DIFFERENTIAL RESPONSES OF GRASSLAND GROUND BEETLE (COLEOPTERA: CARABIDAE) COMMUNITIES TO CONSERVATION RESERVE PROGRAM MANAGEMENT PRACTICES A Thesis by Evan Waite Bachelor of Science, University of Florida, 2017 Submitted to the Department of Biological Sciences and the faculty of the Graduate School of Wichita State University in partial fulfillment of the requirements for the degree of Master of Science July 2019 © Copyright 2019 by Evan Waite All Rights Reserved DIFFERENTIAL RESPONSES OF GRASSLAND GROUND BEETLE (COLEOPTERA: CARABIDAE) COMMUNITIES TO CONSERVATION RESERVE PROGRAM MANAGEMENT PRACTICES The following faculty members have examined the final copy of this thesis for form and content and recommend that it be accepted in partial fulfillment of the requirement for the degree of Master of Science with a major in Biological Sciences. __________________________________ Mary Jameson, Committee Chair __________________________________ Gregory Houseman, Committee Member __________________________________ Andrew Swindle, Committee Member iii “We should preserve every scrap of biodiversity as priceless while we learn to use it and come to understand what it means to humanity.”—E.O. Wilson iv ACKNOWLEDGEMENTS This research project would not have been possible without the contributions of numerous individuals who so kindly dedicated extensive amounts of time, expertise, and unwavering support throughout the entire process. I would first like to thank Dr. Mary Liz Jameson for being the best mentor that anyone could ever ask for; I would not be half the scientist or person I am today without your guidance. Thank you to Alex Morphew and Fraser Watson for being with me on this journey every step of the way; even while knee deep in CRP. Having friends willing to help with anything at the drop of a hat is truly something special, and I would not have made it through this without you two. For assistance with the gargantuan task of sorting the carabids from our traps, I thank Morgan Trible and Katie Slavenburg. Your willingness to sort through mountains of beetle biomass was a blessing. As a new carabidologist, this project would not have gone anywhere without the knowledge and hospitality of Bob Davidson at the Carnegie Museum of Natural History, Dr. Zack Falin at the Snow Entomological Museum Collection, Dr. Dave Kavanaugh at the California Academy of the Sciences, and Dr. Kip Will at the University of California at Berkeley. Without access to their specimens and the verification of my identifications, this project would have stayed where these beetles reside; on the ground. I would like to thank Dr. Andrew Swindle for his guidance on my committee, Dr. Gregory Houseman for his assistance and willingness to help an entomologist embrace his ecologist side, and Molly Reichenborn for her constant support as a friend and project manager that has the super- human ability to make things happen. Finally, I would like to thank the people that have been my support system since day one; my family. My mom, Cindy Connors, as well as my grandparents Peggy and Jim Waite have v always supported every weird, buggy adventure I have chosen to pursue. Even over 1,000 miles away I felt your love and belief in me every single day. The research “Linking CRP grassland management to plant, insect, and bird abundance and diversity”, which is the subject of this thesis, has been financed, in part, with federal funds from the Fish and Wildlife Service, a division of the United States Department of Interior, and administered by the Kansas Department of Wildlife, Parks and Tourism. The contents and opinions, however, do not necessarily reflect the views or policies of the United States Department of Interior or the Kansas Department of Wildlife, Parks, and Tourism. vi ABSTRACT Grasslands of the Great Plains have declined over 70% since European settlement due to conversion of natural habitats to cropland and urban centers. The federally supported Conservation Reserve Program (CRP) was created to improve water quality, reduce soil erosion, and increase native habitats for wildlife. Within these restored grassland ecosystems, ground beetles (Coleoptera: Carabidae) are a keystone invertebrate group that fill several crucial niches and may serve as bioindicators of successful land management strategies. Using pitfall traps at 108 CRP sites across 650 km and a precipitation gradient of 63.5 cm, we examined the response of carabid beetle richness and abundance to a low diversity (CP2) and relatively high diversity (CP25) initial seeding and a grazing treatment (grazed or ungrazed) over two field seasons. We captured 4,841 carabid beetles representing 48 species across this wide gradient. Overall, grazing and CP treatments did not have a detectable effect