Relative Effects of Environmental Variation and Fishing on the Demography and Ecology of Tropical Wrasses
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ResearchOnline@JCU This file is part of the following work: Lowe, Jake R. (2020) Relative effects of environmental variation and fishing on the demography and ecology of tropical wrasses. PhD Thesis, James Cook University. Access to this file is available from: https://doi.org/10.25903/nq18%2Dqr71 Copyright © 2020 Jake R. Lowe. The author has certified to JCU that they have made a reasonable effort to gain permission and acknowledge the owners of any third party copyright material included in this document. If you believe that this is not the case, please email [email protected] Relative effects of environmental variation and fishing on the demography and ecology of tropical wrasses Thesis submitted by Jake R. Lowe BSc, GCertResMeth in November 2020 for the degree of Doctor of Philosophy at the College of Science and Engineering, and the ARC Centre of Excellence for Coral Reef Studies, James Cook University EFFECTS OF ENVIRONMENT AND FISHING ON WRASSES Acknowledgements Foremost, I thank my primary supervisor Garry Russ, I can’t express how grateful I am for his excellent supervision and mentorship over the years. I am also indebted to my secondary supervisor Howard Choat, his enthusiasm, expertise, and perspective has been instrumental in my development as a scientist. The quality of this thesis would not have been possible without Garry and Howards advice and support throughout the entire process. I would also like to thank David Williamson, Daniella Ceccarelli, and Richard Evans for their time, energy, and advice which contributed to the quality of this thesis. I particularly want to express my gratitude to David Williamson for the use of his unpublished data. Immense thanks to Rene Abesamis and Abner Bucol for facilitating my work in the Philippines, the laughs, and most of all the correspondence of data between the Philippines and Australia that ensured the completion of this thesis. I look forward to working with you all again in the future. I am very grateful to have received an Australian Commonwealth Government Research Training Program Scholarship from November 2016 to May 2020. I am also grateful to the College of Science and Engineering for providing me scholarship from May 2020 to October 2020, when I encountered delays due to the global COVID-19 pandemic. Research for this thesis was supported by a grant to Garry Russ from the Australian Research Council Centre of Excellence for Coral Reef Studies and a grant from the College of Science and Engineering. Thanks to the many field and technical staff that have enabled this project. A special thank you to Brock Bergseth, Gerard Ricardo, Haydn Turner, Kyle Hillcoat, Katie Sievers, Sam Smith, Roxie Diaz, and Ramon Dapat Jr. for their outstanding assistance in the field. Thanks also to Dong Chun Lou and Sue Reilly for their expertise and advice in the laboratory. Invaluable logistical support in the Philippines was provided by SUAKCREM. I am particularly grateful to Dr. Alcala and Ma’am Emily, thank you for looking after me during my time in Dumaguete. In Australia, I thank the staff of JCU boat and dive, Orpheus Island Research Station, and the Explore Group Hamilton Island for their assistance in the field. A huge heartfelt thanks to my parents, Linda and John, for always supporting me and encouraging me to follow my dreams. Thanks also to my sister, grandparents, and loyal friends. Finally, a very special thank you to my partner Portia for her love and support throughout the course of this project. i EFFECTS OF ENVIRONMENT AND FISHING ON WRASSES Statement on the Contribution of Others This thesis was funded by an Australian Research Council (ARC) grant to Prof. Garry R. Russ, through the ARC Centre of Excellence for Coral Reef Studies and a grant from the College of Science and Engineering, James Cook University. I received an Australian Commonwealth Government Research Training Program Scholarship from November 2016 to May 2020. I also received a scholarship from the College of Science and Engineering from May 2020 to October 2020, when I encountered delays due to the global COVID-19 global pandemic. I performed Chapters 2 and 3 of this thesis in collaboration with colleagues David Williamson, Daniela Ceccarelli, and Richard Evans. The long-term data used in these chapters were collected through David Williamson’s long-term monitoring program “Assessing the ecological effects of management zoning on inshore reefs of the Great Barrier Reef Marine Park”. I assisted this monitoring program as a benthic observer from 2016 – 2018. Data from 2006 – 2014 used in chapters 2 and 3 were collected without my assistance. David, Daniela, and Richard all appear as co-authors on published