Deep-sea fauna of European seas: An annotated species check-list of benthic invertebrates living deeper than 2000 m in the seas bordering Europe. Holothuroidea

Gordon Paterson

Fifteen morphotypes of deep-sea holothurians were documented by photography or videography at depths of 596–2,566 m, using Remote Operated Vehicles (ROV) video surveys and towed camera transects, during hydrocarbon exploratory activities in the Colombian Southern Caribbean. Most of the morphotypes were identified to the species level based on the images. The species belong to four orders, Apodida (1 species), Persiculida (3 species), Elasipodida (8 species), and Synallactida (3 species). Four species, three genera, and three families are reported for the first time in the Colombian Caribbean Sea. Some of the reports also represent first records for the Caribbean Sea and the Atlantic Ocean.

The holothuroid family Rhopalodinidae currently contains three genera and 16 species, all restricted in their distribution to the East Atlantic Ocean from Senegal to the Cape of Good Hope. Their peculiar morphology is discussed and new information is presented on the southern African forms. A key for the identification of the 16 known species is provided. It is suggested that the appearance of the proboscis was probably a response to the shallow turbu-lent waters of the tropical East Atlantic. Because of the polytentaculate nature of most species, the branched nature of the tentacles of some species, the shape of the calcareous ring, the ambulacral restriction of the pedicels and the form of the body wall tables, the Rhopalodinidae could well be classified within the Dendrochirotida, close to the Phyllophorinae. However, this point of view is not supported by recent work based on analysis of rDNA and numerous morphological characters. According to this analysis Rhopalodinidae should...

Invertebrate Zoology, 2014, 11(1): 156–180 © INVERTEBRATE ZOOLOGY, 2014 Deep-sea fauna of European seas: An annotated species check-list of benthic invertebrates living deeper than 2000 m in the seas bordering Europe. Holothuroidea Andrey V. Gebruk1, Alexey V. Smirnov2 and Antonina V. Rogacheva1 1 P.P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Nakhimovsky Pr., 36, Moscow, 117997, Russia. E-mails: agebruk@ocean.ru antonina@ocean.ru 2 Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, St.-Petersburg, 199034, Russia. E-mail: sav_11@inbox.ru ABSTRACT: An annotated check-list is given of Holothuroidea species occurring deeper than 2000 m in the seas bordering Europe. The check-list is based on published data. The check-list includes 78 species. For each species synonymy, data on localities in European seas and general species distribution are provided. Station data are presented separately in the present thematic issue. How to cite this article: Gebruk A.V., Smirnov A.V., Rogacheva A.V. 2014. Deep-sea fauna of European seas: An annotated species check-list of benthic invertebrates living deeper than 2000 m in the seas bordering Europe. Holothuroidea // Invert. Zool. Vol.11. No.1. P.156–180. KEY WORDS: deep-sea fauna, European seas, Holothuroidea. Глубоководная фауна европейских морей: аннотированный список видов донных беспозвоночных, обитающих глубже 2000 м в морях, окружающих Европу. Holothuroidea А.