentibus lateralibus 3-5 mm. longis

antrorso-flexuosis 3-5 cm. separatim, badius; spina terminali valida supra ad

basim plana ad apicem subulata, 12-20 mm. longa; scapo inflorescentia inclusa

3-4 m. alta, paniculata, 10-14 ramis lateralibus; flori et fructum non vidi.
THE AGAVE FAMILY IN SONORA 149

PN-2164

FIGURE 59.—Agave aktites on sandy island of Lechuguilla. Leaves of this

plant were cut for analysis sample and for herbarium specimens, exposing the broad globose heads, or "cabeza."

near sea level, Jan. 4, 1952, Gentry 1H23. SONORA: Near el faro

in Yavaros, on coastal sand dunes with Jatropha, Crotón, Opuntia,

Oct. 19, 1966, Gentry & Arguelles 22015 (If., photo.).

Although the fruit and flower of this plant are unknown, it

appears to belong in the section Ensiformae. It is distinguished

from the more common A. pacifica by its smaller size; broad, short

globose stem; light-bluish, glaucous, narrow leaves; broad, flat

spine; and narrow panicle. Its small size, remote teeth, and geographic position suggest affinity with A. datylio across the Gulf

of California in the Cape District. Its taxonomic position will become clearer when flowers and fruits can be obtained.

The specific name means coast dweller, from the Greek word

aktes, seashore. A. aktites is known only from the coast of northwestern Sinaloa and northwestern Sonora, where it is scattered

along the mainland littoral and upon the adjacent sandy islands

of Santa Maria and Lechuguilla, the latter taking its name from

the local name for the plant. On Isla Lechuguilla, the plants were

found upon the stationary dunes, scattered in colonies of small

bluish rosettes with large round cabezas (fig. 59), the numerous

leaves stiffly ascending, flat above, rounded below, and with ffaring

bases. The plants suckered out for 3-6 feet in the loose sand. My
150 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

guide, Carlos Preciado, stated that his people (Mayo Indians) had

pit baked as many as 40-50 cabezas at one time on that island.

They are sweet, well flavored, and superior to those of other

agaves on the mainland. Some people had taken small plants from

the island to plant about their houses on the mainland. The guide

also stated that Isla Santa Maria had many more of this agave

than Isla Lechuguilla. Analysis of the leaves of both island and

mainland forms showed only negligible amounts of sapogenin.

Agave schlechtendalii Jacobi, Hamburger Garten-Blumenzeitung

20: 555.1864.

Of the many agaves named by Jacobi, schlechtendalii is the only

one he attributed to Sonora, stating (loc. cit. 22:169. 1866.) that

it was first introduced to Europe by the Botanical Garden in Gottingen. He described the leaves as 7-8 inches long and 2 inches

wide in the middle. The plants were immature.

Berger wrote (5, p. 162) that he had observed two forms of the

plant Jacobi had reference to for several years in the Botanical

Garden at La Mortola. Berger described the leaves as 65 X 12-13

cm., but had not seen the plants flower. He thought the plant was

related to A. aw.ericana.

I do not recognize this plant from Jacobi's or Berger's description, nor do I see any way of applying Jacobi's name to Sonoran

or other agaves.

MANFREDA SALISB., GEN. PL. FRAGM. 78. 1866.

Low perennial herbs with fusiform succulent roots, fleshy underground stems or bulbous rootstocks, and short-lived, pliant, unarmed leaves in 1-several sessile rosettes ; leaves without a terminal spine, the margins entire or finely serrulate, frequently crisped

or undulate, weak, wholly green or spotted reddish or purplish,

mostly reclínate; inflorescence spicate, the flowers mostly single,

lax, proterandrous, the tube cylindrical, usually narrow and deep ;

segments equal or subequal, reflexed or outcurving at anthesis,

green, yellow, to reddish or purple ; filaments mostly long-exserted ;

anthers versatile ; pistil long exserted, cylindrical, tubular, with a

clávate, 3-lobed stigma ; ovary fleshy, 3-celled ; fruit an erect capsule, apically dehiscent; seeds numerous, flat, lunate or oblique,

black. M. virginica (L.) Salisb.

The genus Manfreda was recognized by Rose, Trelease, and

Small, but European horticulturists, including Berger, the monographer of Agave, combined it with Agave as a subgenus. American
THE AGAVE FAMILY IN SONORA 151

floristic men have generally included it with Agave. The writer

believes that this group of small herbs constitutes a natural generic

group, separable from Agave by their unarmed, perennial, tender,

reclinate leaves, bulbous leaf bases, fleshy underground stems,

succulent fusiform roots, and lax spicate inflorescences. Some

seedling agaves have succulent roots, but all adult plants examined

have only wirelike roots about the base of the plant. These main

roots later branch out into feeder rootlets. The flowers of Manfreda are quite variable. The longer tubular form with its midtube flexure, as in M. virginica, is quite different from any Agave

flower, but some other Manfreda flowers with short tubes resemble flowers of some of the agaves. It is, therefore, the sum of

the characters that distinguishes Manfreda from Agave, Manfreda is closely related to Polianthes, another herbaceous genus;

see remarks on page 153.

The presence or absence of reddish spots on the leaves has been

used for separation of species, but this character is generally

valueless as a specific criterion. Many Manfreda populations show

forms with or without leaf spotting and without other correlative

characters for separating them. Leaf spotting appears to be a

Mendelian character, segregating at random and representing a

gene complex common to several species.

Flower tube much longer than the tepals, curved ; filaments shortly

exserted, not surpassing the tepals M, singuliflora

Flower tube shorter than the tepals, straight; filaments longexserted, at least twice as long as the flowers.

Leaves long and narrow, about 1 cm. wide M. jaliscana

Leaves short and broad, 3-6 cm. wide M. planifolia

Manfreda singuliflora (S. Wats.) Rose. U.S. Nail. Herbarium

Contrib. 8: 16. 1903.

Bravoa singuliflora S. Wats., Amer. Acad. Arts Proc. 22: 479. 1887.

Small sessile rosettes, perennial from tuberous bases and succulent roots; leaves few (4-8), reclinate, grasslike, linear, mostly

20-30 cm. long, 7-10 mm. wide, striate, with a thin, smooth or

minutely denticulate, sometimes whitish margin ; spikes 8-12 dm.

tall, long-pedunculate, 8- to 10-flowered near apex ; flowers single,

purplish, remote, short-stipitate ; ovary 8-10 mm. long at anthesis ;

tube very slender, about 30 mm. long, curving, gradually widened

towards apex; tepals recurved at anthesis, 10-14 mm. long; filaments exserted 5-10 mm. beyond tube; anthers 7-8 mm. long;
152 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

pistil exserted 8-10 mm. after anther dehiscence ; capsules orbicular, 15 mm. in diameter, short-stipitate, depressed at apex.

Type—Pringle 1S81, mountains near Chihuahua City, Chihuahua, Aug. 19, 1887. There is one collection from the ChihuahuaSonora border—Sierra Charuco, Rio Mayo, Oct. 1, 1936, Gentry

2915, on a rocky slope with pines. Other collections from adjacent

Chihuahua indicate Manfreda singuliflora as widely and thinly

scattered, associated with pine and oak on the higher slopes of

the Sierra Madre mountains at elevations between 5,000 and 7,000

feet. There is little doubt that the plant occurs along the Sonoran

side of the border in this habitat. It is not an abundant or conspicuous plant and may be overlooked.

M, singuliflora is distinguished from other manfredas by its

very slender, long, curved, tubular flower and the bulbous underground base. It lacks the fleshy stem between the bulbous leaf base

and the succulent roots common to many other manfredas. The

filaments are much shorter than those in most species of the genus.

Manfreda jaliscana Rose, U.S. Nail. Herbarium Contrib. 7: 22.

1903.

Low perennial herb, single or cespitóse, with grasslike, reclínate

leaves; leaves 40-60 cm. long, about 1 cm. wide, conduplicate,

striate, green, smooth, with a narrow pale margin, fraying toward

the tip ; old dry leaf bases of other years persisting ; spike 12-16

dm. tall, with lower bracts leaflike, the upper smaller, scarious,

fragile; flowers terminal, lax, mostly single, greenish yellow (dried

specimens) ; ovary 8-10 mm. long, fusiform; tube 7-8 mm. long,

narrowly funnelform; tepals 10-11 mm. long; filaments 50-60

mm. long, purplish; anthers 8 mm. long; pistil exserted 50-60

mm.; capsules whitish, 12-15 mm. long, short-stipitate, truncate

and short-apiculate.

Type—Pringle 1850, grassy slopes of the barranca near Guadalajara, Jalisco, December 1889. SONORA: Canyon Sapopa, Rio

Mayo, on oak slope in rocks and grass, Feb. 8, 1935, Gentry 1290

(fi., If. & caps.) ; Curohui, Rio Mayo, in loose black loam of oak

forest slope, alt. about 3,000 feet, Apr. 4, 1938, Gentry 36Í0 (fi.).

SINALOA : Mesa Malqueson, Cerro Colorado, rocky slopes on oak

savanna, alt. 2,500 feet, Dec. 8, 1939, Gentry 5183 (caps.). It

occurs widely scattered on the deeper soils of the grassy oak

slopes at elevations between 2,500 and 4,000 feet from southern

Sonora to Jalisco.

Unless fiowering, the plants may go unnoticed because they

resemble clumps of grass. In Sonora and Sinaloa, the people call it
THE AGAVE FAMILY IN SONORA 153

'^mescalito/' but gave me no special use for it. M. jaliscana appears

to be endemic to the western slopes of the Sierra Madre Occidental.

Manfreda planifolia (S. Wats.) Rose, U.S. Nail. Herbarium

Contrib. 7: 22. 1903.

Agave planifolia Wats., Amer. Acad. Arts Proc. 22: 478. 1887.

Perennial rosette herbs with broad leaves ; leaves mostly 10-20

cm. long", 3-6 cm. wide, ovate-lanceolate, fraying at the tips,

broadly clasping at base, finely serrulate; spike 10-12 dm. tall with

whitish, remote, scarious, triangular-apiculate bracts and 8-12

flowers in terminal Vs of shaft; flowers (dried) 30-45 mm. long;

ovary broad, neckless, protruding into tube; tube short, broad,

8-9 mm. deep ; tepals 15-20 mm. long, broadly linear, obtuse, reflexing at anthesis; filaments very long, 6-8 cm.; inserted 5-6

mm. above base of tube.

Type—Pringle llJfl, warm sandy or gravelly banks near

streams in canyons of the Mapula Mountains, Chihuahua, Oct. 19,

1886. SONORA: West slope of the Sierra Madre, east of San

Bernardo, Rio Mayo, Gentry xo. Specimen consisting of a flower

only, brought to camp by an ''arriero'' (muleteer) in oak woods

country.

