Memoirs of Museum Victoria ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs/index.asp MELBOURNE AUSTRALIA Memoir 65 31 December, 2008 Chief Executive Officer J. Patrick Greene Director (Collections, Research and Exhibitions) Robin Hirst Scientific Editor Richard Marchant Editorial Committee Martin F. Gomon David J. Holloway Gary C. B. Poore Kenneth Walker Robin S. Wilson Mi MUSEUMVICTORIA Published by Order of the Museums Board Of Victoria © The Museums Board of Victoria 2008 Typeset by the printer's drawers Printed by the BPA Print Group Memoirs of Museum Victoria SSN 1447-2554 (Online) and ISSN 1447-2546 (Print) Frequency: 1 issue per year Museum Victoria undertakes research in order to contribute to a deeper understanding and appreciation of the origins, diversity and development of Australia's natural, cultural and scientific heritage, and applies this knowledge for the benefit of society. 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A period and dash must separate taxon and author except in the case of reference to the original description, e.g. Leontocaris Stebbing, 1905: 98—99. — Barnard, 1950: 699. Supplementary information (extended lists of material examined, databases etc) should be submitted separately and with the final manuscript to be forwarded to referees. The Editorial Board encourages use of supplementary information to minimise the cost of printing as long as the requirements of the International Code of Zoological Nomenclature are met in the printed paper. Contents Volume 65 2008 1 > Australian species of Psolidium Ludwig (Echinodermata: Holothuroidea: Psolidae) P. Mark O’Loughlin and Dragica Marie 23 > Antarctic and Sub-Antarctic species of Psolidium Ludwig (Echinodermata: Holothuroidea: Psolidae) P. Mark O’Loughlin and Cynthia Ahearn 43 > Marmenuera, a new genus of leptophlebiid mayfly (Insecta: Ephemeroptera) endemic to Tasmania J.C. Dean, G.N.R. Forteath & A.W. Osborn 51 > A new species in the Ophiocoma erinaceus complex from the South-west Pacific Ocean (Echinodermata: Ophiuroidea: Ophiocomidae) Milena Benavides-Serrato and Timothy D. O'Hara 57 > A new species in the genus Ophiomyxa from South-west Australian waters (Echinodermata: Ophiuroidea: Ophiomyxidae) Amanda M. Franklin and Timothy D. O’Hara 63 > Deep-sea majoid crabs of the genera Oxypleurodon and Rochinia (Crustacea: Decapoda: Brachyura: Epialtidae) mostly from the continental margin of Western Australia Bertrand Richer de Forges and Gary C.B. Poore 71 > The Psocoptera (Insecta) of Tasmania, Australia E. R. Schmidt and T. R. New 153 > Distribution patterns and diversity of invertebrates of temperate rainforests in Tasmania with a focus on Pauropoda Penelope Greenslade 1 65 > Hydroids of the BANZARE expeditions, 1929-1 931 : the family Haleciidae (Hydrozoa, Leptothecata) from the Australian Antarctic Territory Jeanette E. Watson 179 > Lobataria Newtoni gen. et sp. nov., (Hydrozoa, Anthoathecata, Tubulariidae) from southern Australia Jeanette E. Watson 185 > Corymorpha Rubicincta, a new Hydroid (Hydrozoa, Anthoathecata, Corymorphidae) from Port Phillip, Australia Jeanette E. Watson 189 > A New Species of the Roughy Genus Hoplostethus (Trachichthyidae) off North-western Australia Martin F. Gomon Memoirs of Museum Victoria 65: 1-22 (2008) ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs/index.asp Australian species of Psolidium Ludwig (Echinodermata: Holothuroidea: Psolidae) P. Mark O’Loughlin 1 and Dragica Maric 2 1 Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne 3001, Victoria, Australia (pmo@bigpond.net.au) 2 Research Associate, Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne 3001, Victoria, Australia (dragicamaric@hotmail.com) Abstract O'Loughlin, PM. and Maric, D. 2008. Australian species of Psolidium Ludwig (Echinodermata: Holothuroidea: Psolidae). Memoirs of Museum Victoria 65: 1-22. Six new species of Psolidium Ludwig from Australia are described: Psolidium berentsae, P. hutchingsae, P. karenae, P. laperousazi , P. marshae, P. mccallumae. The species Psolus parmatus Sluiter and Psolus spinuliferus H. L. Clark are re-assigned to Psolidium. Diagnoses are given for Psolidium granuliferum H. L. Clark, P. nigrescens H. L. Clark, P. minutus (H. L. Clark), P. parmatus (Sluiter), P. ravum Hickman and P. spinuliferus (H. L. Clark). The family Psolidae is ascribed to Forbes as author, not Perrier. The diagnosis of the family Psolidae, assignment of genera, and status of the family are discussed. The genus Psolidium is revised. Keys are provided to the genera of Psolidae, and 12 known Australian species of Psolidium. Keywords Echinodermata, Holothuroidea, Psolidae, Psolidium, taxonomy, new species, keys. Introduction Rowe (in Rowe and Gates, 1995) listed four species of Psolidium Ludwig, 1886 from Australia: P. granuliferum H. L. Clark, 1938 (southern Western Australia), P. minutus (H. L. Clark, 1938) (Lord Howe I), P. nigrescens H. L. Clark, 1938 (New South Wales), and P. ravum Hickman, 1962 (Tasmania). Psolus spinuliferus H. L. Clark, 1938 was described for northwestern Australia, and is re-assigned here to Psolidium. Psolus parmatus (Sluiter, 1901) was described for Indonesia, and was found recently in collections from the continental slope off Western Australia. It is also re-assigned here to Psolidium. In this work six new species of Psolidium are described for Australia. Pawson and Fell (1965), and subsequent authors, have incorrectly nominated Perrier (1902) as the systematic author of the Psolidae. Forbes (1841) was the original author. Materials and methods Live colour photographs of some species were taken in the field by Karen Gowlett-Holmes, and the images lodged in the South Australia Museum with a photoindex registration prefix PK. Some specimen photographs were taken by Leon Altoff and Audrey Falconer, with Mark O’Loughlin, using a Pentax K10D digital SLR with a variety of lenses and lit using two electronic flashes. Preserved specimens were placed on or suspended above black velvet. The live photo was taken in a perspex tank with filtered sea water. Photos of ossicles were taken by Mark O’Loughlin and Chris Rowley using a Leica DM5000 B compound microscope, and Leica DC500 camera with montage software. Some specimen photographs were taken by Simon Hinkley with Dragica Maric using a Leica MZ16 stereomicroscope, Leica DC500 digital camera with montage software. Drawings of tube feet distributions were made by Dragica Maric using stereomicroscope and camera lucida. Abbreviations: AM— Australian Museum, Sydney; MCZ— Museum of Comparative Zoology, Harvard, USA; NMV— Museum Victoria, Melbourne, Australia; SAM— South Australian Museum, Adelaide; WAM— Western Australian Museum, Perth; ZMA— Zoologisch Museum, Amsterdam. Specimen registration number prefixes: AM J, NMV F, SAM K, WAM Z, ZMA V.ECH.H Dendrochirotida Grube, 1840 (restricted Pawson and Fell, 1965) Key to genera of Psolidae Forbes 1. Dorsal and lateral scales imbricating, conspicuous, lacking calcareous towers; scales of ventro -lateral body clearly demarcated from the thin calcareous sole that lacks scales 2 2 P. Mark O’Loughlin and Dragica Marie — Dorsal and lateral multilayered ossicles (scales) embedded in integument; some scales with conspicuous projecting calcareous towers; ventro -lateral body not clearly demarcated from sole Echinopsolus Gutt, 1990 2. Tentacles 15; oral valves 5, radially situated Ceto Gistel, 1848 — Tentacles 10; oral valves situated interradially if present ... 3 3. Mid-body tube feet absent dorsally and laterally Psolus Oken, 1815 — Mid-body tube feet present dorsally and laterally 4 4. Dorsal and lateral scales covered by ossicles that include hour-glass shaped and/or tower ossicles Lissothuria Verrill, 1867 — Hour-glass shaped and tower ossicles not present amongst the dorsal and lateral ossicles 5 5. Mouth and anus lie in the plane of bilateral symmetry Psolidium Ludwig, 1886 — Mouth and anus lie in a plane at right angle to the usual plane of bilateral symmetry Ekkentropelma Pawson 1971b Psolidae Forbes (synonymy of systematic records) Psolidae Forbes, 1841: 201-02,206.— Agassiz, 1845: 11.— Agassiz, 1848: 905. “Psoline sub-family” Bell, 1882: 642 (no family or other sub- family nominated). Psolida (uncredited).— Haeckel, 1896: 380, 441, 442. Psolinae R. Perrier, 1902: 493, 512 (sub-family of Cucumariidae, with Cucumariinae). Psolidae Perrier, 1902.— Pawson and Fell, 1965: 4.— Pawson, 1969a: 129.— Pawson, 1968b: 19.— Pawson, 1968c: 347.— Tommasi, 1969: 8.— Pawson, 1970: 28.— Pawson, 1971a: 33-34.— Pawson, 1971b: 115, 118.— Tommasi, 1971: 3-4.— Pawson and Valentine, 1981: 450.— Carriol et Feral, 1985: 50,-Gutt, 1988: 22-23.-Gutt, 1990: 112-13.— Massin, 1992a: 317.— Massin, 1992b: 179.— Lambert, 1996: 21.— Massin, 1997: 101. Psolidae Ed. Perrier (undated).— Thandar, 2006: 35 (R. Perrier was the author of Psolinae). Psolidae (uncredited).— Mortensen, 1927: 413.— Deichmann, 1940: 206. -Deichmann, 1941: 73, 135-36.-H. L. Clark, 1946: 385, 412-13.— Deichmann, 1947: 336: 336.— Deichmann, 1954: 401.— Hickman, 1962: 60.— Pawson, 1964: 461-62.— Pawson, 1967: 1-2.— Baranova and Belyaev, 1968: 236.— Pawson, 1968a: 142.— Pawson, 1969b: 38, maps 3, 5.— Cherbonnier, 1974: 601, 605.— Dartnall, 1980: 13, 77.— Pawson, 1982: 815.— Cannon and Silver, 1987: 10, 11, 29.— Rowe (in Rowe and Gates), 1995: 317. Diagnosis ( most recently by Lambert, 1996, quoting Pawson, 1970). Body flattened, with well-defined ventral sole. Dorsal surface of body invested by imbricating scales. Ventral sole soft, surrounded by tube feet. Mouth and anus dorsally turned. Type genus. Psolus Oken, 1815 (original designation; = Lepidopsolus Bronn, 1860, and Lophothuria Verrill, 1866; synonymy by Theel, 1886). Other genera. Ceto Gistel, 1848 (= Cuvieria Jager, 1833, Callisto Gistel, 1848, Stolinus Selenka, 1868, Hypopsolus Bell, 1882, and Theelia Ludwig, 1892; synonymy by Pawson, 1971a); Lissothuria Verrill, 1867 (= Thyonepsolus H. L. Clark, 1901; synonymy by Pawson, 1967); Psolidium Ludwig, 1886; Ekkentropelma Pawson, 1971b; Echinopsolus Gutt, 1990. Remarks. The family Psolidae was erected by Forbes (1841), who based his family on the genus Psolus Oken and remarked that Cuvieria Peron (= Ceto Gistel, by Pawson, 1971a) should be included in Psolidae. The family was recognised by Agassiz (1845, 1848). Bell (1882) referred to a “Psoline sub-family”, without reference to family or additional sub-family. Perrier (1902) erected 2 sub-families for Cucumariidae: Cucumariinae and Psolinae. Subsequently no author (including Perrier himself) has referred to Psolinae Perrier, 1902. Perrier (1905), Mitsukuri (1912), and Ohshima (1915) referred species of Psolidium Ludwig, 1886, and Psolus Oken, 1815, to Cucumariidae, without reference to Psolidae. Mortensen (1927) referred Psolus and Psolidium species to Psolidae, without indication of family author. Many authors followed Mortensen (1927). Pawson and Fell (1965) incorrectly nominated Perrier (1902) as the systematic author of the Psolidae. Subsequent authors incorrectly referred to Psolidae Perrier, 1902. Within the history of holothuroid classification we note that Semper (1868) referred Psolus Oken to order II Pneumonophora, family Dendrochirotae, sub-family Dendrochirota Gastropoda. Theel (1886) referred Psolus Oken to order II Pedata, family Dendrochirotae, sub-family Gastropoda. Forbes (1841) distinguished the family Psolidae as “having a soft circumscribed disk like the foot of a Gasteropodous Mollusc on which the suckers are placed for progression”. In his diagnosis of sub-family Psolinae, Perrier (1902) continued emphasis on the distinct sole with its specialised tube feet. Mortensen (1927) added the presence of large imbricating scales dorsally, sharply delimited from the thin-walled ventral sole; dorsal mouth and anus; and 10-15 tentacles. Subsequent diagnoses by Deichmann (1941), Hickman (1962), Pawson and Fell (1965), Tommasi (1969, 1971), Pawson (1970, 1982), Carriol et Feral (1985), Cannon and Silver (1987), Gutt (1988), Rowe (in Rowe and Gates, 1995), and Lambert (1996) have generally agreed with Mortensen (1927). But none has continued to include the significant point made by Mortensen (1927) that there is a sharp demarcation between the dorsal scales and the thin-walled sole. In discussing his new genus Psolidiella, Mortensen (1925) noted “a distinct ventral sole, which is, however, not limited from the rest of the body by a sharp edge”, one reason given by Mortensen for not referring Psolidiella to the Psolidae. A second reason stated by Mortensen (1925) for not referring Psolidiella to the Psolidae was “the fact that the posterior part of the intestine, with its mesentery, is in the left ventral interradius”. He added “the situation of the posterior part of the intestine appears to be of primary importance for the subdivision of dendrochirotes, the cucumariids and the phyllophorids having it in the left, and the psolids having it in the right ventral interradius”. Australian species of Psolidium Ludwig 3 Hickman (1962) noted contradictory observations by Deichmann (1941) who claimed that “Cucumariidae seem typically to have the third mesentery attached on the right side of the midventral muscle band, the Phyllophoridae seem to have it attached to the left, and the Psolidae have it either way”. She explained that for Psolidium and Thyonepsolus (- Lissothuria ) the loop is attached in the left ventral interambulacrum, while in Psolus it is attached in the right. Some cucumariid species were examined in this study: Apsolidium densum O’Loughlin and O’Hara, 1992, Neoamphicyclus mutans (Joshua, 1914), Psolidiella hickmani O’Loughlin, 2000. In these three cucumariid genera and species the posterior intestinal mesentery is attached to the left of the midventral radial muscle, supporting Mortensen (1925) and contradicting Deichmann (1941). O’Loughlin (2000) illustrated this position for the genus Psolidiella. Pawson (1968a) described aright attachment for the cucumariid species Pseudopsolus macquariensis (Dendy, 1896), and Ludwig and Heding (1935) reported a right attachment for their cucumariid species Pseudocholchirus mollis , supporting Deichmann (1941). This evidence indicates that the position of posterior intestinal mesenteric attachment is variable for cucumariids as currently assigned. Some psolid species were examined in this study: Ceto cuvieria (Gistel, 1848), Echinopsolus acanthocola Gutt, 1990, Psolidium poriferum (Studer, 1876) (- incertum ), Psolidium ravum Hickman, 1962, Psolus antarcticus Philippi, 1857, Psolus arnaudi Cherbonnier, 1974, Psolus charcoti Vaney, 1906, Psolus koehleri Vaney, 1914, Psolus paradubiosus Carriol and Feral, 1985. In eight of these psolid species the posterior intestinal mesentery is attached to the right of the midventral radial muscle, supporting Mortensen (1925) and Deichmann (1941) for Psolus species. But in Ceto cuvieria it is attached to the left. This evidence indicates that the position of posterior intestinal mesenteric attachment is variable for psolids as currently assigned. Pawson (1967) noted difficulty with the Psolidae in determining whether or not some species should be considered psolids or referred to another dendrochirotid family. The cucumariid genera Pseudopsolus (see Pawson, 1968a), Apsolidium and Neocnus (see O’Loughlin and O’Hara, 1992), and Psolidiella (see O’Loughlin, 2000) include species with a sole that is not delimited by a distinct junction of ventro -lateral body wall scales with a thin-walled sole lacking scales, and lack conspicuous imbricating dorsal and lateral scales. None has been referred to Psolidae. We support this exclusion. The genus Echinopsolus Gutt, 1990 was referred to Psolidae on the grounds of the species having a distinct sole. We note that Psolus charcoti Vaney, 1906 and Echinopsolus acanthocola Gutt, 1990 lack a sharply demarcated sole, and lack macroscopic imbricating scales dorsally, and should not be referred to Psolidae. Reassignment of these taxa does not belong in this revision of Psolidium , and will be treated elsewhere. A comprehensive revision of the relationships amongst cucumariid and psolid species is required and should be undertaken with supportive evidence from molecular genetic data. Table 1. Distribution of Australian species of Psolidium Ludwig, 1886. Psolidium species Distribution P. berentsae sp. nov. Queensland, Lizard I, 6-18 m P. granuliferum H. L. Clark, 1938 SW Western Australia to SE Tasmania, 4-37 m P. hutching sae sp. nov. Northern New South Wales, 12-15 m P. karenae sp. nov. South Australia, 3-12 m P. laperousazi sp. nov. SE Tasmania to South Australia, 1-10 m P. marshae sp. nov. Western Australia to South Australia, 5-14 m P. mccallumae sp. nov. Western Australia, off Point Cloates, 100 m P. minutus (H. L. Clark, 1938) Eastern Tasman Sea, 1-10 m P. nigrescens H. L. Clark, 1938 NSW, Broken Bay to Batemans Bay, 0-11 m P. parmatus (Sluiter, 1901) Indonesia to NW Western Australia, 95-487 m P. ravum Hickman, 1962 SE Tasmania to W South Australia, 0-15 m P. spinuliferus (H. L. Clark, 1938) NW Australia, Darwin to Perth, 0-22 m 4 P. Mark O’Loughlin and Dragica Marie Key to Australian species of Psolidium Ludwig 1. Dorsal and lateral scales with vertical spires/pillars (not lumps) 2 — Dorsal and lateral scales lacking spires/pillars; some species with surface lumps on the scales 3 2. Dorsal and lateral scales each covered with slightly bulbous pillars; lacking mid-ventral radial series of tube feet; “thorn” ossicles present in body wall P. parmatus (Sluiter, 1901) (NW Australia slope) — Dorsal and lateral scales with predominantly single digitiform spires; mid-ventral radial series of tube feet present; lacking “thorn” ossicles in body wall P. spinuliferus (H. L. Clark, 1938) (N and W Australia) 3. “Thom” ossicles present in body wall; live and preserved colour “black” ,.P. nigrescens H. L. Clark, 1938 (central New South Wales) — Body wall lacking “thorn” ossicles; live and preserved colour not “black” 4 4. Body wall with cupped crosses and/or cups 5 — Body wall lacking cupped crosses and/or cups 11 5. Cupped crosses and/or cups in sole of 2 ranges of size ... 6 — Cupped crosses and/or cups in sole of 1 range of size 7 6. Rosettes present in dorsal body wall and tentacles; smaller cupped crosses and cups up to 32 pm long; sole with irregular thick perforated plates, knobbed on surface and margin P. marshae sp. nov. (SW Australia) — Rosettes absent from dorsal body wall and tentacles; smaller cupped crosses and cups up to 24 pm long; sole with smooth perforated plates P. minutus (H. L. Clark, 1938) (E Tasman Sea) 7. Dorsal and lateral cups shallow, completely covered by fine spinelets, including the cross; lacking mid-ventral radial series of tube feet P. mccallumae sp. nov. (Western Australia slope) — Dorsal and lateral cups and cupped crosses not shallow; spinelets on rim of cup or distally on branches of cupped cross, not on cross; mid-ventral radial series of tube feet present 8 8. Body wall with predominantly cups, fewer cupped crosses 9 — Body wall with predominantly cupped crosses, fewer cups 10 9. Knobbed plates in sole; tentacle rosettes small, up to 40 pm long; body rounded ventrally in transverse section P. berentsae sp. nov. (NE Queensland) — Perforated plates in sole predominantly smooth; tentacle rosettes large, up to 80 pm long; body flat ventrally P. laperousazi sp. nov. (SE Australia) 10. Knobbed perforated plates in sole; irregular perforated plates in tentacles P. hutchingsae sp. nov. (N New South Wales) — Perforated plates in sole predominantly smooth; tentacles lack irregular perforated plates P. ravum Hickman, 1962 (SE Australia) 11. Lacking series of mid-ventral radial tube feet; dorsal and lateral tube feet inconspicuous in mid-body; dorsal and lateral scales coarsely granuliform .... P. granuliferum H. L. Clark, 1938 (southern Australia) — Mid-ventral radial tube feet present as scattered series; dorsal and lateral tube feet conspicuous in mid-body; dorsal and lateral scales finely granuliform P. karenae sp. nov. (South Australia) Psolidium Ludwig Figures le, 2d, 4b-d, 5e, 8f Psolidium Ludwig, 1886: 9.— Mortensen, 1927: 413.— Deichmann, 1941: 141-143.— Deichmann, 1947: 336.— Lambert, 1996: 21. Diagnosis. Dendrochirotid holothuroids; small, up to 40 mm long; mid-body arched dorsally in transverse section, flat ventrally; dorsal and lateral body covered with imbricating scales, usually macroscopically conspicuous, sometimes obscured by integument, scales irregular in size and arrangement; scales decreasing in size ventro-laterally, orally and anally; lacking large oral valves; extensible oral cone, anterior or anterior-dorsal or dorsal orientation; extensible anal cone, posterior or posterior-dorsal or dorsal orientation; tube feet dorsally and laterally in mid-body, pass through scales. Sole distinct, oval to elongate; discrete margin created by junction of small imbricating ventro -lateral scales with thin- walled, usually calcareous sole that lacks scales; peripheral band of tube feet, may be discontinuous across the inter-radii anteriorly and posteriorly; peripheral tube feet frequently of 2 sizes, those of outer series smaller; mid-ventral radial series of tube feet present or absent. Calcareous ring solid, plates sub -rectangular, radial and interradial plates with tapered anterior projections; radial plates with deep notch posteriorly, interradial plates with shallow concave indentation posteriorly; 10 dendritic tentacles, ventral 2 smaller. Dorsal and lateral ossicles: multi-layered or single-layered perforated plates (scales), always some with tube foot canals; integument covering scales may have cupped crosses, cups, “thorn” ossicles (irregular branched rods pointed distally), buttons, perforated plates and rosettes; tube foot small endplates, and tube foot support ossicles that are irregular rods and plates, bent and curved, variably perforated. Sole ossicles: inter-radii with small to large single-layered perforated plates (rarely with multi -layering), smooth to variably knobbed and thickened, sometimes with cupped crosses, cups, thorn ossicles and rosettes; radii with additional tube foot ossicles, large endplates and tube foot support ossicles that are irregular rods and plates, bent and curved, variably perforated. Australian species of Psolidium Ludwig 5 Figure 1. Photos of live specimens of Psolidium Ludwig, 1886 (a-d, in situ by K. Gowlett- Holmes; e, in aquarium by L. Altoff): a, P. granuliferum H. L. Clark, 1938, Port Davey, Tasmania (14 mm long preserved; SAM K2174, PK0259); b, P. karenae sp. nov., Yorke Peninsula, South Australia (17 mm long preserved; holotype SAM K2177, PK0105); c, P. laperousazi sp. nov.. Kangaroo I, South Australia (13 mm long preserved; paratype SAM K2179, PK0253); d, P. marshae sp. nov.. Kangaroo I (10 mm long preserved; SAM K2173, PK0273); e, P. ravum Hickman, 1962, Portland, Victoria (17 mm long preserved; NMV FI 25379). 6 P. Mark O’Loughlin and Dragica Marie Figure 2. Photos of preserved specimens of Psolidium Ludwig, 1886 (a, c-d by L. Altoff; b, e-f by S. Hinkley and D. Marie): a, P. berentsae sp. nov.. Lizard I, Queensland (lateral view; 12 mm long; holotype AM J24098); b, P. granuliferum H. L. Clark, 1938, Smokey Bay, South Australia (dorso-lateral view, with mouth left; 19 mm long; SAM K2176); c-d, P. hutchingsae sp. nov.. Split Solitary I, New South Wales (20 mm long; holotype AM J24107); c, dorso-lateral view; d, 10 tentacles, with 2 small ventral ones bottom; e, P. karenae sp. nov., Adelaide (lateral view, mouth left; 20 mm long; paratype SAM K2185); f, P. laperousazi sp. nov., D’Entrecasteaux Channel, Tasmania (dorso-lateral view, mouth right; 18 mm long; paratype SAM K2339). Australian species of Psolidium Ludwig 7 Figure 3. Photos of preserved specimens of Psolidium Ludwig, 1886 (by L. Altoff): a, P. marshae sp. nov., Geographe Bay, Western Australia (dorsal view; 8 mm long; holotype WAM Z31173); b, P. mccallumae sp. nov., slope off Point Cloates, Western Australia (lateral view; 7 mm long; holotype NMV F126891); c-d, P. nigrescens H. L. Clark, 1938, Gunnamatta Bay, New South Wales (40 mm long; AM J6821); c, lateral view; d, ventral view (sole); e-f, P. parmatus (Sluiter, 1901), Adele slope. Western Australia (9 mm long; NMV F151835); e, lateral view; f, ventral view (sole). 8 P. Mark O’Loughlin and Dragica Marie Figure 4. a, P. spinuliferus (H. L. Clark, 1938), Darwin (lateral view; 10 mm long; AM J24096; photo by L. Altoff); b, generalised form of radial (left) and interradial plates of the calcareous ring of Psolidium species (drawing by M. O’Loughlin); c-d, drawings of sole showing distribution of tube feet (by D. Marie); c, P. granuliferum H. L. Clark, 1938 (SAM K2176); d, P. karenae sp. nov. (SAM K2188). Tentacle ossicles: rods variably perforated, thick to thin, long to short, straight or bent, flat or curved; dendritic tentacle branch endplates are small, irregular in shape, cupped, with a few large perforations and irregular margin; perforated plates may be present; densely branched rosettes may be present. Type species. Psolidium dorsipes Ludwig, 1886. Australian species. Psolidium berentsae sp. nov., P. granuliferum H. L. Clark, 1938, P. hutchingsae sp. nov., P. karenae sp. nov., P. laperousazi sp. nov., P. marshae sp. nov., P. mccallumae sp. nov., P. minutus (H. L. Clark, 1938), P. nigrescens H. L. Clark, 1938, P. parmatus (Sluiter, 1901), P. ravum Hickman, 1962, and P. spinuliferus (H. L. Clark, 1938). Remarks. The descriptive term “thorn” ossicles is used for the body wall ossicles in the form of irregular short branched rods that are pointed distally. They occur in Psolidium nigrescens H. L. Clark and P. parmatus Sluiter. H. L. Clark (1938) referred to them as “triradiate particles/spicules”. Sluiter (1901) did not report their presence. The type of P. parmatus was examined by one of us (PMO), and their presence noted. Pawson and Valentine (1981) reported that their Atlantic species Psolidium prostratum lacked endplates in the dorsal tube feet. Thandar (2006) described Psolidium multipes from South Africa, and among characteristic Psolidium features reported that the sole was not distinct from the dorsum, the ventral tube feet were in radial series, and there were multi- layered ossicles in the sole. These characters are atypical of Psolidium, but may indicate juvenile form. The Australian species of Psolidium are distinguished from each other in the key. The key indicates three morphological groupings of Australian Psolidium species: P. parmatus and P. spinuliferus and with their vertical pillars/ spires on dorsal and lateral scales; species with cupped crosses and cups (as for the type species P. dorsipes)-, and species lacking cupped crosses and cups. Australian species of Psolidium Ludwig 9 Figure 5. Photos of ossicles from Australian species of Psolidium Ludwig, 1886 (by Mark O’Loughlin and Chris Rowley): a-c, P. berentsae sp. nov.; a, dorsal cupped crosses and cups (small specimen, paratype AM J24099); b, dorsal bridged cup (holotype AM J24098); c, dorsal cups and rosettes (holotype AM J24098); d-f, P. hutchingsae sp. nov.; d, dorsal cupped cross (paratype AM J24108); e, dorsal multi-layered ossicle (scale) with tube foot canals (AM J19665); f, tentacle plate and rosettes (AM J19665). 10 P. Mark O’Loughlin and Dragica Marie Figure 6. Photos of ossicles from Australian species of Psolidium Ludwig, 1886 (by Mark O’Loughlin and Chris Rowley): a, P. granuliferum H. L. Clark, 1938, multi-layered ossicles from the sole (SAM K2174); b-c, P. laperousazi sp. nov. (paratype SAM K2179); b, dorsal partly closed cup; c, cup and plate from sole; d-f, P. marshae sp. nov. (paratype WAM Z31165); d, dorsal small cupped crosses and rosettes; e, large cup and knobbed plate from sole; f, dorsal buttons, small endplate, large and small cupped crosses and rosette. Australian species of Psolidium Ludwig 11 Figure 7. Photos of ossicles from Australian species of Psolidium Ludwig, 1886 (by Mark O’Loughlin and Chris Rowley): a-b, P. mccallumae sp. nov. (holotype NMV F126891); a, dorsal cups; b, mount of part of sole body wall, with plates and cups; c-d, P. minutus (H. L. Clark, 1938) (NMV F93176); c, dorsal large cup and small cupped crosses; d, large cup and small cupped crosses from sole; e-f, P. nigrescens H. L. Clark, 1938 (AM J6821); e, knobbed plates from sole; f, tentacle “thorns” and rosettes. 