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A 5 3 G Aoik MULTIPLE PATERNITY: A LIFE-HISTORY STRATEGY FOR PORCELAIN CRABS IN THE FACE OF RISING TEMPERATURES A Thesis submitted to the faculty of San Francisco State University In partial fulfillment of the requirements for A 5 the Degree 3 G a o ik Master of Science In Biology: Marine Biology by Thomas Joseph Yockachonis San Francisco, California May 2016 Copyright by Thomas Joseph Yockachonis 2016 CERTIFICATION OF APPROVAL I certify that I have read MULTIPLE PATERNITY: A LIFE-HISTORY STRATEGY FOR PORCELAIN CRABS IN THE FACE OF RISING TEMPERATURES by Thomas Joseph Yockachonis, and that in my opinion this work meets the criteria for approving a thesis submitted in partial fulfillment of the requirement for the degree Master of Science in Biology: Marine Biology at San Francisco State University. Jon i.D. Pro Eri Professor of Biology Frank Cipriano, Ph.D Professor of Biology MULTIPLE PATERNITY: A LIFE-HISTORY STRATEGY FOR PORCELAIN CRABS IN THE FACE OF RISING TEMPERATURES Thomas Joseph Yockachonis San Francisco, California 2016 Multiple paternity is widespread across many taxa including birds, insects and marine species. Despite multiple paternity being rare among studied crustaceans, a study found that -93% of female porcelain crabs, Petrolisthes cinctipes, mate with more than one male and produce broods of mixed paternity. No explanation has been given to the reason of why multiple paternity is prolific in P. cinctipes, or what function polyandry may serve for crabs in general to my knowledge. In order to identify potential advantages of multiple paternity in P. cinctipes, brood survival differentials were measured under ambient conditions and after a heat-shock. Microsatellite profiling was used to distinguish multiple from single paternity and compared to brood survival, in the presence or absence of a heat- shock. When exposed to a heat-shock, single-sired broods experience a significant drop in mean brood hatching %, compared to multiply-sired broods. Multiply sired broods also show a substantial, but non-statistically significant, mean difference in mean hatching % between conditions, compared to single-sired broods. Results suggest that multiple paternity reduces embryo survival variance between ambient and heat-shock conditions and that multiple mating could be an advantage for P. cinctipes in the high-intertidal zone. I certify 1 correct representation of the content of this thesis. Chair, Thesis Committee Date ACKNOWLEDGEMENTS I would like to thank my advisor, Jonathon Stillman, for giving me the opportunity to fulfill one of the greatest accomplishments of my life thus far. He has provided me the freedom to explore my research interests and his guidance has enabled me to develop my scientific skill-set. Because of his efforts, I have been able to obtain the career I have been seeking for the past seven years. I would also like to thank my supporting thesis committee members, Eric Routman and Frank Cipriano, for their guidance and direction in editing this document. My lab mates Eric, Lindsay, Emily K., Jennifer and lab-tech Adam of course deserve much appreciation for their stable humor and help when I could not come to RTC. I want to express immense gratitude to Seabird McKeon, the most important mentor of my college career. Thank you first of all for funding the vast majority of my graduate research. Also, thank you for allowing me assist on expeditions to Belize when I needed it most and for supporting me academically for the past five years. I would also like to thank my mentors at the Smithsonian, Amanda Windsor and Michele Weber. 1 appreciate you helping me develop my molecular skills, design an experimental approach and for your friendship. 1 also need to thank a fellow Navy EOD Tech and friend, Ike Kanakanui. Our adventures across Sicily by car and many islands aboard “The Freedom Boat” are some of my most valued memories, and taught me to follow my passion. Lastly, I must thank my soon-to-be wife, Katrina Eichner, for her love, patience and loyalty throughout my time in college. You are my biggest source of personal motivation to better myself in every way. Without you, I most likely would have never finished grad-school and 1 would be living in a remote part of a foreign country with no shoes or professional direction. v TABLE OF CONTENTS List of Tables............................................................................................................................viii List of Figures............................................... ix List of Appendices....................................................................................................................... x 1.0 Introduction............................................................................................................................1 2.0 Materials and Methods......................................................................................................... 14 2.1 Porcelain Crab Collection and Heat-shock Treatment ............................................. 14 2.2 Identification of Multiple Paternity.......................................................................... 16 2.2.1 Embryo Preservation................................................................................ 16 2.2.2 Tissue Digestion & DNA Extraction.........................................................16 2.2.3 PCR Amplification................................................................................... 17 2.2.4 Fragment Analysis.................................................................................... 18 2.2.5 Quantifying paternity............................................................................... 19 2.3 Statistical Analysis..................................................................................................20 3.0 Results..................................................... 22 3.1 Control hatching % by brood...............................................................................22 3.2 Heat-shock Effect....................................................................................................23 3.3 Overall brood hatching %: MP vs. SP................................................................ 24 3.4 Mean overall hatching %: MP vs. SP ................................................................ 25 3.5 Mean Proportional hatchings by treatment: MP vs. S P ....................................26 3.6 Brood Hatching % Differences (NHS-HS)........................................................ 27 3.7 Mean Differences in Mean Hatching % ............................................................... 28 4.0 Discussion 29 4.1 Heat-shock Effect....................................................................................................29 4.2 Effect of Multiple Paternity: Ambient and HS Conditions................................ 30 4.3 Multiple Paternity and Survival Variance............................................................31 4.4 Conclusions.............................................................................................................32 4.5 Future Directions.....................................................................................................32 References.................................................................................................................................34 Appendices................................................................................................................................ 38 vii LIST OF TABLES Table 1. Primer sequences: Loci Pc 156s and Pc 170s LIST OF FIGURES Figure Page 1. Low tide habitat temperatures of P. cinctipes and P. eriomerus........................................ 5 2. Thermal survival of P. cinctipes and P. eriomerus........................................................... 6 3. Multiple mating: Geometric mean and survival..............................................................10 4. Multiple mating: Variance and direct fitness gains.......................................................... 12 5. No heat-shock hatching % by brood.............................................................................. 21 6. Heat-shock effect of mean brood hatching %................................................................. 22 7. Overall survival by brood: Multiple vs. Single Paternity................................................23 8. Mean brood overall survival: Multiple vs. Single Paternity............................................24 9. Mean hatching % by heat-shock treatment: Multiple vs. Single Paternity.......................25 10. Brood survival differences between heat-shock treatments.............................................26 11. Mean brood survival differences: Multiple vs. Single Paternity......................................27 LIST OF APPENDICES Appendix 1. Confirmed multiple paternity through fragment analysis 1 1.0 Introduction Stress variation occurs naturally in the earth’s climate system over a range of temporal scales, such as annually, seasonally and daily. This stress variability is reflected in species’ present evolutionary adaptations and can be observed through small and large scale patterns of biogeography (Harley et al. 2006) and physiological response under a shifting physical stress, such as temperature (Helmuth and Hofmann 2001). An thermal stress is a temperature that when changed, alters organismal physiological processes and causes a clear drop in reproductive yield (Hoffmann and Hercus 2000). For a species to have adaptive potential to thermal
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