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JUL 1 9 1982 



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FIELDIANA 
Botany 

Published by Field Museum of Natural History 



New Series, No. 2 



REVISION OF SOUTH AMERICAN 
SAURAUIA (ACTINIDIACEAE) 

DJAJA D. SOEJARTO 



DE 



1980 



May 27, 1980 
Publication 1306 



REVISION OF SOUTH AMERICAN 
SAURAUIA (ACTINIDIACEAE) 



,-. ,* 



FIELDIANA 
Botany 

Published by Field Museum of Natural History 



New Series, No. 2 



REVISION OF SOUTH AMERICAN 
SAURAUIA (ACTINIDIACEAE) 

DJAJA D. SOEJARTO 

Department of Pharmocognosy and Pharmacology, College of Pharmacy 

University of Illinois at the Medical Center 

Chicago, Illinois 

and 

Research Associate 

Field Museum of Natural History 



May 27, 1980 
Publication 1306 



Library of Congress Catalog Card No.: 79-51551 

ISSN 0015-1746 
Printed in United States of America 



CONTENTS 

List of Illustrations vii 

Introduction 1 

History and Orthography 3 

Morphological and Taxonomic Criteria 4 

Ecology, Geography, and Paleobotany 8 

Economic Uses 13 

Systematic Position 14 

Acknowledgments 17 

Taxonomy 19 

Key to Series 20 

I. Ser. Omichlophilae 20 

H. Ser. Gynotrichae 27 

IE. Ser. Laevigatae 28 

IV. Ser. Parviflorae 40 

V. Ser. Pulverulentae 54 

VI. Ser. Macrophyllae 68 

VH. Ser. Lanatae 90 

Imperfectly Known Taxa 100 

Addenda 100 

Appendix: List of Accepted Series and Species 131 

References 132 

Index to Latin Names . . . .139 



LIST OF ILLUSTRATIONS 

1. Altitudinal distribution of South American Saurauia 9 

2. Altitudinal distribution of Mexican and Central American Saurauia 9 

3. Saurauia spinuligera R. E. Schultes 101 

4. Saurauia stapfiana Busc 102 

5. Saurauia caquetensis R. E. Schultes var. caquetensis 103 

6. Saurauia rusbyi Britt 104 

7. Saurauia aequatoriensis Sprague 105 

8. Saurauia glabra (R. & P.) Soejarto 106 

9. Saurauia adenodonta Sleumer 107 

10. Saurauia laevigata Tr. & PI 108 

11. Saurauia strigillosa Tr. & PI 109 

12. Saurauia multinervis Soejarto 110 

13. Saurauia schultesiana Soejarto Ill 

14. Saurauia solitaria Sleumer 112 

15. Saurauia micayensis Killip 113 

16. Saurauia natalicia Sleumer 114 

17. Saurauia pulchra Sprague 115 

18. Saurauia floccifera Tr. & PI 116 

19. Saurauia cuatrecasana R. E. Schultes 117 

20. Saurauia arnoldi Sleumer 118 

21 Saurauia mexiae Killip ex Soejarto 119 

22. Saurauia tomentosa (HBK.) Spreng 120 

23. Saurauia formosa Sleumer 121 

24. Saurauia chaparensis Soejarto 122 

25. Saurauia peduncularis Tr. & PI 123 

26. Saurauia lehmannii Hieron 124 

27. Saurauia scabra (HBK.) Dietr 125 

28. Saurauia tambensis Killip 126 

29. Saurauia excelsa Willd 127 

30. Saurauia brachybotrys Turcz 128 

31. Saurauia meridensis Steyermark 129 

32. Saurauia ursina Tr. & PI. . . .130 



INTRODUCTION 1 

The genus Saurauia, founded by Willdenow in 1801, has about 
280 species (Melchior, 1964) represented in both tropical and sub- 
tropical Asia and America. In the Americas, Saurauia is found from 
Central Mexico in the north to Bolivia in the south, through Andean 
South America. 

Scientific interest in this genus has been almost exclusively 
taxonomic and has consisted largely of descriptive accounts of new 
species as they were discovered. In 1966 Hunter published a revision 
of the Mexican and Central American members of the genus in 
which 22 species were recognized. Prior to Hunter's work, there had 
been only one revisional treatment of the American Saurauia, 
namely that of Buscalioni and Muscatello, which was published se- 
rially in Malpighia, vols. 24-30, between 1912 and 1927, and in 1927 
published in a single volume under the title of "Studio monografico 
sulle species americane del genero 'Saurauia' Willd." Unfortu- 
nately, this work lacks good organization and consistency in 
presentation, contains many errors in spelling of names and places, 
and has been criticized by workers of tropical American floras 
(Standley and Steyermark, 1949; Macbride, 1956). 

Because of the absence of a standard taxonomic work on which 
reliable identification of South American Saurauia can be made, it 
was considered urgent to re-study the South American Group by 
examining the type specimens, specimens examined by Buscalioni 
and Muscatello, and other recent collections. Herbarium studies 
have been supplemented, in part, by field work in several parts of 
Colombia, Ecuador, and Peru during the summers of 1963-1966. 
Nevertheless, more extensive field work is needed because many 

'Part of a doctoral dissertation, originally submitted to the Graduate School of the 
Department of Biology, Harvard University, Cambridge, Massachusetts. Dr. 
Soejarto's present address is Department of Pharmacognosy and Pharmacology, Col- 
lege of Pharmacy, University of Illinois at the Medical Center, 833 South Wood 
Street, Chicago, 111. 60612. An Index to Exsiccatae is available from the author. 



2 FIELDIANA: BOTANY 

species are still known only from a few herbarium collections. In a 
sense, the present work is a sequel of Hunter's 1966 revision of the 
Mexican and Central American species. 

The institutions listed below have loaned specimens of Saurauia 
for study. The realization of this work would not have been possible 
without their generosity and it is acknowledged here with gratitude. 
Citation of the institution follows the standardized herbarium ab- 
breviations according to Holmgren and Keuken (1974) wherever 
possible. 

A Arnold Arboretum, Harvard University, Cambridge, 

Mass. 

B Botanisches Museum, Berlin, Germany. 

BM British Museum of Natural History, London, England. 

COL Institute de Ciencias Naturales, Universidad Nacional, 
Bogota, Colombia. 

ECON Economic Herbarium of Oakes Ames, Harvard Univer- 
sity, Cambridge, Mass. 

F Field Museum of Natural History, Chicago, Illinois. 

FI Herbarium Universitatis Florentinae, Institute Botanico, 

Firenze, Italy. 

G Conservatoire et Jardin Botaniques, Geneva, Switzer- 

land. 

GH Gray Herbarium, Harvard University, Cambridge, Mass. 

HAL Institut fur Systematische Botanik und Pflanzenge- 
ographie der Martin-Luther-Universitat, Halle, Ger- 
many. 

K The Herbarium and Library, Royal Botanic Gardens, 

Kew, England. 

L Rijksherbarium, Leiden, Netherlands. 

MA Institute "Antonio Jose Cavanilles," Jardin Botanico, 

Madrid, Spain. 

MEDEL Herbario de la Facultad Nacional de Agronomia, Medel- 
lin, Colombia. 

MICH University Herbarium, University of Michigan, Ann 
Arbor, Michigan. 

NY The New York Botanical Garden, New York, New York. 

P Museum National d'Histoire Naturelle, Laboratoire de 

Phanerogamic, Paris, France. 

PASTO Institute Tecnologico Agricola, Universidad de Narino, 
Pasto, Colombia. 



SOEJARTO: SA URA VIA 3 

S Department of Botany, Naturhistoriska Riksmuseum, 

Stockholm, Sweden. 
UC Herbarium of the University of California, Berkeley, 

California. 
US United States National Museum, Department of Botany, 

Smithsonian Institution, Washington, D.C. 
USM Herbario San Marcos, Museo de Historia Natural, Lima, 

Peru. 
UV Departamento de Biologia, Universidad del Valle, Cali, 

Colombia. 

YEN Institute Botanico, Caracas, Venezuela. 
Z Botanischer Garten und Institut fur Systematische 

Botanik der Universitat Zurich, Zurich, Switzerland. 

HISTORY AND ORTHOGRAPHY 

The generic name Saurauia was established by Willdenow in 
1801, with a single species, Saurauia excelsa Willd., based upon 
specimens collected by Franz Bredemeyer in Caracas, Venezuela. 
Bredemeyer, the Prussian Kaiser's chief gardener, was sent abroad 
on several botanical expeditions to collect plants for the spring gar- 
den of the emperor. Two of these expeditions were to the Americas 
between 1783 and 1788, under the auspices of Kaiser Joseph II. The 
first, in 1783, was carried out in less than one year. The second, 
begun in 1783, lasted until 1788. Among other regions, Bredemeyer 
visited the Caribbean Islands and Venezuela. He stayed in Caracas 
for two years. Bredemeyer took both living and dried plant speci- 
mens back to Germany and many were forwarded to Willdenow, 
who studied them. Willdenow proposed the name Saurauia to honor 
the private adviser of the emperor, Franz Graf von Saurau, who 
patronized Willdenow's work. Von Saurau was also known as a 
friend and supporter of the natural sciences (Wittstein, 1852). 

In the past, two orthographies were in use, Saurauja and 
Saurauia, partly because in the original Willdenow (1801) paper 
both spellings were used (Saurauja for the species heading, once, 
and Saurauia for the legend of the plate, also once) and partly be- 
cause of an error of W. J. Hooker (1842) who, in a footnote, men- 
tioned that Saurauja was "so named by Willdenow in compliment to 
some botanist of the name of Sauraujo, but who is otherwise un- 
known to fame." Between 1801 and 1934, most botanists used the 
orthography Saurauja, but Gilg (1895) and Gilg and Werdermann 



4 FIELDIANA: BOTANY 

(1925) noted that Willdenow consistently annotated the herbarium 
specimens as Saurauia, which was later conserved in the illustra- 
tion. The diaeresis was used to connect the two vowels, so that the 
name should be pronounced Sau-rau-i-a. Between 1895 and 1934, 
some botanists began to use and accept the orthography Saurauia, 
and since 1934 only this orthography has been in use (Buscalioni 
and Muscatello, 1912-1927; Sleumer, 1934, 1938, 1941; Diels, 1937; 
Schultes, 1943-1963; Steyermark, 1952; Macbride, 1956; Hunter, 
1966; Soejarto, 1968, 1969a, b, 1970). 

In 1968, Paclt proposed to revert to the spelling Saurauja, but this 
proposal was rejected in the 1970 Seattle Congress (Stafleu, 1970). 
Upon Paclt's (1971) insistence and following Nicolson's (Nicolson 
and Brooks, 1974) proposal, the 1974 Leningrad Congress accepted 
the spelling Saurauja, which will become mandatory. 

In view of the fact that the orthography Saurauia has been more 
widely used by modern taxonomists (in South America alone 51 taxa 
have been described under this name between 1912 and 1971, as 
against only 31 described between 1855 and 1934 under Saurauja), 
the orthography Saurauia is used in the present revision. It seems 
that conservation of Saurauia over Saurauja will solve the problem, 
since a reversal to the old orthography (Saurauja) will cause confu- 
sion and inconvenience (cf. Hoogland et al., 1977). 

MORPHOLOGICAL AND TAXONOMIC CRITERIA 

1. Habit 

Members of South American Saurauia are mostly woody shrubs 
and small trees. Several species, however, may attain 30 m. or more 
in height, with a bole of up to 40 cm. in diameter at base. Taxonomi- 
cally, plant habit is of little importance. 

2. Leaves 

The leaves of Saurauia are arranged in a 2:5 phyllotaxy, and in 
most species they are crowded around to clustered behind the tip of 
the branchlets, with internodes barely discernible. In several 
species, however, the leaves are distributed from the tip of to low 
along the branchlets, with the internodes prominent and well- 
defined, and this character has been used principally in the recogni- 
tion of the Series Omichlophilae. 

The shape of the leaf blade varies little, from obovate to elliptic or 
elliptic-oblong, hence it is a poor taxonomic character. The same is 
true of the leaf apex and base, although the leaf apex in S. isoxan- 



SOEJARTO: SA URA VIA 5 

thotricha, which is caudate with acumen up to 4.5 cm. long, and the 
leaf base in S. spinuligera, which is truncate to subcordate, are rare 
exceptions. 

In several species, the leaf margins at base bend abruptly upward 
in the direction of the apex, on the upper leaf surface, and fuse with 
the midrib, and among themselves (at their tips) at a short distance 
above the leaf base, to form a boat-shaped structure, the so-called 
"basal flap" (Schultes, 1963b). This was thought to be an important 
taxonomic character. Field work, however, indicated that such a 
flap, which may be complete or incomplete, depending on the degree 
of fusion, can be found within a species population and, moreover, 
flap- and non-flap-bearing leaves may be found in the same tree, 
indicating that the character has no taxonomic significance. The 
function of the basal flap is not clear, since examination showed that 
none is associated with insect habitation (cf. S. aromatica). 

Leaf margins are mostly serrulate to rarely serrate, while suben- 
tire margins represent a variant of distantly serrulate ones. Leaves 
with entire margins are not represented, but denticulate or double- 
serrated margins may be found. Usually, leaf margins are of little 
taxonomic importance. 

Venation of the leaves is penninerved and the number of lateral or 
secondary veins, which ranges from 5 to 40 pairs, is a useful 
taxonomic character. In most species, the tertiary veins jut out from 
the lower leaf surface and are more prominent than the lesser vena- 
tion ("lesser reticulum," see Hunter, 1966), while in others they are 
immersed and scarcely distinguished from or scarcely more promi- 
nent than the lesser venation. This character is important in group- 
ing the species into series. 

Undoubtedly, the types of trichomes and pubescence, in particular 
those on the lower surface of the leaf blade, are very important 
taxonomic characters. In this respect, members of the South Ameri- 
can group have the same general trichome types and pubescence as 
those from Mexico and Central America, for which Hunter (1966) 
has proposed terminology, adopted in the present revision. 

3. Inflorescence 

The inflorescence ofSaurauia is basically a thyrse and is axillary 
in position. A great number of the South American species have 
many-branched and many-flowered inflorescences (with up to 500 
flowers per inflorescence), but few have little branched and few- 



6 FIELDIANA: BOTANY 

flowered inflorescences (as few as three flowers in S. omichlophila). 
In addition to this character, inflorescence size and pubescence are 
useful for identification. 

The bracts and bracteoles may be foliaceous, linear, triangular or 
subulate. With the exception of S. peduncularis, the presence of 
foliaceous bracts is not a constant feature among members of the 
South American Saurauia. 

4. Flower 

Usually, the flowers of Saurauia are described as bisexual and 
sometimes as polygamo-dioecious. Field and herbarium studies 
showed, however, that they should be described as functionally dioe- 
cious (Soejarto, 1969a). 

Floral parts are normally pentamerous, including the androecium 
(Brown, 1935). However, irregular repetition of the floral parts is 
not uncommon among the South American species, where sextam- 
ery, heptamery, and even octamery of the calyx, corolla, and 
gynoecium are found in the same inflorescence, although in such 
cases the pentamerous flowers always predominate. An exception is 
the tetramerous condition of the perianth parts and the gynoecium 
of S. yasicae, where this feature is constant and is a good taxonomic 
character. 

The size of the flowers vary from less than 10 mm. to 50 mm. 
broad at anthesis. Though interspecific variation is more or less 
continuous, flower size is often useful for plant identification. 

The pubescence of the sepals is an important taxonomic character. 
Among the South American species, there are almost as many 
species with glabrous to glabrescent sepals as those with pubescent 
sepals. The greater number of the species in the Series 
Omichlophilae, Laevigatae, and Parviflorae have glabrous to 
glabrescent sepals, whereas all members of the Series Pulverulen- 
tae, Macrophyllae, and Lanatae have pubescent sepals which are 
glabrous on the inner surface, others pubescent throughout, and still 
others with sepals only partially pubescent, usually on the upper 
half. Further, subdivisions may be distinguished on the basis of the 
distribution of pubescence over the outer surface. This may be (1) 
glabrous, (2) pubescent on the parts exposed and glabrous on the 
parts covered (imbricated) in bud, (3) glabrous on the parts exposed 
and pubescent on the parts imbricated in bud, and (4) two kinds of 
pubescence may be present on the parts exposed in bud, in which case 
the parts imbricated may be either glabrous or pubescent. 



SOEJARTO: SA URA VIA 1 

In addition to pubescence, the types of the trichomes on the sepals 
contribute further to the classification of the species. Those occur- 
ring inside (on the inner surface) and on the outer imbricated sur- 
face in bud are also present on the outer surfaces exposed in bud 
when the vestiture is heterotrichous (of two or more types of 
trichomes). Sericeous, strigose, hirsute, and shaggy trichome types 
may occur on the exposed surfaces in bud. 

The sepals are persistent and the base fused. The margins and 
apex are usually ciliolate to ciliate, rarely subentire. In contrast, the 
petals, which are free for most of their length but fused at base, fall 
as a unit with the stamens, after anthesis. As a result, many flowers 
have been described as "unisexual" or "male" and "female" when 
examined after anthesis. In all species studied, the petals are gla- 
brous and in the great majority the color is white. Usually, the 
petals are larger than the sepals. 

In Saurauia the stamens are free for most of their length, and in 
the South American species the number varies from 13 to 240 per 
flower. Variable as it is, stamen number is a convenient diagnostic 
feature, though tedious to record (Soejarto, 1969a). The filament is 
filiform and its length varies from 1.5 to 4.5 mm. The length of the 
filament may be a useful taxonomic character as an aid to identifi- 
cation, as is also the size of the anther, which varies from 1 to 4 mm., 
depending on the species. The shape of the anther is usually linear, 
but it is obcordate in the species with small flowers. 

5. Fruit 

The fruit of Saurauia, which is a berry, is green when fresh, even 
when mature, the rind fleshy and glossy. It may split upon handling, 
but the septae do not separate from the central column. Fruit size 
and characteristics are not of taxonomic importance. 

6. Cytology 

Chromosome counts of 15 species from South America showed 
that chromosome number may be of little importance in the 
taxonomy of the genus (Soejarto, 1970). In all the species investi- 
gated, it was found that the gametic chromosome number (n) equals 
30. 

7. Species grouping 

Seven series are recognized to comprise the South American 
species. Of the 49 species accepted, only S. yasicae definitely extends 
to Central America and Mexico. The pubescent ovary of S. 



8 FIELDIANA: BOTANY 

loeseneriana justifies its inclusion in the Series Gynotrichae, also 
represented in Central America (Hunter, 1966). 

ECOLOGY, GEOGRAPHY, AND PALEOBOTANY 

Species of Saurauia are characteristically adapted to humid or 
wet habitats, such as near waterfalls, along streams, in gullies and 
ravines, along river banks, in the humid mountain and rainforest 
areas, in mossy forests, and in cloud-covered regions. Some species 
prefer shady and misty habitats, while others prefer sunlit places. 
Among the South American species, there are more adapted to high 
elevations (over 1,500 m. altitude) than to lower elevations (below 
1,500 m.), and there are more species found in undisturbed habitats 
than in disturbed habitats. Although plants often grow in pastures, 
on open hillsides, along roadsides, in sugarcane plantations, in 
second-growth vegetation, and even in gardens, the chances are that 
these plants were part of the natural vegetation found in the area 
and have been left behind deliberately during forest clearing, and 
that diligent search will show that members of the species are still 
found growing in their natural environment. Many species grow on 
sandy soil, others on rich humus and loamy soil, more rarely on 
rocky and lateritic soil. The highest altitude which members of 
South American Saurauia tolerate is 3,600 m., represented by S. 
bullosa, and the lower limit is almost at sea level, represented by S. 
pseudoleucocarpa, S. parviflora, S. mexiae, and S. yasicae. While 
most species from lower altitudes are glabrescent or sparingly 
pubescent, those from high altitudes may be glabrescent or densely 
pubescent. Species with chartaceous leaves and those with coriae- 
ceous to strongly coriaceous leaves are found both in low as well as 
in high altitudes. 

As a result of the altitudinal pattern of distribution, the geograph- 
ical range of the South American species is limited to the Andean 
mountain system, from Venezuela in the north to Bolivia in the 
south. No species are represented in the "puna" vegetation of Peru 
and Bolivia. 

A careful examination of the ecological map of Colombia (Espinal 
and Montenegro, 1963), which is based upon the Holdridge system, 
shows that the distribution pattern for the majority of the Colombian 
species follows closely that of the vegetation types known as humid to 
wet submountain and mountain (montane, see Smith and Johnston, 
1945) forests and subparamo (subalpine vegetation), with a mean 
annual rainfall of 1,000-4,000 mm. and a mean annual temperature 



SOEJ ARTO: SA URA VIA 



Number of 

30 
25 
20 

15 
10 
5 




species 


17 


31 


30 


24 


5 








6 


10 








1 


500 1000 1500 2000 2500 3000 3500 4000 (m.) 



FIG. 1. Altitudinal distribution of South American Saurauia.(68.5 per cent of the 
species are found at altitudes of 1,500-3,000 m.) 

Number of species 



20 



18 



15 










13 














12 


10 - 














7 










5 


4 


























2 
















500 1000 1500 2000 2500 3000 3500 (m.) 



FIG. 2. Altitudinal distribution of Mexican and Central American Saurauia. (69.3 
per cent of the species are found at altitudes of 1,000-2,500 m.) 

of (8-) 10-20 (-24)C. Records of meteorological observations (1954- 
1961) from several stations in Narino and Putumayo, deposited at the 
Geographic Institute of Agustin Codazzi in Bogota (Colombia), con- 
firm the rainfall and temperature ranges cited above. A histogram of 
altitude classes shows that 1,500-3,000 m. represents the optimal 
altitude range for the South American species (fig. 1). Similarly, a 



10 FIELDIANA: BOTANY 

histogram constructed for the Mexican and Central American species 
(fig. 2) shows that the optimal altitude range is 1,000-1,500 m. The 
higher optimal range for the South American species is due to the 
telescoping effect of the equator. 

A strong species concentration is located near the Colombo- 
Ecuadorian frontier (the Narino-Putumayo region), just before the 
Andes splits into the three Colombian Cordillera systems, i.e., the 
Occidental, Central, and Oriental. The great concentration of 
species in this region is due, probably, to the rugged nature of the 
topography and to the diverse types of habitats available which 
have permitted a faster rate of speciation. In a rough sketch, such 
diversity is represented by the complex "nudos de los pastes" with 
their peaks (above 4,000 m.) and ridges and the superandean plane 
and valleys in between, the subalpine and alpine vegetations, the 
western and eastern forested slopes, and the coastal zone with its 
tropical rain forest and mangrove swamps. At least 12 species are 
found in this region, of which six are restricted to the area. 

Another high species concentration is located in the Choco- 
Caldas-Antioquia region of Colombia, where the western and cen- 
tral cordilleras, which are narrowly separated by the Cauca River 
valley, almost meet. The isolation of these two cordilleras may be 
considered incomplete in this part of Colombia. At least 16 species 
are found here, but only four of these (S. spinuligera, schultesiana, 
laevigata, strigillosa) are restricted to the area. It appears that the 
Choco-Caldas-Antioquia triangle is not a center of speciation, but 
rather a melting pot between the western and central cordilleras. 

On the basis of the present data, the distribution of the species of 
Saurauia in the Andean countries can be summarized as follows: 

Country Number of Species Endemic species 

Venezuela 6 1 

Colombia 29 22 

Ecuador 12 7 

Peru 11 5 

Bolivia 5 3 

The high number of species and endemism in Colombia is expected, 
and is believed to be the product of a more rapid rate of speciation, 
brought about by the existing diversity of physical conditions, 
created through the ramification of the Andes. The valleys of the 
Magdalena and Cauca rivers, which run parallel in a north-south 
direction, provide ideal lowland barriers to isolate populations of 



SOEJARTO:SA[/flAC7/A 11 

Saurauia, and it is apparent that the ridges and slopes have pro- 
vided routes of migration. The low number of species found in Peru 
and Ecuador may be deceiving, however, since this may be due to 
the lack of botanical explorations, a condition apparent from the 
herbarium representation. Further field work in these areas may 
turn up new records and, perhaps, new taxa. 

Central and South America are separated by a lowland barrier 
starting at the isthmus of Panama, extending southward to north- 
western Colombia. The only species known, beyond any doubt, to 
occur in both Central and South America is Saurauia yasicae. It is 
suspected that S. brachybotrys may also be represented in Mexico 
and Central America, where it may be known as S. aspera (see 
Galeotti 7235). That no species are known from the Antilles may be 
accounted for by the lack of obvious mechanisms for long-distance 
dispersal (Hunter, 1966; Soejarto, 1969a), whereas their absence in 
the Guiana highlands and in the highlands of Brazil may be due to 
the arid conditions which prevail on these plateaus. In addition, the 
great savannas of the Orinoco in the north and the Gran Chaco in 
the south effectively isolate these highlands from the Andes. 
Further south, frost limits the distribution of Saurauia, as is also 
true at altitudes above 3,600 m. 

In its global aspect, the present-day distribution of the genus goes 
as far north as the Tropic of Cancer, and as far south as the Tropic of 
Capricorn. In Asia, where it is distributed from India to the Fiji, the 
number of species (over 170) is greater than that in the Americas 
(71). A similar disjunct distribution is known for many other genera 
of flowering plants (Good, 1953; van Steenis, 1962; Thome, 1972), 
and it is well established that genera with such a discontinuous 
pattern of distribution (amphi-Pacific tropical) represent survivors 
of once extant arcto-tertiary (boreotropical) flora, which has failed to 
survive in Eurasia and North America (Raven and Axelrod, 1974; 
Wolfe, 1975). Unfortunately, paleobotanical evidence is inconclu- 
sive with regard to Saurauia. 

Three mentions of fossil Saurauia have been made, all of them 
based on vegetative organs (leaves). Hollick (1936) described a 
leaf impression from the Eocene deposits of the Kupreanof Island 
(Hamilton Bay) of Alaska as S. alaskana. The identification of the 
specimen was made by comparing it with photographs of other fossil 
Saurauia from Europe, and not with recent collections. Hunter 
(1966) raises serious doubt as to the validity of such identification, 



12 FIELDIANA: BOTANY 

and, in addition, he questions whether the following characters 
which appear in Rollick's photograph are referrable to Saurauia: (1) 
the rounded apex, and (2) the tertiary veins, which are perpendicu- 
lar to the midrib. Of the five species from Mexico and Central 
America (incl. S. yasicae) with tertiary veins perpendicular to the 
midrib, all have immersed tertiary veins, and none of the Mexican 
and Central American species has leaves with rounded leaf apex. 
Among the South American species, the prominent tertiary veins of 
some, e.g., S. excelsa and S. multinervis, tend to be perpendicular to 
the midrib nearest to the apex, and, in particular, the prominent 
tertiary veins of S. micayensis become perpendicular to the midrib 
on the upper half of the leaf blade. To be sure, all South American 
species have either an acute, acuminate, or, rarely, a caudate leaf 
apex, but a rounded apex is within the range of variation of S. 
arnoldi, S. yasicae, and S. pulchra. Furthermore, the acuminate 
apex of obovate leaves may be broken during handling as to give the 
apex a rounded appearance. The leaf margins in several species, 
notable in S. yasicae and S. laevigata, also tend to be somewhat 
entire in the lower half of the blade. While all these indicate that the 
leaf characters described as S. alaskana may be within the range of 
variation of Saurauia, this identification must be accepted with res- 
ervation. As Hollick himself stated, he was actually uncertain as to 
whether the specimen should be referred to Saurauia, Juglans, or 
Hickoria, especially the latter. Ficus alkalina Lesq. (1883), referred 
to by Hollick as resembling his specimen, appears to match better 
the leaf of Saurauia, notably S. yasicae. At any event, the evidence 
for the presence of fossil Saurauia in the New World is not definite. 

In 1870 Ettinghausen described S. deformis from a leaf impres- 
sion of the Tertiary of Croatia, which was earlier described by Unger 
as Juglans deformis. Ettinghausen referred the specimen as sugges- 
tive of certain Mexican species, but Hunter (1966) refuted this refer- 
ence. More notable, however, is the leaf impression from the Ter- 
tiary (Eocene) formation of Sezanne, France, described by Langeron 
(1900) as S. roborans. Langeron indicated that the leaf impression 
resembles the leaves of S. tristyla, an Asian (Indonesian) species. 

A collection of clay seeds from the upper Eocene flora of Hordle, 
Hants (Great Britain) was described by Chandler (1926) as Ac- 
tinidia crassisperma. These seeds, as illustrated by Chandler, may 
very well be the seeds of Saurauia, since there is little difference 
between the seeds of the two genera. 



SOEJARTO: SA URA VIA 13 

In the light of recent paleobotanical evidence and interpretation 
(Graham, 1972; Wolfe, 1972, 1975), which (1) support the relation- 
ship between the Eocene floras of Europe and western North 
America, and (2) establish that the present Indomalayan flora rep- 
resents a relict of the once widespread northern hemisphere tropical 
flora, it is safe to say that Saurauia may have indeed lived in 
Europe and western North America during the Tertiary. In the 
Americas, climatic deterioration during the Oligocene may have 
forced Saurauia populations to migrate southward. However, the 
arrival of Saurauia in South America probably did not take place 
until after the Central American land bridge (the Panama isthmus) 
was well established in the Pliocene, about 5.7 million years before 
present (Simpson, 1950; Lloyd, 1963; Emiliani et al., 1972; Heezen 
et al., 1973). 

ECONOMIC USES 

1. Fruit. The fruit of Saurauia is a berry filled with numerous 
small seeds embedded in a mucilaginous pulp. When a fruit is ma- 
ture, this mucilage is clear, somewhat sticky and sweet. Standley 
and Steyermark (1949) noted that the "pulp is very thin and in 
consistency much like the white of a raw egg." The natives around 
Pasto (Colombia) collect the fruits to eat and, occasionally, offer 
them with other wild fruits for sale in the local marketplace. The 
most popular species in the Narino-Putumayo region of Colombia 
are S. brachybotrys, S. bullosa, and S. tomentosa, all known as 
"moquillo." In other parts of South America, the fruits of other 
species are also known to be edible (also see Perez- Arbelaez, 1956). 
In the Guatemalan markets and Central American countries, the 
fruits are sometimes offered for sale (Standley, 1937; Standley and 
Steyermark, 1949). 

2. Wood. The use of the trunk and branches for firewood is a 
common practice, as various field workers have noted. For example, 
Cabrera (coll. no. 154} observed that the wood of S. ursina is used for 
making charcoal. The stems of Saurauia have a soft pith which is 
hollowed out and used in Honduras by the natives as blowguns 
(Hunter, 1966). In the Sibundoy Valley of Colombia, the straight 
trunks of S. brachybotrys (3-6 m. long), which commonly grow in the 
area, are used locally in house constructions. 

Although many species grow to 20-30 m. tall with a diameter up 
to 40 cm. at base, and an appreciable volume of wood, Saurauia 



14 FIELDIANA: BOTANY 

wood is not of commercial importance (also see Acosta-Solis, 1960). 
Record and Hess (1943, p. 508) describe the wood of Saurauia as 
follows: 

Wood pale reddish brown throughout; not attractive. Luster rather low. Odor- 
less and tasteless. Rather light, but firm, tough, and strong; sp. gr. (air-dry) 
0.58; weight 36 Ibs. per cu. ft.; texture medium; grain straight; easy to cut, 
saws finely wooly, is rather hairy under the plane; probably not durable. Use- 
ful locally for general carpentry and interior construction. 

3. Other uses. Aside from its edible fruits and its wood, little is 
known as to other uses of Saurauia in the Americas. Uses commonly 
related to folk medicine are generally questionable in efficacy. Ac- 
cording to a native of the Sibundoy Valley, the pith of moquillo (S. 
brachybotrys) may be extracted and applied to snake bite to soothe 
or even cure it. Likewise, Schultes (Schultes 3203, also from the 
Sibundoy Valley) noted that "the bark rasped and the powder 
applied to sores to extract pus." In Central America, Standley (1923, 
p. 814) observed that "the fruit (ofpipicho, a species of Saurauia ) is 
said to be sweet and mucilaginous, and syrup made from it is ad- 
ministered for chest affections." 

SYSTEMATIC POSITION 

The systematic position of Saurauia has long been the subject of 
discussion and has not been settled. Saurauia is closely related to 
Actinidia, and the current taxonomic practice is to refer Saurauia to 
either Actinidiaceae, Dilleniaceae, or Saurauiaceae. 

When Willdenow (1801) established the genus, he placed it within 
Polyandria Pentagynia (Tiliaceae), which De Candolle (1822, 1824) 
later included in Ternstroemiaceae (Theaceae). Actinidia, described 
in 1836 by Lindley, was originally assigned by him to Dilleniaceae. 
Several years later, Siebold and Zuccarini (1843) recognized the 
similarities between Actinidia and Saurauia, when the fruits of the 
former were collected (Lindley did not have fruits). They then as- 
signed these two genera, with some doubt, to Ternstroemiaceae. In 
the same year, Endlicher, who had earlier (1840) treated Saurauia 
in Theaceae and Actinidia in Dilleniaceae, in the third Supplement 
to his Genera Plantarum assigned the two genera within the tribe 
Sauraujeae of Ternstroemiaceae. Bentham (1861) and later Ben- 
tham and Hooker (1867) followed Endlicher. Baillon (1873) sepa- 
rated the two genera again, and placed Saurauia in Theaceae and 
Actinidia in Dilleniaceae. Gilg (1895), while placing the two genera 



SOEJARTO: SA URA VIA 15 

in Dilleniaceae, treated Saurauia under Saurauieae and Actinidia 
under Actinidieae. Van Tieghem (1899) separated Saurauia and 
Actinidia from Dilleniaceae and defined a new family, Ac- 
tinidiaceae, to include these two genera. Hallier (1912) felt that 
Saurauia and Actinidia should be accommodated in Clethraceae (of 
Ericales), and assigned them there together with Clematoclethra, in 
the tribe Saurauieae. Lechner (1915), who made a study of the com- 
parative anatomy of the ovules of Actinidia, Clethra, Clemato- 
clethra, and Saurauia, agreed with Hallier. Schnarf (1924), how- 
ever, on the basis of the comparative embryology of Saurauia and 
Actinidia, did not fully agree with Hallier. On the one hand, he 
recognized the gap between the two genera; on the other, he 
acknowledged a close relationship between them and to both the 
Theaceae and Clethraceae. Gilg and Werdermann (1925) accepted 
Actinidiaceae, but also included in it Clematoclethra and Sladenia. 
Hutchinson (1926, 1959) recognized Actinidiaceae as distinct from 
Dilleniaceae, but, unlike Van Tieghem, he restricted the lianous, 
predominantly dioecious Actinidaceae to the single genus Actinidia. 
He established Saurauiaceae as a separate family comprised only of 
the genus Saurauia. In his system, he placed Saurauiaceae together 
with Actinidiaceae in the Theales, and Dilleniaceae in the Dil- 
leniales. Engler and Diels (1936) again united Saurauia and Ac- 
tinidia under Actinidiaceae, which, along with Dilleniaceae, they 
placed in the Theineae of the order Parietales (followed by Law- 
rence, 1951). Record and Hess (1943), who made extensive studies 
on the wood anatomy of the New World angiosperms, firmly sup- 
ported Hutchinson in accommodating Saurauia in a separate fam- 
ily, Saurauiaceae, which Record (1926) had previously noted was 
related to Theaceae. Metcalfe and Chalk (1950), in their Anatomy of 
the Dicotyledons, again stressed the separation of Saurauia from 
Actinidia into two distinct families. Erdtman's (1952) conclusion on 
the basis of his pollen studies supports the opinion which refers 
Saurauia to Clethraceae. Melchior, in the 1964 edition of Engler's 
Syllabus, treated Saurauiaceae as a synonym of Actinidiaceae, 
which comprises three subfamilies, Actinidioideae (Actinidia), 
Saurauioideae (Saurauia), and Clematoclethroideae (Clemato- 
clethra). He placed the family along with Dilleniaceae and Theaceae 
in the Guttiferales. Hunter (1966) believed that Saurauia should be 
assigned to Clethraceae, but he did not do so, referring it, instead, to 
Dilleniaceae. Cronquist (1968) recognized Actinidiaceae (sensu Van 



16 FIELDIANA: BOTANY 

Tieghem) and placed it, together with Theaceae, in the Theales, 
while Takhtajan (1969) maintained the separation of Actinidia from 
Saurauia as two separate families, Actinidiaceae and Saurauiaceae, 
both of which were placed under Ericales, together with Cle- 
thraceae. 

The inclusion of Saurauia and Actinidia in the Dilleniaceae is 
usually based upon the presence of raphide-bearing cells in these 
two genera, a phenomenon common to members of this family. This 
character has been held by various authors not to be of such sys- 
tematic importance as are the characters of floral structure. Wood 
anatomy does not support the inclusion of Saurauia in the Dil- 
leniaceae, since the wood of Saurauia lacks the broad rays charac- 
teristic of the family (Record, 1926) and since spiral thickenings in 
the vessels of Saurauia are absent in Dilleniaceae (Dickison, 1967). 
In addition, the seeds of Saurauia are non-arillate, as opposed to the 
arillate seeds of all members of Dilleniaceae. 

The inclusion of Saurauia and Actinidia, in Theaceae 
(Ternstroemiaceae) is understood for obvious reasons, among them: 
pentamerous flowers; quincuncial aestivation of sepals; numerous, 
distinct, and epipetalous stamens; one, multi-locular pistil; axile 
placentation; distinct styles (for most of their length but fused at 
base); non-arillate seeds; and a straight embryo embedded in endo- 
sperm. Cytologically, Saurauia also seems to be allied to Theaceae 
(Soejarto, 1970), whereas embryological and anatomical studies 
show close alliance between Actinidia and Theaceae. However, the 
ovules in Theaceae are chiefly bitegmic and crassinucellate, the 
fruit a capsule, as opposed to the unitegmic and tenuinucellate 
ovules, and a berry type of fruit of Actinidaceae (sensu Van 
Tieghem). Nevertheless, the frequent practice of including 
Saurauia and Actinidia in the Theaceae, before Van Tieghem estab- 
lished his Actinidiaceae, indicates a close affinity between the two. 

On the basis of similarities in pentamerous flowers, quincuncial 
aestivation of the sepals, anther orientation at anthesis, unilacunar 
nodal anatomy, axile placentation, tricolporate unornamented pol- 
len, numerous small seeds (with straight embryo) embedded in en- 
dosperm, woody habit, mountainous tropical habitat, and epipetal- 
ous stamens, Saurauia with its close allies Actinidia and Clemato- 
clethra should, according to Hunter (1966), be referred to Cle- 
thraceae. However, as pointed out by Cronquist (1968), the high to 
extremely high number of stamens, and the often wholly separate 



SOEJARTO: SA URA VIA 17 

styles establish a distinct gap between Saurauia and its allies, and 
Clethraceae. In addition, Actinidiaceae (sensu Melchior, 1964) have 
axillary inflorescences, whereas Clethraceae have terminal in- 
florescences (with the exception of Schizocardia, which has the in- 
florescence subterminal). Further distinctions in inflorescence 
structure and type of fruit may be made between these two families. 

The close affinity between Saurauia and Actinidia is demon- 
strated by the following commonly shared characters: numerous, 
distinct, epipetalous stamens; one multi-loculed pistil; unitegmic, 
tenuinucellate ovules; areolate, nonarillate seeds (fruit a berry). 
Basically, these are characters used by Van Tieghem (1899) to de- 
fine his Actinidiaceae. That most members of Saurauia are func- 
tionally dioecious (Soejarto, 1970 and unpublished data) demon- 
strates further a close affinity between this genus and the predomi- 
nantly dioecious Actinidia. 

On the basis of all considerations, Saurauia and Actinidia, with 
their ally Clematoclethra, should be placed together in the same 
family Actinidiaceae. In determining relationship, floral characters 
should have more weight, and herbarium studies of the groups con- 
cerned further confirm this statement. 

Regarding relationships among the higher categories, Cronquist 
(1968, p. 194) said: 

"The Actinidiaceae are clearly out of place in the Dilleniales, because of their 
syncarpous ovary with unitegmic, tenuinucellate ovules, in contrast to the 
apocarpus gynoecium and bitegmic, crassinucellate ovules of the Dilleniales. 
The Theales are also chiefly bitegmic and crassinucellate, but there are some 
exceptions and transitional forms. Both Marcgravia and Hypericum, for exam- 
ple, are essentially tenuinucellate, and Marcgravia also shows stages in the 
fusion of the integuments. It is but a short further step to the single integu- 
ment of the Actinidiaceae. The ovular structure of the Actinidiaceae would be 
perfectly at home in the Ericales, as would the frequently poricidal dehiscence 
of the anthers. The large number of stamens and often wholly separate styles 
would be out of place in the Ericales, however. Altogether the Actinidiaceae 
seem to be most at home in the Theales, where they may be regarded as fairly 
closely related to the Theaceae and near-ancestral to the Ericales." 

ACKNOWLEDGMENTS 

To Dr. Richard E. Schultes, my former academic advisor who 
supervised the present work, I wish to express my sincere thanks 
and appreciation for his support and constant encouragement. 

To friends and colleagues who have helped me directly or indi- 
rectly during the preparation of this work, and to those whose 



18 FIELDIANA: BOTANY 

generous and kind collaboration have made my field work a pleas- 
ant experience, many thanks are due. Special thanks go to Drs. B. G. 
Schubert and L. I. Nevling, who kindly read the manuscript and 
offered useful criticisms. My deepest gratitude goes to Dr. A. J. G. H. 
Kostermans, my academic advisor while I was an undergraduate in 
Indonesia, who introduced me to Botany, and whose support made 
this work materialize. Finally, to my dedicated wife, Mariela, who 
has continuously given me encouragement and who typed the man- 
uscript in the early stages of preparation, I want to express my 
fondest thanks and appreciation. 



TAXONOMY 

SAURAUIA WILLD., Ges. Naturf. Freunde Berlin Neue Schriften 
3: 407. 1801 (Saurauja-diagn., Saurauia-pl.).Type species: S. 
excelsa Willd. 

