SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 9 

Genera of Bamboos 

Native to the New World 

(Gramineae: Bambusoideae) 

F. A. McClure 

(EDITED BY THOMAS R. SODERSTROM) 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1978 
ABSTRACT 

McClure, F. A. (edited by Thomas R. Soderstrom). Genera of Bamboos Native 

to the New World (Gramineae: Bambusoideae). Smithsonian Contributions to 

Botany, number 9, 148 pages, 49 figures, 1973.-Seventeen genera of bamboos 

native to the New World are classified, compared, and illustrated, including 

four new genera and four new species. The introduction of several taxonomic 

characters based on hitherto neglected morphological features, both vegetative 

and reproductive, is undertaken with the objective of improving traditional 

perspectives. Two reforms are urged as essential to the elevation of the level 

of refinement at which future taxonomic treatments of bamboos may be executed. These are (1) the general adoption of improved collecting methods 

correlated with more extensive and sustained field observations, and (2) the 

fostering, through interdisciplinary collaboration, of progressive development and 

integration of diversified studies of documented materials drawn from a common 

source for each individual taxon. 

OFFICIAL PURLICATION DATE is handstamped in a limited number of initial copies and is re. 

corded in the Institution’s annual report, SmBthsonian Year. SI PRESS NUMBER 4780. SERIE~ COVER 

DESIGN: Leaf clearing from the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

McClure, Floyd Alonzo, 1897-1970. 

Genera of Bamboos native to the new world (Gramineae: Bambusoideae) 

(Smithsonian contributions to botany no. 9.) 

Bibliography: p. 

1. Bamboo-America. 1. Title. 11. Series: Smithsonian Institution. Smithsonian contributions 

QKI ~2747 no. 9 [QK495.G74] 581’.08s [584’.93] 72-8954 

to botany no. 9. 

For sale by the Superintendent of Documents, US. Government Printing Office, Washington, D.C. 20402 

Price $2.60 domestic postpaid, or $2.25 GPO Bookstore 
h 

On 15 April 1970, Dr. Floyd A. McClure passed away in his bamboo garden 

while removing a plant to give to a young friend. McClure had devoted his entire 

career to the bamboos, which he described as a “symbol of uprightness, chivalry, 

and devotion.” His first book, The Bamboos: A Fresh Perspective, and the present 

work represent as rich a heritage as any man could hope to leave to the botanical 

world. Of all his numerous publications, these two books represent the synthesis of 

a lifetime study by one of the most productive minds in the botanical sciences, a 

mind which, to our good fortune, had matured to its full potential. 

McClure, not only as a botanist but as a man, inspired and encouraged all 

those with whom he came in contact. My first acquaintance with him was here 

at the Smithsonian Institution in August 1961, since which time I had cherished 

his friendship and advice. In 1967, he persuaded me to extend my interest in the 

anatomy of the monocotyledons to include the vascular architecture of the bamboo culm. Utilizing the optical shuttle system and the data analyzer projector, 

this study confirmed the growing consensus that the structure of the bamboo 

culm, like that of maize, does not represent a typical monocotyledon stem as had 

been previously accepted. 

A little over a year before McClure’s death, Dr. Thomas R. Soderstrom, 

Smithsonian agrostologist, joined him in his bamboo studies; thus we are assured 

that McClure’s life work will be carried on. Daily sessions between the two 

continued to the very day of his death. Soderstrom, following McClure’s wishes, 

has undertaken the difficult and demanding task of seeing to fruition this 

present and final work of a great man and botanist. 

McClure’s original intention had been to revise the bamboos for Die 

Natiirlichen Pflanzenjamilien; but when he realized that this would be too large 

a task, he decided to publish first the genera of the New World as represented 

in this book. Although the bulk of the manuscript was in near-readiness at the 

time of his death, significant portions of the work remained undone. It thus 

fell to Soderstrom to conduct an intensive program of organizing the material. 

Every effort was made to complete the book just as McClure would have, 

drawing upon his copious notes whenever possible to finish the uncompleted 

portions. Fortunately, Mrs. McClure, who had worked closely with her husband 

throughout his career and was completely knowledgable of the details of 

McClure’s manuscript-preparation methods, was able to provide much needed 

help. Mr. Elmer Smith, a botanical illustrator at Harvard University, returned 

to the Smithsonian on two occasions for extended periods to complete all the 

figures. He had worked closely with McClure over the years, and the two had 

developed a personal rapport that transcended the usual scientist-artist relationship. 

I want to congratulate Dr. Soderstrom for accepting the commitment to 

complete this monumental work, which, in its final form, I am sure would have 

pleased Dr. McClure. I also wish to stress that he has had to put aside his own 

research for extended periods to carry out this task. 

iii 
I would also like to thank the National Science Foundation for its continued 

support of this work after Dr. McClure’s death and Dr. Eric Holttum, who 

painstakingly reviewed the manuscript before submission to the Smithsonian 

Institution Press. Mr. H. K. Airy Shaw kindly reviewed the Latin descriptions, 

making a number of useful suggestions, and Dr. Alicia Lourteig compared parts 

of the text that appear in French with the original herbarium notes in Paris. Dr. 

Cleofk E. Calder6n gave assistance and the benefit of her expertise in the final 

stages of manuscript preparation. 

McClure was always concerned with the precise usage of words, and he 

developed over the years a glossary of bamboo terms which first appeared in his 

earlier book. Since publication of that glossary he modified some of the terms 

and wrote definitions for several new ones. Because of the practicality of having 

all terms appear in one reference, the decision was made to include in this 

volume the entire glossary, even though some of these terms are not used in 

connection with bamboos of the New World. 

The archival materials and bamboo library of Dr. McClure are now 

incorporated into the Hitchcock-Chase Grass library in the Department of 

Botany, Smithsonian Institution. These materials will continue to be curated, 

updated, and utilized. Without the extensive holdings of bamboo specimens, 

literature, card files, and cross-references that Dr. McClure had accumulated over 

a lifetime, a book of this magnitude could never have been completed. 

EDWARD S. AYENSU 

Chairman 

Department of Botany 

Smithsonian Znstitution 

Washington, D.C. 

21 Sepember 1971 

iv 
Foreword 

My first acquaintance with Dr. McClure came through his articles on plants 

cultivated at Canton, which I read in Singapore, where I was also making 

myself acquainted with plants grown by Chinese people. We first met in Java 

in 1929, about which time I began to try to take some interest in bamboos, 

provoked by the fact that nearly all the bamboo plants in the Botanic Gardens 

at Singapore had lost their labels, and the consideration that bamboos were 

plants of importance which should not be neglected. This led me to examine 

native Malayan bamboos when I had the opportunity of botanical travel. In this 

rather casual study I was helped and stimulated by McClure’s successive papers 

on his Chinese bamboos, from which I learned much. Our paths met again 

after World War 11, in Singapore, London, and Washington, where I had the 

privilege of his hospitality on two occasions. I have enjoyed reading the text 

of this book, which brings many happy memories into my conscious thought, 

and am honored to be asked to write this Foreword, in which I wish to try to 

assess the significance of this, his last contribution to recorded knowledge of 

the plants to which he devoted so much of his life. 

Most early descriptions of bamboos were based on flowering specimens by 

herbarium botanists who had never seen the living plants. Inevitably such 

descriptions omitted vegetative characters without which field botanists and 

cultivators could not name their plants, many of which flower only at long 

intervals. Inevitably, also, the herbarium botanists, familiar with the plants of 

north temperate regions, compared bamboo spikelets and flowers with those of 

the grasses with which they were familiar, and tried to fit bamboos into the 

grass pattern, not realizing that the reverse process leads to a better understanding of the whole. 

Effective vegetative description, and its correlation with floral description, 

of the great bamboos of Southeast Asia was begun by Kurz, and was continued 

by Gamble in his comprehensive monograph of Indian bamboos (1896). Gamble, 

however, did not extend his thinking on the comparative morphology of the 

inflorescence beyond that of Munro and Bentham, and little further critical 

thought had been given to the subject when in 1924 McClure began his studies 

of the bamboos of southern China (related to those of India but in many 

cases distinct). Between 1924 and 1940 he established a plantation of six hundred 

bamboo plants, collected during numerous travels, so that he could watch their 

growth and flowering, In so doing he studied vegetative branching at culm 

nodes (which Japanese botanists had found important in describing their own 

bamboos) and also made detailed observations on the development of inflorescences. For the first time he correlated vegetative branching with that of the 

inflorescence, and established clearly a basic distinction between the two types 

of inflorescence, referred to in the present book as iterauctant (indeterminate) 

and semelauctant (determinate). Iterauctant branching matches exactly the 

vegetative branching at culm nodes and its precise description involves a more 

careful observation of the sheathing organs at the bases of spikelet-like structures 

V 
than had previously been undertaken. This is one of the keys to the understanding of bamboo classification. Japanese botanists also studied the branching 

of bamboo rhizomes, a subject of great practical importance to growers; McClure 

did the same for the bamboos of China, concerning which no such records had 

been made. 

Thus when McClure went to tropical America in 1942 he had acquired 

an understanding of bamboos as living plants wider and more detailed than 

that of any of his predecessors, and enriched by his own original thought. For 

the first time he applied to American bamboos the same kind of comprehensive 

observation and thought, and this has resulted in a synthesis, presented in this 

book, of unique significance. He examined many native bamboos over a wide 

range in the American tropics and subtropics, collecting specimens with all the 

kind of detailed observation he had found to be necessary; he also examined 

type-specimens and other material of all previously described species, attempting 

to correlate specimens which often failed to show all the characters he believed 

to be significant. He made a new bamboo garden, in which once more he could 

study growing plants. The work is not completed; the imperfections of earliercollected specimens leave gaps in our knowledge. Although clear distinctions 

between species in most genera remain to be established, the present work is a 

firm foundation on which further work can be based, and an indispensible guide 

to further thought and action. The excellent illustrations, designed to show 

clearly a wealth of significant detail, are a very important complement to the text. 

My own knowledge of bamboos as living plants is confined to the species 

of Malaya and New Guinea, and some from India seen in cultivation. The 

tropical American species dealt with in the present book are in the main very 

different from those of the Malayan region, and prompt some thoughts, which 

I hope may be of interest, though I have not an adequate knowledge of Japanese bamboos. These are also important to the following statement concerning 

the evolutionary significance of bamboos as members of the family Gramineae. 

It seems to me possible that the development of woody culms, which is a 

distinctive feature of bamboos, may have originated more than once in the 

family; that is, the bamboos as we know them may not be monophyletic in this 

respect. It seems likely that the Gramineae had developed a reduced and 

specialized type of inflorescence before woody forms appeared (the very specialized woody nature of bamboos is unique and cannot be a relic of any 

proto-monocotyledonous stock). The large bamboos of the genera Bambusa, 

Dendrocalamus, and Gigantochloa (which I believe are closely allied, though 

separated by Munro on fruit-characters for which he had little evidence, but 

which were repeatedly copied by later authors), as well as Schizostachyum and 

its allies, all have six stamens and a well-developed pericarp. They also all 

have a caespitose habit with pachymorph rhizome-elements, and inflorescences 

of the iterauctant type that matches exactly vegetative branching, with a 

prophyllum at the base of every branch, Thus these bamboos, predominant in 

Southeast Asia, in addition to the maximum development of woody habit, also 

show a combination of primitive characters. Of this great group of “primitive” 

bamboos, only Bambusa migrated to the Americas from Asia, presumably at a 

time when the land-bridge to North America from Asia experienced a mild 

climate. 

The other well-known genus shared by America and Asia is Arundinaria. 

One may also postulate for it a similar migration (even as restricted by McClure) 

vi 
since it is much more diversified in Asia than America. This genus has been 

variously interpreted, both in America and Asia, by different authors. Japanese 

botanists have recognized that some of their bamboos, originally included in 

Arundinaria, differed in many ways, and have attempted to recognize new genera. 

But the type-species of Arundinaria is North American, and so the species of 

Asia must be judged by that type. In the present book McClure deals exhaustively with A,. gigantea, and indicates what he regards as the essential characters 

of the genus, from which he excludes many American species formerly included 

in it. Unfortunately he does not indicate which species of the Old World he 

regards as congeneric with the American type. I believe that students of Japanese 

bamboos would find the treatment of Arundinaria in the present work valuable 

in a reassessment of their own species of the segregate genera. 

Arundinaria has a semelauctant inflorescence in which the spikelets are 

grass-like and the branches of the inflorescence lack prophylla at their bases. 

In the Bambusa group of genera there is a prophyllum as first foliar organ on 

every branch, leafy or flowering, right up to the prophylla called paleae which 

enclose the true floral elements. In most grasses there are no prophylla (and no 

bracts) at any branch of the inflorescence, but the paleae persist. There is, thus, 

a gap between the prophylla of vegetative branches and the paleae, so that the 

homology of vegetative prophylla and paleae was long unrecognized. The 

semelauctant bamboos sometimes show intermediate stages, with prophylla 

present at the base of inflorescence branches (Figures 3~, 1 9 ~ ) . T h e presence 

of such intermediate stages may be significant as indicating possible evolutionary 

lines of transition from the Bambusa to the grass types of inflorescence. It seems 

to me likely that the transition has occurred on several lines. The bamboos of 

Japan might provide interesting evidence, if considered from this standpoint. 

In any case, precise observation of bracts and prophylla in relation to flowering 

branches of semelauctant bamboos may indicate significant diagnostic characters 

for recognition of species. 

Arundinaria has leptomorph rhizomes from which new culms arise as 

axillary structures; that is, the growth of the rhizomes is monopodial. I have 

produced evidence (Holttum, 1955) that sympodial branching of the stem predominates in all families of monocotyledons, and monopodial branching is 

certainly the exception in Gramineae. The latter occurs in some genera of 

Paniceae, but I believe it is rare in other divisions of the family, In some other 

families (e. g., Palms) it has developed separately in distinct groups of genera, 

and it may have done the same among woody Gramineae. Possibly therefore the 

occurrence of leptomorph rhizomes in Awndinaria is less important as evidence 

of relationship to other genera than some other characters. 

In this book McClure describes a new species of bamboo which he includes 

in the genus Yushania, previously known only from Taiwan. This looks like a 

third migrant (or could both have evolved from a common northern ancestor 

no longer extant?). Other genera of American bamboos, now more clearly distinguished than formerly, may be found to give indications of relationships 

with Asia. 

One group of species, however, appears to be quite exclusive to the New 

World, and surely originated there, namely those here included in Chusquea and 

its immediate allies Neurolepis and Swallenochloa. Munro (1868: 13, 52) remarked on the resemblance between Chusquea and some panicoid grasses, and 

also to the fact that in essentials of spikelet structure they are not far apart. 

vii 
It is greatly to be regretted that McClure was not able to complete a survey 

of the bamboo genera of the Old World, of which he had such a wide and 

intimate knowledge. He accumulated, however, an immense amount of the 

reference material necessary for such a survey, which, together with his manuscript notes, is available as a basis for further study by his students. I hope such 

study will result in published works, so that more of the fruits of McClure's 

labors may become available to botanists, foresters, and others in Southeast 

Asia, and may be put to practical as well as scientific use. I believe that the 

peculiar and very remarkable properties of the bamboos of tropical Asia could 

find new uses in modern technology, but they have not yet received the attention they deserve. McClure's bamboo plantation at Canton could yet be of 

great service in providing more information about these plants. 

A new study of bamboo anatomy, especially nodal anatomy, is certainly 

a key to understanding the branch patterns which McClure and others have 

shown to be distinctive characters. Plant anatomists have been too content 

to describe only internodal structure (which is, of course, important). Nodal 

anatomy is extremely complex, and doubtless difficult to understand, but on it 

depends the dynamics of growth of bamboos and of some other monocotyledons. 

Because there is not secondary growth of concentric woody tissue in bamboos, 

the primary structure of the node has to be far more precise than in dicotyledons, 

and intercalary growth at the base of each culm internode needs a far more 

complex organization. When this is better understood, respect for bamboos as 

highly specialized organisms may well be greatly enhanced. The question has 

also to be asked: How is the control of all this complexity organized? 

R. E. HOLTTUM 

Royal Botanic Gardens 

Kew, England 

November 1971 

viii 
Acknowledgments 

The laboratory studies and field work that serve as the basis for the following 

treatise cover a period dating back to 1943. In that year I initiated, under the 

auspices of the Smithsonian Institution, a project for the U. S. National Research 

Council: a survey of bamboo species of a special category occurring in Mexico, 

Central American countries, Colombia, Venezuela, and Brazil (McClure: 1944). 

Opportunities to continue the collection and field study of bamboos of the 

Western Hemisphere were afforded during my incumbency (1944-1954) as 

Field Service Consultant on Bamboo with the Office of Foreign Agricultural 

Relations of the U. S. Department of Agriculture. More recently, as Consultant 

on Tropical Forestry to the Maria Moors Cabot Foundation of Harvard University (1956-1959) I was given the opportunity to visit European herbaria to 

study types and critical specimens of Western Hemisphere bamboo genera, 

and to advance the organization of pertinent data from the literature and from 

personal experience. Since 1955, the Department of Botany of the Smithsonian 

Institution, as administrator of grants-in-aid from the U. S. National Science 

Foundation, has fostered the progress of my bamboo studies directed toward 

the revision of the descriptions of the bamboo genera of the world, for the 

second edition of Engler and Prantl’s Die Naturlichen Pflanrenfamilien. 

Facilities for the study of the Gramineae (including the Bambusoideae) now 

available at the United States National Herbarium were greatly enriched by the 

fruits of the many mutually related activities pursued by Dr. Agnes Chase during 

the course of her long and distinguished career as agrostologist. Among these 

fruits are innumerable notes and photographs illuminating types and critical 

specimens of gramineous plants that she investigated in the principal herbaria of 

Europe and America. In many cases these notes and photographs are accompanied by duplicate specimens or fragements from type-collections-specimens 

presented as tokens of appreciation for expert curatorial services (identifications, 

annotations, etc.) freely, but often at great personal sacrifice, bestowed upon 

taxonomically critical materials in extant (but in many cases neglected) classical 

collections. The acquisition of specimens representative of the grass (and 

bamboo) flora of the world was greatly augmented by a system of exchanges 

supported by long sets of duplicates by Dr. Chase on numerous wide-ranging 

field trips carried out in the United States and other New World countries. 

Fruitful discussions with Thomas R. Soderstrom, Associate Curator, Division 

of Grasses, and David B. Lellinger, Associate Curator, Division of Ferns, Depaftment of Botany, Smithsonian Institution, are gratefully acknowledged. 

The skills of several artists went into the preparation of the line drawingssome based on my own original sketches, others made specifically for the present 

work: Elmer W. Smith, Mrs. Gesina Berendina Threlkeld, Mrs. Martha Niepold, 

and Florence Mekeel (Mrs. H. J. Lambeth). 

Essential to the fulfillment of the objectives of this study has been the 

opportunity to examine the types and critical specimens of the genera under 

consideration. In this connection, it is a pleasure to acknowledge the courtesies 
extended by the responsible officials of the following botanical institutions: 

The Herbarium of the Field Museum of Natural History, Chicago; the Herbarium of the New York Botanical Garden; the Herbarium of the Arnold Arboretum, Cambridge; The Herbarium of the Museum National d’Histoire 

Naturelle, Paris; The Herbarium of the Royal Botanic Gardens, Kew; The 

Herbarium of the British Museum (Natural History), London; the Herbarium 

of the Botanische Museum, Berlin-Dahlem; The Herbarium of the University 

of Pisa; The Makino Herbarium, and the Herbarium of the Botanical Institute, 

Tokyo; and the United States National Herbarium (Smithsonian Institution), 

Washington, D. C. 

F. A. MCCLURE 

Smithsonian Institution 

Washington, D. C. 

I March 1970 

I I 

To Conrad Chapman, who from the beginning has manifested a sustained 

and sustaining interest in the objectives and the fruition of my bamboos studies. 

X 
Contents 

Page 

... Preface, by Edward S. Ayensu ..................... 

Foreword, by R. E. Holttum ....................... 

Acknowledgments, by F. A. McClure .................. 

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

Key to the Recognized Genera of Bamboos Native to the New World .... 

Apoclada McClure ................................................. 8 

Arthrostylidium Ruprecht .......................................... 15 

Athroostachys Bentham ............................. ........... 40 

6 

Arundinaria Michaux ............................................. 21 

Atractantha McClure, new genus ............. " . . " " " " 42 

Aulonemia Goudot ................................................ 53 

Bambusa Schreber, Subgenus Guadua (Kunth) Hackel .... 

Colanthelia McClure and E. W. Smith, new genus .................... 77 

Glaziophyton Franchet ............................................. 83 

Chusquea Kunth ............................ 

Elytrostachys McClure ....................................... .... 79 

Merostachys Sprengel .............................................. 87 

Myriocladus Swallen ................................. ....... 94 

Neurolepis Meisner ............ ................................ 97 

Rhipidocladum McClure, new ge ....................... 

Swallenochloa McClure, new genus ..... ................... 

Yushania K. H. Keng .............................................. 113 

Glossary .......................................................... 122 

Literature Cited 140 ............................................. 

xi 

Genera of Bamboos 

Native to the New World 

(Gramineae: Bambusoideae) 

F. A. McClure 

(EDITED BY THOMAS R. SODERSTROM) 

True knowledge can only be acquired piecemeal, 

by the patient interrogation of nature. 

Sir Edmund Whittaker 

Introduction 

It is now more than three quarters of a century 

since the preliminary publication of Hackel’s 

treatment of the bamboo genera of the world, prepared as part of the first edition of Engler and 

Prantl’s Die Naturlichen Pflanzenfumilien (1887). 

The fact that several supplements were published 

shortly after is eloquent of Hackel’s feeling that 

his work on this group was incomplete and of an 

essentially tentative nature. It is sobering to reflect 

that, although the number of bamboo species 

known to science has more than doubled in the 

meantime, the nature of the material collected has 

changed but little. The problem of achieving an 

adequate perspective on the taxonomy of the group, 

however, has become immensely more complicated, 

rather than simplified, by much that has been done 

in this interval. Specimens collected continue to be 

fragmentary, field notes are still brief or lacking, 

F. A. McClure, Honorary Research Associate, Department of 

Botany, Smithsonian Institution, Washington, D. C. 20560 

(Deceased 1970). Thomas R. Soderstrom, Department of 

Botany, Smithsonian Institution, Washington, D. C. 20560. 

and the published descriptions of entities have 

continued to omit reference to features of fundamental taxonomic importance, especially those relating to the nature of the branching habit of all 

axes of the plant. Published supplementary studies 

in the fields of anatomy and cytology have not, in 

general, improved the situation as they might have, 

had the workers in these fields collaborated more 

closely with the taxonomist. 

The present treatment is neither monographic 

nor definitive in its intent or scope. It is offered as 

representing a preliminary step forward in the 

taxonomic conquest of the bamboos of the New 

World. The dominant objective of the studies on 

which this treatise is based has been to facilitate 

the recognition of the generic affiliation of each 

bamboo native to the Western Hemisphere that 

requires identification. To this end-and within 

the limits imposed by available materials-an effort 

has been made to bring into sharper focus the 

image (dependable features) and the apparent 

present boundaries of each genus. It has been possible to improve the concept and the circumscription of some genera by including generally 

neglected morphological features. The elimination 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

of traditionally included features that do not hold 

good has improved the concept and the circumscription of some genera, and has suggested a 

revised taxonomic disposition of others. The diverse 

patterns oE branching habit in the vegetative apparatus and in the inflorescence have been given 

particular attention, both here and in a broad 

survey published earlier (McClure, 1966b). The 

exploitation of the taxonomic potential of these 

patterns is limited to those entities in the available 

specimens of which this feature is represented. 

The incompleteness and disparities (notably the 

incongruent representations of morphological and 

ontogenetic aspects of the plant) that characterize 

the available specimens, field notes, and descriptions of most recorded bamboo species exert serious 

restrictions upon the development of uniformly 

satisfactory circumscriptions of the genera. I have 

been able to improve and olarify the image of 

representative species of some of the genera through 

field studies. Available plants of the type-species 

and/or one or more representative species of fourteen genera of bamboos of the New World and 

the Old World, respectively, have been maintained 

under cultivation for a number of years for progressive collection and observation of their successive stages of development, on a comparative basis. 

Owing to the incomplete and fragmentary representation of some genera in the existing herbarium 

material and the fact that living material of some 

of the genera has not been accessible to me, the 

drafting of generic descriptions that are uniform 

in their coverage of every detail is still an impossibility, In comparison with previous treatments, 

however, the coverage of taxonomically significant 

features has been greatly extended as far as gross 

morphology is concerned. Attention has thus been 

directed to structures and to dynamic aspects of 

the bamboo plant that have hitherto been largely 

or completely neglected. It is hoped that those 

who collect bamboo specimens for identification or 

for description and naming and those who address 

themselves to the description and classification of 

the bamboos will in the future find useful leads 

here, and that the taxonomic conquest of the bamboos will have received a perceptible impetus. The 

new demands for evidence from other disciplines 

means that in order to be up-to-date the collector 

must abandon the idea that the conventional leafy 

flowering specimens are sufficient for purposes of 

description and classification. He must have in 

mind not only the morphologist but the anatomist, 

cytologist, chemist, geneticist, biochemist, ecologist, 

etc., so that material for the study of each entity in 

as many disciplines as possible may be available 

under the same herbarium voucher. 

In some cases, the nomenclatural type of a genus 

(especially where its choice was fortuitous) represents a species that is morphologically peripheral 

to the main body of the currently recognized com 

ponents of the genus. In any case, the depth of 

presently attainable taxonomic perspectives varies 

from genus to genus. For this reason, the emphasis on phylogeny is uneven, tentative, and of necessity extremely limited in this treatment of the 

genera. During the course of the phylogenetic die 

versification of bamboo taxa, some of the characteristic expressions of different individual features 

(conventionally referred to as "characters") have 

recombined (in disregard of quasi-generic boundaries). This may be due to the persistence of 

genetic compatibility within populations of related 

plants, or between groups of related species, some 

of which have already acquired combinations of 

morphological features of such strong divergence as 

to suggest, respectively, either specific or generic 

segregation by the taxonomist. 

The proposed consolidation of currently recognized genera that apparently are not set off from 

each other by clearly disjunct patterns of morphological feature combinations is prompted by 

the desire to facilitate identifications to genus. Uncertainties often arise as a result of feature combinations that constitute intergradations between 

two or more such genera, especially where these 

intergradations have either been ignored or given 

separate generic rank. An effort has been made to 

eliminate these uncertainties by allowing the intergradations to fall within a single genus with an 

expanded circumscription. They then yoke toge ther the recognizable extremes under one generic 

name. Where feasible, the diversities are then given 

informal status as either subgenera or sections. 

However, it is possible that more comprehensive 

morphological studies, supplemented by correlated 

studies in other disciplines and deepened by the 

use of the electron scanning microscope of both 

vegetative and reproductive structures, may force 

the revision of the generic lines proposed here. 
NUMBER 9 

It appears that some groups of closely related 

bamboos flout the intellectual concepts of morphological homogeneity and morphological disjunctions, at either the “species level” or the “genus 

level”-or both, Of course, the terms “genus level” 

and “species level” lack a sound semantic basis, 

since the component elements of both genera and 

species are, in fact, determined primarily on the 

basis of circumscription (fences) that reflect morphological similarities and disjunctions rather than 

levels of evolutionary advancement. As every perceptive taxonomist recognizes, the concepts “relatively advanced” and “relatively primitive” are 

properly focused upon the individual features or 

structures of an organism, rather than upon the 

organism as a whole or a group of organisms. Carlquist (1961:7) states the point very simply in these 

words: “One must remember that only characteristics, not plants, or species, are primitive or advanced.” This is because features commonly evolve 

independently-as though possessing their own individual phylogeny-rather than simultaneously 

with other characters, as though linked with them, 

genotypically. Proposed phylogenetic arrangements 

of taxa, or groups of taxa, can be given meaning 

only by a projection of the relative incidence of 

supposedly “advanced” and supposedly “primitive” 

features in their component elements. 

The temporal and morphological divergencies 

that, in most bamboos, isolate the vegetative state 

from the sexually reproductive state, combine to 

give rise to a physical isolation between the specimens, descriptions, and names that, in many cases, 

independently represent these two states of a given 

species. As a result of the practical difficulties involved in bringing adequate representation of the 

essential morphological features of the vegetative 

state of a bamboo into authentic association with 

the morphological features of its reproductive state, 

a dichotomy similar to that which has produced 

the mycological category “Fungi Imperfecti” has 

developed in the taxonomy of the bamboos. In 

comparison with the fungi, however, most bamboos 

confront both the collector and the taxonomist with 

formidable obstacles to the resolution of these 

difficulties. These obstacles are imposed by a very 

protracted vegetative state, and morphological gradations that appear in acropetal sequence within 

each component axis of the plant. These and other 

gradations appear also in temporal sequence among 

the component parts of the relatively massive body 

of the plant as it develops toward sexual maturity. 

Another stumbling block has its origin in the 

radical changes in appearance and composition 

that often take place in the inflorescences during 

their development, especially in those genera where 

they are of the iterauctant (indeterminate) type 

(McClure, 1966b:93 et seq.). The generic affinities 

suggested by the morphological features of the vegetative apparatus of a given species may be, and 

often are, quite different from those suggested by 

the morphological features of the reproductive apparatus of the same species. Therefore, the full 

integration of the vegetative features with the reproductive features of each species is the only rational approach to the achievement of a realistic 

circumscription of the genera. This calls for drastic 

improvements in collecting methods and in taxonomic studies as well. The morphology and the 

ontogeny of the whole plant must eventually be 

assembled and brought under consideration. Until 

this has been accomplished the taxonomist cannot 

function effectively (or respectably) in collaboration with specialists in integrating the results of 

studies on paramorphological aspects of the bamboos for the progressive clarification of their taxonomy. A realistic approach to the solution of the 

“genus problem” requires the building up, maintenance, and sustained competent comprehensive 

interdisciplinary study of living collections embracing both the vegetative and the reproductive states 

of critical species of each genus. 

In the first sentence of Chapter I of his illuminating review entitled Comparative Plant Anatomy 

(1961), Carlquist reminds us that “comparative 

anatomical studies of angiosperms have achieved 

a remarkable record within the past century, and 

one may safely say that few systematic studies would 

fail to benefit from incorporation of anatomical 

data.” And again, on page 120, “certainly no generic monograph can be said to be complete without studies on leaf anatomy.” This “guide to taxonomic and evolutionary application of anatomical 

data in angiosperms” serves the useful purpose of 

emphasizing the impressive number and diversity 

of the anatomical features that have been brought 

into focus by studies of angiosperms other than 

the bamboos. The sparsity of references to bam- 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

boos incorporated by Carlquist in his book emphasizes the retarded state of our current knowledge 

of the anatomy of this group of plants. It is regrettable, however, that so few of the published 

anatomical studies of the bamboos are sufficiently 

comprehensive or well documented to afford a reliable basis for conclusions as to their taxonomic 

significance for the differentiation of genera. It remains to be seen whether really comprehensive 

studies on their anatomy will be significant for the 

taxonomy of the bamboos as they have been shown 

to be in some other groups of the Gramineae. 

In circumscription of genera, the taxonomic usefulness of features still to be revealed by paramorphological studies of the bamboos remains to be 

effectively explored. For the time being, at least, 

the illumination that is potential in such distinguished exploratory studies as those carried out 

by Shibata (1900), Brandis (1907), Ohki (1932), 

Takenouchi (1931a,b), Freier (1941), Metcalfe 

(1960), and others (cited under the pertinent genera) is eclipsed by the inadequate array of species 

and features covered, and by the shadow of persistent uncertainties with respect to where the truly 

generic boundaries lie and, in some cases, with respect to the identification or the documentation of 

those species that were studied. The extent to 

which the present work may appreciably improve 

the view is limited by the lack of evidence as to 

whether, and at what points, paramorphological 

features will strengthen or modify the image of any 

genus as it is here portrayed. 

Valuable perspective on the incidence and the 

varations of a number of anatomical features of 47 

species listed under 25 genera of bamboos are presented by Metcalfe (1960). Insofar as the bamboos 

are concerned, major attention is given in this 

work to leaf anatomy. The generic perspectives 

afforded by these recorded data are summarized 

by Metcalfe (1960584) as follows: “The anatomical 

data obtained for the leaves of the various bamboos examined by the present writer have been 

brought together in Table I. Perusal of this table 

confirms the opinion already expressed that there 

are no clear-cut distinctions between any of the 

genera that have been examined so far as leaf structure is concerned.” 

Ohki (1934) published a summary of his studies 

of spodograms of the leaf epidermis in representative species of Bambusa, Dendrocalamus, Phyllo. 

stachys, Pleioblastus, Sasa, Semiarundinaria, and 

Sinobambusa. Of Ohki’s results, Metcalfe (1960: 

584) has the following to say: 

The facts recorded by Ohki seem to the present writer to 

support the view that there are no very clear-cut divisions 

between the genera on the basis of characters revealed in 

spodograms. On the other hand, variations in the occurrence 

and distribution of macro-hairs, micro-hairs, and pridtlehairs, variations in the width of the bands of bulliform cells, 

differences in the number and distribution of papillae on the 

long-cells have specific diagnostic value. This applies also to 

quantitave characters such as the dimensions of stomata and 

prickle-hairs. For the indentification of some species it is important to know whether the walls of the basal cells of the 

micro-hairs are smooth or scabrous, and whether or not the 

walls of the prickle-hairs bear protuberances. 

The hazards of attributing to all members of a 

taxon (genus or species) a feature whose presence 

has been demonstrated in a single specimen or a 

single member has not always been heeded by the 

taxonomist. Metcalfe (1960:584) observes: “Thus 

we find uniseriate micro-hairs only in Guaduellu 

oblonga amongst the species examined by the author. Then again, Phyllostachys is the only genus 

from certain species of which fusoid-cells appear 

to be absent.” 

Some of the taxonomic characters most promising for the elucidation of the bamboos at the genus 

level are to be found in features of structures that 

are rarely represented at all in extant herbarium 

specimens-and never adequately so. This lack 

could be remedied either by extensive travel and 

repeated visits to wild stands of representative taxa, 

or by the establishment and maintenance of vast 

living collections for the same purpose. In either 

case, the lifetime of one person will not be enough. 

The traditional and still prevalent disproportionate emphasis on the importance of the diversity 

comprehended by features of the reproductive 

structures as the all-but-exclusive source of fundamental taxonomic characters has resulted in the 

acceptance of mere fragments of the plant as specimens adequately documenting most known taxa. 

It has also given us a vast but relatively sterile 

literary heritage. These unchallenged bad examples, aided by the priority principle embodied in 

the ICBN have tended to perpetuate-even encourage-undue haste in both the collection of specimens for preservation, and the preparation of their 

formal description for publication. The body of 
NUMBER 9 5 

uncorrelated literature has become undigestible. 

The elevation of the level of refinement at which 

future taxonomic treatments of bamboos may be 

executed is conditioned upon the improvement of 

certain procedures. In order to make this progress 

possible, collectors must adopt the objective of 

documenting for the taxonomist a more detailed 

image of each taxon collected. This would involve 

more deliberation in the selection of the components of the conventional array of pressed specimens. It would provide for more extensive and 

more sustained field observations, and the recording of the results in the form of notes and sketches 

and/or photographs to accompany the pressed 

specimens. It would involve the simultaneous collection of other appropriately selected, preserved, 

and correlated study materials by way of fostering 

the progressive development and integration of diversified studies, under interdisciplinary collaboration, of documented materials from a common 

source for each individual taxon (McClure, 1966b: 

6, 147). The study set of any given collector number should be made to represent the plant as 

completely as possible before duplicate specimens 

are segregated for distribution. 

Plant taxonomy is currently in a period of accelerated evolution. Perspectives on its problems 

and its methods are changing. Under the stimuli 

exerted by a developing dissatisfaction with the 

results achievable through traditional methods, and 

by the developing interest of specialists and their 

published studies in several fields ancillary to morphology, perceptive taxonomists are actively seeking new ways to bridge the gaps between the way 

things are in nature and the ways in which they are 

pictured in conventional treatises. It is anticipated 

that, through the gradual accumulation and integration of the results of interdisciplinary collaboration in morphological and paramorphological 

studies, the present work will be superseded by a 

greatly improved and more comprehensive portrayal of the bamboos-one that more adequately 

illuminates the details of their individualities in 

multidisciplinary depth and clarifies their generic 

and phylogenetic relationships, both within and 

beyond the subfamily Bambusoideae. 

ABOUT THE KEY.-A key is by many botanists admired on the basis of the conciseness of its leads. 

Partly for this reason, the array of contrasting features used in botanical keys is, by convenience, 

kept to a minimum. However, the user of such 

keys may be misled where exceptions to the impressions created by unduly abbreviated leads are 

left unmentioned. The negligent perpetuation of 

loose interpretations of structural complexities can 

thwart the desire to make an effective key. Inadvertently incorporated ambiguities may betray both 

the maker of a key and the user of it. 

The relatively lavish use of contrasting characters that appears at some points in this key is motivated by practical considerations. Effective definition of the assumed disjunctions between some 

pairs of closely related bamboo genera demands it. 

Again, specimens typical of extant herbarium material of most bamboos afford such a sparsely diversified and, in many ways, incongruent representation of the taxonomically critical structures that it 

is only by means of either a very broad scoop or a 

net with small meshes that many of them may, 

with any degree of confidence, be directed into a 

labeled pigeonhole. Because of limitations of available documented coverage it has not been possible 

to make genus-wide assessment of the potential of 

some features as sources of contrasting characters; 

however, the taxonomically useful array of contrasting characters has been augmented, both by drawing upon several morphological features hitherto 

neglected and by clarifying others that tradition 

interprets loosely. Some characters elsewhere given 

unqualified generic importance have had to be 

eliminated at certain junctures. 

An innovation that seems to serve a useful purpose here incorporates in either lead of a given 

pair, as occasion arises, mention of one or more 

characters which, while they do not provide additional contrasts, are common to the taxa designated 

by that particular lead for subsequent differentiation from each other. 

Conventional botanical keys commonly use indention as the means of matching members of each 

pair of contrasting leads, and for setting off adjacent pairs from each other, visually. Besides 

being uneconomical in many ways, this usage occasions avoidable inconveniences and disadvantages. It confronts both the preparer of the manuscript and the typesetter with hurdles that are not 

always successfully cleared. A key of the design 

offered here is easier to prepare, easier to set in 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

type, easier to proofread, and less costly to publish, 

than one embodying indentation. After a little 

practice in its use, one needs make no greater effort, either to follow the path that leads to the 

right name, or to retrace one’s steps in case one 

loses his way. 

Key to the Recognized Genera of Bamboos Native to the New World 

la. Culms in the vegetative state normally not developing branches .......................................... 2 

1b. 

2a. Flowering culms leafless, the above-ground internodes exceedingly thin walled and provided with pith septa at close intervals, the lowermost one greatly elongated: branch 

buds solitary at culm nodes: inflorescence never terminal to a culm, short-peduncled, 

the rachis deliquescent: spikelets terminating in a depauperate sterile anthecium ........ 

........................................................................................................................ Glaziophyton 

Flowering culms leafy, the above-ground internodes not exceedingly thin walled, and 

not provided with pith septa at close intervals, the lowermost one shorter than those 

that follow it: branch buds lacking at culm nodes; inflorescence always terminal to a 

Culms in the vegetative state normally developing branches ............................................ 3 

2b. 

3a. 

3b. 

4a. 

4b. 

5a. 

5b. 

6a. 

6b. 

7a. 

7b. 

8a. 

8b. 

9a. 

9b. 

1Oa. 

culm, long peduncled, the rachis excurrent: spikelets terminating in a perfect floret 

................................................................................................................................ ~euro~p~s 

Primary (first-order) branches (elements of the branch complement) at each midculm 

node more than one; stamens 3 .......................... , . , , . , , , , , , . , , . . , , . . , , , , , . . , , , , , , 

Primary (first-order) branches (elements of the bra complement) at e 

node single; stamens 3 ...................................................................... 

Transitional glumes regularly 4, the first two (“empty glumes”) in size and 

distinct from the second two (“sterile lemmas”) ......................................................... ..5 

Transitional glumes 0-1-2, empty glumes and sterile lemmas not distinguishable ....... 6 

Culm internodes lacking a lumen (filled with pith as in Zea mays): initial (primary) 

branch buds and primary (first-order) branches at each midculm node typically many 

(rarely only 3), of constellate insertion, the middle bud or branch several times as 

large as those that accompany it: culms arising from well-developed rhizomes of either 

pachymorph or leptomorph form in a given species, or from rhizomes of both forms 

in the same plant (as in Chusquea fendleri) ............................................. Chusquea 

(primary) branch buds and primary 

(first-order) branches at each midculm node usually 3, 1- (rarely) 5, of level insertion, 

the middle one dominant but not several times as large as those that flank it; culms 

arising from weakly developed pachymorph rhizomes and, in some species, also by 

tillering ....................................................................... Swallenochloa, new genus 

Inflorescences iterauctant: terminal segment of each rachis conspicuously elongate: spikelets terminating in a rudiment borne on a bristle-like prolongation of the rachilla 

Inflorescences semelauctant; terminal segment of each rachis not conspicuously elongate: 

Culms scandent‘; inflorescences capitate ................ ................................ Athroostachys 

Culms self supporting, the internodes typically more or less conspicuously sulcate, 

inflorescences not capitate .......................................................................................... 8 

Inflorescence a raceme: branches at midculm node to 5 ...................................... Apoclada 

Inflorescence a panicle: branches at midculm node to 3 ................................... Yushania 

Culm internodes provided with a lumen: initial 

behind the palea of the solitary perfect floret ...................... Atmctantha, new genus 

spikelets terminating in a depauperate sterile anthecium ........................................... 7 

Primary element of the branch complement at midculm nodes flat, unsegmented, appressed and wholly adnate to the surface of the culm: second-order branches of 

apsidate insertion and displayed in fan-shaped array; inflorescences semelauctant .. , ,I0 

Primary element of the branch complement at midculm nodes terete above its base, 

segmented and diverging from the culm; and dominant over the fasciculate secondMidculm sheaths (as far as known) with the blade very much narrower at its base than 

the apex of the sheath proper, abruptly rounded to a constricted base, and more or 

less strongly reflexed; the surface of the lower internodes in young culms commonly 

shows color mottling; inflorescences typically spicate racemes, the rachis in some species 

(as in Merostachys pluriflora) more or less strongly contracted, sometimes producing 

a congestion of the spikelets; spikelets typically of secund orientation, either (a) 

order branches: inflorescences either semelauctant or iterauctant ............................ 11 
NUMBER 9 7 

containing a single perfect flower (as in Merostachys speciosa) and terminating in a 

rudiment borne on a bristle-like prolongation of the rachilla, the latter more or less 

completely concealed in the narrow, canaliculate sulcus of the palea, or (b) containing two or more perfect flowers (as in Merostachys pturipora), the prolongation 

of the rachilla then usually more robust than bristle-like, the sulcus of the uppermost 

fertile palea broader and not canaliculate, and the terminal structure a depauperate 

(not rudimentary) sterile anthecium; transitional glumes at the base of the spikelet 2, 

strongly differing in size and shape, both empty, the first an “empty glume,” the second 

a “sterile (empty) lemma;” empty glume, sterile lemma and fertile anthecia all of 

brittle, extremely fragile consistency and of a typical grayish color at maturity; 

mature fruit oblong or ovoid, not compressed, the pericarp leathery or crustaceous, 

thickened toward the base and apex, the sulcus and the basal position of the embryo 

not manifest externally ........................................ 

Midculm sheaths (as far as known) with the blade 

apex of the sheath proper, broadly triangular and not at all reflexed; the surface of 

the internodes in young culms typically plain green. Inflorescences typically with 

either spicate or open racemose branching, the rachis either deliquescent or excurrent 

(when excurrent, either straight or more or less strongly geniculate; inflorescences 

with excurrent rachises in some species secund (strongly so in Rhipidocladum maxonii; 

weakly so in R. raceiniflorum and its allies) ; spikelets each containing few to several 

perfect flowers and terminating regularly in a depauperate sterile anthecium; transitional glumes at the base of the spikelet usually 3, rarely 2 or 4, all progressively 

approaching the first fertile lemma in size and shape, the first two empty, the third 

rarely and the fourth always (when present) subtending a depauperate flower; glumes 

and anthecia neither very brittle nor of extremely fragile consistency, at maturity 

usually representing variously tinted versions of stramineous color, in some species 

punctate with minute green dots; mature fruit oblong, usually more or less noticeably 

compressed, the pericarp pergamineous (leathery or crustaceous in R. verticillatum) , 

the sulcus and the basal position of the embryo usually more or less clearly manifest 

externally ............................................ .............. Rhipidocbdum, new genus 

Rhizomes leptomorph; developed midculm branch complements of restricted insertion, 

the primary element dominant-in some species (as in Arundinaria gigantea sp. tecta, 

for example) occasionally either suppressed temporarily or remaining solitary; inflorescences semelauctant; stigmas 3 ........................................................... A~ndina&~ 

Rhizomes pachymorph; stigmas in most cases 2 (often 3 in Bambusa) .......................... 12 

Inflorescences iterauctant, forming pseudospikelets ................................................... 13 

Inflorescences semelauctant, not forming pseudospikelets ................................................ 14 

Culm branches (at least at lower culm nodes) thorny (except in the unarmed northern 

forms of Bam’busa arnpZexifoNa and 8. aculeata) ; spikelets proper not pedicellate (the 

distal segment of the rachis not elongated) ................... Bambusa subgenus Guadua 

Culm branches all unarmed; spikelets proper rendered pediceuate by the elongated 

distal segment of the rachis ....................................................................... Eb’trostahYs 

Primary branch at midculm nodes appressed, not swollen basally, and not bearing buds 

at its congested proximal nodes; culms and branches in most known species characterized by one or more long internodes followed by several nodes closely crowded 

together by the suppression (abortion) of the intervening internodes; the Cuh 

sheaths and branch sheaths as well as the leaf sheaths each thickened and indurate; 

leaf blades sessile or subsessile, typically thick, of leathery texture, and broadly 

rounded at the base (rather grass-like and only perceptibly narrowed at the base in 

Myriocladus maguirei) ; inflorescences typically of elongate profile, with an excurrent 

rachis, of open racemose or paniculate branching (a condensed linear panicle in M. 

cardonae), the primary branches of the rachis in some species forming secund racemes; 

spikelets each containing usually 2 perfect flowers, occasionally 3 and, in at least one 

species, sometimes up to 5 ....................................................................... Myn’ocwus 

Developed branch complements at midculm nodes with the primary element patent, 

more or less prominently swollen basally, and potentially proliferating from buds at 

the congested proximal nodes; the culms, and sometimes their branches, in a few 

species (as in Arthrostylidium schomburgkii and Aulonemia queko) with a long 

internode (usually the first above-ground one of culms) followed by 2 or more nodes 

lob. 

1 la. 

llb. 

12a. 

12b. 

13a. 

13b. 

14a. 

14b. 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

closely crowded together by the suppression (abortion) of the intervening internodes; 

the culm sheaths and leaf sheaths in a few species (as in Aulonernia eflusa and A. depexa) thickened, indurate, and provided wih leathery or parchment-like blades; leaf 

blades with well-developed petioles in A. effusa and A. deflexa; inflorescences either 

spicate, subspicate, open racemose, purely paniculate, or with both racemose and 

paniculate branching in the same plant; spikelets each containing few to many 

perfect flowers ..................................................................................... 

15a. Base of the primary axis typically extending downward beyond the 

of the prophyllum or lowermost circumcingent sheath to form a low, downwardtapered promontory; inflorescence a raceme with zigzag rachis; plants of tropical or 

subtropical climates ................................................................................ Arthrostylidim 

l5b. Base of the primary axis not extending downward into a promontory; inflorescence a 

panicle (sometimes intermixed with racemose branches in CohntheZio) with straight 

rachises; plants of temperate or cold climates ................................................................ 16 

16a. Sheath at midculm nodes provided with scissing from) a conspicuous persistent 

................................................... ............................... Colanthelia, new genus 

Im nodes typically lackin spicuous basal girdle 

Apoclada McClure 

FIGUW 1-5 

Apoclada McClure, in Reitz, ed., 1967:57. 

Plants unicespitose; unarmed, Rhizomes pachymorph. Culm habit unknown, the internodes k 

sulcate. Branch complement at each midculm node 

arising from (what appears to be) more than one 

primary bud, the primary (first-order) component 

consisting of more than one axis (2-5) crowded 

together and inserted in a horizontal line, each 

primary axis dominant over branches of higher orders potentially arising by proliferation from buds 

(when these are present) at its proximal nodes. 

Leaf blades with transverse veinlets typically not at 

all or only weakly manifest externally. 

Inflorescences semelauctant, each typically consisting of but a single spikelet (weakly racemose in 

Apoclada arenicola; some spikelets reduced to a 

single floret in Apoclada diversa). Prophylla usually absent (present in Apoclada diversa when a 

flowering branch is reduced to a solitary spikelet). 

Transitional glumes of conventional nature typically lacking at the base of each spikelet-when 

present (e.g., Figure 3~) typically consisting of leaf 

sheaths bearing reduced blades, but highly diverse 

(as between species) in form, number, and spatial 

relation to the first fertile lemma. Spikelets usually 

made pedicellate by being terminal to a peduncle, 

sometimes (as occasionally in Apoclada diversa) 

sessile; containing few (rarely but one) to several 

to many perfect florets, and terminating in a depauperate sterile anthecium. Lemma (when subtending a functional flower) embracing its palea 

only basally at maturity. Palea broadly sulcate and 

2-keeled dorsally, the margins not at all or only 

slightly and partly overlapping. Lodicules typically 

3, the anterior 2 asymmetrical and paired, the posterior one smaller and symmetrical. Stamens normally 3 (exceptionally as sometimes observed in 

Apoclada diversa, varying from 3 to 6 in spikelets 

from the same specimen), the filaments filiform, 

free. Stigmas 2. Mature fruit (known only in 

Apoclada simplex) an oblong or subfusiform, sulcate, mucronate caryopsis, the pericarp glabrous, 

appreciably thickened and crustaceous, the sulcus 

broad, the basal position of the embryo weakly 

manifest externally, 

ETYMoLOGY.-The name Apoclada (Greek, apo, 

separate, and clados, branch) alludes to the apparently independent origin and insertion of the primary (first-order) elements of the midculm branch 

compiemen ts. 

TYpE-sPECrEs.-Apoclada simplex McClure and L. 

B. Smith. 

RELATIONSHIPS.-Aside from certain morphological features of the individual florets and a few 

features of the vegetative structures that are shared 

by other bamboo genera of the New World flora, 

Apoclada’s only apparent special relationship to 

another genus is one that vigorously stirs one’s 

curiosity. I refer to the strong detailed resemblance 

that links the fruits of Apoclada simplex (Figure 

5 P-R) to the fruits of the in-other-respects very 

different taxon, Rhipidocladum verticillatum (Figure 42 R-T). 
NUMBER 9 9 

DISTRIBUTION.-AS far as the available documentation informs us, all four of the currently recognized species of Apoclada are endemic to Brazil, 

where they are restricted to low elevations near 

the northern limits of the southern temperate zone. 

Two of these species are represented only by specimens collected from humid sites, and the other two 

appear to be restricted to arid habitats. 

Key to the Species of Apoclada 

la. Plants of humid sites; culm internodes antrorse-pubescent: primary axes of branch complement at midculm nodes more or less strongly unequal ................................................ 2 

lb. Plants of and sites: culm internodes glabrous; primary axes of branch complement at 

midculm nodes subequal, slender, appressed ................................................................... .3 

2a. Culm internodes with walls of substantial thickness; primary axes of branch complement 

at midculm nodes robust, divergent; flowering branches all leafy: inflorescences of 

invariable form ........................................... ................................ 4. A. simplex 

2b. Culm internodes thin walled: primary axes of bra complement at midculm nodes 

slender, appressed; flowering branches all leafless; inflorescences of highly variable 

form ........................................................................................................... .3. A. diversa 

Primary axes of midculm branch complements without basal buds: flowering branches 

leafy; keels of palea not perceptibly winged: style densely hispid ................... 

..................................... 2. A. canmvieira, new combination 

buds; flowering branches leafless; keels of palea perceptibly winged: style glabrous.. ......... 

3a. 

........................................ 

3b. Primary axes of midculm branch complements potentially proliferating from basal branch 

1. dpocladu arenicola McClure, new species 

FIGURES 1, 2 

Culmi in speciminibus suppetentibus usque 1.1 

m alti et 3 mm diametro, omnino glabri, internodiis cavis, medianis (quorum paries crassitudinis 

mediocris est) elongatis, versus cicatrices nodorum 

ramiferorum vix vel leviter dilatatis, superficiei 

omnino laevi et dura, primo opaca demum nitida. 

Culmorum et ramorum primariorum vaginae deciduae (in specimine suppetente deficientes, ita incognitae). Rami primarii ad nodos medianos 

culmorum prodientes plures, tenues, subaequales, 

appressi, omnino papillati, praecipue versus basin 

retrorse pubescentes, e nodis suis proximalibus proliferantes. Foliorum vagina arcta, persistens, primo 

praecipue apicem versus subtilissime et antrorse 

scabriuscula et farinosula et papillata, alibi glabra 

laevigata nitidaque, secus marginem externam ciliolata; auriculis minutis vel rudimentariis vel obsoletis; setis oralibus confertis, capillaceis, infime 

cohaerentibus, superne liberis, pallidis, levibus, anfractisque, denique cadentibus; ligula interiore 

perbreve apice convexa, dorso canescenti, margine 

dense ciliolata; ligula exteriore vix ulla, margine 

primo dense ciliolata demum glabrescenti et vel 

denticulata vel erosa vel integra; petiolo circa 1 mm 

~~ 

.............................. 1. A. ardcoka, new species 

longo, basi puberulo, superne glabro, primo glaucescenti demum nitido; lamina angustissima, vel 

planiuscula vel aculeiforrni rigidaque, usque ad 9 

cm longa et circa (in statu siccato) 1 mm lata, 

utrinsecus omnino glabra; nervis perpaucis; venulis 

transversis haud manifestis. 

Inflorescentiae ex culmo ab initio efoliato ortae; 

nonnullae (saltem in specimine suppetente) ad 

spiculam unicam redactae; vel solitariae vel 2-3 

simul insertae a bractea communi subtentae; proPhyllis nullis. Glumae transitionales vulgo 2 (interdum 1 vel 3) usque 3 cm inter se distantes, 

naviculares, apice vel acutae vel acuminatae et 

breviter aristatae, dorso carinatae plurinerves et 

glabrae, substantia subhyalinae fragilesque, I ca 10 

mm longa, 11 ca 11 mm longa, 111 ca 12 mm longa; 

gluma transitionalis suprema nonnunquam florem 

depauperatum subtendens ubi lemmati fertili similis. Spiculae pedicellatae, usque 4 cm longae, 

laxe pauciflorae, lanceatae, compressulae. Rachillae 

segmenta tarde disarticulantia, 2-3 mm longa, prominenter nervosa, clavata, unilateraliter complanata, apice dilatata, vel omnino vel pro parte 

puberula. Lemma pergamineum vel papyraceum, 

usque ad 13 mm longum, lanceatum, plurinerve, 

superne carinatum et apice breviter mucronatum. 