12 P. Mark O’Loughlin and Dragica Marie Figure 8. Photos of ossicles from Australian species of Psolidium Ludwig, 1886 (by Mark O’Loughlin and Chris Rowley): a-c, P. parmatus (Sluiter, 1901); a, dorsal scale with tube foot canal (holotype V.ECH.H1300); b, pillars on edge of part of scale (NMV F109378); c, dorsal “thorns” (holotype V.ECH.H1300); d-e, P. ravum Hickman, 1962 (SAM K2180); d, dorsal cupped crosses; e, cupped cross and plate from sole; f, P. spinuliferus (H. L. Clark, 1938), tentacle rods and dendritic branch endplates (top right) (AM J24096). Australian species of Psolidium Ludwig 13 Psolidium berentsae sp. nov. Table 1, Figures 2a, 5a-c Material examined. Holotype: Queensland, Lizard I., off western side of Palfrey I., washings from coral blocks, 14°40'S 145°28'E, 6 m, P. B. Berents and P A. Hutchings, 12 Jan 1976, stn 76 LIZ 16B, AM J24098. Paratypes: type locality and date, J24099 (2); type locality and date, stn 76 LIZ 16A, J24100 (1). Other material (up to 6 mm long; no cups detected): Chinamans Head, washings from coral blocks, 14°36'S 145°37'E, 6 m, P A. Hutchings and P B. Weate, 10 Jan 1976, stn 76 LIZ 15, J24101 (1); off Chinamans Head, reef rock, 14°40'S 145°28'E, 7 m, P. A. Hutchings and P B. Weate, 27 May 1976, stn 76 LIZ B-00-03-3, J24102 (1); No Name Reef, southwest end, rubble from base of bommie, 14°40'S 145°39'E, 15 m, I. Loch, 19 Dec 1984, J24106 (1). Other material (up to 11 mm long; no ossicles; presumably initial formalin preservation): Lizard I, off Chinamans Beach, coral block, 14°40'S 145°28'E, 7 m, P. A. Hutchings, Apr 1978, stn D15-27-1, J24104 (1); Outer Yonge Reef, 14°37'S 145°38'E, 18 m, P. A. Hutchings, 15 Jan 1977, stn 77 LIZ 47-4, J24105 (3). Description. Psolidium species up to 12 mm long (preserved); body rounded in transverse section ventrally; dorsal and lateral scales thick, up to 1.3 mm wide; oral, anal and mid-body scales frequently tapering to projecting, bluntly pointed distal end, body surface very uneven. Sole with peripheral band of tube feet, outer single series of slightly smaller tube feet, inner series 2 wide; mid-ventral radial series 2 wide; sole not discrete in very small specimens (4-6 mm long), small scales conspicuous. Dorsal and lateral ossicles: multi-layered ossicles (scales), thick, with tube foot canals; buttons numerous, perforated, thick, irregularly oval, thickly knobbed, up to 176 pm long, up to 12 perforations, intergrade with multi-layered ossicles; in larger specimens (12 mm long) cups numerous, deeply cupped, thick cruciform base, rim bluntly spinous, cups sometimes “bridged” with transverse branches from rim joined, cups 56-96 pm long; in small specimens (4—6 mm long) numerous thin cupped crosses and cups, with elongate pointed spines on rim, cups up to 80 pm long; rosettes numerous in larger specimens, densely branched, irregularly oval in form, up to 56 pm long. Sole ossicles: numerous knobbed plates, elongate, irregular in shape, large marginal and surface knobs, 3-12 perforations, up to 240 pm long; rare smooth elongate, perforated plates, up to 240 pm long. Tentacle ossicles include abundant rosettes, densely branched, up to 40 pm long. Colour (preserved). Dorsal and lateral dark to pale brown, sometimes with brown flecking; sole off-white; tentacle trunks brown, dendritic branches off-white; introvert off-white. Distribution. Queensland, Lizard I, 14°36'-14°40'S 145°28'-145°39'E, coral rocks and rubble; 6-18 m. Etymology. Named for Dr Penny Berents (Scientific Officer, Marine Invertebrates, Australian Museum), with appreciation of her contribution to Australian marine invertebrate research, and with gratitude for her gracious assistance with loans and research in the Australian Museum. Remarks. Some of the paratype specimens are small, 4 mm long (J24099 (2)). In this material the cups and cupped crosses are much finer than in the larger type material. This is judged to be a developmental difference. Some of the non-type specimens assigned to the new species are small, with specimen lengths 4 mm (J24101), 5 mm (J24102) and 6 mm (J24106). No cups or cupped crosses were detected, and this was judged to be a sampling inadequacy for this very small material, since fine cups and cupped crosses were found in 4 mm long paratype. The specimens have the same appearance as the types, and are from the type locality and eco-niche. Further collecting and study might reveal the presence of an additional species. Other additional material judged to be P. berentsae sp. nov. has the body form and scale outlines of the species, but lacks calcareous material. The preservation history presumably included time in acidic formalin solution. Specimen lengths are 5 mm (J24104) up to 11 mm (J24105). A rounded ventmm, absence of discrete sole and presence of conspicuous scales ventrally appearto be juvenile developmental characteristics. The distinguishing characters of P. berentsae sp. nov. are the tapered body scales, rounded ventrum, fine cups with long pointed spines in small specimens, and thick cups with blunt spines and sometimes “bridges” in large specimens. Psolidium granuliferum H. L. Clark Table 1, Figures la, 2b, 4c, 6a Psolidium granuliferum H. L. Clark, 1938: 503-04, fig. 49.— H. L. Clark, 1946: 415.— Rowe, 1982: 458, 464, pi. 31.4.— Rowe (in Rowe and Gates), 1995: 318. Material examined. Holotype: Western Australia, Koombana Bay, Bunbury, 9-15 m, C-A-H Expedition, 26 Oct 1929, MCZ 1666. Other material. Cape Nauraliste, Geographe Bay, Map Reef, near Two Rocks, limestone reef, coralline foliose algae, 31°28.365'S 115°33.634'E, 7.6 m, A. Sampey, 18 Jul 2005, WAM Z31164 (1); South Australia, W Eyre Peninsula, Smokey Bay, 37 m, N. Pearsons, 1971, SAM K2176 (2); Tasmania, Port Davey, Bramble Cove, NW end, under rock, 4-6 m, K. L. Gowlett-Holmes, 17 Mar 2003, SAM K2174 (2; live colour photo SAM PK 0259). Description. Psolidium species up to 21 mm long (preserved); body height low to moderately high, dorsal and lateral scales thick, up to 3 mm wide, lumps on surface creating coarsely granuliform appearance; abundant small scales at base of larger scales; tube feet dorsally and laterally inconspicuous, sparse across mid body, usually present near base of scales. Sole: peripheral band of tube feet, outer single series of distinctly separate smaller tube feet; series of inner larger tube feet 2 wide, series not always continuous anteriorly and posteriorly; lacking series of mid-ventral radial tube feet. Dorsal and lateral ossicles: multi-layered perforated ossicles (scales), thick, large surface lumps, canals for tube feet, intergrade with single-layered perforated plates; lacking cups, cupped crosses, rosettes. Ventral ossicles: small to large single-layered perforated plates, up to 22 perforations, thin to moderately thick, irregularly round to oval, up to 312 pm long, plate surfaces with abundant fine to medium knobs and rare large peripheral knobs, plate margins knobbed to bluntly spinous (fewer surface and peripheral knobs in plates from small specimens); large specimens have 14 P. Mark O’Loughlin and Dragica Marie perforated plates with secondary surface developments, intergrading into multi-layered ossicles. Tentacles lack rosettes. Colour. Live. Dorsally and laterally grey with off-white; white ventral ly. Preserved. Dorsally and laterally grey, or pale to darker brown with off-white; off-white ventrally. Distribution. SW Western Australia (Koombana Bay) to SE Tasmania (Port Davey); 4-37 m. Remarks. The distinguishing characteristics of Psolidium granuliferum H. L. Clark, 1938 are the absence of a mid-ventral radial series of tube feet on the sole, inconspicuous dorsal and lateral tube feet, coarsely granuliform dorsal and lateral scales, numerous small scales at the base of the large dorsal and lateral scales, and multi-layered ossicles in the sole of larger specimens. In the smallest specimens the sole has perforated plates with fewer knobs, and lacks perforated plates with secondary surface developments and multi-layered ossicles. Psolidium hutchingsae sp. nov. Table 1, Figures 2c-d, 5d-f Material examined. Holotype: N New South Wales, 50 m west of Split Solitary I, 30°14'S 153°10'48"E, Herdmania momus, rocks, sponges, ascidians, 15 m, P. A. Hutchings and L. C. Rose, 7 Mar 1992, stn NSW 677, AM J24107. Paratype: Type location and date, J24108 (1). Other material. Coffs Harbour, SE Islet, 30°19'S 153°09'E, 12 m, A. Hoggett and D. Johnson, 22 Jan 1982, J15469 (1); Port Stephens, 32°37'S 152°04'E, surface of ascidian, S. Smith, 1985, J19665 (1). Description. Psolidium species up to 18 mm long (preserved); dorsal and lateral body scales thick, up to 2.2 mm wide, body surface uneven; oral and anal scales tapering to narrow rounded end distally. Sole with peripheral irregular band of tube feet, about 4-5 wide, lacking discrete outer series of smaller tube feet; mid- ventral radial series of tube feet, about 2 wide. Dorsal and lateral ossicles: multi-layered ossicles (scales) thick, up to 4 tube foot canals; buttons perforated, irregularly oval, thick, smallest 80 pm long with 3 perforations, intergrade with thickened and knobbed perforated plates and with multi- layered ossicles; crosses abundant, deeply cupped, arms bifurcate, arms distally finely spinous, cupped crosses 56-88 pm long; rosettes rare, up to 32 pm long. Sole ossicles: knobbed to thickened perforated plates, marginal and surface knobs, variable shape from flat crosses to irregular plates with up to 12 perforations, up to 160 pm long; shallow concave crosses rare, arms bifurcate, arms bluntly spinous to finely knobbed distally, up to 72 pm long; shallow cups rare, knobs to short digitiform spinelets on rim, cups up to 96 pm long. Tentacle ossicles include irregular, thick, perforated, plates, up to 320 pm long; abundant rosettes, densely branched, up to 80 pm long. Colour. Preserved. Dorsally and laterally pale to dark grey- brown, some specimens with dark brown patches or spotting; sole off-white; tentacle trunks brown, dendritic branches off- white; introvert off-white. Distribution. Northern New South Wales, Split Solitary I, Coffs Harbour and Port Stephens; rock, sponge, ascidians; 12-15 m. Etymology. Named for Dr Pat Hutchings (Senior Principal Research Scientist, Australian Museum), with appreciation of her contribution to Australian marine invertebrate research and in particular for the collection and documentation of specimens described in this work. Remarks. The distinguishing characteristics of Psolidium hutchingsae sp. nov. are the presence dorsally of abundant deeply cupped crosses with bifurcate arms that are distally finely spinous, and large rosettes and irregular plates in the tentacles. Psolidium karenae sp. nov. Table 1, Figure lb, 2e, 4d Material examined. Holotype: South Australia, Spencer Gulf, Yorke Peninsula, Port Victoria jetty, under rock, 5-6 m, K. L. Gowlett- Holmes and W. Zeidler, 14 Dec 1994, SAM K2177 (live colour photo SAM PK 0105). Paratypes: St Vincent Gulf, Kemps Ground, 12 m, N. Holmes, Apr/May 1985, K2188 (1); Adelaide, Hallet Cove, R. Balfour, 1970, K2184 (3); Moana Beach, reef off S end, 10 m, K. L. Gowlett-Holmes, 4 Apr 1987, K2185 (3); Nuyts Archipelago, just N of centre of W island, Franklin I, 6-8 m, P. Aerfeldt et al., 14 Apr 1983, SAM K2187 (1); St Francis I, W end of N beach, among rocks, 3 m, W. Zeidler, 24 Jan 1982, K2181 (2). Description. Psolidium species up to 32 mm long (preserved); body height low to moderately high, dorsal and lateral body scales thick, finely granuliform, large, up to 4.5 mm wide, with sparse small scales at base; dorsal and lateral tube feet conspicuous, pass through centre of scales. Sole: peripheral band of tube feet, outer single series of distinctly separate smaller tube feet, inner series of larger tube feet, 2-3 wide, series not always continuous anteriorly and posteriorly; mid-ventral radial series of tube feet, irregularly spaced, 1-2 wide. Dorsal and lateral ossicles: multi-layered perforated ossicles (scales), thick, fine surface lumps, canals for tube feet, intergrade with single-layered perforated plates up to 160 pm\ lacking cupped crosses, cups, rosettes. Ventral ossicles: small to large single-layered perforated plates, typically up to 24 perforations, up to 46 perforations in larger specimens, thin to moderately thick, irregularly round to oval, up to 240 pm long, plate surfaces with abundant fine to medium knobs and rare large peripheral knobs, plate margins knobbed to bluntly spinous; perforated plates from small specimens with fewer surface and peripheral knobs; large specimens lack secondary surface developments on perforated plates, and multi-layered ossicles. Tentacles lack rosettes. Colour. Live. Dorsally dark reddish-brown. Preserved. Dorsally pale to dark brown to grey-brown; sole off-white to cream to pale brown. Distribution. South Australia, St Vincent Gulf to Nuyts Archipelago; 3-12 m. Australian species of Psolidium Ludwig 15 Etymology. Named for Karen Gowlett-Holmes (CSIRO Marine and Atmospheric Research; Eaglehawk Dive Centre) in appreciation of her contribution to marine invertebrate research, and with gratitude for her fieldwork and photography that contributed substantively to this work. Remarks. The distinguishing characteristics of Psolidium karenae sp. nov. are the conspicuous dorsal and lateral tube feet, finely granuliform dorsal and lateral scales, sparse small scales at the base of the large dorsal and lateral scales, absence of cups, cupped crosses and rosettes, and presence of separate single outer series of smaller peripheral tube feet around the sole. In the sole of the smallest specimens the perforated plates have fewer knobs. Psolidium laperousazi sp. nov. Table 1, Figures lc, 2f, 6b-c Material examined. Holotype: SE Tasmania, D’Entrecasteaux Channel, Tinderbox, E of boat ramp, 1.5-3 m, K. L. Gowlett-Holmes, 14 Jul 1991, SAM K2172. Paratypes: type locality and date, K2339 (2); South Australia, Kangaroo I, b/wn Western River Cove and Snug Cove, W side, 8-10 m, K. L. Gowlett-Holmes, 24 Feb 2003, K2179 (1, live colour photo SAM PK0253). Description. Psolidium species up to 27 mm long (preserved); body height low to moderately high; dorsal and lateral scales thick, embedded in thick integument, imbricating, up to 1.5 mm wide; dorsal and lateral tube feet conspicuous, abundant. Sole: peripheral band of tube feet, 3-4 wide, not always continuous anteriorly and posteriorly, lacking series of distinctly separate outer small tube feet; regular, close series of mid-ventral radial tube feet, 2 wide. Dorsal and lateral ossicles: multi-layered perforated ossicles (scales), thick, irregular form; deeply cupped crosses and more abundant partially and fully closed cups, 40-72 pm, typical size range 56-64 pm long, cup rim densely, finely spinous; small to large buttons, up to 12 perforations, thick, smooth, irregular form, up to 264 pm long, intergrade with multi-layered ossicles. Ventral ossicles: predominantly large single-layered perforated plates, up to 20 perforations, rarely up to 42 perforations in larger specimens, variable thickening, irregularly round to elongate, smooth, irregular margin, up to 368 pm long; perforated plates with variable surface and peripheral knobs rare; perforated plates with 4 perforations rare; moderately deep to deeply cupped crosses and more abundant partially and fully closed cups, 48-64 pm long, typical size range 48-56 pm long, cup rim bluntly spinous. Tentacle ossicles include rosettes, irregularly oval, densely branched, up to 80 pm long in larger specimens. Colour. Live: dark pink dorsally. Preserved: pale brown to cream dorsally and ventrally; small, discrete, dark brown spots may be present on the sole. Distribution. SE Tasmania (D’Entrecasteaux Channel) to South Australia (Kangaroo I); 1-10 m. Etymology. Named for Thierry Laperousaz (Collection Manager, Marine Invertebrates, South Australian Museum), with gratitude for his prompt and gracious assistance with loans. Remarks. The distinguishing characteristics of Psolidium laperousazi sp. nov. are the embedding of the dorsal and lateral scales in thick integument, the abundant and conspicuous tube feet, the predominance of cups over cupped crosses dorsally and ventrally, large rosettes in the tentacles, and the presence of dark spots on the sole. Psolidium marshae sp. nov. Table 1, Figures Id, 3a, 6d-f Material examined. Holotype: Western Australia, Cape Naturaliste, Geographe Bay, Bunker Bay, granite reef, brown algae canopy, Cystophora, 33°32.152'S 115°01.993'E, 5.4 m, A. Sampey, 15 Apr 2005, WAMZ31173. Paratype: Eagle Bay, granite reef, brown algae canopy, Sargassum, Cystophora, 33°33.387'S 115°04.078'E, 4.9 m, A. Sampey, 2 Feb 2005, Z31165 (1). Other material. South Australia, Kangaroo I, b/wn Western River Cove and Snug Cove, 12-14 m, under rock, on coralline red alga or sponge, K. L. Gowlett-Holmes, 14 Mar 2004, SAM K2173 (1, live colour photo SAM PK0273). Description. Psolidium species up to 10 mm long (preserved); very uneven surface; dorsal and lateral scales thick, up to 1.5 mm wide; oral and anal scales small, tapering to rounded end distally. Sole with peripheral band of tube feet, band up to 4 wide, size variable, outermost series slightly smaller; mid-ventral radial series irregular, up to 4 wide; peripheral series of tube feet variably continuous anteriorly and posteriorly. Dorsal and lateral ossicles: multi-layered ossicles (scales) thick, tube foot canals; buttons numerous, thick, smooth, not knobbed, irregularly oval, 3-9 perforations, up to 216 pm long; crosses small, deeply cupped, most quadri-radiate, some tri- or penta-radiate, abundant, typically 32 pm long, arms distally spinous, spinelets long, spinel ets sometimes joined to close rim to form cups; rosettes abundant, oval, typically 32 pm long. Sole ossicles: knobbed plates numerous, variable form, 4-16 perforations, thickly knobbed on surface and marginally, up to 240 pm long; cups and cupped crosses shallow concave, with digitiform spinelets marginally only; cups and cupped crosses of 2 sizes, larger typically 72 pm long, of variable form, some intergrading with knobbed plates, smaller typically 32 pm long, quadri-radiate or tri-radiate base, marginal rim closed or not. Tentacle ossicles include abundant rosettes, oval, up to 56 pm long. Colour. Live. Predominantly white with some brown patches. Preserved : dorsally and laterally off-white with dark brown-black patches and flecks; sole off-white; tentacle trunks with dark brown markings. Distribution. Western Australia, Cape Naturaliste, Geographe Bay; granite; 5 m; South Australia, Kangaroo I; 12-14 m. Etymology. Named for Loisette Marsh (Research Associate, Marine Invertebrates Section, Department of Aquatic Zoology, 16 P. Mark O’Loughlin and Dragica Marie Western Australian Museum) , in appreciation of her considerable contribution to echinoderm systematic and biogeographical research, and with gratitude for her gracious assistance with loans of echinoderm specimens for systematic research. Remarks. The distinctive characters of Psolidium marshae sp. nov. are the presence of 2 size ranges of cupped crosses and cups in the sole, and presence of rosettes in the body wall. Psolidium mccallumae sp. nov. Table 1, Figures 3b, 7a-b Material examined. Holotype: Western Australia, off Point Cloates, upper continental slope, hard substrate. Southern Surveyor SS10/2005 stn 139, 22.85°S 113.51°E, 100 m, 9 Dec 2005, NMV F126891. Description. Psolidium species up to 7 mm long (preserved); dorsal and lateral body scales thick, up to 1.3 mm wide, oral and anal cone scales tapering to projecting angled spine; dorsal and lateral tube feet conspicuous. Sole with peripheral band of tube feet, outer single series of slightly smaller tube feet, inner single series; lacking mid- ventral radial series of tube feet. Dorsal and lateral ossicles: multi-layered ossicles (scales), up to 2 tube foot canals; cups shallow, 4-8 perforations, cross and rim covered with fine spinelets, cups 40-72 pm long; rosettes rare, small, up to 32 pm long. Sole ossicles: smooth, irregular perforated plates, up to 16 perforations, up to 184 pm long; shallow cups with finely spinous rim, typically 4 perforations, typically 48 pm long; lacking rosettes. Colour ( preserved ). White dorsally, laterally, ventrally. Distribution. Central Western Australia, off Point Cloates, upper continental slope, hard substrate; 100 m. Etymology. Named for Anna McCallum (Marine Biology Section, Museum Victoria) with appreciation of her participation in the collection of this material on the Southern Surveyor, and her careful curation of this holothuroid collection in Museum Victoria. Remarks. The distinctive character of Psolidium mccallumae sp. nov. is the complete cover of small spinelets on the cross and rim of the shallow dorsal cups. Psolidium minutus (H. L. Clark) Table 1, Figures 7c-d Psolus minutus H. L. Clark, 1938: 507-09, figs 51-52.— H. L. Clark, 1946: 414.— Pawson, 1968b: 21 (key).— Cannon and Silver, 1987: 29. Psolidium minutus.— Rowe (in Rowe and Gates), 1995: 318.— Coleman, 2002: 65, fig. Material examined. Paratypes: Lord Howe I, Neds Beach, under rock, Apr 1932, AM J6450 (3). Other material. Neds Beach, under rock, N. Coleman, 20 Jul 2002, NMV F93176 (1); Middleton Reef, shallow reef front, 29°27.2'S 159°06.8’E, sandy bottom, 10 m, 4 Dec 1987, AM J20901 (1). Description. Psolidium species up to 10 mm long (live); dorsal and lateral body scales thick to thin, bluntly toothed on free edge, up to 1.3 mm wide; tube feet pass through scales. Sole with peripheral band of tube feet, outer single series of smaller tube feet, inner single series; mid-ventral radial series of tube feet 2 wide. Dorsal and lateral ossicles: single to multi-layered ossicles (scales) with tube foot canals; large cups shallow to deep, closely finely spinous rim, large cups 48-80 pm long; small cupped crosses and cups deep, closely finely spinous distal arm branches and cup rim, small crosses and cups up to 24 pm long; lacking rosettes. Sole ossicles: perforated plates, irregularly round to elongate, smooth, marginally slightly knobbed to bluntly spinous, up to 36 perforations, up to 360 pm long; shallow cups with finely spinous rim, typically 4 perforations, typically 48 pm long; very small deeply cupped crosses and cups, finely closely spinous on distal arm branches and rim, up to 20 pm long; lacking rosettes. Tentacles lack rosettes. Colour. Live colour white, tentacles yellowish (H. L. Clark, 1938; Coleman, 2002: 65, fig.). Distribution. Tasman Sea, Lord Howe I, Middleton Reef; 1-10 m. Remarks. The distinctive characters of Psolidium minutus (H. L. Clark, 1938) are the presence of 2 size ranges of cupped crosses and cups in the sole, and absence of rosettes in the body wall. H. L. Clark (1938) noted that Dr Deichmann had called his attention to the fact that the ossicles in Psolus minutus resembled those in some species of Psolidium. Rowe (1995) reassigned Psolus minutus to Psolidium on the basis of dorsal tube feet penetrating the scales. Psolidium nigrescens H. L. Clark Table 1, Figures 3c-d, 7e-f Psolidium nigrescens H. L. Clark, 1938: 505-507, fig. 50.— H. L. Clark, 1946: 414-15.— Cannon and Silver, 1987: 29.— Rowe (in Rowe and Gates), 1995: 318. Material examined. New South Wales, Broken Bay, Hawkesbury River, Yeomans Bay, Smith-Maclntyre Grab, mud sediment, G. Phillipson, Oct 1992 (specimen not lodged in museum); Port Jackson, Middle Harbour, N bank W of Spit Bridge, 33°48’S 151°15’E, steeply sloping bottom with telestacean bed, live and dead mussels, Tethia sp. and compound ascidians, 8 m, J. K. Lowry, 9 Jun 1981, J24097 (1); Botany Bay, off Dolls Point, dredged, 7-11 m, D. F. McMichael, Oct 1949, J6789 (4); Port Hacking, Gunnamatta Bay, 34°04'30"S 151°08'54"E, on shells of Anadara, I. Bennett, 2 Nov 1948, AM J6821 (6); Batemans Bay, Clyde R, edge of channel near southern bank downstream from Princes Hwy road bridge, 35°42’34"S 150°H'3"E, 5 m, clump of hairy mussels, Australian Museum party, 30 Mar 2004, J24092 (3). Description. Psolidium species up to 40 mm long (preserved); body elongate; dorsal and lateral body scales thick, up to 4 mm wide, covered by fairly thick integument; scales tapered to projecting rounded point orally and anally; dorsal and lateral tube feet conspicuous, up to 5 pass through a scale. Sole with peripheral band of tube feet, about 5 wide, outermost tube feet not smaller; mid-ventral radial series of tube feet predominantly 2 wide. Dorsal and lateral ossicles: multi-layered ossicles (scales) with tube foot canals; thick knobbed buttons, 3-10 perforations, Australian species of Psolidium Ludwig 17 up to 192 pm long, intergrade with multi-layered ossicles; branched rods with pointed ends (“thorn” ossicles), irregular form, 3-5 arms, flat to 3 -dimensional, up to 56 p m long; rosettes, densely branched, up to 40 pm long. Sole ossicles: perforated plates, slightly concave, irregularly round, knobbed marginally, up to 14 perforations, up to 152 ja, m long; very shallow, marginally knobbed, 4-perforation cups and cupped crosses, typically 96 jim long; rare rosettes; rare “thorn” ossicles. Tentacle ossicles include: rosettes, up to 40 jim long; “thorn” ossicles, up to 40 jim long. Colour ( preserved ). Dorsal and lateral body black to brown to dark grey, with fine white spotting over scales; tube feet off- white; sole pale brown to cream, with some fine brown flecking; tentacle trunks dark brown, tentacle dendritic ends off-white. Distribution. Eastern Australia, central New South Wales, Broken Bay to Batemans Bay, estuarine; 0-11 m. Remarks. The distinguishing characters of Psolidiumnigrescens H. L. Clark, 1938 are the “thorn” ossicles in the body wall, and very dark brown to black colour. Psolidium parmatus (Sluiter) Table 1, Figures 3e-f, 8a-c Psolus parmatus Sluiter, 1901: 102-03, pi. 6 fig. 14. Material examined. Holotype (very poor condition): Indonesia, Banda Sea, Siboga stn 259, 5°29.2'S 132°52.5'E, coral sand, 487 m, V.ECH. H1300. Other material. Western Australia, Ningaloo North, Southern Surveyor SS2005/10 stn 163, 21.94°S 113.84°E, hard substrate, 133 m, 12 Dec 2005, NMV F112156 (1); Adele, SS05/2007 stn 161, 14.5622°S 122.9182°E, hard rubble, 95 m, 4 Jul 2007, F151835 (1); Kulumburu, SS05/2007 stn 176, 13.2247°S 123.3957°E, 400 m, F151836 (1). Arafura Sea, Southern Surveyor SS05/2005 stn 42, 9°7.018’S 133°24.725'E, 204 m, 18 May 2005, F109377 (1); SS05/2005 stn 43, 9°5.312'S 133°2.989'E, 226 m, 19 May 2005, F109378 (1). Description. Psolidium species up to 13 mm long (preserved); dorsal and lateral scales up to 2.0 mm wide, scales each with numerous blunt calcareous vertical projections; dorsal and lateral tube feet not conspicuous amongst projections. Sole with peripheral band of tube feet, single outer series of smaller tube feet, inner single series of larger ones; lacking mid-ventral radial series of tube feet. Dorsal and lateral ossicles: single to multi-layered perforated plates (scales), lace-like secondary surface developments to multi-layering, tube foot canals; each scale with numerous thick, bulbous calcareous spires, constricted basally, rounded distally, about 200 pm high, 136 pm widest diameter, finely spinous surface; irregular branched pointed rod “thorn” ossicles, up to 160 pm long; shallow concave cups, perforations large, typically 4 around central cross, frequently 4 smaller corner perforations, cups knobbed around margin and on cross, cups typically 120 pm long; lacking rosettes. Sole ossicles: shallow concave cups, perforations large, typically 4 around central cross, frequently 4 smaller corner perforations, cups knobbed around margin and on cross, cups typically 112 pm long; “thorn” ossicles up to 96 pm long; lacking rosettes. Colour (preserved). White dorsally, laterally, ventrally. Distribution. Indonesia (Banda Sea), Arafura Sea, to NW Western Australia (Ningaloo North); 95-487 m. Remarks. Tube feet pass through the dorsal and lateral scales of Psolus parmatus Sluiter, 1901. The species is re-assigned here to Psolidium Ludwig. The distinctive species characteristic of Psolidium parmatus (Sluiter) is the presence of numerous blunt vertical calcareous projections on each of the dorsal and lateral scales. These projections resemble “shields” in profile, and hence the species name chosen by Sluiter (parma, Latin for “small shield”). Psolidium ravum Hickman Table 1, Figures le, 8d-e Psolidium ravum Hickman, 1962: 60-61, figs 75-86, pi. 2 fig. 7.