Relaxing in the jungle, French Guiana. Rcpnyduced by courtcsy of Nlary Evans Picture Library.

An Introduction to

Tropical Rain Forests

Second Edition.

T.C. WHITIx/{ORE ”

(Ïixford New York Tolqgt; OXFORD UNIVERSITY PRESS 1998

Oxfirra’ Ï_lítl1!€1‘Sll}-' Press, Great Clarendraiz Stireet, Oxford 0X2 GDF

s Oxford New Yorlz Athens Aiicrkliïnd Bangkok Bogota Bombay Buenos Aires Calentta (Ïïajie Toreo Dar es
Salaam Delhi Florence Horng Kong lstarilsvirl Karaclyi Kuala Liiiízpttr ¿’limites Madrid Mellmttrite
lkdexicr; City Alairobi Paris Siiïgtïfïoïe Taipei Tokyo Toronto» íïlarsaiov

and associciteci corripaizies in Berlin l laadaiz

Oxford is a trade tirarle of (Jxforcl Unit-ersity Press

Published in the U irited States

by Oxfiurd Urtitiersiíy Press, lira, New York

© TC. ‘iïlhiiiriore, l998

All rights TBSEÏUBd. N o part of this pithlieaticnr may by? areprorlirced, stored in a retrieval system, or
traizsmitted, in any ftirm or hy any means, witlïotlï the prior permission in iariting of Oxford Uiriversity
Press. íïlithin the UK, exceptions are czlloreed in resjrect of any fair ¿feeling for the ptirgmse of research or
private study, or cariticism or TGIJÍHMI, a5 permitted ¿trader the Copyright, Designs anal Piztettts Act,
1988, or in tlze case of reprograpbic i-ejirtidiietion in aeeoiïltiiiee with the ternrs cif . licencias issaed by the
Copyrigght ljeeizsing Agency. ljirqiiiries ¿‘oncerszing reproduction mitsíde those terms and in other
ecmntiries should be sem‘ to . the Rights Department, Oxford [lizitieisity Pi-ess, at tlsre address above.

This hoolz is stilo’ subject to tlae condition thai‘ it shall not, by tod)’ oftrade or otherwise, he lüïïf, re-sold,
hired ont, or otherwise cirenlaterl without. the fmhlisliefs {irior eonsent in any form of hincliiig or cover
other than that in iohich it is piihlislyed and mithcnrt a sitriilar cozrdition iltcltldíïlg tlJis condition being
iinposetl on the siihsequent pairchaser.

A catalogitre record for this lzoolx is auciilalale front the Biritisl: Library

Library of Congress (Ïataloging in Publication Data

‘Whitniryize, T. C. {Timothy Charles) An introduction to tropical rain foresïs/ ÏÏC. Wlhitrrzore. -— 2nd ed.
lnclirdes bibliographical references (p. 251) and index. I. Rain forest ecology. 2. Rain forests. l. Title Ql-
l54l.5.R27‘L-Ï745 1998 577.34—dc2I 97-3l35Ü

¡sen o r9 850148 x {Hole} o r9 8501471 (Phlz)

Typeset by
EXPO Holdings, NlrIltn-‘sitt
Piintea’ io Great Britain hy
The Bath —Press, AvonPreface to second edition

There has been a change since 1989 in the per- ception of tropical rain forcsts. Faltering steps are bóllïg taken
to attempt to ccmtain impact on then], and indccd all the tvorld’s forests. The catalyst was the 1992 United
Nations Conference on Environment and Development at Rio de janeiro. Chapter '10 has been com- pletely’
rcwrittcn and cxpandcd into two chap- zcrs to rcllect the numerous national and international moves aímed to
make exploitatioii sustainable. This includes forest Certification and so-called ‘cco-labelling’ of timbcr. The
most recent global assessment of human impact on íïüpicaïl forcsts by FJXO is summarizcd. Timhcr
cxpleoiteltíian, contlnucs to spread, even though the other iralucs of forcsts as rcscrvoirs of bio- logical
diversity and for local people are now morc- widcly accepted. The Virgin Eastern Ffllll forests are nearing
cxhaustion and moves into tropical Anwríca are beginning. Rain forcsts are increasingly being reduced to
fragincnts in anthropcagenic landscapcs. The nature of biodi- vcrslty, its fate, and steps tU be taken to
describe and conserve it, have recently come under the Spotlight. These new dcvelopmcnts are all described.
Topics of particular: concern, and an agenda for action, are highliglatcd. These ¿Ispccrs are pcrheips those
that the layrman hears most about, and perhalas which interest him most. Their lntcrpretation rcsts on biology
and ecology which, as before, are described in the first ¡nue clltlptcrs. Here, change has been mainly an
incremental increase in knowledge and understanding, with a few important break-

throughs, though there are no maior ¡nalradigm- shifts to report.

Information is brought up to date throughout, atytenlpting to incorporarc the main points from what sccms to
lac an increasing ÍOITCIIT of publi- cations. Some: of ‘rhc highlightsare e15 follows. Chapter Z: thc
tlorthcrix limit of rain forest is now increased to 28°, and the castern flank of the Andes is confirmed as the
richest region for both plant and animal species; the record is nmv 307 tree species on one hcctarc. Chapter 3:
there is better understanding of flower-ing and fruiting patterns in scastnml climatcs. Chapter 6': the past
history of rain forcsts and their biogcogrix- ph)’ is clcarer, especially in America and Asia. Chapter 7 on
forest dynamícs, is stlbstantiellly rewrittcn and cxpandcd. After over 20 years of intensivo research rhc
framework is lïrmly in place, continuíng studies ¿Irc now beginning to fill it in. The complexity of the
microclhnatc of Canopy gaps and the adjacciat undcrstotcy has been unraixcllcd. This forms the basis for a
better Linderstanding of shade-tolcrailcc, the ccophysiologícal basis ofwhich can now also be described.
Interest in lour-impact logging, cxploitatloi) within the ecological limito ¡of the forest conccrncd, is
burgconíxlg, and the neces- sary Procedures are described. The near ubiquity of communityr-scalc
dísturbanccs to rain forest, many of thcm human, has bccn incrcasinglyw rcalíscd. Their signs arc, aualyscd.
Chapter 8: advances in Lmdcrstanding the site conditions that determine the (JCCLIIFEHCE of both hcath
and upper montana forest arc described.

vi Preface ¡‘o seconrï edition

The book, as before, aims to introduce tropi- cal rain forests to a wide audiencia, to attempt to Conve}
something of their fascinatíoti and importance, the scientific chailenges they stili pose, and what is really
Iiappening to them. Even at this level there is much new information that clarifies or exrends what was
known 8 years ago, and there are important political changes too. It has: been a challenge to cover
developments without substantialiy increasing the length.

To enhance the usefulness of the book as a starting point fair deepcr enquiry, more numer- ous references to
other books and reviews are included. A fulier description is gixïezi of siivicuïture.

Cczmbridge T.C.\‘I(Í.
Níarch 1997Preface to first edition
The aim of this book is to provide an introduc-

tion to the worlds tropical rain forests for a hroad audience, to describe their structure and functioning, their
value to man, and what he is doing to them. Examples are tlrawil from all parts of the humid tropics. Today
there is Inore research being conducted in tropical rain forests than EVE)!‘ laeftire. Stirne is driven by
curiosity, some by the tlesire to harness these forests to mankind’s needs. What generalizations can be made?
To what extent are there real differences between rain forests in different places, and what are the current
frontiers of knowledge? ls present-deiy concern about man’s impacts on tropical rain forests justified? The
book seeks to provide an ansvirer to these questions, at a simple level accessible to all tvhci x-vant to know
something about these grand forests.”

ln lands where tropical rain forests occur man has lived iu closest dependence on them since time
immemtirial. ELIIOPCHIIS becelme aware of them over two inillennia ago (Chapter 1). Increasing
knowledge since the Renaissance with the Vüjïdges of discovery and then the Cültïl- nial era revealed that
there are in fact Inanv dif- ferent kinds of tropical rain forest (Chapter 2). Plants exist in a luxurizince and a
diversity of ¡Jizarrc forms tindreamed Of in ternperate lati- tudes (Chapter 3). Animal life is also rich and
diverse (Chapter 4}. Modern science continues to unravel the many kinds of ctamplicated ir1ter—

dependence of plants and amimals, for example, in flower pollinatitm and seed dispersal (Chapter S).
Tropical rain forests have waxed and waned in extent through geological time, and the present patterns of
species’ distributions are a result of these historical events. The former idea that these great forests have sur-
viired innnutable ‘since the dawn of time’ is a rmnantic fallacji, as investigations of the last three decades
have shown {Chatpter 6). The forests are also continuously‘ changing at the other end of the time scale, the
life span of an individual tree. The elucidzltion of forest dynam- ics (Chapter 7} has been the other major
lnreak- through of recent years. Wïe now know a great deal about the ecology of individual tree species and
the particular requirements for groi-vth of their seedlings in eanopy gaps formed by the death of big trees.
Silvieulture, the manipnlaritin of forests by man to favour tree species of his choice, is Clepentlent on
undetstatntling these innate characteristics. Tropical rain forests can be a sustained source of timher, renewed
by regrosvth after felling, so long as (and it is a vital proviso) humans work within the limits of their natural
dynamics. Tropical rain forest nutrient cycles (Chapter S) are also now reasonably well LITlClCTSIÏUOCl,
with enough detailed studies made for tentative generalizaticz-ns to be possible. The old ideas of a closed
cycle and with nearly all the nutrients in the plants have not stirvived.

‘* To aid comprchension some of the more technical or specialized terms are defined in a glossarv at the end
of the lJOOk, and

some ttonccpts are (liscussed more fully in the text notes

.viii "

Shifting agricnlture is now mvelHcnrJwrn to be a sustainable form of farming, suitable for infer- tile soils.
Sustainable human utilization of forest lands for crops or trees depends, as with silvicul- ture, on vrorlcing
within the natural limits of the nutrient cycle.

One of the driving forces of science is the puzzles found in nature. Of these one of the great- est is to
understand the phentimenal species rich- ness of tropical rain forests. The extremes so far discovered are a
100 m2 plot in Costa Rica on which grew 233 species of vascular plants (includ-

ing 73 tree species, mostly as seedlings) (Table

2.3), and a 10 000 m2 plot in Peru with 580 trees of over 10 crn in diameter, of which every second stem suas
a different species (Fig. 2.27}. In both samples the study plot did not contain all the species in its locality.
Many different factors already‘ mentioned contribute to this hyper- ríchness; they are brought together in
Chapter .9. The book ends with a considera tion of human. impact on tropical rain forests, Chapter 10. This
final chapter starts tvíth a historical sketch of increasing interventíon through historical time.

Present-day iinpacts are described, and it is

shown that pan-tropical gerteralizations are sim- plistic and misleatling. The very real causes for concern are
analysed. These have led to the increasing public unease, and this in turn has

triggeted a response in governments, interna

Preface to first edition

tional institutions, and businesses which, as this book goes to press, is suddenly accelerating. Because of this
response, the book is able to end on an optimistic note. All is not yet lost of the world’s tropical rain ftarests.
There are still big gaps in our scientific untlerstantlingwvhich Iieed urgently to be addressed. But mankind
has the ability to discover how these self-perpetuating dynamic ecosysterns ‘¡York and then turn that l-
znotvledge to advantage. The present book aims to review the scientific basis for good decisions.

-lf human societíes can then exercise. the self-

restraint to utilize rain forests by working within their ¡iatural iimits the cries of doom and gloom of the late
eighties will have served a useful purpose. This is a hig ‘if’.

Why have tropical rain forests always laeen so thrilling to biologists? Wlhat is their excitement that drives
scíentists throughout the world forward vuhen for most the pay is so poor and the support of their employing
institutes and universities minimal? Is present public concern about rain forest}: justïfied? Are they really dis-
appearing, and if they are, does it matter? Are they so fragile that their sustainable tztilization is impossible?
These are the questions to which the following pages are addressed.

Wfirney, (Jxon
June 1989 TILW’.Ac/erzowledgements

There seemed to be a need for a book that iooked at all aspects of tropical rain forests and rnankind’s impact
upon them, yet in a concise text, written so as to be readily understood by the concerned layman as well as
the non- specialist college student. Here, I have attempted this daunting task, but the wider the subject matter
the further the scientist strays outside his own special field. The opinions and interpreta- tions remain my
responsibility but Í have been dependent on assistance from a large group of peopie, unfortunately too
numerous to mention individually by name. I do mention especially the help of S.C. Chin, R.T. Corlett, P.
Lucas, I. Polunin, l-Iugh Tan, and LM. Turner, who made valuable comments on drafts of the whole text and
those who likewise commented on one or a few Chapters, namely C.P. Burnham, N.M. Collins, WK. Gong,
TJ. Lowery, N. Sizer, and Tamin. For all this assistance I express my deep thanks.

The following acknowledgements are made for the figures: Ambio (Fig. 10.14); American Museum of
Natural History Novitates 23 87 12 Arlarch 1987 (Fig. 2.10); P5. Ashton (Fig. 1.5}; Biological Conservation
(Fig. 10.15); Biotropica (Fig. 10.20); British Ecological Society (Figs. 2.32, 3.26, 8.5); N.D. Brown (Fig.
7.12); E. Brunig (Fig. 2.11); Butterworth (Figs. 6.10, 6.11, 6.12); Cambridge University Press (Fig. 6.1);
Lord Cranbrook (Figs. 2.8, 2.9); DJ. Chivers íFig. 5.10}; Ding Hou (Fig. 3.13); j. Dransfield (Figs. 3.14, 5.2);
Elsevier (Figs. 4.3,. 4.5); Evolution. (Fig. 6.17); Gustav Fischer Verlag ajFigs. 2.1, 2.19); Forest Research
Institute Kepong

(Figs. 3.34, 3.35, 3.37, 3.40); A- Czentry (Fig. 1.6};

' C. HuXley-Lambrick (Fig. 5.11); World Conser-

vation Monitoring Centre (Fig. 4.1); International Palm Society (Figs._6.2, 6,9); Junk (Figs. 7.3, 7.13, 10.9);
LA. MacKinnon (Fig. 4.4); Malayan Nature Society (Figs. 5 .14, 5 .15 , 7.1); ll/Iaiaysian Forester (Fig. 3.25);
S. Mayo (aroids on Figs. 1.8, 2.2, 3.17, 9.1); Nature (Fig. 7.38); Photo- cfaemismv and Phorobiology (Fig.
7.2); I. Polunin (Figa. 3.22, 3.36); H. Rijksen (Fig; 3.2); Rijksherbarium Leiden (Fig. 6.15}; Science (Fig.
5.7); Springer Verlag (Figs. 2.21, 8.8); IVLD. Swaine lFigs. 3.31, 3.33, 7.34, 10.6, 11.6); j. Tan (Figs. 4.6,
10.19); Taxon (Fig. 10.17); Rosemary Wise lFigs. 5.14, 5.15). _

Readers of my earlíer publications will recog- nize some of the illustrations. In particular I have borrowed
generously from my study of the Eastern rain forests (Whitmore 1975, 1984a} which was written to a greater
depth and for a more specialist audience than the present work.

I thank Professor Gloria Lim and Associate Professor S.C. Goh who enabled me to spend one semester at the
National University of Singapore in an ideal environment, and to the undergraduates who took courses A203
and A303 in 1988 and were unwitting guinea-pigs for most of the text. Ivan Polunin provided the ititimate
contact with living rain forest, which varas an indispensable source of inspiration. Christine Brotherton gave
invaluable assistance, especially in chasing references.

Finally, ¡ny wife typed the whole book in all ‘its drafts and has made the whole enterprise possible.
x Acleizowledgemezzrs, Second edition

Many friends and colleagues have continued to contribute ideas and criticisms tvhich have helped to clarify
my understanding. "ln particu- lar, parts of this new edition were read and commented on by S.R. Aiken, R.
Borchert, LA. Bruijnzeel, PD. Hardcastle, R. Motley’, J. Procter, E.V.]. Tanncr and LM. Turner.

I am most grateful to Professor G. Glatzel who provided me with the chance to try out the complete revision
of human impact, which forms the twti final chapters, in a course of lectures at the Institute of Forest
Ecology, Universitat für Bodenkultnr, Vienna, and to the students who came to listen.

The following acknowledgements are made for tiet-v figures: CAB international (Fig. 10.26);

Conservation Biology (Fig. 11.5); Forest Ecology ci" Management (Elsevier Science) (Pig. 7.16); Forestry
Department HQ, Peninsular Malaysia (Fig. 10.16); the late T. lnouc {cover photo- graph); Institute of
Hydrology’ (Fig. 10.30}; ltzrerciencia (Fig. lüfilgflaurnai (JfECÜfKJgy (Fig.

7.21); journal of Tropical Ecoiogy (Cambridge

University Press) (Fig. 7.21}; KT. Lee (Fig. 11.7); Oryx {Fauna ¿c Flora International) (Fig. 10.27); World
Wide Fund for Nature (Fig. 10.25). Although every effort has been made to trace and contact copyright
holders, in a few instances this has not been possible. If notified the puhlishers will be pleased to rectify any
omission in future editions.

Contents
Explartartary ¡totes xív

Reference; and fanfic-ev‘ reading‘ ' xiv Ussits, sgnvnfaerfs, ana’ alabretxiariorts - Xiv 1 A11 introduction to
tropical rain forests 1 2 What are tropical rain fnrests? 10 2.1 Tropical InOÍSt forests and their clinmtes 10

2.2 The Forrest f<_)r11'1a[it)¡1s 13

3.3 Forest nmintenance-{he growth cycle »-. 25

2.4 Floristics . 31

2 5 Nature of the tropical rain forest comlnuttity ___ 35

2.6 Vílhat are tropical rain fOIeStSP-Chapter stlmmary 38

3 Plant life 40
3.1 Climbers and epiphytes 46
3.2 ’l'rees l 51
3.3 Scasünal rhythm}; 58
3.4 Plant life-{Íhapter sunnmary 63
4 Rain forest animals 65
4.1 Riclmcss and diversity of animals 65

4.2 Modas of coexistence 68

4.3 Carrjging capacity of the forest 71

4.4 Rain ftarest aninïalsrljhapter summary 3

S Interconnections between plants and animals-the web

of life 74 5.1 Animals as; pollingators 74 5.2 Anílnals as disperscrs 78 5.3 FOOCl webs, mobile links, and
kcystone species; 80 5 .4 (jo-cx-"olution ‘ 8 1 5.5 Iznplícations for Forest conserx-‘ation 85 5.6 Plant
breeding systems 86

Contents

5.7 Interconnections between plants and animals-the web ' of life—Chapter summary 87 6 Tropical rain
forests through time 89 6.1 Palaeogeogreaphy _ 89 6.2 Palaeoclimates ' , 97 6.3 Pleistocene refrlgia _ 103 6.4
Seasonal climate elements in Malesia ' 106 6.5 High mountain plants 106 6.6 Conservation 107 6.7 Tropical
rain forests through rime—Chapter summary 107 7 Forest dynamics 109 7.1 Forest mieroelimates ' 109 7.2
Pioneer and clímax tree species "l 13 7.3 Seed and seedling ecology 120 7.4 Components of species richness
124 7.5 Growth and shade tolerance anaíysed 126 7.6 Silviculture ' 13'} 7.7 Growth rates and longevity 142
7.8 Community-wide disturbances 145 7.9 Forest recovery after human Llistar-banca 151 7.10 The dynamics
of clímax rain forest 152 7.11 Forest dynamics-Chapter summary 154 8 Nutrients and their cycles 156 3.1
Shifting agricultura? 156 8.2 Nutrient pools and cycles in primary rain forest 159 8.3 Practical implications
164 8.4 Heath forest—a fragile ecosystem _ 169 8.5 The upper montane forest enigma 171 - 8.6 Nutrients
and their CyClES—Chapter summary 172 9 Species richness 174 10 Tropical rain forests yesterday and
today 179 10.1 Indigenous cultures 179 10.2 The colonial era 185 10.3 Minor forest products ,1 186 10.4
Tropical rain forests in modern times " 190 10.5 Forest conversion to other land usages . 191 10.6 Logging
and the tiïnber trade ' 19.9 10.7 Animals in logged forest 206 10.8 Rates of disappearance 207 10.9
"Controlling human impact Zl Ü 10.10 Causes for concern 213

10.11 Tropical rain forests yesterday and traclayn-Chelpter summary 220

(jorzrents xíii

'11 Tropical rain forests at the eusp of the new

nnllenniunr 222 "l 1.1 Management for sustainahility‘ 222 l ‘l .2 Forest Certification 223 11.3 The future
pattern of land usage 224 11.4 Prioríties for elction 233

11.5 Tropical rain forests at the eusp of the new millenniuin-Cliaptttr summary 235 Egfllogize 2.3? ïïrxt notes
239 Referentes 251 Glossary 266 lzzdex to ¡’HÏGIEZS ¿md minor forest ¿Urodzrers 271

G ¿neral index 2.78Explanation; notes

REFERENCES ¿ND FLÉRTIÏER READINC}
This book is an introduction to tropical rain forests and the text aims to tliseuss the main topics. The reader
kvho wants to discover more can do so via the notes at the back of the book Which are referred to by
superscript numbers in the text. In the text notes are named original research papers on which particular
points are based. ln addition, for every chapter these text notes include a selec- tion of books and rei-‘iew
papers which give fullcr information than there is space for here. The sources of data included in Figures and
Tables are given in their captions. All the papers referred to in text notes, Figures, and Tables are cited
together at the end of the lnook.

Knowledge of tropical rain forest fills a great Warehouse. The references cite-d have been care-fully chosen
to help the reader peer through its VJÍflClÜWS and discover the huge store of knoowletlge arranged within
its “Kills.

UNITS, SYMBOLS, AND ¿KBBREKFIATIKDNS

Througbout this book S] units are used. The Convention is followed that values such as metres per year or
stems per hectare are shovm as In year”, stems ha“ not mfyrear, Sttïlïlsfllél.

means ‘greater than’

means ‘equal to or greater than’ means ‘less than’

means ‘equal to or less than’

a A iv v

B.P. means years Before Present

c. = Circa, abüul’

M = mega, 10*‘

n = nano, HT”

s.l. means ‘in the broad sense’

S means SUS1

An introduction to tropical rain forests

n-ain ftirests have crossed a th reshrold of taercep- ion. Reports on television, radio, or in the press of another
piece of destruction, or a new message of gloom for the planet, have become commonplzice.‘ The public
firmly believes that something nasty is happening down on the Equator, even that the- once vast Amazon rain
ïoresrs have all but disappeaireti. Man’s present- iajs‘ impact on tropical rain f-orests is, however, fest the last
stanza of a saga stretching back into the past beyond the begitining of written history.
European knowledge of tropical fcrrests began when Alexander the Great crosscd the Khyber Pass in 327
nc‘, into the Puniab, to esrablish the eastern limits of his short-lived empire on the banks of the Indus? ‘His
army saw mangroiïe sivamps (tvhich upset conventional views on trees), jackfruit, mangoes, bananas,
Cotton, and ïranjrans-«which upset everybody/E views on -.s-hat rtmots are supposed to do.’3 These bizarre
Índings were incorporated in the Earn/tir)‘ into 71321:: of Theophrastus, PhÍlOSIJPlICI‘ and pupil of Plato
and Aristotle, to become part of the general knowledge of plants, copied, corrupted, 2nd no1‘. improved upon
for nearly two thousand years until the great Voyages of discovery of the sixteenth and seventeenth centuries
and the sub- sequent Etiropean colonial expansion.

The word jungle, still often in use, comes from

the Hindi jangal, a reference to the dense im— "

penerrable forest and scrub around settlements.

Tall stories percolated back to Ettrope from the early visitors (Fig. 1.1). For example, in the Fast, poisoned
arrows WETC discovered to be tipped with the sap of the upas tree,‘ accounts of whose identity and
‘preparation mingled fact and fahle. Of this the great Dutch naturalist

GF. Rumpf wrote in 1750:

Under the tree itself no plant, shruh or grass grows- not only within its periphery but, CVCD, nor ‘within a
stoncïs throw of it; the soil is sterile, dark and as if burned. Such poisonousness tioes the tree exhibir that
from the infected air birds perching on the branehes are stupefied and fall dead, and their feathcrs srrew the
soil. So caustic i-vere the branches sent to me in a stout hamboo vessel that when the hand ivas placed on the
txessel, a tingling was produced such as one feels on coming out of the cold into the Wtïrlïïth. Eircryvthing
pcrishes which is affected by its exhala- tion, so that ali animals avoid it, and birds seek not to fly over it. No
man dare approach it unless his arms, legs ¿ind head be protected by clothesÏ

Osbeck, on a voyage from Siveden to China, stopped in east java and, on 20 January 1752, saw a tree with
flowers on its trunk. Cauliflory is unknoivn in northern Europe. He believecl he had found a leafless parasite,
and called it Melia ¡{Jarasitica (Fig. 1.2) naively commenting:

A small herb of barcly a fingefs length groviritig on tree trunks. It is so rare that so far as is known no one
ever saw it before‘;

“An intfodzïxcïfontito "tpfopicqvl rain forest;

Fig; 1.11. Eafly Europcanrravellérs brought back éxaggéráttd tales about njopíhcafrainforests. This
‘engraving fromfïofa Bfásiíiensis ofvon Martins, V .184Ü {plate IX)‘ is a scéñc‘ irï Íhé Atlantic coastufórcst.
of Brazil. " v- ' ' ' V '

‘Pág. 1.2; mmversábnirne 011 the ¿rank of Dyésaxg-rum 1

parasitícum, ‘something unkñbwn iri riortherñ"

Europe. Ñïhcn the Swedish botaníst Osbcck" savií rhiái . specíes_ in javahefichought th; flowers_werc a
lcafless *

parasitg. Solomon Islands.-

VUÍthÍÏÍQICIIÍaIpEEetratícJH scientific ápecimens“ í

began‘ to-fioodv; back to ‘the museums of Europe;

Atïfirstítheïïjlants ‘wérc*t}1é=:iveeds' pleíces, thá1íy'"of whichïhave ividáoécurrencc.‘

Iúdéed, fivhenïgibïïrjáéus‘ rinádkíhis _ greát syrithésis -‘ of ‘theïworld’s_ plantsglthe "Species pfanrafunyof

9'

1 753,11€ ‘belíéveci from this Jzvídence: ‘that; the

tropics hád. á ráther species-poor landïunífo'rm

flora.

An introdoctitirt to tropical rain ¡‘rn-tests ' 3

The tropies had a powerful influence on the ieveltiptnent of loitiltagyr in the HÍHÜTCEDEh century’.
Biogeogreiphyr and ecology are both founded on the iourneys in South America of me German Alexander
von Humboldt, in the Andes {where he recorded how. vegetation nges with climate) and in the loyvland rain
zoresrs of Venezuela. He travelled with the Érenchman Atimé Bonpland. They arrived at Camana, Venezuela,
on 16 July 1799 and the tft-ect of the tropical environment led von i-iulnholtit to write home:

That trees! Coconut trees 50-60 feet high; Poirtcicmt: yrzicfïettrínza? with a foot hi gh houq uet of
Itiagililieetlt

ïrzght red fltfrwers; pisang and a host of trees With :.;OFIT1ÜUS leaves and seenteti flowers, as big as the _
im of a hand, of which we knew‘ nothing . W7C z-¿sh around like the ClCIIIEIIÍBLl; in the first three dajrs
were unable to classify anything; we pick up one ¿ubjectto throw‘ it aiway for the next. Bonpland keeps
telling me he will go mad if the ivonders do not cease.

Perhaps even more important for the (lcvelop- ïnent of biology’ was the stitntilus the tropical rain forest gave
to the minds of Charles Darwin 3nd of Alfred Russel Wlallaec in their indepen- íent expositions of the theory
of evolution hy ïiatural selection. Darwin, as a young man 22 years old, “rent as naturalist on the voyage of
the Beagle, whose first trotaical ' landfall aras Seiivador on the Atlantic coast of Brazil

Fig. 10.11). He went ashore on 29 February,

Leap (lay, 1832 and has recorded:

Delight is a nyeak term to express the feelings of a ZÉElILlIallSÍ Villlo, for the first time, has wandered by
himself in a Brazilian forest. The elegance of the grasses, the novelty’ of the parasitical lalants, the heauty of
the flowers, the glossy‘ green of the foliage, hat above all the general luxuriance of the iregetatiotl, áiliecl me
with ¿Idmiration . The noise from the insects is so loud, that it may he heard even in a vessel JIIChOIECl
several hundred yards from the shore... . To a person food of natural history, such a day as this brings with it
a deeper pleasure than he can ever hope to experience again.

Wallace spent five yrezirs travelling in South America (1848-52) and then eight more (‘1854-62} "in the
Malay archipelago where he

discovered the two distínet faunas of the region, epitomizetl hy the houndary’ WallaCeÏs Line named zifter
him {Chapter 6). He too uvas impressed by the riehness of the forests:

If the traveller notices a particular species and wishes to find more like it, he may often turn his eyes in train
in every direction. Trees of varied forms, dimensions and colours are around him, hut he rarely secs any one
of them rcpeated. Time after time he goes towards a tree which looks like the one he seeks, but a closer
exaamination proves it to he distinct. He may at length, perhaps, meet ‘with a second specimen half a mile
off, or may fail altogether, till on another occa- sion he stumbles on one by accidenta-

Richness in species w215 one of the vivid dis- coveries of these nineteenth century’ explorer naturalists. it is
now belieirecl that about half the world’s species occur in tropical rain forests although they only occupy
about seven per cent of the land area. Herbs familiar in Europe have vroody relatives which gives a whole
new dimension to taxonomy:

Ñearly every natural order of ¡‘Jlants has here trees among its representativos. Here are grasses {hamhoos}
of 40, 60, or or more feet in height, stimetinles growing ercer, sometimes tangled in thorny thickets, through
which an elephant could not penettate. Ven-‘ains-g form spreading trees ivith digitate leaves like the l'l()FSC-
Cl1CStl1Llt.llJ Nlilkvirorts,“ Stout ivoody twiners ascending to the tops of the higher trees, and ornamenting
them with festoons of fragrant floyvers not their own. instead of your periwinkleslï we have here handsome
trees exuding a milk which is some- times saltitiferous, at others a most deadly poison, and hearing fruits of
corresponding qualities. Violets“ of the size of apple trees. Daisies (or what might seem claisíes} borne on
trees like alders.“

Tropical rain forest is eertainly very different from the vegetarion of northern Europe familiar to these
naturalists, and few were ahle to resist recording their impressions in lyrical prose, or to exaggerate (Fig.
1.3). In perhumid climates, on normal tropical soils and at its grandest, as in the western h/Ialay arcliipelago,
it is, to use von Humholdfis phrase, forest piled upon forest, the top-most trees 45 m or occasionally even
taller {Figs 1.4, 1.5, 1.6}, often as solitary

4 ' An introtïtteririiz to

«iuxr: ("ras ,t-i-r,.>.t'1:te m- HTIHS.

Fig. 1.3. The (Hang-titan ¿inf Sumatra and Bornco is in fact tlocile and shy and even if provoked is ¡note
likely to flee than to attack. This, the frontispiece to PoR. Wallaeeï ¡‘Vialnya Arcísipeltïgo, the European
reader the th rill he was expectin".

emergents which stand head and shoouldets above a billowitïg etintinuous Canopy, num): shades of green.
Yifithin the Canopy” there are trees of all different heights, “which scimetimes locally occur in layers or
strata (pp. 29-30), with crowns of tiaany shapes. "Trunks are mostly‘ slender with only a minority exceeding
a metre in girth. The trunks may be buttressed (p. 54}, and the bark variously‘ sculpttired and coloured (p.
57). But the forest is more than just a collec- tion of trees, as has been vividlyr described by’ E.}.H. Corner:

On its Canopy birds and butterflies sip nectar. On its htanches orchids, aroids and parasitie mistletoes offer
flowers to other birds and insects. Among them ferns

tropical? rat-n foreszs

creep, liehetis encrust, and eentipedes and Scorpions lurk. In the rubble that falls among the epiphytic roots
and stems, ants build nests and even earthu Worms and snails find homes. There is a minute ¡nunching of
caterpillars and the silent sticking of plant httgs. On any of these things, plant or animal, a fungus may bie
groiving. Through the branches spread spiders" webs. Frogs wait for insects, and a snake glides. There are
nests of birds, bees and vuasijs. Along a limb ptass: vvary monkeys, a halting Squirrel, or a bear in search of
honey; the. shadow of an eagle startles them. Through dead snags fungus and heetle have attached the wood.
There are fungus braclcets nibbled round the edge and borcd by other beetles. A woodpecker tapa. In a hole a
hornhill broods. Víïhere the main branches diver-ge, a strangling fig finds grip, a bushy epiphyte has
temporary root, and hidden sleeps a leopard. Ïn deeper shade black termites have built earthy turrets and
smothered the tips of a froung creeper. Ï-Ianging from the limhs are cables of lianes which have hoisted
themsclwres through the under- growth and are suspended by their grapnels. On their swinging stems grows
an epiphytic ginger whose red seeds a bird is pecking. Where rain trickles down the trunk filniy ferns, mosses
and slender green algae maintain their delicatc lives. Round the base are frag- ments of bark and coils of old
lianes, on ‘which other ferns are growing. Between the huttressroots a tor- toise is eating toadstools. An
elephant has ruhbcd the bark and, in its deepened foormarks tadpoles, mos- quito larvae and threadworms
swim. Pigs squeal and drum in search of fallen fruit, seeds and ttufflcs. ln the humus and ttndersoil, insects,
fungi and luaeterizi and all sorts of animaleulac participate with the tree roots in tlecomposing everything
that dies.“

During the twentieth century‘ knowledge of trenpical rain forests develoláed in two streams.“ On the other
hand academic scientists continued to collect and identify the plants and animals, and to describe forest
structure. Many of these studies were made on short visits of a few months’ duration. Theft tried to
comprehend the nature of forest irariatilon from place to place. This W215 in ziccord iwt-‘ith the
preoccupation of ecologists in temperare countries at that time with the tiaturc of clímax communities and
with succession. This phase of study culminated with the pubiication of a masterly synthesis by 13W.
Richards of the whole field up to about 1940.18

éaJ

An introduction to tropical rain foresïs v 5

zeeperldently, colonial fotesters began to

"imit blocks of forest to be presetved from ¿ing for agriculture, to control the utílizatiorl forests for timber
production, and to develop ‘ÍCIIlÍLIÏC. This last involved the application of me ccnturies old European
knowledge about what is today sornetilnes called ‘gap-plïase ijrnainics’, nznnely the ability of different
species to tegenerate after different degrees of Canopy íísmtbalïcc.

ln recent decades the two separate streams have mergecl to lead to the new synthesis Which is a nmjor part of
the present book.

Fig. 1.4. Lotvland CVCIÏgTCCII dípterocarp rain forest in profile (Brunei), with ¡Diptercrcarpus crirzítzts
right centre and Shores curtisii to its left. W itho ut the scale object the reader would not realize the huge size
of these trees.

Apart from the growing knowledge of tropi- cal rain forests in the Western scientific tra clition it Inusl: not be
forgotten that within the tropícs man has lived close to Nature and in intimate contact with tropical forests for
millennia. The forests yielded all the products Iieeded for his life, and he learned how to grow crops on inher-
ently infertile rain forest soils, by shifting agrí- eulture, moving the fields every 2 or 3 years and allowíng
forest regrowth to restore site fertility. This discovery was Inacio independently in all parts of the tropical
zone. His numbers were never large.

6 An introduction to tropical rain foresrs

: gravamen

.- J -:.r.r,u'.- . ,1 , , s. , __,,.*. - Jr"- _ -,,. . .-

.-' ;¡- _.-_v. _.— - »'_a.__. _,

v- . ’ ' .. . n.‘ . .

Fig. 1.5. Profile diagram of lowlaintl evergreen dipteroeairp rain forest, Brunei, Ridge crest plot 60 _>< T5
m, all trees over 4.5 m tall shown. Nlature phase forest except for extreme right hand end. {From Ashton
1965, in Vllhitmore 198m1, liig. 1.6; see latter for species‘ identification.)

The era of European exploration of the world

ftxlltiwed by the Industrial Revolution led to increasing human impact, which has increased continually till
the present day lFigs. 1.7, 1.9). The evolution of medical knowledge in the West, plus the development of
powerful drugs, has this century removed most of the health hazards of the humid tropics so that death rates
have diminished and life expectaney increased,

both dramatically. This has added the ¡‘ICW pres- sure on tropical forests of much higher and rapidlv
increasing human populations (Fig. 10.10). The teehnological development of reli- ahle machines for road
building and log hauling, of chain savvs for tree felling, and of hulldozers for land-clearanee since Wltirld
Wlar Il have made it {Jossihle to remove tropical rain forests on a scale and at a rate that “ras previously

An ÍflÉfOdítCÏÍOn to tropical rain forvests 7

ramátmrwwua ww’ e‘
Fig. 1.6. Emcrgcnt kapok, Ceiba pentctridra var. Caribes ; riverine forest near Iquitos, Peru. Subsequently all

fellccl to supply a short-lived plywotid industry. This species is one of the biggest and commonest emergents
of the Amazonian rain forests {Gentry and Vasquez 1988) and occurs also in Africa (Fig. 3.31].

Fig. 1.7. Lovvland semi-evcrgteen rain, forest penetrated by logging road, with S. Cotitinho (1985). Lower
Amazon, jari, Brazil.

decadtï aigopfvare . noivï start-agraria an _ ble; Everrïonljá - _fo':t'y' tyears‘ ago, ‘tvlien 1113i" ‘son

Íïïiitláñd. iïfiíiïïïil.”‘ïfelïícs=- i

Odvs_sey._ ïbegaia,- ‘jtropicalï irainfï ¿forests boundless; I ‘descended "the: Amazonas; '_flai1_1¿._.o

theïAndes in- zEcuadorppon mulebaek‘ Visio iiiïiïïáti‘—i"

gate montafies- forest ‘zonation, pa; journey .

_nofi_viïïtali<es*ari'_ hour} b? gbus; sThen‘Vfilftravelled {ividelv ¿throughÏÍhei-i-Soloïhïíñs in‘ the;

«wesïtesrn- ÏPacificÏ ibyiz; schoorierg} and collected“ arre never ‘seeniljbyrgscience ‘es ‘sparsely inhabired:
islands "xvheregtheÍ rain ‘forest; cameï downïipto" tzheïvild íeoconutsfleáningljoverfithej‘sandy. beach.i_¿_
_

ter ‘I-Ï

._.Aérïairodaéeaaï.te¿vastas gar‘ ‘¡aretes

ï:_.herhs,‘7_h‘ere the”:artiidjColocásia"giga téaÏat i

esksïtzréstïsgijth "iivaiiérsljed asas -oné"'tiiés las .. thoselandsifrcïm Loriclonïrn onljrna. ay o ser"

and ¿sanearsdayïfbgtïajn the: ‘parche go ¡uggie

ïisvhichï" remain; ¡a rriving bj? durïíp truck " long v va 1 s1addy-L__Icgging?re‘adirtherezi;¡usage

eralfiandï; scientific" interest in ?t:o'pic¿1-=raiín ‘jforestsi’ "thaiïrj ever "before anclith_is__ne\v ease
ofïáecess iisípariïofï 7th'e'_‘reas_on. soíletjus:n'¿ss¿¿_'tt1rn__is;Chapter-z to

close} examina tio n“ of "¿the tropical" forest: zone: ‘¿ha then- jcontinueï: __‘withz ¿analysis ¿‘of its;
yplants; animals, ‘dynamiceaad ’ present status. .

Fig. 1.9; Lowiand evergreen

beside anewly built telecommu- nieations toweraccess-track, jMalaya. Emergeut ‘Camarium and Sborea but
no other clistinet strata. Note ¡naty

An introduction to tropical rain forests _ -r ‘ "9

dipreroearp rain forest in profile, ._

2i
What are-tropical rain forests?
2.1. TROPICAL MOIST FORESTS AND
THIÏIR CLIMÁTES

The German latitanist AJÏW. Schimper was one of the great nineteenth century naturalists. In. 1898 he wrote
21 monumental book, translated in 1903 into Fvnglish as Plant geogrïzpby upon om ecologicai imsis, which
built upon the hundred years and more of European scientific cliseoveries in the tropics. To Schimper we owe
the term tropical rain forest {trojiisciie Regenwaid) for the forests of the permanently wet tropics. He
recognized in total ftiur major sorts of woody vegetation in the tropics. Rain fotests, then in progressively
drier and more sea- sonal climate, monstiaon forests, savanna forests, and thorn forests. In still drier climates
non- wotidy vegetatioti, tropical grasslands, and deserts, were recognized. This classification is still useful as
an outline framework in all parts of the tropics. For most purposes Schimpefs great groups provide too eoarse
a classification

- of vegetation and it is useful to hat-‘e more fineiy

defined classes. These are called fotmations, and in all six of Schimpefis groups several can be recognized. A
vegetation formation is defined on its structure and on the physiogrlomy of its com-

-’ ponent plants. For tropical forests the structural

properties include the heightof the trees, whether they tend to have their crovvns in layers, and the presence
of different kinds of

climbers and epiphytes. The phvsiognomic prop- erties include whether the trees are buttressed, their crown
shape, the nature of the leaves, (size, shape, thickness, margin}, whether the forest is evergreen ot, if not,
how stronglv deciduous, and where on the trees flowers and fruits are borne. Using these criteria different
forest for- mations can -be defined and are found to occur in many different places. _ Amongst the various
tropical rain forest for- mations are the most structurally complex and diverse land ecosystems that have ever
existed on earth, with the greatest numbers of co- existing plant and animal species. Ïn species richness they
are only rivalled by coral reefs. These lofty forests are the eipex of creation.

