Extinction Risk and Population Declines in Amphibians
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Extinction Risk and Population Declines in Amphibians Jon Bielby Imperial College London, Division of Biology, A thesis submitted for the degree of Doctor of Philosophy at Imperial College London – May 2008 1 Thesis abstract This thesis is about understanding the processes that explain the patterns of extinction risk and declines that we see in amphibians, how we can use that understanding to set conservation priorities, and how we can convert those priorities into practical, hands-on research and management. In particular, I focus on the threat posed by the emerging infectious disease, chytridiomycosis, which is caused by the chytrid fungus, Batrachochytrium dendrobatidis ( Bd ). Amphibians display a non-random pattern of extinction risk, both taxonomically and geographically. In chapter two I investigate the mechanism behind the observed taxonomic selectivity and find that it is due to species biology rather than heterogeneity in either threat intensity or conservation knowledge. In chapter three I determine which biological and environmental traits are important in rendering a species susceptible to declines, focussing on susceptibility to Bd . I found that restricted range, high elevation species with an aquatic life-stage are more likely to have suffered a decline. Using these traits, I predict species and locations that may be susceptible in the future, and which should therefore be a high priority for amphibian research and conservation. 2 The use of predictive models to set conservation priorities relies on the accuracy of the modelling technique used. In chapter four I compare the predictive performance of both phylogenetic and nonphylogenetic regression and decision trees, and assess the suitability of each technique. Although phylogenetic regression provided the most predictive models of extinction risk and decline, decision trees could be a useful tool in future studies. Once high priority species or locations have been identified as susceptible to Bd , what should be the next step? In chapter five I identify highly susceptible species that have suffered an increase in extinction risk as a result of threat processes other than chytridiomycosis, and investigate the potential role of the disease in their decline. I find that the historical range of one of the four species investigated does harbour Bd at a relatively high prevalence, which suggests that structured surveillance of Bd in this location, and further investigation of the role of chytridiomycosis in the species’ decline would be warranted. In chapter six I describe such a structured, opportunistic screening for Bd on the island of Sardinia. I assess the threat that chytridiomycosis may pose to the threatened and endemic species of Sardinia and discuss future research directions and conservation needs of the island’s amphibians. 3 Declaration All of the work presented in this thesis is mine, with the following acknowledgements: Chapter two This chapter was published as a senior authored paper in Animal Conservation (Bielby, Cunningham, & Purvis, 2006, Animal Conservation, 9,135-143). The main idea behind the paper was my own, although Andy Purvis provided guidance on how to conduct the analyses. Olivia Curno, Sanjay Molur, Shawn McCracken of www.tadpole.org , Ronald Nussbaum of the University of Michigan, Rohan Pethiyagoda of the Wildlife Heritage Trust of Sri Lanka, Mark-Oliver Rödel, staff at Jatun Sacha research centre, Bruce Young of NatureServe helped with gathering site species check-lists. All aspects of data collection and collation, analysis, interpretation and writing are my own. Chapter three This chapter was published as a senior authored paper in Conservation Letters (Bielby, Cooper, Cunningham, Garner & Purvis, Conservation Letters, in press ). Natalie Cooper helped collect the data used in the analysis, and Andy Purvis provided guidance on the statistical techniques used. Tim Coulson provided advice on the statistical methods used. Emmanuel Paradis provided advice on the use of compare.gee function. S. Blair Hedges, Roberto Ibanez, Andres Merino-Viteri, Eugenia del Pino, Santiago Ron, and Tej Kumar Shrestha all helped in compiling data on the number of eggs per 4 clutch. The idea behind the chapter and all aspects of data analysis, interpretation and writing are my own. Chapter four The ideas behind the analyses in this chapter were developed by me with guidance from Andy Purvis. Natalie Cooper and Marcel Cardillo provided some of the phylogenetic comparative analyses presented in the chapter, and are acknowledged accordingly. The amphibian dataset underpinning the analyses were collected by me in collaboration with Natalie Cooper. The mammalian data analysed in the chapter