Direct and Indirect Effects of the Invasive Orconectes Rusticus on Native O
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Direct and indirect effects of the invasive Orconectes rusticus on native O. sanbornii in Ohio streams THESIS Presented in Partial Fulfillment of the Requirements for the Degree Master of Science in the Graduate School of The Ohio State University By Christopher Allen Johnson Graduate Program in Environment & Natural Resources The Ohio State University 2016 Master's Examination Committee: Dr. Lauren M. Pintor, Advisor Dr. Suzanne M. Gray Dr. Stephen N. Matthews Copyrighted by Christopher Allen Johnson 2016 Abstract While the direct effects of non-native species as predators and competitors have been widely studied, the indirect effects of invasive species as novel prey are relatively unexplored. Yet, these novel energetic pathways are of great importance to long-term community response to biological invasions. Novel prey, as supplemental resources for native predators, can positively influence predator abundances. These changes in predator populations, and the associated shifts in predation risk, may result in concomitant effects upon native prey. These indirect effects may impose significant consequences on native prey populations. Aquatic systems are among the most impacted by biological invasions. Globally, freshwater crayfish are among the taxa most at risk to the effects of invasive species. In Ohio, the invasive rusty crayfish (Orconectes rusticus) has been implicated in the decline of a native congener, O. sanbornii. This research aimed to evaluate the influence of indirect effects of invasive O. rusticus on patterns of replacement of the native O. sanbornii in Ohio streams. I used a combination of field observations and experimental manipulations to fulfill three objectives. First, I quantified population demographics of O. sanbornii from streams invaded and not yet invaded by O. rusticus. I also measured and compared the body size of O. rusticus in streams where they currently co-occur with O. sanbornii or have replaced the native species. Second, I conducted a within stream tethering experiment to test for differences in predation risk between these two species. Finally, I used a laboratory experiment to explore the effect ii of predation risk, competitor identity, and the interaction of the two factors on the behavior of both O. sanbornii and O. rusticus. Results of field surveys indicated a significantly smaller average O. sanbornii body size in streams where they are found co- occurring with O. rusticus. However, there was no detectable difference in O. rusticus body size between streams where they currently co-occur versus streams which they have displaced O. sanbornii. There were no detectable differences in sex ratios or risk of predation across populations of either species. Results of the laboratory experiment indicated that both species respond to increased risk of predation with increased refuge use and reduced measures of activity. However, risk effects depended upon competitor identity. When competing with conspecifics, O. sanbornii significantly increased its use of refuge with elevated predation risk, while decreasing measures of activity. However, when competing against invasive O. rusticus, O. sanbornii only increased refuge use at high predation risk and demonstrated increased activity at zero and low risk. In contrast, O. rusticus decreased activity with elevated risk, an effect which was heightened when facing a heterospecific competitor. Native O. sanbornii suffers a tradeoff between managing the risk of predation versus cost of competing with a novel competitor. However, O. rusticus responds to competition from O. sanbornii in ways which are advantageous in risky environments. These results suggest that the behaviors of O. rusticus may allow the species to minimize the trade-off between finding food and avoiding predators and out-compete native O. sanbornii. iii Acknowledgments I’d like to first thank my advisor, Lauren Pintor, for providing me opportunity to embark on my graduate career under her mentorship. Some of my most enjoyable experiences pursuing this degree have been the discussion of research ideas and ecological theory in various meetings over these last few years. Through this, she has taught me the importance of seeing beyond initial appearances, to dig to discover the true nature being presented, but also to step back and see the “big picture”. I would also like to thank my graduate committee, Suzanne Gray and Stephen Matthews for their insight and patience as this work has processed through its numerous iterations. Additionally, I would like to thank all the members of Pintor lab; Lauren Hostert for the encouragement and all the help that first year in acclimating to graduate life, and Jenna Odegard for the never ending positivity and always willing hand whether it be cleaning years old muck out of tanks or impromptu trips to the field. I’d also like to thank Lynn and Doug McCready, Kay Stefanik, Richard Oldham for the all the help (and sunburn) in building, literally, the foundations for the initial iterations of this work. I would also like to thank my friends and fellow graduate students for the encouragement and good conversation. I would like to give a special thanks to all the undergraduates that have aided me over these last few years, in particular, Erin O’Shaughnessey and Scott Meyer. Know iv that this research would not have been completed without your help. Additionally, I’d like to thank my advisors and mentors from FMU, Jeff Steinmetz, Travis Knowles, and Lorianne Turner. Each of you has played an instrumental role in getting me where I am today. Lastly, I would like to thank my family for putting up with all the missed holidays and phone calls, and providing support and encouragement along the way. v Vita May 2001 .......................................................Titusville High School, FL May 2013 .......................................................B.S. Biology, Francis Marion University August 2013 to present .................................Graduate Associate, School of Environment and Natural Resources, The Ohio State University Publications Johnson, C. A. and J. D. Camper. Geographic Distribution. Farancia erytrogramma erytrogramma. Herpetological Review, 2013 Fields of Study Major Field: Environment and Natural Resources vi Table of Contents Abstract ............................................................................................................................... ii Acknowledgments.............................................................................................................. iv Vita ..................................................................................................................................... vi Publications ........................................................................................................................ vi Fields of Study ................................................................................................................... vi Table of Contents .............................................................................................................. vii List of Tables ..................................................................................................................... ix List of Figures .................................................................................................................... xi Chapter 1: Introduction to the effects of biological invasion ............................................ 1 The Invasion Pathway ..................................................................................................... 2 Consequences of Biological Invasion ............................................................................. 4 Impacts of Non-Native Prey ............................................................................................ 5 Animal Behavior and Personality-Biased Invasion ........................................................ 7 Indirect Effects of Novel Prey ......................................................................................... 9 Study System ................................................................................................................. 10 Objectives ...................................................................................................................... 12 Figures ........................................................................................................................... 14 Chapter 2 : Direct and indirect effects of the invasive Orconectes rusticus on native O. sanbornii in Ohio streams ................................................................................................. 15 Abstract ......................................................................................................................... 15 Introduction ................................................................................................................... 16 Methods ......................................................................................................................... 20 Field Survey............................................................................................................... 20 Statistical Methods .................................................................................................... 22 Results ........................................................................................................................... 23 Discussion ....................................................................................................................