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Impacts of Climate Change on Woodland Birds; from Individual Behaviour to Population

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Impacts of Climate Change on Woodland Birds; from Individual Behaviour to Population Impacts of Climate Change on Woodland Birds; From Individual Behaviour to Population Change Thesis submitted in part fulfilment of PhD Jeremy Alexander Smith September 2019 Acknowledgements I would first like to thank my family who have not only supported me through the last four years but for my whole journey to get here. This thesis is dedicated to my mum Penny Smith who has been a rock throughout my life and encouraged me with my decisions even when they have seemed completely impossible or down right stupid. Similar thanks must be given for my supervisors. Dr Rob Thomas has known me since 2006 when he introduced me to Pied Flycatchers, ever since then I haven’t been able to leave them alone and nor the dataset from Cwm Clydach. His continuous pastoral, emotional and supervisory care has been essential to me getting here. I would also like to thank Drs Hefin Jones and Ian Vaughan for all of their personal and professional guidance through the years and the many hours they have spent reading and commenting on chapters. Thanks also go to Dr Sean Rands for his input in the creation of the project, grant application and his help during the project. Although not formally a supervisor, Dr Fred Slater’s assistance throughout the field seasons was invaluable and his kind training of me was exceptionally gratefully received. This thesis would not have been possible without the data so graciously provided by the Gower Ornithological Society and all of the field work conducted by the volunteers over many years, thank you! My particular gratitude to Heather Coats who has been a contact and advisor along the way. The access to Tŷ Mawr Farm by the friendly and kind farmers who own and work the land was very much appreciated. Access to land and conduct recording was also permitted by the Wildlife Trust of South and West Wales even though the data were not used here. Thanks to my office mates and pub mates who took me out for walks towards the end and made me laugh all the way through. Particular mention goes to Rich Facey, Jen Stockdale, Meg and Luke Evans who have been there all along and to Sophie Lee Lane and Mike Shewring who joined the ride a little later! Thanks to my students who helped collect the field data and kept me company during the seasons. My mates have made a huge difference to my time picking me up when I was down, listening to my whinging and moaning, particular mention goes to Ross and Rebecca Price. They were also there to share in the triumphs. My gratitude goes to NERC specifically the GW4+ DTP for funding this project which was hosted by Cardiff University. ii Table of Contents 1 GENERAL INTRODUCTION .............................................................................................................. 2 1.1 MULTI-TROPHIC PHENOLOGICAL MATCHING ........................................................................................... 2 1.2 CLIMATE CHANGE EFFECTS ON PHENOLOGY ............................................................................................ 3 1.3 SPATIAL AND TEMPORAL ADAPTATIONS TO CLIMATE CHANGE............................................................. 4 1.4 DIFFERING BIRD MIGRATION STRATEGIES AND THEIR SENSITIVITIES TO CLIMATE ........................... 5 1.5 TRI-TROPHIC WOODLAND ECOSYSTEM (OAKS, CATERPILLARS AND PIED FLYCATCHERS) ............. 6 1.6 THE PIED FLYCATCHER AS A MODEL STUDY SPECIES ............................................................................. 7 1.7 POPULATION ABUNDANCE AND DECLINES IN PIED FLYCATCHERS ....................................................... 8 1.8 MODELLING OF PATTERNS AND MECHANISMS OF POPULATION CHANGE ............................................ 9 1.9 PREDICTING POPULATION TRAJECTORIES OF A MIGRATORY PASSERINE USING AN INTEGRAL PROJECTION MODEL ................................................................................................................................................ 11 1.10 THESIS STRUCTURE AND HYPOTHESES ................................................................................................... 12 2 POPULATION DECLINE AND DEMOGRAPHIC CHANGES IN A BREEDING POPULATION OF PIED FLYCATCHERS IN SOUTH WALES OVER 26 YEARS ......................................................13 2.1 ABSTRACT ................................................................................................................................................... 13 2.2 INTRODUCTION ........................................................................................................................................... 14 2.3 METHODS .................................................................................................................................................... 18 2.3.1 Study Site ................................................................................................................................................. 18 2.3.2 Data Collection ...................................................................................................................................... 18 2.3.3 Weather data ......................................................................................................................................... 19 2.3.4 Statistical Analysis ............................................................................................................................... 20 2.3.5 Principal Component Analysis ....................................................................................................... 20 2.3.6 Temporal trends ................................................................................................................................... 21 2.3.7 Weather impacts on demography ................................................................................................ 22 2.4 RESULTS ...................................................................................................................................................... 23 2.4.1 Population size and nest box occupancy ................................................................................... 23 2.4.2 Inter-annual variation in April weather and laying date ................................................. 24 2.4.3 Effects of weather on fledging success ....................................................................................... 28 2.5 DISCUSSION ................................................................................................................................................. 32 2.5.1 Population change in a global, long-term & multi-species context .............................. 32 2.5.2 Inter-annual variation in weather conditions ........................................................................ 32 2.5.3 Changes in the timing of breeding ............................................................................................... 33 2.5.4 Variation in clutch size ...................................................................................................................... 34 2.5.5 Variation in hatching success, brood size and number of fledged chicks .................. 34 iii 2.5.6 Associations between weather and breeding parameters ................................................ 35 2.5.7 Drivers of population change ......................................................................................................... 36 2.5.8 The value of long-term studies for understanding the climatic and demographic drivers of population change .............................................................................................................................. 37 3 PHENOLOGY OF INVERTEBRATE ABUNDANCE AND PIED FLYCATCHER BREEDING BIOLOGY -CAUSES AND CONSEQUENCES OF TROPHIC MIS-MATCHES ..................................39 3.1 ABSTRACT ................................................................................................................................................... 39 3.2 INTRODUCTION ........................................................................................................................................... 40 3.3 METHODS .................................................................................................................................................... 44 3.3.1 Study Site ................................................................................................................................................. 44 3.3.2 Weather .................................................................................................................................................... 45 3.3.3 Invertebrates .......................................................................................................................................... 45 3.3.4 Birds ........................................................................................................................................................... 46 3.3.5 Statistical analysis ............................................................................................................................... 47 3.4 RESULTS ...................................................................................................................................................... 50 3.4.1 Weather and invertebrates ............................................................................................................. 53 3.4.2 Bird breeding parameters and phenology ............................................................................... 55 3.4.3 Variation
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