Palea lemmate brevior, papyracea, oblonga, dorso 

NUMBER 9 11 

FIGURE I.-Apoclada arenicola McClure. A, Leafy vegetative 

culm (midculm section), x 0.6; B, midculm bud complement, x 6; c, branch complement (basal part), x 6; D, 

leafy twig, x 1.8; E, apex of leaf sheath and base of leaf 

blade x ca. 18; F, flowering culm, with rhizome, x 0.6. 

Drawings A, c, F based on Chase 11886 (US) and B, D, E on 

Chase 12007 (US). 

latissime sulcata, in apicem variabilem vel hebete 

acuta vel obtusa vel subtruncata et bi- vel quadridentata, in plano mediano valde curvata, inter et 

extra carinas Clare nervosa glabraque, secus carinas 

prominenter alatas ciliis pallidis pulchre ornata. 

Lodiculae vel opacae vel pellucidae vel diaphanae, 

margine suprema ciliolatae alibi utrinsecus glabrae, 

anterioribus 2 sub-semiovatis, posteriore lanceata. 

Antherae usque 6 mm longae, brunneae, apice subpenicullatae. Ovarium glabrum. Stylus glaber, 

typice in stigmatibus 2 divisus, nonnumquam 

B 

(atypice) cum ram0 stigmatic0 tertio abortivo 

praeditus. Caryopsis incognita. 

Culms up to 1.1 m tall and 3 mm in diameter, 

glabrous throughout, the midculm internodes 

elongate, scarcely or only lightly flared toward the 

sheath scar at branch-bearing nodes, hollow, with 

a smooth, hard surface and walls of medium thickness. Sheaths of the culm and sheaths of primary 

branches deciduous (lacking in the available specimens). Primary branches produced at midculm 

nodes several, proliferating from buds at their basal 

nodes, slender, subequal, appressed, papillate 

throughout, retrorse-pubescent toward the base. 

Leaf sheaths tight, persistent, obscurely antrorsescabrous, perceptibly farinose and papillate near 

the apex, elsewhere glabrous, smooth, and lustrous, 

ciliolate on the outer margin; auricles minute, rudimentary or obsolete; oral setae numerous, crowded, 

TICURE 2.-Apoclada arenicola McClure, A, Spikelet, x 1.2; B, transitional glume I, X 7.2; c, 

ransitional glume 11, x 7.2; D, transitional glume 111, x 7.2; E, lemma, x 7.2; F, floret, 

x 7.2; G, palea, abaxial view of distal portion, x 7.2; H, palea, oblique view, x 7.2; I, lodicule 

:omplement, x 15; J, stamen, x 7.2; K, gynoecium, x 30. All drawings based on Chase 11886 

(US) ' 

. 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

hair-like, cohering basally, free, pale, smooth and 

wavy above, falling away at length; inner ligule 

very short, dorsally canescent, apically convex, 

densely ciliolate on the margin; outer ligule a thin 

line, its margin ciliolate at first, later glabrescent, 

then seen as either denticulate, erose, or entire; 

petiole ca 1 mm long, basally puberulous, glabrous 

distally, lightly glaucous at first then lustrous; blade 

very narrow, flattish to aculeiform and stiff, up to 

9 cm long and (when dry) ca 1 mm broad, glabrous 

throughout on both surfaces, the longitudinal 

nerves very few, strong, with transverse veinlets not 

at all apparent externally. Flowering culms leafless 

from the first. 

Inflorescences racemose (often reduced to a single 

spikelet), solitary or emerging in twos or threes 

and subtended at culm nodes and branch nodes by 

a common bract; prophylla none. Transitional 

glumes subhyaline, commonly 2 (sometimes 1 or 

3) up to 3 cm distant from each other and from 

the first fertile lemma, boat-shaped, acute or acuminate and awned at the apex, dorsally keeled and 

glabrous, I ca 10 mm long, 11 ca 11 mm long, 111 

ca 12 mm long, the uppermost transitional glume 

sometimes similar to the lemma and subtending a 

depauperate flower. Spikelets lanceate, pedicellate, 

up to 4 cm long, lax, compressed, comprising few 

to several perfect flowers. Rachilla segments tardily disarticulating, 2-3 mm long, curved, D-shaped 

in cross-section, prominently nerved, flared toward 

the apex, puberulous to subglabrous throughout. 

Lemma pergamineous, up to 13 mm long, severalnerved, keeled toward the acuminate apex and 

terminating in a short awn, dorsally glabrous, 

densely fringed with pale cilia on the outer margin. 

Palea shorter than its lemma, papery, oblong, emarginate and bi- or quadridentate at the broadly 

rounded apex, broadly sulcate dorsally, prominently nerved between and beyond the keels, 

fringed with pale cilia on the keels, elsewhere 

glabrous throughout. Lodicules (in the specimen) 

either opaque or pellucid or diaphanous, ciliate at 

the apex of the margin, elsewhere glabrous 

throughout; the anterior 2 sub-semiovate, the posterior one lanceate. Anthers up to 6 mm long, 

yellow. Ovary glabrous, the style glabrous, typically divided into 2 stigmatic branches, atypically 

bearing (in addition) an abortive stigmatic branch. 

Caryopsis unknown. 

TYPE-CoLLECTIoN.-Brazil, Matto Grosso, between Bonito and Rondonopolis, from a single 

flowering plant among scattered colonies of plants 

in the vegetative state growing in sandy soil, 8 iv 

1930. Agnes Chase 11886 (holotype: US 1500498, 

1500499). Additional specimens seen: between 

Rondonopolis and Santa Rita do Araguaya, from 

a plant in the vegetative state, representative of an 

element common in the local flora for several 

kilometers along the road, 13 iv 1930, Agnes Chase 

12007 (US 1500468 and 1500469). 

2. Apoclada cannavieira (Alvaro da Silveira) 

McClure, new combination 

FIGURE 3 

Arundinaria cannavieira Alvaro da Silveira, 1919: 101, fig. 2. 

TYPE-COLLECTION.-In campis arenosis siccisque in 

Serra do Caparao (as published; ‘Serra do Cabral’ 

as indicated in the field notes that accompany the 

type-collection) Minas Geraes, ubi sub nomine 

‘cannavieira,’ vulgo cognita est. Floret Novembri. 

Silveira 644 (holotype: Silveira herbarium), US 

(duplicate from the type-collection). 

3. Apoclada diversa 

FIGURE 4 

Apoclada diversa McClure and Smith, in Reitz, ed., 1967:62, 

fig. 10z-e.

TYPE-COLLECTION.-Brazil, Santa Catarina, Cacador, Rio Cacador, lugares limidos, 22 i 1946, 

Swallen 8271 (holotype: US 2152479 and 2383418). 

4. Apoclada simplex 

FIGURE 5 

Apoclada simplex McClure and Smith, in Reitz, ed., 1967:59, 

fig. 10s-y. 

FIGURE 3.-Apoclada cannavieira (Alvaro da Silveira) McClure. A, Upper portion of flowering culm, x 0.6; B, apex 

of leaf sheath and base of leaf blade, x 12; C, spikelet, 

x 1.2; D, diagram of longitudinal section of spikelet, x ca. 

2.4; E, transitional glume (sterile lemma), x 3.6; F, depauperate floret attached to rachilla segment (transitional 

glume [E], which subtended bud has been removed), 

x 6; G, floret, x 2.7; H, lemma, x 2.7; I, palea, X 2.7: J, 

stamen complement and lodicule complement, x 3.6; K, 

lodicule complement, x 6; L, stamen, X 3.6; M, gynwcium, 

x 14.4. All drawings based on Alvaro da Silveira $44 (US). 


NUMBER 9 15 

FIGURE 4.-Apoclada diuersa McClure and L. B. Smith. A, 

Upper portion of leafless flowering culm, x 0.6; B, bud complement from node above midculm level, x 3; C, base of 

branch complement, x 3; D, spikelet with antecedent part 

of stem, x 1.2; E, bract that subtends flowering branches, 

x 1.2; F, floret, x 7.2; G, lemma, x 7.2; H, palea, x 7.2; 

I, lodicule complement, x 15; J, stamen, x 15; K, gynoecium, 

x 15; L, fruit (juvenile), x 15. All drawings based on 

Swallen 8271 (US). 

TYPE-coLLEcrIoN.-Brazil, Santa Catarina, Campos Novos, Palmares, pinhal, 900 m, 27 xii 1963, 

Reitz and Klein 16355 (holotype: US 2434535). 

Arthrostylidium Ruprecht 

FIGURE 6 

Arthrostylidiuin Ruprecht, 1839:27; 1840:117.-Hance, 1862: 

235.-Munro 1868:39.-Doell, in Martius, 1880: 172.-Bentham, in Bentham and Hooker, 1883: 1208.-Hackel, in 

Engler and Prantl, ed., 1887:92.-Pilger, in Urban, ed., 

1900-1901 : 336.-Hackel, 1903a: 67.-Hitchcock, 1927b: 307; 

1936:243.-McClure, 1957:199. 

Plants unicespitose, unarmed. Rhizomes pachymorph. Culms either self-supporting or clambering; each midculm node bearing a single initial 

(primary) branch bud, the prophyllum in some 

species elongating simultaneously with the germination of the bud, the primordium producing a 

single segmented, terete primary axis, this dominant 

over axes of higher orders that usually proliferate 

promptly from conventionally subtended buds typically present at its own proximal nodes; the base 

of the primary axis typically extending downward 

beyond the locus of insertion of the prophyllum or 

lowermost circumcingent sheath to form a low, 

downward-tapered promontory, this promontory 

naked in a few species (as in the available specimens of the type-species), elsewhere bearing usually 

two distichously inserted buds or branches each 

subtended by a small, noncircumcingent bract. 

Branch complement (when developed) at midculm 

nodes of either restricted or gremial insertion, the 

primary member typically strongly to weakly dominant; sometimes, however, apical growth in the 

initial primordium fails to take place, even after 

the precocious germination of buds at its proximal 

nodes. Sheaths at midculm nodes with or without 

a persistent girdle. Leaf blades with transverse 

veinlets not at all manifest to more or less clearly 

visible. 

Inflorescences semelauctant, spicate racemes; 

subtending bracts and prophylla always lacking, 

pulvini at the base of the branches of the rachis 

usually lacking; transitional glumes 2 or 3, the first 

and second empty, the third a sterile lemma sometimes subtending a rudiment; spikelets sessile or 

subsessile, containing few to several perfect florets 

and terminating in a depauperate sterile anthecium; lemma of perfect florets subtending its palea 

only basally at maturity, palea gaping antically, 

broadly sulcate and 2-keeled dorsally; lodicules 3, 

the anterior 2 asymmetrical and paired, the posterior one smaller and symmetrical (exceptionally 

none, as recorded for A. cacuminis). Stamens 3, the 

filaments filiform, free. Stigmas 2. Mature fruitas far as known (available examples are rare)-an 

oblong mucronate, sulcate caryopsis, the pericarp 

pergamineous, of even thickness throughout, or appreciably thickened apically to form a distinct cap 

or nodule basal to the stylar column; the sulcus 

and the basal position of the embryo clearly manifest. 

ETYMoLoGY.-The name Arthrosty Zidium (n,), 

derived from the Greek, arthron, joint, and stylidium, column, alludes to the readily disarticulating 

nature of the rachilla, in spikelets of the lectotype 

and numerous other (but not all) species of the 

genus. 

TypE-spEcIEs.-Hitchcock (1927b: 307) designated 

Arthrostylidium cubense Ruprecht (1839:27) as the 

type of the genus (see McClure 1957:199). 

of the 43 trivial names that have previously been 

associated with it, the image of the genus Arthrostylidium still contains a few taxa that embody 

conspicuously deviant features. Since in my present 

perspective I characterize these features as “maverick” (see glossary for definition), I have not given 

them significance at the genus level. Some species 

still retained in the genus (e.g., Arthrostylidium 

schomburgkii) share such a feature with Aulon.emia 

q ueko, Glaziophy t on m irab ile, R h ipidoclad um 

harmonicum, and several species of Myriocladus. 

I refer to a disproportionate elongation of at least 

the first above-ground internode, and usually some 

internodes of primary branches as well. These 

elongated internodes are followed by one or more 

aborted (obsolete) internodes, the successive ones 

being separated from each other by normal nodes 

RELATIONSHIPS.-EVen after the exchsion of 23 

NUMBER 9 17 

FIGURE 5.-Apoclada simplex McClure and L. B. Smith. A, 

Portion of clump showing rhizomes and base of culms, x 

0.12; 6, midculm internode, nodes and culm sheath, x 0.6; 

c, apex of sheath from lower culm node, adaxial aspect, 

x 1.2; D, apex of sheath from higher node, adaxial aspect, 

x 1.2: E, midculm complement of leafy flowering branches, 

x ca. 0.6; F, variant of midculm branch complement, x ca. 0.6; G, apex of leaf sheath and base of blade, x ca. 4.8; 

H and I, spikelets, showing reduced laminiferous foliage leaf 

standing in place of a first transitional glume, X 3; J, 

sterile lemma (either standing empty or subtending a depauperate flower) as a second transitional glume (that an 

be either a sterile lemma or a fertile lemma), x 6; K, floret 

from middle of spikelet, x 6 L, palea, x 6; M, lodicule 

complement, x ca. 18; N, stamen, x 15: 0, gynoecium, x 

15; P, fruit, embryo side, showing weak indication of the 

position of the embryo, x 7.2; Q, fruit, hilum side, showing 

the sulcus, x 7.2; R, fruit, longitudinal section, x 7.2. Drawings A, H based on Klein 4108 (US), B-F and I-R on Reitz 

and Klein 16,355 (US), and G on Klein 4657 (US). 

bearing normal sheaths and branch buds (or 

branches) inserted distichously (cf. McClure, in 

Maguire, Wurdack, et al., 1964:2). Available specimens of Arthrostylidium fimbriatum show several 

vegetative features that suggest introgression from 

(or a possibly recessive genetic heritage shared by) 

some members of the genus Myriocladus. I retain 

this species in Arthrostylidium on the basis of the 

characteristic features of the inflorescence. The 

inflorescence in Arthrostylidium angustifolium 

deviates from the spicate form of the inflorescence 

of the type-species of the genus by substituting a 

spicate raceme for each spikelet. I retain this species in Arthrostylidium on the basis of a characteristic feature complex of the midculm branch 

complement. Arthrostylidium excelsum, A. sarmentosum, and A. venezuelae produce spicate racemes 

with a geniculate rachis (inflorescences technically 

identical with those of Rhipidocladum harmonicum) but the same characteristic feature complex 

of their midculm branch complements keeps them 

in Arthrostylidium. 

In terms of the ontogeny and morphology of the 

midculm branch complements, the taxa I retain in 

Arthrostylidivm display a cline, one extreme of 

which is found in A. cubense and the other in A. 

sarmentosum. The central structure of this feature 

complex is the promontory (Figure SD), a tapered 

bulge that extends downward from the locus of 

initiation of the primoridum of the solitary primary bud at midculm nodes. In available specimens of A. cubense the promontory lacks lateral 

buds; in A. ekrnanii it sometimes bears one bud; in 

A. capillifolium it sometimes bears two buds; in 

A. urbanii it sometimes bears four buds. The buds 

that are borne on this outwardly unsegmented promontory are distichously inserted and subtended 

by noncircumcingent bract-like sheaths, while the 

buds that are borne more distally, on the proximal 

nodes of the segmented, terete primary axis that 

develops from the apical meristem of the initial 

primordium are subtended by circumcingent 

sheaths. As the number of buds on the unsegmented promontory increases, the number of circumcingent sheaths subtending buds on the 

proximal part of the primary axis decreases until, 

in Arthrostylidium sarmentosum, all the buds (excepting the uppermost one) are subtended by noncircumcingent bract-like sheaths. In some specimens 

of Arthrostylidium sarmentosum from Venezuela 

and Puerto Rico the development of the strongly 

dominant primary axis of the midculm branch complement is aborted immediately above the bud or 

branch at its uppermost proximal node. In other 

specimens, the dominance of the primary axis over 

the proximal secondary axes is greatly reduced. In 

such a specimen-as for example in Soderstrom 

1051 from Trinidad-the fully developed midculm 

branch complement is deceptively similar to that 

characteristic of all taxa I include in the genus 

Rhipidocladum. The persistence in arthrostylidioid 

taxa of the promontory (a feature that is entirely 

lacking in rhipidocladoid taxa), however, suggests 

that we have, in the vegetative cline observed in arthrostylidioid taxa, a case of evolution convergent 

toward Rhipidocladum with respect to the described feature complex. This convergence is weakly 

paralleled by the appearance, mentioned above, of a 

geniculate rachis in the inflorescence of Arthrostylidium excelsum, A. venezuelae, and A. sarmentosum, a feature shared by Rhipidocladum solely 

in R. harmonicum. 

As initially delimited by Ruprecht, Arthrostylidium was already highly polymorphic. In the Latin 

notes that follow his brief formal description he 

(Ruprecht, 1840: 117) says: “Genus Chusqueas et 

Arundinarias jungens.” For years, I read this as 

meaning simply that Arthrostylidium is a genus 

that falls between Chusquea and Arundinaria. Although Ruprecht did not actually transfer any 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 9 19 

FIGURE 6.-Arthrostylidium cubense Ruprecht (A-M) and Arthrostylidiurn rnultispicaturn Pilger (N-Q). A, Rhizome and 

lower part of culm, x 0.6; B, leafy flowering midculm nodes, 

x 0.6; c, apex of leaf sheath and base of blade, x a. 3; 

D, insertion of components of midculm branch complement, 

x 3; E, inflorescence, x 6; F, spikelet, x 6; G, transitional 

glumes (3), x 6; H, floret, x 6; I, lemma, x 6; J, palea, 

x 6; K, lodicule complement, x 15; L, stamen, X 15; M, 

gynoecium, x 15; N, fruit, embryo side, x 12; 0, fruit, hilum 

side, x 12; P, fruit, oblique aspect, x 12; Q, fruit, longitudinal section, x 12. Drawings A and H-M based on Brother 

Leon 16214 (US), B-D on Ekrnan 12649 (US), E-G on Wright 

3809 (US) and N-Q on Leonard 3817 (US). 

names from either Chusquea or Arundinaria to 

Arthrostylidium, he continues in a vein that conveys the impression that he does, in fact, consider 

that known members of these three genera actually 

form a continuum. Doell seems to have taken this 

view, since he was led, apparently by Ruprecht’s 

comparison of Chusquea leptophylla to Arthrostylidium trinii, to publish the combination Arthrostylidium leptophyllum (Nees) Doell (in 

Martius, 1880: 175). Ruprecht’s image of Arundin,aria was clearly derived from Nees (1834) and not 

from Michaux (1803). 

It appears that the natural boundary (disjunction) between Arundinaria and Arthrostylidium 

was obscured for Ruprecht (1839)-and has remained so for other taxonomists, down to the present-by the image of the genus Arundinaria which 

Nees produced (1834:478-483) when he incorporated it in such morphologically (and generically) 

divergent species as Arundinaria falcata, A. verticillata, A. amplissima, A. macrostachys, A. pinifolia, A. wightiana, and A. glaucescens. Every one 

of these species is excluded from Arundinaria by 

my circumscription of the genus. Hance (1862:235; 

1876:340) comments upon the weakness of the published bases for the differentiation of Arthrmtylidium and Arundinaria from each other. Major 

treatises by Munro (1868), Bentham (in Bentham 

and Hooker, 1883), Hackel (in Engler and Prantl, 

ed., 1887), and E. -G. Camus (1913) are unanimous 

in their perpetuation of the earlier failure to establish a clear distinction between the respective 

groups of New World bamboos that truly represent 

these two genera. 

The Latin notes that follow Ruprecht’s brief formal description of the genus Arthrostylidium 

(1839:27) read in free translation as follows: 

Natives of tropical America. This genus unites the Chusqueas and the Armdinaria. Among the latter especially, 

fruit-bearing flowers of A[rundinaria] amplissima and A. 

wightiana absciss, while in the rest [of Arundinaria] and in 

Chusquea, the flowers adhere tenaciously to the rachilla or 

“stylidium.” A[rthrostylidium] maculatum, by the sexual difference within the dimorphic spikelets, provides a full notion of the genus, [a notion] gradually obliterated in the 

rest [of the species] by way of A[rthrostylidium] cubense. 

Apparently begun by Ruprecht in 1839, the 

search for a natural boundary (disjunction) between Arthrostylidium and “Arundinaria” has remained fruitless for more than 130 years. The 

sense of futility induced by this perennial failure 

is revealed dramatically in several taxonomic papers that carry the quest into the 20th centuryamong them Pilger (in Urban, ed., 1900-1901: 

336-337), and Hackel (1903a:67-70). Pilger (in 

Urban, ed., 1907:289) gave renewed expression to 

his bewilderment in the following words (translated from the German): 

It now seems to me doubtful whether the genus Arthrostylidiurn has any justification, or should go into Arundinaria. Hackel (1903[a]:67 ff) has already transferred most 

of the genus Arthrostylidium to Arundinaria, leaving in 

Arthrostylidium only the forms with a doniventral inflorescence ([as in] A. Prestwi Munro from the West Indies). I do not believe that this difference is sufficient to 

justify the genus; a future monograph may well have to 

unite the two genera. 

Hackel’s tentative proposal (1903a: 69) to clarify 

the concept of the genus Arthrostylidium by limiting its content to the taxa represented by A. trinii 

Ruprecht, A. racemiflorum Steudel, A. presto& 

Munro, A. pittieri Hackel-“and perhaps also A. 

fim briatum Grisebach’l-has been universally and 

persistently ignored. A wholly different image of 

the genus was adumbrated by Hitchcock’s (1927b: 

307) selection of Arthrostylidium cubense Ruprecht 

as lectotype. But the boundary between Arthrostylidium and Arundinaria was still left undefined. 

Pilger and Hackel both revealed, in print, a mutually held conviction that if the basis for a taxonomic boundary between Arthrostylidium and 

Arundinaria really exists it must be sought amongst 

the morphological diversities that characterize the 

reproductive structures of the respective members 

of the complex. In his key to bamboo genera 

Hackel (in Engler and Prantl, ed., 1887:92) proposed the separation of the then current images of 

Arundinaria and Arthrostylidium solely on the ba- 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

sis of the (unreliable because unstable) number of 

transitional glumes (empty glumes alone or empty 

glumes and sterile lemmas) borne at the base of 

each spikelet. 

The natural boundary between Arthrostylidium 

Ruprecht and Arundinaria Michaux still awaits 

explicit definition. In morphological terms, plants 

typical of Arthrostylidium are distinguishable from 

plants typical of Arundinaria by the possession of 

pachymorph rhizomes, a unicespitose clump habit, 

a spicate inflorescence with sessile or subsessile 

spikelets, and binate stigmas; while plants typical 

of Arundinaria possess leptomorph rhizomes, a diffuse or pluricespitose clump habit, a paniculate or 

racemose inflorescence with pedicellate spikelets, 

and ternate stigmas. The respective areas of the 

natural distribution of plants of the two genera are 

disjunct, both in space and in respect to prevailing 

temperature minima; plants of the genus Arthrostylidium being confined to frost-free sites, while 

the genus Arundinaria is made up of frost-hardy 

plants. 

DISmIBuTIoN.-The twenty named New World 

species I have retained in the genus Arthrostylidium are all endemic to regions with a mesophytic, 

tropical or subtropical climate. Their aggregate 

recorded geographic range extends from Brazil, 

with one known species, to Venezuela, with four 

known species, to Cuba, with eight known species, 

and several other Caribbean islands, each with one 

or two species. Pilger (in Urban, ed., 1900-1901: 

337) stresses the narrowly limited known distribution of some of the taxa he calls “island species.” 

Annotated Checklist of New World Species 

Included in the Genus Arthrostylidium 

As here conceived and circumscribed, the genus 

Arthrostylidium embraces the following named 

taxa, all of which are endemic to the New World. 

1. Arthrostylidium angustifolium Nash, 1903: 172. 

2. Arthrostylidium cacuminis McClure, in Maguire, Wurdack, et al., 1964:3. 

3. Arthrostylidium capillifolium Grisebach, 1862: 

531. 

Arundinaria capillifolia (Grisebach) Hackel, 1903a: 

69. 

4. Arthrostylidium cubense Ruprecht, 1839:28, 

pl. iv:fig. 13 (Figure 6 A-M). 

Arundinaria cubense (Ruprecht) Hackel, 190Ja:69. 

5. Arthrostylidium distichum Pilger, in Urban, 

ed., 1900-1901:342. 

6. Arthrostylidium ekmanii Hitchcock, 1936: 16. 

7. Arthrostylidium excelsum Grisebach, 1864:529. 

Arundinaria excelsa (Grisebach) Hackel, 1903a:69. 

As represented by currently available herbarium 

specimens, Arthrosty lidium excelsum does not appear to be clearly differentiated (disjunct) from 

Arthrostylidium venezuelae. 

8. Arthrostylidium fmbriatum Grisebach, 1862: 

9. Arthrostylidium haitiense (Pilger) Hitchcock 

Arundinaria haitiensis Pilger, in Urban, ed., 1907: 

288. 

10. Arthrostylidium longiflorum Munro, 1868:41. 

Arundinaria longiflora (Munro) Hackel, 1903a:69. 

Guadua exalata Doell, in Martius, 1880:181. 

11. Arthrostylidium multispicatum Pilger, in Urban, ed., 1900-1901:341. 

Arundinaria multispicata (Pilger) Hackel, 1903a: 

69. 

12. Arthrostylidium obtusatum Pilger, in Urban, 

ed., 1900-1901 :340. 

Arundinaria obtusata (Pilger) Hackel, 1903a:69. 

13. Arthrostylidium pubescens Ruprecht, 1839:29, 

Arundinaria pubescens (Ruprecht) Hackel, 1903a: 

69. 

14. Arthrostylidium reflexum Hitchcock and Ekman, in Hitchcock, 1936: 19. 

15. Arthrostylidium sarmen.tosum Pilger, in Urban, ed., 1903:108; Chase, 1914:278, pl. 21. 

16. Arthrostylidium scandens McClure, in Maguire, Wurdack, et al., 1964:4. 

17. Arthrostylidium schomburgkii (Bennett) Munro, 1868:41. 

Arundinaria schomburgkii Bennett, in Schomburgk, 1841:562. 

1 8. A r t h rosty lid iu m simp liciuscu lum (Pilger) McClure, new combination. 

Arundinaria simpliciuscula Pilger, 1920:29. 

19. Arthrostylidium urbanii Pilger, in Urban, ed., 

1900-1901 :339. 

531. 

and Chase, 1917:399. 

pl. iv: fig. 14. 
NUMBER 9 21 

20. Arthrostylidium venezuelae (Steudel) McClure, 

Chusquea ? venezuelae Steudel, 1854:337. 

?Arundinaria standleyi Hitchcock, 1927a:79. 

As illustrated by currently available herbarium 

specimens, Arundinaria standleyi appears to be 

only weakly differentiated from Arthrostylidium 

venezuelae by the sparse dorsal vesture of its lemmas in the form of lustrous white appressed or 

spreading deciduous hairs. 

1942172. 

Checklist of New World Species Here Excluded 

from the Genus Arthrostylidium 

1. Arthrostylidium ampliflorum. See Rhipido2. Arthrostylidium amphimum. See Aulonemia 

3. Arthrostylidium angustiflorum. See Rhipido4. Arthrostylidium aridaturn. See Aulonemia 

5. Arthrostylidium bartlettii. See Rhipidocladum 

6. Arthrostylidium burchellii. See Colanthelia 

7. Arthrostylidium eflusum. See Aulonemia ef8. Arthrostylidium geminatum. See Rhipidocla9. Arthrostylidium haenkei. See Aulonemia 

10. Arthrostylidium harmonicum. See Rhipidocla11. Arthrostylidium leptophyllum is Chusquea 

12. Arthrostylidium longifolium. See Bambusa (sg. 

13. Arthrostylidium maculatum. See Aulonemia 

14. Arthrostylidium maxonii. See Rhipidocladum 

15. Arthrostylidium pittieri. See Rhipidocladum 

16. Arthrostylidium prestoei. See Rhipidocladum 

17. Arthrostylidium purpuratum. See Aulonemia 

cladum ampliflorum. 

amp lcssima. 

cladum angustiflorum. 

setigera. 

bartlettii. 

burchellii. 

fusa. 

dum geminatum. 

haenkei. 

dum harmonicum. 

leptophylla Nees. 

Guadua) longifolia. 

parui flora. 

maxonii. 

pittieri. 

prestoei. 

purpurata. 

18. 

19. 

20. 

21. 

22. 

23. 

Arthrostylidium queko (as quexo). See AulmeArthrostylidium racemiflorum. See Rhipidocladum racemiflorum. 

Arthrostylidium spinosum. See Bambusa (GuaArthrostylidium steyermarkii. See Aulonemia 

Arthrostylidium subpectinatum. See Aulonemia 

Arthrostylidium trinii. See Rhipidocladum parmia queko. 

dua) longifolia. 

stey ermarkii. 

su bpectinata. 

viflorum. 

ArundinarM Michaux 

FICURIB 7-18 

Arundinaria Michaux, 1803, 1:73.-McClure, 1957:200. 

Miegia Persoon, 1805, 1:lOl.-McClure, 1957:205. 

Ludolfia Willdenow, 1808:320.-McClure, 1957:204. 

Macronax Rafinesque, 1808:353.-McClure, 1957:204. 

Triglossum Fischer, 1812:vi.-McClure, 1957:210. 

Pleioblastus Nakai, 1925: 145.-McClure, 1957:207. 

Nipponobambusa Muroi, 1940:89.-McClure, 1960:194. 

Nipponocalamus Nakai, 1942:350; 1951:326.-McClure, 1957: 

206. 

The literature that deals with Arundinaria-even 

when the genus is viewed solely in terms of the few 

New World taxa that I recognize as falling within 

its natural boundaries-comprises many individual 

items. Repeated consultation and attentive study 

of this literature has convinced me that it neither 

contains nor conveys the documented information 

needed for the construction of a clear image of 

the New World components and the natural 

boundaries of the genus. For this reason, instead 

of listing all or any major part of it here, I have 

selected-and reserved for notice in the text-those 

items that appear to have a clear pertinence to the 

thread of the present account and to be least likely 

to confuse the reader. 

Plants of diffuse habit (both diffuse and pluricespitose in some species); all axes unarmed. Rhizomes leptomorph. Culms self-supporting, either 

remaining solitary, or tillering from subterranean 

buds-with or without the intercalation of a metamorph axis, the internodes fistular, either terete 

throughout or more or less markedly sulcateusually so for only a short distance upward from 

the locus of insertion of a bud or a branch complement, rarely (as occasionally seen in plants of 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Awndinaria gigantea ssp. gigantea) lightly so all 

the way from one node to the next. Primary 

branch buds at culm nodes solitary, each containing 

but a single initial primordium. Branch complement at midculm nodes in some species facultativerestricted monoclade, (rarely, either temporarily or 

permanently, lacking throughout) typically unrestricted monoclade and always of restricted insertion, with the primary member dominant. Leaves 

(blades of leaf sheaths) petiolate, with transverse 

veinlets clearly visible on both surfaces initially, 

but sometimes becoming obscure in old leaves. 

Inflorescences semelauctant, generally of open 

racemose or paniculate branching, sometimes assuming both of these forms, or even reduced to a 

single spikelet, all in the same plant. Rachis either 

deliquescent or excurrent, its branches typically not 

prophyllate, the lowermost primary ones always, 

the upper ones commonly, subtended by either a 

small or rudimentary bract, or a line of hairs. 

Transitional glumes at the base of each spikelet, 

usually 2 and empty (the first one sometimes 

lacking), rarely 3, the third (conventionally called 

a sterile lemma) sometimes empty, more commonly 

subtending a depauperate flower. Spikelets comprising several to many perfect florets, and terminating 

acuminately in progressively depauperate sterile 

florets or empty anthecia. Rachilla segments potentionally disarticulating just below the locus of 

insertion of each fertile lemma. Lemma (when 

subtending a functional flower) fully embracing 

the palea only basally at maturity. Palea broadly 

sulcate and 2-keeled dorsally, the margins not at 

all or only slightly and partially overlapping. 

Lodicules 3, the anterior two typically more or less 

strongly asymmetrical and paired, the posterior one 

symmetrical and usually smaller (at least either 

shorter or narrower) than the anterior two. Stamens typically 3, the filaments filiform, free. 

Stigmas 3. Mature fruit an oblong, sulcate, glabrous 

caryopsis, terminating apically in the persistent base 

of the style; the pericap thin, pergamineous or 

coriaceous, in some species appreciably thickened 

at the apex of the fruit; the contours of the basal 

embryo clearly revealed as a rule in the dry fruit 

by the embryotegium. 

ETYMoLoGY.-The name Arundinaria is derived 

from the Latin, urundo (arundin-), reed or cane, 

and the suffix, aria, that signifies belonging to. 

From 1829 (Kunth, 1829:137) at least until 1861 

(Bentham, 1861 :433) predominant usage attributed 

the authorship of the genus Arundinaria to L. C. 

Richard. In the absence of positive evidence to 

the contrary, however, Michaux is now generally 

recognized as the author of it. 

TYPE-spECIEs.-Arundinaria gigantea (Walter) 

Muhlenberg (1813). 

As basionym, Arundo gigantea Walter (1788), 

takes precedence over Festuca grandiflora Lamarck 

(1791), and Arundinaria macrosperma Michaux 

(1803) at the species level, since both Lamarck’s 

and Michaux’s species are here included in a 

comprehensive circumscription of Arundinaria gigantea (ICBN 1966:Art. 57). 

RELATIONSHIPS.-From the polymorphic neotropical genus, Arthrostylidium (q.v.), numerous species 

of which have at one time or another been misplaced in Arundinaria, taxa characteristic of Arundinaria are distinguishable by the following 

invariable combinations of morphological features: 

leptomorph rhizomes, diffuse or pluricespitose 

clump habit, and ternate stigmas. Moreover, the 

respective geographical distributions of the two 

genera are different and disjunct, and are correlated 

with distinctive ecological adaptations. As far as 

known, plants of the genus Arthrostylidium are 

confined to frost-free sites, while the genus Arundinaria comprises plants that are frost-hardy. 

Included by some authors in Arundinaria, the 

recognized components of the Old World genus 

Thamnocalamus are set off from this genus by the 

combination of pachymorph rhizomes, unicespitose 

clump habit, and well-developed, sheath-like bracts 

by which the lower branches of each inflorescence 

are subtended. 

Of the known African species at present allocated 

to Arundinaria, A. alpina K. Schumann (at least) is 

excluded from this genus by its pachymorph rhizomes, a unicespitose clump habit, and binate 

stigmas; cf Yushania sg. Otatea, p. 116. 

DISTRIBuTION.-The known New World components of the genus Arundinaria constitute the polymorphic type-species whose natural distribution is 

limited to continental United States. They are 

represented principally in North American, British, 

and European herbaria by specimens gathered in 

Alabama, Arkansas, Delaware, Florida, Illinois, 

Indiana, Kentucky, Louisiana, Maryland, Missis- 
NUMBER 9 23 

sippi, Missouri, North Carolina, Ohio, Oklahoma, 

South Carolina, Tennessee, Texas, Virginia, and 

West Virginia. 

A map of southeastern United States showing, 

by counties, the observed incidence of stands of 

native Arundinaria-under the name Arundinaria 

tecta applied in a comprehensive sense-is presented 

by West (1935:258, fig. 1). Owing to the difficulties 

involved in reconciling incongruities and inconsistencies between various published taxonomic 

usages-difficulties augmented by the fragmentary 

nature of most of the available preserved specimens-it is not yet possible to construct authentic 

maps of the precise distribution of stands of the 

respective subspecific entities herein recognized. A 

general idea of the pattern of their distribution, 

however, may be derived from Gilly's map (1943, 

fig. 1) by assuming a rough (but not precise) correspondence between Gilly's "Mississippi-type'' and 

my A. gigantea ssp. gigantea; between Gilly's "Atlantic-type" and my subspecies tecta; and between 

Gilly's "Intermediates" and my subspecies macrosperma (cf. Figure 16). 

Known Old World taxa that I recognize as 

members of the genus Arundinaria are found in 

China proper, Hainan Island, northern India, 

Japan, Korea, Madagascar, Nepal, the Ryukyu 

Archipelago, Sikkim, Taiwan, Tibet, and northern 

Vietnam. 

THE NEW WORLD COMPONENT OF THE GENUS 

Awndinaria.-The only known New World bamboos embraced by my circumscription of the genus 

Arundinaria are those included in its polymorphic 

type-species, Arundinaria gigantea (Walter) Muhlenberg, sensu lato. The development of an improved idea of the geographical boundaries of the 

genus and its known content requires, therefore, 

a critical examination of taxa that have hitherto 

been assigned to it from the bamboo floras of the 

Old World, as well as those of the New. 

During the interval since Arundinaria was first 

described and given monotypic status by Michaux 

in 1803, the number of species assigned to the 

genus increased spectacularly. The inventory of 

names linked to it in the taxonomic literature 

grew from the initially solitary entry, Arundinaria 

macrosperma Michaux, to a total of about 376, 

counting only binomials. The inclusion of trinomials swells the number of its published nomenclatural diversities to about 482. Of the New World 

component of this total, I have excluded the taxa 

represented by 50-odd names listed on page 36 

et seq. 

As a result of interim studies and revised judgments published by other authors, many of the 

Old World species originally (or at one time) 

incorporated in the genus Arundinaria have subsequently been made either the respective types, 

or components, of the following genera: Brachystachyum, Chimonobambusa, Indocalamus, Neosasamorphia, Oreocalamus, Pseudosasa, Sasa, 

Sasaella, Sasamorpha, Semiarundinaria, Sinarundinaria, Sinobambusa, Thamnocalamus, and Yushania. All of the bamboos that have since been 

allocated to these genera are endemic to Old World 

areas. 

Annotated Checklist of New World Elements 

Included in the Genus Arundinaria 

(as herein defined) 

Arundinaria gigantea (Walter) Muhlenberg, 1813: 

14. Basionym: Arundo gigantea Walter, 1788:81. 

Arundo tecta Walter, 1788:81. 

"Festuca grandiflora Lamarck, 1791:191. 

Arundinaria macrosperma Michaux, 1803, I:74. 

* Triglossum bambusinum Fischer, 1812:6. 

Arundinaria tecta (Walter) Muhlenberg, 1813: 14. 

"Arundinaria bambusina (Fischer) Trinius, 1820: 

"Miegia pumila Nuttall, 1837: 149 [illegitimate 

"Arundinaria tecta p pumila Ruprecht, 1839:22. 

"Arundinaria tecta y distachya Ruprecht, 1839:22. 

*Awndinaria tecta 8? colorata Ruprecht, 1839:22. 

"Arundinaria macrosperma a arborescens Munro, 

"Arundinaria macrosperma p suffruticosus Munro, 

"Armdinaria macrosperma tecta Wood, 1871: 

"A,.undinaria gigantea tecta (Walter) Scribner, in 

"Armdinaria tecta var. decidua Beadle, in Bailey, 

"Given the present unsatisfactory state of their 

description, their documentation, and their typification, vis-8-vis the highly polymorphic character 

of the populations here included under Arundi97. 

name]. 

1868: 15. 

1868:15. 

404. 

Kearney, 1893: 478. 

1914:446. 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

naria gigantea, sensu lato, it is my judgment that 

proposals for a more refined formal disposition of 

the taxa represented by these names should be 

deferred until comprehensive studies on a broad 

disciplinary spectrum in field and laboratory have 

improved present perspectives. 

Proposed Taxonomic Revision 

of the Type-species of Arundinaria 

The relegation by Hitchcock (1935:29 [1951:27]) 

of the name Arundinaria macrosperma Michaux to 

synonymy under Arundinaria gigantea (Walter) 

Muhlenberg appears to be referable to an earlier 

tentative decision expressed in the following paragraph quoted from page 156 of a paper entitled 

“Types of American Grasses” (Hitchcock, 1908). 

Arundinaria macrosperma Michaux “Gramen altissimum 

ramosum a Virginia ad Floridam & in occidentalibus juxta 

fluviis ab Illinoensibus ad ostium Misissipi [sign for undershrub].” The specimen is fragmentary and one can not be 

certain which species of Arundinaria it represents. Michaux 

probably included the large and the small canes in one 

species. As he described the plants as being very high, we 

may retain this name for the tall cane, as is done in our 

manuals. 

Hitchcock’s disposition of the matter of synonymy 

appears to have been influenced by the reiterated 

allusion to size-in the trivial name, gigantea, of 

one taxon, and in the description of the other, 

macrosperma, as “altissima.” In other words, it is 

based upon an interpretation of Michaux’s description of the monotypic genus Arundinaria, and not 

upon an interpretation of the lectoholotype by 

completing the image of its botanical source (cf. 

Rickett and Camp, 1950). At the same time, however, the taxon represented by Arundinaria tecta 

(Walter) Muhlenberg was maintained by Hitchcock as a distinct species. 

In present perspectives it appears that the taxa 

represented by the binomials Arundinaria gigantea 

(Walter) Muhlenberg, Arundlnaria tecta (Walter) 

Muhlenberg, and Arundinaria macrosperma Michaux embody, respectively, three components of a 

polymorphic array of populations (cf. Figures 16, 

17). As shown by their purest available typifications, 

Arun.dinaria gigantea and Arundinaria tecta are 

clearly set off from each other by distinctive combinations of strongly contrasting morphological and 

ontogenetic features (Figure 15). The lectotype 

and isotype of the taxon Arundinaria macrosperma 

are seen as representing a plant that falls within 

the series of variants (putative nothomorphs) that 

form a cline bridging the morphological and ontogenetic divergency that exists between Arundinaria 

gigantea and Arundinaria tecta (Figures 17, 18). 

Circumstantial evidence of the biological relationships of these three taxa supports the proposal 

that they be treated taxonomically as members of 

a single polymorphic species. 

When the circumscription of Arundinaria macrosperma Michaux is interpreted as including the 

type of Arundo gigantea Walter, the adoption of 

the name Arundinaria gigantea (Walter) Muhlenberg in place of Arundinaria macrosperma Michaux 

for the type-species of the genus Arundinaria 

(Hitchcock, 1935:29 [1951:27]) has the sanction of 

the International Code of Botanical Nomenclature 

(see Lanjouw et al, ed., 1966: Art. 57). This precedent determines the binomial to be adopted when 

the three taxa now under discussion are treated as 

subspecies under one specific name. 

In present perspectives, any specimen that bears 

a recorded origin indicating nativity to continental 

United States, and that shows features of both 

subspecies gigantea and subspecies tecta-even 

though it is otherwise incomplete-may be determined as subspecies macrosperma. 

The subspecific identification placed on incomplete specimens that show only features characteristic of either subspecies gigantea or subspecies tecta 

should be followed by a query (?), since the name 

of the subspecific taxon to which the character of 

the missing structure (s) would carry the identification, by way of the key, must remain uncertain 

in such cases. Where, on account of the incompleteness of a specimen, uncertainties arise as to its 

correct subspecific disposition, it may with propriety 

be filed tentatively as Arundinaria gigantea, sensu 

lato. However, the possibility that more intensive 

field studies in the United States may bring to light 

native plants specifically distinct from Arundinaria 

gigantea, as this taxon is here circumscribed, should 

not be ignored. 

Concerning the Typification 

of Arundo gigantea, Arundo tecta, and 

Arundin,aria macrosperma 

Of all of the many previously published treatises 
%UMBER 9 

Key to the Herein Recognized Subspecies of Arundinuria gigantea, sensu lato 

la. Rhizomes without air canals: typically of steadily diageotropic growth, i.e., normally not 

turning up at the apex to form a culm. Plants of diffuse habit, the culms normally 

solitary and not tillering. Midculm sheaths each shorter than the corresponding internode; deciduous. Primary axis of midculm branch complements not constricted basally, 

lacking a “neck” (see glossary). The lowest bud (or secondary branch) of midculm 

pleioclade) branch complements inserted but a short distance above the base of the 

first internode of the primary branch and (in contrast to those inserted at its succeeding 

nodes) not subtended by a conventional sheath. The primary and secondary axes of 

midculm branch complements relatively short and slender, all appressed basally and 

then curving strongly away from the culm. Culms with their leafy branch complements 

fully developed present a narrowly linear profile. Leaf blades pubescent on the abaxial 

surface; subglabrous on the adaxial. Lemmas dorsally hirsute with antrorse hairs and 

minutely hispidulous or scaberulous; the exposed exterior surface pale green (not tinted 

with wine) and more or less conspicuously glaucous when fresh; transverse veinlets 

clearly manifest externally. Lodicules typically subopaque, fringed with marginal cilia, 

and suppfied with numerous vascular traces. Pistils lacking the uncinate rudimentary 

stylar branch that is regularly present on the pistil in the neotype of subspecies tecta. 

Fruits narrowly conical apically and terminating in the persistent base of the style 

(Figures 7-11) ........................................ ........................... 1. subspecies gigantea 

lb. Rhizomes with air canals forming a continuous and unbroken cylinder; apical growth 

commonly diageotropic (horizontal) for only a short distance, then directed upward 

to form a culm. Plants of diffuse habit (culms arising from lateral buds of the rhizomes 

normally remaining solitary) and also pluricespitose as a result of the tillering of culms 

(principally those that are apical to rhizomes). Midculm sheaths each longer than the 

corresponding internode: not deciduous. Primary axis of midculm branch complements 

constricted basally into an obconical “neck” consisting of usually 2 or 3 very short 

internodes with budless sheath-bearing nodes (Figure 12~) , followed by more elongated 

internodes each bearing basally a secondary member of the branch complement (or a 

bud) subtended by a conventional sheath. The primary and secondary axes of midculm 

branch complements elongate, appressed basally, then ascending and, when fully developed, curving broadly away from the culm. Culms with their leafy branch complements fully developed present a lanceolate profile. Leaf blades densely pubescent on 

both surfaces. Lemmas dorsally glabrous or nearly so, the exposed exterior surface 

tinted wtih wine, and not glaucous when fresh; transverse veinlets not at all manifest 

or only barely perceptible. Lodicules thin, transparent, with glabrous margins and 

vascular traces weakly to scarcely manifest. Pistils all showing the unciform rudiment 

that becomes a conspicuous feature in mature fruits. Fruit dome-shaped apically, 

terminating in a short beak, and regularly bearing (just below the base of the beak, 

and directly above the tip of the sulcus) a usually more or less strongly unciform rudiment-apparently representing a supernumerary style or stylar branch (Figures 12-14) 

................................................................................................. 2. subspecies tecta, new status 

lc. Rhizomes with air canals either present or absent; when present, either continuous or 

discontinuous; other individual morphological and ontogenetic features appearing in 

diverse recombinations of either identical or intermediate expressions of the same contrasting features that (in characteristic combinations described above) clearly differentiate subspecies gigantea and subspecies tecta . , , , , . , . ,3. subspecies macrosperma, new status 

26 

*Edna Rema Walker (1906) reports and illustrates the occurrence of rudimentary stylebranches in a number of species of grasses, including Oryza sativa. See also Arber, 1934, fig. 

88. I have not found any published notice of the occurrence of this feature in any bamboos. 

that have been concerned with the clarification of Munro’s monograph (1868)-Hitchcock‘s publicathe taxonomy of the New’World components of the tions (1905, 1908) proposing lectotypes for Arundo 

genus Arundinaria-including not only local floras gigantea Walter and Arundin.aria macrosperma 

and manuals, but such taxonomic treatises as Michaux, are the only ones that are recognizable 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

as representing a disciplined conformity to the 

spirit and the letter of the pertinent references in 

the International Code of Botanical Nomenclature. 

In the selection of a neotype for Arundo tecta, 

and living plants (hypotypes, sensu Frizzel, 1933: 

653) to supplement the original images of subspecies 

gigantea, tecta, and macrosperma, I have followed 

the stipulations of the International Code of 

Botanical Nomenclature (1966) as set forth in note 

3 under Art. 7; and in paragraphs 3, 4, and 5 

under the heading, Guide for the determination 

of types (Lanjouw, et al., ed., 1966:71). The protolog 

pertaining to each of these three taxa lacks a precise 

indication of type-locality. However, I believe that 

the plants selected present details that authentically supplement the sketchy images of these taxa 

provided by their respective protologs. According 

to Ewan (1969:198) “Thomas Walter’s new species 

in his Flora Caroliniana (1788) are generally assumed to be descriptions of plants that grew about 

his plantation at the mouth of the Santee River. 

But we must remember that John Fraser was 

Walter’s advocate and ‘well spring in the wilderness’ 

who brought plants for Walter to study and 

describe.” 

In the present limited state of our knowledge of 

the component living plants, and in the restricted 

perspective afforded by their fragmentary representation in extant herbarium specimens, the apparently clinal nature (composition) of presumably 

hybrid populations from which these three taxa 

have been selected renders their more refined 

systematic treatment infeasible as yet (cf Bocher, 

1963: 11-12, 1967:258; Stebbins, 1940; Heiser, 1949; 

and Allan, 1949). 

1. Arundinaria gigantea ssp. gigantea 

FIGURES 7-11, 15~, 16~ 

HoLoTY PE.-Walter Herbarium (BM) ; Walter 

s.n. The specimen is a mere fragment of a sterile, 

leafy axis-apparently the tip of a young culmwith the distal branch complement in an early 

stage of development. A ticket attached to the 

specimen bears the name, Arundo gigantea, but no 

other annotation. 

The feature-combination characteristic of Arundinaria gigantea ssp. gigantea-as elaborated in the 

descriptive key-is based on the following specimens: Vegetative state: McClure (LU 15140) collected 18 April 1928 from a homogeneous, 

apparently natural stand growing in and adjacent 

to a swamp at the side of a small stream on the 

Old Shaker Farm, near Lebanon, Warren County, 

Ohio; and McClure 21321 (US) yielded by plants 

from the same source maintained under cultivation as MBG 2784. Flowering state: Winteringer 

7214 (US) “Low ground west of Sandusky, Alexander Co., Illinois. 7 July, 1951.” Fruiting state: 

Bain 117 (US) “River banks, Tennessee. 1892;” 

and Bush I98 (US) “Common in woods. Eagle 

Rock, Missouri. May 31, 1898.” 

Swallen 6717 (US 2078084-2078089)-i11ustrated 

in Hitchcock, 1935 [1951, fig. I], as Arundinaria 

giganten (Walter) Muhlenberg-and D. M. Moore, 

s.n. (US 2206426-2206430) are relatively complete 

specimens that appear at first sight to represent typical A. giganten ssp. gigantea. On closer examination, 

however, both of these collections show evidence 

of minor introgression of genetic influence from 

subspecies tecta. This is expressed inconspicuously 

in such tectoid features as a noticeable vinous 

tinting of the lemma and the palea, and the development, sooner or later, of negative geotropism 

in each rhizome, whereupon it turns upward apically to form a culm from its growing point. Strict 

application of the formula provided by the descriptive key identifies the plants represented by 

these two specimens as members of the clinal 

series embraced by A. gigantea ssp. macrosperma. 

They may be visualized, however, as occupying 

positions morphologically and genetically (and 

therefore taxonomically) very near to A. gigantea 

ssp. gigantea. 

2. Arundinaria gigantea ssp. tecta (Walter) 

McClure, new status 

FIGURES 12-14, 15~, 16c 

Arundinaria tecta (Walter) Muhlenberg, 1813:14: 1817:191. 

Arundo tecta Walter, 1788:81. 

FIGURE 7.-Arundinaria gigantea (Walter) Muhlenberg ssp. 

gigantea. A, Basal section of mature culm with section of 

rhizome attached, x 0.6; B, middle section of mature culm 

at end of first season, x 0.6; c, base of midculm branch 

complement, x 2.4; D, apex of leaf sheath with petiole and 

base of leaf blade, x ca. 6; E, cross-section of rhizome, showing lack of air canals, x ca. 12. All drawings based on 

McClure Bamboo Garden specimen no. 2784 (US). 

28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

NEOTYPE.-Walter’s type has not been found. 

However, the living plant to which his brief characterization led me, has presented both the vegetative 

and the reproductive stages of its ontogeny, and 

many facets of its potential morphological diversity. 

The feature-combination characteristic of A. gigantea ssp. tecta-as elaborated in the descriptive keyis based on McClure 22000 (US), a series of specimens yielded by plants in a natural stand growing 

in and adjacent to a swamp lying between Stony 

Run Creek and the Pennsylvania Railroad, near 

Friendship International Airport, Anne Arundel 

County, Maryland, and by plants from the same 

source maintained under cultivation, as MBG 2762. 

They were collected over a period of years both 

at the original site, and from plants taken from 

the same source and maintained under cultivation, 

since 1955. ‘This series of specimens constitutes the 

exclusive documentation of the characterization of 

subspecies tecta presented in the descriptive key. 

Plants growing in the wooded swamp reach a 

maximum height of about 2.5 m. These have 

remained in a sterile (vegetative) state ever since 

my observations were initiated in 1952. Other sterile 

plants of the same taxon growing in upland clay 

soil on the opposite side of the railroad (until 

they were destroyed by earth fill) reached a maximum height of about one meter. Plants growing 

along the edge of the swamp push their rhizomes 

into the original gravel ballast along the railroad, 

where they produce culms 0.5 to 1.0 m tall. Here 

and there among these latter, little patches of 

culms in a depauperate condition may be found in 

flower every year. However, I have never found 

fruits produced in that situation. The reproductive 

organs in every floret are always more or less completely destroyed by the larvae of insects that pass 

this stage of their life cycle within the anthecia. 

Two plants among those maintained under cultivation in my garden (about 20 miles from the 

natural stand) flowered and fruited freely. The 

production of fruits by these cultivated plants is 

a circumstance attributed to the absence of the parasitic insect at the latter site. In all observed cases 

of flowering, whether in wild plants or in those 

grown under cultivation, the culms that flowered 

died in the same season, along with the rhizomes 

that remained attached to them. 

3. Awndinaria gigantea ssp. macrosperma 

(Michau) McClure, new status 

FIGURE 16a 

Arundinaria macrosperma Michaux, 1803, I:74 [as to type]. 

TYPE (elig. Hitchcock, 1908: 156).- (P) “Herbier 

de 1’Amerique Septentrionale d’Andre Michaux.” 