on carabid abundance, biomass, or diversity. However, we found significant, positive effects of grazing on all three metrics in tallgrass habitats (strongly correlated with precipitation and plant community diversity). There were no effects of CP on the ground beetle community. Additionally, we found that species turnover was correlated with habitat transitions found along the precipitation gradient. Our findings suggest that moderate grazing by cattle over two seasons does not negatively affect carabid communities and that carabid communities respond differently to the effects of land management across a precipitation gradient and grassland habitats. vii TABLE OF CONTENTS Chapter Page 1. INTRODUCTION…………………………….……………….………………….............1 1.1 Introduction………………………………………………………………………..1 2. MATERIALS AND METHODS………………………..………………………..…….....6 2.1 Site Selection…………………….……….……………………………………......6 2.2 Insect Sampling……...…………….……………………………………….……...9 2.3 Plant Sampling…………...……………………..………………………………..11 2.4 Laboratory Methods…..…..……………………………………………………...11 2.5 Habitat Designations……………………………………………………………..13 2.6 Data Analyses…………………………………………………………………….13 3. RESULTS…………………………………..…………………………………………....17 3.1 Carabid Beetles………………………..……………………………..………......17 3.2 Carabid Abundance………………………………..………………..……………20 3.3 Carabid Biomass…………………………………………………………………21 3.4 Carabid Diversity………………………………………………………………...22 3.5 Response of Carabid Community to Habitat……………………………………..24 3.6 Species Turnover…………………………………………………………………29 4. DISCUSSION………..…………………………………………………………….…….35 4.1 Discussion………………………………………..……………………………....35 4.2 Response of Carabid Communities to Grazing and CP Treatments………..…....35 4.3 Species Turnover in Carabid Communities……………………………………...39 4.4 Carabids as Bioindicators and Comparison with European Ecology……………42 4.5 Significance………………………………………………………………………43 REFERENCES…………………………………………………………………………………..45 APPENDICES……………………………………………………………………….…………..57 viii LIST OF TABLES Table Page 1. Taxon table with abundances and feeding guild….……………….……………………..18 2. Most abundant species in each region……..…………………………………………….19 3. Kruskal-Wallis for overall treatment effects…………………………………………….23 4. PERMANOVA results for region………………………………………………………..26 5. PERMANOVA results for two region effects…………..……………………………….27 6. PERMANOVA for grazing and CP combinations….…………………………………...27 ix LIST OF FIGURES Figure Page 1. Plot layout……………………………………..…………………...…………...…………8 2. Overview of pit fall trap design……………………………………………………….....10 3. Rank abundance of carabid species………………………………………...……………20 4. Carabid abundances by treatment and region……………………………………………21 5. Carabid biomass by treatment and region………………………………...………………22 6. Carabid diversity by treatment and region……………………...………………………...23 7. Three-region NMS ordination……………………………….…………...………………25 8. Two-region NMS ordination……………..………………………………………………28 9. Total species turnover..………………………………………...………………………...30 10. Species turnover by Conservation Practice……………………………………………....31 11. Species turnover by grazing treatment…………………………………………………...32 12. Change in the dominant species………………………………………………………….33 13. Habitat designations of each site…………………………………………………………34 x LIST OF MAPS Map Page 1. Kansas map including all study sites……………...………………………………...……..7 xi LIST OF PLATES Plate Page 1. Pitfall trap in-situ………………….……………………………………………………..10 xii CHAPTER 1 INTRODUCTION 1.1 Introduction Over the past few years, a worrying trend has emerged in environmental ecology. Numerous reports demonstrate that insect populations are in severe decline due to anthropogenic stressors (Conrad et al. 2006, Fox et al. 2014, Hallmann et al. 2017, Lister and Garcia 2018, Benslimane et al. 2019). However, insect community reassembly is positively influenced by habitat restoration (Longcore 2003, Littlewood et al. 2006, Albrecht 2007). While restored habitats will never be the same as the original habitat, restoration efforts can mitigate some of the damage done to ecosystem processes. Restoring habitat is more important now than ever before, and focusing restorations on insect communities is crucial to maintaining healthy ecosystems because insects provide many necessary ecosystem services such as pollination, pest control, carrion and waste removal, and serving as food sources for other organisms (Ratcliffe 1996, Larochelle and Lariviere 2003, Kremen and Chaplin-Kramer 2007, Nichols et al. 2008). Ecosystem resiliency, or the ability to recover from disturbance, can be aided by the invertebrate community and the services they provide. One key invertebrate group that provides ecosystem services (Larochelle and Lariviere 2003), is heavily impacted by these human-imposed stressors (Koivula 2011), and responds positively to