The ostracod genus Trachyleberis (Crustacea; Ostracoda) and its type species Simone N. Brandão, Moriaki Yasuhara, Toshiaki Irizuki & David J. Horne Marine Biodiversity ISSN 1867-1616 Volume 43 Number 4 Mar Biodiv (2013) 43:363-405 DOI 10.1007/s12526-013-0163-6 1 23 Your article is protected by copyright and all rights are held exclusively by Senckenberg Gesellschaft für Naturforschung and SpringerVerlag Berlin Heidelberg. This e-offprint is for personal use only and shall not be selfarchived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Mar Biodiv (2013) 43:363–405 DOI 10.1007/s12526-013-0163-6 ORIGINAL PAPER The ostracod genus Trachyleberis (Crustacea; Ostracoda) and its type species Simone N. Brandão & Moriaki Yasuhara & Toshiaki Irizuki & David J. Horne Received: 21 November 2012 / Revised: 12 April 2013 / Accepted: 14 May 2013 / Published online: 21 August 2013 # Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2013 Abstract The Trachyleberididae is one of the most diverse families of the Ostracoda in Mesozoic to present-day marine environments. Its type genus is Trachyleberis Brady, 1898, first described on the basis of specimens collected alive. Unfortunately the identity and morphological characteristics of the type species of Trachyleberis have long been the subject of confusion resulting from misidentifications and misunderstandings in the literature. We review the taxonomic history of this problem, providing extensive synonymies, new diagnoses, descriptions and illustrations of relevant species. We establish the identity of the type species of Trachyleberis, Cythere scabrocuneata Brady, 1880, by reference to a previously-designated lectotype from the Seto Inland Sea, Japan, noting that a second distinct species, Trachyleberis niitsumai Ishizaki, 1971, is represented in the original syntypic series. We consider that in the interests of nomenclatural and taxonomic stability C. scabrocuneata should be retained as the type species of Trachyleberis, despite the fact that the original description of the genus was based on a species misidentified as C. scabrocuneata and which was subsequently described as a new species, Trachyleberis lytteltonensis Harding and Sylvester-Bradley, 1953. The geographical and stratigraphical distributions of Trachyleberis are reconsidered in the light of our revision. Trachyleberis now comprises 18 known species inhabiting shallow marine environments of the Northwestern Pacific from Japan to the northern coast of the South China Sea off Hong Kong. The revised stratigraphic occurrence of Trachyleberis is from the Eocene to Recent. As many as 400 living and fossil species have been previously assigned to Trachyleberis and S. N. Brandão (*) Biozentrum Grindel und Zoologisches Museum, Universität Hamburg, Martin-Luther-King-Platz 3, Hamburg 20146, Germany e-mail: brandao.sn.100@gmail.com T. Irizuki Department of Geoscience, Interdisciplinary Graduate School of Science and Engineering, Shimane University, Matsue 690-8504, Japan S. N. Brandão Senckenberg am Meer, Deutsches Zentrum für Marine Biodiversitätsforschung (DZMB), Südstrand 44, Wilhelmshaven 26382, Germany M. Yasuhara School of Biological Sciences, University of Hong Kong, Hong Kong SAR, China M. Yasuhara Swire Institute of Marine Science, University of Hong Kong, Hong Kong SAR, China M. Yasuhara Department of Earth Sciences, University of Hong Kong, Hong Kong SAR, China M. Yasuhara Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA D. J. Horne School of Geography, Queen Mary University of London, Mile End Road, London E1 4NS, UK D. J. Horne Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK Present Address: S. N. Brandão Laboratorio de Geologia e Geofisica Marinha e Monitoramento Ambiental-GGEMMA, Departamento de Geologia and Programa de pós-Graduação em Geodinâmica e Geofísica, UFRN, Campus Universitário, Lagoa Nova, Caixa-Postal: 1596, 59072-970, Natal, RN, Brazil Author's personal copy 364 they range from the Cretaceous to the Recent; Cretaceous species formerly attributed to Trachyleberis are now reassigned to other genera. We suggest that Trachyleberis most likely evolved from a Cythereis-like ancestor during the Eocene–Oligocene of Japan, the same area that is the centre of its modern distribution. Keywords Biogeography . Type species designation . Marine . Ostracoda . Trachyleberididae . Taxonomic revision Introduction Members of the Trachyleberididae Sylvester-Bradley, 1948, one of the most diverse and abundant ostracod families, have inhabited marine environments since the Jurassic and have a global distribution today. The impressively long and detailed fossil record of trachyleberidid ostracods has an outstanding utility for macroevolutionary research and the potential to facilitate better understanding of the evolutionary history of marine life and testing of evolutionary theories over longer timescales than those available for most biological studies (Liow 2006, 2007), as well as to aid paleoceanographic reconstruction (e.g., Cronin 1991; Brouwers 1994; Irizuki et al. 2007). However, the morphologic and systematic identities of the Trachyleberididae and the closely related family Hemicytheridae remain controversial, in part because of considerable confusion regarding the trachyleberidid type genus (Trachyleberis Brady, 1898) and its type species. Difficulties arise for several reasons: 1. Brady (1898) misidentified the species he used for his description of his new genus Trachyleberis. Harding and Sylvester-Bradley (1953) showed that it was not, as Brady (1898) thought, Cythere scabrocuneata Brady, 1880, and accordingly they described it as a new species, Trachyleberis lytteltonensis; they also designated one of Brady’s (1880) syntypes as a lectotype of Cythere scabrocuneata, and mentioned Cythere scabrocuneata Brady, 1880 as the type species of Trachyleberis (although in 1953 the only way of posteriorly designating a type species was through the Commission of the ICZN). 2. The taxonomic identity of Trachyleberis scabrocuneata (Brady, 1880), the subsequently designated type species of Trachyleberis, is uncertain because the syntype series is now known to include at least two distinct species: Trachyleberis scabrocuneata (Brady, 1880) and Trachyleberis niitsumai Ishizaki, 1971. Unfortunately, Harding and Sylvester-Bradley (1953) did not figure their lectotype (which defines the true Trachyleberis scabrocuneata), but illustrated instead, as T. scabrocuneata, valves from the syntypic series which would now be Mar Biodiv (2013) 43:363–405 assigned to T. niitsumai. Importantly, the designated lectotype still exists even if it was not illustrated. 3. Species that are not conspecific with true T. scabrocuneata have nevertheless been illustrated under that name by various other authors, including an unnecessary neotype designated by Warne and Whatley (1996) in the mistaken belief that the lectotype was lost. 4. As a result of the above issues, the morphological characters that define the type species of Trachyleberis remain uncertain. The main aim of this paper is to revise Trachyleberis scabrocuneata and thereby establish unambiguously the identity of the genus Trachyleberis and to designate its type species. In order to do so, we provide extensive synonymies as well as new diagnoses, descriptions, Scanning Electron Microscope (SEM) and light microscope images and drawings of T. scabrocuneata (including the lectotype), T. niitsumai and T. lytteltonensis Harding and Sylvester-Bradley, 1953 (the lattermost being removed from the genus Trachyleberis sensu stricto), and reconsider the geographical and stratigraphical distribution of Trachyleberis in the light of our revision. Material and methods For taxonomic decisions we followed strictly the Fourth Edition of the International Code of Zoological Nomenclature 1999 (ICZN) (online version http://iczn.org/code). We use only terms recognized by the ICZN (i.e., lectotype, paralectotype(s), syntype(s), topotype(s), type species, type genus), while avoiding unofficial ones (e.g., hololectotype, lectoholotype, genotype). In order to clarify our review of the taxonomic confusion surrounding Trachyleberis scabrocuneata (Brady, 1880), herein we refer to the true identity of this species, as defined herein, as Trachyleberis scabrocuneata sensu stricto (or s.s). All specimens studied are included in the following collections: (1) Zoology, The Natural History Museum, London (NHM); (2) Zoologisk Museum, København (Zoological Museum, Copenhagen) (ZMUC CRU); (3) Micropalaeontology I, Senckenberg Research Institute and Natural History Museum Frankfurt (SMF Xe); (4) Crustacean collection of the Zoologisches Museum Hamburg, University of Hamburg (ZMH K-); (5) Resources Centre of the Great North Museum at the Discovery Museum, Newcastle upon Tyne (NEWHMF); (6) Department of Paleobiology, National Museum of Natural History, Smithsonian Institution (USNM)) (Table 1). Uncoated valves from the collection of the NHM were digitally imaged by SEM using a Zeiss environmental SEM LEO 1455VP in The Natural History Museum, London. Uncoated valves from Osaka Bay (Seto Inland Sea) were digitally imaged using a Philips XL-30 environmental SEM with LaB6 electron Slide no. Material “Trachyleberis” lytteltonensis NHM 1952.12.9.11 1 A M LV, 1 A M RV “Trachyleberis” lytteltonensis NHM 1952.12.9.21 “Trachyleberis” lytteltonensis NHM 1952.12.9.31 “Trachyleberis” lytteltonensis NHM 1952.12.9.41 “Trachyleberis” lytteltonensis NHM 1952.12.9.51 “Trachyleberis” lytteltonensis NHM 1952.12.9.61 “Trachyleberis” lytteltonensis NHM 1952.12.9.71 “Trachyleberis” lytteltonensis NHM 1952.12.9.81 “Trachyleberis” lytteltonensis ZMUC CRU 3630 “Trachyleberis” lytteltonensis ZMUC CRU 3630 “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis ZMUC CRU 3630 ZMUC CRU 3631 “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis ZMUC CRU 3631 ZMUC CRU 3631 ZMUC CRU 3631 “Trachyleberis” lytteltonensis NEWHM 1.54.24 “Trachyleberis” lytteltonensis Trachyleberis scabrocuneata NEWHM 1.54.24 NHM 1948.3.10.3, 767 Trachyleberis scabrocuneata NHM 1948.3.10.4, 768 Trachyleberis scabrocuneata NHM 1952.12.10.1 Trachyleberis scabrocuneata NHM 1952.12.10.2 Trachyleberis scabrocuneata NHM 1952.12.10.8.A Trachyleberis scabrocuneata NHM 1952.12.10.8.B Previously labelled as Trachyleberis lytteltonensis H. & S.-B. 1 A M LV, 1 A M RV Trachyleberis lytteltonensis H. & S.-B. 2 A M LV, 1 A M RV Trachyleberis lytteltonensis H. & S.-B. 1 (?A) F LV, 1 A F RV Trachyleberis lytteltonensis H. & S.-B. 1 J RV Trachyleberis lytteltonensis H. & S.-B. 1 J RV Trachyleberis lytteltonensis H. & S.-B. 1 J LV Trachyleberis lytteltonensis H. & S.-B. 1 A M LV, 1 A M RV Trachyleberis lytteltonensis H. & S.-B. bk. V(s) T. scabrocuneata (Brady, 1898) T. scabrocuneata 18 A F, 43 A M4 (Brady, 1898) 1 M RV, 1 bk. RLV T. scabrocuneata (Brady, 1898) 8 A F LV, 3 A F RV, T. scabrocuneata (Brady, 1898) 4 A M LV, 6 A M RV, 1 J LV, 1 J RV 1 A F LV T. scabrocuneata (Brady, 1898) 1 A M LV T. scabrocuneata (Brady, 1898) 77 A F, 64 A M, 3 J4 Trachyleberis lytteltonensis H. & S.-B. 1 A F LV, 1 A F RV, Trachyleberis lytteltonensis 3 A F RLV, 3 A M RLV, 1A M RV +50 “live” specimens Trachyleberis lytteltonensis 1 A M RV Trachyleberis scabrocuneata (Brady) 1 A F RV Trachyleberis scabrocuneata (Brady) 1 A M LV, 1 A M RV Trachyleberis scabrocuneata (Brady) fragmented SP Trachyleberis scabrocuneata (Brady) fragmented SP Trachyleberis scabrocuneata (Brady) empty (i.e. no valves) Trachyleberis scabrocuneata (Brady) Type Identification by Brady Identification by other authors holotype Cythere scabrocuneata T. scabrocuneata, T. lytteltonensis paratype Cythere scabrocuneata T. scabrocuneata, T. lytteltonensis paratype Cythere scabrocuneata T. scabrocuneata, T. lytteltonensis paratype Cythere scabrocuneata T. scabrocuneata, T. lytteltonensis paratype Cythere scabrocuneata T. scabrocuneata, T. lytteltonensis paratype Cythere scabrocuneata T. scabrocuneata, T. lytteltonensis Cythere scabrocuneata T. scabrocuneata, T. lytteltonensis paratype paratype no3 Cythere scabrocuneata T. scabrocuneata, T. lytteltonensis Cythere scabrocuneata T. scabrocuneata (Brady, 1898) no3 Cythere scabrocuneata 5 no3 no3 Cythere scabrocuneata 5 Cythere scabrocuneata T. scabrocuneata (Brady, 1898) no3 no3 no3 Cythere scabrocuneata T. scabrocuneata (Brady, 1898) Cythere scabrocuneata T. scabrocuneata (Brady, 1898) Cythere scabrocuneata 5 paratype Cythere scabrocuneata Trachyleberis lytteltonensis Author's personal copy Our identification Mar Biodiv (2013) 43:363–405 Table 1 Material studied herein paratype Cythere scabrocuneata Trachyleberis lytteltonensis paralectotype Cythere scabrocuneata S.-B.: T. scabrocuneata paralectotype Cythere scabrocuneata S.-B.: T. scabrocuneata lectotype Cythere scabrocuneata H. & S.-B.: T. scabrocuneata lectotype Cythere scabrocuneata H. & S.-B.: T. scabrocuneata paralectotype Cythere scabrocuneata H. & S.-B.: T. scabrocuneata paralectotype Cythere scabrocuneata H. & S.-B.: T. scabrocuneata 365 366 Table 1 (continued) Our identification Slide no. Material Previously labelled as Type Trachyleberis scabrocuneata NHM 1952.12.10.10.A 3 LV paralectotype Cythere scabrocuneata H. & S.-B.: T. scabrocuneata Trachyleberis scabrocuneata NHM 1952.12.10.11.A 2 RV, 1 bk. V Trachyleberis scabrocuneata NHM 1952.12.10.12.C 2 LV, 1 RV Trachyleberis Trachyleberis Trachyleberis Trachyleberis Trachyleberis Trachyleberis Trachyleberis Trachyleberis Trachyleberis Trachyleberis ZMH K-42870 ZMH K-42870 ZMH K-42870 ZMH K-42870 ZMH K-42870 ZMH K-42870 ZMH K-42870 ZMH K-42870 ZMH K-42870 NHM 1948.3.10.1, 765 1 1 1 1 1 1 1 1 1 1 Trachyleberis niitsumai NHM 1948.3.10.2, 766 1 A M RV Trachyleberis niitsumai NHM 1952.12.10.5.A 1 LV, 1 RV Trachyleberis niitsumai NHM 1952.12.10.5.B SP 3 LV, 1 RV Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata n. sp. Trachyleberis scabrocuneata (Brady) – – – – – – – – – Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) new slide by S.N.B. 1 LV new slide by S.N.B. Trachyleberis niitsumai A F LV A F RV A M LV A M LV A F LV A F RV (A-1) LV (A-1) RV A M RV A M LV paralectotype Cythere scabrocuneata H. & S.-B.: T. scabrocuneata paralectotype Cythere scabrocuneata H. & S.-B.: T. scabrocuneata No No No No No No No No No 2 – – – – – – – – – Cythere scabrocuneata 2 Cythere scabrocuneata S.-B.:T. scabrocuneata 2 Cythere scabrocuneata H. & S.-B.:T. scabrocuneata 2 Cythere scabrocuneata H. & S.-B.:T. scabrocuneata 2 Cythere scabrocuneata H. & S.-B.:T. scabrocuneata 2 Cythere scabrocuneata H. & S.-B.:T. scabrocuneata Cythere scabrocuneata H. & S.-B.:T. scabrocuneata Cythere scabrocuneata H. & S.-B.:T. scabrocuneata 1 LV 4 LV Trachyleberis scabrocuneata 2 new slide by S.N.B. 2 1 1 1 1 1 Trachyleberis sp. cf. T. niitsumai NHM 1952.12.10.3 1 RV Trachyleberis sp. cf. T. niitsumai NHM 1952.12.10.12.B 4155 Trachyleberis sp. cf. T. niitsumai NHM 1974.342 1 bk. LV 1 bk. LV, 1 bk. V – – – – Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata Cythere scabrocuneata Trachyleberis sp. cf. T. niitsumai NEWHM 1.43.22 2 A F RV, 2 A F LV, 1 A M LV, 3 ?A or (A-?1) M? RV, 8 J LV, 5 J RV A M LV A F RV (A-1) LV (A-1) RV A F RV No No No No 2 – – – – Cythere scabrocuneata 2 Cythere scabrocuneata H. & S.-B.:T. scabrocuneata 2 N Cythere scabrocuneata H. & S.-B.:T. scabrocuneata not studied by Brady P. & H.; W. & W.:T. scabrocuneata Cythere scabrocuneata – Cythere scabrocuneata Brady 2 – – – – S.-B.:T. scabrocuneata Mar Biodiv (2013) 43:363–405 NHM 1952.12.10.10.B SNB-1 060 Trachyleberis niitsumai NHM 1952.12.10.11.B SNB-1 061 Trachyleberis niitsumai NHM 1952.12.10.12.A 4131 Trachyleberis niitsumai NHM 1952.12.10.12.D SNB-1 062 Trachyleberis niitsumai USNM 560189 (MY: TRA1044) Trachyleberis niitsumai USNM 560190 (MY: TRA1045) Trachyleberis niitsumai USNM 560191 (MY: TRA1046) Trachyleberis niitsumai USNM 560192 (MY: TRA1047) Trachyleberis sp. cf. T. niitsumai NHM 1948.3.10.5, 769 – – – – – – – – – S.-B.:T. scabrocuneata Author's personal copy scabrocuneata scabrocuneata scabrocuneata scabrocuneata scabrocuneata scabrocuneata scabrocuneata scabrocuneata scabrocuneata niitsumai Identification by Brady Identification by other authors Our identification Slide no. Trachyleberis sp. cf. T. niitsumai NEWHM 1.43.23 Previously labelled as Type Identification by Brady Identification by other authors Cythere scabrocuneata Brady 2 Cythere scabrocuneata – 6 Cythere scabrocuneata H. & S.-B.:T. scabrocuneata 6 Cythere scabrocuneata H. & S.-B.:T. scabrocuneata 6 Cythere scabrocuneata H. & S.-B.:T. scabrocuneata 6 Cythere scabrocuneata H. & S.