POLIANTHES L., SP. PL. 1: 316. 1753.

Small perennial herbs with succulent leaves on short bulbous

stem; leaves radical, unarmed, seasonal; inflorescence bracteate,

racemose, lax, the flowers zygomorphic, bisexual, commonly in

pairs; perianth tube cylindrical to funnel-form, expanded above

the middle, curved, much longer than the segments; stamens 6,

included in perianth, inserted on sides of tube; anthers linear,

erect, dorsifixed; ovary 3-locular; ovules numerous, flat, with a

lax testa ; fruit an ovoid capsule.

This is a small American genus, apparently endemic to Mexico,

consisting of about 12 species. A sketchy account of them was provided by J. N. Rose in 1903 (U.S. Nati. Herbarium Contrib. 8:

8-14). There is still little material available for study. The genus

Polianthes is closely related to the genus Manfreda, but the

zygomorphic flowers with ampliate tubes and included vertical

anthers appear to form a natural generic group to themselves.

Shinners has recently submerged Manfreda under Polianthes

(SIDA 2 (4) : 333-338. 1966), along with several other closely

allied genera, but before I can concurr, I would prefer to study
154 AGRICULTbRE íIANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

the various respective organs, both floral and vegetative, of these

little known taxa.

Polianthes tuberosa L., Sp. PI. 453.1753.

Rosette herb with green, succulent but thin, outcurving leaves,

30-45 cm. long, linear-lanceolate, channeled below, plane above,

frequently brown-spotted on the back, margins smooth ; inflorescence laxly spicate, 6-9 dm. tall with leaflike bracts on the

peduncle; flowers geminate, waxy white, 3.5-6 cm. long; segments

much shorter than the tube, spreading, oblong-lanceolate; filaments 4-5 mm. long, included, inserted on upper part of tube;

flowers strongly fragrant.

The plant is known only as a cultigen and its origin is generally

regarded as Mexico, where it is called "nardo.'' The English name

is tuberose. It is now cultivated for cut flowers in many parts of

the world with more benign climates, as it is only ''half-hardy."

Judging by the great masses of nardo flowers seen in the markets

of Mexico on fiesta days, it must be very popular as a cut flower.

They have good lasting qualities and a penetrating fragrance.

They have been observed in the markets of the larger Sonoran

towns, having been shipped in from the south. They may be locally

cultivated in Sonora, although I find nothing in my notes to that

effect.

YVCCA L., SP. PL. 319. 1753.

Acaulescent or arborescent perennials with a simple or branched

caudex crowned by a rosette of spirally arranged leaves; leaves

pliant or rigid, pungently tipped, denticulate-serrulate or with

filiferous margins or the margin entire; inflorescence a large

panicle borne in the upper axils of the rosettes, rarely apical;

flowers open-campanulate to globose with incurved tepals ; tepals

6, whitish or frequently purple-tinged, subequal, connate towards

base or distinct, fleshy tender ; stamens 6, inserted on an annular

nectary or on base of petals; the filaments enlarged apically,

puberulent, the anthers small, sagitate, close and eccentrally

affixed ; stigma sessile or on a short style, 6-lobed ; ovary superior,

3-locular, with numerous ovules; fruit a loculicidally dehiscent

capsule or indéhiscent, dry and spongy or baccate ; seeds black, flat

and thin or thick, or rounded, rugose or finely wrinkled.
THE AGAVE FAMILY IN SONORA 155

Fruit dehiscent, drying as a persistent capsule; leaves less than

1.5 cm. wide, flexible elata, below.

Fruit fleshy, indéhiscent ; leaves 2-6 cm. wide, stiffer.

Flowers 20-40 mm. long, broader than long ; ovary short, thick,

about 25 mm. long including the pistil; leaves thin, pliant;

plants mostly simple.

Plants surculose, in maturity 3-5 m. tall; leaves 3.5-5 cm.

wide, the margin smooth, filiferous; filaments pubescent

schottii, p. 157.

Plants nonsurculose, 1-2 m. tall; leaves 2-3 cm. wide; margin

nonfiliferous, serrulate; filaments glabrous below the apical

knob madrensis, p. 159.

Flowers 50-100 mm. long ; ovary long and tapering, 40-65 mm.

long; leaves thicker, stiffer, frequently wider; plants usually

developing several trunks.

Large plants branching in the crown; leaves 70-140 X 4-5

cm.; flowers subsessile, erect or divergent; branches of the

inflorescence pubescent grandiflora, p. 162.

Plants not branching in the crown ; leaves 60-80 X 3-4 cm. ;

flowers pedicellate, nodding; branches of the inflorescence

glabrous arizonica, p. 166.

Yucca elata Engelm., Bot. Gaz. Crawfordsville 7: 17.1882.

Yucca angustifolia var. radiosa Engelm. ex King, U.S. Geol. Exploration

40th Parallel Rpt. 5: 496. 1871.

y. angustifolia var. elata Engelm., Acad. Sei. St. Louis Trans. (1873) 3: 50.

1878.

Y. radiosa (Engelm.) Trel., Mo. Bot. Gard. Ann. Rpt. 13: 56. 1902.

Short to medium arborescent plants, mostly 1-3 m. tall, the

older frequently 1- to 3-breached in the crown, and closely surculose forming clumps; rosettes symmetrical, large; leaves numerous, narrowly linear, the blades mostly 40-70 X 0.8-1.5 cm., planoconvex, rounded below, striate, pale green, pliant, divergent, finally reflexing and persisting dry on the trunk; margin white,

finely filiferous; inflorescence a diffuse ellipsoidal panicle, 1.5-2

m. long, the peduncle relatively slender and greatly exceeding the

leaf crown ; flowers creamy white or tinged with green or pink ;

tepals subequal, generally 35-50 X 15-25 mm., the inner broader,
156 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

PN-2166

FIGURE 60.—Yricca elata on the Campbell Avenue Farm, University of Arizona, Feb. 1, 1957. Original seedlings were from greenhouse in 1939.

elliptical or ovate, acute; filaments mostly 18-25 mm. long;

slender; pistil 22-30 mm. long; ovary about 8 mm. diameter,

oblong-cylindrical with deep carpel sutures, abruptly terminating

in style; style 7-10 mm. long, shape variable; capsules smooth

w^hitish, regularly oblong, 5-6.5 X 3-4 cm., the wall rather thin ;

seeds 7-10 x 9-14 mm., thin, dull black, with broad marginal

wing (partly from (57) ).

Type—Engelmann did not specify a type specimen. However,

McKelvey, (33) nominated as lectotype Rothrock 382, U.S. Nati.

Herbarium No. 35891, collected in 1874 near Camp Grant, Graham

County, Arizona. This species commonly occurs on sandy and

gravelly soils at elevations between 1,500 and 6,000 feet from

Pecos County, Texas, to western Arizona and adjacent Mexico.

I have been unable to locate a specimen from Sonora. However,
THE AGAVE FAMILY IN SONORA 157

since Y. elata is well documented along the Sonoran border region

in Arizona and northern Chihuahua, there is little doubt it exists

in adjacent Sonora.

Y, elata is easily recognized in its native range by its elegant

crown of narrow, flexible, finely filiferous leaves with a thin white

margin on well-developed trunks. No other dry-fruited yucca is

known to occur in Sonora. Figure 60 shows a group cultivated

at the University of Arizona, Campbell Avenue farm, in Tucson.

The original seedlings were transplanted from the greenhouse in

1939. The photograph was taken Feb. 1, 1957, so the plants, the

tallest of which are over 6 feet, are 18 years old. This provides

an idea of bulk growth under these arid conditions (average annual rainfall 12-14 inches). The uneveness of growth probably

reflects the genetic variability so common in wild sexual populations. Wild stands have been employed as cattle feed, for making

paper, for jute-class fiber, and for a detergent soap. (See p. 12.)

Kearney and Peebles (30) reported the "Indians eat the young

flower stalks and lower portion of the stem.'' Who will be the first

to collect and document this plant in Sonora ?

Yucca schottii Engelm., Acad. Sei. St. Louis Trans. (1873) 3: 46.

1878.

Plants at first simple, eventually surculose, the trunk short or

to 3-5 m. tall, rarely branched; the rosettes deeply leaved on

young plants, reduced on old ; leaves numerous, mostly 50-80 X

3.5-5 cm., bluish green, thin, pliant, mostly straightly ascending

to declined on trunk, rarely recurving, the margin thin, brown,

friable and exfoliating in age, sparsely filiferous ; terminal spines

small, 5-15 mm. long, weak, reddish brown or fuscous; panicles

short, rarely exceeding leaves, 5-8 dm. long, erect, the rachis and

branches usually densely tomentose ; flowers white, small, globose,

broader than long, truncate at base; tepals 20-40 X 10-20 mm.,

ovate, the outer brownish mucronate ; filaments 14-18 mm. long,

pubescent, outflexed and swollen towards apex ; anthers 3-4 mm.

long, closely and eccentrally fixed ; ovary thick, with pistil about

25 mm. long, abruptly tapering to a 6-lobate stigma; fruit (dry)

60-125 X 25-38 mm., rounded at base, tapering at apex, frequently irregularly constricted ; seeds 5-8 X 7-10 mm., thick, flat

or rounded, black, rough.

Type—Mo. Bot. Gard. 135692. Sierra west of Santa Cruz

County, Arizona. SONORA: El Rancho Roble (loop of the Bavispe

River), northeast of El Tigre, Sept. 2-13, 1941, White Í217 (If.

& fr.) ; Canyon de la Escalera, oak grassland, Aug. 3, 1940, White
158 AGRICULTUEE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

PN-2166, PN-2167

FIGURE 61.—Upper, Yucca schottii in Huachuca Mountains, Arizona; lower,

Yucca grandiflora showing large, divergent, subsessile flowers of the

type.
THE AGAVE FAMILY IN SONORA 159

S128; 21.4 km. north of Nacozari in oak woodland, July 22, 1960,

Feiger 3593, On rocky volcanic and limestone slopes at elevations

between 4,000 and 5,000 feet in oak woodland and pine-oak forests

in southeastern Sonora and adjacent areas in Arizona, New

Mexico, and Chihuahua. (Fig. 61, upper.)

Y, schottii is distinguished by its short inflorescence with small

flowers on tomentose branches, pliant, light bluish-green leaves,

and summer-flowering habit. It shares these characters with Y,

7nadrensis, but the latter has serrulate leaf margins, narrower

leaves, and a simple, shorter habit. Y, schottii is not known to

intergrade with other fleshy-fruited species although they occur

sympatrically. This may be attributable to the spring flowering

of related species and the summer flowering of Y. schottii. Kearney

and Peebles (30) regarded it as, ''A large handsome species, often

cultivated."