was collected as part of the PanTheria project. All aspects of data analysis, interpretation and writing are my own. Chapter five The main idea behind this chapter is my own. Patricia Burrowes, Rafael Joglar, and Abel Vale helped with the field work conducted in Puerto Rico. Tissue samples from amphibians were provided by Stefan Lötters, Trent Garner, and Thomas Doherty-Bone. Barry Clarke allowed access to the NHM collection. Frankie Clare helped to process the laboratory samples. All aspects of data interpretation and writing are my own. Chapter six A grant awarded to Trent Garner, Stefano Bovero from the People’s Trust for Endangered Species provided funding for this chapter. The Sardinian NGO Zirrichiltaggi and their associates provided invaluable logistical help, sausage, 5 cheese and wine that proved essential for the fieldwork conducted in Sardinia. The Forestry workers and managers of Sardinia also provided invaluable support during my time on the island. Frankie Clare helped with the processing of the laboratory samples. All aspects of analysis, interpretation and writing are my own. 6 Acknowledgements Thanks to NERC for the funding that made my doctoral research possible. Also thanks to a grant from PTES which funded the work on Sardinia presented in this thesis. Thanks also to Simon Stuart and Janice Chanson provided help with use of the Global Amphibian Assessment database. To start with, big thanks to my supervision team, Andrew, Trent, and Andy. Before I started, someone pointed out that by the end I’d almost certainly hate my supervisors at least a little bit. Well, I can honestly say that has not happened as yet. Maybe after the viva. Thanks in particular to Andy, who I’ve now known for a long time, and who has given me real guidance, direction and confidence over the past few years. A famous writer once said: “All first drafts are shit. They are, however, necessary shit.” So a big thank you to all of the people who in the last three and a half years have read through the necessary and highlighted my “opportunistic” use of commas, my generally misuse of semi-colons, my chess-like use of paragraphs (“hhhmmm, I’ve not used the knight for a while, now seems like a good time”), my over-use of hyphens, and my generally poor grammar. These people include the many members of the Purvis lab 7 (too many to mention/remember) since 2004, and also Amanda Duffus, and Andrew King. There are, of course, periods of time during a PhD when all sense of reality goes out of the window. In those times, my mates were there to see me through. Thanks to all of those at Silwood, IoZ, and the Real World who have slapped me back down to earth every so often. Charlie, Shaun, Jimmy, Matty and Neil have always given me a sense of perspective when needed; thanks lads. Also, special thanks go to: Guy Cowlishaw who has helped me hold it together on a couple of occasions when I was about to come apart at the seams; 1001 shukrans to Trent and Dina who were basically always there for me, and fed me fish and wine when I dropped by like The Littlest Hobo; Ben and Janna who always had a pie on hand in case of an emergency; and a special firm-handshake to my office-mates for much of my PhD, Kingy and Amber who’ve……well, let’s not get all emotional. But thanks. Finally thanks to all of my family who’ve provided support, interest and encouragement. In particular, thanks to Auntie Jan and Uncle Ron for those days as Little Switzerland catching newts and eating Drewery’s hot-cakes. That’s pretty much where it all started, so thank you for being around and being interested and transferring some of your enthusiasm and energy to me. And finally thanks to Mum, Dad and Ali, because all this PhD really represents is me doing things that I like and care about and enjoy, and that’s what you’ve always encouraged me to do. Thank you. 8 Table of Contents Thesis abstract...............................................................................................2 Declaration .....................................................................................................4 Acknowledgements .......................................................................................7 Table of Contents...........................................................................................9 List of Tables................................................................................................12 List of Figures ..............................................................................................14 List of Appendices.......................................................................................15 List of Acronyms and Abbreviations………………………………………….16 Chapter 1: General Introduction .................................................................18 Chapter 2: Taxonomic Selectivity in