-B.:T. scabrocuneata no no holotype Cythere scabrocuneata – Cythere scabrocuneata – not studied by Brady Trachyleberis raynerae Trachyleberis sp.? NHM 1952.12.10.4 2 A F RLV; 1 A M RLV; 5 A M LV; 4 A M RV; 5 A F RV; 5 LV fragmented SP Trachyleberis sp.? NHM 1952.12.10.6 fragmented SP Trachyleberis sp.? NHM 1952.12.10.7 fragmented SP Trachyleberis sp.? NHM 1952.12.10.9 fragmented SP Pennyella sp. Pennyella sp. Trachyleberis raynerae 4 RV, 2 LV, 2 V 12 V 1 A M RV 1 (A-1) F RV Trachyleberis raynerae paratype not studied by Brady Trachyleberis raynerae 1 RV Trachyleberis raynerae paratype not studied by Brady Trachyleberis raynerae 1 A M LV Trachyleberis raynerae paratype not studied by Brady Trachyleberis raynerae Cytheroidea NHM 1900.3.6.237 NHM 1911.11.8.M3310 PMHULL 3169 (old number HU.63.C.10) PMHULL 3170 (old number HU.63.C.11) PMHULL 3171 (old number HU.63.C.12) PMHULL 3172 (old number HU.63.C.13) NEWHM 1.43.24 Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Cythere scabrocuneata Brady Cythere scabrocuneata Brady Trachyleberis raynerae Cytheroidea NEWHM 2.08.24 Trachyleberis raynerae Trachyleberis raynerae Trachyleberis raynerae Author's personal copy Material Mar Biodiv (2013) 43:363–405 Table 1 (continued) Cythere scabrocuneata T. scabrocuneata, T. lytteltonensis Trachyleberis scabrocuneata no 4 A M RLV, 1 A F RLV, 5 A M RV, Brady 4 A M LV, 2 A F LV, 1 A F RV several valves scabrocuneata 2 ? Locality Type of slide/vial Specimen(s) originally in this slide was(were) transferred by S.N.B. to the following slide Material now in this slide was originally in the following slide “Trachyleberis” lytteltonensis Lyttelton Harbour, NZ – – “Trachyleberis” lytteltonensis Lyttelton Harbour, NZ – – “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis Lyttelton Harbour, NZ Lyttelton Harbour, NZ Lyttelton Harbour, NZ – – – – – – “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis Lyttelton Harbour, NZ Lyttelton Harbour, NZ black/gray paper MPS black/gray paper MPS black/gray paper MPS black/gray paper MPS black/gray paper MPS black/gray paper MPS black/gray paper MPS – – – – Comments Both valves coated for SEM 367 Our identification ? 368 Table 1 (continued) Type of slide/vial Specimen(s) originally in this slide was(were) transferred by S.N.B. to the following slide Material now in this slide was originally in the following slide “Trachyleberis” lytteltonensis Lyttelton Harbour, NZ – – “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis Lyttelton Harbour, NZ Lyttelton Harbour, NZ Lyttelton Harbour, NZ black/gray paper MPS black plastic MPS glass with alcohol black plastic MPS – – – – – – “Trachyleberis” lytteltonensis Lyttelton Harbour, NZ black plastic MPS – – “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis “Trachyleberis” lytteltonensis Trachyleberis scabrocuneata Lyttelton Harbour, Lyttelton Harbour, Lyttelton Harbour, Lyttelton Harbour, Lyttelton Harbour, Chall. Stat. 233b – – – – NEWHM 1.54.24 – – – – – – – Trachyleberis scabrocuneata Chall. Stat. 233b – – Trachyleberis scabrocuneata Chall. Stat. 233b – – Trachyleberis scabrocuneata Chall. Stat. 233b black plastic MPS black plastic MPS glass with alcohol black plastic MPS glass with alcohol brown/gray paper MPS brown/gray paper MPS brown/gray paper MPS glass slide – – Trachyleberis scabrocuneata Trachyleberis scabrocuneata Chall. Stat. 233b Chall. Stat. 233b glass slide glass slide – – – – Trachyleberis scabrocuneata Chall. Stat. 233b Brady’s original MPS 1952.12.10.10.B – Trachyleberis scabrocuneata Chall. Stat. 233b Brady’s original MPS 1952.12.10.11.B – Trachyleberis scabrocuneata Trachyleberis scabrocuneata Chall. Stat. 233b Core OB2, Osaka Bay, Japan Core OB2, Osaka Bay, Japan Brady’s original MPS MPS 1952.12.10.12.D – – – MPS – – Trachyleberis scabrocuneata NZ NZ NZ NZ NZ Comments These specimens were previously in the glass with alcohol with the same collection number and were transferred by S.N.B to a new MPS Five of these valves were previously in the glass with alcohol with the same collection number and were transferred by S.N.B. to this MPS Valve coated for SEM Valve coated for SEM Author's personal copy Locality This material was studied by DJH This material was studied by DJH Slightly broken dorsally and ventrally Fragmented appendages, but entire male copulatory limb Empty. Identified as T. scabrocuneata because of slide 1952.12.10.8.A. S.N.B. added the B S.N.B. added the A after the slide number S.N.B. added the A after the slide number Mar Biodiv (2013) 43:363–405 Our identification Locality Type of slide/vial Specimen(s) originally in this slide was(were) transferred by S.N.B. to the following slide Material now in this slide was originally in the following slide Trachyleberis scabrocuneata Core OB2, Osaka Bay, Japan Core OB2, Osaka Bay, Japan Core OB2, Osaka Bay, Japan Core OB2, Osaka Bay, Japan Core OB2, Osaka Bay, Japan Core OB2, Osaka Bay, Japan Core OB2, Osaka Bay, Japan Chall. Stat. 233b Chall. Stat. 233b Chall. Stat. 233b Chall. Stat. 233b Chall. Stat. 233b Chall. Stat. 233b Chall. Stat. 233b MPS – – MPS – – MPS – – MPS – – MPS – – MPS – – MPS – – brown/gray paper MPS brown/gray paper MPS glass slide glass slide white paper MPS white paper MPS brown/gray paper MPS white paper MPS MPS – – – – – – – – – – – 1952.12.10.10.A 1952.12.10.11.A – – – 1952.12.10.12.C – MPS – – MPS – – MPS – – brown/gray paper MPS brown/gray paper MPS brown/gray paper MPS white plastic MPS – – – – – – – – Trachyleberis scabrocuneata Trachyleberis scabrocuneata Trachyleberis scabrocuneata Trachyleberis scabrocuneata Trachyleberis scabrocuneata Trachyleberis scabrocuneata Trachyleberis Trachyleberis Trachyleberis Trachyleberis Trachyleberis Trachyleberis Trachyleberis niitsumai niitsumai niitsumai niitsumai niitsumai niitsumai niitsumai Trachyleberis niitsumai Trachyleberis niitsumai Trachyleberis sp. cf. T. niitsumai Chall. Stat. 233b grab sample OB-40, Osaka Bay, Japan grab sample OB-40, Osaka Bay, Japan grab sample OB-40, Osaka Bay, Japan grab sample OB-40, Osaka Bay, Japan Chall. Stat. 233b Trachyleberis sp. cf. T. niitsumai Chall. Stat. 233b Trachyleberis sp. cf. T. niitsumai Chall. Stat. 233b Trachyleberis sp. cf. T. niitsumai Chall. Stat. 233b Trachyleberis niitsumai Trachyleberis niitsumai Trachyleberis niitsumai Comments Author's personal copy Our identification Mar Biodiv (2013) 43:363–405 Table 1 (continued) S.N.B. added the A after the slide number S.N.B. added the B after the slide number 369 Valves sorted in the 1970s from the dried sediment of the Challenger station 233b stored in the NHM 370 Table 1 (continued) Our identification Locality Type of slide/vial Specimen(s) originally in this slide was(were) transferred by S.N.B. to the following slide Material now in this slide was originally in the following slide Comments Trachyleberis sp. cf. T. niitsumai Trachyleberis sp. cf. T. niitsumai Trachyleberis sp.? Chall. Stat. 233b Chall. Stat. 233b Chall. Stat. 233b white paper MPS MPS glass slide – – – – – – Trachyleberis sp.? Trachyleberis sp.? Chall. Stat. 233b Chall. Stat. 233b glass slide glass slide – – – – This material was studied by DJH This material was studied by DJH No cover slip. Very fragmented appendages Very fragmented appendages Very fragmented appendages Trachyleberis Pennyella sp. Pennyella sp. Trachyleberis Trachyleberis Trachyleberis Trachyleberis Cytheroidea Cytheroidea Chall. Stat. 233b Porc. Stat. 19 Val. Stat. 12 One Tree Hill One Tree Hill One Tree Hill One Tree Hill Lyttelton Harbour, NZ Chall. Stat. off E. Moncoeur Island glass slide brown paper MPS brown paper MPS white paper MPS white paper MPS white paper MPS white paper MPS MPS MPS – – – – – – – – – – – – – – – – – – sp.? Author's personal copy raynerae raynerae raynerae raynerae Very fragmented appendages Norman collection Norman collection This material was studied by DJH This material was studied by DJH 1. All these specimens (i.e., 6 LV, 7 RV, plus 1 broken RV) are in one single micropaleontolgical slide with a rectangular surface subdivided in 24 quadrants each quadrant numbered from 1 to 24 2. Paralectotypes of T. scabrocuneata herein reassigned to a distinct species 3. These specimens were studied by Brady (1898) but not by Harding and Sylvester-Bradley (1953), therefore they are not types 4. Specimens with soft parts 5. It is not clear if Jellinek and Swanson (2003) studied these specimens 6. The following material was previously included in the paralectotypes of Cythere scabrocuneata, but can not be identified below genus level because only a few, very fragmented limbs are present Mar Biodiv (2013) 43:363–405 Abbreviations: A adult; (A-?1) probably last juvenile stage; bk. broken; Chall. Stat. HMS Challenger station; DJH David J. Horne; F female; GS glass slide; H. and S.-B. Harding and SylvesterBradley 1953; J juvenile; “live” carapaces with soft parts; LV left valve; M male; MPS micropalaeontological slide; MY Moriaki Yasuhara specimen number; NEWHM Resources Centre of the Great North Museum at the Discovery Museum, Newcastle upon Tyne; NHM Zoology collection of the Natural History Museum, London; NZ New Zealand; One Tree Hill One Tree Hill Gingin Western Australia; P. and H. Puri and Hulings, 1976; PMHULL Hull collection of the Paleontology Department (Microfossils sector) of the Natural History Museum, London; Porc. Stat. Porcupine 1869 station 19; RLV closed carapace (i.e., right and left valves); RV right valve; S.-B. Sylvester-Bradley, 1948; SEM scanning electron microscope; SNB Simone Nunes Brandão; SP soft parts; V (either left or right) valve(s); USNM collection number of the National Museum of Natural History, Smithosionian Institution (Washington DC); Val. Stat. 12 Valorous Expedition 1875 station 12; Vial vial with alcohol; W. and W. Warne and Whatley, 1996; ZMUC CRU Crustacea collection of the Zoological Museum, Copenhagen Author's personal copy Mar Biodiv (2013) 43:363–405 source in the National Museum of Natural History, Smithsonian Institution. Soft parts were imaged with a digital camera coupled to a Zeiss microscope in the Universität Hamburg. Except for the valves from Osaka Bay, which were studied by M.Y., comparisons with non-type material (mostly from Japan) were based on the few published illustrations provided by several authors (for details see synonymies in the taxonomic section). Many ostracod species have been described, without clear SEM images, from Taiwan (e.g., Hu 1984; Hu and Tao 2008), causing considerable taxonomic confusion. Taxonomic revision of Taiwanese ostracods is in progress by one of us (M.Y.) and other colleagues, and will be published elsewhere; however, a few Taiwanese records that have already been confirmed by us are included in the synonymy of Trachyleberis scabrocuneata. Abbreviations: A, adult; (A-1), last juvenile instar; bk., broken; Chall. Stat., HMS Challenger station; DJH, David J. Horne; F, female; GS, glass slide; H. and S.-B., Harding and Sylvester-Bradley 1953; ICZN, International Code of Zoological Nomenclature; J, juvenile; LV, left valve; M, male; MPS, micropalaeontological slide; MY, Moriaki Yasuhara; NEWHM, New Hancock Museum, applied to material of the G.S. Brady Collection, formerly at the Hancock Museum, Newcastle upon Tyne, and now housed in the Resources Centre of the Great North Museum at the Discovery Museum, Newcastle upon Tyne; NHM, Zoology collection of The Natural History Museum, London; NZ, New Zealand; One Tree Hill, One Tree Hill Gingin Western Australia; P. and H., Puri and Hulings 1976; PMHULL, Hull collection of the Palaeontology Department (Microfossils sector) of the Natural History Museum, London; Porc. Stat., Porcupine 1869 station 19; RLV, closed carapace (i.e., right and left valves); RV, right valve; S.-B., Sylvester-Bradley, 1948; SEM, scanning electron microscope; SMF Xe, Micropalaeontology I collection, Senckenberg Research Institute and Natural History Museum Frankfurt; S.N.B. Simone Nunes Brandão; SP, soft parts; USNM, Collection of the Department of Paleobiology, National Museum of Natural History, Smithsonian Institution; V, (either right or left) valve(s); Val. Stat. 12, Valorous Expedition 1875 station 12; Vial, vial with alcohol; W. and W., Warne and Whatley, 1996; ZMUC CRU, Crustacea collection of the Zoological Museum, Copenhagen. Results and Discussion Taxonomic review of the genus Trachyleberis Brady, 1898 and its type species Cythere scabrocuneata Brady, 1880 The names Cythere scabrocuneata or Trachyleberis scabrocuneata feature in more than 50 publications, but here we discuss only the most relevant papers (Fig. 1). For information on other papers see Table 2. In reading this review, it may be 371 helpful to keep in mind that we now consider the original material referred by Brady (1880, 1898) to Cythere scabrocuneata Brady, 1880 to include at least three species: Trachyleberis scabrocuneata s.s. (as defined herein) (black in Fig. 1), Trachyleberis niitsumai Ishizaki, 1971 (white in Fig. 1), and “Trachyleberis” lytteltonensis Harding and Sylvester-Bradley, 1953 (gray in Fig. 1). Brady (1880: 103) described as new the species Cythere scabrocuneata from three samples collected during the Challenger Expedition: (1) Station 233b, Seto Inland Sea, Japan, 15 fathoms (∼27 m); (2) Station 162, off East Moncoeur Island, Bass Strait, 38-40 fathoms (∼55-73 m); (3) Wellington Harbour, New Zealand (depth not stated). Brady (1880) did not designate a holotype, and all the specimens referred to in his 1880 publication are considered to be syntypes (which we now consider to include at least three distinct species). Some syntypes of Cythere scabrocuneata Brady, 1880 are housed in The Natural History Museum in London, while other syntypes of Cythere scabrocuneata in the G.S. Brady ostracod collection were formerly housed at the Hancock Museum, Newcastle upon Tyne, but have now been transferred to the Resources Centre of the Great North Museum at the Discovery Museum, Newcastle upon Tyne. A check on the material in this last collection in March 2012 (by DJH) confirmed that the only remaining syntypes of Cythere scabrocuneata are from the Seto Inland Sea, Japan, Station 233b; these specimens are on two slides, numbered NEWHM 1.43.22 (21 valves) and 1.43.23 (3 carapaces, 19 valves). It is worth noting that the labels on both slides indicate a depth of 14 fathoms, not 15 fathoms as stated by Brady (1880). However, the database of the NHM (http://www.nhm.ac.uk) shows 15 fathoms as the depth of station 233b. A multiple species slide, NEWHM 2.08.24, labelled as “off E. Moncoeur Island, Bass Strait, depth 38 fathoms, HMS Challenger 2.4.1874” includes the name Cythere scabrocuneata among others on the label, but it now only contains specimens of other genera and does not contain any specimens that could be assigned to C. scabrocuneata, T. lytteltonensis or T. niitsumai. “T.” lytteltonensis or T. niitsumai. No slide could be found for the third Challenger locality, Wellington Harbour, and a note in the database indicates that this slide once existed but was recorded as missing by P.S. Davis in 1991. Sylvester-Bradley (1948) and Harding and Sylvester-Bradley (1953) took their illustrated specimens and (in the latter case) designated a lectotype from “Challenger” material in The Natural History Museum, then known as the British Museum (Natural History), from “the Inland Sea of Japan at 14 fathoms”, regarding this as the only surviving type material of Cythere scabrocuneata, apparently unaware of the two above-mentioned slides at the Hancock Museum (which are from the same locality and represent part of the same syntypic series). Brady (1880, Pl. 17.5a-f, 23.2a-c) did not provide any information on the collection locality of his figured valves, Author's personal copy 372 Mar Biodiv (2013) 43:363–405 T. scabrocuneata s.s. Description of Cythere scabrocuneata Brady, 1880 from three stations of the HMS Challenger Expedition T. niitsumai Description of Trachyleberis lytteltonensis from Brady’s (1898) material Lectotype designation for T. scabrocuneata s.s. Brady 1898 Brady * illustration of T. niitsumai BUT misidentified as T. scabrocuneata Description of Trachyleberis Brady, 1898 with a single nominal species (=Cythere scabrocuneata). However, the specimens studied actually belong to a distinct species than T. scabrocuneata s.s. 1880 “T”. lytteltonensis 1953 Invalid lectotype designation for T. scabrocuneata [sic] Brady, 1898 First valid fixation of the type species of Trachyleberis, and illustration of Trachyleberis niitsumai and “T”. lytteltonensis 2003 Harding and Jellinek and Sylvester-Bradley Swanson 2013 * Herein Fig. 1 Summary of the taxonomic confusion on “Cythere scabrocuneata sensu lato”. The shades of the asterisks (on the map) and text boxes relate to the three species: black for Trachyleberis scabrocuneata sensu stricto, white for Trachyleberis niitsumai Ishizaki, 1971 and gray for Trachyleberis lytteltonensis Harding and Sylvester-Bradley, 1953. The map was made on the program Ocean Data View (Schlitzer 2012). SEM photos: © The Natural History Museum, London (2012) and it is not possible now to verify the identity of the specimens from Wellington Harbour. Eighteen years later, Brady (1898: p. 444-445, Pl. 47.1-7, 18-25) described the new genus Trachyleberis and included in it only one species: Cythere scabrocuneata Brady, 1880. Although not so designated formally by Brady himself, Cythere scabrocuneata Brady, 1880 would have become, by monotypy, the type species of the genus Trachyleberis, according to Article 68.3 of the ICZN. However, the specimens studied by Brady (1898) and identified as Cythere scabrocuneata were newly