SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 43 

A Revision of the Genus Polylepis 

(Rosaceae: Sanguisorbeae) 

Beryl B. Simpson 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1979 
ABSTRACT 

Simpson, Beryl B. Revision of the Genus Polylepis (Rosaceae: Sanguisorbeae). 

Smtthsontan Contributions to Botany, number 43, 62 pages, 39 figures, 1979.- 

Species of Polylepis are shrubs or trees native to the mid- and high-elevation 

tropical Andes. Some species of Polylepis form woodlands growing well above 

normal treeline within grass and scrub associations at elevations over 5000 m. 

Consequently, Polylepis appears to be the highest natural occurring arborescent 

angiosperm genus in the world. The physiological basis allowing such high altitude growth is not understood, but evidence indicates that woodlands at these 

elevations are restricted to microsites where ecological conditions are similar to 

those of lower altitudes. The genus is distinct from other rosaceous members of 

the tribe Sanguisorbeae and is interpreted here as consisting of three species 

groups that appear to have spread independently north and south along the 

Andean chain. Because of the patchy distributions of populations and disturbances 

by man, species are often polytopic. As a result, numerous forms have been described as distinct taxa in the past. In addition, hybridization appears to occur 

between species making circumscription of some taxa difficult. In this revision, 

evidence from studies of gross morphology, leaflet anatomy, pollen morphology, 

and field observations of the autecology of various populations were used to ascertain the number of species within the genus and their relationships to one another. 

Data from these sources indicate that Polylepis should be considered as consisting 

of 15 species, one of which is first described in this treatment. A key for the identification of the species is provided. Each species is illustrated, discussed, and its 

modern distribution mapped. 

OFFICIAL PUBLICATIONS DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution's annual report, Srnithsonian Year. SERIES COVER DESIGS: Leaf clearing from the 

katsura tree Cei cidipliyllum japoiiicicm Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Simpson, Beryl B. 

A re\ ision of the genus Polylepis (Rosaceae: 

(Smithsonian contributions to botany : no. 43) 

Bibliography: p. 

1. Polylepis. 2. Botany--Andes. I. Title. 11. Series: Smithsonian Institution. Smithsonian 

QKl.SZ745 no. 43 [QK495.R781 581'.08s [583'.372] 79-986 

Sanguisorbeae) 

contributions to botany : no. 43. 
Contents 

Page 

9 

1 

2 

3 

4 

4 

4 

6 

6 

6 

n 

8 

12 

14 

14 

14 

15 

Polylepis Ruiz and Pavon ....................................a* 17 

Key to the Species of Polylepis . . . . . . . . . , . . . . , . . . . . . . , . . . , . . . , , . . . . 18 

1. Polylepis multijuga PiIger . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . 19 

2. Polylepis lanuginosa Humboldt, Bonpland, and Kunth . . . . . . . . . . 21 

3. Polylepis hieronymi Pilger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , . . . . 23 

4. Polylepis pauta Hieronymus , . . . . . . . . . . . . . . . . , , . . . . . . . . . . . . . . . 27 

5. Polylepis sericea Weddell . . . . . . . . . . . , . . . . . . . . . . , . . . . . . . . . . . . . . 28 

6. Polylepis subsericans Macbride . . , . . , . . . . . . . , , , . . . . . . . . . . . . . . . . 31 

7. Polylepis pepei, new species . . . .. . . . . . . . . . . . . .. . . . . . . . . .. . . . . . 32 

8. Polylepis reticulata Hieronymus . . . . . . . . . . , . . . . , . . . . . . . . . . . . . . . 53 

9. Polylepis weberbaueri Pilger . . . . . , . . , . . , . , . . . . . , . . . . . . . . . . , . . . 37 

10. Polylepis quadrijuga Bitter . . . . . . . . . . , . . . . , . . . . . . . . . . . . . . . . . . . 40 

11. Polylepis besseri Hieronymus . . . . . . , . . . . . . . . . . . . . . . . . . . . . . . . . . 42 

12. Polylepis tomentella Weddell . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . . 46 

13. Polylepis incana Humboldt, Bonpland and Kunth . . . . . . . . . . . . . . . 50 

14. Polylepis racernosa Ruiz and Pavon . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 

15. Polylepis australis Bitter . , . . . , . . , . . . . . . . . . . . . . . . . , , . . . . . . . . . . . 56 

Literature Cited , . . . . . . . . . . . . . . , . . . . . . . , . . . . . . . . . . . . . . . . . . . . . . . , . . 60 

Index ............................................................ 62 

Introduction ...................................................... 

Previous Treatments nus .................................. . 

Morphological Characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Habit .......................................................... 

Bark ........................................................... Branching Pattern and Leaf Arrangement . . . . . . . . . . . . . . . . . . . . . . . . . . Stipule Sheaths .................................................. 

Trichome and Vestiture Types . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leaves and Leaflets .............................................. Leaf Anatomy ................................................... Inflorescences and Flowers . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . . Pollen Morphology . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

Acknowledgments .................................... .. * 

Species Groups and Relationships of the Genus . ... . * *. . '.. .. . '.*'** 

1 

. . . . ....... .. ... ............ .. . .............I. .. .. . 

tribution . . . . . . . , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

. . .. . .. ...... . .. ,.. . . ....... ... *.. .. .... 3.. . * ...... 

... 

111 

A Revision of the Genus PolyZepis 

(Rosaceae: Sanguisorbeae) 

Beryl B. Simpson 

Introduction 

Above elevations of 3500 m in the tropical Andes, 

vegetation usually consists of various forms of low 

grass and scrub paramo or puna. Woody elements 

are scarce at such elevations in terms of both numbers of species and individuals. Yet, one arborescent 

genus, Polylepis, is found throughout the high 

tropical Andes. Although some members of the 

genus are components of the upper montane forest, 

others occur in woodlands at elevations as high as 

5200 m, completely surrounded by puna vegetation 

and well isolated from any other type of forest. The 

presence of true trees growing at such altitudes is 

rare anywhere in the world and consequently Polylepis ranks with the conifers of the Himalayan 

Mountains as the highest natural occurring trees. 

The deep red-colored bark and contrasting dark 

green leaves of Polylepis trees makes them even more 

noticeable in the often bleak paramo or puna landscapes. On close examination the bark can be seen 

to be composed of almost an infinite number of 

parchment-thin layers. The name Polylepis is, in 

fact, derived from the Greek words poly (many) 

plus letis (layers), referring to the shredding, multilayered bark that is common to all species of the 

genus. 

The physiological tolerances allowing growth at 

these elevations and the prehuman, natural distribution of Polylepis have been the subjects of considBeryl B. Simpson, Department of Botany, National Museum 

of Naturnl History, Smithsonian Institution, Washington, D.C. 

20560. Present address: Department of Botany, University of 

Texas, Austin, Texas 78712. 

erable debate, but evidence indicates that even before severe decimation by man, high elevation trees 

of Polylepis were limited in their distribtution by 

the presence of specialized microhabitats. 

In addition to its uniqueness as an inhabitant of 

extremely high elevations, Polylepis has played an 

important role in the culture of various Andean 

Indian groups by providing building material and 

firewood (Hueck, 1972; Pulgar, 1967). As pointed 

out by Koepcke (19611, the woodlands themselves 

constitute a distinctive habitat for other organisms. 

A monotypic bird genus, Oreomanes, is completely 

restricted to Polylepis woods, providing the only 

case of an avian genus limited in distribution to a 

single plant genus. It is likely that other endemic 

faunal elements will be found in the future. 

Taxonomically, the genus belongs within the 

Rosaceae to the subfamily Rosoideae, tribe Sanguisorbeae (Focke, 1888; Robertson, 1974 a,b,c). 

Although similar in various morphological traits 

to Acaena Linnaeus and Margyricarpus Ruiz & 

Pavon (including Tetraglochin Poeppig), the genus 

is a natural one and quite distinct from all other 

members of the family. However, the spotty distribution of most species (caused by both natural and 

human factors) has resulted in much interpopulational variation. In the past, local populations were 

often described as a distinct taxa (varieties, subspecies, or even species). Of the previously described 

taxa, only 15 are considered here to represent meaningful biological entities. A new species from Peru 

and Bolivia is described. In the determination of 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

these taxa, I have used habitat data, gross morphology, leaf anatomy, and palynology; chromosome counts were unsuccessfully attempted. Because 

of the specialized environmental conditions required 

by Polylepis and its arborescent, long-lived habit, 

no crossing experiments were undertaken. 

ACKNOWLEDCMENTS.-~ thank the curators of the 

Arnold Arboretum, Botanische Museum (Berlin), 

British Museum (Natural History), Gray Herbarium, Conservatoire et Jardin botaniques (Geneva), 

Chicago Natural History Museum (Herbarium), 

Royal Botanic Gardens (Kew), Missouri Botanical 

Garden, New York Botanical Garden, Museum 

National d’Histoire Naturelle (Paris), Rijksmuseum 

(Stockholm), University of California, National 

Museum of Natural History (Smithsonian Institution), and the Naturhistorisches Museum (Vienna) 

for their patience in extending loans of Polylepis 

for a considerable period. I greatly appreciate the 

preparation of the drawings by Alice Tangerini, the 

measuring of specimens and drafting of the maps 

by Stanley Yankowski, the provision of the pollen 

description by Joan IV, Nowicke, the photography 

of Polylepis in the field by John L. Keff, and help 

with the Latin diagnosis by Marie-Helene Sachet 

and Ray Fosberg. Dan Nicolson and Harold Robinson made numerous suggestions for improvements 

of the text. Beth Lawler and Barbara Dennis carefully typed the various drafts of the manuscript. 

Previous Treatments of the Genus 

Unlike most Andean genera, Polylepis has no 

generic synonyms even though it occurs in six different countries. This lack of synonyms seems attributable to its early and complete description by 

Ruiz and Pavon (1794) and the distinctiveness of 

the group from other members of the Rosaceae. 

In addition, all of the subsequently described taxa, 

with the exception of Acaena ochreata Weddell 

(placed here in P. sericea Weddell), were originally 

correctly described as members of the genus. The uncomplicated generic taxonomy contrasts, however, 

with the infrageneric taxonomic situation. Variation within species, the patchy distribution of most 

taxa, and the poor systematic treatments of earlier 

botanists have tended to produce infrageneric confusion. 

Subsequent to the original circumscription of 

Polylepis and the description of a single species, 

P. racemosa (Ruiz & Pavon, 1798), Humboldt, 

Bonpland, and Kunth (1824) described three additional species: P. incana, P. lanuginosa, and P. villosa. The last of these is considered here as a 

synonym. IVeddell (1861), in his compilation of 

Andean plants added three more species (P. tomentella, P. sericea, and “Acaena” ochreata) as well 

as two varieties of earlier described species. His 

interpretation of P. lanuginosa and his variety, P. 

lanuginosa ,8 microphylla, both appear to be misunderstandings of the true P. lanuginosa and of 

the nature of a set of small-leaved specimens that 

are considered here as belonging to P. weberbaueri. 

Hieronymus (1 895 and 1896) described five species 

of Polylepis from northern South America, which 

he found among the collections of Lehmann (P. 

lehmannii) and Stubel (P. besseri, P. reticulata, P. 

stuebelii, and P. pauta). In 1906 Pilger described 

five more species (P. hieronymi, P. albicans, P. 

weberbaueri, P. niultijuga, arid P. serrata) from 

Weberbauer’s Peruvian specimens sent to the museum at Berlin. 

The only study of the genus as a whole, that of 

Bitter (191 Ib), did little to further understanding 

of the group. Bitter’s typological concept (Mayr, 

1957) and his treatment of aberrant individuals 

rather than distinctive populations as varieties 

(Mayr, 1959) led to such taxonomic fragmentation 

that it became practically impossible to identify 

new collections. In all, Bitter described in his monograph 13 of the 33 species he recognized, as well 

as nine subspecies and 18 varieties. For one species, 

Polylepis incana, he circumscribed nine infraspecific 

taxa. Later, in 1913, Bitter described a new variety 

of Polylepis australis (P. australis var. tucumanica) 

with four new subvarieties. This small group of 

isolated populations in southern Bolivia and northern Argentina, earlier described as P. australis by 

Bitter, had already been split into several varieties 

(as varieties of P. racemosa) by 0. Kuntze (1898), 

and Bitter had added four more in his 1911 monograph. Few of Bitter’s taxa are retained here. It is 

fortunate that there are photographs of the types of 

the Berlin Herbarium, which was destroyed in part 

during World War I1 (the Rosaceae being one of 

the casualties), because it allows an assessment of 

Bitter’s species that might otherwise have been 

totally impossible. Unfortunately, there are no 

isotypes or photographs of many of his varieties 

and subspecies. In these cases, I have relied on de- 
NU.MBER 43 3 

scriptions and locality data as guides to their placement. 

After Bitter’s treatments, only five new taxa 

(Polylepis subintegrifolia, P. subsericans, P. quindiensis, P. boyacensis, and P. cocuyensis) were added. 

An examination of the type material and the range 

of variation present in previously described species 

indicates only one of these (P. subsericans) is actually distinct. 

Species Groups and Relationships of the Genus 

In his monograph, Bitter (1911b) arranged the 

species of Yolylepi.5 into two sections and 11 

“groups” of species. Six of the groups were given 

formal names, but Bitter never indicated whether 

he considered the groups to be subsections or series. 

Bitter’s arrangement of the species of Polylepis is 

shown in Table 1. Table 2 shows the infrageneric 

groupings proposed here. Although there has been 

much reduction in the number of recognized species 

and some rearrangement, the species groups or complexes recognized here agree to some extent with 

Bitter’s sections except that his section Gymnopodae 

is split into two. None of the groupings proposed 

here are given formal recognition. The genus is such 

a tightly knit unit, well isolated from other genera 

in the Sanguisorbeae, and has so few species, that 

formal recognition of subgeneric groups seems unduly complex. Instead, I have placed all of the 

members of Bitter’s section Dendracaena into a 

group called the sericea group and split the section 

Gymnopodae into the reticulata group and the 

incana complex. The last is called a complex beTABLE 1.-Alignment of Polylepis species according to Bitter (1911b) 

Section Group 

Dendracaena PI uri jugae 

Annulatipilosae 

unnamed 

La t ifol iola tae 

unnamed 

Subtustomentosae 

Gymnopodae unnamed 

Supranitidae 

unnamed 

Paucijugae 

unnamed 

Paucijugae 

Bitter’s 

species Present treatment 

P. multijuga P. multijuga 

P. serrata 

P. pauta } P. pauta 

P. annulatipilosa J 

P. ochreata 

P. Stuebeli 

P. h~Pa~rea } P. sericea 

P. sericea I 

P. coriacea 

P. Lehmannii 

P. lan ugin osa 

P. hypoleuca 

P. Hieronymi 

P. microphylla 

P. Weberbaueri 

P. quadrijuga 

P. reticulata 

1 

7 

I 

P. albicans J 

P. lanuginosa 

P. hieronymi 

P. quadrijuga 

1 

i 

1 

1 P. reticulata P. nitida 

P. brachyfihylla 1 ._ . I 

P. australis P. australis 

P. racemosa P. racemosa 

P. villosa 

P. Besseri 

P. triacon tandra 

P. Crista-galli 

P. tenuiruga } P. besseri 

P. subquinquefolia I 

P. rugulosa 

P. pallidistigma 1 

P. incana P. incana 

P. tomentella 

P. tarpacana } P. tomentella 

} 

I 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 2.-Species groups of Polylepis proposed in this 

treatment 

sericea group reticulata group incana complex 

P. multijuga P. reticulata P. racemosa 

P. lanuginosa P. weberbaueri P. incana 

P. hieronymi P. quadrijuga P. besseri 

P. pauta P. tomentella 

P. sericea P. australis 

P. Pepci 

cause of the blurred specific boundaries and the 

indication of hybridization of many of the taxa 

within it. 

The sericea group includes those species with 

leaves usually containing many pairs of leaflets, 

sericeous trichomes on the underleaf surfaces and 

stipule sheaths. Both P. multijuga and P. hieronymi 

have shaggy trichomes on the under surfaces of the 

leaflets but have sericeous stipule sheaths and numerous pairs of leaflets. The one most discrepant element in the group, P. subsericans, forms a connecting link with the incana complex. The trichomes 

of this species are minutely strigose rather than 

truly sericeous and are mixed with glandular multicellular trichomes characteristic of the incana complex. However, because some populations of P. 

sericea (former P. albicans) approach this type of 

vestiture, I have included P. subsericans in the 

sericea group. 

All species placed in the reticulata group have 

nitid, emarginate leaflets with pannose under leaflet surfaces. The three species in the group are 

closely related to one another. Their link to other 

members of the genus appears to be via P. lanuginosa. It is probable that the group was derived from 

an ancestral stock similar to that leading to the 

modern P. lanuginosa and related taxa within the 

sericea group. 

Members of the incana complex have variable 

numbers of pairs of leaflets, but often only one. 

The leaflet lower surface usually has glandular 

trichomes often mixed with a lanose vestiture and, 

in the case of P. racemosa, with scattered, shaggy 

trichomes. Individuals of P. australis with glabrous 

leaflets represent one extreme in this group and 

plants of P. besseri with densely lanose leaves constitute another extreme type. 

Bitter (191 1 b) extensively discussed the relationship of Polylepis to Acaena section Elongata. Particularly members of the sericea group (P. multijuga and P. pauta) share characters of multipinnate 

leaves and long, densely flowered racemes with 

members of this section of Acaena. Bitter argued 

for a derivation of Polylepis from an Acaena-like 

ancestor rather than the reverse. He based this conclusion on Polylepis being more restricted in distribution than Acaena and an inhabitant of more 

specialized habitats. Species of Acaena occur in 

South America, New Guinea, Hawaii, and North 

America, whereas Polylepis is restricted to the 

Andes. Various other authors, while not commenting on the affinities of Polylepis, have placed Polylepis next to Margyricarpus (including Tetraglochin) (cf. Focke, 1888) and more distant from 

Acaena. Hutchinson (1967) placed Polylepis between Leucosidea, which it resembles in many 

vegetative and floral characters, and Tetraglochin 

in his tribe Poterieae (=Sanguisorbeae), but he 

made no statements as to the reasons for his alignment. On the basis of gross and pollen morphology, 

Polylepis appears quite similar to Acaena and 

slightly less similar to Margyricarpus. Since the 

generic relationships of this group of Sanguisorbeae 

are being investigated by Dr. Werner Rauh of 

Heidelberg, I have not attempted to assess the alignment of genera within the tribe. 

Morphological Characters 

HABIT.-AII species of Polylepis are trees or 

shrubs ranging in height from 1 to 27 m (Figure 

1b-d). Although not absolute, there is a loose correlation of habit with habitat. All of the tropical 

taxa that grow at relatively low elevations (e.g., P. 

multijuga, P. pauta) are tall trees (Tosi 1960), 

whereas many of the species growing at elevations 

of 3800 m and above are short trees or shrubs. Both 

P. besseri and P. pepei, which reach elevations over 

4000 m, are shrubby, but P. tomentella, observed 

at 5200 m on Volcan Sajama, Bolivia, retains an 

arborescent habit even at this altitude. In some 

cases, the presence of shrubby individuals in a population has been caused by the activities of man 

or domesticated grazing animals, which prune trees 

and cause the formation of multistemmed individuals through destruction of the apical meristem. 

BARK.-The bark of Poly Eepis consists of numerous layers of thin, dark red exfoliating sheets (Figure la). In some cases, the layered bark can be over 

an inch thick. All of the larger branches have similar shredding bark. No measurements of tempera- 
FIGURE 1.-Growth forms and habitats of Polylepis species: a, trunk of a small tree of P. tonentella with the layered, shredding bark characteristic of all species of the genus; 6, shrubby growth 

form of P. 6essei-i growing in Chapare, Cochabamba, Bolicia: c tree growth form of P. besseri, 

Volcan Chachani, .4requipa, Peru; d, shrubby plants of P. weberbauerj, Canar, Ecuador: e, trees 

of P. raceitiosa planted in the plaza of Cerro de Pasco, Peru: f, cultivated group of trees of 

P. racetnosa around a house, near Tingo Pacha, Junin, Peru. 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tures under the bark relative to the ambient air 

temperatures have been made, but it would seem 

that the bark serves as an insulation from both the 

nightly frosts and the intense diurnal irradiation. 

Temperature readings have been made of plants 

in Afroalpine habitats. Hedberg (1964) showed that 

loose stem coverings such as overlapping scales, mats 

of trichomes, or persistent leaf bases insulated the 

interior of arborescent plants from extremes of heat 

or cold. 

BRANCHING PATTERN AND LEAF ARRANGEMENT.- 

Perhaps, in part because of the harsh environment 

in which many species of Polylepis grow, trees tend 

to have twisted, crooked stems and branches. Contorted growth is often associated with windy, cold, 

or arid habitats. The angular pattern of the 

branches is due primarily, however, to the way in 

which the trees branch and the leaves are borne. 

Growth is sympodial with repeated sympodial 

branching. The lateral branches show a “flush” 

type of growth with long, often naked branch segments and the leaves congested at the branch tips 

(Figure 2a, c). The leaves are borne alternately but 

often appear whorled because of their congestion 

(e.g., Figure 2b). The spurts in growth indicated 

by the long leafless branch segments probably occur 

in the short southern summer when temperatures 

are warm and rainfall is relatively high (a typical 

pattern in most Polylepis habitats). Species that 

occur in the upper portions of the montane rainforest have less pronounced growth flushes than 

those of high elevations. Bitter (1911b) noted the 

tendency of the “less specialized” species to have a 

more even dispersion of leaves along the branches 

than the more “advanced” species. Extremes can be 

seen in P. multiflora (Figure 8) and P. pepei (Figure 20). 

For taxonomic purposes, therefore, the amount 

of congestion of the leaves can be used in combination with other characters to discriminate between 

taxa. The amount of crowding of the leaves near 

the branch tips is variable within most species, 

however, and is not as incisive a character as Bitter 

(191 lb) indicated. 

STIPULE SHEATHS.-Each leaf has a pair of stipules 

fused around the branch forming a sheath. The 

congestion of the leaves results in a pattern of 

stacked, inverted cones due to the overlapping of 

the stipule sheaths (Figure 2a). On the top of the 

sheaths on either side of the petiole there are often 

projections, or spurs. The presence or absence of 

these spurs, their size, and vestiture are important 

taxonomic characters. Extremes can be seen in P. 

multijuga (Figure 8) and in P. incana (Figure 34). 

