Tropical Trees and Forests: An Architectural Analysis

456 Pages·1978·37.96 MB·English

by  Prof. Dr. Francis Hallé, Prof. Dr. Roelof A. A. Oldeman, Prof. Dr. Philip B. Tomlinson (auth.)

F. Halle R. A. A. Oldeman P. B. Tomlinson
Tropical Trees
and Forests
An Architectural Analysis
With 111 Figures
Springer-Verlag
Berlin Heidelberg New York 1978
Prof. Dr. FRANCIS HALLE
Institut Botanique, Universite des Sciences
et Techniques du Languedoc
5, Rue Auguste-Broussonet, F-34000 Montpellier
Prof. Dr. ROELOF A.A. OLDEMAN
"Hinkeloord ", Vakgroep Bosteeit
Generaal Foulkesweg 64, NL-6703 BV Wageningen
(formerly: O.R.S.T.O.M., Apartado 99-B, Quito)
Prof. Dr. Pm LIP B. TOMLINSON
Harvard University, Harvard Forest
Petersham, Mass. 01366, USA
ISBN-13: 978-3-642-81192-0 e-ISBN-13: 978-3-642-81190-6
DOl: 10.1007/978-3-642-81190-6
Library of Congress Cataloging in Publication Data. HALLE, FRANCIS. Tropical
trees and forests. Bibliography: p. Includes indexes. I. Trees- Tropics. 2. Forest
flora - Tropics. 3. Trees - Morphology. 4. Trees - Growth. 5. Plant morpho-
genesis. I. OLDEMAI', R.A.A., joint author. II. TOMLI:--;SOI', P.B .. 1932-. joint
author. III. Title. QK493.5.H36 582'.16'0913 77-15112.
This work is subject to copyright. All rights are reserved, whether the whole
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':J::: by Springer-Verlag Berlin' Heidelberg 1978.
Softcover reprint of the hardcover 1st edition 1978
The use of registered names, trademarks, etc. in this publication does not imply,
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Reproduction of the figures: Gustav Dreher GmbH. Stuttgart.
2131/3130-543210.
ladis
les arbres
etaient des gens comme nous
mais plus solides
plus heureux
plus amoureux peut-etre
plus sages
c'est tout
J. PREVERT
Gedaagde, bodemvaste boschgenooten
boomen die'k, weI vichtig jaren lang
boom wete; en, zoo hooge als nu geschoten
gezien hebbe, op zoo menig wandelg ang
wat ben ik, arme miere, u bijgeleken
die sta en u aanschouwe, 0 hooge boomenreken
GUIDO GEZELLE, "Eeuwelingen"
Botany needs help from the tropics;
its big plants will engender big thinking
E.J.H. CORNER
Preface
This book is not an exhaustive survey of known information
in the manner of a text-book -the subject is much too
big for this to be possible in a relatively concise volume-
but presents a point of view. We are concerned ultimately
with the analysis of tropical ecosystems, mainly forests,
in terms of their constituent units, the individual trees.
Many different approaches are possible in the analysis of
tropical forests. A simple one is to treat the trees as obstacles
which in a military sense intercept projectiles or are a hin-
drance to foot soldiers (ADDOR et aI., 1970). A similar ap-
proach might be adopted by an engineer confronted by
a forest which has to be removed to permit road construc-
tion. The timber merchant is concerned with the ability
of a forest to yield saleable lumber. The interest here is
in the size of the larger trunks with some concern for the
kinds of trees.
At a less destructive level the scientist aims to compre-
hend the forest from many different points of view. The
forester himself, in conjunction with the taxonomist, will
wish to analyze the floristic composition of the forest and
perhaps account for species diversity in an evolutionary
time scale (e.g., FEDOROV, 1966; ASHTON, 1969). The evolu-
tionary biologist in his turn may be concerned with repro-
ductive strategies in forest trees (e.g., BAWA, 1974), espe-
cially in a comparative way.
The approach adopted by the ecologist offers the greatest
scope, since he may combine several different methods of
analysis. Much research has gone into the physiognomy
of tropical forests, size distribution of trees, stratification,
diversity in relation to soil type or soil moisture content
and has been summarized recently by ROLLET (1974).
Phenological studies of tropical forests have produced a
great deal of data which reveals the extent to which flower-
ing, fruiting and leaf fall mayor may not be seasonal (e.g.,
COSTER, 1923; HOLTTUM, 1940, 1953; cf. also LIETH, 1970).
