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Sciurus Carolinensis) in Ireland UCC Library and UCC researchers have made this item openly available. Please let us know how this has helped you. Thanks! Title Ecology of frontier populations of the invasive grey squirrel (Sciurus carolinensis) in Ireland Author(s) Goldstein, Emily Anne Publication date 2014 Original citation Goldstein, E. A. 2014. Ecology of frontier populations of the invasive grey squirrel (Sciurus carolinensis) in Ireland. PhD Thesis, University College Cork. Type of publication Doctoral thesis Rights © 2014, Emily A. Goldstein. http://creativecommons.org/licenses/by-nc-nd/3.0/ Item downloaded http://hdl.handle.net/10468/1927 from Downloaded on 2021-10-11T09:54:00Z Ecology of frontier populations of the invasive grey squirrel (Sciurus carolinensis) in Ireland Emily A. Goldstein MSc, BA This thesis is presented to the National University of Ireland, Cork, in candidature for the degree of Doctor of Philosophy Research supervisors: Dr Fidelma Butler Dr Colin Lawton School of Biological, Earth and Environmental Sciences, College of Science, Engineering and Food Science, University College Cork Head of School: Professor John O’Halloran May 2014 Abstract The rise in invasive species, together with habitat destruction, is associated with worldwide declines in biodiversity and ecosystem functioning. Management and control of invasive species, as well as amelioration of invasion impacts, provide unending challenges to species and ecosystem ecologists as well as conservation managers. All continents and many islands have experienced species invasions, often accompanied by severe and long-lasting ecological and economic costs. Although any species can become invasive if it is transported to, establishes in and spreads in a new environment outside of its native range, rodents are a particularly frequent invader. Rodent introductions are often inadvertent but are also commonly intentional as these animals are traded and transported as pets and may escape from captivity. Tree squirrel species are attractive to humans and are able to establish populations with only a few founding individuals, making them a group well suited to performing the role of biological invaders. The eastern grey squirrel (Sciurus carolinensis) is the most commonly introduced squirrel species worldwide. This research addressed the grey squirrel invasion and frontier population biology. Novel results were generated through diverse research techniques. Public squirrel sighting surveys and hair-tube surveys were used to locate the southern frontier of grey squirrel range expansion in Ireland. A 22-month intensive live trapping study of two frontier populations facilitated the collection of personality and demographic data from squirrels in increasing populations. A systematic literature search on grey squirrel demography provided context for the studied populations, among frontier and established introduced populations, as well as those in the native range. Advanced spatially explicit population modeling techniques predicted future uncontrolled range expansion and objectively compared the outcomes of 12 grey squirrel management strategies. i The current southern extent of the grey squirrel was identified during the two-year citizen science survey that was completed in 2012. Hair-tube surveys recorded grey squirrels in four of the 14 study sites, but were not effective in recording the species close to the leading edge of expansion identified in the citizen science survey. During the live trapping work a total of 130 individual grey squirrels were captured on 1,072 occasions over 3,880 trap days in two frontier sites. Simple measures of personality provided a baseline dataset for the species and indicated that personality is driven by both state dependent and individual variation. Population densities were low and breeding rates high in both populations indicating increase. The annual survival rate was estimated at 31.7% ± 4.7 and was not significantly different between the two sites. After controlling for habitat type, the literature search identified trends in demographic parameters that differentiated between frontier and established introduced grey squirrel populations as well as those from the native range. The grey squirrel invasion has caused varying impact levels in different European countries and these were reflected in demographic parameters. Without intensive and coordinated region-wide management or control, the grey squirrel population is predicted to continue to expand to the south and south west of Ireland. Fatal control measures were predicted to have a greater effect on reducing squirrel abundance and slowing expansion than non-fatal immunocontraception control techniques. Habitat modification measures via broadleaf removal or carrying capacity reduction were also predicted to be effective in reducing abundance and slowing range expansion. The methods and results are discussed in both a basic scientific and applied invasion management context. An improved understanding of the behaviour, population