UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS Island Biogeography in the Anthropocene and Quaternary “ Documento Definitivo” Doutoramento em Biodiversidade, Genética e Evolução Sietze Johannes Norder Tese orientada por: Doutora Margarida Maria Demony de Carneiro Pacheco de Matos Doutor Paulo Alexandre Vieira Borges Doutor Kenneth Frank Rijsdijk Documento especialmente elaborado para a obtenção do grau de doutor 2020 UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS Island Biogeography in the Anthropocene and Quaternary Doutoramento em Biodiversidade, Genética e Evolução Sietze Johannes Norder Tese orientada por: Doutora Margarida Maria Demony de Carneiro Pacheco de Matos Doutor Paulo Alexandre Vieira Borges Doutor Kenneth Frank Rijsdijk Júri: Presidente: ● Doutor Rui Manuel dos Santos Malhó, Professor Catedrático, Faculdade de Ciências da Universidade de Lisboa Vogais: ● Doutor Christoph Küffer Schumacher, Professor, Institut Für Landschaft Und Freiraum da University of Applied Sciences Eastern Switzerland, (Suiça) ● Doutora Ana Isabel Costa Febrero Queiróz, Investigadora, Centro Interdisciplinar de História, Culturas e Sociedades - CIDEHUS da Universidade de Évora ● Doutor Paulo Alexandre Vieira Borges, Professor Auxiliar com Agregação, Faculdade de Ciências Agrárias e do Ambiente da Universidade dos Açores (Orientador) ● Doutora Angelica Crottini, Investigadora Auxiliar, Centro de Investigação em Biodiversidade e Recursos Genéticos – CIBIO/InBIO ● Doutor Jorge Manuel Mestre Marques Palmeirim, Professor Associado com Agregação, Faculdade de Ciências da Universidade de Lisboa ● Doutora Cristina Maria Branquinho Fernandes, Professora Associada com Agregação, Faculdade de Ciências da Universidade de Lisboa Documento especialmente elaborado para a obtenção do grau de doutor Financiado pela Fundação para a Ciência e a Tecnologia em projeto UID/BIA/00329/2013 e bolsa PD/BD/114380/2016 2020 NOTA PRÉVIA A presente tese apresenta resultados de trabalhos já publicados ou em preparação para publicação (Parte 1 e 2), de acordo com o previsto no nº 2 do artigo 25º do regulamento de Estudos Pós-graduados da Universidade de Lisboa, publicado no Diário de República II série nº 57 de 23 de Março de 2015. Tendo os trabalhos sido realizados em colaboração, o candidato esclarece que participou integralmente na conceção dos trabalhos, obtenção dos dados, análise e discussão dos resultados, bem como na redação dos manuscritos. Lisboa, Fevereiro de 2020 Sietze Johannes Norder Island Biogeography in the Anthropocene and Quaternary – PhD dissertation Sietze J. Norder i ACKNOWLEDGEMENTS I often find it hard to believe how lucky I am, to have received the opportunity to spend nearly four years on just one thing: on learning as much as possible about this fascinating world. Doing my PhD was a wonderful experience, but would not have been so much fun without the following people: I have been extremely fortunate with an amazing team of three wonderful supervisors. Guida was an incredible support over the previous years and I am greatly indebted to her for making everything run smoothly. I admire her bright mind and for the kind ways in which she helped me to reflect upon my work. I also thank her for reminding me of the importance to take a break and do something completely different. I am very grateful to Paulo for his daily guidance and for stimulating me to develop my own ideas. His battery seems always fully charged and the number of different things he manages to fit into one week is spectacular. Nonetheless, while juggling with many activities, he would always make time to have a conversation and discuss ideas. I also thank him for the off-road tours across the island, and for taking me with him to the field. I wish to thank Kenneth for his friendship and our refreshing walks together. Dr. Rijsdijk’s regular dose of humour helped me to always look on the bright side of (academic) life. His optimism and enthusiasm is unparalleled, and I highly value our talks about science, the universe, and everything. Furthermore, I would like to thank my collaborators, I have learnt a lot from them and I couldn’t have done my work without them. I also thank the reviewers of my papers who dedicated their free time to read my manuscripts, their constructive comments really helped me to improve my work. I thank my buddies and officemates from the BIODIV doctoral programme, the Azorean Biodiversity Group, and the TCE/BIOMAC group. My stay at the Azorean Biodiversity Group and the TCE/BIOMAC group was very beneficial for the progress of my research. I thank Paulo for inviting Maartje and me to Terceira, we had a great time living on this beautiful island. I also thank Willem Bouten and Daniel Kissling for receiving me as a guest and for welcoming