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HABITAT ASSOCIATIONS OF CARRION BEETLE (COLEOPTERA: SILPHIDAE) COMMUNITIES FOUND ON SMALL MAMMAL CARRION IN THE KANSAS FLINT HILLS A Thesis by Emmy L. Engasser Bachelor of Science, Wichita State University, 2014 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 December 2017 © Copyright 2017 by Emmy L. Engasser All Rights Reserved HABITAT ASSOCIATIONS OF CARRION BEETLE (COLEOPTERA: SILPHIDAE) COMMUNITIES FOUND ON SMALL MAMMAL CARRION IN THE KANSAS FLINT HILLS 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 Liz Jameson, Committee Chair ______________________________________ Leland Russell, Committee Member ______________________________________ Peer Moore-Jansen, Committee Member iii There are two things after death: putrefaction, the work of microbes, and disappearance, the work of insects. -Yovanovitch, 1888 iv ACKNOWLEDGMENTS First, I would like to thank Dr. Mary Liz Jameson. Her passion inspired me to pursue entomology. Without her support and guidance, I would not be the same person I am today. I thank Wichita State University for funding and use of their facilities, and the well-known Maria Martino and Marcia Norton (Wichita State University) for help with putting out fires and anything we needed. Thanks to my committee members, Dr. Leland Russell for his help with experimental design and statistical questions and Dr. Peer Moore-Jansen for providing forensic expertise, the funds needed to initiate this research, and granting access to WSU’S Skeleton Acres Research Facility. Their assistance and constructive criticism during this project was greatly appreciated. Special thanks to Rachel Stone, the other carrion queen, who was beside me through all the laughter, blood, sweat, and tears of this project. We did it! Rachel and I would not have been able to execute this research without the wonderful help of our research technicians: Jackie Baum, Jacqueline Nascimento, Brandon Hein, Breanna Sayers, Ethan Grennan, Niall Horton, Hannah Hoetmer, Faith Hanna, Morgan Trible, Theresa Wolff, and Nicole Brown. Thank you for stepping up to the stinky tasks! My sincere gratitude also goes out to: Drs. Brett Ratcliffe and MJ Paulsen (University of Nebraska-Lincoln) for assistance in identification of beetles; Dr. Josh Perkin (Texas A&M University) for guidance with statistical analyses; Juju Wellemeyer (Missouri Department of Conservation) for helping with statistics, brainstorming and being the best motivator a friend could ask for; Ranger Randy Just and Seth Turner (Kansas Department of Wildlife, Parks and Tourism) for their coordination and approval to conduct research at El Dorado State Park; Joyce Dudeck, Kim and Scott Bays for allowing this research to be conducted on their beautiful v property; Larry Slayton (Sedgwick County Zoo, Wichita) for ordering many rats for this research; and Matthias Seidel and Axel Gonzalez for help with Czech translation (Univerzita Karlova v Praze, Czech Republic). I am also thankful for my supportive family during this time, especially my parents, Mike and Gigi Dudeck; my husband, Kyle Engasser; and my friends that have been here for me throughout the entire process and new friends I have made along the way. I love you all. vi ABSTRACT Necrophagous insects play an important role in decomposition and nutrient recycling of decomposing animals. Ecological studies of carrion-associated beetles enhance forensic investigations by providing information about community assemblages and predictable patterns of succession. Gaps in forensic entomology, include: 1) repeatable, replicable ecological research, 2) research sites across all geographic zones, and 3) research conducted throughout a full annual cycle. To address these gaps and observe habitat associations of carrion beetles, the influence of habitat (woodlands versus grasslands) and abiotic factors on carrion beetle (Coleoptera: Silphidae) communities were examined at three sites located within the Kansas Flint Hills. A total of 3,333 adult carrion beetles in nine species were collected from pitfall traps baited with rat carrion over twelve 4-week collecting periods. Silphid beetle community differed in species composition between grassland and woodland habitats, but communities did not differ significantly in overall mean abundance, mean species richness, or mean species diversity. Six species exhibited strong habitat associations; two associated with grassland habitats (Nicrophorus marginatus and Necrodes surinamensis), and four with woodland habitats (Nicrophorus orbicollis, Necrophila americana, Oiceoptoma noveboracense, and Nicrophorus pustulatus). These results are relevant for predicting patterns of silphid beetle communities in the Kansas Flint Hills and assist in determining corpse relocation in forensic studies. vii TABLE OF CONTENTS Chapter Page I. INTRODUCTION………………………………………………………………………...1 II. MATERIALS AND METHODS………………………………………………………….5 STUDY SITES……………………………………………………………………5 SAMPLING PROTOCOL………………………………………………………...8 MEASUREMENT OF ABIOTIC VARIABLES…………………………………9 LABORATORY METHODS……………………………………………………10 ANALYSES……………………………………………………………………...10 III. RESULTS……………………………………………………………………………......14 IV. DISCUSSION……………………………………………………………………………18 REFERENCES……………………………………………………………………………...