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Abstract Philopatry in Prairie Voles ABSTRACT PHILOPATRY IN PRAIRIE VOLES: AN EVALUATION OF THE HABITAT SATURATION HYPOTHESIS By: Kristen E. Lucia Philopatry, or delayed dispersal of sexually mature offspring, may be due to ecological constraints on dispersal. In this study I manipulated the population density of prairie voles (Microtus ochrogaster) living in experimental enclosures to test the predictions of the habitat saturation hypothesis that philopatry and subsequent group formation in this cooperatively breeding mammal is affected by the availability of suitable territories. A significantly greater proportion of offspring remained philopatric at high densities, with females being more philopatric than males at all densities. This increase in philopatry led to a significant increase in the proportion of social units that were groups as well as a significant increase in group size. These results provide the strongest evidence for a mammal of a causal affect of density on dispersal and group formation and suggest that habitat saturation is at least a partial explanation for philopatry in prairie voles. PHILOPATRY IN PRAIRIE VOLES: AN EVALUATION OF THE HABITAT SATURATION HYPOTHESIS A Thesis Submitted to the Faculty of Miami University in partial fulfillment of the requirements for the degree of Master of Science Department of Zoology By Kristen E. Lucia Miami University Oxford, Ohio 2007 Co-Advisor Reader _____________________________ Brian Keane Co-Advisor _____________________________ Nancy G. Solomon Reader _____________________________ Robert L. Schaefer TABLE OF CONTENTS List of Tables iii List of Figures iv Dedication v Acknowledgements vi Introduction: Cooperative Breeding and Philopatry 1 References 7 Chapter 1: An Evaluation of the Habitat Saturation Hypothesis 12 Introduction 12 Methods 15 Results 22 Discussion 24 References 29 Tables 35 Figures 38 General Conclusions 43 References 45 ii LIST OF TABLES Table Page 1 Vegetative data. 35 2 Prairie vole social structure. 37 iii LIST OF FIGURES Figure Page 1 Trapping schedule for study. 38 2 Mean prairie vole group size (± SE) within enclosures as a function of 39 density and year. 3 Proportion of prairie vole social units within an enclosure that were 40 groups as a function of density and year. 4 Proportion of prairie vole offspring within each enclosure that were 41 classified as philopatric during the three week residency period as a function of density. 5 Proportion of male and female prairie vole offspring within each 42 enclosure that were classified as philopatric during the three week residency period as a function of density. iv DEDICATION To my parents, Maureen and Art Lucia, who always encouraged me to follow my own path. v ACKNOWLEDGMENTS I would like to thank my advisors, Brian Keane and Nancy Solomon for their encouragement, advice, and support throughout my time at Miami. I also thank Bob Schaefer for his patient guidance through the statistical gauntlet that is animal behavior research. I would also like to thank Ann Rypstra, Rodney Kolb, and the ERC staff for providing technical support during my two field seasons. I thank Tom Crist and Todd Levine, who helped with density estimations and program MARK. I also thank the animal care staff for their aid in maintaining the prairie vole colony in Boyd. I thank Ashley Richmond, Michelle Edwards, Tony Fries, John Williams, Lisa Aschemeier, and all the vole wranglers who made this research possible. I also thank Craig Streatfeild who let me bounce countless ideas off him and was always willing to help with the residency saga. I am grossly indebted to my good friends Kathy, Jen, Jenn, Sam, Harry, Will, Andy, Janelle, Padre, and Phill, who supported me through the last three years and provided as much aid as they could (and when all else failed joined me in my insanity). I offer many thanks to Jack Cranford, who helped foster my love for mammalogy, and all those at Virginia Tech who helped mold me into the scientist I am today. I also thank my family for their love and support throughout my graduate career. I thank Maddie for her love and for putting up with my long days at school and in the field and all my friends who have if nothing else put up with the tiresome excuse of “I can’t ______, I’m working on my thesis.” The research was financially supported by Miami University Summer Workshop fund, managed by Dave Berg, the National Science Foundation, and the American Society for Mammalogists Grants-in-Aid fund. vi INTRODUCTION: COOPERATIVE BREEDING AND PHILOPATRY Cooperative breeding is a social system typically defined by three characteristics: philopatry (delayed dispersal of offspring), reproductive suppression (when mature offspring do not reproduce), and provision of care to offspring that are not one’s