SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 44 

The Genera of Bambusoideae (Poaceae) 

of the American Continent: 

Keys and Comments 

Cleofe E. Calderon

and Thomas R. Soderstrorn 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1980 
ABSTRACT 

Calderon, Cleofe E., and Thomas R. Soderstrom. The Genera of Bambusoideae 

(Poaceae) of the American Continent: Keys and Comments. Smithsonian Contributions to Botany, number 44, 27 pages, 1980.-The history of the grass subfamily Bambusoideae is reviewed and nomenclatural problems of subfamilial and 

tribal level are explored. Characters are presented to distinguish the subfamily 

from all other grasses and differentiating features of the two major groups of 

bamboos-herbaceous and woody-are included. Keys are given for the tribes 

and genera of herbaceous American bamboos and genera of woody American 

bamboos, the latter based principally on vegetative characters. A conspectus of 

the subfamily also appears, with a list of all 37 genera recognized in the American continent, each with nomenclatural and taxonomic notes. Comments on the 

morphology of the bamboo plant and the systematic value of some characters, 

especially vegetative, are given in the introduction. 

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

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

the Katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Calderon, Cleofe E.E. 

The genera of Bambusoideae (Poaceae) of the American Continent. 

(Smithsonian contributions to botany : ‘no. 44) 

Bibliography: p. 

1. Bamboo-America-Classification. 2. Grasses-America-Classification. 3. Botany-America 

-Classification. I. Soderstrom, Thomas R., joint author. 11. Title. 111. Series: Smithsonian 

Institution. 

QKLS2747 no. 44 [QK495.G74] 581’.08s [584’.93] 79-14750 

Srnithsonian contributions to botany : no. 44. 
Contents 

Page 

Introduction 1 

History of the Bambusoideae ........................................ 4 

Nomenclature 9 

Characterization of the Bambusoideae versus Other Grasses ............. 11 

Characterization of Woody versus Herbaceous Bamboos ................ 12 

Key to the Tribes of American Herbaceous Bamboos .................. 13 

Key to the Genera of American Herbaceous Bamboos ................. 13 

Key to the Genera of American Bamboos ............................ 15 

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

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

Conspectus of the Grass Subfamily Bambusoideae in the Western 

Tribes of American Herbaceous Bamboos ............................ 16 

Tribes of American Bamboos ....................................... 18 

Genera of American Bambusoideae ................................... 18 

Literature Cited ................................................... 23 

Hemisphere ..................................................... 16 

iii 
Dedicated to Professor C.G.D. Nees von Esenbeck 

(1776-1858) , father of the Bambusoideae 

Bambuseae, quidquid dicant alii, propriam 

subfamiliam s. ordinem graminunt eflormant, 

cum vicinis omnino connexam. 

Ruprecht, 1839 
The Genera of Bambusoideae (Poaceae) 

of the American Continent: 

Keys and Comments 

Cleofe E. Calderon

and Thomas R. Soderstrom 

Introduction 

Bamboos are a unique group of arborescent 

grasses that occur throughout the world, although 

they are generally associated with Eastern cultures, 

where they play such a significant role in the everyday life of the people. The New World shares 

equally in the bounty of bamboos, but they are less 

known. The rich forests of the New World provided 

man with all his needs and bamboo was only one 

of many useful plants at his disposal. While bamboo 

is used by man wherever he comes into contact 

with it, only in a few places of the New World has 

he used it to any great extent, such as the lowlands 

of Colombia and Ecuador, where the native Guaduas are used in construction. 

Many bamboos, though, are herbaceous in nature 

and, looking like palm seedlings or ferns, inhabit 

the dark rain forests. Many of these are diploids, 

unlike the larger tetraploid and hexaploid bamboos. 

The arborescent and the herbaceous genera, however, are similar in basic morphological and anatomical features and their alliance has been demonstrated by many corroborative studies. M’e 

consider all of the genera to comprise a single subfamily, Bambusoideae. 

Cleofe E. Calderon and Thomas R. Soderstrom, Department 

of Botany, National Museum of Natural History, Smithsonian 

Institution, Washington, D.C. 20560. 

1 

The concept of the subfamily Bambusoideae, with 

woody and herbaceous members, has evolved over a 

long period of time as part of the developing 

classification of the grass family. While the concept 

of the subfamily, as we espouse it today, was first 

presented by Ne=s von Esenbeck in the early part 

of the last century, few followed his ideas and only 

now, with additional data to support it, do most 

agrostologists accept the concept. 

Unlike most grasses, the woody bambusoid species 

(bamboos) and herbaceous barnbusoid (which we 

call herbaceous bamboos) have biological peculiarities that have hampered their study. Most bamboos, 

for example, are large and complex plants that remain in a vegetative state €or many years, flowering 

only occasionally. Collectors generally shy away 

from bamboos because of their size and complexity 

and usually because they are rarely in flower. Herbaceous bamboos, on the other hand, flower continuously or at least for several months during the 

year, but their flowers are generally hidden under 

leaves or are inconspicuous, causing the plants to 

look sterile. 

Nevertheless, there are many more collections of 

herbaceous bamboos than of bamboos themselves, 

and most collections of bamboos are incomplete. 

Our earlier field experience was directed especially 

toward the herbaceous bamboos, on which a greater 

number of studies had already been made than on 
2 

the bamboos. As a result we can now assign all the 

genera of herbaceous to tribes, while the state of 

knowledge does not allow this for all the bamboos. 

It was McClure’s opinion also, as evidenced in his 

final paper, published posthumously (1973), that 

the American genera of bamboos were still too imperfectly known to be placed in tribes. 

Arborescent forms occur in a variety of unrelated monocotyledonous families, for example the 

Palmae, Pandanaceae, Strelitziaceae, and Agavaceae. 

This growth form is also known in some members 

of otherwise herbaceous families: Aloe of Liliaceae, 

Puya of Bromeliaceae, and some genera of Araceae. 

In the same category the bamboos are one of the 

remarkable examples of arborescent forms in a predominantly herbaceous family, the Poaceae. All of 

these arborescent monocotyledons exhibit fundamental differences in construction, both from 

dicotyledonous trees and among themselves. Bamboos represent one of the few groups of arborescent 

monocotyledons that compete with dicotyledonous 

trees in size and branching habit. Some species of 

Bambusa in South America may reach over 30 m in 

height and 20 cm in diameter. Examples of this and 

even greater size are known in Asiatic members, 

such as Dendrocalamus giganteus, the largest of all. 

Bamboos, like the majority of arborescent monocotyledons, lack vascular cambia. Some essential 

features of the growth-form and growth-limiting 

characteristics in some arborescent monocotyledons 

have been described by Tomlinson (1964). In the 

example that he gave, branching is in general considered limited, in some cases absent, or more commonly the production of lateral branches occurs 

only in the inflorescence. It has been said that the 

absence of a vascular cambium from the aerial axis 

restricts branching in monocotyledons, while dicotyledonous trees with vascular cambia are typically 

much branched. The bamboos are probably the exception to this rule, and they present us with many 

questions still unsolved. 

Very little is known about branching in bamboos 

from a morphological and physiological point of 

view. The production of lateral vegetative branches 

does not correspond exactly with any of the cases 

cited by Tomlinson. In the aerial axis of a bamboo, 

lateral branches are produced from axillary buds, 

usually one at each node, although multiple buds 

are produced at the node in Chusquea. 

A bamboo culm does not usually produce 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

branches at the lower nodes but only at some distance above the ground, in the manner of a dicotyledonous tree. The number of branches at a node 

in the middle of the plant (“midculm node”) may 

be one or (some species of Merostachys) as many as 

150 or more, In many cases the original number of 

branches is augmented in successive years by rebranching from buds at the base of the branches. 

The adult pattern of this set of branches presents 

very useful diagnostic characters of generic validity. 

For descriptive purposes we refer to this aggregation 

of branches as a “branch complement.” 

The underground axis of a bamboo also branches 

and produces a strong rhizome system. There are 

basically two types, sympodial and monopodial. 

McClure (1966) refers to these as pachymorph and 

leptomorph, respectively. 

For practical descriptive purposes we use the term 

“bud complement” for the set of buds at the midculm node, which represents an early stage of the 

bud development and is typical for the genus. We 

avoid reference, however, to the number of initial 

buds since it can be known only from develop 

mental studies, which are still lacking. 

Bamboos possess a basic pattern of construction 

that consists of a strongly segmented axis with solid 

or hollow internodes and with a particular system 

of ramification. The axis passes through different 

stages during its development. There are basically 

two phases, each recognized by a combination of 

morphological features. A detailed analysis of the 

growth habit of bamboos is outside the scope of this 

paper, mainly because complete developmental 

studies are largely lacking. For this reason we are 

not going to describe the morphological and physiological changes of a bamboo seedling, the very early 

stages of the juvenile phase, or the many intricate 

phenomena that occur with the onset of flowering. 

We will refer very briefly to some features of the 

two major phases of growth form. These are different stages of the vegetative state of a bamboo plant 

and occur in the majority of known genera. As 

Tomlinson (1964) pointed out, bamboos constitute 

a peculiar and distinctive group with regard to their 

growth form and do not conform to any of the 

arborescent types that he defined. 

Stem elongation plays a significant role in the 

development of the plant form and consequently 

in the dimorphism of leaves, During the first phase 

they produce from the rhizome long shoots with 
NUMBER 44 3 

a strong apical dominance, fast internode elongation, and almost full suppression of lateral appendages, The new stem produces only one leaf at each 

node, in the form of a sheathing organ that envelops it completely. This new stem, which is at 

full diameter as it emerges from the soil, grows 

quickly and in many species reaches full height 

within a relatively short time. When the new shoot 

has completed most of its growth, the second phase 

of the vegetative cycle begins with the development 

of the lateral members, branches and foliage, at the 

same time that the soft tissues of the stem harden. 

This phase may extend over a period of several 

years during which the branching and rebranching 

occur along with a moderate increase in height. 

In the majority of grasses, including herbaceous 

bamboos, the development of the plant does not 

follow this pattern, which is associated with the 

arborescent habit. In general, grasses produce short 

shoots with a balance between apical dominance, 

moderate stem elongation, and a simultaneous production of lateral appendages, one leaf at a node in 

alternate fashion but very few or no lateral 

branches. 

Culm leaves and bud complements are the outstanding morphological structures of the first phase 

of the bamboo growth form, while branches and 

foliage leaves are characteristic of the second. These 

structures offer numerous characters of great systematic importance, which merit special comment. 

The extreme dimorphism in bamboo leaves has 

given rise to much confusion in leaf terminology. 

There are two basic types: “culm leaves” and 

“foliage leaves.” Culm leaves cover the new growing 

culm and since their function is to protect the 

growing shoot, the greater part of the leaf is sheath; 

often, in fact, the blade is hardly developed. Because the blade is the less developed part of the 

culm leaf, it has generally been disregarded and the 

whole structure called merely “sheath” in the literature. Kurz (1876), in his perceptive account of the 

growth form of bamboos, drew particular attention 

to the useful characters in this structure to distinguish species. 

Types of culm leaves are characteristic of genera, 

while differences in foliage leaves or in the culm 

leaf itself serve to distinguish species. This structure, which is of paramount importance in identifying bamboos, is often overlooked by collectors. 

As opposed to culm leaves, foliage leaves exhibit 

a greater development in the blade or lamina, the 

sheath becoming more or less proportionate to the 

size of the blade or to the number of leaves in a 

branch. The blade in most Bambusoideae is broad, 

more or less lanceolate, with the base constricted 

and jointed to the sheath by a little stalk, which 

often bears a pulvinus. This stalk is like the petiole 

of dicotyledonous leaves and is often similar in ap 

pearance; for descriptive purposes we use the term 

“petiole” without any interpretive connotation. 

Foliage leaves are produced laterally on the 

branches in most cases and are replaced periodically. The number of leaves on a branch varies 

within genera and species. Sometimes the leaves 

overlap closely, forming a distinct and recognizable 

set of leaves in the distal part of a branch. In our 

descriptions we refer to this group of leaves as a 

“leaf complement.” 