Yucca madrensis Gentry, sp. nov. Figure 62

Stem simple, short, rarely to 2 m. tall, terminated by a thin open

rosette; leaves mostly 50-100 X 2-3.5 cm., linear-lanceolate, thin,

flexible, straight to reflexing in age, nearly plane to slightly conduplicate, bluish glaucous green to yellowish green, smooth-striate

above and below, the margins not filiferous, serrulate with a thin

brown and grayish border ; spine 5-12 mm. long, weak, subulate,

brown to gray ; panicle short, not exceeding the leaves, the central

rachis stout, tapering, sparsely pubescent along sutures near the

branch axils, the branches short, the longer in midpanicle 6-8

cm. long, ascending, pubescent; flowers white, small, about 35

mm. long; outer tepals lanceolate, 30 X 10 mm., inner tepals ovatelanceolate, 30 X 12-14 mm. ; stamens 15 mm. long, the filaments

glabrous below the swollen flexuous apex ; ovary 25 mm. long, the

6-lobate papillate pistil on a short beak ; fruit unknown.

Type—Gentry 21209, Sierra Charuco, along the SonoraChihuahua boundary, Mexico, in pine-oak forest, alt. 4,500-5,000

feet, Sept. 9, 1965 (If. & old dried inflorescence), deposited in U.S.

Nati. Herbarium, No. 2557499. Same locality: Upper Sonoran oak

canyon slope, July 23, 1936, Gentry 23Oi (flower description

Planta simplex brevicaula, 1-2 m. alta; folia 50-100 cm. longa 2-3.5 cm.

medio lata, linearo-lanceolata, palido-viridia flexila, margine brunneo-grisea

serrulata; spina terminali debila subulata, 5-12 mm. longa; inflorescentia

paniculata, ramis lateralibus pubescentibus ad 6-8 cm. longa en medio

inflorescentia; perianthemum album ca. 35 mm. longum, segmentis inaequalibus lanceolatis vel ovato-lanceolatis, ca. 30 mm. longis; filamenta 15 mm.

longa glabra infra apicem clavata; ovarium cylindricum 25 mm. longum;

stylus breviter 6-lobatum; fructum non vidi.
160 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

f.V. -« !

'^ ^.W>. ^-

PN-2168, PN-2169

FIGURE 62.—Yucca madrensis on Sierra Charuco: Upper, Note thick base of

dry persisting inflorescence. Type specimen was made of this plant;

lower, younger long-leaved plant.
THE AGAVE FAMILY IN SONORA 161

drawn from this number) ; high rocky rims around Arroyo Hondo,

alt. 5,000 to 5,500 feet. Mar. 6 and 7,1951, Gentry 102J,2 (If. only).

SONORA : Sierra Murrieta above Toribusi, 15 miles west of Bacanora, oak woodland, limestone, alt. about 4,500 feet. May 17, 1957,

Gentry 16630. A rarely observed plant on rocky slopes at elevations between 4,000 and 5,500 feet in oak woodland and pine forests, southeastern Sonora and adjacent Chihuahua.

This rare yucca, named after the Sierra Madre which it inhabits, cannot be certainly placed in section until the fruits are

known. It resembles Y, schotti with its small flowers and flexible

slender leaves. However, none of the fleshy-fruited yuccas, section

Sarcocarpa, have leaves with serrulate margins. Several of the

dry-fruited yuccas, section Chaenocarpa have serrulate leaf margins. The Sierra Charuco plants were originally reported (13)

as Y. rígida Engelm., which is a long-trunked, stiff-leaved xerophyte of the Coahuilan Desert. Other than the serrulate leaf margins, there is little to support placing the Charucan plants in that

species. The Sierra Murrieta specimens are serrulate along only

the upper margins of the leaves. This, together with the more friable nature of the margin, suggests an infusion of genes from

r. schottii. Y. madrensis, because of the similarities in inflorescence and thin, flexuous leaves, appears to form an alliance with

Y. schottii to the north and Y. jaliscensis Trel. to the south.

Trelease did not cite a type specimen for Y. jaliscensis (Contrib.

U.S. Nati. Herbarium 23: 92. 1920.), only "Jalisco; type from

Zapotlan." However, in his earlier account of Y. schottii jaliscensis

Trel. Mo. Bot. Gard. (Ann. Rpt. 13: 99. 1902.), he mentions

Pringle's collection from Zapotlan. The type of Y. jaliscensis Trel.,

therefore, should be taken as Pringle JÍ392, collected in "Valley

Zapotlan, Jalisco, Mexico, 13 May 1893. Tree-like 15-25 feet.'' It

is a good specimen with leaf and flower. Duplicates of this collection are doubtless in many herbaria, as Pringle collected in large

sets.

The name given to Y. madrensis by the mountain people is usually "soco." The roots were reported as used for soap and the

tender, green fruits as eaten by the Warihio Indians. Analyses

of the leaves by the U.S. Department of Agriculture {53, 5i)

showed about 0.025 percent sapogenin in the leaves and 1.5 percent smilogenin in the roots, which identifies the Indian soap

chemically. The plant forms a low, rather bluish-leaved, ornamental rosette. Sometimes the leaves are unusually long and somewhat arching. As with other yuccas, the leaves on the more elongate old stems are smaller than leaves on young vigorous plants.

Y, madrensis flowers in summer along with the rains.
162 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Yucca grandiñora Gentry, Madroño 14: 51-53.1957.

Arborescent, 4-6 m. tall, branching from the base and toward

the crown, with deep leaf crowns ; leaves 70-100 cm. long, to 140

cm. on vigorous young plants, 4-5 cm. wide in the midblade,

slightly narrowed above base, dark green, smooth, ascending to

descending, persisting dry and deflected in age on the trunk, the

margin narrow, brown, exfoliating on dried specimens, the terminal spine stout, brown or chestnut, broadly grooved ; inflorescence

an irregular open panicle, erect or deflexed, 70-100 cm. long;

peduncle 10-30 cm. long, glabrate below; bracts and bractlets

scarious, dull white, friable; lateral branches densely whitetomentose, flexuous, horizontal or openly angled from the central

rachis; flowers short-pedicellate to subsessile, glabrous, creamy

white, divergent, large; perianth 6-9 cm. long, the segments

spreading, ovate, thin, bluntly mucronate, connate at base, the

outer slightly smaller and thicker than the inner ; filaments hyaline pubescent throughout, 45-48 mm. long, flexed and swollen

apically, anthers 5 mm. long, closely affixed, oblong; ovary elongate, 45-60 mm. long, the style forming a short beak with deeply

lobate stigma ; fruit large, fleshy.

Type—Gentry 11601, above Tierra Negra, Cedros Range, east

of Rio Cedros, Sonora, Feb. 14, 1952, U.S. Nati. Herbarium No.

2089433 and No. 2089434 (fig. 61, lower), SONORA: Sierra de

la Ventana, west side of higher peaks, Rancho Guacoporo, May

27, 1965, Gentry & Gomales 21181; 3 miles south of Huachinera,

alt. 4,100 feet, Oct. 4, 1965, Hastings & Turner 65-55 (If. & fr.) ;

21.4 km. north of Nacozari, Feiger 359Í in foothills with oak

woodland ; vicinity of Rancho San Isidro, canyon de Rio Zatache,

alt. 2,930 ft.. Feiger & Marshall 3387, ^^trunk ca. 10 ft." Tentatively assigned here also from 15 miles south of Bacanora in mountain pass. May 16, 1957, Gentry 16615, On grassy slopes in open

woodland, usually with oaks, alt. 2,000 to 4,500 feet on volcanic

and limestone rocks, through southeastern Sonora.

Yucca grandiflora shows a close relationship to Y, arizonica in

its arborescent habit and large flowers and fruits (fig. 63). Among

the rather many characters that separate it from Y, arizonica are

the high-branching trunks, the open ragged inflorescence, frequently declined, with densely pubescent rachises, subsessile vs.

long-pedicellate flowers, and erect or divergent flowers vs. nodding

flowers. The leaf of Y, grandiflora is wider, thinner, and more

flexible than that of Y, arizonica. Except for the arborescent habit

and pubescent inflorescence, there is little similarity between Y,
THE AGAVE FAMILY IN SONORA 163

FIGURE 63.—Yucca grandiflora south of Bacanora; large succulent fruits

PN-2170
164 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

grandiflora and 7. schottii, its nearest geographic neighbor, which

generally occupies higher elevations in the mountains.

The large yucca population about Bacanora is doubtfully assigned here to Y. grandiflora. (See figure 64.) The glabrate

branches and strong pedicels of the inñorescence are more like

those of Y. arizonica, but the treelike form and large, declined,

ragged panicles are more like Y. grandiflora. It appears probable

that an exchange of genes has occurred in this area. The population as a whole consistently resembles Y. grandiflora. Rather than

attempt subspecific or other definition without more material evidence, it seems prudent to do no more than point out the problem

at this writing.

Y. grandiflora in the Rio Mayo country is called *'sahuiliqui,"

the Spanish spelling of the Warihio name for the plant. Some

Mexicans, including those at Bacanora, call it "datil," a common

misnomer for yuccas in the state. Some of the rancheros about

Bacanora expressed a liking for the sweet fruits, saying they

were even better when roasted than raw. I ate them when hungry,

but was soon satiated, as with a bar of candy. Some of these fruits

reach very large size, 8 inches long and 2 inches in diameter and

estimated to weigh about 2 pounds (fig. 63). They are quite bitter

until fully ripe, when the thick flesh all about the seed core becomes soft. The seeds fill the cells irregularly, many of them undeveloped, apparently for lack of the pollen gamete. When seeds

are remote along the cells, they grow very thick, being about

round. Where they grow closely packed, they are strongly flattened

as is typical of seeds of the Agavaceae, but large and thick like

all seeds of the fleshy-fruited yuccas.

Y. grandiflora is not a common plant. Other than the localities

cited above, I remember it only from the Sierra Murrieta, west

of Sahuarita. "Sahuiliqui,'' however, is also reported to grow near

Jecopaco within the state of Chihuahua in the upper Mayo River

country. The mountain pass south of Bacanora along the road to

Tonichi has many fine examples of this handsome plant. The

flowering season is spring; in the Rio Cedros country Y. grandiflora was in early flower in middle February. Around Bacanora,

farther north, the last flowers and ripening fruits were found in

mid-May. On Sierra de la Ventana only dried flowers were found

in late May.
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FIGURE 64.—Yucca grandiflora south of Bacanora; old plants of a typical population discussed on page 164.

PN-2171

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166 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Yucca arizonica McKelvey, Arnold Arboretum Jour. 16: 270.1935.