own, commonly referred to as alloparental care (Solomon and French 1997). Cooperative breeding is found across the animal kingdom in species of insects and fish as well as birds and mammals (Koenig and Dickinson 2004; Queller 1994; Russell 2004; Taborsky and Limberger 1981). While occurring in only around three percent of bird and mammalian species (Jennions and Macdonald 1994; Koenig and Dickinson 2004; Russell 2004) the distribution of cooperative breeding is not random, but is instead concentrated in certain families, such as Maluridae and Lybiidae in birds and Canidae and Callitrichidae in mammals (Arnold and Owens 1998; Ligon and Burt 2004; Riedman 1982). Why an individual would forgo reproduction and remain at the natal nest to help rear the offspring of others is difficult to explain in light of Darwinian selection. Extensive empirical and theoretical research has been conducted in attempts to determine the factors that led to the evolution of philopatry, and a wide range of hypotheses have been proposed. Almost all of the explanations for philopatry can be categorized into one of three general hypotheses: those focused on life history characteristics, those emphasizing the benefits of philopatry and those based on ecological constraints on dispersal. The life history hypothesis proposes that cooperative breeding evolves in species with similar dispersal patterns and longevity (see review by Hatchwell and Komdeur 2000). The benefits of philopatry hypothesis emphasizes fitness benefits that can be obtained by an individual that remains philopatric (Stacey and Ligon 1991). These benefits can include increased indirect fitness by helping closely related individuals survive and reproduce (kin selection theory – see review by Clutton-Brock 2002). Direct benefits may include the acquisition of an important resource, such as improved survival while on the home territory (Emlen 1995; Kokko and Ekman 2002), access to food stores while on the natal territory (Stacey and Ligon 1987), the development of an important skill such as hunting through practice with group members (Bednarz and Ligon 1988), increased probability of obtaining a territory (Rood 1990), experience raising young or mating access to unrelated opposite sex conspecifics (Jennions 1 and Macdonald 1994). The ecological constraints hypothesis proposes that offspring will remain philopatric when ecological conditions limit the probability of successful dispersal and independent breeding (Emlen 1982). The ecological constraints hypothesis is the only one of the three hypotheses that proposes that dispersal is based on environmental limitations. Emlen (1982) formally developed the ecological constraints hypothesis based on the ideas of Selander (1964), Brown (1969), and Koenig and Pitelka (1981). The ecological constraints hypothesis focuses on four factors that could potentially determine whether an offspring will disperse and attempt to breed or remain philopatric within its natal group: (1) the cost/risk of dispersal, (2) the probability of successfully becoming established on a suitable territory, (3) the probability of obtaining a mate, and (4) the likelihood of successful reproduction once an individual obtains a territory and a mate (Emlen 1982). Many different types of constraints have been investigated in the past. Some researchers have looked at seasonal dynamics like temperature to evaluate changes in dispersal patterns and group size, because individuals may form groups to decrease thermoregulatory costs associated with decreasing temperatures (Getz et al. 1987; Getz et al. 1993). The aridity-food distribution hypothesis states that eusociality in naked mole-rats (Heterocephalus glaber) and Damaraland mole-rats (Cryptomys damarensis) may be due to the constraint on digging imposed by living in an arid environment with patchy food distribution (Jarvis et al. 1994). Tunnel expansion is primarily limited to times immediately following sporadic rainfall events. Group-living individuals work together and make large expansions to the burrow system in the brief window of opportunity a rain provides, thus lowering the cost of foraging and locating patches of food that can be shared among the group members (Jarvis et al. 1994, Lacey and Sherman 1997). A similar hypothesis has been proposed for the gundi (Ctenodactylus gundi), although it remains untested (Nutt 2005). The mate limitation hypothesis states that philopatry occurs due to a limited number of available mates (Marra and Holmes 1997), as seen in the superb fairy-wren, Malurus cyaneus (Pruett- Jones and Lewis 1990). Although poorly studied to date, support for the mate limitation hypothesis is slowly growing as experimental manipulations of available mates are performed (Dickinson and Hatchwell 2004). 2 The best studied
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