collected from Lyttelton Harbour, New Zealand (gray in Fig. 1) and subsequently shown by Harding and Sylvester-Bradley (1953) to belong to a distinct species which they described as Trachyleberis lytteltonensis. It is possible that the Lyttelton Harbour specimens may be conspecific with Brady’s (1880) Wellington Harbour syntypes of Cythere scabrocuneata, but this can no longer be checked since the Wellington Harbour specimens are lost. What is certain is that Brady’s (1898) Lyttelton Harbour specimens are not the same species as the lectotype of Cythere scabrocuneata, selected and designated from among the Challenger station 233b (Seto Inland Sea, Japan) syntypes by Harding and Sylvester-Bradley (1953). According to Article 70.3 of the ICZN, both Cythere scabrocuneata Brady, 1880 (= Trachyleberis scabrocuneata s.s. herein) and Trachyleberis lytteltonensis Harding and Sylvester-Bradley, 1953 are eligible as potential type species of Trachyleberis; the choice and its justification will be discussed below. Sylvester-Bradley (1948) recorded, described and illustrated five subfossil specimens from the syntypic series of Brady’s (1880) Cythere scabrocuneata from the Challenger station 233b (Seto Inland Sea, Japan). According to the study by one of us (S.N.B.), of these five specimens, one belongs to Trachyleberis scabrocuneata s.s. and the other four belong to Trachyleberis niitsumai (more details in the taxonomic section below). Puri (1953: p. 172) studied Trachyleberis species from North America and suggested that “it seems best to take Brady’s 1898 specimens as the actual genotypes.” Although it was published shortly before Harding and SylvesterBradley’s paper in the same year and could therefore take priority, Puri’s opinion seems to have been influenced by his mistaken supposition that the “paratypes” of Cythere scabrocuneata figured by Sylvester-Bradley (1948) were not specimens originally used by Brady (1880) (SylvesterBradley and Harding, 1954). Article 70.3 of the ICZN, Publication Our ID Stratigraphical distribution Locality (general) Locality (details) Trachyleberis scabrocuneata (Brady, 1880) no specimen studied – – – Cythere scabrocuneata n. sp. T. scabrocuneata s. s., T. niitsumai, maybe also additional species T. scabrocuneata s. s., T. niitsumai Recent Japan Type locality: Seto Inland Sea Lectotype and paralectotypes. Recent Japan Type locality: Seto Inland Sea Lectotype designation. Misidentification of some specimens of Trachyleberis sp. nov. 1 as T. scabrocuneata. T. scabrocuneata s. s., T. niitsumai T. scabrocuneata s. s. Recent Japan Recent Japan Lake Nakaumi, Shimane Prefecture Sendai Bay region, Pacific coast T. scabrocuneata s. s. Recent Japan T. scabrocuneata s. s. Recent Japan T. scabrocuneata s. s. Holocene Japan Irizuki et al., 1998 Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata T. scabrocuneata s. s. Holocene Japan Irizuki et al., 2001a Trachyleberis sp. T. scabrocuneata s. s. Holocene Japan Western coast of the Miura Peninsula Osaka Bay, Seto Inland Sea Ishizaki, 1966 Trachyleberis scabrocuneata (Brady) T. scabrocuneata s. s. Miocene Japan Goroku, W of Sendai City Poorly preserved fossil. Specimens figured by Ishizaki (1966, pl. 18.9) is strongly abraded, so it is not possible to see the ornamentation. Ishizaki, 1969 Trachyleberis scabrocuneata (Brady) T. scabrocuneata s. s. Recent Japan Lake Nakaumi Brackish lake previously named Nakanoumi. Ishizaki, 1971 T. scabrocuneata s. s. T. scabrocuneata s. s. Recent Japan Aomori Bay, Northeast Honshu Ishizaki et al., 1993 T. scabrocuneata s. s. Japan Kanazawa City T. scabrocuneata s. s. Early Pleistocene Recent Japan Urauchi Bay T. scabrocuneata s. s. Late Pleistocene Japan Fujisawa City, Kanto District T. scabrocuneata s. s. Recent Japan Suo-Nada, Seto Inland Sea T. scabrocuneata s. s. Recent Japan Suo-Nada, Seto Inland Sea Kajiyama, 1913 Trachyleberis scabrocuneata (Brady 1880) Trachyleberis ishizakii Yasuhara et al., 2002 Trachyleberis ishizakii Yasuhara et al. Trachyleberis ishizakii Yasuhara et al., 2002 Trachyleberis scabrocuneata (Brady, 1880) Cythereis yamigera (Brady) T. scabrocuneata s. s. Recent Japan Aburatsubo Bay Jellinek & Swanson, 2003 Trachyleberis sp. B T. scabrocuneata s. s. Recent Japan Type locality: Seto Inland Sea Jellinek & Swanson, 2003 Trachyleberis sp. C T. scabrocuneata s. s. Recent Japan Aburatsubo Bay, Kanagawa Prefecture notes Revision of the literature Warne, 2008 Records of Trachyleberis scabrocuneata sensu stricto Brady, 1880 Harding & Sylvester-Bradley, 1953 Trachyleberis scabrocuneata (Brady) Ikeya & Compton, 1983 Trachyleberis scabrocuneata Ikeya & Itoh, 1991 Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis sp. 1 Ikeya & Suzuki, 1992 Ikeya & Suzuki, 1992 Ikeya et al., 1987 Irizuki et al., 2006 Irizuki et al., 2009 Irizuki et al., 2010 Irizuki et al., 2011b Author's personal copy Author’s ID Mar Biodiv (2013) 43:363–405 Table 2 Summary of previous records of Trachyleberis scabrocuneata “sensu lato” off Shimane Peninsula, southwestern Japan Sea off Shimane Peninsula, southwestern Japan Sea Kawasaki Poorly preserved fossil. Lectotype 373 374 Table 2 (continued) Publication Kamiya & Nakagawa, 1993 Kamiya & Nakagawa, 1993 Author’s ID Our ID Stratigraphical distribution Locality (general) Locality (details) Trachyleberis scabrocuneata (Brady, 1880) Actinocythereis sp. T. scabrocuneata s. s. Holocene Japan Takahama-Cho, Fukui Prefecture Holocene Japan Takahama-Cho, Fukui Prefecture Trachyleberis scabrocuneata (Brady) T. scabrocuneata s. s. Pliocene and Pleistocene Japan Okubo, 1979 Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis ishizakii Yasuhara et al., 2002 Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis ishizakii T. scabrocuneata s. s. Recent Japan T. scabrocuneata s. s. Recent Japan T. scabrocuneata s. s. Japan T. scabrocuneata s. s. Middle Pleistocene Plio-Pleistocene T. scabrocuneata s. s. Holocene Japan Shimo-iida (Ukari Formation) and Dainichi (Dainichi Formation), Shizuoka Prefecture Hosonosu Sand Bank, Seto Inland Sea Fukaura (near MBS), Seto Inland Sea Naganuma Formation, Kanagawa Prefecture Sogwipo Formation, Cheju Island Tateyama, Boso Peninsula Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata T. scabrocuneata s. s. Recent Japan Type locality: Seto Inland Sea T. scabrocuneata s. s. Recent Japan T. scabrocuneata s. s. Holocene Japan Lake Nakaumi, Shimane Prefecture Yokosuka City, T. scabrocuneata s. s., T. niitsumai T. scabrocuneata s. s. Recent Japan Type locality: Seto Inland Sea Recent Japan Yasuhara & Irizuki, 2001 Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis sp. T. scabrocuneata s. s. Recent Japan Aburatsubo Bay, Kanagawa Prefecture Osaka Bay, Seto Inland Sea Yasuhara & Seto, 2006 Trachyleberis ishizakii T. scabrocuneata s. s. Holocene Japan Hiroshima Bay, Seto Inland Sea Yasuhara et al., 2002a Trachyleberis ishizakii T. scabrocuneata s. s. Holocene Japan Kitan Strait Yasuhara et al., 2002b Trachyleberis ishizakii T. scabrocuneata s. s. Holocene Japan Osaka Bay, Seto Inland Sea Okubo, 1979 Ozawa, 2009 Paik & Lee, 1988 Sasaki et al., 2007 Sylvester-Bradley, 1948 Tanaka et al., 1998 Tanaka et al., 2012a Korea Author's personal copy T. scabrocuneata s. s. Kamiya et al., 2001 notes Paralectotype Kanagawa Prefecture Ueno & Hanai, 1965 Yajima, 1992b Cythere scabrocuneata, Brady Pennyella sp. Recent NW Atlantic Brady & Norman, 1889 Cythere scabrocuneata, Brady Pennyella sp. Recent N Atlantic Brady, 1898 T. lytteltonensis Harding and Sylvester-Bradley, 1953 Acanthocythereis sp. Recent New Zealand Lyttelton Harbour Recent South China Sea – Chapman, 1914 Trachyleberis scabrocuneata (G. S. Brady) Trachyleberis scabrocuneata (Brady, 1880) Cythere scabrocuneata, Brady cf. Trachyleberididae Miocene Central Australia Mallee Chapman, 1926 Cythere scabrocuneata, G. S. Brady Trachyleberididae sp. Upper Eocene New Zealand Weka Creek Egger, 1901 Cythere scabrocuneata Brady not Trachyleberis Recent West of Australia Gou et al., 1981 Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis spp. (maybe T. cuneatelles) Trachyleberis sp. MiocenePliocene S. China Cai, 1982 Guan, 1978 Reizhou Peninsula and others Gray marls. Mar Biodiv (2013) 43:363–405 Wrong records of Trachyleberis scabrocuneata s. s. Brady & Norman, 1889 Publication Harding & Sylvester-Bradley, 1953 Hornibrook, 1952 Hou & Gou, 2007 Hou et al., 1982 Ikeya & Compton, 1983 Ikeya et al., 1985 Irizuki et al., 2001a Irizuki et al., 2002 Irizuki et al., 2005 Irizuki et al., 2006 Irizuki et al., 2010 Ishizaki, 1963 Ishizaki, 1968 Ishizaki, 1984 Jellinek & Swanson, 2003 Our ID Stratigraphical distribution Locality (general) Locality (details) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata Trachyleberis niitsumai, T. scabrocuneata s. s. Trachyleberididae Recent Japan Type locality: Seto Inland Sea Miocene to Recent New Zealand Off southeastern coast of North Island Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata Trachyleberis spp. Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata [sic] Brady, 1898 Jellinek & Swanson, 2003 Trachyleberis sp. A Liu, 1989 Okubo, 1979 Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata Sasaki et al., 2007 Sylvester-Bradley, 1948 Nakao et al., 2001 Tanaka & Nomura, 2009 Wang & Zhao, 1985 notes China Trachyleberis cuneatelles? Trachyleberis niitsumai, T. scabrocuneata s. s. Recent Japan Shibukawa, Okayama Prefecture Trachyleberis niitsumai Recent Japan Omaezaki Trachyleberis niitsumai Pleistocene Japan Trachyleberis niitsumai Holocene Japan Noma Formation, Aichi Prefecture Osaka Bay, Seto Inland Sea Trachyleberis niitsumai Japan Japan Trachyleberis niitsumai? Middle Pleistocene Middle Pleistocene Recent Japan Tahara Formation, Atsumi Peninsula Meimi Formation, Hyogo Prefecture Urauchi Bay Trachyleberis niitsumai Recent Japan Suo-Nada, Seto Inland Sea Trachyleberis sp. Middle Miocene Japan Trachyleberis niitsumai Recent Japan Sunakozaka Formation, east of Kanazawa City, Ishikawa Prefecture Uranouchi Bay Trachyleberis niitsumai M. Pleisto.Holocene Recent New Zealand Lyttelton Harbour Trachyleberis niitsumai Trachyleberis lytteltonensis Harding and SylvesterBradley, 1953 Trachyleberis niitsumai Recent Japan Seto Inland Sea Trachyleberididae Cenozoic East China Sea Donghai Trachyleberis niitsumai Japan Dainichi Formation Trachyleberis niitsumai Early Pleistocene Recent Japan Trachyleberis scabrocuneata Trachyleberis niitsumai Holocene Japan Shibukawa, Tamano-shi, Seto Inland Sea Tateyama, Boso Peninsula Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady) Trachyleberis niitsumai Recent Japan Type locality: Seto Inland Sea Trachyleberis niitsumai Late MiocenePliocene Recent Japan Maja and Aka Formations Trachyleberis niitsumai collected by I. Okubo,lat. 34° 27.1' N, long. 133° 54.2' E. Author's personal copy Irizuki & Hosoyama, 2000 Author’s ID Mar Biodiv (2013) 43:363–405 Table 2 (continued) Types from T. lytteltonensis East China Sea 375 376 Table 2 (continued) Publication Wang, 1982 Wang et al., 1988b Warne & Whatley, 1996 Yamane, 1998 Yasuhara & Irizuki, 2001 Yasuhara & Seto, 2006 Yasuhara et al., 2002a Yasuhara et al., 2005 Zheng, 1994 Dubious records of Trachyleberis scabrocuneata sensu stricto Bodergat & Ikeya, 1988 Brady, 1880 Our ID Stratigraphical distribution Locality (general) Locality (details) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady, 1880) s.s. Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata Trachyleberis spp. Quaternary North China Bohai Gulf Trachyleberis spp. Recent East China Sea Trachyleberis sp. cf. T. niitsumai Trachyleberis niitsumai Recent Japan Type locality: Seto Inland Sea Recent Japan Hiuchi-nada Bay, Seto Inland Sea Trachyleberis niitsumai Recent Japan Osaka Bay, Seto Inland Sea Trachyleberis niitsumai Holocene Japan Hiroshima Bay, Seto Inland Sea Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady) Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis niitsumai Holocene SW Japan Kitan Strait Trachyleberis niitsumai Holocene Japan Hiroshima Bay, Seto Inland Sea Trachyleberis niitsumai Holocene Japan Iyo-nada Bay, Seto Inland Sea Trachyleberis sp. – South China Sea off Hong Kong Recent Bass Straits off East Moncoeur Island Trachyleberis scabrocuneata ? Cythere scabrocuneata n. sp. ? Ise and Mikawa Bays notes No illustration. Brady, 1880 Cythere scabrocuneata n. sp. ? Recent New Zealand Wellington Harbour Brady & Norman, 1889 Cythere scabrocuneata, Brady ? Recent Atlantic coast of France Côtes des Landes This material is not in the NHM, London. Brady & Norman, 1889 Cythere scabrocuneata, Brady ? Recent Atlantic coast of France Bay of Biscay This material is not in the NHM, London. Chapman, 1915 Cythere scabrocuneata, G. S. Brady ? Recent east of Tasmania Chapman, 1915 Cythere scabrocuneata, G. S. Brady ? Recent Cythere scabrocuneata G. S. Brady ? Recent Chapman, 1919 Cythere scabrocuneata G. S. Brady ? Recent Southern Ocean Chapman & Crespin, 1928 Cythere scabrocuneata G.S.B. ? ? Crespin, 1943 Cythere scabrocuneata G.S.B. ? Tertiary Malz & Ikeya, 1982 Trachyleberis scabrocuneata (Brady, 1880) Trachyleberis scabrocuneata (Brady, 1880) Cythere scabrocuneata Brady ? Trachyleberis scabrocuneata (Brady) Okubo, 1980 Puri & Hulings, 1976 Ruan & Hao, 1988 east of Tasmania off S Australia Off SW St. Frances Island Mornington Peninsula No illustration, only lists of species found in distinct depth of the bore hole. Australia Gippsland, Victoria No illustration, only a table of species found in distinct strata. Pleistocene SW Taiwan ? Recent Japan Szekou Formation, near Tsailuhsian Seto Inland Sea ?T. scabrocuneata s. s. Recent Japan Type locality: Seto Inland Sea ?T. scabrocuneata s. s. Holocene East China Sea Okinawa Trough No illustration. In Chinese. Mar Biodiv (2013) 43:363–405 Chapman, 1919 Pacific/Southern Ocean Pacific/Southern Ocean Southern Ocean Author's personal copy Yasuhara et al., 2003 Author’s ID Author's personal copy Holotype and paratypes of T. lytteltonensis Trachyleberis lytteltonensis Harding and SylvesterBradley, 1953 Recent Recent Trachyleberis lytteltonensis sp. nov. Harding & Sylvester-Bradley, 1953 Records of other species Tanaka et al., 2012b ? no specimen studied Cythereis dorsoserrata (G. Brady) 1880 Trachyleberis scabrocuneata Müller, 1912 – Japan New Zealand Lyttelton Harbour Suruga Bay, Pacific coast of Central Japa No illustration. No illustration. many localities around Japan Pacific coast of Japan and Sea of Japan Trachyleberis scabrocuneata (Brady 1880) Ikeya & Cronin, 1993 Recent ? Trachyleberis scabrocuneata (Brady) Hanai, 1961 ? no specimen studied Hanai, 1959 no specimen studied Recent Aburatsubo Cove, Koajiro, Miura-shi, Kanagawa Prefecture Identification not possible, because illustration of only internal view is provided. 377 Pacific coast of Japan This paper consists on a list of part of the Challenger material deposited in the NHM, London. no specimen studied Not taxonomic (lists of species & materials, morphological studies, etc) Bate, 1963 no specimen studied Author’s ID Publication Table 2 (continued) Our ID Stratigraphical distribution Locality (general) Locality (details) notes Mar Biodiv (2013) 43:363–405 which must be cited for a valid fixation of a type species, does not apply to Puri’s work at all, because it only pertains to works published after 1999. Under the preceding editions of the Code, no such independent choice of a type species without recourse to the Commission could have been done. Therefore, what Puri was suggesting was simply not allowed in 1953, without bringing a Case to the Commission. Harding and Sylvester-Bradley (1953) studied the material both from the Seto Inland Sea (i.e., the syntypes from Challenger station 233b studied by Brady (1880)) and from Lyttelton Harbour, New Zealand (i.e., the specimens studied by Brady (1898) when describing the genus Trachyleberis). Harding and Sylvester-Bradley (1953) recognized that the specimens from the Seto Inland Sea belong to a different species than the specimens from Lyttelton Harbour. For the latter specimens, they described and named a new species: Trachyleberis lytteltonensis. Importantly, they designated a male specimen from Challenger station 233b (Seto Inland Sea) as the lectotype of Cythere scabrocuneata Brady 1880. The collection number provided by Harding and Sylvester-Bradley [British Museum (Nat. Hist.) B.M. 1952. 