THE.

,BAMBOOS
.
i
~

.
'.

A FRESH' PERSPECTIVE

py F. A. McClure
."' '

{.,'

..:;.;'"" . ~ .. ~ .
~:.~
.~!

'~\
.".
"~.
"; I'

,
..

I
I
,
 Tree-grass-unique in the plant kingdom

Symbol if uprightness, chivalry and devotion

THE BAMBOOS
Inspiration if poets, artists and philosophers A FRESH PERSPECTIVE
Writing material oJ the ancients;

the stylus of contemporaries

HONORARY RESEARCH ASSOCIATE IN
BOTANY, SMITHSONIAN INSTITUTION
Food, shelter and elothing if the people
Industrial substance qf a thousand uses

Redeemer if waste places-protector if the soil

Harvard University Press . Cambridge, Massachusetts . 1966
 t
1

© Copyright 1966 by the President

and Fellows of Harvard College
All rights reserved To Ruth Drury McClure
Distributed in Great Britain
by Oxford University Press, Landon
Library of Congress Catalog eard Number 66-10126
Printed in the United States of America
Composcd by Graphie Services, Inc. in
Baskerville on the Intertype Fototronic
Printed by the Murray Printing Co.
Bound by Colonial Press, !ne.
Book design by David Fard
Tbls book was prepared under the auspices
afthe Maria Moors Cabot Foundation.
Preface

The presentation of this treatise in book form is the outcome of

my response t6 an opportunity offered by the Maria Moors Cabot
Foundation of Harvard University, The Foundation's Adminis-
trative Committee desired a summary of authentie information
on selected aspects of the bamboos, to place this group of plants
in perspective in relation to the planning of its research activities.
I am deeply indebted to Dr. Paul C. Mangelsdorf, Chairman of
the Committee, for the invitation to initiate this undertaking, and
to his successor in this office, Dr. Kenneth V. Thimann, for
sustained and sustaining interest and support.
My involvement in scientific studies of the bamboos dates from
aperiod of service .(1924 to 1927) under the United States Depart-
ment of Agriculture, as Agricultural Explorer in China I was
instructed to secure living material of useful bamboos fqr trial in
the Western Hemisphere. With facilities made available by the
Lingnan University at Canton, China, budgetary aid from the
Rockefeller Foundation through its "China Foundation," and per-
sonal subsidy generously provided by the late Helen Lyon Jones,
it was possible eventually to assemble, principally from eight prov-
inces of southeastern China, over 600 accessions of living bamboo
plants for observation and experimental study. (Through the inter-
est of Dr. R. E. Holtturn, who recently visited Canton, it is now
known here that this collection is still intact, and that it remains
linder the care of the erstwhile colleague who collaborated in
establishing it.) The United States Department of Agriculture has
accessioned 250 numhers of living plants of species selected from
this collection duririg the 17 years from 1924 to 1940. In the pur-
suit of studies undertaken in that period I became greatly obligated
to my Chinese colleagues for assistance of a most distinguished and
effective quality,
viii PREFACE PREFACE ix

Throughout the years 1933 to 1936 and 1941 to date;the Depart- sense of gratitude for generous hospitality and for much practical
ment of Botany of the Smithsonian Institution has extended benef- help. I feel an obligation of a special nature to David Fairchild,
icent hospitality to my herbarium studies on bamboo. The John Wilson Popenoe, Robert Cunningham Miller, Conrad Chapman,
Simon Guggenheim Foundation supported my taxonomie work on Robert A Young, Ross E. Moore, and Malcolm G. Lyon, each
selected groups during the fiscal year 1942-43. During the fiscal of whom has shared and actively supported a concern for the con-
year 1943-44, under the auspices of the Smithsonian Institution, tinuity, progress, and fruition of my work. The 10yaJ encourage-
I made a survey of useful bamboos of a special category available ment of my efforts by Professor Adolph E. Waller, a favorite
in the United States, Mexico, Honduras, ,Colombillo Venezuela, lecturer in Botany at the Ohio State University, has exerted sus-
Brazil, and Puerto Rico for the Office of Scientific Research and taining influence of a rare quali ty.
Development of the U.S. National Research Council (McClure By her tireless and efficient service in bringing up to date the
1944). The post of Field Service Consultant on Bamboo with the assembling and processing of published papers on bamboo, and mak-
Office of Foreign Agricultural Service of the U.S. Department of ing many translations into English from a polyglot,array of litera-
Agriculture (1944 to 1954) gave me the opportunity to study and ture, besides carrying general secretarial responsibilities, Maude
collect bamboos in six countries of Central and South America, Kellerman Swingle lightened my burden and quickened the pace
as weil as in India, East Pakistan, and the islands of Java and of my labors during the fonnative stage of this book. Mamie Idella
Luzon. Ultimately I was able to establish living collections of elite, Herb gamered many references from the libraries of the Smith-
economic species in Guatemala, EI Salvador, Nicaragua, Costa sonian Institution and the U.S. Department of Agriculture, helped
Rica, Ecuador, and Peru. During this period it was my good for- process theliterature, and transcribed with patient industry and
tune to have the use of research facilities, and the collaboration of gratifying precision, the numerous earlier drafts through which
members of the staff, at the Federal &,:periment Station at Maya- the manuscript has passed. Ruth Drury McClure has given con-
güez, Puerto Rico, in carrying out exploratory studies designed to stant support, enhancing the joys and devotedly sharing the trials
improve methods of bamboo propagation. At the suggestion of of my pursuit of the bamboos through the years. Withc her help,
Donald D. Stevenson, the U.S. Department of Agriculture author- the manuscript was read aloud and critically considered during
ized me to serve, during this period, as its representative in the each revision and the final touches were incorporated in the
capacity of consultant on bamboo to Champion Papers Inc. This definitive draft.
relation afforded an opportunity to make field studies in Jamaica William C. Kennard, erstwhile collaborator in the processing
and Trinidad, to design and supervise the establishment of an of data on bamboo propagation, has read with a critical eye that
experimental bamboo plantation in Guatemala, and to participate part of the manuscript which deals with this subject. For a pains-
'in elaborate studies based on it. Since the tennination, in 1956, taking and cornpetent scrutiny of the whole original draft, and
of my latest period of service with the U.S. Department of Agri- for valuabie suggestions, I am grateful to R. E. Holtturn, fonnerly
culture, financial support made available by the UNESCO Com- Director of the Botariic Gardens at Singapore and author of a
mittee for Biological Studies in the Humid Tropics, the U.S. recently published taxonomie treatise on the bamboos of the Malay
National Science F6undation, and the Maria Moors Cabot Foun- Peninsula..Among other reliable sources of infonnation that have
dation of Harvard University, respectively, has sustained my enabled me to extend beyond my personal e"-"perience the range
bamboo studies. of subjects covered in this book are several distinguished studies
This account would not be complete without an acknowledgment published by Willard M. Porterfield.
of my indebtedness to innumerable persons of good will, in all Permission has been graciously extended by ~he Philosophical
walks of life, both here and in those parts of the world where my Library, New York; Alfred Knopf, lnc., New York; The Univer-
field studies have taken me. These encounters have left a lasting sity Press, Carnbridge; The Clarendon Press, Oxford; The Oxford
x PREFACE

University Press, London; Hiroshi Muroi; Yokendo, Ltd., Tokyo;

the Ministry of Education, Government of India, and Encyclo-
pedia Britannica,"Inc., to reproduce ~extual material or illustrations,
or both, from the copyrighted writings of Alfred North Whitehead,
Arthur Waley, Agnes Arber, C. R. Metcalfe, R. S. Troup, N. L
Vavilov, H. Muroi, Yoshio Takenouchi, P. N. Deogun, and myse1f,
Contents
respectively. These courtesies are acknowledged individually at
appropriate points in the book. The frontispiece, derived frorn
a rubbing of a stone carving based on a painting by Hui Nien
(known to connoisseurs of Oriental art for his skill in painting Foreword by Kenneth V. Thimann XlV
bamboos), was first used in this greatly reduced form by C. LeRoy
Baldridge in the cover design of his book, Turn 10 the East, pri- Introduction 1
vately published and copyright by him in 1925.
Excepting those credited to other sourees, the photographs and PART I.'THE BAMBOO PLANT
the original sketches for the line drawings are my own. With his 1 Vegetative Phase: The Maturing Plant 11
well-known ar'tistic and technical skill, EImer W. Smith has tidied Segmented ·axes. The neck The rhizome system. Two basic farms
my pencil sketches, rendered them in ink, and reconditioned of rhizome. Reinterpretation of the rhizome system. Pachymorph
illustrations adopted from published sourees. Several of the original rhizomes. Leptomorph rhizomes. Metamorph a..xes. Other cornbina-
drawings wem made by him specially for this book. tions of underground growth farms. Key to known farms of the
barnboo rhizome system. Clump habit. The culm. Branch buds at
As a last word, I wish to thank the staff of the Harvard Uni- cu1rn nodes. The ~ranch complement. The sheathing organs. Culm
versity Press, andJoseph D. EIder, the Science Editor, in particular, sheaths. Brauch sheaths. Leaf sheaths. Rhizome sheaths. Neck
for distinguished contributions to the degree of excellence that has sheaths. !,:ophylla. Roots.
been achieved in the production of this volume.
2 Reproductive Phase ' 82
F.A.M. Fruiting behavior in different bamboos. Flowering habits of bam-
Smithsonian Institution boos as related to taxonomy and the problem of field identifica-
Washington, D.C, tion. The infLorescence. The indeterminate infiorescence. The deter-
November 28, 1964. minate infiorescence. The course of development of the indete.rmi-
nate inflorescence. Variations and "anomalies" in the indeterminate
inflorescence. Diversity of form. manifested by the detenninate inflo-
~escence. The barhboo infiorescence from the point'ofview ofphys-
101ogy. The prophyllum. in inflorescence branching. Spikelet, floret,
and flower. The fmi t.

3 Vegetative Phase: The Seedling 122

The seedling in bamboos with pachyrnorph rhizomes. The seedling
in bamboos with leptomorph rhizomes. Recapitulation ofsignificant
events in the ontogeny of a bamboo ,plant.
XlI CONTENTS CONTENTS xiii

PART II. ELITE BAMBOOS AND PROPAGATION METHODS Appendix H. Bamboos'Offered and Nurseries
Offering Them in the Uni ted States 292
4 Selected Species 147
Arundinana amabilis McClure. Bambusa vulgan:s Schrad. e.x Wendland. Glossary 295
Dendrocalamus strictus (Roxb.) Nees. Gigantochloa verticillata (Willd.)
Munro. Guadua angustifolia Kunth. Meloca:nna baccifera (Roxb.) Kurz. Literature Cited
Ochlandra travancorica (Bedd.) Bentham ex Gamble. 319
Index of Scientific Names 339
5 Propagation 202
Propagation by means of seeds. Duration of viability in bamboo Index of Subjects 345
seeds. Planting the seeds. Requirernents relating to scientific and
commercial objectives. Small versus large plants. Two groups of
bamboos requiring different propagation procedures. SignificaDce of
the. differences between the two groups in relation to vegetative
propagation. Conclusions on past work, and future objectives. Facts
and factors affecting propagation procedures. Season of the year as
a "variable" in procedures. Dormancy and development of buds.
Rooting. Need of a new approach to the study of vegetative propa-
gation in bamboos.

PART IIL BASES OF CLASSIFICATION

6 Flowering and Fruiting Behavior in Bamboos

of Different Genera and Species 267
Anmdinaria amabilis McGlure. Arundinan:a aun'coma Mitford. Arundinaria
pumila Mitford. Arundina:ria pygmaea (Miq.) Aschers. & Graebner.
Arundinaria simonii (Garr.) A. & C. Riviere. Arundinaria simonii var.
variegata Hook. f. Arundinaria variegata (Sieb.) Makino. Bambusa cope-
la.ndii Gamble. 8yn.: Sinocalamus copelandii (Gamble) Raizada. Bambusa
lineata Munro. Bambusa multiplex (Lour.) Raeusch. Bambusa tuldoides
Munro. Chimonobambusa falcata (Nees) Nakai. Chimonobambusa hook-
eriana (Munro) Nakai. Chimonobambusa quadrangularis (Fenzi) Makino.
Chusquea abietifoHa Griseb. Dendrocala:mus hamiltonii Munro. Dendro-
calamus strictus (Roxb.) Nees. Guadua ~rJUStifolia Kunth. Guadua tn'nii
(Nees) Nees. Mclocanna baccifera (Roxb.) Kurz. Merostachys jistulosa
DoeH. Nastus elegantissimum (Hassk) Holttum. Ochlandra scriptoria
(Dennst.) C. E. G. Fischer. -Ochlandra stridula Thwaites. Ochlandra
travancorica (Bedd.) Benth. e.xGamble. Oxytenanthera abyssinica(A. Rich.)
Munro. Phyllostachys spp. Sasa tessellata (Munro) Makino et Shibata.
Thamnocalamus falconeri Munro. Thyrsostachys oliven' Gamble.

7 Bamboos from the Point of View of Taxonomy 280

Appendix 1. Generic Key to Bamboos under

Cultivation in the United States and Puerto Rico 289
 FOREWORD xv

selves and for their effeets on the soiL He also studied and eol-
lected in Kwangsi and in -the Hong Kong area. But in 1941 the
war drove him back to the United States, and he has made the
National Herbarium his headquarters ever since. From there he
has visited almost every country in Latin America to study and
Foreword collect bamboos, to direct programs of investigation, .distribution,
~nd propaga:ion of bamboos there, work on bamboo propagation
m Puerto R,CO, and establish field plantings in Guatemala and
five other Latin American countries. He also visited United States
Born on a farm in Ohio, Floyd Alonzo MeClure attended the missions in the Far East, and introduced selected bamboos into
Ohio State University at Columbus, with the intention of return- various Pacific islands for triaL As consultant on bamboos to a
ing to the farm. But upon his graduation, in 1919, an unexpeeted United States paper manufacturer McClure has made field studies
offer of an instruetorship in horticulture at Lingnan University in in Central Ameriea, Florida, and Te"",,s and has collaborated in
Canton, China, presented an irresistibleopportunity for an adven- research on the pulping properties of selected species. In this he
turous young man interested in seeing the world and its peoples has been helped by his earlier studies of bamboo papers in China.
and plants. He stayed on at Lingnan, was made eurator of the It was with a view to making MeClure's lifetime study and
herbarium, and in 1931 became professor of botany there. In the experience with .bamboos available to a wider circle that in 1955
meantime he had returned twice on furlough to· the Uni ted the Maria Moors Cabot Foundation, through Professor Paul Man-
States and had also led numerous field trips in the Chinese interior gelsdorf, invited him to try to set down at least apart of his
and in Indochina, in part as e"1'lorer for the United States Depart- aecumulated knowledge. Surely no one can claim more extensive
ment of Agriculture. The effeetiveness of his collecting was of acquaintance at first hand with every aspect of this fascinating
course greatly helped by the fact that from his first years at Ling- group of plants. We have been fortunate, too, in having the serv-
nan he had studied Cantonese and aequired a fiuent eommand of ices of EImer Smith, who has illustrated many botanical and zoo-
it. It was on these field trips that MeClure became interested in logical publieations, and has given expression both to his botanical
ban;lboos; he very early initiated a Bamboo Garden at .Lingnan knowledge and his artistic feeling in preparing the very numerous
and established there many of the plants he had collected. This illustrations. The book has been many years in preparation, and
constant preoceupation with living plants, rather than preserved it contains not only a mass of information but mueh valuable
material, is characteristic of his interests, and it is that whieh has insight. I have acted informally as editor and critic.
made the present book possible.
In 1932, McClure returned to the United States and Europe KENNETH V. THlMANN

for a four-year period. He studied critical specimens in the bamboo Chairman, Executz've Committee
Marz"a Moors Cabot Foundatz'on
collections at the United States National Herbarium in Washing-
ton, at the Royal Botanie Gardens, Kew, and at the British Mu-
seum; at Leiden he worked on other materiaL He obtained his
doctorate at Columbus in 1935, and returned to China in 1936. At
Lingnan he stimulated his colleagues in chemistry and engineering
to make joint investigations ofbamboo products, and he established
a large experimental planting where 10,000 plants of bamboo
could be studied both for the useful properties of the plants them-
 ,

My new Province is aland of bamboo groves;

Their shoots in spring fiU the valleys and hiUs.
The mountain woodman cuts an cmnfol of them
And brings them down to sell at the early markeL
Things are eheap zn proportion as they are eommon; THE BAMBOOS
For two forthings I bUJ a whole bwulle.
I put the shoots in a great earthen pot,
And steam them over the boiling riee.
Their purpl, nodules broken, they snggest an old brocade,.
Their white skin gleams like new pearls.
-po GRU-I, TRANSLATED BY WA1.EY (1929)
In trod uction

The bamboos :p.ave age-old connections with fishing, papermaking,

landscape gardening, handicrafts, medicine, art, and even poetry.
Having long been creatively exploi ted at the handicraft level, var-
ious kinds of bamboo minister in a comprehensive"manner to the
material needs of native peoples of both the old World and the
New. In some of the heavily populated regions of the tropics cer-
tain barnboos supply the one suitable material that is sufficiently
cheap and plentiful to meet the vast need for economical housing
(McClure 1953). Beyond this, barnboo culms provide the raw
material for hundreds of implements of daily necessity in the horne
and in the pursuit of a livelihood, as well as musical instruments
and countless sorts of toys for children. The young shoots of cer-
tain species are eaten as a vegetable; the living plants of others are
cultivated in decorative or protective hedges about homes and
villages. Oriental scholars, artists, and epicures have celebrated
the importance of bamboo and praised its admirable qualities in
paintings, rhyrned couplets, and lyrical tributes (Po Chu-i, trans-
lated by Waley 1929; Li 1942; McClure 1937, 1956b, 1961a).
The suitability of bamboo tools and bamboo fibers for mak-
ing paper was demonstrated centuries ago by Chinese artisans
(McClure 1928; Sung 1929). As a result of recent fiber-dimension
studies and improvements in mill techniques, fine papers of many
varieties and adaptations can now be made from the pulp of cer-
tain bamboos. High-grade barnboo pulps can be used in the pure
state for coated or uncoated book and magazine papers. The
pulp of certain bam boos exce1s in the field of soft facial tissues
and for thin, India-type papers where opacity is at a premium.
Fiber· dimensions with a high length-to-di-ameter ratio give many
bamboo pulps a special versatility.
As a result of modern technological
,
advances and applied
2 lNTRODUCTION lNTRODUCTION 3

research, new uses for bamboo products have been developed and for nearly two decades with very fruitful results (Shepherd 1952).
new significance found in old uses. Tabashir, precipitated within However, young shoots of many tropical speeies of bamboo con-
the culm internodes of many tropical bamboos, consists largely tain lethai concentrations and amounts of cyanogens (Valt'er et
of arnorphous silica in microscopically fine state. lt has been shown al. 191?, 1911). ~he di~estive processes of the herbivora destroy
to have, both in the purified form and in combination with the pOlson, but In Indla eattle sometimes die when they are
several elements and compounds, excellent properties as a catalyst allowed to eat too freely of the toxie young shoots. Boiling readily
for certain chemical reactions (Netherlands Patent No. 53,471). drives off the volatile cyanide, so there is no risk involved in eating
The most successful of Edison's early electric lamps bad for its eooked shoots.
light-giving element a carbonized filament of barnboo. (McClure Bamboos are set off from other grasses by the predominanee
1948). Bamboo fibers were still used in special-purpose carbon- of certain "bambusoid" structural characters, many of which are
filament lamps as recently as the year 1908 (Bryan 1926). japa- considered to qe ~'primitive.~' The most easily recognizable vegeta-
nese sciehtists have demonstrated that charcoal prepared from tive features that distinguish the bamboos are the prominent
certain bamboos has properties that render it superior to that development of a rhizome system, the woodiness and strong braneh-
derived from conventional ~ources for use in elec~ric batteries ing of the eulms, the presence of petioles on the leaf blades, and
(Miyake and Sugiura 1950, 1951), and in India bamboo charcoal the differenee in form between the sheaths clothing young eulm
is used for pharmaceutical purposes. Oriental jewelers use the shoots and those borne on the leafy twigs. To these may be added.
silica-charged charcoal from barnboos in preference to that from such floral characters as well-developed lodieules, in most species
other sources. From the white powder abundantly produced on three in number, and a style consisting typically of a single eolumn,
the outer surface of young culms of a Chinese bamboo, Chang bearing one, two, or three (rarely more) stigmas (see Figs. 48 and
(1938) isolated, among other substances, a crystalline compound 53). No single eharaeter is diagnostic, however, and the boundary
related to the female sex hormone. Piatti (1947) reported the suc- between the bamboos and the other grasses is not sharp (see Parodi
cessful preparation of liquid diesel fuel from barnboo culms by 1961 ).
distillation. In japan, Kato (1911) extracted from bamboo shoots Of very uneven geographieal distribution, bamboos appear
the enzymes nuclease and deaminase, besides an unnamed enzyme more or less prominently in the natural vegetation of many parts
that dissolves fibrin, and another of emulsin type, capable of of the tropical, subtropical, and mild-temperate regions of the
hydrolyzing salicin. Komatsu and Sasaoka (1927) isolated glucu- world, from sea level to the snow line. Bam boos are found in the
ronic acid and L-":ylose, in a crystalline state, from the juice of greatest abundanee and variety on the southern and southeastern
bamboo shoots. Yoshida and Ikejiri (1950) found an aqueous ex- borders of Asia, from India through China and japan to Korea.
traGt of bamboo shoots superior to conventional media for the In the vast Eurasian eontinent north, west, and northwest of Tibet
culture of certain pathogenic bacteria (Shigella and Brucella). This and China, however, no native bamboo has been found. The
extract, wheh added to tuberculin, increased the intensity of re- island of Madagasear is particularly rich in endemie genera and
sponses to the skin test. speeies, baving more known kinds than all of Afriea. Three en-
The use of dried mature bamboo leaves for deodorizing fish oils demic species have been described frorri Australia. In the Western
has been patented in japan Oapanese Patent No. 175,685). In Hemisphere, the known natural distribution of bamboos extends
both the Eastern and Western Hemispheres, the foliage of many from 39°25' N in eastern United States (personal observation) to
bamboos has long been used as a major or a supplementary source 45°23'30" S in Chile (Dusen, 1903-1906:18f!) and even to 47° S in
of forage (McClure 1958). The improvement of the management Argentina (Parodi 1945:64).
of grazing range in the coastal plain of North Carolina, where a The natural distribution of bamboo in the world has been
native bamboo is the principal source of browse for beef cattle, greatly modified by human intervention. For example, many nat-
is the object of a project in which three state agencies and three ural stands have been more or less completely destroyed in clearing
ageneies of the U.S. Department of Agrieulture have eollaborated land for agriculture. The canebrakes of the UniteCl States have
 4 INTRODUCTION INTRODUCTION 5

been greatly reduced in number and extent. Guadua aculeata; a bring them into fresh perspective in the light of personal experi-
giant species once abundant locally in several countries of Central ence and of published observations judged to be authentic and
America, has been almost completely eliminated in some areas. pertinent to the aspects touched on herein. The fragmentary na-
On the other hand, Bambusa vulgaris, a pantropic species of un- ture, of present knowledge about members of this generally
known origin, has been naturalized in large areas on the island neglected group of plants renders impossible, as yet, the synthesis
of J amaica in the wake of a sort of rriigratory agriculture 'in which of an adequate picture of either their essential character or their
stakes freshly cut from living culms of this bamboo, and stuck in relation to members of other tribes of the Gramineae,1 and to
the ground to support yam vines, take root and produce thriving extragramineous groups such as the Arecaceae, Juncaceae, Lilia-
groves. Sinobambusa too!sik, a Chinese bamboo, once highly prized ceae, Flagellariaceae, and Restionaceae (see Stebbins 1956). A
as a garden ornamental in Honolulu, has shown that it can be- thoroughly satisfactory taxonomic exploration of the relationships
come a troublesome weed there, for it escaped from cultivation of the bamboos, and the exploitation of their technological poten-
and now dominates many acres cf anee pure native vegetation. tialities, await basic studies by specialists in diverse disciplines.
In Mrica, Europe, the British Isles, and the United States, intro- The three principal parts of this account are designed to give
duced bamboos have found an important place as ornamentals the reader a concept of (1) what to look for in studying a bamboo
and as a source cf fishing poles, garden stakes, and fencing, as plant, (2) what is known about bamboo propagation (with sugges-
weil as material for handicrafts and interior trim. Many little- tions for improving conventional procedures), and (3) distinguish-
known exotic species of promise are still confined to introduction ing characters, and known technical properties, of several bamboos
. gardens, where they await basic studies and the prolonged critical that are outstanding in the importance they have been given as
research necessary to test their potential usefulness in the local economic plants. The appendices touch on additional aspects of
economy. the bamboos, and provide supplementary reference material.
Relatively little of a basic nature is known about any of the Graduate students and specialists in the several disciplines
bamboos. Among the disciplines that have neglected this group concerned with original research in the plant sciences will find
more or less completely are those dealing with anatomy, cytology, numerous invitations to undertake studies designed to illuminate
genetics, plant breeding, biochemistry, morphology, mo~hogene­ areas where recorded knowledge of the bamboos is deficient. An
sis, ecology, and the physiological phenomena related to vegeta- effective exploration of the physiological foundations of gross
tive growth, flowering, and fruiting. Even the commonly practiced morphology and anatomy should improve our understanding of
methods of propagation are still restricted to the traditional pro- the evolving diversity of form that is at once the major concern
cedures of the plantsman's art, and current applications of the and, often, the dispair of the taxonomist.
newer knowledge of hormones and the techniques of controlling
vegetative growth and sexual reproduction have not touched the Perspectives
problems involved in the practical management and exploitation Although the following description of the bamboo plant is pre-
of plants of this group. Furthermore, the taxonomic treatment of sented primarily from the point of view of gross morphology, the
the bamboos, which is basic to all other angles of study, is still in lTraditional perspectives upon the morphologieal, physiologieal, and genetic barriers
a very retarded state. An amplified and modernized description that supposedly prevail between major divisions of the grass family cannat fail to be
of this important tribe of the grasses is presented on pp. 232ff. modified by facts brought to light through an increclible feat accomplished by scientists
working independently in India and Formosa, respectivcly. I refer to the apparently
Revised descriptions of the tenable genera are being prepared, successful hybridization between a barnboo (subfamily Barnbusoideae) as "male" parent
with special emphasis on the redefinition of their respective bound- and a sugareane (subfamily Panicoideae) as "female" parent, first reported by Venkatra-
man (1937). Morphological and anatomical studies (Kutty Amma and Ekembaram 1940;
aries, and the systematic use of fundamental characters hi therto Loh cl al. 1950) and cytological studies (Janaki Aromal 1938) appear to have established
generally neglected. 'the hybrid nature of the resulting progenies. However, since the matter seems still to hover
in the rea1m of controversy, further attempts at such crosses and thorough studies of the
One objective of this treatise is to make astart at removing progeny should be carried out before final conclusions are formed wl,th respect to the sig-
the bamboos from the shadow of insufficient ~owledge, and to nificance _of the published evidence.
6 INTRODUCTION INTRODUCTION 7

physiological and biochemical setting of morphogenesis has been botanical entity could be integrated, with confidence in their
constantly in mind during its preparation. 2 common pertinence. As the publication and integration of such
The external form of the homologous parts of a bamboo plant data progressed, the recognition, delineation, and differentiation
varies with their respective positions on the several segmented of critical species could be given greater depth of focus, the clas-
axes, and with the increase in size of the whole plant. These clif- sification of thc bamboos could gradually be made more con-
ferences in outward form refiect underlying physiological processes sistent and more workable (functional), and the approach to their
that may operate through biochemical gradients as well as by economic e"1'loitation could be fruitfully broadened..
discrete steps (see Schaffner 1927; Porterfield 1933; Watson 1943; The hitherto generally neglected branching habit as expressed
Thompson 1944; Thimann 1954; Prat 1954; Wardlaw 1956, 1959; in the rhizome, the culm, and the inflorescence is herein given
Heslop-Harrison 1959; Zimmerman 1961). Possibly it is for this special attention. Also emphasized are the nature and role of the -
reason that some of the outward manifestations of gross morphol- prophyllum, which is very important to an understanding of
ogy in the bamboo plant are not easily amenable to the rigid branching habit, particularly in the indeterminate inflorescence .
. application of a hierarchy of technical terms or reasoned concepts. Takenouchi's excellent illustrations (1931a; 1932) of the forms
On the other hand, the fact that some structures do not yield assumed by the prophyllum in the vegetative axes of the common
readily to a regime of sharply defined categories is, perhaps in J apanese species are a pioneer contribution to the elucidation of
part at least, an indictment of our insufficient knowledge. Again, this category of bamboo structures.
the fault may lie partly in the point of view from which we make Published dissertations on the bamboos often leave basic con-
our observations, Of in the nature of the assumptions on which cepts, observations, and generalizations inadequately documented.
we base our interpretations. A special effort has been made here to avoid this fault by citing
The best picture of any given bamboo that can be pieced illustrative species and, according to the requirements of the case,
together by the taxonomist, on the basis of the morphology of a particular specimen or collection.
conventional herbarium specimens alone, is sadly inadequate. The In my effort to achieve an improved understanding and por-
increasing diversity and the improving integration of recorded trayal of the bamboo plant, I have sought fresh ways of looking at
botanical data have resulted in a swelling chorus of appeals for a farniliar things, and have challenged established concepts and
broader approach to the study of taxonomy as well as other as- usages that cast a shadow on certain areas where more light
pects of biology (see Vavilov 1940; Sprague 1944:235; Bush and is needed. 3 Critical attention directed at (1) the elimination
Lowell 1953; Metcalfe 1954:440;' Prat 1960:60 et passim; Kety of ambiguities and (2) the disciplined communication of my own
1960). For a third of a century, I have had a growing conviction observations has led to the adoption of some new terminology and
that a really successful conquest of the bamboos requires an inter- to the modification of some of the old. Terms in these two cate-
disciplinary approach. Ideally, this would involve the collaboration gories are highlighted by means of asterisks and daggers, respec-
of a group of specialists representing such points of view as anat- tively; in the Glossary.
omy, biochemistry, histology, physiology, morphology, genetics, Attention is directed especially to the taxonomie scope of the
ecology, taxonomy, and utilization, to name the most obviously subject matter of this work. It is essentially restricted to the plants
pertinent disciplines. Instead of working independently, as in the universally recognized as bam boos (the tribe Bambuseac), although
past, on study materials drawn from diverse sources-materials occasional reference is made to features of members of the related
all too often of uncertain identity-they should work together (at tribe, Streptochaeteae. Entirely omitted from the treatment are
least in conscious collaboration) on documented material from a the tribes Olyreae, Phareae, and Parianeae, recently allocated to
common source. In this way, all of the data recorded for a given the subfamily Bambusoideae by Parodi '(1961).
2Takhtajan (1959:129) e."<.Presses the opinion that "thc evolutionary botany cf the
future will be erected on the basis cf a synthesis cf morphology and physiology." Macleod 3Koestler (1964) touches obliquely upon the history and uses of this device in his
and Gabley (1961:viii) aver that "tbe cornerstones of botanical philosophy are still the chapter entitled "Tbe Evolution of Ideas." The taxonornist of refl~ctive temperament
triumvirate of rnorphology, ta:'{onomy and physiology." will discover in Koestler's work much discourse that is instructive as weil as diverting.
 Human knowledge is a process of approximation. In the flcus
0/ experience there is comparative clariry. But the discrimination 0/
this clarity leads into the penumbral background. There are always questions Part! The Bamboo Plant
left over. The problem is to discriminate what we know vaguely.
-ALFRED NORTH WHITEHEAD (1948 )

One must strioe Jor inJallibility without pretending to itl

-MALEBRANCHE
1. Vegetative Phase: The MaturingPlant

As most commonly encountered, a bamboo plant is somewhere

on its way through the usually long, purely vegetative phase of
development (or maturing) that intervenes between the seedling
stage (p. 122) and the reproductive phase (p. 82). The component
parts of the plant as they appear in the more familiar vegetative
phase will be described first. The different forms assumed by the
various kinds of structure as they appear in either of the two more
rarely observed stages will then be more easily understood. Four
major categories cf vegetative structures may be recog~1ized: seg-
mented axes, sheathing foliar organs, buds, and raots.

Segmented axes
The basic frame of the bamboo plant consists of a ramifying sys-
tem of segmented vegetative axes (Fig. 1). These axes may be
differentiated as_ rhizomes, culms, and culm branches. There is
no central trunk cr main axis. Each cornponent axis, whether
rhizome, culm, or branch, consists cf aseries of nodes and inter-
nodes. It is clothed (at least during the period of its active growth)
with enveloping sheaths that face alternate sides of the axis at the
successive nodes. With certain minor exceptions, each sheath
subtends a bud or a branch complement. Adventitious roots are
initiated at the nodes of segmented axes-in certain parts of the
plant to be detailed later (p. 79).
The term "segmented" refers to the regular alternation of nodes
and internodes that characterizes the axes of the rhizome system,
the culms, and the several orders of culm branches. This segmen-
tation is seen as an expression of physiological periodicity. The
mechanism of the origin of this type of structure poses intriguing,
and as yet unsolved, problems relating to morphogenesis. These
 VEGETATIVE PHASE: THE MATURING PLANT 13

consists of aseries of .nonfistulose, usually very short, segments

clothed with persistent, relatively small scalelike sheaths. The
nodes here are always without buds, and usually lack roots and
root primordia. In all axes of the plant; this basal part of each
axis is typically more slender than the part distal to it. Convention
I has already established the use of the term "neck" (Arber 1934:
67) for the constricted form it always assurnes in the l\nderground
parts of the plant (see text, p. 13 and Figs. 2,3, and 6). The super-
ficial resemblance of this transitional structure to a neck is in some
bamboos almost completely l~st elsewhere in the plant. In the
branches of the culm in all species of some genera (Arundinaria
and Phyllostachys, for example) it is very short and is not appreci-
ably constricted (Fig. 27).
As compared with other segmented axes, the "neck" appears
to be less specialized, noül.bly in its internal structure. Superficially
it appears, in its most common form, as a tnmsitional structure
of no special importance. However, the diversity of its manifesta-
tions of. form and behavior, especially in certain of the under-
ground axes, marks the neck as a focus of particular interest. The
basis of this assertion is made plain in the following pages (see
Fig. 1. The basic frarne of <: b~boo plant consists of ramifying systems of seg-
rnented axes. The charactenstlcs of the system of branching shown by the rhi- especially pp. 17 and 128).
z.omes, and of .that shown by the culms, have distinctive value für the recogni-
tlOn of the taxon to which a given plant belongs. The rhizome system
Rhizome axes are typically subterranean, and the rhizome system
problems are ably discussed by Porterfield (1933). They require
constitutes the structural foundation of the plant (Figs. 1-4).
furtber investigation by means of the tools and techniques of
Since this part of the plant is wholly or almost entirely out of
anatomy, biochemistry, and physiology.
sight and is not easily accessible, it is usually ignored by collectors,
The nodes of the segmented axes are important centers cf mor-
generally neglected by taxonomists, and practically unknown to
p~lOgenetic activity. In a growing point, it is at the loci later recog-
contemporary anatomists. 1 Consequently, the rhizome system is
mzable as nodes that sheath primordia are initiated and segmen-
one of the least weil understood parts of the bamboo plant. This
tation first comes into focus (see pp. 61 and 123). Only at the nodes
neglect has unfortunate consequences, because the rhizome system
do roots and branches emerge. The final growth in length of each
performs important functions in the life of the plant. Moreover,
segmented axis takes place önly near the nodes.Variations in the
an understanding ofthe form of therhizome system is prereq;'isite
form and other features of the nodes and internodes (particularly
to an understanding of the clump habit in any bamboo (Figs. 16,
those of the culm) afford useful characters for the differentiation
1 and 17). By the term "clump habit" I refer to the spatial relation
of taxonomie entities,
of the culms that make up the visible part of an individual bamboo
lShibata (1900, Figs. 1-14) illustrated some anatomical features of the rhizome of
The neck Arundinaria hindsii (sensu bot. jap.• non Munro), Phyllostachys pubcsccns (as Ph mitis), Psc.udosasa
(as Arundinaria) Japonica, Sasa (as Bamllusa) nipponica and Sasa (as Ba:mbusa) palmata. McClure
Each segmented axis is a branch of another segmented axis. The (1963) illustrated -anatomical features of thc rhizome that differentiate Arundinaria iecla
basal portion of each segmented vegetative axis of every category from A. gigantca.
14 THE BAMBOO PLANT

plant, whether the plant is caespi tose or diffuse in its spread in

space.
The rhizome system assumes, in plants of different species and
genera, a number of more or less -sharply distinct forms and habits Sud ---.j,

Root -----"<0..

Rhizome neck---4'"-t

+----Rhizome proper
1958

'~#---r--Culm neck
1956

Rhizome proper --:l!t:='---~

primordio--m~#'" __-",

-Sheollh scars----"

Fig. 3. Leptornorph rhizome system of Phyllostachys bambusoides, as seen from

, above, showing the base, of a culm (in cross section) and two new rhizomes, all
produced from lateral buds. Root primordia emerge in a single verticil at each
Fig. 2. Pachymorph rhizome system of Bambusa tuldoidcs, as seen from above, node of the rhizome proper, equally on all sides, and develop promptly. Those
showing individual rhizomes produced in successive years, and the base of the that have been cut off are shown as small circles. Roots (not shown) are also
culm invariably tenninal to each. The sheaths have been rernoved (note sheath borne at the several subterranean nodes of the obconical base of the culm, imme-
scars) to reveallateral buds, root prirnordia, and other details. Raots emerge on diately above (distal to) the culm neck. The smaller, unlabeled figure (from
the lateral and lower surfaces of the rhizome proper in an all-over pattern, while Takenouchi 1931a:PL 1, Fig. 4) is a lateral view based on the same species. Note
z.s a rule the upper surface is less completely covered; in this area the root pri- the abr:upt change in direction of the a.'-Üs on the left, from vertical to horizontal.
rnordia may remain dormant indefinitely.
 VEGETATNE PHASE: THE MATURING PLANT 17

commonly from a bud on another rhizome. Under certain condi-

tions a rhizome may emerge from a culm, starting as a bud either
at a subterranean node or, more rarely, at an aerial node of the
culm (see p. 245 and Fig. 97, 4-6).
The individual rhizome is a segmented axis consisting typically
of two parts: the rhizome proper and the rhizome neck (Figs. 2
and 3). The neck is basal to the rhizome proper, and is the part
that develops first. In its most common form, the rhizome neck
is relatively short and obconical in shape but it may be greatly
elongated and obterete (Fig. 13). Its basal diameter is smaller
than that of either of the axes it joins together. In bamboos with
pachymorph rhizomes (p. 24) the rhizome neck carries the rhizome
primordium that is apical to it downward to a depth that is char-
acteristic for each species, the age of the plant, and the nature
Fig. 4. An early illustration of the type of bamboo rhizome herein nam.ed of the substrate. This behavior of the neck often assumes a dra-
leptomorph, exemplified by Phyllostachys viridis. The terminal bud of the rhizome
in this bamqoo and all other species of the genus Phyllostachys that have ,been BAMBUSA VUlGARJS, Schrad_ CA rhizome plexus seen from
studied is normally diageotropic. The rhizome is pictured here as_turning upw~d bases of vertical aerial axes indicated in black
at length (feft) to form a culm (see metamorph II axes, p. 30). The manifestation
of negative geotropism by the terminal bud of the rhizome is facultative, not
obligate, in bamboos of this genus. The tillering of culms shown at the right is
also facultative, and rare, in species ·of Phyllostachys. It can appear in bamboos
with leptomorph rhizomes only where there are buds at the underground nodes
of the cuIm. From A and C. Riviere 1879:Fig. 3 Cas Ph mitis). See p. 51.

of growth (Fig. 10). These diverse manifestations afford challenging

openings for fruitful studies from the points of view of physiology
and anatomy as weil as gross morphology. They may be used for
the recognition and description and, to a limited extent, the dassi- .
fication of taxonomie entities.
Arber (1934:67) refers to the individual axes (branches) of the
bamboo rhizome system as "rhizome segments." For the rhizome
in Bambusa vulgaris (see Fig. 5) this term appears at first sight to
be quite suitable. However, when the clearly segmented nature
of the individual rhizome axis is considered, Arber's use of the
term is seen to be ambiguous. Its application to the individual
branches of the slender, wide-ranging type of rhizome (see Fig. 3) Fig. 5. This slightly stylized sketch by Agnes Arber C1934:Fig. 23) gives only an
is even more clearly inappropriate. It seems best, therefore, to inkling of the complicated mass ofintervvoven axes that the pachymorph rhizome
system of a bamboo plant becomes in time. Ponion of a specimen (No. 205) of
apply the term "rhizome" to each individual branch or axis of Bambusa vulgaris preserved in the Museum at the Royal Gardens, Kew, seen from
the rhizome system. Each rhizome develops from a bud-most below; greatly reduced. See also Fig. 2 and pp. 24f.
18 TEE BAMBOO PLANT VEGETATIVE PHASE: TEE MATURING PLANT 19

matic aspect in young seedlings; see p. 128 and Fig. 60,i (Dendro- the culms develop from the lateral buds oflong, slender rhizomes,
calamus strictus), also Fig. 61 (Melocanna bacciftra). Again, in large commonly referred to in earlier literature as trar;ant, monopodial,
plants of Guadua angustifolia the rhizome necks show, in their or indeterminate, and hereinafter referred to as leptomorph (Figs.
"start-and-stop" growth and in their physical orientation, a special 3, 4, and 8, B).
adaptation related to the support of the culms (Fig. 6). The rhizome proper is characterized by its typically subterra-
Arber (1934:67) apparently was the first to use the term "neck" nean position, by the presence cf reets cr root primordia and
for this part of the rhizome, attributing its detailed study to Shi- prophyllate buds (always solitary) at all or most of its nodes, and
bata. As a matter of fact, the structure described by Shibata by the reIativeIy simple and uniform character of the sheaths that
(1900:443, Figs. 13, 14) was the neck ("Stiel") of the young culm clothe it (Fig. 34). Each diageotropic rhizome axis normally ori-
("Schössling") and not that of the rhizome. Shibata says in foot- a ents itself so that the plane of distichy in the insertion of sheaths
note: "This 'Stiel' furnishes the sole path for the movement of and buds or branches lies at right angles to the force of gravity.
materials between the rhizome and the culm that develops from
the shoot." This statement is incomprehensible to the reader who Two basic forms of rhizome
is unaware that Shibata was referring only to bamboos in which. The rhizome manifests its character in two divergentforms, each
with important variations. Differences in the gross morphology,
combined with differences in growth habit, give rise to charac-
teristics by which a number of more or les.s clearly distinct forms
of rhizome system and clump habit may be recognized (see p. 32
and Fig. 10).
It appears that in their classic work, Les Bambous (1879:312-322),
the Rivieres were the first to publish (in a Western language, at
any rate) a dear distinction bet:ween the two basic forms assumed
by the bamboo rhizome. These were taken as characteristic of two
distinct groups of bamboos (see p. 209) described aso
"1. Bamboos of autumnal growth, and caespitose dump habit;
"2. Bamboos of spring growth, anda generally spreading
[tra,antJ dump habit" (p. 312).
An unidentified species of Gigantochloa-under the name Bam-
busa macroculmis Riviere and Riviere (1879:Fig. 2)-was chosen as
exemplifying the first group. Its rhizome is described as developing
in a horizontal direction for a short distance, after which it turns
upward to form a cp.lm (as in Figs. 2 and 7).
Phyllostachys viridis (under the name Ph. mitis) was chosen as
Fig. 6. In large plants of Guadua angustifolia~ the lateral buds of the pachymorph exemplifying the second group. Its rhizome is described as ranging
rhizome push prematurely. and make a limited growth to form only the neck widely i'l- a horizontal direction, with more or less widely spaced
portion of the next order of branches, thus providing' a collective "foot" to sup- culms arising from later;al buds at the nodes (as in Figs. 3 and 17).
port the -heavy culm that is produced at the same time [rom the terminal bud of
the mother rhizome. In this way, the tough, elongated necks of as yet incomplete The next contribution to the subject is that of Houzeau de
rhizomes form an indispensable support for the lOO-ft culm. Lehaie (1906'-1908:150), who called the rhizome <;>f bamboos
20 THE 'BAMBOO PLANT

of the first group a "caulo-bulbe," and illustrated tbe rhizome of

Bambusa vulgaris (under tbe name B. thouarsiz) as an example (Fig.
7). Houzeau characterized the rhizome of bamboos of the second
group as "long and slender, with indefinite subterranean develop-
ment." As an example, he illustrated the rhizome cf an unspeci-
fied member of the genus Phyliostachys. This author concluded his
discussion with tbe challenging statement tbat "the essential dif-
ference between caespitose bamboos and running [tra,antJ bamboos

Fig. 8.. Types of rhizome designated by McClure (1925:Figs. 1, 2) as CA) sym-

padial (Bambusa beecheyana); CB) monopodial (Arundinaria a:mabilis). These terms
were later, and perhaps more appropriately, applied by Japanese botanists
(notably Nakai) to the manner of origin of culms associated, respectively, with
the two types of rhizome.

is neither generic nor specific, but is of a: physiological nature."

The recent discovery of the occurrence (in Chusquea Jendlerz) of
botb types of rhizome in tbe same plant (see Fig. 10, 11) appears
to confirm this assertion. However, Houzeau's observation loses
its intended force when we reflect that all morphological features
Fig. 7. Bambusa vu.lgans shows the type of rhizome herein named pachymorph. recomized as "characters" at either the specific or the - generic
b
Under the synonymous name, Bambusa thouarsi~ this species was chosen by
level a"I-e basically physiological in origin.
Houzeau de Lehaie (1906:Pl. ill) to illustrate the development of a typical
"caespitose" bamboo. See Fig. 5. McClure (1925) first proposed the terms monopodial and sym-
22 THE BAMBOO PLANT VEGETATIVE PHASE: THE MATURlNG PLANT 23
podial to characterize the branching habit of the rhizome in the ferent manifestations of the rhizome system. These were then des-
two main groups of bamboos called by the Rivieres "tral:ant" ignated as folIows: "1. Simple clump-forming; 2. Clump-forming
and "caespitose," respectively. Takenouchi (1926:44) adopted from runners; 3. Lateral clump-forming from rhizomes, and
McClure's terminology in listing three "kinds" o(rhizomes, and 4. Irregularly branching" (p. 4). Reproduced in my Figs. 11, 13,
the genera of Japariese bamboos that exemplify them. However, 14, and 3 (insert), respectively, these are interpreted herein as two
four distinct forms of growth (Fig. 9) were actually illustrated. typical and more or less distinctly diverse e,:pressions of each of
Takenouchi later (193Ia:Fig. I, '1-4) again illustrated four dif- the two basic types. Later in this work, however,' Takenouchi
shows signs of faltering between the designations "rhizome" and
"culm base" in his interpretation of the short, thick, determinate
Q type hereinafter .called pachymorph; and in the section captioned
~!I!l1 i "Morphology and characteristics of rhizomes" (Takenouchi 1932:
)? A trans., p. 7) he deals concretely only with the long, slender, inde-
terminate type, hereinafter called leptomorph.
In writing about various Japanese bamboos, Nakai (1933:12),
Makino, and others have rather consistently (and with good reason)
used the terms "monopodial" and "sympodial" in reference to
the manner of origin of the culms rather than to the habit of the
associated rhizome. M'cClure later (1945b, 1946) substituted the
terms ((indeterminate" and "determinate," in the belief that these
are more precise and caU attention to a more basie differenee in
the behavior of the growing point of the rhizome prevailing in the
two forms respectively.
To the distinctive branching habit in the underground part of
the plant which Takenouchi (193Ia:Fig. 1, 3; trans., p. 4) calls
"lateral clump-forming from rhizomes," Keng (1948:2) gives the
name "amphipodial." Its characteristics are described on p. 31.
Shibata (1900:Figs. 1-14) was the first to publish detailed
comparative studies of the anatomy of the rhizome in different
bamboos. However, he dealt only with leptomorph rhizomes, and
made only the briefest reference to their external morphology. He
did not differentiate the two basic forms discussed herein. McClure
(1963) described and illustrated a hitherto unrecorded anatomical
Fig.9. Types ofrhizome system illustrated by Takenouchi (1926:PI. 3, Figs. i-iv). feature, the presence of peripheral air canals in the rhizomes of
CA) Leptomorph rhizomes. with solitary culms, designated by Takenouchi as Arundinana teeta. This feature, paired with the lack of it in the
"Monopodien (Phyllostachys)." (B) Leptomorph rhizomes with tillering culms,
designated by Takenouchi as "Monopodien und Sympodien (Pleioblastus)"; this
rhizomes of Arundinaria gigantea, constitutes a reliable taxonomie
apparently corresponds to the category "amphipodial" of Keng, f. (1948:2). (C) character for differentiating the two species.
Short-necked pachymorph rhizomes, designated by Takenouchi as "Sympodien
(Dendrocalamus~ Bambusa)." CD) Long-necked pachymorph rhizomes with tillering Reinterpretation of the rhizome system
culms, designated by Takenouchi as "Sympodien (Indocalamus)" [= Yushania
niitakqyamensis]. Takenouchi later (1932:Pl. 4, Fig. 1) referred to the long neck in Further comparative studies of the diverse forms of growth in
"Indocalamus" as a "running rhizome." See also Figs. 11, 13, 14, and 3 (insert). the underground parts of bamboos should intrigue the physiologist
 24 THE BAMBOO PLANT VEGETATIVE PHASE: THE MATURING PLANT 25
and the anatomist, as well as the morphologist. The taxonomist podial" (McClure 1925) 'and "determinate" (McClure 1946). In
and the field botanist could use the results to good advantage~ pachymorph rhizomes, the rhizome neck may be short or long.
Meanwhile, improvements in the differentiation , illustration,
description, and terminology of the known modes of expression in Leptomorph rhizomes
the bamboo rhizome system are desirable. The pairs of terms; 1t is proposed to call the rhizome proper leptomorph when it is
"caespitose" and "trafant," "sympodial" and "monopodial," long and slender, and has the following associated characteristics:
"determinate" and "indeterrninate" have proved increasingly a cylindrical or subcylindrical form, with a diameter·usually less
difficult to use in an unambiguous manner, even with the help of than that of the culms originating from it; internodes longer than
Keng's new term, "amphipodial"-probably because they embrace broad, relatively uniform in length, symmetricalor nearly so,
too many undefined ideas. At any rate, in actual use they all rarely solid, typically hollow, the usually narrow central lumen
eventually break down in ways that leave either the original con- interrupted at each node by a diaphragm; nodes in some genera
cepts or the distinctive features of associated variables somewhat usually somewhat elevated or inflated, in others not; lateral buds
blurred., and they cannot easily be modified. It now appears that in the dormant state. boat-shaped, with a distally oriented apex;
a slight shift and a narrowing of the initial focus of attention may see Figs. 3 and 10, 7-10.
improve the interpretation, the terminology, and the descriptions. In leptomorph rhizomes generally (as in the known species of
")
Phyllostachys) every node bears a solitary prophyllate bud and a
This changed view will be presented by characterizing the two
basic forms of the rhizome proper as pachymorph and leptomorph. single verticil of roots. In some bamboos, .however (as in many k
arundinarioid species-especially in the genus Sasa), buds may be
lacking here and there at one to several adjacent nodes, and root
Pachymorph rhizomes
development may be very sparse or even lacking entirely at some
It is proposed to call the rhizome proper pachymorph when it nodes. No sign of dorsiventrality has been noted, but each rhizome
is short and thick, and has the following associated characteristics: axis shows a strong tendency to orient itself so that the plane of
a subfusiform (rarely subspherical), usually more or less curved insertion of sheaths and buds is horizontal.
(rarely straight) shape, with a maximum thickness typically some- Most of the lateral buds of leptomorph rhizomes are tempo-
what greater than that of the culm into which it is always trans- rarily or permanently dormant. The majority of those that ger-
formed apically; internodes broader than long, asymmetrical minate produce culms directly; a few produce other rhizomes.
(longer on the side that bears a bud), solid (apparently never The orientation of the terminal bud in leptomorph rhizomes is
hollow); nodes not elevated or inflated; lateral buds solitary, in consistently diageotropic in most species and under most circum-
the dormant state asymmetrically dome-shaped, with a subcircular stances. The persistence of this diageotropic character suggested
margin and an intramarginal apex. In pachymorph rhizomes the terms "monopodial" (McClure 1925) and "indeterminate"
with a horizontal orientation, varying degrees of dorsiventqJity (McClure 1945a:7 and 1945b:278). In certain species regularly
are manifested, in (1) a more profuse production of roots on the' (as in Arundinaria tecta, Fig. 10, 8), and in others only under certain
lower side and (2) dorsiventral flattening of the axis; see Figs. conditions (Phyllostachys spp., Fig. 4), the leptomorph rhizome turns
6 and 7. upward to form a culm. In leptomorph rhizomes, the rhizome neck
Lateral buds of a pachymorph rhizome produce only rhizomes; is always short.
a culm can arise directly only from the apex of such an axis. A Ueda (1960) has made extensive excavations of the leptomorph
transformation of the internal organization of a pachymorph rhizome systems of plants of several species of J apanese bamboos,
rhizome axis is prerequisite to culm formation. This feature was plotting the pattern and r:"te of their extension in the soil, and
the basis for the characterization of this type of rhizome as "sym- calculating the rate of accumulation of total substance.
 VEGETATIVE PHASE: THE MATURlNG PLANT 27

proper show buds at their nades; rhizome necks have no buds. The scale cf
representation is not uniform. Captions to individual figures follow. See p. 32 for
• key.
1. Bambusa pachinensis here exemplifies a shart-necked parchymorph rhizome
in which there is a marked tendency toward wholly erect orientation, that is,
4 negative geotropism. Takenouchi's illustration of this species, showing the growth
habit resulting from this orientation of the rhizome, is reproduced in Fig. 1l.
2. A typical and widely representative expression of the pachy:norph rhizome
system is that of Ba:mbusa tuldoides, here illustrated. The rhizome neck is shorter
than the rhizome proper. Thc rhizome proper is horizontal in its proximal part,
curves upward in its distal part, and terrninates in an crect culm. Thc resulting
clump is caespitose, that is, a compact tuft of culms. This is the type of growth
first referred to as caespitose (Riviere 1879:312 and Fig. 2), later by McClure
(1925) as sympodial, and still later (McClure 1946:107) as determinate. Sec
Figs. 2 and 8, A.
3. In Arundinaria pusilla the growth of the terminal bud of the short-necked
pachymorph rhizome sometimes deviates from the typical 'c"xpression shown by
Bambusa tuldoides. Beyond the rhizome proper, the axis describes in this case a
broad curve, and its diameter is gradually reduced before it emerges from the
ground as a culm. This transitional portion of the axis is typically rootless, but
it usually has a bud at each node. Observations made in South Vietnam in 1953
5 6 indicate that in this species pachymorph rhizomes with two distinct potentialities
emerge from the subterranean buds. In those a..xes that originate be10w a certain
depth, a long transitional stage (here called a metamorph TI axis) intervenes
between the rhizome proper and the culm proper. In rhizomes originating above
a certain depth, the terminal bud produces a culm directly, without an inter-
vening metamorph TI axis. In the Andean highlands of Ecuador, I have observed,
in the rhizome system of large, old cIumps of Chusquea sca:n.dens) a growth form
similar to the one here described. See Fig. 12 and p. 30.
4. In some bamboos with pachymorph rhizomes. the rhizome neck is elongated
just enough to give the caespitose clump a noticeably open habit. A typical
example of this is afforded by Sina:rundinana nitida, here illustrated. Bambusa
vulgaris shows the same trend.. See Fig. 7.
5. Melocanna baccifera> here illustrated, is one of a number of bamboos that
are characterized by- long-necked pachymorph rhizomes producing solitary
• culms. Here, and in Schizostachyum hainanense) the rhizome neck may be a meter
or more in length. In a bamboo native at Tingo Maria, eastern Peru (Guadua
sp., McClure 21438-A) the rhizome neck reaches a length of 6 m. Such an
e10ngated rhizome neck. is subcylindrical throughout most of its length, c"xcept
11 near the distal end, where it expands into a rhizome proper that is terminal to
it. It is wholly free of buds, and only exceptionally bears a few roots on the
lower side. This type of growth gives rise to a simple,. open clump made up of
widely spaced solitary culms. See Fig. 82 and p. 36.
6. Long-necked pachymorph rhizomes and culms that tiller are characteristic
of Yushania niitako:Jamensis, here illustrated. Buds at the base of culms terminating
Fig. 10. Known forms assumed by the rhizome system in different bamboos are long-necked rhizomes give rise to axes with two distinct potentialities. They
bere illustrated by diagrammatic views from above. Open ends signify the con- produce culms either (1) by way of long neck.s, or (2) by tillering, through
tinuation cf an a.....::is in a horizontal direction. Double cirdes represent basal short-necked rhizomes' that often resemble metamorph I axes. The result is a
cross-sections cf hollow culms. Single circles in<;licate culrns arising by the slow number of spaced-out tufts of culms, joined together by the long-neck.ed rhi-
transformation cf the terminal bud cf a rhizome through a metamo'rph II axis. zomes-a culm habit that may be described as compound-caespitose. See also
Transverse lines indicate sheath scars at nodes; roats are omitted. Rhizomes Fig. 13 and p. 40.
 28 THE BAMBOO PLANT

7. Shibataea kwnasasa apparently -is facultative in respect to the behavior of

the growing point of its rhizomes, and in respect to tillering of the culms. The
rhizome nonnally remains diageotropic in its growth, but in an occasional one
the growing point becomes negatively geotropic and turns upward to form a
culm, through a metamorph TI a-as. Culms arising from lateral buds of the rhi-
zome norrnally remain solitary. Occasionally, however-perhaps predominantly
in age-they tiller to form tufts of additional culms, by way of metamorph I
axes-a condition illustrated in 8. See also Fig. 14 and p. 36.
8. In Arundinaria tect~ shown here, the growing point of the leptomorph rhi-:
zorne, after carrying the growth of the a..ns a certain rather uniform distance in
a horizontal direction, develops negative geotropism, whereupon the axis turns
upward in a broad curve and at the same time gradually modifies its form to
that of a culm, by way of a metamorph TI axis. This behavior of the rhizome
apparently is obligate in Arundina:ria tect~ but is more commonly facultative
elsewhere-as in the example shown in 7. From their subterranean buds, all
culms produce both rhizomes and other culms, the latter by tillering through
metamorph I axes. See pp. 34 and 40.
9. A leptomorph rhizome with a normally diageotropic growing point, and
culms that normally do not tiller, are here exemplified by Phyllostachys bambu-
soides. Characteristic of all species of Phyllostachys that have been studied, this is
perhaps the most widely typical expression of the leptomorph rhizome. The
diageotropic growth of the apex of the rhizome proper extends the latter steadily
in a generally horizontal direction. If and when, for any reason, the apical
meristem ceases to funtion-upon the termination of a season's growth, for
e..xample-the growth of the plant may be carried on in the same general direc-
tion by one or more new rhizomes that emerge from buds near the tip of the old
one (Fig. 3). A lateral bud of the rhizome proper may give rise to either a culm
or a rhizome. On the periphery of plants of mature stature the culms usually
emerge at an open spacing, giving a diffuse clump habit. The culms normally Fig. 11. In some bamboos the pachyrnorph rhizome shows a marked tendency
do not tiller, but may do so when damaged, or when growing under unfavorable toward a wholly erect orientation. Takenouchi here uses Bambusa pachinensis to
conditions . .In Phyllostachys, the apex of the rhizome does not ordinarily develop illustrate this habit of growth, which he represents as typical for the genus
negative geotropism, but it may do so under exceptional conditions (cf. Fig. 4) Bambusa. Perhaps Takenouchi's inclination (1932:trans., p. 7) to interpret the
as when forced upward by natural obstacles, or when it emerges from the gro~d pachyrnorph rhizome as a culm base, instead of a rhizome, grew out of his
upon encountering a steep declivity. See Fig. 16, 2. choice of this example as typical of the whole genus. Redrawn from Takenouchi
10. In Indocalamus sinicus, shown here, the leptomorph rhizome rernains dia- 1932,Fig. 4,. 1. See also Fig. 10, 1.
geotropic, and the formation of tufts or' secondary culms through tillering by
way of metamorph I a.xes appears to be the normal e..'\."}Jression. Takenouchi
(1926:Fig. 3, TI) illustrates a similar habit in Arundinana (as Pleioblastus) simonii, and "culm base," as described above. The anomalous form and
and refers to it as typical of the general Pleioblastus and Pseudosasa. See Figs. 9, B
and 18, p. 34.
behavior of these axes, and their position in the plant, suggest the
11. The association, in the same plant, of both leptomorph and pachymorph general designation "transitionaL" Für convenience, the term
rhizomes, apparently rare among the bamboos, is here e.."ernplified by Chusquea "metamorph" is proposed as a technical designation of such tran-
fendleri. The swollen character of the pachymorph rhizomes that appear here in
association with those of leptomorph habit distinguishes this form of e.."pression
sitional axes. Evidence at hand suggests the characterization of
from the one illustrated in the foregoing figure, where slender, metamorph I two kinds of metamorph axes: metamorph I and metamorph ll.
a."es appear in their stead. See also Fig. 15 and p. 31. This rough classification is based primarilyon the respective posi-
tions in which they appear in the plant.
Metamorph axes A metamorph I axis occupies a positIon between the neck proper
In some bamboos, certain axes or portions ofaxes of subterranean and its culm. In :form, it is intermediate between a typical culm
ongln do not fit exactly into the categories "neck," "rhizome," base and a pachymorph rhizome, as contrasted in Fig. 20. Such
30 THE BAMBOO PLANT

axes often appear where the culms arising from lateral buds of a
leptomorph rhizome tiller to form tufts, as in Figs. 9, B and 14.
A metamorph II axis is intermediate in form and position
between the apex of a rhizome (pachymorph or leptomorph) and
the culm into which the rhizome is transformed apically. It
appears where the transformation of the apex of a rhizome into
a culm takes place gradually and not abruptly. This gradual
change is typical wherever a: leptomorph rhizome is transformed o

apically to form a culm (Figs. 4 and 10, 7 and 8). The interca-
lation of a transition zone (metamorph II axis) between a pachy-
morph rhizome and the culm distal ·to it, as seen in Arundinaria
pusilla (Fig. 10, 3 and 12) and some species of Chusquea, is rela-
Fig. 13. The combination cf long-neck.ed pachymorph rhizomes with culms that
tiller was first illustrated by Takenouchi, on the basis cf Indocalamus niitakayamen-
sis~ now known as Yushania niitakayamensis. The structure that is interpreted
herein as a rhizome neck Is in this case called by Takenouchi a "running rhi-
zome." The culms that terrninate these long necks tiller by metamorph laxes.
Redravm [rom Takenouchi 1931a:Fig. 1, 2. See also Fig. 10, 6.

tively rare, and stands in sharp contrast with the abrupt trans-
formation that is typical of bamboos with pachymorph rhizomes,
as illustrated in Figs. 2 and 7.

Other combinations of underground growth forms

As I underst"nd it, the habit of growth designated by Keng
(1948:2) as amphipodial is found in the underground parts of cer-
tain species of th~ arundinarioid genera (Arundinaria~ Indocalamus~
Pseudosasa, Shibataea, Sasa). It is the result ofthe occurrence together,
in the same plant, of two distinct kinds of subterranean axes. One
of these is of the type described above as leptomorph; the other
r----Mletelmc>rDhlJ axis
corresponds to the concept described above as metamorph I.
Examples are ilIustrated in Figs. 9, B and 14.
#lI\hW~~ ~~~;--paChymOrPh rhizome The capacity to produce both typical pachymorph and typical
leptomorph rhizomes in the ·same plant is represented among the
buds of the subterranean axes of Chusquea fendleri and some other
species of this genus. The resulting form of expression is illustrated
Fig. 12. Metamorph TI a.-.::es distal to MO pachymorph rhizomes, as exemplified
by Anmdinana pusilla. Fie1d sketch, made en route from Saigon to Daht, South in Figs. 10, 11 and 15. This potential is not known to exist in any
Vietnam, December 24, 1953.-See also Fig. 10,3. of the bamboos'mentioned by Keng.
 32 THE BAMBOO PLANT

I
•

Fig. 14. Tillering of the culms in Shibataea kumasasa appears to be a facultative

character in this species. Redrawn from Takenouchi 1931a:Fig. 1, 3. See also
Fig. 10, 7.

Key to known forms of the bamboo rhizome system, with examples

and illustrations
1. Rhizomes proper all pachymp~p.h; culms each
developed from the tip (apic\li meristem) of a
rhizome I

2. Rhizome neck rouch shorter than the rhi-

zome proper emerging from it
3. Rhizome proper nearly erect; clump Bambusa
compact-caespitose (Figs. 10, 1 and 11) pachinensis
3. Rhizome proper typically horizontal Fig. 15. Chusqtaa findlm. The capacity to produce either pachymorph cr lepto-
4.- Rhizome-culm transition always Bambusa morph rhizomes (cr both), is potential in the buds of the subterranean axes of
abrupt; clump compact-caespitose tuldoides this and some other species of Chusquea. The enlarged node above shows the
(Figs. 2 and 10, 2) principal branch bud and, below it, a cluster of smaller, subsidiary buds Ca
4: Rhizome-culm transition often pro- Arundinaria character peculiar to the genus Chusquea) and also several spinelike aerial roots.
longed in a metamorph TI a.,..,üs; pusilla Field sketch from plant represented by specimen under McClure No. 21236.
clump compact- cr open-caespitose See also Fig. 10, I I, and p. 31.
(Figs. 10, 3 and 12)
 34 THE BAMBOO PLANT

2. Rhizome neck rioticeably elongated; clump Szonarundinaria

open-caespitose (Fig. 10, 4) nitida
2. Rhizome neck greatly elongated
5. Culms solitary; clump open, diffuse (Fig. Melocanna
10,5) baccifera
5. Culms tillering via metamorph laxes to Xushania
[Olm tufts; clump compound-caespitose niitakayamensis
(Figs. 10, 6 and 13)
1. Rhizomes proper all leptomorph; rhizome necks
all short; culms arising primarily from lateral
buds of rhizomes
6. Growing apex of rhizome becomes negatively
geotropic and emerges as a culm, by way of
a metamorph II a"(is
7. Negative geotropism in rhizome axis obli- Arundinaria
gate, the reaction usually rather prompt; tecta
rhizomes show peripheral air canals
(Fig. 10, 8)
7. Negative geotropism in rhizome axis Shibataea
facultative, the reaction often much de- kumasasa
layed; rhizomes lack peripheral air canals
(Figs. 10, 7 and 14)
6. Growing apex of rhizome typically remains
diageotropic; it becomes negatively geotropic
and emerges from the ground as a culm only
in response to external forces
8. Culms typically solitary, normally not Phyllostachys
tillering to form tufts; clump diffuse bambusoides
(Figs. 3, 10, 9, and 17)
8. Culms typically forming tufts by tillering Indocalamus
through metamorph laxes; clump com- sznzcus
pound:caespitose (Fig. 10, 10)
1. Rhizomes proper of two types, pachymorph and Chusquea
leptomorph, in the same plant; all rhizome necks findleri
short; clump compound-caespitose (Figs. 10, 11
and 15)

Clump habit
The nature and meaning of the typical form assumed by the
visible part of a given bamboo plant becomes clear only after the
form and growth habit of its rhizome system are fuUy understood.
Bamboos with short-necked pachymorph rhizomes grow in dis-
crete, compactly caespitose clumps (Dendrocalamus membranaceus,
Fig. 16, 1). Pachymorph rhizomes ~th a slightly enlongated neck
produce a less compact clump (Sinarundinaria nitida and Bambusa
 36 THE BAMBOO PLANT

vulgaris). Bamboos with leptomorph rhizomes produce culms typi-

cally in open array (Phyllostachys pubescens, Fig. 17), and the
culms of bamboos with only long-necked pachymorph rhizomes
(Melocanna baccifera, Fig. 82) stand similarly isolated. In writings
about the bamboos of India, all bamboos manifesting this open
type of clump habit are usually lumped together as "single-
stemmed," without distinction and without reference to dispaI1.ties
in subterranean characteristics. Bamboos with long-necked pachy-
morph rhizomes and culms that tiller via metamorph laxes pro-
duce composite clumps made up of compact tufts that appear
separate and distinct but actually are connected below the ground
by elongated rhizome necks (Yushania niitakayamensis, Fig. 13).
This same clump habit is found in bamboos that have both lepto-
morph and pachymorph rhizomes (Chusquea fendleri, Figs. 10, 11
and 15) and also in those that have leptomorph rhizomes and
culms that tiller via metamorph laxes as in Indocalamus sinicus
(Fig. 10, 10), Semiarundinariafastuosa (Fig. 18), and Shibataea kuma-
sasa (Fig. 14). In any particular plant of some of these bamboos,
deviations induced by any influence that interferes with its free
natural growth may obscure the e,,:pression of clump habit pre-
dicated above.

The culm
Bamboo culms are either lateral branches or apical projections of
the rhizome, depending on whether the rhizome proper is lepto-
. morph or pachymorph. In habit, they vary from strictly erect,
erect with pendulous tips, or ascending, through broadly arched
to clambering (Fig. 19) and from nearly straight to strongly zig
zag. Given uniform environmental conditions, each habit charac-
ter is fairly consistent within a given taxonomie entity, such as a
species or a subspecies.
In any given bamboo plant, the culms may originate in one
(or sometimes two) of the following ways: (1) as the transformed
distal end (apex) of a pachymorph rhizome, either with or with-
out the intervention of a metamorph II axis; (2) from a lateral
bud of a leptomorph rhizome; (3) from a bud on the underground Fig. ~ 7. Open diffuse clump habit in a bamboo with leptomorph rhizomes, ex-
base of a culm, by tillering, or by way of a metamorph I axis; emphfied by Phyllostachys. puhescens in the garden cf Mr. Sankichi Ishida, at
(4) as the transformed distal end (apex) of a leptomorph rhizome Komazawa, near Tokyo, Japan. By selective harvesting cf the young shoots, the
average distance between culms is kept at a level corresponding to optimum yield.
through a metamorph II axis. U.S.D.A. photo by P. H. Dorsett. See Fig. 3.
38 THE BAMBOO PLANT

I c

r
~
~~

F H

Fig. 19. Selected- exarnples of culm habit, illustrated diagrarnmatically. (A) Sasa
Fig. 18. Semiarundinariafastuosa (P.l. 112080). Underground part of aplant, show- palmata; CB) Sinocalamus beecheyanus; (C) Phyltostachys nigro,' CD) Schizostachyum
ing a leptomorph rhizome and the characteristic tillering of a primary culm from hainanense; CE) Dinochloa standens (auctt. non Kuntze); CF) Sinocalamus affmis; (G)
a basal budo U.S.D.A. photo by Robert Taylor. See Figs. 10, 10 "and 20. BambuSa textilis; (H) Arundinaria amabilis. Note the result of circumnutation that
took place during the growing stage of the culms in CE). Frofi McClure and
Li 1941:Fig. 1.
 40 THE BAMBOO PLANT

A culm that develops from a lateral bud of a leptomorph rhi-

zome (Semiarundinaria fastuosa, Figs. 18 and 20, right) is an example
of the most complete form. It shows two parts: the aerial culm
proper and the subterranean culm base. The culm base consists
of two structurally distinct parts: the culm base proper and the
culm neck. The culm base proper is a narrow, inverted cone with
each node marked by the presence of a sheath, avertieil of roots,
and, in many cases, a bud. In aplant that tillers, such a bud can
give rise to a new culm. The internodes may be solid but gener-
ally are hollow and short, increasing in length by small increments
from the basal one to the uppermost, which appears at about the
level of the surface of the ground. At this point the culm proper
begins, <itS signalized by a more or less abrupt increase in the
length of the successive intemodes, which here take on a niore
strictly cylindrical form. At its lower end, the inverted cone of
the culm base proper is attached to its point of origin on the rhi-
zome by the slender, curved neck.
A culm arising as the transformed distal end of a leptomorph
rhizome (Figs. 4 and 16, 2) has a metamorph II axis in place of·
the specially differentiated subterranean culm base and culm necK-
just described. Even without direct evidence of its actual relation-
ship to the rhizome, the peculiar origin of such a culm is made
clear by the character of this basal part. Its lower part is broadly
curved in an upward sweep; its lower intemodes are solid Or
nearly so, and shorter than those of culms that emerge from lateral
buds of the rhizome. The change from the small-lumened o;r
rarely solid condition of the internodes of the rhizome axis to the Fig. 20. Lower part of a culm of each of two species-(iq2) Dendrocalamus lati-
. large-lumened condition of the internodes of the culm takes place florus, (right) Semiarundinariafastuosa-both split in two, to illustrate Takenouchi's
gradually, though in some species (Phyllostachys bambusoides) the interpretation of the structures shown: "Culm with [1q2] and without [right]
sharp distinction between the true culm and the underground culm base: CA)
internodes of such a culm may all be solid. culm proper; CB) underground culm base; (C) stipe or stalk." As interpreted
A culm arising within a pachymorph rhiz6me system typically herein, CA) is the culm proper in both; CB) in the culm on the left is a pachymorph
originates only as the transformed distal end~of a rhizome axis, rhizome, and in the culm of the right it is the culm base; (C) in the culm on
the left is a rhizome nec~ and in the culm on the right it is a culm neck. The lepto-
as in Bambusa tuldoides (Fig. 2). In all such culms, the place of the
morph rhizome of the culm on the right, not shown here, may be seen in Fig.
culm base proper is taken by the rhizome itself, to which the 18. The discrepancies between the two interpretations given above illustrate
culm is here terminal. A neck is found only at the base of the the observation made elsewhere herein (p. 6) that "some of the outward forms
rhizome, and there is no culm base proper and no culm neck and behavior of the bamboo plant are not easily"amenable to the rigid applica-
tion of a hierarchy of technical terms or reasoned concepts." Redrawn from
(Figs. 2 and 20, left). Exceptionally, new culms may arise in a Takenouchi 1931a:Fig. 4. . .
bamboo with a pachymorph rhizome by tillering, by way of a
metamorph I axis, as in Yushania niitakayamensis (Fig. 13), or by
42 THE BAMBOO PLANT

way of a metamorph n axis that intervenes between the rhizome 35

and the culm proper as in Arundinaria pusilla (Fig. 12). /\
30
Small culms generally taper gradually from base to tip. In
larger ones, the lower half or so is roughly cylindrical or imper- w 25
(
Cl
ceptibly tapered, andthe rest of the culm more sharply so. In the o
Z
20
a \~~-- "
,
\......>,; •••••••.. .... "
largest ones from vigorous plants of many species (in the genus Ir
,
Phyllostachys, for example) there may be an appreciable increase ....
W
I
;

~ 15
in the diameter for sorne distance, beginning at or near the base, LL
o
:b
then a gradual decrease in diameter, and finally, in the upper % 10r-7'~r·~···_··--~~~_···_··2··~~--~--~
or '!. of its length, the culm is more sharply tapered. ....J:
(!)
I .•••
/./c ~ \
, •.•••• ....
In the cyde of increase and decrease in length, and in the pro~ Z "5 I , .....
lU .. \ ...........
..J
gressive change of shape, of the successive internodes and their O~--~
I
____L -____L-__ __ ~L- ~

associated sheaths (a combination of structures illustrated by 1 5 10 15 20 25

Porterfield, 1930b, Fig. 4, as phytons) bamboo culms and their SERIAL NUMBER OF INTERNODE
branches manifest a periodicity that is weil described and docu-
mented by PorterfieId (1933). Shigematsu (1958) has developed Fig. 21. Graphie representation of the march of increase and decrease in inter-
mathematical formulas, based on detailed measurements, to eluci-
node length in the culms ofthree barnboos: Ca) Bambusa multiplex B. nana var. (as
typica); (h) &nobambusa toots-iJs' and Ce) Arundt"nana (as Pleioblastus) simonii. Redrawn
date the patterns of change in length and diameter of culm inter- from Takenouchi 1932:Fig. 16.
nodes and thickness of culm wall characteristic of 15 species
representing 8 genera of J apanese bamboos. first internode, 0.( two or more nodes without an intervening inter-
The march of increase and decrease in internode length, as node, intercalary growth having been suppressed locally. Here
found in self-supporting bamboo culms, may be graphically repre- the consecutive sheath scars are usually less than 1 inch distant
sented by plotting the length of each internode against its serial from each other. The insertion of branch complements and the
number, beginning with the lowest. The graphs so constructed subtending branch sheaths at such congested nodes is distichous,
(Figs. 21-23) generally take the form of the profile of an asym- and the development of the nodes is normal in other respects as
metrical mountain, with a single acute or more or less rounded,. weil. The congestion of nodes (suppression of internodes) also
crest, or with two or more, usually less well-marked, peaks in the occurs, in conjunction with a greatly elongated first internode, in
median region. It will usually be steeper on the "west" side (as a ~ a number of otherspecies' of this and other bamboo genera of
result of larger length increments in successive internodes in the the Western Hemisphere, including Glaziophyton and Myriocladus.
lower part of the culm) and more gently sloped on the "east" side. Since these elongated first internodes, followed by congested nodes,
In scandent bamboos, of which apparently there are no culm- appear regularly in all culms of the species of which they are
intemode measurements on record, the curve of internode length characteristic, it seems cIear that they are not teratic in nature,
would embrace a broad, nearly flat middle section, representing but have a stable genetic basis.
aseries of mid-culm internodes of more or less equal length. The shape of the component segments (nodes and internodes)
Schromburgk (1841) brought to light the occurrence of a greatly of bamboo culms varies from species to species, and in some cases
elongated first above-ground internode in culms of a bamboo now from genus to genus. Differences in the internodes include the
known as Arthrostylidium schromburgkii. In a culm 15 m tall the degree and extent of the depression (groove) immediately above
first internode may be 5 m long. Another feature of this plant the insertion of a bud or a branch complement. In all bamboos
(McClure 1964:2) is the occurrence, immediately above the long of the genus Phyllostachys, for example (Fig. 27) each culm inter-
 VEGETATIVE PHASE: THE MATURING PLANT 45

INTERNODE I-ENGIH
IN CENTIMETERS ceptibly above the level of the surrounding culm surface. In some
42. bamboos whose culms have prominent culm nodes ("supranodal
40 ridges"), buds at lower nodes are so deeply sunken that they do
not project above this ridge (Phyllostachys arcana; Sasa longiligulata).
It has been suggested (Titkenouchi 1932, trans., p. 56) that pressure
exerted by the bud upon the contiguous developing internode,
while both are closely confined by the enveloping sheath, is re-
sponsible for the channeling of the internode at and above the
insertion of a budo But this channeling is so diverse in its expres-
sion, and so completely lacking in some bamboos (known species
of Melocanna aitd Schizostachyum, for example) where such pressure
can scarcely be wholly absent, that it must have some other cause
22 as welL Unpublished studies by Frank Venning on the young
20 shoots ofbamboos of several genera indicate that there may be
/. fundamental differences in the manner in which the bud primordia
of the culm and its branches are initiated, and that these differ-
ences may be correlated with the absence, or the presence aod
/4
extent, of channeling. Titkenouchi (1931b:154) relates the deep
/2

10 II'ITERNOO!1: ll!:NGTH
IN ClINTIMI';TCItS

4 Culm fzeigMs: 6·7M

6J31'<
2

o 2 4 6 8.'0/2/4 /6 /"20222426 28
30 32 3f 363S
INTERNODE NUMSER.

