SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 69 

A Revision of the Genus Olyra 

and the New Segregate Genus Parodiolyra 

(Poaceae: Bambusoideae: Olyreae) 

Thomas R. Soderstrom 

and 

Fernando 0. Zuloaga 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1989 
ABSTRACT 

Soderstrom, Thomas R., and Fernando 0. Zuloaga. A Revision of the Genus Olyra and the 

New Segregate Genus Parodiolyra (Poaceae: Bambusoideae: Olyreae). Smithsonian Contributions to Botany, number 69, 79 pages, 46 figures, 2 appendices, 1989.-The herbaceous 

bambusoid genus Olyra is revised. Four new South American species are described Olyra 

amapana, 0. latispicula, 0. retrorsa, and 0. tamanquareana. Three species of Olyra were 

found to differ significantly from the rest of the genus in possessing the following combination 

of characters: weak, scandent culms; filiform female spikelet pedicels; female spikelets that 

fall entire, have a pulvinate internode between the inflated glumes, and a stipitate anthecium; 

and a hilum that does not extend for the entire length of the caryopsis. The segregate genus 

Parodiolyra is established to accomodate these species, and the new combinations P. lateralis, 

P. luetzelburgii, and R ramosissima are proposed. The new genus may share affinities with 

Raddiella and Diandrolyra rather than Olyra. 

Keys to the species in both genera are presented, as well as complete descriptions, field 

photographs, and distribution maps. Scanning electron microscope studies of the surface of the 

female anthecia yielded characters of taxonomic significance. These are illustrated and 

incorporated in the key. Two appendices account for all names that have been published in the 

genus Olyra. 

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

recorded in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf 

clearing from the katsura tree Cercidiphyllum japonicum Siebold & Zuccarini. 

Library of Congress Cataloging-in-Publication Data 

Sodastrom, Thomas R. 

A revision of the genus Olyra and the new segregate genus Parodiolyra (Poaceae: Bambusoideae: Olyreae). 

(Smithsonian contributions to botanv : no. 69) <. 

Bibliography: p. 

1. Ofyra - Classification. 2. Parodiolyra - Classification. 3. Grasses-Classification. 

I. Zuioaga, Fernando 0. 11. Title. 1111 Series. 

QKlS2747 no. 69 [QK495.G74] 581 s-dcl9 [584’.93] 88-22646 CIP 
Contents 

Page 

Preface .......................................... iv 

Introduction ........................................ 1 

Methods and Materials ................................. 1 

Acknowledgments ................................... 2 

Olyra Linnaeus ....................................... 2 

Key to the Species of Olyra .............................. 3 

1 . Olyra amapana Soderstrom & Zuloaga. new species ............... 5 

2 . Olyra buchtienii Hackel .............................. 8 

3 . Olyra caudata Trinius ............................... 10 

4 . Olyra ciliatifolia Raddi .............................. 12 

5 . Olyra ecaudata Doell ............................... 14 

6 . Olyra fasciculata Trinius ............................. 17 

7 . Olyra filiformis Trinius .............................. 19 

8 . Olyra glaberrima Raddi .............................. 20 

9 . Olyra holttumiana Soderstrom & Zuloaga .................... 24 

10 . Olyra hum’lis Nees von Esenbeck ........................ 24 

11 . Olyra juruana Mez ................................ 26 

12 . Olyra latifolia Linnaeus .............................. 27 

13 . Olyra latispicula Soderstrom & Zuloaga. new species .............. 35 

14 . Olyra longifolia Humboldt. Bonpland & Kunth ................. 35 

16 . Olyra maranonensis Swallen ........................... 46 

17 . Olyra micrantha Humboldt. Bonpland & Kunth ................. 46 

18 . Olyra obliquifolia Steudel ............................. 52 

19 . Olyra retrorsa Soderstrorn & Zuloaga. new species ............... 54 

20 . Olyra standleyi Hitchcock ............................. 57 

21 . Olyra tamanquareana Soderstrom & Zuloaga. new species ........... 58 

22 . Olyra taquara Swallen .............................. 61 

23 . Olyra wurdackii Swallen ............................. 63 

15 . Olyra loretensis Mez ............................... 43 

Parodiolyra Soderstrom & Zuloaga. new genus .................... 64 

Key to the Species of Parodiolyra .......................... 65 

Zuloaga. new combination ............................. 66 

2 . Parodiolyra luetzelburgii (Pilger) Soderstrom & Zuloaga. new combination ... ........................................... 70 

3 . Parodiolyra ramosissima (Trinius) Soderstrom & Zuloaga. new combination ... ............................................ 73 

Appendix 1: Names Included in Olyra Linnaeus .................... 76 

Appendix 2: Names Excluded from Olyra Linnaeus .................. 77 

Literature Cited ...................................... 78 

1 . Parodiolyra lateralis @Yes1 ex Nees von Esenbeck) Soderstrom & 

... 

111 
Preface 

My first experience with Ulyra came in 1962 on an expedition to Kaieteur Falls in the 

interior of Guyana (then British Guiana) in northern South America. Large clumps of 0. latifolia 

were scattered here and there, mostly at the edge of the forest or in clearings where trees had 

fallen. The unusual grass Pariana, with yellow spike-like inflorescences and spikelets with 

multiple stamens, was also found here. Both genera looked like bamboos in the field, but when 

I returned to Washington I found Olyra filed in the herbarium in the Paniceae and Pariana in 

the Hordeae because of the spike-like inflorescence. I would now include Ulyra and several 

other genera, including Pariana, in the Olyreae (Soderstrom and Ellis, 1987). 

On a collection trip to Suriname in 1963, I was impressed by the growth habit of Ulyra 

luetzelburgii, a clambering plant that grew abundantly at the forest edges abutting a granite 

outcrop. The plants appeared to be related to Ulyra but differed from it. Later, as part of 

continuing studies on the different genera of the Olyreae, it became evident that this species 

and two others that had been included in Olyra, 0. lateralis and 0. ramosissima, formed a 

natural group that deserved recognition as a separate genus. Here, we name that genus 

Parodiolyra in honor of the Argentine agrostologist Lorenzo R. Parodi (1895-1966). 

Several species of Olyra have been transferred to other genera (Arberella, Cryprochloa, 

Piresia, and Sucrea). With the transfer of three more species to the new genus Parodiolyra, 

and with the new species herein described, this leaves twenty-three species in Olyra. 

The visit of Dr. Fernando 0. Zuloaga of the Instituto Danvinion in San Isidro, Argentina, 

to the Smithsonian in 1982-1983, was a fortuitous event. Dr. Zuloaga came to Washington to 

continue his studies of New World Panicum. As part of his investigation he made extensive 

scanning electron microscope (SEM) studies of the surface of the anthecia of all species. He 

found significant differences in the surface texture and ornamentation, which were most useful 

in the taxonomy of the genus. An early key to Olyra that Cleofe E. Calderon and I had prepared 

was based in part on characters of the surface of the female anthecia, but only of those that 

could be seen with the light microscope. Because of the similar type of anthecium in Ulyra and 

Panicum, Zuloaga decided to run SEM studies on all species of Olyra. These proved to be quite 

useful taxonomically, and he extended these studies to the anthecia of other taxa in the tribe 

Olyreae, including Parodiolyra. The present revision is based on classical and SEM studies. 

Characters from both approaches are incorporated in the keys. 

T.R. Soderstrom 

22 April 1987 

iv 
A Revision of the Genus Olyra 

and the New Segregate Genus Parodiolyra 

(Poaceae: Bambusoideae: Olyreae) 

Thomas R. Soderstrom 

and Fernando 0. Zuloaga 

Introduction 

The genus Olyra is found throughout the Neotropics. The 

most widespread species is also the type species, 0. latifolia, 

based on a specimen from Jamaica. Kunth (1815), in his system 

of grass classification, regarded the genus as a distinct group 

with a relationship to the bamboos, but he was later influenced 

by Trinius (1826) who aligned the genus with Panicurn. The 

reasoning of Trinius has persisted into this century, due 

primarily to the uncanny similarity of the female spikelets of 

the two genera; in both, the female anthecium is usually white 

and indurate. In fact, while the female spikelet of Olyra has 

only one floret and two glumes, is monoecious, and has three 

lodicules, in Panicurn two florets are present, with the lower 

usually male and the upper hermaphroditic, there are only two 

lodicules, and the spikelets are bisexual. 

Following Kunth, one of the first to recognize the bambusoid 

affinities of Olyra was Sir Dietrich Brandis, who made an 

elegant anatomical study (1907) of woody bamboo leaves 

including broad-leaved herbaceous grasses such as Olyra, 

Diandrolyra, and Pharus. Later workers firmly established the 

bambusoid affinities of Olyra, especially on the basis of leaf 

anatomy characters. Other characters, such as seedling type, 

have also been useful in grass classification. The bambusoid 

type of seedling as defined by Soderstrom (1981:26) is shared 

by members of the Olyreae and Bambuseae. Along with the 

Thomas R. Soderstrom (deceased), Department of Botany, National 

Musewn of Natural History, Smithsonian Institution, Washington, 

D.C. 20560. Fernando 0. Zuloaga, Instituto de Botainica Daminion, 

Casilla de Correo 22, San Isidro, Argentina. Miembro de la Carrera 

del Investigador Cientijico del Consejo Nacional de Investigaciones 

Cientificas y Tecnicas (CONICET) a2 la Republica Argentina. 

Review Chairman: Mark M. Littler, Department of Botany, Smithsonian Institution. Reviewers: Gerrit Daviake, Missouri Botanical 

Garden; Lynn G. Clark, Iowa State University. 

woody bamboos (Bambuseae), four herbaceous tribes (Buergersiochloeae of New Guinea, and Anomochloeae, Olyreae, 

and Streptochaeteae of the New World) constitute the “core 

group” of the subfamily Bambusoideae as recognized by 

Soderstrom and Ellis (1987). 

The woody bamboos differ from the herbaceous in their 

long-lived woody culms, generally strong rhizomatous nature 

and complex branching, and propensity to flower only at long 

intervals. Olyra and other herbaceous bamboos usually flowcr 

each year for longer or shorter periods of time, but, 

interestingly, one group of related species (0. taquara, 0. 

ecaudata, and 0. standleyi) apparently do not flower every 

year, and may be monocarpic, as are many woody bamboos. 

The phenomenon is like that of the woody bamboos and is 

worth studying in these herbaceous species since it may offcr 

clues as to the origin of this unique phenomenon in the grasses. 

METHODS AND MATERIALS.-Procedures of classical alpha 

taxonomy were used in this study. Field methods included 

photographing the live plants, collecting and drying them for 

herbarium specimens, preserving leaves, spikelets, and other 

structures in FAA (formalin-ethanol-acetic acid) and transferring them to 70% ethanol in the laboratory. 

Laboratory studies were made using a Wild M5 dissecting 

microscope and Leitz Ortholux compound microscope. For the 

scanning electron microscope (SEM) studies anthecia were 

removed from herbarium specimens, secured on stubs, carbon 

coated in a vacuum evaporator, and coated with a goldpalladium alloy. These specimens were then viewed in a 

Cambridge S4-10 or Cambridge Stereoscan 250 Mk 2 scanning 

electron microscope operating at 10-20 kV. 

In addition to the large collections of material at US, we 

consulted specimens lent by the following herbaria: CEPEC, 

CR, F, GH, IAN, MO, NY, R, RB, U, UB, and WIS. The 

abbreviations of these herbaria, as well as others mentioned in 

1 
2 SLMITHSONIANCONTRIBUTIONS TO BOTANY 

the text, follow Holmgren et al. (1981). Collection localities 

listed within each species discussion are quoted or paraphrased 

from specimen labels. Metric equivalents of English distances 

are provided in brackets. 

ACKNoWLEDGMENTs.-Much of the early work on this 

revision was undertaken in collaboration with a student of the 

late Prof. Lorenzo R. Parodi (Facultad de Agronomia y 

Veterinaria, Universidad de Buenos Aires), Cleofe E. Calderon 

(resident of Washington, D.C.), who came to the Smithsonian 

from Argentina in 1964, working with Soderstrom on the 

Olyreae. Calderon subsequently made a number of field trips 

to tropical America and collected many species and specimens 

of Olyra and other bamboos. She is responsible for most of the 

field photographs in this paper. Much of the specimen material 

was collected by Calderon and Soderstrom on field trips 

sponsored by the Smithsonian Institution and National 

Geographic Society. 

Zuloaga is grateful to the Consejo Nacional de Investigaciones Cientificas y Tecnicas de la Republiblica Argentina 

(CONICET) for a grant to spend two years (1982 and 1983) 

at the Smithsonian Institution. Final studies were made in July 

and August 1986 under a Smithsonian Institution short-term 

visitor’s grant. 

The illustrations of Olyra loretensis, 0. micrantha, and 

Parodiolyra lateralis (Figures 24, 26, and 39) were made by 

Gesina Berendina Threlkeld, formerly National Museum of 

Natural History (NMNH), now resident of Los Alamos, New 

Mexico. Those of P. luetzelburgii and P. ramossisima (Figures 

41 and 45) were made by Vladimiro Dudas, Instituto de 

Botnica Darwinion. The others were rendered by Alice R. 

Tangerini, NMNH. Most of the field photographs were taken 

by Calderon. 

We also thank Mary Sangrey (NMNH) and Jon Ricketson 

(Missouri Botanical Garden) for valuable assistance, and Gerrit 

Davidse (Missouri Botanical Garden), Lynn G. Clark (Iowa 

State University), John Wurdack (NMNK), and Emmet J. 

Judziewicz (NMNH) for helpful advice and suggestions, the 

latter in particular for his assistance in preparing the manuscript 

for publication. 

Olyra Linnaeus 

Olyra Linnaeus, 1759:1261. 

Mapira Adanson, 1763:2, 39, 574. 

Cespitose, monoecious perennials, usually from creeping 

rootstocks. Culms erect and sometimes climbing, leaning on 

the vegetation, unbranched at the lower and median nodes, 

branching at the upper nodes to produce inflorescences; 

internodes cylindrical, glabrous to pilose, the nodes thickened 

to compressed, glabrous or pilose. Leaves with ligules small 

to large, membranous, shortly ciliate at the apex: auricles 

present or absent; pseudopetiole short, densely pilose to nearly 

glabrous; blades flat, ovate-lanceolate to lanceolate, asymmetric to symmetric basally, glabrous or pilose, the margins ciliate 

to scabrous. Inflorescence with one to many panicles borne at 

the uppermost nodes, the panicles lax and diffuse to contracted, 

few- to many-flowered, with the lower branches whorled or 

alternate, the upper ones alternate, in a few cases the panicles 

composed of 2-8 conjugate branches; lower branches of the 

panicle with male spikelets and with or without terminal female 

spikelets, the upper ones with male spikelets below and 1-8 

terminal female spikelets; pedicels of the female spikelets 

thickened, the spikelets disarticulating above the glumes (with 

the exception of 0. micrantha). Female spikelets with 2 aristate 

to acuminate glumes, these subequal or unequal (in this case 

with the lower glume longer than the upper), 5-15-nerved, 

usually with transverse veinlets, pilose to glabrous; anthecium 

indurate, the lemma covering the palea with its borders, shorter 

than the glumes, pilose, with variable types of pubescence, to 

glabrous, smooth or pitted; lemma 5-1 I-nerved, whitish to 

olivaceous, shining; palea 2-4-nerved; lodicules 3, conduplicate; ovary glabrous, smooth: stigmas 2, plumose; caryopsis 

brownish; hilum linear, as long as the caryopsis, the embryo ]/5 

or less the length of the caryopsis. Male spikelets aristate to 

acuminate, shorter than the female spikelets (longer in 0. 

wurduckii), glabrous to hispid, membranous; glumes absent: 

lemma 3-9-nerved; palea 2-nerved (4-nerved in 0. holttumiana): stamens 3. 

TYPE SPECIES.-Olyra latifolia Linnaeus. 

DIsTRIBUTI0N.-Twenty-thrfx species, found throughout 

the Neotropics from Mexico and Florida, United States, south 

to Argentina; apparently introduced in Africa (including 

Madagascar) and Fiji. The species typically occur in lowland, 

wet, tropical forests. Exceptions are noted within species 

discussions and in the key, below. 

DIscL!ssIoN.-We consider two names in Olyra to be of 

uncertain application, and do not treat them further herein. 

First, Olyra guineensis Steudel, 1853:37, is an African taxon 

described from the island of Principe in the Gulf of Guinea. 

We have not seen the type, the current location of which is 

unknown, Second, we could not locate the type of Olyra 

speciosa Mez, 1921:7, thought to be in the Berlin Herbarium. 

We could not confidently make an identification from Mez’s 

description. 
NUMBER 69 3 

Key to the Species of Olyra 

1. Female anthecium pitted or granular over part or all of its surface with conspicuous, 

round to oblong excavations ........................... 2 

2. Inflorescence conjugate, male and female spikelets on separate branches; female 

anthecium with rounded excavations and long, stiff hairs toward the apex, 

obovoid, l/4 the length of the spikelet ............ 14. 0. longifolia 

(northern South America) 

2. Inflorescence not conjugate, male and female spikelets together; female 

anthecium never with both rounded excavations and long hairs on its surface 

(occasionally with bicellular microhairs toward the upper surfaces of the 

lemma and palea, or unicellular retrorse hairs on the palea, as in 0. retrorsu), 

ovoid to fusiform, greater than l/4 the length of the spikelet ......... 3 

3. Female spikelets disarticulating below the glumes, the pedicels thin, not 

thickened; lower and upper glumes separated by an internode; female 

anthecium 2.5-3.8 mm long, entirely covered by rounded excavations . . .............................. .17.0. micrantha 

(South America) 

3. Female spikelets disarticulating above the glumes, the pedicels thickened; 

lower and upper glumes not separated by an internode; female anthecium 

at least 6.8 mm long, granular or partly to entirely covered by rounded or 

oblong excavations ............................ .4 

4. Female anthecium with deep, rounded excavations; leaf blades lanceolate, 

11-14 cm long ...................... 7. 0.filiformis 

(Bahia, Brazil) 

4. Female anthecium with shallow, slightly oblong to round excavations; leaf 

blades oblong to lanceolate, 13-35 cm long (10.5-13.2 cm in 0. 

retrorsu), the female anthecium with a unique combination of slightly 

oblong excavations and retrorse hairs on the apex of the palea .... 5 

5. Female anthecium stipitate, the stipe conspicuous, -1 mm long; leaf 

blades asymmetric basally ...................... 6 

6. Inflorescences numerous at the uppermost nodes, contracted, each with 

a terminal female spikelet and male spikelets below; female 

spikelets greater than 33 mm long ................. 

(Amazonas, Brazil) 

6. Inflorescences 2 or 3 at the uppermost nodes, umbelliform or 

paniculiform, open, with 1-5 terminal female spikelets; female 

7. Inflorescences paniculiform, the lower branches whorled, the upper 

ones alternate; female anthecium with oblong excavations; 

female spikelets 22-33 mm long, 2.9-3.2 mm wide; male 

spikelets glabrous ............... 6. 0. fasciculata 

(Panama, Peru, Bolivia, Argentina, and southern Brazil) 

7. Inflorescences umbelliform, with 2-8 conjugate branches; female 

anthecium with rounded to hexagonal, not oblong, excavations; 

female spikelets 13-17 mm long, 3.4-4 mm wide; male 

spikelets scabrous to densely hirsute .... 18. 0. obliquifolia 

(northern South America) 

5. Female anthecium not stipitate; leaf blades symmetric basally ..... 8 

8. Palea of female anthecium with conspicuous retrorse hairs toward the 

apex; lemma with bicellular microhairs toward the apex; panicles 

6-10 cm long, 1-2 cm wide; leaf blades 10.5-13.2 cm long, 

............... .21. 0. tumanquareana, new species 

spikelets 13-33 mm long. ................... .7 
4 SMITHSONAN CONTRIBUTIOXS TO BOTANY 

2.7-3.4 cm wide; ligules conspicuous, 2.5-4 mm long ...... 

(Mato Grosso, Brazil) 

8. Palea of the female anthecium without retrorse hairs; lemma without 

bicellular microhairs; panicles 9-23 cm long, up to 30 cm wide; 

leaf blades 15-35 cm long, 4-8 cm wide; ligules small, 0.6-1.4 

9. Female spikelets ovate, acuminate to shortly aristate, 8-11.6 mm 

long, 3-3.8 mm wide, the anthecium 70-80% as long as the 

spikelet, ovate, with prickle hairs on the margins of the lemma 

......................... .5.0. ecaudata 

(Costa Rica to the Guianas, Peru, Bolivia, and Brazil) 

9. Female spikelets lanceolate, attenuate to aristate, 16-25 mm long, 

the anthecium 8.4-9.5 mm long, 2.2-3 mm wide, at most 50% 

as long as the spikelet, lanceolate, without prickle hairs on the 

margins of the lemma. ................... 10 

10. Female anthecium with numerous bicellular microhairs at the tip 

of the palea; spikelets without purple spots ......... 

(cloud forests, Costa Rica to Venezuela) 

.................... 19. 0. retrorsa, new species 

mm long. ............................ .9 

....................... .20.0. standleyi 

....................... .22. 0. taquara 

10. Female anthecium glabrous; spikelets often with purple spots . 

(gallery forests, central Brazil) 

1. Female anthecium glabrous or pilose, not pitted or granular .......... 11 

11. Female anthecium glabrous (occasionally with a few bicellular microhairs at the 

tip of the palea or sparse prickle hairs at the margins of the lemma); ligules 

membranous, conspicuous, up to 10 mm long ............... 12 

12. Male spikelets (8.9-11.5 mm long) always longer than the female spikelets 

(7-7.5 mm); leaf blades symmetric basally, lanceolate; female anthecium 

a little shorter than the glumes, the palea with abundant bicellular microhairs 

(Amazonian Venezuela and Brazil) 

12. Male spikelets (5-11 mm long) always shorter than the female spikelets 

(14-50 mm); leaf blades asymmetric basally, ovate-lanceolate to lanceolate; 

female anthecium half as long as the glumes, the palea glabrous .... 13 

13. Female anthecium ovate ......................... 14 

14. Female spikelets 30-48 mm long, the glumes subequal, awns 20-30 

mm long; female anthecium 8.2-10 mm long, -l/4 the length of the 

(Costa Rica to Bolivia) 

14. Female spikelets 14-25 mm long, the glumes unequal, awns less than 

20 mm long, usually 10 mm or less; female anthecium 5.6-6.1 mm 

(Florida to Argentina; introduced into the Old World tropics) 

15. Male lemma 7-9-nerved, the palea 4-nerved; male spikelet 10.2-10.9 

mm long, 0.8-1.2 mm wide, purplish, scabrous; female anthecium 

14.5-15.8 mm long; inflorescences with the lower branches in a 

(Panama) 

15. Male lemma 3-nerved, the palea 2-nerved; male spikelet 5 mm long, 1 

mm wide, acute, brownish, hirsute; female anthecium -10.7 mm long; 

inflorescence formed from a fascicle of terminal branches ...... ............................ 2. 0. buchtienii 

and prickle hairs ...................... 23. 0. wurdackii 

spikelet. ........................ 3. 0. caudata 

long, -l/z the length of the spikelet. ......... 12. 0. latifolk 

13. Female anthecium elliptical ........................ 15 

verticil, the upper alternating ........... 9. 0. holttumiana 

(Mapiri region, Bolivia) 
NUMBER 69 5 

11. Female anthecium pilose, with cylindrical, flattened, or prickle hairs over the 

entire surface or at the base and apex of the anthecium; ligules various . . 16 

16. Female anthecium broadly ovate, the lemma gibbous, with conspicuous and 

caducous prickle hairs over the entire surface ................ 

(Bahia, Brazil) 

16. Female anthecium elliptical, the lemma not gibbous, with cylindrical or 

flattened hairs but without prickle hairs over the entire surface ..... 17 

17. Inflorescence on reduced-leafy culms, showy, with white female spikelets 

terminating the branches ................ 15. 0. loretensis 

(Amazonian Peru and Brazil) 

17. Inflorescence on leafy culms, not showy, female spikelets various ... 18 

18. Female anthecium with long, appressed, cylindrical macrohairs covering 

the entire surface. ......................... 19 

19. Female glumes unequal, the lower glume long-aristate with an awn 

10-12 mm long; female anthecium acuminate ........... 

(South America) 

19. Female glumes subequal, acuminate to subulate, not aristate; female 

20. Inflorescences 1-4 (usually 1). borne at the uppermost nodes, open 

(Amazonian Peru and western Brazil) 

20. Inflorescences -10, borne at the uppermost nodes, contracted and 

reaching about the same height. ................ 

(AmapB, Brazil) 

18. Female anthecium with long, flat macrohairs toward the base and apex 

on the ventral surface, otherwise glabrous ............. 21 

21. Male spikelets 3.5-4.7 mm long, hispid; pseudopetiole scaberulous . ........................ 16. 0. maranonensis 

(northern Peru) 

21. Male spikelets 8.5-12 mm long, glabrous; pseudopetiole densely 

hispid on both surfaces ..................... 22 

22. Panicles 4-9 cm long, 1-5.5 cm wide, the lower branches whorled 

to alternate; leaf blades ovate-lanceolate to lanceolate, 4.3-1 1.5 

cm long, 1-2.2 cm wide. ........... 10. 0. hurnilis 

(southeastern Brazil, Paraguay, and Argentina) 

22. Panicles 9-23 cm long, 5-18 cm wide, the lower branches whorled; 

........................ 13.0. latispicula, new species 

.......................... .4. 0. ciliatifolia 

anthecium apiculate or acuminate ............... .20 

.......................... 11.0. juruana 

.................. 1. 0. amapana, new species 

leaf blades ovate-lanceolate, 17-27 cm long, 4-8.5 cm wide . ........................ .8.0. glaberrima 

(San Luis Potosi, Mexico to Honduras; Peru; eastern Brazil) 

1. Olyra amapana Soderstrom & Zuloaga, new species 

FIGURES 1,2, 15 

TYPE SPECIMES.-BRAZIL, AMAPA: Rio Araguari, vicinity 

Camp 13, 1"45'N, 52"W, in shadow of upland forest, J.M. 

Pires, W Rodrigues, and G.C. lrvine 51536 (holotype, IAN; 

isotypes, NY, US sheet no. 2381088). 

Gramen perenne, caespitosum, culmis erectibus; nodi 

incrassati, pilosi. Laminae lanceolatae, 14.5-17 cm longae, 

3.5-4.2 cm latae, glabratae, basi asymmetricae, apice symmeuicae; pseudopetioli 0.3-0.4 cm longi: ligulae membranaceae, 

0.7 mm longae. Paniculae contractatae, parvae, 2.7-4 cm 

longae, 1-1.5 cm latae. Spiculae femineae lanceolatae, 11 

mm longae, 2.3 mm latae, albidae, glumis inter se subaequalibus, anthoecio longioribus; gluma inferior subulata, 7-nervis; 

gluma superior subulata, 7-nervis, anthoecium lanceolatum, 

apiculatum, 9.7 mm longum, 2.4 mm latum, pilosum. Spiculae 

masculae lanceolatae, 3.3-4.8 mm longae, 0.7-0.8 mm latae, 

albidae; lemma subulatum, 3-nerve: palea 3-4 mm longa, 

2-nervis. 
6 SMITHSOMAN CONTRIBUTIONS TO BOTANY 
NUMBER 69 7 

FIGURE 2.-Detail of female anthecium, Olyra ampanu: a, dorsal side (x 50); 6, detail of the ventral side 

showing hairs (X 100); c, base of the dorsal side showing germination lid (x 30); d, apex of the lemma, stomata 

(X 1ooO). All micrographs of Pires et al. 51536 (US). (Reduced to 77% of indicated magnifications for 

publication.) 

Cespitose perennials; Culm erect, these shortly pubescent, 

with whitish, appressed hairs toward one side of the culm; 

internodes cylindrical, hollow; nodes thickened, dark, shortFIGURE I.--Olyra U~M~M: a, flowering branch with inflorescences from 

upper nodes of culm (X 0.6); b, simple inflorescence with terminal female 

spikelets and lateral male spikelets (X 3.5); c, single inflorescence showing 

pedicels of spikelets (x 3.5); d, lower glume of female spikelet (x 7); e, upper 

glume of female spikelet (x 7); f, lemma of female spikelet (x 7); g, palea of 

female spikelet, inside view (x 7); h, female anthecium with internode and 

pedicel, palea side (~7); i, female anthecium with internode and pedicel, lemma 

side (X 7); j, palea of female anthecium, dorsal view (X 7); k, palea of female 

anthecium, ventral view (x 7); 1, gynecium and lodicules (x 10); m, male 

spikelet, side view (X 10); n, lemma of male spikelet, inside view (x 10); 0, 

lemma of male spikelet, outer view (x 10); p, palea of male spikelet, side view 

(X 10); q, palea of male spikelet, inside view (x 10); r, male flower (x 10); s, 

pseudopetiole, lower surface (x 5.5); f, pseudopetiole, upper surface (x 5.5). 

All components based on Pires et al. 51536 (US). Illustration by A.R. 

Tangerini. 

pubescent on one side. Leaves with sheaths stramineous, 

strongly ribbed, short-pubescent with appressed hairs, more 

densely so toward the borders, otherwise glabrous; ligules 

membranous, -0.7 mm long, shortly ciliate at the tip; 

pseudopetiole 0.3-0.4 cm long, short-pubescent on both 

surfaces; blades lanceolate, 14.5-17 cm long, 3.5-4.2 cm 

wide, glabrous, narrowed to an asymmetric and truncate base, 

the apex symmetric, the upper margins scabrous, otherwise 

glabrous, the midnerve prominent and scabrous on the abaxial 

surface. Inflorescences -10, paniculiform, borne from the 

uppermost nodes, long-exserted with peduncles longitudinally 

ridged and scabrous to hispid, all the panicles reaching 

approximately the same height; panicles contracted, small, 

few-flowered, 2.7-4 cm long, 1-1.5 cm wide, the lower 

branches whorled to alternate, the upper ones alternate, with 

numerous long to short-pedicelled male spikelets arranged on 

tertiary branchlets, each panicle with 1, rarely 2 to 4, terminal 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

female spikelets; axis longitudinally ridged, scaberulous, the 

axis of the branches similar to the main axis; pedicels of the 

female spikelets thickened, smooth, glabrous, those of the 

male spikelets smooth and glabrous. Female spikelets fusiform, 

glabrous, whitish, 11 mm long, 2.3 mm wide, the glumes 

subequal, the anthecium a little shorter than the glumes; lower 

glum subulate, 7-nerved, glabrous on the outer surface, shortly 

pilose toward the apex of the inner surface; upper glum 

subulate, 10.3 mm long, 7-nerved, scaberulous toward the apex 

of both surfaces; anthecium fusiform, apiculate, 9.7 mm long, 

2.4 mm wide, smooth and shiny, covered with long, whitish, 

cylindrical macrohairs. Mule spikelets fusiform, 3.3-4.8 mm 

long, 0.7-0.8 mm wide, whitish, scaberulous on the outer 

surface to glabrous; lemma subulate, 3-nerved; palea 3-4 mm 

long, 2-nerved. 

DISTRIBUTION.-Endemic to wet forests of Amapa, Brazil 

(Figure 15); known only from the type collection. 

DIscussIoN.-This species resembles 0. juruana, from 

which it differs by the numerous, long-pedunculate axillary 

panicles present at the upper nodes, each panicle with one, 

rarely two to four, terminal, female spikelets. 

2. Olyru buchtieienii Hackel 

FIGURES 3,4 

Olyra buchtienii Hackel, 1912:20. [Type: "Bolivia: In silvis ad San Carlos 

prope Mapiri, 750 m.s.m., 1907, leg. Dr. 0. Buchfien no. 1157." Holotype, 

W, not seen; isotypes, F, US (sheets no. 1099260 and 2877960).] 

Cespitose perennials, -3 m tall. Culms erect, branched at the 

upper nodes; internodes cylindrical, hollow, glabrous; nodes 

thickened, dark brown, glabrous to shortly pilose. Leaves with 

sheaths glabrous, 13-20 cm long, striate, with membranous 

FIGURE 3.-Detail of female anthecium, Olyra buchtienii: a, apex, dorsal side (x 50); b, apex of palea showing 

rounded excavations (X 100); c, dorsal base of the anthecium showing germination lid (X 40); d, detail of the 

excavations of the palea (x 700). Micrographs a,c of Buchrien 1157 (US) and b,d of Buchtien 40 (US). (Reduced 

to 77% of indicated magnifications for publication.) 
NUMBER 69 9 

FIGURE 4.-Distribution of Oiyra buchtienii, 0. caudata, 0. giaberrima, and 0. hoittwniana. 

margins: auricles membranous, glabrous: ligules membranous, 

conspicuous, 3-5 mm long, brownish, glabrous: pseudopetiole 

0.4-0.6 cm long, dark brown, sparingly pubescent or more 

frequently glabrous; blades lanceolate, 20-30 cm long, 

3.5-7.3 cm wide, narrowing from an asymmetric base to a 

symmetric, subulate apex, glabrous on both surfaces, the 

margins scabrous, the midnerve prominent. Inflorescences 

umbelliform on the upper nodes: panicles lax, -12 cm long, 

15 cm wide, the branches spreading, conjugate to verticillate 

from the base of the panicle, with male spikelets in pairs on 

short pedicels, the pedicels congested in fascicles of 6-8 and 

alternate on either side of a triquetrous axis, 1 or 2 terminal 

female spikelets on each branch; pedicels of the male spikelets 

short-hispid, those of the female spikelets thickened, 0.6-3 

cm long, triquetrous, flattened toward the base, scabrous to 

short hispid: main axis hispid to scabrous, the axils of the 

branches dark brown, hispid. Axillary panicles similar to the 

terminal one. Female spikelets ellipsoid, subulate apically, 

28-38 mm long, 3.6-3.9 mm wide, both glumes longer than 

the anthecium: lower glume long subulate to aristate apically, 

9- 15-nerved, pubescent to hispid on the outer surface, densely 

hispid on the inner surface with short hairs; upper glume 

subulate to aristate apically, 20-29 mm long, 7-9-nerved with 

transverse veinlets, pubescent to hispid on the outer surface, 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

with short appressed hairs on the inner one; anthecium 

lanceolate, acute apically, -10.7 mm long, 3.3-3.5 mm wide, 

glabrous, smooth and shining, whitish to stramineous, with 

rounded excavations on the apex of the palea. Male spikelets 

lanceolate, 4.9 mm long, 1-1.2 mm wide, or less as long 

as the female spikelets, short- to long-hispid; lemma acute, 

3-nerved, short-hispid on the outer surface, otherwise glabrous; 

palea 3.5-4.8 mm long, 2-nerved, sparingly hispid to glabrous; 

anthers 3,2-4 mm long. 

DISTRIBUTION.-Only known from Bolivia, in lowland 

forests of the Department of La Paz (Figure 4). 

ADDITIONAL SPECIMENS SEEN.-BOLIVIA, LA PAZ: San 

Carlos, Buchtien 40 (NY, US); valley between Tipuani and 

Apolobamba, Weddell 4581 (US). Nor Yungas, 13.5 km above 

San Pedro, Gentry and Solomon 44515 (MO, US). 

DIscussIoN.-Olyra buchtienii resembles 0. latifolia, 0. 

caudata, and 0. holttum‘ana, from which it differs by the 

presence of noticeably umbelliform inflorescences similar to 

those of 0. obliquifolia and the panicles with all branches 

whorled and reaching nearly the same height. The male 

spikelets are smaller than in these related species, measuring 

only about */7 the length of the female spikelets, and are borne 

in discrete fascicles along the panicle branches, a distinctive 

character not seen in other species in the genus. 

3. Olyra caudala Trinius 

FrcLJRes 4,5 

Olyra caudata Trinius, 1836:292. [Type: “Peruvia, in sylvis densis, Tocache.” 

Holotype, LE, not seen; fragment of the holotype, US sheet no. 2877958.1 

Olyra dimidia& Hochstetter ex Steudel, 1853:36. [Type: “(Hrbr. Hostm. nr. 

786a). Surinam.” Holotype, P, not seen; fragment of the holotype, US sheet 

no. 2877957.1 

Olyra pittieri Hackel, 1901:461. [Type: “Costa Rica: in planitie silvatica 

Cardoncellol (CordonciUal, near Rio Angel, Prov. Heredia). Piflier 3639.” 

Holotype, W, not seen, fragment of the holotyp, US sheet no. 2877933; 

isotype, MO.] 