me in their group. Of course, my dear friends in Portugal and the Netherlands, my brother and sisters, I thank you for the time we spent together, for all the little things that bring so much joy when shared. Island Biogeography in the Anthropocene and Quaternary – PhD dissertation Sietze J. Norder ii My loving parents, how can I ever thank you enough? For putting me on this beautiful world so full of mysteries, for the warm home to which I can always return to, for the interest and involvement with my activities, and your never-ending loving support. Maartje, four years ago we started this adventure together. We packed our bags and our bikes. We explored new places and met friends. We discovered new sounds, smells, and shades of light. It has been wonderful to make this journey together, I feel very lucky to travel with you, my loving companion. Island Biogeography in the Anthropocene and Quaternary – PhD dissertation Sietze J. Norder iii ABSTRACT The realization that human activities have a major influence on ecosystems from local to global scales has given rise to the concept of the Anthropocene. However, although the influence of human activities on biodiversity is clearly significant, it remains unclear to what extent the rate and magnitude of biodiversity changes differ from pre-human dynamics. Islands are ideal model systems for understanding the relative contribution of environmental and societal variables to biodiversity change because the onset of human activities on islands can generally be clearly defined. The aim of this PhD thesis is to place human-environment interactions on islands in the context of environmental fluctuations over the Quaternary. The thesis consists of two parts. First, I quantify how island area and isolation have changed over the Quaternary as a result of climate-driven sea-level fluctuations, and analyse how these dynamics have shaped modern biodiversity patterns. Secondly, I study how human activities in the past and present have shaped island ecosystems and landscapes, and compare their rate and magnitude to pre-human dynamics. Overall, the findings of my thesis indicate that modern biodiversity patterns show legacies of past human activities but are also imprinted by environmental dynamics in deep-time. Furthermore, the rate of change following human settlement on islands can largely exceed Quaternary background rates. Therefore, my findings are in line with studies that indicate that human activities have become a major driver in shaping biodiversity across scales. Nonetheless, my comparison of islands worldwide also highlights the diverse ways in which abiotic, biotic, and anthropogenic variables have interacted across individual islands. Therefore, future studies should acknowledge that global biodiversity change can manifest differently across localities. Finally, I emphasize the importance of strengthening interdisciplinary approaches in island biogeography to enhance our understanding of biodiversity changes in the Anthropocene, and how they relate to deep- time dynamics. Key words Volcanic oceanic islands, social-ecological systems, glacial-interglacial cycles, human ecodynamics, biodiversity Island Biogeography in the Anthropocene and Quaternary – PhD dissertation Sietze J. Norder iv RESUMO A constatação de que as atividades humanas exercem uma grande influência sobre os ecossistemas, da escala local à global, originou o conceito do Antropoceno. No entanto, apesar da influência das atividades humanas ser claramente significativa, ainda não está claro até que ponto a taxa e magnitude de alterações na biodiversidade diferem da dinâmica pré- humana. As ilhas constituem sistemas modelo ideais para compreender a contribuição relativa de variáveis ambientais e sociais, porque o início das atividades humanas é em geral conhecido. O objetivo desta tese de doutoramento é enquadrar as interações homem-ambiente nas ilhas no contexto das flutuações ambientais no Quaternário. A tese consiste em duas partes. Primeiro, quantifico como a área da ilha e o isolamento mudaram no Quaternário devido às flutuações do nível do mar provocadas pelo clima e analiso como essas dinâmicas moldaram os padrões modernos de biodiversidade. Em segundo lugar, estudo como as atividades humanas passadas e presentes moldaram os ecossistemas e paisagens das ilhas e comparo a sua taxa e magnitude com a dinâmica pré-humana. As conclusões de minha tese indicam que os padrões modernos de biodiversidade mostram legados de atividades humanas passadas, mas também são afectados pela dinâmica ambiental em escalas temporais longinquas. Além disso, a taxa de alterações após o povoamento humano nas ilhas pode exceder largamente taxas quaternárias antecedentes. Os meus resultados concordam com estudos mostrando que as atividades