The Evolution of Male and Female Reproductive Traits in Simultaneously Hermaphroditic Terrestrial Gastropods

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The Evolution of Male and Female Reproductive Traits in Simultaneously Hermaphroditic Terrestrial Gastropods THE EVOLUTION OF MALE AND FEMALE REPRODUCTIVE TRAITS IN SIMULTANEOUSLY HERMAPHRODITIC TERRESTRIAL GASTROPODS Inauguraldissertation zur Erlangung der Würde eines Doktors der Philosophie vorgelegt der Philosophisch-Naturwissenschaftlichen Fakultät der Universität Basel von Kathleen Beese aus Friedrichroda, Deutschland Basel, 2007 Genehmigt von der Philosophisch-Naturwissenschaftlichen Fakultät der Universität Basel auf Antrag von Prof. Dr. Bruno Baur PD Dr. Andreas Erhardt Basel, den 13.02.2007 Prof. Dr. Hans-Peter Hauri Dekan 3 TABLE OF CONTENTS SUMMARY..............................................................................................................................................................9 GENERAL INTRODUCTION............................................................................................................................11 SEXUAL SELECTION AND SEXUAL CONFLICT......................................................................................................11 POSTMATING CONFLICT IN HERMAPHRODITES...................................................................................................12 REPRODUCTIVE MORPHOLOGIES IN STYLOMMATOPHORAN GASTROPODS........................................................13 OUTLINE OF THE THESIS .....................................................................................................................................14 CHAPTER I – EVOLUTION OF FEMALE SPERM STORAGE ORGANS IN THE CARREFOUR OF STYLOMMATOPHORAN GASTROPODS .............................................................................................17 ABSTRACT...........................................................................................................................................................17 INTRODUCTION ...................................................................................................................................................18 MATERIALS AND METHODS ................................................................................................................................21 Sampling and carrefour complexity .............................................................................................................21 Reproductive characters ...............................................................................................................................21 Life history and habitat specificity ...............................................................................................................22 Carrefour length and sperm length ..............................................................................................................23 Molecular data and phylogenetic analyses..................................................................................................24 Ancestral character states and character association.................................................................................25 RESULTS..............................................................................................................................................................28 Phylogeny ......................................................................................................................................................29 Carrefour complexity and spermatheca evolution.......................................................................................28 Carrefour associations with reproductive characters .................................................................................30 Associations with life history and habitat specificity...................................................................................30 Variation in carrefour length and sperm length ..........................................................................................31 DISCUSSION.........................................................................................................................................................33 Reproductive characters and postcopulatory sexual selection ...................................................................34 Influence of life history and habitat specificity............................................................................................36 Carrefour size and sperm length evolution..................................................................................................36 Conclusions ...................................................................................................................................................38 ACKNOWLEDGEMENTS .......................................................................................................................................38 APPENDIX............................................................................................................................................................39 CHAPTER II – COEVOLUTION OF MALE AND FEMALE REPRODUCTIVE TRAITS IN A SIMULTANEOUSLY HERMAPHRODITIC LAND SNAIL........................................................................43 ABSTRACT...........................................................................................................................................................43 INTRODUCTION ...................................................................................................................................................44 5 TABLE OF CONTENTS MATERIALS AND METHODS ................................................................................................................................ 46 Study organism.............................................................................................................................................. 46 General methods ........................................................................................................................................... 47 Reproductive traits........................................................................................................................................ 48 Image proceeding and statistical analyses................................................................................................... 50 RESULTS.............................................................................................................................................................. 50 Intraspecific variation in reproductive traits............................................................................................... 50 Allometric relationships................................................................................................................................ 51 Reproductive character associations ........................................................................................................... 52 DISCUSSION ........................................................................................................................................................ 53 ACKNOWLEDGEMENTS ....................................................................................................................................... 56 CHAPTER III – EXPANDABLE SPERMATHECA INFLUENCES SPERM STORAGE IN THE SIMULTANEOUSLY HERMAPHRODITIC LAND SNAIL ARIANTA ARBUSTORUM........................ 57 ABSTRACT........................................................................................................................................................... 57 INTRODUCTION ................................................................................................................................................... 58 MATERIALS AND METHODS ................................................................................................................................ 60 RESULTS.............................................................................................................................................................. 63 Spermatheca size in relation to sperm storage ............................................................................................ 63 Spermatophore volume and sperm number.................................................................................................. 65 DISCUSSION ........................................................................................................................................................ 66 ACKNOWLEDGEMENTS ....................................................................................................................................... 69 CHAPTER IV – BURSA TRACT DIVERTICULUM IN THE HERMAPHRODITIC LAND SNAIL ARIANTA ARBUSTORUM (STYLOMMATOPHORA: HELICIDAE): MORPHOLOGY, FUNCTION, AND EVOLUTIONARY IMPLICATIONS.............................................................................. 71 ABSTRACT........................................................................................................................................................... 71 INTRODUCTION ................................................................................................................................................... 72 MATERIALS AND METHODS................................................................................................................................ 74 Animals .......................................................................................................................................................... 74 Histology and morphometry ........................................................................................................................
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