Fig. 22. Pattern of increase and decrease in intemode length in two culms from
the same clump of Phyllostachys nidularia (P.1. 67399), based on measurements
made at the U.S.D.A. Barbour Lathrop Plant Introduction Garden, Savannah,
Georgia, by McClure and Ditmeyer, August 12, 1942.

o Z+6 10IZIf161820222426Z8303.t 363f.14042 464f.150~

node is grooved (sulcate) throughout its length above the insertion lNTJ!:RNODE /'lUMBER

of a bud or branch complement; in Shibataea the internodes are

only flattened. The internodes of the culm branches and of the Fig. 23. Pattern of incr:ease and decrease in internode length in six culms from
the same clump of Phyllostachys pubescens (P.1. 80034), based on measurements
rhizome in these genera show this same characteristic.
made at the U.S.D.A. Barbour Lathrop Plant Introduction Garden, Savannah,
The buds at culm nodes generally protrude more or less per- Georgia, by McClure and Ditmeyer, August 12, 1942.
46 THE BAMBOO PLANT

channeling throughout each culm internode in Phyllostaehys to the

simultaneous development of all of the internodes (and buds) of
a young shoot in this genus, while in those bamboos where chan-
neling does not continue throughout the internode, the internodes
and buds are said to develop successively and acropetally.
A characteristic of many bamboos is the appearance of a white
exudate on the surface of the culm internodes. This varies from
a barely perceptible bloom (similar in superficial appearance to
that familiarly seen on some kinds of plums and grapes) to a con-
spicuously abundant, fluffy, flourlike deposit that more or less
completely conceals the green surface of the internodes, as in
Lingnania ehungii (Fig. 24). The time of the appearance of the
exudate in relation to the development of the culm, and its tex-
ture, abundance, distribution, and persistence, are variables that
may, in conjunction with other internode characters, be useful
in the identification of some species of bamboos in the field. Kurz
(1876:252) mentions the "white-pruinous" culms of Dendroealamus
giganteus and cf an unidentified bamboo, "booloo idyooh,'.' and
the occurrence cf ('a white fugaceous meal" on the culms cf "boo-
100 kenneng," a yellow-culmed form of Bambusa vulgaris. The exu-
date collected from young culms of Lingnania (as Bambusa) ehungii
was analyzed at Harvard University by Chang (1938). One com-
ponent, comprising about 25 percent of the powder, was identified
as a triterpenoid ketone, identical with or closely resembling frie-
delin, a constituent of the wax of corks. Two similar compounds,
bambuselin land bambuselin II, were later described from the ,
same source (Chang 1941).
Among other conspicuous internode characters are the surface
texture, the combination and pattern of colors, and the type of
pubescence. The pigskinlike surface of the lower mid-culm inter-
nodes in Phyllostaehys makinoi and Ph. viridis; the papillate surface
of the lower and mid-culm internodes of Chimonobambusa quadrangu-
laris; the color pattern yellow-with-a-green-sulcus of the internodes Fig. 24. Lin;:,rmania chungz"i. Plants of this species are made conspicuous by a layer
of the whole culm in Phyllostaehys bambusoides cv. 'Castillon'; the of flourlike exudate that more or less completely conceals the green surface of
the internodes in culms of the current year's growth. aulms in plants of mature
velvetlike pubescence of the internodes in young culms of Ph. stature may lack buds and branches in the lower % or % of their height. "Gray
pubeseens; and the hard, rough, siliceous surface of the lower culm bloom on green bamboo culms adds an indescribable beauty to this forest
internodes, often clothed with pale appressed hairs, in species cathedral." Au Tsai, Kwangsi Province, China. Reproduced from McClure
1935:203.
of Sehizostaehyum, are examples. Kurz (1876:251-252) lists others.
48 WE BAMBOO PLANT VEGETATIVE PHASE: THE MATURlNG PLANT 49
Arber (1934:77, Fig. 27) describes and figures as "spinous hairs" and in many species of Chusquea, especially those in the seetion
the characteristic retrorse hooks-outgrowths of the epidermis- Rettbergia.
that appear on the slender upper internodes of young culms in a Ve!asquez and Santos (1931) published an excellent portrayal
climbing bamboo, Arthrostylidium multispicatum. of the culm anatomy of five tropical bamboos.
The culm nodes also show more or less strongly marked varia-
tion in appearance as between different species. An important
feature of culm nodes is the sheath scar-which marks the sheath- Branch buds at culm nodes
node of Hacke! (1887, trans., p. 3). The sheath scar is a trans- Branch buds emerge on alternate sides of the culm just above the
verse circumaxial offset in the surface of the culm, marking the sheath scar at successive nodes. Bach bud stands in a position
circumaxial locus of insertion of the culm sheath. The sheath median to the base of the sheath that subtends it. With the excep-
sear may be thin and inconspicuous, without any outgrowths, tion of those of the genus Chusquea, all bamboos studied have the
as in Melocanna, or it .may be thick, with a strongly fiared callus primary buds solitary at culm nodes, as in Lingnania chungii (Fig.
(see p. 67) and conspicuously fringed with brown hairs, as in Ling- 25, A). In Chusquea the principal bud at each culm node (above
nania chungii (Fig. 25, A), Phyllostachys nidularia, and Sinobambusa ground) is fianked by two to many smaller ones. These represent
tootsik. The sheath scar may be symmetrical and level, or it may independent branch primordia, each enclosed in aseparate pro-
take a strong dip on the side that passes under the point of inser- phyllum (Fig. 25, B). In exceptional cases (sometimes, for example,
tion of the branch complement, as it does in Bambusa arundinacea in Ch. pittierz) the small buds emerge in a continuous narrow band
that extends complete!y around the culm. The unusual nature of
the branch buds, and the course of deve!opment of the branch
complement, shown in Merostachys and in some species of Arthro-
stylidium will be described be!ow (pp. 53ft).
Takenouchi (1931b:143) was the first to call attention to the
\\ order of breaking of branch buds on the young culm. In some
species these buds awaken in acropetal order, that is, each bud
breaks as soon as the internode to which it is basal has tompleted
its growth; examples are typical Arundinaria gigantea and all species
of Phyllostachys. In certai:iJ. other species the order is basipetal.
" Here, some or all of the branch bucjs of the full-grown young

i"l culm remain dormant. for a while (sometimes for several weeks,
as in Bambusa textilis) before breaking. In. small plants of Semi-
arundinaria viridis the mid-culm buds break first, and the wave of
branch growth. then spreads simultaneously upward and down-
ward in the culm. In some bamboos, and under' c~rtain conditions,
A B c both the awakening of the primary branch buds of the culm and
Fig. 25. CA) Mid-culm node of Li~anant'a chungi~ showing sheath scar with COll- their subsequent proliferation may be irregular both in time and
spicuous callus and fringe of hairs and typical solitary branch bud; far an in order of incidence. In mature bamboo plants one or more of
illustration of the branch complement of this species, see Fig. 26, 4. (R, C) Mid-
culrn node of Chusquea sp., showing eR) multiple-bud (constellate) complement
the lowest above-ground buds on each culm may remain dormant
peculiar to this genus, and (C) branch complement; both somewhat diagrammatic; indefinitely. In most bamboos, immature plants may bear buds
not drawn to scale. at all of the culm nodes. In many bamboos, culms of progres-
 VEGETATIVE PHASE: THE MATURING PLANT 51
2

sively larger sizes (those approaching mature stature) have a

1 progressively longer series of budless lower (above-ground) nodes.
In Bambusa textilis and Arundinaria amabilis, for example, culms
from plants of mature size may lack buds and branches in the
lower %to ~h or even 0/4 of their length. This 18 a desirable aes-
thetic feature in bamboos planted for ornament--especially such
species as Lingnania chungii (Fig. 24). In culms used in handicrafts
or in industry, the absence of the knots that mark the insertion
of branch complements malms for economy and ease in working
the materiaL In the sterile condition, culms commonly remain
unbranched above the ground in bamboos of the genera Glazio-
phyton, Guaduella, Neurolepis, and Puelia. The incidence of buds at
subterranean nodes of culms arising from lateral buds of lepto-
12 morph rhizomes may differ from species to species in the same
genus (Phyllostachys), or within the same species as the plant ad-
vances toward maturity (Arundinaria amabilis).

b - - - e.g.
The branch complement
In bamboos whose cuhns are branched (the condition prevailing
in most genera) it is at the mid-culm nodes that the typical form
of the branch complement appears. The lowermost one or more
branch complements in the series on a given culm are generally
less weil developed than those at mid-culm nodes, and so cannot
.be taken as typical examples. In most bamboos the branching
at the uppermost culm nodes is usually not strongly distinctive
of a genus or species. However, bamboos with a fixed number of
branches at each culm node, as in Sasa and Phyllostachys, are
exceptional in this respect.
In a few bamboo genera, the number ofaxes in a mid-culm
branch complement is fixed at a characteristic total. In Sasa, the
single primary branch remains solitary because it possesses no
basal buds. In Phyllostachys, the typical primary mid-culm branch
"
complement is binary, the two axes being more or less strongly
, 1.:'
unequal (subequal in Ph. arcana), and subsidiary branching is
Fig. 26. Lingnania chungii. Analysis of parts to illustrate the original de- limited to a single, very much smaller axis that sometimes emerges
scription of tpe species: 1, 2, culm sheath; 3, rhizomeand base of culm'
4, mid-culm branch complement; 5, cross section of culm; 6, two pseudo~
between them from a bud at the base of the smaller of the two
spikelets; 7, diagram of two pseudospikelets (g. b., bract subtending a bud; (Fig. 27).
pr., prophyllum; e.g., empty glume; (, perfect floret); 8, floret; 9, lemma; In many bamboos the branch primordium at each mid-culm
10, palea; 11, -lodicules showing variation in shape; 12, anthers, showing node ramifies precociously before it ruptures the prophyllum. The
variation in the apex; 13, gynoecium. Drawn by Chan Hin-yau.
 VEGETATIVE PHASE: THE MATURING PLANT 53

generic name Pleioblastus (Nakai 1925:145-146) was based On this

character. However, Nakai's description may leave the incorrect
impression that the bamboos included by him in this genus actu-
ally have more than one primary bud at each culm node. The
extent cf this precocious ramification cf the original branch pri-
mordium varies from species to species and from genus to genus.
Usui (1957 b) illustrates the phenomenon in several 'of its mani-
festations (Fig. 28). In Melocanna baccifera, and the kitown species
of Schizostachyum, . the degree of precociousbranching is very pro-
nounced, and the branch complement emerges from the prophyl-
lum a sunburst of juvenile a~es. In these genera, the primary
branch is indistinguishable in size from the subsequent orders
ofaxes, and the result is a dense tuft of slender subequal branches.
In these and other bamboos, the growing out of buds at proximal
nodes of branches of successively higher orders augments still
further the number of component axes in a given branch com-
plement. The tempo of this augmentation, and the length of time
during which it continues, determine the composition and nature
of the mid-culm branch complement found in any culm of a
given age. In many bamboo genera (Bambusa, Dendrocalamus,
Sheoth scor-~:::::::}Supro-noda 1-1~§l,,'if" Gigantochloa) the primary branch emerges, and remains, strongly
ridge dominant. Subsequent orders ofaxes that develop from its basal
buds have progressively diminished diameters and lengths (Fig. 30).
In bamboos of the genus Chusquea, the branch that arises from
the large central bud at each mid-culm node is strongly domi-
nant-sometimes nearly equal to the mother culm in diameter
and length-and is often tardily or not at all rebranched at its
basal nodes. However, this large central bud often remains dor-
mant, and may become completely obscured by the dense cluster
of slender, subequal branches that develop from the small adventi-
tious buds at the same culm node (Fig. 25, B).
Fig. 27. Brauch coroplement from a mid-culm node of Phyllostachys elegans The branch complement in bamboos of the genus Merostachys
(P.I. 128778). The featur~s shown characterize the genus in the following and in some species of Arthrostylidium bears a superficial resem-
r~spects: the .branch-beanng nodes of the culm are marked by a prominent
ndge (sometl:nes referred to by Hayata as a pulvinus, and by others as a blance to that in Schizostach:yum and Melocanna, in that it consists
supranodal ndge) above the sheath scar; the internodes of the culm are of a large number of slender, subequal leafy axes that usually
flattened throughout by grooves above the point of insertion of each bud branch from their basal buds only. Gloser examination, however,
or branch complement (all brauches are similarly flattened); and the pri-
mary branches are borne typically in twas at each node hut sometimes a
reveals a basic difference. In Schizostachyum and Melocanna, the
third ~ne develops between the other two (upper right). From McClure branches at mid-culm nodes radiate from a common point, the
1957:Flg. 2. See p. 13, and Sheath node and Culm node in the Glossary. locus of insertion of the primary branch primordium. In Mero-
 VEGETATNE PHASE: THE MATURlNG PLANT 55

stachys (Fig. 31) and in some species of Arthrostylidium (A. harmon-

icum, Fig. 32), the focus of radiation of the mid-culm branches·
falls within a vacant area. This hitherto undescribed character
has its origin in the peculiar nature of the branch bud, the stage
of development of the internode. when the bud is formed, and the
behavior of the primary branch primordium (primordia?) as
p subsequent development takes place: The material" at hand for

A B c

t i

D
M.A><.

Fig. 28. Diagrammatic cross sections of four types of mid-culm branch buds and
diagrams of corresponding branch complements, presented by Usui (1957:'Figs.
1-4) as typical of bamboos of six genera. CA) The genus Saso,' here the primordial
branch axis re.m~ins solitary, both befare and after the rupture of the prophyllum
t~at enclos~d l~ l~ the bud stage. (B) Represents three genera: Pleioblastus (merged
wlth Anmdmana In the present work), Semiarundinana, and Sirwbambusa; the broken
lines-ostensibly represent branches produced after the rupture of the prophyllum. D E F
(C) Th: genus Sht'bataea. CD) The genus Plryllostaclrys; here the branches emerge
as a palI' of subequal axes, the basal bud on the smaller one of which may remain Fig. 29. Examples of diverse ·degrees of basal proliferation in the mid-culm
dormant, er develop as shown here, to form a third, very rouch smaller one branch complement in representatives of several genera: CA) Sasa palmata; (E)
between the other two. The captions given above are somewhat modified from Phyllostac~s dulcis; CC) Shibataea kumasasa,' (D) Aru:n.dinaria tecta var. decidua; (E)
the originals ..The author's leß"end: "P, prophyll; 1st, the &st foliage leaf; 2ru4 Arundinaria (Pleioblastus) simonii; CF) Semiarundt'na:riafastuosa.
the second fohage leaf [on a glven axis]; M. Ax.) main a.us; L, a leaf on the node
of main axis; the black leaf shows the prophyll and the white leaf shows the
foliage leaf. Branch intemodes not drawn to scale." See Figs. 27_and 29.
 VEGETATIVE PHASE: THE MATURING PLANT 57

Fig. 30. Branch complement of Bambusa rutilC4 illustrated from a mid-culm node
of a 3-year-old culm in the Lingnan Dniversity Bamboo Garden (LDBG 1476)
showing a strongly dominant prirnary branch and thornlike branches of higher
orders. Drawn by M. K. Hoh, 1939.
Fig. 32. Arthrosf)llidium harmonicurn. Basal portion of a fully developed mid-culm
branch complement, showing the relation of the component branches to each
other at the locus of their emergence. Based on a specimen in the U.s. National
Herbarium, collected in Ecuador by W. H. Camp (No. E-1613). Drawn by EImer
W. Smith. See Figs" 31 and 33.

study is i~adequate to a complete elucidation of the development

of this type of branch complement. The earliest stage available
to me is found in a sterile specimen of Merostachys sp. accessioned
in the National Herbarium under US 562138 (Agnes Chase 9458).
A slightly later stage is shown in US 1257740 (Agnes Chase 9466).
A solitary two-keeled prophyllum encloses the primary branch
primordium (primordia?). Several primordia appear later in alter-
nate, distichous insertion (emergence), enclosed individually in
Fig. 31. Diagrammatically reconstructed early stages in the development of a small, bladeless sheaths. The spatial pattern of their insertion
mid-culm branch complement typical of the genus Merostachys~ based principally "suggests separate origins, though this can only be determined by
on a specimen in the D.S. National Herbarium, collected by Agnes Chase (No.
9463). The development of the branch complement in Arthrosf)llid~ian harmonicum
(Figs. 32 and"33) is similar.
58 THE BAMBOO PLANT

anatomical studies. In the early stages, the bud lies within a few
millimeters of tbe level of insertion of the culm sheath by which
it is subtended, and the surrounding culm tissue is_ still in a meri-
stematic condition. When the branch complement has reached
its full development, however, the spot originally occupied by the
bud is vacant, and the primary branches are seen to be inserted
in an arch above its periphery. The fragmentary evidence at hand
suggests that, by tbe elongation of the culm internode, the several
branch primordia are carried forward acropetally and fanned out
along tbe surface of the culm. The branches arising from these
primordia finally emerge at separate loci arranged in a tight arch.
To characterize branch complements with this type of insertion
the term apsidate is proposed. The manner of origin of branch
complements of tbis type remains to be elucidated by anatomical
studies initiated at early stages of their formation.
Branch complements originating in this manner generally COID-
prise numerous slender, subequal axes. In Arthrostylidium harmonicum
(Figs. 32 and 33) the pattern of development appears to be basic~
ally the same as tbat typical in most of tbe known species of Mero-
stachys. Exceptionally, however, one or two of the axes at tbe top
of the arch of insertions may elongate and branch from buds other
than their basal ones. This has been observed frequently in Arthro-
stylidium capillifolium and A. cubense, and once in a sterile leafY
specimen of an unidentified species of Merostachys (US 1910797;
Froes 20060, sheet 2).
In branch complements where tbe primary axis is dominant,
this branch resembles tbe culm itself in varying degrees, and usu-
ally on a very reduced scale. This resemblance is especially notice,
able in many bamboos tbat have pachymorph rhizomes. Here
the base of the prirriary branch at each mid-culm node incorpo-
rates a structure somewhat resembling the pachymorph rhizome
(Figs. 25, C and 30). The internodes of tbis basal part of the
branch are solid, and often bear raots Gr root primordia, as in
Bambusa tulda and B. textilis (Fig. 97), B. vulgaris, and Gigantochloa
apus, for example. The transition to hollow, elongated internodes
is as abrupt here as is tbe transition from rhizome to culm that Fig. 33. Arthrosrylidium harmonicum. Lower and mid-culm nodes, showing early
stages in the development of the absidate, pleioclade branch complement. Docu-
is typical in bamboos with pachymorph rhizomes (Fig. 20, lift). mented by specimens in the D.S. National Herbarium collected in Ecuador by
In some bamboos with scandent culms (Chusquea and Dinochloa) McClure (No. 21416). U.S.DA photo by Jarnes Mitche1L See Figs. 31 and 32.
tbe principal branch at lower or mid-culni nodes may approach
 60 THE BAMBOO PLANT VEGETATIVE PHASE: THE MATURING PLANT 61

the culm itself in diameter and length as weil as form. In some Again, Bambusa dissimulator shows an inconspicuous but appreciably
bamboos with pachymorph rhizomes, the resemblance of tbe pri- more pronounced thorniness, and in B. rutila (Fig. 30) tbe spines
_mary branches to tbe culm is much less spectacular. This is par- are still more strongly developed. Referred to by Arber (1934:77)
ticularly true where the primary branch is not appreciably larger as "spinescent buds, which are eitber dormant or permanently
than the secondary ones at mid-culm nodes, as in- Lingnania arrested" (cf. Kurz 1876:236), these thorns actually show all of
chungii (Fig. 26, 4). In Melocanna baccifera and Schizostachyum spp., the characteristicmarks of segmented axes. The nodes bear simple,
tbe branch complement is made up of numerous, widely radiat- scalelike sheaths, and these sheatbs often subtend buds. The buds
.ing, smalI, slender, subequal axes. Such branches generally do not on primary thorns sometimes produce other, smaller thorns, cr
develop root primordia. However, an exceptional example of tbe leafy twigs, or even infiorescences. Seibert (1947:272) refers to bam-
spontanious production of roots from the basal nodes of such baD branch thorns as "short shoots."
branches, in sitv, is shown in a sterile specimen of Schizostachyum
sp. collected by N. W. Simmonds (s.n.) in Western Samoa, October The sheathing organs
13, 1954, and preserved at tbe Kew Herbarium.
Every node of every segmented vegetative axis of a bamboo plant
In self-supporting culms the pattern of the lengtb of tbe primary
bears a sheatbing organ, which embraces tbe developing internode(s)
branches at successive nodes, from the bottom to tbe top of tbe
distal to its insertion. This sheath is often referred to loosely as a
series, follows the pattern of culm-internode lengtb. Of tbis cor-
leaf. The sheatb proper should, however, be distinguished from
respondence, Porterfield (1933:362) says: "The relation between
its blade, which is the only leafiike part. In the bamboos, a foliage
the lengtb of [culm1 internodes and branch length is significant,
leaf (a leaf proper) is an appendage to a leaf sheath proper (cf.
and strongly suggests a connection with functions of growth."
Leaf and Sheath in the Glossary). Sheath primordia are the first
However, certain scandent bamboos show quite a different pattern.
appendages to emerge behind tbe apex of tbe growing point of a
In Chusquea simpliciflora and Melocalamus compactiflorus, for example,
segmented axis (Porterfield 1930a:Fig. 1). Next to emerge is the
tbe principal branch at mid-culm nodes may approach the culm
primordium of the bud subtended by each sheatb. Greatly in need
itself in diameter and lengtb, but tbe internodes of tbe mother
of further study is the question of the possible infiuence of sub-
culm where these branches originate are not correspondingly
stances elaborated in the developing sheath upon the initiation of
elongated.
bud primordia and upon ensuing events. These later events in-
In some bamboos, certain branches of tbe several orders become
clude tbe development of segmentation in the elongating axis, the
dwarfed, indurated, sharp-pointed, and curved, in degrees that
formation and transverse anastomosis of fibrovascular bundles,
generally correspond to the species. Extreme degrees of these
intercalary growth, and even the initiation of root primordia,
modifications result in a IDeal or more or less general manifes-
each of which takes place only at or just above anode (see pp. 12
tation of thorniness in the plant. Branches in the lower part of
and 123).
the culm are usually the most thorny ones. All of the khown
Witb reference to the relation between a bud and its axillant
species of Guadua, and a uumber in tbe genus Bambusa have thorny
basal branches. Bambusa arundinacea is a classic example. The de- leaf, Arber says (1950:125):
gree of thorniness may vary within a species. For example, in its
It is generally agreed that the vast majority of buds arise in connec-
most northerly distribution, the typically thorny Guadua amplexi- tion with an 'axillant' leaf; but to what, e..-..::actly, this amounts remains
folia is represented by a thornless form. Intermediate degrees of vague, and Warming's criticism, though made in the eighteen-seventies,
thorniness characterize other farms cf this species. In some cases, is still pertinent. He wrote [in French]: "It appears that no one has clari-
the extent to which branches are invested witb thornlikeness is lied the relation between a bud and the leaf called mother-leaf [axillant
so slight that the character may be overlooked (Bambusa malingensis). leafJ·"
62 THE EAMBOO PLANT

Porterfield (1930a) and Sharman (1945) trace the anatomical

development ofaxillary buds and of the corresponding axillant
(subtending) "leaves" in a bamboo and in several nonbambusoid
members of the Gramineae, respectively. Garrison (1949) and
others have made developmental studies of the anatomical relation
between bud and axillant leaf in several species of woody dicoty-
ledonous plants. However, it appears that the respective origins
of the two associated elasses of structures, and their relation to
each other, still await effective illumination from the point of view
of physiology and biochemistty.
The term "sheath" is commonly used in a comprehensive sense,
to inelude both the sheath proper and its appendage~. In the bam-
boos the principal appendages of the sheath proper are the blade,
the ligule, and the aurieles (Fig. 35). These appendages may
assurne very different sizes and forms in the sheaths of different
parts of the plant. The bJade, the ligule, or both may be very'
small or even rudimentary, and the aurieles may be lacking en-
tirely, as in the sheaths of therhizome, for example (Fig. 34).
The sheath proper is the basal, and basic, part of the structure.
Ey their elose, firm application to the axis that bears them, and Fig. 34. Rhizome sheaths, showing typical extreme reduction of the sheath
by their form and structure, the sheaths proper are specially blade, illustrated by Takenouchi (1931a:Fig. 2): 1~ Semiarundina:ria viridis~' 2, Sasa
fitted to perform the function of protecting, supporting, and (as Pseudosasa) kurilensis var. nebulosa," 3, Arundinaria (as Pleioblastus) variegata var.
viridis f. major; and 4, Phyllos~achys hambusoides (as Ph. reticulata) var. marliacea. See
stiffening the intemodes of the young aerial axes during the period'
pp. 75f.
while these are actively elongating. Without this support during
their critical period, the tender meristematic tissues of the inter-
calary zones of a growing culm or branch would be incapable of shape, and texture, and (2) the relative, development of the blade
resisting the transverse mechanical forces that impinge upon them. and other appendages, constitutes an important' basis for differ-
In the rhizome, however, the most obvious ftmction of the sheaths entiating the bamboos from the rest of the Gramineae. The dimor-
proper is the formation of a sharp-pointed resistant shield that phism between the culm sheath and the leaf sheath is especially
protects the tender apical meristem as' it is pushed through the conspicuous in the bamboos (Fig. 35).
soil by the elongation of the rhizome axis. All bamboo sheaths Tax.onomists and anatomists frequently fail to identify for the
have one character in common: the smooth, lustrous inner (adaxial) reader the axis, or the point of origin, of the sheath whose features
surface of the sheath proper. This feature facilitates the adjust- are being described. Too often this structure is referred to indis-
ments made necessary by the rapid elongation of the young inter- criminately either as a "leaf" or a "sheath." For elarity, the follow-
nodes while still tightly embraced by the sheaths. The white exu- ing types of sheath found in the bamboo plant should be differ-
date mentioned earlier (p. 46) may ftmction here as a lubricant. entiated: culm sheaths, branch sheaths, leaf sheaths, rhizome
The strong disparity that prevails between the sheaths found sheaths, neck sheaths, and prophylla. The last-named are often
on different axes of a bamboo plant, in respect to (1) their size, referred to ambiguously as bracts, or bracteoles, or bud scales. '
 64 TIfE BAMBOO PLANT 2

-Petiole

Sheoth proper

Leof sheoth--+

Culm sheoth Folioge leof

Fig. 35. Diagrammatic representation of culm sheath ~d foliage leaf sheath in
the.genus. Bambusa, showing the several component parts; not drawn to a uni-
form scale. The dimorphism that prevails as between these two. types of sheath
is generally much more marked in the Bambuseae than in the other tribes of
the Gramineae. Oral setae are sometimes referred to as shoulder bristles, when
auricles are absent.

Culm sheaths
Fig. 36. Culm sheaths of four congeneric species: 1~ Bambusa tuldoides~ LUBG
The series of sheaths that clothe the successive internodes of a 2664; 2, B. textilis, LUBG 1842; 3, B. pervariabilis, LUBG 1226; and 4, B. tulda,
developing bamboo culm show a progressive change, acropetally, LUBG 1350; all [rom the mid-culm level. As a ruJe, differences between species
in the size, shape, substance, and vesture of both the sheath proper are shown by the culm sheaths more dramatically in bamboos of large stature,
such as these, than in very small species. As demonstrated here, the aurides often
and its appendages. A given culm sheath is therefore in these
contribute useful characters. The main drawing in each case shows the outer
respects characteristic of the position of its origin. In a culm of (aba.xial) aspect of the whole sheath; the insert shows the inner (adaxial) ~pect
mature stature the culm sheaths at nodes in the mid-culm range of the apical por#on, including the ligule at the junction of the blade with the
show the development of their various parts and appendages that sheath proper. The original drawings (here reproduced at about '15 natural size)
where made.by Miss Y. M. Lau, from specimens grown at the Lingnan Uni-
is most highly characteristic of a given species (Fig. 36). In sheaths versity Baniboo Garden, under the accession numbers indicated.
66 THE BAMBOO PLANT VEGETATIVE PHASE: THE MATURING PLANT 67
from near the base and tip of a culm, however, the features typical In a given species, the culm sheaths may be persistent, tardily
of the species are as a rule somewhat obscured by a kind of gener- deciduous, or promptly deciduous. In some species (Guadua amplex-
alization of their expression. At the base of the culm the sheaths ifolia) the sheaths at the lower culm nodes may be persistent while
approach in form those of the rhizome or the neck. At the upper- those above are promptly deciduous. In Semiarundinaria fastuosa
most several nodes the sheath and its blade, by a rather sudden abscission is incomplete, and a culm sheath usually hangs for a
change of proportions, take the form of leaf sheath and petiolate time attached by a narrow sector of tissue at the middle of its
leaf blade. The progressive change in the form and dimensions of base. In bamboos of the genus Phyllostachys the culm sheaths are
sheath proper and sheath blade ·characteristic of aseries of sheaths promptly and completely deciduous; in Sasa they are persistent.
from successive nodes of a single culm of each of two species is The incidence of deciduousness may vary from species to species
shown in Fig. 37. within a given genus, as in Arundinana. As a rule, the deve10pment
of a branch complement dislodges the subtending sheath partially
or completely. Where the culm sheaths are stubbornly persistent,
and particularly where they are closed, as in some species of Chus-
quea, by having the outer edge fused to the outer surface of the
sheath, the emergence of the branch complement ruptures the
sheath locally (Fig. 38).
In manl' bamboos, such as Phyllostachys nidularia, Sinobambusa
tootsik and Lingnania chungii (Fig. 25, A), for example, we find at
the base of each culm sheath in the lower and midculm range a
ring of cor1:y tissue properly called a callus or sheath callus (see
p. 48). U pon the abscission of a sheath most of this tissue remains
attached to the sheath node. Arber (1934:309) attributes to Godron
(1880) the description of a similar structure in the leaf sheath of
Chimonobambusa (as Arundinaria) falcata, and proposed for it the
term collaret. Actually, it is rather at the base of the culm sheaths,
and of the branch sheaths that are proximal to the leaf sheaths, that
this structure is conspicuously developed in Chimonobambusafalcata.
In. Melocalamus compactiflorus and in some scandent species of
Chusquea (Fig. 38) the base of each mid-culm sheath consists of,
or is inserted upon, a conspicuous horizontal band of tissue distinct
in structure from that of the rest of the sheath proper; and prob-
lil iirll i/liI ably related to the callus anatomically. It is e>'l'ansible in all di-
rections as long as the corresponding intercalary zone of the
)1\ '\ il 1
1" 11
I\~ embraced internode is in active growth, and it "takes on an asym-
metrical form at geniculate nodes. Upon the abscission of the
sheath, this band remains attached to the sheath node. Gamble
Fig. 37. The progressive change in form and dimensions 'Of sheath proper and
sheath blade that characterizes the series of sheaths [rom successive nodes of a
(1896:111 and PI. 98, Fig. 10) characterizes this structure as "a
single culm, exemplified by Pseudosasa (as Yadakeya) japonica (above); and Bambusa broad leathery ring left by the sheath." The term sheath girdle
pachinensis (below); all much reduced. Redrawn [rom Takenouchi 1931a:Fig. 16.' seems eminently suitable for it.
 VEGETATIVE PHASE: THE MATURING PLANT 69

Culm Sheath Blade. A culm sheath blade is sessile, and may be

either persistent, incompletely abscissile, or completely abscissile. 2
. In the lowermost sheaths the blade is usually "reduced" to a
Cu Im shealh blade sturdy mucro and· the rest of the appendages are lacking or very
rudimentary.

Branch sheaths
Young branches The series of sheaths clothing each branch of the culm recapit-
ulates, on a reduced scale, that clothing the culm. The dimensions
are progressively reduced, and the form of the sheaths is progres-
sively generalized, in the successive orders of branches. In some
of culm shealh
bamboos with scandent culms, where the principal branch at mid-
culm nodes may approach the culm itself in diameter and length,
its sheaths may be indistinguishable for those of the culm proper
(Chusquea simpliciflora; Melocalamus compactiflorus). Branch sheaths
may be persistent or deciduous. The blade of a mid-branch sheath
may be persistent but more often it soon abscisses. In sheaths at
the basal nodes of a branch the blade is usually represented by a
short-mucro.

Shealh proper, closed

al base by fusion Leaf sheaths
The sheaths that clothe the distal nodes of all culms, and of
branches of all orders, are properly called leaf sheaths. Their
blades, called leafblades, or simply leaves; are the plant's principal'
source of elaborated food. The sheath of a foliage leaf is distin-
guished from other kinds of sheaths by having a petiolate (stalked)
blade, and by being relatively small and inconspicuous in relation
to its blade (Fig. 35). Each one being much longer than the inter-
. node it embraces, leaf sheaths are deeply imbricate, that is, over-
lapping, each one nearly covering the one above it, except in rare
cases where, under weak illumination, the axis bearing them is
atypically elongated. Takenouchi illustrates the variation in auri-
cles and ligules of leaf sheaths as between several Japanese bam-
Fig. 38. Lower mid-culm nodes of yaung" culm of Chusquea sp.~ showing the well- boos in different genera (Fig. 39).
developed "girdle" of the culm sheaths. The lower sheath shows the fusion of
the outer margin to the sheath itself. This closure results in the persistence. of 2A new feature- of vascular anatomy, in the fonn cf transverse bars that take the
the sheath, a condition related to the extravaginal eruption of the branch COID- place cf the end walls of the component cells of the annular vessels in a "sheath leaf'
plement shown ahove. Field sketch, documented by herbarium specimens (culm sheath blade) from a young culm shoot of PhylLostachys nidularz"a is described and
(McClure No. 21440) collected in Ecuador. illustrated by Porterfield (1923).
 VEGETATIVE PHASE: THE MATURING PLANT 71

(Lat., collum) 5 mm long, and characterizes the ligule proper (the

ligula interior of Hackel) in the same species as "lanceolate" and
5-7 cm long.

The Leaf Blade. Leaf blades (see Leaf in the Glossary) differ from
culm sheath blades and branch sheath blades in being stalked or
petiolate. Leaf blades are generally thinner than ~ulm sheath
blades, and often show more marked dorsiventrality. In many
bamboos the petiole has a pair of rather strongly deve!oped pul-
vini at its base (Dendrocalamus latijlorus; Indosasa gibbosa). By means
of these, the petiole orients the blade in response to the dominant
incidence of light, but the mechanism of the functioning of pul-
vini has not been studied in the bamboos.
The order of development of leaf blades is acropetal. They
emerge, singly, through the "mouth" of the preceding leaf sheath.
As it emerges, each one is rolled up tightly along its long axis
(Fig. 40, lift). Brandis (1907:72) makes reference to the convolute
J
Fig. 39. Diverse forms assumed by the ligule and auricles cf the bamboo leaf
sheath, as exemplified by: 1, 2, Bambusa shimada~' 3, 4, Sasa tsuboiana; 5, 6, Arundi-
naria (as Pleioblastus) variegata; 7, 8, Phyllostachys pubescens (as Ph. edulis); 9, 10,
Phyllostachys formosana; 11, 12, Shibataea kumasas~' all dravro at same scale. From .
•e
...
~l .
", ••
h.
Takenouchi 1931a:Fig. 15.
•• 0-1
~
The Ligule. The ligule, in the conventional sense of the term, is an ,.,e ,•
upward extension of a sheath proper, behind and beyond the' boO
•'.'•,•
0'
locus of insertion of the sheath blade. The ligule is to be recog-
nized by its position rather than by a common shape or texture.
In size, texture, and substance, the bamboo ligule ranges from
. oe

•• ,
,
(r'••, •0

0
.
tr
•·....-s
00
•
"

a smalI, thin, and scarious structure similar to that commonly

~. • ,', .'0 d
found in the herbaceous grasses (see Jackson 1949:215) to a struc-
" 0' · e,::: • G
ture of substantial size and thickness. The "leaf scar" left by the 'd.' 0,
abscission of the leaf stalk (petiole) is bounded on its adaxial side' .! '" "
A

by the conventionalligule, and is surrounded on the outer side by

an upward extension of the leaf sheath. The two structures com- Fig. 40. (Lift) Cross section of a developing leaf blade of Phyllostachys pubescens,
bined make a little cup, referred to earlier (McClure 1941 :33-34) showing the convolute vernation characteristic of the foliage leaves of most bam-
as the "cupule" (Lat., pocillum). Hacke! (1899:716 and e!sewhere) , boas; greatly enlarged. From Takenouchi 1931a:Fig. 12 (as Ph. edidis). (Right)
refers 'to the outer rim as an "external ligule" (Lat., ligula externa Diagrammatic magnification of features of the lower (abaxial) surface of a leaf
of Bambusa dolichocla~ showing: (f{) short spinelike hairs; (St) lang spinelike hairs;
and ligula exterior), a designation I have occasionally used. Holtturn (S) stomata; (G) vascular bundles, which appear as longitudinal veins. The
(1958:15, Fig. 3) refers to this structure as a "callus." In describ- transverse veinlets are here not visible, being submerged in the mesophyll tissue.
ing Guadua perligulata, Pilger (1937:57) refers to it as a "collar" From Takenouchi 1931a:Fig. 17. '
72 THE BAMBOO PLANT

vernation of bamboo leaf blades .. Chusquea pinifolia, which Brandis

says may possibly be an exception, conforms in this respect.
The foliage leaves of all bamboos have more or less clearly man-
ifest parallel veins ofthree orders: primary (the midrib), secondary,
and tertiary (in descending order of size). In those bamboos whose
leaf anatomy has been adequately studied, mutually adjacent
veins of all orders are connected with each other by transverse 00

~
vein!ets (Fig. 41). In some bamboos (principally the hardy species),
"""
-
2
these transverse vein!ets may be clearly seen to form, with the
parallel veins, a pattern conventionally called tessellation, cor tes-
sella te venation. In the leaves of most of the physiologically
tender, so-called "tropical," species, on the other hand, tesse1jlation
.....
~
is not at all or, generally, only barely detectable externally. Mit- i'"'
ford (1896:55) calls attention to the remarkably wide incidence, - i""
amon« the bamboos, of a correlation between cold-hardiness, in r--
the plant and' strongly marked tessellate venation in the leaves.
Wisely refraining from drawing a conclusion with respect to all
I'- ...... '"'
1 3
bamboos, Mitford terminates his observations with these words.
Fig. 41. Patterns made by transverse veinlets with the longitudinal ones, as e.x-
1t would be, of course, idle to assert that every bamboo which has emplified in leaf blades of: 1; Bambusa stcnostachya>' 2; Phyllostachys makinoi; 3, Sasa
tessellated leaf veins is hardy; indeed we know that there are many bam- veitchii (as S. albo-margi'nata); all greatly enlarged. From Takenouchi 1931a:Fig,
boos with tessellated venation which, from their habitat, cannat be 20. The transverse veinlets are usually not manifest to superficial examination in
grown in this country [England]. Only one thing is certain, vi~., that no unskeletonized leaves of the genus Bambusa and most other tropical genera with
bamboo introduced up to the present has proved hardy that has not pachymorph rhizomes.
such tessellation.
that are found, usually in unequal abundance, on the two edges
Comparative anatomical studies should reveal structural dif- of each leaf blade in most bamboos. These and other microscopic
ferences that explain the strong diversity of external expressi~n features of the leaf surface, which he illustrated diagrammatically:,
that characterizes the transverse venation in bamboo leaves of dif- are reproduced in Fig. 40, right. Ohki (1932) made a systematic
ferent genera. Brandis (190'i:PI. 12, Fig. 18) presents clear evi- study of these features of the epidermis in the common bamboos
dence that in the tropical bamboo Dendrocalamus gzganteus the of Japan by means of spodograms of the leaf blade, a method
transverse veinlets may be obscured from superficial view by thick first demonstrated by Molisch (1920:262ff, PI. 2, Fig. 9). Ohki
layers of tissue both above and below them, in addition to the (p. 4) recognizes eight "types" of spodograms: "1. the Shibataea
epidermal layer. On the other hand, in the hardy bamboo Pseu- type; 2. the Dendrocalamus type; 3. the Bambusa type; 4. the Sasa
dosasa japonica (ibid.: PI. 11, Fig. 3, as Arundinaria) the transverse type; 5. the Semiarundinaria type; 6. the Sinobambusa type; 7. the
veinlets impinge upon the epidermis on both surfaces of the leaf Pleioblastus type; and 8. the Phyllostachys type." His analytical key
blade. Such structural differences are not often brought into focus (p. 3) to the J apanese genera of bamboos, based on characters
where the leaves being compared are sectioned only in a plane at drawn from his leaf spodograms, folIows:
right angles to the long axis (see Prat 1936:Fig. 21, A and Met-
Silica cells are not found in the epidermis above the assimilation
calfe 1956:Figs. land II). . 1 tissue on the under surface of a leaf . . . . . . 1. Shibataea Makino
Takenouchi (1931a:Fig. 17) illustrates variations in the size, Silica cells are found in the epidermis above the assimilation tissue
shape, and distribution of the short, spinelike, antrorse processes 1 on the under surface of a leaf. . . . . . . . . . . . . . . . . . . . . . . .. 2
 VEGETATNE PHASE: THE MATURING PLANT 75
74 THE BAMBOO PLANT
their spodograms that ~ have. fOU1;d it extremely difficult to distinguish
Silica corpuscles (Kieselkörper) are faund on either cr both sides of
one from the other. Th1S, I th1nk, lS most likely due to the fact that they
the longer rectangular cells of a leaf. . . . . . . . . . . . . . . . . . .. .
came from the same phylogenetic branch.
2 ...................... 2. Dendrocalamus Nees ab Esenbeck
Silica corpuscles are hardly ever found on either cr both sides of the The results of Ohki's survey suggest that the epidermis of the
\ longer rectangular ceHs of a leaf. . . . . . _ . . _ . . . . . . . . . . . . . 3
leaf blade in some bamboos offers characters which, when used in
Silica cells in the upper epidermis above the veins are comparatively
short, and are nearly dUr»b-bell shaped cr rectangular in form conjunction with other morphological features, may be of diagnostic
3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3. Bambusa .schreber value for the recognition of genus or speeies (see Prat 1936;
Silica cells in the upper epidermis above the veins are nearly rec- Jacques-Felix 1955; Metcalfe 1956, 1960).
tangular, ovate er elliptical in shape . . . . . . . . . . . . . . . . . . . . 4 Brandis (1907) produced the first comprehensive comparative
When an articulation band consists of 3 cell-rows, the length of the study of the anatomy of gramineous leaves, having examined
same cells in the middle row is nearly equal to the width, cr rnay
mat~rial of "122 species in 21 genera." Of these, 40 species repre-
4 be shorter cr longer . . . . . . . . . ... . 4. Sasa Makino et ShiBata
5. Pseudosasa Makino sentmg 15 eurrently recognized genera of bamboos are diseussed
6. Sasaella Makino or illustrated by hirn: Brandis eondudes (p. 87) that "the uniform-
When an articulation band consists of 3 cell-rows, the articulation ity of the structure of bamboo leaves is very remarkable, consider-
cells in the middle row are mostly measured more in their length ing the great morphological differences in the order. In spite of
than in their width................................. 5
the great variety in regard to important morphological eharaeters,
When an articulation band consists of 3 cell-rows, the articulation
cells in the middle row are generally 11.6-24.9p. wide, even in an the structure of the leaves is the same in all genera and speeies,
exceptional case measure at least 10p. in width .. .... - ..... . as far as they have been examined."
5 . . . . . . . . . . . . . . . . _ ..... _ ...... 7. Semiarundinaria Makino Metcalfe (1956) made a eomparative study of transverse sec-
When an articulation band consists of 3 cell-rows, the' articulation tions of the foliage leaves of 45 speeies, representing 28 currently
ceUs in the middle row are generally 3.3-16.61' wide; but some of recognized genera, and illustrated the central portion of sections
them are sometimes less than 10p. in width ...... . . . . . . . . . 6
Silica corpuscles in the articulation cells are especially numerous
taken at mid-point of the leaf blades of 14 speeies. This author
............................... 8. Sinobambusa Makino eondudes that "although [in their leaf anatomyJ the bamboos
6 9. Chimorwbambusa Makino exhibit eombinations of charaeters that appear to be diagnostie
Silica corpuscles in the articul.ation cells are few in number cr are for the speeies in which they are exemplified, there are no dear
not found ....................................... 7 anatomical lines of demarcation between the genera." Six refer-
Silica cells in the upper epidermis above the veins are mostly 2.5-10U-
enees that represent the prineipal previously published contribu-
long; ind in some species, in addition to such a short silica cell
stomata measuring more than 28fk in length are found in the tions to the knowledge of this subjeet are listed here by Metealfe
7 under epidermis .................... 10. Pleioblastus Nakai (see also Metcalfe 1960: lvii, 583-584, and 586-589; M. V. Brown
Silica cells in the upper epidermis above the veins are about 4.9-10p. 1958; Jaeques-Felix 1958, 1962). Porterfield (1937) deseribed and
or more long, and stomata in the under epidermis are shorter than illustrated the stages in the deveiopment of stomata in the "sheath
about 271" in length ...... 11. Phyllostachys Siebold et Zuccarini leaf" of Phyllostachys nigra and Ph. pubescens.
At the conclusion of his paper (p. 128) Ohki has the following
to say about· the diagnostie value of leaf spodograms: Rhizome sheaths
GeneraIly speaking, there can be no doubt but that leaves have some Of all the kinds of sheaths borne by a bamboo plant, those of the
characters which may be regarded as criteria for the determination of rhizome proper vary least from node to node within a ·given axis.
different species. Yet it may weIl be said that they never possess all the Aseries of sheaths from the suceessive nodes of a pachymorph
characteristics for the classification of species ... Moreover, some genera
rhizome shows a pereeptible progressive change in size that cor-
such as Sasa, Sasaella and Pseudosasa bear so much resemblance as to
 76 THE BAMBOO PLANT

responds to changes in the diameter of the axis. In a leptomorph

rhizome proper, however, the sheaths are remarkably uniform in
all respects. Examples of sheaths from leptomorph rhizomes of
several species of bamboos are shown in Fig. 34. All of the append-
ages of rhizome sheaths proper are depauperate; auricles are
completely suppressed; the blade is commonly reduced to a hard
mucro. Rhizome sheaths are usually strongly imbricate, and per-
sistent or tardily abscissile.

Neck sheaths
Neck sheaths dothe the basal part of every segmented vegetative
axis in all bamboos (p. 12f). Neck sheaths have the simplest of the
various forms assumed by the sheath on any of the axes of the
bamboo plant; moreover, they never subtend a budo The neck
sheath proper is smalI, broadly triangular' in shape, typically hard /

I
and tough in texture, and generally smooth and lustrous on the
inner surface. The blade is obsolete or reduced to aminute, usu-
ally hard mucro; aurides are lacking and the ligule is obsolete. f
Neck sheaths are strongly imbricate and generally absciss very
tardily if at all.

Prophylla
The distinctive sheath at the basal node of each vegetative branch 9 11
[0
is properly called a prophyllum, or prophyll. Some authors refer
Fig. 42. Prophylla of buds at culm nodes, showing variation in form, fusion of
to it by a general term, such as bract, bracteole, or bud scale. By
the margins, and shape of the wings, from species to species, as exemplified by:
its position, orientation, shape, and function, this sheath is placed 1, S~'no~alamus latiflorus,' 2, Bambusa multiplex (as B. nana var. I:YPica);' 3, Phyllostachys
dearly in a dass by itself. It is addorsed (that is, its back is ap- makino~' 4, Arundinaria (as Pleioblastus) simonii:," 5, Sasa paniculata var. paniculato," ~
pressed) to the axis from which the branch that bears it emerges. Pseudosasa japom"ca; 7, Shibataea kumasasa; 8, Semiarundinan"a fastuosa; 9, Sinobambusa
In its rudimentary state, the branch that bears the prophyllum
/0,
tootsif<;· Chimonobambusa quadrangularis; all much enlarged. From Takenouchi
1932:Flg. 58.
is completely endosed by the latter to form a budo In' contrast
with other' types of sheaths described, the prophyllum lacks the wings may be entirely separate from each other or they may be
conventional appendages, blade, aurides, and ligule. The charac- joined at the apex of the prophyllum. When the bud is dormant
teristic shapes of prophylla representative of the buds at culm these wings are appressed to the axis on which the bud is borne.
nodes in different species are illustrated in Fig. 42. Typical pro- The margins of the prophyllum may be either free or fused to-
phylla have in common two salient vascular bundles (veins), which gether in varying degrees. Some prophylla are in this manner
mark the "keels" along which the margins are folded forward, closed all the way 'Trom base to tip while others are only partly
and which set off the "margins" from the "back." The prominence closed (Fig. 43).
of each keel is generally accentuated by a wing of thin tissue. The As in other types of sheaths, the size, shape, vesture, ~d otlier
 VEGETATIVE PHASE: THE MATURING PLANT 79

Roots
Roots are the only unsegmented vegetative axes in a bamboo plant.
They conform to the popular characterization "fibrous" (in a
sense of the term not found in ]ackson) which probably is best
described as slender, roughly cylindrical, and apparently not
increasing in diameter with age. In some large bamboos, as in
Guadua angustifolia, subterranean root axes of the first oroer may be
relatively massive, with a diameter cf nearly a centimeter.
The ephemeral primary root of the seedling bamboo is soon
overshadowed by the progressive development of adventitious
roots, which take over the functions of anchorage and absorption.
Adventitious roots in the bamboos are initiated only at a level
immediately distal to a sheath node. They develop chiefiy at the
nodes of the rhizomes proper and at the underground nodes of
A s the culms (Fig. 44, A). In certain species with pachymorph rhi"
zomes root primorclia form on the swollen base of primary branches
at lower midculm nodes (see text, p. 244). In many species, roots
also appear normally in a primordial or partially developed state
on nodes at varying levels of the aerial part of the culni (Fig.
44, B). In Bambusa vulgaris, Gigantochloa verticillata, Chusquea pittieri,
and several species of at least one genus with leptomorph rhizomes
(Chimonobambusa) root primordia may form at nodes in the lower
s' half or more of the culm length. These primordia may remain
dormant, or they may develop to a length of one to several cen-
Fig. 43_ Prophylla of buds at culm nodes, closed and open [orms, each shown in timeters. The degree of their development at the successive nodes
aba.xial view and in cross section: CA, A') Sasa veitchii (as S. albo-marginata) with
the prophyllum closed on the aba..xial face by the fusion of the marginsj CB, B') always diminishes acropetally. In Chusquea pittieri and Chimonobam-
Yushania (as Arundinaria) niitakayamensis, "With the prophyllum open on the aba."ial busa quadrangularis, and in other species of both genera, these
face, the margins not fused, greatly enlarged. From Takenouchi 1932:Fig. 59. aerial roots typically become dry, somewhat indurated, and thin
at the tip, assuming the appearance of thorns, though actually
features of the prophyllum vary more or less from species to spe- they are of rather brittle consistency (see Arber 1934:Fig. 28, B).
cies and with its position on the plant. In buds at successive Published information about the subterranean root system in
nodes of a leptomorph rhizome, the prophylla are always very bamboos is very scarce. Apparently unique is an exploratory study
uniform, but in pachymorph rhizomes they may show appreciable by White and Childers (1945) of the extent and subterranean
changes in size and shape from node to node.· In culms and their distribution of the root system in Bambusa tulda, with particular
several orders of branches, the size, shape, and vesture of the pro- reference to its potential stabilizing effect on the soil where erosion
phylla are modified progressively from the base to the tip of a is a problem. The root system of a well-established clump growing
given axis (Takenouchi 1931a:PI. IU). They are smallest in size on in a heavy clay soi! on a hillside of unrecorded gradient was
leafy twigs, and largest on pachymorph rhizomes. For a discussion studied by digging a broad trench 4 ft deep extending radially
of the prophyllum in infiorescence branching, see p. 108. from the base of the clump to a point more than 17 ft away. The
 VEGETATIVE PHASE: THE MATURlNG PLANT 81

number of roots exposed in each square foot of the surface of

the trench wall was counted. Eighty-three percent of the roots
were present in the upper foot of the soi!; 12 percent between 1
and 2 ft; 4 percent between 2 and 3 ft; and 1 percent between
3 and 4 ft from the surface. The longest roots extended 17 ft from
the base of the clump. The growing tips of the uppermost root
branches extended above the soil into the accumulated mulch of
,, fallen bamboo leaves.
,, As a general rule, in bamboos with pachymorph rhizomes the
,
,I underground roots are much more concentrated, both in their
I
I . place of origin and in the space they occupy, and the radial ex-
,I tension cf individual raots is much greater} than in bamboos with
I
I,
leptomorph rhizomes. However, the more widely ranging growth
,?-H of the leptomorph rhizome effects a certain compensation by
I
, dispersing its sparser array. of shorter roots more rapidly into the
I
I
,I previously unoccupied soiL
Shibata (1900:Figs. 23-47) illustrated anatomical features of
the roots of ten different species, representing six genera. In this
work, and later in collaboration with Makino (Makino and Shi-
bata 1901 :Figs. 15-19), Shibata brought into focus a fundamental
anatomical difference between bam boos of the genus Bambusa and
those of Sasa and other arundinarioid genera. In the endodermis of
the roots of Bambusa, exemplified by Bambusa multiplex (as B. nana)
and B. stenostachya, the cell walls are shown to be thickened sym-
metrically on all sides, while in the arundinarioid genera; ex-
emplified by Sasa borealis and members of Arundinaria, Chimono-
.bambusa, Phyllostachys, Sasa, and Shibataea, the cell walls of the
endodermis are conspicuously more heavily thickened on the side
toward the center of the root.
This and other, apparently correlated, root differences noted
by Shibata may now be recognized as associated with gross morpho-
Fig. 44. (A) "The base of a shoat of Phyllostachys nigra from which the sheaths logical differences, such as type of rhizome, branching habit
have· been removed, showing t~e yaung raots in various stages of deve!opment. in vegetative and reproductive axes, the relative salience cf trans-
BB, basal huds; H, culm; HB) culm base; St) 'Steil' or connecting bridge to rhi- verse veinlets in the leaf blades, and even differences in ecological
zome" (see "The neck," pp. 13t). From Porterfield (1935:Fig. 1). (R) Culm base cf
Chimonoba:mbusa quad:rangularzs, showing adventitious roots: normal subterranean adaptation, that separate two large groups cf genera.
roats and spinelike aerial roots. Redrawn from Takenouchi 1931a:Fig. 4. Further comprehensive and detailed comparative studies are
greatly needed to improve our perspective on external and inter-
nal features of the bamboo root system and the range of variation
within and between both species and genera. See also p. 122, where
the primary root is discussed.
 REPRODUCTIVE PHASE 83

reproduction, Bambusa vulgaris remains to this day one cf the most

vigorous of the known bamboos. No evidence of systemic senes-
cence in sterile plants of this species has come to my attention.
In contrast with this apparent potential permanence of the
purely vegetative state in some bamboos, we find in others a
2 Reproductive Phase persistent tendency to flower. Bambusa lineata, for example, is often
described as "constant flowering" (Gamble 1896:47). Plants of
this bamboo have been under observation upwards of a hundred
years in cultivation at Bogor, Calcutta, and Peradeniya. No record
of the production of fruit, or of the death of the flowering plants,
The duration cf the vegetative state and the incidence cf flowering has been published. Holtturn (1946:343, and personal conversation)
and fruiting vary from one species to another. In some bamboos states .that at Singapore Schizostachyum brachycladum and some
the vegetative state shows signs of persisting indefinitely, the plants plants of S. longispiculatum flower continuously, while S. gracile
producing new growth with undiminished vigor year after year, and S. grande constantly flower and fruit a little, and all of the
for example, Arnndinaria variegata, Bambusa vulgaris) and Sasa tes- plants retain their vegetative vigor apparently undiminished.
sellata, as known in cultivation. Whether the plants urider discus- A few bamboos are designated as flowering annually. Indocal-
sion represent physiological strains distinct in this one character amus wightianus (as Arundinaria wightiana), Bambusa lineata, and
from the rest of the population of the species to which they belong Ochlandra stridula are so listed by Gamble (1896:viii); others are
is not known. Perhaps such bamboos may, in same cases, possess a said to flower constantly. In most records there is no clear evi-
physiological stability which, under most conditions, keeps them dence to serve as a basis for distinguishing the behavior described
perpetually in a vegetative state, and which permits individual as "fiowering annually" from that called aeonstant flowering."
plants under the impact cf certain unknown circumstances-even Since, at low elevations in the deep tropics, temperature and
pure accident of a physiological nature-to produce gametes. At light values vary only slightly during the year, it seems likely
any rate, the behavior in Bambusa vulgaris suggests such a condition. that in both cases the active production of new flowers follows
Ever since itwas described and given a scientific name (now more the yearly pattern of rainfall. Dr. Holtturn observes that in Singa-
than 150 years ago) byWendland (1810), Bambusa vulgaris appears pore, where there is no dry season, flowering is eontinuous, not
to have been the most generally recognized of alJ known bamboos, "annual," while in plaees with a definite dry season flowering
and it has become the most widely distributed species in cultiva- may oceur onee a year.
tion. Apparently the general population of this bamboo has re, The majority of the known bamboos for which records are
mained steadily in a vegetative state as long as the plant has been available fall between two physiological extremes (constant steril-
under scientific observation. Each occurrence of flowering (as ity and constant flowering) by manifesting a eydie recurrenee
reported, and as observed personally) has involved at most only of the flowering state at intervals of several to many years. Indi-
a few plants (see Hildebrand 1954:50). It appears that these always vidual kinds are not always consistent in their flowering habits,
die upon flowering. I have found no published record of bamboos but two general modes of behavior are distinguishable. Bamboos
of this species having produced fruits (see Moebius 1898:86). If of some species generally die within one or two years after flower-
they really do not do so, then no seedling progenies are being ing (Bambusa arundinacea) while others do not die, but their vege-
produced to perpetuate or accentuate, in this species, the tend- tative growth slows down during the flowering period. This latter
ency to enter the flowering state. However, in spite of its appar- group includes all species of the genus Phyllostachys and several
ently not having been "rejuvenated" in recent times by sexual species of Arundinaria whose behavior is a matter of record. In
84 THE BAMBOO PLANT REPRODUCTIVE PHASE 85

order to think clearly about the ontogeny of a bamboo plant, manifest behavior that places it in more than one of these groups.
therefore, it is necessary to distinguish between the life cycle and Dendrocalamus strictus, for example, falls within both the second
the flowering cycle, since the two may not be coextensive. and the third groups. Ghinkul (1936:26) conceives three groups as
In most bamboos the flowering cycle is much greater than one folIows: (1) mostly small species, whose life span is undetermined,
year. Because of its great length in many cases, and because of flowering and bearing fruit repeatedly at short intervals without
frequent conflict in details between different accounts, and since dying; (2) plantsdying regularly after the first and only fruition;
the flowering and fruiting of bamboos has in many cases (espe- (3) plants dying in part, and renewing the dead parts after fruition.
cially in India) relieved a local famine brought on by drought 1t is dear that gregarious fiowering"is not the universal rule;
(see Munro 1868:4), the flowering cycle in the bamboos has been nor is it invariable in those species where it is the general rule.
subject to much popular interest and speculation. Published Flowerings, called sporadic, of only a portion of the plants within
accounts designed to fix the length of the flowering cycle for a given population known or presumed to be of the same age are
particular species are rather numerous. Suessenguth (1925) and frequently recorded. The flowering of some of the culms, and not
Blatter (1929-1930) have published the most complete available others, in a single clump has been recorded. And I have in my
summaries of such records. These relate principally to Asiatic own records a note (dated Canton, China, June 19, 1938) register-
species, with extended discussions cf the circumstances, patterns, ing the flowering of a single twig on a plant of Bambusa eutuldoides
and possible causes of flowering in bamboos. The evidence pre- (Lingnan University Bamboo Garden accession number LUBG
sented by such records usually consists primarily of aseries of 1206). No further flowering was seen in the following 18 months
dates at which a given species flowered gregariously in a given during which I was able to observe the plant. Holtturn (1955:408)
region. From this evidence, many species of bamboo appear to says, "I know of no recorded case of mature Malayan bamboos
have a characteristic and more or less sharply defined flowering flowering gregariously and then dying, as is the habit of many
cycle, of the order of roughly 1, 3, 7, 11, 15, 30,48,60, or possibly bamboos in the monsoon region (a plant may be considerably
(Kawamura 1927:341) even 120 years, at the end of which time weakened by the full flowering of several culms, but may yet
all plants of a given seedling generation flower gregariously. gradually regenerate from small basal buds)."
However, the records on which the flowering dates assembled by
these authors were based are in no case documented by first-hand
personal observation of the complete cycle in a· single heredita,y Fruiting behavior in different bam boos
line from seed to seed. Parodi's documentation (1955), based on It is rarely possible for even the most interested and dedicated
his personal observation of one full flowering and fruiting cycle observer to assemble a complete account of the full course and
in Guadua trinii, appears to be unique. Fuller details are given on the outcome offlowering in a given bamboo. Two important items
p. 275. (see also Bambusa copelandii, p. 269). A scientific approach of information most often lacking in available reports are: whether,
to the study of the question of flowering cycles in bamboos calls and in what quantity, mature or viable fruits were produced; and
for the frank recognition of one hard fact. Existing records do not whether any plants survived the flowering period.
provide· a positive basis for the generally held assumption that The Indian Forester (1875- ) contains rather frequent notices,
same bamboos have, as an innate character, a flowering eyde of communicated by members of the Indian Forest Service and
precise and invariable length. others, of the availability, in quantity, of the seeds of certain bam-
On the basis of their flowering behavior, Brandis (1906:662) boos. As a result of these notices and published accounts of the
defined three groups of bamboos. These are briefly interpreted by relief of famine conditions in India by heavy yields of edible fruits
Blatter (1929-1930:136) as folIows: (1) those that flower annually following the flowering of bamboos such as Bambusa arundinacea
or nearly so; (2) those that flower gregariously and periodically; (Munro 1868:4) and Dendrocalamus strictus (Arber 1934:95), there
(3) those that flower irregularly. However, a given species may is abroad a rather widespread impression that heavy fruiting
86 TIfE BAMBOO PLANT REPRODUCTIVE PHASE 87

automatically follows gregarious flowering in the bamboos as a

general rule. This impression may have been strengthened by Flowering habits of bamboos as reJated to taxonomy
and the problem of fieJd identification
accounts of the plagues of rats that deveJop periodically in south-
ern Brazil, when local populations of certain bamboos (Mero- To the taxonomist and to the plant coll~ctor, the bamboos present
stachys spp.) flower gregariously and fruit heavily (Pereira 1941; special difficulties. These difficulties have their origin in several
Giovannoni et al. 1946, see also Udagawa 1958). However, a dif- characteristics common to most species. Rarity of flowering is
ferent impression has been derived from certain published records one of them. Another is the more or less severe suppression of
and from personal observations made during aperiod cf more vegetative activity whicli generally prevails during the flowering
than a third of a century. These observations cover flowering period, and which in some species usually results in the death of
plants from twenty of the recognized genera, and thousands of the plant. In some tropical bamboos, a failure to initiate new
pres~rved flowering specimens cf bamboos representing, in aggre- vegetative growth at the usual season may give advance warning
gate, all of the known genera. They add up to the impression of the impending onset of flowering. This means that intact ex-
that the incidence of maturation of fruits is reJativeJy low in the. amples of the perishable culm sheaths will usually not be avail-
majority of known bamboos. It appears that abundant yieJds able when flowering takes place.
occur in only a reJativeJy few species out of the hundreds that Since the taxonomie differentiation anddescription of all
have been observed in flower; and these few are principally bam- angiosperms have been, from the time of Linnaeus, based primar-
boos not under cultivation. However, this impression should be ily upon the reproductive structures, it is only natural that bam-
disciplined by a recognition of the diverse nature of the evidence. boos not in flower or fruit would be passed over by most collectors.
In bamboos that have a long flowering period, the number of It is also quite natural that little or no effect would be made to
fruits discoverable at a given moment is usually very low. This incorporate a represe~tation cf 'the vegetative parts cf the plant,
suggests that either the set of fruit, or its maturation, is reJativeJy since systematic procedures traditionally concede only minor tax-
infrequent, in both time and space. Again, in most bamboos the onomie value to characters from these structures. Moreover, the
florets bearing mature fruits are more or less promptly JeJeased most neglected vegetative parts of a bamboo plant are usuallY
by the disarticulation of the rachilla, and fall to the ground. Wild bulky and require special treatment.
creatures, particularly certain birds and rodents, harvest them However, certain vegetative structures may be used effectively
either from the plant or from the ground. Hughes (1951) states for the recognition of bamboo entities, without reference to the
that in his experience at Plymouth, North Carolina, the incidence flowers, after the range of their variation in the plant has been
of destruction of the immature fruits by insects is so high in studied. Prominent among these vegetative structures is the culm
flowering natural stands of Arundinaria [tectal that in order to get sheath. Since the production of new culms, and therefore of culm
any mature seeds he found it necessary to dust the inflorescences sheaths, generally ceases entirely during the flowering, and since
regularly with DDT. Of Guadua amplexifolia, which flowered in the old sheaths usually deteriorate rapidly, these struetures are
1954 at the Federal Experiment Station in Puerto Rico, Kennard rarely found associated in herbarium specimens with flowers from
(1955:194) says: "In spite of the fact that many thousands of) the same plant. Branching habit and other features of the several
flowers were produced and that the pollen apparently was viable, categories of segmented axes are also useful for taxonomie purposes.
fruit set was very low. Examination of thousands of spikeJets It is important therefore, in the study of the bamboos, to build
yieJded only 1003 fruits." Only 1 percent of the seeds that were up independently for each recognizable entity a complete repre-
planted germinated. sentation (specimens, notes, sketches, and photographs ) of the
For circumstantial records partially documenting the flowering vegetative struetures of the sterile plant. Then, when flowering
and fruiting behavior of seJected bamboos of different genera and finally occurs, the association of characters from the two phases
species, see Chapter 6. of a given entity will be facilitated. In this connection, it is im-
88 THE BAMBOO PLANT REPRODUCTNE PHASE 89

portant to be aware tbat the morphological expression of com-

ponent parts varies within certain limits, in correlation with the The inflorescence
relative age of tbe different parts of the plant, and tbat the mode The earliest description of tbe bamboo inflorescence as a distinct
of expression found in the youngest (currently produced) culms structural unit apparently is tbat of Nees (1841:460). This was
reflects tbe relative maturity and the current stature of the plant quoted verbatim by Munro (1868:11) as a part of his character-
as a whole (see McClure 1935:193-197, and Holttum 1958b:I-2). ization of the "Bambusaceae." Freely translated it reads as folIows:
The importance of the foregoing considerations relates primarily "The inflorescence [in Tribe xvii, the Bambuseae üf Nees] is a
to tbe fact that, as encountered in the field, most bamboo plants panicle, very rarely witb strongly compound branching, more
are not in a flowering condition; their vegetative structures provide commonly smaller and depauperate; in some bamboos it is re-
the only means of identifying them. duced to the form of a raceme or a spike (in some verticillate, in
Franchet (1889:281) in his remarks following tbe character- otbers simple); it may even be reduced to a peduncle bearing a
ization of a new genus of bamboos asserts that single spike." Gamble (1896:1), in his characterization ofthe tribe,
it is especially in its vegetative characters that Glaziophyton is clearly dis-
simplified it to tbe following phrase: "Inflorescence various, usually
tinguished from all of the known Bambuseae, and perhaps even more so a large, compound panicle with spicate branches." These defini-
from most of the other genera of the group to which it belongs. 1t was, tions approximately cover the usage still generally accepted. The
of course, a long time ago that Kunth noted that, in the Gramineae, situation is not much helped by tbe adoption of such borrowed
most of the genera could he clearly separated only by their vegetative terms as "thyrsoid" and "capitate." The "infiorescence" of Thyr-
organs, as the floral organs furnish only vague elements for distinguishing sostachys is not really like a thyrse. And tbe term "capitate" is
the plants, and even these [elements] are of douhtful value.
applied, without discrimination or qualification, to botb the glo-
Franchet's assertion with regard to Glaziophyton is correct. But bose indeterminate infiorescence of Guadua capitata (a cluster of
tbe statement attributed to Kunth (I have not been able to find pseudospikelets) and the subglobose determinate inflorescence of
the original source of it) goes beyond what can be supported by Athroostachys capitata, which latter is really a short-branched panic-
available evidence. While it is possible to recognize and identify ulate raceme.
individual species of bamboos, Once they are thoroughly known, In the introductory chapter of his monograph of the bamboos,
by an adequate array of vegetative characters, it is not always Ruprecht (1839:15) called attention to the difference between
possible, given the present state of our knowledge, to determine Arundinan·a and Bambusa in respect to tbe "disposition of the
the generic aflinities of an unknown bamboo witb certainty, by spikelets." Howeyer, in setting up the two major divisions of
means of vegetative structures alone. tbe bamboos known to him, Ruprecht differentiated tbem not on
During tbe first half of the present century, and particularly tbis character (tbat is, the branching habit of tbe inflorescence)
in Japan, the description of new species of bamboos on the basis but on the number of stamens in the androecium, thus: '(Bam-
of vegetative structures alone was practiced by taxonomists on an busae verae" with 6 stamens, and "Arundinariae" with 3 stamens.
increasing scale. Where conscientiously executed, this procedure In the introduction to his description of the genus Schizostachyum,
can be sanctioned on the same premises as the naming and de- Nees (1829:534, Obs. IIl) called attention tothe "peculiar struc-
scribing of Fungi Imperfecti from sterile materiaL This expedient ture" (singularem strucMam) of th" "spikelets" in the type species
is in general more satisfactory for bamboos of large stature than of this genus. From his description of this structure, it is evident
for small ones, when characters are drawn from gross morphology that tbe inflorescences in tbe specimen he had before him con-
alone. In any case, the original description should be full enough sisted of pseudospikelets in an advanced state of development
to set forth reliable specific characters which, in combination, are (see pp. 65 and 66). Ruprecht (1839:44; 1840:134), in his own
truly distinctive. description of tbe genus Schizostachyum, used the following words
90 THE BAMBOO PLANT REPRODUCTIVE PHASE 91

(translated from his Latin): "The glumes of Nees's description are every branch of every order. Two basically different forms of
[labeled] in our illustration as lemmas [but tbey are] really bracts inflorescence may be distinguished: indeterminate and determinate
subtending spikelets or buds." Here tbe matter rested until Moe- (Figs. 45-53). These two forms manifest distinctive characteristics
bius (1898:PI. IV, Fig. III) figured an inflorescence of Bamlrusa related to fundamental differences in their manner of development.
vulgaris and, during tbe course of a discussion of morphological
details, made several references to .the continuing nature cf its The indeterminate inflorescence
development. An indeterminate bamboo inflorescence is one the course cf whose
Pilger (1927:Figs. 1 and 2) illustrated an inflorescence of Guadua development is prolonged indefinitely by the progressive elabora-
tessmannii and presented a diagram purporting to show the details tion of its branching. A separate "grand period of growth'" is
cf its organization. Pilger's description cf the dynamic aspects cf initiated and completed, independently, in each flowering axis
the development of this inflorescence is difficult to follow. More- of each successive order of branches. Each flowering aXis is spike-
over, the diagram of its organization (ibid., Fig. 2) is defective in letlike in appearance, and it terminates in aspikelet. However,
respect to a fundamental detail. It fails to show tbat all of the its basal part is a very short rachis, clothed with lemmalike bracts,
branches of the indeterminate inflorescence characteristic of this each of which subtends a prophyllate branch bud instead of a
species recapitulate the organization and tbe dynamic aspect of flower. For this reason, tbese spikeletlike branches have been
the primary aXis. To be explicit, the ultimate branches are not given the distinctive name, pseudospikelets (McClure 1934:Fig. I).
shown to have the basal buds (mentioned in the text) that are Being repositories of meristem, tbe buds at the base of a pseudo-
themselves potential pseudospikelets, and capable of developing spikelet make possible the continued expansion of the indetermin-
into' reproductive branches exactly like tbe ones on which tbeyare ate inflorescence as long as the physiological state of tbe meristem
borne. F or this reason, Pilger's diagram of tbis indeterminately and adjacent tissues favors the breaking of their dormancy and
branching inflorescence portrays it as exactly like the fundamen- the continued development of tbe new axes and tbeir appendages.
tally different inflorescence of Glaziophyton mirabile, in which, while
each branch is prophyllate, and subtended by a bract, the branch-
The determinate inflorescence
ing is determinate. Also notewortby is the fact !hat, in confining
his comparisons to grasses of nonbambusoid genera, Pilger failed A determinate bamboo inflorescence is one the course cf whose
to place this indeterminate type of inflorescence in perspective development is strictly limited to a single "grand period of growtb"
among the bamboos. Pilger's important contribution received no that encompasses the elaboration of the whole complement, of
published notice from students of the bamboos until McClure branches of a solitary rachis. Terminal growtb ceases in all branches
(1934) presented a study ofthe inflorescence in the Chinese species of the inflorescence within a limited time. Each branch terminates
of Schizostachyum, having arrived at the concept "pseudospikelet" in a conventional spikelet; no meristem remains afterward in the
to distinguish a whole (spikeletlike) branch of an indeterminate form of dormant lateral buds.
inflorescence from tbe spikelet proper that terminates it. The
following statement is an attempt to elucidate more fully some Other coritrasts
hitberto generally neglected basic features, particularly the branch- The expansion of an indeterminate inflorescence follows very
ing, of the bamboo inflorescence. The extreme forms of expression closely the pattern of development characteristic of tbe vegetative
are described and given distinctive names. phase of growth. The expansion of the determinate inflorescence,
The bamboo inflorescence is an aXis, or a system ofaxes (asso-
lAccording to Porterfield (1928:327), "The grand period of growth covers not only the
ciated branches) emanating from a common axis, the primar-y enlargement of some part already" in existence, hut the production of a fully fonned organ
rachls. The primary Tachis ends in aspikelet; and so does its or serics of organs with definite parts ::i.nd specific characters."
 REPRODUCTlVE PHASE 93

on the other hand, manifests a high degree of simplification, as in

the panicle of Sasa veitchii (Fig. 46), for example. This change
marks an evolution in the direction of patterns common in many
of the herbaceous grasses of nonbambusoid subfamilies (Poa
pratensis, for example)? The diversity of form manifested by the
determinate infiorescence is discussed on pp. lOlf.