The production ecologist is interested in the forest as an
VIII Preface
efficient system for light interception and yield of dry mat-
ter, both in a relative and a comparative way (e.g., KIRA,
1978; KIRA etal., 1964, 1969; MONSI etal., 1973; BERNARD-
REVERSAT, 1975). Photosynthetic efficiency in terms initially
of leaf and branch orientation but ultimately in competitive
ability is another stimulating approach which is summarized
in the description of trees as "crafty green strategists"
(HORN, 1971).
A universal tendency in these approaches is to treat trees
as equivalent units- as taxonomic, physiological, reproduc-
tive units and so on. Much less attention has been given
to the trees in the forest as individuals. This is our approach.
However, we do not merely regard trees as individuals at
one point in time, but as genetically diverse, developing,
changing individuals, which respond in various ways to
fluctuations in climate and microclimate, the incidence of
insects, fungal and other parasites but particularly to
changes in surrounding trees. The tree is then seen as an
active, adaptable unit and the forest is made up of a vast
number of such units interacting with each other.
In order to understand the adaptive strategies of the tree
in the forest we must first, in a rather paradoxical way,
remove it from its natural habitat and study it in isolation,
more or less free from the natural accidents to which it
is otherwise subjected in a competitive environment. Iso-
lated in this way one can study the tree from the point
of view of the geneticist and developmental morphologist.
The tree develops from a seed which carries the genetic
information which will determine its form. It is only by
studying the form of the tree expressed in a more or less
optimal environment that its genetic potential is clearly
revealed. Briefly, we find out what the tree can "do". This
leads, therefore, to the recognition of what has been termed
the" architecture" of the tree (HALLE and OLDEMAN, 1970),
a concept which is elaborated in great detail in this book.
The concept of architecture involves the idea of form, impli-
citly containing also the history of such a form. The life-
long succession of developing forms in a plant is revealed
by the concept of the architectural model and its reiteration.
By examining large numbers of species in a comparative
way, we are led to the recognition of the existence of similar
,developmental plans among taxonomically dissimilar trees.
Much of this book is, therefore, a description of these
Preface IX
developmental" models" as established by HALLE and OL-
DEMAN, and this provides a framework to which we can
attach a great deal of information about the growth of
trees in the tropics. Recognition of the existence of similar
constructional principles (architectural models) in a great
variety of trees (and our survey has been as cosmopolitan
as we could make it) implies that the "strategy" of a tree
differs from species to species. What significance, in fact,
does architecture have for the success of the tree in a natural
environment?
Here we must return to the forest, away from our opti-
malized environment which has been so productive of new
information. We have learned how a tree is capable of
growing, by virtue of its genetic make-up. Now we can
ask the question, how, in fact, does it grow in the vigorously
competitive environment of the forest itself? This leads
to the recognition of ways in which a "real" tree is
constructed in a natural stressed environment, as distinct
from an "ideal" tree, growing precisely according to its
genetic plan and not subject to environmental stress. This
is not to suggest that we are making a distinction between
theoretical and practical information. Both circumstances
exist, and trees function in both optimal and nonoptimal
environments. The point will become clearly established
that without a knowledge of the potential growth activity
of the tree, it is impossible to recognize its actual growth
expression. Once this is appreciated we are in a position
to reassemble the living forest in terms of its developing
units. From this it should be appreciated that our approach
is a biological one in the purest sense.
Throughout the text we have stressed tropical examples
and our ultimate aim has been to understand trees of the
lowland, humid tropics. What is the reason for this em-
phasis? The answer is really quite simple. Taken in a very
general sense the climate of the wet tropics is uniformly
favorable for plant growth and allows the existence of an
enormous species diversity. Consequently in this environ-
ment there is a wider array of growth expressions in woody
plants than anywhere else in the world. If one seeks funda-
mental principles, it is clear that one should do so where
they are freely and clearly expressed. Once an understanding
of growth processes in tropical trees has been established,
it then becomes possible to look at forests in the more
x Preface
constrammg environments of temperate latitudes, where
diversity is less and is directed towards survival amidst
macro climatic stress. One can understand a temperate forest
better when one has grasped principles of tree growth in
the tropics, but scarcely the reverse.
Unfortunately, botanical history has inverted this logical
sequence. Consequently a degree of justification for our
approach has to be introduced via our initial thumb-nail
sketch of the floristic diversity of the tropics. For the same
reason we have attempted to provide some background
in morphogenetic terms for our subsequent analysis of tree
architecture, since our approach is often different from that
of a temperate forester for whom responses to seasonal
climates seem so important in tree growth. This is by no
means intended as a complete review of organogenesis in
woody plants of the tropics, but it should serve to clarify
further our point of view. Frequently, of course, a topic
can only be discussed with examples of temperate tree
species in mind because no information is available about
tropical species.