In addition, the presence or absence and type of 

trichomes extending along the upper surface of the 

leaf petiole onto the inner surface of the sheath 

and projecting from the top of the sheath are useful for separating species. Young leaves and stipule 

sheaths tend to be much more pubescent than 

older, mature leaves. Since inflorescences are borne 

in the axils of leaves, and young leaves would be 

particularly sensitive to cold air temperatures, this 

dense pubescence would serve to insulate vulnerable 

tissues. When the leaves are shed, the petiole up 

to the point of attachment of the basal pair of 

leaflets from the branch and the stipule sheath persist on the branch (Figure 2a). 

TRICHOME AND VESTITURE TYpEs.-The type and 

placement of trichomes on various surfaces and the 

resultant appearance of the surface (vestiture) of 

the plant parts (especially the stipule sheaths, leaflets, and fruits) are important taxonomic characters in Polylepis. Unfortunately, uses of various 

terms are not consistently employed by various 

authors. Bitter (191 lb) divided the trichomes found 

throughout the genus into two principal types: 

capilli resiniferi or capilli pulverulenti and pili. The 

first included most multicelluar trichomes (usually 

glandular) and the second the various types of 

single-celled trichomes or trichomes with one or 

two basal cells. In this treatment, the first types of 

trichomes are simply referred to as multicellular 

trichomes and designated as glandular if they appear to contain, or have secreted, resin. The singlecelled trichomes or those with a long terminal cell 

subtended by one or two small cells can vary from 

short to long, curled to straight. On various surfaces, they appear downy, woolly, shaggy, rough, or 

silky. Throughout this treatment, the following 

usages for vestiture types have been employed. 

Barbate: with a tuft of long, curled trichomes; usually used 

Hispid: with long, straight, stiff, upright trichomes. 

Lanose: with long, twisted, interivoven trichomes producing 

Pannose: Lvith short, twisted, very matted trichomes producPilose: with short, weak, thin, twisted trichomes; variable in 

for trichomes on the apices of the anther sacs. 

a coarse, woolly effect. 

ing a feltlike appearance. 

density. 
NUhfBER 43 7 

FIGURE 2.-Branching and flowering patterns of Polylepis species: a, congestion of leaves of 

P. racemosa at the branch tips rvith persistent petioles surmounting the stipule sheaths of 

dehisced leaves; b, young leaves at the tips of branches of P. weberbaiteri appear almost whorled 

because of the congestion; c, branch of P. incana with pendant inflorescences; d, an inflorescence 

of P. besseri showing the spreading stigma in the lorver rear right flower and the hairy anthers 

on the exposed long-filamented stamens. 

Puberulous: with very short, curved trichomes closely packed 

and producing a downy appearance. 

Sericeous: with straight, long trichomes lying flat on the surface and presenting a silky surface. 

Strigose: rvith straight, rather stir trichomes appressed to the 

surface and producing a rough appearance. 

Tomentose: covered with long, straight trichomes, the trichomes upright and consequently not producing a silky 

surface; usually used here for trichomes on the apices of 

the anther sacs. 

T’illous: rvith long (over 1 mm), weak, hvisted, shaggy trichomes giving a loosely tangled appearance. 

LEAVES AND LEAFLETS.--A~I species of Polylepis 

have compound, imparipinnate leaves, but the 

number of pairs of leaflets varies within and between species. Within a range of one or two pairs, 

the number of leaflets is a useful character for distinguishing between species or species groups. Of 

greater taxonomic utility, however, are the size, 

shape, thickness, and vestiture of the leaflets. Since 

the flowers have no modifications for animal pollinators (see below), there are relatively few floral 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

characters that can be used to distinguish species. 

Consequently, leaf and leaflet characters take on 

primary importance for the identification of taxa. 

The arrangement of the leaflets and the position 

from the terminal leaflet of the largest pair of leaflets determine the shape of the leaf. In species with 

one pair of leaflets, the outline of the leaf is usually 

rhombic. Depending upon the position of the largest 

pair, the leaf can be trullate to obtrullate in taxa 

with more than one leaflet pair (Figure 14). 

Various taxa differ from one another in the type 

and amount of trichome coverage of the rachis and 

the appearance of the top surface of the rachis 

where the leaflets are attached. The rachis can be 

tomentose, villous, lanose, sericeous, pilose, or 

glandular. Often grandular trichomes are mixed 

with some type of single-celled trichome. Bitter 

(1911b) often used the presence of glandular trichomes as the basis for the circumscription of a 

variety (i.e., P. microphylla var. polyaythrotricha). 

Within a population, however, individuals vary in 

the amount of such trichomes. Nevertheless, some 

species such as P. tomentella or P. incana often have 

a preponderance of glandular trichomes that helps 

to distinguish them from other similar taxa. 

The vestiture of the point of leaflet attachment 

is distinctive in some species of Polylepis although 

the density of trichomes and the amount of glandular trichomes present at this point is often variable 

within a species. Most taxa have some sort of a 

hispid vestiture appearing as a tuft of trichomes 

(e.g., Figure 8c), but leaves of P. weberbaueri have 

no differentiated trichomes on this part of the rachis 

or only a band of dark, glandular trichomes (Figure 24e). The glandular trichomes present at the 

point of leaflet attachment often secrete resin, which 

produces a crust of red or orange material. 

The most important taxonomic characters for 

distinguishing species of Polylepis are provided by 

the leaflets. Leaflet characters are often correlated 

with others, such as the length of the inflorescence 

or protuberance type on the fruits, and present the 

most visible characters for distinguishing taxa. Useful characters include the shape of the outline, the 

size of the leaflet, the apex type, the aspect of the 

base, and the vestiture. Most members of the sericea 

group have ovate, entire, or serrate leaflets with 

acute or obtuse apices. Members of the 1-eticulata 

group have elliptic or almost orbicular leaflets 

strongly emarginate at the apex. Leaflets of the 

incana complex are obovate, crenate, and attenuate 

at the base. 

The vestiture of the under leaflet surfaces, and to 

some extent of the upper leaflet surfaces, is very 

species specific, although the density of trichomes 

can often vary within a species. Members of the 

serz'cea group, for example, usually have long, silky 

trichomes on the under leaflet surface, but the pattern of the location of the trichomes varies from 

one species to another (Figure 3). In the reticdata 

group, all taxa have matted, feltlike trichomes on 

the under leaflet surface but differ in the presence 

or absence of a second type of trichome. The species 

of the incana complex all have, to some extent, small 

glandular trichomes, which secret a yellowish resin, 

dispersed on the under surfaces. 

The appearance of the upper leaflet surface providts an additional character for the separation oL 

taxa. The surface can be nitid, rugose, dull, or variously puberulous (Figure 4). 

LEAF ANAToMY.-In his monograph (191 Ib), 

Bitter emphasized the importance of the internal 

structure of the leaves for systematic purposes. Since 

Bitter tended strictly to apply a typological method 

and apparently examined only one or a few leaves 

of each of his narrowly defined taxa, the anatomy 

of the leaves appeared to be quite constant within 

a taxon. We have examined the anatomy of leaves ot 

populations of every species, including all of the 

forms that would have been given specific status 

by Bitter. LVhile it is true that there are modifications of the internal leaf structure that correlate 

with the varying habitats of the different taxa, 

there is too much variation within one species (Figure 5) and too much overlap between species (particularly species of the same species group) to make 

leaflet anatomy a practical way of distinguishing 

between taxa. 

Basically, the leaves of all species are built on a 

plan of dorsiventral arrangement of cells, with the 

epidermis and palisade layer on the adaxial surface 

and the spongy tissue on the abaxial surface (Figure 

4a). In different species, there are modifications that 

include an increase in the thickness of the cuticle 

(Figure 5), smaller and more regularly arranged 

epidermal cells, a more tightly packed series of 

palisade layers (Figure S), reduction of the spongy 

tissue, and an invagination of the lower leaf surface 

so as to produce stomata1 crypts (Figures 4d, 6). In 

almost all species there are bundle sheaths with 
NUMBER 43 9 

FIGURE 3.-Leaflet surfaces of Polylepis species: a, underleaf surface of P. pauta with trichomes 

restricted to the veins and stomates sunken in the areas appearing as circles (X 50); b. a stomate 

from 3a (x 2500) showing the pappilose subsidiary cells; c, untlerleaf of P. pepei from which 

the trichomes have been scraped away and on which the stomates are deeply sunken and surrounded by irregular subsidiary cells (X 300); d, underleaf surface of P. racetnosa, with accesible 

trichomes scraped aicay re\.ealing the depressions (stomata1 crypts) in which the stomates are 

sunken (x 100). 

extensions. Usually there are rhomboid crystals are adaptations to the physiological drought with 

throughout the cells of the bundle sheath extensions which many taxa are faced (Openheimer, 1960). In 

(Figures 4b, 5b, d, 6b), but their arrangement is the species that grow at the upper parts of the moist 

variable within leaflets from the same leaf. ceja de la montana, xeromorpliic adaptations are 

Many of the features associated with the leaflets less pronounced. All of the members of the sericea 
10 SMITHSONIAN COSTRIBUTIOKS TO BOTANY 

FICL'RE 4.- Leaflet cross sections of Pulylepis species (all x 105) : a, P. sekea (sericea group) 

under normal light microscopy showing the dorsiventral arrangement of the leaflet cells with a 

thin epidermis, a 2- or 3-layered palisade layer and a small amount of spongy tissue; b, another 

leaflet of P. sericea ~iewed under polarized light revealing the crystals in the cells of the bundle 

$heath extensions; c, P. tiiullijzign (sericeci group) with its irregular epidermis and rather 

loosely organized palisade layer; d, P. tomeritella (incam complex) with an invaginated underleaflet surface and prominent bundle sheath extensions. 

species group have comparatively mesomorphic 

leaves. The leaflets are large and thin relative to 

their surface area (Figure 4a, b, c). The cuticle is 

thin. The under surface of the leaflet is flat, although 

usually coi,ered, at least in part, with trichomes. The 

cells of the epidermis and palisade layers are not 

tightly compacted although the) can be quite regular (Figure 4u, b). Other species also found at lower 

eleval ions and/or generally found in relatively 

mesic habitats (e.g., P. australzs, P. Izieronymi) have 

similar leaflet anatomies. Species found at higher 

elevations and in drier habitats show a progression 

of specializations. The cuticle becomes markedly 

thicker (Figure 5), the ledlet becomes thicker relative to it5 surface aiea (Figure 5), the epideimal 

layers become more compact, more regular, and 

composed of smaller cells (Figures 4d, 6). In addition, the undersurface of the leaflets of these taxa 

has iiivaginations known as stomata1 crypts in 

which the majority of the stomates occur (Figure 
NUMBER 43 11 

FIGURE 5.-Cross sections of the leaflets of Polylepis reticulntn (x 105), showing the intraspecific 

variability in the number of cell layers of the epidermis and the thick cuticle characteristic of 

members of the wticulata group: n, 0, the same leaflet under regular light (0) and polarized 

light (6) with a 2-layered upper epidermis; c, d, another leaflet ullder regular (a) and polarized 

(b) light rvith an epidermis primarily one cell layer thick. 

34. Stomates are often sunken (Figure 3c) and/or 

surrounded by papillose subsidiary cells (Figure 3b). 

The dense covering of trichomes or resin on the 

under leaf surfaces help prevent dessication when 

the stomates are closed. 

Bitter (1911b) remarked on the groups of cells 

now knowii as bundle sheath extensions. He called 

these cells "pillar cells" and ascribed a supportive 

function to them. Under the microscope, these 

bundle sheath extensions do look like pillars of 

buildings (Figure 4d), but more recent investigations 

have ascribed other functions to them. It now appears that these cells serve to conduct water to the 

epidermis (IVylie, 1951), rather than to serve as 

supports, but the fact that the walls of these cells 

are often thickened might indeed provide some 

strengthening for the leaflets. In an en! ironnient 

with almost dailj freezing and thawing, leaf cells 

must be subjected to repeated and severe p1i)sical 

stress. Crystals are commonly found in these cells in 

a variety of plants throughout the angiosperms 

(Fahn, 1974). In a few species such as P. reticulafa 

and P. weberbaueri, rhaphites are found scattered 

in the cells of the spong) layer. 
12 SAIITHSOXIAN CONTRIBUTIONS TO BOTANY 

FIGURE B.-Cross sections of Polylepis besseii leaflets under regular (n) and polarized (b) light 

show the in\aginated lower epidermis, the small cell size, and the packed arrangement of the 

cells characteristic of members of the incana complex found at high elevations (X 105). 

INFLORLSCENCES AND FLowERS.-The flowers Of 

all species of the genus are borne on inflorescences. 

In most cases the inflorescences are long enough to 

hang pendantly (Figure Zc), but in the westernmost 

populations of P. tomentella and in at least one 

population of P. pepei, the inflorescence is so reduced that it remains almost hidden in tlie leaf axil. 

Such an extreme reduction (Figure 31) contrasts 

with tlie very long racemes of P. multzjuga (Figure 

8). It appears that the single-flowered racemes represent a reduction that resulted from selection 

against exposed flowers at extremely high elevations. 

Although not investigated in Andean regions, Hedberg (1964) showed that it was common for Afroalpine plants to nestle the flowers in a “protectiLe” 

bed of leaves and trichomes. The reduced inflorescences of Polylcpt~ are surrounded bj denselj pubescent young leaves and stipule sheaths. 

In the species with pendant inflorescences, the 

flowers are borne regularly along the rachis or clustered toward the terminal end. All flowers are subtended by a bract. The flowers themselves are reduced and have many features associated with wind 

pollination (I\71iitehead, 1969). These include: tlie 

absence of petals, green rather than colored sepals, 

an absence of scent or nectar, numeious anthers 

with long filaments, abundant, dry pollen, a large, 

spreading, fimbrillate stigma (e.g., Figure 24, pinnately compound leaLes and the growth of trees in 

stands. The reduced inflorescences of some populations might seem ill-adapted for wind pollination, 

but the reduction must represent a compromise 

between overexposure of vulnerable pollen mother 

cells and adaptations to wind pollination. Species 

with reduced inflorescences normally grow in very 

windy areas. 

The number of stamens is exceedingly variable, 

even within a species. Counts of six to 24 per flower 

have been made for a single species. The anther sacs 

are often red or purple and always have trichomes 

on part or all of the surface. The presence ot 

antheridial trichomes was considered by Bitter 

(191 lb) to be a major character separating Polylzpis 

from Acaena. The trichomes are single celled, 

straight 01- curled, and most commonly restricted to, 

or most dense on, the apex of the sacs. 

The flowers of all species examined are strongly 

protogynous, which suggests that tlie trichomes of 

the anthers serve to insulate tlie developing pollen 

grains while the flower is slightly open and the 

stigma receptive (Figure 26e). After a period of time, 

the flowers open fully, the anthers deliisce longitudinally (Figure 26d) and the stigma withers. Ai 

the pollen is shed, the anther 5acs curl in such a way 

as to turn the anthers “inside out.” The trichomes 

are consequently “inside” the anther at the end of 

anthesis (Figure 2d). Although protogyny would 

tend to increase outcrossing, tlie flowers mature 
NUMBER 43 I3 

FIGURE 7.-Pollen of Polylepis species shows the uniformity of pllen within the genus and the 

similarity of pollen of related members of the Sangisorbeae: a, P. besseri, equatorial view (x 

2100),Simpson 8570 (US); 6, P. besseri, polar view (X 2200); c, P. incana, polar view (x 2100), 

Simpson 8544 (US) ; d, Margyricarpus pinitatus (x 2200) . 

sequentially from the base to the tip of the inflorescence and inflorescences mature at different times 

within a tree. Consequently, unless physiologically 

presented, self-pollination could easily occur. 

The ovary is sunken in a surrounding floral cup 

from which the stigma protrudes. The base of the 

style, where it emerges from the floral cup, often 

bears a ring of trichomes appearing as a tuft after 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

the style has withered. In a few species, trichomes 

extend up the style. Bitter (1911b) described several 

varieties on the basis of the presence of these 

trichomes continuing up the style; however, the 

character is variable within populations of several 

species. 

POLLEN MORPHOLOGY (description provided by 

Joan W. Nowicke).-We have examined the pollen 

of all species of Polylepis recognized here by scanning electron and light microscopy following preparation by acetolysis (Erdtman, 1952). Figure 7a-c 

shows two representative grains. The survey indicated no significant differences between species 

in exine morphology. In addition, widths of pollen 

grains (10 per sample, four samples per species) 

were recorded from acetolyzed grains of four species. 

No significant differences were found in size. In 

fact as illustrated by Figure 7d, pollen of Mal-gyricarpus and Acaena is also exceedingly similar to 

that of Polylepts. Not only would it be impossible 

to separate taxa of Polylepis from one another using 

pollen, but it would also be difficult to separate 

Polylepis from closely related genera on the basis of 

pollen alone. It is possible to distinguish between 

Polylepis and Acaena pollen only by using extremely high magnification (van der Hammen, pers. 

comm.). 

The pollen of Polylepis can be described in the 

following way: Monads, isopolar, more or less 

spheroidal to slightly oblate in shape; (the following 

description of the apertures assumes that the long 

axis is perpendicular to the equator of the grain) 

58-76, wide, 3-colporate, the limits of the endoaperture obscure, the colpus (ectoaperature?) completely covered by a pontoperculuni, the pontoperculum and adjacent sexine protruding at the 

equator to form a pouchlike extension; the nexine 

about equal to the sexine in thickness; the tectum 

slightly undulate to slightly rugose, the surface 

microechinate, the echinae evenly distributed. 

FRuiTs.-The fruits of Polylepis are actually 

achenes composed of the floral cup fused to the 

ovary. Fruits of all species are indehiscent and one 

seeded. The surface of the fruit of different species 

has ridges, knobs, spines, or wings. There are no 

definite sites for the placement of these different 

types of protuberances that appear irregularly over 

the surface. The type of protuberance, wings versus 

spines, or knobs versus wings, is useful for distinguishing between species. 

Bitter (1911b) asserted that the fruits of all species 

must be wind dispersed because members of the 

genus are trees and are thus too tall for animals 

(presumably mammals) to brush against. However, 

the elaboration of spines on the fruits of many taxa 

(e.g., Figure 12e) would argue for animal dispersal 

although wind dispersal undoubtedly predominates 

in P. australis (Figure 38). Numerous birds forage 

or live in Polylepis trees and it is possible that they 

disperse fruits caught in their feathers. There are 

no records of birds eating Polylepis fruits. 

Geographical Distribution 

Except for one series of populations, Polylepis is 

confined to the tropical South American Andes. 

Species range from northern Venezuela to northern 

Chile and adjacent Argentina. One group of extratropical populations is distributed in the mountains 

of northwestern Argentina. Although many species 

grow at extremely high elevations (4000 to over 

5000 m) in habitats that became available for 

colonization only at the end of the Tertiary 

(Simpson, 1975, 1979), several other taxa grow at 

much lower elevations (to 1800 m). The occurrence 

of species at relatively low altitudes mixed with 

montane forest elements indicates that components 

of the genus could have been present in western 

South America in the Miocene or even earlier. 

Radiation and specialization appear to have occurred northward and southward from the vicinity 

of northern Peru and to higher elevations. 

Bitter’s (191 lb) statement that the highest number of species occurs in Colombia, Ecuador, and 

Peru was based on a count of his numerous artificial 

species. Using the taxa recognized here, the concentration of species is in Ecuador/Peru. In the mountains of these two countries, both low elevation, 

unspecialized, and high elevation, specialized, taxa 

occur. Since these groups of species are not closely 

related to one another, it appears that there have 

been several waves of north-south colonization along 

the Andean chain and differentiation has not been 

vertical at points along the Cordillera. 

Geographical distributions of any particular 

species are practically impossible to determine with 

precision for several reasons. First, because of the 

high elevations and specialized microliabitats in 

which species occur, collections are few and tend to 

be concentrated in accessible areas such as R.e.rida 
NUMBER 43 15 

(Venezuela), Pichincha (Ecuador), central Peru, and 

northern Argentina. In addition, and of more serious consequence, trees of most species have been 

cut by man since prehistoric times for firewood or 

building materials. Tfith the recent reexpansion of 

human populations in the high central Andes, cutting has accelerated. In contrast to this destruction 

of natural Polylepis populations is the tendency for 

Andean peoples to plant or shelter trees near houses 

or settlements. These two factors combine to produce an apparent distribution pattern that may 

bear little or no relationship to the original distribution of a species. In the mapping of distributions 

in this treatment, I have used specimen and literature 

data combined with personal observations. Since 

many are old collections (pre-1850), however, current distributions may now differ from those indicated for the various species. 

Ecology 

Without doubt, many species of Polylepis possess 

some of the most unique autecological and synecological relationships of any angiosperm. While 

taxa that grow at relatively low elevations (e.g., 

members of the upper montane forest such as 

P. mziltijiiga, P. pnztta, and P. hieronymi) may have 

no more specialized adaptations than many other 

montane forest trees, those species forming woodlands at elevations between 4000 and 5200 m must 

possess a suite of specializations that allow them to 

withstand the peculiar conditions of these altitudes. 

In addition to formidable elevations, the highest 

habitats also tend to be semiarid since they occur 

on the western Cordillera of the central Andes, 

where mean annual rainfall averages between 200 

and 500 mm. Moreover, tropical areas above 3600 m 

exhibit extreme variation in diurnal temperatures. 

Kight temperatures are often below freezing 

throughout the year while midday temperatures are 

often 10-12°C higher. This kind of climatic regime 

causes the soil below about the top 30 cms to remain 

a constant 24°C (or lower) all year (Walter and 

hledina, 1969). In addition there is no period during the year in which plants can become truly dormant. Instead, they remain active (some are drought 

deciduous) all year, but under exceedingly harsh 

conditions. Theoretically, the daily freezing temperatures, the inability of roots to absorb nutrients 

below the uppermost soil levels that are heated 

during the day, and the presence of a constant, drying wind should preclude tree growth. The occurrence of woodlands at elevations up to 5200 m 

therefore presents a physiological problem. Explanations for the distribution patterns of Polylepis have 

been advanced by several workers. 