Scapha Noron., Rel. PI. Javan. m Verb. Batav. Genootsch Kunsten 
5 (ed. 1), Art. 4: 3. 1790 (nom. nud.). 

Palaua R. & P., Prodr. Fl. Peruv. Chilens. 100, pi. 22. 1794 (non 
Cavanilles, 1785). Type species: Palaua lanceolata R. & P. 

Apatelia DC., Mem. Soc. Phys. Geneve 1: 426. 1822. 

Vanalphimia Leschen. ex DC., Mem. Soc. Phys. Geneve 1: 419. 1822, 
pro syn. 

Marumia Reinw. ex Bl., Cat. Gewass. Buitenzorg 79. 1823. 

Leucothea Moc. & Sesse ex DC., Mem. Soc. Phys. Geneve 1: 419. 
1822, pro syn. 

Davya Moc. & Sesse ex DC., Prodr. 1: 525. 1824, pro syn. 

Reinwardtia Bl. ex Nees, Syll. PI. Nov. Ratisb. 1: 196. 1824 (n. v.) 
(non Spreng., nee Dum., nee Korth.). 

Tonshia Hamilt. ex D. Don, Prodr. Fl. Nepal. 225. 1825, pro syn. 

Blumia Spreng., Syst. Veg. 3: 126. 1826. 

Overstratia Desch. ex R. Br. in Bennet, PI. Jav. Rar. 171. 1840. 

Obelanthera Turcz., Bull. Soc. Imper. Naturalistes Moscou 20 (1): 
148. 1847. 

Draytonia A. Gray, U.S. Explor. Exped. Bot. 1: 206. pi. 15. 1854. 
Trematanthera F. v. Muell., Victoria Naturalist 3: 71. 1886. 

Pubescent trees and shrubs. Leaves simple, spirally arranged, petiolate, pen- 
ninerved, exstipulate. Inflorescences basically thyrsiform (sometimes reduced to a 
single flower in Asiatic species), axillary. Flowers bisexual, actinomorphic, basically 
pentamerous (tetramerous in S. yasicae), bracteolate, buds usually globose (ellipsoi- 
dal in S. adenodonta); sepals 3-6(-8), usually 5, mostly green and somewhat fused at 
base, persistent, aestivation quincuncial, outer ones usually somewhat smaller and 

19 



20 FIELDIANA: BOTANY 

more densely pubescent, all usually ciliolate to ciliate marginally and/or apically; 
petals 3-6(-9), usually white to rarely pink, fused at base, falling as a unit with 
stamens; stamens indefinite, filament white, adnate to corolla base, filiform, pubes- 
cent at base (trichomes filiform, reddish brown to dark deep red in color), anther 
yellow, bifurcate, versatile, extrorse, basally dehiscent by rimiform pores; ovary 
superior, usually globose, 3-7- but usually 5-loculed, sulcate, mostly glabrous; styles 
as many as locules, obsolete to exceeding androecium in height, free, but often 
slightly coherent at base (sometimes coherent throughout in Asiatic species), filiform 
to somewhat fleshy, ovules indefinite, anatropous, unitegmic, placentation axile. 
Fruits baccate, seeds numerous, small, somewhat globose to irregularly ellipsoid, 
areolate, embedded in a mucilaginous pulp; embryo straight, one-third to one-half as 
long as seed, endosperm copious, mealy. 

KEY TO SERIES 

1. Leaves and inflorescences distributed from tip of to low along branchlets/stems; 

shrubs I. Ser. Omichlophilae 

1. Leaves and inflorescences clustered behind or crowded around tip of branchlets; 

shrubs and trees. 

2. Ovary and fruit pubescent II. Ser. Gynotrichae 

2. Ovary and fruit glabrous. 

3. Leaves with tertiary veins immersed beneath, scarcely more prominent 

than lesser venation; plants usually glabrescent III. Ser. Laevigatae 

3. Leaves with tertiary veins elevated beneath, more prominent than lesser 
venation; plants pubescent. 

4. Plants mostly glabrescent to sparingly pubescent, flowers less than 13 
cm. broad at anthesis (except S. schultesiana ), sepals glabrous to partly 
and sparingly pubescent outside, glabrous inside, stamens less than 40 

IV. Ser. Parviflorae 

4. Plants sparingly to copiously pubescent, flowers vary in size, sepals pul- 
verulent or pubescent outside, partly to entirely pubescent inside (except 
S. tambensis and S. chaparensis), stamens over 40. 
5. Sepals pulverulent, trichomes shorter than 0.5 mm.; branched 
trichomes mostly predominate all over the plant. 

V. Ser. Pulverulentae 

5. Sepals pubescent, trichomes 0.75 mm. or longer; unbranched 
trichomes mostly predominate all over the plant. 

6. Leaves sparingly to abundantly pubescent beneath, but not so 
dense as to be velvety or wooly, lower epidermis not obscured by 
pubescence under lower power VI. Ser. Macrophyllae 

6. Leaves densely lanate beneath, velvety and soft to touch, lower 
epidermis obscured by pubescence under low power. 

VII. Ser. Lanatae 

I. Ser. OMICHLOPHILAE Soejarto, ser. nov. 

Brachitrichae Busc., Malpighia 25: 220, p. p. min. 
Gynogynae Busc., 1. c. 220, p. p. min. 
Stenobasicae Busc., 1. c. 220, p. p. min. 



SOE JARTO: SA URA VIA 2 1 

Frutices per folia et inflorescentiae infertis secus ramulos et caules distributis 
caractentur. 

1. Leaves glabrous to glabrescent, subsessile to very shortly petiolate, petioles 0.2-0.7 
cm. long, leaf base truncate 1. S. spinuligera 

1. Leaves sparingly to abundantly pubescent, distinctly petiolate, petioles 1 cm. or 
longer, leaf base cuneate. 

2. Inflorescence densely pubescent, leaves chartaceous, scabrous and opaque 
above, stellate trichomes abundant to scattered beneath 2. S. stapfiana 

2. Inflorescence sparingly pubescent, leaves coriaceous to subcoriaceous, glossy 
and smooth above, radiate trichomes scattered beneath. 

3. Flowers 20-30 mm. broad, leaves broadly elliptic to obovate with length/ 
width ratio of (1.2-)2(-2.3)/l, secondary veins (6-)8-12(-16) pairs 

3. S. omichlophila 

3. Flowers 12-23 mm. broad, leaves narrowly elliptic to obovate with length/ 
width ratio of (1.2-)3(-3.5)/l, secondary veins (9-)13-18(-22) pairs 

4. S. caquetensis 

1. Saurauia spinuligera R. E. Schultes, Caldasia 2: 44. 1943. 
Type: Pennell 10452 (GH, holotype). Figure 3. 

Shrubs, glabrescent to sparingly pubescent. Branchlets very slender, terete, gla- 
brous to sparingly aculeate-hirsute (spinuligerous); trichomes (spinules, cf. Schultes) 
if present yellowish to golden brown, base (mammae, dto.) grayish yellow, elliptic- 
subconoidal, becoming abruptly very slender, straight to flexuous toward apex, to 5 
mm. long. Leaves distributed from tip of to low along branchlets; blades oblong- 
elliptic, shortly and abruptly acuminate at apex with acumen to 5 mm. long, truncate 
to cordate at base, serrulate along margins, 6-12.5 cm. long, 2-4.5 cm. wide, coria- 
ceous in dry state, gray to dark gray-brown above, gray to gray-brown beneath, 
subglossy and smooth above, secondary veins immersed, scarcely more prominent 
than lesser venation, glabrous on both sides but with setulose to setose trichomes 
along midrib and secondary veins beneath; petioles 0.2-0.7 cm. long, 1-2.5 mm. in 
diameter, somewhat canaliculate, glabrous to scattered setose pubescent. Inflores- 
cences distributed from tip of to low along branchlets, ascending to pendulous, 5-13- 
flowered, 7.5-15 cm. long, 2-4 cm. wide, glabrous, primary penduncle very slender, to 
1.5 mm. in diameter, bracts linear, to 5 mm. long. Flowers 12.5-15 mm. broad, buds to 
4 mm. in diameter, pedicels 1.5-5 mm. long, bracteoles subulate, 1-3 mm. long; sepals 
5, outer 2 ovate to elliptic, acute to obtuse, 4-5 mm. long, 2.5-3 mm. wide, imbricate 
one and inner 2 suborbicular to broadly elliptic, obtuse, 5-6 mm. long, 4.5-5 mm. wide, 
all glabrous throughout, marginally entire to irregularly ciliolate; petals 5, white, 
suborbicular to obovate, obtuse to rounded, 8-9 mm. long, 7-8 mm. wide; stamens 
20-25, filament 1.5-2 mm. long, anther 1.5 mm. long; ovary 5-loculed, globose, 5- 
sulcate, glabrous, styles 5, to 4.5 mm. long, subcapitate. Berries 5-loculed, globose, to 
6 mm. (8-10 mm., cf. Schultes) across, 5-sulcate. 

Habitat. Mossy forest on spur (subparamo), at altitudes of 
2,800-3,300 m. 

Distribution. Colombia (Departments of Antioquia and Caldas). 



22 FIELDIANA: BOTANY 

Specimens examined. COLOMBIA, ANTIOQUIA: vie. Medellin, 
Tamesis, Feb., fl. fr., Toro 983 (MEDEL, NY). CALDAS: Cord. 
Occid., Cerro Tatama, fr., Pennell 10452 (GH). 

When this species was described, only fruiting material (Pennell 
10452) with exceedingly young flower buds was available. Pennell 
10452 has what Schultes called "spinules," hence the specific name, 
along the branchlets. These spinules become slender and the swol- 
len base decreasing in thickness higher up the branchlets. In con- 
trast, the flowering material represented by Toro 983 is devoid of 
any epidermal emergencies along the glabrous branchlets and 
petioles. It seems that the spinules may not be typical of this species. 
More characteristic are the oblong-elliptic leaves with truncate to 
cordate base, and the short or almost lacking petioles. In this re- 
spect, the species is unique among South American Saurauia. The 
glabrous sepals and the mode of leaf distribution from the tip of to 
low along the branchlets, together with the high altitudes at which 
the species grows, indicate close relationship to S. omichlophila, the 
reason for its inclusion in the Ser. Omichlophilae. 

2. Saurauia stapfiana Busc., Malpighia 27: 314. 1916. Type: 
Triana 267 (P, lectotype; G. K, isolectotypes; F, GH, photos). Figure 
4. 

Shrubs to 3(-5) m. tall, erect to rarely bushy, copiously pubescent. Branchlets 
slender, obtuse-angled to terete, densely to sparingly strigose, trichomes rusty brown, 
to 4 mm. long. Leaves distributed from tip of to low along branchlets; blades elliptic to 
obovate, acuminate at apex with acumen to 15 mm. long, cuneate to obtuse but rarely 
oblique at base, serrate to serrulate along margins, (8-)10-20(-23) cm. long, (3-)5- 
8(-10) cm. wide, subcoriaceous, dark green above, green with yellow to rose-colored 
veins beneath (in dry state dark brown to sooty above, rusty brown beneath), often 
rugulose and scabrous above, secondary veins (15-)17-23(-27) pairs, tertiary veins 
elevated, more prominent than lesser venation, sparingly strigillose pubescent 
(trichomes often reduced to warts) along secondary and minor veins and densely 
strigose to strigillose pubescent along midrib above, abundantly to sparingly pubes- 
cent, with trichomes of stellate to radiate types along and between veins and mixed 
with setose types along major veins beneath; petioles (0.5-)1.5-2(-3) cm. long, 1.5-2 
mm. in diameter, half-terete to furrowed above, densely to abundantly pubescent 
with trichomes of stellate and strigose types. Inflorescences distributed from tip of to 
low along branchlets, ascending to curving downward but rarely pendulous, (3-)5- 
15(-20)- flowered (lateral cymes often reduced to one flower), (l-)3-13(-20) cm. long, 
(0.7-) 2-4(-6) cm. wide, densely strigose to shaggy-strigose pubescent, primary pe- 
duncle slender, to 10 mm. long. Flowers 12.5-17.5 mm. broad, buds to 5 mm. across, 
pedicels 4 mm. long, bracteoles linear to triangular, to 4 mm. long; sepals 5, green to 
greenish white, ovate to orbicular-ovate, acute to obtuse but very rarely truncate, 
4.5-6 mm. long, 3.5-4.5 mm. wide, exposed parts in bud abundantly strigose to 
shaggy-strigose pubescent, imbricated parts glabrous, marginally ciliate to ciliolate, 



SOEJARTO: SA URA VIA 23 

all glabrous inside; petals 5, white, oblong-elliptic to obovate, rounded and often 
incised, 5-8 mm. long, 4-6 mm. wide, stamens 20-40, filament 2-2.5 mm. long, anther 
2-3 mm. long; ovary 5-loculed, globose, 5-sulcate, glabrous, styles 5, obsolete to 5 mm. 
long, stigmas simple to capitate. Berries green, subglobose, to 6 mm. across, 5- 
sulcate. 

Habitat. Rich soil in wet mountain forest, humid and open 
forest, subparamo (with low shrubs, ferns, and sphagnum), cloud- 
zone vegetation (of low trees, shrubs, and ferns), shady and semi- 
shady places, second-growth areas and open hillsides, at altitudes of 
1,700-3,200 m. 

Distribution. Colombia (Departments of Choco, Antioquia, 
Boyaca, Tolima, Valle and Cauca). 

Vernacular names. Lulumoco (Choco: Barkley & Molina). 

Saurauia stapfiana is characterized by the combination of the 
following features: leaves relatively small and predominantly obo- 
vate with stellate to radiate pubescence on the lower surface, dis- 
tributed from the tip of to low along the branchlets, inflorescences 
small, few-flowered, little- to non-branched, flowers with sepals 
glabrous inside and on the imbricated parts, and with a low to mod- 
erate number of stamens. Two varieties can be distinguished: var. 
stapfiana and var. radiata. 

1. Lower leaf pubescence stellate, trichomes 0.3-1.3 mm. broad ... a. var. stapfiana 
1. Lower leaf pubescence radiate, trichomes 0.2 mm. broad b. var. radiata 

a. Saurauia stapfiana var. stapfiana. 

Specimens examined. COLOMBIA, CHOCO: La Mansa, Jan., fl., 
Molina & Barkley 19-Ch-030 (F, K, US). ANTIOQUIA: Medellin, 
Dec., fl., Sandeman 5522 (COL, K); N. of Medellin, near San Pedro, 
Sept., fl., Lehmann 4031 (K, US); NW of Medellin, Camino Real 
Antiguo (Boqueron de San Cristobal), June, fr., Barkley & Correa 
123 (US); 1 km. S. of Hoyo Rico, Sept., fl. fr., Correa & Barkley 
L-80A-202 (US); road between Medellin and El Retire at vie. of Las 
Palmas, Oct., fl., Diaz 38 (F, MEDEL), Dec., fl., Falla 15 (F, 
MEDEL), May, fl. fr., Gutierrez et al. 119 (MEDEL, US), 120 (US), 
Aug., fl., Hatheway 1546 (B), Sept., fl., J. Uribe 257 (MEDEL); El 
Retire, fl., Daniel 3244 (MEDEL); La Ceja, Alto de Pantanillo, Dec., 
fl., Uribe-Uribe 4191 (COL, ECON); road Medellin-Dabeiba, April, 
fl., Romero-Castaneda 2443 (F, MEDEL). TOLIMA: Mariquita, fl., 
Triana 267 (G, K, P; GH, photo; with the original note: Prov. de 
Mariquita et Antioquia, forets du Quindio et de Antioquia). 



24 FIELDIANA: BOTANY 

b. Saurauia stapfiana var. radiata Soejarto, var. nov. 

Folia subtus distante ad sparse radiata, trichomatibus radiatis minus quam 0.2 
mm. in diametro, brachiis elongate conoideis minus quam 0.1 mm. longis. Typus: 
Soejarto & Rivera 2045 (GH, holotypus; ECON, MEDEL, isotypus). 

Specimens examined. COLOMBIA, ANTIOQUIA: road between 
Prado and Armenia, vie. of Las Partidas, Jan. fl., Uribe-Uribe 2888, 
fl. fr., 2889 (both in COL, ECON, GH); Sonson, below Paramo de 
Sonson, June, fl., Soejarto & Rivera 2045, 2046 (both in ECON, GH, 
MEDEL). BOYACA: Sierra Nevada del Cocuy, along path from 
Laguna Seca to Bachira, Aug., fl., Grubb et al 607 (US). VALLE: 
Cordillera Central, ridge, between Las Brisas and La Marina, 
Cuatrecases 22647 (ECON, F). CAUCA: Cordillera Central, western 
drainage of paramo de Purace, above Quebrada de Rio San Juan, 
Oct., fl., Cuatrecasas & Willard 26315 (US). 

3. Saurauia omichlophila R. E. Schultes, Caldasia 2: 319. 
1944. Type: Schultes 3236 (ECON, holotype; COL, GH, isotypes). 

Shrubs to 5 m. tall, erect to declining or straggling, rarely spreading, little to 
profusely branched; sparingly pubescent. Branchlets terete to obtusely angled, gla- 
brescent to abundantly pubescent, trichomes of hirsute to hirsute-strigose types, 
yellowish white to gray-brown or rose-colored. Leaves distributed from tip of to low 
along branchlets; blades obovate to elliptic, abruptly and shortly acuminate at apex, 
cuneate to broadly cuneate-rotundate at base, serrulate to setaceous-serrulate along 
margins, (2-)4-10(-15) cm. long, (l-)2-4(-6) cm. wide, coriaceous, sublaevigate to 
glossy above, dark green on both sides, secondary veins (6-)9-12(-14) pairs, tertiary 
veins immersed, scarcely more prominent than lesser venation, scattered pubescent 
with setose to hirsute trichomes along veins above, later glabrescent, scattered to 
sparingly sericeous-hirsute pubescent along and between veins, sometimes with scat- 
tered radiate trichomes beneath; petioles 0.5-2 cm. long, 0.75-1.5 mm. in diameter, 
sparingly to abundantly pubescent, trichomes of setose to sericeous hirsute types. 
Inflorescences distributed from tip of to low along branchlets and stems, often hori- 
zontal to pendulous, (3-)7-20(-35)-flowered, (5-)8-13(-18) cm. long, 2-5 cm. wide, 
abundantly setose to sericeous-hirsute pubescent, primary peduncle 4-8 cm. long, 
bracts linear, to 5 mm. long, rarely foliaceous, to 25 mm. long. Flowers 20-30 mm. 
broad, buds to 10 mm. in diameter, pedicels to 5(-6) mm. long, bracetoles linear to 
subulate, 3-5 mm. long; sepals 5-8, ovate to suborbicular, obtuse, 6-8 mm. long, 4-7 
mm. wide, exposed parts in bud sparingly to abundantly pubescent, trichomes of 
hirsute-strigose type, imbricated parts glabrous, all glabrous inside, marginally and 
apically ciliolate; petals 5-9, white to rose-colored, oblong to elliptic-oblong, obtuse to 
incised, (8-)10-15 mm. long, 5-10 mm. wide; stamens (10-) 20-30(-40), filaments 3-4 
mm. long, anther 2.5-3 mm. long; ovary 5-8-loculed, globose, 5-8-sulcate, glabrous, 
styles 5-8, obsolete to 4(-5.5) mm. long, stigmas simple to subcapitate. Berries green 
to purple, (4-)5-8-loculed, globose, to 10 mm. across, 5-8-sulcate. 

Habitat. Wet mountain forest, cloud forest, mist-zone, and sub- 
paramos, usually only in undisturbed, shady, and wet places on rich 
humus, at altitudes of 2,500-3,200 m. 



SOEJ ARTO: SA URA VIA 25 

Distribution. Colombia (Departments of Cauca, Narino, 
Putumayo). 

Vernacular names. Moquillo de paramo (Sibundoy Valley: 
Schultes), Dji-nu-sse (Sibundoy Valley, Kamsa Indian name: 
Schultes), Moquillo de paramo (Pasto: Soejarto). 

Specimens examined. COLOMBIA, CAUCA: between Valencia 
and San Sebastian, west side of the mountain, Core 1018, 1019 (both 
in NY, US); Cordillera Central, east facing slope, Cuatrecasas 23662 
(F, US); El Derrumbo Mountain, Killip 7986 (NY); Purace, vie. of 
Lake San Rafael, Uribe-Uribe 3876 (COL, ECON). NARINO: road to 
Mocoa, between El Encanto and Dolores, Garcia-Barriga et al. 
13023 (COL, MEDEL, US); above Lake La Cocha, Paramo de 
Tabano, Hernandez 79 (ECON, GH, PASTO); Ciudadela, La Cocha 
to Sibundoy, near Paramo de Bordoncillo, Schultes & Villarreal 
7560 (COL, ECON, F, GH, K, NY, US), 7560A (COL, ECON, GH, 
US); between Lake La Cocha and Paramo de Tabano, Schultes & 
Villarreal 7771 (COL, F, GH, K, US); Pasto, above "Bosque Botana" 
Exptl. Garden, Soejarto & Porter 478 (ECON, GH), Soejarto 1500, 
1501, 1509, 1511, 1517, 1598 (all in ECON, GH), 977 (COL, ECON, 
GH, PASTO), 1493 (ECON, F, GH, L, NY, P, PASTO, UC, US), 1497 
(ECON, GH, PASTO), 1502 (ECON), 1521 (ECON, F, G, GH, US). 
PUTUMAYO: between San Antonio de Bordoncillo and Santiago de 
Sibundoy, Cuatrecasas 11790 (COL, US); Paramo de San Antonio, 
between Lake La Cocha and Sibundoy, Schultes 3236 (ECON, 
holotype; COL, GH, isotypes); between La Maria and Paramo de San 
Antonio, Schultes & Villarreal 7550 (COL, F, GH, K, US); Sibundoy 
Valley, Garcia-Barriga 4563 (COL, US; F, GH, NY, photos), 
Schultes & Cabrera 20098 (GH); Paramo de Las Ovejas, between El 
Encanto and Sibundoy Valley, Soejarto 1051 (ECON, GH, PASTO); 
km. 40 east of Pasto, road to Sibundoy Valley, Soejarto 1176 (COL, 
F, G, GH, NY, PASTO, US), 1177 (ECON, GH, PASTO), 1178 
(ECON, GH). 

Saurauia omichlophila is abundantly represented in the moun- 
tain system of Narino and Putumayo, where the plants commonly 
grow in the subalpine and cloud-covered zone on rich black soil. The 
slender stem and branches are often declining and descending to the 
ground and the plants become bushy, or they can grow very long and 
slender and straggling among the dense shrub vegetation, such as in 
the mountain ridges above Pasto. The plants grow either individu- 
ally or in clumps, which are often produced vegetatively, since the 



26 FIELDIANA: BOTANY 

stems and branches which touch the ground produce adventive roots 
in the wet and frequently misty habitat. 

The indument of the shoot, young leaves, and inflorescences is 
often ochre-rosy to reddish or greenish brown in color, but turning 
paler to yellowish white when old. Most members of the species have 
hirsute to sericeous-hirsute trichomes, but Core's collection from 
Cauca has strigose to tuberculate-strigose indument (which is deep 
to reddish brown in color) over the branchlets, leaves, and sepals. 
The sepals are mostly green, but they may be purplish red or rose- 
colored. 

The irregular repetition of the floral parts in this species is worthy 
of note. Pentamery and hexamery are quite common, but hepta- and 
octamery are less frequent. Basically, however, the floral parts are 
pentamerous, and higher series are always found together with pen- 
tamery, in the same plant. 

4. Saurauia caquetensis R. E. Schultes, Caldasia 2: 32. 1943. 
Type: Cuatrecasas 8439 (US, holotype; COL, F, isotypes). Illustra- 
tion: Caldasia 2: 33. 1943. Figure 5. 

Shrubs to 4 m. tall, erect to bending, much branched but rarely bushy; sparingly 
pubescent. Branchlets slender, terete, sparingly pubescent, trichomes of shaggy- 
strigose type, gray to rusty brown in color. Leaves distributed from tip of to low along 
branchlets; blades narrowly elliptic to elongate-obovate, acuminate at apex with 
acumen to 15(-25) mm. long, cuneate to obtuse at base, rarely oblique, very rarely with 
basal flap, serrate to denticulate-serrate along margins, (5-)6-12(-17) cm. long, 
(1.5-)2-5 cm. wide, chartaceous to subcoriaceous, dark green above, light green be- 
neath, glossy and smooth above, secondary veins (9-)13-18(-22) pairs, tertiary veins 
immersed to slightly elevated, scarcely more prominent than lesser venation, glabrous 
to glabrescent with scattered strigose trichomes along major veins above, glabrescent, 
but major and minor veins with sparingly distributed shaggy-strigose and scattered 
radiate trichomes beneath; petioles 7-20 mm. long, 1-1.5 mm. in diameter, sparingly 
strigose. Inflorescences distributed from tip of to low along branchlets, erect to pendu- 
lous, 5-20(-30)-flowered, 5-16 cm. long, 1.5-4 cm. wide, pulverulent to abundantly 
pubescent, trichomes of strigose and stellate to radiate types, primary peduncle 
(1.5-)3-8 cm. long, bracts linear to subulate, to 10 mm. long. Flowers 12-22.5 mm. 
broad, buds 5-10 mm. in diameter, pedicels to 5 mm. long, bracteoles triangular, to 5 
mm. long; sepals 5, green, ovate to suborbicular, obtuse, 5-7 mm. long, 4-6 mm. wide, 
all glabrous throughout, marginally and apically entire to ciliolate, especially at apex; 
petals 5, white, obovate to broadly oblong, rarely spatulate, obtuse to slightly incised, 
6-10 mm. long, 4-7 mm. wide; stamens 20-35, filament 3-4 mm. long, anther 2-2.5 mm. 
long; ovary 5-loculed, subglobose, 5-sulcate, glabrous, styles 5, obsolete to 4(-5) mm. 
long, stigmas simple to subcapitate. Berries green, subglobose, to 5 mm. across, 
5-sulcate. 

Habitat. Cloud-covered and wet mountain forest on rich soil, at 
altitudes of 2,000-2,800 m. 



SOEJARTO-.SAURAUIA 27 

Distribution. Colombia (Departments of Huila, Cauca, 
Putumayo, Caqueta). 

Individuals of Saurauia caquetensis resemble those of <S. 
omichlophila in habit, but they may be distinguished by much nar- 
rower leaves (about 3:1 ratio of leaf length/width) in the former. The 
population in the Putumayo region is characterized by the usually 
erect to ascending, many-flowered, and many-branched inflores- 
cences with small flowers, in contrast to the pendulous, few flowered, 
and non-branched inflorescences with large flowers, represented by 
Cuatrecasas 8439 collected from Caqueta. Two varieties can be dis- 
tinguished: var. caquetensis and var. parviflora. 

1. Inflorescences pendulous, unbranched, 5-10-flowered, flowers ca. 20 mm. broad 

var. caquetensis 

I. Inflorescences erect to ascending, many-branched, 13-30-flowered, flowers ca. 13 

mm. broad var. parviflora 

a. Saurauia caquetensis var. caquetensis. 

Specimens examined. COLOMBIA, CAQUETA: Cordillera 
Oriental, along ridge at Gabinete, March, fl., Cuatrecasas 8439 (US, 
holotype: COL, F, isotypes). 

b. Saurauia caquetensis var. parviflora Soejarto, var. nov. 

Var. caquetensis differt inflorescentiis erectioribus ad ascendentibus, floribus 
13-20(-30) in inflorescentiis, ca. 12.5 mm. in diametro, staminibus 30-35. Typus: 
Cuatrecasas 8532 (US, holotype; GH, NY, photos). 

Specimens examined. COLOMBIA, CAUCA-HUILA: Moscopan, 
Sept., fl., Uribe-Uribe 3861 (COL, ECON). PUTUMAYO: km. 40 
east of Pasto, Sta. Clara, July, fl., Soejarto 1052, 1175 (both in COL, 
ECON, GH, PASTO), 1050, 1513, 1514, 1515, 1516, 1519, 1520, 
1524 (all in ECON, GH); Cerro Portachuelo, road San Francisco of 
Sibundoy to Mocoa, various sites, May, fl., Schultes & Villarreal 
7766 (ECON), July, veg., Soejarto 1130 (COL, GH, PASTO), 1151 
(GH, PASTO), fl., 1148, 1168 (ECON, GH, PASTO), Aug., fl., 1540, 
1541, 1549, 1550 (all in ECON, GH), fr., 1560 (ECON, GH). 

II. Ser. GYNOTRICHAE Busc., Malpighia 25: 220. 1912. 

Brachitrichae Busc., 1. c., p. p. 

5. Saurauia loesneriana Busc., Malpighia 25: 399, pi. 12, fig. 28. 
1913. Type: Jelski 259 (GH, NY, photos). 

Shrubs to 3.5 m. tall, erect; copiously pubescent. Branchlets sulcate to terete, 
abundantly pubescent, trichomes of setose-hirsute to sericeous types, light brown, 
swollen at base and abruptly turning very slender and flexuous towards apex, to 5 
mm. long. Leaves crowded behind tip of branchlets, blades elongate-elliptic to obo- 



28 FIELDIANA: BOTANY 

vate, acute to shortly acuminate at apex with acumen to 10 mm. long, cuneate at 
base, serrulate along margins, 15-25 cm. long, 4.5-7 cm. wide, chartaceous, dark 
brown to black above in dry state, dark olive-brown beneath, subglossy and smooth 
above, secondary veins 23-30 pairs, tertiary veins somewhat elevated, scarcely more 
prominent than lesser venation, sparingly pubescent above with trichomes of setose 
to sericeous or strigillose types along veins, abundantly to sparingly pubescent be- 
neath with trichomes of hirsute to sericeous types along major veins, but of radiate to 
clustered types along and between minor veins; petioles 1.5-3 cm. long, 1.5-2.5 mm. in 
diameter, sparingly pubescent, with setose-hirsute trichomes. Inflorescences 
straight, 6-15-flowered, 5-10.5 cm. long, 2-3 cm. wide, densely strigose to sericeous 
pubescent, primary peduncle stout, 2-4.5 cm. long, bracts linear, to 10 mm. long. 
Flowers subsessile, 20 mm. broad, buds to 8 mm. in diameter, bracteoles triangular, 
to 5 mm. long; sepals 5, pale green with rusty brown trichomes, outer 2 ovate to 
elliptic, 7-8 mm. long, 4-5 mm. wide, subacute, imbricate one broadly ovate to subor- 
bicular, 7-8 mm. long, 6-6.5 mm. wide, obtuse, inner 2 broadly ovate to suborbicular, 
7-8.5 mm. long, 6-7 mm. wide, rounded, exposed parts in bud densely pubescent with 
trichomes of sericeous type, imbricated parts pulverulent with trichomes of stellate 
type, marginally and apically ciliolate, all densely pulverulent with appressed- 
stellate trichomes inside; petals 5, white, oblong to obovate, gnawed to entire along 
margins, praemorse to rarely incised at apex, 10-12 mm. long, 5-7 mm. wide; stamens 
30-50, filament 2.75-3.25 mm. long, anther 2.25-3 mm. long; ovary 5-loculed, globose, 
5-sulcate, sparingly to abundantly pubescent with trichomes of appressed-stellate 
type, styles 5, to 5 mm. long, stigmas capitate. Berries known only in immature state, 
mostly sparingly pubescent on upper portion, glabrescent on lower half. 

Habitat Shrubby land, clay soil, and sunlit area at altitudes of 
2,800-3,000 m. 

Distribution. Peru (Departments of Piura and Cajamarca). 

Specimens examined. PERU, PIURA: Huancabamba, 
Pacaipampa, Tambillo, fl., Jelski 259 (GH, NY, photos of type). 
CAJAMARCA: Cutervo, Arenales, 1 hr. S. of St. Thomas, Dec., fl., 
Stork & Norton 10161 (F, G, K, UC). SAN MARTIN: Huallaga, 
valley of Rio Apisoncho, 30 km. above Jucusbamba, July, fl., 
Hamilton & Holligan 1020 (K). 

The above description has been based primarily upon Stork & 
Horton 10161, which compares extremely well with the type photo- 
graph (Jelski 259) . Both Buscalioni and Macbride (1956) described 
the ovary as glabrous, but careful examination of the floral parts of 
Stork & Horton collection revealed that the ovary is pubescent 
throughout or only on the upper portion. This is an important 
character, because it is the only South American species with pubes- 
cent ovary. 

HI. Ser. LAEVIGATAE Busc., Malpighia 25: 224. 1912; Hunter, 
Ann. Missouri Bot. Gard. 53: 80. 1966. 



SOEJ ARTO: SA URA VIA 29 

Oligotrichae Busc., Malpighia 25: 224, p. p. 
Scabrae Busc., 1. c., p. p. min. 

1. Sepals glabrous to sparingly pubescent outside, glabrous to partly pubescent (usu- 
ally on upper half) inside. 

2. Leaves lanceolate to oblanceolate, length to width ratio of 3 or more to 1. 

3. Blade of leaves small, 3-12 cm. long, 1-3.5 cm. wide, villous-barbate at axils 
of secondary and often tertiary veins beneath; Peru 6. S. briqueti 

3. Blade of leaves large, 10-30 cm. long, 3-10 cm. wide, not villous-barbate 
beneath; Colombia l.S. portachuelensis 

2. Leaves elliptic to obovate, length to width ratio of 2 or less to 1. 

4. Inflorescences and floral parts glabrous throughout; Bolivia . . 8. S. rusbyi 
4. Inflorescences abundantly to sparingly pubescent; Colombia to Peru. 

5. Villous-barbate at axils of secondary veins beneath, stamens 50-80, 
anther less than 0.85 mm. long; Peru d. S. glabra 

5. Villous-barbate pubescence absent beneath, stamens less than 50, 
anther 1 mm. or longer. 

6. Leaves glabrous except for scattered trichomes along major veins be- 
neath, inflorescences 40- to more than 150-flowered, flowers less than 
10 mm. broad, sepals partly pubescent inside, anther 1-1.5 mm. long, 

ovary mostly 4-loculed; Colombia (to Mexico) 10. S. yasicae 

6. Leaves radiate pubescent beneath, with strigose trichomes along 
midrib, inflorescences 4-30-flowered, flowers 10-15 mm. broad, sepals 
glabrous inside, anther 2.75-3.25 mm. long, ovary 5-loculed; Ecuador 

11. S. aequatoriensis 
1. Sepals abundantly pubescent throughout or inside only. 

7. Inflorescences 100-200 flowered, flowers less than 10 mm. broad, buds ellipsoi- 
dal, ca. 3.5 mm. long; Ecuador 12. S. adenodonta 

7. Inflorescences less-than-100-flowered, flowers ca. 15 mm. broad, buds globose, 
to 6 mm. in diameter; Colombia. 

8. Leaves glabrescent, inflorescences 3-20-flowered, sepals heterotrichous out- 
side, trichomes of strigillose and stellate types 13. S. strigillosa 

8. Leaves glabrous, inflorescences 30-100-flowered, sepals homotrichous out- 
side, trichomes of stellate type 14. S. laevigata 

6. Saurauia briqueti Busc., Malpighia 30: 140, pi. 12, fig. 26. 
1927. Type: Weberbauer2177 (F, leaf fragment, isotype; G, GH, NY, 
US, photos of holotype?). 

Shrubs to 4 m. tall; glabrescent. Branchlets slender, terete, reddish maroon, 
smooth, glabrous to glabrescent, younger parts sparingly pubescent, trichomes of 
strigillose to tuberculate-strigillose types. Leaves clustered behind tip of branchlets; 
blades lanceolate to narrowly elongate-obovate, acute to very shortly acuminate at 
apex, cuneate at base, serrulate along margins, 3-12 cm. long, 1-3.5 cm. wide, sub- 
coriaceous, glossy and smooth above, secondary veins 12-16 pairs, tertiary veins 
immersed, scarcely more prominent than lesser venation, mostly glabrous through- 
out, except for scattered strigillose trichomes along major veins, often villous-barbate 



30 FIELDIANA: BOTANY 

at axils of secondary veins beneath; petioles 0.75-2 cm. long, 1-1.5 mm. in diameter, 
glabrescent. Inflorescences straight, moderately branched, 25-100-flowered, 4-11.5 
cm. long, 2.5-8 cm. wide, glabrescent to sparingly strigillose pubescent, often with 
minute stellate trichomes along lesser ramification, primary peduncle 2-5 cm. long, 
bracts linear, to 3 mm. long. Flowers 8-10 mm. broad (buds not available), pedicels 
1-2 mm. long, bracteoles triangular, to 1.5 mm. long; sepals 5, oblong to spatulate, 
subacute to rounded, 3.5-5 mm. long, 2-3 mm. wide, all glabrous throughout, margin- 
ally and apically ciliolate; petals 5, orbicular-oblong, rounded, 4-5 mm. long, 2.5-3 
mm. wide; stamens 20-30, filament 1.5 mm. long, anther 1 mm. long; ovary 3-5- 
loculed, ovoid to globose, 3-5-sulcate, glabrous, styles 3-5, 2.25 mm. long, stigmas 
subcapitate. Berries immature. 

Habitat. Altitudes of 1,800-2,400 m. 
Distribution . Peru (Department of Junin). 

Specimens examined. PERU, JUNIN: Tarma, Huacapistana, 
Cerro Sincapata, Oct., fl., Velarde-Nunez 749 (US, topotype), fl., 
Weberbauer2177 (F, leaf fragment, isotype; G, GH, NY, US, photos 
of holotype?). 

The above description has been based primarily upon Velarde- 
Nunez 749, a topotype. The outstanding features which separate S. 
briqueti from other Peruvian species are its small, narrowly 
elongate-obovate leaves, with a length to width ratio of 3-5:1, and a 
high degree of glabrescence. 

7. Saurauia portachuelensis R. E. Schultes, Caldasia 2:40. 
1943. Type: Miguel de Ipiales 40 (ECON, holotype; F, isotype). Il- 
lustration: Caldasia 2:41. 1943. 

Small trees to 5 m. tall, trunk crooked to straight, crown open; glabrescent to 
sparingly pubescent. Branchlets stout, terete, glabrescent to sparingly hirsute pubes- 
cent, trichomes light to dark silky brown. Leaves clustered behind tip of branchlets; 
blades oblanceolate to elongate-elliptic, cuspidate to acuminate at apex with acumen 
to 15 mm. long, cuneate to obtuse at base, serrate to serrulate along margins, 10- 
20(-30) cm. long, 3-6C-10) cm. wide, strongly coriaceous, dark green above, light to 
dull green beneath, glossy and smooth above, secondary veins (10-)15-25(-30) pairs, 
usually depressed above, tertiary veins immersed, scarcely more prominent than 
lesser venation, glabrous throughout except for setose to appressed-setose trichomes 
along midrib above and along midrib and secondary veins beneath; petioles 0.75-2 
cm. long, 2-3 mm. in diameter, sparingly setose pubescent. Inflorescences erect, 
straight, 50-100(-150)-flowered, (10-)14-20(-25) cm. long, 3-6 cm. wide, glabrous on 
lower parts, pulverulent with appressed strigose mixed with minute stellate 
trichomes on upper parts, primary peduncle 5-10 cm. long, bracts linear-subulate, to 
9 mm. long, glabrous to sparingly strigose pubescent. Flowers 7.5-12.5 mm. broad, 
buds to 4 mm. in diameter, bracteoles triangular, to 3 mm. long; sepals 5, green, outer 
2 and imbricate one ovate to elliptic, obtuse to subacute, 3-3.5 mm. long, 1.5-2 mm. 
wide, inner 2 obovate to orbicular-obovate, 3-4 mm. long, 2-3 mm. wide, all glabrous 
throughout, marginally ciliolate (fimbrilliferous, see Schultes); petals 5, elliptic to 



SOEJARTO: SA URA VIA 3 1 

orbicular-obovate, rounded, sometimes incised, 5-7 mm. long, 3-5 mm. wide; stamens 
15-30, filament 2-2.5 mm. long, anther 2-3 mm. long; ovary 5-loculed, subglobose to 
ellipsoid, 5-sulculate, glabrous, styles 5, obsolete to 3.5 mm. long, stigmas simple to 
subcapitate. Berries dark green, 5-loculed, globose, to 7 mm. across, 5-sulcate. 

Habitat. Wet mountain forest, subparamo, and cloud-covered 
zone at altitudes of 2,000-3,000 m. 

Distribution . Colombia (Department of Putumayo). 

Vernacular names. Moquillo de paramo (Sibundoy: Miguel de 
Ipiales). 

Specimens examined. COLOMBIA, PUTUMAYO: Upper 
reaches of Putumayo River in the Sibundoy Valley, along mountain 
ridge, La Cabana, Jan., fl., Cuatrecasas 11643 (COL, F, US); horse 
trail from San Francisco of Sibundoy to Mocoa, Jan., fl., Miguel de 
Ipiales 40 (ECON, F); road Sibundoy to El Pepino of Mocoa, Cerro 
Portachuelo, Aug., fl., Soejarto 521, fr., 522, July, fl., 1149, fr., 1150, 
fl., 1158, fr., 1167, Aug., fr., 1539, veg., 1542, fl., 1574, (all in 
ECON, GH); above Sibundoy Valley, road to Pasto, La Chorrera, 
Aug., fr., Soejarto 1528 (ECON, COL). 

Individuals of S. portachuelensis are easy to recognize in the field 
from their strongly coriaceous, glabrous, and elongate leaves which 
are shiny above, and the glabrescent, many-flowered inflorescences 
with small white flowers, the glabrous sepals, and the low number of 
stamens. The species is abundantly represented in the wet moun- 
tain forests along Cerro Portachuelo, in particular between the 
Sibundoy Valley and Mocoa. They may be found on black rich soil, 
rocky soil, stream bank, rocky cliff with thick soil, or along gullies, 
but not in open or exposed places. 

8. Saurauia rusbyi Britt., Bull. Torrey Bot. Club 16:64. 1889. 
Type: Rusby 482 (MICH, lectotype; NY, isolectotype). Figure 6. 

