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Demographic Consequences of Rapid Climate Change and Density Dependence in Doctoral Thesis Doctoral Migratory Arctic Geese

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Demographic Consequences of Rapid Climate Change and Density Dependence in Doctoral Thesis Doctoral Migratory Arctic Geese Doctoral theses at NTNU, 2020:156 Kate Layton-Matthews Kate Layton-Matthews Demographic consequences of rapid climate change and density dependence in Doctoral thesis Doctoral migratory Arctic geese ISBN 978-82-326-4660-9 (printed ver.) ISBN 978-82-326-4661-6 (electronic ver.) ISSN 1503-8181 Doctoral theses at NTNU, 2020:156 theses at NTNU, Doctoral NTNU Philosophiae Doctor Department of Biology Thesis for the Degree of Thesis for the Degree Faculty of Natural Sciences Faculty of Natural Norwegian University of Science and Technology Kate Layton-Matthews Demographic consequences of rapid climate change and density dependence in migratory Arctic geese Thesis for the Degree of Philosophiae Doctor Trondheim, May 2020 Norwegian University of Science and Technology Faculty of Natural Sciences Department of Biology NTNU Norwegian University of Science and Technology Thesis for the Degree of Philosophiae Doctor Faculty of Natural Sciences Department of Biology © Kate Layton-Matthews ISBN 978-82-326-4660-9 (printed ver.) ISBN 978-82-326-4661-6 (electronic ver.) ISSN 1503-8181 Doctoral theses at NTNU, 2020:156 Printed by NTNU Grafisk senter … Meanwhile the wild geese, high in the clean blue air, are heading home again. Whoever you are, no matter how lonely, the world offers itself to your imagination, calls to you like the wild geese, harsh and exciting – over and over announcing your place in the family of things. Mary Oliver, Wild Geese A barnacle goose family feeding on the tundra at Ny-Ålesund, Svalbard (©Jasper Doest). Acknowledgements It was a big decision moving to a country I had hardly visited and a city I had never heard of. But the risk paid off. The first week I arrived I met our Svalbard team - Bart, Mathilde, Brage and Vidar - and was whisked off to Mathilde’s cabin and I already started feeling at home. In the last four years, I realised that the group at CBD is really something special. Especially the PhD(+) gang where I have found so many good friends rather than just colleagues. A huge thanks to all you guys for uncountable skiing and climbing trips, meet-ups, parties, cave raves, dinners and the rest… To Vidar and Brage – for being exceptional supervisors, not just with all of the academic support (which I would have been lost without) but the ‘real life’ support too. To Maarten, you are an inspiring person, thanks for your genuine curiosity and enthusiasm in all things. Susanne, my bachelor’s supervisor, for being the first person to tell me I should consider ecology and to Arpat, my master’s supervisor, you were/are a huge inspiration and gave me the best possible career start. Takk til Marta, Hanne og Mari - I was supposed to be helping you guys but in the end you were all a huge support for me. To Mum for saying I’ll be the next David Attenborough (you never know…) and Dad for much-needed advice at times. To friends from home for supporting me even though I left London almost 10 years ago…and for the amazing people I have met along the way in Manchester and Zürich, during fieldwork in Sweden and Svalbard, and on adventures further afield, I am extremely privileged to have met so many inspirational people from different walks of life. Table of contents List of papers ..................................................................................................................................................... 5 Synopsis 1| Summary ......................................................................................................................................... 6 2| Introduction .................................................................................................................................... 8 Arctic climate change .............................................................................................................. 8 Population responses to climate change ........................................................................... 9 3| Thesis objectives ......................................................................................................................... 11 4| Methods ........................................................................................................................................ 13 Study system ........................................................................................................................... 13 Data collection ........................................................................................................................ 14 Integrated population models............................................................................................. 