В. Гебрук, А.В. Смирнов, А.В. Рогачева Институт океанологии им. П.П. Ширшова РАН, Нахимовский просп. 36, Москва, 117997, Россия. E-mails: agebruk@ocean.ru; antonina@ocean.ru Зоологический институт РАН, Университетская наб., 1, Санкт-Петербург 199034 Россия. E-mail: sav_11@inbox.ru РЕЗЮМЕ: Приводится аннотированный список видов Holothuroidea, обитающих глубже 2000 м в морях, окружающих Европу. Список основан на опубликованных данных. Список насчитывает 78 видов. Для каждого вида приведены синонимия, данные о нахождениях в европейских морях и сведения о распространении. Данные о станциях приводятся в отдельном разделе настоящего тематического выпуска. Как цитировать эту статью: Gebruk A.V., Smirnov A.V., Rogacheva A.V. 2014. Deepsea fauna of European seas: An annotated species check-list of benthic invertebrates living deeper than 2000 m in the seas bordering Europe. Holothuroidea // Invert. Zool. Vol.11. No.1. P.156–180. КЛЮЧЕВЫЕ СЛОВА: глубоководная фауна, европейские моря, Holothuroidea. Deep-sea fauna of European seas. Holothuroidea Phylum Echinodermata Class Holothuroidea Order Apodida Family Myriotrochidae Genus Acanthotrochus Danielssen et Koren, 1879 TYPE SPECIES: Acanthotrochus mirabilis Danielssen et Koren, 1879. COMPOSITION: 3 species (Belyaev, Mironov, 1981a). DISTRIBUTION: Arctic — 1, Pacific — 1, Antarctic — 1 species. Acanthotrochus mirabilis Danielssen et Koren, 1879 Acanthotrochus mirabilis Danielssen, Koren, 1879: 115, pl. 3–4, figs. 8–20; Danielssen, Koren, 1882: 35, pls. 5–6, figs. 8–20; Clark, 1908: 130, pl. 8 figs. 1–6; Djakonov, 1933: 158, fig. 83; Heding, 1935: 24, textfigs. 4, 5, pl. 3 figs. 3–10; Östergren, 1938: pl. 2 fig. 11; Belyaev, Mironov, 1981a: 522, fig. 1 figs. 1–3; Belyaev, Mironov, 1982: 108, fig. 17 (map); Madsen, Hansen, 1994: 127, figs. 91–93, map 38. LOCALITIES: Norwegian North-Atlanic Expedition (1876–1878), St. 295 (Danielssen, Koren 1879, 1882); “Sebastopol”, St. 1351 (Belyaev, Mironov, 1982), 1712, 1713, 1714, 1742, 2482 (Belyaev, Mironov, 1982 — only as dots on the map); “Polarstern” ARK XIII/2 cruise, St. 059GKG(bio+geo), 068-GKG(bio), 093-GKG(bio) (det. A.V. Smirnov, unpublished); BIOICE, St. 3637 (unpublished). DISTRIBUTION: Arctic: Norwegian and Greenland Seas, Litke Trough. DEPTH RANGE: 1090–3705 m. Genus Myriotrochus Steenstrup, 1851 TYPE SPECIES: Myriotrochus rinkii Steenstrup, 1851. COMPOSITION: 16 species (Belyaev, Mironov, 1982; Gage, Billett, 1986; Smirnov, 1999). DISTRIBUTION: Arctic — 2, Atlantic — 5, Atlantic and Pacific —2, Pacific — 6, Antarctic — 3 species. Myriotrochus bathybius H.L. Clark, 1920 Myriotrochus bathybius Clark, 1920: 126, pl. 4, fig. 3; Carney, Carey, 1976: 69; Gage et al., 1985: 202; Gage, Billett, 1986: 234, figs. 3–6, 7A, B, 9A, B, 18B; Harvey et al., 1988: 190; Smirnov, 1999: 17. REMARKS: The wheels of the North-East Atlantic specimens described by Gage, Billett (1986) differ somewhat from the wheels of the