Cespitose perennials forming dense clumps, with thick 

adventitious roots. Culms erect, branching at the upper nodes, 

cylindrical, 1-2.5 m tall, arcuate; nodes thickened, shortpilose. Leaves with sheaths pubescent or densely papillosepilose to glabrous, the margins short-ciliate; auricles present, 

membranous; ligules membranous and short-ciliate at the apex, 

conspicuous, 5-10 mm long, pubescent on the abaxial surface, 

the adaxial surface glabrous, smooth; pseudopetiole 0.5-0.8 

cm long, brownish, short to densely pilose on its surface; blades 

ovate-lanceolate, 18-30 cm long, 6-10 cm wide, greenish, 

asymmetric and narrowed basally, symmetric and acuminate 

apically, short-pilose to scabrous, with a prominent midnerve 

and scabrous margins. Panicles umbelliform, lax, diffuse, 

15-20 cm long, 10-20 cm wide, the lower branches whorled, 

spreading and reaching approximately the height of the upper 

ones, these alternate or verticillate, the male spikelets on short 

pedicels below, with the pedicels paired or in fascicles along 

either side of a triquetrous axis, a single terminal female 

spikelet on each branch; main axis longitudinally ridged, 

scabrous, the mils of the branches densely pilose, the axis of 

the branches triquetrous, with one side flattened; pedicels of 

the male spikelets thin, short-pilose to scabrous, those of the 

female spikelets thickened, scabrous to short-pilose. Axillary 

panicles usually present, similar to the terminal one. Female 

spikelets ovoid, long-subulate apically, 30-48 mm long, 

4.5-5.5 mm wide, the glumes subequal and longer than thc 

anthecium (4:l); lower and upper glume long-subulate with it 

geniculate awn 20-30 mm long, 9-nerved with transverse 

veinlets, densely pilose on the inner surface, especially toward 

the apex, short-pilose on the outer surface; anthecium ovoid, 

acute apically, 8.2-10 mm long, 4-5.2 mm wide, stramineous, 

brownish to olivaceous at maturity, smooth and shining, 

glabrous, shortly stipitate at its base, the stipe -0.4 mm long; 

lemma with prickle hairs on the upper margins; palea pitted 

toward the apex and on the basal margins. Caryopsis ovoid, 

brownish, 6-6.8 mm long, 3.2-3.8 mm wide; hilum linear, 

reaching the entire length of the caryopsis; embryo 1/6-1/7 the 

length of the caryopsis; germination lid rounded. Male spikelets 

lanceolate, brownish, 4-6.6 mm long, 0.7-1 mm wide, hispid; 

lemma short-aristate, 3-5-nerved, hispid, more so toward the 

apex and margins; palea 3.2-5.8 mm long, 2-nerved, sparingly 

pilose toward the middle, otherwise glabrous; anthers 3 mm 

long. 

DISTRIBUTION.-Mesoamerica (Costa Rica and Panama) 

and the West Indies (Trinidad) to South America (Colombia, 

Venezuela, Surinam, Guyana, Peru, Bolivia, and Brazil), in 

forests between 150 and 1 lo0 m elevation (Figure 4). 

ADDITIONAL SPECIMENS SEEN.-BOLIVIA, COCHABAMBA: 

Cant6n Chuquioma, Campamento Isarsama de la UMSS al rio 

del mismo nombre, Beck 1562 (US). 

BRAZIL, ACRE: Aldeota between Parangaba and Papagaio, 

Maas et al. PI3110 (F, US); Rio Branco, Colonia Penal, 

Emygdio 1815 (R, US); Rio Branco, Zoobotanical Garden of 

the Universidade Federal do Acre, Lowrie et al. 152 (MO). 

AMAZONAS: Near Tres Casas, Krukoff 6529 (F, MO, NY, US); 

Serra dos ~urucucus, Prance et a/. 10168 (Us). MAT0 GROSSO: 

Rio Aripuanal, Estrada Sta. Elena, Cordeiro 127 (US); Pacas 

Novas, afluente do Mamore, Kuhlmann 513 (RB, US). PARA: 

Parque Indigena do Tumucumaque, Rio Paru de Oeste, Missao

Tiriyo, Cavalcante 2334 (MO, US); Rio Vermelho, Serra do 

Cachimbo, BR 163 Cuiaba-Santarem, km 867, Silva et al. 

209 (MO, US), km 887, Prance et al. 25311 (MO, US). 

RONDONIA: Entre Cururui y Vilhena, Kuhlmunn 1864 (RB). 

RORAIMA: SCrra dos Sur~CucuS, Of Mission Station, 

Prance et al. 10168 (US). 

COLOMBIA, CAQUETA: 16 km from Florencia toward 

Belem, Luteyn et al. 4899 (MO, US); Rio Ortezuaga, Metucha, 

Woronow 6060 (US); Nova Granada, Salitre, Andre 1141 (US). 

COSTA RICA, ~1O16~: A few km west of Bribri, Gomez et 

al. 20316 (CR, WIS). SAN JoSE: Vicinity of El General, Skutch 

4238 (NY, US). 
NUMBER 69 11 

FIGURE 5.-Detail of female anthecium, Olyra caudafa: a, apex of the anthecium, dorsal side (x 50); b, apex 

of the anthecium, ventral side (X 50); c, detail of macrohairs on the margins of the lemma (x 500); d, surface 

of the lemma (X 200). Micrographs a, d of Allard 20195 (US), b, c of Krukoff 6529 (US). (Reduced to 77% of 

indicated magnifications for publication.) 

GUYANA: New Cabalebo, Courantyne River, Jenman 134 

PANAMA, CoCLE: 16.7 km north of turnoff to Coclesito 

from Llano Grande, Hammel 1857 (MO, US); forest around 

Limon, Folsom 5835 (MO). DARIEN: Cerro Pirre, Bristan 577 

(MO), Gentry and Clewell 7119 (MO), Mori and Kallunki 

5447 (MO); vicinity of gold mine at Cana, Croat 37649 (MO). 

PERU, AMAZONAS: Alrededor de la comunidad Kusu, Rio 

Numpatkin, Ancash 76 (F, MO, US); quebrada Kayamas, 

Cenepa, Ancash 171 (F, MO, US); vicinity of Quebrada 

Shimpunts, near upper Rio Cenepa, Berlin 881 (F, MO, US), 

Kayap 1265 (F, MO); trail N of Cenepa to Tuhushiku Creek, 

Berlin 1850 (F, MO, US). HUANUCO: Cueva de Las Lechuzas, 

cerca de Tingo Maria, Ferreyra 13222 (US); Tingo Maria, 

Asplund 12308 (R, US), Allard 20915, 22583 (US); al este de 

Tingo Maria, cerca a1 Cerro Quemado, Schunke 10127 0. 

(US). 

LORETO: Padre Abad, Schunke 5468 (MO); Yurimaguas, lower 

Rio Huallaga, Killip and Smith 27975 (F, NY, US); a1 este de 

Tingo Maria, cerca a1 cem Quemado, Schunke 10127 0; 

Nature Reserve, Barbour 4786 (MO); 39 km SW of Puerto 

Maldonado, S.F.Smith et al. 160, 551 (US). SAN MARTI^^: 

Carretera a1 Rio Tocache, Plowman and Schunke 7461 (F, 

US), Schunke 3587 (F, MO, US); Santa Rosa de Mishollo, 

Schunke 6804 (F, MO, US); puente de Rio Uchiza, Schunke 

7719 (WIS); Quebrada de Ishichimi, cerca de Tocache, 

Schunke 10045 (F, MO, U); trail up Rio Huallaga Valley 

toward Limon, Gentry et al. 25552 (F, MO); trail to Schunte, 

10 km W of Tocache Nuevo, Plowman et al. 11368 (F, MO, 

SURINAM: Maratakka River, without collector, Jan 2,1915 

without locality, Ule 6299 (Us). MADRE DE DIOS: Tambopata 

US). 

(US 1914625); Paramaribo, Wullschlaegel s.n. (US). 
12 SMITHSONIANCONTRIBUTIONS TO BOTAhY 

TRINIDAD: Cedros, Broadway s.n., Nov 4, 1916 (US); 

without locality, Fendler 967 (US). 

VENEZUELA, APURE: Selva de Cutufi, between Cutufi on 

the Rio Cutufi and the Rio Sanare, Davidse and Goruder 

21 724 (MO). BOLIVAR: Near Campamento Las Pavas, Steyermark 117188 (MO). MONAGAS: La Hormiga area, between La 

Pica and Cano Colorado, E of Maturin, Wurdack and 

Monachino 39482 (NY, US). TACHIRA: Slopes of Cerro of 

Cuchilla La Pabellana, W of San Joaquina de Navay, 

Steyermark 119386 (MO). 

DIscussIoN.-”he female spikelets of Olyra caudata 

resemble robust, long-awned versions of those of 0. latifolia. 

As in 0. latifolia, the female glumes disarticulate early, 

completely exposing the anthecium, which becomes brownish 

to olivaceous at maturity. Davidse (1987) has postulated that 

the florets “may mimic edible berrylike fruits that are eaten 

by frugivorous birds.” As in some other species of the genus, 

a short internode is present at the base of the anthecium. 

4. Olyra ciliatifolia Raddi 

FIGURES 6, 15 

Olyra ciliafifolia Raddi, 1823:19. [Type: “prope Rio Janeiro.” Holotype: PI, 

not seen, fragment of the holotype, US sheet no. 2879005.1 

Olyra cuneafifoliu Desvaux, 183 1:210. [Type: “Reperitur in Brasilio.” 

Holotype, P, not seen, fragment of the holotype, US sheet no. 2877953.1 

Perennial in cespitose clumps with 10 to 20 culms in a 

fascicle. Culms erect-ascending, geniculate or not at the lower 

nodes, 50-130 cm tall, unbranched; internodes hollow, shortly 

pubescent with retrorse, whitish and appressed hairs; nodes 

purplish, shortly pilose and thickened. Leaves with sheaths 

mottled, sparsely pilose with short whitish hairs to glabrescent, 

the margins ciliate, more so toward the tip; ligule membranousciliate, small, 0.4 mm long; pseudopetiole 0.2 cm long, 

stramineous to greenish, pilose, the borders long-ciliate to 

scabrous; blades ovate-lanceolate, 9-24 cm long, 3.6-8.5 cm 

wide, ascending, asymmetric and txuncate basally, the apex 

symmetric and acuminate, the margins ciliate, both surfaces 

scabrous, the midnerve prominent toward the base. Inflorescences borne at the uppermost nodes, long-exserted in hispid 

peduncles, paniculiform; panicles pyramidal, lax, 9.5- 19 cm 

long, 3.5-10 cm wide, the lower branches whorled and with 

male spikelets paired, one short, the other long-pedicelled on 

spreading tertiary branchlets, the upper ones alternate and with 

male spikelets on pairs below and a single terminal female 

spikelet on each branch; axis longitudinally ridged, hispid to 

scabrous, the branches and branchlets slender, triquetrous, 

scabrous; pedicels of the female spikelets thickened, smooth, 

glabrous. Female spikelets lanceolate, aristate, 1 1.6-23 mm 

long, 1.8-3 mm wide, the glumes unequal; lower glume 

-10-12 mm long, pilose on the apex of the inner surface and 

toward the outer margins, otherwise glabrous, 5-7-nerved with 

transverse veinlets, long-aristate with the awn scabrous; upper 

glume acuminate, 7.7-10.2 mm long, with a pubescence 

similar to that of the lower glume, 5-7-nerved with transverse 

veinlets; anthecium fusiform, acuminate, 6.5-7 mm long, 

1.6-3 mm wide, smooth and shining, whitish, dark at maturity 

with black spots, sparsely pubescent with long cylindrical, 

whitish macrohairs, more densely so toward the apex and base; 

lemma 5-7-nerved. Caryopsis long-ellipsoid, brownish, 4.3-5 

mm long, 1.4-2 mm wide; hilum linear, reaching the entire 

length of the caryopsis. Male spikelets fusiform, 5.4-10.7 

mm long, reddish to purplish, scabrous to shortly pubescent 

on the outer surface, aristate, awn -3 mm long; lemma aristate, 

3-5-nerved, sparsely pubescent toward the apex, otherwise 

glabrous; palea acute, 4.7-8.6 mm long, 2-nerved, sparsely 

pubescent toward the apex, otherwise glabrous, anthers 2.7-3.7 

mm long, 

DISTRIBUTION.-Colombia, Venezuela, Guyana, Brazil, 

Paraguay, Bolivia, and Argentina (Figure 15), often growing 

in disturbed places, from 100 to 1200 m elevation. 

ADDITIONAL SPECIMENS SEEN,- ARGENTINA, CORRIENTES: Itati, dense woodland on the bank of the River Parana, 

Pedersen 7013 (US); Perichbn, Quariin and Gonzalez 3094 

(US). MISIONES: Ruta 12, arroyo Aguaray Mini, Fernandez el 

al. 53 (US); arroyo Santo Pipo, Schwarz 2595 (MO, US); Santo 

Pipo, Schwarz 3445 (US, WIS); arroyo Yabebiri, Schwarz 

4530 (MO, US); Colonia Nacanguazu, Schwarz 1846 (MO); 

Corpus, Schwarz 1928 (MO, US); Iguazd Falls, Krapovickas 

and Cristobal 13701 (US); El Alcazar, Schwindt 1456 (US); 

breto, Ekman 675 (US), Montes 1779 (F, US), 2047 (MO, 

US), 27410 (F, MO, US); Puerto Santo Pipo, Schwarz 2641 

(MO); Makako, Schwarz 2917 (MO, US); Puerto Mani, 

Schwarz 10295 (WIS); Puerto Aguirre, Parodi 4395 (US); 

Puerto Iguazu, Bridarolli 3619 (US); Puerto Leoni, Schwarz 

7699 (US, WIS); Puerto Mineral, Schwarz 7579 (US); 

Cainguas, Schwindt 592 (WIS); Santa Mari’a, Montes 1645 

(MO, US). 

BOLIVIA, BENI: Huachi, head of Beni River, White 1043 

0; Pampas near lake Rogagua, Rusby 1669 (NY, US). 

BRAZIL, ACRE: Beira do Rio Branco, -5 km N de Boa 

Vista, Black 51-13779 (IAN, US); without locality, Ule 8047 

(US). BAHIA: Rodovia BA-265, 6 km W of Caatiba, Mori and 

Santos 11566 (MO, RB); 19 km N of Esplanada on road to 

Jaquera, Calderon et al. 2411 (US). CEARA: Grangeiro 

Taboleiros, von Luetzelburg 24257 (Us). DISTRITO FEDERAL: 

altos do corrego ‘Mato Grande,’ afluente margem esquerda 

do Ribeirao Papuda, Filgueiras and Pereira 850, 851 (US); 

-30 km S of Brasilia, on road to Belho Horizonte, Irwin and 

Soderstrom 5623 (US); Corrego Landim, -20 km NE of 

Brasilia, Irwin et al. 15679 (MO, US). Brasilia, Pires et al. 

9324 (UB). GOIAS: 12 km W of village of Presidente Kennedy, 

Plowman et al. 8159 (MO); lower slopes of Serra Dourada, -30 

km E of Goias Velho, Irwin et al. 11868 (F, MO, US); 24 km 

NE of Catalao, Irwin et al. 25106 (US); 75 km N of Corumba

de Goias on road to Niquelandia, Irwin et al. 19030 (F, MO, 

RB, US); Rio das Contras, Araguaina, Irwin et al. 21121 (F, 

MO, US); -9 km S of Guara, Irwin et al. 21535 (US); between 
NUMBER 69 13 

FIGURE 6.-Detail of female anthecium, Olyra ciliafifolia: a, anthecium, dorsal side (x 30); b, anthecium, ventral 

side showing the apex of the plea with rounded excavations and lemma (X 200); c, apex of the lanma showing 

cylindrical macrohain (x 200); d, detail of a macrohair (X 1ooO). Micrographs a,b,d of Chase I1146 (US), c of 

Chase 11487 (US). (Reduced to 77% of indicated magnifications for publication.) 

Itabira and Goiabeira, Chase 11487 (US); between Viannapolis 

and Ponta Pora, Chase 11318 (F, MO, NY, RB, US); 

Goyandira, Chase 11575 (US); Santa Rita do Paranahyba, 

Chase 12098 (US); vicinity of Goiabeira, Chase 15022 (US); 

vicinity of Goiabeira, between Annapolis and Goias, Chase 

11506 (F, MO, US); Santa Rita do Araguaya, on Rio Araguaya, 

Chase 11818 (US). MARANHAO: Castanhede, Sucre 9386 (US); 

Caxias to Barra do Corda, Swailen 3522 (NY, US), 3553,3584 

(US), 3555 (R, RB); Grajahu to Pbrto Franco, Swallen 3774 

(US); south of main house of Fazenda Morros, about 35 km 

south of Loreto, Eiten and Eiten 3738A (MO, NY), 4306 (F, 

NY, US), 10633 (US); between the rios Balsas and Parnaiba, 

about 40 km south of Loreto, Eiten and Eiten 3827 (F, NY, 

US); 55 km due W of Barra do Corda, Eiten and Eiten 10276 

(US). Perizes, Black et af. 54-16511 (IAN, US); Palmerinha, a 

74 km de Tuntum, Sifva and Santos 712 (US). MATO GROSSO: 

Agua Quente, prope Palmeiras, Lindman A2597 (US); along 

Rio Salo Lourenco, Barca, Chase 11975 (US); between 

Rondbnopolis and Sao LourenCo, Chase 12001 (US); sitio do 

Dr. Alfredo Neder, Sucre 10380 (RB); without locality, H. 

Smith 206 (Us). MATO GROSSO DO SUL: Dourados, Colonia 

Agricola Federal, Swallen 9406 (US): Aquidauana, Chase 

11059 (US); vicinity of Dourados, Chase 10999, 11009 '12 

(US), 11006 (RB, US); Urucum, Corumba, on Rio Paraguay, 

Chase 11146 (US). MINAS GERAES: Caldas, Widgren 1215 

(US); Faria, serra da Bocaina, Chase 10539 (US); Funao, 

Macedo 1002 (US); Lavras, Chase 8749 (F, MO, NY, RB, 

US), 8788 (US); along railway between Lavras and Formiga, 

Dorsett 214b (US); Serra do Cipo, km 100, Chase 9291 (F, 

MO, NY,US); Serra do Cipo, in cleared field near Vaccaria, 

Chase 9136 (F, MO, NY, US); Serra do Curral, Acaba Mundo, 

Chase 8960 (F, MO, US); Barreiras, Kuhfmann s.n., Feb. 20 
14 SMITHSONIANCONTRIBUTIONS TO BOTAW 

1926 (US); Buritys, near Rio Sao Francisco, Chase 10469 (F, 

MO, US); -10 km NE of Francisco Sa, road to Salinas, Irwin 

et al. 23143 (F, MO, US); -5 km SE of Paracatu, Irwin et al. 

26191 (F, MO, US); between Urubli and Cambuhy, valley of 

Rio Anzol, Dorsett 154b (US); Sa Miguel, NW of Formiga, 

Chase 10550 (NY,US); Santa Terezinha, Macedo I661 (RB, 

US); between Uberlandia and Rio Paranahyba, Chase 1161 7 

(US); without locality, Widgren 914 (R, US); Morro do Pau 

Lavrado, Heringer 3800 (UB); 30 km from Paraopeba, 

Heringer 5901 (UB). PARA: 20 km W of Redencao, near 

3827 and 4306 from Brazil. Hunziker et al. (1982) have 

recorded a chromosome number of n = 11 for the species. The 

common name is recorded as “paja carmzo” in Venezuela. 

5. Olyru ecuudutu Doell 

FIGURES 7, 8a,b, 34 

Olyra ecaudafa Doell in Martius, 1877:326. [Type: “In Guiana gallica 

(Lepriew n. 547)” Holotype, P, not seen, fragment of the holotype, US 

sheet no. 2877952.1 

Chego sao JOaO and ~roncamento Sam Teresa, Plowman Robust perennials with large dense clumps of up to 50 

et 8451 (F* Moao, Rio do Vargem, cubs. Culm up to 3 or 4 tall, frorn a thick, short

Swallen 9001 (Us); km Of Swallen 8741 ps); rhizomatous base, geniculate-ascending and bearing fully 

Parque Nacional de Iguacu, 9 km S of Jardinopolis along the 

road to Medianeiva, Davidse et al. I1222 (MO, US), Falcao 

100 (R); Rio Ivai, Hatschbach 15729 (US); 6 km SE of Sao 

Luis, Davidse et al. 11214 (MO). PIAuI: entre Piaba e Barrinha, 

sucre and silver 9428 (US). No DE JANEIRO: Serra do Baram, 

repre= do pkaquara, Soderstrom and sucre 1921 (CEPEC, 

RB, US), 1922 (CEPEC, US), Sucre 7485 m~). RORAIMA: 5 

developed leaves only toward the top, 5 or 6 of these ascending 

in a cluster, with the flowering culms usually separate from the 

vegetative ones; vegetative culms erect; flowering culms 

bladeless, to 2 m long; internodes thick, hollow, shortpubescent with whitish hairs to glabrous; nodes geniculate, 

thick, prominent, formed of the thickened basal rim of the 

km ao N de Boa‘Vista, Black51-13779 (US); pioximo a boca 

do Igarape Iguapira, Pires et al. 14-633(103) (US). sAo 

PAULO: Fazenda Capi, Clayton 4603 (US); Fazenda of Lauro 

Toledo, 1 km W of Rio Sao Mateus, Clayton 4576 (US); 

Fazenda Sao Jose, 1 km W of Rio Sao Mateus, Eiten et al. 

5889 (MO, US); Itapira, Hoehne 20338 (US); Nova Europa, 

Hoehne 13657 (US); Parque do Estado, Sao Paulo, Clayton 

and Eiten 4186, 4188 (US); Ribeirilo Preto, Lofgren 1544 

(Us). STATE AND LOCALITY UNKNOWN: Gardner 3523 (Us), 

Sellow, Brasil (US), Pohl, Brazil (US), Glaziou 14397 (US). 

COLOMBIA, CAQUETA: 16 km from Florencia toward 

Belem, Eiten and Eiten 10633 (MO). META: Ranch Menegua, 

about 100 km SE of Villavicencio, Schiefer 795 (GH, US); 

without locality, Moritz 685 (US). 

GUYANA: Without locality, Schomburgk 703 (US). 

PARAGUAY: In regione lacus Ypacaray, Hassler 12444 

(F, NY, US), 13021 (US); Pedro Juan Caballero, Morong 505 

(MO, NY, US); Villa Elisa, Pedersen 5909 (US); Estancia La 

Soledad, Santiago, Pedersen 3200 (MO, US); Cordillera de 

Altos, Fiebrig 480 (F, GH); Trinidad, Rojas 3071 (US); Villa 

Rica, Jorgensen 4098 (F, GH, MO, NY, US); in regione cursus 

superioris fluminis Apa, Hassler 81 94 (NY); without locality, 

Anderson I185 (US), Balansa 258 (US), Hassler I587 (NY). 

TRINIDAD: Port of Spain, Botanical Garden, Hitchcock 

10133 (US); without locality, Broadway 1918 (NY, US), 5085 

(MO), 9194 (US), Fendler 968 (NY), Purdie 26 (US). 

VENEZUELA, BARINAS: Ticoporo forest reserve, Breteler 

4015 (US). BOLIVAR: Near El Dorado, Irwin et al. 25106 

(MO); 1 km N of Paragua, Liesner and Gonzalez 5755 (US). 

GUARICO: Altagracia de Orituco-Taguay (carretera hacia 

Carmen de Cura), Aristiguieta and Agostini 6439 (US). 

DISCUSSION.-This is one of the commonest species in 

eastern Brazil, flowering between January and May. There are 

only female spikelets on the axillary panicles of Eiten and Eiten 

sheath above and the thickened upper rim of the internode 

below, flared, dark brown and with remorse hairs to glabrous. 

Leaves 5-7 per complement, clustered above the naked culms; 

sheaths with flattened, caducous, and brownish hairs along the 

overlapping margins, the rest of the surface sparsely pilose to 

glabrous; ligule a tiny brown membrane, 0.6-1.1 mm long, 

shortly ciliate at the tip; pseudopetiole dark brown, 0.3-0.5 

cm long, with brown, thick, flattened hairs on the adaxial 

surface, the abaxial surface with or without these hairs; blades 

ovate-lanceolate, 21-35 cm long, 5-7.5 cm wide, rigid, dark 

green, fully developed on the upper nodes, gradually narrowed 

above to an acute or acuminate, symmetric tip and slightly 

cordate, symmetric at the base, glabrous on both surfaces to 

sparsely pilose on the adaxial surface, the midnerve prominent, 

the basal margins short-ciliate, otherwise scabrous to completely glabrous. Inflorescences paniculiform, 2 or 3 from each 

of the uppermost nodes, some inflorescences short and found 

only at the base of the plant; panicles 10-20 cm long, 10-25 

cm wide, the lower branches verticillate, the upper ones 

alternate to opposite, stiffly spreading, the individual branches 

up to 15 cm long, usually with 2-6 female distal spikelets and 

below with male spikelets on short branchlets or in pairs, one 

long- and the other short-pedicellate, the panicles exserted on 

long, glabrous to sparsely pubescent peduncles; axis anti 

branches sparsely hirsute, longitudinally ridged, scabrous, the 

axils of the branches densely pilose. Female spikelets ovoid, 

acuminate to occasionally shortly aristate, 8-1 1.6 mm long, 

3-3.8 mm wide, the glumes subequal or the lower glume 

occasionally a little longer than the upper one, sparsely pilose 

toward the margins and nerves on the outer surface, shortly 

pilose toward the apex on the inner surface; lower glume 

apiculate to shortly aristate, 5-10-nerved with transverse 

veinlets; upper glume apiculate, 7-1 1-nerved with transverse 

veinlets; anthecium ovoid, 6.6-8.3 mm long, 2.7-3.4 mm 
NJMBER 69 15 

FIGURE 7.-Detail of female anthecium, Olyra ecaudata: a, anthecium, ventral side (x 50); b, anthecium, dorsal 

side (X 30); c, margin of the lemma with prickle hairs and palea with rounded excavations toward the margins 

(X 200); d, surface of the lemma with excavations (x 200). Micrograph a of Cremers 4478 (US), b-d of 

Soderslrom & Calderon 2292 (US). (Reduced to 77% of indicated magnifications for publication.) 

wide, indurate, whitish, with small excavations all over the 

lemma and palea; lemma 7- or 8-nerved, the margins with 

conspicuous, rigid, and long prickle hairs; palea 4-nerved and 

with rounded excavations toward the apex and upper margins. 

Caryopsis ellipsoid, brownish, 4.8 mm long, 2.2-2.6 mm 

wide; hilum reaching the entire length of the caryopsis. Male 

spikelets lanceolate, aristate, 7.5-11 mm long, 0.6-1 mm 

wide, greenish, glabrous to scaberulous; lemma 3-nerved, 

aristate, the awn 5 mm long; palea 2-nerved; anthers yellow, 

3.3-4.3 mm long. 

DISTRIBUTION.-Ranging from Costa Rica and Panama in 

Mesoamerica to Colombia, Venezuela, Surinam, French 

Guiana, Peru, Bolivia, and Brazil in South America (Figure 

34); growing in primarily humid forests, generally at elevations 

below 500 m, although Smith 2700 from Peru was collected 

at 2000 m. 

ADDITIONAL SPECIMENS SEEN.-BOLIVIA, PANDO: Cobija, 

Ule 9146 (US). 

BRAZIL, ACRE: About 80 km from Rio Branco on road to 

Xapuri, Calderon and Soderstrom 2292 (US); 125 km from 

Rio Branco on road to Porto Velho, Calderon and Soderstrom 

2305 (US); 8"54'S-72"51 W, Calderon and Soderstrom 2353 

(US). BAHIA: Bahia State Highway 2, 7 km N of Ibirataia City, 

Fazenda Santo Antonio, Calderon and Pinheiro 2223 (CEPEC, 

US, WIS), 2259 (US); Parque Nacional Monte Pascoal, 14 

km E of BR-101, 13 km N of Itamaraju, Soderstrom et al. 

2197, 2199 (CEPEC, US). ESPIRITO SAVTO: Reserva Florestal 

da Companhia Vale do Rio Doce, Luna Peixoto 416 (RB, 

US). MAT0 GROSSO: Fazenda Cachimbo, Cordeiro 1132 (MO). 

PARA: Alto Tapajos, Vila Nova, perto da Cachoeira do 

Chacorao, Pires 3951 (IAN, US); Belem, Floresta do Utinga, 

Pires 1432 (US); Castanhal, Colonia 3 de Outubro, Froes 
16 SMITHSONIANCONTRIBUTIONS TO BOTAhY 

FIGURE 8.-Field photos of Olyra species. Olyra ecaudafa: a, habit of plant (Brazil, 1972) showing dying culms 

after flowering; b, habit of plant showing plants before flowering. Olyra fasciculata: c, habit of plant (Brazil, 

1968); g, detail of panicle. Olyra lafifoh: d, habit of plant (Costa Rica, 1968); e, uppernodes with inflorescence; 

detail of panicle. All photographs by C.E. Calderon. 
NUMBER 69 17 

24873 (US), Black 49-8609 (US): 50-65 km N Of Gumpi, 

Belem-Brasilia, Prance and Silva 58683 (US); Serra dos 

Carajas, Azul, near camp at Serra Norte, Duly et al. 1937 (US). 

RONDONIA: Porto Velho, silva 391 (US); Rio Contra, 6 km 

from Jaciparana, Calderon et al. 2814 (US, WIS). 

COLOMBIA, VAUPES: Confluence of Macaya and Ajaju 

rivers, Puerto Hevea, Grass1 10019, 10029 (US). 

COSTA RICA, Borders of SAN JOSE and PUNTARENAS: Rio 

Convento, on the Pacific slope, Dodge et al. s.n. (CR). 

FRENCH GUIANA: Sur le Tamp, Saut Koumakou Soda, 

Cremers 4478 (US): without locality, Leprieur 547 (US). 

PANAMA, DARIE: Along ridge N of Ensenada El Guayabo 

that separates the Rio Jaque Valley from the Pacific Ocean, 

Knapp and Mallett 3181 (MO, US). PANAMA: Cerro Azul, 

6-10 km del lago Goofy, Calderon 2087 (US), Croat 17289 

(MO, US), Soderstrom 2003 (MO, US): Cerro Jefe, Calderon 

and Dressler 2122 (US): Zetek Trail 2100, Barro Colorado 

Island, Judziewicz 4437 (MO, US, WIS). 

PERU, LORETO: Bosque Nacional de von Humboldt, region 

of San Alejandro, Gentry and Revilla 16200 (MO, WIS). 

PASCO: 19 km W of Oxapampa, D. Smith 2700 (MO, US). 

SURINAM: 3 km S of Juliana Top, 12 km N of Lucie Rivier, 

Irwin et al. 55055 (US); vicinity of Kayser airstrip, 25 km 

above confluence with Lucie River, Maguire et al. 53950 (US); 

entre Hanover Savannah et Para, Hoock 93 (NY, US): -2 km 

below affluence of Oost Rivier, Maguire et al. 54095 (MO, 

US): near Brownsweg along railway, Wessels Boer 641 (US). 

VENEZUELA, ZULIA: 3 km E of the Rio de Or0 settlement 

on the Rio de Oro, Davidse et al. 18620 (MO, US). 

DISCUSsIoN.--Olyra ecauduta forms large clumps to 3/4 m 

in diameter, of tall erect culms reaching 3-4 m in height, and 

is found only in shaded areas of good, often primary, forest, 

flowering between October and July. The full-sized leaves 

develop only on the upper nodes where 4-6 ascending blades 

are clustered, creating a plume-like effect. Generally the 

flowering culms are borne separately from the sterile ones and 

bear only bladeless sheaths at their nodes. Caldeon noted on 

the label of one of her specimens (Calderon and Dressler 

2122) that the plants on Cerro Jefe in Panama were all sterile. 

Dressler, who had visited the locality over the years, remarked 

that the colony had never been in flower as long as he visited 

it. Two years later I visited the locality in his company and the 

entire colony was in flower with the plants dying (see specimen 

of Soderstrom 2003 from here). This same phenomenon was 

noted in plants collected in Bahia, Brazil (Soderstrom et al. 

2197) and on Barro Colorado Island, Panama (Judziewicz, 

1983, persxomm.). All plants of this species had flowered and 

died in the region. In this case both tall and short inflorescences 

were found, as in another specimen from Acre, Brazil 

(Calderon and Soderstrom 2353). A possible explanation is 

that the plant remains sterile for many years and at time of 

flowering the tall mature shoots flower as well as new small 

shoots that have developed from a weakened base. 

Flowering culms with fully developed leaves (Soderstrom 

2003 from Panama and Froes 24873 from Para, Brazil) are 

rarely encountered in herbarium specimens. 

6. Olyra fasciculata Trinius 

FIGURES 8c,g, 9, 11 

Olyrafasciculata Trinius, 1834:25 [page 113 in published form, 18351. [Type: 

“V. spp. Bahiens.” Holotype, LE, not seen.] 

Olyru heliconia Lindman, 190011, pl. 6. [Type: “Habitat in Brasilia centr., 

Matto Grosso, Matto do Curupira, in silva primaeva, mens. Febr. florens, 

Exp. I. Regnell., A,, 3017.” Holotype, S, not seen; fragment of the holotype, 

US sheet no. 2877949.1 

Perennial, short-rhizomatous grass, in cespitose clumps with 

20-50 culms in a fascicle, usually leaning on trees or shrubs. 

Culms erect, branching on the upper nodes, 1.5-3 m tall; 

internodes hollow, glabrous, cylindrical, up to 1 cm in 

diameter, the lower internodes with a sheath only or with a 

rudimentary or reduced blade; nodes glabrous, thickened, 

purplish. Leaves with sheaths short-pilose with retrorse hairs 

to glabrous, greenish to pale, the margins short-pilose to 

glabrous: ligule membranous, short-ciliate at the apex, 1-4 

mm long, pilose on the abaxial surface: pseudopetiole 0.4-1.3 

cm long, purplish to black, short-pubescent on both surfaces: 

blades ovate-lanceolate, 24-32 cm long, 5-13.2 cm wide, 

ascending, narrowed from an asymmetric base to a symmetric 

or asymmetric tip, the basal margins short-ciliate, otherwise 

scabrous, the midnerve prominent, abaxial surface pilose 

toward the base, otherwise glabrous. Inflorescences: 2 or 3 

borne on the uppermost nodes: terminal panicles 20-30 cm 

long, 16-25 cm wide, the branches divergent from the axis, 

drooping, the lowermost whorled and almost reaching the 

same length as the upper branches, 16-25 cm long, the upper 

ones alternate, sometimes 3-10 branches in one fascicle: 

peduncles densely pilose: axis longitudinally ridged, scabrous, 

the branches spreading, triquetrous, flattened on one side, 

hispid toward the base, otherwise scabrous, with 4-6 male 

spikelets borne alternately on short, scabrous branches on the 

lower portion and 1-5 female spikelets at the summit of each 

branch, the pedicels of the female spikelets thickened, pilose. 

Female spikelets lanceolate, greenish to brownish, 22-33 mm 

long, 2.9-3.2 mm wide, aristate or not, the glumes unequal 

and strongly nerved, pilose to glabrous: lower glume as long 

as the spikelet, acuminate to long-aristate, rigid, 5-9-nerved 

with transverse veinlets, the inner surface densely pilose to 

pilose only toward the apex, the outer surface pilose to scabrous 

toward the apex, otherwise glabrous: upper glume 17-20 mm 

long, acuminate to subulate apically, 7-9-nerved with transverse veinlets, with pubescence similar to that of the lower 

glume, anthecium fusiform, acute, 9-1 1.5 mm long, 2.5-2.8 

mm wide, glabrous, pitted, with oblong excavations over the 

entire surface, stipitate, the stipe 0.8-1 mm long: lemma 

5-nerved. Caryopsis fusiform, 6.8 mm long, 2 mm wide: hilum 

linear, reaching the entire length of the caryopsis. Male 

spikelets fusiform, 8-13 mm long, 0.7-1 mm wide, glabrous, 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 9.-Detail of female anthecium, Olyrafarciculufu: a, surface of the lemma showing oblong excavations 

(X 100); b, anthecium, dorsal side (X 50); c, base of the anthecium showing stipe (X 50); d, anthecium, ventral 

side (X 200). Micrographs a,c,d of Killip & Smith 23452, b, of Pereiru 2294 (US). (Reduced to 77% of indicated 

magnifications for publication.) 

pink to purplish; lemma acuminate apically, purplish, scabrous, 

3-nerved; palea 7-10 mm long, %nerved, scaberulous; anthers 

3,7.6-8.5 mm long. 

DISTRIBUTION.-Panama; South America from Peru and 

Bolivia to Brazil and northwestern Argentina (Figure 11); 

found on edges of forests, usually in full sun, between 200 and 

1600 m elevation. 

ADDITIONAL SPECIMENS SEEN.-ARGENTINA, SALTA: 

Quebrada del Acheral, Castellanos 21 (LIL); Finca El Candado 

Grande, Cuezzo and Legname 572842 (LIL); El Angosto del 

Rio Pescado, Meneses and Vervoorst 18 (LIL); Dept. Oran, 

Finca Yakulika, 1 km antes de Rio Pescado, Turpe 2790 (LIL, 

WIS); Rio Pescado, 9 km de la finca Yaculika, Vervoorst and 

Cuezzo 7841-C (LIL). 

BOLIVIA, BENI: Rurrenabaque, White s.n., Aug 1921 (NY), 

Beck 8219 (WIS); San Juan, Williams 958 (NY). COCHABAMBA: Antahuacana, Buchtien 6433 (US). SANTA CRUZ: 

Buena Vista, Steinbach 2010 (GH, US); Ascension de 

Guarayos, Krapovickas and Schinini 31702 (US). LA PAZ: 

Coroico, Hitchcock 22720 (NY, US); Polo-Polo bei Coroico, 

Buchtien 450 (F, NY), 3622 (US), 3623 (F); Yungas, Rusby 

31 (NY), Bang 508 (NY, US). 

