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Female Dispersal and Inbreeding in the Red-Cockaded Woodpecker

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Female Dispersal and Inbreeding in the Red-Cockaded Woodpecker FEMALE DISPERSAL AND INBREEDING IN THE RED-COCKADED WOODPECKER by Susan J. Daniels Thesis submitted to the Faculty of the Virginia Polytechnic Institute and State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE in Biology Approved by Jeffrey R. Walters, Chair Robin M. Andrews David R. Notter August 25, 1997 Blacksburg, VA Keywords: Picoides borealis, natal dispersal, breeding dispersal, inbreeding depression, inbreeding avoidance, optimal inbreeding, cooperative breeding Female Dispersal and Inbreeding in the Red-cockaded Woodpecker Susan J. Daniels (ABSTRACT) Dispersal is a critical life-history component; it determines gene flow and has profound effects on population structure, demography, social systems, and population viability. To add to our knowledge of dispersal and, in particular, our understanding of the relationship between dispersal and inbreeding, I studied three aspects of the biology of the red-cockaded woodpecker: dispersal of breeding females; the costs, benefits, and frequency of inbreeding; and the effect of inbreeding on natal dispersal. Dispersal of breeding female red-cockaded woodpeckers is strongly associated with inbreeding avoidance and mate choice, weakly associated with site choice, and not found to be associated with social constraints. Estimates of mortality for non-dispersing and dispersing breeding females were 24 and 59 percent per year, respectively—rare evidence of the cost of breeding dispersal. Significant costs of close inbreeding were found. Closely related pairs (kinship coefficient greater than 0.1) had lower hatching success as well as lower survival and recruitment of fledglings than unrelated pairs. Moderately related pairs (kinship coefficient between 0 and 0.1) and moderately inbred individuals had increased hatching success, but did not produce more young. Despite documented costs of close inbreeding and a predictable spatial distribution of closely related males near the natal territory, female fledglings disperse a median of only two territories and a modal distance of one territory. Natal dispersal of females is affected by closely related males on the natal site but unaffected by closely related males or moderately related males that are off the natal site. Dedication To my mother, Ellen. Acknowledgments I thank Jay Carter III, Phil Doerr, and Jeff Walters for the opportunity to use a remarkable data set, and the many people who have participated in the collection of these data. Thanks to R. Meekins and A. Felty for training, assistance, and pleasant company in the field. Thanks to J. Priddey for the method to calculate dispersal distance in numbers of territories crossed, and to R. Barr for his work in estimating territory centers. R. Andrews and D. Notter gave much- appreciated advice throughout this project. Special thanks to Jeff Walters for his superb direction of my Master’s research, and to Jim Lyons for constant personal support. iii TABLE OF CONTENTS ABSTRACT.............................................................................................................................................................. ii DEDICATION ..........................................................................................................................................................iii ACKNOWLEDGMENTS.............................................................................................................................................iii LIST OF TABLES..................................................................................................................................................... vii LIST OF FIGURES..................................................................................................................................................... ix CHAPTER 1: INTRODUCTION................................................................................................................................. 1 BREEDING DISPERSAL .......................................................................................................................................... 1 COSTS, BENEFITS, AND FREQUENCY OF INBREEDING.............................................................................................. 2 NATAL DISPERSAL AND INBREEDING..................................................................................................................... 3 THE RED-COCKADED WOODPECKER...................................................................................................................... 4 Conservation status........................................................................................................................................ 4 Ecology.......................................................................................................................................................... 4 Social system.................................................................................................................................................. 5 Population dynamics ...................................................................................................................................... 5 THE STUDY AREA AND METHODS OF DATA COLLECTION......................................................................................... 6 CHAPTER 2: DISPERSAL OF BREEDING FEMALES ................................................................................................... 7 INTRODUCTION ................................................................................................................................................... 7 Tests of hypotheses......................................................................................................................................... 7 Additional analyses ........................................................................................................................................ 9 METHODS ........................................................................................................................................................... 9 Study species, study area, and methods of data collection .............................................................................. 9 Definitions and measures ............................................................................................................................. 10 Sample sizes ................................................................................................................................................. 11 Statistical analyses....................................................................................................................................... 11 RESULTS........................................................................................................................................................... 12 Annual frequencies of dispersal, site fidelity, and mortality.......................................................................... 12 Frequency of dispersal by circumstance ....................................................................................................... 13 The test for inbreeding avoidance................................................................................................................. 14 Tests of mate choice ..................................................................................................................................... 14 Tests of site choice ....................................................................................................................................... 18 Tests of social constraints............................................................................................................................. 19 Age of the breeding female........................................................................................................................... 21 Distance dispersed by breeding females ....................................................................................................... 23 Mortality rates by circumstance and the cost of dispersal............................................................................. 25 DISCUSSION ...................................................................................................................................................... 26 Multiple causes of breeding dispersal........................................................................................................... 26 Inbreeding avoidance................................................................................................................................... 27 Mate choice.................................................................................................................................................. 27 Site choice.................................................................................................................................................... 28 Social constraints......................................................................................................................................... 30 Age of the breeding female........................................................................................................................... 31 Mortality rates ............................................................................................................................................. 31 Dispersal distance.......................................................................................................................................
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