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Personality in the Brush-Legged Wolf Spider: Behavioral Syndromes and Their Effects on Mating Success in Schizocosa Ocreata

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Personality in the Brush-Legged Wolf Spider: Behavioral Syndromes and Their Effects on Mating Success in Schizocosa Ocreata Personality in the Brush-legged Wolf Spider: Behavioral Syndromes and their Effects on Mating Success in Schizocosa ocreata A thesis submitted to the Graduate School of the University of Cincinnati in partial fulfillment of the requirements for the degree of Master of Science in the Department of Biological Sciences in the College of Arts and Sciences by Trinity D. Walls B.A. Washington University in Saint Louis July 2018 Committee Chair: George W. Uetz, Ph.D. Cincinnati, OH Abstract Recent studies have shown that animal “personality” demonstrates consistent behavioral variation at the individual level that persists across lifestages and contexts. The most commonly measured behavioral syndrome involves a “bold” to “shy” continuum, in which individuals are evaluated based on their willingness or latency to engage in risk-taking behaviors. I examined bold-shy behavioral syndromes in the brush-legged wolf spider, Schizocosa ocreata. Spiders were repeatedly given open field tests and later exposed to simulated predator stimuli. All spiders were tested as juveniles and adults. Results of open field tests showed individual S. ocreata exhibit consistent behavioral patterns associated with either end of the continuum of bold (exploratory) to shy (freeze) behavioral syndromes. These differences persisted across contexts, as well as lifestages (juvenile, adult). Bold spiders exhibited shorter latency to explore in an open field and to resume exploration after a simulated predator than did their shy counterparts, but also showed more variation in latency to resume exploration after a simulated predator. After reaching maturity, females were given a two-choice test using video playback of male courtship to analyze differences in mate choice, while males were exposed to female cues to assess courtship vigor. While open field behaviors and responses to simulated predators were correlated, personality type did not show significant effects on male courtship in the presence of female cues or female mate preference in the context of video playback. Males and females of differing personality types were also paired in a two-by-two factorial design to assess the effect of personality on overall mating success. No differences in mating success were found, suggesting that personality type measured using bold-shy attributes may affect somatic traits but not reproductive traits in this study. ii iii Acknowledgements I would like to thank the University of Cincinnati and the Yates Fellowship program for providing financial support and space to conduct this research. I thank my advisor, Dr. George Uetz, for providing indispensable guidance and direction for the last two years, as well as for his encouragement and confidence throughout this process. I thank the other members of my research advisory committee, Dr. Elke Buschbeck and Dr. Nathan Morehouse, who were also fundamental in helping me revise and improve my ideas and project designs. I thank the post- doctoral researchers Dr. Brent Stoffer and Dr. Alex Sweger for their patience and constructive feedback during my time here. I also thank my lab members, Emily Pickett, Madeline Lallo, and Tim Meyer for their encouragement and advice, and the numerous undergraduates who performed animal husbandry. Lastly, I thank the spiders for being complex enough to display distinct personalities and making this research possible. iv Table of Contents Abstract…………………………………………………………………………………………. ii Acknowledgments …………………………………………………………………....……........ iv List of Figures……………………………………………….………………………………….. vii Introduction………………………………………………………………………………………..1 Methods…………………………………………………………………………………………....6 Collection and Care.………..………………………………………………….……….......7 Establishing Personality………………………………………………………….……......8 Open Field Test……………………...........................................................................8 Correlating Personalities with Behaviors in Other Contexts …………………………......9 Simulated Predator Stimulus Test……..…………………………………………..9 Female Mate Choice………………………………………………………………………9 Male Courtship……………………………...……………………………………………10 Live Mating Trials of Different Personalities…………………………………………10 Differences Between Research Seasons………………………………………………………...11 Statistical Analyses…………………………………….…………………………………11 Results……………………………………………………………………………………………13 Part I: Lab-reared spiders in an open field.......................................................................13 Part II: Lab-reared spiders’ responses to a simulated predator stimulus………………..14 Part III: Field-exposed spiders in an open field.…………..…………………………….15 Part IV: Field-exposed spiders’ responses to a simulated predator stimulus………..