BRAZIL, ESPIRITO SANTO: Vargem Alta, Fruteira, Pereira 

2294 (US). GOIAS: Santa Rita do Araguaya, on Rio Araguaya, 

Chase 12046 (us). MINAS GERAIS: Without locality, Macedo 

4274 (IAN, US). PARANA: Capa0 Bonito, Dusen 18000 (MO, 

US); 7 km E of Londrina, Swallen 8737 (US); Quatro Portoes, 

Pereira 7905 (F, RB, US), Hatschbach 10519 (F, US); 

Maringa Orto Florestal, Hatschbach 12929 (F, US); Parque 

Nacional do Iguacu, W bank of Rio Flonano, Lindeman and 

Haas 3558 (US); Rio Ivai, Hatschbach 15720 (US). RIO DE 

JANEIRO. Rio de Janeiro, Avenida Niemeyer, Chase 10003 (F, 
NUMBER 69 19 

MO, US); Leblon, Rosengurtt PE-3386 (US); Pao de Acucar, 

Chase 8404 (F, MO, NY, US), 9798 (US); Serra do Corcovado, 

Sumare, Calderon 2022 (US); Serra da Mendanha, 22”51’S, 

43”32’W, Soderstrom and Sucre 1952 (CEPEC, RB, US), 

Sucre 6406 (RB, US); Strada de Gavea, proximo a Salo 

Conrado, Calderon 2024 (RB, US); Vale de Bom Sucesso, 

“Caixa d’Agua,” Soderstrom and Sucre 1982 (CEPEC, RB, 

US), Sucre 2004, 2204 (US), Braga 1955 (RB). SAO PAULO: 

Ubatuba, Soderstrom and Sendulsky 1994 (US); without 

locality, Regnell239 (NY), Burchell 954 (US), Gardner 137 

(US). SANTA CATAFUNA: Aguas de Chapeco, Smith and Klein 

13107 (R, US), Reitz and Klein 16713 (US); Nova Teutonia, 

Phumann 459 (RB). STATE AND LOCALITY UNKNOWN: Burchell 6709 (US), Pohl sen. (US), Riedel 160 (US), 934 (GH, 

US), Spruce 4807 (US). 

PANAMA, COLON: Headwaters of the Rio Boqueron near 

fork with Rio Nombre de Diosito, Hammel 3951 (MO, US). 

SAN BLAS: El Llano-Carti road, 9”19’N, 78”55W, de Nevers 

et al. 5254 (MO, US). 

PERU, cuzco: Cerro San Pedro, Vargas 8498 (US); 

Quitemi, Chaver AIfaro 3387 (MO). HUANUCO: Huanuco to 

Tingo Maria, km 100, Rio Chinckao, Seibert 2297 (MO, US); 

Puente Durand, N of Huanuco, valley of Chinckao River, Stork 

and Horton 9574 (F, US), Vargas 5388 (US); along road 

between Chinchao and Puente Durand, McClure 21458 (US). 

JUNINColonia : Perene, Hitchcock 22056 (NY, R, US), 22070 

(US); 28 km N of San Ramon, on road to Oxapampa, Dillon 

and Turner 1439 (F, MO); La Merced, Killip and Smith 23452 

(US), 23788 (F, NY, US); Rio Paucartambo, near junction with 

Rio Perene, Seibert 2194 (MO, US); San Ramon, Killip and 

Smith 24755 (NY). SAN MARTIN: Juan Jui, Alto Rio Huallaga, 

Klug 4197 (F, MO, NY, US); Lamas, Williams 6449 (F, US); 

San Roque, Williams 7792 (US); without department, Pozuzo, 

Macbride 4572 (US); Santa Ana, Cook and Gilbert 1653 (US); 

in montibus sews flumen Mayo, prope Tarapoto, Spruce 480 

(NY). STATE AND LOCALITY UNKNOWN: Andes, Peru, Poeppig 

s.n., 1834 (US), Gardner 137 (NY). 

DISCUSSION.-Olyra fasciculata bears some resemblance 

to 0. obliquifolia but can be distinguished by its paniculate 

inflorescence bearing longer female spikelets on less thickened 

pedicels. Common names include “tacuarilla” (Bolivia) and 

“taquaril” (Brazil). Chromosome numbers of 2n = 14 (Calderbn 

and Soderstrom, 1973) and n = 7 (Hunziker et al., 1982) have 

been recorded. It usually flowers between October and April. 

7. Olyra jiluormis Trinius 

FIGURES 10, 11, 17a,b 

Olyrafiliformis Trinius, 1834:27 [page 115 in 1835 published form]. [Type: 

“V. spp. Bahiens.” Holotype, LE, not seen, fragment of the type, US sheet 

no. 287795 1; isotype, GH.] 

Cespitose perennials, short-rhizomatous, with 10-25 culms 

in a fascicle. Culms geniculate-ascending, occasionally decumbent and rooting at the lower nodes, climbing and arching, 

40- 125 cm tall, many-noded, unbranched, sparingly branching 

at the upper nodes; internodes cylindrical, hollow, sparingly 

pilose to glabrous, the lowest short, followed by two elongated 

internodes bearing a culm leaf without blade; nodes thickened, 

purplish, pilose with whitish retrorse hairs to glabrous. Leaves 

with sheaths strongly ribbed, greenish to purplish, the margins 

membranous, pilose toward the upper portion, otherwise 

glabrous; auricles conspicuous, membranous; ligule membranous, 0.5-1.8 mm long, glabrous on the adaxial surface, 

short-pubescent on the abaxial one; pseudopetiole -0.2 cm 

long, pale to purplish, densely pilose; blades oblong-lanceolate, 

11-14 cm long, 1.2-2.8 cm wide, narrowed and asymmetric 

at the base, the apex symmetric, scaberulous to scabrous on 

both surfaces, the abaxial surface short-hispid, the midnerve 

prominent on the abaxial surface, the margins scabrous. 

Inflorescences paniculiform, contracted, one to several on the 

upper nodes, long exserted, peduncles glabrous; panicles 5-10 

cm long, 0.7-2 cm wide, the branches alternate, spaced and 

appressed to the axis, each branch with male spikelets below 

and a single terminal female spikelet; an’s longitudinally 

ridged, scaberulous, the axis of the branches flattened, hispid 

to scabrous, the male spikelets in pairs or in short branchlets, 

one subsessile, the other short-pedicellate, the female spikelets 

on thick, short-pubescent pedicels. Female spikelets fusiform, 

17-24 mm long, aristate, whitish, glabrous to scaberulous, the 

glumes unequal, the anthecium approximately 1/3 the length 

of the spikelet; lower glume as long as the spikelet, 

long-aristate, the awn -6 mm long, 7-9-nerved with transverse 

veinlets, scaberulous on the outer surface, shortly pubescent 

toward the apex on the inner surface; upper glume 13-20 mm 

long, aristate, 7-9-nerved with transverse veinlets and 

pubescence similar to that of the lower glume, anthecium 

fusiform, acute, 6.8-8.5 mm long, 2.4-2.6 mm wide, pitted, 

with rounded excavations over the entire surface, glabrous; 

lemma 5-nerved. Caryopsis fusiform, brownish, 4.8-5.5 mm 

long, 1.8-2 mm wide; hilum linear, reaching the entire length 

of the caryopsis. Male spikelets lanceolate, acuminate, 4.3-5.7 

mm long, 0.5-0.8 mm wide, 1/3-1/6 the length of the female 

spikelets, hispid to glabrous, whitish; lemma 3-nerved; palea 

3.7-5.5 mm long, 2-nerved; anthers 1.8-2.5 mm long. 

DISTRIBUTION.-Endemic to lowland forests in Bahia, 

Brazil (Figure 11). 

Camaca-Canavieiras, 3 km E de Camaca Belem and Aguiar 

1389 (US); Lagoa Itaipe, at 1 km from the lake and 25 km SE 

of Urucuca, Caldero’n and Pinheiro 2172 (US, WIS); Fazenda 

Joao Elias, cacao plantation 3 km E of Camaca on road to 

Canavieiras, Soderstrom et al. 2123 (CEPEC, US); road from 

Una to Santa Luzia, 10 km SW of Una: Rio da Pimenta o 

Pimenteira, Fazenda S. Jorge, Calderon and Pinheiro 2267 

(CEPEC, US); 9 km from Una on the road to Santa Luzia do 

Salobro, Soderstrom et al. 2229 (CEPEC, US); 6 km N of 

ADDITIONAL SPECIMENS SEEN.-BRAZIL, BAHIA: Rodovia 
20 SMITHSONIAN CONTRIBUTIONS TO BOT.4“ 

FIGURE IO.-Detail of female anthecium, Olyra filiformk: a, apex of the anthecium, dorsal side (x 100); b, 

apex of the anthecium, ventral side (x 100); c, detail of the lemma (x 260); d, detail of the excavations (x 700). 

All micrographs of Soderstrom el al. 2123 (US). (Reduced to 77% of indicated magnifications for publication.) 

Apuarema on road Apuarema-Jaguaquara, Calderon et al. 

2373 (US). 

DIscUssI0N.-This delicate species, growing among weedy 

vegetation in semi-shaded forests, is quite unlike any other in 

the genus. In some areas of Bahia the plants are abundant on 

sandy soil and have a weak, more or less cespitose base. They 

appear to be annuals (not known in the Bambusoideae); 

certainly none of the plants look older than a year. The species 

is easily distinguished by the linear inflorescences with 

appressed female spikelets that consist of very long and 

acuminate glumes that far exceed the anthecium in length. The 

female anthecia are very deeply and conspicuously pitted. 

Apparently, after final flowering the flowering node may 

produce a tuft of new branches causing the topheavy culm to 

bend over to the ground where it takes root to produce a new 

plant (Soderstrom et al. 2123). In flower between March and 

July. 

8. Olyra glaberrima Raddi 

FIGURFS 4, 12,30e, f 

Ofyra glaberrimu Raddi, 1823:19. [Type: “Invenitur in Monte nuncupato 

Corcovado non procul ad Urbe Rio de Janeiro.” Holotype, PI, not seen, 

fragment of the holotype, US sheet no. 2877950.1 

Olyra semiovafa Trinius, 1826:249. [Type: “V. sp. Brad (Lungsdorfn.” 

Holotype, LE, not seen, fragment of the holotype, US sheet no. 2877927.1 

Olyra obliquu Desvaux, 1831:210. [Type: “Habitat in Brasilio.” Holotype, P, 

not seen, fragment of the holotype, US sheet no. 2877938.1 

Ofyra corcovadensis Wawra, 1866:180, pl. 95. [Type: “Corcovado an feuchten 

schattigen Orten; Wwr u. Maly coll. n. 504,” not seen, current location 

unknown.] 

Olyra semiovafa var. pubiflora Hackel, 1904:276. [Type: “In silvis in region,? 

cursus superioris fluminis Apa, Dec. n. 8194 (Hussler, Paraguay).” 

Holotype, W, not seen, fragment of the holotype, US sheet no. 2877925; 

isotypes, MO, P.1 

Olyra yucatana Chace, 1908:178. [Type: “Gaumer 2372, No. 125941 Herb. 

Field Columbian Museum.” Holotype, F, not seen, fragment of the holotype, 

US sheet no. 2877920.1 
NUMBER 69 21 

FIGURE 11.-Distribution of Olyra farciculata, 0. jiliformir, 0. latirpicula, 0. obliquifolk, 0. retrorsa, 0 

tamanquaream, and 0. wurdackii. 

Olyra semiovata var. pubescenr Hackel, 1910:46. [Type:"Paraguay septentr. lower nodes, unbranched; internodes cylindrical, glabrous to a 1909 leg. Fiebrig n. 5299 in Herb. Hassler." Holotype, W, not seen, shortly pubescent with reeorse hairs; nodes thickened, 

fragment of the holotype. US sheet no. 2877926.1 glabrous, purplish. Leaves with sheaths greenish to purplish, 

strongly ribbed, shortly hispid, the margins long-ciliate, more 

densely so toward the upper portion; auricles membranous; 

Cespitose perennials, short-rhizomatous, with 10-20 culms 

in a fascicle. Culms 60-200 cm tall, erect, geniculate at the 
22 SMITHSONIANCONTRIBUTIONS TO BOTANY 

ligule membranousciliate, small, 0.5-0.8 mm long; pseudopetiole 0.3-0.7 cm long, stramineous to purplish, densely 

pubescent with short, appressed whitish hairs on both surfaces; 

blades ovate-lanceolate, 17-27 cm long, 4-8.5 cm wide, 

ascending, asymmetric and truncate at the base, symmetric at 

the acuminate tip, glabrous, the margins scaberulous to shortly 

ciliate, the midnerve inconspicuous or prominent toward the 

base; Inflorescences paniculiform, borne from the uppermost 

nodes, long-exserted, the peduncles longitudinally ridged, 

glabrous: panicles lax, diffuse, pyramidal, 9-23 cm long, 5-18 

cm wide, the branches spreading, the lower ones verticillate 

and with numerous male spikelets only or occasionally with a 

terminal female spikelet, the upper branches alternate, with 

male spikelets below and a single terminal female spikelet in 

each branch; axis and branches longitudinally ridged, scabrous, 

the axils of the branches glabrous; pedicels of the female 

spikelets thickened, sparsely scabrous; pedicels of the male 

spikelets scabrous. Axillary panicles similar to the terminal 

one. Female spikelets fusiform, aristate, 15-23 mm long, 

2.5-3 mm wide, the glumes unequal and longer than the 

anthecium: lower glum long-aristate, scabrous on both 

surfaces or the inner surface glabrous, 5-9-nerved with 

transverse veinlets, the awn 5-11 mm long; upper glume 

acuminate to long-aristate, 10-14 mm long, scaberulous on 

both surfaces, 5-7-nerved: anthecium broadly ellipsoid, acute, 

6.5-8.6 mm long, 2.2-3 mm wide, whitish to stramineous 

with black spots at maturity, smooth and shining, densely 

pubescent toward the lower margins of the lemma, with long, 

flat, whitish to brownish macrohairs, shortly pubescent toward 

the apex: lemma 5-nerved; palea 2-nerved, with sparse, flat 

hairs toward the apex to completely glabrous. Caryopsis 

ellipsoid, brownish, 4.5-5.4 mm long, 2.8 mm wide; hilum 

FIGURE 12.-Detail of female anthecium, Oryra glaberrima: a, apex of the anthecium, dorsal side (X 50); b, 

apex of the anthecium, ventral side showing flattened macrohain (X 100); c, base of the anthecium showing the 

germination lid (x 40); d, detail of the hairs on the base of the lemma (x 100). All micrographs of Chose 12171 

(US). (Reduced to 77% of indicated magnifications for publication.) 
NUMBER 69 

as long as the caryopsis. Male spikelets fusiform, brownish to 

purplish, 7.3-12 mm long, 0.6-0.9 mm wide; lemma 3-nerved, 

aristate, the awn scabrous, -1 mm long, the rest of the surface 

glabrous; palea 6-9 mm long, acuminate to short-aristate, 

2-nerved, glabrous, purplish; anthers orange, 3-6 mm long. 

DISTRIBUTION,-Disjunct between northern Mesoamerica 

(Mexico, Belize, Guatemala, and Honduras) and eastern Brazil 

(Figure 4), growing in moist forests from sea level to 700 m 

elevation; also two collections from southern Peru. 

ADDITIONAL SPECIMENS SEEN.-BELIZE, BELIZE: Gracie 

Rock, 1.5 mi [2.5 km] S of mile 22 on Western Highway, Croat 

23894 (MO, US), Liesner and Dwyer 1511 (US); Gracie Rock, 

Sibun river, Gentle 1639 (US, WIS). COROZAL: San Antonio, 

Gentle 4759 (NY, US, WIS). EL CAYO: El Cayo, River bluffs, 

Bartletr 11475 (NY, US); 9.5 mi [15 km] S of Georgeville on 

road to Augustine, Croat 23478 (US); Belize-Cay0 road, high 

ridge, 41 mi [65 km] section, Gentle 9681 (US); vicinity of 

Cuevas, S of Millionario, Croat 23604 (MO); Valentin, Lundell 

6210 (NY, US); Rio Frio Cave, near Augustine, Spellman 

1586 (MO), Swallen 1580 (MO). TOLEDO: In high ridge, near 

Condemn Branch Hills, Gentle 5253 (US); Yellow flowers, in 

high ridge, near San Antonio, Gentle 5503 (US). DISTRICT 

UNKNOWN: Tower Hill Estate, Karling 56 (US); Roaring Creek, 

Lundell8729 (US). 

BRAZIL, ESPIRITO SANTO: Vargem Alta, Fruteira, Pereira 

2259 (RB, US); Reserva Florestal do Rio Doce, Sucre 8476 

(RB); without locality, Kuhlmunn 143 (US). GOIAS: Without 

locality, Gardner 3523 (US). PARANA: Road BR-2, RibeirBo 

do Cedro, Hatschbach 8719 (US); Fazenda no Rio Vermelho, 

18 km ao noroeste de Bela Vista, Rssmann 3842 (US); high 

bank of Rio Ligeiro, S of Rio Ivai, -15 km E of Sao Tome, 

Lindeman and Haas 914 (US); without locality, Dusen 4416 

(Us). PERNAMBUCO: Tapera, Pickel 2354 WS). RIO DE 

JANEIRO: Andarai, Freire 423 (US); Camino dos Macacos, 

Chase 9971 (F, MO, NY, US); Corcovado, Luetzelbwg 109 

(NY), Chase 8183 (F, MO, NY, US), 8185 (F, MO, US), 9749 

(F, US); Estrada Velha, Rio de Janeiro-Pelropolis, Unia o

Industria, Soderstrom and Sucre 1867a (CEPEC, US); 

Excelsior, Brade s.n., Dec. 2, 1928 (R, US); Fabrica das Chitas, 

Schwacke s.n. (US); Floresta da Covanca, Jacarepagua, Duarte 

5043 (US); Gavea, Parque da Cidade, Calderon 2010 (RB, 

US); Mata da Tijuca, Kennedy and Sucre 872 (US); Morin, 

Peuopolis, Chase 121 71 (US); Petropolis, Gaes 804 (R), 94 

(R); Pic0 da Tijuca, Chase 8487 (F, US); Bico do Papagaio, 

Emygdio 379 (US), Almeida and Laroche 1368 (RB), Almeida 

1537, 2069 (RB), Lutz 1759 (R, US); Rio de Janeiro, Alto da 

Boa Vista, Black 54-18122 (IAN); Serra do Corcovado, entre 

Vista Chinesa e Mesa do Emperador, Calderon 2013 (US), 

Serra do Corcovado, Sumare, Calderon 2021 (US) Estrada da 

Vista Chinesa, Sucre 2132 (US); Tijuca, Chase 12146 (US), 

Brade s.n., Dec. 2, 1928 (US), Kennedy and Sucre 872 (US); 

Tijuca, Bom Retiro, Rosa 80 (R); Tingua, Boa Esperanca,

Mello Filho 1125 (US); without locality, Beetle 190 (US), 

Glaziou 18640 (NY); without locality and collector, Jan. 21, 

23 

1886 (Us 2463835). SANTA CATARINA: Brusque, Azambuja, 

Calderon 2001 (US), Reitz and Klein 870 (NY, US); 

Blumenau, Ule 979 (US); Braco Joaquin, Luis Alves, Itajai, 

Klein 929 (US), Reitz and Klein 3342 (F, US); Guaramirim, 

Reitz and Klein 2395 (US); Ibirama, Luedenvald s.n., 1903 

(US); Horto Florestal I.N.P., Ibirama, Klein 949 (US), Reitz 

and Klein 1169 (US), Smith and Klein 7535 (US), 7537 (R, 

US); Isla de Santa Catarina, Gaudichaud 94 (US); Jaragua, 

Hoehne 24387 (US); Mato de Malucher, Brusque, Smith 5782 

(R, US); Morro do Bau, Lourteig 2359 (US); Morro do Bau, 

Ilhota, Reitz and Klein 18180 (US); Mono Costa da Lagoa, 

Klein 7001 (US); Morro do Ribeirao, Klein 6968, 6971 (US); 

Parque Estadual do Turvo, Tenente Portela, Klein 11885 (US); 

Paulo Lopes, costa do Morro de Paulo Lopes, Klein 9814 (US); 

Sabai, Vidal Ramos, Klein 2285 (NY), Reitz and Klein 4285 

(US); TresBarras, Garuva, S. Francisco do Sul, Reitz and Klein 

5560 (US). SAO PAULO: Alto da Serra, Usteri 9972 (US); Morro 

das Pedras, Brade 7859 (R, US); Parque do Estado, Hoehne 

28592 (NY, US); Piassaguera, Hoehne 7966 (US); Pindorama, 

Estacao Experimental, Carvalho 165 (US); 10 km S of centsr 

of city of Sao Paulo, Instituto de Botanica, Parque do Estado, 

Eiten 5592 (SP), Sendulsky 276 (US); Sao Paulo, Pickel 5880 

(US); Sao Paulo, Semi da Cantareira, Pickel 5894 (US), Coelho 

s.n., May 10, 1947 (US); without locality, Skvortzov 3 (US). 

STATE AND LOCALITY UNKNOWN: Riedel 935 (Us), Burchell 

3320 (US), Glaziou 505 (US). 

GUATEMALA, ALTA VERAPAZ: Montana Yxocubvain, 2l/2 

mi [4 km] W of Cubilquitz, Steyermark 44967 (F, US). PET~N: 

El Paso, Lundell 1574 (US); Hiltum, Lundell 3586 (US); La 

Libertad, Lundell 3375 (US, WIS); Tikal, Cook and Martin 

116 (US), in ramonal covering the ruins, Lundell15441,15808 

(US), ruina templo Las Inscnpciones, Molina 15774 (US); 

along road to Aguada Seco, Lundell 15689 (US); between 

Main Plaza and Aguada Comental, Contreras 3841 (US); 

Lake Yaxha, Gentry 8303 (MO); north shore of Lake Yaxha, 

Croat 24671 (MO); 2 mi [3.2 km] S of entrance of Tikal 

National Park, Croat 24712 (MO); NW of Jungle Inn, 

Contreras 3901 (US); in ramonal on Remate Road, Contreras 

293 (US); vicinity of archeological camp on N shore of Lake 

Yaxha, Croat 24671 (US); Uaxactum, Bartlett 12167 (NY, 

US). 

HONDURAS, CORTES: Calan, Dickson 1405 (US). 

MEXICO, CAMPECHE: Tuxpena, LundelllO61 (F, US, WIS); 

4 km S of Xpujil, Sheperd 32 (WIS). CHIAPAS: Hacienda San 

Pablo, near Champoton, Collins 58, 73 (US); Pixoyal, Ross 

s.n., 18 May 1954 (US); 19 mi [30 km] E of Zapata on the 

Zapata-Balancan road, Davidse 9425 (US); 17 km SE of 

Palenque, on the road to Bonampak, Davidse et al. 20395 

(MO). OAXACA: El Cerro de Nacimiento, E of Cosolapa, Vera 

Santos 2437 (US). QUINTANA ROO: Coba, Lundell 7694 (Us); 

9 km S of San Joe de la Montana,Davidse et al. 20268 (MO); 

8 km N of Union, 110 km SW of Chetumal, Davihe 20162 

(MO); 15 km N of Dzuich, Roe et al. 1340 (WIS); Tancah, 

Swallen 2825 (US). SAN LUIS POTOSI: Tamazunchale, Lundell 
24 SMITHSONIATV CONTRIBUTIONS TO BOTAW 

and Lundell 7238 (NY, US). TABASCO: La Palma, Balanan, 

Math 3244 (US); 5 km del rancho hnta de MontaiIa, 

C.Cowan 3375 (MO). YUCATAN: Chichen Ita, Bequuert 47 

(US), Steere 1511 (US), Swallen 1580 (NY, WIS), 2443 (US); 

Lake Chichankanab, Gaumer and Sons 23683 (US), Swallen 

2758 ll2 (US); Peto, Swallen 2720 (US). 

PERU, MADRE DE DIOS: Province of Manu, road from 

Salvaci6n to Shintuya, 600 m, Plowman and Davis 5085 (US). 

PASCO: Province of Oxapampa, Iscocacin, Palcazu Valley, trail 

to Villa America, Foster 9527 (MO). 

DIscUss1oN.-Mesoamerican plants have long been known 

as Olyra yucatana, but we can find no way to differentiate them 

from the Brazilian populations of 0. glaberrima. Olyra 

glaberrima was formerly known only from Atlantic Brazil, 

from Espirito Santo to Santa Catarina. Conspecifity with 0. 

yucatuna, which occurs from southern Mexico to Hondurus, 

leaves open the question as to how such a disjunct distribution 

came about. 

The Plowman and Davis 5085 collection from Peru may 

possibly represent a new species. It differs from typical 0. 

glaberrima in its more delicate habit with thinner leaves, 

papillose-pilose sheaths, sparsely pubescent pseudopetioles, 

awnless female glumes, and shorter (4 mm), awnless, male 

spikelets. However, female anthecia were not available and 

further study is required to determine whether this plant is 

indeed different from 0. glaberrima. Flowering in Mesoamerica 

is from December to July, and in South America from October 

to April. 

9. Olyra holttumiana Soderstrom & Zuloaga 

FIGURE 4 

Olyra holffumiana Soderstrom & Zuloaga, 1986:722. [Type: "Panama. Prov. 

Panama, La Eneida, regicn of Cerro Jefe," 3 March 1973, Dressler 4288. 

Holotype, US, sheet no. 2685261; isotypes, K, LE, MO.] 

Perennial. Culms arching, -2 m long, branching from the 

upper nodes; internodes cylindrical, hollow, glabrous; nodes 

compressed, purplish, glabrous. Leaves with sheaths stramineous with short-ciliate borders, otherwise glabrous, with 

conspicuous membranous auricles, the neck purplish black, 

short-pubescent; ligule membranous, conspicuous, 7-8 mm 

long, purplish black, short-ciliate above, shining and glabrous 

on the inner surface, short-pubescent on the outer; pseudopetiole 0.4-0.7 cm long, purplish, short-pubescent on both 

surfaces; blades oblong-lanceolate, acuminate, 24-35 cm long, 

6.8-9 cm wide, with an asymmetric base and apex, ascending, 

the upper surface greenish, the lower bluish green, both 

surfaces short-ciliate, the midnerve manifest on both surfaces, 

with scabrous borders. Panicles lax, 15-18 cm long, 20 cm 

wide, with the branches diverging from the axis, the lower 

ones verticillate, the upper alternate along the axis; axis 

longitudinally ridged, scabrous, the branches uiquetrous, 

flattened, hispid and scabrous toward the upper part, the base 

of the branches densely hispid and purplish black, the male 

spikelets in pairs and short-pedicellate, appressed to the 

flattened side, the pedicels thin, hispid, the female spikelets 

terminal, single; pedicels of the female spikelets thickened, 

short-hirsute. Axillary panicles similar to the terminal. Female 

spikelets fusiform, aristate, 35-50 mm long, 5.5-6.2 mm wide, 

greenish, the glumes unequal and longer than the anthecium; 

lower glume aristate, 15-nerved, the nerves tessellate, the inner 

surface hispid toward the apex, the outer surface scaberulous, 

the awn markedly hirsute; upper glume 11-nerved, the nerves 

tessellate with a pubescence similar to that of the lower glume; 

anthecium long-ellipsoid, acute, 14.5-15.8 mm long, 5.6-6 

mm wide, glabrous, smooth, shining, white; lemma 1 1-nerved; 

gynecium with a single style and 2 plumose stigmas; lodicules 

3, -1.8 mm long. Male spikelets lanceolate, 10.2-10.9 mm 

long, 0.8-1.2 mm wide, purplish, scabrous; lemma acuminate, 

7-9-nerved, scabrous on the outer surface; palea 8.2-9.7 mm 

long, 4-nerved, glabrous, purplish; stamens 3. 

DISTRIBUTION.-Known only from Panama (Figure 4). 

ADDITIONAL SPECIMEN SEEW.-PANAMA, PANAMA: Cerro 

Jefe, Nee 13970 (WIS). 

DIscUssI0N.-Olyra holttum'ana is distinguished by its 

very large female spikelets with long-awned, many-nerved 

glumes and male spikelets with an unusually large number of 

nerves in the lemma and palea. Although somewhat resembling 

0. fasciculata and 0. obliquifolia, the habit and smooth female 

anthecium of this species suggest that it is more closely related 

to 0. latifolia, 0. caudata, and 0. buchtienii. 

10. Olyra humilis Nees von Esenbeck 

FIGURFS 13, 15 

Olyra humilis Nees von Esenbeck, 1829:304. [Type: "Habitat ad ripas Taquari 

fluminis (Sellow. Vidi in Herb. Reg. Berol.)." Holotype, B, fragment of the 

holotype, US sheet no. 2877946.1 

Olyra hwnilis var. angustifolia Doell in Martius, 1877:321. [Type: "A cl. 

Riedel (n. 1938) in umbrosis siccis prope S. Carlos prov. S.Pauli lecta." 

Holotype, not seen, current location unknown; isotype, US sheet no. 

2877947.1 

Olyra hwnilis var. lafifolia Doell in Martius, 1877:321. [Type: Based on Olyra 

humilis Nees von Esenbeck, 1839:304.] 

Olyra glaberrima var. humilis (Nees von Esenbeck) Mez, 1912:28. [Type: 

Based on Olyra humilk Nees von Esenbeck, 1839:304.] 

Short- to long-rhizomatous perennial grass in cespitose 

clumps, with 10-20 culms in a fascicle. Culms geniculateascending or erect, many-noded, 25-100 cm tall; internodes 

short-hispid with whitish hairs to glabrous, hollow, cylindrical, 

the basal ones with a sheath only; nodes thickened, dark, 

glabrous. Leaves with sheaths pale with greenish spots, ribbed, 

glabrous or with papillose-pilose hairs, the margins ciliate; 

auricles absent; ligule membranous, short-ciliate at the tip, 

small, 0.4 mm long; pseudopetiole 0.2 cm long, stramineous 

to brownish, densely hispid, with long and whitish hairs; blades 

ovate-lanceolate !o lanceolate, 4.3-11.5 cm long, 1-2.2 cm 

wide, asymmetric and truncate at the base, symmetric at the 
NUMBER 69 25 

FIGURE 13.-Detail of female anthecium, Olyru humill: a, anthecium, dorsal side (x 30); b, apex of the 

anthecium, ventral side (x 50); c, base of the anthecium, ventral side (X 30); d, portion of the palea showing 

flattened macrohairs (x 200). All micrographs of Vulls et ul. 1804 (US). (Reduced to 77% of indicated 

magnifications for publication.) 

tip, scabrous on the adaxial surface to completely glabrous, the 

abaxial surface glaucous, the margins scabrous, the midnerve 

inconspicuous. Injl'orescences borne from the uppermost 

nodes, with 1-6 panicles at each node, the panicles lax, 

pyramidal, 4-9 cm long, 1-5.5 cm wide, few-flowered, the 

branches spreading, the lower ones whorled to alternate, the 

upper ones alternate, each branch with numerous male spikelets 

in pairs (one short- and the other long-pedicellate) below and 

one single terminal female spikelet; axis longitudinally ridged, 

scabrous, the branches triquetrous, scabrous; pedicels of the 

female spikelets thickened, smooth, glabrous. Axillary panicles 

similar to the terminal one. Female spikelets fusiform, aristate, 

14-21 mm long, 2.3-2.8 mm wide, both glumes shortly pilose 

to scabrous on the inner surface, and glabrous to scaberulous 

on the outer one: lower glume aristate, 5-7-nerved with 

transverse veinlets, the awn geniculate and variable in length, 

4-10 mm long; upper glume acuminate to shortly aristate, 

11-11.5 mm long, 5-7-nerved with transverse veinlets; 

anthecium long-ellipsoid, acute apically, 6-9.8 mm long, 

2.3-2.7 mm wide, whitish, smooth and shining, pilose; lemma 

5-nerved, pilose toward the apex and base on the ventral 

surface, the base covered by long, whitish or brownish, 

flattened unicellular macrohairs, the apex with flattened, 

unicellular macrohairs and prickle hairs, the dorsal surface of 

the lemma with sparse hairs toward the apex and base to 

glabrous: palea 2-nerved, with flat hairs toward the apex, 

otherwise glabrous. Caryopsis ellipsoid, brownish, 5 mm long, 

2.8 mm wide; hilum reaching the entire length of the caryopsis. 

Male spikelets fusiform, aristate, 8.5-11 mm long, 0.6-0.8 

mm wide, purplish, glabrous; lemma 3-nerved, aristate, the 
26 SMITHSOMANCONTRIBUTIONS TO BOTANY 

awn scabrous, 2-2.5 mm long, the rest of the surface glabrous; 

palea 5.5-7.8 mm long, 2-nerved; anthers 3,3.6-5.2 mm long. 

DISTRIBUTION.-Northwestern Argentina, Brazil, and Paraguay (Figure 15); from sea level to 1250 m elevation. 

ADDITIONAL SPECIMENS SEEN-ARGENTINA, CORRIENTES: Arroyo Chimiray y ruta 40, Quarin 3413 (US); 5 km E 

of Gdor. Virasoro, Schinini and Carnevali 10490 (US). 

MISIONES: Concepcion, Parodi 6944 (US); San Javier, Quarin 

3456 (US); Santa Ana, Montes 1181 (MO, US); Sierra Imh, 

Bertoni 1964 (US); 5 km ruta 101 desde Deseado a1 SW, 

Hunziker et al. 11028 (SI, US). 

BRAZIL, DISTRITO FEDERAL Acampamento do D.V.O., 

Brasilia, Belem and Mendes 60 (US); Fazenda Vargern Bonita, 

-10 km S of Brasilia, Irwin et al. 12287 (F, RB, US); Parque 

Nacional do Gama, Clayton 4959 (US); Parque Municipal do 

Gama, 25 km W of Brasilia, Irwin et al. 19480 (F, US); 3 km 

N of Sobradinho, Irwin et al. 10096 (MO, US); tributary of 

Rio Sobradinho, Clayton 4871 (US). GOIAS: Serra dos Pirentus, 

20 km E of Pirenopolis, Irwin et al. 34307 (MO). MINAS 

GERAES: Barreiros, Kuhlmann s.n., Feb 20, 1926 (RB, US); 

Caxambu, Sampaio 5946 (IAN, R); Chapeo de Sol, Serra do 

Cipo, Chase 9198 (F, US); Morro do Pilar, Black and 

Magalhaes 51-12125 (IAN, US); Sabara, Hoehne 6680 (IAN, 

R); Serra do Cipo, entre 0s km 130 e 132, Black et al. 51-11989 

(IAN); Serra do Cipo, km 103, Chuse 9286 (F, MO, US); Serra 

do Curral, Chase 8934 (F, MO, NY, US), 8965 (IAN, RB, 

US). PARANA: Florestal, 29 km E de Curitiba, estrada 

Curitiba-Paranagua, Tessmann 2786 (US); Papao Grande, 

Dusen 7752 (US); Ponta Grossa, Swallen 8455 (US); Ssio Josio 

do Triunfo, Hatschbach 14863 (US); Vila Velha, Hatschbach 

Klein et al. 916 (US); Colonia Sao Pedro, -8 km from the 

ocean, Soderstrom et al. 1991 (RB, US); Colonia Sao Pedro, 

estrada BR-101 a Colonia Sao Pedro, Valls et al. 1828 (US); 

Fazenda da Palma, Swallen 7016 (US); Fazenda Leitao, Rio 

Pardo, Jurgens GI 22 (US); Hamburger Berg, Lindman A485 

(US), Malme 190 (US); Morro da Policia, near P6rto Alegre, 

Rambo 37864 (US); Morro do Sabia, near Porto Alegre, Rambo 

43745 (US, WIS); Porto Alegre, Canisio 840 (US); Pareci 

Velho, prope Cai, Rambo 43790 (US); Rodovia Santa 

Rosa-Girua, 13 km apos S. Rosa, Valls et al. 1804 (US); Santa 

Barbara do Sul, estrada Carazinha-Cruz Alta, apos S. Marinho, 

Valls et al. 1676 (US); Sao Leopoldo, Dutra 551 (R), kite 

2081 (GH); Tres Cachoeiras, 6 km from ocean, Soderstrom et 

al. 1988 (CEPEC, US); Panambi, rodovia BR-285, km 300, 

Quarin 1163 (RB). SANTA CATARINA: Forest Seminirio 

Diocesiano, W of Chapeco, Smith and Reitz 12521 (US); W 

of Chapeco, on road to Guatambu, Smith and Reitz 12529 (F, 

MO, R, US); Pinheiral, Faxinal dos Guedes, Smith and Reitz 

9777 (US); Linha Coqueiro, by Rio Peperi-Guacu, Smith and 

Reitz 12662 (US); Passo d’Anta, S of Abelardo Luz, Smith and 

Reitz 12892 (R, US); Peperi, forest above Rio Peperi-Guacu, 

Smith and Reitz I2782 (R, US); Pinheiras by new airport E of 

Porto Uniao, Smith and Reitz 8848 (R, US); Ribeirao, Morro 

14666 (us). RIO GRANDE DO SUL: Chicuta Oliveira, Pelotas, 

da Tapera, Ilha de S. Catxina, Klein and Bresolin 8766 (US); 

Salete, Klein and Bresolin 11580 (US); Vargem Grande, Lauro 

Muller, Reitz and Klein 7233 (US); Vila Rica, by Rio do Peixe, 

Smith and Reitz 12930 (R, US). 