versions of Chapter 2 and 3 which appear in the Journals Marine Biology and Environmental Biology of Fishes, respectively. Funding for monitoring field work that contributed to chapters 2 and 3 of this thesis was provided via grants to David Williamson by the Australian Research Council (ARC); the CRC Reef Research Centre; and the Australian Government Department of Environment and Energy through the Marine and Tropical Sciences Research Facility (MTSRF), the National Environmental Research Program (NERP), the National Environmental Science Program (NESP), and the Reef 2050 Integrated Monitoring and Reporting Program (RIMReP). Field work in the Philippines which contributed to Chapters 4 and 5 of this thesis were made possible thanks to a memorandum of agreement established between James Cook University, Silliman University Angelo King Centre for Research and Environmental Management (SUAKCREM), and the Philippine Bureau of Fisheries and Aquatic Resources (BFAR). This was facilitated via colleagues Rene Abesamis and Angel Alcala at SUAKCREM. Additionally, Abner Bucol interpreted histological data of Philippine wrasses and corresponded data to Australia when my field trip was cancelled due to the COVID-19 global pandemic. Both Abner Bucol and Rene Abesamis will appear as named authors on the separate publication of Chapters 4 and 5 from this thesis. ii EFFECTS OF ENVIRONMENT AND FISHING ON WRASSES Publications Arising from this Thesis 1. Lowe JR, Williamson DH, Ceccarelli DM, Evans RD & Russ GR (2019) Responses of coral reef wrasse assemblages to disturbance and marine reserve protection on the Great Barrier Reef. Marine Biology 166, 119. doi:10.1007/s00227-019-3566-5. 2. Lowe JR, Williamson DH, Ceccarelli DM, Evans RD & Russ GR (2020) Environmental disturbance events drive declines in juvenile wrasse biomass on inshore coral reefs of the Great Barrier Reef, Australia. Environmental Biology of Fishes 103, 1279-1293. doi:10.1007/s10641-020-01022-2. 3. Lowe JR, Russ GR, Bucol AA, Abesamis RA & Choat JH (in prep.) Spatial variability in the early gonadal development and sexual ontogeny of Hemigymnus, Cheilinus, and Oxycheilinus wrasses among Indo-Pacific coral reefs. Journal of Fish Biology. 4. Lowe JR, Russ GR, Bucol AA, Abesamis RA & Choat JH (in prep.) Spatial variation in the demography of fished and unfished Hemigymnus, Cheilinus, and Oxycheilinus wrasse populations among Indo-Pacific coral reefs. Journal of Animal Ecology. iii EFFECTS OF ENVIRONMENT AND FISHING ON WRASSES Abstract One of the greatest challenges faced by fishery managers and ecologists is determining the degree to which natural variation in population dynamics (i.e., species abundance and life history traits) are modified by fishing. Multiscale sampling that encompasses temporal and spatial variation in anthropogenic and environmental factors are key to understanding mechanisms that drive population dynamics. To date, our understanding of these processes has been hindered by a lack of spatially explicit demographic information for many fishery- targeted species. This is particularly true for multi-species, Indo-Pacific coral reef fisheries that are under increasing levels of exploitation. For example, across Indo-Pacific reefs, large- bodied wrasses are common components of reef fish assemblages and are exposed to increasing levels of exploitation, yet little is known regarding their life histories or drivers of population variability. This thesis was designed to assess the relative effects of environmental variation and fishery exploitation on the demography and ecology of tropical wrasses both spatially and temporally, to determine patterns of demographic variation and population drivers among and within species. To assess the relative importance of fishing and environmental factors influencing population dynamics, here I combine data from long-term monitoring of wrasse populations across marine reserve networks of the Great Barrier Reef (GBR), Australia with multiscale, age- based comparisons of sexual ontogenies and age-based demography of the fishery-targeted wrasses Hemigymnus melapterus, H. fasciatus, Cheilinus fasciatus, and Oxycheilinus digramma collected from low-latitude Philippine fish markets and from two unfished populations on the Great Barrier Reef. Firstly, in Chapter 2, I analyse a long-term (2006 – 2018), spatially extensive (≈ 700 km) ‘natural experiment’ quantifying the responses of 11 wrasse taxa to acute environmental disturbance events, predator density, and no-take marine reserve (NTMR) protection on fringing coral reefs in the Palm (18°34′ S, 146°29′ E), Whitsunday (20°08′ S, 148°56′ E), and Keppel Island (23°10′ S, 150°57′ E) groups, Great Barrier Reef, Australia. The responses of wrasse densities to benthic habitat change were taxa-specific and temporally consistent. Disturbance- mediated