?cr. to' eO v. SYSTEMATIC STUDIES OF PELAGIC HYPERIIDEAi\ AMPHIPODS OF THE INFRAORDER PHYS OCEPHALATA (CRUSTACEA: AMPHIPODA: HYPE,RIIDEA) by Wotfgang Zeidler, B.Sc. (Hons), MSc. A thesis submitted for the degree of Doctor of Philosophy within the UniversitY of Adelaide Department of Environmental Biology August 2000 Volume I: Text, Appendices and Bibliography ll Contents Abstract vi Statement vln Acknowledgments ix 1. Introduction 1 2. Evolution/Phylogeny ofHyperiidean Amphipods 6 3. Materials and Methods 9 4. Note on Major pre-2Oth Century Collections 13 5. Systematics 16 Suborder HYPERIIDEA Milne-Edwards, 1 830 l6 Infraorder PHYSOSOMATA Pirlot, 1929 16 Infraorder PHYSOCEPHALATA Bowman & Gruner, 1973 18 5.1 Superfamily VIBILIOIDEA Bowman & Gruner,1973 23 Family VIBILIIDAE Dana, 1852 24 Vibilia H. Milne-Edwards, 1830 26 Vibilioides Cheweux, 1905 74 Family CYLLOPODIDAE Bovallius, 1 887 77 Cyllopus Dana, 1853 78 Family PARAPHRONIMIDAE Bovallius, 1 887 84 P ar ap hr o nim a Claus, | 87 9 85 5.2 Superfamily CYSTISOMATOIDEA new superfamily 92 Family CYSTISOMATIDAE Willemöes-Suhm, I 875 92 Cy s t i s o m a Guérin-Mén ev ille, | 8 42 93 5.3 Superfamily PHRONIMOIDEA Bowman & Gruner,1973 120 Family PHRONIMIDAE Dana, 1852 t22 Phronima Latreille, 1 802 r23 Phronimella Claus, 1 871 t32 Family PHROSINIDAE Dana, 1852 134 Phrosina Risso, 1822 135 Anchyl omer a Mllne-Edwards, 1 8 3 0 t37 Primno Guérin-Méneville, 1 836 t39 111 Family HYPEzuIDAE Dana, 1852 t45 Hyperia Latreille in Desmarest, 1823 t47 Themisto Guérin, 1825 t49 Hyperiella Bovallius, 1 887 151 Hyperoche Bovallius, 1 887 151 P e gohyp er i a B arnard, 1932 t52 LaxohyperiaYinogradov & Volkov in Vinogradov et a1.,1982 153 Family LESTzuGONIDAE new family 153 Lestrigonus Milne-Edwards, 1 830 r57 Phronimops¿s Claus, I 879 t57 Themistella Bovallius, 1 887 158 Hyperioides Chevreux, 1 900 158 Hyperietta Bowman, 1 973 159 Hyperionyx Bowman, I 973 1s9 Family IULOPIDIDAE new family 160 Iulopis Bovallius, 1 887 161 Family BOUGISIDAE new family r66 BougisiaLaval,1966 r67 Family DAIRELLIDAE Bovallius, I 887 t69 Dairella Bovallius, 1 887 t70 5.4 Superfamily LYCAEOPSOIDEA Bowman & Gruner,1973 176 Family LYCAEOPSIDAE Chevreux, 1 91 3 176 Lycaeopsis Claus, 1879 t78 5.5 Superfamily PLATYSCELOIDEA Bowman & Gruner,1973 187 Family PRONOIDAE Claus, 1879 190 Pronoe Guérin-Méneville, I 836 192 Family PARAPRONOIDAE new family 19s Purapronoe Claus, 1879 t97 Eupronoe Claus, 1879 202 Family AMPHITHYRIDAE new family 205 Amphithyrus Claus, 1879 208 Paralycaea Claus, 1879 2t2 Amphithyropsis new genus 215 IV Family BRACIIYSCELIDAE Stephensen, 1923 2r7 B r a chys c elus B ate, 186 | 219 Family LYCAEIDAE Claus, 1879 224 Lycaea Dana, 1852 226 Simorhynchotus Stebbing, I 888 230 Family ANAPRONOIDAE Bowman &. Gruner, 1973 232 Anapronoe Stephensen, 1925 234 Family THAMNEIDAE new family 236 Thamneus Bovallius, 1 887 238 Family TRYPHANIDAE Boeck, 1871 240 Tryphana Boeck, 1871 242 Family OXYCEPHALIDAE Bate, 1861 244 Oxycephaløs Milne-Edwards, 1 830 247 RhabdosomaWhite,1847 2s2 Leptocotis Streets, I 877 2s6 Calamorhynchus Streets, 1 878 259 Glos s ocephalus Bov allius, 1 8 87 262 Tullbergella Bovallius, 1 887 265 Streetsia Stebbing, 1 888 268 Cranocephalus Bovallius, 1 890 272 Family PLATYSCELIDAE Bate, 1862 275 Platyscelus Bate, 1861 279 Hemityphis Claus, 1879 283 Paratyphis Claus, 1879 286 Tetrathyrus Claus, 1 879 289 Family PARASCELIDAE Bovallius, 1 887 292 Parascelus Claus, 1879 295 Thyropus Dana,1852 298 Schizoscelzs Claus, 1879 301 Euscelus Claus, 1879 304 6. Discussion and Conclusion 308 Appendix I 311 Revised Classification of the Infraorder Physocephalata Appendix 2 314 Character List Appendix 3 322 Item Descriptions Appendix 4 326 Paupdata: Cladistic Data Matrix Bibliography 330 VI Abstract The systematics of the hyperiidean amphipod infraorder Physocephalata is investigated, with particular emphasis on the higher classification of the superfamily Platysceloidea, that has proved to be the most problematical superfamily for zooplankton workers. The study is based on collections held by major natural history museums in Australia, Europe and North America. All of these museums were visited personally, and in addition, specimens were borrowed from a number of other overseas institutions. Current studies of the systematics of the Physocephalata support the hypothesis that this group is probably polyphyletic, and genera have sometimes been grouped together in families with no phylogenetic justification. Consequently a number of families are split, resulting in an increase in the number of monogeneric families, giving a classification that reflects the apparently ancient phylogenetic origins of the group. A new key to all families of the sub-order Hyperiidea is provided, as are keys to the genera, and most species, of the infraorder Physocephalata. New diagnoses, reflecting the results of this study, are given for all superfamilies and families, and most genera and species. Twenty-three families,54 genera and 168 species of Physocephalata are recognised in this review. The superfamily Vibilioidea now consists of the families Vibiliidae, Cyllopodidae