At this point we wish to stress the significance of 

the occurrence of two growth forms in bamboos, a 

phenomenon that does not occur elsewhere in the 

grasses. Moreover, the specialized leaves that cover 

the new shoot and later the numerous vegetative 

branches produced from almost every node are features unique among the grasses and rare in monocotyledons. Characters from these structures have 

proved to be of great systematic value at the generic 

and specific levels. Unfortunately these are the 

very structures that are usually not collected. 

Flowers are of equal systematic importance in 

bamboos as they are in other grasses and flowering 

plants, but because they are so seldom available we 

must generally resort to distinguishing bamboos by 

vegetative characters, of which, fortunately, there 

are many. 

After a limited period of vegetative growth the 

reproductive phase starts with the production of inflorescences. In most bamboos this takes place at intervals from few to many years, at which times it is 

generally gregarious. In some others the frequency 

of flowering in a population is almost yearly, as in 

species of Swallenochloa and Myriocladus, both 

genera that inhabit high mountains. It is interesting 

to note that many bamboos cease vegetative growth 

for a long period prior to the flowering and often 

lose all foliage at the peak of this process, which 

may last for a few years or more. In many cases 

the development of inflorescences is not limited to 

the distal parts of the branches or culm, but to 

every part of the plant where meristematic tissue 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

allows it, from the base of the plant up. Thus, the 

complexity of bamboo inflorescences increases, making the traditional terms used in descriptions of 

grasses, such as spike, raceme, and panicle, inappropriate. 

Generally in bamboos and, to a lesser extent, in 

herbaceous bamboos, inflorescences are complex 

systems of ramification throughout which bracts and 

prophylla are well developed. The basic units of 

this system are partial inflorescences, in themselves 

clusters of spikelets. In some bamboos the production of partial inflorescences starts at the base of the 

spikelet by the activity of buds in the axils of the 

glumes, as in the “pseudospikelets” of Bambusa 

(McClure, 1966, 1973). Along very general lines the 

inflorescences of bamboos can be divided into two 

major groups, In one the production of inflorescences is restricted, limited to a single group of 

spikelets. In the other case it is unrestricted, with 

unlimited production of inflorescences brought 

about by rebranching from buds at the base of any 

one of the partial inflorescences. The complexity of 

an inflorescence is increased by the different basic 

types of partial inflorescences themselves. These may 

be of either limited (determinate) or unlimited (indeterminate) growth. 

The use of a very simplified terminology is highly 

desirable until detailed morphological analyses are 

carried out. For general descriptive words, such as 

“bract,” “partial inflorescences,” or the concept of 

“complex inflorescences,” we refer to Troll (1964). In 

his more modern typological terminology the term 

“synflorescence” could be apDlied to the bamhoo 

flowering system, but this would require an elaborate morphological analysis. More details on this 

subject are given in a previous paper (Calder6n & 

Soderstrom, 1973), and for general meanings of 

different bamboo structures we refer the reader to 

the glossaries in two of McClure’s publications 

(1966, 1973). 

In the following keys to bamboos we have given 

particular emphasis to the vegetative characters that 

are generally available, and for the herbaceous 

genera we have used both floral and vegetative 

characters since these are commonly present. In the 

keys to bamboos we have added floral characters as 

supplementary information. Following the keys we 

have listed the genera of Bambusoideae and for 

each genus we have given the author, type-species, 

type-locality, and a few notes on distribution and 

ecology. 

Because the concept Bambusoideae as we understand it today has taken almost two centuries to 

develop, we have followed its history and tried to 

clarify its development and present this in the following chapter. Confusion has reigned not only in 

the systematics of this unusual group of grasses but 

in its nomenclature as well, and only by understanding its history could we identify those responsible 

for conceptualizing the subfamily and the major 

divisions within it. 

Rupert Barneby of the New York Botanical Garden 

and to Dr. Dan Nicolson of the Smithsonian Institution for advice on nomenclatural problems and 

for their careful review of the manuscript, We owe 

a continuous debt of gratitude to Dr. Jose Cuatrecasas of the Smithsonian for his illuminating discussions of morphology, advice on nomenclature, 

and ever-ready help in all botanical problems, big 

and small. 

ACKNOWLEDGMENTS.-we are grateful to Dr. 

History of the Bambusoideae 

In 1763 Adanson, in his treatment of Gramineae 

for the families of flowering plants, divided the 

family into several natural groups on the basis of a 

combination of characters, both floral and vegetative. His major groups are maintained to this day, 

the Poae and Panica being two of the most important, presently called tribes Poeae and Paniceae. 

The beautifully executed and printed grass flora 

of New Holland (Australia) by Robert Brown in 

1810 presented early ideas on relationships within 

the family. Brown remarked that, while grasses were 

connected in a reticulate fashion, he was nonetheless able to recognize three groups, delimited chiefly 

by modifications in the fruiting (spikelet) structure. 

The group of genera that contained Poa was characteristic of temperate regions and the other, which 

contained Panicum, was characteristic of hot and 

tropical regions. He was familiar with both groups, 

the former in his native England and the latter in 

tAle warm regions of Australia. Brown also listed a 

third group that contained a mixture of genera, 

which he admitted was scarcely natural. In 1814 he 

contributed a treatise on the botany of Terra 

Australis (Australia) for the report of the voyage 

during 1801-1803 of the Investigator, of which 

Matthew Flinders was commander and Brown, him- 
NUMBER 44 5 

self, the naturalist. In this account, Brown clarified 

his earlier remarks on grasses and divided the family 

into two tribes, which he called the Paniceae and 

Poaceae. He gave a very good explanation of the 

spikelets in each group and how they differ. Many 

later authors have interpreted these tribes of Brown 

to represent what many regard today as the subfamilies Panicoideae and Pooideae. While Adanson 

first named these two groups and characterized 

them, Brown elaborated upon them. 

The genius of Charles Kunth is apparent early 

in the career of this German botanist who, at the 

age of 27, published a highly significant paper on 

grasses (1815) while he was at Paris. “Having had 

the occasion to examine a large number of species 

of grasses, living [plants] as well as [specimens] in 

the rich herbaria of this capital [Paris],” he was convinced that the genera could be ordered into many 

natural groups. He presented a list of 10 such 

groups, among them the familiar Paniceae (as 

“Gramina Panicea”). His tenth, the bamboos (as 

“Gramina Bambusacea”), to our knowledge marks 

the first appearance of this group in a system of 

grasses. Kunth’s 10 groups were repeated in a later 

treatment of the grass family in his Synopsis Plantarum of 1822. 

During the same year there appeared in Sweden 

the Aphorismi of Agardh, in which nine groups of 

grasses were listed, one of them bamboos. It is likely 

that Agardh’s division of the family was modeled 

on the ideas presented by Kunth in 1815, since the 

groups are almost identical. 

Kunth (1815, 1822b) and Agardh (1822) were thus 

evidently the first authors to regard bamboos as a 

separate category of grasses. They included the few 

genera known at the time. 

In 1827 Link listed the plants in cultivation in 

the botanical garden in Berlin, of which he was 

curator. Then, as now, one of the widely cultivated 

bamboos was Bambusa glaucescens, an oriental 

species with small foliage and slender branches, 

which adapts easily to indoor culture. Link called 

the bamboo Ludolfia and placed it in the Triglossae, one of two sections that he erected for the 

bamboos. His other section was called Bracteiflorae 

and in this he included Nastus and Bambusa arundinacea, the type of the genus, which is commonly 

called the “thorny bamboo of India.” Link’s sections 

were admirably based on characters from the inflorescence and spikelet, but faulty observations regarding stamen number caused him to include 

species of the same genus in each of his sections. 

It was not until 1829 and 1835 that the bamboos 

were to receive a much more detailed treatment by 

the gifted Christian Nees von Esenbeck, a close 

friend of Goethe and a renowned botanist interested 

in all branches of natural history, philosophy, and 

social ethics. His 1829 treatment of the grasses of 

Brazil is a masterpiece. Bamboos constitute his 

tenth group, which he further divided into two 

parts, including in one of them the herbaceous 

genus Streptochaeta. 

The 1835 work of Nees dealt solely with the bamboos of Brazil, a group that must have intrigued 

him during his earlier study of the grasses of that 

country. He now divided the bamboos into three 

groups: Bambuseae (including Bambusa); Arundinariae (including Arundinaria); and a third, without a name, that contained Streptochaeta. However, 

the latter was listed as Streptochaeteae the following 

year (1836) in his enumeration of grass genera in 

Lindley’s Natural History oj Botany. 

In the above system Nees gave the genus Bambusa a detailed treatment and divided it into two 

subgenera, Bambusa and Guadua. The former contained species native to the Old World and the latter species native to the New World. Many authors 

have since regarded them as independent genera, 

but Hackel (1903) and McClure (1973) are among 

those who have followed Nees. 

In our opinion Nees’ 1835 paper was not only the 

first taxonomic monograph to be offered for bamboos but also one that embodied the most natural 

system, as we interpret it today, Bambusa and 

Arundinaria being among our woody bambusoid 

grasses (bamboos) and Streptochaeta an herbaceous 

bambusoid (Calderon 8c Soderstrom, 1973). Further, 

Bambusa belongs to those bamboos that have 

sympodial (pachymorph) rhizomes and pseudospikelets, and Arundinaria belongs to those that 

have monopodial (leptomorph) rhizomes and spikelets of the usual grass type. It is significant that Nees 

recognized the relationship of an herbaceous grass 

like Streptochaeta to the woody bamboos and accorded the genus equal rank. This paper is a testament to the perceptive mind of one of the greatest 

agrostologists of all times, yet it went unheeded 

and unnoticed for over a century after its publication. 

Few authors after Nees accepted his placement of 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

herbaceous and woody members together. Most, in 

fact, followed the lead of the Russian botanist, 

Franz Ruprecht, who, in 1839, published a large 

monograph on the bamboos. In his lengthy introduction, Ruprecht remarked that, whatever others 

might say, bamboos form a natural subfamily or 

order of grasses wholly connected to the other members. Ruprecht placed the 67 species of woody bamboos in two groups, following the system of Nees. 

Ruprecht’s contribution was not to the system of 

classification of bamboos but rather to the extensive 

worldwide treatment that he gave them. He devoted 

several pages to the morphology of the plant, discussed geographical distribution, and gave descriptions and keys to all the species. In its restriction to 

woody bamboos, this monograph had a greater influence on later workers than did that of Nees. 

The excellent monograph of Colonel Munro 

(1868) followed that of Ruprecht in its exclusive 

treatment of woody genera. Distinguished both as 

soldier and botanist, Munro was an outstanding 

agrostologist, who was often consulted by his colleague, George Bentham, on grass problems. Munro 

formulated three groups to accommodate his bamboo genera. The first two were those of Nees, but he 

added a new one, the Bacciferae, to contain the genus 

Melocanna and others from Asia. He divided the 

first group (which he called Triglossae out of personal preference) into three parts, the Arundinariae, 

Arthrostylideae, and Chusqueae, the latter two containing the New World genera with which he dealt. 

Munro’s system rested upon the foundation laid 

by Nees half a century earlier for the genera of 

Brazil but expanded to cover bamboos of the world. 

His system was thus more complete and introduced 

a whole new group, the Bacciferae, of eight genera, 

all Asiatic, All the groups and subgroups that 

Munro recognized have remained in systems to this 

day, albeit at different rank or with different name. 

Like Nees, Munro had a ke,-n perception of natural 

relationships in bamboos. 

In 1881 George Bentham published his classic 

paper, “Notes on Gramineae.” His friend, William 

Munro, had recently died and Bentham lauded the 

colonel upon whom he had depended for ideas regarding natural relationships in the grass family. 

In this paper, Bentham listed four groups of bamboos, adopting the concepts of Nees and Munro, 

with slight modifications in name and rank. 