Climate

Constant high temperature is charaeteristic of

tropical climates, and climates in which the‘

mean temperature of the coldest month is 18 °C or over, is often used as a definition. This excludes ¡some
tropical montane areas and a dif- ference of less than 5 “C between the mean tem- peratures of the warmest
and eoidest months is sometimes used as an altfifnéltiVft definition. Rainfall is the second most important
factor, and its amount, and even more important its distribution through the year, define different
tropical climates. Night is the winter of the:

Tropical ¡worst forests and their climares 11 _

tropics, hecause the diurnal range of tnean daily temperature eXceeds the attnual range and is greater in the
drier months.

Rain forests develop where every month is wet (with 100 mm rainfall or more), or there are only short dr)’
periodsywlïich occur mainly as LIUPICClÍCÍE-‘Ible spells lC-Lsïllïg only a feu-Kcltty-‘s or weeks. Vïlhere
there are sex-‘eral dry Inonths [60 Intn rainfaïll or less} of regular oceurrence, monsoon forests exist. Outside
Asia these are usually called tropical seasonal forests. They are slightly less lofty but with more even Canopy
top, and the lvigger trees are ntostlv deciclututis, although perhaps only hricfly’. They have fewer dependent
climhers and epiphytes, and elreless species rich.

The blanket term tropical moist forest Was coined in 'I9?63‘l to cover both rain and ¡nonsoonlseasonal
forests, Schimpefs first two great groups, which together comprise the tall, closed-cantipy forests of the wet
tropics. Tropical nmist forest has proved a useful group- ing for einalyses of the rate of forest tlisappear- ance
(which will he discussctl in Chapter 10), because the exact houndary between rain and Seasonal forests is
hard to pinptlllït and because individual natíons seldom tlistinguish between them. To the laiologist, however,
there are major differences, and this book is about tropical rain forests, those which occur in the everwet
(perhu- mid) climates, with onljv passing mention of nlonsoon forests.

The climates. in various parts of the rain forest helt are shown in Fig. 2.1 using the useful picto- rial tlevice of
Climate Diatgranis ClÉVlSELÏl by ‘Xïalter and Lieth.“ Note the tlistinction, even

_withii1 the rain forest belt, between places with

no dry season (eng. Inuch of the lndo-hïíalagran forests and the western Anlíïztïlï), and those where there is a
slight dry Season (eg. Wfest ¿áfrica and the eastern tomazon). One savanna forest site is also shown {Telmale
ln Ghana) to tlenmtistrare the much lower and less evenlv dis- tributed rainfall of the seasonal tropics.

The (jlimate Diagrams depíct average climate. For plants and animals extremes are often ¡note important
tiran means, even when onlv of rare

Occurrence. For rain forests tinusually prolonged dry periods are of particular significance, for exaunplc‘ as
triggers for flowering (section 3.3), or because dtought can eictuallv kill plants or make the forest flammable
(section 7.8). Porests, because of their stature, have internal tnícroclimates that differ from‘ the general
climate outside the canopv. These are tliscussed more fully in section 7.1. In general terms, it is cool, ltumicl,
and dark ttcar the floor of a mature patch of forest, progressively alteritig upwards to the canopy top.
Different plant and animal species have specialized to the various forest interior microclimeltes, as Will be
seen lÜClOVk’.

Occurrence of tropical rain forests

Tropical rain forests occur in all three tropical land ¿[C3522 llïíg. 2.1). Most extensive are the Attiericaïi or
neotrtipical rain forests, about half the global total, 4 x '10“ km2 in area, and one- sixth "of the total broad-
lcaf forest of the urorld.“ These occur in th ree parts, of which the largest lies in the Amazon and Orinoco
"hasins. Second is a block which lies across the Artdes on thefacific coasts of Ecuador and Clolomlaia,
extending northwards through middle America as far as Veracruz in southernmttst Mexico (19°N). The
Atlantic coast of Brazil has a third block, a strip less than 50 km svide on the coastal mountains, extending
beyond the trUpÍ-CS to the vicinity of Rio de janeiro (c. 23°C). Totlay only about '12 per cent remains“ (Fig.
10.11). It merges southwards into suhttopical rain forest of sirnpler structure and different flora, but in a wav
we shall never now be able to define. Brazil is the nation with more rain forest than any other. .

The second largest block of tropical rain forest occurs in the Eastern tropics, and is esti- rnared to cover 2.5 x
10“ ltml. It is centred on the Nletlay archipelago, the region known to bütülllsts as Malesia. Indonesia”
occupies most of the archipelago and is second to Brazil in the amount of rain forest it possesses. The Eastern
rain forests extend beyond lvíalesía into the Pacific and southwards as a narrow broken

12 lll/hat are tropical rain forests?

Belem
2277

—-—_.__fi . ‘Ilill

AFRlC
RAIN EST

Ámmflflmh Doula

- 3948

Balehonnur

¡Nno- ‘¡iia a MALAY "t 7 saw EST

The forest formations 13

coastal strip in Queensland, northeast Australia, in a similar position to the Atlantic coast forests of Brazil. As
in Brazil, this forest extends heyond the tropics, southwartls into New South ‘ïïales. Close study of the
merging zone has shown that towards their temperate limit tropi- cal rain forest formations Become
increasingly restricted to the wertest sites and the deepest, most friable, and most fertile soils?“ The X-
lalesian forests extend northurards up the Íxlalay peninsula into continental southeast Asia, Burnra, Thailand,
and lndo-China. Again, :.t the botmdary they probably occur as a mosaic and occupy the ‘best’ sites, though
we iack tletailed studies. There is also an ill-defined change to suhtropical rain forest. Between 92° and
9891:, bClUW’ the great south wall of the Himalaya, in southern China, Assam and upper Burma tropical
rain forest reachcs further away from the Equator than anywhere else, attaitiirtg ZS°N37 There are further
outliers, in southwest Sri Lanka centred on Sitiharaia, and also on the Western Chats of India where
seasonality is strong.
Africa has the smallest block of tropical rain

forest, 1.8 >< 10° km3. This is Centred on the Congo hasin, teaching from the high mountaius et its eastern
limit westwards to the Atlantic Ocean, with outliers in East Africa. It extends as a coastal astrip into Y-(lest
Africa, but wood- lands reach the coast at the so-called Dahotney Gap.” There are tiny parches of rain forest
on the east coast of Nlaclzigascar and in the Ïvíasceirenes. Outside the Congo core the African rain forests
have been extensively’ tlestroyed.

2.2. ‘rue FORFST FCHUVLATIONS

From a satellite or high flying aeroplane the

earth is seen to have a stimbre, dark, blue-green girdle about the Equator, just glimpsed through breaks in the
cloud. There is considerable varia- tioti from place to place in this rain forest mantle due to the distinct forest
formations. A forest formation,'as described above, is reccig- nized by a particular combination of vegetarion
structure and physiognontiy’ regardless of flora. (Éonvergent evolution has occurred because dif- ferent
species in the three land areas have evolvetl similar responses to particular environ- tnents. In many cases we
still do not understand what facer tuf the environment evokes a particu- lar response, but sometimes we do,
as is dis- cussctl further in relation to features of montane and hcath forests in sections__8.4, 8.5’.

The forest formations occupy‘ different physi-

cal habitats and these are mostly’ sharply bounded, but where this is not so there is a Inerging zone. '

The formations can be grouped for conve-

nience aiccording to the main physical character- istics of their habitats (Table 2.1). This is an arbitrary
arrangement but gives a useful hierar- chy. The Iiaming of vegetation types is always problematical. In the
case of tropical rain forest formations the names (listed in Table 2.1} reflect site and the structure and
physiognomyr of the formation.

The first division is between clímates with a

dry season and those that areperhumid. The

Fig. {Left} Tropical rain forest, distribution and climate. {After Wfliitii1ore 1983, 19845:; ‘Walter and

Licth 1967.]

The (Élimzite Diagrams show dry periods as dotted and rainy pcriods as hatehed {or ttïhere monthly rainfall
exceedslüü mm shown black at 11'10 scale). Long-term mean annual rainfall (mm) is shown in figures.
Exrergreen rain forest occurs in perhumid climates, i.e. virtually all of the Indo-Malayran region, the upper
Amazon and Pacific coast South America. Ït is replaced by semi-evergreen rain forest {lljfillnd the fringes of
Indo-Nlalaiya, and in the lower Amazon and in Africa, itichlding the Congo basin, where dry periods oceur
irreguiarly scattcred through the year and do not show up on these diagrains because they are based on

long-term means.
14 - ‘What are tropical rain foresrs?
Table 2.1 The formations of tropical rnoist fotests

Cllfllálli.‘ Soil water Soils l Elevation n Forest fommtion Seasnmaily Strong annual híonsoon forests city
short-age {vziriotis formations] i’ Ii‘

Sligiit annual ' Rain fnrests‘:

shertage Semi-evergrecn rain forest Everwiet Dtyiand Zotml Lowlands - Lowland evetgreen rain forest
(perhumítlïi (mainly: oxísols,

{ultisols} 7 (7502 1200-1500 m 4 Lower monturas rain forest 7 i600] TSÜÜ-EHJÍJÍ] m Upper ¡zic-ntaite
min fui-est fi (33501 m 2‘ SÜUÜ [3350] I'll to 1 Sub-alpine. forest tree iine Podznlized sunnls lkïostly
lon-‘lancls _J Heath forest Limtzstone Mosrly lowlands Fun-est ox-‘t-r Iimestonc Ulttalmsic rocks Pt-‘Iostiy
lourlnntis Forest over uilrzlhasiczs . , ‘i’ "‘

Water Coastal Beach vegetzitiun

Table salL- Mnngrove forest

high Eat ivater Bruckish ivatet‘ ftarest

least [__ ___l

perimii- ln land (Jligotrophic pczits r Pezu; swamp fui-est

rally] fresii- _í

WEIIEI‘ Euwophie {muak and ¿Plïrlnïillfintly wet. Fresllwater Swan-tp forest ‘ mineral} soils y , L _
PWÏUÚÏCBHY Wet Freshivatcr periutiie sivamp forest _l_

Those. shown [mltl art‘ diseussed. in the ¡ex-t

second division is a erude measure of soil RVEIICI’ availability, and distinguishes swamp from dryland
forests. The third clivisitiii is On soils and, tvithin drylantl forests, distinguishes those on parent materials
with atypieal properties- peat, quartz sand, limestone, and LlltrabélSic rocky-from the widespread ‘zonal’
soils, mainly ultisols and ÜXÍSOiS. Finally‘ there is a division of the forests un zonal soils by altitutle.

In this scheme a few formations are defined by physical habitat as Well as vegetatioti struc- ture and
physiogntiniy‘, for example, forest over limestone. The overall result is a set of easily reeogniztztl forest
formations Keith memorable names that say something about the intist dis- tinctive characteristics of the
formation. It is a pragmatie scheme of rain forest classification and the formations can he recognized
through- out the humid tropies HlÏlÏOUglÏ many different regional names have Been applied. For example,

in Brazil fresh-tester swarnp forests are called irarzea or igapo.

This [wok is only a primer or introduction to rain forests and tiescriptions are given just to the more extensive
forest formations.

Tropical lowland evergreen rain forest

Description

(Figs. 1.4—1.7, 22-24} This is the. most luxuri- ant of all plant communities. It is lofty, dense, eirergreen
forest 45 m or more tail, eharaeterized by the large number of tree species which oecur together. Gregarious
dominante (eonsociations) are tmeomintin and tisually ttvo-thirtls or more of the upper-Canopy‘ trees are
ofspeeies individ- ually not eontributing more than 1 per cent of the total number. This formation is
convention- ally regartled 21s having three tree layers (see

‘Tim forest fo-rrrzttrions 1

Fig. 2.2. l_(')\.’\’lí_iI1Cl]Ï¿iÍI1 forest WiIl1lUXUlTlflJ1iÏ aroids; XÏCMISÍBFT! cleiiríosr; icheeseplant, left)
and Plvilodeizdrtuzz sp. (right), showing their Cord-like pendcnt TIJUÍS. (Kerner and Oliver 1895, Hg. 423.)

pp. 29-30}: the top layer of individual or grouped giant emergent trees, over a main strarurn at about 24-36
m, and with smaller, ishade-tlivellíng trees below that. Ground vege- tation is often sparse and mainly of
small trees; her-bs are patchy. Some of the biggest trees have clear boles of 30 In and reach 4.5 in girth, and
may be tleciduotis (Fig. 3.20) or seini-deciduous without affecting the evergreen nature of the Canopy as a
whole. Boles are tisually almost cjrlindricai (Fig. 3.29). Buttresses {Figs 331-333} are Common. Catiliflory
fFig. CLZ] ¿nd rarniflory íFig. 5.2) are Common features. Pinnate leaves

‘un

(Fig. 2.5) are frequent; ieafblades of ntesophyll size (Fig. 3.26) predoniinate. Big i-voody climbers (Fig.
3.15), rnostly free-hanging, are frequent to abundant and sometimes also bole climbers (Fig. 3.17). Shade and
sun epiphytes (Figs. 3.21, 9.1) are raccasional to frequent. Bryophyres are rare.

All other rain forest formations differ from this in hairitig simpler structure, sometimes ivith fewer life forms,
and fewer species.

Hahitat ¿md OCCHÍÏEIICB

Tropical lowland eirergreen rain forest occurs in perhuniid lowiand ciirnates “there water stress is
intermittent or absent, from sea-level to c. 1200 m on. dryland sites. It is the main lowland formation of the
Eastern tropics and also occupies western Amazonia plus the Pacific coast of South America. In Africa it is
restricted to three small blocks near the coast of West Africa, between Guinea and Liberia and Cameroon and
Gabon?” On the Climate Diagrams (Fig. 2.1.}, all sites with this formation have no regular einnual dry
season.

Tropical sembevergrecn rain forest

Description

Semi-evergreen rain forest is a closed, high forest in which the biggest emergent trees some- times attain 45
m in height. lt includes both evergreen and, in the top of the canopy, decidu- ous trees, in an intnnate mixture
but tvith a definite tendency to gregarious (IICCLIITCHCE. Deciduous trees may comprise up to one-third
of the taller trees, though not all are necessarily’ leafless at the same tirne. The number of species is high, but
less so than in etïergreen rain forest. Buttresses continue to be frequent and occur in both evergreen and
deciduous species. Bark tends to be thicker, and rougher and cauliflory’ and ramiflory rarer. The stature tends
to be slightly less than CVCrgÍCEfl rain forest with emergents as seattered individuals which are smnerimes
rare. The Canopy is sometimes locaily stratified. Big woody climbers tend to be very

16 Wfïmr are tro-¿‘Jícal rain forests?

Fig. 2.3. Loivland evergreen dipterocarp rain forest on hill slopes, Selangor, Malaya, at about 800 m

clevation.

The giant cmcrgent trees 50 in or more tall are Sinn-ea cortïsir’, clumped on spur ridges, and showing the
catilifloisrerwlike crowns typical of tnany Dipterocarpaceac.

Fíg. 2.4. Lowiand rain forest, -

Caroní basin, Venezuela.

i This forest has no emergent trees, the Canopy top is fairly flat, and the crowns of the numerous species are
nested closely together but mostly ‘with a small surrounding space.

The forest formotions . 17

Fig. 2.5." Pinnatc leaves are a common feature of lowland rain forest; Canarium sp. Guandong, China.

abundant. Bamboos are present. Epiplïiyres are

oceasional to frequent and include many ferns i

and otchíds.

Habitat and occnrrencc?"

‘Where there is a regulan‘ annual period of mois- ture stress, either due to rainfall seasonaliry’ or particular
soii conditions, this replaces the previ- ous formation. It occupies most of the African

rain forest block, including the tvhole Ctmgt) basin. itoccurs today as a narrow fringe around the main
Eastern rain forest block, and also forms the Outlier in india, and most of the Australian. tropical rain forest
(Fig. 2.6). Ir veas probably more extensive in pyrts of continental southeast Asia in the past but has been
reduced to its present extent by human eictiirities. This is probably the formation that occurs in middle
America. The eastern and southern Amazon (1.8 X 106 km2, about. half of the forested iarea)‘ cio not havea
rain forest climate". The dry season is extremely strong and may last five months. Sembevergreen rain forest,
rather than a tleciduotis monsoon forest, occurs because the very deep soils act as a reservoir and are tapped
for trater by roots that reach doi-vn to 8 rn depth.“ This is probably the formation that OCCUFS in middle
America. The northernni'ost rain forests below the Hirnziiztyar, aswell as the south China Xishtiangbana rain
forests, are mainly the evergreen formation.” The occur- rence of a dry season is elearly shown in the Climate
Diagramas of Fig. 2.1, except in the Congo basin where dry ijeriods are so irregular in occurrence that they
do not show up on these iong—tetm mean records.

Fig. 2.6. The rain forest rnargin, northeast Queensland, Australia. Lowland semi-evergreen rain

forest in the foreground sharply mcrging into wet sclc- rophyll forest with emergent Eucafypttrs grairdis in
the background. The view is looking wcstwards, away from the coast, to iucreasingly‘ dry climates.

18 ' ‘Nils-at are tiropícal rain forests?

The montana rain forests
DEJSCTÍÏJÉÍOH and frabilat” fhigs. 23-110)
As one clirnhs a tropical mountain one SLICCES-

sively encounters forests of different structure and physiognt)ni}-' (Table 2.2). The most dra- matic change,
ruhieh tisually occurs over a short distance, is from mestiphyll-dtïminated forest with an uneven hilloviring
cantipg; surface to a lower, lnore even, often palevcoltittred, micro- phyll-dominared Canopy, of more
slendet trees, usually with gnarled limbs and ver}! dense sub- crowns. This is upper tnontane rain forest. It is
clearly distinctix-‘e hoth from the air and to "the trEWCllCl’ on foot. This formation is often (July 10 III tall
or less, and its shorter facies are some- times called elfin Woodlzmtl. The trees may he heavily Swathed in
bryophyttts and ñlmy ferns and the formation is then known as mossy forest (though-liverworts dominate).
Peat often fiorms, sometimes with the laog-moss Spffihïlgïlïfln. On small mountains upper montane rain
forest

ahuts on low-land rain forest, hut on bigger ones an intermediate formation, lDWEI‘ inontane rain forest,
occurs and this has 21 broad eeotone against the lowland formation. On the highest peaks, upper montane
rain forest itself is replaced upvrards hy a shorter more gnarled for- mation svitlïi even tinier leaves
(Halïüplïyllsl known as suhalpine forest, of which good descriptions exist for New Guinea, hut ‘Nlficlï is
much more extensivo ¡u the Atides. The tree line on the biggest moiintains is at c. 4000 m. It is often
clepressed by fire. Above it, heyond the cli- matie limits of trees, occurs alpine vegetation, an edaphieally
determined mosaie of shrub heath, mtJss tundra, fern I11Ca(l(')\'u¡, ¿Uld grassland, called par-amo in the
Andes. These treeless landscapes extend up to the snowline at c. 4500 m.

On large mountains all the forest formations

occur to higher Elfilïéïtiolï than on small ones and Outlying spurs. This is the so-called hïíassener- ltebmtg or
mass elevatitm effect, first tleserilved for the Alps. III the tropics the compressitm and

Table 2.2 Charaeters of structure and phjrsiogtiomy’ used to define the principetl montane forest

formations Formation Tropical lovvlaand Tropical lower Tropical ilpper‘ et-‘ergreen rain montane rain
montane rain forest’ ' forest forest Canopy height 25-45 m i 15-33 In '1.5—'l S m. Emergent trees
Charaeteristic, to Often ahsent, Usually ahsent, ' SUMO} m tail to 37 m tall to Z6 m tall Pinnare leaixes
l-‘reqtient Rare Very rare Principal leaf size Class of Mesophyll Nlesophyll Microphyll

woody plants:
Buttresses Usually frequent

and large Cauliflorjv’ Frequent Big woody climhers Abundant

Bole climhers Oftcn ahundant

Frequcnr
Oecasional

‘tlascular epiphytes Non wascular epiphy tes

Frotïrvvïhitmore {198421} tahle 18.1

Ïlncluded for comparison
TFollowing Raunkiaer {1934}
Uncommon, small Usually absent

. Rare ¿‘khsent

Usually‘ none None

lïrequcnt to Ver}! few ahundant

Ahundant Frequent
Oceasitanal to. Often ahundanr

ablJÏ-Idfint

The forest formations

19

Fig. 2.7. Forest zones on the main mountains of Malaya. (Whinnore 1984a, fig. 18.1.)

21m Forest formation ' Fluristic zune _ Important groups

u M t [Inniierae --——---- : n ¡me -

18W mpuiiilirane erilzaceuus Enraceae l

Myítaceue l
IEÜÜ l
E Hair-laurel Fagaceae, Lauraceae
E: 120D Lower —
E "wnlane Upper . Shores pfatycladaszS. erïiata. l

fi SUD dipteracarp ïarara, flrpferacarpus retusus 50° Hill As below + LUWÍBBÜ dipterucarp Sfiarea
¿‘HFÍÍSJÏ l

3m Lgwland Numemus dipterucarps especially Dípreracargus

a dipterocarp spa. Shores spp. and flryafiafanaps aramatrca

Fig. 2.8. Upper montante rain forest, Mount Bcnom, Malaya. »-

Note flattish Canopy top, and trees with dense subcmwns of crowded foliage and small leaves. The big trees
are Leptospeflnzrvn flavescerts; their gnarled limbs are another distinctive feature of this forest formation.

20 What are tropical ratio forests?

Eig. 2.9. Leptospertnzam ¡‘lauescens {Krith an under- growth of low bushes, ferns, and sedges. A dry, open

facies of upper montane rain forest; Mount Benom, Malaya. Lord Cranbrook for scale {i967},

depressiort of the forestbelts on small moun- tains seeins mainly associated with loweritig in the level at
which cloud hahitually forms.

e The changes in forests and their eniriron- ment with elevation are aualysed further in

- section 8.5.

Occurrence

The Eastern tropics are very mountainotis and many descriptions exist of the extensive montane forests and
their zonation.“ There are fewer descriptions for the Andes, although the complete series of formations
occurs on both flanks and over large areas. The fullest studies have been made on Jamaica in the
Caribbean \Vl‘l€¡'B-Z0l‘lí1tl()n is compressed. Montane rain forests are least extensive in Africa, where they

Fig. 2.1i} Upper montane rain forest, Mount

Nokilttlaki, Sulawesi, 2275 rn elevarion. The trees are swathed in film}! ferns and hryophytes. This facies is
often loosely called ‘mossy forest’, though the bryopluytes are mostly litterworts.

occur in Cameroon and at the eastern fringe of the Congo river hasin. Africa has the most eX- tensive
seasonally dry tropical montane woody ‘vegetation lforests, hushlands, thickets, and shrublands), whose
description is beyond the scope of this book.”

Heath forest,
Description (Figs. 2.'l 1-213)

Even the luotanicalljr incxperienced casual ‘wanderer xirill notice the change tvheu, he enters the kerangas
forest the store}! formed by large saplings and small poles predominatcs and forms a tidy and orderly but
forhidcling phalanx which is dense and

The forest fomtazions _ 21

Fig. 2.1 1. Heath forest with very even c-anopy top and small-crowns, interdigitated with lowland evergreen
dipterocarp rain forest in interior Sarawak.
Fig. 2.12. (Below left) Open stunted heath forest at Bako, Sarawak. The tree with feathery foliage massed
dense subcroxartis is Casrzarirstr nrfiailfis. See Fig. 2.13.

Fig. 2.13. {Below right) Calopbyllarm incrassarzmz in the Balta heath forest, shoviring its characteristic
shiny, vertically-held leaves. Many species in heath frarest have thick, leathery leaves (sclemphylls) like
these.

Figs. 2.12, 2.13 both show species with atlaiptations to rninimize foliage water loss and heat load (see section
8.4}.

,3; .-

mas.

22

lll/hat ¿ITE tropical rain forests?

Fig.i2._14. h/langrotre forests are a forest formation that occurs on accreting coasts in all parts of the tropics.
They have several peculiar features of structure and phjirsiognoniy, for example stilt roots (seen here on
Rhízopltora. apiculatn in Malaya}, and viviparous reproduction. They’ are floristically much poorer than
most other forest formations, with one family (Rhizrnyhoraceael predominant amongst the trees. Mangrove
plants have specialized physiologicail mechanisms to enable them to live in salt water. hïangrnpves are
disappearing fast as land is reclaimed for building (eg. Singapore), or for prawn ponds or salt pans. They are
the breeding ground for the marine life on which coastal fisheries depend and it is being increasingly rcalized
that their destruction has serious repercussions. Fuller accounts can be found in Clough {i982} on salt
balance, Lugo and Snedaker (1974) on ecology, and Tomlinson (1986) on taxonomy.

often difficult to penetrate. The Canopy is low, uniform and Lisually denselyr closed with no trace of
lay-‘ering hrownish and reddish colours prexrail in the foliage of the upper part of the Canopy and {the
forest) appears considerably brightcr.

Thus wrote E.F. Brunig, foremost student of

the Asian heath forests, about the ¡’salto National Park, Sarawakdf He also recorded the predominance of
microphylls over mesophylls, and that many leaves are sclerophyllous, often held obliquely or vertically,
often in dense clus- ters, and either waxy grey or highly reflective.

Big svoody clirnbers are rare but not slender

wiry ones. Epiphytes are common. Myrme- cophytes (Fig. 5.1i} and insectivorous plants may be ahundant,
including in Asia the pitcher plants Ne-pvenrlaes (Fig. 4.7).

Iïafritar and occurrence

Heath forest occurs ou soils developed from
siliceous sand, either coastal alluvium or weath- ered sandstones. These become podzolized. They are low in
sesquioxides and so are laoorly lauffered, and are highly acidic (pH less than

The forest formatíons . 23

Fig. 2.15. Steep Iimestone hills, here in Malaya in ‘the form of tower karst, are extcnsixfe in the Eastern
tropics and Caribbean. Linjestone cárries a clistínctívc forest formation with several very different facies; for
example, ‘ _ sheltered cnclosed valleys are tich inherbs {especially Gesneriaceae} and epiphytes; and
‘exposed summits, which‘ are prone to desiccation, have a stuntccl Xetomorphieforest with resemblances to
the heath forest for- ‘ mation. For fuller details see Whitman: (1934a).

4.0). They‘ are frequently of coarse textute The most extensive heath forests late in the

= freely clraining. The streams dtaining heath upper reaches of the Rio Negro (appropriately

forest are black, or tea-colourecl. When viewed named) and Rio Orinoto in South America. In. by‘
transmirted light (due to the presence of Brazil they are knowvn as canïpina, camp- organic colloicls), and
are usually acid wítha inarana, caatinga Amazonica, or campina low cation content. . l rupestre. There has
been intensiva study of them

at-the famous San Carlos research centre in southern Venezuela, but they only represent 6 per centïif the
Brazilian Amazon rain forests. The tlescription of*the Guyana heath forest dominated hy vvallaha, Eperua
fizllcata, by T.A. W. Davis and PNC’. Richards, is one of the classic works cif rain forest ecologyfá? There
are extensive heath forests in Kalimantan, Sarawak, and Brunei in Boi-neo, where the)! are called kerangas.
The other principal occurrences are small areas in Malaya, and on coastal sands in Africa in Gabon,
Cameroon, and Ivory Coast.

Heath forest is perhaps the most strikingly distinctive lowland rain forest formation. There has been much
debate on the extent to which its special features are correlated with either nutri- ent status or ivith water
relations. The extremely acid soil is probably toxic to the rcaots of many plants including crops. These topics
are dis- cussed further in section 8.4.

Peat swamp forest

Descripvtitnv:

This is a forest formation defined more on its special habitat than on structure and physiog- nomy. The
Eastern peat swamp forests have a limited and distinctive flora.

Habitat ¿md caccurrence

In parts of Sumatrza, southern Thailand, Nlalaya, Borneo, Mindanao, and west New Guinea a phvsiographic
setting exists which has favoured the formation of peat. Since sea-level rose at the end of the last Glaeial
maximum (section 6.2), the rivers have deposited sílt as levées and on Hood plains. Sivamps developed
behind the levées and became less saline as their soil level was raised by continuing siit deposition at times of
high river level. Mangrove became replaced by inland species whose litter failed to decay in the still salty,
high sulphide, waterlogged condi- tions. This litter turned into peat vahich has con- tinued to form up until
the present day. The result is that now there are domed peat swamps up to 20 km across which are oldest in
the

>24 , - ‘i ' . - lÏ-Ïhat are tropical rain forests?

Fig. 2.16. The highly developed pear su-uamp forests of Sarawak are domed and hear a concentric series of
different forest types (1-5) from edge to centre. {Vïhitmore I984a, fig. 13.5). A core taken through the peat at
the dome ccntre contains pollen of the suecessivc communities, with mangrove at the base. This
CÏCUIOIISIHIÍES that as the dome gets higher by peat accumulation each community‘ is replaced hy the
HCXE in sequence.

centre. These carry up to si}; different forest types; developed concentrically on the dome íFig. 2.16), each
occurs at a particular height above the water table. The innermost type is an open, stunted forest. The outer
forests reach 50 m tall and are a ‘Kaluable timhet’ resourcefï“ Here some silt has been washed in. Further
from the margin all the Iiutrient input has heen from rainfall and the peat is extremely oligotrophic and acid.
The peat, which is semi-liquid, reaches 13 m deep below the most developed dornes. Agriculture is
impossible on peat over 1-3 rn deep and forests should be retained. Unfortunately, Indonesia eliready‘ has
failed examples of the ill-conceixred conversion of deep peat to agricultura” Lowland peat svvamp forests in
rain forest climates also occur in the Americasfiincluding the pegass swamps of Guyana and parts of the
Amazon hasin (eg. Tupinamhara island east of Manaus), as well as in Africa (Kenya, Uganda and Congo) hut
these have not received detailed study. The total arcas of tropical peat are ZZ >< T04 km3 in Asia,

Forest marntennnceethe growth cycle - _- ' i" . . _. _, 25-

,7 X 104 km2 in America, and 3 X 104 km2 in Htfricaw and most of it is tinder rain forest.

Freshwatcr swamp forests

Description (Figs. 2.17, 2.18)

Like peat swamp forests these are defined mainly on habitat. They are a very tliverse assemblage of forest
types fiooded hy river water, hence ivith soils richer in plant ziutrients than peat swamp forest, and of fairly
high pH 26.0 or more). Where flooding is periodic, either daily, monthly, or seasonally, further dimen- sions
of variability are introduced, a group called periodic swamp forests. There ¡hay he a few centimetres of peat,
or an organo-mineral -_muck) soil may occur.

Habitat ¡md occnrrence

The Amazon, which has annual floods and is also influenced by tides to some 900 km from the mouth, has
very extensive and diverse fresh- water permanent and periodic swamp forests, known as varzea and igapo
respectively, for. white water (silt-iaden) and black water

Fig. 2.17. Seasonal freshwater swamp forest. Papua New Guinea.

Fig. 2.18. iireshwatcr swamp forest. Papua New Guinea.

rivers. Ñïihere unstable banks of ¡illuvitiln oceur, which are moved by every annual flood, exten- sive
cphemerai grasslands develop during the ltaw-water season. It is surprising to the visitor to see this mighty
river and its main tríbutaries flanked by meadows and not by forest. The species occurring in swamp forests
along lvlacic- water rivers differ from those along more eiltrophic white or clear waters. The alluvrial plains
of Asia once carried extensiva: swamp forests but littlenow remains as these have mostly been cleared for
wetland rice cultivation. The Congo basin is about one-third occupied by swamp forests, many disturbed by
man, and little studied.

2.3. FORFST NlAÏNTENANCE-—Tl‘lE

GROWTH CYCLE

The discussion so far has been about ¡nature forests. The different formations ali have dis- tinctive structure
related to which are particular physiognomic features and dispositions of epi- phytes and climbers. But the
canopy is not all

Fig. 2.19. Forest Canopy gap formed by the fall of a

single large tree. {Halle et al. 1978, fig. 107.) The medieval. French Word Chablis is sometimes Lised to
describe the Canopy hole, plus the associated damage, and the fallen mass of branehes, leaves, and twigs.

like this, because trees are mortal and eventually dic. In reality, thecanopy is in a continuous state of flux,
with gaps (Fig. 2.19} developing from one of many causes, varying in size from tiny to huge. Gaps are
colonized by seedlings which grow up to become saplings and then poles before attaining mafllfity. In order
to analyse this dynamic nature of the forest Canopy it is convenient to recognize a forest growth

"cycle consistíng of gap, building, and mature

phases, which are arhitrary subdivisions of the continua] process of grovrth.“ It follows that a forest consists
of a mosaic of patches at different phases of the growth cycle (Fig. 2.20). This Pattern-in space reflects the
processes of forest maintenance which continue through time. Where a tree dies" of old age, its crown slowly
dies back and then the limbs and finally the hole disintegrate. In this case a fourth, dcgenerate phase to the
growth cycle exists. Commonly, howcver, a tree clics suddenly, struck by light- ning or hlown over (Fig.
2.21}, of snapped off hy ivind. ‘Oi/ind may hltiw over several trees. In an extreme case, windthrow has
created long narrou’ corridors to over 8 km long in Sarawals: peat swamp forest. On steep slopes landslides
occur, perhaps as a result of an earthquake. Volcanoes create mud flows llahars) or ash or lava flows;
Between 10—2Ü°'n0rth and south of the Equator cyclones occur and these periodi-

Gap Building Pflalure

Fig. 2.20. Canopy phases on 2 ha of tropical lowland cvergreen dipterocarp rain forest at Sungai Nlenyala,
hflalaya, 1.971. Long narrow gaps result from windfaïll of single moribund giant trees. Big gaps result from
multiple windfall increasing an originally small gap. The extensive area at the north end of building phase -
forest results from regrowth after partial clcarance in 1917. A plan like this is a valuahle aid to interpretizig
forest structure and dynamies.

cally clear huge swathes of forest, in the Caribbean, Bay of Bengal region, northeast Philippines,
Queensland, and Melanesia. These various causes create Canopy gaps of grearly dif- ferent síze and some
places are more prone to extensíve destruction than others. The forest structural mosaie varies from very fine
to extremely coarse. ' The Dutch forester F. Kramer created in the lower montane rain forest of west java
artificial

Forest nzaintenartce —- the grrlwtfv cgicle

Pig. 2.21. View along a gap created by a single treefall. Note man to right of the upturncd roots and soil (the
so-calied root plate}. Stilawesi.

gaps of 0.1, 0.2, and 0.3 ha, equivalent to circles of 36, 51, and 62 rn in diameter.“ In the smallest gaps
seedlings grew up which had established below the Canopy, but in the larger two sizes they were replaced by
a new set of dif- ferent species which avere not present before. This is what alwajrs happens as gap size
increases, and the two kinds of species are called cliniaix and pioneer, respecrivelyfïi‘ They are dis- cussed
fully in section 7.2. Their essential char- acteristics are that clímax species can germinate and their seedlings
establish below a forest Canopy, so these species can persist in the same place, the seedlings growing up after
a gap develops. But if the gap is too big the clímax

27

species are replaced by pioneer species which germinate and grow fast after gap formation. The ¡TWO
essential features of pioneer species are that they need full light for both germinarion and seedling
establishment. Their seedlings are therefore. not found below canopy shade. They cannot therefore perpetuate
themselves in the same place. Below pioneer trees, clímax species establish and as the pioneers die off, one
hy one or in small groups, Canopy gaps develop and the next growth cycle is based on these clímax species.
There is a floristic shift from one suite of species to the tither, and this is what is termed succession {Figs
2.22, 2.23).“

Any forested landscape is likely to have a patch somewhere which is recovering from a landslide or front
multiple windthroxy‘ as a forest ofpitineers, so this patch is in a state of succes- SÍOH. Elsewhere clímax
species will occur. These, as a class, are Self-pcrpetuating, so their regrowth in gaps is cyclic replacement
rather than succession. The landscape is in a state of dynamic equilibrium, not changing as a ‘whole although
small parts are in continual flux. It has ¿iptly been terrned a shifting mosaic Steady state.“

Forests differ in the gap-forming processes to which they are prone. For example, Papua New Guinea“ is a
land of cyciones, earthquakes, vol- canic eruptions, and periodic tires, all of which cause catastrophic
tlestruction of big swathes of forest. The mosaic of structural phases is coarse and forests of pioncers are
uridespread. It is esti- rnated that 8-16 per cent per century of the land surface of Papua Nery Guinea is
disturbed by landslides {section 7.8), and 2 per cent of Panama, another earthquake-prtine couhtryd? Malaya
by contrast has very little catastrophic destruction from these natural causes; fine struc- tural mosaics and
stands of clímax species are widespread.

Gap-phase dynamics and the two ecological groups of species, pioneer and clímax, will be described more
fully in Chapter 7. It seems that

anos: of the world’s forests work according to this Inodelfls Strongly seasonal tropical forests, in which fire
is a major factor, are probably an

28 ‘Xfihai are tropical raiizïforests?

a8

¿É

‘1 Jlü‘!

“T ma’ M‘

l¡.

Ü E 12 18 21 3|] 3B Jn

12

Fig. 2.2.2. Forest on the north coast of Kolombangara, Solomon Islands, dominared by overmamrc trees of
the light-dernandiilg clímax species Canzprzosperrartt fvrevipetiofalzon (CAMB) ivhich is not regencraring
itself. {See Whitman: "i974, fig. 2.3 for full species names.) r_

—= This forest resulted from massive disturhance, and unless that is repeared it virill change in composition
to

i resemble that of Fig. 2.23, ‘whose species are already present in the lower part of the Canopy.

Fig. 2.23. Forest on the wvest coast of Kolombangara, Solomon Islands, dominated by the slow-growing,
shaide-ttileraijt clímax species Dillenia salomonezisis iDlLS) and Sclaizoirteria sermm {SCT-IS} which are
regcneraring themselves. (Whitmore "[974, fig. 2.4 and 1984a, fig. 17.20; see either source for full species
list.

Forest mairzretzance — the growtf? cycle

exception. [t is more likely that aifter tlisturbance resprouting rather than regeneration from seed is
important; but we have no detailed ecological knowledge on how they work, which is still a serious
deficiency ¿waiting an ecologist to apply modern apprOaClÏLCS and insights.

Canopy layer-s

ln this book the term forest Canopy is Used for the isrh-zile plant community above the ground. It is
commonly described as laeing layered or

i
29

stratified and this is a useful aid to description or amalysis although, because the forest is dynamic with
parches at all stages of the growth cycle, stratification is a simplification and abstractitm.

A mature lowland rain forest has trees of

many sizes. The rallest stand head and shoulders above the general level of the canopy and are known as
etnergents. They occur either ¿Ilone or as groups. Single kapolc trees {CÏeflJa ¡{Jemïai-idara), which reach
immense size in Amazonia, are a common and conspictious sight viewed from the

Fig. 2.24. Profile showing mature {ends} and building plTdSCS of the lowland evcrgreen dipterocarp rain
forest

iig. 2.1; see larrer for species names.)

st Belalong, Brunei. Plot arca GÜ >< 7.5 m, all trees over 4.5 rn tall shown. {Ashton 1964 in Whitmore ‘l
984g,

Dipterocarps shoïvn harchcd; note how these still have tall, narrow, youthful Inonopodiaïl crowns in the
building phase, which change to sympodial, broader than deep, and with several large limbs in the mature

phase.

30 What are trogüical ‘min forests .9‘

feet 80 170

Fig. 2.25. Profile diagram of temperate deeiduous forest, Sussex, England. F. Pages syfwttica (beach); Co,
Coryhrs aveífana lhazel); Fr, lïraxinzrs excelsítrr (ash); Q, Quercus robar (Oak); Tax, Texas ¿desata {tyew}.

{After Watt 1924, Hg. 3.}

river (Fig. 1.6). By contrast, the rough, hillov- ing eanopy of a west Malesían forest results from groups of
emergen‘: species of

_Dipteroearpaceae (Figs. 2.3, 2.24); Strata have

usually been depicted by a profile diagram, a side-view ofa strip usually c. 60 X 6m. What such a strip
depicts depends on where it lies with respect to the rnosaie of structural phases as can be seen from Fígs. 2.20
and 2.24. Before their use for tropical rain forests, profile dia- grams were used to describe the structure of
the forests of south England, which have a very simple structure, just one layer of trees over a shrulu and a
herb or ground layer (Fig. 2.25). It is probably by analogy to temperate forests that European scientists
sought to see strata in tropi- cal rain forests.