The classic studies of Andean geography are those 

of Carl Troll (see Troll, 1959 for references). In 

many of his expositions, Troll specifically discussed 

Polylepis. In all cases concerning species found 

within the puna zone or at the upper limits of the 

ceja (montane cloud forest), Troll considered Polylepis woodlands a distinctive type of vegetation 

different from true ceja, interandean scrub, or puna 

grassland. The ability of Polylepis to grow above 

the montane forest could, in Troll’s opinion, be 

explained by the presence of microclimatic phenomena that produce “lower elevation” conditions. Such 

phenomena include the formation of cloud layers, 

especially at night, on some slopes and along low 

drainage areas, which prevent nighttime freezing 

of the soil. 

In his discussions of Peruvian vegetation, Rauh 

(1950a,b) also indicated that Polylepis grows in 

specific elevational belts on west-facing slopes of 

(primarily) the Eastern Cordillera of Peru. Pulgar 

(1967) described Polylepis as a natural inhabitant 

of only his siini region. This geographical province 

is confined to eastern and western facing slopes 

betwe-n 3500 and 4000 m and contains no large, 

flat valleys. Tliithiii this zone Polylepis is prevalent 

along water courses. While lamenting the fact that 

there is no active reforestation of qziiniuales (PolyZepis woodlands) where they have been cut, Pulgar 

(1967:92) cited the custom (“almost a superstition”) 

practiced by the Altiplano Indians of never cutting 

down an old Polylepis tree but merely exploiting 

the branches. In his discussion of the punu region 

(defined as the treeless flat plateau at 4000 to 4800 

m), he mentioned the presence of Polylepis but 

asserted that individual trees in this zone had been 

purposefully or inadvertently planted by man. He 

gave an example of the robust growth of Polylepis 

trees within the town and arouiitl houses of Cerro 

de Pasco (Figure Ic) as an example of the former. 

In the latter case, trees near houses have resulted 

from sprouting of branches that were brought from 

lower regions and which were able to grow Tvhen 

sheltered by huts and walls (Figure If). 

In contrast to all of these views is the hypothesis 
16 SMITHSOXIAS CONTRIBUTIONS TO BOTANY 

of Ellenberg (1 958a,b) that the high Andean puna, 

particularly that of the Altiplano of southern Peru 

and Bolivia, is a manmade grassland which was 

originally covered by evergreen tree or shrubland. 

The dominant woody taxa of these natural woodlands would have been Polylepis, Buddleia, Piiya 

i~zimondii, and species of Chitqziiraga. To support 

this hypothesis, Ellenberg first drew a parallel with 

the 1fe:literranean region of Europe, which he 

claimed has a similar climate to that of the puna 

and the vegetation of which has been shown to have 

been caused by man. He cited examples of Eitculyptzis trees groxving successfully at elevations of about 

4000 m in the puna. He argued against the restriction of Polylepis to slopes with special microclimatic 

conditions by citing an example of a few trees he 

saw growing on a level area. The absence of trees 

anywhere in abundance except on “inaccessible” 

slopes is attributable, in his opinion, to the cutting 

of trees for firewood, the clearing of woodlands for 

pastureland, and the destruction of seedlings by 

domesticated animals in all other areas. He considered the apparent natural and integrated vegetation of the puna simply the result of immigration 

and establishment of grasses and forbes from the 

few prehuman tree-free areas over the last 400 years. 

IYhile Ellenberg’s hypothesis cannot yet be disproved, it appears less likely than a hypothesis of 

natural localization of woodlands of Polylepis with 

an additional reduction due to human influence. 

As pointed out by IValter and hIedina (1969), trees 

with a root system over a few centimeters deep cannot normally grow at very high elevations because 

of the inability of roots to penetrate or absorb water 

from soil that is between 0” and 4” C. Special situations are necessary to modify the usual prohibitive 

conditions. IValter and Jledina (1969) proposed that 

Polylepis in Venezuela grows in rock areas rvhere 

the crevices between the rocks allow warm air to 

reach considerable depths and hence allow root 

penetration and function as low as one meter or 

more. Recently, Smith (1 977) tested this suggestion 

by planting seedlings of P. sericea (the native 

species) in open paramo and sheltered rocky areas 

at 3600 m in JIerida, Venezuela. Yo seedling survived one year in exposed areas. Seven trees (147,) 

planted in rocky areas were surviving at the time 

of the report (one year after planting). 

It is obvious that an enormous amount of decimation and vegetative change has been caused by man 

in the high central Andes, but Ellenberg’s arguments are untenable on several other grounds. His 

examples are, for the most part, taken from artificial 

situations. Eucalyptus is an introduced genus in 

South America and the trees growing at high elevations have been carefully nurtured and protected. 

His comparison with the Mediterranean region is 

misleading in view of the enormous differences in 

climate. Despite the similarities in mean annual 

rainfall and mean annual temperature (the two 

parameters he gives) between the two areas (Walter 

et al., 1975), the dispersion of the precipitation and 

the annual versus diurnal temperature patterns are 

completely different. Around the Mediterranean, 

most of the precipitation falls in the winter and the 

area is typically temperate in that summers are 

warm and winters cool or cold. In the puna, rainfall 

is primarily during the southern summer. At any 

given hour of the day at a locality on the Altiplano, 

temperature is practically constant throughout the 

year, while within any one day there is as much 

as a 12OC temperature difference. Bliss (1956) has 

pointed out that there are great physiological differences between arctic and alpine plants (i.e., 

latitude and altitude vegetation types with similar 

physiognomies), even though they are similar in 

visual aspect. 

Descriptions of many botanists and geographers 

who have worked in the Andes (Jameson, 1845; 

Hertzog, 1923; Weberbauer, 1945; Troll, 1959; 

Hueck, 1972) clearly indicate that Polylepis woodlands are, in the opinions of the authors, a specialized type of vegetation restricted to certain areas 

such a5 rocky slopes, river courses, valley bottoms, 

or slopes normally covered by cloud banks. Koepcke 

(1 961) particularly opposed Ellenberg’s views. He 

logically pointed out that many areas (such as shown 

in his fig. 59 and in Troll, 1959, figs. 17 and 19) are 

accessible to man but still have flourishing Polylepis 

woodlands. He interpreted the surviL a1 of trees 

around houses on the Altiplano not as an indication of their ability to grow naturally at these elevations, but rather as a duplication by man of the 

normal specialized microhabitat conditions (i.e., protection from the wind, relatively warm soil conditions and, usually, increased water availability) of 

areas in which they naturally occur. 

The only studies that shed any light on the possible paleohistory of Polylepis are those of van der 

Hammen and colleagues (van der Hammen, 1974 
NUMBER 43 

and references therein) in Colombia. Because Polylepis species are all wind pollinated, pollen cores 

provide a comparatively accurate record of their 

history in a given area. Cores from the Sabana de 

Bogota and Laguna Fuquen, Colombia, spanning 

almost three million years provide such a record. 

The core at Bogota clearly shows the first appearance of Polylepis in the area at about 600,000 years 

ago. Since that time, the fluctuating climates of the 

Pleistocene caused expansions and contractions of 

local Polylepis populations. During a warm wet period (about 30,000 years ago), the area from which 

the pollen cores were taken was dominated by Polylepis woodlands. Shortly afterward, as conditions 

became colder, low paramo elements became more 

abundant until, in drier periods, from 21,000 years 

ago, trees were so rare that essentially no grains 

appear in the cores. In the latest cores from Fuquen 

(10,820 YBP to present), the climate was warmer, 

but Polylepis never regained dominance, in part, 

perhaps, because the climate was already dry but 

also because of cutting and burning by man. Xevertheless, it is obvious that even at the low elevation 

(2650 m) of Bogota, the distribution of Polylepis 

fluctuated during the Pleistocene and that a complete coverage of this high Andean area by woodlands was an uncommon event and has not occurred 

since 30,000 YsP-long before the active influence 

of man. 

In addition to ecological arguments, it is hard to 

explain the differentiation of distinct taxa on the 

Eastern and Western Cordilleras as well as northsouth along the Andean chain if there were a 

continuous woodland of Polylepis covering the 

mountains. Under circumstances of a more or less 

continuous cover, clinal variation should, at most, 

develop. However, until the physiology of Polylepis 

is more completely understood and the microclimatic conditions of the areas in which it now occurs 

more thoroughly investigated, the question of the 

potential distributional limits of the genus will remain unsettled. In any case, it will be impossible 

ever to know with certainty the natural prehuman 

distribution of the genus. 

Polylepis Ruiz and Pavon 

Polylrpis Ruiz atid Pavon, 1794:80, Icon 15. [Type-species: 

Polylepis racenosa.] 

17 

Deciduous or evergreen trees or shrubs, 1-27 m 

tall. Bark of the trunk and larger branches deep 

red, many layered, the layers thin and exfoliating. 

Branching repeatedly sympodial with long, often 

bare branch segments and leaves slightly to strongly 

congested at the tips of branches. Leaves alternate, 

imparipinnate with 1-9 pairs of leaflets, the first to 

the third pair from the apex the largest, rhombic, 

trullate or obtrullate in outline, 0.8-9.1 cm wide, 

2-19 cm long, rachises sericeous, villous, with scattered twisted trichomes, glandular or glabrescent; 

point of leaflet attachment with a tuft of straight 

or twisted long trichomes, a ring of glandular trichomes, or both; stipules fused around the branches 

forming a sheath; the sheath with spurs on either 

side of, or sloping away from, the petiole, abaxial 

surface sericeous, villous, pubescent, glandular or 

glabrescent, apex with long trichomes often projecting from the inner surface. Leaflets lanceolate, 

ovate, oblong, or obovate in outline, 0.2-1.7 cm 

wide, 0.6-3.7 cm long, margins entire, revolute, 

crenate or serrate, apex acute, obtuse, emarginate 

or tridentate, base unequally cordate or attenuate; 

upper surface glabrous, glabrescent, minutely sericeous, hispidulous or with scattered niulticellular 

trichomes (often most dense in the midvein depression); lower surface sericeous, minutely sericeous, 

hispidulous, villous, pannose or glandular, often 

with glandular trichomes mixed with another type, 

sometimes with a pannose covering under another 

layer. Inflorescences upright or pendant to 36 cm 

long, bearing 1 to over 60 flowers, the flowers closely 

packed or spaced along the rachis; floral bracts 

lanceolate to ovate, to 3 mm long, scarious to variously covered with trichomes; raches sericeous, villous, pilose, pannose, glabrescent, or glandular. 

Flowers perfect (perhaps functionally female in 

some plants due to abortion of the anthers), 0.4-1.0 

cm wide, sepals 3 or 4, lanceolate to ovate in outline, sericeous, villous, pilose or glandular on the 

outer face, sometimes pannose on the tips of the 

inner surface, persistent; stamens (4)) 6-36, basifixed, inserted on the floral cup at the base of the 

style; tomentow. or barbate on the anther sacs, 

longitudinally dehiscent; style villous or hispid at 

the base, occasionally villous along most of the style; 

stigma peltate, orbicular, flat, expanded, fimbrillate; 

ovary inferior, rarely protruding from the floral 

cup; carpel 1, ovule 1, pendulous. Fruit an achene, 

cylindrical, fusiform, turbinate or globose, variously 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

adorned with protuberances, spines, or wings, body 

and protuberances sericeous, pilose, lanose, villous, 

with scattered irregular single-celled or multicelled 

trichomes, glandular or glabrescent, from 0.2-1.4 cm 

wide including protuberances, 0.2-1.3 cm long. Seed 

pendulous, the testa membraneous and the cotyledons flat or slightly convex. 

DISTRIBUTION.-h the Andes at elevations of 

1800-5200 m from Venezuela in the state of Lara 

south to Tarapaca, Chile, and Cordoba, Argentina. 

Key to the Species of Polyepsis 

1. Undersurface of leaflets covered with shiny, silky trichomes, the sericeous trichomes sometimes most prominent on the veins: or covered with dense, straight, short, appressed trichomes, 

or with long (> 2 mm) shaggy trichomes sometimes covering a shorter layer, not glabrous. 

2. Undersurface of leaflets with silky trichomes or short, straight, dense, appressed trichomes. 

3. Trichomes very silky and covering entire under leaf surface, long, dense. 

4. 

4. Pubescence golden red or white; leaflets obovate or broadly oblong: apex 

the veins of the undersurface of leaflets 

or short, straight, appressed, and overlying a lower layer of very short, matted trichomes. 

5. Lea!.es with 4-9 pairs of leaflets; leaflet$ acute or obtuse at the apex, ovate, oblong 

5. Leaves with 1-3 pairs of leaflets; leaflets emarginate at the apex, obovate or narrowly oblong, margins entire. 

6. Undersurface of leaflets dull with short, appressed, straight trichomes over entire 

surface and olerlying a layer of short, twisted trichomes; leaflets robust, yellowish margins crenate toward the apex; fruit more or less cylindrical with irregularly flattened spines ............................................................... 6. P. subsericans 

6. Undersurface of leaflets shiny with short, silky trichom 

on the veins and leaf margins; leaflets thin, green, margins entire, fruit elongate, 

2. Undersurface of leaflets with loose, shaggy, long (> 2 mm) trichomes that can cover 

shorter twisted or multicellular glandular trichomes. 

7. Leaves with 6-8 pairs of leaflets: inflorescences 17-36 rm long, flowers numerous, 

usually over 20 per inflorescence, densely packed; stipule sheaths strongly spurred ........ 

7. Leaves with 3-4 pairs of leaflets; inflorescences 3-12 cm long, flowers fewer than 20 

(occasionally 24) scattered: stipule sheaths lacking true spurs (although trichomes at 

the apex can produce the impression of a pointed spur). 

8. Stipule sheaths with prominent long, silky, white trichomes on the outer surface 

extending down the internode; leaves with 3-4 pairs of leaflets; undersurface of leaf. 

lets densely covered with shaggy white scattered trichomes ................ 3. P. hieronyrni 

8. Stipule sheaths sparsely villous on the outer surface and with long twisted trichomes 

projecting at the apex from the inner face near the petiole; leaves with 1-3 pairs 

of leaflets; undersurface of leaflets sparsely covered wth shaggy fulvous trichomes 

originating from the veins and with scattered glandular trichomes or protuberances 

......................................... 14, P. racernosa 

I. Undersurface of leaflets covered with 2 mm), twisted, matted, feltlike, white 

or yellowish trichomes or with scattered, short (< 2 mm) twisted, 1-celled trichomes, the 

short trichomes never covered by longer trichomes, but in cases often mixed with multicellular glandular trichomes, or glabrou?. 

9. Undersurface of leaflets lacking scattered yellow, multicellular glandular trichomes, pannose or lanulose, not glabrous; plants of Colombia and Ecuador or northern Peru. 

10. Revolute leaflet margins and undersurface of leaflet apex with short, straight trichomes 

distinctive from the pannose covering of most of the undersurface of the leaflet: 

leaves with 2-3 pairs of leaflets ............ ............. 8. P. reticulafa 

10. Revolute leaflet margins and undersurface of the leaflet apex uithout a distinctive 

type of trichome; leaves with 3-3 pairs of leaflets. 

Pubescence silvery or whitish; leaflets ovate or elliptic; apex obtuse ........ 5. P. sericea 

emarginate .................... .......................................... 2. P. Zanugimsa 

3. Trichomes confined to, or 

or lanceolate, margins serrate ................ .................................... 4. P. pauta 

slightly twisted, knobbed around the middle ............................................ 7. P. pepei 

..................................................................................................... 1. P. rnultijuga 

.................................................... 
NUMBER 43 19 

11. 

11. 

Upper surface of leaflets very nitid, when young covered with long, shaggy white 

trichomes; stipule sheaths with acute spurs, inflorescences 5.8-14 cm long with 

8-18 flowers; fruits with flattened spines: small trees or shrubs (less than 2 m tall) 

of southern Ecuador and northern Peru ....................................... .9. P. weberbaueri 

Upper surface of leaflets rugose or slightly nitid, occasionally with scattered long 

trichomes; young leaflets without conspicuous white trichomes; stipule sheaths 

obtuse at the apex; inflorescences 2.1-5.4 cm long with 3-7 flowers; fruits with 

thin, terete spines; trees 5-8 m tall of Colombia and Ecuador .... 10. P. quudrijuga 

9. Undersurface of leaflets with yellow multicellular glandular trichomes and resinous exudate 

mixed with short matted trichomes, or glabrous; plants of Ecuador, Peru, Bolivia, or 

Argentina. 

12. Undersurface of leaflets glabrous. 

13. Fruits strongly winged, the wings as wide as the body of the fruit; leaflets broadly 

lanceolate, almost serrate ...................... 15. P. australis 

13. Fruits with irregular ridg leaflets obovate or oblong, 

crenate or entire. 

14. Fruit with irregular ridges, sometimes flattened into narrow wings; plants 

of Bolivia .......................................................................................... 12. P. tomentella 

14. Fruit spiny: plants of Colombia, Ecuador or Peru. 

15. Apex of leaflets obtuse or acute, iptic to narrowly ovate, 

..................... 5. P. sericea 

15. Apex of leaHets emarginate, leaflet shape broadly ovate, fruits with irreg12. Undersurface of leaflets with short matted trichomes, glandular trichomes, or both. 

16. Undersurface of leaflets with dense, matted trichomes usually mixed with glandular trichomes most prominent on the veins and leaf margins; plants of southern 

Peru, Bolivia, northern Chile, and northern Argentina. 

17. Undersurface of leaflets with matted gray or white trichomes; stipule sheaths 

pubescent: fruit with spines prominent at the apex and projected upward, 

sometimes flattened at the base ............................................................ 11. P. besseri 

17. Undersurface of leafiets with a dense covering of matted white or yellowish 

trichomes; stipule sheaths with glandular trichomes at the apex, glabrous 

below; fruits with short, irregularly positioned blunt spines or blunt spines 

flattened and coalesced into a narrow, wavy wing extending the length of 

the fruit .............................................................................................. 12. P. tomentella 

16. Undersurface of leaflets with sparse, scattered trichomes and irregularly scattered 

yellow glandular trichomes; plants of Ecuador and central Peru ........ 13. P. incam 

fruits with knobs or stout spines 

ular thin spines ........................................................................ 10. P. quadrijuga 

1. Polylepis multijuga Pilger 

Polslepis rriultijuga Pilger, 1906:536. [Type collection: at 

Chugar near Hualgayoc, Cajamarca, Peru, 2700-3000 m, 

May 1904, Weberbauer 4098. Type destroyed, isotypes 

WRSL, G!] 

Trees 4-15 m tall. Leaves only slightly congested 

at the ends of the branches (Figure 8a), imparipinnate with 6-8 pairs of leaflets, obtrullate in outline, 

6.4-9.1 cm wide, 12.2-19.5 cm long; rachises lanose, 

point of leaflet attachment (Figure 8c) with a tuft 

of long, straight trichomes (some with stalked basal 

cells) mixed with small, clavate, multicellular glandular trichomes; stipular sheaths acute at the apex 

with spurs reaching 1 cm in length, densely sericeous on the outer surface. Leaflets broadly lanceolate or ovate in outline, second or third pair from 

the terminal leaflet the largest, one of this pair 

1.1-1.7 cm wide and 3.0-3.6 cm long; margins serrate; apex acute, base unequally cordate; upper leaf 

surface sparsely villous in the midvein depression; 

lower surface densely lanose with individual trichomes 1-2 cm long (Figure 8b). Inflorescences pendant, 17.3-36.0 cm long, bearing numerous (often 

more than 45) flowers; floral bracts small, sericeous 

on the outer surface; rachises sericeous. Flowers perfect, 0.4-0.8 cm in diameter (Figure 84; sepals 3 

or 4, ovate, lanose on the outer surface; stamens 

5-16, circular in outline, tomentose; style hispid at 

the base. Fruit more or less fusiform, sericeous, with 

irregular spines up to 4 mm long (Figure 8e), sometimes with long trichomes on the spines, 0.6-1.1 cm 

wide including spines and 0.3-1.0 cm long. 

DIsTRisuTIoN.-Northern Peru in the provinces 
20 ShfITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE S.--Polylepis multijugn: a, branch (X 0.5); 6, undersurface of leaIiet (x 3); c, point of 

leaflet attachment (X G): d, flower (X 6); e, fruit (x 6). 
NUMBER 43 21 

of Amazonas and Cajamarca (Figure 9). Flowers 

recorded in May, July, and from September to 

December. 

COMMOIV NAME.-Quinua.

UsEs.-The red heartwood is reported to be used 

for furniture manufacture. 

REPRESESTATIVE SPECIMESS EXAMINELL-PERU, AMAZONAS: 

Chacapoyas, Matthews s.n. (K). CAJAMARACA: Hualgayoc, 

Hacienda Taulis, 13 km heyond Palmito Junction toward La 

Playa, 2900 m, 2 Sep 1964, Hutchison & Bisniark 6463 (F, 

MO, SY, P, UC), 3200 m, 15 Oct 1954, Rauh & Hirsch P2249 

(XY). 

DIscussIoN.-In the small restricted area of northern Peru in which it grows, Polylepis multijuga 

occurs as low as 2500 m mixed with species of 

tree ferns, Podocarpus, Chusquea and Berberis, its 

branches often covered with Tillandsia and epiphytic orchids. Bitter considered this species to be the 

most primitive of the genus. Numerous characters 

involving morphology, anatomy, and ecology seem 

to reinforce this conclusion. The leaves are large, 

thin, and reminiscent of those of some species of 

Acaena. The inflorescences are large and pendulous, 

presumably the ancestral type for the genus. The 

I~URT 9.-Collection localities of Poly!epis multijuga. 

fruits, like those of all of the members of this 

species group, and species of Acaena, are spiny and 

appear to be dispersed by animals. More advanced 

taxa have become increasingly dependent on wind 

as a dispersal agent. Finally, the species is an integral member of the upper montane rainforest 

(ceja de la montana), rather than a fringe species 

or one that grows completely isolated from the 

montane forest in the higher elevations. 

