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Patia M. Connell Abstract PREDATOR OUT OF PLACE: THE DIFFERENTIAL IMPACTS OF NATIVE AND NON- NATIVE CRAYFISH ON BUFO TADPOLES By: Patia M. Connell June, 2010 Chair: Dr. Jeffery McKinnon Department of Biology Predators have been shown to alter important fitness components (larval period, mass at metamorphosis, and overall survivorship) of larval anurans. Uncommonly studied predators of tadpoles include crayfish. Both native and introduced crayfish occur in NC. To gain a better understanding of the effect of introduced crayfish in NC, we examined the impact of native (Procambarus acutus) and introduced (P. clarkii) crayfish on Fowler’s toad larvae (Bufo fowleri). We hypothesized P. clarkii would have a greater effect on fitness components of Bufo fowleri because it is larger, more aggressive and invasive in many parts of the world. Because these two species can co-occur, we conducted an experiment that allowed us to evaluate the independent and interactive effects of the two crayfish species at different densities (0, 1 or 2 individuals present). We found that crayfish did not alter toad larval period, but reduced toad survival and caused toads to metamorphose at larger masses. The effect of crayfish on toad survival and mass at metamorphosis increased with crayfish density. Interestingly, native crayfish caused a stronger reduction in survival and a stronger enhancement in mass at metamorphosis than did introduced crayfish when present at similar densities. Although toad survivorship in ponds with both crayfish species was comparable to survivorship in ponds containing only native crayfish, toads metamorphosed at larger sizes when both crayfish species were present. We caution, however, that P. clarkii can achieve higher densities than native species in some situations and likely have a more important effect under high-density conditions. PREDATOR OUT OF PLACE: THE DIFFERENTIAL IMPACTS OF NATIVE AND NON- NATIVE CRAYFISH ON BUFO TADPOLES A Thesis Presented To The Faculty of the Department of Biology East Carolina University In Partial Fulfillment of the Requirements for the Degree Masters of Science by Patia M. Connell June, 2010 ©Copyright 2010 Patia M. Connell PREDATOR OUT OF PLACE: THE DIFFERENTIAL IMPACTS OF NATIVE AND NON- NATIVE CRAYFISH ON BUFO TADPOLES by Patia M. Connell APPROVED BY: DIRECTOR OF THESIS:________________________________________ Dr. David R. Chalcraft, PhD COMMITTEE MEMBER:__________________________________________________ Dr. Derek Alderman, PhD COMMITTEE MEMBER:__________________________________________________ Dr. Kyle Summers, PhD COMMITTEE MEMBER:__________________________________________________ Dr. Heather Vance-Chalcraft, PhD CHAIR OF THE DEPARTMENT OF BIOLOGY ________________________________________________ Dr. Jeffery McKinnon, PhD DEAN OF THE GRADUATE SCHOOL: ________________________________________________ Paul J. Gemperline, PhD ACKNOWLEDGEMENTS I would like to thank my advisor, David Chalcraft, for his guidance over the past few years and for teaching me the careful methodology that good science requires. I’d like to thank the rest of my committee (Derek Alderman, Kyle Summers, and Heather Vance-Chalcraft) for their support and guidance as well. My lab mates have been an invaluable support system and I would like to specifically thank Jon Davenport and Natalie Amoroso for keeping me alive and sane. I am grateful to several people who helped with the project: Zac Aardweg, Freddy Herrera, Matt Prentice, Brooks Lowe, Robby Deans, John Haywood, Bob Connell, Jessica Kingsley, and Jason Hernandez. I am especially grateful to Heather Vance-Chalcraft for the use of her lab and equipment, and for keeping me apprised of opportunities for outreach to the community. I hope to continue working with her in the future. Thanks also to Melissa Meyers and Weyerhaeuser for allowing me to use your facilities at Cool Springs and for the volunteer opportunities you have provided. I would like to thank the teachers that have taught me so much about Eastern Carolina and its flora and fauna: Trip Lamb, Claudia Jolls, and of course David Chalcraft. Your lessons have already benefited me in many ways and will continue to do so in the future. Thanks also to those that helped my grim financial situation with odd jobs and babysitting commitments: Claudia Jolls, Jason Bond, Barbara Beltran, Jon Davenport, and any I have forgotten. Special thanks go to Barbara Beltran, without whom I may have never gotten registered for classes, paid for teaching, assigned labs to teach, or been recommended for many of the opportunities that I have had here at ECU. She has been a constant and reliable source of information, a sounding board, and a friend. I’d like to thank Barb for being there for all of the Biology graduate students. I’d like to thank friends and family who have been nothing but supportive through my graduate experience, even if it was usually from 500 miles away. Specific thanks to my parents who have provided endless support, encouragement and love throughout my graduate school career. This project was funded in part by a grant from the National Science Foundation, DEB- 0716558 TABLE OF CONTENTS ABSTRACT …………………………………………………………………………. i LIST OF FIGURES …………………………………………………...