PARAGUAY: In regione fluminis Alto Parana, Fiebrig 

5436 (US); in altaplanicie Sierra de Amambay, Hassler 12051 

DIscUssI0N.-This species resembles 0. glaberrima, from 

which it differs by the size of plants, inflorescences, and 

panicles. The common name in Brazil is “bambu-fraco.” It 

flowers between October and May. 

(US). 

11. Olyra juruana Mez 

FIGURES 14, 15, 17e 

Olyra juruuna Mez, 1917:45. [Type: “Brasilia: Dept. Amazonas ad flumen 

Jurua prope Minas Geraes (Ule n. 5469)” Holotype. B, not seen, fragment 

of the holotype, US sheet no. 2877945.1 

Perennial, short-rhizomatous in cespitose clumps with 5-10 

culms. Culms simple, 35-1 10 cm tall, geniculate-ascending; 

internodes compressed, hollow, glabrous, the lower internodes 

with a sheath only; nodes compressed, dark brown and shortly 

hirsute. Leaves 5-7 per complement; sheaths strongly ribbed, 

hispid over the entire surface to ciliate toward the margins, 

with or without long hairs toward the upper portion; ligule 

membranous, 0.4-0.7 mm long, shortly ciliate at the apex, 

brownish; pseudopetiole 0.3-0.5 cm long, -0.2 cm wide, 

densely to sparsely pubescent with short, appressed hairs, 

brownish to purplish; blades ovate-lanceolate, 14-20 cm long, 

4.5-5.9 cm wide, narrowing from an asymmetric and truncate 

base to a symmetric or asymmetric, apiculate apex, glabrous, 

occasionally hispid toward the base on the adaxial surface, the 

midnerve prominent toward the base, the margins ciliate or 

not. Inflorescences paniculiform, 1-4 borne from the uppermost nodes or usually a single terminal panicle present, the 

peduncles hirsute; panicles pyramidal, 6-14 cm long, 1-4 crn 

wide, the branches alternate, with numerous male spikelets 

short-pedicelled on tertiary branchlets, 5 to numerous terminal 

female spikelets present on each panicle; axis and branches 

densely hispid to scabrous, longitudinally ridged; pedicels of 

the female spikelets thickened, smooth. Female spikelets 

fusiform, 13.5-18.5 mm long, 2.5-3.5 mm wide, whitish to 

stramineous, with the lower and upper glume subequal, 

subulate apically, scabrous toward the apex, otherwise 

glabrous; lower glume as long as the spikelet, 7-11-nerved 

with transverse veinlets; upper glume 12-16 rnm long, 

7-1 1-nerved with transverse veinlets; anthecium fusiform, 

8.8-10.7 mm long, 2.2-3.2 mm wide, whitish, acuminate 

apically, pilose all over the surface with long, appressed, 

cylindrical macrohairs; lemma 5-nerved, rugose at the upper 

margins; palea 2-nerved, excavated at the apex. Caryopsis 

long-ellipsoid, 6-6.5 mm long, 2 mm wide, brownish. Male 

spikelets fusiform, 4.2-6.7 mm long, 0.6-0.8 mm wide, 
NUMBER 69 27 

FIGURE 14.-Detail of female anthecium, Olyru juruunu: (I, apex of the lemma showing prickle hairs, long 

macrohairs, and excavations (x 200); b, anthecium, ventral side (x 200); c, upper portion of the lemma with 

rugose surface (x 1ooO); d, detail of macrohairs and prickle hairs (x 400). Micrograph u of Vurgus 15246 (US), 

6-d, of Schunke 19 (US). (Reduced to 77% of indicated magnifications for publication.) 

whitish to stramineous, hispid, occasionally glabrous; lemma 

?-nerved, subulate apically; palea 2-nerved. 

DIsTRIBUTION.-Growing in forests of eastern Peru and 

western Brazil (Acre); occasional as far east as Para (Figure 

15); found at 120-800 m elevation. 

ADDITIONAL SPECIMENS SEEN.-BRAZIL, ACRE: 52 km 

from Rio Branco on road to Sena Madureira, Calderdn and 

Soderstrom 2290 (US); 8’543, 72’51 W, Calderdn and 

Soderstrom 2352 (US); Porangaba, Rio Jurua-Mirim, Maas et 

al. PI2957 (US). PARA: Aguas Boas, Rio Pixuna, Kuhlmann 

1958 (RB, US). 

PERU, CUZCO: Entre Pilcopata y Keres, Vargas 15246, 

16353 (US). LORETO: Prov. Maynas, Explorama Tourist Camp, 

Rio Amazonas above mouth of Rio Napo, Gentry et al. 27478 

(MO, US), 28975 (MO, US), 37167 (US, MO); Gamitanacocha, Rio Mazan, Schunke 19 (F, NY, US). PASCO: Prov. 

Oxapampa, Iscozacin, Palcazu Valley, Foster 7968 (MO). 

DIscUssI0N.-Flowers between January and July. The 

Gentry et al. collections from northern Peru differ from typical 

material of 0. juruana in having pseudopetioles densely 

pubescent over the entire surface; (I-) 4-7 panicles borne on 

the uppermost nodes, these small (2-3 cm long) and 

short-exserted; female spikelets sparsely hispid on the outer 

surface; and male spikelets only 4.2 mm long. More collections 

are needed to determine if these plants represent a new species. 

12. Olyra latifolk Linnaeus 

RGLTES 8d-f, 16 

Olyru lutifolia Linnaeus, 1759: 1261. [Type: “Jamaica.” not seen, current 

Olyru puniculutu Swartz, 1788:21. [Type: “Incolit Jamaicae.” Holotype, S, 

location unknown.] 

not seen, current location unknown.] 
28 SMITHSONIANCONTRIBUTIONS TO BOTAhT 

FIGURE 15.-Distributim of Olyra amapana, 0. ciliatifolia, 0. hwnilis, 0. juruana, 0. loretensir, and 0. 

maranonensis. 

Olyra arundinacea IIumboldt, Bcmpland & Kunth, 1816:197. [Type: “Crescit Olyru pubescens Raddi, 1823:18. [Type: “In montibus ubique in Provincia 

in crepidinibus opacatis Andium Quinduensium pope El Azufral.” Rio-Janeiro.” Holotype, PI, not seen, fragment of the type, US sheet no. 

Holotype, P.] 2877930.1 

Olyra cordifolia Humboldt, Bonpland & Kunth. 1816: 198. [Type: “Crescit in Oiyra brevifolia Schumacher. 1827: 176. [Type: “Guinea, Africa.” Holotype, 

convalle frigida Bogotensi inter Santa Fe et Zipaquira.” Holotype, P. not not seen, current location unknown; fragment of the holotype. US sheet no. 

seen.] 2877961.1 
NUMBER 69 29 

Oiyra scabra Nees von Esenbeck, 1829:306. [Type: “ad Almadem prov. 

Bahiensis (Martius).” Holotype, M, not seen, fragment of the holotype, US 

sheet no. 2877928.1 

Olyra brasiliemb Desvaux, 183 1:210. [Type: “Ex Brasili.” Holotype, P, not 

seen.] 

Oiyra media Desvaux. 1831:lM. [Type: “Crescit in Brasilio.” Type, P, not 

seen.] 

Olyra latifoiia var. arundinaceu Grisebach, 18&4:535. [Type: Based on 0. 

arundinaceu Humboldt, Bonpland, and Kunth, 1816:197.] 

Oiyra latifolia var. glabriuscula Doell in Martius. 1877:316. [Type: Based on 

0. arundinacea Humboldt, Bonpland, and Kunth, 1816:197.] 

Olya iufifolia var. pubescens Doell in Martius, 1877:316. [Type: Based on 0. 

pubescem Raddi, 1823:lg.l 

Olyra cordifolia var. scubriuscula Dcell, in Martius, 1877:317. [Type: “A 

beato Martio in silvis provinciae Minarum lecta.” Holotype, M, not seen, 

fragment of the holotype, US sheet no. 2877954.1 

Olyra iafifoliu var. vesfifa Henrard in Amshoff and Henrard, 1948:321. [Type: 

“Suriname R., Berg en Dal (Focke n. 358).” Holotype, U.] 

Cespitose perennials, with the plants forming clumps of up 

to 20 culms from a hard, knotty crown. Culms 1-6 m tall, 

erect, climbing and leaning in shrubs, or decumbent, branching 

usually from the middle and upper nodes; internodes hollow, 

cylindrical, up to 1 cm in diameter, glabrous to shortly pilose, 

with or without retrorse hairs just above the nodes; nodes 

constricted, dark, shortly pilose with retrorse, appressed hairs, 

to glabrous. Leaves with sheaths covered by papillose-pilose 

hairs to glabrous, the margins short- to long-ciliate, the lower 

ones deciduous, the upper ones usually persistent; auricles 

present, membranous; ligules membranous, shortly pilose at 

the apex, 0.7-5 mm long, glabrous on the adaxial surface, 

shortly pilose on the abaxial; pseudopetiole dark brown, 

0.4-0.7 cm long, shortly pilose; blades lanceolate, 13-30 cm 

long, 2-10 cm wide, asymmetric and narrowed to subcordate 

at the base, symmetric to slightly asymmetric and acuminate 

apically, densely hispid over the entire surface to glabrous, the 

basal margins short-ciliate to glabrous, the midnerve prominent, hirsute or not toward the base, the lateral nerves 

anastomosing. Inflorescences lax and diffuse, pyramidal, 

12.5-21 cm long, 4-14 cm wide, with a ridged, hispid to 

glabrous peduncle, the branches divergent from the axis, the 

lower ones verticillate and only with male spikelets, the upper 

ones alternate and with male spikelets below and 1 to many 

terminal female spikelets; pedicels of the male spikelets thin, 

scabrous to densely hispid, those of the female spikelets 

thickened, purplish, short-pilose; aris longitudinally ridged, 

scabrous to densely hirsute, the branches uiquetrous, hirsute 

to scabrous, the axils of the branches densely pilose. Female 

spikelets broadly lanceolate, aristate, 14-25 mm long, 2.8-3.1 

mm wide, the glumes unequal, pubescent on the inner surface, 

the outer surface short-pilose, with the margins hirsute, to 

scabrous or glabrous; lower glum long-aristate, 5-1 l-nerved, 

with transverse veinlets; upper glum 9.4-12 mm long, 

short-aristate, 5-9-nerved, with transverse veinlets; anthecium 

ovate, obtuse to acute apically, 5.6-6.1 mm long, 2.6-3.1 

mm wide, whitish, olivaceous at maturity, smooth and shiny, 

with rounded excavations toward the upper margins of the 

palea and lower margins of the lemma; lemma with short 

prickle hairs on the lower margins. Caryopsis ellipsoid, 4-4.5 

mm long, 2.3-2.5 mm wide; hilum as long as the caryopsis; 

embryo */6 the length of the caryopsis. Male spikelets fusiform, 

aristate, greenish to purplish, 5.5-8.1 mm long, 0.7-0.9 mm 

wide; lemma aristate, 3-nerved, with strong nerves, glabrous 

on the inner surface, scabrous on the outer; palea 4.4-6.4 

mm long, glabrous, purplish, 2-nerved. 

DISTRIBUTION.-This is the most widespread and weedy of 

any species in the genus, and is found essentially throughout 

the Neotropics from Florida (not verified in recent times), the 

West Indies, and central Mexico (Nayarit and Tamaulipas) 

south to northern Argentina (Comentes and Misiones). Olyra 

latijiolia is found in forest margins, mostly below 1000 m 

elevation. The species has also become naturalized in Africa, 

including Madagascar. 

ADDITIONAL NEW WORLD SPECIMENS SEEN.- ARGENTINA, CORRIENTES: Desembocadura del arroyo Garape en el 

Rio Parane, 45 km E de Ituzaingo, Quarin et al. 2852 (US). 

MISIONES: Loreto, Mutinelli s.n., Jan 1940 (US). 

BELIZE, EL CAYO: Over Cohune River, Bartlett 11566 (US); 

Cohune RidgeLundell 6428 (NU, US); Hummingbird Highway, S of Belmopan, Gentry 8585 (US); Mountain Pine Ridge, 

San Agustin, Lundell 6755 (NY); Pine Ridge, Duck Run, 

Bartlett 11538 (US); Reed Broken Ridge, Gentle 9661 (US). 

STANN CREEK: Little Cocquericot, Belize River, Lundell3874 

(US); Maskall, Northern River, Gentle 1020 (NY, US, WIS); 

Maya Mounds, vicinity of Cockscomb Mountains, Shipp 531 

(NY); Stann Creek Valley, Big Eddy Ridge, Gentle 3406 (NY, 

US); Temash River, Schipp 1374 (NY). TOLEDO: Between 

Machaca and Camp 2, Gentle 6930 (US); between Orange 

Point and Moho River, Gentle 7683 (US); Edwards Road 

beyond Columbia, Gentle 7370 (US); Swasey Branch, Monkey 

River, Gentle 3983 (NY, US). 

BOLIVIA, BENI: Guayamerin, Krapovickas and Schinini 

35105 (US). LA PAZ: Charopampa, Buchtien 1155 (US); Mapiri, 

Rusby 209 (US); San Carlos, Buchtien 7 (US). PANDO: SW of 

Cobija on Rfo Naraneda, Sperling and King 6446 (US). SANTA 

CRUZ: Ichilo, Montero to Puerto Grether, Renvoize and Cope 

3948 (MO). 

BRAZIL, AMAPA: Around Porto Platon, Pires et al. 51024 

(US); Rio Amapari, frequent along road to Porto Terezinha, 

Cowan 38486 (RB). AMAZONAS: Santa Isabel, Rio Negro, 

Baldwin Jr. 3401 (US); Suomata, Tabatinga, Pires and Black 

837 (US). BAHIA: Aea do CEPEC, Huge 210 (CEPEC, US), 

Santos 3421 (RB); vicinity of Salvador, Chase 7862 (US); 

Brotas, Chase 7878 (US); ramal que liga a rodovia IlheusItabuna ao povoado de Japu, krn 1, Santos and Mattos Silva 

3223 (CEPEC); rodovia Olivenca-Maruim, 12 km S de 

Olivenca, margem do Rio Acuipe, Mattos Silva et al. 1503 

(CEPEC). CEARA: Serra do Baturie, Eugenio 281 (RB). 

ESPIRITO SANTO: Morro Born Retiro, Soderstrom and Sucre 

1958 (US); Reserva Florestal de Linhares, Soderstrom and 

Sucre 1885 (US); Jaticoba, Brade 18499 (RB). GOIAS: Estrada 
30 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

FIGURE 16.-Detail of female anthecium, Olyru lu!ifoliu: u, anthecium, dorsal side (x 30); b, anthecium, ventral 

side showing excavations on apex of palea (x SO); c, detail of prickle hairs on margin of the lemma and rounded 

excavations on the tip of the palea (X 500); d, base of the anthecium showing rounded papillae and excavations 

(X 1OOO). Micrograph u of Reeder & Reeder 2422 (US), b-d of Schunke I930 (US). (Reduced to 77% of indicated 

magnifications for publication.) 

Leonino-Mozarlindia, prox. a Leonino, Guimarcies 89 (US); 

between Annapolis and Goyaz, Chuse 11497 (US). MARANHAO: 

Carolina to San Antonio de Balsas, Swallen 4041 (RB, US). 

MATO GROSSO: Entre 17 de Febrero e Mingao, Kuhlmunn 

2676 (RB); 20 km S of Xavantina, Irwin et al. 17038 (RB); 

km 274 of Xavantina-Cachimbo road, Philcox et al. 3115 

(RB); km 264 of Xavantina-Cachimbo road, Philcox and 

Fereira 4234 (RB); 30 km ENE of Barra do Garcs, Anderson 

9778 (RB); Bandalta, Pereira et al. 467 (RB). MINAS GERAIS: 

Fazenda da Cachoeira, Municipio Tombos, Barreto 1646 (US); 

Fazenda da Tabunha, Dist. Ilheu, Mexia 4963 (WIS); without 

locality, Widgren 914 (US). PARA: Belem, Archer 7977, 8200 

(US), Black 54-16154 (US), Pires and Black 164 (RB, US); 

margem esquerda do Rio Sao Manoel, abaixo Cachoeira do 

Caldeirao, Pires 3855 (US); Castanhal, woods along Braganza 

railway, Goeldi 302 (US); colonia 3 de Outubro, Guedes 282 

(US). PARANA: Horto Florestal near Maringa, Lindemn and 

Haas 13233 (US); Morretes, Dombrowski 7269 (US); Parque 

Nacional Iguassu, Duarte 1878 (RB). PARAIBA: Preia (ESCOla), 

Vasconcellos 5859 (US). PERNAMBUCO: Tapera, Pickel I642 

(WIS); Bonito, Tenorio 67-247 (Us). RIO DE JANEIRO: COSmC 

Velho toward Corcovado, Soderstrom et al. 1857 (CEPEC, 

US); Alto da Boa Vista, Sucre 2112 (RB); Jardim Botanico, 

Sucre 2062, 2126 (RB, US); Campo Grande, Serra do 

Mendanha, Lima 67 (RB), Sucre et al. 6388 (US); Floresta dii 

Covanca, Jacarepagua, Duarte 5042 (RB); ItaipuaG6, Pic0 Alto 

Moirao, Andreata 163 (RB); Sumare, Sucre 6234 (RB). RIO 

GRANDE DO SUL: Torres, Tres Cachoeiras, Mattos 20 (US); 
NUMBER 69 31 

S teinkopf, pres Sao Leopoldo, Rambo 42283 (WIS). RONDONIA: 

Rio Jaruary, Kuhlmunn 1865 (RB); km 214-215 MadeiraMamore railroad near Abuna, Prance et al. 5973 (US); vicinity 

of Santa Barbara, 15 km E of km 117, Prance and Rams 

71 70 (US); 1 km N of Ribeirao, road Abuna to Guajara-Mirim, 

Prance et al. 6450 (US); track between Mutumparana and Rio 

Madeira, Prance et al. 54Yl (US). RORAIMA: Fernando Costa, 

Black51-13719 (US); Canto Galo, Rio Mucajai, Prance et al. 

3960 (Us). SANTA CATARIhA: s. Ludjero, Orth 3100 (Us). SAO 

PAULO: Loreto, Sampaio 15948 (US); vicinity of Sao Vicente, 

Smith 2082 (US). 

COLOMBIA, ATLANTICO: Los Rendales, near laguna de 

Sabalo, Dugand and Jaram'llo 2714 (US). ANTIOQUIA: 

Vicinity of Planta Providencia, valley Rio Anori between Dos 

Bocas and Anori, Denslow 2296 (MO). BOLIVAR: Island of 

Mompos, Lands of Loba, Curran 370 (US). CALDAS: Santa 

Cecilia, van Sneidern 5247 (US). CAQUETA: Rio Orteguaza, 

Woronow and Juzepczuk 6136 (US). CHOCO: Bahia Solano, 

near Ciudad Mutis, Killip and Garcia 33597 (US); Rio San 

Juan, Cuutrecusus 21382 (US). GUAJIRA: S of Carraipia, 

Haught 4276 (US). MAGDALENA: Rio Frio to San Andres de 

la Luna, Pitrier 1700 (US); Sierra de Perija, Grant 10898 (US, 

WIS); Santa Marta, Smith 2173 (US). META: 4 km NW of 

Villavicencio toward Bogota, Luteyn et al. 4799a (US). 

PUTUMAYO: Puerto Ospina, Schultes and Cabrera 18946 (US, 

WIS). SANTAmER DEL SUR: Vicinity of Puerto Berrio, Haught 

1809 (US). TOLIMA: Mariquita, Uribe 3324 (MO). VALLE DEL 

CAUCA: Cisneros, Dagua Valley, Killip 11481 (US); El 

Aguacate, Rio Yurumangui, Cuutrecasas 16147 (US). VAUPES: 

Yurupari, Allen 3282 (MO, US); Puerto Hevea, Grass1 10032 

COSTA RICA, ALAJUELA: Vicinity of Los Chiles, Rio Frio, 

Holm and Iltis 967 (US); La Catarata de San Pedro de San 

Ramon, Brenes 20579 (NY); Rio Naranjo, Tonduz 7641 (US); 

2 km W of La Marina, Pohl and Davidse 11266 (US). 

CARTAGO: Vicinity of Pejivalle, Standley and Valerio 47133 

(US); Turrialba, near Reventazon River, Calderon 2104,2105 

(US), Holm and Iltis 57 (NY), King 5347 (US), Pohl and 

Davidse 10827, 10836 (US). GUANACASTE: Bebedero, Brenes 

12586 (NY); between Bebedero and Paso Hondo, Brenes 

13110 (NY); Bebedero, Hacienda Catalina, Jimenez 719 (US); 

5 km N of Canas, Finca La Pacifica, Pohl and Davidse 10542 

(US); 1 km W of Canas Dukes, Pohl and Davidse 10575 (US); 

between Colonia Carmona y Buena Vista, Jim'eez 377 (US); 

along the Rio Corobici, about 4 km NW of Canas, Taylor 

18049 (US); 10 km S of La Cruz along the Carretera 

Interamericana, Pohl and Davidse 10600 (US); 6 km NE of 

Las Juntas, Pohl and Davidse 10962 (US); Naranjos Agrios, 

Standley and Valerio 46451 (US). HEREDIA: On road to Puerto 

Viejo, 20 mi [32 km] N of San Joe, Lathrop 5557 (US). 

LIMON: Near RiO Barbilla, Cufodonti 659 (US); Rio Hondo, 

Calderon 2114 (US); 4 km SW of Rio Hondo, Soderstrom et 

al. 1211, 1215 (US, MS); Rio Madre de Dios, Calderon 

2112 (US). PUNTARENAS: Finca Los Helechales, between 

(US). 

Buenos Aires and Cerro Pittier, Hatheway 1685 (US); 5 km 

NW of the Carretera Interamericana in Boruca area, Pohl and 

Davidse 11148 (US); Finca Volcan Angel, Schubert and 

Rogerson 770 (US); 5 km SW of Rincdn, Osa Peninsula, Pohl 

and Davidse 10715 (US); Santo Domingo de Golfo Dulce, 

Tonduz 10082 (US). SAN JOSE: 22 km E of Dominical, Pohl 

and Davidse 11233 (US); vicinity of El General, Skutch 4055, 

4237 (NY, US); San Jose, Hitchcock 8503 (US). 

CUBA, CAMAGUEY: Vicinity of La Gloria, Shafer 13 (US); 

Sierra Cubitas, Shafer 528 (US). HABANA: Caibarien, Ansovin 

387 (US); Junto, Baker and Wilson 584 (US); Loma de Coca, 

Ehn 1000 (US), 1217 (US); Anafe, Ekman 1040 (US). ISLA 

DE PINOS: Sierra San Juan del Mar, Britton et al. 14648 (US); 

Nueva Gerona, Curtiss 293 (US); Swetland Ranch, San 

Francisco de las Piedras, Killip and Swetland 41637 (US); La 

Ceiba region, Killip 43647 (US); Sierra Las Casas, Killip 

44181 (US); Managua, Palmer and Riley 1058 (US). 

MATANZAS: San Miguel de 10s Banos, Killip 13864 (US). 

ORIENTE: El Cristo, Ekman 1450 (US); Baracoa, Pollard et al. 

53 (US); Rio Matamoros S of Holguin, Shafer 1348 (US); Alto 

Cedro to Paso Estancia, Shafer 1630 (US); Piedra Gorda, 

Shafer 3488 (US); Coastal cliffs near Rio Yamuri, Shafer 

7840 (US); La Perla, Shafer 8537 (US). PINARDELRIO: Vinales, 

Killip 13533 (US); Sierra San Juan, above San Blas, Morton 

4140 (US); Santa Cruz to Rangel, Morton 4436 (US); El 

Guama, Palmer and Riley 21 6 (US); Limestone hills between 

Rio Cayaguateje and Sierra Guane, Shafer 10455 (US); vicinity 

of Sumidero, Shafer 13499 (US); Buenaventura, Wilson 9247 

(Us); Sierra de Anafe, Wilson I1582 (Us). SANTA CLARA: 

Limones, Jack 6234 (US), Pringle 70 (US); Banao, Leon 

3990 (US); Sancti Spiritus, Sierra de Gavilanes, Leon and 

Clement 6621 (US). PROVINCE UNKNOWN: Herradura, 16 Mar 

1906, Hitchcock s.n. (US); Marianao, hon 583 (US). 

DOMINICAN REPUBLIC, AZUAY: Without locality, Rose 

3986 (US). BARAHONA: Paradis, Abbott 1636 (US). DUARTE: 

Los Franceses, near Gaspar Hernandez, Jime'nez 2451 (US). LA 

VEGA: Cotuy, Abbott 737b (US); Piedra Blanca, Allard 13331 

(US), 13602 (US); Sierra Prieta, Jimenez 1169 (US); El Pino, 

Jimenez 4404 (US). PUERTO PLATA: La Rosa, Eggers 2466 

(US). SAMANA: Sanchez, Abbotr I45 (US), Rose 4343 (US); 

Pilon de Azucar, vicinity of Laguna, Abbott 429 (US). SANTO 

DOMINGO: Rio Ozama, Ekmun 12522 (US). SEIBO: San Pedro 

de Macons, Rose 4203 (US). TRUJILLO: Valdesia valley, Allard 

17315 (US); La Nassa, Faris 269 (US). 

ECUADOR, AZUAY: Balao, Eggers 14407, 14669 (US). 

BOLIVAR: 5 km W of Echeandia on road to Guaranda, Young 

11 (US). ESMERALDA: Environs of Lita, on the Ibarra-San 

Lorenzo railroad, Madison et al. 5049 (US). GUAYAS: Rio 

Guayas, west of Guayaquil, Hitchcock 19962 (US). LoS RIOS: 

Hacienda Clementina on Rio Pita, Asplund 5403 (US); Rio 

Palenque Biological Station, Dodson and McMahon 5032 

(US). MANABI: El Recreo, Eggers 15068 (US); near Guale, 

Haught 3065 (Us). NAPO-PASTAZA: Without locality, Asplund 

8917 (NY); environs of Limoncocha, Madison et al. 5446 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

(US). ORO: Between Santa Rosa and La Chorita, Hitchcock 

21126, 21134 (US); Portovelo, Holway and Holway 1002 

(US). SANTIAGO-ZAMORA: Rio Upano valley, Camp 1438 (US); 

hilis W of Rio Upano, along Rio Chupiangas, Prieto 12 (US). 

EL SALVADOR, CHALALTENANGO: along road to Chalaltenango, 3 km E of Hwy. 4, Pohl and Davidse I1879 (MO). 

CUSCATLAN: Hacienda Chanqueso sudl. Suchitoto, Bachufer 

and Lehmboden 2430 (MO). LA UNION: Vicinity of La Union, 

Standley 20810 (US). SAN SALVADOR: San Salvador, S. 

Calderon 491 (US), Hitchcock 8856 (US), Standley 23199 

(US); Volchn de Guazappa, Pohl and Gabel 13626 (MO). 

SANTA ANA: Santa Ana, Hitchcock 8842 (US); 2 km SE of 

Metapan, Pohl 12593 (MO). SAN VICENTE: Vicinity of San 

Vicente, Standley 21691 (US). SONSONATE: Vicinity of 

Sonsonate, Standley 22355 (US). PROVINCE UNKNOWN: Cerro 

de La Olla, near Chalchuapa, S. Calderon 1047 (US); San 

Andrts, Kovar 1002 (US); Santa Carlota, S.Calderon 941 (US). 

FRENCH GUIANA: Chemin des Emerillons, km 6, Lescure 

204 (US); Maroni, Melinon 121 (WIS). 

GUATEMALA, ALTA VERAPAZ: S of Cubilquitz, Steyermark 44545 (US); Cubilquitz, von Turckheim II 845, 4037, 

8619 (US), 8478 (NY); vicinity of Secanquim, Pittier 256 

(US). HUEHUETENANGO: Between Ixch and Rio Ixcan, Sierra 

de 10s Cuchumatanes, Steyermark 49332 (US). LZABAL 

Vicinity of Puerto Darrios, Standley 24813 (US); vicinity of 

Quirigua, Standley 24676 (US); N of Quirigua, Weatherwar 

84 (US); Sierra del Mico, between Los hates and Izabal, 

Kellerman 7346 (US). JUUTIAPA: 8 mi [13 km] SW of San 

Cristobal, Dunn et a[. 23274 (MO). LIVINGSTON: Livingston, 

von Turckheim II 1224 (US). PETEAN: Dolores, on iio Mopan 

trail, Contreras 3052 (US); La Libertad, Aguilar 47 (US, WIS), 

Lundell 3610 (US); Tikal National Park, Tikal, Contreras 

3725 (US), Lundell 15741 (US). RETALHULEU: Retalhuleu, 

Kellerman 6274 (US). SUCHITEPEQUEZ: Mazatenango, Heyde 

and Lux 6400 (US). 

GUYANA: Kanuku Mountains, near Moco-Moco village, 

Maas et al. 3921 (US); Kanuku Mountains, forest trail to Mt. 

Iraimakipang, Goodland and Maycock 471 (US). 

HAITI: Massif de la Selle, Riviere Froide, Ekmun 6622 

(US); Gonave Island, Eyerdam 127 (US); La Table, vicinity 

of Anse Galette, Gonave Island, Leonard 3241 (US); vicinity 

of Pikmi, Gonave Island, Leonard 5122 (US); vicinity 

Port-au-Prince, Holdridge 1868 (US), Leonard 5269a (US); 

vicinity Cap-Haitien, Leonard 5292 (US); vicinity Michel de 

l’Atalaye, Leonard 7273 (US), 8039 (US); vicinity of La 

Vallee, Tortue Island, Leonard and Leonard 11452 (US). 

HONDURAS, ATLANTIDA: Lancetilla Valley, Chickering 

191 (US); vicinity of La Ceiba, Yuncker et al. 8285 (NY, 

WIS), 8476 (NY), 8822 (NY); Puerto Sierra, Wilson 352 (NY, 

US); near Tela, Standley 52707, 52937, 54131 (US); 3 mi [5 

km] S of Tela, Miller 12677 (US). CHOLUTECA: 1 km W of El 

Corpus, Pohl and Davidse 12383 (MO). COLON: 4.5 mi [7 kml 

NE of Trujillo on road to Castilla, Saunders 588 (MO). 

COMAYAGUA: Esquias, 75 km E of Comayagua, Hernandez 

112 (MO); alrededores del lago Yojoa, Duenus 86 (MO). 

COPAN: Hacienda Ocoteseco, 20 km NE de Santa Rosa de 

Copan, Portillo I (MO); 4 mi [6.5 km] NE of Santa Rita, 

Molina and Molina 30834 (MO). CORTES: NE shore of Lake 

Yojoa, Williams and Williams 18741 (US); San Pedro Sula, 

matorrales of Rio Piedras, Molina 5227 (US); road San Pedro 

Sula-corks, near Choloma, Molina 6669 (US); Pimienta, Rio 

Ulua, Molina 5640 (US). EL PARAISO: Road to Danlii, woods 

along Rio San Francisco, Swallen 11245 (US). FRANCISCO 

MORAZAN: Rio Orilla, faldas del Cerro Majicaran, Molina 

I534 (MO). ISLAS DE LA BAHIA: Roatan Island, 4 km E of 

Coxenhole, Molina 20740 (US). OLANCHO: 5 km de Juticalpa, 

Pereira 44 (MO); 22 km ENE of Juticalpa, Pohl and Davidse 

12420 (MO). VALLE: Amapala, Valerio Rodriguez 3397 (US). 

PROVINCE UNKNOWN. Cuyamel, Carleton 445 (NY, US). 

JAMAICA: Ferry River, Harris 11320 (US); above Gordon 

Town, Harris 11346 (US), 11436 (US); Louis River, near 

Castleton, Harris 11537 (US); Troy, Harris 12571 (US); Bog 

Walk to Spanish Town, Hitchcock 9298 (US); Ewarton to 

Linstead, Hitchcock 9410 (US); near Ipswich, Hitchcock 9623 

(US); Bath, Maxon 2367 (US); near Ipswich, St. Elizabeth, 

Maxon and Killip 1522 (US); Stoney River Base camp, 

Portland, Morley and Whitefoord 680 (US). 

LEEWARD ISLANDS, ANTIGUA: McCarthy valley, Box 

77 (US). DOMINICA: Near Carholm Estate, Layou River 

Valley, Ernst 1943 (US); banks of Layou River across from 

Clarke Hall, Stern and Wasshausen 2391 (US); Clyde River 

Valley, SW of Melville Hall airfield on road to Governor 

Estate, Ernst 1037 (US); Layou River Valley, Clarke Hall 

Estate, Ernst I712 (US). GUADELOUPE: Without locality, 

Duss 3147 (US); Ste. Rose, Questel 851 (US); Petite 

Montagne, Questel 4097 (US); Roche Blanche, Stehle I787 

(US); Cote de Baillif, Stehle 5942 (US); Calumet, Stehle 6244 

(US). MARTINIQUE: Without locality, Hitchcock 16442 

(US); Montagne, Stehle 3731 (US). 

MEXICO, CHIAPAS: Escuintla, Matuda 341 (US); Finca 

Irlanda, Purpus 7401 (NY, US); N of Mapastepec, Sharp 

X-176 (US); near the junction of the Rio Perlas and Rio Jatate 

at San Quintin and near Laguna Miramar, Sohns I705 (US); 

Sierra Madre, Tateoka 1133 (US). GUERRERO: Carretera a El 

Paraiso, 21 km al NE de Atoyac, near Rio Santiago, Koch et 

al. 79303 (US). JALISCO: 12 km NE of Nacastillo, Iltis and Nee 

1536 (IBUG, US, WIS); 10-13 km SE of El Tuito, McVauglt 

25405 (US); 5 km W of Puerto Vallarta, Soreng 2186 (US). 

MICHOACAN: Aquila, Hinton et al. 16137 (NY, US); 32 km N 

of Playa Azul, King and Soderstrom 4924 (NY, US). NAYARIT: 

6-7 mi [-lo km] S of Las Varas, road to La Cucaracha, 

McVaugh 19192 (US); 1-1.5 mi [-2 km] above La Cucaracha, 

12-13 mi [-20 km] S of Las Varas, McVaugh 19218 (US); 

along the road to Miramar and Santa Cruz, 15 km SE of San 

Blas, Feddema 1006 (US); vicinity of San Blas, trail to 

Ensenada de Matanchen, Ferris 5366 (US); 5 mi [-8 km] E 

of San Blas, Reeder and Reeder 2421, 2422 (US). OAXACA: 

Vicinity of Cafetal Concordia, Morton and Makrinius 2700 
NUMBER 69 

(US); Chiltepec, Martinez-Caldero'n 45 (US); Huantla de 

Jimenez, Schulres 824 (US); 5 km N of Matias Romero, King 

799 (US, WIS); between Ojillan and Jalapa de Diaz, Contatti 

3764 (NY, US); San Pedro de Tepinapa, Santiago de Jocotepec, 

Vera Santos 3410 (US), 3411 (NY, US); Ubero, Williams 

9335 (Us). QUINTANA ROO: Coba, Lundell 7698 (Us). SAN 

LUIS POTOSI: Micos, Pringle 3795 (NY, US); between 

Narango-Platinito-Salto de Agua, Sohns 1412, 1413 (US); 

Tamasopo, Pennelll8013 (US). TABASCO: Arroyo de Tapijuhiya, 

Rovirosa 43 (US). TAMAULIPAS: Rancho El Cielito, valle del 

Rio Sabinas, McDonald 890 (MO). VERACRUZ: Misantla, 

Purpus 5976 (US); Barranca de Pancaya, Purpus s.n. (US); 

vicinity of Rio Tonto, Vera Santos 2270 (NY). YUCATAN: 

Chichen Itza, Swallen 2428, 2467 (US). 

NICARAGUA, BOACO: N slope of Cerro Mombacho, 

Stevens and Grijalva 14711 (MO); 1-2 km SE of Santa Maria, 

along Rio Fonseca, Nee et al. 28009 (MO). CARAZO: 13 km 

SW de Diriamba, Moreno 24314 (MO). CHINANDEGA: Ameya, 

Maon et a/. 7190 (NY, US). CHONTALES: Hacienda Veracruz, 

including Cerro La Batea and Cerro Los Charcos, Stevens 

22253 (MO), 23277 (MO). GRANADA: Low hills near Granada, 

Maxon et al. 7611 (NY, US); Finca Cutirre, NW de Volcan 

Mombacho, Sandino 1272 (MO); 5 km N de Nandaime, 

Moreno 9905 (MO). LEON: Quesalguague, Baker 2105 (US). 