holotype of M. bathybius (see Smirnov, 1999, fig. 4). Only after examination of new findings of M. bathybius from the type locality (eastern tropical Pacific) it will be 157 possible to justify the determination of the NorthEast Atlantic material as M. bathybius. LOCALITIES: “Challenger” (1973–1985), St. ES 06, ES 27, ES 56, ES 118, ES 129, ES 137, ES 147, ES 152, ES 164, ES 169, ES 172, ES 180, ES 184, ES 185, SBC 188, ES 190, ES 204, ES 207, ES 231, AT 267, AT 282, ES 283, ES 289 (Gage et al. 1985; Gage, Billett, 1986; Harvey et al. 1988); “Discovery” (1977–1980), St. 9638#2, 9756#14; “Challenger, 50304, 50812#1, 50910 (Gage, Billett 1986); BIOICE, St. 2863 (unpublished data). DISTRIBUTION: Cosmopolitian. In the NorthEast Atlantic: Rockall Trough and Porcupine Seabight. Described from the East Tropical Pacific (Clark, 1920), also found in the North-East Pacific (Carney, Carey, 1976). DEPTH RANGE: 1800–4310 m. Myriotrochus giganteus H.L. Clark, 1920 Myriotrochus giganteus Clark, 1920: 127, pl. 4, fig. 4; Gage et al., 1985: 203; Gage, Billett, 1986: 239, figs. 7C, 8, 9C, 10–12, 24B; Harvey et al., 1988: 190. Myriotrochus sp. ex gr. macquoriensis — giganteus Belyaev, Mironov, 1982: 102, figs. 13, 14, pl. II figs. 6, 7. LOCALITIES: “Challenger” (1977–1985), St. AT 131, ES 137, ES 164, ES 169, ES 207, ES 231, ES 283 (Gage, Billett, 1986; Harvey et al., 1988). DISTRIBUTION: Eastern Tropical Pacific and North-East Atlantic. In the North-East Atlantic: Rockall Trough. DEPTH RANGE: 2898–3665 m. In the NorthEast Atlantic: 2898–2946 m. Myriotrochus clarki Gage et Billett, 1986 Myriotrochus clarki Gage, Billett, 1986: 247, figs. 7D, 9D, 13–17, 18A; Harvey et al., 1988: 190; Smirnov, 1999: 17; Rogacheva et al., 2013: 612–613, fig. 17l. Myriotrochus vitreus — Cherbonnier, 1970: 1269. Myriotrochus sp. — Gage et al., 1985: 203. LOCALITIES: “Challenger” (1975–1983), St. ES 34, ES 185, ES 197, ES 200, ES 218, ES 231, ES 232 (Gage et al., 1985; Gage, Billett, 1986; Harvey et al., 1988); “James Cook” ECOMAR, Sts. JC011/75, JC011/101, JC037/15, JC037/19, JC037/ 27, JC037/61, JC037/67, JC037/70, JC048/49, JC048/50, JC048/52 (Rogacheva et al., 2013). DISTRIBUTION: North-East Atlantic: Rockall Trough, coast of Spain and northern Mid-Atlantic Ridge. DEPTH RANGE: 480–2907 m. Genus Prototrochus Belyaev et Mironov, 1982 TYPE SPECIES: Myriotrochus zenkevitchi Belyaev, 1970. COMPOSITION: 20 species and subspecies (Belyaev, Mironov, 1982; Gage, Billett, 1986). 158 A.V. Gebruk et al. DISTRIBUTION: Arctic — 1, Atlantic — 5, Pacific — 10, Indian Ocean and Pacific — 1, Antarctic — 3 species. Prototrochus zenkevitchi rockallensis Gage et Billett, 1986 Prototrochus zenkevitchi rockallensis Gage, Billett, 1986: 252, figs. 7E, F; 18C–E; 19–23; 24A; Harvey et al., 1988: 190. P. zenkevitchi (Belyaev, 1970) subsp. — Gage et al., 1985: 204. LOCALITIES: “Challenger” (1973–1985), St. ES 06, ES 10, ES 34, ES 56, ES 57, ES 118, ES 129, ES 135, ES 137, ES 147, SBC 159, SBC 160, SBC 163, ES 169, ES 172, ES 176, ES 184, ES 185, ES 190, ES 197, ES 200, ES 204, ES 207, SBC 215, SB C216, ES 218, ES 231, ES 232, ES 283, ES 