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Eleanor Velasquez Thesis UNIQUE ISLAND HABITATS – A COMPARISON OF COMMUNITY ASSEMBLY IN MARINE AND TERRESTRIAL CONTEXTS Eleanor Margaret Velasquez Bachelor of Science (Botany and Ecology) The University of Queensland Submitted in fulfilment of the requirements for the degree of Doctor of Philosophy School of Earth, Environmental and Biological Sciences Science and Engineering Faculty Queensland University of Technology 2019 Keywords community assembly, critically endangered regional ecosystem, epibiont, urban forestry, general dynamic model of oceanic island biogeography, propagule pressure, pumice rafting, species–area relationship, theory of island biogeography. Unique island habitats – a comparison of community assembly in marine and terrestrial contexts i Abstract Biodiversity loss has far-reaching implications for the survival of both human and other species on Earth. Countless studies have linked biodiversity to essential processes or ‘ecosystem services’ such as the provision of fresh water and purification of air. As human-modified ecosystems are now estimated to dominate the Earth, with the loss of biodiversity being far greater than natural background processes. Understanding the abiotic (nonliving) and biotic (living) factors that influence the formation of communities over time and space has therefore never been more urgent. Thus, the main aim of the research presented in this thesis is to document and understand the abiotic and biotic factors driving the formation of insular biotic communities within two contexts. This PhD research also re- examines a fundamental theory in ecology, the theory of island biogeography (TIB), and its central premise the species–area relationship (SAR), as predictors of ecosystem community formation. The first insular community examined in this research was that of biota forming on floating pumice rafts in the Pacific Ocean. Each pumice stone was treated as a mini-island, as per the TIB and SAR. The patterns of richness forming on these stones were assessed against the key abiotic factors of age, area and climatic zone of collection, and the biotic factors of competition and facilitation. The second insular community examined was that of remnant forest fragments of the critically endangered regional ecosystem Melaleuca irbyana (R. T. Baker). Each remnant forest was treated as a human-made ii Unique island habitats – a comparison of community assembly in marine and terrestrial contexts island-like community as per the TIB and SAR, situated within the context of a highly modified human-made environment comprising either farmland or peri-urban environments. Here, the establishment success of M. irbyana seedlings was assessed against abiotic factors of remnant forest area and isolation, soil conditions and habitat disturbance (e.g. fire) and biotic factors of overstory and understory richness. The examination of these two, rarely studied and different communities for my PhD research, was done purposefully to provide an opportunity to compare the differences between the two contexts. Firstly, the examination of the two systems allowed contrasts to be made between the influence of abiotic and biotic predictors of community assembly in marine versus terrestrial contexts. Secondly, I examined the differences in abiotic and biotic predictors of community assembly in connection to one of the most fundamental theories in ecology the TIB and associated lines of thinking such as the SAR. This examination was particularly relevant to contributing new knowledge to my field of study, as the communities were formed via substantially different processes. Pumice rafted island-like communities are formed by the colonisation of marine animals and plants, under unassisted conditions, in the open ocean. Whereas, island-like forest fragments of M. irbyana represent the human-made remaining fragments of a native community that has been extensively cleared for farming and housing developments. In Chapter 3, I report that elements of the TIB, namely the SAR, are beneficial for predicting the richness forming on pumice stones as habitat area had one of the strongest correlations to species richness on pumice Unique island habitats – a comparison of community assembly in marine and terrestrial contexts iii clasts. However, pumice invertebrate communities were also found to change depending on the age of pumice clasts and climatic zone of collection. In Chapter 4, I found more evidence in support of the TIB and SAR within pumice rafted communities. As pumice community functional trait diversity was not only found to increase as epibiont richness increased, but also in relation to age and habitat area and then also altered depending on the most influential climatic zone encountered. In Chapter 5, I examined the