MANAGUA: Vicinity of Managua, Garnier 81 7 (US); camino a 

Salamina, Hacienda Santa Cruz, Guzmdn and Castro I51 

(MO). MATAGALPA: 11 km NE de Muy Muy, Moreno 24431 

(MO); Quebrada Santa Cruz, between Puertas Viejas and 

Esquipulas, Stevens 22343 (MO). NUEVA SEGOVIA: 7 km NE 

de El Jicaro, Moreno 1672 (MO). RIO SAN JUAN: 5 km SE of 

Rio Oyate, Nee and Miller 27584 (MO). RIVAS: Isla de 

Ometepe, Moreno 22064 (MO), Robleto 961 (MO), 1349 

(MO). ZELAYA: Rio Escondido, Bahia de Bluefields, Molina 

1909 (US). PROVINCE UNKNOWN: Jinotepe, Hitchcock 8721 

(US); El Rodeo, Garnier 1518 (US). 

PANAMA, BOCAS DELTORO: Isla Colon, vicinity of Chiriqui 

Lagoon, von Wedel 1343 (US). CHIRIQUI: 2 mi [3.2 km] SW 

of Puerto Armuelles, Croat 22060~ (MO); San Bartolo Aniba, 

Croat 26697 (MO). COCE:: El Valle, Croat 25272 (MO). 

COLON: Vicinity of Ancon, Piper 5226 (US); Balboa, Standley 

25437 (US); Barro Colorado Island, Bailey and Bailey 17 (US), 

Calderon 2101 (US), Ebinger 145 (US), Standley 31335, 

41009 (US), Stimson 5288 (US); Chiva-Chiva Trail, Red Tank 

to Pueblo Nuevo, Piper 5229 (US), Maxon and Harvey 6580 

(US); Pedro Miguel to Culebra, Hitchcock 7919 (US); around 

Frijoles, Pittier 2686, 3759 (NY, US); hills N of Frijoles, 

Standley 27533 (US); Gatun, Hitchcock 7983 (US); Madden 

Dam, Dwyer and Elias 7486 (US); vicinity of Rio Cocoli, 

Road K-9, Stern et al. 310 (US); along the Rio Culebra, above 

Santa Isabel, Pittier 4151 (US); Rio Pedro Miguel, near East 

Paraiso, Standley 29937 (US); trail toward Chico, Steyermark 

and Allen 17458 (US); Portobello, Las Cruces Trail, Ebinger 

106, 114 (US, WIS); vicinity of Palenque, Pittier 4126 (NY, 

US), DARIEN: Vicinity of Campamento Buena Vista, Rio 

33 

Chucunaque above confluence with Rio Tuquesa, Stern et al. 

854 (US); near helipad at Hydro Camp on Rio Morti, Duke 

15418 (US); vicinity of Paya, Rio Paya, Stern et al. 207 (US). 

HERRERA: 10 km N of Las Minas on road to El Toro, Sytsma 

and D'Arcy 3242 (MO); 18 km W of Las Minas, N slope of 

Cerro Alto Higo, Hamme14224 (MO). PANAMA: Agricultural 

Experimental Station at Matias Hernandez, Pittier 6643 (US); 

Alhajuela, Dwyer 1038 (US), Pittier 2339 (NY, US), 3470 

(US); 3 km from Canitas toward the Bayano Dam, Calderdn 

and Dressler 2151, 2157 (US); Juan Diaz, Killip 4089 (US), 

Standley 30607 (US); vicinity of Panama, Hitchcock 8101 

(NY, US); Pedro Gonzalez, Panama Bay, Perlas Islands, Allen 

2596 (US); -2 mi [3.2 km] up Rio La Maestra, Kennedy 1183 

(US); about 3-4 mi [5-6 km] upstream on Rio Pasiga, 

Kennedy 1205 (US); near Rio Mato Puerco, Erlanson 185 

(US); San Jose Island, Pearl Archipelago, East Harbor, 

Erlanson 250 (NY, US), 305 (US), Johnston 100, 462 (US), 

Stimson 5316 (US). SAN BLAS: Cangandi, de Nevers and 

Herrera 5665 (MO). VERAGUAS: 4 km W of Colonia Penal, 

Antonio 2230 (WIS); vicinity of Santiago, Stern et al. 991 

(US). PROVINCE UNKNOWN: Vicinity of Bella Vista, Piper 

5290 (US); road to Comzal, Gervais 143 (US); Camino de Las 

Sabanas, Heriberto 194 (NY, US); near Old Fort Lorenzo, 

mouth of RIo Chagres, Piper 5949 (NY, US); vicinity of San 

FElix, eastern Chiriqui, Pittier 5206 (NY, US). 

PARAGUAY: Picada Chiriguelo, Sierra de Amambay, 

Rojas 6861 (US); Santa Teresa, Bertoni 1718 (US). 

PERU, AMAZONAS: Quebrada Huampami, Berlin 703 (US). 

AYACUCHO: RIo Apurimac Valley, near Kimpitiriki, Killip and 

Smith 22985 (US). CUZCO: Cordillera Vilcabamba, 15 km 

walking distance from RIo Apurimac and Hacienda Luisiana, 

Dudley 10055 (MO). HUaNUCO: Bosque Nacional de Iparia, 

RIo Pachitea, near campamento, Miel de Abeja, Schunke 1306, 

1364, 1930, 2919 (US); Tingo Maria, Asplund 12226 (US). 

JUNIN: E of Quimiri bridge, near La Merced, Killip and Smith 

24002 (US); Rio Paucartambo valley, near PerenC bridge, 

Killip and Smith 25330 (US). LORETO: Iquitos, Williams 3752 

(US); Rio Nanay, upland forest near Zamito, Plowman 2513 

(US); Santa Maria, about 25 km below Yurimaguas on Rio 

Huallaga, Allard 22484 (US); trocha Yurimaguas-Tarapoto, 

Ferreyra 4959 (US); Rio Ampiyacu, environs Puca urquillo, 

Davis et a/. 811 (Us). MADRE DE DIOS: Explorer's Inn, 30 km 

SW of Puerto Maldonado, Londono 15 (US). SAN MARTIN: 

Alto Rio Huallaga, William 5824 (US); Gamitanacocha, Rio 

Mazan, Schunke 210 (US); Lamas, Williams 6465 (US); lower 

Rio Huallaga, Williams 3802,4375 (US); San Roque, Williams 

7682 (US); Tarapoto, Williams 5491, 5873 (US); camino a1 

Pueblo Viejo de Tocache, Schunke 3777 (US). UCAYALI: 

PuCdlpa, Soukup 3060 (Us). DEPARTMENT UNKNOWN: La 

Merced, McBride 5521 (US). 

PUERTO RICO: Vicinity of Mayaguez, Chuse 61 75 (US), 

61 77 (US), Heller 4583 (US), Sintenis 138 (US), Soderstrom 

1802 (US); vicinity of Maricao, Chuse 6200 (US); NE of 

Arborito, Chase 6334 (US); vicinity of Vega Alta, Chase 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

6416 (US); vicinity of Quebradillas, Chase 6570 (US); between 

Guainabo road and Bayamon, vicinity of San Juan, Chase 

6645 (US); Sierra de Luquillo, slopes of El Yunque, Chase 

6732 (US); Toa Baja, McKee 10601 (US); Cayey, Sintenis 

2396 (US); Panuelas, in montibus Llano, Sintenis 4764 (US); 

Santa Marta Hills near San German, Stimson 1229 (US); Rio 

Piedras, Wetmore 172 (US). 

SURINAM: Kayser airstrip, 45 km above confluence with 

Lucie %vier, Irwin et al. 55986,57696 (US); 3 km S of Juliana 

Top, 12 km N of Lucie Rivier, Maguire et al. 54423, 54424, 

54441,54858 (US). 

TOBAGO: Scarborough, Broadway 4362 (US), Main Ridge, 

Cowan 1488 (US), 1528 (US), Hitchcock 10221 (US), 10264 

(US); Green Hill, Purseglove 6261 (US); Bacolet, Sandwith 

1629 (US). 

TRINIDAD: North Post to Maqueripe, Britton et al. 876 

(US); Caparo Woods, Broadway 4933 (US); Arima Valley, 

Cowan and Simmonds I195 (US); Tabaquite, Cowan and 

Foster 1340 (US), Soderstrom 1072 (US); Port-of-Spain, 

Hitchcock 9968 (US), 9981 (US), 10195 (US), 10326 (US), 

10327 (US); San Fernando, Hitchcock 10105 (US); Aripo 

Valley, beginning of trail to Aripo Caves, Kennedy et al. 

1705 (US); near Simla, Smith 10085 (US); St. Joseph 

Savannah, Soderstrom 1088 (US); Erin Savanna, Soderstrom 

1127 (US). 

UNITED STATES, FLORIDA: Tampa Bay [mixed with 

Pharus lappulaceus], Herb. Chapman (photograph at US of 

herbarium sheet from unknown source). 

VENEZUELA, AMAZONAS: between Rio Mawariuma and 

Rio Baria, Gentry and Stein 47240 (US). ANZOATEGUI: Cerro 

La Danta, NE of Bergantin, Steyermark 61101 (US). APURE: 

Departmento Paez, 25 km by car E of El Nuba, van der Werff 

and Gonzalez 4821 (MO). ARAGUA: Trail behind Biological 

Station at Rancho Grande, Luteyn and Luteyn 5182 (US); 

northern slope of Parque Nacional Rancho Grande, McClure 

21264 (US); near Maracay, Soderstrom 1359 (US). BOLIVAR: 

17 km W of Rio Caura along Hwy. betweem Caicara and 

Ciudad Bolivar, Davidse 4442 (WIS); Rio Reforma, Steyermark 88060 (US). CARABOBO: Guaremales, road from Puerto 

Cabello to San Felipe, Pittier 8923 (US). COJEDES: Rio Pao, 

Paraima, Trujillo 5435 (MO); entle Hacienda Mata Clara y El 

Baula, Aymard et al. 2303 (MO). DELTA AMACURO: 13 km ESE 

of town of Sierra Imata, Davidse and Gonzalez 16620 (MO). 

DISTRITO FEDERAL: Cerro Naiguaa, Steyermurk 92143 (Us). 

FALCON: Maparari, 11 km E de Churuguara, Wingfield 6509 

(MO). MERIDA: near Bolero, along Rio Onia, Steyermurk 

(WIS); Guinand Estate, Siquire Valley, Pittier 5941 (US). 

MONAGAS: Quiri Quiri to Caripito, Chase 12581 (Us). NUEVA 

ESPARTA: Juan Griego Trail, Isla Margarita, Johnston 314 (US). 

PORTUGUESA: 30 km W of Guanare by air, along Rio Tucupido, 

Liesner et al. 12638 (MO); 17 km NW of Tucupido, Davidse 

et al. 21469 (MO). SUCRE: Cerro Patao, N of Puerto Hierro, 

Steyermark and Agostini 91165 (US). TACHIRA: The Balcon, 

56708 (us). MIRANDA: 3 km sw of Aragaita, Davidse 4105 

vicinity Cristobal Colon, Broadway 445 (US). YARACUY 

Departmento San Felipe, Municipio Veroes, 5 km S de Bella 

Vista, Agostini et al. 795 (MO). ZULIA: Vicinity of Mene 

Grande, Pittier 10647 (US). STATE UNKNOWN: Eleanor Creek, 

lower Orinoco, Rusby and Squires 357 (US). 

VIRGIN ISLANDS, ST THOMAS: Eggers 289 (US), 673 

(US). TORTOLA: Britton and Shafer 835 (US). 

WINDWARD ISLANDS, GRENADA: Morne Delice, 

Beard 571 (US); Azimar Mountains, 18 Oct 1905, Broadway 

s.n. (US). SANTA LUCIA: Hitchcock 16357 (US), 16479 

(US). SAINT VINCENT: Kingshill, Beard 1371 (US). 

ADDITIONAL OLD WORLD SPECIMENS SEEN.-ANGOLA: 

Dundo, poximum flumen Luachimo, Gossweiler 14211 (US); 

River Luovwa, Moxico District, Milne-Redhead 41 76 (US). 

BURUNDI: Bururi, Rumonge, foret de Kigwena, Reekmans 

1519 (MO), 4849 (MO). 

CAMEROON: Mbalmayo, Jacques-Felix 9142 (US); Mimbia, Zenker 164 (US); Yaounde, Zeuillen 220 (US). 

CENTRAL AFRICAN REPUBLIC: Manovo-Gounda-St. 

Floris National Park, 0.5 mi [ 1 km] downstream of KoumbalaPende confluence, Fay 6061 (MO); Niao River 4 km S of 

Sakala Road, Fay 7412 (MO). 

COMORO ISLANDS: Mayottem Hachiroungou massif, 

Lorence and Saida 2861 (MO). 

CONGO: Paysoles M'brous, Chevalier 5766 (US). 

ETHIOPIA: 6 km SE of Tippi, 7"09", 35'232, Meyer 

GABON: Ndjole, Thollon 723 (US). 

GHANA: Ejura, Vigne 3454 (US); Kumasi, Yamoah 623 

GUINEA: Mt. Nimba, Adam 3155 (MO); Dendano, Adam 

3619 (MO); Nzerekore, Adam 5343 (MO); Bogbozou, Mum 

5564 (MO); Mt. Kouyo, Adam 5798 (MO); Singuedou, Adam 

6205; Kindia, Seguea, Adam 12635-1 (MO). 

IVORY COAST Several km SE of Zie-Noa bridge, near 

Kange Yanze, Fosberg 40485 (US). 

KENYA: Meru Dist., lower Imenti Forest, on MeruMikinduri road, Faden and Faden 78-894 (MO). 

LIBERIA: Nyaake (Webo), Web0 Dist., Baldwin 6086; 

Jabroke (Palipo), Web0 Dist., Baldwin 6698 (US); near 

Soplima, Vonjama Dist., Baldwin 10059. 

MADAGASCAR: Bassin de la Manampanihy, Humbert 

6042 (US). 

MALAWI: Lujeri Estate, Mlanje, Wiehe 714 (US). 

MOZAMBIQUE: Amatongas forest near Gondola, Fisher 

and Schweickerdt 271 (US); Garuso, Fisher and Schweickerdt 

288 (US); Tzengannedose, Gomes and Sousa 4375 (US); Vila 

de Joa0 Belo, Chipenhe, na floresta de Chirindzene, de Lemos 

and Balsinhas 49 (US). 

8026 (US). 

(US). 

NIGERIA: Abugi, Kabba Prov., Clayton 623 (US). 

SENEGAL: Tambacounda, Niokolo koba, Adam I7425 

SIERRA LEONE: Jala, 20 Jan 1927, Fairchild s.n. (US). 

SOUTH AFRICA: Ngoya, Zululand, 18 Apr 1895, Wylie 

(MO); Ziguinchor, Bayottes, Adam 13713 (MO). 
NUMBER 69 35 

sen. (US). 

TANZANIA: Magenga forest, Faulkner 1201 (US); Languga, Usambaras Dist., Greenway 424 (US); Amani, Hitchcock 

24509 (US); Moshi, Hitchcock 24555 (US). 

UGANDA: Musole forest, Barkley 3231 (US); Kyewaga 

forest, Busiro County, Dawkins 369 (US); Nipayo, Dummer 

667 (US); Budongo forest, Loveridge 168 (US). 

ZAIRE: Yambata, De Giorgi 7422 (US); Parc National 

Garamba, Terr. Dungu, Troupin 790 (US); Dima, Vanderyst 

5237 (US); Mokaba, Vanderyst 5600 (US). 

ZIMBABWE: Chirinda Forest, Crook 1088 (MO), Pardy 

and Armitage 26 (MO); Inyanga, Pungwe, Wild 5274 (MO). 

DISCUSSION.-Olyra latifolia is a common, coarse, scandent 

species of forest margins and secondary forests throughout the 

Neotropics. It is naturalized and widespread in tropical AfriThere are both diploid and tetraploid populations. Recorded 

chromosome counts include n = 22 (Davidse and Pohl, 1974, 

1978; Hunziker et al., 1982), 2n = -30 (Gould and Soderstrom, 

1967, as 0. cordifolia), 2n = 22 (Reeder, 1969, Gould and 

Soderstrom, 1970, Kammacher et al., 1973), n = 11 (Pohl and 

Davidse, 1971, Davidse and Pohl, 1972a), and n = 11, 22 

(Davidse and Pohl, 1972b, Olorode, 1975, Dujardin, 1978). 

13. Olyra latispicula Soderstrom & Zuloaga, new species 

FIGURES 11. 17c, 18, 19,20 

TYPE SPECIMLV-BRAZIL, BAHIA: Municipio Porto Seguro, 

Parque Nacional Monte Pascoal, located 14 km E of BR 101 

at a point 13 km N of Itamaraju, 320 m, 13 May 1976, 

Soderstrom, Russell, and Huge 2208 (holotype, CEPEC; 

isotype, US). 

Gramen perenne, rhizomatibus breviter, culmis in fasciculis 

densis, erectis -0.70 m longis; nodi incrassati, pilosi. Folia per 

complementum 10-15; laminae ovatae-lanceolatae, 9-1 1 cm 

longae, 2.2-3.8 cm latae, glabratae, basi asymmetricae valde, 

truncatae, apice symmetricae; pseudopetioli 0.3-0.4 cm longi, 

pilosi; ligulae membranaceae-ciliatae, -0.3-0.4 mm longae. 

Paniculae laxatae, parvae, 5-9 cm longae, 1-3 cm latae. 

Spiculae femineae lanceolatae, aristatae, 14.8-15.8 mm 

longae, 3.7 mm latae, virides, glumis inter se inaequalibus; 

gluma inferior aristata, 5-nervis, nervis tessellatis; gluma 

superior acuta, 6.6 mm longa, 5-nervis, nervis tessellatis; 

anthoecium late ovatum, acutum, 5.5 mm longum, 3.7 mm 

latum, pilosum; lemma 5-nerve, nervis tessellatis. Spiculae 

masculae lanceolatae, acuminatae, 4.3 mm longae, 0.7 mm 

latae, scabrae; lemma acuminatum, 3-nerve; palea 3.5 mm 

longa, 2-nervis. 

Short-rhizomatous perennial, 0.70 m tall. Culms borne 

closely together, geniculate-ascending, unbranched at the lower 

and median nodes; internodes compressed, hollow, shortpubescent with appressed and retrorse hairs; nodes thickened, 

purplish, glabrous. Leaves 10-15 per complement, drooping, 

rarely ascending in the upper portion of the plant; sheaths 

striate, greenish to purplish, short-pubescent to glabrescent; 

ligules membranous-ciliate, - 0.3-0.4 mm long; pseudopetiole 

0.3-0.4 cm long, brown, long- to short-pubescent with 

thickened, whitish hairs on both surfaces; blades ovatelanceolate, 9-1 1 cm long, 2.2-3.8 cm wide, the upper leaves 

becoming smaller, strongly asymmetric and truncate at the 

base, with a symmetric, acuminate tip, glabrous on both 

surfaces, the margins ciliate to scabrous, with or without short 

hairs toward the base, the midnerve prominent. Inflorescence 

paniculiform, 2 or 3 borne at the upper nodes of the culms: 

peduncles long, -8-10 cm long, with short, retrorse hairs; 

panicles few-flowered, 5-9 cm long, 1-3 cm wide, the 

branches divergent from the axis, the lowermost whorled, 

occasionally alternate, the upper ones alternate, the lower 

branches bearing male spikelets only, with the spikelets paired, 

one short- and the other long-pedicelled, the upper branches 

with male spikelets at the base and a terminal female spikelet 

or with female spikelets only; axis longitudinally ridged, hispid 

on the lower part, otherwise glabrous, pedicels of the male 

spikelets purplish, scabrous, those of the female spikelets 

thickened, smooth and glabrous. Axillary panicles similar to 

the terminal one but smaller. Female spikelets lanceolate, 

aristate, 14.8-15.8 mm long, 3.7 mm wide, greenish with 

unequal glumes; lower glume long-aristate, scaberulous, 

5-nerved with transverse veinlets; upper glume acute, 6.6 mm 

long, glabrous, 5-nerved with transverse veinlets; a short stipe 

present at the base of the anthecium; anthecium broadly ovoid, 

strongly compressed, acute apically, 5.5 mm long, 3.7 mm 

wide, with conspicuous, caducous, long prickle hairs over the 

entire surface; lemma gibbous, depressed on the dorsal face 

along either side of the midnerve, 5-nerved with transverse 

veinlets. Male spikelets lanceolate, acuminate, 4.3 mm long, 

0.7 mm wide, reddish, scabrous; lemma acuminate, 3-nerved 

with the midnerve scabrous toward the apex; palea 3.5 mm 

long, 2-nerved, scabrous toward the apex; anthers 2.2 mm long. 

DISTRIBUTION.-Known only from the type locality, a 

lowland wet forest on the slopes of Monte Pascoal in southern 

Bahia, Brazil (Figure 11). 

DISCUSSION.-Olyra latispicula is a very distinctive new 

species, unusual in its broad, slightly apiculate female 

anthecium, with the lemma gibbous and the whole surface 

covered by conspicuous prickle hairs. The plants are delicate 

and small, with asymmetric leaves that have truncate bases. Its 

affinities with other species in the genus are not obvious. 

14. Olyra longifolia Humboldt, Bonpland & Kunth 

FIGURES 21, 22,23 

Olyra longifolio Humbldt, Bonpland & Kunth, 1816:198. [Type: “Crescit in 

humidis Guyanae ad ripam fluminis Guaviare, pmpe San Fernando de 

Atabapo, in sylvis Orinocensibus prop Santa Barbara.” Holotype P, not 

seen, fragment of the holotype, US sheet no. 2877943.1 

Olyra swinamensis Hochstetter ex Steudel, 1853:36. [Type: “Hrbr. Dr. 
36 SMITHSONIANCONTRIBUTIOXS TO BOTAhT 

FIGURE 17.-Field photos of Olyru species. 0.filiformk: u, detail of panicles; b, habit of plant (Bahia, Brazil, 

1972). 0. hfkpiculu: c, habit of plant (Bahia, Brazil). 0. micrunfhu: d, habit of plant (Brazil, 1968);J detail of 

panicles. 0. juruunu: e, habit of plant (Brazil, 1976). Photographs by C.E. Calderh, except c by TR. Scderstrom. 

Hostmunn N. 863. Surinam.” Holotype, P, not seen, fragment of the 

holotype, US sheet no. 2877922; isotype, MO.] 

Ofyru longifolia var. grundifoliu Doell in Martius, 1877:325. [Type: Based 

on 0. longifolia Humboldt, Bonpland & Kunth, 1816:198.] 

Olyra longifolia var. purvifolia Doell in Martius, 1877:325. [Type: “In vicinis 

provinciae Manaos do Alto Amamas a cl. R. Spruce lecta (n. 880 et 1145, 

nomine Olyrae acutincdae in ejus herbario signata).” syntype Spruce 880 

not seen; syntypes of Spruce 1145, NY. P, US. fragment of syntype, 1145, 

US sheet no. 84571.1 

Ofyru kegefii Mez, 1921:6. [Type: “Holl. Guyana (Kegef n. ZSI).” Holotype, 

B, fragment of the holotype, US.] 

Perennial in cespitose clumps, short-rhizomatous. Culms 

0.30-5 m tall, simple below, branching at the upper nodes, 
FIGURE 18.--Olyra latirpicula: a, habit (X 0.5); b, inflorescences from upper nodes (x 0.5); c, pseudopetiole 

and ligule (x 5). Based on Soderstrom et al. 2208 (US). Illustration by A.R. Tangerini. 

NUMBER 69 37 
38 SMITHSONIANCONTRIBUTIONS TO BOTANY 
KUMBER 69 39 

FIGURE 20.-Detail of female anthecium, Olyra latirpicula: a, apex of the anthecium, ventral side (X 50); 6, 

detail of the surface showing caducous prickle hairs (x 200); c, base of the anthecium, ventral side (X 50); d, 

prickle hairs (X 1OOO). All micrographs of Soderstrom et 01. 2208 (US). (Reduced to 77% of indicated 

magnifications for publication.) 

erect or rarely clambering; internodes cylindrical, hollow, 

shiny, glabrous; nodes thickened, short-pubescent with whitish 

hairs. Leaves with sheaths strongly ribbed, densely pubescent 

toward the summit, the rest of the surfaces short- to 

long-pubescent to glabrous; auricles absent; ligules membranousFIGURE 19.41yra latirpicula: a, habit and inflorescence (x 4); b, female 

spikelet (X 8); c, lower glume of female spikelet (x 8); d, upper glume of 

female spikelet (x 8); e, female anthecium, palea side (x 8);J lemma of female 

spikelet, inside view (X 8); g, lemma of female spikelet, outside view (x 8); 

h, palea of female spikelet, outside view (x 8); i, palea of female spikelet, inside 

view (X 8); j, same as i, apex opened (x 8); k, gynecium (x 8); I, male spikelet 

(x 12); m, lemma of male spikelet, inside view (x 12); n, palea of male spikelet, 

with stamens enclosed (X 12); 0, palea of male spikelet, inside view (x 12): p, 

andrecium and lcdicules (X 16.5); q, lodicules of male spikelet (x 24). All 

components based on Soderstrom et al. 2208 (US). illustration by A.R. 

Tangerini. 

ciliate, -0.6 mm long, brownish, short-pubescent on the abaxial 

surface; pseudopetiole 0.2-0.5 cm long, pale to purplish, 

pubescent with short, appressed hairs to glabrescent; blades 

lanceolate, 5.5-21 cm long, 1.2-5.7 cm wide, ascending, with 

a subcordate to narrowed and symmetric base, the apex either 

symmetric or asymmetric, glabrous on the adaxial surface, 

glaucescent and hispid on the abaxial one, the upper margins 

ciliate, the midnerve prominent. Inflorescencex racemiform, 

usually with 2 to many panicles borne from the uppermost 

nodes, all the panicles almost reaching the same height, the 

peduncles short-pubescent; panicles 3.5-7.5 cm long, 1-2 cm 

wide, with male and female spikelets borne in different 

branches, usually with 2-5 branches with male spikelets at the 

base, one of these branches longer than the other ones; pedicels 

of the male spikelets slender, scabrous; female spikelets 4-12, 

disposed alternately on one branch; pedicels of the female 
40 SMITHSOMAN CONTRIBUTIONS TO BOTAh?' 

FIGURE 21.-Detail of female anthecium, Olyra longifolia: a, base of the anthecium. ventral side (x 70); b, 

detail of the papillae at the base of the anthecium (x ZOO); c, papillae (X 1OOO); d, apex of the palea showing 

rounded excavations and unicellular macrohairs (x 500). Micrographs u-c of Black48-2447 (US), d of Schultes 

& Cabrera 15386 (US). (Reduced to 77% of indicated magnifications for publication.) 

spikelets thickened; axis of the branches scabrous to sparsely 

pubescent. Female spikelets lanceolate, aristate, 17-22 mm 

long, 2.4-3 mm wide, greenish to purplish, the glumes 

subequal, scabrous, anthecium '/2 or less the length of the 

spikelet: lower glume short-aristate, 7-13-nerved with transverse veinlets, acuminate, glabrous, scabrous toward the apex 

of the inner surface: upper glume 15-18 mm long, 7-13- 

nerved with transverse veinlets, acuminate, glabrous; anthecium obovoid, 5.2-6.6 mm long, 1.9-2.4 mm wide, narrowed 

basally, acute apically, pale, with rounded to irregular 

excavations spaced throughout the surface and thickened, 

caducous, fragile and unicellular hairs toward the apex: lemma 

7-nerved, palea 4-nerved. Caryopsis obovoid, brownish, 

3.8-4.2 mm long, 2 mm wide: hilum as long as the caryopsis: 

embryo l/6 the length of the caryopsis. Male spikelets 

lanceolate, short-aristate, 5.3-7.3 mm long, 0.6-0.8 mm wide, 

scabrous, pale to purplish: lemma 3-nerved, short-aristate: 

palea 2-nerved; anthers 3,2-3 mm long. 

DISTRIBUTION.-Northern South America from Colombia 

to Bolivia and Brazil, from sea level to 400 m elevation: 

especially common in the Guianas (Figure 23). 

ADDITIONAL SPECIMENS SEEN.- BOLIVIA, BENI: 2 km 

NW of Guayamerin, Anderson et al. 12119 (MO, US). 

BRAZIL, AMAPA: Araguari River, Pires et a/. 50581 (NY); 

Oiapoque, Froes 25743 (IAN). AMAZONAS: Cahoeira Caranguejo, River Cuabury, Holt and Blake 422 (US); Cucuhy, Rio 

Negro, Holt and Gehriger 356 (NY, RB, US), 399 (US); near 

Tunui, Icana River, Froes 28411 (IAN); Barcelos, Maguire et 

al. 60041 (US); Rio Negro, -120 km above Barcelos, Madison 

et al. 6181 (US); between Moreira and Rio Quinini, Prance et 
NUMBER 69 41 

FIGURE 22.-Detail of female anthecium, Olyra longifolia: a, apex of the anthecium, dorsal side (x 50); b, apex 

of the anthecium, ventral side (X 50); c, detail of the palea (x 200); d, anthecium, ventral side (x 50). Micrographs 

a-c of Sfeyermark 512431 (US), d of Schultes & Cabrerra I5386 (US). (Reduced to 77% of indicated 

magnifications for publication.) 

al. 16253 0; Camunde, Rio Negro, Baldwin 3396 (US); 

Negro River, Ilha do Piranade, above Barcelos, Baldwin 3297 

(RB, US); Negro River, Ilha do Matupiri, Froes 28261 (US); 

Parana de Sao Jose de Ariraha, Baldwin 3298 (US); Parawa, 

below Sao Felipe, Rio Negro, Baldwin 3572 (US); Tapuruquara, Rio Negro, Black 48-2477 (US), Pires 274 (NY); Rio 

Negro, between Moreira and Rio Quinini, Prance et al. 16253 

(US); Rio Negro, between Ilha Jacari and Airao, Prance et al. 

15098 (US); basin of Rio Demeni, vicinity of Tototobi, Prance 

et al. 1021 7 (US); regiao do Rio Jari, Estrada do Munguba, 

Silva 1642 (US). MARANHAO: Pindare River, Moncao, Froes 

20328 (US). PARA: Belem, Soderstrom 1193 (US); Bon 

Jardim-Itaituba, Swallen 6943 (US); Braganca railroad, Goeldi 

79 (RB, US); Ilha do Rocado, 4 km NW de Cachoeira Porteira, 

Marfinelli 7130 (MO); Moju River, Estate Conceicao, Goeldi 

150, 15I1/z (US); Moju Fabrica, Kuhlmann 2128 (R, RB, US), 

Black 54-16211 (US), 54-16319 (IPEAN); Gurupa, Igarape

Jacupi, Pires and Silva 4690 (US); River Cumina, Cuminamiri, Silva 1265 (US); 1 km N of Fazenda Urucurituba, on 

Tapajos River, Archer 8397 (US); Paragominas, River Uraim, 

Silva 404 (US); Piratininga, Almeirim, Duck s.n., April 16, 

1923 (RB, US); Serubin, Itacaiuna River, Froes and Black 

24540 (US). RORAIMA: Baixo Rio Branco, Froes 23011 (NY, 

US); Posto Mucajai, Rio Mucajai, Prance et al. 11074 (US). 

COLOMBIA, BOYACA: Orocue, Haught 2821 (F, NY, US). 

cHocO: Quebrada La Sierpe, Forero et al. 3915 (MO). EL 

VALLE: Veneral, Yurumangui River, Cuatrecasas 15873 (F, 

US); Quebrada del Corosal, Cajambre River, Cuatrecasas 

I7751 (F, US). VAUPES: Cachivera de Jirijirimo, Apaporis 

River, Schultes and Cabrera 12424 (US); between the rivers 
42 

FIGURE 23.-Distribution of Olyra fongifofia. 

SMITHSO?JIANCONTRIBUTIONS TO BOTAhT 

Pacoa and Kananari, Apaporis River, Schultes and Cabrera 

13817 (US); Raudal Yayacopi and vicinity, Schultes and 

Cabrera 15386 (US); Ininda River, San Joaquin, Fernandez 

1999 (F, US); Mitu, Arbelaer and Cuatrecasas 6784 (US); 

Pacoa River, tributary of Rio Apaporis, Schultes and Cabrera 

15259 (US); selvas del Tui-Igarape, Cuatrecasas 6869 (US). 

FRENCH GUIANA: Acarouany, Melinon 231 (US, WIS); 

Haut Itany, Hoock s.n., 1 Jan 1956 (P); Itany, Tolinga, Hoocli 

s.n., 30 Dec 1955 (P). 

GUYANA: Akwero, Maas 2409 (US); Bonisika Landing, 
NUMBER 69 43 

Arawau River, Archer 2336 (US); Bunsika Ck., Aruwau Ck., 

without collector, March 28, 1945 (NY); Cuyuni River, Tufin 

970 (US); Essequibo River, near mouth of Mazaruni River, 

vicinity of Penal Settlement, Hitchcock 17241 (US); SE of 

Georgetown, East Coast Water Conservancy, Hitchcock 16979 

(US); vicinity of Issorora, on Aruka River, Hitchcock 17577 

(US); Kamakusa, Leng 16 (NY); Kaieteur Fall, left bank of 

Potaro River, Cowan and Sodersrrom 2001 (F, US); 3 mi [5 

km] above Kaieteur Fall, Cowan and Soderstrom 2108 (US); 

Kanuku Mountains, in drainage of Moku-moku Creek, A.C. 

Smith 3410 (F, MO, NY, US); Mazaruni Station, Turin 138 

(RB, US); Moruka River, Santa Rosa, De la Cruz 998 (NY, 

US); Pomeroon River, De la Cruz 3167 (F, MO, NY, US); 

Potaro River, Jenman 7528 (US), Sandwith 1395 (US): 

Rockstone, on the Essequibo River, Hitchcock 17332 (NY, 

US); vicinity of Tumatumari on Potaro River, Hitchcock I7363 

(F, MO, NY, US); Wismar, on the Demerara River, Hitchcock 

17423 (US): banks of Yawakuni River, Abraham 189 (US); 

without locality, Jenman 5963 (NY, US), 6737 (US). 

PERU, LORETO: Rio Nanay, lowland forest near River at 

Shiriara, Plowman 2554 (US); Timbuchi, on the Rio Nanay, 

Williams 1043 (F, US). 

SURINAM: Kayser airstrip, 25 km above confluence with 

Lucie Rivier, Maguire et al. 53952 (US); Kayser airstrip on 

the Zuid-Rivier, Kramer and Hekking 2978 (NY); Nickerie 

River, Kamisa falls, Maas 1966 (US); Wayombo River, near 

the mouth, Lindeman 6322 (US); N of Nickerie River, Lanjouw 

and Lindeman 3404 (NY); without locality, Hostmann 97 (US). 

VENEZUELA, AMAZONAS: Along Orinoco River, at mouth 

of Rio Sanariapo, below Sanariapo, Steyermark 58431 (F, 

US); Cunucunuma River, along river 0.5 mi [l km] above 

Culebra Rapids, Maguire et al. 30393 (NY, US); near Santa 

Barbara del Orinoco, Steyermark 117163~ (MO); vicinity of 

Rio Coro-Coro, near airport of Yutaje, Liesner ef al. 10980 

(MO); 5 km S of San Carlos de Rio Negro, Liesner 6205 

(MO); W side of Isla Sebastian, Rio Casiquiare, above 

Chapezon, Liesner and Clark 8890 (MO); Cafio Jotajana, 

tributary of Cano Guiniquina, Steyermark 115125 (MO); 

Ibaruma, Tamayo 3617 (F, MO); upstream from San Victor, 

past Quebrada Piedradero and Rio Matanaima, Steyermark 

87188 (NY): between La Margarita and Puerta Miranda, Acure 

River, Steyermark 87721 (US); 5-14 km ESE of Los Castillos 

de Guayana, Davidse and Gonzalez 16275 (MO). APURE: 4.5 

airline mi [7 kml ESE of San Carlos del Meta along the banks 

of the Rio Meta, Davidse and Gonzalez 13828 (MO); E of the 

southern tip of the Galeras de Cinaruco, Davidse and Gonzalez 

14616 (MO), 15640 (MO, US); near the Rio Meta at Fundo 

El Algarrobo, Morichal de la Madera, Davidse and Gonzalez 

14200 (MO). BOLIVAR: Rainforest along Rio Apacara,

Apacara-tepui, Steyermark 74642 (F, NY, US); Sierra Imataca, 

Rio Toro, between Rio La Reforma and Puerto Rico, 

Steyermark 87837 (NY); Ichun River, below the Salto Maria 

Espuma, Steyermark 90463 (NY, US); near Campamento Las 

Pavas, Steyermark I17199 (MO); Paragua River, between La 

Paragua and Salto de Auraima, Killip 3 7317 (US); Torono 

River, Indian camp above junction with Paragua River, Killip 

37424 (US); vicinity of Los Patos, Steyermark 86996 (US): 

Guaiquinima, dense forest east of River Paragua toward Cerro 

Guaiquinima, Killip 37481 (US); SE of Canaima, Steyermark 

Epana and Cano Araguabisi, Steyermark et al. 115125 (WIS). 

GUARICO: Parmana, Tamayo 4030 (F, MO, US). 