and Paraphronimidae. The family Vibiliidae is reviewed. Cyllopus is placed in the family Cyllopodidae, while Vibilioides is recognised as a valid genus' The family Cystisomatidae is reviewed, and placed in a new superfamily, Cystisomatoidea, because it has a number of characters that preclude it from the superfamily Vibilioidea. The superfamily Phronimoidea is reviewed at the family and genus level. The families Phronimidae, Phrosinidae and Dairellidae remain unchanged but the family Hyperiidae is split into four. The genera Hyperia, Themisto, Hyperiella, Hyperoche, Pegohyperia vlt and Laxohyperia are retained in the family Hyperiidae. The genera Lestrigonus, Phronimopsis, Themistella, Hyperioides, Hyperietta and Hyperionyx are transferred to the new family, Lestrigonidae. Iulopis and Bougisia do not fit into either of these families, nor do they resemble each other, and they are therefore placed in two new families, Iulopididae and Bougisidae. The superfamily Lycaeopsoidea is maintained on the basis of the morphology of the antennae, the mouthparts and the extreme sexual dimorphism. Genera of the superfamily Platysceloidea are diagnosed using the taxonomic database program DELTA (Dalwitz et al. 1999). This database is also used for a phylogenetic analysis of the genera using PAUP (Swofford 2000). Taxonomic changes as a result of this study are summarised as follows. The family Pronoidae is restricted to the monotypic genus Pronoe. Amphithyrzs has characters in common with Paralycaea and the new genus Amphithyropsis, and together they form the new family Amphithyridae. The family Brachyscelidae is recognised for the genus Brachyscelus. Thamneus has a number of characters that differ considerably from any other genus and is therefore placed in the new family Thamneidae. Euprono¿ is like Paraprono¿ in the morphology of the moutþarts, antennae and gnathopods, and together they form the new family Parapronoidae. The family Lycaeidae is limited to Lycaea and Simorhynchotus. The status of the family Anapronoidae is confirmed. The family Oxycephalidae seems to be polyphyletic but more work is required to resolve the systematic status of the eight genera recognised. Metalycaea globosa, previously included in the Oxycephalidae, is regarded a species of Lycaea. The family Platyscelidae is restricted to four genera, Platyscelus, Paratyphis, Tetrathyrus and Hemityphis. Amphithyrus is removed to the new family Amphithyridae. The family Parascelidae is also restricted to four genera, Parascelus, Thyropus, Schizoscelus and Euscelus. Hemiscelus is regarded a slmonym of Hemityphis. All records of associations with gelatinous zooplankton are documented. It is suggested that a more detailed study of this unusual relationship might help to resolve the evolutionary origins and phylogeny of the Hyperiidea. vrl_l- Statement This work contains no material that has been accepted for the award of any other degree or diploma in any university or other tertiary institution and, to the best of my knowledge and belief, contains no material previously published or written by another person, except where due reference has been made in the text. I give consent to this copy of my thesis, when deposited in the University Library, being made available for loan and photocopying, Wolfgang Zeidler August,2000 IX Acknowledgments I am indebted to my supervisor, Em. Prof. Bill Williams, and my co-supervisor for the latter half of my studies, Dr Derek Duckhouse, for providing invaluable guidance and encouragement. I am very grateful to the South Australian Museum and the Department of Zoology at the University of Adelaide (now the Department of Environmental Biology) for providing the facilities and administrative support to enable me to undertake my research. This study would not have been possible without the cooperation and assistance of numerous people in charge of collections, or facilities, at the major institutions of the world. All were exceptionally generous with their time, searching for specimens, arranging loans, or making me feel welcome and providing access to collections during personal visits, or helped in other ways. I am most grateful to all of them. There are so many that they are listed by continent and institution as follows. In Europe: at the BIVINH, Ms J. Ellis for the loan of specimens, Ms M. Lowe, Mr P. Clark and Prof. G.A. Boxshall for providing access to the collections and for the loan of specimens, the BMNH also gave permission to reproduce the drawings by Sydney 'Wheeler Parkinson for one of my papers (Zeídler 1995a) and Mr A. gave valuable advice, and Mr R. Banks and Mr P. Cooper organised copies of Parkinson's drawings for publication; at the MNHN, Dr D. Defaye for the loan of specimens and information on the collections; at the MOM, Dr C. Carpine for the loan of specimens; at the NMW, Dr V. Stagl for the loan of specimens; at the SMNH, Mr L. Sandberg for the loan of specimens, Ms K. Sindemark, Dr A. Waren and Prof. C. Erseus for their kind hospitality and for providing access to the collections; at the Z}i4.P, Dr H. E. Gruner for the loan of specimens, Dr C.O.