— Dartnall, 1980: 51, 77,-Rowe, 1982: 458, 464, fig. 10.29c.-Rowe (in Rowe and Gates), 1995: 318-19. Material examined. Syntype: SE Tasmania, Derwent Estuary, Ralphs Bay, dredged, 13 m, 28 Aug 1959, AM J7202 (1). Other material: N point of Trial Bay, near Kettering, 3 m, under rock, 31 Mar 1991, SAM K2180 (1); Hobart, AM G248 (3); Frederick Henry Bay, Primrose Sands, Renard Point, under rock, 3-4 m, K. L. Gowlett Holmes, 25 Aug 1999, K2178 (1); Bass Strait, Lulworth, Black Rock Point, M. O’Loughlin et al„ 22 Nov 1982, NMV F97439 (1); Tamar R., Greens Beach, M. O’Loughlin, 7 Mar 1981, NMV F97440 (1). Victoria, Bass Strait, Ninety Mile Beach, 10 km W of Seaspray, J. Watson, 1977, AM J11171 (1); Portland, 500 m E of Whalers Point Lighthouse, 10 m, under rocks, P. Berents and R. Springthorpe, 10 May 1988, AM J21980 (1); Saxon Reef, 10 m, J. Watson, 24 Feb 2007, NMV FI 25379 (1). South Australia, Spencer Gulf, Yorke Peninsula, Port Elizabeth, 7 m, K. Sheard, 23 Feb 1941, SAM K2182 (1); Pt Turton jetty, 3-4 m, N. Holmes, 31 Aug 1986, SAM K2186 (1); W Eyre Peninsula, Streaky Bay, Pt Westall, littoral, M. O’Loughlin, 15 Jan 1991, NMV F 9743 8 (1); Nuyts Archipelago, Franklin I, 12-15 m, W. Zeidler and P. Aerfeldt, 23 Feb 1983, SAM K2183 (1). Description. Psolidium species up to 40 mm long (preserved); body form oval to elongate; body height low to moderately high; dorsal and lateral scales thick, up to 2 mm wide, covered with thick integument; dorsal and lateral tube feet conspicuous, numerous. Sole: peripheral band of tube feet, 4-5 wide, not always continuous anteriorly and posteriorly, lacking series of distinctly separate outer small tube feet; regular, close series of mid-ventral radial tube feet, 2 wide. Dorsal and lateral ossicles: multi-layered perforated ossicles (scales), thick, irregular form; deeply cupped crosses with distal spinous to digitiform spinelets, rare partially or fully closed cups, 32-72 pm long, typical size range 40-56 pm long; small to large buttons, up to 12 perforations, thick, smooth, irregular form, up to 216 pm long, intergrade with multi-layered ossicles. Ventral ossicles: predominantly large single-layered perforated plates, up to 16 perforations, rarely up to 38 perforations in larger specimens, variable thickening, irregularly round to elongate, smooth, irregular margin, up to 360 pm long; 18 P. Mark O’Loughlin and Dragica Marie perforated plates with variable surface and peripheral knobs rare; perforated plates with 4 perforations rare; moderately deep to deeply cupped crosses with distal spinous to knobbed spinelets, rare partially or fully closed cups, typical size range 4CM-8 p m long, rarely up to 56 pm long. Tentacle ossicles include rosettes, irregularly oval, densely branched, up to 96 pm long in larger specimens. Colour. Live: grey dorsally and laterally. Preserved: dark brownish-grey to off-white dorsally and laterally; sole brown to grey-brown. Distribution. SE Tasmania (Derwent Estuary), Bass Strait, to W South Australia (Nuyts Archipelago); 0-15 m. Remarks. Although not reported by Hickman (1962), the syntype and all of the specimens determined here to be P. ravum have rosettes in the tentacles. The specimen from the Eyre Peninsula (NMV F97438) ventrally has more abundant plates with 4 perforations, but the external morphology and presence of cupped crosses dorsally and ventrally are typical of P. ravum. The specimens from Frederick Henry Bay (SAM K2178) and Kettering (K2180) have similar morphological appearances to P. laperousazi sp. nov., but the dorsal and ventral ossicles are typical of P. ravum. Knobbed to spinous plate margins in the sole of the specimens from Port Elizabeth (SAM K2182) and Portland (NMV F125379) are similar to those of P. karenae sp. nov. and P. granuliferum H. L. Clark, but the external morphology and presence of dorsal and ventral cupped crosses and cups are typical of P. ravum. Small ossicles were absent from specimens from Nuyts Archipelago (SAM K2183) and Yorke Peninsula (K2186) (probably formalin preservation history), but the specimens have the typical external morphology of P. ravum. The distinguishing characteristics of Psolidium ravum Hickman, 1962 are the conspicuous dorsal and lateral tube feet, the predominance of cupped crosses over cups dorsally and ventrally, large rosettes in the tentacles, and absence of dark spots on the sole. Psolidium spinuliferus (H. L. Clark) Table 1, Figures 4a, 8f, 9a-d Psolus spinuliferus H. L. Clark, 1938: 509-11, fig. 53.— H. L. Clark, 1946: 414.— Cannon and Silver, 1987: 29.— Rowe (in Rowe and Gates), 1995: 319. Material examined. Northern Territory, Darwin Harbour, North Shell I, 12°2948"S 130°53'12"E, coral rubble covered with sponges and some algae, 5 m, P. A. Hutchings, 16 Jul 1993, stn NT 346, AM J24096 (1). Western Australia, Perth, Cottesloe, Mudurup Rocks, c70 m S of groyne, reef flat, Sargassum zone, on reef flat under thin veneer of sand, 31°59'51"S 115°45'01"E, 0-1 m, J. Keesing, 6 Feb 2007, WAM Z37479 (1); Trigg I, clOO m N of ‘island’, inshore mixed algal zone mid-platform with thin veneer of sand overlaying reef, 31°52’29"S 115°45’04"E, 0-1 m, J. Keesing, 19 Feb 2007, Z37478 (1); Waterman, Sargassum zone, mid-platform, 31 0 51'15"S 115°45'05"E, 0-1 m, J. Keesing, 14 Feb 2007, Z37468 (5); from mixed localities, Cottesloe and Trigg I, inter-tidal platforms, on reef flat under thin veneer of sand, J. Keesing, Feb 2007, Z37469 (1). Description. Psolidium species up to 20 mm long (preserved); dorsal and lateral body scales thin, single-layered, with spires, scales up to 1.5 mm wide; tube feet dorsally and laterally pass through scales, not conspicuous amongst spires. Sole: peripheral band of tube feet, outer single series of smaller tube feet, inner single to zig-zag series; mid-ventral radial series irregular, double to zig-zag to scattered. Dorsal and lateral ossicles: single-layered, thick, perforated plates (scales), irregularly oval, some with secondary thickening, most with vertical digitiform spire near margin; spires up to 400 pm long, 120 pm diameter, distally spinous. Sole ossicles: knobbed plates, numerous, predominantly regular 4-holed thin plates, smooth to finely knobbed marginally, typically 80 pm long; lacking cupped crosses, cups, rosettes. Tentacle ossicles include: thick perforated plates, elongate, variable form, some with secondary layer development, up to 352 pm long; numerous rosettes, large to small, frequently with 4 central perforations, densely branched, oval to elongate and distally rounded, up to 160 pm long, intergrade with elongate plates. Colour(live and preserved). White, dorsally and ventrally. Distribution. Northern Territory (Darwin), to Western Australia (Perth); 0-22 m. Remarks. The dorsal and lateral tube feet are not conspicuous, and were not noticed by H. L. Clark (1938). Psolus spinuliferus H. L. Clark, 1938 is reassigned here to Psolidium Ludwig. The distinguishing characters of Psolidium spinuliferus (H. L. Clark) are the predominantly single-layered scales with vertical digitiform marginal spire. The type specimen (MCZ no. 1669) was taken off the Eighty Mile Beach near Broome in northwestern Australia, at 18-22 m. Acknowledgments We are most grateful to the following for their gracious assistance: CynthiaAhearn (Smithsonian Institution; literature); Leon Altoff and Audrey Falconer (Marine Research Group, Field Naturalists Club of Victoria; photography); Ben Boonen (photoshop of images, format of figures); Mary Catherine Boyett (MCZ; loan); Karen Gowlett-Holmes (CSIRO, Hobart; live colour photos, field work); Simon Hinkley and Chris Rowley (NMV; photography); John Keesing (CSIRO, Perth; data), Thierry Laperousaz (SAM; loans); Loisette Marsh and Mark Salotti (WAM; loans and data); Stephen Keable and Roger Springthorpe (AM; loan, data); Jan Watson (Marine Research Group, Field Naturalists Club of Victoria; fieldwork). We are most appreciative of the helpful suggestions made by referees Dr D. L. Pawson and Dr F. W. E. Rowe. References Agassiz, L. 1845. Nomina systematica generum echinodermatum tarn viventium quam fossilium, secundum ordinem alphabeticum disposita, adjectis auctoribus, libris in quibus reperiuntur, anno editionis, etymologia et familiis, ad quas pertinent. Jent et Gassmann: Soloduri: Switzerland. Australian species of Psolidium Ludwig 19 0.25 mm 0.05 mm Figure 9. Photos of ossicles from Australian species of Psolidium Ludwig, 1886 (by Mark O’Loughlin and Chris Rowley): a-d, P. spinuliferus (H. L. Clark, 1938) ( WAM Z37468); a, part of dorsal scale with marginal vertical spire; b, dorsal scale with tube foot canal and base of lost spire; c, tentacle rosettes; d, tentacle plates. Agassiz, L. 1848. Nomenclatoris zoologici index universalis, continens nomina systematica classium, ordinum, familiarum et generum animalium omnium, tarn viventium quam fossilium, secundum ordinem alphabeticum unicum disposita, adjectis homonymiis plantarum. Jent et Gassmann: Soloduri: Switzerland. Baranova, Z.I., and Belyaev, G.M. 1968. Echinodermata. In: Zenkevich, L. A. (ed.). La vie des animaux. Izdatel stvo Proveshchenic. Moscow 2: 197-297. (in Russian) Bell, J.F. 1882. Studies in the Holothuroidea. I. On the genus Psolus and the forms allied thereto. Proceedings of the Zoological Society, London. 1882: 641-50, pi. 158. Bronn, H.G. 1860. Die klassen und ordnungen des thier-reichs, wissenschaftlich dargestellt in wort und bild. 2. Aktinozoen. C. F. Winter’sche Verlagshandlung: Leipzig and Heidelberg. 434 pp., 49 pis. Cannon, L.R.G., and Silver, H. 1987. Sea Cucumbers of Northern Australia. Queensland Museum: Brisbane, Australia, i-viii, 60 pp. Carriol, R-P, and Feral, J-P. 1985. Reexamen de quelques Psolidae (Holothurioidea, Echinodermata) antarctiques et subantarctiques. Description de deux nouvelles especes du genre Psolus. Bulletin du Museum national d’Histoire naturelle, Paris 4(7) A(l): 49-60. Cherbonnier, G. 1974. Invertebres marins des Xlleme et XVeme expeditions antarctiques Fran£aises en Terre Adelie. 15. Holothurides. Tethys 5(4): 601-10. Clark, H.L. 1901. Holothurians of the Pacific Coast of North America. Zoologischer Anzeiger 24: 162-71. Clark, H.L. 1938. Echinoderms from Australia. An account of collections made in 1929 and 1932. Memoirs of the Museum of Comparative Zoology at Harvard College 55: 1-596, 28 pis, 63 figs. Clark, H.L. 1946. The echinoderm fauna of Australia. Its composition and its origin. Carnegie Institution of Washington Publication 566: 1-567. Coleman, N. 2002. Lord Howe Island Marine Park. Neville Coleman’s Underwater Geographic. 95 pp. 20 P. Mark O’Loughlin and Dragica Marie Dartnall, A. 1980. Fauna of Tasmania Handbook 3. Tasmanian echinoderms. University of Tasmania: Hobart. 82 pp., 36 figs, 5 pis, 18 maps. Deichmann, E. 1940. Report on the Holothurians, collected by the Harvard-Havana expeditions 1938 and 1939, with a revision of the Molpadioida of the Atlantic Ocean. Memorias Sociedad Cubana Historia Natural, Felipe Poey 14(3): 183-240, 10 pis. Deichmann, E. 1941. The Holothurioidea collected by the Velero III during the years 1932-1938. I. Dendrochirotida. Allan Hancock Pacific Expeditions 8(3): 61-194, pis 10-30. Deichmann, E. 1947. Shallow water holothurians from Carbo de Hornos and adjacent waters. Anales del Museo Argentino de Ciencias Naturales. Invertebrados Marinos 8: 325-51. Deichmann, E. 1954. The holothurians of the Gulf of Mexico. Fishery Bulletin of the Fish and Wildlife Service 89(55): 381-410. Dendy, A. 1896. Observations on the Holothurians of New Zealand; with descriptions of four new species, and an appendix on the development of the wheels in Chirodota. Journal of the Linnaean Society of London, Zoology 26: 22-52. Forbes, E. 1841. A history of British starfishes and other animals of the class Echinodermata. John van Voorst: London. 267 pp. Gistel, T. 1848. Echinodermata. Class 11. Pp. 174-77 in: Naturgeschichte des Thierreichs fur hohere Schulen. Stuttgart. xvi+216 pp., 32 pis. Grube, A.E. 1840. Aktinien, Echinodermen und Wiirmer des Adriatischen und Mittelmeeres. J. H. Bon: Konigsberg. 92 pp. Gutt, J. 1988. Zur Verbreitung und Okologie der Seegurken (Holothuroidea, Echinodermata) im Weddellmeer (Antarktis). Berichte zur Polar for schung 41: 1-87. Gutt, J. 1990. New Antarctic holothurians (Echinodermata) —I. Five new species with four new genera of the order Dendrochirotida. Zoologica Scripta 19(1): 101-17. Haeckel, E., 1896. Systematische Phylogenie der Echinodermen. Pp. 348-504 in: Systematische Phylogenie der Wirbellosen Thiere (Invertebrata). Zweiter Teil des Entwurfs einer systematischen Stammesgeschichte. Reimer: Berlin. 720 pp. Hickman, V.V. 1962. Tasmanian sea-cucumbers (Holothuroidea). Papers and Proceedings of the Royal Society of Tasmania 96: 49-72, 2 pis, 186 figs. Jager, G.F. 1833. De Holothuriis. Dissertatio Inauguralis. Turici. 40 pp., 3 pis. Joshua, E.C. 1914. Victorian Holothuroidea, with descriptions of new species. Proceedings of the Royal Society of Victoria. New Series 27(1): 1-11. Lambert, P. 1996. Psolidium bidiscum, a new shallow-water psolid sea cucumber (Echinodermata: Holothuroidea) from the northeastern Pacific, previously misidentified as Psolidium bullatum Ohshima. Canadian Journal of Zoology 74: 20-31. Ludwig, H. 1886. Die von G. Chierchia auf der Fahrt der Kgl. Italianische Corvette Vettor Pisani gesammelten Holothurien. Zoologische Jahrbiicher 2: 1-36, 2 pis. Ludwig, H. 1892. Echinodermen. Pp. 32-61, pis 15-17 in: Bronn,H. G., Klassen und Ordnungen des Thier-Reichs 2(3) 3 Das System. C. F. Winter’sche Verlagshandlung: Leipzig and Heidelberg. 460 pp., 17 pis. Ludwig, H., and Heding, S. G. 1935. Die holothurien der Deutschen Tiefsee-Expedition. 1. Fusslose und dendrochirote formen. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition aufdem dampfer Valdivia 1898-1899 24: 123-214. Massin, C. 1992a. Holothurians (Echinodermata) from Marion and Prince Edward Islands: new and little-known species. Zoologica Scripta 21(3): 311-24. Massin, C. 1992b. Three new species of Dendrochirotida (Holothuroidea, Echinodermata) from the Weddell Sea (Antarctica). Bulletin de Tlnstitut Royal des Sciences Naturelles de Belgique, Biologie 62: 179-91. Massin, C. 1997. First record of a Psolidae (Holothuroidea, Echinodermata) in the Mediterranean Sea (Sicilian Channel). Bulletin de Tlnstitut Royal des Sciences Naturelles de Belgique, Biologie 67: 101-06. Mitsukuri, K. 1912. Studies on Actinopodous Holothurioidea. Journal of the College of Science, Tokyo Imperial University 29(2): 284 pp., 8 pis. Mortensen, T. 1925. Echinoderms of New Zealand and the Auckland- Campbell Islands. 4. Holothurioidea. Pp. 322-86 in: Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-1916. 29. Videnskabelige Meddelelser Dansk Naturhistorisk Forening, Kqbenhavn 79. Mortensen, T. 1927. Handbook of the Echinoderms of the British Isles. Oxford University Press: London. 471 pp. Ohshima, H. 1915. Report of the holothurians collected by the United States Fisheries Steamer Albatross in the northwestern Pacific during the summer of 1906. Proceedings United States National Museum 48: 213-91, pis 8-11. Oken, L. 1815. Lehrbuch der Naturgeschichte. 3. Zoologie. Jena: Germany. Pp. xxxviii+850+xviii. O’Loughlin, PM. 2000. A review of the cucumariid genus Psolidiella Mortensen (Echinodermata, Holothuroidea). Memoirs of Museum Victoria 58: 25-37. O’Loughlin, P.M., and O’Hara, T.D. 1992. New cucumariid holothurians (Echinodermata) from southern Australia, including two brooding and one fissiparous species. Memoirs of the Museum of Victoria 53: 227-66. Pawson, D.L. 1964. The Holothuroidea collected by the Royal Society Expedition to Southern Chile, 1958-1959. Pacific Science 18(4): 453-70. Pawson, D.L. 1967. The psolid genus Lissothuria. Proceedings of the United States National Museum 122(3592): 1-17. Pawson, D.L. 1968a. Some holothurians from Macquarie Island. Transactions of the Royal Society of New Zealand (Zoology) 10(15): 141-50. Pawson, D.L. 1968b. The echinozoan fauna of the New Zealand subantarctic islands, Macquarie Island, and the Chatham Rise. Bulletin of the New Zealand Department of Scientific and Industrial Research 187: 9-33, 3 figs, 1 pi. Pawson, D.L. 1968c. A new psolid sea cucumber from the Virgin Islands. Proceedings of the Biological Society of Washington 81: 347-50. Pawson, D.L. 1969a. Holothuroidea from Chile. Report No. 46 of the Lund University Chile Expedition 1948-1949. Sarsia 38: 121-45. Pawson, D.L. 1969b. Holothuroidea. Pp. 36-38, 1 pi. in: Bushnell, V.C., and Hedgpeth, J.W. (eds). Distribution of Selected Groups of Marine Invertebrates in Waters South of 35° S Latitude. Antarctic Map Folio Series (II). American Geographical Society: New York. Pawson, D.L. 1970. The marine fauna of New Zealand: Sea cucumbers (Echinodermata: Holothuroidea). Bulletin of the New Zealand Department of Scientific and Industrial Research 201: 7-65, 10 figs, 2 pis. Pawson, D.L. 1971a. The Western Australian psolid holothurian Ceto cuvieria (Cuvier). Journal of the Royal Society of Western Australia 54(2): 33-39. Pawson, D.L. 1971b. Ekkentropelma brychia n.g., n.sp., an Antarctic psolid holothurian with a functionally lateral sole. Proceedings of the Biological Society of Washington 84(14): 113-18. Pawson, D.L. 1982. Holothuroidea. Pp. 813-18; figs pp. 791, 792; pis 136, 137 in: Parker, S.P. (ed. -in-chief). Synopsis and Classification of Living Organisms. McGraw-Hill: New York. Australian species of Psolidium Ludwig 21 Pawson, D.L., and Fell, H.B. 1965. A revised classification of the dendrochirote holothurians. Breviora 214: 1-7. Pawson, D.L. and Valentine, J.F. 1981. Psolidium prostratum, new species, from off the east coast of the USA. (Echinodermata: Holothuroidea). Proceedings of the Biological Society of Washington 94(2): 450-54. Perrier, R. 1902. Holothuries. Pp. 273-554, pis 12-22 in Expeditions scientifiques du Travailleur et du Talisman pendant les annees 1880, 1881, 1882, 1883. Masson et Cie, Editeurs: Paris. Perrier, R. 1905. Holothuries antarctiques du Museum d’Histoire Naturelle de Paris. Annales des Sciences Naturelles. Zoologie 1: 1-146, 5 pis. Philippi, A. 1857. Vier neue Echinodermen des Chilenschen Meeres. Archiv Naturgeschicht 23: 130-34. Rowe, F.W.E. 1982. Sea-cucumbers (class Holothurioidea). Pp. 454-76, figs 10:26-10:37, pis 29-32 in: Shepherd, S.A., and Thomas, I.M. (eds). Marine Invertebrates of Southern Australia. Part 1. South Australian Government Printer: Adelaide. Rowe, F.W.E., and Gates, J. 1995. Echinodermata. In: Wells, A. (ed.). Zoological Catalogue of Australia 33: i-xiii, 1-510. CSIRO: Melbourne. Selenka, E. 1868. Nachtrag zu den Beitragen zur Anatomie und Systematik der Holothurien. Zeitschrift Wissenschaftliche Zoologische 18: 109-19, pi. 8. Semper, C. 1868. Holothurien. Reisen imArchipel der Philippinen. 2. Wissenschaftliche Resultate. Erster Band. Engelmann: Leipzig. 288 pp., 40 pis. Sluiter, C.P 1901. Die Holothurien der Sz'Zwga-Expedition. Siboga- Expeditie 44: 1-142, 10 pis. Buchhandlung en Druckerei v. E. J. Brill: Leiden. Studer, T. 1876. Uber Echinodermen aus dem antarktischen Meere und zwei neue Seeigel von den Papua-Inseln, gesammelt auf der Reise SMS Gazelle um die Erde. Monatsberichte der Koniglich Preussichen Akademie der Wissenschaften. Berlin. Pp. 452-65. Thandar, A.S. 2006. New species and new records of dendrochirotid and dactylochirotid holothuroids (Echinodermata: Holothuroidea) from off the east coast of South Africa. Zootaxa 1245: 1-51. Theel, H. 1886. Report on the Holothurioidea dredged by HMS Challenger during the years 1873-1876. Report on the scientific results of the voyage of HMS Challenger, Zoology 14(39): 1-290, 16 pis. Tommasi, L.R. 1969. Lista dos Holothurioidea recentes do Brasil. Contributes Avulsas Instituto Oceanografico, Sao Paulo 15: 1-29. Tommasi, L.R. 1971. Equinodermes do Brasil. I. Sobre alguncas especies novas e outras pouco conhecidas para o Brasil. Boletim Instituto Oceanografico, Sao Paulo 20(1): 1-23. Vaney, C. 1906. Holothuries. Expedition Antarctique Fran§aise (1903-1905). Sciences Naturelles: Documents Scientifiques. Masson et Cie, Editeurs: Paris. 30 pp., 2 pis, 1 map. Vaney, C. 1914. Holothuries. Deuxieme Expedition Antarctique Fran£aise (1908-10). Sciences Naturelles: Documents Scientifiques. Masson et Cie, Editeurs: Paris. 54 pp., 5 pis. Verrill, A.E. 1866. On the Polyps and Echinoderms of New England. Proceedings of the Boston Society of Natural History 10: 333-57. Verrill, A.E. 1867. Notes on the echinoderms of Panama and the west coast of America, with descriptions of new genera and species. Transactions of the Connecticut Academy of Arts and Sciences 1(2): 251-322. Memoirs of Museum Victoria 65: 23-42 (2008) ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs/index.asp Antarctic and Sub-Antarctic species of Psolidium Ludwig (Echinodermata: Holothuroidea: Psolidae) P. Mark O’Loughlin 1 and Cynthia Ahearn 2 1 Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, 3001, Victoria, Australia (email: pmo@bigpond. net.au) 2 Department of Invertebrate Zoology, MRC-163, National Museum of Natural History, Smithsonian Institution, PO Box 37012, Washington DC, 20013-7012, USA Abstract O’Loughlin, P.M., and Ahearn, C. 2008. Antarctic and Sub-Antarctic species of Psolidium Ludwig (Echinodermata: Holothuroidea: Psolidae). Memoirs of Museum Victoria 65: 23-42. Five new species of Psolidium Ludwig from Antarctica and Sub-Antarctica are described: Psolidium emilyae sp. nov., P. normani sp. nov.; P. pawsoni sp. nov.; P. schnabelae sp. nov.; P. whittakeri sp. nov. Psolidium incertum (Theel) is a junior synonym of Psolidium poriferum (Studer). Psolidium tenue Mortensen is raised out of synonymy with P. incertum. Six species of Psolidium Ludwig from Antarctica and Sub-Antarctica are diagnosed: P. disciformis (Theel); P. dorsipes Ludwig; P. gaini Vaney; P. incubans Ekman; P. poriferum (Studer); P. tenue Mortensen. A key is provided for the 11 Antarctic and Sub-Antarctic species of Psolidium Ludwig. Keywords Echinodermata, Holothuroidea, Psolidae, Psolidium, taxonomy, new species, key Introduction This study is based on the extensive collections of Antarctic and Sub-Antarctic holothuroids held in the National Museum of Natural History (NMNH) of the Smithsonian Institution in Washington DC, the New Zealand Oceanographic Institute (NZOI, now National Institute of Water and Atmospheric Research [NIWA]), and Museum Victoria (NMV), South Australian Museum (SAM) and Tasmanian Museum and Art Gallery (TMAG) in Australia. The material is representative of the Antarctic Ocean off both Eastern and Western Antarctica, and the Sub-Antarctic Islands in the Atlantic, Indian and Pacific oceans. The NMNH collection came from the United States Fish Commission Albatross expeditions, and the United States Antarctic Research Program (USARP), and collections by the research vessels Eastwind, Edisto, Eltanin, Glacier, Hero and A/as Orcadas (See O’Loughlin and Ahearn, 2005). The NIWA collections were made in the Ross Sea region by the research vessels Endeavour and Tangaroa. The NMV and SAM collections came from the Australian National Antarctic Research Expedition (ANARE), and were collected by the research vessels Aurora Australis and Nella Dan. Heard Island collections in TMAG were made for the Australian Antarctic Division by F/V Southern Champion. Some BANZARE holothuroids remained on loan to USNM, and were included in our study. Some specimens from the British Museum (The Natural History Museum) (BM[NH]) that were collected by the Discovery, Challenger and William Scoresby were also available for our study. Some specimens taken by R/V Verna were donated to NMNH by the American Museum of Natural History. Recently collected holothuroids were donated to NMV from the AMLR R/V Yuhzmorgeologiya (2003) and Icefish (2004) expeditions to the southern Atlantic. Holothuroid specimens and data collected recently by the New Zealand International Polar Year Census of Antarctic Marine Life Project expedition to the Ross Sea by R/V Tangaroa were made available for collaborative study at NIWA in Nelson (New Zealand). The type specimen for Psolidium poriferum (Studer, 1876) was borrowed from the Humboldt-Universitaet museum fur naturkunde in Berlin (ZMB). We have reviewed the systematics of species of Psolidium Ludwig, 1886, described to date from Antarctica and Sub- Antarctica, and recognise six species. O’Loughlin (2002) has previously synonymised Psolidium navicula Ekman, 1927, and Psolidium bistriatum Ludwig and Heding, 1935, with Psolidium (Cucumaria) coatsi Vaney, 1908, and referred P. coatsi to his new genus Psolicrux O’Loughlin, 2002. We have added five new species of Psolidium from Bouvet Island, South Sandwich Islands, South Shetland Islands, Weddell Sea, Ross Sea, Prydz Bay and Enderby Land. In the past some specimens from Antarctica and Sub-Antarctica have been determined as Psolidium incertum (Theel, 1886). These specimens, that are similar morphologically, represent a complex of five species. 