…………...… x LIST OF PLATES ……………………………………………………..…..…...…… xi CHAPTER 1: REVIEW OF SPECIES INVASIONS ………………....……….……. 1 References……………………………………………………………………... 17 CHAPTER 2: EXPERIMENTAL EVALUATION OF THE EFFECTS OF TWO CRAYFISH SPECIES ON FOWLER’S TOADS...…………………………………. 22 Introduction……….…………………….…………………………………...... 22 Methods…………………………………………………………………..…… 28 Statistics…………………………………………………………………..…… 32 Results……………..………………………………….…………………..…… 33 Discussion…………………………………………………… …………..…… 35 References…………………………………………………………………….. 41 Figures………………………………………………………………………… 44 CHAPTER 3: ASSESSMENT AND IMPLICATIONS OF THIS STUDY…………. 51 References…………………………………………………………………….. 58 APPENDIX A: ANIMAL USE PROTOCOL APPROVAL..………….……………... 59 APPENDIX B: PLATES…………………...………………………………………….. 61 LIST OF FIGURES Figure 1: Average effect of crayfish presence on Bufo fowleri survivorship, mass, and larval period.…………………………………………………………………………………...44 Figure 2: Average effect of crayfish density on Bufo fowleri survivorship, mass, and larval period……………………………………………………………………………………………45 Figure 3: Average differences in survivorship, mass, and larval period of Bufo fowleri as affected by Procambarus clarkii at low and high density………………………………………46 Figure 4: Average differences in survivorship, mass, and larval period of Bufo fowleri as affected by Procambarus acutus at low and high density………………………………………47 Figure 5: Average differences in survivorship, mass, and larval period of Bufo fowleri as affected by crayfish at low density…….………………………………………………………..48 Figure 6: Average differences in survivorship, mass, and larval period of Bufo fowleri as affected by crayfish at high density……………………………………………….………….…49 Figure 7: Actual vs. expected percentages of Bufo fowleri killed when exposed to a high density of crayfish…………………………………..……………………………………………………50 LIST OF PLATES Plate 1: Photographs of Procambarus acutus and P. clarkii……………………………………61 Plate 2: Range maps for Procambarus acutus and P. clarkii……………………...……………62 Plate 3: Images of Bufo fowleri adults and larvae……………………………………………….63 Plate 4: Range map of Bufo fowleri……………………………………………………………..64 Plate 5: Pictorial representation of experimental treatments……………………………………65 Plate 6: Photograph of mesocosms………………………………………………………………66 Plate 7: Representation of treatments used to calculate effect of crayfish presence……………67 Plate 8: Representation of treatments used to calculate effect of crayfish density………………68 Plate 9: Representation of treatments used to calculate effect of Procambarus clarkii at low and high density………………………………………………………………………………………69 Plate 10: Representation of treatments used to calculate effect of Procambarus acutus at low and high density…….……………………………………………………………………………70 Plate 11: Representation of treatments used to calculate different effects of Procambarus acutus and P. clarkii at low density…………………………………………………..…………………71 Plate 12: Representation of treatments used to calculate effect of Procambarus clarkii and P. acutus together and separately…………………………………………………………………..72 CHAPTER 1: Review of Species Invasions Humans have been moving species around the planet for as long as they themselves have been moving around the planet. In the beginning, when we were constrained by technology, the species we moved were few and portable: our diseases, our parasites, our domesticated livelihoods. As the human race developed, so did our ingenuity. While we were busy exploring the planet by land and water, the organisms we transported (with or without our knowledge) were exploring the places we took them. They lived and thrived or died out, according to their ecological needs and what was available in their new homes. The evidence of this global transplantation is sometimes apparent, but in some cases the immigrants fit in so well that they go unnoticed. Species such as dandelions, redbuds, daylilies, daffodils, and English ivy are so common in neighborhoods and along roadways that we may forget that they escaped someone’s garden and proliferated. As benign as these species seem, there are many organisms that can cause much damage when transplanted. Organisms that have been transplanted out of their natural range are known as nonindigenous, nonnative, exotic, or introduced species (Mac, Opler et al. 1998). When introduced species become
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