DIscUss1oN.-This is a variable species, with great variation 

in the size of plants and panicles. 

Olyra longifolia is characterized by having the female and 

male spikelets in conjugate branches of the same panicle, these 

being contracted and borne on the uppermost nodes of the 

culms. In this character the species parallels the inflorescence 

structure of the olyroid Cuban herbaceous bamboo genera 

Ekmanochloa Hitchcock and Mniochloa Chase. The obovoid 

anthecium is greatly reduced in relation to the size of the 

spikelet and has rounded excavations and long, stiff, caducous 

hairs over the entire surface (more so toward the apex). 

Flowering occurs from December to June. 

Clayton and Renvoize (1986) cite this species as a link 

between Olyra and Cryptochloa but it should be pointed out 

that there is no stipe at the base of the female anthecium as in 

the latter genus. Cryprochloa also differs in its usual 

arrangement of female racemose inflorescences from the lower 

nodes and female paniculate inflorescences from upper nodes 

of the culms. 

The common name is recorded as “taboquinha” in Brazil. 

106401 (MO). DELTA AMACURO: Cano Jotajana, between 

15. Olyra loretensis Mez 

FIGURES 15.24.25 

Olyru lorefemis Mez, 1917:47. [Type: “Peruvia, Dept. Loreto, prope Leticia 

in silvis (Ule n. 6224):’ Holotype, B, fragment of the holotype, US sheet 

no. 2877942.1 

Cespitose perennials from creeping rootstocks and forming 

dense clumps. Culms 30-60 cm tall, geniculate-ascending, 

cylindrical, hollow, short-pubescent, biform: leafy culms 

bearing leaves only, with the leaves of the basal nodes 

consisting of a sheath only or with a rudimentary blade, the 

sheaths shorter than the internodes; nodes thickened, formed 

of the thickened basal rim of the sheath above and the thickened 

upper rim of the internode below, brownish, short-pubescent 

with retrorse hairs. Leaves 5-15 per complement; sheaths 

sparsely papillose-pilose with short and caducous, appressed 

hairs, the margins ciliate, more so toward the upper part; ligule 

membranous-ciliate, small, 0.5 mm long: pseudopetiole 

stramineous, 0.2-0.3 cm long, covered by long whitish hairs; 

blades ovate-lanceolate to lanceolate, 11-15 cm long, 3.5-5.8 

cm wide, acuminate, asymmetric and truncate basally, the 

midnerve conspicuous toward the base, transverse veinlets 

present, the adaxial surface glabrous, the abaxial one pilose 

toward the base, the basal margins shortly ciliate, otherwise 
44 SMITHSONAN CONTRIBUTIONS TO BOTAhY 

FIGURE 24.4lyra loretensis: u, habit (X 0.5); b, portion of a branch (x 3.2); c, male spikelet, side view (x 

13.5); d, ligule and pseudopetiole (x 3.2); e, lower glume, inner side (x 3.2);J upper glume, inner side (x 3.2); 

g, anthecium, dorsal side (x 6.5); h, anthecium, ventral side (X 6.5). Component u, a composite: base from Froes 

31193 (IAN), inflorescence from L. Will- 3001 (US); habit and inflorescence from Cowan and Maguire 

38104 (US). Components b,d,e-h based on Cowan and Maguire 38104 (US); c on Pires and Black 845 (US). 

Illustration by G.B. Threlkeld. 
NUMBER 69 45 

FIGURE 25.-Detail of female anthecium, Olyru loretensir: a, apex of the anthecium, dorsal side (X 50); b, apex 

of the anthecium, ventral side (X 50); c, base of the anthecium, dorsal side (X 50): d, detail of the surface of the 

lemma with cylindrical macrohairs (x 200). All micrographs are of Grass1 10089 (US). (Reduced to 77% of 

indicated magnifications for publication.) 

scabrous. Non-leafy culms with leaf sheaths only; sheaths 

stramineous to purplish, wide, strongly ribbed and overlapping, 

sparsely pilose. InJ'orescences paniculifor m, terminal, long-to 

short-exserted, the young panicles included in the upper 

sheaths, these pyramidal, lax, 6-12 cm long, 5-14 cm wide, 

the lower branches whorled and with male spikelets only, the 

upper ones alternate to opposite, with numerous and spaced 

male spikelets in long tertiary branchlets, the female spikelets 

terminal and solitary in each culm; axis and branches 

longitudinally ridged, scabrous to glabrous, the ails of the 

branches pilose, brownish. Female spikelets lanceolate, aristate, 13-18 mm long, 2.5-3 mm wide, whitish, the glumes 

subequal, longer than the anthecium, scabrous to sparsely 

pilose toward the apex, otherwise completely glabrous, tipped 

by purple awns; lower glume 5-9-nerved with manifest 

transverse veinlets, long-aristate, the awn of 6-8 mm long; 

upper glum 5-9-nerved with transverse veinlets, aristate, awn 

of 4-6 mm long; anthecium long-ellipsoid, narrowed at the 

base, short-stipitate, 7.8-8.1 mm long, 2.1-2.5 mm wide, 

acuminate apically, whitish, pilose with long, appressed and 

cylindrical macrohairs over the entire surface; lemma 3-nerved, 

palea 2-nerved. Caryopsis fusiform, brownish, 4.5 mm long, 

2.2 mm wide; hilum linear, as long as the caryopsis. Male 

spikelets lanceolate, 2.5-3.3 mm long, 0.5-0.7 mm wide, 'Is 

the length of the female spikelets, scabrous to scaberulous; 

lemma 3-nerved, aristulate; palea 2.2-2.6 mm long, 2-nerved. 

DISTRIBUTION.-This is an Amazonian species that grows 

in the dense forests of equatorial Brazil, Colombia, and Peru 

(Figure 15), at elevations from 0 to 400 m. 

ADDITIONAL SPECIMENSEEN.-BRAZIL, AMAPA: Serra 
46 SMITHSONIANCONTRIBUTIONS TO BOTANY 

do Navio, Emmerich and Andrade 712 (R, US), Serra do 

Navio, Cumca Ore Body, Cowan and Maguire 38104 (NY, 

RB, US). AMAZONAS: Esperanca, Pires and Black 845 (IAN, 

US), Krukoff 7502,7512,751 7 (NY); Estrada Humaia-Labrea, 

Vieira et al. 53 (MO); Sao Francisco, Rio F’unu, Kuhlmann 

913 (IAN, RB, US); road Humaia to Porto Velho, km 10, 

Prance et al. 3446 (F, MO, US); Rio Solimoes, Fodes 24058 

(IAN); Solimoes river, Barrio de Odorio, Sao Paul0 de 

Olivenca, Lleras et al. PI7389 (US); Manaus-Porto Velho 

Highway, km 380, 2 km S of Rio Jut& Prance et al. 22865 

(US); Rio Purus, between Lago Quati and Lago Arima near 

Rio Jacare, Prance et a1 13427 (US); km 620 on ManausHumaita road, Lowrie et al. 73 (MO); Rio Javari, noninundated disturbed forest behind village of Paumari, Gentry 

and Revilla 20559 (MO). PARA: Santarem, Estrada para o rio 

Curua-Una, acampamento do Juarana, Cavalcante and Jouc 

1470 (US); Taperinha, parana do Ituqui, Froes 31193 (IAN). 

RONDONIA: 8 km NE of Porto Velho, Prance et al. 8246 (MO, 

COLOMBIA, AMAZONAS: Rio Amaca-Yacu, 20 km de son 

embouchure sur I’Amazone, Sastre and Echeverry 612 (US); 

Leticia, Black and Schultes 46-1 (IAN, US); 18 km N of 

Leticia, Gilleft I6471 (MO); Loretoyacu River, 3”46‘S,7Oo23‘ 

N, Grass1 10089 (US). 

PERU, JUXIN: Puerto Bermudez, Killip and Smith 26568 

(US). LORETO: San Fernando, frontier military post on Rio 

Jauari, above Santa Rita, Gentry and Revilla 20914 (F, MO); 

La Victoria, Williams 3001 (F, US). 

D1scUssIoN.-Flowering occurs between August and February. Olyra loretensis is unique in the genus in possessing 

inflorescences borne on specialized reduced-leafy culms which 

completely lack blades. It is a handsome species, in which the 

white color of the body of the female glumes contrasts 

strikingly with the purple color of the panicle branches, female 

spikelet pedicels, female glume awns, and male spikelets. 

Gould and Soderstrom (1970) have recorded a chromosome 

number of 2n = 22. 

US). 

16. Olyra maranonensis Swallen 

FIGURE IS 

Olyru marunonemis Swallen, 1966:86. [Type: “Type in the U.S. National 

Herbarium No. 2382316, collected in rainforest on ridge crest of Quebrada 

Chuivi, valley of Rio Maranbn near Cascades de Mayasi, elevation 500-550 

m., Amazonas, Peru, September 20, 1962, by John J. Wurduck (No. 1936)” 

Holotype, US sheet no. 2382316.1 

from an asymmetric base to an acuminate, symmetric tip, 

ascending, glabrous, the margins scaberulous, the midnerve 

conspicuous toward the base on the adaxial surface. Inflorescences paniculiform; panicles terminal, lax and pyramidal, 13 

cm long, 4 cm wide, the lower branches whorled, the upper 

whorled to alternate, spreading, the lower branches naked at 

the base and with male spikelets only, the upper branches with 

male spikelets arranged in tertiary branchlets below and one 

terminal female spikelet in each branch or the female spikelets 

borne solitary from the axis in the upper portion; axis 

longitudinally ridged, scabemlous, purplish, the branches 

triquetrous, scabrous, the axils of the branches glabrous; 

pedicels of the female spikelets thickened, scaberulous to 

glabrous. Female spikelets fusiform, aristate, 11-25 mm long, 

2.5-2.7 mm wide, glabrous, whitish; lower glume 11-25 mm 

long, aristate, 5-nerved with transverse veinlets, glabrous on 

the inner surface, short-scabrous toward the apex and margins 

of the outer surface, the awn of -10 mm long; upper glume 

acuminate, 10-13 mm long, 7-nerved with transverse veinlets, 

glabrous on the inner surface, short-scabrous toward the apex 

of the outer surface; anthecium ellipsoid, acute, 7.5 mm long, 

2.7 mm wide, smooth and shining, whitish to stramineous, 

with long, flattened macrohairs toward the apex and base of 

the lemma on the ventral surface, the margins of the lemma 

with prickle hairs, dorsal surface glabrous. Caryopsis not seen. 

Male spikelets fusiform, short-aristate, 3.5-4.7 mm long, 

0.6-0.7 mm wide, hispid; lemma 3-nerved; palea 2.8-4 mm 

long, 2-nerved. 

DISTRIBUTION.-Known only from the type locality in 

northern Peru (Figure 15). 

DISCUSSION.-Olyra maranonensis resembles 0. glaberrim, differing from it by the smaller, hispid male spikelets and 

scaberulous pseudopetioles. 

17. OZyra micrantha Humboldt, Bonpland & Kunth 

FIGURES 17dJ 26-28 

Olyru micrunfha Humboldt, Bonpland & Kunth, 1816:199. [Type: “Crescit in 

umbrosis. humidis ad ripam Orincci prope cataractam Maypurensium.” 

Holotype, P, not seen, fragment of the holotype, US sheet no. 2877940.1 

Olyru hirsufu Trinius, 1826:250. [Type: “Brasil. (Langsdorff).” Holotype, LE, 

not seen.] 

Olyru scorbiculufu Schrader ex Nees von Esenbeck, 1829:303. [Nomen 

nudum.] 

Olyru venrricosu Nees von Esenbeck, 1829:303. [Type: “Var. alpha in Mont. 

Corcovado, haud procul a Rio de Janeiro urbe. (Mart. Langsdorff).-Var. 

bela in paludibus ad Paraiba. (Seren. Princ. Muximil. Neovid., nec non in 

ripa lacas Canomensis, in sylvaticis, et alibi in pro, Paraensi, juxta fluv. Cespitose perennials* Culms erect, -’ unbranched; Amamnum (Mar[,),- Vidi in Herb, Regio Berol, a Sel[owjo lecbm).-var, 

internodes cylindrical, hollow, glabrous, stramineous to in sylvis aboriginibus ad Sebastianoplin aliisque in locis prov. Rio 

purplish; nodes thickened, purplish, glabrous. Leaves with 

sheaths pale to greenish, mottled with dark spots, the margins 

de Janeiro. Floret Martio-Majo, et Novembr.” Holotype, B, not seen.] 

seen.] 

Manaos prov. do Alto Amazonas (Spruce).” Holotype, B, not seen.] 

olyru micrunth var. lunceolutu Doell in Martius, 1877:324. [Type: ”prOpe 

Rio de Janeiro (Guudichaudn. 297)’’ Holotype, P. not seen. fragment of the 

Olyru wvillei Steudel. 1853:36. [Type: “ex Hrbo. Urville.” Holotype, P, not 

ciliate to glabrous, the rest of the surface shortly pubescent to olyru micrudha var. ~cu~vufu Doell in Martius, 1877:324, [Type: g*prope 

glabrous; ligule membranous-ciliate, -0.5 mm long; pseudo~etiole -0.3 cm long, dark, scaberulous on both surfaces; blades 

lancedate, flat, 18-23 Cm long, 3.6-4 cm long, narrowing 
NUMBER 69 47 

FIGURE 26.-OIyru micrunthu: u, inflorescence (x 0.5); 6, leaf blade (x 0.5); c, base of the plant (x 0.5); d-J 

caryopsis side view (x 6); g, female anthecium, dorsal side (x 6); h, female anthecium, ventral side (x 6); i, 

lemma of the male spikelet, side view (X 6); j. palea of the male spikelet, inner view (x 6); k, lower glume of 

the female spikelet (X 6); 1, upper glume of the female spikelet (x 6); m, portion of a branch of the inflorescence 

with a male spikelet (x 6); n, female spikelet (x 6); o, male spikelet (x 6). Components u and b based on Irwin 

el ul. 54892 (US), c on Alfson 496 (US), and d-o on Belem 1814 (US). Illustration by G.B. Threlkeld. 
48 SMRHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 27.-Detail of female anthecium, Olyra micrunthu: a, apex of the anthecium, dorsal side (X 50); b, 

detail of the surface of the lemma with rounded excavations (x 200); c, detail of the surface of the palea (x 200); 

d, anthecium, ventral side (x 50). Micrographs a,b of Chose 10186 (US), c,d of Anderson 867 (US). (Reduced 

to 77% of indicated magnifications for publication.) 

holotype, US sheet no. 2877940.1 

Olyru micrunthu var. subvelutina Doell in Martius, 1877:324. [Type: “prope 

Rio de Janeiro (Riedel. sub nomine Olyrae pubescentis).” Holotype not 

located.] 

Olyru d’Urvillei Doell in Manius, 1877:323. [Nomen nudum.] 

Cespitose perennials in dense clumps of as many as 20 culms 

borne close together from a short, creeping rhizomatous base. 

Culms erect, 1-4 m tall, climbing and leaning on the 

vegetation, densely branching at the upper nodes; internodes 

cylindrical, hollow, glabrous; nodes compressed, dark, shortly 

pubescent to glabrous. Leaves with sheaths stramineous, 

densely pubescent with long and appressed to erect hairs; ligule 

membranous-ciliate, conspicuous, 2.4- 15 mm long, glabrous 

on the adaxial surface, sparsely pubescent on the abaxial one; 

pseudopetiole 0.4-1.2 cm long, pale to purplish, shortly pilose; 

blades oblong-lanceolate, 13-32 (-55) cm long, 2.4-1 1.5 

(-13) cm wide, acuminate apically, the base symmetric and 

subcordate, the adaxial surface greenish, glabrous to occasionally hispid to scabrous, the abaxial surface glaucous, glabrous 

to hispid, the midnerve prominent, the margins shortly ciliate. 

Inflorescences paniculiform, borne from the uppermost nodes; 

panicles lax and diffuse, pyramidal, 10-32 cm long, 4-14 cni 

wide, the branches spreading, the lower ones whorled and 

bearing male spikelets only, the upper ones alternate with male 

spikelets below and terminal female spikelets; axis longitudinally ridged, hispid, the branches angled, hispid; pedicels of 

the female spikelets thin, not thickened, similar to the ones of 

the male spikelets, scabrous to hispid. Female spikelets 
NUMBER 69 49 

FIGURE 28.-Distributim of Olyro microntho. 

disarticulating below the glumes, ovoid, aristate, 5.4-12.8 lower and upper glume; lower glum aristate, 5-nerved; upper 

mm long, 1.8-2.6 mm wide, hispid to scabrous, the glumes glum aristate to subulate apically, 3.7-9.2 mm long, 

separated by an internode, falling with the anthecium, unequal, 3-5-nerved; anthecium ovoid, acute, 2.8-3.5 mm long, 

papillose-pilose with long to short hairs on the outer surface 1.3-2.1 mm wide, minutely scabrous toward the apex of the 

to scabrous, the inner surface of the glumes densely hispid, lemma to completely glabrous, brownish, pitted, with rounded 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

excavations over the entire surface; lemma 5-nerved. Male 

spikelets fusiform, aristate, 7.4-9.3 mm long, 0.8-1.1 mm 

wide; lemma long-aristate, scabrous, sparsely pilose; palea 

4.67.1 mm long, 2-nerved, scabrous; anthers 3, 3.7 mm long. 

D1sTRIBunoN.-Widely distributed in South America, from 

Colombia and Venezuela eastward to the Atlantic Coast and 

westward to the Andes (Figure 28), at elevations mostly below 

500 m; the southwestern boundary of its range apparently is 

determined by rainfall, with this species occurring in the wetter 

areas of Peru, Bolivia, Argentina, and Paraguay. There is also 

a nineteenth century collection from the Fiji Islands that must 

represent a casual introduction. 

San Pedro, S of Tobuna, Renvoize et al. 3232 (US). 

BOLIVIA, BENI: Chacobo village of Alto Ivon, Boom 4091 

(US). LA PAZ: San Carlos, Buchtien 38, 39, 65, 77, 78, 1158 

(US), 76 (NY). PANW: Rio Arroyo, Sperling and King 6554 

(US). SANTA CRUZ: Rio Vibora, Steinbach 7572 (NY). 

BRAZIL, ACRE: Vicinity Sena Madureira, Cid and Nelson 

25410 (MO). AMAPA: Rio Jari, base of Serra da Arumanduba, 

Egler and Irwin 45967 (NY, US). AMAZONAS: Near M~MUS, 

Spruce 1533 (NY, US); Manaus, estrada do igarape do 

Tabatinga, Rodriguez and Chagas 2823 (US); Manaus, estrada 

antiga de S. Raimundo, Chagas and Coelho 3773 (US). BAHIA: 

Fazenda Barra do Manguinho, Mattos Silva et al. 1067 

(CEPEC); Sb Bento dos Lages, Lutzelburg 137 (NY); Castelo 

Novo, Velloso 1099 (US); Jaguaquara, Pinto 619 (US); Marau, 

Belem 1814 (CEPEC, US); Rodovia BR 5, 16 km S of 

Eunapolis, Bele'm and Pinheiro 2675 (CEPEC, US); Rodovia 

Camacan-Canavieira, 3-30 km de Camacan, Belem et al.

1392 (CEPEC, US); Santa Cruz do Cabralia, Eupunino 280 

(CEPEC, US); Dunas de Itapue, Noblick 1625 (CEPEC); 

Belmonte, Santos 1128 (CEPEC, US); estrada Urucuca-Serra 

Grande, Comes 101 (RB); Rodovia Belmonte-Itapebi, km 26, 

Mattos Silva and Huge 590 (CEPEC); Rodovia MarauUbaitaba, km 3, Calderon 2036 (US); Riveiab de Fora, 4 km 

S of Itamaraju city on road Itamaraju-Teixeira de Freitas, 

Calderon and Pinheiro 2213 (CEPEC, US); Semnha, Capanema s.n. (US 950601). CEARA: Baturite to Guaramirango, 

Swallen 441 7 (US). ESPIRITO SANTO: Est. Pedro Palacios, Boa 

Vista, Vieira 52 (RB); between Barra de Santa Angelica and 

Burarama, Soderstrom and Sucre 1971 (US); Reserva Biologica 

de Sooretama, Mata do Cupido, Soderstrom and Sucre 1899 

(CEPEC, US); road from Cachoeiro de Itapemirim to Itabira, 

Soderstrom and Sucre 1966 (US); Serra do Capara0, Chase 

IOlll (US); SW of Vitoria, Soderstrom and Sucre 1875 

(CEPEC, US). MARANHAO: Rio Pindare, Moncao, Froes 20306 

(US). MATTO GROSSO: Entre Cururu y Vilhena, Kuhlmann 

1867 (RB); margem esquerda do Rio Juruena, Rosa and Santos 

213 9 (RB). MINAS GERAES: Juiz de Fora, Schwacke s.n., 1895 

(RB); without locality, Lindberg 512 (US); Caldas, Regnell 

III-1377 (P, US); Lavras, Black 394b (US); Vicosa, Chase 

10186 (US), Mexia 5089 (US, WIS), 5139a (P, US), 5113 

(NY, P, US, WIS), 5153 (NY, WIS); road to Barroso, about 

ADDITIONAL SPECIM EVS SEEN.-ARGENTINA, MISIONES: 

km 6, Mexia 5183a (US, WIS). PARA: Thome Assu, Agua 

Branca, Mexia 5952 (NY, US); S slope of Akarai Mountains, 

in drainage of Rio Mapuera, A. Smith 2934 (NY, US); Boa 

Vista on the Tapajos River, Dahlgren and Sella 82 (US), 

Swallen 3131 (US); Breves, Pires and Silva 6682 (US); 

Castanhal, Braganca railroad, Goeldi 300 (NY, RB, US); 

Fordlandia, Tapajos River region, Krukoff 1047 (NY); Maraba, 

Serra dos Carajas, Cavalcante 2117 (US); Obidos, Swallen 

5081, 5090, 5092 (US); Sete Varas airstrip on Rio Curua,

Strudwick and Sobel 4311 (WIS). PARANA: Cadeado, Hatschbach 14715 (US); Camp0 Mofao, Swallen 9010 (US); Cerro 

Azul, Hatschbach 5037 (US); Mono do Farol, Caoiba, 

Imaguire 158 (US); Parana River near Porto Byington, 

Lindeman and Has s I71 7 (US); Pedra Branca de Araraquara, 

Hatschbach 12878 (US); Porto de Cima, Dusen 10257 (NY, 

US). PmAMBUCO: Without locality, Gardner 1198 (US); 

Antonio, Saito 291b (US); Fazenda Arabari, Enorio 66-198 

(US). RIO DE JANEIRO: Alto Macahe, Nova Friburgo, Glaziou 

18620 (NY, P, US); Alto da Boa Vista, Calderon 2028, 2030 

(RB, US); Cascadura, Holway and Holway 1473 (US); Cosme 

Velho toward Corcovado, Morro Novo Mundo, Soderstrom et 

al. 1859 (CEPEC, US); Corcovado, Fluster 1141 (US), Rose 

and Russell 20216 (NY, US); Gavea, Chase 9981 (US); Mt. 

Itatiaia, road between Monte Serrat and Ponte Maromba, 

L.Smith 1439 (US); Itatiaia National Park, S face of Mt. Itatiaia, 

Eiten and Eiten 6499 (US); Madalena, Feio and Castro Faria 

s.n. (US 2463828); Mage, Fazenda do Cortume, VidulI1-3997 

(US); Monte Serrat, Serra de Itatiaia, Chase 8353 (US); Morro 

Novo Mundo, Soderstrom et al. 1856 (US); Restinga de 

Jacarepaga, at east side of Pedra de Itauma, Soderstrom and 

Sucre 1915 (CEPEC, US); Rio de Janeiro, Holway and Holway 

1077 (US), Chase 10004 (NY, US), Calderon 2026 (US), Nee 

3400 (US); Serra do Corcovado, entre Vista Chinesa e Mesa 

do Emperador, Calderon 2014 (RB, US); Serra da Mendanha, 

Soderstrom and Sucre 1951 (US); Teresopolis, Brade 9677, 

9690 (US), Emygdio 81 (US); Tijuca, Chase 12152 (US); Vale 

de Bom Sucesso, Caixa d'Agua, Soderstrom and Sucre 1983 

(CEPEC, US), Sucre 5844 (US), Sucre 5850 (US); Vista 

Chinesa, Schwacke s.n. (US 2463831); Mata do Rumo, Sucre 

7684 (RB); Vista Chinesa, Sucre 7659 (US); Alto da Pedra da 

Givea, Sucre 4335 (RB); Jacarepagua, Sucre 9606 (RB); alto 

da Boa Vista, Sucre 1120,1136 (RB); restinga da Tijuca, Sucre 

1057 (RB); Parque Nacional da Tijuca, Lima 686 (RB); Mesa 

do Imperador, Almeidu 1985 (RB). RORAIMA: Mt. Roraima, 

ridge NW of Paulo, Tate 162 (NY); Mt. Roraima, along 

Kukenam River at Roraima ford, Paulo, Tate 182 (NY); Serra 

Tepequem, in quebrada near airstrip, Maguire 40054 (NY); 

foothills of Serra Tepequem, near Igarape Paparu, Prance et 

al. 4341 (US); Semnha, Rio Mucajai, Prance et al. 4187 (US); 

Serra dos Sururucus, S of Mission Station, Prance et al. 10053 

(US). sAo PAULO: Butantan, Kuhlmann 557 (US); Campinas, 

Novaes 1280, 1281 (US); Campinas, Fazenda Santa Ana, 

Viegas 5036,SI 02 (US), Fazenda Riqueza, Viegas 2876 (US); 

25 km NW de Mogi Guacu, Gibbs and Leitao Filho 6098 
NUMBER 69 

(US); Sao Paulo, Parque do Estado, Calderon 2007 (US), 

Fonseca 4 (US), Skvortzov 167 (US), Gehrt 12892 (US); 

Pirassumunga, Pick1 5820 (US); Santos, Vaccari s.n. (US 

1625216); Santos, Guaruja, Holway and Holway 2014 (US); 

Santos, Monserrat, Rosengurtt PE-3401 (US); S. Jose dos 

Campos, Eiten and Mimura 3350 (US), Lofgren 3831 (US); 

Sao Paulo, Usteri 9975 (US), Hoehne 29798 (NY, US); 2 km 

E of Sete Barras, Clayton and Eiten 4670 (US); Xiririca, 

Hoehne 24313 (US). SANTA CATARINA: Azambuja, Calderon 

2004 (US), Klein 534 (NY, US), Reitz 2214 (US); Blumenau, 

Reitz and Klein 2201 (NY), Ule 893 (US); Bom Retiro, 

Bresolin 807 (US), Klein and Souza 10519 (US); Braco do 

Norte, Rio do Medio, Mattos 7162 (US); Brusque, Reitz 5732 

(US); Mato de Hoffmann, Smith and Veloso 5669 (US); 

Ibirama, Klein 613 (US), Reitz and Klein 3702 (US), Smith 

and Klein 7569 (RB, US); Ilha de Santa Catarina, Morro da 

Lagoa, Rohr 484 (US); Jordao, Klein 9672 (US); Morro do 

Cambirela, Bresolin 314 (US); Morro do Cavalo, Reitz and 

Klein 955 (NY, US); Morro da Cruz, Valls et al. 2734 (US); 

Morro da Fazenda, Calderon 2005 (US); Morro da Quebrada, 

Klein and Bresolin 8355 (US); Morro no Rio Tavares, Klein 

and Bresolin 6244 (US); Pantano do Sul, Klein et al. 7950 

(US); Piloes, Reitz and Klein 3802 (US); Praia Braba, Reitz 

and Klein 1078 (US), 1082 (NY, US); Saco Grande, Klein and 

Bresolin 8399 (US). 

FIJI ISLANDS: Sandalwood Bay, anno 1838-1842, Wilkes 

s.n. (US). 

FRENCH GUIANA: Layon point de Trijonction-Mitaraka, 

km 7.5, Sastre 1585 (US); St. Jean, Benoist 1284 (US); Saint 

Laurent, Hoock 96 (NY, US); Route de St. Laurent, Hoock 

s.n., 15 Apr 1961 (P); Tumuc Humac, Hoock s.n., 23 Jan 1956 

GUYANA: Maicwac River, Altson 496 (US); Alubia Falls, 

Kamerang River, Leaky 217 (NY, US); upper Demerara 

River, Jenman 4119 (US); Kanuku Mountains, forest trail to 

Mt. Iraimakipang, Goodland and Maycock 465 ( US); 

Essequibo River, Moraballi Creek, near Bartica, Sandwith 

282 (US); Rockstone, Cleason 624 (NY, US), Hitchcock 

17283 (NY, P, US), Jenman 7548 (US); Mt. Roraima, 

McConnell and Quelch 708 (NY, US). 

PARAGUAY: Ibitimi, Rojus 6110 (US); Paraguari, Rojas 

12938 (US); Pedro Juan Caballero, Rojas 6398 (US); Santa 

Teresa, Bertoni 1688 (US); Sierra de Amambay, Hassler 

11314 (NY, P, US). 

PERU, cuzco: Consuelo, Asuncion, Vargas 7361 (US). 

HUANUCO: Rio Azul, near Tingo Maria, Ferreyrd 12737 (US); 

without locality, Anderson 867 (US). JUNIN: Chanchamayo 

Valley, Schunke 631 (US). LORETO: Vicinity Aguaytia, Croat 

20968 (MO). MADRE DE DIOS: Atalaya, vicinity of Hacienda 

Ammonia, 2-3 km west of village, Foster and Wachter 7378 

SURINAM: Arrowhead Basin, Maguire 24456 (NY); 

Brownsberg, Lindeman 12504 (US); Coppename River near 

Raleighfalls, Lanjouw 785 (US); 2 km S of Juliana Top, Irwin 

(PI. 

040). 

51 

et al. 54645 (US); near summit of Juliana Top, Irwin et al. 

54891,54892 (US); area of Kabalebo Dam project, Lindemun 

et al. 317 (US); hills 9 km N of Lucie and 12 km W of Oost 

riviers, Maguire et al. 54232 (US); vicinity of Moengo, Cottica 

River, native trail to Adjoema Kondre village, Cowan 39004 

(NY); Nassau, Lanjouw and Lindeman 2554 (NY); Saramacca 

River headwaters, Jacob kondre, Maguire 23826 (NY, US); 

Zanderij 11, Maguire 23707 (NY, US). 

VENEZUELA, AMAZONAS: Cerro Duida, Steyermark 58075 

(US); Cerro Huachamacari, Rio Cunucunuma, vicinity of 

Camp 11, Maguire et al. 29941 (NY, US); Cerro Sipapo, 

vicinity of Base Camp, Maguire and Politi 27987 (NY, US); 

Serrania Paru, Rio Paru, Cano Asisa, Rio Ventuari, Cowan and 

Wurdack 31440 (NY, US); Yavita, Williams 14007 (US). 

ANZOATEGUI: Medanos de Puerto La Cruz y El Morro, Tamayo 

20 99 (MO). APURE: Reserva Florestal San Camilo, Steyermurk 

et al. I01 756 (MO). BOLIVAR: Rio Chizca, region de Uriman, 

Bernardi 768 (NY); Gran Sabana, between waterfall at 

Rue-menu and Divina Pastora on Rio Kukenan N of Santa 

Elena, Steyermark 59224 (NY, US); Hato de la Divina Pastora, 

Gran Sabana, Tamayo 2893 (US); Hacha, Bernardi s.n. (NY); 

Sierra Ichun, cercanias del Salt0 Maria Espuma del Rio Ichun, 

base of the Sierra de Ichun, Steyermark 90351 (NY, US); SE 

of La Queina, Wurdack and Guppy 157 (NY, US); river banks 

of Raudal Maracca, about 110 river km from mouth, Wurdack 

and Monachino 41046 (NY, P, RB, US); Isla El Casabe, Rio 

ParagUa, Killip 37304 (NY, us). FALCON/LARA: CerrO SOCOpa, 

Liesner et al. 8336 (MO). MEWDA: Along Rio Onia, near 

Bolero, N of Mesa Bolivar, Steyermark 56737 (NY, US). 

TACHIRA: region of La Fundacion, Liesner and Gonzalez 

10188 (MO), Liesner et al. 9543 (MO), van der Werff 4940 

(MO). ZULIA: Deparahento Colon, Casigua El Cubo, Bunting 

et al. 6704; Departamento Bolivar, entre Las Tres Marias y el 

Rio Chiquito, Bunting and Stoddart 9045 (MO); Depatamento 

Perija, Est. Hidropogica Aricuaisa-Pie de Monte, Bunting and 

Panapera 11024 (MO). 

DIscUssION.-Olyra m'crantha is atypical of the genus in 

having thin-pedicellate female spikelets in which the glumes 

fall attached with the anthecium, and in the conspicuous 

internode between the lower and upper glume. This peculiarity 

is also present in the genera Raddiella and Parodiolyra, but 

these genera have a different type of caryopsis (in which the 

hilum does not reach the entire length of the caryopsis), female 

anthecium (not pitted), and inflorescence. Robust inflorescences of 0. micranthu probably bear more spikelets than any 

other olyroid grass. 

This is a weedy species with considerable morphological 

variability in the size of plants, leaf blades, panicles, and 

female glume size and pubescence. Many specimens from 

northern South America bear large leaves while in the South, 

as for example in Sao Paulo, Brazil, they are small and narrow. 

There is also variation in the ligule length with some 

populations from around Rio de Janeiro having very long ones. 

However, study of the specimens available showed that 
52 SMITHSONIANCONTRIBUTIONS TO BOTANY 

intermediate types are frequent. An unpublished study made 

by Dr. Lynn G. Clark confirmed that large-leaved and 

small-leaved plants represented extremes of the same species. 

Chromosome counts of n = 20 (Davidse and Pohl, 1974, 

1978) and 2n = 40 (Gould and Soderstrom, 1967) have been 

reported for the species. 

18. Olyra obliquifolia Steudel 

FIGURES 11,29,30a,b 

Olyra obliqui~oliu Steudel, 1853:36. [Type: "Hochst. Hrbr. Kappler nr. 1472 

(non H.B.). Surinam." Holob, P, not seen, fragments of the type and a 

photograph, US sheet nos. 2877937,2975664; isotype, MO.] 

Perennial, short-rhizomatous, borne in fascicles of 10-20 

culms. Cufms 0.3-2 m tall, erect or slightly geniculate at the 

base, simple, many-noded; internodes densely pilose with 

retrorse hairs to glabrous, cylindrical, hollow; nodes thickened, 

purplish, short-pubescent. Leaves 5-9 per complement; 

sheaths short and densely pilose to glabrous, strongly ribbed, 

the margins ciliate to glabrous, membranous; auricles present, 

small; ligule membranous, short-ciliate at the apex, 3-3.5 

mm long, short-pubescent on the abaxial surface; pseudopetiole 0.2-0.4 cm long, densely to sparsely pilose, with whitish 

and stiff hairs to glabrous; blades narrowly oblong, 16-30 cm 

long, 4-9 cm wide, glabrous, narrowing to an asymmetric tip, 

narrowed and asymmetric basally, the margins ciliate to 

scabrous, the midnerve prominent on both surfaces, the lateral 

nerves anastomosing. Inflorescences umbelliform, borne from 

the uppermost nodes, panicles 12-16 cm long, 12-20 cm wide, 

each with a fascicle of 2-8 branches, the branches spreading 

and reaching approximately the same height, falling at 

maturity; peduncle hispid, the branches hispid toward the basal 

part, otherwise glabrous, triquetrous with one side flattened, 

with 1-4 terminal female spikelets and congested male 

FIGURE 29.-Detail of female anthecium, Olyra obliquifolia: a, apex of the anthecium, ventral side (x 50); b, 

apex of the anthecium, dorsal side (x 40); c, detail of the excavations on palea (x 600); d, base of the anthecium 

showing stipe (X 30). Micrographs a-c of Muguire 40720 (US), d of Cavalcante 847 (US). (Reduced to 77% of 

indicated magnifications for publication.) 
NUMBER 69 

FIGURE 30.-Field photos of Olyru species. 0. obliquifolia: u, habit of plant in forests of Brazil; b. detail of 

urnbellifom panicle. 0. standleyi: c, habit of plant, Panama; d, panicles; g, detail of a panicle. 0. glaberrima: 

e, habit of plant, Brazil;f, detail of male and female spikelets. All photographs by C.E. Caldeoh, 1968. 

53 
54 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

spikelets below, the male spikelets borne alternately in pairs 

(or in groups of 4) along the branches; pedicels of the male 

spikelets short, hispid, those of the female spikelets thickened, 

short-pubescent; axils of the branches densely pilose. Female 

spikelets lanceolate, acuminate to short-aristate, 13- 17 mm 

long, 3.4-4 mm wide, disarticulating above the glumes, 

brownish to purplish, densely papillose-pilose to sparsely 

pilose, occasionally glabrous; glumes acuminate to subulate 

apically, longer than the anthecium, 7-9-nerved with transverse veinlets, the nerves strong, hispid on the outer surface, 

more so toward the margins and apex to glabrescent, the inner 

surface densely pilose on the upper portion; upper glume 

13-15 mm long; anthecium fusiform, acute, 9.2-11.5 mm 

long, 3.1-3.4 mm wide, glabrous, pale, separated from the 

glumes by a conspicuous stipe 0.6-1 mm long, the surface of 

the anthecium with hexagonal to rounded excavations. 