285, ES 289 (Gage et al., 1985; Gage, Billett, 1986; Harvey et al., 1988). DISTRIBUTION: North-East Atlantic: Rockall Trough, Porcupine Seabight, Whittard Canyon, continental slope of northern Bay of Biscay. DEPTH RANGE: 1000–2946 m. Prototrochus mediterraneus Belyaev et Mironov, 1982 Prototrochus mediterraneus Belyaev, Mironov, 1982: 89–90, fig. 5a, pl. 1 figs. 5, 6. LOCALITIES: “Vityaz”, St. 7930 (Belyaev, Mironov, 1982). DISTRIBUTION: Mediterranean Sea. DEPTH RANGE: 2890 m. Prototrochus theeli (Östergren, 1905) Myriotrochus theeli — Östergren, 1905: CLIX–CLXI; 1938: taf. 3 figs. 1–4; Belyaev, Mironov, 1982: 86–87; Madsen, Hansen, 1994: 124, fig. 87, map 36. Prototrochus theeli — Smirnov, Smirnov, 2006: 105–106. LOCALITIES: Swedish Zoological (Kolthoff’s) Polar Expedition (1900), St. 29 (Östergren, 1905; 1938); “Polarstern” ARK XI/1 cruise (1995), St. 23a, 44-GKG (bio1), 44-GKG (bio2), 49-GKG (bio2) (det. A.V. Smirnov, unpublished). DISTRIBUTION: Arctic: Norwegian and Greenland Seas, Laptev Sea, Amundsen Basin. REMARKS: Hansen and Madsen (1994) indicate that this species “has been… found in several localities in the Norwegian Basin, between Iceland and Norwey, during the “Håkon Mosby” expeditions 1981–1984. (T. Brattegard, personal coomunication)” Unfortunatelly they published the findings of P. theeli only as dots on the map, without station data. DEPTH RANGE: 600–3900 m. Genus Siniotrochus Pawson, 1971 TYPE SPECIES: Siniotrochus phoxus Pawson, 1971. COMPOSITION: 3 species (Pawson, 1971; Belyaev, Mironov, 1981b; Belyaev, Mironov, 1982; Gage, Billett, 1986). DISTRIBUTION: North Atlantic and North Pacific; Atlantic — 2, Pacific — 1 species. Siniotrochus myriodontus Gage et Billett, 1986 Siniotrochus myriodontus Gage, Billett, 1986: 266, figs. 28C–E, 29–31. LOCALITIES: “Discovery” (1978), St. 9756#14; “Challenger” (1979–1982), 50603#1, 50604#1, 51415#1 (Gage, Billett, 1986). DISTRIBUTION: North-East Atlantic: Porcupine Seabight. DEPTH RANGE: 3490–4000 m. Genus Parvotrochus Gage et Billett, 1986 TYPE SPECIES: Parvotrochus belyaevi Gage et Billett, 1986. COMPOSITION: 1 species (Gage, Billett, 1986). DISTRIBUTION: North Atlantic. Parvotrochus belyaevi Gage et Billett, 1986 Parvotrochus belyaevi Gage, Billett, 1986: 263, figs. 24C–F, 26, 27. LOCALITIES: “Challenger” (1976–1985), St. ES 57, ES 135, ES 143, ES 147, ES 152, ES 169, ES 172, ES 180, ES 204, ES 207, ES 285 (Gage, Billett, 1986; Harvey et al., 1988). DISTRIBUTION: North-East Atlantic: Rockall Trough. DEPTH RANGE: 1160–2921 m. Family Synaptidae Genus Protankyra Östergren, 1898, emend. Rowe et Pawson, 1967 TYPE SPECIES: Synapta abyssicola Théel, 1886 [=Protankyra brychia (Verrill, 1885)]. COMPOSITION: 35 species (Östergren, 1898; Clark, 1908; Heding, 1928). DISTRIBUTION: Atlantic — 5, Indian Ocean — 10, Indian Ocean and Pacific — 4, Pacific — 15, Atlantic and Pacific — 1 species. Protankyra brychia (Verrill, 1885) Synapta brychia Verrill, 1885: 539. Protankyra brychia — Östergren, 1898: 116; Clark, 1908: 25, 105, pl. 4 figs. 12–14; Deichmann, 1930: 209; Deichmann, 1940: 229, pl. 41 figs. 1–3; Deichmann, 1954: 