In addition to structural layering, which

becomes Confused by the growth eyele, different tree species habitually mature at different heights, and
whole genera and even some fami- lies reach up to different parts of the Canopy. For example, most
Ebenaceae and Euphorbi- aceae are small trees, most Burseraceae, Leeythidaceae and Sapotaceae reach the
top of the. canopy, and amongst emergents are many Leguminosae and nearly all rain forest

Dipterocarpaceae. Far Eastern Pvïyristieaceae are

nearly all small trees, but in the neotropics there are larger canopy-top species, e.g. the important timber
producing genera Dialyarztíyeira and lïírola. Young individuals, or trees dwarfed because of unfavourable
growth conditions, obscure such layer-ing and there is no evidence that habitual Inature height falls into a
small number of discrete classes. Strarificarion is most prominent in species-poor forest where groves occur
of one or a few species, which is why semi-evergreen lowland rain forest is more con- spicuously stratified
than evetgteen (p. 15). Crown form (p. 51) changes from Inonopodial, deeper than broad, to sympodial,
broader than deep, as a big tree matures (Fig. 2.24}. Some small tree species remain monopodial to matu-
ríty, eg. Annonaceae, Ebetielceae, and Myristic- aceae. It follows that layering of crown shape also occurs,
and this is also easily seen on a profile diagram. The forest microclimate alters Lipvvards through the Canopy
(Fig. 7.1). This probably triggers the meramorphosis of crown {section 3.2); it also leads to yet another kind
of stratifieation, that of epiphytes and elimbers (Fig. 3.18).

Floristícs
2.4. FLORISTICS

The humid tropics are extremely rich in plant species. Of the total of approximately 250 D00 species of
floweríng plants in the world, about two-thirds (170 O00) occur in the tropics. Half of these are in the New
World south of the MexicofUS frontier, 21 0.00 in tropical Africa (plus 10 000 in Madagascar} and 50 OOO
in tropical and subtropieal Asia, tvíth 36 O00 in Nlalesia. A few plant families are confined to humid tropical
climates, e.g. Myristieaceae, the Iiutmegs; others are strongly concentrated there tvith a few temperate
(Jutliers, eg. Annonaeeae, the soursop family, Musaceae, the bananas, and Ebenaceae, the ebonies (whose
temperate species provide the ftuits called persimmonlfi”

There are similarities, especially at family level, between all three blocks of tropical rain forest, but there are
fewer genera in com- mon and not many species?" All three regions have abundant Leguminosae of
subfarnily Caesalpinoideae. Other big families include Annonaceae, Euphorbiaceae, Lauraceae, Mor- aceae,
Myristicaceae, Rubiaceae, and Sapotaceae. On small areas Annonaceae, Euphorbiaceae, and Rubiaceae are
nearly altvayrs among the ten most species-rich families in all three regions. America is characterized by
numerous Lecythidaceae, the Brazil nut family, Vïíiïl} '11 genera, and about 120 species. The most dis-
tinctive and unique feature of western Malesia is the abundance and species richness of Dipterocarpaceae.”
Borneo for example has 287 species and 9 genera, and in many places most of the big forest trees belong to
this single family. Conifers have many species in the East (Fig. 2.26) and are found at all altitudes. So fat
only one conifer has been found in the lowland rain forests of Africa and another in tropical America.

.ln flora Africa has been called ‘the odd man out’;52 there are fewer families, fewer genera, and fewer species
_in her rain forests than in either America or Asia. For example, there are 18 genera and 51 speciesof native
paltns on Singapore islandÏ-as many as on the whole of
31

Fig. 2.26. The conifer Podocarpus rzeriífoliusis a common and valuable timber trce found in lowland and
lower montane rain forests throughout lVÏalGSÍFl. Note the big, permanent, radial limbs on this mature
specimen.

mainlaudï Africa (15 genera, 50 species); Africa has only 4 species of bamboo, and MtKinabalu (4101 m) in,
northern Borneo has almost the same number of ferns as the African continent. There are also differences
Awithin each rain forest region. Not all species occupy the whole available area despite the absence of
physical bar- riets; for example Tbeobroma, the cocoa genus, is confined to northwest Amazonia; and many
species found in Sumatra do not reach New Guinea. These patterns are thought to have his- . torical causes
which will be discussed in Chapter 6. Other patterns are believed to relate to climate. For example, the rain
forest flora of Ghana, herbs

32 lïïbal are tropical rain ftirests?

oc . 3DG- ¡Y I 25D — xKw x x X ZÜD-r _ ' x I X s ‘ U q) a. m o É 15o — x x 25 x E K I :1 z X >< u >< ><

x X l x ' 141D- . o U _ x c X 1 l U u 50- u ¡ l K K I u __ I , l o ‘l 2 Area {ha}

and trees, ‘dllÏErS clinally from west to east across the country, a distance of c. 300 km,“ in patallel with
increasing climatic seasonalityt.

To obtain a more precise picture of differ- ences in species richness, Fig. 2.2.7 shows the Iiumbers of tree
species of 0.1 m in diameter or larger on small plots. The numbers vary from 23 had in Nigeria to 307 had in
the Ecllaílürliltl Amazon at Cuyabeno.

Eig. 2.27. Species richness among trees of 0.1 In in diameter and over on small plots in tropical lowland rain
forest: n America; x Eastern tropics; I Africa. Lines connect sample plots that lie close together. {Data of
Wïhitmote 'l984a, f-ig. 1.5; Gcntry 19885; Víïhitlïiore and Sidijrasal 1986; ‘Whitmore er al. 1987;

Valencia er al. 1994.)

Species numbers rise Keith increasing plot arca as is shown on Fig. 2.28. In ziddition to this basic partcm, the
African forests are species-poor except for one at Korup {K}. Most of the Eastern tropics are species-rich but
the richest ftJrests of all are in America where, however, some are also very species-poor. The Yanamono
{Y} and Cuyabcno {C} plots lie in the upper ¡‘hnazon near the Andes {see Fig. 6."! 6h) and this region from
Colombia to Peru is now believed to be richest in the tropics for birds, frogs and primates, as well as
flovsrering plants, exceeding cx-‘en BÉJrneo. (Kw, Kalimantan, Víïanariset-Samhoia.)

On the richcst rain forest plots every second tree on a hectare belongs to a different species. It is difficult to
conceive a forest much richer than the Ecuadorian plot, where 76 per cent of the species present had only one
or two trees. ‘This plot had ‘JÜ per cent of the entire tree flora of Amazonian Ecuador {and '16 Pltr cent
amongst trees 2 5 cm diameter).

Species-atea eurves have been ctïensttucted for several forests, and species numbers continue to rise over
several hectares (Fig. 2.28}.
On a slightly more extended scale a single SD- ha plot at Pasoh, Malaya, contained c. 830 species, 2Ü—3Ü
per cent of the total tree flora of the country drawn to the size measured (10 mm diameterl.” An enumeratítin
of 6.6 ha spread across 5 >< 2.5 km of forest at Bultit Raya, Satan-ak, had 711 Species over 0.1 m in diame-
ter, which is about half the trees of this size found in all of Sarawalc.

To set these figures in context, the WlIOlC of ELITÜPE north of the Alps and west of Russia has only 50
indigentius tree species and eastern north America has 171.5“

One component of the extreme species rich- ness of many rain forests is the local endemics,

Floristrcs . 33

mol: Yananlono, Peru 0.1 rn

280
2BÜ_ lavanarisei.
240- Kaiimantan 0.1 m +
220— /f
20D i fi

. .51?

18D xBukit hagongl Malaya

15D. fzfigungal E0] m

+ ¡f Menyala

srïJaro, ¡(alimentan 0.3 m Papua 0.1 m

Nu rnber of species
10D
Papua G.3 m
I I lLi
G.2 (14 0.6 0.8 1.0 1.2 1.4 1.6
Area lhal

Eig. 2.28. Species area curvcs for tropical lowland evcrgreen rain forests. {After Xïïhitmore 1984:: and
Gentry 19885).

The Yamamono ftirest is the second richest yet found, every second tree on the hectarc plot ivas a different
species. These eurves xirere mostly made by adding together the number of species found on contiguous
subplots. See also Figs. 2.29, 6.164’).

57

namely species of ‘very limited range. For exaimple, many trees collected in Perak, north- west h/Íalaya, in
the late ninereenth century have tiever been found elsewhere and, because the loivlancls were soon
afternrards largely converted to ruhher plantations, are almost certainly extinct. The monospecific genus
Burfeillitiderzdron flllfïitfn, known from one col- lection from a litnestone hill in Perak, has never been seen
again and its habitat has been used for road Inetal. By contrast, ‘although the broadleaf forest cover of
England is reduced to c. 4 per cent, no species of woody plant has been lost from the flora.

Despite the great interest eitery-‘one has in the amazing plant species richness of tropical rain forests, there
¿NE very few total species counts. On a single 100 m3 plot in wet lovuland rain forest in Costa Rica, 233
vascular plant species were recorded {Table 2.3). This is hy far the richest plant community ever enumerared
on

Earth. In three forests in «western Ecuador, sample ‘areas ten times bigger (0.1 ha}, and each from ten non-
contiguous subpiots, had totals of 365, 173, and 169 species in perhumid, dry, and slightly seasonal forest
respeetiirely-‘ÏS Even if the trees are excludetl, all these four samples are amougst the richest plant
communities ever encountered with 153, 25”}, 125, and 105 non- tree species, respeetii-‘ely. The Rio
Palenque forest of western Ecuador has a total of 1030 species in 80 ha, one-quarter of them endemic; the
VRIhOlC of the British Isles have 1380 seed plant slaecies, ivhich helps put this figure in perspective.

The. most nearly comparable species-rich plant communities outside the tropics are the Is-"Iediterranean
heathlancls of the Cape of South Africa} {known as fynbos) and of southwest Australia. Here total species
ziumhers are similar to trees alone in rain forest. Chalk grassland in England is extremely rich on a small
scale and may have 32-33 species in a 0.5 m3 quadteït, hut the total flora of one site of a few hectares is tmly
5Ü*5S species”

a É ¡a e qJ CL U1 u.- Ü 3 b E El 3 Z

Area

Fig. 2.29. The species-area curves of the different kinds of species diversity.

The two species-poor communities (a) and (b) have low alpha diversity and low and high, hera diversity,
rcspectively‘. Communities (c) and (d) hy contrast are species-rich and have high alpha diversity and low and
high beta, respeetively’. Community {e} differs from all the iithers in having two distinet

l ctinstituent parts, namely it has higher ganuna diversity; the parts each resemble {b}. See also Fig. 2.28.

34 l-Wbrtt are tropical rain forests?

Table 2.3 Numbers of species and individuals in the different synusiae on a 100 m3 plot in ever green rain
forest near Las Horquetas, Costa Rica

individuals Species Total Cumulative total Vascular plants: {a} Independent {free-standing} plaints Plants :
3" "l m tall '134f3 132 132 {of ivhich tree seedlings} [566] (64) . Plants "¡-3 m tall 144 36 14Ü {of WlIÍClÏl
trees) (134) (S) Plants B 3 m tall 38 18 144 {of which trees) (35) (4) (b) Dependent plants Climbcrs bole 233
21 165 woody (free-hangítlgl (f8 Z0 174 Epiphyrtes 339 61 233 (of which aroids} (90) [17] ¡[of which
bromeliads] '49] {S} (of which ferns) 7] ' L9] Total all vascular 2'] l 233 Bryophyta: Liverworts 3 25 iVlosscs
3 ‘7 Total all Bryophytzi 32

From Wilhitmore et.‘ al. (1986) table ‘I

The count was made by a team of ‘11 petiple; removing one synusizï ata time, and took a total of 192 man-
hours. At this rate it would take 1.0 man-years to extend the plot to a full hectare.

These various (lata show that meaningftil dis-

cussions of species richness must specify scale.“ Por example, we Inay usefullji’ compare richness ivithin
rain forests hy counting tree species on plots of c. 1 ha. This ivithin-commutiity’ diver- sity has been called
alpha diversity. At the other extreme we can record species richness of a ivhtilelantlscape made up of several
communi- ties, and this has been called gamma diversitjv. The fynhos is very rich with 8500 species on 8.9
00D km3. It is Inade up of a mtisaic of differ- ent flo-ristic communities, each of "virhich has rather few
species. That is to say fynbos has

low alpha and high gamma diversity. Within a single florisrie community species replace each other from
place to place. This gives a third component to richness, known as beta diverm sity. Por example, within
lowland rain forest there are differences in species within a single community between ridges, hillsides, and
iralleys. Figure 7.38 demonstrates how in Amazonian Peru species composition changes as the forest gets
older and this is another example of beta diversity. The species richness of chalk grassland is due to high
alpha diver» sity, beta diversity is low.

hieture Of the tropical rain ¡‘tn-est comntertity

These different kinds of diversity can be shown on species-area curves (Fig. 2.29}. Where numerous species
occur on a small area the curve rises steeply. ‘Where the minimum area is iarge the curve continues to rise a
long way before flattening. These represent high alpha and beta diversity, respectively’. A change in slope of
the curve reflects transition to a different com- munity, so if the curve has a series of steps before it flattens
out this shows its full richness contains a component ofgamma diversity.

2.5. NATURE OE THE TROPICAL RAIN

FOREST CoMiv-¡IHNITYÓÏ

Now that the main kinds of variaritm in tropical rain forests have been described we can consider 3. question
that has been very much discussed by tropical ecologists: namely, whether tropical rain forest is one huge
floristic association varying haphazelrdly from place to place or whether distinct communities exist.”- It has
been shown in this chapter that there are in fact numerous kinds of isariation. It is possible to strange them
roughly into a hierarchy of dimin- ishing importance.

Biogcography

‘Jariatitm can only operate on the species that are present, and over-riding all other reasons for it is the
availability of flora. Por example, diptero- carps dominate the rain forests of western Íxialesia and give those
forests their unique char- acteristics which are referred to repeatedly throughout this book.
Disturbance

The. influence of nlassive disturhance is the second most important factor. Forests regrow- ing after a
cyclone or human destruction, for example, are dominated by pioneer or near- pioneer species (Fig. 2.22)“
and because trees live a century or more (Fig. 7.35) rare catastro- phes can have long-lasting effects.

Lu
U1
Habitat

Major physical habitats bear different forest formations (Table 2.1) and this-third cause of variation is of
similar importance to major dis- turbance.

Variation within the formation

‘Topograpbyr

Further down the hierarchgi, once these three factors have been allou-‘ed for, comes irariatitan Within a
formation. This teneis to be continuous, without sharp boundaries. It results from various causes. One kind is
linked to geology, which manifests itself in itarious ways relating both to topography anti to the chemical and
physical properties of the soii. Within a rain forest formation it is common for some species to be associated
with different topographic situ- ations, especially with ‘valleys or ridge erests.

Fig. 2.30. Homonoia riparia, a x-viry shrub of swiftly

flowing rocky‘ riircrs, is a typical rheophyte, with

linear-lanceolare (vrillovirlike) leaves called stenophylls.

Yui-man, SOUÍlNRJCSI China.

Rheophytes are a highly distillütfilü sjrnusia or life-form ‘community, They are commonest in the tropics,
especially in Nlalesizi Where they reach greatest abundance in northern Borneo. The life-form is an
adaptation to a very diffieult habitat, an otherwise empty ecological niche, which has been colonized by only
a few untelatcd plant families. Van Steenis {i981} gave a full account of these peculiar plants.“

36 ‘Xïímt are tropical rain forests?

Fig. 2.31. The dipterocarp
Shotter: ctrrtisíi and the big,

prickly, stemless bertam palm Extgeissona tristis commonly grow together in Ivialaya.

Fig. 2.32. (Right) Species patterns on a 50 ha plot at Barro Colorado Island, Panama. Trees over 0.2 rn in
diametcr are shown. {a} Trichilia tzrfaezctiïaïa, the commonest bi g tree, is ubiquittius. {b} Poaisenia
afflïrtfid is largely confined to steep slopes. (c) The pioneer species Cecrtapirt insígrtís occurs in cluinps not
related to topography but which have developed in cantaapy’ gaps. (Hubbell and Foster in Sutton et al. 1983.]
The distribution of trees within a rain forest results from many factors ‘which may íntcract and are not
always easy to discover.

Nature of the tropical rain forest commmzitgr

{al

{b}

A. -.(¿á;{i«¿i'(¡[¡
Cecropia insignia

37

_ wet Season of extractahlc soil phosphorus.

38 ‘Mii-rat are tro-pica! rain forests?

For exelmple, at the edge cif swiftly’ Ílovxiing rocky rivers and on their Siïllïgle beinks, firmly rooted in
crevices, is found the sjvnusizi of rheo— phytes (Hg. 2.30). On alluvial ttalley floors is found another set of
species. For example, in Sarawzik the ¡Jccasionally floodecl levées of satell- watered, friable, deep, fertile
soil support a very tall forest which includes the group of Siiorea species producing iliipe nuts, plus the
Borneo ironwood {Husideroxjiiraiz ztr/ageri). An illustra- tion of species that are commonest on ridge crests
is the grey-crowned dipterocarp Sborea ttitarrisií (Fig. 2.3) and its associate the stemless, prickly’, giant
bertam palm {Ettgetssortcz ïristis) in Malaya (Fig. 2.31}. Howexrer, not all species are stronglyr linked to
topograpby; Fig. 2.32 gives examples from Panama.

Stiil

Most rain forest soils are low in plant Ilutritïlïlïs and physical factors have a more important role than
fertiiityt in determining species rainges. Soil physical factors probably determined which species were more
alauïideint over either granite or shale in a survey of 26 628 trees on 676 ha in Ulu Kelantan, Malafrafií At
Ktirup, Cameroon, there large emergent species of Leguminosaïei Caesalpinoicieae {Mic-roberiiitiri
ínisnicata, Tetraberliitia bifolioiatrt, T. inoreiiann) form groves c. 600 m across svhich have been shown

to be associated with low’ concentratitms in the se.

PTESGHCG of" seedfing's

Another more subtle level of variation in tree species composition is dependent on the ¡Jresence of seedlings
on the forest floor at the time a gap develops. Different species establish their seedlitigs better on different
microsites {section 7.3}. Species also differ in how frequently they produce fruit and hence replenish their
seedling bank {section 3.3}; for example, the diptercicarp Borneo Camphürflïüüd (Dryofaaioiiops ¿ii-mirat-
ica}, xaihich dominttted certain hilalayati forests, probably‘ ÜWCLl its elhundance to the ‘reproduc- tive
pressure’ of more frequent ftuiting than tither dipterocarps. At this lowest lev-‘el of varia-

tion, which has been named gap-phase replace- ment, chance plays a strong role: trees Inust flower, the
flotvers must set fertile fruit, dispersal must operate, seeds germinate, seedlings estab- lish and survive
damage by falling debris or from herbivores, and then a Canopy‘ gap must develop over them tvithin a few
years before they die in the dimly illuminated forest untiergroivth.

As new studies are made to investigate the fundamental problem of the nature of variation from place to
place in tropical rain forest, they confitm the model of a hierarchical Series of kinds of variatitm, and
illuminai te the part of the hierarchy the particular study happens to relate to. Nowatlayïs, ¿mention has
increasingly‘ swung away from this problem to address others, which form, the subject of the next few
Chapters.

2.6. ‘ïil-IAT ARE titomcat RMN

FORESTS?—CHAPTER SUI»=1I\-I;—\RY

l. The wetter tropical climates have closed forests, collectively’ described as tropical moist forests. Tropicttl
rain forests occur where there is only a short dry season or none. Monsoon {tropical seasonal) forests occur
where the dry‘ season is Stronger.

2. Distinct forest formations occur in differ- ent habitats ("Fable 2.1), which are usually sharply houndetl.
They differ mainly in structure and physiogntamy and the same formations OCCUI’ in similar sites
throughout the humid tropics.

3. Forests consist of a mosaic of gap-phase, btiilding-phase, and ntature-pheise forest {Figs 2.20, 2.24}.
These phases are arbitrary subdivisions of a continuous forest growth cycle. The spatial pattern of structural
phases represents processes of change with time.

4. There are two contrasting ecological species groups whose key features are that clímax species can
germinate and establish seedlings below a canopy’, where-as pioneer species require full light. Clima):
species grow up in small gaps and pioneer species in big gaps.

Wfilïat ¿are tJTJpÍCcIÍ ‘rain forests? — Calsapíer sïuíïnraïy - 39

5. There are differences in species composi- tion at all scales, between the three tropical areas and xarithin
them. Tree species numbers per heetare on snmll plots vary from c. 20 to over SÜÜ. ¡‘Xfriea is poorest.
These factors have his- ïnrieal explanations {Chapter 6).

6. Tropical rain forest eonunulfitíes show ïrariation at a range of seales which ean be arranged in a rtnugh
hierarehy. Availalnility’ of

Hora eumes firsz, then" differences lvetween for- mations and that resultmg From nmssive distur- bEIÜCC
such ¿s cyelones. Xï-‘itlïíiï formations variation is more eentínuous and less obvíous and is related to
topography {riverine forests and I'l]EÜ]Jl1}’ÉES are especially cÏÍstíncth-‘el, to soil; as well a5 to seedling
elrrixral and success at ¿1 particular spot, for which there is a strong element of chance.

3
Plant life

The ttaveller Xvalks out of an abandone-d resthrause on to ‘what was a íawn ÍS months before, to find self"
sown saplings of Tectona grandis [teak] 5 rn high. They have long internodes, and leaves the size of dinner
places. He feels like Alice in ‘Wonderland. Or he noticcs the tveeds growing on abandoned farm land,
including such objects as Venzolnia cotrferta, a composite with vcgetative parts like a rosettc of Taroxacun:
(ifiïcinale [dandelion] 3 m across, on a

stalk up to 4 m high...“

The exuberance of plant life in the humid

. tropics continues to dazzlc scícntists from the

more sober temperate bíomes. More ‘purple passages’ have been penned on lowland ever- green rain forest
than any other vegetation type. Trees have a great diversity of form and size and some, have the Lmiquely
tropical attributes of huge buttresses or trunk-borne florvers. They support a wealth of climbers and
epiphytes: one stands and marvels. Bamboos (Fig. 3.1), palms (Figs. 02.31, 3.4, 6.3),‘ cyclanths, pandansfig
(Fig. 3.2), stranglers (Fig. 3.3-—3.5, 10.4), and banyans add peculiar extra dimensíons. As acquaintance
deepens some genera are found to have numerous species growing together, distin- guished in the forest by
details of bole, back, buttress, and leaf. Dípterocarpaceae in western Malesiaiis the extreme example. Of
‘¿Sborea M. Jacobs has svrittenfïi’

complete fugucs could he composed from‘ the leaves ..._, varying in size, thickness, venation and haïrs; with

Fig. 3.1. Scbízostachyugn grande showing the chat-

acteristíc clumped growth of bamboos. Ivíalazyia. This elegant species becomes ahundant in seriously
disturhed lowland estergrcen rain forest in the mountains at 600-800 m elevatíon.

Plant life - 41

Fig. 3.2. Jïararzmga sirmoscz, showing the typical open branchíng and Strap-like leaves of the Pandanaceae,
dendroid monocotsof the Old World tropics. Solomon Islands. ' ‘

Fig. 3.3. (Below left) Strangling fig, showing lnultiple descending and anastonnosing roots wholly obscuring
the trunk of the host tree. North Sutnarra.

- ¿fr Ca ¡Él-o í Fig. 3.4. {Below right) Strangling figjwith fan palms, Licuado rrzmsayi. Mission Beach,
Queensland. © Ted Nleadïwoodfall Wild Images. - e i

tw eden“) 7* .4-
: '\

42 Plant life

",9

n“?

"\

aii“. "-

J_¡-5\:CI'

_ «. a-

u,

i‘.

Fig. 3.5. The. development of a strangling fig (From Corner 1940 as fig. 2.25 in ‘Whitniore "l 984o].
Strangling figs occur in all parts of the tropics. ln Asia, lïïíghtia and ivíetrosidezros and in the New World,
Classic; also have strangling species.

Fig. 3.6. Fruits of ¿l miscellany of lfipterocarpaceac.

{FPtO 1.985.) The family ís named from the nvo-winged fruits of Diprerocarpus [Greek di—douhle, ptcroir-
witig, Ramos-fruit]. The wings are the flower scpals which become elongated and may be 2, 3, or 5 in
number or ahsent.

Plant life _ 4.3

Fig. 3.7. The monopodial undergrowth treelet Agrostisracbgrs íongifzilitt, here seen ‘with an erect tuft of pale
green young leaves. Singapore.

Fig. 3.8. {Below left} Eucaíyptus deghtprtz, twelfth tallesttree species in the Wrorld, here c. 60 lTl tall. New
Britain.

This is the only rain forest species of Jjtacïafgipttcs. lt is a pioneer of riverine‘ talluvium from Sulawesi east
to New Britain and is now commonly grovvn in planrations throughout the humid tropics.

Fíg. 3.9. {Below right) Ezzcaiyïxtzis degiarpm has very attractive hark, sloughing as long papety scales to
reveal pale green fresh surfacesthat darkcn through shades of green to become tvarrn recldish brown.
44 Plant life
Table 3.1 The Worlds tallest trees
Hcight Species {family} Local name Origin

{m}

143 Eucalygirus regnans Blue gum Victoria, Australia (Nlyrtaceael

115 Seguros}: sempervirens Californian redwood California, USA (Coniferae) '

107 Ettcalyprns registres Blue gum Victoria, ¿Kustralia (Myrtaceael

96 Seqnoiadendiron gigantenm Xïïellingtonia California, USA (Coniferac)

89 Araucanía hunsteiitíí Klin ki pine New Guinea (Coniferael

87 Kooinpassía excelsa h/Ienggaris, tualang Sabah, ivlalaysiai (Legumínosae)

85 Abies izobilis Noble fir Washington, USA

{Conifetae}

76 ' Dryobafaitops Ianceolïam Kapur Sarawak, hrlalaysía {Dipterocarpaceaej

76 Eucalyfittrs grande‘ Plooded gum New South Xlïalcs, (Ivlyrtaceae) _ Australia

75 Siborea supverba Balau Saball, lvïalaysia (Dipterocarpaceae)

75 Agarbis austmiís Kau ri New ‘Zealand

(Coniferae) _ -

71 Ettcalypttts deghtpta Kamerere New Britain (Myrtaceae)

70 Agatbis datirmara Sulawesi, Indonesia

(Coniferae)

70 Ceiba pentrmdra var. Kapok Africa caribaea y (Bombacaecac)

Various SOLUSCCS, including Richards 13.00’. (1996) Rain forest species shown in bold lRelict tree on
BAL Estate near Tawati

countermclodies of fruits and stipules; a single fer- menta [flourishl for a particularly large flower; and a
trcmolo for an aherrant calyx. And from time to time the recurrcnt tnelody of the flowers’ exquisite scent.

Just so, Fig. 3.6. If the kaleidoseope of striku ing inipressions is analysed ‘¡N8 see that trees are the
predominant life form, in all sizes from unhranched pygmies with a single apical tuft of leaves (Fig. 3.7),
which never get taller than 1-2 m, to eniergent giants, which include some

of the world’s tallest (Table 3.1; Figs. 3.8, 3.10}. Shrubs (Le. woody plants with several main stems) are rare.
Forest floor herbs are patchy, and much of the surface is hare except for a, usually thin, layer of leaf litter. On
landslips and along rivers giant herbs often form thíckets, gingers and Marantaceae everywhere, bananas

in Asia (Fig. 3.11}, Heliconia in America and,

lvfelanesia, bamboos in slightly seasonal

climates.

Plant life 45

Éig. 3.10. A giant SÍJorea czzrtisii. Note the two men the tree. Brunei. RS. ¡‘tshtan gives instructions i957}.

w‘

,.¡

Fig. 3.11. Extensíve Clumps nf the wild banana ÍVÏIISG ¿runcam colonizing a bared roadside, ivith two tree
ferns, Cyrzthea ccnrtarriinzrns. 1500 m clcvatican, ¿‘Vïaltwa ‘with PJ. Grubh (1977}.

Dependcnt on the Free-living QIIJÉOICrOPhÍC forest piants are a few Iieterotrophs. There are many
Loranthaceae (Inistietraes), as hemipara- sitic epíphytcs. Santalaceae, which includes the sandal-wvnod
(Santaírtrwz), is a family of smalï hemiparasitic trees. Full parasires include BakmOpÍJrJra (Fig. 3.12) and
Rafflesia (Fig. 3.13), ‘which in R. arnolidíi, has. the uroridï; largest HOWJCI‘, 1 m in diameter. Saprophgites
are Lmcommon and of only a few families, (Lg. Butmanníaceac (Fig. 3.14).

46 ' . Plant life

Fig, 3.12. Balanopbora Iafisepafa showing a male _ Fig; 3.14. Tíiisnzia aseroe. Tiny saprophytes of the

inflorescence. Lower montane rain forest, Malaya. ráin forest floor which live on dead plant ports and

i Balanophoras are heterotrophs, they have no '_ have onlythe flowers above groúnd. Lower montane leaves
or chlorophyil and are parasitic on tree _ ' rainforest; Malaya. " v i ' i" ‘

i¿ roots. 7 _ :

3.1.. CLIMBERS AND EPIPÍ-IYTES"

There are tivo different and distinct sorts of

‘cliniber. Big woody climbers are those that han}; freely from trees,—for example Gnetum {Fígs 3f15,
v3.16) with its hooped Stern,“ gnarled Bankinter, and in the Old World, especially Asia, the spiny climbing
palms with scaly fruits'called_ rattans. Big woody climbers have their leaves up- ín the top ‘of the carïopy,
and some have crowns as big‘ as" any tree. They are strongly lightí demanding and grow up in Canopy gaps
besides" being abundant on forest fringes. The other kind" of clirnber adheres to the tree trunks by special-
ized roots and may completely" hide the bark

of the woody elimbing ‘vines Tetrastigma from beneath mig‘ 3'17)‘ These are called bole iwhichonly the
flowers emergeThese ¡ire blotchcd Clímbefs‘ They are especially Coïnmon in Ve“? redand white; smell of
Carrion, and are pollinated humid Climate-S ¿"d uncümmon OI rare in

by flícs. . y - , _ , - - _ forests that experience a strong dry season. Bole —. V climbers are usually mixed with
epiphytes and with a group of species cailled"hemi-epiphytes

Fig,.3.13i.—- Rafifflesia pricei: ¿Note flyyfor scale. Sahehiv

"Rafflesias are parasites which" live inside the stems

which commence life 21s bÜlC cliznhers but‘

Ciianfaers ¿md epíplajrres . 47

Fig. 3.16. Cauliflory: Grzeïmn cuspidcatam: with its seeds borne on the Stein {note swollcn, hooped nodes).
Malaya.

showing the distinctivelyr Iiooped ‘stem of this gemis.

Éalalaya.

Free-hanging big woody climhers like this are a - charactc ristic life-form of many lowland rain forest fo
rmatítins.

Hg. 3.17. (Right) Eole totally elcithed by aroid bole i slinibcrs, (prohah|y' Scindnfysirs 5p.}. Big woody
¿limbcr to right. Sumatra.

48 ' Plant life .

-3Üm Ñ _25 ¿l _ _ E, photophyïes N ‘ j Freycmetia ‘g _2Ü ‘ safornonense g \ d) \ z“ - 11 a 15 L z E :5 m i

Mixed Araeeae E“: 5 i" 5 . ' * " Ï k‘ h e o l Srenaphfaena 3 spp. _. C s mp yt S

Fíg. 3.18. hole clinihers occiur- in zones. Lowland

rain forest, Kolomhángara, Solomon Islands. (Whitmore 1974.)

big. 3.19. lïatyceriunr, the stag’s horn fern, an epiphytc of trcc crowns; on Campuosperrïzcz auricatïatzmz.
Singapore. ' '

This fcrn has two sorts of fronds, one sort form a ncst in which humus collccts, the others, the fertile - fronds,
are pcndent and much divided.
Cïlinzbers and epipbytes , 49

Hg. 3.20. Dryrzaria sparsisora, another nest-forming _ epiphytie fern of tree crowns, forms gigantic masses, _
seen here in the crown of a Tacna attstraíís cmergent tree flowering after ir has shed its leaves. Queensland. '

become epiphytes by dying —oiff behind; by this

-eurious"proceditre they slowly climh the trunk ¿of the supportingtree. Bole clímbers and epi- gphytes are
¿zoned (Fig. 3.18), being—.specialized {to the different forest interior microclimares- "

zgFig. 7'.1). In the "very humid, dark lower layers shade-dwellers, or skiophytes, occur. Some, for

example, ‘the filmy ferns, are poikilohydric: they

have thin leaves which have the capacity to rehydrate without damage after desiccatíon. On the upper boles,
in the crowns and in Canopy gaps photophytes occur (Figs. 319-323). These are homoiohydric; they resist
desiccation by various adaptations. The leaves are leather-y with thick cuticles, there may he water-storage

Fig. 3.21. Two epiphytic orchids Maxillariá ehzznzeg (front) and Diotfaonerz ímbriieata (behind) from the
montanerain forests of Guyana. (Lindlkïjfis Siertum Orcbidacearun: 1838-, plate 40.) _ _i u i p These are
photophytic: ‘epiphytes-of tree-crowns. Crown epíphytes livenin a periodicallyhot, dry microclimate and
have many xeromorphic '_ i features. In these orchids the leaves pare thick‘ and ' leathcry _and the stems
have swollenwater-storing . leafibases called pseudobulbs. '

organs, e.g. the pseudobulbs ofsome orchids; and bromeliads, abundan: in theineotfopics, have-leaf bases
arranged to enclosea space or rank which collects water. lMany photophytes have} crasstilacean acid
metabolism (CAM). Some epiphytes trap nutrients by colleeting falling detritus amongst their leaf bases.
Hurnus then" deveiops, and roots push out into it, for example, the stag’s horn and bird’s nest ferns
(Piatyceríttm, Asplenizcm: Figs. 3.19, 3.23).

50 Plant life
Eig. 3.22. Taeaiophgrütrm is a bizarro genus of Old
XWorld epiphgrtic orchids in Xvhieh there are no leaves

and photosysithesis takes place in green, flattened roots. The name Ineans ‘tapewonn Ieaf’. h/lalaya.

Fig. 3.23. Asplenirrm nidos, birdïs nest fcrn, Old World tropics. l-iilmus collects in the ncst-forming frond
bases. java.

Trees 5 1

Other epiphytes may colonize and ‘aerial gardens’ form. The numbers of epiphyte species increase with
wetness of climate. (Jounts in forests in northwvesrern tropical. America have found 9-24 species in dry
forests and 238-668 species in very wet forests, respectively 2 per ¿ent and 23-24 per cent of the local flora.”

On old leaves a film of a special class of epi- phytes, called epiphylls, often grows. These are mostly
cyranobacteria, green algae, bryophytes, Éichens or small filmy ferns. They must impair Ïeaf function, and
have laeen shown to do so in coffee plantations.

3.2. TREES
Crowns

Crowns hax-‘c a single leading apical shoot or many, rcprcscnting n1o11(,)p()tiial and sympodial
construction, respectivcly. Nlost trees have znonopodial crowns tvhen they are young; many change at
maturityr. The plant with just an apical tuft of leaves (Fig. 3.7) has the extreme monopodial construction, a
form best exhibited by palms (Fig. 6.3). There are no branches, just the terminal ieaves. In palms the leaf
bases :onceal the delicate, usually edible, single apical had and are often spíny or fibrous. Very few palms can
resist frost and the family is essen- rially’ tropical. if the bud goes, so does the stem: ‘like a foolhardy
gambler the palm stakes all on a Single Card’. Dvlany palms grow’ by a tufting Éïabit, as a result of basal
synipotiial branching, 2nd this is common amongst Monocotyledons as a whole, as found also, for example,
in gingers, bamhoos, and bananas" (Figs. 3.1, 3.11). The apical leaf tu ft amay trap falling detri- rus, and this
has been claimed to provide a source of humus and nutrients for the tree. Other Inonopodial crowns have
lateral hranches, and these are. often whorled. ‘Where the lateral branches are in tiers, the crown has a
¿’iagtida-lil-(e appearance (Pig. 3.24). In the monopodial juveniles of forest giants the lateral hranches are
shorolived and eventually fall off.

Eig. 3.24. Pagmea cremskxtrt has a tiered, pagoda-

like crown. Singapore. i _ Tropical trees have a huge diversity of crown form whose construction has been
shown to result from the interaction of three basic processes. Crown form is a valuablc aid to species
identification, together with bark, buttress, and bole: there is no need to rcly on flowers or fruit.

They behave like giant pinnate leaves. In these big trees at maturíty, permanent lateral limbs develop fhígs.
2.26, 3.25, 3.29), and on them the juvenilc tree form may be rcpeated—so— eailed ‘reiteraritm’. The
metamorphosis from juvcnile to mature form OCCUIS in big trees when the Cfiflüp)’ top is reached,”
presumably trig- gered by the change in niicroclimate (Fig. 7.1). Knot-free timber is produced below the
sympo- dial crown. Trees that are tail in Virgin forest are

52 Iïtïïnt life?

mm)

Fig. 3.2.5. Crown construction of two dipterocarps. Slïorea nrecísropïerfivx (A, B} and Dryoihalaizryps
aromarica (C, D}. (Halle and Ng ‘l 981, in Wïhitmore 1984.52, fig. 2.9.} The juvenile tree (A, C) has
montiptidial crown structure. At ITÏHÍUÏTÜ’ the crown is metamorphosed into a sjrmpoclial structure (B, D)
which has numerous small suhcrtuxarns. In S. n-secistopieigvc these are reiteraritms of the juvenile tree.

Lisually’ much shotter Where grown as speci- meus, in arhoreta, or in forest regroxxrn on open land.

Tree croïvns in fact alvaays have a precise con- struction” determined by the intetaction of three main
factors: apical versus lateral growth, as just discussed; radially symmetrical versus bilaterally symmetrical
lateral itieristems; and imermittent versus continuous growth. The con- struction is most easily observed in
juveniles before metamotphosis. Twenttt-three different crown ‘models’ have been recognized. For example,
the papaya (Critica papaya) has apical growth with lateral inflorescences, is radially symmetrical, and gtoxvs
coutinuouslv. There is

little correlation between crown architecture amahïsed this ‘way and tatxonomy; for example, Euphorbiaceae
have numerous models but ¿’Xtmonaeeae and Ehenaeeae all hetve Inonopo- dial, radially hranehed erowns
of intermittent growth. Nor is there an}; etirrelation With ecologia The adapïlk-‘C signifieanee of architec-
tutal Inodels is unrestilvcd.

Leaves

The first imptession is of gloouiyr, Llull green, tiniform foliage. This is quickly dispelled by closer
inspection. There is "actualljv a great diver- sity in leaf size. shape, netve-pattern, margín, texture, and
CUlÜLIT. Lamina sizes amongst lourland evergreen rain forest trees are mainly but not eutirely notophjyll
and mesophyll (Fig. 3.2.6}, but as vras destrihed in (Ïhapter 2 micro- phylls are ptedominant iu heath and
tipper

I)‘

s4

[al 55:3! su 25 2T ‘.3 1a s . Tía ,0,0¡ —'—‘¡'J ¡e o, 22-22%:- ï-ÉEÉÉÉ EE-ÉÏLÉ EEQQÉQ 2g 322 Oesgeg
a¡__:ïg_ _;¿o u ¿‘Ceïazs :5“???

Fig. 3.26. Leaf size spectrum. {al Lower montane

rain forest [elevaïion ÍTlÜ m ‘i. {b} Lowland rain

forest (520 m). Anlazonían slope of the ¡‘males in

Ecuador. {Aifter Gruhh et al. 1963. fig. 5.} (Iharacteristicallï. the ltiwland forest has larger leaves. The leaf
size spectrum has proved a useful tool in the analysis of rain forest phjssiogtiomyt. Here the percentage of
species with lezn-‘es of different size is shown [sometimes number of individuals is used]. The spectrum
‘uvas proposed by Raunkiaer {19341 and modified h)’ Webb (1959) who introduced notophtlls {2025-4500
mml] for part of the mestiphjvll class. They should be consultes] for further details. Leaf area can be
estimated With sufficient rtecuraey as 2B lamina length (to base of drip tipl >< hreadth.

Trees _ 53

¿h .

Fig. 3.27. The giant palmate leaf of Travesia cheirarztha, ‘Ghosfs Fotitprint’. Malaya.

montane forests. In lowland rain forests, species tvith larger leaves are found in the lower part of the
eanopyïwhere it is most humid and cool and with less water stress. Pinnarely ancl palmately compound
leaves are frequent in lowland rain forest (Figs. 2.5, 3.27). The biggest leaves are found amongst giant
monocotyledonous herbs, the aroicls (Figs. 1-8, 5.6), gingers and their rela- tives (Zingiberales), ancl palma.