“Gramen altissimum ramosum 5 [Michaux’s sym. 

bol for undershrub]. Hab. a Virginia ad Flondam 

et in occidentalibus [a phrase is here crossed out 

and illegible] juxta fluvios ab Illinoensibus ad 

ostium Misissipi -f [Michaux’s symbol for perennial plant.]” The specimen is a leafy flowering 

branch with leaf blades and florets detached. 

ISOTYPE (nunc elig.).-A specimen that quite 

clearly represents a duplicate from the same Michaux collection as that which yielded the lectotype 

is preserved at the Paris Herbarium. A printed 

label on the sheet bears the following information: 

MUSEUM D’HISTOIRE NATUWLE DE PARIS 

herbier d’Antoine Laurent de Jussieu. 

Donneau Museum par les enfants d’Andrien 

de Jussieu en 1857. Catal. no. 2881. 

The label (made in Paris by the elder Jussieu) 

bears the following annotations: 

Arundo gigantea Walter 

M. Michaux le dit tres different de 1’Arundo. 

il a tout le port du Nastus de l’inde et paroit 

etre la meme plante, mais M. Michaux dit qu’il n’a 

que 3 etamines et pour I’ordinaire 3 styles ou 

peut-etre un style divisC profondement en 3. Caroline 

et Rives du Mississipi jusqu’a 1’Ohio. Donne par M. 

Michaux 1797. 

Arundinaria Mich. en Dew. 

Another specimen, preserved in the Richard 

Herbarium (P) and labeled “ex Herb. E. Drake” 

supplements slightly the fragmentary representation 

(and the diversity) of the feature-combinations that 

appear in the image of the taxon Awndinaria 

macrosperma Michaux provided by the lectotype 

and the isotype. The species label bears the inscripFIGURE 8.-Arundinaria gigantea (Walter) Muhlenberg ssp. 

gigantea. A, Midculm section of young culm, x 0.6; B, tip 

section of young culm, x 0.6; c, culm sheath from midculm 

node, with blade not reflexed, x 3; D, culm sheath from 

higher level, with blade reflexed, x 3; E, Walter’s type of 

Arundo gigantea (from British Museum photo), x ca. 0.4. 

All drawings based on McClure Bamboo Garden specimen 

no. 2784 (US). 
NUMBER 9 29 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tion “Arundinaria macrosperma [ Michaux] Caroline. m.” With the possible exception of the word 

“macrosperma”-which appears to have been inserted by a different hand-this inscription is in the 

handwriting of Michaux. The specimen gives the 

plant from which it came a recognizable relationship to subspecies tecta. 

A sterile specimen, McClure 21320 (US), and a 

fruiting specimen, McCZure 21664 (US), represent 

a plant taken from the wild near the Ogeechee 

River, about 12 miles south of Savannah, Georgia, 

in the vegetative state, by David Bisset. I received 

a division of it in 1942 and maintained it under 

cultivation (as MBG 2803) until August 1948, when 

it flowered, bore fruits and died. The inflorescences 

match those of the lectotype and isotype of Arundinaria macrosperma. The fruits of this plant are 

similar in shape to the fruits of the neotype of 

subspecies tecta but differ in their much larger 

average size and in the infrequent incidence of the 

unciform rudimentary style branch. 

McClure 22018 (US) represents a plant secured 

from the wild near Travelers Rest, South Carolina, 

through the good offices of Robert A. Young, and 

maintained under cultivation (as MBG 2782) from 

1948 to the present. The plant, which has remained 

steadily in the vegetative state, combines expressions of morphological features characteristic, respectively, of subspecies gigantea and subspecies 

tecta, and some that are intermediate. The rhizomes 

(as in ssp gigantea) lack air canals, but (as in ssp. 

tecta) turn up apically to form a culm. The culms 

(as in ssp. tecta) proliferate from subterranean 

buds to form either culms or rhizomes. The primary axis of midculm branch complements (as in 

ssp. gigantea) lacks a basal “neck” but the component members of the complement (as in ssp. 

tecta) are more elongate and not so strongly curved 

away from the culm. The leaf blades are, on the 

average, larger than those of either typical subspecies gigantea or typical subspecies tecta. 

The revised taxonomic dispositions embodied in 

the nomenclature proposed here for these three 

subspecific components requires a postscript to my 

paper (McClure, 1963a) entitled “A New Feature 

in Bamboo Rhizome Anatomy.” The plant therein 

called Arundinaria gigantea (Walter) Muhlenberg, 

whose rhizomes lack air canals (cf. Figure 7~), is 

herein classified (and described in the key) under 

the name Arundinaria gigantea ssp. gigantea. Its 

documenting number is incorrectly given (McClure, 

1963) as MBG 2792; however, the origin of the 

plant is correctly given there; its correct number is 

MBG 2784. The plant (MBG 2762) therein called 

Arundinaria tecta (Walter) Muhlenberg, whose 

rhizomes show continuous air canals (cf. Figure 

13~) is herein classified (and described in the key) 

as Arundinaria gigantea ssp. tecta. The unnamed 

specimens (Biltmore 1405, and Radford and Wood 

6879-A) cited therein and described as having air 

canals discontinuous in the rhizomes, are now 

recognized as falling within my circumscription of 

Arundinaria gigantea ssp. macrosperma. 

In the limited perspective provided by their 

fragmentary representation in extant specimens 

of the conventional sort, these three subspecies are 

usually not clearly differentiated from each other 

in herbaria, For the identification of complete 

plants or complete specimens, however, the descriptive key is believed to be adequate and reliable. 

As they appear in nature, and as they are characterized in the descriptive key (p. 25), plants of 

subspecies macrosperma probably comprise the 

numerically dominant element of Arundinaria gigantea, sensu lato. This subspecies is here interpreted (circumscribed) as embracing plants (all 

apparently of hybrid origin) that embody, respectively, diverse recombinations of either identical or 

intermediate expressions of the same contrasting 

features that (in characteristic combinations) 

clearly differentiate subspecies gigantea and subspecies tecta, the putative genetical parents. 

The union, under one specific name, of the 

morphologically and genotypically distinct but obviously (at least ostensibly) interbreeding members 

of sympatric populations has the sanction of precedent. In the present case it has other defenses as 

well. It affords a framework for facilitating the 

practical disposition (identification and filing) of 

the fragmentary specimens of this species that 

FIGURE 9.--Arundinaria gigantea (Walter) Muhlenberg ssp. 

gigantea. A, Young leafy sterile culm with section of rhizome 

attached, x 0.6; B, young leafless flowering culm, x 0.6; c, 

whole sheath on midculm node of young culm shoot, x 0.6; 

D, apex and blade of culm sheath from upper node of young 

culm, showing reflexed sheath blade, x ca. 1.8. All drawings 

based on D. M. Moore s.n., Arkansas, USA, 2 August 1947 

(US) ’ 

32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

predominate numerically in herbaria. Assigning 

subspecific status to the three components portrayed in the descriptive key provides a means of 

giving greater depth of focus to floristic and ecological field studies such as those carried out by West 

(1935). For example, West’s identifications and 

descriptions of the geographic areas within which 

the existence of “canebrakes” is recorded by him 

are extremely valuable. Apparently, however, no 

effective effort was made, either to document the 

observed “ecological forms” by means of annotated 

voucher specimens or to correlate their distribution 

with that of the nine different “habitats” (ecological complexes) and the eight different floristic associations (“forest types”) within which they were 

observed to occur. Hopefully, such ecological and 

floristic emphasis will also encourage collectors to 

cultivate an awarneess of the potential taxonomic 

importance of morphological and ontogenetic diversities that await documentation, and to exercise 

the discipline necessary to give due attention to 

them in preparing specimens, whether these are 

intended for identification only or for permanent 

preservation. 

West (1935:255) maintained that the Arundinaria gigantea and Arundinaria tecta of the manuals 

are indistinguishable. “It is believed that the two 

so-called species of the manuals are in reality 

ecological forms of the same species , . . when both 

types are planted side by side in the greenhouse 

or in the open, they soon become indistinguishable.” 

I have had under observation for at least ten 

years, both in the wild and in cultivation under 

essentially identical conditions, plants I recognize 

as representing Arundinaria gigantea, Arundinaria 

tecta, and Arundinaria macrosperma, respectively. 

They have retained their distinguishing features as 

described in the key and as documented by voucher 

specimens cited here under the corresponding subspecies. 

Plants from different natural stands of either 

subspecies gigantea or subspecies tecta rarely match 

each other in all details of their ontogeny and their 

gross morphology. The persistence of observed 

minor differences between a number of variants of 

both subspecies gigantea and subspecies tecta when 

the plants were grown for a number of years under 

essentially identical conditions appears as evidence 

that the genotypic heterogeneity that characterizes 

hybrid populations may embrace the whole content 

of this polymorphic species-lightly so in the most 

divergent taxa (ssp. gigantea and ssp tecta) and 

more profoundly so in the diverse components of 

subspecies macrosperma that form a cline between 

the first two subspecies. 

Observations Concerning the 

Taxonomic Utility of Certain Structures 

and Their Contrasting Features 

I find that certain features that have been used 

by others to differentiate taxa identified, respectively, as Arundinaria gigantea and A. tecta are 

taxonomically undependable at both the species 

level and the subspecies level. 

MAXIMUM CULM HEIGHT.-The impression given 

by many local floras and incidental treatises is that 

plants of two recognizable bamboo taxa native to the 

United States may be distinguished from each other 

by the maximum height attained by their culms. 

Maximum culm heights given for plants referred 

to in the literature-either as gigantea or as macrosperma-range from 7.6 to 12 meters (Nuttall, 1837: 

149; Mitford, 1896:165; Hitchcock, 1936: 19 [1951: 

271) and those given for plants-referred to either 

as tecta or as macrosperma range up to 4.6 meters 

(Nuttall, 1818, I:39; Mohr, 1901: 103). However, 

I have not found published culm heights above 5.5 

meters documented in any case by reference to 

field notes accompanied by voucher specimens. 

Among the field notes accompanying specimens of 

more than 300 collections examined personally, I 

found less than a dozen that bear an indication of 

the height of the culms of the plant from which 

the specimen was taken. And in no case have I 

found field notes recording a maximum height 

greater than 5.5 meters, regardless of how the 

specimen was identified. 

The existence today of canebrakes containing 

culms at least 7.6 meters tall is not in doubt. During 

FIGURE lO.--drundinaria gigantea (Walter) Muhlenberg ssp. 

gigantea. A, Leafy culm tip, x 0.6; B, branch complement 

from node high on culm, with leafy and leafless flowering 

branches (with many spikelets shattered), x 0.6; c, base of 

midculm branch complement, x 2.4; D, apex of leaf sheath 

with petiole and base of leaf blade, x ca. 4.8. All drawings 

based on D. M. Moore s.n., Arkansas, USA, 2 August 1947 

(US). 

34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

the second world war I saw in a hardware store 

in Savannah, Georgia, 7.6-meter-long culms of a 

native bamboo on sale as fishing poles. I also have 

a photograph taken in 1905 or 1906 which shows 

a stand of native bamboo in Louisiana, the culms 

of which are estimated (on the scale of a man on 

horseback included in the picture) to be at least 

7.6 meters tall. In the absence, however, of documentation by means of adequate voucher specimens, it is impossible to assess the taxonomic 

pertinence of these published and unpublished 

observations. In the following paragraph C. A. 

Brown (1929:317) discusses what he considers to 

be the ecological significance of differences in culm 

size observed in different stands-all of which he 

includes in a comprehensive “Arundinaria tecta.” 

One might inquire as to the reason for the difference in 

size between the small woody canes the size of a lead pencil 

and only a few feet [ca. lm] tall and the large woody canes 

one to one and a half inches [2.5-3.8 cm] in diameter and 

up to thirty feet [9m] tall. It has been noticed that there 

are many patches of this small type. The only explanation 

that can be offered is that there must be some difference in 

soil or moisture conditions which hinders the growth of 

these plants. 

West(1935:255 and fig. 1) follows Brown in recognizing only one genotype, in adopting for it the 

name Arundinaria tecta (in a comprehensive 

sense), and in suggesting that plants producing 

tall culms owe their superior stature to a phenotypic response to favorable environmental conditions, especially those conditions that are related 

to edaphic factors. It appears that plants with very 

tall culms (illustrated by Harper 1928, fig. 18) have 

been reported only from the Gulf States. The answer to the question “Does culm size have any 

taxonomic significance?” remains to be discovered 

by means of additional field studies, documented 

by comprehensive, fully annotated specimens. 

There is no significant difference between subspecies gigantea and subspecies tech with respect to 

the observed maximum heights of culms found in 

stands of plants of these two taxa which are growing under similar ecological conditions at the northern limits of their currently known distribution. 

I have had these stands under intermittent observation for 40 years and 16 years, respectively. 

In the light of morphological and ontogenetic evidence that hybridization has taken place between 

plants representing these two taxa, it seems possible-even probable-that at least some of the plants 

with very tall culms may embody what is commonly known as hybrid vigor. 

APPENDAGES OF CULM SHEATHS AND LEAF 

SHEATHS.-The morphological expression of the appendages-ligules (inner and outer), auricles, oral 

setae, and blades-apical to culm sheath and leaf 

sheaths is often more or less variable even as betlveen specimens taken from the same plant. This 

variability is due, in large part, to ontogenetic factors related to the age of the whole plant, the age 

range as between young and old culms in the same 

plant, and the point of origin within the plant and 

on the culm from which the specimen was taken 

(see hlcclure, 1966b:6, par. 1). 

dence, patterns of distribution, and persistence of 

trichomes in the bamboos are subject to influence 

by so many (generally unrecorded) variables that 

it is often difficult to find consistent patterns of 

vesture in a given structure, as represented by preserved specimens. Pertinent variables that are generally ignored during the collection of bamboo 

specimens are age (or stature) of the plant; environment and relative vigor of the plant; position 

on the culm and relative age of the culm from 

which a given specimen is taken. Weathering, spontaneous abscission, or abrasion, may erase trichomes 

that were originally present on some structures. 

cently (Hitchcock, 1935:29, key) supposedly distinct differences in the loci of insertion of 

inflorescences was commonly offered as a means 

for the differentiation of two elements of the New 

World component of Arundinariu recognized (but 

inadequately characterized) under the names A. giVESTURE OF VEGETATIVE ~TRumuREs.-The inciLOCI OF INSERTION OF INFLORESCENCES.-Until reFIGURE ll .-Arundinaria gigantea (Walter) Muhlenberg ssp. 

gigan tea. A, Spikelet with pedicel and inflorescence branch 

subtended by a small bract, x 1.8; B, base of spikelet showing transitional glumes I and 11 and sterile lemma, x 1.8; c, 

transitional glume I (normal) , x 3.6; D, transitional glume 

11 (variant), x 3.6; E, transitional glume 1x1 (sterile lemma), 

variant, x 3.6; F, floret, x 3.6; G, fertile lemma (normal), x 

3.6; H, fertile lemma with stipule-like appendages, x 3.6; I, 

stipule-like appendage, X 10.8; J, palea, x 3.6; K, lodicule 

complement showing dorsal surface, x 7.2; L, lodicule complement from another flower showing adaxial surface, x 7.2; 

kr, gynoecium and androecium, x 7.2; N, fruit, hilum side, 

x 7.2; 0, fruit, embryo side, x 7.2. All drawings based on 

D. M. Moore, s.n., Arkansas, USA, 2 August 1947 (US), 
NUMBER 9 35 

J 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

gantea (or A. macrosperma) and A. tecta. It appears that C. A. Brown (1929:316) was the first to 

focus attention upon the confusion that may result 

from reliance upon this feature as a source of contrasting characters to support a traditional pattern 

of taxonomic segregation. I have observed that this 

is one of the features whose expression-in Arundinaria as well as in many other genera-may 

change progressively during successive stages of the 

flowering of a plant or may differ as between very 

young and very old parts of the plant. Its reliability for the differentiation of taxa of any category 

must be tested by means of exhaustive and sustained field studies. 

TRANSITIONAL GLUMES INCIDENT AT THE BASE OF 

EACH SPIKELET.-variable also are the transitional 

glumes at the base of the spikelets. Gilly (1943: 

301, figs. 2-A, 2-M) illustrated, as characteristic (typical) of his Atlantic type and his Mississippi type, 

empty glumes that are distinguished, respectively, 

by size, shape, vesture, marginal ciliation, and venation. These differences, which really exist, are ostensibly of genotypic origin. However, I have 

found the empty glumes to be not consistently uniform, in their number or in their morphological 

expression, either in duplicates from the collections 

cited by Gilly, or in the plants I have chosen as 

typical, respectively, of the subspecies I distinguish 

under Arundinaria gigantea, sensu lato. 

LoDIcuLEs.-Michaux (1803, I:74) gave the genus 

Arundinaria but 2 lodicules, which he designated 

as “appendices.” He, and those who subsequently 

have quoted this number, apparently overlooked 

the posterior member of the trio that I have always 

found to be present in members of this genus. On 

account of its slightly more distal insertion, however, the posterior lodicule often remains attached 

to the palea and hidden inside of it, when a floret 

is dissected. Members of the lodicule complement 

in Arundinaria (and in other genera as well) are 

subject to such a diversity of unpredictable variabilities that it is often very difficult to discover 

features that embody expressions sufficiently reliable for descriptive purposes. 

THE STYLAR AxIs.-In the neotype of Arundinaria gigantea ssp. tecta, the fusion of two of the 

three stylar branches sometimes extends upward to 

a point noticeably beyond the level at whidh the 

first one became free. The appearance produced 

by this deviation from the usual manner in which 

branches of the style are disposed may have lent 

deceptive support to a description that characterizes the genus Arundinaria as embracing taxa having two styles (or stylar branches) and taxa having 

three styles (or stylar branches). Published 139 

years ago by Nees (1834:478) this incorrect characterization of Arundinaria is still widely copied in 

the current literature. The error appears to have 

had its origin in Nees’ allocation to the genus 

Arundinaria (where the number of stylar branches 

or stigmas is always 3) of several species of other 

genera (see below the list entitled New World 

species excluded from the genus Arundinaria as 

herein defined) where the number of stylar 

branches or stigmas is always 2. 

The array of morphological, anatomical, and 

ontogenetic features whose taxonomic utility I have 

explored may not complete the list of potentially 

useful ones. It remains to be seen whether comprehensive studies in anatomy and other disciplines, 

correlated with intensive field studies, will improve 

the present taxonomic perspectives on this heterogeneous complex of closely related but distinguishable entities between which biological (genetic) 

isolation apparently has not yet been fully established. 

Reeder (1957, fig. 29) illustrated, in cross-section 

and sagittal section, the embryo of a fruit from a 

plant of bamboo native in Ann Arundel County, 

Maryland, documented under MBG 2762, and identified as Arundinaria tecta. Could (1960:873) reported for a plant from a stand of bamboo native 

in Walker County, Texas, identified as Arundinaria 

gigan.tea, a chromosome count of “n equals 24.” 

Checklist of New World Species Excluded 

from the Genus Arundinaria 

(as herein defined) 

The apparently general acceptance of the conFIGURE 12.-Arundinaria gigantea ssp. tecta (Walter) McClure. 

A, Young culm shoot with segment of rhizome, x 0.6; B, 

lower part of sterile leafy culm in the second season of its 

growth, with solitary leafy branches fully developed, x 0.6; 

c, midculm section of culm in its second season of growth, 

showing persistent sheaths and developing branch complement, x 0.6; D, base of unrestricted monoclade midculm 

branch complement, x 1.2; E, seedling with anthecium still 

attached, x 1.2. All drawings from McClure Bamboo Garden 

specimen no. 2762 (US). 
NUMBER 9 37 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ventional circumscriptions of the genus Arundinaria that were published after that of Nees (1834) 

has postponed the recognition that, of the approximately fifty distinct neotropical taxa that have at 

one time or another been assigned to this genus, 

not one falls within its natural boundaries. 

1. Arundinaria acuminata. See Yushania acumi2. Arundinaria amplissima. See Aulonemia am3. Arundinaria aristulata. See Aulonemia aristu4. Arundinaria ? attenuata Doell; species sedis 

5. Arundinaria burchellii. See Colanthelia bur6. Arundinaria cannavieira. See Apoclada canna7. Arundinaria capillifolia is Arthrostylidium 

8. Arundinaria cubensis is Arthrostylidium cu9. Arundinaria ? decalvata Doell; species sedis 

nata. 

p lissima. 

lata. 

mihi incertae etiam nunc manet. 

chellii. 

vieira. 

capillifolium Grisebach. 

bense Ruprecht. 

mihi incertae etiam nunc manet. 

10. Arundinaria deflexa. See Aulonemia deflexa. 

11. Arundinaria distans. See Colanthelia distans. 

12. Arundinaria effusa. See Aulonemia effusa. 

13. Arundinaria excelsa. See Arthrosty lidium venezuelae. 

14. Arundinaria fimbriata is Arthrostylidium fimbriatum Grisebach. 

15. Arundinaria Pabellata; species sedis mihi incertae etiam nunc manet. 

The name Arundinaria flabellata (Fournierj McClure (in Maguire, Wurdack, .et al., 1964:182) is 

based on Guadua ? Pabellata Fournier 1881:131. 

The plant is known only by nonflowering specimens (Liebmann 132 [“131”]) really too fragmentary for confident generic allocation at present. 

Since it clearly does not belong in Arundinaria (as 

the genus is defined herein) the taxonomic disposition of this plant awaits the realization of its thorough study and documentation, preferably at the 

type-locali ty. 

16. Arundinaria glaziovii. See Aulonemia glazi17. Arundinaria glaziovii var. macroblephara. See 

ovii. 

A u 1 on em ia ramosissima. 

18. Arundinaria goyazensis. See Aulonemia goya19. Arundinaria haenkei. See Aulonemia haenkei. 

20. Arundinaria haitiensis is Arthrostylidium haitiensis (Pilger) Hitchcock and Chase. 

21. Arundinaria herzogiana. See Aulonemia herzogiana. 

22. Arundinarict hirtula. See Aulonemia hirtula. 

23. Arundinaria humillima. See Aulonemia hu24. Arundinaria leptophylla is Chusquea lepto25. Arundinaria longiflora is Arthrostylidium 

26. Arundinaria longifolia. See Bambusa (Gua27. Arundinaria macrostachya. See Colanthelia 

28. Arundinaria maculata. See Aulonemia parui29. Arundinaria microclada is Chusquea abieti30. Arundinaria mirabilis is Glaziophyton mira31. Arundinaria mucronata. See Aulonemia aris32. Arundinaria multiflora is synonym of Arundi33. Arundinaria multispicata is Arthrostylidium 

34. Arundinaria obtusata is Arthrostylidium obtu35. Arundinaria parviflora. See Rhipidocladum 

36. Arundinaria patula. See Aulonemia patula. 

37. Arundinaria pinifolia is Chusquea pinifolia 

38. Arundinaria pittieri. See Rhipidocladum pit39. Arundinaria prestoei. See Rhipidocladum preszensis. 

millima. 

phylla Nees. 

longiflorum Munro. 

dua) longifolia. 

macrostachya. 

flora. 

folia Grisebach. 

bile Franchet. 

t u la ta, 

naria trianae, q. v. infra. 

multispicatum Pilger. 

satum Pilger. 

parviporum. 

(Nees) Nees. 

tieri. 

toei. 

FIGURE 13.-Arundinaria gigantea ssp. tecta (Walter) McClure. 

A, rhizome turned upward apically and terminating in a 

simple inflorescence, x 0.6; B, leafy and leafless flowering 

branches from tip of rhizome, x 0.6; c, apex of sheath and 

reflexed blade from midculm node, x 3; D, apex of leaf 

sheath showing petiole and base of blade, x 6; E, crosssection of rhizome, showing one of the peripheral air canals, 

x ca. 12. All drawings based on McClure Bamboo Garden 

specimen no. 2762 (US). 
NUMBER 9 39 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

40. 

41. 

42. 

43. 

44. 

45. 

46. 

47. 

48. 

49. 

50. 

51. 

52. 

53. 

54. 

55. 

Arundinaria pubescens is Arthrostylidium puArundinaria queko. See Aulonemia queko. 

Arundinaria radiata. See Aulonemia radiata. 

Arundinaria ramosissima. See Aulonemia raArundinaria rhizantha. See Colanthelia rhizanArundinaria schomburgkii is Arthrostylidium 

Arundinaria setifera. See Aulonemia haenkei. 

Arundinaria setigera. See Aulonemia setigera. 

Arundinaria simpliciuscula. See ArthrostylidArundinaria sodiroana. See Aulonemia sodiArundinaria standleyi. See Arthrostylidium 

Arundinaria trianae. See Aulonemia trianae. 

Arundinaria ulei. See Aulonemia ulei. 

Arundinaria urbanii. See Arthrostylidium urArundinaria verticillata. See Rhipidocladum 

Arundinaria viscosa. See Aulonemia viscosa. 

bescens Ruprecht. 

m osiss ima . 

tha. 

schomburgkii (Bennett) Munro. 

ium simpliciusculum. 

roana. 

venezuelae. 

banii. 

verticillatum. 

Athroostachys Bentham 

FIGURES 19, 20 

Athroostachys Bentham, in Bentham and Hooker, 1883: 1208,- 

Hackel, in Engler and Prantl, 1887:93.-McClure, 1957:200. 

Achroostachys [sic] Bentham, 1881: 134 [nomen nudum].- 

McClure, 1957:199. 

Plants unicespitose, unarmed. Rhizomes pachymorph. Culms clambering (teste Gardner), the internodes terete and fistular. Primary branch buds 

at culm nodes apparently solitary (not seen). Initial branch primordia at each midculm node more 

than one. Branch complement at a midculm node 

unrestricted pleioclade, the initial component axes 

(three in Glaziou 13325; illustrated in Figure 19~) 

each arising from an apparently independent initial 

primordium, all inserted at approximately the same 

level, subequal or the middle one noticeably dominant. Leaf blades with transverse veinlets not 

manifest externally. 

Inflorescences semelauctant, solitary, contracted, 

short-peduncled panicles of capitate or subcapitate form, each inserted at or near the apex of a 

usually leafy twig, and subtended by a laminiferous 

sheath; the primary branches of the percurrent 

rachis inserted distichously in obscurely secund 

orientation, very short, not prophyllate, either unbranched or bearing one or two subequal short 

branches; the lowermost primary branches each 

subtended by a bract, the uppermost one or more 

not so subtended, the branches of higher orders 

mostly so subtended; the bracts varying from the 

liguliform caudate unawned foliar organ up to 3 

cm long that subtends the lowermost primary 

branch, down to short, awned scales 0.5 mm long 

that subtend branches of the higher order. Transitional glumes 2, empty, approximate. Spikelets 

pedicellate, each containing a single perfect flower, 

the rachilla potentially disarticulating just below 

the locus of insertion of the fertile lemma, distally 

prolonged behind the palea and bearing a depauperate sterile anthecium. Lemma of perfect florets 

fully embracing its palea only basally at maturity. 

Palea gaping antically, narrowly sulcate and 2- 

keeled dorsally. Lodicules 3, the anterior 2 asymmetrical and paired, the posterior one symmetrical 

and much smaller. Stamens 3, the filaments filiform, free. Stigmas 2. Mature fruit unknown. 

ETY MoLocY .-The name A throostach ys (derived 

from the Greek, athroos, crowded, and stachys, 

spike) ostensibly alludes to the condensed paniculate branching of the capitate inflorescence. 

TYPE-sPECIES.-AthroostachyS capitata (Hooker) 

Bentham (in Bentham and Hooker, 1883:1208). 

Merostachys capitata Hooker (1840:pl. 273-274); 

Munro (1868:50); Doell (in Martius, 1880:216); 

Ekman (1913:64). Chusquea fimbriata Steudel 

(1854:338). Cotypes of Merostachys capitata: 

Gardner 136 (BM); Tweedie 1324 (BM); type of 

Chusquea fimbriata: Riedel s.n. (P). 

As a result of my own experience, I can understand the difficulty evidently encountered by Bentham in determining the categories to which to 

FIGURE 14.-Arundinaria gigantea ssp. tecta (Walter) McClure. 

A, Inflorescence of racemose branching, x 0.6; B, infloresence 

of paniculate branching, x 0.6; c, immature spikelet, x 1.8; 

D, abnormally elongate spikelet, X 1.5; E, floret, X 7.2; F, 

fertile lemma, x 7.2; G, palea, x 7.2; H, lodicule complement, x 15; I, stamen, x 7.2; J, gynoecium x 7.2; K, fruit, 

hilum side, x 7.2; L, fruit, embryo side, x 7.2; M, fruit, 

embryo side, the embryotegium inconspicuous, X 7.2; N, 

fruit, longitudinal section, x 7.2. All drawings based on 

McClure Bamboo Garden specimen no. 2762 (US). 
NUMBER 9 41 

L 

U 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

assign the successive sections of each ultimate 

branch of the inflorescence and the several sheathing structures that precede the fertile lemma in this 

bamboo. Having identified as a first empty glume, 

the bract that subtends the pedicel of a spikelet, 

and having equated the pedicel, so subtended, to 

an elongated rachilla, Bentham (in Bentham and 

Hooker, 1883: 1209) wrote as follows: “spiculis . . . revera sessilibus, sed ob rachillam supra 

glumam infimam productam pedicellatae apparent.” That is to say “the spikelets are really 

sessile, but they appear to be pedicellate on account 

of the elongation of the rachilla beyond the first 

glume.” 

Doell encountered similar difficulties. In his description (in Martius, 1880:216) of the type-species 

of the genus Athroostachys (under Merostachys 

capitata Hooker) Doell not only referred to the 

spikelets as sessile, he gave them two rudimentary 

empty glumes and two sterile lemmas! 

Munro (1868:50) recognized the spikelets of 

Merostachys capitata as pedicellate, but he saw the 

two empty glumes as sterile lemmas-by convention 

called, at that time, “flosculi steriles.” 

In his original description of Merostachys capitutu, Hooker confessed uncertainty as to its generic 

affinity, but his recorded observations do not contain the errors noted above. 

Under Merostachys capitata, but without designating types, Doell (in Martius, 1880:217) described 

two variants of Athroostachys capitata, distinguishing them on the basis of width of leaf blades: (Y 

latifolia (presumably representing the typical form 

of the species) and @ angustijolia. Munro (1868: 

50) intimates that the type of Chusquea frnbriata 

Steudel represents a plant with the character of the 

latter variant. He adds, however, that it is not 

clear whether the sterile, leafy branches of the 

Riedel specimen belong with the flowering branches 

or not. 

RELATIONS HIPS.-^ do not find clear evidence of 

a close affinity between Athroostachys and any other 

known genus. The strong resemblance of the very 

prominent radiate oral setae, borne on its leaf 

sheaths (Figure I~D), to those of many species of 

Merostachys may have exerted an influence in 

favor of the initial allocation of the type-species 

of Athroostachys to the genus Merostachys. This 

superficial resemblance, however, plus noticeable 

similarities in a few technical features of the reproductive apparatus, does not add up to a clear 

indication of close phylogenetic affinity between the 

type-species of Athroostachys and members of the 

genus Merostachys. 

Drsm1suTIoN.-Recorded only from Brazil, the 

single known species of Athroostachys has been collected in the immediate vicinity of the city of Rio 

de Janeiro; near Jacarehy in the state of Parani; 

and near Villa Maria in the state of Mato Grosso. 

Atractantha McClure, new genus 

FIGURES 21-23 

The defects of conventional procedures prevalently followed in the documentation of bamboos 

for taxonomic purposes are impressively demonstarted by the extreme incompleteness of the first 

two (and the only available) collections representing this very distinct, hitherto undescribed genus. 

Representation, in herbaria, of the individual component taxa apparently has not been improved in 

comprehensiveness during the thirty years that 

have passed since the first specimen was brought 

to light! 

Plantarum habitus non adhuc relatus. Rhizomata non adhuc visa (?pachymorpha). Culmi in 

speciminibus suppetenibus haud satis representati; 

internodiis teretibus, vel inanibus vel efistulosis. 

Ramorum complementarum (in speciminibus suppetentibus) axes componentes primum proditi 

plures, ad circa libram inserti, unusquisque ex primordio distinct0 enascens (gemmis intactis primordios primos includentibus non adhuc visis); uno 

alios plus minusve valde dominanti; ramorum comFIGURE 15.-Young culms and roots of Arundinaria gigantea 

(Walter) Muhlenberg ssp. gigantea (A) and A. gigantea ssp. 

tecta (Walter) McClure (B) compared. Aa, Base of culm and 

section of rhizome, with roots, x 0.3; Ab, young culm, full 

height at age 3 months, showing branches already well 

developed, x 0.15; AC, cross-section of root from rhizome, 

to show air canals, x 15; Ad, cross-section of root from culm 

base, to show air canals, x 15; Ba, base of culm and section 

of rhizome, with roots, x 0.6; Bb, young culm, full height, 

with branches not yet developed, x 0.3; BC, cross-section of 

root from rhizome, to show air canals, x 30; Bd, cross-section 

of root from culm base, to show air canals, x 30. Drawings 

Aa-Ad based on McClure Bamboo Garden specimen no. 2784 

(US), Ba-Bd on McCture Bamboo Garden specimen no. 

2762 (US). 
NUMBER 9 43 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

plemento ubi plene evoluto composito-pleioclado. 

Foliorum lamina venulis transversis extus vel infirme vel haud manifestis. 

Inflorescentiae iterauctantes, vel capitatae vel 

diff usae, earumdem ramorum insertione aut disticha aut sympodiali, rhachidum ramificationibus 

omnibus et bracteatis et prophyllatis, omnis rhachidis segment0 terminali pro pedicel10 spiculae consistenti. Glumae transitionales ad basin spicularum 

vulgo praeditae hic nullae. Spiculae deciduae, infra et proxime locum lemmatis insertionis disarticulantes, unaequaeque flosculam perfectam 

solitariam habentes, rhachilla post paleam prolongata in setam gracilem anthecio rudimentario 

terminatam. Lemma fertile maturitate saltem paleam suam basi tantum circumplectans. Palea apice 

bifurcata, dorso sulco angustissimo instructa, antice leviter imbricata. Lodiculae 3, anteriores 2 

sese aequales, plus minusve valde asymmetricae 

atque geminatae, posteriore symmetrica et anterioribus aliquanto minore. Stamina 3, filamentis filiformibus liberisque. Stigmata typice 2 (atypice et 

rarissime 3). Fructus maturus adhuc non inventus. 

Plants of as yet unrecorded habit. Rhizomes not 

yet seen, ?pachymorph.l Culms not adequately represented in the available specimens; the internodes 

terete, either fistular or efistular. Branch complements (at least those seen) containing several initial 

components, each arising from a distinct initial 

primordium.2 The branch complement at midculm 

nodes unrestricted pleioclade when fully developed, 

one of the primary components more or less 

strongly dominating the others, all inserted in close 

order at approximately the same level. Leaf blades 

with transverse veinlets weakly to not at all manifest, externally. 

‘The pachymorph character of the rhizomes (which are 

neither represented in the available specimens nor described 

by the collector) may be surmised from the strong resemblance of the inflated base of the primary axis of a branch 

complement (exemplified in the isotype specimen of A. radiata) to a pachymorph rhizome (see McClure 1966b358, 

last paragraph). 

‘It has not been psible to ascertain with certainty from 

the available specimens whether the independent primary 

primordia incident at each culm node are originally enclosed 

in a common prophyllum as one primary bud, or whether 

each independent initial primordium corresponds to an independent primary bud. 

FIGURE 16.-Arundinaria gigantea, sensu lato. Semidiagrammatic sketches showing the extremes and intermediate expressions of the principal features that characterize the 

recognized components of the Arundinaria gigantea complex, including subspecies gigantea (A) , subspecies macrosperma (B), and subspecies tecta (c), as to growth habit 

(a), rhizome in cross-section (b), spikelet (c). fertile lemma 

(d) , and transitional glumes (e) , Subspecies gigantea is 

characterized by the combination of expressions Aa, Ab, AC, 

Ad, and Ae. Subspecies tecta is characterized by the combination of expressions ca, cb, cd, and ce of the same 

features. Nothomorphs of the presumed hybrid swarms (of 

which subspecies gigantea and subspecies tecta are the 

putative parents or grandparents) herein given the status 

subspecies macrosperma, show diverse combinations OE the 

same contrasting expressions of these features, or of intermediate expressions of them. Some of these intermediate 

expressions are illustrated by Ba,- sd, and ~e (transitional 

glumes) in the intermediate expression shown in the lectotype of Arundinaria macrosperma Michaux. The transitional 

glumes often vary more or less widely in size and shape 

within any given specimen, whether of otherwise typical subspecies gigantea or typical subspecies tecta, or of the polymorphic subspecies macrosperma. Moreover, in any given 

specimen of any one of the subspecies the lowermost transitional glume may be lacking entirely, or it may be inserted 

at some distance below the second one. A third transitional 

glume sometimes appears in the guise of a “sterile lemma.” 

Inflorescences iterauctant, of either capitate or 

diffuse form, their branching pattern either distichous or sympodial, each axis the bracteate and 

prophyllate rachis of a pseudospikelet, the terminal 

segment of each rachis serving as the pedicel of an 

abscissile spikelet. Transitional glumes at the base 

of each spikelet none.3 Spikelets each made pedicellate by the segment of the rachis it terminates, 

each spikelet typically containing but a single perfect flower, and disarticulating promptly at maturity just below the locus of insertion of the 

lemma, the rachilla prolonged behind the palea in 

a bristle-like structure bearing a minute rudiment 

of a sterile anthecium. Lemma in functional florets 

embracing its palea only basally at maturity. Palea 

apically bifurcate, lightly imbricate antically, dorsally sulcate, the sulcus very narrow or subtubular. 

Lodicules 3, the anterior 2 more or less strongly 

asymmetrical, paired and equal, the posterior one 

symmetrical and somewhat smaller (at least either 

shorter or narrower). Stamens 3, filaments filiform, 

The ostensible (approximate) position of the transitional 

glumes is here (and in Elytrostachys, q.v.) occupied by what 

I now classify as bracts. 

46 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

ss 

Arundinaria gigantea, sensu Lato 

9iyantea ssp. tecta 

No thomorph s 

? (ssp. macrosperma) ~~~ ? 

FIGURE 17.-Arundinaria gigantea, sensu lato. Diagrammatic 

representation of the putative phylogenetic relationships of 

the principal New World components of the genus Arundinaria. The arrows suggest putative contemporary potential 

for further introgression or hybridization. The question 

marks and the broken lines raise the question, “Are there 

any existing stands of ‘pure’ gigantea and ‘pure’ tecta?” 

Arundinaria 

in rhizomes \ Air canals continuous I canals not present / in rhizomes 

t 

Air canals discontinuous 

in rhizomes 

FIGURE 18.-Arundinaria gigantea, sensu lato. Schematic portrayal of some of the quantitative and qualitative relationships 

visualized as existing between the strongly divergent populations herein categorized as subspecies gigantea and subspecies tecta, respectively, and the populations of intermediates 

(putative nothomorphs) herein given taxonomic status as 

subspecies macrosperma, based on Arundinaria macrosperma 

Michaux (as to nomenclatural lectotype) . The apparent 

absence of either a clear morphological discontinuity, or an 

absolute barrier to the exchange of genes, as between the 

populations comprising these three groups of plants, they 

are herein interpreted as constituting a polymorphic species, 

and given the formal, comprehensive name, Arundinaria gigantea, sensu lato. 

free. Stigmas typically 2 (atypically and very rarely 

3 in A. radiata, q. v,), Mature fruit not yet found. 

ETYMoLoGY.-The name A tractantha (f.) Greek 

atraktos, spindle, and anthos, flower, alludes to the 

spindle-like form of the anthecia. 

TYPE-SPECIES.-A tractantha radiata McClure. 

RELATIoNSHIPS.-The obvious natural affinity of 

the known bamboos of the genus Atractantha is 

toward bamboos of the genus Elytrostachys, with 

which they share the following features: inflorescences are pseudospikelets of a peculiar form characterized by rachises with long terminal segments, 

each of which serves as the pedicel of an abscissile 

spikelet; the absence of the transitional glumes 

commonly found at the base of each spikelet, and 

the presence, in their stead, of a bract inserted on 

each of the two nodes that precede the elongated 

terminal segment of each rachis; each spikelet typically containing but a single perfect floret, which 

is followed by a rudiment of a sterile anthecium 

borne on the tip of a bristle-like prolongation of 

the rachilla. Insofar as currently available specimens inform us, bamboos of the genus Atractantha 

differ from bamboos of the genus Elytrostachys in 

the following respects: midculm branch complements containing several axes of primary order, 

each developed from one of several distinct initial 

branch primordia each with its own prophyllum; 

leaf sheaths with auricles and oral setae not only 

much less conspicuously developed but of different 

morphological configuration; empty bracts preceding the base of the terminal segment of each rachis 

none or one; the slender, spindle-like form of the 

anthecia; and the number of stamens in each flower 

limited to 3. While the recorded geographical 

range of bamboos of the genus Atractantha is limited to the state of Bahia, Brazil, the recorded 

FIGURE 19.-Athroostachys capitata (Hooker) Bentham. A, 

Basal portion of young plant in the vegetative state, showing 

two culms and the rhizomes to which they are terminal, 

x 0.G; B, midculm node and branch complement showing 

three primary branches of independent origin, derived from 

the primordial meristem enclosed within the prophyllum 

of a solitary primary bud, x 3; c, tip of a leafy flowering 

branch with its terminal inflorescence, X 0.6; D, apex of 

leaf sheath which subtends a flowering branch and the 

base of its blade, x 0.9; E, base of a flowering branch 

(schematic). Drawing A based on Glaziou 13325 (P), B, on 

Weddell 152 (P) , and c, D, E on Glaziou 20155 (F) , 
NUMBER 9 47 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

geographical range of bamboos of the genus Elytrostachys extends from Venezuela to Nicaragua. 

bution of the two known species of the genus Atractantha is limited to the state of Bahia, Brazil. It 

appears that nothing concerning the ecological features of their respective natural habitats was reDISTRIBUTIoN.-The recorded geographical distri- corded by the collectors of the only available 

specimens. 

Key to the Known Species of Atratantha 

la. Leaf blades up to 10.5 cm long and up to 1.0 cm broad; leaf sheaths hispid and hispidulous at first; internodes of leafy twigs hirsute; inflorescences diffuse, their ultimate 

branches curved; rachises of sympodial insertion, each bearing a bud at a single proximal node; bud-subtending bracts laminiferous ................................ 1. A. falcatu, new species 

lb. Leaf blades up to 19.5 cm long and up to 3.5 cm broad; leaf sheaths glabrous from the 

first; internodes of leafy twigs glabrous; inflorescences capitate, their ultimate branches 

straight; rachises of distichous insertion, each bearing a bud at each of"2 proximal 

nodes; bud subtending bracts without leaf blades .................... 2. A. radiafu, new species 

1. Atructantha fulcutu McClure, new species 

FIGURE 21 

Plantarum habitus non adhuc relatus. Culmi in 

specimine suppetenti neque satis representati neque 

descripti, itaque ignoti. Vaginae culmi et vaginae 

ramorum primariorum in specimene suppetenti deficientes. Vagina foliorum arcta, initio hispida hispidulaque demum subglabrescens; versus apicem 

tantum nervis interdum debiliter manifestis; auriculis vel subnullis vel sat evolutis glebosisque; setis 

oralibus paucis, e basi ad apicem sensim angustatis, 

omnino glabris vel basin versus sparsissime et antrorse scaberulis, fragilissimis, vel mox vel postea 

effractis; ligula interiore subnulla, truncata; ligula 

exteriore truncata, margine primo sparse et minutissime ciliolata denique glabrescenti integraque; 

petiolo vix 2 mm longo, pagina adaxiali hispidulo 

pagina abaxiali hirsuto; lamina usque 10.5 cm 

longa et 1.0 cm lata, lineari-lanceata, apice acuta, 

basi rotundata, basin versus utrinsecus sparse hirsuta, alibi utrinsecus primo antrorse hispidula, 

denique sensim glabrescenti; costa mediana et tribus paribus nervorum secundariorum pagina 

abaxiali sat prominulis; pagina adaxiali inter 

omnes nervos propinquos profunde et angustissime 

sulcata, prope marginem externam costa alata penitus instructa; venulis transversis extus haud manifestis. 

Inflorescentiae ramos ramulosque vel foliigeros 

vel aphyllos terminantes, ob rhachidibus ramificationibus insertionis sympodialis falcatae diffusaeque. Bracteae e quoque rhachidi proditae 

pleraeque 2, altera gemmam subtendenti altera 

vacua, approximatae, laminiferae, paucinerves, 

substantia indurata sed fragili, dorso sparsim puberulae. Spiculae usque 15 mm longae, maturitate 

cadentes; ubi unaquaque spicla delapsa relictum 

est segmentum terminale rhachidis a bractea persistenti fere omnino vestitum. Glumae transitionales nullae. Anthecia papyracea, glabra nitidaque, 

nervis extus vix vel haud manifestis. Lemma usque 

15 mm longum, lanceatum, attenuate acuminatum. 

Palea lemmate brevior, secus margines sulci angustissimi antrorse ciliata. Lodiculae diaphanae, 

angustae, margine apicem versus sparsim ciliolatae. 

Antherae apice emarginatae, basi profunde divisae, 

ubi desiccatae brunneae. Ovarium vel omnino 

glabrum vel apice tantum hispidulum. Stylus antrorse hispidulus. Fructus maturus adhuc non visus. 

FIGURE 20.-Athroostachys capitata (Hooker) Bentham. A, 

Diagram of the branching system of the inflorescence; B, 

a primary branch of the inflorescence, terminating in a 

spikelet, and bearing two secondary branches each terminating in a single somewhat immature spikelet, x ca. 3; c, 

a secondary branch subtended by a bract and bearing a 

solitary spikelet, x ca. 3; D, transitional glumes of a mature 

terminal spikelet, x ca. 3; E, hermaphrodite floret and (at 

left) depauperate (sterile) anthecium of a mature terminal 

spikelet, x ca. 3; F, component structures of a secondary 

branch of the inflorescence (schematic) ; G, floral parts in 

immature state (gynoecium not visible), x ca. 12-i H, mature 

lodicule complement (the narrow symmetrical one is the 

posterior member), x ca. 18; I, stamen, x ca. 12; J, gynoecium, unmodified, x ca. 12; K, gynoecium, immature 

fruit stage, x ca. 12. All drawings based on Glaziou 20155 

(F). 
NUMBER 9 49 

K 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Plants of unrecorded habit. Culms neither rep 

resented nor described in the available specimen. 

Culm sheaths and sheaths of the primary axes of 

branch complements lacking in the specimen. Leaf 

sheaths tight, hispid and hispidulous at first, at 

length subglabrous; nerves sometimes weakly discernible toward the apex only; auricles either subobsolete or somewhat developed and lumpy; oral 

setae few, tapered, glabrous throughout or sparsely 

and antrorsely scaberulous basally, very fragile, 

sooner or later broken off; the inner ligule truncate, not at all or barely exserted; the outer ligule 

truncate, sparsely ciliolate on the margin at first, 

at length glabrescent and entire; petiole scarcely 2 

mm long, hispidulous on the adaxial surface, and 

hirsute on the abaxial; the blade up to 10.5 cm 

long and 1.0 cm broad, linear-lanceolate, apically 

acute, basally rounded, sparsely hirsute on both 

surfaces at the base, elsewhere antrorsely hispidulous at first, then gradually glabrescent on both surfaces; the conventional midrib and three pairs of 

secondary nerves moderately prominent on the 

abaxial surface, narrowly and deeply sulcate between all adjacent nerves of the adaxial surface, 

with a winglike rib from base to tip near the outer 

margin and, at the same time, the midrib obsolete 

toward the apex on this surface; transverse veinlets 

not at all manifest on either surface. 

Inflorescences terminating leafy and subleafless 

branches of all orders; of diffuse, falcate branching, 

the rachis branches of sympodial insertion. Bracts 

at the base of each rachis commonly 2, approximate, laminiferous, few-nerved, of indurate but 

brittle substance, sparsely puberulent dorsally, one 

subtending a bud, the other empty. Spikelets up 

to 15 mm long, falling away at maturity, each at 

length leaving the ca 5 mm long terminal segment 

of its rachis almost completely clothed in a persistent bract. Transitional glumes at the base of 

each spikelet none. Anthecia papyraceous, glabrous 

and lustrous, without externally discernible nerves. 

Lemma up to 15 mm long, lanceate, attenuately 

acuminate. Palea shorter than the lemma, the sulcus very narrow, with its edges marked by a row of 

antrorse cilia. Lodicules diaphanous, narrow, 

sparsely ciliate on the margin apically. Anthers 

dark brown when dry, sagittate basally and emarginate apically, the tip of each half curving outward. 

Ovary glabrous throughout or antrorsely hispidulous apically. Style antrorsely hispidulous. Mature 

fruit not yet seen. 

ErYMoLocY.-The trivial name, falcata, alludes 

to the curvature that characterizes the branches of 

the inflorescences. This curvature is construed as 

induced by the sympodial nature of the branching 

of the individual rachises. 

HoLorYPE.-In the U.S. National Herbarium 

(no.2040296) collected in a “carrascal” [forest of 

small trees] at Esplanada, Bahia, Brasil, sometime 

during 1950-51, by Gerald0 Pinto (no. 0681). 

2. Atructantha radiata McClure, new species 

FIGURES 22, 23 

Plantarum habitus collectore neglectus. Culmi 

in specimine suppetenti neque satis representati 

neque descripti, ita ignoti. Vaginae culmi et vaginae ramorum primariorurn in specimine suppetenti 

deficientes. Vagina foliorum arcta, dorso glabra, 

opaca, nervis vix aut haud manifestis; auriculis vel 

nullis vel ad lineam reductis; setis oralibus crebris, 

superne saltem anfractis, omnino glabris, propter 

substantiam fragilissimam mox efiactis; ligula interiore ca 1.0 mm longa, dorso granulata, apice 

asymmetrica, margine integra; ligula exteriore arcuata, angusta, margine primo minute ciliolata 

demum glabrescenti; petiolo usque 5 mm longo, 

gracili omnino glabro; lamina usque 18 cm longa 

et 3.5 cm lata, lanceata, apice acuminata, basi inaequaliter vel rotundata vel subdeltoidea, utrinsecus 

glabra vel faciei abaxiali pro parte subtiliter puberula; nervis pluribus, validis, tertiariis a secundariis haud facile distinguendis, venulis transversis 

perpaucis inter se late distantibus. 

Inflorescentiae capitatae, in ramis ramulisque foliiferis insertae ubi et terminales et laterales, rhachidum ramificationes insertione distichae. Rhachidum segmentum terminale usque 10 mm longum, 

FIGURE 21.--A tractantha falcata McClure. A, Culm internode 

with branch complement of leafy and leafless flowering axes, 

x 0.6; B, base of branch complement, x ca. 3; c, apex of 

leaf sheath and base of leaf blade, x ca. 6; Da and Db variations in structures forming terminal portion of a flowering 

twig, diagramatic; E, laminiferous bract from base of a 

branch of the inflorescence; F, floret, x 6, G, lemma, x 6; H, 

palea (right) and prolongation of the rachilla (left), x 6; 

I, lodicule complement, x 15; J, stamen, x 6; K, gynoecium, 

x 30. All drawings based on Pinto 0681 (US). 
NUMBER 9 51 

H 
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 9 53 

apice usque 1.0 mm diametro dilatatum, omnino 

glabrum laevigatum nitidumque. Bracteae gemmam subtendentes in nodis basalibus omnis rhachidis vulgo 2, glabrae, substantia delicatissima, 

infima 2-3 mm, superiore usque 5 mm longa, non 

laminiferae. Spiculae usque 19 mm longae, omnes 

maturitate caducae, rhachidi sua tum conspicua 

relicta apicaeque nuda. Glumae transitionales 

nullae. Anthecia papyracea, dorso glabra nitidaque, 

nervis extus haud manifestis. Lemma usque 17 mm 

longum, lanceatum, attenuate acuminatum. Palea 

lemmate parum longior, seas latera sulci angusti 

ciliolis antrorsis crebris ornata. Lodiculae minimae 

angustae diaphanae acuminatae secus marginem 

superne ciliolatae. Antherae apice obtusae basi 

conspicue sagittatae, ubi desiccatae vinaceae. Ovarium superne pilosum. Stylus vel conspicue vel 

inconspicue hispidulus. Stigmata typice 2 (atypice 

et rarissime 3). Fructus maturus adhuc non visus. 

Plants of unrecorded habit. Culms neither adequately represented nor described in the available 

specimens. Culm sheaths and sheaths of the primary axis of branch complements lacking in the 

specimen. Leaf sheaths tight, glabrous, dull, the 

nerves scarcely or not at all visible externally; 

auricles either lacking or reduced to a line; oral 

setae crowded, curved distally, glabrous throughout, very fragile and soon broken off; the inner 

ligule ca 1.0 mm long, dorsally granulate, apically 

asymmetrical, the margin entire; the outer ligule 

arcuate, narrow, the margin minutely ciliolate at 

first, then glabrescent; petiole up to 5.0 mm long, 

slender, glabrous throughout; blade up to 18 cm 

long and 3.5 cm wide, lanceate, acuminate, rounded 

or subdeltoid at the asymmetrical base, either 

glabrous on both surfaces, or partly and minutely 

puberulous on the abaxial surface; many-nerved, 

the nerves strong, the tertiary ones not easily distinguished from the secondary ones; transverse veinlets few and far removed from each other. 

Inflorescences capitate, terminal and lateral to 

leafy branches and leafy twigs, the branches (raFIGURE 22.-Atractantha radiata McClure. A, Leafy flowering 

branch, x 0.6; B, apex of leaf sheath and base of leaf blade, 

x 3; c, base of branch complement, x 4.8; D, E, variants of 

flowering axes, x 0.6. All drawings based on Frdes I9947 

(US) * 

chises) of distichous insertion. Bracts at the base 

of each rachis typically 2, approximate, glabrous 

and of very delicate substance, not laminiferous, 

soon disintegrating, the lowermost one 2-3 mm, 

the uppermost up to 5 mm long; each subtending 

a bud. Spikelets up to 19 mm long, readily falling 

away at maturity, each leaving conspicuous and 

naked the apex of the terminal segment of its 

rachis; the latter glabrous, smooth and lustrous, up 

to 10 mm long and (at its slightly flared apex) up 

to 1 mm in diameter. Anthecia papery, glabrous 

and shining throughout, the nerves not at all visible 

externally. Lemma up to 17 mm long, lanceate, 

attenuately acuminate. Palea a little longer than 

the lemma, the margins of the sulcus close together 

and densely fringed with antrorse cilia. Lodicules 

diaphanous, very small, narrow, the margin apically 

ciliate. Anthers sagittate basally, obtuse apically, 

dark purple when dry. Ovary apically pilose. Style 

conspicuously or inconspicuously hispidulous. Stigmas typically 2, atypically and very rarely 3. Mature fruit not yet seen. 