Figure 65

Aborescent, commonly forming clumps with unbranched trunks

up to 3 m. tall; leaves mostly 60-80 X 3-4 cm. thick, straight,

deflexing and persistent on trunks, concave above, convex below,

light green to yellowish green, with a thin, brown, smooth margin,

rarely exfoliating, and a light reddish to gray terminal spine 1-1.5

cm. long; panicle 5-10 dm. long, erect, the flowers borne above

the apical leaves, long obovoid in outline, the laterals strong and

spreading; rachises glabrous; flowers mostly 60-100 mm. long,

long-pedicellate, creamy white, not expanding, campanulate, the

bracts and buds frequently purple-tinged; tepals lanceolate, 70-

60 X 24-22 mm., acute, firm; filaments pubescent, about 35-40

mm. long; ovary long, narrow, 40-45 X 4-6 mm., with a tapering

style 5-6 mm. long; fruit 15-20 X 4-6 cm., mostly oblong, short

or longbeaked; flesh sweet and edible when ripe; seeds large, thick

flat, dull black.

Type—'Trobably near Nogales, Ariz., Mo. Bot. Gard. No.

135693'' {56), SONORA: 3 miles north of Rancho Tinaja, near

El Datil, Feb. 27, 1951, Gentry 10215; about 5 miles northeast of

El Datil, Altar District, Mar. 22, 1952, Gentry & Fox 11628; 6-7

miles north of Matape, Nov. 4, 1958, Gentry & Arguelles 17619

(If.) ; talus slope of basaltic cliff toward south end of Baviso

Mountains, 17 miles southeast of Magdalena, Sept. 11, 1934, Wiggins 71SU (If. & fl.) ; Barranca 13 miles south of Divisadero, elevation 1,880 feet, Sept. 25, 1934, Wiggins 7U6U B; 21.8 miles east of

Puerto Libertad, alt. 1,550 feet, Feb. 20, 1965, Hastings & Turner

65-17 (If. & fl.) ; south end of Baviso Mountains, 17 miles southeast of Magdalena, Sept. 11, 1934, Wiggins 71SU (If. & fl.) ; same

locality, Shreve 6638; crossroads about 22 miles southeast of La

Union, Feiger & Gentry 7850; 16.4 miles northeast of Mazucahui

along road to Moctezuma, alt. about 2,950 feet, June 11, 1960,

Feiger & Marshall 33^8 (If. & fr.) ; about 15 miles southeast of

Magdalena on road to Cucurpe, Feiger 79U; Rancho San Gorge

(northwest of Hermosillo), alt. about 2,700 feet. Feiger & Russell

6902. Generally ranges at elevations between 1,000 and 4,500 feet

on grassy slopes and along arroyos in the upper desert in southern Arizona and northern Sonora.

The Sonoran plants are generally taller and in smaller clumps

than those observed in the populations of southern Arizona. The

latter are commonly only 1 to 2 m. tall, appearing relatively

stunted, due perhaps to the higher elevations and shorter growing

season. On the granodioritic hills near El Datil, Y, arizonica was
THE AGAVE FAMILY IN SONORA 167

//.. -

PN-2172, PN-2173

FIGURE 65.—Yucca arizonica in northwestern Sonora : Upper, short-stemmed

clone or colony; lower, tall clone near Rancho Dátil,
168 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

associated with Cercidium ('*palo verde"), Carnegiea (*'saguaro"), Olneya ("palo tésota"), and Fouquieria ("ocotillo").

In Sonora, this species is usually recognizable by its long, narrow, rather stiff leaves and tall, unbranching trunks (fig. 11).

The long, narrow tepals and glabrous ñoral rachises distinguish

it further from its arborescent neighbors, Y. schottii and Y.

grandiflora. The plants noted were always scattered; no dense

populations were observed.

Weber (57) in southeastern Arizona found intergrading characters between Y, arizonica and Y, baccata and interpreted this

as hybridization. Out of this complex McKelvey (3S) earlier had

separated Y. thomberi and Y. confinis, distinguished by the characters acaulescent vs. caulescent and the angle of the filament

tip at anthesis. Weber found these characters undefining and recognized the two taxa as hybrids. He mentions the ^'typical blue

form'' character of the leaves for identifying populations to be

associated with Y, arizonica. This hybridization has not been observed in Sonora where all populations seen are outside the natural

range of Y. baccata.

However, in the locality around Bacanora in eastern Sonora,

an extensive population of a tree yucca shows affinities to Y,

arizonica and grandiflora. This Bacanora population resembles

the former in its smaller flowers and nearly glabrous inflorescences, and resembles the latter in its larger size, stem branching,

larger, more pliant leaves, and declined panicles. Both McKelvey's

and Weber's painstaking studies illustrate the striking difficulties

in the taxonomy of these genetically promiscuous plants. The

Sonoran members are no exception and an adequate concept of

relationships still depends upon detailed field population studies.

In Sonora, Y, arizonica is called ''datil" and the name Rancho

Dátil commemorates its presence there. The sugary ripe fruits of

Y. arizonica may grow up to 6-8 inches long and are eaten by the

local inhabitants. The remarks of Bell and Castetter (3) regarding food use of the yucca by the Papago Indians are generally

more applicable to this species than to Y, baccata Engelm. The

latter scarcely occurs in the Papago territory, wheras Y, arizonica

is common through their mountains. Bell and Castetter were apparently unaware of McKelvey's species when writing their

treatise.

Bird peckings have been observed in the ripe fruits of Y, arizonica.

Insect larvae, presumably of the Prónuba moths, frequently bore

through the seeds, leaving but a small percentage of viable seeds.

Because of these and other environmental hazards, only a minute
THE AGAVE FAMILY IN SONORA 169

fraction of any given sexual generation ever matures beyond the

seeding stage, perhaps one seed in a million. Young plants are

always rare. Weber (57) gives 6 years and 1 month as the earliest

age at which Y. arizonica flowered from seed planting. In nature,

the plants do not flower every year because inflorescence apparently depends upon favorable rains. The plants are very long lived

and Weber reasons about their age as follows: ''Y. arizonica occasionally produces stems 2.4 m. tall (Sonoran plants commonly reach

this height). The annual growth rate of seedling stems in this

species was shown to be 0.27 cm. Based upon these measurements,

(for 6-year period) the older stems of Y. arizonica would approximate 880 years old.'' Clumps of this yucca, which gradually build

out by rhizomatous offsets, may represent a generation more than

a thousand years old.

Several analyses of the leaves of Y. arizonica for sapogenin

content were as follows: from El Datil, 0.95 percent and 1.1 percent sarsapogenin; from Nogales, 0.25 percent; and from near

Ruby, Ariz., 0.0 percent. Analysis of the seeds showed 28.4 percent

oil and 14.4 percent protein {53-56).

Yucca sp.

A rare Yucca was collected in sterile condition on Sierra Charuco, Mar. 7, 1951, Gentry 10250, 102h7 (fig. 66). The plant had

a short, simple trunk and a small crown of divergent, yellowishgreen leaves. The leaves have a smooth, glossy surface, unlike

any other Yucca known to me. It reminds me of another colony

of yuccas that occupied the top of Sierra Orejón when I climbed

that peak in the winter of 1934-35. The Rio Mayo skirts the foot

of Sierra Orejón, which rises steeply as an isolated peak in the

Rio Mayo Valley about where the Sonora-Chihuahua boundary

crosses the Rio Mayo. The Warihio Indians told me the plants bore

large fruits Hke bananas, but sweeter, which they ate (i5). It is

probably an undescribed species and botanists in the region should

try to collect it.

HESPERALOE ENGELM. EX. WATS. EX KING, U.S. GEOL.

EXPLORATION 40TH PARALLEL RPT. 5: 497. 1871.

Cespitóse perennials with bulbous fibrous bases forming grasslike clumps ; leaves succulent, linear elongate, fibrous, with narrow

filiferous margins and graying tips; inflorescence racemose to

paniculate, the flowers in indeterminate clusters on unequal pedicels; flowers stipitate, nonsequential, 6-merous, narrowly campanulate with connivent tepals; tepals with fleshy keel, about
170 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

FIGURE 66.—Anonymous species of Yucca on Sierra Charuco.
THE AGAVE FAMILY IN SONORA 171

equal, essentially free but united on a fleshy nectariferous receptacle; filaments inserted on receptacle or adnate to base of

tepals; anthers dorsifixed, sagitate, introrse, included in perianth ;

ovary ovoid to oblong, trigonous, 3-celled ; ovules numerous in 6

ranks; style elongate but included in perianth; stigma papillate,

perforate ; fruit a dehiscent capsule, stipitate, beaked, transversely

rugose, persistent; seeds large, lunate, thin, flat, black; cotyledon

arched in germination.

A small genus of northern Mexico and adjacent United States,

consisting of three species. Hesperaloe nocturna Gentry is knov^n

only from Sonora. H, parviflora (Torr.) Coult. occurs in Texas

and H. funifera (Koch) Trel. in northeastern Mexico. The strong

fibers of H, funifera are used locally for cordage, v^hile H, pariflora is in ornamental culture in Texas. Trelease gave a historical

account of these tv^o species in 1902 (i6). The follov^ing synoptical

key shows some contrasting characters of the three species and

identifies our Sonoran subject, H. nocturna.

Leaves large and coarse, 3-6 cm. wide near the base and to 2 m.

long, nearly straight, with coarse white marginal threads ; flowers

greenish tinged with purple funifera

Leaves smaller, 1-3 cm. wide near base and 1-1.5 m. long, arcuately spreading, with very fine marginal threads.

