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FEMALE PERSPECTIVES on BREEDING PHENOLOGY in EASTERN ONTARIO GRAY TREEFROGS by HAYLEY JENNIFER ROBERTS a Thesis Submitted To

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FEMALE PERSPECTIVES on BREEDING PHENOLOGY in EASTERN ONTARIO GRAY TREEFROGS by HAYLEY JENNIFER ROBERTS a Thesis Submitted To FEMALE PERSPECTIVES ON BREEDING PHENOLOGY IN EASTERN ONTARIO GRAY TREEFROGS by HAYLEY JENNIFER ROBERTS A thesis submitted to the Department of Biology in conformity with the requirements for the degree of Master of Science Queen’s University Kingston, Ontario, Canada (September 2015) Copyright © Hayley Jennifer Roberts, 2015 This work is licensed under a Creative Commons Attribution 4.0 International License Abstract Studies of the forces that shape local adaptation contribute to our understanding of origins of range-wide variation, speciation, and potential responses to changing environments. Phenology is an important component of such local adaptation. Phenological research is central to studies of geographical range limits and species distributions, migration, speciation, response to climate change and ex situ conservation and re-introduction programs. Despite over 22,000 phenological studies published thus far, there is a lack of phenological information on reproductive patterns of temperate anurans, particularly for females. My thesis therefore focuses on gaining insights into temperate frog phenology primarily of females, and patterns of female choice. I focus on Hyla versicolor (Gray Treefrogs), an Eastern North American temperate frog. My data come from perimeter transect surveys done at two wetlands at the Queen’s University Biological Station over the entire 2014 breeding season. I compared male and female peak abundance using an N- mixture model. Both males and females peaked on Julian Day 154 (June 3rd 2014). Females arrived three days after males in my focal wetlands, and left three days before. Timing of arrival is most likely based on a combination of females responding to male advertisement calls, and both sexes responding to abiotic factors (e.g. temperature, precipitation) across the season. Additionally, I looked at trends in male size over the season and found a weak positive relationship between male size and Julian day. However, females showed no preference for larger males, implying that female choice may focus on other things like genetic quality. I did find evidence of positive assortative mating, perhaps because of biophysical limitations of small females carrying larger males during oviposition. A fuller knowledge of female phenology enhances our understanding of local adaptation, and potentially provides insights into speciation and responses to climate change. ii Co-Authorship Dr. Stephen Lougheed This thesis was formatted in the manuscript format as outlined in the guidelines provided by the Department of Biology. Both data chapters are co-authored by Dr. Stephen C. Lougheed, who contributed financially and intellectually to the design and development as well as the editing of the thesis. Dr. Marc J. Mazerolle Both data chapters are co-authored by Dr. Marc J. Mazerolle, who contributed intellectually to the design and development as well as the editing of the thesis. Publications arising from and included in this thesis: Roberts, H. J., M. J. Mazerolle, and S. C. Lougheed. Phenology of male and female arrival and peak abundance in Gray Treefrogs, Hyla versicolor. Will be submitted to Copeia. Roberts, H. J., M. J. Mazerolle, and S. C. Lougheed. Female preference and temporal trends in male size within Gray Treefrogs, Hyla versicolor. Will be submitted to the Canadian Journal of Zoology. iii Acknowledgements Firstly, I would like to thank my supervisor Dr. Stephen C. Lougheed for all his help, encouragement and assistance for the past two years of completing my Masters. It has been an amazing experience being part of the Lougheed lab, and I will never forget what I have learned here. I would additionally like to thank my committee members Laurene Ratcliffe, Shelley Arnott and Marc Mazerolle for all of their incredible input in helping me to create my thesis. There are many people who helped make my thesis a possibility. Firstly, the Queen’s University Biological Station and its staff were vital in providing me with the ability to focus entirely on my research for my field season. I specifically would like to thank Veronica for accommodating my nocturnal schedule. I would also like to thank my SWEP student Amanda Cicchino for assisting me throughout my entire field season during long humid nights searching for frogs. I had many other field assistants who helped me at night including: Steven Davis, Melissa Bredow, Jackie Leonard, Amelia Cox, Carmen Gemmell, Olivia Gemmell, Oliva Gowen, Erin Kelly, Joanna Znotins, Henry Xie, and Emma Gandy. Additionally, I would like to thank the other members of the Lougheed lab for their knowledge and assistance. For helping me to make my box traps I thank Tice and Ron, as well as the entire class of Biol 307 for helping me to enact one of my drift fences. Financial support was provided by the Queen’s University Graduate Award as well as the Natural Science and Engineering Research Council (NSERC) for a Canadian Discovery Grant awarded to Dr. S.C. Lougheed. Finally, I would like to thank my family and friends, for without their encouragement and support this Masters could not have been completed. Specifically to my parents, Valya Roberts iv and Steve Roberts for not only coming to help me transect at night, but also for helping me make my traps, and giving me the emotional support I required to finish my Master’s. v Table of Contents Abstract .......................................................................................................................................... ii Co-Authorship .............................................................................................................................. iii Acknowledgements ...................................................................................................................... iv Table of Contents ......................................................................................................................... vi List of Tables ................................................................................................................................ ix List of Figures ................................................................................................................................ x Chapter 1 General Introduction .................................................................................................. 1 1.1 Phenology Overview ............................................................................................................. 1 1.2 Range Limits and Species Distributions ............................................................................... 1 1.3 Migration............................................................................................................................... 3 1.4 Ex-situ Conservation and Phenology .................................................................................... 4 1.5 Speciation .............................................................................................................................. 5 1.6 Climate Change ..................................................................................................................... 5 1.7 Breeding Phenology in Frogs and Differing Strategies for Males and Females ................... 8 1.8 Phenology Patterns of Frogs of Eastern North America ..................................................... 10 1.9 Study Species ...................................................................................................................... 12 1.10 Thesis Objectives .............................................................................................................. 13 1.11 Literature Cited ................................................................................................................. 14 Chapter 2 Phenology of male and female arrival and peak abundance in Gray Treefrogs, Hyla versicolor. ............................................................................................................................ 32 2.1 Abstract ............................................................................................................................... 32 2.2 Introduction ......................................................................................................................... 33 vi 2.3 Materials and Methods ........................................................................................................ 36 2.3.1 Study Sites .................................................................................................................... 36 2.3.2 Data Collection ............................................................................................................ 36 2.3.3 Data analysis ............................................................................................................... 37 2.4 Results ................................................................................................................................. 38 2.4.1 Frog counts .................................................................................................................. 38 2.4.2 Frog abundance ........................................................................................................... 39 2.5 Discussion ........................................................................................................................... 40 2.6 Literature Cited ................................................................................................................... 45 Chapter
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