The species most similar to Polylepis multijuga, 

P. pauta, can be distinguished from it by several 

leaf characters. The leaves of P. multijuga are larger 

than these of P. pauta, the lower surfaces are covered 

by wooly rather than silky hairs, and the tricliomes 

at the point of leaflet attachment often have multicelled bases. In addition, the apices of the stipule 

sheaths of P. multijuga are larger and more rounded 

than those of P. pauta and the inflorescences, in the 

specimens I have seen, are much larger, both in 

length and in the number of flowers they bear. This 

species differs from P. lanuginosa in having more 

numerous leaflets, obtuse rather than emarginate 

leaflet apices, and a lanose rather than sericeous 

under leaflet covering. 

2. Polylepis lanuginosa Humboldt, Bonpland, 

and Kunth 

Polylepis lanuginosa Hurnboldt, Bonpland, and Kunth, 1824: 

228. [Type collection: near Calpi, Chimborazo, Ecuador, 

June, Bonpland 2191. Holotype P!, isotype P!] 

Polylepis lehrnannii Hieronymus, 1895:29. [Type collection: 

west of Cuenca, Azuay, Ecuador, Lehtnarin 6487. Type 

destroyed, Rockefeller photograph no. 3387, types of the 

Berlin Herbarium, isotypes GH! NY!] 

Polylepis coriacea Bitter, 191 lh:603. [Type collection: I’alley 

of Pangor, Ecuador, Spruce s.72. Holotype IV!] 

Small trees (?) to 4 m tall. Leaves fairly congested 

at the ends of the branches, imparipinnate with 

2-3 pairs of leaflets (Figure IOU), obovate in outline, 

2.9-4.2 cm wide, 4.5-7.2 cm long; rachises villous; 

point of leaf attachment with a small tuft of 

long straight trichomes; stipular sheath with acute, 

densely sericeous spurs, the hairs red gold in color. 

Leaflets (Figure 10a) obovate or oblong in outline, 

first or second pair from the apex the largest, one 

of this pair 0.9-1.4 cm wide, 2.0-3.1 cm long; margins crenate or revolute; apex emarginate; base 

unequally cordate, upper leaf surface rugose or dull, 

sometimes with trichomes scattered in the vein de- 
22 SSIITHSONIAN CONTRIBUTIOSS TO BOTANY 

FIGURE IO.--Polylepis lanuginosa: a, branch (x 0.5); b, lower leaflet surface at point of leaflet 

attachment (x 3); c, flower (x 6); d, fruit (x 6). 
NUMBER 43 23 

pressions; undersurface densely sericeous, the trichomes white or golden red, often with a covering of short wooly trichomes under the silky hairs. 

Inflorescences (Figure 10a) pendant, sometimes 

branched, 0.9-4.4 cm long, bearing 4 to 20 flowers; 

floral bracts lanceolate, sericeo-lanose; rachises lanose. Flowers (Figure 1Oc) perfect, about 0.7 mm 

in diameter, sepals 4, ovate, outer surface villous; 

stamens about 12, lanose at the apex; base of style 

hispid, lower part villous. Fruit (Figure 10d) fusiform, irregularly spined, the spines flattened or thin, 

usually villous, body of the fruit pilose; 0.5-0.9 cm 

wide including spines, 0.3-0.6 cm long. 

DIsTRIsvTIoN.-Apparently restricted to central 

and southern Ecuador in the provinces of Chimborazo, Azuay, and CaNar (Figure 11). Flowering 

recorded April to June. 

COMhlON 1\JAME.-QUifiNl. 

Usm-None specifically reported. 

REPRESENTATIVE SPECIMENS ExAMINED.-ECUADOR. AZUAY: 

west of Patul, 3 km between Huahualcay and Rio Patul 

below Pasas de Pinglion, 267CL3275 m, 19 May 1943, Steyermark 52599 (F). CARAR: between Tamho and Suscal, north 

rim of the valley of the Rio de Caniar, 2000-3000 m, 23 Apr 

1945, Giler 2773 (F, K, NY); Canar, 15 Sep 1918, Rose 6 Rose 

23789 (NY, US). 

FlCURr 11 .-Collcction localities of Polylepis lanuginosa. 

DIscvssIoN.-Although similar in many morphological characters and probably derived from Polylepis multijuga, P. lanuginosa differs from it and all 

other members of the genus in its possession of 

branched inflorescences. In addition to this character, it differs from P. multijuga in leaflet pubescence, leaflet shape, and vestiture of the point of 

leaf attachment. The leaflets of P. lanuginosa are 

emarginate, the upper surface is rugose, the lower 

surface has a mixture of short wooly trichomes overlain by dense silky hairs. Leaflets of P. multijuga 

are acute, the upper surface flat, and the lower surface densely lanose. The point of leaflet attachment 

in the former is a small tuft of straight trichomes 

and in the latter a ring of glandular hairs mixed 

with long straight trichomes. The two species also 

have fruit differences: P. multijuga has terete spines 

on the fruit and P. lanuginosa has flattened spines. 

There has been some confusion in the characterization of this taxon attributable in part to the 

specimens cited by Weddell in the Chloris Andina. 

Weddell’s brief verbal description could be applied 

to many members of the genus, but he cites two 

specimens, neither of which belong with the type 

cited by Humboldt, Bonpland, and Kunth. Weddell’s first specimen was the one he presumably 

considered the “true” P. lanuginosa; the second 

specimen corresponds to the variety j3 microphylla. 

Bitter correctly realized that neither of these specimens corresponded to the original P. Eanuginosa 

and consequently made the first specimen the type 

of his new species, P. quudrijuga, and the second the 

type of P. m icrophylla. Nevertheless, Weddell’s 

older application of the name P. lanuginosa was 

followed by many botanists and has often been misapplied to specimens of P. quadrijuga or P. weberbaueri. 

3. Polylepis hieronymi Pilger 

Polylepis hieronymi Pilger, 1906:535. [Type collection: Salinas, 

Cuesta de Pollo, l’alle del Tambo, Bolivia, June 1873, 

Lorentz 6 Hiel-onymus 938a. Type destroyed, Rockefeller 

photograph no. 3355, types of the Berlin Herbarium at NY.] 

Polylepis iacemosa p hypoleuca Weddell, 1861:238. [Type 

collection: hetween the cities of Tarija and San Luis, 

Tarija, Bolivia, Jul-Aug, 1846, Weddell 4607. Lectotype P!] 

Polplepis racemosa var. albotometitella 0. Kuntze, 1898:77. 

[Type collection: Sierra de Cordoba, Los Gigantes, 

Gordoba, Argentina, Kurtz 6926. lsotypes SY!] 

Polylepis hypoleuca (Weddell) Ritter, 191 lb:607. [Type col- 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 12.--Po[ylepis hieronpmi: a, branch (X 0.5); b, upper (right) and lower (left) leaflet 

surfaces (x 3); c, lower leaflet surlace at point of leaflet attachmeiit (X 6); d, flower (X 6; e, 

fruit (x 6). 
NUMBER 43 25 

lection: based on the same type as P. racemosa p hypoleuca 

Weddell.] 

Polylepis hieronymi var. dolicholopha Bitter, 191 lb:609. [An 

illegitimate name since it includes the type of the species 

and, under the present International Code of Botanical 

Nomenclature, the correct name must be P. hieronymi var. 

hieronymi.] 

PolyZepis hieronsmi var. saltensis Bitter, 1911h:609. [Type 

collection: near Pampa Grande, the pass “El Alizar,” Salta, 

Argentina, 2400-2600 m, 1900, h’elson 12584. Holotype S!] 

Polylepis australis var. bijuga Bitter, 191 1 b:624. [An illegitimate name, based on the type of P. racemosa var. albotomentella 0. Kuntze.] 

Small trees to 3 m tall. Leaves (Figure 12a) congested at the ends of the branches, imparipinnate 

with 3 to 4 pairs of leaflets: obtrullate in outline; 

variable in size, from 1.9-5.2 cm wide and 2.9-9.7 

cm long; rachises sericeolanose; point of leaflet attachment with a band of small, dark-colored multicellular glandular trichomes, appearing as a dark 

ring around the rachis or seemingly undifferentiated 

from the rest of the rachis but with small glandular 

trichomes under the lanose covering, sometimes with 

a few long trichomes with one or two basal cells: 

stipular sheaths usually sloping away (downward) 

from the leaf petiole but sometimes with a slight 

shoulder, without spurs but sometimes appearing 

spurred because of the projection of long trichomes 

on the abaxial surface, sericeous on the outer face 

with the long, white silky hairs sometimes extending 

down the internode. Leaflets (Figure 12a) ovate in 

outline, the first pair below the terminal leaflet the 

largest, one of this pair 0.5-1.2 cm wide and 1.2-3.2 

cm long; margins crenate, apex obtuse; base attenuate or cordate, the pairs of leaflets often slightly 

FIGURE 13.-Collection localities of Polylepis hieronymi. 

obliquely positioned so as not to appear paired, 

terminal leaflet petiolate; upper surfaces (Figure 

12b) rugose, usually sparsely villous; lower surfaces 

(Figure 12b) densely lanose with long, twisted trichomes. Inflorescences pendant, 4.9-12 cm long, 

bearing 8-24 flowers; floral bracts conspicuous but 

small, to 1 mm, strigose on the outer surface; 

rachises tomentose. Flowers (Figure 12d) perfect, 

0.6-0.9 cm in diameter; sepals 4 (sometimes 3), oval, 

sericeous on the outer face; stamens 9-13, orbicular, 

upper half with a dense beard of straight, yellow or 

white trichomes; stigma spreading; point of style 

attachment slightly hispid. Fruit (Figure 12e) almost 

cylindrical with long, sometimes hooked, spines; 

body of fruit tomentose, spines occasionally tomentose, 0.3-0.9 cm wide including spines, 0.5-1.0 cm 

long. 

DIsTRIBuTIoN.-SoUthern Bolivia (Tarija) and 

northern Argentina on the upper slopes of the 

Eastern Andes (Figure 13). Flowering recorded in 

March, June, July, August, and September. 

COMMON NAME.-QQuenoa.

UsEs.-None specifically reported. 

REPRESENTATIVE SPECIMEXS EXAIfINED.-BOLI\’IA. Valle 

del Tamho, Salinas, 10 Jun 1873, Lorentz L- Hieronymus 938 

(G, NY, S). TARIJA:O’Connor; Quebrada de Canaletas, between Tarija and Entre Rios, 2300 m, 15 Ju1 1954, Meyer 

18213 (LIL); without precise locality, Mar 1890, Bang 769 

(F, GH, MO, NY, US). ARGENTIN’4. JUJUY: Estacion 

L’olcan, Tiraxi, 2300 m, 27 Jan 1927 Castillon 388 (LIL): 

Tumbaya, Volcan, 3000 m, 13 Feb 1927, Venturi 9002 (US). 

SALTA: Dept. Guachipas Pampa Grande, lomas del Zapallar, 

1810 m, 1 Sep 1949, Hueck 260 (LIL). 

DIscussIoN.-Collections of Polylepis hieronyrni 

are scarce despite the statement by Hueck (1972) 

that this species is a very common tree forming a 

band of woodlands at the upper reaches of the ceja 

of the central parts of the Eastern Cordillera. An 

examination of the few available specimens indicates that different trees vary in leaf size. The Jujuy, 

Argentina, specimens I have examined have the 

smallest leaves, whereas those from Salta have the 

largest. 

On the basis of the long, relatively unmatted, 

cottony pubescence of the under leaflet surface, the 

presence of clavate multicellular glandular trichomes at the point of leaflet attachment, and the 

large, narrowly spined fruits, P. hieronynii would 

appear to belong to the P. sericea group. The 

two species, while similar in some respects can, how- 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ever, easily be told apart. Unlike P. multipora, there rather than straight, spines on the fruit of P. hierare no spurs on the stipule sheaths of P. hieronymi. onymi seem unique in the genus. Bitter (1911b) 

The leaves of P. mullijuga also have many more pointed out the similarity of the hooked spines of 

pairs of leaflets and there are many more this species to the glochids of some Acaena species, 

flowers per inflorescence than in P. hjeronymi. The but it is doubtful if there is any homology in the 

exceedingly narrow fruit and sometimes hooked, structures. The spiny rather than alate fruits and 

FIGURE 14.--PoZylepis pauta: a, branch (X 0.5): b, underleaflet surface (x 2); c, point of leaflet 

attachment, upper surface (X 6); d, point of leaflet attachment, lower surface (x 6); e, flower 

(X 6); 1, fruit (X 6). 
NUMBER 43 27 

the densely sericeous stipule sheaths distinguish this 

species from P. besseri and P. racemosa. 

4. Polylepis pauta Hieronymus 

Polylepis pauta Hieronymus, 1896:313. [Type collection: Corredor Machai, Ecuador, 3900 m, 1871, Stubel 23% Type 

destroyed, Rockefeller photograph no. 3390, types of the 

Berlin Herbarium at GH, SY.] 

Polylepis serrata Pilger, 1906:536. [Type collection: Huamalics, southeast of Monzon, Huanuco, Peru, 3400-3500 m, 

1903, Weberbuuer 3354. Type destroyed, Rockefeller photograph no. 3394, types of the Berlin Herbarium at GH, NY, 

Polylepis serrntn var. psilanthera Bitter, 1911b:593. [An illegitimate name since the original circumscription includes the 

type of the species.] 

Poljlepis serrnta var. paricipiln Bitter, 191 1b:593. [Type collection: Convencion, Yanamanche, between Cuzco and 

Santa Aria, Cuzcu, Peru, 3500-3800 m, Weberbauer 4854. 

Type destroyed, isotype WRSL.] 

Polylepis annulatipilosa Bitter, 191 lb596. [Type collection: 

Andes of Quito, Pichincha, Ecuador, Jameson 16. Lectotype 

W!, isotypes G! GH!] 

Trees (2) 5-27 m tall. Leaves slightly congested at 

the ends of the branches, imparipinnate with 4-9 

pairs of leaflets (Figure 14a), obtrullate in outline, 

2.4-6.6 cm wide and 3.4-11.7 cm long; rachises 

lanose; point of leaflet attachment hispid on the 

upper surface, the long (to 3 rnm) trichomes (Figure 

14c) usually mixed with multicellular, glandular 

trichomes and resinous exudate; stipular sheaths 

red brown, sericeous on the outer face. Leaflets 

(Figure 14b) ovate, acute at the apex, oblong or 

lanceolate in outline, second pair from the terminal 

leaflet usually the largest, one of this pair 0.6-1.4 cm 

wide and 0.8-3.3 cm long; margins serrate; apex 

acute; base unequally cordate; upper leaflet surface 

glabrous or slightly tomentose, particularly in the 

midvein depression; lower surface sericeous with the 

white or gold trichomes completely restricted to, or 

most conspicuous on, the veins. Inflorescences pendant, 6-21 crn long, bearing 16 to over 60 flowers; 

floral bracts conspicuous, pilose on the outer surfaces; rachises sericeous. Flowers (Figure 14e) perfect 

except possibly for some plants from Peru (Cuzco), 

which appear to be male sterile, 0.3-0.6 cm in diameter; sepals 3 or 4, ovate, green, outer surface sericeous; stamens 5-12, red, sericeous; stigma expanded, 

style hispid at the base. Fruit (Figure 14f) irregularly 

shapzd, rhomboid to cylindrical with numerous, 

unequal, glabrous to villous, thin spines, 0.3-1.4 cm 

wide including spines, 0.3-1.2 cm long. 

US.] 

DISTRIBUTION.-From north-central Ecuador (Pinchincha) to southern Peru (Cuzco), primarily in the 

eastern Andes (Figure 15). Flowering and fruiting 

recorded in February, May, July, August, September, and December. 

COMMON NAMES.-Pauta, quiniual, quena, queuna. 

UsEs.-None recorded. 

REPRESE~TATIVE SPCIMENS ExAhfINED.-ECUADOR. PI’APO: 

Mount Guamani, 4000 m, 22 Sep 1939, Asplund 8767 (S). 

PERU. SAN MARTIN: Huallaga, Valley of the Rio Cipisoncho, 

30 km above Jucusbamba, io55’S, 77OlO’W, 3600 m, 22 Aug 

1965, Hamilton 2 Holltgun I49 (K, S). Cuzco: Paucartamho, 

Chacapampa, 1800-2000 m, Dec 1950, no collector 9909 (510); 

Uruhamba Valley, Torontoy, 1 May 1915, Bingham 557 (US); 

Quispicnachi, Marcapata, 15-16 Feb 1929, Weberbauer 7803 

(BM, F, G, GH, K, NY, US); Convencion, Panticalla, 3800 

m, Ju1 1949, Martn i608 (UC, US). APURIMMAC: Cordillera 

Salcantyl, 3600 m, 30 May 1969, no rollector TI467 (NY). 

DIscussIox.-Like Polylepis multijuga, P. pniita 

exhibits many unspecialized characters such as the 

size of the plants (large trees), thin, large, and numerous leaflets and long inflorescences with numerFIGURE 15.-Collection localities of Polylepis pauta. 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ous flowers. This species appears to grow in relatively inesic habitats, usually mixed with, or at the 

upper edge of, the montane forest. Specimens have 

been collected as low as 1800 m, the lowest elevation 

recorded for any species of the genus within the 

tropics. 

RIorphologically, Polylepts pauta is intermediate 

in many characters between P. rnultijuga and P. 

lariugtnosa on the one hand and P. sericea on the 

other. It differs from the former two taxa b) having 

smaller leaves and inflorescences, a reduced pubescence on the sheaths and rachises of the leaves and 

on the inflorescences and a restriction of the pubescence on the underleaf surfaces to the veins. Although 

very similar in almost all characters to P. serzcea, the 

sympatiy of P. pauta with P. sertcea (on Volcan 

Pichincha, Ecudor, and in the Urubamba Valley, 

Torontoy, Peru) with no intermediates indicates 

the two are distinct taxa. The two species can be 

distinguished by the differences in leaf margins 

(seirate in P. pauta) and the fruit morphology (fruit 

bod) thinnei and the spines thinner and longer in 

P. paiita than in P. seiicea). 

5. Polylepis sericea Weddell 

Polylepis sericea Weddell, 1861:238. [Type collection: Sierra 

Sevada, Iierida, T’enezuela, 11,000 ft (3500 m), June 1847, 

Funck L- Schlirn 1546. Lectotype P! isotype G!] 

Acnena ochrenta Weddell, 1861 :240. [Type collection: Andes 

around Quito, west slopes of Pichincha, Pichincha, Ecuador, 

12,000 ft (3600 m), April to May 1856, Jaineson 73, Lectotype P! isotypes GI GH!] 

Polylepis Stuebelii Hieronymus, 1896:313. [Type collection: 

wesern slope (if Mt. Quilindana near Barnhasacha, Ecuador, 

3700 m, Stuibel 204. Type destroyed, Rockefeller photograph 

no. 3399, types of the Berlin Herbarium at NY.] 

Polylepis albicans Pilger, 1906:535. [Type collection: Cordillera Blanca above Caraz, Ancash, Peru, June 1903, Weberbutler 3229. Type destroyed, Rockefeller photograph no. 

3383, types of the Berlin Herbarium at GH, SY.] 

Polylepis ochreata (lVeddel1) Bitter, 1911a:5. 

Polylrpis ochreata ur, integra Bitter, 1911b:998, fig. 4. [Type 

collection: T‘olcnn Mojanda, Ecuador, 30004000 m, Sodiro 

s.n. Holotype FI, not seen, possible isotype IT!] 

Polyiepis hypargyrea Bitter, 1911b:600, taf. 5. [Type collection: Sierra Se\.ada, Paramo de la Culata, T’enezuela, December, ilfor.itz 1120. Type destroyed, Rockefeller photograph iio. 17985, types of the Berlin Herbarium at GH.] 

Polylepis sttbintegrn Benoist, 1934:326. [Type collection: 

western slopes of Mt. Pichincha, Taurichupa, Pichincha, 

Ecuador, 4000 m, 28 Nov 1930, Benoist 3356. Lectotype P!] 

Polylepis qitinrliensis Cuatrecasas, 1941 :343. [Type collection: 

Central Cordillera, Sevado del Ruiz, western slope of 

macizo del Quindio, Caldas, Colombia, 3400-3500 m, 5 May 

1940, Cuatrecasas 9327. Holotype US! isotype us!] 

Deciduous trees (2) 5-15 m tall. Leaves slightly 

congested at the ends of the branches (Figure I&), 

imparipinnate with 2-6 pairs of leaflets obtrullate 

in outline, 2.5-5.9 cm wide, 2.4-11.9 cm long; 

rachises slightly tomentose, point of leaf attachment 

hispid or with a ring of multicellular glandular 

trichomes and resinous exudate, or both; stipular 

sheaths spurred or truncate at the apex, outer surface and top sericeous. Leaflets elliptic or ovate in 

outline, second pair from the terminal leaflet the 

largest, one of this pair from 0.5-1.1 cm wide and 

1.4-3.8 cm long; margins entire or slightly serrate; apex acute or obtuse; base unequally cordate; 

terminal leaflet petiolate; upper surface glabrous, 

lightly sericeous, or hispidulous; lower surface with 

a dense layer of gray, white, or gold silky trichomes 

(Figure 16b, c), but occasionally almost glabrous. 

Inflorescences pendant, 3-1 8 cm long, bearing 8-35 

flowers; floral bracts conspicuous, green or orange, 

scarious; rachises tomentose. Flowers perfect, 0.5-1 

cm in diameter; sepals 3-5, oval, green, outer surface 

sericeous (Figure 16d); stamens 6-16, purple or deep 

red brown, covered with long white trichomes; base 

of style hispid. Fruit irregularly shaped, more or 

less turbinate, pilose, with a wide variety of knobs 

and stout spines (Figure 16e), 0.2-0.7 cm wide including protuberances, 0.2-0.8 cm long. 

DIsTRr~vTIos.-Scattered in populations from 

Venezuela in the state of Lara along the Andes to 

Central Bolivia in the province of La Paz (Figure 

17). Flowering recorded every month except December. 

COMMON NAMEs.-coloradi to, keshua, quinua,

quinual, quinuar, quinhuar, queuna.

UsEs.--In Peru an infusion made from the bark 

was used to counteract colic. 