408; Madsen, 1953: 151, fig. l; Sibuet, 1977: 554; Gage et al., 1985: 201; Harvey et al., 1988: 190. Synapta abyssicola Théel, 1886a: 14, pl. 1 fig. 11. Protankyra abyssicola — Perrier, 1902: 538; Clark, 1908: 25, 105, Pl. 4 figs. 8–11; Clark, 1913: 227; Clark, 1920: 124; Clark, 1924: 496, pls. 11 figs 6–7, 12 fig. l; Hérouard, 1923: 140; Ludwig, Heding, 1935: 146, figs. 12–13. Protankyra abyssicola var. pacifica Ludwig, 1894: 174, pl. 18 figs. 13–19. Protankyra pacifica — Clark, 1908: 25, 105; Clark, 1920: 124; Heding, 1928: 252; Ludwig, Heding, 1935: 149; Carney, Carey, 1982: 69; Maluf, 1988: 163. Synapta sp. — Théel, 1886b: 20. Deep-sea fauna of European seas. Holothuroidea LOCALITIES: “Princesse Alice II” (1910), St. 2986 (Hérouard, 1923); “Challenger” (1975– 1982), St. ES 32, ES 56, ES 118, ES 152, ES 164, ES 169, ES 172, ES 180, ES 185, SBC 188, ES 207 (Gage et al., 1985; Harvey et al., 1988; Pawson et al., 2003). DISTRIBUTION: Atlantic, Pacific. In the NorthEast Atlantic: Rockall Trough, Bay of Biscay. DEPTH RANGE: 869–4990 m. In the NorthEast Atlantic: 2871–4879 m. Genus Labidoplax Östergren, 1898, sensu Heding, 1931 TYPE SPECIES: Synapta tenera Norman, 1864 (nomen nudum) = Synapta buski McIntosh, 1866. COMPOSITION: 5 species (Östergren, 1898; 1905; Heding, 1931a; Gage, 1985; Smirnov, 1997). DISTRIBUTION: North Atlantic — 4, Pacific — 1 species. Labidoplax southwardorum Gage, 1985 Labidoplax southwardorum Gage, 1985: 255, figs. 1, 3a, d (left and bottom), e, f; Gage et al., 1985: 200; Harvey et al., 1988: 186, fig. 3a. LOCALITIES: “Challenger” (1973–1985), St. ES 10, ES 27, ES 28, ES 34, SBC 48, ES 53, ES 56, ES 57, SBC 58, ES 59, SBC 61, ES 111, ES 118, ES 129, ES 135 (Gage, 1985; Gage et al., 1985), ES 137, ES 140, ES 143, ES 147, ES 152, ES 164, ES 169, ES 172, SBC 174, ES 176, ES 180, ES 184, ES 185, ES 190, ES 197, ES 200, ES 204, ES 218, ES 244, ES 283, ES 285, ES 289 (Gage, 1985; Gage et al., 1985; Harvey et al., 1988). DISTRIBUTION: North-East Atlantic: Rockall Trough. DEPTH RANGE: 1000–2946 m. Labidoplax similimedia Gage, 1985 Labidoplax similimedia Gage, 1985: 259, figs. 2, 3b, d right and top; Gage et al., 1985: 201; Harvey et al., 1988: 187. LOCALITIES: “Challenger” (1975–1985), St. ES 34, ES 118, ES 129, ES 137, ES 143, ES 147, ES 164, ES 169, ES 172, ES 176, ES 180, ES 185, ES 190, ES 200, ES 204, ES 218, ES 244, ES 283, ES 285, ES 289 (Gage, 1985; Gage et al., 1985; Harvey et al., 1988). DISTRIBUTION: North-East Atlantic: Rockall Trough. DEPTH RANGE: 1101–2946 m. Order Elasipodida Family Laetmogonidae Genus Benthogone Koehler, 1896 TYPE SPECIES: Benthogone rosea Koehler, 1896. COMPOSITION: 3 species. 