Flowers diurnal, dark red to light red, tubeless; tepals appressed at anthesis; filaments elongate (13-14 mm.) with

small anthers (4 mm.) ; ovary small (4-5 mm. long), ovoid,

free of receptacle; style elongate (12-13 mm.), slender.

parviflora

Flowers nocturnal, greenish lavender, with a short tube;

tepals reflexed at anthesis; filaments short (8-9 mm.) with

large anthers (8-9 mm.) ; ovary large (8-9 mm. long), oblong, imbedded in receptacle; style short (8 mm.), stout.

nocturna

Hesperaloe nocturna Gentry, Madroño 19: 74.1967. Figure 67

Acaulescent, densely cespitóse rosettes forming large clumps

about 1.5 m. tall and 1-2 broad; leaves narrowly linear, 1-1.5 m.

long, 1-2 cm. broad near the base, striate, plane towards the base

and deeply concave upwards on the upper side, deeply rounded

below, tip acicular and pungent, fraying in age; margin narrow,

brown, irregularly and finely filiferous; inflorescence slender,

1.5-2 m. tall, a simple or 2- to 3-branched arching raceme on long
172 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

FIGURE 67.—Hesperaloe nocturna with flower (N) compared with flower

and inflorescence section of H. parviflora (P).
THE AGAVE FAMILY IN SONORA 173

peduncles ; bracts of the peduncle dry, lanceolate, 6-4 cm. or less

long; bracteoles chartaceous, 3-5 mm. long, obtuse to acute;

flowers 24-30 mm. long, fasciculate on unequal pedicels 5-18 mm.

long, 4-6 or 8 at the node, noctural, pruinose pink to lavender in

the bud, greenish below, stipitate on a pedicellar joint; tube 2-3

mm. long; tepals 15-22 mm. long, reflexing at nocturnal anthesis,

other times appressed, whitish within, the outer tepals pink or

lavender on the back and narrower than the inner, the inner with

a broad flat keel and greenish pink or greenish lavender on the

back ; filaments equaling the pistil, 8-9 mm. long, attached to base

of tepals; anthers sagitate, versatile, 8-9 mm. long; ovary 8-9

mm. long above tube base, green, trigonous, oblong, roundly

angled at the apex, ringed at the base with nectary ; style 8 mm.

long, stout, stigma capitate, papillate; capsules depressed ovoid

or oblong, 3-4 cm. long, 2.5-4.5 cm. broad, apiculate, rugose;

seeds large, black, 8X11 mm.

Type—Gentry & Feiger 19988, 15 miles southeast of Magdalena

along road to Cucurpe, by Sierra Baviso, Sonora, Mexico, alt.

3,200 to 3,500 feet. May 21, 1963, deposited in U.S. Nati. Herbarium, No. 2503567. Also at the type locality: Gentry 19890, Apr.

1 (sterile) ; Feiger 3^58, July 16, 1960. Talus slope of basaltic

cliff toward south end of Baviso Mountains, 17 miles southeast

of Magdalena, Sept. 11, 1934, Wiggins 7182 (If. & cap.). The description of the flowers was made with fresh material; dried

flowers are smaller.

In both H. nocturne and H, parviflora, the flowers appear one

by one at any given node, so there is little apparent progression

up or down the raceme. The strictly noctural flowering of H.

nocturna is of particular importance in separating it from the other

two species with diurnal flowers. Speculatively, the reflexing of

the tepals with their whitish interior are correlative to pollination by night-flying insects; while the red, tubular form of the

diurnal flower is probably correlative to pollination by birds.

Other structures, such as the large nocturnal anthers and the

small diurnal anthers, may also reflect biotic adaptations.

H. nocturna is patently rare; it is not known from other collections or places. However, since much of northeastern Sonora

has not been botanized, it may be expected in other Sonoran localities. The Sonoran collections record the genus from west of the

continental divide. In appearance, the plants closely resemble the

clumped "bear grass,'' Nolina microcarpa, which occurs in the

same region. The habit of Hesperaloe nocturna in figure 68 can

be compared with that of N. microcarpa in figure 72, page 183.
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PN-2176

FiGtTRE 68.—Hesperaloe nocturna in nataral habitat. Large clumps look like "bear grass," Nolina microcarpa, but large capsules

(right) are characteristic of genus Hesperaloe.

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THE AGAVE FAMILY IN SONORA 175

Transplants of H, nocturna have responded well in Nogales and in

the Desert Botanical Garden near Phoenix, Ariz.

DASYLIRION ZUCC. EX OTTO & DIETR., ALLG.

GARTENZEITUNG 6: 258. 1838.

Short, arborescent perennials v^ith thick single stems crowned

with dense rosettes of serrulate or prickly, linear, hard fibrous

leaves ; inflorescence dioecious, narrowly paniculate, elongate, conspicuously bracteate; pedicels jointed at base of flowers; flowers

small, whitish, persistent, congested, on ascending lateral branches ;

stamens 6, vestigial on pistillate flowers ; ovary 3-celled, 3-ovuled,

2 aborting ; stigmas 3 in apical notch of ovary ; capsules 1-seeded,

small, 3-winged, thin-walled. The only general account of this

genus is that written by Trelease in 1911 (^9).

Dasylirion wheeleri S. Wats, ex Rothr., in Wheeler, U.S. Survey

West 100th Meridian Rpt. 6: 378. 1878.

Bushy perennial with stocky stems 0.5-1.5 m. tall, skirted by

persistent, dry, declined leaves ; leaves 1.5-3 cm. broad above flaring clasping bases, 6-10 dm. long, numerous, armed with sharp

slender, straight to antrorse teeth, yellow to brownish, 2-5 mm.

long, 5-15 mm. apart ; panicles slender, 3-4 m. long, the peduncle

extending above the leaves ; flowers on numerous, ascending, short

lateral branches, subtended by broad, scarious, fimbriate bractlets ;

laterals of the staminate flowers longer (6-9 cm. long) and more

densely flowered than the fruiting laterals; fruit obovate, 5-6

mm. wide, 6-7 mm. long, the apical notch about 1 mm. deep, apices

of the wings acute to obtuse, seed 3.5-4 mm. long, brownish;

flowers May-August. (Fig. 69.)

Type—Ash Creek, Ariz., Rothrock 329 and 655, SONORA:

El Rancho del Roble, northeast of El Tigre (Loop of the Rio

Bavispe), alt. 6,000 feet, Sept. 2-13, 1941, White Í216 (fr.) ; western foothills of Sierra de la Caballera, about 10 miles east of

Colonio Morelos, alt. 3,400 feet, Sept. 22, 1941, White U80 (fr.) ;

Arroyo de la Galera, White 306i; Canyon de Bavispe in oak grassland. White 323i; Santa Rosa Canyon near Bavispe, White 575,

Sierra Charuco, Sonora-Chihuahua boundary, pine-oak forest,

Sept. 13, 1935, Gentry 1822. Cerro Jojoba, about 50 miles south of

Caborca, May 20, 1963, Feiger & Gentry 7868 (If.) ; 15.4 miles

southeast of Magdalena on road to Cucurpe, alt. about 3,400 feet.

Feiger 3459. Widely scattered in oak woodland and pine-oak forests at elevations between 3,000 and 6,000 feet. The western geo-
176 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

PN-2176

FIGURE 69.—Dasylirion wheeleri on Sierra Charuco. Thick trunk is hidden by

dry, persistent leaves.

graphic limit is on Sierra del Viejo in northwestern Sonora, east

to western Texas and Chihuahua.

The only other plant with which this species may be confused

in our area is D. leiophyllum Engelm. ex Trel. The latter, however,

can be distinguished quickly by the teeth that bend downward

rather than upward on the lower half of the leaves. There is no

record of D. leiophyllum in Sonora, but it may yet be found in

northeastern Sonora adjacent to New Mexico and Chihuahua.

D. wheeleri is commonly known by an Indian name "sotol."

Some tribes, however, have quite different names for it ; the Warihio call it "sawo" (13). The Indians have many uses for this plant.

The leaves were employed for thatching and for mats and baskets

after the teeth were stripped off. The trunks are now employed
THE AGAVE FAMILY IN SONORA 177

for posts in houses and corals. The fiber in the tender leaf buds

is used as cordage. Like many other members of the Agavaceae,

the soft meristematic tissue in the stem of D, wheeleri is cooked

for food and drink. The hard liquor ''sotoF' is distilled and consumed locally. It is of inferior quality, having a strong taste and

leaving a raspy feeling in the throat after swallowing. Some individuals, addicted to its use, stink like the primitive stills in which

it is made. Standley reported (i3) that the trunks are sometimes used as cattle feed. After the leaves are burned oif, the

trunks are split open to permit hungry cattle to eat the spongy

interior.

Except in botanical gardens, D, wheeleri is infrequent in cultivation. However, because of its singular rosette appearance,

drought resistance, cold hardiness, and long life, it would be adorning and suitable for landscaping public buildings and highways

in our Southwest. The lower part of the leaves with their broad,

ñaring, white bases are frequently seen in southwestern novelty

stores as decoration arrangements and are known as *'desert

spoons."

NOLINA MICHX., FLORA BOREALA AMER. 1: 208. 1803.

Acaulescent or arborescent perennials with the trunk sometimes

swollen at base or below ground ; leaves linear, hard, fibrous, rough

or serrulate on margins ; inflorescence a diffuse racemose panicle,

the branches subtended by bracts ; bractlets minute, scarious, more

or less lacerate-cilate; pedicels jointed; perianth 6-merous, small,

persistent, spreading; stamens 6, usually abortive in fertile

flowers ; filaments short, slender, or dilated at base ; ovary sessile

or short-stipitate and abortive in flowers with fertile stamens,

deeply 3-lobed; ötyle very short; ovules 2 in each cell; fruit a

papery tripartite capsule, tardily dihiscent or rupturing ; seeds 1-3

in a capsule, globose to oblong, light-colored.

The species in this genus are not well known and there are

problems in specific relationships. The record of the genus in Sonora is feeble, as shown by the paucity of specimens cited below.

I have included two taxa, Nolina bigelovii and N. texana compacta,

which apparently have not been collected within Sonora, but which

have been collected along the Arizona-Sonora boundary. There is

little doubt that they occur in Sonora. A systematic revision of

this genus would be an excellent Ph.D. thesis for a student and

would fill a present void in the taxonomy of the Agavaceae. The

last general treatment was by Trelease, "The Desert Group

Nolineae," published in 1911 (í9). The genus consists of about
178 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

20 species and is best developed in Mexico, but several species

occur across Southern United States from California to Florida.

Plants v^ith v^ell-developed trunks, 1-several m. tall; leaves 10-

35 mm. v^ide.

Trunks slender above the base and branched; leaves mostly

10-15 mm. wide at base, persistent serrulate; capsules much

broader than long, 8-9 X 5 mm.; seeds globose, dark gray,

eastern Sonora . matapensis, below.

Trunks short and thick, unbranched; leaves mostly 2-2.5 cm.

wide, at first serrulate, then filiferous ; capsules as long as broad,

8-12 mm. ; seeds ovate to oblong, whitish, northwestern Sonora.

bigelovii, p. 180.

Plants acaulescent ; leaves usually less than 12 mm. wide.

Panicles elongate, about equaling the leaves; capsules inflated,

the seed not protruding early ; bracts subtending lower branches

shorter than the branches microcarpa, p. 181.

Panicles short, within the rosette; capsules not inflated, early

ruptured by growing seed ; lower bracts long caudate, surpassing the lower branches texana compacta, p. 182.

Nolina matapensis Wiggins, Dudley Herbarium Contrib. 3: 65.

1940.