REPRESEXTATIVE SI~ECIMENS Ex.~~~I~ED.-T’ENEZUELA. LARA: 

TZ’estern slope of the Paramo del Jabon, 3100-3400 m, 2 h’ov 

19139, Citatrecasaa, Ruiz-Ter-an L* Lopez-Figueiras 28216 (US); 

Cordillera de Norte, Paramo tle Mucuruba, Quebrada de 10s 

Colorados, Cueva de 10s Colorados, 3900 m, 20 Oct 1969, 

Cuatrecases, Ruiz-Teran L- Lopez-Figueiras 28145 (US). MERIDA: 

Laguna Mucubaji, aboke Los Apartaderos, 3625-3655 m, 

21 Ju1 1944, Steyermark 57513 (F, SY); Paramos de Laguna 

Grande, 21 Jan 1929, Pittier 13253 (A, F, G, NO, SY, US); 

Laguna Segra, 20 >far 1951, Gines 2210 (US), Sierra Nevada 

tie Santo Domingo, N\V slope, Laguna Negra, 3600-3700 m, 

21 Sep 1952, Hitrnbert 26443 (P, S. US); 67 km northeast of 

M6rida along the road to Barinas, 3500 m, 19 Nov 1963, 
NUMBER 43 29 

FIGURE I6.-PoZyZepis sericea: a, branch (X 0.5); b, lower leaflet surface (x 3); c, point of 

leaflet attachment, lower surface (X 4); d, flower (x 6); e, fruit (x 6). 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 17.--Collection localities of Polylepis sericea. 
NUMBER 43 

Breteler 33ff (Nu); Sierra Nevada above Merida, 3900 m, 

2 Mar 1939, Alstoti 6803 (BM, US); Quebrada El Yoyo, 3880 m, 

12 Apr 1930, Gehriger 73 (A, F. MO, KY, UC, US); Sierra 

Nevada de Santo Domingo, Laguna de Los Pastos, 3800 m, 

17 Oct 1969, Cuatrecasas, Ruiz-Teran C- Lopez-Figueiras 28121 

(US); Paramo Piedras Blancas, 4000 m, 27 Mar 1945, Jahn 

fQ3 (US): “Merida,” 3900-4000 m, 13 Ju1 1953, Beruordi 608 

(NY); Merida, Nov 1958, Schreider s.n. (K). COLOMBIA. ANTIOQUIA: I’ulgo, Paramo de Kerveo, 4000 m, 1851-1857, 

Triaiia 1866 (G). CALDAS: Cordillera Central, head of the 

river Otun, paramos between Laguna del Mosquito and 

Plan del Vilar, 3650-3750 m, 26 NOT 1946, Cuatrecasas 23257 

(F, P, US), southeast of Ruiz, El Aprisco, 3500-3600 m, 5 May 

1940, Cuatt-ecasns 932i (US), Paramo del Quindio, 4200-1400 

m, 15-20 Aug 1922, Pennell i9 Hazen 9912 (GH, K, NY); 

high valleys near Solamina, 3000 m, Sep 1945, Tomas 2428 

(US). NARINO: Yacuanquer, north of Yacuanquer, 4 Jan 1943, 

Garganta 472 (F). ECUADOR. CARACHI: above Tufinio, 4000 

m, 23 Ju1 1935, Asplund 17037 (S). PICHINCHA: Chaparro de 

Sebritana, 3400-3500 m, 15 Ju1 1944, Solis 8379 (F): western 

slopes of I’olcari Cayambe, 3900 m, 27 Ju1 1967, Sparre I7789 

(S): Paramo de Guamani, slope toward Papallacta, 3900 rn, 

31 Oct 1955, Asplund 18244 (S); northwest slope of Mt. Corazon, 3300 m, 27 Ju1 19.55, Asplund I7103 (S), 3000 rn, 22 Jun 

1876, Andre 373.5 (K, NY); west of Pichincha, 12 Ju1 1931, 

Betioist 4549 (l’, S): western slope of Pichincha, 12,000 ft, 

Jatlleson 16 (G). NAPO: Paso de Guamani about 4 km east of 

pass on road to Papallacta, 3 Mar 1967, Sparre 15029 (S). 

CHIMBORAZO: Cordillera Oriental, sotobosque de Cubilliri, 

Hcda, Alas, 3350-3300 m, 1 Mar 1944, Solis 7524 (F). CANAR: 

Canar, 15 Sep 1918, Rose 6 Rose 23789 (NY, US). PERU. 

AXCASH: Carhuas, Cordillera Blanca, above Vicos, 3600 rn, 

11 Mar 1964, Hutchison L- Wright 4325 (F, MO, NY, UC, US). 

CUZCO: Urubamba \’alley, Torontoy, May 1915, Heller 2183 

(US): Lucumayo \‘alley, 2600 m, 18 Jan 1915, Cook & Gilbert 

I280 (US). BOLII‘A. LA PAZ: between Sorata and Mapiri, 

Fola, Aug 1933, Cardenas I202 (NY): Murillo, 8 krn after 

Palca on the road to Iquico, 4000 rn, Vuilleumier 342 (GH). 

DIscussIox.-Tlie geographical distribution of 

Polylcpis sericea spans the greatest latitude of any 

species in the genus, but populations (at least at the 

present time) are discontinuous along a north-south 

Andean axis. Correspondingly, the species consists 

of a series of relatively pronounced forms, many of 

which have been described as distinct taxa. However, examination of a large number of specimens 

from throughout the range shows that plants similar 

to a form characteristic of one area frequently occur 

in a population from another part of the range. 

The form upon which the name Polylepis sericea 

was based occurs predominantly in Venezuela and 

has elliptic leaflets that are glabrous above and 

densely covered beneath with silvery, silky trichomes. Similar plants are common in Ecuador 

(Pichincha). In Venezuela, however, there are also 

plants with almost entirely glabrous leaves. In fact, 

the amount, length, and color of the pubescence on 

the undersurface of the leaflets varies greatly 

throughout the range. In Colombia, the trichomes 

of the lower leaflet surfaces tend to be slightly 

shorter than those most commonly found in Venezuela, while in Ecuador plants from Coraoh, and 

in Peru plants from Caraz, have pronounced pubescence on the upper leaflet surfaces as well, similar 

to that of P. subsericans. 

‘There is also some variation in the shape of the 

top of the stipule sheaths. Most populations have 

spurs on either side of the leaf petiole but some 

specimens from Pichincha have the spurs so reduced 

as to make the top of the stipule sheath appear flat. 

Morphologically, Polylepis sericea is most similar 

to P. pauta and the two share most characters. The 

two differ in that the pubescence of the underside 

of the leaflets of P. sericea usually covers the entire 

surface arid thus appears as a solid layer of silky 

hairs. The trichomes of the underleaflet surfaces of 

P. pauta on the other hand are always much more 

conspicuous on, or restricted to, the veins. In addition, the leaflets of P. pauta are more ovate and 

larger than those of P. sericea and generally have 

serrate rather than entire margins. 

6. Polylepis subsericans Macbride 

Polylepis subsericaiis Macbride, 1934:367. [Type collection: 

Huamanga, Hacienda Totorabaniba, Ayacucho, Peru, 3500- 

3600 m, May 1910, Weberbauer 5487. Holotype F!, isotype F!] 

Polylepis incatia var. pavipila Bitter, 1911b:640. [Type collection: Castro-Virreyna, western slopes of the Andes above 

Pisco, Huancavelica, Peru, 39001000 m, May 1910, Weberbauer 5433. Type destroyed at Berlin, Isotypes F! GH! 

US!] 

Trees to 8 m tall. Leaves congested at the ends 

of the branches, imparipinnate with 1 pair of leaflets; trullate in outline, 1.7-3.3 cm wide, 3.1-3.4 

cm long; rachises pilose with scattered multicellular 

glandular trichomes; point of leaflet attachment 

(Figure 18b) with a small tuft of long, straight, 

single-celled trichomes; stipular sheaths sloping 

away from the petiole, appressed hispid on the 

upper portion and the outer face. Leaflets obovate 

in outline, 0.5-0.7 cm wide, 1.7-2.0 cm long, margins slightly revolute, more or less entire or slightly 

crenate toward the tip, apex slightly emarginate 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 18.-PoZylepis sztbsericans: a, branch tip (X 1.5); 

b, point oE leaflet attachment, upper surface (X 6). 

with the trichomes from the lower surface projecting into the notch, upper surface dull, sparsely 

pilose; lower surface (Figure 18a) densely hispid, 

the trichomes covering a lower layer of short, 

twisted, several-celled trichomes. Inflorescences 

pendant, 3-4 cm long, bearing about 3 or 4 flowers: 

floral bracts lanceolate, pilose on the outer surface; rachises pilose. Flowers perfect, about 0.8 cm 

in diameter, sepals 4, ovate, outer surface pilose 

with multicellular glandular trichomes extending 

to the tips on the inner face; stamens 12 or 13, 

purple, covered with long, straight white trichomes; 

base of style sparsely sericeous. Fruit more or less 

cylindrical in outline with irregularly flattened 

spines; mature fruit not seen. 

DlsTRIBvTIoh’.-Central Peru in the provinces of 

Ayacucho and Huancavelica (Figure 19). Flowering 

recorded only from the month of May. 

COMMON NAMES.-xOne specifically reported. 

Usm-None specifically reported. 

REPRESENTATIVE SPECIMESS ExAMrNED.-The Only 

collections seen are those listed as the type of the 

species and its synonym (Figure 19). 

DIscvssIoN.-Although originally described as a 

variety of Polylepis incana by Bitter, P. subseiicans 

appears to be more closely related to members of 

the sericea group than to the incana complex. Presumably Bitter placed the taxon within P. incana 

because of the single pair of leaflets and the yellowFIGURE 1 S.-Collectioq localities of PoZylepis subsericans. 

ish cast of the leaflets due to a resinous exudate 

from the glandular trichomes. However, the dense, 

straight trichomes covering the under surface of the 

leaflets are, as pointed out by Macbride, more 

similar to those of populations of P. sericea. As in 

the case of P. pepci, P. subsericans would seem to 

have arisen from an isolate of an ancestral stock 

similar to P. sericea. 

Because of the coarse, appressed covering of 

straight trichomes on the leaflets and the single 

pair of leaflets, this taxon is easily distinguished 

from other members of the genus. 

7. Polylepis pepei, new species 

TYPE CoLLEcTIos.-Chapare, Cochabamba, Bolivia, 77 km after Chapare on the road to Todos 

Santos, 4200 m, 4 Jan 1968, Vitilleztmier 465 (Holotype US! Isotypes GH! P!). 

Frutex nanus. Folia imparipinnata; foliola unijugata ad trijugata; foliolum obovatum ad oblongum, apice emarginatum vel tridentatum, base inequale attentuatum, margine integrum; supra breviter sericeum subtus breviter sericeum apprime 

secus margines et nervos. Racemi pauciflori. Flores 
NUMBER 43 33 

perfecti, staminibus (4) 10-12, apice barbatis. 

Fructus fusiformis leviter tortilis, nodosus vel spinuloses ad medium. 

ETYMoLoGY.-This species is named for Jose 

Cuatrecasas, called Don Pepe by his contemporaries, 

who has devoted his life to further an understanding of the flora and phytogeography of the tropical 

Andes. 

Shrubs to 1.5 m tall. Leaves very congested with 

the area of overlapping stipule sheaths swollen 

(Figure 20a), imparipinnate, with 1-3 pairs of leaflets, obtrullate in outline, 1.0-2.4 cm wide, 0.8-2.8 

cm long; rachises glabrescent or with variable 

amounts of short, straight, white, single-celled, appress-d trichomes; point of leaflet attachment (Figure 20c) with a ring of short, straight, single-celled 

trichomes; stipule sheaths with short, obtuse spurs 

on either side of the petiole, outer face glabrescent 

to strigose. Leaflets oblong to obovate, second pair 

from the top the largest if more than one pair is 

presmt, one of this pair 0.2-0.4 cm wide, 0.6-1.2 cm 

long, margins entire, ciliate, apex emarginate or 

tridentate due to a projection of the midvein, base 

unequally attenuate; upper surface sparsely to 

densely covered with short, appressed, straight 

single-celled trichomes; lower surfaces (Figure 20b) 

minutely sericeous, the trichomes often most conspicuous on the veins and margins. Inflorescences 

upright or pendant, to 4 cm long, bearing 14 

flowers; floral bracts lanceolate, to 3 mm long, 

outer surface strigose; rachises densely covered with 

similar trichomes. Flowers (Figure 204 perfect, 

sepals 3 or 4, lanceolate to ovate, outer surface 

covered with short, straight, appressed trichomes; 

stamens (4?) 10-12, anthers circular, tomentose at 

the apex; base of style sparsely hispid or glabrous. 

Fruit (Figure 20e) fusiform, often slightly twisted, 

with an irregular number of blunt protuberances or 

spines around the middle, 0.2-0.3 cm wide including 

protuberances, up to 0.7 cm long. 

DIsTRIBuTl0N.-Southeastern Peru in Cochabamba south along the Eastern Cordillera to Bolivia in the province of Cochabamba (Figure 21). 

Flowers recorded in July and January. 

COMMO~ NAMEs.-None specifically reported. 

Usm-None specifically reported. 

REPRLSE\TATIVE SPECIME~S EXAMINED -PERU. Cuzco: 

Pinasnoij, Panticalla Pass, 3600 m, 15 Ju1 1915, Cook Q GzZbert 1837 (US), Arma, 4350 m, July 1915, Bingham 2068. 

BOLIVIA. Cordillera Real, Alaska Mine, 13,700 ft, 14 Mar 

1926, Tate s.n. (NY). 

DIscussmN.-Although collections are sparse, 

Polylepis pepei appears to have a fairly large distributional range along the Eastern Cordillera from 

southern Peru to central Bolivia. My own collections (Vuilleumier 465) and those of Cook and 

Gilbert indicate that the species grows above the 

cloud forest on the east facing slopes. While Cook 

and Gilbert do not actually mention the habitat 

of the species, one of their specimens is covered 

with lichens and Spanish moss. 

On morphological grounds, Polylepis pepei is 

placed in the sericea species group. The type collection, with three pairs of leaflets and with trichomes predominantly on the veins and leaf margins of the underleaf surface, appears similar to 

small-leaved individuals of P. sericea from Volcan 

Pichincha, Ecuador. The collection from Peru has 

only one pair of leaflets, reduced inflorescences, and 

more dense covering of trichomes on the underleaflet surfaces. Despite these similarities to P. 

sericea, P. pepei constitutes one of the most distinctive species in the genus. Plants are true shrubs, 

not dwarfed trees, the fruit is elongate, twisted and 

knobbed around the center. The tendency for the 

leaflet apex to appear tridentate through projection 

of the midvein is also unique in the genus. 

8. Polylepis retkulata Hieronymus 

Polylepis reticdata Hieronymus, 1896: 312. [Type collection: 

near Las Calderas de Pasachoa and Ruminaqui, Pichincha, 

Ecuador, Stubel 20a. Type destroyed, Rockefeller photograph no. 3393, types of the Berlin Herbarium at GH, NY.] 

Polylepis nitida Bitter, 1911b3615, fig. 9, taf. 7. [Type collection: Minza at Volcan Tungurahua, Tungurahua, Ecuador, 

3600 m, Stubel287. Type destroyed, Rockefeller photograph 

no. 3388, types of the Berlin Herbarium at GH, NY.] 

Polylepis brachyphylla Bitter, 1911b:616, fig. 10 taf. 8. [Type 

collection: lower region of the paramos between Chimborazo and Guaranda, Chimborazo, Ecuador, 4 Nov 1856, 

Remy s.n. Holotype P! isotype P!] 

Trees 4-8 m tall. Leaves congested at the ends of 

the branches (Figure 22a), imparipinnate with 2-3 

pairs of leaflets, oblanceolate in outline, 1.8-3.5 cm 

wide, 2.0-5.0 cm long; rachises lanose; point of 

leaflet attachment (Figure 22c, d) with a small tuft 

of silky trichomes on the upper side or pilose with 

small multicellular, stacked glandular trichomes 

forming an annulus around the rachis; stipular 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 20.-Polylepis pepei: a, branch (X 0.5); b, leaf, lower surface (x 3); c, point of leaflet 

attachment, upper surface (X 12.5); d, flower (T%ith stamens missing) (X 4.5); e, fruit (X 4.5). 
NUMBER 43 35 

FIGURE 21.-Collection localities of Polylepis pepez. 

sheaths with short spurs, outer surface glabrescent 

to shortly lanose, with various amounts of long, 

straight trichomes projecting from the underside 

above the apex. Leaflets oblong or oblanceolate in 

outline, first or second pair from the terminal leaflet 

the largest, one of this pair 0.4-0.9 cm wide, 1.2-2.1 

cm long, margins revolute, sometimes crenate, the 

edges of the revolute leaf margins and the apices of 

the underside with a rim of rather long, straight, 

appressed trichomes, apex strongly emarginate, base 

unequally cordate, terminal leaflet petiolate; upper 

surfaces nitid, sometimes with a few scattered long 

white trichomes primarily in the depression of the 

midvein; lower surface pannose (Figure 224. Inflorescences pendant, 2.0-12 cm long, bearing 3-7 

flowers; floral bracts ovate, orange, usually scarious 

and glabrous but occasionally outer surfaces pilose. 

Flower (Figure 22f) perfect, 0.6-0.7 cm in diameter; 

sepals (3) 4, ovate, outer surface densely lanulose; 

stamens 6-1 7, globose in outline, the anthers tomentose, particularly at the junction of the anther sacs 

and along the edges of the anther sacs; base of style 

villous. Fruit (Figure 22e) globose, adorned with 

irregular, flattened spines, the spines with scattered 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

e 

f 

FIGURE 22.--Polylepis reticulala: a, branch (x 0.5); b, leaf, upper surface (x 1.5); c, point of 

leaflet attachment, upper surface (x 6); d, point of leaflet attachment, lo\ver surface (X 6); 

e, fruit (x 6); f, flower (x 6). 
NUMBER 43 

trichomes, the body of the fruit densely lanose, 

0.2-0.8 cm wide including spines, 0.3-0.9 cm long. 

DI~TRIBc'TI~N .-Central to southern Ecuador, primarily in the Eastern Cordillera (Figure 23). Flowers 

recorded in September, November, December, and 

May. 

COMMON NAMES.-Quinua, quinoar, yagual. 

UsEs.-None specifically reported. 

REPRESENTATIVE SPECIXIEKS ExAMINED.-ECUADOR. PICHINCHA: Quito, 1845, Jameson 204 (G, I$'); Imbabura, 4000- 

4300 m, 5 May 1957, Hirsch EZ13 (NY). TUNCURAHUA: Mount 

Caruhuayrayo, 4200 m, 22 Aug 1939, Asplund 8f6f (S); Aka 

de Passa, 3500 m, 28 Oct 1944, Solis 8736 (F); southwest slopes 

of Tungurahua, below paramo of hlinza, 3400 m, 14 .4pr 

1939, Pentlatid 2 Summers 420 (F. US). CHIMBORAZO: Chimborazo, 4000 m, 4 Ju1 1846, Andre 3933 (F, US); Eastern 

Cordillera of Riobamba, 3200 m, Rimbach s.n. (K); woods 

of Eastern Cordillera, Riobamba, 3400 m, December, Rimbach 

156 (GH, P, L.S); ibid 498 (S); .%zuay: along the Rio Matatiero west of Cuenca, 10,000-10,500 ft, 3 Mar 1945, Camp 

2000 (GH, MO, KY). IVITHOUT PRECISE LOCALITY: Eastern 

Cordillera, Kirnbach 26 (A, F); ibid, 3000-3400 m, May 1930, 

Rinibach I4 (A, F, GH, NY, US). 

DIscussIorv.-From accounts by collectors, Polylepis reticulata is the most common tree on the relatively humid slopes of the Ecuadorian Andes at the 

upper reaches of the montane forest. It appears to 

grow primarily in the cloud forest, but is also found 

isolated from true forest in valleys and along ridges. 

In general, the species exhibits a constant morphology throughout its range. The only variant populaFIGURE 23.-Collection localities of Polylepis reticulata. 

tions appear to be near Canar. Plants from this 

region resemble P. weberbazieri more than those 

farther east. 

The most conspicuous feature of this species 

(which it shares with P. weberbazieri and, to a lesser 

extent, with P. quadrijuga) is the shiny upper surface and emarginate apex of the leaflets. Polylepis 

reticulata and P. weberbaueri differ in the pubescence of the stipule sheaths and the underleaflet 

surfaces. The former has silky tricliomes on the 

outer face of the stipule sheaths and along the 

revolute margins of the leaflets, The latter has a 

dense wooly covering on both parts and thus no 

differentiated trichomes along the leaf margins. 

Although further collections may show these two 

taxa to be conspecific, I have kept them distinct because of the apparent consistent differences in morphology and the differences in habitat. It is possible 

that P. weberbaueri is derived from P. i.eticzilata 

and represents a taxon that evolved adaptations 

toward drier conditions following colonization from 

the more moist eastern slopes. 

9. Polylepis weberbaueri Pilger 

Polylepis weberbaueri Pilger, 19063535. [Type collection: 

from Yanganuco to Yungay, Ancash, Peru, 3700-3800 m, 

16 Jun 1903, Weberbauer .3287. Type destroyed, Rockefeller 

photograph no. 3399, types of the Berlin Herbarium at 

GH, NY.] 

PolyEepis Zanuginosa Humboldt, Bonpland k Kunth, p 

microphplla TVeddell, 1861 :238. [Type collection: Chirnborazo, Chimborazo, Ecuador, Bonpland 3Z4Z. Holotype P! 

isotypes A! F!] 

Polylepis ttiicrophylla (Weddell) Bitter, 1911b:611. [Based on 

P. lanuginosn p niicrophylla IVeddell.] 

Polplepis nzicrophplla var. polyarthrotricha Bitter, 1911h3612. 

[Locality not specified, Goudot 1. Holotype TY!] 

Shrubs or small trees to 1.5 in tall. Leaves very 

congested at the end of branches, sometimes producing an appearance of a whorled arrangement 

(Figure 24a) iniparipinnate with 3-5 pairs of leaflets, obtrullate in outline, 0.9-2.4 cin wide and 

1.5-5.0 cm long; rachises denszly lanose or lanose 

with dark red glandular trichomes; point of leaflet 

attachment (Figure 24e) with a tuft of matted multicellular glandular trichomes; stipule sheaths with 

short obtuse spurs, outer surface glabrous with long, 

straight silky trichomes protruding from the undersurface. Leaflets (Figure 24b) ovate to obovate in 

outline, first (rarely second) pair from the apex the 

NUMBER 43 39 

largest, one of this pair 0.4-0.9 cm wide, 0.4-1.6 cni 

long, margin revolute, apex strongly emarginate, 

base unequally cordate; upper leaf surface (Figure 

24c, d) nitid, rugose, or pilose, particularly in the 

depression of the midvein; lower surface pannose. 

