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Ecology of Phasmids (Phasmatodea) in a Moist Neotropical Forest: a Study on Life History, Host-Range and Bottom-Up Versus Top-Down Regulation

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Ecology of Phasmids (Phasmatodea) in a Moist Neotropical Forest: a Study on Life History, Host-Range and Bottom-Up Versus Top-Down Regulation ECOLOGY OF PHASMIDS (PHASMATODEA) IN A MOIST NEOTROPICAL FOREST: A STUDY ON LIFE HISTORY, HOST-RANGE AND BOTTOM-UP VERSUS TOP-DOWN REGULATION Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Technischen Universität Kaiserslautern vorgelegt von Jürgen Berger Kaiserslautern, im Mai 2004 Cover: A female of Metriophasma diocles displays all its beauty (on Piper marginatum; © by the Author). ECOLOGY OF PHASMIDS (PHASMATODEA) IN A MOIST NEOTROPICAL FOREST: A STUDY ON LIFE HISTORY, HOST-RANGE AND BOTTOM-UP VERSUS TOP-DOWN REGULATION Vom Fachbereich Biologie der Technischen Universität Kaiserslautern zur Verleihung des akademischen Grades „Doktor der Naturwissenschaften“ genehmigte Dissertation D 386 vorgelegt von Dipl.-Biol. Jürgen Berger Betreuer der Dissertation: Prof. Dr. Burkhard Büdel Kaiserslautern 2004 Tag der wissenschaftlichen Aussprache: 13. Juli 2004 Vorsitzender der Prüfungskommission: Prof. Dr. Helmut Joseph Schmidt 1. Gutachter: Prof. Dr. Burkhard Büdel 2. Gutachter: PD Dr. Jürgen Kusch Phasmids, the most peculiar beasts, ‘between the Devil and the Deep Blue Sea’ (Lawton & McNeill 1979) Acknowledgements I Acknowledgements Many people and organizations contributed to the work presented in this dissertation, and I am grateful for all of their help. I thank my advisors Rainer Wirth and Burkhard Büdel and all the members of the Department of Botany at University of Kaiserslautern for years of stimulating discussion leading to the ideas presented here. I am grateful to Donald Windsor for all of his generous support during my stay in Panamá. Tanja Lakatos, Sergio Bermudo, Manuel Redling, Florian Pankewitz, Nicole Saverschek, Britta Hartard, Sebastian Meyer and Kim Beisel never got tired in field or lab work. Jennifer Lauterbach helped making my written words sound english. Gracias. I also thank all visiting scientists, assistants and volunteers at the Smithsonian Tropical Research Institute and all staff I had the unforgettable opportunity to interact with. Special thanks go to Christiane Bröker and to my family who provided emotional, mental and physical support throughout the project. – Por la vida y el amor. Funding was provided by the Smithsonian Tropical Research Institute (STRI), the Landes- graduiertenförderung Rheinland-Pfalz (LGFG) and the German Academic Exchange Board (DAAD). Table of Contents II Table of Contents Acknowledgements ..................................................................................................................... I Table of Contents .......................................................................................................................II List of Figures.............................................................................................................................V List of Tables ............................................................................................................................ VI List of Abbreviations ..............................................................................................................VII 1 Introduction..............................................................................................................................1 1.1 General aspects....................................................................................................................1 1.2 Current knowledge about tropical phasmid ecology...........................................................2 1.2.1 Design and aims of the study.....................................................................................................3 1.3 Site characterization and general methods ..........................................................................5 1.3.1 Study site, vegetation and climate .............................................................................................5 1.3.2 Maintenance of phasmid species in the laboratory....................................................................7 1.3.3 Identification of phasmid and plant species...............................................................................7 1.3.4 Data analysis..............................................................................................................................8 2 Community structure and host range of phasmids on BCI .................................................9 2.1 Introduction .........................................................................................................................9 2.2 Materials and methods.......................................................................................................11 2.2.1 Line-transects ..........................................................................................................................11 2.2.2 No-choice feeding trials...........................................................................................................12 2.2.3 Data analysis............................................................................................................................12 2.2.3.1 Estimating phasmid community parameters........................................................12 2.2.3.2 Estimating niche dimensions and overlap ...........................................................15 2.3 Results ...............................................................................................................................16 2.3.1 Phasmid diversity and density .................................................................................................16 2.3.2 Seasonality...............................................................................................................................17 2.3.3 Food niche ...............................................................................................................................19 2.4 Discussion .........................................................................................................................24 2.4.1 Phasmid diversity, density, and distribution............................................................................24 2.4.2 Seasonality...............................................................................................................................26 2.4.3 Niche breadth and habitat choice.............................................................................................27 2.4.4 Conclusions .............................................................................................................................30 Table of Contents III 3 Life cycle, potential population growth, and egg hatching failure of Metriophasma diocles .............................................................................................................31 3.1 Introduction .......................................................................................................................31 3.2 Materials and methods ......................................................................................................33 3.2.1 Assessing demographic population parameters and life history traits.................................... 33 3.2.2 Modeling potential population growth and the effect of limited hatching success................. 35 3.3 Results ...............................................................................................................................36 3.3.1 Demographic population parameters and life history traits.................................................... 36 3.3.2 Potential population growth and effects of limited hatching success ..................................... 39 3.4 Discussion .........................................................................................................................40 3.4.1 Demographic population parameters and life history traits.................................................... 40 3.4.2 The biotic potential of Metriophasma diocles ........................................................................ 41 3.4.3 A first insight into M. diocles population control................................................................... 42 3.4.4 Conclusions............................................................................................................................. 43 4 Preference of female adults and performance of larval stages of M. diocles on single food plant species............................................................................................................................44 4.1 Introduction .......................................................................................................................44 4.2 Materials and methods ......................................................................................................47 4.2.1 Measuring leaf traits ............................................................................................................... 48 4.2.1.1 Measuring leaf toughness ................................................................................... 48 4.2.1.2 Leaf phenol and tannin content........................................................................... 48 4.2.2 Dual-choice feeding trials with M. diocles adult females....................................................... 49 4.2.3 Problems inherent with dual-choice feeding trials.................................................................. 50 4.2.3.1 Conditioning due to prior feeding experience..................................................... 50 4.2.3.2 Choice of reference plant species ......................................................................
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