Small trees with simple or 3- to 4-branched trunks, dilated at

base, with checked bark and rather small leaf crowns; leaves

linear, 10-15 mm. wide near base, 7-12 dm. long, recurving, hard,

striate, greenish yellow and somewhat glaucous, finely serrulate

on margins; panicles 1.5-3 times longer than the leaves and variously angled from the stem apex ; primary branches of the panicles

15-35 cm. long; branchlets mostly 10-15 cm. long; pedicels filiform, 8-10 mm. long, jointed 1-2 mm. above base; perianth segments oblong-linear, 2 mm. long, the outer spreading, inner more

erect; capsule much broader than long, 5 X 8-9 mm., depressed,

the apical notch very broad, the wall thin, tardily dehiscent ; seeds

pale brownish, nearly round, 2.5-3 mm. in diameter; flowers-May.

Type—Near Matapé, between Hermosillo and Suaqui, Sonora,

Sept. 29, 1934, Wiggins 7515, in Dudley Herbarium No. 266078.

(A specimen of this number in U.S. Nati. Herbarium is labeled

'^Gravelly washes and hills near Matapé.") SONORA: 9 miles

northeast of Matapé on road to Batuc, north pass in Sierra Batuc,

alt. 3,510 feet, Sept. 9, 1941, Wiggins & Rollins Í39 (fr.) ; same

locality, Sept. 29, 1934, Shreve 6786 (If. & fr.) ; 20 miles southeast of Cananea, May 8, 1948, Wiggins 11699 (fl.) ; Sierra Tecurahui, southeast Sonora, in pine-oak forest, alt. 4,000 to 4,500 feet.
THE AGAVE FAMILY IN SONORA 179

Oct. 26-28, 1961, Gentry, Barclay, & Arguelles 1H29 (fr.) ; Sierra

Orejón, Rio Mayo, Sonora-Chihuahua border, on oak slope, Jan.

1, 1935, Gentry 1209 (fr.) ; El Rancho Roble, northeast of El

Tigre, alt. 6,000 ft., Sept. 2-13, 1941, White 4200 (If. & fr.). Occurs in open stands widely scattered on grassy slopes in oak woodland and with the lower pines at elevations between 3,000 and

5,500 feet; southeast Sonora and adjacent Chihuahua. (Fig. 70.)

N. matapensis is not likely to be confused with any other plant

in Sonora when it is observed in its native habitat. Its narrow

leaves and small flowers and capsules distinguish it from the tree

yuccas, the only other arborescent members of the family in eastern Sonora. Its near relative N. microcarpa (fig. 72, upper) is

PN-2177

FIGURE 70.—Nolina matapensis on mountain above Guasaremos, Chihuahua.

Plants are partly screened by Ipomoea chilopsis and oaks, but black trunk

shows at base of photo.
180 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

trunkless and resembles coarse clumps of bunchgrass. However,

herbarium collections may be hard to separate because there is

some overlap in the relative len^h of pedicels and thickness of

capsule v^alls. A collector's note on plant habit would facilitate

identification of herbarium specimens.

N. matapensis is generally called "palmita'' or ''palmito," but

the Warihio name is "tuya." They use the new leaves in the long,

conic, terminal buds for weaving baskets. The trunks may be used

locally for posts. As far as I know, this tree has never been brought

into horticulture. It could perhaps be used to advantage for landscaping in the semiarid southwestern areas. Nothing is known

about the longevity of the trees nor the reproductive cycle. It seeds

very well in certain years.

Nolina bigelovii (Torr.) S. Wats., Amer. Acad. Arts Proc. 14: 247.

1879.

Dasylirian bigelovii Torr., U.S. Exploration Miss. Pacific Rpt. 4: 151. 1857.

Coarse truncate plants 1-3 m. tall with a large crown of stiff

leaves persisting dry and reflexed on trunk; leaves 80-120 X

15-35 cm., linear, at first serrulate then filiferous ; panicles 6-10

dm. long; bracts deltoid-lanceolate, thin, 4-10 X 1-2 cm., attenuate, soon deciduous ; primary branches of the inñorescence slender

10-20 cm. long, ascending, glabrous, smooth, the secondaries much

shorter ; bractlets chartaceous, lacerate ; perianth segments oblonglinear, 2.5-3 mm. long, the outer introrsely short-apiculate, reflexed in pistillate flowers, the inner erect or ascending ; filaments

of abortive stamens dilated below; stigma lobes sessile, capsule

narrowly emarginate at apex and base, 8-12 X 9-12 mm. ; seeds

ovate to oblong, 2.5-3.5 mm. long, whitish, coat wrinkled.

Tyi)e—Biglow 155S-Í, Bill Williams Fork, Ariz. Trelease (^7)

doubtfully referred Schott lUl from an unspecified locality in

Sonora to this species—''with fruit scarcely 8 mm. in diameter."

The following specimens are from the Arizona-Sonora border

and indicate this species in northwestern Sonora. ARIZONA:

Tinajas Altas, Yuma County, Mar. 21, 1933, Shreve 6233; same

locality, Kearney & Peebles 10911; arroyo head. Tule Tank Canyon, Cebeza Prieta Range, Yuma County, Apr. 14, 1947, Tinkham

s.n,; Tule Mountains, Mexican boundary line, Feb. 11, 1894,

Mearns 2797, On rocky slopes and gravelly bajadas of mountains

in northern part of the Sonoran Desert. Although Nolina bigelovii

presents a rank growth aspect, it is one of the most xerophytic of

thenolinas. (Fig. 71.)
THE AGAVE FAMILY IN SONORA 181

PN-2178

FIGURE 71.—NoUna bigelovii in western Arizona near type locality. The two

large flowering plants in foreground are about 9 feet tall.

Nolina microcarpa S. Wats, Amer. Acad. Arts Proc. 14: 247.1879.

Acaulescent cespitóse perennial forming large, dense, grasslike

clumps; leaves 6-12 mm. wide, 6-12 dm. long, linear, arching,

serrulate on margins, flat keel beneath, the tips spliting into

brushlike tufts; panicle 1 m. or more long, the lateral branches

15-50 cm. long with numerous branchlets about i/è as long ; bractlets 1-2.5 mm. long, ñmbriate; pedicels slender, 4-5 mm. long;

perianth segments 1.5-2 mm. long, lance-oblong, spreading, whitish; capsule subglobose, 4-5 mm. long, 6-7 mm. broad, deeply

notched at apex, papery, the walls rupturing at maturity; seeds

ovoid to obovoid, 3-3.5 mm. in diameter, light brown, finely

wrinkled.

Type—Rocky Canyon, Ariz., Rothrock 278. SONORA : Tepopa,

Sierra Saguaribo, June 4, 1936, Gentry 2228; mainly in grama
182 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

grassland in oak woodland at elevations between 3,500 and 5,500

feet, from western Arizona to western Texas and adjacent Mexico.

(Fig. 72.)

*'Bear grass'' in the wild pasture lands of Arizona and New

Mexico is familiar to every stockman. The only name I have for

the plant in Sonora is ''sotol chiquito'' in the Rio Mayo country.

It is very rare there and this name sounds quite spurious. It is

probably abundant in northeastern Sonora, and the lack of specimens again calls attention to the general lack of botanizing in

that region.

Nolina texana S. Wats., var. compacta (Trel.) I.M. Jtn., Arnold

Arboretum Jour. 24: 90.1943.

Nolina erumpens compacta Trel., Amer. Phil. Soc. Proc. 50: 418. 1911.

N. texana S. Wats., Amer. Acad. Arts Proc. 14: 248. 1879.

N, caudata Trel., Amer. Phil. Soc. Proc. 50: 417. 1911.

Acaulescent, lov^, spreading, cespitóse, grasslike perennial ;

leaves narrov^ly linear, 70-120 X 0.3-0.4 cm., deeply rounded

belov^, margins somev^hat serrulate or sometimes smooth ; panicle

short, slender, 4-5 dm. long, the primary branches 10-15 cm. long,

slightly scaberulose in lines; branchlets 2-5 cm. long; bracts

caudate-attenuate, surpassing the branches, dry, yellowish, persisting; capsules 4-5 mm. high, 5-7 mm. wide, ruptured early

by enlarging seeds ; seeds globose, 3 mm. in diameter.

Type—Vincinity of New Braunfels, Tex., Lindheimer 550 in

18Í6 and 712 in 1847. Several Arizona collections show this plant

near the Sonoran border : Mule Mountains, Cochise County, July

20, 1894, Tourney, s.n. (type of N, caudata Trel.) ; Monument No.

40, Mexican Boundary line, May 9, 1892, Mearns 258 (If. & fl.) ;

cited by Trelease (loc. cit.) from Nogales are Ferris 1902, Coville

1624 in 1903, and Thompson 1911. These localities are all in the

oak woodland-grama grassland community that characterizes both

sides of the border from Nogales eastward. It is remarkable to

find no collections of this plant during the last 50 years in Arizona

and Sonora. It should be looked for and collected in both the

United States and Mexico; ñowers May-June. (Fig. 72.)
THE AGAVE FAMILY IN SONORA

Éi

A^'».Âi«âkaw.:r

PN-2179, PN-2180

FIGURE 72.—Upper, Nolina microcarpa in Harquahala Mountains of western

Arizona ; lower, Nolina texana near Comstock, Tex., along highway 90.
184 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

LITERATURE CITED

(1) ANDERSON, EDGAR.

1949. INTROGRESSIVE HYBRIDIZATION. 109 pp. John Wiley & Sons,

New York.

(2) BAKER, J. G.

1888. HANDBOOK OF THE AMARYLLiDEAE. 216 pp. London.

(3) BELL, W. H., and CASTETTER, E. F.

1941. ETHNOBIOLOGICAL STUDIES IN THE AMERICAN SOUTHWEST. VII.

THE UTILIZATION OF YUCCA, SOTOL, AND BEARGRASS BY THE

ABORIGINES IN THE AMERICAN SOUTHWEST. N. Mex. Univ. Bul.

372, Biol. Ser. 5, 74 pp.

(4) BENNETT, W. C, and ZINGG, R. M.

1935. THE TARAHUMARA, AN INDIAN TRIBE OF NORTHERN MEXICO. 412

pp. The University of Chicago Press, Chicago.

(5) BERGER, ALWIN.

1915. DIE AGAVEN. 285 pp., illus. Jena. Verlag von Gustav Fischer.

(6) BREITUNG, A. J.

1968. THE AGAVES. Cactus and Succulent Jour. Yearbook. 107 pp.

[Reprint from Cactus and Succulent Jour. vols. 31-38.] Abbey

Garden Press, Reseda, Calif.

(7) CAMP, W. H., and GILLY, C. L.

1943. THE STRUCTURE AND ORIGIN OF SPECIES. Brittonia 4: 325-385.

(8) CASTETTER, E. F., BELL, W. H., and GROVE, A. R.