Inflorescences pendant, 2.1-3.4 cm long, bearing 3-7 

flowers clustered at the terminal end; floral bracts 

small, 0.1-0.2 cm long, lanose or scarious; rachises 

lanose. Flowers (Figure 240 perfect, 0.4-0.6 cm in 

diameter; sepals four, ovate, sericeo-lanose on the 

outer surface; stamens 6-16, purple, locules barbate 

to glabrous, base of the style villous. Fruit globose 

in outline, irregularly spined, the spines flattened, 

body of the fruit and spints shortly villous; about 

0.29-0.57 cm wide including spines, 0.31-0.65 cm 

long. 

DIsTRInu?.IoN.-Central Ecuador in the Province 

of Pinchincha south to northern Peru in the Department of Ancash, primarily on dry slopes of the 

Andes (Figure 25). Flowering recorded in September, Novemb-r, and December. 

COMMON NAMES.-Quenuar, quinual.

UsEs.-Sone reported. 

REPRESENTATIVE SPECIIIENS ExAIIINED.-ECUADOR. PICHINCHA: Andes of Quito, 1855, Couthouy s.n. (GH). CHIMFIGURE 25.-Collection localities of Pol)lepis weberbaueri. 

BORAZO: Chimborazo, 12000 ft, Dec 1846, Jameson 523 (BM, 

G); eastern side of Chimborazo, 10 Nov 1845, Jameson 25 

(K); border with Canar (western escarpment) near El Tambo, 

10,000-11,500 ft, 6-9 Jul 1845, Camp 4088 (F, NY). CANA R: 

Canar, 16 Sep 1918, Rose, Pachano & Rose 23792 (GH, NY), 

vicinity of Canar, 15 Sep 1918, Rose 22722 (NY, US), ibid, 

16 Sep 1918, Rose C- Rose 22733 (SY); “Ecuador,” Dec 1890, 

Sodiro 421 (G, Herb DC). PERU. PIURA: Huancabamba, on 

the road to Huancabamba just below the summit, 38 km 

above Canchaque, 3120 m, 13 Sep 1964, Hutchison & Wright 

6578 (F, GH, MO, NY, P, UC). CAJAMARCA: Huambos, 2000 m, 

11 Sep 1956, Sozikup 4457 #(F, US). ASCASH: Cordillera Blanca, 

Quebrada Tangamico, 30 Ju1 1959, without collector PI970 

(hY. 

DIscussroN.-In maay respects, Polylepis weberbaueri appears to be the ecological equivalent in 

the drier Ecuadorian Cordillera of P. reticulata. It 

is probable that P. weberbaueri is derived from 

P. reticulata following an establishment of a population on the western chain. In areas of apparent 

sympatry near Canar, Ecuador, specimens intermediate in morphology could be interpreted as 

hybrids between the two taxa. Although this hybridization might suggest that P. weberbaueri is 

only a geographic variant of P. reticulala, consistent 

character complexes over extensive geographical 

areas indicate that the two should be considered as 

independent, but closely related, species. The two 

can be distinguished most easily on the basis of the 

vestiture of the underleaflet surfaces and the stipule 

sheaths. The lower leaflet surface of P. reticulata 

is lanose over most of the surface, but distinctly 

sericeous along the slightly revolute leaflet margins. 

The entire underleaflet surface of P. weberbaueri 

is lanose and the leaf margins glabrous (Figure 24d). 

In addition, the exterior surface of the stipule 

sheaths are sericeous in P. reticulata and lanose in 

P. weberbaueri. 

Included within this taxon are populations originally described as a variety of Polylepis lanztginosa 

(P. lanuginosa p microphylla Weddell) and subsequently as P. microphylla by Bitter (Figure 24a). In 

our collections in southern Ecuador (Simpson 8532) 

on the slopes of mountains near Canar, we found 

plants that have mature, lower leaves characteristic 

of P. weberaueri and young, upper leaves like those 

of the specimens placed in P. microphylla (Figure 

24c). In both “P. rnici-ophylla” and the branch tips 

of our specimens from Canar, the “paired” leaflets 

are often somewhat skewed so as to appear alternately arranged on the rachis. All of the specimens 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

that have been previously referred to P. miciophy lla 

are sterile, precluding the use of flowers or fruits as 

a taxonomic guide. Until further evidence indicates 

otherwise, the small-leaved form would seem most 

logically placed in synonomy with P. weberbaueri. 

In addition to the typical variety, Bitter circumscribed a second variety, P. rnicrophylla var. polyarthrotricha, which he considered to merit separation on the basis of its possession of dark red multicellular glandular trichomes on the leaf raches and 

underleaf surfaces. He recognized (and listed) intermediate forms between the purely lanose type and 

the lanose-glandular type. Since a gradation of 

glandular trichome density exists within the populations from Chimborazo, formal recognition of a 

separate variety seems unnecessarj. In all of the 

specimens of the species examined, at least a small 

cluster of glandular trichomes exists at the point of 

leaflet attachment. 

10. Polylepis quadrijuga Bitter 

Polylepis qztadrijziga Bitter, 1911b:613, taf. 6. [Type collection: Cordillera Oriental, Paramos de Bogota, Muzo, Cundinamarca, Colombia, May 1844, Goiidot s.n. Holotype PI] 

Polylefiis boyarenais Cuatrecasas, 1941:343, figs. 13, 14. 

[Type collection: Cordillera Oriental, Paramo de Santa 

Rcsita between Belen and Susacon, Boyaca, Colombia, 3300- 

3400 m, 3 Aug 19.*0, J. Cuatrecases 10371. Holotype PI, 

isotype US!] 

Polylepis rocuyensis Killip 8: Cuatrecases-Cuatrecasas, 

1942:33, fig. 19. [Type collection: Nebado del Cocuy in the 

Quebrada of San Paulino, El Morroti, Boyaca, Colombia, 

3800 m, 11 Sep 1938, J. Cuatrecasas 1105. Holotype US! 

isotype P!] 

Tree 5-8 m tall. Leaves (Figure 26a) only slightly 

congested at the ends of the branches, imparipinnate with 3-5 pairs of leaflets, obtrullate in outline, 

2.6-4.5 cm wide and 3.8-9.4 cin long; rachises 

densely lanose; point of leaflet attachment (Figure 

26c) with a distinct zone of glandular trichomes and 

resinous exudate; stipulate sheaths with acute spurs, 

outer surface lanose, with long, straight trichomes 

emerging from the inside surface at the apex. Leaflets (Figure 26b) ovate to obovate in outline, second 

or third pair from the apex the largest, one of this 

pair 0.6-1.3 cm wide and 1.3-2.4 cm long; margins 

entire or slightly crenate; apex emarginate with 

long straight trichomes visible in the indentation; 

base unequally cordate; upper leaflet surface rugose 

or nitid, sometimes with scattered long trichomes; 

lower leaflet surface densely pannose or lanulose. 

Inflorescences pendant, 5.8-14 cm long, bearing 8-18 

flowers; floral bracts lanceolate, exterior surface 

lanose rachises sericeous. Flowers (Figure 26d) perfect, 0.6-0.7 cm in diameter, sepals 4 or 5, oval in 

outline, outer surface lanose; stamens 8-20, globose, 

covered with long, curled trichomes, especially at 

the apex; base of style villous. Fruit (Figure 26f) 

globose, covered with irregularly positioned long, 

thin spines, the spines glabrous or lanose, body of 

the fruit lanose, 0.3-0.7 cm wide including spines 

and '0.2-0.i cm long. 

DIsTRIBuTIox.-Colombia in the provinces of 

Santander, Boyaca, and Cundinamarca (Figure 27). 

Flowers recorded in May, August, and September. 

Coxmros r\'AhrEs.-None recorded. 

Usm-None recorded. 

REPRESE~TATIVE SPCIMENS ExAMIsED.-COLOMBIA. SANTANDER: Eastern Cordillera, Paramo de las Coloradas, aboi e 

La Baja, 3900-4100 m, 27 Jan 1927, Kirlip L? Smith I8170 

(A, GH, YY, US): Paramo de la Rusia, 3300-3500 m, 4 Aug 

1940, Cuatrecasns 19428 (F, US). BOYAAA: Nevado del Cocuy, 

I'alle tle la Cueta, 3700 m, 10 Sep 1938, Cuatrecases 1294 

(CS); I'alle de Cocuy, southwest slopes, 3100@3750 m, 8 Sep 

1938, Cuntrecasas 1259 (F, US); Chorreon de San Paulino, 

37.50 m, 10 Sep 1938, Cuatiecasas 1355 (F, US); valley of Rio 

del Corallites, 4000 ni, 9 Jun 1957, Gmbb. Curry, FernandezPerez 755 (K, US); I'alle tle las Lagunillas, 4000 m, 13 Sep 

1957, Grubb, Ciirrg, Feriiandez-Perez 839 (L'S); Paramo de 

la Rusia near La Opera along the road between Duitama and 

Charata. 3643 m, 20 Aiig 1953, Longeriheim 3518 (C-C, US); 

Avenitla tle Penas Segras (Buenos .4ires), 3350 m. 11 Sep 1969, 

Ciicitreca.tes Roclrigrter 27711 (US); Guican, Hoya del Rio 

Tabor, in the direction of Ritacuva, 3530-3370 m, 18 Sep 

1969, Rod? iguez 2 Cuatrecasas 27834 (us). CUNDINAVARCA: 

Macizo de Bogota, eastern slopes of Paramo de Chisaca, 

Quel)rada de Santa Rosa, 3300-3350 m, 16 Sep 1961, Cuatrecmas L* Jnrainilia 25999 '(US). 

Discr:ssIo~.-Although isolated far north of 

similar members of the Polylepis sel-icea species 

group, P. quadyijzign exhibits many characters that 

indicate its close affinity with P. 1anzcgino.m These 

include the densely sericeo-lanose stipule sheaths, 

the large, multiflowered racemes and the presence 

of multicellular glandular trichomes with a large 

terminal cell at the point of leaflet intersection. In 

herbaria the two taxa have often been confused 

with each other, but they can be distinguished on 

the basis of several characters. The pubescence of 

the undersurface of the leaflets is sericeous in P. 

Zanziginosn and lanose in P. qzradrijuga. The latter 
NUMBER 43 41 

FIGURE 26.--PolyZepis qzradrijuga: a, branch (x 0.5): b, undersurface of leaflets (x 3): r, point 

of leaflet attachment, upper surface (x 6); d, e, flowers (x 6); f, fruit (x 6). 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 27.-Collection localities of Polylepis quadrijuga. 

lacks any long, single-celled trichomes at the point 

of leaflet intersection, whereas the former has these 

mixed with the glandular trichomes. Presumably 

P. quadrijuga ha5 differentiated from a northern 

isolated population of P. lanuginosa or resulted 

from establishment of a population from a southern 

propagule with subsequent differentiation. 

The habitat of this species is humid, often boggy 

paramos or along streams, where trees can form 

dense stands at elevations as high as 4000 m. 

11. Polylepis besseri Hieronymus 

Polylepis besseri Hieronymus, 1896:312 [Type collection: 

Capi, Bolivia, Mar 1890, Bang 769. Lectotype G! isotypes 

GH! MO! NY! US! W!] 

Polylepis raceniosa var. tomentosa 0. Kuntze, 1898:77 [Type 

collection: between Challa and Tapacari, Cochahamba, 

Bolivia, 3600-4600 m, 18 Mar 1892, Lorentz 0 Hieronymus 

s.n. Type destroyed at Berlin, isotypes NY! UC! US!] 

Polylepis raceniosa var. lanata 0. Kuntze, 1898:77. [Type 

collection: Tunari, Bolivia, 3WO-4000 m, Kuntze s.n. Type 

destroyed at Berlin, isotype NY!] 

Polylepis besseri var. abbreviata Bitter, 1911b:628. [Illegitimate name since it was based on P. ruce?nosa var, tornentosa 

0. Kuntze.] 

Polylepis besseri var. longipedicellata Bitter, 1911h:629. [Type 

collection: Puna of Mizque, Cochabamba, Bolivia, d’Orbigny 

s.n. Holotype G!, isotypes NY! US!] 

Polylepis triacontandra Bitter, 191 lh:630. [Type collection: 

Larecaja near Sorata, Cochipata, La Paz, Bolivia, 3300 m, 

9 Oct 1818, Mandon 674. Lectotype G!, isotypes NY! P! 

Polylepis crista-galli Bitter, 191 1b:633. [Type collection: Tucurnilla, Tarija, Bolivia, 2500 m, Fiebrig 2020. Lectotype 

G!, isotypes A! P!] 

Polylepis crista-galli var. Eongiracemosa Bitter 1911b3634. 

[Type collection: Pinos, between Tarija and San Luis, 

Tarija, Bolivia, 2500-2700 m, 1 Mar 1903. Fries 2296. 

Holotype S!] 

Polylepis tenuiruga Bitter, 191 1h:635. [Type collection: 

“Chile” without precise locality, Besser s.n. Type destroyed 

at Berlin, Rockefeller photograph no. 3397, types of the 

Berlin Herbarium at GH, NY.] 

Polylepis subquinquefolia Bitter, 1911b:636. [Type collection: 

Sandia above Coyoenyo, Puno, Peru, 3600 m, Weberbauer 

931. Type destroyed at Berlin, Rockefeller photograph 

no. 3391, types of the Berlin Herbarium at GH, NY.] 

Polylepis rzcgulosa Bitter, 1911h3639. [Type collection: Pampa 

behind train station, Arequipa to Puno line, Arequipa, 

Peru, 3800 m, Weberbauer 4882. Type destroyed at Berlin, 

isotype WRAT.] 

Polylepis incana ssp. subtusalbida Bitter, 1911b:640. [Type 

collection: Cuesta de Duraznillos. Bolivia, 2400-2600 m, 

Dec 1907, Hertzog 712. Holotype WRSL.] 

Polylepis inccrna ssp. incarum Bitter, 1911h:643. [Type collection: Lago Titicaca, Isla del Sol, near Challa, Bolivia, Seller 

118. Holotype B!] 

Polylepis incana ssp. brachypoda Bitter, 191 1b:644. [Type 

collection: Lago Titicaca, Isla del Sol, near Challa, Bolivia, 

Seller 148~. Type not seen, possibly destroyed at Berlin.] 

Polylepis pallidistigma Bitter, 1911h:645. [Type collection: 

Azangaro, Munani, Puno, Peru, 3650 m, Weberbauer 1369. 

Type destroyed at Berlin, isotype WRAT.] 

S! US!] 

Trees 1-8 m tall. Leaves congested at the ends 

of the branches, imparipinnate with 1-3 pairs of 

leaflets (Figures 28a, 29a); rhombic or obtrullate in 

outline, 1.4-4.9 cm wide, 1.8-7.9 cm long, rachises 

densely lanose mixed with glandular trichomes; 

point of leaflet attachment (Figures 28b, 29b) with 

a small tuft of yellowish or white trichomes; stipule 

sheaths with a slight shoulder or sloping away from 

the petiole, densely pubescent at the base. Leaflets 

broadly obovate to almost orbicular, first pair from 

the apex the largest, one of this pair 0.5-1.5 cm 

wide, 0.9-3.4 cm long; margins crenate or more or 

less entire, often revolute, apex obtuse or emarginate, base abrupt, unequal; upper surface dull, often 

rugose, sometimes with scattered twisted trichomes; 

lower surface (Figure 28c, 29b) pannose or lanose, 

the vestiture white, often with glandular trichomes 

very dense along the veins and leaflet margins, occa- 
NUMBER 43 43 

FIGURE 28.--Polykpis besseri from Colomi, Bolivia: a, branch (X 0.5): b, point of leaflet attachment, upper surface (x 3); c, point of leaflet attachment, lower surface (X 3); d, flower (X 3); 

e, fruit (X 3). 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

sionally glabrescent. Inflorescence pendant (or more 

or less upright if very short), 3.0-1 1.0 cm long, bearing 3-12 flowers; floral bracts lanceolate, glandular 

on the outer surface; rachises glabrescent with scattered glandular trichomes or lightly lanose. Flowers 

(Figure 28d, 29c) perfect, 0.9-1.0 cm in diameter; sepals 3-4, ovate, shortly lanose on the outer surface and 

on the tips of the inner surface, outer surface often 

with variable amounts of glandular trichomes; 

stamens up to 24 (12-24) per flower, anthers orbicular, tomentose at the apex; base of style white 

tomentose. Fruit (Figure 28e, 29d) very turbinate, 

with variable numbers and placement of flattened 

spines often coalesced into a spiny wing, body of 

fruit and spines densely lanose; 0.2-0.9 cm wide 

including protuberances, 0.4-1.3 cm long. 

DIsTRIUvTIoN.-Southern Peru from X!Ioqueque 

and Cuzco south to Arequipa and Puno (the Besser 

specimen label “Chile” was probably collected in 

Tacna, Peru) and in Bolivia in the provinces of 

La Paz, Cochabamba, Oruro, Chuiquisaca, and 

Tarija (Figure 30). Flowers recorded in January to 

June and October and November. 

COMMON NAME.-Quenua. 

USES.-None specifically reported. 

REPRESENTATIVE SPECIVESS ExAMINED.-PERU. CUZCO: 3km 

east of Cuzco, valleys and open slopes, 3600 m, 1 Oct 1936, 

West 8053 (GH, 110, UC); Cuzco, 3540 m, 19 May 1958, 

Zrifantes 5649 (P); Kavia, Infantex 6225 (P); near Cuzco, 3300- 

3800 m, 19-21 May 1958, Humbert 30526 (P): without localit\-. IM-1940, Gay s.n. (GH). MOQUECUA: Cordillera above 

Torata, 39004000 m, 14-15 Feh 1929, Weberbauer i470 (BM, 

F, G, S, US). TACSA: 4 km north of Torata, 3840 m, 27 Jan 

1952, Pearson 33 (F, UC). Puxo: Yolcan Chachani, ca. 4000 m, 

16 Jan 1977, Sirripson 8570 (US); ihid, Rauh P556 (XY); ihid, 

4000-4200 m, 14 Apr 1925, Pennell 13295 (F, GH, US); 

Sihuata, La Cumhre hetrveen Arquipa and Puno, 2600-4000 

m, 13 Nov 1947, Ferreyra 259f (GH, L‘C, US); hetween Lake 

Salina and Arequipa, 13,000 ft, 7 Feh 1943, Sandernan 3816 

(F, K); Pro\. Carahaya, between Macusani and Sunoa, 4000 m, 

29 Feb 1948, Vargas 7139 (MO); Puno, 2 Jan 1920, Shepard 

150 (GH, SY). BOLI1’1.4. LA P.iz: Dept. La Paz, 3840 rn, 

Mar 1910, Buchtien 4243 (GH, SY): Titicaca, d’Orbigny s.n. 

(P). COCHABAMBA: Chapare, Quehrada de Colomi, 11,500 ft, 

E. k’. Bails B62ff (BM, K, UC, US); Colomi, 12,200 ft, Brooke 

5075 (F, SY); Ahra de Colomi, 3450 m, 20 Jan 1929, Steinborli 9869 (F, GH, K, JIO, s); ibid, Feb 1947, Cardenas 3906 

(S, ITS); ihid, Jan 1949, Cardenas f333 (US); Prov. Cochahamha, Liriuni, 24 Feh 1950, K?-aprovickas & Fz~cl?s 6690 

JLIL): 100 mi northwest of Cochahamha across the Tunari 

Range, Choro, 11,000 ft. 18 Jan 1950, Brooke 5984 (F, SU); 

Pro\-. Qnillacollo, 4 km after Cochahamba on the road to 

Morochata. 5 Jan 1968, Vuilleurnier 465 (GH, P, US); 81 miles 

out of Cochahamba on the road to Santa Cruz, 13 Nov 1959, 

,\faguire & dlaguire 44f82 (GH, K, SY, US). CHUQUISACA: 

Guerraloma, 2800 m, May 1958, Carderias 572f (US.) 

DIscussIoN.-1.Vithout doubt, Polylepis besseri is 

comprised of the most polymorphic and confusing 

set of populations of any of the species of the genus. 

The same confusion I experienced in the treatment 

of these populations is evident in the treatments of 

earlier workers. Several specimens described as 

varieties of Polylepis wcemosa by Otto Kuntze 

(1898) are included here and were transferred to 

components of this species by Bitter (1911b). In 

the original description of the species by Hieronyinus, several collections were cited from various 

parts of Bolivia. Bitter specifically stated that Hieronymus’ species was an unnatural one containing 

more than one species and he separated part of the 

collections out at P. tenuii-uga. Another specimen 

annotated by Hieronymus as belonging to P. besseri 

was made the type of P. crisia-galli by Bitter. In addition, Bitter described several other species on the 

basis of one or two collections. Even within the collections designated as the types of his various species, 

however, there is tremendous variation. The specimen labeled “Type” of P. triacontandra has leaflets over 3 cm long, whereas other specimens from 

the same collection have leaflets about 1 cm long. 

A large series of collections hp various botanists 

from Colomi, Bolivia, has several herbarium names 

indicating that the collectors considered describing other new species. 

While final resolution of this series of populations 

must wait until more collections are made in the 

Eastern Bolivian mountains, the treatment proposed 

here is the only logical one possible lvith the material now present in herbaria. The assignment of 

the various populations to a series of species as 

proposed by Bitter makes no sense in the light of the 

number of intermediate populations and variation 

within populations. 1Vithin a population, the greatest amount of variation is in the size of the leaflets 

and the density of the trichomes on the underleaflet surfaces. Between populations, the variation is 

in amount of creriation or revolution of the leaflet 

margins, the amount of rugosity of the upper leaflet surface, the extent of the glandular trichomes 

on the veins of the underleaflet surfaces and the 

amount of coalescence of the spines of the fruits to 

form spiny wings. All of these characters can vary 

to some extent, however, within a population. 

