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The Pennsylvania State University The Graduate School Department of Geography REVISITING MESOPREDATOR RELEASE: CARNIVORE DYNAMICS ALONG A GRADIENT OF LANDSCAPE DISTURBANCE A Thesis in Geography by Andrew T. Townsend 2014 Andrew T. Townsend Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science August 2014 The thesis of Andrew T. Townsend was reviewed and approved* by the following: Robert P. Brooks Professor of Geography and Ecology Thesis Advisor Joseph A. Bishop Instructor in Geography, Research Associate for Riparia Thomas L. Serfass Professor of Biology at Frostburg State University Cynthia A. Brewer Professor of Geography Head of the Department of Geography *Signatures are on file in the Graduate School iii ABSTRACT Human induced habitat loss and predator persecution caused severe declines in apex carnivores throughout the North American continent. Removal of apex predators allowed smaller, lower rank predators from the Order Carnivora to become prominent. These "mesopredators" flourished, destabilizing ecosystems by driving many prey species toward extinction. However, some suggest that mesopredators still benefit from contemporary vegetation changes and fragmentation by thriving in disturbed areas. Many worry the versatility of these mesopredators could further threaten their prey species by leading to increased predation in anthropogenically- disturbed areas. This study seasonally sampled predator distributions along land cover gradients in forested, riparian corridors in Appalachia to identify whether landscape modification results in changes in carnivore community structure in the region. The study area consisted of randomly generated sites along streams in central Pennsylvania. I gathered data from camera traps and field surveys to catalogue the spatial ecology of mesopredators. I analyzed these data with landscape metrics to test the hypothesis that as forest contiguity decreases, both the abundance and richness of the predator community increases, possibly adding pressure on vulnerable prey populations. Through the analysis of these habitat metrics and carnivore occurrence data, this study found that carnivore species richness and abundance do generally increase with human disturbance in rural settings. However, this pattern is not due to the behavior of every species as many mesopredators are present across these rural landscapes and exhibit different responses to disturbance. Nevertheless, a few important generalists, namely the canids and raccoons, do show preferences toward more human disturbed areas and thus, are most accountable for this observed pattern. Keywords: spatial ecology, mesopredator, predation, forest decline, landscape gradients, riparian corridors, Appalachia, ecological cascades, landscape ecology iv TABLE OF CONTENTS List of Figures ........................................................................................................................ vi List of Tables ......................................................................................................................... vii Acknowledgements ................................................................................................................ viii Chapter 1 The Rise of the Mesopredator ................................................................................ 1 Mesopredators and Landscape Disturbance.................................................................... 4 The Newfound Mesopredator Community ..................................................................... 6 A Search for Solutions ................................................................................................... 8 Significance and Further Study ...................................................................................... 10 Chapter 2 The Mesopredators of Central Pennsylvania ......................................................... 12 Coyote (Canis latrans) ................................................................................................... 16 Opossum (Didelphis virginia) ........................................................................................ 18 Bobcat (Lynx rufus) ........................................................................................................ 19 Fisher (Martes pennanti) ................................................................................................ 21 Striped Skunk (Mephitis mephitis) ................................................................................. 22 Mink (Neovison vison) ................................................................................................... 24 Raccoons (Procyon lotor) .............................................................................................. 25 Gray Fox (Urocyon cinereoargenteus) ........................................................................... 27 Black Bear (Ursus americanus) ..................................................................................... 28 Red Fox (Vulpes vulpes)................................................................................................. 30 Chapter 3 Case Study: Mesopredator Community Dynamics along a Gradient of Landscape Disturbance in Central Pennsylvania ............................................................ 32 Landscape Change and Corridors ................................................................................... 32 Study Objectives ............................................................................................................ 37 Study Area ..................................................................................................................... 40 Methods ......................................................................................................................... 42 Results ............................................................................................................................ 62 Chapter 4 Implications, Discussion, and Conclusions ............................................................ 96 Appendix A Study Tables ............................................................................................. 105 Appendix B Survey and Assessment Forms .................................................................. 112 Literature Cited .............................................................................................................. 114 v LIST OF FIGURES Figure 2-1. A representation of the mesopredator community food web. .............................. 15 Figure 2-2. Canis latrans at a nonforested, heavily disturbed site. ......................................... 16 Figure 2-3. Didelphis virginia at a moderately disturbed site. ................................................ 18 Figure 2-4. Lynx rufus at a less disturbed, interior forest site. ................................................ 19 Figure 2-5. Martes pennanti at a less disturbed, interior forest site. ....................................... 21 Figure 2-6. Mephitis mephitis at a highly disturbed, agricultural site. .................................... 22 Figure 2-7. Neovison vison at a less disturbed, interior forest site. ......................................... 24 Figure 2-8. Procyon lotor at a less disturbed, interior forest site. ........................................... 25 Figure 2-9. Urocyon cinereoargenteus at a highly disturbed site. .......................................... 27 Figure 2-10. Ursus americanus at a moderately disturbed, forest-edge site. .......................... 28 Figure 2-11. Vulpes vulpes at a highly disturbed, agricultural site. ........................................ 30 Figure 3-1. Map of riparian study sites (n = 23) in the Ridge and Valley Province of central Pennsylvania. ...................................................................................................... 42 Figure 3-2. Example sites for forested (left) and nonforested (right) riparian zones. ............. 43 Figure 3-4. Example site buffers (1 km radius) with land cover. ........................................... 51 Figure 3-5. The approximate relative abundance of species in the predator community. ....... 63 Figure 3-6. Example scatterplots of several important metrics and species richness with best fitting regression lines. ............................................................................................ 67 Figure 3-7. A graph denoting site abundance in the predator community by season.. ............ 71 Figure 3-8. A scatterplot with a regression line showing predator abundance by edge density. ........................................................................................................................... 73 Figure 3-9. A scatterplot with a regression line showing opossum site abundance by edge density. ........................................................................................................................... 74 Figure 3-10. A scatterplot with a regression line showing carnivore site abundance by edge density, this time without opossums or raccoons. .................................................. 75 Figure 3-11. Percent Nonforest and the occupancy of both fox species. ................................ 78 vi Figure 3-12. Locations of red and gray fox occurrences in the study landscape..................... 79 Figure 3-13. Coyote occurrence locations detected by camera traps across study landscape. ......................................................................................................................