Taxonomy, biodiversity and biogeography of the brittle stars (Echinodermata: Ophiuroidea) of South Africa. Jennifer M. Olbers Thesis presented for the degree of Doctor of Philosophy Universityin the Department of of Biological Cape Sciences Town University of Cape Town November 2016 Supervisor: Prof. Charles L. Griffiths University of Cape Town Co-supervisor: Dr Yves Samyn Royal Belgian Institute of Natural Sciences The copyright of this thesis vests in 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 only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University of Cape Town ii “The sea, once it casts its spell, holds one in its net of wonder forever” - Jacques Yves Cousteau “…echinoderms, a noble group especially designed to puzzle the zoologist”. - Libbie Henrietta Hyman iii DECLARATION - PLAGIARISM & FREE LICENCE I, Jennifer M. Olbers, declare that: (a) I know the meaning of plagiarism and declare that all of the work in the thesis, with the exception of which is properly acknowledged, is my own; (b) the above thesis is my own unaided work, both in conception and execution, and that apart from the normal guidance from my supervisors, I have received no assistance except as stated below; (c) neither the substance nor any part of the thesis has been submitted in the past, or is being, or is to be submitted for a degree at this University or at any other University, except as stated below; (d) the University is granted free licence to reproduce the above thesis in whole or in part, for the purpose of research. (e) Several of the chapters in this thesis have appeared in published form, in co-authorship with the other researchers listed below. Below is an account of the contributions made by the candidate, and each of the co-authors, to each of these jointly-published papers / chapters. Author abbreviations: JO: Jennifer M. Olbers YS: Yves Samyn MO: Masanori Okanishi CG: Charles L. Griffiths FR: Frank W.E. Rowe TF: Toshihiko Fujita Chapter 2 published as: Olbers, J.M., Samyn, Y. and Griffiths, C.L. 2015. New or notable records of brittle stars (Ophiuroidea: Echinodermata) from South Africa. African Natural History, 11: 83-116. Author contributions: JO conceived the paper with CG. JO collected the data, examined the specimens, analysed the data and wrote the initial paper. CG and YS contributed valuable comments to the manuscript. Chapter 3 published as: Olbers, J.M. and Samyn, Y. 2012. The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa. Western Indian Ocean Journal of Marine Science, 10(2): 137-154. Author contributions: JO conceived the paper with YS. JO and YS collected data, examined and identified specimens and analysed the data. JO wrote the initial manuscript with YS contributing comments and valuable input. Chapter 4 published as: Olbers, J.M., Rowe, F.W.E., Griffiths, C.L. and Samyn, Y. 2014. The rediscovery of a collection of echinoderms including two holotypes in the Durban Natural Science Museum, South Africa. Novitates, 36: 1-13. Author contributions: JO conceived the paper with FR. JO collected data. JO and FR examined and identified specimens. JO analysed data and wrote the initial paper. CG, FR and YS contributed valuable comments to the manuscript. iv Chapter 5 published as: Okanishi, M, Olbers, J.M. and Fujita, T. 2013. A taxonomic review of the genus Asteromorpha Lütken (Echinodermata: Ophiuroidea: Euryalidae). The Raffles Bulletin, 61(2): 461-480. Author contributions: MO conceived the paper. JO and MO collected data. JO and MO examined and identified specimens. JO and MO analysed data and wrote the paper. TF contributed valuable comments to the manuscript. _________________ Jennifer M. Olbers November 2016 v TABLE OF CONTENTS ACKNOWLEDGMENTS......................................................................................................................................viii PHOTOGRAPHIC CREDITS..............................................................................................................................xi ABBREVIATIONS..............................................................................................................................................xii ABSTRACT..........................................................................................................................................................1 CHAPTER 1: GENERAL INTRODUCTION..........................................................................................................3 BACKGROUND TO ECHINODERMATA AND OPHIUROIDEA..............................................................3 GENERAL TERMINOLOGY AND MORPHOLOGY OF OPHIUROIDEA.................................................4 AIMS AND OBJECTIVES........................................................................................................................9 OUTLINE OF CHAPTERS.......................................................................................................................9 CHAPTER 2: NEW OR NOTABLE RECORDS OF BRITTLE STARS (ECHINODERMATA: OPHIUROIDEA) FROM SOUTH AFRICA........................................................................................................................................12 ABSTRACT............................................................................................................................................12 INTRODUCTION...................................................................................................................................12 MATERIALS AND METHODS................................................................................................................13 RESULTS..............................................................................................................................................14 TAXONOMIC ACCOUNT..........................................................................................................17 Family ASTEROSCHEMATIDAE Verrill, 1899.............................................................17 Family GORGONOCEPHALIDAE Ljungman, 1867.....................................................18 Family OPHIOMYXIDAE Ljungman, 1867...................................................................20 Family OPHIURIDAE Lyman, 1865...............................................................................21 Family AMPHIURIDAE Ljungman, 1867......................................................................24 Family OPHIACTIDAE Matsumoto, 1915.....................................................................27 Family OPHIOCOMIDAE Ljungman, 1867...................................................................29 Family OPHIOTRICHIDAE Ljungman, 1867................................................................32 Family OPHIACANTHIDAE Ljungman, 1867................................................................38 Family OPHIODERMATIDAE Ljungman, 1867............................................................40 Family OPHIOLEPIDIDAE Ljungman, 1867.................................................................47 DISCUSSION........................................................................................................................................48 ACKNOWLEDGMENTS........................................................................................................................48 PLATES.................................................................................................................................................49 CHAPTER 3: THE OPHIOCOMA SPECIES (OPHIURIDA: OPHIOCOMIDAE) OF SOUTH AFRICA..............................................................................................................................................................58 ABSTRACT............................................................................................................................................58 INTRODUCTION...................................................................................................................................58 MATERIALS AND METHODS................................................................................................................59 TAXONOMIC ACCOUNT......................................................................................................................60 Ophiocoma brevipes Peters, 1851 [=Ophiocoma (Breviturma) brevipes].....................................61 Ophiocoma dentata Müller and Troschel 1842 [=Ophiocoma (Breviturma) dentata]..................63 Ophiocoma doederleini de Loriol, 1899 [=Ophiocoma (Breviturma) doederleini].......................64 Ophiocoma erinaceus Müller and Troschel 1842.......................................................................65 Ophiocoma pica Müller and Troschel 1842................................................................................66 Ophiocoma pusilla (Brock, 1888)...............................................................................................67 Ophiocoma scolopendrina (Lamarck, 1816) ..............................................................................68 Ophiocoma valenciae Müller and Troschel 1842.......................................................................69 DISCUSSION.........................................................................................................................................70 ACKNOWLEDGMENTS........................................................................................................................70