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Thesis Fulltext.Pdf (7.003Mb) MATING STRATEGIES AND SPERM COMPETITION IN NEW ZEALAND GECKOS (FAMILY GEKKONIDAE) A thesis submitted in fulfilment of the requirements for the Degree of Doctor of Philosophy in Zoology in the University of Canterbury by Amanda C. Todd University of Canterbury 2003 Contents ○○○○○○○○○○○○○○○○○○○○○○ Abstract○○○○○○○○○○○○○○○○○○○ 1 ○○○○○○○○○○○○○○○○ Chapter 1: General Introduction○○○○○○○○○○○ 2 ○○○○○ Taxonomy of New Zealand Geckos ○○○○○○○○○○○○○○○○ 6 Chapter 2: Testis Size as a Predictor of Mating System ○○○○○○○○○○○○○ 7 ○○○○○○○○○ Introduction○○○○○○○○○○○○○○○○○○○○○○○○○○ 7 Methods○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 10 Results ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 14 Discussion ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 22 ○○○○○○○○○○○○○○○○○○○○○○○○○○○ Conclusions ○○○○○○○○ 26 Chapter 3: Male Reproductive Characteristics and Sperm Competition ○○○ 27 ○○○○○○○○○○○○○○○○○○○○○○○○○○○○ Introduction ○○○○○○○ 27 ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ Methods ○○○○○○ 31 ○○○○○○○○○○○○○○○○○○○○○○○○○○○○ Results ○○○○○○○○○○ 41 ○○○○○○○○○○○ Discussion ○○○○○○○○○○○○○○○○○○○○○○○○○ 48 ○○○○○○○○○○○○○○○○○○○○○○○○○○○ Conclusions ○○○○○○○○ 51 Chapter 4: Female Reproductive Morphology and Sperm Competition ○○○○ 52 ○○○○○○○○○○○○○○○○○○○○○○○○○○○○ Introduction ○○○○○○○ 52 ○○○○○○○○○○○○○○○○○○○○ Methods ○○○○○○○○○○○○○○○○○ 55 Results ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 58 ○○○○○○○○○○○ Discussion ○○○○○○○○○○○○○○○○○○○○○○○○○ 67 ○○○○○○○○○○○○○○○○○○○○○○○○○○○ Conclusions ○○○○○○○○ 72 Chapter 5: The Social Mating System of Hoplodactylus maculatus ○○○○○○ 73 ○○○○○○○○○○○○○○○○○○○○○○○○○○○○ Introduction ○○○○○○○ 73 ○○○○○○○○○○○○○○○○○○○○ Methods ○○○○○○○○○○○○○○○○○ 75 Results ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 82 Discussion ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 94 ○○○○○○○○○○○○○○○○○○○○○○○○○○○ Conclusions ○○○○○○○○ 98 ○○○○○○ Chapter 6: Sperm Morphology of Hoplodactylus maculatus ○○○○ 99 Introduction ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 99 ○○○○○○○○○○○○○○○○○○○○○○○ Methods ○○○○○○○○○○○○○○ 100 ○○○○○○○○○ Results ○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 102 ○○○○○○○○○○○○○○○ Discussion ○○○○○○○○○○○○○○○○○○○○ 107 Chapter 7: Effect of Temperature on Sperm Motility in ○○○○○○○○○○○○○○○○○○○○○○ Hoplodactylus maculatus ○○○○○○ 111 Introduction ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 111 ○○○○○○○○○○○○○○○○○○○○○○○ Methods ○○○○○○○○○○○○○○ 113 ○○○○○○○○○ Results ○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 116 ○○○○○○○○○○○○○○○ Discussion ○○○○○○○○○○○○○○○○○○○○ 121 ○○○○○○○○○○○○○○○○○○○○○○○○○○○ Conclusions ○○○○○○○○ 125 ○○○○○○○○○○○○○○○○○○○○○ Chapter 8: General Discussion ○○○○○○ 126 ○○○○○○○○○○○○○○○○○○○ Future Research ○○○○○○○○○○○○○ 131 Acknowledgements ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 134 ○○○○○○○○○○○○○○○○○ References ○○○○○○○○○○○○○○○○○○○○○○ 136 ○○○○○○○○○○○○○○○○○○○○○○○○○○○○ Appendix A ○○○○○○○○○○ 175 ○○○○○○○○○○○○○○○○○○○○○○○○○○○ Appendix B ○○○○○○○○○○○ 182 List of Figures Figure Page 2.1 Relationship between left and right testis volume in New Zealand geckos 14 2.2 Variation in residual testis volume between months for Hoplodactylus ○○○○○○○○○○○○○○○○ maculatus from Turakirae Head ○○○○○ 15 2.3 Relationship between mean testis volume