The Ecology of Trees in the Tropical Rain Forest

The Ecology of Trees in the Tropical Rain Forest

The Ecology of Trees in the Tropical Rain Forest

314 Pages ·2004·3.01 MB ·English

The Ecology of Trees in the Tropical Rain Forest

This page intentionally left blank TheEcologyofTreesintheTropicalRainForest


Currentknowledgeoftheecologyoftropicalrain-foresttreesislimited,with


detailedinformationavailablefor perhapsonly a few hundredof the many


thousands of species that occur. Yet a good understanding of the trees is


essential to unravelling the workings of the forest itself. This book aims to


summarisecontemporaryunderstandingoftheecologyoftropicalrain-forest


trees.Theemphasisisoncomparativeecology,anapproachthatcanhelpto


identifypossibleadaptivetrendsandevolutionaryconstraintsandwhichmay


alsoleadtoaworkableecologicalclassiWcationfortreespecies,conceptually


simplifying the rain-forest community and making it more amenable to


analysis.


Theorganisationofthebookfollowsthelifecycleofatree,startingwith


thematuretree,movingontoreproductionandthenconsideringseedgermi-


nation and growth to maturity. Topics covered therefore include structure


andphysiology,populationbiology,reproductivebiologyandregeneration.


ThebookconcludeswithacriticalanalysisofecologicalclassiWcationsystems


fortreespeciesinthetropicalrainforest.


IANTURNERhasconsiderableWrst-handexperienceofthetropicalrainforests


of South-EastAsia,havinglived and worked in the region for morethan a


decade.AftergraduatingfromOxfordUniversity,hetookupalecturingpost


attheNationalUniversityofSingaporeandiscurrentlyAssistantDirectorof


theSingaporeBotanicGardens.HehasalsospenttimeatHarvardUniversity


asBullardFellow,andatKyotoUniversityasGuestProfessorintheCenter


forEcologicalResearch. MMMM CAMBRIDGE TROPICAL BIOLOGY SERIES


EDITORS:


PeterS.AshtonArnoldArboretum,HarvardUniversity


StephenP.HubbellPrincetonUniversity


DanielH.JanzenUniversityofPennsylvania


PeterH.RavenMissouriBotanicalGarden


P.B.TomlinsonHarvardForest,HarvardUniversity


Alsointheseries


Tomlinson,P.B.Thebotanyofmangroves


Lowe-McConnell,R.H.EcologicalstudiesintropicalWshcommunities


Roubik,D.W.Ecologyandnaturalhistoryoftropicalbees


Benzing,D.H.Vascularepiphytes


Endress,P.K.DiversityandevolutionarybiologyoftropicalXowers MMMM The Ecology of Trees in


the Tropical Rain Forest


I.M. TURNER


SingaporeBotanicGardens uf770uf775uf762uf76cuf769uf773uf768uf765uf764 uf762uf779 uf774uf768uf765 uf770uf772uf765uf773uf773 uf773uf779uf76euf764uf769uf763uf761uf774uf765 uf76fuf766 uf774uf768uf765 uf775uf76euf769uf776uf765uf772uf773uf769uf774uf779 uf76fuf766 uf763uf761uf76duf762uf772uf769uf764uf767uf765


The Pitt Building, Trumpington Street, Cambridge, United Kingdom


uf763uf761uf76duf762uf772uf769uf764uf767uf765 uf775uf76euf769uf776uf765uf772uf773uf769uf774uf779 uf770uf772uf765uf773uf773


The Edinburgh Building, Cambridge CB2 2RU, UK


40 West 20th Street, New York, NY 10011-4211, USA


477 Williamstown Road, Port Melbourne, VIC 3207, Australia


Ruiz de Alarcón 13, 28014 Madrid, Spain


Dock House, The Waterfront, Cape Town 8001, South Africa


http://www.cambridge.org


©I. M. Turner 2004


First published in printed format 2001


ISBN 0-511-04148-9 eBook (netLibrary)