Caryopsis broadly ovoid, 5.8 mm long, 2.6 mm wide, dark 

brown; hilum linear, as long as the caryopsis. Male spikelets 

lanceolate, acuminate, 5.5-8.7 mm long, 0.8-1.3 mm wide, 

brownish to purplish, densely hirsute to scabrous; lemma 

acuminate, 3-nerved, densely hispid, more so toward the 

margins to scabrous; palea 5-8.1 mm long, 2-nerved, sparsely 

hispid with margins short-ciliate to glabrous; anthers 4 mm. 

DISTRIBUTION.-Northern South America from Par& Brazil, 

and Surinam east to Maranhao, Brazil (Figure Il), from sea 

level to 700 m elevation. 

ADDITIONAL SPECIMENS SEEN.-BRAZIL, AMAPA: Mapati, 

km 116, Black and Lobato 50-9686 (US); Rio Oiapoque, 5 

km SE of Clevelandia, Irwin et al. 47430 (NY); Rio Araguari, 

lo26'N, 51°58W, Pires et al. 50803 (NY, US), vicinity of 

camp 12, loll", 52'8W, Pires et al. 51367 (F, NY, R, US); 

40 km SW of Serra do Navio, Rio Cupuxi, Dembros 304 (RB). 

MARANHAO: Rio Pindare, Moncalo, Froes 20302 (US); Rio 

Alto Turiacui, Barraquinha, Jangoux and Bahia 149 (MO); 

Turiacui, Rosa and Vilar 2746 (MO); S of Fazenda Guarany, 

km 133 of BR-316, Duly et al. 166 (MO). PARA: Belem 

Soderstrom 1192 (US), Belem I.P.E.A.N., Calderon 2062 

(US); 20 km S of Belem, Irwin 5025 (US); Castanhal, Braganca 

railroad, Goeldi 42, 74 (US), 301 (F, MO, NY, RB, US); 

Castanhal, Colonia 3 de Outubro, Black 49-8592 (US); 

cabeceiras do Alumine, Sampaio 5815 (R); road BR 22, 

Capanema to Maranha0, Prance and Pennington 1870, 1895 

(US), km 60, Prance and Silva 58765 (F, MO, US), km 64, 

vicinity of Piritoro, Prance and Silva 58872 (F, MO, US), 

Prance and Pennington 1946 (F, US); km 167-173 da Estrada 

Belem-Brasilia, Oliveira 541 (IAN); Rio Capim, Caratateua, 

Black 48-2368 (NY); Maguary, Snethlage I1 7 0; entre a fonte 

do Rio Mapua e o Rio Piria, Black 50-9826 (US); Marituba, 

Granja Imperial, Pereira 5044 (US); Rio Paru de Oeste, 

Cavalcante 847 (US); Igarape Pixuna, Black 48-2978 (US); 

Regiao Igarape Pitoro, Froes 34731 (IAN); Santa Isabel, 

Swallen 5045 (R, RB, US), Pires and Black 1330 (RB). 

FRENCH GUIANA: Itany, Village Tolinga, Machiri, Hoock 

97 (NY, US); Gregoire, Deward 125 (US); sommet des Monts 

Atachi Bacca, de Granville 785 (US); Inselberg, Cremers 

7451 (MO); Haut Maroni, Sastre and Moretti 3849 (MO); 

Region de Camopi, Montagne Alikene, de Granville B.5063 

(US); Riviere Sinnamary a Petit Saut, Cremers 5382 (US); 

Montagnes de la Trinite, de Granville 5969 (US, WIS). 

SURINAM: Kabelstation, Essed 51 (MO, US); Lely 

mountains, 175 km SSE of Paramaribo, Mori and Bolten 

8427 (RB, US), Lindeman et al. 50 (US); in montibus, qui 

dicuntur Nassau, Lanjouw and Lindeman 2533 (NY); Nassau 

mountains, Marowijne River, Plateau A, 1 km E of Camp on 

line A, Cowan and Lindeman 39032 (NY, US), occasional in 

forest on Line 12, Plateau A, Maguire 40720 (F, NY, RB, 

US); Anelofoesie, Saramacca River, upstream from Nieuw 

Jacob Kondre, J. and P. Florschutz 1655 (NY); Saramacca 

River, near Posoegronoe, Maguire 24032 (F, MO, NY, RB, 

US); Tafelberg, 0.5 km S of East Ridge, Maguire 24595 (NY, 

US); Zuid Rivier, 2 km above confluence with Lucie Rivier, 

Maguire et al. 53971 (US), Irwin et al. 55736 (F, MO, US); 

without locality, Schwenitz s.n. (NY). 

DISCUSSION.-A remarkable feature of Olyra obliquijolia 

is the presence of umbelliform inflorescences, with all of the 

branches of the panicles whorled and reaching approximately 

the same height; every branch has male spikelets in pairs along 

one side of the axis. It is interesting to note that in this species, 

as in 0. tamaquareana, all of the branches of the panicle fall 

at maturity, only the peduncles remaining at the top of the culm. 

Olyra obliquijolia resembles 0. fasciculata and 0. tamaquareana. The three species are distinguished by the 

presence of a peg-like extension of the base of the anthecium 

that extends downward into a small cavity formed by the outer 

wall of the rachilla internode. Davidse (1987) reported that this 

stipe is full of oil and serves as a functional elaiosome for the 

dispersal of the seeds by ants. 

Chromosome numbers of 2n = 23, 44 (Gould and 

Soderstrom, 1967) have been recorded for the species. Flowers 

throughout the year. 

19. Olyra retrorsa Soderstrom & Zuloaga, new species 

FIGURES 11,31.32 

TYPE SPECIMEN.-BRAZIL, MAT0 GROSSO: Rio Verde, Apr 

1918, Kuhlmann 1868 (holotype, RB; isotypes, IAN, RB). 

FIGURE 31.-U[yra retrorsa: a, inflorescence terminating culms (x 0.6); b, 

branch of inflorescence (X 3.6); c, female spikelet (x 7); d, lower glume of 

female spikelet (X 7); e, upper glume of female spikelet (x 7); female 

anthecium (x 7); g, lemma of female anthecium, inside and outside view (X 

7); h, palea of female spikelet, outside view (x 7); i, palea of female spikelet, 

inside view (X 7);j, palea of female spikelet with margins enclosing the flower 

(X 7); k, enlargement of tip of palea showing retrorse hairs (x 30); [, gynecium 

and lodicules (X 7); m, lodicules of female spikelet (x 15); n, base of pistil 

with lodicule and staminodes (X 15); 0, male spikelet, side view (X 7); p, lemma 

of male spikelet (X 7); q, palea of male spikelet, empty (x 7); r, palea of male 

spikelet enclosing flower (X 7); s, andrecium and lodicules (X 11); 1, ligule and 

upper surface of pseudopetiole (X 4.2). All components based on Kuhlmann 

1868 (RB). Illustration by A.R. Tangerini. 
NUMBER 69 55 
56 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 32.-Detail of female anthecium, Olyru retrorsu: a, apex of the anthecium, dorsal side showing sparse 

bicellular microhairs (x 50); b, apex of the anthecium, ventral side (x 40): c, detail of reworse unicellular hairs 

and bicellular microhairs on the surface of the palea (x 100); d, detail of retrorse, unicellular macrohairs (x 500). 

All micrographs of Kuhfmunn 1868 (RB). (Reduced to 77% of indicated magnifications for publication.) 

Gramen perenne. Culm's erectibus, non ramosus ad basis; 

nodi incrassati, glabri. Laminae lanceolatae, 10.5-13.2 cm 

longae, 2.7-3.4 cm latae, basi symmetricae, subcordatae, apice 

symmetricae, subulatae; pseudopetioli 0.2-0.3 cm longi, pilosi 

breviter; ligulae membranaceae, apice ciliatae breviter, 2.5-4 

mm longae. Paniculue oblongae, contractatae, 6-10 cm 

longae, 1-2 cm latae. Spiculae femineae lanceolatae, subulatae, 

13.8 mm longae, 3.1 mm latae, glumis inter se subaequalibus, 

glabris; gluma inferior subulata, 5-7-nervis, nervis tessellatis; 

gluma superior subulata, 11.6-12.8 mm longa, 5-7-nervis, 

nenis tessellatis; anthoecium lanceolatum, acuminatum, 8.5- 

8.7 mm longum, brunneum, excavatum minute; lemma 

5-nerve, apice pilosum breviter; palea 2-nervis, apice pilosa. 

Spiculae masculae lanceolatae, 9-9.7 mm longae, 0.8-1 mm 

latae, aristatae, glabratae; lemma 3-nerve, aristatum; palea 

acuminata, 5.3-7.5 mm longa, 2-nervis. 

Perennial. Culms erect, unbranched; internodes cylindrical, 

hollow, pubescent; nodes thickened, brownish, glabrous. 

Leaves with sheaths stramineous, papillose-pilose with appressed, caducous hairs; auricles membranous; ligules membranous, conspicuous, 2.5-4 mm long, short-ciliate at the apex, 

glabrous and shiny on the adaxial surface, short-pubescent on 

the abaxial; pseudopetiole 0.2-0.3 cm long, short-pilose; 

blades lanceolate, 10.5-13.2 cm long, 2.7-3.4 cm wide, 

subcordate and symmetric basally, subulate and symmetric 

apically, drooping, the midnerve not prominent, papillosepilose toward the base of the blade on the abaxial surface, the 

rest of the surfac: glabrous, the margins scabrous to glabrous. 

Inflorescences oblong, contracted; panicles borne from the 
NUMBER 69 57 

uppermost nodes, 6-10 cm long, 1-2 cm wide, the branches 

alternate; axis longitudinally ridged, hispid, the branches 

triquetrous, scabrous to short-hispid, the lower branch bearing 

only male spikelets, the upper ones with pairs of male spikelets 

on either side of the triquetrous axis, one subsessile, the other 

short-pedicelled, 1-4 terminal female spikelets on each branch, 

pedicels of the female spikelets thickened, short-hispid. Female 

spikelets lanceolate, subulate apically, 13.8 mm long, 3.1 mm 

wide, the glumes subequal, glabrous; lower glume subulate, 

5-7-nerved with transverse veinlets, scabrous toward the apex 

on the outer surface, pilose toward the apex on the inner 

surface; upper glume subulate, 11.6-12.8 mm long, 5-7- 

nerved with transverse veinlets, the pubescence similar to that 

of the lower glume; anthecium lanceolate, acuminate apically, 

8.5-8.7 mm long, 2.9 mm wide, brownish, slightly pitted with 

oblong excavations; lemma 5-nerved, with bicellular microhairs and long, unicellular macrohairs toward its apex; palea 

2-nerved, with long, retrorse unicellular hairs toward the 

middle portion and the apex. Male spikelets lanceolate, 9-9.7 

mm long, 0.8-1 mm wide, aristate, glabrous; lemma 3-nerved, 

glabrous, brownish, aristate, the awn scabrous; palea acuminate, 5.3-7.5 mm long, 2-nerved, sparsely pilose toward the 

middle, otherwise glabrous, brownish; anthers 3.1 mm long. 

DISTRIBUTION.-Endemic to Mato Grosso, Brazil (Figure 

1 I), and known only from the type locality. 

DIScUSsIoN.-While 0. retrorsa shares with 0. taquara the 

same type of blade morphology and ornamentation of the 

female anthecium, the retrorse hairs on the female palea and 

bicellular microhairs at the tip of the female lemma of the 

former species have no counterpart in the genus. Due to the 

lack of material it is not known whether the species flowers 

irregularly, as does 0. taquara. 

20. Olyra standleyi Hitchcock 

FIGURES 30c,d,g, 33.34 

Olyra standleyi Hitchcock, 1927:86. [Type: “In the US. National Herbarium, 

no. 1307238, collected in moist forest, at El Muiieco on the Rio Navarro, 

Province of Cartago, Costa Rica, alt. 1400 to 1500 m, March 6 to 7, 1926, 

by Paul C. Sfandley and Ruben Torres (no. 50932):’ Holotype, US sheet 

no. 1307238.1 

Robust arching or clambering perennials from thick creeping 

short-rhizomatous rootstocks. Culms up to 3 m tall, decumbent 

and geniculate at the lower nodes, branching sparingly from 

the middle and upper nodes; internodes glabrous, hollow, up 

to 35 cm long and 1 cm diameter; nodes dark brown and with 

short, retrorse whitish hairs to glabrous, flared, formed by the 

thickened basal rim of the sheath above and the thickened 

upper rim of the internode below. Leaves with sheaths strongly 

ribbed, glabrous to sparsely papillose-pilose, with caducous, 

glassy hairs; auricles membranous, glabrous; ligules membranous, -1 mm long; pseudopetiole broad, brownish, flat, 

0.3-0.5 cm long, -0.5 cm wide, glabrous to sparsely pilose 

on both surfaces; blades ovate to oblong-lanceolate, 14-30 

cm long, 4-7 cm wide, greenish to purplish at the base, the 

base symmetric, cordate, narrowed abruptly to an acute to 

subulate and symmetric to slightly asymmetric apex, the 

abaxial surface glaucous, the midnerve prominent, the margins 

short to long-ciliate toward the base to completely glabrous. 

Inflorescences paniculiform, borne on the 8 or 9 upper nodes; 

panicles 9-17 cm long, 5-25 cm wide, with simple branches 

stiffly ascending or spreading from the axis, with 1-3 female 

spikelets borne terminally on each branch and numerous, 

paired (or in groups of 4) male spikelets below, one subsessile, 

the other short- to long-pedicelled, the panicles exserted on 

long, glabrous to short-pilose peduncles -40 cm long, the lower 

branches whorled, the upper ones whorled to alternate or 

opposite; axis longitudinally ridged and sparsely to densely 

pubescent, flattened on one side with the spikelets all borne 

on the other side; axils of the branches pilose. Female spikelets 

18-25 mm long, 2.2-3 mm wide, pale green to brownish or 

purplish, the glumes attenuate, unequal and longer than the 

anthecium, glabrous to occasionally hispid; lower glume 

5-7-nerved with transverse veinlets, glabrous to sparsely 

hispid toward the base and more so toward the apex to hispid, 

the inner surface pilose toward the apex; upper glume shorter 

than the lower, 12.5-19 mm long, 5-7-nerved with transverse 

veinlets, with a pubescence similar to that of the lower glume; 

anthecium fusiform, 8-9.5 mm long, 1.9-2.5 mm wide, 

whitish to stramineous, pitted with oblong to ovate excavations, the apex of the palea covered with bicellular microhairs 

and short prickle hairs; lemma 5-7-nerved; palea 4-nerved; 

lodicules -1 mm long. Caryopsis fusiform, brownish, 6.3 mm 

long, 2 mm wide; hilum as long as the caryopsis. Male spikelets 

lanceolate, aristate, 10-14 mm long, 0.9-1.5 mm wide, 

purplish, glabrous to hispid; lemma aristate, 3-5-nerved with 

transverse veinlets, glabrous to scabrous or hispid; palea 5-10 

mm long, 2-nerved; anthers orange-yellow, 5-7 mm long. 

DISTRIBUTION.-Cloud forests of Costa Rica, Panama, and 

Venezuela (Figure 34), at elevations from 900 to 2700 m 

(occurs at lower elevations in Venezuela). 

ADDITIONAL SPECIMENS SEEN.-COSTA RIGA, CARTAGO: 

Along primitive road, 1 km S of El Muneco, Pohl and Davidse 

I1698 (US); 8 km S of Cartago by air, Finca El Chaparral, 4.5 

km S of bridge on Agua Caliente at Lourdes, Liesner and 

Judziewicz 14606 (MO, US, WIS); El Muneco, S of Navarro, 

Standley 33878 (Paratype, US). 

PANAMA, PANAMA: Cerro Jefe, Calderon 2081 (US), 

Calderon and Dressler 2123 (US), Folsom et al. 2009 (US), 

Soderstrom 2008 (US), J.Z and F. Witherspoon 8482 (US); 

vicinity of Cem Jefe, kwis and Dressler 7558 (US). 

VENEZUELA, AMAZONAS: Cano San Miguel, Rio Guiania, 

Williams 3391 (MO). DISTRITO FEDERAL: Along ridge of the 

Avila, McClure 21246 (US); cordillera del Avila, between Los 

Venados and Pic0 Oriental, Steyermark 55597 (US); near 

summit of El Junquito, Chase 12447 (US), MrClure 21239 

(US); en selvas nubladas de Galipanh, Tamayo 1381 0. 

GUARICO: En )as orillas de la Laguna del Salvaje, Tamayo 
58 SMITHSONIANCONTRIBUTIONS TO BOTANY 

FIGURE 33.--Detail of female anthecium, Olyra sfandfeyi: a, apex of the anthecium, dorsal side (X 50); b, detail 

of the apex of the plea showing numerous prickle hairs (x 250); c, detail of the apex of the plea showing 

numerous bicellular microhairs (x 400); d, detail of the surface of the lemma with oblong excavations (X 200). 

All micrographs of McClure 21246 (US). (Reduced to 77% of indicated magnifications for publication.) 

3391 0. ZULU: Sierra de Perija, loma arriba de la quebrada 

del Rio Omira-Kuna (Tumuriasa), SE de Pishikao e Iria hacia 

la mision Sucurpo, Steyermark and Dunsterville 105686 (MO). 

DIscuss1oN.-When collecting Olyra standleyi in Venezuela, McClure (herbarium label) stated that the plants showed 

signs of dying after flowering, at least the aerial parts. A similar 

observation was made by Sodersuom on Cerro Jefe, Panama, 

where the whole population of the species flowered and died 

in 1973, with seedlings produced. The plants often grow in 

huge, sterile clones. 

The elevations (1400 m in Costa Rica, lo00 m in Panama, 

and 2700 m in Venezuela) at which this species occurs are the 

highest for any olyroid grass, and, indeed, for any herbaceous 

bambusoid. The Amazonas and Zulia, Venezuela plants are 

from lower elevations. Flowering has been recorded from 

October through April. 

The spikelets of Calderon 2081 are densely hispid, while 

in the rest of the specimens examined they are nearly glabrous. 

21. Olyra tamanquareana Soderstrom & Zuloaga, 

new species 

FIGURES 11,35 

TYPE SPECIMEN.-BRAZIL, AMAZONAS: Rio Negro, Ilha 

Tamanquare, downstream near Santa Isabel do Rio Negro 

(Tapuruquasa), Sep 12, 1979, K. Kubitzki, C.E. Calderon and 

H.H. Poppendieck 79-252 (holotype, INPA; isotypes, K, MO, 

LE, NY, SI, US). 

Gramen perenne, caespitosum. Culmis 10-20 in fasciculis 
NUMBER 69 59 

FIGURE 34.-Distribution of Olyra ecaudata, 0. sfandleyi, and 0. taquora. 

densis, non ramosus ad basim, 60-120 cm longis; nodi 

compressi, glabri. Folia per complementum 3-7; laminae 

lanceolatae, 13-25 cm longae, 3-5.5 latae, basi asymmetricae, 

angustatae, glabrae; pseudopetioli -0.4 cm longi, pilosi; ligulae 

membranaceae, apice ciliatae breviter, -3 mm longae. Paniculae contractatae. Spiculaefem’neae lanceolatae, longe aristatae, 

35 mm longae, 4.2 mm latae, glumis inter se subaequalibus, 

anthoecio longioribus; gluma inferior 9- 13-nervis, nervis 

tessellatis; gluma superior 9- 1 1-nervis, nervis tessellatis, 

stipite prominentis, 1-1.2 mm longis; anthoecium lanceolatum, 13-13.2 mm longum, 3.5-3.8 mm latum, glabrum, 

excavatum valde; lemma 5-nerve. Spiculae masculae lanceolatae, acutae, 5 mm longae, 1 mm latae, pilosae; lemma 3-nerve; 

palea 2-neMs. 

Cespitose, robust perennials from creeping rootstocks, 

forming dense clumps with 10-20 culms in a fascicle. Culms 

erect, geniculate at the base, 60-120 cm tall, unbranched at the 

lower and median nodes, branching toward the upper nodes; 

internodes hollow, glabrous, the lower ones with a sheath only 

or with a rudimentary blade; nodes dark, compressed, glabrous. 

Leaves 3-7 per complement; sheaths strongly ribbed, with the 

margins shortly ciliate, otherwise puberulous; auricles membranous; ligule membranous, -3 mm long, shortly ciliate at the 

tip; pseudopetiole dark brown, -0.4 cm long, short-to 
SMITHSONTAN CONTRIBUTIONS TO BOTAW 

FIGURE 35.-Detail of female anthecium, Olyra fatnanquareana: a, apex of the anthecium, ventral side (x 100); 

b, detail of excavations (X 200); c, base of the anthecium showing stipe, dorsal side (x 40); d, base of the 

anthecium showing stipe, ventral side (X 40). All micrographs of Kubitzki ef al. 79-252 (US). (Reduced to 77% 

of indicated magnifications for publication.) 

long-hispid on both surfaces; blades lanceolate, flat, 13-25 

cm long, 3-5.5 cm wide, narrowing to an asymmetric base, 

the apex acuminate and aymmetric, glabrous, the margins 

smooth, scabrous toward the apex, the midnerve conspicuous, 

the lateral nerves anastornosing. Inflorescences paniculiform 

and numerous (5-7), borne at the uppermost nodes, longexserted, peduncles longitudinally ridged, shortly hispid, 

panicles falling completely at maturity, only the peduncles 

remaining, contracted, formed by a single branch (occasionally 

two conjugate branches present), each branch with the axis 

flattened or triquetrous, with male spikelets shortly pedicelled, 

spaced and alternate in groups of 4 below, and one terminal 

female spikelet per branch; axis shortly hispid, pedicels of the 

male spikelets short, hispid, those of the female spikelets 

thickened, shortly hispid. Female spikelets lanceolate, longaristate, 35 mm long, 4.2 mm wide, the glumes subequal, 

scaberulous to shortly hispid on the outer surface, densely 

hispid toward the apex on the inner surface, the anthecium less 

than '/2 the length of the spikelet; lower glum 9-13-nerved 

with transverse veinlets; upper glum 9-1 l-nerved with 

transverse veinlets; stipe conspicuous at the base of the 

anthecium, -1-1.2 mm long; anthecium lanceolate, 13-13.2 

mm long, 3.5-3.8 mm long, glabrous, pitted, with rounded 

excavations over the entire surface, mottled at maturity; lemma 

5-nerved. Caryopsis fusiform, brownish, 9 mm long, 2.5 mm 

wide; hilum linear, as long as the caryopsis. Male spikelets 

lanceolate, acute, 5 mm long, 1 mm wide, shortly hispid on the 

outer surface, smooth and shining on the inner surface; lemma 

3-nerved, palea 2-nerved; anthers 1.8 mm long. 

DISTRIBUTION.-Known only from the type locality, a dense 

forest along the Xi0 Negro in Amazonas, Brazil (Figure 11); 

Prance el al. I5797 (NY, US), from the same locality, is an 
hUMBER 69 61 

immature plant that may also be referable to this species. 

DISCUSSION.-Olyra tamanquaream resembles 0. obliquifolia, from which it differs in having panicles with but a single 

(occasionally two) branch present with only one terminal 

female spikelet per branch, and in the larger female spikelets. 

22. Olyra taquara Swallen 

FIGURES 34,36,31 

Olyra fuquuru Swallen, 196636. [Type: “In the US. National Herbarium, No. 

2434254, collected in swamp forest at Ribeao Grande, Municipio Jatai, 

Goias, Brazil, July 25, 1956, by A. Mucedo ( No. 4586).” Holotype, US 

sheet no. 2434254; isotype, US sheet no. 2434255.1 

Robust erect perennials in dense clumps of as many as 30 

culms from a thick creeping short-rhizomatous base. Culms 

up to 3.5 m tall, unbranched at the base, and branching toward 

the upper nodes; internodes cylindrical, glabrous, hollow and 

smooth, up to 1 cm in diameter, the basal internodes with a 

sheath only or with a rudimentary or reduced blade; nodes 

conspicuous, flared, dark brown and glabrous to shortpubescent, formed by the thickened basal rim of the sheath 

above and the thickened upper rim of the internode below. 

Leaves with sheaths strongly ribbed, glabrous to pubescent 

with glassy, caducous and brownish hairs all over the surface, 

pale green and strongly mottled with dark purple spots; ligule 

membranous, brownish, 0.7-1.4 mm long; pseudopetiole 

brownish, broad, flat, 0.3-0.4 cm long, shortly pubescent, 

usually recurved and positioning the blade downward; blades 

ovate to oblong-lanceolate, 21-32 cm long, 5.6-8 cm wide, 

narrowed abruptly to an acuminate and symmetric apex, 

cordate and symmetric basally, pale to purplish around the 

midnerve at the juncture with the pseudopetiole, glabrous on 

the adaxial surface, the abaxial surface glabrous to sparsely 

pubescent with short, appressed hairs and covered with 

FIGURE 36.-Detail of female anthecium, Olyru fuquuru: u, apex of the anthecium, dorsal side (x 50); b, apex 

of the anthecium, ventral side (x 50); c, detail of the surface of the lemma showing excavations (X 100); d, 

detail of the base of the anthecium (X 50). Micrographs u, d of Irwin and Soderstrom 5873 (US), b,c of Richardr 

6804 (US). (Reduced to 77% of indicated magnifications for publication.) 
62 SMITHSONIAN CONTRIBUTIONS TO BOTmY 

FIGURE 37.-Field photo of Olyra taquara (photograph by TR. Soderstran, Brazil, 1964), with Brazilian 

botanist Romeu Belem. 

brownish, caducous hairs toward the base, about the same color 

as the upper one (purplish in young leaves), the margins 

scabrous. Inflorescences paniculiform, borne from the uppermost nodes, 10-23 cm long, 10-30 cm wide, the lower 

branches whorled, -13-20 cm long, the upper ones alternate, 

ascending or spreading from the axis, the panicles exserted on 

long glabrous peduncles, these becoming sparingly pubescent 

toward the panicle; axis strongly angled and sparingly pilose, 

the branches sparingly pilose and flattened on one side, the 

axils of the branches pilose, each branch with male spikelets 

in pairs, one short- and the other long-pedicellate and to several 

on fine tertiary branchlets all along the branches: the terminal 

female spikelets 2 to 4 on each branch. Female spikelets 

fusiform, 16-20.5 mm long, 2.2-3 mm wide, pale green to 

stramineous, mottled with purple spots, shining, the glumes 

subequal and subulate to aristate, longer than the anthecium; 
NUMBER 69 63 

lower glum aristate, 5-nerved with transverse veinlets, the 

inner surface shortly and densely pilose, the outer surface 

scabrous toward the apex, otherwise glabrous: upper glume 

12.6-17.7 mm long, subulate to aristate, 5-nerved with some 

transverse veinlets, with a pubescence similar to that of the 

lower glume; anthecium fusiform, 8.4-9.2 mm long, 2-2.7 

mrn wide, whitish, indurate, irregularly pitted with small 

excavations; lemma 5-nerved. Caryopsis fusiform, brownish, 

6.2 mm long, 2.6 mm wide; hilum 5.6 mm long, almost 

reaching the entire length of the caryopsis. Male spikelets 

lanceolate, purplish, 10.4-14.1 mm long, 0.9-1.3 mm wide, 

aristate, glabrous: lemma awned, 3-nerved: palea 7.8-10.3 

mm long, 2-nerved, glabrous. 

DISTRIBUTION.-Found in the shade of gallery forests in 

central Brazil (Figure M), from 300 to lo00 m elevation. 

ADDITIONAL SPECIMENS SEEN.-BRAZIL, DISTRITO FEDERAL: Colegio Agricola de Brasilia, Pires and Mattos 9865 

(US), Mattos and Heringer 351 (RB, US); 35 km E of Brasilia, 

Irwin and Soderstrom 5420 (US); Brasilia, terrenos do Country 

Club, Sucre 446 (RB, US); Parque Municipal de Gama, -20 

km S of Brasilia, Irwin and Soderstrom 5873 (F, MO, US): 

slopes of Chapada da Contagem, -20 km E of Brasilia, Irwin 

and Soderstrom 5203 (US). GoIAA: Ribeiab Grande, Macedo 

4580 (US): Rio da Prata, vicinity of Posse, Irwin et al. 14569 

(F, MO, RB, US). MATO GROSSO: Serra do Roncador, vicinity 

of Chavantina, margin of Rio Mortes, Prance et al. 59092, 

59108 (F, MO, US, WIS); Ponte de Pedra, Kuhlmann 1869 

(IAN, RB); -270 km N of Xavantina, 12’54’S, 51’52W, Ratter 

et al. 1505 (US): 5 km W of the Rio Suia-Missu Ferry, -290 

km N of Xavantina, Ratter et al. 1425 (US): 260 krn along new 

road NNE of village of Xavantina, Eiten and Eiten 8615 (US). 

MATO GROSSO DO SUL -5 km N of Xavantina, Irwin and 

Soderstrom 6327 (US): Base Camp Square, 12”49’S, 51’46W, 

Richards 6804 (RB, US). MINAS GERMS: Horto Florestal de 

Paraopeba, Heringer 9171 (UB, US). PARA: Serra do 

Cachimbo, Pires et al. 6261 (IAN). 

DIscussIoN.-This species resembles Olyra ecaudata of the 

tropical forests of northern South America and 0. standleyi of 

Venezuela, Panama, and Costa Rica. In all three, the leaf blades 

are conspicuously symmetric at base (and often subcordate 

with a small purple spot), while the female anthecium is 

completely pitted with small excavations over its entire surface. 

Also, the whole plant when in flower, between May and 

September, produces numerous inflorescences but few leaves. 

Soderstrom examined large clumps near Brasilia in 1983 and 

found no signs of flowering on the culms: as these were old, 

mature culms it seemed apparent that flowering is not annual 

and may be a one-time event ending with death of plant. This 

phenomenon should be studied in 0. ecaudata, 0. standleyi, 

and 0. tuquara. 

The common name is recorded in Brazil as “taboquinha.” 

The chromosome number is 2n = 20 (Gould and Soderstrom, 

1967). 

23. Olyra wurdackii Swallen 

FIGURFS 11,38 

Olyra wurahckii Swallen, 196635. [Type: “Type in the US. National 

Herbarium, No. 2307269 and 2307270, collected at edge of open rock on 

crystallime dome on right bank of Rio Siapa just below Raudal Gallineta, 

Territory Amazonas, Venezuela, altitude 300-500 m, July 21, 1959, by John 

J. Wurahck and L. S. Adderley (No. 43540).” Holotype, US sheet nos. 

2307269.2307270.1 

Perennial in cespitose clumps. Culms 1-4 m tall, branching 

at the upper nodes, erect: internodes hollow, short-pubescent: 

nodes thickened, conspicuous, densely pilose with short, 

whitish, appressed hairs. Leaves with sheaths greenish, 

short-pubescent, the borders ciliate toward the upper portion, 

otherwise glabrous; ligule membranous, short-ciliate or not at 

the tip, 4-8 mm long, brownish; pseudopetiole short, dark 

brown, glabrous; blades lanceolate, 26 cm long, 5.2 cm wide, 

the apex symmetric, subulate, with a symmetric, subcordate 

base, glabrous on both surfaces, glaucous (to purplish near the 

base), the midnerve inconspicuous, the basal borders ciliate. 

Inflorescences paniculiform: panicles axillary and terminal at 

the upper nodes, -30 cm long and 20-25 cm wide, 

many-flowered, the branches divergent to ascendent, the lower 

ones whorled, the upper alternate to opposite, each branch with 

many male spikelets below and 2-5 terminal female spikelets 

above: pedicels of the male spikelets short, disposed in short 

tertiary branchlets; axis of the panicles longitudinally ridged, 

densely hirsute, the axis of the branches triquetrous, hispid, 

purplish, pedicels of the female spikelets thickened, hispid. 

Female spikelets lanceolate, acuminate, 7-7.5 mm long, 

2.2-2.5 mm wide, the glumes subequal, densely to sparsely 

hispid on the dorsal surface to glabrescent, densely pilose 

toward the apex on the ventral surface: lower glume acuminate, 

9-nerved, with transverse veinlets: upper glume acuminate, 

7-7.5 mm long, 7-9-nerved, with transverse veinlets; 

anthecium ovoid, acute, 5.5 mm long, shortly stipitate, smooth 

and shining, whitish; lemma with long prickle hairs toward the 

basal margins: palea with short prickle hairs and numerous 

bicellular, bottle-like microhairs on the upper portion. Male 

spikelets 8.9-11.5 mrn long, 0.8-1 mm wide, greenish to 

purplish: lemma 3-nerved, scabrous on the dorsal surface, 

glabrous on the ventral one, long-aristate, the awn 2.5-4 mm 

long; palea 7.5-8.8 mm long, 2-nerved, glabrous, shortaristate; anthers 3,4-4.3 mm long. 

DIsTRIBuTIoN.-Known only from two collections from the 

Amazonian region of Venezuela and Brazil (Figure 11). 

Manaus-Porto Velho Highway, km 246,4 km S of Igapo Acu, 

Prance et al. 20531 (US). 

DIscussIoN.-The affinities of 0. wurdackii are not clear. 

The species is unusual in that the male spikelets are larger than 

the female, and the female antheciurn is completely smooth 

and shining with conspicuous bicellular microhairs and prickle 

hairs at the apex of the palea. 

ADDITIONAL SPECIMENS SEEN.-BRAZIL, AMAZONAS: 
64 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 38.-Detail of female anthecium, Olyra wurdockii: a, apex of the anthecium, ventral side (x 100); b, 

detail of the apex of the palea showing prickle hairs and short bicellular microhairs (x 500); c, base of the 

anthecium, ventral side (x 50); d, detail of hairs on margin of the lemma (x 400). All micrographs of Prance et 

al. 20531 (US). (Reduced to 77% of indicated magnifications for publication.) 

Parodwlyra Soderstrom & Zuloaga, new genus 

Grumen herbaceum scandens, perenne. Culmi supra nodi 

medium profuse ramificantes profuse. Lamina ovato-lanceolata 

vel lanceolata, ad basim asymmeuica truncata, ad apicem 

asymmeuica, acuminata; pseudopetiolus brevis, pilosus; ligulae breves, membranaceo-ciliatae. Inflorescentiae paniculatae, 

laxae, diffusae, e nodis superionbus productae, ramis inferioribus masculis, ramis superioribus cum spiculis femineis infra 

et spiculis masculis supra (vel rarnis superioribus cum spicula 

feminea 1); pedicelli feminei filiformes. Spiculae 1-florae, 

unisexuales. Spicula fem'nea decidua; glumae 2 subequales, 

brevipilosae, quam anthoecium longiores, inflatae vel induratae; rachilla inter glumas incrassata, pulviniformis; gluma 

inferna 5-9-nervata amplectens; gluma superna 3-6-nervata; 

anthoecium ellipsoideum, induratum laeve, nitidum, apice 

acuto vel obtuso; lemma 5-nervatum tesselatum; palea 

2-nervata; lodiculae 3; stylus singularis; stigmata 2. Caryopsis: 

hilum lineare quam caryopsis 1/2-3/4 longius. SpicuIae 

masculae fusiformes, acuminatae, hispidae vel laeves; lemma 

3-4-nervatum; palea 2-nervata; stamina 3. 

Perennial monoecious herbs. Culms vine-like, arching and 

clambering, profusely branching from the median and upper 

nodes; internodes hollow, cylindrical; nodes thickened, pilose 

with remrse hairs. Leaves with sheaths glabrous to pilose; 

ligule membranous-ciliate, small; pseudopetiole short, pilose; 

blades ovate-lanceolate to lanceolate, flat, stiff, asymmetric and 

truncate at the base, tapering to an acuminate, asymmetric apex. 

Inflorescences borne from the uppermost nodes, paniculiform, 

the panicles lax and diffuse, the lower branches with male 

spikelets only, the upper ones with male spikelets below and 
NUMBER 69 65 

terminal female spikelets above or with female spikelets only; 

pedicels of the female spikelets not thickened, the female 

spikelets disarticulating below the glumes. Female spikelets 

ellipsoid, shortly pilose on the outer surface of the glumes; 

glumes subequal, inflated to indurate at maturity; rachilla 

prominent between the lower and upper glume and thickened, 

giving the spikelet a pulviniform shape at its base; lower 

glume 5-9-nerved with transverse veinlets, completely embracing the upper glume; upper glume 3-6-nerved with 

transverse veinlets; small stipe present at the base of the 

anthecium; anthecium, indurate, shorter than the glumes, 

smooth and shining, acute to obtuse apically, with or without 

conspicuous bicellular, bottle-like microhairs at the tip of the 

lemma and palea; lemma 5-nerved; palea 2-nerved; lodicules 

3; style 1, stigmas 2. Caryopsis ellipsoid, brownish, the hilum 

linear, 1/2-3/4 the length of the caryopsis. Male spikelets 

lanceolate, acuminate, hispid to glabrous; lemma 3-4-nerved; 

palea 2-nerved; anthers 3. 

TYPE SPECIES.-Parodiolyra ramosissima (Trinius) Soderstrom & Zuloaga. 