…..15 Part V: Lab-reared vs field-exposed ……………………………………………………..15 Part VI: Female responses to a two-choice test.………………………………................16 Part VII: Male courtship on female silk.…………………………………………………17 Part VIII: Mating personalities together..………………………………………………..17 Part IX: Lab-reared spiders in extended open field test.………………………………...18 Part X: Lab-reared spiders’ responses to a simulated predator stimulus…………………18 Discussion……………………….……………………………………………………………….19 Future Directions.………………………………………………………………………………..25 References……………………………………………………………………………………….29 v Figures…………………………………………………………………………………………...35 vi List of Tables and Figures Figure 1: Lab-reared Penultimate Latency to Explore in Open Field across Trials Figure 2: Lab-reared Mature Latency to Explore in Open Field across Trials Figure 3: Distribution of Initial Open Field Responses in Lab-reared Penultimate Females Figure 4: Distribution of Initial Open Field Responses in Lab-reared Penultimate Males Figure 5: Distribution of Initial Open Field Responses in Lab-reared Mature Females Figure 6: Distribution of Initial Open Field Responses in Lab-reared Mature Males Figure 7: Lab-reared Penultimate Female Latency to Recover from Predator Stimulus Figure 8: Lab-reared Penultimate Male Latency to Recover from Predator Stimulus Figure 9: Correlation between Open Field Latency and Predator Stimulus Latency in Lab-reared Penultimates Figure 10: Lab-reared Mature Female Latency to Recover from Predator Stimulus Figure 11: Lab-reared Mature Male Latency to Recover from Predator Stimulus Figure 12: Correlation between Open Field Latency and Predator Stimulus Latency in Lab-reared Matures Figure 13: Field-exposed Penultimate Latency to Explore in Open Field across Trials Figure 14: Field-exposed Mature Latency to Explore in Open Field across Trials Figure 15: Distribution of Initial Open Field Responses in Field-exposed Penultimate Females Figure 16: Distribution of Initial Open Field Responses in Field-exposed Penultimate Males Figure 17: Distribution of Initial Open Field Responses in Field-exposed Mature Females Figure 18: Distribution of Initial Open Field Responses in Field-exposed Mature Males Figure 19: Field-exposed Penultimate Female Latency to Recover from Predator Stimulus Figure 20: Field-exposed Penultimate Male Latency to Recover from Predator Stimulus Figure 21: Correlation between Open Field Latency and Predator Stimulus Latency in Field- exposed Penultimates vii Figure 22: Field-exposed Mature Female Latency to Recover from Predator Stimulus Figure 23: Field-exposed Mature Male Latency to Recover from Predator Stimulus Figure 24: Correlation between Open Field Latency and Predator Stimulus Latency in Field- exposed Matures Figure 25: Lab-reared Mature Female Latency to Recover from a Predator Stimulus using Boldness Scores Figure 26: Lab-reared Mature Male Latency to Recover from a Predator Stimulus using Boldness Scores viii Introduction Behavioral variation and its drivers have been the subject of considerable research, particularly within lower animal taxonomic levels. Changes in animal behavior have been attributed to weather, age, habitat, food abundance, predator proximity, resource abundance, breeding season, etc. (Relyea 2001), although responses may vary among taxa. Ecotypic behavioral variation between populations of the same species is also found, e.g., coastal garter snakes accepting slugs as prey while inland garter snakes reject them (Arnold 1981), and differences in attack latency values for the same prey between grassland and riparian spider populations (Hedrick and Riechert 1989). However, even more specifically and in some instances surprisingly, variation between individuals in the same population has also been recently shown to have a sizable influence on animal responses. Over the last few decades, ethologists and behavioral ecologists have begun addressing a previously understudied facet of animal behavior that may have substantial impact on animal responses in different contexts: behavioral syndromes and “personality” (Gosling 2001; Réale et. al. 2007). In the past, personality has primarily been considered in humans and other complex animals with large mental capacities (McGuire et. al 1994; Capitanio 1999) because such traits are generally thought of as arising from characteristic patterns of thinking, behaving, and feeling (Pervin and John 1997) that have been shaped through individual experience. While this may be true, animal personality has been defined more simply as consistent behavioral differences between individuals that persist across time and contexts (Dall, Houston, and McNamara, 2004; Kralj-Fišer and Schuett, 2014). This simplified definition allows for personalities to be quantified in less complex animals and also allows for consistent individual variation to be examined rather than average population
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