Towards an Unravelling of the Taxonomy of Chrysaora (Scyphozoa; Semaeostomeae;

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Towards an Unravelling of the Taxonomy of Chrysaora (Scyphozoa; Semaeostomeae; Towards an unravelling of the taxonomy of Chrysaora (Scyphozoa; Semaeostomeae; Pelagiidae) from around South Africa VERENA RAS Department of Biodiversity & Conservation Biology, University of the Western Cape, Private Bag X17, Bellville, 7535, South Africa A THESIS SUBMITTED IN FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE Supervisor: Professor Mark J. Gibbons Co-supervisor: Dr. Adriaan Engelbrecht November 2016 i I declare that “Towards an unravelling of the taxonomy of Chrysaora (Scyphozoa; Semaeostomeae; Pelagiidae) from around South Africa” is my own work, that it has not been submitted for any degree or examination at any other university, and that all the sources I have used or quoted have been indicated and acknowledged by complete references. Signature: Date: 06/02/2017 ii TABLE OF CONTENTS Acknowledgements ................................................................................................ .................... 1 Section A: Towards an unravelling of the taxonomy of Chrysaora (Scyphozoa; Semaeostomeae; Pelagiidae) from around South Africa .......................................... 2 Abstract ................................................................ ...................................................................... 2 Chapter 1: Introduction .............................................................................................................. 3 Chapter 2: Materials and Methods ........................................................................................... 14 2.1 Morphology .................................................................................................... 14 2.2 Cnidome ......................................................................................................... 18 2.3. DNA Analyses ............................................................................................... 20 Chapter 3: Results .................................................................................................................... 24 3.1 Quantitative Results........................................................................................ 24 3.2 Comparitive Statistics ..................................................................................... 26 3.3 Genetic Analyses ............................................................................................ 31 3.4 Summary......................................................................................................... 34 Chapter 4: Formal descriptions ................................................................................................ 35 4.1 Chrysaora agulhensis sp. nov. ....................................................................... 35 4.2 Descriptions of Additional Taxonomically Important Features for C. fulgida and C. africana .................................................................................................... 41 Chapter 5: Remarks.................................................................................................................. 44 References ................................................................................................................................ 53 Tables ....................................................................................................................................... 70 Figure Headings ....................................................................................................................... 85 Figures...................................................................................................................................... 90 Appendices ............................................................................................................................. 113 Section B: The identification of a South African Rhizostoma (Scyphozoa: Rhizostomeae) 126 Abstract .................................................................................................................................. 126 Chapter 1: Introduction .......................................................................................................... 127 Chapter 2: Materials and Methods ......................................................................................... 131 2.1 Specimen Collection .................................................................. 131 2.2 Morphological Data Analyses ................................................... 132 2.3 DNA Analyses ........................................................................... 134 iii Chapter 3: Descriptions.......................................................................................................... 137 3.1 Rhizostoma sp. 1 ........................................................................ 137 3.2 Rhizostoma sp. 2 ........................................................................ 141 Chapter 4: Remarks................................................................................................ ................ 144 References ................................................................ .............................................................. 149 Tables ..................................................................................................................................... 156 Figure Headings ..................................................................................................................... 165 Figures.................................................................................................................................... 168 Appendices ............................................................................................................................. 178 iv Acknowledgements Sincere and special thanks are given to my parents, Verena and Jerome Ras, who have provided the encouragement, support and help throughout all difficulties and emotional trials experienced throughout this degree. To my siblings for accepting my incredibly erratic working hours which prevented them from functioning optimally. To Professor Alan Channing and Dr. Adriaan Engelbrecht for their supervision and advice concerning all molecular laboratory work and analysis. To Dr. Martin Hendricks for his supervision on all histological work and technical support throughout the years. The Biodiversity and Conservation Biology department for this amazing research opportunity and to the National Research Foundation for their continued financial support which enabled me to complete this degree. To all the friends within the BCB department who made going to the office and all fieldwork an unforgettable experience, particularly Aseeqah Davids and Robyn-lee Wales who over-extended themselves to help me gather data and specimens or simply keep my sanity. Special thanks are given to Thasrik Esack whose love and support (and tenacious editing of multiple drafts) enabled me to complete the last and final hurdle in this journey to submission. Lastly, but perhaps most especially, to Professor Mark J. Gibbons for his continuous supervision, advice, support and guidance, without whom this project would not have been a success. 2 Towards an unravelling of the taxonomy of Chrysaora (Scyphozoa; Semaeostomeae; Pelagiidae) from around South Africa Abstract Historically, two species of Chrysaora are known from the Benguela Current Ecosystem: C. fulgida (Reynaud 1830) and C. africana (Vanhöffen 1902). However a third morphotype is now seen, which bears a resemblance to both. Thus a complete qualitative and quantitative analysis of the morphometric and meristic data of these three species was conducted, along with an in depth study into the cnidome as a potential tool of identification. These findings are supplemented by a genetic analysis using cytochrome c oxidase subunit I and internal transcribed spacer 1 gene markers. Three species were unambiguously identified. The genetics and morphology showed considerable divergence, with some of the features used to separate them including: tentacle number and shape, colouration and shape of the oral arm, shape of the gastrovascular pouches and the number and shape of the marginal lappets. Although the mtDNA indicated clear separation of the three Chrysaora, the nucDNA displayed some ambiguity. The cnidome showed considerable divergence and succeeded in separating these species, while the rhopalia of the three species also showed distinct differences in the lengths of the rhopalal canals and basal stems. Much of the confusion surrounding jellyfish taxonomy has been the result of observations made on predominantly preserved specimens that are in subpar quality, a hinderence which this study endeavored to overcome. Keywords: Agulhas, Benguela, COI, ITS1, Nematocysts, Scyphomedusae, Systematics 3 1. Introduction Cnidarian history: Medusozoa vs. Anthozoa Cnidaria is a diverse phylum comprised of approximately 11 000 relatively “simple” species that are characterized by their ability to synthesize a highly complex cellular organelle known as the cnida (Daly et al. 2007). The cnida is a diagnostic characteristic for the Cnidaria (Daly et al. 2007), and is often regarded as an encapsulated stinging cell possessing an eversible tubule (Weill 1934). In addition to the cnida, radial symmetry and the presence of a polyp (benthic attached phase) and planula (short-lived larval phase) stage within their development have also been considered diagnostic for this phylum. However, the latter mentioned features can prove problematic as many Cnidaria
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