24 P. Mark O’Loughlin and Cynthia Ahearn Three are new and are described here. Two belong to previously described but unused or synonymised taxa: Psolidium poriferum (Studer, 1876) and Psolidium tenue Mortensen, 1925. Specimens of Psolidium gaini Vaney, 1914, showed significant variations in ossicle size and form and the possibility of another species complex is suggested. Molecular data will be an important additional source of evidence for further refinement of our work. Preserved specimens of Psolidae typically have tentacles withdrawn, and there has been some historical reluctance to dissect to sample tentacles for ossicle study. We found tentacle ossicle form to be essential for diagnosis within the Psolidium poriferum species group. To assist in our diagnoses we describe “rod-plates”, derived from rods with branches joined or with side-connections to create perforations (Figures 6e, f). These are distinguished from smooth perforated plates that show no evidence of rod derivation. Materials and methods SEM and some digital photography were done by Cynthia Ahearn. Specimens were photographed directly using a Nikon Coolpix 995 digital camera with flash lighting. Ossicles were photographed using the same camera with a Wild M20-34504 compound microscope. SEM images were taken using a Leica Stereoscan 440 with LaB6 electron source. Some specimen photographs were taken by Leon Altoff and Audrey Falconer, with Mark O’Loughlin, using a Pentax K10D digital SLR with a variety of lenses and lit using two electronic flashes. Preserved specimens were placed on or suspended above black velvet. Photos of some ossicles were taken by Chris Rowley, with Mark O’Loughlin, using a Leica DM5000 B compound microscope, and Leica DC500 camera with montage software. Specimen registration number prefixes are: NMV F, SAM K, TMAG H, USNM E or without prefix (See O’Loughlin and Ahearn, 2005). Dendrochirotida Grube, 1840 (restricted Pawson and Fell, 1965) Psolidae Forbes, 1841 Remarks. A synonymy and diagnosis for Psolidae are provided by O’Loughlin and Marie (2008). Pawson and Fell (1965), and subsequent authors, have nominated Perrier (1902) as the systematic author of Psolidae. We recognise Forbes (1841) as the original author. Key to genera of Psolidae (See O’Loughlin and Marie, 2008) Key to Antarctic and Sub-Antarctic species of Psolidium Ludwig, 1886 1. Mid-ventral (sole) radial series of tube feet present 2 — Lacking mid-ventral (sole) radial series of tube feet 3 2. Ossicles in sole include irregularly oval to round disc-like smooth plates, variably with few minute, closed to almost closed, perforations P. disciformis (Theel, 1886) (S Chile) — Irregularly oval to round, disc-like smooth plates, variably with few minute, closed to almost closed, perforations, not present in sole P. dorsipes Ludwig, 1886 (S Chile, S Argentina, Strait of Magellan, Burdwood Bank, Falkland Is) 3. Dorsally with deep goblet-like cups, closed across rim, vertical spines above rim; brood-protects in folds of sole . P. incubans Ekman, 1925 (South Georgia) — Dorsally lacking deep goblet-like cups closed across rim; no brood-protection in folds of sole 4 4. Dorsal and lateral scales conspicuous; dorsal and lateral tube feet inconspicuous; lacking numerous dorsal and lateral tube feet support ossicles; crosses and small perforated plates rare to absent in sole 5 — Dorsal and lateral scales inconspicuous; dorsal and lateral tube feet numerous and conspicuous; dorsal and lateral tube foot support ossicles numerous; sole with numerous perforated plates 9 5. Largest tentacle trunk ossicles predominantly thick, narrow rod-plates and perforated plates with prominent rod thickenings P. tenue Mortensen, 1925 (Antarctic Peninsula, Ross Sea, Wilkes Land, Antipodes I., Prydz Bay) — Largest tentacle trunk ossicles predominantly smooth perforated plates 6 6. Dorsal scales frequently with well-developed, radiating- to-margin, linear, rounded thickenings between rows of marginal perforations .. P. poriferum (Studer, 1876) (Marion, Kerguelen, Heard, McDonald Is; MacRobertson Land) — Dorsal scales rarely with linear thickenings between rows of marginal perforations 7 7. Dorsal scales frequently with smooth white thickening; perforations reduced, many almost closed P.pawsoni sp. nov. (Weddell Sea, Ross Sea, Enderby Land) — Dorsal scales with irregular lumpy or reticulate thickening, rarely with smooth white thickening and almost closed perforations 8 8. Largest tentacle trunk ossicles predominantly plates with large irregular perforations P. whittakeri sp. nov. (South Sandwich, Bouvet, South Shetland Islands) — Largest tentacle trunk ossicles predominantly plates with large perforations centrally grading to numerous small close perforations marginally P. schnabelae sp. nov. (off Prydz Bay, MacRobertson Land) 9. Ossicles in sole numerous crosses; rare perforated plates .. P. emilyae sp. nov. (Prydz Bay) — Ossicles in sole perforated plates 10 Antarctic and Sub-Antarctic species of Psolidium Ludwig 25 10. Dorsal scales single-layered only perforated plates; perforated plates in sole lacking irregular thickenings P. gaini Vaney, 1914 (South Georgia, S Orkney Is, Weddell Sea, Antarctic Peninsula, Ross Sea, Prydz Bay) — Dorsal scales single-layered with some secondary layering; sole with irregularly thickened, scale-like perforated plates P. normani sp. nov. (Adelie Land, Prydz Bay, MacRobertson Shelf, Enderby Land) Table 1. Distribution of Psolidium Ludwig, 1886. Antarctic and Sub-Antarctic species of Psolidium species Distribution P. disciformis (Theel, 1886) Chile, Comau Fiord to Strait of Magellan; 8-448 m P. dorsipes Ludwig, 1886 S Argentina, Falkland Is, S Chile; 10-483 m P. emilyae sp. nov. E Antarctica, Prydz Bay, Four Ladies Bank; 450-556 m P gaini Vaney, 1914 South Georgia to Ross Sea to Prydz Bay; 19-646 m Weddell Sea; 260-795 m (Gutt, 1988) P. incubans Ekman, 1925 South Georgia, Cumberland Bay; 12-38 m P. normani sp. nov. Adelie Land, Prydz Bay, Enderby Land; 105-193 m P. pawsoni sp. nov. Weddell Sea, Ross Sea, Enderby Land; 137-920 m P. poriferum (Studer, 1876) Marion, Kerguelen, Heard, McDonald Is; 100-600 m Eastern Antarctica, MacRobertson Land; 177 m P. schnabelae sp. nov. MacRobertson Land, slope off Prydz Bay; 1266 m P. tenue Mortensen, 1925 Antarctic Peninsula to Ross Sea to Prydz Bay; 90-923 m New Zealand, Antipodes I.; 2010-2100 m P. whittakeri sp. nov. Bouvet, S. Sandwich, S. Shetland Is; 146-759 m Psolidium Ludwig Psolidium Ludwig, 1886: 9.— Mortensen, 1927: 413.— Deichmann, 1941: 141-43.— Deichmann, 1947: 336,-Lambert, 1996: 21. Diagnosis (see O’Loughlin and Marie, 2008). Dendrochirotid holothuroids; small, up to 40 mm long; mid-body arched dorsally in transverse section, flat ventrally; dorsal and lateral body covered with imbricating scales, usually macroscopically conspicuous, irregular in size and arrangement; scales decreasing in size ventro -laterally, orally and anally; lacking large oral valves; extensible oral cone, anterior to anterior- dorsal to dorsal orientation; extensible anal cone, posterior to posterior-dorsal to dorsal orientation; tube feet dorsally and laterally in mid-body, pass through scales. Sole distinct, oval to elongate; discrete margin created by junction of small imbricating ventro -lateral scales with thin- walled, usually calcareous, sole that lacks scales; peripheral band of tube feet, may be discontinuous across the inter-radii anteriorly and posteriorly; peripheral tube feet frequently of 2 sizes, outer series smaller; mid-ventral radial series of tube feet present or absent. Calcareous ring solid, plates sub -rectangular, radial and interradial plates with tapered anterior projections; radial plates with deep notch posteriorly, interradial plates with shallow concave indentation posteriorly; 10 dendritic tentacles, ventral 2 smaller. Dorsal and lateral ossicles: multi-layered or single-layered perforated plates (scales), always some with tube foot canals; integument covering scales may have cupped crosses, cups, “thorn” ossicles (irregular branched rods pointed distally), buttons, perforated plates and rosettes; tube foot small endplates, and tube foot support ossicles that are irregular rods and plates, bent and curved, variably perforated. Sole ossicles: inter-radii with small to large single-layered perforated plates (rarely with multi-layering), smooth to variably knobbed and thickened, sometimes with cupped crosses, cups, thorn ossicles, and rosettes; radii with additional tube foot ossicles, large endplates and tube foot support ossicles that are irregular rods and plates, bent and curved, variably perforated. Tentacle ossicles: perforated plates, rod-plates and rods, thick to thin, long to short, straight or bent, flat or curved; dendritic tentacle branch endplates are small, irregular in shape, cupped, with a few large perforations and irregular margin; densely branched rosettes may be present. Type species. Psolidium dorsipes Ludwig, 1886. Antarctic and Sub-Antarctic species. Psolidium disciformis (Theel, 1886); P. dorsipes Ludwig, 1886; P. emilyae sp. nov.; P. gaini Vaney, 1914; P. incubans Ekman, 1925; P. normani sp. nov.; P. pawsoni sp. nov.; P. poriferum (Studer, 1876); P. schnabelae sp. nov.; P. tenue Mortensen, 1925; P. whittakeri sp. nov. Psolidium disciformis (Theel) Figures la, b; 4a Psolus disciformis Theel, 1886: 85, pi. 9 fig. 6. Theelia disciformis— Ludwig, 1892: 350.— Perrier, 1902: 494. Theelia (?Psolidium) disciformis.— Ludwig, 1894: 136.— Ludwig, 1898: 52. Psolidium disciformis.— Deichmann, 1947: 337.— Pawson, 1969: 38, map 3. Material examined. Syntypes (2): Chile, Challenger stn 311, 52°45’30"S 73°46’W, 448 m, BM(NH) [18]86.10.2.86. Diagnosis. Psolidium species up to 18 mm long (25 mm in Theel, 1886); dorsal and lateral scales conspicuous, up to 1 mm wide; dorsal and lateral tube feet numerous, conspicuous, up to 2 penetrate each scale. 26 P. Mark O’Loughlin and Cynthia Ahearn Figure 1. Photos of preserved specimens of species of Psolidium Ludwig, 1886 (a-c, e, f, by Cynthia Ahearn; d, by Leon Altoff and Audrey Falconer), a, b, P. disciformis (Theel, 1886), Chile, 18 mm long, holotype BM(NH) [18]86.10.2.86: a, dorsal; b, ventral, c, P. dorsipes Ludwig, 1886, Argentina, ventral, 14 mm long, USNM 1114252. d, P. emilyae sp. nov., Prydz, Bay, dorsal (above) and lateral (oral end right), 14 mm long, holotype SAM K2221. e, P. gaini Vaney, 1914, Victoria Land, lateral, 25 mm long, USNM E40556. f, P. incubans Ekman, 1925, South Georgia, lateral, 8 mm long, BM(NH) 2008.3190. Antarctic and Sub-Antarctic species of Psolidium Ludwig 27 Figure 2. Photos of preserved specimens of species of Psolidium Ludwig, 1886 (a, b, by Leon Altoff and Audrey Falconer; c, by Cynthia Ahearn; d, by Mark Darragh). a, b, P. normani sp. nov., Prydz Bay: a, lateral (above) and ventral, 24 mm long, holotype NMV F157400; b, close-up of sole with scale-like plates (left), and of lateral scales (right), specimen 13 mm long, paratype NMV F69118. c, P. pawsoni sp. nov., Weddell Sea, dorso-lateral, 23 mm long, holotype, USNM 1112364. d, P. poriferum (Studer, 1876), Kerguelen I., dorsal, 20 mm long, holotype ZMB 2259. Sole: outer peripheral single series of smaller tube feet; inner peripheral single series of larger tube feet; zig-zag series of mid-ventral (sole) radial tube feet. Sole ossicles: numerous irregularly oval to round, disc-like smooth plates, variably with few small perforations, up to 220 pm long; rare shallow cupped crosses and more numerous 4-holed regular shallow cups with blunt marginal spines, cups 150 pm long. Colour (live). White (P. Lambert, pers. comm.). Distribution. SE Pacific Ocean, Chile, Comau Fiord to Strait of Magellan (42° to 56°S), on rock; 8-448 m (P. Lambert, pers. comm.). Remarks. Theel (1886) noted the presence of dorsal and lateral tube feet in his new species, but Ludwig (1886) erected the genus Psolidium for Psolidae with this diagnostic character in the same year. Deichmann (1947) assigned Psolus disciformis Theel, 1886, to Psolidium Ludwig, 1886. The form of the ossicles in the sole is diagnostically distinctive, and ossicles from dorsum and tentacles were not examined. Recent intensive field work on the Chilean coast under the auspice of the San Ignacio del Huinay Foundation has discovered an abundance of P. disciformis on the rocks in the coastal shallow from 8 to 27 m (P. Lambert, pers. comm.). Psolidium dorsipes Ludwig Figures lc; 4b Psolidium dorsipes Ludwig, 1886: 9-10, pi. 2, fig. 9.— Ludwig, 1892: 349.— Ludwig, 1894: 135 (passim).— Perrier, 1904: 16.— Perrier, 1905: 48, pi. 2 figs 5-15, text figs G-H.-Ekman, 1925: 112-13, text fig. 26.— Deichmann, 1941: 1 42-44, pi. 29 figs 1-12.— Deichmann, 1947: 28 P. Mark O’Loughlin and Cynthia Ahearn Figure 3. Photos of preserved specimens of species of Psolidium Ludwig, 1886 (a, b, by Cynthia Ahearn; c-f, by Leon Altoff and Audrey Falconer), a, P. schnabelae sp. nov., Prydz Bay, dorso-lateral (oral end right), 26 mm long, holotype SAM K2345. b-d, P. whittakeri sp. nov.: b. South Sandwich Is, ventral, 28 mm long, holotype USNM 1112365; c. Deception I., lateral, 28 mm long, NMV F104834; d, tentacles, specimen 28 mm long, NMV F104834. e, f, P. tenue Mortensen, 1925, Prydz Bay, 28 mm long (two ventral small, bottom), NMV F107416; e, lateral; f, sole. Antarctic and Sub-Antarctic species of Psolidium Ludwig 29 Figure 4. Photos of ossicles of species of Psolidium Ludwig, 1886 (a, b, f, by Cynthia Ahearn; c-e, by Chris Rowley), a, P. disciformis (Theel, 1886), shallow cup (bottom centre left, 150 pm long) and disc-like plates from sole, holotype BM(NH) [18]86.10.2.86. b, P. dorsipes Ludwig, 1886, button (left, 90 pm long) and cup from sole, USNM 1114251. c-e, P. emilyae sp. nov., holotype SAM K2221: c, dorsal scale; d, dorsal tube foot support plates; e, crosses and plates from sole, f, P. incubans Ekman, 1925, deep goblet-like dorsal cups, 70-90 pm wide, 80-110 pm high, BM(NH) 2008.3190. 30 P. Mark O’Loughlin and Cynthia Ahearn 336-37.— Hickman, 1962: 60.— Pawson, 1969: 38, map 3.— Pawson and Valentine, 1981: 453.— Lambert, 1996: 28-30, table 1. Material examined. South Atlantic Ocean, Argentina, Santa Cruz, E of Grande Bay, 92 m, USNM E2177 (2 specimens); E of Cabo Virgenes, 101 m,USNM 1114251 (5); near Strait of Magellan, 66 m, 1 1 14252 (l);Tierra del Fuego, Staten I., E40823 (2); Cape Penas, 81 m, E40824 (2); R/V William Scoresby stn 834, Cape Virgenes, 0-38 m, BM(NH) 2008.3182 (1); Scotia Sea, E of Burdwood Bank, 419^-83 m, E40825 (1). South Pacific Ocean, Chile, Strait of Magellan, 36 m, E2178 (4); Cape Froward, 82 m, USNM 1081593 (1). Diagnosis. Psolidium species up to 17 mm long; preserved form variably with low to high profile, short to elongate oval sole; dorsal and lateral scales conspicuous, smooth, up to 1.6 mm wide; dorsal and lateral tube feet conspicuous, numerous. Sole: outer peripheral single series of smaller tube feet; inner peripheral single to double series of larger tube feet; mid-ventral (sole) radial single to zig-zag to double series of tube feet. Dorsal ossicles: multi-layered plates (scales) thick, 0-2 canals and 0-2 marginal indentations for tube feet per scale; intergrade with large thick knobbed plates with developing secondary layers; perforated plates with thickenings (buttons), 100-150 pm long, 3 to many perforations; thick smooth perforated plates, 150-200 pm long; numerous shallow cups, oval, marginally with round knobs, biggest cups knobbed on cross and rim, 2 large central and 2 smaller end perforations, cups 40-60 pm long. Sole ossicles: shallow cups, oval, 2 large central and 2 smaller end perforations, some imperfectly formed, variably finely knobbed on margin to thickly knobbed on margin and cross, cups 40-70 pm long; thickly knobbed, regular, 4-holed buttons, 80-100 pm long, some irregular, up to 150 pm long; shallow concave perforated plates variably knobbed on surface and margin, up to 16 perforations, up to 200 pm long. Distribution. Argentina (E of Grande Bay), Burdwood Bank, Falkland Is; Chile, Tierra del Fuego, Strait of Magellan; 10-483 m. Remarks. One Psolidium dorsipes specimen (USNM 1114251) has shallow cup-shaped indentations covering the sole that appear to indicate brood-protection under the body. Psolidium emilyae sp. nov. Figures Id; 4c-e Material examined. Holotype: Eastern Antarctica, Prydz Bay, Four Ladies Bank, ANARE 1991, Aurora Australis stn 25B, 68°31’S 77°27’E, 450-556 m, K. L. Gowlett-Holmes and W. Zeidler, 3 Feb 1991, SAM K2221. Diagnosis. Psolidium species 14 mm long (preserved); elongate tubular body, transversely rounded form, sole narrower than body width; dorsal and lateral tube feet conspicuous, numerous, cover body closely; dorsal and lateral scales inconspicuous. Sole: peripheral single series of tube feet; some slightly smaller outer tube feet in an incomplete series (partly concealed by projecting ventro -lateral scales); lacking mid-ventral (sole) radial series of tube feet. Dorsal ossicles: thick, smooth, single-layered perforated plates (scales), lacking secondary developments, small perforations, tube foot canals, scales up to 0.7 mm wide; irregular, curved tube foot support ossicles, up to 4 perforations, up 128 pm long. Sole ossicles: numerous crosses, predominantly regular, elongate, 4-armed, thick, with distal pairs of swellings on arms, branches rarely joined to create perforations, up to 144 pm long; few thick perforated plates, up to 10 perforations, margin variably bluntly toothed to slightly knobbed, up to 224 pm long. Colour ( preserved). Off-white. Distribution. Eastern Antarctica, Prydz Bay, Four Ladies Bank; 450-556 m. Etymology. Named for Emily Whitfield (Marine Research Group of the Field Naturalists Club of Victoria) who first noticed the distinctive cross ossicles in the sole of this species, and in appreciation of her research assistance in Museum Victoria. Remarks. The cross ossicles in the sole of Psolidium emilyae sp. nov. are diagnostically distinctive among Antarctic and Sub-Antarctic Psolidium species, and tentacle ossicles from the unique small holotype specimen were not examined. P. emilyae is similar morphologically to Psoldium normani sp. nov. from Prydz Bay (see below), but the depth of occurrence of P. normani (105-193 m) is shallower than P. emilyae (450-556 m). A single specimen from Prydz Bay (98-301 m) of the morphologically similar Psolidium gaini Vaney, 1914, also occurred at a shallower depth than P. emilyae. Ossicles from the sole of specimens of P. gaini and P. normani that were similar in size to the small type specimen of P. emilyae were examined, and found to be similar to those in larger specimens of the two species, and not the crosses typical of P. emilyae. Psolidium gaini Vaney Figures le; 5a-c; 6a, b Psolidium gaini Vaney, 1914: 18-20, pi. 1 figs 7-9, pi. 4 figs 6-14.— Ekman, 1925: 5, 117-19, text-fig. 28,-Ekman, 1927: 414-15.— Grieg, 1929a: 13.— Grieg, 1929b: 9.— Pawson, 1969: 38, map 3.— Arnaud 1974: 651,-Cherbonnier, 1974: 608,-Gutt, 1988: 23, 28, 30, 32, 65, 73, 77,-Gutt, 1991a: 147, 149, 152, 153,-Gutt, 1991b: 324. Psolidium sp. MoV 2081.-O’Loughlin et al„ 1994: 552, 554. Material examined. Antarctic Ocean, South Orkney Is, 298-302 m, USNM E40558 (1 specimen); South Shetland Is, Aspland I., 163-180 m, A MLR- 03 stn 90, 20 Mar 2003, NMV F104812 (2); Antarctic Peninsula, Joinville I., 210-220 m, USNM E40559 (10); Graham Land, 91 m, E40561 (2); Palmer Archipelago, 85-130 m, E40552 (2); 102 m, E40553 (1); 38-70 m, E40554 (2); 70-150 m, E40555 (2); 55 m, E40562 (1). Ross Sea, Terra Nova stn 340, 76°56’S 164°12’E, 293 m, BM(NH) 1932.8.11.253 (2); NZOI Endeavour stns A461, 0-550 m, NIWA 43882 (1); A534, 366 m, NIWA 43884 (2); A537, 546 m, NIWA 43885 (1); E209B, 163 m, NIWA 43886 (2); Discovery stn 1660, Pennell Bank, 0-351 m, BM(NH) 2008.3183-3189 (7); Tangaroa stn 0802/100, 451-447 m, NIWA 45696 (4). Antarctic and Sub-Antarctic species of Psolidium Ludwig 31 Figure 5. Photos of ossicles of species of Psolidium Ludwig, 1886 (a, SEM by Cynthia Ahearn; b-f, by Chris Rowley), a-c, P. gaini Vaney, 1914: a, dorsal scale and tube foot support plate (bottom left), USNM E40551; b, dorsal tube foot support plates, NMV F68112; c, plates from sole, NMV F68112. d-f, P. normani sp. nov.: d, dorsal scale, paratype SAM K2341; e, dorsal tube foot support plates, paratype NMV F68662; f, plate from sole, paratype SAM K2341. 32 P. Mark O’Loughlin and Cynthia Ahearn Figure 6. Photos of ossicles of species of Psolidium Ludwig, 1886 (a-c, SEM by Cynthia Ahearn; d, e, by Chris Rowley; f, by Cynthia Ahearn). a-b, P. gaini Vaney, 1914, USNM E40551: a, plate from sole; b, tentacle ossicle, c-f, P. tenue Mortensen, 1925: c, dorsal scale, USNM E40783; d, dorsal scale, with tube foot endplate in tube foot canal, and absence of tube foot support plates, NMV F107442; e, tentacle rod-plates, NMV F68113; f, tentacle rod-plates, largest 780 fim long. Antarctic and Sub-Antarctic species of Psolidium Ludwig 33 Victoria Land, 640-646 m, USNM E40556 (2); 598-613 m, E40560 (1); 573-576 m, USNM 1082084 (1); 598-613 m, USNM 1112624 (1); Balleny Is, 55-146 m, E40551 (19); 150-157 m, E40557 (1); Tangaroa stn 0602, 140-150 m, NIWA 44712 (1). Adelie Land, BANZARE stn 90, 66°21’S 138°28’E, 640 m, SAM K2340 (1). Prydz Bay, Four Ladies Bank, 298-301 m, NMV F68112 (1). Diagnosis. Psolidium species up to 27 mm long (up to 40 mm in Vaney, 1914); body elongate, transversely rounded form (preserved), sole narrower than body width; dorsal and lateral tube feet conspicuous, numerous, cover body closely; dorsal and lateral scales inconspicuous. Sole: outer peripheral single series of smaller tube feet; inner peripheral single series of larger tube feet; lacking mid- ventral (sole) radial series of tube feet. Dorsal ossicles: thick, smooth to irregularly thickened, single-layered, perforated plates (scales), up to 950 p m long, perforations small, up to 3 canals or marginal indentations for tube feet per scale; irregular, asymmetrical, curved tube foot support plates, 4-8 perforations, digitiform to bluntly spinous (South Shetland Is) to pointed spines (Prydz Bay) on one margin, opposite margin lacking projections, plates up to 176 pm long. Sole ossicles: thick, regular, 4-holed buttons, variably knobbed to bluntly spinous marginally, 160-190 pm long; intergrade with thick, smooth, perforated plates, sometimes slightly concave, sometimes slightly knobbed marginally, sometimes margin upturned, number of perforations variable, size of plates variable; plates up to 14 perforations, up to 184 pm long, many knobbed marginally (South Shetland Is); plates up to 19 perforations, up to 255 /mi long (Ross Sea); plates up to 17 perforations, up to 224 (rarely 272) p m long, slight swellings marginally (Prydz Bay). Distribution. South Georgia, South Orkney Is, South Shetland Is, Weddell Sea, Antarctic Peninsula to Ross Sea to Prydz Bay; 19-795 m. Remarks. Vaney (1914) reported the sizes of the two type specimens as 40 and 30 mm long. It is incongruous that none of the numerous specimens seen in this study is longer than 26 mm. Ekman (1925) noted some differences between the specimen that he examined and the description by Vaney (1914), and made a point that what Vaney described as “cups” dorsally, were for him tube foot support ossicles. We agree with Ekman (1925) that there are numerous tube foot support ossicles dorsally, that have the form of irregular curved perforated plates that are superficially cup-like. The dorsal and lateral tube feet in Psolidum gaini, P. emilyae sp. nov. (above) and P. normani sp. nov. (below) are conspicuous because of the presence of support ossicles, whereas the dorsal and lateral tube feet of Psolidium poriferum (below), P. pawsoni sp. nov. (below), P. schnabelae sp. nov. (below), P. tenue (below) and P. whittakeri sp. nov. (below) have at most rare tube foot support ossicles and are inconspicuous. We noted plates from the sole of Antarctic Peninsula specimens (type region) up to 200 pm long with up to 13 perforations. Sole plates from Adelie Land specimens were up to 300 pm long with up to 23 perforations. Other variations are noted in the diagnosis above. The variations are significant, and suggest that there may be more than one species. Grieg (1929a, b) reported Psolidium gaini from South Georgia (55 m) and Graham Land (60-90 m), and noted reports of P. gaini from the South Shetland Is (19 m) and Emperor William Land (360 m). Gutt (1988) reported P. gaini from the Weddell Sea (260-795 m). Psolidium incubans Ekman, 1925 Figures If; 4f Psolidium incubans Ekman, 1925: 5, 113-116, text fig. 27.— Arnaud, 1974: 584. -Gutt, 1988: 73. Material examined. South Georgia, Discovery stn MS67, E Cumberland Bay, 38 m, 28 Feb 1926, BM(NH) 2008.3190 (1 specimen). Diagnosis. Psolidium species up to 12 mm long; wrinkled body surface, created by high cups; dorsal and lateral scales not conspicuous; dorsal and lateral tube feet conspicuous. Sole: outer peripheral single series of smaller tube feet; inner peripheral single to zig-zag to double series of larger tube feet; lacking mid-ventral (sole) radial series of tube feet; sole soft, with folds for brood-protection. Dorsal ossicles: perforated plates with anastomosing secondary developments and multi -layering (scales), up to 700 pm long, 0-2 tube foot canals per scale; shallow cups with 2 large central and 2 smaller end perforations, marginal knobs, cups 90 pm long; deep goblet-like cups closed across rim, vertical blunt spines above rim, cups 70-90 pm wide, 80-110 pm high. Sole ossicles: shallow cupped crosses, shallow cups with small marginal knobs, cups 80 pm long; shallow concave perforated plates, upturned marginal knobs, up to 14 perforations, plates up to 150 pm long. Distribution. South Georgia, Cumberland Bay; 12-38 m. Remarks. As noted by Ekman (1925) this small species of Psolidium brood-protects in folds of the sole. Psolidium normani sp. nov. Figures 2a, b; 5d, f Psolidium cf. gaini.— O’Loughlin et al., 1994: 552, 554 (non Psolidium gaini Vaney, 1914). Material examined. Holotype: eastern Antarctica, MacRobertson Shelf, ANARE 1993, Aurora Australis stn 127, 67°16’S 65°26’E, 109-121 m, M. O’Loughlin, 12 Feb 1993, NMV F157400. Paratypes: Type locality and date, F68661 (1 specimen); F69118 (1); Prydz Bay, Fram Bank, stn 130, 67°32’S 69°02’E, 105-114 m, F68662 (1); ANARE 1991, Aurora Australis stn 100, 67°28’S 68°50’E, 145-150 m, C. C. Lu andT. N. Stranks, 28 Feb 1991, F68111 (1); K. L. Gowlett-Holmes and W. Zeidler, SAM K2220 (1); Enderby Land, BANZARE stn 41, 65°48’S 53°16’E, 193 m, SAM K2341 (2); Adelie Land, BANZARE stn 90, 66°21’S 138°28’E, 640 m (possibly “much shallower”, according to BANZARE records), SAM K2349 (1). Diagnosis. Psolidium species up to 30 mm long; body elongate, transversely rounded form (preserved), sole narrower than 34 P. Mark O’Loughlin and Cynthia Ahearn body width; dorsal and lateral tube feet conspicuous, numerous, cover body closely; dorsal and lateral scales inconspicuous. Sole: outer peripheral single series of smaller tube feet; inner peripheral single series of larger tube feet; lacking mid- ventral (sole) radial series of tube feet. Dorsal ossicles: predominantly single-layered perforated plates (scales), some secondary layering; up to 4 tube foot canals or marginal indentations per plate; scales up to 800 pm wide; numerous tube foot support plates, curved, spinous on one side (rarely both sides), marginal spines sometimes bifurcate, up to 19 perforations, up to 136 pm long; tube foot ‘endplates’ small irregular mesh-like, up to 48 pm wide, few perforations, irregular marginal projections, not regularly perforated plates. Sole ossicles: scale-like perforated plates, intergrading with buttons with 4 perforations; plates bluntly to sharply spinous to knobbed on one side, projections sometimes bifurcate; pronounced surface thickenings with ridges or lumps; up to 48 perforations per plate; plates up to 352 pm long. Distribution. Eastern Antarctica, Adelie Land; Prydz Bay, Fram Bank; MacRobertson Shelf; Enderby Land; 105-193 m (? 640 m). Etymology. Named for Mark Norman (Senior Curator, Marine Biology Section, Museum Victoria), with admiration of his enthusiastic engagement with marine invertebrate studies, with gratitude for his personal support, and in recognition of his contribution to collecting from Prydz Bay and Heard I. Remarks. Psolidium normani sp. nov. is similar externally to P. emilyae sp. nov. (above) and P. gaini Vaney, 1914, but is distinguished by the scale-like plates in the sole and secondary thickening of the dorsal scales. Psolidium pawsoni sp. nov. Figures 2c; 7a, b Psolidium incertum — Gutt, 1988: 1, 3, 23, 27-77.— Gutt, 1991a: 147-153.