12.10.1-2] refers to two slides, still available for study at The Natural History Museum, London: NHM 1952.12.10.1 (micropalaeontological slide with both valves) and NHM 1952.12.10.2 (glass slide with fragmented soft parts, including the male copulatory limb). As noted by Warne (2008), this lectotype designation is valid because it followed the ICZN (especially Articles 74.2, 74.3 and 74.5) and therefore the name Cythere scabrocuneata Brady, 1880 (independent of its generic assignment) is fixed for the species from the Seto Inland Sea. However, one significant problem is that the Japanese specimens, which Harding and Sylvester-Bradley considered one single species (i.e., Trachyleberis scabrocuneata), actually belong to two distinct species (i.e., Trachyleberis niitsumai and Trachyleberis scabrocuneata s.s.). Even more problematic is that the valves they illustrated (Harding and Sylvester-Bradley 1953, pl. 1.5, 1.6, 1.8, 2.5, 2.6, 2.9, 2.10) belong to Trachyleberis niitsumai and not to Trachyleberis scabrocuneata s.s., and they did not illustrate their lectotype. This is probably the main cause of subsequent taxonomic confusion about Trachyleberis scabrocuneata. Additionally, Harding and Sylvester-Bradley (1953) discovered that the supposedly “empty valves” of Cythere scabrocuneata from Japan mentioned by Brady (1880) did in fact contain (dried) soft parts (see Fig. 5f herein), which they re-hydrated, dissected and illustrated in their text-figs 2025. These last specimens with the dried soft parts actually comprise the lectotype and one paralectotype of Trachyleberis scabrocuneata s.s. and six other paralectotypes that have been herein reidentified as Trachyleberis niitsumai. Unfortunately, two books widely used for ostracod generic identification, i.e., the Treatise on Invertebrate Paleontology Author's personal copy 378 (Moore 1961) and Post-Paleozoic Ostracoda (van Morkhoven 1963), illustrate Trachyleberis scabrocuneata using valves of Trachyleberis niitsumai instead of Trachyleberis scabrocuneata s.s. (Moore 1961: 334, Fig. 259; van Morkhoven 1963: 176, Fig. 271). Similarly, A Stereo-Atlas of Ostracod Shells, another widely used publication for ostracod identification but at species level, includes a supposedly definitive paper on Trachyleberis scabrocuneata, as the type species of Trachyleberis, by Ikeya and Compton (1983), which illustrates both T. scabrocuneata s.s. (their specimens IGSU O-107 and IGSU O-108) and T. niitsumai (their specimens IGSU O-109 and IGSU O-110). As noted by Ikeya and Compton (1983), Puri and Hulings (1976: p. 289) considered incorrectly that the lectotype of Cythere scabrocuneata Brady, 1880 was lost. Puri examined the sediment of the Challenger sample 233b and picked out two topotypic valves (both are now broken). Puri and Hulings (1976: pl. 26.6, 26.8) illustrated the internal and external views of one of these valves, a left valve, which probably belongs to Trachyleberis scabrocuneata s.s. Warne and Whatley (1996) designated one of these topotypes as a neotype of Trachyleberis scabrocuneata (Brady, 1880), in the mistaken belief that the lectotype was lost. However, as already discussed above, the lectotype is not lost, so the neotype is unnecessary and its designation is invalid. While trying to clarify the identity of the type species of the genus Trachyleberis, Jellinek and Swanson (2003: 20–22) added even more confusion to this topic. First, they considered, as the type species of the genus Trachyleberis, the species from the Lyttelton Harbour (Trachyleberis lytteltonensis Harding and Sylvester-Bradley, 1953), but named it erroneously as “Trachyleberis scabrocuneata BRADY 1898 [sic] (non Cythere scabrocuneata BRADY, 1880)”. The name Trachyleberis scabrocuneata Brady 1898 is invalid, because Cythere scabrocuneata was described by Brady in 1880 and, therefore, the only correct citation for the authorship of this nominal species is “Brady, 1880”. The invalid citation by Jellinek and Swanson was clearly deliberate, not a mistake, because they took care to distinguish between it and what they referred to as “Trachyleberis scabrocuneata BRADY 1880 sensu auct.” Second, Jellinek and Swanson (2003: 20—22) did not provide a list of the specimens of “Cythere scabrocuneata sensu lato” studied by them. However, in their two pages of remarks on Trachyleberis scabrocuneata, they stated that they studied “all the remaining material of Tr. scabrocuneata and Tr. lytteltonensis” housed in the Zoological Museum of Copenhagen and in the Natural History Museum, London. Based on this last statement and on their illustrations, we conclude that they studied the following material: Mar Biodiv (2013) 43:363–405 1. At least part of the syntypes of Brady’s (1880) Cythere scabrocuneata from Challenger station 233b (see their pl. 5.1—3), which Jellinek and Swanson (2003) called: (a) ‘“Trachyleberis” sp. A [= Trachyleberis scabrocuneata (BRADY 1880) sensu SYLVESTER-BRADLEY 1948]’; (b) ‘“Trachyleberis” sp. B. [= Trachyleberis scabrocuneata (BRADY 1880) sensu HARDING & SYLVESTER-BRADLEY 1953]’, in which they included the lectotype of Cythere scabrocuneata (= Trachyleberis scabrocuneata s.s.) (see below). 2. Specimens from Lyttelton Harbour, New Zealand, studied by Brady (1898), that Harding and SylvesterBradley (1953) described as Trachyleberis lytteltonensis and that Jellinek and Swanson (2003) named erroneously as “Trachyleberis scabrocuneata BRADY 1898” (see their pls. 3.1, 4.1—4). 3. ‘“Topotypic’ material from the [sic] Sea of Japan” (i.e., Recent, Aburatsubo Bay, Kanagawa Prefecture, Japan) that Jellinek and Swanson (2003, pls. 5.4, 6.1—2) call ‘“Trachyleberis” sp. C [= Trachyleberis scabrocuneata (BRADY, 1880) sensu IKEYA & COMPTON, 1983].’ Jellinek and Swanson (2003) identified as “Trachyleberis” sp. B the lectotype of Cythere scabrocuneata Brady 1880 (their pl. 5. 3). However, a lectotype is by definition a species name bearing specimen and therefore cannot be left in open nomenclature. In fact, Jellinek and Swanson (2003) dealt with at least three species: 1. Trachyleberis scabrocuneata s.s. from (a) the type locality (Challenger station 233b) (their “Trachyleberis” sp. B), and (b) subsequently collected specimens from Aburatsubo Bay, Kanagawa Prefecture, Japan (their “Trachyleberis” sp. C) (their pls. 5.3, 5.4, 6.1, 6.2). Here, we include both Jellinek and Swanson’s (2003) “Trachyleberis” sp. B and “Trachyleberis” sp. C in one single species because of the following similar morphologies of: (1) carapace; (2) male copulatory limb; (3) strongly modified claw on the distal margin of the first segment of the fifth limb (their “thoracic leg P1 showing knee-point clasper” in pl. 6.1a, 6.2a) (see Fig. 7a herein). 2. Trachyleberis lytteltonensis from Lyttelton Harbour (their “Trachyleberis scabrocuneata BRADY 1898”) (their pls. 3–4). 3. Trachyleberis niitsumai from the Challenger station 233b (their pl. 5.1, 5.2) (their “Trachyleberis” sp. A). There seems to be a mislabelling on the specimen figured in their plate 5.1, which is probably the NHM 1948.3.10.1 and not 1948.8.3.10.1 (this last number does not exist in the NHM collection). Author's personal copy Mar Biodiv (2013) 43:363–405 Jellinek and Swanson’s (2003) designation of the type species of Trachyleberis must be considered invalid for the following reasons. Firstly, a valid fixation of the type species of Trachyleberis must explicitely cite: (a) the ICZN’s Article 70.3; (b) the name previously cited; (c) the valid name of the species selected (Art. 70.3.2). Jellinek and Swanson (2003) only accomplished the second of these tasks. Secondly, they mixed (1) the lectotypification of Trachyleberis scabrocuneata s.s. (which had already been done validly in 1953) with (2) the fixation of the type species of Trachyleberis and failed to use the valid name (i.e., Trachyleberis lytteltonensis) for the species they wanted to fix as the type species. They wrongly used the term “lectotypes”, stating (op. cit.: 22): “the Lyttelton Harbour specimens must be treated as lectotypes of Trachyleberis scabrocuneata (sensu BRADY 1898)”. As noted by Warne (2008), a lectotype of Trachyleberis scabrocuneata s.s. had already been designated by Harding and Sylvester-Bradley (1953) and all subsequent designations are invalid (see ICZN’s Article 74.1.1). Fixation of the type species of Trachyleberis Brady 1898 As mentioned above, if Brady (1898) had described and illustrated specimens of Trachyleberis scabrocuneata s.s., this species would have been unquestionably fixed as the type species of Trachyleberis. However, because Brady’s 1898 specimens actually belong to Trachyleberis lytteltonensis Harding and Sylvester-Bradley, 1953, the ICZN’s Article “70.3 Misidentified type species” becomes relevant. This Article states that an author “may select, and thereby fix as type species, the species that will, in his or her judgment, best serve stability and universality, either: 70.3.1. The nominal species previously cited as type species [i.e., Cythere scabrocuneata Brady, 1880 (= Trachyleberis scabrocuneata s.s. herein)] or 70.3.2. The taxonomic species actually involved in the misidentification…” [i.e., Trachyleberis lytteltonensis Harding and Sylvester-Bradley, 1898] Here we choose the first option (i.e., 70.3.1) and hereby fix Cythere scabrocuneata Brady, 1880 (= Trachyleberis scabrocuneata s.s. herein), as defined by its lectotype from Challenger station 233b in the Seto Inland Sea, Japan, as the type species of the genus Trachyleberis Brady, 1898 (Figs. 4, 5, 6, and 7), thus confirming the designation (by monotypy) by Harding and Sylvester-Bradley (1953). Our reasons for this choice are explained below. The two species available as type species actually belong to two distinct genera, one of them being new to science and yet to be named. Therefore, the first advantage of choosing Cythere scabrocuneata Brady, 1880 379 (= Trachyleberis scabrocuneata s.s. herein) is that the genus Trachyleberis thus remains what most researchers consider it to be, and our choice serves to stabilize the application of the generic name of Trachyleberis. It could be argued that Trachyleberis lytteltonensis was nevertheless the species on which Brady actually based his description of Trachyleberis, and should, accordingly, be designated as the type species; however, since the genus to which it belongs is not Trachyleberis (as understood generally in the literature) but a new, currently monotypic genus (that will be described and named in a separate publication by Yasuhara et al.), designating lytteltonensis as the type species of Trachyleberis would require generic reassignment of species currently assigned to Trachyleberis, which would almost certainly result in further confusion. In our opinion such a decision would not best serve the interests of stability. The monotypic nature of the genus to which “Trachyleberis” lytteltonensis belongs does not facilitate understanding of common generic characters. This last genus is, moreover, only known from Recent sediments, so that no stratigraphical or paleobiogeographical information is available. Cythere scabrocuneata Brady, 1880 (= Trachyleberis scabrocuneata s.s. herein), on the other hand, has 17 fossil and Recent congeneric species (see details below). Therefore, choosing the latter as the type species of Trachyleberis provides a richer generic concept in terms of morphology, biogeography and paleobiogeography; it will also make comparisons of Trachyleberis with other trachyleberidid genera, including the possible ancestor Cythereis, easier. This is important because not only is the Cythereis–Trachyleberis relationship unclear at present, but also the generic taxonomy of Trachyleberididae in general is still confusing. Furthermore, several other Mesozoic to Recent trachyleberidid ostracod genera (e.g. Actinocythereis, Acanthocythereis), are in need of revision which will require comparisons with the type genus, Trachyleberis. An unfortunate drawback of choosing Cythere scabrocuneata Brady, 1880 (= Trachyleberis scabrocuneata s.s. herein) as the type species of Trachyleberis is the lack of soft parts in good enough condition to be described in detail (see detailed discussion below). Only a few, mostly strongly sclerotized structures (e.g., outline and strong claws of limbs, and most of the male copulatory limb) have been or can be well described and figured from the dried specimens available (Harding and Sylvester-Bradley 1953; Jellinek and Swanson 2003; Figs. 5f and 7 herein). Conversely, the appendages of “Trachyleberis” lytteltonensis are well known (Harding and Sylvester-Bradley 1953, text-figs 2–19; pls. 1.1–1.4, 1.7, 2.1–2.4, 2.7–2.8). We tried hard, unfortunately without success, to find specimens of Trachyleberis scabrocuneata s.s. with soft parts in wet scientific collections. Therefore, we are only able to provide two drawings and one photo of the few type specimens and Senckenberg specimens with dried soft parts. Author's personal copy 380 Nevertheless, it is now possible to define the genus Trachyleberis unequivocally on the basis of its valve characteristics, and we consider this to be the most important consideration since it permits many other fossil trachyleberidid genera and species (for which soft parts are also unknown) to be compared with it. The soft parts of Trachyleberis species Harding and Sylvester-Bradley (1953) illustrated and described in detail the soft parts of the paratypes of Trachyleberis lytteltonensis, but according to our definition of the genus this species can no longer be assigned to Trachyleberis. They also provided a description and drawings of specimens they assigned to Trachyleberis scabrocuneata (Brady, 1880), which included two species (T. scabrocuneata s.s. and T. niitsumai). Unfortunately, this last description and illustrations were based on fragmented limbs, which had been preserved dry since the 1870s. Additionally, according to Harding and Sylvester-Bradley’s (1953) legend “segments and setae shown lightly stippled were missing in the specimens seen, and have been added by analogy with the other species”; each of their drawings is a compilation of the observation of two or three specimens (the catalogue numbers of which they did not specify). Therefore, these illustrations cannot be used for the characterization of either T. scabrocuneata s.s. or T. niitsumai. Okubo (1979, Fig. 4) drew the female limbs (antennula, antenna, mandibula, maxillula, fifth limb, sixth limb, seventh limb and the reduced furca) of one specimen (MO-664) of the form he assigned to Trachyleberis scabrocuneata (Brady, 1880), which includes both T. scabrocuneata s.s. and T. niitsumai. Unfortunately, the figured valves (their Fig. 7a–e; SEM images) are from other specimens (MO-1072 to MO1075) collected from a different location (Shibukawa, Tamanoshi) than specimen MO-664 (from Hosonosu Sand Bank). Therefore, it is once again not possible to ascertain to which of the two species the figured and described limbs belong. Finally, Jellinek and Swanson (2003, pl. 6) provided eight optical photos of two specimens assigned by them to their “Trachyleberis” sp. C (= T. scabrocuneata s.s. according to our definition), which show the outline and details of the male fifth limb and of the male copulatory limb. We borrowed these two specimens, but on examining them we discovered that unfortunately the permanent medium of both slides containing the soft parts had dried out, making further detailed study impossible. Tanaka (2008) described Trachyleberis ikeyai from the southeastern part of the Sea of Japan (116 m depth). Remarkably, T. ikeyai resembles T. scabrocuneata s.s. in three unique characters (observation of T. scabrocuneata s.s by S.N.B.): (1) the modified, strongly sclerotized claw on the distal margin of the segment I of the male fifth limb (Fig. 5f herein) (other trachyleberidid species tend to show a simple Mar Biodiv (2013) 43:363–405 seta in this location); (2) the furca developed as four well developed, strongly barbulated setae (while T. lytteltonensis, for example, displays four reduced setae); (3) the male copulatory limb, including the (3.1) sub-rounded basal capsule, (3.2) the very elongate and slightly curved, well sclerotized ventro-distal process and (3.3) proximally curved copulatory tube. Tanaka’s (2008, Fig. 6j) drawing does not include the delicate, small, poorly sclerotized dorso-distal process of the males copulatory limb, which may either really be absent in T. ikeyai or may have been overlooked because of its very delicate nature. Despite these striking similarities in soft part morphology, the two species cannot be synonymized because their valves show conspicuous differences. The valves of females of T. ikeyai show a very similar lateral outline to T. scabrocuneata s.s., while valves of males of both species differ more conspicuously in that the former species is higher in relation to length and has a broader posterior margin than the latter species. Finally, valves of both males and females of T. ikeyai differ from T. scabrocuneata s.s. in the surface ornamentation: the former species shows distally flattened spines, while the last species shows spines with pointed tips. Unfortunately, we were unable to locate any other specimens of T. scabrocuneata s.s. and T. niitsumai with well preserved soft parts. Therefore any further description of the chaetotaxy (number, kind and location of setae and claws) of the soft parts of T. scabrocuneata s.s. awaits the discovery of new material. Revision and (paleo)biogeography of the genus Trachyleberis Brady, 1898 Trachyleberis was erected in the nineteenth century and, according to Kempf (1986, 1995), 366 species and subspecies plus 10 subgenera have been assigned to it. Another 50 species and subspecies were only tentatively included in