Shrubs?; sparingly pubescent. Branchlets slender, terete, glabrescent to sparingly 
hirsute pubescent. Leaves clustered behind tip of branchlets; blades obovate, shortly 
acuminate at apex with acumen to 10 mm. long, cuneate at base, serrate (aristate- 
serrate, see Britten) along margins, 10-20 cm. long, (3-)5-8 cm. wide, chartaceous, in 
dry state sooty above, olive-brown beneath, smooth and glossy above, secondary veins 
10-14 pairs, tertiary veins immersed, scarcely more prominent than lesser venation, 
sparingly strigillose between and along veins and abundantly sericeous pubescent 
along midrib above, sparingly hirsute pubescent along major veins beneath; petioles 
1-2 cm. long, 1-2 mm. in diameter, glabrescent to sparingly hirsute pubescent. In- 
florescences slender, 15-30-flowered, 8-15 cm. long, 4-7 cm. wide, mostly glabrous 
except for ciliate to fimbriate bracts and bracteoles, primary peduncle very slender 
(1-1.5 mm. in diameter), 4.5-8 cm. long, bracts subulate to triangular, to 4 mm. long. 



32 FIELDIANA: BOTANY 

Flowers 10-15 mm. broad, buds to 3.5 mm. in diameter, pedicels extremely slender, 
7.5-15 mm. long, bracteoles subulate, to 1.5 mm. long; sepals 5, broadly elliptic to 
suborbicular, rounded, 3-4 mm. long, 3-3.5 mm. wide, all glabrous throughout, mar- 
ginally ciliolate; petals oblong-obovate, rounded to incised, 5-8 mm. long, 4-6.5 mm. 
wide; stamens 30-40, filament 2.5 mm. long, anther 1-1.25 mm. long; ovary 5-loculed, 
subglobose, 5-sulcate, styles obsolete to 3.5 mm. long, stigmas simple to subcapitate. 
Berries unknown. 

Habitat. Altitudes of 1,500-1,800 m. 
Distribution. Bolivia (Province of La Paz). 

Specimens examined. BOLIVIA, LA PAZ: Mapiri?, fl., Rusby 
481B (GH, US); near Yungas, fl., Rusby 482 (MICH, lectotype; NY, 
isolectotype). 

It appears that, because of mislabeling, different collections from 
different plants have been given the same number: Rusby 481 . After 
careful examination, it was possible to separate the material into 
three groups of specimens, which represent three separate collec- 
tions: Rusby 481, 481 A, 481B. The specimens referred to Rusby 481 
(F, G, NY, US; collected at Mapiri, 1,500 m., April, fl.) and Rusby 
481 A (MICH, NY, US; collected at Yungas, 1,800 m., fl) belong to 
Saurauia spectabilis, whereas those referred to Rusby 481B (GH, 
US; collected at Mapiri?, 1,500 m., fl.) belong to S. rusbyi. The US 
collection of Rusby 481B was originally labeled Rusby 482, but 
floral characters reveal that this specimen belongs to a functionally 
staminate plant with obsolete styles. In contrast, the true Rusby 482 
belongs to a functionally pistillate plant with long styles. 

S. rusbyi differs from S. spectabilis in its glabrous, few-flowered 
inflorescences with laxly distributed, long- and filamentous- 
pedicelled flowers, glabrous pedicels and sepals. In addition, S. rus- 
byi has a lower stamen number (30-40) than S. spectabilis (45-85). 

9. Saurauia glabra (R. & P.) Soejarto, comb. nov. Basionym: 
Palaua glabra R. & P., Syst. Veg. Fl. Peruv. et Chilens. 182. 
1798. Type: Pavon s.n. (G, lectotype; GH, photo). Figure 8. 

Apatelia glabrata DC., Mem. Soc. Phys. Geneve 1: 428. 1822; 
Prodr. 1: 526. 1824. 

Sauravia glabrata Steud., Nomencl. Bot. ed. 2, 2: 516, nom. nud. 
(excl. syn. Sauravia serrata DC.). 

Saurauja glabrata Choisy, Mem. Soc. Phys. Geneve 14: 117. 1855. 

Saurauia aequatoriensis Sprague var. boliviana Busc., Malpighia 
30: 35. 1927, p. p. 



SOEJARTO: SA URA UIA 33 

Shrubs or trees to 15 m. tall; glabrescent. Branchlets slender, terete, glabrescent to 
sparingly strigillose pubescent, smooth. Leaves clustered behind tip of branchlets; 
blades elliptic to elliptic-obovate, acute to very shortly acuminate at apex, cuneate to 
obtuse at base, serrulate along margins, 10-18 cm. long, 5-8 cm. wide, chartaceous, 
smooth and glossy above, secondary veins 10-15 pairs, tertiary veins immersed, 
scarcely more prominent than lesser venation, scattered strigillose pubescent to gla- 
brescent along and between veins above and beneath, villous-barbate at axils of 
secondary veins beneath; petioles 1-2.5 cm. long, 1-1.5 mm. in diameter, glabrescent. 
Inflorescences somewhat straight, (15-)25-50(-80)-flowered, 5-13 cm. long, 2.5-6 cm. 
wide, abundantly to sparingly pubescent, trichomes of stellate (especially along lesser 
ramification) mixed with tuberculate-strigillose types, primary peduncle 2-8 cm. long, 
bracts linear, to 5 mm. long, rarely foliaceous, to 25 mm. long. Flowers 10 mm. broad, 
buds to 3 mm. in diameter, pedicels to 5 mm. long, bracteoles linear, to 3 mm. long; 
sepals 5, elliptic to orbicular-obovate, obtuse to rounded, 2.5-4 mm. long, 2-2.5 mm. 
wide, exposed parts in bud pulverulent with trichomes of appressed-stellate mixed 
with shaggy -strigillose types, imbricated parts glabrous, all glabrous inside, margi- 
nally irregularly ciliolate, especially along apex; petals 5, obovate to orbicular- 
obovate, rounded and often deeply incised, 5-6.5 mm. long, 3.5-5 mm. wide; stamens 
50-80, filament 2 mm. long, anther 0.5-0.85 mm. long; ovary 4-5-loculed, ovoid, 4.5- 
sulcate, glabrous, styles 4-5, 1-1.5 mm. long, stigmas subcapitate. Berries unknown. 

Distribution. Peru (Department of Huanuco). 

Specimens examined. PERU, HUANUCO: Mima (?), fl.,Pavon s. 
n. (G, GH, photos), fl., Pavon s. n. (F, fragment; G, P), veg., Pavon s. 
n. (F, leaf fragment); Chinchao, July, fl., Sawada 97 (F); no loc., fl., 
Pavon s. n. (Fl), ? Pavon s. n. (BM). 

For technical reasons, it has been necessary to propose the above 
new combination. Were Palaua glabra R. & P. synonymous with 
Saurauia serrata DC. (1822), the correct name for this taxon must 
be S. serrata DC., as it was treated by Sprengel (1827), not S. gla- 
brata Steud. (1841). However, S. serrata DC., a Mexican species, is 
taxonomically distinct from P. glabra R. & P. Since S. glabrata 
Steud. is a nomen nudum, while S. glabrata Choisy is a later 
homonym, they must be rejected according to the Code. 

10. Saurauia yasicae Loes., Bot. Jahrb. 23: 125. 1896; Hunter, 

Ann. Missouri Bot. Gard. 58: 84. 1966, pro syn. Type: Rotschuch 

246 (not seen). Illustration: Ann. Missouri Bot. Gard. 58: 85. 1966. 

Saurauja herbert-smithii Rusby, Descr. 300 New Sp. S. Amer. PL 

57. 1920. Type: H. H. Smith 857 (A, BM, F, G, GH, K, L, 

MICH, NY, P, S, US). 

Saurauia leucocarpa Schlecht. var. smithiana Busc., Malpighia 
29: 232. 1922. Type: Turckheim 1445 (not seen). 



34 FIELDIANA: BOTANY 

Saurauia yasicae var. laevigatae Busc., 1. c. 232. Type: Tonduz 
11453 (G, L). 

Saurauia yasicae var. laevigatae fina. veranii Busc., 1. c. 413. 
Type: Tonduz 13147 (G, P). 

Saurauia smithiana Busc., 1. c. 445. pi. 3, fig. 1. Type: H. H. 

Smith 857. 
Saurauia pseudopittieri Busc., 1. c. 30: 97. 1927. Type: Pittier 

11247 (not seen). 

Saurauia zetekiana Standl., J. Arnold Arbor. 11: 124. 1930. 
Type: Bangham 578 (G, fragment). 

Saurauia belizensis Lundell, Field & Lab. 13: 7. 1945. Type: 
Gentle 4439 (not seen). 

Trees to 30 m. tall, diameter to 30 cm. at base; glabrescent to sparingly pubescent. 
Branchlets slender, terete, glabrescent to scattered pubescent, trichomes tuberculate. 
Leaves crowded behind tip of branchlets; blades obovate to elliptic, shortly acuminate 
at apex with acumen to 10 mm. long, cuneate to subattenuate at base, serrulate to 
serrate along margins, 6-22 cm. long, 2-10 cm. wide, membranaceous to rarely char- 
taceous, in dry state dark brown to sooty above, olive to olive-brown beneath, some- 
what glossy and smooth above, secondary veins 5-15(-18) pairs, tertiary veins im- 
mersed, scarcely more prominent than to indistinguishable from lesser venation, 
glabrescent above, scattered pubescent with trichomes of clustered and tufted to 
setose-tufted types along major veins beneath, lower epidermis glabrous and some- 
what pustulate; petioles (0.5-)l-3 cm. long, (0.5- )2. 5 mm. in diameter, terete, glabres- 
cent. Inflorescences straight and often spreading, (20-HO-150-flowered, (3-)5-20 cm. 
long, (1.5-)3-10 cm. wide, glabrescent to abundantly stellate to tufted pubescent (par- 
ticularly along lesser ramification and pedicels), primary peduncle (1.5-)3-10 cm. 
long, bracts subulate, to 5 mm. long, rarely foliaceous, to 35 mm. long. Flowers 7-10 
mm. broad, buds to 3 mm. in diameter, pedicels 1-6 mm. long; sepals (3-H(-5), 
oblong-ovate to suborbicular, obtuse to rounded, 2-3 mm. long, 1.5-3 mm. wide, ex- 
posed parts in bud abundantly stellate pubescent, imbricated parts glabrous, all 
glabrous medially inside, but stellate pubescent laterally and on upper portion, mar- 
ginally ciliolate; petals (3-H(-5), white, oblong-obovate to obovate, rounded, 3-4.5 
mm. long, 2-4 mm. wide; stamens (18-)20-30(-35), filament 1.5-2.5 mm. long, anther 
1-1.5 mm. long; ovary (3-H(-5Moculed, globose, (3-H(-5)-sulcate, glabrous, styles 
(3-H(-5), obsolete to 4 mm. long, stigmas simple to capitate. Berries green to dirty 
white, (3-H(-5)-loculed, subglobose, 2-3 mm. across, 5-sulculate. 

Habitat. Virgin and deep forest, advanced secondary forest, 
wooded valley, in coffee plantation with original forest trees, thicket 
along stream, forested riverbank, hilltops and slopes, roadside, 
shady as well as sunlit areas, at altitudes of 30-1,300 m. 

Distribution. Mexico (Chiapas, Oaxaca, Tabasco), British Hon- 
duras (Cayo, Stann Creek, Toledo), Guatemala (Alta Vera-paz, 
Quezaltenango, Retalhuleu, Suchitepequez), Honduras (Atlantida, 



SOEJARTO: SA URA VIA 35 

Cortes, Santa Barbara, Yoro), Nicaragua (Matagalpa), Costa Rica 
(Alajuela, Cartago, Limon, Pantarenas, Tilaran), Panama (Canal 
Zone, Panama), Venezuela (Merida, Barquisimeto), Colombia (An- 
tioquia, Magdalena, Cundinamarca, Tolima). 

Vernacular names .-^Jahoncillo, Wild orange (British Honduras: 
Gentle), Chulindron (Honduras: Von Hagen). 

Specimens examined. BRITISH HONDURAS, CAYO DISTR.: 
Humming Bird Hwy., 31-mile section, Nov., fr., Gentle 8907, 8923 
(both in G). STANN CREEK DISTR.: Humming Bird Hwy., July, 
fr., Gentle 8254 (G). 

GUATEMALA, QUEZALTENANGO: Colomba, Sept., fl., Skutch 
1335 (G). 

COSTA RICA: forests of Tuis, Nov., fl., Tonduz 11453 (G, L); Las 
Vueltas, Tecurrique, Jan., fl., Tonduz 13147 (G, P). 

PANAMA, COCLE: Anton Valley, vie. of Finca Tomas Arias, 
Aug., fl., Allen 3630 (G, K, U), N. of Anton Valley, Sept., fl., Allen 
3706 (G). No locality, July, fl., Aviles 999 (G), Sept., fr., Bangham 
578 (G). 

VENEZUELA, MERIDA: El Vigia to Cano Amarillo, Feb., fl., 
Bernardi 1905 (Fl, NY). BARQUISIMETO: S. of Barquisimeto, 
Cabudare, Serranias of Terepalma, Lara, Aug., fl., Saer 637 (VEN). 

COLOMBIA, MAGDALENA: Sierra Nevada of Santa Marta, Cin- 
cinnati region, Cerro Quemado, fl., Espina & Giacometo A55 (F), 
Bellavista region, fl., A70 (F); trailside near San Andres, NW. slope, 
June, fl., Kernan 132 (NY, P); Santa Marta, ? Cacagualito, June, fl., 
Smith 857 (A, BM, F, G, GH, K, L, MICH, NY, P, S, US), March, fl., 
1774 (A, BM, F, G, GH, L, NY, P, S, U, UC, US). ANTIOQUIA: 
Anori, Providencia, between the hydroelectric plant and Dos Bocas, 
July, fl. fr., Soejarto 2847, 2934, 2941 (all at COL, F, UNIV. OF 
ANTIOQUIA*), May, fl., 4030, 4031, 4032, 4140, 4142-4146, 4149 
(all at UNIV. OF ANTIOQUIA), fl. fr., 4148 (UNIV. OF AN- 
TIOQUIA); no precise locality, Nov., fl., Espinal 769 (ECON, UV). 
CUNDINAMARCA: Pacho, La Palma Highway, Aug., fl., Haught 
6081 (US); about 75 km. by highway NW. of Bogota, at bridge across 
Rio Negro on highway between Pacho and La Palma, March, fl., 
Little, Jr. 7370 (NY, US). TOLIMA: Mariquita, road to Fresno, July, 
fl., Uribe-Uribe 3000 (COL, ECON, GH). NO PRECISE LOCALITY: 
Nueva Espana, fl., Pavon s. n. (G). 

Department of Biology, Univ. of Antioquia, Medellin, Colombia. 



36 FIELDIANA: BOTANY 

Although Hunter (1966) treated S. yasicae as a synonym of S. 
laevigata, these two species are treated here as two separate taxa, on 
the basis of leaf and floral characters. For further details, please 
refer to S. laevigata. 

11. Saurauia aequatoriensis Sprague, Trans. Proc. Bot. Soc. 
Edinburgh 22: 426. 1904. Type: Spruce 4989 (K, lectotype; F, G, 
GH, NY, P, S, iso-lectotypes). Figure 7. 

Saurauia aequatoriensis var. glabrata Busc., Malpighia 30: 39. 
1927, p. p. Type: Spruce 4989. 

Saurauia aequatoriensis var. glabrata fina. veranii Busc., 1. c. 30: 
39. 1927. Type: Spruce 4989. 

Saurauia aequatoriensis var. glabrata fina. longepetiolata Busc., 
1. c. 30: 40. 1927. Type: Sodiro 151, ex descr. 

Saurauia rhamnifolia Killip, J. Washington Acad. Sci. 16: 572. 
1926. Type: Rose & Rose 22377 (NY, US). 

Shrubs to small trees to 8 m. tall, trunk straight, bark fissured, red-brown, wood 
reddish white, branches spreading, rarely squarrose; glabrescent. Branchlets slender, 
terete, often smooth, glabrescent to scattered strigose to strigillose pubescent. Leaves 
crowded behind tip of to distributed somewhat low along branchlets; blades elliptic to 
obovate or oblong-obovate, acuminate to rarely cuspidate at apex with acumen to 10 
mm. long, cuneate to rarely obtuse at base, distantly serrate with very fine serrations 
along margins, (7-)10-15(-20) cm. long, 3.5-8 cm. wide, subcoriaceous, green on both 
sides but slightly darker above (in dry state sooty above, dull brown beneath), glossy 
and smooth above, secondary veins 11-2CK-24) pairs, tertiary veins irregular, some- 
what immersed, scarcely more prominent than lesser venation, mostly glabrous except 
for scattered strigillose to minutely tuberculate trichomes along major veins above and 
scattered radiate to clustered and strigillose trichomes along major veins beneath; 
petioles 1.5-2.5 cm. long, 1.5 mm. in diameter, sparingly pubescent, trichomes of 
strigose to strigillose types. Inflorescences often erect, very little-branched, (4-)10-30- 
flowered, 4-15 cm. long, 2-4 cm. wide, scattered strigose to strigillose pubescent, 
primary peduncle 1.5-5 cm. long, bracts triangular to linear, to 10 mm. long. Flowers 
10-15 mm. broad, buds ellipsoidal to subglobose, to 3.5 mm. in diameter, pedicels 1-3 
mm. long, bracteoles triangular, to 3 mm. long; sepals 5, green, ovate to elliptic to 
obovate, subacute to obtuse, 6-7.5 mm. long, 4-5.5 mm. wide, exposed parts in bud 
glabrous to scattered pubescent, trichomes of strigillose and radiate-stellate types, 
imbricated parts glabrous to scattered pubescent, trichomes of radiate-stellate types, 
all glabrous inside, marginally and apically ciliolate; petals 5(-6), white, elliptic to 
oblong, retuse to rounded, 7-8 mm. long, 4-5 mm. wide; stamens 20-40, filament 3.5-4 
mm. long, anther 2.75-3.25 mm. long; ovary 5-loculed, ovoid, 5-sulcate, glabrous, styles 
5, obsolete to 5 mm. long, stigmas simple to capitate. Berries green, often with reddish 
purple tinge, 5-loculed, subglobose to obovoid, to 10 mm. across, 5-sulcate. 

Habitat. Rain forest, humid mountain forest, cleared slope, 



SOEJARTO: SA URA VIA 37 

humid rocky riverbank, road cut, shady and open areas, at altitudes 
of 800-2, 000(-3, 600?) m. 

Distribution. Ecuador (Provinces of Tungurahua, Napo-Pastaza, 
Santiago-Zamora). 

Vernacular names. Moco (Tungurahua: Heinrichs). 

Specimens examined. ECUADOR, TUNGURAHUA: vie. of Am- 
bato, Aug., fl., Rose & Rose 22377 (NY, US); Banos and vie., Jan., fl., 
Heinrichs 268 (G, NY, Z), March, fl., Mexia 6999 (F, UC), fl., Pen- 
land & Summers 99 (F, US), July, fl., Sandeman 23 (K), April, fl., 
Benoist 4229 (P), 4200 (P), fr., Sodiro s. n. (P); road Banos to Puyo, 
bank of Pastaza River, Aug., fl. fr., Soejarto & Hernandez 1342 
(ECON); Mt. Tungurahua, southwest slope, headwater of Agua de 
Oro, April, ft., Heinrichs 855 (B, G, NY, Z), northern slope, Jan., fl., 
Rimbach 46 (K, MICH), Oct., fr., 270 (F, S); valley of Pastaza River, 
road Banos to Cashurco, Sept., fl., Hitchcock 21753 (GH, NY, US); 
no locality, Hacienda La Merced, Feb., fr., Mexia 7008 (UC, US), 
Jan., fl. fr., Rimbach 496 (S). SANTIAGO-ZAMORA: Huamboya, 
Feb., fl., Solis 7518 (F). 

NO PRECISE LOCALITY: In Andibus Ecuadorensibus, fl., 
Spruce 4989 (BM, F, G, GH, NY, P, S). 

S. aequatoriensis is fairly well represented in the areas surround- 
ing Banos (Tungurahua and Napo-Pastaza), where the plants com- 
monly grow in shady and humid places, particularly in the forested 
area along the Pastaza river, at altitudes of 1,300-1,800 m. This is 
probably the area where the type was collected by Spruce. The al- 
titude given by Heinrichs (Heinrichs 855) as 3,600-3,800 m. (head- 
water of Agua de Oro) is questionable, because this is too high for 
the species. Solis 7518, the only record from Santiago-Zamora, dif- 
fers from specimens collected in the Tungurahua and Napo-Pastaza 
areas in its glabrous, moderately branched inflorescences, and its 
smaller flowers with completely glabrous sepals. 

12. Saurauia adenodonta Sleumer, Notizbl. 15: 372. 1941. 
Type: Schultze-Rhonhof '2914 (K, lectotype). Figure 9. 

Shrubs; sparingly pubescent. Branchlets terete, densely tuberculate-strigose 
pubescent. Leaves clustered behind tip of branchlets; blades oblong-elliptic to 
oblong-obovate, shortly and abruptly acuminate at apex with acumen to 10 mm. long, 
broadly cuneate to rarely oblique at base, finely and distantly serrulate along mar- 
gins, 12-23 cm. long, 5-8 cm. wide, chartaceous, glossy and smooth above, secondary 
veins 16-20 pairs, tertiary veins immersed, scarcely more prominent than lesser 
venation, glabrous except for distantly scattered tuberculate trichomes along veins 



38 FIELDIANA: BOTANY 

above, scattered to sparingly pubescent with trichomes of radiate to clustered (along 
veins) and tuberculate-strigillose types (along midrib) beneath; petioles 1-2.5 cm. 
long, 1-2.5 mm. in diameter, densely tuberculate pubescent. Inflorescences somewhat 
straight, 100-ca. 200-flowered, 9-22 cm. long, 2-7 cm. wide, abundantly pubescent 
with trichomes of radiate to stellate mixed with shaggy -strigillose to shaggy-strigose 
types, primary peduncle 4.5-9.5 cm. long, bracts linear, to 3.5 mm. long, very rarely 
foliaceous, to 11 mm. long. Flowers 7-9 mm. broad, buds ellipsoidal (sepals become 
fully open and extended, and curved outward in mature buds), to 3.5 mm. long, to 2.5 
mm. in diameter, pedicels to 3 mm. long, bracteoles linear, to 2 mm. long; sepals 5, 
elliptic to oblong-elliptic, subacute to obtuse, 3-3.5 mm. long, 2-2.5 mm. wide, exposed 
parts in bud densely pubescent with trichomes of stellate and strigillose types, imbri- 
cated parts stellate pubescent, all abundantly to densely stellate pubescent on upper 
half but glabrous on lower half inside, marginally ciliolate; petals 5, oblong to 
elliptic-oblong, rounded, 3.5-4 mm. long, 1.5-2.5 mm. wide; stamens 20-30, filament 2 
mm. long, anther 1-1.5 mm. long; ovary 3-5-loculed, subglobose, 3-5-sulculate, gla- 
brous, styles 3-5, obsolete to 3.5 mm. long, stigmas simple to subcapitate. Berries 
unknown. 

Distribution. Ecuador (Province of Napo-Pastaza). 

Specimens examined. ECUADOR, NAPO-PASTAZA: Region of 
Pastaza River, Mera, March, fl., Lugo 11 (S), fl., Schultze-Rhonhof 
2914 (K). 

S. adenodonta is easily distinguished from all other Ecuadorian 
species by its small flowers (less than 10 mm. broad), its small and 
ellipsoidal flower buds, and its glabrescent leaves. The species is 
apparently related to S. laevigata. 

13. Saurauia strigillosa Tr. & PI., Ann. Sci. Nat. ser. 4, 18: 266. 
1862; Prodr. Fl. Nov. Granat. 1: 263. 1862. Type: Triana s. n. (G, 
lectotype; F-fragment, P, isolectotypes; GH, US, photos). Figure 11. 

Saurauia strigillosa var. microphylla Busc., Malpighia 30: 107. 
1927. Type: Goudot s. n. (P; US, photo). 

Saurauia floribunda Benth. ex Sprague var. laevigata Busc., 1. c. 
28: 113. 1917. Type: Goudot s. n. (P; US, photo). 

Trees; sparingly pubescent. Branchlets slender, terete, sparingly strigillose pubes- 
cent. Leaves crowded behind tip of branchlets; blades elliptic to oblanceolate or 
oblong-obovate, abruptly and very shortly acuminate at apex, narrowly cuneate at 
base, serrulate along margins, 6-15 cm. long, 2-5 cm. wide, chartaceous to subcoria- 
ceous, glossy and smooth above, in dry state sooty above, dark olive-brown beneath, 
secondary veins 10-15 pairs, tertiary veins immersed, scarcely more prominent than 
lesser venation, scattered to sparingly strigillose pubescent along major veins and 
scattered stellate pubescent to glabrescent along minor veins with glabrous and 
pustulate epidermis above and beneath; petioles 1-2 cm. long, 1-1.5 mm. in diameter, 
terete, sparingly strigillose pubescent. Inflorescences little branched, often somewhat 
flexuous, (3-)10-20-flowered, (2-)5-ll cm. long, 2-4 cm. wide, abundantly pubescent 



SOEJARTO: SA URA VIA 39 

with trichomes of strigillose and stellate types, primary peduncle (l-)2-6 cm. long, 
bracts subulate, to 4 mm. long. Flowers 15-18 mm. broad, buds to 5 mm. in diameter, 
pedicels 3-6 mm. long, bracteoles triangular, to 2 mm. long; sepals 5, elliptic to 
oblong, obtuse, 4-5 mm. long, 2.5-3.5 mm. wide, exposed parts in bud abundantly to 
densely pubescent with trichomes of strigillose and stellate types, imbricated parts 
densely stellate pubescent, all densely stellate pubescent inside, marginally ciliolate; 
petals 5, white, oblong to oblong-obovate, rounded to often incised or tridentate, 6-8 
mm. long, 4-5 mm. wide; stamens 20-30, filament 2.5 mm. long, anther 2 mm. long; 
ovary 5-loculed, subglobose, 5-sulcate, glabrous, styles 5, 1-3.5 mm. long, stigmas 
simple to capitate. Berries unknown. 

Habitat. Mountain forest, subparamo, along creeks, at altitudes 
of 1,500-2,800 m. 

Distribution. Colombia (Departments of Antioquia, Quindio, To- 
lima). 

Vernacular Names. Chachafruto (Quindio: Goudot). 

Specimens examined. COLOMBIA, ANTIOQUIA: Paramo of 
Sonson, Jan., fl., Daniel 3463 (US). QUINDIO: Dec., fl., Goudot s. n. 
(FI, P; US, photo). TOLIMA: Mariquita, fl., Triana s. n. (G, lec- 
totype; F-fragment, K, P, isolectotypes; GH, US, photos); Ibague, La 
Palmilla, Nov., fl., Goudot s. n. (P; US, photo). 

14. Saurauia laevigata Tr. & PL, Ann. Sci. Nat. ser. 4, 18: 267. 
1862; Prodr. Fl. Nov. Granat. 1: 264. 1862; Hunter, Ann. Missouri 
Bot. Gard. 53: 84. 1966, p. p. min. Type: Triana s. n. (G, lectotype; 
P, isolectotype; GH, photo). Figure 10. 

Trees; sparingly pubescent to glabrescent. Branchlets slender, terete, scattered 
tuberculate pubescent to glabrescent. Leaves obovate to oblong-obovate, very shortly 
and abruptly acuminate at apex, cuneate at base, rarely oblique, distantly and finely 
serrulate along margins, 10-24 cm. long, 5-10 cm. wide, chartaceous, glossy and 
smooth above, in dry state sooty above, dark dull gray-brown beneath, secondary 
veins 12-16 pairs, tertiary veins somewhat immersed, scarcely more prominent than 
lesser venation, scattered floccose pubescent along major veins but mostly glabrous 
above, scattered tuberculate pubescent to glabrescent along midrib but mostly gla- 
brous with pustulate epidermis beneath; petioles 2-4 cm. long, 2-3 mm. in diameter, 
somewhat terete, smooth and glabrescent. Inflorescences straight, 30-ca. 100- 
flowered, 13-22 cm. long, 4-6 cm. wide, glabrescent along lower portion, abundantly 
pubescent with stellate trichomes along lesser ramification, primary peduncle 5-10 
cm. long, bracts subulate, to 3 mm. long. Flowers 15 mm. broad, buds to 5 mm. in 
diameter, pedicels to 5 mm. long, bracteoles subulate, to 3 mm. long; sepals 5, outer 
two oblong to oblong-ovate, obtuse, 3-3.5 mm. long, 2 mm. wide, imbricate one and 
inner two suborbicular, rounded, 3-4 mm. long, 3-3.5 mm. wide, all abundantly stel- 
late pubescent throughout, marginally and apically ciliolate; petals 5, oblong- 
obovate, rounded, 12-14 mm. long, 4-5.5 mm. wide; stamens 20-30, filament 3.5-4.5 
mm. long, anther 2.5 mm. long; ovary 5-loculed, globose, 5-sulcate, glabrous, styles 5, 
1.5 mm. long, stigmas simple. Berries unknown. 



40 



FIELDIANA: BOTANY 



Habitat. Forest, at altitudes of ca. 1,400 m. 

Distribution. Colombia (Departments of Antioquia and Tolima). 

Vernacular names. Dulumoco (Antioquia: Uribe-Uribe). 

Specimens examined. COLOMBIA, ANTIOQUIA: Narino, 
Quebradita El Oso, Jan., fl., Uribe-Uribe 1912 (COL, ECON). TO- 
LIMA: Mariquita, fl., Triana s. n. (G, lectotype; BM, K, P, isolec- 
totypes; GH, photo). 

Based upon studies of the type specimen, S. laevigata is here de- 
fined in a narrower sense than that treated by Hunter (1966), who 
included in it S. yasicae and its synonyms as well. The two species 
are taxonomically distinct and their separation is warranted by the 
following characteristics: 



Leaves: 

Lateral veins: 
Foliar tri- 

chome types: 
Flowers per 

inflorescence: 
Flowers: 

Sepals (both surfaces): 

Filament: 
Anther: 



S. laevigata 
Usually large, subcoriaceous 

to chartaceous 
10-15 pairs 
Strigillose and stellate 

(3-UO-20 

ca. 15 mm. broad, 

pentamerous 
Abundantly pubescent 

3.5-4.5 mm. long 
Linear, ca. 2.5 mm. long 



S. yasicae 
Usually small, 

membranaceous 
5-15(-18) pairs 
Clustered to tufted 

or setose-tufted 
(20-HO-150 

7-10 mm. broad, 

tetramerous 
Glabrous to sparingly 

pubescent 
1.5-2.5 mm. long 
Cordate, 1-1.5 mm. long 



IV. Ser. PARVIFLORAE Soejarto, ser. nov. 

Brachitrichae Busc., Malpighia 25: 220. 1912, p. p. min. 
Gynogynae Busc., 1. c. 221, p. p. min. 
Stenobasicae Busc., 1. c. 221, p. p. min. 

Plantae glabrescentes flores parvos habentibus, sepala omnino glabri intus et om- 
nino glabri vel partim et parce pubescentes extus. 

1. Plants glabrous throughout, secondary veins 35-37 pairs; Colombia 

15. S. multinervis 

1. Plants sparingly pubescent (S. chiliantha copiously pubescent), secondary veins 
less than 30 pairs. 
2. Flowers 20 mm. or more broad, 6-20 per inflorescence, secondary veins 15-18 

pairs 16. S. schultesiana 

2. Flowers 7 to less than 20 mm. broad. 

3. Leaves villous-barbate at axils of secondary (often tertiary) veins beneath; 

Peru 17. S. peruviana 

3. Leaves not villous-barbate beneath. 



SOEJARTO: SA URA VIA 41 

4. Lower leaf surface densely to abundantly stellate pubescent, epidermis 
usually obscured by pubescence 18. S. chiliantha 

4. Lower leaf surface glabrescent to sparingly pubescent, trichomes mostly 
not of stellate type, epidermis not obscured by pubescence. 

5. Texture of leaves strongly coriaceous, often very rigid and brittle, leaf 
margins aculeo-serrulate with hard and sharp-pointed serrulation. 

6. Leaf blades 10-23 cm. long, 3.5-6.5 cm. wide, secondary veins 
14-16 pairs, major veins abundantly pubescent beneath, trichomes 
to 4 mm. long, flowers 5-8 mm. broad; Peru 19. S. solitaria 

6. Leaf blades 20-41 cm. long, 13-17 cm. wide, secondary veins 20-25 
pairs, major veins sparingly pubescent beneath, trichomes to 2 
mm. long, flowers 10-15 mm. broad; Colombia 20. S. micayensis 

5. Texture of leaves chartaceous to membranaceous, rarely subcoria- 
ceous, leaf margins serrulate, serrulations not spiny, often terminat- 
ing in setae. 

7. Sepals glabrous throughout. 

8. Inflorescences 15-60-flowered, stamens 20-35, leaf texture 
chartaceous to membranaceous; Peru 21. S. natalicia 

8. Inflorescences 40-200-flowered, stamens 40-70, leaf texture 
subcoriaceous; Colombia 22. S. parviflora 

1. Sepals partly to completely pubescent outside. 

9. Inflorescences 20-150-flowered, flowers 7-10 mm. broad, sta- 
mens 15-20, leaves scattered radiate pubescent beneath; Col- 
ombia 23. S. pseudoleucocarpa 

9. Inflorescences 50-500-flowered, flowers 10-19 mm. broad, sta- 
mens 45-85, leaves sparingly sericeous-strigose pubescent; 
Bolivia 24. S. spectabilis 

15. Saurauia multinervis Soejarto, Bot. Mus. Leafl. 22: 266. 
1969. Type: Idrobo & Fernandez 54 (US, holotype; COL, isotype). 
Figure 12. 

Shrubs to 2 m. tall, many-branched; glabrous. Leaf blades elongate-obovate, cuspi- 
date at apex, broadly cuneate to obtuse at base, dentate-serrate with minute and 
sharp-pointed serrations along margins, 30-42 cm. long, 12-14 cm. wide, coriaceous, 
dirty dark olive-brown above in dry state, brownish olive beneath, laevigate and 
smooth above, secondary veins 35-37 pairs, tertiary veins elevated, more prominent 
than lesser venation, both surfaces glabrous; petioles 1-2 cm. long, 3-5 mm. in diame- 
ter, glabrous. Inflorescences straight, ca. 150-flowered, 23 cm. long, 15 cm. wide, 
glabrous, primary peduncle 1.5 cm. long, bracts linear, to 5 mm. long. Flowers not 
known. Fruits laxly distributed, berries 5-loculed, globose, to 6 mm. in diameter, 
5-sulcate, glabrous; pedicels 4-8 mm. long, bracteoles linear to 3 mm. long; persistent 
sepals 5, oblong-elliptic to ovate, obtuse, 5-6.5 mm. long, 3-4.5 mm. wide, all glabrous 
throughout, marginally entire to irregularly ciliolate; persistent styles 5, 4.5-5 mm. 
long, stigmas capitate. 

Habitat. Altitude of 2,000 m. 



42 FIELDIANA: BOTANY 

Distribution. Colombia (Department of Cauca). 
Vernacular names. Lulumoco (Cauca: Idrobo & Fernandez). 

Specimens examined. COLOMBIA, CAUCA: Cordillera Occi- 
dental, eastern slope, El Tambo, Aug., fr., Idrobo & Fernandez 54 
(US, holotype; COL, isotype). 

16. Saurauia schultesiana Soejarto, Bot. Mus. Leafl. 22: 267. 
1969. Type: Pennell 10501 (NY, holotype). Figure 13. 

Trees; copiously to sparingly pubescent. Branchlets terete, abundantly pubescent 
with rusty strigose to setose trichomes, young leaves and shoot deep brown in dry 
state. Leaves crowded around tip of branchlets; blades elliptic to oblong-obovate, 
acuminate at apex with acumen to 15 mm. long, cuneate at base, serrulate along 
margins, 14-18 cm. long, 7-8 cm. wide, subcoriaceous, in dry state dark dirty brown 
above, grayish olive-brown beneath, scarcely scabrous above and beneath, secondary 
veins 15-18 pairs, tertiary veins elevated, more prominent than lesser venation, 
sparingly to abundantly pubescent beneath, with trichomes of setose to setulose 
mixed with radiate types along and between minor veins and of strigose types along 
major veins; petioles ca. 2 cm. long, 2-3 mm. in diameter, half-terete, abundantly 
rusty strigose pubescent. Inflorescences straight, 6-20-flowered, 13-30 cm. long, 2-7 
cm. wide, pulverulent, abundantly scurfy-strigose pubescent, primary peduncle 8-15 
cm. long, bracts broadly triangular to suborbicular, 10-15 mm. long, 5-7 mm. wide, 
glabrous. Flowers 20-25 mm. broad, buds to 9 mm. in diameter, pedicels 5-10 mm. 
long, bracteoles broadly triangular, to 10 mm. long; sepals 5, outer two trullate to 
oblong-ovate, subacute, 10-12 mm. long, 6-7 mm. wide, imbricate one suborbicular, 
10-12 mm. long, 8-9 mm. wide, inner two suborbicular to orbicular-oblong, 10-14 mm. 
wide, all glabrous to medially sparingly strigillose pubescent outside, glabrous in- 
side, marginally entire to apically ciliolate; petals 5, white, subquadrangular to 
spatulate, rounded, 15-19 mm. long, 9-11 mm. wide; stamens 25-35, filament 4-5 mm. 
long, anther 2.5-3 mm. long; ovary 5-7-loculed, globose, 5-7-sulcate, glabrous, styles 
5-7, to 7.5 mm. long, stigmas capitate. Berries 5-7-loculed, globose, to 6 mm. across 
(immature), 5-7-sulcate. 

Habitat. Mossy forest at altitudes of 3,200-3,400 m. 
Distribution. Colombia (Department of Caldas). 

Specimens examined. COLOMBIA, CALDAS: Cordillera Occi- 
dental, Cerro Tatama, Sept., fl. fr., Pennell 10501 (NY, holotype). 

The floral characters of S. schultesiana very much resemble those 
of S. omichlophila, but the inflorescence approaches that of S. bul- 
losa, whereas the leaf pubescence is similar to that of S. excelsa. 
This combination of characters, therefore, is unique and easily sets 
it apart from all other species. In this connection, the following 
comment by Schultes (1943, p. 45), when he described S. spinuligera 
from the same locality as that of S. schultesiana, is noteworthy: 'The 
presence of such an extraordinarily distinct species of Saurauia on 



SOEJ ARTO: SA URA UIA 43 

Cerro Tatama, which is in fact, an island-mountain isolated from 
neighboring elevations, would seem to suggest that more interest- 
ing endemic species or varieties of this genus might be found there." 

17. Saurauia peruviana Busc., Malpighia 27: 319. pi. 8, fig. 15. 
1916. Type: Haenke s. n. (COL, GH, photos). 

Saurauja scabra Poepp. ex Choisy, Mem. Soc. Phys. Geneve 14: 
116. 1855, excl. syn. Palaua biserrata R. & P. and P. glabrata 
DC. Type: Poeppig 127 (also numbered Poeppig 1694) (G, 
holotype; F, GH, P, U, isotypes; COL, NY, photos). 

Saurauia pseudoscabra Busc., Malpighia 28: 1. 1917. Type: 
Weberbauer 842 (COL, GH, NY, photos; photo in NY labeled 
as 942) 

Saurauia scabra Poepp. ex Choisy var. boliviano Busc., Mal- 
pighia 27: 501. 1916. Type: Weddell 4206 (P, lectotype). 

Saurauia floribunda Benth. ex Sprague var. peruviana Busc., 
Malpighia 28: 119. 1917. Type: Mathews s. n. (G, K). 

Saurauia weberbaueri Busc., Malpighia 30: 148. pi. 5, fig. 8. 
1927. Type: Weberbauer 4454 (COL, NY, photos). 

Saurauia pseudoparviflora Busc., Malpighia 30: 158, 1917, p. p. 
maj. typ. incl. Type: Bang 387 A (G, lectotype; F, MICH, 
NY, US, isolectotypes). 

Saurauia coroicoana Busc., Malpighia 30: 197. pi. 9, fig. 17. 
1927. Type: No collector s. n. (P, lectotype). 

Saurauia pyramidata Sleumer, Notizbl. 12: 145. Type: Buchtien 
2205 (US, lectotype; NY, isolectotype). 

Saurauia scabriuscula Macbr., Fl. Peru 3A(2): 683. 1956. A new 
name for S. scabra Poepp. ex Choisy. 

Shrubs to 3 m. tall; copiously pubescent. Branchlets slender, terete, abundantly to 
sparingly strigose to shaggy-strigose pubescent. Leaves clustered behind tip of 
branchlets; blades obovate to oblong-obovate, very shortly and abruptly acuminate to 
acute at apex, cuneate at base, serrulate along margins, 10-18(-25) cm. long, 3-6C-10) 
cm. wide, chartaceous, somewhat scabrous above, secondary veins 13-20 pairs, ter- 
tiary veins elevated, more prominent than lesser venation, sparingly to abundantly 
setulose to strigillose pubescent along and between minor veins (trichomes often 
reduced to mere warts) and abundantly appressed-setose to strigose pubescent along 
major veins above, abundantly pubescent beneath with trichomes of setulose to setose 
mixed with stellate to stellate-setose or shaggy types along and between veins, 
villous-barbate at axils of secondary veins beneath; petioles 1.5-3(-6) cm. long, 1.5-2 
mm. in diameter, abundantly appressed-setulose pubescent. Inflorescences somewhat 
straight, (25-)50-ca. 200(-300)-flowered, 3-18 cm. long, 2-7 cm. wide, densely to abun- 
dantly setulose to shaggy-setulose pubescent, primary peduncle 1-7 cm. long, bracts 



44 FIELDIANA: BOTANY 

triangular, to 3 mm. long. Flowers 5-8 mm. broad, buds to 3.5 mm. in diameter, 
pedicels 1-2 mm. long, bracteoles triangular, to 1.5 mm. long; sepals 5, orbicular 
ovate to suborbicular, subacute to rounded, 1.5-2.5 mm. long, 1-2 mm. wide, exposed 
parts in bud usually glabrous to rarely pubescent with trichomes of stellate to shaggy 
types, imbricated parts glabrous, all glabrous inside, marginally ciliolate to promi- 
nently dilate; petals 5, white, oblong, rounded, 3-5 mm. long, 2.5-5 mm. wide; sta- 
mens 15-30, filament 2 mm. long, anther 1 mm. long; ovary 3-5-loculed, ovoid, 3-5- 
sulculate, glabrous, styles obsolete to 3 mm. long, stigmas simple to subcapitate. 
Berries 3-5-loculed, subglobose, to 4 mm. across, 3-5-sulcate. 

