FERTILIZATION AND PLANT LITTER EFFECTS ON THE PLANT AND EPIGEAL ARTHROPOD COMMUNITIES A dissertation submitted to Kent State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy by L. Brian Patrick December 2009 Dissertation written by L. Brian Patrick B.A., University of Kansas, 1998 M.A., University of Kansas, 2001 Ph.D., Kent State University, 2009 ________________________________, Chair, Doctoral Dissertation Committee Mark W. Kershner, Ph.D. ________________________________, Members, Doctoral Dissertation Committee James L. Blank, Ph.D. ________________________________ Alison J. Smith, Ph.D. ________________________________ Lauchlan H. Fraser, Ph.D. ________________________________ Randall J. Mitchell, Ph.D. Accepted by ________________________________, Chair, Department of Biological Sciences James L. Blank, Ph.D. ________________________________, Dean, College of Arts and Sciences John R. D. Stalvey, Ph.D. ii TABLE OF CONTENTS LIST OF FIGURES ……………………………………………………………………..vii LIST OF TABLES …………………………………………………………………..........x ACKNOWLEDGMENTS ………………………………………………………………xii CHAPTER I. Introduction………………………………………………………………..1 Biodiversity-productivity theory…………………………………..3 Plant litter …………………………………………………………5 Trophic relationships in the epigeal arthropod community ………7 Dissertation organization …………………………………………9 References ……………………………………………………….11 II. Large-scale manipulation of plant litter and fertilizer in a managed successional temperate grassland ………………………………………..23 Abstract…………………………………………………………..23 Introduction………………………………………………………24 Materials and Methods …………………………………………..26 Study site and experimental design ……………………...26 Plant community sampling ……………………………...28 Abiotic sampling ………………………………………...29 Statistical analyses ………………………………………30 Results …………………………………………………………...33 General trends …………………………………………...33 Species-level analyses …………………………………...37 Plant group responses …………………………………...42 Ecosystem response ……………………………………..46 Discussion …………………………………………………….....49 References …………………………………………………….....56 III. Beetle (Coleoptera) responses to fertilization and plant litter in a temperate old-field grassland …………………………………………………….....63 Abstract ……………………………………………………….....63 Introduction ……………………………………………………...64 Materials and methods …………………………………………..67 Study site and experimental design ……………………...67 Plant community sampling ……………………………...69 Beetle community sampling ………………………….....70 Statistical analyses ………………………………………71 iii Results …………………………………………………………...72 General beetle responses ………………………………...72 Beetle species-level responses …………………………..78 Discussion …………………………………………………….....84 References …………………………………………………….....91 Appendix A ……………………………………………………...98 Appendix B …………………………………………………….106 Appendix C …………………………………………………….109 Appendix D …………………………………………………….110 Appendix E …………………………………………………….111 Appendix F ……………………………………………………..112 Appendix G …………………………………………………….113 IV. Epigeal predator responses to fertilization and plant litter: testing biodiversity theory at the ground level ………………………………...117 Abstract ………………………………………………………...117 Introduction …………………………………………………….118 Materials and methods …………………………………………123 Study site and experimental design …………………….123 Spider community sampling …………………………...125 Statistical analyses ……………………………………..126 Results ………………………………………………………….128 General trends ………………………………………….128 Spider species-level analyses …………………………..139 Aggregate ecosystem-level analyses …………………...146 Discussion ……………………………………………………...150 References ……………………………………………………...159 Appendix A …………………………………………………….168 Appendix B …………………………………………………….171 Appendix C …………………………………………………….172 Appendix D …………………………………………………….173 Appendix E …………………………………………………….174 Appendix F ……………………………………………………..175 V. Review of the Nearctic genus Scyletria Bishop and Crosby (Araneae, Linyphiidae), with a transfer of S. jona to Mermessus O. Pickard- Cambridge ……………………………………………………………...177 Abstract ………………………………………………………...177 Introduction …………………………………………………….177 Materials and methods …………………………………………178 Taxonomy ……………………………………………………...180 Genus Scyletria Bishop and Crosby 1938……………...180 Scyletria inflata Bishop and Crosby 1938 ……………..180 iv Mermessus jona (Bishop and Crosby 1938) new combination …………………………………………….187 References ……………………………………………………...198 VI. Fertilization effects on predator diversity in a terrestrial detrital food web …………………………………………………………………………..202 Abstract ………………………………………………………...202 Introduction …………………………………………………….203 “Green” world vs. “Brown” world and cascading effects ……..................................................................................206 Materials and methods …………………………………………209 Study site and experimental design …………………….209 Arthropod community sampling ……………………….210 Statistical analyses ……………………………………..212 Results ………………………………………………………….213 Discussion ……………………………………………………...221 References ……………………………………………………...227 VII. Summary and importance of this study ………………………………..237 Future directions ……………………………………………….241 References ……………………………………………………...243 A I. First record of the genus Myrmedonota Cameron (Coleoptera, Staphylinidae) from North America, with descriptions of two new species …………………………………………………………………………..248 Abstract ………………………………………………………...248 Introduction …………………………………………………….248 Materials and methods …………………………………………250 Taxonomy ……………………………………………………...250 Genus Myrmedonota Cameron, 1920 ………………….250 Myrmedonota aidani Maruyama and Klimaszewski, sp. nov. ……………………………………………………..253 Myrmedonota lewisi Maruyama and Klimaszewski, sp. nov. ……………………………………………………..259 Modified key to species of Zyras group of genra in America North of Mexico ………………………………………………………265 References ……………………………………………………...266 A II. Eight new Ohio state records of true bugs (Hemiptera, Heteroptera) from pitfall traps ……………………………………………………………..268 Abstract ………………………………………………………...268 Introduction …………………………………………………….269 Materials and methods …………………………………………269 Collection sites ………………………………………....270 Results and discussion …………………………………………271 v Hebridae ………………………………………………..271 Miridae …………………………………………………271 Nabidae ………………………………………………...275 Reduviidae ……………………………………………..275 Rhyparochromidae ……………………………………..279 Thyreocoridae ………………………………………….283 References………………………………………………………285 vi LIST OF FIGURES CHAPTER 1. Introduction. Figure 1. Line graph of the number of journal article titles and “topic” hits containing the word “biodiversity” in a database search of ISI Web of Knowledge …………………………………………………………………………………......2 CHAPTER 2. Large-scale manipulation of plant litter and fertilizer in a managed successional temperate grassland Figure 2. Average plant biomass, Shannon’s H, percent soil organic content and moisture, and PAR ………………………………………………………………35 Figure 3. Ordination diagrams of the first two axes of canonical correspondence analysis (CCA) with the four treatments as environmental dummy variables ….41 Figure 4. Average species richness and average biomass of each growth form plant group within each treatment …………………………………………….....44 Figure 5. Average species richness and average biomass of the native and non- native plant groups within each treatment ………………………………………45 Figure 6. Two-dimensional ordination of ecosystem-level properties from 2005 in 24 experimental plots from nonmetric multidimensional scaling (NMS) ……47 CHAPTER 3. Beetle (Coleoptera) responses to fertilization and plant litter in a temperate old-field grassland Figure 7. Average effective Shannon’s H’ (eH’) of beetle species in each treatment ………………………………………………………………………...73 Figure 8. Species richness, abundance, and biomass of all beetles, carabids, and staphylinids ……………………………………………………………………...77 Figure 9. Regressions of all beetles, carabids, and staphylinids against abundance, biomass, plant species richness, and standing crop biomass …….....80 Figure 10. Average abundance of selected species by year ………………….....82 CHAPTER 4. Epigeal predator responses to fertilization and plant litter: testing biodiversity theory at the ground level vii Figure 11. Average effective Shannon’s H’ (eH’) of spider species in each treatment ……………………………………………………………………….137 Figure 12. Species richness, abundance, and biomass of all spiders, linyphiids, and lycosids …………………………………………………………………….138 Figure 13. Regressions of all spiders, linyphiids, and lycosids against abundance and biomass …………………………………………………………………….140 Figure 14. Regressions of all spiders, linyphiids, and lycosids against plant species richness and standing crop biomass …………………………………...141 Figure 15. Average abundance of selected species by year …………………...143 Figure 16. Two-dimensional ordination of ecosystem-level properties from 2005 from nonmetric multidimensional scaling (NMS) ……………………………..148 CHAPTER 5. Review of the Nearctic genus Scyletria Bishop and Crosby (Araneae, Linyphiidae), with a transfer of S. jona to Mermessus O. Pickard- Cambridge Figure 17. Scyletria inflata Bishop and Crosby 1938, male palpus…………....185 Figure 18. Mermessus jona (Bishop and Crosby 1938) new combination, male palpus and female epigynum………………………….......................................192 Figure 19. Geographic distribution of Scyletria inflata Bishop and Crosby 1938 and Mermessus jona (Bishop and Crosby 1938) new combination ……………193 CHAPTER 6. Fertilization effects on predator diversity in a terrestrial detrital food web Figure 20. Abundance, biomass, and species richness of all arthropods, predators, and prey ………………………………………………………………………...216 Figure 21. Regressions