Trachyleberis: many researchers tentatively assigned many questionable species to Trachyleberis, pending taxonomic revision of this genus (e.g., Howe and Howe 1975). As a result, the geographical and age distributions of this genus have been considered as global and from the late Cretaceous to Recent (e.g., Moore 1961; van Morkhoven 1963; Neale 1972; Bate 1975). However, the taxonomic confusion regarding the type species of Trachyleberis is now resolved, and here we revise the taxonomic concept of this genus and propose a relatively narrow concept. As a result, Trachyleberis now contains only 18 species and ranges from the Eocene to Recent (Fig. 2). Note that the taxonomy of the Taiwanese species described by Hu (e.g., Hu and Cheng 1977; Hu 1984; Hu and Tao 2008) is confused. These species are therefore only tentatively included in the list (“included species”) below, awaiting future revision. Author's personal copy Mar Biodiv (2013) 43:363–405 381 50 N Peter the Great Bay Hokkaido 1 40 N Sea of Japan 38 3 Bohai Sea Northern Limit 2 45 Yellow Sea 39 39 39 10 6 11 Honshu 36 7 Korea 21 12 1513 98 22 20 1614 2347 19 17 25 28 27 26 24 18 29 37 30 Kyushu 31 32 33 35 R 43 yu ky u Is 41 nd 42 la East China Sea 34 Oldest record s 40 41 30 N Taiwan 45 20 N 46 Ocean Data View 44 South China Sea Southern Limit 110 E 120 E 130 E 140 E 150 E Fig. 2 Map showing the revised Trachyleberis distribution. Northern and southern limits of Trachyleberis distribution and oldest Trachyleberis record are shown. Key locality names are also shown, for explanation on locality numbers (Loc. No.) see Table 3. This map was made on the program Ocean Data View (Schlitzer 2012). Northern limit: Aomori Bay (Ishizaki 1971); Southern Limit: Northern coast of the South China Sea off Hong Kong (e.g., Cai 1988); Oldest record: Saga Prefecture, Kyushu (Eocene–Oligocene: Yamaguchi et al. 2006) Comprehensive diagnosis A genus of the family Trachyleberididae distinguished by the following valve characteristics: in lateral view, valves subrectangular to subtrapezoidal, anterior margin rounded, posterior margin angulate, dorsal margin slightly concave to straight, ventral margin slightly convex to straight. Carapace strongly dimorphic, males more elongate in lateral view. Carapace laterally compressed in dorsal view. Ornamentation conspicuous, spinose, tuberculate, with prominent nodules and/or spines. Ocular ridge present. Inconspicuous ventrolateral ridge; posteromedian lateral ridge absent; antero- and postero-marginal rims absent; subcentral tubercle subdued; anteroventral cluster of four spines/tubercles inconspicuous; numerous spines on the lateral surface. In addition, (A-1) juveniles have conspicuous and parallel ventrolateral and median lateral ridges; continuous anteromarginal ridge; and more conspicuous primary reticulation than the adult. Hinge holamphidont; right valve with anterior and posterior teeth strongly projecting but not denticulate; left valve with median element a narrow, crenulate bar and a rounded projecting tooth at its anterior end. Internal snap-knob structure present at mid-length ventrally. Four elongate to suboval vertical adductors with a V-shaped frontal scar. Remarks Based on these characters, Trachyleberis is clearly distinguishable from closely related trachyleberidid genera such as Acanthocythereis Howe 1964, Actinocythereis Puri Author's personal copy 382 Mar Biodiv (2013) 43:363–405 1953 and Cythereis Jones 1849, while several species previously assigned to Trachyleberis are now easily excluded from this genus. It is beyond the scope of the present paper to discuss further details on non-Trachyleberis taxa. These will be discussed in a separate paper. According to our observations, the juvenile morphology seems to be useful for the identification of similar species of Trachyleberis and also similar trachyleberidid genera. For example, although adults of T. scabrocuneata s.s. and T. niitsumai are very similar, their (A-1) instars are clearly distinct. The (A-1) of T. scabrocuneata s.s. has subdued primary reticulation and prominent and spinous lateral ridges. In contrast, that of T. niitsumai has a distinct primary reticulation and subdued lateral ridges and spines. Additionally, in the possible ancestral genus Cythereis, the adults are very similar to their penultimate (A-1) instars, while one of the few differences between these developmental stages is the size of the valves (Szczechura 1965). This is in contrast with Trachyleberis that shows morphological differences between the adult and (A-1) stages. Included species (listed by original binomen) Holocene–Recent. Cythere scabrocuneata Brady 1880, Trachyleberis niitsumai Ishizaki, 1971, Trachyleberis ikeyai Tanaka, 2008, Trachyleberis straba Frydl, 1982, Trachyleberis verrucifera Zhao, 1985 (in Zhao et al. 1985). Plio–Pleistocene. Trachyleberis retisculpta Huang and Gou, 1982, Trachyleberis lungkangensis Hu and Cheng, 1977, Trachyleberis cuneatelles Hu, 1984, Trachyleberis fujyh Hu and Tao, 2008, Trachyleberis kingshui Hu and Tao, 2008, Trachyleberis yangtii Hu and Tao, 2008, Acanthocythereis lobulia Hu and Tao, 2008, Trachyleberis uncuneatelles Hu, 1984. Miocene. Trachyleberis leei Huh and Whatley, 1997; Trachyleberis praeniitsumai Huh and Whatley, 1997; Trachyleberis mizunamiensis Yajima, 1992; Trachyleberis shukunohorensis Yajima, 1992. Eocene–Oligocene. Trachyleberis inouei Yamaguchi, 2006 (in Yamaguchi et al. 2006). Excluded species Although several species from Upper Cretaceous strata have been assigned to Trachyleberis (Bate 1972; Neale 1975), their morphology does not justify their inclusion in Trachyleberis. For example, the Upper Cretaceous species T. raynerae Neale 1975 (Fig. 3) is distinguishable from the newly defined Trachyleberis by: (1) lacking the ocular ridge; (2) having clear anterior and posterior marginal rims; (3) showing relatively few spines on the lateral surface (especially in the anterior and posterior areas); (4) showing a distinct ventrolateral ridge continuous into anterior marginal rim; (5) showing a anteroventral cluster of four spines. In our opinion, this species should be assigned to Cythereis Jones, 1849. Fig. 3 Types of Trachyleberis raynerae Neale, 1975. Copyright: The Natural History Museum, London (2012). Holotype (NHM PMHULL 3169, old collection number HU.63.C.10): a, male RV external view. Paratype (NHM PMHULL 3172, old collection number HU.63.C.13): b, b’, male LV- b, external view; b’, internal view. Paratype (PMHULL 3170, old collection number HU.63.C.11): c, c’, (A-1) female RV- c, external view; c’, internal view. White scale bar valid for all valves Author's personal copy Mar Biodiv (2013) 43:363–405 Fig. 4 Trachyleberis scabrocuneata (Brady, 1880). All specimens from the type locality: Challenger station 233b, Seto Inland Sea, 14 fathoms. Copyright: The Natural History Museum, London (2012). Adult male lectotype* (NHM 1952.12.10.1): a, RVexternal view; b, LVexternal view. Paralectotypes: c-l. c, c’, female RV (NHM 1948.3.10.4 768)-c, external view, c’, dorsal view; d, d’, female LV (NHM 1952.12.10.10.A)- d, external view, d’, dorsal view; e, e’, male RV (NHM 1952.12.10.11.A)e, external view, e’, dorsal view; f, f’, male LV (NHM 1952.12.10.10.A)- f, 383 external view, f’, dorsal view; g, (A-1) juvenile (NHM 1952.12.10.11.A), RV external view; h, (A-1) juvenile (NHM 1952.12.10.10.A), LV external view. * The low quality of the SEM photos of the lectotype of Trachyleberis scabrocuneata is due to fact that these valves are only partly coated with gold. To avoid damage to this important specimen, we and Ms Miranda Lowe (curator of the Challenger collection in the Natural History Museum) decided not to re-coat the lectotype. Better photos of some paralectotypes are provided herein 384 Table 3 Localities where the revised genus Trachyleberis occurs (see Fig. 2) Loc. no. Locality Formation Age T. scabrocuneata T. niitsumai Recent Middle Miocene Late Pleistocene Middle Miocene Recent Late Miocene Late Miocene Middle Pleistocene + + NE Honshu, Japan NE Honshu, Japan NE Honshu, Japan NE Honshu, Japan NE Honshu, Japan NE Honshu, Japan NE Honshu, Japan C Honshu, Japan 8 8 8 9 9 9 9 10 11 11 11 11 12 13 14 14 14 15 Boso Peninsula Boso Peninsula Boso Peninsula Kawasaki Miura Peninsula Miura Peninsula Aburatsubo Bay Noto Peninsula Kanazawa Kanazawa Kanazawa Kanazawa Fukui Nagano Kakegawa Omaezaki Lake Hamana-ko Gifu C C C C C C C C C C C C C C C C C C 15 15 16 16 17 Gifu Gifu Chita Peninsula Atsumi Peninsula Osaka Bay C Honshu, Japan C Honshu, Japan C Honshu, Japan C Honshu, Japan SW Honshu, Japan surface deposit Kadonosawa F. Anden F. Moniwa F. surface deposit Hanawa F. Hanawa F. Jizodo F., Yabu F. (drilling core) Numa F. Numa F. Kamiiwahashi F. drilling core drilling core Naganuma F. surface deposit Hiradoko F. Omma F. Omma F. Omma F. Sunakozaka F. Takahama F. Arakida F. Dainichi F. surface deposit surface deposit Akeyo F., Agi F. Tōyama F. Shukunohora F. Shukunohora F. Noma F. Tahara F. drilling core 17 17 17 Osaka Bay Osaka Bay Osaka Bay SW Honshu, Japan SW Honshu, Japan SW Honshu, Japan surafce deposit drilling core drilling core Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Holocene Holocene Middle Pleistocene Holocene Holocene Middle Pleistocene Recent Late Pleistocene Early Pleistocene Early Pleistocene Early Pleistocene Middle Miocene Holocene Early Miocene Early Pleistocene Recent Recent Early Miocene Middle Miocene Middle Miocene Middle Pleistocene Middle Pleistocene Middle PleistoceneHolocene Recent Holocene Holocene + + + (+) + (+) (+) + + + + + + (+) (+) + + + (+) + + + + cf. + (+) (+) (+) + + + + + + + + (+) + + + + + + Ishizaki, 1971 Irizuki and Matsubara, 1994 Ishizaki and Matoba, 1985 Ishizaki, 1966 Ikeya and Itoh, 1991 Irizuki et al., 2001b Yamaguchi and Hayashi, 2001 Irizuki et al., 2011a Sasaki et al., 2007 Frydl, 1982 Yajima, 1978 Ikeya et al., 1987 Irizuki et al., 1998 Ozawa, 2009 Kajiyama, 1913 Kamiya et al., 2001 Ishizaki et al., 1993 Ozawa, 1996 Kamiya et al., 1996 Ishizaki, 1963 Kamiya and Nakagawa, 1993 Yamada et al., 2001 Nakao et al., 2001 Ikeya et al., 1985 Ikeya and Hanai, 1982 Irizuki et al., 2004 + + + Yajima, 1988 Yajima, 1992a Irizuki and Hosoyama, 2000 Irizuki et al., 2002 Ishizaki, 1984 +, cf. + + Yasuhara and Irizuki, 2001 Yasuhara et al., 2002b Yasuhara et al., 2004 Author's personal copy Aomori Bay Ninohe Oga Peninsula Sendai Sendai Bay Fukushima Fukushima Saitama Reference Mar Biodiv (2013) 43:363–405 1 2 3 4 5 6 6 7 others Loc. no. Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Honshu, Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Japan Formation Age T. scabrocuneata T. niitsumai drilling core Meimi F. driiling core Yoshino F. surface deposit surface deposit surface deposit surface deposit surface deposit short core drilling core drilling core surface deposit short core surface deposit Holocene Middle Pleistocene Holocene Middle Miocene Recent Recent Recent Recent Recent Recent Holocene Holocene Recent Recent Recent + + + + Recent Recent Late Eocene-Early Oligocene Late Pliocene-Early Pleistocene Late Pliocene-Early Pleistocene Recent Early Pleistocene Late Miocene-Pliocene Recent Middle Miocene Middle Miocene Middle Miocene Middle Miocene Late-Middle Pleistocene Middle PleistoceneHolocene Middle PleistoceneHolocene Holocene Shikoku Shikoku Kyushu, Japan surface deposit surface deposit Kishima F. 31 Miyazaki Kyushu, Japan Sadowara-Takanabe F. 31 Miyazaki Kyushu, Japan Takanabe F. 32 33 34 35 36 36 36 36 37 38 Urauchi Bay Tanega-shima Island Kume-jima Island Sekisei-sho Pohang Pohang Pohang Pohang Cheju-do Island Bohai Sea Kyushu, Japan Kyushu, Japan Ryukyu Islands, Japan Ryukyu Islands, Japan Korea Korea Korea Korea Korea N China surface deposit Masuda F. Maja F., Aka F. surface deposit Chunbuk F. Chunbuk F. Chunbuk F. Chunbuk F. Sogwipo F. drilling core 38 Bohai Sea N China drilling core 39 Jiangsu C China Donghai Group (drilling core) + + + + + (+) + (+) + + (+) + + + + + + + + (+) + + + (+) Yasuhara et al., 2002a Irizuki et al., 2005 Irizuki et al., 2001a Yajima, 1988 Ikeya and Suzuki, 1992 Ishizaki, 1969 Tanaka et al., 1998 Okubo, 1979 Yamane, 1998 Yasuhara et al., 2003 Yasuhara and Seto, 2006 Yasuhara et al., 2005 Irizuki et al., 2010 Irizuki et al., 2011b Tanaka, 2008 + (+) + Ishizaki, 1968 Irizuki et al., 2008 Yamaguchi et al., 2006 + Iwatani and Irizuki, 2008 + + + cf. + + + + ? Reference Iwatani et al. 2011 (+) + (+) + + + + + + Irizuki et al., 2006 Irizuki, 2004 Tanaka and Nomura, 2009 Tabuki et al., 1987 Huh and Paik 1992 Huh and Paik, 1992 Huh, 1994 Huh and Whatley, 1997 Lee and Paik, 1992 Wang, 1982 + Wang, Q. et al., 1988 ? Hou et al., 1982 385 SW SW SW SW SW SW SW SW SW SW SW SW SW SW SW 29 29 30 Kitan Strait Hyogo Hyogo Tsuyama off Shimane Lake Nakaumi Lake Nakaumi Seto Inland Sea Hiuchi-Nada Hiroshima Bay Hiroshima Bay Iyo-Nada Kasado Bay Suo-Nada off Yamaguchi, Tsushima Uranouchi Bay Uranouchi Bay Saga others Author's personal copy 18 19 19 20 21 22 22 23 24 25 25 26 27 27 28 Locality Mar Biodiv (2013) 43:363–405 Table 3 (continued) Author's personal copy + Brady, 1880 ? ? Gou et al., 1983 + ? ? ? ? + + + + + Zheng, 1987 Fujian Institute of Oceanography, 1988 Cai, 1982 Zhao et al., 1985 Zheng, 1994 Cai, 1988 Gou et al., 1981 + + ? Wang, P. et al., 1988 Liu, 1989 + + + Numerous species from Paleogene strata in New Zealand and southeastern USA (e.g., Hornibrook 1953; Howe and Howe 1973; Ayress 1995) have been included in Trachyleberis. Here again, we exclude these species from the newly diagnosed Trachyleberis. For example, Howe and Chambers reported Trachyleberis? montgomeryensis (Howe and Chambers, 1935) from the Eocene–Oligocene of Mississippi, USA, but its adult specimens lack the ocular ridge and have relatively few spines on their lateral surfaces. This species also shows distinct ventrolateral ridge and subcentral tubercle, and the juveniles lack primary reticulation, median lateral ridge, and ocular ridge (Howe and Howe 1973; 1975). The Recent New Zealand species T. lytteltonensis Harding and Sylvester-Bradley, 1953 differs from Trachyleberis by: (1) its oval-shaped frontal scar; (2) its weakly developed multifurcate tubercles on the lateral surface; (3) lacking the ocular ridge; (4) a distinctly different form of male copulatory limb, as previously noted by Harding and Sylvester-Bradley (1953). T. thomsoni Hornibrook, 1952 differs from Trachyleberis by: (1) lacking the ocular ridge; (2) having relatively few spines on the lateral surface; (3) showing the distinct ventrolateral ridge; (4) showing the anteroventral cluster of four spines. Japan RSeto Inland Sea 47 (+) shows that species names are only listed in tables; + shows that photographs of species are present Pliocene Wanglougang F. (drilling core) Recent S China 46 S S S S S China China China China China South China Sea South China Sea South China Sea South China Sea Reizhou Peninsula and others Reizhou Peninsula 44 44 44, 45 45 46 C China C China Fujian Taiwan Strait 42 43 + Recent Recent Recent Recent Miocene-Pliocene surface deposit surface deposit surface deposit surface deposit drilling core Holocene Donghai Group (drilling core) drilling core surface deposit C China East China Sea East China Sea 40 41 Holocene Recent Age Formation Locality Loc. no. Table 3 (continued) T. scabrocuneata Reference others Mar Biodiv (2013) 43:363–405 T. niitsumai 386 Questionable assignments Trachyleberis haikangensis Gou and Zheng, 1983 (Pliocene), Trachyleberis awajiensis Yamaguchi, 2005 (in Yamaguchi et al. 2005) (Eocene), Cythere scabra Münster, 1830 (Eocene), Trachyleberis wenzhouensis Chen, 1990 (in Yang et al. 1990: Eocene). Possible synonymies Costa sinensis Hu, 1981 (Pleistocene) and Trachyleberis costus Hu, 1983 (Plio–Pleistocene) may be juveniles of a Trachyleberis species. Invalid binomens (nomina nuda because they were introduced in an unpublished thesis): Trachyleberis koreanica Lee, 1990 (Pleistocene), Trachyleberis sejongi Lee, 1990 (Pleistocene). Geographic and Stratigraphic occurrence The genus Trachyleberis as defined herein includes 18 species (instead of more than 400) and is restricted to shallow marine environments of mid-latitudes of the northwestern Pacific (Fig. 2). Its southernmost occurrence is the northern coast of the South China Sea off Hong Kong (Zhao et al. 1985; Cai 1988; Zheng 1994) and there are no confirmed records from the tropical Western Pacific (Fig. 2). For example, no certain Trachyleberis records are reported from Hainan Island (Gou 1990), Vietnam (Tanaka et al. 2009), Indonesia (Dewi 1997), and any other lower latitude areas (e.g., Kingma 1948; Hanai et al. 1980; Whatley and Zhao 1987, 1988; Reeves et al. 2007). Kingma (1948) identified Cythereis dacyi Howe and Author's personal copy Mar Biodiv (2013) 43:363–405 Fig. 5 Valves and male copulatory organ of Trachyleberis scabrocuneata (Brady, 1880), Trachyleberis niitsumai Ishizaki, 1971 and “Trachyleberis” lytteltonensis Harding and Sylvester-Bradley, 1953. Copyright: The Natural History Museum, London (2012). Trachyleberis scabrocuneata - Paralectotypes: a-e. a, e, female LV (NHM 1952.12.10.10.A), a, internal view, e, adductor muscle scars; b, female 387 RV (NHM 1948.3.10.4 768); c, male LV (NHM 1952.12.10.10.A), internal view. d, g, male RV (NHM 1952.12.10.11.A), internal view, g, adductor muscle scars. Lectotype: f, male copulatory limb (NHM 1952.12.10.2). Trachyleberis niitsumai - h, female RV (USNM 560190, TRA1045), adductor muscle scars. “Trachyleberis” lytteltonensis Paralectotype: i, female LV (NHM 1952.12.9.2), adductor muscle scars Author's personal copy 388 Fig. 6 Trachyleberis scabrocuneata (Brady, 1880). Osaka Bay, Seto Inland Sea, Holocene sediment core OB2 (ZMH K-42870): a, b, g, adult male; c, d, h, adult females; e, f, (A-1) juveniles. a, c, e, RV Mar Biodiv (2013) 43:363–405 external view; b, d, f, LV external view; c’, RV internal view, c”, c”’, details of hinge of c’; g, LV internal view. White 500 μm scale bar valid for all entire valves Author's personal copy Mar Biodiv (2013) 43:363–405 389 Fig. 7 Trachyleberis scabrocuneata (Brady, 1880). Copyright: The Natural History Museum, London (2012). Abuaratsubo Bay, Kanagawa Prefecture (SMF Xe 20217): a, b, adult male. a, fifth limb; b, furca (barbulae of 3 setae not shown) a b 100 µm 10 µm © The Natural History Museum, London (2012) Law 1936 in Java Island, Indonesia; Hanai et al. (1980) considered this identification to be questionable and cited the Javanese species as “Trachyleberis? sp.” However, this generic assignment is questionable, as Hanai et al. (1980) suggested. Kingma’s sketch shows the Javanese species to be more similar to other trachyleberidid genera such as Legitimocythere or Echinocythereis. Herrig (1977: pl. 2.6– 2.7) reported Trachyleberis scabrocuneata from the Plio– Pleistocene of northern Vietnam, but according to his plate Herrig’s specimens are not even Trachyleberis. The northern limit of the revised Trachyleberis’ geographic distribution is Aomori Bay, in the northeastern Honshu Island, Japan (Fig. 2; Ishizaki 1971). No certain Trachyleberis records are known from any areas further north, including Hokkaido (Ozawa 2004; Ikeya and Cronin 1993: http://geochange.er.usgs.gov/data/PRISM/ostracodes/ pacific/), Alaska (Brouwers 1993), and the Peter the Great Bay (Schornikov 2006) (Fig. 2). Within this latitudinal range, Trachyleberis is quite common, and known widely from various Miocene–Holocene strata as well as modern bottom sediments in Honshu Island (Ishizaki 1968, 1971; Hanai et al. 1977; Ikeya and Cronin 1993; Ozawa and Kamiya 2001; Yasuhara and Irizuki 2001; Irizuki et al. 1998, 2001a, 2001b, 2004, 2008, 2010, 2011a, 2011b; Tanaka 2008), Kyushu Island (Irizuki et al. 2006; Iwatani and Irizuki 2008), the Ryukyu Islands (Tabuki et al. 1987), South Korea (Huh and Whatley 1997), East China Sea (Wang and Zhao 1985; Wang et al. 1988b), South China Sea (Cai 1982, 1988; Zheng 1994) and Taiwan (Hu and Tao 2008) (see Fig. 2 for locality names). Evolution of Trachyleberis The earliest records (Eocene– Oligocene fossils from Kyushu, Japan) occur in the centre of the distribution of the genus (Fig. 2), suggesting that Trachyleberis may have originated in Kyushu or its surrounding region and dispersed in both south and north directions. Trachyleberis most likely evolved directly or indirectly from Cythereis that is well known from the Tethyan Cretaceous. However, fossil ostracod records are rare in the Paleogene of the northwestern Pacific (Yamaguchi and Kamiya 2007) and almost absent in the Cretaceous (Ishizaki T. scabrocuneata 0.60 Male Female LV height (mm) Fig. 8 Size of Trachyleberis scabrocuneata sensu stricto, “T.” lytteltonensis Harding and Sylvester-Bradley, 1953 and T. niitsumai Ishizaki, 1971 The bathymetric distribution of Trachyleberis is deeper towards its southern limit in the South China Sea, where it is reported only from water depths greater than 50 m (Cai 1982, 1988; Li 1985). In contrast, it is fairly common over a range of depths from>50 m water depth (Tanaka 2008) to as shallow as a few meters in Japanese waters and the East China Sea (e.g., Ishizaki 1968, Wang et al. 1988b; Yasuhara and Irizuki 2001). This bathymetric distribution may be related to the temperature preference or tolerance of the genus. The earliest fossil record of the genus is Trachyleberis inouei from Eocene–Oligocene strata in Kyushu Island, Japan (Fig. 2; Yamaguchi et al. 2006). This early Trachyleberis species is more regularly reticulated with less prominent spines than extant species such as T. scabrocuneata s.s. and T. niitsumai, as are some Miocene species (e.g., T. leei and T. mizinamiensis) (e.g., Yajima 1992a; Irizuki and Matsubara 1994; Huh and Whatley 1997; Irizuki et al. 2004). These morphological characters may be regarded as primitive and are still shared by juveniles of Recent Trachyleberis (Figs 4g–h, 6e–f, 9e–f). Juvenile 0.50 “T.” lytteltonensis Male Female 0.40 Juvenile T. niitsumai 0.30 Male 0.65 0.75 0.85 0.95 LV Length (mm) 1.05 Female Juvenile Author's personal copy 390 Fig. 9 Trachyleberis niitsumai Ishizaki 1971. Paralectotypes of T. scabrocuneata herein reidentified as Trachyleberis niitsumai. Copyright of a, b: The Natural History Museum, London (2012). Challenger station 233b, Seto Inland Sea, 14 fathoms. a, a’, b, b’, (NHM 1952.12.10.10.B ); a, a’, male RV: a, external view, a’, internal view. b, b’, female LV: b, external view, b’, internal view. Osaka Bay, Seto Mar Biodiv (2013) 43:363–405 Inland Sea, modern grab sample OB-40: c, c’, adult female (USNM 560190, TRA1045); d, d’, adult male (USNM 560189, TRA1044); e, f, (A-1) juveniles (USNM 560192 , TRA1047 and USNM 560191, TRA1046, respectively). c, e, RV external view; c’, RV internal view; d, f, LV external view; d’, LV internal view Author's personal copy Mar Biodiv (2013) 43:363–405 Fig. 10 “Trachyleberis” lytteltonensis Harding and Sylvester-Bradley, 1953. Copyright of a-f: The Natural History Museum, London (2012). All specimens from the type locality: Lyttelton Harbour, New Zealand, 2–9 m (1–5 fathoms). Male holotype (NHM 1952.12.9.1): a, RVexternal view; b, LV external view. Paratypes: c-f. c-d, single male specimen (NHM 391 1952.12.9.2)-c, RVexternal view, c’, RV internal view, d, LVexternal view, d’, LV internal view; e, female (NHM 1952.12.9.6) RV external view; f, female LV (NHM 1952.12.9.7) external view. Not types but from the type locality (ZMUC CRU 3631): g-i. g, male RV dorsal view; h, male LV dorsal view; i, female LV dorsal view Author's personal copy 392 Mar Biodiv (2013) 43:363–405 (Tables 1, 2 and 3, Figs. 4, 5a-g, 6, 7 and 8) 1880 in part non 1889 non 1898 non 1901 1912 in part 1913 non 1914 Fig. 11 Pennyella sp. from the northern Atlantic studied by Brady and Norman (1889). Copyright: The Natural History Museum, London (2012). NHM 1900.3.6.237: a, LV external view; b, LV internal view 1992). Thus, it is not possible at present to discuss further the origin of Trachyleberis. In particular, more research is needed in the tropical western Pacific region, where fossil ostracod data are sparse, to bridge the gap between the well-studied fossils from Japan and those of New Zealand–Australia. Independently, Holden (1964) recorded a trachyleberidid species from the Upper Cretaceous of California, USA, which is similar to Trachyleberis in lateral outline and in having an ocular ridge. However, this Californian species (Actinocythereis allisoni Holden, 1964) differs from Trachyleberis in having: (1) conspicuous anterior and posterior marginal rims; (2) a distinctive ventrolateral ridge, subcentral tubercle, and anteroventral cluster of four tubercles; (3) relatively few spines on the lateral surface. Therefore, it is still uncertain whether the ancestor came from the south or from the north. Systematics Class Ostracoda Latreille, 1806 (Higher classification based on Horne et al. 2002) Subclass Podocopa Sars, 1866 Order Podocopida Sars, 1866 Suborder Cytherocopina Baird, 1850 Superfamily Cytheroidea Baird, 1850 Family Trachyleberididae Sylvester-Bradley, 1948 Genus Trachyleberis Brady, 1898 (Tables 1, 2 and 3, Figs. 1, 2, 3, 4, 5, 6, 7, 8, 9, 10 and 11) Trachyleberis scabrocuneata (Brady, 1880) (= Trachyleberis scabrocuneata sensu stricto herein) ? 1915 ? 1919 non 1926 ? 1928 ? 1943 1948 in part non 1952 1953 in part ? 1961 1963 in part? non 1963 1965 in part non 1966 non 1968 1969 1971 ? 1976 Cythere scabrocuneata sp. nov.: Brady, 103, non pI. 17, figs. 5a-f, non pI. 23, figs. 2a-c. Cythere scabrocuneata Brady, 1880: Brady and Norman, 154-155. Trachyleberis scabrocuneata (Brady, 1880): Brady, 444-445, pI. 47, figs. 7, 18-25. Cythere scabrocuneata Brady, 1880: Egger, 441, pl. 8, figs. 1-3. Cythereis dorsoserrata (G. Brady,1880): Müller, 351. Cythereis yamigera [sic] (Brady): Kajiyama, 12, pl. 1, figs. 64-66. [typographical error for hamigera] Cythere scabrocuneata Brady, 1880: Chapman, 40, pI. 5, fig. 24. Cythere scabrocuneata Brady 1880: Chapman, 43. Cythere scabrocuneata Brady, 1880: Chapman, 28. Cythere scabrocuneata Brady, 1880: Chapman, 98, 102. Cythere scabrocuneata Brady, 1880: Chapman and Crespin. Cythere scabrocuneata, G. S. Brady: Crespin : 100. Trachyleberis scabrocuneata (Brady, 1880): Sylvester-Bradley, 794-795, non pl. 122, figs. 13-18. Trachyleberis scabrocuneata (Brady, 1880): Hornibrook, 32-33, pl. 3, figs. 38, 39, 48. Trachyleberis scabrocuneata (Brady, 1880): Harding and Sylvester-Bradley, 11-15, ?textfigs. 20-25, non pl. 1, figs. 5, 6, 8, non pl. 2, figs. 5, 6, 9,10. Trachyleberis scabrocuneata (Brady, 1880): Hanai, 373, text-fig. 14, fig. 2. Cythere scabrocuneata Brady, 1880: Bate, 84, Table 3. Trachyleberis scabrocuneata (Brady, 1880): Ishizaki, 31-32, pl. 2, figs. 10, 20a-d. Trachyleberis scabrocuneata (Brady, 1880): Ueno and Hanai, 455, fig. 423.3, non fig. 423.1-2 Trachyleberis scabrocuneata (Brady, 1880): Ishizaki, 156, pI. 18, fig. 9. Trachyleberis scabrocuneata (Brady, 1880): Ishizaki, 38, pls. 9, figs. 13–14. Trachyleberis scabrocuneata (Brady, 1880): Ishizaki, 221-222, pl. 26, fig. 8. Trachyleberis scabrocuneata (Brady, 1880): Ishizaki, 92-93, pl. 4, fig. 16. Cythere scabrocuneata Brady, 1880: Puri and Hulings, 289, pl. 26, figs. 6, 8. Author's personal copy Mar Biodiv (2013) 43:363–405 non 1977 non 1978 1979 in part ? 1979 ? 1980 ? 1981 non 1981 ? 1982 non 1982 non 1982 non 1982 non 1983 1983 in part non 1984 non 1985 non 1985 ? 1986 1987 1988 non 1988b ? 1988 ? 1988 non 1989 1991 1992 Trachyleberis scabrocuneata (Brady, 1880): Herrig, 1261-1262, pl. 2, figs. 6, 2-7. Trachyleberis scabrocuneata (Brady, 1880): Guan, 291, fig. 85, pl. 75, fig. 20. Trachyleberis scabrocuneata (Brady, 1880): Okubo, 149-151, figs. 7a–b, ?figs. 4, 7e, non figs. 7c-d. Trachyleberis scabrocuneata (Bradley) [sic]: Hu, 66. Trachyleberis scabrocuneata (Brady, 1880): Okubo, 412. Trachyleberis scabrocuneata (Brady, 1880): Hu, 67. Trachyleberis scabrocuneata (Brady, 1880): Gou et al. 175, pl. 84, figs. 20-23. Trachyleberis scabrocuneata (Brady, 1880): Malz and Ikeya, pl. 1, figs. 4-5. Trachyleberis scabrocuneata (Brady, 1880); Wang, pl. fig. 29. Trachyleberis scabrocuneata (Brady, 1880): Cai, pl. 3, figs. 40-41. Trachyleberis scabrocuneata (Brady, 1880): Hou et al. 223-224, pl. 83, figs. 1-5. Trachyleberis scabrocuneata (Brady, 1880): Gou et al. 81-82, pl. XII, figs. 14-23. Trachyleberis scabrocuneata (Brady, 1880): Ikeya and Compton, 119-126, pl. 10, 120, figs. 1-4, pl. 10, 126, figs. 1-4; non pl. 10, 120, fig. 5, pl. 10, 122, figs 1-5, pl. 10, 124, figs. 1-5, pl. 10, 126, fig. 5. Trachyleberis scabrocuneata (Brady, 1880): Ishizaki, 42, pl. 1, fig. 5. Trachyleberis scabrocuneata (Brady, 1880): Ikeya et al., pl. 3, fig. 13. Trachyleberis scabrocuneata (Brady, 1880): Wang and Zhao, pl. 7, fig. 6. Trachyleberis scabrocuneata (Brady, 1880): Hu, 125. Trachyleberis scabrocuneata (Brady, 1880): Ikeya et al. fig. 5. Trachyleberis scabrocuneata (Brady, 1880): Paik and Lee, 550, pl. 2, fig. 11. Trachyleberis scabrocuneata (Brady, 1880): Wang et al. 245, pl. 42, figs. 14-16. Trachyleberis scabrocuneata (Brady, 1880): Bodergat and Ikeya, 424. Trachyleberis scabrocuneata (Brady, 1880): Ruan and Hao, 357, pl. 64, fig. 32. Trachyleberis scabrocuneata (Brady, 1880): Liu, 153, pl. 168, fig. 11. Trachyleberis scabrocuneata (Brady, 1880): Ikeya and Itoh, 126, fig. 24C. Trachyleberis scabrocuneata (Brady, 1880): Ikeya and Suzuki, pl. 9, fig. 3. 393 1992 1992b 1993 1993 1993 ? 1993 1993 non 1994 non 1996 1998 1998 non 1998 non 2000 2001 2001a non 2001a non 2001 2001 non 2001 2002a non 2002a non 2002b 2002b non 2002 non 2003 Trachyleberis sp.: Ikeya and Suzuki, pl. 9, fig. 4. Trachyleberis scabrocuneata (Brady, 1880): Yajima, 235, figs. 1–7 in unlabeled plate. Actinocythereis sp.: Kamiya and Nakagawa, 129, pl. 4, fig. 8. Trachyleberis scabrocuneata (Brady, 1880): Kamiya and Nakagawa, 129, pl. 4, fig. 7. Trachyleberis scabrocuneata (Brady, 1880): Ishizaki et al. fig. 7c. Trachyleberis scabrocuneata (Brady, 1880): Ikeya and Cronin, Table 1, text-fig. 4, appendix 1. [no taxonomy, but paleoceanography]. Actinocythereis sp.: Kamiya and Nakagawa, 129, pl. 4, fig. 8. Trachyleberis scabrocuneata (Brady, 1880): Zheng, pls. 58, figs. 17-18, 59, figs 1, 2. Trachyleberis scabrocuneata (Brady, 1880) s.s.: Warne and Whatley, 163-164. Trachyleberis scabrocuneata (Brady, 1880): Irizuki et al. 7, fig. 2.13. Trachyleberis scabrocuneata (Brady, 1880): Tanaka et al. pl., fig. 24. Trachyleberis scabrocuneata (Brady, 1880): Yamane, pl. 12, fig. 5. Trachyleberis scabrocuneata (Brady): Irizuki and Hosoyama, fig. 3.10. Trachyleberis scabrocuneata (Brady, 1880): Kamiya et al. 103, fig. 18.18. Trachyleberis sp.: Irizuki et al. figs. 3-8. Trachyleberis scabrocuneata (Brady, 1880): Irizuki et al. fig. 3.7. Trachyleberis scabrocuneata (Brady, 1880): Nakao et al. fig. 11.16. Trachyleberis sp.: Yasuhara and Irizuki, 95, pl. 12, figs. 9-13. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara and Irizuki, pl. 12, figs. 1-6. Trachyleberis ishizakii sp. nov.: Yasuhara et al. 93, figs. 7.1–10. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara et al. fig. 3.22. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara et al. fig. 3.3. Trachyleberis ishizakii Yasuhara et al. 2002: Yasuhara et al. fig. 3.4. Trachyleberis scabrocuneata (Brady, 1880): Irizuki et al. pl. 1, fig. 3. Trachyleberis scabrocuneata Brady 1898 [= Trachyleberis lytteltonensis Harding and Sylvester-Bradley 1953] [sic]: Jellinek and Author's personal copy 394 non 2003 2003 2003 non 2003 non 2005 non 2005 non 2006 2006 2006 non 2006 non 2007 non 2007 2007 non 2008 ? 2008 2008 non 2009 2009 2009 Mar Biodiv (2013) 43:363–405 Swanson, 20-23, pl. 3, figs. 1-3, pl. 4, figs. 1-4. “Trachyleberis” sp. A [= Trachyleberis scabrocuneata (Brady, 1880) sensu SylvesterBradley, 1948]: Jellinek and Swanson, pl. 5, figs. 1-2. [= T. niitsumai] “Trachyleberis” sp. B [= Trachyleberis scabrocuneata (Brady, 1880) sensu Harding and Sylvester-Bradley 1953]: Jellinek and Swanson, pl. 5, fig. 3. [= T. scabrocuneata s.s.] “Trachyleberis” sp. C [= Trachyleberis scabrocuneata (Brady, 1880) sensu Ikeya and Compton, 1983]: Jellinek and Swanson, pl. 5, fig. 4, pl. 6, figs. 1, 2. [= T. scabrocuneata s.s.] Trachyleberis scabrocuneata (Brady, 1880): Yasuhara et al. fig. 2.14. Trachyleberis scabrocuneata (Brady, 1880): Irizuki et al. fig. 4.2. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara et al. fig. 4.8. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara and Seto, fig. 5e–f. Trachyleberis ishizakii Yasuhara et al. 2002: Yasuhara and Seto, fig. 5g–h. Trachyleberis ishizakii Yasuhara et al. 2002: Irizuki et al. fig. 7.3. Trachyleberis scabrocuneata (Brady, 1880): Irizuki et al. fig. 7.4. Trachyleberis scabrocuneata (Brady, 1880): Hou and Gou, pl. 172, fig. 13, pl. 176, figs. 1, 2, 4–8, pl. 227, fig. 30. Trachyleberis scabrocuneata (Brady, 1880): Sasaki et al. fig. 5.20. Trachyleberis ishizakii Yasuhara et al. 2002: Sasaki et al.: fig. 5.13. Trachyleberis scabrocuneata (Brady, 1880): Hu and Tao, 267, pl. 17, figs. 10, 11, 13, 14, 20, pl. 36, figs. 9, 13, 14, pl. 46, fig. 15, pl. 55, figs. 1-3, 6, 15, pl. 69, figs. 10, 11, pl. 83, figs. 4, 7, 8, 10, 13-15, 20, pl. 90, fig. 16, not pl. 104, fig. 21, pl. 129, fig. 8, pl. 161, figs. 2, 3, 5, 7, 9, 15-17, 22, pl. 186, figs. 2, 3, 10, 13, 16, pl. 210, fig. 18. Trachyleberis scabrocuneata (Brady, 1880): Hu and Tao, 267, text-fig. 160A, B, pl. 46, figs 17-22, pl. 162, fig. 1. Trachyleberis scabrocuneata (Brady, 1880): Warne, 341-343. Trachyleberis scabrocuneata (Brady, 1880): Tanaka and Nomura, fig. 4.26. Trachyleberis ishizakii Yasuhara et al. 2002: Irizuki et al. fig. 3.13. Trachyleberis ishizakii Yasuhara et al. 2002: Ozawa, fig. 4.20. 