How to predict the future? [Elektronische Ressource] : on niches and potential distributions of amphibians and reptiles in a changing climate / vorgelegt von Dennis Rödder

How to predict the future? [Elektronische Ressource] : on niches and potential distributions of amphibians and reptiles in a changing climate / vorgelegt von Dennis Rödder

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How to predict the future? On niches and potential distributions of amphibians and reptiles in a changing climate Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrichs-Wilhelms-Universität Bonn vorgelegt von DENNIS RÖDDER aus Troisdorf Bonn, 2009 Angefertigt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn Erstgutachter: Prof. Dr. Wolfgang Böhme Zweitgutachter: Prof. Dr. Johann-Wolfgang Wägele Tag der mündlichen Prüfung: 21.10.2009 Diese Dissertation ist auf dem Hochschulschriftenserver der ULB Bonn http://hss.ulb.uni-bonn.de/diss_online elektronisch publiziert. 2 Rheinische Friedrich-Wilhelms Universität Zoologisches Forschungsmuseum Alexander Koenig Sektion Herpetologie Adenauerallee 160 53113 Bonn Dipl. Biologe Dennis Rödder Bonn, den 15.05.2009 ERKLÄRUNG Hiermit erkläre ich an Eides statt, dass ich für meine Promotion keine anderen als die angegebenen Hilfsmittel benutzt habe, und dass die inhaltlich und wörtlich aus anderen Werken entnommenen Stellen und Zitate als solche gekennzeichnet sind. _____________ Dennis Rödder 3 “Geographic distributional areas are the shadows produced by taxa on the geographical screen.

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How to predict the future? On niches and potential
distributions of amphibians and reptiles in a changing climate








Dissertation
zur
Erlangung des Doktorgrades (Dr. rer. nat.)
der
Mathematisch-Naturwissenschaftlichen Fakultät
der
Rheinischen Friedrichs-Wilhelms-Universität Bonn
vorgelegt von
DENNIS RÖDDER
aus
Troisdorf
Bonn, 2009
Angefertigt mit Genehmigung der Mathematisch-Naturwissenschaftlichen
Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn
















Erstgutachter: Prof. Dr. Wolfgang Böhme
Zweitgutachter: Prof. Dr. Johann-Wolfgang Wägele
Tag der mündlichen Prüfung: 21.10.2009


Diese Dissertation ist auf dem Hochschulschriftenserver der ULB Bonn
http://hss.ulb.uni-bonn.de/diss_online elektronisch publiziert.
2
Rheinische Friedrich-Wilhelms Universität
Zoologisches Forschungsmuseum Alexander Koenig
Sektion Herpetologie
Adenauerallee 160
53113 Bonn
Dipl. Biologe Dennis Rödder Bonn, den 15.05.2009











ERKLÄRUNG

Hiermit erkläre ich an Eides statt, dass ich für meine Promotion keine anderen als die
angegebenen Hilfsmittel benutzt habe, und dass die inhaltlich und wörtlich aus anderen
Werken entnommenen Stellen und Zitate als solche gekennzeichnet sind.



_____________
Dennis Rödder



3
“Geographic distributional areas are the shadows produced
by taxa on the geographical screen. To study them one
needs to measure ghosts”
A. RAPOPORT






“Most ecologists would agree that niche is a
central concept of ecology, even though we
do not know exactly what it means”
L. REAL and S. LEVIN






