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Analysing Patterns and Processes in Anthropogenically Altered Tropical Forests Studies from the Guiana Shield and West Africa

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Analysing Patterns and Processes in Anthropogenically Altered Tropical Forests Studies from the Guiana Shield and West Africa ANURAN COMMUNITIES ON THE CUTTING EDGE: ANALYSING PATTERNS AND PROCESSES IN ANTHROPOGENICALLY ALTERED TROPICAL FORESTS Studies from the Guiana Shield and West Africa Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Bayerischen Julius-Maximilians-Universität Würzburg Anuran Communities on the Cutting Edge: Analysing Patterns and Processes in Anthropogenically Altered Tropical Forests Studies from the Guiana Shield and West Africa Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Bayerischen Julius-Maximilians-Universität Würzburg vorgelegt von Raffael Ernst aus Bad Hersfeld Würzburg 2006 Eingereicht am............................................................................................................05.05.2006 Mitglieder der Promotionskommission: Vorsitzender: Prof. Dr. M. J. Müller Gutachter: Prof. Dr. K. E. Linsenmair Gutachter: Prof. Dr. W. Hödl Tag des Promotionskolloquiums.......................................................................................... Doktorurkunde ausgehändigt am.......................................................................................... CONTENTS PREFACE Page 1 LIST OF PUBLISHED CONTRIBUTIONS 2 CHAPTER 1 GENERAL INTRODUCTION 3 1.1 Communities: The search for basic patterns and elementary processes 3 1.2 Two realms: The origins of similarities and differences in two tropical forests 7 1.3 The cutting edge: Logging and the history of anthropogenic disturbance 13 1.4 Set up of thesis 25 CHAPTER 2 METHODS AND MODEL SYSTEMS 28 Methods & Model systems 2.1 Measuring and monitoring amphibian diversity in tropical forests. I. An evaluation of methods with recommendations for standardization 28 M.-O. Rödel and R. Ernst 2.2 On the cutting edge – The anuran fauna of the Mabura Hill Forest Reserve, Central Guyana 43 R. Ernst, M.-O. Rödel and D. Arjoon CHAPTER 3 PATTERNS 60 The ‘ground’ state - general community patterns 3.1 Community assembly and structure of tropical leaf litter anurans 60 R. Ernst and M.-O. Rödel Crossing the line - anthropogenically induced community pattern changes 3.2 Anthropogenically induced changes of predictability in tropical anuran assemblages 89 R. Ernst and M.-O. Rödel 3.3 Disturbance related predictability patterns of community composition in two tropical treefrog assemblages 106 R. Ernst and M.-O. Rödel CHAPTER 4 PROCESSES 126 Explaining the transition – factors influencing pattern changes 4.1 The impacts of selective logging on three sympatric species of Leptodactylus in a central Guyanan rainforest 126 R. Ernst, T. Konrad, K. E. Linsenmair and M.-O. Rödel 4.2 Diversity erosion beyond the species level: Dramatic loss of functional diversity after selective logging in two tropical amphibian communities 153 R. Ernst, K. E. Linsenmair and M.-O. Rödel CHAPTER 5 EXTENDED SUMMARY AND SYNTHESIS 183 5.1 Amphibian communities in disturbed forests: lessons from the Neo- and Afrotropics 183 R. Ernst, K. E. Linsenmair, R. Thomas and M.-O. Rödel CHAPTER 6 ALLGEMEINE ZUSAMMENFASSUNG 216 ACKNOWLEDGEMENTS 224 CURRICULUM VITAE 226 EHRENWÖRTLICHE ERKLÄRUNG 228 FIELD GUIDE CD-ROM inside back cover PREFACE (…)”The theory of ecosystems contains an important distinction between what is complicated without any Gestalt or unifying principles (…) and what is complex (…) A multiplicity of more or less lawful, interacting factors may operate together to form a unity, a system (…) Organisms, ways of life, and interactions in the biosphere in general, exhibit complexity of such an astoundingly high level as to color the general outlook of ecologists. Such complexity makes thinking in terms of vast systems inevitable. It also makes for a keen, steady perception of the profound human ignorance of biospherical relationships and therefore of the effect of disturbances” (…) ARNE NAESS (Phil. Founder Of Deep Ecology, 1972) Complexity [not complication] and the effects of disturbances on complex systems are the very objects of concern addressed in the present thesis. An investigation of complex systems usually begins with the search for patterns, unifying principles if you like. In the following studies, these patterns are consequentially analysed on the basis of modern theories on biodiversity and community ecology with a special emphasise on the role of anthropogenic disturbance (logging) and its influence on general community properties. Timber lot Mabura Hill, Guyana R. E. 1 LIST OF PUBLISHED CONTRIBUTIONS At the time of submission of this thesis, the following contributions have either been accepted for publication or were submitted to the peer reviewed journals listed in parentheses. Other