and SVL in New ○○○○○○○○○○○○○○○○○○○○○ Zealand geckos ○○○○○○○○○○ 16 2.4 Relationship between mean testis volume and SVL across species of ○○○○○○○○○○○○○○○○○○○○○ lizards and snakes ○○○○○○○○ 16 ○○○○○○○○○○○○○ 2.5 Relative testis size of lizards and snakes ○○○○○○ 18 3.1 Diagram of inverted hemipenis showing the location of measurements 32 3.2 SEM micrographs of hemipenes of Hoplodactylus maculatus Canterbury: A. Covered with mucus; B. With mucus removed 35 3.3 SEM micrographs of hemipenes of Hoplodactylus maculatus prepared using: A. Potassium hydroxide - good condition; B. No treatment; C. Potassium hydroxide - damaged; D. Saponin ○○○○○○○○○○○○ 37 3.4 Relationship between everted hemipenis length and mean testis volume residual in New Zealand geckos ○○○○○○○○○○○○○○○○○○○○○ 42 3.5 Relationship between residual testis volume, hemipenis size and season in New Zealand geckos: A. Length; B. Width ○○○○○○○○○○○○○ 43 3.6 SEM micrographs of hemipenis of Hoplodactylus maculatus: A. Sulcal surface; B. Asulcal surface ○○○○○○○○○○○○○○○○○○ 45 3.7 SEM micrograph of calyces covering hemipenis showing the location ○○○○○○○○○○○○ of measurements ○○○○○○○○○○○○○○○○○○ 45 3.8 Sperm from two New Zealand gecko species: A. Hoplodactylus pacificus; B. H. maculatus Canterbury ○○○○ 46 3.9 Relationship between mean testis volume residual and sperm length in ○○○○○○○ New Zealand geckos ○○○○○○○○○○○○○○○○○○○○○ 48 4.1 Longitudinal dissection of oviduct of Naultinus elegans punctatus ○○○ 56 4.2 SSTs in the uterine tube of Hoplodactylus maculatus: ○○○○○○○○○○ A. Whole-mount; B. Histological section ○○○○○○ 59 4.3 Transverse section of the vagina of Hoplodactylus maculatus ○○○○○○ 60 4.4 Relationship between number of folds in the vagina and mean relative testis volume in New Zealand geckos during: A. Sperm storage season; B. Non-sperm storage season ○○○○○○○ 65 5.1 Study site at Devils Knob, Banks Peninsula: A. Location map; B. Photograph ○○○○○○○○○○○○○○○○○○○○○ 76 ○○○○○○○ 5.2 Hoplodactylus maculatus ○○○○○○○○○○○○○○○○○○○○○ 77 5.3 Diagram of enclosure design during: A. 2001; B. 2002 ○○○○○○○○○○ 80 5.4 Mean number of Hoplodactylus maculatus captured across months ○○○ 82 ○○○○○○○○ 5.5 Size distribution of Hoplodactylus maculatus ○○○○○○○○ 84 5.6 Mean temperature during 2001 and 2002: A. Maximum; B. Minimum 85 5.7 Mean number of courtships/h across months for ○○○○○○○○○○○○○○ Hoplodactylus maculatus ○○○○○○○○○○○ 86 5.8 Relationship between SVL size rank and mean courtship rate in male Hoplodactylus maculatus during: A. 2001; B. 2002 ○○○○○○○○○○ 88 5.9 Relationship between total body length size rank and mean courtship rate in male Hoplodactylus maculatus during: A. 2001; B. 2002 ○○○○○ 88 5.10 Relationship between total body length size rank and mean courtship rate in female Hoplodactylus maculatus during: A. 2001; B. 2002 ○○○ 89 5.11 Temporal variation in mean fight rate of Hoplodactylus maculatus during: A. 2001; B. 2002 ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 91 5.12 Relationship between time of day and mean fight rate in Hoplodactylus ○○○○○○○○○○○○○○○○○○○○ maculatus across months ○○○○○ 92 5.13 Relationship between size rank difference and mean attack rate in ○○○○○○○○○○○○○ Hoplodactylus maculatus ○○○○○○○○○○○○ 93 6.1 SEM micrographs of two sperm morphs from Hoplodactylus maculatus: A. Long; B. Short ○○○○○○○○○○○○○○○○○○○○○○○○○○○○○○ 