— Gutt, 1991b: 315, 320, 324-25 (non Psolidium incertum (Theel, 1886) = Psolidium poriferum (Studer, 1876) below). Material examined. Holotype: Weddell Sea, Coats Land, Caird Coast, off Brunt Ice Shelf, USARP, R/V Glacier , cr 2 stn 1, 74°28’06"S 30°31’42"W, 513 m, 24 Feb 1969, USNM 1112364. Paratypes: type locality and date, USNM E40798 (4); type locality and date, NMV F157430 (2). Other material: Weddell Sea, 430 m, USNM E33925 (1). Ross Sea, Tangaroa 0802, 67-69°S 170°E-178°W, 329-334 m, NIWA 38273 (1); 329-334 m, NIWA 38281 (1); 137-150 m, NIWA 38604 (1); 300-340 m, NIWA 38624 (6); 547-605, NIWA 39084 (12); 565-920 m, NIWA 39181 (2); 479-480 m, NIWA 39311 (6); 456-540 m, NIWA 45695 (8); 447-455 m, NIWA 45699 (1); Tangaroa 0602, stn 394, 67°S 179°E, 540-600, NIWA 44667 (5). Enderby Land, BANZARE stn 34, 66°21’S 58°50’E, 603 m, SAM K2342 (2). Diagnosis. Psolidium species up to 28 mm long; body form elongate, narrow, high; dorsal and lateral scales conspicuous, thin, smooth, up to 2.0 mm wide; dorsal and lateral tube feet inconspicuous. Sole: outer peripheral single series of smaller tube feet; inner peripheral single series of larger tube feet; lacking mid- ventral (sole) radial series of tube feet. Dorsal ossicles: multi-layered perforated plates (scales), single-layered marginally; plates frequently with smooth white thickening and reduced/small perforations aligned perpendicular to margin in parallel series; plates with reticulate thickening and multi-layered centrally, lacking frequent radiating linear thickenings between marginal perforations; up to 4 tube foot canals per plate; dorsal and lateral tube feet lacking support plates. Sole ossicles: throughout most of sole rare, small, smooth, thin, irregularly shaped rods and perforated plates, sometimes as regular 4-holed plates with marginal knobs or digitiform projections, sometimes surface knobs, up to 200 pm long; near margin of sole and peripheral tube feet thicker, irregular, elongate, perforated rod-plates, surface with pronounced linear thickenings (suggesting branched rod origin), up to 420 pm long, intergrading with oval to elongate thick perforated plates, slightly concave, some with secondary layering, up to 320 pm long. Tentacles: largest tentacle trunk ossicles irregular round to oval to elongate, smooth perforated plates, some thin with large perforations, some thicker with smaller perforations, plates up to 550 pm long, rarely with any secondary layering; few narrow perforated plates, lacking evidence of rod derivation, up to 650 pm long. Colour (preserved). Body pale brown to off-white; scales with slightly discontinuous haloes; fine red-brown microscopic flecking over dorsal body. Distribution. Weddell Sea, 260-795 m (Gutt, 1991b); Ross Sea, 137-920 m; Enderby Land, 603 m. Etymology. Named for Dr David Pawson (USNM, Senior Research Scientist), with gratitude for his provision of the opportunity to pursue this research, and appreciation of his constant interest and encouragement. Remarks. Massin (1992) examined specimens of Psolidium poriferum (as P. incertum) from both the Sub-Antarctic islands of the Indian Ocean and the Weddell Sea, and observed that there were significant differences in the structure of the dorsal scales. He judged that the Weddell Sea material “probably represents another species”. We agree, and refer the Weddell Sea, Ross Sea and Enderby Land material to Psolidium pawsoni sp. nov. Gutt (1988) reported Psolidium incertum (Theel, 1886) from the Weddell Sea, and synonymised Psolidium tenue Mortensen, 1925, with P. incertum. We reject a synonymy of P. incertum with P. tenue (see our treatments of P. poriferum and P. tenue). We assume that the Weddell Sea material seen by Gutt is synonymous with our Psolidium pawsoni sp. nov. from the Weddell Sea, but acknowledge that Psolidium tenue might also occur in the Weddell Sea. Psolidium pawsoni sp. nov. is distinguished diagnostically by the frequent occurrence on the dorsal and lateral scales of smooth white thickening with consequent reduced/small perforations. Antarctic and Sub-Antarctic species of Psolidium Ludwig 35 Psolidium poriferum (Studer) syn. nov. Figures 2d; 7c-f Cuvieria porifera Studer, 1876: 452-53.— Studer, 1879: 123. P solus poriferus.— Bell, 1882: 644.— Lampert, 1889: 854.— Studer, 1889: 163. Psolus poriferus (Cuvieria). —Theel, 1886: 130.— Lampert, 1885: 122. Psolus incertus Theel, 1886: 86-87, pi. 8 fig. 4, pi. 6 fig. 5.— Studer, 1889: 163. (new synonymy). Theelia incerta.— Ludwig, 1892: 350. Theelia (? Psolidium) incerta.— Ludwig, 1894: 136.— Ludwig, 1898: 52. Theelia porifera (part).— Ludwig, 1898: 52.— Pawson, 1969: 38. Psolidium incertum.— Pawson, 1969: 38, map 3.— Pawson, 1971: 34, 37.— Guille, 1982: 70,-Massin, 1992: 317, 320-21, figs 8-9.— Branch, Jangoux, Alva, Massin and Stampanato, 1993: 56, 61-65.— Thandar, 1999: 368,-Thandar, 2006: 38. Psolidium poriferum.— Pawson, 1971: 34, 37.— O’Loughlin and Marie, 2008: 3 Material examined. Psolidium poriferum (Studer, 1876) holotype: Kerguelen, Royal Sound, Gazelle , 119 m, ZMB 2259. Psolidium incertum (Theel, 1886) syntypes (3; very poor condition): Kerguelen, Royal Sound, Challenger stn 149c, BM(NH) [18]86.10.2.89 (1); Heard I., stn 150, BM(NH) [18]86.2.88 (1); stn 151, BM(NH) [18]86. 10.2.90 (1). Other material: Indian Ocean, McDonald Is, 210-234 m, USNM 1081576 (2 specimens); S of Kerguelen I., BANZARE stn 47, 150 m, SAM K2344 (23); Heard I., ANARE Aurora Australis AA90, 200 m, SAM K2189 (1); 260-380 m, K2190 (1); 260 m, K2191 (1); 266-380 m, K2205 (1); ANARE Aurora Australis AA92, 185-204 m, NMV F66849 (1); 229 m, F66850 (1); Heard I., F/V Southern Champion , cr SC26, Apr/ May 2003, Western Plateau, 26A406 m, TMAG H3303 (1), H3306 (2), H3308 (2), H3317 (1), H3320 (4); Aurora Bank, 222-292 m, H3304 (4), H3305 (3), H3307 (4), H3309 (2), H3313 (2), H3314 (2), H3315 (1), H3318 (3), H3321 (6), H3322 (1); Coral Bank, 293-297 m, H3310 (3), H3311 (4); Shell Bank, 259-329 m, H3312 (1), H3316 (3), H3319 (2). Eastern Antarctica, MacRobertson Land, BANZARE stn 107, 66°45’S 62°03’E, 177 m, SAM K2343 (1). Diagnosis. Psolidium species up to 33 mm long; body form elongate, narrow, high; dorsal and lateral scales conspicuous, thin, smooth, up to 1.8 mm wide; dorsal and lateral tube feet inconspicuous. Sole: outer peripheral single series of smaller tube feet; inner peripheral single series of larger tube feet; lacking mid- ventral (sole) radial series of tube feet. Dorsal ossicles: multi-layered perforated plates (scales), single-layered marginally, frequently with well developed radiating-to-margin linear rounded thickenings between series of perforations, reticulate thickenings towards centre of plates, multi-layered centrally; up to 3 tube foot canals per plate. Sole ossicles: throughout most of sole rare, small, smooth, thin, irregularly shaped rods and perforated plates, 0-10 perforations, bluntly spinous around margin, sometimes surface knobs, up to 220 pm long; sometimes as regular 4-holed plates with marginal knobs, sometimes surface knobs, up to 160 pm long; near margin of sole and peripheral tube feet some thicker, irregular perforated rod-plates, surface with linear thickenings and large knobs, rod-plates up to 370 pm long, intergrading with oval to elongate perforated plates with secondary layering, up to 400 pm long. Tentacles: largest tentacle trunk ossicles irregular round to triangular to oval, thick perforated plates, irregular large perforations, some small plates with secondary layering, plates up to 650 pm long; some plates with irregular linear thickenings between perforations (suggesting rod origin), some rod-plates up to 800 pm long. Colour (preserved). Body off-white; scales not haloed. Distribution. Marion, Prince Edward, Kerguelen, Heard and McDonald Is, 100-600 m (Massin, 1992), 150-406 m (this work); Eastern Antarctica, MacRobertson Land, 177 m. Remarks. Pawson (1971) indicated that the evidence was “quite strong” for a synonymy of P. incertum with P. poriferum, and O’Loughlin and Marie (2008, this volume) inferred this synonymy. Both species were described from the same type locality. Based on an examination of the type materials and the literature, Psolidium incertum (Theel, 1886) is synonymised here with Psolidium poriferum (Studer, 1876). Psolidium poriferum occurs abundantly on the Sub -Antarctic Islands of the Indian Ocean, with the exception of a single BANZARE specimen found on the MacRobertson Shelf on the Antarctic coast. Psolidium poriferum (Studer) is distinguished diagnostically by the frequent occurrence on the scales of well developed radiating-to-margin linear rounded thickenings between series of perforations. Psolidium schnabelae sp. nov. Figures 3a; 8a-c Material examined. Holotype: Eastern Antarctica, MacRobertson Land, slope off Prydz Bay, BANZARE stn 29, 66°28’S 72°41’E, 1266 m, 25 Dec 1929, SAM K2345. Other material: type locality and date, SAM K2346 (2). Diagnosis. Psolidium species up to 25 mm long; body form elongate, high; dorsal and lateral scales conspicuous, thin, smooth, up to 2.5 mm wide; dorsal and lateral tube feet inconspicuous. Sole: outer peripheral single series of smaller tube feet; inner peripheral single series of larger tube feet; lacking mid- ventral (sole) radial series of tube feet. Dorsal ossicles: multi-layered perforated plates (scales), single-layered marginally, reticulate thickenings extend from margin towards centre of plate, multi-layered centrally, lacking frequent radiating linear thickenings between marginal perforations, lacking smooth white thickening with small perforations, margin with irregular thickenings and perforations, not smooth; up to 4 tube foot canals or marginal indentations per plate; small mesh-like tube foot endplates, about 5 perforations, 40 pm wide; lacking dorsal and lateral tube foot support plates. Sole ossicles: throughout sole rare, small, smooth, thin, irregularly shaped rods and perforated plates, up to 10 perforations, up to 240 pm long, sometimes as regular 4-holed plates with bluntly spined margin, sometimes surface knobs, up to 180 pm long; near margin of sole and peripheral tube feet thicker irregular rod-plates and very thick irregular perforated plates, up to 300 pm long. 36 P. Mark O’Loughlin and Cynthia Ahearn Figure 7. Photos of ossicles of species of Psolidium Ludwig, 1886 (a, d, SEM by Cynthia Ahearn; b, e, f, by Cynthia Ahearn; c, by Chris Rowley), a, b, P. pawsoni sp. nov., paratype USNM E40798: a, dorsal scales; b, tentacle ossicles, larger 520 pm long, c-f, P. poriferum (Studer, 1876): c, dorsal scale, holotype ZMB 2259; d, dorsal scale, SAM K2344; e, dorsal scale, SAM K2344; f, tentacle ossicle, 640 pm long, SAM K2344. Antarctic and Sub-Antarctic species of Psolidium Ludwig 37 Figure 8. Photos of ossicles of species of Psolidium Ludwig, 1886 (a-d, f, by Chris Rowley; e, SEM by Cynthia Ahearn;). a-c, P. schnabelae sp. nov.: a, dorsal scale, holotype SAM K2345; b, tentacle ossicles, holotype SAM K2345; c, plate from sole, paratype, SAM K2346. d-f, P. whittakeri sp. nov.: d, dorsal scale, NMV F104834; e, dorsal scale, paratype USNM E40795; f, tentacle ossicles, NMV F104834. 38 P. Mark O’Loughlin and Cynthia Ahearn Tentacles: largest tentacle trunk ossicles predominantly smooth perforated plates, irregularly oval, large central perforations grading to numerous small close ones near margin, finely denticulate margin, plates up to 520 pm long; some thicker plates with large perforations, plates up to 300 p m long; few rod-plates, up to 300 pm long. Colour ( preserved ). Body pale brown to off-white; scales haloed, with pale outer edge (single-layer perforated edge of scales with white thickening). Distribution. Eastern Antarctica, MacRobertson Land, slope off Prydz Bay; 1266 m. Etymology. Named for Kareen Schnabel (NIWA), with gratitude for her generous and gracious assistance with loan material and data from NIWA. Remarks. The original BANZARE lot from station 29 comprises three specimens. We are not confident that the two smaller specimens are conspecific with the holotype, and they are assigned to “Other material” with reservation. We note that this species is recorded for a significantly greater depth than the other species of the Psolidium poriferum group. Psolidium schnabelae sp. nov. is distinguished diagnostically by the predominant form of the largest tentacle ossicles with large central perforations grading to small peripheral ones. Psolidium tenue Mortensen Figures 3e, f; 6c-f Psolidium tenue Mortensen, 1925: 8-9, text figs 4-5. Psolidium incertum.— O’Loughlin et al., 1994: 552, 554 (non Psolidium incertum (Theel, 1886) = Psolidium poriferum (Studer, 1876)). Material examined. Antarctic Peninsula, 90 m, USNM E40789 (1 specimen); 265 m, E40790 (1); 100-200 m, E40791 (2); 270 m, E40792 (6); 330 m, E40793 (2); 212 m, E40794 (1); 311 m, E40796 (1); 325 m, E40801 (1); 670 m, USNM 1005859 (2); Discovery stns 181, Palmer Archipelago, 160-335 m, BM(NH) 2008.3191^ (1); 182, 278-500 m, BM(NH) 2008.3192 (1); 187, 0-195 m, BM(NH) 2008.3193 (1); 190, 0-250 m, BM(NH) 2008.3194-3196 (3); 599, W of Adelaide I., 0-150 m, BM(NH) 2008.3197-3198 (2). Ross Sea, 923 m, USNM E40776 (1); NIWA Tangaroa 0802, 67-74°S 167°E-179°W, 420-866 m, NIWA 36003 (1); 36027 (20); 38102 (1); 38545 (1); 38815 (4); 38842 (1); 38872 (1); 38884 (18); 38956 (9); 45698 (3); 45727 (2); 45728 (1); Discovery stns 1644, 626 m, BM(NH) 2008.3199 (1); 1649, 695 m, BM(NH) 2008.3200-3201 (2); 1652, 0-500 m, BM(NH) 2008.3202-3205 (4); 1660, 0-351 m, BM(NH) 2008.3206-3208 (3); 2200, Balleny Is, 512-532 m, BM(NH) 2008.3209-3218 (11); NZOI Endeavour stns A461, 0-550 m, NIWA 43870 (many specimens); A466, 555 m, NIWA 43871 (many); A521, 569 m, NIWA 43872 (5); A531B, 348 m, NIWA 44705 (1); A537, 546 m, NIWA 43873 (4); D216, 463 m, NIWA 43874 (4); E199, 278 m, NIWA 43875 (2); E200, 646 m, NIWA 43876 (1); E203, 187 m, NIWA 43877 (4); E220A, 371 m, NIWA 43883 (many); E220B, 371 m, NIWA 43878 (many); E212B, 91 m, NIWA 43879 (1). Wilkes Land, 183 m, USNM E40799 (1); BANZARE stn 97, 474 m, SAM K2347 (1). Prydz Bay, 320-768 m, NMV F68113 (2); F68116 (1); F68117 (2); FI 0741 5 (1); F107416 (1); F105417 (1); F107437 (1); F107438 (1); FI 07442 (1); F107443 (1); F107447 (1); SAM K2219 (1); BANZARE stn 103, SAM K2348 (4). New Zealand, Antipodes I., 2100 m, USNM 1004885 (1). Diagnosis. Psolidium species up to 50 mm long; body form elongate, narrow, high; dorsal and lateral scales conspicuous, thin, smooth, up to 3.0 mm wide; dorsal and lateral tube feet inconspicuous. Sole: outer peripheral single series of smaller tube feet; inner peripheral single series of larger tube feet; lacking mid- ventral radial series of tube feet. Dorsal ossicles: multi-layered perforated plates (scales), single-layered marginally, reticulate thickenings from near margin towards centre, multi-layered centrally; rarely any development of smooth white thickening with smaller perforations; rarely any development of radiating-to-margin thickenings; up to 3 tube foot canals per plate; rare tube foot support plates; endplates small, mesh-like. Sole ossicles: throughout most of sole rare, small, smooth, thin, irregularly shaped perforated plates, 0-8 large perforations, up to 230 pm long; sometimes as regular 4-holed plates with slightly knobbed margin, sometimes with surface knobs, plates 100-200 pm long; near margin of sole and peripheral tube feet some thicker irregular perforated rod- plates, up to 570 pm long, intergrading with irregularly oval thick perforated plates, with secondary layering, up to 480 pm long. Tentacles: largest tentacle trunk ossicles predominantly long, narrow, thick, rod-plates (rods joined to create perforations) and perforated plates with prominent rod thickenings; not predominantly smooth perforated plates; rod- plates variably straight or bent or curved, up to 800 pm long. Colour (preserved). Body variably dark brown to pale brown to off-white; coarse red-brown flecking dorso-laterally; scales with a light-coloured “haloed” marginal appearance. Distribution. Antarctic Peninsula, Ross Sea, Wilkes Land, Prydz Bay, 90-923 m; New Zealand, Antipodes I., 2010-2100 m. Remarks. We raise Psolidium tenue Mortensen, 1925, out of a synonymy by Gutt (1988) with Psolidium incertum (Theel, 1886), synonymised above with Psolidium poriferum (Studer, 1876). P. poriferum is an almost exclusively Sub-Antarctic species. The single type specimen for Psolidium tenue Mortensen, 1925, was taken north of Discovery Inlet in the Ross Sea from 640 m. There are no inconsistencies between the original description and the material studied here from the Ross Sea. The length (preserved length of 30 mm), paucity of ossicles in the sole, single small knob sometimes present on the ossicles in the sole, and size of ossicles in the sole are all consistent with the Ross Sea material. Psolidium tenue Mortensen, 1925, is distinguished diagnostically by the rod- plate form of the largest tentacle ossicles. Psolidium whittakeri sp. nov. Figures 3b-d; 8d-f Psolidium incertum.— Ludwig and Heding, 1935: 162-64, text figs 28-29 (non Psolidium incertum (Theel, 1886) = P. poriferum (Studer, 1876) (above). Antarctic and Sub-Antarctic species of Psolidium Ludwig 39 Material examined. Holotype: Scotia Sea, South Sandwich Is, Saunders L, 57°39’24"S 26°26’42"W, 415-613 m, R/V Islas Orcadas, USARP cr 575 stn 52, 26 May 1975, USNM 1112365. Paratypes: type locality and date, USNM E40795 (39 specimens); NMV FI 57431 (6). Other material: South Sandwich Is, 146-527 m, USNM E40817 (1); 355-468 m, E40818 (1); 302-375 m, E40819 (2); 360-486 m, E40827 (2); 452-759 m, E40828 (1). Bouvet I., Icefish 2004 stn 59, 408-656 m, NMV F104857 (3); stn 71, 193-207 m, F104867 (1); stn 73, 316 m, F104896 (1); stn 76, 650-646 m, FI 04837 (7) South Shetland Is., Elephant I., 326-382 m, AMER-03 stn 38, NMV FI 04849 (1); Deception I., 460-484 m, AMUR- 03 stn 88, FI 04834 (3); F104835 (1). Diagnosis. Psolidium species up to 42 mm long; body form elongate, narrow, high; dorsal and lateral scales conspicuous, thin, smooth, up to 2.0 mm wide; dorsal and lateral tube feet inconspicuous. Sole: outer peripheral single series of smaller tube feet; inner peripheral single series of larger tube feet; lacking mid- ventral (sole) radial series of tube feet. Dorsal ossicles: multi-layered perforated plates (scales), single-layered marginally, multi-layered centrally; reticulate thickenings extend from near margin towards centre of plate, thickenings near margin sometimes parallel to edge of plate, sometimes coalesce into smooth thickenings with smaller perforations, lacking frequent radiating linear thickenings between marginal perforations; up to 4 tube foot canals or marginal indentations per plate; dorsal and lateral tube feet lack support ossicles. Sole ossicles: throughout sole rare, small, smooth, thin, irregularly shaped rods and perforated plates, 0-9 large perforations, bluntly spined around margin, sometimes surface knobs, up to 260 pm long; sometimes as regular 4-holed plates with bluntly spined margin, sometimes surface knobs, up to 230 pm long; near margin of sole and peripheral tube feet thicker irregular rods and elongate perforated rod-plates, up to 420 pm long, intergrading with some round to oval, perforated thick plates up to 350 pm long, some with secondary layering. Tentacles: largest tentacle trunk ossicles; perforated plates, irregularly round to oval to elongate to narrow, thick, smooth, lacking rod-like thickenings between perforations; perforations large, irregular in size, rare incipient secondary developments; plates up to 750 pm long; rare rod-plates, up to 440 pm long. Colour (preserved). Dorsal and lateral body off-white (type material) to pale brown to brown (Bouvet and South Shetland Is material); scales faintly haloed, with pale outer edge (single- layer perforated edge of scales with white thickening); small dark brown markings; sole off-white to pale brown to brown. Distribution. South Sandwich Is, South Shetland Is, Bouvet I.; 146-759 m. Etymology. Named for Scott Whittaker (USNM), with appreciation and gratitude for his generous and gracious assistance to Cynthia Ahearn in preparing SEM images. Remarks. We judge that the 10 specimens from Bouvet I. (567 m) that were referred to Psolidium incertum (Theel, 1886) (- P. poriferum above) by Ludwig and Heding (1935) do not differ in any diagnostic characters from material from Bouvet I. examined here, and hence from Psolidium whittakeri sp. nov. In particular, the smooth perforated plate form of the tentacle trunk ossicles and the pale brown body and brown sole are the same. The preserved colour of the type material of Psolidium whittakeri sp. nov. (collected in 1975) is off-white. The recent material from Bouvet and the South Shetland Islands (collected in 2003 and 2004) is brown. It is difficult to judge whether this significant colour difference is due to preservation history or species difference. No significant morphological differences have been detected and it remains for current work on molecular genetic evidence to confirm species status. Of the series of poriferum- like species of Psolidium, we recognise that P. whittakeri sp. nov. (Bouvet, South Sandwich, South Shetland Is) is closest to P. pawsoni sp. nov. (Weddell Sea), but judge that there is significantly more frequent development of smooth white thickening on the scales of P. pawsoni. We note also that the P. whittakeri specimens are significantly larger (up to 42 mm long) than the P. pawsoni specimens that were available (up to 28 mm long). Acknowledgments We are most grateful for the generous assistance of: Leon Altoff and Audrey Lalconer (Marine Research Group of the Lield Naturalists Club of Victoria; photography); Ben Boonen (Photoshop, format of figures); Andrew Cabrinovic (BM(NH); loans, registration data); Mark Darragh (photography); Niki Davey (NIWA, Nelson; collaborative research); Phil Lambert (Chilean data); Thierry Laperousaz (SAM; loans, registration data); Susie Lockhart (AMLR R/V Yuhzmorgeologiya [2003] and Icefish [2004] material and data); Carsten Lueter (ZMB; loan); Kirrily Moore (TMAG; registration data); David Pawson (USNM; research facilitation); Chris Rowley (NMV; photography); Kareen Schnabel (NIWA; station data); Emily Whitfield (Marine Research Group of the Lield Naturalists Club of Victoria; research assistance); Scott Whittaker (USNM; assistance with SEM). Access to specimens and data collected by the New Zealand International Polar Year Census of Antarctic Marine Life Project, and to the NIWA invertebrate collections, is gratefully acknowledged. We are appreciative of the helpful review provided by Dave and Doris Pawson (USNM). Mark O’Loughlin records with sadness the death of his esteemed colleague Cynthia Ahearn. References Arnaud, P.M. 1974. Contribution a la bionomie marine benthique des regions antarctiques et subantarctiques. Tethys 6(3): 465-656. Bell, J.F. 1882. Studies in the Holothuroidea. I. On the genus Psolus and the forms allied thereto. Proceedings of the Zoological Society, London. 1882: 641-50, pi. 48. Branch, M. L., Jangoux, M., Alva, V., Massin, C. and Stampanato, S. 1993. The Echinodermata of subantarctic Marion and Prince Edward Islands. South African Journal of Antarctic Research 23(1, 2): 37-70. Cherbonnier, G. 1974. Invertebres marins des Xlleme et XVeme expeditions antarctiques Fra^aises en Terre Adelie. 15. Holothurides. Tethys 5(4): 601-10. 40 P. Mark O’Loughlin and Cynthia Ahearn Deichmann, E. 1941. The Holothurioidea collected by the Velero III during the years 1932-1938. I. Dendrochirotida. Allan Hancock Pacific Expeditions 8(3): 61-194, pis 10-30. Deichmann, E. 1947. Shallow water holothurians from Carbo de Hornos and adjacent waters. Anales del Museo Argentino de Ciencias Naturales. lnvertebrados Marinos 8: 325-51. Ekman, S. 1925. Holothurien. Further zoological results of the Swedish Antarctic Expedition 1901-1903 1(6): 1-194. Ekman, S. 1927. Holothurien der deutschen Siidpolar-Expedition 1901-1903 aus der Ostantarktis und von den Kerguelen. Deutsche Siidpolar-Expedition 19 (Zoology 11): 359-419. Forbes, E. 1841. A history of British starfishes and other animals of the class Echinodermata . John van Voorst: London. 267 pp. Grieg, J.A. 1929a. Echinodermata from the Palmer Archipelago, South Shetlands, South Georgia and the Bouvet Island. Scientific Results of the Norwegian Antarctic Expeditions 1927-1928 and 1928-1929 , 1(2): 1-16. Grieg, J.A. 1929b. Some echinoderms from the South Shetlands. Bergens Museum Arbok 1929. Naturvidenskapelig rekke 3: 1 - 10 . Grube, A.E. 1840. Aktinien, Echinodermen und Wiirmer des Adriatischen und Mittelmeeres. J. H. Bon: Konigsberg. 92 pp. Guille, A. 1982. A new genus and species of ophiacanthid brittlestar (Echinodermata: Ophiuroidea) from the Kerguelen Islands, with new taxonomic, biogeographic and quantitative data on the echinoderm fauna. Australian Museum Memoir 16: 67-87. Gutt, J. 1988. Zur Verbreitung und Okologie der Seegurken (Holothuroidea, Echinodermata) im Weddellmeer (Antarktis). Berichte zur Polarfors chung 41: 1-87. Gutt, J. 1991a. On the distribution and ecology of holothurians in the Weddell Sea (Antarctica). Polar Biology 11: 145-55. Gutt, J. 1991b. Are Weddell Sea holothurians typical representatives of the Antarctic benthos? Meeresforschung 33(4): 312-29. Hickman, V.V. 1962. Tasmanian sea-cucumbers (Holothuroidea). Papers and Proceedings of the Royal Society of Tasmania 96: 49-72, 2 pis, 186 figs. Lambert, P. 1996. Psolidium bidiscum , a new shallow-water psolid sea cucumber (Echinodermata: Holothuroidea) from the northeastern Pacific, previously misidentified as Psolidium bullatum Ohshima. Canadian Journal of Zoology 74: 20-31. Lampert, K. 1885. Die Seewalzen. Holothurioidea. Eine Systematische Monographie. 310 pp, 1 pi. in C. Semper (ed.) Reisen im Archipel der Philippinen 2(4)3: 1-310, 1 pi. Lampert, K. 1889. Die wahrend der expedition SMS Gazelle 1874-1876 von Prof. Dr. Th. Studer gesammelten holothurien. Zoologische Jahrbiicher. Abtheilung fur Systematik, Geographie und Biologie der Thiere 4(4): 806-58, pi. 24. Ludwig, H. 1886. Die von G. Chierchia auf der Fahrt der Kgl. Italianische Corvette Vettor Pisani gesammelten Holothurien. Zoologische Jahrbiicher 2: 1-36, 2 pis. Ludwig, H. 1892. Echinodermen. Pp. 325-61, pis 15-17 in: Dr H.G. Bronn’s Klassen und Ordnungen des Thier-Reichs 2(3) 3 Das System. C. F. Winter’sche Verlagshandlung: Leipzig and Heidelberg. 460 pp, 17 pis. Ludwig, H. 1894. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the US Fish Commission steamer Albatross, during 1891. XII. The Holothurioidea. Memoirs of the Museum of Comparative Zoology, Harvard University 17(3): 1-183, pis 1-19. Ludwig, H. 1898. Holothurien. Ergebnisse der Hamburger Magalhaensischen Sammelreise 1892/93 1: 1-98, 3 pis. Ludwig, H. & Heding, S.G. 1935. Die holothurien der Deutschen Tiefsee-Expedition. 1. Fusslose und dendrochirote formen. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition aufdem dampfer Valdivia 1898-1899 24: 123-214. Massin, C. 1992. Holothurians (Echinodermata) from Marion and Prince Edward Islands: new and little-known species. Zoologica Scripta 21(3): 311-24. Mortensen, T. 1925. On a small collection of echinoderms from the Antarctic Sea. Arkiv for zoologi 17A(31): 1-12, 8 figs. Mortensen, T. 1927. Handbook of the Echinoderms of the British Isles. Oxford University Press. London. 471 pp. O’Loughlin, PM. 2002. Report on selected species of BANZARE and ANARE Holothuroidea, with reviews of Meseres Ludwig and Heterocucumis Panning (Echinodermata). Memoirs of Museum Victoria 59(2): 297-325. O’Loughlin, PM. and Ahearn, C. 2005. A review of pygal-furrowed Synallactidae (Echinodermata: Holothuroidea), with new species from the Antarctic, Atlantic and Pacific oceans. Memoirs of Museum Victoria 62(2): 147-79. O’Loughlin, P.M., Bardsley, T.M. and O’Hara, T.D. 1994. A preliminary analysis of diversity and distribution of Holothurioidea from Prydz Bay and the MacRobertson Shelf, Eastern Antarctica. Pp. 549-55, 1 fig., 2 tbls in: David, B., Guille, A., Feral, J-P. and Roux, M. (eds), Echinoderms through Time. Proceedings of the Eighth International Echinoderm Conference, Dijon, France, 6-10 September, 1993. Balkema, Rotterdam. O’Loughlin PM. and Marie, D. 2008. Australian species of Psolidium Ludwig (Echinodermata: Holothuroidea: Psolidae). Memoirs of Museum Victoria 65: 1-22. Pawson, D.L. 1969. Holothuroidea. Pp. 36-38, 1 pi. in: V.C. Bushnell and J.W. Hedgpeth (eds). Distribution of Selected Groups of Marine Invertebrates in Waters South of35°S Latitude. Antarctic Map Folio Series (II). American Geographical Society, New York. Pawson, D.L. 1971. The Western Australian psolid holothurian Ceto cuvieria (Cuvier). Journal of the Royal Society of Western Australia 54(2): 33-39. Pawson, D.L. 1982. Holothuroidea. Pp. 813-18; figs pp. 791-92; pis 136, 137 in: S.P. Parker (editor-in-chief). Synopsis and Classification of Living Organisms. McGraw-Hill, New York. Pawson, D.L. and Fell, H.B. 1965. A revised classification of the dendrochirote holothurians. Breviora 214: 1-7. Pawson, D.L. and Valentine, J.F. 1981. Psolidium prostratum, new species, from off the east coast of the USA (Echinodermata: Holothuroidea). Proceedings of the Biological Society of Washington 94(2): 450-54. Perrier, R. 1902. Holothuries. Pp. 273-554, pis 12-22 in: Expeditions scientifiques du Travailleur et du Talisman pendant les annees 1880, 1881, 1882, 1883. Masson et Cie, Editeurs, Paris. Perrier, M. R. 1904. Holothuries du cap Horn. Bulletin du Museum d’Histoire Naturelle 1: 13-17. Perrier, R. 1905. Holothuries antarctiques du Museum d’Histoire Naturelle de Paris. Annales des Sciences Naturelles. Zoologie 1: 1-146, 5 pis. Studer, T. 1876. Uber Echinodermen aus dem antarktischen Meere und zwei neue Seeigel von den Papua-Inseln, gesammelt auf der Reise SMS Gazelle um die Erde. Monatsberichte der Koniglich Preussichen Akademie der Wissenschaften. Berlin. Pp. 452-65. Studer, T. 1879. Die fauna von Kerguelensland. Archiv fur Naturgeschicht 45(1): 104-41. Studer, T. 1889. Die Forschungsreise SMS Gazelle in den Jahren 1874 bis 1876. 