The treatment of grasses that appeared in 1887 

in Die Naturlichen Pflanzenfamilien by the Austrian agrostologist, Eduard Hackel, has had a tremendous influence up to the present time. The major divisions of grasses that he enumerated were 

followed by most subsequent investigators. He divided the bamboos into four groups, which were 

exactly the same as those of Bentham. Hackel’s expertise in grasses such as Andropogon and Festuca 

did not extend to the bamboos, especially not to the 

herbaceous ones, which he dispersed among several 

distantly related groups. 

The most recent treatment of grasses in Die 

Naturlichen PfZanzenfamilien was prepared by 

Robert Pilger (1954), who omitted the woody genera 

of Bambusoideae. He did, however, assemble the 

majority of herbaceous monoecious bambusoid 

genera in one subfamily, which he called Olyroideae, and established another just for the herbaceous 

bamboo, Anomochloa. In this context it is interesting to note that Pilger recognized nine subfamilies, 

three of which are based on genera that we regard 

as bambusoid. 

At the time of Hackel’s paper, Franchet (1887) 

described three new herbaceous genera of grasses 

from tropical IVest Africa: Atractocarpa, Guaduella, 

and Puelia, all of which he stated to be bambusoid. 

The superficial similarity in spikelets of one of the 

genera to those of the American bamboo, Guadua, 

in fact prompted its diminutive name. 

The most complete monograph on Indian bamboos was published in 1896 by James S. Gamble, 

who was director of the Imperial Forest School at 

Dehra Dun, India, in the latter part of the past 

century. Although the monograph is a highly valuable treatise, it did not contribute anything new 

to the system of classification since Gamble employed the same four groups as Bentham. 

In 1913 Edmond-Gustave Camus published a 

monograph on bamboos, which was little more than 

a compilation of the literature to that time, although it did present a slightly modified system. 

His major groups were those already recognized 

by Nunro except for the inclusion of Franchet’s 

herbaceous genera and some alterations in group 

names and rank. 

Aimee Camus,like her father, Edmond-Gustave, 

worked at Paris and published many new genera 

and spccies of grasses, especially from the French 

colonies of the time. Her classification of bamboos 

(1935) differed little from her father’s except for 
NUMBER 44 7 

the rearrangement of some existing tribes and the 

addition of two new subdivisions to accommodate 

recently described genera of Madagascan bamboos. 

Both the Camusian systems incorporated woody 

genera. Interestingly, the only herbaceous ones 

were, however, those of their compatriot, Adrien 

Franchet. 

The characters used in systems of classification 

up through the time of A. Camus were for the 

most part floral, little importance being attached 

to vegetative structures. Probably for this reason 

Nees included Streptochaeta with the bamboos; the 

hermaphrodite spikelet, three lodicules, six stamens, and three stigmas were similar enough to 

those of bamboos to justify inclusion with them. 

A few other authors (Meisner, 1843; Steudel, 1855; 

Doell, 1880) followed Nees and placed that genus, 

along with Streptogyna (from tropical America, 

Africa, and Asia), next to bamboos. Trinius (1820), 

however, aligned Streptogyna with Bromus, and it 

remained with that genus through recent systems, 

an indication of his strong influence. 

Other herbaceous broad-leaved forest grasses 

like Leptaspis, Olyra, Pariana, and Pharus, although closely resembling the above genera, were 

seldom associated with them. Grisebach (1864) did 

recognize the relationship of Pariana to bamboos, 

but most other authors misinterpreted its spikelike 

inflorescence as evidence of a close affinity to members of the Hordeae. The unisexual spikelets and 

monoecious condition, common to all of these 

genera but not features of any woody bamboo, 

probably account for the fact that these grasses 

were all placed elsewhere in most systems. 

’IYhile Kunth, in his 1815 paper, regarded Olyra 

as representing a distinct group of grasses related 

to the bamboos, he was soon thereafter influenced 

by his contemporary, Carl Trinius, who treated 

the genus with Panicum in his monograph on that 

and allied genera (1826). Superficially the anthecia 

of Panicum and Olyra are indeed remarkably similar and many astute botanists since the time of 

Trinius have accepted without question the opinion of this influential Russian agrostologist. In fact, 

this disposition was endorsed by Hackel (1887) 

and adhered to by his many disciples to the present. Some other authors felt that the monoecious 

condition of these genera was an indication of a 

natural alliance between Olyra, Zea, and Zizania, 

the latter two genera belonging to widely unrelated groups by our present understanding. While 

not linked in early systems with bamboos, some of 

these monoecious genera were placed together 

since early times: Pharus and Leptaspis, for example, had been associated with Olyra by Kunth 

(1815), Agardh (1822), and Link (1827). 

It is well to recall the early words of Adanson 

(1763), who remarked that botanists were prejudiced in favor of divisions based on floral characters. He considered this a pitfall and, in establishing his natural groups of grasses, felt that all 

parts of the plant should be considered together. 

But his advice was not heeded until 1931, when 

Avdulov brought together data from many lines 

of investigation that had been developing in the 

grasses since the end of the nineteenth century. 

Study of the anatomy of the grass leaf has shown 

that there are great structural differences between 

major groups, and anatomical characters are now 

used widely in grass systematics. Of the numerous 

anatomical papers that we have previously cited 

(Calderon & Soderstrom, 1973:36), that of Brandis 

(1907) is of particular interest here. In his welldocumented paper, Brandis referred to the similarities of the structure between the leaves of bamboos and those of several herbaceous grasses such 

as Olyra, Diandrolyra, and Pharus. A short time 

later, on the basis of leaf anatomy, Krause (1909) 

divided the family into three major groups, of 

which one was the Bambusoideae. The findings 

of Brandis were corroborated by Michaud-Page 

(1947) and Metcalfe (1956), who pointed out that 

leaves of Streptochaeta, as well as some other herbaceous genera, had the same anatomical structure 

as those of bamboos. 

The embryo, endosperm, and seedling have also 

provided characters of systematic value. Van Tieghem (1897) studied the embryo and seedling and 

was the first to point out their value in distinguishing major groups of genera. His work has been 

added to by the embryo studies of Yakovlev (1950), 

Kinges (1961), and Reeder (1962). In the latter 

study, the author pointed out similarities between 

embryos of the “olyroid” genera and bamboos. 

Characters of the endosperm have provided the 

basis of a system of classification developed by 

Hayek (1925). 

The detailed morphological and anatomical 

studies that became fashionable toward the end of 

the last century, and have continued since, were 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

directed as well to the grass inflorescence and 

spikelet. The unusual spikelet of Streptochaeta has 

intrigued most serious agrostologists since the 

genus was described by Nees in 1829 and placed 

by him with the bamboos. Celakovsky (1889) published detailed morphological studies of the spikelet and interpreted the flower as the most primitive 

of any grass, an idea that has persisted to the 

present. The classic paper on the morphology of 

the grass flower by Goebel’s student Schuster (1909) 

corroborated the opinion of Celakovsky. In his 

phylogenetic scheme for the family, Schuster regarded Streptochaeta as the type closest to the 

hypothetical ancestor of the family. Several other 

investigators, such as Arber (1929) and MichaudPage (195 I), have tackled this well-studied genus 

and published still further scholarly accounts. 

The appearance of significant papers by Avdulov 

and Prat in 1931 marks this year as a milestone 

in the development of a modern system of classification for the grasses. Nikolai Pavlovic Avdulov 

was a cytologist at the Institute of Applied Botany 

and New Cultures in Leningrad, of which N. I. 

Vavilov was director. Henri Prat was a French 

anatomist, who had recently completed his thesis 

on a study of grass leaf epidermis. 

Avdulov, working under the cytologist G. A. 

Levitskii, made a survey of the karyotype of all 

the grasses he was able to obtain. He studied the 

chromosome number, size, and form, in addition 

to the seedling and leaf anatomy. Combining his 

extensive data with those of others, he was able 

to develop an original system of classification. 

Prat (193 1) showed that the grass leaf epidermis 

also provided numerous characters of systematic 

value. He expanded upon the data presented in 

that paper and in 1936 offered a new system, incorporating his own data with those of Avdulov 

and others. While Avdulov had presented little 

data on bamboos-due to lack of study materialPrat recognized the bambusoid as a major division 

of the family. 

Avdulov had set a precedent for evolutionary 

considerations in the grasses and made them the 

basis of a new phylogenetic scheme. His colleague, 

Roman Yulievic Roshevits, an agrostologist at the 

Botanic Garden in Leningrad, aided Avdulov in 

identifying the grasses that the latter used in his 

studies. In 1937 Roshevits published a large treatise on grasses of the world. 

In his classification Roshevits followed Avdulov 

with some modifications and presented ideas on 

evolution in the family derived mainly from 

Schuster’s phylogenetic interpretation of Streptochaeta and other bamboo genera. In 1946 he presented another scheme in which his subfamily 

Bambusoideae contained six tribes, five consisting 

of herbaceous and one of woody genera. Hubbard’s 

system (1934), in which many new tribes were described, had appeared between the two papers of 

Roshevits and in the second the author adopts 

Hubbard’s new tribal names. In the 1946 scheme 

of Bambusoideae, Roshevits did not give a description of the subfamily nor his reasons for the tribal 

arrangement, stating that he would do so in a 

later paper. 

Jacques-Felix (1955) published a paper on primitive African grasses, which he brought together in 

his bambusoid series. The author stressed the leaf 

anatomy common to all the genera, which he allocated to four tribes that included both woody and 

herbaceous genera. This paper established the 

modern concept of the Bambusoideae, based on a 

combination of natural relationships. 

A few years later Parodi (1961) formalized the 

nomenclatural status of the subfamily Bambusoideae and in his diagnosis added further substantiating characters from embryo structure, 

chromosome type and number, geographical distribution, and ecology. In this particular paper 

Parodi dealt only with the genera of grasses from 

Argentina so that his subfamily Bambusoideae circumscribed only the tribes with Argentinian representatives. In agreement with this concept, we 

are following Parodi’s system, extending it to 

cover, in the present paper, the whole of the 

American continent. 

A few systems of Bambusoideae have, however, 

confined themselves to the woody genera. Stebbins 

& Crampton (1961), in their system of grasses for 

temperate North America, felt that the commingling of herbaceous and woody members would 

result in a heterogeneous subfamily. They therefore recognized only Arundinaria as bambusoid 

and relegated to the Oryzoideae the herbaceous 

genera Olyra and Pharus. Grosser & Liese (1973) 

presented a classification of the woody bamboos, 

based on characters from culm anatomy of a number of Asiatic genera they studied. 

The most eminent bamboo specialists of recent 
NUMBER 44 9 

times have been Richard E. Holttum, of England, 

and Floyd A. McClure, of the United States, both 

of whom have made significant contributions to 

our knowledge of woody bamboos. Perhaps because 

both had such extensive experience with bamboos 

in the field, the former in Singapore and Malaysia 

and the latter in China and tropical America, they 

stressed the value of vegetative characters in this 

group, whose members flower so seldom, without 

devaluating those of the inflorescence, some of 

which are unique in the grass family. 

Holttum (1956) commented upon some morphological characters that previous authors had used 

in their systems of classification and divided the 

bamboos into four groups, using characters from 

the gynoecium He did not attach formal names 

to the groups nor did he bring herbaceous genera 

into his discussion. 

McClure (1961) limited his detailed description 

of the subfamily Bambusoideae to his own group 

of expertise, the woody bamboos. He did, however, 

later agree with the concept of Bambusoideae offered by Parodi in 1961, to which he refers in a 

revision of bamboos of Santa Catarina, Brazil 

(McClure & Smith, 1967). 

N. N. Tsvelev, of the Komarov Botanical Institute, has published a number of papers on evolution in the grass family. Of particular interest to 

us is his system of classification proposed in 1968, 

in which he divided the family into two subfamilies, the Bambusoideae and Pooideae, each further 

divided into a number of tribes. Tsvelev recognized the unique set of characters shared by genera 

of the Bambusoideae, which collectively form a 

group of equivalent rank to that formed by all 

other members of the family. In his revision of the 

grasses of the Soviet Union, Tsvelev (1976) employed this new system. 