“Of course not. After all, I may be wrong.”
B. RUSSELL




4
CONTENTS
CONTENTS 5
ACKNOWLEDGEMENTS 9
AIMS AND OVERVIEW 10
SECTION 1: GENERAL INTRODUCTION 15
1.1. Species in a changing world 16
1.2 Climate niches and the spatial distribution of species 17
1.3. What is needed? 20
Species records 20
Climate data 21
1.4. How do CEMs work? 24
1.5. Evaluation of results 26
1.6 CEM applications in taxonomy and ecology: afrotropical reed frogs as examples 27
Hyperolius cinnamomeoventris sensu lato 27
Hyperolius mitchelli and H. puncticulatus 30
1.7. Strengths of CEMs and potential ‘pitfalls’ when interpreting results 32
SECTION 2: STRUCTURE OF CLIMATE NICHES 38
2.1. Translating natural history into geographic space: a macroecological perspective on the
North American Slider, Trachemys scripta (Reptilia, Cryptodira, Emydidae) 39
Introduction 40
Methods 41
Presence data points of Trachemys scripta 41
Climate data 42
Assessing the explanative power of variables 42
Results and discussion 44
Monthly temperature variation at Slider records 44
Which climatic factor best predicts the Slider's geographic range? 45
Impact of winter temperatures 49
Possible strategies for compensation of climatic variation 50
Conclusion 51
2.2. Alien invasive Slider in unpredicted habitat: a matter of niche shift or variables studied? 52
Introduction 53
Methods 56
Slider record data 56
Climate data 57
5 Contents
Selection of climate variables 57
Computation of CEM 58
Results 60
Discussion 67
Conclusions 68
2.3. Niche shift versus niche conservatism? Climatic characteristics within the native and
invasive ranges of the Mediterranean Housegecko (Hemidactylus turcicus) 70
Introduction 71
Methods 73
Studied species 73
Species records 74
Climate data 76
Climate Envelope Models 77
Results 79
Bioclimatic conditions at native and invasive records 79
Single variables: niche overlap, similarity and equivalency 79
CEMs computed with sets of variables 81
Sets of variables: niche overlap, equivalency and similarity 82
Discussion 87
Niche conservatism versus niche shift 88
Habitat selection versus background effects 89
Methodical caveats 89
Conclusions 90
SECTION 3: HOW DO ACCESSIBILITY AND BIOTIC INTERACTIONS SHAPE
REALIZED DISTRIBUTIONS? 91
3.1. Predicting the potential distributions of two alien invasive Housegeckos (Gekkonidae:
Hemidactylus frenatus, Hemidactylus mabouia) 92
Introduction 93
Material and methods 94
Species records 94
Climate data 95
Climate Envelope Models 96
Results 96
Discussion 99
Do the current ranges of the two geckos reflect equilibrium or are they still
spreading? 101
Competitive exclusion 102
Conclusions 103
3.2. ‘Sleepless in Hawaii’ – does anthropogenic climate change enhance ecological and
socioeconomic impacts of the alien invasive Eleutherodactylus coqui Thomas 1966 (Anura:
Eleutherodactylidae)? 104
Introduction 105
Material and methods 106
Climate and computation of Climate Envelope Models 106
Species records 108
Results 108
Discussion 112
6 Contents
3.3. Will future anthropogenic climate change increase the potential distribution of the alien
invasive Cuban treefrog (Anura: Hylidae)? 114
Introduction 115
Material and Methods 117
Species records 117
Climate data 117
Computation of Climate Envelope Models 118
Results 119
Discussion 122
3.4. Potential distribution of the alien invasive Brown tree snake, Boiga irregularis (Reptilia:
Colubridae) 125
Introduction 126
Material and Methods 128
Species records and climate data 128
Variable selection 129
Climate Envelope Models 129
Results 131
Current potential distribution 131
Discussion 133
Biotic interactions and/or accessibility 133
Climate 135
Which regions are at high risk of invasion by the Brown tree snake? 135
Conclusions 138
3.5. Human Footprint, facilitated jump dispersal, and the potential distribution of the
invasive Eleutherodactylus johnstonei Barbour 1914 (Anura: Eleutherodactylidae) 139
Introduction 140
Material and methods 142
Species records 142
Variable selection 142
Computation and evaluation of the Climate Envelope Model 143
Results 144
Discussion 147
Methodical caveats 148
Conclusions 149
SECTION 4: NICHE DYNAMICS IN SPACE AND TIME 150
4.1. Environmental niche plasticity of the endemic gecko Phelsuma parkeri from Pemba
Island, Tanzania: a case study of extinction risk on flat islands by climate change 151
Introduction 152
Material and methods 154
Climate data 155
Modelling algorithm 156
Results 156
Potential distribution and changes in climatic conditions 157
Discussion 160
Possible threats 160
IUCN status and potential sustainable use 163
7 Contents
4.2. Reinforcing the predictions of the disturbance vicariance hypothesis in Amazonian
harlequin frogs: a molecular phylogenetic and climatic envelope modelling approach 165
Introduction 166
Methods 169
A central Amazonian distribution gap 169
Nested monophyly of eastern Amazonian Atelopus 171
Divergence in climate envelopes and allopatry 173
Results and Discussion 175
A central Amazonian distribution gap 175
Nested monophyly of eastern Amazonian Atelopus 177
Divergence in climate envelopes and allopatry 177
Conclusions 180
5. SUMMARY 183
6. REFERENCES 191
7. APPENDIX 227
Appendix 2.2-S1 228
Appendix 2.2-S2 229
Appendix 2.3-S1 230
Appendix 2.3-S2 232
Appendix 2.3-S3 233
Appendix 4.1-S1 234
Appendix 4.2-S1 235
Appendix 4.2-S2 243
Appendix 7.1. Species records used in this thesis 246
Atelopus, eastern 246
Atelopus, western 247
Boiga irregularis, native 247
Boiga irregularis, invasive 252
Eleutherodactylus coqui, native 252
Eleutherodactylus coqui, invasive 253
Eleutherodactylus johnstonei, native 253
Eleutherodactylus johnstonei, invasive 254
Hemidactylus frenatus, native 255
Hemidactylus frenatus, invasive 259
Hemidactylus mabouia, native 261
Hemidactylus mabouia, invasive 263
Hemidactylus turcicus, native 265
Hemidactylus turcicus, invasive 279
Osteopilus septentrionalis, native 282
Osteopilus septentrionalis, invasive 283
Phelsuma parkeri 284
Trachemys scripta, native 285
Trachemys scripta, invasive 289