1938. THE EARLY UTILIZATION AND DISTRIBUTION OF AGAVE IN THE

AMERICAN SOUTHWEST. Univ. N. Mex. Bul. 335, 92 pp.

(9) CORRELL, D. S., SCHUBERT, B. G., GENTRY, H. S. and HAWLEY, W. O.

1955. THE SEARCH FOR PLANT PRECURSORS OF CORTISONE. Econ. Bot.

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(10) CRUSE, R. R.

1949. A CHEMURGIC SURVEY OF THE DESERT FLORA IN THE AMERICAN

SOUTHWEST. Econ. Bot. 3:111-131.

(11) EHRLER, W. H.

1967. AGAVE PLANT, EFFICIENT WATER USER. U.S. Dept. Agr. Agr.

Res. 16 (4) : 11.

(12) ENGELMAN, GEORGE.

1875. NOTES ON AGAVE. Acad. Sei. St. Louis Trans. 3: 291-322.

(13) GENTRY, H. S.

1942. RIO MAYO PLANTS. Carnegie Inst. Wash. Pub. 527, 328 pp.

(14)

(15)

(16)

1949. LAND PLANTS OF THE CALIFORNIA GULF REGION. Allan Hancock

Foundation. Pacific Expeditions 13: 81-180.

1963. THE WARIHIO INDIANS OF SONORA-CHIHUAHUA : AN ETHNOGRAPHIC SURVEY. Bur. Amer. Ethnographic Bul. 186: 61-144,

pis. 28-38.

1967. PUTATIVE HYBRIDS IN AGAVE. Jour. Hered. (Washington) 58:

32-36.

(17) GONCALVES DE LiMA, OSWALDO.

1956. EL MAGUEY Y EL PULQUE. 278 pp., illus. Fonda de Cultura

Económica, Mexico.
THE AGAVE FAMILY IN SONORA 185

(18) GRANICK, ELSA B.

1944. A KARYOSYSTEMATIC STUDY OF THE GENUS AGAVE. Amer. Jour.

Bot. 31: 283-289.

(19) GRANT, VERNE.

1963. THE ORIGIN OF ADAPTATIONS. 606 pp. Columbia Univ. Press,

New York.

(20) HARLAN, J. R., and DEWET, J. M. J.

1963. THE COMPILOSPECIES CONCEPT. Evolution 17: 493-497.

(21) HARRINGTON, M. R.

1933. GYPSUM CAVE, NEVADA. Southwest Museum Papers, Southwest

Museum, Los Angeles.

(22) HODGE, CARL, ed.

1963. ARIDITY AND MAN. Amer. Assoc. Adv. Sei. Pub. 74: 309-337.

[Especially, AGAVE USES, pp. 325-326; and A. SISALANA, A.

ATROVIRENS, A. VILMORINIANA, A. LECHEGUILLA. STEROIDS &

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(23) HoYER, B. H., MCCARTHY, B. J., and BOLTON, E. T.

1964. A MOLECULAR APPROACH IN THE SYSTEMATICS OF HIGHER ORGANISMS. Science 144: 959-967.

(24) JACOBI, G. A.

1864. VON VERSUCH ZU ENNER SYSTEMATISCHEN ORDNUNG DER AGAVEEN.

296 pp. Hamburg. Separabdruck der Hamberger Garten- und

Blumenzeitung.

(25)

1864-68. Hamburger Garten- und Blumenzeitung. [This is {2J^) in

serial journal form. The later years contain supplementary

notes to his Systematischen Ordnung Agaveen; plant species

entered by numbers referring to those as given originally in

book. Additional species are given A and B numbers.]

(26)

(27)

1868. NACHTRAGE I ZU DEM VERSUCH ZU ENNER SYSTEMATISCHEN

ORDNUNG DER AGAVEEN. pp. 138-174.

1870. NACHTRAGE II OR ZWEITER NACHTRAGE ZU DEM VERSUCH ZU ENNER

SYSTEMATISCHEN ORDNUNG DER AGAVEEN. pp. 147-175.

(28) JACOBSEN, J.

1960. HANDBOOK OF SUCCULENT PLANTS. 1: 64-130. Blandford Press.

[An English translation of Berger's work, but without his keys

and only partial descriptions. Cites some additional horticultural names, but not the later species of Trelease and other

American authors.]

(29) JOHNSTON, I. M.

1924. EXPEDITION OF THE CALIFORNIA ACADEMY OF SCIENCES TO THE

GULF OF CALIFORNIA IN 1921—THE BOTANY. Calif. Acad. Sci.

Proc. 12: 951-1218.

(30) KEARNEY, T. H., and PEEBLES, R. H.

1942. FLOWERING PLANTS AND FERNS OF ARIZONA. U.S. Dept. Agr.

Misc. Pub. 423, 1069 pp.

(31) LANGMAN, IDA K.

1964. A SELECTED GUIDE TO THE LITERATURE ON THE FLOWERING PLANTS

OF MEXICO. 1015 pp. Univ. Pennsylvania Press.
186 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

(32) LITTLE, ELBERT.

1943. In Benson, Revision of Status of Southwestern Desert Trees

and Shrubs. Amer. Jour. Bot. 30: 235.

(33) MCKELVEY, SUSAN D.

1938 and 1947. YUCCAS OF THE SOUTHWESTERN UNITED STATES. Part

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(34) MAYR, ERNST.

1959. ISOLATION AS AN EVOLUTIONARY FACTOR. Amer. Phil. Soc. Proc.

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(35)

1966. ANIMAL SPECIES AND EVOLUTION. 797 pp. Harvard Univ. Press,

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(36) and others.

1957. THE SPECIES PROBLEM. Amer. Assoc. Adv. Sei. Pub. 50, 395 pp.

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(37) MILLER, C. D., and HAMILTON, R. A.

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(38) PENNINGTON, C. W.

1958. TARAHUMAR FISH STUPEFACTION PLANTS. Econ. Bot. 12: 95-102.

(39) ScHUCHERT, CHARLES.

1935. HISTORICAL GEOLOGY OF THE ANTILLEAN—CARIBBEAN REGION.

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1920-26. TREES AND SHRUBS OF MEXICO. Contrib. U. S. Nati. Herbarium 23: 87-142.

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THE AGAVE FAMILY IN SONORA 187

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GLOSSARY OF SPECIAL TERMS

abaxial, the side of a lateral organ away from the central axis.

acaulescent, stemless or without visible stem below the leaves.

acicular, needle-shaped.

adaxial, the side of a lateral organ next to the central axis.

adventitious buds, those produced in areas without visible bud

initials, as from the stem instead of the axils of the leaves.

allopatric, applied to allied species or populations inhabiting separate geographic areas. Compare sympatric.
188 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

antrorse, directed forward or towards the apex, as the prickles on

a leaf margin.

arcuate, moderately arched or curving.

ascending, of leaves pointed upward and outward at about 20°

or more from the horizontal.

attenuate, gradually narrowed or prolonged.

bud printing, of leaves when the margin of one leaf is impressed

upon the surface of the next leaf.

hulbiferous, producing bulbils.

hulhil, small plant reproduced vegetatively in the axils of the inflorescence. A form of asexual reproduction.

cabeza, Spanish, head, applied to the thick, short stem of agaves.

campanulate, bell-shaped.

castaneous, chestnut-colored.

caulescent, having a stem or trunk below the leaves.

cespitóse, as applied to succulents, growing in clusters by the production of basal branches, suckers, or offsets.

chartaceous, papery, dry, and thin.

circadian (L. circa, about & dies, day), relating to biologic variations or rhythms with a cycle of about 24 hours.

clone, a group of individual plants reproduced asexually from a

single original parent.

conduplicate, folded together lengthwise.

conic, cone-shaped.

contingent perennial, a plant living more than 2 years and

whose flowering is contingent upon the proper climatic conditions, i.e., rainfall and higher temperature.

crenate, applied to leaf margin strongly and abruptly undulate

with large teats.

cucullate, hooded at the apex.

cultigen, a plant known to exist only in cultivation. Compare

cultivate.

cultivate, as a noun, a cultivated plant with known wild ancestors.

deflexed, bent downward.

descending, of leaves, directed below the horizontal.

ensiform, sword-shaped.

explánate, flattened, spread out flat.

exserted, exceeding the corolla, as the filaments extending beyond

the tepals.

filiferae, threadlike structures along the leaf margins.

filiferous, having threadlike structures.

friable, said of teeth or leaf margins that are easily brushed or

rubbed off.
THE AGAVE FAMILY IN SONORA 189

funnelform, funnel-shaped.

glaucous, whitened with a waxy coating over the epidermis.

guttered, having the sides of the leaves raised to form a troughshaped leaf, partly conduplicate.

haft, handle, applied to the narrowed lower part of the leaf, which

is usually the least prickly part for grasping with the hand.

keel, the fleshy midrib on the tepal or the leaf.

laterals, the main branches of the paniculate inflorescence.

maguey, an American Indian name for agave. It was picked up

by the earliest Spaniards and appears to have been in general

use in the Caribbean region and Mexico. Cortes in "Historia de

Mexico'' wrote, ''miel de unas plantas, que llaman en las otras, y

estas maguey, que es muy mejor que arrope; y de estas plantas

hacen azúcar, y vino, que es asi mismo venden'' (17).

mescal, an American Indian name applied to agave plants, to the

cooked parts of the same, and to the distilled liquor made from

the meristem. Used more in northern Mexico. Also applied

loosely to Manfreda, Tillandsia, and other monocots.

metl, a Náhuatl name for agave, still in use among native tribal

people in central Mexico; ''pápalo metl" (butterfly agave, A.

potatorum), "tlaca metl" (A. salmiana).

monocarpic, a plant or rosette that flowers once and dies. Compare polycarpic.

multiperennial, a plant that flowers once and dies, but requires

several to many years to mature.

neck, apical portion of ovary between the ovarian cells and the

base of the tube. (See fig. 3, p. 22.)

panicle, the branched inflorescence of the subgenus Agave with

flowers borne in umbellate clusters on lateral branches.

paniculate, like a panicle.

patulous, standing open, spreading.

plane, applied to leaves having the upper surface flattened as compared with guttered or explánate.

pohjcarpic, a plant or rosette that flowers repeatedly, but not necessarily every year. Compare monocarpic.

proterandrous, the condition of a perfect flower when the anthers

dehisce before the pistil is receptive. Hence, the flower cannot

normally be self-fertilizing.

pruinose, having a waxy exúdate on the surface of the leaf.

pulque, the fermented juice of the larger agaves. Word derived

from Náhuatl "poliuhqui" or "ocli poliuhqui," but which was

applied to soured or spoiled "ocli," fid Nunez Ortega, the chronicler of Hernando Cortes (17).
190 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

raceme, an inflorescence in which the flowers are borne on pedicels

along a central axis.

