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Hybridisation, and the Conservation of the Grey Duck in New Zealand

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Hybridisation, and the Conservation of the Grey Duck in New Zealand Hybridisation, and the Conservation of the Grey Duck in New Zealand Wiebke Muller Supervisors: Dr. James Briskie Dr. Neil Gemmell 2 Dedication To my mother, who, ravaged by dementia, might no longer be able to grasp what the completion of my thesis signifies, but who resolved after her diagnosis that her illness would never restrict the options and choices in education and life of either my brother or myself; and to my father, who proudly supported that resolution every step of the way. 3 Acknowledgements Many people have helped me complete my work over the last three and a half years. I am indebted to all of them, and they appear in no particular order. My warmest thanks go to all the following individuals and organisations: To my supervisors, Jim and Neil, for taking me as a student when I found myself ‘orphaned’ just two month into my PhD, for their support and encouragement at that difficult time, and for all help and suggestions through the duration of the project; most of all for letting me pursue my project my way, while being quick to react whenever I did require information or help. To Jim for his sense of humour and rating me more valuable than a trap, as ‘lost students cause more paperwork’, to Neil for leaving all important equipment in the lab for two extra weeks so I could finish my work when he moved labs, to both for encouraging me that ‘if you can’t import the Drosophila into New Zealand, it just means you have to go where the flies are’. To all the labmates and technicians who have helped me find my way around the department, the labs, the library, and the paper work. In particular, Christine, Sophie, Melanie, Nicola, Ben and Tommy for pleasant days at Kaikoura, where they taught me how to handle birds, band them, and perform such procedures as venipuncture. Having mastered the art of taking blood from the miniscule veins of passerines, ducks surely seemed rather surprisingly simple, although a lot more difficult to restrain safely! 4 Also, and just as much, the patient people in the Molecular Ecology lab, who would tirelessly explain each procedure and each piece of equipment to someone entirely new to practical molecular work. To name just the most important, Margie, Iris, Amy and Jawad for guiding me through my first PCR and genotyping run, for showing me how to make and run a gel, and for recommendations on how to analyse the data. Maggie for being so wonderfully ready to help, never being fed up with last minute demands for reagents or autoclave runs, and her as well as Craig for putting up with a lot of questions on genotyping. Tammy for helping me with DNA extraction protocols and computer analysis of the microsatellite data. Everyone for helping me over the inevitable disappointments of molecular work, the failed amplifications and the mishaps. Dr. Terri Markow of the University of Arizona (now University of San Diego), for allowing me to come to her laboratory to pursue my study using desert Drosophila. Her and everyone in her lab for making my stay enjoyable and productive, for teaching me all I know about Drosophila husbandry, gathering, housing and handling. To Vanessa for the outbred flies, to Erin, Luciano, Tom, and Jeremy, for the explanations and hands-on help, for getting more Thanksgiving dinner invitations than I could attend, for a wonderful afternoon at the Desert Museum, for being taken out to sample flies in the desert and mountains, and for tolerating my family’s private phone calls to the laboratory! To all my flatmates and friends who supported me in my work as I tried to support them in theirs, for tolerating prototypes of duck traps to be erected in living rooms, and for living through monologues on duck hybrids every time we entered a park with a 5 pond! (Well, I lived through all those monologues on earthquake proof buildings, milk protein chemistry and wireless communications). In particular, to Nathan for driving me to Westport so I could see the ducks and field facilities there, and to Amber for reading all those jubilant and discouraged emails, and still finding time and energy for my worries, which rather paled against hers over her battle with that renegade bunch of cells in her spine. For the funding received from the Stocker Scholarship of the Forest and Bird protection Society (twice), from the Laura J Clad Memorial Scholarship and from the NZ Study Abroad Awards, as well as through the School of Biological Sciences, and Landcare Research’s FRST funded Outcome Based Initiative “Sustaining and Restoring Biodiversity”. To Patrick Jean-Guay for the samples of DNA of grey duck, mallard and domestic duck needed for the molecular work once I realised that I could not rely on the purity of any samples taken in New Zealand. To the farmers and hunters of HariHari and Haast, who did not just tolerate my work, but proved unexpectedly enthusiastic both about my work and making my stay alone for months in the big ‘green elephant’ a lot more enjoyable. For each and every farmer who allowed me to put up a trap on their pasture, and for the fact that nobody ever declined me that wish and that some even changed their grazing regimes for me, to the hunters who shared their catch and their knowledge. I am still amazed at that wonderful black 6 labrador returning a frightened duck alive, safe and sound, if somewhat stunned, after a noisy chase! To the quilting group that fed me fantastically every time they had their meetings in the field station (I looked forward to those meetings!), to everyone who invited me into their homes for dinners and tea, or just had a chat over a fence. To the family who let me look at their disabled ‘pet’ duck, which the cat brought in as a chick. To Craig who helped me in the first field season, and bore with me the frustration of the first weeks in which the ducks just refused to enter the traps. To Jack van Berkel who let me have a cleaner in the field station (and to Jeanette herself), for the internet connection there, for organising pots fit to cook for one or two, rather than twenty, for the washing machine, and for the mouse traps. They made such a phenomenal difference! To all the other guests who shared the field station with me, who tolerated blood samples in the fridge, duck dissections in front of the building, and who often fed me and always entertained me. And of course, to my family, who, if they grew tired of ducks, never showed it. To my father for encouraging and enabling my studies, to my brother and to my grandmother, and all the aunts and uncles and cousins. Some days, phone calls and emails really do make a difference. 7 Table of Contents Abstract ............................................................................................................................ 17 Introduction ..................................................................................................................... 21 Hybridisation in animals: the historical view ......................................................................................22 Hybridisation in animals: a paradigm shift .........................................................................................23 Hybridisation as a natural evolutionary process .................................................................................23 The wide range of consequences of hybridisation ...............................................................................25 Exogenous and endogenous selection ..................................................................................................27 Introgression: a matter of direction and amount .................................................................................28 Consequences at population level ........................................................................................................29 Transfer of adaptations ........................................................................................................................30 Non-genetic aspects: overlooked and understudied.............................................................................31 Hybridisation as a facultative, adaptive response ...............................................................................31 Hybrid speciation .................................................................................................................................32 Hybridisation as a conservation issue .................................................................................................33 The diverse range of issues with hybrids in conservation ....................................................................35 Hybrid swarms .....................................................................................................................................36 Setting conservation guidelines ...........................................................................................................38 Hybridisation in waterfowl ..................................................................................................................39 The mallard and grey duck hybridisation in New Zealand ..................................................................42 A history of the mallard and grey duck hybridisation ..........................................................................43 Prior studies aiming to understand the causes of the mallard and grey duck hybridisation ...............44 The aims of this thesis ..........................................................................................................................45
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