Morphologically, Polylepis besseyi most closely 
NUMBER 43 45 

FIGURE PQ.-Polylepis besseri from Chachani, Peru: a, branch (x 0.5); b, point of leaflet attachment with one leaf turned to show lower surface (X 6); c, flower (X 6); d, fruit (X 6). 
46 SiMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 30.-Collection localities of Polylepis besseri. 

approaches P. tomentella, but the latter never has 

the dense pure white covering of the underleaflet 

surfaces that is so common in P. besseri. In addition, the fruits of P. tomentella are ridged or irregularly winged, whereas those of P. besseri have flattened spines projecting upward at the apex of the 

fruit or coalesced into spined wings irregularly 

placed on the fruit. The stipule sheaths of P. besseri 

are almost always pubescent and those of P. tomentella glabrescent or glandular. 

12. Polylepis tomentella Weddell 

Polylepis tomentella Weddell, 1861:237, taf. 78. [Type collection: Chuquisaca, Cinti, Bolivia, Jan 1846, Weddell 3927. 

Lectotype P! isotype P!] 

Polylepis tavapacana Philippi, 1891:Zl. [Type collection: 

near Cana, Tarapaca, Chile, 3900 m, Philippi S.?L Lectotype SGO, isotypes GH! SGO.] 

Polylepis toment ,lla var. pilosior Bitter, 1911b:647. [Type 

collection: Chuquisaca, Cinti, Bolivia, Jan 1846, Weddell 

3947 (in part). Holotype P!] 

Polylepis tomentella ssp. pentaphylla Bitter, 1911b:648. [Type 

collection: Laguna Tres Cruces, Jujuy, Argentina, Dec 1900 

to Feb 1901, Clarin 11688. Holotype S!] 

Polyl pis tometitella ssp. tetragona Bitter, 1911b:649. [Type 

collection: Salinas Grande near Jujuy, Jujuy, Argentina, 

3500 m, Hauthal 141. Type not seen, apparently destroyed 

at Berlin.] 

Polylepis tomentella ssp. dentatialatu Bitter, 191 lb:650. [Type 

collection: in a valley between Chorolque and Tacna, 

Potosi, Bolivia, 3600-3800 m, Hauthal 117. Type not seen, 

apparently destroyed at Berlin.] 

PolykPis tarapacana var. multisquama Bitter, 1911b3654. 

[Type collection: Tarapaca, Chile, Philippi s.n. B, apparently destroyed. Probably an illegitimate name since it 

was based on a Philippi specimen that was most likely 

part of the type collection of P. tarapacana.] 

Polylepis tarapacana var. sajamensis Bitter, 191 1b:654. [Type 

collection: Sajama, Oruro, Bolivia, 4500 m, Stubel 1. Type 

not seen, probably destroyed at Berlin. Cited by Bitter as a 

unicate.] 

Polylepis tarapacana var. bievifilamentosa Bitter, 191 1b:654. 

[Type collection: near the Bolivian province of Oruro, 

Tacna, Peru, Stubel 112. Type not seen, apparently destroyed at Berlin.] 

Polylepis tavpacana var. pycnolopha Bitter, 1911b:654. [Type 

collection: between La Paz and Tacna, Bolivia, 12,300- 

13,400 ft, 1838, Pentland s.n. Holotype P!] 

Small trees 1-4 m tall. Leaves congested at the 

ends of the branches or continuously borne all along 

the branches, composed of 1 (occasionally 2) pair 

of leafl-ts (Figures 31a, 32a), trullate in outline, 

0.8-3.9 cm wide, 1.9-2.4 cm long, rachises glabrous 

or covered on the abaxial side with glandular trichomes and long, white, single-celled trichomes, 

point of leaflet attachment sericeous; stipule sheaths 

with a slight shoulder at the apex or sloping downward, glabrous at the bottom of the outer surface, 

the top of the outer surface often with glandular 

trichomes, and with long trichomes projecting from 

the inside upper sheath surface (Figures 31b, 32b). 

Leaflets ovate to obovate, 0.3-0.6 cm wide and 

0.7-2.0 cm long, variable in thickness, margins 

entire to crenate, apex obtuse, base unequal; upper 

surface glabrous, dull green or yellowish green (Figures 31c, 32c); lower surface variable, densely pilose, 

often with glandular multicellular trichomes giving the surface a matted, yellow appearance or glabrescent with scattered glandular protuberances. Inflorescences upright or pendant; if upright, bearing 

1 or 2 flowers, if pendant, with 1-8 flowers, 0.1-5.1 

cm long; floral bracts hidden or lanceolate, denticulate, about 2 mm long, sparsely covered with 

glandular trichomes; rachises glabrescent with appressed, twisted trichomes mixed with glandular 

protuberances and resin. Flowers (Figures 31d, 32d) 

perfect, 0.4-0.6 cm in diameter, sepals 3, ovate, 

sometimes adnate at the base, green, outer surface 

with twisted white trichomes and glandular pro- 

48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

d e 

FIGURE 32.-PoZyZepis tomentella from Cochabamba, Bolivia: a, branch (x 0.5); b, cluster of 

leaves (x 3); c, upper leaflet surface (x 12.5); d, flower (x 6); e, fruit (x 6). 
NUMBER 43 49 

tuberances, stamens 6-14, ovoid, red, tomentose at 

the apex; base of style hispid. Fruit turbinate, with 

3-4 ridges, the ridges sometimes expanded into wings 

with entire or toothed margins or merely with a few 

knobs, body of the fruit with twisted trichomes 

(Figures 31e, 32e), 0.2-0.78 cm wide, including 

ridges, 0.3-0.9 cm long. 

DISTRIBUTION.-SOUthern Peru from 1 acna to 

Puno southward to northern Chile in the province 

of Tarapaca and across western and central Bolivia 

to northern Argentina in the province of Jujuy 

(Figure 33). Flowering recorded in July, October, 

and December to March. 

COMMON NAME.--Quenoa.

UsEs.--None specifically described, but personally 

observed to be used as firewood. 

REPRESENTATIVE SPECIMESS EXAMIKED.-PERU, PUNO: Santa 

Rosa, 50 miles south-southwest of Llave, 4600 m, 25 Jul 1946, 

0. PearSon 16 (US). TACNA: Cordillera de Tacora, Chulunquaiani, Weddell s.n. (P); Tacora, Chislluma, 4500 m, Apr 

1926, Werdernzanii 1143 (A, BM, F, G, GH, K, MO, NY, 

UC, US). BOLII‘IA. ORURO: Province Carangas around the 

base of I’olcan Sajama, 4350 m, 18 Oct 1967, Vuillrurnier 316 

FIGURE 33.-Collection localities of Polylepis tomentella. 

(GH, US); Challapata, 4000 m, 1 Apr 1921, Asplzrnd 6169 

and 6170 (US). POTOSI: Province Frias, 39 km northwest of 

Potosi on the road to Challapata, 3680 m, 14 Dec 1967, 

Vuilleurniel- 450 (GH, US); Province Chichas, near Quechisla 

on the road to Chonolque, 3600 m, Dec 1931, Cardenas 24 

(GH); Province Sud Chichas, Mina Isca Isca above the village 

of La Torre north of Tupiza, 3875-3880 m, 2 Dec 1967, 

Vuilleumier 402 (GH, US); near Santa Barbara on the road 

from Quechisla, 4200 m, 22 Feb 1936, West 6110 (GH, MO, 

UC). COCHABANBA: Mizque, D’Orbigny 495 ‘(P); between 

Monte Punco and Siberia, 3500 m, 28 Jan 1958, Meyer 20335 

(LIL). CHUQUISACA: Province Sor Cinti, along the road 

between Camargo and San Lucas, 3300 m, 10 Dec 1967, 

T’uilleuiniel- 427 (GH, US); Quebrada Honda, 3100 m, 28 hfar 

1934, Harniriarlund 395 (S). ARGESTINA. JUJUY: Humahucaca, Mina Aquilar, 4500 m, 16 Mar 1953, Sleurner 

3375 (S). 

DIscussIoN.-Included here within Polylepis tomentella are essentially all of the forms originally included in the taxon by Weddell. Bitter considered 

Weddell’s taxon to be an artificial assembly of at 

least two distinct species, and he consequently recognized both P. tomentella and P. tarapacana, a 

species described by Philippi. In his treatment, Bitter effectively typified P. tomentella by excluding 

all of the specimens cited by M’eddell except Weddell’s own collection from Cinti, Bolivia. Consequently, the specimens cited by T7eddell from 

Tacna fall within the circumscription of P. tampacana if it is recognized. In addition to the treatment of P. tornentella as two species, Bitter also 

erected numerous subspecific taxa within both 

species. Despite his fragmentation of the entire complex, however, he stated that an examination of 

more specimens might eventually show P. tomentella and P. tarapacana to be conspecific. My examination of a variety of specimens, including numerous 

personal collections from western Bolivia, indicates 

that this is indeed the case. While it is true that 

populations from Tarapaca, Chile, and adjacent 

Bolivia look very different from those in Cochabamba, Bolivia, there is a gradual transition from 

west to east across the altiplano with no populations 

that appear as “hybrids.” If it were not for the 

clinal change exhibited by the populations across 

Bolivia, the eastern and western forms could be 

considered separate taxa. In the west, plants have 

reduced thick leaflets with almost entire margins 

and densely pubescent underleaf surfaces (with a 

large number of glandular trichomes mixed with 

tightly twisted, short tricliomes). Eastward in Bo- 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

livia toward Oruro and southward toward Jujuy, 

Argentina, the leaves become larger, more crenate, 

and thinner. Especially in central Bolivia, the pubescence of the underside of the leaves becomes less 

dense. In Cochabamba, the leaflets are quite thin, 

almost glabrous, and very crenate. In addition, the 

inflorescences are longer and bear more flowers in 

the eastern populations than in those in the west. 

All of these morphological changes from west to 

east appear to be related to the changes in environmental conditions. Populations in the west are at 

4000-5200 in elevations in extremely arid areas. 

Eastward across the altiplano, the elevation is lower 

and the rainfall higher. Morphological characteristics of the westermost populations are common 

features of desert trees, which become less pronounced as aridity decreases. 

Not only does it seem to be most meaningful to 

consider P. tomentella and P. tarapacana as one 

taxon, but it alscj seem unnecessary to retain the 

subspxific taxa Bitter erected. In general, the characters that separate these subspecies and varieties 

are the number of pairs of leaflets, the amount of 

dentation of the leaflet margins, and number of 

fruit ridges. All of these characters are variable 

within populations of many species of the genus 

and not even consistent within the collections designated as types of these subspecific entities by Bitter. 

A more difficult problem with Polylepis tomentella lies in an assessment of its affinity with P. 

besseri and P. incana, particularly in central and 

southern Bolivia. In eastern BoliLia, populations of 

P. toinentclla are very similar to those of P. incana. 

Both species in this region normally have three 

leaflets, crenate leaflet margins, glabrous or sparsely 

pubescent under leaflet surfaces, and few-flowered 

racemes. The fruits of both usually have irregular 

wings. It is possible that the two taxa approach each 

other morphologically because of the similarity of 

the habitats in which they occur. Since this is the 

only area where the two species come into contact, 

it is also possible that they are forms of one species 

that replace each other geographically. The morphological characters that can best separate the two as 

treated here are the pubescence of the underleaflet 

surfaces and the shape of tlie leaflet. In P. incana, 

the leaflets are long, rather narrow, and distinctly 

oblanceolate, where in P. tornentella they are 

broader and often more ovate. The underleaf surfaces of P. incana have a dusky yellow cast, whereas 

those of populations of P. tomentella in areas of 

sympatry (Cochabamba, Bolivia) do not. Polylepis 

tomentella in western Bolivia does have yellowish 

underleaflet surfaces, but in these populations the 

leaflets are very thick and have a dense covering of 

twisted white trichomes mixed with the yellow 

glandular trichomes. These two characters represent 

the most easily described of a set of features that 

gives the foliage of P. incana a very different aspect 

from that of P. tomentella. The two are, however, 

sometimes difficult to distinguish from each other. 

In eastern Peru, there are also some specimens 

that indicate that I’olylepis tomentella might hybridize with P. besseri (e.g., iMeyer 20335). Such 

specimens have crenate, nonemarginate leaflets like 

P. tomentella, but the characteristic densely wooly 

pubescence of P. besseri. The two are usually separated ecologically with P. besseui, inhabiting the 

moist eastern slopes and P. tomentella tlie interior 

drier regions. In the valley around Cochabamba, the 

two could naturally meet or have been brought into 

close proximity by man. 

13. Polylepis incam Humboldt, Bonpland, 

and Kunth 

Polylepis iricaiia Humboldt, Bonpland, and Kunth, 1824:227. 

[Type collection: Los Paston, Guachucal along the Rio 

Blanco, Ecuador, 9684 ft, Dec, Bonpland 2191. Holotype 

PI, isotype PI] 

Polylepis incam \ar. uillosistyla Bitter, 1911b:642. [Type 

collection: El Altar, Ecuador, Meyer 17i. Type not seen, 

presumed destroyed at Berlin.] 

Trees 5-8 m tall. Leaves congested at the ends of 

branches (Figure 34a), imparipinnate with one pair 

of leaflets, trullate in outline, 1.4-3.9 cm wide, 

1.9-6.4 crn long; rachises with short, twisted trichomes and small, multicellular glandular trichomes 

mixed with resinous exudate; point of leaflet attachment (Figure 34c) with a small tuft of long, 

curled trichomes; stipular sheaths sloping away from 

tlie petiole, outer surface with glandular trichomes 

and resinous exudate; inner basal part of petiole 

and inside of the top of tlie sheath villous. Leaflets 

obovate in outline, 0.4-1.4 cm wide (0.5), 1.2-4.0 cm 

long, margins crenate, apex obtuse to slightly emarginate, base attenuate; upper leaflet surface glabrous; lower surface (Figure 34b) densely covered 

with short, twisted, irregular multicellular trichomes 

mixed with resinous exudate and often with sparse, 

52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

bent, long, single-celled trichomes. Inflorescences 

pendant, 2-7 cm long, bearing 3-10 flowers; floral 

bracts to 3 mm long, narrowly lanceolate, glabrous 

or pilose on the outer surface; rachises with glandular trichomes and exudate. Flowers (Figure 344 perfect, 0.3-0.7 cm in diameter; sepals 3 (4), ovate, outer 

surface covered with appressed small multicellular 

trichomes; stamens 8-24, anthers orbicular, barbate; 

base of style villous. Fruit (Figure 34e) turbinate to 

fusiform, irregularly ridged and/or winged with 

pointed wings, 0.2-0.7 cm wide, including wings, 

0.2-0.5 cm long. 

DISTRIBUTION.-Centra1 Ecuador in Pichincha to 

southern Peru in the Department of Cuzco (Figure 

35). Flowering recorded in January, May, June, 

and August to November. 

COMMON NAMES.-Quinuar, quinual, quenuiiia, 

cceuna, manzanita, qqueuna. 

USES.-T~C wood is used by Indians to manufacFIGURE 35.-Collection localities of Polylepis incana. 

ture instruments for cultivation, for house construction, and as firewood. Recently man has used the 

wood in construction of mining shafts at high elevations. Tannin extracted from the branches is used 

for tanning. 

REPRESENTATIVE SPECIMEM ExAMMINED.-ECUADOR. PICHINCHA: Quito, 1845, Jaineson 17 (G, W); l-olcan Iliniza, northern 

slopes, 4000 m, 19 Apr 1969, Sparre I5625 (S); El Chaupi on 

the road to I’olcan Iliniza, 3300 m, 29 Apr 1967, Sparre 

15654 (S); between Pifo and Papallacta, Jan 1955, Fagerlind 

+ Wibonz s.n. (S). COTOPAXI: railroad station Cotopaxi, 

3400 m, 26 May 1939, Asplund 6476 ‘(G, S, US). TUXCURAHUA: 

ricinity of Ambato, 24-28 Aug 1918, Rose L* Rose 22397 

(GH, NY). Azu.4~: between Molleturo and Toredor, 2590- 

3900 m, 14 Jun 1943, Steyermark 538038 ‘(F, XY). PERU. 

ANCASH: Cordillera Raura, 4300 m, Ranh L- Hirsh 22.754 (XY); 

Chiklin, 28 Oct 1927, Sawada P83 (F). HUANCCO: Llata, 7000 

ft, 21 Aug 1922, Macbride 2 Featherstone 2249 (F, G, GH, 

S, US). PASCO: near Oyon, Rio Huaura, 3800 m, 29 Nor 1964, 

h’oepcke 1861bd (UC); below Cerro de Pasco on the road to 

Ambo, 24 Nov 1945, Seibert 2202 (MO, US); between Cerro de 

Pasco and La Quinua, 5 km along the highway above La 

Quinua, 3600 m, 21 Jun 1940, Asplund 11847 (G, S, US). 

Cuzco: Pisac, 3500-3600 m, 1 May 1925, Pennell 13731 (F, 

GH); 3 km east of Cuzco, 3600 m, 1 Oct 1936, West 8053 

(GH, MO, VC); Paucartambo \’alley, Chatica, 3800 m, Aug 

1926, Hrrreya I129 (F, US). 

DiscussIoN.-Undoubtedly Polylepis incana has 

been the most misapplied epithet in the genus. It 

has been used for almost any specimen with a single 

pair of leaflets as well as for most specimens for 

which a name was not readily apparent. Part of the 

problem is that P. incana forms the center of a complex of species, many of which appear to hybridize 

in central Peru. An additional element of confusion 

was introduced by Bitter’s treatment of the taxon. 

Bitter never mentioned Bonpland’s type of the 

species under any of his subspecific taxa, and it thus 

appears possible that he was actually not certain as 

to the correct typification of the name. Kevertheless, 

he circumscribed three varieties and six subspecies 

within P. incana. As correctly pointed out by 

Macbride (1934:367), many of the varieties and subspecies are elements that belong within P. racemosa. 

These include P. incana ssp. micyanthera, P. incana 

var. connectens, P. incana subsp. icosandra and 

P. incana var. primovestita. The type of Bitter’s 

P. incana var. primouestita is the same as the type 

of P. racemosa of Ruiz and Pavon. The variety P. 

incana var. flauipila was the basis on which Macbride erected his new species, P. subsericans. Several other of Bitter’s infraspecific taxa, P. incana 

var. incarum. P. incana ssp. .mibtusalbida, and P. 
NUMBER 43 53 

incana ssp. by-achypoda belong within P. besseri. 

The only one of Bitter’s varieties or subspecies that 

truly appears to belong to P. incana is P. incana 

var. villosirtyla, but it is not considered here to 

merit varietal status. While this assignment of 

Bitter’s subspecific taxa appears to be correct, absolute certainty is impossible because most ot tliz 

types of his taxa were apparently destroyed at 

Berlin. 

As circumscribed here, Polylepis incana includes 

only those individuals with three leaflets having 

short pilose vestiture on the underleaflet surfaces, 

usually mixed with multicellular glandular trichomes and resin. Individuals with shaggy, villous 

underleaflet surfaces are referred to P. racemosa 

and those with a panose covering to P. besseri. In 

central Peru, however, especially in the Pampa de 

Junin, intermediate specimens do occur. It is probable that human planting of this taxon with P. 

racemosa has fostered hybridization. Personal observations indicate that populations include trees 

with an upright, narrow branching pattern as well 

as some with an expanded crown. Intermediate 

specimens have also been seen from La Libertad, 

Peru. Under natural circumstances it appears that 

P. incana was distributed from Ecuador to southcentral Peru on the dry slopes of the Andes (westfacing slopes of the Eastern Cordillera and eastfacing slopes of the TVestern Cordillera), whereas 

P. racemosa inhabited upper valleys of rivers on the 

Tl‘estern Cordillera and the central valleys of Peru 

from Huanuco. 

In Cochabamba, Polylepis incana and P. tomentella are T ery similar morphologically. The difference in leaflet pubescence serves to distinguish the 

two: P. tornetella in this region often has glabrous 

underleaflet surfaces, whereas P. incana has pilose, 

yellowish underleaflet surfaces. Still farther south, 

either P. bessen’ or P. rtigulosa replaces P. incana. 

The dense, tomentose (pannose) covering of the 

undersurface of the leaflets of these two taxa is 

never found in P. incana. 

14. Polylepis racemosa Ruiz and Pavon 

Polylepis raceinosa Ruiz and Pavon,1798: 139. [Type collection: 

Caxamarquilla and Pillao, Peru, Ruiz s.n. Holotype P! 

isotype G!] 

Polylepis uillosa Humboldt, Bonpland, and Kunth, 1824:228. 

[Type collection: Caxamarca, Peru, 8784 ft. Aug, Bonpland 

3685. Holotype P! isotype P!] 

Polylepis incann ssp. icosatidra Bitter, 191 lb:641. [Type of 

collection: Cajatambo, between Tallenga and Piscapaccha, 

Ancash, Peru, Weherbauer 2886. T)pe not seen, presumably 

destroyed at Berlin.] 

Polylepis incana ssp. niicranthera Bitter, 1911b:642. [Type 

collection: Sierra, Caxamarquilla, Huanuco, Peru, without 

collector. Holotype IV!] 

Polylepis incana var. primovestita Bitter, 191 lb:645. [Type of 

collection: Caxamarquilla, Peru, Kuiz s.n. An illegitimate 

name since the same collection is the type of P. racernosa.] 

Polylepis incana var. connectens Bitter, 1911b:645. [Type collection: in the mountains of Huanuco, Peru, Haenke s.n. 

Holotype PR.] 

Trees 4-15 m tall. Leaves congested at the ends 

of the branches (Figure 36a), imparipinnate with 

1-3 pairs of leafleflts; rhomboid to obtrullate in outline, 2.3-5.0 cm wide and 3.5-8.8 cm long, rachises 

villous, especially along a line on the upper surface near the base of the leaf and extending into 

the inside of tlie stipule sheath; point of leaflet attachment (Figure 36c) with a tuft of long white, 

straight trichomes; stipular sheaths with a slight 

shoulder or sloping away from the petiole, glabrous 

on the outer surface but with long trichomes protruding at the apex from the inside surface. Leaflets 

obovate to almost oblong in outline, second pair 

from the terminal leaflet the largest when present, 

a leaflet of this pair from 0.7-0.6 cm wide and 2.1- 

3.7 cm long, margins crenate, especially at the apex, 

apex slightly to deeply emarginate, base attenuate 

or unequal; upper leaflet surface glabrous, dark 

green, sometimes with scattered trichomes in the 

midvein depression; lower surface (Figure 36b) with 

long, shaggy trichomes inserted on the veins but 

loosely covering the entire undersurface, these trichomes mixed with a variable number of short, 

twisted, several-celled, often glandular trichomes. 