DIsTRIEUn0N.-Three species; low- and mid-elevation 

forests and savannas from Costa Rica south to Bolivia and 

Bahia, Brazil. 

DIscUssI0N.-Species of Olyra are erect plants that 

generally occur in shaded forest. They are generally unbranched at the lower nodes and have female spikelets on clavate 

pedicels in which the glumes are persistent (except for 0. 

micrantha). The glumes are not inflated and there is no swollen 

internode between them. The hilum is linear and extends for 

the full length of the caryopsis. 

Three species previously included in Olyra (as 0. lateralis, 

0. luetzelburgii, and 0. ramosissima) do not possess the above 

features but rather share a suite of unusual characters that 

clearly set them apart. We are establishing Parodiolyra to 

accomodate them. Parodiolyra includes decumbent, vine-like, 

clambering, or declining plants that branch and often root from 

the lower or from the median and upper nodes. They may grow 

in shaded forest but also often in full sunlight when enough 

moisture is available. The female spikelets fall entire, 

disarticulating below the glumes, and a conspicuous, thickened 

internode is present between the lower and upper glume in all 

species. There is also a prolongation of the rachilla between 

the upper glume and the anthecium, in the form of a slender, 

thin stipe. Perhaps the most striking feature of this new genus 

is the presence of a shortened hilum that extends for only '/z 

to 3/4 the length of the caryopsis, never reaching its entire 

length (as in Olyra and most other genera of the tribe Olyreae). 

In Parodiolyra, the shortened hilum, female spikelet 

disarticulating entire from a filiform pedicel, and the conspicuous, thickened internode between the glumes are reminiscent 

of Raddiella Swallen and Diandrolyra Stapf, rather than Olyra. 

However, plants of Diandrolyra differ from Parodiolyra in 

being erect with spike-like inflorescences borne on specialized 

culms and with the male spikelets bearing only two stamens. 

Raddiella, a genus that inhabits moist areas such as savannas, 

wet cliff faces, and bases of waterfalls, also has decumbent 

culms, but it differs from Parodiolyra by the disposition of the 

inflorescences and male and female spikelets within the plants. 

In Parodiolyra the inflorescences are paniculiform, borne only 

from the uppermost nodes, lax, diffuse, and many-flowered, 

with the lower branches bearing male spikelets and the upper 

branches bearing male spikelets below and female spikelets 

above. In contrast, in Raa'diella there are numerous axillary 

inflorescences borne at the median and upper nodes of the 

plants, and these panicles are small, few-flowered, and usually 

bear female spikelets only, with the male spikelets confined 

to terminal panicles. 

The species of Parodiolyra are also characterized by their 

inflated, membranous glumes (indurate at maturity in P: 

lateralis), with the lower member amplectent. The female 

anthecium is smooth and shining, with the tip of the palea with 

rounded excavations and the upper portion of lemma and palea 

covered by conspicuous bottle-like, bicellular, secretory 

microhairs in P. luetzelburgii and P. ramosissima (completely 

glabrous in P. lateralis). These hairs contain a resin that is 

secreted when the anthecium is mature and become dark, 

making the anthecium viscous. In P. lateralis the glumes are 

indurate and purplish and shining at maturity, and Davidse 

(1987) has suggested that their dark coloration might attract 

birds, while the induration of the glumes might serve as 

mechanical protection for the anthecium and its enclosed 

caryopsis. 

Key to the Species of Parodiolyra 

1. Female spikelets at maturity black, indurate, 2-2.6 mm long, the anthecium 

glabrous, obtuse apically; leaf blades 2.5-5 cm long . . . . . . , , , . . . . . . . . . . . , . . . , , . . . . . , . . . , . , . . , 1. P. lateralis, new combination 

(widespread, Costa Rica to Bolivia and Brazil) 

1. Female spikelets at maturity light-colored, chartaceous, 3-4.6 mm long, the 

anthecium with conspicuous bicellular microhairs at the acute apex; leaf blades 

4-13 cm long. . . , . . . . . . . . . . . . . , , . . . . . , . . . . . . . . . .2 
66 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

2. Female spikelets 3-3.6 mm long, 1.5-1.7 mm wide; male spikelets 4.1-5 mm 

long; panicles with the lower branches whorled; ovary glabrous; lower margins 

of the blades long-ciliate . . . . . . . . 2. €! luetzelburgii, new combination 

(northern South America from Venezuela 

to Maranhb and Mato Grosso, Brazil) 

2. Female spikelets 5-5.8 mm long, 1.9-2.8 mm wide; male spikelets 3.5-4.3 mm 

long; panicles with the lower branches alternate or opposite; ovary pilose 

towards the summit; lower margins of the blades glabrous or scabrous, not 

ciliate . . . . . . . . . . . . . . . . . . 3. P. ramosissima, new combination 

(Bahia, Brazil) 

1. Parodiolyra lateralis (Presl ex Nees von Esenbeck) 

Soderstrom & Zuloaga, new combination 

FIGURES 39,40,44 

Panicwn laferale var. a Presl ex Nees von Esenbeck, 1829:213-214. [Type: 

“In montibus Orinocensibus lectam.” Holotype, PR, not seen, fragment of 

holotype and photograph, US sheet no. 2877929. Nees saw this specimen 

in the Haenke Herbarium in PR where one of the labels gives as the locality 

“Peruanae montanae huanoccenses [Department of Huanuco, Peru],” the 

latter word of which Nees misread as “Orinocences.”] 

Panicwn laterale J.S. Presl, 1830:305. [Type: “Hab. in Peruviae montanis 

huanoccensibus.” By the habitat that Presl cites, he clearly indicates that his 

I? laferale is the Haenke specimen from Peru, var. a of Nees von Esenbeck]. 

Olyra sarmenfosa Doell in Martius, 1877:319. [Type: “A cl. Poeppig (n. 3055) 

in Brasilia Amazonica, et in republica Peruviana a cl. Dombey (n. 146) lecta.” 

Syntypes, B, not seen, fragments of the syntypes Poeppig 3055, US sheet 

no. 2877929, Dombey 146, US sheet no. 2877929.1 

OIyra laferalk (Presl ex Nees von Esenbeck) Chase, 1908:179. [Type: Based 

on Panicwn laterale var. a Presl ex Nees von Esenbeck, 1829:213-214.1 

Roddiella frwafa Swallen, 194839. [Type: “Guyana, Waratuk Falls, Potan, 

River Gorge,” Maguire and Fanshawe 23035. Holotype, US sheet no. 

171359; isotypes, NY, US sheet no. 1914990.1 

Sprawling perennial. Culms up to 8 m or more long, trailing 

and clambering on vegetation, branching profusely at the 

median and upper nodes; internodes sulcate, hollow, glabrous, 

the basal ones elongated, 11-20 cm long and with sheaths 

only or sheaths with rudimentary blades; nodes thickened, 

formed by the thickened basal rim of the sheath above and the 

thickened upper rim of the internode below, pilose with whitish 

retrorse hairs. Leaves with sheaths overlappping, ribbed, 

sparsely pilose with retrorse hairs, more densely so along the 

margins, long-ciliate or not at the summit; auricles present, 

long-ciliate or not; ligule membranous-ciliate, -0.2 mm long; 

pseudopetiole, indurate, 1-2 mm long, pale to dark brown, 

densely pubescent on both surfaces; blades ovate-lanceolate, 

2.5-5 cm long, 0.5-1.3 cm wide, rather stiff, asymmetric and 

truncate basally, the apex asymmetric and apiculate, sparsely 

ciliate on the margins towards the apex and base, the rest of 

the surface scabrous or shortly pubescent to glabrous, the 

abaxial surface glaucous. Inflorescences paniculiform, borne 

profusely from the uppermost nodes, occasionally with 

panicles produced at the lower nodes, long-exserted, the 

peduncles 5-9 cm long, glabrous; panicles lax, pyramidal, the 

branches spreading, 1.2-3 cm long, 1.5-3.5 cm wide, the 

lower branches opposite or alternate and bearing male spikelets 

only or paired male spikelets (one long and the other 

short-pedcelled), the upper branches alternate and with female 

spikelets only, or some specimens with panicles bearing only 

female spikelets; hxis and branches cylindrical, flexuous, 

scabrous to sparsely pilose, the mils of the branches pubescent; 

pedicels smooth, sparsely pubescent to glabrous. Female 

spikelets elliptical, acute, 2-2.6 mm long, -1.2 mm wide, 

greenish to purplish or black at maturity; glurnes subequal and 

longer than the anthecium, indurate at maturity and shining, 

the lower glume embracing completely the upper one; lower 

glume 5-7-nerved, scabrous to shortly and densely pilose, 

more densely so towards the base; upper glume 3-5-nerved, 

hispid to glabrous; anthecium obovoid, obtuse apically, 

indurate, whitish, smooth and shining, glabrous, shortly 

stipitate at the base; lemma 7-nerved; palea 2-nerved. 

Caryopsis ovoid, brownish, 1 mm long, 0.6 mm wide, the 

hilum linear, I12 the length of the caryopsis. Male spikelets 

lanceolate, acute, 3-4 mm long, 0.6-0.8 mrn wide, shortly 

pilose to glabrous, brownish to purplish, the glumes occasionally present; lemma 3-nerved; palea 2-nerved; anthers -2 mm 

long, purple. 

DISTRIBUTION.-This is the most common and widespread 

species of Parodiolyra. It is distributed from the mountains of 

Mesoamerica (Costa Rica) into South America, south along the 

eastern foothills of the Andes to Bolivia and east at the edge 

of the Guyana highlands to Surinam (Figure 44). The plants 

inhabit wet savannas, riverbanks, bluffs, and cliffs in full shade 

or sun, at elevations from 400 to 1800 m above sea level. 

ADDITIONAL SPECIMENS SEEN.-BOLIVIA, LA PAZ: Hacienda Simaco, sobre el camino a Tipuani, Buchtien 5333 (F, 

MO, NY, US); San Carlos, Buchtien 25 (US); Ticunhaya, Tate 

1079 (NY, US). 

BRAZIL, AMAZONAS: Rio Negro, Serra Curicuriari, Madison and Kennedy 6565 (US), Prance ef al. 16125 (MO, US). 

RORAIMA: NE of Mission station, Serra dos Surucucus, Prance 

et al. 10011 (MO, US). 

COLOMBIA, CHOCO: Cerca a1 Rio Atrato, en 10s alrededores de Quibdo, Molina and Barkley 19C142 (NY, US). 

CUNDINAMARCA: “Pipiral,” SE Of Quetame, Penneii 1685 

(NY, us). MAGDALENA: santa Marta, H.H.Smith 2149 (F, 

MO, US, WIS). META: Las Lagartijas, plateau between Rio 

Papaneme and Rio Duda, 11 km SW of Uribe, Fosberg 19512 
NUMBER 69 67 

FIGURE 39.-Parodiolyra laterulb: a, flowering branches with inflorescences (x 0.6); b, base of leaf blade (X 

3.7); c, female spikelet, view of lower glume side (x 15); d, female spikelet, view of upper glume side (x 15); 

e, female anthecium, dorsal view (x IS);$ female anthecium, ventral view (X 15); g-i, caryopsis (X 15); g, dorsal 

view; h, ventral view; i, side view; j, gynecium (x 15); k, male spikelet (x 15). Component a based on Skutch 

2254 (US); b based on Smith 2149 (US); c-f,k based on MacBride 4990 (US); g-i based on Buchtien 25 (US); 

j based on Steyermurk 89552 (US). All illustrations by G.B. Threlkeld. 
68 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 40.-Detail of female anthecium, Parodiolyra laferalir: a, apex, ventral side (x 200); b, apex, dorsal 

side (x 100); base, c, base, ventral side (X 200). All micrographs of Steyermark ei al. 92399 (US). (Reduced to 

77% of indicated magnifications for publication.) 

(US); Sierra de La Macarena, Central Mountains, Approach 

Ridge, Philipson and Idrobo 1877 (US), 

COSTA RICA, HEREDIA: Cordoncillal, Pittier 3641 (US); 

Cienaga de Agua Buena, Canas Gordas, Pittier 11008 (US). 

SAN JOSE: Helechales du General, Vallee du Diquis, Pittier 

12058 (US): vicinity of El General, Skutch 2254 (MO, US). 

PROVINCE UNKNOWN: Cienaga de Agua Buena, Pittier 11008 

(US). 

GUYANA: Mure-Mure Savanna, Kaieteur Plateau, Cowan 

and Soderstrom 2141 (US), 2165 (US). 

PANAMA, c~CLE: El Valle, Soderstrom 2015 (US). 

HERRERA: 18 km W of Las Minas, trail to top of Alto Higo, N 

slope of Cerro Alto Higo, Hamme14213 (MO, US). PANAMA: 

Cerro Azul, Dwyer 1504 (MO, US), Calderon 2088 (US); 

Cerro Jefe, Calderon and Dressler 2139 (US), Witherspoon 

8498 (MO, US); Cerro Campana, Calderon and Dressler 

2129 (US), Calderon 2092 (US); 7 mi 111 km] N of Cerro 

Azul on road to Cerro Jefe, Blum et al. 1789,1809 (US). 

PERU, HUASUCo: Andes, Poeppig 976 (F, US). PASCO: 

Vilcabamba, hacienda on Rio Chinchao, Macbride 4990 (F, 

US). SAN MARTIN: San Roque, Williams 7111 (F, US). STATE 

UNKNOWN: Andes, Poeppig 976 (US). 

SURINAM: Tafelberg, West Grace Falls, Arrowhead Basin, 

Maguire 24501 (F, NY). 

VENEZUELA, AMAZONAS: entre Shimada-Wochi y el alto 

Rio Matacuni, Sierra Parima, Huber and Colchester 8390 

(MO, US). BOLIVAR: 3 km S of El Puajl, Liesner and Holst 

18818 (MO, US); Gran Sabana, between Mision of Santa 

Teresita de Kavanayen, NW of Rio Karuai, Steyermark 59380 

(F, NY, US); between Rio Karuai and Salto de Iraba-raima 

along Rio Karuai, at SW base of Ptari-tepui, Steyermark 60705 

(F, NY, US); Chimana Massif, along base of southeast-facing 

sandstone bluffs of Chimanta-tepui (Toroo6-tepui), Steyermark 

75506 (F, NY, US); Steyermark Falls, on upper reaches of Rio 

Tirica, Steyermark 76004 (F, NY, US); Sierra de Lema, 

cabeceras de Rio Chinacan 80 km SW of El Dorado, 

Steyermark 89552 (F, NY, US): Cerro La Danta, NW of Cerro 

Venamo, Steyermark and Nilsson 32 (F, NY); Chimanta 

Massif, Amuri-tepui, Steyermark and Wurdack 1348 (MO, 

NY, US): Cerro Venamo, subiendo el Rio Venamo, Steyermark 

et al. 92399 (US); Rio Venamo, Steyermark 92778 (US). 

DISTRITO FEDERAL: Cerro Naiguaa, arriba del pueblo de 

Naiguata, Lomas de Las Delicias, 9-12 km SW de Hacienda 

Cocuizal, Steyermark 91998 (US). ZULU: Perija, Gines 1517 

(US). 

DIscussI0N.-In flower all year. One group of specimens 

is peculiar in that all panicles of the plant contain only female 

spikelets. These compare favorably with the type specimen of 

Raddiella truncata Swallen, however close examination of the 

spikelets reveals that they are like those of Parodiolyra 

lateralis and that the flowering system is that of Parodiolyra. 

In Raddiella, the flowering system of the lower nodes produces 

peduncles, each of which bears only 2-3 female spikelets, 

while the flowering system of the terminal node produces only 

panicles bearing male spikelets. 

The chromosome number has been recorded as n = 18 

(Davidse and Pohl, 1978). 

FIGURE 41 .-Parodiolyra luefzelburgii: a, habit; b, male spikelet; c, male 

lemma; d, male palea; e, male fl0wer.L female spikelet; g, longitudinal section 

through pulvinus at base of female spikelet; h, lower female glume. dorsal 

view; i, lower female glume, ventral view; j, upper female glume, dorsal view; 

k, upper female glume, ventral view; 1, female anthecium, dorsal view; m, 

female anthecium, veitral view; n, female flower. 0, caryopsis, dorsal view; 

p, caryopsis, ventral view. All based on Irwin et al. 55516 (US). All 

illustrations by Vladimiro Dudas. 
NUMBER 69 69 
70 SMITHSONIANCONTRIBUTIONS TO BOTANY 

2. Parodiolyra luekelburgii (Pilger) 

Soderstrom & Zuloaga, new combination 

FIGURES 41,42,430-c, 44c 

Olyra luelzelbwgii Pilger, 1930:1049. [Type: “Nord-Brasilien: lgarape, 

Caican; auf Sand (Oktober 1927), von Luetzelbwg, Brasilianische 

Grenzexpeditionen General Rmdcm, n. 21354. ” Holotype, B, not seen, 

fragment of the holotype, US sheet no. 2877941; isotypes, IAN, R.] 

Cespitose, short-rhizomatous perennial. Culms 70- 160 cm 

tall, erect and geniculate, climbing and leaning on the 

vegetation, unbranched towards the base, densely branching 

at the median and upper nodes; internodes cylindrical, hollow, 

glabrous, the basal internodes with a sheath only or sheaths 

with a rudimentary or reduced blade: nodes dark, thickened, 

pubescent with remrse whitish hairs. Leaves with sheaths 

shorter than the internodes, densely pilose all over the surface 

to pilose towards the margins and apex, the rest of the surface 

glabrous, usually mottled with green spots: ligule membranous, 

short-ciliate at the apex, -0.4 mm long: pseudopetiole 

stramineous, densely villous, -2 mm long: blades ovatelanceolate, 8-11 cm long, 1.6-3.1 cm wide, stiff, glaucous, 

flat, basally truncate and slightly asymmetric, the apex 

asymmetric; both surfaces densely hispid to scabrous or the 

adaxial surface scabrous and the abaxial one densely pubescent: abaxial surface silvery blue green, the margins longciliate towards the base, otherwise scaberulous, cartilaginuous 

at the tip: midnerve inconspicuous. Inflorescences paniculiform, borne from the uppermost nodes, the peduncles ribbed, 

elongate, glabrous: panicles pyramidal, lax, 5-8 cm long, 

4.5-8 cm wide, the lower branches whorled, with male 

spikelets only (these arranged in pairs or in groups of 3 on 

long pedicels), the upper branches alternate to opposite, with 

male spikelets below and terminal female spikelets above or 

only with female spikelets: axis scabrous to hispid, longitudiFIGURE 42.-Detail of female anthecium, Porodiolyru luetzelbwgii: u, apex, dorsal side (x 200); b, apex, ventral 

side (X 200); C, detail of bicellular microhair on apex of the palea (X 1ooO); d, base, ventral side (x 200). All 

micrographs of Bfuck 54-16672 (IAN). (Reduced to 77% of indicated magnifications for publication.) 
NUMBER 69 71 

nally ridged, the branches cylindrical, purplish, the axils of the 

branches pilose; pedicels smooth, hispid; axillary panicles 

similar to the terminal one. Female spikelets ellipsoid, whitish 

to purplish, inflated, 3-3.6 mm long, 1.5-1.7 mm wide; 

glumes subequal, acute, scabrous with short, retrorse hairs, 

hirsute or not with stiff hairs towards the apex and with short 

hairs towards the base, separated by a conspicuous, thickened 

internode, a stipe present at the base of the anthecium; lower 

glume chartaceous, strongly 7-9-nerved, completely embracing the upper glume; upper glume chartaceous, strongly 

5-7-nerved; anthecium lanceolate, shorter than the glumes, 

1.9-2.5 mm long, 1-1.2 mm wide, whitish to stramineous, 

smooth and shining, with numerous bicellular bottle-like 

microhairs at the acute apex; lemma 5-nerved, with rounded 

excavations towards the apex; palea 2-nerved, with rounded 

excavations towards the apex and on the inner margins; ovary 

glabrous. Curyopsis 1.4 mm long, 0.9 mm wide, the hilum -3/4 

as long as the caryopsis. Mule spikelets lanceolate, 4.2-5.1 

mm long, 0.8 mm wide, purplish, acuminate, hispid; lemma 

3-4-nerved, acute; palea 4.1-5 mm long, 2-nerved, sparsely 

hispid on the middle portion; anthers 3-3.2 mm long. 

DISTRIBUTION.-Northern South America from western 

Venezuela east and south to Maranhao and Mato Grosso, 

Brazil (Figure 44), growing in open areas, on sandy slopes, 

ridges, and quebradas, from 100 to 600 m above sea level. 

ADDITIONAL SPECIMENS SEEN.-BRAZIL, AMAPA: Colonia 

do Torrao, Pires and Cavakante 52636 (MO, US). AMAZONAS: 

Maues, across from Guarana factory, Campbell et al. P21984 

(US). MARANHAO: Itapicunu, Black et al. 54-16672 (IAN). 

MAT0 GROSSO: Municipio de Colider, 30 km de Guarantan, 

FIGURE 43.-Detail of female spikelets of Parodiofyra species. R luetzelburgii: a, apex of the spikelet, ventral 

side, showing cucullate apex (X 70); b, apex of the lower glume (x 70); c, detail of the surface of the lower glume 

(X 400). P. ramosirsima: d, detail of the surface of the lower glume showing simple papillae, prickle hairs, and 

bicellular microhairs (X 400). Micrographs a-c of Wurduck and Adderley 43514 (US); d of Calderon and 

Pinheiro 2198 (US). (Reduced to 77% of indicated magnifications for publication.) 
72 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 44.-Distribution map of Purodiolyru species. 

(IAN); Rio Capim, Aproaga, Black 48-2385 (IAN, NY, US). 

RORAIMA: Igarape Arapapo, Luetzelbwg 21481 (R); Valle do 

Croe-Cry, Luetzelburg 21359 (R). 

FRENCH GUIANA: Montagnes de la Trinite, de Granville 

et al. 6075 (MO). 

SURINAM: Wilhelmina Gebergte, 5 km E of confluence 

of Oost and Lucie riviers, 3"36'-3"41'N, 56O30'-56O34W, 

Irwin et al. 55516 (MO, NY, US); trail line Lucie RivierWilhelmina Mts., slope of unknown peak near km 14, Schulz 

VENEZUELA, AMAZONAS: -33 km a1 NE de Puerto 

Ayacucho, Huber 2732 (MO); on right bank of Rio Siapa, 8 

km below Raudal Gallineta, Wurdack and Adderley 43514 

10462 (US). 

(MO, NY, US). BOLIVAR: Morichal El Caballo, -225 km S of 

Caicara del Orinoco by road, Hoist and van der Wersf2610 

(MO, SI); rapids of Rio Apacara, l/4 mile down stream from 

the mouth of Rio Abacapa to the mouth of Rio Abacapa, 

western side of Ap5cara-tepui, Steyermark 74702 (F, NY, US); 

Sierra Auraima, en la margen terminal norte sobre el margen 

oeste del Rio Paragua, en la zona del raudal de El Perro, 

Steyermark 90817 (F, NY, US). TACHIRA: Monraila de 

Guafitas, just W and N of El Pinal, Steyermark et al. 119502 

(MO, US). 

DIscussIoN.-Flowers throughout the year. This species 

shows a closer resemblance to P. ramsissima than to P. 

lateralis. 
NUMBER 69 73 

3. Parodiolyra ramosissima (Trinius) Soderstrom & Zuloaga, 

new combination 

FIGURES 43d, 44,45,46 

Olyra ramsirsirno Trinius, 1834:28 [page 116 in published form, 18351. 

[Type: “Bahiens,” Riedel s.n. Holotype, LE, not seen, fragment of the 

holotype, US: isotype, GH.] 

Olym blancherii Mez, 1917:46. [Type: “Brasilia, Bahia, loco accuratiore non 

indicato (Blancher n. 273O).”Holotype B, not seen, fragment of the holotype, 

US.] 

Cespitose perennial. Culms 5-30 per clump, 30-150 cm 

tall, geniculate at the lower nodes, the young culms erect, 

unbranched at the lower nodes, profusely branching at the 

median and upper nodes, climbing and leaning on the 

vegetation; internodes cylindrical, hollow, maroon to purplish, 

glabrous, the basal ones with sheaths only or sheaths with 

rudimentary blades; nodes thickened, densely pilose with long 

retrorse whitish hairs. Leaves with sheaths strongly ribbed, 

greenish to purplish, glabrous to pubescent with retrorse hairs, 

the margins ciliate; ligules membranous at the base and 

short-ciliate at the tip, -0.3 mm long; pseudopetioles 

stramineous to purplish, scabrous to densely pubescent with 

long, whitish hairs, -2-3 mm long; blades lanceolate, 3.6-13 

cm long, 0.9-1.7 cm wide, ascending, stiff, narrowed and 

asymmetric basally, tapering abruptly to an apiculate tip, 

glabrous to strongly scabrous, the margins scabrous, the 

midnerve prominent towards the base, the abaxial surface 

greenish to somewhat purplish or dark maroon. Inflorescences 

numerous, long-exserted, borne from the uppermost nodes, 

3-8 cm long, 1-4 cm wide, the lower branches alternate to 

opposite and with paired male spikelets (one short and the other 

long-pedicelled), the upper branches alternate and with male 

spikelets below and one terminal female spikelet above or with 

female spikelets only; an’s longitudinally ridged, scaberulous 

to short-pubescent, the branches cylindrical, puberulent to 

scabrous, the axils of the branches pubescent; pedicels of the 

female spikelets similar to those of the male spikelets, 

scabrous; axillary panicles similar to the terminal one. Female 

spikelets broadly ellipsoid, inflated, 5-5.8 mm long, 1.9-2.8 

mm wide, whitish, obtuse to acute; glumes exceeding the 

anthecium in length, shortly hispid to scabrous with short, 

retrorse hairs, glabrous and shining on the inner surface, the 

internode conspicuous and thickened between the lower glume 

and the anthecium; lower glume 4.1-5.1 mm long, 7-!+nerved 

with transverse veinlets; upper glume 3.9-4.6 mm long, 

5-6-nerved with transverse veinlets, acute; anthecium ellipsoid, 3-3.3 mm long, 1.5-2 mm wide, acute, whitish to 

olivaceous at maturity, smooth and shining, with numerous 

bicellular, bottle-like microhairs at the apex, shortly stipitate 

at its base, the stipe -0.3 mm long.; lemma 5-nerved, with 

rounded excavations towards the tip; palea Znerved, with 

rounded excavations towards the tip; ovary pilose towards 

summit. Caryopsis ellipsoid, brownish, 2-2.5 mm long, 

1.4-1.8 mm wide; hilum linear, 1.7-1.9 mm long, 3/4 the 

length of the caryopsis. Male spikelets lanceolate, 3.5-4.3 

mm long, 0.7-1 mm wide, purplish, densely hispid with long, 

whitish hairs; lemma 3-nerved, acute; palea Znerved, pilose 

in the middle portion to glabrous; pistilode usually present. 

DISTRIBUTION.-Endemic to Bahia, Brazil (Figure 44), 

where it grows on sandy soils within or at the edges of open 

forests, frequently in disturbed areas, at elevations below 150 

m. 

ADDITIONAL SPECIMENS SEEN.-BRAZIL, BAHIA: 3 km w 

of Itacare, Soderstrom et al. 2216 (CEPEC, US); rodovia 

Camamu-Ibirapitanga, 11 km SW of Camamu City, Calderon 

and Pinheiro 2237 (CEPEC, US); Reserva Biologica Pau 

Brasil, 16 km W of Porto Seguro, Calderon and Pinheiro 

2198 (CEPEC, US), Soderstrom et al. 2189 (CEPEC, US), 

Eupunino 19 (US), Huge 67 (CEPEC, US); ramal para 

Mogiquicaba, com entrada no km 23 da rodovia BelmonteItapebi, Mattos Silva and Brito 974 (RB, US); -18 km W of 

Valen`ca on road to Zona Orobo, Soderstrom et al. 2162 

(CEPEC, US); between Itapebi and Belmonte, 10-12 km E 

of the Estacao Experimental of CEPLAC, Soderstrom er a1 

2136 (CEPEC, US); rodovia Camamu a Ibirapitanga, Pinheiro 

1847 (MO, US); km 10, rodovia Porto Seguro para Eunapolis, 

Calderon 2050 (US); Marau, Belem 1810,1821 (CEPEC, US): 

entre 0s kms 45-46 da rodovia Eunapolis-Porto Seguro 

(BR-367), Mori et al. 10960 (CEPEC, MO, US); 17 km W of 

Pbrto Seguro, Mori et al. 10797 (CEPEC); 4 km N of OlivenGa, 

on the road from Olivenca to Ilheus, Mori et al. 13678 

(CEPEC, MO); Nova Vicosa, Guimcaaes 1022 (RB); estrada 

Olivenca-Maruim, krn 4-5, Mantone 1006 (RB); antiga 

rodovia que liga Estacao Ecologica de Pau Brasil a Santa Cruz 

da Cabrala, kms 2 a 9, Mattos Silva and Brito 898 (RB); 

rodovia Canavieiras/Camaca (BA 270), a 20 km W de 

Canavieiras, dos Santos and Mattos Silva 3279 (MO); without 

locality, Calderon 2448 (US). 

DISCUSSION.-In flower between January and October. Until 

1966, when Romeu Belem recollected it in Bahia, P. 

ramosissima had been known only from Blanchet and Riedel 

type collections made there in the 19th century. 
74 SMFHSONIANCONT!UBUTIONS TO BOTANY 
NUMBER 69 75 

FIGURE &.-Female anthecium of Parodiolyra rmosksima: a, apex, ventral side (x 150); b, apex, dorsal side 

(x 100); c, base, ventral side (X 100); d, detail of microhair on top of the palea (x 1500). Micrograph a of Mori 

et al. 10960 (US), b-d of Calderon and Pinheiro 2198 (US). (Reduced to 77% of indicated magnifications for 

publication .) 

FIGURE 45.-Parodiolyra ramosksima: a, habit; b,c, male spikelet; d, male 

palea; e, male flower, f, branchlets bearing female spikelet and male spikelet 

(fallen); g, female spikelet, view of back of lower glume; h, female spikelet, 

view of back of upper glume; i, longitudinal section through pulvinus at base 

of female spikelet; j, lower female glume, ventral view; k, upper female glume, 

ventral view; 1, female anthecium, dorsal view; m, female anthecium, ventral 

view; n, female flower, 0, caryopsis, dorsal view; p, caryopsis, ventral view. 

Component a based an Calderdn 2050 (US), b-p based on Calderon 2448 

(US), both from Bahia, Brazil. All illustrations by Vladimiro Dudas. 
Appendix 1 

Names Included in Olyra Linnaeus 

(Names accepted here are in italics.) 

0. amapana Soderstrom & Zuloaga 

0. arundinacea Humboldt, Bonpland & Kunth = 0. latifolia 

0. brasiliensis Desvaux = 0. latifolia 

0. brevifolia Schumacher = 0. latifolia 

0. buchtienii Hackel 

0. caudata Trinius 

0. ciliatifolia Raddi 

0. corcovadensis Wawra = 0. glaberrima 

0. cordifolia Humboldt, Bonpland & Kunth = 0. latifolia 

0. cordifolia var. scabriuscula Doell in Martius = 0. latifolia 

0. cuneatifolia Desvaux = 0. ciliatifolia 

0. dimidiata Hochstetter ex Steudel = 0. caudata 

0. d’Urvillei Doell in Martius = 0. micrantha 

0. ecaudata Doell in Martius 

0. fasciculata Trinius 

0. filiformis Trinius 

0. glaberrima Raddi 

0. glabemma var. humilis (Nees von Esenbeck) Mez = 0. 

0. heliconia Lindman = 0. fasciculata 

0. hirsuta Trinius = 0. micrantha 

0. holttumiana Soderstrom & Zuloaga 

0. humilis Nees von Esenbeck 

0. humilis var. angustifolia Doell in Martius = 0. humilis 

0. humilis var. latifolia Doell in Martius = 0. humilis 

0. jwuuna Mez 

0. kegelii Mez = 0. longifolia 

0. latifolia Linnaeus 

0. latifolia var. arundinacea Grisebach = 0. latifolia 

0. latifolia var. glabriuscula Doell in Martius = 0. latifolia 

0. latifolia var. pubescens Doell in Martius = 0. latifolia 

humilis 

0. latifolia var. vestita Henrard in Amshoff & Henrard = 0. 

0. latispicula Soderstrom & Zuloaga 

0. longifolia Humboldt, Bonpland & Kunth 

0. longifolia var. grandifolia Doell in Martius = 0. longifolia 

0. longifolia var. parvifolia Doell in Martius = 0. longifolia 

0. loretensis Mez 

0. maranonensis Swallen 

0. media Desvaux = 0. latifolia 

0. micrantha Humboldt, Bonpland & Kunth 

0. micrantha var. decalvata Doell in Martius = 0. micrantha 

0. micrantha var. lanceolata Doell in Martius = 0. micrantha 

0. micrantha var. subvelutina Doell in Martius = 0. micrantha 

0. obliqua Desvaux = 0. glaberrima 

0. obliquifolia Steudel 

0. paniculata Swartz = 0. latifolia 

0. pittieri Hackel = 0. caudata 

0. pubescens Raddi = 0. latifolia 

0. retrorsa Soderstrom & Zuloaga 

0. scabra Nees von Esenbeck = 0. latifolia 

0. scorbiculata Schrader ex Nees von Esenbeck = 0. micrantha 

0. semiovata Trinius = 0. glaberrima 

0. semiovata var. pubescens Hackel = 0. glaberrima 

0. serniovata var. pubiflora Hackel = 0. glaberrima 

0. standleyi Hitchcock 

0. surinamensis Hochstetter ex Steudel = 0. longifolia 

0. tamanquareana Soderstrom & Zuloaga 

0. taquara Swallen 

0. urvillei Steudel = 0. micrantha 

0. ventricosa Nees von Esenbeck = 0. micrantha 

0. wurdackii Swallen 

latifolia 

76 
Appendix 2 

Names Excluded from Olyra Linnaeus 

(Current names are in italics.) 

Olyra axillaris Lamarck = Lithachne pauciflora (Swartz) 

Olyra blanchetii Mez = Parodiolyra ramosissima (Trinius) 

Olyra brasiliensis (Bertoloni) Sprengel = Raddia brasiliensis 

Olyra capillata Trinius = Cryptochloa capillata (Trinius) 

Olyra capillata var. segregata Doell in Martius = Cryptochloa 

Olyra concinna Hooker f. = Cryptochloa concinna (Hooker f.) 

Olyra flaccida Doell in Martius = Arberellaflaccida (Doell in 

Olyra floribunda Raddi = Raddia brasiliensis Bertoloni 

Olyra floribunda var. microphylla Doell in Martius = Raddia 

Olyra hoehnei Pilger = Raddia brasiliensis Bertoloni 

Olyra lancifolia Mez [This species appears to belong to a 

different genus from Olyra. The poor material available does 

not allow a decision on its generic disposition to be made at 

this point.] 

Olyra lateralis (Presl ex Nees von Esenbeck) Chase = 

Parodiolyra lateralis (Presl ex Nees von Esenbeck) Soderstrom & Zuloaga, new combination 

Olyra luetzelburgii Pilger = Parodiolyra luetzelburgii (Pilger) 

Soderstrom & Zuloaga, new combination 

Olyra malaccensis Wallich = Scleria caricina (Cyperaceae), 

teste Chase and Niles (1962) 

Olyra malmeana Ekman = Raddiella malmeana (Ekman) 

Swallen 

Palisot de Beauvois 

Soderstrom & Zuloaga, new combination 

Bertoloni 

Soderstrom 

capillata (Trinius) Soderstrom 

Swallen 

Martius) Soderstrom & Calderon 

brasiliensis Bertoloni 

Olyra nana Doell in Martius = Raddiella esenbeckii (Steudel) 

Olyra orientalis Loureiro = not a member of the Poaceae, teste 

Olyra ovata Hamilton = Zchnanthus panicoides Palisot de 

Olyra pauciflora Swartz = Lithachne paucifora (Swartz) 

Olyra pineti Wright ex Grisebach = Lithachne pineti (Wright 

Olyra podachne Mez = Cryptochloa capillata (Trinius) 

Olyra polypodioides Trinius = Raddia distichophylla (Schrader) 

Olyra ramosissima Trinius = Parodiolyra ramosissima (TrinOlyra sampaiana Hitchcock = Sucrea sampaiana (Hitchcock) 

Olyra sarmentosa Doell in Martius = Parodiolyra lateralis 

Olyra strephioides Grisebach = Mniochloa strephioides (GriseOlyra strictiflora Fournier = Cryptochloa strictiflora (Fournier) 

Olyra sympodica Doell in Martius = Piresia sympodica (Doell 

Olyra urbaniana Mez = Raddia guianensis (Brongniart) 

Calderon & Soderstrom 

Chase and Niles (1962) 

Beauvois 

Palisot de Beauvois 

ex Grisebach) Chase 

Soderstrom 

Chase 

ius) Soderstrom & Zuloaga 

Soderstrom 

(Presl ex Nees von Esenbeck) Soderstrom & Zuloaga 

bach) Chase 

Swallen 

in Martius) Swallen 

Hitchcock 

77 
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