Habitat. Wet wooded slope, edge of brushy ravine in clearing, 
forest remnant in cleared land, Andean forest, edge of forest on 
sandy soil, at altitudes of 750-2,700 m. 

Distribution. Ecuador (Province of Santiago-Zamora), Peru (De- 
partments of Cajamarca, Amazonas, Huanuco), Bolivia (Provinces 
of La Paz and Cochabamba). 

Specimens examined. ECUADOR, SANTIAGO-ZAMORA: 18 
km. ENE. of Loja, Rio San Francisco, Canillones, Feb., fl., Fosberg 
23145 (NY, P, US). 

PERU, CAJAMARCA: Colassay, Oct., fl., Woytkcwski 6943 (GH). 
AMAZONAS: Bongara, Jalca Zone along Shipasbamba-Pomacocha 
trail, June, fl., Wurdack 1112 (F, NY, UC, US); Chachapoyas, fl. fr., 
Mathews s. n. (BM, G, MA). HUANUCO: Cuchero, fl., Poeppig 127 
(=Poeppig 1694} (BM, F, G, GH, HAL, L, NY, P; COL, photo); Chin- 
chao, veg., Rivero s. n. (P). NO PRECISE LOCALITY: fl., Haenke s. 
n. (COL, GH, photos); fl., Weberbauer 842 (COL, NY, photos), 4454 
(NY, photo). 

BOLIVIA, LA PAZ: Murillo, in ravine of Coroico, fl., No Collector 
s. n. (P); Yungas, fl. fr., Bang 387A (F, G, K, MICH, NY, US), Dec., 
fl., Mexia 4310 (GH, UC, US), fl., Rusby 483 (F, MICH, NY, US), 
Weddell 4206 (P). COCHABAMBA: Chapare, Palmar, April, fl., 
Cardenas 5992 (US); Antahuacana, Espiritu Santo, 160 km. NE. of 
Cochabamba, June, fl., Buchtien 2205, fr., 2222 (both in NY, US). 
NO PRECISE LOCALITY: Unduavi, Oct., fl., Rusby 506 (MICH, 
NY). 

NO LOCALITY: fl.,Pavons. n. (Fl). 

S. scabra Poepp. ex Choisy is a homonym of S. scabra (HBK.) 
Dietr. (1843), and must be rejected according to the Code. The 
specific name S. scabriuscula proposed by Macbride (1955) is 
nomenclaturally superfluous, because an earlier name, S. peruviana 
Busc. (1916), based upon a different type (Haenke s. n.) , but which is 



SOEJ ARTO: SA URA UIA 45 

taxonomically synonymous, is available. Therefore, the correct 
name of this taxon is Saurauia peruviana Busc. 

Bang 387 is the type of S. pseudoparviflora Busc. Another collec- 
tion, also labeled Bang 387, was described by Buscalioni as S. rus- 
byi (non Britton) var. glabrata Busc., and still another Bang 387 
was described as S. rusbyi (non Britton) var. spectabilis Busc. fma. 
macrophylla Busc. Another collection, Bang s. n., was described by 
Rusby as S. brevipes, treated here as a synonym of S. spectabilis 
Hook. Careful examination of all Bang's collection revealed that, 
first, there appear to be two different elements erroneously labeled 
Bang 387; second, S. rusbyi (non Britton) var. glabrata Busc., S. 
rusbyi (non Britton) var. spectabilis Busc. fma. macrophylla Busc., 
and S. brevipes Rusby are synonymous. For the purpose of clarity, a 
new number, Bang 387 A, has been assigned to the first group of 
specimens that represents the type of S. pseudoparviflora Busc., 
while another number, Bang 387B, has been assigned to the second 
group that represents the type of S. rusbyi var. glabrata Busc., S. 
rusbyi var. spectabilis Busc. fma. macrophylla Busc., and S. brevipes 
Rusby. 

Due to the wide distribution of S. peruviana, a considerable mor- 
phological variation exists between the populations, one of the rea- 
sons for the long synonymy. This variation is summarized as follows. 
Poeppig 127 has relatively small leaves with unbranched multicellu- 
lar trichomes (setulose type) on the lower surface, short, little- 
branched, few-flowered inflorescences (less than 50 flowers per in- 
florescence), and flowers with glabrous sepals. Other Peruvian collec- 
tions have branched trichomes (shaggy type) on the lower leaf sur- 
face, little- to moderately-branched, few- to many-flowered inflores- 
cences (50-100 flowers per inflorescence), and flowers with sepals 
glabrous to sparingly or abundantly pubescent outside; the leaf size 
also varies from 1 1 cm. long by 4 cm. wide to 24 cm. long by 8 cm. wide. 
Fosberg 23145, the only known Ecuadorian collection, perfectly 
matches Poeppig 127, but the leaves are slightly larger and the 
inflorescences are many -branched and many-flowered. The Bolivian 
collections, particularly those referred to as S. pseudoparviflora 
Busc., have slightly larger leaves in general than the Peruvian col- 
lections, and the lower leaf surface also varies from glabrescent (with 
setulose trichomes along the major veins) to abundantly shaggy 
pubescent, with many-branched and many-flowered inflorescences, 
and flowers with glabrous sepals. Buchtien 2205, described by 



46 FIELDIANA: BOTANY 

Sleumer as S. pyramidata, has few-branched and few-flowered in- 
florescences, and, although Sleumer mentioned that "ovarium parce 
pilosum" examination of the duplicate types (NY, US) shows that the 
ovary is glabrous. 

Notwithstanding the variable nature of the populations, they do 
have the following combination of features, which characterize the 
species: (1) leaves villous-barbate pubescent on the axils of the secon- 
dary and often tertiary veins beneath (especially the Bolivian collec- 
tions), (2) flowers small (5-8 mm. broad when open); (3) sepals lacking 
unbranched multicellular trichomes outside (glabrous on the unex- 
posed surface in bud, and inside); (4) stamen number low (15-30). 

18. Saurauia chiliantha R. E. Schultes, Caldasia 2: 32. 1943. 
Type: Dugand & Jaramillo 2988 (COL, holotype; GH, US, isotypes). 
Illustration: Caldasia 2: 35. 1943. 

Saurauia kail ima R. E. Schultes, Mutisia 3: 5. 1952. Type: Daniel 
3425 (COL, holotype; MEDEL, US, isotypes). 

Shrubs to small trees to 10 m. tall, crown open; copiously pubescent. Branchlets 
robust, terete, prominently scarred, densely setose to hirsute pubescent, often wooly to 
touch, trichomes very slender, long and weak, yellowish to reddish brown. Leaves 
clustered behind tip of branchlets; blades obovate to oblong-obovate or elongate- 
obovate, mucronulate-cuspidate to rounded at apex, cuneate at base, distantly 
setulose-serrulate with fine, stiff and sharp-pointed serrulations along margins, (10-) 
17-28 cm. long, (5-)7-13(-16) cm. wide, strongly coriaceous, often stiff and brittle, dark 
green above, gray to reddish brown beneath (in dry state light to dark gray-brown or 
sooty above), secondary veins 20-28 pairs, tertiary veins elevated, more prominent 
than lesser venation, scattered to abundantly pubescent with trichomes of 
tuberculate-setose, strigose and villous types along veins above, abundantly to densely 
pubescent with trichomes of stellate to dendroid mixed with setose to hirsute types 
along and between veins beneath; petioles stout, 1-3 cm. long, 3-5 mm. in diameter, 
canaliculate, abundantly to densely pubescent with trichomes of strigose to villous 
types. Inflorescences straight, often spreading, 100- to more than 500-flowered, 
(6-UO-20 cm. long, (2-H-10 cm. wide, densely to abundantly pubescent with trichomes 
of setose to villous mixed with silvery-stellate types, primary peduncle 0.5-4(-5.5) cm. 
long, bracts triangular to linear, to 10 mm. long, very rarely foliaceous, to 20 mm. long. 
Flowers 8-10 mm. broad, buds to 4 mm. in diameter, pedicels to 3 mm. long, bracteoles 
linear to triangular, to 3 mm. long; sepals 5, pale green to white, suborbicular to 
orbicular-obovate, acute to rounded, 2.5-3.5 mm. long, 2.5-3 mm. wide, all glabrous 
throughout, but sometimes medially scattered strigillose pubescent outside, 
marginally irregularly ciliolate; petals 5, white to creamy yellow, oblong to 
obovate-oblong, rounded and often incised, 4-5 mm. long, 2-3 mm. wide; stamens 20-30, 
filament 2-2.5 mm. long, anther 1.5 mm. long; ovary 5-loculed, globose, 5-sulcate, 
glabrous, styles 5, obsolete to 4.5 mm. long, stigmas simple to subcapitate. Berries 
5-loculed, subglobose, to 6 mm. across, 5-sulcate. 

Habitat. Humid mountain forest and subparamo, at altitudes of 
(1,900-) 2,300-2,900 m. 



SOEJARTO: SAt/flA[//A 47 

Distribution. Colombia (Departments of Antioquia, Quindio, 
Caldas, Tolima, and Valle). 

Vernacular names. Dulumoco (Antioquia: Daniel). 

Specimens examined .COLOMBIA, ANTIOQUIA: Road Sonson 
to Alto Capiro, May, fl., Core 770 (NY, US); road Sonson to Abejorral, 
May, fl., Ewan 15756 (NY, US); road Sonson to Mesopotamia, June, 
ft., Soejarto & Rivera 2044 (ECON, GH, MEDEL); Paramo of Sonson, 
Jan., fl., Daniel 3425 (COL, MEDEL, US), March, fl., Scolnik et al. 
19An215 (US), June, fl., Rivera 313, 314 (both in MEDEL), June, fl., 
Soejarto & Rivera 2047, 2048 (both in ECON, GH, MEDEL); Amaga, 
Jan., fl., Uribe-Uribe 1435 (COL). QUINDIO: between Circasia and 
Pereira, vie. of Alto El Roble, Aug., fl., Dugand & Jaramillo 2988 
(COL, GH, US). CALDAS: Vic. of Manizales, Quebrada La Minita, fl. 
fr., Soejarto 2027 (ECON, GH); road Manizales to Nevado del Ruiz, 
vie. of Miraflores, fl., Soejarto 2031 (ECON, GH); San Felix, near 
Salamina, July, fl., Tomas 2092 (US), Oct., fl., 3121 (MEDEL). TO- 
LIMA: Road El Libano to Murillo, km. 11 to 22, at Alto de Penones, 
July, fl., Garcia-Barriga 12263, 12269 (both in US). VALLE: Cordil- 
lera Occidental, El 18 (Cali), Aug., fl., Espinal 1898 (ECON, US); 
San Antonio (W. of Cali), March, fl., Killip & Garcia-Barriga 33929 
(US). NO PRECISE LOCALITY: Novilleros-Quindio, March, fl., 
Andre 2318 (F, K, NY). 

S. chiliantha is distinguished from all other Colombian species by 
the following combination of characters: (1) leaves strongly coria- 
ceous (often very rigid) and densely pubescent beneath with 
trichomes of stellate mixed with setose to hirsute types; (2) inflores- 
cences very shortly pedunculate with profuse small flowers; (3) sepals 
glabrous or at least glabrous throughout except for the sparingly 
distributed strigillose trichomes on the exposed parts in bud; (4) 
stamens low in number (20-30). The species is abundantly 
represented in the lower range of the Paramo of Sonson (Antioquia) 
at altitudes of 2,500-2,900 m. Members of the populations from this 
area (described by Schultes as S. kallima) differ from those from 
Caldas (locality of the type of S. chiliantha) primarily in their more 
rigid leaves. 

Several individuals collected from Sonson (Scolnik et. al. 19An215, 
Rivera 314, Soejarto & Rivera 2041) have unusually glabrescent 
leaves (sparingly radiate pubescent on the lower surface), whereas 
Garcia-Barriga's collections (12263 and 12269) from Tolima and 
Killip and Garcia-Barriga's (33929) from Valle have unusually 



48 FIELDIANA: BOTANY 

dense pubescence on the outer surface of the sepals. Aside from these 
differences, other characters appear constant. 

19. Saurauia solitaria Sleumer, Notizbl. 12: 147. 1934. Type: 
Weberbauer 7067 (F, lectotype). Figure 14. 

Shrubs to 2 m. tall; copiously pubescent. Branchlets slender, terete, densely and 
coarsely hirsute pubescent; trichomes brown to deep reddish brown, silky, upward 
curved, slightly swollen at base and turning gradually very slender towards the tip, 
to 7 mm. long. Leaves distributed from tip of to somewhat low along branchlets, 
blades elliptic to elongate-obovate, cuspidate to shortly acuminate at apex with hard- 
and sharp-pointed acumen to 15 mm. long, cuneate to obtuse at base, spiny-serrate 
with serrations hard- and sharp-pointed along margins, 10.5-23 cm. long, 3.5-6.5 cm. 
wide, firmly coriaceous, sooty to black above in dry state, reddish brown beneath, 
glossy and somewhat smooth above, secondary veins 14-16 pairs, tertiary veins 
somewhat elevated, scarcely more prominent than lesser venation, sparingly pubes- 
cent with trichomes of setulose to strigose to setose types along and between veins 
above and beneath; petioles 1-1.5 cm. long, 2 mm. in diameter, abundantly hirsute 
pubescent with trichomes of similar types and color to those along branchlets. In- 
florescences straight, 40-60-flowered, 15-16 cm. long, 4.5-6 cm. wide, densely setose 
pubescent, primary peduncle slender, 8-9 cm. long, bracts linear, to 5 mm. long, 
glabrous. Flowers 8-10 mm. broad, to 3 mm. in diameter, pedicels very slender, 3-7 
mm. long, bracteoles linear, to 3 mm. long, glabrous; sepals 5, elliptic to oblong- 
elliptic (ovate-elliptic, cf. Sleumer), obtuse to rounded, 3-3.5 mm. long, 1.8-2.5 mm. 
wide, all glabrous throughout, marginally entire to rather irregularly ciliate around 
apex; petals 5, rosy, oblong-obovate (elliptic-rotundate, cf. Sleumer), rounded and 
slightly incised, 4-4.5 mm. long, 2.5 mm. wide; stamens 18-22, filament 2-2.25 mm. 
long, anther 1-1.25 mm. long; ovary 5-loculed, ovoid, 5-sulculate, glabrous, styles 5, 
1.5 mm. long, stigmas simple. Berries not known. 

Habitat. Riverbank and mountain forest at altitudes of 1,700- 
2,000 m. 

Distribution. Peru (Department of La Libertad). 

Specimens examined. PERU, LA LIBERTAD: Pataz, valley of 
Mixiollo, one of the left tributaries of Rio Huallaga, Aug., fl., Weber- 
bauer 7067 (F, lectotype). 

20. Saurauia micayensis Killip, J. Wash. Acad. Sci. 16: 570. 
1926. Type: Killip 7932 (US, lectotype; A, photo). Figure 15. 

Saurauia rigidissima R. E. Schultes, Bot. Mus. Leafl. 13: 283. 
1949._Type: Cuatrecasas21716 (GH, holotype; COL, F, US, 
isotypes). 

Small trees to 8 m. tall; sparingly pubescent. Branchlets stout, terete, glabrescent to 
abundantly pubescent with trichomes of setose to strigose types, trichomes upwards 
curved, gray to gray-brown, to 3 mm. long; black in dry state. Leaves clustered behind 
tip of branchlets; blades broadly elliptic to oblong-obovate, cuspidate to acuminate at 
apex with acumen to 10 mm. long, broadly cuneate to obtuse at base, serrate along 



SOEJARTO: SA URA VIA 49 

margins with serrulations rigid and sharp-pointed to prickly, 20-41 cm. long, 13-17 cm. 
wide, strongly coriaceous, extremely rigid and brittle, dark gray -brown above, light to 
ochraceous green beneath (in dry state dark olive-brown to dull gray above, dark 
yellowish to grayish brown beneath), subglossy and somewhat scabrous above, secon- 
dary veins strongly elevated, strongly conspicuous, (15-)20-25 pairs, tertiary veins 
elevated, strongly more prominent than lesser venation, glabrescent to scattered 
pubescent with trichomes of strigose to setose types especially along major veins above, 
glabrescent to scattered pubescent with trichomes of strigose to tuberculate-strigose 
(along major veins) mixed with clustered to stellate types (along and between minor 
veins) beneath; petioles robust, 2.5-5.5 mm. long, (2-H-7 mm. in diameter, sparingly 
setose pubescent. Inflorescences straight, many-branched, (75-)150-500-flowered, 
(12-)20-42 cm. long, 9-20 cm. wide, glabrescent to sparingly pubescent with trichomes 
of setose to strigose types, primary peduncle 5.5-15 cm. long, very stout and conspicu- 
ously woody, to 7 mm. in diameter, bracts linear-subulate to triangular, to 11 mm. 
long. Flowers 10-15 mm. broad, buds to 4 mm. in diameter, pedicels 1-2 mm. long, 
bracteoles linear to triangular, to 4 mm. long; sepals 5, elliptic to oblong-obovate, 
2.5-3.5 mm. long, 2-2.5 mm. wide, subacute to rounded, all glabrous throughout, 
marginally entire to ciliolate (especially at apex); petals 5, white to pinkish white, 
oblong, 5-7 mm. long, 3-4.5 mm. wide, obtuse; stamens 15-25, filament 2-3 mm. long, 
anther 1.5-2 mm. long; ovary 5-loculed, subglobose, 5-sulculate, glabrous, styles 5, 
obsolete to 3.5 mm. long, stigmas simple to subcapitate. Berries 5-loculed, subglobose, 
to 6 mm. across, 5-sulculate. 

Habitat. Humid mountain and submountain forest, at altitudes 
of (1,500-) 2,000-2,900 m. 

Distribution. Colombia (Departments of Valle, Cauca, Narino). 
Vernacular names. Moquillo (Narino: Mora). 

Specimens examined. COLOMBIA, VALLE: Cordillera Occiden- 
tal, Los Farallones, July, fr., Cuatrecasas 21716 (COL, F, GH, US), 
fl., 21802 (ECON, F); Rio Pichinde, between Quebrada de Juntas 
and El Recreo, Aug., fl., fr., Cuatrecasas 8321 (US). CAUCA: Macay 
Valley, La Gallera, July, fl., Killip 7932 (US; A, photo). NARINO: 
Ricaurte, road Chucunes to La Planada, June, fr., Mora 2662 
(ECON, PASTO). 

The most remarkable features of this species are: (1) the strongly 
coriaceous to extremely rigid leaves with very prominent venation 
beneath, (2) the many -branched, many -flowered glabrescent in- 
florescences, and (3) the flowers with glabrous sepals. S. micayensis 
is apparently related to S. paruiflora, from a lower mountain zone 
(1,000-1,500 m.), and S. pseudoleucocarpa, from the Colombian 
Pacific coastal zone (0-300 m.). All three species have glabrescent 
leaves and inflorescences, and small flowers with complete absence of 
unbranched multicellular trichomes on the sepals. 



50 FIELDIANA: BOTANY 

21. Saurauia natalicia Sleumer, Notizbl. 12: 144. 1934. Type: 
Weberbauer 7867 (F, lectotype; COL, photo). Figure 16. 

Saurauia pseudoparviflora Busc. var. rusbyana Busc., Malpighia 
30: 160. 1927. Type: Kuntze s. n. (NY, lectotype). 

Shrubs to 3 m. tall; glabrescent. Branchlets slender, terete, sparingly strigillose 
pubescent. Leaves clustered behind tip of branchlets; blades elliptic to obovate-elliptic, 
acuminate at apex with acumen to 20 mm. long, cuneate at base, serrate to serrulate 
along margins, 8-25 cm. long, 3-10 cm. wide, chartaceous to membranaceous, scarcely 
scabrous above, secondary veins (15-)20-30 pairs, tertiary veins elevated, more promi- 
nent than lesser venation, sparingly muricated to strigillose pubescent along and 
between minor veins and sparingly strigose pubescent along major veins above, spar- 
ingly pubescent with trichomes of tuberculate-strigillose (along minor veins) and 
strigose mixed with stellate types (along major veins) beneath, with pustulate epider- 
mis; petioles slender, terete, 1.5-3.5 cm. long, 0.5-1.5 mm. in diameter, sparingly 
strigillose pubescent. Inflorescences loose, sparingly to abundantly strigillose pubes- 
cent, primary peduncle slender, (2-)5-8 cm. long, bracts linear, to 5 mm. long. Flowers 
10-15 mm. broad, buds to 3.5 mm. in diameter, pedicels 1-5 mm. long, bracteoles linear, 
to 2 mm. long; sepals 5(-6), elliptic to oblong to obovate-oblong, acute to rounded, 3-4 
mm. long, 2-3 mm. wide, all glabrous throughout, marginally irregularly ciliolate; 
petals 5(-6), white, oblong to obovate-oblong, rounded to rarely deeply incised, 6-7 mm. 
long, 3.5-5.5 mm. wide; stamens 20-35, filament 3 mm. long, anther 1.25-1.75 mm. 
long; ovary 5(-6)-loculed, subglobose, 5(-6)-sulcate, glabrous, styles 5(-6), 1-4.5 mm. 
long, stigmas simple to subcapitate. Berries 5(-6)-loculed, globose to obovoid, to 7 mm. 
across, 5(-6)-sulcate. 

Habitat. Thicket and bushwood, at altitudes of 1,600-2,500 m. 
Distribution. Peru (Department of Cuzco) and Bolivia. 

Specimens examined. PERU, CUZCO: Calca, Lacco Valley, road 
to Huairurum, July, fl., Vargas 11099 (US); Quispicanchi, Mar- 
capata Valley, near Chilechile, Feb., fl., Weberbauer 7867 (F, lec- 
totype; COL, photo). AYACUCHO: Choimacota Valley, Feb., fl., 
Weberbauer 7567A (BM). 

BOLIVIA. NO PRECISE LOCALITY: San Miguelito, June, fl. fr., 
Herzog2238 (L, S, Z); Sta. Rosa, April, fl., Kuntze s. n. (NY). 

S. natalicia is closely related to S. glabra (R. & P.) Soejarto, differ- 
ing from it in the following characters: leaves without villous-barbate 
pubescence in the axils of secondary veins beneath, flowers with 
glabrous sepals, stamens low (20-30) in number. S. natalicia also 
appears to be allied to S. putumayonis, but the latter has larger and 
broader leaves, with sparingly distributed radiate to stellate 
trichomes on the lower surface, and flowers with pubescent sepals. 

22. Saurauia parviflora Tr. & PL, Ann. Sci. Nat. ser. 4, 18: 268. 
1862; Prodr. Fl. Nov. Granat. 1: 265. 1862. Type: Triana s. n. (BM). 



SOEJARTO: SAURA VIA 51 

Saurauia parviflora var. lehmanniana Busc., Malpighia 30: 81. 
1927. Type: Lehmann 9027 (K; GH, NY, photos). 

Shrubs to small tree to 5(-10) m. tall, sometimes bushy; glabrescent. Branchlets 
mostly slender, terete, glabrous to sparingly strigillose pubescent, somewhat smooth, 
dark brown to black in dry state, shoots and young leaves of same type of pubescence. 
Leaves clustered behind tip of branchlets; blades narrowly to broadly elliptic to obo- 
vate, shortly acuminate at apex with acumen to 10(-25) mm. long, cuneate to obtuse at 
base, serrulate to rarely serrate along margins, (10-)15-30(-40) cm. long, (5-)8-14 cm. 
wide, subcoriaceous, dark green above, light green beneath (in dry state dark, deep 
brown above, light to olive-brown beneath), glossy and smooth above, secondary veins 
(9-)16-20(-22) pairs, tertiary veins elevated, more prominent than lesser venation, 
glabrous to occasionally scattered strigose pubescent along midrib above, glabrescent 
with scattered strigose trichomes along major veins beneath; petioles (1.5-)2-4(-5.5) 
cm. long, 1.5-2 mm. in diameter, furrowed above, glabrous to glabrescent. Inflores- 
cences straight, (25-)40-100(-200)-flowered, (5-)12-27 cm. long, (2-)4-14 cm. wide, gla- 
brescent, primary peduncle (2-)3.5-7 cm. long, bracts linear, to 5 mm. long, rarely 
foliaceous, to 20 mm. long. Flowers 12.5-17 mm. broad, buds to 5 mm. in diameter, 
pedicels 0.5-1.5C-3) mm. long, bracteoles linear, to 5 mm. long; sepals 5, white to 
greenish white, outer two elliptic to oblong, obtuse, 3-4 mm. long, 2.5-3.5 mm. wide, 
imbricated one and inner two suborbicular to orbicular-obovate, obtuse, 5-6 mm. long, 
4.5-5.5 mm. wide, all glabrous throughout, marginally entire to ciliolate (especially 
along apex); petals 5, white, broadly oblong to oblong-obovate, usually incised to very 
rarely tridentate, 6-8 mm. long, 5-6 mm. wide; stamens 40-70, filament 2-2.5 mm. long, 
anther 1.25-1.75 mm. long; ovary 5-loculed, ellipsoid to globose, smooth to 5-sulculate, 
glabrous, styles 5, obsolete to 3.5(-4) mm. long, stigmas simple to capitate. Berries 
green, 5-loculed, globose, to 7 mm. across, 5-sulcate. 

Habitat. Wet mountain forest, mist zone, road cut, and clearing in 
wet mountain forest, at altitudes of (5-)100-l,500(-2,000) m. 

Distribution. Colombia (Departments of Valle, Cauca, Narino), 
mostly restricted along the western side of the Cordillera Occidental. 

Vernacular names. Moquillo (Narino: Checa). 

Specimens examined. COLOMBIA, VALLE: Pacific coast, Rio 
Yurumangui, El Aguacate, Feb., fl., Cuatrecasas 16114 (F), fl. fr., 
16144 (COL, GH, US; F, fragment). CAUCA: Rio Timbiqui, Nov., fl., 
Lehmann 9027 (COL, K; GH, NY, photos). NARINO: El Diviso, road 
to Tumaco, fr., Calvache 76 (ECON, PASTO); Ricaurte and vicinity, 
fr., Checa 17 (ECON, PASTO), Aug., fl., Soejarto 1444 (ECON, GH, 
PASTO); Barbacoas, between Junin and Altaquer, May, fl., Mora 
3059 (ECON, PASTO), fl., Soejarto 1450 (ECON, GH), 1453 (K), veg., 
1454 (ECON, GH); Altaquer, Alto de Cayambe, July, fl., Soejarto & 
Pinkley 945 (COL, ECON, GH), fl., 947 (COL, ECON, GH, PASTO); 
Tumaco, La Guayacana, July, fl., Soejarto & Pinkley 954 (ECON, 



52 FIELDIANA: BOTANY 

GH, PASTO); Buesaco, Santa Maria, July, fl., Recalde s. n. (ECON, 
PASTO); no. loc., fl., Corella s. n. (ECON, PASTO). NO PRECISE 
LOCALITY: Gneshi, fl., Lehmann BT-805 (K, NY); fl., Triana s. n. 
(BM). 

S. parviflora is abundantly represented in western Narino, in par- 
ticular in the area between Ricaurte and Tumaco. Field characters 
include subcoriaceous, glossy, glabrous to glabrescent leaves, with 
pale green to yellow veins, white to yellowish white inflorescences 
(peduncle and lateral cymes are of this color) with laxly distributed 
white flowers. The flowers are often infected by insects (weevils), 
which burrow their way in before the flowers open. This causes such 
infected flowers to decay and turn black, making them conspicuous. 

The range of S. parviflora overlaps with that of S. peduncularis in 
the Ricaurte region, and plants of both species may be found growing 
in the same place. Natural hybrids may have been produced in such 
cases, and several collections of this nature have been treated as S. 
peduncularis, due to the stronger influence of this species. 

23. Saurauia pseudoleucocarpa Busc., Malpighia 29: 342. 
1922. Type: Lehmann BT-1173 (G, lectotype; A, F-fragment, K, 
NY, isolectotypes; COL, photo). Illustration: Malpighia 29: pi. 7, fig. 
12. 1922. 

Saurauia micans R. E. Schultes, ined. 

Small trees to 10 m. tall; glabrescent. Branchlets slender, terete, ashy to dark brown 
in dry state, sparingly pubescent with trichomes of tuberculate-strigose to setulose 
types. Leaves crowded around tip of branchlets; blades elongate-elliptic to elongate- 
obovate, acuminate at apex with acumen to 20 mm. long, cuneate at base, distantly 
serrulate with fine and sharp-pointed serrulations along margins, (10-)15-28(-30) cm. 
long, (3-)5-10(-12) cm. wide, chartaceous, in dry state dark dull brown to sooty above, 
light to dark reddish brown beneath, glossy above, secondary veins 20-30 pairs, 
tertiary veins elevated, more prominent than lesser venation, glabrous but occasion- 
ally with strigose to strigillose trichomes along major veins above, scattered pubescent 
with setose to strigose (along midrib) and radiate trichomes (along and between veins) 
beneath; petioles 2-3(-3.5) cm. long, 1.5-2 mm. in diameter, sparingly strigose to setose 
pubescent. Inflorescences straight, (20-)50-ca. 150-flowered, (5-) 10-20 cm. long, 3-10 
cm. wide, sparingly pubescent with setose to strigose mixed with radiate to clustered 
trichomes, primary peduncle slender, 1-10 cm. long, bracts linear, to 20 mm. long. 
Flowers 7-10 mm. broad, buds to 3.5 mm. in diameter, pedicels extremely slender, to 3 
mm. long, bracteoles linear, to 3 mm. long; sepals 5, narrowly elliptic to obovate, 
obtuse, 2.5-3.5 mm. long, 1.5 mm. wide, exposed parts in bud sparingly radiate pubes- 
cent, imbricated parts glabrous, all glabrous inside, marginally ciliolate; petals 5, 
white, spatulate to elongate-obovate, rounded, 4-6 mm. long, 2-2.5 mm. wide; sta- 
mens 15-20, filament 2 mm. long, anther 1.5 mm. long; ovary 5-loculed, obovoid, 
5-sulculate, glabrous, styles 5, obsolete to 3.5 mm. long, stigmas simple to subcapi- 
tate. Berries 5-loculed, obovoid to globose, to 4 mm. across, 5-sulcate. 



SOEJARTO: SA URA VIA 53 

Habitat. Rain forest, at altitudes of 0-300 m. 

Distribution. Colombia (Departments of Choco, Valle, Cauca), 
restricted along the Pacific coastal zone. 

Specimens examined. COLOMBIA, CHOCO: Off San Juan river, 
Quebrada El Taparal, riverbank, May, fr., Cuatrecasas 21494 
(ECON, F, K, US), Aug., fl., Hugh-Jones 309 (K); Rio Calima in the 
Choco region, between Quebrada de Aguaclara and Quebrada La 
Brea, May, fl., LehmannBT-1173 (A, F-fragment, G, K, L, NY; COL, 
GH, photos). CAUCA: Cordoba, Dagua Valley, Pacific coastal zone, 
Dec., fl., Pittier 545 (GH, NY, US). 

24. Saurauia spectabilis Hort. ex Hook., Bot. Mag. 69: no. 3982. 
1842. Type: Ex icon. Illustration: Bot. Mag. 69: pi. 3982. 1842. 
Saurauia brevipes Rusby, Descr. 300 New Sp. S. Am. PI. 57. 

1920. Type: Bang 387B (NY, S, UC) (specimen in NY 

labeled as Bang s. n. ). 
Saurauia aequatoriensis Sprague var. boliviano Busc., Malpighia 

30: 35. 1927, p. p. 

Saurauia rusbyi auct. non Britt.: Busc., 1. c. 173. 1927 (S. rusby). 
Saurauia rusbyi auct. non Britt. var. glabrata Busc., 1. c. 175. 

1927. Type: Bang 387B. 
Saurauia rusbyi auct. non Britt. var. spectabilis Busc., 1. c. 178. 

1927. Type: Pentland 116 (P). 
Saurauia rusbyi auct. non Britt. var. spectabilis fma. mac- 

rophylla Busc., 1. c. 181. 1927. Type: Bang 387B. 
Saurauia rusbyi auct. non Britt. var. spectabilis fma. ueranii 

Busc., 1. c. 184. 1927. Type: Rusby 481 A (MICH, NY, US). 

Shrubs to small trees to 10 m. tall, glabrescent to sparingly pubescent. Branchlets 
sparingly pubescent, trichomes strigose to sericeous, ferruginous. Leaves crowded 
around tip of branchlets; blades obovate to elongate-obovate, acuminate at apex with 
acumen to 15 mm. long, narrowly cuneate to subrotundate at base, often oblique, 
serrate to serrulate (biserrate, cf. Hooker) along margins, (13-)18-30(-35) cm. long, 
(4.5-)5-10(-11.5) cm. wide, chartaceous, in dry state dark olive-brown to dark brown 
above, light olive- to gray-brown beneath, glossy to subglossy above, secondary veins 
(15-)18-25(-28) pairs, tertiary veins elevated, more prominent than lesser venation, 
glabrescent to scattered pubescent with trichomes of sericeous-strigillose type along 
veins above, scattered to sparingly pubescent with trichomes of sericeous-strigose 
type along veins with pustulate epidermis beneath; petioles l-3(-3.75) cm. long, 
2.5-3.5 mm. in diameter, abundantly pubescent with sericeous-strigose trichomes. 
Inflorescences straight, many-branched, spreading, (50-)100-300(-500)-flowered, 
(12.5-)18-35(-42) cm. long, (6-)9-15(-21) cm. wide, abundantly pubescent with 
sericeous-strigose trichomes, primary peduncle (5-)8-12(-14.5) cm. long, bracts linear 
to triangular, 2-10 mm. long. Flowers 10-19 mm. broad, buds to 7 mm. in diameter, 



54 FIELDIANA: BOTANY 

pedicels 5-10 mm. long, bracteoles triangular, 2-3 mm. long; sepals 5, ovate to subor- 
bicular, obtuse, 4-7 mm. long, 3-5 mm. wide, exposed parts in bud sparingly strigose 
pubescent, imbricated parts glabrous, all glabrous inside, marginally ciliate; petals 5, 
spatulate to obcordate, slightly to deeply incised, 6-8 mm. long, 5-7 mm. wide; sta- 
mens (45-)65-85, filament 2-3 mm. long, anther 1-1.65 mm. long; ovary 5-loculed, 
globose, 5-sulcate, glabrous, styles 5, obsolete to 5(-6) mm. long, stigmas simple to 
capitate. Berries immature. 

Habitat. Mountain forest, garden, and roadside, at altitudes of 
750-2,500 m. 

Distribution . Bolivia (Province of La Paz). 

Specimens examined. BOLIVIA, LA PAZ: Unduavi Valley, fl., 
Badcock 94 (K), Julio 467 (US); Larecaja, vie. of Mapiri, Jan., fl., 
Bang 1742 (A, F, G, GH, K, MICH, S, US), fl., Buchtien 1849 (US), 
Sept., fr., 1992 (US), April, Rusby 481 (F, G, K, NY, P, US); road 
Tipuani, hacienda Simaco, Feb., fl., Buchtien 5461 (BM, F, G, GH, 
K, NY, P, US, Z); Tipuani Valley, hacienda Casana, Jan., fl., Buch- 
tien 7377 (P, S, US), fl., Weddells. n. (P, sheet no. 87-29-66); vie. of 
Onanea, Cerro de Uacani, May, fl., Mandon 832 (K, P); North Yun- 
gas, Coroico, fl., Pentland 116 (P); Yungas, fl., Rusby 481A (MICH, 
NY, US); basin of Rio Bopi, San Bartolome, near Calisaya, July, fl., 
Krukoff 10356 (A, F, G, K, NY, S, U), fl. fr.,Bang387B (K, L, NY, S, 
UC). NO PRECISE LOCALITY: Pearce s. n. (K); from cultivation, 
Kew Garden, fl., Stove s. n. (K). 

W. J. Hooker (1842) described S. spectabilis on the basis of a plant 
"raised by Mr. Knight, of the Exotic Nursery, King's Road, Chelsea 
(England), from seeds, imported from the Republic of Bolivia, in 
1838." As stressed by Hooker, the species is characterized by the 
narrow elongate (obovate-lanceolate) leaves and the large, spread- 
ing, many -branched and many-flowered inflorescences. 

V. Ser. PULVERULENTAE Busc., Malpighia 25: 222. 1912, stat. 
nov. 

Tomentosae Busc., 1. c. 220., p. p. 
Laevigatae Busc., 1. c. 224., p. p. 

1. Stamens 20-50, at most not more than 80. 

2. Flowers 20 mm. broad or larger, at least not less than 18 mm. broad, when open. 
3. Sepals subglabrous outside, glabrous on the lower two-thirds to three-fourths 
and sparingly pulverulent on the upper one-third to one-fourth inside, in- 
florescences 10-30(-50)-flowered, pedicels 1.5-4 mm. long ... 25. S. pulchra 
3. Sepals pulverulent and tuberculate pubescent outside, pulverulent inside, 



SOEJARTO: SA URA UIA 55 

inflorescences (25-)40-100-flowered, pedicels 0.25-1 mm. long (subsessile). 

26. S. aromatica 

2. Flowers 10 mm. broad, at most not more than 18 mm. broad, when open. 
4. Leaves glabrescent to sparingly pubescent, floccose trichomes predominate. 

5. Blade length to width ratio of (1.2-)2(-2.5) to 1, smooth above, floccose 
trichomes scattered throughout, epidermis not pustulate beneath, sta- 
mens 20-30 27. S. floccifera 

5. Blade length to width ratio of (2-)3(-4) to 1, somewhat scabrous above, 
floccose and radiate trichomes scattered beneath, epidermis pustulate 
beneath, stamens (30-)35-50(-80) 28. S. choriophylla 

4. Leaves abundantly to sparingly pubescent, tufted and dendroid trichomes 
predominate. 

6. Blades 20-50 cm. long, 6-25 cm. wide, flowers 10-13 mm. broad; stellate, 
dendroid, and tufted trichome types predominate . . 29. S. cuatrecasana 

6. Blades 15-30 cm. long, 6-12 cm. wide, flowers 15-18 mm. broad; radiate 
and minute stellate trichome types predominate 30. S. arnoldi 

1. Stamens 100-ca. 200, but not less than 80. 

7. Leaf pubescence dense and velvety beneath, with trichomes of dendroid and 
stellate types, lower epidermis obscured by pubescence . . 31. S. tomentosa 

7. Leaf pubescence sparser and not velvety beneath, with trichomes of other 
types but not dendroid nor stellate, lower leaf epidermis not obscured by 
pubescence. 

8. Flowers 15 mm. broad, pedicels less than 5 mm. long; 1,200-2,000 m. 

altitudes 32. S. pseudostrigillosa 

8. Flowers 20-25 mm. broad, pedicels 5-12 mm. long; ca. 100 m. altitude. 

33. S. mexiae 

25. Saurauia pulchra Sprague, Trans. & Proc. Bot. Soc. Edin- 
burgh 22: 426. 1904. Type: Sprague 323 (K, COL, NY, US, photos). 
Figure 17. 

Small trees to 9 m. tall, diameter to 30 cm. at base, bark gray-brown, smooth, trunk 
crooked; sparingly pubescent. Branchlets moderately thick, terete, pulverulent, and 
tuberculate pubescent. Leaves crowded around tip of branchlets; blades oblong- 
obovate, rounded to very shortly and abruptly acuminate at apex, cuneate to obtuse at 
base, distantly serrate to serrulate along margins, 11-25C-35) cm. long, 5-12.5(-16) cm. 
wide, firmly coriaceous, in dry state dark brown above, dull brown beneath, glossy and 
somewhat smooth above, secondary veins 12-24 pairs, tertiary veins irregular, some- 
what elevated, scarcely more prominent than lesser venation, scattered tuberculate 
pubescent along and between veins above, scattered pubescent with trichomes of 
radiate to stellate or clustered (along veins) mixed with tuberculate-strigose types 
(along midrib) beneath; petioles 1-1. 5 cm. long, 1.5-2 mm. in diameter, furrowed above, 
glabrescent. Inflorescences often ascending, somewhat straight, 10-30(-50)-flowered, 
6-20 cm. long, 3-10 cm. wide, glabrescent to pulverulent along lesser ramification and 
pedicels, primary peduncle slender, 1-7 cm. long, bracts triangular to linear, to 6 mm. 
long. Flowers laxly distributed, 20-25 mm. broad, buds to 6 mm. in diameter, pedicels 
robust, to 3 mm. (4 mm. in fruiting state) long, bracteoles triangular, to 3 mm. long; 
sepals 5, broadly elliptic to orbicular-ovate, obtuse to rounded, 7-10 mm. long, 5-8 



56 FIELDIANA: BOTANY 

mm. wide, outer two scattered to abundantly minutely stellate pubescent (especially 
medially), imbricate one and inner two glabrous on exposed parts in bud, sparingly to 
abundantly minutely stellate pubescent on imbricated parts, all glabrous on lower 
two-thirds to three-fourths and sparingly to abundantly minutely stellate pubescent 
on upper one-third to one-fourth inside, marginally irregularly ciliolate; petals 5, 
white, oblong-obovate to broadly spatulate, rounded, 9-14 mm. long, 7-10 mm. wide; 
stamens 25-45, filament 3-4.5 mm. long, anther 3-3.5 mm. long; ovary 5(-7)-loculed, 
subglobose, 5(-7)-sulcate, glabrous, styles 5(-7), 0.8-5 mm. long, stigmas simple to 
capitate. Berries 5(-7)-loculed, globose, to 10 mm. across, 5(-7)-sulcate. 

Habitat. Wet mountain and submountain forest, moist soil, at 
altitudes of 1,600-2,300 m. 

Distribution. Colombia (Departments of Cauca and Huila). 

