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i BIOENERGETICS AND HABITAT SUITABILITY MODELS FOR THE CHINESE MYSTERY SNAIL (BELLAMYA CHINENSIS) by Danielle M. Haak A DISSERTATION Presented to the Faculty of The Graduate College at the University of Nebraska In Partial Fulfillment of Requirements For the Degree of Doctor of Philosophy Major: Natural Resource Sciences Under the Supervision of Professors Kevin L. Pope and Valery E. Forbes Lincoln, Nebraska May, 2015 ii BIOENERGETICS AND HABITAT SUITABILITY MODELS FOR THE CHINESE MYSTERY SNAIL (BELLAMYA CHINENSIS) Danielle M. Haak, Ph.D. University of Nebraska, 2015 Advisors: Kevin L. Pope and Valery E. Forbes Relatively little is known about the invasive Chinese mystery snail (Bellamya chinensis). This research aims to elucidate some of the mystery surrounding this species. First, we place the species in context with other invasive freshwater snails of the USA and Canada, identifying current information gaps, categorizing shared characteristics among families and species, and comparing functional roles and ecological effects of freshwater snails. We conclude that more focus needs to be directed to regulating the aquarium, pet, and food trades if we are serious about preventing future invasions. Next, we develop a bioenergetics model for the species by quantifying and comparing consumption, egestion, respiration, and production of the Chinese mystery snail at varying water temperatures. We observed differences in these values across different water temperatures, indicating that temperature affects growth and reproductive strategies of this species. Then we focused on analyzing a specific case study to identify physical, chemical, and biological lake characteristics that help predict where the Chinese mystery snail is found. The top predictor model found that Chinese mystery snail presence is correlated with Secchi depth, latitude, and the presence of other aquatic invasive species. Finally, we use network analysis to develop a method for coupling social and ecological network models so they may be used in tandem to assess how humans aid the movement of the Chinese mystery snail, as well as how the snail affects an ecosystem after invasion. This was achieved through the adaptation of the framework of infectious disease network modeling. iii i Author’s Acknowledgments Where do I start? A PhD is so much more than just this document. To my co- advisors, Dr. Kevin Pope and Dr. Valery Forbes: thank you. There were ups and downs and surprises along the way, and you two guided me through each step with patience and encouragement (and tough love when I needed it). Not only have I changed as a scientist, I've changed as a person as a result of the past 4 years and for that I will always be grateful. Thank you for taking a chance on me. To my committee members, Dr. Chad Brassil, Dr. Amy Burgin, and Dr. Mark Pegg: thank you for enduring the snail ride as it evolved. Your suggestions, thought-provoking meetings, and inspiration were critical to the completion of this project. To Dr. Craig Allen and the IGERT committee: thank you for the unique opportunity of being an IGERT student. This program is an incredible launching pad and opened doors I didn't even know existed. To the University of Nebraska Foundation: thank you for the financial support enabling me to focus on my dissertation full time as a Presidential Graduate Fellow. To Dr. Brian Fath and the IIASA staff: thank you for giving me the opportunity to spend a terrific summer in Austria with i a magnificent international team. YSSP was a highlight of grad school, without a doubt. A very special thank you to the wonderful staff that keeps the NE Coop Unit and IGERT moving forward, especially Valerie Egger and Caryl Cashmere. I will be forever appreciative for the hours of work you both saved me and for your patience when I made things more difficult than they needed to be. Valerie, thank you for all of the help organizing 2 European adventures; I definitely didn't make things easy, but your support resulted in two flawless trips. I was lucky to join a great group of people comprised of SNR faculty, the NE Coop ii Unit, and the IGERT crew. I was even luckier to be part of the Pope Lab, and I cannot thank you guys enough. I am especially grateful to Chris Wiley, Robert Kill, Jason DeBoer, Brian Hammond, Kelly Turek, Luke Kowalewski, Nathan Stewart, Dr. Dustin Martin, and Dr. Chris Chizinski: we had some awesome (and a few not-so-awesome) times and my memories of graduate school will always be dominated by this beautiful, motley crew we were lucky to be a part of. To the Forbes Lab: I loved our talks on science and on life – thank you! To my fellow IGERTs, especially Shelli Hellman, Hannah Birgé, Maggi Swilinski, Ilonka Zlatar, and Noelle Hart: nothing bonds a group faster than being plopped down in a new country where no one speaks the native language. I am continuously inspired by your enthusiasm, optimism, encouragement, and quests for self-improvement, and I am so blessed to be able to call you all friends. To my fellow snailer, Bruce Stephen: thank you for the snail enthusiasm and the endless edits – I would have been lost without your snail expertise! Alec Wong: thank you for all of your dedication to field and lab work! I don’t have room to name everyone who contributed to this process but know you are in my heart and I am thankful for the role you played as ii advisor or friend or combination of the two. Last, but definitely not least, to my wonderful family, Dad, Mom, and Jonathon: thank you for not forcing me to grow up too fast. I've been able to wander, explore, question, and figure things out as I go, and none of that would be possible without an unbreakable support system. You taught me to keep striving for growth (and to remember there's always room for improvement), while still providing encouragement and unconditional love. And to my friends who became family, Lindsay Schaffner and Jereme Gaeta: thank you for everything you both are (On Wisconsin!). iii Table of Contents Chapter 1: Introduction ............................................................................................1 Chapter 2: Non-native and invasive freshwater snails of the USA and Canada....................................................................................................................15 Chapter 3: Carbon budget of the adult Chinese mystery snail (Bellamya chinensis) at various water temperatures ....................................................................................76 Chapter 4: Identifying environmental predictors of presence or absence of the adult Chinese mystery snail ..........................................................................................112 Chapter 5: Coupling ecological and social network models to assess “transmission” and “contagion” of an aquatic invasive species ..........................138 Chapter 6: Conclusions, management implications, and directions for future research ................................................................................................................186 iii Appendix A. Wet weight snail data .....................................................................194 Appendix B. Dominant fish species found in each water body ...........................199 Appendix C. Assessing pH and calcium requirements of the Chinese mystery snail ......................................................................................................................205 Appendix D. Ecological network development ...................................................212 iv List of Tables Table 2-1. Taxonomy and native ranges of the invasive freshwater snail species included in this review, as well as the number of species from each family native to the USA and Canada. An * indicates a “native transport” rather than an exotic species ............................................................................74 Table 2-2. Taxonomy of both native and non-native gastropods of North America ............................................................................................................75 Table 3-1. Summary data for adult snails at each of 3 water temperatures. Significant differences are indicated by different superscript letters ...............................104 Table 3-2. Mean (±SD) carbon and nitrogen content (%) for adult body tissue, fecal matter, and juveniles produced at each water temperature. Statistically significant differences within each category are indicated by different superscript letters ..............................................................................................................105 Table 3-3. Estimated carbon budget for Chinese mystery snails at each of three water temperatures. Numbers are mean specific rates with units of mg Carbon/gram body mass/day. Egestion values include both unassimilated material as well as excretion. .........................................................................106 Table 4-1. Environmental variables collected for each lake in the case study example. An * indicates variables that were available for > 75% of lakes and thus, included in all analyses ...................................................................129 v Table 4-2. Aquatic invasive species present in each lake included in case study. CMS = Chinese mystery snail (Bellamya chinensis), BMS = Banded mystery snail (Viviparus georgianus), RCF = Rusty crayfish (Orconectes rusticus), EWM = Eurasian water-milfoil (Myriophyllum spicatum), CLP = Curly-leaf pondweed (Potamogeton crispus),
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