3. Zoologie und Geologie. Foreword + 322 pp., 33 pis. Mittler und Son, Berlin. Thandar, A.S. 1999. Deep-sea holothuroids taken by the RV Africana II in 1959, from off the west coast of the Cape Peninsula, South Africa. Annals of the South Africa Museum 105: 364-409. Antarctic and Sub-Antarctic species of Psolidium Ludwig 41 Thandar, A.S. 2006. New species and new records of dendrochirotid anddactylochirotidholothuroids(Echinodermata: Holothuroidea) from the east coast of South Africa. Zootaxa 1245: 1-51. Theel, H. 1886. Report on the Holothurioidea dredged by HMS Challenger during the years 1873-1876. Report on the scientific results of the voyage of HMS Challenger, Zoology 14(39): 1-290, 16 pis. Vaney, C. 1908. Les holothuries de l'Expedition Antarctique Nationale Ecossaise. Transactions of the Royal Society of Edinburgh 46: 405-41, 5 pis. Vaney, C. 1914. Holothuries. Deuxieme Expedition Antarctique Fran 5 aise (1908-10 ). Sciences Naturelles: Documents Scientifiques. Masson et Cie, Editeurs, Paris. 54 pp., 5 pis. Memoirs of Museum Victoria 65: 43-50 (2008) ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs/index.asp Marmenuera , a new genus of leptophlebiid mayfly (Insecta: Ephemeroptera) endemic to Tasmania. J.C. Dean 1 , G.N.R. Forteath 2 & A.W. Osborn 2 1 Environment Protection Authority, Freshwater Sciences Unit, Ernest Jones Drive, Macleod, Victoria, 3085, Australia (john.dean@epa.vic.gov.au) 2 Honorary Research Fellow, Queen Victoria Museum and Art Gallery, Wellington Street, Launceston, Tasmania, 7250, Australia Abstract Dean, J.C., Forteath G.N.R. and Osborn, A.W. 2008. Marmenuera, a new genus of leptophlebiid mayfly (Insecta: Ephemeroptera) endemic to Tasmania. Memoirs of Museum Victoria 65: 43-50. The genus Marmenuera gen. nov. is established to accommodate two species of leptophlebiid mayfly, both of which are endemic to Tasmania. Adults and nymphs of M. ida (Tillyard) and M. tillyardi sp. nov. are described and figured. Keywords Mayflies; Australia; taxonomy; Marmenuera ; M. tillyardi sp. nov.; M. ida (Tillyard); Leptophlebiidae. Introduction The generic taxonomy of leptophlebiid mayflies of Australia, although still incomplete, has been advanced in recent years. A preliminary generic key to nymphs (Dean, 1999) included 15 described and eight undescribed leptophlebiid genera. Of the eight undescribed genera, three have since been described (. Loamaggalangta Dean et ah, 1999, Kaninga Dean, 2000 and Manggabora Dean and Suter, 2004), a fourth is described below and “Genus S” from southwest Australia has previously been referred to Loamaggalangta (Dean, 2000). The present paper erects a new genus to accommodate two species of Leptophlebiidae from Tasmania. The nymphs of these species were designated Genus W sp.AVl and Genus W sp.AV2 by Dean (1999). Recent biological studies in Lake Pedder and several inflowing streams provided an opportunity to rear out adults of both species. Repositories of type and other material are abbreviated as follows: Museum of Victoria (MV); Australian National Insect Collection, Canberra (ANIC); British Museum of Natural History (BMNH). Abbreviations of life stages of examined material are: N, nymph; MI, male imago; FI, female imago; MS, male subimago; FS, female subimago. Taxonomy Order Ephemeroptera, Family Leptophlebiidae, Subfamily Atalophlebiinae Genus Marmenuera gen. nov. Diagnostic features. Imago. Forewing (figs 1, 3, 17, 19): membrane hyaline, cells in apical third of wing translucent white; veins brown, many crossveins suffused with dark brown pigmentation; length-width ratio of forewing slightly less to slightly greater than 3.0; four to seven costal crossveins basal to the bulla, 11-19 distal to the bulla; MA forked at 0.40-0.43 wing length; MP 2 attached by crossvein to MP t at 0.18-0.20 wing length; base of ICUj linked to both CuA and CuP by a crossvein; ICu t and ICu 2 strongly diverging as wing margin approached. Hindwing (figs 2, 4, 18, 20): 0.22-0.26 length of forewing; length/width ratio 1. 6-1.9; costal margin with shallow concavity at about midlength, costal space broader both basal and distal to the concavity; vein Sc joining costal margin at 0.8-0.85 wing length; 7-13 costal crossveins and 6-11 subcostal crossveins. Legs: yellow or medium brown, all femora with two strongly developed dark brown bands, one at midlength and the other near the apex; tarsal claws similar, each claw with an apical hook and opposing ventral flange (fig. 9); forelegs of male with ratios of segment lengths 0.81-0.86, 1.00, 0.05-0.07, 0.34-0.35, 0.34-0.38, 0.26-0.30, 0.11-0.23. Male genitalia (figs 7, 8, 24, 25): claspers three-segmented, basal segment narrowing abruptly at about midlength; penes lobes fused in basal half, widely separated apically; each lobe relatively broad and approximately parallel-sided, with three moderate sized ventral spines close to inner margin and slightly posterior to where the lobes separate (fig. 8). Female 44 J.C. Dean, G.N.R. Forteath &A.W. Osborn sternum nine with apical margin deeply excised (Fig. 10). Subimago. Wings pale fawn/white, brown blotches and/or suffusions similar in distribution and intensity to corresponding imago. Mature nymph. Head prognathous; antennae at least half length of body. Mouthparts: clypeus with lateral margins sub-parallel; labrum (figs 12, 27) slightly wider than clypeus, width 1.7-1. 9 times length along median line, anterior margin with shallow to moderate mesal concavity, frontal setae arranged as single row, sub-apical setae arranged in a single row set back from anterior margin at about 0.9x labrum length; mandibles with outer margin somewhat angular at midlength, gently curved basally and distally, a tuft of long setae around midlength and a sparser series of long setae basal to the tuft, incisors slender, prostheca strongly developed; maxillae quadrate, subapical row of 30-35 pectinate setae, palp moderately short, three segmented, terminal segment almost as long as middle segment, middle segment bearing simple setae only; labium with glossae elongate, turned under ventrally, palp three-segmented, terminal segment almost as long as middle segment and with series of small spine -like setae along inner margin, submentum with fringe of long setae along lateral margins. Legs banded; femora moderately broad (figs 13, 28); tarsal claws smooth, without ventral teeth (figs 14, 29). Abdominal segments without setae on lateral margins, postero-lateral spines on segments 7-9; posterior margins of abdominal terga with series of long, robust spines interspersed with minute spines (fig. 15). Gills present on abdominal segments 1-7; each gill lanceolate, lateral tracheae strongly developed (fig. 16). Caudal filaments about Iff times length of body; each segment with apical whorl of both stout spine-like setae and fine hair-like setae, spine-like setae predominating near base of each filament but becoming shorter towards midlength where the hair-like setae predominate. Type species. Marmenuera tillyardi sp.nov. Etymology. The generic name is derived from “mar.me.nuer”, the western Tasmanian aboriginal word for “country” or “tribal territory” (Plomley 1976, p.191), and pertains to the endemicity of the genus to Tasmania. Remarks. The genus Marmenuera can be distinguished from all other leptophlebiid genera by the following combination of characters. Imago: (1) basal half of forewing with costal crossveins heavily suffused with dark brown, often with brown pigmentation more extensive; (2) forewing with ICUj attached to CuA and CuP by crossveins; (3) ICiq and ICu 2 strongly diverging as wing margin approached; (4) hindwing with shallow concavity at about midlength, and Sc joining wing margin at 0.8-0.85 wing length; (5) tarsal claws similar; (6) male genitalia with penes fused in basal half and widely separated apically, with three moderate-sized ventral spines situated slightly posterior to where the lobes separate; (7) female sternum nine with apical margin deeply excised. Nymph: (1) labrum slightly wider than clypeus; (2) width of labrum 1.7-1. 9 times length along median line; (3) labium with glossae elongate and turned under ventrally; (4) terminal segment of labial palp with series of small spine -like setae along inner margin; (5) submentum with fringe of long setae along lateral margins; (6) tarsal claws smooth; (7) gills lanceolate, moderately broad; (8) posterior margins of abdominal terga with row of stout spines interspersed with small spines. Marmenuera does not fit into any of the leptophlebiid lineages defined by Pescador and Peters (1980), and Towns and Peters (1980), and its closest relatives are probably here in Australia. The genus appears most closely related to Loamaggalangta Dean, Forteath and Osborn and Bibulmena Dean. The three genera share many characters, including attachment of the base of ICu to the CuA-CuP crossvein in the forewing, similar tarsal claws in the imago, a deeply excised sternum nine in the female, similarities in the nymphal mouthparts (in particular the elongate and underturned glossae, the fringe of setae on the lateral margins of the submentum and the small spine-like setae along the inner margin of segment 3 of the labial palps) and smooth tarsal claws in the nymph. Christidis (2005) placed Bibulmena and Loamaggalangta at the base of a large clade containing many of the Australian leptophlebiid genera, but found no support for the grouping of these two genera into a monophyletic clade. The character states listed above are perhaps primitive and not synapomorphies. Marmenuera tillyardi sp.nov. Figures 1-16 Genus W sp.AVl Dean 1999, p. 86, figs 239-41. Types. Holotype: Male imago (reared from nymph), Tasmania, un- named creek flowing into Lake Pedder (Tasmap 1:25 000 series. Serpentine 4026, Grid reference 55G 4184 52606), collected 28 Jan 2006, emerged 30 Jan 2006, A. Osborn and N. Forteath (MV, T-20000). Paratypes: Male imago and female imago (both reared from nymph), same location as holotype, collected 28 Jan 2006, emerged 19 Mar 2006 and 23 Feb 2006 respectively, A.Osborn & N.Forteath (MV, T-20001 and T-20002 respectively); male imago and female imago (both reared from nymph), same location as holotype, collected 5 Jan 2006 and 28 Jan 2006 respectively, emerged 14 Jan 2006 and 13 Feb 2006 respectively, A.Osborn and N.Forteath (ANIC). Material examined. Tasmania. 6 MI, 12 FI (all reared from nymphs), un-named creek flowing into Lake Pedder (type locality), 28 Jan 2006, emerged various dates, A. Osborn and N. Forteath; 3 MI, 1 FI (all reared from nymphs), same location, 25 Jan 1998, N. Forteath; 1 FI (reared from nymph), same location, 23 Feb 1998, N. Forteath; 1 MI, 1 FI (both reared from nymphs), same location, 4 Mar 1998, N. Forteath; 2 MI, 2 FI (all reared from nymphs), same location, 5 Jan 2006, A. Osborn and N. Forteath; 1 MS, 1 FS (both reared from nymphs), 4 N, same location, 27 Dec 1997, N.Forteath; 2 N, Piney Creek, Heamskirk Rd, 27 Oct 1994, MRHI; 1 FI, 1 FS (both reared from nymphs). Lake Pedder, Trappes Bay, 4 Mar 1998, N. Forteath; 1 MI, 2FI (all reared from nymphs). Lake Pedder, Serpentine Arm, Apr 2005, N.Forteath. Description. Male imago: Length: body 8.8-10.2 mm., forewing 8.8-10.1 mm., caudal filaments 12.4-14.2 mm. Eyes: upper lobes pale pink/brown, separated mesally by a distance approximately 1/8 of the diameter of the upper lobe in dorsal view; lower lobes grey. Thorax: medium-dark brown; pronotum with narrow raised carina along median line, dark brown markings medially on posterior half of segment, dark brown Marmenuera, a new genus of leptophlebiid mayfly (Insecta: Ephemeroptera) endemic to Tasmania. 45 Figures 1-4. Marmenuera tillyardi. Male imago: 1, forewing; 2, hind wing. Female imago: 3, forewing; 4, hind wing. Figures 5-10. Marmenuera tillyardi. Male imago: 5, abdomen, dorsal; 6, abdomen, ventral; 7, penes, dorsal; 8, penes, ventral, slide preparation; 9, foretarsal claw. Female imago: 10, sternum nine. 46 J.C. Dean, G.N.R. Forteath &A.W. Osborn Figures 11-16. Marmenuera tillyardi. Nymph: 11, abdomen, dorsal; 12, labrum; 13, forefemur; 14, foretarsus; 15, posterior margin of abdominal segment four; 16, gill, abdominal segment four. along lateral margins and curved dark brown band parallel to lateral margin approximately midway between margin and medial carina. Legs: predominantly medium brown, tending to paler in apical half of tibiae and tarsi; all femora with two broad dark brown bands, one just beyond midlength and the other sub-apical; tarsal claws similar, each claw with terminal sclerotised hook and opposing ventral process. Forewing (fig. 1): membrane hyaline, stigmatic region opaque, white; costal crossveins in basal half of wing heavily suffused with dark brown, subcostal and R x -R 2 crossveins in basal half of wing less strongly suffused with dark brown, crossveins near MA fork not suffused with brown. Abdomen: terga generally reddish-brown with dark brown markings, tergum 6 with four conspicuous dark brown longitudinal bands extending over most of the segment (fig. 5), segment 7 with similar bands but restricted to anterior half of the segment, segment 8 with dark markings only adjacent to the anterior margin; sterna reddish brown, inconspicuous paler markings on segments 2-5 (fig. 6), darker brown markings poorly defined. Genitalia (figs 7, 8): penes lobes fused basally, widely separated apically; each lobe relatively broad and approximately parallel-sided, apically truncate, outer margin somewhat angular just before apex; three moderate sized ventral spines close to inner margin posterior to where the lobes separate. Female imago. Length: body 10.1-12.2 mm; forewing 10.0-11.8 mm; caudal filaments 13.4-16.4 mm. General colour similar to male imago. Forewing: membrane slightly opaque; costal, subcostal and R t -R 2 crossveins all surrounded by suffusion of dark brown, more strongly developed than in male. Abdomen: medium reddish-brown, terga with darker brown markings, sterna medium reddish-brown, no strong markings; sternum nine deeply cleft (fig. 10). Subimago. Wings pale fawn/yellow, brown blotches and/or suffusions similar in distribution and intensity to corresponding imago. Mature nymph. Body length ranging from about 10 to 13 mm.; antennae about half length of body; cerci about IV 2 times length of body, terminal filament a little longer. General colour yellow with dark brown markings; abdomen with conspicuous pattern of dark brown and yellow, abdominal segments 6 and 9 darker than remaining segments (fig. 11); all legs with dark Marmenuera, a new genus of leptophlebiid mayfly (Insecta: Ephemeroptera) endemic to Tasmania. 47 banding. Mouthparts: labrum (fig. 12) a little broader than clypeus; maximum width 1.7-1. 9 times length along median line; maximum width at about 3/4 labrum length, basal to this the lateral margins relatively straight and diverging; anterior margin shallowly concave, with five well developed medial denticles; single row of setae close to frontal edge of labrum, sub-apical setal fringe also consisting of a single row set back from anterior margin at about 0.9 labrum length. Legs relatively slender; forefemora length 3L — 1 times width (Fig. 13), outer margin with moderate number of short, spine -like setae and also longer, hair-like setae; foretarsus (fig. 14) with 30-40 ventral spines; tarsal claws smooth, without ventral teeth. Postero -lateral spines on abdominal segments 7 (small) and 8-9 (relatively large); posterior margins of abdominal terga with series of large, conspicuous spines interspersed with the occasional shorter spine (fig. 15). Gills lanceolate, lateral tracheae moderately developed (fig. 16). Etymology. The species is named for R.J.Tillyard, who first recognised that this was probably an undescribed species (see below). Remarks. When describing Atalophlebia ida, Tillyard (1936) referred to specimens from Cradle Mountain which were smaller and less spotted than the type series. It now seems likely that these belong to the species herein described as M. tillyardi, and Tillyard’s decision not to describe the male imago of A. ida based on the Cradle Mountain material has been proven taxonomically astute. Scholes (1961) recommended that the imago and subimago of A. ida should be referred to as the “Large Speckled Spinner” and “Large Speckled Dun” respectively, and we suggest that “Lesser Speckled Spinner” and “Lesser Speckled Dun” are appropriate common names for the imago and subimago of Marmenuera tillyardi. This reflects both the smaller size of the new species and the reduced wing pigmentation. Marmenuera ida (Tillyard) Figures 17-29 Atalophlebia ida Tillyard, 1936: p. 42, fig. 7, plate 1(10). Massartellopsis ida Scholes, 1961: p. 30, plate 2. Genus W sp.AV2 Dean, 1999: p. 86, figs 242, 243. Types. Holotype: female imago, Tasmania, Dee Bridge, 7 Feb 1933, R. J. Tillyard (BMNH). Paratypes: five female imagos, collected with holotype (BMNH); one male subimago. Lake St Clair, 6 Feb 1933, R. J. Tillyard (BMNH). Types not examined. Material examined. Tasmania. 5N, Lake Pedder, Trappes Inlet, February and April 1997, N. Forteath; 3FI (2 reared), 1FS (reared), same location, 1 Feb 1998, N. Forteath; IMS (reared), same location, 12 Jan 1998, N. Forteath; IMS (reared), same location, 4 Mar 1998, N. Forteath; 6MI,5FI (all reared), 21N, same location, Nov-Dee 2006, N. Forteath and A. Osborn; IN, Lake Lilia, Cradle Mountain, 25 Mar 1998, J. Dean; IN, Dip River Falls, 31 Oct 1998, D. Cartwright; 4FI, Huon River Crossing, 16 Feb 1971, A. Neboiss; 1MI, IMS, 1FS, D'Entrecasteaux Passage, Exit Cave, Janl993, A. Clarke. Description. Male imago. Length: body 1 1 .0-12.4 mm, forewing 11.2-12.0 mm, caudal filaments 18.4—24.2 mm. Eyes: upper lobes pale pink/brown, in contact on meson of head, lower lobes grey. Thorax: medium-dark brown; pronotum with narrow raised carina along median line, dark brown markings medially on posterior margin, along lateral margins and parallel to lateral margin approximately midway between margin and medial carina. Legs: predominantly yellow, all femora with two broad dark brown bands, one just beyond midlength and the other subapical; tarsal claws similar, each claw with terminal sclerotised hook and opposing ventral process. Forewing (fig. 17): membrane hyaline, stigmatic region slightly opaque, white; all costal, subcostal and R x -R 2 crossveins heavily suffused with dark brown, crossveins in vicinity of MA fork also strongly suffused with dark brown; large brown blotch filling area between veins Sc and R 2 at midlength of wing. Abdomen: strongly contrasting pattern of dark brown and yellow; terga 2-7 yellow in anterior half and dark brown in posterior half, yellow colouration extended back as narrow band along median line (fig. 22), extent of yellow colouration increasing from segments 6 to 8, segments 9 and 10 predominantly brown; sterna reddish brown, pale yellow along lateral margins, segments 2-7 with two pairs of small pale spots, the anterior pair more widely eparated than the posterior pair (fig. 23). Genitalia (figs 24—25): penes lobes fused basally, widely separated apically; each lobe relatively broad, apex rounded and turned slightly inwards, sub- apically with outer margin evenly curved, three moderate sized ventral spines posterior to where the lobes separate. Female imago. Length: body 11.6-14.8 mm.; forewing 12.0-14.2 mm; caudal filaments 18.2-22.2 mm. Forewing: pigmentation more extensive than in male; dark brown suffusions around costal, subcostal and R t -R, crossveins broad, almost circular; large brown blotch surrounding MA fork. General colour and abdominal markings similar to male imago. Mature nymph. Body length ranging from 12-16 mm; antennae a little over half length of body; cerci about 114-1 Vi body length, terminal filament a little longer. General colour yellow with brown markings; abdomen with conspicuous pattern of dark brown and yellow, segments 2-8 yellow towards anterior margin and dark brown to the posterior (fig. 26); all legs yellow with dark banding. Mouthparts: Labrum (fig. 27) a little broader than clypeus; maximum width 1.7-1. 9 times length along median line; maximum width at about 2/3 labrum length, basal to this the lateral margins relatively straight and diverging; anterior margin moderately concave, with five well developed medial denticles. Legs moderately broad (fig. 28); forefemora length a little over 3 times width, outer margin bearing moderate length spine-like setae and also some longer, hair-like setae; fore-tarsus with 15-20 ventral spines (fig. 29); tarsal claws smooth, without ventral teeth. Postero-lateral spines on abdominal segments 7 (small) and 8-9 (relatively large); posterior margins of abdominal terga with series of large, conspicuous spines interspersed with the occasional shorter spine. Gills lanceolate, lateral tracheae moderately developed. Remarks. Although the holotype has not been examined the description and, in particular, the image of the female forewing presented by Tillyard (1936) leave no doubt as to the identity of this species. The species is readily distinguished from M. tillyardi in the adult by the abdominal colour pattern, the more extensive pigmentation of the forewing and the structure of the 48 J.C. Dean, G.N.R. Forteath &A.W. Osborn Figures 17-21. Marmenuera ida. Male imago: 17, forewing; 18, hindwing. Female imago: 19, forewing; 20, hindwing; 21, forewing, holotype. male genitalia, and in the nymph by the abdominal colour pattern and the structure of the labrum and foreleg. Scholes (1961) referred this species to the genus Massartellopsis, presumably based on advice from Edgar Riek (CSIRO Division of Entomology), but it is now known that the genus Massartellopsis is endemic to South America. Scholes (1961) suggested the common names “Large Speckled Spinner” and “Large Speckled Dun” for the imago and subimago respectively of Marmenuera ida, and we consider this appropriate. Keys to species 1. Adults 2 - Nymphs 3 2. Forewings heavily pigmented, crossveins around MA fork suffused with brown (figs 17,19); male imago with outer margin of penes lobes evenly rounded (fig. 24) Marmenuera ida - Forewings lightly pigmented, crossveins around MA fork not suffused with brown (figs 1,3); male imago with outer margin of penes lobes angular (fig. 7) Marmenuera tillyardi 3. Foretarsus with 15-20 ventral spines (fig. 29); forefemur with spines along outer margin relatively long (fig. 28) Marmenuera ida - Foretarsus with 30-40 ventral spines (fig. 14); forefemur with spines along outer margin relatively short (fig. 13) Marmenuera tillyardi Marmenuera, a new genus of leptophlebiid mayfly (Insecta: Ephemeroptera) endemic to Tasmania. 49 Figures 22-29. Marmenuera ida. Male imago: 22, abdomen, dorsal; 23, abdomen, ventral; 24, penes, dorsal; 25, penes, ventral. Nymph: 26, abdomen, dorsal; 27, labrum; 28, forefemur; 29, foretarsus. Acknowledgments We are indebted to the Tasmanian Hydroelectric Commission and the W. D. Booth Charitable Trust for support of field work undertaken on Lake Pedder by A. Osborn and N. Forteath. The Council and Editor of the Royal Society of Tasmania are thanked for permission to reproduce Plate I Figure 10 from Tillyard (1936). Preparation of the manuscript was supported by the Environment Protection Authority of Victoria. 50 J.C. Dean, G.N.R. Forteath &A.W. Osborn References Christidis, F. 2005. Phylogenetic relationships of the Australian Leptophlebiidae (Ephemeroptera). Invertebrate Systematics 19: 531-39. Dean, J.C. 1999. Preliminary keys for the identification of Australian mayfly nymphs of the Family Leptophlebiidae. Identification Guide no. 20. Cooperative Research Centre for Freshwater Ecology, Albury, New South Wales. Pp iii + 91. Dean, J.C. 2000. Descriptions of new Leptophlebiidae (Insecta: Ephemeroptera) from Australia. II. Kaninga, a new monotypic genus from south-western Australia. Records of the Western Australian Museum 20: 87-94. Dean, J.C.; Forteath,G.N.R. and Osborn, A.W. 1999. Loamaggalangta pedderensis gen. & sp.nov.: A new mayfly from Tasmania (Ephemeroptera: Leptophlebiidae: Atalophlebiinae). Australian Journal of Entomology 38: 12-16. Dean, J.C. and Suter PJ. 2004. Descriptions of new species and a new genus of leptophlebiid mayflies (Insecta: Ephemeroptera) from the Northern Territory, Australia. Memoirs of the Museum of Victoria 61: 111-18. Pescador, M.L. and Peters, W.L. 1980. Phylogenetic relationships and zoogeography of cool-adapted Leptophlebiidae (Ephemeroptera) in southern South America. Pp. 43-56 in: Flannagan, J.F. and Marshall, K.E. (eds). Advances in Ephemeroptera Biology. Plenum Press: New York. Plomley, N.J.B. 1976. A word-list of the Tasmanian Aboriginal Languages. Foot & Playsted: Launceston. Pp xv + 486. Scholes, D. 1961. Fly-fisher in Tasmania. Melbourne University Press: Carlton. Pp xiv + 208. Tillyard, R.J. 1936. The trout-food insects of Tasmania. Part II: A monograph of the mayflies of Tasmania. Papers and Proceedings of the Royal Society of Tasmania. 