Leaves commonly possess clrip tips (Fig. 3.28). Recent research has demonstrated these really do hasten
dtainage of water films, which

probably therefore retards the growth of epi- ' _ phylls, as well as reducing loss of soluble nutri-

ents by leaching.

A Hash of colour gïimpsed up in the Canopy, as seen from the forest floor, or a eoloured crown seen looking
out over a hillside, is likely to he young leaves not flowers. Before full expansion the leaves of many species
are tinged reddish, though they may be white and in a few rare cases are blue. This very striking character-
ístie of tropical rain forests is commonest in shade-toletant species. It may be a defence

"Fig. 3.28. Drip tips of Ficus beccarii. Ivialaya.’

against hetbivores.” The young leaves have not yet become tough and inedible (cf. p. 82). They do not yet
have ehlorophyll and their delayed greening may make them less tiutritititts, and also may save theíplant
potential predation of valuable resources. Even so, young leaves are the site of nearly all herbivory. Unlike
temperate decíduous forests, there are no spectacular autumn colours in the tropical rain- forest. In fact,
hrightly coloured senescent leaves are sufficiently uncommon to be auseful aid to identification, for example,
Efaeocarptts leaves wither a clear red.

' Boles ‘

The interior of rain forest is often likencd to a.

Cathedral with giant eolumns teaching up to a

54 Plant life

Fig. 3.29. Cylinclrical bole of the iraluable timber species Agarbis roIJusta ssp. rzesogibfla. Highlands of
Papua New Guinea.‘ ‘ The smooth bark With distant papery scales has scattered whitish microliehens. The
big permanent‘ limbs "of the mature crown are visible, so are scars left by the ternporary hranches of the
juvenilc tree.

leafy roof (Figs. 2.24, 3.29). Boles in fact com- monly taper slightly, forgwhich foresters Inake allowance
when constructing volume tables.

They may be flutecl, and occasionally take on a‘

bizarre form (Fig. 3.30).

Buttresses

Buttresses‘ íFigs. 3.31, 3.32) are prorninent in some forest formations. They differ in shape,
Fig. 3.30. ‘ No one can fail to rccognize a mature Pertusadïrtrï tree because of its totally hizarre latticed
trunk. Malaya.

size, and thickness between species and fami-

’ lies; for example, they are particularly marked

in Bombacaceae. Buttresses are tension struc« turegtesonaring ‘When struek with an axe, and aire mainly
found on uphill sides of trees and cottnterbalancing asymmetric or epiphyte laden crowns. An analysis of the
stresses to whicha tree is subject and the strength of its wood shows. buttresses are irtdeed of structural
importance, helping to support the tree.”

Trees _ 5 5

..........4..-e.h..-.u._.....a,..a..sm.ui..uwmnuum.z.mmfiñnmw.._..eauum..-e.__._‘ v__— “qm, r ,-—,4;_4;; 4..

Fig. 3.31. Steep plank butttesseï, of a big kapok tree, Ceiba pentandra. Ghana.

Kapuk reaches .70 rn tail in Africa; this variety (var. caríbaea) is the tailest rain forest tree in the cínntinent
and also occurs in South America (Fig. 1.6} together with the othci‘ three species bf Ceiba, It is iikeiy that

C. ¡bentandra ‘was dispersed on sea currents to Africa where it now grows wild from Senegal to the Great
Rift ‘Jaiiey and Angsziia. The ¡rscfui fibre Ïcapok from inside thexfruit is derived from a smail spineiess
variety (var. perztandra} which is cuitivated throughout the tropics (Fig. 20.2);

56
Plant life

Fig¿ 3.32. Btimhacaccac are characterized by particularly‘ huge buttresses, which artain Erobdingnagian-size
in Hufieroderzdrcan duckei. Brazilian Amazon.

‘Trees _ 5 7

Hg. 3.33. Flying lnuttresses and stilt roots of Llapaca ‘

gnizzeensis. Ghana.

Flying buttresses occur in some families (Fig. 1 J

.33), and are common in swamp Fiat-esta. Stilt

roots are another feature. They are very char- actcristic of pandans, and are also Found in pellms, some
mangroves (Fig. 2.11.4}, and a few other trees.

B ark

Bark is extremely diverse.“ Colour varies from coal black in ebony (Diospgrros) and some Ïx-Iyristicaceae to
white in Trisrartíopsásfï through bright rufous hrown (Ertgenirt). Calycopbgirlhszn of the neotropies has;
hark like hurnished copper. The lazo-h surface may be

smooth, scaly, fissured, scrolled, or dippled (Figs. 1.4, 2.26, 2.31, 3.9, 3.29}. Smooth bdflCS have a ltang
persistent surface and these often house lichens which may he specific, e.g. Dfsogayrtis commonly has a dark
green micro= lichen. The hark may contain a latex, resin, or coloured sap.

The botanist, confused hy jacohs’ eacophony of fallen leaves, uses the crown, hole, huttresses, and bark to
help him With identification. Flowers are seldom present and are hard to see or to examine high in the eanopy
but the Chill’- acters of the living tree are so diverse they provide ample clues for identification. The most
Liseful forest Floras are based on these ‘field’ characters and use them entirely For their kevsÏ“

Roots

These are, foi’ ÜlTVÍOUS reasons, less ‘¿vell known than the parts ahove the ground. Sonle rain Forest
species have a deep rap root. Others hai-‘e ‘sinkets’, roots that tlescend xrertically’ from spreacling roots or
buttresses. lvíost of" the root hiomass is in the top 0.3 m or so of the soil and there is sometimes a
concentratitm or root mat at the surface. The relatively‘ Few deeper roots Inay he important in gathering
DUÍTÍCHÉS from deeomposing rock or which have Éeziched down; species with such roots have
putitentiaillyr an important role in levar-input sustainable agricul- turc hased on tree and crop" mÍXEUFGS
(p. 163). Deep roots are important in tapping water in dry weather (see p. 17}. Fine roots atc difficult to
study; it is hard to extract them from the soil and tvhen exrracted to see which are alive. Roots up to 2 mm in
dianneter form 20-50 per cent of the total root biomass” ¿ind their believed rapid turnover is probably a
significant part of ecosystem nutrient cycles (Chapter S).

It has recently been realized that, as in other forests, in the rain_ forest nearly all trees have mycorrhiza.
These are mainly cndotrophic, vesic- ular arhuseular in typeghut a tien! groups, some trery
important‘(Dipterocarpaceahe, Fagaceae, Etzcalygitzrs, LegumitiosaefCaesalpinoideae), have ectotrophic
myrcorrhiza. Nitrogert-fixing nodules

58 Plant life

occur in most Leguminosae {the bacterium Rbiztibium), and rllntrs and Casuarina (the fungus Frankfa}.

3.3. seasoNAL RHYTHNIS

Shoot and Ieaf growth“

Tropical rain forests are evergreen and the climate is benign to plant life, always warm, and moist for all or
most of the time. One might expect growth to be continu ous. A few trees and rather plants have indeed been
found to grow all the time, as measured by production of nodes, each hearing one or a few leaves. The
pioneer tree h/lacaranga tendrias is an example (Fig. 3.34). Internode elongation rate, another measure of
growth, is constant in this species but in trees of other species, despite continuous production of nodes at a
Steady rate, elongarion rate, and hence height growth rate, increases through time (Fig. 3.35). The exact
measure used for growth matters; even with Macari/enga
3.8
3.6

314 130 _ 3.2 12o ‘ 3.o 11o _. 2.a mo __ -— ‘ 2.6 2.4 2.2 2.0 1.8 1.a 1.4 1.2 1.o 0.3 0.5

Curnulative number of new nodos .

Final height of nodes im] {al

D12 3 4 5 Ei 7 8 91Ü1112131i415
March June
19?5 Mmmm 19??

Fig. 3.34. In Mncaranga ¿arterias height growth is steady. New nodes form continuously and regularly and
every internodc clongatcs by the same amtiunt.

{After Ng 1984, fig. 3}.

tanaritrs we do not know if meristeln activity is continuous. Palms exhibir conrinuous growth and the hoop-
like leaf scars left on the trunlc show elongation is also Steady, though steni diameter waXes and wanes with
growth condi- tions (Fig. 3.36).

Most rain forest trees, however, exhibir inter- mittent shoot growth (Fig. 3.37}. A number of nodes form with
their associated flush of new leaves, sometimes brightly coloured at first, and often as pendent, hanging
tassels; followed by a testing period. The intermittent growth of the shoot tips is seldom reflected by growth
rings in the wood, and where it is these are not annual and often not annular either. Rain forest trees, unlike
those of seasonal climates, cannot be aged by counting wood rings {section 7.7}.

Intermittent growth inhibits the build-up of fungi or insects which attack young leaves. In Malaya young
leaves of Heuer: are often attached by two fungi, a mite, and a thrip any of which ciccasionally causes
defoliation, even though the tree produces leaves in flushes.

-—- 4.9

j’ —- 2.6

/ f 1:3

_/.J' 1.5

,.=/' y 1.4 í ‘¡f 1.2 ,2’

É8

\ I h.) Lt Final height of nodes 4m] [el

1.0
Cumulative number of

new nodes o in‘

IEl||II"IIÍ|l||
2 3 r1 5 8 7 8 Q1G111213141516
June
19??
Ü ‘I

fiflarch 191G

Months

Fig. 3.35. In Üroxyluin iizdicnm height growth incrcascs through time. New nodes form continuously and
rcgularly’ lcf. Fig. 3.34] but the internodes continue to elongate. Growth rate measured by node production is
linear but measured by height it accclerates. {After Ng 1984, fig. 5.]

Seasonal rhythzns

Hg. 3.36. This old flowcring Corypba ¡unbiztczriífera [ïiïlipüt palm] in the strongly’ seasonal climate of east
java has experienced gotid and had growing periods.

In a study of 81 species of dipterocarps at Kepong, is/Íalely-‘a, 70 produced leaves in flushes and in the
nearby Ulu Gomhak forest 38 out of 41 species flushed.

Leaf falls’

The sharp distinction hetween deciduuus and evergreen species of temperate climates does not exist in the
humid tropics.

Trees with continuously growing shoots suffer continuous leaf fall. Leaf life span xraries from about 3 to '15
months. A young palm ¡hay accu- niulate leaves and develop an oblong crown

59

(Fig. 6.3), but at Tïlaïtlritj,’ for every new leaf unfurletl, an old one drops off.

Amongst the majority of species that grow in Hushes we may distinguish three patterns of leaf fall. Vïhere
leaves are shed well before bud- break the crown will be bare for a period, perhaps only a few days. Such
trees are termed deciduous. ln the humid tropics such species fltnver on the hare crown, for example,
Bombrix, Firmíana (Fíg. 5.1), Pterocyïnbium, and ‘Toona (Fig. 3.20), whose bird pollinators are probably
assístetl by the lack of leawes.
ln other species leaf fall occurs at about the same time as bad-break (Pig. 3.38), but may be a few days earlier
or later, depending on partic- ular weather conditions or on the local climate. For example, the rain tree
(Sanranea suman) is ever-green in Singapore, but is bare for a few tlays in central America where it is native.
En Singapore it briefly’ has two co-existent leaf flushes and in America it is hriefly deciduous. Species Vvitl]
this behaviour are called leaf exchaltgers. The gardener whose tasle. is to sweep up the fallen leaves only has
to do so once or twice a year.

Fiilzllly‘, leaf fall may occur Well after bud break. Species With this pattern are evergreen. Leaves live about
7 to '15 months and leaves of several flushes sometimes (JCCIII together. In ytïung trees of the dipterocarp
Neobczfaszticargurs fJEÍTITÍÍ up to seven leaf generarions have been found to be present simultaneously’.

Synchronization

Leaf life üpans on isolated relict and planted specimen trees observed in Singapore at the tip of the Malay
Peninsula were found to be 12 months in four species and 6 months in another three species.“ One might
adduce a climatic trigger for leaf fall except that 17 further species had lifespans with no relation to the year,
and were mote or less constant in each species but ivith individual trees out of phase. This stiggests "there is
a genetically determined life span; leaves become old and less efficient as they ¡become

60 Plant life

‘JUOr- 9D__ G Branch developed at node

7D
6D
40-
30
2D
10
Curnulative number of new nodes -

(TI É t

Ü1
Dec
19?5

Final height of nodes [mi le)

2 3 4 5 6 ir’ 3 9 1011 12131415l617i18192Ü212223

Dec
19??
Months

' Fig. 3.3 7. In Harpullia confusa height growth is in flushes. New nodes form intermittently with testing
periods between. The internodes continue to elongate after initial formatiom-[After Ng 1984, fig. 10.)
more shaded and accurnulate epiphylls and are eventually dropped.

These specimen trees all behaved as individu- als. A few species showed lack of synchrony within the crown
between different limbs— mango (Mangífera) is a good example-«and this has been called ‘manifold
growth’. It was not until a study was made of trees in their tiatural habitat that it was realized that even in the
almost aseasonal climate of Malaya (Fig. 2.1} there is an annual rhythm in phenology. The study covered 9
years at the Ulu Gomhak” forest and 61 treesof 46 species were observed monthly from a platform in the
crown of an emergent. There were two leaf flushes per year, in February to April, and in September to
October. Most leaf change occurred then, though different individuals of a species were often not in exact
synchrony. The main peak of leaf growth came after the driest time of year, the second lesser peak began
before but extended into the wettest time of year. Most

trees flowered annually, mainly at the early leaf flush, but some at the later one. The Dipterocarpaceae
however only floivered twice, in years which also saw heaviest flowering of the other trees. It is well known
in Malaya that durians and other forest fruits are on sale twice a year, and there are sporadic bumper harvests.
In more strongly seasonal tropical climates than Malaya there are more numerous decidu- ous species.
Crowns are bare in the dry season, but flowering and leaf flushing show complex patterns that are not all
easily relatahle to clí- matic seasonality. An interesting study in the tropical seasonal forest at Guanacaste,
Costa Rica,“ found that flushing and flowering were controlled by tree water status. On dry sites leaves were
lost early in the dry season. In most species rehydration and bud break occurred fol- lowing heavy rain, but in
a few leaf loss allowed rehydration and was followed by bud break, sometimes Xvith flowering, without rain
having fallen. On EVEÍ sites the species remained ever-

Sensorial rhjxtbms _ 61

Fig. 3.38. Dyera costtriara (jeiutoilg) is a leaf- exchanging species. On this tree a flush of new lCaVCS seen.
The new leaves are expanding immcdiaatcly after the old ones were shed. Note Picztyceriunz crown
epiphyrtes. NIaIaji-‘a.

green or just exchanged their leaves. Later, experimental irrigation of clusters of trees confirmed these
deductions.35 In the semi-ever- green rain forest of Barro Colorado Island =BCÏ}, Panama, only a small
minority of tree 3nd woody climber species tested had leaf fall delayed by maintenance of moist soil by
irriga- tion during the four-month dry seasonfs Computer modelling supports the hypothesis that flushes of
shoot growth, leafing and tlowering result from genetically controlled endogentius periodicity being
entrained to sea- sonal changes in water stress, progressiveiy more strongly the stronger the dry season."
Thus, in Malaya endogenous controls doniinate, ¿It BCi soil moisture buffers most species against the
moderare seasonal drought, whilst in Costa Rica the strong drought is important. It is seen that the interaction
of internal rhythmic processes and external drought can have various

consequences. HOWCVCI‘, one patterri, common in Inonsoon-climate Asia, still remains enig- rnatic. In the
dry deeiduotts forests of Thailand, for example, new leaves are produced more or less synchronously Kiefer-
e the first rains break the annual 4-5 month drought. How can shoot tiCtÍVlty resume in water-stressed twigs?
Further studies are needed.”

{e‘¿,‘}=_xr0rx_i_ 2,_l_.(_ Fla\'k(—y'\;r\

Flowering patterns

Climax species in rain forests mostly‘ flower once a year and at about the same time. Some figs (Ficus)
become fertile very irregularly in Contradictiolï to this generalization, and With important consequences for
their dispersers {section 4.3}. There have been suggestions, which remain unsubstantiared, that ‘small trees
within the Canopy flower more continually. By contrast, pioneerspecies, and hence secondary forests, are in
continua] flower, some species several times a frear and others all the time (eg. in Asia Adinrtrzdrfl dumosa,
rliiacarartga fJej-vzei, and Diiienia suffrnticosa and in Africa Vernonia conferir: and Trenes orienraiis).

Monocarpy

A fewspecies, mostly monocotyledonous plants such as some palms and bamboos, exhibir monocarpic or
‘big bang‘ flower-ing. Usually the plants grow for many years and then (aver a whole district simultaneously
flower, fruit, and die. This is best developed in the sea- sonal tropics, for example, many bamboos in
continental Asia, including (Zhina, whose eopious fruitfali is followed by ¡Jlagues of rats, and then a dearth
of materials for house eon— struction until fresh clumps grow up. One of botany’s mysteries is how
subdivisions of a single bamboo clump grown apart time them- selves to flower simultaneotisly’. One
striking example concerns Arttndinarfia ¡Caicara which flovirered 35 ‘years after introduction from India in
Algeria, France, Luxembourg, and Irelanddïg Most spectacular of all monocarpic species is

“the talípot palm (Cris-aguilar: izmbraciriifern),

which grows the largest inflorescence in the llvlalaya.

'62 .- - ü : : ‘Plaiztlife

' Fig. 3.39. Corypha irmbractrlifera, the giant talipot palm, is monocarpic. It fltiwersoncc and thcnidies.

An early European traveIler-wrote back from Indiain 1681 ‘one single leaf (is) so hroad that it will cover _ ‘
some fifteen to twenty men, and keep, them drywhen it rains’. _

worldlFigs. 3.36,, 3.39): a gigantic apical candela“? brum, ‘which ‘then develops hundreds of thou-

sands of fruitsbeforeithe palm dies. A whole avenue of. talipot» the -I’eradeniya. Botanic Garden, gSri
‘Lanka flowered and then died

simnlraneously. _ _ _

' «There are fewer monocarpic woody dicotyleá ‘donous plants. The shrub Strobiíarvrtbes with" several
species" through the Asian ‘rain forests is

one; Tacbigali," with" '24.’ species through the

neotropics, Harnrsiopartiaxi"ingens in .the moun: tains. of New" Gtihiea,9" Cerberiopsis "(3 spp., New
Caledonia} and some Spatfaelia spp., (tropi— cal America) are thefonly trees so farknown which have this
habit. ' ' i
Massflorvering in russian: il/Ialesia
Everyyear iniMalaya and Borneoithere “is a.

single period of heavy flowering and fruiting. A5

"deseribed-abtive, careful studies (Jver 9 "years alt

Plant life -— Chapter sttmmary

RaípfalJ-Kepong

ur.

Leaf fIushing-Kepong [No data fer July 19m . ' ¡

lïI1

zlI1III11I1I

No. of trees

l|||IIIÍ

b C3

No. of trees

l l l l l l l l É L. 19?? i inean

Fig. 3.40. Leaf flushing and flowering of dipterocarps, and rainfall. Kepong, A/lalaya. (After Ng, 1984, fig.
12.}

Ulu Gombak, Malaya, showed there is a second lesser episode and that production of new leaves alsta occurs
mainly at these times. These two annual events coincide with two slightly drier peritads, a few ‘weeks of
little rain. Once or twice a decade the early dry spell is unusually strong and is followed by heavier than usual
flower- ing and fruiting, and in these years there is gregarious heavy flosvering and fruiting of
Dipterocarpaceatt (Fig. 3.40).

Different species of dipterocarp flower after each other over a period of several weeks but fruiting is
simultaneous, at about the time heavy rains begin. The fruits are highly favoured by pigs and mass fruiting is
believed to satíate these marauders as well as seed-attacking beetles. There is strong selection pressure
against any tree out of phase, so mass fruiting is a self-reina forcing phenomenon. Htwvever, mass frniting in
this family is confined to the west Malesian rain forest species, and we have no explanation as to why the
gregarious dipterocarp Anisoptera tburifera of New Guinea flowers and fruits every year.

63
There are no other tropical rain forest plants that show such massive occasional fertility. It is this property
that makes di pterocarps and the rain forests they dominate unique (see section 7.6).

.-

Triggers for flowering

Chilling of already formed young flower buds, triggering them to complete their development, has long been
known to be the flowering stimulus for some species.” For example, Zephyranthes rosca, the rain lily, flowers
after the stimuius of rain and associated Cooling; some (Jrchids and trees (eg. the pigetm orchitl Dendrobizan
crumenatum, a common epiphyte of wayside trees of Malaya, and the angsana tree Pterocarpus indices) burst
into bloom some days after a midday storm causes sudden chiliing. Dipterocarpaceae may show some such
trigger. As mentioned above, mass flowerings ctiineide With exceptionally dry weather. Up to abcuut half of
all dipterocarp trees burst into flower simultaneously. Later, the corollas Carpet the ground like fallen, tinted
confetti and are followed after 3-4 months by a heavy fruit fall. There has been much slaeculation as to the
exact nature of the trigger. Dronght, or an increase in hours of bright sunshine, or an unusually big difference
between night and day temperatures because of the cloudless weather occurring at that season, have all been
Suggested. The problem has still not been finally resolved.”

3.4. PLANT LIFF.——CHAPTER

SUMMARY

1. There is a great richnessof life forms in trop- ical rain forests. The trees provide support for elimbers and
epiphytes. There are species of all life forms adapted to the different forest interior rnicroclimates; the
broadest classification is into photophytes and skiophytes. 2. The first impression of a uniform phalanx “of
trees is quickly dispelíed. There is enormous variety in crown, bole, buttresses, bark, roots, and leaves.
Flowers are ephemeral and species

64 Plant life

are usually identified from these characters of the living tree.

3. Only a few species grow’ continuouslyrlïor most, shoot and leaf growth is intermittent, in flushes. Young
leaves are eommonly’ highly coloured. Leaf fall follows various patterns, and often occurs after a new flush
has grown, but there may be a bate period and this is more pro- nounced in seasonal climates.

4. Frarests show synchrotiized leaf flnshing, and usually bear flower and fruit annnally. This is in response to
climatic cues interacting with

internal plant physiology, and becomes increas- ingly Inarked the more seasonal the climate. ‘West Malesian
rain forest Dipterocarpaceae are unique in their gregarions flowering and fruiting only a few times a decade.

S. Pioneer species flower more continually.

6. A few species, especially in seasonal forests, are monocarpic, floivering then dying.
7. Triggers to flowering include low night temperature and the ehilling of bud initials early in their
development.

4
Rain forest animals”

Animal life is, on the whole far more alaundant and varicd xyithin the tropics than in any other part of the
globe, and a great ilumher of peculiar forms are found there which never extend into remperatc regions.
Endless cccentricities of form and extreme richness of colour are its most pronrinent features, and these are
manifcstcd in the highest degree in those equarorial lands where vegetation ‘acquircs its greatest beauty and
fullest development.

,-\.R. Vï/allace ‘Tropical nature ¿Ir-id ¿Jtlser esseys, 1878.

4.1. RICTINESS AND DIVERSITY or

AN lMALS

The casual visitor enter-ing a rain forest is oï-‘er- whelmed by the lush diversity of the plants but
tlisappointed hy the aptgireng absence- of _ani_rr_1g_ls, _eg<ce.p_t_..for_the rising-i and falling reso-s-
fiant _t_rill of cicadaïs, and very soon tooby the attentions of-bitinginsects, and__in_.thec,Easter_n
ttopifi_of_hlood;s_u_ckiug leeches. The observant naturalist hoivever, deteets the signs of bigger amimals. ln
Borneo for instance he may notice an ti_r_ang_nta11’_s,nocturnal sleeping platform of branches roughly
thrown together high in the Canopy; scratches on a tree trunk x-vhere a big cat has sharpened its claws or
where a sun bear has tried to break into a_bees’ nest; part of the forest floor swept clean fiofflleaf litter as the
display ground of an cargos pheasant; the muddy

holloxvs or disturbed soil surface Of.\«'v'21ll()\VlI'lg ot rootling To see the animals and not just their signs he
will walk quietly through the forest at dawn or dusk- when lgjgls are most active, or at night when many
lmïnlniïlíeïnd frtgs are awake and can be seen in the light of a head-lamp.

Tropical rain forests are irldeed fabulously rich in ¿Inimeil life as AR. Wallace and the other nineteenth
century naturalist explorers reported. Just how many species the world’s rain forests contain is still, a century
after their pioneering studies, only a matter of rough conjecture. For manïmzals, birds, and (ithcr larger
animalls there are rtiughly twice as many species in tropical regions as tempetalte ones (Fig. 4.1). These
groups are fairly ‘avell studied, insects and other invertehrares much less so, and if the same pro- portion
hetx-veetl tempetatc and tropical species holds for them then the tropics as a whole may hold a total of
Lfiïlnílliongnaimal species; the great majority! of these are undesiïrinhflefd insects, with their greatest
ahundance in rain forests.

The richness‘ of various animal groups in different patches of rain forest is shown in Table 4.1. Most of the
hyper-rich sites, ali of them world records, are in the. Lipper Amazon close to the Arides.,Tl1ese include
birds at Limoncticha,__Cochahgïgslflmfllfïablc 4.1) and

“fiiïiïlïñpfaÏt-a, each-nvith about 500 species in

uÏder-‘SO km1, amphihians {frogs and toads) in

66 ¡‘Rain forest aizimals

á? Laïítude° Latitude“ Qee” 5 añ’ ,. 57mm? ‘b,

_ l7p\ 108 ' 123 4o Ü x ' 5|] J_LJ_I4_I_LJ _ ° Ü 20 49 BU SU Ü 2G 4Ü 6D Ü 20 ¿Ü BU 80100120 Number

of species Number uf species Number m‘ species Fig. 4.1. Decrease away from the tropics in species
numbers of vswalluwtaail butterflíes exemplifies a very con-unen biogeograïphíc pattern. (Collins and Morris
1985, fig. 2, in Ixmglnan ¿nd jenik 198?.) Table 4.1 Species richncss of ‘various groups in some well-studied
lowland rain forests Number of species Area Mammals Birds Reptiles Amphibians Moths Vascular {km3}
ípriznates) l plants [non-ílynagj Neutropice: Panama - ‘ Barro Colorado Is. 15 _ 97{.‘ï){39} 443 S1 52 ”'
1320 Costa Rica _ La Selva c’. 15 c. 117(4)[52} 410 86 43 c. 4000 "1 668 Ecuador _ ' Límuncocha '15 ‘*
480 * * ‘ “ Santa Cecilia 3 > * ‘* 92 93 ’° * Peru ' Cocha Cïashu :2 50 W] 3)Íc. 70] 554 54 77 '"‘ 1370
Africa: Gabon Makokou _ 2000 _ '199(14) 342 63 38 * ”‘ Far East: n I Malaysia Pasoh 8 89(5) 212 > 20 25 "
‘ Papua New Guinea _ . .

Gogol 10 , 27(lÜ] 16.2 34 23 " ‘l

Various sources, including (lcntry (1990), ÏVïcDade c: al. (19.94)

*‘ N0 dara available

Richness and diversity ofanimals _ ,67

Ecuador (Table 4.1), and a single tree at Tambopata, Peru, with 43 ant species in 26 genera-about the same
number as the whole British Isles or Canada.

The question of how mativ species live in the ivorld’s tropical rain forests is currently under debate, as part‘
of the concern for their adequate Conservation to be discussed in Chapter 10. The huge figure of 2-3 million
species, or two-thirds of all forms of life on earth, may even, perhaps, be a substantial underestimate. T.L.
Erwin“ has argued that the global total count may be nearer 30 million species, based on his analysis of the
insects living in the crown of Luehea seemannii in a scrubby tropical seasonal forest in Panama. Nineteen
trees, sampled over three seasons, vielded 955 species of beetles not counting wcevils. They were extracted
by fogging the treecrovirns with knockdown insec- ticidcs. Erivin’s study breaks away from earlier loose
arguments about species numbers by pro- viding a new focused approach, and although one might disagree
with the steps in his extra- polation he shows the way objective estimates can he obtained.”

Taxonomic description, mainly of vertebrates, has proceeded further than ecological studies, and
investigations on the ecology of animals in rain forests have developed more recently than of plants. ln this
chapter something is said about the differences in vertebrate communities from place to place, followed by
discussion of the main means ivhereby so many species are able to co-exíst, and then on what factors may
limit animal numbers. In Chapter 5 we go on to describe interactions between animals and plants, cohabiting
components of the rain forest ecosystem. Invertebrates are important as decomposers of dead organic matter
on the forest floor and this subject is discussed further in section 8.2.9“

Differences and sirnilarities can be seen between the faunas of the three tropical regions. Let us consider
primates as an example. In trop- ical America and parts of Africa up to 14 sym- patric prirnate species co-
exist, but there are nowhere more than 9 in the Asian forests.

Primate hiomass is highest in African rain forests where it may reach (iver 2 tonne km4. In Africa, and also in
Asia, there are numerous species specializing in eating leaves {folívores} which they are able to digest using
commensal bacteria that inhabit Inodified, extended parts of the gut. America has a larger number of
frugivore-insectivore species than elsewhere, none which are so dependent on leaves, and some species
which are much smaller than are found elsewhere.” ln both Africa and America mixed-species primate troops
occur, but none have been found in Asia. The propottitins of pri- mates to other fruit- and leaf-eating Canopy-
dwellers varies from place to place. New Guinea has more frugivorous birds than elsewhere because there is
only one Computing matnmal, a tree-kangartio, and no primates or sqtiirrels. South America is sometimes
called the bird continent. It has a particularly rich bird fauna, followed by Asia, with Africa trailing behind. In
the Old ‘iVorld tropics hornbills (Fig. 4.2) form a guild of specialist Canopy-top frugivores. This niche is
filled in America by toucans, slightly smaller in size but with similar beak construc- tion, which evolved
convergentiy to feed on the same sorts of fruit (section 5.2). Tropical America is also much the richest region
in bars,

Big. 4.2. Rhinoceros hornbill {Buceros rhinoeeros}. Hornbills are specialist frugivores of Old World tropical
forests.

68 Hair: ¡‘tn-est animrzis

and the fhITlElZÜH has aibout half 211i lcntawn Species of freshwater fish.

For many rain forest vertebrates fruits are an important component of the diet. This differs from the situation,
in tropical savantias where herbivores pretlominate and occur at higher biomass. For example, at the Cjocha
Cashu rain forest in Peru, mammal and bird biomass ‘NAS c. 2 tonne km4, whereas it can reach 12 tonne
km4 in African savanna. Fruits are variously‘ supplemented by other foods. For example, frugi- vorous birds
ziuglïletlt their diet with insccts, which are richer in protein, especially during the nesting and moultitig
season.

Besides differences betiveeti the three tropical regions there are other differences within them. One major
pattern is that within the African anti American rain forests there are areas of especially high species richness,
set like islands in a sea of relative poverty. This is shown vireil by ¿Xfrican birds and is paralleletl in America
by many groups, including birds and butterflies (Figs. 6.'l6,6.1?_l. The upper ¿‘rmazon near the Andes in
Colombia, Ecuador and Peru is excep- tionally rich. No such patchiness has been detected in Asia where the
Inajor ptittern is set by \Wallace’s Line, one of the sharpest zoogeti- graphical boundaries in the worid and
which delirnits the continental Asian faunas from the

IOÜr
Insectívores
80"
Si

‘a 6D- CJ 3 "5 g ‘lÜ- Frugívores 2B _ Nectarivores Ü|ï.,_l_ l l I l l I l 400 8ÜÜ 1200 i500 ZÜÜÜ 2400 280D
3200 3500
Eievation im]

Fig. 4.3. Reduction in bird species numbers of three feeding guilds with increasing elexratitm. Peruvian
Andes. {Data of Tcrborgh in Bourliete 1983.)

Atistralasian (Fig. 6.5). These patterns are now realized to have explanations based on Earth history as will
be discussed in Chapter 6.

Finaily’, in» all parts of the tropics there is a general diminutioiï in species number with increasing elevaticin,
from lowland to mtïntanc forest (Fig. 4.3}.

4.2. MODES or COEXISTENCE

Studies have now been made in many parts of the tropics to elucidate hoiv so Inany einimals co-cxist in the
same forest. They do so by special- ization. Rain forest is lofty and provides a diverse threc-dimensitnïal
living space and a huge variety‘ of fOOLlS.

Specialízation in time and space

Some animals are aictii-‘e by day and others lay’ tiight, and many species live mainly in a single layer of the
canopgr. Figure 4.4 shows this vividly for the mammals in a rain forest in Borneo. The canopy laycrs
occupictl by the dif- ferent birds at La Selva, Costa Rica, are shown in Table 4.2 and hy nine sy-‘mpattic
squirrcls at NFPassa, Gabon, in Fig. 4.5. Invertebrados have also developed particular nichos in time and
space. lll-l. Murphy has describetl how at the Bukit 'l‘imah forest, Singapore, there are morning and evcning
rush hours when ctypto- zoans move from the soil to feed in the Canopy and back, aivaited at the foot of the
trees by a predattiry SCÜÏPÍOII, frog, and spider, while during the tlay these dawn 2111€] dusk predators are
replaced by‘ others, including a flying lizaird. Murphy has also tlisctivered that insects do not fly at random
through the forest canopy but follow particular, ¡areferretl flight paths, and that along these are concentrated
the web- making spidets which prey on them.

The water-filled tanks of epiph_vtic bromeliads are a particularly special niche in the ÜCÜÍFOPÍCS (Fig.
4.6}. They are ‘vetitable aerial aquaria. [n jamaica 68 species have been found to inhabit them, including
mosquito larvae. in fact,

Modas of coexistence . 59
Day
Hylubatid [l]

Bay and night

Pan

/__q_—= 4k ¿q?

Scíuridial

É
CH
Tr;
É
Sciurid . ,. w s
Eewidfl) ' " = _

.5 í Ennaceid {l} É Vivenidlïl fi- .

Uivenid EZ}

MLISIEII-d

Suricidlí]

Ï Suid [l]
Histricid {2}
Tragufifl
Álfiurid {'13}

Fig. 4.4. Spacc- end tinlc-partitionirlg of ¡Ion-Ílyirlg mammals inche lowlancl rain forest of Sabah. (Data of
lvíacKinnon in Whitlnorc 1984a, fig. 3.1.)

Tupa i id {3}

Felíd E4}
Table 4.2 Bird zonation at La Selva forest, DBCIEHSÍRQ body Size

' Costa Rica 3o-

Above Canopy Vultures, hawks, swífts 25

Canopy top Toucans, coringas, parrots,

cacique birds É 20

15-25 rn Wïoodpeekers, woodhewcrs, E

large rrogons, jacamars, 15

_ puffbirds

Understorey" Most hummingbirds, 10

antíbirds, nmnakins, flycarchers, tanagers 5

Forest floor Tinamous, great eurasmw,

ground doves, wrens Ground

Data Of Slüd ¡f1 BOUFÏÍÉFC (1933) Fig. 4.5. Canopy height preferencias of nine sympatric

Squirrel species in lowland rain forest, Gabon. (Data

of Ernmuns in Bourliére 1983.) Note how similar heights in the Canopy are oecupied" by squirrels of very
different lmdy weight.

70 Rain forest (¡nin-tab

Fig. 4.6. Epiphytic htoïneliads are a very tiistinctix-‘e

feature of the neorropics. ¡‘tmazonían Eeuaidcit. Wïarer collects En tanks at the overlapping leaf bases and is
ahsorhed hy the plaïnt‘. This pool provides home for many aniniailcules, including mosquito larvae.

malaria-control in the New World, unlike the Old, can never he hy the eliminarior] oF stagnant Water
hecause of the presence of these ‘water- tank epiphytes. '

The pitcher plants hiepentfaeis of Madagascar, Seychelles, and the Eastern tropics provide another special
niche for animals (Fig. 4.7). In Maiafm 55 insect species have been recorded as inhahitants of the pitchers,
two-thirds of which live and lnreed there. Amongst the nïore Surpris- ing occasitmal inhahitants of hoth
laromeliad tamtks and Nepentbes” pitchers are species of Íreshwater crah.

Fig. 4.7. Nepembtïs sairgreirïea, hilalajxra. These pitcher plants are a feature of the Eastern tropics. Small
invertehrares, which fall into the pitcher, drown and are digested by" the plant.

Food preferenees

Many cases have now been z-inalysed virhich show how the animal inhahitants hdVC evolved to share out
the food I”(_‘.S(')L1['CCS of a forest.

Neotropical hats are hy far the most complex assemhleiges any-‘virhete in the world of sympattic mammals,
with 35-f50 species co-existing in a few square kilometres of forest. Amongst the 35 sjrmpatric species of
Barro Colorado Island, Panama, nine fotidndefined guilds can he distin- guished (Tahle 4.3), each ‘with from
one to nine species. Vkïithin each guild there is spccialization rnaitily for size of food particle, which is itself
proportional to body weight.

In northeast Gabon in. Central Africa there are five synipatric nocrutnal species of iorises (prosimian
primates), a porto (Perodicïrus porto), three bush-bahies [Gaiago aliens", G. (Íenrfidoviï, Ezroliciss
elegantuhrs}, and an angwemtiht) {Arctocebtts calabarensis). These avoid competition hy pairtitioning hoth
food and space. In the Caflüpy G. demidottii is mainly insectivtirous, the potro mainly eats fruits, and Euticus
feeds on plant gtims. The FWD other

(Íaarrying capacity Off-tiñe forest _ 71

Table 4.3 The nine feeding gnilds found amongst the 35 hat species at Barro Colorado Island, Panaima
1 Frugittores ln canopy’

l " Near the ground

3 ” As scavezigcrs

1' Omnivtires lieeding on nectar, ptyllen, fruit,

and insccts

5 Sanguivores The iranipires, feeding on blood Carnivores (‘Which glean}

7‘ Piscivores {Fish eaters)

S Insectivorcs Sirius-flying hawkitlg
9 " Fast-flying hawking

Data of Barrocorso in Bourliére {i983}

species inhabit the undergrowth being mainly Ítugivortius (G. flffíïíif) and insectivorous (the atigwantilaru),
respectively. There are additional differences in hunting technique, with speeializa- zion on slow- or fast-
moving insects.

Amongst symparrie primates diet is always related to body size. The smallest species are insectivores, but
insects cannot supply’ enough nutrition for medinm-sized einirnals in relation to the hunting effort, and fruit
is the most important sort of food. Por the largest species, ieaves and seeds are ITIÜSt important and they
also eat some fruit. This is well exemplified at (Ïïocha Cashu described below.

There are also some instantes of intricate co- existence between insects and their plant feeds.“ One is Uraizm
fitigens, a day-flying moth of Central America, which is well known hecause it Inigrares in dense clouds over
long distantes. Research has shown that these moths are in search of a fresh food source. Their cater- pillars
feed on the leaves of the forest climher Omfhhaiea. After (-1 few generations the plant counterartacks by
producing a toxic zilkaloid, DB/IDP, structurally‘ similar to fructose and a powverfnl insect-feediïig
deterrent, that inrerferes with sugar metabolism. These discoveries, start- ing from pure cnriosity to explain
why a rnoth migrates, have led to investigations into the pharmaceutical ptitential of DMDP for the treatment
of cancer, ohesity, diabetes and AIDS.

A second example is the South American leaf- cntter ant Aira, whose species tend to tavtnid taking to their
fungns-gardens the leaves of species that are flingicirlalggi’

Breeding sites

Another yvay species may live together is demonstrared by the extremely rich frog and road community of a
forest at Santa Cecilia in Amazonian Ecuador, 110W’ nnfortunately destroyed. There the 74 species, all living
within 3 km3, were tnainly’ OPPOIIÏUIIÍSÍÏC feeders and differed mainly hy a wide ClÍVEISÍEY of
hreerling sites. Ten modes of egg-laying hehaviour were distinguished, including deposition in water, on
vegetarion, in tree cz-ivities, in a depression in the soil made by the Inale, in foam nests, and on the back of
the female. There was also some teni- ptiral tliivisicin, only ten ot fewer species being found to hreed
simultaneously.

4.3. CARRYINC} (SAPACITY or THE

FOREST

Ultimately, hy whatever manner the animals have evolved to co-exist within a particulzir forest, their
numbers are limited by the amount of food av-ailalwle.

Data have been gathered from several rain forests which show how there are periodic lean periods when
animals may’ go htlngry’. ’l'he (Ïlocha Cashu area of the ivlanu National Park in Amazonian Peru may be
used hy way of Example to show how‘ animals suhdiisitle the resources of the forest and cope with hungry
pericidsdü” At Cocha Cashu the dominant inamtnals and birds are ffUgiVÜfCS, contributing about three-
qnarters of the total ‘vertehrate hiomass. Amongst the mammals, primates then rodents and pcccaryv’ are
commonest. Fruit pro- duction was estimated to he 2 tonnes ha” year”, but with a 3-month lean pcritïtl, Balay
to

July, ivhen alanndance feli to lverween 2 amd 5

per cent of its peak tlnantities. During the lean period frugivores are forced either to migrate or

7,2 Rain forest animals

to alter their diet. Five of the eleven primate species were studied in detail. These were found to shift to
different foods and what they are depended on their size. The two largest species, the capuchin monkeys
(Cebus ' albífrons, C. apella; 3000-5000 g) concentrate on palm nuts tvhich they are able to crack x-vith their
strong jaws, but only the larger species is really strong enough. The squirrel nionltey {Saimiri sciuretrs; 800-
1200 g) eats figs, but stimctinies goes a sveel-c with no food or just insects, using up more energy than it
gains. The two tiniest species, the tamarins (Saguinus fuscíctïvllis, S. inzperator; 400-500 g) were observed
to spend 90 per cent of their feeding time sippitig Iiectar. Only about ‘l per cent of the 2000 tree species in
the area fruit during the lean period, and these are essential to sustain the frugivorotls animals.

