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Changes to Spider Community Ecology Mediated By CHANGES TO SPIDER COMMUNITY ECOLOGY MEDIATED BY ALTERED FOREST UNDERSTORY VEGETATION by Andrew P. Landsman A dissertation submitted to the Faculty of the University of Delaware in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Entomology and Wildlife Ecology Spring 2016 © 2016 Andrew P. Landsman All Rights Reserved ProQuest Number: 10157859 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. ProQuest 10157859 Published by ProQuest LLC (2016). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, MI 48106 - 1346 CHANGES TO SPIDER COMMUNITY ECOLOGY MEDIATED BY ALTERED FOREST UNDERSTORY VEGETATION by Andrew P. Landsman Approved: __________________________________________________________ Jacob L. Bowman, Ph.D. Chair of the Department of Entomology and Wildlife Ecology Approved: __________________________________________________________ Mark Rieger, Ph.D. Dean of the College of Agriculture and Natural Resources Approved: __________________________________________________________ Ann L. Ardis, Ph.D. Senior Vice Provost for Graduate and Professional Education I certify that I have read this dissertation and that in my opinion it meets the academic and professional standard required by the University as a dissertation for the degree of Doctor of Philosophy. Signed: __________________________________________________________ Jacob L. Bowman, Ph.D. Professor in charge of dissertation I certify that I have read this dissertation and that in my opinion it meets the academic and professional standard required by the University as a dissertation for the degree of Doctor of Philosophy. Signed: __________________________________________________________ W. Gregory Shriver, Ph.D. Member of dissertation committee I certify that I have read this dissertation and that in my opinion it meets the academic and professional standard required by the University as a dissertation for the degree of Doctor of Philosophy. Signed: __________________________________________________________ Douglas W. Tallamy, Ph.D. Member of dissertation committee I certify that I have read this dissertation and that in my opinion it meets the academic and professional standard required by the University as a dissertation for the degree of Doctor of Philosophy. Signed: __________________________________________________________ April M. Boulton, Ph.D. Member of dissertation committee ACKNOWLEDGMENTS My loving wife and daughter, whose support and patience helped me immensely throughout this endeavor. iv TABLE OF CONTENTS LIST OF TABLES ....................................................................................................... vii LIST OF FIGURES ....................................................................................................... ix ABSTRACT ................................................................................................................. xii Chapter 1 INDIRECT INTERACTIONS BETWEEN FOREST SPIDERS, WHITE- TAILED DEER, AND INVASIVE PLANTS .................................................... 1 White-tailed Deer Impacts to Vegetation and Vertebrate Wildlife .................... 1 Invasive Plant Impacts to Vegetation and Vertebrate Wildlife .......................... 3 Effects to Cursorial Arthropods .......................................................................... 5 Factors Influencing Forest Spider Ecology ........................................................ 7 Potential Impacts to Forest Spiders Mediated by Habitat Structure and Prey Availability ......................................................................................................... 9 2 ECOSYSTEM-ENGINEERING EFFECTS FROM HERBIVORES AND INVASIVE PLANTS ALTER SPIDER COMMUNITIES ............................. 13 Introduction ...................................................................................................... 13 Methods ............................................................................................................ 17 Study Area .................................................................................................. 17 Vegetation Sampling .................................................................................. 18 Arthropod Sampling ................................................................................... 19 Statistical Analyses ..................................................................................... 20 Results .............................................................................................................. 23 Vegetative Diversity and Structure ............................................................ 23 Insect Prey Abundance and Biomass ......................................................... 24 Spiders ........................................................................................................ 25 Discussion ......................................................................................................... 27 Acknowledgements .......................................................................................... 38 TABLES ........................................................................................................... 40 v FIGURES ......................................................................................................... 49 3 HABITAT STRUCTURE ALTERS INVERTEBRATE NUTRITIONAL DYNAMICS AS INDICATED BY CHANGES IN STABLE ISOTOPES .... 57 Methods ............................................................................................................ 61 Study System .............................................................................................. 61 Habitat Structure ......................................................................................... 63 Results .............................................................................................................. 64 Discussion ......................................................................................................... 64 Acknowledgements .......................................................................................... 68 FIGURES ......................................................................................................... 69 4 MANAGEMENT IMPLICATIONS ................................................................ 76 REFERENCES ............................................................................................................. 81 Appendix A PLANT SPECIES FOUND IN STUDY AREA .............................................. 98 B SPIDER SPECIES ............................................................................................ 99 C ISOTOPIC ENRICHMENT SAMPLES ........................................................ 101 vi LIST OF TABLES Table 1 Geographic locations of forest patches within study area. Coordinates represent approximate centroid of paired plots per forest fragment. Latitude is listed as decimal degrees North and Longitude listed as decimal degrees West. All coordinates reported in North American Datum of 1983 (NAD 1983). ................................................................... 40 Table 2 Classification of spider families into functional groups used in analyses. Groups are based on predominant prey capture method and natural history. Functional group designations were modified from Cardoso et al. (2011). .............................................................................. 41 Table 3 Jaccard community dissimilarity for herbaceous and woody vegetation throughout study area in July, 2013. Presence of vegetative species was pooled within forest patches and compared between each patch. .... 42 Table 4 Spider species richness and raw abundance in response to habitat structure and prey availability in August, 2013. Functional group abundances were analyzed using GLM with negative binomial probability distribution to account for overdispersion. Richness was analyzed using GLM with Poisson distribution. Likelihood ratio test statistic G2 was used to determine P-value. ............................................. 43 Table 5 Multivariate linear regression results for raw and relative abundance of spider functional groups in response to environmental variables. The orb web weaving, space web weaving, and hunting spider functional groups were included in analysis. Wilk’s Λ test statistic was used to assess significance. Spider data were collected in August, 2013. ........................................................................................................ 44 Table 6 Univariate linear regression results for relative abundance of individual spider functional groups in response to environmental variables. Spider data were collected in August, 2013. .......................... 45 vii Table 7 Univariate analysis of raw abundance of individual families in the orb web weaving, space web weaving, and hunting functional groups. Analyses were conducted using GLM with Poisson distribution. Likelihood ratio test statistic G2 was used to determine P-value. Members of the Clubionidae,
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