reclínate, reclining, applied to leaves pressing downward upon

lower leaves or on the ground.

recurved, recúrvate, curved backward or downward.

reflexed, bent sharply downward.

retrorse, directed towards the base.

rhizome, underground stem or shoot.

rosette, a closely spaced group of radiating leaves limited to a

portion of the stem, usually at the base of the inflorescence.

sapogenin, a compound derived by hydrolysis from saponin. A

large group of such compounds have been found in plant tissues

and named according to their speciñc molecular configuration,

e.g., diosgenin, smilogenin, hecogenin.

shaft, in Agave the central axis of the inflorescence including the

peduncle and the central rachis of the flowering portion or

branches.

spike, an inflorescence with the flowers more or less sessile along

a common or single peduncle or shaft.

spine, terminal spine, in Agave the pungent indurated tip of the

leaf.

spreading, of leaves extending outward less than 20° from horizontal.

subulate, awl-shaped, long tapering.

surculose, producing suckers or offsets.

sympatric, applied to related species or populations inhabiting the

same geographic area.

teat, fleshy prominences under the teeth on the leaf margins. (See

fig. 26, p. 85.)

teeth, the prickles along the leaf edge.

tepal, a combination of sepal and petal, applied when the segments

of the perianth are not differentiated into two dissimilar ranks,

(common in monocotyledons).

trigonous, three-angled with plane faces.

tubular, tube-shaped.

umbel, a flat-topped or low-rounded flower cluster with the pedicels

of unequal length from a common point, like an inverted

umbrella.

umbellate, having the inflorescence in umbels or in similar form.

undulate, applied to leaf margins with low teats, wavy, as compared to straight margins.

urceolate, urn-shaped.

vallecullate, little valley-shaped, as applied to agaves having folds

in the leaf towards apex, plicate.
THE AGAVE FAMILY IN SONORA 191

INDEX

(Page number in boldface type indicates taxon description)

Agave, 41

abrupta, 84

aktites, 1,147,148

americana, 21, 82, 150

var. expansa, 80

angustifolia, 143,147

applanata, 19,119

var. huachucensis, 119

asperrima, 19

bergeri, 147

bovicornuta, 14,19, 27, 85, 91

bracteosa, 20, 21

candelabrum, 143

cerulata, 19, 134

chihuahuana, 120

chrysantha, 104

chrysoglossa, 2,14, 69

coUina, 147

colorata, 19,104,117

consociata, 134,135

deserti, 19, 134

desmettiana, 19, 94

ellemeetiana, 65

elongata, 143, 147

expansa, 80

falcata, 58

felgeri, 58, 60

filifera, 48

flexispina, 104, 115

fortiflora, 122,128

fourcroydes, 138

franceschiana, 97

geminiflora

var. sonorae, 14

goldmaniana, 8,19,124

guiengola, 20

gutierreziana, 143

hartmanii, 14, 53

huachucensis, 1,119

inaequidens, 27, 86

jaiboli, 3, 89

kirchneriana, 147

lecheguilla, 9, 22

lophantha, 22

marmorata, 128

mayoensis, 62

mescal, 14

miradorensis, 94, 97

multifilifera, 3, 46

murpheyi, 99

neglecta, 97

ocahui, 3, 72

owenii, 14,143

pacifica, 3, 139,142,143

palmeri, 1,18,101, 115

parryi, 19, 119,120

var. hauchucensis, 119

parviñora, 2,14, 54

subsp. flexiflora, 8, 56

subsp. parviflora, 54

pelona, 2, 3, 31, 76,130

planifolia, 153

polianthiflora, 14, 51

potatorum, 21

pumila, 56

regeliana, 94, 97

rhodacantha, 141

roezlii, 8

salmiana, 10, 22

schidigera, 48

schechtendalii, 14, 150

schottii, 10, 57

seemanniana, 88

shawii, 124
192 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

shrevei, 18, 104,107, 111, 119

subsp. matapensis, 2, 115

subsp. shrevei. 111

sisalana, 19, 97

sonorae, 14

spectabilis, 143

striata, 8, 21

subsimplex, 3, 14,131

sullivani, 138

utahensis, 21, 80

vilmoriniana, 3, 10, 62

weberi, 97

wocomahi, 19, 88, 105

yaquiana, 14,143

zebra, 2, 80, 126

Agave,

associates of, 6

capsules of, 21

characters of, 17

collectors of, 15

diseases of, 8

enemies of, 6

flowers of, 20, 28

foods of, 9, 10

fossils of, 5

habit of, 17

inflorescence of, 21

karyotypes of, 25

pollinators of, 8

populations of, 2, 26

prickles of, 19

products of, 10

seedlings of, 2

seeds of, 21

stomates of, 17

uses of, 9

uses of agave fiber, 9

agua miel, 11

ahmmo, 134

amolé, 60, 69, 72

amoliyo, 60, 76

Arguelles, Juan, 15,16

ascorbic acid, 12

Aztec codices, 11

B

Baker, John G., 24

Barclay, Arthur S., 15

bats, 7, 9

bear grass, 182

bees, 8, 57

beetles, 8

Berger, Alwin, 24

Bezy, R. L., 16

bighorn sheep, 6,138

Bravoa singuliflora, 151

bulbils (See glossary)

bumblebees, 8

cabezas, 9

caracara, 7

Carnegiea, 168

cattle food, 10

Ceiba, 76

Cercidium, 168

Chaenocarpa, 161

chahuiqui, 48

Citellus, 6

Colletotrichum, 8

Coreidae, 8

cortisone, 10

Cretaceous, 3

cultigen (See glossary)

D

Dasylirion, 175

bigelovii, 180

leiophyllum, 176

wheeleri, 175

datil, 168

den ; kl, 134

Dioscorea, 11

diploids, 25
THE AGAVE FAMILY IN SONORA 193

Diploidia, 8

Dipodomys, 6

dove, 7

E

endemics, 2, 5

Engelmann, George, 24

evolution,

agave, 2

factors of, 3

physiographic, 3, 5

fairy rings, 2

Feiger, Richard S., 15, 16

flicker, 7

flies, 8

flower measurements, 28

flower parts, 36, 37

flycatcher, 7

Fouquieria, 168

fungi, 8

Furcraea, 20

generation, 3

generative period, 3

Gentry, Howard S., 15

Goldman, E. A., 15

ground sloth, 6

H

hawks, 7

Hemiptera, 8

henequén, 139

Herman, Frederick, 17

Hesperaloe, 169

funifera, 171,173

noctura, 5,171

parviflora, 171,173

homologous variation, 20,23, 25

Hughes, Regina 0., 17

hummingbird, 7

Hymenoptera, 8

ideographs, 36, 39

Indians

Apache, 105

Mayo, 9, 115, 119, 150

Papago, 80,101,105,126,

138

Pima, 105

Seri, 80,134

Tarahumara, 11

Warihio, 54, 91,107,112,

161, 164,169,180

Yaqui, 119

Yuma, 138

Jacobi, G. A., von, 24

jaiboli, 91

Johnston, I. M., 15

K

key to Agave species, 42

key to genera of Agavaceae, 40

Kleine, Oda, 16

lechuguilla, 105,126

lechuguilla ceniza, 112

lechuguilla verde, 88

Littaea, 19, 21

Lowe, Charles, 17

M

McKelvey, Susan D., 15

maguey, 148

majahui, 76
194 AGRICULTURE HANDBOOK NO. 399, U.S. DEPT. OF AGRICULTURE

Manfreda, 20,150,153

jaliscana, 152

planifolia, 153

singuliflora, 151

virginica, 150

Marshall, Joe T., 16

Marshall, Peter, 16

Mayr, Ernst, 24

Megathymidae, 8

mescal (See glossary), 80, 147

mescal bacanora, 147

mescal ceniza, 119

mescalero, 16, 80

mescalito, 54, 62

Miocene, 3

Miocene shore line, 4, 5

Moser, Becky, 16

Moser, Cathy, 16

Pajarito Mountains, 56

Palmer, Edward, 15

palmita, 180

palmito, 180

palo tesota, 168

palo verde, 168

parrot, 7

Pennsylvanian, 3

Pericos plantation, 139

Pleistocene, 3

Polianthes, 52, 151,153

tuberosa, 154

polyploids, 25

Prodoxidae, 8

Prónuba moth, 168

pulque, 11

N

Nelson, E. W., 15

Neotoma, 6

Nolina, 177

bigelovii, 5, 177, 180

caudata, 182

compacta, 182

erumpens compacta, 182

matapensis, 178

microcarpa, 173, 179, 181

texana, 182

var. compacta, 177,182

noriba, 89

O

Obregon plantation, 139

ocahui, 76

ocotillo, 168

ojahui, 76

Olneya, 168

opuntia, 10

quids, 9

R

raven, 7

Rollins, Reed C, 15

root system, 151

Rose, J. N., 15

Russell, Alexander, 16

Russell, Paul, 15

Russell, Robert, 16

sahuiliqui, 164

sapogenins, 13

Sarcocarpa, 161

sawo, 176

Schott, Arthur, 15

seed oil, 13

seedlings, 2

sex hormones, 10

Shreve, Forest, 15

sobali, 57
THE AGAVE FAMILY IN SONORA 195

sobari, 57

soco, 161

sotol, 176

sotol chiquito, 182

species concept, 23

Standley, Paul C, 15

Stronak, Dirk D., 16

subspecies, 27

sugars, 9

surculose (See glossary)

Sylvilagus, 94

taiehcholi, 54

tauta, 57

tautilla, 57

temeshi, 91

tequila, 11, 148

Tertiary, 4

Thomas, Robin, 16

Thomomys, 6

totasali, 112

Trelease, William, 15, 24

Triassic, 3

trinomials, 27

tuya, 180

U

umbel (See glossary)

V

varieties, 27

vitamins, 12

volcanics, 3

vulture, 7

W

wasps, 8

Weber, John H., 15

White, Stephen S., 15

Wiggins, Ira L., 15

wocomahi, 88, 107

Yucca, 12, 154,169

angustifolia

var. elata, 155

var. radiosa, 155

arizonica, 5, 12, 162, 166

baccata, 13, 168

brevifolia, 12

confinis, 168

elata, 12, 155

elephantipes, 12

filifera, 13

glauca, 12

grandiflora, 12,162,168

jaliscensis, 161

madrensis, 159

products of, 12

radiosa, 155

rígida, 161

schidigera, 13

schottii, 157,161, 164

thornberi, 168

uses of, 12

•Ù U.S. GOVERNMENT PRINTING OFFICE: 1972 <