We recognize the Bambusoideae as a subfamily 

of the Poaceae, with a set of characters that binds 

together a large number of herbaceous and woody 

genera. To most agrostologists today the Bambusoideae is one of six or seven subfamilies of 

grasses; to Tsvelev it is one of two. 

Nomenclature 

In 1812 Palisot de Beauvois described 69 new 

genera and 640 new species of grasses and proposed 

many new binomials and new names. He divided 

the grasses into families, tribes, cohorts, and sections, and while all of these appeared in his 

“Tabula Methodica,” none of them is given formal 

description. Trinius, a short time later (1820), in 

his Fundamenta Agrostog-raphiae, continued to use 

the old Linnean system, and his genera of grasses 

are to be found under such categories as “Triandria Monogynia” and “Monoecia Hexandria Monogynia.” 

As the number of described genera of grasses 

increased, especially during the first half of the 

last century, the idea of tribes was developed in 

order to bring together genera with similar characters. Robert Brown, in fact, used the term “tribe” 

for the Poaceae and Paniceae, which he characterized in 1814. In the following year Kunth stated 

that “there is not at all a natural distribution of 

genera into tribes as has been done with success 

in other plant families. However, the possibility 

of this distribution should not be denied, which 

is the principal object of this memoir.” Kunth 

listed 10 tribes, for which he used such Latin 

group names as “Gramina Panicea” and “Gramina 

Stipacea.” In a later paper (1822b), he repeated 

his 10 groups of 1815, but called each a “Sectio” 

and gave new endings, for example, “Sectio 1. 

Paniceae and Sectio 2. Stipaceae.” Link (1827) 

offered a more elaborate system than his predecessors and divided the grasses into sections, but 

these were not comparable to those of Kunth, for 

his sectional category was superior to those of 

family, division, and suborder, terms that he used 

interchangeably. Thus the Paniceae was a suborder of a section and the Stipaceae a division of 

his section called “Paniculatae Uniflorae.” 

It would serve no purpose to review other systems of the time except to point out two items 

that are already obvious: (1) agrostologists recog 

nized that the genera of grasses could be grouped 

together on similar characters and (2) there was no 

agreement on names for these groups. For example, in the case of the well-known genus Panicum, 

Adanson established a section called “Panica.” 

Another section was his “Poae,” which included 

several genera he felt related to Poa. Robert Brown 

recognized the Paniceae, calling it a “tribe,” characterized it further, and added more genera. Kunth 

recognized the same group, as one of his sections, 

but called it “Gramina Panicea” in 1815 and later 

just “Paniceae” (1 82213). To Link, the Paniceae was 

a suborder of a section. 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Regardless of what authors called the group of 

genera that included Panicum, they agreed that it 

was one of the natural divisions of grasses. Today 

we call a group of related genera a “tribe” and 

for this group we use the tribe Paniceae. 

The first author to recognize and establish that 

Panicum and allies represented a natural group 

was Adanson. By present definition we regard this 

group as tribe Paniceae, of which the authorship 

must be credited to Adanson. Article 19 of the 

Zn t e 1.n a t i o n a 1 Code of B o t a n ica 1 No m e n c 1 a t u re 

(ZCBN), note 2, states that when an improper termination has been published, it should be changed 

to accord with the present rules. We thus change 

“Panica” to “Paniceae,” but still give Adanson 

credit for establishing the tribe. 

In the subfamily of special concern to us, the 

Bambusoideae, there is little agreement in the 

literature regarding authors of tribes such as 

Olyreae and Bambuseae, or even of the subfamily 

itself. For example, Butzin (1973) credited Munro 

(1868) as author of the subfamily, while McClure, 

in his various publications, gave credit to others. 

In our opinion credit should go to Nees von Esenbeck (1835), who first recognized the bamboos and 

allies as constituting a major natural division of 

the family. 

In order to decide who was responsible for each 

category, we have followed the development of the 

concept of the bamboos as a subfamily of grasses. 

In tracing the ideas from one author to another, 

we have attempted to follow the development of 

the concept of natural groups of bambusoid genera 

(which we call “tribes”) and the aggregation of 

smaller groups into a large one (which we call 

“subfamily”). 

In presenting the history of the Bambusoideae 

above, we referred to these aggregates of genera 

as “groups,” to avoid using the actual terms of the 

authors, since there was no uniformity in these 

terms. For the tribe Bambuseae, we thus give 

credit to Kunth who, in 1815, recognized the bamboos as a natural group. Nees von Esenbeck, in 

1835, went further and established a larger group 

-subdivided into three parts, two containing 

woody genera and one an herbaceous genus-similar to the system we recognize today at the subfamily level. Tl’e thus give credit to Nees for the 

recognition of the bamboos as a subfamily of 

grasses. 

Later authors recognized even further groups of 

genera of bamboos. Munro’s monograph of 1868 

covered the Bambusoideae in the sense of our subfamily, even though he called it a tribe and named 

it “Bambusaceae.” Munro divided his Bambusaceae 

into sections and subsections, two of which, Arthrostylidiae and Chusqueae, were recognized as 

natural groups for the first time. Today these 

groups are regarded as tribes and we give credit 

to him as their author. 

Article 35 of the ICBN states that “a new name 

published on or after 1 Jan 1953 without a clear 

indication of the rank of the taxon concerned is 

not validly published. For such names published 

before 1 Jan 1953 the choice made by the first 

author who assigned a definite rank is to be followed.” 

As we have pointed out, there was no uniformity 

in the names of rank applied to grass taxa in the 

older literature. Since different names were used 

interchangeably for the same intended rank, they 

cannot be accepted in a literal sense. For example, 

Bambusa and allied genera constituted a section 

in the systems of Kunth (1822b) and Link (1827). 

Several years later, however, Link (1833) made 

them a family “Bambusaceae,” which he listed as 

a subdivision of the section. On the other hand, 

Meisner (1843) called this group a subtribe. 

We must look to the first author who recognized the taxon, whatever rank name he gave it, 

and assign the rank name that applies to the group 

by our present understanding. We have followed 

this way of thinking in our interpretation of the 

nomenclature of bambusoid taxa prior to 1 January 1953, when guidelines were lacking. Since that 

date the rules for valid publication, as set forth 

in the ZCBX, must be followed. 

Sometimes the selection of the authorship of a 

subfamily was based solely on the use of the first 

correct spelling by modern rules, an interpretation 

with which we do not agree. For this reason, 

McClure (1966) cited Rehder (1945) as author of 

the subfamily Bambusoideae of the “Gramineae” 

and Ascherson & Graebner (1902) as authors of the 

subfamily Bambusoideae of the “Gramina.” It was 

not until late in the last century that the term 

“subfamily” was in fact used as a category for a 

major subdivision of the grass family. h’ees von 

Esenbeck’s concept of the bamboos as constituting 

a major natural division of the grass family thus 
NUMBER 44 

preceded both the term we use for that group 

today and the rules that govern its correct Latin 

spelling. 

Characterization of the Bambusoideae versus 

Other Grasses 

BAMsusoIDEAE.-Perennial herbaceous or woody 

and arborescent, but without secondary thickening, 

rhizomatous; culms when woody branched at the 

nodes, the branches in some species with thorns or 

spines; leaf blades usually flat, broad, lanceolate or 

linear-lanceolate, articulate with the sheath by a 

petiole that twists and moves the blade in different 

positions; blades with tessellate venation strongly 

or weakly manifest superficially; flowering not seasonal, in some cases (bamboo) in cycles of long intervals of several years or almost continuous through 

several months of the year; inflorescences of different types, usually composed of complex systems of 

partial inflorescences with a limited or unlimited 

and continuing ramification and production of new 

partial inflorescences of restricted or unrestricted 

type (pseudospikelets), usually with very welldeveloped bracts (subtending leaves and prophylla); 

spikelets or pseudospikelets 1- to many-flowered, 

without glumes or with 2 to several “transitional 

glumes;” lemma (3-)5- to many-nerved, not awned 

or only seldom so, and the awn then not geniculate; 

palea 2- to many-nerved, keeled or rounded dorsally, exceptionally bifid; lodicules generally 3 

(rarely 0 to 6 or many), usually large and with 

hairs of different types and well-developed vascularization; stamens 3 to 6 or many (rarely 2), sometimes partially fused or monadelphous; stigmas 2 

or 3 (rarely 1); fruit of variable type, dry (caryopsis 

or achene) or fleshy (drupe or berry-like), sometimes 

very large and many times the size of the original 

gynoecium; hilum linear, almost as long as the 

fruit; embryo several times smaller than the fruit. 

Seedling: Coleoptile short and not elevated from 

the caryopsis by an internode, first two to several 

leaves bladeless or with a reduced blade, the first 

expanded blade broad, ovate or lanceolate, horizontal in position. 

Leaf Anatomy: Leaf blade (lamina) with a very 

conspicuous midrib containing a complex vascular 

system of several bundles in two rows, strongly 

developed sclerenchyma and ground tissue; mesophyll with cells arranged in horizontal layers par11 

allel to the epidermis, not radiate; chlorenchyma 

composed of arm cells and translucent fusoid cells 

at each side of the vascular bundle and in between 

layers of arm cells; bundle sheaths always double 

and well developed, the outer sheath with very few 

chloroplasts; transverse veinlets connecting the 

longitudinal vascular bundles present. 

Leaf Epideymis: With short cells in pairs or 

sometimes in rows over the veins; silica bodies 

usually cross-shaped, saddle-shaped, of olyroid type, 

or of intermediate forms; microhairs nearly always 

present, bicellular with both cells of about the 

same length and with rounded apex, microhairs of 

3 or 4 cells sometimes occurring; papillae common 

and abundant on the long cells and overarching 

the stomata; long cells with thick sinuous walls; 

stomata usually with low dome-shaped or sometimes triangular subsidiary cells. 

Physiology: C, pathway of CO, fixation. 

Cytology: Chromosomes small, x = 12, with most 

bamboos tetraploid (2n=48) or hexaploid (2n=72), 

and herbaceous bamboos diploid or polyploid, with 

n numbers of 7, 9, 10, 11, and 12 reported. 

Ecology and Geographical Distribution: Plants 

growing in association with trees or shrubs, usually 

in warm temperate woodlands, tropical rain forest 

or, if herbaceous, in the shaded understory of warm 

forests, also along streams or, if in rather open 

areas, always in shade of taller vegetation; usually 

dependent on humidity, shade, and warm temperatures in the environment; abundant in the 

tropics and subtropics of the world, only some 

woody members reaching temperate-cold areas of 

both hemispheres. 

GRASSES (excluding Bambusoideae). - Perennial 

or annual herbs, sometimes tall and robust but 

neither with lignified culms nor complex ramification at nodes of the culm, nor thorns, nor spines; 

cespitose, stoloniferous or rhizomatous, but then 

not with strong complex systems; nodes with only 

one bud or none; plants without a special growth 

pattern, i.e., not producing differentiated elongated 

new culms, the leaves all alike and produced progressively as the plant grows; leaf blades usually 

linear and if lanceolate petiolate, not deciduous; 

outer ligule never present; flowering seasonal, with 

one flowering period each year, either in spring or 

summer, according to the group; inflorescence spiciform, racemiform, or paniculiform, usually lacking 

bracts and prophylla and not forming complex sys- 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tems of ramification; pseudospikelets not formed; 

spikelets 1- to many-flowered, usually with 2 

glumes; lemma awnless or with an awn, this sometimes geniculate; palea usually 2-nerved and 2- 

keeled; lodicules usually 2, relatively small; stamens 

usually 3; stigmas 2. Fruit a caryopsis, exceptionally 

of a different type, but then not fleshy: hilum punctiform, ovate or linear, short or long; embryo small 

or large. 

Seedling: Of different types according to the 

group, but not bambusoid. 

Leaf Anatomy and Epidermis: Of different types 

but not bambusoid. 

Physiology: C, and C, pathways of CO, fixation 

according to the group. 

Cytology: Chromosomes small or large, x=7, 9, 

10, and sometimes 12. 

