The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgementTown of the source. The thesis is to be used for private study or non- commercial research purposes only. Cape Published by the University ofof Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University Taxonomy, Systematics and Biogeography of South African Cirripedia (Thoracica) Aiden Biccard Town A thesis submitted in fulfilment of the degreeCape of Master of Science in the Department of Zoology, Faculty of Science, University of Cape Town Supervisor Prof. Charles L. Griffiths University 1 Town “and whatever the man called every livingCape creature, that was its name.” - Genesis 2:19 of University 2 Plagiarism declaration This dissertation documents the results of original research carried out at the Marine Biology Research Centre, Zoology Department, University of Cape Town. This work has not been submitted for a degree at any other university and any assistance I received is fully acknowledged. The following paper is included in Appendix B for consideration by the examiner. As a supervisor of the project undertaken by T. O. Whitehead, I participated in all of the field work and laboratory work involved for the identification of specimens and played a role in the conceptualisation of the project. Figure 1 was compiled by me. Town Whitehead, T. O., Biccard, A. and Griffiths, C. L., 2011. South African pelagic goose barnacles (Cirripedia, Thoracica): substratum preferences and influences of plastic debris on abundance and distribution. Crustaceana, 84(5-6): 635-649. Cape of SIGNATURE: _________________ DATE: _________________ University 3 Contents Abstract ____________________________________________________________________ 5 Acknowledgements ___________________________________________________________ 6 General Introduction _________________________________________________________ 7 Chapter 1 Additions to the South African Cirripedia (Thoracica) ____________________________ 10 Chapter 2 A Guide to the South African Cirripedia (Thoracica) _____________________________ 35 Plates ____________________________________________________________________Town 110 Chapter 3 Biogeographic patterns in species richness and endemicityCape within South African Cirripedia (Thoracica)________________________________________________________________ 152 of Synthesis__________________________________________________________________ 171 Appendix A _______________________________________________________________ 173 Appendix B _______________________________________________________________ 177 University 4 Abstract The South African Cirripedia (Thoracica) are reviewed for the first time in 88 years, since that of Barnard (1924). Data collection consisted of records from the South African Museum, previously published literature, and specimens collected in the field. The current state of knowledge of the taxon in South Africa has been considerably raised. Thirteen new records are added to the fauna, of which 84.6% are cosmopolitan and 15.4% introduced species. This raises the total number of South African Cirripedia (Thoracica) to 86, of which 64.06% are cosmopolitan, 11.24% introduced and 24.7% endemic. Excluded from this, are three unknown species, which are likely to be new to science. Descriptions of these species will be undertaken outside of this thesis and published; however, they were classified as endemic and included in the analyses presented in Chapter three. Eleven of the new fauna can be described as “offshoreTown benthic” and represent well known deep-water taxa. The remaining five are “coastal inshore” species. The Agulhas bioregion displays the highest species richness, with a total of 53 species recorded. It also contains the highest number of endemic species, 15 in total, outCape of all nine bioregions in South Africa. Lowest species richness was observed in the deep water, offshore bioregions (apart from the South-west Indian offshore) as a result of a lackof in samples from these areas. High endemicity rates in the Agulhas and the South-west Indian offshore bioregion are most likely attributable to inadequate sampling of the surrounding deep water habitat. High discovery rates of new species to the region are indicative of many years without the work of fulltime taxonomic experts. Additional expertise and work is required in order to improve on the current state of knowledge of marine biodiversityUniversity in South Africa. 5 Acknowledgements A great deal of value was added to this project by the following people who contributed material or data is some form or another: Clova Jurk, Dr. Ken Hutchings, Debbie Robertson-Andersson, Dr. Kerry Sink, Dr. Peter Ryan, Dr. Lara Atkinson, Megan Clair Laird, Matthew Melidonis, Otto Whitehead, Katharine James, Oscar Kathide, Rebecca Milne, Jennifer Olbers, Laura Tang, Dane- Lee Marx and Lousie Lange – thank you to all of you. I would like to acknowledge Elizabeth Hoenson of the Marine Invertebrate Collection at the Iziko South African Museum for her assistance in extracting data from historical records and for allowing me to access type specimens – this project would not have been possible without you. Thank you to all the cirripedologists around the world for Townanswering emails and offering to receive specimens for identification – Prof. Yair Achituv, Dr. Francis Kerckhof, Prof. John Buckeridge, Prof. William Newman and Dr. Harold Voris. You were all a tremendous help. Special mention must be made to my lab-matesCape Rebecca Milne and Megan Laird for their support and for constantly fielding questions.of Megan, you were particularly helpful during the closing stages of this project and I really appreciate everything you have done. I also acknowledge Karen Tunley for her assistance in the production of figures 14, 15 and 18. Last and certainly not least, thank you to my supervisor, Prof. Charles L. Griffiths, for offering this project to me and believing that I could succeed. Thank you for all the ideas borne out of the many thought-provoking conversations over a cup of tea. This project wasUniversity funded by a National Research Foundation grant to Prof. Charles Griffiths, and the Marine Biology Research Centre in the Zoology Department at the University of Cape Town. 6 General Introduction The Cirripedia (Thoracica) represents a highly diverse taxon within the marine Crustacea with species living on a wide range of substrata from the intertidal zone down to the deepest parts of the ocean. We are most familiar with those frequently encountered in the intertidal and coastal inshore zones, however, these represent only a fraction of the existing diversity, the majority of which is located in pelagic and deeper water offshore. South African Cirripedia were first mentioned in the literature by Christian Ferdinand Friedrich Krauss in 1848. The two species he described can be found in his volume on molluscs, as they were considered to be members of this phylum at the time. They were Asemusporosus and Coniarosea – now known as Tetraclita serrata (Darwin, 1854) and Tetraclita rufotincta (Pilsbry, 1916) respectively. It was J. V. Thompson (1823) who established that the Cirripedia were in factTown not molluscs but crustaceans. Darwin (1852, 1854) later produced an extensive work on the subject, which included several type specimens from South Africa and Cape Town. He recorded eleven species in total, of which five types were classified as endemic, these includedCape Balanus capensis (Austromegabalanus cylindricus), Tetraclita serrata, Cthamalus ofdentatus , Octomeris angulosa and Scalpellum ornatum (Ornatoscalpellum ornatum). The first major contribution to the crustacean fauna of South Africa was that of Stebbing (1910) who documented 26 species of barnacles. This was later increased to 67 species of Cirripedia (Thoracica) by K. H. Barnard who wrote a detailed guide to the fauna in 1924. Since then only several other publications have reported additional species and no updated version of Barnard’s 88 year oldUniversity guide to the Cirripedia exists. The sole purpose of this thesis is to provide such a guide to the scientific community in South Africa. This will be achieved in the following three chapters: Additions to the South African Cirripedia (Thoracica) which accounts for all the species recorded for the first time within our political borders. A diagnosis and abbreviated description is provided for each of the new species to aid in their identification. Furthermore, they are classified as being either cosmopolitan, endemic or introduced based on known global 7 distribution ranges and invasive tendencies as defined in the literature (Mead et al., 2010; Carlton et al., 2011). A guide to the South African Cirripedia (Thoracica) provides an illustrated account of every species presently recorded in South Africa, and is structured to be used as a much needed tool for the accurate identification of species by the scientific community. It includes updated synonymies and classification of species based on their current inferred phylogenetic relationships using alpha-taxonomy. Distribution ranges for each species are also established and are provided as a series of maps together with a photographic account of the fauna all in the interest of assisting the user in narrowing down an identification of a species. All data, apart from the pelagic taxa, are displayed as points rather than ranges as these are more informative