Inflorescences pendant, 4.0-1 1 cm long, bearing 3-1 1 

flowers; floral bracts lanceolate, to 3 mm long, 

scarious; rachises with scattered, curled trichomes. 

Flowers (Figure 36d) psfect, 0.9-1.0 crn in diameter, 

sepals 4, ovate, outer surface villous especially at the 

tips, pilose on the tips of the inner surface, often 

with intermixed several-celled glandular trichomes, 

stamens 10-20. anthers orbicular, barbate; style often 

enlarged near the base due to projection of tlie ovary 

from the floral cup, villous at the point of projection. Fruit turbinate (Figure 36e) with 4-5 irregular, 

flattened wings projecting into a point at the top, 

body of the fruit covered with curled, white trichomes mixed with glandular multicellular tri- 
54 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 36.--Polylepis racemosa: a, branch (X 0.5): b, leaf lrom the undersurface (x 1.5); c, point 

of leaflet attachment, upper surface (X 6); d, flower (X 4.5); e, fruit (4.5). 
NUMBER 43 55 

chomes, 0.2-0.9 cm wide including wings, 0.3-0.8 

cm long. 

DIsTRIsUTloN.-Northern Peru in the Province of 

La Libertad and in humid heads of valley on the 

western slopes of the Peruvian Cordillera south to 

Lima and in central Peru from Huanuco across 

the Pampa de Junin to Huancayo (Figure 37). 

Flowers recorded November to February, April, 

May, June, and August. 

COMMON NAME.--Quinuar. 

USES.-AS an ornamental and as firewood. 

REPRESENTATIF E SPECIME~S ExAMIKED.-PERU. LA LIBERTAD: 

Province Santiago de Chuco, above Cachicadan toward the top 

of hills north of Cachicadan, 2900 m, 25 Nov 1938, Stork S 

Hortorz 9972 (F, G, UC), ibid, 3100-3250 m, 8 Jun 1953, Lopez 

0983 (US). HuAxuco: near Cerro de Pasco, 12,000 ft, 4 Feb 

1927, Kanehiyn 5 (F); Province Huainuco, km 321 on Route 3 

north of Quinua, 11 Jan 1977, Simpson 8554 (US). LIMA: 

Rio Blanco, 3000-3500 m, 15-17 Apr 1929, Killip & Smith 

21745 (F, GH, NY, US); Province Huarochiri, Rio Blanco, 

4 Jun 1940, .4splttnd 11374 ‘(G, S, US); ibid, Infiernillo, 3400 

m, 3 Jun 1940, Asplurid I1343 (G, NY, P, S, US); km 111 

on Route 20A between Lima and La Oroya, 12 Jan 1977, 

Simpson 8559 (US). JUNIN: Province Cerro, between La 

Quiniua and Huariaca, Estacion La Aurora, 3300 m, 24 Jun 

1940, Asplurid 119tf (G, S, US); Tarma, 3300-3500 m, 20-22 

Apr 1929, Killip Smith 21899 (F, GH, NY, US); Tarma, 

Ayabamba, 1944, Soilkup 2536 (F, US): between Tarma and 

Oroya, climbing to the summit, 29 Jun 1948, Ferreyra 3792 

FIGURE 37.-Collection localities of Polylepis racemosa. 

(US); Province Jauja, Parco between Jauja and Oroya, 3200- 

3300 m, Sov 1947, Ferreyra 2833 (US); Huancayo, Pampas 

Valley, 3300 m, 24 May 1948, .4nderson 695 ,(US); ibid, Concepcion, 10,800 ft, Jul 1945, Sazilcman 4921 (K). 

DIscussIos.--This type-species of the genus described by Ruiz and Pavon was redescribed 26 years 

later by Humboldt, Bonpland, and Kunth as Polylepis nillosa, but the type specimens of the two taxa 

are practically identical and come from the same 

area, Of the two names, P. uillosa has been most 

frequently used and, next to P. incana, is the name 

given to most of the specimens that fall within the 

incuna complex. Since Bitter’s monograph, the name 

P. racemosa has tended to be given to specimens 

with more than one pair of leaflets and P. uillosa 

to those with one pair, The type specimens of both 

names, however, have only one pair. The collections 

of Killip and Smith (21745) from Rio Blanco above 

Lima show that within one population, plants can 

have leaves with one, two, or three pairs of leaflets. 

An extreme morphological type with large leaflets 

in three pairs is seen in the collection of Asplund 

(11?74), which probably came from a particularly 

favbrable environment (planted?). The leaflets of 

these specimens are quite glabrous-a character 

often found when water is abundant. The flowers 

are also large and numerous. The major part of the 

range of P. racemosa (as outlined in Pulgar, 1967) 

is in the valley of the Rio hlontaro and across the 

Pampa de Junin in central Peru. Enormous populations cover the broad, flat valley from Cerro de 

Pasco to Huancayo. In addition, populations occur 

down to Huanuco, down the valleys toward Lima 

and southward toward Cuzco. More than any other 

species, this taxon is cited as being planted and used 

as an ornamental by man. The trees around Cerro 

de Pasco have all been planted. Polylepis incana, 

also much planted, has been collected from the same 

area. Cerro de Pasco is at a higher elevation, and 

more exposed, than areas where P. racemosa normally grows. Because of this influence of human 

activities, it is difficult to determine the natural 

distribution of P. racemosa, but it seems safe to 

infer that it is a natural inhabitant of the Montaro 

River valley. It is possibly because of man’s influence that hybrids with P. incnna occur. Polylepis 

incnna tends to be a species of drier regions, growing from Ecuador south on the eastern side of the 

Western Cordillera of Peru. If artificially watered, 

P. racemosa could grow in these areas. In southern 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Peru where P. rugulosa is the native species, similar 

overlap may occur due to human planting of P. 

racemosa. 

Morphologically, P. racemosa differs from P. 

incana in the size of the leaflets and the pubescence 

of the underleaflet surface. The former has leaflets 

2.7-4 cm long, whereas in the latter the leaves range 

from 1.2 to 4 cm but usually shorter than 2.7 cm. 

The underleaf surfaces of P. incana have a relatively 

sparse covering of yellow glandular protuberances 

or trichomes and the upper leaf surfaces are often 

dull and yellowish. The leaflets of P. racemosa are 

green above and have villous trichomes on the veins 

of the lower leaflet surfaces. In some populations, 

P. racemosa has emarginate leaflets. Possible hybrids 

include specimens from Huancayo (Hodge 6221) 

and La Libertad (Stork Q Horton 9972.) 

15. Polylepis australis Bitter 

Polylepis australis Bitter, 1911b:619. [Type collection: Sierra 

Santa Barbara, Jujuy, Argentina, 2500 m, 11 Ju1 1901, 

Fries 261. Lectotype S!, isotype US!] 

Polylepis racemosa var. glabra 0. Kuntze, 1898:77. [Type collection: Sierra de Cordoba, Cordoba, Argentina, Schnyder 

s.n. Type presumably destroyed at Berlin, possible isotype 

Polylepis rucemwa var. glabrescens 0. Kuntze, 1898:77. [Type 

collection: Cienaga, Catamarca, Argentina, Lorentz 310. 

Type destroyed at Berlin, collection duplicate at NY!] 

Polylepis racemosa var. albotomentella 0. Kuntze, 1898:77. 

[Type collection: Lm Gigantes, Cordoba, Argentina, F. 

Kurtz, not seen, presumably destroyed at Berlin.] 

Polylepis racemosa var. pubescens 0. Kuntze, 1898:77. [Type 

collection: Cuesta de Copina, Sierra de Achala, Cordoba, 

Argentina, Hieronymus s.n. Type not seen, presumably 

destroyed at Berlin, collection duplicates at BM! G! K! 

NY!; specimen at G! annotated by Bitter.] 

Polylepis racemosa var. subresinosa 0. Kuntze, 1898:77. [Type 

collection: Cuesta de Copina, Sierra de Achala, aodoba, 

Argentina, Hieronymus s.n. Type not seen, presumably 

destroyed at Berlin, collection duplicate at G!] 

Polylepis racernosa Lar. pubit~ervia 0. Kuntze, 1898:77. [Type 

collection: Cerro Champaqui, Argentina. Kurtz s.n. Type 

not seen, presumably destroyed at Berlin.] 

Polylepis racemosa var. fuscotornentella 0. Kuntze, 1898:77. 

[Type collection: Oyada, Argentina, Lorentz 442. Type not 

seen, presumably destroyed at Berlin.] 

NY!] 

Polylepis australis var. gla6ra (0. Kuntze) Bitter, 1911b3622. 

Polylepis australis var. glabrescens (0. Kuntze pro parte) 

Bitter, 191 1b:623. 

Polylepis australis var. subcalva Bitter, 1911b:623. [Type 

collection: Cienaga, Catamarca, Argentina, Lorentz 310 

pro parte. Type presumably destroyed at Berlin, collection duplicate at NY!] 

Polylepis australis var. oblanceolata Bitter, 1911b:623. [An 

illegitimate name since Bitter cites P. racemosa var. pubescens as a synonym.] 

Polylepis australis var. latifoliolata Bitter, 1911b:624. [Type 

collection: north of the Cuesta de Copina, Cordoba, Argentina, Hi: rony71ius s.n. Type not seen, presumably destroyed 

at Berlin.] 

Polylepis austrnlis var. fuscitoinentella (0. Kuntze) Bitter, 

19 I 1 b: 625. 

Polylepis australis \ar. crcnulata Bitter, 1911b:625. [Type 

collection: Sierra Santa Barbara, Jujuy, Argentina, 2500 m, 

11 Ju1 1901, Fries 26f. Holotype S!] 

Polylepis australis var. tucumanica Bitter, 1913b:478. [Type 

collection: Canada del Munoz, Tafi de I’alle, Tucuman,

Argentina, Castillon 22523. Lectotype destroyed at Berlin, 

Isotype G!] 

Polylepis australis var. tucumanica subvar. rnajuscula Bitter, 

1913:478. [Type collection: La Quenoa, Tucuman, Argentina, 2600 m, 11 Mar 1912, Lillo 11257. Type not seen, 

presumably destroyed at Berlin.] 

Polylepis australis var. tucumanica subvar. latifrons Bitter, 

1913:479. [Type collection: Canada del Munoz, Tafi del 

Yalle, Tucuman, Argentina, Castillon s.n. Type destroyed 

at Berlin, Isotype LIL!] 

Polylepis austrnlin tar. tucumanica subvar. gracilescens Bitter, 

1913:479. [Type collection: Angostura, Tafi del Valle, 

Tucumin, Argentina, Castillon s.n. Type presumably destroyed at Berlin, Isotype G!]. 

Polylepis austrnlis var. tucumanica subvar. breviuscula Bitter, 

1913:479. [Type collection: La Cienaga, Tucuman, Argentina, 2500 m, 19 Dec 1908, Lillo 8767. Type not seen, presumably destroyed at Berlin, isotype G!] 

Trees 1.5-10 m tall, Leaves very congested at the 

ends of branches with the area of overlapping 

stipule sheaths swollen (Figure 38a); leaves composed of (1) 2-3 pairs of leaflets, obtrullate in outline, 1.8-5.3 cm wide, 2.0-7.2 cm long; rachises 

glabrescent to densely glandular; point of leaflet 

attachment with a tuft of long white trichomes 

(Figure 38b); stipule sheaths glabrescent, glandular 

or pilose on the outer surface and with long straight 

trichomes projecting from the inside surface near 

the petiole. Leaflets broadly lanceolate, obovate, first 

or second pair from the terminal leaflet the largest, 

one of this pair 0.3-1.5 cm wide, 1.0-4.0 cm long; 

margins crenate, sometimes ciliate, apex obtuse, 

base unequally abrupt or attenuate; upper surface 

glabrescent with a few scattered, twisted trichomes, 

glabrous or densely glandular (Figure 38b); lower 

surface glabrous, glabrescent, glandular or with 

scattered, appressed straight trichomes (Figure 38b). 

Inflorescence pendant, 1.8-7.3 cm long, bearing 2-12 

flowers, floral bracts lanceolate, scarious, sparsely 

glandular, 3-6 mm long, rachises with scattered 
NUMBER 43 57 

FIGURE 38.--PoZyZepis australis: a, branch (x 0.5); h, point of leaflet attachment, upper surface 

(x 3): c, flower (x 6); d, fruit (x 6). 
58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

glandular trichomes and resin. Flowers (Figure 38c) 

perfect, 0.7-1.0 cm in diameter, sepals 3 or 4, ovate, 

outer surface glandular or pilose, stamens 8-16, 

anthers with a tuft of straight trichomes at the apex; 

base of style glabrous or with a few straight trichomes. Fruit (Figure 38d) turbinate, winged, the 

wings irregular and pronounced, 0.2-0.6 cm wide 

including wings, 0.2-0.9 cm long. 

DIsTRIBuTIoix.-hTorthern Argentina in the Provinces of Jujuy, Salta, Catamarca, Tucuman, and 

Cordoba (Figure 39). Flowering recorded June, 

July, October to April. 

COMMON NAMES.-Tabaquillo, quenoa, quenua, 

q ueii ura, 

UsEs.-The wood is used for the manufacture of 

farming implements. 

REPRESENTATIVE SPECIX%ENS EX.~MINED.-ARGENTINA. 

SALTA: Dept. Santa I’ictoria, Santa I’ictoria, 2385 m, 28 Jan 

1942, hleyer 46Q4 (UC); ibid Rodeo Pampa, 3000 m, Jan 1943, 

Meyer 4876 (F, LIL, VC); between Guachipas and Pampa 

Grande, 9 Jan 1961, Meyer 21861 (LIL); Dept. La. Candelaria, 

Cuesta occidental, Sierra del Castillyo, 2000 m, 7 Sov 1931, 

Schreiter 6639 (LIL, S. UC). JUJUU: Laguna de Yala, 13 Jan 

1970, hleyer, Legname h Koyama 126 (LIL). CATAMARCA: 

FIGURF, 39.-Collection localities of Polylepis australis. 

Dept. Andalgaa, Los Quenoales, above the Mesaco de las 

Rosas, 15 Jan 1952, Sleumer 2245 (US); Andalgala, 1800 m, 

12 Mar 1915, Jogensen If55 (GH, MO, US). TUCUMAN: Dept. 

Chicligasta. La Estancia 10s Pavos, 2000 m, 13 Dec 1925, 

T’enturi 3YYO (MO, S, US); ibid, 22 SOT 1926, I’enturi 4591 

(F, GH, UC, US); Saladillo, 1000 m, 18 May 1948, dfeyer 

If063 (MO, FY); Dept. Trancas, Pie a la Cuesta, 21 Apr 1926, 

I’ejttuii +j53 (A, S) ; Dept. Tafi, Cumbre de .infama toward 

La Aguadita, 26 Oct 1934. Sch,.eiei 981f (S, US) ; La Cienaga 

to Tafi del I’alle, 2OOC-2500 m, 15 Jun 1923, Schwiter 3f590 

(A, US); La Cienaga, 2700 m, 6 Oct 1924, Schreiter 89339 

(US); Tafi del I’alle, 24 Sep 1949, Pdncios, Bnrkley, Bnlegno 

19‘4rIO6 (MO, S); Cumbre Anfama, 2700 m, 20 Sov 1927, 

I’enturi 5f97, (A, MO, S); Quebrada Chada del Neunoy, 

23 Mar 1944, Lourteig 553 (.I, US); Quebrada de 10s Glisus, 

Jan 1912, Cnstillon 893ff (US); San Jose, Chaque\il, 2650 m, 

4 Jan 1953, Petersen 6 Hjerting 876 (S); Dept. Burroyaco, 

Cerro del Campo, 2500 m, 30 Dec 1928, Venturi 9171 (BM, 

US); El Saldillo, 12 Sep 1916, Joi-gensen ?I (GH, US); La 

Quenua, 3000 m, 27 Jan 1933, Parodi I0732 (GH); Laguna 

del Tesoro, 18 Oct 1969, Legume 5 l’aca 3741’ (LIL). 

CORDOBA: Dept. Calmuchita, Cerro Champaqui, 2600 m, 14 

Mar 1952, Ki.apovickas 7690 (LIL): ibid, without date, Kurtz 

68f2 (XY); Dept. San Alberto, Mina Clatro, 11 Nov 1952, 

Krapovickns iifi (LIL); Dept. Punilla, la barranca of the rio 

Yuspe, 1780 m, Dec 1949, Meser ck Sleumer 15627 (B); Sierra 

Grande (Achala), 13-15 Jan 1876, Hieronyinus 317 (F, US); 

Pampa de Achala, 2200 m, 13 Dec 1945, Hunriker I398 (MO.) 

DIscussIow.-Because of its isolated geographical 

position in northern Argentina, Polylepis australis 

is a convenient unit for circumscription. As indicated by numerous authors (Bitter, 191 Ib, Zardini, 

1973), it may be only a geographically isolated form 

of either P. racemosa or P. tomentella. Otto Kuntze 

(1898) described as varieties of P. mcemosa many 

populations that have been placed in P. australis by 

Bitter (1911b) and subsequent workers. In morphology, plants from populations included within 

P. australis are distinct enough to be easily recognized. In general, the fruits of this species are more 

winged than those of any other in the genus. In 

addition, the leaflets are usually glabrous or glabrescent; however, there is enormous variation 

within and between populations. Plants even within 

a population can have leaflets varying from small to 

large and glabrous to densely glandular. This 

variability has, unfortunately, been the cause of a 

taxonomic muddle in the treatment of the species. 

When Bitter described Polylepis australis (1911b), 

he neglected to cite any specimens that could be 

associated directly with the specific epithet. Instead, 

after the description of the species, he gave a general 

distributional range and proceeded to circumscribe 

eight varieties. Bitter did not name a variety P. 
NU!vIBER 43 59 

australis var. austruliJ nor did he indicate which 

of the varieties he would consider to contain the 

type of the species. In choosing a lectotype, I have 

chosen the only variety for which there is an extant 

holotype. The type of this variety thus becomes the 

type of the species. Further complications in the 

taxonomy of the species arise because Bitter used 

different specimens of the same collections as types 

of some of his varieties (e.g., Lorentz 310, Hieronymus s.n.). Since the actual specimens used by Bitter 

were destroyed at Berlin, and since he obviously 

considered the collecLioris to be polytypic, it is impossible to refer duplicates of these collections to 

his various varieties. LVhile the descriptions of the 

varieties might provide some clue as to his designation of particular specimens, there is so much variation that designating parts of the collections not 

seen by Bitter would be potentially misleading. Also 

there is no apparent need to recognize subspecific 

taxa, and I have merely given the types as Bitter 

listed them and made no attempt to designate 

lectotypes from the isotypes. 

A similar situation pertains in the case of Polylepis 

australis var. tucumanica and its subvarieties described by Bitter in 1913. The description of the 

variety has no specimens associated with it; however, there are extant isotypes of the collections that 

he cited for the various subvarieties. One of these 

is designated here as the lectotype of the variety 

tucumanica. Still, as in the case of the other varieties 

of the species described by Bitter, there appears to 

be no justification for the retention of this variety 

or its subvarieties. 
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Index 

(Synonyms in italics) 

Acaena ochreata, 28 

Polylepsis albicans, 28 

annulatipilosa, 27 

australis, 56 

australis var. bijuga, 23 

australis var. menulata, 56 

australis var. fuscitomentella, 56 

australis var. glabra, 56 

austrulis var. glabrescens, 56 

australis var. latifoliolata, 56 

australis var. oblanceolata, 56 

australis var. subcalva, 56 

australis var. tucumanica, 56 

australis var. tucumanica subvar. breviuscula, 56 

australis var. tucumanica subvar. <gracilescens, 56 

australis var. tucumanica subvar. latifrons, 56 

awtralis var. tucumanica subvar. majuscula, 56 

besseri, 42 

besseri var. abbreuiata, 42 

besseri var. longipedicellata, 42 

boyacensis, 40 

brachyphylla, 38 

cocuyensis, 40 

coriacea, 21 

crista-galli, 42 

crista-galli var. longiracemosa, 42 

hieronymi, 23 

hieronymi var. dolicholopha, 23 

hieronymi var. saltensis, 23 

hypargyrea, 28 

hypoleuca, 23 

incana, 50 

incana ssp. brachypoda, 42 

incana var. connectens, 53 

incana var. flavipila, 31 

incana ssp. icosandra, 53 

incana ssp. incarum, 42 

incana ssp. micranthera, 53 

incana var. primovestita, 53 

incana ssp. subtusalbida, 42 

incana var. villosistyla, 40 

lanuginosa, 21 

lanuginosa p microphylla, 57 

lehmannii, 21 

microphylla, 87 

microphylla var. polyarthrotricha, 37 

multijuga, 19 

nitida, 33 

ochreata, 28 

ochreata var. integra, 28 

pallidistigmu, 42 

pauta, 27 

pepei, 32 

quadrijuga, 40 

quindiensis, 28 

racemosa, 53 

racemosa var. abbreuiata, 42 

racemosa var. albotomentella, 23 

racemosa var. fuscotomentella, 56 

racemosa var. glabra, 56 

racemosa var. glabrescens, 56 

racemosa p hypoleuca, 23 

racemosa var. lanata, 42 

racemosa var. pubescens, 56 

racemosa var. pubineruia, 56 

racemosa var. subresinosa, 56 

racemosa var. tomentosa, 42 

reticulata, 33 

rugulosa, 42 

sericea, 28 

serrata, 27 

serrata var. parcipila, 27 

serrata var. psilanthera, 27 

stuebelii, 28 

subintegra, 28 

subquinquefolia, 42 

subsericans, 31 

taraplcana, 46 

tarapacana var. brevifilamentosa, 46 

tarapacana var. multisquama, 46 

tarafiacana var. pycnolopha, 46 

tarapacana var. sajamensis, 46 

tenuiruga, 42 

tomentella, 46 

tomentella dentatialata, 46 

tomentella ssp. pentaphylla, 46 

tomentella var. pilosior, 46 

tomentella ssp. tetragona, 46 

triacontandra, 42 

uillosa, 53 

weberbaueri. 37 

62