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Evolution, Systematics and Distribution of Desmosomatidae (Isopoda, Peracarida, Crustacea) in the Deep Sea

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Evolution, Systematics and Distribution of Desmosomatidae (Isopoda, Peracarida, Crustacea) in the Deep Sea Evolution, Systematics and Distribution of Desmosomatidae (Isopoda, Peracarida, Crustacea) in the deep sea Dissertation Zur Erlangung des Doktorgrades im Department Biologie der Fakultät für Mathematik, Informatik und Naturwissenschaften der Universität Hamburg vorgelegt von Saskia Bianca Brix aus Ellerau Hamburg 2006 According to ICZN article 8.3 all names and nomenclatural acts in this thesis are disclaimed for nomenclatural purposes. Table of contents Table of contents Summary i Abbrevations iii 1 Introduction 1 1.1 Deep-sea Isopoda 1 1.2 Desmosomatidae Sars, 1897 1 1.2.1 South Atlantic 3 1.2.2 Southern Ocean 6 1.2.3 Systematic problems 7 1.3 Phylogenetic cladistics 8 1.4 Aims and Questions 9 2 Material and methods 10 2.1 Sampling 10 2.2 Taxonomic methods 12 2.3 Collection Material 15 2.3.1 University of Hamburg: Zoological Museum 15 2.3.2 Smithonian Institution: Natural Museum of Natural History (Washington D.C., U.S.A. 16 2.3.3 American Museum of Natural History (New York, U.S.A.) 17 2.3.4 Museum Victoria (Melbourne, Australia) 17 2.3.5 Australian Museum (Sydney, Australia) 18 2.3.6 New Zealand Institution of Oceanographic and Atmospheric Research (NIWA) (Wellington, New Zealand) 18 2.3.7 Zoologisk Museum Kopenhavn (Kopenhagen, Danmark) 18 2.4 SEM: handling of species used for pictures 19 2.5 Phylogenetic Methods 19 2.6 List of species used for phylogenetic analysis 21 2.7 Characters used in phylogenetic analysis 24 3 Results 25 3.0 Zoogeographic aspects 25 3.0.1 Desmosomatidae of DIVA-1 25 3.0.2 Species composition at the seven EBS stations of DIVA-1 26 3.0.3 Desmosomatidae of ANDEEP I & II 29 3.0.4 Species composition at the ANDEEP stations 30 3.0.5 Distribution of the family Desmosomatidae 33 3.1 Taxonomy 38 3.1.1 Systematic overview 38 3.1.2 Groundpattern of Janiroidea (modified after Ax 1999; Gruner 1965; Hessler, Wilson & Thistle 1979; Wilson 1987 and Wägele 1989) 38 3.1.3 Characters of Desmosomatidae referring to the groundpattern of Janiroidea 39 3.1.3.1 Body 39 3.1.3.2 Anus 42 3.1.3.3 Sexual dimorphism 42 3.1.3.4 Cephalic rostrum 43 3.1.3.5 Eyes 43 3.1.3.6 Antennula 43 3.1.3.7 Antenna 44 3.1.3.8 Mandible 44 3.1.3.9 Maxillae 45 Table of contents 3.1.3.10 Maxilliped 46 3.1.3.11 Face of cephalon 47 3.1.3.12 Coxae 47 3.1.3.13 Pereonites 47 3.1.3.14 Pereopods 48 3.1.3.14.1 Pereopod I 49 3.1.3.14.2 Pereopods II and III 52 3.1.3.14.3 Pereopods IV to VII 52 3.1.3.15 