connections between the TIB and SAR and establishment of M. irbyana seedlings into the understory of remnant forest areas. Here, my tests of the TIB and SAR (i.e. remnant area and isolation) did not directly aid in understanding the drivers of regeneration within this community. This was because seedling establishment of M. irbyana was linked to habitat disturbance and had little relationship to remnant forest area or isolation. However, the SAR did provide information on herbaceous native understory diversity, which increased with increasing remnant forest area. These research findings show that even the smallest remnant forests of M. irbyana provide increased chances of establishment success of this critically endangered community and, therefore, are worth preserving even when competing land uses such as development of housing estates, parks or roads threaten their conservation. The research conducted in this thesis contributes to understanding the dynamics of community assembly in different contexts and further the use of the TIB and its central principles the SAR in aiding this. Although the TIB and SAR provide useful predictive elements in some communities (e.g., pumice iv Unique island habitats – a comparison of community assembly in marine and terrestrial contexts rafting), they were not helpful for explaining the drivers behind regeneration in remnants of M. irbyana forests. These results are important because many studies simply examine insular communities in terms of their relationship to the SAR. However, even in the pumice rafted community where the TIB and SAR provided useful predictive elements, collection of additional abiotic and biotic data increased the power of models. For example, pumice community assembly was also influenced strongly by age, climatic zone of collection (as a measure of relative isolation) and the presence of foundation species such as barnacles of the genus Lepas. Overall, I find that age, isolation and area of ecosystems matter for species richness, functional trait diversity and specific needs of target species that may be critically endangered but that the ‘quality’ of an ecosystem is also important for predicting species establishment. Unique island habitats – a comparison of community assembly in marine and terrestrial contexts v Publications during candidature Peer-reviewed papers Eleanor Velasquez, Scott Bryan, Merrick Ekins, Alex G. Cook, Lucy Hurrey and Jennifer Firn. 2018. Age and area predict patterns of species richness in pumice rafts contingent on climatic zone encountered Ecology and Evolution 8: 5034-5046. https://doi.org/10.1002/ece3.3980. Conference abstracts Velasquez, E. M., Bryan, S. and Firn J. (2018) Plenary speed talk and poster: Pumice rafting: A hitchhiker's guide to marine biodiversity. Ecological Society of Australia, September 2018. Brisbane, Australia. Presented by Eleanor Velasquez. Velasquez, E. M., Bryan, S. and Firn J. (2016) Speed talk: Pumice rafting: A hitchhiker's guide to marine biodiversity. Society for Conservation Biology 4th Oceania Congress, July 2016. Brisbane, Australia. Presented by Eleanor Velasquez. Velasquez, E. M. and Firn, J. (2016) Poster presentation: Melaleuca irbyana: does remnant forest area drive seedling establishment in a critically vi Unique island habitats – a comparison of community assembly in marine and terrestrial contexts endangered tree? The 2nd International Conference on Urban Tree Diversity, February 2016. Melbourne, Australia. Presented by Eleanor Velasquez. Velasquez, E. M., Bryan, S. and Firn, J. (2015) Contributed Talk: Havre pumice rafts: A unique test of the theory of island biogeography. The Student Conference on Conservation Science, February 2015. The University of Queensland, St Lucia, Brisbane. Presented by Eleanor Velasquez. Velasquez, E. M., Bryan, S. and Firn, J. (2015) Contributed Talk: Havre pumice rafts: A unique test of the theory of island biogeography. The International Biogeography Society – 7th biennial Conference, January 2015. Bayreuth, Germany. Presented by Eleanor Velasquez. Publications included in this thesis I have chosen to incorporate one publication into my PhD thesis as follows: Eleanor Velasquez, Scott Bryan, Merrick Ekins, Alex G. Cook, Lucy Hurrey and Jennifer Firn. 2018. Age and area predict patterns of species richness in pumice rafts contingent on climatic zone encountered Ecology and Evolution 8: 5034-5046. https://doi.org/10.1002/ece3.3980. Contributor Statement of contribution Author Eleanor Velasquez (Candidate) Designed experiment (60%) Wrote and edited the paper (60%) Statistical analysis (70%) Unique island habitats – a comparison of community assembly in marine and terrestrial contexts vii Contributor Statement of contribution Conducted field and lab work (50%) Author Jennifer
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