Ecology and Geographical Distribution: Forming 

dense covers in open, sunny places, being the main 

component of prairies, steppes, savannas, and 

mountain slopes; common elements of the herbaceous vegetation from dry or arid zones to humid 

ones, from cold, temperate to warm, tropical, 

reaching the extreme limits of vegetation in the 

mountains or near the poles of both hemispheres; 

relatively few species adapted to the dark conditions of the woodlands or tropical forests. 

Characterization of Woody versus 

Herbaceous Bamboos 

WOODY BAMBoos.-Plants with a woody, strongly 

segmented cauline axis, often arborescent and with 

a prominently developed rhizome system; plants 

with a special growth pattern, producing at the 

initial period unbranched, elongated new shoots 

covered by overlapping modified leaves (“culm 

leaves”), and at the nodes one to multiple buds arranged in different patterns, leading when developed to complex systems of ramification; leaves 

dimorphic, those covering the new culm (“culm 

leaves”) with a greater deyelopment in the sheath, 

the blade usually smaller and of variable shape and 

orientation, caducous or persistent, the petiole 

sometimes lacking, auricles and oral setae often 

greatly developed; foliage leaves usually with a 

lanceolate blade, articulate to the persistent sheath 

by a petinle and deciduous, and often with tessellate 

venation strongly or weakly manifest, outer ligule 

commonly developed, auricles and oral setae 

usually well developed; flowering in cycles of 

several-year intervals, the individual usually monocarpic; inflorescences of different types in complex 

systems of ramification, with production of partial 

inflorescences either limited (one) or unlimited as 

the result of continuous ramification and production of several partial inflorescences of restricted or 

unrestricted type (pseudospikelets); partial inflorescences composed of 1 to many spikelets arranged 

in racemiform or paniculiform fashion; spikelets or 

pseudospikelets 1- to many-flowered, the flowers all 

hermaphrodite; fruit a caryopsis, achene, or fleshy. 

Plants common in tropical or subtropical lowlands, 

some reaching the temperate or cold regions and 

altitudes up to 4500 m, usually associated with 

woody plants. 

HERBACEOUS BAMBoos.-Herbaceous, very small 

and delicate to robust plants, but without lignified 

culms and with rhizomes only weakly to moderately 

developed, never so strongly as in the bamboos; 

plants without a special pattern of development, 

not producing tall, unbranched new shoots; nodes 

with only one bud, not developing vegetative 

branches or if exceptionally so, then the branch 

solitary; culms never armed; leaves not dimorphic, 

with only foliage leaves formed; blades lanceolate 

or exceptionally linear, not deciduous or exceptionally so, and rarely with manifest tessellate venation; 

outer ligule generally lacking; auricles lacking or 

weakly developed; oral setae lacking except in the 

tribe Parianeae; flowering almost continuous or for 

several months of the year, not in cycles of several 

years; inflorescences of different types, sometimes 

in complex systems resulting from the continuous 

ramification and production of new partial inflorescences; pseudospikelets not produced; spikelets 

l-flowered, hermaphrodite or unisexual (the plants 

then monoecious) or many-flowered, the flowers 

then hermaphrodite or rarely polygamous, the female distal to the male; fruit a caryopsis, rarely an 

achene. Plants common in forests of the tropics 

and subtropics, not reaching the temperate or cold 

regions or generally elevations greater than 1000 

m, inhabiting the shade of forest or in more open 

places such as savannas, campos, or stream banks, 

sheltering under taller vegetation. 
NUMBER 44 13 

Key to the Tribes of American Herbaceous Bamboos 

1. Spikelets hermaphrodite ................................................ ................ 2 

as in the majority of grasses; outer ligule developed .......................................... ST~PTOGYNEAE 

Spikelets unisexual and 1 -flowered, dimorphic; pla us .............................................. 4 

2. Spikelets many-flowered; leaves lacking a petiole between sheath and blade, ribbon-like 

Spikelets 1-flowered; leaves with a petiole between sheath and blade; outer li 

developed; blade with strong tessellate venation ............................................................ 3. Leaves mostly basal, the sheaths strongly compressed-keeled, overlapping, and 

a fan-shaped fascicle; petioles up to 25 cm long; inflorescence of cymoid type, with large 

bracts developed; spikelets 2 or 3 in the axil of each bract; lodicules 0; a ring of hairs 

present around the base of the flower; stamens 4; stigma 1 (Bahia only) .......A NOMOCHLOUE 

Leaves distributed evenly along the culm, the sheaths rounded, not overlapping; petioles 

no longer than 1 cm when developed; inflorescence spicate, without large bracts developed; 

spikelets solitary at each node of the axis, without subtending bracts; lodicules 3, long 

lacking around the base of the flower; stamens 6; stigmas 3 

................................................................................... STREPTOCHAETEAE 

1 to but diverging from midvein; culms forming compressed 

fascicles, leaves long-petiolate, all in one plane in a fanlike arrangement; inflorescence 

paniculate, disarticulating and falling in sections of several branches; lemma of female 

spikelet papery or slightly indurated; stamens 6; stigmas 3 ........................................ PHAREAE 

Veins of blades parallel to the midvein as normal in Poaceae; plants otherwise; leaves 

5 

5. Rhizomes of monopodial type, spreading, the culms not forming very dense clumps; inflorescences terminal and solitary, usually not rebranching or producing secondary partial 

inflorescences; lateral inflorescences from nodes at axils of leaves not produced; inflorescence spiciform, disarticulating and falling in segments; stamens 2 to many; stigmas 2; 

...... PARIANEAE 

Rhizomes of sympodial type; culms appressed, forming dense, small or large clumps; 

inflorescences terminal and lateral, rebranching at base and producing 2 to several partial 

inflorescences of higher order in succession, exceptionally only 1; partial inflorescences 

paniculate or racemiform; stamens 3 ,(rarely 2); oral setae lacking .......................... OLYREAE 

short-petiolate; lemma of female spikelet leathery to indurate ............................... 

oral setae of leaves usually well developed 

Key to the Genera of American Herbaceous Bamboos 

1. Spikelets hermaphrodite .......................................................................................................................... 2 

2. Spikelets many flowered; leaves without a blade, ribbon-like as in 

the majority of grasses; outer ligule produced ...................................................... Streptogynn 

Spikelets 1 -flowered; leaves petiolate; outer ligule not developed 

3. Petiole of leaves up to 25 cm long; blade with a cordate base; 

large bracts; lodicules 0; stamens 4; stigma 1 ............................................................ Anomochloa 

Petiole less than 1 cm long; blades lanceolate; inflorescence without large bracts; lodicules 

4. Veins of blade not parallel to the midvein but diverging from it; culms forming compressed 

fascicles, the leaves all displayed in one plane in a fanlike arrangement; leaves longpetiolate; inflorescence paniculate, disarticulating and falling in sections of several 

branches ........................................................................................................................................ p~ 

Spikelets unisexual; plants monoecious .......................................... 4 

very long, 3; stamens 6; stigmas 3 .................................... .................................. ..Streptochueta 

Veins of blade parallel to the midvein; plants otherwise; leaves short-petiolate ..... 5. Inflorescence spiciform, disarticulating and falling in segments; oral setae us 

developed (Parianeae) .................................................................................................................... 6 

Inflorescence paniculate or racemiform; oral setae lacking (Olyreae) .......................................... 7 

6. Inflorescence consisting of several segments, disarticulating from each other and falling 

separately at maturity; inflorescences produced only at the apex of the main culms or 

on short bladeless culms; male spikelets with numerous stamens; stigmas plumose; sheaths 

....... ..Pariana 

Inflorescence completely covered by several bracts and only 1 terminal segment disarticulating at maturity; inflorescences produced as in Pariana in addition to subterranean 

(cleistogamous) ones; male spikelets with 2 stamens, their filaments fused; stigmas not 

with one or more scarlike marks toward the summit (Amazonia) ...... 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

10. 

11. 

12. 

13. 

14. 

15. 

16. 

17. 

18. 

19. 

plumose: sheaths with no scarlike marks (Bahia and northern Espirito Santo) ........ Eremitis 

Inflorescences usually produced on specialized culms (bladeless or with few blades), at the 

base of the main one, the inflorescence rarely developing at the apex of the main culm ..8 

Inflorescences always produced on main, leafy culms .................................................................... 11 

Spikelets of both sexes mixed in each inflorescence 

Female and male spikelets separated into 2 di 

Specialized culms decumbent, shorter or 

Cuba) .............................................................. 

to the West Indies) ...................................................................... .................... Piresia 

blades and several partial inflorescences; anthecium appressed-pilose (Bahia and Amazonia 

Specialized culms erect, much shorter than the main axis, besring only 1 fully developed 

blade at its apex and a single inflorescence; anthecium glabrous (Sao Paulo to Bahia)

Lemma of female spikelet obtuse or subacute, awnless; plant 

spreading, 5-15 cm tall ........................................ ................................................. niochloa 

Lemma of female spikelet acuminate, awned; nts firm, erect, 50-100 cm tall 

Ekmanochloa 

Inflorescence consisting of several partial inflorescences, each containing 2 to 4 clusters 

of 1 large female and several smaller male spikelets, the clusters falling as a unit; filaments 

of stamens fused into a tube (northeastein Amazonia and French Guiana) .... Froesiochloa 

12 

Basal internode of culms thickened, cormlike; pedicel of male spikelets lon he 

Basal internode of culms not thickened; pe r than the female 

spikelet .................................................................................................................................................... 13 

Female spikelets disarticulating below the glumes; glumes with a cushion-shaped, pulviniform base; pedicel of female spikelet not thickened at the summit; delicate plants with 

weak, decumbent, or trailing culms .................................................................. Raddiella 

Female spikelets disarticulating above the glumes; glumes wit 

pedicel of female spikelet thickened at the summit; plants otherwise .......... 

Sexes separated in different inflorescences; glumes of the female spikelet wit 

........................ 15 

Sexes mixed in the same inflorescence: glumes without cartilagi 

................................ 16 

Leaves many, crowded and distally imbricate, forming a definite leaf complement; partial 

inflorescences all lateral, the male predominant, these small and paniculiform, 2 or more 

developing at each node of the leaf complement; female inflorescences small and racenose, few developing mostly at the middle nodes of the culm (Venezuela, Guianas, 

d terminal, the 

female predominant, these racemose, several developing acropetally at each node; male 

inflorescences few, large and open paniculiform, developing at the summit of culm 

(eastern Brazil: Pernambuco to Rio de Janeiro) ..................................... Internode of rachilla between the glumes and anthecium elongated; 

produced at the summit of the culm and from several nodes below 

Internode of rachilla between the glumes and anthecium not elongate 

Anthecium laterally compressed, triangular in outline, zygomorphic (pi 

Anthecium dorsally compressed, fusiform, actinomorphic .................................. 

Inflorescence dense and contracted, borne singly at the summit of the culm; lemma of male 

spikelet similar in texture to glumes, 7- to 9-nerved: stigmas with short appressed processes, 

not plumose and curling at maturity; culms with only 1 to 3 blades developed (Panama) 

Inflorescence composed of several partial inflorescences produced at the summit of the 

culm and from 1 to few nodes below: lemma of male spikelet unlike that of female 

glumes, thin and papery, 3-nerved: stigmas with long processes, plumose, not curling 

at maturity; culms with most of the blades developed .............................................................. 19 

Inflorescence open, paniculate or digitate; female lemma many-nerved; female anthecium 

not mottled at maturity (Mexico to northern Argentina and West Indies) Olyra 

Inflorescence small, condensed: female lemma 5-nerved; female anthecium mottled at ma- 

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

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

Inflorescences of different types, never disarticulating and falling partially ......... 

female spikelet (Amazonas; Surinam) ...................... Rehia 

undulating margins; apex of blades asymmetrically apiculate 

not asymmetrically apiculate ...................................................... 

northern Brazil, and Bahia) ......................................... .......... Strephium 

Leaves few, evenly spaced along the culm; partial in 

................................................................................................................................ Maclurolyra 

turity (southeastern Brazil) ...................................................................................................... Rei~%ia 
NUMBER 44 15 

Key to the Genera of American Bamboos 

1. Rhizomes monopodial (leptomorph); midculm branch complement basally appressed, 

composed of 3 axes, the central one slightly dominant, all later rebranched: inflorescence 

solitary, restricted, of limited branching; stamens 3; stigmas 3 (temperate regions of the 

Rhizomes sympodial (pachymorph) (subtropical and tropical regions of both hemispheres)..2 

2. 