ETYMoLocY.-The trivial name, radiata, alludes 

to the radiate orientation of the rachises of the 

capitate inflorescence. 

HOLOTYPE.-h the U. S. National Herbarium 

(nos. 1910764, 1910765, and 2146781) collected at 

“Estrada de Bom Gosto a Olivenca, Estado da Baia, 

Brasil, March 15, 1943,” by Ricardo de Lemos 

Froes (no. 19947). Additional material examined: 

an isotype from the same collection on loan from 

the herbarium of the Instituto Agronomico do 

Norte, Belem, Para, Brazil (IAN no. 15373). 

Aulonemia Goudot 

FICURE~ 24-26 

Aulonemia Goudot, 1846:75. 

Matudacalamus Maekawa, 1961:344. 

Plants unicespi tose, unarmed. Rhizomes pachy 

morph. Culms of small or medium stature and 

self-supporting, to very tall and upright, the apex 

erect, nodding, pendulous or scandent; each midculm node bearing a single initial (primary) branch 

bud, the prophyllum elongating simultaneously 

with the germination of the bud. Branch complement at midculm nodes typically comprising but a 

single initial (primary) axis, this dominant over 

axes of higher orders that may proliferate promptly 
I 

m 
NUMBER 9 55 

or tardily from usually functional buds borne at 

its own proximal nodes. Sheath at midculm nodes 

typically lacking a well-marked (conspicuous) basal 

girdle; the auricles and oral setae (varying from 

species to species) entirely lacking to well developed 

and scaberulous to entirely glabrous. Leaf blades 

with transverse veinlets not at all to weakly to 

sometimes strongly manifest externally. 

Inflorescences semelauctant, forming typically 

open panicles, the typically solitary, pulvinate or 

epulvinate primary ramifications with subtending 

bracts generally small, obsolete or lacking entirely. 

Spikelets few-flowered to many-flowered, terminating in a depauperate sterile anthecium. Transitional glumes 2 or 3; when three the uppermost one 

usually recognizable as a “sterile lemma.” Lemma 

in functional florets embracing its palea only basally at maturity. Palea broadly sulcate and 2- 

keeled dorsally, the margins not at all or barely 

imbricate. Stamens 3, the filaments filiform, free. 

Stigmas 2. Mature fruit (as far as known) an 

oblong or subfusiform, sulcate, apically mucronate 

caryopsis, the pericarp pergamineous, glabrous, of 

uniform thickness throughout or sometimes appreciably thicker apically, the sulcus and embryotegium clearly manifest. 

ETYMoLoGY.-The name Aulonemia, a Latinized 

combination of the Greek words, aulos (ancient 

name of a flute made from a shin bone) and nemos 

(a forest with pasture for cattle) alludes to (1) the 

musical instruments (called, in the vernacular, 

“queko”) made by the native inhabitants from 

elongated culm internodes of plants of the typespecies, and (2) the browse made accessible to 

herbivorous animals by the pendulous leafy culm 

tips of plants of the same species (cf. Goudot, 

1846:75, et seq.; R. W. Brown, 1954:112, 554). The 

terminal element, nemos, makes the natural gender of the word Aulonemia neuter; however, the 

vernacular trivial name, queko, does not reveal a 

FIGURE 23.-Atractantha radiata McClure. A, Primary pseudospikelet (early stage of development), x 15; B, two 

pseudospikelets, showing the prophyllum at the base of the 

one on the left, x 3; c, cluster of pseudospikelets, with two 

pedicels naked, x 3; D, diagram of a typical pseudospikelet; 

E, floret, x 9; F, lemma, x 9; G, palea (left) and prolongation of the rachilla (right), x 9; H, diagrammatic crosssection of palea, x 18; I, lodicule complement, x 15: J, 

stamen, x 6; K, gynoecium, x 15. All drawings based on 

Froes 19947 (US). 

choice of gender by Goudot. In the absence of 

any other precedent, Aulonemia was given the 

feminine gender by McClure and L. B. Smith (in 

Reitz, ed., 1967:47) when they attached to it the 

trivial name, lancipora. 

TypE-spEcIEs.-Aulonemia queko Goudot. 

RELATIoNSHIPs.--The array of 24 species here incorporated in Aulonemia imparts to the genus an 

admittedly polymorphic content. Some of the arms 

of marked diversity recognizable within the genus 

as I have circumscribed it involve species that show 

individual features suggesting the presence of genetic elements shared by several other genera. Aulonemia humillima alone is superficially grass-like 

in appearance. My study of fragmentary specimens 

and brief field notes provided by the solitary extant collection of Aulonemia hirtula suggests to me 

that it probably joins Aulonemia queko in being 

distinguishable from the other known species of 

the genus by the possession of relatively large culms 

with greatly elongated internodes, each followed by 

two or more nodes, the adjacent ones separated 

from each other by an aborted subobsolete internode. This peculiarity-without apparent linkage 

with any other stable attribute-also appears as a 

stable attribute of each plant in Glaziophyton mirabile, in a few species of Arthrostylidium (cf. 

McClure, in Maguire, Wurdack, et al., 1964:2), and 

quite generally in the known species of Myriocladus. Aulon,emia eflusa apparently is joined by Aulonemia deflexa in combining other vegetative 

features that suggest a phylogenetic heritage shared 

by members of Myriocladus, viz., the frequently delayed germination of primary branch buds at culm 

nodes and/or the branch bud at the proximal nodes 

of primary branches (a feature ostensibly related 

to the strongly indurated vegetative foliar appendages) and the closely appressed culm branches, each 

comprising conspicuously elevated numbers of short 

internodes and crowded foliiferous nodes. Aulonemia subpectinata shows a facultative but not obligate tendency to delay the germination of midculm 

branch buds, and to postpone proliferation from 

buds at the proximal nodes of the initial branch at 

midculm nodes. Aulonemai laxa and A. viscosa 

share this tendency. These two species also share 

with other species of Aulonemia, and with some 

species of both Aythrostylidium and Bambusa, a 

more or less conspicuous vinous tinting or macula- 
56 SMMITHSONIAN CONTRIBUTIONS TO BOTANY 

tion of either vegetative structures alone or reproductive structures alone, or both. To accommodate 

one of these last two species, Maekawa has proposed 

(196 1 : 344) the recognition of a monotypic genus, 

Matudacalamus. In the perspective resulting from 

my studies in the field and in various herbaria, 

Matudacalamus laxus Maekawa appears to be a 

member of one of the several nondisjunct arms of 

diversity that make up the polymorphic natural 

genus, Aulonemia. Maekawa expresses the opinion 

that the “wavy, smooth” oral setae of the leaf 

sheaths, and the “one-at-a-node” culm branches are 

two critical features that set off Matudacalamus 

from other genera he mentions as related to it. 

However, neither one nor both of these featuresnor any other combination of morphological features at present available to me-appears to supply 

either a reliable (stable) basis, or a justification, for 

segregating Matudacalamus from Aulonemia at the 

genus level. The mingling of racemose branching 

with the paniculate branching of the inflorescences 

of Aulonemia purpurata gives it a striking resemblance to the branching typical of the inflorescence 

in Colanthelia cingulata. The type-specimen of A. 

purpurata, however, does not include any feature 

that shows other affinities in that direction. 

DIsTRIBu’rIoN.-The named species here incorporated in Aulonemia gives the genus an aggregate 

recorded geographical and altitudinal range extending from Brazil, with 10 species at 800-2800 m, 

through Bolivia, with one species at about 3200 m; 

Peru, with four species at 1500-3000 m; Ecuador, 

with one species at 1800-3000 m; Colombia with 

three species at 2800 m; Venezuela, with three species at 1900-2760 m; Guyana, with one species at 

about 2700 m; Costa Rica, with one species at 1800- 

2700 m; to Mexico, with one species at about 

2200 m. 

Annotated Checklist of 

Bamboos Here Recognized 

as Belonging in the Genus Aulonemia 

1. Aulonemia amplissima (Nees) McClure, new 

combination. 

Arundinaria amplissima Nees, 1834:479. 

Arthrostylidium amplissimum (Nees) McClure, in 

Steyermark, et al., 1951:33. 

My dissections of florets from the type-collection 

reveal the number of stigmas to be regularly 2- 

instead of 3 as recorded by Nees (1834:479) and as 

reiterated by Ruprecht (1839:27). The number of 

stigmas is correctly given by Munro (1868:26) and 

by later authors. 

2. Aulonemia aristulata (Doell) McClure, new 

Arundinaria aristulata Doell, in Martius, 1880: 165. 

Sieglingia aristulata (Doell) Kuntze, 1898, I11 (3): 

341. 

Arundinaria mucronata Munro ex E.-G. Camus, 

19 12: 244. 

3. Aulonemia deflexa (N. E. Brown) McClure, 

combination. 

new combination. 

4. Aulonemia eflusa (Hackel) McClure, new combination. 

Arundinaria deflexa N. E. Brown, 1901:75. 

Arundinaria eflusa Hackel, 1903a:71. 

Arthrostylidium eflusum (Hackel) McClure, in 

Steyermark, et al., 1951:31. 

5. Aulonemia glaziovii (Hackel) McClure, new 

combination. 

6. Aulonemia goyazensis (Hackel) McClure, new 

combination. 

Arundinaria glaziovii Hackel, 1903a: 72. 

Arundinaria goyazensis Hackel, 1903a:71. 

7. Aulonemia haenkei (Ruprecht) McClure, new 

combination. 

Arthrostylidium haenkei Ruprecht, 183927. 

Arundinaria haenkei (Ruprecht) Hackel, 1903:69. 

Arundinaria setifera Pilger, 1905: 145. 

8. Aulon,emia herzogiana (Henrard) McClure, 

new combination. 

Arundinaria herzogiana Henrard, in Herzog, 1921 : 

75. 

9. Aulonemia hirtula (Pilger) McClure, new combination. 

Arundinaria hirtula Pilger, 192 1 :445. 

FIGURE 24.-Aulonemia queko Goudot. A, Leafy flowering 

branch, x 0.6; B, junction of leaf sheath and leaf blade, 

x ca. 1.8; c, insertion of lowermost branch of inflorescence, 

x ca. 6; D, spikelet, x 1.5; E, transitional glumes I, 11, and 

III, x 3.6; F, sterile lemma, x 3.6; c, floret, x 3.6; H, fertile 

lemma, x 3.6; I, palea, x 3.6; J, lodicule complement, X 

7.2; K, stamen, x ca. 6; L, gynoecium, x ca. 12: M, stigmatic 

processes, greatly enlarged. .411 drawings based on Goudot 

no. I (P). 

58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

10. Aulonemia humillima (Pilger) McClure, new 

Arundinaria humillima Pilger, in Engler and 

11. Aulonemia laxa (F. Maekawa) McClure, new 

Matudacalamus laxus F. Maekawa, 1961:345. 

12. Aulonemia panriflora (Presl) McClure, new 

Guadua parviflora, J. S. Presl, in K. B. Presl, 1825- 

Bambusa parviflora (Presl) J. A. and J. H. Schultes, 

in Roemer and Schultes, 1830, 7 (2):1350. 

Arthrostylidium maculatum Ruprecht, 1839:28. 

Arundinaria maculata (Ruprecht) Hackel, 1903a: 

69. 

13. Aulonemia patula (Pilger) McClure, new combination. 

Arundinaria patula Pilger, 1898:719. 

14. Aluonemia purpurata (McClure) McClure, 

Arthrostylidium purpuratum McClure, 1942: 170. 

15. Aulonemia queko Goudot, 1846:76, pl. 4 (Figures 24-26). 

Arthrostylidium queko (Goudot) Hackel, in Engler 

and Prantl, 1887:93 [sphalm. quexo]. 

Arundinaria queko (Goudot) Hackel, 1903a:74. 

Available specimens from collections supplementing the holotype represent this taxon as a 

species embracing plants somewhat diverse in respect to mature stature and habit, size, shape, 

texture, and vesture of leaf blades, disposition of 

the peripheral branches of the inflorescence, and 

degree of development of awns and dorsal pricklehairs on the lemmas. The wide altitudinal range of 

the natural distribution of plants, which I recognize 

as belonging to this species, suggests that some of 

the phenotypic diversities noted may be ecotypic 

in origin. More detailed studies in field and laboratory may, however, reveal the existence of genotypic 

bases for some of the observed diversities. 

In 1950 I had an opportunity to visit the typelocality of Aulonemia queko near La Trocha in the 

region of Colombia called Quindio. I found no 

sign of either the original forest, or the bamboo 

described by Goudot as Aulonemia queko. The 

combination. 

Prantl, 1906: 100. 

combination. 

combination. 

1835, I (4-5):257. 

new combination. 

mountainside is clothed instead with a beautiful 

carpet of Pennisetum clandestinum Hochstetter ex 

Chiovanda, a grass native to Africa. I eventually 

decided that specimens of a plant I had collected 

in Ecuador (McClure 21429) and in Peru (McClure 

21450) represent Aulonemia queko, in spite of 

minor deviations with respect to some of the 

features given this species by Goudot. This ended 

my own quarter-century of puzzlement over the 

riddle presented by Goudot’s deliniation of Aulonemia queko. It appears that Hackel’s action (in 

Engler and Prantl, 1887) in making Aulonemia a 

synonym of Arthrostylidium was induced by Ruprecht’s (1839) broad circumscription of Arthrostylidiunz. Again, Hackel’s (1903a) transfer of 

Alonemia queko to Artmdinaria apparently was 

suggested by Nees’ (1834) image of Arundinaria. 

The anguished efforts of Hackel (1903a) and Pilger 

(in Urban, 1900-1901) to find bases for distinct 

and stable generic images of the diverse elements 

that had been, at one time or another, placed in 

Arthrostylidium and Arundinaria are brought into 

focus elsewhere (p. 19). 

I identify as representing a species of Elytrostachys the specimen Fendler 2496 (a leafy twig in 

the vegetative state, preserved at Kew) cited by 

Munro (1868:46) in his treatment of Aulonemia 

queko. It seems likely that Munro’s modification of 

Goudot’s spelling of the specific epithet from queko 

to quexo misled Hackel (in Engler and Prantl, 

1887). In 1903a, however, Hackel restored the original spelling in the combination, Arundinaria 

queko. 

16. Aulonemia radiata (Ruprecht) McClure and 

Arundinaria radiata Ruprecht, 1839:25. 

17. Aulonemia ramosissima (Hackel) McClure, 

Arundinaria 1-amosissima Hackel, 1903a:74. 

Arundinal-ia glaziouii var. macroblephara E.-G. 

18. Aulonemia setigera (Hackel) McClure, new 

L. B. Smith, in Reitz, ed., 1967:56. 

new combination. 

Camus, 1913, I:41. 

combination. 

FIGURE P5.-8ulonemia queko Goudot. A, Seedling plant, 

x 0.6; B, twinned culm sheaths, x ca. 0.6: c, sterile leafy 

branch, x 0.6; D, apex of leaf sheath and base of leaf blade, 

x 1.5. Drawing A based on McClure 214298 (US), B, c, D on 

McClure 21429 (US). 
NUMBER 9 59 

NUMBER 9 61 

Arundinaria setigera Hackel, 1903a:73. 

Arthrostylidium aristatum Glaziou ex E.-G. Camus, 

19. Aulonemia sodiroana (Hackel) McClure, new 

Arundinaria sodiroana Hackel, 1930a:70. 

On the basis of the similarity of the inflorescences 

in their respective type-specimens, Arundinaria 

sodiroana was by Hitchcock and Chase made a 

synonym of Arundinaria patula. Plants of the two 

taxa may be distinguished, however, even in the 

nonflowering state by the distinctly prominent 

fimbriation of the outer margin of the leaf sheaths 

present in A. sodiroana but lacking in A. patula. 

20. Aulonemia steyermarkii (McClure) McClure, 

new combination. 

Arthrostylidium steyermarkii McClure, in Steyermark, et al., 1951:31. 

21. Aulonemia subpectinata (0. Kuntze) McClure, 

new combination. 

Arthrostylidium subpectinatum 0. Kuntze, 1891, 

II:760. 

22. Aulonemia trianae (Munro) McClure, new 

combination. 

Arundinaria trianae Munro, 1868:25. 

Arundinaria multiflora Doell, in Martius, 1880: 166. 

23. Aulonemia ulei (Hackel) McClure and L. B. 

Arundinaria ulei Hackel, 1903a:75. 

Species sedis mihi etiam nunc incertae manet. 

24. Aulonemia viscosa (Hitchcock) McClure, new 

Arundinaria viscosa Hitchcock, 1927a: 79. 

1913, I:67. 

combination. 

Smith, in Reitz, ed., 1967:57. 

combination. 

Checklist of New World Species Here 

Excluded from the Genus Aulonemia 

FIGURE 26.-Aulonemia queko Goudot. A, Upper culm nodes 

with buds and base of branch bud, x 0.6; B, buds enlarged, 

x 1.2; c, bud in advanced development, x 1.2; D, base of 

young branch complement, x ca. 1.2; E, tip of leafy branch 

terminating in an inflorescence, x 0.6; F, spikelet, x 2.4; 

c, transitional glume I, x 6; H, transitional glume 11, x 6; I, 

sterile lemma, x 6; J, fertile lemma, x 6 K, floret, showing 

palea, x 6; L, mature fruit (hilum side), x ca. 15; M, mature fruit (embryo side), x ca. 15; N, mature fruit, longitudinal section, x ca. 15; 0, lodicule complement, x ca. 6; 

P, gynoecium (without stigmas), x ca. 6. All drawings based 

on McClure 21429 (US). 

1. Aulonemia cingulata. See Colanthelia cingu2. Aulonemia intermedia. See Colanthelia inter3. Aulonemia lancipora. See Colanthelia lancilata. 

media. 

flora. 

Bambusa Schreber, Subgenus Guadua (Kunth) 

Hackel 

FIGURES 27-29 

Genus Cuadua Kunth, 1822a: 150; 1822b:254.-Ruprecht, 

1839:38 [1840: 1281.-Munro, 1868:70.-Doell, in Martius, 

1880: 176.-Bentham, in Bentham and Hooker, 1883: 1210.- 

McClure, 1957:203. 

Bambusa (Guaduae) Nees, in Martius, 1829:532. 

Bambusa Sect. I1 Guadua, Hackel, in Engler and Prantl, 

Bambusa (Guadua) Hackel, 1903c:194. 

1887:95. 

Plants typically unicespitose and typically armed. 

Rhizomes pachymorph, the rhizome neck in some 

species more or less elongated (exceptionally, as 

in McClure 21438 and 21438-A, to 5 m or more). 

Culms self-supporting below and apically nodding, 

pendent, or clambering; internodes cylindrical or 

terete, in some species shallowly to more or less 

deeply sulcate above the point of insertion of a 

bud or a branch complement; hollow, occasionally 

subsolid (teste Doell, in Martius, 1880: 186), exceptionally (as commonly in Bambusa amplexifolia) 

multifistular in the lower part of the culm. Branch 

buds at culm nodes typically solitary, lacking in the 

midculm range (in plants of some taxa at mature 

stature). Midculm branch complements lacking in 

mature plants of some species (as in some strains 

of Guadua angustifolia; see Bambusa guadua), 

facultative restricted monoclade in others; in most 

species unrestricted monoclade, of restricted insertion (the insertion rarely gremial, as in Guadua 

spinosa; see Bambusa swalleniana) the component 

axes more or less strongly unequal, the primary 

axis always dominant. Branches (at least those at 

lower nodes of the culms) typically thorny in all 

known species (see exception noted under Bambusa 

amplexifoZia). Leaves (leaf sheath blades) with 

transverse veinlets not at all visible to more or 

less clearly manifest externally. 

Inflorescences iterauctant, developing by way of 

pesudospikelets, the primary pseudospikelets either 

pedunculate or sessile, and forming sparse to more 
62 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

or less densely crowded, sometimes capitate aggregations, lateral or terminal to leafy or leafless axes. 

Spikelets typically comprising several perfect florets 

(only 2 in Bambusa capitata) and either tapering 

apically through progressively reduced sterile 

anthecia, or terminating abruptly in a single depauperate anthecium; the rachilla potentially 

disarticulating immediately below the locus of 

insertion of each fertile lemma, and in some species 

also below a sterile lemma. Transitional sheathing 

organs inserted between the uppermost bud-subtending bract and the first fertile lemma 0-2 (-3). 

[In some species of Bambusa subgenus Guadua, 

and in other genera in which branches of the inflorescences form pseudospikelets, sheathing structures occasionally reported as “sterile lemmas” 

sometimes prove to be bracts that are inserted at 

a level below the base of the spikelet proper, and 

subtend prophyllate buds that are easily mistaken 

for abortive florets. Cf. Munro 1868, pl. 5: fig. 4 

(such a bud) and fig. 5 (a floret) both referred to 

by Munro (1868: 153) as “flosculi fertilis.”] Lemma 

in functional florets fully embracing its palea only 

basally at maturity. Palea sulcate and 2-keeled 

dorsally, the keels typically more or less prominently 

winged (not more noticeably winged in Burchell 

7642 [cotype of Guadua refracta Munro] than in 

some species of the Old World genus Bambusa), 

the margins typically not at all or only barely 

imbricate. Lodicules typically 3, the anterior 2 

asymmetrical and paired (sometimes connate}, the 

posterior one symmetrical and smaller. Stamens 6 

(3 in Guadua spinosa; see Bambusa swalleniana), 

the filaments filiform, free. Style terminating in 2 

or 3 (rarely 4) stigmas or (exceptionally) undivided. 

Mature fruit an oblong or lagenoid, mucronate or 

subrostrate, in some species strongly dorsiventral 

caryopsis, the pericarp pergamineous or coriaceous, 

somewhat thickened apically, the sulcus and the 

basal position of the embryo clearly manifest. 

The name Guadua is the Latinized version of a 

vernacular term used by aboriginal natives of 

Ecuador and Colombia to designate plants of this 

subgenus. 

TYPE-sPECIEs.-McClure (1957:203) designated 

Guadua angustifolia Kunth as type-species of the 

genus Guadua. This species therefore becomes the 

type of subgenus Guadua of the genus Bambusa, 

but under the name Bambusa guadua Humboldt 

et Bonpland (Figure 27). 

RELATIoNsHIPs.-Bamboos hitherto allocated to 

the genus Guadua have no close relatives among 

known members of other New World genera. However, their common technical image is very similar 

to that of members of the Old World genus Bambusa. Kunth (1822a:150) failed to cite any morphological feature to differentiate his genus Guadua 

from Schreber’s genus Bambusa. The first promising effort in this direction was made by Munro 

(1868:76) in‘ these words: “The upper palea in 

Guadua has the keels much more distinctly and 

broadly winged than in any Bambusa.” However, 

the trivial taxonomic significance Munro attached 

to this feature is clearly apparent in the following 

sentence: “This genus is kept up by most authors; 

but I am unable, in the absence of a perfect fruit, 

to point out any good distinction, except a wellmasked one in geographical distribution, between 

it and Bambusa.” Munro then lists the features 

proposed by other authors as distinctive of Guadua 

as a genus, and shows that not one of them provides 

Guadua with a clearly distjunct morphological relation to Bambusa. My own observation is that 

the keels of the palea are not more noticeably 

winged in Burchell 7642 (cotype of Guadua refracta Munro) than in some species of the Old 

World genus, Bambusa. It also appears that subgenus Guadua and subgenus Bambusa are not 

sharply disjunct, with respect to features of the 

gross morphology of either their vegetative or 

reproductive structures or with respect to their 

geographical endemism. 

By 1829 Nees (in Martius, p. 532) had already 

made “Guaduae” an uncategorized division of the 

genus Bambusa. This proposal was more fully 

developed by Nees in Linnaea (1834:465). 

In 1887 Hackel (in Engler and Prantl, p. 95) 

proposed the formalization of the status of Guadua 

as Section I1 under Bambusa; in 1903 he (1903~: 

194) continued the subordination of Guadua under 

FIGURE 2T.-BamBusa guadua Humboldt and Bonpland, 

sensu lato. A, culm sheath, adaxial aspect, x 0.3; B, same, 

detail at base of blade, enlarged; c, upper portion of culm 

sheath, abaxial aspect, x 0.3; D, portion of first-order branch 

bearing a leafy twig with thorns, x 0.6; E, apex of leaf 

sheath and base of blade, x ca. 0.6; F, branch bud at culm 

node, x ca. 0.6; G, short-shoot thorny branch at lower node 

of culm, x ca. 0.6. Drawings A-D based on Gutierrez and 

Rarkley 17 C 609 (US), and E-G on Fosberg I9831 (US). 
NUMBER 9 63 
64 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

the genus Bambusa and proposed the elevation of 

its status from section to subgenus. 

DISTRIBUTION.-AS documented at present, the 

recognized New World components of this arm of 

the genus Bambusa exhibit an aggregate recorded 

distribution extending from Mexico to all countries 

of Central and South America, excepting Chile. 

They reach their greatest spontaneous development 

under mesophytic conditions in frost-free areas at 

elevations not known to greatly exceed 1500 m. 

Some species show a remarkable tolerance of, or 

ability to thrive in, soil that is saturated with water. 

I have observed that, in the general vicinity of 

Chinchina, Department of Caldas, Colombia, plants 

of Bambusa guadua occupy, spontaneously, practically every available type of ecological niche. 

Guadua philippinensis Gamble, an Old World 

species of diverse, as yet incompletely known, 

affinities, is known only by the type-collection (C. V. 

Piper 475). This collection was made 15 May 1911, 

at Mati, District of Davao, island of Mindanao. No 

information concerning the ecological setting of the 

plant has come to light. 

Agrostologists who find it difficult to accept 

Hackel’s disposition of the genus Guadua here 

adopted may be helped by the following words of 

Davis and Heywood (1963:106) quoted from their 

treatment of the category “genus”: “When in doubt 

as to whether to accord generic rank to a group, 

there is much to be said for the subgenus as a 

suitable category; it draws attention to the group 

in the classification, and at the same time allows 

people to continue to use the old binomial.” Final 

emphasis added. 

Annotated Checklist of Binomials 

Recognized Herein as Belonging to 

the Subgenus Guadua 

NEW WORLD BAMBOOS 

Improved perspectives resulting from the development of more complete images of the taxa 

represented by some of the binomials in the suggested synonymies may give occasion for future 

revision in some cases. 

1. Bambusa aculeata (Ruprecht) Hitchcock [as 

Guadua aculeata Ruprecht ex Fournier, 1881: 130. 

Guadua aculeata var. liebmanniana E.-G. Camus, 

Bambos], 1913:387 (Figure 28 k, 1). 

1913, I:112. 

Guadua inermis Ruprecht ex Fournier, 1881.129. 

Guadua intermedia Ruprecht ex Fournier, 1881: 130. 

Bambusa aculeata appears to represent a northern extension of Bambusa guadua (Guadua 

angustifolia) clearly distinguishable from the latter 

only by means of vegetative features. I have not 

been able to distinguish Guadua inermis from 

Guadua aculeata by means of inflorescences in their 

respective type-collections. Contrary to an assertion 

in the original description of Guadua inermis, the 

lemmas in the type-specimen are entirely glabrous, 

just as they are in the type of Guadua aculeata. If 

Fournier’s descriptive term “inermis” and the 

vernacular name “caiia mansa” quoted by him are 

authentic, then Guadua inermis may prove to be 

an unarmed form of Guadua aculeata from the 

northern range of that species. I do not at present 

see any basis for giving a distinctive taxonomic 

status to either Guadua inermis or Guadua 

aculeata var. liebman.niana. 

In 1877 Fournier (p. 198) published a paper 

entitled “De la modification des envelopes florales 

des GraminCes suivant le sexe de leurs fleurs” in 

which he referred to the spikelets of Guadua 

aculeata Ruprecht as embodying “a form of 

polygamy.” When, however, Fournier cited Guadua 

aculeata as the source of an illustrative example, 

this binomial had not yet been validated or given 

taxonomic content by the publication of a formal 

description. 

For descriptions of this same phenomenon as 

observed in bamboos of the genera Schizostachyum 

and Gigantochloa, among others, see McClure 

(1934:544ff; 1966b:100, 117) and Arber (1934:133). 

Instead of being considered as a reliable source of 

taxonomic characters, however, this morphological 

deviation should perhaps be seen simply as representing progressive transitional morphogenetic 

FIGURE 28.--Bambusa guadua Humboldt and Bonpland 

(A-J) and B. aculeata Ruprecht ex Fournier (K-L) . A, Leafy 

flowering twig, x 0.6; B, sterile leafy twig, x 0.6: c, pair of 

pseudospikelets, x 1.2; D, diagram of longitudinal section 

of pseudospikelet; E, bract and bud subtended by it at base 

of pseudospikelet, x 1.8; F, floret, x 3; G, lodicule complement, x ca. 5.4; H, stamen, x 6; x, gynoecium, x ca. 6; 

J, diagram of cross-section of floret; K, fruit, embryo side, 

x ca. 6; L, fruit, hilum side, x ca. 6. Drawing A based on 

Cobin 1167 (US), B-J on McCEure 21228 (US), and K, L on 

iMcClure 21556-A (US) . 
NUMBER 9 65 
66 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

changes in sexual expression that become potential 

during the later stages of the ontogeny of indeterminate (iterauctant) inflorescences in several 

bamboo genera. 

2. Bambusa amplexifolia (Presl) Schultes f., in 

Roemer and Schultes, 1830:1348. 

Guadua amplexifolia Presl, 1830:256. 

Plants of Bambusa amplexifolia are very thorny 

throughout most of the known range of the species 

(which extends from Venezuela and Colombia to 

Mexico) but progressively less thorny forms appear 

from El Salvador northward, and a completely 

unarmed form is found in the State of Sinaloa 

(specimen at US under McClure 21200). No mention of thorns appears in the original description 

of this species, the type of which was collected by 

Haenke from an unrecorded locality in Mexico. 

Cf. similar notes regarding Guadua inermis under 

Bambusa aculeata. 

3. Bambusa barbata Trinius, 1835:627. 

Nastus barbatus (Trinius) Ruprecht, 1839:41, pl. 17. 

4. Bambusa capitata Triflius, 1835:626; 1836, 

Schizostachyum capztatum (Trinius) Ruprecht, 

Guadua ? capitata (Trinius) Munro, 1868:81. 

Chromosome counts of 2n=ca. 36 and 2n=46 

were reported by Gould and Soderstrom (1967, 

under Guadua) for Brazilian collections of this 

species. 

Guadua distorta (Nees) Ruprecht, 1839: 131, pl. 16; 

111: PI. 337. 

1839 [1840: 136-137, pl. 17, fig. 46.1 

5. Bambusa distorta Nees, 1834:470. 

fig. 59. 

6. Bambusa glaziouii Hackel, 1903c: 194. 

glaziowii], 1913, I: 108. 

combination. 

Guadua glaziouii (Hackel) E.-G. Camus [as 

7. Bambusa glomerata (Munro) McClure, new 

Guadua glomerata Munro, 1868:79. 

Non invalidatus by Bambusa glomerata Royle ex 

iMunro (1868:147) [nom. nud. ex sched.] as synonym of Dendrocalamus strictus Nees. Cf. ICBN, 

1961, Art. 64. 

8. Bambusa guadua Humboldt et Bonpland, 

1808:63, pl. 20 (Figures 27, 28a-j, 29). 

Guadua angustifolia Kunth, 1822b: 253. 

Nastus guadua (Humboldt et Bonpland) Sprengel, 

‘3. Bambusa latifolia Humboldt et Bonpland, 

Guadua latifolia (Humboldt et Bonpland) Kunth 

The description under the name Guadua latifolia 

in Munro (1868:78), and the illustration under the 

same name in Doell, in Martius, (188O:pl. 49), are 

based on Spruce 1954, a collection that appears to 

represent a species quite distinct from the plant 

represented by the nomenclatural type of G. 

latifolia. 

Nastus latifolia [!I (Humboldt et Bonpland) 

10. Bambusa longifimbriata (E.-G. Camus) McGziadzra Zongifimbriata E.-G. Camus, 1913, I: 113. 

A specimen from the type-collection of this 

species (Glaziou 5717) (US from C, where it is 

labeled Guadua refracta Munro) differs from the 

type of the latter species by having the keels of the 

palea prominently winged. 

11. Bambusa longifolia (Fournier) McClure, new 

The basionym of the combination Arthrostylidium 

longifolium (Fournier) E.-G. Camus (1913, 1:68) 

is Arundinaria ? longifolia Fournier (1881: 131) the 

type of which is a sterile Liebmann specimen cited 

by Fournier as “Jicaltepec, Aprili, Lieb.” In his 

description of Arthrostylidium longifolium, Camus 

(1913, I:68) combines with his verbatim French 

translation of Fournier’s description of Arundinaria 

longifolia a description, in French, of a flowering 

1825, vol. 2:113. 

1808:67, pl. 21. 

1822: 254. 

Sprengel, 1825, 11: 113. 

Clure, new combination. 

combination. 

FIGURE 29.-Barn busa guadua Humboldt and Bonpland, sensu 

lato. A, Base of culm with its rhizome and the precociously 

developed necks that help the root system support the culm, 

which may be as much at 37 meters tall and 20 cm in 

diameter, x ca. 0.06; B, tangential slice from internode and 

nodes v and VI of a culm ca. 3.5 cm in diameter, x 0.6; C, 

sector of cross-section of culm wall at internode IV, x 0.6; 

D, apex of culm sheath, adaxial aspect, x 0.6; E, basal part 

of first-order branch from midculm node, with buds still 

intact, x ca. 0.6; F, second-order branch with buds still 

intact, x ca. 0.6; 6, third-order branch showing spines and 

foliage, x ca. 0.6; H, thorn complement with leafy twig 

arising from proximal node of thorn, x ca. 0.6; I, J, 

examples of extreme forms of leaves, x ca. 0.6; K, apex of 

leaf sheath and base of blade, enlarged. Drawings A, B based 

on McClure 21232 (US) and C-K on McClure 21215 (US). 
NUMBER 9 67 
68 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

specimen which he illustrates and cites as “Las 

Sedas. Altitude 2000 m. (Pringle).” Available 

material from the type and other collections of 

Arundinaria longifolia Fournier matches specimens 

of Arthrostylidium spinosum Swallen (1938:6) 

whose type I identify as a species of Bambusa 

subgenus Guadua. The name now becomes Bambusa 

longifolia. The Pringle specimen cited by Camus 

(of which I have three sheets available at US) 

turns out to be a duplicate of Pringle 6742. 

This I identify as belonging to the newly described 

subgenus Otatea of the genus Yushania. Meanwhile 

numerous specimens of bamboos pertaining to this 

new subgenus are filed in herbaria around the 

world under the name “Arthrostylidium longifolium 

(Fournier) E.-G. Camus.” 

12. Bambusa macrostachya (Ruprecht) McClure, 

new combination. 

Guadua macrostachya Ruprechb, 1839:39, pl. 15: 

Guadua dioica Steudel, 1854: 334, teste Doell, in 

Martius, 1880: 182. 

13. Bambusa maculosa Hackel, 1903c: 196. 

Guadua maculosa (Hackel) E. -G. Camus, 1913, 

I: 106. 

14. Bambusa paniculata (Munro) Hackel, 1903c: 

195. 

Guadua paniculata Munro, 1868:85. 

Bambusa munroi Hackel, 1909b3374. 

Non invalidatus by Bambusa paniculata Willdenow ex Munro (1868:123) [nom. nud. ex sched.] 

as synonym of Nastus borbonicus Gmelin (cf. ICBN 

1966, Art. 64). 

15. Bambusa paraguayana (Doell) Bertoni, 1918: 

159. 

Guadua paragunyana Doell, in Martius, 1880: 179. 

16. Bambusa refracta (Munro) McClure, new 

Guadua refracta Munro, 1868:84. 

17. Bambusa spinosissima Hackel, 1903c: 197. 

Guadua spinosissima (Hackel) E.-G. Camus, 1913; 

I: 112. 

18. Bambusa superba (Huber) McClure, new combidation. 

Guadua superba Huber, 1904:479. 

19. Bambusa swalleniana McClure, new name. 

Guadua spinosa (Swallen) McClure, 1954:82 (not 

fig. 35. 

combination. 

Bambusa spinosa Roxburgh, 1832: 198.) 

Arthrostylidium spinosum Swallen, 1938:6. 

20. Bambusa tagoara Nees, in Martius, 1829:532. 

Guadua tagoara (Nees) Kunth, 1834:611. 

21. Bambusa tessmannii (Pilger) McClure, new 

Guadua tessmannii Pilger, 1924: 124. 

combination. 

22. Bambusa tomen.tosa (Hackel and Lindman) 

Guadua tomentosa Hackel and Lindman, in 

McClure, new combination. 

Lindman, 1900:20, pl. 12. 

23. Bambusa trinii Nees, 1834:469. 

Guadua trinii (Nees) Nees ex Ruprecht, 1839:40, 

pl. 15, fig. 38. 

Guadua trinii var. p scabra Doell, in Martius, 

1880: 179. 

Bambusa tacuara Arechavaleta, 1897:550, pl. 72; 

teste Parodi, 1936:239. 

24. Bambusa iiograndensis Dutra, 1938:147, fig. 1. 

Guadua riograndensis (Dutra) Herter, 1941:49. 

Guadua ribbentropii Herter, 1940: 148 (based on 

Bambusa tacuara Arechavaleta). 

25. Bambusa venezuelae (Munro) McClure, new 

Guadua venezuelae Munro, 1868:86. 

combination. 

26. Bambusa uirgata Trinius, 1835:624. 

Guadua uirgata (Trinius) Ruprecht, 1839:40. 

27. Bambusa weberbaueri (Pilger) McClure, new 

Guadua weberbaueri Pilger, 1905: 152. 

combination. 

OLD WORLD BAMBOOS 

28. Bambusa philippinensis (Gamble) McClure, 

Guadua phi lip pinensis Gamble, 191 3:203. 

new combination. 

Checklist of 

Names of Congeneric New World Taxa 

of Unresolved Specific Status 

1. Bambusa chacoensis Rojas, 1918:157. 

2. Guadua fascicularis Doell, in Martius, 1880: 186. 

3. Guadua lindmani E. -G. Camus, 1913:1113. 

4. Guadua polyclados Doell, in Martius, 1880: 182. 

Guadua sp., Lindman, 1900:22, pl. Ilc. 
NUMBER 9 

Annotated Checklist of 

New World Species Here Excluded 

from the Subgenus Guadua 

1. Guadua exalata Doell (in Martius, 1880:181) 

is Arthrostylidium longiflorum Munro (1868: 

4 1). 

2. Guadua ? flabellata Fournier (1881: 131). 

Arundinaria flabellata (Fournier) McClure (in 

Maguire, Wurdack, et al., 1964: 162), species 

sedis mihi etiam nunc incertae; fortasse ad 

genus Yushaniam subgenus Otateam attinit. 

Guadua pallescens Doell (in Martius 1880: 186) 

syn. Bambusa pallescens (Doell) Hackel (1908: 

160), identified by Bentham (in Bentham and 

Hooker, 1883: 1210) as Bambusa pallida Munro, 

is Bambusa tuldoides Munro (1868:93), an Old 

World species naturalized in Brazil. 

Guadua parviflora J.S. Presl (in K. B. Presl, 

1830:257). See Aulonemia paroiflora (Presl) 

McClure. 

Guadua perligulata Pilger (in Diels, 1937:8) is 

Chusquea perligulata (Pilger) McClure, new 

combination. 

Chusquea Kunth 

FICURFS 30, 31 

Chusquea Kunth, 1822a: 151; 1822b:254; 1829:138; 1833:427.- 

Nees, 1834:484.-Endlicher, 1836: 102.-Ruprecht, 1839:30 

[1840: 1201.-Munro, 1868:52.-Doell, in Martius, 1880: 194.- 

Bentham, in Bentham and Hooker, 1883: 1209.-Hackel, in 

Engler and Prantl, 1887:93.-Arechavaleta, i8!37:542.- 

McClure, in Swallen, 1955:86; 1957:202. 

Rettbergia Raddi, 1823: 17.-Nees, in Martius, 1829:535.-Nees, 

1834:486.-McClure, 1957:207. 

Dendrugrostis Nees [as subgenus], 1834:487.-McClure, 1957: 

202. 

Plants of unicespitose or multicespitose or diffuse 

habit; either unarmed throughout or (as in C. 

Jendleri Munro; cf. McClure, 1966b, fig. 15) the 

culms armed by a ring of short, sharp, aborted roots 

at several of their lower, above-ground nodes. 

Rhizomes either wholly pachymorph (as in most 

known species) or wholly leptomorph (as in C. 

simpliciflora Munro) or both pachymorph and 

leptomorph in the same plant (as in C. fendleri). 

Culms either self-supporting below and broadly 

arched or clambering above (as in C. scandens 

Kunth), or wholly dependent upon external support from other vegetation (as in C. simpliciflora 

69 

Munro); the internodes pithy to the very center 

and (as in Zea mays L.) lacking a natural lumen, 

cylindrical or terete, in most species shallowly 

sulcate for some distance above the locus of insertion of a complement of buds or branches-rarely 

(commonly in some variants of C. pinifoliu Nees) 

sulcate or noticeably flattened all the way from 

one node to the next. [In some species, as in C. 

pittieri Hackel, the pith in old culms may become 

shrunken or broken down, leaving an irregular 

central passage without a membrane-lined inner 

surface. This same change may explain discrepant 

descriptions of C. uruguayensis Arechavaleta, whose 

culms are described by Arechavaleta (1897:546) as 

solid, and by Parodi (1941:334) as hollow.] Branch 

complement at midculm nodes unrestricted pleioclade, the initial components-developed from 

separate primary buds of two size categories in 

constellate insertion, the smaller ones usually many 

(rarely only 2)-comprising axes of two size categories (major and minor) the major ones solitary 

and (when developed) strongly dominant (similar 

in form to the mother culm and, particularly in 

scandent species, often approaching it in size) the 

minor ones (rarely only 2) usually numbering 

several to many and, when many hiding the large 

central bud from sight when it does not germinate. 

Leaves (leaf sheath blades) with transverse veinlets 

in some species (as in C. lanceolata Hitchcock) 

clearly evident, in others obscure, and in most 

species not at all manifest, externally. 

Inflorescences semelauctant, terminal to leafy or 

leafless branches, or to the culm itself, typically 

with a strong, variously branched central axis, 

usually paniculate, rarely either racemose or 

capitate; prophylls and bracts usually lacking entirely, bracts subobsolete in a few species (e.g., C. 

bambusaeoides (Raddi) Hackel). Transitional 

glumes at the base of the spikelet usually 4 (-5, as 

sometimes in C. bambusaeoides): I and II “empty 

glumes” (these in some species subobsolete), III, 

IV and (-v) “sterile (empty) lemmas.” Spikelets 

typically containing but one perfect floret-occasionally 2 in some species (e.g., C. andina 

Philippi, teste Munro (1868:58); and C. tenella 

Nees, teste Nees (1834:493); specimens under 

McClure 21292 (C. oxylepis (Hackel) Ekman) commonly show an aborted floret in the axil of transitional glume IV (“sterile lemma” II)-the rachilla 
70 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 9 71 

FIGURE 30.-Chusquea scandens Kunth. A, Leptomorph rhizome turning up to form a culm, x 0.6; B, lower segment of 

a small culm with buds, x 0.6; c, midculm node showing 

complement of branch buds, front view, x 1.2; D, same, in 

profile, x 1.2; E, apex of culm sheath, abaxial view, x 1.2; 

F, apex of culm sheath, adaxial view, showing ligule; G, 

midculm node with fully developed branch complement, 

x 0.6; H, auxiliary branch, showing leaves at apex, branch 

sheath, and a complement of branches emerging extravaginally, x 0.G; I, apex of leaf sheath with petiole and base 

of leaf blade, x 12. All drawings based on McClure 21414 

(US). 

very short and not prolonged behind the palea of 

the terminal perfect floret. [My personal impression in this regard agrees with that expressed by 

Bentham (in Bentham and Hooker, 1883:1209) in 

Latin: “According to Doell, the rachilla in C. 

tenella and other species [of this genus] is prolonged beyond the apical floret a condition which 

we have not seen in our specimens.” (Emphasis 

added by F. A. M.)] Fertile lemma fully embracing 

its palea only basally at maturity. Palea gaping 

antically, in most species dorsally convex, narrowly 

sulcate and more or less noticeably 2-keeled toward 

the usually emarginate or bidentate or bimucronate 

apex, rarely (as in C. capitata Nees) very similar 

to its lemma, i.e., compressed and l-keeled (not 

sulcate) dorsally and apically entire. Lodicules 3, 

the anterior two appreciably asymmetrical and 

paired, the posterior one symmetrical and smaller. 

Stamens 3, the filaments filiform, free. Stigmatic 

branches 2, one of them rarely (as sometimes in C. 

scandens Kunth, for example) divided above the 

point of its union with the other one. Mature 

fruit (rarely found) a small, oblong or fusiform 

cuspidulate or bicorniculate caryopsis with a 

pergamineous pericarp of uniform thickness, the 

sulcus and the embryotegium showing different 

degrees of conspicuousness in different species. 

ETYMoLoGY.-The name Chusquea is the Latinized 

version of a vernacular term used by aboriginal 

natives of Ecuador and Colombia to designate 

plants of this genus. 

TYPE-sPECIEs.-ChUsqUeU scandens Kunth (1822b: 

254); Nastus chusque Humboldt, Bonpland, and 

Kunth (1816, I:201). 

genus Neurolepis share a spikelet structure (Figure 

41 c-L, N) that is basically similar to that universal 

in Chusquea, members of the latter genus diverge 

RELATIoNsHIPS.-Although all members of the 

conspicuously from Neurolepis by their bambusoid, 

deciduous leaf blades, their branched culms with 

efistulose internodes, and their more prominently 

developed rhizomes. 

Plants of the closely related genus Swallenochloa, 

are distinguished from plants of the genus Chusquea by the relatively weak development of their 

rhizomes: the usually stiff, erect culm habit; the 

hollow structure of the culm internode; the typically horizontal and level (not constellate) insertion 

of branch buds at culm nodes, and the correspondingly different branching habit of the culm; the 

more or less strongly xeromorphic (leathery) texture of the leaf blades. The very narrow profile of 

the inflorescences of Swallenochloa is not diagnostic, 

however. 

The existence of noticeable affinities or similarities between members of the genus Chusquea and 

members of the genus Arundinaria is suggested by 

Trinius (1835:617), Baillon (1894:253), and Beadle 

(in Bailey, 1914:449). As far as the reproductive 

apparatus is concerned, similarities between members of these highly dissimilar genera are limited to 

some aspects of the lodicules and the stamens. 

As far as elements of the vegetative apparatus are 

concerned, the only noticeable similarities are 

shown by those species of the respective genera 

that share the following three features: (1) leptomorph rhizomes; (2) leaf blades with conspicious 

transverse venation; and (3) frost-hardiness. These 

three features are not known to be associated in 

any species of any one of the other bamboo genera 

of the New World. I do not construe their association together in exceptional species of Chusquea 

to be evidence of a close phylogenetic relationship 

or affinity between the two genera, Chusquea and 

Arundinaria. 

Nees (1834:467, 468) gave Chusquea Kunth four 

subgenera: (1) Platonia (Kunth) with C. (Platonia) 

elata (Kunth) listed as type; (2) Chusquea with 

C. scandens Kunth listed as type; (3) Rettbergia 

(Raddi) with C. (Rettbergia) gaudichaudii Kunth 

listed as type; and (4) Dendragrostis Nees with C. 

(Dendragrostis) pinifolia Nees listed as type. Subsequent authors have not maintained taxonomic 

status for the subgenera proposed by Nees. The 

originally monotypic content of Platonia Kunth 

(non Platonia Martius) was restored by Meisner 

NUMBER 9 73 

FIGURE 31.-Chusquea scandens Kunth (A-M) and C. oxylepis 

(Hackel) Ekman (N-P). A, Leafy flowering complement of 

branches of secondary order, x 0.6; B, inflorescence terminating a primary branch, x 0.6; c, terminal ramifications of 

inflorescence with persistent transitional glumes (“empty 

glumes” I and 11), x 7.5; D, spikelet with its pedicel, x 7.5; 

E, transitional glumes 111 and IV (”sterile lemmas”), x 7.5; 

F, lemma, x 7.5; G, palea, x 7.5; H, lodicule complement, 

x 9; I, stamen, x 7.5; J, gynoecium, x 9; K, terminal 

branches of inflorescence, x 7.5; L, fruit, embryo side, x 

7.5; XI, fruit, hilum side, x 7.5; N, fruit, embryo side, x 7.5; 

0, fruit, hilum side, x 7.5; P, fruit, embryo side, x 7.5; Q, 

diagram of spikelet. Drawing A-J based on McClure 21414 

(US), K on Bonpland 3386, type-species (P) , L, M on Fosberg 

22386 (us), and N-P on McClure 21292 (us). 

(1843, I:426; 1843, II:325) to its original status as 

genus, with a new name, Neurolepis. 

As I visualize the natural boundaries of the 

genus, Chusquea embraces taxa that’give it a polymorphic composition. One arm of this diversity 

induced Raddi to publish Rettbergia as a monotypic genus. As observed by Nees (in Martius, 

1829:536) a number of the features attributed to 

the type-species of Rettbergia by Raddi (1823:18, 

pl. 1: fig. 1) suggest faulty observation on the part 

of its author. Subsequent study of numerous available specimens collected from the type-locality 

confirms the opinion-first expressed tentatively by 

Nees (in Martius, 1829:536) and tacitly affirmed 

by Kunth (1830:331), Ruprecht (1839:34), Munro 

(1868:67), and Doell (in Martius, 1880: 198)-that 

the type-species of Rettbergia does not fall outside 

the natural boundaries of the genus Chusquea. 

However, the trivial name of the type-species of 

Rettbergia (R. bambusoides [as bambusaeoides] 

Raddi) was not formally incorporated in the genus 

Chusquea until its claim to priority over C. gaudichaudii Kunth was given expression in the combination C. bambusoides (Raddi) Hackel (in Wettsteii, ed., 1908:81). 

Chusquea bambusoides is joined by C. capituliPora, C. sellowii, and C. capitata to form a clearly 

recognizable but apparently not clearly disjunct 

arm of diversity, in which C. capitata stands most 

sharply distinguished from other members of the 

genus by the form of the palea, which here approaches that of the lemma. Another arm of diversity is rendered clearly disjunct from Chusquea 

proper principally by distinctive features of the 

vegetative apparatus. It is represented by Chusquea 

depauperata, C. spicata, C. subtessellata, and C. 

weberbaueri which now constitute the published 

content of the genus Swallenochloa, q.v. 

Chusquea pinifolia (Nees) Nees (1834:4WO), 

Awndinaria pinifolia Nees (in Martius, 1829:525), 

represents a taxon of special interest. Its polymorphic character finds expression in the following 

validly published names: C. baculifera Alvaro da 

Silveira (1919:99), C. heterophylla Nees (1835:488) 

with three variants named by Doell (in Martius, 

1880:207), and C. pinifolia var. heterophylla (Nees) 

Hackel (in Wettstein, ed., 1908:82). Since, however, 

they apparently have not been shown to represent 

morphologically disjunct entities, they are currently 

regarded as synonyms of a highly polymorphic 

C. pinifolia. In the habit of the plants, the form of 

the midculm branch complement, and the form of 

the inflorescences, some of the divergent forms of 

C. pinifolia bear superficial resemblance to members of the genus Swallenochloa. Their basic 

morphology, however, keeps plants of this taxon 

clearly within the natural limits of the genus 

Chusquea, their aggregate range in distribution 

extending from slightly above sea level to elevations 

(in the midtropics) reaching 3200 m, in generally 

mesophytic sites between about 47OS latitude and 

Mexico, and on islands adjacent to this area. 

DIsTRIsuTIoN.-Documented in herbaria by specimens of more than 100 described species, the genus 

Chusquea is represented by one or more of these 

in the indigenous flora of every country of the 

New World from Mexico to Chile and Agentina, 

and that of many of the islands as well. 

The aggregate altitudinal coverage recorded for 

its members carries the genus from approximately 

sea level to the lower limit of perpetual snow. 

According to Munro (1868:61) Jameson (in the 

field notes to a collection under Jameson 9) recorded the altitudinal range of Chusquea fendleri 

in Ecuador as “a litore usque ad 12,000 ped. s. m.” 

Chusquea scandens Kunth was collected by W. H. 

Camp (E-1778 and E-1824, US) in Ecuador at an 

elevation of 11,000 ft (3300 m) and C. andina 

Philippi was collected by Germain in Chile “ad 

limitem nivis perpetuae” (Munro, 1868:58). 

Ranges of tolerance and of requirements, in 

terms of ecological factors, are not adequately 

recorded either in the available field notes or in 

the literature; but the entire gamut covered by 

members of the genus Chusquea must be very 

comprehensive. According to Brade (1956) as re- 
74 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

viewed by F. Markgraf (1958) the zone occupied 

by Chusquea pinifolia (Nees) Nees is recognizable 

on Itatiaia at elevations ranging from 2000 to 2400 

meters. [Hackel (in Wettstein, ed., 1908:82) gives 

the altitudinal range of C. pinifolia on Itatiaia 

as 1300 m to 2750 m.] The meteorological station 

at 2200 meters recorded an annual rainfall (as the 

average for a period of 20 years) of 2417 mm 

(roughly 95 in) and a temperature range between 

an average low of 7” and an average high of 11 O C; 

extremes recorded: -6O and 35” C. Brade lists the 

following taxa that appear along with Chusquea 

pinifolia as characteristic of the vegetation of this 

high tableland: Cortaderia modesta, Paepalanthus, 

Xyris, Sisyrinchium, Clematis, Fuchsia, Buddleia 

speciosa, Fragaria chiloensis, Valeriana, Mikania, 

and Erigeron. 

Weberbauer (191 1 [ 1945: 1461) summarizes his 

notes on the ecological role of Chusquea in the 

vegetation of Peru as follows (trans. by FAM): 

Chusquea is a genus characteristic of the cloud forest 

(“Ceja de IaMonta‘iia”); in the north it extends over to the 

western side of the Andes. In its true center of distribution 

it sends out extensions that reach down as far as the tropical 

forest. The shrubby species of the genus Chusquea exert a 

conspicuous influence on the physiognomy of the plant formations of the cloud forest, and here and there occupy large 

areas, suppressing other plants. Their slender branches, from 

whose nodes emerge masses of twigs and dense foliage, support themselves on other woody plants, from which they 

hang down in beautiful curves. 

It should be borne in mind that Weberbauer’s 

concept of the genus Chusquea included the species 

treated herein under Swallenochloa, q. v. These 

are referred to by him as “shrubby species” (vide 

infra.). On the other hand, his concluding sentence 

refers to species I retain in Chusquea. 