1935: 23-59. Towns, D.R. and Peters, W.L. 1980. Phylogenetic relationships of the Leptophlebiidae of New Zealand (Ephemeroptera). Pp 57-69 in: Flannagan, J.F. and Marshall, K.E. (eds). Advances in Ephemeroptera Biology. Plenum Press: New York. Memoirs of Museum Victoria 65: 51-56 (2008) ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs/index.asp A new species in the Ophiocoma erinaceus complex from the South-west Pacific Ocean (Echinodermata: Ophiuroidea: Ophiocomidae) Milena Benavides- Serrato 1 and Timothy D. O'Hara 2 1 Echinoderms Curator, Colombian Marine Natural Museum, Institute of Marine and Coastal Research “Jose Benito Vives de Andreis” INVEMAR, Cerro Punta de Betfn 1016, Santa Marta, Colombia (milena_benavides@invemar.org.co) 2 Museum Victoria, GPO Box 666, Melbourne, Victoria, 3001, Australia (tohara@museum.vic.gov.au) Abstract Benavides-Serrato, M. and O'Hara, T.D. 2008. A new species in the Ophiocoma erinaceus complex from the South-west Pacific Ocean (Echinodermata: Ophiuroidea: Ophiocomidae). Memoirs of Museum Victoria 65: 51-56. A new species is described from the Ophiocoma erinaceus complex of tropical ophiuroids, reliably distinguished by the almost complete lack of granulation on the ventral disc surface. The new species is currently known only from the south-west Pacific Ocean, ranging from the northern section of the Great Barrier Reef to Tonga. Keywords Echinodermata, Ophiuroidea, Ophiocoma Introduction Ophiuroids of the genus Ophiocoma, common on coral or rocky reefs throughout the tropics, are difficult to identify and many historical records are inaccurate (Devaney 1968, Devaney 1970). Part of the problem has been the lack of distinctive morphological characters and the ability of some species to change colour pattern, from being consistently dark during the day to variegated at night (Hendler 1984, O’Hara et al. 2004). Devaney (1968, 1970) revised the taxonomy of the genus by introducing new taxonomic characters based around the shape of the oral and dental plates and their associated papillae. However, some problems were left unresolved. One taxonomically problematic group contains black- coloured specimens with alternating numbers of arm spines from the Indo-Pacific Ocean generally known as Ophiocoma erinaceus (Muller & Troschel, 1842). The concept of this ‘species’ has varied widely from being considered a variety of the variegated species O. scolopendrina Lamarck, 1816 (eg Matsumoto, 1917) to being divided into species based on the number of tentacle- scales; two in Ophiocoma erinaceus and one in O. schoenleinii Muller & Troschel, 1842 (eg see Clark & Rowe, 1971). Devaney (1970) considered them ‘polymorphs of the same species having phenotypically linked morphological characters’. Field work across northern Australia led one of the authors (O’Hara) to consider that two species were indeed present, one with red-tube feet ( O . erinaceus) and one with grey tube feet (i O . schoenleinii ). However, subsequent pilot molecular data suggested that three clades were present: with red-tube feet and two tentacle scales ( O . erinaceus), grey-tube feet and one wide tentacle scale ( O . schoenleinii), and grey-tube feet and two tentacle scales (undescribed) (O’Hara et al., 2004). Materials and methods This paper is the formal scientific description of the third species in the complex, using a suite of morphological characters that include those of the dental plates introduced by Devaney (1970). The three known species in the complex are distinguished. Material examined for this paper is lodged in Museum Victoria, Melbourne (MV). The abbreviation d.d. is used for disc diameter. Ophiocoma cynthiae sp. nov. Figures 1, 2c, 2f Material examined. Holotype: Queensland, Great Barrier Reef, Raine Island, shallow rocks off tower, 11°35.5’S, 144°2.3’E, 1-2 m, 5 Dec 2001, MV FI 12641 (ethanol, 17 mm d.d.). Paratypes: Australia, Queensland, Great Barrier Reef, Raine Island, shallow rocks off tower, 11°35.5’S, 144°2.3’E, 1-2 m, 5 Dec 2001, MV F91535 (5 individuals, ethanol, d.d. 21 mm, 22 mm, 20 mm, 18 mm, 13 mm); Fiji, Naviti, 18°12’S, 178°11’E, 0-2 m, 7 Jul 2002, MV F101821 (1, ethanol, d.d. 21 mm); Tonga, Vaua’u, Keitahi, 18°36’S, 173°55’W, 0-2 m, 5 Oct 1983, MV F91614 (1, dry, d.d. 15 mm). Comparative material examined. Ophiocoma schoenleinii Muller & Troschel. Australia, Queensland, Raine Island, shallow rocks off tower, 12 Dec 2001, 11°35.5’S, 144°2.3’E, 1-2 m, MV F101823 (1, ethanol, d.d. 15 mm); Moulter Cay, 11°21.4’S, 144°01.5’E, 1 m, 11 Dec 2001, MV F91534 (2, ethanol, d.d. 12 mm, 13 mm); Lizard Island, Coconut Beach, 14°40.8’S, 145°28.5’E, 2-5 m, 25 Oct 2005, MV F109814 (1, ethanol, d.d. 13 mm); Bird Island Reef, 14°4.6’S, 145°28’E, 5-15 m, 29 Oct 2005, MV F109813 (3, ethanol, d.d. 20 mm, 20 mm, 18 mm); Papua New Guinea, Bora Bada, MV F91583 (2, ethanol, d.d. 12 mm, 13 mm); Fiji, Naviti, 18°12’S, 178°1’E, 0-2 m, 7 Jul 2002, MV F93796 (1, ethanol, d.d. 20 mm); Vuda Point (between Nadi and Handtoka), 17°30’S, 177°25’E, 4 Jul 1981, MV F91593 (2, dry, d.d. 13 52 Milena Benavides-Serrato and Timothy D. O'Hara mm, 17 mm). Ophiocoma erinaceus Muller & Troschel. Western Australia, Ashmore reef. West lagoon, 12°14.29’S, 123°0.77’E, 3-9 m, 2 Oct 2002, MV F93633 (1, ethanol, d.d. 18 mm); Ashmore reef, west lagoon, 12°13.26’S, 122°59.28’E, 6-11 m, 1 Oct 2002, MV F93624 (5, ethanol, d.d. 13 mm, 11 mm, 13 mm, 6 mm, 5 mm); Queensland, Raine Island, shallow rocks off tower, 11°35.5’S, 144°2.3’E, 1-2 m, 5 Dec 2001, F91536 (2, ethanol, d.d. 15 mm, 20 mm); Moulter Cay, reef rim, 11°24.8’S, 144°01.9’E, 1-5 m, 10 Dec 2001, MV F91544 (1, ethanol); Lizard Island, First beach, 14°39.7’S, 145°26.8’E, 2^1 m, 28 Oct 2005, MV FI 09808 (1, ethanol, d.d. 19 mm). Description. Holotype. Disc 17 mm d.d, pentagonal with slightly incised interradial margins. Dorsal disc densely covered with rounded granules 0.15 mm diameter, 13-20 per mm 2 , sparse or patchy near the arms bases and interradial margins, covering radial shields. Ventral disc without granules, covered in thick wrinkled epithelium, disc plates obscured; a few elongated spinelets occur in a row on either side of the genital slit adjacent to the oral shields. Oral shields oval, longer (2.2 mm) than broad (1.9 mm), widest point a little distal of the midline, slightly depressed in the distal centre. Adoral shields triangular, much smaller than the oral shield, slightly lobed radial angle, restricted to the lateral edge of the oral shield, not meeting radially or interradially, margins obscured by epidermis (Fig. lc). Four oral papillae on each side of jaw angle; inner is spiniform, twice as high as wide; second is triangular, wider than high; third is largest, two to three times as wide as high, longest proximally; last (buccal tentacle scale) is wide and low, up to four times as wide as high, proximal end extending under the third oral papilla. Ten to 11 dental Figure 1. Ophiocoma cynthiae sp. nov. holotype (22 mm d.d.): a, dorsal surface of the whole animal; b, ventral surface; c, detail of oral frame; d, detail of the dorsal disc and arm surfaces. New species in the Ophiocoma erinaceus complex from the South-west Pacific Ocean 53 Figure 2. Dental plates of Ophiocoma erinaceus (MV F109808, 22 mm d.d.): a, external surface, d, internal, Ophiocoma schoenleinii (MV F109813, 21 mm d.d.); b, external; e, internal. Ophiocoma cynthiae sp. nov. (MV F91535, paratype 22 mm d.d.): c, external; f, internal. 54 Milena Benavides-Serrato and Timothy D. O'Hara papillae on each jaw, placed in three vertical rows near teeth, inner row slightly smaller, and an irregular transverse row or cluster across the jaw margin between the inner oral papillae; four hyaline tipped teeth in a vertical row, twice as wide as high. First ventral arm plate much smaller than succeeding plates, with a straight or rounded proximal margin, straight to convex lateral sides and a convex distal margin, longitudinally grooved; second plate as wide as long, widest distally, with a convex distal edge, rounded lateral angles, recurved lateral sides and truncate proximal edge, often sunken proximally so that the distal edge of the first plate projects over the proximal edge of the second; succeeding plates 1.5 times broader than long, widest in the distal half of the plate, convex distal margin, sharp lateral angles, recurved lateral sides around tentacle pore, concave proximal edge overlain by preceding plate, plates in contact for more than half of arm length. Upper arm plates fan-shaped, usually wider than long, with a convex to truncate distal margin, straight divergent lateral sides and a slightly concave proximal border overlain by the preceding plate, plates contiguous throughout the arm, plates forming a zig-zag row after the fifth segment past the disc, displaced away from the enlarged alternating upper arm spines. Lateral arm plates separated from each other for all arm length. Number of arm spines on each side of first twenty segments as follows (clockwise from the madreporite): 3 3 3 4 3 3 3 4 3 3 3 4 3 3 3 3 3 3 4 4 3 3 3 3 2 3 3 4 3 3 3 4 2 3 3 4 3 3 3 4 4 4 4 4 4 4 4 5 4 4 4 4 4 4 4 5 4 4 4 5 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 5 4 4 4 4 5 4 4 4 4 5 4 4 5 4 4 4 4 4 4 4 4 5 4 4 5 4 4 4 4 4 4 3 5 4 5 4 4 5 4 4 5 4 4 4 3 4 3 4 4 4 4 3 4 3 4 3 4 4 4 4 4 3 4 3 4 4 3 4 4 3 4 3 4 4 4 4 4 4 4 3 4 4 4 4 4 4 3 4 4 3 4 3 4 3 4 3 3 4 3 4 4 3 4 3 3 4 3 4 4 3 4 3 3 4 3 4 4 3 4 3 3 4 3 4 Arm spines of the first 5-7 segments flattened or curved, with truncate tips, subequal or uppermost longest, mostly longer than the ventral arm plate (except on first segment); spines on succeeding segments becoming progressively more cylindrical; spines start alternating 3-4 after segments 12-16, alternating uppermost spines become elongated and thickened, cigar- to club-shaped, longest in the middle of the arm, to three segments in length, 4-4.5 mm long, lower spines elongate and tapering; distal arm segments with three slender non- alternating spines. Two oval subequal tentacle scales on each pore, sitting side by side on the lateral arm plate, reducing to one near the arm tips. Colour. Ethanol: Disc and arms uniform dark brown colour. Upper side spines of five first segments of arm spines with some white longitudinal bands along the edge; tube feet dark, with a pale tip near the mouth, oral tentacles pale; mouth frame basically tan with small dark spots, madreporite white. Paratypes. A dental plate, dissected from one paratype (F91535, 22 mm d.d., Fig 2c), was 4.9 mm long and 1.7 mm wide, with two foramina, the lower septa twice as thick as the upper one, and dental papillae confined to the upper third of the plate, which is slightly narrower than the rest of the plate. Paratypes generally had more elongate granules or spinelets along the genital slit, extending to 1/2R. On paratype F91614 the dorsal disc granules form a very regular sharp boundary at the edge of the disc and (since it is preserved dry) small thin overlapping ventral disc scales (0.20-0.25 mm wide) can be clearly seen extending from the lateral margin to the oral shield. The second oral papillae can sometimes be larger than the third, but this is not consistent on a specimen. There can be up to 17 dental papillae. Paratype F101821 differs in having ventral arm plates with a concave or slightly notched distal margin except near the arm tip, the dorsal arm plates are regularly arranged in a series, not alternating from side to side, and the maximum arm spine length is 8.6 mm. Paratypes F101821 and F91614 from Fiji and Tonga respectively, have a lighter colour than the holotype and paratypes (F91535) from Australia, the mouth frame is white and the arms have broad bands of dark and pale segments. This may represent the “night” colour form. Distribution. Northern Great Barrier Reef, Fiji, Tonga, 0-2 m; under coral slabs or within dead coral matrix. Etymology. This species is named after friend and tireless colleague Cynthia Ahearn, the former Museum Specialist in Echinoderms at the Smithsonian Institution who passed away in August 2008. Remarks. The new species is clearly an Ophiocoma Agassiz as defined by Devaney (1970), possessing both dental and oral papillae, the latter in a continuous row, small adoral plates restricted to the lateral sides of the oral plate, a covering of spherical granules on the dorsal disc surface, and rounded arm spines in the middle of the arm. It belongs to the ‘scolopendrina’ group of species (Devaney 1970) with alternating numbers of arm spines and the uppermost arm spine enlarged. Within this group it shares many features in common with O. erinaceus including the dark colour, the presence of two to three arm spines on the first segment, the coarse disc granulation (10-20 per mm 2 ) and the size of the upper arm spines, which are longer on the side with four spines. Within, the O. erinaceus complex, O. cynthiae is distinguished by the lack of granules on the ventral disc surface (except for a few sparsely distributed along the genital slit near the oral shield). In O. schoenleinii they persist as a wedge near the ventral margin and extend almost to the oral shields in O. erinaceus. Furthermore, O. schoenleinii is distinguished by having only one wide tentacle scale on most pores past the first few segments, and denser dorsal disc granules (>20 per mm 2 ). Ophiocoma erinaceus can be further distinguished by the colour of the tube feet: bright red in live specimens and white in ethanol-preserved ones. Comparison of the dental plates of the three species (Fig. 2) indicates other possible differentiating characters. The dental plate of the new species is somewhat larger on specimens of similar size (21-22 mm d.d.), and is less narrowed on the region supporting the dental papillae. There are only two teeth foramina and the septum of second is much wider than the New species in the Ophiocoma erinaceus complex from the South-west Pacific Ocean 55 first. The dental plates of the other two species have three foramina, but the third is much larger on O. schoenleinii. However, due to the low numbers of specimens of the new species we are reluctant to dissect additional material and the intraspecific or size-related variability of these characters is unknown at this stage. This new species is similar to the description of the nominal species Ophiocoma tartarea Lyman, 1861, from Hawaii which was considered a synonym of O. erinaceus four years later by the same author (Lyman, 1865). In both his description of O. tartarea (1861) and O. erinaceus (1865) he refers to the absence of granules on the ventral disc surface, but without distinguishing between a wedge of granules and no granules at all. We have examined images of Muller & Troschel’s type specimens in the Museum fur Naturkunde Humboldt-Universitaet zu Berlin of both O. erinaceus (ZMB Ech 922) and O. schoenleinii (ZMB Ech 930, 4658). The type specimen of O. erinaceus clearly has a wedge of granules on the ventral interradial margin. The type specimens of O. tartarea appear to be lost. They were originally deposited in the museum of the Boston Society of Natural History (now called the Museum of Science), which transferred the majority of their collection in the 1940s to the Museum of Comparative Zoology in Harvard and some specimens to the United States National Museum. Downey (1969) does not list this species in her catalogue of US ophiuroid-type specimens, and there is no record of them in these museums today (Kirdahy, Aheam & Boyett pers. comm.). Given the lack of available type material, the early synonymy of O. tartarea with O. erinaceus by the same author, and the lack of known records of specimens without any ventral granules from Hawaii, we proposed to erect a new species, O. cynthiae. Acknowledgments We would like to thank Chris Rowley (Museum Victoria) for taking the camera images of the new species; Joan Clark (University of Melbourne) and Kate Naughton (Museum Victoria) for assisting with the SEMs; Dr Carsten Lueter (Museum fur Naturkunde, Humboldt-Universitaet zu Berlin) for providing images of type specimens of Ophiocoma erinaceus and O. schoenleinii ; Carolyn Kirdahy (Boston Museum of Science), Cynthia Ahearn (formerly of the United States National Museum) and Mary Catherine Boyett (Museum of Comparative Zoology, Harvard) for attempting to trace the type specimens of Ophiocoma tartarea-, the Raine Island Corporation and Environment Australia for supporting the fieldwork to Raine Island and Ashmore Reef; Dr Maria Byrne and Mark O’Loughlin for collecting material from Fiji and Tonga respectively. References Clark, A.M. & Rowe, F.W.E. 1971. Monograph of Shallow-water Indo-west Pacific Echinoderms. Trustees of the British Museum (Natural History). London. 238 pp, 31 pis. Devaney, D.M. 1968. The systematics and Post-Larval growth changes in Ophiocomid brittlestars. PhD thesis. University of Hawaii. 272 pp. Devaney, D.M. 1970. Studies on ophiocomid brittlestarts.l. A new genus ( Clarkcoma ) of Ophiocominae with a reevaluation of the genus Ophiocoma. Smithsonian Contribution to Zoology 51: 1-41. Downey, M.E. (1969). Catalog of recent ophiuroid type specimens in major collections in the United States. Bulletin of the United States National Museum 293: 1-239. Hendler, G. 1984. Brittlestar colour-change and phototaxis (Echinodermata: Ophiuroidea: Ophiocomidae). Pubblicazioni Della Stazione Zoologica Di Napoli I ( Marine Ecology) 5: 379-401. Lyman, T. (1861). Descriptions of new Ophiuridae. Proceedings of the Boston Society of Natural History 8: 75-86. Lyman, T. (1865). Ophiuridae and Astrophytidae. Illustrated Catalogue of the Museum of Comparative Zoology, Harvard University 1: 1-200, pis 1-2. Matsumoto, H. (1917). A monograph of Japanese Ophiuroidea, arranged according to a new classification. Journal of the College of Science, Imperial University Tokyo 38(2): 1-408, pis 1-7. Miiller, J. & Troschel, EH. 1842. System der Aster iden. Von Friedrich Vieweg und Sohn, Braunschweig. 134 pp., 12 pis. O’Hara, T.D., Byrne, M., & Cisternas, P. 2004. The Ophiocoma erinaceus complex: another case of cryptic speciation in echinoderms: pp. 537-42 in Heinzeller, T. & Nebelsick, J.H. (eds). Echinoderms: MUnchen: Proceedings of the 11th International Echinoderm Conference, Munich, Germany, 6-10 October 2003, A.A. Balkema, The Netherlands. Memoirs of Museum Victoria 65: 57-62 (2008) ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs/index.asp A new species in the genus Ophiomyxa from South-west Australian waters (Echinodermata: Ophiuroidea: Ophiomyxidae) Amanda M. Franklin and Timothy D. O’Hara Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia (tohara@museum.vic.gov.au) Abstract Franklin, A.M. and O'Hara, T.D. 2008. A new species in the genus Ophiomyxa from South-west Australian waters (Echinodermata: Ophiuroidea: Ophiomyxidae). Memoirs of Museum Victoria 65: 57-62. A new species is described from the genus Ophiomyxa collected from south-west Australia. It is distinguished by the lack of marginal plates, three arm spines, the uppermost reaching up to 2 segments in length, and a characteristic pattern of reduced dorsal arm plates. Current evidence suggests it is endemic to waters around 400m deep off the coast of south-western Australia. Keywords Echinodermata, Ophiuroidea, Ophiomyxa Introduction Knowledge regarding the diversity of ophiuroids from the south-west of Australia is rather limited. A recent survey coordinated by Australia’s Commonwealth Scientific and Industrial Research Organisation (CSIRO) in 2005 sought to validate multiple use management frameworks and characterise marine ecosystems in this area. Epibenthic sleds were used to collect invertebrate epifauna. Through this sampling process, many ophiuroid specimens belonging to an undescribed species in the family Ophiomyxidae were collected. The Ophiomyxidae are a small taxonomically problematic family of ophiuroids that were originally placed with basket and serpent stars in the order Phrynophiurida (Matsumoto 1915), but more recently with the other simple-armed brittle- stars in the order Ophiurida (Smith et al. 1995, Janies 2001). The characteristic covering of thick skin and reduction of skeletal plates have been considered a primitive (Matsumoto 1917) or derived trait (Byrne pers. comm). This paper is the formal scientific description of the new species of Ophiomyxa Muller & Troschel, 1842. A suite of morphological characteristics are used to distinguish the new species including the presence and shape of dorsal arm plates, the length and number of arm spines and the location of ossicles. The material examined in this study is lodged in the Museum of Victoria, Melbourne (MV) and the Western Australian Museum (WAM). The abbreviation d.d. is used for disc diameter. Taxonomy Ophiomyxa crinita sp. nov. Figures 1 & 2 Material examined, (all from the RV Southern Surveyor expedition SS10/2005 to south-western Australia). Holotype: Off Lancelin, 31° 0.75'S, 114° 49.5'E, 394-393m, 1 Dec 2005, (stn 75), WAM Z21290. Paratypes: as holotype, MV F111566(n=199). Other material: Off Zuytdorp, 27°8.01'S, 112°45.06'E,414-405m,5 Dec 2005, (stn 105), MV FI 11 951, (2). Off Bunbury, 33° 0.504'S, 114° 34.26'E, 42 1-4 14m, 20 Nov 2005, (stn 13), F111561(103). D’Entrecasteaux, 35° 4.176'S, 115° 20.16'E, 378-379m, 21 Nov 2005, (stn 17), MV FI 1 1562(2). Off Albany, 35° 21.882'S, 118° 18.42'E, 398-407m, 23 Nov 2005, (stn 25), MV F111563(l). Two Rocks, 31° 37.08'S, 114°58.32'E,364-404m, 19Nov 2005,(stn4),MVF111559(15); 31° 36.528'S, 114° 58.86'E, 329-370m, 19 Nov 2005, (stn 6), MV FI 11560(2). Off Bunbury, 33° 0.582'S, 114° 34.2'E, 423-397m, 29 Nov 2005, (stn 67), MV F111564, (100). Perth Canyon, 31° 59.544'S, 115° 10.98'E, 508-478m, 29 Nov 2005, (stn 68), MV F111565(l). Off Jurien Bay, 29° 52.062'S, 114° 23.22’E, 414-401m, 2 Dec 2005, (stn 78), MV FI 11567(5); 29° 50.5 14'S, 114°21.72'E, 408-427m, 2 Dec 2005, (stn 80), MV FI 1 1568( 1); MV F111935(28). Off Abrolhos Islands, 28° 59.4’S, 113°45.9'E, 389-407m, 3 Dec 2005, (stn 90), MV FI 11661(5). Holotype description. 19 mm d.d. with slightly indented interradial margins. Disc covered with a thick, fairly smooth epithelium obscuring the limits of the radial shields, oral and adoral shields and oral plates (fig. 1C). No series of interradial marginal plates. Bursal slits 5 mm long, ending 1 mm before 58 Amanda M. Franklin and Timothy D. O’Hara the disc margin at the end of the 4 th arm segment, not bordered by spines or papillae. Arms are approximately 110 mm long and 3 mm wide at the base. The dorsal, ventral and lateral arm plates are hidden from view by a thick epithelium which extends to cover the arm spines but does not create any webbing (fig. 1B&C). Segments have 3 spines per side, reducing to 2 distally, positioned ventro-laterally. The dorsal-most spine resides on the lateral midline and is the widest (0.8 mm at the arm base). At the base of the arm the dorsal spine is longest (3.4 mm), about 2 segments long, the middle spine is just longer than a segment and the ventral spine is slightly shorter. Midway down the arm the dorsal spine shortens to about 1.5 segments long, the middle spine is slightly longer and the ventral spine equal to a segment. Distally, the ventral spine is longest; approximately a segment long. The dorsal spine is about 0.5 segments long and when the middle spine is present it is slightly shorter than a segment. Damage to the specimen has removed skin from some spines. Without skin, proximal spines are conical and smooth, gradually changing to become slightly serrated on 2 sides by mid-arm and becoming serrated and comb-like (but not hooked) on 1 side distally. No tentacle scales; the tube feet do not have a calcareous tube around the base. Each side of the jaw angle has 3 oral papillae which are separate from each other. Two jaw angles also bear a 4 th very small, triangular oral papilla, positioned distally. The middle papillae are 2 times higher than wide and generally triangular, sometimes rounded, in shape with denticulated tips. The innermost papillae are 2-3 times higher than wide, ovoid and also with a denticulated edge. The apical papillae are largest, generally about as high as they are wide, much rounder and not as denticulate. The distal oral tube feet are located in the mouth, near the ventral surface and are not covered by protecting scales. Paratypes. Range from 11 mm - 22 mm in d.d., juveniles resemble small adults. Dorsal and ventral skin from a 19 mm d.d. paratype were partially dissolved in bleach. This disc epithelium contained 2-3 layers of transparent, overlapping oval plates, ranging in size from 182 pm - 303 pm wide and 136 pm - 409 pm long, flat and smooth, showing varying degrees of perforation (fig. 2E). Perforations usually cover the entire plate but some are only perforated in the middle. The perforated plates are more common near the radial shields and the bursal slits. C-shaped ossicles are scattered throughout the stomach wall, occasionally in small clusters, typically 76 pm long and 38 pm wide (fig. 2F). Figure 1: Ophiomyxa crinita sp. nov. holotype (19 mm d.d.) in A, dorsal view; B. detail of dorsal disc; C, detail of ventral disc and mouthparts. A new species in the genus Ophiomyxa from South-west Australian waters (Echinodermata: Ophiuroidea: Ophiomyxidae) 59 Figure 2. Ophiomyxa crinita sp. nov.. A, dental plate; B, arm spines, i, dorsal at base of arm, ii, middle at proximal mid arm, iii, middle at distal mid arm, iv, ventral at tip of arm; C, dorsal view of arm, skin removed; D, ventral view of arm and mouth parts, skin removed; E, plates from disc skin; F, ossicles from stomach wall. Scale bars A-D: 1 mm, E&F: 0.1 mm. 60 Amanda M. Franklin and Timothy D. O’Hara Radial shields are approximately 2 mm long, 1/10 d.d., 3 times longer than wide, rectangular in shape, with curved edges and angled inwards at the proximal end. Ventrally, the adradial genital scale articulates with the radial shield, but it is not visible. The abradial genital scale is visible and shaped like a two pronged fork (fig. 2D). Proximal to the abradial scale is a line of small irregularly shaped, perforated plates terminating in an L-shaped plate partially overlapped by the oral shield. The dorsal arm plates do not extend onto the disc, are rhombic in shape, 2.25 mm wide and 1.15 mm long proximally, about 2 times wider than long. The lateral corners are rounded, the distal edge is convex, the proximal edges are slightly concave and the proximal tip is indented and overlain by another pentagonal plate, 0.7 mm wide and 0.5 mm long (fig. 2C). Proximal to this plate are two small, round plates which are slightly pointed on their inner edges. Residing on either side of the midline, they do not touch any other plates. Ventral plates, 1.20 mm long and 1.70 mm wide, are about 1.5 times wider than long, shaped like slightly flattened heptagons with proximal ends pointed and distal ends indented, the angles are rounded and edges slightly concave (fig. 2D). The proximal edge curves around the lateral plates and the tentacle pores. The 1 st ventral plate is shaped like a flattened hexagon, 3 times wider than long. Lateral arm plates extend from near the ventral midline to just past the lateral midline touching the edge of the dorsal plates. They bear a slight dome on the edge of the arm, the distal edge of which has three small articulation ridges, one for each arm spine. Oral shields are oval, with a small point proximally and slightly wider distally, about 2 times wider than long (fig. 2D). Adoral shields are pentagonal, about as wide as long, and follow the proximal and lateral edges of the oral shield and do not touch radially or interradially. Dental plates are divided by transverse fissures into 4 pieces each supporting a tooth about the same size and shape as the apical papillae, or first tooth. The middle two pieces are square with rounded lateral edges, the end two pieces are triangular, and all have curved corners (fig. 2A). Each has a rectangular groove in the middle where the tooth sits and the ventral piece also supports the apical papilla. Oral plates are about two times longer than high and are axe shaped in lateral view with the ventral edge curving up as it nears the jaw apex. Vertebrae have a V-shaped aboral groove and a smooth aboral surface. Variations from the holotype include a thicker, wrinklier epithelium, slightly pinker on the dorsal surface. Oral papillae number either 4 or 3 on all jaw angles and some may touch (fig. 2D). They vary in how triangular or broad they are, generally more broad apically. The dental plate may bear a 5 th pointed, small tooth dorsally. Colour ( Live specimens ). The dorsal disc is peach-brown in colour, slightly lighter on the arms and fading to cream at the tips. There are some tan and some cream spots on the dorsal disc surface, with the cream spots being smaller. The ventral disc surface, mouth frame and spines are a bright white and the tube feet are a transparent white. Tests for bioluminescence were negative (Mallefet & O’Hara, unpublished data). Colour ( Preserved in ethanol ). The disc is pale pink-brown dorsally and white ventrally. The middle of the disc is pale grey due to the internal organs. The tube feet are tan whilst the mouth frame is cream. The spines when covered with skin are cream, but transparent and glassy where the skin has been removed. Etymology, crinita (Latin, f.), long haired, in reference to the relatively long arm spines. Distribution. South-western Australia from off Albany to Zuytdorp, 329-508 m. Remarks. This species belongs to the genus Ophiomyxa due to the presence of a thick skin covering the disc and arms, denticulate glassy oral papillae, fragmented dorsal arm plates, and the second oral tentacle opening within the jaw slit (Matsumoto 1917; Fell 1960). Within Ophiomyxa, the species falls into a group of species previously distinguished as the genus Ophiodera Verrill, 1899 that are characterised by the absence of a row of marginal interradial plates that run between the pairs of radial shields. Ophiodera is not currently recognised, as the designated type species O. serpentaria has the intermediate condition of a few rudimentary marginal plates near the radial shields (Mortensen 1927). However, there may be some merit in a grouping of these species, with other potential synapomorphies including similar shaped oral plates and long pointed arm spines that become serrated on distal segments. Whatever the status of Ophiodera, these characters are useful diagnostically within Ophiomyxa. Of the species without marginal plates, Ophiomyxa crinita is most similar to Ophiomyxa neglecta (Koehler, 1904), sharing a similar number of arm spines, similar shaped dorsal and ventral plates and denticulate oral papillae. However, Ophiomyxa neglecta has small arm spines that are less than one segment long, ventral plates of similar length and width, radial shields about 1/5 of the disc diameter, and C-shaped ossicles in the disc and arm epithelium (Koehler 1922). Ophiomyxa anisacantha H.L. Clark, 1911, Ophiomyxa bengalensis Koehler, 1897, Ophiomyxa compacta (Koehler, 1905), and Ophiomyxa punctata (A.H. Clark, 1952) all have four, often five arm spines (see also Irimura 1982; Imaoka et al. 1990). The dorsal arm plates of O. anisacantha, O. bengalensis and O. punctata are divided into several small, oval scales whereas in O. punctata this only occurs on the first 2 segments and the other segments bear no plates. The arm spines of O. anisacantha are shorter than O. crinita and also differ in having the ventral arm spine of equal length to the dorsal spine. Ophiomyxa punctata has short black lines parallel to the genital slits and O. compacta has no genital scale and oral papillae that are not denticulated. All of these species are from the tropical Indo-Pacific region. Two other species have been referred to Ophiodera from the Atlantic Ocean: O. serpentaria Lyman, 1883 and O. stimpsonii (Lyman, 1875). They both have at least a few marginal disc plates and lack dorsal arm plates (Verrill 1899; Paterson 1985). Ophiomyxa stimpsonii has more than four short arm spines and five oral papillae. Ophiomyxa serpentaria has lateral plates that may be fused to the ventral plates (Verrill 1899). The new species is sympatric with the widespread Indo- Pacific species Ophiomyxa australis Liitken, 1869, which can be distinguished by the presence of the marginal row of disc A new species in the genus Ophiomyxa from South-west Australian waters (Echinodermata: Ophiuroidea: Ophiomyxidae) 61 plates, up to seven short arm spines often webbed with skin, and a different pattern of dorsal plate fragmentation (Mortensen 1924; Irimura 1982). Acknowledgements We thank Alan Williams (CSIRO, voyage leader), Karen Gowlett-Holmes (CSIRO, voyage photographer) and other scientific staff and crew of the Southern Surveyor for the successful SS10/2005 expedition, Jerome Mallefet (Catholic University of Louvain, Belgium) for bioluminescent tests; David Staples (Museum Victoria) for the type photographs, and David Collins (Museum Victoria) for assistance with the electronic line drawings, and Maria Byrne (University of Sydney) for discussions about the systematic position of the Ophiomyxidae. References Clark, A.H. 1952. Echinoderms from the Marshall Islands. Proceedings of the United States National Museum, 102: 265-303. Clark, H.L. 1911. North Pacific ophiurans in the collection of the United States National Museum. Bulletin of the United States National Museum, 75: 1-302, figs 1-144. Fell, H.B. 1960. Synoptic keys to the genera of Ophiuroidea. Zoological Publications of the Victoria University Wellington, 26: 1-44, 6 figs. Imaoka, T., Irimura, S., Okutani, T., Oguro, C., Shigei, M. & Horikawa, H. 1990. Echinoderms from continental shelf and slope around Japan Vol. 1. Japan Fisheries Resource Conservation Association, Tokyo: 159 pp. Irimura, S. 1982. The Brittle-stars of Sagami Bay. Biological Laboratory, Imperial Household, Japan: 95 pp, 15 pis. Janies, D. 2001. Phylogenetic relationships of extant echinoderm classes. Canadian Journal of Zoology 79: 1232-50. Koehler, R. 1897. Echinodermes recueillis par T Investigator dans l’Ocean Indien. I. Les ophiures de mer profonde. Annales des Sciences Naturelles, Zoologie, 8: 277-372, pis 5-9. Koehler, R. 1904. Ophiures de mer profonde. Siboga-Expeditie, 45: 1-176, pis 1-36. Koehler, R. 1905. Ophiures littorales. Siboga-Expeditie Monographs, 45: 1-142, pis 1-18. Koehler, R. 1922. Contributions to the biology of the Philippine Archipelago and adjacent regions. Ophiurans of the Philippine seas and adjacent waters. Bulletin of the United States National Museum, 100: 1-486, pis 1-103. Liitken, C.F. 1869. Addimenta ad historiam Ophiuridarum. 