Pleopods 53 3.1.3.16 Uropods 53 3.1.3.17 Pleotelson 54 3.1.4 Diagnoses 54 3.1.4.1 Desmosomatidae Sars, 1897 54 3.1.4.2 Diagnoses of the subfamilies 56 3.1.4.2.1 Austroniscinae subfam. nov. 56 3.1.4.2.2 Desmosomatinae Hessler, 1970 57 3.1.4.2.3 Eugerdellatinae Hessler, 1970 57 3.1.4.2.4 Nannoniscinae (Hansen, 1916) 57 3.1.4.2.5 Pseudomesinae (Hansen, 1916) 57 3.1.4.3 Diagnoses of the genera 58 3.1.4.3.1 Austroniscus Vanhöffen, 1914 58 3.1.4.3.2 Chelator Hessler, 1970 58 3.1.4.3.3 Cryodesma Svavarsson, 1988 58 3.1.4.3.4 Desmosoma Sars, 1864 59 3.1.4.3.5 Disparella Hessler, 1970 59 3.1.4.3.6 Echinopleura Sars, 1899 60 3.1.4.3.7 Eugerda Meinert, 1890 60 3.1.4.3.8 Eugerdella Kussakin, 1965 61 3.1.4.3.9 Exiliniscus Siebenhaller & Hessler, 1981 61 3.1.4.3.10 Hebefustis Siebenhaller & Hessler, 1977 61 3.1.4.3.11 Micromesus Birstein, 1963 62 3.1.4.3.12 Mirabilicoxa Hessler, 1970 62 3.1.4.3.13 Momedossa Hessler, 1970 63 3.1.4.3.14 Nannoniscoides Hansen, 1916 63 3.1.4.3.15 Nannonisconus Schultz, 1966 63 3.1.4.3.16 Nannoniscus Sars, 1870 64 3.1.4.3.17 genus novum 64 3.1.4.3.18 Oecidiobranchus Hessler, 1970 65 3.1.4.3.19 Panetela Siebenhaller & Hessler, 1981 65 3.1.4.3.20 Paradesmosoma Kussakin, 1965 65 3.1.4.3.21 Prochelator Hessler, 1970 66 3.1.4.3.22 Pseudogerda Kussakin, 1965 66 3.1.4.3.23 Pseudergella gen. nov. 66 3.1.4.3.24 Pseudomesus Hansen, 1916 67 3.1.4.3.25 Rapaniscus Siebenhaller & Hessler, 1981 67 3.1.4.3.26 Reductosoma Brandt, 1992 67 3.1.4.3.27 Regabellator Siebenhaller & Hessler, 1981 68 3.1.4.3.28 Saetoniscus Brandt 2002 68 3.1.4.3.29 Thaumastosoma Hessler, 1970 68 3.1.4.3.30 Torwolia Hessler, 1970 68 3.1.4.3.31 Whoia Hessler, 1970 68 Table of contents 3.1.5 New species 70 3.1.5.1 DIVA-1 70 3.1.5.1.1 Desmosoma renatae sp. nov. 70 3.1.5.1.2 Eugerdella theodori sp. nov. 78 3.1.5.1.3 Momedossa longipedis sp. nov. 87 3.1.5.1.4 Torwolia tinbienae sp. nov. 95 3.1.5.2 ANDEEP I & II 103 Eugerdella serrata sp. nov. 103 3.1.5.3 Collection material 114 3.1.5.3.1 gen. nov. sp. nov. A 114 3.1.5.3.2 Prochelator maorii sp. nov 116 3.1.5.3.3 Pseudomesus satanus sp. nov. 125 3.1.5.3.4 Paradesmomsoma australis sp. nov. 132 3.1.5.3.5 Oecidiobranchus slopei sp. nov. 140 3.1.5.3.6 Disparella kensleyi sp. nov. 146 3.1.5.3.7 Echinopleura cephalomagna sp. nov. 153 3.1.5.3.8 Whoia victoriensis sp. nov. 161 3.1.5.3.9 Pseudogerda anversense (Schultz, 1979) 196 3.1.5.4 Redescriptions 170 3.1.5.4.1 Chelator brevicauda (Menzies & George, 1972) 170 3.1.5.4.2 Disparella neomana (Menzies & George, 1972) 176 3.1.5.4.3 Disparella funalis (Menzies & George, 1972) 185 3.1.5.4.4 Eugerdella rotunda (Menzies & George, 1972) 193 3.1.5.4.5 Pseudogerda anversense (Schultz, 1979) 196 3.1.5.4.6 Desmosoma dolosa (Menzies & George, 1972) 203 3.1.5.4.7 Mirabilicoxa acuta (Menzies & George, 1972) 210 3.1.5.4.8 Mirabilicoxa similipes (Menzies & George, 1972) 217 3.2 Phylogeny 225 3.2.1 List of characters used in the phylogenetic analysis 225 3.2.2 Consensus trees 233 3.2.3 Character distribution in the trees 237 4 Discussion 248 4.0 Zoogeographic aspects 248 4.1 Are Desmosomatidae monophyletic? 