Ecology of the Green Sea Turtle (Chelonia Mydas L) in a Changing World
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Ecology of the green sea turtle (Chelonia mydas L) in a changing world Submitted by Ana Rita Caldas Patrício to the University of Exeter as a thesis for the degree of Doctor of Philosophy in Biological Sciences August 2017 This thesis is available for Library use on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. I certify that all material in this thesis that is not my own work has been identified and that no material has been previously submitted and approved for the award of degree by this or any other University. Signed: ________________________ Ana R. Patrício Green turtle hatchling moving towards the ocean at Poilão, Guinea-Bissau 2 Abstract Climate change is threatening biodiversity, causing populations and species to adapt, or otherwise, become extinct. Sea turtles have survived dramatic climate changes in the past, however, due to a history of intense human exploitation, and the current anthropogenic threats, their current resilience may be jeopardized. The main pursuits of this thesis were to i) evaluate the resistance of green turtles to predicted climate change impacts, using a globally significant rookery, in Poilão, Guinea-Bissau, as a case study; and ii) assess key population parameters to inform the conservation management of this resource. As the work developed I additionally had the opportunity to study the dynamics of an emerging disease in a juvenile foraging aggregation from Puerto Rico, which contributed to a broader understanding of resilience in this species. Specifically, I investigate the nest site selection behaviour of green turtles, their nesting environment, and the outcomes for their offspring, at Poilão, and apply this information to infer on the resilience of this population under future scenarios of climate change. I explore the connectivity established by the dispersal of post-hatchlings from Poilão, followed by their recruitment to foraging grounds, to set the geographical context of this major population. Lastly, I model the dynamics of Fibropapillomatosis, which affects juvenile green turtles globally, and examine the potential for disease recovery. The green turtle rookery in Poilão shows some resilience to expected climate change impacts. This significant population likely contributes to all juvenile foraging aggregations along the west coast of Africa, and to some extent to those in South America. Currently, green turtles are capable of recovery from Fibropapillomatosis, however, the incidence of disease may be enhanced by climate change. 3 Acknowledgements I am most thankful to my wonderful supervisory team; Brendan Godley, Paulo Catry and Annette Broderick, for their guidance, advice and support. Thank you Brendan for taking me as your student without knowing me, for being so optimistic and supportive, and for pointing me in the right direction throughout my PhD. Thank you Paulo for being so enthusiastic about this project, for the incredible opportunity of conducting research at the most spectacular sea turtle rookery in the world, and for believing in me. Thank you Annette for being inspirational, and always caring. Thank you to the amazing duo, Carlos Diez and Robert van Dam, the first sea turtle gurus I met, in 2006, with whom I collaborated for my masters, and have continued to collaborate to this day, including the present thesis. A big thanks to Carlos Carreras, for his support with the genetic analyses, and for his friendship. Thank you also Lucy Hawkes, Rui Rebelo, and Angela Formia, for the important contributions to this thesis. Throughout my PhD I was funded by a scholarship from the Foundation for Science and Technology of Portugal (FCT). Additional financial support for fieldwork and laboratory analyses was granted by the MAVA Foundation, the Rufford Foundation, NOAA's National Marine Fisheries Service, US Fish and Wildlife Service, Chelonia, and Widecast. Logistics and research support at Guinea-Bissau was provided by the Institute of Biodiversity and Protected Areas (IBAP), and I am very pleased to have met and worked with IBAP’s technicians and nature rangers, ever so welcoming. I am most thankful to Aissa Regalla, coordinator of the monitoring of species and habitats; Castro Barbosa, focal point for sea turtles in Guinea-Bissau; and Quintino Tchantchalam, Director of the PNMJVP, for facilitating fieldwork and research permits, and for sharing their knowledge on the sea turtles from the Bijagós with me. Thank you also to Bucar Indjai, Amadeu Mendes de Almeida, Emanuel Dias, António José Pires, Betânia Ferreira Airaud, Dominic Tiley, Ana Marques, and Mohamed Henriques, for their contribution to data collection, and for making the time in Poilão even better. I am forever grateful to the Bijagós 4 people; the IBAP park rangers César Banca, Paulino da Costa, Carlos Banca, Santinho da Silva, Saído Gomes, Seco Cardoso, João Pereira and Tio Zé, and the participants from Canhabaque Island; Sana, Beto, Matchu, Rapaz, Sene, Correia, Tó, Cândido, Neto, Iaia, Chiquinho, Mário, and Joaquim, for their help in the fieldwork, and for sharing their culture with me, an invaluable gift. At Puerto Rico support was provided by the Department of Natural and Environmental Resources (DNER-PR), and numerous national and international volunteers, and DNER biologists participated in the fieldwork. A special thanks to my partner Miguel Varela, for his continuous support, his contribution for this thesis, and for joining ‘my world’ of sea turtle research and conservation. Last but not least, I thank my mother, the woman I most admire and my role model, for teaching me, among many other things, to love and respect all living creatures, and my daddy who encouraged and supported my first living abroad experience in Australia, which serendipitously led me to sea turtle conservation and thus to this PhD. 5 Table of contents Abstract .............................................................................................................. 3 Acknowledgements ............................................................................................ 4 Table of contents ................................................................................................ 6 List of tables and figures .................................................................................... 7 Author’s declaration .......................................................................................... 21 List of notations and abbreviations ................................................................... 24 General introduction ......................................................................................... 27 Chapter 1: Balanced primary sex ratios and resilience to climate change in a major sea turtle population ............................................................................... 31 Chapter 2: Nest site selection repeatability and success of green turtle Chelonia mydas clutches ................................................................................................. 75 Chapter 3: Climate change resilience of a globally important green turtle population ....................................................................................................... 113 Chapter 4: Dispersal of green turtles from Africa’s largest rookery assessed through genetic markers ................................................................................. 149 Chapter 5: Novel insights into the dynamics of green turtle fibropapillomatosis .......................................................................................... 185 Final remarks .................................................................................................. 213 Conservation implications............................................................................... 213 Future research .............................................................................................. 215 6 List of tables and figures Chapter 1: Balanced primary sex ratios and resilience to climate change in a major sea turtle population Figure 1a. Map of the Bijagós Archipelago, Guinea-Bissau: the João Vieira and Poilão Marine National Park is represented by the striped area, and the black frame depicts Poilão Island; b. Map of Poilão Island showing the four green turtle nesting beach sections monitored in this study (1-Farol, 2-Acampamento Oeste, 3-Acampamento Este, 4-Cabaceira). Pie charts present the mean nesting distribution across three habitats: ‘open sand’ (OS: white), ‘forest border’ (FB: grey), and ‘forest’ (F: black), in each section. Estimated mean proportion of males (M) and females (F) produced in each section are given (average across 2013 and 2014). Section 5-Praia Militar, was not monitored in this study due to difficult access and the small proportion of nests hosted there. (Maps created using www.seaturtle.org/maptool). Figure 2a, b. Mean bi-weekly air temperature (open circles) and precipitation (bar) at Bolama Island (http://cdo.ncdc.noaa.gov/CDO/cdo); c, d. estimated mean incubation temperature during the thermosensitive period (TSP) experienced by green turtle clutches laid from 15 June to 15 December at Poilão Island, at three habitats (OS-‘open sand’, FB-‘forest border’, F-‘forest’); e, f. bi-weekly proportion of green turtle nesting distribution at Poilão. Figure 3. Logistic function (solid curve) and 95% confidence intervals (CI, dashed curves) showing expected proportion of green turtle male hatchlings, as a function of