Vernacular names. Moco (Huila: Sprague), Moquillo (Huila: 
Schultes & Villarreal). 

Specimens examined. COLOMBIA, CAUCA: Near Cauca-Huila 
border, Inza, Nov.,ft.,Lehmann2192 (BM, K, US); Tierra Dentro, Rio 
Paez Valley, Indian Village, Jan., fl., Pittier 1276 (US). HUILA: 
Cordillera Oriental, 25 km. NE. of Algeciras, between Vega Temopila 
and Altagracias, March, fl. fr., Little, Jr. 7540 (NY, US); west-facing 
slope of the Cordillera, above Guadalupe, Resina, March, fl., Perez- 
Arbelaez & Cuatrecasas 8367 (COL, US); E. of Neiva, Aug., fl. fr., 
Rusby & Pennell 598 (NY); "Balsillas" on Rio Balsillas, Aug., fl. fr., 
Rusby & Pennell 712 (NY, US; GH, photo); San Agustin and vie., 
Nov., fl.,Romero-Castaneda 6566 (COL), Jan., fl., Schultes & Villar- 
real 5330 (COL, ECON), March, fl., Sprague 323 (K; COL, NY, US, 
photos). 

26. Saurauia aromatica R. E. Schultes, Caldasia 2: 28. 1943. 
Type: Cuatrecasas 9105 (F, US). Illustration: Caldasia 2: 29. 1943; 
Bot. Mus. Leafl. Harvard Univ. 20: 223, pi. 3. 1963. 

Saurauia echinosepala R. E. Schultes, 1. c. 34. 1943. Type: 

Perez-Arbelaez & Cuatrecasas 6473 (US, holotype; F, iso- 

type). 
Saurauia alvaroi R. E. Schultes, Bot. Mus. Leafl. Harvard Univ. 

20:221. 1963. Type: Schultes 22551 (ECON, holotype; COL, 

F). 

Small trees to 15 m. tall, diameter to 10 cm. at base, bark gray-brown, lightly 
fissured; sparingly pubescent. Branchlets robust, terete, sparingly tuberculate to 
tuberculate-strigose pubescent. Leaves clustered behind tip of branchlets; blades obo- 
vate to elongate-obovate, broadly and very shortly acuminate at apex, cuneate to 
obtuse but rarely oblique at base, frequently with well-developed basal flap, distantly 
serrulate to serrulate-denticulate along margins, 16-45 cm. long, 8-20 cm. wide, char- 



SOEJARTO: SA URA UIA 57 

taceous to coriaceous, glossy and smooth and dark green above, light green beneath, 
secondary veins (15-)20-27(-33) pairs, strongly prominent, tertiary veins elevated, 
more prominent than lesser venation, glabrescent to tuberculate or tuberculate- 
strigose pubescent along and between veins above, scattered pubescent with trichomes 
of stellate to radiate types (along and between veins) mixed with strigose to 
tuberculate-strigose types (along midrib) beneath; petioles robust, 3-7 mm. in diame- 
ter, furrowed above, tuberculate to tuberculate-strigose pubescent and/or lenticellate, 
often with minute floccose to clustered trichomes. Inflorescences straight, often erect, 
25-100-flowered, 15-30 cm. long, 7-15 cm. wide, densely pubescent with trichomes of 
shaggy- to tufted-tuberculate or -echinulate types and rusty brown in color, primary 
peduncle stout and woody, 7-15 cm. long, bracts linear, to 7 mm. long, rarely foliace- 
ous, to 20 mm. long. Flowers 17-22 mm. broad, buds to 6 mm. in diameter, pedicels 
0.25-1 mm. long (subsessile), bracteoles linear to triangular, to 2 mm. long; sepals 5, 
orbicular-ovate to oblong or subquadrangular, pale green to white with rusty brown 
pubescence, obtuse, 5.5-8 mm. long, 4-6 mm. wide, all pulverulent throughout, 
trichomes sparingly distributed, of shaggy-tuberculate to -echinulate on exposed 
parts in bud, marginally ciliolate; petals 5, white, oblong-obovate to oblong or sub- 
quadrangular, rounded to somewhat truncate, 7-9 mm. long, 3-6 mm. wide; stamens 
30-50, filament 2-2.5 mm. long, anther 2.5-3 mm. long; ovary 5-loculed, subglobose, 
5-sulcate, glabrous, styles 5, obsolete to 6 mm. long, stigmas simple to capitate. 
Berries 5-loculed, globose, to 7 mm. across, 5-sulcate. 

Habitat. Wet rain forest, submountain forest, open forest, river- 
banks, border of sugarcane plantation, road embankment, cultiva- 
tion areas, thickets and gardens, at altitudes of 500-2,600 m. 

Distribution. Colombia (Departments of Caldas, Huila, Caqueta, 
and Putumayo). 

Vernacular name. Moquillo (Putumayo: Soejarto). 

Specimens examined. COLOMBIA, CALDAS: Highway of 
Quindio, between La Gabriela and La Linea, July, fl., Perez- 
Arbelaez & Cuatrecasas 6473 (F, US). HUILA: Archeological Park, 3 
km. W. of San Agustin, April, fl., Little, Jr. & Little 7616 (NY, US). 
CAQUETA: Cordillera Oriental, east facing slope, Quebrada del Rio 
Hacha, Cajon de Pulido, March, fl., Cuatrecasas 8722 (US); Sucre, 
April, fl., Cuatrecasas 9105 (F, US); between Sucre and La Portada, 
April, fl., Cuatrecasas 9150 (F, US). PUTUMAYO: Quebrada del Rio 
Mulato, Dec., fl., Cuatrecasas 11282 (F, US); road from San Fran- 
cisco to Mocoa, 10 km. above Pepino, July, fl., Schultes 22551 
(ECON, F, GH); Pepino of Mocoa and vie., Aug., fl. fr., Soejarto 523, 
524, 528, 534, 544 (all in ECON, GH); Road Sibundoy to Pepino of 
Mocoa, La Mesa and vie., Aug., fl. fr., Soejarto 1575-1578, 1582, 
1583, 1586 (all in ECON, GH), veg., 1579, 1580 (both in ECON), 
veg., 1584, 1585, 1587, 1588, 1590 (all in ECON, GH). 

When Schultes (1963, p. 222) described S. alvaroi, he noted that 



58 FIELDIANA: BOTANY 

"the species may be distinguished from all other known concepts of 
the genus by a most curious canal-like pouch which is formed by two 
conspicuous vexilliform flaps arising perpendicularly from the base 
of the leaf blade along the midrib and which are joined together at 
their apical and basal ends." Field studies, however, could not con- 
firm the taxonomic importance of this basal flap, since it occurs 
rather widely, though not frequently, in many other unrelated 
species. Among others, such a flap was recorded in S. tomentosa, S. 
brachybotrys, S. cuatrecasana, S. caquetensis, and S. pseudostrigil- 
losa. Furthermore, members of the population of S. aromatica from 
the same locality as that of the type collection, in the Mocoa region, 
possess both flap- and non-flap-bearing leaves. Also, flap- and non- 
flap-bearing leaves may even be found in the same tree. A series of 
herbarium collections of S. aromatica and S. brachybotrys, from the 
Narino and Putumayo regions, establish that the basal flap is actu- 
ally formed by the continuation of the leaf margins, probably a 
product of abnormal development. This structure is so well de- 
veloped in certain individuals that it is boat-like in shape. 

27. Saurauia floccifera Tr. & PI., Ann. Sci. Nat. ser. 4, 18: 267. 
1862; Prodr. Fl. Nov. Granat. 1: 264. 1862. Type: Triana s. n. (G, 
lectotype; F-fragment, K, P, US, isolectotypes; GH, photo). Figure 18. 

Small trees to 10 m. tall, crown spreading; copiously pubescent. Branchlets stout, 
terete, dark grayish brown to sooty in dry state, sparingly pubescent; trichomes tufted 
and dendroid to dendroid-tufted. Leaves crowded behind tip of branchlets; blades 
broadly elliptic to obovate, acute to acuminate to cuspidate at apex with acumen to 10 
mm. long, narrowly to broadly cuneate at base, rarely oblique, serrulate to distantly 
serrate with fine serrations along margins, (8-)10-25(-30) cm. long, (6-)8-16 cm. wide, 
chartaceous to rarely membranaceous, in dry state dark reddish brown to sooty above, 
dull gray-brown beneath, smooth and glossy above, secondary veins (18-)23-30(-33) 
pairs, tertiary veins elevated, more prominent than lesser venation, scattered floccose 
pubescent above, scattered pubescent with trichomes of floccose (along veins) mixed 
with tufted to dendroid-tufted types (along midrib) beneath; petioles 1.5-3 cm. long, 
2-3 mm. in diameter, scattered to sparingly floccose pubescent. Inflorescences 
straight, (25-)30-100(-ca. 150)-flowered, 9-24 cm. long, 3.5-8.5 cm. wide, lower parts 
sparingly floccose pubescent, upper parts and lesser ramification pulverulent, 
trichomes of clustered and dendroid to tufted types, primary peduncle 3-12 cm. long 
(16 cm. long, cf. Schultes), bracts linear to triangular, to 8 mm. long. Flowers 10-15 
mm. broad, buds to 3-5 mm. in diameter, pedicels to 4 mm. long, bracteoles triangu- 
lar, usually upwards curved, to 2 mm. long; sepals 5, white to pale green, elliptic to 
obovate, obtuse, 4-5 mm. long, 2.5-3.5 mm. wide, all pulverulent throughout, and 
densely pubescent with trichomes of stellate, clustered, tufted, and dendroid types on 
exposed parts in bud, but of stellate type on imbricated parts, marginally ciliolate, 
especially on apex; petals 5, white, obovate to oblong, rounded, 4.5-6 mm. long, 3.5-5 
mm. wide; stamens 20-30, filament 2.5 mm. long, anther 2 mm. long; ovary 5-loculed, 



SOEJARTO: SA URA VIA 59 

ovoid to subglobose, 5-sulculcate, glabrous, styles obsolete to 3.5 mm. long, stigmas 
simple to subcapitate. Berries 5-loculed, obovoid to subglobose, to 6 mm. across, 
5-sulcate. 

Habitat. Rain forest, steep wooded slope, rich soil, sugarcane 
field, secondary forest, wet forested slopes, and mist-covered zone, at 
altitudes of 600-2,000 m. 

Distribution. Venezuela (State of Tachira) and Colombia (De- 
partments of Choco, Antioquia, Norte de Santander, Cundinamarca 
and Meta). 

Vernacular names. Dulumoco (Choco: Archer), Chupahuevo 
(Cundinamarca: Triana). 

Specimens examined. VENEZUELA, TACHIRA: Alto de Lirio, 
between Bramon and Las Delicias, July, fl. fr., Steyermark 57460 (F, 
NY). NO PRECISE LOCALITY: Alicia del Rio Aponcito, Tulia, Jan., 
fl., Vareschi 3113 (YEN). 

COLOMBIA, NORTE DE SANTANDER: Sarare region, basin of 
Margua River, between Junin and Cordoba, Nov., fl., Cuatrecasas 
13367 (US). CHOCO: Between La Oveja and Quibdo, April, fl., 
Archer 1733 (US). CUNDINAMARCA: Ubala, Aug., fl., Triana 
5414 (BM, COL, US); Caqueza, canyon of Rio Negro, halfway be- 
tween Guayabetal and Serovita, Aug., fl., Fosberg & Bellis 22066 
(NY, US); road Guayabetal to Quetame, Monteredondo, June, fl., 
Uribe-Uribe 4842, 4843 (both in COL, ECON); Guayabetal, Aug., fl., 
Richters. n. (GH, US); SE. of Quetame, Susumuco, Sept., fr.,Pennell 
1741 (NY); Quetame, Aug., fl., Schultes 5684 (COL, F, GH, K, NY, 
US), Triana s. n. (G, K, P; F, fragment; GH, US, photos). META: 
Horse trail between Guayabetal and Acacias, Rio Manzanares, 
Aug., fl., Garcia-Barriga 15397 (COL, ECON); vie. of Villavicencio, 
Jan., fl., Naught 2554 (A, COL, S, US). 

Like S. tomentosa, members of S. floccifera are characteristically 
beset with multi-cellular branched trichomes, which are, in the case 
of the latter, early caducous (floccose), particularly along the 
branchlets, the petioles, the primary peduncle and the leaf surfaces, 
thus leaving these organs glabrescent to sparingly pubescent when 
mature. However, S. floccifera differs greatly from S. tomentosa in 
floral characters, especially in flower size and stamen number. Also, 
other aspects of pubescence in these two species are different. 

As to the ecology of the species, the following remarks by Schultes 
(1945, p. 252) on the species population in the Quetame area are 



60 FIELDIANA: BOTANY 

noteworthy: "A careful examination of Saurauia floccifera in the 
gorge through which the highway runs indicates that the leaves are 
rather generally smaller than those of the type. It is my belief that the 
type may have been collected from a forest where the plant grew in a 
shaded situation. At the present time, the hills along this gorge are 
partly, or, in many places, completely deforested, and the individuals 
of Saurauia floccifera are usually found in isolated areas on grassy 
slopes. Near the lower range of the species, there are a number of 
individuals which occur in the forest, and it was noted that the leaves 
of these tend to be larger and thinner (cf. O. Haught 2554 ). Similarly, 
the collection of L. Richter s. n., from a relatively low altitude in the 
same general region, has larger, more membranaceous leaves." 
Further, from the field notes of collectors, it appears that S. floccifera 
may be easily noted in the field by its showy white inflorescence. 

28. Saurauia choriophyla R. E. Schultes & Gutierrez, Caldasia 
3: 251. 1945. Type: Toro 845 (MEDEL, holotype; NY, isotype). 

Trees to 20 m. tall, diameter to 40 cm. at base, bole somewhat straight, bark fissured, 
brown, crown spreading to somewhat conoidal; glabrescent. Branchlets terete, dis- 
tinctly scarred, pulverulent, abundantly to densely pubescent with trichomes of tuber- 
culate to shaggy-strigillose types, rusty brown in color. Leaves crowded behind tip of 
branchlets; blades oblong-obovate to elongate-obovate but rarely oblong to elongate 
elliptic, acute to shortly acuminate at apex, with acumen to 10 mm. long, cuneate to 
rarely oblique at base, subentire to serrulate with very fine serrulations along mar- 
gins, 15-25C-30) cm. long, 4.5-10(-12) cm. wide, subcoriaceous to chartaceous, some- 
what scabrous throughout, in dry state sooty above, dark to light dull brown beneath, 
secondary veins 23-30 pairs, tertiary veins elevated, more prominent than lesser 
venation, glabrescent with pustulate epidermis above, sparingly pubescent with floe- 
cose and radiate trichomes, and with pustulate epidermis beneath; petioles 2-5 cm. 
long, 1-2 mm. in diameter, half-terete, pulverulent. Inflorescences straight, 20-100- 
flowered, 5-20 cm. long, 3-8 cm. wide, lower parts glabrescent, upper parts and along 
lesser ramification densely pubescent, trichomes of shaggy-strigillose type, yellowish 
to rusty-brown in color, primary peduncle 3-8 cm. long, bracts linear, to 5 mm. long. 
Flowers 10-15 mm. broad, buds to 5 mm. in diameter, pedicels to 3 mm. long, bracteoles 
triangular to linear, to 1.5 mm. long; sepals (4-)5, suborbicular to oblong or ovate, 
rarely spatulate, acute to rounded, 4-5 mm. long, 2.5-5 mm. wide, exposed parts in bud 
abundantly to densely pubescent with trichomes of radiate to shaggy or shaggy- 
tuberculate types, imbricated parts densely stellate pubescent, all densely stellate 
pubescent inside, marginally ciliolate; petals (4-)5, white, oblong-obovate, 5-7 mm. 
long, 4.5-5 mm. wide; stamens (30-)35-50(-80), filament 2.5-3 mm. long, anther 1.5-2 
mm. long; ovary (4-)5-loculed, globose, (4-)5-sulcate, glabrous, styles (4-)5, obsolete to 
4.5 mm. long, stigmas simple to capitate. Berries unknown. 

Habitat. Secondary forest, fringe of forest, wet forested river- 
bank, humid coffee plantation and waysides, at altitudes of 1,000- 
1,800 m. 



SOEJARTO: SA URA VIA 61 

Distribution. Colombia (Departments of Antioquia, Caldas, To- 
lima, and Valle). 

Vernacular name. Dulumoco (Valle: Herrera). 

Specimens examined .COLOMBIA, ANTIOQUIA: Pueblorico, 
June, fl., Espinal 79 (MEDEL); Venecia, June, fl., Soejarto & Rivera 

2054 (ECON, GH, MEDEL); Fredonia, June, fl., Soejarto & Rivera 

2055 (ECON, GH, MEDEL), Dec., fl., Toro 845 (MEDEL, NY). CAL- 
DAS: Chinchina, Cuatrecasas 23375 (F, GH); NE. of Armenia, July, 
fl., Pennell et al. 8666 (GH, NY, US). TOLIMA: El Fresno, Alto de 
Aguila, Dec., fl., Garcia-Barriga 8227 (COL, US); El Libano, La 
Trinidad, Dec., fl., Pennell 3306 (NY, US). VALLE: Cordillera Occi- 
dental, basin of Rio Calima, between Darien and Mediacanoa, El 
Cairo, Jan., fl., Cuatrecasas 13892 (F, GH); Sevilla, June, fl., Herrera 
922 (US). 

S. choriophylla is closely allied to S. floccifera, but the former has 
more elongate, pustulate, and somewhat scabrous leaves (particu- 
larly the lower surface), with a higher stamen number (35 or more, 
but less than 80), in contrast to the broad, smooth, and somewhat 
glossy (above), and non-pustulate leaves, with a lower stamen 
number (20-30), in the latter. 

29. Saurauia cuatrecasana R. E. Schultes, Caldasia 2: 315. 
1944. Type: Cuatrecasas 9209 (COL, holotype; F, US, isotypes). 
Figure 19. 

Trees to 10(-30) m. tall, diameter to 20(-40) cm. at base, bark thick, brittle, irregu- 
larly cracked, light gray-brown, living bark ochre in color, wood soft, with ochre-rosy 
tint, crown spreading; copiously pubescent. Branchlets stout, robust, terete, distinctly 
scarred, mealy white to yellowish, densely scurfy pubescent with tuberculate 
trichomes, scabrous. Leaves clustered behind tip of branchlets; blades broadly obovate 
to elliptic to oblong, acute to obtuse but rarely very shortly acuminate at apex, cuneate 
to obtuse at base, often oblique, rudimentary basal flap occasionally present, serrulate 
with very fine serrulations along margins, 20-4CK-50) cm. long, (6-)10-20(-25) cm. wide, 
chartaceous, scabrous above and beneath, in dry state dark brown to sooty above, light 
reddish to olive-brown beneath, secondary veins 23-30(-33) pairs, tertiary veins ele- 
vated, more prominent than lesser venation, scattered to sparingly pubescent with 
trichomes of tuberculate (along and between major veins) to strigillose types (along 
midrib) above, densely pubescent with trichomes of stellate to shaggy or dendroid or 
dendroid-tufted types and deep reddish violet in color beneath (scurfy in appearance 
along major veins); petioles robust, 3-10(-13) cm. long, 2-5 mm. in diameter, somewhat 
furrowed above, densely to abundantly scurfy pubescent with tuberculate trichomes. 
Inflorescences straight and often spreading, (50-)100-(more than)300-flowered, 15- 
30(-35) cm. long, (3-)7-15 cm. wide, densely pubescent with trichomes of shaggy to 
tufted types, mealy and yellowish to rusty brown in color, primary peduncle 4-16 cm. 
long, bracts linear to linear-triangular, to 8 mm. long. Flowers 10-13 mm. broad, buds 



62 FIELDIANA: BOTANY 

to 4 mm. in diameter, often obovoid, pedicels to 2(-3) mm. long, bracteoles minute, 
linear -triangular, to 1-5 mm. long; sepals 5(-6), pale yellowish green to white, oblong 
to oblong-elliptic to suborbicular, subacute to rounded, 3-4.5 mm. long, 2-3.5 mm. 
wide, all densely scurry pubescent with trichomes of shaggy-tuberculate type on 
exposed parts in bud and of radiate to stellate types on imbricated parts, and densely 
stellate pubescent (sometimes limited to lower one-fourth only) inside, marginally 
ciliolate; petals 5, white, oblong to oblong-obovate, rounded and rarely somewhat 
incised, 5-6 mm. long, 2.5-3.5 mm. wide; stamens 25-40, filament 2.5-3 mm. long, 
anther 1.5 mm. long; ovary 5(-6)-loculed, globose, 5(-6)-sulculate, glabrous, styles 
5(-6), obsolete to 4 mm. long, stigmas simple to capitate. Berries unknown. 

Habitat Forest, fringe of forest, open hillsides, submountain 
forest, at altitudes of 1,900-2,900 m. 

Distribution. Colombia (Departments of Caldas, Tolima, Valle, 
Cauca, and Huila). 

Specimens examined. COLOMBIA, CALDAS: Vic. of Manizales, 
Gallinazo, May, fl., Cuatrecasas 9209 (COL, F, US); Cordillera Cen- 
tral, Otun River basin, between Peria Bonita and Las Delicias, Nov., 
fl., Cuatrecasas 23352 (ECON, F, GH); Miraflores, on road Manizales 
to Nevado del Ruiz, June, fl., Soejarto 2030 (ECON, GH), July, fr., 
Pinto 423 (COL). TOLIMA: Along Quindio highway, between 
Cajamarca and summit of Divide, March, fl., Killip & Varela 34547 
(BM, US). CAUCA: Cordillera Central, headwaters of Rio Palo, 
Quebrada de Santo Domingo, Dec., fl., Cuatrecasas 19172; Rio Mun- 
chique, July, fl., Garcia-Barriga et al. 12951 (US); Alto de Punicias, 
near Jambolo, Rio Palo basin, Tierra Dentro, Feb., fl., Pittier 1457 
(US). HUILA: Road to La Plata, region of Moscopan, Santa Leticia, 
July, fl., Garcia-Barriga & Hawkes 12871 (US). 

30. Saurauia arnoldi Sleumer, Notizbl. 12: 143. 1934. Type: 
Schultze 814 (not seen), ex descr.; Kernan 125 (NY, neotype; P, US, 
iso-neotypes). Figure 20. 

Small trees to 6 m. tall; sparingly pubescent. Branchlets slender, angular to terete, 
glabrescent to sparingly strigose pubescent. Leaves crowded behind tip of branchlets; 
blades obovate to oblong- or elliptic-obovate, rotundate to cuspidate to broadly acumi- 
nate at apex with acumen to 20 cm. long, serrate-dentate along margins with serra- 
tions terminating into setae, 15-20(-30) cm. long, 6-12 cm. wide, chartaceous, some- 
what scabrous above, secondary veins 16-22 pairs, tertiary veins elevated, more 
prominent than lesser venation, scattered strigillose pubescent along minor veins 
(pubescence denser along major veins) above, scattered to sparingly pubescent with 
trichomes of radiate to stellate (along and between veins) mixed with strigose to 
strigillose types (along major veins) beneath; petioles (l-)1.5-4(-6) cm. long, 1.5-3 mm. 
in diameter, half-terete, sparingly to abundantly strigose pubescent. Inflorescences 
somewhat straight, moderately-branched, 35-100-flowered, 7-20 cm. long, 4-8 cm. 
wide, abundantly scurfy pubescent with trichomes of stellate to radiate (especially 



SOEJARTO: SA URA VIA 63 

along lesser ramification) mixed with strigose to shaggy-strigose types, primary 
peduncle 6-12 cm. long, bracts linear, to 6 mm. long. Flowers 15-18 mm. broad, buds to 
5 mm. in diameter, pedicels to 5 mm. long, bracteoles triangular to linear, to 3 mm. 
long; sepals 5, often white, elliptic to oblong or oblong-obovate, acute to rounded, 5-7 
mm. long, 3-4 mm. wide, all densely to abundantly stellate pubescent throughout, but 
occasionally shaggy-strigose trichomes are found on exposed parts in bud, and lower 
portion inside glabrous medially, marginally to submarginally ciliolate; petals 5, 
white, oblong to oblong-elliptic, obtuse to rounded, 7-8(-9) mm. long, 2-5 mm. wide; 
stamens 25-40, filament 2-3 mm. long, anther 2.5 mm. long; ovary 5-loculed, ovoid, 
5-sulcate, glabrous, styles 1-7 mm. long, stigmas simple to capitate. Berries imma- 
ture. 

Habitat. Damp forest, thicket near stream, old clearing, ravines 
and riverbanks, at altitudes of 1,300-2,000 m. 

Distribution . Colombia (Department of Magdalena). 

Specimens examined .COLOMBIA, MAGDALENA: Between 
San Pedro and the headwaters of Rio Sevilla, La Cebolleta, Jan., fr., 
Barclay & Juajibioy 6797 (ECON); Colonia Hurtado, San Lorenzo 
Mts., NW. exposure, June, fl., Kernan 125 (NY, P, US); Santa 
Marta, Sierra del Libano, March, veg., Smith s. n. (NY); Las Nubes, 
Dec., fl., Smith 809 (A, BM, F, G, GH, K, L, MA, MICH, NY, P, S, U, 
UC, US); Rio Domachui Basin, Cancurua, Oct., fr., Cuatrecasas & 
Romero-Castaneda 24804 (US); no. loc., Linden 789 (BM). 

The distribution of S. arnoldi is restricted to the Sierra Nevada of 
Santa Marta, where the type, Arnold Schultze 814, was collected in 
the San Lorenzo area, at an altitude of 2,200 m., on March, 1927. 
Since the original type specimens have possibly been destroyed 
(Sleumer, in litt.), Kernan 125 has been selected as a neotype because 
of the excellent match with the original specific description. In addi- 
tion, Kernan 125 also represents a topotype. 

With regard to field characters, the following note of Harriet 
Barclay (Barclay & Juajibioy 6797) is noteworthy: "Herb to 4-5 m. 
tall; stem red, with brown hairs above. Leaves darker green above, 
red veins, rough. Flowers white in bud, sepals white, petals white, 
filaments yellow-green, anthers yellow, pistil green. Flowers resem- 
ble Begonia. " This is, strangely, quite a contrast to the following note 
by Arnold Schultze: "Peculiar gnarled tree of scandent .habit. In- 
florescences and flower buds pink, petals white inside, light pink 
outside." 

31. Saurauia tomentosa (HBK.) Spreng., Syst. Veg. ed. 16. 4. 
Cur. Post. 2: 211. 1827. Based upon Palava tomentosa HBK. Illus- 
tration: HBK., Nov. Gen. Sp. PI. 7: pi. 650. 1824; Rhodora 65: 14. fig. 
8. 1963. Figure 22. 



64 FIELDIANA: BOTANY 

Palava tomentosa HBK. in Kunth, Synops. PL Aequinoct. 3: 213. 
1922; Nov. Gen. Sp. PL 7: 222. pi. 650. 1824. Type: Bon- 
pland 3206 (P, lectotype; F, isolectotype; NY, US, photos). 

Saurauia ruiziana Steud. var. tomentosa (HBK.) Choisy, Mem. 
Soc. Phys. Geneve 14: 116. 1855. (Based upon P. tomentosa}. 

Saurauia pseudoexcelsa Busc., Malpighia 25: 235. 1912, p. p. (type 
excl.). 

Saurauia sprucei Sprague, Trans. & Proc. Bot. Soc. Edinburgh 22: 
427. 1904. Type: Spruce 6195 (BM, G, K, P). 

Saurauia tomentosa (HBK.) Spreng. var. chillanea Busc., Mal- 
pighia 27: 4. 1914. Type: Sodiro 153 (not seen), ex descr. 

Saurauia pruinosa R. E. Schultes, Bot. Mus. Leafl. 16: 81. 1953. 
Type: Schultes & Villarreal 7651 (ECON, holotype: COL, 
NY, US, isotypes). 

Trees to 30 m. tall, diameter to 40 cm. at base, trunk to 15 m. long, straight, bark 
deeply fissured, corky, brown to ashy -brown, wood light brown, coarsely fibrous, crown 
open; copiously pubescent. Branchlets stout, terete, strongly scarred, ashy to hoary 
tomentose pubescent and mealy in appearance, with trichomes of tufted to dendroid 
types (aloeform, cf. Schultes), brown to whitish brown. Leaves crowded behind tip of 
branchlets; blades obovate to elliptic, acuminate at apex with acumen to 15 mm. long, 
cuneate to rotundate, sometimes pseudoauriculate at base, often oblique, basal flap 
infrequently present, finely denticulate to serrulate-denticulate along margins, (12. 5-) 
20-35(-45) cm. long, (4-)7-12(-15) cm. wide, strongly coriaceous, dark brown to green 
above, gray to ashy-brown or brown beneath, rarely deep red-brown, scabrous above, 
secondary veins (25-)28-37(-42) pairs, tertiary veins elevated, more prominent than 
lesser venation, pubescent with trichomes of tuberculate to tuberculate-strigose (be- 
tween veins) and strongly strigose to shaggy-strigose types (along veins) above, 
densely hoary- to velvety-brown tomentose pubescent with trichomes of dendroid to 
tufted or dendroid-tufted mixed with stellate (between veins) and strongly tufted to 
dendroid-tufted to shaggy-strigose types (along veins) beneath; petioles (1.25-)2-4.5(-6) 
cm. long, 2.5-5.5 mm. in diameter, pubescence similar to that along branchlets. In- 
florescences straight, 25-70(-120)-flowered, 14-26 cm. long, 5-12(-15) cm. wide, pubes- 
cence similar to that along branchlets, primary peduncle (3-)5-13 cm. long, bracts 
subulate, 4-10 mm. long. Flowers (18-)20-25(-30) mm. broad, buds to 10 mm. in 
diameter, pedicels to 15 mm. long, bracteoles triangular, 2-4 mm. long; sepals 5, 9-13 
mm. long, 6-9 mm. wide, outer two ovate to elliptic, acute to obtuse, imbricate one ovate 
to elliptic, obtuse, inner two oblong to suborbicular, obtuse to rounded, all densely 
stellate pubescent throughout, but exposed parts in bud also with tufted trichomes, 
marginally and apically ciliolate to ciliate; petals 5, white to creamy white, oblong to 
suborbicular, obtuse, often deeply incised, 10-13 mm. long, 8-10 mm. wide; stamens 
(87-)100-186(-200), filament 3-4 mm. long, anther 1.5-2 mm. long; ovary 5-loculed, 
globose, 5-sulcate, glabrous, 3-6 mm. in diameter, styles 5, fleshy, obsolete to 6.5(-7.5) 
mm. long, stigmas simple to capitate. Berries green, often with purple or maroon tint, 
globose to ovoid, to 15 mm. across, 5-loculed, 5-sulcate. 

Habitat. Wet mountain forest, submountain forest, cloud-covered 



SOEJARTO: SA URA VIA 65 

zone, secondary forest, edge of primary forest, open and wind-swept 
hillside, steep slope, along shady wet places and streams, sugarcane 
plantation, gardens, abandoned fields, roadside embankment and 
meadows, at altitudes of 2,000-2,800 (-3,000) m. 

Distribution. Venezuela (States of Falcon, Trujillo, Merida, 
Tachira), Colombia (Departments of Norte de Santander, Cun- 
dinamarca, Narino, Putumayo), Ecuador (Provinces of Pichincha, 
Tungurahua, Bolivar, Chimborazo). 

Vernacular names. Babosa (Tachira: Steyermark), Cujaro 
(Merida: Hahn), Moquillo (Narino-Putumayo: Soejarto), Carron 
(Bolivar: Rimbach, Solis), Tation (Bolivar: Solis). 

Specimens examined. VENEZUELA, FALCON: Zumuro coast, 
road between Toronday and Mucuchies, May, fl. fr., Badillo 853 
(VEN). TRUJILLO: Above Jojo, towards La Morita, Aug., fl., Aris- 
teguieta & Medina 3412 (NY, VEN); vie. of Visun (Las Mesitas), Aug., 
R.,Aristeguieta & Medina 3676 (NY, VEN); La Lagunita, Sept., fl., 
John 1146 (US, VEN). MERIDA: 35 km. W. of Merida, along road to 
La Carbonera, Feb., fl., Breteler 3606 (VEN). TACHIRA: Between 
Villapaez and Betania, along Rio Tachira, near Colombo-Venezuelan 
boundary, July, fl. fr., Steyermark 5761 (F, VEN). 

COLOMBIA. NORTE DE SANTANDER: Culaga Valley, near 
Tapata (N. of Toledo), March, fl. fr., Killip & Smith 20347 (A, COL, 
GH, NY, US); Cordillera Oriental, basin of Samaria River, Oct., fr., 
Cuatrecasas & Schultes 12802 (GH). CUNDINAMARCA: Vic. of 
Salto de Tequendama, in the Sabana of Bogota, fl., Humboldt 5761 
(COL, G, NY, photos). NARINO: Municip. of Piedrancha, along Rio 
Guabo, upper reaches of Rio Guiza, July, fl., Soejarto & Pinkley 973 
(COL, ECON, GH), 1437, 1438, 1467, 1468 (all in ECON, GH), fl. fr., 
1469 (ECON, GH, PASTO), 1470 (ECON, GH), fl., 1471 (ECON, GH). 
PASTO: Cantaclaro, near roadside Pasto to la Florida, July, fl., 
Soejarto 1042, fl. fr., 1043 (both in COL, ECON, GH, PASTO); above 
Pasto, road to Buesaco, July, veg., Soejarto & Vogelmann 1201 (COL, 
ECON, GH, PASTO, US); road Pasto to Chachagui, July, veg., 
Soejarto & Hernandez 1207 (COL, ECON, GH, PASTO); above In- 
stitute Tecnologico Agricola Experimental Station, Aug., fl., Soejarto 
1476, veg., 1477 (both in COL, ECON, GH), veg, 1481, 1482 (both in 
ECON, GH); Corota Island on Lake La Cocha, fl., Soejarto 2009 
(ECON, GH); vie. of El Encano, April, fr., Hernandez 10 (ECON, 
PASTO), fl., 80, 81 (both in GH, PASTO, US), fl., Soejarto & Porter 
503, 504 (both in COL, ECON, GH), fl., 508, veg., 508A, 509 (all in 



66 FIELDIANA: BOTANY 

ECON, GH); below Paramo de Tabano, July, ft.,Soejarto 1047 (COL, 
ECON, GH), fl., 1053 (COL, GH, PASTO), veg., 1055 (ECON, GH), fl., 
1432A, 1433 (both in COL, ECON, GH); above Lake La Cocha, on 
road to Sibundoy, near Paramo de Bordoncillo, May, fr., Schultes & 
Villarreal 7564 (COL). PUTUMAYO: Two to 5 km. N. of San Pedro 
de Sibundoy, Aug., fl., Chindoy 180 (ECON); Sibundoy Valley, May, 
fl., Schultes & Villarreal 7651 (ECON, K, NY, US). 

ECUADOR, PICHINCHA: Quito, Bonpland 3206 (F, P; NY, US, 
photos). TUNGURAHUA: NW slope, April, fr.,Bell 736 (EM); Bafios, 
April, fl., Benoist 4178 (P). BOLIVAR: Cerro de Pucarra, trail to 
Telimbela, Nov., fl., Soils 6832 (F); vie. of Chillanes, Nov., veg., Soils 
6668 (F); between Capillaucu and Los Illanes, Oct., fl., Soils 6314 (F); 
Cordillera Occidental, Valle de Limon, Oct., fl., Soils 6398 (F); Bal- 
sapamba and vie., Oct., Rimbach 232 (F, G, NY), fl., 373 (S). CHIM- 
BORAZO: Chimborazo, fl., Spruce 6195 (BM, G, K, P; US, photo). 
GUAYAS: Guayaquil, fl., Ruiz & Pavon s. n. (F, fragment; G). NO 
EXACT LOCALITY: In Andibus Ecuadorensibus, fl., Spruce 5174 
(BM, F-fragment, G, K, NY, P); Palmira, Feb., fl., Benoist 3822 (P); 
San Jose de Minas, March, fl., Benoist 3958 (P). 

S. tomentosa is one of the most beautiful of the South American 
species and is worthy of cultivation. It is commonly found in dis- 
turbed habitats as low, many-branched trees, 3-5 m. tall, but with a 
trunk of up to 30 cm. in diameter, as a result of pruning. The low and 
profusely branched tree with a pink pruinose indument on the 
leaves and large white flowers is a most pleasing sight. 

The original habitat of the species is wet mountain forest with 
rich humus, at altitudes of 2,400-3,000 m., such as that found on the 
Corota Island (2,800 m. alt.) in the middle of Lake La Cocha. The 
vegetation on this island still remains intact because of government 
protection. Here, individuals of S. tomentosa are rather abundant, 
and the trees grow to a height of 15-20 m., with a bole straight and a 
diameter of up to 40 cm. at base. The only other species growing on 
this island is S. bullosa. 

Individuals collected from lower altitudes (800-1,800 m.) are usu- 
ally influenced by other species found in that locality, and can be 
distinguished by the presence of trichome types not found in S. 
tomentosa, the less dense indument, and the smaller size of the 
flowers. This is the case with many of the specimens collected from 
Merida, Norte de Santander, Piedrancha (Narino), and Bolivar. 
However, the presence of dendroid to dendroid-tufted trichome types, 



SOEJARTO: SA URA VIA 67 

combined with the high number of stamens (100-200) identify them 
as members of S. tomentosa. 

32. Saurauia pseudostrigillosa Busc., Malpighia 28: 125. 
1917. Type: Sodiro 152 (COL, GH, NY, photos). Illustration: Mal- 
pighia 28: pi. 6, fig. 11. 1917. 

Saurauia floribunda Benth. ex Sprague, Trans. & Proc. Bot. Soc. 
Edinburgh 22: 426. 1904, p. p. (type incl.) Type: Spruce 
5540 (F, fragment; BM, G, GH, K, NY, P, US) 

Erect shrubs to small trees; sparingly pubescent. Branchlets stout, irregularly 
angular, distinctly scarred, abundantly scurfy-pubescent, trichomes of tuberculate to 
tuberculate-strigose types. Leaves crowded behind tip of branchlets; blades obovate to 
elongate-obovate, rotundate to obtuse but rarely cuspidate at apex, cuneate to rarely 
obtuse at base, sometimes with rudimentary basal flap, serrulate along margins, 
(15-)20-35(-37) cm. long, (6-)8-15(-16) cm. wide, chartaceous, somewhat scabrous 
above, secondary veins (16-)20-35 pairs, tertiary veins elevated, more prominent than 
lesser venation, muricated to tuberculate pubescent along and between veins above, 
scattered to sparingly pubescent with trichomes of radiate to stellate to clustered 
(between veins) and tuberculate-strigose types (along veins) beneath; petioles (1.5-)2- 
3.5(-4.5) cm. long, 2-4 mm. in diameter, slightly furrowed above, glabrescent to spar- 
ingly pubescent along furrow. Inflorescences straight, many-branched, (20-)40-80 
(-lOO)-flowered, (10-)15-30 cm. long, 5-15 cm. wide, densely to abundantly scurfy- 
pubescent with trichomes of strigose to tuberculate-strigose mixed with clustered 
types, primary peduncle 6-14 cm. long, bracts triangular, to 4 mm. long. Flowers ca. 15 
mm. broad, buds to 5 mm. in diameter, pedicels to 5 mm. long, bracteoles triangular, to 
2 mm. long; sepals 5, outer two ovate, 6-8 mm. long, 5-6 mm. wide, acute, imbricate one 
suborbicular, 8-10 mm. long, 6-8 mm. wide, inner two suborbicular-oblong, 8-10 mm. 
long, 7-8 mm. wide, exposed parts in bud densely scurfy-pubescent with trichomes of 
tuberculate type, imbricate parts densely stellate pubescent, all densely stellate 
pubescent inside, marginally and apically ciliolate; petals 5, oblong, obtuse to rounded, 
ca. 10 mm. long (open flowers not seen); stamens 100-200, filament 2.5-3 mm. long, 
anther ca. 1.5 mm. long; ovary 5(-6)-loculed, globose, 5(-6)-sulcate, glabrous, styles 
5(-6), obsolete to 5.5 mm. long, stigmas simple to capitate. Berries 5(-6)-loculed, 
subglobose, to 6 mm. in diameter, 5(-6)-sulcate. 

Habitat. Andean forest, at altitudes of 1,200-2,000 m. 
Distribution. Ecuador (Province of Pichincha). 

Specimens examined. ECUADOR, PICHINCHA: Guajalito, 
Saloya, western descent of the Cordillera, fl., Solis 5676 (F), fr., 5677 
(F); Saloya, "km. 50 to 70 of the road," Aug., fl., Solis 1096 (F). NO 
PRECISE LOCALITY: Western Andes, Las Maquinas, Sept., fr., 
Anthony & Tate 269 (US); in Andibus Ecuadorensibus, Pallatanga, 
fl., Spruce 5540 (F, fragment; BM, G, GH, K, NY, P; US, photo); near 
Canzacoto and Morascocha, fl., Sodiro s. n. (P); no locality, fl., Sodiro 
152 (COL, GH, NY, photos), July, fl., Benoist 4480 (P). 



68 FIELDIANA: BOTANY 

Anthony & Tate 269 and Soils 5677, both of which are in fruiting 
condition, match very well the specific original description and the 
type photograph. On the other hand, Spruce 5540, the type of S. 
floribunda Benth. ex Sprague (a homonym of S. floribunda Lind. & 
PI.; see S. excelsa), has too high a number of flowers per inflores- 
cence (over 100) and too low a number of stamens per flower (34-50), 
but other characters fall within the range of S. pseudostrigillosa. 

33. Saurauia mexiae Killip ex Soejarto, Bot. Mus. Leafl. 22: 268. 
1969. Type: Mexia 8488 (US; holotype; F, K, NY, S, U, UC, 
isotypes). Figure 21. 