Annotated Checklist of 

Bamboos Here Recognized 

as Belonging to the Genus Chusquea 

1. Ch usquea abietif olia Grisebach, 1864: 529. 

Arundinaria (Arthrostylidium?) microclada Pilger, 

in Urban, ed., 1907:289. 

Apparently this taxon is known only by vegetative material from the type-locality; the nomenclatural type-collection remains undesignated. Pilger’s 

uncertainty as to the correct generic disposition of 

this taxon is patent. 

2. Chusquea acuminata Doell, in Martius, 1880: 

204. 

Chusquea tenuis Glaziou ex E.-G. Camus, 1913, 

I:90. 

3. Chusquea afinis Munro ex E.-G. Camus, 1913, 

I:80, pl. 60: fig. B. 

4. Chusquea andina R. A. Philippi, 1858:103. 

[In rough notes McClure compares this species 

with Chusquea culeou Desvaux, perhaps indicating 

that it is synonymous with it. Munro (1868:58-59) 

expressed the opinion that this species is very 

probably only an alpine form of C. cu1eou.-T.R.S] 

5. Chusquea anelythra Nees, 1834:491. 

6. Chusquea anelytroides Ruprecht ex Doell, in 

Martius, 1880: 206. 

7. Chusquea argentina Parodi, 1941:339, pl. 24: 

8. Chusquea bambusoides (Raddi) Hackel, in 

Rettbergia bambusaeoides Raddi, 1823: 18, 57, pl. 

Chusquea gaudichaudii Kunth, 1829: 138. Nomen 

Chusquea gaudichaudii Kunth, 1830:331-332, pl. 

Nastus bruneus A. N. Desvaux, 1831:211. 

Nastus bruneus and some other collections have 

the awns of the empty glumes antrorse ciliate or 

antrorse scabrous on the dorsal keel of the awn. 

This is a variable character. 

fig. 4. 

Wettstein, ed., 1908:81. 

1: fig. 1. 

nudum. 

78. 

9. Chusquea bambusoides var. minor McClure 

and Smith, in Reitz, ed., 1967:25. 

10. Chusquea bilimekii Fournier, 1881: 132. 

11. Chusquea capitata Nees, 1834:489. 

Rettbergia capitata Nees was cited as a synonym 

of C. capitata in Munro (1868:69) and Doell (in 

Martius, 1880: 196). Nees had regarded Rettbergia 

as synonymous with Chusquea and published the 

species as C. [K (ettergia)] capitata. 

12. Chusquea capituliflma Trinius, 1835:613. 

The spikelets sometimes contain two florets in 

this species. 

13. Chusquea capituliflora var. pubescens McClure 

and Smith, in Reitz, ed., 1967:28. 

14. Chusquea carinata Fournier, 1881: 132. 

15. Chusquea culeou Desvaux, in Gay, 1854:450. 
NUMBER 9 75 

16. 

17. 

18. 

19. 

20. 

21. 

22. 

23. 

24. 

25. 

26. 

Chusquea culeou forma longiramea Parodi, 

1941 :343. 

Chusquea cumingii Nees, 1835:487. 

Chusquea paroifolia R. A. Philippi, 1864:299. 

Chusquea decolorata Munro ex Parodi, 1945: 

Chusquea deficiens Parodi, 1941:335-338, figs. 

2,3, and pl. 22. 

Chusquea delicatula Hitchcock, 1927b:309-3 10. 

Chusquea discolor Hackel, 1903b: 155. 

Chusquea dombeyana Kunth, 1832:553, pl. 191. 

Chusquea fasciculata Doell, in Martius, 1880: 

Chusquea fendleri Munro, 1868:61. 

Chusquea fernandeziana R. A. Philippi, 1873: 

577-578. [May be the same as C. Zigulata 

Munro.] 

Chusquea galeottiana Ruprecht ex Munro, 

1868: 59. 

65-66. 

202-203, PI. 54. 

Chusquea galeottiana Ruprecht, in Galeotti, 

1842:246. Nomen nudum. 

27. Chusquea gracilis McClure and Smith, in Reitz, 

ed., 1967:43-44, pl. 8i-k. 

28. Chusquen heydei Hitchcock, 1927a:80-81. 

29. Chusquea huantensis Pilger, 1920:29-30. 

30. Chusquea ibiramae McClure and Smith, in 

Reitz, ed., 1967:40-42, pl. 8d-f. 

31. Chusquea inamoena Pilger, 1905: 150-151. 

32. Chusquea jamesonii Steudel, 1854:337. 

33. Chusquea juergensii Hackel, 1909a:325-326. 

34. Chusquea lanceolata Hitchcock, 1935: 145-146. 

35. Chusquea lehmannii Pilger, 1899:35-36. 

Chusqziea pilgeri E. -G. Camus, 1913, 1:83. [Type 

cited as “C. leibmanni [sic!] Pilger . . non Fournier,” error for C. Zehmannii Pilger which is valid, 

as is C. liebmannii Fournier.] 

36. Chusquea leptophylla Nees 1835:489. 

Arthrostylidium trinii sensu Steudel, 1854:336, 

in part, non Ruprecht, 1839:119. 

Arthrostylidium (?) leptophyllum (Nees) Doell, 

37. 

38. 

39. 

40. 

41. 

42. 

in Martius, 1880:175-176. 

Chusquea liebmannii Fournier, 1881: 132. 

Chusquea ligulata Munro, 1868:62. 

Chusquea linearis N. E. Brown, 1901:76. [Probably equals C. pinifoh (Nees) Nees.] 

Chusquea longifolia Swallen, 1940:210. 

Chusquea longipendula Kuntze, 1898, I11 (3): 

Chusquea lorentziana Grisebach, 1874:249-250. 

348-349. 

43. Chusquea macrostachya R. A. Philippi, 1896: 

44. Chusquea mexicana Hackel, 1902:256. 

45. Chusquea meyeriana Ruprecht ex Doell, in 

Martius, 1880:203-204. 

C. meyeriana is closely related to C. acuminata 

Doell but differs in the pubescent leaf blades, 

prominent external ligule, reflexed inflorescence 

branches, and smaller spikelets. 

46. Chusquea mimosa McClure and Smith, in 

Reitz, ed., 1967:37. 

47. Chusquea montana R. A. Philippi, 1864:298- 

299. 

48. Chusquea muelleri Munro, 1868:65. 

49. Chusquea nelsonii Scribner and Smith, 1897: 

16. 

50. Chusquea nigricans R. A. Philippi, 1865:323- 

324. 

51. Chusquea oligophylla Ruprecht, 1839: 124, pl. 

7: fig. 23. 

52. Chusquea oxylepis (Hackel) Ekman, 1913:65, 

pl. 4: fig. 6 (Figure 31~-~). 

Chusquea bambusoides (Raddi) Hackel subsp. 

oxylepis Hackel, in Wettstein, ed., 1908:81-82. 

53. Chusquea palenae R. A. Philippi, 1896:350. 

54. Chusquea pallida Munro, 1868:65. 

55. Chusquea paruipora R. A. Philippi, 1896:349- 

350. 

56. Chusquea perligulata (Pilger) McClure, new 

combination. 

Guadua (7) perligulata Pilger, in Diels, 1937:57-58. 

57. Chusquea peruuiana E. -G. Camus, 1913, I:88. 

Chusquea ramosissima Pilger, 1905: 145, not Lindman 1900:24. 

Chusquea sandiensis Pilger, 1920:29. 

58. Chusquea picta Pilger, 1905:151. 

59. Chusquea pinifolia (Nees) Nees, 1834:490. 

Arundinaria pinifolia Nees, in Martius, 1829:525. 

.? Ludolfia pinifolia (Nees) A. Dietrich, 1833, II:25. 

Chusquea heterophylla Nees, 1834:488. 

Chusquea heterophylla var. elonguta Doell, in 

Martius, 1880:207. 

C h usq uea heterophy lla var. microphylla Doell, 

in Martius, 1880:207. 

Chusquea heterophylla var. squumosa Doell, 

in Martius, 1880:207. 

Chusquea pinifolia var. heterophylla (Nees) Hackel, 

in Wettstein, ed., 1908 [1906:21]. 

60. Chusquea pittieri Hackel, 1903b: 153-154. 

350-351. 
76 

Chusquea maurofernandeziana 

1892: 64, nomen nudum. 

C h usq uea maurof ernanderiana 

Camus, 1913, I:86, pl. 56c. 

Hackel ex Pittier, 

Hackel ex E.-G. 

61. Chusquea polyclados Pilger, 1905: 147. 

62. Chusquea pubescens Steudel, 1854:337. 

63. Chusquea pubispicula Pilger, 1905: 148-149. 

64. Chusquea purdieana Munro, 1868:56. 

65. Chusquea quila Kunth, 1830:329-330, pl. 77. 

Nastus prolifer Desvaux, 1831:211. 

Coliquea quila Steudel, in Bibra, 1853: 115. 

Chusquea quila var. laxiflora Desvaux, in Gay, 

Chusquea intermedia Steudel, in Lechler, 1857:52, 

Chusquea quila var. longipila E.-G. Camus, 1913, 

66. Chusquea ramosissima Lindman, 1900:24. 

C h usquea phacellophora Pilger, 1923:456. 

67. “Chusquea” rollotii Berry, 1929: 2-3. 

This is a fossil species described from fragmentary 

remains in lithified gray shale of the late Tertiary 

(Colombia). 

68. Chusquea scandens Kunth, 1822:254 (Figures 

Nastus chusque Humboldt, Bonpland, and Kunth, 

Bambos chusque Poiret, 1817:494. 

Chusquea jamesonii Steudel, 1854:337. 

Chusquea quitensis Hackel, in Sodiro, 1889:484, 

Chusquea quitensis Hackel, 1903b: 154. 

Chusquea quitensis var. patentissima Hackel, 

Chusquea meyeriana var. patentissima (Hackel) 

1854:447. 

nomen nudum. 

I:198, fig. 61a, nomen nudum. 

30, 31 A-M). 

1816, I:201. 

nomen nudum. 

1908:161. 

69. 

70. 

71. 

72. 

E.-G. Camus, 1913, 1:94. 

Chusquea sclerophylla Doell, in Martius, 1880: 

200. 

Chusquea sellowii Ruprecht, 1839 [1840:125, 

pl. 11: fig. 261. 

Chusquea serrulata Pilger, 1898: 71 9-720. 

Chusquea simpliciflora Munro, 1868:54. 

Chusquea simplicifolia Munro ex Hemsley, in 

Godman and Salvin, ed., 1885:587, error for 

C. simpliciflora Munro. 

73. Chusquea sneidernii Asplund, 1939:797-799, 

fig. 2. 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

74. Chusquea spadicea Pilger, 1899:35. 

75. Chusquea spencei Ernst, 1872:262. 

76. Chusquea spinosa Fournier, 1881: 131. 

77. Chusquea straminea Pilger, 1905: 147-148. 

78. Chusquea sulcata Swallen, 1940:209. 

79. Chusquea swallenii McClure and Smith, in 

Reitz, ed., 1967:44-45, 48, figs. a-c. 

80. Chusquea tarmensis Pilger, 1905:151. 

81. Chusquea tenella Nees, 1834:492-493. 

82. Chusquea tenella var. latifolia Dutra, 1938: 146. 

83. Chusquea tenuiflora R. A. Philippi, 1859:206. 

Chusquea ciliata R. A. Philippi, 1864:299. 

84. Chusquea tenuiglumis Doell, in Martius, 1880: 

199-200. 

85. Chusquea tenuiglumis var. laxiuscula Doell, in 

Martius, 1880:ZOO. 

86. Chusquea tenuiglumis var. subcytindrica Doell, 

in Martius, 1880: 199-200. 

87. Chusquea tonduzii Hackel, 1903b: 155. 

88. Chusquea tuberculosa Swallen, 1931: 14. 

Chusquea hispida McClure, 1942: 179, fig. 7. 

89. 

90. 

91. 

92. 

93. 

94. 

95. 

96. 

1. 

2. 

3. 

4. 

5. 

6. 

7. 

8. 

Chusquea uliginosa R. A. Philippi, 1859:207. 

Chusquea unifiora Steudel, 1854:337. 

Chusquea urelytra Hackel, 1903b: 158. 

Chusquea uruguayensis Arechavaleta, 1897: 

546-547. 

Chusquea valdiviensis E. Desvaux, in Gay, 

1854: 446. 

Chusquea virgata Hackel, 1903b: 156-157. 

Chusquea ? wettsteinii Hackel, in Wettstein, 

ed., 1908 [1906:21-221. 

Chusquea wilkesii Munro, 1868:63. 

Checklist of 

Species Here Excluded from 

the Genus Chusquea 

Chusquea amplopaniculata Steudel (1854:337) 

is Dinochloa scandens (Blume) 0. Kuntze. 

Chusquea aristata. See Neurolepis aristata. 

Chusquea depauperata. See Swallenochloa deChusquea fimbriata. See Athroostachys capitata. 

Chusquea glomerata Munro (1868:50) (as synonym of Merostachys capitata Hooker). See 

Athroostachys capitata. 

Chusquea humilis. See Swallenochloa spicata. 

Chusquea simplissima. See Swal1en.ochloa spiChusquea spicata. See Swallenochloa spicata. 

pauperata. 

cata. 
NUMBER 9 77 

9. Chusquea subtessellata. See Swallenochloa sub10. Chusquea tessezlata. See Swallenochloa tessel11. Chusquea venezuelae. See Arthrostylidium 

12. Chusquea weberbaueri. See Swallenochloa 

t esse 1 la ta. 

lata. 

venezuelae. 

weberbaueri. 

Invalid Species of Chusquea 

1. Chusquea caamanoi Sodiro (1881:ll). Nomen 

nudum. 

Colanthelia McClure and E. W. Smith, new genus 

FIGURE 32 

Plantae unicespitosae, inermes. Rhizomata pachymorpha. Culmi habitu staturaque late diversi, aut 

parum alti et erecti vel “decumbentes” (teste Hackelii sub Arundinaria rhizantha) aut “altissimi scandentes” (teste Trinii sub Arundinaria distans), 

nodis valde elevatis et anguste cristatis, nodo medio 

unoquoque juventute gemmam solitariam gerenti, 

prophyllo gemmae et gemma ipsa in germinatione 

simul elongatis. Ramorum complementum ad nodos 

culmorum medianos axem ordinis primarii solitarium continens, idem basi vulgo mox proliferatum, ramis ordinis superioris axem primarium 

plus minusve valde superantibus. Vaginae internodia culmorum foventes basi cingulis latis post 

delapsum vaginarum persistentibus praeditae. Foliorum laminae venulae transversae extra vulga haud 

manifestae. 

Inflorescentiae semelauctantes, in speciebus plerisque vel infirme paniculatae vel racemosae, formis 

intermediis variis in ipsa planta interdum praesentibus; prophylla omnino haud ulla, bracteis 

dempta infima vel parvis vel obsoletis. 

Glumae transitionales vulgo 2, rarissime vel 1 

vel 3. Spiculae raro pauciflorae (ut in Colanthelia 

lanciflom) pleraeque pluriflorae, angustissimae et 

fragilissimae, apice in anthecio sterili plus minusve 

valde depauperato terminantes. Lemma fertile in 

maturitate paleam suam basi tantum circumplectans. Palea dorso 2-carinata et late sulcata, marginibus vix vel haud imbricatis. Rachillae segmenta 

ob fragilitas articulorum (nodorum) suorum in 

maturitate facilissime disarticulantia. Lodiculae 

typice 3, duae anticae plus minusve asymmetricae 

atque geminatae, postica symmetrica et vulgo 

minore. Stamina 3, filamentis filiformibus liberisque. Stigmata 2. Fructus non adhuc suppetens. 

Plants unicespitose, unarmed. Rhizomes pachymorph. Culms of small to medium stature and erect 

or decumbent, to very tall and scandent, the nodes 

prominent with a narrow crest, each midculm node 

giving rise to but a single initial bud; the prophyllum of the bud growing while the bud germinates, 

the primordium producing a single (segmented, 

terete) primary axis, this dominant over the axes 

of secondary order usually proliferating promptly 

from buds typically present at its own proximal 

nodes. Sheath at midculm nodes provided with (and 

abscising from) a conspicuous persistent girdle 

(basal expansion zone). Leaf blades with transverse veinlets as a rule not manifest externally. 

Inflorescences semelauctant, either paniculate or 

racemose (sometimes showing in the same specimen 

forms intermediate or intermixed between these 

two forms), part or all of the inflorescences (or 

even whole flowering branches) in a given specimen 

sometimes reduced individually to barely more 

than a solitary spikelet. Transitional glumes typically 2, rarely 1 or 3. Spikelets usually pedicellate, 

typically many-flowered (few-flowered in Colanthelia lanciflora), very narrow and fragile, terminating apically in a more or less strongly depauperate 

sterile anthecium. Fertile lemma fully embracing its 

palea only basally at maturity. Palea 2-keeled and 

broadly sulcate dorsally, gaping ventrally. Rachilla 

segments (owing to their fragility at the nodes) 

easily disarticulating at maturity. Lodicules typically 3, the anterior 2 more or less asymmetrical 

and paired, the posterior one symmetrical and 

usually smaller. Stamens 3, the filaments filiform 

and free. Stigmas 2. Fruit not yet available. 

ETYMoLoGY.-The name Colanthelia (f.) is coined 

from the Greek kolos, shortened, and anthele, 

“plume or panicle of a reed” (R. W. Brown, 1954). 

It alludes to a tendency common to all of the 

currently recognized species of the genus. There 

is a noticeably to strongly manifested tendency 

toward a progressive reduction of the reproductive 

structures from weak panicles to simple racemes 

and, in some cases, even from whole flowering 

branches all the way to little more than solitary 

spikelets. 
78 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 9 79 

FIGURE 32.-CoZanthelia cingulata (McClure and L. B. Smith) 

McClure. A, Rhizome and culm bases, x 0.6; B, branch complement from midculm range, in flowering state, x 1.2; 

c, culm sheath from above midculm range, with developing 

branch complement, x 0.9; D, apex of leaf sheath and base 

of leaf blade, x 7.2; E, paniculate inflorescence terminating 

a leafy flowering branch, x 0.6; F, progressively depauperate 

leafy and leafless flowering branches, x 0.6; G, flowering 

branch terminating in an inflorescence with mixed paniculate and racemose branching, x 0.6; H, spikelet with its 

pulvinate pedicel, x 2.4; I, transitional glumes, x 6; J, 

floret, x 6; K, lumma, x 6; L, palea, x 6; M, lodicule complement, x 12; N, stamen, x 6; 0, gynoecium, x 12. Drawings A-E, G-o based on Reitr and KEein 9679 (US), and F on 

Duse'n 18011 (US). 

TYPE-sPEcIEs.-Colanthezia cingulata (McClure 

and L. B. Smith) McClure. 

RELATIONSHIPS.-Affinity toward Aulonemia is 

weakly suggested by the nature of the midculm 

branch complement; this possible affinity is reinforced by the occurrence of paniculate inflorescences in some species. Among available examples, 

a specimen of Aulonemia haenkei (US 1256334) 

from Peru shows an occasional flowering branch 

reduced to little more than a solitary spikelet. The 

feature to which the name Colanthelia alludes also 

comes to light here and there in members of other 

genera. Among bamboos of the Old World genera 

a few similarly reduced flowering branches sometimes appear in specimens of Chimonobambusa 

marmorea. Other attributes (e.g., racemose inflorescences) suggest affinity toward Arthrostylidium. 

Such divergent attributes are connected by intermediate expressions to form clines, both from one 

species to another and (in some cases) within the 

same specimen. In the available material representing the genus Colanthelia, the morphological 

gamut of the reproductive apparatus reaches from 

leafy to leafless branches, each terminating in either 

a panicle, a raceme, a combination between those 

two forms, or a single spikelet. Pertinence to the 

genus Colanthelia (among the known bamboo 

genera of the New World) is made plain in its 

known members (even in their vegetative state) by 

the overall delicacy of the plant, with distinctively 

small leaf blades, combined with pachymorph 

rhizomes, prominent narrow-crested midculm 

nodes, a wide girdle at the base of the sheath at 

each node, and the unarmed complements with 

the primary element clearly dominant. With the 

exception of Colanthelia lanciflora, all known members of the genus have more or less distinctively 

long and narrow spikelets with more or less clearly 

exposed slender, elongate segments. 

DISTRIBUTION.-AS far as their natural distribution is a matter of record, all of the known species 

of Colanthelia are confined to Brazil. Three species have been reported from Santa Catarina, and 

one each from Minas Gerais, Paran$ Rio Grande 

do Sul, and S5o Paulo. The recorded altitudinal 

range extends from 30 m for Colanthelia cingulata, 

to 550 m for C. intermedia, to 710 m for C. gracillima, and to 1600 m for C. lanciflora. 

Annotated Checklist of 

Recognized Species of the Genus CoZantheZia 

1. Colanthelia burchellii (Munro) McClure, new 

combination. 

Arthrostylidium burchellii Munro, 1868:43. 

Arundinaria burchellii (Munro) Hackel, 1903a:69. 

2. Colanthelia cingulata (McClure and L. B. 

Smith) McClure, new combination (Figure 32). 

Aulonemia cingulata McClure and L. B. Smith, in 

Reitz, ed., 1967:50. 

3. Colanthelia distans (Trinius) McClure, new 

combination. 

Arundinaria distans Trinius, 1835:621. 

Trinius (1836, III:622) states, incorrectly, that a 

terminal rudiment is lacking in Arundinaria distans 

Trinius, q.v. 

4. Colanthelia intermedia (McClure and L. B. 

Aulonemia intermedia McClure and L. B. Smith, 

in Reitz, ed., 1967:52. 

5. Colanthelia lanciflora (McClure and L. B. 

Aulonemia lanciflora McClure and L. B. Smith, in 

Reitz, ed., 1967:47. 

6. Colanthelia macrostachya (Nees) McClure, new 

combination. Arundinaria macrostachya Nees, 

1834:48 1. 

7. Colanthelia rhizantha (Hackel) McClure, new 

combination. Arundinaria rhizantha Hackel, 

1909a: 323. 

Smith) McClure, new combination. 

Smith) McClure, new combination. 

Elytrostachys McClure 

FIGURES 33-35 

Elytrostachys McClure, 1942: 173, figs. 4-6; 1957:ZOZ. 

Plants unicespitose; unarmed. Rhizomes pachymorph. Culms self-supporting below, typically weak 
80 SMMITHSONIAN CONTRIBUTIONS TO BOTANY 

and pendulous or clambering above; the internodes 

cylindrical, hollow. Branch buds at culm nodes 

solitary, each containing but a single initial primordium. Branch complements at midculm nodes 

of either restricted or gremial insertion, unrestricted monoclade, consisting typically of one 

strong, central axis and a tuft of more slender ones 

arising from lateral buds at proximal nodes of the 

initial primordium, these latter buds sometimes 

germinating precociously while the apical growth 

of the initial primordium remains inhibited. Leaves 

(leaf sheath blades) typically with transverse veinlets not at all or only weakly manifest externally; 

oral setae few, rigid, erect or spreading, slender 

except at the bulbous base, scabrous. 

Inflorescences iterauctant, terminating leafy or 

leafless axes, of diffuse form, each axis the bracteate 

and prophyllate rachis of a pseudospikelet, the terminal segment of each rachis serving as the pedicel 

of a spikelet. Transitional glumes at the base of 

each spikelet none. [The conventional position of 

the transitional glumes is here (and in Atractantha, q. v.) occupied by what I now classify as bracts. 

When, in 1942, I first described and illustrated the 

genus Elytrostachys, I identified and labeled these 

structures as being glumes. However, in the perspective achieved more recently through a re-study 

of the structures in this transitional zone in the 

inflorescence of species of Elytrostachys in comparison with their counterparts in species of A tractantha, I now see them in both genera as empty bracts 

that are terminal to the series of bracts that subtend buds, rather than as empty glumes that precede the lemmas. The elongated terminal segment 

of the rachis that follows the two short internodes 

on which the emtpy bracts are inserted serves as the 

pedicel of the spikelet that is terminal to it. Having noted this, we can confirm. the lack of empty 

glumes by recalling that the natural loci of their 

insertion would be found immediately preceding, 

and close to, that of the first lemma.] Spikelets 

each made pedicellate by the segment of the rachis 

it terminates, and each typically containing but a 

single perfect flower (sometimes 2 in E. czavigera), 

promptly disarticulating at maturity immediately 

below the locus of insertion of a fertile lemma, the 

rachilla prolonged behind the terminal perfect palea in a bristle-like segment bearing a minute rudiment of a sterile anthecium. [When the spikelet 

contains but a single perfect floret, the prolongation of the rachilla is bristle-like and bears a minute 

rudiment; but when the spikelet contains more 

than one perfect floret, the prolongation of the 

rachilla behind the palea of the terminal perfect 

floret may be more robust than bristle-like, and 

then will bear a sterile anthecium more substantial 

than rudimentary.] Lemma in functional florets 

fully embracing its palea only basally at maturity. 

Palea gaping antically, dorsally canaliculate in 

spikelets containing but a single perfect floret, 

broadly sulcate in the lower floret when the spikelet 

contains two perfect florets. Lodicules 3, subequal 

or unequal, the anterior 2 asymmetrical and paired, 

the posterior one symmetrical and smaller. Stamens 

6, the filaments filiform, free. Stigmatic branches 

2. Fruit a fusiform or lagenoid, rostrate, sulcate 

caryopsis, the pericarp coriaceous, of even thickness 

below, thickened at the apex, the basal position of 

the embryo clearly manifest. 

ETYMoLoGY.-The name Elytrostachys, formed 

from the Greek elytro (combining form of eZytron) 

sheath, cover (elytra = wingcover of beetles) and 

stachys, spike, alludes to a fancied resemblance of 

the lemmas to the wing-cover of a beetle. 

TYPE-sPECIES.-E~ytrostachys typica McClure. 

RELATIONSHIPS.-The only known New World 

genus to which Elytrostachys appears to be closely 

related is Atractantha. These two genera share the 

following features: inflorescences of a peculiar form 

characterized by rachises with long terminal segments, each of which serves as the pedicel of an 

abscissile spikelet; the absence of the transitional 

glumes commonly found at the base of each spikelet, and the presence, in their stead, of a bract inserted on each of the two short internodes that 

precede the elongated terminal segment of each 

rachis; each spikelet typically containing but a 

single perfect floret followed by a rudiment of a 

sterile anthecium borne on the tip of a bristle-like 

prolongation of the rachilla. From known members of Atractantha, known members of Elytrostachys differ in the following features: midculm 

FIGURE SJ.--Elytrostuchys clavigera McClure. A, Portion of a 

seedling plant, x 0.6; B, culm sheath at midculm node of 

young culm shoot, x 0.6; c, midculm node and basal part 

of a typical branch complement, x 0.6; D, leafy twig with 

terminal inflorescence, x 0.6; E, two nodes from upper 

part of leafless flowering culm, x 0.6. All drawings based 

on McClure 21478 (US), 

J 

D 

0 

B 

~ 
NUMBER 9 83 

FIGURE 34.-Elytrostachys clauigera McClure. A, Leafy twig 

with pseudospikelet after the leaf blades and the first spikelet 

have fallen away, x 1.5; B, diagrammatic array of the parts 

of a flowering branch, showing pseudospikelets in various 

stages of development; c, pseudospikelet with the terminal 

spikelet mature and disarticulated, x 1.5; D, laminiferous 

bract (blade lacking) that subtended a young pseudospikelet, 

x 1.5; E, young pseudospikelet; F, transitional glumes, x 1.5; 

G, pedicel of the one-flowered spikelet, x 1.5; H, diagram 

of cross-section of a floret; I, lemma, lateral aspect, x 1.5; 

J. palea, lateral aspect, x 1.5; K, lodicule complement, x 3; 

L, stamen, x 3; M, gynoecium, x 3; N, mature fruit, hilum 

side showing sulcus, x 1.5; 0, mature fruit, embryo side 

showing embryotegium, x 1.5. All drawings based on 

McClure 21478 (US). For detailed caption for B, see McClure. 

1942:178, fig. 6. 

branch complements of restricted or gremial insertion, each arising from a solitary branch bud containing but a single initial primordium; auricles 

and oral setae of leaf sheaths not only much more 

conspicuously developed, but of different morphological configuration; in the presence of at least two 

empty bracts preceding the elongated terminal segment of each rachis; in the loosely convolute, more 

or less inflated form of the anthecia; and in the 

presence of twice as many stamens in each flower. 

As far as known to date, the respective geographical 

ranges of the two genera are widely disjunct. 

DISTRIBUTION.-of the two recognized species, E. 

typica has been reported only from the borders of 

a forest at the type-locality, El Limbn, Venezuela. 

Elytrostachys clauigera McClure has been collected 

in Colombia and all countries northward to Honduras, and from situations at reported elevations 

from 200 to 1500 m, described as river banks, old 

river terraces, wet ravines, crest of range, edge of 

forest, opening in forest. Both species are represented as flowering and fruiting freely. 

Key to the Species of Elytrostachys 

la. Leaf blades scabrous toward the base on the upper surface; laminiferous bracts silkypubescent in part; bladesless empty bracts (originally called empty glumes) I to 14 mm, 

11 to 18 mm long; terminal segment of rachis to 17 mm long and pubescent in part; 

lemma to 17 mm long; ovary hispidulous toward apex ........................................ 1. E. clavigera 

lb. Leaf blades glabrous on both surfaces; laminiferous bracts glabrous; bladeless empty 

bracts (originally called empty glumes) I to 15 mm, II to 30 mm long; terminal segment 

of the rachis to 6.5 mm long and entirely glabrous; lemma to 23 mm long; ovary entirely 

glabrous ...................................................... 

Glaziophyton Franchet 

FIGURE 36 

Gl~ziophyt~n Franchet, 1889:277.-McClure, 1957203. 

Arundinaria sect. Glaziophyton (Franchet) Hackel, in Engler 

and Prantl, 1897:46.-Lamson-Scribner, 1899:Q. 

Plants multicespitose, unarmed. Rhizomes pachymorph, the neck of an occasional one much elongated, thus creating an opening between tufts of 

culms arising either by tillering, or from a cluster 

of rhizomes with short necks. Culms strictly erect, 

the. internodes (in leafless, juncoid culms) cylindrical, the hollow interior partitioned by numerous 

thin but firm septa of pith without vascular tissue, 

the first above-ground internode disproportionately 

elongate, the upper ones very short to obsolete; 

juncoid culms in the vegetative state normally 

branchless and leafless-only after burning (teste 

Glaziou) do small secondary culms arise by tillering 

from the stumps and emit leafy branches. Branch 

buds at the nodes of these secondary culms solitary. 

....... ........................................................ 2. E. t~pica 

Vegetative branch complements of restricted insertion, unrestricted or facultatively restricted monoclade, with the median (primary) one dominant 

(those present in the only extant leafy specimens, 

Glaziou 8999 and 17914, not fully developed). Leaf 

sheaths with auricles and oral setae weakly developed or obsolete. Leaf blades with transverse veinlets manifest to superficial view, more prominently 

so on the abaxial surface. Culms in the flowering 

state leafless; the very slender, elongate, subequal, 

highly ramified flowering barnches borne in basally 

appressed pleioclade complements each arising from 

a solitary bud and inserted at one of the close-set 

distal nodes of the culm and producing gigantic, 

nodding, apically pendulous inflorescences (up to 

1 m long, teste Franchet). 

Inflorescences semelauctant, effusely branched 

panicles with a deliquescent rachis, the branches of 

all orders solitary, very slender and wiry, each subtended by a bract, and each bearing a 2-keeled 
84 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 9 85 

FIGURE 35.-Elytrostachys clauigera McClure. A, Midculm 

branch bud in early stage of germination, x 0.6; Ba, basal 

portion of young midculm branch complement, x 0.6; 

eb, basal portion of mature branch complement at upper 

node of culm, x 0.6; c, branch complements at upper nodes 

of culm in early stage of development, x 0.6; D, middle 

portion of sterile leafy first-order axis of midculm branch 

complement, x 0.6. All drawings based on McCEure 21478 

(US) * 

prophyllum at its basal node. Transitional glumes 

at the base of the spikelet 0-3; I and 11 “empty 

glumes”; III (when present) a “sterile lemma” that 

is either empty or subtends a rudimentary flower. 

Spikelets petiolate, each containing 2 (1-3) perfect 

flowers and terminating in a small, loosely convolute sterile anthecium (an occasional spikelet 

contains several progressively depauperate florets 

above the perfect ones); the rachilla segments somewhat elongate, potentially disarticulating immediately below the locus of insertion of each lemma 

including the terminal, reduced one (s). Palea gaping antically, dorsally sulcate and 2-keeled. Lodicules 3, the anterior two asymmetrical and paired, 

the posterior one symmetrical and smaller. Stamens 

3, the filaments filiform, free. Stigmas 2. Fruit 

unknown. 

ETYMoLoGY.-The name of this bizarre, monotypic genus honors its discoverer, Auguste Fransois 

Marie Glaziou (1833-1906), a French botanist and 

plant collector, who made excellent specimens and 

critical observations of this plant in the field during 

a period of 15 years. Glaziou passed on his observations to Franchet, apparently by oral communication. 

TYPE-sPEcIEs.-Glaziophyton mirabile Franchet, 

the sole known representative of the genus. 

World bamboos endemic on mountaintops in New 

Caledonia) resembles Glaziophyton superficially in 

the branching habit of the inflorescence (with each 

branch of the inflorescence subtended by a bract 

and bearing a prophyllum at its basal node) and in 

the manifest tessellate venation of the leaf blades, 

but differs profoundly in having the flowering 

cuhs leafy, with efistular internodes, and in the 

presence of 6 stamens and 3 stigmas in each flower. 

The disproportionate elongation of a single lower 

internode of every culm-a feature of the typespecies of Glaziophyton-is a vegetative character 

shared by most species (including the type) of 

RELATIONSHIPS.-GreSlania Balansa (genus of Old 

Myriocladus and several species of Arthrostylidium 

(e.g., A. schomburgkii). The peculiar, juncoid nature of the septate lumina of the culm internodes 

in the type-species of Glaziophyton apparently has 

no close counterpart in any known species of bamboo. Glaziophyton shows spikelet characters similar to those of Myriocladus, and features within the 

florets that link it in a general way with Merostachys, Myriocladus, and Arthrostylidium. The 

genus Glaziophyton, however, stands without any 

obviously close affinities among the bamboos. 

Glaziou assumed the existence of juncoid affinities in this plant, and on the field label of his first 

collection (Glaziou 8999, 7 June 1877) he wrote 

a tentative (unpublished) binomial, placing it in 

the genus Juncus. Franchet, however, in his discussion of its relationships, compared Glaziophyton 

mirabile to cyperaceous plants, such as Scirpus lacustris and Cyperus articulatus. It is of interest to 

note that both Juncus eflusus Linnaeus and Glaziophyton mirabile Franchet have been recorded from 

the same geographical area and at approximately 

the same elevation, in Brazil. 

DISTRIBUTION.-EXtant specimens indicate an endemism restricting this species to open, dry mountain tops in the general vicinity of Rio de Janeiro. 

Glaziophyton mirabile poses interesting problems 

for the serious student of the genetic, as well as 

the physiological, bases of morphogenesis. During 

a period of 15 years, Glaziou observed repeatedly 

that the sterile culms were all typically juncoid, 

without branches and without foliage leaves. When 

the sterile culms were destroyed by burning, however, secondary culms or tillers arising from the 

stumps produced leafy branches. Flowering was observed on leafless juncoid culms only. 

We have here what appears to be an extremely 

exaggerated expression of an ontogenetic feature 

found in some other bamboos (e.g., Bambusa arundinacea) where culms in the flowering state 

regularly lack foliage leaves from the first. In 

Glaziophyton mirabile, however, the phenetic gap 

between the flowering state and the vegetative state 

of the plant is very much more extreme morphologically, more persistent, and apparently deeprooted genetically. The only ecological condition 

that is known to trigger the production of shoots 

bearing leafy ramifications is a bizarre one, namely, 

burning of the aerial part of plants previously re- 
86 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 9 87 

FIGURE 36.--Glaziophyton rnirabile Franchet. A, Habit of 

underground part of plant: rhizomes with elongated necks 

(shown in broken outline), double circles represent the 

stumps of severed culms, greatly reduced; B, sterile leafless 

culm (partly in longitudinal section, to show pith diaphragms) and two young shoots, diameter x 0.6, length 

x ca. 0.9; c, tlp of leafless sterile culm, showing close-set 

budless nodes, x ca. 3; D, E, culm sheaths (apical portion 

only) from nodes 8 and 13 of sterile culm, abaxial and 

adaxial aspects, respectively, x 4.8; F, stumps of burned 

leafless plant, with leafy culms that developed by tillering, 

after burning, x 0.6; c, foliage leaf and part of its sheath, 

abaxial aspect, x 1.8; H, junction of foliage leaf and its 

sheath, adaxial aspect, x 1.8; I, portions of two mature 

flowering branch complements, the latter with an intact 

subtending sheath, x 1.2; J, bract subtending a middle-order 

branch of the inflorescence, x 6; K, prophyllum of a 

middle-order branch of the inflorescence, x 9; L, diagram of 

the structure, in longitudinal section, of the ultimate-order 

branching of the inflorescence. Each branch terminates in 

a spikelet. The axis of the flower proper is represented by 

a narrow cone, bearing the prophyllum-like palea; hl, spikelet, showing extreme development in length and in number 

of functional florets. Spikelets with 2 or 3 functional florecs 

are typical, x ca. 2.4; N, transitional glumes, x ca. 12; 0, 

floret, x 9; I>, diagram of floret Structure in cross-section; Q, 

lemma, x 9; R, palea, x 9; s, lodicule complement, x 9: 

T, stamen, x 9; IJ, gynoecium, x 15; v, branch of stigma, 

x ca. 120. Drawings A, B based on Glaziou 8999 (A based on 

Glaziou’s notes incorporated in Franchet’s description of the 

plant), C-E on Glaziou 11383 (P) , and F-v on Glaziou I7914 

(P). 

maining in the unbranched, perennially leafless 

state. How long will the “satiable curiosity” of 

technically equipped persons ignore the challenge 

to investigate this enigma-in depth! 

Merostachys Sprengel 

FIGURE 37 

Merostachys Sprengel, 1825, 11: 132.-Endlicher, 1836-1840: 

102.-Ruprecht, 1839:36.--Munro, 1868:46.-Doell, in Martius, 1880:207.-Bentham, in Bentham and Hooker, 1883: 

1209.-Hackel, in Engler and Prantl, 1887:93.-McClure, 

19573205. 

Plants unicespitose, unarmed. Rhizomes pachymorph. Culms self-supporting in the lower part, 

the slender, sometimes long-attenuate upper part 

nodding, pendulous or clambering; internodes 

cylindrical, typically hollow, exceptionally, as in 

some as yet unidentified species, efistulose or filled 

with pith. Primary branch buds solitary at midculm nodes, each bud giving rise to but a single 

initial (primary) axis. Branch complement at midBradocalumus Nakai, 1933: 10.-McClure, 19573201. 

culm nodes monoclade, with its primary element 

solitary, flat, unsegmented, roughly ovate or cordate to triangular in contour, thin, adnate to the 

surface of the culm; secondary branches slender, 

subsequal, of apsidate insertion, arising from primordia formed on the margin of the primary element, and displayed in fan-shaped array. Leaves 

(blades of leaf sheaths) with transverse veinlets usually not manifest externally. 

Inflorescences semelauctant, spicate racemes terminating usually leafy twigs, the rachis excurrent, 

bearing spikelets of solitary (in a few species paired 

or ternate) insertion and secund orientation, the 

distal 1 or more spikelets typically depauperate or 

rudimentary; bracts subtending spikelets (excepting 

the Jowrrmost one) generally reduced, rudimentary, 

or obsolete. Transitional glumes usually 2 and 

very unequal, both usually empty, but one or the 

other sometimes subtending what appears to be a 

rudimentary bud which when present, is the lemma 

of a sterile floret. Spikelets sessile or subsessile, 

usually containing but a single perfect flower (in 

some species now and then 2-as in an isotype of 

Merostachys speciosa, teste Munro (1868:48)-in 

others regularly 2; rarely-as in M. pluriflora-2-5 

and exceptionally, as in M. polyantlza, even up to 

10) and terminating in a rudimentary or more or 

less depauperate sterile anthecium. Rachilla segments potentially disarticulating immediately below the locus of insertion of a fertile lemma. 

(When the spikelet contains but a single perfect 

floret, the prolongation of the rachilla is bristle-like 

and bears a minute rudiment; when the spikelet 

contains more than one perfect floret the prolongation of the rachilla behind the palea of the terminal 

perfect floret may be more robust than bristle-like, 

and then may bear a sterile anthecium more substantial than rudimentary.) Florets more or less 

strongly inflated, the lemma fully embracing the 

palea only basally at maturity, the palea gaping 

antically, narrowly to broadly sulcate and &keeled 

dorsally. Lodicules 3, in some species subequal and 

subsimilar, elsewhere the anterior 2 asymmetrical 

and paired, the posterior one smaller and sym. 

metrical. Stamens 3, the filaments filiform, free. 

Stigmas 2. Mature fruit a mucronate or weakly 

rostrate, oblong or ovoid caryopsis with a thin, 

coriaceous or crustaceous pericarp of even thickness 

or more or less strongly thickened toward the base 

and toward the apex; in the absence of both a sul- 
88 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

cus and an embryotegium, the respective positions 

of the hilum and the embryo are not manifest 

externally. 

ETYMoLoGY.-The name, Merostachys, formed 

from the Greek, meros, part (partial, or incomplete) and stachys, spike, alludes to the secund (onesided) profile of the inflorescence. 

TYPE-SPECIES.-Merostachys speciosa Sprengel. 

RELATIoNsHIPs.-RhipidOcladum is the only bamboo genus toward-whose members bamboos of the 

genus Merostachys show any strong or striking resemblance. This resemblance is shown most spectacularly by the manner of development and form 

of the branch complement at midculm nodes (Figures 37c, 42~). However, the form of the sheaths 

(Figures 37~, 42~) at midculm nodes of the young 

culms in active growth (insofar as these are known 

for plants of either genus) provides a convenient 

means of establishing the generic affinity of a given 

plant. Otherwise a deceptive technical resemblance 

between members of Merostachys and members of 

Rhipidocladum is manifested only where certain 

features of the reproddctive structures appear in 

exceptional combinations. The fact that in Merostachys the thickening of the pericarp of the fruit 

proceeds simultaneously from both apex and base 

may provide an additional item of difference between Merostachys and Rhipidocladum. Thus occasions for confusion or uncertainty as to generic 

affinity or generic discontinuity may involve those 

species of Rhipidocladum having pectinately secund, spicate inflorescences (e.g., R. maxonii) or 

those species of Merostachys (e.g., M. pluriflora and 

M. polyantha) having spikelets that contain several 

perfect florets and terminate in a robust (not bristle-like) rachilla segment bearing a sterile anthecium more substantial than rudimentary. Specimens 

of such mutually imitative members of Merostachys 

and Rhipidocladum, however, may be sorted to 

genus if, in addition to the midculm branch complement, they both contain intact or complete examples (in good condition) of sheaths from 

midculm nodes. 

I do not find any support for the opinion expressed by Hackel (1909a:327) that Merostachys is 

related to Chusquea. 

DISTRIBuTIoN.-The 25 species of Merostachys 

tentatively recognized here give the genus an aggregate but discontinuous distribution extending 

from Argentina to Guatemala and British Honduras, at moderate elevations and up to 1500 meters. 

As far as known today, the genus reaches its fullest 

development and greatest diversity in Brazil, where 

20 described species are found. Outlying stands of 

single species have been reported from Argentina, 

Uruguay, Paraguay, Peru, Guatemala, and Honduras; two species have been recognized in Venezuela. 

Annotated Checklist of 

Species of the Genus Merostachys 

1. illerostachys anomala Dutra, 1938:151, fig. 3. 

[See Merostachys multiramea.] 

2. Merostachys argyronema Lindman, 1900:22, pl. 

15. 

The Guatemalan plant referred to this species by 

McClure (in Standley and Steyermark, ed., 1955: 

207) is known only in the vegetative state. It probably represents a species distinct from M. argyronema, whose vegetative features are, as yet, insufficiently known. 

3. Merostachys bradei Pilger, 1927: 114. [See 

4. Merostachys brevispica Munro, 1868:49. 

5. Merostachys burchellii Munro, 1868:51. 

M erosta ch ys p lurifloru.] 

This species was initially described by Munro on 

FIGURE 3’i.-Merostachys sp. (A,o) and Merostachys speciosa 

Sprengel (B-N) . A, Seedling plant, basal part, showing 

caespitose clump habit and pachymorph rhizomes (tip 

of young culm shoot at left), x 1.2; B, culm sheath (apical 

portion only) from midculm node of mature culm, x 0.6; 

c, branch complement (base only of the component axes) at 

midculm node, x ca. 0.9; D, leaf sheath (apex only) showing 

ligule, base of oral setae, petiole, and base of leaf blade, 

x 4.8; E, flowering branch, terminating in a secund spicate 

inflorescence, x 0.6; F, single branch (a sessile spikelet) 

of the inflorescence, x 2.4; G, diagram in longitudinal 

section of a spikelet, showing two transitional glumes followed by a segment of the rachilla, a bristle-like prolongation of the rachilla (terminated by a rudiment), the lemma, 

palea, and the floral axis - the last shown as a narrow 

cone; H, diagram in cross-section of the floret; I, lemma in 

two aspects, x ca. 3.1; J, palea in three aspects, and the 

bristle-like prolongation of the rachilla in two aspects, x ca. 3.1; K, lodicule complement, x 6; L, stamen, x 4.8; M, 

gynoecium, x 6; N, branch of a stigma, greatly enlarged; 

0, mature fruit (in two aspects, x 3.6) after it has produced a seedling plant (shown at A). Drawings A, o based 

on Swallen 8134 (US), B on Chase 9466 (US), c on Lofgren 

1653 (US), D-F on SelEow 1286 (US ex B) and DN on Reitz 

2268 (US). 
NUMBER 9 89 
90 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

the basis of a single collection consisting of sterile 

specimens under Burchell 3243 (K). Dutra (1938: 

151) described, under the same name, a flowering 

specimen purporting to supplement Munro’s image 

of that species without, however, citing any specimen to document his description. But, in his introductory statements, Dutra made reference to the 

flowering, in 1907, of a plant identified by him as 

Merostachys burchellii Munro. A flowering specimen collected in that same year by Dutra under 

no. 513 bears (at US) a tentative identification as 

Merostachys burchellii. It may represent the same 

collection as the uncited specimen on which Dutra 

based the (emended) description of M. burchellii 

referred to above. Judged by its vegetative features, however, Dutra 513 is very different from 

Burchell 3243, the type of Merostachys burchellii 

hlunro. It is closely related to (but not identical 

with) Dutra 518 (US), type-collection of Merostachys anomala Dutra (9. v.), collected in 1906 in 

the same general locality. 

6. Merostachys ciliata McClure and Smith, in 

7. Merostachys claussenii Munro, 1868:48. 

Reitz, 1967:71, pl. ~ZD-H. 

? Merostachys claussenii p mollior Doell, in MarEkman (1913:64) expressed the opinion that 

Doell’s type of 6 mollior is identical with Munro’s 

type of M. claussenii. Ekman supports this opinion 

by the assertion that Claussen 399 “is indeed a 

duplicate of the original plant,” i.e., is actually a 

duplicate from the Claussen collection cited by 

Munro. This assertion is neither verified nor supported by my observation that Munro’s specimen of 

“Claussen, Minas Geraes” (frag. ex K at US) is 

clearl:! distinguishable from Doell’s type (Regnell 

III n. 1425-frag. et photo ex S at US) on the basis 

of features of the spikelets alone. The discrepancy 

between Ekman’s interpretation and the way things 

look to me suggests the need for a re-evaluation of 

the evidence as to whether more than one recognizable taxon is involved here. 

8. Mcrostachys exserta Munro ex E. -G. Camus, 

This is a species of which I have not seen a specimen. The type (P) cited by Camus as “Auguste de 

Saint-Hilaire, 1816 B 1821; Cat. D, no. 713,” is annotated by Munro (teste Camus) with the words 

tius, 1880:214. 

1913, 1~74, pl. 44A. 

“Spica exserta ab affinibus distat; spicula brevi M. 

Fischerianae approximat.” 

9. Merostachys fischeriana Ruprecht ex Doell, in 

iMartius, 1880: 2 15. 

10. Merostachys fistulosa Doell, in Martius, 1880: 

209, pl. 55. 

11. Merostachys glauca McClure and Smith, in 

Reitz, 1967:74, pl. 12N. 

12. Meyostachys kunthii Ruprecht, 1839:37, pl. 10: 

fig. 30. 

Aderostachys speciosa sensu Kunth, 1830:333, pl. 79. 

[Non Sprengel, teste Ruprecht.] 

13. Meyostachys maguireorum McClure, in Maguire, Wurdack, et al., 1964:5. 

14. Merostachys multiramea Hackel, 1909a:326. 

Merostachys anomala Dutra, 1938: 151, fig. 3. 

15. Merostachys neesii Ruprecht, 1839:37, pl. 10: 

fig. 31. 

Merostachys speciosa sensu Nees, in Martius, 1829: 

527. “on Sprengel, teste Ruprecht.] 

16. Merostachys pauciflora [as pauciflorus] Swallen, 1943:469, fig. 1. 

17. Merostachys petiolata Doell, in Martius, 1880: 

216. 

18. Merostachys pluriflora Munro ex E. -G. 

Camus, 1913, I:77. 

Bambusa pubescens Doell, in Martius, 1880: 189, pl. 

51: E. -G. Camus, 1913, I:124. (Non Loddiges 

ex Lindley, 1835, III:357.) 

Biasilocalarnus pubescens (Doell) Nakai, 1933: 10. 

Merostachys hradei Pilger, 1927: 114. 

A catalog of the hitherto unassembled series of 

amazing and amusing taxonomic inadvertencies by 

which the history of this species has become complicated may prove instructive to the prospective 

student of the bamboos. Having projected “BamDztsa” as the proper repository for a specimen from 

Gaudichaud’s collection no. 97, Doell (in Martius, 

1880) proceeded (certainly by inference rather than 

on the basis of direct evidence) to include in the 

illustration of his Bambusa pubescens a floral diagram showing six stamens. When Doell’s otherwise 

authentic plate, and his description (including the 

statement “stamina 6”) caught the eye of Dr. Nakai, 

the latter saw what he took to be a new genus, and 

proceeded (Nakai 1933) to christen it Brasilocalamus. Not having examined a specimen of the type- 
NUMBER 9 91 

species, however, Nakai failed to detect Doell’s error 

in giving it six stamens. At an earlier (unrecorded) 

date, Munro had already annotated (at P) another 

specimen from the same Gaudichaud collection 

with the following words: “Merostachys sp. indescr. 

M. plurifloya nov. sp. M. Kunthii Rupr. affinis differt spiculae flosculis hermaphrod [itis] 3-5.” E. -G. 

Camus (1913, I:77) published Munro’s manuscript 

name, with a description consisting solely of the 

Latin annotation associated with it. Camus failed 

to take account of the fact that this name is based 

upon a specimen from the same collection that 

yielded the type of Bnmbusa pubescens Doell. Consequently, the latter name also appears in the Camus work (1913, I:124) along with a full 

reproduction of Doell’s description of it translated 

into French. Camus apparently overlooked the fact 

that Bambusa pubescens Doell (in Martius, 1880) 

is a later homonym of Bambusa pubescens Loddiges 

in Lindley (1835) which name appears (Camus, 

1913, I:152) in a list of synonyms under Dendrocalamus strictus. In 1927 Pilger described Merostachys bradei as new to science, on the basis of a 

specimen from a different collection of M. pluriflora. An English translation of Pilger’s notes in 

German reads as follows: 

The new species is distinguished by the short, thick, spikes, 

as well as by the 2-3 fully developed flowers with normal 

lemma and palea [in] short, thick, downy spikelets. E. -G. 

Camus (1913, I:77) mentions a M[erostachys] pluriflora 

Munro without a full description (‘les kpillets pluriflores’) ; 

but that species, which cannot be considered as properly 

published, can hardly be identical with my new species because that one is considered to be nearly related to M. 

Kunthii. The rudiments of glumes below the two empty 

glumes cited by Eichler in Flora Brasiliensis were not observed in our species. 

A formal description of the following species, 

apparently new to science, is included here because 

by adding a member in which the number of anthecia in its apically indeterminate spikelets sometimes reaches the unprecedented figure of ten, it 

dramatically extends the current circumscription of 

a genus whose type-species has one-flowered spikelets terminating in a bristle-like prolongation of the 

rachilla bearing a rudiment. 

19. Merostachys polyantha McClure, new species 

Culmi 6-8 m alti (teste A. Lima); internodia 

teretia, cava, parietibus tenuibus, extus infra unumquidque nodum in zona angusta pilis albidis adpressis confertim vestita, alibi papillis retrorse 

aculeiferis praedita, demum luteo-viridia et punctis 

vinosis minutis elongatisque omnino variegata. 

Culmorum vaginae deciduae, tum ex omnibus cingulo pilis albidis retrorse adpressis vestito remanenti; auriculis et setis oralibus valde evolutis; 

ligula perbrevis, margine irregulatim denticulata 

ciliolataque; lamina decidua (non adhuc visa). 

Foliorum vagina striato-sulcata, secus marginem 

externam setis lustratis ornata, vaginis infimis dorso 

et ut videtur omnino glabris, supremis papillis retrorse uncinatis punctatis; auriculis et setis oralibus 

circa ut in vaginis culmorum evolutis; ligula interiore haud exserta, exteriore anguste lineari, margine vel subglabra vel dense ciliolata; petiolo usque 

8 mm longo, compresso, vel omnino glabro vel 

utrinque dense scabro; lamina usque 150 x 35 mm, 

oblongo-lanceolata, basi asymmetrice vel rotundata 

vel cordata, apice asymmetrice et abrupte acuminata; laminis infimis et eis supremis in quoque 

axi florifero ovatis acuminatissimisque, omnibus 

utrinque concoloratis et fere glabris; nervis longituclinalibus omnibus in laminarum facie abaxiali 

tantum salientibus; costa tantum basin versus laminae distinguenda; nervibus secundariis utroque 5-8; 

tertiariis in quoque spatio costae contiguo usque 8, 

versus marginem laminae gradatim paucioribus ita 

in quoque spatio marginali saepius ad 2 redactis; 

venulis transversis in facie abaxiali tantum distinguendis, vicinis 1.5-4.0 mm sese distantibus. 