There is evidence that palrns and figs also play an important role as farnine foods in other South ¿american
forests. Such plants have been called keysttine species because of their vital function. They have obvious
importance in forest conservation, and we shall discuss them further in section11.3.

Figs as keystone species

The bustling activity at a large Hg is one of the unfor- gettable spectacles of the [Cocha Cashu] forest.
Nionkey troops arrive from all directions as if guided by some rnysterious perception. ‘We have seen over
"E00 monkeys of five species and 20 to 30 species of birds fecding simultaneously’ in a single Ficus perfo-
reto. Htiw is it ¡itissilnle that so many animals inde- pendently discover a tree almost the first day the fruit
ripens? ‘We believe they are sunnnoned by the shrill din of the niyriads of parakeets (Brotogerus spp.) that
quickly eonvergc on the scene. These birds are fig spe- cialists as much as Saimiri, and the sound of them in
numbers is almost certain indicaïtmn of a fruiting treedf"
Certain of the big tree fig species of western Malesia, mainly stranglers and banyans (Ficus sect. Urostígma;
Figs. 3.3, 3.5) are just as attrac- tive and important for birds and mammals. Fifty hornbills of four species
have been ObSCYVCLl

feeding simultaneously in a single fig tree at Gunung Mulu, Sarawak. At Kuala Lompat, Ixíalaya, 60 bird
species feed mainly on 38 Ficus species in an area of 2 km2. A studymï at Kutai, east Kalimantan, found that,
as at (Jocha Cashu, certain figs were the keysmne resource for many mammals and birds, and provide the
baseline food because they bear fruit throughout the year. Fig fruits are supplernented by fruits of clirnbcrs of
the Annonaceae and by some Meliaceae and Nljrristicaceate. At both Ktlala Lompat and East Ktltai
individual fig trees, and indeed species, fruited sporadically, but collect- ively these figs were always fertile.

There is no record of figs as keystone species in the African rain forests. In a Ciabonese forest it was found
that figs were rare, and mainly‘ fed on by wide-ranging hats. Mon-cover, they fruited sparsely and
sporadically. ivïonkeys and large birds relied on two Dulyristicaceae and an Annonaceae for keystone famille
frnits.1°5

Carrying capacity and Conservation

Some animal species exist at low population densities either because of the intense competi- tion for food or
living space or from the con- stant attentions of predators. This is true of many invertebrates, and also of
lower canopy birds, whose total bioinass is not dissimilar from that in a temperate forest but is divided
between numerous rare species instead of fewer abun- dant ones. Rain forest birds of the lower canopy
characteristically’ breed seasonally when insect food is particularly abundant, lay’ only one or two eggs, and
have a low rate of success in raising fledglings. Although each species occurs at a low density of only a few
pairs per hundred hectares of forest, subadults can be seen wan- dering around trying to find living space.
Adults are long-listed. It seems likely that a spell of lean years could lead to local extinction of rare species,
followed later lay ¡‘e-invasion from adja- cent forest when better conditions return. XWe have little data to
substantiare this specularion, and long-term monitoring of populations is required. lt is a reason for concern
because as

Rain forest nninrals — Cfzrzpïet‘ srmtrrtrtr-y 73

rain forests become fragmented into isolated, small, relict parches, there may not be enough space for the
survival of those species that exist near their lower sustainable limit of density (see p. 227}.

4.4. 1mm FORFST Anos-mts-

. CHAMER SUMMARY

l. Rain forests are very rich in animal species. Numbers diminish with elevatitin and away’ from the rropics.
There are examples between zhe three regions of convergenr cvoluricin, for example, amongst specialist
frugivtirtius birds. There are also differences, for example, neorrop- ical bat diversity has no parallel
elsewhere.

2. ¿‘Xmerictïil and African rziin forests have parches of species richness and of relative poverty. In the
Eastern tropics the main zoogeo- grz-iphiczil pattern is associated with Wallaceïs Line which separates Asian
and Australasian Saunas.

3. These many animals co-exist in the same forest by sharing the resources in numero-us ways. This is shown
by‘ considering related groups, for example, primates or hats. Many exhibir preference for a particular layer
in the forest Canopy’, and are ¿active either by day or by‘ night (Fig. 4.4}. There is strong specializarion for
particular foods. There may be differences in time and place of reproduction.

4. For many rain forest verrebrates fruits are an important food. The amount of fruit varies through the year
and the number of animals the forest can support is determined by the amount produced during the lean
.period. A few so- called key-stone species fruit all the year round and provide the baseline food. In America
and Asia certain figs are important keystone species.

5. Many animals occur ar vwery low popula- tion density and probably suffer local CXEÍIICÉÍOH during
exceptionally lean periods, but re-invade when conditions inlprtive. Such species are likely ro" disappear if
rain forests are reduced by man ro small relict parches.

Inter/connections between plants and anímals—

the web oflife y

Animals are dependenr on plants for food, consumers on producers, heterotrophs on autotrophs, and forests
also provide diïrerse litring space for their animal denizens, as was outlined in Chapter 4. But tropical rain
forests are more than just a restaurant and a home. In this chapter we explore aspects of the complex web of
l1‘1t{:I'C()l'l]'lCCElCJflS ivithin rain forests which scientists have begun to disenrangle over recent years.
Plants make use of ¿‘inimals as pol- linartirs (very few are wind-pollinared in con- traSt to temperate
forests), and many use them also as dispersers (though wind tlispersal also does occur}. lt has recently been
faShi()I’13l)lE3.t{) invoke the terrn co-evolution, in a (ine-to-one relationship of specializarion, as a driving
force for species evolution. Some symhicitic relation- ships do bear this stamp, but in most cases the forces
of natural selection are now realized to be more diffuse.

5.1. ANissiALs AS 1*oLLii\-';t‘1‘oits“’4 The same combinarions of flower and inflores- cence features
which artract particular pollin- ators, so-called character ‘sy-aidroines’, OCCLII’ in all parts of the tropics.
The plant atrracts its animal visitors with both nectarm and pollen.

Bird flowers are mhust and harsh in colour- reds, oranges, ycllows, and greens; the necrar is warery and
copious. Firmirmu (Pig. 5.1) is a good example. This syndrome, as cithers, is rec- tngnized by the local birds
when a [ilant is culti- vared OLIÉSÍClB its place of origin. Bird flowers are held awajv from the leaves, or
are hornc when the crown is deciduous.

e»

Fig. 5.1. Firnriflrtn rriafngvtntt. Malaya. The robust inflorescence home on the hare crown, with XFÍVlCl
(Jrange flow-ers is typical of species adapted to bird-pollination.
Animais as jiollínators _ 75

Eig. 5.2. Kosrersnarssia maiayarxa like many other Bomhacaceae has flourers that are nocturna] and
ramifloroizs. They are pollinated by bars ‘which brush pollen on to their body while seeking at. the peral
bases for ncctar. (See also Fig. 5.3.} Malaya.

Fig. 5.3. Paebira macrocarpaof tropical Africa, and also in the Bombacaecae, exemplifies the other kind of
bat-pollinated flower (ef. Fig. 5.2). Mengla Botanic Garden, Yunnan, China.

Bat pollination involves two different syn- dromes. Bat flowers open at night, have viscid nectar, often a
musry’ or sour scent, and are often dull cream sin colour. They are either deepiy cup-shaped, as in Dario and
its relative Kostermansia (Fig. 5.2) and pollen is conveyed on the hafs head, or else they have a mass of
exposetl stamens, like a shaving btush, which deposits pollen on to the hatïs breast; examples are Parkia and
Pacbíra (Fig. 5.3), Bats, like

Fig. 5.4. harina spcciosa showing the pendulous

inflorescenccs which hang free, where the poiiinators

can easily reach them i-vithout coliision with the

crown. Planted fur its garlic-like seeds. hdalaya. All species of this pantropical genus [Leguminosag
subfamily Mimosoideae} have drumstick-likc infloresccnces held free of the foliage. This species is ba[-
>p0llÍ.T'I€ltt3(l. ln others, pollinarioil has been recorded by hees, buttcrflies, birds, or in Africa by the porto,
a noerurnal primate.

birds, cannot easily penetrate foliage and hat flowers are either borne outside the crowvn, sometimes on long
proeesses-so-ealled penduliflory shown Well hy bananas (Mr/rra), Oroxyhtm, and Parkia (Fig. 5.4), or are
bornc behind the leafy tivigs on the limbs (ramiflory; eg. Kostermansïa, Fig. 5.2) or on the trunk (cauliilory,
Fig. 1.2).

ü There is sometimes over-lap between hat and bird pollination. MHCIÜIG, Spatbodea, and

76 Issïercortnertions ¿’Jetzueen plants ¿ïïíd (¡animals ——- the web offife

fig. 5.5. Ixora‘ Irabbií showing the ¡ong ctwrolla tube characteristic of moth-pollinared flowers. lvïalajra.

pethaps Erytfsrina ‘ere visited (though perhaps not pollinatecl) hy both groups. o '

Moth flow-ers, like bat flowers, are mostly

nocturnal. They are commonly pale in colour, sweetlyr scented, and have a corolla tube which is often very
long and ClÜWWJ ‘¿which the rnoth inserts its proboscis to reach nectar produced at the base. Rairdicï and
other Ruhiaceae are good examples (Fig. 5.5}.

Beetle flowers, many of them highly’ fragrant, are as characteristic of the tropics as bee floveers are of
temperate and semi-arid lancls. Beetles pollinate by clumsy scrambling, eating pollen as they go, and many
heetle flowers are open tiishes or bowls tisually set ¿unongst the leaves.

Fig. 5.6. The giant South Amcricail aroid Dracontiurvn gigas with a single

multipaltnate leaf {A} 4 m tall, and inflorescence (( ) 2 m tall. (A. Engler and

Prantl, Pflanzenfi/wtíliert II {3} 1889.}

The Old World counterpart is Anrorpkvoplsoifrcs some of whose species are

¿ilso gigantic.

jlníznals ras polliszrztors _ 77

Ánnonaceae, Magnoliaceae, iVlyristicaceae, and zhe palm falïïil)’ provide Inany eïamplesloü

Bee flowers, so common in tempcratc cli-

rnates, are also extremely common in the humid ïropics and are perhalas the major flower-tjirpe. Bee flowers
are diurnal, often zygomorphic, Ïorightly’ coloured, sometimes ivith lines of colour {which may only he
visible in ultra- ‘¡iolct light to which hees are sensitive}, and ¿re provided Vvith fOOIÍhOlClS. Orchids, and
íeguminosae subfamily Papiiionatae are prime examples. Some orchids have evolved exceed- Ïngly Conlplex
mechanisms, which in some species the intended pollinator has learned to ‘¿jr-pass by chewing through the
corolla base to teach the nectar.

Poul smelling, mottled, hrown and Carrion-

¿oloured flowers attract beetles and flies by ieception and often trap them; zin-iorpbopballus ¿nd
Draeorztizmz (Fig. 5.6}, Arisrolocbirï and Rafflesia (Fig. 3.13) are examples.

A recent tlisctávery is of sequential flowering

7.o species which share the same pollinators. The

Ííkclihooci of successful cross-pollinarion is enhanced, firstly by eonspecific plants all bloom-

SimLlltaHEOuSl)’, and additionally, by hloom- at a different time from other species. Four ‘Jr the five i-
l"elier)nicz species at La Selva, Costa Rica, share their humming-laird poliinators in zhis manner and
withother food-plants (Fig. 5.7}. Six closely related Show/ea in Nialaya simi- Zarly share the same thrip
pOllÍHQÍKJFÍLIÜ? Also at Pasoh, Malaya, ten unrelated species, pollinated h)‘ the same carpenter bees
(Xgrlocopa), have

been discovcretl to have staggered flowering times (Fig. 5.8}.-

Deeper intricacies, to promote cross- rather than self-pollination, continue to be tliscovered. For example,
Oroxylum in Nlalaya iaroduces c. 18 ml nectar per night hut in hursts of 0.05 mi each.“ This stimulates its
bat pollinator to travel from plant to plant, periodically rctttrrl- ing, a mode of foraging that has been evoca-
tively’ termed ‘trap-lining’ hy analogy to a hunter x-vho sets and monitora a line of snares.

A distinction has been discovered between the

three nectarivoroias [wars of Malaya which matches them to the phenolrggy of the species they depend
onfmg Two hats feed on the nectar of plants that Ílox-ver continually, banana and Son-trataría (a mangrove
genus}. These hats, A/Iacrogiossus solJrintrs and M. ntiníímrs, respec- tively‘, roost singly or in small
groups. The third

Maimiirl A slo N e
JF

Fig. 5 .7. Sequential Ílowerng of the tindetgroxarth species fed on and pollinated by herinit humming lairds.
Lotvlanti rain forest, La Selva, Costa Rica. The five HÏEÏÍCOHÉG are started. {From Stiles 1977.] 1, i-
leliconia pogormrba; 2, PrJsSÍflrJI-a ‘LJÍÉÍÑLÏÍCI; 3, H. wagneriana; 4, ¿Ïacobinia aerea; 5, Cositas y rather;
6, Heliconie sp. 18; 7, Helieonirt sp. 16; 8, Apfseíartdra sincíairicmcr; 9, (Ïrastus ntaíortïeanns; 10,
Helirronict sp. 3.

Xanthophyffum discolor
Xanrhophyfíum rufum

,.

Connarus sp.
Síndera veluhha
Síndora cortacea
Pa/lesua ¡’errea
Calhearpa mamgayí
Millerila atropurpurea
Cannarus sp.

él...

March April May

i I “ June

Big. 5.8. Scquential flower-ing of ten species all pollinatcd by the same Carpenter hces lXyrlocra-yirt).
Malaya. (relppanah 1985.}

78 lnterconnecïions between gilants and animals — the zrxeb oflifr’

hat, Eonyctei'ís spelaea feeds on tree species that only flower once or twice a year, including the important
fruit trees durian and petai (Dario, Paiulia Fig. 5.4). Eonycrei'is inhahits caves and lives in huge colonies,
ivhich emerge at dusk, like a plume of smoke, from crevices in the rock, and may have to travel many
kilometres to feed. Their gregaritius living is helieved to enable them to communiceite to one another the
loca- tion of patchy food sources. Destruction of forest within the bars’ range reduces the food supply, made
up of many species each essential for part of the year.

ÁNlLVlfXLS AS DÍSPERSERS Two kinds of bird fruits can be recognizedd” Non-specialist ftugivores feed
on fruits with watery, sugary flesh and small seeds, which provide only part of their diet, rnostly carhohy-
drates. These ftuits are common in secondary forest, for example, Melastomaraceae and Trama, and the birds
that feed on them, often congregate in flocks. By contrast, fruits evolved for dispetsal by specialist
frugivorous birds provide a higher quality diet, rich in fats and proteins; the seed is large; many are drupes.
Lauraceae contain many examples in all three rain, forest regions, plus many palms and Burseraceae in Asia
and America. Other special- ist bird fruits are hrightly coloured, dehiscent, and arillate. This syndrome is
found in many families. It is well exemplified by the nutmeg family, Myristicaceae, in which a yellovirish
peri- carp splits open to exposc a vivid red edihle aril, partly concealing a hig black seed (Fig. 5.9). Both
types of bird fruit occur in Elaeocarpaceae: Elaeocarpirs has an oily drupe, Slomzea fruits are dehiscent and
arillate.

At Ivíakokou, Cíahonfi” fruit type was found to be more closely related to mode of foraging and energy
needs than to animal taxonomy. This forest had 3.9 species of fruit consumers {seven large eantipyr birds,
eight small and two large rodents, nine squiiïrels, seven ruminants and six monkeys). Analysis of 122 species
of

Fig. 5 .9. ¿’big-visited fmgmns, the nutmeg, has fruits

adapted for dispersal hy specialist frugivotous birds

{Blume "l 835, plate 55.] The fawn fruits dehisce to exposc a single, hig black seed, laartially’ enclosed in a
finely divided, Scarlet aril.

The Dutch East India Company tried in the y seventeenth century to control the spice trade hy confining
nutmeg cultivation to just a feiv islands of the Molucczis, hut iverc thivairted hy nutmeg- dispcrsing pigeons.

fruits eaten by these showed, as prime distinc- tiotis, that the birds amd monkeys favoured brightly coloured
fruit with succulent pulp or arillate seeds, whereas the ruminants and large rodents favoured large,
indehiscent, fihrous fruits. .

There are no well-defined character .syn- dromes for mammal-dispersed fruits (Fig. 5.10}.

Animals as dispersas _ 79

Civets, rnongooses, and bears are probably attracted by a strong smell. Colours are often - áull and nmted.
Durían {Dario zíbethinus}, ‘ most famous fruit of the East, certainly has a áístinctivc pungcnt aroma. Tigers
are notorious Ïor their passion for durians. All these animals _ are mainly nocturna]. The close study of five
primate species at Cocha Cashu, Peru {section 4.3), found that, although all fed on ftuits, the only common
feature the fruits possessed was ¿rellow to orange colounm

Fig. 5.10. The siamaltg

{Hylofjntes syndactylus), a large

y ape of Malaya and Sumatra {weight 9-12 kg), seen here feeding on Agïaía, includes much fruit in its diet.

It has recently eorne to be realized that fish are major dispcrsers of fruits of the. riverine forests of
AHHIZOIIÍELlH They feed during the high tvater season in the riparian, periodic swamp forests. During the
low water season they retreat to lagoons and live largely on accu» mulated fat. No particular features
characterize fishwdispersed fruits. Two species of rubber tree, Hezxea brasilíensis and H. sprucearta, have
seeds that are a major food source. for certain Arïnazonian fishÏFhey occur in seasonal swamp

80 hiterconiiectioizs herzueen ¡Giants ana’ animals — the rare!) of life

forest {and in the case of H. brasiliensis also on dry land), and have big fruit capsules that dehisce
explosively to disperse the 20- to 40- mm-dianieter seeds for '10 m or more. Some seeds fall on dry land.
Fish congregate below the trees to catch the rest. Seeds that escape preda- tion float until the floodwaters
recede, at which time they reach the soil surface and germinate.

The relationship of particular animal dis- perscrs to particular fruit-bearing plant species is less close than
with pollination. Temporal partitioning has ÏIOWEVBI‘ been discovered. For example, in Trinidad there
occur19 bushy epi- phytic Mieoizicz species, Melastomataceae, which bear hrightlyr coloured sugary fruits,
typical of the felmily. Each species only bears fruit for a short period and these perinds do not Overlap,
Which enhances the likelilïood of díspersal for any particular species.“ It is, hOWCVEI‘, difficult to
determine in this and other examples, how much selection pressure there has been to evolve non-overlapping
fruiting periods because, with nincteen species each of which only fruits for a short tvhile, there is a strong
prolaability peritids will not (iverlap purely by chance and, as described above (section 3.3), flowering hnd
fruiting are tindet the COIITHJl of a plexus of internal factors iuteractiug with climatefl“ [n the Makokou
study LlÍSCLISSCLl above, climatic factors were paramount.

5.3. PLANT WEBS, MOBILE LINKS,

AND KEYSTONE SPECIES

A rain forest may be universally green and monotonous to out eyes hut a herhivore sees it as a poisonotts
place with parches of alkaloid, saponin, or siliceous spicules, and seeks out what it can safely eat.“ Plants
have evolved toxins that deter herbivores, and related groups are constrained by their genetic constitution to
manufacture similar Chemicals. Thus, for example, Apocynaceae are rich in alkaloitlsf” especially in the
milky latex that nearly all possess. Drug companies exploit these similar-

‘ities when screening forest plants for potential

new medicines. instead of an expensive blind search they concentrate von those families most likely to be of
interest.

Herbivores haire evolved adaptations that enable them to detoxify particular chemicals. Thus there have
¿IÍSCH guilds of insects all spe- cialized to feed on a particular group of related, and therefore chemically
similar, plants. Such a
group may be called a plant webfil” In the forest '

many parallel insect guilds co-exist on these dif-

ferent plant webs, svithout tuverlap of feeding ‘

sites?" In some cases the insects utilize the chemicals they ingest. Birthving hutterflies and some other
Papilitinidzie feed on the herbaceotis climbers of family Aristolochiaceae, which provide them with a
distasteful chemical and

render them Lmpalatable to predators. The_ butterflies can therefore afford to congregate v

near the plant, and this facilitates mating. They are mimicked by other butterflies and also by day-flying
moths.

Two complex plant tvebs have beendisentan- gled in the neotmpical rain forests, one involv- ing Solanaceae
and the other Passifloraceaem

There are e. 500 species of Passifloraceae in the New ‘¿World tropics, as ‘wiry climbers and small trees;
Passiflora is the biggest genius. The main insect feeders are Heliconiine butterfhes, along with flea beetles,
coreid bugs, and diopter- inc moths as less important herhivores. At any one locality there are between ten,
and fifteen passiflores of various habits and tuccttpying dif- ferent fiabitats. These are pollinated and dis-
persed by generalist bees, birds, hats, and moths. The pollinarors and dispersers cannot obtain their food
solely from Passifloraceae because flowering is sporadic. Instead they feed on species of different plant webs
at different times of year, and thus may be called mobile links because they bridge (wtherwise ¡Jnconnected
components of the forest ecosystem.

Ifa forest is degraded partly to pasture, svhich is happening extensively in parts of the neotrop- ics {Chapter
lO), some of the local passifltire species disappear along with their hatbitat. Likewise, if an area becomes
entirely clímax ftirest, there is also a loss of diversity in the

(Ïta-evolutíran - _ 81

plant webs and a parallel loss in the dependent insect guild. Moreover, Heliconiine butterflies are the
principal model for major nlimicry com- plexes so loss of any of these butterflies dimin- ishes the numbers of
the (edible) mimics.

The plant web and associated inscct guild on neotrtipical Solanatceae is similar. ln this case ithoniiine
butterflies are the principal feeders and mimetic model.

Antither dimension of complexity is that the mobile link species which are essential to the reproductive
success of the plants of several sep- arate plant tvebs, are themselves comrnonlyr totally dependent on a few
keystonc species for prot-‘iding their food at times when nothing else És avatilable. The importance of some
figs as key- stone species for birds and mammals was described in section 4.3. The continuously tiovvcring
pioneer forest-fringe shrub illlelasroma malabarhriciim is today a keystone species at Pasoh, ivlzilaya, for
carpenter bees which are heavily’ dependent on it during lean periods. ‘Cïhcn the primary forest burst into
heavy flower in late 1981 these bees were observed to migrate from the secondary scrub with which the
remaining primary‘ forest is now SUlTOUfldCLl on to the new ephemeral rain forest food sourcem

In rhnerica, EUglOSSÍIIL’ bees are important mobile links. Thirty to fifty species may occur together and
any one bee species may pollinate a ¿tiren plant species, including orchids and aroids. Trap-líning (p. 77) is
Common. Orchids provide pheromones to male Euglossines vrhich are essential for courtship and hence
mating. The loss or zibsence of orchids thus causes the ecosystem to start to unravel. The faílures to set fruit
of plantation-grovirn Brazil nut trees is a good example of these intricate interactionsdï‘

The Bratzil nut (Bertbolletia excelsa) is ¡Jolli- nated by Eugltissines. I-Iosvever, these bees ¡reed some other
source of pollen for the 11 months when it is not in bloom, and they also need epi- phytic orchids to complete
their own life cycle. “Wild Brazil nut trees occur scatteretl through the forest. When concentrated in isolated
mono- specific plantations or remaining as relict trees after forest destruction they do not have the

linkages essential for their success. The huge, thickly tvwïiody fruits uveigh several kilograms and fall
heavily to the ground. They contain lÜ—2{) strongly ¿irmoured seeds, the Brazil nuts of Commerce. The
final link in the Brazil out life cycle is the agouti {lkisyprracïa}, a giant rodent, which gnaws open the outer
carapace and stores the nuts in caches scattered through the forest. Agoutis forget some of their nut boards,
which is how the scattered distribution of Brazil nut trees arises.

If any one plant-iveb becomes particularly abundamt, insects of its feeding guild ‘Will fhitl it easy to locate
in"the forest and may increase in numbers. ‘Pest pressure’ is likely alwayrs to keep any particular plantwvel)
at a low density‘, below a th rcshold at which it becomes too ‘apparent’ to its herbivores. This is o-ne
mechanism that maintains high species tiiversity in many eco- systems, including tropical rain forests. One
particular case of the reduction of tree sccdling population density ís discussed later in section 7.3.

5.4. CIO-EVOLUTION

Extreme richness in species of plants and animals is one of the best known features of tropical rain forests.
How it has arisen is one of the questions that scientists find perpetually’ fas- cinaring eibout these
ecosystems. lt is a question that has therefore sparketl off a plethora of dis- cussion and specularion. i

CÏo-evoltltion, i.e. close relationships between individual plant and animal ispccics, has been invoked as a
Inajtir motor of species evolu- tion.“ An arms race has been postulated in yvhich an insect species succeeds
in detoxifyring a protective chemical, feeds ‘well, and multiplies until the plant responds to this pest pressure
by evolving a new toxin. This is chemical warfare and its extreme form has sometimes been called the Red
Queen Hypothesisd-lvecause ‘it takes all tjhe running you can do to stay in the same placeïm But such
stepwise co-evolution seems, on inspection and reflection, to be improbable.

82 Inter/connections barragan plants and animals — the tve!) ciflife

In fact, pest pressure from the planes point of view is general, and toxic chemicals are devel- oped against all
its herbivores. ‘Arms races’ are unlikely to proceed as relentless and progressive bouts of defence and
counrer-dcfence. They are likely to be rnuch more erratic and to lead to the development of groups of
mntually dependent plants and animals, the plant-webs and their associated, herhivore gnilds ivhích were
dis« cussed above.

Speculation that particular ecological species groups of trees are more likely to have their leaves chemicallv
defended against herhivores than others has also not surviired scrutiny, Chemicals that are ptissibljv toxic are
indecd commonly present, but it now seems likely that ¡mechanical toughness rather than chemical corn-
position is the most important deterrent to insect herbivory. Most loss of leaf area in fact occurs before full
expansion of the lamina has taken place, because young leaves, which are soft and not fully formed, are
especially susceptible.

A study at Barro Colorado Island, Panama, demonstrates these general ctmclusionsdzó There, 70 per cent of
the 46 species ínvestigated had young" leavesdamaged more than mature ones. Even though their content of
phenolic cornpounds was 2.3 times higher, they were less tough and fibrous. In the same study mature leaves
of the pioneer species were grazed six times more rapidly than those of climeïx species, and were also found
to be less tough and fihrous.

Symbiosis

There are vrarious different rain forest examples of symhiotic associations between ants and plants.“ About
9D species of Hydrtopbytum, Myrniecodia (Fig. 5.11} and relatives, family Rubiaceae, centred on New
Guinea, have swollen tuhers, like irregular knobbly hand grenades, with chamhered, hollow intcriors that are
ant-inhabited. The excretions of the ants, and the humus that they accumulate nourish the plantdz“ Other
epiphytes may grow mi the humus. In the New World tropics certain tree-

Fig. 5.11. The aut-plant Myrmecoditï tirherosa with its swollcn srem cut open to show the internal cham- hers
inwvhich the antlridomyrmex cordarns lives. Note covered annruns on tree trunk. New Ireland.

living ant species collect seeds of various epi- phytes and plant them on their earthen nests to form aerial ‘ant
gardensïï” They collect animal faeces to nourish the plants which reward them by production of starch graíns
or sugary secre- tions from extrafloral nectaries. Ant gardens are particularly abundant in heath forest.

Certain climhing rattan palms are associated writh ants. Some Korthalsia species have a swollen modified
woody ligule, or ochrca, within whose concave interior the ants dwell. Should one accidentally brush against
an ant- Korthalsïa it emits a curíous whispering, rnstling, rattling noise, as the ants all rush out and heat their
mandibles on the hOllOW box against the attack. In other rattans, species of Calaïnus and Daemonorops, ants
make nests amongst the bases of interlockíng platcs of spincs on the leaf sheaths (Fig. 5.12), hut no benefit to
the palm has been demonstrared.

The Wiest African rain forest treelet Bar/taria fistulcisa is defended against large browsing her- hivores by the
aut Pacbysimrt aellfiíops whose stíng can penetrate elephant hide and which numhs lesset mainmals,
including humans, for several days.

The genera of pioneer trees Cecropfa (New

World; Fig. 7.6) and Macas-miga (Old World, mainly h/Ialesia; Figs. 7.7, 7.8} hoth have numer-

Co-euoltrtion . 8;»
5g. 5.12. The climbing rattan palm Dfl€’—?71t‘)?l()?’{)pS ÏÍPEÍÍKI of Borneo and Snmatrzi, showing the
inter- écking coinbs of spines in whose bases ants make ïïeir nests. {Blume "i835, plate 136.)

sus species that harbour ants, related groups of ¡‘azteca and Cremaïogasrer, respectively. ln ¿Ïecropírt the
trunk, branches ¿ind tixrigs are all hollow and prone to inhabitatirgn; in Macamnga ‘zst the apical few
internodes of the twigs (Pig. 5.13), ivhere the ant keeps scale insects (mainly ¿Ïoeears penangensis} which
tap the tree’s sap. The trees produce starch grains on the edges of ÍÏICÍI‘ recurved stipules on which the ants
feed. Experimental removal of ants has led to these lhieazranga spp. being attacked by leaf-eating ínsects.
Furthermuïire, these ant macarangas are ‘¿stiallyi free of epiphytes and climbers because

Fig. 5.13. ¿’Mcrcczranga trilobcr showing the holes made in the internodes by cohabiting ants. Note also the
recurvcd stipules on whose margins starch grains form which the ants feed on. Wïest Malesiaa.

the ants bite off any part of a foreign plant that comes into contact with their host (33 per cent of uninhabited
il/Ittcaraziga tríioha in Malayia had climbers on them compared to only 5 per cent of ant-inhabited onesLm
The ant- Maeai-anga story has a further mrist. Plants of several ¿‘r/Iacaranga species have been found with
their leaves badly eaten despite the presence of ants. Very careful searching has shown that the culprits are
tiny catcrpíllars of the Lycaenid hutterfly Arhopala, with never more than a few on any one tree. The
caterpillars produce a sugary secretion when touched by an ant, so are

84 fI1Ï(??'COPIIí8CÍÍOIÍ5 between pfflHÍS ¿md ¿animals — the reel} ofïlife

rudimeniary ' wings

wasp

escapes from gall ÏlOVJK-ÏIT wasp inside gali flower ¡_ Female l} fig wasp I G. seed develops l C. virasps
mate inside fig along style short style ¿‘nl D. Fenilized female F. female flower: H. gall flowen Graw“; Out
past pollinaied egg Iaid IHSIClEI Scam a,

mouth

E. Flies tu tree
Enters young fig «;,____—____ _ ,

“nm young “gs Male flower, usually near mouth

Fig. 5.14. The pollinaritm of fíg flonrers is one of the most inrímare plaint-tmimiïl relationships known.
(Partly

after Corner "i988, figs. 162, 164.] Special fig vi-‘asps, a different species for every species of fig, develop
inside grill glowcrs {el}; and bitc their xsiay out {B}; they mate inside the fig (C); the male dies but the
female eran-ls out of the mouth of the Hg, getting pollen on her body as she passes the Inale flowers (D); she
flics to another tree. probably EllÏlIFElCtCd by the sccnt of ytiung figs (E); the young fig either contains
female flowers with long styles and these are pollinated (F) so develop a seed (C), or the {ig Indy contain
Stcrlle gall flovrcrs with short styles tlmvn which the female inserts her OVipOSÍÍCJT and lays an egg (H);
this harchcs. the gruh feeds on the gall flower ovary and dcvelops into either a male or female xxiasp, which
then rcpeats the cycle.

lnzfnïications for forest Conservation _ 85

tolerated. They are coloured like the stipules or leaf nerves of the Macau-mago and in this way escape
ptedation. The hutterlly has evolvecl to parasitize the ¿int-iïríaccïranga syinibitisisfl“

Other equally complex mutualisms have also

Ïoeen cliscovered that do not involve ants. One such is between Passifloreiceaie and Heliconiine butterfiies.“
The butterfly lays its eggs on young shoot tips and the caterpillars feed there. The ntother butterfly’ seeks out
Linoccupied plants but the plants have evolveci either yellow‘ extrafltiral nectaries that tnimic eggs, or tii-
ange- yellotv modified stipules that imitate young caterpillars. Helieoniine butterflies have Eïitïlvetl an acute
sense of vision and an ability to learn. it can he observed that they counter these defences by prohing
potential layfing sites With their antennae and forelegs to see if they are really occupied. The Passifloreiceaie
have excep- tionally polyrnorphic leaves, variable looth srithin and lnewveen species; it is claimed this
enables them to escape detection by the butterflies. What lends credence to the claim for parallel evolution
with considerable mutual influence between buttetflíes and passiflores, is that the more complex avoidance
and detection systems occur in hoth the inseet and the plant in those species which taxonomy suggests are
more highly evolved, whereas the most priinitive species have the simplest aclaptations.

A particularly close symbiosis is that lnetwccn ¿and the tasps that pollinate them. Etrery species of fig has its
ox-vn wasp, and ‘where a tig tree is cultivated rïiutside the range of its pollina- tor fertile seeds fail to
develop. This intricate microcosm of evolution is described in Fíg. 5.14. As a further complication some
wasps have a species-specific parasite (Fig. 5.15).

5.5. IMPLICATIONS FOR. FOREST

(Í ONSEIÜEATIC} N

Tropical rain forests are hecoming fragmented into small relict parches or sirnplified in struc- ture and
composition hy exploitation for timber,

Fig. 5.15. The lig to vvasp relationship is rcnclered even more complex by parasitic insects called Inquilines
which lay their eggs inside the developing Hg xyasp grub by pushing their very long oviposter through the
wall of llalf-gtïfflit-‘lï fig fruits right into a gall flower. {Corner 1988, fig. 164.)

as will be discussed further in Chapters 10 and ‘l "l. lnterconncetiotls between plants and animals in the eco-
system, Which are only now beginning to he studied, are essential for its functioning, and are prone to
distuption. The ecosystem may then slowly’ collapse. Trees live for perhaps a century’ or inore (Fig. 7.35), so
are likely to persist for decades after potential polli- nators and dispersers have gone. At the Sungai Menyalei
forest in Malayra, which ctJnsists of only a few relict hectares as an island in a sea of tubber plantation,
fleshy tnzimmal-dispersetl ftuits of Cbrgrsopiaylluni aroxburgbis" nousadays lie in piles around the parent
treeÑÏl-l That forest, which to the causal Observer still looks fine, is in fact breaking down. HOVUEVCI’, as
was discussed above, plant species are not always tlepcndent on only a single animal species for ¡aollination
or dispersal; where relict forest parches ÜCCUI which are big enough to retain some suitable animals and
where hunting has not extermi- natetl them, the ecosystem may continue to function even though it has an
inconïplete species assemblage. lt is a matter of high priority for biologists today {section 11.4) to explore
and monitor the changes that take place in forest fragrnents, tvith a view to manipulation if necessary to
conserve species richness, because in a century or so from now much of the svorltlï; tropical rain forest svill
exist as frag- inents. Can these hold the full species comple- ment? As yet we do Ilot know.‘

86 Irziïercoimecfions beat/een {altar-its ¿md aztizncils — the suela of life

5.6. PLANT BRÉEDÏNG SïKïTlih-‘lslll

Plants have evolved the stations syndromcs of floxsrei‘ and inflorescetrce characters tlescribed above, to
attract particular animal ptallinators which then convey pollen from flosvei‘ to Íltiwer. Where pollinarion
occurs between different plants the amount of generic recombinatioti is ltigh. These are so—called
(Juthreedittg species. Genetic recombination is less in self-pollinarcd species, so-called itibreeders, in which
flowers on the same-Julant are invohretl, and is zero in apomicts, viz. those species whose reproduction
bypasses the sexual process {zilthough many apomicric species do also reproduce sexually on rare
occasions}.

The question has been delaated \\«'l'l(:l'.l]€l' the prevalence of one ¡Jartictilar bfEBCliflg sy-‘srem is

the driving force that accounts for the enormous

richness in plant species of tropical rain forests. A. Fedorov observed that many rain forest species occur as
scatreretl ÍIILlÍïFÍIÏlLIBlS, and spec- ulated rhar cross-pollination is low or negligilale with the result that
ínbreeding or aptnnixis pre- vails. As mutr-irions occur over the course of time the isolated populations
evolve into differ- ent species. RS. ¡‘ishton countered Pedorov x-vith the opposite vieiv. He belietred cross-
pollínarioti ¡vrevails to give high gene flow between individ- uals. bach generation is genetically diverse as a
result of (Jutbreeding and is operatetl on lay selection, with resulting strong specialization for different niches
in the forest ectysysrenïïi-‘i

There are in tropical rain forests some genera x-vith ntany’ syrnpatric species and these could have arisen
from either mechanism. One of the best known, species-rich families is Diptero- carpaceeie, and in any one
west Malesian lowland rain forest up to twenty or so species mainly of Digiremcargms, Hopea, and Sistirea
commonly co-exist. Other lvialesiati examples are Polyaltlsia (Annonaïceae), Calophgrlhrm and Gzzrcínia
(Gtitríferae), and S}'Z}’g'ÍMWI (Nlyttzïcciïe). Further instances are Eschzueilera, Inga, and the forest floor
palms Geononta of the neorropics, and in XWest Africa Diospyros and Drypetes.

Veirionus studies on breeding syvstems in rain forest trees now cast light on the quesritins raised by fedorm’
and Ashton. An investigation of tliprerocari) breeding systems in h/Ialayam demonstrated two eipomictic
species, hire prob- able apomicts, sixteeti with slight or only tenta- tive evidence, and seiusn with negative
evidence for apomixis. ln addition rcn species were cross- pollinatcd and a further two prtibzalalyv so.
Apomixis is xsrell-ltntixsrn in culrivated rain forest fruit trees, namly in cirrus, clove (Sj-‘Zygirtfïí aro-
marictim}, mangosrrseu {Cïrrrcinfcï naeitgosraiza; Fig. 5.16), dtiltti, langsar lzls-(¡ÏTISÏILÜÍ donreszicimi],
and Inango (iï/Iazïgifera}, and probably {JCCUES in wild Garantia and Sjtrzygiren-z, both of which genera
have numerous syïnp-atric forest species.

By contrasr dioecism, the occurrence of male and female flowers on different plants, ensurcs that cross-
piallination must occur. lt is found ln many rain forest species, e.g. many Ebenaceae, Euphorbiaceae, and
Myristicaccae. Arieliaceae and Sapintlaceae svhcrise flowers look hermaph- rodite are in fact fufltïtlünélllfy’
dioecious. At La Selva, (LÏosta Rica, 23 per cent of 333 tree species were (liDECiÜILIS, and 26 per cent of
the 71'] tree species with stems 0.’! m in tllaïlïíilïfii‘ or over were diotrcious on an 8.8 ha study area in
Sarawak. But in the flora as a tirhole, when herbs and climbers are included, dioecy is less common: only 17
per cent of 507 species at La Selva and 9 per cent of “i320 species at Barro Colorado IslantlÉ-‘Ï

Apomicts are obligate inbrceders. Dioecitius species are tibligate ÜLIElJIÜEKÍlCFS. Hovirexrer, in a
species that is otttbreeditig, ttiost gene flow is in fact restricted to short Llistatlces. "This has been
denlonsttzlted in rain forest in a study‘ of isoen- zymes and of morpholtigiczil variatiian in the leaves of the
small tree Xterosperrinuiri inter- mediztnz, and isoenzg-‘mes of the emergen: tree Shorty: leproszrla at
Pasoh, h-‘Izilayta; ir was found rhar the further trees are apart the more they differd” lt appears rhar most
pollinzïritin occurs between near neighbotirs and most seeds fall near the parent tree.

The conclusion from all these studies isïhat rain forests, like other ecosystems, hats-e a diver-

The web of life — Clsrapter strmmarjr ' - 87

Fig. 5.16. Garcínía rsmngostana lmangosteen} of Malesia ha.s extremely uniform fruits; the species
reproduces apomictically. Malayan wayside fruit Stall.

il sity of brecding systems and therefore no single A one is the driving force of species BVOIHÍÍOH. All

kinds of breeding systems contribute to species richness, none offers an exclusive mcchanism. The patterns
of genetic varíation in the forest confirm, howevet, that there is restricted gene How in the few species
studied.