Culms in the vegetative state usually branched ........................ 

3. Culms highly modified, juncoid in both vegetative and flowering stages, with a few bladeless leaves developed only in the basal part: the greater part of each culm consisting 

of a single elongated internode, thin-walled and with pith septa at close intervals; large, 

paniculiform inflorescence: stamens 3: stigmas 2 (Serra dos OrgAos, state of Rio de 

Janeiro, Brazil) .................................................................................................................. Gla%~~hy~on 

Culms leafy, of a coarse grasslike appearance, with unusually large leaves up to 2 m long, 

the sheath and blade not strongly differentiated: internodes with more or less thick walls 

and without septa: inflorescence terminal to the culm, open or contracted: spikelets with 

1 fertile floret, 4 tra 

4. Branch complement at 

of limited branching 

Branch complement at 

Northern Hemisphere: 1 species native to continental United States) ............ Arundinaria 

Culms in the vegetative state usually not branched ................... 

5. Culm characterized by 1 or 2 usually elongated internodes separated by several short ones; 

culm leaves very thick and leathery with sessile or subsessile blade: sheath persistent, the 

blade deciduous: inflorescence long and narrow; spikelets usually 2-flowered (isolated 

tepuis in Venezuela) ............................................................................... Myriocladus 

Culms without elongated internodes, these evenly spaced and separated from each other 

by 1 node: culm leaves herbaceous, not leathery, with a petiole between the blade and 

sheath, tardily deciduous: inflorescence 

flowered ......................................................... 6. Branches of the branch complement slend .......................................... 7 

Branches of the branch complement of two sizes, a dominant larger branch and subsidiary 

ones, several times smaller 

7. Branches arising at differen 

leaving a triangular, unseg 

thick-walled ........................................................................... Branches arising all at the same level in a more or less horizontal line ...................................... 9 

8. Culm leaf petiolate: base of blade much narrower than upper edge of the sheath: blade 

reflexed, caducous; spikelets usually 1- or 2-flowered, shortly pedicellate and secund along 

the unbranched axis: fruit an achene .......................................................................... Meros~uc~~s 

Culm leaf without a petiole, falling entire: blade erect, appressed, sessile, at the base as 

wide as the upper edge of the sheath, the edges of both in a continuous line: spikelets 

many-flowered, solitary and sessile, alternat 

............................................................................. ................................ cladum 

tropics 

known as “pAramo”: rather small, erect bamboos with stiff culms, very short internodes 

and very thick leaves; inflorescences contracted, dense, usually narrow: spikelets with 1 

fertile floret and 4 transitional glumes (Costa Rica to northern Peru) ............ Swnllenochloa 

Plants of low to high elevations but never of “pAramo”: habit of plants otherwise; inflorescences usually loose, open; transitional glumes 2 .......................................................................... 10 

10. Midculm branch complement of 3 subequal axes with a common prominent or rounded, 

tapered base extended downward: culms very thick-walled with a small lumen: inflorescence paniculiform with numerous spikelets, lacking subtending bracts: transitional glumes 

not widely separated from the first anthecium (Mexico to Honduras) Otutea 

Midculm branch complement usually of 5 branches without a common rounded base: culms 

hollow or solid; inflorescence racemose, consisting of 1 to few spikelets, usually subtended 

by a large bract: transitional glumes widely separated from the first anthecium (central 

11. Plants with spines or thorns: culms hollow, sometimes robust: culm leaf deciduous, without 

a petiole: blade erect, sessile, more or less triangular: inflorescence unrestricted, of un1s ................ 

9. Plants characteristic of the vegetation above t 

and southern Brazil) .......... A poclada 
16 SMITHSONIAN CONTRIBUTIONS 

limited branching; pseudospikelets many-flowered; stamens 6; stigmas typically 3 

.................................................................................................... Bambusa subg. GuadM 

Plants without spines ................................................................... I2 

12. Culms solid .............................................................................. Culms hollow, with thin walls or, if the walls thick, wi 

13. Culms in cross section with air canals in a ring near the epidermis; young stages of bud 

complement with 3 subequal buds; branches developing intravaginally, i.e., the branches 

growing upward emerging from the summit of the sheath; culm leaf-blade reflexed, 

caducous; inflorescence unrestricted, of unlimited branching; pseudospikelets I-flowered; 

transitional glumes 0; stamens 3; stigmas 2 ................................................................ Atractantha 

Culms in cross section without manifest air canals; young stages of bud complement with 

more than 3, usually several buds, the central one several times larger than the lateral; 

each bud within a separate prophyllum; branches developing extravaginally, i.e., the 

branches emerging through the base of the sheath; culm leaf-blade erect, persistent; 

inflorescence restricted, single, of limited branching; transitional glumes 4; spikelets with 

14. Plants with a midculm branch complement of 3 subequal branches, the central one slightly 

dominant; culm leaves and foliage leaves with dense, long oral setae; inflorescence condensed, capitate; spikelets I- or 2-flowered; transitional glumes 2 (state of Rio de Janeiro 

Plants with a midculm branch ent of more than 3 branches, the central one very 

15. Nodal region delimited by 2 ridges, the upper one more prominent than the lower, forming 

a narrow crest on the opposite side of the branch complement; nodes with a girdle; 

secondary branches crowded at the base and sides of the main branch; blade of culm leaf 

petiolate, reflexed; inflorescence loose, open, racemiform or paniculiform; spikelets many- 

.............................................. Colanthel~a 

a girdle; branch complement with 

many secondary branches verticillate at the base of the central, dominant axis; inflorescence 

otherwise ................ ....................................... 16. Blade of culm leaf sessile, erect, without a petiole, the base as wide as the upper edge of the 

sheath; oral setae short; plants climbing or hanging from trees, usually with thin culms 

and a very small lumen, sometimes hollow but the walls then thick; inflorescence restricted, 

of limited branching, spicate-racemose; spikelets sessile, many-flowered; transitional glumes 

2 or 3; stamens 3; stigmas 2 

Blade of culm leaf petiolate, strongly reflexed, several times narrower than the top of the 

sheath; oral setae exceptionally long and stiff plants cespitose, the culms erect below and 

arching over vegetation, the larger ones with very thin walls; inflorescence unrestricted, of 

unlimited branching; pseudospikelets 1-flowered; transitional glumes 0; stamens 6; stigmas 

2 ..................................................................... 

1 fertile floret and 4 transitional glumes .................... .......... Chusquea 

to Bahia, eastern Brazil) .......... ....................................................................... ~~hroostachys 

much larger and dominant over the secondary ones .................................................................... 15 

flowered (southern and eastern Brazil) 

Nodal region without a prominent cres 

............................................. .Arthrostylidium 

TO BOTANY 

Conspectus of the Grass Subfamily Bambusoideae 

in the Western Hemisphere 

SUBFAMILY BAMBUSOIDEAE NES, 1835 

Tribe ANOMOCHI.OEAE: Anomochloa 

Tribe ARTHROSTYLIDIEAE: Arthrostylidium, Rhipidocladum 

Tribe ARUNDINARIEAE: Arundinaria 

Tribe BAMBUSEAE: Bambusa subg. Guadua 

Tribe CHUSQUEEAE: Chusquea, Newolepis, Swallenochloa 

Tribe OLYREAE: Cryptochloa, Diandrolyra, Ekmanochloa, 

Froesiochloa, Lithachne, Maclurolyra, Mniochloa, Olyra, 

Piresia, Raddia, Raddiella, Rehia, Reitzia, Strephium 

Tribe PARIANEAE: Eremitis, Pariana 

Tribe PHAREAE: Pharus 

Tribe STREPTOCHAETEAE: Streptochaeta 

Tribe STREPTOCYNEAE: Streptogpa 

GENERA NOT YET ASSIGNED TO TRIBES 

A poclada 

A throostachys 

Atractantha 

Aulonemia 

Colanthelia 

Elytrostachys 

Glaziophy ton 

Merostaehys 

Myriocladus 

Otatea 

Tribes of American Herbaceous Bamboos 

TRIBE ANOMOCHLOEAE C. E. HUBBARD (in 

Hutchinson, 1934:219).-This is a monotypic tribe 

based on the genus Anomochloa, which was described in 185 1 by Alphonse Brongniart, the director of the Museum d’Histoire Naturelle in Paris, 
NUMBER 44 17 

on the basis of plants cultivated in the Jardin des 

Plantes. The plants had been grown from seeds 

sent to France in the early 1840s by M. Pinel, a 

Frenchman in Brazil. The bizarre grass, which 

looks more like a member of the Maranthaceae than 

Poaceae (hence its specific name, A. marantoidea), 

was cultivated in the great European conservatories 

during the 1850s. By the early part of the following 

decade it had disappeared from cultivation and 

remained unknown from the wild until 1976, when 

the senior author discovered a colony of plants 

growing in an undisturbed, densely shaded forest 

in Bahia, Brazil. 

TRIBE OLYREAE KUNTH (1815:75).-The Olyreae is one of the oldest recognized tribes of grasses 

and is the largest of the herbaceous bambusoids, 

with 14 described genera. The tribe is endemic to 

tropical America, except for one species, Olyra 

latifolia L., that occurs also in tropical Africa and 

Madagascar, where it is apparently introduced. 

Olyra, with about 22 species, is widespread and extends from Mexico and the West Indies to northern 

Argentina. Some genera are quite restricted, such 

as Ekmanochloa and Mniochloa, found only in 

Cuba, and Maclurolyra, known only from Panama. 

While most species grow in shaded forests, their 

habitats are diverse. Species of Raddia, a genus of 

eastern Brazil, often occur in scrub forests associated with bromeliads and cacti and experience 

long periods of dryness. On the other hand, Froesiochloa inhabits the continuously wet rain forests 

of French Guiana and the neighboring states of 

Amapa and Para, Brazil..l. 

TRIBE PARIANEAE (HACKEL) HUBBARD (inHutchinson, 1934:219-220).-Hackel (1887:88) had 

recognized the distinctiveness of Pariana and established a subtribe Parianeae for it, but he included this in a distantly related group, the 

Hordeae. 

This is an American tribe of two described 

genera, Pariana and Eremitis. The larger of the 

two is Pariana with about 30 species in tropical 

lowland rain forest, primarily of the Amazonian 

basin, but extending north to Costa Rica and 

south to Peru, Bolivia, and Ecuador, the distribution following the natural outline of the basin. 

Pariana, like Hevea, is a rather good indicator 

plant of the Amazonian hylaea. The other genus, 

Eremitis, has been known from the single described 

species, E. paivifEora, from Bahia. In recent years 

we have found a number of further species of this 

genus, all from rain forests in Bahia and northern 

Espirito Santo. 

TRIBE PHAREAE STAPF (in Thiselton-Dyer, 

1898:319).-Whnn Stapf established the tribe 

Phareae in Flora Capensis, he included in it only 

Olyra, since that genus occurs in South Africa and 

Pharus does not. ‘That he considered Pharus to be 

a member of the tribe, although obvious from the 

name, is confirmed by Brandis (1907:87), who states 

that Stapf “has kindly informed me that three 

other genera [besides Olyra] . , . in his opinion belong to the same tribe.” From the list following 

that statement in Brandis, the three genera are 

Diandrolyra, Leptaspis, and Pharus. 

The tribe contains two genera. Pharus embraces 

about eight species that inhabit shaded tropical 

forests of the New World from central Mexico (San 

Luis Potosi) and the West Indies to Argentina (as 

far south as Corrientes and Tucuman). The other 

genus, Leptaspis, was originally described by 

Robert Brown on the basis of a species from Australia. There are about six species in the genus, one 

occurring in tropical Africa and the others in 

southern India and Ceylon, Southeast Asia, Indonesia, Australia, and the Philippines. 