…...21 APPENDICES…………………………………………………………………………………...26 viii CHAPTER 1 INTRODUCTION Carrion, or decomposing animal tissue, is an important ephemeral, nutrient resource for many organisms, including microbial, plant, invertebrate, and vertebrate communities (Barton et al. 2013, Benbow et al. 2016). Necrophagous insects, primarily Diptera and Coleoptera, play a large role in decomposition and nutrient recycling of these decaying animals (Payne 1965, Ratcliffe 1996, Kočárek 2003). Studies observing the environmental influences upon these insect assemblages have applications for community ecologists and forensic entomologists. An important concept of community ecology is that biotic and abiotic factors in the environment limit community assemblages and species interactions. Many insect species exhibit strong habitat associations (Bajerlein et al. 2011, Parry et al. 2016), and reasonable evidence supports the hypothesis that habitat types influence community composition of Silphidae species (Walker 1957, Anderson 1982, Shubeck 1983, Kentner and Streit 1990, Lingafelter 1995). Kočárek (2003) found that rat carrion in an open meadow site decayed much faster than rat carrion in a forest site, which suggests that these habitats play a significant role in rates of decomposition. This information is relevant in predicting patterns of insect communities with forensic importance. For example, some species of necrophagous beetles are used to determine if a corpse has been relocated after death in forensic investigations (Matuszewski et al. 2013, Madra et al. 2014). Forensic entomology relies on experimental data of necrophagous insects in the United States as evidence in criminal cases to determine postmortem interval (PMI) or period of insect activity (PIA) (Tomberlin et al. 2011, Harvey et al. 2016). However, lack of standardized research protocols and credibility of scientific information derived from these studies requires 1 wholesale changes to serve society more effectively (National Research Council 2009). Many gaps remain within forensic research involving necrophagous insects, including: 1) repeatable, replicable ecological research, 2) research sites across all geographic zones, and 3) research conducted throughout a full annual cycle (Tomberlin et al. 2011, Michaud et al. 2012, Tomberlin et al. 2012). An unreplicated treatment (or simple pseudoreplication; Benbow et al. 2016) is the most common experimental design error in forensic science articles published between 1985- 2009 (Michaud et al. 2012). This leads to inflated power of statistical tests and makes it more likely that researchers will conclude that an effect exists when it may not (Hurlbert 1984). Another problem with pseudoreplication is that the treatment level is confounded with natural variation that occurs within that treatment level, such as site effects. Pseudoreplication is avoided by conducting and designing experiments that have adequate replications using multiple carcasses within replicated treatments. Forensic entomology research that encompasses natural variability should be performed in all biogeographic regions, ecosystems, and habitats; multiple study sites within broad regions are needed to measure site effects and variability between sites (Michaud et al. 2012). Studies that are designed in this fashion will add validity to evidence at crime scenes that may take place in areas with no previous record of necrophagous insects. To address these gaps in forensic ecological research in the Kansas Flint Hills, we developed a repeatable framework that lacks pseudoreplication and includes natural variability for one full year. Existing research on carcass decomposition has primarily revolved around the succession and composition of fly species (Diptera) (Bala and Singh 2015, Nadeau et al. 2015). Beetles (Coleoptera) have received less consideration for estimating PMI within forensic science (Midgley et al. 2010),