5.7. LNTERCONNECTIONS,BETWEEN
PLANTS AND ANIMALs-«THE WEB os

' Lite-CHAPTER SLIMIvÍAI-{Y

1. Numerous animals co-exist in tropical rain forests by subdividing its food resources. There has been loose
co-evolution between plants and animals.

2. Flowers have developed the same syn- dromes of features for particular groups of polliu nators in all parts
of the tropies, and we can distinguish bat, bird, beetle flowers, etc.

3. There are also syndromes of fruit charac— ters, likewise pantropical, each adaptivc to par- ticular
dispersers. i x '

4. Some plant species that have the same pol- linators or dispersers co-exist by flowering or fruiting at
different times.
S. Related plants have similar defensive- ' _ Chemicals and form plant webs, each one fed on

by a particular guild of insects evolved to detoX— ify those particular compounds. ‘Pest pressure’ is likely to
prevent members of any one plant web bcconiing particularly abundant. Scveral plant webs may share the
same pollinators or dispersers: these have been calied ‘mobile links’. At lean Seasons these depend for food
on plants that are fertile when littie else is, the so-called ‘keystone species’. Both mobile links and key- stone
species are essential for the ¡maintenance of full ecosystem diversity.

6. Closerco-evolution is less common. Ants and rain forest plants have evolved several kinds of mutualism.
New World Heliconiine butterflies and Passifloraceae have CO-BVCIiVCLl and the sirnpler adaptations in.
both partners are in the more primitive species. Figs and their pol- linating wasps show one—to-one species
relation- ships (Fig. 5.14). '

7. All kinds of plant breeding systems found in other biomes exist side-by-side in tropical rain forests. No
single hreeding system can be invoked as the mechanism that has generated high species richness. ' i 4 8. '
Rain forests are becoming fragmented, or changed by exploitation. It is likely that there is a loss of
complexity and diversity and the web

88 i)":trarcomzeïctírïzts bettyeert "plants aim’ airimals — the irreal? (iflffe

of intcrconncctitnns is unravclling. For example, AnTazíJnía11 fish need swamp forest Fruits; durians are
pollínated by hats that need Other pollen sources for the ll months Xvllfill durians are not in flower; guilds of
I-ïeliconiitie buttertlies and their Inimics become irnpover-

ished if their food-plant Passifloraccac begin to disappear because of loss of habitat. Because increasingly
more rain forests are being altered, biologists izrgetttlv need to monitor these changes with a view, if
necessary, to manipula- tiün in order to ctJnservc biodívcrsity.

6
Tropical rain forests through time

The great contrast between the two tlivisions of the zrchipclago is noi-vrhere so abruptly exhibited as on
¿passage from the island of Bali to that of Lombock, where the two regions are in closest proximitv. ln Bali
ave have ba rbets, fruit-thrushes, and woodpeckers; on ¿missing over to Lombock these are seen no more, but
we have an abundance of cockaroos, honey-stickers, ¿nd brush-turkcys which are equally unknown in Bali or
any island further WTSY. The strair is here fiftcen miles wide, so that we may pass in two hours from one
great division of the earth to another, differing as esscntially in their animal life as Europe does from
America.

A. R. ‘Xlallacc The Malay arcbipelago, {3rd ecln.), i373.

Patterns of distribution of plants, animals, and vegetation have long fascinated biologists. Their detection is
part of the search for general- izations about Nature, the reduction to simplic- ity of initially bemusing
variety. The scientist then goes on to seek for the causes of the pat- terns hc has detected. Over the past few
decades two major causes have been discovered for many of the present-day patterns seen amongst rain
forest plants and elnimals. These are conti- nental drift, and past fluctuations of climate; both have
contributed to a recent revolution in our understanding of tropical rain forest laio- geography.

6.1. PALAEOGEOGRAPI-ÏY

The occurrence of separare continental plates that have driftecl by ocean-floor spreading is an established
factd” Plate teetonics caused the supercontinent Pangaea to break up from about

180 million years ago (mid-Jurassic) onwards"

(Table 6.1, Fig. 6.1). The northern and isouth- ern halves, Laurasia and Güfldwaflalafld, con- tinued to
fragment and today’s arrangcmcnt of land and sea uvas not ¿achieved until about late Tcrtiary, about 10
million years ago.“° The break-up of Gondwanaland is of special significance for the tropical biogeographer
because all three tropical regions contain frag- ments of it. Thus, continental drift has had pow- erful
influences on patterns of distribution of families, genera, and species.

Gondwan an ranges

Sitnilaritics in flora between the three regions of tropical rain forest, described in general terms in Chapter 2,
OCCLIÏ because all are mainly parts of old Cïondwanaland. "The major evolution of the flowering plants
had occurred before Gondwanalazid began to break up and i135 con- tinued on the different fragments.
Today 334

90 Tropical ¡rain forests through time

Table 6.1 Geological periods and epochs mentioned in the text '

N111‘ T“; _ , i

l ylmlgn Permd cpoch Ï/Plelstocene 3 Q CÏ - Andes mountains Completed -'. m . ‘ J. l Pliourne h-iiocene —
collision at Sulasvesi 241’. _ i: I creares ¡nodern ¡‘vlalesia Oligocene 38 - 53 — Atlantic nearly formed
‘lïocene [l-"ig. 6.1L‘) 54.‘) - Palaeocene 65 ___. Upper í 97.5 ‘ 1.5 Lower — Gondwnnalz-intl and Laurasizi
144 1m" starting m break up F" . . b Late E ig 6 l i- .2 1.53 _- -——‘ M” Mid , i _, ISÜ — Pangaea starts to
break up

(Fig. 6,13]

Q = Quatertiziry i Last Il)‘ years oi Quaternary is the Holocene

genera and S9 families of fiowering plants are

essentially pantropiczll. Looking mote closely, other patterns can also be seen.

Campmïisjrernza has: a typical Citmdiyanam (lis- tríhution. This genus of light-demanding loivland
raimforest trees ÜCCUTS in Panama, Colombia, Brazil, Madagascar, Seychelles, and from India to
Micronesia. Ahsence from conti- nental Africa is believed to reflect massive extinctions there due to periods
of strongly dry climate.
Disjunct ranges, ivith one part of a plant- griïups (iecurritig on the Gtiyana Shield of north- ern South
America and the other in Nlalesiei, are also laelieved to reflect deseent from a pan- Gtmdwvanan ancestor,
with extinction in Africa. Examples ¿te Bonnetiaceae, a family of small trees (Bonnetfio America,
Ploiastiuriz Asia), and Tetrameristaceae, with lkntanrerista,‘recently- discovered in Guyana, as a relative of
the WCll-

known peat swamp forest genus ‘Tetramerfsra (punah) of western Malesia.

The geographical distribution of Diptero- eargyaceele dlsf} reflects a Gondwanan ancestory. SLIb-family
Dipterocatpoideae contains the important hig timber trees, pre-eminent in today’s international timber trade
and on vuhich so much of the discussion in this book is focused. They range from Malesia ivestwartls to
India and Sri Lanka, with one outlying endemic genus in the Seychelles and fossiis in Stamalia and Uganda.
Subfamily Mtmotoideae has three genera. Two are small ttees of stih-Sahatam ¿Xfx-iea, mainly of seasonal
forests. The third, diseoveted in the mid 19805, is a big rain forest tree ‘of Colombia in South Americad“
South America is also home to the third suhfamilir, Pakaramoideae, comprisiiïg one species, a shrub ot small
tree of the Guyana Shield rain forests, in Guyana and Venezueia, discovered in the 19705.

There are other groups besides Diptero- carpaccae which span the southern Indian Ocean, occutring on
ftagments of old Gondwana. The fan palms Be-rassas and its allies are one example {Figs (5.2, 6.3),
Nepentbes, the pitcher plants (Fig. 4.7}, found in B/Iadagzisceit‘ and Seychelles as Well as Malesia, ‘are
another.

Another common pattern is a geographícal range that spans the southern ¿‘ttlantíc Ocean, with represent-
atives in hoth tropical America and tropical Africa?” Twtelve families of floiveríng ¡Jlants are essentiallja
iimited to these two regions, and in many Cases are much mote numerous in one; for example, Bromeliaceae
and Cactaceae ‘which are mainly American hut have a Few Old ‘Cïiorld species. One hundred and ten genera
haire the same amphi-¿‘ltlantic range, eg. AIIIIOFIG, Chlorofihrhtar, Ocoteo, and the palms lïiaeis and
Ralybia {oil tweilm, raffia).

Closure of the eastcrn Tethiurs Oceanm

Gondwanaland and Lexur-asia were separated by the great Tethys Ocean. Tethys was closed by the
northwards movement of parts of Gondwanaland lFig. 6.1L First Africa and then

¡ai 18€ million gears (mid-Jurassic!

lI'ii

¡bnoo míilion years {lower Cretaceousl

¡‘im

Fig. 6.1. Continental drift. (a) IVÍid-Jurassic: Laurasia {north} and Gondivarlaland (south) in contact as
Pangaea, separated in the east by the Tethys Ocean. {b} Lower Cretaceous: Gondivanaland breaking up and
Laurasia starting to do so; Tethys Ocean being closed; AustraliafNew Guinea still jolned to Antarctica. (c)
Late Eocene: Atlantic Ocean nearly formed; Tcthys Ocean mugh smaller; Africa and India have nearly
collided with Laurasia; AustraliafNew Guinea has scparated firom Antarctica but the Malay archipelago has
not yet formed. (Smith and Briden 1977; present-day coastlines and continental shelvcs shown, Mercator
projection.)

92 Tropical rain forests through time

.. "° wsaïïmo ‘h’ 34..., 9°"WM»;;:Q s»?

"o, wm.» a» «q, N M ’°‘ Qggggg kvvwe::"“ñ».x M N“ M, A“ W v

Fig. 6.2. (Above) The fan palms of trihe Borasseae DTÏgÏTIQÏCCl on Gondwanaland. They continued to
evolve as ir broke up and occur today on its Old World ftagments. Outline, Bomssus and l-iyplzaene (10 spp.
each), both ividespread and probably with rangos extended by n1an. Bi, Bistnarkia (1 5p.);

Bo, Btirassotïendrorz (2 spp.); La Letonia (3 spp); Lo, Lodtiicea {1 5p., the double Coconut;

1M), Medemia {l 5p., last seen 1964, probably extinct.) (After Uhl and Dransfield 1987.)

Fig. 6.3. (Left) Borasszrs flabellffer, the lontar palm, here seen planted in east Malaya and exhibiting the
deep, oblong crown of youth.

Pczlaeoge ogro); by . 9 s:

india drifted north and collided with the south- A

ern‘ margin of Laurasia. Further east the continental plate which comprised Antarctica/ Australialstiuthern
New‘ Guinea moved north- wzirds, iaroke in two leaving Antarctica behind, and, as a simplificarion, collidetl
with the south- east extremity‘ of Laurasia, ar about 15 million years ago, the mitl-Miocetie; this created the
Nlalay archipelago (h/Ialesia) as it exists today. Both super-continents had their own set of plants and
animals.

India rafted Gondwanan plants northwards and after COllÍSÍOH these mixed ivith Laurasian ones to give the
dual-origin flora of that region today. The collision within ivialesia was at what is the modern island of
Sulawesi. Western and eastern Malesia have very different animals, dcmarcared hy a very sharp boundary,
‘Ü-{ïallaceïs Line (Figs. 6.4, 6.5). The strength of Wallactïs Line as a boundary varies with the ease with
which the animal group Linder consideration disperses over salt water. Present (lay zoogeo- graphy is a clear
reflection of the plate tectonic history. Plants do not divide so sharply at Ñïlallaceïïa Line, but groups of
ntirthivestern and southern lGtmdwanan) origin can he recognized rjFigs. 6.6, 6.7’), centred in i-vestern and
eastern Nlalesia, respectively. The northwestern group

- 1 3D‘ EGIJ’

- * - Wallactïs Line

---- Ïflflm line {edge of curuinenlal shelves]

has Laurasian plants, such as Magnoliaceae (Fig. 66h), ivell as others that come from the Indian part of
Gondivana, such as DÍPCEIÍJ- carpaceae (Fig. 66a), Ctenolophtnt, Dorit), iízrgeisstana and Gorzystylusyl“

Both Latlrasian and (Ïíondvyanan floristíc groups occur in the tropical rain forests of Queensland, northeast
Australia. Recent research has shown that Laurasian elements were well represented there before the mid-
Miocene collision and there was no sudden influx. This is because the evolution of the Mailay archipelago
was in fact more complex than a single collision.“5 Various shards pro- gressively luroke off Gtindivana
from the Jurassic onwards, drifted northwards, and became embeddetl in what is ntiw continental rïsia (Fig.
6.8). Then the easternmost limit of Lauralsia, southwest Sulawesi, broke off from Borneo at aboutr-IÜ
million years ago taking its Laurasiain flora with it, including Crerzoíopfron, Dario and Gonystylusfl“ This
flora is thought to have subseqtïently colonized land further east, including further fragments rafted off
(Íiondwana. More land progressively arose, both as island arcs and further fragments, and at about 15 million
years collision occurred, facili- tating further exchanges of plants and animals.

Fig. 6.4. \‘i¡allace’s Line, which runs bCÍWEZCII Borneo and Sulawesi, marks the boundary between the
Asian and Australasiarl faunas of Laurasian and Gondwanzïn ancestry, respectiirclyn (After Whitmore 1984a,
i-ig. 1.9.)

94 Tropical rain forests tfarozzgh ÍÍHïE

¿IÍACTERESTIC ÁEMEALE.

(b)

Fig. (.5 . ‘kïïallaceïn: Selection of animals to demonsttate the two distinct Faunas that occur in NIalesia.
These are {left to right and top to bottom) (a) Borneo: the ivestern tarsict Ïïzrsixrs rÏaancgnrrs, (me flying
lemur Clynoceplaahts variegnttts in flight and another one seated, the pentnil treeshrew PÉÍÏKJCCFHS
iozafi, the nlalay tapir Tapims indicas and a couple of lesser mousedeer ‘Tragnlns javamïcrrs; {b} New
Guinea: a tree kangaroo Deizdrolagzts inustis, the fairy lory Cbarmosyrra pcïpon, the twelvewvired bird of
paradise Seleucítfes nrelanoleuca, the common paradise kingfislïer Tarzjrsipiera gniatea, and a crovizned
pígeon Goirm crístata {COFOPJCIÍLI}. {Wallace 1876, Vol. l.)

Fíg. 6.6. (Right) Plant geography of Ivlalesia clerix-"ed from plate teetonics. {a} Dipternncarpaceae, ¿md

{b} Magnoliaceae are centre-d on the western, Laurasiaai, part of Malesia, They originan: from the Indian
patrt of Gonchvamaland and from Lautasia respectively. (c) Styphelia, Epacridaceae, has an eastern,
Gondwanan, centre of distribution. [Whitnmrc 1981, 8.3, 8.4, 8.9.) -

Palaeogeragrflplfly

An Anisoprera

[lo Corvfclobiunr

D Dípre roearpus
Dr Üryobalanops
H Hopea
N Neabalan ocarpus
Pa Parashoma
S Shama
St Steman oparus
U
V
Va

Alcitnandra
Efmenïlfa
Km eria i

Liriodendron lN. America also] Magnolia {America aisu} Mangiíeiía ' firfichelfa Paramích ella

Pa: chyia max

Tsaongiodendror:
U1

96 ‘Tr-optical’ rain forezsts terrestre}: time

Pinus ¡Julien

‘í F. ltesiya _ "'* P. rnerkusii

Fig. 6.7. In Dulalesia coniiers have either a northern, Lautasinn or a southern, GÜHLÏWIS-llliïll, centre of
LlÍSÏIFÍlÏJIIKlÜII (cf. Fig. 6.6].

la] Pinzis, {b} Pbylloclarïrrs, celery píne. (Kïihitniore 1981, fivs. 8.1, 8.6.)

The northern Andes

Another result of plate teetonics, which has left its mark on neotropical rain Forest hiogeogts- ph}; is the
Creation of the Andes which contin- ued till the end of the Pliocene, 2 million years ago. The Andes arose Ivy
the progressive uplift of the westttrn edge of South Pimerica as it over- rides the eclge of the Pacific
continental plate which is being DUSlIÜCl under, a process known as SLlbClUCÍlÜn. One eonsequettce has
been to

¿lis-ide in two the ranges of lox-irland rain forest species, So that today’ they are (lisjunct With one part in
the Pacific coast rain forests ¿Incl the other part in the Amazonian fiarests. In many groups there has been
evolution since the range became laroken so that the two parts are no longer identicsl. mi example, the
vegetable ivory palms, suhfamily Phytelephantrgicleeie, is shown in Fíg. 6.9.

Pelczeoclimntaes .97

c7 li
<1" , ‘7 _ '\ ‘lïmerá? TanImbarÉ-L/ï r__ __ ¡z-¡«f-Lfihï? ¡”K T l x J" h, x \\\ Australia \ l» K\ —x , ‘I.

Fig. 6.8 Plate tectoiïics and the eirolutimi of the Mala}; lfClllpfllagtl. From the late Jurassic (‘150 million
years BP} onvrards at the eastern end ot the Tethvs Ocean (cf. Fig. ójl) fragments hroke off northwest
Australia and drifted IIOIÍhWHFLlS. "This map shows the situa- :ion at the Early Cretaeetnus (100 million
years BP). After Metcalfe 1996, fig. 15h.}

The full lviogeograiphical implications of these

‘stepping stones‘, and earlier ones that ¿also Llrifted

northwaïrds, remains to he worked out.

6.2. l’AI.AliC)Cl.li\'1A'l'lÉ5

The climate of the tropics has i706!) continuallv changing. The ÜlCl idea of fixitv is quite wrong; eliinatic
changes have had profound íntluences on species rangos.

The Quaternary

Nltist. knowledge ahout past climates is for the last 2 million years, the Quezternary period, during which
there has been repeatetl alternar

"mars-n Ammandra Pelar: dra

Phytelephas
Ámman dra —'
Phytelephas

Eig. 6.9. The vegetahle ivory palnis, subfamily Phjttelephantoideaie, are confined to northivest South
America where their range is disjtltict. The Lipliit of the Andes, which continued to the end of the Plioeene 2
million years BT’, split the range. Evolution continued. ‘Today no species is found in the rain forests on hoth
sides of the mountains. {After Uhl and Dransfield 1987.] Nlany groups show this pattern, ‘evith one set of
species eentred on the Choco region of Colombia, the other in the firmaron, for example Cozzeggrirt,
Lictmia, and Partimzrí of the Chrysobalanaceziet

tion at high latitudes near the poles laetween Ice Ages or Glacial periods and lnterglaeials. During Glaeial
periods tropical climates “vere slightlv cooler ¿ind drier, with lex-ver and more seasonal rainfall. During
these times rain forests became less extensive and seasonal forests expanded. Most of the (Quaternatv uvas
like that; present-day elitnates are extreme and not typical of the period as a whole. "Today we live

A at the height of an interglacial.

The last Glacial ¡‘Hz-IXÍÏNHIÏ! (the ‘Oiliseonsin- Würm Glaeiatíon} was about 18 000 years ago

.98 Tropical rain forests tlartiargla time

(before present, BP). Since then the climate has got warmer, and during the last 10 000 years, the period
known as the Holocene, was warmest 5000-8000 years agoüAt the Glaeial maxima sea levels were lower by
as much as 180 rn because much water was frozcn in the greatly extended polar ice caps. In Malesia the
Sunda shelf, extending from the Asian continent, and the Sahul shelf, teaching out from New Guinea, were
both exposed (Fig. 6.4), so the separate islands west of Wallace’s Line became joined as one land mass and
those to the east as another. The islands of the Caribbean also became par- tially joined. Sea surface
temperature was cooler than today, by 5 °C or motel“ at 18 O00 BP in the tropics.

The evidence for past differences in climate in the tropics is two-fold, from the record of past vegetation
clecluced from palvnology, and from direct physical traces.

Prtlytiologyillg

There are montane peat deposits in former lakes in all three tropical regions. Pollen from nearby vegetation
gets incorporated in peat as it forms and can be extracted for study. The picture revealed by pollen analysis of
peat from a site at 2600 m near Bogota, Colombia, in the northern Andes (Fig. 6.10] is typical. The repeated
oscil- lations in the relative abundance of the pollen of the three different species groups shown on Fig. 6.1.0
are a reflection of fluctuating climate. In cold epochs vegetation belts were depressed so that the site lay in
paramo, high mountain inoorland, in which grasses and the high moun- tain shrubs Acaena and Polylepis
occur. By con- trast, in lnterglacials the tree line rose above the site and pollen of trees and shrubs predomi-
nated. Similar oscillations have been found in African and Malesian montane peat deposits, though none
stretch so far back in time. Radiocarbon dating has shown that the oscilla- tions were simultaneous at
different places and also coincide with the altcrnating Glacials and Intergleïcials of temperate latitudes. The
oscilla- tions of the tree line in all three tropícalregions, rcconstructed from numerous pollen profiles

[:1 Trees and shrubs

Herbs (Gramineael

' - ácaenalFafylspís

‘t’

0') Jh-
I

t]; E! l UÉper Guate

m ¡D | _L

Depth lmetresl
K3 a‘
C3 G

l I__L I Middle Quaternarv

IIl___|,
14o
P:
É
16D E
2
CJ
180 5
E.

—l 2m

Fig. 6.10‘. Pollen diagram of a 200 tn core of lake sediments and peat, extending for the whole Quaternary‘.
Northern South America, at Etigota, Ctxlombia, 2600 m altitude. (Data of ‘¿an der llammen in Flenley 1979,
fig. 4.12.} The repeated lluctuations in ahundance of the three species groups is a reflection of continual
clitnatic change.

that have now been zinalvsed, are shown in Fig. 6.2| 1. The reduction in temperature during Glacial periods
tivas greater at high clevations than in the loxrvlands and conseqttently vegeta- tion zones were compressed
and tlepressecl (Fig. 6.12).

There are fewer records of changes in vegeta- tion from the losvlands because suitable peat is uncommtm
and, although pollen in mineral soil can he Lised instead, few suitable pollen dcposits have been found. Two
profiles from the low- lands of northern South America are reproduced in Fig. 6.13. These show repcated
oscíllatitins between dominance by savanna herbs (including grasses} and by trees, which reflect drier and

Allllurlu ¡{menu-ul

fkillluiiti [rriulruu]

Allitudfi: imanes]
4500

¡iüüüi-

350o É-

ia]

¡El Super páramo

iïi Páramo

mm: Suhpáramo 2 Andean forest

É Suhandean

bou ndary v W Forest limit

fi; Andoanfsubandean forest boundaw

_ Recent forest cfearance eminem‘

Páram oISupcr páramo

forest

Palaeocfimates

Lagunillas
Los Bohos
Palacio
Bogota

‘ Fuquene

u‘ Pedro Palo

450o ,-
4000 -
3500 -

ibl

Age [years HP]

I‘::I Afroalpine bell

¡:1 EricaceousfAfrnalpine boundaw 1mm Ericaceous baát

mw msm Forest limit

2 lflontane ínrest bell

Recent forest clearance annïted

¡nIIonInhm-mnmgnnlnumllnumlfllll

4500 «-
4000
3500
300D
lcl

í Tundta

Age [years BP]

iii Tropicalpine grassland mm Suhalpíne grassland 1 llflontane and Subalpine forest

.\\\\\ Forest limit

Recent forest cfcarance omified

GrassiandfTundra houndary

Lower mountain forest

20 DUB
Age lyears EP}

{lnferred age ar Sífunkí)

‘IÜÜÜÜ

DEE

\\\\\\\\‘

. Kitandara

Bujuku
Rutundu
Mahoma
Kaisungar

[Cherangani] Kilimanjaro

Sacred
Muchoya
Brass

lmbuka

Komanimambun.
Iriim _
‘ Sirunki

.99

Fíg. 611. Oscillarions in the upper limit of forests during

the late Quaarernarp’ in

(a) the Colombian Andes,

{bl the mountains of east Africa,

and (c) the New Guinea _ highlands. (After Flenley 1.979, figs. 4.26, 3.2.4, 5.23.]

/ÜO

600D

la) PHESENT DAY

EÜÜD

b: C) Q D

3-300

Trotaícal rain fiarests thi-onggh tam»

[bl 14 ÜÜÜ-ZÜ 000?." e Xerophyti:

‘regotation

/\\Pcrennial snow

Super Páramo
2ÜÜD -
Attitude [metres

‘Páramo 7 Sub- Páramo

Savanna

and en;

‘HÏIÜÜ '

tropical
Ü.

Fig. 6.12. Vegetation zones on the Andes near Bogota, ñÏolornláia. Compra red to la] the present da)’, the
zones were (b) hoth depressed and CÜIIIPICSSCLl at the last Glacial maximum. (¿Uter- van der l-Ïammen
in Flcnlcy‘

1979, fig. 4.27.)

3630i

ai lo] Ï

wetter conditions, respectively. Racliocarbtun (lares show rhar the ciscillatititis correspond to Cilacial and
Inrerglacíal pcriocls. In central Ivlalayva a micLPleistocene peat ClCPOSÍÍ helow what is now the nun-vay
of Suhang airport,

Fig. 6.13. Pollen diagrams front lovulatid northern

South Atticricza (a) easterti Coltimhia and {hi Guyana.
Radiocarlwon dates ¿re ShÜVüTl in years BP. (After
Hatter in Vïhítmore and Prance "i981 Hg. '13.)
The repeated ÍlUCIUEItÍÜIIS in ahundnnce of the

three species groups shows an alrernation between dricr and wetter periods, with herhs and trees dominant,
respeetiiuelir. The recent sharp ciccline in trees Inay rellect the influence of latiming by‘ hunmns.

Kuala Lumpur, “ras clominelted by‘ Pinus and

gratas pollen. Pine Sehtanna is Confined to strongly’ seasonal climates and is not found in Nit-lliure today
(Fig. 6.721}. At a site ¡r1 lowland ¡‘tmaztinia pollen in rixrerine alluxfial deposits shows‘; tepeatetl episodes
of ahundant grasa at 4000, Z100, and 700 BP. This, hovtrex-‘er, proh- ahly reileets local itariatitms in “alter
level, hence in vegetation, and only indireetly‘ indíceïtes climatie tluctuatitan.

ln the Irïivuland rain forests of Queensland, pollen ¡Jrofiles have been described From peat deposited in
‘sen-‘eral voleanic crater lakes. Retaeated oseiliatitm between dominance by rain forest and by ‘tarot
scletophyrll Forest with tnueh Ettcrrlyptrss (Fig. 2.6) has occurred. The present- day laoundzary between
these TWO forest forma- tions is only a few kilolnetres from the sites. Althtiugh there have undouhtetll}!
heen changes in vegetation these might he only slight and ma)‘

Paíaeoelimates

zot reflect more than small fluctuations in :Éimate.

Éeoscieiztific eixidence

Previous seasonal climates in places that are at zhe present day perhurnid can also he detected r-y physical
signs. Soils containing true laterite, which only tlevelop in seasonal climates with ínctuating soil water
conditions, have been Ïound below the South China Sea between Malaya and Borneo, part of the Snnda
contin- ental shelf. This sea-bed is also dissccted by deep ÏÏXÏCI‘ vallevs. On the Sahul shelF lfllnlflll’
modules, which similar-ly reflect a seasonal climate, have. been found. The implication is that at Glacial
maxima, when sea-level was lOVVCT and these mntinental shelves were exposed, the climate aras ¡ndecd
seasonal, as the pollen record indi- zates. Off the coast of South America sea-bed ¿Ïeposits also indicate past
seasonal climate.

ln northern South America sand dunes have been detected in the llanos‘ savannas of ‘Colombia, a region that
today has a humid .‘limatc. The southern part of the Congo rain forest of ¿Xfrica grows over sands that have
¿Íñnity with the Kalahari desert to the south. In both these regions the sand is thought to have blown on the
wintl and deposited in places that avere open and very dry.

Land-forms provide further clues of past climate. Braided streams choked with debris

101

develop in climates. in which rivers Hood during periodical wet set-isons. Steep slopes form, sur- rounded by
gentle plains with grave] heds, as periodic heavy rains erocle the hills. In Amazonia, aerial survey has
identified areas under rain forest today with these geomorpho- logical signs that the}! developed in a seasonal
climate. By contrast, in perhtiniid climate-s that do not have marked seasonality, rivets develop meanders,
and hills are ronnded and with tlecper soils that are formed by chemical ‘weath- ering and less tnechanicaí
erosi(aii.“9

Today the. very highest equatorial mountains of New Guinea, East Africa, and the Andes have a permanent
snow and ice czip, above c. 4500 m. During Glacial maxima the SHOWF-llflc was depressed by as much as
"i500 rn (Fig. 6JÍ2). Signs of glaciation can be seen today below the present snow-line and on some
Inountains that at present have no snow cap; for example Mt. Kinahalti, 4101 rn, in ¡iorthern Borneo was
glaciated down to c. 3000 m (Fig. 6.14).

The Tertiary

Less is known about Tertíary climates, i.e. before about 2 million BP, but they too fluctu- ated. Cores from
lvcltnv the South China Sea between hríalayra and Borneo, which are used by oil companies for
stratigraphical analysis, include a band which contains pollen of the

Fig. 6.14 The ice-smoothed

rocky Summit of Mt. Kinahaln, Sabah, 4101 rn, which was covered by glaciers during the Quatcrnary ice
agcs.

1 U2
Tropical rain forests Íhrtmgb tinte

Fig. 6.15. The aroid genus Spzitfsipbyfltttzn occurs on hoth sides of the tropical Pacific Ocean in

lowla nd rain forest.

Eighty-eight other genera show the same amphi-Pacific disiunction, including Gireirtarda, Heficonicz,
Saurauia, Siocnrett, and Trigormfaalarzzzs. (Map 5, van Stecnis 1962}.

conifers Abies, Keteieeria, Pinus, and Tsuga, and of the xeromorphic shrnh lipfzrediïi, all of which are today
confined to the much cooler, driet climates. of continental Asia. This horizon spans 14 million years from the
Oligocene into the early Miocene (about 35 to 21 million BP). Rain forests were more CXÉEHSÍVE than at
any time in the Quaternary during- the early Plioccne, parts of the Miocene, and especially the early Eocene;
so these were all warm pericids. Then, in the late Tertiary, fluctuations similar to those of the Quaternaiy
occurred. The pollen record near Bogota discussed above was truly remarkable in extending back 3.5 million
years, during which period it recorded 27 com- plete clirnatic tiscillationsdfll

Tertiary climates. have left their mark on

present-day ranges of flowering plants in several ways. There are 89 genera of floweting plants represented
on both borders of the Pacific Ocean in hoth tropicatl America "and tropical Asia (Fig. 6.15)."5' These so-
called ‘amphi- Pacific tropical disiuncts’ can hardly be explained by continental drift. They are in fact
believed to have QClIÍEVSIIl their present range by migration north-viards around the Pacific rirn

via the Bering Straits; during warm periods of ¿z

the Tertiary the climatic gtadient polewards from the Equator was less steep than today. This is snhstantiated
hy Tertiaryr fossil flotas in Alaska which contain many subtropical and tropical plants, e.g. Aiangium,
Cinnrtirtomum, Firmíazza, ¡‘i/Iacaranga, and Saurauiti. . ¡‘Xfrica has a much poorer flora than the othe two
rain forest rcgionsÍ-ïf This is helieved to be because it was much more strongly desiccated during the
Tertiary. We noted in Chapter 2 Africzïs poverty in hamhoos, ferns, ancl palms. ¿Xnstralia too Suffered
strong Tertiary desicca- tion. At that time its lncsic vegetation became mainly confined to the eastern
seahüard. The strip of tropical rain fOTESÍS found today in north Queensland is only 2—3Ü‘l<n1 wide and
is of par- ticular interest because it contains the relicts of the old rnesic flora. This includes the ancestots

from which many modern Australian species adapted m hot dry climates are helieved to have

evolved, eg. Brachycfaíton, Flindeirsie, Gcziríenia, in some cases by explosiva speciation, e.g. Acacia. New
Caledonia is a shard of Gondwanalzïnd which drifted away eastwards from northeast Australia starting in the
Upper Cretaceous 82

Pleistocene refitgin

million BP. Because it is an island its vegetation has suffered less from the drier Glacieil climates so more of
the old flora has stiritived.

The lands bordering the western Pacific have the greatest concentration of prirnitive ‘rlowering plants found
anywhere, includ- ing rhnborellaceeie, Atlstrohailqiaceae, Caly- canthaceae, Degeneriaceae,
Eupomatiacexae, Himantandreiceae, Laïrdizahalaceae, Magnoli- aceae (Fig. 6.65}, Tetracentraceae, Trocho-
clendraceae, and Winteraceac. [t is most likely that they vsurviive here as relicts. Vülinteraceae, for example,
is a very ancient family with fossils back to the Barremian and was cosmopolitan in the mid-Cretaceous.
6.3. PLEISTLÜCENP. REFUGIA

The pollen record gives direct evidence for iluctuations in vegetatiran which reflect fluctu- ations in climate,
and there are also irarious dif- ferent clues in the physical environment from ivhich the same climatic
fluctuations can bc dednced. it has loeen discovered that rain ftirests have vvaxed and waned in extent during
the Quatertiary, and probably in the Tertiary too, and are not the eincient and irnmuta ble bastions where life
tiriginated which populist writ- ings still sometimes suggest. In the present interglacial they are as extensivo
as they have ever been, or nearly so. At Glacial maxima. itiwland rain forests are heliei-‘ed to have con-
tracted and only to have persisted in places VÍITCTC conditions remained favourable for them, as parches
surrounded by tropical seasonal forests, like islands set in a sea. ln subsequent hiterglzicials, perhumitl
conditions returned, the rain forests expanded out of these parches, which, have come to be called Pleistocene
refugia.

In the late 1.960s it was shown that Xvithin Amazonia birds have areas of high species endemisnl and
richness ivhich are surrounded by relatively poorer areas. The same was soon demonstrated for lizards.”3
Subsequentlyi many groups of zinimals have been shovx-‘n to exhibit

103

such patchiness, notably butterflies, and so have flowering plants, though for them only a few families were
included in the analysis. The centres of concentration more or less coincide with each other (Fig. 6.1621}.
The existence of these overlapping centres of species richness and endemisni is not seriously disputetl,
though for flowering plants some (eg. the centre around iVlanaus city} have been shown to he merely
reflections of high collecting intensityd“ These loci Ovfirliïp with areas that geoscieiïtific evi- dence
‘stlggests retained rain forest during Pleistocene glaciatiems (Fig. tílñb}. The original interpretation was that
rain forests have repent- edly fragmented to become islands in a sea of savanna and then expanded again, and
that the overlapping centres of high species richness and entlemism are the places that continuouslyr
remainetl rain forestfl” This interpretation has been criticizetl. Firstly’, it is more likely that rain forest uvas
replaced by seasonal forest than by savanna. Secondly, it has been questioned whether the geoseientific
evidence really does refleet past seasonal climates. The alternative postulation to counter the second
CTÍÍÍCÍSII] is that patchiness in species‘ tangos and richness is a consequence of diverse past fluctuatitnns in
the environment across the vast 35IÜÍJ-kn1 width of the Amazon basin. These fluctuations include basin-
wide lower temperature, rainfall and atmospheric carbon dioxide ctmcentration, and increased seasonality
during Glacial maxima, with in addition local forest disruption by mobile rivers {see section 7.8) and by
flotading, wind and firedsf‘ It is also now realized that species have evolved and migratetl within the Amazon
basin in an etivironment that has been fluctuating on a much longer time scale than the past two million years
of the Pleistocene.

In the African rain forests four groups of loci of species richness and endemism are now recognized by
students of mammals, birds, reptiles, ziniphibians, butterflies ¿ind flower- ing plants.“ These are centred on
Upper Guinea, on Cameroon and neighbouring Crabüïï and Congo, on the central Congo basin, and fourthly
on its eastern rim (Fig. 6.17). These are

104 Trrqpícaz’ min forests tíamtïgïa time

1000 km

m Butterflíes

1ÜÜDkm

2 or30ver|apping

endemiccentres

[RW WM -:

Palaeoecologícal

forest refuges

Üverfap of endernísm

and refuges lfilefstoeene refztgícz

Upper Guinea
Central Congo
Cameroon-Gabon-Congo '

Fresentdavrainfcrest - Pleistocene refuges

Fig. 6.17. Present-day extent of tropical rain forest

in rïfrica. Areas of high species richness and endemism, which are postular-ed locations of Pleistocene forest
refugia, also shown. (¿after Nïaley in Alexander et al. 1.996, fig. 5.)

the areas to which it is belieived rain forest retreated at the Glacial maxima. Tree species per hectare of over
0.1 rn in diameter pick up this pattern. The Cameroon refuge had ‘138 species on the richest of several O.64
—ha sample plots at-Korup and contrasted to less than 100 species per hectare for non-refugia] sites in the
African rain forests (Fig. 2.27).

The Pygmies who divellvsrithin the central

African rain forest fall into three groups Vïïlth

distinctive language and culture and these groups have genetic differences that suggest ÍÜ-ZÜ 000 years of
isolation. It is belíeved that as the forest fragmentcd into three parts, so did the Pygmy population; later to
spread out again nvith the Holocene climatic amelioration.153

Most parts of Malesia are today about as

equally rich in species, irlcludizig endemics, as the Pleistocene refugia of Africa and America. At the Glacial
maxima the Sunda and Sahul
105

continental shelves were exposed by falling sea- level. Rain forests were likely to have become confined to
the more mountainous places where there was more, orographic, rain. The main development of seasonal
forests in this region is likely to have heen on the newly exposed low- lands, and when sea-level rose again at
the next Irlterglacial these and the physical signs of sea- sonal climates (tlescribed above) were drowned. The
parts of Pvlalesia that are above sea-level today probably remained, largely perhumid and covered by rain
forest, which explains their extreme species richness and their lack of geo- scientific evidence of seasonal
past climates. Present-day lowland rain forest communities consist of plant and animal species that have
survived past clirnatic vicissitudes or have immi- grated since the climate ameliorated. Th us many species
co-exist today as a result of historical chance, not because they CO-EVOlVECÍ together. Their communities
are neither immutable nor finely tuncd. This point is of great importance to the ideas scientists have
expressed concerning plant-animal interactions which were discussed in Chapter 5, as well as to theoretical
discus- sions on the nature of plant communities. Those parts of the worldïs tropical rain forests that are most
rich in species are those that the evidence shows have been the most stable, where species have evolved and
cont- inued to accumulate vvith the passage of time without episodes of extinction caused by

_ unfavourable climatic periods. This is similar to

the pattern seen in other forest biomes, for example in the warm tcmperate forests of

Fig. 6.16. (Left) Pleistocene refugia in the New ‘sXforld rain forests. {a} Centres of endemism of butterflies,

plants and birds. (b) The overlap of these with palaeoecological forest refuges as deduced from
palaetwclimares,

topography’, geomorpholtugy, soils, and vegetation structure. (Whitmore and France 1987, figs. 7.4, 7.5.)
Note the cxtcnsivc refugia in the western Amazon near the Andes. This is the region of the world's richest
rain forests in terms of numbers of both animal and plant species (see Chapter 2). The locations of the two
hyper-rich forest plots at Cuyalaeno {Ecuador} C and Yanomonci (Peru) Y are shown (see fig. 2.27). There
are not yet sufficient sample plots spread across Amazonia to confirm that tree species numbers on small
plots rcflect the pattern of refugia detected from other evidence. The strong overla p between butterflies,
plants and birds shown in {a}, as well as the ranges of other groups of animals, arc interpreted to reflect a
wide range of past environmental disturhances stretching hack through much of the Pleistocene and

possibly into the Tertiary.

106

China“ where there are species-rich parches with a concentration of relict conifers inelud- ing the dawn
rethvood (B/Ietaseqiroiai gbqoto- srroboídes); and in the contrast between the species-rich Tertiary-relict
Colchic forest at the eastern end of the Black Sea in Transcaucasian Russia with the rest of the European
temperate hroadleaïf forest.

6.4. SEASONÁL CJLÏNÏATE ELEMENTS

IN MALFSIA
Seasonal climate plants in Malesia today have disjunct distributions, Confined to continental Asia and
é\ustralia; their extents clepend on how strongly scasonal a clinlate they require (Fig. 6.18), and on the two
population areas being out of dispersa-il range of each (Jther. [n Glacial times when there was more. seasonal
forest, as described above, these ranges would have extended and become continuous or nearly so.

Elephant and gaur (Elephas Jnoxímus, Bos gfliïlïf/{Sl ttiday have races living in the rain forests of western
Malesizi though they are mainly eininials of the seasonal forests of conti- nental Asia. Elephant browse on
forest fringe and gap plants and gaur are commensral with shifting cultivators, inhahitiug early regrowth
forest on fallowed fields. lt seems likely that these animals adapted to live in rain forest when their preferred
seasonal vegetation tlisappearetl as the climate ameliorated and sea-level rosefló” The same may apply to the
Sumatran rhinoceros {Didermocerus sumarreasis).