TRIBE STREPTOCHAETEAE NEES (in Lindley, 

1836:378).--Nees (1835) regarded Streptochaeta as 

forming one of the three divisions of his bambusoid 

group. In his monograph he did not give a name 

to this division, which included only the genus 

Streptochaeta. The formal name Streptochaeteae 

appeared for the first time in Lindley (1836:378) in 

a list of grass genera prepared by Nees and dated 

12 December 1835. 

This is a monotypic American tribe consisting of 

Streptochaeta, the first herbaceous genus to be associated with the bamboos by Nees (1835) and the 

subject of various morphological studies (Celakovsky, 1889; Schuster, 1909; Arber, 1929). Of the two 

described species one, S. spicata, is widespread, 

ranging from southern Mexico through Central 

America, Trinidad, and in South America as far 

south as Misiones, Argentina. 

The more restricted species is S. sodiroana, which 

inhabits forests from Belize (British Honduras) 

through Costa Rica and Panama, into the lowlands of Ecuador. A third, undescribed species has 

been found in a single locality in Espirito Santo, 

eastern Brazil. 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

All species occur in shaded tropical forests, 

usually on hilly terrain. 

TRIBE STREPTOGYNEAE C. E. HUBBARD 

EX CALDOR~N & SODERSTROM 

Gramina perennia; foliorum ligula externa 

evoluta; racemi erecti, elongati, lineares, rigidi, 

secundi; spiculae subteretes, multiflorae; lemmata 

fertilia convoluta et maturitate teretia, indurata; 

lodiculae 3, lanceolato-acutae; stamina 2; stylus 

singularis, longissimus; stigmata 2 vel 3, longissima, 

persistentia, demum spiraliter torta et firma; caryopsis linearis; hilum angustissime lineare, caryopsi 

aequilongum; embryo ca. 1 / 10 partem caryopseos 

aequans. 

Pilger (1954:311) placed Streptogyna in a new 

subtribe of the Festuceae but did not give a Latin 

description. Hubbard (1956: 1) elevated the subtribe 

to tribe Streptogyneae but did not add a Latin 

description, which we are herein doing in order to 

validate the name. 

Streptogynn is a monotypic genus with two species, 

one (S. americana C. E. Hubbard) occurring in the 

New World and the other (S. crinita P. de Beauv.) 

in the Old World. The American species inhabits 

primary forests, where it forms rare patches and 

ranges from southern Mexico (Chiapas) through 

Central America to Trinidad, northern South 

America, and eastern Brazil (Bahia to northern 

Espirito Santo). The Old World species is found in 

forests in tropical West Africa, from Congo to 

Sudan, and again in southern India and Ceylon. 

Tribes of American Bamboos 

TRIBE ARTHROSTYLIDIEAE (MUNRO) E.-G. 

Camus (1913:16). - Munro (1868:13) first used the 

name Arthrostylidiae for a subsection of the section Triglossae and included Arthrostylidium and 

a couple of other genera. E.-G. Camus, in 1913, 

elevated this subsection to the rank of tribe. 

The tribe includes at least Arthrostylidium and 

Rhipidocladum, a genus recently segregated from 

it by McClure (1973). 

TRIBE ARUNDINARIEAE NEES (1835:466),- 

Nees von Esenbeck established the Arundinariae as 

one of his three major groups of bamboos and included in it Arundinaria and three other genera. 

In our present limited knowledge of the tribe, 

the only genus of the Western Hemisphere is 

Arundinaria, restricting the tribe therefore to 

North America. 

TRIBE BAMBUSEAE KUNTH (1815:75).-This 

was one of the original tribes of Kunth, in which 

he included all of the bamboo genera known to 

him. 

In the New World the tribe includes only the 

species that belong to the subgenus Guadua of 

Bum bum. 

TRIBE CHUSQUEEAE (MUNRO) E.-G. Camus 

(1913:16). - Munro (1868:13) named Chusqueeae as 

one of his subsections of the section Triglossae and 

included in it the genera Chusquea and Neurolepis 

(as Platonia). E.-G. Camus elevated this subsection 

to the rank of tribe in 1913. 

As presently understood, the tribe is endemic to 

the New World and contains the genera Chusquea, 

Neurolepis, and Swallenochloa. It is the most 

widely distributed tribe of Bambusoideae in the 

New World. 

Genera of American Bambusoideae 

Anomochloa Brongniart, 185 1 : 368-370 

TYPE-SPECIES: A. marantoidea Brongniart, 1851: 

TYPE-LOCALITY: Bahia, Brazil. 

COMMENTS:ame as those on tribe. 

TYPE-SPECIES: A. simplex McClure 8c L. B. Smith, 

TYPE-LOCALITY: Santa Catarina (Campos Novos), 

Brazil. 

COMMENTS: Four described species from southern 

Brazil, two of which are known from the cerrados of Mato Grosso, Goias, and Minas 

Gerais, and two from mountainous regions of 

Santa Catarina. 

368-372, 1 plate. 

Apoclada McClure, in McClure & Smith, 1967:57 

1967: 59-60. 

Arthrostylidium Ruprecht, 1839:27 

TYPE-SPECIES: A. cubense Ruprecht, 1839:28. Of 

the species enumerated by Ruprecht in his 

original description of the genus, Hitchcock 

(1927b3307) designated the above as the type 

of the genus. 

TYPE-LOCALITY: Province of La Habana (Ramon 

de la Sagra), Cuba. 

COMMENTS: A widespread genus of about 20 

species of clambering or climbing bamboos 

that range from the Caribbean Islands and 

Mexico to Venezuela and northern Brazil. 

Arundinaria Michaux, 1803, I:73 
NUMBER 44 19 

TYPE-SPECIES: A. macrosperma Michaux = A. 

gigantea (Walter) Muhlenberg, 1813: 14, based 

on Arundo gigantea Walter, 1788:81. 

TYPE-LOCALITY: Eastern United States (probably 

South Carolina). 

COMMENTS: A large genus of bamboos of temperate and cold regions of the Northern Hemisphere. Only one species, A. gigantea (Walter) 

Muhlenberg, is represented in the New World, 

where it grows in marshes and wet sandy 

banks of streams and rivers in eastern and 

southern United States. 

Athroostachys Bentham, in Bentham & Hooker, 

1883: 1208 

TYPE-SPECIES: A. capitata (Hooker) Bentham, 

based on Merostachys? capitata Hooker, 1840, 

plates 273-274. 

TYPE-LOCALITY: Rio de Janeiro, Brazil. 

COMMENTS: A monotypic genus, the single 

clambering species occurring in forests of the 

Serra do Mar of eastern Brazil, from Bahia 

south to Rio de Janeiro. 

Atractantha McCIure, 1973:42-48 

TYPE-SPECIES: A. radiata McClure, 1973:50-53. 

TYPE-LOCALITY: Bahia, Brazil. 

COMMENTS: A genus of 9 or 10 species of climbing bamboos of humid tropical forests and lowtree coastal forests on sandy soils of Bahia in 

eastern Brazil, one species extending into 

northern Espirito Santo. 

Aulonemia Goudot, 1846:75 

TYPE-SPECIES: A. queko Goudot, 1846:76. 

TYPE-LOCALITY: Quindio, Colombia. 

COMMENTS: A genus of over 20 species, many occurring in high-alti tude forests from Mexico 

and Central America through the Andes to 

Bolivia and Peru, eastward to the Guianas, and 

southward to the Serra do Mar of eastern 

Brazil. 

Bambusa Schreber, 1789:236, nom. cons. subgenus 

Guadua (Kunth) Nees, 1835:465, based on 

Guadua Kunth, 1822a:150-151 

TYPE-SPECIES OF THE SUBGENUS: Guadua angustifolia Kunth, 1822b:253, based on Bambusa 

guadua Humboldt & Bonpland, 1808:63, pl. 20 

(for an explanation of this lectotype and 

synonymy, see McClure, 1957:203). 

TYPE-LOCALITY: Probably Quindio, Colombia. 

COMMENTS: This is a large genus with representatives in the tropics of Asia and America. All 

of the New World species, of which there are 

about 30, belong to the subgenus Guadua. 

These are erect and usually robust, clumpforming bamboos that grow along rivers, 

mixed with tall trees in dense, humid forests, 

or occasionally in drier areas, where the plants 

occupy locally wet and more protected sites. 

TYPE-SPECIES: C. scandens Kunth, 1822b3254, 

based on Nastus chusque H.B.K., 1816:201. 

TYPE-LOCALITY: Bogota, Colombia. 

COMMENTS: The largest genus of American bamboos, with over 90 species that occur in every 

country from Mexico to Chile and Argentina 

and on many of the islands of the Caribbean. 

Some species reach the southernmost latitude 

of any Bambusoideae in the continent in the 

Andes, between Argentina and Chile at ca. 

47" S latitude. The majority are typical of cool 

forests at high elevations, but a few occur in 

forests on slopes at low elevations of eastern 

Brazil. 

Colanthelia McClure & E. W. Smith, in McClure, 

TYPE-SPECIES: C. cingu2ata (McClure & L. B. 

Smith) McClure, based on Aulonemia cinguZata McClure & L. B. Smith, 1967:50. 

TYPE-LOCALITY: Santa Catarina (Sombrio, Garapuvu), Brazil. 

COMMENTS: Seven species described from forests 

at low to medium altitudes of southeastern 

Brazil northwest to Minas Gerais. 

Cryptochloa Swallen, in Woodson & Schery, 1942: 

TYPE-SPECIES: C. variana Swallen, in Woodson & 

TYPE-LOCALITY: Cocle (Valle de Anoh), Panama. 

COMMENTS: About 15 species of shaded, mesophytic forests, extending from southern Mexico 

to the Pacific lowlands of northern South 

America, through Amazonia, and to eastern 

Brazil. 

Chusquea Kunth, 1822a: 151 

1973~77-79 

3 17-3 18 

Schery, 1942: 3 18-320, 1 figure. 

Diandrolyra Stapf, 1906:204 

TYPE-SPECIES: D.bicolor Stapf, 1906:204-205. 

TYPE-LOCALITY: "Native country unknown" 

(raised at Kew from fruits communicated by 

Messrs. Sander & Son). 

COMMENTS: About 6 species of humid rain 

forests of southeastern Brazil, from Bahia to 

Sao Paulo. 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Ekmanochloa Hitchcock, 1936:374-375 

TYPE-SPECIES: E. subaphylla Hitchcock, 1936: 

TYPE-LOCALITY: Province of Oriente (Sierra de 

Nipe), Cuba. 

COMMENTS: Two species known only from the 

province of Oriente, Cuba, on limestone hills 

and pinelands. 

375-377, fig. 343. 

Elytrostachys McClure, 1942: 173 

TYPE-SPECIES: E. typica McClure, 1942: 174-175, 

TYPE-LOCALITY: Distrito Federal, Venezuela. 

COMMENTS: Two species of wet lowland and 

midaltitude forests, recorded from Honduras 

to Venezuela. 

fig. 4. 

Eremitis Doell, in C.F.P. von Martius, 1877:338 

TYPE-SPECIES: E. parviflora (Trinius) Calderon & 

Soderstrom, based on Pariana paruiflora 

Trinius, 1834: 17. Doell (in Martius 1877:338- 

339, pl. 48) described the species as Eremitis 

monothalamia, but this name is illegitimate 

since Trinius’ earlier diagnosis constitutes 

valid publication. Presumably Trinius had 

examined the same specimen-Riedel 1591- 

in the Leningrad herbarium as had Doell, although he does not cite it as Doell did. 

TYPE-LOCALITY: Bahia (Castelnovo [Castelo 

Novo]), Brazil. 

COMMENTS: About 7 species known only from 

Bahia and northern Espirito Santo in eastern 

Brazil, inhabiting forests and forest margins, 

sometimes in disturbed places. 

Froesiochloa G. A. Black, 1950:29-30 

TYPE-SPECIES: F. boutelouoides G. A. Black, 

1950:30-31, pl. (estampa) 1. 

