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Early Life History of a Landlocked Amphidromous Fish: Migration, Critical Traits and Ontogeny

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Early Life History of a Landlocked Amphidromous Fish: Migration, Critical Traits and Ontogeny Early life history of a landlocked amphidromous fish: migration, critical traits and ontogeny Jason M. Augspurger A thesis submitted for the degree of Doctor of Philosophy in Zoology At the University of Otago Dunedin, New Zealand 2017 Abstract Life-history plays a key role in determining species distributions and population structuring. Many studies evaluate the role of adult characteristics when predicting species distribution and population structuring, but very few evaluate the role of early life-history which is often the most vulnerable, and therefore limiting, phase of life-history. This thesis examines the role of early life-history in distribution and population structuring leading to divergence, and potentially speciation, by evaluating short and long term population structuring mechanisms, and the role of early life-history alteration in a facultatively amphidromous fish. Landlocked populations of Galaxias brevipinnis were found to be genetically divergent from diadromous populations, despite no obvious physical barriers to population connectivity. On shorter timescales, catchment level meta-populations were formed within lakes and along coasts. Larvae were found to be highly rheotactic through lab trials, and the majority of larvae were found to be retained within river plumes through larval trawling and otolith microchemistry, suggesting a simple behavioural mechanism responsible for both the long term isolation leading to divergence and short term meta-population structuring patterns seen here. Egg and larval size was larger in diadromous populations than in landlocked populations suggesting a potential pathway for the formation of landlocked populations and adaptation to local conditions. Despite size differences, responses to behavioural cues were similar in both diadromous and landlocked populations likely due to retaining a bipartite fluvial-pelagic life-history pattern. A review of amphidromous fish shows landlocking and life-history plasticity is common both within species and families, suggesting patterns seen here may be applicable across many amphidromous fish taxa. Further, the consistent use of pelagic rearing environments highlights amphidromy as an obligate benthic-pelagic migration where the various life- history patterns are a response to landscape centered on rapidly moving larvae to a pelagic environment. This thesis shows landlocked populations of an amphidromous fish diverging from their diadromous counterparts, due to a simple behavioural response, resulting in adaptation of early life-history in a novel environment, allowing fecundity optimization and providing the potential for context dependent isolation, adaptation, divergence and speciation. Acknowledgements First, thank you to my primary supervisor Gerry Closs. Gerry was always keen on a discussion and remained positive, supportive and enthusiastic no matter the results. I was also given free rein to pursue any questions I found interesting and pursue ideas that were likely to fail inevitably, but serve as a great learning experience. Without these attributes, this thesis would not have been possible. Thank you also to Travis Ingram who was also always keen to chat and give advice based on his work with koaro and prior field experience. I would imagine it is relatively rare for a student to have two supervisors with doors open all the time for anything from a chat about weekend plans to how to frame an introduction. Graham Wallis, Bruno David, and Mark Kaemingk also offered feedback and opinions on components in their respective disciplines and often took on roles similar to those of an adviser. While not recognized as advisers on university paper work, they all also deserve credit for advising this thesis. Similarly, I would like to thank the Closs lab which helped set me on the right track when I arrived. Without Manna Warburton, Andy Hicks and Peter Jones, the frame work of this thesis linking amphidromy to fluvial life-histories would have taken significantly longer to conceive. By suggesting potential questions continuing on from their work, I was guaranteed a solid base from which to build once I had a better grasp of New Zealand’s systems. I would also like to thank those who helped with fieldwork. Matt Jarvis is one of the few who would ever happily agree, and remain positive, after being told the ideal sampling time would be in the middle of a west coast flood and that we would be camping near the top of a pass in the middle of winter just to save 30 minutes off the drive the next day (if we are honest, we likely lost that amount of time waiting for the water to boil every time). While conditions were generally good, often times they were far from it and no one else would ever agree to go on those trips; only someone who truly loves ecology and being out there. Thank you also to Tony Stumbo who drove for the first field trips allowing more time to think when we were all convinced that maybe koaro were just a myth. Thank you also to the zoology department members and support staff. Matt Wylie and Erin Damsteegt were crucial in helping me to spawn and rear fish. Kim Garrett was called upon to drive the boat at night and to sit in Wanaka for two to three weeks waiting for the weather to cooperate. Murray McKenzie built the larval flume tank used for larval swimming trials. Ken miller helped produce many of the maps used in this thesis. Scott Jarvie and Luke Easton offered expertise and experience on stats. The doctoral scholarship from the University of Otago funded my research and ensured I could live comfortably throughout my Ph.D. I also owe a special thank you to the landowners who allowed me to sample fish on their property. I was understandably turned down by many landowners as findings could end up negatively affecting their daily routines, and potentially profit. Those who allowed me to sample did so knowing I would be looking for galaxiids and that results may have implications for their farming practices. These landowners were always keen to hear what I had found and what the best conservation practice was to ensure galaxiids remain in streams for years to come and deserve credit for actively supporting conservation. Thank you also to my family and friends who were, and are, ever supportive regardless of choices or outcomes and will have routinely helped maintain the work-life balance; especially when it was hard to find. Table of Contents 1 General introduction .................................................................. 1 1.1 How does life-history influence species distributions? ...................................................... 1 1.2 What is the role of migration in species distribution? ....................................................... 1 1.3 How do populations become isolated? .............................................................................. 2 1.4 Behaviour as an isolation mechanism ................................................................................ 3 1.5 Life-history adaptation ....................................................................................................... 3 1.6 Amphidromy ....................................................................................................................... 5 1.7 Study species ...................................................................................................................... 6 1.8 Thesis aim ........................................................................................................................... 8 1.9 Thesis structure .................................................................................................................. 8 2 Landscape biogeography and population structuring of a facultatively amphidromous fish .................................................... 10 2.1 Summary ........................................................................................................................... 10 2.2 Introduction ...................................................................................................................... 10 2.3 Methods ............................................................................................................................ 12 2.3.1 Study species............................................................................................................. 12 2.3.2 Sites, sampling design and method selection ........................................................... 13 2.3.3 Genetic techniques and analysis ............................................................................... 17 2.3.4 Otolith micro-chemistry techniques and analyses ................................................... 18 2.3.5 Larval trawling techniques and analyses .................................................................. 19 2.4 Results ............................................................................................................................... 20 2.4.1 Genetics .................................................................................................................... 20 2.4.2 Otoliths ..................................................................................................................... 24 2.4.3 Trawling .................................................................................................................... 28 2.5 Discussion ......................................................................................................................... 28 2.5.1 Long-term population structure
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