250 4.1.1 Relationships of the families 250 4.1.2 Molecular hints 253 4.1.3 Two families or one? 254 4.2 One family – five subfamilies 257 4.2.1 Discussion of the subfamilies and genera 257 4.2.1.1 Austroniscinae subfam. nov. 257 4.2.1.1.1 Nannoniscoides Hansen, 1916 257 4.2.1.1.2 Nannoniscella (Hansen, 1916) 258 4.2.1.2 Desmosomatinae Hessler, 1970 259 4.2.1.2.1 Balbidocolon Hessler, 1970 259 4.2.1.2.2 Desmosoma Sars, 1864 261 4.2.1.2.3 Echinopleura Sars, 1897 263 4.2.1.2.4 Eugerda Meinert, 1890 263 4.2.1.2.5 Mirabilicoxa Hessler, 1970 270 4.2.1.2.6 Momedossa Hessler, 1970 271 4.2.1.2.7 Pseudogerda Kussakin, 1965 272 4.2.1.2.8 Torwolia Hessler, 1970 272 Table of contents 4.2.1.3 Eugerdellatinae Hessler, 1970 273 4.2.1.3.1 Chelator Hessler, 1970 273 4.2.1.3.2 Chelibranchus Mezhov, 1986 275 4.2.1.3.3 Cryodesma Svavarsson, 1988 276 4.2.1.3.4 Disparella Hessler, 1970 278 4.2.1.3.5 Eugerdella Kussakin, 1965 278 4.2.1.3.6 Leutziniscus George, 2001 283 4.2.1.3.7 Oecidiobranchus Hessler, 1970 284 4.2.1.3.8 Paradesmosoma Kussakin, 1965 285 4.2.1.3.9 Prochelator Hessler, 1970 286 4.2.1.3.10 Reductosoma Brandt, 1992 298 4.2.1.3.11 Thaumastosoma Hessler, 1970 298 4.2.1.3.12 Whoia Hessler, 1970 300 4.2.1.4 Nannoniscinae (Hansen, 1916) 302 4.2.1.5 Pseudomesinae (Hansen, 1916) 304 4.2.1.5.1 Pseudergella gen. nov. 304 4.2.1.5.2 Pseudomesus Hansen, 1916 306 4.2.2 Discussion of characters used in phylogeny 307 4.2.2.1 Setation 308 4.2.2.2 Characters 309 4.3 Results of phylogenetic analysis 343 4.3.1 Topology and tree data 343 4.3.2 Results compared to previous phylogenetic studies 344 4.3.3 Problematic characters and clades and their plausibility 345 5 Outlook 348 6 References 349 7 Acknowledgments 361 8 Appendix 363 Summary Summary The present thesis deals with one deep-sea family of Isopoda: Desmosomatidae Sars, 1897. It contains three major topics: 1) taxonomy, 2) biogeographic and distributional aspects (aspects of zoogeography) and 3) phylogeny. Based on morphological characters, a revision of the family Desmosomatidae Sars, 1897 is presented. In the taxonomy part four new species, Desmosoma renatae sp. nov., Eugerdella theodori sp. nov., Momedossa longipedis sp. nov. and Torwolia tinbienae sp. nov. are described from material of the DIVA-1 expedition and Eugerdella serrata sp.
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