The Ecological and Functional Roles of Commercially Important Rabbitfish Species in a Marine Environment
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The ecological and functional roles of commercially important rabbitfish species in a marine environment Ameer Ebrahim BSc (Hons) A thesis submitted for the degree of Doctor of Philosophy at The University of Queensland in 2019 School of Biological Sciences Abstract Herbivorous fishes have consistently been found to have a significant positive impact on the resilience of coral reefs. Their feeding and foraging actions shape the capacity of reefs to both resist, and recover from, disturbance. Relatively little is known, however, about which ecological features in particular influence their distributions, movements, and foraging behaviours. Furthermore, the manner in which their biomass is altered by human actions remains poorly understood. Understanding and appreciating these key factors may help improve global management strategies of fisheries and coral reef habitats. Overall, this thesis provides a framework to establish the ecological functions of commercially important herbivorous fishes. This will not only help authorities establish effective management strategies to curb fishing pressure, but will also provide an indication of the capacity of reefs to resist or recover from disturbances that lead to algal dominance. At present, there is a noticeable gap in the literature pertaining to the ecological role that individual herbivorous fish species play; contemporary research instead focusses on their overall contribution. To bridge this gap and to render this research relevant to management efforts in the Seychelles, this thesis explored the distribution, ecological functions, and spatial ecology of key rabbitfish species in this archipelago of the Indian Ocean. Rabbitfishes are a crucial food source to the majority of the Western Indian Ocean region, comprising the bulk of their artisanal fisheries. In the Seychelles, in particular, they constitute over 60% of the total artisanal catch and, notably, there are no measures aimed at management of stocks currently being implemented or enforced. This thesis demonstrates that the overall distribution of herbivorous fishes is shaped by the habitat characteristics of individual reefs, and not by the presence of marine reserves. Species level analyses on rabbitfishes reveal that they were also influenced by the features of reef habitats, but not the conservation status of individual reefs. Furthermore, this research reveals that fishes from different functional groups occupy distinct reef habitats, suggesting that this may lead to spatial separation in the ii distribution of different forms of herbivory, and possibly reef resilience, in the Seychelles. This thesis investigated the ecological functions of rabbitfishes, focussing on their foraging rates and substrate preference, among diverse sites with varying benthic habitats. It reveals that rabbitfishes display both browsing and grazing traits, indicating possible functional complementarity between species. This is an important distinction as it implies that the conservation of different species may result in distinct shifts in the competitive dominance of coral and algae. Finally, this thesis examines the spatial ecology of the most commercially important fish species of the artisanal fishery in the Seychelles: the shoemaker spinefoot (Siganus sutor); it investigated whether they act as a mobile link between networked habitats, and whether their movements differed between day and night. Detection patterns reveals them to be diurnal herbivores, with only rare nocturnal movements. Furthermore, their movements are influenced by seagrass and coral abundance. Identification of such links between networked habitats are relevant for the purposes of implementing effective fisheries and habitat management strategies, and have a crucial bearing on sites around the archipelago which are to be declared marine reserves. iii Declaration by author This thesis is composed of my original work, and contains no material previously published or written by another person except where due reference has been made in the text. I have clearly stated the contribution by others to jointly-authored works that I have included in my thesis. I have clearly stated the contribution of others to my thesis as a whole, including statistical assistance, survey design, data analysis, significant technical procedures, professional editorial advice, financial support and any other original research work used or reported in my thesis. The content of my thesis is the result of work I have carried out since the commencement of my higher degree by research candidature and does not include a substantial part of work that has been submitted to qualify for the award of any other degree