Trees to 10 m. tall; copiously pubescent. Branchlets somewhat terete, distinctly 
scarred, abundantly to sparingly strigose pubescent, pubescence mealy. Leaves 
crowded around tip of branchlets; blades somewhat broadly obovate, abruptly acumi- 
nate with acumen to 15 mm. long, obtuse to rounded at base, rarely oblique, serrulate 
along margins, 17-28 cm. long, 9-15 cm. wide, chartaceous, in dry state dark olive- 
brown above, gray-green beneath, somewhat scabrous above, secondary veins 16-22 
pairs, tertiary veins elevated, more prominent than lesser venation, shaggy-strigillose 
pubescent along veins above, abundantly pubescent with trichomes of stellate (along 
and between veins) mixed with strigillose types (along major veins) beneath, epidermis 
somewhat pustulate throughout; petioles 1.5-3 cm. long, 2-3 mm. in diameter, half- 
terete, densely strigillose to strigose pubescent. Inflorescences straight, lax, and 
somewhat spreading, ca. 100-flowered, 18-26 cm. long, 10-17 cm. wide, abundantly to 
densely pubescent with trichomes of shaggy to shaggy-strigillose types, pubescence 
mealy, primary peduncle 8-11 cm. long, bracts triangular to linear, to 6 mm. long. 
Flowers 20-25 mm. broad, buds to 8 mm. in diameter, pedicels 5-12 mm. long, brac- 
teoles minute, subulate, to 2 mm. long; sepals 5, orbicular, rarely oblong, rounded to 
obtuse, 4-6 mm. long, 4-5 mm. wide, exposed parts in bud abundantly pubescent with 
trichomes of stellate to shaggy types, imbricated parts abundantly stellate pubescent, 
pubescence mealy, all abundantly to densely stellate pubescent on upper portion and 
glabrous on lower half inside, marginally and apically ciliolate to ciliate; petals 5, 
white, oblong to oblong-obovate, rounded, 9-11 mm. long, 5-7 mm. wide, stamens 
100-200, filament 2.5-4.5 mm. long, anther 1.5 mm. long; ovary 5-loculed, globose, 
5-sulcate, glabrous, styles 5, 1 mm. long, stigmas simple. Berries unknown. 

Habitat. Riverbank, at an altitude of 95 m. 
Distribution. Ecuador (Province of Esmeraldas). 

Specimens examined. ECUADOR, ESMERALDAS: Parroquiade 
Concepcion, below Playa Rica, Dec., fl., Mexia 8488 (US, holotype; F, 
NY, S, U, UC, isotypes). 

VI. Ser. MACROPHYLLAE Busc., Malpighia 25: 218. 1912. 

Strigosae Busc., 1. c. 218, p. p. min. 
Mesophyllae Busc., 1. c. 218, p. p. 
Veranianae Busc., 1. c. 219, p. p. 
Brachitrichae Busc., 1. c. 220, p. p. min. 



SOEJARTO: SA URA VIA 69 

Gynogynae Busc., 1. c. 221, p. p. min. 
Stenobasicae Busc., 1. c. 221, p. p. min. 

1. Stellate or radiate trichome types predominate over lower leaf surface. 

2. Leaves conic-strigose pubescent above, pitted to lacunose beneath. 

34. S. formosa 
2. Leaves not conic-strigose pubescent above, neither pitted nor lacunose beneath. 

3. Inner surface of sepals glabrous to partly stellate pubescent (usually only on 
upper portion). 

4. Villous-barbate trichomes present at axils of secondary veins beneath, 
stamens more than 100 35. S. chaparensis 

4. Villous-barbate trichomes absent at axils of secondary veins beneath, 
stamens less than 60. 

5. Inflorescences usually pendulous, 5-30-flowered, bracts usually 
foliaceous, flowers 15-25 mm. broad, stamens 40-50. 

36. S. peduncularis 
5. Inflorescences straight, 40-200-flowered, bracts minute and not 

foliaceous, flowers 10-13 mm. broad, stamens 13-30. 

37. S. putumayonis 

3. Inner surface of sepals densely stellate pubescent. 

6. Leaves sparingly to abundantly pubescent with trichomes of setose to 
setulose mixed with stellate types beneath. 

7. Inflorescences 6-15(25)-flowered, flowers 15 mm. broad, stamens 
35-40, leaf blade smooth above 38. S. herthae 

1 . Inflorescences 30-100-flowered, flowers 12.5 mm. broad, stamens 
25-45, leaf blade puncticulate and scabrous 39. S. lehmannii 

6. Leaves glabrescent, scabrous above, trichomes of strigose to strigillose 
mixed with radiate types beneath, inflorescences (5-)30-150-flowered, 

flowers 10-18 mm. broad, stamens (30-)50-85(-100) 40. S. scabra 

1. Unbranched and/or branched trichome (but not of stellate or similar) types pre- 
dominate over lower leaf surface. 

8. Leaf pubescence distinctly heterotrichous beneath with trichomes of 
strigose, setose or clustered, sometimes mixed with stellate or radiate types. 

9. Texture of leaves membranaceous, trichomes of strigillose mixed with 
slender-stellate types beneath, inflorescences slender and flexuous, 20- 
50-flowered, flowers 15 mm. broad, stamens 30-40 . . . .41. S. tambensis 

9. Texture of leaves chartaceous to coriaceous, trichomes of setose or hir- 
sute or clustered, often mixed with radiate types beneath, inflorescences 
robust, straight, more than 50-flowered, flowers 10-25 mm. broad, sta- 
mens over 50. 

10. Inflorescences (15-)30-70(-150)-flowered, flowers 10-18 mm. broad, 
stamens (30-)50-85(-100) 42. S. excelsa 

10. Inflorescences (20-)50-200-flowered, flowers 15-25 mm. broad, sta- 
mens (65-)100-200(-240) 43. S. brachybotrys 

8. Leaf pubescence distinctly homotrichous beneath with trichomes of un- 



70 FIELDIANA: BOTANY 

branched types, branched trichomes mostly absent or nearly so, then only 
restricted along midrib. 

11. Inflorescence and lower leaf pubescence hirsute, trichomes of up to 5 
mm. long, branched trichomes absent. 

12. Apex of leaves caudate (rarely acuminate), acumen to 45 mm. long, 
inflorescences compact, flower pedicels 6 mm. long. 

44. S. isoxanthotricha 

12. Apex of leaves acuminate, acumen to 10 mm. long, inflorescences 
lax, flower pedicels 10-15 mm. long 45. S. meridensis 

11. Inflorescence and lower leaf pubescence setose to strigose, stellate 
trichomes sometimes present along major veins beneath, inflorescences 
compact, flower pedicels 1-4 mm. long 46. S. prainiana. 

34. Saurauia formosa Sleumer, Notizbl. 12: 144. 1934. Type: 
Weberbauer 6637 (F, lectotype: GH, US, isolectotypes). Figure 23. 

Shrubs; copiously pubescent. Branchlets angular, stout, abundantly to densely 
pubescent with trichomes of sericeous-strigose type, yellowish brown, to 5 mm. long. 
Leaves clustered behind tip of branchlets; blades elliptic to obovate or oblong-obovate, 
acuminate at apex with acumen to 15 mm. long, cuneate at base, rarely oblique, 
setaceous to ciliate-serrulate along margins, (23-)30-45 cm. long, (10-)13-19 cm. wide, 
chartaceous to subcoriaceous, in dry state dark yellowish brown above, light yellow- 
brown to grayish yellow-brown beneath, strongly scabrous above, somewhat soft 
beneath, secondary veins 24-28 pairs, tertiary veins elevated, more prominent than 
lesser venation (reticulation prominent beneath), sparingly to abundantly pubescent 
with trichomes of conic-tuberculate to conic-strigose (between veins) and strigose to 
sericeous-strigose types (along major veins) above, pitted to lacunose between minor 
veins and abundantly pubescent with trichomes of stellate (along and between veins) 
mixed with setose to sericeous types (along major veins) beneath; petioles robust, 
(3-)6-ll cm. long, 5-7 mm. in diameter, half-terete, densely sericeous-hirsute pubes- 
cent. Inflorescences straight, robust, 50-100-flowered, 20-33 cm. long, 5-15 cm. wide, 
densely to abundantly pubescent, trichomes of sericeous mixed with thin, minute and 
white stellate types, peduncle 10-16 cm. long, bracts triangular, to 10 mm. long. 
Flowers 15-20 mm. broad (22 mm., cf. Sleumer), buds to 7 mm. in diameter, pedicels to 5 
mm. long, bracteoles subulate to triangular, to 7 mm. long; sepals 5, oblong to oblong- 
obovate, obtuse to rounded, 8-10 mm. long, 4-6 mm. wide, exposed parts in bud densely 
strigillose pubescent, imbricated parts densely stellate pubescent, all abundantly to 
densely stellate pubescent on upper portion and glabrous on lower half inside, 
marginally ciliolate; petals 5, white, oblong-spatulate, rounded, 10-12 mm. long, 
3-4.5 mm. wide; stamens 60-80, filament 3-4 mm. long, anther 1.5-2 mm. long; ovary 
5-6-loculed, ovoid to globose, 5-6-sulcate, glabrous, styles 5-6, to 4 mm. long, stigmas 
capitate. Berries not known. 

Habitat. Mountain forest (?), at altitudes of 2,800-2,900 m. 
Distribution. Peru (Departments of Junin and Huancayo). 

Specimens examined. PERU, JUNIN: Jauja, vie. of Rio Masam- 
eri, tributary of Rio Pangoa, April., fl., Weberbauer 6637 (F, lee- 



SOEJARTO: SA URA VIA 71 

totype; GH, US, isotypes). HUANG AYO: Oxapampa, Aug., fl., 
Soukup 2433 (F, US). 

35. Saurauia chaparensis Soejarto, Bot. Mus. Leafl. Harvard 
Univ. 22: 270. 1969. Type: Steinbach 8920 (GH, holotype; F, K, NY, 
S, isotypes). Figure 24. 

Shrubs; copiously pubescent. Branchlets moderately stout, terete, prominently 
scarred, abundantly pubescent with trichomes of shaggy to strigose types, but some- 
times glabrescent. Leaves crowded behind tip of branchlets; blades obovate to 
oblong-obovate, blunt to very shortly acuminate at apex, cuneate at base, rarely 
oblique, serrulate with very fine serrulations along margins, 10-18 cm. long, 3-6 cm. 
wide, chartaceous, in dry state sooty above, olive-brown beneath, distinctly scabrous 
throughout, secondary veins elevated, more prominent than lesser venation, spar- 
ingly pubescent with trichomes of clustered and strigose types (along and between 
veins) and with pustulate epidermis above, abundantly pubescent with trichomes of 
stellate to tufted (along and between veins) and shaggy to strigillose types (along 
major veins) beneath, villous-barbate pubescent at axils of secondary veins beneath; 
petioles 2-3 cm. long, 1.5-2 mm. in diameter, terete to half-terete, scabrous and 
abundantly pubescent with trichomes of shaggy-strigillose type. Inflorescences 
straight, somewhat loose, 30-80-flowered, 12-20 cm. long, 5-12 cm. wide, densely 
mealy-pubescent with tuberculate trichomes, primary peduncle 5-10 cm. long, bracts 
linear, to 6 mm. long, very rarely foliaceous, to 25 mm. long. Flowers ca. 15 mm. 
broad, buds to 6 mm. in diameter, pedicels 5-15(-20) mm. long, bracteoles triangular, 
to 2 mm. long; sepals 5, outer two oblong-obovate, acute, 3.5-4.5 mm. long, 3-3.5 mm. 
wide, imbricate one suborbicular, rounded, 4-5 mm. long, 3.5-4 mm. wide, inner two 
suborbicular to orbicular-oblong, 4-5 mm. long, 3.5-4.5 mm. wide, exposed parts in 
bud densely shaggy pubescent, imbricated parts glabrous, all glabrous inside, margi- 
nally subentire, apically irregularly ciliolate; petals 5, oblong to oblong-obovate, 
rounded to truncate and often incised, 7-9 mm. long, 5-7.5 mm. wide; stamens 100- 
150, filament 2-3 mm. long, anther 1-2 mm. long; ovary 5-7-loculed, globose, 5-7- 
sulcate, glabrous, styles 5-7, obsolete to 4 mm. long, stigmas simple to capitate. 
Berries 5-7-loculed, globose, to 5 mm. across (immature), 5-7-sulcate. 

Habitat. Forest, at altitudes of 2,200-2,400 m. 
Distribution. Bolivia (Department of Cochabamba). 

Specimens examined. BOLIVIA, COCHABAMBA: Chapare, In- 
cachaca, Jan., fl. fr., Steinbach 8920 (GH, holotype; F, NY, S, 
isotypes), March, fl., 9513 (BM, F, G, GH, K, NY, S). 

The following features characterize S. chaparensis: (1) leaves 
small, abundantly stellate pubescent with villous-barbate trichomes 
at axils of secondary veins beneath, (2) inflorescences loose, (3) flow- 
ers of medium size (ca. 15 mm. broad) and long-pedicellate (to 20 mm. 
long), (4) stamens high in number (more than 100). 

A rather remarkable feature of S. chaparensis is the frequency of 
irregular repetition of some floral parts. Flowers with 6-7-loculed and 



72 FIELDIANA: BOTANY 

6-7-styled ovaries are found in almost equal proportions as those with 
5-loculed and 5-styled ovaries, in the same inflorescence. 

36. Saurauia peduncularis Tr. & PI., Ann. Sci. Nat. Ser. 4, 18: 
267. 1862; Prodr. Fl. Nov. Granat. 1: 264. 1862. Type: Triana 3252 
(A, COL, GH, NY, US, all photos). Figure 25. 

Shrubs to 3 m. tall, erect to curving, sometimes trailing, rarely bushy; sparingly 
pubescent. Branchlets slender, terete, glabrescent to abundantly pubescent, trichomes 
of setose to hirsute types, yellowish to bright brown, to 4 mm. long. Leaves clustered 
behind tip of branchlets; blades narrowly to broadly elliptic to obovate, cuspidate to 
acuminate at apex with acumen to 15(-20) mm. long, cuneate to obtuse at base, rarely 
oblique, serrate to biserrate along margins (7-)10-25 cm. long, (3.5-)5-10(-15) cm. wide, 
chartaceous to subcoriaceous, light to dark green on both sides, scarcely scabrous 
above, secondary veins (9-)13-20(-23) pairs, tertiary veins elevated, more prominent 
than lesser venation, scattered to abundantly pubescent with trichomes of strigillose 
(between veins) to strigose types (along veins) above, sparingly pubescent with 
trichomes of stellate (along and between veins) mixed with strigose types (along 
midrib) beneath; petioles 1.5-4 cm. long, 1.5-3 mm. in diameter, sparingly setose 
pubescent. Inflorescences usually pendulous, rarely ascending, lateral cymes squar- 
rose, loose, (5-)10-25(-30)-flowered, 7-30 cm. long, 4-13 cm. wide, abundantly pubes- 
cent with trichomes of setose to hirsute mixed with radiate to stellate or clustered 
types, primary peduncle 4.5-18 cm. long, bracts foliaceous, elliptic to elongate- 
elliptic, acuminate to acuminate-cirrhous at apex, cuneate to attenuate at base, 
setaceous-serrulate along margins, to 45 mm. long, to 15 mm. wide, sparingly strigil- 
lose pubescent throughout. Flowers 15-25 mm. broad, buds to 7 mm. in diameter, 
pedicels 4-10 mm. long, bracteoles linear-subulate to foliaceous, 4-15 mm. long; sep- 
als 5, pale green, outer two ovate to elliptic, subacute, 5-7 mm. long, 3-4.5 mm. wide, 
imbricate one and inner two suborbicular to broadly elliptic, obtuse, 5-8 mm. long, 4-6 
mm. wide, exposed parts in bud abundantly to sparingly pubescent with trichomes of 
stellate and strigose types, imbricated parts abundantly stellate pubescent to gla- 
brous, all glabrous to partly stellate pubescent (on upper portion) inside, marginally 
ciliolate; petals 5, white, spatulate to orbicular-obovate, rounded to retuse, 8-13 mm. 
long, 6-9 mm. wide; stamens 45-50, filament 2-3 mm. long, anther 2-3 mm. long; 
ovary 5-loculed, globose to ellipsoid, 5-sulcate, glabrous, styles 5, obsolete to 4(-5) 
mm. long, stigmas simple to subcapitate. Berries immature. 

Habitat. Mountain forest, secondary forest, along creek and 
shady, wet riverbank, on rocky soil, at altitudes of 1,000-1,600 
(-2,400) m. 

Distribution. Colombia (Department of Narino). 

Vernacular names. Moquillo (Ricaurte: Soejarto; Tuquerres: 
Triana & Planchon). 

Specimens examined. COLOMBIA, NARINO: Cartagena, along 
Guiza River, July, fl., Soejarto 935, 937, 938, 943 (all in ECON, GH, 
PASTO), fl. 936 (COL, ECON, GH, PASTO), veg., 1456 (ECON, GH); 
W. of Ricaurte, El Palmar, Aug., fl., Soejarto 1446 (ECON, GH); 



SOEJARTO: SA URA VIA 73 

Cordillera of Tuquerres, fl., Triana 3252 (A, COL, GH, NY, US, all 
photos). 

NO PRECISE LOCALITY: Rio Cuaiquar, May, fl., Andre K1000 
(K); El Tena, Sept., fl., Benoist 4773 (P). 

The type specimen of S. peduncularis (Triana 3252) was collected 
from "forest de la cordillere de Tuquerres, alt. 2400 metres," which is 
rather high for the species. All the recent collections were made from 
the Ricaurte area at measured altitudes of 1,000-1,600 m. Although 
only the photograph of the type was available for study, the identity 
of the species is unmistakable from the following characteristics: (1) 
loose and pendulous inflorescences with lateral cymes normally per- 
pendicular to the main axis, (2) foliaceous bracts, and (3) relatively 
long primary peduncle (up to three times the length of the flower- 
bearing part), hence the specific name. The above amplified descrip- 
tion has been based upon the recent collections. 

37. Saurauia putumayonis R. E. Schultes, Caldasia 2: 42. 
1944. Type: Cuatrecasas 11431 (COL, holotype; F, US, isotypes). 
Illustration: Caldasia 2:42. pi. 1. 1943. 

Shrubs to small trees to 5 m., rarely bushy, branches slender, often curving; spar- 
ingly pubescent. Branchlets slender, terete, glabrescent to scattered pubescent with 
trichomes of strigillose to strigose types, white to light brown. Leaves clustered behind 
tip of branchlets; blades obovate to oblong-obovate, cuspidate to shortly acuminate at 
apex with acumen to 15 mm. long, cuneate to rarely oblique at base, serrulate along 
margins, (10-)13-25(-30) cm. long, (5-)7-15 cm. wide, chartaceous to membranaceous, 
somewhat scabrous above, secondary veins 16-22 pairs, tertiary veins elevated, more 
prominent than lesser venation, scattered to sparingly pubescent with trichomes of 
tuberculate to setulose (along and between minor veins) and strigillose to setose types 
(along major veins) above, scattered to sparingly pubescent with trichomes of stellate 
to radiate or clustered (along and between veins) mixed with strigillose to strigose or 
setose types (along major veins) beneath; petioles (l-)2-5 cm. long, 2-3 mm. in diameter, 
furrowed above, glabrescent to sparingly strigillose pubescent. Inflorescences straight, 
(20-HO-100(-200)-flowered, 15-20(-42) cm. long, 8-20 cm. wide, sparingly to abun- 
dantly scurfy -pubescent with trichomes of strigillose to setulose types, primary pedun- 
cle 5-15 cm. long, bracts triangular to linear, to 10 mm. long, rarely foliaceous, to 25 
mm. long. Flowers 10-13 mm. broad, buds to 4 mm. in diameter, pedicels 3-10 mm. long, 
bracteoles triangular to linear, to 3 mm. long, sepals 5, pale green to rarely white or 
reddish maroon, ovate to elliptic or suborbicular, acute to obtuse or rounded, 4-5.5 mm. 
long, 2.5-4 mm. wide, exposed parts in bud sparingly scurfy-pubescent with trichomes 
of setulose to strigillose to shaggy mixed with radiate types, imbricated parts glabrous, 
all glabrous to scattered radiate pubescent on upper portion inside, marginally entire 
to ciliolate; petals 5, white to rarely pink, oblong to oblong-obovate, rounded to rarely 
incised, 6.5-7.5 mm. long, 4-5.5 mm. wide; stamens (13-)18-30(-33), filament 1.5-2 mm. 
long, anther 2-2.5 mm. long; ovary 5-loculed, globose, 5-sulcate, styles 5, obsolete to 5.5 
mm. long, stigmas simple to capitate. Berries 5-loculed, globose, to 10 mm. across, 
5-sulcate. 



74 FIELDIANA: BOTANY 

Habitat. Subparamo, wet mountain forest, wet rain forest, bor- 
der of rather densely forested mountain, secondary forest, steep and 
rocky slopes, road cut and riverbank, at altitudes of 1,400-2,900 
(-3,000) m. 

Distribution. Colombia (Departments of Meta, Huila, and 
Putumayo). 

Vernacular names. Moquillo (Putumayo: Schultes & Villarreal). 

Specimens examined. COLOMBIA, META: Rio del Nevado, 
below the confluence of Rio Arroz, south slope of Paramo of Sumapaz, 
Sept., fr., Fosberg 20941 (NY, US). HUILA: Rio Villalobos, region of 
the confluence of Rio Cauchos, Jan., fl., Schultes & Villarreal 5199 
(COL, GH, US). PUTUMAYO: Road San Francisco of Sibundoy to 
Mocoa, Feb., fl., Bristol 539 (COL, ECON, GH); east-facing slope of 
the Cordillera, between Mocoa and Sachamates, Dec., fl., Cuat- 
recasas 11416 (COL, US), 1 1431 (COL, F, US); Mocoa River drainage, 
between Sachamates and San Antonio, Jan., fl., Ewan 16692 (BM, 
NY, US); San Francisco of Sibundoy, Feb., fl., Miguel 63 (F); Cerro 
Portachuelo, road Sibundoy to Pepino of Mocoa, Aug., fl., 
Fernandez-Perez 5837 (COL, ECON, GH), March, fl., Schultes & 
Cabrera 19028 (ECON, F, GH, NY, US), July & Aug., fl., fr.,Soejarto 
520, 1552, 1553, 1554, 1558, 1559, 1561, 1573, 1593 (all in ECON, 
GH), fl.fr., 1135, 1141, 1143,1145,1164, and veg., 1144 (all in ECON, 
GH, PASTO), fl. fr., 1048, 1121, 1140 (all in COL, ECON, GH, 
PASTO), veg., 1117,1547 (both in GH); above Santiago of Sibundoy, 
La Chorrera, July, fl. fr., Soejarto 1525, 1526, 1529 (all in ECON, 
GH). 

38. Saurauia herthae Sleumer, Feddes Repert. 45: 9. 1938. 
Type: Schultze-Rhonhof 2165 (K, lectotype). 

Shrubs to 6 m. tall; copiously pubescent. Branchlets often crooked, slender, terete, 
abundantly strigose to setose pubescent, trichomes yellowish to rusty brown. Leaves 
clustered behind tip of branchlets; blades obovate to elliptic-obovate, acuminate at 
apex with acumen to 25 mm. long, cuneate to obtuse at base, serrulate along margins, 
serrulations widely separated around base, (12-)20-30 cm. long, (4.5-)7-12 cm. wide, 
chartaceous, in dry state sooty-green above, deep grayish olive-brown beneath, some- 
what smooth above, secondary veins 14-22 pairs, tertiary veins elevated, more promi- 
nent than lesser venation, scattered to sparingly pubescent with trichomes of setulose 
type (especially along major veins) above, sparingly to abundantly pubescent with 
trichomes of stellate and dendroid (along and between veins) mixed with strigose to 
setose types (along major veins) beneath; petioles 1-2.5 cm. long, 1.5-2 mm. in diame- 
ter, sparingly strigose pubescent. Inflorescences somewhat straight to ascending, 
small, little-branched, 6-15-flowered, 6-8 cm. long, 1.5-3.5 cm. wide, densely pubescent 
with strigose to shaggy trichomes, primary peduncle 3-4 cm. long, bracts linear, to 5 



SOEJARTO: SA URA VIA 75 

mm. long. Flowers ca. 15 mm. broad, buds to 4 mm. in diameter, pedicels to 4 mm. in 
fruiting condition, bracteoles linear, to 4 mm. long; sepals 5, oblong-elliptic to subor- 
bicular, acute to obtuse, 6-7.5 mm. long, 4.5 mm. wide, exposed parts in bud abundantly 
pubescent with trichomes of strigose and stellate types, imbricated parts densely 
stellate pubescent, all densely stellate pubescent inside, marginally ciliolate; petals 5, 
white, oblong-obovate, rounded, 7-8 mm. long, 4.5 mm. wide; stamens 35-40, filament 
2-2.5 mm. long, anther 2.5-3 mm. long; ovary 5-loculed, globose, 5-sulcate, glabrous, 
styles 5, to 5 mm. long, stigmas capitate. Berries 5-loculed, subglobose, to 7 mm. across, 
5-sulcate. 

Habitat. Rain forest, riverbank, at altitudes of 350-2,600 m. 

Distribution. Ecuador (Provinces of Napo-Pastaza, Tungurahua, 
and Santiago-Zamora). 

Vernacular names. Shauri (Santiago-Zamora: Cazalet & Pen- 
nington). 

Specimens examined. ECUADOR, NAPO-PASTAZA: Canelos 
(Oriente), Jan., fl., Schultze-Rhonhof 2165 (K, lectotype). TUN- 
GURAHUA: valley of Pastaza River, between Banos and Cashurco, 
Sept., fl., Hitchcock 21883 (NY, US). SANTIAGO-ZAMORA: Taisha, 
bank of Guaguayme River, Jan., fr., Cazalet & Pennington 7543 (NY, 
UC, US); canyon of Paute River, near confluence of Yubal River, 
Hacienda Frutillas, fl., Fosberg & Prieto 22740 (US); eastern region 
between Rio Sordo and La Esperanza, trail to Huamboya, descent of 
Cordillera Oriental, Feb., fl., Solis 7375 (F). 

S. herthae is allied to S. prainiana, but the former can be distin- 
guished in the predominantly stellate trichomes on the lower leaf 
surface and in the small, few-flowered inflorescences, in contrast to 
the predominantly setose trichomes on the lower leaf surface and the 
larger, many -branched inflorescences, in the latter. 

39. Saurauia lehmannii Hieron., Bot. Jahrb. Syst. 20, Beibl. 49: 
47. 1895. Type: Lehmann 6673 (K, lectotype; F, US isolectotypes; 
GH, NY, photos). Figure 26. 

Small trees to 8 m. tall, ramification squarrose and open, branches thick; copiously 
pubescent. Branchlets terete, abundantly pubescent with setose to hirsute trichomes. 
Leaves clustered behind tip of branchlets; blades elongate-obovate to obovate, cuspi- 
date at apex with acumen to 10 mm. long, cuneate at base, setaceous-serrate to 
-biserrate along margins, 15-25(-30) cm. long, 6-14 cm. wide, subcoriaceous, brownish 
green above, grayish green beneath (in dry state dark to dull reddish brown above, 
ashy brown beneath), puncticulate and scabrous above, secondary veins 15-20 pairs, 
tertiary veins elevated, more prominent than lesser venation, sparingly and minutely 
muricated (along and between minor veins) and abundantly pubescent with setose 
trichomes (along major veins, especially midrib) above, sparingly to abundantly 



76 FIELDIANA: BOTANY 

pubescent with trichomes of stellate (along and between veins) mixed with setose types 
(along midrib) beneath; petioles 2-4 cm. long, 1.5-3 mm. in diameter, sparingly to 
abundantly pubescent with trichomes of setose to hirsute types. Inflorescences 
straight, lateral cymes often squarrose, 30-100-flowered, ll-20(-24) cm. long, (4-)6-12 
cm. wide, abundantly to densely pubescent with trichomes of setose to hirsute mixed 
with stellate types, primary peduncle 6-12 cm. long, bracts subulate, to 7 mm. long. 
Flowers ca. 12.5 mm. broad, buds to 3 mm. in diameter, pedicels 2-5 mm. long, 
bracteoles 1.5-3 mm. long; sepals 5, green to reddish, elliptic to oblong, 5-6.5 mm. long, 
3-4.5 mm. wide, exposed parts in bud densely scurfy-pubescent with trichomes of setose 
mixed with stellate types, imbricated parts densely stellate pubescent, all densely 
stellate pubescent inside, marginally to submarginally ciliate; petals 5, creamy white, 
oblong, rounded to incised, 6-7.5 mm. long, 4-5 mm. wide; stamens 25-45, filament 
1.5-2 mm. long, anther 2-3 mm. long; ovary 5-loculed, ovoid to subglobose, 5-sulculate, 
glabrous, styles 5, 1-4 mm. long, stigmas simple to capitate. Berries not known. 

Habitat. Wet tropical forest, rich dense jungle, forested slopes, at 
altitudes of 1,000-2,300 m. 

Distribution. Ecuador (Provinces of Pichincha, Azuay, El Oro). 

Specimens examined. ECUADOR, PICHINCHA: Santo Domingo 
de Colorado, 0.5 km. N. of town, April, ft., Little, Jr. 6169 (F, K, US); 
Cordillera Occidental, Saloya, Aug., fl., Solis 10964 (F). AZUAY: W. 
Andes of Cuenca, fl., Lehmann 6673 (F, K, US; GH, NY, photos); 
between Chacanceo and Rio Blanco, on road to Molletura (between 
Putucay and Norcay rivers), June, fl., Steyermark 52814 (NY). EL 
ORO: Palma River (tributary of Amarillo River), near Pampa de los 
Cedros, S. of Cerro Chivo-Turco, Aug., fl., Steyermark 53784 (NY). 

40. Saurauia scabra (HBK.) Dietr., Synops. PI. 3: 272. 1843 (non 
Poepp. ex Choisy, 1855). Based upon Palava scabra HBK. Illustra- 
tion: HBK., Nov. Gen. Sp. PI. 7: pi. 648, 649. 1824, Caldasia 2:43, fig. 
7-11. 1943. Figure 27. 

Palava scabra HBK. in Kunth Synops. PI. Aequinoct. 3: 213. 
1822; Nov. Gen. Sp. PI. 7: 221. 1824. Type: Bonpland s. n. 
(P, lectotype; F, fragment, isolectotype). 

Saurauja scabra (HBK.) Tr. & PL, Ann. Sci. Nat. ser. 4(18): 266. 
1862; Prodr. Fl. Nov. Granat. 1: 263. 1862 (non Poepp. ex 
Choisy, 1855). Based upon P. scabra HBK. 

Saurauia humboldtiana Busc., Malpighia 27: 305. pi. 11, fig. 23. 
1916, p. p.; 1. c. 28: pi. 1, fig. 23. 1917. Type: Bonpland s. n. 
(F, fragment; P). Same type specimen as that of Palava scabra 
HBK. 

Saurauia humboldtiana var. bonplandiana Busc., Malpighia 27: 
311. 1916, p. p. (type incl.). Type: Bonpland s. n. (F, frag- 
ment; P). Same type as that of Palava scabra HBK. 



SOEJ ARTO: SA URA VIA 77 

Saurauia prainiana Busc. var. humboldtiana Busc.,Malpighia25: 
396. 1913. Type: Triana s. n. (n. v.), ex descr. 

Saurauia narcissifragrans R. E. Schultes, Caldasia 2: 39, 43 (figs. 
7-11). 1943. Type: Schultes & Villarreal 5169 (ECON, 
holotype; COL, F, L, NY, S, US, VEN, isotypes). 

Trees to 7(-15) m. tall, diameter to 20 cm. at base, trunk crooked to straight, bark 
slightly fissured, corky, brown, crown open; sparingly pubescent. Branchlets terete, 
glabrescent to scattered strigose pubescent, trichomes gray to rusty brown. Leaves 
crowded behind tip of branchlets; blades elliptic-oblong to narrowly obovate, acute to 
very shortly and abruptly acuminate at apex, cuneate to narrowly cuneate at base, 
very rarely oblique, serrulate to setaceous-serrulate along margins, (10-)15-24(-31) 
cm. long, (3.5-)5-8(-l 1) cm. wide, chartaceous to subcoriaceous, green to pale green and 
scabrous throughout, secondary veins (9-)15-22(-27) pairs, tertiary veins elevated, 
more prominent than lesser venation, scattered pubescent with trichomes of strigillose 
to rarely strigose types along and between veins above, scattered to sparingly pubes- 
cent with trichomes of strigillose to strigose (along major veins) mixed with radiate 
types (between minor veins) beneath; petioles (l-)1.5-3.5(-4.5) cm. long, 1.5-3.5 mm. in 
diameter, glabrescent. Inflorescences crowded behind tip of branchlets, erect to ascend- 
ing, (5-)15-40(-60)-flowered, (5-)10-20(-29) cm. long, (l-)3-10(-14) cm. wide, scattered 
pubescent with trichomes of strigose to sericeous-strigose types, primary peduncle 
(l-)3.5-8(-ll) cm. long, bracts subulate to triangular, 3-10 mm. long. Flowers 10-18 
mm. broad, buds to 8 mm. in diameter, pedicels to 3 mm. long, bracteoles triangular, 
2-4 mm. long; sepals 5, elliptic to obovate, obtuse to acute, 5-8 mm. long, 3-6 mm. wide, 
exposed parts in bud densely shaggy-strigose pubescent, imbricated parts densely 
stellate pubescent, all abundantly to densely stellate pubescent inside, marginally to 
submarginally ciliate; petals 5, white, elliptic to oblong, obtuse, 7-15 mm. long, 6-10 
mm. wide; stamens (20-)28-45(-70), filament 2-4 mm. long, anther 1.5-3.5 mm. long; 
ovary 5-loculed, globose to ovoid, 5-sulculate, glabrous, styles 5, obsolete to 6(-7) mm. 
long, stigmas simple to subcapitate. Berries green, persistent, styles purple-red, 5- 
loculed, globose, to 10 mm. across, 5-sulcate. 

Habitat. Dense forest, cloud forest, submountain forest, secon- 
dary forest, open hillside, thicket, meadows, along streams and 
rivulet, shady and humid places, roadside and gardens, at altitudes of 
1,400-2,800 m. 

Distribution. Venezuela (State of Merida), Colombia (Depart- 
ments of Santander, Cundinamarca, Caldas, Tolima, Boyaca, Huila, 
Caqueta). 

Vernacular names. Chupa-chupa (Merida: Bernardi), Moquillo 
(Cundinamarca: Soejarto). 

Specimens examined. VENEZUELA, MERIDA: El Carrizal, Los 
Granates, ridge of Say-say, Feb., fl., Bernardi 2065 (NY). 

COLOMBIA, SANTANDER: Salto de Tequendama (Tequen- 
damita?), July, n., Barkley & Torres-Romero 35338 (US); Surata 



78 FIELDIANA: BOTANY 

River Valley, above Surata, Jan., fl., Killip & Smith 16596 (A, F, 
GH, NY, US); vie. of California, Jan., fl., Killip & Smith 16933 (A, 
GH, NY, US); vie. of Charta, Feb., fl., Killip & Smith 19024, 19103 
(both in A, GH, NY, US); vie. of Tona, Feb., fl. fr., Killip & Smith 
19475, 19480 (both in A, GH, NY, US). CALDAS: Cordillera Cen- 
tral, Salento, July, fl., Pennell 8878 (NY). BOYACA: Sierra Nevada 
del Cocuy, near Boavita, Aug., fl., Grubb et al. 686 (US). CUN- 
DINAMARCA: Sabana de Bogota, fl., Ariste-Joseph A-223, B-125 
(both in US); Cordillera Oriental, east facing slope, between El Salto 
and El Colegio, March, fl., Cuatrecasas 8196 (COL, F, US); El Tab- 
lazo between Subachoque and San Francisco, Finca "El Carmen," 
Jan., fl., Garcia-Barriga 11012 (US); Alban, roadside, 1 km. before 
village, Aug., fl., Garcia-Barriga 12544 (US); road to Fusagasuga, 
between Aguaclara and La Aguadita, Oct., fl., Garcia-Barriga 
16121 (COL, ECON), May, fl., Schneider 1056 (S); Nemocon, Cerro 
del Mortino, Dec., fl., Garcia-Barriga 17717 (COL, ECON); road 
Mosquera to La Mesa, Dec., fl., Garcia-Barriga 17738 (COL, ECON), 
Oct., fl., Uribe-Uribe 4124 (A, COL, ECON, GH, K); San Isidro, 8 
km. S. of Gachala, May, fl., Grant & Fosberg 9323 (NY, US); 
Facatativa highway, Sept., fl., Haught 6140 (US); June, fl., Uribe- 
Uribe 4816 (COL, ECON, GH); Tocaima, Dec., fl., Perez-Arbelaez 
2438 (US); Jute, on road Bogota to La Mesa, Aug., fl. fr., Soejarto 
283, 318, 319 (all in COL, ECON, GH); Salto de Tequendama, vie. of 
waterfall, fl. fr., Soejarto & Idrobo 912, 913 (both in COL, ECON, 
GH), Soejarto 2058 (ECON, GH); March, fl., Killip 33977 (A, COL, 
F, BM, NY, S, US); Bojaca, road to Lake Pedro Palo, June, fl., 
Uribe-Uribe 1764 (COL); Pacho, Oct., fl., Uribe-Uribe 1828 (COL, 
ECON); Dec., fl., Nujos. n. (K); Zipacon, vie. of Boca de Monte, Feb., 
fl., Uribe-Uribe & Villarreal 2555 (COL, ECON); Santandercito, 
below Salto de Tequendama, June, fl., Uribe-Uribe 5239 (COL, 
ECON). TOLIMA: La Palmilla (Quindio), Prov. de Mariquita, fl., 
Triana 3253A (G: the left element on the sheet); same locality, fl., 
Trianas. n. (NY, P) (note on sheet: Saurauia scabra HBK.). HUILA: 
Mountains, 10 km. SE of Santa Anna, Feb., ft., Little, Jr. 7272 (NY, 
US); Rio Villalobos, vie. of Rio Suazita, Quebrada Guayabo, Jan., fl., 
Schultes & Villarreal 5169 (COL, ECON, F, NY, S, U, US, YEN), 
53/7 (COL). NO PRECISE LOCALITY: Santa Anna, fl., Bonpland 
s. n. (F, fragment; P) (annotated as: Palava scabra HBK. [N.G. & 
Sp]); no loc., fl., Mutis 1182, 4598, 4599, 5773 (all in US); fl., no 
collectors, n. (K); no loc., f[.,Purdies. n. (K). 



SOEJARTO: SA URA VIA 79 

When Humboldt et al. (1824) published the amplified description 
ofPalava scabra (Nov. Gen. Sp. PI. 7: 221. pis. 648 and 649. 1824), 
two descriptions were provided. The first was based upon Bonpland's 
collection from Santa Anna (?Huila, plate 648), Colombia; the sec- 
ond was communicated by Mutis and depicted in the illustration on 
plaie 649. It was noted that the second description is "proxima 
Saurauia excelsae" but origin was not indicated. Sprengel (1827) 
considered P. scabra HBK. as synonymous with Saurauia excelsa, 
but Dietrich (1843) recognized Saurauia scabra (HBK.) Dietr. as a 
distinct species, as also did Triana & Planchon (1862a,b). Buscalioni 
(1916) retained the second description as S. scabra HBK. and segre- 
gated the first description under the new name Saurauia hum- 
boldtiana Busc. According to the Code, the name S. humboldtiana is 
nomenclaturally superfluous and should be rejected, since an earlier 
legitimate name, S. scabra (HBK.) Dietr. (not of Poepp. ex Choisy, 
1855 = S. peruviana Busc.), is available. Taxonomically, little dif- 
ference exists between the two descriptions supplied by Humboldt et 
al., thus, there is no merit of their separation into different species. 

41. Saurauia tambensis Killip, J. Wash. Acad. Sci. 16: 571. 1926. 
Type: Hitchcock 21281 (US, holotype; NY, isotype). Figure 28. 

Shrubs; sparingly to copiously pubescent. Branchlets slender, terete to angular, 
prominently scarred, sparingly tuberculate to strigose pubescent. Leaves crowded 
behind tip of branchlets; blades oblong-obovate, acuminate at apex with acumen to 20 
mm. long, cuneate at base, setaceous-serrulate along margins, 18-25 cm. long, 7-9 cm. 
wide, membranaceous, somewhat smooth above, secondary veins 18-20 pairs, tertiary 
veins elevated, more prominent than lesser venation, scattered to sparingly strigillose 
to strigose pubescent above, sparingly to abundantly pubescent with trichomes of 
strigose to strigillose mixed with silvery slender-stellate types (especially along veins) 
beneath; petioles 2-2.5 cm. long, half -terete, 1.5-2 mm. in diameter, abundantly to 
sparingly strigose pubescent. Inflorescences slender and somewhat flexuous, loose, 
20-50-flowered, 12-20 cm. long, 4-5.5 cm. wide, pulverulent and densely pubescent 
with trichomes of setulose to shaggy-setulose and slender-stellate types, primary 
peduncle slender, 7-10 cm. long, bracts linear, to 5 mm. long. Flowers 15 mm. broad, 
buds to 4 mm. in diameter, pedicels to 10 mm. long, bracteoles linear, to 2 mm. long; 
sepals 5-6, oblong to oblong-obovate, obtuse to rounded, 4-5 mm. long, 3-4 mm. wide, 
exposed parts in bud sparingly pubescent with trichomes of setulose to shaggy-setulose 
and slender-stellate types, imbricated parts glabrous, all glabrous inside, marginally 
ciliate; petals 5, white, oblong-obovate, rounded and sometimes incised, 7-9 mm. long, 
5-6 mm. wide; stamens 30-40, filament 3-4 mm. long, anther 2 mm. long; ovary 
5-loculed, globose, 5-sulcate, glabrous, styles 5, 2.5 mm. long, stigmas simple. Berries 
not known. 

Habitat. Altitudes of 600-1,200 m. 

Distribution. Ecuador (Provinces of Tungurahua and El Oro). 



80 FIELDIANA: BOTANY 

Specimens examined. ECUADOR, EL ORO: between Portovelo 
(gold mine near Zaruma) and El Tambo, Sept., fl., Hitchcock 21281 
(NY, US). 