The course of development of the indeterminate inflorescence

The following description of the development of the indeterminate
inflorescence departs, in some minor details, from that of Holttum
(1956a:82; see "spikelet tufts") and an earlier one (McClure 1934)
and it is more complete than either (see also Nees 1829; Moebius
1898; and Pilger 1927). The basic struetural unit of the indeter-
minate inflorescence is the pseudospikelet. For clarity it is necessary
to think' in terms of two categories of pseudospikelets: primary
and secondary. Primary pseudospikelets have their origin on seg-
mented axes that are themselves vegetative, in a morphological
sense at least. Secondary pseudospikelets originate from other
pseudospikelets.
Primary pseudospikelets differ in minor structural details which
are correlated with their respective positions on the vegetative
axis that bears them. Lateral ones are sessile; a terminal one is
made pedicellate by the distal internode of the vegetative axis
that bears it. Closer examination reveals further that, whereas the
lateral ones have a two-keeled prophyllum as the first (proximal)
foliar appendage, the terminal one has an unkeeled bract in place
of this two-keeled structure. (Fig. 47, A). The prophyllum that
embraces this axis is located far below, at the base of the vegetative
branch itself. This is the only structural difference between a
terminal pseudospikelet and a lateral one. Secondary pseudospike-
lets of all orders are of the sessile category. A lateral primary
pseudospikelet is therefore the prototype of all secondary pseudo-
spikelets. And since secondary ones are by far the most numerous
form in any flowering specimen, adescription of the develop-
ment and structure of its prototype will be most generally usefuL
Fig.45. Bambusa multiplex var. n:viereorum: (A) base of plant (showing pachymorph Initiating the sequence of events, a solitary prophyllate infio-
rhizome and caespitose clump habit); (E) leafy branches terminating in primary
2No report of the occurrence of indeterminately brancmng inflorescences in any grass
pseudospikelets (young inflorescences of indeterminate branching). Drawn by outside the subfamily Bambusoideae has yet come to light.
EImer W. Smith.
 REPRODUCTIVE PHASE 95

Fig. 47. Diagrammatic comparison of a branch of (A) Ba:rnbusa multiplex ter-

minating in an indeterminate inflorescence with a branch of (B) Sasa veitchii ter-
minating in a determ.inate inflorescence. The two dark objects at the base of the
infiorescence proper in (A) are prophyllate buds, each capable of duplicating the
pseudospikelet of which they are apart. This is a basic distinguishing feature of
the indetermtnate inflorescence. The broken lines in the infiorescence proper in (B)
Fig. 46~ Sasa veitchii. Portion of aplant, showing leptomorph rhizome, diffuse indicate fully developed spikelets like the apical one shown. Here no buds or
clump habit, and leafy culm hearing solitary branches (monoclade branch com- other repositories of meristem remain to initiate additional branches. This is a
plements) with determinate inflorescenees. Adapted from Makino and Shibata basic distinguishing feature of the determ.inate barnboo inflorescence. Cf. Fig. 48
(1901:PI. I). (A, B).
96 THE BAMBOO PLANT REPRODUCTIVE PHASE 97

rescence bud lateral to a segmented vegetative axis develops into short-often not more than 1 or 2 mm in length. This being
a short primary reproductive branch. This branch is spikeletlike the case, the inflorescence cluster tends to become congested as
in superficial appearance, but may be recognized as a pseudo- the number of orders (successive "generations") of pseudospikelets
spikelet by the presence of a two-keeled prophyllum at its base, increases. When the spikelets are all persistent, a globular head is
followed by one or more bud-subtending bracts (Figs. 48, Band formed, as in Dendrocalamus strictus. Commonly, however, the spike-
49, B). The presence of branch buds (revealed only by careful
dissection) characterizes this part of the axis of the pseudospikelet
as arachis; the distal part is the rachilla of the spikelet proper.
The transition between the lower, bud-bearing part of the pseudo- >I~-
spikelet and the spikelet proper that follows is marked (1) by the
initiation of the development of abscission layers at the· nodes
of the axis and (2) often by the intercalation of one or two empty \

~
glumes (see p. 109). With or without intervening empty glumes,
the transformation of the rachis to arachilla is made plain by ~
the appearance, at the next level, of a floret-the beginning of the

~ ~
spikelet proper. 11
B E F
Sometimes, as in Schizostachyum terminale (teste Holttum 1958:51,
52), Bonia tonkinensis, and Arundinaria prainii (firsthand observation),
the buds subtended by bracts on the rachis of the primary pseudo-
l~
~B'
spikelet may remain dormant. In such cases, the individual inflo- )$
>

rescence consists of hut a single pseudospikelet. In most bamboos,

however, these buds break more or less promptly, and each re-
sulting branch becomes a secondary pseudospikelet, whose basal
buds produce tertiary pseudospikelets, and so on. The promptness
with which the successive orders of pseudospikelet buds formnew
pseudospikelets is characteristic and relatively uniform in each
bamboo. In Oreobambos buchwaldii, an embryo pseudospikelet cluster
forms precociously, emerging from the primary prophyllum with
several orders of branches already well advanced. In .Dendrocalamus
A.' »' E' F
• i71
~~ ~I1
strictus, tbe development from the primary pseudospikelet to a Fig. 48. Components of the infiorescence in Bambusa multiplex (A-F) and Sasa
full-blown, spherical head, consisting of many pseudospikelets, vcitchii (A'-F). CA) Brauch of the indeterminate inflorescence-a I:seud~spikelet
with a 2-keeled prophyllum at its base. (A~) Branch of the determmate l~flores­
proceeds at a spectacular rate, but apparently no record of the cence-a spikelet with 2 empty glumes at 11s base. (B) Base of (A), showmg. the
duration of the active growth of these quickly maturing pseudo- prophyllum, the two bud-subtending bracts, and the empty glume t~at precede
spikelet clusters has been published. In Bambusa vulgaris, inflo- the first lemma of the spikelet (not shown). (E') Base of the flowenng branch
that bears (A'), showing the 2-keeled prophyllum--:-here (in contrast) far removed
rescence clusters have been observed to continue to produce new from the spikelet. (C, C') Floret. (D, D') Lemrp.a. (E, E') Palea. (F, F') the com-
pseudospikelets related to the same primary axis through two full ponent parts of the flow~: lodicules: st~.ens (androeci~), pistil (~noecium),
years (Moebius 1898), while in Bambusa multiplex cv 'Alphonse and fruit. Based on speclmens depoSlted 10 the D.S. NatlOnal HerbarlUffi .. (A-F)
US 516519 eolleeted by G. T. Lane (sn) February 25, 1898, in The Royal
Karr' they apparently may remain active for astilI longer period. Botanie Garden, Caleutta. (A'-F) US 2241716, eolleeted by T. Makino (sn)
The internodes of the rachis of the pseudospikelet are very June 1909, near Ihagi-mura, Musashi, Japan.
 REPRODUCTIVE PHASE 99

nidularia in two stages of development: "before anthesis" arid

"after anthesis." The capitate form resulting from the progressive
branching from the basal nodes of short axes in the successive
orders of branching is strikingly shown (Fig. 50) but the author
makes no reference to the indeterminate (continuing) nature of
the branching habit of this inflorescence. Porterfield (p. 259) says
only that "the characteristic capitate cluster [inflorescence1 of

B
Fig.49. The indeterminate inflorescence in Dendrocalamus sikkimensis: CA) fiower-
ing branch "about one month after"anthesis," with a cluster of pseudospikelets Fig. 50. The inflorescence of Phyllostachys nidularia, as illustrated by Porterfield
at each nodej CE) pseudospikelet Crecognizable as such by the prophyllum at its (1926:Figs. 1, 2). (A) An early stage, ('before anthe:?is," showing the form of the
base on the left) terminated by aspikelet with three florets. A solitary stigma primary branch (a pseudospikelet) with the spikelet tips of pseudospikelets of
(tip of the style) is seen projecting from the apex of the lowermost floret. Six higher order protruding beyond the uppermost bracts. The primary pseudo-
stamens have emerged from each of the upper two florets. Pollen is discharged spikelet in this illustration appears to be terminal to the internode shown below
from the anthers through apical pores. From Arber 1934:Fig. 32A. it, but the sheath scar that precedes the first bract suggests that it may be a
lateral branch at the node that terminated the internode shown. If it actually
was subtended by a leaf sheath (now fallen away) of which the scar is shown,
lets terrninating the earlier pseudospikelets in a given cluster fall
then a prophyllum preceded the small bracts shown at its base. This prophyllum
away more or less promptly as they mature, leaving the cluster of could not be shown in this view because it would lie behind the pseudospikelet
pseudospikelets sparser, and irregular in shape. This is character- as shown here. It would be addorsed to the next internode of the axis on which
istic of some bamboos of the genus Bambusa, all known species of the pseudospikelet is borne, and which is hidden by the latter. (E) A later stage,
('after anthesis," showing tbe capitate form attributed by Porterfield (p. 259) to
Elytrostachys, and many species of Schizostachyum. To the experienced the "telescoping" of the inflorescence. Actually, this is a tuft of pseudospikelets,
eye, the naked rachis tips that remain afford a clue to the loss of an indeterminate form of inflorescence. Its congested form results from the rapid
spikelets that has occurred, and an intimation of the relative age consecutive branching of the very short pseudospikelet axes of successive orders
arising from buds in the axils of basal bracts of each pseudospikelet. It appears
of the inflorescence.
that tbe indeterminate nature of the inflorescence in bamboos of the genus
Porterfield (1926) illustrated the inflorescence of Phyllostachys Phyllostachys has not hitherto been reported in tbe literature.
 100 THE BAMBOO PLANT REPRODUCTIVE PHASE 101

Ph. nidularia is in effect a cluster of spikelets in which the branch- The combination of prophyllate lower branches subtended by
ing inflorescence has been telescoped." bracts with upper branches neither prophyllate nor subtended by
bracts, found in the inflorescences of Thamnocalamus spathiflorus,
suggests a stage of evolution perhaps still further removed from
Variations and "anomalies" in the indeterminate inflorescence
the indeterminately branching condition.·
The pseudospikelets of successively higher orders may become
progressively depauperate. In the spikelet terminating a primary
Diversity of form manifested by the determinate inflor~scence
pseudospikelet the flowers are generally perfect and, potentially
at least, functional. In pseudospikelets that are branches of succes- The brief general statement (p. 91) characterizing the determinate
sively higher orders, the florets and flowers may become smaller, bamboo inflorescence as having but a single grand period of
and the development of the sexualorgans may become progres- growth is elaborated here by a few additional observations. Its
sively restricted. The gynoecium is the first part affected, and punily form may be analyzed for diversity in terms of several features,
staminate flowers may result. Other changes may be noted, as the more important of which are the following: the presence or
well. In Schizostachyum lima, for example, fully developed perfect- absence of a strong central axis (rachis); the spatial disposition
flowered spikelets of the primary pseudospikelets are rather cf the Ioci of insertion of the rachis branches; the dimensions,
promptly deciduous, while those bearing rudimentary perfeet physical proportions, and habit of the rachis and its branches;
flowers absciss much more tardily, and those bearing purely starn- the number of orders of branches; the absence, or presence and
inate flowers are persistent. In plants that have been flowering degree of development, of pulvini at the base of branches; the
for same time, this behavior results in a growing preponderance, absence, or presence and degree of development, of bracts sub-·
in pseudospikelet clusters, of spikelets with rudimentary perfect tending branches; and the presence or absence of prophylla.
flowers and rudimentary staminate ones (McClure 1934:544ft). The rachis .may be strong and excurrent (Sasa veitchi,), or it
When preserVed in herbaria, specimens with such a history may may be deliquescent (Aulonemia queko). A strong central axis with
give the impression that the species they represent has uniseA"Ual a single order of widely spaced, very short, solitary branches pro-
flowers and the sexual pattern called polygamy (see p. 117), both duces an elongate spicate or subspicate raceme in Arthrostylidium
as typical expressions. cubense. In such inflorescences the pattern of emergence of the
Again, the inflorescence branches (either those of higher order primary branches is usually somewhat obscured by torsion of the
or those higher on the rachis) may fail to develop buds at basal rachis. This twisting produces a secund, subspicate raceme in
nodes. This puts a stop tothe further production of pseudospike- most known species of Merostachys. In Merostachys retrorsa and M.
lets. Such a movement away from the typical indeterminate form pauciflora most of the spikelets are retrorsely oriented. Each reflexed
of expression (clustered pseudospikelets) might. be construed as a pedicel bears a pair of minute pulvini inserted adaxially at its
step in the direction of (or a sign of the potential for) evolution base. In other species of this genus the spikelets are antrorsely
of the bamboo inflorescence toward a determinate form (raceme oriented and pulvini appear to be ladung. In Athroostachys capitata
or panicle). The essentially determinate inflorescences in Glazio- the infiorescence assurnes a capitate aspect, hut on dissection it
phyton mirabile and Greslania montana appear to be the result of a is seen to be a very compact raceme which may be designated as
strang trend in this direction. Here, the inflorescence branches "paniculate" because the rachis bears its branches. mostly in fas-
bearwell-developed prophylla and are subtended by well-developed cicles of two or three, of which each primary one appears to have
bracts. The occurrence of an occasional dormant prophyllate bud given rise, subcutaneously, to one or two secondary ones. Each
on the ultimate branches of an inflorescence, especially in the fascicle is subtended by a single, narrowly acuminate bract bearing
latter genus, suggests a relict tendency toward inde!erminateness. a-long awn. In Arundinaria teeta the usually racemose inflorescence
 102 THE BAMBOO PLANT REPRODUCTIVE PHASE 103

may occasionally combine the branching habits of both a raceme of a physiological state that corresponds either to dormancy or
and a panicle or it may assume a typically paniculate form. to the loss of the meristemmatic potential-or possibly to the
A streng, elongate, e~current central axis with more than one exhaustion of stored nutrients. In the case cf an indeterminate
order of branches produces a "typical" panicle. The primary infiorescence comprising but a single pseudospikelet, the potential
branches of a panicle may be solitary, as in. Sasa veitchii (Fig. 46), for subsequent branching might remain, in the form of dormant
or two to several branches may emerge from a' single node of the buds at the base of the primary pseudospikelet (as in Bonia ton-
rachis, as in Arundinaria amplissima. Branches of higher orders are kinensis and Arundinaria prainiz). On the other hand, 'when a de-
nearly always solitary. All orders of branches may remain more terminate infiorescence stops growing after producing one spikelet,
or less appressed or only slightly divergent (Arthrostylidium sub- that usually is theend ofthe story. The development of additional
pectinatum). In Indocalamus sinicus the individual branches of the spikelets is no langer possible, since no meristem remains. How-
panicle are very lang and stiff; as they mature, they are given a ever, it sometimes happens in an inflorescence with determinate
wide angle with the rachis by the operation of basal pulvi:di. In branching that the spikelets themselves do not stop growing at
all but one of the species mentioned, as in most bamboos witli the usual point. The continued activity of the apical meristem
determinate infiorescences, the bracts subtending branches of the may extend the length of some spikelets to alm ost a foot, as I
inflorescences are rudimentary cr absent, often being represented have observed in fiowering plants of Arundinaria dolichantha. This
only by a mere line or ridge below the locus of insertion of primary suggests the steady persistence, in the meristem at the tip of a
branches. As a rule, however, the bract subtending the lowest spikelet, of a physiological state corresponding to active reproduc-
primary branch of determinate infiorescences is more noticeably tive morphogenesis.
developed than any of the others. As long as the signal that determines the length of the spikelet
is thus inhibited, the spikelet continues to grow apically. H. Fung
695, a specimen of Arundinaria dolichantha collected at Hoh Tung
The bamboo infiorescence from -the point of view of physiology
in Tonkin (now North Vietnam) shows another striking morpho-
The results of determinateness and indeterminateness that appear ' logical anomaly. After producing fiowers for a length of abaut
in the bamboo infiorescence are commonly interpreted by the' 3 in., the rachilla in some of the spikelets reverted to a purely
taxonomist only in terms that will most simply describe the super- vegetative state and terminated in a leafy twig.
ficial aspect of the resulting growth forms. These may be more From the point of view of morphology, the component parts
fully understood when viewed also through the eyes of the phys- of the bamboo infiorescence are conventionally seen as the result
iologist, particularly if attention is given to the extreme range ·af a more cr less radical "modification" of the structures (nodes,
of expression often to be found within the infiorescences of a single internodes, and sheathing appendages) characteristic of the seg-
plant. mented axes of the plant in the vegetative state. The specialized
It sometimes happens (nowand then in Arundinaria simonii, for (or "reduced") sheathing organs of the infiorescence-and even
example) that a determinate infiorescence is reduced to a single the androecium and gynoecium-are commonly referred to as
spikelet borne at the apex of the rachis. Reference was made "leaves" or "sheaths" in modified form (see Watson 1943 and
earlier (p. 96) to a similar reduction in infiorescences whose branch- Arber 1950:68). In some bamboos with indeterminately branching
ing is indeterminate (that is, normally of a continuing nature). infiorescences (Bambusa multiplex and Phyllostachys bambusoides, for
The use cf the term "reduction" in this sense is traditional in example) the transition from the vegetative phase to the repro-
taxonomy that is based solelyon gross morphology. However, ductive phase may be marked by such gradual changes in the
the physiologist probably would see behind the visible phenomenon form of the sheathing structures that the correspondence is, indeed,
(extremely early cessation of growth) the premature development striking (Figs. 45 and 51). The indeterminately branching infio-
 REPRODUCTIVE PHASE 105

tionally, however, the subtending bracts, and prophylla, may be

weil developed (as in Glaziophyton mirabile and Gieslania spp.;
see Pilger 1945:22, 24, on Greslania).
From the physiological point of view, the similarities between
carnponent elements of vegetative and reproductive shoots, espe-
cially in respect to the form and pattern of insertion of certain
cf the foliar organs, da not indicate the lingering infiuence, in
the inflorescence, of the form in which the corresponding (homo-
logous?) organs were cast during the vegetative phase of the plant's
life. They appear to be rather a sign, or the result, of features
common . to the corresponding inner (physiological) states of the
respective bodies of meristem. The distinctive forms and functions
that characterize the inflorescence, particularly at the level of the
androecium and gynoecium, speak eloquently of strang monitor-
ing factors of a physiological nature peculiar to the flowering
state (cf. Heslop-Harrison 1959:269 et passim and Wardlaw 1961).
A Wilson (1942:759) says, "It is becoming widely recognized that
the generally held concept of the essential organs of the flower
as modified foliar appendages rests upon a most insecure foun-
dation." Nozeran (1955:13) credits Thompson (1944) with having
Fig. 51. In same bamboos with indeterminate inflorescences, the bracts that
precede primary pseudospikelets or subtend branches thereof bear more or less
proposed the following ideas, based on studies of ontogeny: (1) the
strongly developed, leaflike blades. This condition prevails in different degrees parts of the flower have no apparent relation to leaves; (2) they
in four speeies of PJ:tyllostachy:s, as illustrated by these figures: (A) Ph. nigra cv. are outgrowths that cannot be equated 100 leaves; (3) all of the
hmon (as var. hmonis); (B) Ph. bambusoides; (C) Ph. aurea; (D) Ph. pubesems (as Ph. primordia of the floral parts are mutually homologous; and (4) their
mz"tis). This leaflike character of the bracts is lacking in same species of this genus,
as in Ph. nidularia (Fig. 50) but is sometimes seen in Bambusa multiplex (Fig. 45). differentiation is monitored solely by the physiological condi-
From Kawamura 1927:Fig. 6. tions that prevail in thefloral axis at the time of their develop-
ment. In this perspective, the plausible traditional designation of
rescence is, in fact, like the vegetative system in being made up all of the floral parts as "modified leaves" loses some of its appeal.
essentially of ramifying axes that are all clearly segmented and This whole subject is admirably treated from the historical and
bear subtending sheathing organs and prophyllate primordia philosophical points of view by Arber (1950:chap. V; see also
(buds) at all points of branching. The determinate inflorescence, Watson 1943; Wardlaw 1956; and Takhtajan 1959:chap. 1).
on the other hand, in the extreme form of its expression, as in The physiological point of view also places in fresh perspective
Sasa veitchii (Figs. 46 and 47, B), has the rachis of the inflorescence several fundamental aspects and features of the reproductive
and its branches of all orders below the spikelet level segmented phase: (1) the sudden change in phyllotaxy, from the distichous
only weakly if at all. These axes are often more strongly paren- expression that prevails elsewhere in the plant (except in the
chymatous in substance than the corresponding branches of an branching of some determinate inflorescences), to the trimerous
indeterminate inflorescence. Subtending foliar organs (bracts) are arrangement of parts that prevails within the flower; (2) the radi-
usually rudimentary or absent, and amplectant prophylla are cal and abrupt changes that take place in the very short axis of
typically lacking entirely in determinate inflorescences. Excep- the flower, giving rise to farms so highly specialized, and so di-
106 THE BAMBOO PLANT REPRODUCTIVE PHASE 107

verse, as palea, lodicules, stamens, and pistil within the space of Now how can we rationalize a comparison between the graup of short
1 or 2 mm; (3) the "anomalous" manifestations in the bamboo flowering branches at one node faund in a bamboo with a panicle of
spikelets as [aund in a grass? If all the lower intemodes o( a tuft of short
flower that appear in the form of teratic structures intermediate
flowering branches [pseudospikelets] in the bamboo were much elon-
between contiguous categories of th~ floral structures just mentioned gated, as [ar as the empty glumes in each case, ..:Ne should have some-
(cf. Munro 1868:152 et passim; Arber 1934:Fig. 202-210); (4) the thing very like a panicle of spikelets, each spikelet on astalk and either
general lack, in the determinate inflorescence (as contrasted with terminal on a branch or in the axil of a bract with a prophyll immedi-
the indeterminate form), of well-developed subtending bracts and ately above the bract. If we eliminate these branch-bearing bracts and
the prophylls adjacent to them and reduce the empty glumes at· the base
of prophylla at the points of branching, and the corresponding
of each spikelet to two, we have exactly the condition of a grass panicle.
loss (below the spikelet level) of both dearly marked segmentation
and distichous branching; (5) the diverse behavior of different As Pilger did earlier (see p. 90), Holtturn here sets off the in-
species and strains of bamboos in respect to the ini tia tion, patterns determinate infloreseence ("short flowering branches") of a bam-
of development, and fruition of the reproductive phase; (6) the boo, against the determinate inflorescence ("panide of spikelets")
corresponding patterns of behavior in the vegetative life pf the' of a nonbainbusoid grass. Holtturn states elsewhere (1956a: 85) that
plant, both during and after flowering; and (7) inflorescences the primitive condition would seem to be that now normal in several
intermediate between the two extremes described above as deter- species of Malayan Schizostachyum, in which every branchlet is leafy
minate and indeterminate. Studies of the bamboos, along the lines (with normal blades), the spike!et tufts occurring atthe few most distal
illustrated by Zimmerman (1961), should improve our under- nodes on many branches, flowering being often continuous. This con-
standing of the genesis of some of these ~anifestations. dition could easily be modified to that of a terminal panicle on each
leafy branchlet.
The development of a comprehensive perspective with regard
to the characteristics of the intermediate (transitional) morpho- It appears that conventional preoccupation with observations
logical expressions just mentioned requires a dose examination of made at the spikelet and flower levels has diverted attention from
the whole range of the known genera of bamboos. Such a broad important characters that can be brought into focus only by ex-
perspective is not yet available in published dissertations, partly amining the structure of the inflorescence as a whole, and studying
because of the hitherto incomplete coverage of the known forms, its development in the light of facts relating to physiology and
but chiefly because the dose examination of the materials has morphogenesis (cf. Thompson 1944, especially pp. 66 and 68, and
generally been confined to the level of the flower and the spike- the discussion that fol!ows). Systematic studies carried out in the
let. As a rule, published observations on individual species have present century have resulted in the description of hundreds of
not penetrated deeply enough to reveal the basic nature and origin new species and a number of new genera, but the fundamental
of the branching habit of either the vegetative or the flowering nature of the branching habit of the bamboo inflorescence as
axes. pictured herein has been general!y neglected, in spite of its phy-
Comparative studies of the branching habit of the bamboo logenetie signifieanee, and its taxonomie value. Arber, whose
inflorescence carried out personally indicate that the evolution, numerous anatomical studies have so greatly helped to elucidate
by stages, from the less specialized form of the typical indetermi- the structural diversity manifested in the spikelet and flower of
nate inflorescence, as seen for example in Bambusa multiplex (Figs. the Gramineae, has dealt more ful!y with the inflorescence, as
45 and 48, A-F), to the more specialized form of the typical such, in other plants than with the branching habit of the bamboo
determinate inflorescence, as seen in Sasa veitchii (Figs. 46 and 48, inflorescence and its significance (cf. Arber 1926-1929, 1934, and
A' -F), corresponds to the general pattern hypothesized by Holtturn 1950). Holtturn (1956a, b, 1958a) has made fruitful preliminary
(1958:18-19) in the following words: studies of the principal vegetative organs, the infiorescence, and
 108 THE BAMBOO PLANT REPRODUCTIVE PHASE 109

the fruit (where available) in bamboos of seven genera represented Spikelet, floret, and flower
in the flora of Malaya. In all of these genera the rhizome is pachy- The spikelet is a basic structural unit of the bamboo inflorescence.
morph. In six of them (Bambusa, Dendrocalamus, Dinochloa, Gigan- Essentially, it represents a distinct and characteristic aggregation
tochloa, Schizostachyum, and Thyrsostachys) the branching of the of flowers and structures intimately associated with them. The
inflorescence is indeterminate. In the seventh, Racemobambos~ the spikelet consists of a specialized axis, the rachilla, and its branches,
inflorescence is determinate in its brandung habit. the flowers. The rachilla is clothed in aseries of imbricate sheath-
ing appendages. Of these there are two maJor categories, called
The prophyllum in inflorescence branching by American agrostologists glumes (or empty glumes), and lemmas
Two-keeled prophylla are generally lacking on the branches of (or flowering glumes). Empty glumes are characterized (1) by
determinate inflorescences. In typical indeterminate inflorescences, occupying the basal (physiologically transitional) portion of the
on the other hand, each branch develops from a prophyllate budo spikelet axis; (2) by being "empty," that is, not subtenoing either
In gross dissections of such inflorescences, the distinctive features a branch bud or a flower bud; and (3) by having, in determinate
of the two-keeled prophyllum make it a marker useful in following inflorescences generally, a smaller size and slightly different shape
the sequence of branching, especially where the branches of suc- in comparison with the lemmas. The term lemma, or flowering
cessive orders are extremely short (as in Schizostachyum blumii; see glume, stands for a glume that subtends a flower. A lemma is
McClure 1934:546). referred to as sterile if its subtended flower is rudimentary or
One-keeled prophylla appear in inflorescences that embrace lacking. In some bamboos, the basis for a clear distinction between
structural features intermediate between the extremes of determin- empty sterile lemmas and immediately adjacent empty glumes is
ate and indeterminate branching, as in the transitional middle not easy to establish.
part of the inflorescence of Thamnocalamus spathiflorus for example, The internode preceding the first sheathing appendage (usually
the prophylla are sometimes one-keeled and very narrow, and do an empty glume) of the spikelet is designated a pedicel. In pani-
not completely enclose the branch primordia on which they are· culate and racemose determinate inflorescences (Indocalamus sinicus;
inserted. In such cases, the branch promordium apparently does Aulonemia queko) the pedicel usually is fairly long. In some determi-
not have the true resting stage that is found in typical indetermin- nate inflorescences (the spicate raceme of Arthrostylidium cubense)
ate inflorescences, where it is always completely enclosed by a the pedicels are extremely short. In indeterminate inflorescences
two-keeled prophyllum. (made up of pseudospikelets) the spikelet has no otherpedicel
In flowering axes having one-keeled prophylla, the counterpart than the very short uppermost internode of the rachis. Being
of the other half of a two-keeled prophyllum usually appears on covered by the uppermost bract of the rachis, this pedicel is visible
the approximately opposite side of the axis, at the second node only upon dissection of the pseudospikelet.
(following a very short internode), and the second structure is Gommon agrostological usage designates as florets the units
usually one-keeled like the first. This pattern, and the independent into which aspikelet breaks up when the rachilla segments disar-
origin (from distinct primordia) of the two parts of the divided ticulate. But regardless of whether the rachilla segments are ab-
palea in Streptochaeta (see Page 1951:28 and Figs. 1 and 2), support scissile or not, a floret consists of (1) a lemma, (2) the rachilla
the theory that the vegetative prophyllum andits reproductive segment immediately above the node which bears the lemma that
homologue, the palea, may have had their evolutionary origin in embraces it, and (3) a flower, consisting of a branch of the ra-
the fusion of two adjacent sheatlling organs, made possible by the chilla subtended by the lemma .and embraced by a prophyllum
suppression of the intervening internode. The spontaneous suppres- called the palea. Besides the. palea, the floral axis bears lodicules
sion of culm internodes in Arthrostylidium schomburgkii is verified (when these are present), stamens, or a pistil (or both). The floret
on p. 43. thus includes structures from axes of two orders. The floret should
110 THE BAMBOO PLANT

be clearly differentiated from the flower proper in descriptive

statements. The form and organization of the spikelet, floret, and Stig.
flower of Phyllostachys nidularia are very effectively illustrated by
L
Porterfield (see Figs. 52 and 53).
lt appears that the incidence of two empty glumes at the base
of the spikelet is generally accepted by agrostologists as a norm
for the interpretation of spikelet structure in the nonbambusoid
grasses. When only one empty glume is present in aspikelet, it is
interpreted as being the second, or upper one, of two. Each of
the sheathing structures above the second empty glume is con-
ventionally called a lemma, whether it subtends a flower or not.
In some bamboo genera with a determinate inflorescence (Arundi-
naria and Sasa, among others): the spikelet generally conforms to

c
Fig. 52. Details of the spikelet in Phyllostaclrys nidularia, as illustrated by Porter-
field (1926:Figs. 3, 4b, 7, 8). (A) Single spikelet. (E) Diagrammatic sagittal sec- Fig. 53. The floret and flower of Phyllostachys nidularia, redrawn after Porterfield
tion of spikelet, showing arrangement of the florets, of which 1 and 2 are func- (1926:Figs. ~ 5, 6). (A) Floret in cross seetion. CB) Floret (anterior aspect) with
tianal, 3 and 4 are rudimentary and sterile. (C) Plan Slf spikelet in cross section, lemma. removed (p~bescent pale~ with notched ape.-..:: seen at back). (C) Flower
shoVl;ing florets 1-4. CD) Longitudinal section of tip of spikelet, showing the (postenor aspect) Wlth dorsallodicule removed and the anthers and stigmas cut
rudimentary third and fourth florets. All much enlarged. L~ lemma; P, palea; o~. All much enlarged. L, lemma; P, palea; Lod.~ lodicule; Stam., stamen; Stig.~
G, glume; R rachilla segments. stIgma.
114 THE BAMBOO PLANT REPRODUCTIVE PHASE 115

buds (cf. Munro 1868:Pl. 5, Figs. 4 and 5, and p. 153). In the high and very variable (Beddome 1873:235 records 50-60; Gamble
vegetative axes of the plant, and in the spikelet itself, the insertion 1896:125 records up to 120 in a single fiower of Ochlandra traDan-
of the appendages is two-ranked, and distichous-alternating in a con"ca). Even when small, the number of stamens is sometimes
single plane. The axis of the fiower, on the other hand, has its irregular (Pseudosasajaponica, with 3, 4, or 5; Dendrocalamus hamiltonii
appendages inserted in whorls of three. The first whörl distal to with 6 or 7). Astamen consists of two parts: the anther and the
the palea consists of lodicules-typically three, rarely two, some- filament. The filaments may be threadlike and free (Phyllostachys),
times lacking (in most known species of Gigantochloa and in some fiat, and free (or more or less coherent at first as· in Bambusa
species of Dendrocalamus and Schizostachyum), occasionally more Dulgaris), or fused into a tube, in species of Schizostachyum (as
numerous (as in Ochlandra traDancorica). The lodicules vary in size Neohouzeaua), Oxytenanthera, and Gigantochloa. Normally the filaments
and texture, but are generally small, thin, and diaphanous, with elon!!ate
b
at anthesis and thrust~the anthers out of the
.
fiower. Each
venation irregular and often obscure. They are often noticeably anther consists initially of four locules (pollen-mother chambers)
unequal in size and shape. The posterior one that is addorsed to in two pairs. (cf. Arber 1934:Figs. 201-210). The two pairs are
the palea is generally symmetrical and narrowly lanceolate. The joined through most of their length by the connective. The con-
anterior two, located near the margins of the palea, are in same nective is sometimes prolonged above, inta a smooth, or penicil-
genera broadened asymmetrically (subsemiovate) and paired. I late, appendage between the two apical lobes of the anther. The
have encountered no reference to this characteristic as a sign of apical lobes of the anthers, usually subacute and rounded, are
zygomorphy in the bamboo fiower. The appearance of zygomorphy sometimes nipple-shaped (Dinochloa scandens), or divergently cor-
is made even more noticeable in the fiowers of a number of bam- niform and (exceptionally) twisted (Fig. 26, 12). As the pollen
boo genera (Chusquea and others) by the occurrence of only two matures, the two locules of each pair unite to form a sac. After
stigmas, these being matclied with the paired lodicules. the anther is exserted and pendent, the pollen spills out of each
In some bamboo species the anterior lodicules are thicker in sac through an apical pore (Fig. 49) or, in some bamboos, by way
substance than the posterior one and become swollen and turgid of a lonab
slit that follows the suture between the adjacent edges
at anthesis. By pushing the lemma away from the palea, this of each pair of locules.
causes the fioret to open (Bambusa multiplex). The "forked" lodicules The gynoecium, or pistil,is referred to by Arber (1934:163)
of Bambusa arundinacea (Arber 1934:Fig. 106, F), are so characterized as "simple and uniform" in members of the grass family. On
because of their appearance in cross section. They are said (ibid.: p. 152 of the same work the author says, "When we turn to the
117) to have a "fish-tail-shape," and are examples of the form gynaeceum we ... receive no help from the ontogeny in deciding
sometimes assumed by these structures after anthesis. As they lose how many carpels are present ... The mature structure, however,
their turgor and the fioret doses, one edge of each lodicule is harmonizes perfectly with the descriptive convention that it con-
caught and pressed between the lemma and palea. What was a sists ofthree carpels uni ted edge to edge." And elsewhere (ibid.: 120)
"belly" in the turgid state may become a thin fiap in the dry state, Arber states that "the gynaeceum, though best interpreted as
as often seen in herbarium specimens. In some species of Schizo- formed by three carpels, has an ovary with only a single cavity,
stachyum with one-fiowered spikelets, and elsewhere, lodicules are with one ovule attached to its back wall." In the gynoecium, three
lacking, and the fiorets do not open at anthesis. Hackel (1881) more ·or less dearly distinct categories of component structures
discusses the lodicule complement of several bamboos and other are externally discernible: ovary, style, and stigma. The ovary is
Gramineae. the basal and basic structure. Its shape usually changes rapidly
Next in order above the lodicules come the stamens, which during the early stages of its growth. The diversity thus produced
appear typically in one or two whorls of three, according to the in the various specimens gives rise to discrepancies between dif-
genus. However, in Ochlandra, the number of stamens may be very ferent descriptions and different interpretations of the form
 116 THE BAMBOO PLANT REPRODUCTIVE PHASE 117

characteristic of the ovary; and of the fruit, of a given species. The shorter and sometimes is divided almost to its base. In such ex-
ovary has been described as stipitate (stalked) in some bamboos. treme cases, as in Bambusa multiplex, for example, the three branches
(Phyllostachys; see Siebold and Zuccarini 1843:746; Munro 1868:36), are sometimes interpreted as three styles. Arthrostylidium cubense
but this is an error due to misinterpretation of the dried form exemplifies a condition common in its genus, in which the style
of an immature state of the ovary (often found in herbarium is very short and bulbous, and bears two somewhat elongate
specimens) in which, owing to the thickening and induration branches, each terminating in a stigma of the form here designated
of the apical part of the pericarp, this part retains its size and by a new term, bottlebrush. Stigmas are variable in form as
shape, while the soft lower part of the ovary shrinks to asiender between genera as weil as between species within a genus. Their
stalklike form upon drying. N akai interpreted this same phenom- branching often varies within a given species, and teratic mani-
enon in Bambusa as a thickening of the base of the style, and used festations of several kinds are common. Floral histogenesis in
it incorrectly, along with thorniness, another character shown Bambusa arundinacea is effectively described and illustrated by
variably in some species, to separate this genus from Leleba (cr Barnard (1957:2-7, Figs. 1-12) in what is apparently the pioneer
McClure 1946b: 106). contribution to our knowledge of this aspect of the Bambuseae.
In most herbaceous grasses the ovary bears two styles separately Perfect fiowers are the normal reproductive ·e,,--pression in most
inserted, while in the bamboos the ovary narrows at its apex bamboos. However, offrequent occurrence are two states in which
into a usually single stylar column, or style (for alternative a secondary polygamous condition is produced:
descriptions of the style, see p. 287f). In external aspect, the style (1) In many bamboos one or two so-called male fiowers, in
commonly appears as an upward extension ofthe ovary. However, which the gynoecium is absent or rudimentary, appear in the
~s Holtturn has suggested, the relation of the style and the ovary transition zone just above the empty glumes at the base of the
lS fully revealed only by a study of their inner structure. Holtturn spikelet. These are followed by perfect fiowers, as in Bambusa
(1956a:Fig. 9) illustrates a completely hollow style in Oxytenanthera multiplex, ofwhich (as B. nana) Munro (1868:90) says, "The lower-
(species not designated); and in Schizostachyum brachycladum (ibid.: most 1 or 2 and the uppermost 2 or 3 fiowers imperfect (either
Fig. 10) a style that is hollow except for a central strand of tissue. male or female), the intervening 3 to 6 fiowers perfect." Of Phyl-
Arber (1934:120) refers to this centralstrand as the "stylar core," and lostachys heteroclada, Oliver (1894) says "the upper fiorets of each
illustrates it (ibid.:Figs. 47 and 48) as it appears in Ochlandra seti- spikelet appear to be staminate. "
gera, O. beddomei, and O. stridula. The style may be greatly eiongated (2) In the one-fiowered spike lets that terminate the depauperate
and unbranched, with a single long stigma, as in J)endrocalamus pseudospikelets that sometimes develop late in the life of an
st~ctus. Here the style is slender and fragile above; ·only the indeterminate infiorescence (Schizostachyum lima) the gynoecium
th1cker, harder basal part persists as a short point on the fruit. In may have become obsolete, while the androecium is either weil de-
Phyllostachys, Ochlandra, and Schizostachyum the greatly elongated veioped or more or less rudimentary (see p. 100). Arber (1934:133)
and more or less hardened style terminates in three short mentions other examples of secondary or fortuitous polygamy as
stigmas (four Or five in some species of Ochlandra). Here, the occurring in Gigantochloa (as Oxytenanthera) albociliata and Gigantochloa
style persists on the mature fruit as a long beak (Fig. 55). This maxima. In the determinate infiorescence of Puelia and Atractocarpa,
character is often correlated with a tightly convolute and some- the lowermost fiower in aspikelet may be purely staminate and
what hardened condition in the lemma and palea. In such cases the uppermost purely pistillate.
the floret i~ terete, or fusiform, with aminute apical opening
through WhlCh the anthers and stigmas are exserted. In Bambusa The fruit
Dulgaris, the style is elongated and apically delicate (as in Gi- The bamboo fruit (Fig. 55) is indehiscent, and the single seed
gantochloa), and may bear one, two, cr three stigmas at its apex. usually fills the pericarp completely. In external features the bam-
In other species of Bambusa the style is commonly very much boo fruit embraces a wide range of forms: a caryopsis furnished
 REPRODUCTIVE PHASE 119

with a thin pericarp and shaped like a grain of wheat but larger
and with the short, two- or three-pronged stylar column (or the
base of it) persistent at the apex (Arundinaria and arundinarioid
genera); a similar fruit with the pericarp thickened at the apex
only (Bambusa, Dendrocalamus, Nastus); a long-beaked structure with
either a moderately thickened hard pericarp (Cephalostachyum and
Schizostachyum), a thick, hard pericarp (Ochlandra), or·a very thick,
tough pericarp (Melocanna baccifera); a subspherical chestnutlike

. 7. D. strictus. Original; based on fruits collected by Dr. L D. Cleme~t in

March 1957, from plants cultivated at ~he'Atkins Institution ofHarvard Vruver-
sity, Soledad, Cuba; X 2"%.
8. Dendrochloa distans. Original; based on fruits from India, supplied by the
V.S. Department of Agriculture, under P.l. 117532, in an early 'stage of germina-
~XI% .
9. Elytrostacftys clCaJigera, dorsal and lateral aspects. Original; based on fnnts
from EI Recreo, Nicaragua, collected by McClure (No. 21478); X 1:Y;.
10. Gigantochloa nigro-ciHata. Redrawn from Kurz 1876:PL II, Fig. 16a, b,- X 1%.
11. Guadua aculeata, dorsal and lateral aspects. Original; based on fruits col-
lected from plants cultivated at Chocolä, Guatemala, by McClure (No. 21591);
X3.
12. Melocanna baccifera.. Original; based on fruits supplied by Denis Koester
from plants cultivated at Rosario, Alta Verapaz, Guatemala, under P.l. 164567;
XY:,.
13. Nastus elegantissimus. The fruits not seen. Copied from an unpublished
sketch by J.S. Gamble (as Oreiostachys pullei Gamble) based on one of ten.fruits
collected by KAR. Bosscha (s.n..) in Java (Koorders 1908:130); courtesy ofthe
Keeper of the Herbarium at the Royal Botanical Gardens, Kew; X 2%.
14:. Ochlandra travancorica. Original; based on fruits supplied by the V.S.
Department of Agriculture under P.l 190905; XL
15. Oxytenanthera alyssinic~' dorsal and ventral aspects. Original; based on
fruits from Eritrea, supplied by the V.S. Department of Agriculture, under P.L
22776; X 1' /,.
Fig. 55. Bamboo fruits, drawn by EImer W. Smith. 16. Phytt~stachys pubescens, ventral aspect. Redrawn from Muroi 1956:PL 49,
1. Bambusa blumeana, dorsal aspect, showing the hilum or "sulcus." Redrawn,
Fig. 1 (as Ph. heterocycla var. pubescens); X%,. .
slightly modified, from Kurz 1876:Pl. II, Fig. 14b; X2:Y;. 17. Pseudosasa japonic~ ventral and lateral aspects. Reruawn from MurOl
2. B. multiplex; dorsal and lateral aspects, showing hilum and embryotegium, 1956:PI. 51, Fig. 12; X2'/.. .
respectively. Redra"WIl from Muroi 1956:Pl. 49, Fig. 2 (as Leleba multiplex); Xl 'h. 18. Melocalamvs compactifiorus (Kurz) Bentham. Original; based on fruits col-
3. B. longispiculata, dorsal and lateral aspects. Original; based on fruits from lected in South Vietnam by McClure in Deceniber 1953; X Yz.
India, supplied by the V.S. Department of Agriculture under P.L 117530; Xl %. 19. Sasa nebulosa, dorsal and ventral aspects. Redrawn from Muroi 1956:Pl. 51,
4. Cephalostachyu,m sp.,- persistent a.us of flower shown at base. Original; based Fig. 11; X3. .
on fruits from Burma, supplied by the V.S. Department of Agriculture .(without 20. Schizostachyum gracile (teste Holttum), dorsal aspect. Redrawn from Kurz
P.l. number); X 1Yz.
1876:PI. II, Fig. 2 (as S. chilianthum); ca. Xly:'. See also Fig. 63.
5. Chimonobambusa marmorea, dorsal aspect. Original; based on fruits collected
21. Sinobambvsa tootsik, dorsal and ventral aspects. Reruawn from Muroi
by Dr. Jisiburo Ohwi (s.n.) near Osawa, Saitama prefecture, Japan, May 15,
1952; X 21'2. 1956:Pl. 49, Fig. 3; ca. X 2'/z. . .
22. Bambvsa copelandii, dorsal and ventral aspects, the latter shoWlng the
6. Dendrocalamus asper. Redrawn from Kurz 1876:Pl. II, Fig. 15b (as Bambusa prominent embryotegium 'at the base. Redrawu. from Raizada 1948:Pl. 1, Fig. C
aspera in legend, Gigantochloa aspcra in the te..xt, p. 221); X4.
(as Sinocalamus copelandiz); X 2.
120 THE BAMBOO PLANT REPRODUCTIVE PHASE 121

fruit with a very short or obsolete beak and a thickish leathery no endosperm. This statement is verified by Stapf (1904:408) für
pericarp (Melocalamus compactiflorus), and numerous variants of at least one species, in an exhaustive study of the deveJopmental
these main types. See also Kurz 1876:266-7, and Holttum 1956a: anatomy of the fruit of Melocanna baccifera (as M. bambusoides). No
Figs. 1-14, 1958:Fig. 10, C-E. comparable description of the fruit of any other bamboo has been
In spite of the striking diversity of external features manifested encountered. Holttum is of the opinion (expressed in personal
by the fruits of different bamboos, and the usefulness of these conversation) that in the genera mentioned by Brandis as having
features for the recognition of some genera, the inadequacy of the no endosperm the embryo has no resting stage, and the endosperm
gross morphology of the bamboo fruit alone for the purposes of is absorbed through the scutellum as fast as it is formed.
comprehensive classification is evident in the weak and inconclusive
role it has played, even when a conscientious attempt has been
made to use it in traditional systems. Recent anatomical and
systematic studies published by Holttum (1956a, 1958a) have
greatly improved our perspectives on the. form and gross anatomy
of the fruits of the bamboo genera of Malaya. Holttum has
brought into focus fundamental similarities and differences that
apparently are of dependable value für characterizirig sQme genera,
and for uniting related genera into groups of major phylogenetic
importance. Further studies of this nature are greatly needed. The
vast majority of preserved fiowering specimens of the known
species of bamboo do not present mature fruits. For this reason,
published descriptions and illustrations cover only very incom-
pletely the fruits of the known genera. A sustained effort to bring
together documented studies of all of the existing examples will
be very rewarding, and will gradually illuminate an important
aspect of the nature and relationships of the different kinds of
bamboos. Usui (1957a) published a study of the anatomy of the
embryo in Sasa nipponica and Arundinaria (as Pleioblastus) chino. The
characteristics of the embryo of the fruits of bamboos of different
genera are also being studied by Reeder (1961, 1962) particularly
in comparison with the embryo of the fruits of other genera of
the Gramineae (cf. also Kennedy 1899). Such studies may reveal
features useful, in combination with other characters, for showing
hitherto neglected affinities, or sharpening the differentiation of
the bamboo genera from each other, and from reJated genera
of the nonbambusoid Gramineae-particularly if they embrace
the formative stages of the ontogeny of the embryo (cf. M. V. Brown
1960:218) as weil as its dormant stage.
Brandis (1907:87) states that in the Indian species of the genera
Dinochloa, Melocalamus, Melocanna, and Ochlandra the ripe seed has
 VEGETATIVE PHASE: THE SEEDLING . 123

just above each of several of the lower sheath nodes of the primary
culm (see p. 79). According to Shibata (1900:444ff), as the prox-
·imal part of each adventitious root matures (in bamboos of Phyl-
lostachys and Arundinaria at least) the ·cells of the epidermis and
some parts of the underlying tissues die progressively and even-
3 Vegetative Phase: The Seedling tually disintegrate, leaving the endodermis as the protective
outer layer. In at least four species of Bambusa, on the other hand,
the root epidermis is described by Shibata as persistent.
The primary culm (developed from the plumule) is a seg-
Published accounts of the Course of development of the bamboo mented axis of clearly negative geotropic reaction, bearing a foliar
seedling are so fragmentary in nature, and are limited to such a appendage at each sheath node. The course of development, and
small array of entities, that scarcely any basis exists for perspectives the mechanism of growth, in a young bamboo culm shoot taken
of either particular or general application. Brandis (1899) pointed from aplant of mature stature (exemplified by Phyllostachys nigra;.
out that in his day the remarkable process of the development of see Fig. 56) are described and illustrated by Porterfield (1930a).
the seedling into a mature clump had not been sufficiently studied The apex of the growing point is protected by many layers of
and, as Arber observes (1934:65), this unfortunately still holds overlapping sheathing appendages (culm sheaths), which are the
true. first lateral organs to be differentiated.
Between the time of germination of the embryo and the ac- The segmented axes of· a bamboo plant elongate principally
quisition of its full complement of vegetative structures, the bam- during a "grandperiod of growth" (Porterfield 1928). This elonga-
boo plant may be called a seedling. Leaving aside detailed char- tion is effected by means of intercalary growth, a process described
acteristics of the embryo as not pertinent to the present account, in great detail by Porterfield (1930b) as it occurS in the bamboo
we may describe the initial stage of the seedling (Figs. 57-66) as culm (see also Holttum 1955:403). In intercalary growth the
consisting essentially of a root (the primary root) and a shoot immature axis increases in length by the elongation of cells ih
(the primary culm). zones of secondary meristem each located just above anode (Fig.
First to emerge, the primary root (developed from the radicle) 56). As defined by Jackson (1949:199), an intercalary vegetative
is asiender, unsegmented axis, cylindrical cr nearly so, with a zone of growth lies between zones of mature tissue. In the elon-
subapical body of meristem that produces new cells both prox- gating segmented axes of a bamboo plant the locus of each zone
imally and distally. The tissue that results from the proximal of intercalary growth is just above the locus of insertion of a sheath.
increase causes the elongation of the root, and that resulting from There is empirical evidence that the sheath may be the origin of
the distal increase makes the root cap. No experimental evidence substances that control, or at least influence, the process of inter-
has been found as to whether the primary root in any bamboo calary growth and possibly also the initiation of root and branch
is actuated in its initial orientation by positive geotropism, or primordia. When Chinese gardeners wish to dwarf a bamboo,
partly or wholly (if only temporarily) by negative phototropism. they remove each culm sheath prematurely, beginning with the
In the zone of active elongation just proximal to the apical meri- lowest, before the elongation taking place above its node is com-
stem, many of the epidermal cells produce root hairs that establish pleted. Upon the removal of a sheath, the elongation above its
effective contact with the ambient medium. Although the primary node ceases. The initiation of branch buds and root primordia
root develops a system of vascularized lateral branches, it is essen- on any segmented axis always takes place withiri a zone of inter-
tially limited in its growth, and is soon overshadowed and super- calary growth, before the tissues lose their meristematic potential,
seded by raots of an "adventitious" origin, whose primordia emerge and while the subtending sheath is stillliving (see also pp. 12 and
 VEGETATIVE PHASE: THE SEEDLING 125

reaching a few inches or at most a foot or so in height. Elsewhere

(as in Melocanna bacciftra, for example) it may reach the phenomenal
height of 18 ft, as observed by Dr. Ernest Imle on a visit to the Fed-
eral Experiment Station in Puerto Rico (personal communication).
In some bamboos the primary shoot (or culm) produces other culms
or rhizomes only from buds at its lowermost nodes (Figs. 58 and 62),
Region oF cell
str-et'chin!f or it mayaiso ranlify freely at upper nodes aswell, to form leafy
branches. The primary culm in some seedlings of Melocanna
bacciftra grown under artificial (fluorescent) light and controlled
temperature (25°C) in quartz-pebble culture irrigated with a
standard nutrient solution, produced lateral branches from buds
at mid-culm nodes before producing secondary culms from their
:zone - -41--~;;J1
IntercaZary· basalbuds. More commonly secondary culms may develop by
tillering before leafy branches appear. Occasionally, a secondary
, shoot may emerge when the primary one is only a few inches long
and still lacks foliage leaves (Fig. 62). The emergence of a rhi-
zome proper is usually delayed until several erect culms have
emerged by tillering, as in those species of Arundinaria whose
seedlings have been studied (Figs. 64 and 65). The first brancb
.Mature ,L to develop from a basal bud of the primary culm of a seedling is
interno""" --+- usually a culm that arises by tillering, without an intervening
rhizome. However, it sometimes happens in a seedling cf Melocanna
baccifera that the first basal branch to develop may be a rhizome
Fig. ?6. Diagrammatic longitudinal section cf a young bamboo culm shoat, (Fig. 61).
shoWlng stag.es of deve~opm.ent and zones of intercalary growth. Stippled areas After the successful establishment of a rhizome axis a seedling
represent .regIons of actlve dlfferentiation and growth of tissues; unstippled areas,
mature tlssues. In the bamboos, all segmented a.xes elongate by intercalary bamboo plant may be said to be adolescent, since as a rule it has
growth. Redrawn [rom Porterfield 1930b:Fig. 1. by this time developed all of the structures that will characterize
it a.s vegetatively mature. However, as a plant increases in stature,
61). In the culms of many banlboos (all known species of Phyl- successive generations of its culms will manifest a gradual change
lostachys and Shibataea, for example), each zone of intercalary in the shape, dimensions, vesture, and texture of constituent parts.
growth is marked at branch-bearing no des by a transverse thick- The attainment of mature stature may require from 3 to 20 years
ening or "supranodal ridge," that is, the culm nade, sensu stric~o or more, depending on the genetic constitution of the plant and
~Fig, 27). At nodes that do. not bear buds or branches, this ridge the nature of its environment. The leaf blades of the seedling may
15 usually much less consplCUOUS, and in some bamboos it may' be smaller than those of the mature plant (Arundinaria teeta) or
be absent, It is lacking throughout or inconspicuous (even at they may be much larger (Melocanna baccifera). The culm sheaths
branched nodes) in the culms of Melocanna baccifera, and in those in small plants may be provided with auricles or' oral setae or
of most known species of Schizostachyum. both. In rare cases these become, progressively smaller in sub-
In the seedlings of some bamboos (species of Arundinaria, Bambusa, sequent orders of culms and, as the plant develops stature, dis-
and Dendrocalamus) the primary shoot is generally rather short, appear altogether (Phyllostachys viridis). More common isthe case
 126 TEE BAMBOO PLANT

where the aurides and oral setae are lacking in the culm sheaths
in small plants but become weil developed in the sheaths of sub-
sequent ordersof culms, as the plant increases in stature (Phyl-
lostachys bambusoides).

The seedling in bamboos with pachyrnorph rhizomes

Bambusa arundinacea Retz. On the postgermination history of seed-
lings of Bambusa arundinacea we possess but few observations. Arber's
account (1934:65), largely a condensation from Brandis (1899:4-5),
folIows: .
It is said. that in the first stages of their existence the young plants
are very dehcate and,· except under the infiuence of plenty of moisture,
they are unable to resist the scorehing effect of the sun's rays; on the
other hand, excess of water about their reets ~auses them to die off rap-
idly. Moreover, they are incapable of competing with the roinor grasses,
by which they are easily and speedily choked and destroyed. Brandis's
account of the stages in their development is that in March, 1882, he
[ound large patches of yeung seedlings [rom seed which had been pro-
duced in 1881 and had germinated during the rains of that year. The
youngest plant consisted of one shoot, about ~ in. long, bearing two or
three leaves at the tip and, below these, a sheath with a small imperfect
blade. Near the -ground the shoot bore a short, membranous-pointed
sheath, at the base of which were two rootlets, about 3 in. long. At a
later stage, several conical side shoot~_ made their appearance, just below
the surface of the ground; they were bent, first dO'WIlwards, then upwards,
and were covered with numerous membranous, white sheaths. These
side shoots, which would ramify later, were the beginnings of the rhi-
zome. They are destined to turn· upward at the tip, thus forming leaf-
bearing sterns, eac4 rooting from the bend. Besides these underground
side shoots, with short internodes, others arose which had moderately Fig. 57. Bambusa multiplex. Seedlings in two stages of development: (left) froit
with radicle and plumule just beginning rapid elongation; (right) fruit with
long internodes, and rooted at the nodes, sending up leaf-bearing sterns
attached young plant showing branchin,g root, and culm shoot with first foliage
from these points also_ In this manner it came about that seedlings, not
leaf. Redrawn from Muroi 1956:Fig. 47 (as Leleba multiplex).
quite a year old, had an underground rhizome of complicated build,
pushing numerous rootlets into the soil, and bearing a number of shoot.s,_
Takenouchi (1932:Fig. 127) presents illustrations of two stages
of which the first to be forrned were short-lived. The other bamboos
which Brandis examined showed a general similarity to Bambusa arundi- in the germination of seeds of Bambusa arundinacea, without textual
nacea. discussion.

A remarkably rapid rate of growth in seedlings of Bambusa Bambusa multiplex (Lour.) Raeusch. (Fig. 57). Muroi (1956:Fig. 47)
arundinacea is recorded by White in connection with his account illustrates two early stages in the development of the seedling of
ofthe longevity ofbamboo seeds stored under controlled conditions Bambusa (as Leleba) multiplex. Muroi's notes, in Japanese, have not
(see p. 203f). been translated.
 128 THE BAMBOO PLANT

Chimonobambusafalcata (Nees) Nakai (Fig. 58). Troup 1921:Fig. 373)

illustrates the early stages of the development of a seedling of
Chimonobambusa (as Arundinaria) falcata to the age of 4 months. Of
this species (ibid.:985) he says only this: "A tufted growth may
commence at an earIy age." At 4 months this seedling had tour
sterns, with a flfth already initiated as a small shoot. This shoot
barely shows, basally, the beginning of the horizontal growth that
will characterize the" successive rhizome axes more conspicuously
as the plant develops more culms.

Dendrocalamus sikkimensis Gamble (Fig. 59). Arber (1934:Fig. 22)

presents sketches illustrating an early stage in the development of
a seedling of Dendrocalamus sikkimensis.

Dendrocalamus strictus (Roxb.) Nees (Fig. 60). Troup (1921:985) de-

scribes the development of the seedling in Dendrocalamus strictus
in the following words:
The plumule emerges in the form of a pointed conical bud with
sheathing scale-like leaves, which rapidly develops inta a thin, wiry stern
hearing single foliage leaves arising altemately at the nodes, the bases of
the leaves sheathing the stern. Meanwhile, fibrous raots develop from the
base of the young shoot. The tufted form of the young plant commences
to show itself at an early stage. This is effected by the production on the
rhizome of successive pointed buds, from which are developed shart rhi-
zomes which curve upwards and form aerial shoots. The buds and rhi-
zomes, and the shoots arising from them, become successively larger
and larger [Fig. 74]. The earlier shoots are thin, wiry, and grass-like;
but subsequently a time comes when woody culms are produced, which
bear some resemblance to the adult culms in form, in the shape of the
sheaths, and in other particulars.
The most dramatic feature brought out by Troup is the man-
ner in which the strong positive geotropism of the neck of the
pachymorph rhizome operates in young seedlings to carry the
successive rhizome axes deeper and deeper into thesoil. Troup
observes: "In this species the new rhizomes of seedlings take a
decided bend downwards before curving upwards to form aerial
Fig.58. Chimonobambusafalcata. Fruit, and stages in the development ofseedlings:
shoots; thus successive shoots, besides being larger than the pre- Ca) froit, in dormant state; Cb) early stage of germination; Ce) seedling 1 month
ceding ones, arise from rhizomes deeper in the ground." This oId, with fruit still attached., showing leafy culm and a second shoot arising, by
behavior stands in striking contrast with the gradualness of the tillering, from a bud at the base of the first culm; Cd) well-rooted seedling 4
months oId, with four leafy culms and a fifth already visible as a small shoot.
manner in which leptomorph rhizomes usually penetrate the soi! Redrawn from Troup 1921:Fig. 373 (as Arundinariafaleata).
130 THE BAMBOO PLANT

A II----plumule

f[:;~!iii?--caryopsis

Plumule~

scutellUm----q
primary
reat :..-_----t;(-

B, B,

f
Fig. 59. Dendrocalamus sikkimensis. (A) Seedling, natural size, the root incomplete.
The fruit or ~'seed"-"caryopsis"-is still enclosed in a glume and the lemma and
palea. As Arber indicates in her caption to the figure, the vigorous first root has
burst through the glume. (BlJ B 2 ) Two views of another seedling, with the glume,
lemma, and palea removed to reveal the plumule, scutellum, and base of pri~
mary root. Redrawn from Arber 1934:Fig. 22.

to greater depths (see discussion of rhizome neck behavior, p. 17f).

Troup adds: .
The rate of,development of the clump depends very largely, even in
the same speCles, ~n the conditions under whic~ it has been grown
[p'. 985] . . . In v'gorous nursery-grown seedlings of Dendrocalamus
stnctus as many as ten shoots have been counted at the end of the first
season : .. Under favorable conditions the shoots of bamboo seedlings
may :he b~ck for . s~me y:ars in succession before the plant finally
estabhshed Itself; thlS IS partlcularly common in the case of Dendrocalamus
strictus in dry situations. Even under more or less favorable conditions
the first shoot of the seedling ~ay die off at the end of the first seaso~ Fig. 60. Dendrocala:mus strictus: (a-g) froit in dormant state, and stages in the
development of seedlings ('under more or less favorable conditions"; (h) seedling
[po 987] ... A clump [of bamboo] may be said to have attained matu- at 1 year; a second culm has ansen from the base of the primary one, the latter
rity" when it commences to produce full-sized culms [po 990]. now dead; (t) nursery seedling 14 months old. Note that the neck of each succes-
sive rhizome a...as carries its rhizome deeper into the earth. The rhizome then
The age of the plant when this point is reached will vary, how- takes a horizontal course, only to turn upward when a change in the physiolegi-
ever, since the manifestations of maturity are as much a function ca! state ef the growing point, induces a changed pattern in the tissues sub-
sequently produced. The axis then takes on the form. of a developing culm, with
lClear thinking requires that we discriminate between vegetative maturity (here
intended) and sexual maturity in bamboos.
negative geotropism and ether distinctive physiological behavior strongly devel-
FAM. oped. Redrawn from Troup 1921:987 and Figs. 374 and 375.
 132 . THE BAMBOO PLANT

of the site where the plant grows, and of forces that impinge upon -Bronch bud
the clump from without, as of the innate potentialities of the
Primory _ _
plant. Deogun (1937:PI. IU) illustrates later stages in the develop-
culm
ing clump (see Fig. 74) .

. Melocanna baccifera (Roxb.) Kurz (Figs. 61 and 62). According to Kurz

(1876:266), the seed of Melocanna baccifera often germinates before Young culm
the fruit falls to the ground. It is a curious fact that, in the long shoa!
period since this bamboo was first made known to science by
Roxburgh (1819), the only published notes on the course of
development of the seedling are the following observations by
/
Troup (1921:1012):

Germination commences vvith the first heaVy- showers of the rainy Frui! - - - -
season, reots and ShOOt5 being produced from the thick end of the fmit;
roots often begin to appear before the fruit falls. The seedlings, unlike Rhizome
those of most bamboos, roake vigorous growth from the commencement. neck
By the end of the first season each fruit will usually have produced about
five shoots, of which the latest may be as much as 10 ft. high; these
shoots are crowded together in a clump. During the second season more
shoots are produced, the clump expands somewhat, and the largest
culms reach a height of about 20 ft. By the fifth season the culms attain
almost their maximum height, but .are still thin and crowcled together,
and it is not until later that they become spaced out with the gradual
extension of the rhizomes. [Troup (Fig. 397) shows aplant seven years
old from seed; see Fig. 82.]
This bamboo spreads to a remarkable extent by its long vigorous
rhizomes. [Actually, the neck is the only markedly elongated part of the
rhizome in this bamboo.] At the last general fruiting in Arakan it was
also observed to spread, owing to the rolling of the heavy fmits down
the hill-sides, to places where it did not e.xist before, and was found
springing up on savannahs and in beds of streams. It does not thrive
weIl under shade, but springs up readily in gaps. Fig. 61. Meloca7/,TI.a baccifera. Fruit with basal portion of att~ched seedl~ng,
As an instance of the great vigour and vitality of the rhizomes, it in the second year of its growth at the Federal Ex~nment StatlOn,
may be mentioned that Mr. W. D. Turner of Hurbanswala, Dehra Dun, Mayagiiez, Puerto Rico, under P.I: 164567. The pnmary culm had
obtained seeds from Assam in 1912, of which six gerrninated successfully reached a height of 5.4 m (18 ft) according to Dr. Ernest .Imle, wh.o
and produced strong plants, which grew and spread rapidlyon moist supplied the annotated specimen. The de-;relopment of th~s plant IS
fertile ground.In 1917 he was able to dig up no fewer than 400 offsets unusual in the great size attained by the -pnmary c:um and m ~he pre-
for transplan#ng elsewhere, after which a fairly large grove of bamboos cocious emero-ence of long-necked" rhizomes. The rhizome neck IS much
'still remained in the parent crop. elongated in btms species, and its development ~shown here in str:mgl.r
geotropic orientation) preceded that _of the rhIzome proper, Whl~h 15
terminal to it. Four of the roots of the culm have been cut off (ordes
For additional observations on the seedling stage of Melocanna with dark centers); four dormant root primordia are shown between
baccifera see p. 125. the two young culm shoots. Fruit, 6 cm in diameter.
 134 THE BAMBOO PLANT

4 5

?:~:.
..
'C' '

6 8

Fig. 62. Melocanna baccifera. Fruits collected by Denis Koester from plants culti-
vated at Rosario, Aha Verapaz, Guatemala, under P.I. 164567: (above) sectioned
[roit, showing the thickness of the pericarp to be as great as the diameter of the
seed cavity; (lower right) froit hearing two shoots. The precocious emergence of
additional shoots from the primary one, befare the development of foliage leaves
(rare elsewhere) occurs frequently in this species. This apparently is related to
the abundance of food stored in the scutellum and pericarp of these fruits as
compared to that found in the small thin-walled caryopses of most other bam-
boas. Drawing by EImer W. Smith, [rom McClure photo. See Stapf 1904.
11 12 13 14
Schizostachyum acutijlorum Munro. Takenouchi (1932:Figs. 127 and 128)
presents illustrations of three stages in the germination of seeds of
Schizostachyum acutijlorum, and two stages of the further development
of the seedling, without textual discussion.
Fig. 63. Schizostachyum gracile (teste Holttum): 1, the fruit, as it falls from the
Schizostachyum gracile (Munro) Holttum (teste Holttum) (Fig. 63). Kurz plant, still enclosed basally by the lemma and palea; asiender prolongation of
(1876:PI. II, Figs. 6-12) iHustrates the early stages of the germin- the rachilla, terminated by a minute rudiment of a floret, is seen at the left;
2, the same, with the lemma and palea removed, now seen as a caryopsis with a
ation of the seed and thegrowth of the seedling of Schizostachyum· conspicuous sulcus, accompanied by the three lodiculesj 3, the same, in longitu-
gracile (as S. chilianthum) to the 35th day. On pp. 267-268 of the dinal section, the endosperm shown in white; 4, the ungerminated seed, in
same work, Kurz supplements the captions to the figures by the profile, showing the protruding embryo; 5, the seed, front view; 6, partial view
of the germinating seed, showing the emerging coleorhiza on the 4th day of
following brief account of the development of the seedling during germination; 7, the same, on the 5th day, showing the coleoptile (a) and coleo-
its first year. It is at some points difficult to follow with confidence rhiza (b); 8~ the same, on the 6th day; 9~' the same, on a smaller scale, including
Kurz's terminology and the sequence of events as he describes the seed, with the first sheath of the primary culm exposed by the removal of a
them in this account: portion of the coleoptile; 10 and 11, two views ofthe same on the 9th day; 12,
the same, on a smaller scale, on the 11th day; 13~ the whole plant, w~th the
Here the lower b1unt end of the cotyledon [coleorhiza, Fig. 63, 6] fruit still attached, on the 15th day; 14~ the same, on the 35th day, shoWlng the
protrudes through the pericarp about the fourth day after sowing, and first leaf blade. Reclrawn from Kurz 1876:Pl. 2 (as Schizostachywn chilianthwn).
 136 THE BAMBOO PLANT

is followed the next day by.its upper part [coleoptile, Fig 63, 7]. Already
the following day the primary rootlet, which is hairy, forces its way
downwards to a considerable length, while the upper part has enlarged
and separated into two equally large lobes which are separated from the
downward growing part by a more cr less distinct constriction. These.
two lobes enclose in their axil the plumule" which is stiff, hairy and
striped and quickly protrudes from between them, as can be seen in
Fig. 9 (3 days later) [Fig. 63, 10, 11]. The subsequent stages of develop-
ment of the young plant are represented in Figs. 10-11 [Fig. 63, 12, 13],
as observed on the "eleventh and fifteenth day after sowing. At the latter
stage the grovvth of the plantlet becomes considerably slower, and
although still connected with the seed, the cotyledon [endosperm?] was
entirely absorbed already before the fifteenth day of [after] sowing, and
thus the young plant is left to itself for further nourishment from the
soil alone. On the thirtieth day after sowing, the halm-sheaths and a
leaf are fully developed (see Fig. 12) [Fig. 63, 14], but instead of seeing
the groWih now accelerated, it becomes considerably slower, so rouch
so, that after a lapse of a year the plants reached only 2-2% feet in
height.

The seedling in bamboos with leptomorph rhizomes

Arundinaria nikkoensis Nakai (Fig. 64). Hisauchi (1949) describes arid

illustrates a few details of the development of the external features
of two seedlings of Arundinaria nikkoensis. The following interpre-
tation of Hisauchi's account is based on a translation, by David
Ray, of the ]apanese text.

Each ge:r.minating seed produced OOt of an a single primary root and

a single upright, leafy culm. In the course of the first year the primary
root sent out many short branches, and the culm sent out a single leafy
branch from each of two successive nodes near the ground. Early in
the second year, an underground bud situated at the upperrnost of
several nades crowded near the base of the culm gave rise to an upright
branch which is interpreted as a culm (culm no. 2). Then, later in the
same year, a bud still lower on the first culm Cnear its very base) de-
veloped into a horizontal axis Ca rhizome) which, after growing 'for a
considerable distance laterally, turned up at the tip to form an un-
branched culm, whose leaves wer:e larger than those of the aerial branches
of the primary culm [Fig. 64, B]. From the proximal bud of the hori- Fig. 64. Arundina:ria niklwensis: (A) 1-year-old seedling; (B) 2-year-old seedling,
zontal a......::is (rhizome) a horizontal axis of the second order arose. Again, showing two leptomorph rhizome axes; (C) subterranean part of a 2-year-old
from the proximal bud of the second culm another horizontal axis arose. seedling, enlarged to show details and relation ofthe segmented.<L,-:es ofdifferent
orders and categories that have developed. The horizontal Or downward-pointing
This latter gave rise, from its proximal bud, to_ still another horizontal
segmented a...xes bearing sheaths are rhizomes. The unsegrnented dovroward-
axis of the second order. pointing axes are ro.ots. Rearranged from Hisauchi 1949:Figs. 1-3_
 138 THE BAMBOO PLANT

The subsequent formation of culms from these rhizomes is not

recorded by Hisauchi, who concludes his notes with tne observation:
1t must be confessed that, even" after such a long period of observation,
the rhizome's manner of branching is yet to be explained.

Arundinaria simonii (Carr.) A. & C. Riviere (Fig. 65). Muroi (1956:Fig.

48) illustrates seedlings of Arundinaria (as Pleioblastus) simonii at
three stages of their early development. At 150 days from germi-
nation, two rhizome axes were already making strang growth, and
showing oblique geotropic orientation. Here, the rhizome itself
performs the depth-regulating function that in bamboos with
pachymorph rhizomes, is effected by the rhizome neck (see p. 17).

Arundinana teeta (Walt.) Muh!. [det. F.A.M.]. Hughes (1951:118) says:

The first rhizome on a seedling [as Arundinaria without specific
identification] was ohserved in August, 1949, on a plant grown from
seed [sown] in 1947. Thls initial root stock [rhizome] elongated for a
distance cf five inches horizontally through a layer cf mulch, developed
three nocles each having nurnerous raots ... and then turned directly
upward inta an aerial leafy stern.

It has been established, by personal observation, that in this

species the transformation of the terminal bud of a rhizome axis
into a culm, as illustrated in the young seedling (ibid.:Fig. 3) is
a character that is retained by the plant thereafter as the normal
form of growth in the rhizome. See Fig. 10, 8 and p. 34.

Sasa nipponiea (Makino) Makino et Shibata (Fig. 66). Usui (1957a)

illustrates the basal portion of a seedling of Sasa nipponiea, still
attached to the fruit.

Recapitulation of significant events in the ontogeny

of a bamboo plant
Fig. 65. Arundinaria simonii. Seedlings: 1, at 10 days; 2, at 30 days; 3, at 150 days
Differentiation between tissues emerging from the distal and the [rom germination, showing at this stage two leptomorph rhizome axes descending
obliquely. From Muroi 1956:Fig. 48 (as Pleioblastus simonizJ.
proximal facies of the apical meristem of the radicle.
Emergence and elongation of the unsegmented radicle to form a
primary. root.
 VEGETATIVE PHASE: THE SEEDLING 141

Differential development of successive internodes of the primary

shoot in respect to their shape and dimensions, the character
of the foliar appendages, and the incidence ofaxillary buds.
Development of plain sheaths (without appendages) at the lowest
nodes (the neck) of the primary culm.
2nd_ Differentiation of a ligule at the apex of sheaths proper inserted
............ at levels above basal ones on the primary culm .
Differentiation of (sessile) sheathblades from the sheath proper
at all nodes (except several basal and several apical ones) of
s.c. each aerial axis. .
Differentiation of foliage leaves with petiola.te blades on sheaths at
several nodes at the apex of each aerial axis (culm or branch)_
Differentiation (in most species) of auricles or oral setae, or both,
at the apex of sheaths proper (except those at basal nodes) on
s ____ _ all aerial axes.
Development of abscission in certain foliar appendages, including
petiolate leaf blades, and in twigs -bearing senile leaf blades_
Development or suppression of (usually solitary) vegetative buds in
the axils of sheaths on all segmented axes.
Fig. 66. Sasa nipponica. Basal portion of very young seedling, with adjacent part Duplication of the primary culm from its basal buds, by tillering,
of the fmit. The author's legend: "2nd., second leaf; Ist., first leaf of the plumule;
clp., coleoptilej cLr., coleorhiza; r., prirnary root; S.c., seed coat; s., scutellum." From
and the differentiation of an important new organ, the culm
Usui 1957a: Fig_ 4_ neck.
Differentiation of leafy aerial branches from mid-culm nodes_
Differentiation of a pachymorph or a leptomorph rhizome axis
Development of root hairs On the "piJiferous zone" of the primary from a basal bud of a culm.
root. Duplication of rhizome axes from (always solitary) buds at nodes
Development of branch roots on the newly matured part of the of the rhizome.
primary root_ Development of culm axes from lateral buds, or the growing tip,
Initiation of sheath primordia from apical meristem in the plu- of a rhizome.
mule and in the primary shoot into which it develops_ Development of adventitious roots at nodes of a rhizome.
Initiation of segmentation (differentiation of nodes and internodes) Development of WaA-Y exudate as epidermal vesture, particularly
following the initiation of sheath primordia from apical meri- on e"-1'0sed parts of segmented axes.
stem in all axes except the roots_ Development of pubescence -as epidermal vesture here and there,
Initiation of branch-bud primordia axillary to developing sheaths_ particularly on sheaths or their blades, or both.
Emergence and elongation of the segmented plumule to form the Differentiation of reproductive axes, of either determinate cr
primary shoot. indeterminate branching, marked by:
Development of adventitious roots from the zone of intercalary Changes in interval and degree of development of segmentation
growth at basal nodes of the young culm_ cf the axes, and changes in the dimen~ions, form, and texture
142 THE BAMBOO PLANT VEGETATIVE PHASE: THE SEEDLING 143

of the axes and of their foliar appendages, from those character- the nature and the sequence of the physiological events that pre-
istic of the vegetative state to those characteristic of the flower- cede or accompany the flowering of the plant, as well as the
ing state (inflorescence). factors that determine the length of the period of vegetative de-
Differentiation of pseudospikelet and spikelet as structural units. velopment; and whether they are innate in the plant, or dependent
Differentiation cf determinate inflorescence branching. upon the fulfillment of aseries of specific stimuli from without,
Differentiation of bracts, glumes, lemmas, paleas, lodicules, andro- or both. See Seifriz 1920 and Wardlaw 1961.
ecium, and gynoecium.
Anthesis, pollination, and fertilization.
Maturing of fruit, or withering of pistil containing unfertilized
ovule.
Withering and drying of the floral parts.
Development of abscission at the nodes of the rachilla axis, or at
the apex of arachis, which releases the partes) of the inflores-
cence containing the ripe fruit.

The principal objective here is the provision of an aid to the

visualization of the more important stages cr advances in the
ontogeny of the individual plant. Since many of these changes
occur simultaneously, or overlap each other, it is not possible to
list them in a linear sequence that corresponds to the chronolog-
ical order of their appearance. In view of the dichotomies that
develop sooner or later as between the respective ontogenies of
plants of the great groups of genera and species (dichotomies
resulting from diversities in the manner of branching of the rhi-
zome anci of the inflorescence, and in the respective timing of
initiation and termination cf growth in various axes, für example)
an attempt to make an exhaustive list of foci of attention (char-
acters) would run the account into a degree of complexity beyond
the limits established for the present work. The repeated occur-
rence of certain of the events (such as the development of a culm
from a lateral bud, or the growing tip, of a rhizome), combined
with the progressive increase in the size cf the culms that emerge
from the successive orders of branches of the rhizome system,
results in the progression of the plant toward its mature stature.
As indicated elsewhere, gametic reproduction in a bamboo
plant usually is consummated only after a relatively long period
of vegetative development, but may, in certain well-authenticated
cases, erupt within a few months \}r a year or so of the germi-
nation of the seed (see p. 276). We are still in ignorance of both
To talk qf the habits if Bamboos, and qf the management qf Bamboo plants,
has little meaning and is of no practical use. Each species
has its own peculiarities and its own requirements . . . and without
Part n Elite Bamboos and Propagation Methods
a re!iable guide, the study qf Bamboos ... would be hopeless.
-BRANDIS (,899:3)
4 Selected Species

lt has been intimated that, from the points of view of both pure
and applied science, more comprehensive studies of the bamboos
are urgently needed. The successful and fruitful execution of a
program of bamboo studies, however, calls for the establishment
of a source of documented materials appropriate to the needs of
each pertinent discipline. Dried herbarium specimens may yield
material suitable for certain studies such as leaf anatomy and the
morphology of the reproductive structures. Fiber-dimension
determinations relating to papermaking require selected and COffi-
parable portions of the culm. Certain studies in anatomy and
cytology require material preserved in liquid. If arrangements are
made to have such material prepared by the general collector,
he should be provided with suitable vials and preservatives and
be given very fuH written instructions concerning the selection,
processing, documentation, and labeling of the material, and how
to pack it for shipment. The documentation ofthe material should
embrace an adequate system of cross references, connecting the
study material with the herbarium vouchers, special notes, and
photographs. Above all, the collector's field number corresponding
to the herbarium .specimens should be associated with all of the
study material taken from the same plant (er. p. 6).
Stern and Chambers (1960) have recently stated the reasons, and
described the procedure, for the use of herbarium specimens
suitable for identification by a taxonomist, as vouchers for the
documentation of wood sampies for anatomical studies. Actually
the principles and requirements described apply to all plant
materials intended for study from the point of view of any botan-
ical disci pline.
For work in physiology, genetics, breeding, propagation and silvi-
culture, living plants are essential. The development and disci-
 148 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPEClES 149

plined application of a routine for the testing and screening of agricultural economy is stressed by Jones and Wolff (1960:57) in
the many bamboos living plants of which are already available the foIIowing words:
in this country is long overdue. There are in our collections bam-
Results will not be quickly or easily achieved. None of our present
boos of evident superiority for various purposes, some of which crops, we can be reasonably sure, attained an established niehe in agri-
were introduced a fuII half-century ago, but which have not yet culture without at least decades of directed endeavor. Many athers have
been subjected to any properly monitored basic investigations. failed to attain crop status perhaps more because of a lack of sustained
Moreover, there are almost certainly in existence elsewhere other research interest than lack of economic potential in the speCies involved.
bamboos that for special purposes may prove superior to any of A carefully planned program, based from the beginning on the best
possible information and with provisions for frequent refinements of
the numerous kinds 'immediately available in the United States.
evaluations of promising species, as new information is brought to light,
Some of these could readily be introduced from countries stilI offers the best formula for success.
accessible and cordial to the free exchange of plants and infor-
mation. Others are to be had only in areas where, for political On studying the accounts of the bamboos included here, the in-
reasons, the accomplishment of such objectives is rapidly becoming quiring mind probably will notice at once the alm ost total omission
more difficult. . of data on yields. It is characteristic of "the backward state of
The importance of establishing, maintaining, and studying existing knowledge of this group of plants that data on annual
living coIIections of bamboos as a means of advancing the knowl- yields from a given site are hardly available for any bamboo-
edge necessary to their taxonomie disposition and their economic even those that have assumed supreme importance in local
exploitation has been stressed by McClure (1935) and Holtturn economies elsewhere in the world. Some very limited yield data
(1958). There exists an urgent need for the development and are summarized below under B. vulgaris, G. verticillata and M.
discriminating application of criteria more suitable than those baccifera. Reliable figures of this sort on other species might
used in the past to select and introduce bamboos for agriculture, afford a key to the economic promise of commercial ventures.
certain domestic industries, and the agencies for soil and wateT Therefore, in the list of suggested studies offered in the following
conservation. Basic studies in the various biolo21cal
o sciences can pages, first place may weil be given to a project for determining,
be fully revealing and fruitful only when living plants of the on a long-term basis, the pattern of the per-acre-per-year yields
species most suitable for the particular objectives are available. of plantings of biometrically optimum size, located on ecological
FoIIowing systematic exploratory studies carried out coIIaboratively sites representative of areas likely to be available in a given land-
over a 10-year period in laboratory and field, along lines adopted use program. Different harvesting regimes, and other devices for
under my direction, a progressive domestic paper company has discovering how to get long-term sustained yields, should be
. receritly developed a plantation of over 3000 acres of selected incorporated.
species of bamboos as a first step in establishing a permanent and In the selection of bamboos for individual treatment here, no
adequate source of paper pulp of proved superiority. attempt has been made to give special emphasis to the needs of
The bamboos treated in the following pages were selected for the domestic situation in the United States. Of the seven species
discussion because they possess features of outstanding scientific discussed, only one (Arundinaria amabilis) is considered to be
interest and industrial promise. Living plants of aII of them are eminently eligible for major attention in this economy. The aim
available in this country. Yet none of them has ever been made has been , rather, to demonstrate and recommend a rational ap-
the object of comprehensive or sustained studies here. The im- proach to theselection, out of the hundreds of available kinds,
portance of a competently directed program of studies in relation of those bamboos that stand out in ways that clearly relate to the
to the general· problem of bringing new crop plants into any needs and the conditions of a given economy.
 150 ELITE BAMBOOS AND PROPAGATION METHODS 1

The brief description of each species is intended solely to convey

a general idea of the size, habit, and over-all appearance of the
plant in the sterile condition in which it would usually be en-
countered. It is not intended as a means of specific identification. ®
Arundinaria amabilis McClure (Figs. 67-70 and 92)
Because of the superior technical properties of its culms, Arundi-
naria amabilis held apre-eminent position among bamboos in
world trade over aperiod of about 50 years, beginning late in
the last century. During this period, this bamboo supplied the
preferred material for split-and-glued fishing rods in England and
America, and for hop poles in Germany. Its culms are still in
demand in the United States for rug poles, and for fine handicraft
productions, induding fishing rods. While this report was being
prepared, an inquiry was received from a leading manufacturer
of fishing rods, who desired to place an order for $10,000 worth
of select culms of this bamboo.
The following properties are responsible for the esteem in which
the culms of Arundinaria amabilis are held: natural straightness,
slight taper, and freedom from branches in the commercial cuts,
stiffness and resiliency (slowness to take a set when held under
strong flexure), lack of prominence in the nodes, and high density,
toughness, and strength of the wood (McClure 1944:38-40, 51).
The high commercial value of Arundinaria amabilis was sustained
by the maintenance, at the source, of a rigorously executed regime
cf selection, processing, storage, and special packing cf the culms
for shipment. Until the supply was cut off by political events on
the Chinese mainland, this bamboo was without a significant
competitor on the world market.