15 Path analysis............................................................................................................................ 15 E-SURGE ................................................................................................................................... 15 Integral projection models ................................................................................................... 16 5| Results and discussion .............................................................................................................. 17 6| Conclusions and prospects ....................................................................................................... 22 7| References .................................................................................................................................... 24 Papers I–IV List of papers I. Layton-Matthews K, Loonen MJJE, Hansen BB, Coste FD, Sæther B-E, Grøtan V (2019) Density- dependent population dynamics of a high Arctic capital breeder, the barnacle goose. Journal of Animal Ecology. doi: 10.1111/1365-2656.13001 II. Layton-Matthews K, Hansen BB, Grøtan V, Fuglei E, Loonen MJJE (2019) Contrasting consequences of climate change for migratory geese: predation, density dependence and carryover effects offset benefits of high-arctic warming. Global Change Biology. doi: 10.1111/gcb.14773 III. Fjelldal MA*, Layton-Matthews K*, Lee AM, Grøtan V, Loonen MJJE, Hansen BB (2020) High-arctic family planning: earlier spring onset advances age at first reproduction in barnacle geese. In press, Biology Letters. doi: 10.1098/rsbl.2020.0075 IV. Layton-Matthews K, Grøtan V, Hansen BB, Loonen MJJE, Fuglei E, Childs DZ. Environmental change reduces body mass, but not population growth, in an Arctic herbivore. Manuscript. Declaration of contributions Paper I: KLM, BBH and VG designed the study. MJJEL collected goose data. KLM analysed the data with input from VG and CFDC. KLM wrote the manuscript, which was reviewed and commented on by BES, BBH, MJJEL and VG. Paper II: KLM designed the study with input from MJJEL, BBH and VG. MJJEL provided goose data and EF provided fox data. KLM analysed the data with input from BBH, MJJEL and VG. KLM wrote the manuscript, which was reviewed and commented on by all co-authors. Paper III: KLM, BBH and VG designed the study. MJJEL provided the data. MAF analysed the data, with input from AML and KLM. KLM and MAF wrote the manuscript, which was reviewed and commented on by all co-authors. MAF and KLM contributed equally to the study and are both co-first authors (*). Paper IV: KLM and DZC designed the study, with input from BBH and VG. MJJEL provided goose data and EF provided fox data. KLM analysed the data, with input from DZC. KLM wrote the manuscript, which was reviewed and commented by all co-authors. 5 1| Summary Climate change effects are being felt across affected egg production and hatching success ecological levels and most strongly in the Arctic. through rapid advancement of spring onset. Scaling up from effects on demographic rates to However, this was offset by reduced pre-fledging the population-level, and even community-level, survival due to increasing predation by Arctic is a daunting task. This is particularly the case for foxes, an indirect consequence of climate change migratory species that experience different linked with changes in abundance of climate and density regimes over their annual overwintering herbivores. In this way, climate cycle. However, increasingly sophisticated effects on one population can cascade statistical tools provide us with the means to do throughout a community. Climate and density so. In this thesis, I used state-of-the-art analytical also influenced reproduction and survival during approaches to provide a mechanistic and holistic the non-breeding season, via carryover effects. understanding of the impacts of climate change Overall, counteracting density, carryover and and density dependence on the population (direct and indirect) climate effects stabilised dynamics of a migratory Arctic herbivore, the breeding population size. In paper III, I analysed barnacle goose (Branta leucopsis). The study variation in an important life-history trait: age at population of barnacle geese breeds in high- first reproduction (AFR). As established in paper arctic Svalbard but spends the winter at Solway II, advancing spring onset increased the Firth, UK, with a spring stopover on mainland probability of females producing goslings in Norway. I investigated the dynamics of a local general, but importantly, this effect was stronger population breeding on western Svalbard, close for first-time than experienced breeders. The two- to the settlement of Ny-Ålesund, using 28 years week advance in spring phenology led to an of individual-based data. In paper I, I used an earlier AFR, by more than doubling the integrated population model to quantify
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