Inflorescentiae e racemis spicatis secundisque sistentes, solitariae, axes saepius laminiferos terminantes; pedunculo vel haud vel parum exserto, 

apice dense canescenti; rhachidi 25-65 mm longa, 

curvula, dense antrorse-pubescenti; bractea spiculam basilarem subtendenti ovata, apice acuta apiculataque, multinervia et valde carinata, dorso 

marginibusque omnino glabra, usque 10 mm longa, 

succedentibus gradatim redactis, distalibus vel rudimentariis vel obsoletis. Spiculae compressulae, 

anguste triangulae, apice indeterminatae, insertione solitariae, confertae, alternae, distichaeque, 

pleraeque 1.5-2.0 mm (infimae usque 4.0 mm) sese 

distantes; spiculis in quoque racemo usque 26, infima 55 mm longa, sequentibus gradatim brevioribus, tum distalibus 7-8 vel depauperatis vel 

rudimentariis; pedicel10 sub-nil, latere externo prominenter pulvinato; glumis transitionalibus saepius 
92 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

2, alia vel alia aliquando rudimentum (velut gemmam dormientem) foventi; I ca 4 mm longa, anguste triangula, acuminatissima, valde carinata, 

cetera enervosa, dorso invalide appresso-hirtula, 

seas margines valde ciliata; II 6-9 mm longa, ovata, 

acuminata, 5-nervia et valde carinata, extus valde 

hirtula, marginibus versus apicem tantum incalide 

ciliolata. Flosculi in spiculis proximalibus usque 

10, eis in spiculis succedentibus gradatim paucioribus; flosculo terminali vel plus minusve depauperato vel etiam in statu rudimentario dirempto. 

Anthoeciorum elementae utraeque crustaceae, extus 

plus minusve dense strigosae, sews margines apicem versus antrorse-ciliatae. Rhachillae segmenta 

usque 4mm longa, semiclaviformia, omnino molliter antrorse pubescentia. Lemma flosculi infimi 

usque 20 mm longum, flosculorum sequentium gradatim redactum, anguste lanceolatum, apicem verus 5-nervium naviculareque, alibi enervium. Palea 

vulgo vel lemmam aequans vel parum brevior, raro 

parum longior; oblongo-lanceolata, late sulcata, 

omnino enervia, apice et profunde et anguste bilobata. Lodiculae dimensionibus formaque variabilissimae, textura tenuissimae, deorsum interdum 

opacae, sursum semper translucidae, vel enerviae 

vel invalide paucinerviae, omnino glabrae vel margine (apice saltem) obscure ciliolatae. Antherae 

atropurpureae, versus basin apicemque parum 

attenuatae. Ovarium, stylus et rami styli infra stigmata omnino glabri. Fructus non adhuc inventus. 

Culms 6-8 m tall (teste A. Lima) the internodes 

terete, hollow, thin-walled, more or less heavily 

coated at first with white powder and densely 

clothed in a narrow zone immediately below each 

node with retrorsely appressed lustrous white hairs, 

elsewhere sparsely strewn with papillae each tipped 

with a minute retrorse hook; ultimately seen as 

yellowing green in color, variegated with longitudinally elongated vinaceous stipples. Culm 

sheaths deciduous, upon abscission leaving a rather 

prominent persistent girdle densely clothed outside 

and fringed basally with retrorsely appressed lustrous white hairs; the sheath proper ciliate on the 

outer margin, apparently entirely glabrous abaxially; auricles and oral setae prominently developed, 

the auricles narrow, tortuous, dull, and glabrous 

abaxially, densely hirtellous adaxially, the oral 

setae 10-12 mm long, spreading from crowded loci 

of insertion on the margin and on the adaxial surface of the auricles, lustrous, very sparsely and 

obscurely antrorse-scabrous throughout, crisped 

distally; ligule short, truncate or slightly arcuate 

apically, minutely and irregularly denticulate and 

ciliolate on the margin; blade deciduous (not seen). 

Leaf sheaths proper (represented on flowering twigs 

only in the available specimens) minutely corrugated (quasi-plicate) longitudinally, densely fringed 

with lustrous hairs on and near the outer margin; 

the proximal sheaths in a given series glabrous, the 

distal ones strewn (on the exposed surface at least) 

with retrorsely uncinate papillae; auricles and oral 

setae approximately as seen on the culm sheaths; 

inner ligule not exserted, the outer ligule narrowly 

linear, subglabrous to densely ciliolate on the margin; petiole up to 8 mm long, dorsiventrally 

compressed, entirely glabrous to densely antrorsehispidulous throughout on both surfaces; blade up 

to 150 x 35 mm, oblong-lanceolate, asymmetrical 

and broadly rounded or cordate at the base, asymmetrically and abruptly acuminate at the apex, 

abruptly becoming ovate-acuminate and greatly reduced in size at the proximal and distal extremes 

of every leafy flowering twig, concolorous and glabrous or nearly so on both surfaces; longitudinal 

nerves all salient on the abaxial surface only, the 

midrib distinguishable only toward the base of the 

blade; secondary nerves 5-8 on each side of the 

midrib; tertiary nerves up to 8 in the first space on 

each side of the midrib, the number in each successive internerve space reduced progressively, 

commonly 2 next to each margin of the blade; 

transverse veinlets visible only on the abaxial surface, adjacent ones in a given space 1.5 to 4.0 mm 

distant from each other. 

Inflorescences congested secund spicate racemes 

terminal to usually leafy axes; peduncle scarcely to 

slightly exserted, densely canescent apically; rachis 

25-65 mm long, lightly curved, densely strigose; 

bract subtending the lowermost spikelet ovate, 

acute and apiculate apically, dorsally many-nerved 

and prominently keeled, glabrous throughout on 

the back and on the margins, up to 10 mm long, 

those subtending successive spikelets progressively 

reduced, the distal ones either rudimentary or obsolete. Spikelets somewhat compressed, narrowly 

triangular, of indeterminate growth apically, of 

solitary but crowded, alternate and distichous insertion at intervals of 1.5 to 4.0 mm from the base 
NUMBER 9 93 

of the raceme upward in each row; spikelets in 

each raceme up to 26, the basal one up to 55 mm 

long, the successive ones progressively reduced in 

length to about 20 mm, followed by sometimes 

seven or eight abruptly and progressively depauperate to rudimentary ones; pedicels almost obsolete, each prominently pulvinate on the external 

side only; transitional glumes normally two, one or 

the other sometimes subtending a bud-like rudiment; I about 4 mm long, narrowly triangular, 

acuminate, prominently keeled, otherwise without 

visible nerves, weakly appressed hirtulous dorsally 

and strongly ciliate on the margins; 11 6-9 mm long, 

ovate, acuminate, &nerved and strongly keeled, 

hirtulous dorsally and weakly ciliolate on the margin toward the apex only. Florets in the lowermost 

spikelets up to 10, progressively fewer and shorter 

in succeeding spikelets, the terminal floret either 

more or less strongly depauperate or stalled at a 

rudimentary stage in its development. Elements of 

the anthecia both crustaceous, more or less densely 

strigose on the outer surface, antrorse-ciliolate on 

the margins apically. Rachilla segments up to 4 

mm long, semiclaviform, softly pubescent throughout with antrorse hairs. Lemma in the lowermost 

floret up to 20 mm long, progressively reduced in 

the succeeding florets, narrowly lanceolate, 5- 

nerved and strongly boat-shaped toward the apex, 

elsewhere without manifest nerves. Palea commonly as long as the lemma or a little shorter, 

rarely a little longer; oblong-lanceolate, broadly 

sulcate, lacking manifest nerves, deeply and narrowly bilobed at the apex. Lodicules of variable 

form and dimensions, of delicate texture, sometimes 

opaque at the base, always translucent above, either 

weakly few-nerved or nerveless, glabrous throughout or obscurely ciliolate on the margin toward 

the apex only. Anthers dark purple, lightly attenuate toward the base and at the apex. Ovary, 

style and branches of the style (below the stigmas 

at least) entirely glabrous. Fruit not yet found. 

COTYPES (= syntypes, sensu Frizzell 1933:647).- 

Herbarium Bradeanum no. 22857, leg. Flavia Torgo 

(s.n.) 19 viii 1926; and Herb. Kew, leg. A. Lima 

(= Dartlano de Andrade-Lima) no. 62-4122, 24 

viii 1962 (ex Instituto de Pesquisas Agronomicas 

Secao de Botanica, Sao Paulo, no. 13192), both

collections taken simultaneously from the same 

flowering plant (teste A. Lima): “Serra de Mongagui, pr. Praia Grande, Municipio Mongagua, Sao 

Paulo, Brasil, a sombra da mata no alto da serra 

(100-120 m). 6-8 m. F1. roxoescuro. Caule c/peq. 

aculeos cuticulares (?) volt. p/base.” Plants shaded 

by forest at the summit of a hill 100-120 m above 

sea level-culms 6-8 m tall, with minute cuticular 

hooks on the lower internodes; flowers dark red. 

[Vernacular name:] ‘taquara miju.’ ” Notes ex A. 

Lima. Both cited specimens seen. Photographs of 

both cotype specimens, and a duplicate of the specimen collected by A. Lima are on deposit at the 

U.S. National Herbarium. 

RELATIONSHIPS.-Merostachys polyantha differs 

from the closely related M. pluriflora in the more 

robust development of the inflorescences, spikelets, 

anthecia and the auricles, and oral setae; as well as 

in the greater maximum number (up to 26) of 

spikelets in each raceme; the greater maximum 

number (up to 10) of florets in each spikelet; the 

shape and vesture of the parts of the anthecia; and 

the much broader, oblong-lanceolate shape of the 

midrange leaf blades. 

DIsTR1suTIoN.-Merostuchys polyantha is known 

as yet by only two collections taken, simultaneously 

(teste A. Lima), from a single source: Serra de Mongag&, prope Praia Grande, Municipio SPo Paulo, 

Brasil. Its nearest known relative, Merostachys 

PlzirifZora, has been collected near Iguape, State of 

S5o Paulo, and on the Island of Santa Catarina, 

State of Santa Catarina. 

20. Merostachys retrorsa McClure, in Maguire, 

Wurdack, et al., 1964:6. 

21. Merostachys riedeliana Ruprecht ex Doell, in 

Martius, 1880: 2 13. 

22. Merostachys sellovii Munro, 1868:51. 

23. Merostachys sparsiflora Ruprecht, 1839:37, pl. 

10: fig. 32. 

24. Merostachys speciosa Sprengel, 1825:249 (Fig 

ure 37~-~). 

Under this name Munro (1868:48) writes ‘‘I find 

in one of Sellow’s specimens the second hermaphrodite flower which I had first observed in M. Clausseni.” 

25. Merostachys speciosa sensu Kunth, 1830: 333, 

334, pl. 79. [See Merostachys kunthii.] 

26. Merostachys speciosa sensu Nees, in Martius, 

1829:527-531. [See Merostachys neesii.] 
94 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

27. Merostachys ternata Nees, in Martius, 1829: 

28. Merostachys vestita McClure and Smith, in 

The following authors offer additional information on some of the species of Merostachys listed 

above: Doell, in Martius, 1880:207-218; McClure, 

in Swallen, 1955:207-209; McClure and Smith, in 

Reitz, ed., 1967:63-75; Metcalfe, 1960:564, 586-588; 

Munro, 1868:47-52; Nees, in Martius, 1829:527- 

531; Parodi, 1936:242-244; Ruprecht, 1839:36-38. 

529. 

Reitz, 1967:72, pl. 12 i-j. 

Annotated Checklist of Excluded Species 

Erroneously Allocated to the Genus Merostachys 

1. Merostachys capitata Hooker, 1840:273. [See 

A throostachys capitata.] 

Merostachys capitata latifolia Doell, in Martius, 

1880:2 17. 

Merostachys capitata @ angustifolia Doell, in Martius, 1880:217. 

2. Merostachys race.mipora (Steudel) Fournier, 

1881: 131. [See Rhipidocladum racemiflorum.] 

3. Merostachys sikokianus (Makino) Nakai, in 

Hara, 1935:74. [A Japanese plant of the genus 

Sedum-Crassulaceae.] 

Myriocladiis Swallen 

FIGURES 38-40 

Myriocladus Swallen, in Steyermark, et al., 1951:34, fig. 4; 

in Maguire, Wurdack, et. al., 1957:237-239.-McClure, 

195$:205. 

Plants, unicespitose, unarmed. Rhizomes pachymorph. Culms self-supporting, the internodes terete, in some species hollow, in others efistular, the 

lowermost above-ground one usually disproportionately elongated and followed by one to several more 

or less completely suppressed (obsolete) ones with 

the intervening nodes crowded closely together. 

Branch buds at culm nodes solitary. Branch complements restricted monoclade, the successive ones 

sometimes brought deceptively closely together by 

the suppression of culm internodes. The primary 

element of branch complements apparently never 

proliferating from any bud at its proximal nodes, 

this perhaps due to suppression of the buds by 

powerful physical pressure exerted by the very 

thick, rigid sheaths that subtend them. [A proximal 

internode of the single component branch is also 

(as in the culm) commonly elongate and followed 

by one to several suppressed (obsolete) ones with 

the intervening nodes crowded closely together, 

making the corresponding sheaths strongly imbricate.] Leaves (blades of leaf sheaths) petiolate or 

subsessile, the transverse veinlets usually obscure or 

invisible externally or (as in M. paludicolus) more 

or less conspiculously manifest on both surfaces. 

Inflorescences semelauctant, terminating leafy or 

leafless branches and (in some species) the culms as 

well; generally long and narrow, of diverse branching habit, racemose to paniculate, usually with part 

or all of the branches of secund orientation and 

lacking both prophylla and subtending bracts. 

Spikelets either pedicellate, subpedicellate or sessile, containing 2 (1-5) perfect florets, and terminating in a depauperate sterile anthecium. Rachilla 

segments short, potentially disarticulating immediately below the locus of insertion of the first only, 

or (in some species) each of the fertile lemmas. 

Transitional glumes typically 3: I and 11 “empty 

glumes,” 111 a “sterile lemma” that is either empty 

or subtends a rudimentary flower. Fertile lemma 

fully embracing its palea only basally at maturity, 

the palea broadly sulcate dorsally, the margins gaping basally and lightly overlapped above. Lodicules 

3, either subsimilar in shape or the anterior 2 more 

or less noticeably asymmetrical and paired, the 

posterior one symmetrical and slightly smaller. Stamens 3, the filaments filiform, free. Style very short, 

terminating in two stigmas. Mature fruit a mucronulate or weakly rostrate, fusiform caryopsis with a 

thin membranaceous pericarp of even thickness, the 

sulcus and the basal position of the embryo clearly 

manifest. 

ETYMoLoGY.-The name Myriocladus, formed 

from the Greek, myrios, numberless, and cludos, 

branch, ostensibly alludes to the profuse ramification of the inflorescences. 

TYPE-sPF.cIEs.-Myriocladus virgatus Swallen. 

RELATIONSHIPS.-Certain of the striking generic 

features of Myriocladus are shared by individual 

FICURF 38.-MyriocZadus confertus Swallen. Aa, Ab, Leafy 

flowering plant, x 0.6; B, seedling, x 1.2; c, first-order 

branch of inflorescence, x 1.2; D, spikelet, x ca. 9.6; E, 

apex of leaf sheath and base of blade, x 1.8; F, two nodes of 

culm with shortened internode and an intact branch bud, 

x ca. 6. All drawings based on Steyermark and Wurdack 

1220 (US). 

96 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 9 97 

FIGURE 39.-Myriocladus virgatus Swallen (Aa-AC) and hlyriocladus confertzts Swallen (B). Aa, Leafy tip of sterile culm, 

x 0.6; Ab, apex of leaf sheath and base of blade (abaxial 

aspect), x 1.2; AC, apex of leaf sheath and base of blade 

(adaxial aspect), x 1.2; B, leafy tip of sterile CUh, X 0.6; 

Drawing Aa based on Cowan and Wurdack 31376 (US), Ab, 

AC on Steyermark 58293 (F), and B on Steyermark and 

IVurdack 1220 (US). 

members of other genera. A disproportionately 

elongate proximal above-ground culm internode 

followed by two or more obsolete internodes is a 

feature shared by Glaziophyton mirabile, by Aulonemia queko, and by a number of species of 

Arthrostylidzum, including A. schomburgkii. The 

strongly thickened and indurate culm sheaths, 

branch sheaths and leaf sheaths with leathery 

blades, and the (sometimes irregular) incidence of 

solitary and appressed branches at culm nodesfeatures characteristic of all known species of 

Myiiocladus-are found also in Aulonemia de-' 

pexa and A. eflusa, and in all known species of 

Greslania. Probably because some species of both 

genera have sessile leaf blades Myriocladus has been 

said to resemble (be related to?) Neurolepis (Swallen in Steyermark, et al., 1951:34). However, in 

its over-all combination of vegetative and reproductive features, Myriocladus is clearly disjunct from 

all other bamboo genera. 

DISTRIBUTIoN.-The twenty known species of Myriocladus are known only from the sandstone tablelands region of Venezuela. They occupy generally 

mesophytic, rarely swampy habitats, at recorded altitudes ranging from 1025 m to 2500 m. 

Checklist of New World Bamboos 

Described Under the Genus Myriocladus 

1. Myriocladus afinis Swallen, in Maguire, Wurdack, et al., 1957:244, fig. 3f. 

2. Myriocladus cardonae Swallen, in Steyermark, 

et al., 1951:35. 

3. Myriocladus confertus Swallen, in Maguire, 

Wurdack, et al., 1957:248, fig. 4c; in Maguire, 

Steyermark, Wurdack, et al., 1957:397. (Figures 38, 39~). 

4. Myriocladus churunensis Swallen, in Steyermark, 1967: 132. 

5. Myriocladus distantiflorus Swallen, in Maguire, 

Wurdack, et al., 1957:248, fig. 4d. 

6. Myriocladus exsertus Swallen, in Maguire, 

7. 

8. 

9. 

10. 

11. 

12. 

13. 

14. 

15. 

16. 

17. 

18. 

19. 

20. 

Wurdack, et al., 1957:242, fig. 3c. 

Myriocladus gracilis Swallen, in Maguire, Steyermark, Wurdack, et al., 1957:393, fig. 74. 

Myriocladus grandifolius Swallen, in Maguire, 

Wurdack, et al., 1957:245, fig. 3h. 

Myriocladus longiramosus Swallen, in Maguire, 

Wurdack, et al., 1957:243, fig. 3e. 

Myriocladus maguirei Swallen, in Maguire, 

Wurdack, et al., 1957:239, fig. 3b. 

Myriocladus neblinaensis Swallen, in Maguire, 

Wurdack, et al., 1957:240, fig. 1. 

Myriocladus paludicolus Swallen, in Maguire, 

Wurdack, et al., 1957:246, 248, fig. 4b. 

Myriocladus paraquensis Swallen, in Maguire, 

Wurdack, et al., 1957:246, figs. 3i, j. 

Myriocladus paruensis Swallen, in Maguire, 

Wurdack, et al., 1957:244, fig. 3g. 

Myriocladus purpureus Swallen, in Maguire, 

Wurdack, et al., 1957:248, fig. 4e; in Maguire, 

Steyermark, Wurdack, et al., 1957:397. 

Myriocladus simplex Swallen, in Maguire, 

Wurdack, et al., 1957:242, fig. 2. 

Myriocladus steyermarkii Swallen, in Maguire, 

Wurdack, et al., 1957:247, fig. 3a; 395. 

Myriocladus uariabilis Swallen, in Maguire, 

Wurdack, et al., 1957:248, fig. 4a; in Maguire, 

Steyermark, Wurdack, et al., 1957:396. 

Myriocladus virgatus Swallen, in Steyermark, 

et al., 1951:34, 35, fig. 4. (Figures 39~a-~c, 40) 

Myriocladus wurdackii Swallen, in Maguire, 

Wurdack, et al., 1957:248, fig. 4f; in Maguire, 

Steyermark, Wurdack, et al., 1957:398. 

Neurolepis Meisner 

FIGURE 41 

Neurolepis Meisner, 1843, I:426, II:325.-Pilger, in Engler 

Platonia Kunth, 1829: 139 [non Rafinesque, 1808, nec Martius, 

Planotia Munro, 1868:70.-Bentham, in Bentham and Hooker, 

and Prantl, 1906:21.-McClure, 1957:206. 

18291,- Nees, 1834:486. 

1883: 1209.-Hackel, in Engler and Prantl, 1887:93. 

Plants typically unicespitose, exceptionally-as in 

N. aristata (Munro) Hitchcock-forming dense, 

more or less extensive thickets; unarmed. Rhizomes 

pachymorph, relatively slender. Culms self-supporting, typically unbranched in the vegetative state, 

the internodes fistular and cylindrical or terete. 

Leaf blades typically coarse and grass-like, petiolate 

or sessile, persistent or articulated with the leaf 
98 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 9 99 

FIGURE 40.--Myriocladus virgatus Swallen. A, Tip of leafy 

flowering culm, x 0.6; 6, spikelet, x 12; c, transitional 

glumes, x 12; D, floret I, showing dorsal aspect of palea, 

x 12; E, floret 11, showing terminal depauperate floret lying 

in the sulcus of t&e palea, x 12; F, lemma I, dorsal aspect, 

x 12; c, palea, ventral aspect, x 12; H, lodicule complement, 

x 12; I, stamen, x 12; J, gynoecium, x 12. Drawing A 

based on Cowan and Wurdack 31376 (US), and B-J on 

Steyermark 58293 (F) . 

sheath, the transverse veinlets not at all visible to 

clearly manifest, externally. 

Inflorescences semelauctant, each terminal to an 

unbranched culm, broadly to narrowly paniculate 

with a strong excurrent rachis, the branches appressed or spreading, commonly only one order of 

the branches (the first or the last or, in some species, 

none) showing secund orientation; prophylla lacking; subtending bracts commonly lacking or 

obsolete. Transitional glumes at the base of the 

spikelet 4 (-5, as in N. angusta Swallen): I and 11 

“empty glumes,” 111 and IV (-v) “sterile (empty) 

lemmas.” Spikelets pedicellate, typically containing 

but one perfect floret (the terminal one), the 

rachilla very short, articulated immediately below 

the locus of insertion of either the first sterile 

lemma only, or both the fertile and the sterile 

lemmas, and not prolonged behind the palea of the 

terminal perfect floret. Fertile lemma sometimes 

embracing its palea fully, but as a rule only basally 

so at maturity. Palea gaping antically, dorsally 

convex, in some species with an entire apex and not 

at all sulcate, elsewhere showing a very short, 

narrow sulcus terminating in an emarginate or 

bimucronulate apex. Lodicules 3, subequal or 

the anterior two appreciably asymmetrical and 

paired, the posterior one symmetrical and smaller 

(at least either shorter or narrower). Stamens 

typically 3-exceptionally -4, -5, -6 as in Cuatrecams 14813 (US) (AT. aristata (Munro) Hitchcockthe filaments filiform, free. Stigmas 2. Mature 

fruit an oblong, straight caryopsis with a uniformly 

thin, glabrous, dry pericarp, the sulcus abbreviated 

distally and proximally, the basal position of the 

embryo clearly to weakly manifest. [Examples of 

mature fruits known to me only from a specimen 

(US) identified by Soderstrom as Neurolepis elata 

(Kunth) Pilger, collected in Colombia by Cuatrecasas (no. 11522). These are extremely minute 

2.0-2.5 mm long) but well formed.] 

ETmfoLom.-The name Neurolepis, coined from 

the Greek words, neuron, nerve, and lepis, scale, 

apparently alludes to the strong, keel-like median 

nerve of the empty glumes in the type-species of this 

initially monotypic genus. 

(Kun th) Pilger 

(in Engler and Prantl, 1906:Zl); Platonia elata 

Kunth (1830:327, pl. 76). 

RELATIONSHIPS.-Plants comprised by Neurolepis 

share the following features with plants comprised 

by the genus Swallenochloa hollow clum internodes, a similar spikelet structure, and inflorescenses with branches in secund orientation. Plants 

of the genus h’eurolepis differ from plants of the 

genus Swallenochloa most conspicuously in their 

coarse, grass-like appearance and generally restricted 

radial development, besides bearing very much 

larger inflorescences that are confined to the terminal position on culms devoid of lateral branches. 

Ostensibly allied to Chusquea by a similar spikelet structure and (in some species) by certain 

features of the inflorescence, members of the genus 

A’eurolepis differ from members of the genus 

Chusquea most conspicuously in bearing an inflorescence only at the tip of an unbranched culm, 

and in the incorporation of the following vegetative 

features: the generally coarse, grass-like (not bamboo-like) appearance of the plant (due to the shape 

and dimensions of the leaf blades, plus the lack of 

a strong differentiation between culm sheaths and 

leaf sheaths); the relatively slender form of the 

pachymorph rhizomes; and culms with hollow internodes but no lateral branches. 

When Kunth (1829: 139) published the genus 

Platonia, he stated that it differs from Chusquea 

only in a peculiar habit. However, on page 327 of 

the same work (1830), he said of Platonia elata 

Kunth-basionym of Neurolepis elata (Kunth) 

Pilger-“Cette plante prksente tous les caracteres 

du genre Chusquea; mais elle en diffkre tellement 

par son port, que je n’ai pas hCsitC B en former 

un genre particulier.” Nees (1834:486) reduced 

hTeurolepis (as Platonia Kunth) to the rank of 

subgenus under Chusquea. Ruprecht (1839:30), 

who did not include Neurolepis (Platonia elata 

Kunth) in his monograph, expressed the opinion 

that Platonia in its true nature is very far from 

Clausquea: “Platonia vero natura sua valde a Chusquea distat.“ 

TY PE-SPEC~E~. -New0 lep is e la ta 

NUMBER 9 101 

FIGURE 41.--NeuroZepis aristata (Munro) Hitchcock (A) and 

Neurolepis pittieri McClure (B-N). A, Flowering plant, x 0.6; 

B, branch of an inflorescence, x 2.4; c, spikelet, x 4.8; D, 

E, transitional glunies, x 4.8; F and G, sterile lemmas, x 4.8; 

H, fertile lemma, x 4.8; I, palea, x 4.8; J, lodicule complement, x 4.8; K, stamen, x 4.8; L, gynoecium, x 4.8; M, apex 

of leaf sheath, x 1.2; N, diagrammatic cross-section of floret. 

Drawing A based on Cuatrecasas 14813 (US) and B-N on 

Pittier I0067 (US). 

Freier’s study (1945: 103) of Neurolepis pittieri 

and N. aristata reveals a bambusoid type of leaf 

anatomy in both species. It seems to me, however, 

not unlikely that intensive comparative studies of 

the known members of the genus Neurolepis, undertaken on a broader interdisciplinary basis, may 

result in their being given a revised disposition 

within the subfamily Bambusoideae, possibly as a 

tribe distinct from the “true bamboos.” 

The genus was treated briefly by Soderstrom (in 

Maguire, et al., 1969) and a key given to the nine 

species which he recognized. Gould and Soderstrom 

(1970) reported a chromosome count of 2n = 48 

for a specimen of N. aperta from Colombia. 

DIsTRIBUTIoN.-Species of Neurolepis have been 

reported from an area that embraces sites in Peru, 

Ecuador, Colombia, Venezuela, and the island of 

Trinidad. All are reported from relatively high 

elevations (2900-4500 m); some of them inhabit 

ecological formations characterized as “pAramo.” 

According to Munro (1868:73) Chusquea aperta 

Munro [= Neurolepis aperta (Munro) Pilger] was 

collected in Colombia by Goudot, who recorded 

for it the local name “Chusquea de PAramo.” 

Notes by Jameson, collector of the type-specimen 

of Chusquea aristata Munro [= Neurolepis aristata 

(Munro) Hitchcock] are quoted by Munro (1868: 

61) as follows: “It is a tall reedy grass, found only 

on the Eastern chain of the Andes. At 13,000 feet 

[ca. 4000 m] it makes its appearance, in irregular 

patches; at 15,000 feet [5000 m] it completely covers 

the whole surface, forming what the natives call 

a ‘carizal,’ impenetrable to man or beast. It continues upwards nearly to the limits of perpetual 

snow.” 

Annotated Checklist of Recognized Species of 

the Genus Neurolepis 

1. Neurolepis angusta Swallen, in Maguire, Wurdack, et al., 1957:249. 

Neurolepis densiflora Swallen, in Maguire, Steyermark, Wurdack, et al., 1957:399. 

2. Neurolepis aperta (Munro) Pilger, in Engler 

and Prantl, 1906:21. 

Planotia aperta Munro, 1868:73. 

Planotia ingens Pilger, 1898:721. 

Neurolepis ingens (Pilger) Pilger, in Engler and 

Prantl, 1906:21. 

3. Neurolepis aristata (Munro) Hitchcock, 1927b: 

313 (Figure 41~). 

Chusquea aristata Munro, 1868:61. 

Planotia acuminatissima Munro, 1868:72. 

Planotia stiibelii Pilger, 1898:720. 

Planotia tesselata Pilger, 1898:720. 

Neurolepis acuminatissima (Munro) Pilger, in 

Neurolepis stiibelii (Pilger) Pilger, in Engler and 

Neurolepis tesselata (Pilger) Pilger, in Engler and 

Neurolepis weberbaueri Pilger, 1921:446. 

Engler and Prantl, 1906:21. 

Prantl, 1906: 21. 

Prantl, 1906:Zl. 

4. Neurolepis diuersiglumis Soderstrom, in Maguire, et al., 1969:16-18, 20. 

5. Neurolepis elata (Kunth) Pilger, in Engler 

and Prantl, 1906:21. 

Platonia elata Kunth, 1830:327, pl. 76. 

Planotia elata (Kunth) Munro, 1868:71. 

Planotia nobilis Munro, 1868:72. 

Nezi?‘olepis nobilis (Munro) Pilger, in Engler and 

6. Neurolepis glomerata Swallen, in Maguire, 

Prantl, 1906:21. 

Steyermark, Wurdack, et al., 1957:399. 

7. Neurolepis mollis Swallen, 1931: 14. 

8. Neurolepis pittieri McClure, 1942:181, fig. 8 

(Figure 41~-N). 

9. Neurolepis virgata (Grisebach) Pilger, in 

Engler and Prantl, 1906:Zl. 

Platonia virgata Grisebach, 1864:530. 

Planotia uirgata (Grisebach) Munro, 1868:71. 

Neurolepis nigra Swallen, 1957b: 400. 

Rhipidocladum McClure, new genus 

FIGURE 42 

Plantae unicaespitosae, inermes. Rhizomata 

pachymorpha. Culmi infra saltem medium sese 

sustinentes erecti, sursum vulgo vel scandentes vel 
102 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

apice penduli; internodiis cylindratis, intus fistulosis. Ramorum complementum ad nodos medianos 

culmorum insertum; eiusdem axi primario applanato, subtriangulato, ad culmi superficiem appresso 

adnatoque, neque cylindrato neque segmentato; 

axibus secundariis numerosis, gracilibus, subaequalibusque, margine axis primarii insertis, ita ordinatione rhipidiforme sese ostendentibus. Foliorum 

lamina venulis transversis in speciebus plerisque 

extus vulgo haud manifestis, alibi plus minusve 

Clare visibilibus. 

Inflorescentiae semelauctantes, ramificatione 

typice racemosa, rachidi vel percurrenti vel deliquescenti; spiculae insertione et orientatione diversae; prophyllis bracteisque nullis. Glumae 

transitionales (glumae vecuae et lemmata sterilia) 

numero diversae atque nonnumquam inconstantes. 

Spiculae vel sessiles vel subsessiles vel pedicellatae, 

flosculos hermaphroditos vel paucos vel plures continentes, apice in anthecio depauperato sterilique 

terminantes. Lemma fertile maturitate saltem paleam suam basi tantum circumplectens. Palea dorso 

2-carinata et late sulcata, antice hians. Lodiculae 

(nonnumquam -2, -1, -0) typice 3, duabus anterioribus plus minusve asymmetricis atque geminatis, 

posteriore symmetrica sat minoreque. Stamina 3, 

filiamentis filiformibus liberisque. Stigmata 2. 

Fructus maturus caryopsidens; caryopsis oblonga 

sulcataque, leviter compressa, apice cuspidata; 

pericarpio vel pergamineo vel crustaceo, glabro, 

vel opaco vel nitido, crassitudine uniformi; sulco 

et embryotegio aut valde aut debiliter manifestis. 

Plants unicespitose, unarmed. Rhizomes pachymorph. Culms self-supporting below, commonly 

either scandent or pendulous above, the internodes 

cylindrical, hollow. Primary branch buds (when 

present) at midculm nodes, solitary, each containing 

a single initial primordium. Branch complement at 

midculm nodes monoclade, of restricted insertion, 

the primary element neither cylindrical nor segmented, but thin, roughly triangular, appressed and 

more or less strongly adnate to the surface of the 

culm, the secondary axes numerous, slender, subequal, arising from primordia formed on the margin 

of the flat, primary element, of close-set apsidate 

insertion, with a fan-like pattern of orientation. 

Leaves (blades of leaf sheaths) with transverse 

veinlets in most species not at all (or scarcely) 

manifest, but in an occasional one more or less 

clearly visible externally. 

Inflorescences semelauctant, typically of racemose 

branching, the rachis either deliquescent, or percurrent and straight or zigzag, the spikelets inserted 

and oriented in diverse patterns. Excepting the 

lowermost one, bracts subtending primary branches 

usually obsolete or lacking; typical prophylls lacking. Transitional glumes (empty glumes and sterile 

lemmas) 2 or 3 (rarely 4) the uppermost one often 

subtending a depauperate flower, the others empty. 

Spikelets comprising few to several perfect florets, 

and terminating in one or more progressively depauperate sterile anthecia. Rachilla segments possessing at maturity the potential for disarticulating 

just below the locus of insertion of each fertile 

lemma. Lemma (when subtending a functional 

flower) fully embracing the palea only basally at 

maturity. Palea broadly sulcate and 2-keeled dorsally, the margins not at all or only slightly and 

partially overlapping. Lodicules typically 3, the 

anterior two asymmetrical and paired, the posterior 

one smaller (at least shorter or narrower) and 

symmetrical (all three symmetrical and subequal in 

R. parviflorum; exceptionally -2, -1, -0 as recorded 

for R. ampliflorum). Stamens 3, the filaments filiform, free. Stigmas 2. Mature fruit (as far as 

known; available examples are rare) an oblong, 

subfusiform, sulcate, apically mucronate caryopsis 

lightly compressed and lightly curved or asymmetriFICURI: 42.-Rhipidoclrrdurn harmonicurn (Parodi) McClure 

(A-Q) and Rhipidoclnduna vet?ici/laturn (Nees) McClure 

(R-T) , A, Rhizome and base of its culm, x 0.6; B, culm internode between nodes v and 1’1, showing first-order element of 

the branch complement with second-order elements developing from buds on its margin, x 0.6; c, first-order element of 

branch complement from 8, removed and shown from the 

adaxial aspect, x 1.8; D, fully developed midculm branch 

complement, basal portion only, in situ, x 0.48; E, midculm 

sheath, adaxial aspect, x 0.6; F, sterile leafy twig, x 0.6; 

c, leafless flowering twig: x 0.6: H, apex of leaf sheath and 

base of blade, x ca. 3; I, spikelet, sessile with portion of 

rachis attached, x ca. 1.8; J, transitional glumes from I, 

x ca. 6; K, floret, x ca. 3.6; L, lemma, x ca. 3.6; M, palea, 

x ca. 3.6; N, lodicule complement, x ca. 7.2; 0, stamen, 

x ca. 6; P, gynoecidm, x ca. 14.4; Q, stigmatic processes, 

greatly enlarged; R, fruit, embryo side, x 7.2; s, fruit, hilum 

side, x 7.2; T, fruit, longitudinal section, x 7.2. Drawings 

A, B based on McClure 21398 (US), c, E, F, H on McClure 

21416 (US), D on Camp E-1613 (US), G, I-M on Killup and 

Smith 25622, N-Q on Sodiro s. n., Oct. 1900 (US), and 

R-T on Glaziou 22424 (P) , 
NUMBER 9 10s 
104 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

cal in the dorsiventral plane; pericarp pergamineous (crustaceous in R. verticillatum), glabrous, of 

uniform thickness; the sulcus and embryotegium 

weakly to strongly manifest. 

ETYMoLoGY.-The name Ripidocladum, derived 

from the Greek, rhipis, fan, and klados, branch, 

alludes to the fan-like pattern formed by members 

of a mature midculm branch complement in all of 

the bamboos herein allocated to the genus. 

TYPE-sPEcIEs.-RhiPidocladum harmonicum (Parodi) McClure, new combination. Arthrostylidium 

harmonicum Parodi (1944:478, fig. 1). 

RELATIONSHIPS.-The combinations of morphological features used herein to define Rhipidocladum 

places this genus between Merostachys and Arthrostylidium, with some of its features indicating 

affinities toward Merostachys and other features 

indicating affinities toward Arthrostylidium. The 

taxa I have allocated to Rhipidocladum are perfectly uniform in terms of the ontogeny and morphology of the midculm branch complements and are, 

in this aspect, indistinguishable from the taxa I 

have retained in Merostachys. However, a comparison of the shape and orientation of the blade 

borne on culm sheaths representing their midculm 

range, insofar as these features have been documented, suffices to distinguish plants of taxa allocated to the genus Rhipidocladum (Figure 42~) 

from plants of taxa retained in the genus Merostachys (Figure 37~). 

As Merostachys and Rhipidocladum are here 

circumscribed, the component species of both 

genera show a considerable range of variation in 

morphological features of their inflorescences. The 

range of these diversities is the more marked in 

Rhipidocladum, where at one extreme it embraces 

deceptive simulations of Merostachys and at the 

other, Arthrostylidium. Fournier (1881: 131) was 

misled by this latter case into transferring Steudel’s 

Arthrostylidium racemiflorum to Merostachys. 

As shown in the key below, the species currently allocated to Rhipidocladum embrace, in 

their inflorescences, six distinct patterns of insertion and orientation of the spikelets. An intimation 

of ostensible relationship between Rhipidocladum 

and Merostachys appears in the noticeable to 

marked secund orientation of the spikelets in the 

spicate racemes of seven species of Rhipidocladum. 

Rhipidocladum and Arthrostylidium coincide 

here and there (compare R. harmonicum with 

Arthrostylidium venezuelae, and R. urbanii with 

A. cubense) with respect to certain forms assumed 

by the inflorescences. As mentioned above, however, 

the respective members of these two genera are 

readily distinguishable by the characteristics of 

their midculm branch complements. 

DISTRIBUTIoN.-The eleven named species comprehended by the present circumscription of the 

genus Rhipidocladum represent taxa that occupy 

open or forested mesophytic situations at low to 

moderate altitudes in frost-free areas, from Mexico 

to Brazil and Bolivia. 

Key to Recognized Species of Rhipidocladum 

la. Rachis deliquescent; spikelets pedicella te . , , , . , , , , , . , 11. R. verticillatum, new combination 

1b. Rachis excurrent; spikelets sessile or subsessile ................................................................. .2 

2a. Spikelets inserted oh the rachis in predominantly binate clusters ...................................... 

......................................................................................... 4. R. geminatum, new combination 

2b. Spikelets of monate (solitary) insertion ............................................................................ ..3 

3a. Rachis ,geniculate (at least in its distal portion) ; spikelets in distichous orientation,,,,. .. 

.......................................................................... 5. R. harmonicurn, new combination 

3b. Rachis not genticulate; spikelets in secund orientation,, ................................ 

4a. Spikelets oriented within a single plane giving a banneret-like form to each inflorescence 

................................................................................... 6. R. maxonii, new combination 

4b. Spikelets not oriented within a single plane ........................................................................ 5 

5a. Spikelets 4-5 cm long; the rachilla not disarticulating promptly at maturity ...................... 

5b. Spikelets not exceeding 2.5 cm in length; the rachilla disarticulating promptly at maturity 

3. R. bartlettii, new combination; 7. R. pawiflorum, new combination; 8. R. pittkd, new 

combination; 9. R. prestoei, new combination; 10. R. rocemiflorum, new combination. 

.................................................................................... 1. R. ampliflorum, new combination 

(the following six species) : ......................... 2. R. angustiflorum, new combination; 
NUMBER 9 

The six names (in couplet 5b) represent Hackel’s 

(1903a:69) image of the genus Arthrostylidium as 

tentatively adumbrated but not typified by him 

(see p. 19). The corresponding binomials label 

several closely related taxa that are poorly represented by their type-specimens, and but weakly 

differentiated from each other by their protologs. 

Numerous currently available specimens represent 

these and similar taxa too incompletely and too 

incongruently to support confident decisions with 

respect to the existence of either suspected synonymies or possible specific or subspecific novelties. 

Annotated Checklist of Recognized Species 

of Rhipidocladum 

1. Rhipidocladum ampliporurn (McClure) McArthrostylidium ampliflorum McClure, 1942: 167, 

HoLoTYPE.--Karsten s. n. (US). Specimens examined include the holotype and elements of the 

holotype collection (Us ex LE, US ex W). 

Clure, new combination. 

fig. 1. 

DISTRIBUTION .-Venezuela. 

2. Rhipidocladum angustiflorum (Stapf) McClure, new combination. 

Arthrostylilium angustiflorum Stapf, 1913:268. 

H0LoTYPE.-“Communicated by Messrs. Sander 

8e Sons, Bruges, 3rd May, 1912” (K). Specimens 

examined include the type-collection (US ex K). 

DrsTRIBLJTIoN.-“Tropical America.” 

3. Rhipidocladum bartlettii (McClure) McClure, 

Arthrostylidium bartlettii McClure, 1954:81.-McHOLOTYPE.--Bartlett 12154 (us). Specimen exnew combination. 

Clure in Swallen 1955:38. 

amined, the holotype. 

DIsTRIBuTI0N.-Guatemala. 

4. Rhipidocladum geminatum (McClure) McArthrostylidium geminatum McClure, 1942: 169, 

HoLoTYPE.-“Jahn 125” (=Jahn 11, teste Dra. 

Zoraida Luces de Febres) (VEN). Specimen examined, the holotype. 

Clure, new combination. 

fig. 2. 

DIsTRIBLJT1oN.-Venezuela. 

5. Rhipidocladum harmonicum (Parodi) McClure, new combination (Figure 42~-~). 

Arthrostylidium harmonicum Parodi, 1944:479, fig. 

1.-McClure, in Sohns and Swallen, 1955: 133. 

HoL0TYPE.-Vargas 3260 (Parodi herbarium). 

Specimens examined include McClure 21416 (US); 

Sodiro “25-Pilger” (US ex Sodiro herbarium); 

Vargas 3260 (US ex Parodi). 

6. Rhipidocladum maxonii (Hitchcock) McClure, 

D1sTRIBuTIoN.-Ecuador; Peru. 

new combination. 

Arthrostylidium maxonii Hitchcock, 1927a:SO. 

HoLoTYpE.-Maxon 8154 (us). Specimens examined include the holotype and Standley 33643, 

33812, 39096, 39105, 39179, 39411 (US); Standley 

and Torres 50981 (US); Standley and Valerio 

49781 (US). 

DISTRIBUTION .-Costa Rica. 

To the midculm branching habit that gives to 

the culms of all species of Rhipidocladum a strong 

merostachyoid appearance, R. maxonii adds another merostachyoid feature - the conspicuously 

secund orientation of the branches of its banneretlike inflorescence. Other less conspicuous but clearly 

divergent morphological features, however, testify 

(in the key) to the authenticity of a generic disjunction between Rhipidoclalum and Merostachys. 

7. Rhipidocladum parviflorum (Trinius) McClure, new combination. 

Armdinaria parviflora Trinius, 1835: 6 19. 

1891 (LE). 

Arthrostylidium trinii Ruprecht, 183929. 

HOL0TYPE.-“in sylvis Brasiliae pr. Ypenema 

Bras. Riedl!” [I891 (LE). Specimens examined include Macedo 4555 (US); Regnell ser. IIZ no. 1420 

(US ex Regnell herbarium); Riedl189 (US ex LE); 

Soderstrom 979 (US); Tamayo 2718 (US ex VEN). 

8. Rhipidocladum pittieri (Hackel) McClure, 

HOL0TYPE.-“in sylvis pr. Ypenkma Bras.” [Riedl 

DIsTRIBLJTIoN.-Brazi~; Venezuela. 

new combination. 

Arthrostylidium pittieri Hackel, 1903a:75. 

HoLoTYPE.--Pittier 7193 (W). McClure, in Swallen, 1955:40. 

Arundinaria pittieri (Hackel) E. -G. Camus, 1913, 

I:40. 

Specimens examined include Grant 1036 (US ex 

A); Heyde and Lux 4302 (US); Pittier 7193 (US); 

Standley 78356 (US). 
106 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

DISTRIBUTION.-costa Rica, Guatemala, Nicaragua. 

In his original description of Arthrostylidium 

pittieri, Hackel (1903a:76) gives this species only 

2 lodicules but 3 stigmas; however, a specimen 

from the type-collection-Pittier 7193 (US)-shows 

lodicules typically 3 and stigmas typically 2. Some 

lodicules in this specimen show an abnormality 

in the form of a style-like apical prolongation 

which is terminated by several (often 3) bristles. 

It seems unlikely that this structure was misconstrued by Hackel as the pistil. The occasional appearance, however, of morphological anomalies in 

the pistil itself (e.g., a supernumerary stigmatic 

branch) might be responsible for the erroneous 

impression recorded by him for this species. 

9. Rhipidocladum prestoei (Munro) McClure, 

new combination. 

Arthrostylidium prestoei Munro, 1895: 186. 

HoLoTYPE.--Prestoe [Trinidad Botanical Garden 

Herbarium no. 16751 (K). 

Arundinaria prestoei (Munro) Hackel, 1903d:516. 

Specimens examined include Broadway 4922 (US 

ex TRIN); Prestoe [Trinidad Botanical Garden 

Herbarium no. 16751 (US ex B [compared with 

type at Kew by Agnes Chase]). 

10. Rhipidocladum racemiflorum (Steudel) McArthrostylidium ? racemiflorum Steudel, 1854:336. 

Merostachys racemiflora (Steudel) Fournier, 188 1 : 

Specimens examined include Brother Angel 721 

(US); Camp E-3814 (US); Curran 198 (US); Davidson 1297 (F); Ghiesbreght 234 (US ex LE, US ex 

P); Liebmann 144 (US ex C); McClure 21221 (US); 

Molina 8072 (US ex EAP); Skutch 3806 (US). 

DIsmIBuTIoN.-~ana~ Zone, Colombia, Costa 

Rica, Ecuador, Honduras, Mexico, Panama. 

11. Rhipidocladum verticillatum (Nees) McClure, 

new combination. 

Arundinaria verticillata Nees, in Martius, 1829: 

523.-Kunth, 1834:483, pl. 155, 156; 1835:348.- 

Ruprecht, 1839:25, pl. 3: fig. 7.-Munro, 1868: 

23.-Doell, in Martius, 1880: 166. 

Ludolphia verticillata (Nees) Willdenow, 1833:25. 

HOLOTYPE.-Sellow s.n. (€3). Specimens examined 

DIsmIBuTIoN.-Trinidad. 

Clure, new combination. 

HoLoTYPE.-Gh iesbregh t 234 (?LE). 

131. 

include Glaziou 20248 (US ex P; US ex C), 

Sellow s.n. (US ex B; US ex LE). (F); 

DIsTRIBUTIoN.-Brazil. 

Swallenochloa McClure, new genus 

FIGURES 43-45 

Plantae unicespitosae, inermes. Rhizomata pachymorpha. Culmi sese sustinentes, in speciebus plerisque adhuc saltem cognitis rigide erecti; internodiis 

fistulosis, supra locum insertionis vel ramorum vel 

gemmarum complementi tantum vel breviter vel 

prorsus sulcatis, alibi teretiblis. Ramorum complementum ad nodos culmorum medianos insidens 

raro absens, nonnumquam monocladum, typice 

solutim pleiocladum, axibus primariis vulgo 3 

(usque 5), ad libram confertissime insertis et ex 

gemmis discretis, unoquoque mox vel demum ex 

gemmis ad nodos proximos suos proliferanti. Foliorum lamina venulis transversis in speciebus nonnullis haud visibilibus alibi plus minusve Clare 

manifestis. 

Inflorescentiae semelauctantes, typice confertim 

paniculatae, lineares et secundae; rhachidi percurrenti; ramis omnibus brevibus, plus minusve valde 

appressis; prophyllis bracteisque nullis. Glumae 

transitionales imo spiculae insidentes 4: I et 11 

“glumae vacuae,” 111 et IV “lemmata (vacua) sterilia.” Spiculae pleraeque florem perfectam unicam 

terminalemque continentes; rhachilla post paleam 

haud producta. Anthoeciorum fertilium maturorum 

lemma paleam suam imo tantum circumnectans. 

Palea antice hians, saltem apicem versus vulgo et 

sulcata et nonnumquam plus minusve manifeste 

bicarinata, dorso alibi convexa. Lodiculae 3, duabus 

anterioribus aequalibus, plus minusve valde asymmetricis et congruenter geminatis, posteriore symmetrica et aliquanto minore. Stamina 3, filamentis 

filiformibus liberisque. Stigmata 2, quorum unum 

nonnumquam sed rarissime supra basin suam in 

duas partes divisum. Fructus maturus non inventus. 

FIGURE 43.--Swallenochloa subtesseliata (Hitchcock) McClure. 

A, Leafy flowering branch complement, x 0.6; B, apex of 

leaf sheath and base of leaf blade, x 3; c, base of branch 

complement high on culm, x 3; D, tuft of ultimate branches 

of inflorescence with transitional glumes attached, x 7.2; E, 

spikelet, x 7.2; F, sterile lemma I, x 7.2; G, sterile lemma 11, 

x 7.2; H, fertile lemma, x 7.2; I, palea, x 7.2; J, lodicule 

complement, x 25; K, stamen, x 15; L, gynoecium, x 30. 

All based on Tonduz 3367, type-species (US). 
NUMBER 9 107 

B 
108 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Plants unicespitose, unarmed. Rhizomes pachymorph. Culms self-supporting, the internodes typically fistular, either terete or lightly to strongly 

sulcate above the locus of insertion of a complement of buds or branches. Branch complement at 

midculm nodes rarely lacking, occasionally monoclade, typically unrestricted pleioclade, the initial 

components most commonly 3 in number (rarely 

to 5), unequal (each of independent origin, i.e., 

developing from a separate primary bud), of level 

insertion (with their bases close together in a 

horizontal row) the middle one dominant, each 

usually proliferating from buds at its proximal 

nodes. Leaves (leaf sheath blades) with transverse 

veinlets not at all evident or more or less clearly 

manifest externally. 

Inflorescences semelauctant, commonly terminal 

to the culm as well as to leafy or leafless lateral 

axes, typically linear in profile, compact-paniculate, 

the rachis percurrent, the primary branches of 

secund orientation, those of all orders usually 

short and more or less strongly appressed; prophylls 

and bracts lacking throughout. Transitional glumes 

at the base of the spikelet 4: I and 11 “empty glumes” 

in some species sub-obsolete; 111 and IV “sterile 

(empty) lemmas.” Spikelets typically containing 

but one perfect floret, the rachilla very short, potentially disarticulating immediately below the 

locus of insertion of the first sterile lemma and of 

any fertile lemma, and not prolonged behind the 

palea of the terminal floret. Fertile lemma fully 

embracing its palea only basally at maturity. Palea 

gaping antically, sulcate and weakly to strongly 

bicarinate for a short distance at the usually emarginate or bidentate or bimucronate apex, dorsally 

convex elsewhere. Lodicules 3, the anterior two 

typically asymmetrical and congruently paired, the 

posterior one smaller (at least either shorter or 

narrower) and symmetrical. Stamens 3, the filaments filiform free. Stigmas 2, one of them sometimes (but rarely) divided above the point of its 

origin as observed in one flower of Weberbauer 

4415, (type-collection of S. weberbaueri). Mature 

fruit unknown. 

ETYMoLocY.-The name Swallenochloa commemorates the initial recognition of this genus by Dr. 

Jason R. Swallen, well known for his extensive 

field studies of New World grasses, and for his numerous contributions to the taxonomy of the 

Gramineae. 

TYPE-SPECIES.-SWa~lenoch~oa subtessellata (Hitchcock) McClure. 

RELATIoNsH1Ps.-Members of Swallenochloa share 

with members of Chusquea and members of Neurolepis a basically similar spikelet structure. Beyond this, Swallenochloa touches each of the other 

two at a few points where one or two technical 

features exceptional in one genus are sliared by 

one of its members with a member of one of the 

other genera. Examples follow. A specimen under 

Pittier 3069 (US 677457)-cited by Hitchcock 

(1927a181) as Chusquea subtessellata Hitchcock, now 

Swallenochloa subtessellata (Hitchcock) McClure, 

from which it differs in several technical detailsappears to conform to the other members of Swallenochloa in all generic aspects except the profile 

and dimensions of the strongly chusqueoid inflorescence. Specimens of an undescribed species of 

Chusquea, under McClure 212?4 (US) have the 

satellite buds and branches at midculm nodes reduced to two, as is usual in Swallenochloa, but in 

all other technical respects the material clearly falls 

within the natural boundaries of Chusquea. A 

whole plant under Fosberg 20856 (US 273832) 

shows unbranched culms with hollow internodes 

and a terminal inflorescence as in Neurolepis, but 

the nature of its foliage places it clearly in Swallenochloa. 

Bamboos of the genus Swallenochloa differ most 

conspicuously from bamboos of the genus Chusquea by combining inflorescences of linear profile 

with distinctive features of the vegetative organs, 

including hollow culm internodes, primary complement of buds or branches at midculm nodes comprising fewer members, presenting several markedly 

distinctive morphological aspects, and foliage leaves 

FIGURE 44.-?wallenochloa subtessellata (Hitchcock) McClure. 

A, Habit of leafy flowering branch showing tessellation of 

leaf blades, x 0.6; R, sheath subtending flowering twigs 

(left) and detached blade (right), x 1.2; C, apex of leaf 

sheath and base of leaf blade, x 7.2; D, tuft of ultimate 

branches of inflorescence with transitional glumes attached, x 

7.2; E, spikelet (without pedicel and transitional glumes) , x 

7.2; F, sterile lemma I, x 7.2; G, sterile lemma 11, x 7.2; H, 

fertile lemma, x 7.2; I, palea, oblique aspect; J, palea, dorsal 

aspect, x 7.2; K, lodicule complement, x 15; L, stamen, 

x 15; M, gynoecium, x 30. All drawings based on Pittier 

3069, a variant form of the species, (US). 

110 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

generally showing definitely xeromorphic structure, 

texture, and habit. 

Swallenochloa subtessellata (as Chusquea subtessellata) is by Niilo Virkki (1963) given a chromosome number 2n = 36. Janaki Ammal (1959) gives 

three species of Chusquea the chromosome number 

of 2n = 48. 