The Plant (Vegetative Characters). Rhizome leptomorph; dump Fig. 67. Arundinaria amabilis: (A) tip of the leptomorph rhizome; CE) mid-culm
open; culms distant, strictly erect to tip, straight or nearly so; node of yaung culm shoat showing tip and base of culm sheaths; (C) base of
branches typically 3 at each mid-culm node, slender, stiff, wiry, initial branch complement at mid-culm nodej (D) leafy sterile twigs; CE) flower-
ing twigs associated on same branch with leafy twig; (F-JI.!) reproductive parts;
the central one dominant, appressed; the incidence of branches (F) empty glumes I (left) and II, from the base of a splkelet; (GJ floret; (H)
and branch buds at culm nodes retreating from the base of the lemma, abaxial aspect; (l) palea, adaxial aspect; (f) lodicules (one ofth: antenor
culms as the plant develops, branches lacking in the lower 1;,', pair at Lift, -and the smaller, posterior one); (K) ~taffien; (L) ~oeclUm; (M)
fruit, lateral aspect (Lift), and dorsal aspect showmg embryoteglUffi. Based on
to % of the height of culms in plants of mature stature; leaf specimens from the type plant, LUBG 1880, in the Lingnan University Bamboo
blades dark green above, paler (glaucescent) below, oblong-lanceo- Garden.
 152 ELITE BAMBOOS AND PROPAGATION METHODS

late, subcoriaceous in texture, durable, the larger ones generally

pendent when mature, with transverse venation conspicuous on
both surfaces.
In a fully established plant, as seen under plantation manage-
ment at Au Tsai in Wai-tsap District, Kwangsi Province, China
(Fig. 68), the dump habit is open, the culms reach a height ap-
proaching 40 ft and a diameter that may exceed 2 in. The lower
half of such culms is only slightly tapered, if at all, and branches
or branch buds are lacking entirely (McClure 1931a).
The chromosome number of Arundinaria amabilis apparently has
not been determined.

Infraspecific Variation. No variants have been described.

Flowering and Fruiting. Flowering in Arundinaria amabilis is cydic

and gregarious within a given population of plants with a common
origin; the length of the flowering cyde is unrecorded. As observed
in a planting established at Canton, China, under LUBG 1880,
the flowering period lasted for 10 years (1929-1939); its termination
was accompanied by a gradual recovery of vegetative vigor. In-
dividual flowering culms eventually died after losing their leaves,
but the rhizomes remained viable. Flowering was active through-
out the period, but fruit production was meager and only a few
hundred mature fruits were discovered, in the course of much
tedious searching.

Distribution and Ecology. As far as published records are concerned,

Arundinaria amabilis is known only in cultivation. The only known
area 6f commercial production is the Kwang-ning District of
Kwangtung Province, and the adjacent part of Wai-Tsap District
of Kwangsi, China. The persistence of the name tonkin cane in
the American trade suggests a possible primordial origin in Tonkin
Province ofIndochina (northern Vietnaam), whence the first sup- Fig. 68. Arundirw.ria amabilis. A typical cluster of characteristic culms in aplant
of medium stature (20 ft taU) in the midst of a commercial plantation at Au
plies of this bamboo to reach the Western market may have come. Tsai, Kwangsi province, southem China. The superior technical properties of
At Canton, China, plants grown in unfertilized "Iateritic" soil the bamboos known in commercial circles as tsingli or tonkin cane have been
derived from granite, with about 75 in. (1880 mm) of rain falling recognizcd in Europe and in the Uni ted States for three-quarters of a century.
Product of the plant known to science as Arundinaria amabilis (McClure 1931a),
in the six warrnest months of the year, and a temperature pattern these canes were prepared for the Western market (McClure 1931b), by disciplined
similar to that of southern Florida, were slow in establishing procedures characteristic of the sophisticatcd craftmanship of China-the old
themselves. During the first 10 years they developed a wide- China, long respected for its mature cultural traditions and its distinguished
artistic productions. McClure photo, 1928.
154 ELITE BAMBOOS AND PROPAGATION METHODS

Fig. 70. Arundinaria amabilis, village scenes at Au Tsai, Kwangsi province, south-
ern China, the principal center of production of culms of this bamboo for export:
(above) the stack of culros at the right represents culls or current overproduction;
Fig. 69. Aru:n.dinan:a amabilis, vicinity of Au Tsai, Kwangsi province, China: the walls of the granary are woven [rom culled culms; the roof is thatched with
(above) bundles of freshly harvested culms being assembled into rafts for trans- the bark of China fir (Cunninghamia lanceolata); (below) sturdy fences woven with
port to the scouring beach; (below) the beach where the culms are cleansed at culled culms guard the sunning yards where hourglass-shaped bundles of culms
onee by scouring with sand. selected for export are being dried in the sun.
 156 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPECIES 157