contributions have previously been published in peer reviewed journals as indicated by full reference. Measuring and monitoring amphibian diversity in tropical forests. I. An evaluation of methods with recommendations for standardization – Ecotropica 10: 1-14. On the cutting edge – The anuran fauna of the Mabura Hill Forest Reserve, Central Guyana – Salamandra 41: 179-194 Community assembly and structure of tropical leaf litter anurans – submitted to Ecotropica. Anthropogenically induced changes of predictability in tropical anuran assemblages - Ecology 86: 3111-3118 Disturbance related predictability patterns of community composition in two tropical treefrog assemblages – not yet submitted The impacts of selective logging on three sympatric species of Leptodactylus in a central Guyanan rainforest - submitted to Amphibia-Reptilia Diversity erosion beyond the species level: Dramatic loss of functional diversity after selective logging in two tropical amphibian communities – accepted Biological Conservation Amphibian communities in disturbed forests: lessons from the Neo- and Afrotropics – in Tscharntke et al. (eds.): Stability of tropical rainforest margins linking ecological, economic and social constraints. Springer – Environmental Science Series (in press) 2 CHAPTER 1 GENERAL INTRODUCTION 1.1 Communities: The search for basic patterns and elementary processes Just as evolutionary biology can be regarded as the all-embracing and interconnecting discipline in all modern fields of biology, so can the study of communities be seen as the overarching concept in ecology, without which, patterns observed at lower organizational levels appear out of context. The following statement by Victor Shelford (1929) may hence serve as a guiding principle to be kept in mind throughout the following paragraphs of this thesis. It highlights the discipline’s function as a connecting theme. (…) “ecology is the science of communities. A study of the relations of a single species to the environment conceived without reference to communities and, in the end, unrelated to the natural phenomena of its habitat and community associations is not properly included in the field of ecology”(…) VICTOR SHELFORD (Laboratory and Field Ecology, 1929) Despite its metamorphosis from a purely descriptive discipline preoccupied with identifying and listing species found in a particular location to a “hard” analytical science with significant methodological advances that makes use of a wealth of sophisticated mathematical models and statistical tools, fundamental questions in community ecology have changed little in the past decades. And these questions are astoundingly simple: Who lives with whom and why? This in turn leads directly to another basic question and an important goal that community ecologists try to achieve, namely to understand the origin, maintenance, and consequences of biological diversity within local communities. It is this search for basic and complex community patterns and the processes that create them that drives modern community ecology. Very often these patterns and processes are simply experimentally intractable and experimental manipulations of entire communities are impossible. As community patterns are the consequence of a hierarchy of interacting processes, the 3 information to be acquired for meaningful interpretation must come from various sources such as mathematical models, statistical tests, and experiments. However, in revealing patterns and identifying the underlying processes, community ecology will prove an indispensable tool providing the necessary knowledge in order to wisely manage ecological systems that we as biological organisms depend on, natural eco systems that are increasingly stressed and fragmented and that we have damaged through either habitat destruction or overexploitation. Box 1.1: Community patterns – the hierarchy of interacting processes Physiological Geological Evolutionary constraints History History The species composition of a Longitude given local community is the result of Altitude various hierarchically interacting processes. The upper limit on the Climate & species composition of a local Rainfall community is set by the composition of the regional species pool. Membership in Regional the regional species pool is constrained species pool by, e.g. geological history of respective regional regions, and the evolutionary processes local of speciation within different taxonomic groups. .Membership in the local species pool, on the other hand, is more Habitat selection Natural disturbance Dispersal ability constrained by local site filters related to e.g. natural disturbances, microclimatic Interspecific interactions factors and potential interspecific interactions. Local community Fig.1.1: Simplified diagram of community assembly processes (modified after Morin 1999) Few of the major controversies in ecology, if any, have been
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