The genus Swallenochloa shares the following 

features with the genus Neurolepis: hollow culm 

internodes, a similar spikelet structure, and a strong 

tendency to secund orientation in the branches of 

the inflorescences. However, members of Swallenochloa differ from members of Neurolepis most 

conspicuously in the dense and extensive development of individual plants, the freely branched 

culms; the smaller size, bamboo-like shape, xeromorphic texture, and upright habit of the foliage 

leaves; and the very much smaller and generally 

more compact inflorescences, which are not confined 

to the terminal position on the culms. 

Among the many available specimens that clearly 

belong to the genus Sz~lallenochloa but remain unidentified as to species, certain features of the reproductive structures show a high incidence of 

intergradation. For this reason, the differentiation 

and definition of species or other subgeneric taxa 

on the basis of typical conventional herbarium 

specimens presents difficulties at many points. The 

high incidence of abortive or distorted structures 

in the spikelets of many of the variants, and the 

scarcity of normal, mature fruits suggest (1) the 

possibility of a hybrid origin for the genus or the 

continued but only partially successful introgression between its members, and (2) the existence 

of insects or fungi whose parasitic activities may 

be responsible for the widespread incidence of distortion shown by the spikelets. As matters now 

stand, so great is the incidence of variation and 

distortion in the spikelets, and so obscure are the 

bases for these diversities, that it is difficult to determine with confidence a suitable taxonomic disposition for many of the available collections. 

The genera Chusquea, Neurolepis, and Swallenochloa represent groups of species that are 

closely related in the essential features of the reproductive apparatus, members of Chusquea leading in the diversity of form expressed in the 

branching of the rhizomes, culms, and inflorescences. From the other bamboo genera of the New 

World flora, these three are set off, as a group, by 

the regular appearance in their members of two 

usually empty “sterile lemmas” following the second “empty glume” at the base of the spikelet, and 

by the termination of the spikelet in a perfect 

floret, without a prolongation of the rachilla behind it. Chusquea is unique among the three in 

the efistulose structure of all segmented vegetative 

axes; in the universal presence of more than one 

branch bud at midculm nodes; and in the constellate insertion of the branch buds and members of 

the branch complement at midculm nodes. Species 

of Chusquea show, in aggregate, the most extreme 

range of ecological tolerance, or preference. Neurolepis recedes from the other two members of the 

group in the coarse, grass-like character of the 

plant; the usually large, persistent leaf blades; the 

universal lack of branching in the aerial range of 

the vegetative part of the culm; and the usually 

rather weak development of the rhizome system. At 

least some species of Neurolepis are found at relatively high altitudes and in ecological formations 

characterized as pkramo. Species of Swallenochloa 

form dense thickets, consisting of erect culms with 

hollow internodes, branch buds and branches at 

midculm nodes in level (not constellate) insertion. 

Their natural distribution is confined to high altitudes, commonly in ecological formations some of 

which are characterized as pPramo. 

The sharing by Swallenochloa, Chusquea, and 

Neurolepis of features not found elsewhere among 

New World bamboos raises questions as to the phylogenetic relationship of these three genera. Could 

Swallenochloa be the product of introgression between members of Chusquea and Nezirolepis that 

FIGURE 45.-Szoallenochloa tessellata (Munro) McClure. A, 

Small leafy flowering culm, x 0.6; B, leafy flowering culm 

with its pachymorph rhizome, x 0.6; c, midculm complement of three branch buds, x 3; D, base of midculm complement of three independent primary branches, x 3; E, 

apex of leaf sheath and base of leaf blade, x 6; F, secondary 

branch of inflorescence, bearing three pedicellate spikelets, 

x 3; G, spikelet, x 6; H, two transitional glumes, X 6; I, two 

“sterile (empty) lemmas,” x 6; J, “fertile lemma,” x 6; 

K, palea, x 6; La-c, lodicule complement, x 12; M, stamen, 

x 6; N, gynoecium, x 24; 0, rhizome and base of culms, 

x 0.6; P, cross-section of rhizome, x 5.4. Drawhg A based 

on Triana 306, type-species (US) ; B, F-N on Schuttes 18771 

(US), c, P on Garcia-Barriga 11643-A; D, E on GarclaBarriga 11684 (US), and o on Soderstrom 1339 (US), 

112 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

are, or have been, sympatric in the high altitudes 

where the three genera meet today? 

DIsTRIBUTION.-~rigina~~y allocated to the genus 

Chusquea, the five bamboos currently recognized as 

species of Swallenochloa, together with numerous 

collections representing as yet undescribed congeneric diversities, are all endemic to the New 

World. Their aggregate geographic range of distribution extends from Bolivia and Brazil to Costa 

Rica, and embraces areas distinguished by characteristic complexes of ecological factors locally 

given distinctive names such as “piramo,” “subpiramo,” etc. Recorded altitudinal limits fall between 2700 m and 4000 m above sea level. The 

highest density of incidence appears in Colombia. 

Fosberg (1944) describes in extensive and vivid 

detail the plant life and the ecological features of 

the Piramo de Sumapaz of Colombia where he collected a species of Swallenochloa referred to as follows (1944:230). 

Another grass (Chusyuea weberbaueri) is locally abundant and conspicuous. It, too, is very stiff, but it usually 

forms colonies rather than bunches, and its blades are flat 

and lanceolate. It is not climbing like other species of its 

genus. In exposed situations it is low but erect, 6 or 8 

decimeters tall when not in fruit, and in very unfavorable 

[situations] it even fruits at 1 or 2 decimeters. Usually the 

spikes make it somewhat taller. In wetter places it becomes 

a meter or more tall, and in sheltered spots, it forms small 

cane-brakes up to 3 or 4 meters deep and very hard to 

push one’s w‘ay through. 

The field notes that accompany the specimen 

(Fosberg 20856, US 2181972) read as follows: Colombia: “Cordillera Oriental, Dept. Cundinamarca, 

Cordillera de las Cruces, Piramo de Sumapaz. Common everywhere on piramo slopes; rough piramo 

plateau with wet forest running up the sides to 

about 3500 m, brush, up to 3650 m. [Plants] varying greatly in size. Aug. 19, 1934.” The specimen 

represents an apparently undescribed species of 

Swallenochloa. 

Weberbauer in his monumental work entitled 

El Mundo Vegetal de 10s Andes Peruanos (1911 

[1945: 1461) says of three species of Swallenochloa 

(under the names Chusquea spicata, C. depauperata, and C. weberbaueri) that “these dwarf forms 

occupy peat bogs and open grasslands at elevations 

between 3200 and 3500 meters” and that “with 

them are associated members of the curious genus 

Neurolepis, whose ensiform leaves emerge from the 

base of the stem and reach, in N. weberbaueri, a 

length of 2.5 meters.” 

The present published content of the genus 

Swallenochloa is limited to five recognized species 

that were originally allocated to the genus Chusquea. 

la. 

lb. 

2a. 

2b. 

3a. 

3b. 

4a. 

4b. 

Key to Currently Recognized Species of Swallenochloa 

Transitional glumes I and 11 (“empty glumes”) subobsolete; branches of inflorescence entirely glabrous ............................... 1. S. depauperata, new combination 

Transitional glumes I and 11 (“empty glumes”) about 1 mm or more in length; branches 

of inflorescence more or less noticeably to densely strigose ........................... 

Branches of inflorescence heavily waxy; spikelets tinted with wine , , , , . , ,. , , , , , ,, 

Branches of inflorescence not noticeably waxy; spikelets of stramineous hue 

Transitional glume IV (upper sterile lemma) almost as long as the lemma, a 

well-developed veins on each side of the midrib ... 3. S. subtessellata, new combination 

Transitional glume IV (upper sterile lemma) less than 3/4 as long as the lemma, and show4. S. tessellata, new combination 

Anthecium dull; pale stramineous; awn of lemma glabrous ... 2. S. spicata, new combination 

Anthecium lustrous; dark stramineous: awn of lemma antrorse-scabrous ............... 

................................................................................... 5. S. weberbaueri, new combination 

ing only one well-developed vein on each side of the midrib .................... 

......................................................................... 

Annotated Checklist of Recognized Species of the 

1. Swallenochloa depauperata (Pilger) McClure, 2. Swallenochloa spicata (Munro) McClure, new 

Chusquea depauperata Pilger, 1905: 149. 

HoLoTYPE.-Weberbauer 3709 (B). 

Genus Swallenochloa DISTRIBUTION.-Peru. 

new combination. combination. 

Chusquea spicata Munro, 1868:60. 
NUMBER 9 113 

HOLOTYPE.-Lechler 2154 (K) the principal element. 

Chusquea humilis Lechler ex Munro, 1868:60. 

ment. 

Chusquea simplicissima Pilger, 1905: 145. 

HoLoTYPE.-kchkr 2694 (K) the principal eleHoLoTYPE.-Weberbauer 2217 (B). 

Hitchcock (1927b:310) made Chusquea simplicissima Pilger a synonym of Chusquea spicata 

Munro. I cannot with confidence verify this disposition of the matter on the basis of the documenting material available to me. Dr. Hitchcock’s 

decision, however, was based on a study of the intact type-specimens of which I have seen only fragments in which the spikelets do not exemplify a 

clearly representative norm. 

3. Szuallenochloa subtessellata (Hitchcock) McClure, new combination (Figures 43, 44). 

Chzisq uea subtessellata Hitchcock, 1927a: 8 1. 

DISTRIBUTION.-PerU. 

HoLOTYPE.-TOnduz 3?67 (us). 

DIsTRIBUTIoN.-cOsta Rica. 

Pittier 3069 (US) cited by Hitchcock (1927a:81) 

as belonging here deviates noticeably from the holotype in both vegetative and reproductive features 

(cf. Figure 44). 

4. Swallenochloa tessellata (Munro) McClure, new 

combination (Figure 45). 

Chusquea tessellata Munro, 1868:60. 

97 (K). 

TYPE-iVATERIAL,-Tria?’Ia [306] (K) and Holton 

DIsTRIBuT1oN.-Colombia. 

Munro gives the altitude at which Triana’s collection was made at 3000 feet. In Triana’s field 

notes the same figure is given in meters. 

5. Swallenochloa weberbaueri (Pilger) McClure, 

Chusquea weberbaueri Pilger, 1905: 146.-Fosberg, 

new combination. 

1944: 230. 

HoL0TuPE.-Weberbauer 4415 (B). 

~IsTRIBuTIoN.-Co~ombia to Peru. 

Yushania K. H. Keng 

FIGURES 46, 47 

Yushania Keng f., 1957:355. 

Plants unarmed, compactly unicespitose at first; 

mature clumps of pluricespitose habit in Y. niitakayamensis and of diffuse (open) habit in Y. aztecol-um. Rhizomes pachymorph, the rhizome neck 

in Y. niitakayamensis and in Y. aztecorum becoming, in plants of mature stature, up to ca 50 cm 

long. Culms of small or medium stature, self-supporting; each midculm node bearing a single initial 

branch bud; margins of the prophyllum not fusea; 

the initial primordium giving rise (simultaneously 

or nearly so and apparently before the germination 

of the bud) to typically three growing points, these 

becoming three subequal primary axes with a common base. The germination of each primary branch 

bud is accompanied by the apparently simultaneous 

elongation of its prophyllum. Branch complement 

arising at midculm nodes with the first three component axes typically dominant over any axes that 

proliferate from buds at their proximal nodes. 

Sheath at midculm nodes typically lacking a wellmarked basal girdle. Leaf blades with transverse 

veinlets not at all to weakly to sometimes strongly 

manifest externally. 

Inflorescences semelauctant, forming contracted 

to more or less prolifically branched panicles, their 

branches epulvinate and eprophyllate, with subtending bracts (excepting the somewhat well-developed basal one) small to rudimentary to obsolete. 

Spikelets few-flowered to several-flowered, each 

terminating in a depauperate sterile anthecium. 

Transitional glumes typically 2, empty. Lemma of 

fertile florets embracing its palea only basally at 

maturity. Palea broadly sulcate and 2-keeled dorsally, the margins not at all or barely imbricate. 

Lodicules 3, the anterior 2 more or less asymmetrical and paired, the posterior one smaller and symmetrical. Stamens 3, the filaments filiform, free. 

Stigmas 2. Mature fruit, as far as known, a subfusiform, sulcate apically mucronate caryopsis, the 

pericarp pergamineous, glabrous, of uniform thickness, the sulcus and the embryotegium clearly manifest. 

ETuhtoLoGY.-The name Yushania, yu, jade, and 

shan, mountain, is a Latinized form of the two 

components of the Chinese name (in the National 

dialect) of the type-locality of Y. niitakayamensis, 

type-species of the genus. A renowned geographical 

feature of Formosa (the Island of Taiwan), Yushan 

is known in Japanese as Niitakayama, and in English as Mount Morrison. 

TYpE-spEcIEs.-I’ushania niitakayamensis (Hayata) Keng f. (see McClure, 1959:209). Based on 
114 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Amndinaria niitakayamensis Hayata (1907:79). 

RELATIONSHIPS.-The simultaneous or subsimultaneous emergence, at midculm nodes, of more than 

one primary branch in Yushania recalls a similar 

feature in Apoclada. In Apoclada, however, the 

several primary axes of a midculm branch complement apparently do not have a common base, and 

the respective primordia from which they arise are 

apparently not initially enclosed in a common 

prophyllum (bud scale) as they are in Yushania. 

Members of Yushania as the genus is herein circumscribed, resemble members of the genus AuZonernia in the paniculate form of the inflorescence. 

Plants allocated to Yushania, however, differ most 

basically from plants allocated to Aulonemia in 

terms of ontogeny (and certain details of the form) 

of the midculm branch complement. In members 

of the genus Yushania, the primordium of the 

initial bud at midculm nodes, while still enclosed 

in the prophyllum, gives rise to typically 3 initial 

growing points in quick succession. This situation 

suggests the presence of the phenomenon conventionally referred to as the “telescoping” of an axisactually the shortening of the interval between the 

loci of emergence of the successive growing points 

produced by a body of primordial tissue. A satisfactory description of the distinctive sequence of 

events that produces the midculm branch complement that is characteristic of members of the genus 

Yushania awaits the intervention of anatomical 

studies designed to supplement the results achievable by way of gross morphology. 

DISTRIBuTIoN.-In the Chinese text that follows 

his Latin diagnosis of the genus, K. H. Keng (1957: 

356) says of the Old World component of Yushania: 

“Of this genus only the type species is known at 

present. It is native of Taiwan, and is found also 

on Luzon, P.I., and in the mainland Chinese provinces of Szechuan and Yunnan.” The known distribution of the New World component of the 

genus (subgenus Otatea) extends from Mexico to 

Honduras. Both the Old World component and 

Key to Subgenera of Yushania 

la. Culms proliferating from buds at their subterranean nodes; inflorescences with few spikelets: lemma weakly or not at all awned; palea acutely bifid apically; bamboos endemic 

to areas of the Old World Subgenus Yushania 

lb. Culms not proliferating from buds at their subterranean nodes; inflorescences with numerous spikelets; lemma conspicuously awned; palea obtuse apically; bamboos endemic to 

areas of the New World Otatea, new subgenus 

FIGURE 46.-Yushania aztecorum McClure and E. W. Smith. 

A, Diagram of a portion of the rhizome system (greatly 

reduced) of a plant of mature stature, with culms and roots 

omitted from the rhizomes proper: the rhizome necks are 

naturally budless and rootless. In young plants of the species 

illustrated, the necks are shorter than the rhizomes proper, 

but as a plant approaches its mature stature the necks of 

successive orders (generations) of rhizomes become progressively more elongated. [This feature has not yet, however, 

been established as generic in Yushania.] B, Basal portion 

of a culm, showing the characteristically short internodes and 

wire-like fibrovascular remains of the weathered, nonabscissile culm sheaths, x 0.6; c, midculm internodes (greatly 

reduced) showing the shallow groove (sulcus) that extends 

upward from the locus of insertion of each primary bud; 

D, midculm node, showing the solitary primary branch bud 

with its prophyllum intact, x 1.5; E, primary branch bud at 

a midculm node, with the prophyllum removed, revealing 

three growing points, each a prophyllate but unsubtended 

bud, x 1.5: F, diagram of the pattern of the loci of insertion 

of the prophylla of the three branch buds portrayed in E, 

in relation to the prophyllum of the initial primordium 

(the body of special meristematic tissue from which the 

three buds emerge), x 4.8; G, base of a midculm branch 

complement, showing the initial components in an early 

stage of development, with sheaths removed, x 0.9; H, midculm branch complement with the initial components fully 

developed, x 1.2; I, base of midculm branch complement 

in a later stage of development, after the production of 

axes of higher orders following tht germination of buds 

at the proximal nodes of the initial component axes, x 1.2; 

J, nonflowering leafy branches (distal portion only) from 

culms of relatively advanced age, showing solitary nature 

of ramification at this level, leaf blades in profile, x 0.6; K, 

leaf sheath apex with petiole and base of a narrow leaf 

blade, x 7.2; L, culm sheath from midculm range (apex 

only), adaxial aspect (left) and abaxial aspect (right) , 

x 0.9. All drawings based on McChre 21204 (US). 
NUMBER 9 115 

C 
116 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

the New World component of the genus Yushania 

embrace taxa still requiring intensive study in field 

and laboratory to determine their infrageneric 

status. The much-needed monographic treatment 

of the genus should, in my opinion, be deferred 

pending the completion of such studies, 

Subgenus Otatea McClure and E. W. Smith, 

new subgenus 

Culmi e gemmis suis subterraneis haud proliferentes; inflorescentiae spiculas multas gaudentes; 

lemma aristatum dorso dense strigosum; palea apice 

obtusa; plantae in terris occidentalibus indigenae. 

ETYMoLoGY.-The name Otatea is a Latinized 

form of the vernacular name otate (corrupted form 

of a word from the Nahuatl, language of the Aztecs) by which some native bamboos (more particularly those pertaining to this subgenus) are 

currently known and referred to in Mexico. 

TYPE-SPECIES.-YUS~U~~U aztecorum McClure and 

E. W. Smith, 

1. Yushunia (Otatea) aztecorum McClure and 

E. W. Smith, new species 

FIGURES 46, 47 A-M 

Plantae initio conferte caespitose demum sensim 

aperientes. Rhizomatum collum initio (in plantis 

staturae juventae) rhizomate proprio suo brevius, 

demum (in plantis staturae adultae) rhizomate 

proprio multo longius. Culmi usque ad 6 m alti 

et 2.5 cm diametro; internodia glabra, in plantis 

juvenibus intus farcta, in plantis adultis fistulosa, 

ea supra nodos medianos ramiferos sulcata. Ramorum complementum ad nodos medianos oriens 

initio axes primarios 3 subequales subcoaetaneosque vulgo capiens; axes ordinum superiorum a 

primarii valde superati. Vagina internodios culmi 

fovens dorso initio setis castaneis fragilis vel sparse 

vel dense vestita, demum calvescens sed papillis 

persistentibus conspersa; auriculae et setae orales 

vel nullae vel invalide evolutae; ligula arcuata, 

margine pro parte ciliata et pro parte crasse fimbriata; lamina appressa et late vel anguste triangula; vaginae ad nodos inferiores culmorum 

orientes haud deciduae, in situ demum fatiscentes. 

Vagina foliorum prominenter nervoso-striata, inter 

nervos plus minusve dense usque haud papillata, 

vel omnino glabra vel hic illic vel omnino 

puberula; auriculae et setae orales vel vix ullae vel 

nullae; ligula interior et ligula exterior dimensionibus, forma, margine vesturaque maxime variabiles; 

lamina foliorum vel lanceolata cum nervis secundariis usque 3-4, vel lineari-lanceolata usque linearis et nervos secondarios vulgo 1-2 praebens; 

venulae transversae extus haud usque Clare manifestae; pubescentia laminae charactere, crebretate, 

et distributione maxime variabilis. 

Inflorescentiae ramos et ramulos vel foliiferos vel 

efoliatos terminantes; pedunculus, rhachis et radii 

scaberuli; bracteae sat minimae nonnunquam obsoletae. Spiculae omnino pallide strigosae, laxae, 

usque ad 4 cm longae, flosculos 3-7 fungentos capientes, ubi maturae sat friabiles. Glumae transitionales typice 2, lanceolatae, valde carinatae, apice 

in aristam scabram ca 2.5 mm longam terminantes; 

I (dempta arista): 2.5-3.5 mm longa, nervos secundarios 1-2 praebens; 11 (dempta arista): 3.5-4.5 mm 

longa, nervos secundarios 2-3 praebens. Rachillae 

segmentum unumquidque dimidium lemmatis sui 

superantia, omnino scabrosa, gracilia, apice subito 

dilatato conspicue ciliata. Lemma floscularum 

FIGURE 47.-Yushania artecoruna McClure and E. W. Smith 

(A-M) and Yushania sp. (N) . A, Examples of leaf blades 

showing the range in size and shape encountered within a 

single population. The largest lanceolate one was taken from 

a young flowering culm. The long, narrow one represents 

an associated form borne by older culms in both the vegetative state and in a weakly flowering state. The several 

progressively smaller ones were taken from flowering 

branches borne on culms of progressively more advanced 

age, x 0.6. B, Leaf sheath apex with petiole and base of a 

broad leaf blade, x 7.2; c, flowering branch-one of three 

initial component axes of a branch complement high on a 

young culm-with several contracted and leafless flowering 

branches arising from buds at the proximal nodes of the 

initial axes, x 0.6 D, tip of a flowering branch from a culm 

of relatively advanced age, characterized by a pronounced 

reduction in the size of the foliage leaf and the inflorescence 

proper, x 1.2; E, spikelet, x 1.8; F, transitional (empty) 

glumes, x 6; c, floret, x 6 H, lemma, x 6; I, palea, x 6; 

J, lodicule complement, x 12; K, stamen, x 6; L, gynoecium, 

x 12; M, seedling, x 0.6; N, fruit, embryo side, showing 

embryotegium (left), and hilum side (right), showing the 

sulcus (conventionally referred to as a hilum) , x 7.2. Drawings A-M based on McClure 21204 (IJS) and N on Ortega 

4341 (US). 
NUMBER 9 117 
118 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

fungentium lanceolatum usque (dempta arista scabra 2.5 mm longa) 9 mm longum, prominenter carinatum, alibi nervos secundarios 2+ 3 praebens. 

Palea usque 9 mm longa, oblonga, apice integra 

truncata, inter et secus carinas strigosa, nervis secundariis extus haud manifestis. Lodiculae subopacae, nervosae, utrinque glabrae, apice ciliis albidis 

ca 0.5 mm longis fimbriatae; duae anticae (demptis 

ciliis) usque ad 1.5 mm longae, leniter asymmetricae et gemmatae, quasi pulvinatae; postica (demptis 

ciliis) usque ca 1.0 mm longa, elliptica. Antherae 

maturitate stramineae, 5-6 mm longae. Ovarium 

glabrum. Fructus non adhuc inventus. 

Plants at first compactly cespitose, becoming diffuse in habit as they mature. Neck of rhizomes at 

first shorter than the rhizomes proper, becoming 

longer in successive annual innovations, and eventually much longer than the rhizomes proper as the 

plant approaches its mature stature. Culms up to 

6 m tall and 2.5 cm in diameter, with glabrous 

internodes pith-filled in young plants and hollow in 

plants of mature stature; internodes above ramiferous midculm nodes sulcate. Branch complement 

at midculm nodes at first typically comprising 3 

subequal primary axes that emerge simultaneousiy 

or nearly so and dominate axes of higher orders 

that develop subsequently from buds at their own 

proximal nodes. Culm sheaths at lower (unbranched) nodes of the culms (Figure 46 B, L) 

not at all deciduous, slowly disintegrating in 

place. Leaf sheaths prominently nerved-striate, 

more or less densely to not at all papillate between 

the nerves; glabrous throughout to puberulous here 

and there or puberulous throughout; auricles and 

oral setae weakly developed to lacking entirely; 

inner and outer ligules highly variable in size, 

shape, margin, and vesture. Blade of leaf sheath 

varying in shape from lanceolate with up to 3-4 

secondary nerves, to linear with 1-2 secondary 

nerves, the transverse veinlets not at all to clearly 

manifest externally; pubescence of the leaf blade 

highly variable in character, density and distribution. 

Inflorescences terminating leafy or leafless 

branches and twigs; peduncle, rachis, and ramifications scaberulous; bracts rather small and 

sometimes obsolete. Spikelets up to 4 cm long, 

comprising 3-7 functional florets, obscurely and 

pallidly strigose throughout, open, more or less noticeably zig-zag at maturity. Transitional glumes 

typically 2, lanceolate, strongly keeled, terminating 

in a scaberulous awn about 2.5 mm long; I (without the awn): 2.5-3.5 mm long, with 1-2 secondary 

nerves; 11 (without the awn): 3.5-4.5 mm long, 

with 2-3 secondary nerves. Rachilla segments 

slender, each more than half as long as its lemma, 

conspicuously ciliate at the abruptly dilated apex, 

elsewhere slender and scaberulous; readily disarticulating at maturity. Lemma of functional florets lanceolate, up to 9 mm long (without the ca 

2.5 mm long scaberulous awn), prominently keeled, 

with 2-3 secondary nerves. Palea up to 9 mm 

long, oblong, entire and truncate apically, strigose 

along the keels and between them; secondary 

nerves not manifest externally. Lodicules subopaque, nervose, glabrous on both faces, fimbriate 

apically with cilia about 0.5 mm long, the two 

anterior ones about 1.5 mm long without the cilia, 

weakly asymmetrical and paired, apparently pulvinate at anthesis, the posterior one elliptical, up to 

about 1.0 mm long without the cilia. Anthers stramineous at maturity, 5-6 mm long, the connective 

not exserted. Ovary glabrous. Fruit not yet found. 

HOLOTYPE.--F. A. McClure 21204 (US), collected 

at about 450 m on La Dispensa, an elevated area 

inland from Rosario, State of Sinaloa, Mexico, 5 

August 1943. Vernacular name Otatk. 

RELATIONsHIPS.-The foregoing description is 

based principally upon the most complete available 

specimen, McCZure 21204 (US). It is supported in 

this and that detail by specimens from thirty-five 

other collections, the bulk of which are too fragmentary for confident infrageneric taxonomic disposition. 

DIsTRrnuTIoN.-Field notes accompanying available specimens of thirty-six collections of members 

of the subgenus Otatea examined are generally 

meager. Most commonly encountered in herbaria 

are specimens from Mexico where, as observed by 

the senior author of this subgenus, plants of this 

subgenus often occupy relatively (seasonally?) dry 

sites, sometimes in association with cacti. Specimens collected in El Salvador and Honduras are 

more rarely seen. Plants we recognize as belonging 
NUMBER 9 119 

to the subgenus Otatea have been collected at reported elevations from 200 m to 2700 m in Mexico 

and Central America. 

2. Yushunia (Otatea) acuminata (Munro) 

McClure, new combination 

Arundinaria acuminata Munro, 1868:25. 
120 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 48.-Chart of Generic Characters. Fruit: view from embryo side (left), view from hilurn 

side (right). Stigma: number of stigmas. Number in parentheses indicates that the number is 

of rare occurrence. Stamen: number of stamens. Number in parentheses indicates that the 

number is of rare occurrence. Spikelet Apex: Diagrammatic representation of the apical element 

of a spikelet. Male-female sign indicates a perfect floret. All others indicate empty florets 

reduced to varying degrees, in some cases quite depauperate. Br Subt: branches of the inflorescence (at least the lowermost ones) subtended by a bract. Br Proph: Branch prophyllum; plus 

indicates each branch of the inflorescence is subtended by a prophyllum at its basal node. 

Znfl Br: type of inflorescence branching. Pan, panicle or paniculate; Rac, raceme or racemose; 

Con to Ope, congested to open; Sec, secund; Sol, solitary; Spi, spicate; Div, diverse: Cap, capitate; 

Dif, diffuse; Dig, digitate. Infl Dew inflorescence development. S, semelauctant; I, iterauctant. 

Rhizome: L, leptomorph; P, pachymorph; LP, leptomorph and pachymorph condition existing 

within the same plant. Dotted lines indicate that the rhizome system has not been seen but 

assumed to be pachymorph. Midculm Prophyllum and Primordium: Primary buds at midculm 

node consisting of the initial primordium (solid black triangle) covered by the prophyllum 

(cone). The single initial primordinm represented by zigzag lines indicates that it gives rise to 

a specialized primary element with buds of apsidate insertion. Dotted lines indicate what the 

situation is assumed to be, although not seen. Initial (Pritnary) Branch: Either lacking, a single 

dominant one, three or more subequal ones, or many subequal ones arising from a primary 

element in apsidate fashion. Open triangles represent buds capable of germination while the 

black dots represent buds which do not germinate. L/S Znt: Long/short internodes. In plants 

of those genera marked with both a plus and a circle, there is a tendency for culms to have 

elongated internodes followed by shortened internodes. A question mark within the circle 

indicates that this feature possibly may not be present in some culms. 
NUMBER 9 121 

Midculm Initial 

Prophyllum and (Primary) Primordium Branch Fruit Genus 

QO 

QO 

Arundinuriu 

Chuspea 

Not yet 

known 

sec 

Con 

Pan 

Swatlemchloa 

Pan 

Muriocladzu J 

Notyet 

known 

08 Pan 

Pan Aubmmia A 

Spi 

Rae 

Div 

see 

Spi 

Sol 

RE c 

_I 

(See 

- descr, 

&&, 

- 

- 

Arthrosty lidhm 

Not yet known 

Not yet known [ 

f 

D if 

Pan 

Not. yet known 

Dig 

&p 
The following terms are defined with special 

reference to the usages adopted by the author for 

describing various characteristics of the bamboo 

plant. Most of these first appeared in the glossary 

of McClure (1966b:295-318), reprinted in Taxon 

(McClure 1966a: 220-235). In that glossary newly 

coined terms were indicated by an asterisk (*); 

others, indicated by a dagger (f), were given new 

or extcnded meanings to describe conditions not 

satisfactorily covered by conventional usage, or by 

botanical glossaries. 

Some of the following definitions incorporate 

more 01 less drastic modifications that, in the present dissertation, are given precedence over the 

definition of the same terms that appear in the 

above publications. For example, the content given 

the terms monoclade and pleioclade in the earlier 

glossary has been drastically changed. When unqualified (standing alone) these terms now refer 

exclusively and to the first order element (s) of a 

branch complement. The qualifying adjectives “restricted” and “unrestricted” refer to their potential 

for proliferation from buds (if any exist) at the 

proximal nodes of their first order elements. Any 

modified, or newly added, terms are indicated by a 

double asterisk (”*) in the present glossary. 

ABAXIAL (L. ab, away from; axis). Away from 

an axis; designates that surface of an appendage 

(such as a culm sheath or a sheath blade) which 

faces, in a structural sense, away from the axis on 

which it is inserted or relies for support. The terms 

abaxial and adaxial (q.v.) should be employed 

instead of upper and lower, or dorsal and ventral, 

whose use-with reference to the two faces of a 

foliage leaf or a reflexed sheath blade, for exampleis apt to perpetrate ambiguity, and so cause confusion. This is because the orientation of such a 

structure may vary-in time, and from one botanical 

entity to another-especially among the bamboos. 

Therefore, neither upper and lower, nor dorsal and 

ventral, will always be correlated reliably with 

Glossary 

122 

anatomical dorsiventrality, to which important 

structural basis all pertinent observations must be 

oriented unmistakably in order to satisfy the requirements of scientific discipline, dependability, 

and clear thinking. Cf. ADAXIAL. 

ABSCISSILE (L. abscissilis, from abscissus, cut off). 

Susceptible of being cut off or disarticulated by the 

formation of an abscission layer; applies to the 

petiole of a deciduous leaf or, as in many bamboos, 

to the segments of a rachilla. See DECIDUOUS. 

ACROPETAL (Gr. ak~on., the highest point; L. peto, 

I go toward). Progressing toward an apex, as when 

the development or emergence of appendages on 

any axis takes place serially in the direction of the 

apex of the axis. Cf. BASIPETAL. 

ACROPETALLY (adv.) Toward an apex. 

ADAXIAL. Toward an axis; designates that surface of an appendage which faces, in a structural 

sense, toward the axis on which it is inserted, or 

relies for support. Cf. ABAXIAL. 

“ADNATE (L. adnascor, I grow to). “Attached 

the whole length” (Jackson, 1949). “Congenitally 

grown together; said especially of unlike parts” 

(Webster, 1959). Used herein as applying to any 

degree or extent of union between the tissues of 

unlike parts. See CONNATE. 

* ‘ADVESTITIOUS (L. adventitius). Occurring in 

a location other than the usual one. In the bamboos, and in other grasses as well, the principal 

complement of roots is adventitious, arising at the 

nodes of culms and rhizomes, and not from the 

primordial root. In the genus Chusquea; for example, adventitious branch buds arise independently, 

both at the right and at the left of the one at the 

usual site, which is median to the base of a subtending culm sheath. See CONSTELLATE. 

““AMPHIMORPH (Gr. amphi, both; morphe, 

forms). Embracing both of two distinct forms. 

Example: The same plant of Chusquea fendleyi 

gives rise to culms from both pachymorph and leptomorph rhizomes. 
NUMBER 9 123 

* *ANANTHOUS (L. anathus). Lacking a flower; 

“transitional glumes,” and “sterile lemmas” are 

ananthous. 

(Gr. anastomGsis, a new outlet). 

“Union of one vein with another, the connection 

forming a reticulation” (Jackson, 1949). 

ANDROECIUM (Gr. andros, male; oikos, house). The 

stamen complement of a flower. I have not followed 

Arber (1934:152 et passim) in adopting the spelling (andraeceum) advocated by Kraus (1908). R. 

W. Brown (1954), Jackson (1949), and Nybakken 

(1959) all give androecium. Cf. GYNOECIUM. 

* “ANTHECIUM (Gr. anthos, flower; oikos, house). 

The protective structure in the Gramineae, which 

is formed by the lemma and palea and which, typically, encloses a single flower. 

ANTICALLY, anticously (L., forward, in front). 

Toward the front; introrsely or adaxially. Jackson 

(1949) records diverse and even antonymous definitions oi antical, mentioning the synonymous use of 

introrse as “occasional.” The bamboo palea often 

gapes antically. 

* *APOCLADA (Gr. apo, separate; klados, branch). 

A bamboo genus; the name alludes to the independent origin of the primordial elements of midculm branch complements, a feature that appears 

in distinctive expression in this taxon. 

In convex arcuate 

array; characterizes the pattern of insertion of the 

secondary component (second-order axes) of a monoclade branch complement, as exemplified by 

members of the genera Merostachys and Rhipidocladum, qq. vv. See CONSTELLATE; GREMIAL; LEVEL; 

RESTRICTED. 

ASSIMILATION TISSUE. Mesophyll; the interior 

ground tissue of a leaf blade. 

AWNED. Bearing a bristlelike apical appendage 

called an awn (OE). The empty glumes of the 

spikelet in the type and some other species of 

Neurolepis are awned. In Streptochaeta (a genus 

of grasses related to bamboos), the lemma terminates above in a very long, apically coiled structure 

called an awn (L. arista) by Nees (1834), the author 

of the genus; Trinius called it a tail (L. cauda). 

The distinctions between the ways in which the 

different types of sheathing structures or their foliar 

appendages terminate apically are not sharp. Technical terms (such as apiculate, awned, cuspidate, 

and mucronate) and their conventional definitions 

do not clearly distinguish the diverse expressions 

ANASTOMOSIS 

““APSIDATE (L. apsis, arch). 

that are a part of the morphological intergradation 

that baffles efforts at hard-and-fast categorization 

here, as in many other facets of plant structure. 

BAMBOO (bumbu, a vernacular word of undetermined Oriental origin). The wood of bamboo 

culms; a plant so classified. 

BAMBOOS. A taxonomic group of plants comprising the tribe Bambuseae of the Bambusoideae, 

a subfamily of the Gramineae; living plants, or 

culms (stems) severed from plants of this group. 

BASIPETAL (L. basis, base; peto, I go toward). 

Progressing toward a base, as when the development or emergence of appendages on any axis takes 

place serially in the direction of the base of the 

axis. Cf. ACROPETAL. 

BASIPETALLY (adv.). Toward a base. 

* “BINATE (as applied to pleioclade midculm 

branch complements). Typically consisting of only 

two axes or branches, e.g., typical midculm branch 

complement in the Old-world genus Phytlostachys. 

BLADE (AS. blued, leaf). “The limb or expanded 

portion of a leaf” (Jackson, 1949). In the bamboos, 

a thin, expanded, chlorophyll-bearing, sessile or 

petiolate, apical appendage (lamina) of a sheath 

proper. See LEAF. 

BOTTLEBRUSH (bottle brush, a brush of cylindrical shape, with the bristles uniformly distributed on 

the axis and oriented approximately at right angles 

to it). A borrowed term, given a slightly modified 

form, and used here (in the adjectival sense apparently fiot recorded in dictionaries) to describe the 

approximate pattern of distribution and orientation 

of the proliferations of the stigmatic surface in certain bamboos, a pattern loosely called plumose or 

aspergilliform by some authors. Cf. DENDROID. 

**BRACT (L. bructea, a thin plate of metal). A 

foliar organ of the rachis of bamboo inflorescences. 

Circumcingent bracts subtend branch buds and 

branches of the rachis in all bamboo genera with 

iterauctant inflorescences, but at one or more of the 

basal nodes immediately above the prophyllum they 

are usually empty. Bracts are usually rudimentary 

or obsolete in most bamboo genera with semelauctant inflorescences; in Glaziophyton, however, they 

are well developed and circumcingent. 

““BRANCH COMPLEMENT.he monoclade or 

pleioclade array of primary branches (first-order 

axes) that develops at a single midculm node. A 

complete definition of each recognized type of 

branch complement includes (1) the mode or pat- 
124 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tern of insertion of its primary (or secondary) components or axes (see INSERTION); and (2) its 

potential (restricted or unrestricted) for proliferation from buds at the proximal nodes of its primary 

(first-order) component (s). 

BRANCH SHEATHS. The sheathing organs borne 

singly at each node of an aerial vegetative branch 

of any order (except the culm itself), excluding the 

neck sheaths and the leaf sheaths. 

“The nascent state of a flower or branch” (Jackson, 1949). In the usage here 

adopted, the term bud is applied only to those 

primordial vegetative or reproductive branches that 

(1) are enclosed in a prophyllum, and (2) have a 

resting stage. Those (such as branch primordia in 

determinate inflorescences generally; and root primordia, for example) that lack either one or both 

of these features are referred to simply as primordia. 

Branch primordia in the determinate inflorescences 

of Glaziophyton, Greslania, and some other genera 

are prophyllate, but they apparently have no resting stage. 

**BUD, PRIMARY. A primary bud is one whose 

initial primordium produces the primary (firstorder) element (axis) in any series of branches of 

ascending order. It appears that, in the members 

of some bamboo genera a solitary primary bud may 

incorporate more than one initial primordium, and 

that each of these initial primordia normally produces an independent first-order element of a pleioclade branch complement. See PRIMORDIUM, INITIAL. 

**CALLUS. A growth consisting internally of 

iindifferentiated pith-like parenchyma that in some 

bamboos forms an external ridge at the junction of 

two structures (as, for example, at the locus of 

insertion of a sheath upon its axis). Holttum (1958: 

15, fig. 3) applies the term to what Hackel (1899: 

716) calls the “exterior (outer) ligule” of the leaf 

sheath, and what Pilger (in Diels 1937:57) calls a 

collar. In that structure the tissue is usually of a 

firmer nature, and is brought into prominence by 

the abscission of the petiole, See CUPULE; GIRDLE; 

and “outer ligule” under LICULE. 

**CARYOPSIS (Gr. karyon, a nut; opsis, likeness). 

“A one-celled, one-seeded, superior fruit, with a 

pericarp united to the seed” (Jackson, 1949, as 

cariopsis). A caryopsis is indehiscent, i.e., the pericarp does not open to liberate the seed. In some 

bamboos (Melocanna baccifera, for example) the 

seed is free from the pericarp at maturity. The 

BUD (ME. budde). 

separability of the pericarp from the seed has been 

much used by agrostologists as a taxonomic feature, 

but mature caryopses of bamboos are rarely available for comparison. 

**CATAPHYLL (Gr. kata, down; phyllon, leaf). 

“The early leaf forms of a plant or a shoot, as cotyledons, bud-scales, rhizome scales, etc.” (Jackson, 

1949). “Any rudimentary scalelike leaf which precedes the foliage leaves, as a bud scale, etc.” (Webster, 1959). As herein used, the term designates the 

sheaths that are incident at the lowermost nodes of 

any vegetative axis of a bamboo plant. In cataphylls 

the sheath proper is smaller in all dimensions than 

it is in the sheaths borne at the more distal nodes 

of the same axis, and its appendages are all rudimentary or obsolete. 

CENTRIFUGALLY (L.centrum, center; fugo, I flee 

from). Outward from a center. 

CENTRIPETALLY (L. centrum, center; peto, I go 

toward). Toward a center. 

**CESPITOSE (L. caespes, a sod; caespitosus, forming a more or less compact clump or tuft). Describes the normal clump habit of bamboos with 

pachymorph rhizomes, except where the rhizome 

neck Is very much elongated and the primary culms 

proliferate by tillering or by way of short-necked 

rhizomes as in the Old World bamboo currently 

known as Yushania niitakayamensis. See DIFFUSE; 

PLURICESPITOSE; UNICESPITOSE. 

**CHARACTER (L.). “An instrument for marking, character” (Webster, 1959). “The technical 

difference whereby allied forms are distinguished, as 

ordinal, generic, specific, and so on” (Jackson, 

1949). “A member of mutually exclusive dates” 

(Kendrick and Proctor, 1964:66). Any distinctive 

form 01 behavior, or characteristic feature of demonstrrible reliability (constancy) for purposes of 

recognition or for the composition of a formal description or for the construction of a key for the 

differentiation of similar or related taxa. Constant 

features serve in some cases alone as recognition 

characters, in other cases as members of a set of 

regularly associated constant features. Webster’s 

definition of the term character, as used by biologists (“Bot. & Zool. A feature or peculiarity (as of 

structure, color, form or size) possessed by the individuals of a variety, species or other category, 

and by means of which they may be recognized or 

differentiated; as a specific character, a generic character, etc. An inherited character is regarded as the 
NUMBER 9 

observable effect of a gene or genes”) makes feature 

a synonym of character. I differentiate these two 

terms with the objective of promoting clear thinking in relation to taxonomic procedures. It is my 

position that a feature doesmot become a character 

in the taxonomic sense until it can be shown to be 

dependable for use in the differentiation of two similar or related taxa. See FEATURE. 

CHARTACEOUS (L. charta, paper; -aceus, made of, 

or belonging to). Paperlike; papery. 

CIRCUMAXIAL (L. circum, around; axis). Completely around an axis; describes the typical reach 

of the locus of insertion of a sheathing appendage, 

or an assemblage of adventitious roots. Jackson’s 

(1949) definition, under Circumaxile, is limited to 

the special sense, “surrounding a central axis which 

separates when the fruit splits open.” 

“*CIRCUMCINGENT (L.circum, around; cingens, 

girding). Characterizes a sheathing appendage (e. 

g., a bract or a prophyllum) with respect to the 

reach of its locus of insertion when this completely 

encircles an axis. 

CIRCUMNUTATION (L.circum, around; nutans, 

wavering). The circular movement that commonly 

takes place in the distal portion of a growing axis, 

causing the apex to follow a spiral path as elongation proceeds. 

**COLLARET (Arber 1934:309). The girdle or 

callus at the base of a leaf sheath. See CALLUS; 

GIRDLE. 

**CONGRUENT (L. congruens, conformable). In 

agreement; consistent; harmonious. Applied to 

herbarium specimens, of different but allied bamboo species, which show a range of corresponding 

characters that can be compared. Incongruent specimens show different ranges of characters and thus 

are sometimes useless as evidence of distinctions between species. 

CONNATE (L. connatus, born at the same time). 

“United, congenitally or subsequently” (Jackson, 

1949), in reference to the union of members of a 

set of homologous parts (such as the filaments in 

a stamen complement) that takes place in some 

bamboos. See ADNATE; DIADELPHOUS; MONADELPHOUS; TRIADELPHOUS. 

CONNECTIVE (L. connectivum). A very narrow 

strip of tissue upon the approximately opposite 

sides of which the two parts of an anther (that is, 

the two pairs of locules) are inserted longitudinally. 

Jackson (1949) describes the connective as “distinct 

from the filament,” of which it appears to be an 

apical extension. 

* “CONSTELLATE (L. constellatio, a cluster of fixed 

stars; a constellation). Arranged in a constellationlike cluster; characterizes (1) the pattern of insertion of the primary components (first-order axes) 

of a pleioclade branch complement as exemplified 

by members of the genus Chusquea (where the 

central element is several times as large as the 

auxiliary ones that flank it), as well as where component first-order elements are subequal, as in 

Athrocstachys; and (2) the pattern of insertion of 

the bud complement from which such a branch 

complement arises. See APSIDATE; GREMIAL; LEVEL; 

RESTRICTED. 

CORNIFOKM (L. cornu, horn). Shaped like a horn. 

CULM (L. culmus, stalk, stem). A segmented 

aerial axis that emerges from a rhizome, and forms 

a part of a gramineous plant; the term is used most 

commonly with special reference to bamboos. Syn.: 

halm, haulm haum. 

“The ‘knot’ of a 

grass stem” (Jackson, 1949). The term node is generally applied in a loose, comprehensive sense to 

that complex locus-the junction of adjacent internodes in a segmented axis of a gramineous plant. 

Hackel (in Lamson-Scribner and Southworth, 1890: 

2, 3) stressed the importance of the “constantly overlooked” difference between the culm nodes and the 

sheath nodes in non-bambusoid grasses. However, 

neither Hackel’s description nor his illustration 

does justice to the distinctive features of these structures as they appear in the bamboos. Culm node is 

here defined with special reference to that level 

within the node (sensu lato) where secondary elongation (intercalary growth) takes place, and where 

branch buds and adventitious roots are inserted. 

Each culm node (sensu stricto) is located just above 

a sheath node (q.v.), from which it is usually distinguishable by the transverse thickening or ridge 

(the “supranodal ridge” of some authors) that appears at the level of insertion of a bud or a branch 

comp!ement. Branch node and rhizome node should 

be similarly differentiated, respectively, from the 

corresponding sheath nodes. See SHEATH NODE. 

One of the sheathing organs 

borne singly at each node of a bamboo culm, below 

the level at which the sheaths of foliage leaves take 

their place. Kurz (1876:268) anticipates this usage 

with “halm-sheath.” The transition between culm 

CULM NODE (L. nodus, knot). 

**CULM SHEATH. 
126 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

sheaths and foliage leaves may be rather abrupt or 

very gradual. 

CULM SHOOT. Bamboo shoot; a young culm in 

any stage of its development short of maturity in 

height. See SHOOT. 

* “CUPULE. The cup-shaped depression that 

marks the locus of insertion of the petiole upon 

the leaf sheath (McClure, 1941:33-34). The cupule 

is apparent, of course, only after the leaf has fallen 

away. See CALLUS; LIGULE. 

* *CUSPIDATE (L. cuspidatus, from cuspis, a point, 

especially the point or head of a spear [Marchant 

and Charles, 19581). “Tipped with a cusp” (Jackson, 1949). Terminating in a small, hard point; 

used herein in this sense to characterize the apex 

in bamboo caryopses where the indurated conical 

base of a slender style persists at maturity. 

**CYMBIFORM (L. cymbiformis; cyrnba, boat). 

Boat-shaped, like the lemma in many bamboos, especially when it is keeled. 

“DECIDUOUS (L. deciduus). Abscissile; falling 

away; “falling in season” (Jackson, 1949). The term 

applied especially to organs (the culm sheaths in 

some bamboos) or parts that become cut off at or 

near the locus of their insertion by development of 

a special layer of cells (an absciss layer), or by the 

formation of a cleavage plane along a well-defined 

line of separation. In many bamboos several of the 

distal internodes of leafy twigs (as well as the individual leaves) are deciduous. They fall away after 

the leaf blades have fallen, upon the formation of 

a transverse rift at a node-usually the most distal 

node bearing a bud-this bud having already produced a new leafy twig. See PERSISTENT. 

DECLINED (L. declinatus, turned aside). “Directed obliquely” (Jackson, 1949); describes the habit 

or orientation of the culms in some species of 

bamboos. 

DELIQUESCENT (L. deliquescens, melting away, 

disappearing). Becoming dispersed or expended, 

as when the main axis of any structure (the rachis 

of an inflorescence of Aulonernia quexo, for example) loses its identity among its own branches, or 

“loses itself by repeated branchings” (Jackson, 

1949). See EXCURRENT. 

DENDROID (Gr. dendron, tree, -oid, having the 

form of). Treelike; bearing branches of more than 

one order; describes the pattern of proliferation of 

the stigmatic surface in certain bamboos. 

*#DEPAUPERATE (L. depauperatus, impoverished). Reduced in size or functional efficiency, or 

both; the term characterizes whole plants, or individual parts or organs thereof such as florets, pseudospikelets, whose development is impeded or restricted by adverse conditions, external or internal. 

DERMOGRAMME (Gr. derrna, skin; gramrna, something drawn). A French term used by Prat (1936: 

178) ta designate any example of his diagrammatic 

illustrations of the cellular details of the structure 

of the epidermis of any grass or bamboo. See 

SPODOGRAM. 

DETERMINATE (L.determinatus, having fixed limits). Of limited growth; characterizes an axis, or a 

system of immediately related axes (a determinate 

inflorescence, for example), whose development or 

potential for development is confined to a single, 

definitely limited “grand period of growth,” after 

which no meristem usually persists. The occasional 

persistence of meristem at the tip of individual 

rachillas sometimes prolongs the grand period of 

growth in a determinate inflorescence atypically. A 

rachilla may then continue to produce new florets 

until the spikelet is greatly elongated-to as much 

as a foot, while its normal length would be 3 to 4 

inches. Again, this meristem may revert to the 

vegetative state, prolificating to produce a leafy 

axis. Both of these teratic phenomena have been 

observed in Arundinaria dolichantha. See PROLIFERATION; SEMELAUCTANT. Cf. INDETERMINATE. 

DIADELPHOUS (Gr. dis, twice; adelphos, brother). 

Having “two groups of stamens”; cognate with Diadelphia, the name of “a Linnean class having the 

stameiis in two bundles of brotherhoods” (Jackson, 

1949). Diadelphous describes the condition of a 

stamen complement of a flower when its members 

have their filaments connate to form two distinct 

groups. In the bamboos, the relative numbers of 

stamens in the respective groups may vary from 

one genus to another. Cf. MONADELPHOUS; TRIADIAGEOTROPIC. Adjectival form derived from 

diageotropism, which is defined under GEOTROPISM. 

DIAPHRAGM (Gr. diaphragma, a partition wall). 

The transverse internal layer of parenchyma found 

at the level of every sheath node. It is reinforced by 

the crossing over and anastomosis of vascular bundles, and forms a rigid structural element that lends 

strength (mechanical resilience) to the segmented 

vegetative axes of bamboos. “A dividing membrane 

or partition” (Jackson, 1949). 

DELPHOUS. 
NUMBER 9 127 

““DIFFUSE (L. diflusus, spread out). Growing in 

open array; characterizes the normal mature clump 

habit of most bamboos with leptomorph rhizomes, 

and also those whose pachymorph rhizomes have a 

greatly elongated neck and solitary culms, e. g., 

Bambzisa sp. (McClure 21438-A) of Peru. (See 

McClure, 196Gb:27, fig. 10:5). 

DISTAL (L. disto, I stand apart). Remote; far 

out; designates loci of insertion, or structures, situated at or near the tip of an axis. Cf. PROXIMAL. 

DISTICHOUS (L. distichus, consisting of two rows). 

“Disposed in two vertical ranks, as the florets in 

many grasses” (Jackson, 1949). Distichous also describes the pattern of insertion of buds and 

branches, and the sheathing appendages on all segmented axes, both vegetative and reproductive, of 

the bamboo plant. As tacitly recognized by agrostologists, but generally not mentioned in published 

definitions, the two ranks are normally inserted on 

opposite sides of an axis but are sometimes secund. 

DISTICHY (substantive of L. distichus). Arber’s 

(1934) word for the two-ranked arrangement of 

buds, branches, and sheathing appendages generally characteristic of segmented axes in the Gramineae. See DISTICHOUS. 

““DOMINANT (L. dominans). Superior in size 

among several associated objects of a class or kind, 

as the primary axis among members of a pleioclade 

midculm branch complement in Bambusa vulgaris, 

or the trees that overtop all others in a forest. 

* “EMBRYOTEGIUM (Gr. embryon, a foetus, tega, 

a covering). The embryotega of Stearn (1966). 

Literally, embryo cover; the “little shield” (Sp. 

escudete) of Parodi (1961); to be distinguished from 

the scutellum, which is a part of the embryo itself. 

“A callosity in the seed coat of some seeds near 

the hilum, and detached by the protrusion of the 

radicle on germination” (Jackson, 1949). Since 

Jackson’s definition is based on the nature of the 

embryotegium in the seed proper, it is not strictly 

applicable to its nature in a caryopsis. Hackel (in 

Lamson-Scribner and Southworth, 1890: 20, fig. 6) 

describes and illustrates the embryotegium in a 

caryopsis (without, however, giving it a technical 

name) as “a place where the embryo lies covered 

only by the pericarp, and plainly visible on the outside.’’ The position of the embryotegium is clearly 

visible in caryopses with a thin pericarp as in bamboos of certain genera such as Bambusa and Arundinaria. In some genera, including Meiocanna (Stapf, 

1904a, pl. 45: fig. 3, pl. 47: fig. 40), and Ochlandra, 

where the pericarp is thick, the embryotegium is 

ordinarily not externally recognizable (cf. Figure 

““EMPTY GLUMES. Revision of McClure’s 

(1966a:ZZj) definition. See TRANSITIONAL GLUMES. 

* “ENDOSPERM (Gr. endo, within; sperma, seed). 

“The nutritive tissue formed within the embryo sac” 

[of a seed] (Webster, 1959)-referred to by earlier 

writers as “albumen” (Jackson, 1949). In some 

bamboos the embryo consumes the endosperm as 

fast as it is formed (Stapf, 1904a). The stored endosperm in some bamboos (Arundinaria gigantea) is 

granular and starchy, in others (Apoclada simplex) 

it appears to be waxy or glutinous, not granular. 

EXCURRENT (L. excurrens, running out). Extending through, as when the main axis of any 

structure (the rachis of the paniculate inflorescence 

in Indocalumus sinicus, for example) maintains its 

identity among its own branches; “where the stern 

remains central, the other parts being regularly 

disposed round it” (Jackson, 1949). Cf. DELIQUES- 

““FACULTATIVE (Fr. facultatif, from L. facultas, 

capability). “Of such a character to admit of existing under various forms or conditions; of happening or not happening, etc.” (Webster, 1959). 