spreading system of rhizomes whose course was marked by widely ment, with the branches cut back to a length of 3 or 4 in. The
spaced slender culms from 0.3 to 2 m in height. In the eleventh material used embraced an age range of about 1-5 years. As is
year the number of new culms initiated jumped to double what the rule in the vegetative propagation of bamboos, the resulting
it had been previously, and the maximum height of new culms plants consist cf spontaneously rooting new shoots arising from
increased to about 4 ffi. pre-existing buds. Success depends upon keeping the cuttings under
After repeated failures, living plants of Arundinaria amabilis were conditions that will awaken dormant buds and cause the new axes
successfully introduced into the United States in 1936, accessioned to root.
by the U.S. Department of Agriculture under P.l. 110509, and In 1954, I saw at the Bogor (formerly Buitenzorg) Botanic Gar-
established in cultivation at the U.S.D.A. Plant Introduction den living plants of Arundinaria amabilis that had been propagated
Garden at Savannah, Georgia. These plants have sustained, with- from seeds I sent there from Canton, China, in 1937. The retarded
out damage, temperature minima that frequently reach 17°F, and state of the development of these plants suggests that the climate
culms 30 ft tall were measured in 1959. prevailing at Bogor may be unsuitable for this species.
This species has been established in cultivation at the McIlhenny
Jungle Gardens, Avery Island, Louisiana; at San Andres, EI Sal- Suggested Studies. Arundinaria amabilis should be given a place in any
vador; at the Imperial College of Agriculture, Saint Augustine, field trials that may be set up in mild temperate or cool tropical
Trinidad, B.W.l.; and at the Federal Experiment Station, Maya- regions. Studies to develop efficient methods for its propagation
güez, Puerto Rico. on a large scale are in order, especially since it has proved rel-
atively refractory in the conventional large-scale procedures. Suc-
Propagation. From the single plant available at Mayagüez in 1948 cessful development of a superior procedure for its propagation
(Fig. 92) about 200 plants were propagated by means of clump would increase the likelihood that large-scale, or at least widespread,
divisions and rhizome cuttings. These were used to initiate three cultivation of this bamboo might develop eventually. This species
experimental field plantings in Puerto Rico: at Mayagüez (sea should also be included in studies aimed at the artificial combi-
level) and at two field stations, Maricao (elev. 2000 ft) and Toro nation of desirable technical and ecological characters by con-
Negro (elev. 3000-3500 ft). Owing to more adequate rainfall and trolled hybridization.
soi! fertility prevailing at the latter two stations, the plants estab-
lished themselves promptly, and have shown excellent growth. Bambusa vulgaris Schrad. ex Wendland (Figs. 5, 7, 71, 72, and 99)
Arundinaria amabilis has proved very refractory to propagation
by means of rhizome cuttings without attached culms. The best Bambusa vulgaris is a bamboo remarkable for a number of reasons.
results have been achieved by the use of clump divisions with one Among these are the high strength of its culms and their adapt-
or more culms attached to a relatively long section of rhizome. ability to a wide variety of uses, the readiness with which the
The larger the propagule, the more quickly it will establish itself. plant responds to propagation by vegetative means, the rarity and
Protection from intense insolation and wind is important. The restricted extent of the incidence of flowering in plantations, and
practical upper limit of propaguie size is set by decreasing con- the prompt recovery of plants after severe harvesting-even clear-
venience in handling, and by the limited availability of material. cutting of the clump. However, the susceptibility of the harvested
However, Dr. John Creech (in an unpublished study) succeeded, culms to invasion by the powder-post beetle (Dinoderus spp.) was
by the use of controlled temperature and moisture conditions (in rated as the highest of all among a dozen important economic
a propagating box maintaining high atmospheric humidity and bamboos studied by Plank at Mayagüez (1950:8). This suscepti-
with sand as the medium) in securing rooted plants from cuttings bility limits its value for many conventional purposes, but its rep-
consisting of culm segments each bearing a full branch comple- utation is redeemed by pulping studies that rate Bambusa vulgaris

~~~~-~~~-----------------------------------------------
 SELECTED SPECIES 159

very high among nearly 100 species selected from those available
in the Western Hemisphere. 1 According to the records of C. T. B.
Ezard, the General Manager, in February 1946, of the now defunct
Trinidad Paper and Pulp Co., Ltd., well-established plants of Bam-
busa vulgaris produced, at Saint Augustine, Trinidad, QVer 4 tons cf
pure, dry cellulose pulp per acre per year on a 3-year cutting
cyde. Indications are that this could have been substantially in-
creased by the use of a longer cutting cyde (McClure 1948:735).
After trying many devices (induding posts of galvanized iron
pipe, and stays of galvanized iron wire, as weil as supports made
from the culms of several species of bamboo) for the prevention
of lodging in their banana plantations, the United Fruit Company
selected Bambusa vulgaris as the most satisfactory source of banana
props in its Central American plantations.

The Plant (Vegetative Characters). Rhizome pachymorph; dump

caespitose, rather open; culms erect or suberect, generally more cr
less curved, commonly 20-50 ft, rarely to 60 ft tall, and up to 4
in. in basal diameter, cornmonly hearing a ring cf reets at several
lower nodes, and sometimes up to weil above the middle of the
culm; branches unarmed, several at each node, the middle one of
each complement strongly dominant; in large plants the branch
bud may remain dormant at several of the lower culm nodes; leaf
blades narrowly to broadly lanceolate, concolorous or nearly so on
the two surfaces, the transverse venation sometimes visible, espe-
cially on the abaxial surface.
According to Hubbard and Vaughan (1940:30), the somatic
chromosome number of Bambusa vulgaris is 72.
5

1 Infraspecific Variation. Kurz (1876:339-340) lists as varieties of Bam-

89101112
busa vulgaris the following color forms distinguished by the Malays:
"Tlle natural species,' bamboo hower hedyoo [aur hijou, in current
Fig. 71. Bamhusa vulgaris: 1, leafy twig; 2, flowering branch bearing tufts of
pseudospike1ets; 3, seetion of a yaung culm shoat; 4, culm sheath, aba.xial aspect;
Malay, teste Holtturn ], also called bamboo hower gullies and
5, yaung brauch cr small culm (green-striped yellow form); 6, top of leaf sheath bamboo ampel, with uniformly green culms and bTanchlets; bam-
and base of leaf blade; 7, spikelet; 8, empty glumcj 9, lemma; 10, palea; 11, boo hower kenneng, also called bamboo koonieng or yellow
lodicule; 1~ stamen; 13, gynoecium. Redrawn from Gamble 1896:Pl. 40. bamboo, with culms uniformly yellow, or rarely with an occasional
one gr,een with yellow stripes; bamboo hower seh-ah, also called

lUnpublished results of tests by technicians of Champion Papers, Ine.

160 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPECIES 161

Fig. 72. Bambusa vulgaris. 1. When a representative clump of 60-ft culms (from
the grove in the background) growing on alluvial land at the Raheen Estate,
Jamaica, was clear-cut, the culrns racked up e.xactly a cord, and weighed 2576lb
in the fresh, green statc (data courtesy of Champion Papers, Irre.). 2. Base of a
well-developed clump showing that in large plants thc branch huds may remain
dormant at several of the lower nodes in culms of mature stature. 3. A elose-up
view featuring the sheaths that clothe thc internodes of the yaung culms while
they are in a growing condition. AY, each internode reaches its full development,
an absciss layer forms at the base of the shcath that clothes it, and the sheath
gradually dries up and falls away. 4. The type form of the species, with green
culms, gives rise by spontaneous mutation to several color forms in which yellow
occurs alone or in combination with green stripes. These forms are generally
rather unstable, and the color pattern often reverts to green in new culms of the
same clump. In the commonest form (the one with typically green-striped yellow
culms), individual culms may be almost pure yellow, with only a suggestion of
green (lefl) or an occasional one may be half green and halfyellow (right). Speci-
mens photographed in EI Salvador. 5 and 6. B. vulgaris, cultivar "Wamin," cv.
nov., is characterized by a shortening and basal inflation of the internodes,
particularly those of thc lower part of the culm. The habit photo (5) was taken
at the National Botanic Garden, Calcutta; the near view (6) represents aplant 5
in the Royal Botanic Garden, Port-of-Spain, Trinidad.

6
 162 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPEeIES 163

bamboo kooda, with culms green beautifully striped with yellow; Bambusa vulgaris thrives under a wide range of soil and moisture
bamboo tootool, or blotched bamboo, with culms green at first conditions. The Inter-American Institute of Agricultural Sciences,
then becoming blotched with black on aging." The first three at Turrialba, Costa Rica, has a clump growing weil on an island
apparently are unstable, and reciprocal reversion takes place spon- in a small lake, where the water table is within 1 ft of the surface
taneously between them. According to personal observation, the of the soil. On the other hand, in EISalvador, the green-striped
green-striped yellow form predominates on the Central American yellow form thrives in areas where the dry season is so severe that
mainland, while the green form predominates in frost-free areas the plants become completely defoliated. In continental United
of continental United States and on the islands of the Caribbean States, the plant is common only in frost-free .or nearly frost-free
area, at least in Cuba, Puerto Rico, J amaica, and Trinidad. parts of Florida .and California, but it abounds in the populated
Bambusa Dulgaris, cultivar 'Wamin' cv. nov. (Fig. 72, 5, 6) is parts of Puerto Rico where, since its introduction early in the 19th
distinguished from the type form of the species principally by the century, it has been an important source of building material for
greatly shortened and basally inflated internodes in the lower part temporary use. In the Spanish colonial period, strips split from
of the culms. This bamboo was first described by Brandis (1906:685) the mature culms were effectively used, and proved very durable,
under the Burmese vernacular name Wamin, without assigning in a special lath-and-plaster construction of house walls. U nless
it to a genus, though he alluded to the resemblance of the culm given special protection, however, it is soon destroyed by xyloph-
sheaths to those of Bambusa Dulgans. Camus (1913:135) listed it agous insects.
under Bambusa as "B ? Wamin Brandis," and simply repeated
Brandis's English description in French translation. This provi- Propagation. The propagating of Bambusa vulgans is discussed under
sional name is invalid. The geographical origin of this apparently various headings in Chapter 5, and illustrated in Figs. 94 and 95.
quite stable form (referred to by Brandis as possibly a deformity)
is still a matter of conjecture. It may be assumed to have origi- Suggested Studies. Because of its vegetative vigor, ease of vegetative
nated in cultivation, however, since the native horne of Bambusa propagation, wide ecological tolerance, the ability of established
Dulgaris, from which it clearly is derived, is still unknown. clumps to withstand severe harvesting, and the high rank given
the culms as a source of paper pulp, Bambusa vulgans should be
Flowering and Fruiting. The flowering and fruiting habits of Bam- included in any program of study aimed at the development of a
busa Dulgaris are treated on p. 82. tropical source of cellulose fiber. According to personal experience
in Guatemala, it should be possible, under favorable ecological
Distribution and Ecology. As far as published records are concerned, conditions, to begin harvesting this bamboo, by clear-cutting of
Bambusa Dulgaris is known only in cultivation, although according the clumps, within 5 or 6 years after the establishment of a plant-
to Trimen (1900:314) Kurz regarded it as indigenous in Java and ing. Optimum planting distances and optimum patterns and cycles
Thwaites treated it as a native of Ceylon. The consensus of modern of harvesting have to be determined for each distinct set of field
published opinion is that the native horne of Bambusa Dulgaris is conditions.
unknown. It is quite pantropic in its distribution today, but its Hybridization studies and a search for the causes of self-sterility
penetration into the temperate zones is limited by its vulnera- in the flowers of the common strain, which is the typical form of
bility to cold (its culms may be killed to the. ground at 32°F). the species, could be fruitfuL One or more flowering plants can
The green form has become naturalized in large areas on the usually be found here and there in the wide range of its distribu-
island of J amaica in the wake of a sort of migratory agriculture. tion, at the beginning of the rainy season.
Stakes freshly cut from culms of the living plant and used to
support yam vines take root and, where neglected, produce ex- Additional References. Wendland (1810) contains the original de-
tensive groves. scription and the earliest known illustration of the species; A. and
 164 ELITE BAMBOOS AND PROPAGATION METHODS

C. Riviere 1879:191-203, 328-331; Gamble 1896:43-45, PI. 40;

Moebius 1898; Arber 1934:64, 66, 67 (Fig. 23), 68; McClure
1955:60, Fig. 11.

Dendrocalamus strictus (Roxb.) Nees

(Figs. 55, 7, 60, 73, 74, and 93, 1-3)
Dendrocalamus strictus has assumed great importance in the economy
of India, chiefly as the foundation of that country's paper industry.
It is India's principal accessible source of paper pulp of acceptable
quality. Its pulp is used wholly for blending with inferior pulps
from other domestic botanical sources. By the papermaker's stand-
ards, pulp from Dendrocalamus strictus is inferior in some respects
(fiber length,z for example) to pulp from some other bamboos,
but this species owes its dominance in India's paper industry to
the abundance of natural stands of it within reach of transporta-
tion. Another reason für giving special attention to this species
here is its possession of a property apparently rare among the
bamboos, namely, a high degree of drought resistance (Deogun
1937:83). On this account, it could become an important source
of germ plasm to impart drought resistance to synthetic bamboos
through hybridization. Again, the scope and volume of published
data on the silviculture and management of Dendrocalamus strictus
in India has set it off, in. this respect, from all other bamboos.
Familiarity with the long history of the efforts of the Indian Forest
Service toward the economic conquest of this bamboo should, in

l'
spite of their inconclusive nature (Deogun 1937:vii, 147) provide
a useful background for the design of similarly oriented studies
1 n tf
of other species, in other settings.
In evaluating data published under the specific name of this 1,11
(; 8' 5
(or any) bamboo, it should be remembered that distinct (though
unrecognized and unnamed) strains or clones from which the data
were drawn may differ in important characteristics (see p. 170).
Fig. 73. Dendrocalamus strictus: 1, leafy twig; 2, fiowering branch, bearing heads of
It is weil to keep this consideration in mind also in selecting living pseudospikelets; 3, culm sheath; 4, spikelet; 5, lemma; 6, palea; 7, stamen; 8,
material for silvicultural studies related to culm production for gynoecium; 9, 10, fruits in different stages of devel0pJ;llent; 11, top of leaf sheath
particular purposes. Deogun refers frequently to a strong tendency and base of leaf blade_ Redrawn form Gamble 1895:PL 58'_
toward congestion in the cl um ps of Dendrocalamus strictus. Conges-
tion in this species is due primarily to a shortness of the rhizome
2 An average fiber length of 1.32 mrn was found in culm material studied by tech~
nicians of Champion Papers, Ine.
 SELECTED SPECIES 167

axes, but it may be complicated by an excessive development of

long branches in the lower part of the culms. Congestion in a clump
renders the extraction of culms very difficult. Deogun's opinion
that some strains of Dendrocalamus strictus are more or less com-
pletely free from 'the tendency to congestion is confirmed by my
own observations. Deogun (1937:Pls. N, a, b, and V) illustrates
clumps of Dendrocalamus strictus before and after thinnii:J.g to relieve
congestion.

The Plant (Vegetative Characters). Rhizome pachymorph; clump

more or less densely caespitose; culms glaucous at first, up to
about 60 ft tall and 5 in. in diameter, of variable habit, broadly
arched to erect, strongly curved to fairly straight; branches several
at each culm node, unequal, the central one strongly dominant,
of variable length and habit, buds and branches present at all
nodes or (in full-grown plants of some strains such as P.l. 254921
lacking in the lower Yz to % of the culm; leaf blades of variable
size, small in some strains to large in others (apparently of inter-
mediate size in the type form), usually pale green above, paler
below, typically pubescent on both surfaces, the obscure transverse
venation, visible only in transmitted light, is described by Gamble
(1896:78) as consisting of "pellueid dots."
Richharia and Kotwal (1940:1033) give the somatic chromosome
number of Dendrocalamus strictus from root-tip counts as 72; Par-
thasarathy (1946:234) gives the somatic number as 70, and the
"n" number at metaphase as 35, both numbers based on exami-
nations of root-tip sections.

Infraspecific Variation. Dendrocalamus strictus is a polymorphie species.

Plants in seedling populations vary, apparently both phenotypically
and genotypically. As observed at the Federal Experiment Station,
Mayagüez, Puerto Rico, and elsewhere (without benefit of com-
parison on uniform habitats), seedling populations are seen to vary
widely in ultimate stature and habit of culms, thickness of culm
wall, texture and pubescence of the culm sheaths, branching
Fig. 74. Dendrocalamus strictus. A young clump in the process of formation, show- habit, size of foliage leaves, and disposition toward congestion of
ing the successive development of culms. The oldest one is on the left; the young- culms in the clump. Two named varieties have been based on
est on the right. From Deogun 1937:Pl. IH.
extremes of variation in the pubescence of the flower-subtending
lemmas: var. prainiana Gamble, with lemmas nearly glabrous, and
168 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPECIES 169

var. sericeus Gamble, with the lemmas silky pubeseent. On the It is not dear whieh of these is the typieal form of the speeies
basis of the variegation that appears as silvery white lines on the nor whieh variations are phenotypie and whieh genotypie. Else-
leaf blades and dark-green and yeJlowish stripes on the eulm where (ibid:163) Deogun deseribes in the following words what
sheaths, A. and C. Riviere (1879:681) deseribed a third variant, he apparently eonsiders to be a distinet form, with stable ehar-
"Bambusa stricta var. argentea" (a name not yet transferred to acteristics:
Dendrocalamus). Numerous seedling strains, only one of whieh (P.I.
The bamboo in certain localities, e.g., Nauri, Lansdowne division in
254923) has been formaJly distinguished and doeumented, are United Provinces, has long internodes and is practically free from side
under eultivation at the Federal Experiment Station in Puerto branches to a great height. It is of superior quality and congestion is
Rieo. One of these strains (undesignated) is established at the never seen. Even clumps growing on a camping ground, [which are]
Atkins Institution, Soledad, Cuba, where it began flowering in always rubbed by bulloeks and buffaloes, showed no signs of congestion.
The clumps are quite big, but not eongested.
1956 (Clement 1956) and fruited abundantly in 1957 and 1958.
Deogun (1937 :79-80) has the foJlowing to say about variation Flowering and Fruiting. Deogun (ibid:l07-109) assembled the exist-
in Dendrocalamus striclus as observed in the field: ing reeords of the gregarious flowerings of Dendrocalamus slrictus
Growth forms.-Different conditions of soil, aspect and drainage con- by provinees and loealities, and summarized his diseussion of its
ditions appear to have resulted in the differentiation of recognisable flowering eyde in the following words (p. 111):
growth forms whieh may possibly be more or less inherited through the
seed, though this needs further examination. The general eonclusion is that the eycle for Dendrocalamus strictus in
Three main forms have been found to be recognisable in the course one loeality is more or less constant but differs in localities remote from
of the present study. one another and with appreeiable climatie and soil differenees. More
than two generations ago Kurz noted the [fiowering] eyde [of Dendro-
1. The eommon type. calamus strictus] as between 25 and 35 years (Ind. For., 1876, pp. 256-257),
This type can further be subdivided into three minor vanatlOns; and after the lapse of so many years with a number of fiowerings, we
(a) Culms with moderately thiek walls. This is the ordinary form are still unable to be more preeise. We ean onIy say that there may be
producing mediurn-sized culms and is met with everywhere. a bigger variation, viz., 20-40 years or so.
(b) Culms hollow with relatively thin walls. Generally found in de-
pressions, on cooler aspects, and where conditions are more fa- Sporadie flowering in parts of a eontinuous population, and
vourable. It attains quite a big size. even in parts of individual c1umps, is of frequent occurrence in
(c) Culms solid or nearly so. Generally found growing on ridges and this speeies (Mathauda 1952). Moreover, a number of eases of the
on hotter aspects. On the whole this variation does not attain preeoeious flowering of seedlings of Dendrocalamus slriclus are on
a big size.
reeord. See p. 274 for further details.
2. Large type.
This type of bamboo is met with in forests in the United Provinces, Distribution and Eeology. Aeeording to Troup (1921:1006),
in Bihar and Orissa, and to a limited extent elsewhere, where growth
conditions are optimum. This is the best known, eommonest, and most widely distributed of
This type has practically no side branches to a great height and seldom all Indian bamboos, occurring in deeiduous forests throughout the
shows signs of congestion. It attains a very big size with long, straight, greater part of India, e.xeept in northem and eastem Bengal and Assam;
and smooth internodes. it is eommon in the drier types of mixed forest throughout Burma. It is
found typically on hilly eountry, aseending to 3500 ft and oeeurring
3. Dwarf type. gregariously sometimes almost to the exclusion of tree growth, but usu-
This [type] is of a small size and only exeeptionally forms dumps. ally forming an under-story to, or a mixture with, deciduous trees. It
Typical examples are met with in Balaghat division of the Central Prov- is not uneommon in eertain types of sal forest on hilly eountry. It is
inces, where it is known as karka, and to a limited extent elsewhere abundant in many parts of the Siwalik traet and outer Himalaya from
.. . This is the poorest form of D. strictus. the Punjab eastward to Nepal, occurring most plentifully between the
170 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPECIES 171
Ganges and Ramganga rivers. It is also common in most of the hilly A certain amount of overhead cover is necessary during the earliest
parts of the Indian Peninsula, except in very maist regions. In Burma stages of development of bamboo seedlings before they actually form
it is the typical bamboo of the drier types of upper mixed deciduous clumps. Natural regeneration of bamboo is conspicuously absent from
forest with cr without teak; it also extends inta dry dipterocarp forest. areas entirely exposed to the sun ... A judicious amount of shade min~
This is the hardiest of all Indian bamboos, thriving in, regions which imizes the effect of frost ... The largest number of bamboos is obtained
suffer periodically from excessive drought. Within its habitat it is frost~ when the light falling on bamboo crops is at the maximum. But ... a
hardy. In the abnormal frost of 1905 in northern India it withstood the certain amount of overhead cover improves the quality of the bamboo
effects of the frost better than almost any of the tree species; only isolated at the e.xpense of its quantity.
clumps suffered, and in these only the yaunger and more tender culms
were affected. In the abnormal drought of 1899-1900 in the Indian Penin- Figures from three experimental plots in Lansdowne Division,
sula it escaped damage, although other species of bamboo and many tree United Provinces, quoted by Deogun (1937:92) to document the
species suffered severely. unfavorable effects of shade on number of culms produced do not
take into account the competition of the trees for other require-
Deogun (1937 :79) describes Dendrocalamus strictus as generally
ments of the bamboos such'as water and nutrients that come from
decidumis except on moist sites, adding the following notes (pp.
the soil. The effects of wind are described as unfavorable in pro-
82ft) on its ecological tolerance, without reference, however, to
portion to its intensity. Deogun (p. 163) says that "in areas ex-·
possible infraspecific genotypes or ecotypes that may be involved:
posed to strong winds the individual culms produce more side
A minimum of about 30 inches and a maximum of 200 inches average branches and so become congested."
rainfall, and a maximum shade temperature of 1l6°F, and a minimum Dendrocalamus strictus has been introduced into the Uni ted States,
of 22°F, are recorded from bamboo~bearing localities. Atmospheric hu· on several occasions, by means of seeds from India. 1t is commonly
midity is said to be one of the determining factors in the distribution
of the species. Thus, in Orissa [according tol J. W. Nicholson (Ind. For. found in cultivation in Florida and southern California but is
1922, p. 425) it flourishes in regions where the rdative humidity of the rare elsewhere, exeept in Puerto Rieo, where numerous seedling
air is low, as in the interior tracts which are beyond the influence of sea strains are rather widely disseminated.
breezes, but as the humidity rises, it becomes scarcer ... In general it
may be said that it can stand drought better than any other species of Propagation (Figs. 60 and 93, 1-3). A summary of experience in
bamboo.
1t is a common e.xperience that frost and drought, although of no
the vegetative propagation of Dendrocalamus strictus by various
consequence to grown-up bamboo, may be very harmful to seedlings and methods is given in Chapter 5.
young plants, and this has been confirmed in the course of experiments
on artificial regeneration of bamboo in Denra Dun Division, Uni ted
Suggested Studies. Search for elite strains.
Provinces.
1t appears that, within its dimatic habitat, Dendrocalamus strictus grows
on practically all types of soils, provided there is good drainage. It does Additional References. Arber (1926:458, Fig. 8, A, B; 461) illustrates
not grow on water-Iogged or heavy soils, such as pure day or day mix· the anatomy of the florets, by cross sections.
ture with lime. Well·drained localities with sandy loam overlying boulders Bhargava (1946:Pt. I, 1ntroduction) describes the use of this bam-
are the best. . .
boo for paper pulp in India.
Dendrocalamus strictus prefers hilly ground and is generally more pre·
dominant, and of better quality on cooler aspects. Blatter (1929-30:459ft) discusses possible "causes" of flowering in
bamboo.
Deogun (ibid.:91) quotes from the observations of M. D. Cha- Brandis (1874:569, Pl. 70) presents a description in English, and
turvedi (1928) on the effects of light and shade on Dendrocalamus the first good illustration of the inflorescence.
strictus as shown by an experiment set up by P. C. Kanjilal: Church (1889:283) gives principal constituents of the fruits.
 172 ELITE BAMBOOS AND PROPAGATION METHODS

Gamble (1896:68, Pis. 68-69) presents the first illustration of the

culm sheath; includes a good, brief treatise on distribution and
uses.
Gupta (1952:547) presents a treatise on the flowering habits of trus
species; of interest for comparison with Matbauda's paper on
the same subject.
Kadambi (1949) deals with the ecology and silviculture of bamboo
forests which have been set apart for exploitation of Dendro-
calamus strictus for papermaking.
Mathauda (1952:86) presents abrief but provocaiive discussion of
reeorded data and observations on the highly divergent flower-
ing behavior manifested by different plants (strains?) of this
species.
Mooney (1933, 1938) makes interesting observations on the loeal
distribution of Dendrocalamus strictus in Orissa and in western
Singhbhum, India, as affected by pedologieal faetors.
Plank (1950) reports results of eomparative studies of this and
eleven other bamboos in respect to faetors influencing attaek and
control of the bamboo powder-post beetle (Dinoderus minutus).
Rebseh (1910): "The chief object of the present article is to draw
attention an.cl invite discussion on the management and work-
ing of bamboo forests in a locality where tbe demand is great
and the working eonsequently intense. The past history of these
forests is reviewed, and the gradual evolution of the present
working method, and changes in system of management are
diseussed." Exp. Sta. Rec. 23:644 (1910).
Riviere (1879:675) presents tbe first deseription giVing detailed
attention to vegetative eharaeters of the plant (as Bambusa
stricta).
Trotter (1922) eompletes the reeord begun by Troup (1921) of the
development of plants of Dendrocalamus strictus from natural
seedlings (from seeds that germinated in 1911) to exploitable Fig. 75. Gigantochlaa verticillata: 1, leafy twig; 2, flowering branch hearing tufts cf
age. pseudospikelets; 3, top cf culm sheath, dorsal aspect; 4, top cf culm sheath, lateral
aspect; 5, pseudospikelet; 6, spikelet; 7, lemma; 8, palea; 9, androecium, showing
Gigantochloa verticillata (Willd.) Munro, at least in part3 (Figs. 75 and monadelphous st"amens with stamen tube; 10, stamen; 11, gynoecium; 12) top of
leaf sheath'and base of leaf bladej 13, lorucule. Redrawn from Gamble 1896:Pl.
76). Syn.: Bambusa verticillata Willdenow, Linn. Sp. PI. 2:245 (1799). 52.

Among tbe twenty-odd introdueed speeies that eonstitute the

3That part which pertains to plants accessioned by the V.S. Departroent of Agri~
culture under P.l. 79568.
 SELECTED SPECIES 175'

Fig. 76. Gigantochloa verticillata. (ToP) A plant of mature stature, with culms up
to 65 ft taU and 5 in. in diameter, that died from an unknown cause.. The habit
of culms and clump are more clearly shown here than in a plant in fuH leaf.
Taken at the Agricultural Experiment Station of the Tela Railroad Company,
Lancetilla, Honduras. (Middle) A yaung, vigorous clump with culms up to 35 ft
taU, at thc Escuela Agricola Panamericana, Zamorano, Honduras. (Bottom) The
lower nades of culms of mature stature are characteristically ringed with a
verticil of rather prominent racts and root primordia. Plant cultivated at Rosa-
rio, Alta Verapaz, Guatemala, under P.L 79568.

2
 176 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPECIES 177

principal known bam boo flora of Java, Giganwchloa verticillata on the upper surface, pubescent at first and later glabrescent on
ranks third in importance (after G. apus and Dendrocalamus asper) the lower surface; transverse veinlets distant, irregularly spaced,
as a source of building material and edible shoots. Since, according oblique, weakly manifest, on the lower surface only, as a rule.
to Heyne (1950:300), the culms are durable only as long as they The chromosome number of Gigantochloa verticillata apparently
are not e>'l'0sed to the weather, the standing that this species has has not been determined.
acquired in Indonesia is due not to the durability of its culms,
but rather to their versatility and their good natural shape, straight- Infraspecific Variation. Gigantochloa verticillata apparently is a highly
ness, and workability. Heyne says that in Java, Bali, and else- variable species. Ochse (1931:Fig. 204) following Gamble (1896:61)
where the posts, walls, ceilings, and rafters of houses are made from illustrates as the typical form Awi gombong, which is charac-
the culms of this bamboo, when the more durable species are not terized by green and yellow striping of the lower culm intemodes.
available. Ochse (pp. 323ff) ascribes to it as minor variants three bamboos
Gigantochloa verticillata is included here, however, primarily be- under the vemacular names Awi andong, Awi leah, and Andong
cause of its remarkable qualities and performance in relation to kekes (Figs. 205-207). Heyne (1950:300) lists and weakly char-
papermaking, brought to light only recently. During the course of a acterizes these and several other variants. However, the taxo-
survey of the pulping properties of bamboos available in the West- nomic disposition of the numerous forms of Gigantochloa under
ern Hemisphere, this bamboo stood in general performance among cultivation in Java and adjacent areas is in need of critical study.
the top half-dozen of about a hundred species tested comparatively
in the laboratories of Champion Papers, Inc. But it is in the field Flowering and Fruiting. Although the flowers of Gigantochloa verticil-
(at Rosario, in the Polochic Valley, Guatemala) that it made its lata have been described, the fruit has not. Evidently plants of this
most spectacular showing. One plant, clear-cut after growing in species flower rarely and sporadically, while fruit production, as
the nursery 3 years from the time it was transplanted from the in many bamboos known only in cultivation, is sparse or nil.
propagating bed (that is, 4 years from the time the single-culm
offset from which it originated was prepared) showed a total in- Distribution and Ecology. Gamble (1896:62) says of Gigantochloa
crement of substance in the harvested culms corresponding to a verticillata:
yield of more than 4 tons of oven-dry cellulose per acre per year Wild, or more usually cultivated, in the Malay Peninsula and through-
from plants spaced at 7 X 7 m. 4 out the Malay archipelago, probably extending northwards to Tenas-
serim; cultivated 'in the Calcutta Botanic Garden.
The Plant. Rhizome pachymorph; clump caespitose; culms closely
spaced, exceeding 15 m (50 ft) in height and 10 cm (4 in.) in di- However, it appears that its precise origin has not yet been identi-
ameter, erect, nodding apically; the nodes perceptibly fiared and fied and, according to Holtturn (personal communication),
fringed with halrs at the sheath scar, the lower ones ringed by a Gamble never saw specimens cf this species from the Malay Penin-
rather prominent verticil of roots or root primordia just above sula. Specimens [from this areal reported by Roxburgh las this bamboo]
the sheath scar; internodes plain green, remarkably cylindrical, were another species. '
at first appressed pubescentwith dark hairs, later glabrescent; the
In fact, the typical form of the species may no longer exist in a
wood (culm walls) tapering upward gradually, commonly from a
primeval state. Holtturn (1958:4) says:
basal thickness of about % in; branches more or less completely
lacking in the lower 'h to % of the height of the culm in plants 1t is fairly clear that the roain center cf distribution of Gigantochloa
of mature stature; leaf blades large, oblong-Ianceolate, glabrous lies north o'f Malaya, probably in Lower Burma., where no intensive
study of bamboos has been made. So that the Gigantochloas of Java, and
4Data courtesy Champion Papers, Inc. some planted in Malaya, probably came from the Burma region, and
 178 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPECIES. 179
may be interesting records of thc migra.tion of men; but in thc absence strains resistant to rot fungi and xylophagous insects is indicated.
of a good knowledge of the bamboos of Burma, those records cannot be
Studies on the propagation of each bamboo of economic promise
interpreted. It is strange that the most distinctive and most useful Gi-
gantachlaas of Java seern hardly known in Malaya (G. apus and G. maximal are always in order.
... I have seen no native plants of that genus in southern Malaya. The
first appears in thc neighborhood of Tampin (at thc southern end of thc Additional References. Backer 1928:275-276); Holttum (1956b;
Main Range) and as one travels northwards the variety is quite bewilder- 1958:114).
ing, and I believe that hybrid swarms exist.

Holttum intimates that early migrants probably carried with Guadua angustifolia Kunth (Figs. 6 and 77-80). Syn.: Bambusa guadua
them bamboos of mixed heredity. The multiplicity of closely re- Humboldt & Bonpland
lated forms of bamboos found today in cultivationin Java and Among the bamboos native to the Western Hemisphere, Guadua
elsewhere in Malaysia certainly suggests the occurrence of in- angustifolia is outstanding in the- stature, mechanical proper~ies
trogression, or the segregation that takes place in the F z generation (strength and workability), and durability of its culms, and in the
following the union of heterozygous gametes. importance their many uses have given this species in the local
The U.S. Department of Agriculture accessioned, under P.l. economy wherever it is available.
79568, plants of the plain green form of Gigantochloa verticillata . On the occasion of describing the species and glvmg it a
collected by Fairchild and Dorsett at the Sibolangit Botanie Gar- scientific name, Humboldt and Bonpland (1808:65-66) made the
den, Sumatra, in May 1926. This bamboo has become established following observations (here given in free ·translation from the
in cultivation in the Western Hemisphere, at the U.S.D.A. Plant original French text):
Introduction Garden, Coconut Grove, Florida; at Lancetilla, Hon- "In America the bamboos offer the same benefits as they do in India.
duras, in the arboretum founded· for the United Fruit Company Bamlw.sa guadua is used alone far building entire houses [Fig. 78]. The
by Dr. Wilson Popen oe; and at Chocolä and Rosario in Guate- walls are made of the oidest and largest culms; the first layer of the
mala. The account given above, relating to the outstanding per- roof (the foundation of rafters) is made with sn:aller 0r:es, while the
formance of this species in relation to papermaking, both in the second layer is thatched with young branches still beanng leaves. In
using the culms of this plant instead of the hard timber of the taU trees
laboratory and in the field, is based on plants derived from P.l. that often surround it, the native Americans avail themselves ofthe follow~
79568: ing advantages: (1) the ease with which they can be cut ~nd transported
over lonO' distances; (2) the slight amount of labor reqUlred to prepare
Propagation. No mention of any experience in propagating this bam- them, siI~ce they are used either whole or ·split i~ two; (3) the durabi~ity
boo by any means other than conventional clump division has of their wood, whie;h may compare favorably wlth that of the best t~m­
been encountered in the literature. ber; finally (4) open construction of their houses: and ,the protectl;)ll
from the buming rays of the sun afforded by the Wlde, thlck roof, mmn-
Suggested Studies. It would be worth while to assemble all available tains a cool and agreeable temperature during even the hottest part of
variants of Gigantochloa verticillata and other outstanding species of the day."
Gigantochlaa, to pave the way for comparative studies of the per- The diversity of uses and the general importance of Guadua
formance of their cellulose pulp in the paper laboratory, and angustifolia appear to have increased rather than diminish.ed. in
comparative trials of living plants on selected ecological sites and modern times. In addition to supplying almost all of the bUlldmg
under diverse patterns of harvesting. In view of the generally ex- materials for rural dwellings, and other uses mentioned by Hum-
cellent form and workability of the culms of many bamboos of boldt and Bonpland, whole culms, boards, and lath from Guadua
this genus in relation to the requirements for structural and handi- angustifolia are used today in large ·structures such as a~artment
craft material prevailing in certain local economies, a search for houses and factories in CaU, Guayaquil, and other large eitles. The
 SELECTED SPECIES 181

scraps of waste from the large-scale fabrication of bamboo boards

and lath are used by the bakers of Cali as fuel to heat their ovens.
The scaffolding that supports the operations of building, painting,
and repair of tbe highest structures is erected with the tall, slender,
strong, and resilient culms of Guadua angustifolia. The rafts by
which many of the culms are transported to the markets by river
are also utilized for shipping farm produce and for carrying passen-
"erS (Fig. 79). Although not ranking with the best, the cellulose
~anufactured from the culms of Guadua angustifolia makes very
good paper. Large quantities of the culms are imported by Peru,
particularly for use in building purposes in the city of Lima. These

A
Fig. 77. Guadua angustifolia: CA) rhizome, with base of its culm, and thc necks of Fig. 78. Guadua angustifolia. (Lift) Culm material of this species "is used alone
new rhizome branches; (B) mid-culm branch complement; (C) apex of sheath for building entire hauses. Thc walls are made of thc oidest and largest culms;
from mid-culm nade, in adaxial aspect, showing thc ligule at thc base of thc the first layer of the roof (the foundation of rafters) is made with smaller ones."
sheath bladcj (D) yaung inflorescence, consisting of two pseudospikelets, thc one In these words, Humboldt and Bonpland (1808:65) described the place of Guadua
on thc right developed from a bud at thc base ofthe ether; (E) diagram showing angustifolia in the economy ofNew Grenada more than l~O years ago: That these
thc relation of thc constituent parts of a primary pseudospikelct; CF) diagram- words remain true today is due to thc superior techmcal propcrtIes and the
matic cross section of a floret. incomparable durability of its culms. (Right) In Colombia and Ec~a~or ~oards
like these, fashioned by opening out large culms of Guadua angustifoha wlth an
axe, replace in a hundred ways the sawn boards of timber woods in conventional
applications. McClure 1953.
 SELECTED SPECIES 183

have their ongm chiefly in the littoral of Ecuador, where tl:lis

species is possibly even more important than it is in the economy
of the Department of Valle, Colombia. Everywhere in rural areas
where they are available, one sees the culms of Guadua angustifolia,
either whole or as unsplitlengths, serving as electric power-
line poles, as fence posts, and in the construction of .sturdy gates
and durable woven fences, corrals, bridges, and water pipes. In
Colombla and Ecuador, boards fashioned by opening large culms
out flat replace in a hundred ways the sawn boards of timber
woods in' conventional use elsewhere (Fig. 78). These boards serve
in place of concrete for surfacing the floors of houses, and the
sunning floors where seeds of coffee and cacao are dried. Seasoned
culms of Guadua angustifolia apparently have a relatively high re-
sistance to both rot fungi and wood-eating insects. It has been
observed repeatedly that ordinary hardwoods used in conjunction
with this bamboo have had to be replaced because of insect dam-
Fig. 79. Guadua angustifolia. (Upper) The vast forests of pure or nearly pure age, while the bamboo still remains serviceable. The original un-
stands ofthis bamboo "severalleagues in extent" mentioned by Humboldt
and Bonpland as existing in Quindiu, New Grenada (now the Department treated siding, consisting of boards of this bamboo, in a 40-year-old
of Caldas, Colombia) have in recent decades been severely reduced in plantation house at Pichilingue in the Department of Los Rios,
extent by clearing to establish cattle farms. Very thorny branches form Ecuador, was still in a serviceable condition in 1945, long after
an effective barrier around thc base of the culms in this typical form of
the hardwood floors had been replaced because of insect damage
the species. (Lower) Rafts, by which a largc share of the culms of this bamw
boo are transported to thc markets by river, are utilized for shipping (McClure 1953:37).
farm produce and for carrying passengers as weIl.
The Plant (Vegetative Characters; Figs. 6, 77, 79, and 80). Rhizome
pachymorph, very thick, the neck somewhat elongate; cJump open-
caespitose; culms commonly to 60 ft, sometimes approaching 100
ft, in height, commonly 4-6 in., exceptionally 8 in., in diameter,
erect, broadly arched above; internodes hollow, usually perceptibly
sulcate above the point of attachment of a branch complement,
lower ones very short, the wood up to 1 in. thick at the base of
the culm; culm sheatbs deciduous in the upper part of the culm,
usually more or less persistent at the lower nodes, denseJy and
minuteJy tomentose on the back, especially toward tbe base, with
smalI, brown, persistent, hairs, and more or less densely strewn
with longer, stiffer, coarser, sharp, antrorse-spreading, persistent
or easily detachable hairs; auricJes and oral setae usually lacking
entireJy in the lower sheaths; ligule very variable, usually more
or less strongly convex, sometimes truncate or humped; culm
sheatb blade roughly triangular, about as broad at the base as the
 SELECTED SPECIES 185

apex of the sheath proper, persistent, appressed to the culm.

Branches (in large culms, suppressed throughout the lower y" or
% of the height except the basal 6-10 nodes) solitary and very
thorny at the basal nodes, usually 1 + 1 or 1 + 2 above the mid-
dIe of the culm and progressively more fasciculate above. Leaf
blades extremely variable in size and shape, those on young growth
ovate-lanceolate to oblong-lanceolate, up to 7 X 2 in., those on
old wood oblong- to linear-lanceolate, up to 8 in. long and y" in.
broad, commonly glabrous or nearly so on the abaxial surface,
sparsely strewn with coarse white bristles or more rarely glabrous
on the adaxial surface, sometimes glabrous on both surfaces;
transverse ridges between the veins often visible here and there
on the lower surface.

Infraspecific Variation. A number of interesting variants of Guadua

angustifolia have come to my attention. The Milagro strain, men-
tioned on p. 275, is characterized by annual flowering. Another
strain found at Pichilingue, Ecuador in 1945 under the vernacular
name "caiia mansa" (McClure 1955:152) differs from the typical
form in the following respects: (1) weaker development of the
branches and leaves at the base of the culms; (2) fewer, shorter,
and blunter spines on the lower branches; (3) the generally some-
what broader, shorter leaf blades; (4) stronger tendency of the
mid-culm branches toroot spontaneously. To these may be added
the difference between a reputed 3-year life of service given by
boards made from culms of "cafia mansa" when used to make
dryingfloors, and the reputed 5 year service given by boards made
from culms of the typical form of the species ..
A third strain was observed at finca Santa J ulia, 24 miles south
of Vinces, Ecuador, in 1945 (Fig. 80). This plant is distinguished
most strikingly by the alm ost complete absence of the long thorny
branches that constitute such a formidable barrier around the
base of culms of the common form of the species (Fig. 79, 1).
While stationed in Cuba, Dr. Frank Venning secured from Ecua-
Fig. 80. Guadua angustifolia. A fine clump of a thornless or nearly thornless strain
of this species observed at finca Santa Julla) 24 mi south of Vinces, Ecuador. dor propagules alleged to have been taken from this nearly thorn-
The culms reach an estimated height of 80-90 ft, and a basal diameter of about less strain. These propagules were presented to the Horticulture
5 in. In this plant the culms are relatively free of the long thorny branches- that Department of the Estaci6n Experimental Agronomica at Santiago
make a formidable baITier around the base of the clump in the typical form of
the species (Fig. 79). U.S.D.A. photo by McClure. de las Vegas, near Havana. Ing. Julian Acuiia Gale, Chief of the
Department of Economic Botany at the Station, reports (in letters
 186 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPECIES 187

dated j uly 14 and August 11, 1960) that the two young plants, of rooted plants (see Triana 1950). For results of similar trials at
which are growing weil in Matanzas clay soil, show some small the Federal Experiment Station in Puerto Rico, see Table 5 and
thorns on basal branches of the culms. It remains to be seen text, p. 234.
whether these plants will grow up to be as free from thorny basal In the variant form "cafia mansa" described above, the primary
branches as those observed at finca Santa julia. branch at mid-culm nodes, and to some extent the secondary
branches, bear root primordia on their swollen, rhizomelike base.
Distribution and Ecology. Native to northeastern South America, In trials in Guatemala in 1950, I found that this bamboo could be
and extending into Panama in a cultivated state at least, the propagated readily by means of cuttings consisting of intact p.ri-
typical form of Guadua angustifolia is especially common in well- mary branches alone, or whole branch complements, from mld-
watered, fertile regions at elevations below 5000 ft, particularly culm nodes.
in Colombia (where it is known by the vernacular name "guadua")
and Ecuador (where it is known as "caiia brava"). Parodi (1936: Suggested Studies. Representatives ofthe available variants of Guadua
235), reports the spontaneous occurrence of this species in Ven- angustifolia should be included in any silviculture studies that may
ezuela, Brazil, Paraguay, and as far south as northern Argentina. be carried out in tropical regions to discover plants of superIor
Through the agency of the V.S. Department of Agriculture, capacity to store the sun's energy in a form of special value in the
the typical form of Guadua angustifolia has been established in human economy. judging by personal observations made in its
cultivation in Florida, but the plants suffer from frost injury at native habitat, it seems likely that Guadua angustifolia may embrace
temperatures below 27 or 26'F and are cut to the" ground at forms outstanding among the bamboos in their tolerance of poor
temperatures 2 or 3 degrees lower, while at 17'F they may be drainage, and in the natural durability of their culms.
killed outright (Young 1946b:360). Comparative studies of the chemistry and physiology of sterile
The natural habitat of Guadua angustifolia corresponds to supe- plants of the typical form of the species with fiowering plants of
rior farm land. The vast forests of pure or nearly pure stands the annually fiowering strain should yield information of value
several leagues in extent mentioned by Humboldt and Bonpland in relation to the problem of inducing fiowering in Guadua an-
as existing in Quindiu, now the Department of Caldas, Colombia, gustifolia and perhaps in other bamboos by artificial means.
have in recent decades been severely reduced in extent by clearing Comparative studies of the pulping qualities of bamboos grow-
to establish cattle farms (Fig. 79). The rhizome appears to be very ing in the Western Hemisphere should include all available strains
persistent. According to ranchers interviewed personally, repeated of Guadua angustifolia.
cutting and burning in at least three successive years is required
to eradicate the plant.
Additional References. McClure (1944:65-66, Tables 18-25) gives
distribution and availability, source and documentation of test
Flowering and Fruiting. The fiowering and fruiting habits of Guadua
material, with results, and comments relating to performance
angustifolia are treated on pp. 274f.
tests of ski poles made from Guadua angustifolia; McClure (1945c,
1953) gives a picture of Guadua angustifolia as it appears in the
Propagation. When new plantings of Guadua angustifolia are started,
present-day economy of Ecuador and Colombia.
in the area of its natural occurrence, single-culm clump divisions
are used by tradition. In trials of whole culms (aged 1 and 2 years,
and 3 years or more) as cuttings, set up in collaboration with Dr. Melocanna baccifera (Roxb.) Kurz (Figs. 55, 12, 61,62,81-85: and ~9,
Alberto Machado in December 1949 at Chinchina, Colombia, it 1 and 2). Syn.: Bambusa baccifera Roxb.; Melocanna bambusozdes Trm.
was found that only culms aged 3 years or more gave good yields Melocanna baccifera is most widely known in the Western Hemi-
188 ELITE BAMBOOS AND PROPAGATION METHODS

Fig. 81. Melocanna baccifera: 1, leafy flowering branch, showing some cf the
flowers with stigmas exserted; 2, leafless flowering branch, showing some flowers
in anthesis; 3, culm sheath (abaxial aspect); { upper part cf a pseudospikelet
(the prophyllum is lacking); 5, floret, showing androecium and gynoecium; 6,
lodicule; 7, stamen; 8, fruit, much reduced; 9, fruit in longitudinal section.
Redrawn from Gamble 1896:Pl. 109.
Fig. 82. Melocarma baccifera. Aplant 7 years oId, "artificially raised from seed" at
Dehra Dun, India. From Troup 1921:Fig. 397 (as M. bambusoides).
190 ELITE BAMBOOS AND PROPAGATION METHODS

sphere as the bamboo with a large, thick-walled fruit commonly

referred to as "Iike a pear" in size and shape (Figs. 55, 12 and
62; cf. Stapf 1904:PI. 45, Fig. 3). In form and structure, however,
the fruit actually is more like the fruit of some species of Seehium.
In its native horne (East BengaI, Pakistan), Meloeanna baeeifera
occurs in vast, often pure or nearly pure stands whose yield of
culms constitutes the principal.material available for economical
housing. In this area it is also the principal source of withes
for making basketry and matting. In pulping studies made in the
laboratories of Champion Papers, Inc., the culms of Meloeanna
baeeifera were rated among the top half-dozen of nearly 100
selected species of bamboo available in the Western Hemisphere.
For a good many years, earlier in this century, the Annual Report
of the Forest Research Institute at Dehra Dun, India, was pub-
lished on excellent paper made from Meloeanna baeeifera. In 1954,
a paper mill with a capacity of 100 tons per day was put into
operation at Karnaphuli, near Chittagong, East Bengal, to utilize
local natural stands of bamboos, of whose yield Melocanna baeeifera
provides 85 to 100 percent (McClure 1956a: 182); see Fig. 83. The
same report states that
The existing sources of sl,lpply (in areas of which cutting rights are
held by the Pakistan Industrial Development Corporation) comprise ab0l't
696 square miles of bamboo (either in pure stand or admixed with forest
trees) situated in the drainage basin of the Kamaphuli river above the
mill. Thc potential annual yield of air-dry bamboo [from this areal on a
3-year, selective cutting eyde, is estimated at 351,360 tons. Ofthis abaut
85 percentjs the muli [Melocanna baceifera] .
.With regard to the density of pure stands of Meloeanna baeeifera;"
Troup (1921: 1011) records that enumerations carried out in Aiakim
in East Bengal, on two separate plots of 1 acre each, gave respec-
tive totals of 10,575 culms weighing 27,404 Ib and 6,855 culms
weighing 33,248 Ib. The moisture content of theculms when
weighed is not indicated.
In preliminary small-scale field trials, carried out in the Polochic
Valley, Guatemala, two plants of Meloeanna baeeifera increased Fig.83. Melocanna baccifera. (Above) A harvesting station on the Karnaphuli River,
during the first 4 years of their growth from single-culm offseto at East Bengal, Pakistan. Bundles of culms intended for paper pulp are rafted to
the Karnaphuli mill site from various parts of this bamboo-producing area, over
an average rate of 24,714 Ib of air-dry culms (equivalent to about distances ranging from about 5 to 100 mi. (Below) From the same area, millions
6 tons of air-dry cellulose) per acre per year, calculated at a of culms selected for structural purposes are delivered annually to Chittagong,
7 X 7-m spacing (unpublished data developed in collaboration East Bengal, Pakistan, by raft. '
 192 ELITE BAMBOOS AND PROPAGATION METHODS

between the U.S. Department of Agriculture and Champion

Papers, Inc.). The rate of recovery of Melocanna baccifera after clear-
cutting was also phenomenal, especially in yaung plants. However,
no long-range data relating to sustained yields under control!ed
harvesting rates have been developed.
The Plant (Vegetative Characters) (Figs. 61,81,82, and 29,1,2). Rhi-
zome pachymorph, the slender neck strongly elongate (up to 3 ft
long in mature plants); young clumps usual!y densely caespitose
at first, becoming more open on the periphery as they develop;
culms distant, erect or ascending, up to 70 ft tal! and 3 in. in
basal diameter (more commonly 50 ft tal! and 1.5-2 in. in diam-
eter), with nodding tip, the internodes cylindrical, glabrous, to 20
in. long at mid-culm, hol!ow, thick-wal!ed for 3 or 4 ft at the base,
thin-wal!ed above, the nodes marked by a thin sheath scar only,
not at all pulvinate or swollen above the sheath scar, glabrous,
those in the lower 'h-% of the culm usual!y without buds or
brancl,:es in plants of mature stature; branches in each complement
numerous, slender, subequal, the point of attachment of the Fig. 84. Melocanna baccifera. (Above) A prefabricated wa~l f~r a hause is not. a
modern innovation in East Bengal. This traditional verslOn 15 a 10 X 30-ft UUlt,
branch complement small; leaf blades oblong-Ianceolate, glabrous woven from fle..xible strips-flattened-out whole culms-of this bamboo, which is
on the upper surface, glaucous and pubescent on the lower surface called muli in Bengalese. (Below) No special decoration is needed, when the
when young, at length glabrescent except at the tip, the inner edge natural pattern of units of construction has its own aesthetic merit.
densely fringed with slender hairs, the outer edge sparsely spinu-
lose with short, sharp, thick-wal!ed teeth; transverse veinlets
(called "pellucid glands" in the literature) sometimes apparent on.
both surfaces, especial!y in young leaf blades, obscure in older
ones. In shape and texture, the culm sheath is quite distinctive
(see Fig. 81).
No reference to the chromosome number ofMelocannabaccifera
has been found in the literature.
Infraspecific Variation. No infraspecific taxa have been descr'ibed, but
fruits of three slightly different shapes that have .come to my
attention are presumed to be those of Melocanna baccifera. Field
observations indicate that in the vegetative condition the plant is .
remarkably uniform in its morphological and physiological ex-
pression.
Natural Distribution and Ecology. Gamble (1896:119) gives the
natural distribution of Melocanna baccifera as "throughout Eastern
 SELECTED SPECIES 195
·~Pl'...-.
Bengal and Burma from the Garo and Khasia Hills to Chittagong
and Aracan, and again in Tenasserim. In parts of the above region,
and certainly in Chittagong, this is the most common species, and
the one most universally used for building purposes." He adds,
"Major Lewis says that white ants [termites] do not touch it."
The plant apparently is in its natural horne in the Hill Tracts
of East BengaI. I have observed that it thrives there almost equally
on the well-watered sandy clay loam alluvial soils and on the well-
drained residual soils consisting of almost pure sand, even to the
summits of the low sand stone hills. It springs up in practically
pure stands after repeated clearings for agricultural purposes have
destroyed the natural forest. The rhizomes are remarkably tena-
cious of life, and survive the burning in situ of the felled culms,
leaves, and branches. The annual rainfall in the Hill Tracts of
East Bengal where this bamboo thrives ranges from 78 to 120 in.,
Fig. 85. Melocanna baccijera. (Above) This bamboo !las culms so versatile for with a long dry season each year from November to March. Aver-
str:uctural purposes that no.other material is needed-even for nails or hinges or age minimum temperatures reach a low of 50'F in January and a
shmgles-to make a beautlful and comfortable home, such as these in Chitta- high of 77'F in July and August. Average maximum temperatures
gong, East Bengal: Paki~tru:. (Below) At the International Housing Exhibition
held at Ne",: De~hl, I~dl~, m ]anuarr. 1954, the delegation from West Bengal reach a high of 97'F in April and a low of 75'F in December.
presented thlS pnze-wmnmg home, WhlCh has many attractive features. Except- Nursery trials made in Florida with plants of Melocanna baccifera
mg the fr<:mework .of durabl.e hardwo?ds, it was constructed of two locally abun- from fruits introduced by the U.S. Department of Agriculture
dant and mexpenslve materials:· plastlc earth and the culms of the muli bamboo.
early in the 20th century were unsuccessful.
In 1948, I secured propagules from a large clump of Melocanna
baccifera near the central kiosk at the old Castleton Garden in
Jamaica. Fifty of these were sent to Guatemala. Six others were
sent to the U.S. Department of Agriculture, where they were
accessioned under P.I. 164567. U pon their release from quarantine
at Glenn Dale in 1949, the surviving plants were sent to the Fed-
eral Experiment Station in Puerto Rico. In the. somewhat unfavor-
able soil and climate at Mayagüez the development of the plant
has been less than spectacular. The material taken to Guatemala
throve in the rich volcanic soil at Chocola, where the maximum
annual rainfall approaches 200 in. Plants for the field studies on
rate of increase carried out in the Polochic Valley (see p. 190)
were drawn from this SQurce.

Flowering and Fruiting. The plants at the old Castleton Garden and
those established in Guatemala and in Puerto Rico all began to
flower in late 1957, produced a large crop of fruits in 1958, and
 196 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPECIES 197

continued to flower and bear fruit in 1959. The plant is known dicate that its cellulose fiber ranks elose to that of Bambusa vulgaris
to be monoperiodic, however, and all of the flowering and fruiting by the standards of the paper technologist.
stands have doubtless died. Fruits produced in Puerto Rico and
Guatemala have been widely disseminated to centers of agricul- Additional References. Roxburgh (1819:col. 37-38, PI. 213) presents
tural research in tropical parts of the Western Hemisphere. the original description and illustration of the species, as Bambusa
Plants of Melocanna bacciftra established in Honolulu, from an baccifera Roxb. Trinius (1821:43) first described the genus Melocanna
independent introduction of unrecorded history, flowered and on the basis of Bambusa bacciftra Roxb. Stapf (1904:401-425, PIs.
fruited in 1948 and 1949 and died .afterward. This lot of plants .45-47) gives a detailed description and illustration of the ,:,orphol-
must represent a hereditary line distinct from that of the Castleton ogy and anatomy of the fruit. Troup (1921:1011-1013, FlgS. 396-
Garden plant. However, no difference has corne to light, other 397). Kurz (1875) published the correct binomial.
than the date of initiation of flowering, which in this species
heraIds the approach of the end of a life cyele. Ochlandra travancorica (Bedd.) ex Garnble (Figs. 55, 14 and 86). Syn.:
On the basis of other recorded occurrences of gregarious flower- Beesha travancorica Beddome
ing and monoperiodic fruiting in Melocanna baccifera, its life cyele Since the individual plant itself is unimpressive, Beddome's ref-
has been variously estimated at about 30 years (Gamble 1896:120), erence to it (1873:235) as "a magnificent species" and Ga:o::ble's
30-35 years (Kurz 1876:257), and about 45 years (Troup 1921:984). characterization of it (1896:125) as "magnificent and most mter-
Blatter 1929-1930:916) says: "The data at present avaiIable are esting" probably allude to the bizarre and noteworthy ~eatures of
not sufficient to justify any definite conelusions." The plant dis- the flowers and fruits, and perhaps to the vast extent of ItS natural
covered at Castleton Garden, Jamaica, was received there, the head stands.
gardener asserted, "around the turn of the century, as a little Ochlandra travancorica first attracted the attention of the paper
seedling I could hold in my hand." Its flowering and fruiting in world by the remarkable maximum length of 9 mrn to which,
1957-1959 gives it a reproductive cycle of about 60 years. according to Bhargava (1946:App. 1, p. 20), individual cellulose
fibers of its culm tissues attain, and by the vastness and exuberance
Propagation (Figs. 61, 62, and 89, 1, 2). When fruits are locally. of the natural stands that occur in South Travancore and. South
available, they afford the best means of propagating Melocanna Tinnevelly mountains. Its culms were found by the Technological
bacciftra. However, since the seed usually germinates very promptly Laboratory of the Indian Central Cotton Comrnittee to yield, by
upon maturing (Fig. 62), even while the fruit is still in place on the sulfate process, a pulp containing 92.66 percent of alpha cellu-
the branch, procurement of fruits from a distance presents special lose with an ash content as low as 0.2 percent, characteristics
problems. The development of the plant from seed is described on indicatinrr that this pulp· has possibilities for rayon rnanufacture.
p. 132. The vegetative propagation of Melocanna baccifera is treated Same ye:rs aga, it was announced in an anonymous article pub-
brieflyon pp. 216f, and illustrated in Fig. 89; 1, 2. lished in Fibres (8:82-84, March 1947) that a rnill operated by
Travancore Rayons, Ltd., and equipped to produce 5 tons of
Suggested Studies.. Because. of its great vegetative vigor, and the rayon and I 1j, tons of transparent paper daily, had recently been
rapidity with which the plant establishes itself from seeds or single- established in Travancore to utilize the natural stands of Ochlandra
culm elump divisions (offsets) giving high early yields, and in travancorica found there. No recent notice has corne to light, either
view of the versatility of the culms in industrial and handicraft of this rnill, or of the one rnentioned before the turn of the cen-
applications, Melocanna baccifera should be ineluded in any study tury by Bourdillon, then Conservator of Forests in the Travancore
of the development of a tropical source of buiIding material, handi- State. Bourdillon is quoted by Garnble (1896: 126) as saying of
craft materials, or cellulose. Unpublished comparative studies in- Ochlandra travancorica: "lt rnakes a splendid paper, and we have a
 SELECTED SPECIES 199

mill which uses it almost exelusively. The fibre has been pro-
nounced superior to 'Esparto.' Our only difficulty in connection
with it is the great cost of the chemieals required."

The Plant. Rhizome unknown (judged by descriptions of the clump

habit, the rhizome should be pachymorph); the elump (according
to Beddome 1873:235) elose and impenetrable; culms commonly
6-8 ft, but in favorable habitats to 20 ft tall, and 1-2 in. in diam-
eter, straggling (even climbing) except where self-supporting by
virtue of the denseness of the clump habit; branching habit un-
known; intem~des gray-green, rough; nodes somewhat swollen
and marked with the base of fallen sheaths; 1.5-2 ft or more in
length (Gamble), sometimes even to 5 ft and very thin walled
(Bourdillon); leaf blades broadly oblong-lanceolate, 6-18 in. long
by 2-4.5 in. broad, glabrous or slightly rough on both surfaces;
rounded (often unequally) at the base into a thick, broad, some-
what concave petiole, the apex long-acuminate, twisted, bearing
stiff hairs, and often scabrous; transverse veinlets submerged but
often showing on the abaxial surface as "oblique pellucid glands."
(Except as noted, the characters for the foregoing description
were drawn from Gamble 1896:125).
Darlington and Wylie (1956:458) cite "EKJ" Uanaki-Arnmal)
as authority for a 2n chromosome count of "about 72" for Och-
landra travancorica.

Infraspecific Variation. Ochlandra travancorica var. hirsuta, described

and illustrated by Gamble (1896:126, PI. 111) from specimens
collected by Beddome in the Travancore HilIs, is differentiated
from the typical variety by thicker leaves, wi th their margins more
cartilaginous, their sheaths strewn wi th appressed hairs emerging
from bulbous bases; and spikelets thickly elothed with light-brown
velvety pubescence.
Fig. 86. Ochlandra travancorica: 1, leafy fiowe.ring branch; 2, culm sheath; 3, pseu-
dospikelet, showing one keel of the lowermost bract (prophyllum); 4, the one- Flowering and Fruiting. Of the incidence of flowering in Ochlandra
flowered spikelet, with two bracts still attached, and the fiower in anthesis; travancorica, Brandis (1906:685) says: "Beddome collected it in
5, 6, bracts; 7, lemma; 8, palea; 9, lodicules; 10, stamen; 11, gynoecium; 12, 13, flower about 1868. I found it in flower on the Tinnevelli ghats in
stigmas, much enlarged; 14, fruit, still surrounded by the lemma, palea, and
bracts. Redrawn from Gamble 1896:Pl. 111. February, 1882, and in Travancore the species was expected to
flower in 1905. Believed to die down after flowering." Gamble
(1896:126) quotes Beddome, the discoverer ofthe species, as saying
 200 ELITE BAMBOOS AND PROPAGATION METHODS SELECTED SPECIES 201
that "it flowers almost every 7 years and dies down." Fischer Suggested Studies. The culms of Ochlandra travancorica and those of
(1934:1863) notes that it "flowers at long intervals and dies down." its congeners should be studied in the paper-technology laboratory,
Ochlandra travancorica apparently is unique among the bamboos in to determine their papermaking characteristics for comparison
the degree of duplication of its essential floral parts, 'especially the with results already obtained from other bamboos available in the
stamens, of which Gamble reports that as many as 120 have been Western Hemisphere:
found in a single flower. Other known species of Ochlandra show The high maximum fiber length of9 mm reported by Bhargava
duplication of the floral parts in a more moderate degree. In (vide supra) for Ochlandra travancorica certainly suggests 'further fiber
common with other members of the genus the fruit in Ochlandra studies. However, this recorded maximum, and the minimum of
travancorica is characterized by (1) a very thick pericarp that is at 1.0 mm, standing alone, carry little weight with the discerning
first fleshy, later indurescent, and (2) a long stiff basally conical paper technologist. It is only the proportion of fibers falling within
persistent beak. Gamble describes the fruit proper in Ochlandra each length elass that has significance.
travancorica as reaching a length of 2 in., with a beak of equal Also needed are field studies to determine whether Ochlandra
length (Fig. 55, 14). When mature, the fruit retains the rachilla, travancorica is a desirable silvicultural subject, in relation to the
the very persistent glumes, lemma, lodicules, and palea-the whole high- and sustained-yield requirements of a paper mill. The repu-
of the one-flowered spikelet (Iess the stamens )-firmly attached to tedly short flowering cycle of Ochlandra travancorica, and the sub-
its base. sequent death of the flowered culms, may, if true, be a disadvan-
tage to its cultivation as a source of cellulose pulp. However, the
Distribution and Ecology. Beddome (1873:235) says: plant apparently fruits freely, and there is the possibility that the
development of a seedling progeny would be sufficiently rapid to
This magnificent species is most abundant on the South Travancore
restore the stand to a productive state by the time the dying
and South Tinnevelly mountains, [at1 3000-5500 feet elevation, where
it covers many miles of the mountains, often to the entire e..'l:clusion of flowered culms had been harvested-if the stands maintained are
all other vegetation; in open mountain tracts it generally only grows to of sufficient extent to provide several years' supply from a single
6-8 feet in height, but most elose and impenetrable, elephants even not progressive-harvesting cycle. Flowered culms of Dendrocalamus
attempting to get through it; inside sholas [small-ravines] and their out- strictus are usable, and give a slightly enhanced cellulose yield of
skirts it grows to 15 feet high and is much more straggling. 1t is called undiminished quality, after standing in the field 4 years (Deogun
Irul by all the natives, and by Europeans the Elephant grass.
1937:115).
Gamble records (p. 125) that Ochlandra travancorica is planted in
Madras and at Peradeniya in Ceylon. Additional Reference. Gamble (1896:121-128) is the most compre-
Attempts to introduce it into the United States by means of hensive available treatise on the known species of this genus.
seeds have been unsuccessful, but seedling plants sent by Dr. Boshi
Sen from Almora, United Provinces, India, in 1951 and accessioned
by the U.S. Department of Agriculture under P.L 198012 have
been established in cultivation at the Federal Experiment Station
in Puerto Rico.

Propagation. No recorded experience in the propagation of Ochlandra

travanconca has been encountered.
 PROPAGATION 203

either that the event will be discovered or that the availability

of seeds will become known to those most interested. It is at pres-
ent, therefore, usually not possible to seeure, on demand, seeds of
a particular bamboo that may be desired for study or exploitation.
The genetic constitution of bamboo seeds from open-pollinated
5 Propagation wild plants is also unpredictable. It may be heterogeneous, and
often appears to be so, in the light of variation shown in Some
seedling populations. Seeds of Bambusa longispiculata (P.L 93573),
Dendrocalamus membranaceous (P.L 74229), and D. strictus (P.I.
Bamboo heads the list of eleven new crops selected in 1957 by 77061) from -natural sources were introduced by the V.S. Depart-
the President's bipartisan Commission as meriting special attention. ment of Agriculture. The seedling populations were grown to
In the words of the V.S. President's Commission on increased in- maturity at the Federal Experiment Station in Puerto Rico. The
dustrial use of agricultural products (1957:IV-4): "Bamboo has plants of each population showed great diversity in vigor, habit of
been the subject of little research in this country. Major efforts growth, and ultimate stature. Probably present also were variations
will have to be made in every phase of the pro gram if large acre- in features of interest from the technieal point of view. Natural
age production and use is to be developed rapidly." seedling populations of bamboos showing genotypie diversity may
The economic success of large-scale exploitation will depend to afford an opportunity for seleeting, as clones, individual superior
an important extent on the cultivation of elite bamboos, selected plants.
for outstanding quality and high productivity. Such traits will -be Several forms of bamboo "seeds" (one-seeded fruits) are shown
found associated only in an occasional individual clone. When it in Fig. 55. Seedling stages of several bamboos are described on
is realized that the establishment of plantings to supply.a modern pp. 126ff and illustrated in Figs. 57-66.
paper mill of a size suitable for economical operation may eall for
the prompt production of at least a million rooted propagules from Duration of viability in bamboo seeds
a single elite plant, the importanee of developing the most effi-
cient means of propagation becomes clear. The production of Nothing approaching the duration of viability normal to the
bamboo plants of uniform stature and of a size suitable for lab- common cereals has been reeorded for the seeds of any bamboo.
oratory studies will also require the development of special tech- Published information on the experimental storage of bamboo
niques cf propagation. seeds to preserve their viability is, however, very meager. The two
New bamboo plants may be produced by means of seeds, vege- items of experience reported below are purely exploratory.
tative fractions, or layers. Experience has shown that each of these White (1947a) summarizes the results of his studies of seeds of
methods has eertain advantages, and each may in certain circum- Bambusa arundinacea as folIows:
stances be subject to limitations, for the propagation of any
particular bam boo. In these tests the most practical method cf preserving viability of bam-
boa seed was storage Qver calcium chloride at room temperature. Storage
aver hydrated lime cr aver charcoal was also a good method if refrig-
Propagation by means of seeds erated. Drying of the seed to a moisture content of abaut 12% definitely
increased longevity when stored Qver hydrated lirne under refrigeration.
The availability of bamboo seeds is conditioned on unpredictable In ather cases there was little or na advantage gained by drying. When
events and circumstanees. The time of fructification of any given no drying agent was used, exposed seed retained viability longer than
stand eannot be foretold with certainty, and there is nO assurance seed sealed ai~ight.
 204 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 205
Deogun (1937: 11 7) reports briefly the results of experiments on need of studies to ascertain the best season for planting-should
the storage of tbe seeds of Dendroealamus strictus carried out at the seeds of any bamboo beeome available in suflicient quantities. The
Forest Research Institute, Dehra Dun. Seeds showing an initial history of the conditions under which the seeds were stored, from
germination capacity of 56 percent were stored dry in sealed tins. the time of colleetion until they are sown, should of course be a
Deogun does not give either the moisture content of the seeds part of the documentation.
when sealed up or the temperature at which they were stored. The potential praetieal advantages of propagating bamboos by
Germination after 1, 2, and 3 years was 54, 43, and 5 percent, means of seeds of known genetic constitution should not be
respectively. No seeds germinated after storage in gunny sacks for neglected. The artificial induction of flowering and fruiting in
1 year, under conditions not otherwise described. this group is therefore a problem that challenges the creative
The viability of corn seed has been successfully extended by imaaination and merits the active eollaboration of persons suitably
refrigeration at the Plant Industry Station, Beltsville, Maryland, "
qualified and equipped for the task. Until tbis challenge is suc-
and elsewhere. Methods for extending the period of viability in cessfully met, however, the handicaps inherent in the "natural"
bamboo seeds should be extensively tested as material becomes situation will continue to prevail.
available. The establishment of a successful "germ-plasm bank" Dendroealamus strictus-the principal bamboo of economic
of fully documented bamboo seeds would do much to encourage, interest in India-is the one species coneerning whieh somewhat
and make possible, the initiation of a wide range of much-needed extensive notes on establishing plants from seeds have been
descriptive and experimental studies of valuable bamboos. published. Deogun (1937:118-119) gives general directions for the
preparation of sites and treatment of the soi!, botb for direct sow-
Planting the seeds ing (to produce plants in situ, without transplanting) and for
nursery practice. Details pertaining to quantity of seed required,
Hughes (1951:119 and personal conversation) reports that seeds of
shading, irrigation, weeding, and thinning are also given, but
a species of Arundinaria native to North Carolina (probably A.
teeta) germinated when gathered in the late dough stage, if planted
without referenee to loeality. Troup (1921:994) gives a more
circumstantial, illustrated account of personal experience in the
and watered immediately. Fully formed seeds of Chimonobambusa
same matters, acquired at the Forest Research Institute at Dehra
"marmorea will also germinate promptly while still green and soft,
when planted in a moist medium (personal experience). White Dun.
(1948:13) relates his experience with another species in the foll00-
ing words: Requirements relating to sciEmtific and commercial objectives
Seed of Ba:mbusa arundinacea were sown in soil at a depth of %inch and Two specifications, whose importance in relation to the require-
abaut an inch apart in rows 3 or 4 inches apart. Germination occurred ments of scientific or commercial objectives must be recognized,
in abaut one week and the seedlings grew quite rapidly. When the plants are: uniformity of genetic constitution, and uniformity of size,
were 6 to 8 inches high they Were transplanted to individual containers
in tbe plants used.
holding abaut one gallon of soil. Transplanting to the field was done
after the plants were 2'!z to 3 feet in height. Growing the plants from "The chief reason why such objectives call for vegetatively prop-
seed is undoubtedly the most economical and convenient method of agated material is tbe requirement for uniformity of genetic con-
propagating large numbers of plants. stitution. In eontrast with seedling progenies, which are generally
heterogeneous with respect to one or more characters, the vege~
The Rivieres (1879:463) advise that, in tbe absence of a knowl- tative progeny from a given clone may be assurried, in the absence
edge of the viability of bamboo seeds aequired from any souree, of evidence to the contrary, to be identical with the original plant
it is best to plant tbem at once. At the same time, they stress the in all respeets, including genetie eonstitution. Such evidence to the
 206 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 207

contrary has come to my personal attention in only two cases. Both Stations in Latin America. They were to form a nucleus for ex-
involved the appearance of minor spontaneous mutations consisting perimental work on the potential utility of these species of bamboo
of distinctive color patterns on the culms of plants propagated by in relation to the local economy of the respective countries. The
the late David Bisset from rhizome cuttings of the plain green director then in charge of the station at Mayagüez informed me
form of Phyllostachys viridis (P.I. 77257) at the U.S.D.A. Barbour that to fil! this request would exhaust the station's supply of some
Lathrop Plant Introduction Garden, Savannah, Georgia (McClure of these bamboos, and deplete its bamboo budget as weil, since one
1957:64-65). In any case, vegetative propagation is the most re- man-day of labor was required to prepare five propagules (clump
liable means available for multiplying bamboo clones to secure divisions), and the air-freight rates on such heavy propagules as
genetically uniform plants for critical study or exploitation. were then being produced by the station made the cost of shipment
Whether the somatic' chromosome complement in bamboos is by air prohibitive.
always uniform throughout the body of the plant, either in number Collaborativestudies were later carried out at Mayagüez, with a
cr in constitution, is a question that still awaits systematic inves- view to producing rooted plants in the size and numbers required.
tigation. Discrepant records as to somatic chromosome number Well-rooted little plants of several species, weighing from 1 to 4
have been reported for both Bambusa arundinacea and Dendrocalamus ounces, were produced in large quantities. Packages containing 100
strictus, and for hybrids between B. arundinacea and a sugarcane to 200 of such plants, isolated from each other in moist sphagnum,
variety produced at Coimbatore, India Oanaki Ammal 1938:925; and with the roots protected by a coating of colloidal material,
Richharia and Kotwal 1940:1033; Parthasarathy 1946:234). were light enough to be handled and shipped cheaply. Moreover,
Uniformity in size of plants is an important-really essential- the thousands of pla~ts produced made no appreciable inroads on
specification for material to be used for replicated treatments. in the available supply of propagating material. We did not achieve
all experimental studies, whether carried out in the laboratory or the goal of uniform size, and we did not find wholly satisfactory
in the field. Moreover, uniformity of size has a financial value in methods for propagating some of the species selected for trial.
relation to the problem of packing and handling large numbers of Brief progress reports on some of these results were published in
plants. the Annual Report of the Federal Experiment Station for the years
1950-1953, and by McClure and Kennard (1955). Selected details
are presented on pp. 229ff. A standardized procedure for producing,
Small versus large plants at will, very small plants of uniform and specified size, by veg-
The principal advantage oflarge, rooted propagules (such as clump etative means of propagation, remains to be perfected. It seems
divisions) over small ones, is in the economy of time required for likely that the method ultimately adopted for certain species will
them to reach mature stature. Personal experience confirms this include: (1) the use of single-bud propagules with a very much
in situations where the prompt establishment of a few large plants reduced amount of accompanying mother tissue; (2) the use of an
is an urgent objective. However, as the distance through which the artificial nutrient medium with the possible addition of synthetic
plants must be transported becomes greater, certain disadvantages hormones; and (3) the provision of accurately controlled conditions
of large propagules become more noticeable. And, of course, large of incubation, including at least temperature and moisture. As
plants are unsuitable for most laboratory studies. compared with "natural" conditions-that is, where relatively large
The advantages of smal!, rooted plants, and the importance of propagules are subjected to only nominal or partial controls-this
their potential role in experimental studies, seem not to have been level of refinement of methods should reduce significantly the in-
discussed in print. In 1946, I requested of the FederalExperiment fluence upon the end results that ordinarily is exerted by variables
Station at Mayagüez 50 plants each of several species of bamboo such as inna:te differences between species and between groups of
to be shipped, by air, to each of five Agricultural Experiment species, and even differences due to topographical origin of buds
 208 ELITE BAMBOOS AND PROPAGATION METHODS Table 1. Two groups of bamboos, requiring different propagation pro-
cedures.
and time of year when the propagating m~terial is severed from
the mother plant. Characters Expression of characters

Group I Group n
Two groups of bamboos requiring different propagation procedures
Climatic adaptation in Tender plants that Frost-hardy plants that
Most of the relatively well-known bamboos have been observed relation to temperature thrive best under thrive best in
to fall into two large groups that contrast more or less sharply with frost-free conditions; climates wi th a
some species are marked, 'but not
each other in relation to details of procedure appropriate to their known to survive extremely cold,
vegetative propagation. The Rivieres (1879:460ff) were the first to temperatures a few de- winter; a few species
call attention to the existence of this natural grouping of the bam- grees below 32°F can survive tempera-
without serious tures a little below
boos, which they discovered during the course of their studies in QOF without serious
damage
propagation at Harnrna, Algeria. The two groups are characterized damage
by the Rivieres as folIows: Culm initiation (active Takes place typically Takes place typically
growth following during the summer or in the spring, at the
1. Caespitose bamboos, with autumnal growth, and rhizomatous breaking of buds autumn, or at the onset of favorable
branches in fascicles. [Examples:] Bambusa macroculmis, vulgaris, vulga:ris on the rhizome) under beginning of a rainy levels of temperature;
vittata, hooker~ spinos~ stricta, argentea, gracilis, scriptoria. natural conditions season following a apparently is con-
2. Running (rarely caespitose) bamboos, with spring growth and relatively dry period; trolled primarily by
apparently is con- temperature levels
branches paired or in fascicles. [Examples: 1Bambusa or Phyllostachys mitis,
quilio~ nigra, viridi-glaucescens, aurea, jlexuosa, violascens; Arundinan"a simon~
trolled primarily by
moisture levels
japonica, falcata, fortunei. Pachymorph, i.c., short, Leptomorph, i.c., long,
Rhizome
Form of the constitu- thick.; internodes asym- slender; internodes
Some of the species listed in each group now bear other specific symmetrical, longer
ent axes metrical, broader than
names, and are disposed in other genera than those indicated here. long (Fig. 2) than broad (Fig. 3)
The synonymy is listed in the index to scientific names. Form of the lateral Dome-shaped; the apex Boat-shaped; the apex
intramarginal distal
The differences between the two groups as enumerated by the buds Indetenninate
Growth habit of indi- Determinate
Rivieres relate to the particular array ofbamboo species with which vidual axes
they were working, and to the ecological conditions prevailing at Clump habit typically Caespitose; the plant a Diffuse; plant with
single dense tuft of culms distant from
their experiment station. The enumeration of the characteristics of each other
culms
the two groups given in Table 1 is based on personal experience Culm origin Distal to the rhizome Normally lateral to the
acquired through experimental studies carried out in both the rhizome'
Culm branches (mid- Basally swollen; recapitu- Basally not swollen; not
Eastern and the Western Hemispheres, and under both tempentte lating the form of the recapitulating the
culm range)
and tropical conditions. rhizome; the dominant form of the rhizome;
Among the genera whose constituent 'species possess character- ones often bearing root not known to bear
primordia of spontane- root primordia of
istics that prevent their being classified as belonging strictly to ous origin in situ spontan~ous origin in
either of the two groups described above are Chimonobambusa, Chus- silu
quea, and Sinarundinaria, as currently defined. Usually obscure Clearly manifest
Transverse venation
of leaf blades
Bambusa, Dendroealamus, Arundinaria, Phyllo-
Typical genera
Significance of the differences between the two groups m relation Elytrostachys, Giganto- stachys, Sasa, Semi-
ehloa, Guadua, arundinaria, Shibataea,
to vegetative propagation Sinobambusa
Oxytcnanthera
In general, given material of the proper age, bamboos of both
 210 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 211

groups respond most favorably to propagation by means of clump SIZE OF PROPAGULE. The simplest method is the division of a
divisions, since such propagules are, cr may be, in ev«ry respect clump into two equal parts, retaining the root system, branches,
complete plants. Of the vegetative fractions that are not complete and foliage of each part as fully intact as possible. Provided they
plants, rhizomes of the right age range may generally (but not are properly set out, such propagules generally give the highest
always) be counted on to give the greatest assurance of favorable degree of success in terms of survival rates and rates of subse-
response to conventional procedures. Other types of vegetative quent development, and are least exacting in respect to care after
fractions, which range in size from whole culm cuttings to branch planting. However, as the scale on which propagation is needed
cuttings, represent larger or smaller arrays of buds from the aerial increases, the prodigal use of large propagules sets a limit to the
parts of the plant. It is in such propagules that the differences in scale of operations by exhausting the reservoir of materiaL
behavior as between bamboos of thetwo groups are most marked At the other extreme of size in clump divisions are propagules
under traditional methods of propagation, where the control of the each composed of the lower part of a single culm with the rhizome
environment is nominal. axis basal to it. These are designated by Deogun (1937:121) as
When we have learned enough about the physiology of bud "offsets." Offsets have many advantages over larger propagules;
dormancy and awakening, nutrition, and tissue differentiation in namely, ease of preparation, accessibility of material of optimum
the bamboos to be able to provide the right cultural conditions, it age, and economy of material. Their principal disadvantages are
may be possible to establish a new plant from any viable bud from their more exacting demands in respect to care after planting and
any bamboo whatsoever. lower survival rate. Single-culm clump divisions (Fig. 89, 1) are
the traditionally preferred method of propagation for certain bam-
boos: Dendrocalamus strictus (cf. Deogun 1937:121) and Bambusa
Clump Divisions. Clump division is the traditional, and perhaps the tuldoides (see pp. DOOf!). For some bamboos, however (for example,
most generally prevalent, method of propagating bamboos vege~ Bambusa textilis), single-culm propagules have not, in my experience,
tatively. given as good survival rates or yields as offsets consisting of two
culms (2-year-old mother and l-year-old daughter) left attached
SEASON OF PREPARATION. Active growth of young shoots from buds together.
on the rhizome usually is initiated during the summer in bamboos
of Group I, and during the spring in bamboos of Group H. The
commonly recommended practice is to process vegetative propa: AGE OF MATERIAL. In small clump divisions, the age of the rhi-

gules just before the initiation of the annual period of active bud zome appears to be of critical importance.
growth in the axes involved. This applies particularly to clump According to Deogun (1937:121) 1- or 2-year-old offsets of
divisions. Apparently there is some sentiment in Great Britain Dendrocalamus strictus give superior results, while propagules con-
(Thomas 1957:248) in favor of dividing the clump after the buds sisting of material 3 years or more in age give progressively poorer
on the rhizome have begun to push, or at any time when the results.
plant is in active growth. There are no published data of a scien- Hageman et al. (1949) describe the successful use of dynamite