Our greatest problem, and one which is encountered by
all who have tried to describe tropical vegetation to an
audience which has never visited it, is that the majority
of plants are likely to be unfamiliar to the reader. We
have used examples of common or commercially valuable
trees, where appropriate, especially in the introductory
chapters. We have otherwise made frequent reference to
existing accounts of the more common tropical species,
and to the earlier account of HALLE and OLDEMAN (1970),
but in order that this book may carry as much new informa-
tion as possible most of the illustrated examples are new.
If the reader still retains a sense of the monstrous, the
fabulous or the unreal in using this book he should try
to make a mental reversal of the situation, which is botani-
cally more appropriate. How strange is the temperate tree,
leafless for a large part of the year, with such marked
synchrony in its development, its brief period of extension
growth, its ability to flower only once each year and with
its peculiar annual radial increments of growth in the wood.
Here is a bizarre object indeed! Organizational understand-
ing of woody plants must come to terms with growth princi-
ples in parts of the globe where they are most readpy com-
prehended, that is in the tropics. It is our intention to
Preface XI
make the future investigators' task an easier one by pre-
senting a rational basis for future research.
Acknowledgments. Organizations which have assisted in
the preparation of this book include the Maria Moors Ca-
bot Foundation and Atkins Fund, both of Harvard Univer-
sity, which have provided direct financial assistance. Dr.
OLDEMAN was able to visit Harvard Forest in October, 1973
as a Bullard Fellow of Harvard University and was granted
leave from the Office de la Recherche Scientifique et Tech-
nique Outre-Mer to take up this appointment. We are
indebted to Dr. MARTIN H. ZIMMERMANN, Director of the
Harvard Forest, for facilities made available during this
and other visits.
Numerous other individuals have played important roles.
Dr. HENRY S. HORN, Princeton University and Dr. JAMES
WHITE, University College, Dublin (both at one time tempo-
rary residents of Harvard Forest) read a preliminary draft
of the manuscript and made numerous constructive and
enjoyably pungent comments. They may still be dissatisfied
with the final product, but this is simply because it reflects
our prejudices and inadequacies. Several people have pro-
vided us with useful information about tree architecture,
including 1.M. VEILLON, MARIE FRANyOISE PREVOST and G.
CREMERS. Photographic and technical assistance in Peter-
sham was provided by ANNE FAULKNER, MONIKA MATT-
MULLER, USHER POSLUSZNY and REGULA ZIMMERMANN.
Typing and editing of manuscripts has been the respon-
sibility of SANDY WEIDLICH and DOTTY SMITH.
In general the willing support of our differing institutions
has been of great benefit during a task made difficult by
our wide geographical separation and the continuing need
to carry out our normal duties. Weare thankful that our
publishers have been patiently appreciative of this circum-
stance.
January, 1978 FRANCIS HALLE
ROELOF A.A. OLDEMAN
PHILIP B. TOMLINSON
Note: Since frequent reference is made to HALLE and
OLDEMAN (1970), for conciseness this is referred to as
"H.O., 1970" in the text.
Contents
Chapter J Introduction
A. What is a Tree? . I
I. Definitions. . I
II. Tree Making. 2
III. Apical Meristems and Tree Construction 4
I. Trees Built by One Meristem. . . . . 4
2. Trees with Modular Construction. . . 4
3. Trees with Trunk-Branch Differentiation 5
4. Trees with Changes in Orientation of Axes . 5
B. The Botanical World of the Tropics . . . . . . . 6
I. Distribution and Size of Flowering Plant Families 6
II. Tropical Floras as Tree Floras . . 8
III. Floristic Richness in Limited Areas. . . 10
IV. Geographic Locations. . . . . . . . . ]0
V. Climate and Tree Growth in the Tropics 10
Chapter 2 Elements of Tree Architecture
A. The Initiation of the Tree. . . . . . 13
I. Seedling Morphology . . . . . 13
II. Juvenility and Phases of Development. 16
B. Apical Meristems and Buds. 18
I. Terminal Buds. . 18
II. Bud Composition. . . 22
III. Lateral Buds. . . . . 22
IV. Secondary Bud Complexes. 23
C. Extension Growth in Tropical Trees 24
I. Introductory Remarks. . . . 24
II. Rhythmic Growth . . . . . 25
I. Rhythmic Growth in Hevea 25
2. Preformation and Neoformation 32
3. Further Examples of Rhythmic Growth 32
III. Continuous Growth. 36
I. Palms ........ . 37
2. A Dicotyledon. . . . . 38
D. Phyllotaxis and Shoot Symmetry 39
I. Primary Orientation. . 39
II. Secondary Orientation. . . 39