42. Saurauia excelsa Willd. Ges. Naturf. Freunde Berlin Neue. 
Schriften 3: 407. pi. 4. 1801. Type: Bredemeyer s. n. (COL, GH, NY, 
all photos). Figure 29. 

Saurauia excelsa var. moritziana Busc., Malpighia 25: 7. 1912. 

Type: Moritz 290 (K, P, US). 
Saurauia excelsa var. xanthotricha Busc., Malpighia 25: 7, 232. 

1912. Type: Funck & Schlim 106 (BM, F, K, P). 
Saurauia excelsa fina. veranii Busc., Malpighia 25: 218. 1912. 

Type: unknown, ex descr. 
Saurauja moritziana Turcz., Bull. Soc. Imper. Naturalistes 

Moscou 31(1): 243. 1858. Type: Moritz 290 (K, P, US). 
Saurauja floribunda Lind. & PL, Trois. Voy. Linden 1: 57. 

1863. Type: Funck & Schlim 1615 (G, P). 
Saurauja schlimii Sprague, Trans. Proc. Bot. Soc. Edinburgh 22: 

427. 1904. Type: Schlim 789 (F, G, K, P, US; COL, GH, 

photos). 
Saurauia brachybotrys Turcz. var. macrantha Busc., Malpighia 

28: 31. 1917., p. p. min. (typ. excl.). 

Saurauia scabra Busc., non Dietr., nee Poepp. ex Choisy, Mal- 
pighia 28:8. 1917, p. p. (incl. fma. veranii Busc., 1. c. 11). 
Saurauia rusbyi Britt. var. spectabilis Busc., Malpighia 30: 178. 

1927, p. p. (typ. excl.). 

Trees to 15 m. tall, diameter to 30 cm. at base, trunk crooked to straight, crown 
spreading; copiously pubescent. Branchlets subterete to angular, abundantly strigose 
to hirsute pubescent, rarely glabrescent; trichomes rusty brown. Leaves crowded 
behind tip of branchlets; blades oblong to obovate to broadly oblong-obovate, acute to 
very shortly and abruptly acuminate at apex, cuneate to narrowly cuneate at base, 
rarely oblique, finely serrulate to setaceous-serrulate along margins, (10-)15-35(-40) 
cm. long, (4-)8-15 cm. wide, chartaceous, medium to dark green above, dull to pale 
green beneath, scabrous above, secondary veins (15-)20-28(-30) pairs, tertiary veins 
elevated, more prominent than lesser venation, sparingly setose to hirsute pubescent 
along veins above, scattered to abundantly pubescent with trichomes of hirsute to 
tufted or shaggy mixed with radiate to stellate types (along and between veins) 
beneath; petioles (1.5-)2.5-4.5(-6) cm. long, 2-4 mm. in diameter, abundantly strigose to 
hirsute pubescent. Inflorescences somewhat straight, (15-)30-70(-150)-flowered, (8-) 
15-25(-30) cm. long, (3-)5-10(-15) cm. wide, abundantly to densely strigose to hirsute 
pubescent, primary peduncle (4-)7-14(-19) cm. long, bracts linear to subulate, 5-25 mm. 
long. Flowers 10-18 mm. broad, buds to 8 mm. in diameter, pedicels to 8 mm. long, 
bracteoles subulate to triangular, 3-5 mm. long; sepals 5, ovate to ovate-oblong, obtuse, 
6-10 mm. long, 3-4 mm. wide, exposed parts in bud densely pubescent with trichomes of 
strigose to hirsute mixed with stellate types, imbricated parts stellate pubescent, all 



SOEJARTO: SA URA VIA 81 

densely stellate pubescent inside, marginally to submarginally ciliolate to ciliate; 
petals 5, white to very rarely rose-white, oblong to obovate, obtuse, 5-8 mm. long, 3-5 
mm. wide; stamens (30-)50-85(-100), filament 2-3 mm. long, anther 2-3 mm. long, 
stigmas simple to capitate. Berries green to pale red, 5-sulcate, globose, 5-10 mm. 
across, 5-sulcate. 

Habitat. Cloud forest, temperate forest, secondary forest, moun- 
tain rain forest, residual forest, forest clearing and edge of forest, 
among rocks in moderately dense forest, along streams, in meadows 
and in parks, at altitudes of 500-2,500 m. 

Distribution. Venezuela (States of Distrito Federal, Aragua, 
Merida, Barinas), Colombia (Departments of Magdalena, Norte de 
Santander, Santander). 

Vernacular names. Cura (Merida: Gehriger), Moco or Moquillo 
(Venezuela: Perez- Arbelaez), Moquillo (Caracas: Williams). 

Specimens examined. VENEZUELA, DISTRITO FEDERAL: 
Caracas, fl.,Bredemeyer s. n. (COL, GH, NY, all photos), Moritz s. n. 
(FI); San Sebastian, ft., Funck 306 (G, P); Galipan, Jan., fl., Funck & 
Schlim 106 (F, K, P); Colonia Tovar, fr., Karsten s. n. (F, GH); 
Chacaito gorge around Caracas, April, fl., Pittier 9494 (US, VEN); La 
Mesa, above Quamita, National Park, Feb., fl., Pittier & Nakichen 
15734 (VEN); El Junquito, June, fl., Lasser 1096 (US, VEN), June, ft. 
fr., Steyermark 57028 (F, NY, US); Avilla, March, fl. fr., Delgado 44 
(US, VEN), April, fl., Lasser 73 (VEN); Los Infiernitos, May, fl., 
Tamayo 145 (US, VEN); S. slope of Cerro Avilla, Aug., fl., Vogl 138 
(F); Las Flores of Avilla, March, fl., Williams 9925 (F). ARAGUA: 
Colonia Tovar, Dec., fl., Allart 391 (A, NY, US, VEN), fl., Fendler 5 
(K), Jan., fl., Lasser & Foldats 4280 (VEN), March, fl., Pittier 9384 
(NY, P, US), Dec., fl., Pittier 10056 (NY, US, VEN); Alto de Choroni, 
Oct., fl., Chardon 202 (US, VEN), 8348 (VEN); highway between 
Maracay and Choroni, Jan., fl., Lasser 171 (US, VEN), Feb., fl., fr., 
Pittier 13926 (F, US, VEN); Chuao, Valle de En Medio, March, fl., 
Pittier 12132 (A, NY, US, VEN). MERIDA: Cerro de Las Flores, 
April, fl., Bernardi 452 (FI, NY, P); Chorros de Milla, May, fl., Ber- 
nardi 536 (NY); Sto. Domingo, Sept., fl., Bernardi 992 (NY, VEN); 
La Mucuy, fr., Breteler 3460 (VEN), April, fl., Gines 4770 (US), 
Sept., fl., Lasser 531, 606 (both in VEN), Sept., fl., Little, Jr. 15507 
(VEN), April, fl., Steyermark 55942 (F, VEN); Jaro(?), July, fl., 
Funck & Schlim 898 (G); Sierra de Merida, June, fl. fr., Funck & 
Schlim 1615 (G, P); Mucuruba, July, Gehriger 316 (A, F, NY, US, 
VEN); between La India and El Morro, May, fl., Ijjazz-Madriz 438 



82 FIELDIANA: BOTANY 

(YEN); Paramo de Aricagua, March, fl., Jahn 1023 (US); ? Colonia 
Tovar, Nov. f[.,Moritz 290 (K, P, US), Jan., fl., Smith et al. 3478 (F). 
MARINAS: vie. of Baruta, April, fl., Lasser 51 (F); vie. of Barinitas, 
April, fl., Lasser 51A (US). NO PRECISE LOCALITY: fl., Kuntze 74 
(F, NY); fl., Linden 38 (G, P); March, fl. fr., Pittier 44 (F); fl., Rohl 
s. n. (YEN). 

COLOMBIA, MAGDALENA: Rio Hacha, Sierra Nevada de Santa 
Marta, March, fl., Schlim 789 (F, G, K, P, US; COL, GH, photos). 
NORTE DE SANTANDER: Sarare region, basin of Rio Margua, 
between Campohermoso and Rio Negro, Nov., fl., Cuatrecasas 12902 
(COL, F, US), 13110 (F, US; COL, photo). SANTANDER: vie. of Las 
Vegas, Dec., fl., Killip & Smith 16081 (US), 16118 (A, NY, US); vie. 
of La Baja, Jan., fl., Killip & Smith 18322 (F, NY, US); NO PRE- 
CISE LOCALITY, Aug., fl., Uribe-Uribe 2433 (COL, ECON). 

NO PRECISE LOCALITY: fl., Moritz s. n. (GH, P, US). 

Members of S. excelsa are normally small trees of 4-10 m. tall, but 
Lasser and Foldats have observed a tree of 20 m. tall, and Pittier 
(Pittier 2132) has noted a trunk of 30-35 cm. in diameter. The great 
variability in the size of the inflorescences is remarkable; the usual 
size is 15-25 cm. long and 30-70-flowered, but a specimen (Lasser 
8351) with an inflorescence of 40 cm. long and ca. 150-flowered is 
represented among the collections. In the latter case, the peduncle is 
thick and woody (8 mm. in diameter), and the lateral cymes are 
widely spreading. 

43. Saurauia brachybotrys Turcz., Bull. Soc. Imper. 
Naturalistes Moscou 31(1): 245. 1858. Type: Linden 972 (BM; F, 
fragment; G, K, P; GH, photo). Figure 30. 

Saurauia brachybotrys var. macrantha Busc., Malpighia 28: 31. 

1917, p. p. maj. Type: Langlassee 57 (G, P, US). 
Saurauia brachybotrys var. scabra Busc., 1. c. 33, p. p. maj. 

Type: Linden 972. 
Saurauia goudotiana Lind. & PL, Trois. Voy. Linden 1: 58. 

1863. Type: Linden 972. 
Saurauia peduncularis Tr. & PI. var. veraniana Busc., Malpighia 

26: 26. 1913. Type: Karsten s. n. (BM) (Colombia: Bar- 

bacoas?). 

Trees to 15 m. tall, diameter to 25 cm. at base, trunk straight, bark narrowly 
splitting into irregular pieces, gray to brown, wood whitish yellow to light brown, 
crown open; copiously pubescent. Branchlets stout, terete to subangular, prominently 
scarred, abundantly strigose to setose pubescent, rarely aculeate, trichomes gray to 



SOEJARTO: SA URA VIA 83 

deep brown. Leaves crowded behind tip of branchlets; blades obovate to elliptic, acute 
to shortly acuminate at apex, cuneate to obtuse and often oblique but rarely with basal 
flap at base, serrate to serrulate along margins, (10-)15-30(-40) cm. long, (3.5-)8- 
15(-18) cm. wide, coriaceous, dark green above, green beneath, scabrous above, secon- 
dary veins (22-)24-30(-35) pairs, tertiary veins elevated, more prominent than lesser 
venation, sparingly pubescent with trichomes of strigose to setose types along and 
between veins above, abundantly pubescent with trichomes of shaggy to tufted mixed 
with hirsute and radiate types along and between veins beneath (epidermis pustu- 
late); petioles (1.5-)2-5(7.5) cm. long, 2.5 mm. in diameter, abundantly setose to hir- 
sute pubescent, rarely aculeate. Inflorescences usually straight, showy white, (20-) 
50-200-flowered, (8.5-)15-30(-37) cm. long, (3-)5-15(-25) cm. wide, densely to abun- 
dantly pubescent with trichomes of strigose to shaggy or tufted types, rarely aculeate, 
primary peduncle (5-)7-10(-13) cm. long, bracts triangular to subulate, to 8 mm. long. 
Flowers 15-25 mm. broad, buds to 7 mm. in diameter, pedicels to 10(-13) mm. long, 
bracteoles triangular, to 5 mm. long; sepals 5, greenish white to green, elliptic to 
ovate, obtuse, 7-8.5 mm. long, 4-6 mm. wide, exposed parts in bud densely pubescent 
with strigose to shaggy or tufted trichome types, imbricated parts pulverulent and 
densely stellate pubescent, all densely stellate pubescent within, marginally ciliolate 
to ciliate; petals 5, white to rosy, oblong to obovate to obcordate, frequently incised, 
often ciliolate apically, 9-11 mm. long, 4-7 mm. wide; stamens (65-)100-200(-240), 
filament 3-4 mm. long, anther 1.5-2.5 mm. long; ovary 5-loculed, ovoid, 5-sulcate, 
glabrous, styles 5, 0.5-5. 5(-7) mm. long, stigmas simple to capitate. Berries green, 
often with purplish red tinge, globose to obovoid, to 15 mm. across, 5-sulcate. 

Habitat. Dense forest, residual forest, edge of forest, wooded 
ridgetop, thickets, along streams, secondary forest, brushy second- 
growth slope, roadside, cornfield, meadow and open hillside, gardens, 
and wet mountain forest, at altitudes of 1,500-2, 500 (-3 ,000?) m. 

Distribution. Colombia (Departments of Antioquia, Caldas, 
Quindio, Tolima, Valle, Cauca, Narino, Putumayo). 

Vernacular names. Dulumoco (Caldas: Dryander; Valle: Cuat- 
recasas, Espinal), Moco (Cauca: Figueiras), Mosquito (Cauca: 
Perez-Arbelaez & Cuatrecasas), Moquillo (Sibundoy Valley: Bristol, 
Schultes, Soejarto), Yunush, Yenesha, or Je-nuss (Sibundoy Valley, 
Kamsa Indian names: Bristol, Schultes). 

Specimens examined. COLOMBIA, ANTIOQUIA: Alto de Minas, 
June, fl., Espinal 66 (MEDEL); Rio Force, Nov., fl., Espinal 774 
(ECON, UV); Cordillera Central, valley of Rio Nechi, 6 km. S. of 
Yarumal, March, fl., Fosberg 21597 (NY, US); between Sonson and 
Medellin, June, fl., Soejarto 2042, 2049, 2052 (all in ECON, GH, 
MEDEL); Angelopolis, Jan., fl., Toro 910 (NY); Titiribi, June, fl., 
Toro 1204 (MEDEL, NY). CALDAS: Salamina, June, fl., Tomas 1846 
(MEDEL, US); Sta. Rosa de Cabal, Los Alpes, Aug., fr., Dryander 
2740 (F, US); Cordillera Occidental, 1 km. N. of San Clemente, 13 km. 



84 FIELDIANA: BOTANY 

E. of Anserma, March, fl., Fosberg 21673 (NY, US). QUINDIO: La 
Palmilla, Nov., fl., Goudot s. n. (P: No. 87-103-66). TOLIMA: 
Mariquita, Jan., fl., Linden 972 (BM; F, fragment; G, K, P; GH, 
photo); El Libano, Alto de San Jose, July, fl., Garcia-Barriga 12250 
(COL, US). VALLE: basin of Rio Calima, El Cairo, between Darien 
and Mediacanoa, Jan., fr., Cuatrecasas 13871 (F, GH, UC, US); Re- 
presa del Calima, Aug., fr., Espinal 1964 (ECON, UV); basin of Rio 
Sanquinini, La Laguna, Dec., fr., Cuatrecasas 15583 (F); basin of Rio 
Cali, Pichinde, Alto de Las Brisas, Oct., fl., fr., Cuatrecasas 18250 (F), 
Jan., ft., Espinal 2324 (ECON, UV); Los Carpatos and El Olivo, July, 
fl., Cuatrecasas 21729 (ECON, F, UV); ridge of Cordillera, La Car- 
bonera, between Las Brisas and Alban, Oct., fl., Cuatrecasas 22155 
(ECON), 22282 (ECON, F); basin of Rio Dagua, La Elsa, March, fl., 
Cuatrecasas 23997 (F); vie. of Filiquis, Oct., fl., Dryander 1612(US); 
vie. of El Saladito, Cali, La Cumbre, Feb., fr., Cuatrecasas 19605 (F, 
GH, US, UV), June, fl., Soejarto 2056, 2057 (both in ECON, GH), 
May, fl., Pennell 5721 (US), June, fl., Killip et al. 39211 (F, US); El 
Silencio, Yanaconas, Killip & Garcia 33755 (A, COL, F, GH, S, US). 
CAUCA, CORDILLERA OCCIDENTAL: Oct., fr., Dryander 2072 
(US); Timbio, Hatoviejo, July, fl., Perez-Arbelaez & Cuatrecasas 
6091 (F, US); Carpinterias between Munchique and Altamira, July, 
fr., Perez-Arbelaez & Cuatrecasas 6148 (COL, F, US); El Tambo, fl., 
Yepes-Agredo 294 (F, US), Aug., fl., Idrobo & Fernandez 55 (COL, 
US); CORDILLERA CENTRAL: West facing slope, between 
Popayan and Purace, Dec., fl., Cuatrecasas 13769 (F, GH), July, fr., 
Garcia-Barriga & Hawkes 12700 (COL, US); Moscopan, basin of Rio 
San Jose, Aguabonita, Jan., fl., Cuatrecasas 235 13 (F); Tunia, Queb- 
rada de Bermejal, ranch of La Primavera, May, fl., Figueiras 8510; 
Cauca Valley, Rio Hondo to Popayan, July, fl., Killip 8258 (GH, NY, 
US); La Capilla (25 km. N. of Popayan), May, fl., Killip 38485 (US); 
vie. of Popayan, July, fl., Kjell von Sneidern 5645 (US), May and 
June, fl.,Lehmann 5537 (F, K, US); Miraflores, above Palmira, Jan., 
fl., Pittier 889 (US). NARINO: vie. of Ricaurte, road Paste to 
Tumaco, June, fl., Espinosa E2929 (NY), Oct., fl., Kjell von Sneidern 
4519 (F, US), Aug., fl. fr., Soejarto 1441, 1447, 1458, 1459, 1460, 
1461, 1462 (all in ECON, GH); La Union, July, fr., Soejarto et al. 
1204 (ECON, GH, PASTO); between La Union and Buesaco, July, 
fl., Soejarto et al. 1206 (ECON, GH, PASTO); Sandona, July, fl. fr., 
Soejarto 1180, 1181 (both in ECON, GH, PASTO); between Sandona 
and La Florida, fl., Soejarto 1182 (ECON, GH, PASTO); Pasto to El 
Encano, Paramo de El Tabano, fl., Mora 2893 (PASTO), 



SOEJ ARTO: SA URA UIA 85 

PUTUMAYO, VALLE DE SIBUNDOY, Jan., fl., Bristol 459 
(ECON, GH, PASTO), May, fr., 1048 (ECON, GH), Feb., fl., Chindoy 
105 (ECON), Feb., fl., Schultes 3203 (COL, ECON, GH); headwaters 
of Putumayo River, San Francisco, Jan., fl., Cuatrecasas 11578 (US); 
above Santiago of Sibundoy, Aug., fl. fr., Soejarto & Porter 513 
(ECON), 510, 512, 514, 518A, 519 (all in ECON, GH), 511, July, fl., 
Soejarto 1174 (ECON, GH, PASTO), Aug., fr., Soejarto 1533 (ECON, 
GH). NO PRECISE LOCALITY: Cordillera Occidental, Nov., fl., 
Langlassee 57 (G, K, P, US), March, fl., Marin 8 (MEDEL), Karsten 
s. n. (BM). 

MEXICO, OAXACA: fl., Galeotti 7235 (G; US-photo)(?). 

Saurauia brachybotrys is most commonly found in disturbed 
habitats: one reason for the rich herbarium representation. Because 
of its wide range, the species is often influenced by other species with 
which it is in contact, or sympatric in distribution, and this is re- 
flected in the variable nature of the indument. In the Antioquia 
region, for example, it is influenced by S. ursina, in the Valle-Caldas 
area by S. cuatrecasana, while in the Putumayo-Narino region by S. 
tomentosa. In several respects, S. brachybotrys is related to S. 
scabra and S. excelsa. However, the presence of setose to strigose, 
shaggy to tufted and stellate to radiate trichomes on the lower leaf 
surface, and the extremely high number of stamens (100-200) 
characterize S. brachybotrys. 

44. Saurauia isoxanthotricha Busc., Malpighia 25: 410. 
1912. Type: Mathews 1216 (K, holotype). Illustration: Malpighia 
25: pi. 5, fig. 6. 1912; Caldasia 2:33, figs. 7-12. 1943. 

Saurauia anolaimensis R. E. Schultes & Garcia-Barriga, Caldasia 
2: 27. 1943. Type: Garcia-Barriga 8994 (COL, holotype; US, 
isotype). 

Saurauia intonsa R. E. Schultes, Caldasia 2: 318. 1944. Type: 
Fosberg 19914; ex descr. 

Shrubs to 4 m. tall, erect to ascending or curving or straggling, moderate to much- 
branched, often spreading, rarely bushy; copiously pubescent. Branchlets slender, 
terete, abundantly to densely pubescent with trichomes of hirsute to sericeous-hirsute 
types, trichomes golden-yellow to golden-brown, sometimes dark reddish to purplish 
brown, moderately rigid, to 10 mm. long. Leaves clustered behind tip of branchlets; 
blades elliptic to obovate-elliptic, caudate at apex with acumen to 45 mm. long, broadly 
cuneate to obtuse at base, serrate to hirsute-serrulate along margins, (4.5-)10-25(-35) 
cm. long, (2-H-9(-15) cm. wide, chartaceous to membranaceous, dark green to dark 
reddish brown above, pale green to golden-reddish brown beneath, somewhat smooth 
above, secondary veins (7-)9-15(-25) pairs, tertiary veins elevated, more prominent 
than lesser venation, abundantly to densely pubescent with trichomes of hirsute to 



86 FIELDIANA: BOTANY 

sericeous-hirsute or hirtellous types along and between veins above, abundantly to 
densely pubescent with trichomes of hirsute to sericeous-hirsute or hispid types (to 5 
mm. long along midrib, to 3 mm. long between veins) beneath; petioles (0.75-)2- 
3.5(-4.5) cm. long, 1-4 mm. in diameter, densely hirsute pubescent. Inflorescences 
straight, may be erect, (5-)10-50-flowered, (6-)10-18(-23) cm. long, 3.5-6 cm. wide, 
densely hirsute to sericeous-hirsute pubescent, primary peduncle 3-10 cm. long, bracts 
linear to subulate, to 15 mm. long. Flowers 15-25 mm. broad, buds to 6 mm. in 
diameter, pedicels to 6(-8) mm. long, bracteoles linear, to 7 mm. long; sepals 5, elliptic 
to oblong, subacute to obtuse, 7-10 mm. long, 6-8 mm. wide, exposed parts in bud 
densely pubescent, trichomes of setose to sericeous-hirsute and often mixed with 
stellate types, imbricated parts scattered to abundantly stellate pubescent, all spar- 
ingly stellate pubescent on upper half and glabrous on lower half inside, marginally 
ciliolate; petals 5, white to rosy, elliptic to narrowly ovate to oblong, obtuse, 12-15 mm. 
long, 6-8 mm. wide; stamens 20-30(-35), filament 2-3 mm. long; ovary 5-loculed, 
globose, 5-sulculate, glabrous, styles 5, obsolete to 4.5 mm. long, stigmas simple to 
capitate. Berries 5-loculed, green with purplish red tinge, globose, to 8 mm. across, 
5-sulcate. 

Habitat. Cloud forest, wet mountain forest, wet riverbank, sub- 
paramo, mountain forest clearing, wet scrub forest, at altitudes of 
2,000-3,000 m. 

Distribution. Colombia (Departments of Santander, Cun- 
dinamarca, Huila, Putumayo), Ecuador (Province of Imbabura), 
Peru (Department of Amazonas). 

Specimens examined. COLOMBIA, SANTANDER: vie. of 
Charta, Feb., fl., Killip & Smith 19319 (NY, US). CUNDINA- 
MARCA: Santana station, above Sasaima, July, fl., Dugand & 
Jaramillo 3875 (US); San Francisco, ranch of El Carmero, between 
Subachoque and San Francisco, Jan., fl., Garcia-Barriga 11014, 
11016, 11022 (all in US); San Bernardo of Sasaima, between Que- 
bradas La Maria and La Victoria, Jan., fl., Garcia-Barriga 12565 
(US); Sabaneta of San Francisco, W. of Sabana de Bogota, May, fl fr., 
Uribe-Uribe 4801 (COL, ECON, GH), 4802 (COL, ECON). 
BOYACA: Chiquinquira, July, fl., Ariste-Joseph A967 (P, US). TO- 
LIMA: road El Libano to Murillo, km. 11 of the highway, July, fl., 
Garcia-Barriga 12253 (US); Buenavista to Azufral, old Quindio 
trail, Aug., fl., Killip & Hazen 9851 (NY). HUILA: Cordillera Cen- 
tral, 30 km. NW. of Palermo, Oct., fl., Little, Jr. 8801 (P); Quebrada 
Guache, 3 km. SW. of Acevedo, canyon bottom, Aug., fl., Little, Jr. 
8482 (NY, US). PUTUMAYO: Paramo de San Francisco (La Depre- 
sion), road San Francisco to Mocoa, Jan., fl., Fray-Miguel 32 (F); 
Cerro Portachuelo, vie. of Buenos Aires and Alto de Siberia, July, fl., 
Soejarto 1065, 1146, 1169 (all in ECON, GH, PASTO), Sept., fl., 
1531, 1532, 1534, 1535, 1538, 1548, 1551, 1563, 1565, 1566 (all in 



SOEJARTO: SA URA VIA 87 

ECON, GH), veg., 1536, 1537, 1555 (all in ECON, GH); below 
Paramo de Capuchino, Santaclara, roadside Pasto-Sibundoy, fl., 
Soejarto 1510, 1522, 1523 (all in ECON, GH). NO PRECISE LO- 
CALITY: fl., Bonpland s. /i.(P); fl., Mutis 2213 (US). 

ECUADOR, IMBABURA: Cordillera Oriental, Alegria, E. of Vol- 
can de Cayambe, May, fl., Drew E-237 (US). 

PERU, AMAZONAS: Bagua, along Quebrada Tambillo, valley of 
Rio Maranon, above Cascadas de Mayasi, Sept., fl., Wurdack 2052 
(NY, US). NO PRECISE LOCALITY: Pangsa (?), fl., Mathews 1216 
(K). 

The collection Soejarto 1551 from Cerro Portachuelo (Putumayo) is 
remarkably identical in practically every respect to Mathews 1216, 
collected from Peru, which was described by Buscalioni as S. isoxan- 
thotricha. It is characterized by the apically caudate leaves and the 
complete absence of any multicellular branched trichomes, except on 
the sepals. Other Colombian collections, notably Soejarto 1555, 1563, 
1565, 1566, are without question referrable to this species, due 
primarily to highly similar type of indument, the apically caudate 
leaves, and the floral characters. Several other collections, however, 
vary with regard to the size and density of the leaf indument between 
the veins beneath. The characteristic hirsute to setaceous-hirsute 
trichomes along the branchlets, the petioles, and along the midrib 
and the inflorescences are very useful for identification. The color of 
the branchlets, shoots, and young leaves of the plants collected in the 
Cerro Portachuelo region is usually dark to bright maroon-red, due to 
the dense indument of the same color, on these parts of the plant. 

Several of the specimens collected from Putumayo (e.g., Soejarto 
1146, 1536, 1537) have rather short and somewhat stiff (hispid) 
trichomes on the lower surface of the leaves, in contrast to the rela- 
tively weak and long (hirsute) trichomes found in Soejarto 1551. In 
addition, they have scattered radiate to stellate trichomes on the 
lower leaf surface. Field observations seem to indicate that these 
individuals, and others found along the road cut, may represent 
hybrid segregates, probably involving S. putumayonis. The speci- 
men Fosberg 19914, described by Schultes as S. intonsa, has acumi- 
nate leaf apex and somewhat attenuate leaf base, but other charac- 
ters agree very well with the range of variation of S. isoxantho- 
tricha. 

45. Saurauia meridensis Steyermark, Fieldiana (Botany) 28(1): 



88 FIELDIANA: BOTANY 

368. 1952. Type: Steyermark 56344 (F, holotype; YEN, isotype). 
Figure 31. 

Shrubs to 3 m. tall, copiously pubescent. Branchlets abundantly setose to hirsute 
pubescent (young leaves and shoot densely sericeous pubescent); trichomes red- 
brown, to 4 mm. long, golden brown. Leaves clustered behind tip of branchlets; blades 
obovate, shortly and abruptly acuminate at apex with acumen to 10 mm. long, obtuse 
to cuneate at base, rarely oblique, setaceous- to ciliate-serrulate along margins, 15-23 
cm. long, 6-10.5 cm. wide, chartaceous to subcoriaceous, in dry state grayish 
maroon-brown and scabrous above, light olive-brown beneath, secondary veins 15-24 
pairs, tertiary veins elevated, more prominent than lesser venation, abundantly to 
sparingly setose to hirsute pubescent along and between veins above and beneath 
(trichomes denser and longer along major veins, but branched trichome types absent 
over both surfaces); petioles 1-3 cm. long, 2-3 mm. in diameter, half-terete, densely 
setose to hirsute pubescent. Inflorescences crowded near tip of branchlets, spreading, 
somewhat straight, lax, 20-80-flowered, 10-19 cm. long, 7-11 cm. wide, pubescence 
similar to that on leaves, primary peduncle 7-9 cm. long, bracts linear-triangular, 
often outcurved to revolute, to 20 mm. long. Flowers ca. 15 mm. broad, buds to 8 mm. 
in diameter, pedicels 10-15 mm. long, bracteoles triangular, to 4 mm. wide, sepals 
5(-7), exposed parts in bud densely pubescent, trichomes of setose mixed with minute 
stellate types, imbricated parts pulverulent and stellate pubescent, all densely stel- 
late pubescent inside, marginally to submarginally ciliolate; petals 5(-7), white, ob- 
long, obtuse to rounded, 5-6 mm. long, 3-4 mm. wide; stamens 30-40, filament 3 mm. 
long, anther 2.5-3 mm. long; ovary 5-loculed, globose, 5-sulcate, glabrous, styles 5(-7), 
1 mm. long, stigmas simple. Berries not known. 

Habitat. Woods along canyon, at altitudes of 1,065-1,820 m. 
Distribution. Venezuela (States of Merida and Tachira). 
Vernacular names. Guaco morado (Merida: Steyermark). 

Specimens examined. VENEZUELA, MERIDA: Canagua River, 
near Canagua, between Mucuchachi and Canagua, May, fl., 
Steyermark 56344 (F, VEN). TACHIRA: Headwaters of Quinimari 
River, below Cerro Las Copas, 20 km. S. of San Vicente de la Re- 
vancha, Jan., fl., Steyermark 100952 (ECON, VEN). 

S. meridensis appears to be closely allied to S. isoxanthotricha , 
particularly with respect to leaf pubescence. The former, however, 
has a non-caudate leaf apex and lax inflorescences with long- 
pedicelled flowers, in contrast to the caudate leaf apex and more 
compact inflorescences with short-pedicelled flowers of the latter. 

The following field notes by Steyermark (Steyermark 56344) may 
be helpful in field identification: "Shrub 10 ft. tall; petals full white; 
leaves firmly membranaceous, dull olive-green and rugose above, 
pale green below; midrib below brownish at base; hairs brick red or 
rufous-buff." 



SOEJARTO: SA URA VIA 89 

Steyermark 100952 has rugulose upper leaf surface and somewhat 
lacunose lower leaf surface, as that in Saurauia formosa, but other 
characters fall within the range of variation of S. meridensis. 

46. Saurauia prainiana Busc., Malpighia 25: 248, pi. 6, fig. 10. 
1912. Type: Ule 6530 (K, lectotype; L, isolectotype; COL, GH, NY, 
photos). 

Saurauia pastasana Diels, Biblioth. Bot. 116: 107. 1937. Type: 

Diets 1039 (n. v.), ex descr. 
Saurauia sydowii Sleumer, Notizbl. 15: 374. 1941. Type: Sydow 

891 (K, S). 

Saurauia consimilis Sleumer, Notizbl. 15: 373. 1941. Type: 
Sydow 264 (K). 

Shrubs to small trees to 10 m. tall, often bushy; copiously pubescent. Branchlets 
slender, terete, abundantly strigose to rarely setose pubescent; trichomes light to 
reddish brown, to 3(-5) mm. long. Leaves crowded behind tip of branchlets; blades 
obovate to elliptic-obovate, cuspidate at apex with acumen to 10 mm. long, 
setaceous-serrate to -serrulate along margins, setae rigid and sharp-pointed, 15.5-30 
cm. long, 6.5-14 cm. wide, chartaceous, pale green above, grayish green beneath (in dry 
state dark brown to sooty above, light to dark olive-brown beneath), somewhat 
scabrous above, secondary veins 15-22 pairs, tertiary veins elevated, more prominent 
than lesser venation, sparingly setulose to strigillose pubescent along and between 
minor veins and abundantly strigose to appressed-setose (rarely setose) pubescent 
along major veins above, sparingly to abundantly setose pubescent along and between 
minor veins and abundantly strigose to appressed-setose or rarely setose pubescent 
along major veins (with stellate trichomes often present) beneath; petioles furrowed, 
1.5-4 cm. long, 2-3 mm. in diameter, sparingly to abundantly strigose to 
appressed-setose pubescent. Inflorescences straight, (7-)15-40(-50)-flowered, 
6.5-20(-30) cm. long, 2-6(-10) cm. wide, abundantly to densely strigose to setose 
pubescent, primary peduncle 2.5-8.5 cm. long, bracts subulate, to 10 mm. long. Flowers 
12-17 mm. broad, buds to 5 mm. in diameter, pedicels 1-4 mm. long, bracteoles 
triangular, to 2.5 mm. long; sepals 5, pale green, suborbicular to broadly ovate to 
elliptic, 4-5 mm. long, 3-4.5 mm. wide, exposed parts in bud abundantly pubescent with 
trichomes of strigose to shaggy-strigose types, imbricated parts densely stellate 
pubescent, all densely stellate pubescent on upper portion and glabrous on lower half 
inside, marginally to submarginally ciliolate to ciliate; petals 5, white, oblong, 
rounded and slightly incised, 11-12 mm. long, 3.5-5.5 mm. wide; stamens 20-40, 
filament 3-3.5 mm. long, anther 2.5-3.5 mm. long; ovary 5-loculed, subglobose to ovoid, 
5-sulcate, glabrous, styles 5, obsolete to 5 mm. long, stigmas 5, simple to capitate. 
Berries green, 5-loculed, globose, to 7 mm. across, 5-sulcate. 

Habitat. Rain forest, humid forest, fringe of forest, riverbank, 
road cut, at altitudes of 400- 1,600 (-2 ,500) m. 

Distribution. Ecuador (Provinces of Pichincha, Napo-Pastaza, 
Tungurahua, and Santiago-Zamora), Peru (Department of Loreto). 



90 FIELDIANA: BOTANY 

Specimens examined. ECUADOR, PICHINCHA: Unido, Oct., 
fr., Sydow 264 (K). NAPO-PASTAZA: Cerro Antisana, Shinguipino 
forest between Napo and Tena rivers, 8 km. SE. of Tena, Sept., fl., 
Grubb et al. 1619 (K, NY); Canton Napo, overhanging Masaguali 
River, April, fl., Mexia 7227 (UC, US); Pastaza River, May, fl. fr., 
Rimbach 261 (F, MICH, NY, US); valley of Pastaza River, vie. of 
Mera and Shell Mera, Feb., fl. fr., Rimbach 490 (S), fl., Schultze- 
Rhonhof2948 (K), Sept., fl. fr., Soejarto & Hernandez 1343, 1345 
(both in COL, ECON, GH, PASTO), 1344 (ECON, GH, PASTO), 
1346, 1347, 1349 (all in ECON, GH), 1348 (GH); Puyo, Feb., fl., 
Sydow 891 (K, S). TUNGURAHUA: Between Mapoto and Mar- 
garitas rivers, along old Canelos trail, March, fl., Penland & Sum- 
mers 184 (F, US); between Rio Verde and Rio Negro (on Banos to 
Puyo route), Sept., veg., Soejarto & Hernandez 1350 (ECON, GH), 
1351 (ECON, GH, PASTO). 

PERU, LORETO: Pumayacu, between Balsapuerto and 
Mayobamba, Aug. -Sept., fl.,Klug3176 (A, F, G, GH, K, NY, S, US); 
Cerro de Escaler, Nov., fl., Ule 6530 (K, L; COL, GH, NY, photos). 

VII. Ser. LANATAE Soejarto, ser. nov. 

Strigosae Busc., Malpighia 25: 228. 1912, p. p. min. 
Ruitzianae Busc., 1. c. 

Folia dense lanata vel velutina subtus, epidermes inferiores per pube occulti mag- 
nificatione inferior. 

1. Inflorescences usually compact with densely aggregated and shortly pedicelled 
flowers, flowers less than 25 mm. broad, stamens usually less than 90. 

2. Trichomes along branchlets and petioles mostly straight and silky, flowers 15-25 
mm. broad, stamens 45-90 47. S. biserrata 

2. Trichomes along branchlets and petioles curved, often coarse and prickly, flow- 
ers 20-25 mm. broad, stamens (25-)30-40(-50) 48. S. ursina 

1. Inflorescences rather loose and spreading, flower pedicels up to 18 mm. long, 
trichomes mostly long and flexuous, flowers (20-)25-40(-50) mm. broad, stamens 
100-200 49. S. bullosa 

47. Saurauia biserrata (R. & P.) Spreng., Syst. Veg. ed. 16, 4, Cur. 
Post. 2: 211. 1827; Macbr., Fl. Peru 3A(2): 679. 1956. Based upon 
Palaua biserrata R. & P. Illustration: DC., Mem. Soc. Phys. Geneve 1: 
pi. 4. 1822. 

Palaua lanceolata R. & P., Syst. Veg. Fl. Peruv. Chilens. 181. 
1798. Type: Ruiz & Pavon s. n. (G: sheet annotated 662). 
Palaua biserrata R. & P., 1. c. Type: Pavon s. n. (G, lectotype). 
[Note on sheet: Palaua biserrata Peru; Palava biserrata R. 
& P.; Apatelia biserrata DC., Prodr. 1: 526.] 



SOEJARTO: SA URA VIA 91 

Palaua hirsuta R. & P. in Ruiz, Travels of Ruiz, Pavon and Dombey 

in Peru 78. 1940, nom. nud. [Note: Field Museum specimen 

no. 845498 is annotated as Palaua hirsuta.] 
Apatelia lanceolata (R. & P.) DC., Mem. Soc. Phys. Geneve 1: 427. 

1822; Prodr. 1: 526. 1824. Based upon Palaua lanceolata 

R. &P. 
Apatelia lanceolata var. peduncularis DC., Mem. Soc. Phys. 

Geneve 1: 427. 1822. Type: Pavon s. n. (G: sheet annotated 

192). 
Apatelia biserrata (R. & P.) DC., Mem. Soc. Phys. Geneve 1: 428. 

1922; Prodr. 1: 526. Based upon Palaua biserrata R. & P. 
Saurauja lanceolata DC., Mem. Soc. Phys. Geneve I: pi. 4, not the 

description on p. 421 (=S. lanceolata, Java). 
Sauravia ruiziana Steud., Nomencl. Bot. ed. 2: 516. 1841, nom 

nud. (excl. syn. S. lanceolata DC., Java). 
Saurauia ruiziana var. weberbaueri Busc., Malpighia 25: 442. 

1913. Type: Weberbauer 2020 (COL, GH, NY, all photos). 

Saurauia scabra Poepp. exChoisy var.prainiana Busc., Malpighia 
27: 493. 1916. Type: Ruiz & Pavon s. n. (P) (Peru: Chin- 
chao). 

Saurauia pseudoruitziana Busc., Malpighia 30: 244. 1927. Type: 
Pearce s. n. (K) (specimen purchased from South America). 

Shrubs to small trees to 5 m. tall, erect, very rarely spreading; copiously pubescent. 
Branchlets terete, leaf scars prominent, pubescence sparse to dense, trichomes of 
sericeous type, to 8 mm. long, silky brown. Leaves crowded behind tip of branchlets; 
blades elliptic to obovate, acuminate at apex with acumen to 10 mm. long, narrowly 
cuneate to subrotundate at base, finely serrulate to setaceous-serrulate along margins, 
(9-)15-30(-33) cm. long, (4-)5-10(-13) cm. wide, coriaceous, dark green to brown above, 
brown to reddish brown beneath, scarcely scabrous above, secondary veins elevated, 
more prominent than lesser venation, sparingly sericeous to sericeous-strigose pubes- 
cent along veins above (pubescence denser along midrib), densely lanate with 
trichomes of sericeous to loriform (to 10 mm. long) mixed with stellate types along and 
between veins; petioles (l-)2-3.5(-4.5) cm. long, 3-5 mm. in diameter, densely sericeous 
pubescent. Inflorescences straight, (8-)10-40)(-75)-flowered, (6-)10-20(-35) cm. long, 
1.5-5(-8) cm. wide, densely sericeous pubescent, primary peduncle (2-)6-10(-18) cm. 
long, bracts subulate to foliaceous, 8-30 mm. long. Flowers 15-25 mm. broad, densely 
aggregated, buds to 8 mm. in diameter, pedicels 1-8 mm. long, bracteoles linear to 
triangular, to 10 mm. long, sepals 5-6, 8-12 mm. long, 5-9 mm. wide, elliptic to 
suborbicular, acute to obtuse, exposed parts in bud densely sericeous pubescent, imbri- 
cated parts densely stellate pubescent, all densely stellate pubescent inside, margi- 
nally ciliolate to ciliate; petals 5-6, white, spatulate, rounded, 9-12 mm. long, 6-8 mm. 
wide; stamens 45-90(-150), filament 3.5-5 mm. long, anther 2.5-3 mm. long; ovary 
5-6-loculed, ovoid, 5-6-sulcate, glabrous, styles 5-6, obsolete to 5 mm. long, stigmas 
simple to capitate. Berries 5-6-loculed, globose, 8-15 mm. across, 5(-6)-sulcate. 



92 FIELDIANA: BOTANY 

Habitat. Cloud forest, humid mountain forest, dense forest, bor- 
der of rain forest, evergreen subxerophytic bush- wood, shrubbery in 
open places, along riverbank, full exposure and semi-shade, at al- 
titudes of 800-2,300 m. 

Distribution. Peru (Departments of Huanuco, Junin, and 
Ayacucho). 