3. Beskrivende og kritiske Bidrag til Kundskab an Slangestjernerne. Kongelige Danske Videnskabernes Selskabs Skrifter, 5(8): 22-109. Lyman, T. 1875. Zoological results of the Hassler Expedition. 2. Ophiuridae and Astrophytidae. Illustrated Catalogue of the Museum of Comparative Zoology, Harvard University, 8: 1-34, 5 pis. Lyman, T. 1883. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Caribbean Sea in 1878-79, and among the Atlantic coast of the United States during the summer of 1880, by the U.S. Coast Survey steamer Blake, Commander J.R. Bartlett, U.S.N., commanding. XX. Report on the Ophiuroidea. Bulletin of the Museum of Comparative Zoology, Harvard University, 10: 227-287, pis 1-8. Matsumoto, H. 1915. A new classification of the Ophiuroidea: with descriptions of new genera and species. Proceedings of the Academy of Natural Sciences of Philadelphia, 67: 43-92. Matsumoto, H. 1917. A monograph of Japanese Ophiuroidea, arranged according to a new classification. Journal of the College of Science, Imperial University Tokyo, 38: 1-408, figs 1-100, pis 1-7. Mortensen, T. 1924. Echinoderms of New Zealand and the Auckland- Campbell Islands. II. Ophiuroidea. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 77: 91-177, figs 1-36, pis 3-4. Mortensen, T. 1927. Handbook of the Echinoderms of the British Isles. Oxford University Press, London: 471 pp. 269 figs. Muller, J. & Troschel, F.H. 1842. System der Asteriden. Friedrich Vieweg und Sohn, Braunschweig: 134 pp. 12 pis Paterson, G.L. J. 1985. The deep-sea Ophiuroidea of the North Atlantic Ocean. Bulletin of the British Museum ( Natural History), 49: 1-162, figs 1-59. Smith, A.B., Paterson, G.L .J. & Lafay, B. 1995. Ophiuroid phylogeny and higher taxonomy: morphological, molecular and palaeontological perspectives. Zoological Journal of the Linnean Society of London, 114: 213-243. Verrill, A.E. 1899. Report on the Ophiuroidea collected by the Bahama expedition in 1893. Bulletin of the Laboratories of Natural History of the State of Iowa, 5: 1-88, pis 1-8. Memoirs of Museum Victoria 65: 63-70 (2008) ISSN 1447-2546 (Print) 1447-2554 (On-line) http://www.museum.vic.gov.au/memoirs/index.asp Deep-sea majoid crabs of the genera Oxypleurodon and Rochinia (Crustacea: Decapoda: Brachyura: Epialtidae) mostly from the continental margin of Western Australia Bertrand Richer de Forges 1 and Gary C.B. Poore 2 1 UMR 7138/UR R148 “Systematique, Adaptation, Evolution”, Institut de Recherche pour le Developpement, B.R A5, 98848, Noumea Cedex, New Caledonia (bertrand.richer-de-forges@noumea.ird.nc) 2 Museum Victoria, GPO Box 666, Melbourne, Victoria, 3001, Australia (gpoore@museum.vic.gov.au) Abstract Richer de Forges, B., and Poore, G.C.B. 2008. Deep-sea majoid crabs of the genera Oxypleurodon and Rochinia (Crustacea: Decapoda: Brachyura: Epialtidae) mostly from the continental margin of Western Australia. Memoirs of Museum Victoria 65: 63-70. Two species are newly described from the continental margin of Western Australia: Oxypleurodon wilsoni and Rochinia annae. Oxypleurodon luzonicum, Rochinia carinata, R. pulchra, R. sibogae and R. strangeri are newly reported from Western Australia. Rochinia fultoni and R. mosaica from south-eastern Australia are refigured. Keywords Decapoda, Brachyura, Epialtidae, new species, Introduction The decapod crustacean fauna along the south-western and central continental margin of Western Australia has been recently revealed to be highly diverse and novel (Poore et ah, 2008). These discoveries result from recent sampling, part of a project mounted by CSIRO Marine and Atmospheric Research (CMAR) and Museum Victoria entitled “Mapping benthic ecosystems on the deep continental shelf and slope in Australia’s South West Region”. This contribution on majoid crabs is based on the collections made during the southern phase of this project (cruise SS10-2005) and on those taken during a second phase along the northern Western Australian continental margin (cruise SS05-2007). Our classification of the Majoidea follows Ng et al. (2008). Poore et al. (2008) reported on 14 species of Epialtidae (3 probably new species), two species of Hymenosomatidae (one new), 20 species of Inachidae (two new) and 13 species of Majidae (three new). Of these 49 species, nine were reckoned to be new records of Indo-West Pacific species for Australia. This paper reports only on the epialtid genera Oxypleurodon Miers, 1886, and Rochinia Milne Edwards, 1875, other members of this family from these collections being relatively well known. We remark on other species of Rochinia in the collections of Museum Victoria (NMV). Types are lodged in the Western Australian Museum (WAM) and Museum Victoria. Indian Ocean, Australia Measurements are given as greatest length (without pseudorostrum) and greatest width. Epialtidae MacLeay, 1838 Remarks. Poore et al. (2008) listed 14 species in this family from the south-western Australian collections. Of these, Austrolibinia gracilipes (Miers, 1879), Hyastenus convexus Miers, 1884, Naxioides robillardi (Miers, 1882), N. taurus (Pocock, 1890), N. tenuirostris (Haswell, 1880), Phalangipus filiformis Rathbun, 1916, and P. hystrix (Miers, 1884) were new records for southern or all Western Australia. They also reported Griffinia lappacea (Rathbun, 1918) and Lahaina agassizii (Rathbun, 1902) which were previously known. One specimen, which could not be identified at the time, belongs to a species of Thacanophrys. Oxypleurodon Miers, 1886 Remarks. Oxypleurodon was redefined and separated from other similar genera by Tavares (1991) and several species from the Indo-West Pacific were reviewed by Richer de Forges (1995). Ng and Richer de Forges (2007) listed the 17 species known. Davie (2002) included the genus in the subfamily Pisinae of family Majidae. Following Ng et al. (2008) it is now in the family Epialtidae MacEeay, 1838, included inside the superfamily Majoidea. One species has been previously recorded from Australia, O. stimpsoni Miers, 1886, from Queensland (Davie, 2002). Here, a second species is newly 64 Bertrand Richer de Forges and Gary C.B. Poore recorded for Australia and a third species is described as new from Western Australia. Oxypleurodon luzonicum (Rathbun, 1916) Figure la Sphenocarcinus luzonicus Rathbun, 1916: 539.— Estampador, 1937: 552 . —Estampador, 1959: 112,-Griffin, 1976: 211, fig. 11a.- Serene and Vadon, 1981: 124, pi. 4E.— Guinot and Richer de Forges, 1986a: 138, fig. 19A, B, 21C, D, pi. 8 figs A-F.— Guinot and Richer de Forges, 1986b: 29.— Richer de Forges, 1992: 4. Rochinia luzonica.— Griffin and Tranter, 1986a: 180.— Tavares, 1991: 161.— Webber and Richer de Forges, 1995: 514. Oxypleurodon luzonicus.— Richer de Forges, 1995: 48, fig. IB, pi. 2A. Oxypleurodon luzonicum.— Ng and Richer de Forges, 2007: 63. Material examined. Australia. WA. Feveque F27 transect (15°00.52'S-14°59.05'S, 121°38.08'E -121 o 39°10'E), 205-211 m, 25 Jun 2007 (stn SS05/2007 099), NMV J58221 (2 females 11.7 x 9.7 mm, 11.8 x 10.1 mm; juvenile 5 x 3.9 mm). Remarks. These specimens from the northern part of the Western Australian continental slope fit well with the description of O. luzonicum from the Philippines. These records extend the distribution of the species considerably south, to 15°S in the western Indian Ocean. The species’ distribution does not overlap with that of another Oxypleurodon from the southern part of the Western Australian slope which is described below as a new species. Oxypleurodon wilsoni sp. nov. Figure lb Rochinia aff. luzonica.— Poore et al., 2008: 56 (colour fig.). Material examined. Holotype: Australia. WA. Off Two Rocks (31°36.32'S-31°37.02'S, 114 0 58.52'E-114°58 16’E), 329-370 m, 19 Nov 2005 (stn SS10/2005 006), WAM C400529 (ovigerous female 15.7 x 15.4 mm). Paratypes. Collected with holotype, WAM C400530 (ovigerous female, 16.4 x 14.0 mm). WA. Off Abrolhos (29°00.46'S-29°01.23 , S, 1 13°46.44’E -113°47.06'E), 419-439 m, 03 Dec 2005 (stn SS10/2005 088), NMV J54070 (male, 18.5 x 19.6 mm). Off Two Rocks (31°37.05'S-31°37.23'S, 114°58.19'E -115°14.39'E), 364-404 m, 19 Nov 2005 (stn SS10/2005 004), NMV J54069 (3 males, 14.4 x 13.5 mm, 13.9 x 13.4 mm, 9.6 x 8.8 mm). Diagnosis. Carapace to 18.5 mm, long, pyriform. Carapace, including pseudorostral spines, covered with short setae. Carapace bearing several elevated plates and tubercles arranged as follows: 1 cardiac plate, round and forming a conical point in its middle; 2 branchial plates, thin, sharp and pointing laterally; 2 epibranchial plates, thin, sharp and oriented obliquely; 1 ovoid mesogastric plate; 2 hepatic plates touching the postocular plates forming together a L-shape; 1 small subbranchial oblong plate on the lateral border; 2 supraocular plates, sharp anteriorly; 1 anterogastric tubercle; 1 tubercle on each side of mesogastric plate; posterior border of the carapace thick, forming medially a large tooth pointing backward. Pseudorostral spines long and sharp, diverging in a V. Eyes small, completely inserted in the orbits, the postocular plate forming a cup. Basal antennal article fused with carapace. Cheliped shorter than pereopod 2. Articles of ambulatory legs cylindrical. Female abdomen of 7 segments. Distribution. South-western WA slope (29°-31.5°S), 329-439 m depth. Etymology. Dedicated to Dr Robin Wilson in recognition of his enthusiasm and hard work during the cruise when this species was collected, and at all times. Remarks. Oxypleurodon wilsoni sp. nov. belongs in the group of species of Oxypleurodon having long and sharp branchial plates: O. luzonicum (Rathbun, 1916), O. stuckiae (Guinot and Richer de Forges, 1986b), O. karubar Richer de Forges, 1995, and O. lowryi (Richer de Forges, 1992). In O. wilsoni the supraocular plate is sharp anteriorly whereas it is rounded in O. luzonicum. The cardiac plate is small, round and elevated in the middle (round and flat in O. luzonicum ). The epibranchial plates are oblong, pointing externally in O. wilsoni but small and triangular in O. luzonicum. The mesogastric plate is ovoid but lozenge-shaped in O. luzonicum. Between the mesogastric spine and the hepatic spine lies a small tubercle absent in O. luzonicum. O. wilsoni is distinguishable from O. stuckiae by the presence of a supraocular spine. The pseudorostral spines are longer and less diverging in O. wilsoni than in O. stuckiae. An anterior gastric tubercle in O. wilsoni is not seen in O. stuckiae. O. wilsoni differs from O. karubar in the following characters: the epibranchial plate is straight but curved in O. karubar, the pseudorostral spines are straight but enlarged basally in O. karubar-, the anterior part of the supraocular plate is spiniform in O. wilsoni but rounded in O. karubar, the hepatic plate touches the postocular plate forming together an L-shape in O. wilsoni but these are several disjointed pieces in O. karubar. The shape of the carapace is pyriform in O. wilsoni while it is large and rounded posteriorly in O. lowryi. The supraocular plate points forward in O. wilsoni but laterally in O. lowryi. The branchial spines are straight in O. wilsoni while they are curved in O. lowryi. The ambulatory legs of the holotype and paratype from the same sample are mixed and it is not possible to be precise about the relative lengths of pereopods. The species was collected over narrow latitudinal and depth ranges. Rochinia Milne-Edwards, 1875 Remarks. Rochinia was redefined by Tavares (1991) updating the extensive revision by Griffin and Tranter (1986a) who provided a key to 29 species of a more inclusive genus. Davie (2002) included the genus in the subfamily Pisinae of family Majidae. Five species have been previously recorded from Australia (Davie, 2002). Ng and Richer de Forges (2007) listed the 24 species known from the Indo-West Pacific and ten from the north-western and western Atlantic and eastern Pacific Oceans. Here, we add three species described originally from the Indo- West Pacific to the Australian fauna, comment briefly on three Deep-sea majoid crabs from Western Australia 65 Figure 1. Lateral and dorsal views, scale = 5 mm. a, Oxypleurodon luzonicum (female, NMV J58221). b, Oxypleurodon wilsoni sp. nov. (holotype, WAM C400259). c, Rochinia annae sp. nov. (holotype, WAM C400531). d, Rochinia carinata (male, NMV J53872). 66 Bertrand Richer de Forges and Gary C.B. Poore others, and describe a new species from Western Australia. Rochinia annae sp. nov. Figures lc, 3 Rochinia sp. MoV51 19.— Poore et al., 2008: 56. Rochinia fultoni.— Poore et al., 2008: 56 (colour fig.). Material examined. Holotype: Australia, WA. Off Two Rocks (31°37.05'S-31°37.23'S, 114 0 58.19'E-115 0 14.39'E), 364-404 m, 19 Nov 2005 (SS10/2005 004), WAM C400531 (male, 11.9 x 8.6 mm). Paratypes. Same data as holotype. WAM C400532 (11 specimens), NMV J54179 (87 specimens, smallest is a male of 7.3 x 4.6 mm). Othermaterial. Australia. WA.OffKalbarri(27°55.43'S-27°56. 01’S, 113°08.17'E-113°08.38'E), 252-253 m, 04 Dec 2005 (stn SS10/2005 099), NMV J54058 (1 male). Off Two Rocks (31°36.32'S-31°37.02'S, 114 0 58.52'E-114°58.16'E), 329-370 m, 19 Nov 2005 (stn SS10/2005 006), NMV J54178 (2 ovigerous females 10.5 x 8.2 mm, 8.6 x 6.9 mm). Jurien Bay (29°52.04'S-29 o 52.26'S, 114 0 23.13'E-114°23.53'E), 414-401 m, 02 Dec 2005 (stn SS10/2005 078), NMV J54253 (7 juveniles). Off Lancelin (31°00.45'S-31°00.17'S, 1 14°49. 3 0 'E-l 14°49. 23 'E) , 394-393 m, 01 Dec 2005 (stn SS10/2005 075), NMV J54238 (1 male). Off Bunbury (33°00.30'S-33°00.07'S, 114°34.16'E-114°34.30'E), 421-414 m, 20 Nov 2005 (stn SS10/2005 013), NMV J54073 (male 8.3 x 5.7 mm), J54238 (1 female 6.0 x 4.6 mm). Off D’Entrecasteaux (35°04.11'S-35°04.14'S, 115 o 20.10'E-115°20.53'E), 378-379 m, 21 Nov 2005 (stn SS10/2005 017), NMV J54916 (1 male 9.5 x 10.7 mm, 2 ovigerous females 8.4 x 6.1 mm, 10.5 x 7.4 mm, 2 females 7.6 x 4.7 mm, 7.2 x 5.2 mm). Diagnosis. Carapace to 11.9 mm long, pyriform with long diverging pseudorostral spines (one -third length of carapace). Dorsal surface spiny, longest spines as follows: 2 longest and thin branchial spines twice as long as other spines; 1 intestinal spine; 1 long cardiac spine; 1 long mesogastric spine; 2 hepatic spines; in protogastric area, a line of 3 short blunt spines, 2 other spines alongside mesogastric spine; 2 short spines on epigastric area; 2 short spines on lower side of cardiac area. Cheliped of male short with chelae inflated; merus cylindrical, smooth, with 5 blunt teeth on inferior border, upper anterior angle forming 2 teeth; carpus short and triangular, forming a carina at its interior border; propodus inflated and carinate on upper border; fingers thin with serrulated interior border; dactylus border forming a bump in its first third. Ambulatory legs long and thin; articles cylindrical, pereopod 2 longer than pereopod 1 (pereopod 2 merus reaching little beyond ridge on reflexed carpus of pereopod 1). Eyes small, protected by cupped postocular tooth; large supraocular tooth. Basal antennal article with long tooth on its antero- external angle; article fused to carapace. Antenna flagellum longer than pseudorostrum; antennule fossae sharp anteriorly. Border of buccal frame expanded laterally. Abdomen of 7 segments. Pleopod 1 of pisid type, distally expanded to an oblique truncate spinose margin, 16 setae in groove (fig. 3). Distribution. South-western WA slope (27°-35°S), 252-424 m depth. Etymology. The species is named in honour of Anna McCallum, who sorted and made preliminary identifications of many of the Decapoda of the Western Australian cruises. Remarks. Of the 35 species described in the genus Rochinia few have long branchial spines. The only species comparable to R. annae sp. nov. are: R. pulclira (Miers, 1886), R. riversandersoni (Alcock, 1895), R. sibogae Griffin and Tranter, 1986, R. galathea Griffin and Tranter, 1986, R. griffini Davie and Short, 1989, and R. paulayi Ng and Richer de Forges, 2007. R. galathea has a long branchial spine but a long and flat hepatic spine, very different from the spine of R. annae. R. sibogae has two very long branchial spines pointing laterally but two other long spines, one intestinal and one cardiac (one long gastric spine in R. annae). Also, the postocular tooth is distinct from the hepatic spine in R. annae whereas it is a fused plate in R. sibogae. R. pulchra has long branchial spines but a different pattern in the other long spines of the carapace: four spines in the median line (three in R. annae). R. pulchra possesses pseudorostral spines about three-quarters as long as the carapace but these are one third as long in R. annae. R. riversandersoni is a large species with long branchial spines and long epibranchial spines. R. annae has only a granule at this place. R. paulayi and R. griffini are also large species, very spiny and showing a totally different arrangement of spines from that in R. annae. Although there is abundant material no specimen is intact and it is not possible to be precise about the relative lengths of pereopods. The species occurs over a wide geographic range along the south-western WA coast. Rochinia carinata Griffin and Tranter, 1986 Figure Id Rochinia carinata Griffin and Tranter, 1986a: 178, figs 56, 64e, f, pi. 12.— Ng and Richer de Forges, 2007: 62. Rochinia sp. MoV 5136.— Poore et al., 2008: 56. Material examined. Australia. WA. Off Barrow Island (21°00.24'S-21°00.02'S, 114 o 22.52'E-114 o 22.30'E), 399-408 m, 13 Dec 2005 (stn SS10/2005 172), NMV J53872 (1 male 28.2 x 20.7 mm). Off Ningaloo North (21°58.13'S-21°58.45'S, 113 0 47.35'E-113°47.28'E), 356-324 m, 11 Dec 2005 (stn SS10/2005 157), NMV J53873 (1 female juvenile 14.2 x 9.9 mm). Off Red Bluff (23°59.12'S-23°59.43'S, 112 o 32.02'E-112°31.44'E), 411 m, 08 Dec 2005 (stn SS10/2005 130), NMV J54067 (1 juvenile 10.5 x 6.5 mm). Remarks. Rochinia carinata was described from the Kei (Kai) Islands in Indonesia and has not been recorded anywhere else since. The specimens from the south-western Australian coast differ from the typical material from Indonesia (Griffin and Tranter, 1986a: pi. 12). The merus of pereopods 2 and 3 is cylindrical in cross section vs carinate in the Indonesian specimens. The mesogastric plate (islet) looks more ovoid in R. carinata from Western Australia whereas it is more rounded in the typical material. The pseudorostrum spines are more elongated in the specimens from south-western Australia than in the specimens from Indonesia. The species’ range is extended from 5°S in Indonesia to 24°S, near Shark Bay, WA. The depth range in Indonesia is 204-325 m and 324-411 m in WA. Deep-sea majoid crabs from Western Australia 67 Figure 2. Lateral and dorsal views, scale = 5 mm. a, Rochinia fultoni (female, NMV J4730). b, Rochinia pulchra (female, NMV J55947). c, Rochinia sibogae (male, NMV J58142). d, Rochinia strangeri (female, NMV J55427). Bertrand Richer de Forges and Gary C.B. Poore Figure 3. Rochinia annae sp. nov. Male left pleopod 1 (paratype WAM C400531). a, anterior view, b, detail of tip, anterior view, c, detail of tip, posterior view. Rochinia fultoni (Grant, 1905) Figure 2a Hyastenus Fultoni Grant, 1905: 313, pi. 11 fig. 1. Rochinia fultoni.— Griffin, 1966: 280.— Griffin and Tranter, 1986a: 176.— Poore, 2004: 387, fig. 1 18f. — Ng and Richer de Forges, 2007: 62. Material examined. Australia. Tas. Eastern Bass Strait, 100 km NE of North Point, Flinders I., (38°52.36'S, 148°25.12'E), 140 m, 15 Nov 1981 (stn BSS 170 S), NMV J4730 (2 ovigerous females 15.0 x 10.0 mm, 11.3 x 7.5 mm; male 11.7 x 7.0 mm). 70 km ENE of North Point, Flinders I. (39°28.24'S, 148°41.48'E), 110 m, 28 Mar 1979 (stn BSS 35), NMV J23091 (juvenile). 37 km NE of Cape Tourville (41°52.52'S-41°59.34'S, 148°37.56'E-148°31 08'E), 124 m, 30 Oct 1988 (stn SLOPE 85), NMV J16059 (male). Vic. W of Cape Nelson, 183 m, 06 Jun 1969, NMV J8610 (male, ovigerous female) Remarks. This species has already been mentioned from the east and south coasts of Australia. Most records in Museum Victoria are from the continental margin of eastern Bass Strait. Rochinia mosaica (Whitelegge, 1900) Pugettia mosaica Whitelegge, 1900: 141, pi. 35 figs 5-7. Doclea profunda Rathbun, 1918: 16, pi. 7 figs 1, 2.— Hale, 1927: 134, fig. 134. Rochinia mosaica.— Griffin and Tranter, 1986a: 185, figs 58, 62e, f.— Griffin and Tranter, 1986b: 363.— Poore, 2004: 387.— Ng and Richer de Forges, 2007: 62. Material examined. Australia. Tas. Eastern Bass Strait, 100 km NE of North Point, Flinders I. (38°52.36'S, 148°25.12'E), 140 m, 15 Nov 1981 (stn BSS 170 S), NMV J4731 (female 9.3 x 6.6 mm). Vic. Eastern Bass Strait, S of Waratah Bay (38°59.54'S, 146°00'E), 64 m, 26 Nov 1973 (stn K7-73-63), NMV J12157 (2 males 13.2x9.4 mm, 12.7x8.7 mm). Central Bass Strait, 100 km SSE of Cape Liptrap (39°45.54’S, 145°33.18'E), 74 m, 13 Nov 1981 (stn BSS 156), NMV J10642 (juvenile); NMV J4732 (female 8.1 x 5.1 mm). SA. Great Australian Bight (33°15.52'S-33 o 16.03'S, 130°37.50'E-130°37 07'E), 139-141 m, 09 May 2000 (stn SS01/00 334), NMV J52198 (3 males 8.5 x 5.2 mm, 7.3 x 4.8 mm, 7.2 x 4.3 mm). Great Australian Bight (33 o 16.00'S-33 o 16.36'S, 130°43.09'E-130°48.38'E), 134—130 m, 10 May 2000 (stn SS01/00 351), NMV J52196 (2 ovigerous females 11.3 x 7.7 mm, 10.9 x 7.4 mm). Great Australian Bight (33 o 16.00'S-33 o 16.01'S, 130°36.35'E-130°38.02'E), 143-140 m, 10 May 2000 (stn SS01/00 350), NMV J52197 (3 males 9.1 x 5.8 mm, 9.4 x 5.9 mm, 7.7 x 4.5 mm; ovigerous female 9.1 x 5.7 mm; female 7. 1 x 4.5 mm; juvenile 4.8 x 3.3 mm). Remarks. This material from south-eastern Australia fits well with the redescription of R. mosaica by Griffin and Tranter (1986a). Whitelegge described this species in the genus Pugettia in which the carapace has a similar pattern but the first pleopod of Pugettia is very different from that of Rochinia. The records are well within the reported distribution of the species from central Queensland, through Bass Strait to the Great Australian Bight. The most westerly is 130°W near the SA-WA border. Rochinia pulchra (Miers, 1886) Figure 2b Anamathia pulchra Miers, 1886: 26, pi. 4 figs la-c. Scyramathia pulchra.— Alcock, 1895: 202.— Doflein, 1904: 84, pi. 27 fig. 12. Rochinia pulchra.— Sakai, 1938: 278, fig. 35, pi. 37 fig. 4.— Sakai, 1976: 223, pi. 79 fig. 1.— Serene and Lohavanijaya, 1973: 56, figs 119-122, pi. 11A.— Takedaand Kurata, 1977: 144, fig. 4a, b.— Griffin, 1976: 210.— Griffin and Tranter, 1986b: 363.— Davie and Short, 1989: 181.— Ng and Richer de Forges, 2007: 62. Not Rochinia pulchra.— Griffin and Tranter, 1986a: 187 (part, NSW specimen) ( =Rochinia griffini Davie and Short, 1989) Material examined. Australia. WA. Ashmore L30 transect (12°31.46'S-12°30.50'S, 123 0 25.38'E-123°25.22'E), 401-404 m, 07 Jul 2007 (stn SS05/2007 192), NMV J58172 (female 17.3 x 10.7 mm). Kulumburu L29 transect (13°15.54'S-13°16.2rS, 123°22.27'E-123°21.24'E), 394-390 m, 07 Jul 2007 (stn SS05/2007 180), NMV J58025 (female 31.5 x 20.1, male 34.5 x 21.7 mm). Leveque L27 transect (14°49.02'S-14°48.32'S, 121 0 27.33'E-121°29.34'E), 407-392 m, 27 Jun 2007 (stn SS05/2007 107), NMV J55947 (female 23.3 x 16.6 mm). Leveque L27 transect (14 o 51.12’S-14 o 50.43'S, Deep-sea majoid crabs from Western Australia 69 121°25.53'E-121°27.0rE), 403-396 m, 03 Jul 2007 (stn SS05/2007 144), NMV J58220 (ovigerous female 28.7 x 21.9 mm). Lacepede L26 transect (15°47.34'S-15°48.30'S, 121 o 03.30'E-121°02.53'E), 119-111 m, 01 Jul 2007 (stn SS05/2007 129), NMV J55681 (male 12.9 x 10.2 mm). Remarks. This species, characterised by its very long and numerous spines, has already been recorded from the north- west slope of Australia. The other records are from Japan to the Philippines. All these specimens are from extreme north WA. Griffin and Tranter (1986a) tentatively identified a female from NSW as this species but it was described as Rochinia griffini Davie and Short, 1989. Rochinia sibogae Griffin and Tranter, 1986 Figure 2c Rochinia sibogae Griffin and Tranter, 1986b: 363, fig. 12. Rochinia riversandersoni.— Griffin, 1976: 211.— Griffin and Tranter, 1986a: 187.— Davie and Short, 1989: 181.— Ng and Richer de Forges, 2007: 62. Mtften'tf/examme