or diploma in any university or other tertiary institution. I have clearly stated which parts of my thesis, if any, have been submitted to qualify for another award. I acknowledge that an electronic copy of my thesis must be lodged with the University Library and, subject to the policy and procedures of The University of Queensland, the thesis be made available for research and study in accordance with the Copyright Act 1968 unless a period of embargo has been approved by the Dean of the Graduate School. I acknowledge that copyright of all material contained in my thesis resides with the copyright holder(s) of that material. Where appropriate I have obtained copyright permission from the copyright holder to reproduce material in this thesis and have sought permission from co-authors for any jointly authored works included in the thesis. iv Publications during candidature Peer-reviewed papers Ebrahim, A., Martin, T. S. H., Mumby, P. J., Olds, A. D., Tibbetts I. R. (2020) Differences in diet and foraging behaviour of commercially important rabbitfish species on coral reefs in the Indian Ocean. Coral Reefs. https://doi.org/10.1007/s00338-020-01918-6 Ebrahim, A., Bijoux, J. P., Mumby, P. J., Tibbetts I. R. (2020) The commercially important Shoemaker spinefoot, Siganus sutor connects coral reefs to neighbouring seagrass meadows. Journal of Fish Biology. https://doi.org/10.1111/jfb.14297 Conference abstracts Ebrahim, A., Bijoux, J. P., Mumby, P. J., Olds, A. D., Tibbetts I. R. Reef features, not marine reserves, shape the distribution of herbivorous fishes on coral reefs in the Seychelles (Oral presentation). Workshop on status of Seychelles coral reefs, reef resilience and coral reef restoration, Seychelles. Ebrahim, A., Bijoux, J. P., Mumby, P. J., Olds, A. D., Tibbetts I. R. Reef features, not marine reserves, shape the distribution of herbivorous fishes on coral reefs in the Seychelles (Oral presentation). The Eleventh Western Indian Ocean Marine Science Association (WIOMSA) Scientific Symposium, Mauritius. Manuscripts in prep for peer-review Ebrahim, A., Bijoux, J. P., Mumby, P. J., Olds, A. D., Tibbetts I. R. Reef features, not marine reserves, shape the distribution of herbivorous fishes on coral reefs in the Seychelles v Manuscripts in prep for peer-review included in this thesis Ebrahim, A., Bijoux, J.P., Mumby, P.J., Olds, A.D., Tibbetts I.R. (submitted to the Journal of Fish Biology) ‘Reef features, not marine reserves, shape the distribution of herbivorous fishes on coral reefs in the Seychelles This work was incorporated as Chapter 2 Contributor Statement of contribution Ameer Ebrahim Designed study (70%) (Candidate) Conducted fieldwork (80%) Conducted analysis (80%) Wrote and edited paper (70%) Jude P. Bijoux Designed study (20%) Wrote and edited paper (5%) Peter J. Mumby Designed study (5%) Conducted analysis (20%) Wrote and edited paper (5%) Andrew D. Olds Wrote and edited paper (10%) Rodney Melanie Conducted fieldwork (20%) Ian R. Tibbetts Designed study (5%) Wrote and edited paper (10%) vi Publications included in this thesis Ebrahim, A., Martin, T. S. H., Mumby, P. J., Olds, A. D., Tibbetts I. R. (2020) Differences in diet and foraging behaviour of commercially important rabbitfish species on coral reefs in the Indian Ocean. Coral Reefs. https://doi.org/10.1007/s00338-020-01918-6 This work was incorporated as Chapter 3 Contributor Statement of contribution Ameer Ebrahim Designed study (70%) (Candidate) Conducted fieldwork (80%) Conducted analysis (60%) Wrote and edited paper (70%) Tyson SH. Martin Conducted analysis (30%) Wrote and edited paper (5%) Peter J. Mumby Designed study (20%) Conducted analysis (10%) Wrote and edited paper (5%) Andrew D. Olds Wrote and edited paper (10%) Rodney Melanie Conducted fieldwork (20%) Ian R. Tibbetts Designed study (10%) Wrote and edited paper (10%) vii Ebrahim, A., Bijoux, J. P., Mumby, P. J., Tibbetts I. R. (2020) The commercially important Shoemaker spinefoot, Siganus sutor connects coral reefs to neighbouring seagrass meadows. Journal of Fish Biology. https://doi.org/10.1111/jfb.14297 This work was incorporated as Chapter 4 Contributor Statement of contribution Ameer Ebrahim Designed study (70%) (Candidate) Conducted fieldwork (60%) Conducted analysis (70%) Wrote and edited paper (75%) Jude P. Bijoux Designed study (20%) Conducted fieldwork (20%) Conducted analysis (20%) Peter J. Mumby Designed study (10%) Conducted analysis (10%) Wrote and edited paper (5%) Rodney Melanie Conducted fieldwork (10%) Ian R. Tibbetts Wrote and edited paper (20%) viii Contributions by others to the thesis Tibbetts IR, Mumby PJ, Bijoux JP, and Olds AD contributed to the conception and design of this research, advised on methods and analyses, and provided critical comments on the thesis and all the associated manuscripts that have been submitted for peer-review.