6.5. HIGH MOUNTAIN PLANTS

A larger part of the land area of the humid tropics of Nlalesia is mountainous than is the

‘case in Pifrica or America. European botanists

living in Malesizi, and perbaps nostalgic for home, have given much attention to the herbs and shruhs ftnind
EIbOXWE the tree line. There are two floristic elements, one northern, epitomized by Prirnitfa and
Rarumcultts [primrosc and lïiut-

Tropical rar"): forests ¿larottgf? time

Fig. 6.18. Geographícal rangcs of two scasonal climate species in ivlalcsia. {a} The two big cores of ever-wet
rain forest as barricrs for distribution of seasonal plants; thc hatchcd ¿areas either dry or consisting of a
mosaic of ever-wet and dry areas.

(h) ¡Pycnospom lzitescens, requiring some degree of dry season. {c} Rbyrtcrfirasïa tnirtintrt, recitliring a
Inuch stronger dry season; the disjunction is much more marked. After van NlfltLlvr-‘EH, Nooteboom and
van Steenis 1.96] in Vïhitniote l984rl, Hg. 14.4.)

tercugn}, the (rther southern, eg. Gonnet-a and Nerteraïf“ At the present deiy, populations on different
niountains are outside dispersa] range of each other and of source areas in the

Tropical rain ‘forests through time —«= Chapter smnnzary

tán.

Fig. 6.19. Coprosnztï ¡socket! at 4000 in on Mt. Kinabailu, Sabah. Cuojzroshscz is a genus of south
temperate latitudes which extends into the tropics on high itiouaitailis.

Himalaya and Australasia. This no longer pre- sents the enigma it fornierly did, now it is real- ized that all
montane xregetation helts haivc been stronglyi and repeatedly’ dcpressed during Glacial maxima (cf. Fig.
(5.12), with the consequence that populations would have been much closer. Ïvít. Kinabalu, the only really
high mountain in Borneo, however, seems ahvays to have been very isolated?” Nevertheless, it possesses a
rich IEITIPCFSJZC flora (Fig. 6.19}.

Parches of paramo nioorland in the high

Andes which are today widely sepatated islands ‘NÜlIlCi likewise have been closer together, if not
continuous, at times in the past.

6.6. CONSERVATION

The central objective of rain forest Conservation is to preserve adequate samples of the total diversity of
species and ecosystems. Interactions between species and fluctuaring clitnates on short, medium, and long
time-scales are one motor of continuing evolution. The discovery that some rain forests have areas of high
endemism provides criteria for the design of Conservation areas. The hiologist can state with Conviction that
it is necessary to include parts of these centres of endcrnism and of their boundn

107

aries, and also of surrounding species-poor forest, in order to conserve samples of the full diversity‘ of the
forest.

6.7. TROPICAL RAIN FORESTS

TITROUGH TINÍE—_CHAPTEIK SUNIA/LÁRÏ’

1. In his search for generalizations the natural- ist looks for repeatíng patterns of distrihutitnn and then for
their interpretation. Two causes explain many patterns of tropical rain Forest hiogeography.

2. Coritinents have dtifted. Sotne plants have ranges that arise from their occurrence on Gondwanalantl and
its fragnients as these have moved apart (Fig. 6.1). Biuogctigraphy of the Eastern tropics (Figs. 65-67]
reflects the con- vergence, then collisitm, of east Gondsvanatlaiui with Laurasia (Figs. 6.1, 6.8}. Northern
South ¿Xmerican rain forest higeographv reflects the Liplift of the Andes (Fig. 6.9).

3. Climatic history explains many other pat- terns. On the continental scale it explains some tlisjunct ranges,
for example genera that occur in the lourland hurnid tropics on both sides of the Pacific (Fig. 6.15). Tropical
cliniates have oscillared throughout the Quaternary, and the later Tertiary, between warm, wet periods, such
as today, and cooler, drier, more seasonal periods. These correspond, respectiveiy, to Interglzicials and
Glacials of high latitudes. Rain forests today are at or near their maxima! Quatetnziry extent. During Glacials
their area was reduced and tropical seasonal forests became more extensive.

4. Evidence fm fluctuaring climates comes from two sources. Firstlg’, pollen analysis shows vegetarion zones
have fluctuared. Most data are from mountains and show ‘vegetation types hzïve moved up and down as
climate has changed (Fig. 6.11). lu parts of the lowlands, rain forest and more Seasonal vegetation have
alternatecl (Fig. 6.13}. Secondly, direct physical evidence from geomorïnhology and soils, shraws some rain
forests occur toda)’ in places that once had sea- sonal climates.

T08
5. ln the neotropics many zmimals and some ¡Jlants have arez-is of high species richness and enclemisau.
These more or less coincide for dit» Ferent groups, and are beiiexïed to be refugia to WhlCh rain fiirest
retreated at Glacial maxima, Geoscieïitific evidence for ref-Ligia is also mote or less coincident (Fig. 6.16}.
In Africa there is evi- dence ot four such refugia (Fig. 6.17).

Tro-¡Jica! ¡‘aii-z forests tlyrougi’: ÍÍWIB

6. Disjunct distributions of seasonal climate plants in ivïalesie! (Fig. 6.13) and widely selva- rated
populations of high Inountain plants would have been ctmtinuous or more nearlv so in. Cilacial tueriods.

7. These patterns of distribution provide a ratíonal basis ‘for the design of conservation areas.

7
Forest dynamics

ln Chapter 6, tropical rain forests avere exam- A , «lc-c mi) med on the long-time scale ofïens of thousands
of’ years and ,sl_1oyvn to have fluctuated in area. " " ‘.'-u'-'1' v“"-‘»f4 ‘ñ hace, . . Dshitngfurtlig sti over
millions of years, con- tinental and evolution sacïíptnppfinigenptlkgfir, Present day fauna and flora _t;a_r
yigtness to these long-term changes. On these very long time scales rain _forests have been unstable and
have changed profoundly in extent and co1npo- sition, as indeed have all other hiomes.

In the present chapter we look at temporal

change at the other extreme, time scales

’ el Ü, ‘ . qhfiuea‘ ts that the forest growth cycle and its

ffia few,‘

. x ,‘-¿¿.7J({o/ï ¡Lt ‘i’ ‘i?

gt t e beyvildering diversity! of Nature, and it

ramificaritms is one SÉ sgt of generalizarions. This is a topic on which understanding has advanced rapidly
and a good figpïiework has been built. It has now rgitchízd a mature stage nvhere the ¡{fiieulgitgdinous
continuing research is essentially in details, ‘flgglgjggiggitit the picture’.

The chapter then goes on to discuss silvicul-

ture: the ways that hurnans make use of forest dynamic processes to manipulate forest composi-

. . 5- s win ¡«rk . . . years to a few centuries, up to the life plahs of tion for their own benefit. It conciudes
with a

one or a few generations of trees. Change is examined in the context of development and disintegration of the
forest Canopy, the forest growth cycle, introduced in section 2.3. Two spatial scales of Canopy dynamics can
be distin- guished: patch disturbancenwhïich lflVÜlVESJJqE

Í‘ ÍÏ 4912.21 l.

or a few trees, and cprnrnunfiyéxïiziïïiïifldilsiïr- bance. Patch disturhance is sometimes called ‘forest gap-
phase dynamics’ and since about the n1id—197Üs has been one of the main interests of forest scientists in
many parts of the worldÜÍ-Ï

In the first part of this chapter the processes of

regeneration and maintenance in rainflfyrorejglizs gire,“ described. They have close parallels"‘elsew;herel '

There seems to lhpïa general model of forest

¿dynamics which holds in many different biomes, m; avatar.“ . . a . .

al lbeit w‘1th, loc. lvariants. hcology as a science IS

r1‘. ‘.1 t fr . . always Sflfifklfig HDI’ generahzarions f0 make SCHSE

\u

consideration of community-wide disturbances to rain forests, now realized to be very and including human
disturbance and recovery

l“ ‘7

7.1 roassr MICROCLIMATES

Species differ in the microclirnate in which they f

successfully regenerate. It is important therefore ¿ar-itv f"

to gain an tlnderstanding of the rnicroclimates

L“! e.“ ‘e "0

within a rain forest. These are mainly deter-

by size of cantipy gap.

which is similar to that in a large cleïïringfïs substantiallfi}; dciyfferent from that near the floor z I" (_ l’ .

below phase forest. Figure 7.1 shows

readings taken in the Canopy at various heights

L’ tf 4' - _‘ï1g¿¡

wi’ esÏpVrÉad,

The microclimate ahove the forest canop)’: 3056i

"HU

Wind amd ¡m s")

Air teimeraium ¡[TJ

Relativa humidily {la}

U’!
É

i:

CÜÏ cmutntiatiun ¡ppm!

La L
E8

ll

89x

r1 a...

2D {JU
Time th}

Fig. 7.1. The microclimate ahovc thc forest Canopy strongly differs from that at ground level as is shown
here by the daily march of wind speed, air temperature, relative humidity, and CO2 etmcentration at various
Canopy levels in lowland rain forest at Pasoh, hïlalaya, 21-22 Notïemher 1973. Night ti me shadcd. {After
Aoki et al. 1978, in ‘Jllhitlntire "I 984m Hg. 4.7.}

rirrdülk above the ground. Outside, wind speeds during the day are higher, as is efitetnmïrgtute, while

.3...

lose to fhe ground

within the forest, carbon dioxide content of the air remains high all the time but up in the canopy it drops
during theïday‘ ¿{pg-Ito uptake by photosyuthesis. '

The light climate within a forest is complex.

There are four components, skylight coming through canopy holes, direct sunlight, seen as sunflecks on the-
forest floor, light transmittttd

Forest (Ïj-‘HHIJIÍCS

¿‘E S- ‘Sn. n: co u: 93.-: C ‘sw g XV) s‘? 05 ><°= .._ 37: “E 04 “É o- . c.” «Ü o BE SÉ‘ n“;- 22a.: (LU

‘CFC?

v I _.._: 200 Still) 400 EKIJÜ 600 70D SCIJÜ 90D ‘Iülüüllüüwavelengtlï {nrr

Ultra- I Visible and Infra-red 1

violet PAR j ywavebam} Al T Wavelengths for j phytoehrome

85D 3'30

red far-red

Fig. 7.2. The light in the open (‘white lighf} and in deep ftarest Canopy shade compared. There is a change in
hoth quantity’ and quality. [Clurves based on fig. "I in .\-'lacl.cllan and Frankland 1985.)

through leaves, and light reflected-‘frtim leaves, trunks and other surface-s. The sky ustially’ has some clouds
and the first two components have aluproximeitely’ the same spectral composition. Light transmitted through,
or reflected from, leaves is greenish because the orange to red wavelengths have been ahsorhed and utilized
for phorosynthesis (Fig. 7.2). The waveband 400 to 700 nm (which is approximately the visible spectrum) is
utilized for photosynthesis and is known as photosytitheticallyj active radiation or PAR. "The forest floor
only rcccives up to c. 2 per cent of the PAR incident on the fin-est canopy’ (Fig. 7.2}, and 50-80 per cent of
this is contained in sunfleclts [Table 7.1). Plants living below closed forest probably rely on the sunfleck
component of the light climate for photo- synthesis, the other components are too small to be utilizable.
Compared to the open, most sunflecks have low energy (Fig. 7.3). Very bright sunflecks are rare and may
have more energy than shade-adapted leaves can utilize.

In addition to reduction in quantity of PAR within the forest cantipy, PAR also changes in quality svith a shift
in the ratio of red to far—red wavelengths (66Ü and 730 nm) (Fig. 7.2}. This is because the Canopy
preferentially absorbs the

Forest nzicroelíiuates
Table 7.1 Forest floor light conditions

Location A-ilean per cent lvíaiximum % latitude) transmission total PAR due

PAR and range to sunllecks

Queensland, 0.5 (OA-ll) 62
¿Australia

(Zíicl STS) Oahu, Haxvaaii {A2-1°30'N i

La Selma,
Costa Rica

(IÜEZÉN)

Singapore e.

n “zow;

2.4 (15-33) s0

1.U—2.0 55-77

2.0 c. 50

Llainly after Chaztlon and Fetchcr {i984} table "l. ‘(cry lirtlc photosyntheticallv active radiarion (PAR)
penctrzites the Canopy and trios: of what does is contained in sunflccks.

red Waveband. At, La Selx-‘a, Closta Rica, the change was from a IIIEClÍiIIÏI value of the redffar- red ratio
of about 1.2 in the open to 0.99 in sunllecks and 0.42 in forest Shade.

Canopy gaps have an aerial l'¡"¡ÍCl‘()ClÍ111fltt3 like that above the forest but the smaller the gap the íess
different it is from the forest interior (Table ÍZ, Fig. 7.4). In particular, the amount of phow zosynthetically
active radiarioti is reduced ("Table 7.1) and its distribution through the day alters. Both the quantity and
quality of light reaching the plant is knoivn to be of profound import- ance in the mechanisms of gap-phase
dynamics, ss will be shown later in this chapter. Due to the increase iirsolar radiation, the soil of gaps is more
prone to tlrjring our in rainless spells than soil below the cazii_)¡7y15‘l and the temperature of ïhc top few
millimetres of soil may become as hot as 35 °C compared to only c. 25 “C below the canopy’ {Table 7.2).
The manner in which different species are adaptfid to different dimen- sions of Inicroclimare, especially‘ the
light climate, are described below. The microclimare has turned out to be rnore complex than had been
expected. Gaps were at first regelrded as having a microclirnate vzirying with their size, tu be contrasred
ivith closed-forest microclimare.

111
8D - {a} Forest floor
60
40

l\) É

CD
3o-
3

- (bl 400 m2 canopv gap

RJ
Cl

D
Proporlíon (es: of Iomminute averages
U1
D

- {CJ 500D m7 Clearing

25

e10 i s i 010091000 PARclasstgrmolm”s l}

Fig. 7.3. Ten minute ¿average values of pl1()t()S}’ntl'1&tiC¿lll}’ active radiatizizi {PAR} at La Selva forest,
(Iosta Rica. inside the forest {a} most ivalues are small with a few big ones due to sunflecks. ln a Canopy
gap (h) xralues are larger, but less than in a large cleaning {c}. ¡[After Chazdon and Fetcher 1984, fig. 2.)

But this is a simplicariondfi" In microclirnaric terms ‘forests are not just Swiss Cheese’, gaps are neither
homogeneous holes nor are they sharply boundedd“ ‘Within a gap the microclimare is most extreme in the
centre amd changes out- wards to the physical gap edge and beyond (Figs. 7.5, 7.16). The larger the gap the
more extreme the microclimare of its centre. There is likely to be a larger arca of srnall-gap-centre
microclimare around the peripherv of large gaps than in the centres of small gaps.

‘Furthermore, there is much more irariability between small gaps than large (ines in microcli- mare. The
amount of direct sunlight controls the

112
Forest dynamics

Table 7.2 Microclimates of open, Canopy gap, and closed lowland rain forest compared. (Danum, Sabah)

Cllosed forest

« Open Big gap Small gap

Per cent sky visible“ 74 31 10 6

Aerial microclimare:
Temperature, maxJrnin. ("CF 30“f22.5 38121.5 34.52215 28.3721
Relativa: humidity, min. (‘Vold 50 52 67 85
Photosynthetically’ active radiarion 35.0 19.2 4.9 0.5
{mol m’¿day'"ll'i

Soil mean temperature ("(3):

"lO mm Inaxzlmin. .35.5i"25 3532.3 305/23- 25122.5 50 mm maxalmin. 29.5Í25.3 3l.5i"24.5 25.1394
245/235

[lnpublishetl data of N. Brown and D. Kennetly Measurements made in gap centres

The smaller the gtip the more elosely it rescmbles closed forest except that minimum air and soil
temperaturas are

similar everywhere "‘ Assessetl from a hemisphere photograph (Fig. 7.4}

l‘ Mean of. weekly‘ shadecl value for 2 rveeks every month for '12 months

L‘ Open maximum temperature kept low by river lzireezes

d Mean daily value, from measurements of 2‘\’VCCk pieriods for 1.2 months

° Daily total, recorded 2 weeks every Inonth for 12 months

Fig. 7.4. Hemisphere photographs of (a) big gap, (b) small gap and {c} closed nlature phase canopy, with 31,
10 and 6 per cent sky visible, respeetively. These are the sites in the Danum forest, Sabah, whose
microclimares are shown in Table 7.2. Photographs ta ken With a 180° fish-eye lens. Niprth at top. Sky
percentage computed with an image analyzer; the human eye is very deceptive. Note cloud in (a).

maximum temperature, relative humidity and PAR (Table 7.2}. Large gaps are cipen to direct sunlight for
most of the day, for most or all of the year. Small gaps are only open for the much

shorter periods when the sun is opposire them and the position of the gaps becomes important. XWhere (as at
Danum, Sabah) afternoons are cloudy, small gaps west of the Zenith let through

Pioneer a-nd clímax tree sgiecies

MAX
Attemperature PC}
1.13
PAR [mol riff!
HAWDif

Distance {m}

75 _ i 25
Distance lmJ

Fíg. 7.5. Mícroclimate along Ilortlï-Soulïl‘! transects through a big and tiny gap Danum, Sabah). Daily
values.

Mea su rernents made over a year. {BFOWTI "1 993, fig. 12.}

’Relative attnospheric water demand.

The large gap is that also shown on Table 7.2 and Fig. 7.4. Note that there is no sharp change in microcli-
rnate at the physical edge of the gap and that its effect extends over 5 m into the forest.

less direct sunlight ¡md have a less extreme mieroelimare than those to the east, nrhieh the sun shines through
during the morning.

Thus, gap size is a poor surrogate measure of

nlieroclimate, most markedly over short periods. These complications were revealed by experi- ments
designed to unravel the growth of seedlings of different tree species in different microclimares. This is an
example of how a science advances. Here we started with the interesting, vital and apparentlyr simple
question of the extent to which seedlings of different tree species have different responses to different sizes of
gap and diseovered that this in fact was based on a very naive view of the forest floor microelimate.

7.2 PIONEER AND cuts-tax TREE

SPECIESÏG?

it is quickly discovered, even by the novice, that tree species differ, in thqamounp of solar radi- ation required
for their rélghlïerfdflon. At one extreme are species only ever found to grow up

along roadsides, in old fields and in big Canopy gaps. At the other extreme are species that grow up in tiny
Canopy gaps, or, in the ultimate case of Lmdergrowth treelets, in no perceptible gap. Ecologists and foresters
etmtinue to engag in

¿y gg.

vigorous debate as to whether species along this spectrum of light climates. can he divided into clear,
separate groups, as any glance at the research literature soon shows. -

As described in section 2.3, some strong light-

demanders require full light for both. seed germ- ination and seedling estahlishmetlt. These are the pioneer
species, sgiagagfrom all others by these two features.i'úgi By eontrast, all other species have the capacity to
germinate and establish below Canopy shade. These may he called clímax species. They are able to perpetu-
ate in the same place, but are an extremely diverse group. Amongst clímax species are some that, although
they can germinate in deep Shade, have seedlings that die unless they get full light within about a year,
examples being some Entandrophragma (Fig. 7.10} of ‘West Africa, Swietenia macropbyllrt of tropical
America

.39 a al";
"l "l 4 Forest‘ djmaraiics

Table 7.3 The main characters of pioneer and clímax tree species in tropical rain forests

Pioneer Clin-tax

Clommon alternative Light-demande; (shade—) intolerant, Shade-bearer (shade-l tolerant, primary names
secondary '-—- i’; i _ Germination Only in Canopy gzïps open to the sky Usually below Canopy’ ivhieh
receive some full sunlight Scgdlings z Cannot survive below canopy in shade, (Ian survive below Canopy,
forming a l" " ‘"7 ‘W-Tb‘ l o‘) never found there ‘seedling bank’ «Wïcvr. 5a,; f'¡f?’2“- “ al , Seeds Usually
small, produced copíouslfv and Often large, not copious, often produced more or less continuotisly‘, and
from early annually’ or less frequently and only on in life trees that have (almost) reached full _ __ ___height
__ _ _ ___ Soil seed bank brian}: species i fica-tv species Dispersal By wind or animals, often for a By
diverse means, including grairity, considerable distance sometimes only a short distance Dormancy Capable
of dormancy (‘orthodox’) Often with no capacity for dormancy l v commonly abundanr in forest soil as a
(‘rceaïlcitrarlt’), seldom found in soil seed '__ ,7 ¡V! _._.¿'(_r_¡-,,hf_, “y 966d bank bank f._l"“ij-p-_-lfl.'.lnq '
A 4 Growth rate. Carbon fixatitm rate, unit leaf rate, and These raites lost-eri" l l l” l l l“ ' ' ' relative growth
rates high Compensation point High Low I-Ieight growth Fast (Fften slow Branehing Spnarsc, few orders
(Jften copious, often several orders Growth periodicity‘ Ïndeterminate llsyilüpticll, no testing Determinate
(‘proleptic’), with testing buds buds ‘Leaf Jlife" Short, one generation present, raiz. high Long, sometimes
several generations f, i}, turn-over rate present so slow turn-over rate Ï-Ïerbivory Leartes susceptible, soft,
little chemical Leaves sometimes less susceptible due to defencc mechanical ttmghness or toxic Chemicals
Wood Usually pale, low density‘, not siliceous Xlariable, pale to very dark, low to high density, sometimes
siliceous Ecological range Vílide Sometimes FIGITOVL’ Stand table Negative Positive Longevity Oftcn
short Sometimes very long

From Sxvaine and lïlïlhitmcire [1 988]; "Whitmore [1991]

"' Reich e! al. {1991}

These sets of characters separate two ecological species groups, but exceptions can he found. Clima): species
htive a wide range of characters such that, except for the first ‘m-‘o listed, some are ver)‘ similar to tiioneers.
Seetext for fuller details.

(p. 142}, and some Partesborecï and Shorea of sion (as described in section 2.3), in other Nlalesia. Apart
from the absence of the two key respects pitaneers are just the extreme end of a characters that distinguish
true pioneers, these continuum of species. HOWBVEI‘, for the pur- species have many of their other features
(Table poses of making sense of the bewildering diver- 7.3}. To the forester, for example, their rapid sity of
the rain ftirest, the recognition of pioneer groivth and pale, low density timlaer, shared and clímax ecological
species groups is useful.

with true pioneers, is important. Thus, although Besides their light requiremcnt for seedling the pioneer
versus clímax distinction has critical success, pioneer and clímax species have a consequences for forest
regenerarion and succes- whole group, or syndrome, of rather associated lïoneer and dianas: tree species
characters {Table.7.3), but these are just tenden- cies and exceptions are xyell-known; For example, Aleurires
is a strong light-demander but has huge seeds (c. 8 g); Milicia exceisa and M. regia are African pioneers with
dark, dense, durable Wood, iroko, a tcak substituir: (much used for garden furniture); and. Casuarina and
Éecrtriríega "arc likewise hCavy-timbcrcd Eastern pioncers. i i

Pioneer tree species

Pioneer species germinnte and cstablísh in ajgap

after its creation (Figs; ,7.6-7.9). Threy grow fast

in height (Fig. 7.9} and laggards are suppressed

so the Canopy grows up with a strong tendency to he one-layered. Below the Canopy seedlings of

'¿ clímax species estahlish and, as the pioneer

‘seedling requirements for solar

115
Canopy breaks up after the ‘death of individual

trees, these clímax species are ‘released’ (to use foresters’ parlance) and grow up as a second grovyth cycle.
Succession has occurred as a group of clímax species replaces the group of pioneer species. Secondary
tropical rain forests have only a few trcc species per hectare, and sometimes only "one or tivo, by contrast to
primary forests which are usually ycry rich (Fig. 2.27).

Pioneer species are also called light-deman-

ders or (shade-} Íutolerants in reference to their i radiation.

Sometimes they are called secondary species because they form secondary or regroiyth forest on cleared
surfaces. Theterminology is confus- ing, and there is no general agreement. ' '

Amongst pioneers, differentspecies dattain dif-

ferent-heights at maturity, the larger-ones are

Fig. 71'.’ Cecrojiia obtusrt (left) and C. scsiadophgilía (right). Brazilian Amazonpwith j.N.1‘v1. Silx-‘a
(1985); = (Iecropia-ís the biggest geuus of pioneer trees in the ilcotropics, three or four species. mayoccur
together.

‘116

7.7. Macararzga gigantes. East Kalimantan. This Fig. 7.8. Macartmga tríioba. Nialaya.

east Borneo variant has perhaps the largest palmate leaves ofvany plant.
l J f f, longerdiyed, Table 7.4 lists common pioneer specie from the different parts of the tropics. Refere e is
often made to small or shortdived and to big -or long-lived pioneers, but it must be recognized that these are
only arbitrary subdivi- sions of contiñupus ivariation. A big gap is sometimes simuïtaiieously colonized hy
pioneers of different mature heights. After the short-lived pioneers die a taller Canopy develops of the larger,
long-lived ones, sometimes together with light-demanding clímax species (see below} that have become
establisheddág This is particularly noticeable in tropical America virhere. there is a big group of the latter
(eg. Cavaniflesia spp., Cedrefa odorata, Simaroziba amare, Swietenia

Forest dynamics
This gcnus ranges from West Africa to Polynesia.
Most of its c. 250 species are pioneers, which

makes it easily thc largest genus of picaneer trees in the xsrorld. Up to about ten species occur together.

macrophjyrlla, Vochysia spp.}. All these species

have similar tirnber (see below) and these tall sec- - ondary forests are of increasing commercial '

‘importance. Viewed from a roadside one sces two stages, early secondary {early seral) forests later

turn into late secondary (late serallvonesdfi) Each '

has a different set of species dominant and suc- cession seems to have occurred, but if one entera an early
seral forest and makes an enumeration it is found that all the species colonized early on.

Climax tree species

Clima): species usually germinate and establish

below a Canopy, therefore they can perpetuate

‘w ‘i,

Pioneer and clímax tree species

Fíg. 7.9. Ocbroma iagopiis, balsa. A Ïní-ycar-old

tree at La Selva, Costa Rica. Most famous tropical pioneer tree species, wídespreaid in the New World
tropics, renowncd for its very low density timber, where grown fast, which is much used for model-building
and was formerly used for aircraft.

iii situ. They are sometimes called primary species or, with reference to their seediíngs, {shade-l tolerants or
shade-bearers. This is the group of which clímax (primary) forest is com- posed, and clímax plant
communities are definedvas those that are self-perpetuating, in a state of dynamic equílíbrium. Seedlings on
the forest floor only grow slowlyi, and in most species seldom reach tailp than about 1 rn because they
eventuallyifi iii unless released. ’ Clímax species may be suhdívíded ín various ways, for example, by Lisual
mature height," but
onerof the most useful subdivisíons is ori the

117

degree of seedlíng shade toleranee, discussed

, ic ya?‘ esfera

above. The small trees that never reach the . surakxn .

Canopy probably only require entipg gpopyuig

space above\and below ground toiiiiidble them m w010“!

slowly to attain maturity. Strongly shado- t es typically grow slowly and have

dark, dense, hard and often siliceous timber which is often naturally durable. Light-demand- ing clímax
species (Figs. 7.10, 7.11) grow faster and most have paler, lighter, spfte timber . . , . . ‘ÍCÏ (“x-w which is
rarely siliceous and is se dom durable without chemical preservation. The range of di f- ferent sorts of clímax
species can often be found

in a single forest (Table 17.5}.

Character syndromes of pioneer and clímax species

It can be seen from Table 7.3 that in essence pioneer species are aggressíve. Typícally they produce a large
volume of low density wood by

mrúlwpïsfitgl íuïufiïvth, with opecib-bgpnched crowns

\ ere y they rapidly pÏe-erhpt competition by fillíng a large space. They start t_ reproduce - - Üigvv-¿‘xx .
early in lifeiand produce c pious seeds fre- quently, which are small and easily dispersed. By these means
pioneers e ficíeiitly exploit new - aiii f3“ big gaps as these develop cat ered through the forest. Growth is
opportunístic and indeterm- ínate, and continues so long as mineral nutrients and water are available. Leaves
are short-lived,

often prcgfihiigpd ¡cpntínually, and as their

LI men"

efficiency ¡soon tiirnmishïeis their nutrients are recycled tg fresh flashes, so there is no great need to
ilniihqsli in mechanical (tough, fibrous leaves) or chemical protection ¿‘gïïíiïsr herbi- vores. Píoneers are
liiiia É to regeiierate in situ, a mature stand has few small individuals, and the stand table is negative (Fig.
7.1221).
Clirnax species, by contrast, are slower groivíng and with denser timber and denser crowns. In order to
establish successfully below a canopy ‘NlICIE photosynthetically active radi- ation is low the scïeds contain
sufficient reserves for the seedlíng to build a root—system and the first photosynthetic organs. Seeds“ are
fewer and

larger, produced later and less often. There ís L———.t\-—-HJ

p-rxíovxc}; ' {INLLuxJ-v» (1 H

118
Stature
Small,
2—7.9 111

tall

Medium,
8-«29 m

tall

Large,

> 30 m tall

FrJrest dynamics
Table 7.4 Some common rain forest pinmgzer trcc species?
Neotropics
V.
Afrzcaï
Cordia nítida
Ocotea atirrerssis

some Piper Vemoniz: patens Vismia baccifera

Tremra
Aiciïornea Iripiínervia
Cecropia, c. 100 spp.
Cordia‘. spp.
Jacaranda copaia

fitïuntingia caiabura Ocbróma iagopus Sciaefflera {Didymopanax}

ñiororoïoni

Ceiba penïandïa
Cedreiitiga catenaefomxis
Gcqupia gIabr-a
Lactia ¿{Jrocera
Ficus capensis
Leer: gtiineehsis
1’h3-'iiaru:f91¿s

flaueiieriarzus

Raiwoifia vomitoria‘
Trama _
Anthocieista nobiiis
Psydrax amoidiana
Cieistcafnïmiis parens
M acararzga
MaesofJsis enzirrii
Mirsanga cecropioides
Spathodecr campzmuiata
Vemonia ¿Grifería
Vismia gitirteensis
Aucomnea ¡eiaineana
Ceiba pentandra
Lopbira aiaïa V
Miiicia exceisa
M ¿línia regia
Nauciea diderrichii
RicirzrJdendrtar-i

bczidelmïii Tenninalia iucorer-zsis ‘Terminalia supe-rba

Eastern tropics
Comnrersonia bartramia
Giociaidíór: spp.
Macczranga‘ >> 1.00 spp.

some Maiiotus spp. some ¡‘l/leiastonza spp.

Phyifazzthus spp.

PÍIJÍMTI-is spp.

Trichosgmrmua-rz, S spp.

Trama
Acacia aulamcarpa
Acacia‘ mangium
Adinanzira ¿famosa
Afpbiirlnic: ¡’mm
Azzthocepísalus, 2 spp.

few Macaranga spp.

Ploiariurrz aiternifiïiiirm

Ezicaiyptus ¿{agita-pta
Ociomeies
Sumarrana
Parras eriasztiaes
TAibizia) faiccztaria

All lzlle larga arid some of thc medium sizcd species ¡are importan: for timbcr l African species from NLD.
Swainc; American spccics from Fíncgan (1992) and G.5. Hartshorn ‘f For descriptions sec Havvthornc
(1995)

Fig. 7.1.0. Pole of Eiïïaïldrílfflkfflgflleï angolense, a clímax species whose seedlings gmw fast in high solar
radiation but soon die in deep Shade. Ghana.

Pioneer andciimax tree species

Fig. 7.11. Pole of Campnospemza attricziiarzmz, a clímax species with similar high light requirements to
Emaizdropbiragrrza (Fig. 7.10). Bilalaya

not the impcrativc need for efficient dispersal over long distanccs because rcgeneration can occur below a
Canopy. Rcgcnerarion in situ means juveniles and adults ttsually occur together. In the jargon of forcstcrs the
stand- tablc is positive (Fig. 7.1 2h).

Succession

The ¿shift from a secondary forest of pionecrs to a primary forest of clímax species is sometimes called
succcssion by ‘relay floristics’ by analogy

to a race.“ However, in some big gaps pioneer *

ancl clímax species grow up together, the former from seed thc latter either from seedlitigs which

119
Number of trees

h.) C2

ll] [I r-i amy-LTL Lïl-FH

0-3 n-a 1-5 21'25 nÏsínleÜiÏsírifizi-i
Birth [ml

Fig. 7.12. Stand tables of {a} the pioneer Endospermzirti msdzrilosum and {bj the clímax species Parinari
papziantt. Lowland rain forest, Kolombangara, Solomon Islands. {Wïhitmore 1974, fig. 7.2.} Pioneer species
cannot reproduce within the forest canopy so the population consists mostly of big

trees, whereas clímax species reproduce in sim and .

have juveniles and adults present together and hence what foresrers sometimes call a ‘reverse l’ or positive
stand-table. See Figs. 2.22, 2.23.

survivecl gap formation or from stem or root sticker shoots. In this case, which is likely where the forest floor
has not been completely dis- rupted, succession is by ‘simultancotis coloniza- tion’, tvith the pioneers
grosving fastest and initially dominant. Both modes of succession can be found in the same forest, dependent
on the severity of disturbance.

Clima): species arrive and establish seecllings ttnder the Canopy of a secondary forest. As the mature phase
cantapy of pioneers ages, individ- ual trccs or’ small groups die and create small gaps. ln these the clímax
seedlitigs are released and grow up as a second growth cycle below whose Canopy clímax species establish
again. As that mature phase canopy breaks up these seecllings are released as another growth cycle. Clima):
species as a group thus perpetuate them- selves irr rsiru, there is no directíonal change in species
composition. This is called cyclic regen- eration or replacement. o í

In a small gap, pre-existing clímax seedlings are released. In a large gap píoncers, which appear after gap
creation, Form thc next forest growth cycle. One of the puzzles which remains unsolved is what determines
gap-svtrítclï size.

120
Table 7.5
Forest dynamics
Regeneration behaviour of the common big tree species, Kolomhangara, Solomon
Islands
Conditions
Group Species to establish
Conditions

to grow up Timber

High forest
High forest
High fonrest
High forest
Climax Dilleltia snlomoflensís

¿‘idarmztbes corymbose Perínarí perpetuar Scbízomeria serrana

CÏalopÍJïlIarn peekelíi
C. neoefnzrrficttm High forest
High forestfsmall gaps
High forest Heavy hardwoods:
High forest high density, hard,
High forest dark, often with
High forest silica
High fcirestfsmall gaps
High forestfgaiïs

v, Nlerlium hardwoods: medium (lEnSÍty,

Pon-terio primera Highfclisttirhed forest High lowest/P small gap ‘I hairdness and colour Elaeocarpns
¿mgustifolizrs High forest Caps Light hardwoods: Cúíïípílflspeïififl lnreyipartiolnrtnn High forestlgaps Gaps
low detisitv, soft, pale T'erïiziíralia crtlorrtarrsarrai High forestfgaps Caps

Pioneer Endospernnirn mediellosanz: Mostly gaps Gaps

Gtareliixa nzolticcancr Mostly gaps Gaps
From Wïhitmore [1974]

The degree of Canopy {listurbance needed hy seedlings increases down the Table. Four ecological species
groups Can

be recognised.

Casual observations in western Malesia on

dipterocarp seedlings show that in big gaps clímax species become prone to attack by shoot borers or to
partial defoliation or to leaf galls. This suggests that the plants become stressed and unable to resist insect
attack, and that in these conditions pioneers replace them. A speci- rnen tree of the extremely shade-tolerant
di- pterocarp Neobalanocarpttts beimii {chengal}, a huge timher species, Was planted in 1955 to cel- ebrare
the creation of Templer Park near Kuala Lumpur, Malaya. In this open site repeated shoot attack by boring
insects has kept it the size of an apple tree, a sort of bonsai diptero- carp. Chengal has also been shown to
have its maximum rate of photosynthesis at intermedi- ate levels of PAR and also to have a very low light
compensatitm ¡joint (cf. section 7.5). In Kramer’s experiment, discussed in section 2.3, which was conducted
in loiver montane rain forest, gap-switeh size was at between 1000 and 2000 n13. ln the Itiwlands, where the
climate is hotter and less humid, pioneers are commonly observed in gaps above about 200 m2.

7.3 SEED AND SEEDLING Ecoroorl”

Pioneer species
Pioneer species germinate from seed in a big gap

after it forms. The seeds may be borne in after gap creation, or niïiy already be present in the soil. (LF.
Symington in 1933173 reported that at Kepong, Malaya, he had placed some ftirest soil in the open and
observed germinarion of pioneer species from it. The experiment tiras repeated by RQVÜÉ]. Keay in Nigeria
in 1960174 and many times again since the earlv 19705 in all parts of the tropicslï‘; ‘With the same results:
wherever a soil seed bank has been sought under lovvland rain forest one has been found. Ivloretiver, it is
always mosthï of pioneers.

Seed physiologists have discovered that

pioneer species have seeds which can withstand desiccation and then become dormant; longevity is extended
at low temperatures. They have called this hehaviour ‘(Jftl1(_)(.lOX1[?6 (cf. Table 7.3). It is this capacity for
dormancy “which enables seeds to accumulare in the soil.

Seed and seedhrt g ecology

I-Iowever, beyond the demonsttarion that seed banks are (so far) ubiquitous, little is yet known about their
veiriatitm in either space or time. These are still topics ripe for investigation. Pioneers have small, easily
tlispersed, copious

seed and the seed bank may tlevelop by seed i

rain, and be continually’ augmeïited. In this case a seed bank uniforni over large areas, the more so with
increasing seed longevity, should develop. Experimenta such as Sy1nington’s tell us nothing about how long
seeds have been in the soil. ln north Thailand the soil bank con—_ tained more seed than one year’s
production of nearby pioneers. Several Cecropia species in tropical America have seeds that live ITIÜFC
than one year in the soil.

The alternative possibility is that the soil seed bank is a record or archive of pioneer trees ivhich formerly
occupied the site. Despitc good aclaptatíons to dispersal many seeds are likely to fall below and near the
parent tree. In this case the spatial pattern will be a series of patches or ‘footprints’.

At Danum, Sabah, the seed bank below primary forest contained almost none of the usual expected pioneer
trees (E-ndospermum, Macarmzga, ll/Irzllotus, Tren/za), but the shrLibs and climbers of adiacent unstable
earth stream banks instead. This site had only recently become accessible by road and the nearest seed source
of the Sabah pioneer tree flora was several kilometres distant, out of dispersal range.

Triggers to germinate

The dormancy of the seeds of tropical pioneer species has been shown to be broken by one of two features of
gap microclimare. Many, so» called photoblastic, species are triggered to ger- minate by exposure to light in
which the photon flux density of the red (660 nm) wavelength exceeds the far-red i730 nm). This is mediated
by their possession of the pigment phytochtome. Photoblastic germinarion is easy to denionstrate by
experiment (see Fig. 7.13}; Red light is depleted below a forest canopy by absorption and reflection (Fig.
7.2) and far-red exceeds red.

121

‘IÜÜ
(I) 1D

G1

(J

“¡la Germinalien 8

Closed
Canopy
Canopy
GBP

Fig. 7.13. The small pioneer tree Utero caracasaira has photoblastic seeds, which getminate at high ratios of
red to far-red light, i.e. in gaps, but not below thc forest Canopy. Los Tuxtlas, Mexico. [Orozco-Segovia er al.
1987, fig. I; the bars indicate the standard errors.)

The seeds have evolved to detect the occurrence above them of a canopy gap with full white light, in which
red exceeds far-red. Moreover, short bursts of white light, as in passing sunflecks, do not trigger germination.

Other species, which include Ocbronza lagopus (balsa), have seeds triggered to germi- nare by either high
temperature?’ or by alter- nating high and low temperature. This so-called thermoplastic response enables
them to detect the change from closed Canopy to gap (Table 7.2). In parts of the neotropics, pure extensive
stands of balsa develop where burning has fol- lowed forest clearance. lt has been discovered that Ocbroma
seeds have an impermeable layer in the testawhich is ruptured by heat to allow the seeds to imbibe water and
to germinate.