102 6.2 Light micrographs of long and short sperm morphs from ○○○○○○○○○○○○○ Hoplodactylus maculatus ○○○○○○○○○○○○ 103 6.3 Two sperm morphs from Hoplodactylus maculatus stained with Hoescht 33342: A. Long; B. Short ○○○○○○○○○○○○○○○○○○○○ 105 6.4 Two sperm morphs from Hoplodactylus maculatus stained with ○○○○○○○○○○○ ethidium bromide: A. Long; B. Short ○○○○○○○ 106 7.1 Relationship between sperm tail beat frequency and time in samples ○○○○ from Hoplodactylus maculatus ○○○○○○○○○○○○○○○○○○ 117 7.2 Mean proportion of motile short sperm in samples from Hoplodactylus maculatus at three temperatures ○○○○○○○○○○○○○○○○○○○○○ 118 7.3 Relationship between sample number and mean proportion of motile short sperm in samples from Hoplodactylus maculatus ○○○○○○○○○○○ 118 7.4 Mean tail beat frequency of short sperm in samples from Hoplodactylus maculatus at three temperatures ○○○○○○○○○○○○○○○○○○○○○ 119 7.5 Effect of time on short sperm motility at three temperatures in samples ○○○○ from Hoplodactylus maculatus ○○○○○○○○○○○○○○○○○○ 120 List of Tables Table Page 2.1 Sex differences in SVL for New Zealand geckos ○○○○○○○○○○○○○○ 19 2.2 Sex differences in head dimensions for New Zealand geckos ○○○○○○○ 20 4.1 Intraspecific variation in number of folds and number and size of SSTs in regions of the oviduct in Hoplodactylus maculatus ○○○○○○○○ 61 4.2 Interspecific variation in number of folds and number and size of SSTs ○○○○○ in regions of the oviduct in New Zealand geckos ○○○○○○ 63 4.3 Relationship between number of folds and number and size of SSTs in the oviduct, season and mean testis volume residual ○○○○○○○○ 64 4.4 Relationship between number of folds and number and size of SSTs ○○○○○○○○○○ in the oviduct, season and sperm length ○○○○○○○ 66 7.1 Effect of temperature, sample number and time on the proportion of motile short sperm in samples from Hoplodactylus maculatus ○○○ 118 7.2 Effect of temperature, sample number and time on tail beat frequency of short sperm in samples from Hoplodactylus maculatus ○○○○○○ 119 1 Abstract Most species of reptile studied to date have polygynandrous mating systems and possess specialised sperm storage regions. Consequently, there is a high potential for sperm competition in this group. Using comparative analyses, I examined the level of sperm competition in New Zealand geckos and how this has influenced the evolution of their reproductive morphology. Across lizards and snakes, there was more than a 40-fold variation in relative testis size. New Zealand geckos fell in the middle of this range and lacked sexual dimorphism in head size, suggesting that most species have polygynandrous mating systems. I confirmed this for one species, Hoplodactylus maculatus, which is gregarious, lacks territoriality and has a courtship pattern that suggests a high level of promiscuity for both sexes. I found that hemipenis size in New Zealand geckos was positively correlated with relative testis size, suggesting that sperm competition has resulted in the evolution of larger intromittent organs. However, the surface features of the hemipenis were relatively conservative across species. Although there was no relationship between sperm length or putative sperm storage site (SST) morphology and relative testis size, species with fewer
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