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Thesis Text Inactive Mendeley Fields Final Edition Downloaded from orbit.dtu.dk on: Sep 23, 2021 Do traits follow taxonomy? Biodiversity and community assembly in marine ecosystems Dencker, Tim Spaanheden Publication date: 2019 Document Version Publisher's PDF, also known as Version of record Link back to DTU Orbit Citation (APA): Dencker, T. S. (2019). Do traits follow taxonomy? Biodiversity and community assembly in marine ecosystems. Technical University of Denmarik. General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. Users may download and print one copy of any publication from the public portal for the purpose of private study or research. You may not further distribute the material or use it for any profit-making activity or commercial gain You may freely distribute the URL identifying the publication in the public portal If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Do traits follow taxonomy? Biodiversity and community assembly in marine ecosystems Ph.D. thesis by Tim Spaanheden Dencker 2019 Supervisors: Martin Lindegren Mark R. Payne Peter Grønkjær ii Preface This thesis was submitted as part of the requirements to fulfil the Doctor of Philosophy Degree (PhD) at the Technical University of Denmark (DTU). The research presented in the thesis was carried out at the Centre for Ocean Life under the National Institute for Aquatic Resources (DTU Aqua) between September 2015 and February 2019 under supervision by Dr. Martin Lindegren (DTU Aqua), Dr. Mark R. Payne (DTU Aqua) and Dr. Peter Grønkjær (Aarhus University). In addition, two external research stays were carried out at Rutgers University, USA, in collaboration with Dr. Malin Pinsky and his research group, and at Oldenburg University, Germany, in collaboration with Dr. Helmut Hillebrand and Dr. Dorothee Hodapp. The Centre for Ocean Life is a VKR Center for Excellence, funded by the Villum Foundation. Additional funding for participation in conferences in South Korea, Latvia and Canada was received from the Otto Mønsted Fond. Kongens Lyngby, February 27 Tim Spaanheden Dencker Vector-based animal silhouettes used in figures and front cover art were obtained from phylopic.org under a Creative Commons Public Domain Dedication 1.0 License iii Acknowledgements ”To cope alone and unaided with a subject so vast, so complex, and so infinitely mysterious as the sea would be a task not only cheerless but impossible, and I have not attempted it.” Rachel L. Carson, The Sea Around Us, 1951. Even though a single name appears on the front cover of this thesis, I owe gratitude and thanks to several people who have contributed their time and energy in one way or another over the last three years to help me arrive at this point. First, Martin Lindegren for supporting me in every possible way throughout my time at DTU Aqua. Your enthusiasm and passion have been contagious, and you have shown me not only how to conduct science, but also how to balance a life in science. A great deal of appreciation for Mark R. Payne as well. Your knowledge and clear analytical mind have helped me when I have struggled with statistical analyses or coding. A thanks goes out to Peter Grønkjær as well. Throughout my time at the Centre for Ocean Life and DTU Aqua, I have been fortunate to work with a fantastic group of people. I will especially look back with fondness on the times spent with friends and colleagues in Martin Lindegren’s group, including Aurore, Esther, Neil, Anna, Laurene, Daniel and Jeremy. Our constant discussing and chatting, peaking over computer screens to share code, trait references and bounce ideas off each other will forever be an academic highlight of mine. It has been a privilege working with all of you on multiple collaborations and projects. To Mridul for being a kindred spirit when it came to having discussions outside our little bubble of trait-based sciences. To Lillian and Chris, two wonderful colleagues and people, who have helped me with countless tiny, yet challenging problems. A PhD has a way of seeping into every crevice of one’s life, and I owe my friends and family a great deal. To Sannyas, the best place to call home; to Ross, Peter, Ole, and Martin, I have a deep gratitude for your friendships; to Theis and Signe, your relationship, friendship and love is inspirational, and you have been solid anchors for the last three years. Bjørn and Frode, one day I will explain everything in this thesis to you. For now, it will have to be a boring bedtime story. Last, but not least, my mother and father for supporting me whenever I have faltered. I could not have done this without you! iv Abstract More than 2/3 of the Earth’s surface is covered by oceans, encompassing vast environmental gradients and variability in habitats. Thousands of species inhabit this seemingly endless space, yet we know surprisingly little about its stunning diversity, how it is shaped, assembled and structured. Consequently, understanding these mechanisms and how they vary in space and time is a key objective in ecology. These questions have traditionally been approached with a species-centric focus. However, this taxonomic approach is inherently limited, as the mere presence of a species reveals little to nothing about why the species is present and able to persist. Instead, species can be characterized by their traits, describing phenotypical characteristics that determine the species’ response to environmental conditions, its interactions in a food web and ultimately its effect on ecosystem functioning. This trait-based approach has emerged as a promising field of research allowing for a more causal and mechanistic understanding of marine biodiversity and ecosystems. In this thesis, I have explored marine demersal (bottom-living) fish communities across spatial and temporal scales in the North Atlantic and Northeast Pacific with regard to patterns and trends in biodiversity, community assembly processes, and environmental and anthropogenic drivers. Structuring of communities has been suggested to follow deterministic processes associated with responses to abiotic factors (environmental filtering) and interactions with other species (limiting similarity). An interplay between these two assembly processes shape and maintain community compositions. We used spatially and temporally resolved survey data on species abundances and traits to investigate temporal spatial patterns of species and trait diversity and the underlying community assembly mechanisms in the North Sea. Our results show that overall temporal trends in species richness and trait richness were highly correlated but varied considerably in space, indicating different degrees of either environmental filtering and limiting similarity acting on community assembly at local scales in the North Sea. In addition to deterministic assembly processes, communities can be structured by neutral, stochastic processes, such as dispersal, speciation, ecological drift and local extinctions. Deterministic and neutral theories have been considered antithetic to each other, despite a common historical origin. To study the effect and relative importance of these assembly processes acting on community composition across spatial scales we compiled a unique high-resolution dataset of scientific bottom- v trawl surveys from the North Atlantic and North-east Pacific, encompassing geo-referenced occurrences and abundances of >1000 species along with information on six life-history traits. Total taxonomic and trait beta-diversity, demonstrating the degree of similarity in species and trait composition between communities was calculated and compared at multiple spatial scales. Overall, our results show that both taxonomic and trait beta-diversity were mainly driven by deterministic assembly processes, primarily channeled through environmental filtering along gradients relating to temperature, depth and available energy. Human activities have left a considerable footprint on marine ecosystems worldwide with ensuing biodiversity loss at global scales. Halting further loss require quality assessment of biodiversity changes and consensus on how to measure such change. We tracked multiple components of biodiversity across time in seven marine ecosystems from both Atlantic and Pacific oceans, incorporating species, abundance and trait information for ~600 species. Despite no systematic trend in species richness over time, we observed significant directional changes in species identities and abundance structures over both short and long time-scales, while trait compositions remained relatively stable. These changes were linked to biotic homogenization, suggesting biotic impoverishment at local scales over time. The research presented in this thesis emphasizes the need to incorporate multiple components of biodiversity in spatial and temporal assessments of community changes and community assembly processes. The results are not only relevant for ecology research. The trait-based approach applied in this thesis provides important information furthering our mechanistic understanding and predictive capabilities; for use in research-based conservation and ecosystem-based management; and ultimately increasing our chances of halting further loss of biodiversity. vi Resumé
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