Specimens examined. PERU, HUANUCO: Between Huanuco 
and Pampayacu, W. side of the Andean Mts., Jan., fl., Kanehira 31 
(A; F, fragment); Huacachi, vie. of Muna, May-June, fr., Macbride 
4181 (F, US); Huanuco, vie. of Carpish, Nov., fl., Sandeman 3491 
(K); above Chinchao, along road Huanuco to Tingo Maria, Aug., fl., 
Soejarto & Wild 1424, 1426 (both in ECON, GH), Chicoplaya, fl., 
Pavon s. n., fr., Pavon s. n. (both in G, with note: Palaua lanceolata 
sp. nov. de Chicoplaya. Fl. Peruv. sin lam. no. 11 by Pavon?); 
Cuchero, fl., Pavon s. n. (F); Muna, veg., Ruiz & Pavon s. n. (F, 
fragment; sheet no. 843573); Pillao, fl., Ruiz & Pavon s. n. (F), fr., 
Ruiz & Pavon s. n. (F). JUNIN: Tarma, vie. of Oreja Capelo, on road 
between Tarma and San Ramon, valley of Palca River, Jan., fl., 
Hodge 6252A (US: sheet no. 2056174); Utcuyacu, on road Palca to 
San Ramon, Jan., fr., Hodge 6252B (US: sheet no. 2268219); Pichis 
trail, between San Nicolas and Azupizu, July, fl., Killip & Smith 
26093 (F, NY, US); Chanchamayo Valley (Tarma), June, fl., Con- 
stance & Tovar2322 (US), Dec., fl., Martinet 1272 (P), fl., Raimondi 
12493 (USM); Huacapistana, June, fl., Killip & Smith 24203 (US), 
24183 (F, NY, US), Oct., fl., Sandeman 4563 (F, K), June, fl., 
Sandeman s. n. (K; sheet no. H2668/66-18); La Merced, Aug., fl., 
Soukup 3369 (F, US); Tarma, Weberbauer 2020 (COL, GH, NY, all 
photos); between Huacapistana and Palca, fl., Weberbauer 2040 (G). 
AYACUCHO: Ayma, between Huanta and Apurimac river, May, fl. 
fr., Killip & Smith 22768 (F, NY, US); Choimacota Valley, vie. of 
Quillomicto, Feb., fl., Weberbauer 7545 (F, US). NO PRECISE LO- 
CALITY: ? Junin, fl., Dombey 952 (P: sheet numbered 87-34-66); 
South America (Peru?), fl. fr.,Pearce s. n. (K: purchased April 1884); 
fl., Poeppig 125 (G: annotated as Saurauja ruiziana Steud. , Apatelia 
lanceolata DC. var. ft. Icon; Choisy scripsit); fl., Poeppig 1 395 (BM, L); 
Andean shrubbery, June, fl., Woytkowski 6346 (GH); fl. fr., Pavon s. 
n. (Fl, G; GH-photo) (Annotation on G sheet: 1. Polyandr. 5-gyn.; 
Palava lanceolata R. & P., Apatelia lanceolata DC.; Perou, M. Pavon, 
1827; 2. Palaua lanceolata ex Peru); fl., Pavon s. n. (P: sheet num- 
bered 87-33-66); fl., Pavon s. n. (G: this has been selected as a lec- 
totype of P. biserrata R. & P.); fl., Pavon s. n. (G) (sheet annotated: 



SOEJARTO: SA URA VIA 93 

Peruvia, herb. Pavon); fl., Pavon s. n. (G) (sheet annotated: 192 Gen 
Nov. ex polyadelphiis Perou, Pavon; Apatelia lanceolata (3 DC., scrip- 
sit APDC.); fl., Pavon s. n. (G: sheet annotated 11); veg., Pavon s. n. 
(F: fragment, sheet numbered 695109); fr., Pavon s. n. (F: fragment, 
sheet numbered 686267); fl., Ruiz & Pavon s. n. (F; sheet numbered 
844793); veg., Ruiz & Pavon s. n. (F: sheet number 844821); fr.,Ruiz 
& Pavon s. n. (F: sheet numbered 845498, annotated as Palaua 
hirsuta, P. lanceolata)', fl., Ruiz & Pavon s. n. (G; NY, photo) (G sheet 
is annotated as Palaua lanceolata sp. n. by Ruiz & Pavon? and 
Palava lanceolata R. & P., Apatelia lanceolata a DC., scripsit APDC; 
this specimen is designated as lectotype of Palaua lanceolata R. & P.); 
fl., Ruiz & Pavon s. n. (F: no. 845432); fr., Pavon s. n. (K). 

The materials collected and, presumably, examined by Ruiz & 
Pavon, and later examined by De Candolle and Buscalioni, made it 
possible to clarify the correct taxonomy and nomenclature of this 
species. Because no types were ever mentioned by Ruiz & Pavon or 
by later authors, it has become necessary to select lectotypes from 
these specimens. In the process, the most useful clues were hand- 
written annotations by Ruiz & Pavon (as Palaua, instead of Palava), 
by De Candolle, and by Buscalioni. Since most of these specimens do 
not bear collector's or any sequential number, any other identifying 
number and/or annotation found on the herbarium sheet has been 
indicated in citing the botanical institution, for the sake of clarity. 

Taxonomically, S. biserrata is characterized by the dense sericeous 
pubescence, by the often aggregated flowers on the inflorescences, 
and by the moderate to high number of stamens. Young leaves are 
often reddish to purplish red. 

48. Saurauia ursina Tr. & PL, Ann. Sci. Nat. ser. 4, 18: 265. 

1862; Prodr. Fl. Nov. Granat. 1: 262. 1862. Type: Triana s. n. (K, 

lectotype; G, P, iso-lectotypes; F, fragment of leaf; COL, US, photos). 

Figure 32. 

Saurauia ursina fma. strigosa Busc., Malpighia 25: 417. 1913. 

Type: Jervise s. n. (K). 
Saurauia ursina fma. veranii Busc., 1. c. 418. 1913. Type not 

indicated ex descr. 
Saurauia ruitziana auct. (non Steud.), 1. c. 425. 1913, p. p. 

Small trees to 7 m. tall, trunk straight to crooked, crown open; copiously pubescent. 
Branchlets slender, terete to subangular, densely pubescent, trichomes coarse to 
prickly, of paleaceous to rarely sericeous types, light to dark reddish brown in color, to 6 
mm. long, curved towards apex of branchlet, thickened to flattened at base and turning 



94 FIELDIANA: BOTANY 

very slender and pointed or bent at apex. Leaves crowded to rarely clustered behind tip 
of branchlets; blades elliptic to elongate-obovate, acuminate at apex with acumen to 15 
mm. long, cuneate to rarely obtuse or oblique at base, serrulate to setaceous-serrulate 
along margins, (7-)10-25(-35) cm. long, (3.5-)4.5-10(-13.5) cm. wide, coriaceous to 
strongly coriaceous, dark greenish brown to maroon above, gray-brown to deep reddish 
brown beneath, moderately scabrous above, secondary veins (16-)20-25(-29) pairs, 
tertiary veins elevated, more prominent than lesser venation, abundantly to sparingly 
setose to hirsute pubescent along and between veins above, densely velutinous pubes- 
cent with trichomes of loriform frequently mixed with stellate to dendroid types 
beneath, petioles (0.75-)2-4.5(-6) cm. long, 2-4.5 mm. in diameter, densely hirsute to 
prickly-hirsute pubescent. Inflorescences straight to ascending, (7-)15-30(-60)- 
flowered, (3.5-)6-12(-15) cm. long, 1.5-6 cm. wide, densely hirsute to prickly-hirsute 
pubescent (particularly along peduncle), primary peduncle (1.5-)-5 cm. long, bracts 
subulate to triangular, to 10 mm. long, very rarely foliaceous, to 25 mm. long. Flowers 
often aggregated on inflorescence, 20-25 mm. broad, buds to 7 mm. in diameter, 
pedicels 0.5-3 mm. long (to 10 mm. in fruiting stage), bracteoles triangular, to 3 mm. 
long; sepals 5, suborbicular-ovate to elliptic, obtuse to acute, 7-9 mm. long, 5-7 mm. 
wide, exposed parts in bud densely pubescent with trichomes of sericeous to hirsute 
types, imbricated parts densely stellate pubescent, all densely stellate pubescent 
inside, marginally ciliolate; petals 5, white, oblong to obovate-oblong, obtuse to in- 
cised, 10-12 mm. long, 4.5 mm. wide; stamens (25-)30-45(-50), filament 4-5 mm. long, 
anther 3.5-4 mm. long; ovary 5-loculed, globose, 5-sulcate, glabrous, styles 5, obsolete 
to 5(-6) mm. long, stigmas simple to capitate. Berries green, 5-loculed, globose, to 10 
mm. across, 5-sulcate. 

Habitat. Humid mountain forest, open forest, woodland border, 
scrub forest, subparamo, secondary forest, riverbank, creek, open 
areas, gardens, at altitudes of 1,500-2,600 m. 

Distribution. Colombia (Departments of Antioquia, Santander 
and Cundinamarca). 

Vernacular names. Dulumoco, Guasco (widely used names in 
Antioquia). 

Specimens examined .COLOMBIA, ANTIOQUIA: Guadalupe, 
Nov., fl., Daniel 2622 (MEDEL, US); El Penol, Aug., fr., Daniel 405 
(US); Guarne, Piedras Blancas and vie., July fl., Cabrera 154 
(MEDEL), March, fr., Espinal 418 (MEDEL), Aug., fl. fr., Espinal 
1237 (MEDEL), Sept., fl. fr., Soejarto 2104, 2111 (UNIV. OF AN- 
TIOQUIA), Aug., fl., Soejarto 4239, 4242, 4244, 4246, 4252, 4253 
(all in UNIV. OF ANTIOQUIA), fr., 4243, 4247 (both in UNIV. OF 
ANTIOQUIA); Valle de Aburra, vie. of Medellin, Dec., fl., Archer 
1089 (MEDEL, NY, US), Oct., fl., Domingo-Penazos 3820 (MEDEL, 
US), Oct., fl., Molina 34 (MEDEL, US), fl., Soejarto & Rivera 2053 
(ECON, GH, MEDEL), Oct., fr., Torres 313 (MEDEL); Sta. Helena, 
above Medellin on road to Rionegro, June, fl., Soejarto 4531 (UNIV. 
OF ANTIOQUIA); Las Palmitas, fl., Barkley et al. 22 (US); road 



SOEJARTO: SA URA VIA 95 

Boqueron to San Cristobal, June, ft., Barkley & Correa 93 (US), Oct., 
fl., Hodge 6599 (MEDEL), May, fl., Hodge 6856 (MEDEL); road 
Caldas to Primavera, Oct., fr., Ruiz s. n. (MEDEL); Alto de 
Boqueron, Sept., fl., Toro 30 (MEDEL); Rionegro, fl., Jervise s. n. 
(K), Dec., fl., no collector 309 (MEDEL); La Ceja and vie., Dec., fl., 
Daniel 2170 (MEDEL, US), June, fl., Soejarto & Rivera 2040 
(ECON, GH, MEDEL); Pozo Rico, June, fl., Soejarto & Rivera 2051 
(ECON, GH, MEDEL); vie. of El Retiro, fl. fr., Uribe-Uribe 4346 
(COL, ECON); vie. of Hoyo Rico, Sept., fl., Valbuena & Barkley 
18A213 (NY, US); Angelopolis, Nov., fl., Barkley & Gutierrez 1681 
(MEDEL); Tamesis, Feb., fr., Toro 956 (MEDEL, NY); Fredonia, 
June, fl., Toro 1039 (MEDEL, NY); Salto de Buey, Jan., fl., Daniel s. 
n. (MEDEL); Sonson and vie., May, fl., Core 745 (US), Aug., fl., 
Johnson & Barkley 18C846 (MEDEL, US), June, fl., Rivera 315 
(MEDEL), fl., Rivera 316 (MEDEL), March, fl. fr., Scolnik et al. 
19An306 (BM, US), June, fl., Soejarto & Rivera 2043 (ECON, GH, 
MEDEL); no loc., April, fl., Triana 3251 (K); Rio Negro, fl., Trianas. 
n. (F, leaf fragment; G, K, P; COL, US, photos). SANTANDER: E. of 
Bucaramanga, Dec., fl., Molina & Barkley 18S342 (US). GUN- 
DIN AM ARC A: Chiquinquira, July, fl., Felix s. n. (L); road Bogota to 
Honda, Guadua, Aug., fl., Garcia-Barriga 16103 (COL, ECON); 
Bogota, Quebrada del Chico, April, fl., Schneider 156 (S); 
Fusagasuga, July, fl., Soejarto & Schultes 216 (COL, ECON, GH), 
218A (COL, ECON, GH); Junin to Gama, Rio Sucio, Sept., fl., 
Garcia-Barriga 17534 (COL, ECON); Gacheta to Ubala, Laguna 
Verde, July, fl., Garcia-Barriga 17487 (COL, ECON). NO PRECISE 
LOCALITY: Alto de Lasca, May, fl., Goudot s. n. (P; US, photo); 
Aug., fl., Tomas 218 (MEDEL). 

The region of Antioquia appears to be the center of distribution of 
S. ursina, where it is abundantly represented in the mountains sur- 
rounding the city of Medellin. This species is distinguished in the 
field by the following characteristics: (1) dense and reddish maroon 
pubescence on the leaves and shoots; (2) coarse, curved, and long 
trichomes, which are sometimes prickly, along the branchlets and 
petioles; (3) aggregated flowers on the inflorescence; and (4) the low 
number of stamens. 

49. Saurauia bullosa Wawra in von Mart., Fl. Brasil. 12(1): 286. 
pi. 56, fig. 1. 1886. Type: ex descr., ex icon. 

Saurauia pseudoexcelsa Busc., Malpighia 25: 236. 1912, p. p. 
Saurauia spragueana Busc., Malpighia 26: 10. pi. 4, fig. 4. 
1913. Type: Sodiro 154 (A187?) (COL, NY, photos). 



96 FIELDIANA: BOTANY 

Saurauia leoi Busc., Malpighia 30: 121. pi. 3, fig. 3. 1927. Type: 

Karsten s. n. (n. v.), ex descr, ex icon. 
Saurauia hypomalla R. Benoist, Bull. Soc. Bot. France 80: 334. 

1933. Type: Rivet 184 (P). 

Saurauia mojandensis R. Benoist, 1. c. Type: Benoist 4001 (P). 
Saurauia garcia-barrigae R. E. Schultes, Mutisia 3: 1. figs. 1-6. 

1952. Type: Garcia-Barriga 12276 (COL, holotype; F, GH, 

US, isotypes). 
Saurauia roseotincta R. E. Schultes, Bot. Mus. Leafl. 16: 83. 

1953. Type: Macbride 3652 (US, holotype; F, isotype). 

Trees to 10(-20) m. tall, diameter to 20(-40) cm. at base, bark irregularly cracked, 
corky, gray-brown, wood white to light brown, brittle, trunk straight to crooked, 
frequently with branches inserted in a wheel-like pattern, crown open; copiously 
pubescent. Branchlets stout, terete, densely hirsute pubescent or wooly, trichomes 
slender, slightly flexuous, gradually swollen or flattened towards their base, golden- 
yellow to -brown to deep reddish brown, to 12 mm. long. Leaves crowded behind tip of 
branchlets; blades elliptic to elongate-obovate, acute to acuminate at apex with acu- 
men to 15 mm. long, cuneate to obtuse to subcordate and frequently oblique at base, 
serrulate to setaceous-serrulate along margins, 15-30(-38) cm. long, (3-)6-12(-16) cm. 
wide, coriaceous, dark green above, hoary to grayish to rusty-brown beneath, rugose to 
rugulose and scabrous to strongly scabrous above, soft and wooly to touch beneath, 
secondary veins (15-)20-27(-32) pairs, tertiary veins elevated, more prominent than 
lesser venation, abundantly setose to strigose pubescent along and between veins 
above, densely lanate pubescent with trichomes of loriform and flexuous mixed with 
stellate (between veins) and curved paleaceous types (along veins) beneath; petioles 
2-4.5(-6) cm. long, 3-4.5 mm. in diameter, pubescence similar to that along branchlets. 
Inflorescences erect to ascending, 9-40(-60)-flowered, 15-33 cm. long, 7-12 cm. wide, 
pubescence similar in type and density to that along branchlets and petioles, primary 
peduncle 7-17 cm. long, bracts linear to subulate, to 15 mm. long. Flowers (20-)25- 
4CK-50) mm. broad, buds to 15 mm. in diameter, pedicels 3-10(-18) mm. long, bracteoles 
triangular to 5 mm. long; sepals 5, mostly pale green, ovate to oblong to obovate, 
subacute to obtuse, 10-18 mm. long, 6-11 mm. wide, exposed parts in bud densely 
pubescent with trichomes of hirsute to shaggy-strigose mixed with minutely stellate to 
tufted types, imbricated parts densely stellate pubescent, all densely stellate pubes- 
cent inside, marginally ciliate; petals 5, broadly oblong to elliptic, obtuse to incised, 
10-22 mm. long, 8-15 mm. wide; stamens (70-)100-200(-240), filament 3-5 mm. long, 
anther 2-2.5 mm. long; ovary 5(-6)-loculed, globose, 5(-6)-sulcate, glabrous, styles 5(-6), 
obsolete to 7 mm. long, stigmas simple to capitate. Berries green, ovoid, to 20 mm. 
across, 5-sulcate. 

Habitat. Wet mountain forest, cloud forest, subparamo, bushy 
forest, moist Cinchona forest, Podocarpus forest, slopes with secon- 
dary forest, rolling hills, wet ravines and gullies, along stream, 
grassy slopes, cultivated potato fields, gardens, at altitudes of 
(1,500-)2,700-3,600 m. 



SOE JARTO: SA URA VIA 97 

Distribution. Colombia (Departments of Santander, Caldas, 
Quindio, Tolima, Valle, Cauca, Huila, Narino, Putumayo), Ecuador 
(Provinces of Carchi, Imbabura, Pichincha, Santiago-Zamora, Loja), 
Peru (Departments of Lambayeque and Huanuco). 

Vernacular names. Dulumoco (Valle: Cuatrecasas), Moquillo 
(Narino: Soejarto; Carchi: Mexia), Pururuju (Santiago-Zamora: 
Solis). 

Specimens examined. COLOMBIA, SANTANDER: Quebrada de 
Paris, N. of La Baja, Jan., fl., Killip & Smith 18825 (A, GH, NY, 
US). CALDAS: basin of Otun River, above Penas Blancas, Nov., fl., 
Cuatrecasas 23316 (ECON, F); Salento, Rio Boquia, July, fl., Killip 
& Hazen 8845 (GH, NY, US); between Manizales and Nevado del 
Ruiz, June, fl., Soejarto 2032 (ECON, GH). QUINDIO: no. loc., Jan., 
fl., Holton 23 (NY); El Roble, near Cartago, Nov., fl., Holton 24 
( = 792) (K, NY). TOLIMA: El Libano to Murillo (km. 11 to 12), Alto 
de Penones, July, fl., Garcia-Barriga 12276 (COL, F, GH, US). 
VALLE: Cordillera Central, headwaters of Tulua River, Quebrada 
de Las Vegas, March, fl., Cuatrecasas 20414 (ECON, F); basin of 
Bugalagrande River, Cuchilla de Barragan, between Las Azules and 
Las Violetas, April, fl. fr., Cuatrecasas 20805 (ECON, F), fl., 20997 
(ECON, F, US); Cordillera Occidental, Quebrada Las Nieves, below 
El Diamante, July, fl. fr., Cuatrecasas 21804 (ECON, F); basin of 
Amayme River, La Albania, Feb., fr., Cuatrecasas et al. 26809 (US). 
CAUCA: road Popayan to Totoro, Alto de Angosi, 11 km. from Totoro, 
July, fl., Garcia-Barriga 12721 (US); Paletara to Calaguala, June, fl., 
Pennell 7097 (US); Tierra Dentro, Las Escaleretas, Moras Valley 
along Paez River basin, Feb., fl.,Pittier 1371 (US). HUILA: Balsillas 
River, Aug., fl., Rusby & Pennell 713 (NY, US). NARINO: Tuquer- 
res, below El Espino, Aug., fl., Soejarto 1472 (ECON, GH); 
Chachagui, E. of Pasto River, 18 km. N. of Pasto, Oct., fl., Fosberg 
21254 (NY, US); Pasto, volcano El Galeras, fl., Castro 78 (ECON, 
PASTO), July, fl., Soejarto & Hernandez 1015, 1045(both in COL, 
ECON, GH, PASTO); Alto de Tangua, road to Yacuanquer, July, fl., 
Uribe-Uribe 5278, 5328 (both in COL, ECON); Sapues, vie. of El 
Espino, Aug., fl., Soejarto et al. 1334, 1435 (both in ECON, GH); 
Guachucal, road Guachucal to Aldana, Aug., fl., Soejarto et al. 1335 
(ECON, GH, PASTO); road El Encano to Pasto, Paramo del Tabano, 
Jan., fl., Cuatrecasas 11966 (US); Lake La Cocha, Corota island, 
July, fl., Garcia-Barriga etal. 13044 (MEDEL, US); Bosque Botana, 
above Institute Tecnologico Agricola Exptl. Station, April, fl., 
Fajardo 81 (ECON, PASTO), fl. fr. veg., Soejarto & Porter 495, 496, 



98 FIELDIANA: BOTANY 

500 (all in ECON, GH), Soejarto 1473, 1474, 1475, 1478, 1479, 1480, 
1483, 1484, 1491, 1494, 1496, 1504, 1508, 1595, 1596 (all in ECON, 
GH). 

ECUADOR, CARCHI: Tulcan, valley of Pun, Aug., fl., Mexia 7599 
(F, UC, US); 12 km. from Tulcan in the direction of Juliandrade, 
Aug., fr., Soejarto et al. 1336 (ECON, GH, PASTO). IMBABURA: 
Shanshipamba, Nov., fr., Solis 14178 (F). PICHINCHA: Cordillera 
de Pichincha, N. of Quito, Machinqui to Pomasqui, Aug., fl., Hitch- 
cock 20888 (GH, NY, US); Nono, upper reaches of Nanegal River 
(tributary of Guayllabamba River), fl., Sodiro 154 (COL, NY, 
photos). SANTIAGO-ZAMORA: Cordillera Oriental, Campanas to 
the east of El Pan, July, fl., Solis 5043 (F), July, fr.,Steyermark 53542 
(NY). LOJA: Villenaco, Oct., fl., Espinosa E-713 (F, NY); between 
Loja and San Lucas, Sept., fl., Hitchcock 21458 (NY, US); San Lucas 
to Ona, Sept., fr., Hitchcock 21513 (NY, US); vie. of Loja, July, fl., 
Penland & Summers 1136 (F, US), Nov., fl., Andre 4413 (F, K, NY, 
US). NO PRECISE LOCALITY: vie. of Zaragura, Sept., fl., Rose et 
al. 23165 (GH, NY, US); E. of Mojanda, fl., Sodiro s. n. (P). 

PERU, LAMBAYEQUE: Chiclayo, Oct., fl., Rank P-2199 (F), 
HUANUCO: Chaglla, May, fl., Macbride 3652 (F, US); upper region 
of Huallaga River, Pampayacu, July, fl., Sawada 53 (F); Pillao, 
March, fl., Woytkowski 34178 (F, US); between Huanuco and Tingo 
Maria, Chinchao, July, fl. veg., Soejarto & Wild 1421, 1422, 1428, 
1429 (all in ECON, GH). NO PRECISE LOCALITY: fl., Martinet 
1640 (P). 

S. bullosa is abundantly represented in the Narino mountain sys- 
tem at altitudes above 2,700 m. At the volcano El Galeras, above 
Pasto, individuals of this species are found up to 3,600 m., probably 
the highest limit of tolerance of members ofSaurauia. Young leaves 
and shoots are often deep purplish red in color, but they turn some- 
what gray or whitish gray when old. Other distinguishing charac- 
ters of this species are (1) leaves densely lanate pubescent beneath, 
with trichomes of loriform to paleaceous mixed with stellate types; 
(2) flowers large (up to 50 mm. broad); (3) stamens high (over 100) in 
number. The heterotrichous nature of the lower leaf indument, 
however, can only be seen clearly by selective focusing using no less 
than 10 times magnification. 

Although members of this species are often found in disturbed 
habitats, such as secondary forest, abandoned fields, and gardens, 
the original habitat appears to be humid or wet mountain forest, as 



SOEJ ARTO: SA URA VIA 99 

that still found on the Corota Island in the middle of Lake La Cocha, 
in Narino. The center of distribution of the species appears to be 
located in the Colombo-Ecuadorian frontier. 



IMPERFECTLY KNOWN TAXA 

1. Saurauia raimondiana Sleumer, Notizbl. 12: 145. 1934. Type: 
Raimondi 4758 (not seen). 

Saurauia raimondiana var. caxamarcensis Sleumer, 1. c. 146. 
Type: Raimondi 5394 (not seen). 

2. Saurauia rhodosma Sleumer, 1. c. 147. Type: Schultze 1141 
(not seen). 

3. Saurauia trolliana Sleumer, 1. c. 148. Type: Troll 1518 (not 
seen). From the original specific description, it appears that this 
species is closely related to S. rusbyi Britt. 

4. Saurauia schultzeorum Sleumer, Notizbl. 15: 374. 1941. Type: 
Schultze-R hon hof 3058 (not seen). 

ADDENDA 

42. Saurauia excelsa Willd. 

Saurauja xanthotricha Turcz., Bull. Soc. Imper. Naturalistes 

Moscou 31(1): 243. 1858; Knuth, Init. Fl. Venez. 479 Type: 

Funck & Schlim 106 (BM, F, K, P). 
Saurauja pycnotricha Turcz., I.e. 244; Busc., Malpighia 25: 240. 

1912; Knuth, I.e.; Schultes, Caldasia 3: 253. 1943. Type: 

Funck 306 (G, P). 

Saurauja spectabilis auct., non Hook.: Turcz., I.e. 
Saurauja peduncularis auct., non Tr. & PI.: Ernst, Expos. Nac. 

Venez. 1883. 217. 1884; Pittier, PI. Usu. Venez. 300. 1926; 

Knuth, I.e. 



100 








FIG. 3. Saurauia spinuligera R. E. Schultes, Toro 983 (NY), collected in Tamesis of 
Antioquia (Colombia). 



101 








FIG. 4. Saurauia stapfiana Busc., Triana 267 (P), a type specimen, collected in 
Mariquita of Tolima (Colombia). 



102 








FIG. 5. Saurauia caquetensis R. E. Schultes var. caquetensis, Cuatrecasas 8439 (F), 
a type specimen. 



103 




/> 2. 



- ; 



FIG. 6. Saurauia rusbyi Britt., flusftjy 482 (MICH), a type specimen. 



104 




FIG. 7. Saurauia aequatoriensis Sprague, Spruce 4989 (G), a type specimen. 



105 








FIG. 8. Saurauia glabra (R. & P.) Soejarto, Pavon s.n. (G), a lectotype of Palaua 
glabra R. & P. 



106 












Typus! 



Museum botamcum Berolinense. 



FIG. 9. Saurauia adenodonta Sleumer, Schultze-Rhonhof 2914 (K), a lectotype. 



107 



FIELDIANA: BOTANY 




FIG. 10. Saurauia laevigata Tr. & PI., Triana s.n. (G), a lectotype, collected in 
Mariquita of Tolima (Colombia). 



108 



SOEJARTO: SA URA VIA 




FIG. 11. Saurauia strigillosa Tr. & PL, Triana s.n. (G), a lectotype, collected in 
Mariquita of Tolima (Colombia). 



109 





FIG. 12. Saurauia multinervis Soejarto, Idrobo & Fernandez 54 (US), a holotype. 



110 





FIG. 13. Saurauia schultesiana Soejarto, Pennell 10501 (NY), a holotype. 



Ill 




FIG. 14. Saurauia solitaria Sleumer, Weberbauer 7067 (F), a lectotype. 



112 




FIG. 15. Saurauia micayensis Killip, Killip 7932 (US), a lectotype. 



113 




PLANTS Of PEBU 



FIG. 16. Saurauia natalicia Sleumer, Weberbauer 7867 (F), a lectotype. 



114 




FIG. 17. Saurauia pulchra Sprague, Sprague 323 (K), a lectotype. 



115 




FIG. 18. Saurauia floccifera Tr. & PI., Triana s.n. (P), an iso-lectotype. 



116 




c" 



ISOTYPUS 

FIG. 19. Saurauia cuatrecasana R. E. Schultes, Cuatrecasas 9209 (F), an isotype. 



117 







FIG. 20. Saurauia arnoldi Sleumer, Kernan 125 (NY), a neotype, collected in San 
Lorenzo Mountains of Magdalena (Colombia). 



118 




. 

' 

' : 



FIG. 21. Saurauia mexiae Killip ex Soejarto, Mexia 8488 (US), a holotype. 



119 




FIG. 22. Saurauia tomentosa (HBK.) Spreng., Bonpland 3206 (P), a lectotype. 



120 





FIG. 23. Saurauia formosa Sleumer, Weberbauer 6637 (US), an iso-lectotype. 



121 





FIG. 24. Saurauia chaparensis Soejarto, Steinbach 8920 (GH), a holotype. 



122 




FIG. 25. Saurauia peduncularis Tr. & PL, Soejarto 938, collected in an area west of 
Tuquerres of Narino (Colombia), the same general area where the type was collected. 



123 




FIG. 26. Saurauia lehmannii Hieron., Lehmann 6675 (F), an iso-lectotype. 



124 




FIG. 27. Saurauia scabra (HBK.) Dietr., Soejarto & Idrobo 913, collected in the 
Salto of Tequendama area (Cundinamarca, Colombia). 



125 





FIG. 28. Saurauia tambensis Killip, Hitchcock 21281 (US), a holotype. 



126 




FIG. 29. Saurauia excelsa Willd., Bredemeyer s.n. (type specimen), formerly de- 
posited at the Berlin Herbarium. 



127 




"!3- 

\OM. 



FIG. 30. Saurauia brachybotrys Turcz., Linden 972 (K), a type specimen. 



128 





' 



IM.AMS 01 VIM /I ! 1 i 



jhn> 10 r-t uili IWMII ii uui - 
ror mUrtaUow !-]! ior . ill*: 
!** . iralj ! infco. Jail &!! 



>>osso 1 1 1 1 1 ii ii ri 

FIG. 31. Saurauia meridensis Steyermark, Steyermark 56344 (F), a holotype. 



129 




FIG. 32. Saurauia ursina Tr. & PL, Triana s.n. (P), an iso-lectotype, collected in Rio 
Negro of Antioquia (Colombia). 



130 



APPENDIX 

LIST OF ACCEPTED SERIES AND SPECIES 
OF SOUTH AMERICAN SAURAUIA 

Arabic numerals in front of the Latin binomial refer to the species 
number used in the present revision. 



L Ser. Omichlophilae Soejarto 

1. S. spinuligera R. E. Schultes 

2. S. stapfiana Busc. 

3. S. omichlophila R. E. Schultes 

4. S. caquetensis R. E. Schultes 

II. Ser. Gynotrichae Busc. 

5. S. loeseneriana Busc. 

III. Ser. Laevigatae Busc. 

6. S. briqueti Busc. 

7. S. portachuelensis R. E. Schultes 

8. S. rusbyi Britt. 

9. S. glabra (R. & P.) Soejarto 

10. S. yasicae Loes. 

11. S. aequatoriensis Sprague 

12. S. adenodonta Sleumer 

13. S. strigillosa Tr. & PI. 

14. S. laevigata Tr. & PL 

IV. Ser. Parviflorae Soejarto 

15. S. multinervis Soejarto 

16. S. schultesiana Soejarto 

17. S. peruviana Busc. 

18. S. chiliantha R. E. Schultes 

19. S. solitaria Sleumer 

20. S. micayensis Killip 

21. S. natalicia Sleumer 

22. S. parviflora Tr. & PL 

23. S. pseudoleucocarpa Busc. 

24. S. spectabilis Hook. 



V. Ser. Pulverulentae Busc. 

25. S. pulchra Sprague 

26. S. aromatica R. E. Schultes 

27. S. floccifera Tr. & PL 

28. S. choriophylla R. E. Schultes 

29. S. cuatrecasana R. E. Schultes 

30. S. arnoldi Sleumer 

31. S. tomentosa (HBK.) Spreng. 

32. S. pseudostrigillosa Busc. 

33. S. mexiae Killip ex Soejarto 

VI. Ser. Macrophyllae Busc. 

34. S. formosa Sleumer 

35. S. chaparensis Soejarto 

36. S. peduncularis Tr. & PL 

37. S. putumayonis R. E. Schultes 

38. S. herthae Sleumer 

39. S. lehmannii Hieron. 

40. S. scabra (HBK.) Dietr. 

41. S. tambensis Killip 

42. S. excelsa Willd. 

43. S. brachybotrys Turcz. 

44. S. isoxanthotricha Busc. 

45. S. meridensis Steyerm. 

46. S. prainiana Busc. 

VII. Ser. Lanatae Soejarto 

47. S. biserrata (R. & P.) Spreng. 

48. S. ursina Tr. & PL 

49. S. bullosa Wawra 



131 



REFERENCES 

ACOSTA-SOLIS, M. 

1960. Maderas economicas del Ecuador y sus usos. Editorial Casa de la Cultura 
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BAILLON, H. E. 

1873. Histoire des plantes, vol. 4, p. 258. 

BENOIST, R. 

1933. Descriptions d'especies nouvelles de phanerogames sud-americaines. Bull. 
Soc. Bot. France, 80, pp. 333-335. 

BENTHAM, G. 

1861. Notes on Ternstroemiaceae. J. Proc. Linn. Soc., Bot., 5, pp. 53-61. 

BENTHAM, G. and J. D. HOOKER 

1867. Genera plantarum, vol. 1, p. 184. 
BRITTON, N. L. 

1889. An enumeration of plants collected by Dr. H. H. Rusby in South America, IV. 
Bull. Torrey Bot. Club, 16, p. 64. 

BROWN, E. G. S. 

1935. The floral mechanism ofSaurauia subspinosa Anth. Trans. & Proc. Bot. Soc. 

Edinburgh, 31, pp. 485-497. 
BUSCALIONI, L. and E. G. MUSCATELLO 

1912-1927. Studio monografico sulle specie americane del genero "Saurauia" Willd. 
Malpighia, 24, pp. 281-412; 25, pp. 1-16, 106-118, 187-250, 389-436; 26, pp. 1-32, 
97-144, 281-312, 389-420; 27, pp. 1-32, 131-158, 293-324, 487-502; 28, pp. 1-48, 
107-138, 223-238, 315-330, 371-402, 473-488; 29, pp. 1-32, 97-112, 230-245, 319- 
366, 411-458; 30, pp. 35-444. 

1927. Studio monografico sulle specie americane del genero "Saurauia" Willd. 
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INDEX TO LATIN NAMES 

New names and combinations are in boldface type, synonyms in 
italics, and all other names in regular Roman type. Names with (*) 
are mentioned incidentally, with (**) imperfectly known taxa, and 
with (t) fossil representatives. Page numbers in boldface type indi- 
cate descriptions, in italics plates/illustrations, in parentheses 
taxonomic keys, and in regular type all others. 

Macrophyllae (Ser.) 6, (20), 68 
Marcgravia* 17 
Marumia 19 
Mesophyllae 68 
Obelanthera 19 
Oligotrichae 29 

Omichlophilae (Ser.) 6, (20), 20 
Overstratia 19 
Palaua 19 

biserrata 43, 90 

glabra 32, 33, 106 

glabrata 43 

hirsuta 91 

lanceolate. 19, 90 
Palava 

biserrata 90 

scabra 76 

tomentosa 63, 64 
Parietales 15 

Parviflorae (Ser.) 6, (20), 40 
Pulverulentae (Ser.) 6, (20), 54 
Polyandria Pentagynia* 14 
Reinwardtia 19 
Ruitzianae 90 
Saurauia (see also Saurauja 

and Sauravia) 1, 3, 4, 8, 9, 13, 
14, 19 

adenodonta 19, (29), 37, 107 

aequatoriensis (29), 36, 105 
var. boliviana 32, 53 



Actinidia 14 

crassispermat 12 
Actinidiaceae 14 
Actinidioideae 15 
Apatelia 19 

biserrata 90, 91 

glabrata 32 

lanceolata 91 

var. peduncularis 91 
Blumia 19 

Brachitrichae 20, 27, 40, 68 
Clematoclethra 15 
Clematoclethroideae 15 
Clethra 15 
Clethraceae 15 
Davya 19 
Dilleniaceae 14 
Dilleniales 15, 17 
Draytonia 19 
Ericales 16, 17 
Ficus alkalinat 12 
Guttiferales 15 
Gynogynae 20, 40, 69 
Gynotrichae (Ser.) 8, (20), 27 
Hickoria* 12 
Hyper icum* 17 
Juglans deformist 12 
Laevigatae (Ser.) 6, (20), 28, 54 
Lanatae (Ser.) 6, (20), 90 
Leucothea 19 



139 



140 



FIELDIANA: BOTANY 



var. glabrata 32, 36 

fina. longepetiolata 36 

fma. veranii 36 
alaskanat 11, 12 
alvaroi 56, 62 
anolaimensis 85 
arnoldi 12, (55), 62, 118 
aromatica 5, (55), 56 

aspera* 11 
belizensis 34 
biserrata (90), 90 

brachybotrys 11, 13, 58, (69), 82, 128 
var. macrantha 80, 82 
var. scabra 82 
brevipes 45, 53 
briquet! (29), 29 
bullosa 8, 13, 42, 66, (90), 95 
caquetensis (21), 26, 58, 703 
var. caquetensis (27), 103 
var. parviflora (27), 27 
chaparensis (69), 71, 122 
chiliantha (41), 46 
choriophylla (55), 60 
consimilis 89 
coroicoana 43 

cuatrecasana (55), 58, 61, 85, 117 
deformist 12 
echinosepala 56 

excelsa 12, 19, 42, 68, (69), 79, 80, 127 
var. moritziana 80 
var. xanthotricha 80 

fma. veranii 80 
floccifera (155), 58, 61, 116 
floribunda 38, 43, 67, 68 
var. laevigata 38 
var. peruviana 43 
formosa (69), 70, 89, 121 
garcia-barrigae 96 
glabra (29), 32, 50, 106 
glabrata 32 
goudotiana 82 
herbert-smithii 33 
herthae (69), 74 
humboldtiana 76 

var. bonplandiana 76 
hypomalla 96 
intonsa 85 

isoxanthotricha (70), 85, 88 
kallima 46 



laevigata 10, 12, (29), 36, 38, 39, 108 

lehmannii (69), 75, 124 

leoi 96 

leucocarpa var. smithiana 33 

loeseneriana 8, 27 

meridensis (70), 87, 129 

mexiaeS, (55), 68, 119 

micans 52 

micayensis 12, (41), 48, 113 

mojandensis 96 

multinervis 12, (40), 41, 110 

narcissifragrans 11 

natalicia (41), 50, 114 

omichlophila 6, (21), 24, 42 

parviflora 8, (41), 49, 50 

var. lehmanniana 51 
pastasana 89 
peduncularis 6, 52, (69), 72, 123 

var. veraniana 82 
peruviana (40), 43, 44, 45, 79 
portachuelensis (29), 30 
prainiana (70), 75, 89 

var. humboldtiana 11 
pruinosa 64 
pseudoexcelsa 64, 95 
pseudoleucocarpa 8, (41), 49, 52 
pseudoparviflora 43, 45, 50 

var. rusbyana 50 
pseudopittieri 34 
pseudoruitziana 91 
pseudoscabra 43 

pseudostrigillosa (55), 58, 67 

pulchra 12, (54), 55, 115 

putumayonis 50, (69), 73, 87 

pycnotricha 100 

pyramidata 43, 46 

raimondiana** 100 

var. caxamarcensis** 100 

rhamnifolia 36 

rhodosma** 100 

rigidissima 48 

roboranst 12 

roseotincta 96 

ruitziana 93 

ruiziana var. tomentosa 64 

ruiziana var. weberbaueri 91 

rusby 53 

rusbyi (29), 31, 53, 104 
var. glabrata 45, 53 



SOEJARTO: SAURAUIA 



141 



var. spectabilis 45, 53, 80 
fma. macrophylla 45, 53 
fma. veranii 53 
scabra 43, 44, (69), 76, 85, 125 

var. boliviano 43 

var. prainiana 91 
ftna. veranii 80 
scabriuscula 43, 44 
schultesiana 10, (40), 42, 111 
schultzeorum** 100 
serrata* 33 
smithiana 34 
solitaria (41), 48, 112 
spectabilis 32, (41), 53 
spinuligera 10, (21), 21, 42, 101 
spragueana 95 
sprucei 64 
stapfiana (21), 22, 102 

var. radiata (23), 24 

var. stapfiana 23 
strigillosa 10, (29), 38, 109 

var. microphylla 38 
sydowii 89 

tambensis (69), 79, 126 
tomentosa 13, (55), 58, 59, 63, 85, 120 

var. chillanea 64 
tristyla* 12 
trolliana** 100 
ursina 85, (90), 93, 130 

ftna. strigosa 93 

frna. veranii 93 
weberbaueri 43 
xanthotricha 100 



yasicae 7, 8, 11, 12, (29), 33, 40 
var. laevigatae 34 
fma. veranii 34 

zetekiana 34 
Saurauiaceae 14 
Saurauioideae 15 
Saurauja 3, 19 

floribunda 80 

glabrata 32 

lanceolata 91 

moritziana 80 

sca&ra 43, 44, 76 

schlimii 80 
Saurauia 

glabrata 32 

ruiziana 91 

serrate 32 
Scapha 19 
Scabrae 29 
Schizocardia* 17 
Sladenia 15 
Stenobasicae 20, 40, 69 
Strigosae 68, 90 
Ternstroemiaceae 14 
Theaceae 14 
Theales 15, 17 
Theineae 15 
Tiliaceae* 14 
Tomentosae 54 
Tonshia 19 
Trematanthera 19 
Vanalphimia 19 
68 




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