2012a 2012b Trachyleberis scabrocuneata, Tanaka et al.: fig. 4R, table 2. Trachyleberis scabrocuneata, Tanaka et al.: fig. 17. Diagnosis Trachyleberis species characterized by spinose carapace, clavate spines, short ocular ridge, absence of primary reticulation, and distinct eye tubercle. Male copulatory limb with a very elongate, heavily sclerotized ventrodistal process. Junior synonym Trachyleberis ishizakii Yasuhara et al. 2002. Material examined Four specimens with soft parts (i.e., collected alive), 19 empty valves (= subfossil specimens), including the lectotype and paralectotypes housed in the Zoology collection of The Natural History Museum, London. Lectotype. Cythere scabrocuneata Brady, 1880; this male specimen was designated lectotype by Harding and SylvesterBradley (1953), who also dissected its dried soft parts. Two slides contain this specimen: 1. One RV, one LV on a MP slide labelled “Trachyleberis scabrocuneata (Brady) ♂, LECTOTYPE, 179.5, ♂, Seto Inland Sea, Chall. Exp., 1952.12.10.1,”. Both valves are illustrated herein in Fig. 4a, b. 2. Fragmented soft parts on a glass slide labelled “179.5, ♂, Trachyleberis scabrocuneata (Brady), LECTOTYPE, Seto Inland Sea, Chall. Exp., 1952.12.10.2”. One of the male copulatory limbs on this slide is illustrated herein in Fig. 5f. Paralectotypes. Cythere scabrocuneata Brady, 1880, one specimen with soft parts and ten empty valves on five micropaleontological and two glass slides: 3. One ♂ RV (slightly broken dorsally and ventrally) on a MP slide labelled “179.5, Trachyleberis scabrocuneata (Brady), RV ♂, figd Bradley 1948, PARATYPE [sic], Challenger Station 233B, 1948.3.10.3. 767”. This specimen was studied by Sylvester-Bradley (1948) and Harding and Sylvester-Bradley (1953). 4. One ♀ RV on a MP slide labelled “179.5, Trachyleberis scabrocuneata (Brady), RV ♀, figd Bradley 1948, PARATYPE [sic], Challenger Station 233B, 1948.3.10.4, 768”. This specimen was studied by Sylvester-Bradley (1948) and Harding and Sylvester-Bradley (1953), it is herein illustrated in Figs. 4c, c’, d, d’ and 5b. 5. Fragmented male soft parts (with copulatory limb) on a glass slide labelled “179.5, ♂, Trachyleberis scabrocuneata (Brady), Seto Inland Sea, Chall. Exp., paratypes [sic], 1952.12.10.8 (pt).A”. This specimen was studied by Harding and Sylvester-Bradley (1953); S. N. Brandão added the “A” at the end of the NHM collection number. Author's personal copy Mar Biodiv (2013) 43:363–405 6. Empty glass slide labelled “NO COVER SLIP, SPECIMEN MISSING, 06/2000, 179.5, ♂, Trachyleberis scabrocuneata (Brady), Seto Inland Sea, Chall. Exp., paratypes [sic], 1952.12.10.8.B”. This slide was studied by Harding and Sylvester-Bradley (1953). S. N. Brandão added the “B” at the end of the NHM collection number. 7. One ♀ LV, one ♂ LV, and one juvenile LV on an original Brady MP slide labelled “179.5, Trachyleberis scabrocuneata Brady, Paratype [sic], 1952.12.10.10.A, “CHALLENGER”, No. 233B, Depth 14 faths., Inland Sea, Japan, G. S. Brady.” These specimens were studied by Harding and Sylvester-Bradley (1953). S. N. Brandão added the “A” at the end of the NHM collection number. Three other LV and one other RV were transferred from this slide to a new slide, now labelled 1952.12.10.10.B (see below). This material is illustrated herein in Figs. 4d, d’, f, f’ and 5a, c, e. 8. One ♂ RV, 1 juvenile RVone broken Von an original Brady MP slide labelled “179.5, Trachyleberis scabrocuneata n. sp., 1952.12.10.11.A, [sic] Paratype, “CHALLENGER”, No. 233B, Depth 14 faths., Inland Sea, Japan, G. S. Brady. 26/5/75, 173”. These specimens were studied by Harding and Sylvester-Bradley (1953). S. N. Brandão added the “A” at the end of the NHM collection number. One other LV was transferred from this slide to a new slide, now labelled 1952.12.10.11.B (see below). Two of these valves are illustrated herein in Figs. 4e, e’, g and 5d, g. 9. One ♂ LV, one broken juvenile LV, one juvenile RVon an original Brady MP slide labelled “179.5, Trachyleberis scabro-cuneata n. sp., 1952.12.10.12.C, Paratype [sic], “CHALLENGER”, No. 233B, Depth 14 faths., Inland Sea, Japan, G. S. Brady. 26/5/75, 173”. These specimens were studied by Harding and Sylvester-Bradley (1953). S. N. Brandão added the “C” at the end of the NHM collection number. Four other LV were transferred from this slide to a new slide labelled 1952.12.10.12.D, and one other broken LV was transferred from this slide to a new slide, now labelled 1952.12.10.12.B (see below). 395 2. Soft parts on a glass slide with dried medium and labelled “Aburatsubo Bay, Kanagawa Prefecture, Inland Sea, Japan, Trachyleberis sp. C, ♂ C, Aburatsubo Bay, SMF Xe 20217”. This specimen is herein illustrated in Fig. 7. 3. One LV, one RV on a MP slide labelled “Trachyleberis sp. C pl. 6 fig. 2, SMF Xe 20218, Aburatsubo Bay, Kanagawa Prefecture, Inland Sea, Japan, Recent”. 4. Soft parts on a glass slide with dried medium and labelled “Aburatsubo Bay, Kanagawa Prefecture, Inland Sea, Japan, Trachyleberis sp. C, ♂ C, Aburatsubo Bay, SMF Xe 20218”. Material not found in the NHM slide and spirit collections. One slide (NHM 80.38.108) was not found in the collection of the NHM. Stratigraphic and geographic distribution (details in Table 2) Pliocene to Recent. Japan and Korea, littoral. Valve measurements (Fig. 8) Lectotype adult male LV L 0.92 mm, H 0.42 mm; RV L 0.91 mm, H 0.40 mm. Paralectotypes. Selected adult males: LV L 0.94 mm, H 0.42 mm; selected adult females: LV L 0.86 mm, H 0.48 mm; RV L 0.88 mm, H 0.47 mm; selected (A-1) juveniles: LV L 0.72-0.74 mm, H 0.38-0.42 mm; RV L 0.73 mm, H 0.39 mm. Two male specimens studied by Jellinek and Swanson (2003) on four slides: Description (Modified after Ikeya and Compton 1983; Yasuhara et al. 2002a) Carapace large. In lateral view, valves subtrapezoidal tapering posteriorly, highest at anterior cardinal angle. Anterior margin rounded; dorsal margin slightly concave; ventral margin broadly convex; posterior margin angulate. Surface ornamented with clavate spines, subcentral tubercle, and ocular ridge. Ocular ridge starting at the eye tubercle and terminating above the midpoint of the anterior margin. Eye tubercle conspicuous, subcentral tubercle inconspicuous. Marginal pores straight, numerous, along anterior and posteroventral margins. Calcified inner lamella moderate in width along anterior and posterior margins. Vestibules absent. Hinge holamphidont. Internal snap-knob structure present at mid-length ventrally. Muscle scars consist of a V-shaped frontal scar and a row of four adductor scars (the second from the top may be subdivided). Sexually dimorphic, males more elongate than females. Male fifth limb with a very strongly developed claw on distal margin of segment I. Male copulatory limb with sub-rounded basal capsule; very elongate and slightly curved, well sclerotized ventro-distal process; delicate, small, poorly sclerotized dorso-distal process; and proximally curved copulatory tube. Furca developed as four well-developed, very barbulate setae. 1. One LV, one RVon a MP slide labelled “Trachyleberis sp. C, pl. 5 fig. 4, pl. 6 fig. 1, SMF Xe 20217, Aburatsubo Bay, Kanagawa Prefecture, Japan, Recent”. Remarks Trachyleberis scabrocuneata s.s. differs from the closely similar T. niitsumai in: (1) having smaller carapace with more sub-triangular and less elongate lateral outline; Additional specimens (not types). Nine subfossil valves studied by M.Y.: Two adult female LV, two adult female RV, one adult male LV, one smaller adult male LV, one adult male RV, one juvenile (A-1) LV, one juvenile (A-1) RVon MP slides, Osaka Bay, Seto Inland Sea, Holocene sediment core OB2, 34º 35' 48" N, 135º 9' 29.1" E, 21.91 m water depth (ZMH K42870). This material is herein illustrated in Fig. 6a-g. Author's personal copy 396 (2) shorter, less regular ocular ridge; (3) lacking the surface reticulation in the adult (but the juvenile (A-1) carapace is weakly reticulate). Jellinek and Swanson’s (2003: pl. 6, figs. 1b, 2b) photographs suggest that the distal process of the copulatory limb of T. scabrocuneata s.s. is asymmetrical— the right one is longer (their pl. 6, Fig. 2b) and more slender than the left (their pl. 6, Fig. 1b). However, Jellinek and Swanson photographed one copulatory limb each of two distinct specimens, so we cannot be sure whether this difference is an asymmetry between right and left sides, or whether it is an inter-individual variation. Paralectotypes of T. scabrocuneata herein reidentified as Trachyleberis niitsumai and Trachyleberis sp. cf. T. niitsumai. For details see the Table 1 and lists of specimens in Trachyleberis niitsumai and Trachyleberis sp. cf. T. niitsumai taxonomic sections): (1) 1 ♂ LV of Trachyleberis niitsumai on the MP slide NHM 1948.3.10.1, 765. (2) 1 ♂ RV of Trachyleberis niitsumai on the MP slide NHM 1948.3.10.2, 766. (3) 1 ♀ RV of Trachyleberis sp. cf. T. niitsumai on the MP slide NHM 1948.3.10.5, 769. (4) 1 RV of Trachyleberis sp. cf. T. niitsumai on the MP slide NHM 1952.12.10.3. (5) 1 LV with 2 holes, 1 RV of Trachyleberis niitsumai on the glass slide NHM 1952.12.10.5.A. (6) very fragmented and dissected soft parts (no copulatory limb) of Trachyleberis niitsumai on the glass slide NHM 1952.12.10.5(pt)B. (7) 3 LV, 1 RV of Trachyleberis niitsumai on the MP slide NHM 1952.12.10.10.B (these valves were originally on slide 1952.12.10.10.A and were transferred to a new MP slide by S.N.B.). (8) 1 LV of Trachyleberis niitsumai on the MP slide NHM 1952.12.10.11.B (these valves were originally on slide 1952.12.10.11.A and were transferred to a new MP slide by S.N.B.). (9) 1 LV of Trachyleberis niitsumai on the MP slide NHM 1952.12.10.12A, 1971. (10) 1 broken LV of Trachyleberis sp. cf. T. niitsumai on the MP slide NHM 1952.12.10.12B, 1971. (11) 4 LV of Trachyleberis niitsumai on the MP slide NHM 1952.12.10.12.D (these valves were originally on slide 1952.12.10.12.C and were transferred to a new MP slide by S.N.B.). (12) 1 RV of Trachyleberis niitsumai on the MP slide NHM 1952.12.10.12.A 4131. (13) 21 V on the MP slide NEWHM 1.43.22. (14) 3 RLV, 19 V on the MP slide NEWHM 1.43.23. (15) several valves on the MP slide NEWHM 2.08.24. Mar Biodiv (2013) 43:363–405 Paralectotypes of Trachyleberis scabrocuneata that cannot be identified positively below genus level because only a few, very fragmented limbs are present This material is included in the collection of the NHM: (1) No cover slip, no soft parts on a glass slide labelled “Paratype [sic], 1952.12.10.4., 179.5, Trachyleberis scabrocuneata Brady, Cotypes. 526.1, Seto Inland Sea., Challenger Exp.”. (2) Very fragmented soft parts (not possible to identify) on a glass slide labelled “179.5, Trachyleberis scabrocuneata (Brady), ♀, Inland Sea, Japan., Chall. Exp., Paratype [sic], 1952.12.10.6.”. (3) Very fragmented soft parts (not possible to identify) on a glass slide labelled “179.5, Trachyleberis scabrocuneata (Brady), ♀, Inland Sea, Japan., Chall. Exp., Paratype [sic], 1952.12.10.7.”. (4) Very fragmented soft parts (not possible to identify) labelled “179.5, Trachyleberis scabrocuneata (Brady), ♀, Inland Sea, Japan., Chall. Exp., Paratype [sic], 1952.12.10.9.”. Other material The following specimens did not belong to the syntype series of Cythere scabrocuneata but where assigned to that species by Puri and Hulings (1976) and designated neotype by Warne and Whatley (1996). As mentioned above, this neotype designation is invalid because the lectotype was not lost. These two valves do not belong to Trachyleberis scabrocuneata, but to another species of Trachyleberis. Additionally, there is a typing mistake in the NHM crustacean collection number cited by Puri and Hulings (1976) and Warne and Whatley (1996) (i.e., 1974.324 instead of 1974.342). However, the correct number 1974.342 is given in the caption of plate 26 of Puri and Hulings (1976). The NHM crustacean collection number 1974.324 refers to Xestoleberis, not to T. scabrocuneata. (1) 1 broken LV, 1 broken V of Trachyleberis sp. cf. T. niitsumai on a MP slide labelled “Cythere scabrocuneata, 1974.342, Topotype, Sediment sample M-298 (Stat. 233B, 15 fms., Inland Sea, Japan), ‘Challenger’, 26.5.1874”. These specimens were sorted from the sediment sample and studied by H. S. Puri. Trachyleberis niitsumai Ishizaki, 1971 (Tables 1 and 2, Figs. 5h, 8 and 9) 1880 in part 1948 in part 1953 in part Cythere scabrocuneata sp. nov.: Brady, 103, pl. 17, figs. 5a-f, pl. 23, figs. 2a-c. Trachyleberis scabrocuneata (Brady, 1880): Sylvester-Bradley, 794-795, pl. 122, figs. 13-18. Trachyleberis scabrocuneata (Brady, 1880): Harding and Sylvester-Bradley, 11-15, ?text- Author's personal copy Mar Biodiv (2013) 43:363–405 1963 in part 1971 ? 1981 ? 1982 ? 1982 ? 1982 1983 in part ?1983 ?1985 1985 1985 1985 non 1987 1988b ?1988a 1988 ?1988 1992 1992 non 1992 non 1994 ?non 1994 1998 1998 figs. 20-24, pls. 1.5, 1.6, 1.8, 2.5, 2.6, 2.9, 2.10, non text-fig. 25. Cythere scabrocuneata Brady, 1880: Bate, 84, Table 3. (a list of Challenger Expedition material). Trachyleberis niitsumai sp. nov.: Ishizaki, 73, 93, Table 1, pl., fig. 5, pl. 4, figs. 15, 18, pls. 5, fig. 3, pl. 6, fig. 10, pl. 7, fig. 9. Trachyleberis niitsumai Ishizaki, 1971: Gou et al. 175, ?pl. 84, figs. 4-6. Trachyleberis niitsumai Ishizaki, 1971: Cai, ?pl. 3, figs. 38, 39. Trachyleberis niitsumai Ishizaki, 1971: Hou et al. p. 225, ?pl. 83, figs. 6-11, ?pl. 88, fig. 20. Trachyleberis niitsumai Ishizaki, 1971: Wang, pl. fig. 30. Trachyleberis scabrocuneata (Brady, 1880): Ikeya and Compton, 119-126, pl. 10, 120, fig. 5, pl. 10, 122, figs. 1-5, pl. 10, 124, figs. 1-5, pl. 10, 126, fig. 5; non pl. 10, 120, figs. 1-4, pl. 10, 126, figs. 1-4 Trachyleberis niitsumai Ishizaki, 1971: Gou et al. ?pl. 15, fig. 29. Acanthocythereis niitsumai (Ishizaki, 1971): Zhao et al. ?pl. 19, fig. 13. Acanthocythereis niitsumai (Ishizaki, 1971): Wang and Zhao, pl. 7, fig. 9. Trachyleberis niitsumai Ishizaki, 1971: Ishizaki and Matoba, pl. 6, figs. 9, 10. Trachyleberis scabrocuneata (Brady, 1880): Ikeya et al. pl. 3, fig. 13. Trachyleberis niitsumai Ishizaki, 1971: Zheng, 201, pl. 6, fig. 5. Acanthocythereis niitsumai (Ishizaki, 1971): Wang, P. et al. pl. 43, figs. 4, 5. Acanthocythereis niitsumai (Ishizaki, 1971): Wang, Q. et al. pl. 1, fig. 4. Acanthocythereis niitsumai (Ishizaki, 1971): Zhao and Wang, pl. 1, fig. 21, 1988 Trachyleberis niitsumai Ishizaki, 1971: Bodergat and Ikeya, 424. Trachyleberis niitsumai Ishizaki, 1971: Lee and Paik, pl. 4, fig. 2. Trachyleberis niitsumai Ishizaki, 1971: Ikeya and Suzuki, pl. 9, fig. 2. Trachyleberis niitsumai Ishizaki, 1971: Huh and Paik, pl. 2, figs. 6, 7. Trachyleberis niitsumai Ishizaki, 1971: Huh, pl.2, fig. 5. Acanthocythereis niitsumai (Ishizaki, 1971): Zheng, Z., pl. 45, figs. 4, 5. Trachyleberis scabrocuneata (Brady, 1880): Yamane, pl. 12, fig. 5. Trachyleberis niitsumai Ishizaki, 1971: Yamane, pl. 12, fig. 4. 397 1998 1998 2001 2001 2002a 2002b ?2003 2003 2005 2006 2006 2008 2009 Trachyleberis sp. 1: Yamane, pl. 12, fig. 6. Trachyleberis niitsumai Ishizaki, 1971: Tanaka et al. pl., fig. 23. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara and Irizuki, pl. 12, figs. 1-6. Trachyleberis scabrocuneata (Brady, 1880): Nakao et al. fig. 11.16. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara et al. fig. 3.22. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara et al. fig. 3.3. Trachyleberis sp. A: Jellinek and Swanson, pl. 5, figs. 1, 2. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara et al. fig. 2.14. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara et al. fig. 4.8. Trachyleberis scabrocuneata (Brady, 1880): Irizuki et al. fig. 7.4. Trachyleberis scabrocuneata (Brady, 1880): Yasuhara and Seto, fig. 5e–f. Trachyleberis niitsumai Ishizaki, 1971: Tanaka, fig. 2.l. Trachyleberis scabrocuneata (Brady, 1880): Tanaka and Nomura, fig. 4.26. Diagnosis Trachyleberis species characterized by spinose carapace, clavate spines, long ocular ridge, and weak primary reticulation. Material examined 1 “live” (= carapaces with soft parts) specimen, 12 subfossil valves. 1. One ♂ LV on a MP slide labelled “179.5, Trachyleberis scabrocuneata (Brady), LV ♂, figd Bradley 1948, [sic] PARATYPE, Challenger Station 233B, 1948.3.10.1, 765”. This specimen was studied by Sylvester-Bradley (1948) and Harding and Sylvester-Bradley (1953, pl. 1, fig. 5, pl. 2, figs. 5, 9). 2. One ♂ RV on a MP slide labelled “179.5, Trachyleberis scabrocuneata (Brady), RV ♂, figd Bradley 1948, [sic] PARATYPE, Challenger Station 233B, 1948.3.10.2, 766”. This specimen was studied by Sylvester-Bradley (1948) and Harding and Sylvester-Bradley (1953, pl. 1, Fig. 6). 3. Two V on a glass slide labelled “179.5, ♂, Trachyleberis scabrocuneata (Brady), Seto Inland Sea, Chall. Exp., [sic] paratypes, 1952.12.10.5.A”. This specimen was studied and dissected by Harding and SylvesterBradley (1953); S. N. Brandão added the “A” in the end of the NHM collection number. 4. Fragmented dissected soft parts (no copulatory limb) on a glass slide labelled “179.5, ♂, Trachyleberis scabrocuneata (Brady), Seto Inland Sea, Chall. Exp., Author's personal copy 398 5. 6. 7. 8. Mar Biodiv (2013) 43:363–405 [sic] paratypes, 1952.12.10.5(pt)B”. This specimen was studied and dissected by Harding and Sylvester-Bradley (1953); S. N. Brandão added the “B” in the end of the NHM collection number. Three LV, one RV on a MP slide labelled “Trachyleberis, Challenger #233b, 14 fathoms, Inland Sea, Japan, SNB-1 060, (these valves were originally on slide 1952.12.10.10.A and were transferred to a new MP slide by S.N.B.), 1952.12.10.10.B”. This specimen was studied by an unknown colleague. S. N. Brandão transferred these specimens from the 1952.12.10.10.A. This material is herein illustrated in Fig. 9a, a’, b, b’. 