8
ACKNOWLEDGEMENTS


This thesis at hand is the result of two years working, exposing myself to many
intersteing and exciting - but also sometimes exhausting - analyses of the interactions
between macro-climate and distribution patterns of amphibian and reptile species. My
studies would not have been possible without the help of many people and institutions.
With few exceptions, most people I would like to thank are listed in the acknowledgements
of the corresponding chapters or listed as co-authores. Anyway, I will mention some of
them here.

First of all I would like to thank Prof. Dr. WOFLGANG BÖHME (Bonn), Dr. STEFAN
LÖTTERS (Trier), Prof. Dr. SEBASTIAN SCHMIDTLEIN (Bonn) and Prof. Dr. MICHAEL VEITH
(Trier) for numerous furitfull discussions and advises during my studies. I am indebt for
their critical reading of the chapters and their constructive criticisms during the last years.
Furthermore, all of them contributed a lot to get my thesis supported by the
‘Graduiertenförderung des Landes Nordrhein-Westfalen’.

Regarding my collaboration with colleagues of which many have become friends,
and I am grateful to Prof. Dr. WOFLGANG BÖHME (Bonn), JOHANNES DAMBACH (Bonn),
Dr. FRANK GLAW (München), OLIVER HAWLITSCHEK (München), Dr. AXEL KWET
(Stuttgart), Dr. STEFAN LÖTTERS (Trier), Prof. Dr. SEBASTIAN SCHMIDTLEIN (Bonn), Dr.
SUSANNE SCHICK (Mainz), Prof. Dr. MIRCO SOLÉ (Ilhéus), Prof. Dr. MICHAEL VEITH
(Trier) and FRANK WEINSHEIMER (Bonn) who greately improved as co-authors the design
of my thesis. JOS KIELGAST (Copenhagen) has fascilitated in numerous ways the access to
literature.

Also ‘Thank you!’ to all the people I have forgotten to mention here. Finally I would
like to thank KATHRIN GRIES (Bonn), my parents and grandparents for their great support
during the last years.


9
AIMS AND OVERVIEW


Predictions of species distributions derived from correlative models can help to
understand the spatial patterns of biodiversity and identification of possible threats for
populations caused by climate change. The amount of available data and software is
rapidly increasing as well as the number of studies applying niche model techniques.
However, a discrepancy between increasingly complex studies and the understanding of
underlying processes, derivation of valid assumptions, and the development of conceptual
backgrounds is still a problem (JIMÉNEZ-VALVERDE et al. 2008). The aim of this thesis at
hand is an assessment of the relative importance of macro-climate, biotic interactions and
accessibility shaping realized distributions of amphibian and reptile species. The results are
comprised in sections each with two or more complementary chapters linked with the
research theme but distinct in the questions elucidated. It needs to be noted that the use of
slightly different analysis methods used in this thesis mirror the improvements in
modelling techniques and the wider availability of GIS data over the period this thesis has
been conducted.


Section 1
The first section provides a general overview over the current knowledge concerning
impacts of climate change on biota, niche concepts, availability of both climate and species
occurrence data and the methods used herein. Additionally, potential ‘pitfalls’ when
applying environmental niche models or climate envelope models are highlighted,
illustrated and discussed using examples.


Section 2
The second section focuses on the structure of climate niches. Climatic variability
within species ranges and habitat choice are analysed and discussed in the context of
natural history properties and corresponding constrains on habitat choice of the respective
species. In chapter 2.1, the hypothesise that climatic requirements allowing successful egg
incubation and balanced sex ratios are the major driver for the geographic distribution of
10