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Habitat Selection and Response to Restoration by Breeding

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Habitat Selection and Response to Restoration by Breeding HABITAT SELECTION AND RESPONSE TO RESTORATION BY BREEDING WESTERN SNOWY PLOVERS IN COASTAL NORTHERN CALIFORNIA By Stephanie D. Leja A Thesis Presented to The Faculty of Humboldt State University In Partial Fulfillment of the Requirements for the Degree Master of Science in Natural Resources: Wildlife Committee Membership Dr. Mark A. Colwell, Committee Chair Dr. Matthew D. Johnson, Committee Member Dr. William T. Bean, Committee Member Dr. Alison O'Dowd, Graduate Coordinator December 2015 ABSTRACT HABITAT SELECTION AND RESPONSE TO RESTORATION BY BREEDING WESTERN SNOWY PLOVERS IN COASTAL NORTHERN CALIFORNIA Stephanie D. Leja Habitat loss and degradation by invasive species is a primary limitation to the recovery of the Western Snowy Plover (Charadrius nivosus nivosus), a federally threatened shorebird that resides on coastal beaches in Humboldt County, California. This habitat threat posed by European Beachgrass (Ammophila arenaria) is often mitigated through dune restoration. Some habitat features altered by restoration (e.g., beach width) may influence plover breeding habitat selection. Further study was needed to determine which physical and social features (e.g., presence of conspecifics) influence plover nesting locations. I evaluated this response of plovers to restoration and identified characteristics that influence nest site selection. In an Information-Theoretic framework, I compared nests (n = 81) and random locations within habitat using logistic regression and Generalized Linear Mixed Models to produce Resource Selection Function analyses and conduct model selection analyses. Plovers nested on wider, less sloped beaches, with greater coverage of natural debris (e.g., driftwood, shells) and more conspecifics than at random locations. Plovers nested primarily (84%) in restored habitats, although this was influenced by one human-restored site with 33% of nests. These findings can guide coastal dune system managers to generate the features in restoration that improve nesting habitat and facilitate survival and recovery of this threatened Snowy Plover population. ii ACKNOWLEDGEMENTS Thank you to Dr. Mark Colwell for his invaluable guidance, advising, and opportunities for intellectual growth and experience. Thank you to my committee members Dr. Matthew Johnson and Dr. Tim Bean for their guidance and essential thesis reviews. Many thanks to the extensive group of Humboldt State student surveyors and volunteers who collected data as part of the larger Snowy Plover project, and especially to those who helped with my data collection. I thank J. Watkins from the U.S. Fish and Wildlife Service, J. Irwin of the U.S. Bureau of Land Management, and S. McAllister. Many thanks to the California State Department of Parks and Recreation: A. Transou, C. Wilson, M. Morrissette, J. Harris, T. Kurz, and C. Ryan. I am grateful to A. Desch and the HSU Wildlife Department for providing field equipment. I also extend my gratitude to the countless others who make this monitoring project happen and thus helped facilitate my research. This research was funded by the California Department of Fish and Wildlife, the California Department of Parks and Recreation, Humboldt State University, the U.S. Bureau of Land Management, and the U.S. Fish and Wildlife Service, and I thank these entities for their support. I also thank fellow shorebird graduate lab mates A. Patrick, D. Herman, M. Lau, T. King, and the Pink House roommates for their methodological conversations, draft revisions, and fellow graduate student empathy. I give a huge thank you to my wonderful parents, Mark and Hope Leja, and to Evan Blair for their limitless and essential support and advice throughout this experience. iii TABLE OF CONTENTS ABSTRACT ........................................................................................................................ ii ACKNOWLEDGEMENTS ............................................................................................... iii LIST OF TABLES ............................................................................................................. vi LIST OF FIGURES .......................................................................................................... vii LIST OF APPENDICES .................................................................................................. viii INTRODUCTION .............................................................................................................. 1 Plover Nest Site Selection ............................................................................................... 4 METHODS ......................................................................................................................... 7 Study Area ...................................................................................................................... 7 Field Methods ............................................................................................................... 10 Geospatial Methods to Obtain Additional Covariates .................................................. 11 Beach width .............................................................................................................. 13 Three habitat types .................................................................................................... 13 Conspecific attraction ............................................................................................... 13 Statistical Analyses ....................................................................................................... 15 Resource Selection Function analyses ...................................................................... 16 Habitat type analysis ................................................................................................. 18 RESULTS ......................................................................................................................... 19 Comparison of Restored and Unrestored Habitats ....................................................... 19 Comparison of Nest and Random Site Characteristics ................................................. 22 DISCUSSION ................................................................................................................... 31 Plovers Selected for Restored Habitats ......................................................................... 31 iv Physical Features are Important to Habitat Selection ................................................... 32 Social Attraction Influences Nest Location Selection .................................................. 35 Management Recommendations ................................................................................... 38 LITERATURE CITED ..................................................................................................... 41 APPENDICES .................................................................................................................. 49 v LIST OF TABLES Table 1. Definitions of variables used in logistic regression and other statistical analyses to evaluate habitat selection and response to habitat restoration of Snowy Plovers. Bracketed terms represent each of these covariates in the analyses. .................... 12 Table 2. Summary of the Chi-square goodness-of-fit analysis that evaluated selection between three habitat types by Snowy Plovers along 80 km of ocean-fronting beach in Humboldt County, CA in 2014. ............................................................. 20 Table 3. A comparison of average (± SD) physical and social characteristics at 81 Snowy Plover nests in three habitat types along ocean-fronting beaches in Humboldt County, CA in 2014. ............................................................................................. 21 Table 4. Summary of the comparison of habitat covariate values (mean ± SD) at nest locations and random points within plover nesting habitat in Humboldt County, CA, in 2014, derived from Student’s t-test statistics. ........................................... 23 Table 5. A summary of 19 competing models examining relationships between physical and social variables influencing habitat selection of nesting Snowy Plovers in Humboldt County, CA in 2014, showing the top five models with ΔAICc < 6 ranked by Akaike weight (wi), that explain 99% of selection and which represent hypotheses combining nine covariates, included the random effect of site (see Appendix E for full candidate model set). ............................................................ 24 Table 6. A summary of 18 competing models after removing one pretending model (which included the width covariate) examining relationships between physical and social variables influencing habitat selection of nesting Snowy Plovers in Humboldt County, CA in 2014, showing the top five models with ΔAICc < 6 ranked by Akaike weight (wi), that explain 99% of selection and which represent hypotheses combining nine covariates, included the random effect of site. ......... 25 Table 7. A summary of 19 models evaluating relationships between physical and social variables influencing habitat selection of a subset of 22 unique nesting male Snowy Plovers and 22 random points in Humboldt County, CA in 2014, ranked by Akaike weight (wi) and which represent hypotheses combining nine covariates, included the random effect of site. ........................................................................ 30 vi LIST OF FIGURES Figure 1. Map of study area in Humboldt County, California,
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