TYPE-LOCALITY: Territory of Amapa (Igarape 

Nataia, a tributary of the rio Oiapoque), 

Brazil. 

COMMENTS: Two or 3 species from wet, densely 

shaded primary forests of French Guiana and 

northern Para and Amapa in Brazil. 

Glariophyton Franchet, I889:277 

TYPE-SPECIES: G. mirabile Franchet, 1889:277-278, 

TYPE-LOCALITY: Vicinity of Rio de Janeiro, 

Brazil. 

COMMENTS: A monotvpic genus known only from 

the mountaintops of the Serra dos Orgaos near 

Rio de Janeiro. 

fig. A. 

Lithachne Palisot de Beauvois, 1812: 135 

TYPE-SPECIES: L. pauciflora (Swartz) Palisot de 

Beauvois, 1812:135, pl. 24: fig. 11, based on 

Olyra pauciflora Swartz, 1788:21. 

TYPE-LOCALITY: Jamaica. 

COMMENTS: Four species of which one is widespread, ranging from Mexico (San Luis Potosi) 

and the West Indies to northern Argentina. Of 

the others, one is endemic to Cuba, one to 

Honduras, and one to eastern Brazil near the 

boundary of the states of Rio de Janeiro and 

Minas Gerais. 

Maclurolyra Calderon & Soderstrom, 1973:6-12 

TYPE-SPECIES: M. tecta Calderon & Soderstrom, 

TYPE-LOCALITY: Province of Colon (Santa Rita), 

COMMENTS: One species known only from rain 

1973~6-12, figs. 4-7. 

Panama. 

forests of Panama. 

Merostachys Sprengel, 1825: 132 

TYPE-SPECIES: M. speciosa Sprengel, 1825: 132. 

TYPE-LOCALITY: None given for type, but Brazil 

according to Kunth, 1829: 139. 

COMMENTS: A genus of 40 or more forest species, 

especially common in Bahia, Brazil, but ranging as far north as Guatemala and as far south 

as northeastern Argentina. 

Mniochloa Chase, 1908: 185-186 

TYPE-SPECIES: M. pulchella (Grisebach) Chase, 

1908: 186, pl. 4, based on Digitaria pulchella 

Grisebach, 1866:231. 

TYPE-LOCALITY: El Yunque de Baracoa, Cuba. 

COMMENTS: Two species endemic to Cuba, one 

on shady limestone rocks in the eastern part 

of the country, and the other on shady banks 

and ravines in the western part. 

Myriocladus Swallen, in Steyermark, 1951 : 34 

TYPE-SPECIES: M. virgatus Swallen, 1951:34-35, 

TYPE-LOCALITY: Cerro Duida (Territorio Federal 

Amazonas), Venezuela. 

COMMENTS: About 20 described species endemic 

to the sandstone tabletop mountains of Venezuela in mesophytic habitats at elevations of 

1000 to 2500 meters. 

fig. 4. 

Neurolepis Meisner, 1843, vol. 1:426, vol. 2:325 

TYPE-SPECIES: N. elata (Kunth) Pilger, in Engler 

TYPE-LOCALITY: Loja, Quito, Ecuador. 

COMMENTS: About 10 species in the Andes of 

northern South America at high elevations 

& Prantl, 1906:21. 
NUMBER 44 

(2900-4500 m), from Venezuela and Colombia 

to Peru, some species occurring in the paramos. 

Olyra L., 1759: 1261 

TYPE-SPECIES: 0. latifolia L., 1759: 1261. 

TYPE-LOCALITY: Jamaica. 

COMMENTS: A genus of about 22 species widespread throughout tropical and subtropical 

America, from Mexico and the West Indies to 

northern Argentina, in shaded forests. One 

species, 0. latzfolza L., which is especially weedy 

and polymorphic and inhabits forest margins 

and disturbed sites, has spread from the New 

World to Africa and Madagascar. 

Otatea (McClure & Smith) Calderon & Soderstrom, 

based on Yushania K. H. Keng, 1957:357, subgenus Otatea McClure & E. W. Smith, in McClure, 1973: 116 

TYPE-SPECIES: 0tatea aztecorum (McClure & 

Smith) Calderon & Soderstrom, based on 

Yushania aztecorum McClure & E. W. Smith, 

in McClure, 1973:116-119, fig. 47. 

TYPE-LOCALITY: Rosario (state of Sinaloa), 

Mexico. 

COMMENTS: Two described species of erect bamboos that inhabit seasonally dry chaparral, 

edges of canyon walls, and limestone cliffs in 

central and southern Mexico, locally to Nicaragua and El Salvador. McClure originally 

established Otatea as a distinct genus and prepared the text and illustrations of it for inclusion in his revision of the New World genera 

of bamboos. According to his dated and unpublished manuscripts and notes, which we 

have seen, it was less than two months before 

his death (15 April 1970) that he decided, with 

some hesitation, to make Otatea a subgenus 

of the Old World bamboo, Yushania. With 

new perspectives on generic concepts of the 

Bambusoideae of the Western Hemisphere, we 

feel that McClure’s original concept of Otatea 

as a genus was correct and are so recognizing it 

in this treatment. In addition to the typespecies, a second was listed by McClure, based 

on Arundinaria acuminata Munro (1868:25), 

which now becomes Otatea acuminata (Munro) 

CaIderon & Soderstrom. 

Pariana FusPe-Aublet, 1775, vol. 2:876-877 

[Regarding the name Fusee-Aublet, see Bernardi, 

TYPE-SPECIES: P. campestris Fusee-Aublet, 1775, 

1976.1 

VO~. 2:877-879, VO~. 4, pl. 337. 

TYPE-LOCALITY: Loyola, French Guiana. 

COMMENTS: About 30 species of the rain forests of 

northern South America, especially abundant 

in the Amazon basin, whose southern and eastern boundaries mark the limits of the genus. 

Only a couple of species reach Central America, 

where they are of rare occurrence in Panama 

and Costa Rica. 

Pharus P. Browne, 1756:344, pl. 38: fig. 3 

TYPE-SPECIES: P. latifolius L., 1759:1269. [In the 

second edition (1789) of P. Browne, the plates 

are re-engraved and reversed and carry the 

Linnean binomials, according to Stafleu & 

Cowan (1976: 372).] 

TYPE-LOCALITY: Jamaica. 

COMMENTS: A genus of about 8 species of densely 

shaded tropical and subtropical forests, from 

southern Mexico and the West Indies to northern Argentina. 

Piresia Swallen, 1964: 152 

TYPE-SPECIES: P. goeldii Swallen, 1964: 153. 

TYPE-LOCALITY: Para (Maguary-assu, Braganca 

Railroad), Brazil. 

COMMENTS: Two described species from the rain 

forest of the Amazonian basin, reaching north 

to Surinam, Venezuela, and Trinidad and west 

to Peru. Several undescribed species, endemic 

to rain forests of Bahia and Pernambuco in 

eastern Brazil, exhibit a disjunct distribution 

for the genus. 

Raddia A. Bertoloni, 1819:410-411 

TYPE-SPECIES: R. brasiliensis A. Bertoloni, 1819: 

41 1. 

TYPE-LOCALITY: Province of Rio de Janeiro, 

Brazil. 

COMMENTS: About 9 species found only in eastern 

Brazil, from Pernambuco to Rio de Janeiro, in 

wet and seasonally dry forests. 

Raddiella Swallen, in Maguire et al., 1948:89 

TYPE-SPECIES: R. esenbeckii (Steudel) Calderon & 

Soderstrom, based on Panicum esenbeckii 

Steudel, 1854:90. [The type-species, cited by 

Swallen as R. nana (Doell) Swallen, nom. illeg., 

was based on Olyra nana Doell (in Martius, 

1877:329), based in turn on the same specimen 

as P. esenbeckii, which provides the earlier 

and, therefore, valid epithet.1 

TYPE-LOCALITY: Province of Rio Negro (Ega), 

Brazil. 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

COMMENTS: Eight to 10 described species that 

occur in wet depressions of savannas in the protection of vegetation, on wet rocky slopes, and 

on cliffs near waterfalls, from Trinidad and 

Panama to Mato Grosso, Brazil. The plants are 

among the smallest and most delicate of the 

subfamily. So diminutive are some of the 

species that grow among mosses on wet cliff 

faces that they may easily be confused with 

other grasses of dwarf stature, as was the case 

with R. molliculma (Swallen) Calderon & 

Soderstrom, originally described as Panicum 

molliculmum Swallen, in R. E. Schultes, 1953: 

57-58. 

Rehia Fijten, 1975:416 

[As pointed out by Fijten (1975:416), Bulbulus 

is not admissible under Article 20 of the ZCBN, 

as it is a technical term, nor under Article 64, 

as it is an orthographic variant and thus later 

homonym of Bulbilis Rafinesque, 1819.1 

TYPE-SPECIES: R. nervata (Swallen) Fij ten, based 

on Bulbulus neruatus Swallen, 1964: 154. 

TYPE-LOCALITY: Maranhao (Caxias to Barra do 

Corda), Brazil. 

COMMENTS: A monotypic genus known only 

from primary rain forests of the eastern Amazon basin and the Guianas. 

Reitzia Swallen, 1956:7-8 

TYPE-SPECIES: R. smithii Swallen, 1956: 8-9, I 

plate. 

TYPE-LOCALITY: Santa Catarina (Azambuja), 

Brazil. 

COMMENTS: One species from subtropical forests 

of Santa Catarina and Sao Paulo in southern 

Brazil. 

Rhifiidocladum McClure, 1973:lOl-104 

TYPE-SPECIES: R. harmonicum (Parodi) McClure, 

1973: 104, fig. 42, based on Arthrostylidium 

harmonicum Parodi, 1944:478, fig. 1. 

TYPE-LOCALITY: Alturas de Pintobamba, Peru. 

COMMENTS: About 10 species in forests at low to 

moderate elevations, from Mexico to Brazil and 

northwestern Argentina. 

Strephium Schrader ex C.G.D. Nees von Esenbeck, 

TYPE-SPECIES: S. distichophyllum Schrader ex 

TYPE-LOCALITY: Bahia (Felisberto), Brazil. 

COMMENTS: Four or 5 species of small, delicate, 

fernlike plants that occupy extremely wet spots 

along streams in dark forests, exhibiting a disjunct distribution between the rain forests of 

Bahia in eastern Brazil and the Guianas (including Amapa and northernPara, Brazil,il), 

Trinidad, and Venezuela. 

Streptochaeta Schrader ex C.G.D. Nees von Esenbeck, 1829:536-537 

TYPE-SPECIES: S. spicata Schrader ex C.G.D. Nees 

von Esenbeck, 1829:537-538. 

TYPE-LOCALITY: Bahia (Felisberto), Brazil. 

COMMENTS:ame as those on tribe. 

Streptogyna Palisot de Beauvois, 1812:80 

TYPE-SPECIES: S. crinita Palisot de Beauvois, 

1812:80, pl. 16: fig. 8 (Atlas). 

TYPE-LOCALITY: Old World. The specimen that 

the author described has 2 stigmas. He mentioned other specimens he had seen from the 

New World that have 3 stigmas and, while he 

did not describe these as distinct, he mentioned 

a S. guyanensis in the index but not in the text. 

Since the Old World species has 2 stigmas and 

the New World species has 3 stigmas, the name 

Streptogyna applies to the Old World species. 

1829:298 

C.G.D. Nees von Esenbeck, 1829:298-299. 

COMMENTS: Same as those on tribe. 

Swallenochloa McClure, 1973: 106-1 12 

TYPE-SPECIES: S. subtessellata (Hitchcock) McClure, 1973:108, fig. 44, based on Chusquea 

subtessellata Hitchcock, 1927a: 81. 

TYPE-LOCALITY: Cerro de la Muerte, Costa Rica. 

COMMENTS: The genus contains about 9 species, 

which are mostly small, clump-forming bamboos with stiff, erect culms, characteristic of the 

vegetation above treeline in the biome known 

as paramo. One species occurs in Costa Rica 

and the others in Venezuela and Colombia to 

Peru. 
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