1 LV on a MP slide labelled “Trachyleberis, Challenger #233b, 14 fathoms, Inland Sea, Japan, SNB-1 061, (these valves were originally on slide 1952.12.10.11.A and were transferred to a new MP slide by S.N.B), 1952.12.10.11.B”. This specimen was studied by an unknown colleague. S. N. Brandão transferred these specimens from the 1952.12.10.11.A. One RV on a MP slide labelled “Trachyleberis scabrocuneata, RV ♂, 4131, [sic] PARALECTOTYPE, “Challenger” Sta. 233B, 1952.12.10.12A, 1971”. This specimen was studied by an unknown colleague. Four LVon a MP slide labelled “Trachyleberis, Challenger #233b, 14 fathoms, Inland Sea, Japan, SNB-1 062, (these valves were originally on slide 1952.12.10.12.C and were transferred to a new MP slide by S.N.B.), 1952.12.10.12.D”. This specimen was studied by an unknown colleague. S. N. Brandão transferred these specimens from the 1952.12.10.12.C. Additional specimens studied by M.Y.. One adult female RV (USNM 560190: TRA1045), one adult male LV (USNM 560189: TRA1044), one (A-1) LV (USNM 560191: TRA1046), one (A-1) RV (USNM 560192: TRA1047) on micropalaeontological slides, Osaka Bay, Seto Inland Sea, modern grab sample OB-40, 34°25.60' N 135°06.83' E, 31 m depth. This material is herein illustrated in Figs. 5h and 9c, c’, d, d’. Stratigraphic and geographic distribution Late Miocene– Recent, Japan and East China Sea, littoral. Valve measurements (Fig. 8). Adult males: LV L 1.10 mm, H 0.52 mm; RV L 1.05-1.10 mm, H 0.47-0.50 mm; adult females: LV L 0.92 mm, H 0.52-0.54 mm; (A-1) juveniles: LV L 0.73-0.83 mm, H 0.41-0.45 mm. Remarks The synonym list provided above is abbreviated, with only important references or references with clear SEM image(s) (see Table 2 for other references). The species considered herein as Trachyleberis niitsumai is the one that was long considered (erroneously) to be Trachyleberis scabrocuneata, and illustrated in the two books and numerous papers (see discussion above). However, our specimens differ slightly from the type specimens of Trachyleberis niitsumai. According to the original illustrations provided by Ishizaki (1971), the specimens from the type locality (Aomori Bay, northeastern Japan) have relatively distinct primary reticulation and less spinose carapaces (Ishizaki 1971), for example, compared with the illustrations by Yasuhara and Seto (2006), Yasuhara and Irizuki (2001), and the present paper (Fig. 9). However, these morphological differences are subtle and Ishizaki’s (1971) specimens look relatively poorly preserved. Thus we considered the less spinose nature and the distinct primary reticulation as a preservation artifacts and/or intraspecific variation. The northern population of T. niitsumai may have more distinct primary reticulation than the southern one, however the northern population is known as yet only from the type locality. Trachyleberis sp. cf. T. niitsumai Ishizaki, 1971 (Tables 1 and 2) Material examined Five subfossil valves. 1. One RV on a MP slide labelled “Trachyleberis scabrocuneata (Brady), ♀, [sic] PARATYPE, 179.5, RV and fragments, Inland Sea, Japan, 1952.12.10.3, ”. This specimen was studied by Harding and SylvesterBradley (1953). 2. One ♀ RV on a MP slide labelled “179.5, Trachyleberis scabrocuneata (Brady), RV ♀, figd. Bradley 1948, [sic] PARATYPE, Challenger Station 233B, 1948.3.10.5, 769”. This specimen was studied by Sylvester-Bradley (1948) and Harding and Sylvester-Bradley (1953, Pls 1.8, 2.6, 2.10). 3. One broken LV on a MP slide labelled “Trachyleberis scabrocuneata (Brady), 1880, LV ♂, specimen photographed on SEM at Leicester, 4155, “Challenger” Sta. 233B, 1952.12.10.12B, 1971”. This specimen was studied by a unknown author. 4. One broken LV, one broken V on a MP slide labelled “Cythere scabrocuneata, 1974.342, Topotype, Sediment sample M-298 (Stat. 233B, 15 fms., Inland Sea, Japan), ‘Challenger,’ 26.5.1874”. These specimens were sorted from the sediment sample and studied by H. S. Puri. Remarks This section has been included mostly for the sake of completeness, as it contains paralectotypes of T. scabrocuneata herein reidentified as Trachyleberis sp. cf. T. niitsumai. “Trachyleberis” lytteltonensis Harding and SylvesterBradley, 1953 Author's personal copy Mar Biodiv (2013) 43:363–405 (Tables 1 and 2, Figs. 5i, 8 and 10) ?1880 in part 1898 ? 1952 1953 2003 Cythere scabrocuneata sp. nov.: Brady, 103. non pI. 17, figs. 5a-f, non pl. 23, fig. 2a-c. Trachyleberis scabrocuneata (Brady, 1880): Brady, 444-445, pl. 47, figs. 1-7, 18-25. Trachyleberis scabrocuneata (Brady, 1880): Hornibrook, 32-33, pl. 3, figs. 38, 39, 48. Trachyleberis lytteltonensis sp. nov. Harding and Sylvester-Bradley, p. 4, text-figs. 2–19; pl. 1, figs. 1–4, 7; pl. 2, figs. 1–4, 7–8. Trachyleberis scabrocuneata Brady, 1898 [sic]: Jellinek and Swanson, 20-23, pls. 3, 4. 399 Paratypes: 2. One male LV, one male RV; this valve is herein illustrated in Figs. 5i and 10c, d 3. One male LV, 1 male RV both coated 4. Two (?adult) female LV, one female RV 5. One juvenile RV 6. One juvenile RV. This valve is herein illustrated in Fig. 10e. 7. One juvenile LV, one broken juvenile RV. This valve is herein illustrated in Fig. 10f 8. One male LV, one male RV NHM, second to fifth slides with paratypes: Diagnosis In lateral view, male valves sub-rectangular, female valves sub-trapezoidal. Lateral pore canals bordered by irregularly shaped, robust tubercles, except for the antero-dorsal region, which is smooth. Postero-dorsal margin with a series of approximately five long spines, plus a few leaf-shaped spines more laterally. Anterior margin with approximately 15 leaf-shaped spines. Anteriorly and posteriorly, just lateral to leaf-shaped spines, are several shorter and more delicate spines. Valve surface with thin, delicate reticulation, bordered by very thin and delicate ribs, which are denser ventrally, weak dorsally and absent anteriorly. Male copulatory limb roughly subtriangular, with rounded basal capsule and irregularly shaped distal lobe. Material examined Two hundred twenty “live” (= carapaces with soft parts) specimens, 29 subfossil valves. Type material. The holotype and paratypes are in The Natural History Museum: 12 specimens collected alive, plus 2 subfossil valves. The holotype and some of the paratypes are divided between one MP slide (valves or carapaces) and alcohol-filled glass vials (soft parts); some paratype soft parts are dissected on glass slides. Additional paratypes are in the G.S. Brady Collection (Great North Museum). NHM, first slide. One paper, MP slide with a total of 6 LV, 7 RV (plus 1 broken RV) with a rectangular surface subdivided in 24 quadrants, each quadrate numbered from 1 to 24. This last slide is labelled “Trachyleberis lytteltonensis Harding and Sylvester-Bradley, 179.5, Lyttelton Harbour, New Zealand, 1952.12.9.1-6(pt.)7,8, holotypes+paratypes”. Only the following cells contain specimens: Holotype: 1. One male LV, one male RV. The soft parts of this holotype are in the glass vial NHM 1952.12.9.1 (see below). This specimen is herein illustrated in Fig. 10a, b. 1. Female soft parts on a glass slide labelled “179.5, Trachyleberis lytteltonensis H. and S-B, Lyttelton Harbour, New Zealand, Paratype, female, 1952.12.9.5.pt.” 2. Male soft parts on a glass slide with many air bubbles in the medium, labelled “49018, 179.5, Trachyleberis lytteltonensis H. and S-B, Lyttelton Harbour, N. Z., male, Paratype, from 1952.12.9.10.pt.” 3. Male soft parts on a glass slide labelled “49058, 179.5, Trachyleberis lytteltonensis H. and S-B, male, Lyttelton Harbour, New Zealand, Paratype, 1952.12.9.11.” 4. Female soft parts on a glass slide labelled “49060, 179.5, Trachyleberis lytteltonensis H. and S-B, female, Lyttelton Harbour, New Zealand, Paratype, 1952.12.9.12.” NHM, Jar with holotype and paratypes. this glass jar (from the spirit collection of the NHM) is filled with alcohol and is labelled “179.5, Trachyleberis lytteltonensis Harding and Sylvester-Bradley, Lyttelton Harbour, New Zealand, 1952.12.9.1, holotype, 1952.12.9.2-6, paratypes”. This larger jar contains eight smaller glass vials, each one containing one even smaller vial inside. Each of these small vials contains a label and a specimen, as follows: Holotype: First small glass vial—soft parts of one specimen, label “1952.12.9.1, Holotype, Trachyleberis lytteltonensis H. and S-B.” Paratypes*: Second small glass vial—soft parts of one specimen, label “1952.12.9.2” Third small glass vial—soft parts of one specimen, label “1952.12.9.3” Fourth small glass vial—fragmented soft parts, label “1952.12.9.4” Fifth small glass vial—fragmented soft parts, label “1952.12.9.5” Sixth small glass vial—fragmented soft parts, label “1952.12.9.6” Author's personal copy 400 Seventh small glass vial—two “live” (= carapaces with soft parts) females, two “live” (= carapaces with soft parts) males, label “1952.12.9.9” Eighth small glass vial—two decalcified valves, label “1952.12.9.10” According to the ICZN’s Article 72.4.6, because Harding and Sylvester-Bradley (1953) did not explicitly exclude any specimen from the type series of T. lytteltonensis, all specimens they studied are paratypes (except the holotype). This includes material they referred to as “40 specimens preserved in spirit from the Brady collection, Hancock Museum, Newcastle-on-Tyne”, from which they selected the holotype and paratypes now kept at the NHM, although the number of specimens mentioned seems to be an underestimated, since a recent check (DJH: March 2012) on the G.S. Brady Collection in Newcastle (now housed in the Resources Centre at the Discovery Museum) confirmed the presence of over 50 specimens. Of these, nine specimens are on a plastic MP slide, NEWHM 1.54.24, labelled “Trachyleberis lytteltonensis paratypes, Lyttelton Harbour, New Zealand. Removed from Brady Spirit Collection by D. Horne, February 1985. c. 50 specimens remaining in the spirit jar)”. This slide contains three male and three female carapaces, plus one male RV, one female RV and one female LV. The specimens remaining in the spirit jar appear to be mostly adult males and females; material in the Brady Spirit Collection does not currently have catalogue numbers, although work is in hand to rectify this. Additional material. From the collection of the Zoologisk Museum, København (Zoological Museum, Copenhagen): 208 “live” (= carapaces with soft parts) specimens, plus 27 subfossil valves (see details below). Material studied by Jellinek and Swanson (2003). The following specimens contained either in slides or in glasses were studied and identified as “Trachyleberis scabrocuneata (Brady 1898)” [sic] by Jellinek and Swanson (2003). Although Jellinek and Swanson (2003) labelled some of the specimens below as “non Tr. lytteltonensis Harding and Sylvester-Bradley 1953”, these specimens do belong to “T.” lytteltonensis. Fragments of valves on a black, plastic micropaleontological slide labelled “Trachyleberis scabrocuneata (BRADY 1898), empty valves, Lyttelton Harbour, New Zealand, 1-5 fathoms, Recent, ZMUC-CRU-3630”. Sixteen “live” (= carapaces with soft parts) (= with soft parts) females, 41 “live” (= carapaces with soft parts) (= with soft parts) males, 2 female LV with soft parts, 2 male LV with soft parts, 2* male RV without soft parts, 1* male RLV (= closed carapace without soft parts) with a rounded hole on a glass with alcohol with two labels: (1) “ZOOLOGISK MUSEUM, KØBENHAVN. Trachyleberis lytteltonensis Hard., Mar Biodiv (2013) 43:363–405 Lyttelton Harbour, New Zealand, 2-10 m, Datm: 1.Sept.1897, Legit: Henry Suter”; (2) “CRU-3630”. *Both male RV and the holed RLV from the glass above were transferred to a new black, plastic micropaleontological slide labelled “These valves were on alcohol together with other specimens, Trachyleberis lytteltonensis, Lyttelton Harbour, New Zealand,, ZMUC-CRU-3630”. One female LV, plus fragments of valve (s) on a black, plastic micropaleontological slide labelled “Trachyleberis scabrocuneata (BRADY 1898), non Tr. lytteltonensis Harding and Sylvester-Bradley 1953, female, Lyttelton Harbour, New Zealand, 1-5 fathoms, Recent, ZMUC-CRU3631” One male RV on a black, plastic micropaleontological slide labelled “Trachyleberis scabrocuneata (BRADY 1898), non Tr. lytteltonensis Harding and SylvesterBradley 1953, male, Lyttelton Harbour, New Zealand, 1-5 fathoms, Recent, ZMUC-CRU-3631” Three female LV, two female RV, four male LV, five male RV, one juvenile LV, one juvenile RV (plus the empty valves described below) on a black, plastic micropaleontological slide labelled “Trachyleberis scabrocuneata (BRADY 1898), empty valves, Lyttelton Harbour, New Zealand, 1-5 fathoms, Recent, ZMUC-CRU-3631”. Seventy-two “live” (= carapaces with soft parts) (= with soft parts) females, 63 “live” (= carapaces with soft parts) (= with soft parts) males, 5 female LV with soft parts, 1 female RV with soft parts, 1 male RV with soft parts, 3 “live” (= carapaces with soft parts) (= with soft parts) juveniles, 5* female LV without soft parts, 1* female RV without soft parts, 1* male RV without soft parts on a glass with alcohol with four labels: (1) Brady’s original label “Cythereis (Cyther) scabrocuneata Brady=? C. dorsoserrata Brady”; (2) with a distinct hand writing “Trachyleberis lytteltonnensis Harding and Sylvester-Bradley 1953, Topotypes”; (3) “Cythere scabrocuneata Brady, Lyttelton Harb.” (plus a couple of illegible words); (4) “CRU-3631”. *All empty valves were transferred to a new black, plastic micropaleontological slide described above. Stratigraphic and geographic distribution Only known from the type locality: Recent, Lyttelton Harbour, New Zealand, 2–9 m depth. Valve measurements (Fig. 8) All “live” (= carapaces with soft parts) specimens (ZMUC-CRU-3060): Adult males: LV L 1.04-1.11 mm, H 0.50-0.53 mm; adult females: LV L 0.85-0.92 mm, H 0.48-0.53 mm; (A-1) juveniles: LV L 0.78-0.81 mm, H 0.41-0.46 mm. Remarks. As discussed above, because of its distinctive carapace and male copulatory appendage morphology, “Trachyleberis” lytteltonensis Harding and SylvesterBradley, 1953 cannot be assigned to Trachyleberis and Author's personal copy Mar Biodiv (2013) 43:363–405 deserves a new genus, which is the subject of a new description by Yasuhara et al. (in preparation). Genus Pennyella Neale, 1974 Pennyella sp. (Fig. 11) 1889 in part Cythere scabrocuneata: Brady and Norman, 154-155, non pI. 47, figs. 1-2, ?pl. 47, figs. 3-7, 18-25. Material examined Twenty valves. Four RV, two LV, two V on a paper MP slide of Norman’s collection (NHM 1900.3.6.237) labelled “Trachyleberis scabrocuneata, 1900-3-6-237, Porcupine 1869, Stat 19, Lat 54°53 N, Long 10° 56 W, 1360 fath”. These specimens were studied by Brady and Norman (1889), and are herein illustrated in Fig. 11. Twelve V on a paper MP slide of Norman’s collection (NHM 1911.11.8.M3310) labelled “Brady, Trachyleberis sp., Lat. 59°11’N, Long 37. 41. W, 1450 fath, Valorous 1875, 1911.11.8.M3 310”. These specimens were studied by Brady and Norman (1889), and are herein illustrated in Fig. 11. Remarks The specimens included in the present section were collected from the North Atlantic and were studied by Norman (for the publication of 1889) and misidentified as Cythere scabrocuneata. Acknowledgements Miranda Lowe (NHM, London) carefully assisted the first author during five visits to the NHM and also kindly provided access to and permission for studying the Challenger and other ostracod specimens housed in the NHM. Dan Gordon (Discovery Museum) kindly facilitated the re-examination of the Brady collection material by DJH. Mark J. Grygier (Lake Biwa Museum) and Philippe Bouchet (MNHM) helped with correct interpretations of articles of the ICZN. Jørgen Olesen and Tom Schiøtte (Natural History Museum of Denmark) loaned the specimens of Trachyleberis lytteltonensis. Alan Lord loaned the specimens from the Senckenberg Research Institute and Natural History Museum Frankfurt. Gene Hunt and Carlita Sanford assisted depositing specimens in the Smithsonian collection. Stephen Eager (Victoria University of Wellington), Katsura Yamada (Shinshu University), John Neil (La Trobe University), Robin J. Smith (Lake Biwa Museum), Hayato Tanaka (Shizuoka University), Mark Warne (Deakin University) kindly provided valuable information and/ or papers on Trachyleberis. The suggestions of the editor Pedro Martínez Arbizú and four anonymous referees greatly improved the present publication. The first author thanks Dietmar Keyser (ZMH, Universität Hamburg), Angelika Brandt (ZMH, Universität Hamburg) and Pedro Martínez Arbizú (DZMB, Senckenberg Institute) for their mentorship. M.Y. thanks Gene Hunt (Smithsonian Institution) for discussion in the initial stage of this project, and Scott Whittaker (Smithsonian Institution) for help in SEM imaging. S.N.B. is/was financially supported by Alexander von Humboldt Foundation, Encyclopedia of Life and Hansische Universitäts-Stiftung. 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