tific nature to support this practice, but it should be investigated, to reduce the labor of breaking up old clumps of large bamboos
since a fuller knowledge of the species involved, the details of for propagation. The indiscriminate use of such material, without
procedure, and the underlying reasons for the practice may reveal discarding the older parts, gave very poor results in terms of sur-
useful leads. Ecological conditions peculiar to some parts of the vival rates in afforestation projects carried out by the Tropical
British Isles may be a critical factor favoring the practice. Forest Experiment Station in Puerto Rico. This is to be expected,
212 ELITE BAMBOOS AND PROPAGATION METHODS

since many of the propagules consisted wholly of material much

more than 3 years old.

PREPARATION OF CLUMP DIVISIONS. The critical point to keep in

mind in separating a clump division from the mother plant is the
importance of.severing the rhizome at the right place in its com-
plex branching system. Since the rhizome system is usually com-
pletely hidden under the soil, the first step to be taken is to leam
the position and orientation of the various connected axes of the
rhizome that are to be included in the propagule. This involves
exploratory excavation. With experience, one learns where and how
to find the place where a cut is to be made.
Clump divisions taken from the periphery of the clump are apt
to give superior results, because they can be selected so as to con-
tain material of optimal age, and because they lend themselves
more readil y to the use of proper methods of extraction.
In the preparation of clump divisions, a distinct procedure in
cutting the rhizome is required by bamboos of each of the two
groups. In Group I, the rhizome should be severed only at one
point-at the neck (Fig. 2) of the oldest rhizome axis in the propa-
gule. The cut should be made at the slender neck in order to
minimize the damage to the rhizome and keep the ra w surface as
small as possible. Moreover, the tissues at this point appear to·have
a greater resistance to decay. This may be due to the presence of
a higher proportion of lignified tissue in relation to parenchyma,
and a smaller amount of stored food, in the tissues of the neck
than in the tissues of the rhizome proper. c
In Group II, either one or two cuts may be needed (Fig. 87, C),
depending upon the orientation of the propagule to the rest of Fig. 87. Propagation cf bamboos cf Group II. Phylfostachys viridi-gla:uccscens~
propagated (A~ B) by means cf rhizomes alone: CA) .r~:zome se~ent ready for
the plant, and the rhizome need not be severed at a neck. the propagating bed, as illustrated by A. and C. RIVlcre 1879:Fl~. 12; CB) the
Once the composition of thepropagule to be lifted has been same propagule, represented (ibid.~ Fig. 13) as the "res~lt" cf tbis method cf
determined, and the rhizome has been severed as described, the propagation. As explained on p. 226, the propagule as :llustrated had not 'y::t
produced .a rooted plant. (C) Propagation by ~eans cf smgle-culm clump dlVl-
aerial part of the propagule may be. pruned before the roots are sion. The details illustrate certain charactenS!lCS cf bamboos 0: ?:oup II that
severed-or promptly thereafter-in order to minimize loss of have significance in relation to the method of prepar~n~ clump d1':'lslQns of bam-
water through the leaves. It is advisable to retain as much foliage boos with leptomorph rhizomes. From A. and C. Rlvlere 1879:Flg. 10.
as may, by ample irrigation and protection of the propagule from
sun and wind, be kept from wilting until the root system is re-
established.
The root system of the propagule should. be preserved as nearIy
214 ELITE BAMBOOS AND PROPAGATION METHODS

intact as possible. The roots are best preserved and protected by

keeping them in a ball of earth when the propagule iso taken from
the mother plant.

PRACTICAL EXPERIENCE-GROUP I. Bambusa tuldoides. Notes on the

course of development, for aperiod of 6 years, of a small experi-
mental planting of l-year-old, single-culm propagules (offsets) of
Bambusa tuldoides (as B. breviflora Munro, Lingnan University Bam-
boo Garden No. 2669) were published by McClure (1938b). Eighty-
seven out of 90 propagules (96 percent) survived. The over-all
annuaI increase in number, and the maximum dimensions, ofnew
culms produced each year, are shown in Table 2.
At the end of the second season the ratio of new culms to older
ones stood at a high of 4.77. The ratio fell off sharply in the
following year, and continued to decline. The rem oval of 167
single-culm propagules in 1933, and 125 in 1934, apparently was
responsible for an appreciable exaggeration of .the rate of decline
in the ratio of new culms to oId, but even assuming that the ones
that were removed would (if they had not been removed) have
produced new culms at the same rate as the ones left behind, the

Table 2. Annual increment of culms from 87 single-culm clump divisions

Fig 88 Propagation of bamboos of Group I. (Lift and right) ~n unidentified
of Bambusa luldoides planted in April 1931 at Canton, China; data from
specles 'of Gigantochloa, illustrated by A. and C. Riviere 1879:FlgS, 6 and 7 (as
McClure 1938b. Bambusa macrocalamis), propagated by means of culm segments: (~efl) c~lm seg-
ment bearing a young branch complement, prep~ed f~r propagation; (r.zght) the
Maximum culm Total Number rinci al branch has developed roots and has gtven nse, from one of l.ts ?asal
dimensions number ofsingle- Ratio of bUds, ~o a young shoot which may become a rooted culm, t~us estabhshmg a
Number of culms culm numbers of
new p 1ant. S ee p. 258 . ( Center) Propagation by means . of
' smgle-culm Ch'
clump
Height Diameter ofnew at year offsets new·culms divisions: Bambusa tuldaides, cultivated at Lingnan Umverslty, Canton, ma,
Year (m) (ern) culms end removed to eId under LUBG no. 2669. Original.
1931 3.0 2.2 87 87' 1.00
1932 6.0 4.0 415 502 167b 4.77
1933 7.5 5.3 348 683 125 b 1.04 . glven
ratlos . . the table
In . cl'or the years 1934 and 1935 would still
1934 12.0 5.8 333 891 .59 have declined progressively. . .
1935 12.0 5.8 281 1172 .31 The performance of a single propagule IS shown m Table 3.
1936 12.0 6.1 193 1365 .16
1937 12.0 6.2 258 1623' .18 A1though this plant performed outstandingly. in the fi~t 2 years,
it showed in later years a steady decrease m the ratIO of new
aThis figure represents new culms only, and excludes the original propagules; the ratio of culms to old that is similar to that shown by the whole ,:,"ray.
new culms to propagules happens to be 1 for the season during which the planting was made.
b Culms rcmovcd in the following March, that is before thc season's production of ncw It appears that, for this species, under the particular condltlO':s
culms of that year began. of this experiment, (1) l-year-old, one-culm propagules of thls
c Counting 292 one~culm propagules removed, thc total number of culms produced is 1915.
 PROPAGATION 217

Table 3. Annual increment of culms from one single~culm clump division the culm, and the long, slender rhizome neck, may be discarded,
of Bambusa tuldoides planted in April 1931 at Canton, China. Data from for convenience. This leaves a propagule scarcely distinguishable
McClure, 1938b. ' from a rhizome cutting. Figure 89, 1, 2 shows a typical propagule,
and the growth produced by such a propagule in 24 months.
Ma'timum culm Total
nurnber Ratio of
Forty-five out of 50 such propagules planted in May 1948 at
dimensions
Nurnber of culms nurnbers of Chocolä in Guatemala survived and produced vigorous plants.
Height Diameter ofnew at year new culms Oxytenanthera abyssiniea, The Nyasaland Forestry Department,
Year (rn) (ern) culms end to old
Zomba (1944:11-12) gives an excellent account of the techniques
1931 3,0 3.0 4 4" 4 developed locally for the successful propagation of Oxytenanthera
1932 5.0 4.6 12 16 3 abyssiniea by single-culm clump divisions.
1933 7.5 5.0 18 34 1.1
1934 12.0 5.8 18 52 0,5
1935 12.0 5.8 21 73 .4
Rhizomes Alone. GRQUP 1. Published knowledge of the details of the
procedures and precautions relating to the propagation of bam-
"'This figure represents new culms only, and excludes the original propagule. boos of tbis group by means of rhizomes alone is very meager. The
following references are typicaL Dabral (1950:313) says: "Th~ ~est
bamboo give excellent results, and (2) the best time to break up method established by comparative experiments and stanstlcal
the plants for further propagation by clump division would be in analysis' by the Provincial Silviculturist of Madras, is by rhizome
the spring, exactly 2 years after the propagules were put in. This planting." The results of these experiments apparently have not
indication should, of course, be checked by repetitions of the ex- been published. Dabral does not mention any species or any de-
periment. tails of the method he recommends, without reservation, as the
Dendroealamus strietus. Deogun (1937:121) says that offsets (clump best,
divisions) provide the only method that has given success in the Ahmed (1956:530) indicates tbat Bambusa tulda is successfully
vegetative propagation of Dendrocalamus strictus. propagated in connection with afforestation pr.ojects ~; S~ngb~um
The offsets should be prepared from one-season-old culms as far as (Bihar, India) by means of rhizomes planted zn sztu, W,th shght
possible and, never from more than 2-season-old culms. It is necessary care 80% success has beeIl achieved; 95% survival is not uncom-
that some portion of the rhizome with a bud be kept and care taken, mon." No details of procedure (for example, age of material,
when planting, not to injure the 'eyes.' The culm may be cut down to preparation of propagules, season of planting, or type of care) are
2' or 3' ... The degree of success attained is very variable, hut may be
as high as 100% , .. The success of the method depends in part on the given.
In the recorded cases where propagation of bamboos of Group
vitality of the rhizome stock used and the time of year when it is planted.
If the rhizomes are taken from young healthy stock and planted immedi- I has been accomplished by means of rhizomes alone, these were
ately at the break of rain, success may be expected, but if the rhizomes planted directly in the field. For this purpose rhizomes have the
are taken from old stock and planted much before the rains, complete advantacre of ereater convenience, being lighter and less bulky tban
failure may result. The weather of the year must also be an important " "
clump divisions. However, systematic trials should be carne . d out
factar. to establish whether offsets may have certain advantages-far ex-
Meloeanna baeeifera. The vegetative fraction of the plant that has ample, in the possible retention of some foliage, and the presence
proved most convenient for the propagation of Meloeanna baeeifera of stored food in the tissues of the culm to nourish new growth.
is a single-culm clump division. These should be made from the My own interest was directed to the use of rhizomes alone for
youngest culms, while the lateral buds of the rhizome are still propagating bamboos of tbis group when I observed the spo,:tane-
dormant, or before they have pushed more than 2 or 3 in. Most of ouS production of small rooted shoots on fragmentary and mutzlated
 218 ELITE BAMBOOS AND PROPAGATION METHODS

Fig. -89. Propagation of bamboos of Group l.

i, 2. Propagation by means of single~culm clump divisions, Melocanna bacciftra.
1. One of 50 propagules brought from Jarnaica in May 1948 for cultivation at
Chocola, Guatemala, under P.1. 164567. 2. Plant developed from a similar propa~
gule in 24 months; in time, the long rhizome necks will give rise to an open
clump habit (see text, p. 36).
3-6. Propagation by means of rhizomes alone. 3, 4. -Bambusa tulda, cultivated
at the Federal Experiment Station, Mayagüez, Puerto RieD, under P.l 21002;
two rhizome cuttings, both removed from the mather plant at approximately
the same age (under 1 year) that have been in the propagating bed for 6 months.
Typically, as shown here, the uppermost bud of each cutting Ca bud actually
borne at the base of the lowermost internode of the culm) awakens promptly
and produces a plant, the first ans of which usually is poody rooted. As a rule,
the buds borne on the rhizome proper awaken more tardily, but produce plants
all culms of which, including the first, are usually well rooted. The divergent
behavior of buds at different levels of the propagule is construed as being
related to differential levels of auxin concentration. The disparity in the inci-
dence (and promptness) of pushing of the buds, in the rhizome proper of these
two ostensibly identical propagules is not understood. Conceivably, it could be
related to an unrecorded difference in the actual age of the two propagules at
the time {)f the removal from the mother plant. 5, 6. Gigantochloa apus, cultivated
at the Federal Experiment Station, Mayagüez, Puerto Rico, under P.l. 99573.
Rooted culms from the top bud of rhizome cuttings (actually thc lowcst bud of
the culm itself) removed at 9 months from the time the rhizomes were put in
the propagating bcd. 5. Plants that were topped and defoliated at 6 months,
while still attached to the rhizome. 6. "Check"-plants that were not topped
or defoliatcd at 6 months. The defoliation of the culms was intended to break
the dormancy of thc buds of the rhizome proper. It did not havc thc desired
3 effect. The difference in root development as between the two lots of plants is
typical also of other pairs of lots from other species similarly treated, and is
construcd as being related to the active production of auxin by thc foliage of
plants in the check (untopped) lots, and the cesSation of auxin production in
the defoliated lots. Both lots of plants shown here were exposed to darkness for
12 days just prior·to the taking of these pictures. This treatment stimulated (or
at least was accompanied by) fresh root 'development in both lots-Icss in the
defoliated lot (5). It will be notcd that thc superior root development in thc
check lot (6) is associated with thc recent awakening of basal buds in plants
1 and 4. This is construed as possibly related to the active removal of au.xin in
the vicinity of basal buds during the recent active production of new roots.
 220 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 221
rhizomes of Bambusa longispiculata (P.I. 93573) that had been embracing more than Olle axis would be cumbersome to handle
discarded when the clump divisions of this and. other large and uneconomical of space in the propagating bed.
species of clump bamboos were prepared by the wholesale method In the present study the age of the material was automatically
formerly employed by the Federal Experiment Station in Puerto fixed at less than 1 year, because the peripheral rhizomes must be
Rico. The viable buds of these fragments developed into small processed before their lateral buds have begun to push-and event
plants while the rhizomes were lying in the open or partly covered that takes place about 1 year after their own emergence. The rhi-
by debris. zomes used were lifted in March and April, and were, therefore,
In 1949, I set up an experiment (with F. Montalvo) at the Fed- about 9 or 10 months old.
eral Experiment Station in Puerto Rico to determine how rhizom:} The principal concern in this experiment was the production
less· than 1 year old (with all buds dormant) would respond as of small, rooted plants for shipment by air to experiment stations
propagules for the production of smalI, rooted plants. Data on the on the mainland of tropical America. As it turned out, the process-
array of material tested, dates planted, initial survival rates, and ing of the plants proved to be such a pressing task that the de-
so forth, are given in Table 4. Figures 90 and 91 illustrate the tailed statistical aspects of the study had finaUy to be abandoned
p:eparation and handling of the rhizomes of Bambusa longispiculata. to give way to the seiection, processing, and shipping of the plants
Flgure 89, 3-6 shows some of the responses of two other species that met the specifications "smalI" and "weH rooted," and the
to propagation by rhizomes alone. . recording of notes on a smaU ,;,rray of plants given special treat-
It seemed logical to begin with single rhizomes, taken from the ment.
periphery of the clump. Apart from the matter of convenience The foUowing major problems were encountered in the course
there is. no reason to limit the size cf the propagule; in fact, unit~ of this exploratory study: (1) meager development of roots on
embracmg two or more axes left attached to each other might culms arising from the distal buds of the rhizomes; (2) decay of
weIl give better results in same bamboos. However, anyone wbo the rhizomes; (3) slowness of the more proximal buds of the rhi-
has excavated the rhizome system of a large bamboo will under- zomes to break dormancy.
stand that th~ simplest procedure is to take away single rhizomes Meager development rif roots. The distal buds, located at the point
from the penphery of the clump. Further penetration into the at which the rhizome axis is transformed into a culm, were the
mass of rhizomes greatly complicates the task, and increases the first to develop. The growth of buds at more proximal positions on
labor required to prepare each propagule. Rhizome propagules the rhizome was very much deiayed, and most of the shoots came
up singly, at long intervals. Most surviving rhizomes still had
Table 4. Initial response of five species to propagation by rhizomes alone. dormant buds after having been under observation for 3 years.
One of the first difficulties encountered was the weak rooting
Average of the first shoots to appear from the rhizomes. These first shoots
height of appeared promptly and had made a good showing by June 10,
Number of Date Sur- shoots
rhizomes planted, vival
1949 (see Table 4). However, they were to~ sparsely rooted to be
June 10, 1949
Species P.I. No. planted 1949 (percent) trusted to survive shipment without first having been established
(ft)
Bambusa longispi- 93573 150 in a nursery bed and allowed to produce additional, more ade-
Mar. 18,31 98.10 6.5
culata quateiy rooted culms.
B. textilis 80872 150 Apr. 1-7 85.33 As a device to get new, more fuUy rooted culms, a new treat-
B. tulda 21002 2.5
156 Apr. 1-7 94.23 4.5 ment was introduced into the experiment June 10-18. A few plants
B. tuldoides 21349 102 Apr. 1-7 99.00
Gigantochloa apus 99573 4.0 in each species were topped and defoliated by cutting back each
100 Apr. 1-7 100.00 3.5
culm (while it was still attached to its rhizome) just above the
222 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 223

Fig. 90. Preparation of rhizome cuttings for the propagation of bamboos of Fig.91. Preparation ofrhizome cuttings for propagation ofbamboos ofGroup Ii
Group .~; Bambusa lonp'spiculata, cultivated at the Fecleral Experiment Station, Bambusa longispiculata cultivated at the Federal Experiment Station, Mayagüez,
J

Mayaguez, Puerto RIeo, under P.l. 93573. (Above) First step; severing the rhi- Puerte Rieo, under P.L 93573. (Above) Third step; trimming the raots on the
zome at the slender neck region. (Below) Second step; separating the rhizome lower side of the rhizome so that it will lie: flat and stahle in the propagating
from the culm. bed. (Below) Fourth step; arranging the rhizomes on a layer of the propagating
medium. The covering layer is still to be added.
224 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 225

second abO'Ve-ground node and by trimming the remaining branches drainage in the propagation beds, this susceptibility to decay fungi
to a length of about 2 in. This left the plants entirely devoid of is attributed primarily to the relatively immature condition of the
foliage. This was done on the theory that it would reduce the whole array of rhizomes. However, there are indications cf inter-
amount of auxin reaching the base of the plant, and thus release specific differences in susceptibility.
the buds of the rhizomes of these axes from the inhibition that Gigantachlo a apus~ whose rhizome tissues are relatively soft, proved
auxin is known to induce in vegetative buds-in the dicots, at to be the most highly susceptible to invasion by rot fungi, and
least. The remaining plants were left intact in a11 respects, as a many of its rhizomes were also invaded by some unidentified
check. The material was then rearranged in the beds so as to in- microorganisms that may have been responsible for the death of
troduce another variable into the environment of the defoliated many of the young rooted plants that occurred while they were
material: partial shade vs. fuIl sun. Partial shade was provided by still attached to the mother rhizomes. Bambusa longzspzculata was
alternating the rows of defoliated material with rows of leafy more susceptible in these respects than the other sp~ci~s of B~m­
material that overtopped and partiaIly shaded it. busa, but somewhat less so than Gigantochloa apus. It 1S mterestmg
The plants were examined again 14 weeks Iater (mid-October that these were the two species that, by their superior vigor and.
1949) and the foIlowing observations were made: the relative promptness with which their buds pushed, seemed most
(1) No appreciable number ofnew shoots had emerged from the suited for propagation by rhizomes alone.
rhizome of either the intact or the defoliated culms; In later trials we found that sterilizing the freshly cut surface
(2) The branch buds on the defoliated culms in partial shade at each end of the rhizome wi th a 10-percent aqueous solution of
had all remained dormant, while those exposed to full sun had Chlorox for 5 minutes and then, after they were dry, sealing the
produced a dense canopy of leafy branches, in aIl of the species; ends with melted paraffin reduced the losses from rotting and from
(3) There had been no appreciable· change in the number of disease to a negligible percentage, and also increased the period
roots on either the defoliated or the intact culms. during which the rhizomes continued to produce new plants.
Fourteen defoliated single-culm plants of Gigantochloa apus (P.I. Slowness if rhizome buds to break dormancy. With the exception of
99573), with branch buds still dormant, and 17 intact single-culm occasional rhizomes, as exemplified in Fig. 89, 4, the outgrowth of
plants of the same species, aIl with dormant branch buds, were the buds in all of the species studied was very slow (Fig. 89, 3),
removed from the rhizomes and stored in complete darkness, dragging out to as much as 3 years. This resulted in the expendi-
wrapped in burlap and plunged in moist cocopeat. After 1 week ture of much time and labor in lifting the rhizomes repeatedly to
the plants were removed and examined. It was observed that the remOve single young rooted plants before they became too larg~.
intact plants had dropped their leaf blades, and a considerable The problem of breaking the dormancy of the buds of the rhI-
number of branch buds of both groups had begun to push. The zome is one that remains to be solved. A treatment that would
culms of the two lots of plants were then partiaIly uncovered and induce all the buds to grow out promptly after the rhizomes are
given subdued light and aeration. At this time there was only a put into the propagating medium probablywould solve the ~roblem
slight intimation of the differences that showed up dramatically of decay as weil. Moreover, causing several plants to anse from
I week later, by which time the defoliated plants had made almost each rhizome at one time should decrease the average size of the
no additional root growth, while the intact controls had made a r~oted plants produced, and reduce the labor involved in the re-
conspicuous development of new root branches (Fig. 89, 5, 6). peated lifting of the rhizomes to remove rooted plants. These are
Decay of the rhizomes. The decay that graduaIly developed in both desirable objectives where the shipment of large numbers of
rhizomes of each group necessitated the discarding of some prop- smalI, rooted plants by air, or the establishment of a very large
agules each year. Since adequate provision had been made for commercial planting, is contemplated.
226 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 227

PRACTICAL E?<PERIENCE-GROUP 11. The Rivieres (1879:505) were Accordina to Tsuboi, gravelly sand is preferred to loamy or clay
the first to describe-in a Western language, at least~the propa- soil for the propagating bed, as the heavier soils are likely to in-
gation of bamboos of Group II by means of rhizomes. In their duce decay. The moisture-holding capacity of the sandy soil is
very brief dissertation, they stress the following general conditions said to be improved by the use of liquid manure, whlCh should
for success: (1) the use of only young material; (2) the use of pieces be applied as the young shoots develop. In view of damage to the
at least 15 to 20 cm long, with three or four nodes, each with an young roots that may result from the movement of the culms by
intact bud; (3) irrigation and cultivation as needed. Planting the wind, the author recommends topping the culms; leaving only
times suggested are April for cold cIimates; ('winter" for warm 5 or 6 branches. Removal of any plant from the nursery must be
climates. A planting depth of 10-15 cm and a spacing of 25 cm deferred until it has developed its new rhizome. ..
are suggested. No data relating to performance of different species Tsuboi's directions are probably the most complete In pnnt
are given. Figure 87, A shows a rhizome section of Phyllostachys and appear sound in all respects. They need to b~ qualified-o r
viridi-glaucescens illustrated by the Rivieres as ready to plant. Figure rather supplemented-in only one important detaIl. !hey apply
87, B shows an allegedly rooted plant ("the result") produced by only to those bamboos of Group II that res pond readüy to prop-
this method. However, careful scrutiny reveals that the propagule agation by rhizomes alone (for example, ~pecie~ of Phyllostachys).
pictured here is far from having produceda truly rooted plant. On the other hand for A rundinarza amabzlzs (Flg. 92) and some
The roots below the leafy shoot have emerged from the rhizome species of Sasa and' Indocalamus, rhizomes alone, without rooted
only. Not until a new rhizome axis has been developed and is culms attached, give very indifferent results at ??st, and often
giving rise to other rooted leafy shoots (culms) will a new plant fail entirely to respond to the conventional condltlOns of propa-
have been established. gation described by Tsuboi. Bamboos of Group II that do . not
Tsuboi (1913:trans. 2-4) describes the ]apanese method of propagate readily from rhizomes alone are generally charact~nzed
processing rhizomes for the propagation of bamboos that, by by (1) the frequent occurrence on the rhizome of nodes ~'thout
inference, can belong only to our Group II, although he mentions buds and (2) a generally marked sparsity of r?o:s on the rhIzome.
neither genus nor species. Two Sources or rhizomes are given: (1). The underlying physiology of these charactenstlcs. may be related
those that are found in their normal position under the ground, to the poor reproductive performance of the rhIzomes of these
and (2) those that emerge from the ground either within the bamboos.
grove or at the edge of a grove where the land falls away sharply
at the brink of a ditch or a river bank. Those portions of an . Whole Culms. It is not now known when the first trial or demon-
intact rhizome that have been exposed to light and air for some stration of this method was made, but Kurz (1876:268) says that
time are described as the easiest to get and safest to use. The use the propagation cf bamboo from a practical point of view can. be
of healthy material 2-3 years old with intact buds is recommended. effected ... by taking whole halms [culms] with their roots a~d burymg
The rhizomes are cut into pieces 45-60 cm long. Tsuboi here them length-wise in the ground. By this pr?cess the altemanng branch
stresses an important point, namely, keeping the material maist clusters send [orth yeung branch shoots WhlCh gradualI>: become trans-
formed into stronger and stronge,::f halms in the proportlOn as roots are
from the time it is dug up until it is put into the propagating bed.
formed. Thus large areas can be planted with Httle trouble.
When the propagating bed is ready, the cuttings are planted in a
horizontal position at intervals of 17.5 cm and covered to a depth Kurz adds that) the propagation must be carried ?ut a~ the onset
of 7.5 cm with fine soi! which is mixed with straw ash as a fertil- of, or during, the rainy season. He glves no detatls. Hls observa-
izer, then firmly packed, and watered. Straw mulch is added to tions were confined to bamboos of Group I.
conserve soil moisture, and a shade is constructed over the bed rt is not clear, from Kurz's ward "with their roots," whether
to protect the young shoots from the full force of the sun's rays. he meant that the root-bearing rhizome was left attached to the
 PROPAGATION 229

culm, or simply that the roots normally produced at the lower

nodes of the culms of many tropical bamboos would be allowed to
remain on. Presumably he meant culms without rhizomes, since
culms with rhizomes attached are mentioned elsewhere by him as
~ another category of propagating material. Since very !ittle exact
information has been published on the propagation of particular
bamboos by means of whole culms without rhizomes, this method
was explored for practical reasons.

GROUP 1. Exploratory trials of three lots of whole-culm cuttings of

Dendrocalamus strictus (P.l. 254923) and one lot of whole-culm
cuttings of Bambusa tulda (P.l. 21002)1 gave the following results:
Fig. 92. Propagation of bamboos of Group TI by means of rhizomes alone' Arun-
di~aria amabilis, eultivated at the Federa1 Experiment Station, Mayagüez, Puerto Of three age groups of Bambusa tulda put in in August and lifted
Rleo, ~nder P.L 110509. (Above) Untreated rhizome -euttings being distributed 260 days later, the 2-year-old culms gave the best over-all score,
on a 2-m. layer of eoeopeat. They were later eovered with a similar layer of the with an average of7.16 rooted plants, with a total of 10 culms, per
sa~e material: The 95 pieces of rhizome used in this particular trial were from
18 m. to 5 ft m len~h; the shorter ones were unbranched and the longer ones
whole-culm cutting. It is of interest that, in the later experiment,
mos:ly bran0ed. ThiS and numerous other trials with this and other propagating 2-year-old culms of this species again made the best over-all score,
~edla ~nd wlth other, more elaborate, treatments of the materialIeft the strong which comes out at an average of 12 rooted plants, with a total of
Impr~slOn that, under uneontrolled eonditions, the propagation of Arund~r/ana
77 culms, per whole-culm cutting (Fig. 93, 4).
~abzlts by means of rhizomes alone is very waSteful of material, since very low
Ylelds wer.e the r~le. The mother plant from which these rhizome cuttings were Whole-culm cuttings of Dendrocalamus strictus (Fig. 93, 1-3)
re:noved IS seen In the foreground. (Btlow) Owing to the crowded state of the processed in March and lifted in less than 400 days later made the
rhIzomes, and the h~avy, compact nature of the "lateritie" day soil in which best over-all score with an average of 6.3 rooted plants, with a
they had developed, It was necessary to e.'xercise great care in order to extricate
them in an undamaged eondition. See p. 156. total of 15.9 culms, per whole-culm cutting; those put in in July
and lifted 290 days later came second, with an average of 4.66
rooted pl,mts, with a total of 12.8 culms, per whole-culm cutting;
those planted in August and lifted 267 days later made the poorest
score
Three-year-old culms made the best over-all score, their average
for the three lots being 5.83 rooted plants, with a total of 13.16
culms, per whole-culm cutting. Four-year-old culms made the
second best over-.all score, their average for the three lots being
4.48 rooted plants, with a total of 11.88 culms, per whole culm
cutting. Two-year-old culms were a elose third, with 4.46 rooted
plants, with a total of 10.16 culms, per whole culm cutting.
The results of a later experiment (McClure and Kennard 1955)
give 2-year-old culms of this species the top score, with l-year-old
culms in second place (Table 5).
lUnpublished experiments carried out at the Federal Experiment Station, Pucrto Rico,
1949-50, in collaboration with F. Montalvo.
 230 ELITE BAMBOOS AND PROPAGATION METHDDS

,Fig. 93, Propagation of bamboos of Group I by means of whole-culm cuttings;

pr~cedure developed, and results obtained, in trials of P.L 76641, Dcndrocalamus 3. Base of same plant. 4. Base and root system of a propagule produced from
stnctus ~1-3) and P.L ~1002, Bambusa tulda (4) at the Federal Experiment Station, a bud on a branch in thc lowcr third of a whole-culm cutting. After it was
~ayaguez, Puerto RIeo. 1. Covering culms with earth after they have been removed from the propagating bcd, the plant was plunged for 1 week in coco-
tnmmed aIfd staked down. 2. Plant developed from one bud in 384 days. peat to promote the restoration of root growth. Each propagule bears a piece of
the mother culm, .sawed off to release the plant in preference to disturbing the
root system to make the cut at the point of its origin.
 PROPAGATION 233

The diverse performance observed, as between species and as

Table 5. Plant production by eleven bamboos of Group I, propagated between individual culms, in the several lots emphasizes the
by whole-culm cuttings. (Adapted from McClure and Kennard 1955 importance of using ample arrays of material in such studies.
Table 1.) . ' Attention is directed particularly to (1) the last column ofTable
5, which gives the score of each age group für each species in terms
Culm Culms Total Plants produced at Plams
age put yield per 10 ft·
of tbe average number of plants produced per 10 ft of culm, and
Species P. 1. No. (yr) ;n (plants) base middle tip of culm (2) columns 6, 7, and 8, which show the actual numbers of plants
Bambusa produced in tbe lower, middle, and upper parts, respectively, of
polymorpha 61373 8 101 2 26 73 U the culms of each age group.
3.4
2 9 104 5 56 43 3.5
3< 9 36
The number of culms used is, in most cases, too small to give
9 23 4 1.2
lextilis 80872 1 14 8 o o 8 0.3 statistically significaut results. Moreover, arepetition of the ex-
2 16 o o o 0 0.0 periment, under different ecological conditions-planting the
3< 274 109 o 8 101 0.2
tulda 21002 1 66 2 o 2 0 0.0 cuttings after the initiation of tbe rainy season, for example-with
2 73 1229 282 366 581 3.9 more culms in each age group, probably would bring out a dif-
3< 34 767 154 310 303 6.8
tuLdoides 21349 1 36 72 o 10 62 0.6
ferent pattern of behavior. However, tbe results obtained strongly
2 40 261 9 15 237 2.4 suggest (1) that each species shows a distinct pattern of response
3< 47 190 17 16 157 1.6
vtntdcosa 77013 1 7 75 14 45 16 2.5
to propagation by means of whole culm cuttings, and (2) that age
2 8 223 15 120 SB 7.4 of culm, and position within the culm, are variables of importauce
3< 11 107 25 18 64 3.5
Cephalostachyum in relation to yield.
pcrgracile 64808 1 7 6 3 o 3 0.2 Bambusa textilis gave very poor results in this trial. Out of 304
2 9 o o o 0 0.0 culms put in, only 25 produced rooted plants. In an earlier trial
3< 13 2 2 o 0 0.1
under more favorable weather conditions, with adequate soil
126493 1 8 109 20 44 45 3.6
2 10 128 89 16 23 4.1 moisture, this species gave an average for all ages of 7 rooted plants
3< 13 15 8 6 1 0.5
Dcndrocalamus per culm, or a score of about 5 rooted plants per 10ft of culm.
strictus 77061 1 12 49 6 24 19 1.0 The contrast between· this score and the very low score made in
2 12 77 o 33 44 1.5 the 1952-1953 experiment is attributed to the extremely dry, hot
Gigantochloa
3< 12 14 11 o 3 0.3
conditions that prevailed in soil and air during the first week after
apus 99573 1 6 36 36 o 0 1.8
2 7 108 44
the culms were put in for the second trial of tbis species, when
24 40 5.5
3< 8 187 74 43 70 9.5 irrigation facilities had not yet been instalIed. All of the lots of
Guadua
angustifolia 132895 1 7
other species in the 1952-1953 experiment (Table 5) were protected
99 19 19 61 3.9
2 6 222 30 99 93 9.1 by irrigation from the time they wer;' put in the ground (1 week
3< 7 166 65 45 56 7.4 after B. textilis) until the rains started.
Sinocalamus
oldhami 76496 1 9 31 3 o 28 1.1
Bambusa ventricosa responded weil to propagation by whole-culm
2 9 128 2 24 102 4.9 cuttings, with an average score for all three age groups of 4.46
3< 8 206 31 68 107 8.2 rooted plants per 10ft of culm, aud a high of 7.4 for 2-year-old
nproduction of 2.5 plants or m:ore per 10 ft of culm was rated as excellent·, .
15-24 0_
.,....., culms. This is in sharp contrast with the almost complete failure
goo cl.
of branch cuttings in three age groups which, in .another experi~
ment, lay in the propagating bed for 2 years, with a bud dormancy
persisting at more thau 95 percent.
 234 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 235
Cephalostaehyum pergraeile manifested a marked difference in re- The Rivieres (1879:473) describe culm "cuttings" as pieces of
sponse as between the respective lots of material from the two culm, bearing fascicles of branches (Fig. 88): They call this the
clones compared (P.l. 64808 and 126493). most rapid method of propagation. They deal with only a few
Gigantoehloa apus (Table 5) made the highest single age-group large bamboos of Group I: Bambusa vulgaris, and unidentified
score-an average of9.5 rooted plants per 10 ft of3-year-old culms. species of Gigantochloa (as Bambusa macroculmzs), and Dendrocalamus
Guadua angustifolia made the second highest single age-group hookeri (as Bambusa haokerz), aIl species in which root primordia
score-an average of9.1 rooted plants per 10 ft of2-year-old culms. appear readily on the base of the branches. As obse'rved by the
This species also made the highest over-all score-an average of Rivieres, these "need only make contact with the soil in order to
6.6 rooted plants per 10 ft of culm for all three age groups. deveIop." The authors direct attention to the importance of leav-
Since the outgrowth of buds on the cuttings continued serially ing a part of the culm intemode attached to the branch comple-
aver a lang period, there was a wide divergence in the size of the ment, the object of which they describe as "to support life i~ t.he
rooted plants recovered. No comprehensive record was made, branch cluster while waiting for the roots to deveIop." It lS m-
either of the number of culms in each plant harvested or of the dicated that, once the branches of the cluster are rooted, "they
range of maximum heights achieved. Many of the plants were may be separated into several individuals."
still very small when the material was lifted, but others had reached Dendroealamus strietus. Some bamboos of Group I apparently do
impressive dimensions. Rooted plants from Bambusa ventricosa, for not respond favorably to this method of propagation in all circ.um-
example, had produced culms 18 ft tall in about 400 days. It stances, however. Deogun (1937:121) says of Dendroealamus strzctus:
would be worth while to ascertain whether it is feasible to remove
Culm cuttings have never struck at Dehra Dun in spite of the fact
the rooted propagules individually as they reached the required that cuttings from eulms of all ages, planted horizontally, v~rtically,
height.
notched, ete., were tried. Orissa reports 20-30 percent suecess Wlth such
euttings and Sunder Lai Pathak [reports] from Pinijaur, Patiala State
GROUP 11. The Rivit,res (1879:461) indicate that their trials of (Ind. Far. 1899, p. 307) a suecess of about 95% after one year and ~O%
whole-culm cuttings of bamboos of this group gave negative after 2 years, the rest having been killed by drought. In thlS case cuttmgs
results. They do not list the species actually tried. I have not with 2 nodes were made from culms 3-5 years old and [these] were
attempted to propagate any bamboos of this group by means of planted Out between 15th January and 15th February in nursery beds,
whole-culm cuttings. weIl manured with leaf mould and stahle manure and sunk below the
ground level ... The beds were flooded twice a week ar:d the cuttings
Culm Segments. GROUP I. Culm segments of bamboos of Group I sprouted by the middle of March. Ninety-five percent glve 1-3 shoots
embracing one or, usually, two or mOre nodes bearing buds or which grew 3' during the first rains.
branches, constitule a form of propagule in casual use in both Dabral (1950) reports confiicting conclusions from his experience
hemispheres. The branches are usually pruned to a length of a in the propagation of Dendroealamus strietus by culm segments. It is
few inches to a foot; no foliage is retained. Such cuttings are stated (p. 313) that this species gave a poor response, and on the
usually set upright or at an angle, with at least one node weIl cov- following page the author says that
ered. AIthough there is very littIe in print relating to this method1 the indications are that für Dendrocalamus stn'ctus~ three-foot long cuttings
of propagating bamboos, personal observation indicates that it taken from two-year old culms plan ted horizontally, one inch below the
is commonly used for certain species such as Bambusa vulgaris soil surface, offer prospects of successful regeneration of the species, under
which propagate readily by almost any of the conventional means. the soil and climatic conditions of -Dehra Dun.

lWhile this book was in galley proof, I camc upan thc experimental results of two Similar cuttings of Bambusa arundinaeea gave no response.
fruitful studies of thi$ method published by W. C. Lin (1962, 1964), Specialist and Chief, Dabral also reports general indications of ·success, without de-
Lu-kwei Brauch, Taiwan Forestry Research Institute, Lu-kwci, Kaoshiung, Taiwan.
tails of performance, in the propagation of Bambusa polymorpha,
 236 ELITE BAMBOOS AND PROPAGATION METHODS

Bambusa tulda, Dendrocalamus longispathus, and Thyrsostachys oliven, -----,----'1

from segments of 2-year-old culms, planted at Dehra Dun during f------ ABOVT ONE METER
(ABOUT 40 INCHES)

the third week in June 1949. The slender tips of the culms were
,discarded and the remaining portion divided into 3-ft segments.
The segments contained, in most cases, abaut three nodes each.
The branches were trimmed to a length of 3 or 4 in., with care
not to injure the buds. The segments were plan ted horizontally in
trenches and covered with 1 in. of soil, weil tamped.
Troup (1921:993) used culm segments of undesignated species
of bamboos (all probably of Group I); he says:
Stern cuttings without rhizomes attached are very uncertain. Vertical
BAMBOO
stern cuttings are usually taken by cutting the culm down as low as pos-
sible) where it joins the rhizome, so as to include the lowest nodes, which
tend to produce rootlets. Culms one year old should be employed.
The superiority of l-year-old material was not borne out by 'my
Qwn experience with culm segments of Bambusa vulgaris var. vittata)
an account of which folIows.
Bambusa vulgans var. vittata. In connection with the establish-
ment, in 1948, of a 20-acre experimental field planting of Bambusa
VJEW FROM SIDE
vulgans var. vittata at Teleman, in the valley of the Polochic River,
Guatemala, I incorporated in the routine an experiment to test Fig. 94. Propagation of bamboos of Group I by means of culm segments with
buds but no branches; Bambusa vulgaris var. vittata. Design developed. for an
the infiuence of age of material on the performance of culm seg- experimental planting at Teleman, Alta Verapaz, Guatemala, showmg the
ments as cuttings. orientation given the cutting as it is planted. Meter-Iong basal cuts of cul~s,
One thousand nineteen of the 1696 culm segments planted were in six age groups, were used for determining. the effect of age of culm material
on yield of rooted plants. See Table 6 and Flg. 95.
basal cuts, and the performance record presented here is confined
to this array. The material was divided into the following six age
Table 6. Performance of I-rn basal segments of culms of Bambusa vulgaris
groups: (1) less than 2 months; (2) about 6 months; (3) 12-18
var. vittata put in as cuttings November-December 1948; data from field
months; (4) 24-30 months; (5) 36-40 months; (6) 48-60 months.
notes completed in July 1949; the calculated percentages were rounded
It is impossible to determine the age of bamboo culms accu-
to nearest whole number.
rately, unless they have been marked in the year of their emer-
gence. However, familiarity with certain visible characteristics Produced Produced
associated with aging makes it possible to arrange them with re- Age rooted unrooted
group NO.of shoots shoots Died
spect to age in aseries that gives meaning to the performance (mo) cuttings (percent) (percent) (percent)
records of the several groups.
The method used in planting the cuttings is shown in Fig. 94. <2 110 20 22 58
ca. 6 256 19 18 63
A severe drought that followed planting resulted in a high per- 12-18 283 26 19 55
centage of fatalities. The infiuence of age on survival, however, 24-30 228 30 19 51
and on the performance of the material is clearly shown in Table 36-40 98 51 29 20
48-60 44 50 34 16
6, and in Fig. 95.
 PROPAGATION 239
70r-__ ~ ____ ~ __ ~ ____ ~ __- ,
this apparently is the first record of tbe successful propagation of
a bamboo by means of the slender distal branches of tbe culm.
It is also the first record of the artificial stimulation of bamboo
root growtb with moist air as the propagating medium. The high
Produced humidity prevailing in tbe propagating frame is presumed to
rooted
50 hoots have been responsible for activating the growth of the existing
w root primordia. This demonstration suggests further experimen-
Cf)
z tation. Propagules rooted out of contact with asolid rooting
0
CL
Cf) 40 medium would be especially desirable for certain experimental
W
0: studies. And such a convenient source of clean root tips would
i-
Z
have important .advantages for chromosome studies.
w 30
"
0:
W
CL
GROUP II. The Rivii:res intimate (1879:504) that bamboos ofGroup
H are capable of giving rise to new plants from the basal segment·
of a culm, planted as a propagule. Their claim is documented by
an illustration (p. 503, Fig. 11) here reproduced as Fig. 96, lift.
Although their claim is true for some bamboos of Group H, it is
10
obvious that the conditions of success in this type of cutting are
not demonstrated by the figure. The branches tbat have arisen from
a bud just at the surface of the ground have no roots of their own.
~~2-----ca~6~--~12~-~18~~24~-~3~0--~3~6·-4~0~-4~8-60 Not having produced roots while the basal nodes were still in a
AGE (MO) meristematic condition, such axes never could have racts. Neither
Fig. 95. Performance cf 1019 culm sections (basal cuts only) by age groups; could they in the given situation, ever produce other culms that
Bambusa vulgaris var. vittata; Teleman, Polochic Valley. Guatemala, December will root themselves, since they have no buds below the surface of
1948-June 1949. the ground. And, most important of all, there is no bud on any
part of the original propagule itself that is covered with earth.
With the permission of Dr. J. Van Overbeek, reference is made There must be a bud in this position, and it must give rise to a
heFe to unpublished results of an exploratory study he carried out rooted culm; and from a rooted culm there must emerge a rhi-
at Mayagüez, Puerto Rico, in the spring of 1945, and reported to zome-the only permanent source of new culms. Until this con-
me by letter, wi th photographs, at that time. dition has been fulfilled, it cannot be said that a complete plant
Representative cuttings, including branch-bearing segments of has been produced. I observed such a defective propagule of
culms, of Bambusa vulgaris were placed in sand in a humid propa- Phyllostaehys pubeseens in my garden at Lingnan University for
gating frame, in late April or early May, without treatment with several years. It remained throughout that period just such a
auxin. Thirty-six days later, rooting had taken place in numerous "plant" as the Rivieres illustrate in their Fig. 11, never producing a
branches, principally those above the level of the sand (Fig. 99). rhizome or a new culm.
In exceptional cases, a few slender branches that were covered A propagule fulfilling the conditions for success can be secured
with sand produced roots_ from bamboos of Group H, only (1) as the basal cut of a culm of
In nature, of course, the principal mid-culm branches of Bam- small stature, where buds are produced at even the basal nodes,
busa vulgarz"s form root primordia spontaneously, in situ. However, or, better, (2) by using the underground part of tbe culm, where
 PROPAGATION 241

the nodes often have dormant buds (for example, some species Of
Phyllostachys). In some bamboos of Group II, however, even this
underground part of the culm is sometimes entirely devoid of buds
(for example, Arundinaria amabilis). In view of the uncertainties
that plague the search for suitable propagules of this type in bam-
boos of Group II, it is advisable to depend upon other methods of
propagation.
The Rivieres (1879:461) signalize their failure to propagate
bamboos of Group II by means of culm segments bearing branch
complements by the statement that "bamboos with summer growth
bear only nonrhizomatous branches which cannot be used for
propagation. "
Since no other reference to the use of culm segments or branches
of bamboos of this group for vegetative propagation could be
found, and since recorded experience in bamboo propagation is
generally based on procedures of a relatively primitive nature, the
limitations ascribed to propagating material from certain sources
is probably a function rather of the crudeness of the methods used
than of the source of the material. This view is supported by the
fact that Arundinaria amabilis, a typical member of Group II, has
been piopagated successfully by means of culm segments bearing
full complements of branches when these are exposed to a con-
trolled range of temperature and humidity. The conditions and
results of this exploratory study by Dr. John Creech have not been
published.
Fig. 96. Lift· ~~~pagation by means of a culm segment with branches. Group II:
Phyllostachys vmdz-glaucescens. A rooted propagule, consisting of the basal segments Single Branches. GROUP I vs. GROUP II. No record of the successful
of a cul~ cut off at ground level, as illustrated by A. & C. Riviere 1879:Fig. 11. propagation of any bainboo of Group II by means of single
As e.xplamed elsewhere, this propagule has not given rise to a complete plant branches as cuttings has come to my attention. The discussion
(See p. 239).
that follows concerns only bamboos of Group 1. These are charac-
Center ~~? right. Propagation by means of single branches. Group I. ('Bambusa
macroculmzs. (Center) Youn~ singl~ branch cutting prepared for propagation. The terized (p. 209) by the resemblance of the mid-culm branches to
~,heat~s co~ceal the root pnmordla on the swollen "rhizomatous" base. Note the the mother culm in having a swollen basal portion that "recapit-
heel of t.ls.sue from the mother culm, whose retention is stressed by the Rivieres ulates" the rhizome. In certain bamboos of this group (species of
as a condltlOn ?f s,:ccessful propagatiof;l by single branch cuttings. (Right) A
young shoot (stIll WlthOut roots) has arisen from one of the basal buds. When authentic Cephalostachyum and Schizostachyum) the branch comple-
s~ch a shoot becomes .a rooted culm, a new plant will have been established. ments in the mid-culm range are composed of numerous smalI,
Flgures at center and nght from A. & C. Riviere 1879:Figs. 8 and 9. See p. 226. subequal branches, in which thecresemblance to the mother culm
and rhizome is not strikingly manifest. In other genera, however
(Bambusa, Dendrocalamus, Gigantochloa, Guadua,' Sinocalamus), the
primary branch generally greatly exceeds the other members of
242 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 243

the branch complement in size and in resemblance to the mother a_naphthylacetamide, (4) 0.1 mg/mI 2,4-dichlorophenoxy acetic
culm and its rhizome. In same species this resemblance includes acid. The lifth sublot was used as a contro!. The March, J une, and
the spontaneous generation of root primordia on the rhizomelike September lots, each totaling 50 processed ~uttings, were planted
basal part of the primary branch, in situ. The Rivieres (1879:476) in moist sand in a greenhouse; the lots slmüarly processed In
refer to such branches as "rhizomatous." As mentioned repeatedly December were planted in sand outside under partial shade.
elsewhere, the formation of root primordia can take place only White says (p. 393):
while the tissues of the branch base are still in a meristematic No differences in rooting were faund which could be associate? with
condition, and not after they have matured. root-promoting treatments. Therefore, the data fr~m [all] 50. cuttlngs of
For propagation by conventional methods, single-branch cutting~ each species obtained at each date. were summ~zed as umts for COID-
should be made, as a rule, onIy from such rootwbearing dominant parison of differences amang specles and the tIme of year they were
central axes of branch complements in the mid-culm range. In obtained.
the experiments reported below, branches that were not already The results, by species and by months, are shown in Table 7.
rooted did not produce roots. The Rivieres state that in their ex- The author's summary folIows:
perience failure always resulted from the use of the smaller branches
(a) Treatment with root-promoting subst.an~s at .the ~sual con-
of the complement. .
centrations has no effect; (b) considerable variatl?n In ro~tlng e.XlSts a~ong
Single-branch cuttings were lirstillustrated by the Rivieres species; (e) rooting varies with the month [in WhlC~] cuttlngs ar.e obtalned;
(see Fig. 96, center and right). However, they conline their discussion Cd) the best month for rooting cuttings vari~ wüh the specles; ~nd Ce)
(1879:469-476) chiefly to culm segments bearing whole branch rooting may be associated with rainfall dunng the month prevlOus to
complements, and give scant attention to single-branch cuttings. obtaining the cuttings:
It is probable that the Chinese farmers were the pioneers in this White could have added a further observation drawn from his
lield. Certain bamboos of Group I are still regularly propagated results, namely, that in his experiment the best score was. made
in China by means ofsingle-branch cuttings (McClure 1938a:PI. 6). by those bamboos in which the most abundant productlOn of
Taking a cue from the Chinese farmers, I succeeded in 1925 in
establishing single rooted branches of Sinocalamus beecheyanus, in
sphagnum, for shipment to the United States. Table 7. Performance (percentage rooted) of single branch cuttings of
Cobin (1947 reported the successful use of branch cuttings to nine species of bamboo put in at four different seasons of the year.
propagate Sinocalamus oldhamiz~ Bambusa vulgaris) B. vulgaris var. (Adapted from White 1947b:393.)
vittata, and Gigantochloa verticillata, all species in which root pri- Season
mordia appear spontaneously in abundance on the swollen part of
Mar. June Sept. Dec. Ave.
the principal branch, in situ, at mid-culm nodes. White (1947b) Species
studied the effect of season of the year and of root-promoting sub- 2 0 0 0 0.5
Bambusa longispiculata
0 0 0 12 3.0
stances on the rooting ofbranch cuttings ofnine species ofbamboos. polymorpha
10.5
textilis 16 25 2 0
Branch cuttings 12-18 in. long from 2-year-old culms were 2 0 0 12 3.5
tulda
processed in lots of 50 each at four seasons: March, June, Septem- 0 0 0 0 0.0
tuldoides
ber, and December. Just before being planted, each lot of 50 Cephalostachyum pergraeile 0 0 0 0 0.0
20 0 20 52 23.0
cuttings was divided into live sublots of 10 each; the basal portions Dtndrocalamus asper
50 46 0 50 36.5
Gigantochloa apus
only of the cuttings of four of the sublots were dipped for 5 sec- Sinocalamus oldhami 36 24 8 16 21.0
onds into ;,!eoholic solutions (Cooper) of, respectively, (1) 5 mg/mI 14 10.5 3.3 15.8
Averages, by season
indole-3-acetic acid, (2) 2 mg/mI indole-3-butyric acid, (3) 2 mg/mI
 244 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 245
root primordia takes place spontaneously, in situ, before the conditions prevailing in the conventional propagating bed. Dif-
branches are removed from the mother culm. Cuttings that have ferences between species in the natural propensity toward spon-
no root primordia when taken from the mother .plant will not taneous production cf root primordia on branches, in situ) are
themselves produce roots. Rooted plants can be secured from shown in the following personal observations (where available,
branch cuttings only through the sprouting of buds On the under- U.S.D.A. plant introduction numbers are given): Bambusa longispic-
ground portion.
ulata, P.l. 93573, O-few; B. multiplex, O-few; B. multiplex var.
T:vo hund~ed .branch cuttings (consisting of the basal foot-Iong riviereorum, P.l. 77014, O-few; B. polymorpha, P.l. 61373, O-few;
~ornon .of prmclpal branches from mid-culm nodes) of each of B. textilis, P.L 80872, O-few; B. tulda, P.l. 21002, O-few; B. tuldoides,
SIX speCIes of bamboos of Group I Were put in at Mayagüez in P.L 21349, O-?; B. ventricosa, P.l. 77013, o-?; B. vulgaris, numerous;
]uly 1~49 (McClure and Montalvo 1950) with the objective of B. vulgaris val'. vittata) numerous; Cephalostachyum pergracile) P.l.
produc.mg small, rooted plants for shipment by air.. In terms of 64808, O-?, PJ. 126493, O-few; Dendrocalamus asper, P.L 71258,
the umformly small size and adequate rooting of the plants se- numerous; Gigantochloa apus, P.l. 99573, numerous; Gnadua angusti-
cured, branch cuttings were found to produce the best results of folia, P.L 132895, O-?; Guadua angustifolia (caiia mansa), numerous;
any of the several meth~ds of propagation tried (Fig. 97, 1-3). Sinocalamus oldhami, P.l. 76496, numerous.
H?:vever, the slow and Irregular breaking of dormancy in the U nder certain circumstances spontaneous rüoting of branches,
cntIcal buds rendered the yield uneven and in some cases en- in situ, is stimulated extraordinarily. Cobin (1947:183) notes that
tirely unsatisfact~ry. Of the six species, Gigan;ochtoa apus gay; the a plant of Bambusa textilis, whose branches normally root very
b:st }'leld by thls method; Bambusa ventricosa gave the poorest, sparsely, showed an abundance of roots on its. branches~ ~ cir-
wlth less than I-percent response. It is noteworthy that most of the cumstance which he attributes to the poorly dramed condItlOn of
buds o.f the cuttings of B. ventn'cosa remained alive-and presum- the soil in which it was growing at the Fairchild Gardens, Coconu t
ably vlable-after having been in the propagating medium (1:1 Grove, Florida. I have observed the same phenomenon in the
cocopeat and basal for 2 full years. No effective method for break- same species under circumstances that are, perhaps, more suggestive.
ing the dormancy of the branch buds in refractory species was Spontaneous production 01 rooted plants in place 01 branches. A smalI,
found.
single-culm rooted plant of Bambusa textilis set. in the bamboo
Spontaneous rooting, in situ, '!! bamboo branches. It may be of interest nursery of the Instituto Agropecuario Nacional at Chocola, Gua:e-
~o discuss this ch~racteristic briefly from the point of view of phys- mala, failed to produce new culms, but several branches at ItS
1010gy. T?ree pomts may be· noted as foci of attention: (1) as lowermost branched node bore long roots, most of which pene-
obse:ved In ~atur~,_ the "propensity" toward spontaneous rooting, trated the soil (Fig. 97, 5). After the branches had been removed
partlcularly m pnmary branches, is more strongly developed in (Fig. 97, 6) and set in the ground as propagules, the plant itself
some bamboos of Group I than in others; (2) bamboos that nor- was excavated. The rhizome was found to be devoid of viable buds.
mally show no root primordia, or only a few, On the principal This observation supports Cobin's suggestion that it should be
branc~es may, under certain circumstances, be induced to produce possible, through suitable procedures, to induce or increase de-
them m abundance; and (3) under either normal or certain ab- velopment, in situ, of root primordia on the branches of bamboos
normal conditions in nature, complete rooted plants may be of this group, in anticipation of the use of such branches as
produced spontaneously in pI ace of apart or all of a branch cuttings. .
complement in certain bamboos of Group 1. A similar occurrence was observed in a small single-culm prop-
The relative abundance, Or lack, of root primordia in branches . agule of Bambusa tulda, another species whose branches normally
of the ba:nboos teste.d in connection with propagation studies is root very sparingly. However, in this case, a complete rooted plant
reflected m the readmess with which they strike root under the took the place of the branch complement at the lowermost node
 246 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 247

(Fig. 97, 4). This was discovered in a field planting at Mayagüez. accumulation of materials could, conceivably, result in the spon-
Upon the excavation of the mother propagule it was discovered o taneous production of Iittle plants from the adjacent branch buds.
that, again, there were no viable buds on the rhizome. In 1953 I This typical form of Guadua angustifolia flowers with extreme rarity,
observed a similar occurrence in Bambusa arundinacea at New Delhi and is not known to have produced viable seeds. It seems quite
India. ' likely, therefore, that its natural dissemination to new areas is
It appears that the food- and growth-regulating substances accomplished primarily by means of these litde rooted plants
e1aborated in the upper part of the propagule accumulate in the which, being attached by a single very slender and brittle "neck,"
lower part of it, without findirig an oudet in the normal produc- would easily be carried away by a stiff wind. Some of those that
tIOn of new growth. Apparendyas a result of the presence of .this happen to land in a stream could, on occasion, find a Iod ging
accumulated material, the lowest viable branch primordium place in stream-bank sites. .
responds by producing a complete rooted plant in place of a Chaturvedi(1947:543, PI. 33, Fig. 1) reports and figures the
branch complement of the conventional form. occurrence of rooted aerial rhizomes, in situ, at the lower branching
Other examples of the occurrence of complete, rooted plants in nodes of old ("parent") culms in congested plants of Dendrocalamus
the place of part or all of a branch complement have been strictus at Baria, Bhind District, India.
obsen:ed . The first invo~ves a bamboo in which natural rooting of Bambusa multiplex is another bamboo in which the spontaneous
the mld-culm branches IS abundant. In a variant of Guadua anuusti- . production of root primordia on mid-culm branches is generally
folia known locally in Ecuador, as "caiia mansa," whole mid-~ulm sparse or nil. In 1948 I saw, in EI Salvador, an old plant of a
branch complements often take on the form of a small rooted dwarf form of this bamboo in which several little rooted plants
plant. Ir this propensity has a genetic basis, it might weil be had emerged [rom mid-culm branch complements of the oldest
possible, and useful, to undertake to incorporate it in other elite culms. The base of the branches had taken on a wholly rhizome-
bamboos through cross breeding, in order to facilitate veg~tativ~ like form and behavior, even to the positive geotropism of the
propagation. This form is much less thorny than the typical form "neck," and the diageotropism of the rhizome proper. .
of the species. These -observations make it possible to read with comprehensIOn
The second example involves the typical form of the same the following passage in Satow (1899:75) about another small-
sp~cies, ~n which the branches at mid-culm nodes rarely bear root leaved form (J apanese: Ho-o-chiku) of the same species:
pnmordla, and apparently have never been observed producing In neglected hedges, roots [rhizomes] are faund hanging down in the
complete plants in situ. It was in the course of a survey of the fonn of a fish-hook. The upper part lengthens into a culm. From the
local production of culms of this species in the Department of root [rhizome] other roots [rhizomes] branch out, gradually increasing. in
Caldas, Colombia, that I found, in the very top of some of the number so as to forro,a bole [the rhizome "nexus" cf Arber], from WhlCh
oldest culms, perfectly formed Iitde rooted plants, in the midst of fine hair-like roots grow downward, but as they cannot reach the soil,
they stop growi~g after attaining a sun Uapanese inch] in length. From
very much overcrowded and degenerated branch complements. It the bole a number of stems grow closely together, ofwhich the inner ones
seemed that the production of these litde rooted plants might be bear branches. This bole attains the size of a half-bushel measure, and
related to a deterioration (state of senescence?) in the phloem of yet is held on to the parent stern by a single root-fibre [the ((neck" of the
very old culms-more especially, perhaps, in the anastomosing primary branch]. Should it be hit with violence, it comes away suddenly,
bundles at the node at which the old branch complement that and if stuck in the ground will give rise to a dense growth.
produced Iittle plants was attached. Faulty functioning of the In 1955 a plant of the Chinese Goddess 'bamboo (Bambusa
phloem could obstruct the downward movement of e1aborated multiplex var. riviereorum) was grown in nutrient solution in a pebble
food and auxins and cause them to accumulate at the base of the medium, irrigated at 8-hour intervals. After 2 months many of the
branch complement in whose leaves they were e1aborated. Such an branch complements on several of the culms had proliferated to
 248 ELITE BAMBOOS AND PROPAGATION METHODS

5 6

Fig. 97. 1-3. Propagation of bamboos of Group I by means of single branch

cuttings: 1, branch cuttings being plantcd in the propagating bed and pertinent
data being recorded, in connection with experiments carried out at thc Federal
Experiment Station, Mayagüez, Puerto Rieo; 2, Bambusa textilis (P.l. 80872), a
well-rooted plant developed from thc distal bud of thc cuttingj 3, Gigantochloa
apus, P.l. 99573; this array of four cuttings demonstrates a tendency that appears
to be general, namely, that ,buds at thc base of branch cuttings in which thc
swollen, "rhizomatous" natural base of thc branch is retained are generally
slower to germinate, but usually produce a better root system, while thc buds at
more distal nodes push more promptly and the resulting culms generally are
slower to develop a good root system. See p. 245.
4-6. Spontaneous production of rooted plants in place of "normal" branches:
4, Bambusa tulda (P.L 21002) at Federal Experiment Station, Mayagüez, Puerto
Rico, December 1948; 5, Bambusa textilis (P.L 80872) at Chocola, Guatemala,
May 1951; 6, four little rooted plants removed from the mother plant shown
in 5.

3
 250 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 251

produce little rooted plants. This exploratory trial was carried out attached to the mother plant, without notehing, satisfactorily pro-
through the collaboration of tbe late Dr. Robert Withrow, at the duced little rooted plants. However, this method was to~ cumber-
Smithsonian Institution. seme to be considered seriously except in unusua.1 circumstances,
Natural propagation i?Y means qf bulbils. Propagation by "bulbilIes," or for very small bamboos, such as the dwarf cultivars of Bambusa
a French term for bulbils or bulbules, formed in connection with multiplex, and Bambusa multiplex var. riviereorum, which responds
the infiorescence after the manner prevailing in some other grasses favorably but does not produce. abundantly.
(for example, Poa bulbosa) is reported by Dutra (1938:145, 150). Method 2. Stump layers may be prepared by cutting off one or
The author's words, freely translated, follow: more of the culms in a clump, leaving one or two nodes with a
bud or branch complement, and covering the stumps so prepared
In Bambusa riograndensis Dutra [that is, Guadua trinii (Nees) Nees' ex
Rupr.] the plant usually propagates itself by bulbils ... During the with a suitable mulch.
period of flowering, the plant loses all of its leaves and covers itself.with In J anuary 1949, exploratory trials of this method were initiated
a multitude of inflorescences, of which not the thousandth part produces at Mayagüez witb Bambusa longispiculata, B. textilis, B. tulda, B.
fruits, which fact is due, I think, to the faulty disposition of the stamens tuldQides, and Dendrocalamus strictus. Whole clumps were prepared for
in relation to the pistils. The latter, being very small, remain enclosed, treatment by severing all culms at about 2 in. above the second
while the former, each equipped with a long filament, emerge while the
node above ground. A bamboo barrier was constructed around
floret remains almost closed. The plant seeks to make up for this defect
by producing, here and there, bulbils that will insure reproduction. each clump to hold the propagating medium. The number of
stumps in each clump was recorded, and half of the stumps in
Layering. In contrast with propagation by means of vegetative each clump were treated with indolebutyric acid, by placing a
fractions, propagation by layers delays the rem oval of the propa- handful of cocopeat impregnated with an aqueous solution
gule from the motherplant until after it has established roots in (200 ppm) of IBA in the axil of each branch complement. The
a propagating medium. The following riletbods may be employed. branches in the other half of the stumps were left untreated, as a
(1) Either a whole culm or only the branch-bearing part of it is check. Finally, the stumps were covered with a 1: 1 mixture of
bent down to the ground and into a shallow trench, and fastened cocopeat and "basa" (Fig. 98, 1-2).
in place by means of hooked or crossed stakes, with or without One clump of each of the live species was treated in this manner
notehing it below each branch-bearing node, and covering it with in January 1949, and a second clump of each in March 1949.
earth or any other suitable propagating medium; the subsequent Botb clumps of Dendrocalamus strictus died outright, possibly of
routine is similar .to that used for whole culm cuttings. (2) The anoxia, which may have resulted from theexcessive depth of the
stumps of severed culms arecovered with a suitable propagating culture medium. Of tbe remaining species, only Bambusa longispicu-
medium. (3) A culm is kept erect, with or without notehing it lata produced rooted plants (Fig. 98, 3). The groups of rooted plants
below each branch complement, and the base of each branch produced by the two clumps of this species were removed in June
complement in the mid-culm range is surrounded with a suitable 1949, with the results shown in Tables 8 and 9.
propagating medium, held in place by a suitable receptacle. In Table 9, an analysis of the details recorded in Table 8 shows
Method 1. The Rivieres (1879:476) recommend layering of whole that the treated stumps gave the better rooting, but produced
culms only for Group I. They call it the "easiest" method of fewer culms than the untreated ones. Such behavior agrees with
propagation, but they do not give data on the performance of.any that observed elsewhere among the fiowering plants, where the
particular species. development of buds is inhibited, but the initiation of root pri-
In preliminary trials, carried out in Guatemala, I found that mordia is stimulated, by the local application, or presence, of
1-year-old culms of Bambusa textilis and Guadua angustifolia (clone auxin.
Cafia Mansa) when bent down and covered with earth while still This is apparently the first recorded trial showing a difference
252 ELITE BAMBOOS AND PROPAGATION METHODS

Fig. 98. Propagation of bamboos of Group I by means of stump layers. Results

achieved at Federal Experiment Station, Mayagüez, Puerto Rico. 1 and 2. Prep-
aration of a clump of Bambusa tulda, P.L 21002: I, propagating medium (1:1
cocopeat and "basa") being added to the level of the tops of the stumps; 2,
stumps in foreground treated with IBA; background, untreated "check." 3. Bam-
busa longispiculata, P.L 93573, e..xamples of smalI, rooted plants, propagated by
thc method illustrated above, and selected for shipment by air. Average weight
of 56 plants of this lot, 4 oz; average weight of thc 94 cornponent culms, 2.5 oz.

3
 PROPAGATION 255

Table 8. Number .of rooted and unrooted culms from two stump layers branch complements of l-year-old culms of Bambusa tuldoides and
of Bambusa longispiculata treated (as shown in Fig. 99, 1, 2) with indole- current year (developing) branch complements of Semiarundinaria
butyric acid in January and March 1949, and excavated in June 1949. Jastuosa, without achieving any success with either species. In the
(Adapted from McClure and Montalvo 1949, Table 1.) first case, the propagating medium, cocopeat, was held in place
Date treated, 1949 by fumiel-shaped cups made of tarred paper. In the second case,
moist sphagnum was confined in a sheet of pliofilm tied about the
January March Total culm. No root-promoting substance was used in eith"er case, and
. Total stumps in clump the culms were not notched .
Treated (number) 27 25 52
Cheek (number) 26 35 61 Conclusions on past work, and future objectives
Total culms from above-ground nodes 122 126 248
Treated: Rooted (number) 20 17 37 It appears that.most of tbe published observations about the vege-
(percent) 54 27 37 tative propagation of bamboos have grown out of experience of an
Not rooted (number) 17 45 62 empirical or exploratory, rather than a scientific, nature. Even
Check: Rooted (number) 21 11 32
(percent) 25 17 21 where a conscious effort has been made to adopt a systematic
Not rooted (number)_ 64 53 117 approach to the accumulation of knowledge relating to the propa-
Total culms from upper nodes of gation of particular species, and to particular objectives, un-
rhizomes 72 52 124
Treated: Rooted (number) 10 15 25 assessed and uncontrolled variables have marred the quality of the
(pereent) 67 80 74 documentation and prevented the achievement of conclusive re-
Not rooted (number) 5 4 9 sults. Consequently, knowledge and practice are still largely in a
Check: Rooted (number) 29 25 54 retarded stage of development, and satisfactorily documented data
(pereent) 51 76 60
Not rooted (number) 28 8 36 relating to the vegetative propagation of particular bamboos
apparently have not yet been produced.
Published discussions of methods of vegetative propagation of
between untreated propagating material of bamboo and that bamboos are invariably concemed primarily with the particular
treated with a root-promoting substance. fraction of tbe plant used: clump divisions of various dimensions,
Method 3. This method is really a form of air-layering, and is from half the clump down to offsets consisting of one or more
widely used in China in the propagation of woody dicotyledonous culms taken from the periphery of the plant; whole rhizome axes
fruit trees. No reference to its use in the propagation of bamboos or segments thereof; whole culms bearing a bud or a branch com-
has been found in the literature. plement at each node; segments of a culm, each containing one or
In exploratory studies, I have tried air-layering of matured more nodes, and each node bearing either a bud or a whole
branch complement; and, finally, cuttings consisting of individual
Table 9. Performance of treated and untreated stump layers of Bambusa branches.
longispiculata) P.l. 93673-an interpretation of date from Table 8. In these different kinds of propagules the mass of tissue surround-
ing, or communicating with, a given bud or group of buds is suc-
Ratio of Culms cessively reduced. This involves areduction of the stored food,
Stump Number New culms new culms rooted
layer of stumps produced to stumps (percent) hormones, and enzymes available to a bud and to the new shoot
that develops from it. In progressively smaller propagules, the
Treated 52 123· 2.34 50 chemical constituents and physiological gradients of the tissues are
Untreated 61 239 3.91 36
exposed more and more intimately to external influences, which
 256 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 257

may be favorable or unfavorable to success. This makes progres- Wi th a few minor exceptions, every node of every segmented
sively more urgent the necessity of supplying, artificiaIly, basic axis of a bamboo plant bears a bud-or a branch which, in turn,
nutrition and critical physiological stimuli to the individual bud has a bud at every node. TheoreticaIly, at least, each one of these
and to the shoot that emerges from it. buds is a potential plant. Studies in vegetative propagation should
Reduction in the mass of the individual propagule makes for include methods for transforming as many as posslble of these
economy of propagating material, simplifies the labor of preparing innumerable buds into littIe rooted plants.
it, and reduces the requirements of space and other facilities. As It is important to remember that the origin of each rooted
the bud is deprived more and more completely of the maternal plant resulting from vegetative propagation can be traced ~o a
tissue that supports it, the control is perfected, the number of single bud (Figs. 97, 2 and 99, lefl). This fact is n?t at once ObVlOUS
unassessed and uncontrolled variables is reduced, and the pros- when one considers relatively large plants denved from cIump
pects of establishing pertinent basic principles and determining divisions consisting ori«inally
o of several rooted culms. Once .this
the optimal conditions of vegetative progagation for each kind of fact is grasped, however, the nation cf "rooting" bam?oo cuttln.gs
bamboo improve. (often referred to in the literature) is seen in a new hght, and.'ts
limitations are apparent. A branch cutting or a whole-culm cuttmg
often bears root primordia at its lower nodes. Only in such a case
Facts and factors affecting propagation procedures
In the stems of woody dicots and gymnosperms, meristem is
typically distributed in a continuous layer-the cambium. In the
bamboos, on the contrary, ll)eristematic tissue is ~onfined to
certain discontinuous and more or less widely isolated regions. The
principal foci of meristematic tissue in the vegetative body of the
bamboo plant are: (1) the tip of every growing axis; (2) the zone·
of intercalary growth just above each node of a11 actively e1onga-
ting segmented axes; and (3) dormant buds and dormant root
primordia, as long as they remain viable.
Activelygrowing roots are repositories of meristem but, in the
bamboos, it appears that this meristem does not, under natural
conditions, give rise to any kind of organ other than roots. On
the other hand, the meristemof the intercalary zones of actively
growing segmented axes (rhizomes, culms, or branches) may give
rise to root primordia or functional roots in addition to buds (rudi-
mentary leafy axes). There is evidence for the existence of more
or less persistent islands or strands of meristem in other places
within the segmented axes of the bamboo plant after elongation
ceases. As far as we know at present, however, these do not play
any part in the regeneration of whole organs-a process that is Fig. 99. Propagation' of hamboos cf Group I by mearis of culm segments with
branches; Bamhusa Dulgans. Raots developed on brauches cf culm segments after
the principal feature of vegetative propagation. For the purposes 36 days in a humid propagating frame: (left) reets developed under the san?,
of vegetative propagation in bamboo, viable dormant buds should from a yaung brauch on asiender culm sectionj (right) raats that developed In
be made the primary focus of attention. thc humid air above thc sand. Sketches based on photos by Van Overbeek.
 258 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 259

can a bamboo cutting itself be rooted, since root primordia can Experience suggests that, under conventional conditions ofprop-
be produced onIy from meristematic tissue, never from mature agation, processing of propagules should be limited to the period
bamboo wood. However, no new plant is produced by such a when all buds are in a dormant state. Traditional practice favors
rooted cutting unless and until a bud gives rise to a new axis which processing propagules of bamboo just. prior to the initiatio:, of
develops roots of its own. The above fact was demonstrated its annual period of active growth. Th,s penod may be descnbed
dramatically in the large-scale trial of basal cu Im segments of as "late autumn (or early winter) to early spring" for the hardy
Bambusa vulgaris var. vittata referred to earlier (p. 236). bamboos (group II) and "the end of the dry season" for the tender,
Personal observation and published experience suggest that or tropical, bamboos (group I). However, it may prove. :-"orth
there is a definite, optimum age range for propagules of each while linder certain circumstances, to study the feaslblhty of
category (rhizomes, single branches, and so forth) from each proce~sing certain types of propagules during the perio~ .cf active
botanical entity. The best rcsults generally do not came from either growth, by providing controlled environmental condltlOns and
the youngest or the oldest 'material. It is not dear, however, suitable nutrition (cf. p. 210).
whether thc basis for the differential behavior of propagules of Table 7 shows that even dormant buds of propagules represent-
different age resides in thc maternal tissue or in the buds them- ing a given part of the plant may give differen: responses to the
selves, or both. same routine of propagation when processed. at dlfferent seasons of
On the one hand, material bearing very young buds, from the year. The natural assumption is that these responses are due to
which the best results might naturally be expected, often gives seasonal differences prevailing in the physiological state of the
poorer results under conventional treatment than that bearing material-differences presumably related to stored nutrients,
older ones (cf. pp. 236f). This may be because the younger buds hormonal and enzymic activity, and so forth. However, since the
are not receiving optimum levels of nutrients in the communicating control of conditions surrounding the material in the propagatmg
matern al tissues. Or perhaps the hormone levels in the tissues of bed has been in all recorded cases only nominal, the observed
very yaung propagules of certain categories are not always favor- differences in response could be in part a function ~f seasonal
able to the awakening of the buds. On the other hand, the sudden differences·2 or differential daily fluctuations, in envlronmental
falling off, with age, in the yield from some types of propagules- cond~tions' such as light, temperature, and moisture prevailing in
in bamboos ofGroup II at least, as suggested by]apanese authors and above the propagating bed.
(cf. Tsuboi 1913:trans., p. 2)-may be due to a loss of viability in
the buds themselves. A study of this subject would be worth while Dormancy and development of buds
from the point of view of pure science, as weil as thai of practical The natural or characteristic behavior of buds in respect to
application. dormancy and the breaking of dormancy varies w~th their position
in the plant, and with the species. In many specles (for example,
Season of the year as a "variable" in procedures Bambusa vulgaris), the removal of a propagule (say a branch of a
";ven age) from the mother plant, at a given season of t?e ~ear,
As pointed out by the Rivieres (1879:461), practicallimitations on
fOllowed by its exposure to the conditions routinely mamtamed
the propagation of bamboos must be considered in relation to the in an outdoor propagating bed, is suflicient to break the. dormancy
season in which the initiation of the annual period of growth takes of one or more buds on the propagule. In other specles (for ex-
place. Published observations and personal experience agree in ample, Bambusa ventricosa), the buds of the same kind of a propa-
showing that, onee active growth has started, the tender shoots
involved are apt to be interrupted in their development if the 2Similar wide differences with season of the year are found in many woody cuttings.
parts on which they are borne are removed from the mother plant. Numerous examples are listed by Thimann and Behnke~Rogers (1950).
 260 . ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 261

gule of the same age, processed at the same time and exposed to centrations have not, to date, given results differing significantly
the same conditions, may remain dormant for as much as 2 years. from those wi th untreated plants.
Whole-culm cuttings show a similar diversity as between species, Because of the inexact nature and incomplete range cf the con-
in regard to the breaking of dormancy in their buds. trols usually maintained, the question of the best treatments and
No published account of the successful use of artificial means the best combination cf environmental conditions for economic
to break the dormancy of bamboo buds at will has come to my vegetative propagation of any bamboo, by means of any particular
attention. A satisfactory degree of success in the vegetative propaga- type of propagule, remains unanswered. Satisfactory' answers can
tion ofbamboos can be achieved only when routines effectively solv- be discovered only by means of aseries of e":periments in which
ing this problem have been established. U nder certain circumstances the environmental conditions surrounding the propagules are
(for example, when propagating material has to be transported) defined and precisely controlled. Furthermore, the propagules must
it would be useful to know also how to preserve the dormancy be processed and treated in such a manner as to break the dor-
and the viability of the buds for some time following the severing mancy of the buds. And finaIly, the rooting of the resulting new
of cuttings from the mother plant. . axes must be assured.
Propagules of any of the categories, from cIump divisions to
branch cuttings, sometimes fail for no obvious reason. In such Need of a neW approach to the study
cases it often turns out that, either through senescence, damage by of vegetative propagation in bamboos
insects, or accident, no viable buds were present on the propagule
The vegetative propagation of many bamboos can be made more
when it was put in the ground.
efficient by the accumulation of further experience through sys-
tematic studies of improved design, based on certain of the con-
Rooting ventional types of propagules. But the conquest of elite bamboos
New axes developed from buds on the buried part of a propagule that are refractory to conventional methods of vegetative propa-
da not always produce roots. That this may constitute a major gation requires the application of more knowled~e of a basic
problem has been demonstrated by data on whole-culm cuttings nature and more refined methods. Basic data reIatmg to varlOUS
and' culm seetions of Dendrocalamus strictus) culm seetions cf Bam- unsolv~d problems (such as the breaking of buu dormancy~ the
busa vulgans var. vittata, and stump layers of Bambusa longispiculata. rootin a of developing axes, and getting plants from a hlgher
The same difficulty was experienced in connection with rhizome perce~age of available buds) are needed in order to .Iift :he routine
cuttings consisting of the determinate rhizomes of caespitose procedures from the empirical level to a more sClentlfic one. It
tropical bamboos. The primary shoots that developed from the seems worth while, therefore, to make a serious attempt to de-
uppermost one or two buds at the distal end of these rhizomes velop routines and techniques designed to reduce the number of
usually produced no roots or, at most, an inadequate number. unassessed and uncontrolled variables to aminimum. The foIlow-
Therefore, one prerequisite to success in the vegetative propa- ina sc:hedule of studies is merely suggestive.
gation of bamboos is a knowledge of how to insure that each new 01. Selection of budwood (documented by name, age, position
axis will give rise to an ample supply of roots while the tissues in the plant, season of the year);
in the intercalary zones of its lower nodes are stilI in a condition 2. Treatment to break dormancy of buds;
to produce meristems. With a single exception (see Table 9 and 3. Isolation ofbuds, using sterile technique if it proves necessary;
pp. 251f), applications of various hormones, such as trichloracetic, 4. Incubation of buds in suitable media impregnated with
indoleacetic, and indolebutyric acids, at a wide range of con- nutrient solution;
262 ELITE BAMBOOS AND PROPAGATION METHODS PROPAGATION 263

5. Application of auxin, if necessary, to induce rooting of the Additional References. Gamus 1913:190-191; Houzeau de Lehaie
growing axis; 1906-1908:45-51, 216-218; Ferrer Delgado 1948; Mltford, 1896:
6. Progressive transfer of rooted propagules to potting media 12-18' McGlure, 1938a, 1945a; McGlure and Montalvo 1949,
and eultural eonditions favoring rapid subsequent development. 1950; , van Overbeek 1944; Pathak 1899; Piedallu 1931; Ueda
In any extensive investigation undertaken, the following vari- 1960:41-54.
ables should be studied individually:
1. Species or clone (begin with "elite" bamboos that are re-
fractory to conventional methods of propagation);
2. Part of plant from which the buds are taken;
3. Age of buds (with refinements to embrace season of year at
which buds are taken);
4. Methods of breaking dormancy in buds;
5. Time of application of treatment to break dormancy (in-
cluding application bifOre and after removal of budwood sticks
from the plant; and bifOre and ajler the removal of buds from
budwood);
6. Stage of development of the buds;
7. Gonditions of incubation (including temperature, moisture,
and quality and quantity of irradiation);
8. Nature of culture medium (substrate);
9. Formula of the nutrient solution;
10. Gonstituents of atmosphere (a tentative suggestion);
11. Time, manner, and rate of application of auxin;
12. Time of rem oval of rooted propagules to new culture
medium;
13. Gonditions for subsequent development of plants;
14. State of development of rooted propagules considered
optimum for various laboratory or field studies, and for nursery
or field plantings.
The development, or adaptation, of the appropriate refinements
of these procedures will require experience in the routines of
sterile technique, tissue culture, the breaking of donnancy in buds,
the use of hormones for stimulating root initiation, and so forth.
Equipment for the automatie control of environmental conditions
will be needed. Gomparable studies have been made on a number
of woody plants of temperate climates (see Thimann and Behnke-
Rogers 1950).
 We are now entmng an epoch I!f differential tCologica4 physwlogi.cal
and genetic classifieation. 1t is an immense work. The "cetm I!f knowledge is praetically
untouehed by biologists. 11 requires lhe joinl<labars I!f many different specialisl!i-
Physiologists, 'iJ'tolugisls, geneticists, systcmatists, and biochemis:/s. Part IU Bases of Classification
11 require, the mtcrnati01UJ.I spirit, Ihe eoupemtive work I!f investigators
throughout the whok world < •• it will bring us logical!y
to the n&Xt slep: integratim and S)7ltheru.
-N. I. VAVlLOV (1940)
1ncreased knowledge t1r:mugh improved methodologieal procedures
and a larger force I!f proper!y trained seimtist, working in the systcmatic area
0/ bwwgy ace the real k1ys to an improved classificatio1l.
-REED C. ROLLlNS (1965)
6 Flowering and fruiting behavior
in bamboos of different genera
and species

It is rarely poss; ble to observe the whole course of the reproductive

phase of the life of a bamboo plant. The recording of a ful! ac-
count of the significant aspects of the event is extremely difficult.
Parodi's simultaneous documentation of a complete life eyde and
complete reproduetive eyde of Guadua trinii (p. 275) is unique in
that it eomprises fust-hand observations recorded by a single in-
dividual on events from three successive generations of a given
hereditary !ine. Raizada's report (pp. 269f) of an equally weil
documented ease history involving events from three successive
generations of Bambusa copelandii is second to Parodi's only in the
cireumstance that it depends for its completeness upon thc
integration -of notes made independently and at different times and
places by three different persons. Seifriz obscrved two sueeessive
ftowerings of Chusquetl abietifolia (pp. 272f), but there is only cir-
cumstantial (not documented) cvidence that the same hereditary
line was involved in both events.
The follo,,~ng limited selection of published accounts is intended
to exemplify the di versity of recorded patterns of reproductive
behavior manifested by different kinds of bamboo-and even by
the same kind under different conditions and circumstances. Vvith-
out e"ception, they are fragmentary and disparate in respect to
details covered. In this regard the selected array of examples is
representative. More elaborate pcrspectives on the reproductivc
phase in the bamboo plant are offere<! on pp. 82ff. It will be helpful
to remember that in order to think clearly about the ontogeny of
a bamboo plant, it is necessaty to distinguish between its life
eyde and its fiowering cyde, since the two may not be eoe"tensive.
 268 BASES OF CLASSIFlCATION FLOWERING AND FRUITING BEHAVIOR 269
Arundinaria amabilis McClure eventually died, but the vegetative vigor ofthe plants apparently
Brief notes on the flowering and fruiting behavior of Arundinaria was not affected, and new culms 18 ft tall were produced simul-
amabilis are recorded on p. 152. taneously. Flowering continued in this way every year for 14 years.
Then in 1904-05 every remaining culrn flowered, and every plant
Arundinaria auncoma Mitford died. Every culm of plants of the same spedes growing in tbc
Bean (1907:229) says that flowers were noticed on plants of Arun- Ternperate House at Kew flowered in 1903, produced seed abun-
dinaria auricoma growing in the Bamboo Garden at Kew in 1898, dantly, and died. .
and that flowers were seen every year for nine years afterward. Mitford (1896:11) records in the follo"ing words his impression
Only a few of the culms were in flower at one time and, although of the behavior of this spedes upon flowering and fruiting:
these died, the vegetative vigor of the plants apparently was not Arundinaria sirrwni [sie] furnishes One exception of a bamboo which
adversely affected by the flowering. flowers in England without dying. It has not infrequently borne seed in
this country, and has been apparently none the worse. Last year (1895)
Arundinaria jmmila Mitford it flowered and seeded in more than one English g'arden. I myself gathered
seed from the culm of a large elump in a garden in Surrey. The remain-
It is recorded by Miss Rebecca Jones in a letter addressed to the ing culms were all in their normal condition, and there was no sign of
U.S. Department of Agriculture, May 6, 1935, that her plant of the leafy stem, being replaeed by flower-bearing branchlets, or of any
Arundinaria purnila ceased all vegetative activity and f10wered heavily injury to) or exhaustion of~ the plant.
in 1934. The flowering culms died, but the rhizomes survived and Parodi (1955:134) records that, in Argentina, plants of Arun-
the plant began to recover its vegetative activity the following dinaria simoni; (as Pleioblastus simonit) flower partially, and recover
year. Plants of the same origin and lineage f10wered at the same their vegetative vigor afterward.
time in Golden Gate Park, San Francisco, and fruited sparingly.
It is not known whether. they survived. Arundinaria simon;; var. variegata Hook. f.
Bean (1907:231) refers to thc record (Bot. Mag. [London] t. 7146)
Arundinaria pygmaea (Miq.) Aschers. & Graebner
of the flowering of Arundinaria simonii val'. variegata in 1877, and
No published record of the f10wering of Arundinaria pygmaea since states that it was ftowering again at Veitch's Nursery, Coombe
the description of the sterile plant, as Bambusa pygmfUa by Miquel Wood, England, in .1907, after 30 years.
in 1866, has COme to my attention. Perhaps because of having to
depend wholly upon vegetative characters for its taxonomie dis- Arundinaria vanegata (Sieb.) Makino
position, various botanists have in the meantime placed it under Arundinaria variegala was first described from sterile living plants by
the genera Arundinaria, Sasa, and Pleioblastus, -with either specific or Van Houtte in 1863, under the provisional (and therefore ille-
varietal status. It now appears that two different spedes, both gitimate) name Bambusa fortunei. Although this Chinese plant is
persistantly sterile, have been grown in ornamental plantings and still popularly referred to by the trivial name "Fortunei," its
in Iiving collections of bamboos, under the name Arundinar;a legitimate name i, Arundinaria variegata, based on Bambusa vanegala
pygmaea. The original plant was described as having solid culms. Siebold. I have not found any published record of lts having
Arundinaria vagans Gamble is the name now applied to the one flowered during the century since it first became known to science.
with hollow culm internodes.
Bambusa copelandii Gamble. Syn.: Sinocalamus copelandi,: (Gamble)
Arundinaria simonii (Carr.) A. & C. Riviere Raizada
Bean (1907:231) records that odd culms ofthis .pedes Were f1ower- According to earl y correspondence made public by Raizada (1948),
ing in the Bamboo Garden at Kew in 1892. The flowering culms Bambusa copelandii flowered and fruited in the Northern Shan
 270 BASES OF CLASSIFICATION FLOWERING AND FRUITING BERAVIOR 271

States, Upper Bunna, in 1896. Fruits eollected by Copeland, and etative growth, the plants produced a lighter crop of flowers and
docwnented by herbarium speeimens deposited in the Herbarium fruits the second year, then died.
at Dehra Dun, were planted by Gamble in the same year. The In the United States, the cultivar "Alphonse Karr" of this
resulting plants fiowered in November 1943, and produeed mature species has been observed on several occasions to remain in a
fruits in May 1944 (Fig. 55, 22). This indicates a fiowering eyde flowering condition, with noticeable vegetative growth and a
of about 48 years. From the fresh material available to hirn, sparse production of fruits over long periods. Julian Nally, in a
Raizada prepared an ernended description and the first illustration letter to the U.S. Department of Agriculture, records that his
of this interesting barnboo to be published. plant ofthis cultivar began flowering and fruiting at Gotha, Florida,
When Raizada was contemplating the transfer of this species in April 1937, and was stin alive and in a reproductive state when
to thc genus Sinacalamus, he sent his rnanuscript and a flowering the letter was "'ritten, in March 1953.
spadrnen to E. D. Merrill, asking for an opinion on the advis-
ability of the proposed action. Merrill referred the inquiry to rne. Bambusa tuldoides Munro
I wrote to Raizada indicating that the proposed transfer appeared Plants of a given species may show diverse flowering behavior
to be reasonable in the light of the resemblanee of thc inflorescence under different environmental conditions. In Bambusa tuldoides, as
of Bambusa copelandii to that of B. oldham;, which I had already observed in southern China, death of the plant usually follows
placed in the genus Sinocalamus. Raizada then published the promptly upon flowering, even though very few seeds are produced.
transfer and added, inadvertently as his own (1948: 10), thc last But individual plants of this spedes from southern China that
paragraph of rny letter to hirn. These are rny words: "I am, how- were introduced in 1925 to the United Fruit Company's arboretum
ever, not at a1l sure that the genus Sinacalamus as it is known at Lancetilla, Honduras, have ShO"'iIl some culms in a flowering
today will remain sharply set off frorn either Dendrocalamus Or state more or less consistently ever since, 'with no apparent diminu-
Bambusa when more of the bamboos of southeast Asia are studied tion of their vegetative vigor. There is no record of any of the
in the field." Later studies have indeed improved my perspectives, plants having either produced fruits or died.
and I now no longer recognize (maintain) the genus Sinacalamus
which I set up in 1940. Chimonoliambusa fa/cata (Nees) Nakai
Munro (1868:26) !im Chimanobambusa (as Arundinan'a)falcata among
Bambusa lineata 1\'1 unro the bamboos that flower annually. Gamble (1888:309) confirms
According to Gamble (1896:47), Bambusa lineata remains in the this flowering habit. Houzeau de Lehaie (1906-1908:33) reports
flowering state constantly at Peradeniya, Ceylon, and at rhe Royal complete flowering, and the fruiting of a small number of 30-year-
Botanic Garden, Calcutta. I saw the plants at Calcutta still in old plants in 1904, adding that the plants almost always die after
flower in 1954 and in the same year at Bogor, where they are said fruiting (cf. Gamble 1921).
by the Director of the Gardens to have been in flower constantly Bean (1907:230) writes of a doubtfully distinct form of this
since their introduction, which took place at least 100 years ago. speeies, under the name Arundinaria falcata var. glomerata, that a
Rhind (1945:14) characterizes this speeies as "constant-flowering" certain nwnber of culms of a plant growing in the Temperate
in its native habitat. House at Kew flower almost every year, and that the plant as a
whole does not suffer.
Bambusa multiplex (Lour.) Raeusch.
Plants of Bambusa multiplex, representing two different accessions ChimoMbambusa hookeriana (Munro) Nakai
tO my bamboo garden at Canton, China, flowered and produeed a . The first recorded observation on the events relating to thc be-
heavy crop of viable seeds. Without making any further veg- havior of Chimonobambusa hookeriana at and after flowering is that
 272 BASES OF CLASSIFlCATION FLO"VERING k'U) FRUITING BEHAVIOR 273
of its discoverer, for whom it was named Arundinaria hookcriana to die down and apparently every plant in the ;sland UamaicaJ
by Munro (1868:29). Hooker writes (1854:1, 29): died, root and all." Bean continues: "It is a curious fact that a
Near the top [of the pass from the Teesta to the Great Rungeet plant [of the same speeies] growing in the Palm House at Kew
Sikkim] I found a plant of "Praong" (a .mall bamboo), in fnll seed~ flowered at exaetly the same time and died."
this sends up many flowering branches from the root [rhizome], but fC',,~ Seifriz (1920) describes, from personal observation, the general
leaf~bearing ones; and after maturing its seed, and giving off suckers flowering and death of plants. of the same speeies in the same
from the root, the parent plant dies. The fruit is a dark, long grain, like general locality in 1918-19. This cireumstance suggests that the
rice; it is boiled and made into cakes~ or inte beer, like Murwa.
life eycle of Chusquea abieliftlia is about 33 years, but the evidence
Bean (1907:230) rccords that plants of this species (as Arundi- that the plants involved in the two flowerings are of the same
naria hookcriana) growing in the Temperate House at Kew flowered hereditary line is only circurnstantial. Both flowerings were followed
in 1899, bore seed, and died, while at Glasnevin, Dublin, where by an abundant production of frnits and the wholesale death of
it flowered at the same time, some plants died while others sur- thc fruiting plants.
vived and recovered their vegetative vigor. The disc,?very of a few non-flowering plants of the same species
was recorded by Seifriz. This is circumstamial evidence that not
Chimonobambusa quadrangularis (Fenzi) Makino all plants of Chusquea abieliftlia are monoperiodic. However, the
Since early times the curious squarish stern and spinous culm possibility exists that these plants did not represent the same
nodes of Chimonobambusa quadrangularis have attracted the attention hereditary line as the one whose members flowered gregariously,
or that they came from seeds produced late in thc previous gen-
and interest of botanists and plant fanders in China and Japan.
eral flo1,.vering.
Native of China, it was carried to Japan long ago, probably during
Flowering material of Chusquea abietiftlia from living plants
the period when Buddhist missionaries from China were exercising
from] amaica growing at Kew is described and illustrated by Hooker
their first great influence in] apan. It was introduced into Europe
(1885).
early in the 19th century; thus it has been known to modern
botanisrs of both hemispheres for weil over 100 years. There is
no record of its having flowered during this time. Although the Dendrocalamus hamilionii :Nlunro
flowers of the square bamboo were unknown to hirn, Nlakino
MUffiO (1868:152) credits Hooker with the observation that plants
plaeed this species in the genus Chimonobambusa on the strength of
of Dendrocalamus hamiltonii flower every year. M unro states that in
certain vegetative morphological characters and the habit, rare
the variety named edulis the flowers occasionally show the most
in hardy bamboos, of sending up new culm shoots in winter or
capricious variations. "Some flowers contain 3 stamens tolerably
late autnmn. This physiological peculiariry is the basis of the
name Chimonobambusa (Gr. cheimön, winter). perfcct, the remaining 3 being converted into sryle-like processes;
sometimes there are two distinct sryles, and sometimes the style is
4-c1eft; oceasionally there are 7 stamens."
Chusquea abietiftlia Griseb.
Two suecessive flowerings of Chusquea abieliftlia in the Blue Moun-
tains of ]amaica have been recorded. They are cireumstantially Dendrocalamus strictus (Roxb.) Nees
related to the same hered; tary line. In reference to the first The flowering habits of Dendrocalamus striclus have been given much
(1884-85), Bean (1907:231) quoted the following statement from attention, and many fragmentary notes have been published
the writings of an eyewitness, Sir Danie! Morris (Gard. Chron., (Suessenguth 1925:513; Blatter 1929:913). There is no documented
OCt. 23, 1886, p. 524): "V,'hen the seed was set, the sterns began record of successive flowerings of the same heredi tary line, but the

- - - ._---- ._~ ._-_._- ._-

 214 BASES OF CLASSIFICATION FLOWEFlNG Al'lD FRUTING BEHAVIOR 275

flowering eyde has beenestimated variously as 20-65 years. Troup propensities has recently eOme to light. It was secured at Milagro,
(1921:1006) says: Ecuador, in 19241 by Johannson and was established in eultivation
at Summit, Canal Zone. Although at the time of its introduetion
This species cerumonly flowers sporadicallYr in isolated dumps cr in it was not suspected of differing in any way from the 'last popu-
small groups, almost every year; it abo flowers gregariously over large
lation of plants of Guaduo. anguslifolia that ranges through the low-
tracts at lang intervals, the gregarious flowering usually taking SOme
years to compiete and often progressing in adefinite direction in succes- lands and up to elevations of 5000 ft in Ecuador and Colombia,
sive years. ActuaUy it is often difficult to distinguish bet'V\.recn sporadic the Milagro plant was discovered later by Walter R. Llndsay to he
f1o ..;ering on a plentiful seale and gregarious flowering, there being all flowering very frequently, often in successive years, at Summit.
stages between thc two ... Thc correct periodicity) however, 15 difficult Plants from the Milagro strain, established under my direction at
tO determine in view of the irregular manner of fiowering. Choco!ä, Guatemala, and at Tingo Maria, Peru, have shown the
same disposition to flower annually. Kennard (1955:193) rccords
Fruiting is generally abundant during gregarious flowering. The
the fact that Guo.duo. anguslifolia flowered and set a few froits on
plant usually dies after heavy fruiting but may survive partial
1- and 2-year-old culms of YOlmg plants vegetatively propagated
flowering (Houzeau de Lehaie 1906-1908;33). Munro (1868:148)
on the grounds of the Federal Experiment Station at l'vfayagilez,
states that the plant "does not die after flowering."
Puerto Rieo, in 1944 and 1945. This record obviously refers to
Several easeS of preeocious flowering in Dendrocalamus slriclus are
on record. Brandis (1899;22) quotes from the 1897 Report of P. "" .
the Milaf!To strain whieh is the onlv introduction of this species
under cultivation at Mayagilez. In April 1948, I saw flowers on
Sunder Lal Pathak, Conservator of Forests, Patiala State, India,
the original plant introduced there, and also on plants representing
the following details. Seeds collected in J une 1894 were planted
1-year-old vegetative propagules (offsets) taken from it.
early in March 1895. In February 1896, all of the seedlings were
lifted and planted in baskets. In April 1896, live of the plants
began to flower at the age of about 13 months. It is not scated Guadua Irinii (N ees) N ees
whether any seed was produced, but the death of at least three of Dutra (1938:147-150, Fig. 1) was impressed by the scant production
the plants within 3 or 4 months after flowering is recorded. Birbal of viable seeds by abundantly fiowering wild plants of Guadua
records the discovery of 5-year-old seedlings in flower in a nursery Irin;; (referred to by the author as G. riograndensis Dutra, but the
where they had been abandoned. Fertile seeds were produced. identity of the plant is weil documented).
Ahmad (1937) reports that, among the 2-year-old progeny from Parodi (1955) personally observed the flowering and fruiting of
a lot of seeds of this species collected in May 1935 and sown in a plants of Guadu.a lrinii representing twO successive generations of
nursery at Karnpur Royal F orest, India, at the break of the mon- the same hereditary line. Tbe elapsed time from seed to s""d was
soon in J uly of the same year, a single seedling was found in a exaetly 30 years. Parodi's unique data are doeumented by preserved
flowering condition in 1937. (See also Sen Gupta 1939.) Apparently specimens of the flowers and fruits. After flowering and fruiting
there is no record of. any strain of this species in whieh precoeious heavily one year and flowering again in the seeond year with
flowering is a heredita.ry character. See Oxytenonthera abyssinica, little or no production of fruits, the parent plants died.
p.276.
Melocarma bacciftra (Roxb.) Kurz
Guaduo. anguslifolil1 Kunth See pp. 195f.
Although the vast majority of the living plants of Guaduo. angusti- l.Not 1929, as pUblished by Higgins and Lindsay (1939:51), according to Walter R.
folia appear to flower rarely (if ever), a clone with quite different Lindsay (personal ccmmunication).
 276 BASES OF CLASSIFICATION FLOWERING AND FRUITING BEHAVIOR 277
;Yferostachys fistulosa Doell obtained from the Forestry Departtnent was sown in 1930 and produced
Under the vemacular name Tacuara lixa, !vferostachys flStulosa, a a fuw seedlings. These seedlings were planted out in the field in the
follm.v1ng rainy season and two of them bore seed for three successive
bamboo native tO" Brazil, is given a reproductive cycle of about years immediately followiog the planting. After IM third year setding ceaud
32 years by Pereira (1941:193). The plants are said to die com- hut the plant! did not die. All of the seed so produced was sown and about
pletely after fruiting heavily. The pertinent observations were made 1 percent proved to be viable. The resulting plant. behaved io exactly
in connection with the study of the plagues of rats that develop the same way as their parents, that i5, they bore seed for the first three
when bamboo seeds became available in quantity, a phenomenon years of their life. By collect.ing and sowing seeds regularly from each new
recurrent in southern BraziL plot, Mr. l'-.1acFadyen has carried out a progressive establishment of
bamboos over several acres of land. Tms seeding of young plants has
never been reported before and is of great interest. \'Vhether it is duc to
Nastus elegantissimum (Hassk.) Holttum seme heritable trait in thc individual plants first SQVffi, or whether It is
due to some Ioeal climatic or soil factors, is not known.
Kurz (1876:257) writes of Nastus (as Schizoslachyum) eleuantissimum
"Remarkable is bamboo ul-ul of Bandäng UavaJ, which flowe~
and dies off every third year." Phyllostachys spp.
Signs of unlimited longevity in bamboos are not restricted to
Ochlandra scriptoria (Dennst.) C. E. C. Fischer plants that have never been observed to flower. The flowering of a
Gamble (1896:122) quotes Bourdillon as saying of Ochlrmdra scrip- number of species of the genus Phy/lostachys has come under my
toria (as O. rheedz"l) that it flowers annually, not dying down after personal observation. In all cases the plants remained in a flower-
flowering. ing state, without maturing any appreciable amount of seed,
for periods varying from 5 to 10 years. The onset of flowering was
Ochlandra stridula Thwaites generally sudden, and was accompanied by a more or less complete
Gamble (1896:123) says, on the authority of Dr. Trimen, that suppression of the vegetative gtowth of the plant beyond the
plants of Ochlandra stridula flower regularly every year in Ceylon. production of flowering twigs from existing buds on the various
segmented axes. Following the period of exclusive flower production
Ochlandra travancorica Benth. and the death of the flowering culms, vegetative activity was
Gamble (1896:126) quotes Col. Beddome, the discoverer of "graduallv, revived , with the emergence of new culms from the rhi-
ZOrne system and a corresponding recession of flower production.
Ochlandra travancorica, as sa}~ng that "i t flowers almost every 7
Unless recorded indications of continual flowering in any species
years and dies down" (see p. 199).
of bamboo are accompanied by evidence as to the period during
which a particular plant was under observation, the statement
Oxytenanthcra abyssmica CA. Rich.) Munro
should be viewed with reservation, A c1ump of Phyllostachys nidularia
According to an anonymous report issued by the F orestry Depart- (P.L 66718) prnduced flowers at the U.S.D.A. Plant Introduction
ment at Zomba (Kyasaland Agt. Quart. Jour. 4: 10. 1944), Oxy- Garden at Savannah, Georgia, during the years 1943-1955. If the
tenanthera abyssinica is generally regarded by the inhabitants of plant had been seen by me only during this flowering period and
Kyasaland as having a flowering cycle of 30 years. A case of if I had not seen it in a vegetative condition either before or after-
precocious flowering and fruiting (paedogenesis) in this species is ward, I might have assumed it to be a species that flowers con-
recorded in the same report in the following words: tinually. Porterfield (1926:257) says: "Every year for the past three
A curious seeding feature of the eommon bamboo has been reported years at Wushih, Soochow, and on :Mokansan [China] the writer
from the estate of R. MacFadyen, Esq. in the Lilongwe dismet. Seed has collected flowering culms of Phyllostachys nidularia. In every
 2i8 BASES OF CLASSIFlCATION FLO'WERlNG AND FRUITING BEHAVIOR 279

case they were isolated culms with many normal vegetative culms is on record as having ftowered at the same time in the wild in
about them." Sikkim. Some plants at Trentham flowered in 1875 and others at
There is evidence that certain environmental (ecological) com- Holland House did not flower until 1877. The cultivated plants
plexes may be responsible for keeping plants of a given speeies all died after producing viable seeds freely, giving rise to a new
constantly at or near the threshold of ftowering, whereas in other generation. Odd culms among the progeny flowered at Kew duri,:g
conditions plants of the same species exhibit gregarious flowering 1893 and 1894. Flowering of this species occurred elsewhere m
only at long intervals. According to personal experience and a England in 1903, 1904, and 1906. Bean states further (p. 233) that
few published records, Phyllostatf!Jis aurea generally shows a long the ftowering of the [then] present generation of this species had
purely vegetative phase (15 to 30 ycars) followed by gregarious already (in 1907) extended over five seasollS. Whether any of the
flowering and subsequent restoration of vegetative activity. This lants of this rreneration survived is not indicated. Stapf (1904a:306)
P "
bamboo is widely used as a hedge plant in Guatemala City. Every states that thisspecies ftowers at an age of 28 to 30 years. D unng
. a
time I have been there, perhaps a dozen times over aperiod of personal conversation at Kew in 1928, Stapf informed me that
10 years, I have seen in these hedges, here and there, individual seeds of Thamnocalamusfalconeri were received at Glasnevin, Dublin,
culms in ftower. No fruits have been found and no gregarious in 1873. Plants propagated from them flowered and fruited in 1900
ftowering has been observed in these hedges. No flowering was and 1901.
observed in the several dumps of this bamboo seen elsewhere in
Guatemala. Thyrsostachys oliverii Gamble
Kawamura (1927) summarized recent and historical records According to Bor (1941), seeds and specimens of Thyrsostachys
pertaining to the ftowering of several species of bamboo in Japan. oliver;i from the Katha Distriet in Upper Burma were sent by
The author conc1udes that the ftowering eyde for Phyllostaehys J. W. Oliver to J. S. Gamble in 1891. Some of these seeds were
bambusoides, Ph. pubeseens (as Ph. edulis), and Ph. nigra cv Henon planted at Dehra Dun and some at the Royal Botanic Garden,
(as Ph. nigra var. henonis) is roughly 60 years. For Pf!Jillostachys aurea, Caleutta. The dates are not indicated in Bor's account, but pre-
he indicates that only one ftowering period (1916-1921) is on sumably the plantings took place in the same year. Toward the
record in Japan. He shows that the ftowering state in a given end of November 1938, Bor observed signs of ftowering in all of
species often extends over a number ofyears, and that the duration the clumps at Dehra Dun. The first inrucation noted was the
of the flowering state varies from speeies to species. fading and falling of the leaves. On December 23, 1938, the first
ftowers were in anthesis, and ripe seeds were found on J anuary
Sasa tessellata (Munro) ::Makino ct Shibata 31,1939. The length ofthe reproductive eyde for this species may
Bean (1907:232) says of Sasa (as Bambusa) tessellata that it had therefore be calculated at about 48 years.
been in cultivation for "probably over sixty years" without ftower- Bor took advantage of the unusual opportunity afforded by this
ing. And to this day, 115 years after its introduction into cultiva. event to record many interesting features of the flowering and
tion, no published record of the ftowering of this species has come fruiting of thc plants. The reproductive structures were also de-
to light. scribed anew, and illustrated in great detaiL

Thamnocalamus faleoneri Munro

Bean (1907:230) records that plants of Thamnocalamus (asA1"!<ndinaria)
falconeri from seeds sown in 1847 flowered simultaneously in 1876
in cultivation at Kew and at Edgecumbe in England; the species
 BAMBOO TAXONOMY 281

show notable variation in form within a given segmented axis.

Moreover, the bamboos usually are very fragmentarily and un-
evenly represented by prepared specimens, even in the best col-
lections. These eonditions greatly complicate the task of comparing
7 Bamboos from the point of view them with each other and with the other grasses, and satisfying
thc requirements of our accepted system of plant dassification.
of taxonomy This is a task to which the traditional methods, materials, and
techniques of taxonomy are not perfect!y adapted. For these
reasens, seasoned agrostologists recognize the tentative nature of
even the most conscientious and sophisticated treatment of bam-
000 classification.
The bamboos have not as yet been set off sharply, by definition, The original description of the subfamily Bambusoideae (Asch-
from the rest of tbe grass family. The seareh for the bases, and erson and Graebner 1902:769) fr~."ly translated from the German
the line, of demareation between the bamboos and the other mem- text, folIows:
bers of the Gramineae still continues. It is mOre than 170 years
sinee the first bamboo genus, Bambusa (as Bambos Retzius, Obs. Usually stately to large grasses, in the Tropics often gigantic (with us
. Bot. 5:24, 1789) was describcd on the basis of a single species, now shrubby or semishrubby). Rhizome usually short, creepmg, t?e plants,
called Bambusa arundinacea. During this time 75 effectively published because of this, making a thick ground cover. Sterns usually stiffiy erect,
the tops as well as the branches usually nodding to pendulous. Branche..
generic names and over a thousand specific names have been usually numerous1 orten several to many arising from one node, the u~tl~
attached to plants generally recognized as bamboos. However, many mate twigs solitary, in distichous~ horizontal arrangement. Leaves. Wlth
of these names arc either "illegitimate" or destined to fall into long-persistent sheaths, the articulated blade usually lanceoIate to linear-
the limbo of synonymy. lanceolate. Inflorescences often very !arge, paniculate} racemose or, more
The bamboos have been treated in the past by a few botanists rarely, spicate, often eIustered at the nodes. Spikelets 2-many-flowered.
Emptv g1umes [Deckspelzen] 2-several, the lowest ones ,mallest, progres-
(most recently by thc late T. Kakai) as a separate plant family,
sively' !arger acropetally, the uppennost shor:er than the ,1':'West lemn:a.
under thc name Bambusaccac. However, most agrostologists agree the lowermost emptv glurnes sometimes beanng sp,kelets tn thelr a.xiIs.
in retaining them in thc Grass Family (Gramineae). Not all con- Lemmas many-neIV~, awnless or, more rarely, with short tenninal awns.
temporary agrostologists agree with those who segregate the .bam- Palea 2-many-nerved, rareIy lacking. Lodicules usually 3, usually ver;
. boas from the other grasSes at the subfamily level. Some, perhaps large. Stamens 3, 6, or many. Styles 2-3, often fused at the base. Frult
the majority, prefer the more conservative procedure of'retaining not tightly eneIosed by the lemma and palea.
this and the other major divisions of the grass famHy at the tribe
The late Dr. Robert Pilger, in preparing his preliminary treatise
leveL Such a treatment connotes less profound, or less sharp,
on thc Grass Family for the new edition of the dassic German work
lines of demarcation. It also gives less taxonomie weight to the
of Engler and Prant!, Die Natürlichen l~flanzmfamilim, divide~ the
observed differences between groups. At the same time, i t accords
Gramineae into 9 subfamilies, of WhlCh the bamboos const,tute
greater recognition to thc many lines of similarity that form a
one as the Bambusoideae. However, Pilger (1954:290) gave only
complex network of relationships, through which comrnon charac-
abrief characterization of the bamOOos, which he ernOOdied in a
teristics link many genera of the Gramineae to two or more of the
proposed major divisions of the family. lIn bamhoos with the indeterminately branching type of inflot'escence dcscribed ~n
Most bamboos have a long and highly complicated life history, a:e
pp. 91:ff, of the present work, the so-<:alled "spikelets/' which tbe authors say borne In
the axi1s cf the lowest "empty glumes.," are acrua1ly spikelet·like brancltes (tne pseudo-
and an extensive plant body in which certain vegetative structures spikelets of McClure 1934).

- - .... ~-~ ~._-_ ... _~ . - .... _----_ ......- .. _- -

.. --~ ...
282 BASES OF CLASSIFlCATION BAMBOO TAXONOMY 283

key to the 9 suhfamilies. Translated from the German text, this aerial ,md subterranean segmented axes; insertion of adventitious
reads as folIows: roots circumaxial, principally subterranean, at nodes of rhizomes
and culm bases (in some species occurring also at aboveground
Culms persistent, woody, often very tall; leaf blades flat, mostly broad,
usually narrovved at thc base, petiolate, and articulated with the sheath; nodes of culms, and even on proximal internodes of midculm pri-
spikelet with two or more empty g1umes, [inflorescence axis (omitted)] mary branches).
often branched and deliquescent; lemma many~ner"V'ed) not awned) or Individual plants spreading either by the production of culms
with a shott, not geniculate awn; Iodicules usually three; stamens 3-6- from abranching system of rhizomes, Or by tillering from sub-
many. Fruit a caryopsis or drupe, to berrylike. terranean buds of culms without intercalated rhizomes. Rhizomes
A fuller view of the group is presented here in a somewhat generally within a given' genus consistently either pachymorph
modified equivalent of thc brief grammatical style that has been or leptomorph, rarely appearing in either one form or the other
made conventional in plant taxonomy by the example ofLinnaeus. in different species of the same genus, and more rarely mani-
Since many of the users of this description may not have a de- festing both forms in the same plant; the rhizome system sending
tailed knowledge of the family characteristics of the Gramineae, a up culms in either caespitose or diffuse array. Buds and branches
disciplined usage of taxonomy is disregarded here, in that charac- always solitary at rhizome nodes. Culms in some species very short-
ters common to the hamhoos and the other grasses are included. lived; more commonly persisting 5 to 10 years; in at least one
species (Phyllvstachys bambusaides) known to have produced a new
JlAMJluso'iDEAE (GRAMINEAE) crop of leafy twigs each year for at least two decades. Culms
Gramineae Suhfam. Bambusdideae-Rehder (1945:78; 1949: ranging in length (by speeies and in plants of mature stature)
635); Pilger (1954:290); Keng (1959:1); McClure (1961b:321). from about 0.5 to 45 m (exceptionally, to 90 m in Dinochloa
Gramina Subfam. Bambusoideae-Ascherson and Graebner andamanico, teSte Blatter 1929:602); of diverse habit: in some species
(1902:769); Prat (1936:217). self-supporting and wholly erect, more commonly nodding or more
Gramineae, Tribe Bambuseae-Kees (1829:520); Ruprecht or less pendulous at the tip; in other species weak and Gither de-
(1840:91; 1839 preprint, p. 1); Bentham (1883:1094); Hackel clined or straggling, or twining by the circumnutation of the grow-
(1887:89); Camus (1913:15); Hubbard (1948:300). incrb
shoots, or clambering, for which habit they may be equipped
Gramina Class X, Bambusacea-Kunth (1815:75). with special morphological adaptations. Culms branched (or bear-
Bambusaceae-Link (1833:308); Trinius (1835:613); Munro ing branch buds) at every node from base to tip in very young
(1868:10); Nakai (1933;11). plants of most species; culms in mature plants either branched
Perennial plants eonsisting of segmented aerial and subterranean throughout 6r, more commonly, unbranched and sometimes ev:n
axes bearing prophyllate buds (or branches) sheathing foliar organs, lacking buds (either at lower or mid-culm nodes or both) or, III
and branched or unbranched adventitious roots. Initiation of new some species, remaining unbranched throughout in the vegetative
vegetative growth typically seasonal, in temperate climates either state (Neurolepis). Each node of every segmented axis marked ex-
vernal, aestival, or autumnal; in tfopical climates~ commonly ternally either solely by the locus of insertion of a sheath (thc
sequ~ntial to the onset of a rainy- season; in warm regions with "sheath-node" of Haekel) Or also by an encircling supranodal
precipitation at frequent intervals throughout the year, often more ridge (the "culm-node" of Hackel) at the (slightly higher) level
or less continuous. Each segmented vegetative axis generally more of the insertion of a bud or a branch complement; nodes of all
Or less strongly lignified, increasingly so toward its surfaee; weakly segmented axes also marked internally by a diaphragm of paren-
so in part or throughout the plant in some taxa. Insertion of chyma reinforced by the crossing over and anastomo~ing of vascu:ar
prophyllate buds and the sheaths that subtend them distichous traces from the culm walls. Lower mid-culm nodes In some specles
(that is, ineident on alternating sides at successive nodes) on both las in Chimonobambusa quailrangularis) bearing, just above the sheath
 284 BASES OF CLASSIFICATION BAMBOO TAXONOMY 285

node, a ring of thom-like adventitious roots in arrested develop_ ligule regularly absent or rudimentary in the sheaths of certain
ment. Branch buds at culm nodes uniformly salitary in most genera axes (for example, necks), and elsewhere throughout the plant in
(more than one in all species of the genus Chusquea), a 2-keeled some spedes. Culm sheaths and branch sheaths always inserted on
amplexicaul prophyllum completely enclosing each branch pri- thcir respeetive axes at levels proximal (not distal) to leaf sheaths.
mordium. Branches at culm nodes usuallv, unarmed , in some Culm sheaths persistent, abscissile or, in some species, incompletely
species more or less strongly thorny, the thoms consisting of dwarfed abscissile; the lowest Ones bladeless but mucronate, the upper ones
and indurated twigs; thorny branches found only on the lower laminiferous, the blades sessile, persistent or abscissile, ·and usually
culm nodes in some spedes; in others appearing at higher levels as lacking a midrib. Leaf sheaths borne at the distal nodes of aerial seg-
weiL Branch complements most highly typical in form and com- mented axes of all orders; generally differentiated sharply from
position at mid-culm nodes; in some species monoclade: in others typical culm sheaths and branch sheaths in features that include
pleioc1ade; component axes in plcioclade branch complements she, shape, texture, and loeus of insertion of the sheath proper,
either subequal or unequal, thc primordial branch in the ,letter and the form of the blade; leaf sheaths proper normally deeply
case more or less strongly dominant; the branch complement at inbricate and persistent; laminiferous. Leaf blades petiolate and
mid-culm nodes either remaining static basally after the completion usually at length abscissile at the level of insertion of the petiole
of its initial grand period of growth or, more comrnonly, augrnented on the sheath (sessile and abscissing only very late, if at all, in some
by the subsequent developmenr of additionalleafy axes from buds species of Neurolepis and .Myriocladus); during development tightly
at basal nodes of one or more of the component branches. Culm rolled individually, on an "axis" parallel to the midrib, with one
internodes in SOme taxa sulcate for some distance above the in- margin at the center of the roll, afterward usually Hat or nearly so
sertion of a bud or a branch complement (grooved all the way from (commonly need1elike in Arthrostylidium capillifolium); a midrib usu-
one node to the next in all spedes of Ph)'llostach)'s); rarely some- aIly distinguishable (at least basally) and more prominent on the
what .square in cross seetion (as in mature plant, of Chimonobambusa abaxiaI surface, the midrib in some species depressed on the adax-
quadrangulans), otherwise cylindrical Or nearly so; either e;/istulose ial surface (strongly so in Neurolepis). Parallel veins (secondary and
with a pithy center (as in Chusquea) or fistulose, the lumen (the tertiary) of all leaf blades (as far as known) connected by trans-
"pith cavity" of Hackei) containing pith in diverse forms and in verse veinlets, these weakly to strongly manifest on one or both
different degrees of abundance, generally. negligible; fibrovascular surfaces, or entirely obscured to superficial view by overlying tis-
bundles of the culm wall set in a matrix of parenchvma , , . most sues. Foliage in plants with branched culms renewed periodically
c10sely crowded together toward thc surface, progrcssively more (common!y once a year) from buds at subdistal nodes, whieh give
Wldely spaced toward the center of the culm; successive internodes rise to leafy twigs replacing those lost by abscission; the renewal of
increasing
. in length (and sometimes also in diameter), below the foliage generaIly diminished or arrested upon the initiation of
mlddle of the culm; those above the middle decreasing in both flowering; plants of same otherwise evergreen species completely
length and diameter, acropetally. Each node of every segrnented deciduous under conditions of extreme drought (as in Bambusa vul-
a: is bearing a sheathing foliar organ (sheath); sheaths generally garis and Deiuirocalamus strictus).
Wlth the outer edge free throughout, but exceptionally (as in the Sexual reproduction monoperiodie or polyperiodic. Flowering
culm sheaths of some spedes of Chusquea) with the Outer edge adnate cycle, according to numerous but mostly weakly documented rec-
to the outer surface of thc sheath for some distance from thc base ords, varying (as between species) from 1 to 60 years Or more in
upvv~ard; sheaths proper either laminiferous, awned, or mucronate; length. Flower production commonly gregarious; in some popula-
laminiferous ones commonly "ith a bristle-bearing auricle Or a tions erratic as to timing; in some cases involving only a part of
tuft of bristles inserted at one Or both lateral extremities of the the plants, or only apart of the culms in individual plants of a
insertion of the blade; most sheaths as a rule apically ligulate, the given population; Howering in some polyperiodie species (as in
286 BASES OF CLA.SSIFICATION BAMBOO TAXONOMY 287

Arumlinaria amabilis and Phyllostachys nidularia under personal obser- the type species of the genera Schizostachyum and E1,ytrostachys)
vation) sometimes continuous for 5-15 years of a given eycle, full tubelike, a sulcus usually lacking entirely in the palea of a fune-
vegetative vigor being restored gradually as flower production sub- tional floret terminating aspikelet, exceptionally (as in some spe-
sides. Anthesis progressing acropetally in each spikelet. Froh pro- eies of Chusquea) a shallow groove appearing only near the apex of
duction abundant to sparse or nil-partial to apparently eomplete the palea; the palea ci ther more or less infiated and gaping anti-
self-sterility eommon in many species. cally Cas in most genera) or enc10sing the floral parts entirely; in
lnflorescences terminal or lateral to vegetative axes; consistently some genera (Oxytenanthera, for example) tightly eonvolute, wit? a
of either determinate 01' indererminate branching within a given small aperture at the apex only. Flowers generally hermaphrod,:e;
genus, Flowering branches leafy or leafiess. Determinate infiores- in somc spedes regularly, and in some spedes only under spec,~l
cences typically pedunculate (exceptionally reduced to a single eonditions, either the androecium or the gynoecium, or both, rod,-
spikelet) the branching habit panieulate, racemose, or spicate- mentary or lackihg entirely in part or all of the flowers of a given
raeemose, usually symmetrieal, rarely secund. Indeterminate infio- spike!et. Lodicules in some species (all of Gigantochloa sen~ str.)
rescences typically consisting of sessile pseudospikelets, these often lacking entirely, in others variable in number, in most sp~c,es 3;
forming' more or less dense tufts or heads. Bracts subtendihg pri- all alike in some, unequal and unlike in others, the antmor two
mary branches of determinate infiorescences generally w,eakly commonly asymmetrie al and palred (these in some speaes func-
dcveloped, Or (excepting the lowest) lacking entirely, those sub- tloning by turgidity to open the fiorets at anthesis, in ot?ers. not),
tending the lowest primary branch typically more strongly devel- the posterior onc symmetrieal, in some species reduced m Slze or
oped than those of the more distal ones, but in some genera lacking; all as a role showing vascular traces; all thin and trans-
reduced to a transverse !ine; bracts subtending the branches of all lucent in some species, somewhat thickened and opaque in others;
orders in indeterminate infiorescenees always weil developed, in glabrous on both surfaces or pubescenl abaxially, the margin usu-
some species all, or the lower ones only, laminiferous. Spikelets ally ciliate above, rarely glabrous. Stamens usually 3 or 6 (varymg
sessile or pedicellate, one-to-many flowered, terminating in either a between 3 and 6 in some speeies, as in Pseudosasa and Sasaella;
functional floret, a sterile floret, or asIender prolongation of the 6-120 in Ochlandra) generally in whorls of 3; the anthers usually
rachilla, this usually hearing a discernible rudiment at its tip; the either yellow or greenish yellow, rarely maroon when fresh and
lowermost floret in the spikelet usually preceded by one-to-several afterward brown (as in Bambusa vulgaris); the filaments either
(commonly 2) empty glumes, these unequal, increasing in size threadlike and free, or flat and either triadelphous or dladelphous,
acropetally, one or more sometimes lackihg or rudimentary. Ra- or all lightly connate-in some species monadelphous (inserted at
ehilla segments in several-flowered spikelets either short and not the mouth of a delleate tube, as in Gigantochloa); the anlhers at
disarticulating between florets or, more commonly~ somewhat first 4-10cular, the two locules of each pair uniting upon the ma.tu-
elongate, and in most genera abscissile immediately below the ration of the pollen, each cavity or sac thus formed then openmg
locus of insertion of the lemma of each functional floret (the dis- by an apical pore or by a slit that fo11o"'"8 a lateral suture; e~ch
artieulated florets stipitate in Guaduella). Lemmas usually mtiero- half of each anther extending as a lobe beyond the eonnectlve
nate or cuspidat:e, in some species muticous; fertile lemmas either proper above and below; the connective often exserted above, in
embraCing the palea fully only basally in funetional fiorels (the some species prominently, between the two halves of the anther,
florets then either open or c10sed at anthesis) or convolute and and terminating in a glabrous or penicillate point. Ovary nor-
tightly enclosing thc palea exeept for an aperture at it. very apex mally soJitary (sometimes duplicated in teratic flowers), 1-celled
in functional fiorets (the florets then not opening at anthesis). (with but a single' caviry) with 1 ovule, and terminating above in
Pa1ea in funetional florets usually with a longitudinal dorsal sulcus 1-3 styles, these (when more than one) united at the base and
extending from base to tip, the sulcus in some species (for example, upward for a distance varying with the spedes m some genera, in

----- ----,--
283 BASES OF CLASSIFICATION

others uniform; each style terminating in 1,2, or 3 (rarely 4 or 5)

stigmas. 2 Proliferations of the stigmatic surface of diverse character
and arrangement: usually capillary in form; smooth or papillate;
simple or branched; of bottlebrush, pinnate, or secund insertion
when simple; dendroid when branched; predominantly white, in
some species purple or red (exceptionally, white with purple ones Appendix r
Generic Key to
intermixed, aceording to Kur" 1976:266). Fruit a caryopsis, or Bamboos U nder Cultivation in the
caryopsisJike, normally one-seeded and indehiscent, sessile (rarely
quasi-pedunculate as in Cephalostachyum capitatum); usually falling United States and Puerto Rico
at maturity with the floral envelope still attached basally; differ-
ing widely in size, shape, and anatomy; the attaehment of the secd
clearly shown by a longitudinal groove or sulcus (the "hilum" of
HackeI) in 30me species, invisible or only weakly discernible exter- As currently classified in taxonomie literature~ the bamboos known to.be
nally in others; thc embryo relatively short-lived among the bam- under cultivation in the United States represent 21 genera. The followmg
boas as compared to that in other grasses generally-a d~rmant key indieates the characteristic features by which these g~era may be
stage apparently lacking in some species, as in Melocanna bacclfera, distinguishcd, The classification of many of these gcner~ stlll rests, how-
where germination often takes place before the fruit falls to the ever on tenuOUS and uneven criteria. Some of the genene concepts were
ground. originaIly based wholly Or largely on characters of their rareJy a vailable
Type genus: Bambusa Schreber (nom. eonserv.); type species: flowers and fruits. Other genera were segregated wholly on the basis of
Bambusa (as Bambos) .arundinacea Retzius. vegetative characters. . .
Pilger (1854:291) made a tentative suggestion that the eontro- Thc following key is based on vegetative charneters as far as IS posslble
versial genus Streptochaeta, eomprising the tribe Streptochaetae of at present. However, "it has been nccessary to rely upon cha.n:cters of ~e
reproductive organs for differentiating certain g~era. TI:1S n~cesslty
Hubbard, might be included in the subfamily Bambusoideae.
reflects the uneven charactenzation of genera that still prevaüs. Wlth the
Parodi (1961) has proposed thc addition of the tribes Olyreae, resent lack of popular knowledge of the barnboos, the identification of
Phareae, and Parianeae to this subfamily. However, these tribes '1'ISt.
prnany kinds must still be left to the specla
were excluded from consideration in the preparation of the fore.. Ov.ing to the still tenuous bases of their segregation as ?en~a} the
going deseription, whieh embraces only the "true" bamboos in names of Sasamorpha Cfrorn Sasa) and Plewblastus (from Arundlluma) have
the traditional sense: the tribe Bambuseae sensu lalG. In the taxo- not yet been aecepted for general use in the United States.
norme literature of this group there are 76 effectively published
la. Rhizomes pachvmorph, i.e.) the rhizome proper short, thiclc.er than the culm
generic names and over 1000 specific names. Ofthese names, many that emerges irom its tip, and not "running. " . .
are destined to fall into syoonymy. 2a. Rhizome neck much langer than the rhizome proper; clump habn dlffuse~
i.e.~ with widely'spaced culms; midculm branches slender. subequal.
250m<: agrostologists may prefcr thc following form of description for this part of the MELOCANNA.
bambusoid pisti1: Ovary . , . contracted above illto a stylar column; the stylar column in 2b. Rhizome neck shorter than the rhizome proper; clump habit caespitose, i.e,}
same species very. short (e'len subobsolete), in othet"S (some species of Arthros!yb'dium) but~
with culms rather dose together. ."
baus, more commonly slenrler and more or [ess elongate; rarely unbranched and terminat-
:Ja. Infiorescence of indetemrinate growth. i.e" the splkelet-hke . ulttmate
ing in a single 'stigma; usuaUy divided iota 2 or 3 (rarely 4 or 5) stigroatic branehcs.
branche, (tl'pical prophyllate pseudospikelet,) dothed basallyw.tb ";lall.
overlapping amplectant bracts, the bracts subtencling buds that contmue

lWith ruy permission, the material in ws Appendix was published by Young <L.'?d Haun
(1961:12-14).
 290 APPENDIX I GENERle KEY 291

for seme time to produce other branches (pseudospikelets); stamens 6; Branches at midculm nodes typica1ly in pairs~ robust, strongly
transver:sc veinlets of leaf blades not -visible or only weakly manifest. uncqual; occasionally with a smaU} third one devcloped betwecn
4a. Brauches at midculm nooes a11 stender~ subequal. them. Pln't.LOSTACHYS.
Culrn sheath proper ancl shea.th blade conspicuously thickened hard Brauches at midculm nodes typically 3 to 5 in each complementt
and brittl.c; the ~uricles of the culin sheath prominently dev~loped; slender. very short. subequal. SHIBATAEA.
style heanng 2 stIgmas. CEPHALOSTACHYUM. Sb. Internodes of rhizomes, Cl,llms, and branches cylindrica1 or nearly
~ulm sheath proper and sheath blade not conspicuously thickened or so.
mdurated; thc auricles of the eulm sheath not prominently developed' Culrn sheaths fugacious, abscissing completely; auricles and oral
style hearing 3 stigmas. SCHIZOSTACHYC~ setac typically very prominently developed (but sometimes l~ck­
4b. Branches at midculm nooes very llnequal, the primal)~ (middle) one in ing entirely) on culm sheaths; bracts at the base of pseudosplke-
each complement more or less strongly dominant, lets smaller than the lemmas~ without leailike apical appcndage.
5a. Culms weak, clambering, not self-supporting, the miclculm internodes SINOBAMBUSA.
very long ~nd ""Cl")" thin~walled, fragile and easily crushed- spikelets Culm sheaths abscissing promptly but incomplete1y, remaining
typieally with a single pencct fiower. ' lightly attached at the middle of the base for a shon: time after
Spikclets pediceHate, the lemmas and paleas loosely convolute' fruit they dryj aurides and oral setac lac~g on culm sheaths; b!acts
sulcate~ the pericarp moderatcly thickenoo at the very apex only, thc at the base of pseudospikdets larger than the lem:rnas, often
apex: crowncd by the pubescent, rather fragile, base of thc style. hearing a small leaflike apical appcndage. St:MIARUNDfXARJA.
ELYTROSTACHYS. ibo Inflorescence of dcterminate growth} thc ultimate branches tenni~
Spik.elcts not pedicel1ate, t~e lemmas and paleas tightly convolute; nating in pedicellate spike1ets.
frUlt not sulcate, the pencarp greatly thick:encd and indurescent 9a. Branches typically solitary at midculm nodcs.
throughout, thc ape.~ narrowcd and prolonged in a tapered persistent, lOa, Culm nodes mOre or less prominently int1ated above the
glabrous beak, OCHLANDRA. sheath sear; culrns commonly tillering, usually ascending,
Sb. Cu]ms se1f-supporting~ not elambering~ thc mideulm internodes not not strictly erect; oral setae of le.af sheaths scabrousj sta-
conspicuously elongate, the walls not fragile or easily c.""UShed. mens 6 in each flower. SAM.
fu. Ra:hilla segments very mort. nOt disartieulating. . lOb. Culm nocks not prominently swollen aoove the sheath $CaI';
Splkelets few-flowcroo, the terminal floret of each spike1et usual1y culms normal1y not tiIlering, more or less strictly erect.
weH developed, perfect; stamen filaments free. DEJ:>..'DROCALAMUS Oral setae wholly lacking in lea[ sheaths; stamens 6 in each
Spikclets severaJAlowered, thc terminal fioret- of each spilreJ.et rudi~ flowcr, SASAMORPHA,
mentary; stamen filaments fused or connccted by a membrane. Oral setae of leaf sheaths few or none, glabrous; stamens 3
GIGANTOCHLOA. (rareIy 4.or 5) in each flower. PSEUDOSASA.
6b. RachiUa segments elongate-up to half thc length of the lemma- 9b. Branchcs typically more than one at each mideulm node; sta-
promptly or tardily disartieulating. mens 3 in each fiower.
KeeIs of palea winged. GUADUA. lla. Culm shcath blade minute; rapid growth of shoots (new
Keeh of palea not wingcd. BAMBUSA. culms) initiated in autumn or late summer.
3b. Inflorescenee of determinate growth, i.e" emerging completc, the ultimate CHIMOXOBAMBUSA.
br~nehcs no: typical pscudospike1ets, and not continuing to b!,anchj l1b. Culm sheath blade weH developed; rapid growth of shoots
splke1ets pedicellate; stamens 3; transverse veinlets ofleaf blades clearlv (new culms} initiated in spring tO early summer.
manifest (obseure in Thamnocalamu.s falconenl ' Oral serac of leaf sheath seabrous. A.ltmrDINAIUA.
Lower branehes of the infiorescence prophyllate, cach subtended by a Oral setac of leaf sheath glabrous. PLElOBLASTUS.
weH-developed bract, THAMNOCALAMUS.
Lower. brauches of infioTCscence not prophyllate, not subtended bY a
well~aeveloped bract. SlNARUNDIN:UUA.
lb. Rhizomes leptomorph) Le., the rhizome proper long~ Hrunning," generally
more slender than thc culms that arisc from its lateral buds.
7a. Inflorescence of indetenninate growth, the ultimate brauches pro~
phyllate pseudospikclets terminating in nonpedicelIate spikelcts.
8a. Intemodes of rhizomes~ culms, and brauches each strongly flat-
tened Or channeled throughout its length on one side-above thc
point of insertion of a bud or brauch complcmcnt.
 BA.'\1BOO NURSERIES 293

B. ventrirosa. Buddha bamboo, 10, 16

B. uulga:ris. 7, 16
Dendrocalamus striclw:. Male baroboo. 16
Phyllostacf!Jis aurea. Fishpole bamboo, 2,4,5,6,7,9, 14, 16, 20
Ph. a.u:rcosulcata (misidentified in some earlier records as Ph. neviniz). Yellowff
Appendix II Bam boas Offered groove bamboo. 4, 9, 13
Ph. bambusoidßs (Pb.. quiliai; misidentified in some earl;er records as Ph.
and Nurseries Offering Them In reticulata). Madake; giant timber bamboo; Japanesc timber bamboo.
4, 8, 11, 16, 22
the United States Ph. bambusoidßs, cv. 'Castillon' (Pb.. castillom). Castillon bamboo, 4, 14
Ph. duleis (misidentifie,d in some earlier records as Ph. henry'). Sweetshoot
bamboo. 14
Ph. pubeseens (Ph. edulis). Moso bamboo. 14
Pb.. jlexuosa. 14, 21
Ph. meyeri. Meyer bamboo. 11, 11
Thc following list is adapted, 'wirh some minor ehanges, from the CUTrenr Ph. nigra (Bambusa nigTa), Black bamboo. 4, 14, 17
(1960) revision ofYoung [1960]. For a fuHer list of bamboo names likely Ph. nigra, cv. 'Henon' (Ph. pubernla; Ph. herumi.<; Ph. nigra var. h,"onis),
to be encountered in horticultural literature, the reader is referred to Henon bamboo; haehiku. 4
Kelsey and Dayton (1942:40-41). Ph. nucla. 3, 19
Synonymous names and alternative names sometimes found in nursery Ph. viridis (Ph. sulphurea var. viridis). 14
eatalogs appear in parentheses; common names in current use follow. Ph. viridis) cv, 'Robert Young' (misidentified in some earlier records as
(Synonymies listed here are not repeated in the Index of Scientific Names,) Ph. sulphlUea). 14
Numbers refer to nurseries Iisted below, Pseudosasa japoni,a (Arundiruria japoniea; Sasa japrmica). :Metake; Yadake;
arrow bamboo, 4, 6, 12, 14, 16, 18, 21
BAMBOOS Sasa palmata (Arnndinaria palmata; Bambusa palmata; misidentified in some
Arundinaria amabilis. Tokin bamboo; tonkin cane; tsingli cane. 14 earlier records as Sasa senanensis). 1, 13,21
A. gigantea (A. rruu:rosperma). Canebrake bamboo; Southern cane. 11 S. pygmaea (Bambusa pygmaea). Pygmy bamboo. 4, 21
A. simonii (Pleiablaslw: simoni,). Simon bamboo. 13, 14 S. tessellata (Sasamurpha tessellata). 4
A. virid:"-striata. 1 SemiarundinariafastUtJsa (Arnndino.riafast!IIJsa; A, nar!hira). Narihira bamboo.
Bambusa rrwltiplex var. riviereornm. Chinese goddess bamboo. 18 4,21
Shibatea kumasasa (S. kumasaca; Bambusa kumasa,a; B. ruscifoli.a; Phyliosta,hys
B. multiplex; typical form. (B. llalla var. 1lormalis; B. argenteu.; B. naP..a argen~
kumasa,a; Ph. ruscifolia). Okamezasa. 4, 21
I<a). Hedge bamboo; oriental hedge bamboo. 6, 14, 16
B. mult,plex, cv. 'Alphonse Karr' (B. nana Alphans< Kam; B. Alphanse Kam).
NURSERIES1
Alphonse Karr hedge bamboo. 6, 7, 12, 14, 16
B, multiplex, cv. 'Fernleaf' (B. 1U1JU1; B. nana dislic/w.; B, nana var. gracillima). 1. j, Herbert ."Jexander, Mlddleboro, Massaehusetts
Fernleaf hedge bamboo. 4, 11, 15 (listed by thi. nursery as B, falseal", 2. Armstrong Nurseries, Ontario) California
error for B. falcata, a misident;fication), 16, 18 3. George M. Darrow, Route I, Glenn Dale, Maryland
B. multiplex> cv. 'Silversmpe' (B. argentea striata; B. nana argentca striat~·, B. 4, Fruitland Nurserles) Augusta, Georgia
nana var. varlegata). Silverstripe hedge bamboo. 6, 14, 16, 18 . 5, Gardens of the Blue Ridge, Ashford, McDowell County, North
B. 7lUI.ltiplex, cv, 'Stripestem Fernleaf', Stripestem femlcaf hedge bamböO. Carolina
18 6. GIen Saint :Mary Nursery Co., GIen Saint Mary, Florida
B, oldhami (Sinocalamus old/w.mi; commonly misidentified as Dmdrocalamus 7. Griffing Nurseries, Beaumont, Texas
latiflorus). Oldham bamboo. 2, 12, 16 .
l"W'hile it rs irnpracdcable to provide a cornplete list of dealers, this partial list !s fur-
B. tuldoUks (B. paUescem; misidentified in some earlier records as B. Thou-
nished with the understanding that no discrimination is inte:tded and DO guaraotec of
am). Puntingpole bamboo. 12, 16 teliabiHty implied}> [Disclaimcr clausc requ:ired by 1aw. f.A.M.].
294 APPENDIX n
8, H, G. Hastings Co" P.O. Box 4088, Atlama2, Georgia
9. Jones Ornamental Nursery, Hobb, Road Nashville Tennessee
10. Batter King, Naples, Florida ' 1

11. ~. M. Kirk, McRae, ""'hite County, Arkansas

12. Lmk Floral Co., Weslaco, Texas
13, Lowland Rhododendron Farm, Kingston, Rhode Island (Iocal sale
only) Glossary
14. McIlhenny Nursery, Avery Island, L<>uisiana
15. McKee's Nursery, P.O. Box 749) Covington, Loulsiana
16. Reasoner', Tropieal Nurseries, 2433 Sixth Ave,. Bradenton, Florida
(Ioeal sale onlyl '
• There's a glory for you!"
17. Ja,,;,es A. Bateman, P.O, Box 114, Slidell, Louisiana H

18. Juhus R~rs Co., Rutherford, New Jersey "I don't knciw .what you mean by ~glory,'" Alke 'said.
19. Joseph W. Showalter, Pon Republic Maryland Humpty Dumpty srolled contemptuously, HOf course you don't-till I tell
you. I meant ~there's a nlce knock-down argum.ent for )'Ou!' l>
20. Charles W, Stegall Farm, Route 5, B~x 314, Montgomery, Alabama
(local sale only) "But 'glory' doesn't mean t a nice knock~down argument,'" Aliee objectoo.
~;When I use a word," Humpty Dumpty said, in rather scomful tone> "it
21. Upper Bank Nurseries, Ine., Media, Pennsvlvania means jUt>t: what I choöse it to mean-neither more nor less,"
22. Wilson Nursery, Magnolia Highway, Gulfp"rt, Mississippi (tThe question is/' said Alice , "whether you can make words mean so many
different things."
"The question 1S," said Humpty Dumpty, ~<which is to be master-that's alt"
LE'Wl'$ CA:RROLL

Tbe folloV\ring terms are defined with special reference to the usages I
have adopted for describing various characteristics of the bamboo plant.
Several of these terms (indicated by an asterisk *) are newly coined; some
others (indicated by a dagger t) are given new or extcnded meanings to
describe conditions not satisfactorily covered by conventional usage: or
by botanical glossaries. Published sources give additional (alternative)
meanings for some of these same terms-meanings that are in more gen-
eral (or special) use elsewhere.
It will be obvious to any perceptive person who undertakes to prepare
a glo,;sary that his path is beset with many pitfalls. Some of these traps
have been laid by uncritical conventional interpretations, or by the
inadvertent misuse, of established terminology in technical papers1 glos-
saries: and dictionaries. Other dangers lurk in the labyrinths of semamics
and syntax. -The improved light cast by the growing body of recorded
observations may have changed, progressively, the mental image of what
is to be described. 'Tbe diversity of the products of morphogcnesis, as
they appear in the different botanical groups, demand, alertness to the
need for critical discrimination in the use} or thc modification, of the
traditional sense of conventional terms. Ambiguities and solecisms may
subvert the most determined eifort to produce definitions that are both
accurate and cle;ar. The end product of thls eifort may never be pushed
beyond the reach of constructive criticisrn and revision. In any case>

..
"-_ _- .. __ ... _~-._'------
296 GLOSSARY GLOSSARY 297

unremitting scholarship will not go unrewarded in thc long run, for a arise independently, both at the right and at the left :,f the one at the
clear definition of tenns is an .aid to dear thinking. usual site, which is median to the base of a subtendmg culm sheath.
Lengthy annotations arc appended to some of thc definitions in this See Const,llatc.
glossary. These are designed tO promote the reader's understanding of the AnasttmWs1S (Gr. anastomiisis, a new outlet). "Union of one vein . . . .-ith
special pertinencc of the terms tO the morphology of the bamboo plant. another, the connection forming a reticu1ation)j Gackson).
Thcy should also bring thc bamboos into clearer- perspective in relation Androetium (Gr. aTu1:tOs) male; oikos) house). The stamen complement of a
to thc other Gramineae. However, onIy a first-hand study of the bam- fiower I have not followed Arber (1934:152 et passim) in adopting the
hoos themselves can adequately iIluminate some of the features found spelli~g (andrae<:eum) advocated by Kraus (1908). Brown (1954),
exc!usively in this group of plants. Jackso n , and Nybakken all give androedwn. Cf. Gynoeczum• .
t AnticaliY; a.w. anticoosly (L., forward, in front). Toward the front; m-
Abaxial (L. ab, away from; axis). Away from an axis;-designates that trorsely or adaxially. Jackson reeords diverse and even a~tonymous
surface of an appendage (such as a culm sheath or a sheath blade) definitions of antical, mentioning the synonymo~ use of Introrse as
which faces, in a structural sense, away from the axis on which it is "occasional." The bamboo palea often gapes anucally.
inserted or relies for support. The terms abaxial and adaxial (q.v.) • Apsidate (L. apsis, areh). In convex arcuate array;-indicates the pattern
should bc employed instead of upper and lower, or dOTllal and ventral, of insertion of the components of a pleioclade branch ~mplement)
whose usc--with reference, to the !WO faces of a foliagc leaf or a where the individual axei of thc primary array emerge man, arch;
reflexed sheath blade, for example-is apt to perpetrate ambiguity, having developed from buds that arise distichously on the penphery
and so cause confusion. This is because the orientation of such a strue- of a dllated primordium that remains adnate to the surface of the
tUfe may vary-in time, and from one botanical entity to another- culm-as in all known spccies of the genus _A.1f!TQslachys, and some
especia1Iy among the ba:rp.boos. Therefore, neither upper and lower, spede, of ArWQs!Jlidium. See pp. 53ft and Figs. 31 and 32. Cf. Con-
nor dorsal and ventral, w.,l1 always be correlated reliably with ana- ,tellate' Gt-emial: Restrieted.
tomical don;ivcntrality, to which important structural basis all pert;- Assimilation tisS1Jß: Mesophyll;-the imerior ground ti,sue of a leaf blade.
nent observations must be oriented unmistakably in order to satisfy Awned. Bearing a bristlelike apical appendage called an awn (OE.): The
the requirements of scientific discipline, dependability, and c1earthink- empty glumes of the spike1et in the type and some. other specl~ of
ing. Cf. AiW;iaL Neurolepis are awned. In Streptoehaeltl, the lemma tcnnmates above ~n a
s
Abscissile (L. abscissilis,. fr. abSCIssus, cut off). Susceptible of bcing cut off very long, apically eoiled structure calle~ an a-:n (L. Mista) by Ne: ,
or disarticulated by the fonnation of an abscission la'yer;-applies to the author of the genus; Trinius called It a tall (L. ,""da). Thc dls-
the petiole of a deciduous Icaf or, as in many bamboos, to the seg- tinetions between the ways in which the different types of sheathmg
ments of a rachilla. See 1)eciduous, strucrures or their foliar appendages terminate apically are not sharp.
Acropeta!. (Gr. wem, the highest point; L. peto, I go toward). Progressing Technical terms (such as apiculate, awncd, euspidate? ~d ~ucronat~)
toward an apex, as when the deve10pment or emergence of append- and their conventional definitions do not clcarly dlstlngulsh the .dI-
ages on any axis takes place scrially in the dlrection of the apex of verse expressions that are apart of the morpho!ogical inter~datlon
thc axis. Cf. Basipetal. . that baffies efforn at hard-and-fast categorizauon here , as J,ll rrumy
Acropetally (adv.). Toward an apex. other faeets of plant structure. . . ' .
Adaxial. To,,-ard an a.xis;-designates that surface of an appendage which Bamboo (bamb", a ve.rnaeular word of undeternuned Onental ongm). The
faces, in a structural sense, toward the axis on which it is inserted, or wood of bamboo culms; a plant so c1assified.
reHes for support. Cf. A baxial. Bamboos. A taxonomie group of plants comprisin~ the tn.b~ BambUlleae
AWentitious (L. adventiciu.s). ·Occurring in a Iocation other than the usual of the Bambusoideae, a subfamily of the Gramlncae; hVlng plants, or
one. In the bamboos, and in other grasses as weil, the principal com- eulms (sterns) severed from plants of this group ..
plement of roots is adventitious, arising at the nodes of culms and Basipeta! (L. basis, base; peto, I go toward). Progressmg toward a base, as
rhizomes, and not from the primordial root. In the genus Chu.squea when the development or emergence of appendages o~ any axlS takes
(apparently uniquely among the bamboos) adve"titious branch buds place serially in the direetion of the base of the axlS. Cf. Acropetal.
 298 GLOSSARY GLOSSARY 299

Basijmally (adv.). Toward a base. the seed is free from thc pericarp at maturity. See Figs. 55 and 62.
Blade (AS. bl<2tf, leaf). "The lirnb or expanded portIon of a leaf" (Jack- Centrifogally (L. c",trum, center; lugo, I flee from). Outward from a center.
SOll!.In the ~amboos, a thin, expanded, chlorophyll-bearing, sessile or Cmtripetally (L. tmtrum, center; petG, I go toward). Toward a center.
. penolate, apIcal appendage (lamina) of a sheath proper. See Leaj Chartaceeus (L. charta, paper; -aceus, made of, 01' belonging to). Paperlike;
TBottlebnish (bottle. brum, a brush of cylindrical shape, with the bristles papery.
umfonnlY,dlstnbuted on the axis and oriented approximately at right t Circumaxial (L. circum) around; axis). Completely around an axis;-de-
angles to It). A borrowed tenn, gIven a slightly modified form. and scribes the TYpical reach of the locus of insertion of a sheathing ap-
used here (in the adjectival sense apparently not recorded in di~tion- . pendage, 01' an assemblage of adventitious reots. Jackson}s definition,
aries) to describe the approxIrnate pattern of distribution and oricn- under Circumaxile) is limited to the special sense, '"sulTounding a
tation of r:he proliferations of the stigmatic surface in certain bamboo~ central axis which separates when the fruit splits open."
a pattern 10ose1y called plumose or aspergilliform by some authors: CirCUlflml,tatwn (L~ circum, around; nuta:ns, wavering). The circular movement
Cf. Dmdroid. that commonly takes place in the distal portion of a growing axi8,
Bract CL. bracten, a thin plate of metall. A sheathing organ, bladeless or causing the apex to follow a spiral path as elongation proceeds.
larmmferous, found within cr immediately below a bamboo infiores- Congruent ,(L. congruens) confonnable). In agreement; consistent; harmoni-
cence. A bract may be empty or it may subtend a braneh bud or a ous.
branch. In ban:boo inflorescenees of determinate branching, bram Conrtate (L. cmmal:us, born at the same time). (~United, congenitally 01' sub-
are usually rudlmcntary or lacking. Prophylla are by some authors sequently" Oackson), in reference tO the union of members of a set
referred to as braets, or braeteoIcs. of homologous parts (such as the filaments in astamen comp1ement)
f Branch compiement. The array of branehes mat develop at a sin~le culm that takes place in some bamboos. See Mvnadelphvus; Diadelphous; Tri-
node, includi~g the p:rimordial one and any that arise by proliferation adelphous.
from buds at Its proxImal nodes. See Figs. 26, 4, 27, 29,30, and 32; see Omnectwe (L. connectivum). A very narrow strip of tissue upon the approxi"
also MonocladeJ' Pleioclade. mately opposite sides of which the two parts of an anther (that is, the
t Branch sheaths. The sheathing organs borne sIngIy at each node of an two pairs of Iocules) are inserted IongitudinalIy. Jackson describes the
aerial vegetative branch of any order (ex:cept the culm itself), excluding connective as t<distinet from the :fila~ent/' of which it appears to be
the neck sheaths and the leaf sheaths. an apical extension.
f Bud (ME. Imdde). "The naseent state of a flower or branch" Uackson). In 'Constellate (L. constellatio, a cluster or group of fixed stars; a conste11ation).
the usage here .adopted, the tenn bud Is applied only to those pri- Arranged in a constellationlike c1uster.lv, defined here (in an adjectivaI
mordlai vegetatrve or reproductive branches that (1) are enclosed in a sense not found in dictionaries)~ constellate characterizes the pattern of
prophyllum, and (2) have a resting stage (Figs. 28 and 42). Those insertion of branch buds (or branches) where more than one primary
(such ~ bran~h primQrdia in determinate inflorescences generally; and bud emerges Independently at a midculm node-a condition thus far
root pnmordIa, for example) that lack either one or both of these found among the bamboos only in the genus Ch'/1.SqlJ.ea. Cf. Apsidatc)'
features are referred to simply as primordia. Branch primordia in the Grcmz·al; Restricted.
determinate inflorescences of G/aJ::iophyten, Grcslania, and some other Corniform (L. comu, horn). Shaped Hke a horn. See Fig. 26, 12.
genera are prophyllate, but they apparently have no resting stage. Culrn (L. culrn'/1.S, stalk, stern). A segmented aerial axis that emerges from a
Caespztose (L. caesp1.t(J~.. fr. caespes a rod), "Growing in tufts, like grass"
J
rhizome, and fot:n1s apart of a gramineous plant;-the term is used
Uackson);.-descnbes the normal elump habit ofbamboos with pachy- most commonly with special referenee to bamboos (Fig. 1). Syn.: halm,
morph rhlZom~ (FI?, 16, 1), except where the rhizome neck is very haulm haum.
l

much e1ongated, as m Melocanna baccifcra. Cf. Diffuse. t Cu/rn node (L. nodus, knot). "The 'knot' of a grass stern" Uackson). The
fCaryopsis (Gr. karyon, a nut;opsis, likeness). "A one-celled one-seeded ter:m node is generaHy applied in a loose, comprehensive sense (sensu
sup:nor fruit, v.~th pencarp united to the seed" Gackson: under Cari~ latv) to that comple:< Iocus (see text, p. 12)-the junction of adjacent
~pslS). A caryopsis i8 indehiscent) that is~ its pericarp does not open to intemodes in a segmented axis of a gramineous plant. Hacke! (Lamson~
IIberate the seed. In some bamboos (Me/oeaTma baccifera, for example) Scribner and Southworth 1890:2, 3 and Fig. 1) stressed the impor-

--------~~~--_ ..... _~ .... _-----

 300 GLOSSARY GLOSSARY 301

tance cf thc "constantly overlooked" differencc betvveen the culm nodes t Determinate (L. determinatus~ having fixed limits). Of limited gro'Wth;-
and the sheath nodes in nonbambusoid grasses. However, neither eharacterizes an axis, or a system of immediately related axes (a
Hackel's description nor his illustration does justice to the distinctivc determinate inflorescence, for example), whose development or po-
features of these structures as they appear in the bamboos. Culm node tential for development is confined to a single, definitely limited
is here defined with special rcference to that level within the node "grand period of growth," after whieh no meristem persists, as a rule.
(sens" lato) where secondary elongation (intercalary growth) takes The occasional persistence of meristem at the tip of individual rachillas
place, and where branch buds and adventitious roots are inserted. sometimes prolongs the grand period of growth in a detc::nninate in-
Each culm node (sensu stricto) is located just above a sheath node floreseence atypieally. A raehilla may then eontinue to produce new
(q.v.), from which it is usually distinguishable by the transverse thick- florets until the spikelet is greatly elongated-to as mueh as a foot,
ening cr ridgc (the "supranodal ridge" cf seme authors) that appears while its nonnal length would be 3 to 4 inches. Again, this meristem
at the level of insertion of a bud or a branch complement. Branch may revert to the vegetative state, prolifieating to produce a leafy
node and rhizome node should be similarly differentiated, respectively, axis. Both of these teratie phenomena have been observed in Arundi-
from the eorresponding sheath nodes. Cf. Sheath node. naria dolichantha (see p. 103). See Proliferation (where prolification is
Culm sheath. One of the sheathing organs borne singly at eaeh node of defined), and Semelauctant; cf. Indeterminate.
the eulm proper, below the level at whieh the sheaths of foliage leaves Diadelphous (Gr. dis, twiee; adelphos, brother). Having "two groups of
take their place. See Figs. 36 and 37. Kurz (1876:268) anticipates stamens";-eognate with Diadelphia, the name of "a Linnean elass
this usage with "halm-sheaths." having the stamens in two bundles of brotherhoods" Gaekson). Di-
Culm shoot. Bamboo shoot; a young culm in any stage of its development adelphous deseribes the eondition of astamen complement of a flower
short of maturity in height. See Shoot. when its members have their filaments connate to form two distinct
Cuspidate (L. ,cuspis~ a .point). Terminating apically in "a sharp, rigid groups. In the bamboos, the relative numbers of stamens in the re-
point" Gackson) in placc of ablade (as in some sheathing organs). spective groups may vary from one genus to another. Cf. Monadelphous;
Cf. Mucronate. Triadelphous.
Deciduous (L. deciduus). Falling off; "falling in season" Gackson). Cf. Diageotropic. Adjectival fonn derived from diageotropism, whieh is defined
Persz·stent. under Geotropism (q.v.).
Declined (L. declinatus~ turned aside). ~'direeted obliquely" Gackson);- Diaphragm (Gr., dz'aphragma, a partition wall). The transverse internal
deseribes the habit or orientation of the eulms in some species of bam- layer of parenchyma found at the level of every sheath node. It is
boos. reinforced by the erossing over and anastomosis of vaseular bundles,
Deliquescent (L. delz'quescens, melting away, disappearing). Beeoming dis- and fonns a rigid structural element that lends strength (meehanical
persed or expended, as when the main axis of any strueture (the rachis resilience) to the segmented vegetative axes' of bamboos. "A dividing
of an inflorescenee of Aulonemia quexo~ for example) loses its identity membrane or partition" Gackson).
among its own branehes, or "loses itself by repeated branehings" Diffuse (L. diJfusus, spread out). Growing in open array;-characterizes the
Uackson). Cf. EXcuTTent. normal mature elump habit typical of most bamboos with lepto-
Dendroid (Gr. dendron, tree; -oid, having the form of). Treelike; bearing morph rhizomes, and those whose paehymorph rhizomes have a
branehes of more than one order;-deseribes the pattern of proliferation greatly elongated neck, as in Melocanna baccifera, for example. This
of the stigmatie surfaee in eertain bamboos. clump habit was refcrred to earlier (McClure 1945b:278) as dumetose.
Depauperate (L. depauperatus, impoverished). Redueed in size or functional Cf. Caespitose.
effieiency or both;-charaeterizes pseudospikeIcts, florets, individual Distal (L. dislo, I stand apart). Remote; far out;-designates loci of in-
organs, or even whole plants whose development is impeded or re- sertion, or struetures, situated at or near the tip of an a",is. Cf. Proximal.
strieted by adverse eonditions, external or internal. Distichous (L. dz'stichus, eonsisting of two rows). "Disposed in two vertical
Dermogramme (Gr. derma) skin; gramma~ something drawn). A French term ranks, as the florets in many grasses'" Gackson). Distiehous also de-
used by Prat (1936:178) to designate any example of his diagrammatic scribes the pattern of insertion of buds and branches, and the sheathing
illustrations of the cellular details of the strueture of the epidermis of appendages on all segmented axes, both vegetative and reproduetive,
any grass or bamboo. See Spodogram. of the bamboo plant. As taeitly reeognized by agrostologists, but
 302 GLOSSARY GLOSSARY 303

generally not mentioned in published definitions, thc two ranks are Fistulos< (L. fistula, a pipe). Hollow; having a lumen;-characterizes thc
normally insertcd on opposite sides of an a.'Xis. internodes of eulms and branehes in most bamboos.
Di,tich} ,Ca substantive apparently not yet found in any dictionary). t Floret (dim. of OF. j1or, 1I0wer). One of the units into which aspikelet
Arber ~ word for thc two-ranked arrangement of buds, brauches, and breaks up when the rachilla segments disarticulate. Regardless of
sheathmg appendages generally characteristic of segmented a.xcs in whether the rachiUa segments are abscissile or not, a floret consists cf
the Gramineae. See Dist'l'dwus. (I) a segment of the rachilla, (2) the lemma that is insertcd upon it,
Embryotcgium (Or. embryonJ a foetus; tega; a covering), Literally, embryo (3) a branch (the axis of a flower) subtendcd by the lemma, (4) a
~over;-the "little shield" (Sp., escudete) of Parodi (1961). "A callosity prophyllum (the palea) of the axis of the flower, and (5) the parts of
In the seed coat of some seeds near the hilum, and detached by the the flower that are inserted on its axlS. Attention is directed espedally
protrusion of thc radicle on gcrminationn Gacksen). Slnce Jackson~s to the fact that the gramineous floret includes structureS from axes of
definition is based on the nature of thc embryotegium in deciduous tVVQ orders, while the flower i5 confined to a single axis. The distinction
~eeds) it is not ~trictly applicable to its manifestation in a typical gram- between floret and flower i8 not always observed in published works
meous caryopSls. Haekel (Lamson-Scribner and Southworth 1890:20, on agrostology. The loose use of the two terms interchangeably is to
Fig. 6) describes and iIlustrates it (without, however, using a teehnical be avoidcd. Cf. Fwwer.
name) as "a pl~ce where the embryo lies covercd only by the pericarp t Flower (OF./lor). That portion of a branch of the rachilla that is distal to
and plamly vunble on the outside." The position of the embryotegium its own prophyllum (the palea)} together with the reproductive organs
(as describcd by Hacke!) is clearly visible in carvopses with a thin (androecium or gynoecium or both) borne by it (Fig. 53, B, C). The
periearp; as in the eommon cereal grains) and in bamboos of cert:ain Jodicules, when present, are ineluded in this concept of the flower, but
gener~ such as Aru:ndinaria and Bambusa, In other genera, including the palea is not. Cf. Floret.
MeloclZtlfUl (Stapf 1904:Figs. 3 and 40) and Ochlandra, where the pcri- Flowtri.ng branch. A leafy or leafiess segmented axis that bears one or more
earp is thickened, the embryotegium is ordinarily not externallv rec- inflorescences. A flowering branch 15 distinguishable from an inffores-
ognizable. See Fig. 55. . cence proper in that it retains all of the morphological characteristics
Empty glumes. Sheathing structures, normally present in every bamboo of the vegetative state, with the sole exception that in some cases it
spikelet (~sually 2, sometimes 1 or 3, exceptionally more; rarely ladung does not produce foliage leaves, The inftorescence proper:> on the other
enurely) ffisertcd on the rachilla, immediately below the lowest lemma. hand, always has morphological features peeuliar to it that are not
Empty glumes do not subtend either branch buds or flowers. In some found on the flowcring branch proper. In describing and illustrating
bamboos, the distinction between empty glumes and sterile lemmas bamboos, some authors fail to distinguish the !wo. Arber (1934: 108)
is not clear. See Glumt;· cf. Lemma. characterizes a leafless flowering bamboo as a Htrul y gargantuan in-
Endasperm (Gr. endo, within; spermo., seed). "The nutritive tissue formcd florescence.)~ Cf. lnjloresccnce.

within the embryo sac" of a seed (Webster)-referred to by earlier Foliar organ. See Sheathing organ.
writers as "albumen') Gacksen). In some bamboos, as in Melocarma Fugaceous (L.fogax, fieeting). Promptly deciduous; "soon perishing" (jack-
baccflera> the embryo consumeS the endosperm as fast as it is formed son).
(Stapf 1904). Fusiform (L.ftsus, spindIe). Spindle-shapcd; eircular in cross section, thick-
Exe.menl (L. excu.rrtnS" running out). Extending through, as when the main est in the middle and tapering toward cach end. See Subfosiform.
axis of any structure (the rachis Qf the paniculate infiorescence in Ge:nieulaU (L. geniculatus, with bended knee). "Bent abruptly at an angle,
Indocalamus sinicus> for example) maintains its identity among its own like the bent knee" (Wcbster). When a growing culm adjusts the
branches; '.'where the stern remains central, the other parts being direction of its orientarion (as in situations where negative geotropism
regularly dlSposed round it" Gackson). Cf. Deliquesemt. operates to effect the restoration of a fallen or deflected culm to an
t Fibrous (L.Jibra, thread, filament). :'-Iumerou" ,Iender, and not conspicu- upright posture), the adjustment takes place through differential
ously tapered;~the meaning of the term as conventionally used to elongation in the upper and lower portions, respectivcly, of the zone
describe the gross aspect of the roots of bamboos and other grasses: a cf inte:rcalary growth immediately above anode, and the node in-
definition a pparetltI y not recorded in dietionaries or glossaries, Jackson~s volved becomes geniculate.
definition of the tenn is not applicable here. tGeQt:ropism (Gr. ge, the earth; plus tropism, defincd by Webster as "the
 304 GLOSSARY GLOSSARY 305

innate tendency o'f an organism to react in adefinite manner to ex~ of its early stages are needed 10 clarify the fundamental details of
~emal sti:nuli"). A physiologicaI potential, and the eonsequent aCt) the mature stage of this peculiar structure, but the distinct tuft of
mvolved In the assumption and maintenance of a partieular direction branches appears to be independent of those arising distichously from
of growth by an axis or organ with reference to the force of o-ravity the divergent: elearly segmented, proximal portion of the primary
or to the centrifugal force genera ted by rapid circular motion. A,~ branch proper. Cf. Apsi(/al.e; Grms/ellate; Restnetcd. . . . .
commonly used, geotropism ambiguously embraces (and confuses) two (~moecium (Gr. gyne, female; aikos, house). The complete pLStII, conslstmg
component phenomena: the visible physical response, and thc mOre ofthe ovary (the ovulary of Schaffner 1934), the ovulc(s),.style(s), and
fund~~enu;;~ invisible respo~se, which i5 physiological. Jackson~s the stigma(s), of" single flower (Fig. 53). I have not followed Arber
definitlon: thc force of gravlty as shovm by curvature in nascent (1934: 120 cl passim) in adopting the spclling (gynaeceum) advocated by
organs of plants" is incomplete, Three distinct classes of geotropic Kraus (1903). Jackson and Nybakken both give gynoecium. Cf.
responses are recognized: positive. negative, and transverse. The last- A ndroecium.
named is called diageotropism. Halm haulrn haum (terms used in British agrostology). See Culm.
t Girdle (AS. gyrt.k~ belt). A conspicuous horizontal band of tissue inserted Herm~phrodi;t, Having both androecium and gynoeci~m;-designates a
circumaxiaUy at the sheath nodes of some bamboos (Melocanna com- flower cr fiowers, so equipped: commonly symbohzed by Q or y.
P~ctiflrn:" and:ome species of Ghusquea, for examp!e; see Fig. 38). The CL.
Imbricate imbricatus, covered with tiles). "Qverlapping like roofing tiIes
gxrdle 15 a vlslble and anatomieally distinct basal part of each sheath and ,hingle," (Brown 1954). Sheathing organs at succes-slve nodes of
proper on any vegetative axis of tbc pJant wherein 1t occurs' it is an axis are imbricate when each one exceeds, in length, the intemode
particularly co.nspicuous at the nodes of the culm. Ai; long ,:" thc it embraces.
correspondmg mterealary zone of the embraced internode is 1n active t lndeterminate (L. indeterminatus) unlimited). "Not t~inat:d .a~solutely:}
gro;nh} a girdle is e~pansile in alI of the dimensions that correspond Gackson). Continuing apical growth characterures an mdlVldual ax:lS
to l'ts surface? and 1t takes on an asymmetrieal form at geniculate as indetenninate. The term is here given another connotation (name1y,
nodes. L pon the abscission of the sheath, the girdle remains attached iterauctant' with particular reference to the continuing rammcation
at thc sheath node. Thc numerous definitions of girdle drawn by of a systen: of immediately reiated "xes, as e.xemplified by the inde-
J~ckson from botanicaI literature cOVer a wide diversity of appIieations terminate inflorescences of certain bamboos. In such inflorescences) the
gIven the term, but no Olle of thern corresponds to thc one added buds at thc proximal node, of each new order of branches bear fresh
here. Cf. Snealh cal/us. bodies of meristem. The deveIopment of new branches from these
Gtwr.. (L. gtwna, a hull or husk). "The chaffy, two-ranked members of buds may be continuous (or interniittent and reactivated on ~ seasonal
the inflorescence of grasses and similar plants" Gackson). See Emp!)i basis) for aperiod lasting in some cases for several su.cce~lve. }·ears.
glumes. Each branch of an indetenninate inflorescence (an aXlS v,rlth ItS ap-
Gregarious CL, gregatiUS of or pertaining to a Bock or herd). aGrowing in
J pendages) has the appearance of aspikelet. Because of this, it is called
com pany" Gackson). Gamble (1896:viii) makes gregarious mean simul- a pseudospikelet. Each p,eudospikelet has its own independent grand
taneous, to describe the Bowering behavior manifested when alI cf the period of growth. The active period of the me:",:",,: at the apex .of
members of a given generation of bamboo plants (from seeds of a the rachilla that terminates each pseudospikelet :tS Imllted. Thus, while
common origin) enter the reproductive state at approximately the the branchinO' ofindeterminate bamboo infiorescences is of a continuing
same time. Cf. Sporadic. nature) each '='individual branch 1S apically detenninate in its gro~1:h.
t Gremial CL. grerm·al~ of a rap 01' bosom). "Growing in a poIIard-like clustern See lteraucta1'.t; Pseudospikelet; cf. Determinatc.
(Brown 1954). The botanical application ofthe term is extended here lriflorescenct (XL. inflorescentia, fr. L. iTfflorescens, f1ower~g). A dis~ete
to c~aracterize thc insertion of a pleiocladc braneh complement in ~
arrgrezation of spikelets associated .....ith a common pnmary rachls .or
~

certam bamboos (Guadua spinosa, for example) where a tuft of ,evera! a common peduncle. E.xceptionally, an inflorescence may compnse
slender, sub~qual branches arises from a basal extension of the primary but a single spikelet. The inflorescence i.n the bamboos of n;ost genera
branch. ThlS apparently unsegmentcd bodv of ramiferous tissue re~ is usually either strictly determinate or mdeternunate as t? lts b:rar:ch-
maius low and adnate to the surface of the culm. Anatomical studies
I habit. The occurrence of individual features of an mtennedmte
 306 GLOSSARY GLOSSARY 307

nature is rare, Cf. Flowm:ng braru:h; see also Dctenninate)· Inrktemdnatc~. ligule/' especially when it is conspicuously devc1oped, as it i3 in some
Pseudospikelet. variants of Arundinaria tecta, for example. The term '<inner ligule" is
Insertion CL. Vtscrti(l; a putting in). <'Tbe mode or place of attachment of then applied to the adaxial structure ronventionally known simply as
an ?rga.n or a.part: as the parts of a flower" Gackson). A locus of in~ the Iigule (see Fig. 35). The ligule is reduced to a mere line, or is even
scrtlOn 15 apomt, lIne, cr area that is coo.:tensive with the connection lacking cntirely, in the sheaths of certain species of barnboos.
be~vee? a. give? axis Or organ and any appendage cr structure to Locus (L. place). The locale in which some event takes place~ or wh.ere
WhlCh It glves nse. See Locu.s. something is to be found. Locus is here given the connotatJOn, pomt,
>I< lterauctant CL. itero, I rc?eat; auctans;> increasing, growing). Ernbracing more .
or line , or area , principaUv in the phrase "locus of in"sertion." See
~an on~ grand ~lod of growth. The branching system, cr thc branch- Insertion.
~ng habIt, of an mdeterminate bamboo infiorescence may be said to be Lodicule (L. lodicula, a small coverlet). One of thc small, usually thin,
lterauctant. Cf. Semelauetant; see Indeterminate; Pseudospikelet. delicate and transparent structures (hy some authors ref~ to as
t LeaJ (AS.). Foliage leaf; bamboo leaf; the chlorophyll-bearlng, usually ('scales") inserted usual1y in a single whorl of 3, immediately helo,,: the
petio]ate, blade of .a leaf shcath proper. 'Ibis concept of leaf, in the stamens in the bamhoo ftower (Fig. 53). The lodicules are relatIvely
sense bamboo leaf, mcludes thc petiole when it is present, but exeludes large, opaque, and parchmentlike in the ftowers of known species of
the leaf sheath proper. Leaf IS here defined in trus circumscribed St:reptochaeta.. Their number is variable in seme bam~oos (up to l~ or
sens,e in. or?er ,tO .avoid a commonly encountered ambiguity occasioned more in Ochlandra stridula); in known species of Gtganti)Cht~ typlcal
by ltS Indlscnnunate ure in the literature. The word is cornmonlv 10dicuJes are lacking entirely.
used by agrostologists) without modification or qualificatiofi~ to ref~, Lumen CL. opening). "The space which is bounded by the walls of ~n
to any one of the morphologically, anatomically, and functionally organ; as the ccntral cavity of a ceU" Qacks?n); the central cavrty
dIvergent forms of sheathing organs borne on the reveral kinds of of a hollow intern ode of any segmented axlS of a bamboo plant
~eg~tatjve axes of ~ bamboo plant. This imprecise use of the term) (Fig. 20). • .. ...
In dlsregard of the Importance of the manifest differences thus looselv ]v[eristem (Gr. meristos, divisible). A body cf tlssue In Whlch cell rnVl,slon
eovered by it, apparently is due to the generalized and unqualHied and differentiation are active· or potential.
nature of the definition of "leaf" given by Jackson: Uthe principal • MetamtJrph (Gr. meta, implying change; I1VJrpM, form). Of intermediate
appendage or lateral organ borne by the stem or axis." Cf. Skeat/:; fonn;-a term proposed here to designatecertain underground portions
leaJ dual/:; see also Sluathing organ. of segmentcd axes, whose transitional character cannot be clearly
LeaJ snealk One of the leaf-bearing sheaths inserted at the distal nodes of indieated by the use of existing conventional tenns. See .Metamorph I;
ea~h aerial vege:ative axis of a bamboo, whcther eulm~ brauch, or Metamorph IL
twlg. Tbc conspl(;uous part of a leaf sheath is the usuallv petiolate * Metamorph L A transitional a,."ds that occupies a position bet\veen the culm
blade inserted at the apex of the leaf sheath proper. Cf. i.afl Sluath. neck and the base ofthe culm proper, where no clearly defined rhizome
Lemma (Gr. a husk). The glume that subtends a flower. A lemma that intcrvenes;-in same Gases douhtfully distinct from an elongated culm
~ubtends a functional (or fully formed) flower is called afertile lemma neck (Fig. 14).
~n ~rder to distinguish it from a sterile temtlU1;, whose subtcnded fiower * Metamorph 11. A transitional axis intermediate in form and position bet-ween
• I, elther depauperate, rudimentary, or lacklng. Cf. Empty glumes. the apex of a rhizome (either pachymorph or leptomorph) and the
Lepto"'.orph (Gr. leptas, thm; morphe, form). A term coined especially to culm into which the rhizome 18 transformcd apically. It appears where
deSlgnate a. slender, e10ngate type of rhi7.ome proper described on the transforruation of the apex of a rhizome into a culm takes place
p. 25 (see Flg. 3). Cf. Pat;hyrTVJrph. gradually and not abrnptJy (Hgs. 12 and 16,2). . .
Li,,"Ulate. Having a ligule. *' . lvfelamorphological (Gr. meta, beyond) plus morphologu;af), Pe:w:m~ng to the
Ligule CL. ligula, a little tongue). A thin, apical extension of a sheath techniques, the afeas of investigation) or the data of dlsclplmes other
proper, adaxial to the locus of insertion of thc sheath blade (or Icaf than morphology.
petiole): The outer drn of the little cup revealed by the absd;sion of Monade/phou.s (Gr. monos, one; adelphos, brother). Uriited in a single brother-
the pctlOle of a fohage leaf 15 somctimes referred to as the Houter hood;-an adjective cognate with Monadelphia, the name of !(a
 308 GLOSSARY
GLOSSARY 309
Linnean class in which the anthers are unlted by thcir filaments into
a single brotherhood" (Jackson), and conventionally used to describe *Neck sheath. One of the reduced, bladeless sheaths that dothe the con-
the stamen complemcnt of a fiower when aII of its mernbers are united strieted proximal part) the neck, of various vegetative a.-res in the bam-
by their connare filaments, Cf. Diadelpiwus; Triadelphous, 000 plant. Neck sheaths are sometimes referred to as cataphylL, (Gr,
}vf.onocarplc. Sec Monoperiodic. kata, do,,'n; phyllo., leaf).
*' MCl1'loclade (G.r, memos" single; klados:; brauch), Comprising hut a single Obsolete CL. obsoletus.} worn out}. "Wanting er ~di~;nta~; usecl cf an
branch (FIg, 29, A);-a term proposcd here to designate branch com. organ which is scarcely apparent or has vanrshed (Jacl:son),
plements so constltuted. Thc monodade state is facultative when Obterete. Circular in cross seetion, tapering progresSlvely frol? one ~nd to
?Uds arc present at the proximal nodes of the primordial brauch; it the otherl and smallest at the proximal end (for example, a rh1zome
15 obligate whcn buds are lacldng at this level. The monodade state neck), Cf. Tm/c. . .
cf bra.n~ complen:cn~ is obligate in most known speeies of the genus • Pacltymorph (Ge. pachys, thick; morphe, form). A tenn <>:med especrally :0
Sas~; 1t 15 facultatlve In some species of the genus Amndinaria. Cf. designate a short, thick type of rhIzome proper (Flg, 2) descnbed m
Pletocladt;" see, also Branch complem.ent, detail on p. 24, Cf. Leptomorpk . ,
*..:Vfon:periodz'c (Or, m.oMS; one; ~eriodos, a completcd course), Having but a Pa/ca (L., [a piece of] chaff), The prophyllum of the a;(lS d a grammeous
sIngle reproductlve cyde wuhin the lifetime of a plant;-characterizes fiower. Jackson's definition: ('the inner bract or glume 1n grasses, called
?amboos (SUC? as Bambusa arundinacea) that fiower but onee, then per- 'Palee bv North Arnerican v..'Titers u is too vague to bc useful. See Pro~