Microsite for establishment

Soil seed banks characteristically contain many species. For example, small samples (2 m3 >< 50 mm deep)
from six different forests in Ghana contained from '17 to 38 species each, nearly all of them pioneers.
Equally characteristically, pioneer forests commonly comprise only one or a few tree species, a subset of the
seed bank, and

122
Forest dynamics

Table‘ 7.6 Different microsite preferencias for establishment of seven pioneer species in a cleared Venezuelan
forest site

5-1

ss 3 ‘a a u, s s E 3 E 8 ‘5 E u) JL.’ cn h LL. Ü t" E É P .5 a ‘¡_- ._, ___ o É k2 = m .57. __ É 3 "eg É c’ 53 E

LÏ g ._ -« o o o R u se a S B: T: 55-3‘ Í‘:- É ‘i’ ÏÉ E e e e. s e e s 3 s2 5 B ñ .3 h? E .3 g '-“ E U a EL c: l‘: O
"" -— U s: b ‘v- 3 É‘ ‘e É e x a a. a W 4 <3 s. -... e ‘i: 8 u e S. TE»: s» e 2 5 tu w- Q h: ,3 D f3 2 u, E o 2:- v;
(.2 ua m m Without dead litter of leaves, twigs and branchcs: Root mat 50 y 8.9 — — — w — — Charcoal
54 6.9 — — — — — vr — Bare soil” 44 14.8 — - — — — r a With litter: Root mat 39 4.3 — — — — — e
- Charcoal- - > e. 35 2.9 e s s a‘ te + -— Bare soill’ c. 35 6.7 e s + + e F —

From Uhl et al. (1981)

Established success: "" very good; + good; waeak or nÍl 9 XVater loss from a can of soil at field capacity in
grams during an 8—h (lay

l‘ Soil was a cream coloured sand

we do not yet fully understand why. Part of the answer probably lies in differential success in seedling
establishment, dependent on weather at the time of germination, or on mierosite. Table 7.6 shows results of
an experiment in Venezuela where six of the microsites commonly oecurring in felled and burned forest were
planted with seven of the local pioneer species. The microsites differedsubstantíally in soil surface
temperature and Water loss. Only one species, a forb (viz. broad-leaved herb), established on al] six. Only the
grass established on bare soil. All species except the grass established 011 the two microsites that were
coolest and most moist.

Climax species

Many rain forest clímax specie haïe seeds that germinare immedately ordevrltilin a few days. They cannot
¿nod desicearion or low tem- petatures. These seeds have been callecljreealci- ttant’173 because they cannot
beïsfdfédfand lfliilis pose a problem to plant breeders and nursery- men. Examples are many fruit species,
e.g. Citrus, Cocoa, mango, and durian (Citrus, Tbeobroma cacao, Mengífera, Derio zibetfafinus) and many
timber species, including all rain forest dipteroearps. Not all clímax species

have recalcitrant seeds, for example, many

Seed ¿md seedling ecology

Leguminosae have hard testas and ráilpfiaged dormancy. No pioneer species has yet been found uvhose seeds
are reealcitrantd?”

Many clímax species have bigger seeds than pioneers. By immediate germination these escape seed etaters
apd alsp degradaríon of their

n») ¿€.7-7ln‘.,5t.’\,k-°’! . . usually fiitty storlage tissues by micro-organisms. Seedlings develop, and clímax
species have a ‘seedling bank’ on the forest floor in contrast to

the soil seed bank of pioneers.

n/licrosire for esteialisbment

As with pioneers, different clímax species may be more successful on some parts of the forest floor. A close
study of the seedling ecology of some Malayangipterocarpaceae by P.F. Burgess explained lsibyieral
previously enigmaric aspects of the distribution of this important family?“ Dipterocarps are confined to the
lowlands and do not penetrare lower montane rain forest. The upper limit taras shown to be set by the
inability of the radicle of a germinating seed to penetrare peat, which develops on the surface above 1050 m
elevation. Seedling establishment is most successful on flat microsites; seedling stocking diminishes ‘with
increasing slope, rapidly at microsites steeper than 45° and falling to nil at 65° slope. This explains the
decrease in numbers of dipterocarps with elevation where the land becomes more rugged. The important
rimber species Sho-ree cn-rtisií (seraya, Fig. 2.3) was studied in detail. After the 1968 seed year seedlings
were recruited markedly better on granite-derived soils, which are coarse and sandy, rather than on the more
Clay-rich soils derived from shale, probably because the latter develop a hard surface skin under the impact
of heavy rain. However, after several years both sorts of soil had about the same stocking of seedlings; the
heavier mortality on granite- derived soils could be because their coarser structure makes them more prone to
drought. This study serves to reveal the great complexity of intertx-vined factors which control seedling

123

establishment and survival and that are impor- tant in determining the ecological range of dif- ferent species.
At Lambir, Sarawak, the dipterocarp Dryobalanops aromarica occurs mainly on ridges and upper slopes with
sandy soil whilst D. Ianceolara occurs on clay soils and in sandy-soil valleys. This pattern has been shown to
result from differences in seedling establishment. For, although both species germi- nated everywhere, D.
lanceolete suffered root predation on ridges and had higher establish- ment success in valleysѓ g

_-_ t, 10x‘.

30k), ¿»y ve" E_.».-\ c. L, ‘la. ¡a lt.‘ «s. =.

Seedlintg szrrviual

Many seeds fall near the parent tree and dense carpets of seedlings form. In several forests mor- tality has
been shown to be density dependen‘: and, therefore, is greatest near the parent, due to pathogens or
herbivoresm The fewer seeds which tlisperse to a greater distance are most likely to grow into seedlings that
survive. This so—called ‘escape hypothesism” has been invoked as a mechanism which preventa rain forest
trees forming single-species stands, although there are exceptions. Por the diptero— carps Shores leprosuia
and S. macroptera it was found“ that mortality depended more on microsite, and had no relationship to
density or to distance from the parent. In the case of S. CMTÉÍSÍÍ ants destroyed isolated seedlings, thus
accentuating the clumping created by seed falling mainly near the parentÏS-ï

By whatever means they arise and are main- tained, species patterns amongst seedlings on the forest floor are
a powerful, though little studied, determinant of floristic pattern in the following mature phase forest.

Dipterocarps bear fruit heavily only once every several years (p. 63}. Seedling populations are then
augmented and many attain a density of over one million per hectare. There is a sub- sequent decline, which
may reach zero, before the next gtegarious fruiting event (see Fig. 7.14).

124 Forest dynamics.

su v- {a} 12o — (si r T000 m \ a ' "s Egg l- NISEE recruits \ EL] —- 4IJBB E 1957 g y a 3 195i] recruits
grecruits ¿f i g '*- ‘a c; E I: Éan- En JÜDUG 2 2 i’ E I u J I I I I I I I n . i ___l,._1___.,_l_ ._¿_.__ _..J I I I a
1961 i952 1Hli3 i964 1955 1365 IBS? 19GB 1959 ‘(BTU
Year ul measurement
1951 1562 1963 1964 1365 i966 i967 1968 1969 19M

‘fear of measurement

Fig. 7.14. Fluctuation of seedling populations of the light hardwood diptetocarp Parasborea tomentella at
Sepilok, Sabah. (a) Fate of individual flushes; [b] total population. Numbers on eight plots of 4 m2. (Fox
1972

in Whitmore 1984a, fig. 7.7.)

Ïn Tvialaya, by contrast to Sabah, the populations of dipterocarp seedlings commonly drop to zero laetween
successixre flushes. This has important consequences for silviculture.

7.4 COMPONENTS or SPECÏFS

RICI-INESS
The dynamic nature of forest canopies provides

many different regenerarion niches to which dif- ferent species have become specialized. A major niche is the
light climate and there are species with all degrees of shade tolerance, as discusscd above, divisible into
pioneer and clímax ecologi- cal groups, which are themselves heterogeneous but less clearly divisible.

In all tropical rain forest flotas there are fewer

pioneer than clímax species, and they mostly belong to a few families; for trees these are Euphorbiaceae,
Malvaceae, Moraceeie, Sterculiaceae, Tiliaceae, Ulmaceae and Urticaceae. The Eastern tropics are richest in
pioneers, the biggest pioneer genus, Macarcmga (Euphorbiaceae,.Figs. 7.7, 7.8), is concentrated there though
with a few outliers in Africa. Most of its c. 250 species are pioneers and many are confined to only one or a
few islands of the Malay archipeleigo. Cecropía (Moraceae, Fig. 7.6} is the second largest pioneer tree
genus. lt has c. 100 species, most very ‘veidespread, and ís entirely neotropical. Musnnga cecro-{sitiides
(Moraceae, Fig. 7.15) of Africa has similar crown form _ to these two. It is monotypic. Trama {Ulmaceae},
with 10415 species, occurs

Fig. 7.15. Mitsanga cecropíoides [palmate leaves}

and Anthocfeista rtobífis {oblong leaves}, pioneers of West iïfrican rain forests. Ghana. '

Components of species richness

in all three rain forest regions. [Jioneer genera do not have neirrtiwly endemic species of very restricted
geographic distribution, and most of them have only a few species. These features are probably a reflection
of efficient dispersal which militares against species evolution iria the devel- opment of localized generically
distinct popula- tions. ivloreover, some pioneers, including the largest two genera Cecropia and iïflncararzga,
are dioecious, which has the same result. It is ¡iote- worthy that the region with the greatest number of
piüflccl’ species is the Eastern tropics. This is probably because partial isolation on the islands that comprise
this region has allowed more species evolution than has occurred on the con- tinuous landmzisses of the
African and American tropics.

There has been much specularion about the

extent to which the co-existence of numerous species in a given forest is due to speciailization to different
facets of the regenetation niche, thus eivoiding direct competition. A little evidence of specialization has been
garhered. ‘When a tree blows over its roots are upended and a holloiv is

125

formed plus a mound of mineral soil, sometimes called the root plate (Fig. 2.21). Ceci-opa: abrirse in Guyana
and Tirema tomenrosa in Penang have been. shownw’ to establish preferenrially on root plates. It has also
been shown at La Selva and at Los Tuxtlas, b/lexico, that (liffcteilt species successfully’ establish seedlings in
differ- ent parts of treefall gaps, the crown, trunk or butt regions”? Differential Seedling perform- ance in the
changing Inicroclilnate from gap centre (Jutwartls was shown by rooted cilttings of seven fvliconia and Pijier
species at La Selvall“ and by wild scedlings of three diptero- carps at Dannin (Fig. 7.16}.

Degrees of disturbance
The most species-rich forested landscape will be

one that includes both parches of secondary forest recovering from a big disturbance and consisting of
pioneers, and also parches of primary forest composed of clímax species. Wïhen succession is completed and
the whole laindscape is primary forest again there will be

14 r al 14

l - l 12v —; 12 I E _, Hopea nervesa , r 1g ___ Ig l Psrashereaniaïaanonan 7% ._ '. _ . .. ' . l g .Í:É

Shnreaiaharensis 7 3 g —— PAR 5 g s ‘ se V’ , I a A e 4 Í U ‘é — 2 .. ' «- Ü 0-5 m Ei-IÜ m 1Ü—15 m 15-20
m 20-25 m 30-35 4- Gap H Closet! forest

Distance from gap centre {m}

Fíg. 7.16. Differcntial seedling heightgrowth of three dipterocarps along a transect from gap centre to closed

forest. Dainum, Sabah. (Brown "1 996, fig. ‘l; error bars are 95 per cent confidence level.)

‘Nleans Iieight growth over 6 years was significantly larger than of same species in closed forest control
plots. Hopea grew faster in both the gap and its penumbra, ikzrasboiren and Siuoren in the ga p [though the
latter W115 badly damaged by an insect, see text). Mean daily PAR up to 10 m from the gap edge was
signifigantlyv’

greater than under closed forest.

126
no pioneers and landscape-scale floristic richness will he reduced. This is "the inrermediote dis- turbance
bgrpotbesis of LH. Connelljlg” The concept is bound to be ‘true at certain scales of dimension. But if we
were to look at tiny parches of the landscape we would probably find species-poor stands of pioneers and
also stands of high species richness where the more numerous clímax species occur in greater mixture and
which have, moreover, accumu- latcd dependent synusiae of epiphytes and climbers, so at this smaller scale
the clímax parches are richest. The catch-all hypothesis of Connell includes intermediate intensity and fre-
quency of disturbance, as ‘well as intermediate stage of recovery. It caused scientists to look at species
composition in a particular “ray and was a salutary correction to prevalent ideas tying diversity to maturity
and stability. But it is not so much an hypothesis as an inevitable property of vegetation at certain scales of
space and time.

7.5 GROWTH AND sesos TOLCERANCE

ANALYSED

The physiological basis of growth rates and of shade tolerance amongst tropical rain forest tree species is
now fairly well LÍHLÏBISÉDÜCLIQ“ This is the interface between ecology and physiology, so-called
ecophysiology. lt is important to conduct experiments in the forest, despite the great complexity of the natural
environment, as well as in the laboratory and shade-house where conditions can be specified and controlled
but are inevitably somewhat unnaturalÉ-gl

The basis for the fast growth of pioneer trees is their high unitleaf rate, E, the rate of dry sveight created by
photosyuthesis per unit area of leaf. Table 7.7 shows that of six species studied at La Selva, Costa Rica,
Ocbroma lagopus (balsa) had unit leaf rate not significantly lower than of the herb Helíanrbus annuus
(sunflower) grown as a control. Other species, subiectively ranked as progressively more shade-tolerant, had
progressively lower unit leaf rate. Relative growth rate (dry weight

FCJFBSÍ dynamics

increase per unit of dry weight} also decreased with increasing shade tolerance hut is a less basic measure
because it changes with age. Other research has shown that the rate of carbon dioxide uptalce is also higher
in light- demanding than in shade—tolerant species, despite high variability due to leaf age and past history
(Fig. 7.17}. This is due to higher appar- ent quantum efficiency and mesophyll conduc- tance (Table 7.8 a,b).
Pioneers also have higher dark respiration and light compensarion point than clímax species (Table 7.8 cal}.
Numerous experiments [rave shown that, overall, pioneers are more flexible in adapting to changing-condi-
tions: they arc better able to adjust architecture, morphology, leaf auatomy or various ecophysio- logical
attrihutes. ‘

Within the forest, most of the photosyntheti- cally active radiation (PAR) is in the sunflecks (section 7.1).
Researchl” on the herb Alocasía mocrorrbizos and on seedlings of the tree Toona ausrrafís, both shade-
tolerant species from the Australian rain forests, showed that photosyn- thesis took 25-40 minutes to reach
Steady state in bright light after a prolonged, low-light period. That is to say, there is an induction period. For
Alocasia, loss of induction when the

30
T: _ 4..-»: Oehromafagopus A
N 25 ,K'
I x‘?
E

g 2D I”; 3' .2 2 15 X . ,3 ¿x fr, Hampea appendtcufata I f. i‘ _____¿-,.__a Bursera simarouba B 8 lo " "
Dfpreiïxpanamensfs l E; _. Gnerum ieyboldr" z viroía koschnyí C

Pentacleflira macmfoba .

40o són ' 15m3

12W
Photon flux density Lumol m’? {ll

Fig. 7.17. Photosynthesis rate of seven tree species at La Selva, Costa Rica. A, pioneer; B, gap-dependent; C,
shade-tolerant Canopy species. (After Oberbauer and Strain 1984, fig. 1.} Rate of photosynthesis (Ineasured
as nct CO2 uptake) increases to an asymptote with increasing light. The maximum decreases with increasing
species shade tolerance.

Growth and shade tolerancia anafysed ‘l 27

Table 7.7 Growth in an open site expressed as unit leaf rate {E} and relative growth" rate (RGR) of seedlings
of six tree species of increasing shade tolerance compared to sunflower Helionrbus annzms. (La Selva, Costa
Rica)

É E as ‘Pía RCR

(grtrï WCClÜl] of Hefiantkzis i815” wcelfl) Ifáïfiairflflrts omziurs (i) 57: 1.00 (ii) 65"’ 0.75 Pioneer species

Ocbroma lagopos (i) S?" 87 0.85

(¡n 52s v9 0.81

Hefiocarfarss {Ippendícuíarus 425’ 65 0.54

Climax species

‘Gap-dependent spp.’

Cor-dia cziliodora 41"’ 64 0.48 Ternrirzalio obloazgz: 10“ 15 0.13 ‘Shade-tolerant Canopy spp.’

Brosímrrm alicastrttm 26'“ 40 0.15

PEÍIIÉLÏCÏEÍÏJÏLI nracroiolao‘ 22*" 3 4 0.15
From Oberhauer and [ïonnelliï (1986) table 2

taba: Different letters folloivíng E. show significantly tlífferent values

Table 7.8 Respirarion and photosynthesis of seetlling leaves of nine species, Gambari forest,
Nigeria
Photosynthesis Respira tion

a b c d Species group Apparent quantum NIcscJphyll Dark respiration Light compensation efficiency
conduetance (¡LLIÏIOl mi s4) point CO¿ (cía)

{mol CO2 photon") (mol CO3 m’: s71]

‘Xíeedv shruhs of

gaps (2 spp.) 0.037 0.028 1.75 4T" Pioneer trees _

(3 spp.)" 0.034 0.028 1.10 32 Lighedemandíng

clímax trees (2 sppfi 0.018 0.010 0.35 ‘Í 9 Sha de-tcilerant

clímax trees (2 spp.) 0.028 0.008 0.5 18

‘bïhitmore (1996} table l, laaseil on data of Riddoch et m‘. (11.991)

* Ceiba, fvlilicia, Riciziodenriror:

ïPres/ygoriz, Srercrrffii:

Weeds and pioneers have more efficient photosynthesis (a, h), higher dark respirarinn (c): and Eight
compcnsaríon point (d). They contrast in these rcspects with clímax Species.

128

plants were put back into low light conditions took over 60 minutes. Furthermore, the rate of carbon gain
varíed with past history and inter- mittent sunflecks could be used more efficiently than COHÍÍHUÓUS high
light. The conclusion is that the temporai pattern of sunfleck distribu- tion through the day (cf. Fig. 7.3) is of
impor- tance, not just the daily total PAR. Further studies like this are needed to analyse inter- specífic
differences of shade tolerance.

Seedling release of clímax species

The light climate

As he gets to know a forest the ecologíst or forester soon develops a feeling for how light- demanding the
different tree species are. Most

"obvious is the difference between pioneer and

clímax species. For the latter, there are also his perceptions on the size of gap in which seedlings are seen to
have grown up from the seedling bank. General observations on the natural history of the forestsuch as these
are an import- ant part of more rigorous scientific analysis, ini- tiaiiy as precursor to help the intelligent
design of experiments, and later to assist interpretation of the results. Nowhere is this general back-
IÜÜ

_ 3D E .5 _ v’: 4a — a? 2D - Ü [J 500 IÜÜU 150D 2000 250G Time (days)

o Hopsa bacteriana

A Hopsa nervosa

<> Parasharea maiaarranan I Shoreajohorensis

} Shade tolerants

} Light demanders

Fig. 7.18. Differentíal survival of dipterocarp

seedlings on the forest floor over 6 years, Danum,

Sabah.

Seedlings of shade-tolerant species, in this-case the two Hopea spp., charactcrístícally survive better in deep
shade.

Forest dynamics

ground knowledge better developed than in the dipterocarp rain forests of Malaysia, where foresters have
now nearly a century of accumu- lared experience. It has been found that amongst dipterocarps there are
species whose seedlings grow up in smail Canopy gaps and others that are successful in big gaps. Correlated
properties are that the first group have lengthy persistence in the seedling bank (Fig. 7.18) and, as trees, have
slow diameter growth and high density, dark coloured, naturally durable timber. Seedlings of the second
group die sotmer (Fíg. 7.18} and trees grow faster and have líghter, paler, non-durable tímber. The two
groups are known as shade-bearers and light-demanders or as medium to heavy hardivoods and light
hardwoods, respectively. The silvicultura] systems invented for dipterocarp rain forests, described later in
section 7.6, depend on these responses to gaps. Medium/heavy hardwood species can be successfully
regenerated by small Canopy disturbances; light hardwoods by more substantíal disturbances. Similar
categories are recognized in other rain forests, and índeed the whole of rain forest silviculture everywhere
depends upon it. It is the practical working out of the wide range of shade tolerance shown by different
clímax species that was discussed above in section 7.2.

Why is there a gap switch size between these ecological groups? For the dipterocarp species that occur in the
Danum forest, Sabah, this has now been elicited by observing over 6.4 years the growth, in artificial Canopy
gaps of different size, of seedlings from the forest floor seedling bank (Fig. 7.19}. lt can be seen in summary
that the light-demander species Shorea jolsrorensis was able to utilize for height growth the extra solar
radiation received in larger gaps and over a ltmger period of time. This is the ecologica] consequence at
population level of its ecopliysi- ological flexibility at the leaf and single seedlíng level. By contrast, the
shade-bearer Hopea Neruosa lacked the flexibility to adjust to and make use of the extra PAR of the larger
gaps. Success, however, is determined by many factors and Parashorea performed hadly because it

Growth and shade trafeiranee anaiïysed

[alïfimonths ‘es; s IEJ ‘É 5 g S s ‘E ‘ U1 .. 2 -+ E 8 U 4-0 4: I Ü’ . ‘a B? .2 s l: g 4- E A .

I} " ---------

S
CG
Mean height (mi

. ._ 47’

5 iii 35"" ¿o 25 3a lviean Canopy openness {Val

- a - Hopea nerviosa - I- - Parashorea mafaanonan

-—.¡— Shores‘ johorensis

occurred in very dense populations and these succumbed to insect damage.

N ti trients

The role of irradiance in. seedling growth and release is easy to observe and has been much investigared. By
contrast, little attention has been given to the potential role of plant mineral nutrientssl” The extent to which
these increase in a gap will vary virith the kind of gap independ- ent of its size (e.g. root plates expose subsoil
and elíminate-living roots). So far, nutrients seem unimportant compared to radiation. For example, four
independent experimente; on dípterocarp seedlings have shown no effect of added nutrients on growth; only
one of four

129

Fig. 7.19. Growth of dí pterocarp seedlings in

canopy gaps of different size over (a) 53 months,

(b) 77 months. Danum, Sabah. (Whítmore and

Brown 1996, fig. 2.}

In closed forest and tiny gaps, height growth was similar but then there was a switch and in larger gaps
Shores‘ foloorensis grew progressively faster with increasing gap size. The effect was stronger at 77 than 53
months. Parasborea, which natural history, tirnber density and leaf ecophysioltigy ' suggest has similar
ecology to Shorea, grew slowlg: because of serious repeated damage to its apex by the Cricket Nisirrus
z/ittatrts.
clímax species fertilized at Bukit Timah, Singapore responded; upper montane species in Jamaica failed to
respond; and in Ghana only three of fifteen species tested showed a clear response to soil fertílíty.

Overall the shadeinutrient ínteraction story remains unresolved. One part of the picture is likeiy to be that
there is no response to nutrients in dark conditions where irradiance is limíting, but a response at higher
irradiances.

Up to 50 species of Piper can co-exist in a single central American rain forest. These are sorted mainly on
radiation gradients with soil moisture and nutrients sometimes playing a role.

San ‘T. c»,

The concept of seedling hhade tolerance dissected

The preceding discussion has shown. that shade tolerance is a useful and easily understood concept that helps
frame analysis of how the forest works. It does, however, have thrcc dis- tinct, though overlappíng,
meanings. Seedlíngs may cliffer. in the levels of PAR required for seediing establishment (as in the clímax :
pioneer dichotomy), or they may have different longevity at a given level of PAR (Fig. 7.18), or they may
differ in the." amount of PAR required

for release. This third meaning has now been

analysed (Pig. 7.19) in an experiment which

revealed that shade-bearers do not increase. their -

130

growth much as gap size increases, wherease

light-demanders do, so are likely to ivin the.

growth race in all but the tiniest gaps.

Growth of saplings, poles and trees

The relative performance of the seedlings of dif- ferent species at the gap phase is the major determinan‘: of
the composition of the grosvth cycle. For example, at Danum by 6.4 years from gap creation seedlings had
become substantial polesm in the larger gaps, and the forest was into building phase, its species composition
probably determined for the next century or more. Thus it is natural that there has been such strong attention
paid to seedling behaviour.

It has now been shown several times that species may change their ranking as they grow’ up. For example, in
Queensland six species of the pioneer, ‘intermediate’ and ‘shade-tolerant’ guilds planted as seeds in gaps of
various sizes changed their rank order of height over seven yearsd” In Ghana there is a group of species that
need high light at first but later grow in deep shade.” Observations on Kolombangara, Solomon Islands, of
1700 trees of 12 species over 30 years has shoivn they change their eco- logical characteristics in a complex
way through life (Fig. 7.20).

Seedlings may show a burst of growth then stagnate as the Canopy closes over them and by the time they
reach tree size will have had several such sports. This phenomenon is called multiple release. ln temperate
forests it is easy to detect from changes in the EVÍdth of annual growth rings in the wood. It is more diffictilt
to demonstrare in tropical rain forests but undoubtedly occurs. Polycylic silviculture (section 7.6), for
example, makes use of it. Side light as well as light from gaps overhead has been shown to stimulate
groivthï”

Ecophysiology of dipterocarpsm

Those dipterocarp species investigated have been found all to have similar maximumphoto- synthesis rates at
low levels of irradiance, with

Forest dynamics

High growth rate

—4 ‘- ¿h High mortality rata :3 High crown expnsure I.|.I Low timber density

—3 -

2*
9

"5

É _2 _ Elaeacarpus
É Campncrsperma
E‘ ¡gfiiiienia m

f D o Endnspermum :1 f

D -l — z - - a

_ A Terminafra ,_.

‘a Gmefina g

E’ E

‘¡'- Ü - "- a Scfifzamaría 3

E i‘ Ü. peekeiii 3

a ' Parinan S
1 ‘Pamena m

4—' + _ . - . _ . v ' ' ' . " o,

É ‘ ' ‘C CJieo-abudicum

'+—

É
C7

(f) .

E +2 —
8
CD
Low growth rate

+3 _ 3 Low mortality rate _ Maranmgs 5 Low crown exposure High tumbar density 5-40 30-50 21H]
Diameter {cm}

Fig. 7.20. Changes in successional status through

life, Kolombangara, Solomon Islands, twelve species

monitored 1964-93. {Burslem and Whitrnore

unpublished.) This complex diagram shows how the 12 common big tree species change successional rank
through life. The four ecological groups they form as seedlings (Table 7.5) have already lost their integrity by
5-10 cm diameter, hut by 7G cm there is still a roughly’ similar rank order.

the light-demanding, light hardwood species responding more strongly to high irradiance. Ten species tested
were, as a group, less respons- ive to irradiance than the pioneer Macaranga degüressa, but much more
responsive than a shade-divelling Begonía.

Dryobafanops faziceolata, a shade-tolerant, has been shown to have a higher carbon fixation capacity at low
irradiance than Sborea ¡lepra- sufa, a well-known iight-demander. This was due to lower compensation point
and dark res- piration (cf. Table 7.8), and higher mean assimi- lation rate. Dryobalanops has different

Silvicultura
131

Fig. 7.21. Seedling architecture of a Iight-demander and a shade-bearer, the dipterocarps Shores ieprosuia
and

Dryobaianops larzceolara. {Zipperlen and Press 1996, fig. 2.}

These were grown at c. "E 0 per cent Canopy openness and have the same total axis length. These tivo
grmvth forms are common amongst rain forest seedlings. They have been dubbed optirnísts and pessimists,
respec- tively, by Kohyama {i987} and represent a trade-off between success after a gap opens in the future
and maxima! interception of scattered sunflecks in the present.

architecture (Fig. 7.211) putting much more of its groivth into lateral branches, an efficient seedling shape for
exploiting patehy sunflecks. Similar differences in architecture existed between the species shown in the
growth experi- ment in Fig. 7.19.

Different dipterocarps have also been found

to differ in their ability to tolerate drought periods. At the physiological level, Dryobafanops lanceoiara
seedlings were shoivn to have ioiver stomatal conductance than other species yet about the same level of
carbon assimilation. Over a day the stornata were open for a shorter period and so the plants lost less water
by tran- spiration. Thus, the species had higher water-use efficiency. Foresters know that nursery-raised
Dryobalanops seedlings survive better on plant- ing out. Ability to survive drought periods has been shown to
be the major factor determining the success of Sfcrtnrea curtisii on coastal hills and inland ridges in Malaya
(Fig. 2.3) and differen- tial mortality of three Shores species in the regular early year drought has been
demon- strated in Penang. In the fairly seasonal rain

forest climate of Sri Lanka the interplay of irra- dianee and drought has been shown to account for the
partitioning of four Shores species

between ridge, siope and valleyxl”

7.6 SILVICULTURE
Rain forest silviculture is the manipularion of

the forest to favour certain species and thereby to enhance its value to humans. Por example, around the turn
of the twentieth century the Malayan rain forest was treated to increase the abundance of Palaquium guita
(Sapotaceae) because there uvas a strong market for its latex, gotta percha, which was used as the insulator in
submarine telegraph cables?“ More commonly, manipulation favours particular timber species. Timber
properties, whether heavy or light, dark or pale, durable or not, are strongly correlated with growth rate and‘
thus to the extent to which the species is light-demanding (Table 7.5). Thus, the ecoiogicai basis of natural
forest silviculture is the manipularion of the forest

132

eanopy. The biological principle of silviculture is that hy eontrolling Canopy gap "size it is P055? ible to
influence species composition of the next growth cycle. The bigger the gaps the more fast- grtiwing light-
denianders will be favoured. This concept has been known in continental Europe since at least the tweith
century. lndeed, our knowledge of forest dynamics, the forest groivth cycle and the significance of Canopy
gap size, varas discovered by foresters einpiricatlly before the formal science of ecology came into exist-
encedf” The practice of silviculture reached tropical rain forests via India and Burma then Malaya-i and was
strongly influenced hy Germanic tra dition?“
Silvicultura] systems

The silvicultural systems that have been zigiplied to tropical rain forests belong to one of two kinds: the
polycyiclic and lïlíïfllücyiïlic systems, respectively (Figs. 722-724). As the name implies, polycyclic systems
are based on the repeated removal of seiected trees in a continu- ing series of fclling cycles, whose length is
less than the time it takes the trees to mature (which forcsters call the rtitation age). The aim is to remove
trees before they begin to stagnare and

Forest ¿Iyizamics

deteriorate from old age, leaving all zippreciating stems to swell the future yield. Because of the very
species-rich nature of most tropical rain forests, and the relaitivelyy small number of species with timber that
is commercial by‘ current standards, extraction on a polycyclic system tends to result in the formation of
seat- tered smali gaps in the forest canopy.

By contrast, monocyclic systems remove all saleable trees at a single (iperatiran, and the length of the cyrele
more or less equals the rota- tion age of the trees. Except in those cases where there are few saleable trees,
damage to the forest is more drastic than under a polycyclic system, the Canopy is more extensively
destroyred, and bigger gaps are formed. Furthermtire, new crop trees all grow up together and this can leag to
oirercrowding, competition between their crowns, and reduced growth rate.

it can be seen at once that the ÍVVO kinds of system will tend to favour shade-bearing and light-demanding
species, respectively, but the extent of the. difference will depend on hour many trees are felled at each cycle
in a pcily- cyclic system. The nub of the difference is that, on the one hand, polyíïyclic systems retain
ativanced growth of half-groivn trees to produce marketable trees at subsequent intermediate

Fig. 7.22. Lowlzind rain fcnrest,

Xïiindsor Tableland, Queensland,

twelire years after logging on a

[Jolycyclic system. Only selected big trees were felled. The Canopy is more broken than in Virgin forest, and
the lcigging road {ivhich will be used again next feiling cjrcle] is clearly visible.

Silvicultura

felling cycles within thc rotation. This is; done by taking good care ofthese ‘adolescent’ trees at each felling.
On the other hand, monocyclic systems forego the increment already accumu- lated in these adolescents and
rely almost entirely onvseedlings to produce the next crop,

kwhich will be ready to harvest only at the. fuil

totation age. If there does happen to be any advanced growth left from the previous forest it is abonos, over
and above the seedling growth on which monocyclic systems are based. f Under the best conditions,
polycyclic systems can greatly increase the timber yield over a full rotation. Their success is" ‘utterly
dependent on‘
Fig. 7.23. Lowland dipterocarp

rain forest, inPahang, Peninsular Malaysia, shortly after logging v. on the monocyclicMalayan Uniform
System. _ All mature commercial stems have been removed, mostly . ‘dipterocarps, and the yield would have
been 20-30 m3 h a“. The forest matrix remains « intact. Note lush regrowth along the logging track.

advanced growth and this meansthat the forest must be adequately stocked VJÍÍlLEÏIÓOlBSCÉHÍS
beforefelling begins, and damage to them by

" loggingkept acceptably low, If loggÏing damage at each felling exceeds a iow minimum, regener- _ ation
will be lost and the yield will progressiveiy

fall off through a series of cycíes.,Co-ntrolcan be achieved either by rnarking trees foriretention beforefelling
begins, ivhich is labour-intïensiire, or by felling down to a rather high diameter

"limit.

4 The often-ciaimed disadvantage of polycyclic systems is that the faster-growing genotypes are

“progressivelyremoved;'because at each felling

134

many or all of the stems that have reached a minimum prescribed girth are removed. The small stems
retained Inay bc either young and potentially fast-growing or old, stunted, and unable to respond to release.
In fact, both fast and slow growth are also determined partly by phenotype (i.e. microsite) as well as by geno-
type. The magnitude ‘of the dísadvantagezüï‘ is therefore anybodfs guess. It can be mitigatecl, as was tlonc in
Queensland, by flexibílity in the rules used to determine whether a tree is felled or not. In Queensland every
species had a specified diameter at which it was xiormallyr felled, but vígorous individuals were retained for
a later cycle and poorly foorrned feeble ones were felled at smaller size.

Silvicultural treatment before felling has sometimes been practísed in the past in order to reduce numbers of
clímbers or to boost the growth of juveniles. Post-felling silviculture to remove deformed stems or to free
regrowth from competitionwvas also wídely practised until recently. At its fullest development several
treatments were given over the first decade-or so after felling. Post-felling treatments have been largely
abandoned, because of the high cost. It was demonstrated in Saranïak that a given sum of money gave a
better return in terms of growth increment in the regroïving forest if

Forest dynamics

Fig. 7.24. Lovvland dipterocarp rain forest in Sabah, East. Malaysia, shortly after logging ton the ira riant of
the monocyclic Malayan Uniform silvicultural system used there,-l\/1.D. Swaíne observes (1987). i p The
forests of this region have a very high proportiotl of commercial stems. Removal of these, as here, commonly
v yields 100 m3’ or more of ti m berha‘l, and only sca ttered big trees remain. '

invested in reducing damage by the logging operation itself rather than in post-felling silvi- culture.
Nowadays it is generally the case that the felling operation provides ‘the only Canopy manipuladora the
foresters can afford.

Criteria for success

Silviculture (Figs. 7.22——7.24) will be successful so long as it is practised within the biological límits of
the forest. The desired species must be capable of growing in the size of gap that is created.

Forests are a potentially renewable source of timbcr, and their use for timber production is not simply a
quarrying operation. The success of silviculture may be defined as the bringing to maturity of a stand of
timber trees with regener- ation below them to form the next silvicultural cycle, and without site
deterioration, such as soil erosion or degradation.

The failure of silviculture follours from working beyond the límits of the inberent dynamic capabilities of the
forest ecosystem. This is commonly beca use rules drawn up by silvi- culturists are not enforced, often
because of political intervention. It may‘ also be because economists, eager to enrich a Iiation, enforce their
dismal pseudoscienee to override basic bio-

Silvicultttre l

logical principles ¿ind tlictate the removal of a larger harvest than the forest can sustain without tlegradaition.

iïïoocïy climbers

Univanted ‘tveed’ tree species can be a problem. So too can woody climbers. The Iiatural behav- iour of
many of the latter is to grow vigorously‘ in strong illuminarion. Thus they are commoon at ftirest fringes,
stich as river banks (and give rise to the myth of the irnpenetralale jungle}, and in big Canopy gaps. ln many
tropical rain forests the removal by logging of many trees per hectare creates large openings ideal for
vigorous climber growth. In many Eastern rain forests dense carpets of the big-leaved tviry convolvtil-
aceotis climbers Merremia spp. smorher the residual forest and form clirnber towers up remaining isolated
trees (Fig. 7.25). Trees even- tually grow up and carry the carpet skywards but forest recovery is delayed and
the trees may develop kínked stems (Fig. 7.42). Very little is known about the ecology of woody climbers.
For example, in the Solomon Islands climbers do- not proliferate excessively’ after cyclones but become
extremely abundant after logging which causes similar Canopy damage. In Sabah, lidera-etnia, Mezoneuron
and Uncarïa, xxrhicb are very serious impediments to forest regeneratitan,

ba U1

Fig. 7.25. Climber tower-s,

Enoghac, Solomon Islands.

This very heavily logged forest has become CaJÏpCtí-Jtl with the wiry laig-leai-‘cd climber Meri'eírtírt.

are uncommon in Virgin forest, perhaps because in this country massive Canopy opening does nor occur
naturally.
Damage

The amount of damage to the forest tlepends more on how many trees are felled than on timber volume
extracted. lt is commonly the case that for every’ tree removed‘ for timber {logged} a second tree is totally
smashed and a third tree receives damage from which it will recover (Fig. 7.2 l. ln Queensland (Fig. 7.27) the
number of stems sittashed was a similar pro- portion but before strict rules avere introduced

in 1982 the proportion damaged Vx-‘as much.

higher.

The pattern of damage differs between extrac- tion by tractor and by overheatl cable, so-called high-lead
logging. The latter causes extreme devastation along the paths of the cables and little between. Seedlings
survixre better than bigger plants. In tractor logging the damage is spread more uniformly and secdling
tiumbers can be very seriously depleted.

For successful recovery from logging the single most important "Criterion is to minimize damage to the
forest floor, for several reasons. On the floor occur the seedlings of clímax species. Within the surface soil
lies the seed

1988
Sabah
1953
Sabah
CD

h Cl

a stems of commercial

specieseüé‘ mín diameter h) G

For key see Fig. 7.25

Forest dggnantics
196G
Nigeria
19??
Sarawak

Fig. 7.26. Effects of logging [total commercial stems ha” in parcntheses} in various forests. {After Nicholson

and Keys unpttblishcd.)

tits the number of stems lost per hectare (by logging or smashed during Ítigging) dccreases the number that

survive [lntact or damaged) increases. Note that in the first three forests as many trees were smashed as ' were
logged and that the Nigerian site, which had fewest commercial stems, had fcwest smashed or

damaged trees.

la] Commercial species [bl All species Before 1982 Alter 1982 Before 1982 After 1982 i156] i136] i335] Key

"¡a Stemsa-ÜZ rn in diameler

Lost Survived

l resídirai stantll

Fig. 7.27. Effects of selective logging, Queensland, on (al commercial species and [bl all species, before and
after introduction of strict silvicultural rules in 19'82. Total stems ha” in parentheses. [From Nicholson and

Keyra‘ Impublisheti.)

The new rules reduced the numbers of stems smashed or damaged. Tlïicir effect was about the same on the

whole forest on itist the commercial species.

bank. Most of the roots and mineral nutrients are usually in the top O.1—0.3 m of the soil. The surface root
and humus layers allow good infiltration of rainfall. It has been clearlyr demonstrated several times that
damage to the cancipy‘ and forest floor can be minimized by the Careftil planning of extraction roads and
tractor paths {so-called snig tracks). In Sarawak such planning enabled 36 per cent more timber to be moved
per hour; overall

costs were '19 per cent down and open spaces in the forest were reduced from 40 to 17 per cent, with 6D
commercial stems per hectare stirtrixring rather than 40. In Queensland the new strict rules of 1982 (above)
not only reduced the tiumbers of trees smashed or damaged {hence increasing the number that survived
íntact; Fig. 7.2?) but aiso reduced snig tracks to 9 per cent of the area, only 2 per cent with su bsoil exposetl.

Silvicultura 137

Soil compaction along tracks is an ecological problem. Infiltration of rainfall is strongly reduced. Plants are
unable to establish on these surfaces which they do not encounter naturally. Tracks can be raked to break up
the compacted soil, or specially designed tyres used on wheeled vehicles which have a low load per unit area.
Wheeled vehicles cause more compaction than tracked onesxwl '

Erosion and bydrology

Another problem. created by logging is erosion, which can be disastrous (Fig. 7.28) and whose extent has
now been well studieddü‘; Streamflow increases for 5-«10 years after logging and is greater after rain than in
unlogged forest because overland flow rises. Peak tvater dis- charges also increase after logging, but the
increase is rarely statistically significant. There is a pulse of increased sediment of two- to ten» fold by road
construction and up to 2Ü—fold by logging, but by 5 years sediment loads are only twice those from the
original forest. Simple pre- cautions can reduce extra sediment down to a quarter of these figures, ‘for
example, by appro- priate alignment and design of cxtraction tracks, including the construction of cross
drainsdll“ lt

is also important not to block streams or the forest is killed in the lakes that then tlevelop.

Low impact logging

The management of the rain forests of Queensland in the early 19805. most nearly approached these ideals
Lmtil, after intense pres- sure from environmentalists, all logging viras stopped at the time they were
declared a World Heritage area (Table 10.15}. i

In the mid—1990s a change in sentiment occurred and throughout _tbe tropics the will began to emerge to
harvest timber with minimal

_ damage to the forest. The rising international

concern about the heavy human impact on trop- ical forests and the mechanisms evolving to control it are
analysed in Chapters IO and 11. Good logging practice, larocedures well known to foresters but previously
usually unenforce- able, are ideas whose time had now come (Figs. 7.29, 7.30}. Pre-planning of snig tracks,
direc- tional felling to minirnize damage to the canopy and facilitate log removal, and winching in lUgS by
20-30 rn of cable to reduce damage to the forest floor, are essential. Climber cutting some months before
felling can" reduce damage to forests with abundant big Kvoody-climbers but is

Fig. 7.28. Erosion along tracks built in rain forest climates can be spectacular as here in Nialaya.