A CRITICAL ASSESSMENT OF THE DENDROCHIROTID SUBFAMILIES, SCLERODACTYLINAE AND THYONINAE, WITH THE TAXONOMIC MANAGEMENT OF THE “SUPERGENUS” THYONE (ECHINODERMATA: HOLOTHUROIDEA) by PREYAN ARUMUGAM Submitted in fulfilment of the academic requirements for the degreeof Master of Science in the School of Life Sciences, College of Agriculture, Engineering and Science, University of KwaZulu-Natal 2011 Supervised by Prof. Ahmed S. Thandar ABSTRACT The key character separating the dendrochirotid families Sclerodactylidae (sensu Pawson & Fell, 1965) and the Phyllophoridae (sensu Pawson & Fell, 1965), i.e. entire or undivided radial processes to the calcareous ring in the former and sub-divided processes in the latter, is unjustified since most sclerodactylid species also have sub- divided processes. It is here assumed that the basis of elevating the subfamily Sclerodactylinae Panning to family level was established on a misinterpretation or mistranslation of the original diagnosis of this subfamily or a lapsus calumni meaning “plates” instead of “processes”. Panning (1949) categorically states that the processes in the Sclerodactylinae are composed of 3–4 large pieces of calcite and only as an exception they are unbroken. Since Pawson & Fell gave no other distinction between the Sclerodactylidae and the Phyllophoridae, the former is here considered an invalid taxon and its three current subfamilies (Sclerodactylinae, Sclerothyoninae Thandar and Cladolabinae Heding & Panning) are re-assigned to the Phyllophoridae. This family now includes six subfamilies: Cladolabinae, Phyllophorinae Östergren, Sclerodactylinae, Sclerothyoninae, Semperiellinae Heding & Panning and Thyoninae Panning. The diagnosis of the Sclerodactylinae, restricted by Thandar (1989), is now modified to include also those forms whose radial and interradial plates may be slightly sub-divided but still form a short tube. Of the eleven genera placed within this subfamily subsequent to its erection, only ten of these remain. Neothyone Deichmann is a preoccupied name for which Lisacucumis is here proposed as a replacement. Thandar’s (1989) diagnosis of the Thyoninae is here accepted, however, the genus Thorsonia Heding is transferred to the Sclerodactylinae. Of the 66 nominal species which currently stand in the “supergenus” Thyone Jaeger, 10 are transferred to Havelockia Pearson within the Sclerodactylinae, while one species is regarded as a synonym of H. herdmani Pearson. In addition, six species are transferred to Stolus Selenka within the Thyoninae. Finally, three species are transferred to Sclerothyoninae, two within Sclerothyone Thandar and one within Temparena Thandar. Two species show an uncertain affinity to Thyone and are temporally removed from the genus. Furthermore, two species currently classified within Havelockia are transferred to Thyone. The now remaining 46 species are separated into seven groups based on the ii composition of their introvert deposits: tables only (8 spp.), rosettes only (5 spp.), tables and rosettes (21 spp.), tables and plates/?reduced tables (2 spp.), rosettes and plates/?reduced tables (3 spp.), plates only (2 spp.), or introvert deposits absent or unknown (5 spp.). Regrettably, no other character could be used in conjunction with the above to suggest at least sub-generic levels. Within the genus Havelockia, Cucumaria redimita Sluiter indicates an affinity with Pentamera Ayres. It is here transferred to this genus within the Thyoninae. Havelockia, now containing 17 species, is also revised. Keys, diagnoses and figures are provided for all nominal species now included in Thyone and Havelockia. Key words: Lisacucumis; Havelockia; new genus; Phyllophoridae; sea cucumbers iii PREFACE The experimental work described in this dissertation was carried out in the School of Biological & Conservation Sciences, University of KwaZulu-Natal, Durban, from January 2010 to December 2011, under the supervision of Professor Ahmed. S. Thandar. These studies represent original work by the author and have not otherwise been submitted in any form for any degree or diploma to any tertiary institution. Where use has been made of the work of others it is duly acknowledged in the text. The format and list of references of this dissertation are in accordance with the instructions to authors of the on-line journal Zootaxa, in which, it is anticipated, this investigation will be published. iv DECLARATION - PLAGIARISM I, Preyan Arumugam, declare that 1. The research reported in this dissertation, except where otherwise indicated, is my original research. 2. This dissertation has not been submitted for any degree or examination at any other university. 3. This dissertation does not contain other persons’ data, pictures, graphs or other information, unless specifically acknowledged as being sourced from other persons. 4. This dissertation does not contain other persons' writing, unless specifically acknowledged as being sourced from other researchers. Where other written sources have been quoted, then: a. Their words have been re-written but the general information attributed to them has been referenced. b. Where their exact words have been used, then their writing has been placed in italics and inside quotation marks, and referenced. 5. This dissertation does not contain text, graphics or tables copied and pasted from the Internet, unless specifically acknowledged, and the source being detailed in the thesis and in the References sections. Signed: ………………………………………………………… v To my fiancé, parents, and sister, Thank you for being my pillar of strength and my inspiration. I love you. “I can do all things through Christ which strengtheneth me” Philippians 4:13 vi LIST OF CONTENTS Abstract........................................................................................................................... ii Preface............................................................................................................................ iv List of Contents............................................................................................................. vii Acknowledgments......................................................................................................... xii List of Acronyms......................................................................................................... xiii List of Figures.............................................................................................................. xiv Introduction.................................................................................................................... 1 Materials and Methods.................................................................................................. 6 Systematic Account......................................................................................................... 7 PART I Order Dendrochirotida Grube, 1840................................................................................. 7 Key to the subfamilies of the Family Phyllophoridae Östergren, 1907.................... 8 Subfamily Sclerodactylinae Panning, 1949..................................................... 10 Key to the genera of the Subfamily Sclerodactylinae Panning, 1949..... 10 Genus Apentamera Deichmann, 1941............................................. 12 Genus Athyone Deichmann, 1941................................................... 13 Genus Deichmannia Cherbonnier, 1952......................................... 15 Genus Engeliella Cherbonnier, 1968............................................... 15 Genus Eupentacta Deichmann, 1938.............................................. 16 Genus Havelockia Pearson, 1903.................................................... 17 Genus Lisacucumis nom. nov. ........................................................ 21 Genus Pachythyone Deichmann, 1941............................................ 22 Genus Pseudothyone Panning, 1949............................................... 23 vii Genus Sclerodactyla Ayres, 1857................................................... 23 Genus Thorsonia Heding, 1940....................................................... 24 Subfamily Thyoninae Panning, 1949.............................................................. 25 Key to the genera of the Subfamily Thyoninae Panning, 1949............... 26 Genus Allothyone Panning, 1949..................................................... 26 Genus Pentamera Ayres, 1852........................................................ 27 Genus Stolus Selenka, 1867............................................................. 29 Genus Thyone Jaeger, 1833............................................................. 31 Genus Thyonina Thandar, 1990....................................................... 33 Subfamily Sclerothyoninae Thandar, 1989..................................................... 36 Key to the genera of the Subfamily Sclerothyoninae Thandar, 1989...... 36 Genus Neopentamera Deichmann, 1941......................................... 36 Genus Sclerothyone Thandar, 1989................................................. 37 Genus Temparena Thandar, 1989................................................... 40 PART II Subfamily Thyoninae Panning Genus Thyone Jaeger, 1833..................................................................... 43 Key to the genus Thyone Jaeger, 1833............................................ 45 T. anomala Östergren, 1898.................................................. 143 T. aurea (Quoy & Gaimard, 1834) ......................................