l~. Monoeyche suggests itself for use in the sense hcre given the newiy phyllum, ,
comcd t~rm, monop~odie, but it aJready bears the disquaIifying t Panicle (L. panieula, a tuft), A determinate infloresc,,:,ce ;"Ith branc~es of
connotatlon annual, wlth reference to the life span of a plant-besides more than one order. Jackson}g definition of pan1cle LS not apphcable
indicating sets cf parts (such a, scpals and petals) that comprise but a here.
single whorl or ey::le..In the interest of clarity, thc term monoearpic, Pa.ieulate (L, paniculatus, tufted), Having some or all of the characters of
commonly uscd to lndicatc the monoperiodic eharaeter of certain bam- a paniclc (q,v,), . , . ,
boos, should Ix: reseI"V'cd (with its orthographie variants; see Jaekson) Parenclryma (NL' l fr. Gr. parenchei~ to pour In beslde). F;mdamental tl~ue,
for rc~erencc to a pistil comprising but a single carpc1. Cf. Pofyperiodic. ground tissue; a tissue, such as pith, composed of thm-walled) undlffer-
Monopodzat (Gr. monos~ one; poas, podo~ foot). Having thc form of a mono- entiated, isodiametric cells (adapted, from Jackson)., .
podium, which is defined by Jachon as ~(a stern of a single, and con- t Pedicel (L, pediccllus, dirn. of pes, foot)", The stalk of aspikelet, that lS, the
tinuous axis, ,) This tcnn was used carrier (McCIure 1925) to designate distal segment or internocle of the a.xis immediately below the glum:s
the type of rhizome described as leptomorph (q,vJ CE SympodiaL that mark the base of aspikelet. The ultirnate branches of a determl-
t MorphogeneslS (Gr. morphe~ form; gentsi~ beginning or origin). (The nate infiorescence are a11 pedicels, since each tenninates in aspikelet.
prodUction o~ morphologicaI eharacters" Gackson). As dcfined here In spicate racemes, and in pseudospikeIets, the pedicels are usually
morphogenesls eonnotes not onIy its externaI manifestation..;;~ but also,l very short, Cf. Peduncle; Rachis.
an~ prim~rily.~ the natur: an~ sequence of events in the prograrn of Pedicellate. Having a pedical. .
gemc monltonng of phYS10loglcal states associatcd ""ith the emergence t PedWlcle (L. pedunculus, dirn, of pes, foot), Thc stalk of a:' Inflores,cence,
of the structural features that determine the charactcristic form of a that is~ the unbranched segment of the infiorescence axlS that 15 unme-
plant. diatel; below the rachi..;;. In determinate inflorescences, the first sheath~
~"4u.cronate CL. mucronatus., pointed), apossessing a short, straight point, as beari~g node below thc first branch of the inflorescenee is here taken
same leaves)) Gackson). Cf. Cuspidate~' }r1uticous. arbitrarHy as marking the base of the peduncle l and. the locus of
;:\1utic01l.$ (L. rnzc. ticus) ?ocked). ~:Blunt, a:"vnless" Oack.son);-used primarHy insertion of the first branch of the inflorescence as marking the apex.
(when apP!lcablel to descnbe thc apex in ,mall sheathing appendages In those rare cases where a determinate inflorescence includ~s only
on the vanous axes cf the bamboo infloreseence. one spikelet, the peduncle is not distingnishable from the podlcel. In
t Neck (AS. Imecca). The constrieted basal part, charaetcristic of all, or most, indetcrminately branching inflorescences, the peduncle JS ~suall~ ver)'
cf the segmented vegetative axes of a bamOOo plant. See pp, 12f and short. An exception appears where the primary pseudosplkelet lS te:-
Figs, 2 and 3, minal (instead of lateral) to a flowering bra~ch. Here the poduncle !S
more e10ngate (see Fig. 45). CL Pedre<!; Rach!S.
 310 GLOSSARY
GLOSSARY 311
PedunC'<.date. Having a peduncle.
unit} as by branching or interpolation of members" Oackson);-a gen-
Pergamineous (L. pergam~ parchment). "Like parchment in te>...'1:ure" eral term for the particular examples described under ProliferatiCIII and
(Jackson).
Pleioclade, as pertaining to vegetative axes, and under Indeterminate, as
Persistent (L. pcrsz'stensj> remaining in place). Not deeiduous;-applies to pertaining to reproductive axes.
organs that rernain in place after they have fulfilled their natural PlUate (L. p!icatus, folded). "Folded into plaits, usually lengthwise" (Jack-
functions (for example, the culm sheaths of Arundinan:a tecta). ..on); marked by longitudinal ridges suggesting a prior condition of
t Phototropism (Gr. photos, light; tropos, a turning). "A tropism in which being folded like a collapsible Chinese fan;-an appearancc shown by
light 15 the orienting stimulus, as in the tuming toward a light of a some bamboo leaf blades.
plant shoot or a tube worm, and in the creeping away [rom a light of Polycarpic. See Polyperiodie.
a blowBy larva') (Webster). '"'The act of turning towards thc sun or a * PO{yperz'ocfit (Gr. polys, many; periodos, a complete course). Having many
a
SOurce of light" ackson, under Heliotropism, of which he gives Photo~ reproductive periods} alternating with vegetative periods, vvithin the
tropism as a synonym). As understood and defined here, the term lifetime of one plant;-characterizes bamboos (such as plants of the
phototropism encompasses a complex of phenomena (visible and known species of PhyllostachJs) that flower repeatedly (and usually
invisible) involved in certain biotic responses to inddent light. Whe:n periodically) during an indefinite life span. Polycyclic suggests it,elf
the corresponding physiologlcal potential is prellent in the appropriate for use in the sense here given polyperiodie, but is rejected because of
living tissue of an axis~ an organ: or an organism, it induces a move- its long-established use with rererenee to the occurrence of an indefinite
ment,. or. the assumption of a particular orientation, with respect to number of whorls of parts) such as: sepals or petals in a flower. Pleio~
the prmcrpal souree of incident light. Two contrasting classes of photo- cyclic is already burdened with the connotation perennial, which is
trOplC responses are recognized: positive and negative. Most published incompatible aS a meaning alternate to that here given polyperiodie.
definitions of this and other tropisms faH to make elear the distinction Moreover, monOCYcIic, which would pair with the pleiocyclie, already
between thc two component phenomena, namely, thc visible physical means annual, in reference to the life span of a plant. In the interest
response, and thc more fundamental, invisible response~ whlch is phys- of clarity) the term polycarpic) sometimes used to indicate the poly-
iologicaL .
periodic character of eertain bamboos, should be reserved (Vv'ith its
t Pirmate (L. PinMtus, feathered). "Featherlike" (Webster): having lateral ortl:J.ographic variants) for reference to ovaries vvith numerous carpels
appenda~es ~istributed i~ two eontinuous series inserted, respectively) (see Jaekson). Cf. MOIWperiodic.
on opposlte sldes of an a.xJ.s and antrorsely oriented;-a borrowed tenn Prirmrrdial (L. primordialis). "First in order of appearance" (Jackson). See
used here to describe the pattern of insertion and orientation of the Primardium.
proIiferations ofthe stigmatic surface in certain bamboos.Jackson and Primrrrdi:um (L., the beginning). An axis or an outgrowth of an axis in
Webster both confine the botanical application of the term pinnate to its earliest recognizable eondition) or in an early donnant statc. In
the arrangement of the leaflets on the rachis of a compound leaf.. the bamboos, a branch primordium borne on a segmented vegetative
Pistil (L. Pistillum, a pesde). See GyMeciUJ'l!. axis is always encIosed in a prophyllum; an adventitious root primor-
P.l Abbreviation of "Plant Introduction," prefixed to a permanent iden- clium is neVer so enelo,ed. See Prophyllum.
tifying number assigned by the U.S. Department of Agriculture to Proliferat;"n (L. poles, off,pring; /ero, I bear). "Bearing progeny as off-
each lot of living plant material (seeds, plants, Or euttings) aere.'lSioned shoots)) Gackson). The term is brought into foeus here with reference
in its r~cord o~ plant introductions. These numbers are published} to the rapid multiplication of members of a branch complement by
aIong "nth pertment names and documenting notes in a connnuincr the prompt awakening of buds at the proximal nodes of the compo-
series of Plant Inventories. , "
neut members. The proliferation of the culm itself by thc same proccss
~ Pleioclade (Gr. pleion) more; klados~ branch). Consisting of more than one (wi,hout the intercalation of a rhizome) is called stooling or tillering.
branch (Fig. 29, B-F);-a term here proposed for designating branch Proliferation should not be confused ,,~th prolification, which Jackson
complements SO eonstituted. Cf. Monoelade; see also Braneh complement' differentiates as "The producrion of tenninal or lateralleaf buds in a
Pleiogeny. '
flower. " An example of prolification is cited under Determinate. See also
Pleiogerr; (Gr. plet'iir., more; genos, birth). uAn increase from the parental Plciogeny.
 312 GLOSSARY
GLOSSARY 313
Prop~lIatß. Bearing a prophyllum (q.v.).
or indirectl y from buds at the proximal nodes of the primordial
Prophyllum (L., first leaf); a.w. proP~li. A sheathing organ, usually 2-keeled branch, whether this: axis is dominant or not. Cf, Apsidaw.: Conslcllate;
and inserted circumaxially at the first (proximal) node of a brauch. In Gremial.
vegetative axes;> and in inflorescences of indeterminate branching, the Rhizome (Gr. rhiz&r.tZ; a mass of roots). An individual component branch
prophyllum at first surrounds the branch primordium to form a bud cf the subterranean system of segmented a..'Ces that constitute the
(Figs. 42 and 43). Jackson's definition of prophyllum limits it to what ~'chassisH (populady referred to as the '"rootstock") of a bamooo plant
is now generally <alle<:! a palea. Prophylla are by 30me authors called (see Fig. 1 and p. 19). A rhizome consists of two parts: ~e rhizome
bracts. or bracteoles. See Pal,a; Sud. proper and the rhizome neck. 1'''''0 disrinct types of rhizome are differ-
Proxim:ll (L. proximus~ nearest). Basal; situated at or near the base of an entiated: leptomorph and pachymorph (qq.v.).
axis or an organ;-designates 100 of insertion, cr structures, so situ- Rhizome shea!'" The husklike sheathing organ inserted at each node of a
ated. Cf. Dis/ai.
rhizome proper (as distinct from the rhizome neck); see Fig. 34.
* Pscudospikelet (L. psev.do~ faIse; by e.xtension, superfically resembling; spicula, Secund (L. secundus). following). Unilateral, as describing (1) the arrange-
spikele~). A. spikeletlike branch of an indeterminate (indeterminately ment) Or pattern of insertion; of appendages on an a.. . .is, and (2) the
branching) mflorescence (see p. 91 and Fig. 50, A, S). See Irufet,rmina!,. form thus given to the resulting ensemble. Thus, the unilateral arrange-
PulV1:nus (L" cushion). A tröpica1 organ CL. triJ..bicus7 pertaining to a tUrn- ment of the spikelets on the rachis in Merostae~s is described by
ing) .associated with movement cr differential growth. Pulvini usuaUy Sprengel (1825, I: 132) as a =und spike (spie. secwula).
ITlanIfest themselves as dome-shaped eminences) commonly in pairs, *Sernelauctant (L. serne4 enee; aucta~ increasing) growing). Embracing hut a
strategically Iocated in relation to the ,organ with whose orientation single grand period of growth. A detenninate bamboo infiorescence
they are concerned-at or near the base of the leaf petiole (Fig. 35), may be said to be semelauctant. See Dettmnir.tltl!; lterauctant.
for example, or a branch of an inflorescence. Pulvini function th!,ough Sensu lato; sensu stricto (L.). In a broad sense; in a restricted sense;-e."<pres-
one or the other of two physiological mechanisms: changes cf turgiditv sions used to indicate the intended scope of a given application of a
and differential growth. '
seientific name or a scientific term.
t Rareme (L., ra"mus, a bunch of grapes). A determinate gramineous inflo- t Sheath (ME. schetke;. A sheathing organ, the basal part of which, the
rescence Vvith a single order of- (usually solitary) branches. ]ackson's sheath proper, completely surrounds the vegetative axis on which it is
definition is not appIicable here. A raceme in which some of the borne, i1$ locus of insertion being circuma.x.ia1. In its simplest form (as
branches emerge in fasciclcs of 2 or 3 mal' be characterized (inter~ in typical neck sheaths and rhizome sheaths) the sheath proper tenni-
preted) as a panicu1ate raceme) if the extra branches in each faseide nates apically in a short, hard point, In its fuHy elaborated form-that
are taken to be secondary ones arisinoet subcutaneouslv, from the ba"'" ~ characteristie or' examples in the middle of any series-the sheath
of a primary one. Athroostac~s capitata bears paniculate racemes in proper usually terroinates above in a Egule, at the base of whi~ i8
\->..hieh all branches are relatively short This gives the inflorescence a
inserted a more or le$S expanded laminar appendage referred to In a
capitate superficial appearance, A spieate raceme is one in which the comprehensive sense as the sheath blade. In addition, the sneath proper
pediceIs are so short that the inHotescence resembles aspike, commonly bears~ at or near each extremity of thc loeus of insertion of
t Rachil/a; a.w. rhachilla (dirn. of rachis, q.v.). The axis of aspikelet in any the blade, a tuft of bristles (oral setae). These may be borne either
gramineous plant. Jackson defines rhachilla vaguely as "a secondary directly on the sheath proper or on the margin of an auride. \\'hen
axis in the inflorescence of grasses. n
there is a pair of amides (one on cach side) the two may be similar
Rachr's)' a.w. rhachis (Gr. rhachis~ backbone). The primary a.,,<is of an infio- and subequal, or they may be unequal in size and dissimilar in shape.
rescence;-its position is terminal to the peduncle. To avoid thc ambiguity occasioned by a commonly encountered indis-
RedW::l!~ Subnormal in size;--connotes also (in some applications) either criminate usage of the term leaf (q.v.) to refer to any one of the diverse
a fallure to fulfill a nonnal function, or a diminution in the expected forms of sheaths borne on the vegetative axes of the bamboo plant~
number of parts in a set (of stamens, for example). Cf. Depauperate. two expedients are followed: (1) the adoption of the differential terms
* Rt:stncled (L. restrietus). Confined;-describes the insertion of a pleioclade rhizome sheath, neck sneath, culm sheath, branchsheath, leaf sheath,
branch complement, when its subsidiary components all arise directly and prophyllu~ (qq.v.); and (2) the differentiation of the sheath

--'.- ~-~"'~~--"-"------ ~-~._-~ .. _ .. _-

 GLOSSARY 315
314 GLOSSARY

proper from its appendages (Fig. 35). A c1ear precedent for this effort alone, to a sheath proper with all of its appendages, to the blade only,
to avoid ambiguity is found in the now fairly common adoption of a or to a bladelcss sheathing structure, such as a prophyllum. See Leof.
precise terminology to differcntiate the several types of sheathing SMot (noun, Fr. AS. sciOtan, to move rapidly). "(1) A young growing
structures borne on the reproductive axes of the bamboo plant, as brauch or tV/ig; (2) the aseending axis; when scgmented into dissimilar
bracts, prophylla, glumes, lemmas, and paleas. See Sheathing orgtm. members it becornes astern" (Jackson). See Cu/m shoot.
t Shealh Made. A distinct foliar part, the lamina, that is appended apically Siliea bodies; also silica corpuscles (G. KieseIMrper). Bodies of siliea that are
on the laminiferous culm sheaths proper and branch shcaths proper in sccretcd and persist within the cells of various tissues .of the plant,
any series. A sheath blade is disdnguishable from a leaf (q.v.), first of particularly any epidermal layer. In both size and sru:pe, they range
a11 by the relative!y proximal position of the sheath proper of the widely. At one extreme, they are smaIl, lack charaetenl:o'tlc shape, and
former on any aerial vegetative a.xis) while leaf sheaths are aIways occur solitarily or in small numbers within a given eell. At the other
inserted at the distal nodcs of any eulm or branch. Sheath blades extreme, they may be large enough indi,1dually to fill a cell more or
are always sessile, while the leaf blades are petiolate (in all known less eompletely (see Siliea cells) and have a eharacteristic shape. Met-
bamboos except same species of .Neurolepis). Moreover) characteristic calfe (1960:"lii) lists 20 "types" (fonns) of silica bodies, some of which
differences in shape are usually quite markcd. The strong divergcnce are illustratcd in Fig. 1 of the same work. He states (p. xx) that "the
in form, as bet,,-een Jeaf blades and sheath blades in the barnboos silica~bodies in silica~cells assume very characteristic forms when the
is One of the eomrnonest and most useful of the grass morphologicai grass tcaf is mature, and are of considerable value for diagnostic and
distinctions between members of the Bambuseae and those of the taxonomie purposes." Ohki's key to Japanese genera of bamboos
other gramineous tribes. See Leof. (reproduced on pp. 73f) refers to the oecurrence of "silica eorpusdes" in
tSheath ."/lw; (L. • a/lw, hard skin). A somewhat prominent ring of paren- thc long epidermal cells of the leaf in Dendrocalomus [D. latiJWrusJ and
chymatous (not hard) tissue which remains at a sheath node after the in thc articulation (bulliform) cclls of thc lcafin Si1wbambusa [So loornk]
abseission of a sheath (of a eulm, especially) in some bamboos (Phyl- and Chimanobambusa [Ch. quadran"uutaris].
lostachys "idulario, for example). Cf. Girdle. . Silica cells. Epidermal short cells each of which is more or less completely
t Sheath "ode. The circumaxial locus of insertion of a sheath on any vegeta- filled by a single silica body (Metcalfe 1960:xx).
tive axis of a gramineous plant (elaborated after Hacke!; Lamson- tSpicaJe (L. spieatus, bearing spikes or can;). Having some or all of the
Scribller and Southwonh 1890:Fig. 1). Thc sheath node is marked , characters of aspike (q.v.). A spicate raceme is a raceme in which the
externally by a more or less prominent offset in the surface of its axis. pediccls are so short that the inflorescence resembles aspike.
At the level of each sheath node the crossing Over and anastomosis of tSpike (L. spico, an ear of grain). In the bamb",:s, aspike is a deterrninate
fib~vaseular bundles take place through a diaphragm whieh marks
infloreseencc in which the sessile or subsesinle spIkelets are lnserted
the mterruil boundary between adjaeent internodes. CL Culm node; see on a solitary rachis. Sincc the distinction between aspike and a spi-
Diapnragm. catc raceme is. in the terms of their definitions, only a matter of
t Sheath sear (Gr. eseherra, mark). J aekson defines Sear as "a mark left on a the relative lengm of "subobsolete" pediee!s, it is sometimes difficul t to
stern by aseparation of a leaf, or a secd by its detachmcnt; a cicatrix." choose between these terms for the description of a given inflorescence
In the bamboos) a sheath scar i5 a narrow, transverse. crrcumaxiaf (as in some species of the genus Merostachys, for examplc). According to
Jackson's definition (not applic.able hete) aspike is an "indeterminate
trace, the loCHS of abscission of a sheath proper. A sheath sear marks

I
inflorescence with flowers sessile on a common elongated axis." A loose
the position of a sheath node (q.v.).
tSheathing organ. Any sheathing structure inserted at anode of auy vegeta- interpretation of Jackson's definition of aspike has resulted in the
tive or reproductive axis in a gramineous plant. Among the bamboos~ oecasional misapplication, in the literature, of the tcrm spike tO a
distinguishable types of sheathing organs are rhizome sheaths, neck bamboo spikelet (q.v.).
sheaths eulm sheaths, branch shcaths, leaf sheaths, prop~ylla, bracts,
j
t Spike/ei (L. spicula). A basic strucrural eomponent of every normal gra-
ernpty glumes, lemmas, and paleas. Unless used) and interprcted} ,,·:ith mineous inflorescence, comprising a segmented axis (the rachilla) and
diseretion, the often~eneountered term foUar organ is apt to be ambig- its appendages (Figs. 48 and 52). The appendages (heginning with thc
UOUS, since it may be construed as referting either to a sheath prOper
lowennost) are: empty glumes (usually 2, rarely more, sometimes only
 316 GLOSSARY
GLOSSARY 317
1, rarely lacking entirely), lemma. (either variable or invariable in sympodium; for example, a system of related .axes wherein successive
number~ according to the taxon involved), and brauches of the rachilla branches aSSume the role or position of effcctive leadership or domin~
(one subtended by each lemma) each hearing a palea and the parts of anee;-a term used earlier (McClure 1925) to designate the branching
a ßower, In some taxa, oue or mOre of either the distal or the proximal habit of the type of rhizome described herein as pachymorph (q.v.).
lemmas,. or b01;h, may be sterile by virtue of either being empty or Cf. Mrmopodio.L
subtending an Incompletely developed ftower. See Floret; Flower. Taxon (neo-Gr.l' from Gr. taxis.. arrangement). "A taxonomie group cr
Spar.kJg~am (Gr. spo~o~ ashes; gramma, something drawn). A graphie illus- assemblage of plants or animals having certain characte:t:istics in com-
tration o~ thc mlcro~copic details of epidermal anatomy; or the special mon, \vhich we take as evidence of genetic relationship, and possessed
prep."'ratl?n on whlch it is based (Molisch 1920; Ohki 1923). Con- of $Ome degree of objeetive reality" (Rickett 1958). Rickett adds, ''We
ventlOnahzed or dlagrammatic illustrations of these anatomical fea- can use the word wherever we can use ~Taxonomic group~ in referring
tu",:, are the "d;,rmogram=" of Prat (1936: 178). to the characteristics, dynarnics, distribution, or uses of such an assem M

Sporod« (Gr. sporad,kos,. dispersed). "Widely dispersed or scattered" Gack- blage." Morton (l957) cautions that "where the words 'taxonomie
son); dJsperscd, or Irregular, in time;-as when the individual plants group' cannot be appropriately substituted, the word taxon is mis-
of a given generation of bamooos (from seeds of a COmmon orioin"\ used."
ent~ the reproductive phase at different times, or at irregular inte;als: Teratic (Gr. teras~ teratos, monster, wonder). Abnormal; teratologieal, in
In elther ,;"se, thc ftowering is said to be sporadic (Gamble 1396:viii). the sense of conventional usage, with particular refe:rence to marked
Cf. Gregarwus.
deviations from the normall or e."i:peeted, morphological expression.
tStoma (Gr., mouth~ opening; pI., stomata). A functional organ found com- See' teratic e.....ample of Proiijicatiofl: under Determmate.
~only, but not evc~here, in the epidennis that covers photosynthetic Terete CL., teres; wen~turned). Circular in cross section, tapered progres-
tLssues. Astoma CODSlstS of two guard cells and (in the bamboos as sively from one end to the other, and smallest at the distal end or tip
far as known) two subsidiary cells (Porterficld 1937). Changes in 'the (for example, a small bamboo culm). Cf. Obtmt&.
turgor of the guard cells result in the opening and c10sing of a s1itlike Testa (L., covering, in a poetic.al sense of the substantive). The outer coat
aperture (the stoma of Jackson) between them. of a seed.
Suh- (L.). A preß" indicating eüher an approximation or some reservation Tillering. HThrowing Out sterns from the base of astern" Gackson); pro-
er Illmtatlon In the use of the term ",,':ith which it i5 combined a" liferation of a culm /'rom its basal (subterranean) buds, without the
· , SIn
subdIstal} suberect} subfamily, subfusifonn. 'intercalation of a rhizome proper (Fig. 14).
Suhfosiform. Of a shape suggesting the concept fusifo rm. but not corre- Tra,ant (F., running, creeping). Ranging widely and freely;-a term used
sponding to it precisely; the approximate general s~pe of a paehv- by the Rivieres (1879:321 et passim) to characterize (1) the slender,
morph rhizome. See Furiform. '
elongate type of rhizome herein called leptomorph (q.v.), and (2) the
t Suhsidary orancllcs. Branches of higher order arising from buds at the base bamboos that spread by this means.
of a dominant primary branch.
Trt:adeiphous (Gr. treis) three; adelphos, brother). Having ~'filaments in three
Suhtend (L. suoUodo, I stretch underneath). To precede on a common axis brotherhoods";-an adje-.ctive cognate 'With Triadelphia} thename of"a
as a foliar organ precedes (subtends) a bud or branch inserted .imme~ Linnean order of plants with their stamens in three setsH Gacksen))
diately above it.
and conventionally used to describe the stamen complement of a
Snlcate (L. suicatus, fUITowed). "Grooved or fUITowed" Gackson);-as flower when its members have their filaments connate to form three
whc:e the otherwtse approximatcly cylindrical shape of the sunaee of distinct groups. In bamboos with triadelphous: androecia, the three
the mternode of a segmented axis is modified by one or more longi- HbrotherhoodsH commonly eomprise one, t\vo, and three stamens,
tudina~ depressions: In most bamboo genera the palea in funetional
ftorets I.sulcate, Wlth a single longitudinal dorsal depression.
Suture (F" fr. L. sutura" fr. S1,lere~ to sew). HA junction er seam of union' a
respectively. Cf. Morw.delphous; Dio.dciphous.
Turgidity (L. bdgUhJs, inftated). Turgor, turgescenee; the fumness imparted
by "the distention of a eell or cellular tissue by water or ether liquid')
line of ppening or dehiseence" Gackson). '
Gackson).
Sympodial (Gr. fYm or;J'1l, together; pous, pados, foot). Having the form of a Um·iateral CL. uniiateralis, fr. u:nus" one; latus, a side), One-sided. See Secund.

- - - - - - - - - - _ ......................_.....................................-
 318 GLOSSARY

t Vemation {L. vem(ltio~ rencwal). ('The disposition or method of arrange-

ment of follage leaves Vv1thin the bud" (Webster). Jackson's definition)
'~the order of unfolding of leaf buds,a is not applicable here. In most
knawn bamboos, the developing leaf blades are individually rollod up
tightly along their long axis 1 with onc edge at thc center of the roll.
Zygomorphic (Gr. zygos, yake; morph~ form). Symmetrically divisible by a
single planc;-((used of fiowers which arc divisible into equal halves in
onc plane onlY:I usually the anteropostenor)' (Jackson). ~ormal bam-
boo flo ....."'Crs conform to thi.. restricted enterion for zygomorphy (see Literature Cited
p. 114). According to JackSOIl, "Sachs extends the meaning to such
flowers as may be bisected in any one plane."

Numbers within brackets that follow each entry indicate thc pages and the
ngure legends where that partieular authority is cited. Hitherto unpublished
personal communications are also listcd here.

Acuiia Gale, Julian

1960 Persona! communication. [185]
Ahm.d, S.
1937 "Two years oid bamboo sced[ing/' [rulian Forester 63, 856-857. [274]
Ahmed, M.
1956 "Notenn the comparative success of SOille wmmon forest speCleS in
afforestation in Bihar/~ Indian. Forester 854 528-530. [217J
Alston, A, H. G.
1931 See Trimen 1900.
Arber, A.
1926 .!;Studies in thc Grarnineac. I. The flowe-..rs of certaln Bambuseae,"
Ann. Botan; (Lvndon) 40, 447-469. [107, 112, 171J
1927a "Studies in thc Gramineae, 1I. Abnonnalities in Ccphalostw;hytt.m.
viTgatum) Kurz, and their bearing on the interpretation of the bam-
boo flower," Ann. Botany (Lorulon) 41, 47-74. [101, 112]
1927b HStudies in the Gramineae. IIL Outgrowths of thc reproductive
shoat, and their bearing on the sign:ficance of lodicule and epiblast,n
Ann. Bota'!J (London) 41, 479-488. [107, ll2]
1928 "Studies in the Gramineae, IV, 2. Stamen-lodicules in Schizostachyum,
3. The terminal kaf of G(;;an.!Mhlou./' Ann. Botany (London) 42, 181-
187. [107, 112]
1929 HStudies in the Gramineae. VIII. On the otganization of the nower
in the bamboo," Ann. Bq,.", (Londm) 43, 765-781. [107, 112]
1934 Tltt Gramineae~ a study <if Cf:rca~ bamboo and grass (Cambridge Universlty
Press~ Cambridge, England). Some illustrations and passages of text
are reproduced with the publisher's permission. [13, 16) 17 {Fig. 5),
18,18, 61, 67, 79,85,98 (Fig. 49), 106, 107, 112, 113 (Fig. 54), 114,
115,116,117,122,126,128,130 (Fig. 59),164,297,303,305]
1950 Natural philosophy qf plant form (Cambridge Univcrsity Press~ Cam~
bridge, England). [61, 103, 105, 107]
Ascherson, P. F. A., and P. Graebncr
1902 Synopsis der miueleuropl:iischen Flora (Engelmann, Leipzig)) vor. 2, part L
[281, 282]
 320 LITERATURE CITED
LITERATURE CITED 321
Backer} C. A.
Bush, G. P., and H. H. LöwelI, cds,
1928 Htmdbock dc Flora van Jaza (All 2, Ruygrok, Batavia)~ bambOos,
V()(}T 1953 Teamwork in rcsta:reh (American University Prl"':SS. Washington, D.C,).
pp. 1-6,260-289. [l79J
[6J
Barnard, C.
Carous) E. G,
1957 "Floral histogenesis in thc monocotyledons. 1. The Gramineae," Aus- 1913 Lu Bambusies-rr,qnbgraphü:, ln'ol()git~ culture, princi'paux usages (Lechc-
tralian J. BoL 5, 1··20. [l17J valier, Pari,). (162, 263, 282]
Bean,W.J.
GalToll, Lewis
1907 wThe flowering of cultivated bamboos," Kew Bul4 Mise. In/arm. 1895 Through [he [ooxing glass and what Altee found there (Altemus, Phila-
(1907),228-233. :268, 269, 271, 272, 278J delphia), chap. vi. [295]
Beddomc, R. H.
Champion Papcrs~ Inc.
PS73] F<Yfcsü:r's manual Gi oota7!Y JOT Southem lruiia (Gantz Eros., Adelphi Personal communications. IHiO. 164, li6, 192]
Prc.-.;;s, Madras), bamboos, pp. re.x.xix-ccx.'XXvi; pI. x,'(viiL [115, 197,
Chang, F. C.
199,200J
1938 "A crysralline compound from the whire powder fauod on Bambusa
Bentham, G. chun"ii" Lingnan Sci. J. 17, 617-622. (2, 46)
1883 "Gramineae/' in G. Bentham and J. D. Hooker) Gerura plantarum 1941 ~(L Alkylation of quinones with esters ..~; tetrav~ent lea.d. IJ, !ri-
(Recvc, Landon), vol. 3, part 2. [282J terpenoid ketones from Bambusa chungu~ unpubhshed dissertatIOn,
Bhargava, M. P. Harvard University [not seen]. [46J
1946 "Bamboo for pulp and paper manufacture. I-lIIt" Indian Forat BulL Chaturvedi, B.
(n.s., "Utilization"), No. 129 (1945 [1946]). [171, 197J 1947 HAerial rhizomes in bamboo culms/' Indian Fort!ster 73.. 543; Pt 33,
Bibral) B. Fig. I. [247]
1899 "The flowering of secdlings of Dmdrocalamus strt'ctus," Indian FQr(s[cr Chaturvedi, ?o.'!, D.
25, 305-306. [274J 1928 ('Influencc of ovenvood on the deve10pmenr of bamboo (Drndr()-
Blatter; E. calamus striclus)/' United Provinees Forcst DelJartment Bulletin, No. 1, 1-8.
1929 "Indian bamböos brought up to date," Indian Forestc 55, 541-562; [170]
586-612. i28SJ Church, A, H,
1929,
1889 "Food grains of India/ 1 Kew Bull. Mise. Inßmn. (1889)} 283-284.
1930 "Flowering of bamboo, I-III,') J. Bomhay Nat. Hist. Scc. j3~ 899-921; [! 71]
34, 135-141,447-467. [84, 171, 1%, 273J
Glement, l D.
Bor, N. L. 1956 [Flowenng of Dmdrocalamus stt:'ctus at Atkins Garden, So!edad, Cien-
1941 "Thyrsostachys oliven Gamble/~ Indian Forest Rec. (nevl series) fuegos, Cuba,J Scien" 124;. 1291, [I68]
Bntany vnl. 2, pt. 2, 221-225, pI. 65-66. [279]
Cobin, ::\1,
Brandis, D. 1947 aNotes on the propagation of tbe sympodiai or clump type of bam~
l874 Forest F!(ffi;l of nort/um:Si ar.d ccn.l:raJ Indio. (Allen, London); bamboos, 000'," Proc. Fla. Stak Hort. Soc. (1947) 60, 181-184. [242, 245J
pp. 560-570. [171J
1899 Crcech, J. L.
"Biological notes on Indian bamboos," Indian Fvreseer 25, 1-25. (122, 1950 Personal communication. p56, 2411
126, 144, 274J
1906 Indian trees (Archbold Gonstable 1 London); bamboos, pp. 600-685) Dabral, S. N.
719-720, [84, 162, 199] 1950 "A preliminary note on propagation of bamboos from culm seg-
1907 <IRemarks on thc structure of bamboo le~wes," Trans, Lir.1J.. Soe, Lrm- ments," Indian F{J'fcstn' 76; 313~314. [217, 235J
drm. [2, Bot.nyJ 7,69-92. [71, 72, 75, 120] Darlington, C. D., and A. P. Wylie
Brown, R. W. 1956 Chroml)somc atlas:::I jlowtring plants (Macmlilan, New York); bamboos,
1954 Compositirm ofscientific words (published by the aurnor). [297, 304. 305J pp, 457-458. [199J
Brown) W. V~ Deogun" P. N. .
1937 "The silvlculture and management of the bamboo DmdroeawJllus
1958 "Leaf anatomy in gt""" ')'Stem.des," Boia", G=. 119, 170-118. (75J
strietus Necs 1 Indian Forest Rec. (o,s., Silviculture) 2, 75-173. (132,
ll

1960 "The morphology of !he grass embryo," Phytorrwrplu;logy 10, 215-223.

(120] 164,166 (Fig. 74),167-171,201,204,205,211,216, 235J
Bryan, G. S. Dusen, p,
1903-
1926 Edison: Th, man 11M Ais work (Knopf, New York), [2]
1906 {{The vegetation of \Vestern Patagonia," in J. B. Hatcher and W. B,
322 LITERATURE CITED LITERATURE CITED 323

Seott, Reparts oJ the frinccton Universi{y Expeditions to Patagonia, 1896- Godron, D. A.

1899, voL 8, pt. 1, "Botany/) pp. 1-34. (Publishcd by Princeron 1880 "Les bourgeons axillaires et les r.ameaux des Graminees," Ra:. Sei.
Gniversity, Princeton, N.].) [3] Not. viii [n.s., .], 429-442. [67J
Durra, J. Gupta, :M.
1938 "Bambus-ees de Rio Grande du Sud/' Rev, Su.damtricana Satan. 5, 1952 '"Gregarious fiowcring of Dtndrotalamw; striclus/' Ina'ian FMester 78>
145-152. [250, 275] 547-550. [172]
Ezard, C. T. B. Hacke], E.
1946 Personal communication, [159] 1881 "'Untersuchungen ueber die Lodiculae der Graeser,'" Engler Botan..
Federa.1 Experiment Station, ~fayagücz) P. R. Jahrb. I, 336--361; bamboo. p. 357. (114J .
1950- 1887 "GramineacJ " in Engler and Prand} Die natÜrlic.hen PflaTl.Zen.fomilitTi
1953 An.ual Rep...L [207] (Engelmann, Leipzig)) Teil 1I) Abt, 2, "Bambuseae,') pp. 89-97. In
Ferrer Delgado. R. the presem work, citations of this pub:ication rder to the English
1948 "La propagaci6n dcl bambu por csquejcs," Rev. Ap. Pw:rto Rieo ~9" trans)ati?n by Lamson~Scribner and Southworth (q.v.). [48,282,299,
3-7. [263] 302, 314J
Fibres 1899 "Enumeratio graminum japoniae/' 8ull. de l1lerbier Boissier ~old seL)
1947 ;;Bamboo pulp: its possibilities for rayon rnanufacture," Fibres [Eng- 7, 701-726. [70]
land! 8, 82-84. (197] Hageman, R. H.) R. FeITer Delgado, and N. F. Childers . .
Fischer. C. E. C. 1949 "The use of dynamitc in lifting bamboo c1umps for propagatIOn,"
1934 Graminew:, part X of J. S. Gamble, Flora 0/ Iht Presidmcy 0/ Madras Tlop. Agr. 26, 122-123. [211J
(Published under thc authority of thc Secretary of 8tate in Council, Hes1op~HaITison, J.
London), pp. 1690-1864. [200] 1959 HGrowth substances and flower morphogenesis,') J. Linn. Sot. Landon
Franchet, M. A. 56, 269-281. [6, 105J
1389 "Note SUT dcux nouveaux genres de B,ambusees/~ J Botanique 3J Heyne, K.
277-284. :88J 1950 De nutti'ge Pla."lten van lndofltsie (cd. 3~ van Hocve, Thc Hague, 1950);
Freeman-Mitford, A. B. bamboo" vol. 1, pp. 285-304. :176, 177]
1896 The bam!;oo garden (Macmilian, London). [72, 263; 297] Higgins, J. E., and W. R. Lind'.,
Gamble,]. S. 1939 Annual rt:purts qf tlu etmal Zone Experimtnt Gar-dtrzs [oT the fisc.al yea.rs
1888 "Notes on thc sma:l bamboos of the genus Af'.uuiinan-a/' Indian For- 1935 and 1936 (Panan;ta Canal Press, Mount Hope, C.Z.). [275]
'SI" 14, 306-314. [271] Hildebrand, F. H,
1896 "Bambuseae of British India," Ann.. Roy. Botan. Garden Calculta 7~ 1954 HAantekeningen ova Javaansc B.ambu-Soorten. Notes on Javanese
1-133. Atlas, platcs 1-119. [67,83,89, 115. 158 (Fig. 71), 164, 165 bamboo species J ' [in Dutch; English summary], Laprnatl Balai Ptn-
(Fig_ 73), 167, 172, 173 (Fig. 75), 177, 188 (Fig. 81), 192, 196, 197, jelidikan. Kehutanan. Indonesia Forest Res. Inst. Rept. 66. [82]
198 (Fig. 86), 199, 200, 201, 270, 276, 304] Hisauchi J K,
1921 "Flowering of Arundinariafalcata in the Tcmperate Housc/' Kew Bult. 1949 NSeedlings and deve10pment of rhizomes of a bamboo (Arundinaria
Mise. Infom.. (1921),302-306. [271] nikkocnsis l'lakfJ.1)" [in Japanesel~ J. JapWL Botatrl 24~ 24-26. {l36, 137
Unpubllshed sketch, courtesy of thc Keeper of the Herbarium, Royal (Fig. 64)]
Botanic Garden" Kew. [119 (Fig. 55, 13)J
Holttum, R. E.
Ganison, R. 1946 "The classification cf rVialayan bamboos," J Amald Arbcrctum 27,
1949 i!Origin and development ofaxHlary buds: Syringa vulgaris L.," Am. 340-346. [83J
J.BQsany 36, 205-213; '<Belu.la papyriftra and Euptelea polyandra," ibid., 1955 ';'Growth~habits of monoootylcdcns-variations on a theme/' Pa/ta-
379-389. :62] morpholollJ' 5, 399-413. [85, 123]
Ghinkul, S. G. 1956a (!Classification of bamboos," Phytomorpholo,?Y 6, 73-90. [93~ ~07, 116,
1936 "]\fass blossoming of bamboo in thc light of phasic development 1~ .
theory" [in Russian; EngHsh summary], Soviel SubtTopicJ 10 (26), 1956& uThe type species of the genus Gigantothlofi and some other notes on
24-29. [85] the genus/' Taxon 6, 28-30. [179]
Giovannoni, M., L. G, C. Vellozo, and G, V, L, Kubiak 1958 I'The bamboos of the Malay Peninsula," Gard. Bull. (Singapore) 16,
1946 "Bobre as 'ratadas' da primeiro p:analto paranaense/' Arq. BioZ. 1-135. [70,96,106,120,148,177,179]
TtdmoL 1, 185-195. [86] Personal communications. [134, 177]
 324 LITERATURE CITED LITERATURE CITED 325
Hooker) J. D. Janaki Ammal, E. K.
1854 HimalayanJoumals Oohn Murray. Lendon); bamboos) vot L [272] 1938 "Chromosome numbers in sugercane X bamboo hybr:d.s/' Nature 11:1J
1885 "Chusql1.t:a abictiJolia," Cu.rtis's Botan, JY1ag. [3] 41" Tab. 6811. !273J 925. [5. 206]
HOMn Japanese Patent No. 175 1685
1959 ('Thc debut of the picture review," Honztm 1) pt. 4, pp. 68-69. Based 1948 "Deodorization cf ehrysalis eil or fish on" [in JapaneseJ} issued 10
on a selection of pictures and texts taken [rom Peter Pollaek's Picluri: C. Oyama, Febroary 27, 1948. [2J
Hiswry 0/ PhQtographYJ and featuring an interview between G. F. Jones, Q.., and I. A. Wolff
Tournachon (as photographer, with his son at the camera) and 1960 "The search for new industrial erops," Ecan. Botany J4, 56··68, [149]
Marie-Eugene Chevreul: on the eve of thc latter's one hundred first Jones, Rebeeca
birthday. On this occasion, Chevrcul quoted words [rom Male- 1935 Letter addrcssed to U. S. Department of Agriculture. [268}
branche as e."\:pressing his own "basic philosophieal pnnciplc." Chev~
roul refen: to :Malebranche by surname only; hut 1t seems reasonable Kadambi, K.
to suppose that the Freneh philosopher Nicolas Malebranche (1638- 1949, HOn the eeology and silviculture of Dendrocalamus strictus in the
1715) is the author of the passage, "One must strivc for infallibility bamboo forests of Bhadravati Division, Mysore State, and compara~
without pretending to it." [8J live notes on the species Bambusa aruna'inacea, Ochlan.d:m travancorica,
0:!9'tcna:nthera monost~gma, and O. stocksi~" Indian Forester 75~ 289-299~
Houzeau de Lehaie, J., ed.
334-349, 398-426. [172]
1906-
1908 Le bam/x;u, Nos. 1-10. [19, 20 (Fig. 7), 263, 271, 274J Kato, K.
1911 (tUeber fermente in Bambusschösslingen," Ho.bpe~ScylC"fs Z. Physiol.
Hubbard, G. E.
ehern. 456-474. [2J
1948 "Gramineae in J. Hutchinson> BrÜish fowmng plants (Gawtho~.
j "
Kawamura~ S.
London). ,282J
1927 "On the periodieal flovtering cf the bamboo/' Japan. J. Botany 3,
Hubbard) C, E,~ .'lnd R. E. Vaughan
335-349. [84, 104 (Fig. 51), 278J
1940 Thc grasscs af A1aUTitius and Rodriguc:z (Royal Crown Agents fot the
Gelerncs, Lenden). [159J Kelsey, H. and \I\r, A. Dayton
1912 Standardized plant names (American Joint Committee on Horticultural
Hughes, R. H.
Nomendaturc; j, Horace McFarland Co.) Harrisburg, Pennsylvania,
1951 ('Observations of cane (Arona'irzaria) flowers~ seed and seediings in the cd. 2). [292]
North Carolina Coastal Plain," Bull. Torr", Botan. Club 78, !l3-121.
[86, 138, 204J Keng, K. H. [A. W.: Keng Pai--chieh and Keng, f.J
1948 "Preliminarv study on the Chinese bamboosn [in Chinese], NalL For-
Humboldt, A., and A. Bonpland
tstry Res. BUr. China. Tech. Bull., No. 8. [22 (Fig. 9), 23, 31]
1808-
1809 Plo:ntae aegu.ir-.octiales (F. Schoell) Paris); bamboos) voL 1) pp. 63-69,
pI. 20,?1. [179, 181]
, Keng, Y. L" cd.
1959
" f;
Flora illustralis pla:niarum primaro.m stm,t;(ffum. G:ammeae. [Agency. or
Sdenee PubLications, Peoples Republic of Chma, ,Pek;ngJ. hoov :-
1181 p') illus., index; te..xt in Chinese; title and sctcntific names m
Personal communication. [125]
in Latin. [282J
Indian FMcster
1875- Notices. [85J Kennard, W. C, . , . , n
1955 ('Flowering of thc bamboo Guadua. amplcxifo/w. Presl In Puerto RlCO,
JaOOon, B. D. Lloydia 18, 193-196. [86, 275)
1949 A gWssary of betan;,;al terms (cd. 4, Duckworth, London). [70 " passimJ
Kenncdy, P. B,
Jacques-Filix, H.
1899 "Thc structure of the caryopsis of grasses with refere~cc to th;i.r
1955 "Notes sur les Graminees d'Afrique tropicale. VI. Les Graminees morphology and elassification," U. S Department oJ Agncutture) D,Ol-
Africaines de type arehaique," J Agr. Trop. BollJJ4 Appl. 2, 423-430. sioff. if Agrostolog}', Bull. 19; bamboos, pp. 35--37, [120]
[75J
1958 "Notes sur les Graminees d'Afrique tropicale. xn, Structute foliaire, Kety, S. S.
"oologie et systematique," J. Agr. Trap. Bo_ Appi. 5, 809-825. [120J 1960 "A biologist examines the mind and bchavior/' Sr;ieru;c 132, 1861-
1870. (6J
1962 '"Lei Graminees cl'Afrique tropicale. L Generalites, classification~
description des genres," Inst. Res. Agron. Trap. Paris BulI. Sci.~ No. 8. Koestler, A,
[75J 1964 TM oel of mali•• (Macmillan, New York). [7J
 326 LITERATURE CITED
LITERATURE CITED 327
Komatsu S.) and Y. Sasaoka
j

1928 «(Chinese handmade paper." unpublished dissertation, Ohio State

1927 "Studies in Japancse plants, 'VIII. The OCCUrTence of free pentose in
University, Columbus Ohio. (11
bamboo ,hoots," Eu/I. ehern. Soe. Japan 2, 57-60. [2J j

1931a "Studies of Chinese bamboos. I. A new spedes of Anmdinaria from

Koorders, S. H.
southem China, Pt, 1. Diagnosis," Lingnan Sti. J. 10,5-10. [152) 153]
1908 "Contribution Da. 1 to the knowledge of the flora of Java (third 1931b "Studies of Chinese bamboos. I. A new species of Arundittaria from
continuation)," Proc. (Sec. Sci.) Roy. Acad. Amslerdam 11, 129-132. southem China. Pt. 2. Notcs on culture~ preparation for thc market
[119 (Fig. 55, 13)]
and uses," Lingr.n.n Sci. J. Jo, 295-305. [153]
Kraus; G. 1934 "Tbe infiorescence in SchizQstachywn Neest J Wash. Acad. Sc~', 24~
1908 "Gynaeceum oder Gynoecium?" Verhatu1!. Physik-ivfed. Gescllsch. Wur:- 541-548. [90,91,93, 100, 108, 281J .
burg (N. F.) 39, 9-14. [FrOfi Arber 1934; not seen.] [297, 305J 1935 "Bamboo,-a taxonomie problem and an economic opportunity,"
Run th~ Charlcs Sei Mon'hly 41, 193-204. [47 (Fig. 24), 88, 148)
1815 "Coruiderations generales sn! les Grarninecs," };fcm. Mus. d'Hist, Nat. 1937 "Bamboo, ~~ a tribute to bamboo published in the program annüunc-
(Paris) 2, 62-75. [282] ing thc Lingnan University Research Lecture for 1936-37, entitled
Kurz, S. «Sam boo as a ficld für research. n The lines reproduced herein were
1875 composed a.s an Hinvitatiün" to thc lecturc. [li}
Preli'mitulry reptrrt on the jfA'tst a;ul ether vegetation of Pegu. Appendix B 1938a "Notcs on bamboo cu~turc with special referen~ tö South China,"
(Baptist Mission Press, Calcutta), bamboos pp. 91-95 and index
[19~ , . Hong Kong NaL !I, 4-18. [242, 263]
1938b "Diary of a small experimental barnboo planting/' Lt'ngnan Sei. J 17,
1876 '(Bamboo and its usc," Indüm Furester l~ 219-269, pt I-li; 335-362,
473-476. [214, 216]
pI. III-IV. J46, 61, 118 (Fig. 55, J, 0), 119 (Fig. 55, 10), 120, 132 1941
134, 135 (Flg. 63), 159, 196, 227, 276, 288, 300] , "On same new and imperfectly known specles of Chinese bamboos,"
Sunyalsmia 6, 28-51. [70J
Kutty ,:!'unma, P. R. Bhagavathi, and T. Ekambarnm 1942 "Bamboo genera/' in H. Kelsey and W. A. Dayton, eds. StarUic.rdtzed
1940 "Sugarcane X bamboo hybrids," J. Indian BQIan. Soe. /8, 209-229. [5] plant name,! (00. 2j j, Horace McFarland Co., Harrisburg, Pa.); ham-
Lamson-Scribner, P.} and E. A. Southworth boos, pp. 40-41. (292]
1890 The 'rue gr~ses~ by Eduard Hackel, translated tram Dt'c natürHchcn 1944 Western Hemisphere bamboo$ as subst1.'tutes Jor OrUntal bamboos Jor thc mfJ.1'aJ,-
Pflar.zctifamd,cn (Holt, New York); Bambuseae, pp. 193c211. Refer- /atMe cif ,rki pole shafts (U. S. NatI. Res, Councit Committee on
ences to .Hackel (18~7) in thc present work ure given as pages in Quartermaster Problems. Final Rcpt. QMC-24). (150, 187]
the Enghsh translatIOn by Lamson~Scribner and Southworth. [48 1945a "Bamboo culture in thc Americas,H Agr. Am. 5, 3-7, 15-16. r25) 263]
282, 299, 302, 314) , 1945b leThe vegetative charactcrs of thc bamboo genus Phyllostachys and
Li; Hui-lin descriptions of cight new species introduced from China/' J Wasn.
1942 HBamboo und Chinese civllization,~' J. New YM'k Botan. Gartkn 43,· Acad, Sei. 35, 276-293. [1, 23, 25, 116, 301]
213-223. [1] 1945e üBamboo in Ecuador's lowlands," Agr. Am. .5;. 190~192, 194. [187]
Lin) W.C. 1946 :'The genus Btlmbusa and some of its first~known specics," Blwnea,
1962, Supp. III, 90-112. [23,25, 27J
1964 1948 HBamboos for farm and horne," U. S, Dept. Agr. Yearbook 1948, 735-
"Studies on the propagation by level CUttings of various bamboos"
740. [2, 159)
[in. Ohinese with Engiish titles und summary], pts. 1 and 2. Rul/. Bamboo as (1 bw'lding material (U. S. Dcpartrnent of Agricultwe, For-
1953
Ta,wan Forcstry Res. Institule Nos. 80 aruf J05. [234J
eign Agricultural Service, Washington, D.C.). [1, 181 (Fig. 78), 183J
Lind,ay, W. R.
1955 HBamboos/' inJ. R. Swallen, HGrasses ofGuatemala/' in P. C, Stand~
Personal communication, (275] ley and J. A, Steycrmark. eds. 1 Fwra Q[ Guatemala;, pt. TI, Fieldia.na
Link, H. F. (Botany), vaL 24, pt. II; bamboos, p. 2 (key to genera) el pa.mm;
1833 Hort1J.s regius botanicus bcrolinensis (Reimer, Berlin). voL 2. [282J the genera of bamboos and other grasses arrangcd in one alpha-
Loh, C. S,) T., H .. Hu l P. T! Ma, and P. M. Tseng betical sequcnce. [164, 185]
1950 <CA report on Saccharum X bamboo hybrids,'~ Arm. Pro<:. Agr. Sec. China 19500 HBamboo utilization in Eastern Pakistan," PakistanJ. Forestry 6, 182-
[TaiwanJ 5, 13-33, pI. I-IV. [5] 186. Some problems relating to long-term procurement of bamboo
Maclood; A M, and L. S. Cobley, eds. for the KamapbuJi Paper Mi!!. [190)
1961 Con!r:mporary botaniea/ tlwu,ght (Oliver ancl Boyd, London). [6] 1956b '~Bamboo In thc economy of oriental peoples," Eton. Botany 1~ 335-
McCiuTe, F. A 361 [1]
1957 Bambcos q[ tne genus Phyl!ostachys U1Ider cultivation in the United S!ales
1925 "Some observations on the bamboos of Kwangtung [China]," Li~ana.n
with a kr.;y Jer tkeir fieM identilicaLion (USDA Agr. Handbook 114). [52
Ag>'. Rev. 3, 40--47. [21 (Fig. 8), 25, 27, 308, 317J
(Fig. 27), 206]
 328 LITERATURE CITED
LITERA"IüRE CITED 329
1958 "'Bamboo as a SQurce of forage/' Pacific Sei. Assoe., 8th Pacific Sei. MOliseh, H.
Con,gr. Proceedings (Manila, R. P.) 1953), vol. iv B, 609-644. [2] 1920 41 Aschenbild und Pflanzenverwandschaft/' Sitzber. Abd. Wiss. Wein,
1961a "B~nboo:" EncyclopaedJ'(J, Bt7:tannica, val. 3, pp, 17-18. Reprodueed in Math.-NatUl'w. Kl., Abi. I, 129, 261-294; bamboo, p. 277 and PI. II,
revlsed and emcnded form~ with thc publisher's pennission. [1] Fig. 8. [73, 316]
1961 b "Toward a fuller description of the Bambusoideae (Gramineae)/t Kcw
Buil. 15, 321-324. [282J Moonev, H. F.
1933' "Thc fOTe!>"ts of thc Orissa States/' Indian Forester 59, 200-221. [172]
1963 ('A new feature in bamboo rhizome anatomy/' RhodrJra 65~ 63-65.
[13, 23J 1938 <'A synecological study of the forests of western Singhbhum," Indiat1.
Foml Rec. [n.s., Silvieulture] 2, 259-356. [172J
1964 "Bambusoideac," in B. Maguire, j, J. Wurdac-.k, and col1aborators
('Thc botany of the Guayana highland-pt, V,'; i'Jem. New York Bota: Morton, C. V.
Card. 10, 1-6. :42] 1957 "Thc misuse of the term taxon/ l Taxon 6, 155. [317}
MeClure, F. A., and W. C. Kennard Munro, W,
1955 (Propagation of bamboo by whole-eulm cuttings/' Pr«, Amer. Soc. 1868 "A monograph of the Bambusaeeae~ including deseriptions of al1 the
Ho,",. Sci. 65, 283-288. [207, 229, 232 (Table 5), 244J species," Trans. Linn. Soc. Londen 26, 1-157. [84,85, 89, 106, 114, 116,
117,271,272,273J
McClurc, F, A.} and P. F. Li
Muro!, H,
1941 ["ruustrated guide to selected bamboo gcnera~~ (in Chinese)J, Scimce
(China) 25, 285-292. [39 (Fig. 19)J 1956 Takt to sasa [in ]apanese: "Bamboos and sasas"j (Tokyo). [118 (Hg.
MeClure, F. A., and F. Montalvo 55,2),119 (Fig. 55, 16, 17, 19, 21), 127 (Fig. 57),138,139 (Fig. 65),
274J
1949 "Bamhusa lfFngispicu!.n.ta easily propagated from stump layets," Puerto
Rico Fed. Expt. Sta. Q!.wt. Rept. Apr.lJM', 1949, pp. 102-104, Table 1. Nakai, T. d
1925 'ITwo new genera of Bambusaceae with special remarks on thc _relate
[254 (Table, 8, 9), 263J
1950 "Propagation studies," P>.u:rlo Rieo Fcd. Expt. SIe.. Ann... Rept. /950,
species growing in eastern Asia," 1:. Amc~a' ~rboretum ~ 145-1.:>3, {53]
1933 Flma sylrlatica koreana. Pars 20~ 1-5.); pI. i-XlV (Forest Expt. Stauon,
pp. 27-28. [263]
KeijY01 Japan [Korea]). [Text in japanese and Latio, with some
Makino, T.> and K. Shibata
English. J [23, 282J
1901 "On Sast; a new genus of Bambusaceae and its affinities," Balcm.
Mag. Tokyo 15, 18-31. [81, 94 (Fig. 46)J Nally, Julian
[1953.] Letter to U.S. Department of Agriculture. [271 J
~tfalebranche. See HiYfi=on
Nees, C. G.
Mathauda, G. S.
1829 "Agrostologia brasiliensis," in C. F. P. :b.fartiusl Flora brasilitnsis
1952 HF10wering habits of bamboo-Dcna'rocalamus strictus/' Ina'~'an F'orC$ter (Engclmann, Leipzig), val. 1, part 1; bamboos, pp. 520-538. [89, 93,
78, 86-88. [169, 172J 282J
Metcalfc, C. R.
1954 "An anatomist's viCY\'S on angiosperm c1assifieation,'> Kew BulL (1954),
, 1841 Florae Africac australioris (Prausnitz, Glogau), vol. 1, Gramineae~' bam-
000', pp. 460-464. [89J
427-440. [6J Netherlands Patent No. 53,471
1956 "Some thoughts on the strueture of bamOOo leaves," Bettm. Mag. 194'2 «Werkwijze voor het uitvoeren van katalytische reaeties/' issued to
Tokyo 69, 391-400. [72, 75J N. V. de Bataafsche Petroleum Maatschappij, The Hague, November
1960 Anatomy of tht mClW)cctyledons, val. 1, Gramincae (Clarendon Press, Ox~ 16, 1942. [2]
[ord, England). Some passages from the text are reprodueed with
Nicholson, J. W. .
the permission of both thc author and tbe publisher, [75, 315] 1922 '''Xotes on the distribution and habit of Dendrocalamus stnctus and
~fitford. See Freeman-Mitford
Bambusa arundirwcea in Orissa," lr.dian Forestet 48> 425~28. [170]
Miyake, I., and G. Suglura Nozeran, Rene .
1950, 1955 HContribution a 1'6tude de que1ques structures florales (essal de mor~
1951 ~(Researeh for thc activitive [st'c] charcoal ofNemagari-bamboo (Sasa phologie florale comparee)," Arm. Sei. Nat., Botan. Biol. 16, 1-224.
panicuiata) materials. Special researeh abom thc carbon of air elec- [l05J
trie battcry. r, II," Japan. FoJ"eStry Soc. J. 32, 181-185; 33, 207-21l. Nyasaland Forestry Department, Zomba
[2J
1944 "The common local bamboo," .1vyasalar.d Agr. Quart. J. 4 (3), 8~13.
Moebius) 1\1. [217,276J
1898 "'Deber ein eigentümliches Blühen von Bambusa vulgaris \Vendl.t Nybakken, O. E.
SenckerWetgische Naturforsck Ces. (Frankfurt a. M.), 81-39. [82. 90, 93, 1959 Greele and Latin ir. scient~'ic terminology (Iowa Statc Dniversity Press,
96, 164J Ames, lowa), [297, 305]

... ~ .. ~- .. -~--_ ....- ~~~- .. _~ ..

 330 LITERATURE CITED LITERATu"RE CITED 331

Ochse, J. j., in coHaboraüon with R. C. Baekhuizen van der Brink 1954 "Das System der Gramineac," En.gler botanische Jakrbuclur 76~ 281-384.
1931 Vegttables'!f thc Duten East Irufies (Arehipel Drukkerij, Buitenzorg, Published posthumously; edited by Dr. Eva Potztal; reviewed by
Java); bamboos, pp. v-xii; 301-327. [177J Jacq\les-Felix (1955) and Parodi (1955). [281, 282, 288J
Ohki, K. Plank, H. K.
1950 aFaetors influcneing attack and control of the bamboo powder-post
1932 "On thc systcmatic importance of spodograms in thc leaves of Japa~
nese Bambusacea.e, u J. Tokyo Imper. Unw, FacuL Sd. (111 Bot.) 4~ beetle," Bull. Pucr'a Rico_ Fd. Exp'. Sta., "0. 48, pp. 1-39. [157, 172)
1-130. [73, 316J Po Chu-i (A.D, 772-846); see A. Vvaley
Oliver. D. Portcrfie1d, W, M. . .
1894 "Phyllostach)'s hctt:roclada Oliv., sp. nov.,') in \V. J. Hocker, 00,> Icones 1923 'lA new feature in vascular anatom)' as disp~ayed by bamboo) parti-
plu:ntarwn (Dulau, London), vol. 23~ pI. 2288 and one page of text, cularly by the young sheath Jeaf," China}. Sei. A.rts 1, 273-279. [69J
[1l7) 1926 '<The morphology of thc bambQo flower with special rcferenee to
Page, V. M. Ph)'llostach)'s nidularia, Munro," Chi1Ul J. Sei Arts 5, 256-260. [93, 99
1951 HMorphology of the spikelet in Strepto$hact~" Bul!. Torrey Bot, Clu./; 78~ (Fig. 50), 110 (Fig. 52), 111 (Fig. 53), 277)
22-35. [108) 1928 "A study cf the grand period of gtowth in bamboo," Bul!. Tarrty Botan.
Parod!, L. R. Club 55, 327-405. [91, 123)
1930a "Morphology of thc growing point in bamboo," Bull. Ytnching Unw.
1936 ;'Las Bambuseas indigenas en 1a Mesopotamia Argentina, " Rm
Arg",ti,," Agroo. 3, 229-244. [186)
Dept. Bi,l. I, 7-15. [61, 62, 123]
1930b "The mechanism cf growth in bamboo," China}. 13, 86-91, 146-153.
1945 "Sinops!s de las gramlneas chilcnas del genera Cr.u.squ<a," Rev. Uni,.
(Univ. CatoL de Chile) 30, 61-71. [3] [42,123,124 (Fig. 56)]
1933 <'Thc periodicity of-bamboo culm Structures," China J. 18> 357-37L
1955 "La fioraci6n cle la Tacu.ara brava. ('G1Mi1uc, trin.it~)," Rel!. ArgentilUl
Agran. 22, 134-136. [84, 269, 275] [6, 12,42, 60) . .
1935 "Thc relation of shoo! roots to shoat elongauon 10 the bamboo
1961 '"La ta.'Xonomla de las Gramineae Argentinas a 1a luz de las investi-
Ph;yllostachys nigra," Am. J. Bata", 22, 878-888. [80 (Fig. 44, All
gaciones mas recientes/' Rectnt advfl:nces in bota.n; (from lectures and 1937 "Histogenesis in the bamboo with special reference to thc epidermis,"
symposia presented to the International Botanieal Congress5 :r\tlo:p.~ Torr!)' Bota... Club Bull. 64, 421-432. [75, 316)
treal, 1959; University of Toronto Press, Toronto), voI. 1, pp, 125-
130. [7, 302J Prat, H.
1936 "La systematique des graminees," A,m, Sei. Nat. LX, Botan·1 18" 165-
Partha..,arathy, N, 258; bamboos, p. 170 ct ,bassim. [n, 75, 282, 300, '316)
1946 "Chromosome numbers in BambU$eae/~ Cur. Sei. 15# 233-234. [167, HGradients histo~physiologiques et petfectionnement organique"
1954
2061 (Travaux de l'Instirute de Biologie Generale et de Zoologie cle
Pathak, S. L. PUniversitc de Montrcal , No. 62); exuait des Rapports cl ClJmmulI:icatians
1899 <'Thc propagation cf the common male bamboo by cuttings in the du VIIl" Congres Intematirmal de Bomniqu.e, Pari$~ 1954, Sec. 8, pp.
Pinjaur-Patiala forest nurseries," Indlan Forester 25, 307 ~308, [263, 294-296. [6)
279] 1960 "Vers une classifieation naturelle des Graminees,'J Bult. Soc. Botan.
Pereira, C, Fr""" 107, 32-79. [6]
1941 "Sobre as ratadas no sul Brastl e 0 eic10 vegetativo das taquaras/' Raizada, M. B.
Arq. IosL BiaL (SiitJ Paulo) 12, 175-196. [86, 276) 1948 "A litde-known Burmese bamboo (Sinocalanrus ropelandlJ." India."'I. Forester:
Piatt1> Luigi N, 7-10. [119 (Fig. 55, 22), 269, 270J
1947 '''Flüssige Brennstoffe aus bambus," Schweiz. Arch. angew. Wiss. Teck. Ray, David
13, 370-376. [2J Translation. [136]
Piooallu, A. " Rebsch, B. A.
1931 "La cwture des bambous," }. Agr. Prat. (n.s.) 56, 293-297. [263) 1910 'IThe bamboo (Dcndrocalanrus strictus) forests of the Ganges Division,
Pilger, R, t:. P.," Indian Forest'" 36, 202-221. [1721
1927 'CU"cber die Blütenstände und Aerchen der Bambuseen~Gattung Reeder, J. R.
Gu.adua Kunth/' Ber. Deut. botan. Geseluch, 45, 562-570. [90, 93) 1961 "The grass embryo in systematics/' Recent advances in ootarry (from lee-
193i "Gramineae," in L. DieJs, Beiträge zur Kamtniss dtr Vegetation und tures and symposia pre,sented to the IX International Botanical Con~
Flora von EcutJdor (Biblioteca Botanica) Heft 116); bamboos, pp. 57-58. gress, Montreal, 19.59; Univcrsity of Toronto Press, Toronto), vol. 1,
[70J pp. 91-96. [120J
1945 ('Additamenta agrostologica," Engler botaniscM jahrbucher 74~ 22, 24. 1962 "The bambusoid embryo: a reappraisal," Am.}. BotaJrY 49, 639-641.
[1~ .
[105J

- - _.. _ .. ~ ... - _ -_._--

..
332 LITERATURE CITED LfrERATURE CITED 333

Rehder, A. Seifriz, W.
1945 "Xotes on some cultivated trees and shrubs," J Amold Arboretum 26, 1920 "Thc length of thc life cyclc of a climbing bamOOo. A striking case of
67-78. (282) sexual periodidty in Chusquea. abietifllia Griseb.," Am. J. BOlarry 7"
1949 Biblzography 0/ cultz'vated trces and skrubs U\rnold Arboretum of Harvard 83-94. [143, 273]
University, Jamaica Plain, Mass,), [282] Sen Gupta, M. L.
Rhind, D. 1939 "Early flowering of Derulrocalamus stn:ctus/' Indian Forester 65, 583-585.
1945 Thc gr(lSS$$ qf Burma (Baptist Mission Press~ Cakutta); bamboos, pp. [274]
1-26. (270) Shannan, B. C,
Riehharia) R, Ho, and J. p, Kotwal 1945 "Leaf and bud initiation in the Gramineae," Bot, Ga:::. 106, 269~289.
1940 ((Chromosome number in bamboo (DeruiTlXalanw.s sirictus)," Indian. J [62]
Agr. Sei. 10, 1033. [167,206] Shepherd, W, O. .
Rickett, H. W. 1952 "Highlights of furest grazing research in thc Southeast l " j, Forestry 50,
1958 ({So what is a taxon?P Ta:um. ;: Si-3B. [317] 280~283. [3]
Rivicre, A. and C. Shibata, K.
1879 "Les bambous," BulL Soc. Ace/im. [mI 5, 221-253, 290-322, 392-421, 1900 "Beiträge zur Wachstumsgeschichte der Bambusgewächse,'t ], Tokyo
460-478,501-526,597··645,666-721, 758--828, Preprinted (1878) as a Imp. Unw. Gall. Sei. 13,427-496. [13, 18,23,81, 123J
separate, continuously paged publication; citations in the present work Shigematsu~ Y. . ,
refer to thc 1379 edition) which contains a fcw minor corrections and 1958 "Analytical investigation of the stern fann of the lmportant SpeCl~
emendation•. [16 (Fig, 4), 19, 27,164,168,172,204,208,213 (Fig. 87), of bamboo" [in Japancse:; English summary), Bull. Fac, Agr. Unw.
215 (Fig. 88), 226, 234, 235, 239, 240 (Fig. 96), 241, 242, 250, 258, MiJazaki 3, 125-135. [42]
317] Siebold, Fr. de, and J G. Zuccarini
Rollins, Reed C. 1843 "Plantarum quas in japonia coIlegit Dr. Ph. de Siebold, Genera nova,
1965 "On tbe bases of biological c1assification," Ta_ 14, 6. [264] notis characteristicis dclineationibusque illustrata proponunt. Fase.
Roxburgh, W. primus," Abhandl. Boy". Akad. Wiss. Math,-Phys. KL 3 (3), 745-749,
1795- pL 5, [116]
1819 Plant:; IIfth< coast IIfCo,tmw.rldel (Nicol, Lendon); barnboas, vol. 1 (1795), Sprague) T. A. . .
col. 56-58; vol. 3 (1819), coL 37-38. [132, 197] 1944 In C. R. Mctcalfe et al.~ '''On the taxonomlc value of the anatomlcal
Ruprecht, F. J. structure of the vegetative organs of the dicotyledons. Art. 7. A review,"
1839 Bambuseas monographicecxponit (Aead. St. Petersburg), pp. 1-71, pl. 1-18; Proe. Li= Soe. lAnd"" 155th Session (1942-43), pt, 3, 232-235. (6J
reprimed, HBambuseae/' Atem. Acad. St. Pctersb. (ser. VI) Sci. Na!. 3 Sprengel~ K,
(pt. 1), 91-165 (1840). Citations in the present work refer to thc .'j
1825 C(Uo[i Linrwei !fJstcma ocgetabilium (cd. xvi; Dieterich! ('n>ettingen); bam-
1839 edition. [89, Z82] boas, vaL 1, pp. 132,249, vaL 2, pp. 112-113. [313]
Satow, E. Stapf, O.
1899 (The cultivation of bamboos in Japan;' Trans. Asiatic So:;, Japan 27) 1904. ~'Ön thc fruit of :.\tclocanna bambusoides) Trin) endosperrnless, viviparo us
pt. Irr, 1·~127J i-viL According to 5atow, the text of this paper consists genus of Bambuseae," T,(1JlS. Lirm. Soc. Loruitm [2> Botany] 6.401-425.
mainly of a translation of a Japanese work by Nawohito Katayama) [121, 190, 197J
Nihrm ehiku.fo (Manual of Japanese bamboo,) (1885). [247] 1904& '~Himalavan bamboos. Arundinaria falconeri and A. falcata"j Gard..
Schaffner, J, H. Ghron. 35; 304-307. [279]
1927 "'Control of sex reversal in thc tassel of Indian rornr)! Botan. Ga.z. 84, Stebbins, G. Ledyard
440··449. [6] 1956 «Cytogenetics and evolution of the grass family,>' Am. J. Botany 43,
1931 "Phylogenetie taxanomy of plaots," Quart. Rev. Bat. 9, 129-160. [305] 890-905. [5]
Schomburgk~ R. H. ; Stern, W, L. and K. L. Chambers
1841 Reisen in Cu"",", und 11m &i.tu>I',t; wahrend dcrJahrm 1835-1839, herausgege- 1960 ((The dtation of wood specimens and herbarium vouchers in anatomical
ben von O. A. Sehomburgk (Georg. Wigand, Leipzig). [42] researcht Taxon 9, 7-13. [147]
Seibert, R. J. Suessenguth1 K.
1947 BA study cf Hevea (with its economic aspects) in thc Republic of 1925 "Ceber die Blüteperiode der Bambuseen/' Flora oder Allgem. Bntan. 2tg.
Peru," AM. Missa",i Baum. Garden 34, 261-352. [61] 118, 503~535. [84, 273]
334 LITERATURE CITED LITERATURE ClTED 335
Sung, YingRhsing, cd. teca deI Cemro Nadonal de Investigaciones de Care). No. 11) 26--3L
1929 T'ier. kung k"ai wu (Commercial Press, Shanghai); a compendium cf [187J
illustrated artie1es (in Chinese) describing the arts and industries prac- Trimen, H.
ticed in China during the Ming Dyna5ty (1368-1644); "Chinese paper-
1900 Handbook qf ehe flora 0/ Gglan (Dulau, London)1 pt. V, Eriocaul(Jf/.acae-
making," voL 2, an. 13. [1J
Gramineae, ~'eontinued by Sir J. D. Hooker; barnboos, pp. 308-319;
Takenouchi, Y. pt. VI, Supplement, by A. H. G. A1ston (1931); bamboos, pp. 341-342.
1926 "On the rhizome of J.apanese bamboos U [injapanese], Trans. lilat. H:'st. [162]
Soc. Fomwsa 16, 37-46. [22 (Fig. 9)J Trinius, C. B.
1931a "Systematisch~vergleichende Morphologie und Anatomie der Vegeta- 1821 "Agrostographisehe Beyträge," in K. Sprengel) Neue entdeckungen Pflan-
tionsorgane derjapanischen Bambus-Arten/' Ta/wh Imp. Univ, (FormDsa) ;enku..de (Fleischer, Leipzig), voL 2, pp. 33-94. [197J
Fac. Set: Mem. 3, :::So. 1 (Botan.~ No, 2), pp. 1-60. Citations in the pre- 1835 "'Bambusaceas quasdam novas describit/" Mem. AcatL Sr.. Petersburg
sent work refer to thc pagination in a manuscript English translation [sero VIJ Sci. Nat. 1, 613. [282J
by B. Y. Morrison deposited in the Iibrary of the U, S. Dc,partx""ol. __ _
of Agrieulture. [7, 15 (Fig. 3), 22, 23, 31 (Fig. 13\, 32 (Hg. 14), 41 Trottert H.
(Fig. 20), 6'3 (Fig. 34), 66 (Hg. 37), 70 (Fig. 39), 71 (Fig. 40), 72, 73 1922 "Development of bamboos from natural sbedlings (Dmdroc:alamu.r
(Hg. 41), 78,80 (Fig. 44)] strictus)," Im/iM Forester 18, 531-536. [172J
1931b "Morphologische und entwicklungsmechanische Untersuchungen Troup, R. S.
bei japanischen Bambus-Artent Mem. ColL Sei. Kyoto Imp. Univ. 1921 Tlu silvicu1iure 0/ Indian trces (Clarendon Press, Oxford, England); bam-
(5er. B) 6, 109-160. [45, 49] boos) vol. 2, p. 725; vol. 3, 997-1013. Some illustrations and passages
1932 Take 110 ken.lf;v1'1 [Bamboo studies; in Japancsc; with bamboo names in of text arc reproduced with the publisher~s permission. (128, 129 (Fig.
LatinJ (To1.7o), Citations in thc present work rcfer to thc pagination
in a manuscript EngHsh translation bcgun by F. A. McClure and Kam
,. I
58), 130, 131 (Fig. 60), 132, 169, 189 (Fig. 82), 190, 196, 197, 205,
236, 274]
Hok..chau and completed by Sabum Katsura in coUaboration with Tsuboi, Isukc
R, A. Young and Mary &hoff, and deposited in the Llbrary of thc 1913 Jikkcn chikwin zosei-ho [in Japanese: Experimental mcthod ofestablish-
U.S" Department of Agriculture. Some illustrations and passages oftext ing bamboo groves] (Forestry Association, Gifu PrefectuTe, Japan).
(in translation) are reprodueed with the publlshei's permission. f7, 22 Q:.totations uscd herein are drawn frorn a manuscript translation of
(Fig. 9), 23, 29 (Fig. 11), 43 (Fig. 21), 45, 77 (Fig. 42), 78 (Fig. 43), pp, 13-23 of theJapanese text by Saburo Katsura, with the collabora-
127, 134] don ofR. A. Young (1936) under the tide I>Methods ofbamboo propa-
Takhtajan, A. J. [a. W.: Takhtadzhian] i gation," deposited in the library of the.U.S. Department of Agricul-
1959 EssffYs on thc evoi.utionary morphology of pLa'f'.ts (Amer. Inst. Biol. Sei., J ture. [226, 258] .
WashingtDn) D.C.), An English translation of the Russian original Udagawa, T" T. Mizuno} and M. Seki
(Leningrad University, Lcningrad, 1954) by Olga Hess, cd, by G. L. 1958 «A fie1d experiment on some rat-poisons)' [in Japanese; English sum-
"1.
Stebbins. [6, 105J mary], Japan. Min. Ag:". and Fl7festry (Forest E.pt. Sta. BuH. l'io. 105),
Thimann) K. V. pp, 1-10. [86J
1954 "Correlatlons of growth by humoral infiucnces H (Harvard University, Uada, K.
BiologicaJ Laboratories, Cambridge 38, Mass.); reproduced from 1960 Studüs on tlu physiology qf bambtw~ w~·th rifcmlCe trJ practical application
Rapports et Communitations du VlIr Cor.gres Interno.tionaL dc Botardque~ Pan's~ (Bull. Kyoto Univ. For..ts, l'io. 30). [25, 263]
1954, Sec. 2, pp. 114c-128. [6] U.S. President)s Commission on Increased Industrial Uze of Agricu1tural Products
Thimann, K V" and J, Bchn1."'C-Rogers 1957 Interim rep01t to The CtmgteSS~ pursuant ,0 Puhlic Lew 540 (84th Congress)
1950 The use 0/ auxins in (he rooting of woody C'.ätmgs (Maria Moors Cabot Washington, D.C.), x, 6, 10,69, and 14 p. [202]
Foundation Pub1ication No. 1; Harvard University) Harvard Forest,
j i Usui 1 H.
Petersham, Mass.). [259, 262J 1957a "Study of thc embryo on Sasa nipponica and Pleioblastus (;hino~" J. Japan.
Thomas, G. S.
"Bamboos," J. Roy. HI7ft. So,. 82, 247-255. [210]
, Botany 32, 193-200. [120, 138, 140 (Fig. 66)]
1957 195ib '~Morphological studies on the prophyll of Japanese bamboos." Bo/an.
Thompson, J. McL. Mag. 1Okyo 70, 223-227. [53, 54 (Fig. 28)]
1944 "'Tov.:ards a modern ph)'"Siological interpretation of tloweringl " Proc. Valt'er, 0" T, Krasnosel'skaya1 N. :Maksimov, and V. Mal'chevskiy
Linn. SO<. LOIJdim. 156th Session (1913-#), pt. 2, 46-69. [6, 105, 107] 1910 "Uebcr den Blauzäuregehalt der Bambusschösxlinge," Bulletin du.
Triana, J V. Departement de t'Agricul1ure au.x Indes .Nicrlandaiscs 42, 1-4. This paper is
1950 HAlgunos metodos de propagaeion y medidas de crecimiento en las a brief7 pre1iminary eommunication ("vorläufige Mitteilung"), in
espccies Guadua angustifoUa. y Bambusa vulgads~ Bolet:"n lnformativa (Biblio-
J) German., of results more e.'<-tensively elaborated in the 1911 paper by

-- ~------""""""'--
336 LITERATURE ClTED LITERATURE CITED 337

the same authoTS, in Russian. The full name of rois apparently !'are 1948 Bw"nboo cuiturc and utilization in l'uerto R':co (Puerto Rieo Fed, Expt.
publication is taken from its tide page. f3] Sta. Gir., No. 29). [204J
1911 ["The hydroeyanic acid eontent of bamboo, and its distribution"; in
Russian; scientific names in LatinJ~ Bull. Imp. Acad. St, Ptkrsbwg (1911), White, D. G.) and N. F. Childers
1945 HBamboo for controlling soil erosion," J. Am. Soc. Agron. 37, 839-847.
397-426. [3J
[79]
Van Overbeek, J.
1944 "Growth-regulating suootances in plants/' Ann. Rev. Biochem. 13, Whitehead, A. N.
1948 Seienee and philosoph) (Philosophical Library, !\ew York), p. 131. Pas~
631-666. [263J sage reproduced with the publisher's pennission. (81
[1945] Pen;onal communication. [238,257 (Fig. 99)J
Wi1son,G. L.
Vavilo-v, N. 1. 1942 "'Thc te10me theory aud thc origin of tbc stamen," Am. J Botany 2~
1940 HThe nevi systematics of cultivated plants/ l in J. Huxley, The new
759-764. [105]
.rystematics (Oxford University Press, London), Passage from p. 565
quoted with the publisher's permissIon. [6, 264] Yoshida, N., and S. Ikejiri
1950 üBamboo~sprouts extracts as growth-prompting (sie! substances for
Velasquez, G. T., and J. K. Santo, miero-organisms. Studies on the gro;-rch-proroonng ~ubstances for
1931 "Anatomieal study of five Philippinc bamboos/~ BulL Nat. App!. Sc:'. micro-organisms. I" [in Japancsc; Enghsh summary]~ SJHkoku Acta Med.
Univ. Philippincs 1, 281-318. [49]
1, 25-28. [2J
Venkatraman) T. S.
1937 "Sugarcane-bamboo hybrid'," Indian] Agr. Sei. 7, 513-514. [5] Young, R. A. 360
1946 I'Bamboos in Amcncan Hortieu1ture (V)," NaCl. Hort. Mag, 25~ .
Venning, Frank D. (186]
Personal cotnmumca tion [45] [19601 Amcricatl nurseries h'sting bam.boos (mimeographed, issued irregularly;
Waley,A. New Crops Research Brauch) Crops Resea:ch Division, A~cultural
1929 A hundrcd and stVetily CMnesc poems (Knopf, New York), p. 214. The Research Service, U.S. Department of Agriculture) Beltsvllle~ Mary-
first ten Unes cf Waley's translation of a Chinese poem by Po Chu-i~ land). [292J
entitled '~Eating bamboo shoots, H are reproduced with the publisher's
Young, R. A. and J. R. Haun . . ... .
perrn.ission. [xvi~ 1] .1961 Bamboo in the Unitcd Statcs: descnptwn, cu/ture and utzlt.Zatum~ "...:th key to
Wardlaw, C. W. genera by F. A. McGlure (Handbook No. 193; U.S. Department of
1956 "The floral meristem as areaction system," Proc. Ro)al Soc, Edinburgh Agriculrure, Washington, D.G.). [289]
[B, Bio1ogy] 66, 394-408. [6, 105] Zimmcrrnan, W.
1959 "Methods in plant morphogenesis," J. Linn. Soc. Lorukm 56~ 154-160. 1961 "Phylogenetic shifting of organs, tissues and phases in Pteridophytes."
[6J . CanadiM}. Boto.'}Y 39) 1547-53. The author discusses other groups of
1961 "Morphology.'~ in A, M. ?v1acleod and L. S. Cobley, eds., Contemporary plants besides the Pteridophytes. [6, 106]
botanical t"eught (Oliver and Boyd, London). [105, 1431
Watson l E. V.
1943 (IThe dynamic approach to plant structure and its relation to modern
taxonomie botany," Biol. Rcv. (Gr. Bri"'in) 18, 65-77. [6, 103, 105]
[\Vebster, N.J
1959 Webster's new intcmatWnai dict~'onary of the English language (ed. 2) una-
bridged; Merriam, Springfie1d~ Mass,), In the 1959 printing here cited}
and used, the section entitled "Addenda,n expanded to ineorporate
definitions of manl' words not found in earlier printings, was copy-
righted in 1954. [302 cl passim]
Wendland, J. G.
1810 Celleetie plant.rum (Hahn, Hannover), voL 2,p. 26, pL 47. [82, 163J
V>'hite, D. G.
1947a ~'Longevltl' of bamboo seed under different storage eondidons,'1 Troft.
Ag<. 24) 51-53. Citations in the present work refer tc thc reprint
repaged, 1-4. [203)
1947b HPropagation cf ba.mboo by branch cutungs,)I Proc. Am. Soc. Hort. &i.
50, 392-394. [242, 243 (Fig. 7)]
Index of Scientific Names

Par each scientific na.rne mmtioned in the text, the 2..uthorlty is given in th;s index. :Many
of thc bamboos listed are burclened with an extensive synonym)". Thc indusion of synonyms
here is restricted to those essential tO the purpOiles of thi.s treathe. Thc word "see" cr
\<equals" follov>"S an obsolete name (one relegated lO synonym)'), ane: direets attention to thc
name currently adopted in its stead. ':'See also>1 indicates cases where both the obsolete
name and the accepted name rer a gi:ven speeies appear in thc text.

Arth.rostylitir.mt Ruprecht, 49, 53, 112, 288, AtU1tdinana japonica Siebold and Zueearini.,
297 208. See Pseu.d.osastt}aponi<:a (Siebold and
Anhros:.yNdium ~apillifolium Grisebach. 58, 285 Zuccarini.) Makino
Arthrostyb,äium cubcnsc Ruprecht. 58. 101, Anmdinaria niJdwctlSis Nakai, 136-137 '(Fig.
109, 117 64)
Arthrostylidiwn Mwumit:l1m Parodi, 55-59 Arundirurria prainii (Ga.'".l1ble) Gamble, 96,103
(Pi@' 31-33) A'I'UlUiinaria pu,..,1.ita (Mitford) :Mitford, 268
ArtJl'tos~lidium multüpic(h'Um, Pilger, 48 Anmdinaria pu.tilla A. Chevalier and A. Camus,
ArtJrrosty!.iditlJn sdwmlw:rgkii (Iknnett) Munro, 26-27 (Fig, 10), 30 (Fig. 12), 32, 42
42, 108 Arutldinaria PYl,maca (Miquel) Aseherson anc
Arthwstylidittm .su.bptctir.a.,tum Kuntze, 102 Gracbncr, 268
Anmdinaria Michaux, 13, 31, 54 (Fig. 28), AnwIinaria sinwnit' (Carriere) A. and C.
67, 81, 83. Sg, 110, 119, 123, 124. 125, Riviere, 28, 43 (Fig. 21), 55 (Fig. 29), 77
209,291, 302, 308 (Fig, <2), 102, 13B, 139 (Fig. 65), 208,
A1lUldinaria amabilis McClure, 21 (Fig. 8), 39 268,292
(Fig. 19),51, 149, 150--;57 (Figs, 67-70)) Arwzdinan'a simrmii var. Qaritgata Hooker f.,
227-228 (Fig. 92), 241, 268, 286, 292 55 (Fig, 29), 269
Arwzdi"aria amplissi'7lfJ ~ees, 102 Arundir:ana tff:ta (Walter) Muhlenberg, 13,
Arwzdif"'.aria fJuricoma Mirf-ord, 268 23,25, 2';'·2B (Fig, 10), 34, 86, lOl, 125,
A7U11d~'nW'ia ckiJw (Franchct and Savatier) 138, 204, 3! 0
Makino, 120 Anmtiinarül tccta vax. deciduo. Beadle, 55
Anmdinana dnlicha'l'l.lAa Keng, 103, 301 (Fig. 29)'
A1"UfItllnaria foltaf.a Nees, 20S. St;:e Ch.i.tnorw- Arun.dinaria va,;:ar.s Gamble 268
bambusafalcakJ (Nee!l) Nakai Arnndinaria varicga4: (Siebold) !>.1akino, 70
A1'IOI.din.a:nafolcata var, glonuraJaGamblc, 271- (Fig. 39), 82, 208, 269
See ChimorwbmrJJlJsa fauata (Nees) Nakai Arur.dinaria vancgata var. viridis f. major
Anmdinariafml1flct (Van Houttc) A. and C. Makino, 63 (Fig. 34) - .
Riviere, 208. See Anmtll""ana varicgaf.a A'tWUlinar:L1 viridi~striala (Siebold) Makino,
(Siebold) Makino :292:. See Arwulmaria w..rno1'/l.a Mitford
Anmdinan-a gi;antca (WalteT; Muhlenberg, AWo(lslacnys capitat,a (Hooker) Bentham c:x
13,23,49, m Benth:am and Hooker f., 89, 101, 312
Aru.'I'l.dinar:n hintisii semu aueu.. jap. (non Auaclocarjxt Franchet, 117
Munro), 13 Autol/.tmw qucr.c Goudot, 101, 109,300

..... _ - - -
340 DIDEX OF SCIENTIFIC NAMES INDEX OF SCIENTIFIC NAMES 341

Bambusa Retzius corr. Schrcber, 22 (Fig. 9\ Bambusa scriptoria sensu A. and C. Riviere Chimorwbamlr'..stl iwakaiana (Mu:nro) Nakai, Elytrostac!.ys McClure, 98, 209, 290
53, 60, 64 (Fig, 35), 73, 74, 81, 89, 98, (non Denn,stedt), 208. See Bambusa 271 272
j ElJtwstad.ys cllWlgera McClure, 118--119
:08) 116, 119, 123. 124, 209, 241, 270, multipJcx var. rim'aC()Tl1.m R. Maire Ckim(maoambusa rr..armcmr. (Mitford) Makino. (Fig. 55), 287
280, 288, 290, 302 BamlJu.sa shimadrn' Hayata, 70 (Fig. 39). See 118 (ng. 55), 204:
Bambustl argtl'llCa Hort. ex A. aod C. Riviere, a.lso Bambusa 1T'.ulliPlcx var. skim(ldai ChimrmobalrJus a qu.adrangularis(Fen:.t;i) Makino,
208. See Bambusa multiplex (Loureiro) Hayata 46. 77 (Fig. 42), 79, 00 (Fig. 44), 272, Gigtmrochloa Munro, 53, 108, 114, 115, 116,
Raeusehel Bamlmsa spirwsa Roxburgh, 208 283, 284, 315 209,215,235,241,287 1 290,307
Bambusa anJJ7.dmacra Retziu~> 48, 60, 83, 85, BtJ:mbusa slrnost.ach)"a Hackd, 73 (Fig. 41),81 ChusquttJ Kunth. 30. 33, 48 (Fig. 25), 49, Gigtmioehtoa al/:1c-citiata (Munro) Kur", 117
114, 117, 126, 127, 203,204) 206, 235) Eambusa str:'ela Roxburgh. 208, See Drnc'rr;- 53, SB) 67, 112) 114, 2OB~ 284, 287, 299, GiganlachlfJa apus (Schuhes) Kurz) 58) 219
246, 280, 288 r:a1amus slrictw (Roxburgh) Kees 304 (Fig. 89), 220, 224, 225, 232, 234, 243,
Bambusa buchgaM Munro, 21 (Fig. 8). See Bambusa strict(l var, argrntetl A. and C. Chu.tlJuea abutljolitJ Grise:bach, 267, 272, 273 244, 245, 249 (Fig. 97 ~ 5)
also Si:n.ocalam!lS buche.ya:nus (Munro) Rlviere, 168. See DnuirQc(llamus strir:tus Cw..csqlJC(1 fmdkn" Munro, 21, 26 (Fig, 10), Gigantothbamaxima Kurz, sensu Holttum, 117
McClure 2R, 31, 33 (ng. 15), 34, 36 Giganloc!rl(lf; nipo-ciliata (Buse) K\.l.rz, 118
(Roxburgh) :Kees, 16$
Bambusa blu.mcana Schultes f,~ 118 (Fig. Bamfnt.sa textitis McClure, 39 {Fi:g. 19). Ckusqw:a pinifclia (Nees) Nccs t n (Fig.55), 119
Bamhusa copciaNlii Ga.-nble, 84, 118 (Fig. 49, 51, S3, 65 (Fig. 36), 211, 220, 232, Cku.tqw:tJ pitticri Haclid, 49, 79 Gigtmior:lll?ß tJCrticiUat.a (WUldenow) Munro.
119.267, 269, 270 233, 243, 245, 24$-249 (Fig. 97), 250, ChlCiqura ~'(arule1t1 Kunth, 27 79, 149. 172-179 (Figs. 75, 76), 242
BamblJ.'ill dissim:ula0r McClure. 61 ChlCifJuea simp1iciflvro Munro, 60, 69 Giazivpr.yton Franchet) 43, 51, 88~ 298
251
Bamoosa dnJicnocladtt Hayata) 71 (Fig. 40) Bam!Jus(l f'.<lda Roxburgh, 58) 65 (l"ig, 36), Clrusquca Sp,; 43 (Fig, 25), 6S (ng, 38) Gla....-wpkyton mirabile F:ranchet, 90~ 100, 105
Bambusa cutuldoitles McClure. 85 79,217,219 (Fig. 89), 220, 229, 230-231 Gramineac Necs, 5, 107 ~ ZOO, 282
Ba:mbusa gracilis Hort. ex A, and C. (Fig. 93), 232, 236. 243, 245, 24$-249 Gralam'a Ba~ans:a, 105,298
200; equals Ar'.L'Viinaria gra;:ilir (Fig. 97), 251, 252 (Fig. 98)
Gralarna m!mlana Ruprecht, 100
Btrmbusa nookai (Munro) A and C. Rivi6re, Barn.busa tuldaides Munro, 14 (Fig. 2). 26-27 Gr-:s!.an:ia spp" 105
Dm.tfrm:alamus l'\ees 22 (Fig, 9;, 53, 73, 74 r
j
Guadua Kunth, 27, 60, 209, 241, 290
208. See Dendracalarr.us }«JIJf.eri Munro (l:'ig. 10), 32, 40, 65 (Fig. 36), 211, 214, 108, 114, 119, 124,209, 241, 270, 290
Bambu.sa. lir..:ata Munre. 83, 270 215 (Hg. aa), 216, 220, 232, 243, 245, Guadua aculr:ata Ruprecht cx Fourn~er, 4,
Dmdrocala..vw.s affiYa's Rendle; 39 (FiE;, 19).
Bambusa !tmgispicuIata Gamble ex Brandis. 251,255.271,292 118-119 (Fig. 55)
See SinocatarTOO' qffinis (Ren dIe) McClure
118 (Fig. 55), 203, 220, 222 90), Bambusa vPZlrnoset McClure, 232, 233, 234, Guadua ampltxifoliaJ. and C, Presl, 60, 67, 86
Dmaracalamus aspcr (Schultes) Backer, 118
223 (Hg. 91), 225, 243 1 245; 252 244, 245, 259, 293 Otu:u1WJ angustifl)/ia Kunth, 18 (Fig, 6), 79,
(Fig. 55), 243, 245 179-187 (Fi!!'. 77-80), 232, 234, 245,
(Fig. 98). 254, 260 Bamlntsa vuigaris Schrader ef(: Wendland, 4, DmarlXa.lamus gigantt"W 1,1 unro, 46, 72
Bam!Jusa rr.acroculmis A. and C. Rivicre. 19, 16; 17 (Flg. 5), 20 (Fig. 7), 27, 36, 46, 246,247, 250, 274
Dmdroca.lrunus hamiltortii Munro; 115, 273
208, 215 (Fig. 88), 235, 240 (Fig. 96) Guadua capitata (Trinius) Munro, 89
58,79,83,90,96,115. 116, 149~ 157-164 DendtlJr:a.lamI!.$Mmiltrmii var. :dulis Munro. 273
Bambusa mtJlingt:nJu MeClure, 60 Guadua perLigu(ala Pilgcr, 70
(Figs, 72), 200, 234, 235, 238, 242, Dendrocaiamus nookcn Munro, 208, 235
Guadu.a spinosa (Sw.allen) McClure. 304
Bambusa mu1tipl$x (Loureiro) Raeuschcl, 43 245, (Fig. 99), 259, 285, 287, 293 Dendroca/amus latifiorus :vfunro, 41 (Fig, 20)~
Bambusa vulgaris va:, vi.lla/a A. 2.."1d C. Riviere, Guadua lusmannii Pilger, 90
(Fig. 21), 77 (Fig. 42). 81,95 (Fig. 47), 71, 315. See abo Sinocalamus latifiarus Guadua trinii (Nees) Nees ex Ruprecht, 84,
96 (Fig. 48), 103, 104, 106> 114, 117> 118 200, 236, 237 (Fig. 94), 238 (Fig. 95), (Munro) McCl~ 250,267, 275
(~'ig.
270, 292
55), 127 (Fig. 57), 245, 247, 251,

Bambusa multiplex cv. <. Alphonse Karr: 96; 292

242, 245, 260
B:.rmbusa 't'Ulgarii cv. '\,llamin' stat. nov., 162
(Fig. 72)
1 Dmdrocaltzmus l~ngispalkus (Kurz) Kurz, 236
Dmdroca.lamus mcmbrl1naCr:US Munro, 34, 35
(Fig. 16), 203
Guaduttla Frnnchet, 51, 286

Bambu.ro 1llllltiplex cv. 'Fernleaf/ 292 Barnbusa ? Wamtn Brandill ex Camus, 162. Dmdrocalamus sikJcimensis Gamble eX Oliver, Indocalamus Nakai, 22 (Fig, 9), 31, 227
Bamhusa multipkx cV. <Silverstrlpe,' 292 See Bamlntsa vulgari.s cv. ,\\ramin' ' 98 (Fig. 49), 128, 130 (Fig. 59) lr.t1wtl.lrJmus :nHroka)lammsis (Hayn.ta) NakaiJ
Ba.mbuJa multipLex cv. "Stripestem Fern~ Bambl.:sa.ccae Link, 280, 282
lea.f: 292 Bambu,o;e'Ac Nees (sensu lat.; cf. Parodi I Dr:ndrocalamus strictus (Roxburgh) Nces) 18.
85, 96, 116, 118-119 (Fig. 55), 128, 131
sensu Takenouchi, 22 (Fig. 9~. Sec Yu~
shania niitak'!Jamtnsi.r (Hayata; Keng f.
Bamhusa mul1ipkx var. n't:iereonun R, :vfaire, :9(1),7, 117.282 (Fig. 60), 164-172 (Figs. 73, 74), 203, IndacalA.'1Ius sinicus (Hance) Nakai, 26 (Fig, 10).
92 (Fig. 45), 245, 247, 231, 292 Bambuooideae Ascherson and Graebner, 5, 204, 205, 206, 200, 211, 216, 229-232 28, 34, 36, 102) 109
Ba7rJJlt$tJ mll.l!ipltx VaT • .skimrul.c.i (Hayata) 7, 93, 281, 232, 288 (Fig. 93), 235. 247, 251, 260, 273, 285, lr.t1octJ1a:mus wigktiant1.S (Nees) Nakai, 83
Sasa1ci 70 (Fig. 39). See also B:.rmb1J.sa
j
BClIia tonkinensis Balansa, 96, 103 293 !r.r.WStl..ltJ gibbosa (MeClure) McClurc, 71
snimadai Hayata ,; Dmdroc(llmnus strictus va!". prtJ.irr.iar.a Gamble,
BmrJ;l!.${l nutaru Wallich ei{ Munro, 112 167
BambustJ I)ldhamii Munro, 292. See also Cephalosta.e'fJu.m Munro, 119, 241, 290 Dtndrrxalamus strir:tus Val'. st:Ticr..s (!viunm) l.ingrumia ch:ungii (tvicClurej McClurc, 46,
Sinotatamus oldhamii (Munro) McClure Gcphalastae'fJum capilatum Munro, 288 Gamb:e,168 47 (Hg. 24),4il (Fig. 25),49,50 (Fig. 26),
Bambusa packirutlSis Hayatu. 26-27 (Fig. 10), Ccphalostar:'fJum pcrgmdlr Mumo, 232, 234, Dcm1ror:klo(l distans Pa.r'..:inson, 118-119 (Fig. 51, 60. 67
29 (Fig. 11), 32, 66 (Fig, 37; 243, 245 55;
Ba/lIousa pC'fIJariabz'lis McClure, 65 (Fig, 36) Ccphalo~'ror:fl:Jum sp., 118 (Fig. 55) _ Din?r:kloa Buse, 58, 108, 120
Bambusa fMlym01pka Munro, 232, 2.35, 243, Chimorwbamlntsa Makino, 74, 79, 81, 208~ 291 Dz'lIcchloa ar,dmwmica KUl"z, 283 MdacaJemus Bentham, 120
245 Ckimonobarnbusa falcata (Nccs) Nakai, 67, Dinochloa setmdcns auctt. (non Kuntzc), 39 Mdacalemus ccmpactifit;ro.s (Kurz) Bcnth:un,
Bambusa rutila McClure, 56 (Fig. 30), 61 128, 129 (Fig. 58), 208, 271 (Fig. 19), 115 60,67,69, Hll-119 (Fig. 55), 120, S04
342 INDEX OF SOIENTIFIC NAMES J I"-'DEX OF SOlENTIFIO NAMES 343

Mclocanna Trinius, 45. 48, 53, 120, 289, 302 36,37 (Fig. 17), 4n, 103, 104 (Fig. 51), Racemobamhos Holttum, 108 SemiarrnvJiMrio.fastuosa (MarHac) Makino, 36,
Mclocanna baccifcra (Roxburgh) Kurz, 18, 126. 278, 283. 293 38 (Fig. 18),40,41 (Fig. 20), 55 (Fig. 29),
26-27 (Fig. 10), 34, 36, 53, 60, 118-119 Phyllimachys bamfJu.soirks cv. 'CastiUon; 35 67, 77 (Fig, 42), 255, 293
(Fig. 55), 121, 124, 125, 132, 133 (F'g. 61), (Fig. 16), 46, 293 Sase Makino and Shibata. 25. 31, 51, 54: Stmianma'inaria viridis (Makino) M:aki.no,49,
134 (Fig. 6Z)t 149, 187-197 (Fi&", 81-85), Phyllos4U;};yl hambusoidcs var, ma:rtiacca (Mit~ (Fig. 28), 57, 73, 74, 81, 110, 209, 227, 53 (Fig. 34)
216,218-219 (Fig. 89), 275, 28i3, 298, 301 ford) Makino, 63 (Fig. 34) 291 Shibataea lYfakino, 31, 44, 54 (Fig. 28), 73,
}"fcrostathJ's Sprengel, ~9) 53, 56 (Fig. 31), PhylliJstacl:Ys d:ulcis McClure, 55 (Fig. 29),293 Sasa bortalis (Hackel) Makino and Shibata. 81, 124,209,291
58, 101,297, 313, 315 pr.yLlostacr,ys ctcgans McClure, 52 ~Fig, 27) SI Shihataca kwnasasa (Zollinger) Makino, 26
MCrflstathys f.s:ulosa DocH, 276 Phytiostad!JJ jlcxuosa A and C. Rlviere) 208, Sasa kuriknsuvar, ntlrulosa (Makino) Makino, (Fig_ 10), 28, 32 (Fig.. 14), 34, 36, 55
Mcrostachys ftauci/fMC. Swallen, 101 293 63 (Fig. 34) (Fig. 29;, 70 (Fig. 39), 77 (Fig, 42), 293
Mcrostachys rctrorsa :McClure, 101 Ph.yJJosIat:~s flrmosana Hayata, 70 (FiS, 39) Sasa kmgiligulata McClure, 4:5 SiMnma'i7l.cria Nakai, 200, 290
Mcroswhys sp., 57, 58, 86 P~tliJ.ttacllY:" ht!('fOdadn n Oliver ex Hooker) Sa,sa nclrutosa (Makino and Shibata)' Ohkl, Sirummdinaria nitida (Mitford) Nak.ai 26-27 j

Myt1ctladus Swalien, 43, 285 117 118-119 (Fig. 55) (Fig, 10), 34
Phyllost(L;h.ys makinGt: Hayata, 46, 73 (Fig. 4n Sasa nipponica (Makino) Makino and Shibata, Si7l.ohamlr.J.Sa ß.-hlino, 54 (Fig. 28), 73, 74, 209,
77 (Fig.42) 13, 120, 138, 140 (Fig. 66) 291
Phyllosuu:hys rruy<ri l\tfeClurc, 293 Sasa pai't]'l.().ta (Mar1iac) Nakai. 13,39 (Fig. 19), Si....obambusa iQorsiJ: (Makino) Makino, 4, 43
Naslus Gmelin, 112, 119
Phyllos!achys 1lll'tis A. and C. Riviere, 208. 55 (Fig. 29~, 293 (F.lg. 21;, 48. 67,77 (Fig.42), !18-119
,Vast'Us ckganllssimus (Hasskarl} Holttum.
See Phyllostacnys uiridis (Young) MeClure SaJa patliculata (Schmidt) Makino and Shi~ (Fig. 55), 315
118-119 (Fig. 55), 276 Sinoca!cmus McClure, 241, 270
,Phyaosttur.ys mieis sensu KawarnurJ. t 104 bata, See following entry, as tO type
NClJ.rolepis Mcisnet, 51, 283, 285, 297, 314 variety Sinocalamus Cfifi.... is (Rendie) McClurc, 39
NeoJwuzcauo. A, Camus, 115. Sec Scki::cslachywr. (Fig. 51), equals
Phylloslachys pu-bcscrns Mazcl ex Houzeau de Sasa paniculaiP. V3.t\ panicu(ataJ 77 (Fig, 42) (Fig. 19)
Nees Sinorulamus heccJu;ya.,us (Munro) McClure, 39
Lehaie Sasa pygmaca (Miquel) E. Q, Camus. 293;
P':Jtlostachys nitlutaria Munro, 44 (F:ig. 22), 'cquals Aruna'inaria pygrr.aea (Miquel) (Fig. 19),242. See also Bambusa fx(cMyo.na
48, 67, 69, 99 (Fig. 50), 100, 104, 110 Ascherroo aod Graebner Munro '
Ochlandta Thwa.ites, 114, 116, 119, 120. 287, (Fig. 52), 111 (Hg. 53), 277, 236, 314 Sasa ttsscliata (Munro) Makino a.>1d Shibata., Sinocall1:Qf'dS copdandii (Gamble) Rai2;"ada, See
290,302 Phyllostachys nigra (Loddlges) Munro, 39 82, 278, 293 ßamhusa copda7l.a'i.:" Gamblc
Ochkmdra hcdt1ofl'.d Gamble. 116 (Fig. 19), 75, 80 ~Fig. 44), 123, 208, 293 Sasa. !s',JbtJiana Makino, 70 (Fig, 39) Sinocaü::mus latifiorur (Munro) McClurc, 77
OchlmuJra smptona (Dennstedt} C. E. C. Pl;yUosuunys nigra cv. <Henon,' 104 (Fig. 51)~ Sasll vn'/chii (Carriere) Rehder, 73 (Fig. 41), (Fig. 42). See also Dm.drocalamus lattJiffrUS
F"Cher, 276 278, 293 78 (Fig. 43), 93, 94 (Fig. 46), 95 (Fig. 47), Munro
OchltJndra. snigm'f. Gamble, 116 P/;JliostacJ!Js rnvia McClure, 293 97 (Fig. 48), 101, 102, 104, 106 Sirlocaltzmus oldhamii (Munro) McClure, 232,
Och/andra str:duta Thwaitcs, 83, 116, 276, Phyllostachys pubcsccr.s Mazet ex Houzeau de Sasaclla Makino, 74, 287 242, 243, 245, 270. See also Bam.ousa old-
307 Leh.ic, 13,45 (Fig. 23), 36, 37 (Fig. 17), Sasam.:npM Nakai, 289, 291 Jwmii Munre
Och.landm travancorit:(! (BeddQme) Bentham ex 46, 70 (Fig. 39), 7! (Fig. 40), 75, 104 Schi:mstacfr;um Nees, 45, 46, 53, 89, 90, 98, Strcptaduuta Schrader ex Nees, 108,288,297,
Gamblc; 112, 114, 115, 118-119 (Fig, 55), (Fig. 51),118-119 (Fig. 55), 239, 278, 293 107, 108, 114, 115, 116, H9, 124:, 241. 307
197-201 (Fig. 86), 276 Phyllostachys quiliai A. and C. Riviere, 2OS, 287. 290 Streptochaaeae Hubbard, 283
OcAlandra travanconca. VM. }u'rslJta Gamblc, 199 See Phyl/osLaehys bamfrJSGides Siebold and Schi:!ostJ1chyum aa;.tiflorum Munre, 134
Olyreae Pilger, 288 Zuccarini Schi;mtachyum hlu:mü Nees) 108
Oreohambus htd::wakHi K. Schumann, 96 Phylloslachys vwlnsce71S A. and C. Rivie~ 208 SchiuS'.cchywn lrrachycladum. Kurz, 83, 116 Thamnocalamus Munro, 290
Oxylr.rumthcra Munro, 115, 116, 209, 287 Plryll(utathys uin.tli-glauccscms (Carriere) A .. SdU'::(ISliU).:Jttm gracilc (Munro) Hohtum, 83, ThmTmocntamus jaltoruri Hocker f, cx MUDro,
Oxytman!herll al?Ys:n'tdcll (A. Richard) Munto, and C, Riviere, 208, 213 (Fig. 87), 226, 113-119 (Fig. 55), 135 (Fig. 63) 278,290
llfi-.119 (Fig, 55), 217 274,276
j 240 (Fig. 96) Schizwtat;hyum grar.dc Ridley, 83 Tho.m,ocalamus spathiftaros (Trinius) )Aunro,
PhyllosttMnys uiridis (Young) McClure, 16 Sch:i::ostachyum luJü'IarMr.se ~.tferrill ex McCiure, 101, 108
(Fig. 4), 19, 46, 125, 206, 293 27, 39 (F;g. 19) Th:JlfsostatfErs Gamble, 89, 108
Pari.eneae Hubbard, 288 Pl>:ylwstachYJ .riridit CV. 'Ro bert Ywagt' 293 Schi:ost.achy'Um lima (Blanco) Merrill, 100, 117 ThyrsostiKftys oliven Gamb1e. 236, 279
Phareae Stapf, 288 Plcioblastus Nakai. 22 (Fig. 9), 28, 53, 54 Schi.::ost.achyum kmgispiculatum Kurz, 83
Phy[[ostad-,:!s S:lcböld and Zuccarini, 13, 16, (Fig. 28), 73) 74, 289; 291; equaJs Aru."l~ Schizcsuuhyum terminale Holtturn, 96
22 (Fig. 9), 25, 28 (Fig. 10), 42, 43, 46, dinaria M.i:chaux Sch:izoslaiJvum sp" 60 YusJ..allia niitakayamcnsis (Hayata) Kcng f., 22
~9j 51, 54 (Fig. 28), 67, 731 "74, 81) 83, POIl ln.:J!Josa Linnaeus, 250 ScmifJ'fliJIdinaria MakinQ, 54 (Fig. 28), 73, 74, (Fig. 9), 26-27 (Fig. 10), 31 (Fig. 13),34,
99,104,115,116,123.124,209,227.241, P(!t1 pratmsis Linnaeus. 93 209,291 36. 40, 77 (Fig. 43)
277, 2M, 291) 311 Pscudosasa Makino, 28, 31, 74, 118-119 (ytg,
P'rlllostachys arcana MeClurc, 45, 51 55), 287, 291
Phylbutru:hys aurCil A and C, Riviere, 104 Pscu.dosasa jaJmlica ~Siebold and Zueearini)
(Fig. 51), 208, 278, 293 Makino, 13,66 <Fig. 37), 72, 77 (Fig. 42),
PlryllGstacl!fs cwtGsulcala McClure) 293 115, 118~1l9 (Fig, 55), 293. Sec also
Phy1tostac~s bambusoidts Sicbold and Zuc· Arundinariajaponica Siebold and Zuecarinl
earini, 15 (Fig. 3), 26 (Fig. 10), 28, 34, Puclia Frnnchet, 51, 117
 I
•

Index of Subjects

Owjng to thc abundanee of figures related to cerwn index-ed subjects~ di:rcct refercnce
to illustrations is omitted in thc interest of economy and simplicity. For this rtason, same of
the page refereru:.es indicate illustrations onl)'; athers indicate text onIy, or hath pertinent
text and figure(s).

Brunboo as plant and reproductive struetulCS, 103-105

a"{cs, segmented, 11 hybridiwtion with 5ugarcane, 5
basic frame, 11-12 life eyc1e vs. Bowering eycle. 84•.267
eharacters: bambusoid) 3, 112~113; n:cog~ matunty, vegetative vs, sexual, ISO
ni.tlon, 16, 52, 6<;-65, 75, 81,87; ta.."o- meristem: definition, 307; fod of, 91,103,
nomic, 12, 16, 23, 50, 52. 64-65, 81~ 123, HO, 256
120, 282-288 morphogcnesis~ 3, 11, 30B
chromosome numbers, 1f)J. 199 mutations, 206
clump: caespitose, 298; diffu.o;e, 301; habit, naturalization of emtics. 4:
13, 34, 37, 283 nodes as foci of tissue differentiation, 12,
cold-hardiness, 72 123, 140
cyeIle phenomena, 84, 285-286 ontogeny, 84, 105. 122-143
death after flowering, 82-83, 85 periodicity> ph)">lologteal, in vegetative

i
l
distribution, :3
ßowering: annual vs. cons~ant vs. incon-
,sistent, 83; duration, 286; gregarious,
ll...'XCS, 11
recovery after flowering. 85, 286
relation to other plant groups.5
84, 285, 304; partial, 85. 285; polyga- n;productien, sexual: monoperiodic, 308;
mcus, 100, 117; spor«.dk, 85 1 316 periodicity, 285; pbysiological aspects,
flovrerlng cycle: length, 84; monoperiodic, 83, 102; po!yperiodic, 311
285, 308; polyperiodie, 285, 31 I reproductive phase, S2-ö4
flowering habits: as related to field iden ti- scedlings, development of, 126-143
fication .and taxonomy) 87; by :>peci.es, spceics, selected: pcrspcctivcs, 147-149;
267-279 portraya:s, 150-201
fruition, 8.5-86) 202, 286, selection of bamboos elite for specific
genera, se1ectcd. key, 289 neecls, 148-149
genctic ronstitution, 203, 205 sources, commercial: ef planw, 293-294;
growth: continuous w. seasona1, 282; eOfi- spccies offered, 292-293
trol by hiochemical graclients, 6; inde- sterility, eonsta."ll, 82
terminate, 305; intercaiary, 123; iter- strains, ph)">iologiea!; 82

1 auctant, 306; grand period of, 91, 123;

by pleiogeny, 310; by proliferation and
proli5cation, 311; start~and~stop, in
strucrures: homologous:, variation in, 6;
rcproductivc, sec Inflo:rcscence, FIoret,
F1owcr, Fruit. Seed, Spikcict; vegeta-
rhizome necks, 18; to mature stature, tive, Set Culm, Neck, Rhizome, Roots,
125; by tillering, 16,22,31-32.35; 38, Sheathing argans
40, 283, 315 studies, exploratory, praerical Te$ults, 148
homotogies; alleged, benvcen vcgctathle study materia:s: assemhly of, technique.s

- - - - - - ....... _~._---.
346 L'IDEX OF SL'BJECTS I:\'DEX OF SL'BJECTS 347

for. 81; doeumentatioo. 147; living, 148; delphous, 287, 307; tnadelphous, 287, Group I, 240-244; spontaneous roodng Root(s)
preservcd, 87 317 in. situ, 244-250; bulbUs 1 natural propa- adventitious. :J, 79-80
trnnsition between vegetative and repro- gynoedum (pistil), 111, 117, 287, 305 gation by, 250, Group Ir, 241; dump anatomy, lItudies, 81
cuctive expression, 103. 106, 286 lodicules, 112-114, 287, 307 divisions, 210-218; culm segments: distribution: on rhizomes. 14-15. 79; in
vegetative aetivity, suppression during ovary (ovulary), 115-1I6, 287-288 Group I, 234-239) 257, Graup 1I, 239- wil, 79, 81
flowcring, 83 stigma. 2SS 241; culm.s, whole: Group I, 227-234) primary, 79, 122
vegetative phase: maturing ;:>kmt, 11; style, 116. 287-288 Group II, 234; !aycring; Group I, 250- pnmordia on culm branches, 79
seed!ing, 122 zygomorphy, 114, 318 255; rhizomes alone: Group I, 217-226, thornlike, on culm nodes. 79-80, 175, 284
yields of culms, by species, :49, 159-160~ Flowering bnmch, 303 Graup 11, 226-228; summary (eone1u~ SeedC'), 117,20:/-204
176, 190 Fruit(s), 117-121, 288 sions) 255-256 embryo. comparative studies; 120; endo-
Bud(,), 298 destn.:.ction by natural agents, 86 objectives, 202, 205-206, 25 7; requirements sperm.; 121, 302
relation, physlological, LO suhtendw.& embryotegium, 118-119) S02 in rcla~ion (0, 202, 205-208, 2'10; 239, genie <:onstitution, 203
sheath, 62 studie:;, 107-108 257,261 germination, 134-135, 204, 238
Consti!Uents and exudates su also Seed(s) problems: decay of rhizomes, 224-225; viability, 203
cyanogens. 3 Inflorescencc. 89, 108, 286, 305 dormancy ofbud:;, 225, 259, 261,,; root~ stt alse Fruit($)
l;:ozymes ,2 bracts and buds, 90-91, 95-96, l00~lOZt ing, promotion of, 221, 260; study by Sbeathing organs, 61 ·78
glucuronk acid, 2 286,298 nev.- 2.tlproach sugges:ted, 261-262; vari- categories: differentiated, 6.3, 314; repro-
powder on culm intemodes, 2, 46-47, 62 branching: determinate, 91, 94-95. 97, ables, uncontroHed, 210, '258-261 duetive, su Fioret, Flower, Infieres-
tabashir.2 101-102,286,301, p:lnlcu1ate, 102,309, seale of operations. 20-2 cence Spikelet
j

GuIr.1, 36 racemose, 101, 312, secund, 101, 313, techniques .and procedures: convcntionaJ, vegetative: branch sheaths, 69; culm
anatomy, studios, 49 spicate, 101, 315; indetetminate, 90. limitations of. 207; fact:> and factors, sheaths, 64-68, 285; leaf, 306; leaf
bast:" interpretation, 23, 40-41 9~95, 97-100, 105, 305, pseudospikdet, 257-261 sheaths, 69-70, 306; neck sheaths, 76,
branch buds, 48-49,51,54, 284j relation 91-93, 97-100, 236, 312 variables requiring consideration: age 0: 309; prophyHa) 57,76-78, loa. 312; rhi-
to channcling of intemodes, 43-45 dlstinction from flowering braneh, .303 material, 211, 258; ambient conditions 7.ome sheaths, 75-76, 313
branch eomplement, 48, 50-53, 55-59, evolution, t01, 106, 112 for propagules, 207; ch:arncteristics of component parts, 62; auricles, 64, 70;
284; mouodade" S08; pleioclade, 310; form.and ontogeny, physiologieaJ aspects, plant, physiological and morphological, blade, 64-66, 69; callus, 48. 67; gird;e,
pdmordia, pre::oeious proliferation. 51, ]02-106 208--2:0; part ofplant uscd 255; ph~i~
j
67-68, 304; iiguk, 70, 306; oral setae,
5.3-54, insertion: apsidatc, 297, corutd- growth, grand period of, 91, 123 olog::cal state of propag-.l!es, 210, 258; 64-65) 70; sheath proper) 62
tate, 299, gremial, 304, restricted, 312 pediccl.. :09, 309 season of preparation of propagules, disparities: berween ax("s, 62; within a.xt:S
branclies: length pattern, 60; thornlike, pedunele, 286, 309 210; size of propagules, 211, 255--256 63-54
60-61; 284 prophyl!a, 90,93,95-97,99,108,119,912 Rhizome primordia, 61
dimensions at maturity, 283 pulv:ini, 101-102, 312 anatomy~ studles, 13, 23 Spikeleo.:, 109-110. 2B6, 315
fiber leng:h) 164, -2.01 rachis, 90, 96, 101-102, 312 buds at nodes. 19 anatomy, studies, 107
habit, 36, 39; 283 re!ation to vegetative axis, 286 componem struetures: neck, 1.3; neck compone.nt structures:: glumes, empty}
imernodes, 204; elongation and suppres- Leaf, 308 sheaths, 309; rhizome proper, 19; rhi- 109, 112, 286, 302; lemmas (ftowering
sion, atypical, 42; exudate, 2,46-41,62; anatomy, studies, 72, 75 zome sheaths, 62-63, 75, 3:3 glun:es) 109, 236, 306; rnchilla, 96, 109,

II
length pattern, 42-45; lumen, 284, 307; biade, 71-75, 285 femu: (types): detenninate and indeter~ 286) 312
shape. 284; texture and vestun; 46 deciduousness and renewal, 285 minate, 25; l.eptomorph, 15-16,25, 31, polygamy, 100, 117
life span, 283 epi dermis, 71-75 33. 35. 94, 306; monopodial, 21, 308; proliferation and prolification~ 103
nodes, 48. 283) 299 petlole and pulv.ru, 64, 71, 312 pachymorph, 14. 17-!8, 20. 24, 29. 31, pseudospikclet 'IS. spikelet, 90
origins, 32) 35~S6, 38, 40 sheath, 64, 69-70, 73,306, 316 33, 40-41, 92, 309; sympodial, 21, 316 Studies suggested, 6, 11, 12, 13, 16, 23~ 58,
primat)'. 123 spodograms and key to Japanese genera. neck, funetiMal behavior, 17-·18, 26-27, 61,62 1 72,81.102, 106,107,120,148)
shape, 42 73-7. 131. 133 149,157,163,171) 178, 187,201,207,
ilieath, 64-68, 285, SOO venation, 72-73, 285 Rhizome system, 13-35 258,264-
sheath nodc and sheath sCllT, 48, 52, 283,
S14
shoot, 124,300,315
vernation, 71. 318
Neck, 1'2, 308 ! forms, known, 26, 32 (key)
interpreta;:':ons, 16-35
Uses and utilization, 1-2, 150, 1:>4-155, 159,
164,176) 179-181. 183.190-191,193-
194, 197
supranodaJ ridge 52, 283
yields (production). 149, 159-160, 176, 190
Propagation, 202
auxins in relation to. 242-243, 251-254,
260
I metamorph axes, 28, 307; metamorph I,
26, 29; me;:amorph 1I, 26, 30, 35
relation to <::lump habit, 13, 283
Floret, 97. 109. 111, 286, SOS materials and metho<ls: groups I and H,
stip,itate, 286 differenees in relation to appropriate
palca. and prophyllum, 108, 1:S, 286-287, procedures, 208··209
309, 312 seeds, 202-204; deve!opment ofseed~
Flower, 97, 109, 111-112, 28";, S03 lings, 126-140.157; diffieulties prevai1~
anatomy, studics, 112 ing; 202-204; planting, 204-205; viabil~
androecium (stamen complcment). 111~ ity and storage, 203-204
115,287; diaddphous, 287, 301; mona.- vegetative fractions: branches, single,