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Intra- and interspecific variability in the distribution patterns and diet of the two most common catsharks caught in demersal trawls off the West and South coasts of South Africa: Evidence for habitat and resource partitioning? By Grant Mark van der Heever A thesis submitted in fulfilment of the requirements for the degree of Magister Scientiae in the Department of Biodiversity and Conservation Biology, University of the Western Cape. July 2017 i Intra- and interspecific variability in the distribution patterns and diet of the two most common catsharks caught in demersal trawls off the West and South coasts of South Africa: Evidence for habitat and resource partitioning? Grant Mark van der Heever Keywords: By-catch Catsharks Competition Demersal Diet Distribution Intraspecific Interspecific Resource Stable isotope analysis ii http://etd.uwc.ac.za/ Abstract Intra- and interspecific variability in the distribution patterns and diet of the two most common catsharks caught in demersal trawls off the West and South coasts of South Africa: Evidence for habitat and resource partitioning? G.M van der Heever MSc Thesis, Department of Biodiversity and Conservation Biology, University of the Western Cape. The objective of this thesis was to elucidate intra- and interspecific variability in the distribution patterns and diet of the two most common demersal catsharks taken as by-catch by the commercial trawling industry targeting the two Cape hake species off the West and South coasts of South Africa. Samples and data were collected during routine demersal hake biomass surveys conducted by DAFF, with distribution data collected from 1994 to 2015 and stomach content and stable isotope data collected from 2014 to 2015. On examination of the distribution data, Holohalaelurus regani was found to be more abundant on the West Coast and Scyliorhinus capensis was found to be more abundant on the South Coast. Both catsharks were observed to display size-based segregations, with catshark size increasing with depth in both species. Differences in the distribution patterns of male and female H. regani were also noted, with female catsharks inhabiting inshore areas and male catsharks inhabiting offshore areas. The two catsharks appeared to display high levels of dietary overlap, with individuals feeding on the most abundant crustaceans and cephalopods caught as by-catch in the trawl on each coast. However, iii http://etd.uwc.ac.za/ although both catsharks consumed similar prey groups across the coast, the abundances of the individual prey species in their diets appeared to differ. Ontogenetic shifts in the diets of both catshark species were also noted, with small catsharks being observed to primarily feed on small prey items, and large catsharks being observed to primarily feed on large prey items. A similar trend was noted for stable isotope data, with δ15N isotope values increasing with catshark size in both species. Differences in δ15N isotope values were noted between coasts as well, with both catshark species displaying significantly higher δ15N values on the West Coast than on the South Coast. The results appear to indicate a strong relationship between habitat and diet, with food separation appearing to be largely a reflection of habitat separation. Based on this, I argue that these morphologically similar catshark species partition their habitat and food resources to limit co-existence, and thus reduce the intensity of intra-and interspecific competition between them. Alternatively, these differences may also be a reflection of differences in environmental tolerance owing the unique environmental and biological factors evident off each coast; or they may simply be due to size-based changes in habitat use that may be associated with parturition or nursery grounds, changes in the habitat of their preferred diet items, or a means to prevent predation by larger sharks. iv http://etd.uwc.ac.za/ DECLARATION I declare that: Intra- and interspecific variability in the distribution patterns and diet of the two most common catsharks caught in demersal trawls off the West and South coasts of South Africa: Evidence for habitat and resource partitioning? is my own work, that it has not been submitted for any degree or examination in any other university, and that all the sources I have used or quoted have been indicated and acknowledged by complete references. Grant Mark van der Heever July 2017 Signed: v http://etd.uwc.ac.za/ Acknowledgements Firstly, I would like to express my sincere gratitude to my supervisor, Professor Mark John Gibbons, for his continuous support and patience throughout this thesis. He gave me the opportunity to fulfil my childhood dream, and I am forever grateful. I would also like to express my thanks to my co-supervisors, Dr Carl van der Lingen and Dr Rob Leslie, for their insightful comments, knowledge and guidance. Their expertise and passion for marine science most certainly increased my love for marine life. Besides my supervisors, I would like to acknowledge and thank the Department of Agriculture, Forestry and Fisheries (DAFF) demersal team, which includes; Dr Deon Durholtz, Robert Cooper, George Julius, George Kant, Clifford Hart, James Peterson and Joe Mhlongo. Their support and guidance, and enthusiasm and humour made the month long research surveys something to look forward to. Special thanks are also due to the captain, officers and crew of FV Andromeda; thanks for your hospitality and safe ventures. Additionally, I would like to thank Tracey Fairweather (DAFF) for her technical support with GIS, as well as Ian Newton and John Lanham from the light stable isotope unit at the University of Cape Town, for processing my muscle tissue samples. I am also grateful to Dr Lara Atkinson (SAEON) for encouraging me to persevere and write. Special thanks are also due to DAFF and NRF for financial support. Most of all, I would like to thank my family, Joy, Mark and Tammy for their love and support. They gave me the opportunity to achieve my life-long dream of vi http://etd.uwc.ac.za/ becoming a marine biologist, and I will forever be indebted to them. Last but not least, I would like to thank my partner, friend and ad-hoc research assistant, Danielle Laubscher, for her love and patience. Thank you for the encouragement, motivation and for putting up with my constant “does this sentence make sense to you” question. It really meant a lot, and I thank you for that. vii http://etd.uwc.ac.za/ Table of Contents Chapter One: General Introduction ................................ ................................................................... 1 Chapter Two: Methods ................................................................ ...................................................... 15 General Methods ............................................................................................................................ 15 Study location .............................................................................................................................. 15 Survey design .............................................................................................................................. 15 Analyses ...................................................................................................................................... 17 Distribution Chapter Methods ...................................................................................................... 18 Study location and data collection ............................................................................................... 18 Data analysis................................................................................................................................ 18 Diet Chapter Methods ................................................................................................................... 19 Study location .............................................................................................................................. 19 Sample collection ........................................................................................................................ 20 Prey identification ....................................................................................................................... 20 Analysis of diet ............................................................................................................................ 21 Statistical analyses ....................................................................................................................... 23 Stable Isotope Chapter Methods ................................................................................................... 25 Study location .............................................................................................................................. 25 Data collection ............................................................................................................................. 25 Laboratory analysis ..................................................................................................................... 26 Analysis of stable isotope data .................................................................................................... 27 Statistical analyses ....................................................................................................................... 27 Chapter Three: Spatial and ontogenetic variability in the distribution pattens of two co- occurring