ISBN 0-521-80183-4 hardback CONTENTS


Preface pagexi


Acknowledgements xiii


1 Introduction 1


Thetropicalrainforest 1


Tropicaltrees 2


Species 2


Tropicalrain-forestdiversity 6


Adaptation 9


Theimportanceofphylogeny 10


TheecologicalclassiWcationoftropicalrain-foresttrees 13


2 Thegrowingtree 15


Trees:form,mechanicsandhydraulics 15


Treestature 15


Wood 18


Tropicaltimber 21


Themechanicaldesignoftrees 24


Buttresses 28


Leaningtrees 34


Treefall 35


Sapascent 35


Treehydraulics 37


Treearchitecture 40


Allometry 41


Bark 46


Roots 46


Roothemi-parasites 48


vii Contents


viii


Mineralnutrition 50


Mycorrhizas 52


NitrogenWxation 56


Othermethodsofobtainingextranutrients 57


Comparativeuseofnutrients 58


Elementaccumulation 60


Leafformandphysiology 60


Size,shapeandotherstructuralcharacteristics 62


Leafprotection 67


Thedefencesoftropicalforesttrees 69


ClassiWcationofplantdefences 81


Foliarbacterialnodules 82


Leafdevelopment:colouredyoungleaves 83


Leaflongevity 86


TheinXuenceofshade 90


Highlightconditions 94


OtherfactorsinXuencingleafperformance 95


Speciesgroupsbasedonleafcharacteristics 96


3 Treeperformance 103


Age,sizeandgrowthintropicalrain-foresttrees 103


Thedynamicsoftreepopulationsintherainforest 105


Mortalityintrees 105


Treegrowthintheforest 108


Treeperformanceinrelationtolightclimate 111


Mortality,growthandadultsize 114


Theuseofgrowthandmortalitydatatorecognisespecies


groups 116


Relativeperformanceofspeciesofsimilarlifehistory 118


Whatlimitstreegrowth? 119


Herbivory 121


4 Reproductivebiology 122


Reproduction 122


Vegetativereproduction 122


Sexualsystems 123


Geneticdiversity 125


WhentoXower? 127


Pollination 130


Pollinationsyndromes 131


TropicalXowers 144


This page intentionally left blank TheEcologyofTreesintheTropicalRainForest


Currentknowledgeoftheecologyoftropicalrain-foresttreesislimited,with


detailedinformationavailablefor perhapsonly a few hundredof the many


thousands of species that occur. Yet a good understanding of the trees is


essential to unravelling the workings of the forest itself. This book aims to


summarisecontemporaryunderstandingoftheecologyoftropicalrain-forest


trees.Theemphasisisoncomparativeecology,anapproachthatcanhelpto


identifypossibleadaptivetrendsandevolutionaryconstraintsandwhichmay


alsoleadtoaworkableecologicalclassiWcationfortreespecies,conceptually


simplifying the rain-forest community and making it more amenable to


analysis.


Theorganisationofthebookfollowsthelifecycleofatree,startingwith


thematuretree,movingontoreproductionandthenconsideringseedgermi-


nation and growth to maturity. Topics covered therefore include structure


andphysiology,populationbiology,reproductivebiologyandregeneration.


ThebookconcludeswithacriticalanalysisofecologicalclassiWcationsystems


fortreespeciesinthetropicalrainforest.


IANTURNERhasconsiderableWrst-handexperienceofthetropicalrainforests


of South-EastAsia,havinglived and worked in the region for morethan a


decade.AftergraduatingfromOxfordUniversity,hetookupalecturingpost


attheNationalUniversityofSingaporeandiscurrentlyAssistantDirectorof


theSingaporeBotanicGardens.HehasalsospenttimeatHarvardUniversity


asBullardFellow,andatKyotoUniversityasGuestProfessorintheCenter


forEcologicalResearch. MMMM CAMBRIDGE TROPICAL BIOLOGY SERIES


EDITORS:


PeterS.AshtonArnoldArboretum,HarvardUniversity


StephenP.HubbellPrincetonUniversity


DanielH.JanzenUniversityofPennsylvania


PeterH.RavenMissouriBotanicalGarden


P.B.TomlinsonHarvardForest,HarvardUniversity


Alsointheseries


Tomlinson,P.B.Thebotanyofmangroves


Lowe-McConnell,R.H.EcologicalstudiesintropicalWshcommunities


Roubik,D.W.Ecologyandnaturalhistoryoftropicalbees


Benzing,D.H.Vascularepiphytes


Endress,P.K.DiversityandevolutionarybiologyoftropicalXowers MMMM The Ecology of Trees in


the Tropical Rain Forest


I.M. TURNER


SingaporeBotanicGardens uf770uf775uf762uf76cuf769uf773uf768uf765uf764 uf762uf779 uf774uf768uf765 uf770uf772uf765uf773uf773 uf773uf779uf76euf764uf769uf763uf761uf774uf765 uf76fuf766 uf774uf768uf765 uf775uf76euf769uf776uf765uf772uf773uf769uf774uf779 uf76fuf766 uf763uf761uf76duf762uf772uf769uf764uf767uf765


The Pitt Building, Trumpington Street, Cambridge, United Kingdom


uf763uf761uf76duf762uf772uf769uf764uf767uf765 uf775uf76euf769uf776uf765uf772uf773uf769uf774uf779 uf770uf772uf765uf773uf773


The Edinburgh Building, Cambridge CB2 2RU, UK


40 West 20th Street, New York, NY 10011-4211, USA


477 Williamstown Road, Port Melbourne, VIC 3207, Australia


Ruiz de Alarcón 13, 28014 Madrid, Spain


Dock House, The Waterfront, Cape Town 8001, South Africa


http://www.cambridge.org


©I. M. Turner 2004


First published in printed format 2001


ISBN 0-511-04148-9 eBook (netLibrary)


ISBN 0-521-80183-4 hardback CONTENTS


Preface pagexi


Acknowledgements xiii


1 Introduction 1


Thetropicalrainforest 1


Tropicaltrees 2


Species 2


Tropicalrain-forestdiversity 6


Adaptation 9


Theimportanceofphylogeny 10


TheecologicalclassiWcationoftropicalrain-foresttrees 13


2 Thegrowingtree 15


Trees:form,mechanicsandhydraulics 15


Treestature 15


Wood 18


Tropicaltimber 21


Themechanicaldesignoftrees 24


Buttresses 28


Leaningtrees 34


Treefall 35


Sapascent 35


Treehydraulics 37


Treearchitecture 40


Allometry 41


Bark 46


Roots 46


Roothemi-parasites 48


vii Contents


viii


Mineralnutrition 50


Mycorrhizas 52


NitrogenWxation 56


Othermethodsofobtainingextranutrients 57


Comparativeuseofnutrients 58


Elementaccumulation 60


Leafformandphysiology 60


Size,shapeandotherstructuralcharacteristics 62


Leafprotection 67


Thedefencesoftropicalforesttrees 69


ClassiWcationofplantdefences 81


Foliarbacterialnodules 82


Leafdevelopment:colouredyoungleaves 83


Leaflongevity 86


TheinXuenceofshade 90


Highlightconditions 94


OtherfactorsinXuencingleafperformance 95


Speciesgroupsbasedonleafcharacteristics 96


3 Treeperformance 103


Age,sizeandgrowthintropicalrain-foresttrees 103


Thedynamicsoftreepopulationsintherainforest 105


Mortalityintrees 105


Treegrowthintheforest 108


Treeperformanceinrelationtolightclimate 111


Mortality,growthandadultsize 114


Theuseofgrowthandmortalitydatatorecognisespecies


groups 116


Relativeperformanceofspeciesofsimilarlifehistory 118


Whatlimitstreegrowth? 119


Herbivory 121


4 Reproductivebiology 122


Reproduction 122


Vegetativereproduction 122


Sexualsystems 123


Geneticdiversity 125


WhentoXower? 127


Pollination 130


Pollinationsyndromes 131


TropicalXowers 144


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