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Studies on the Diversity and Distribution of Marine Ichthyoparasites in Southern Africa Muhammad Azher Irfan Nunkoo A thesis submitted in fulfilment of the requirements for the degree of DOCTOR OF PHILOSOPHY Town in the Department of Biological Sciences Faculty of Science University of CapeCape Town of January 2019 University Supervisors: Dr. Cecile C. Reed 1, Assoc. Prof. Sven E. Kerwath 1, 2, Dr. Carl D. van der Lingen 1, 2, 3 1 Department of Biological Sciences, University of Cape Town, Private Bag X3, Rondebosch, 7701, South Africa 2 Fisheries Research and Development, Branch: Fisheries Management, Department of Agriculture, Forestry and Fisheries, Private Bag X2, Vlaeberg, 8018, Cape Town, South Africa 3 Marine Research Institute, University of Cape Town, Private Bag X3, Rondebosch, 7701, South Africa The copyright of this thesis vests inTown the author. No quotation from it or information derived from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes Capeonly. of Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University Declaration I know that plagiarism is wrong. Plagiarism is to use another‘s work and to pretend that it is one‘s own. I have used the Harvard convention for citation and referencing. Each contribution to and quotation in this project from the works of other people has been attributed, and has been cited and referenced. This project is my own work. I have not allowed, and will not allow, anyone to copy my work with the intention of passing it off as his or her own work. I acknowledge that copying someone else‘s work, or part of it, is wrong and I declare that this is my own work. M. A. Irfan Nunkoo January 2019 ii Table of Contents Table of Contents ..................................................................................................................... iii Abstract ..................................................................................................................................... ix Acknowledgements .................................................................................................................. xii Chapter 1. General Introduction ................................................................................................ 1 1.1. Marine Parasitology ........................................................................................................ 1 1.2. Marine Parasites: Life Cycles, Roles and Functions ...................................................... 2 1.3. Marine Parasites and Humans ......................................................................................... 5 1.3.1. Impacts of Marine Parasites on Humans ................................................................. 5 1.3.1.1. Implications in Capture Fisheries ..................................................................... 5 1.3.1.2. Implications for Human Health ........................................................................ 7 1.3.1.3. Implications for Aquaculture ............................................................................ 9 1.3.2. Impacts of Humans on Marine Parasites ............................................................... 10 1.3.2.1. Marine Fisheries.............................................................................................. 10 1.3.2.2. Marine Pollution ............................................................................................. 11 1.3.3. Applied Marine Parasitology ................................................................................. 12 1.3.3.1. Parasites as Biological Markers ...................................................................... 12 1.3.3.2. Parasites as Bioindicators ............................................................................... 13 1.4. Thesis Overview ........................................................................................................... 14 Chapter 2. Study Region: Currents, Oceanography and Biogeography .................................. 16 2.1. Currents of Southern Africa .......................................................................................... 16 2.1.1. The Benguela Current ............................................................................................ 16 2.1.2. The Agulhas Current .............................................................................................. 17 2.2. Marine Ecosystems of South Western Africa ............................................................... 18 2.2.1. The Northern Benguela .......................................................................................... 19 iii 2.2.2. The Southern Benguela .......................................................................................... 20 2.2.3. The South Coast ..................................................................................................... 21 2.2.4. The East Coast ....................................................................................................... 22 Chapter 3. A Checklist of Metazoan Parasites Associated with Marine Fishes of South Africa .................................................................................................................................................. 24 3.1. Introduction ................................................................................................................... 24 3.1.1. Marine Parasitology in South Africa ..................................................................... 24 3.1.2. Checklists: Benefits and Constraints ..................................................................... 26 3.1.3. Study Aims............................................................................................................. 27 3.2. Materials and Methods .................................................................................................. 27 3.2.1. Study Area ............................................................................................................. 27 3.2.2. Generating the Checklist ........................................................................................ 28 3.2.3. Data Analysis ......................................................................................................... 29 3.3. Results ........................................................................................................................... 30 3.3.1. Discovery, Diversity and Distribution ................................................................... 30 3.3.1.1. Ichthyoparasites .............................................................................................. 30 3.3.1.2. Host taxa ......................................................................................................... 32 3.3.1.3. Host-Parasite Pairs .......................................................................................... 34 3.3.1.4. Parasitological Publications ............................................................................ 35 3.3.1.5. Acanthocephala ............................................................................................... 47 3.3.1.6. Annelida .......................................................................................................... 48 3.3.1.7. Arthropoda ...................................................................................................... 49 3.3.1.8. Cnidaria ........................................................................................................... 51 3.3.1.9. Nematoda ........................................................................................................ 52 3.3.1.10. Platyhelminthes ............................................................................................. 53 3.4. Discussion ..................................................................................................................... 55 iv 3.4.1. Parasitological Research Effort .............................................................................. 56 3.4.2. Parasite Diversity and Distribution ........................................................................ 59 3.4.2.1. Acanthocephala ............................................................................................... 59 3.4.2.2. Annelida .......................................................................................................... 59 3.4.2.3. Arthropoda ...................................................................................................... 60 3.4.2.4. Cnidaria ........................................................................................................... 61 3.4.2.5. Nematoda ........................................................................................................ 62 3.4.2.6. Platyhelminthes ............................................................................................... 62 3.4.3. Current Challenges and Future Research ............................................................... 63 3.5. The Parasite-Host Checklist .......................................................................................... 65 Chapter 4. Community Ecology of Metazoan Parasite Communities of Selected Fishes of the Southern Benguela ................................................................................................................. 136 4.1. Introduction ................................................................................................................