dynamics, and future scenarios at the frontier of species invasions is crucial for managers worldwide and this is provided here for the grey squirrel in Ireland. ii Acknowledgements That I was able to create and present the body of work that follows is due in no small part to the significant support I have received over the last few years and longer. The skills and experience that I have gained through this project have made all the tough times worthwhile. Thank you to everyone whose presence and advice helped me reach the finish line. Most special thanks, first of all, are due to my supervisors, Colin Lawton and Fidelma Butler. I learned so much from both of you over the last few years about the whole research process from design, techniques, analysis, writing, and about being a scientist in general. Colin, you taught me to not take myself so seriously, how to think positively and how to turn problems into solutions. Fidelma, you taught me to think creatively and to always question what I thought I knew. You both guided me through the system masterfully and were great role models. Thank you both for your experience and stories. The Irish Research Council funded the majority of this project. Thanks are due for additional funding from UCC School of BEES, NUIG Department of Zoology and COFORD. Thanks to Coillte, for permission to carry out fieldwork on their properties. Thanks to Marie Roche for excellent Coillte crop maps. I wouldn’t have gotten far without the UCC technical staff. Special thanks to Luke Harman for the sturdy construction of hair tubes and squirrel traps, Stuart Warner for GIS genius, and Allan Whitaker for facilitating the vehicles. Thanks also to John Quinn, a brilliant scientist and teacher, for his interest and collaboration. Warm thanks to Pádraig “Pat” Whelan for his support. Special thanks go to my “big squirrel sister”, Emma Sheehy, the one who went before me. You are an inspiring scientist and you taught me so much over the last few years. Thank you for being there and for all the advice you generously shared. Thanks to Amy Haigh for advice on innumerable topics. To my officemates in the ERI: Tad, Eugene, Rosalyn, Maria, Fabio, Erin, Marion, and Roz. And thanks to iii Margaret in NUIG, your turn is next! Fieldwork was made easier and more enjoyable by the presence of many: especially Aisling (fieldwork champion), Tad, Eugene, Darragh, Amy and Erin. Huge appreciation goes to the OPW tour staff (Patricia, Mary, Margaret and Karen) at the Swiss Cottage in Cahir and the caretaker Ollie (for emergency trap repair skills). Margaret, thank you for letting me stay with you while trapping. Karen, I always enjoyed the days you were out with me the most! I am grateful to the other postgrads in UCC, especially Megan, and colleagues in the ERI for distraction, commiseration, and especially all of the tips and tricks. I owe a great debt to my first scientific mentors, Gary Gillis and Stan Rachootin, professors at Mount Holyoke College, MA. Gary, you first showed me how to critically read a scientific paper, what research actually was, how to suture and how to make a killer Powerpoint presentation. Stan, you showed me the beauty in the natural world and the depth and breadth of your knowledge is awe-inspiring. Thank you for knowing I was destined for squirrels long before I did, and for the generous gift of Monica Shorten’s “Squirrels” as I left MHC in 2007. Thanks, with love and affection, to the rodents who have passed through my life. To Pepper, Salty, Pete and Lance – the chinchillas that started my interest in rodents. Special thanks to Jerry the degu, my constant companion these last few years. I have nothing but unending thanks and gratitude for both of my parents for their constant support (emotional, financial and otherwise) and belief in my ability to accomplish anything. You placed the utmost value on education, always pushed me to do my best and taught me that it’s always worth working a little bit harder. This thesis is my tribute to you. Grateful thanks to my younger brother, David, for constant encouragement, humour, and the generous offer to search Yale’s electronic library for those hard to access PDFs. And finally, my ultimate thanks go to Vince Murphy. You encouraged me the whole way along and helped in more ways than I could ever list. Thank you from the bottom of my heart and know that this wouldn’t have happened without you. iv Declaration This thesis has not been submitted, in whole or in part, to this or any other university in candidature for any other degree and is, except where otherwise stated, the original work of the author. Chapter 2, except where otherwise stated, has been accepted in its current form by Wildlife Research (doi: 10.1071/WR13197). The author list is as follows: Goldstein EA, Lawton C, Sheehy E, Butler F. Input of each author to chapter 2: Author 1: Experimental design, all field and laboratory work, all analysis, and all writing except for the following: Author 3: Design of the citizen science questionnaire and website.
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