“Occasional, incidental, as opposed to obligate” 

(Jackson, 1949). Describes the freedom, or potential 

faculty, to give expression to one or the other of 

two or more states, forms, products or features; 

possessed by (1) the proximal buds of the primary 

branch in certain bamboos with respect to germination or continued dormancy (Aulonemia effuso) (see Key, p. 8); or (2) the apical meristem of 

the developing axes in certain bamboos (Arundinaria gigantea ssp. tecta; Semiarundinaria fastuosa) 

to produce either a culm or a rhizome. 

““FEATURE (ME. feture; L. factura). “The . . . form or outward appearance of anything” (selected 

from Webster, 1959). In conventional discourse 

(and in technical papers as well) the term “feature” 

is often used in the sense here given “structure” 

(e.g., eyes, nose, mouth, chin-individual component structures of a countenance). In the present 

work the technical meaning of feature is restricted 

to characteristic aspects (such as form, size, number, 

color, behavior) of various structures or parts thereof. It is my position that a feature does not become 

a cliaracter in the taxonomic sense until it can be 

37 0). 

CENT. 
128 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

shown to be invariable (dependable) and either 

unique to a given taxon or a member of a set of 

contrasting features that distinguish two similar or 

related taxa. Cf. CHARACTER. 

Numerous, 

slender, and not conspicuously tapered; the meaning of the term as conventionally used to describe 

the gross aspect of the roots of bamboos and other 

grasses, a definition apparently not recorded in dictionaries or glossaries. Jackson’s (1949) definition 

of the term is not applicable here. 

FISTULOSE (L. fistula, a pipe). Hollow; having 

a lumen; characterizes the internodes of culms and 

branches in most bamboos. 

FLORET (dim. OFr. flor, flower). One of the units 

into which a spikelet breaks up when the rachilla 

segments disarticulate. Regardless of whether the 

rachilla segments are abscissile or not, a floret consists of (1) a segment of the rachilla, (2) the lemma 

that is inserted upon it, (3) a branch (the axis of 

a flower) subtended by the lemma, (4) a prophyllum (the palea) of the axis of the flower, and (5) 

the parts of the flower that are inserted on its axis. 

Attention is directed especially to the fact that the 

gramineous floret includes structures from axes of 

two oi-ders while the flower is confined to a single 

axis. The distinction between floret and flower is 

not always observed in published works on agrostology. The loose use of the two terms interchangeably is to be avoided. Cf. FLOWER. 

FLOWER (OFr. fEor). That portion of a branch of 

the rachilla that is distal to its own prophyllum 

(the palea), together with the reproductive organs 

(androecium or gynoecium or both) borne by it. 

The lodicules, when present, are included in this 

concept of the flower, but the palea is not. Cf. 

FLORET. 

A leafy or leafless segmented axis that bears one or more inflorescenses. A flowering branch is distinguishable from an inflorescence proper in that it retains all of the morphological characteristics of the vegetative state, with the 

sole exception that in some cases it does not produce foliage leaves. The inflorescence proper, on 

the other hand, always has morphological features 

peculiar to it that are not found on the flowering 

branch proper. In describing and illustrating bamboos, some authors fail to distinguish the two. Arber 

(1934: 108) characterizes a leafless flowering bamboo 

FIBROUS (L. fibra, thread, filament). 

FLOWERING BRANCH. 

as a “truly gargantuan inflorescence.” Cf. INFLORESCENCE. 

FOLIAR ORGAN. See SHEATHING ORGAN. 

““FREE (ME. fre). “Not adhering, the reverse 

of adnate” (Jackson, 1949). Here given the sense of 

non-connate, in reference to the filiform filaments 

of the stamens in certain bamboos. 

FUGACEOUS (L. fugax, fleeting). Promptly deciduous; “soon perishing” (Jackson, 1949). 

FUSIFORM (L. fusus, spindle). Spindle-shaped; 

circular in cross-section, thickest in the middle and 

tapering toward each end. See SUBFUSIFORM. 

GENICULATE (L. geniculatus, with bended knee). 

“Bent abruptly at an angle, like the bent knee” 

(Webster, 1959). When a growing culm adjusts the 

direction of its orientation (as in situations where 

negative geotropism operates to effect the restoration of a fallen or deflected culm to an upright 

postuie), the adjustment takes place through differential elongation in the upper and lower portions, 

respectively, of the zone of intercalary growth 

immediately above a node, and the node involved 

becomes geniculate. 

GEOTROPISM (Gr. ge, the earth; plus tropism, 

defined by Webster (1959) as “the innate tendency 

of an organism to react in a definite manner to 

external stimuli”). A physiological potential, and 

the consequent act, involved in the assumption and 

maintenance of a particular direction of growth by 

an axis or organ with reference to the force of 

gravity, or to the centrifugal force generated by 

rapid circular motion. As commonly used, geotropism ambiguously embraces (and confuses) two 

component phenomena: the visible physical response, and the more fundamental, invisible 

response, which is physiological. Jackson’s (1949) 

definition: “the force of gravity as shown by curvature in nascent organs of plants” is incomplete. 

Three distinct classes of geotropic responses are 

recognized: positive, negative, and transverse. The 

last named is called diageotropism. 

**GERMINATION (L.germen, an offshoot, sprout, 

bud). A term usually restricted to the beginning of 

growth of the embryo of a seed; here extended to 

the beginning of growth of a temporarily dormant 

axillary bud on a bamboo culm. 

*“GIRDLE (AS. gyrdel, belt: Latin equivalent, 

cingulum). A conspicuous horizontal band of special tissue occupying a position basal to the culm 

sheath proper in some bamboos (e.g., Melocalamus 
NUMBER 9 129 

compactiflorus and some scandent species of Chusquea). It is expansible in all directions as long as 

the corresponding intercalary zone of the embraced 

internode is in active growth, and it takes on an 

asymmetrical form at geniculate nodes. Upon the 

abscission of the sheath, this band remains attached 

at the culm nodes. The distinction between a girdle 

and a sheath callus (q.v.) is not always clear, being 

based principally upon their relative vertical dimension (area). Cf. CALLUS; SHEATH CALLUS. 

**GLUME (L. gluma, a hull or husk). “The 

chaffy, two-ranked members of the inflorescence of 

grasses and similar plants” (Jackson, 1949). Excepting prophylla, this very general definition ostensibly embraces all of the sheathing appendages of 

the inflorescence, including the bracts that, in some 

cases, subtend its branches. As shown elsewhere, the 

application (meaning and usage) of the terms 

lemma and palea and bract has become rather 

generally standardized. However, in those cases 

where it is difficult to distinguish empty glumes and 

sterile lemmas from each other, I have propose‘d to 

cover both categories with the comprehensive term 

transitional glumes, q.v. 

GREGARIOUS (L. gregarius, of or pertaining to a 

flock or herd). “Growing in company” (Jackson, 

1949). Gamble (1896:viii) makes gregarious mean 

simultaneous, to describe the flowering behavior 

manifested when all of the members of a given 

generation of bamboo plants (from seeds of a common origin) enter the reproductive state at approximatelv the same time. Cf. SPORADIC. 

**GREMIAL (L. gremialis, of a lap or bosom). 

“Growing in a pollard-like cluster” (R. W. Brown, 

1954); characterizes the mode of insertion of a tuft 

of subequal second-order axes that develop from 

buds OR a decurrent extension of the primary (firstorder) axis of an unrestricted monoclade branch 

complement (as in Bambusa swalleniana [Guadua 

spinosa] and in numerous species of the genus 

Aithrostylidium), Cf. PROMONTORY. 

GYNOECIUM (Gr. gynb, female; oikos, house). 

The complete pistil, consisting of the ovary, the 

ovule (s), style (s), and the stigma (s), of a single 

flower. I have not followed Arber (1934:120 et 

passim) in adopting the spelling (gynaeceum) advocated by Kraus (1908). Jackson (1949) and Nybakken (1959) both give gynoecium. Cf. ANDROECIUM. 

HALM, haulm, haum (terms used in British agrostology). See CULM. 

**HERMAPHRODITE. Same as PERFECT, q.v. 

**HILUM (L. hilum, a trifle). “The scar left on 

a seed where formerly attached to the funicle or 

placenta” (Jackson, 1949). This definition confines 

the connotation of the term hilum to a scar on a 

deciduous seed that has been freed from its attachment on a dehiscent carpel. However, in the 

literature of agrostology, the term hilum is quite 

generally applied to the longitudinal furrow or 

groove that marks, on the exterior of the indehiscent gramineous caryopsis, the locus of attachment 

of the indeciduous seed that remains confined within an indehiscent pericarp. In the present treatise, 

this external furrow is referred to as a sulcus. See 

SULCATE. 

IMBRICATE (L. imbricatus, covered with tiles). 

“Overlapping like roofing tiles and shingles” (R. 

W. Brown, 1954). Sheathing organs at successive 

nodes of an axis are imbricate when each one exceeds, in length, the internode it embraces. 

INDETERMINATE (L. indeterminutus, unlimited). 

“Not terminated absolutely” (Jackson, 1949). Continuing apical growth characterizes an individual 

axis as indeterminate. The term is here given another connotation (namely, iterauctant) with particular reference to the continuing ramification of 

a system of immediately related axes, as exemplified 

by the indeterminate inflorescences of certain bamboos. In such infloresences, the buds at the proximal nodes of each new order of branches bear fresh 

bodies of meristem. The development of new 

branches from these buds may be continuous (or 

intermittent and reactivated on a seasonal basis) 

for a period lasting in some cases for several successive years. Each branch of an indeterminate inflorescence (an axis with its appendages) has the 

appearance of a spikelet. Because of this, it is 

called a pseudospikelet. Each pseudospikelet has 

its own independent grand period of growth. The 

active period of the meristem at the apex of the 

rachilla that terminates each pseudospikelet is 

limited. Thus, while the branching of indeterminate bamboo inflorescences is of a continuing 

nature, each individual branch is apically determinate in its growth. See ITERAUCTANT; PSEUDOSPIKELET; cf. DETERMINATE. 

**INFLORESCENCE. A discrete aggregation of spikelets associated with a common primary rachis or a 

common peduncle. Exceptionally, an inflorescence 

may comprise but a single spikelet. I know of no 
130 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

exception to the rule that the inflorescence in a 

given bamboo genus is uniformly either semelauctant or iterauctant (q.v.) in its manner of development. The occurrence of individual features of an 

intermediate (incongruous) nature is rare. Cf. 

FLOWERING BRANCH; see also INFLORESCENCE, ITERAUCTANT; INFLORESCENCE, SEMELAUCTANT. 

* *INFLORESCENCE, ITERAUCTANT. An inflorescence ir, which typically both the initiation and the 

termination of growth takes place in serial sequence 

in the flowering axes of successively higher orders, 

each branch possessing residual meristematic tissue 

in the form of basal branch primordia (buds) that 

continue the expansion of the inflorescence as long 

as physiological conditions in the adjacent tissues 

are favorable to their germination and growth. 

**INFLORESCENCE OF MIXED BRANCHING. An inflorescence with peduncles proliferating (duplicating themselves indefinitely from nascent or dormant 

prophyllate branch buds) the rachises of determinate branching (without nascent or dormant 

branch buds) and with the development of all 

spikelets simultaneous or nearly so (e.g., the Old 

World genus Chimonobambusa). 

* “INFLORESCENCE, SEMELAUCTANT. An inflorescence in which typically the initiation of growth is 

simultaneous or nearly so in all axes, and the termination of growth is likewise simultaneous or nearly 

so; and no meristematic tissue remains afterwards 

in the form of dormant buds. 

#*IKSERTION (L. insertio, a putting in). The 

mode, pattern, or locus of attachment of an organ 

or a part. With a view to characterizing the several 

patterns of insertion herein distinguished with respect to the component first- (or second-) order axes 

of a midculm branch complement, the following 

terms have been selected and defined: APSIDATE, 

CONSTELLATE, GREMIAL, LEVEL, and RESTRICTED. 

* WITERAUCTANT (L. iterum, again; auctans, growing). Embracing more than one grand period of 

growth, as the mode of development of the successive orders of the branches of an inflorescence composed (as in bamboos of the genera Bambusa, 

Elytrostachys, and Atractantha) of pseudospikelets 

(q.v.). Cf. SEMELAUCTANT; INDETERMINATE. 

**LEAF (AS). Foliage leaf; bamboo leaf; the 

chlorophyll-bearing, usually petiolate, blade of a 

leaf sheath proper. This concept of leaf, in the 

sense bamboo leaf, includes the petiole when it is 

present, but excludes the leaf sheath proper. Leaf 

is here defined in this circumscribed sense in order 

to avoid a commonly encountered ambiguity occasioned by its indiscriminate use in the literature. 

The word is commonly used by agrostologists, without modification or qualification, to refer to any 

one of the morphologically, anatomically, and functionally divergent forms of sheathing organs borne 

on the several kinds of vegetative axes of a bamboo 

plant. This imprecise use of the term, in disregard 

of the importance of the manifest differences thus 

loosely covered by it, apparently is due to the 

generalized and unqualified nature of the definition 

of “leaf” given by Jackson (1949): “the principal 

appendage or lateral organ borne by the stem or 

axis.” The use of the term “leaf” for the petiolate 

leaf blade as distinct from the leaf sheath in bamboos h2s ample precedent in Munro’s “folia” which 

is described separately from “vagina.” I think that 

we may rationalize this usage by postulating that, 

in the grasses, the sheath is a distinct organ or 

structure inserted between the leaf proper and the 

axis frcm which it emerges (or on which it is inserted). We may continue with the postulate that 

the girdle (where it appears) is a structure inserted 

between the sheath proper and the sheath node. 

Cf. SHEATH; LEAF SHEATH; see also SHEATHING OROne of the leaf-bearing sheaths 

inserted at the distal nodes of each aerial segmented 

axis whether culm, branch, or twig. The conspicuous part of a leaf sheath is the usually petiolate 

blade, inserted at the apex of the sheath, just below 

the ligule. Cf. LEAF; SHEATH. 

**LEMMA (Gr., a husk). The glume that subtends n flower. Lemmas that subtend normally constituted and developed flowers are conventionally 

called fertile lemmas, to distinguish them from 

sterile lemmas, whose subtended flowers are depauperate or obsolete. See TRANSITIONAL GLUMEs. 

**LEPTOMORPH (Gr,, Zeptos, slender, and 

morphe, form). Characterizes the rhizomes proper 

(of a bamboo plant) when they are long and slender, and have the following associated characteristics: a cylindrical or subcylindrical form, with a 

diameter typically less than that of the culms 

originating from it; internodes longer than broad, 

relatively uniform in length, symmetrical or nearly 

so, rarely solid, typically fistulose, the usually narrow central lumen interrupted at each node by a 

diaphragm; nodes in some genera usually somewhat 

GAN. 

**LEAF SHEATH. 
NUMBER 9 131 

elevated or inflated, in others not; lateral buds in 

the dormant state boat-shaped, with a distally 

oriented apex. Cf. PACHYMORPH. 

**LEVEL (L. libella, a water level or plumb level). 

Even, horizontal. Characterizes the pattern of insertion of the prima_ry components (first order 

axes) of a pleioclade branch complement, as exemplified by members of the genus Apoclada. 

LIGULATE. Having a ligule. 

LICULE (L. Zigula, a little tongue). A thin, apical extension of a sheath proper, adaxial to the 

locus of insertion of the sheath blade (or leaf 

petiole). The outer rim of the little cup revealed 

by the abscission of the petiole of a foliage leaf is 

sometimes referred to as the “outer ligule,” especially when it is conspicuously developed, as it is 

in some variants of Arundinaria gigantea ssp. tecta. 

The term “inner ligule” is then applied to the 

adaxial structure conventionally known simply as 

the ligule. The ligule is reduced to a mere line, or 

is even lacking entirely, in the culm sheaths of 

certain species of bamboos. 

Locus (L., place). The locale in which some 

event takes place, or where something is to be 

found. Locus is here given the connotation, point, 

or line, or area, principally in the phrase “locus o€ 

insertion.” See INSERTION. 

One of 

the small, usually thin, delicate and transparent 

structures (by some authors referred to as “scales”) 

inserted usually in a single whorl of 3, immediately 

below the stamens in the bamboo flower. The 

lodicules are relatively large, opaque, and parchment-like in the flowers of known species of 

Streptochaeta. Their number is variable in some 

bomboos (up to 12 or more in OchZandra stridula); 

in known species of Gigantochloa, typical lodicules 

are lacking entirely. 

LUMEN (L., opening). “The space which is 

bounded by the walls of an organ, as the central 

cavity of a cell” (Jackson, 1949); the central cavity 

of a hollow internode of any segmented axis of a 

bambco plant. 

**MAVERICK. Nonconformist (see Webster, 

1959). A term borrowed here (see page 15 et passim) for the purpose of characterizing ontogenetic 

and/or morphological features that appear to range 

alone, as if moving in independent trajectories. 

Maverick features appear at random as discrepancies (phenotypic diversities) among the other feaLODICULE (L. lodicula, a small coverlet). 

tures of one or more species of different genera 

that are otherwise more or less distantly related, 

or at least phenotypically disjunct. I was at first 

inclined to credit their presence in such discrepant 

settings as due to recent introgressions of unlinked 

genes. More recently I have considered the possibility that they come from a common ancestor by 

way of recessive genes. 

‘“MBG. McClure Bambo Garden, Bethesda, 

Maryland, U.S.A. See also P.I. 

MERISTEM (Gr. meristos, divisible). A body of 

tissue in which cell division and differentiation are 

active or potential. 

METAMORPH (Gr. meta, implying change; morpht?, 

form). Of intermediate form; a term proposed here 

to designate certain underground portions of segmented axes, whose transitional character cannot 

be clearly indicated by the use of existing conventional terms. See METAMORPH I; METAMORPH 11 

A transitional axis that occupies 

a position between the culm neck and the base of 

the culm proper, where no clearly defined rhizome 

intervenes; in some cases doubtfully distinct from 

an elongated culm neck. 

A transitional axis intermediate 

in form and position between the apex of a rhizome 

(either pachymorph or leptomorph) and the culm 

into which the rhizome is transformed apically. It 

appears where the transformation of the apex of 

a rhizome into a culm takes place gradually and not 

abruptly. 

METAMORPHOLOGICAL (Gr. meta, beyond, plus 

morphological). Pertaining to the techniques, the 

areas of investigation, or the data of disciplines 

other than morphology. 

MONADELPHOUS (Gr. monos, one; adeZphos, 

brother). United in a single brotherhood; an adjective cognate with Monadelphia, the name of 

“a Linnean class in which the anthers are united 

by their filaments into a single brotherhood” (Jackson, 1949), and conventionally used to describe the 

stamen complement of a flower when all of its 

members are united by their connate filaments. 

Cf. DIADELPHOUS; TRIADELPHOUS. 

METAMORPH I. 

METAMORPH 11. 

MONOCARPIC. See MONOPERIODIC. 

““MONOCLADE (Gr. monos, one; klados, branch). 

Characterizes those branch complements that 

contain only one primary axis. See BRANCH 

COMPLEMENT; MONOCLADE, RESTRICTED; MONOCLADE, 

UNRESTRICTED. Cf. PLEIOCLADE. 
132 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

* *MONOCLADE, RESTRICTED. Characterizes those 

monoclade branch complements whose primary 

axis is incapable of proliferating basally, for lack 

of buds at its proximal nodes. When buds are present at the proximal nodes of the solitary axis of a 

monoclade branch complement but remain dormant, such a branch complement is referred to 

herein as facultative restricted monoclade. 

““MONOCLADE, UNRESTRICTED. Characterizes those 

monoclade branch complements whose solitary primary axis is capable of proliferating basally from 

buds at its proximal nodes. 

MONOPERIODIC (Gr. monos, one; periodos, a completed course). Having but a single reproductive 

cycle within the lifetime of a plant; characterizes 

bamboos (such as Bambusa arundinacea) that 

flower but once, then perish. Monocyclic suggests 

itself ior use in the sense here given the newly 

coined term, monoperiodic, but it already bears 

the disqualifying connotation annual, with reference 

to the life span of a plant-besides indicating sets 

of parts (such as sepals and petals) that comprise 

but a single whorl or cycle. In the interest of 

clarity, the term monocarpic, commonly used to 

indicate the monoperiodic character of certain 

bambcos, should be reserved (with its orthographic 

variants; see Jackson, 1949) for reference to a pistil 

comprising but a single carpel. Cf. POLYPERIODIC. 

MONOPODIAL (Gr. monos, one; pous, podos, foot). 

Having the form of a monopodium, which is defined by Jackson (1949) as “a stem of a single, and 

continuous axis.” This term was used earlier (McClure, 1925) to designate the type of rhizome 

described as leptomorph (q.v.). Cf. SYMPODIAL. 

MORPHOGENESIS (Gr. morphd, form; genesis, beginning or origin). “The production of morphological characters” (Jackson, 1949). As defined 

here, morphogenesis connotes not only its external 

manifestations but also, and primarily, the nature 

and sequence of events in the program of genic 

monitoring of physiological states associated with 

the emergence of the structural features that determine the characteristic form of a plant. 

MUCRONATE (L. mucronatus, pointed). “Possessing a short, straight point, as some leaves” 

(Jackson, 1949). Cf. CUSPIDATE; MVTICOVS. 

“Blunt,* awnless’’ (Jackson, 1949); used primarily (when applicable) to describe the apex in small sheathing apMurIcous (L. muticus, docked). 

pendages on the various axes of the bamboo inHorescence. 

The constricted basal part, 

characteristic of all, or most, of the segmented 

vegetative axes of a bamboo plant. 

NECK SHEATH. One of the reduced, bladeless 

sheaths that clothe the constricted proximal part, 

the neck, of various vegetative axes in the bamboo 

plant. Neck sheaths are sometimes referred to as 

CATAPHYLLS. 

” *NOTHOMORPH (Gr. nothos, hybrid; morphos, 

form). “Applied to any hybrid form, whether F1, 

segregate or back-cross“ (International Code of 

Botanical Nomenclature, Article H.5). Cf. Figures 

17 and 18. 

OBSOLETE (L. obsoletus, worn out). “Wanting 

or rudimentary; used of an organ which is scarcely 

apparent or has vanished” (Jackson, 1949). 

OBTERETE. Circular in cross-section, tapering 

progressively from one end to the other, and 

smallest at the proximal end (for example, a 

rhizome neck). Cf. TERETE. 

“Having the solid figure or lengthwise 

outline of an egg; popularly elliptical or ellipsoidal. 

Bot. broadly elliptical” (Webster, 1959). “A body 

or figure in the shape of the long section of an egg, 

or popularly an ellipse” (Webster, 1959). 

“Shaped like an egg; oval” (Webster, 

1959). “With an outline like that of a hen’s egg cut 

in two lengthwise, the broader end downward” 

(Bailey, 1914; Jackson, 1949). 

**OVOID AND OVATE. The terms “ovoid” and 

“ovate” are used in diverse senses. The original 

botanical sense of ovate appears to be [that of] 

Bailey (1925) and Jackson (1949) but it has (under 

popular influence) drifted to mean elliptic or ellipsoid (Stearn, 1966; Webster, 1959). 

Abbreviation of “Plant Introduction,” prefixed to a permanent identifying number assigned 

by the U.S. Department of Agriculture to each lot 

of living plant material (seeds, plants, or cuttings) 

accessioned in its record of plant introductions. 

These numbers are published, along with pertinent 

names and documenting notes, in a continuing 

series of “Plant Inventories.” See also MBG. 

**PACHYMORPH (Gr. pachys, thick; morphd, 

form). Term proposed by McClure to describe 

the rhizome proper (of a bamboo plant) when it is 

short and thick, and has the following associated 

characteristics: a subfusiform (rarely subspherical), 

NECK (AS. hnecca). 

““OVAL. 

““OVATE. 

P.I. 
NUMBER 9 

usually more or less curved (rarely straight) shape, 

with a maximum thickness typically somewhat 

greater than that of the culm into which it is always 

transformed, apically; internodes broader than long, 

asymmetrical (longer on the side that bears a bud), 

solid (apparently never fistulose); nodes not elevated or inflated; lateral buds solitary, in the 

dormant state asymmetrically dome-shaped, with a 

subcircular margin and an intramarginal apex. 

Cf. LEPTOMORPH. 

PALEA (L., [a piece of] chaff). The prophyllum 

of the axis of a gramineous flower. Jackson’s (1949) 

definition: “the inner bract or glume in grasses, 

called ‘Palet’ by North American writers” is too 

vague to be useful. See PROPHYLLUM. 

PANICLE (L. pun.icuZu, a tuft). A determinate inflorescence with branches of more than one order. 

Jackson’s definition of panicle is not applicable 

here. 

* ‘PANICULATE (L. paniculutus, tufted). Inflorescences with branches of more than one order, the 

rachis excurrent or deliquescent. See PANICLE. 

**PARAMO (Sp., from L. purumus, teste Larousse, 

in AugP: and Gisbert, 1949:697). “Extensive treeless regions that dominate the summits of the Andean cordillera from the upper limit of the forestan elevation of 3800 m (locally 3220 m)-up to the 

level of permanent snow (4700 m);” teste Cuatrecasas (1959:249) where distinctive ecological, vegetational, and floristic features of the piramo are 

described at length-in Spanish. 

PARENCHYMA (Gr. parenchein, to pour in beside). 

Fundamental tissue; ground tissue; a tissue, such as 

pith, composed of thin-walled, undifferentiated, 

isodiametric cells (adapted, from Jackson, 1949). 

PEDICEL (L. pedicellus, dim. of pes, foot). The 

stalk of a spikelet, that is, the distal segment or 

internode of the axis immediately below the glumes 

that marks the base of a spikelet. The ultimate 

branches of a determinate inflorescence are all pedicels, since each terminates in a spikelet. In spicate 

racemes, and in pseudospikelets, the pedicels are 

usually very short. Cf. PEDUNCLE; RACHIS. 

PEDICELLATE. Having a pedicel. 

PEDUNCLE (L. pedunculus, dim. of pes, foot). 

The stalk of ah inflorescense, that is, the unbranched segment of the inflorescence axis that is 

immediately below the rachis. In determinate inflorescences, the first sheathbearing node below the 

first branch of the inflorescence is here taken arbitrarily as marking the base of the peduncle, and the 

locus of insertion of the first branch of the inflorescence as marking the apex. In those rare cases 

where a determinate inflorescence includes only 

one spikelet, the peduncle is not distinguishable 

from the pedicel. In indeterminately branching 

inflorescences, the peduncle is usually very short. 

An exception appears where the primary pseudospikelet is terminal (instead of lateral) to a flowering branch. Here the peduncle is more elongate. 

Cf. PEDICEL; RACHIS. 

PEDUNCULATE. Having a peduncle. 

* “PERCURRENT (L.percurrens, passing through). 

Used here in a sense identical with that given 

EXCURRENT. 

**PERFECT. Signifies the presence, within an 

anthecium, of a flower containing a normally developed complement of sexual elements, both 

staminate and pistillate. Same as hermaphrodite. 

PERCAMINEOUS (L. pergamenu, parchment). 

“Like parchment in texture” (Jackson, 1949). 

PERSISTENT (L. persistens, remaining in place). 

Not deciduous; applies to organs that remain in 

place after they have fulfilled their natural functions (for example, the culm sheaths of Arundinaria 

gigantea ssp. tecta). 

PINNATE (L. pinnutus, feathered). “Featherlike” 

(Webster, 1959). Having lateral appendages distributed in two continuous series inserted, respectively, on opposite sides of an axis and antrorsely 

oriented; a borrowed term used here to describe the 

pattern of insertion and orientation of the proliferations of the stigmatic surface in certain bamboos. 

Jackson (1949) and Webster (1959) both confine 

the botanical application of the term pinnate to 

the arrangement of the leaflets on the rachis of a 

compound leaf. 

PISTIL (L. pistillurn, a pestle). See GYNOECIUM. 

* “PLEIOCLADE (Gr. pleion, more; Itlados, branch). 

Characterizes those branch complements that contain more than one independent primary (firstorder) axis. See BRANCH COMPLEMENT; PLEIOCLADE, 

RESTRICTED; PLEIOCLADE, UNRESTRICTED. Cf. MONOCLADE. 

* *PLEIOCLADE, RESTRICTED. Characterizes those 

pleioclade branch complements whose component 

axes are incapable of proliferating basally for lack 

of buds at their proximal nodes. 

* “PLEIOCLADE, UNRESTRICTED. Characterizes those 

pleioclade branch complements whose component 
134 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

axes are capable of proliferating basally from buds 

at their proximal nodes. 

PLEIOGENY (Gr. pleion, more; genos, birth). “An 

increase from the parental unit, as by branching or 

interpolation of members” (Jackson, 1949); a general term for the particular examples described 

under PROLIFERATION and PLEIOCLADE, as pertaining 

to vegetative axes, and under INDETERMINATE, as 

pertaining to reproductive axes. 

PLICATE (L. plicutus, folded). “Folded into 

plaits, usually lengthwise” (Jackson, 1949); marked 

by longitudinal ridges suggesting a prior condition 

of being folded like a collapsible Chinese fan; an 

appearance shown by some bamboo leaf blades. 

* “PLURICESPITOSE (L. pluris, of more; caespitosus, 

from caespes, a tuft). Characterizes the clump 

habit of a bamboo plant that embraces more than 

one tuft of culms, all tufts belonging to an individual plant being connected by subterranean axes 

as in Arundinaria gigantea ssp. tecta. Cf. UNICESPITOSE. 

POLYCARPIC. See POLYPERIODIC. 

POLTPERIODIC (Gr. polys, many; periodos, a complete course). Having many reproductive periods, 

alternating with vegetative periods, within the lifetime of one plant; characterizes bamboos (such as 

plants of the known species of Phyllostachys) that 

flower repeatedly (and usually periodically) during 

an indefinite life span. Polycyclic suggests itself for 

use in the sense here given polyperiodic, but is 

rejected because of its long-established use with 

reference to the occurrence of an indefinite number 

of whorls of parts, such as sepals or petals in a 

flower. Pleiocyclic is already burdened with the 

connotation perennial, which is incompatible as a 

meaning alternate to that here given polyperiodic. 

Moreover, monocyclic, which would pair with the 

pleiocyclic, already means annual, in reference to 

the life span of a plant. In the interest of clarity, 

the term polycarpic, sometimes used to indicate the 

polyperiodic character of certain bamboos, should 

be reserved (with its orthographic variants) for 

reference to pistils with numerous carpels (see 

Jackson, 1949). Cf. MONOPERIODIC. 

**PRECOCIOUS (as applied to the proliferation of 

the branch primordium at midculm nodes). Elaborating secondary branch primordia before the 

rupture of the bud-scale (prophyllum) that accompanies the “germination” of the branch bud. 

““PRIMARY. The first (initial) element in a 

series of related axes or structures of a given category, as branch buds or branches. 

**PRIMITIVE. “Ancestral” in the sense of L. L. 

Forman (1964:390 footnote). 

PRIMORDIAL (L. primordialis). “First in order 

of appearance” (Jackson, 1949). See PRIMORDIUM. 

PRIMORDIUM (L., the beginning). An axis or an 

outgrowth of an axis in its earliest recognizable 

condition, or in an early dormant state. In the 

bamboos, a branch primordium borne on a segmented vegetative axis is always enclosed in a 

prophyllum; an adventitious root primordium is 

never so enclosed. See PROPHYLLUM. 

An initial primordium 

is a body of undifferentiated meristem which, when 

enclosed by a prophyllum and remaining dormant, 

forms part of a primary bud; it then constitutes an 

incipient axis of the first order. When inserted at 

a culm node, its first potential is to give rise to a 

primary element of a branch complement. See 

BUD, PRIMARY. 

PROLIFERATION (L. proles, offspring; fero, I bear). 

“Bearing progeny as offshoots” (Jackson, 1949). 

The term is brought into focus here with reference 

to the rapid multiplication of members of a branch 

complement by the prompt awakening of buds at 

the proximal nodes of the component members. 

The proliferation of the culm itself by the same 

process (without the intercalation of a rhizome) is 

called stooling or tillering. Proliferation should not 

be confused with prolification, which Jackson 

(1919) differentiates as “The production of terminal or lateral leaf buds in a flower.” An example 

of prolification is cited under DETERMINATE. See 

also PLEIOGENY. 

**PROMONTORY (a physiographic term described 

by Webster, 1959). I have borrowed this term to 

describe the tapered bulge that extends downward 

from the locus of initiation of the primordium of 

the primary bud at midculum nodes in all of the 

known species I here include in the genus Arthrostyfidium. See GREMIAL. 

PROPHYLLATE. Bearing a prophyllum (q.v.). 

PROPHYLLUM (L. prophyllum, first leaf). A sheathing organ, usually 2-keeled and inserted circumaxially at the first (proximal) node of a branch. In 

vegetative axes, and in inflorescences of indeterminate branching, the prophyllum at first surrounds 

the branch primordium to form a bud. Jackson’s 

* *PRIMORDIUM, INITIAL. 
NUMBER 9 135 

(1949) definition of prophyllum limits it to what 

is now generally called a palea. Prophylla are by 

some authors called bracts, or bracteoles. See 

PALEA; BUD. 

PROXIMAL (L. proximus, nearest). Basal; situated at or near the base of an axis or an organ; 

designates loci of insertion, or structures, so situated. Cf. DISTAL. 

PSEUDOSPIKELET (L. pseudo, false; by extension, 

superficially resembling; spicula, spikelet). A spikeletlike branch of an iterauctant (indeterminately 

branching) inflorescence. See INDETERMINATE. 

PULVINUS (L., cushion). A trBpical organ (L. 

trGpicus, pertaining to a turning) associated with 

movement of differential growth. Pulvini usually 

manifest themselves as dome-shaped eminences, 

commonly in pairs, strategically located in relation 

to the organ with whose orientation they are concerned-at or near the base of the leaf petiole, for 

example, or a branch of an inflorescence. Pulvini 

function through one or the other of two physiological mechanisms: changes of turgidity and differential growth. 

*#QUINATE (as applied to pleioclade midculm 

branch complements). Typically consisting of only 

five axes or branches. 

RACEME (L. racemus, a bunch of grapes). A 

semelauctant gramineous inflorescence with a single 

order of (usually solitary) branches. Jackson’s 

(1949) definition is not applicable here. A raceme 

in which some of the branches emerge in fascicles 

of tw:, or three may be characterized (interpreted) 

as a paniculate raceme, if the extra branches in 

each fascicle are taken to be secondary ones arising 

subcutaneously from the base of a primary one. 

Athroostachys capitata (Figure 19c) bears paniculate racemes in which all branches are relatively 

short. This gives the inflorescence a capitate superficial appearance. A spicate raceme is one of which 

the pedicels are so short that the inflorescence resembles a spike. 

RACHILLA, rhachilla (Gr. dim. of rachis, q.v.). 

The axis of a spikelet in any gramineous plant. 

Jackson (1949) defines rhachilla vaguely as “a secondary axis in the inflorescence of grasses.” 

RACHIS; rhachis (Gr. rhachis, backbone). The 

primary axis of an inflorescence; its position is 

terminal to the peduncle. 

REDUCED. Subnormal in size; connotes also (in 

some applications) either a failure to fulfill a normal function, or a diminution in the expected 

number of parts in a set (of stamens, for example). 

Cf. DEPAUPERATE. 

*#RESTRICTED (L. restrictus). Characterizes (1) 

a branch complement-either monoclade or pleioclade-as unable to proliferate from buds at proximal nodes of its first-order component member (s); 

and (2) the locus and the mode of insertion of a 

monoclade branch complement, regardless of 

whether the primary component (first-order) axis 

is dominant or not. 

RHIZOME (Gr. rhizbma, a mass of roots). An individual component branch of the subterranean system of segmented axes that constitute the “chassis” 

(popularly referred to as the “rootstock”) of a bamboo plant. A rhizome consists of two parts: the rhizome proper and the rhizome neck. Two distinct 

types of rhizome are differentiated: leptomorph and 

pachymorph (q.v.). 

RHIZOME SHEATH. The husklike sheathing organ 

inserted at each node of a rhizome proper (as distinct from the rhizome neck). 

Of higher order in a given system of related axes. Example: the branches that 

arise by proliferation from buds basal to the primary branch at midculm nodes in Arthrostylidium. 

Cf. SUBSIDIARY. 

* #SECUND (L. secundus, following). “Arranged 

on one side only; unilateral, as flowers in some 

racemes, spikes, etc.” (Webster, 1959). Applied to 

“parts or organs directed to one side only, usually 

by torsion” (Jackson, 1949). The term comes into 

use in the description of bamboo inflorescences 

with an excurrent rachis, where the modification of 

the rachis, either by torsion or by asymmetrical 

development, causes all of the spikelets, or branches, 

to be oriented more or less strongly toward each 

other on one side of the rachis, and away from 

each other on the other side of it. In actual practice, the term may be applied either to the inflorescence as a whole, or to the parts (spikelets or 

branches) whose orientation is the focus of attention. 

SEMELAUCTANT (L.semel, once; auctans, increasing, growing). Embracing but a single grand 

period of growth. A determinate bamboo inflorescence may be said to be semelauctant. See DETERMINATE; ITERAUCTANT. 

SENSU LATO; SENSU STRICTO (L.). In a broad 

sense; in a restricted sense; expressions used to in- 

**SECONDARY. 
136 SMITHSONIAN CONTRIBUTIONS TO BOTAKY 

dicate the intended scope of a given application of 

a scientific name or a scientific term. 

SHEATH (ME. shethe). A sheathing organ, the 

basal part of which, the sheath proper, completely 

surrounds the vegetative axis on which it is borne, 

its locus of insertion being circumaxial. In its 

simplest form (as in typical neck sheaths and 

rhizome sheaths) the sheath proper terminates 

apically in a short, hard point. In its fully elaborrated form-that characteristic of examples in the 

middle of any series-the sheath proper usually 

terminates above in a ligule, at the base of which 

is inserted a more or less expanded laminar appendage referred to in a comprehensive sense as the 

sheath blade. In addition, the sheath proper commonly bears, at or near each extremity of the locus 

of insertion of the blade, a tuft of bristles (oral 

setae). These may be borne either directly on the 

sheath proper or on the margin of an auricle. When 

there is a pair of auricles (one on each side) the 

two may be similar and subequal, or they may be 

unequal in size and dissimilar in shape. To avoid 

the ambiguity occasioned by a commonly encountered indiscriminate usage of the term leaf (q.v.) 

to refer to any one of the diverse forms of sheaths 

borne on the vegetative axes of the bamboo plant, 

two expedients are followed: (1) the adoption of 

the differential terms rhizome sheath, neck sheath, 

culm sheath, branch sheath, leaf sheath, and prophyllum (q.v.); and (2) the differentiation of the 

sheath proper from its appendages. A clear precedent for this effort to avoid ambiguity is found 

in the now fairly common adoption of a precise 

terminology to differentiate the several types of 

sheathing structures borne on the reproductive axes 

of the bamboo plants, as bracts, prophylla, glumes, 

lemmas, and paleas. See SHEATHING ORGAN. 

SHEATH BLADE. A distinct foliar part, the 

lamina, that is appended apically on the laminiferous culm sheaths proper and branch sheaths 

proper in any series. A sheath blade is distinguishable from a leaf, first of all by the relatively proximal position of the sheath proper of the former on 

any aerial vegetative axis, while leaf sheaths are 

always inserted at the distal nodes of any culm or 

branch. Sheath blades are always sessile, while the 

leaf blades are petiolate (in all known bamboos 

except some species of Neurolepis). Moreover, characteristic differences in shape are usually quite 

marked. The strong divergence in form, as between 

leaf blades and sheath blades in the bamboos, is 

one of the commonest and most useful of the gross 

morphological distinctions between members of the 

Bambusoideae and those of other grasses. See LEAF. 

SHEATH CALLUS (L. callus, hard skin). A somewhat prominent ring of parenchymatous (not hard) 

tissue which remains at a sheath node after the 

abscission of a sheath (of a culm, especially) in 

some bamboos (Phyllostachys nidulnria, for example). Cf. GIRDLE. 

SHEATH NODE. The circumaxial locus of insertion of a sheath on any vegetative axis of a gramineous plant (elaborated after Hackel, in LamsonScribner and Southworth, 1890: fig. 1). The sheath 

node is marked externally by a more or less prominent offset in the surface of its axis. At the level of 

each sheath node the crossing over and anastomosis 

of fibrovascular bundles take place through a diaphra,m which marks the internal boundary between adjacent internodes. Cf. CULM NODE; see DIASHEATH SCAR (Gr. escharu, mark). Jackson 

(1949) defines Scar as “a mark left on a stem by a 

separation of a leaf, or a seed by its detachment; a 

cicatrix.” In the bamboos, a sheath scar is a narrow, 

transverse, circumaxial trace, the locus of abscission 

of a sheath proper. A sheath scar marks the pmition of a sheath node (q.v.). 

Any sheathing structure inserted at a node of any vegetative or reproductive 

axis in a gramineous plant. Among the bamboos, 

distinguishable types of sheathing organs are rhizome sheaths, neck sheaths, culm sheaths, branch 

sheaths, leaf sheaths, prophylla, bracts, empty 

glumes, lemmas, and paleas. Unless used, and interpreted, with discretion, the often-encountered 

term foliar organ is apt to be ambiguous, since it 

may be construed as referring either to a sheath 

proper alone, to a sheath proper with all of its 

appendages, to the blade only, or to a bladeless 

sheathing structure, such as a prophyllum. See 

LEAF. 

SHOOT (noun, from AS. sczotan, to move rapidly). 

“ (1) A young growing branch or twig; (2) the ascending axis; when segmented into dissimilar members it becomes a.stem” (Jackson, 1949). See CULM 

SILICA BODIES; also silica corpuscles (G. Kieselkdrpe,). Bodies of silica that are secreted and persist within the cells of various tissues of the plant, 

PHRAGM. 

SHEATHIKG ORGAN. 

SHOOT. 
NUMBER 9 

particularly any epidermal layer. In both size and 

shape, they range widely. At one extreme, they are 

small, lack characteristic shape, and occur solitarily 

or in small numbers within a given cell. At the 

other extreme, they may be large enough individually to fill a cell more or less completely (see 

SILICA CELLS) and have a characteristic shape. 

Metcalfe (196O:xlii, fig. 1) lists 20 “types” (forms) 

of silica bodies, some of which are illustrated. He 

states (p. xx) that “the silica-bodies in silica-cells 

assume very characteristic forms when the grass leaf 

is mature, and are of considerable value for diagnostic and taxonomic purposes.” Ohki’s (1932: 73f) 

key to Japanese genera of bamboos refers to the 

occurrence of “silica corpuscles” in the long epidermal cells of the leaf in Dendrocalamus [D. Zatiflorus] 

and in the articulation (bulliform) cells of the leaf 

in Sinobambusa [S. tootsik] and Chimonobambusa 

[ Ch . q uadrangu laris]. 

SILICA CELLS. Epidermal short cells each of 

which is more or less completely filled by a single 

silica body (Metcalfe, 1960:xx). 

SPICATE (L. spicatus, bearing spikes or ears). 

Having some or all of the characters of a spike 

(q.v.). A spicate raceme is a raceme in which the 

pedicels are so short that the inflorescence resembles 

a spike. 

SPIKE (L. spica, an ear of grain). In the bamboos, a spike is a determinate inflorescence in which 

the sessile or subsessile spikelets are inserted on a 

solitary rachis. Since the distinction between a spike 

and a spicate raceme is, in the terms of their definitions, only a matter of the relative length of “subobsolete” pedicels, it is sometimes difficult to choose 

between these terms for the description of a given 

inflorescence (as in some species of the genus 

Merostachys, for example). According to Jackson’s 

(1949) definition (not applicable here) a spike is 

an “indeterminate inflorescence with flowers sessile 

on a common elongated axis.” A loose interpretation of Jackson’s definition of a spike has resulted 

in the occasional misapplication, in the literature, 

of the term spike to a spikelet (q.v.) of bamboo. 

A basic structural component of every normal gramineous inflorescence, comprising a segmented axis (the 

rachilla) and its appendages. The appendages (beginning with the lowermost) are: empty glumes 

(usually two, rarely more, sometimes only one, 

rarely lacking entirely), lemmas (either variable or 

SPIKELET (L. spicula, little spike). 

invariable in number, according to the taxon involved), and branches of the rachilla (one subtended 

by each lemma), each bearing a palea and the parts 

of a flower. In some taxa, one or more of either 

the distal or the proximal lemmas, or both, may be 

sterile by virtue of either being empty or subtending 

an incompletely developed flower. See FLORET; 

FLOWER. [Empty glumes= transitional g1umes.-ed.] 

* “SPODOGRAM (Gr. spodos, ashes; gramma, something drawn). The “ash picture” of a plant part, 

such as leaf epidermis, which is revealed on a slide 

prepared by the following method: the plant part 

is burned to the white-ash stage and the product 

transferred intact to a glass slide and mounted in 

an agent such as aniline or Canada balsam. Spodograms were used for purposes of plant classification 

by Molish (1920) and for identification of bamboos 

by Ohki (1932). 

SPORADIC (Gr. sporadikos, dispersed). “Widely 

dispersed or scattered” (Jackson, 1949); dispersed, 

or irregular, in time, as when the individual plants 

of a given generation of bamboos (from seeds of 

a common origin) enter the reproductive phase at 

different times, or at irregular intervals. In either 

case, the flowering is said to be sporadic (Gamble, 

1896:viii). Cf. GREGARIOUS. 

STOMA (Gr. mouth, opening; pl., stomata). A 

functional organ found commonly, but not everywhere, in the epidermis that covers photosynthetic 

tissues. A stoma consists of two guard cells and (in 

the bamboos, as far as known) two subsidiary cells 

(Porterfield, 1937). Changes in the turgor of the 

guard cells result in the opening and closing 06 a 

slitlike aperture (the stoma of Jackson, 1949) between them. 

**STYLE. The description of the style as a separate or distinct structure encounters difficulties 

which grow out of peculiarities in the ontogeny of 

the gynoecium. In many bamboos, the stigmas are 

at first sessile at the apex of the ovary and there 

is no discernible style. Usually, a style develops 

later, but rarely (as in Bambusa multiplex), the 

stigmas remain sessile. It does not apeear useful 

or logical to call these separate styles. The style 

when it appears seems to be an outgrowth of the 

pericarp; but only by means of anatomical studies 

of the pistil (gynoecium) at different successive 

stages in its development can the details of the 

origin, structure, and relationships of the compo- 
138 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

nent structures traditionally given independent 

names and status be clarified. 

A prefix indicating either an approximation or some reservation or limitation in the use 

of the term with which it is combined, as in subdistal, suberect, subfamily, subfusiform. 

#*SUBEQUAL (as applied to the transitional 

glumes of a spikelet, or the component axes of a 

midculm branch complement). Nearly, but not 

quite of equal size. 

Of a shape suggesting the concept 

fusiform, but not corresponding to it precisely; the 

approximate general shape of a pachymorph rhizome. See FUSIFORM. 

Of the same order but of smaller 

size; example: smaller members of the complement 

of initial (primary) buds or branches at midculm 

nodes in Chusquea and Swallenochloa. Cf. SECONBranches of higher order 

arising from buds at the base of a dominant primary 

branch. 

SUBTEND (L. subtendo, I stretch underneath). 

To precede on a common axis, as a foliar organ 

precedes (subtends) a bud or branch inserted immediately above it. 

“Grooved or 

furrowed” (Jackson, 1949); as where the otherwise 

approximately cylindrical shape of the surface of 

the internode of a segmented axis is modified by one 

or more longitudinal depressions. In most bamboo 

genera the palea in functional florets is sulcate with 

a single longitudinal dorsal depression. The furrow 

or groove that marks, on the exterior of the caryopsis in many bamboos, the interior locus of the 

attachment of the seed to the pericarp. Cf. HILUM. 

SUTURE (Fr., from L. sutura, from suere, to sew). 

“A junction or seam of union; a line of opening or 

dehisccnce” (Jackson, 1949). 

““SYMPODIAL (Gr. sym or syn, together; pous, 

podos, foot). Having the form of a sympodium; for 

example, a system of related axes wherein successive 

branches assume the role or position of effective 

leadership or dominance so that the axis of a 

sympodium is composite; a term used earlier (McClure, 1925) to designate the branching habit of 

the type of rhizome described herein as pachymorph 

(q.v.). Cf. MONOPODIAL. 

**TARDY (as applied to the proliferation of the 

branch primordium at midculm nodes). ElaboratSUB- (L.). 

SUBFUSIFORM. 

*“SUBSIDIARY. 

DARY. 

SUBSIDIARY BRANCHES. 

*+SULCATE (L. sulcatus, furrowed). 

ing secondary branch primordia after the rupture 

of the bud scale (prophyllum) that accompanies the 

germination of the branch bud. 

TAXON (neo-Gr., from taxis, arrangement). “A 

taxoncmic group or assemblage of plants or animals having certain characteristics in common, 

which we take as evidence of genetic relationship, 

and possessed of some degree of objective reality” 

(Rickett, 1958). Rickett adds, “We can use the 

word wherever we can use ‘Taxonomic group’ in 

referring to the characteristics, dynamics, distribution, or uses of such an assemblage.” Morton (1957) 

cautions that “where the words ‘taxonomic group’ 

cannot be appropriately substituted, the word taxon 

is misused.” 

TERATIC (Gr. teras, teratos, monster, wonder). 

Abnormal; teratological, in the sense of conventional usage, with particular reference to marked 

deviations from the normal, or expected, morphologica! expression. See teratic example of prolification under DETERMINATE. 

TERETE (L., teres, well-turned). Circular in 

cross-section, tapered progressively from one end to 

the other, and smallest at the distal end or tip (for 

example, a small bamboo culm). Cf. OBTERETE. 

TERNATE (as applied to pleioclade midculm 

branch complements). Typically consisting of only 

three axes or branches. 

TESTA (L., covering, in a poetical sense of the 

substantive). The outer coat of a seed. 

TILLERING. “Throwing out stems from the base 

of a stem” (Jackson, 1949); proliferation of a culm 

from its basal (subterranean) buds, without the intercalation of a rhizome proper. 

TRAFANT (Fr., running, creeping). Ranging 

widely and freely; a term used by the Rivieres 

(1879:321 et passim) to characterize (1) the slender, 

elongate type of rhizome herein called leptomorph 

(q.v.), and (2) the bamboos that spread by this 

means. 

A term newly coined 

to designate those sheathing structures that occupy 

the physiologically transitional zone located immediately below the first fertile kmma of a bamboo 

spikelet. In many of the tribes of conventional 

grasses, and in some bamboos, it is possible to 

focus on one or more features that reveal a clearly 

perceptible distinction between two categories of 

structures in this zone, namely, empty glumes and 

sterile lemmas (q.v.). Agrostologists generally have 

**TRANSITIONAL GLUMES. 
NUMBER 9 139 

made the most (in some cases too much) of these 

structures as sources of taxonomic characters. In 

many bamboos, either the total number or the 

number of members of each category (or both of 

these features) found in a given spikelet may vary 

as between spikelets in a single specimen. Again, in 

some species, one may look in vain for a clear 

distinction (disjunction) between “empty glumes” 

and “s:erile lemmas.” In such cases, their taxonomic 

value may be greatly reduced. For this reason, in 

describing bamboo genera, I have adopted the 

expedient of treating both categories as members 

of a single morphological category, namely, transitional glumes. Besides avoiding a certain psychological difficulty that arises in some cases this 

emphasizes the existence of physiological gradients, 

and places in proper perspective the frequent appearance of morphological gradients, in place of 

the sharp (abrupt) morphological transitions sought 

by the taxonomist. 

TRIADELPHOUS (Gr. treis, three; adelphos, brother). Having “filaments in three brotherhoods”; an 

adjective cognate with Triadelphia; the name of “a 

Linnean order of plants with their stamens in three 

sets” (Jackson, 1949), and conventionally used to 

describe the stamen complement of a flower when 

its members have their filaments connate to form 

three distinct groups. In bamboos with triadelphous 

androecia, the three “brotherhoods” commonly 

comprise one, two, and three stamens, respectively. 

Cf. ~IONADELPHOUS; DIADELPHOUS. 

““TCTFTED (as applied to the component pleioclade branch complements at midculm nodes of a 

bamboo). Indicates more or less profuse branching 

from the basal nodes of the primordial branch. The 

component axes of tufted branch complements may 

be subequal, or unequal (with the primordial axis 

[branch] more or less strongly dominant). 

TURGIDITY (L. turgidus, inflated). Turgor, turgescence; the firmness imparted by “the distention 

of a cell or cellular tissue by water or other liquid” 

(Jackson, 1949). 

““UNEQUAL (as applied to the transitional 

glumes of a spikelet, or the component axes of a 

midculm branch complement). Of more or less 

strongly disparate size. 

* ‘UNICESPITOSE (L. unus, one; caespitosus, from 

cmspes, a tuft). Characterizes the clump habit of a 

bamboo plant that embraces but a single, moresr 

less compact, tuft of culms-as in Bambusa (Guadua) amplexifolia. Cf. PLURICESPITOSE. 

UNILATERAL (L. unilateralis, from unus, one; 

latzis, a side). One-sided. See SECUND. 

“‘UNISULCATE. Provided with, or marked by, a 

single horizontal groove. See SULCATE; HILUM. 

**UNRESTRICTED. Characterizes a branch complement (monoclade or pleioclade) as possessing the 

potential for proliferating by way or buds at the 

proximal nodes of its component first-order member (s). 

“The disposition or method of arrangement of foliage leaves 

within the bud” (Webster, 1959). Jackson’s (1949) 

definition, “the order of unfolding of leaf buds,” 

is not applicable here. In most known bamboos, the 

developing leaf blades are individually rolled up 

tightly along their long axis, with one edge at the 

center of the roll. 

**VERSATILE (L. versatilis, adaptable, turning 

with ease from one thing to another). Applied 

herein to a rhizome system capable of producing 

both leptomorph and pachymorph axes, as in 

some species of Chusquea. Versatile is elsewhere 

(Jackson, 1949) “turning freely on its support, as 

many anthers on their filaments.” 

ZYGOMORPHIC (Gr. zygos, yoke; morphb, form). 

Symmetrically divisible by a single plane; “used of 

flowers which are divisible into equal halves in 

one p!ane only, usually the anteroposterior” (Jackson, 1949). Normal bamboo flowers conform to this 

restricted criterion for zygomorphy (see McClure, 

19GGb: 114). According to Jackson (1949), “Sachs 

extends the meaning to such flowers as may be 

bisected in any one plane.” 

VERNATION (L. vernatio, renewal). 
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