159 DISTRIBUTION: Indian Ocean — 1, Indonesia — 1, cosmopolitan — 1 species. Benthogone rosea Koehler, 1896 Benthogone rosea Koehler, 1896: 114–117, figs. 2, 3, 36, 46; Perrier, 1902: 399–405, pls. XIV: 1–2, XIX: 8–14; Grieg, 1921: 5–6; Hérouard, 1923: 38–39; Heding, 1940: 369; Madsen, 1947: 15–16; Pawson, 1965: 219–221, pl. 5. Benthogone rosea var. cylindrica Perrier, 1896: 900. Benthogone rosea var. 4-lineata Perrier, 1896: 900. Benthogone quadrilineata — Heding, 1940: 369; Heding, 1942a: 15. Non Benthogone quatrolineata — Augustin, 1908. LOCALITIES: “Princesse Alice II”, St. 2290; “Michael Sars”, St. 25A; “Talisman”, St. 35, 58, 59 (Perrier, 1902); Porcupine Abyssal Plain (Billett, 1991); BIOGAS, St. 1, 2, 3 (Sibuet, 1977); BIOGAS VII CP 26, BIOGAS IX CP 33 (Massin, 1984); numerous “Discovery” stations (Billett, 1988). DISTRIBUTION: Eastern Atlantic from Ireland to the Cape Verde Islands; western Indian Ocean; north of New Zealand. DEPTH RANGE: 1103–2480 m. Genus Laetmogone Théel, 1879 COMPOSITION: 12 species. TYPE SPECIES: Laetmogone wyvillethomsoni Théel, 1879. DISTRIBUTION: Atlantic — 2, Pacific — 7, Antarctic and Pacific — 1, Pacific and Indian — 1, cosmopolitan — 1 species. Laetmogone billetti Rogacheva et Gebruk in Rogacheva et al., 2013 Laetmogone billetti Rogacheva et al., 2013: 595–598, figs. 5, 6, 17h, i, 18k, 19g–i. LOCALITIES: “James Cook” ECOMAR, Sts. JC048/24 Dive 165, JC048/16 Dive 162, JC048/56 Dive 180 (Rogacheva et al., 2013). DISTRIBUTION: Known from its type locality on the Northern Mid-Atlantic Ridge, Charlie-Gibbs Fracture Zone area. DEPTH RANGE: 2272–2758 m. Family Psychropotidae Genus Benthodytes Théel, 1882 TYPE SPECIES: Benthodytes typica Théel, 1882. COMPOSITION: 10 species. DISTRIBUTION: Atlantic and Indonesia — 1, Atlantic — 2, Indian — 3, Pacific — 1, Indonesia — 1, cosmopolitan — 2 species. Benthodytes valdiviae Hansen, 1975 Benthodytes valdiviae Hansen, 1975: 82–84, figs. 30– 31; Thandar, 1999: 384–386, fig. 7; Gebruk, 2008: 50, 51. LOCALITIES: “Valdivia”, St. 33 (Hansen, 1975); “G.O. Sars”, MAR-ECO cruise, St. 46/372. (Gebruk, 2008). DISTRIBUTION: North and South Atlantic (off South Africa). 160 A.V. Gebruk et al. DEPTH RANGE: 2480–3050 m. Benthodytes lingua Perrier, 1896 Benthodytes lingua Perrier, 1896: 902; Perrier, 1902: 456–461, pls. XII: 1–2, XXI: 1–9; Deichmann, 1930: 124– 125; Deichmann, 1940: 200–201, pl. XXXV: 3–4; Heding, 1942a: 15; Deichmann, 1954: 384; Rogacheva et al., 2013: 599, fig. 18b. Benthodytes janthina von Marenzeller, 1882 — Grieg, 1921: 11; Heding, 1942a: 15. Pannychia glutinosa Hérouard, 1902: 32, pl. IV: 17. LOCALITIES: “M. Sars”, St. 10; “Ingolf”, St. 18, 20; “Talisman” and “Travailleur” St. 38, 39, 44 (Perrier, 1902); “Valdivia”, St. 33 (Heding, 1942a); BIOGAS St. 2 (Sibuet, 1977); “Discovery” St. 11121#10 (Billett et al., 1985); “G.O. Sars”, MARECO cruise, St. 40/367, 46/372, 50/373 (Gebruk, 2008); “James Cook” ECOMAR, Sts. JC037/27, JC048/36 Dive 171, JC048/48 Dive 176 (Rogacheva et al., 2013). DISTRIBUTION: North and South Atlantic. DEPTH RANGE: 860–3192 m. Benthodytes typica Théel, 1882 Benthodytes typica Théel. 1882a: 103–104, pls. XXVII: 7, XXXV: 4, XXXVIII: 5, XLIV: 8; Théel, 1886: 2; von Marenzeller, 1893b: 12; Grieg, 1921: 10, fig. 8, pl. III: 6–7; Hérouard, 1923: 101–102, pl. VI: 4; Deichmann, 1930: 123–124; Deichmann, 1940: 200, pl. XXXV: 1–2; Heding, 1940: 368; Madsen, 1953: 160–161, fig. 8; Deichmann, 1954: 384. Benthodytes papillifera Théel, 1882a: 102–103, pl. XXXIV: 14. Benthodytes glutinosa Perrier, 1896: 902–903; Perrier, 1902: 462–465, pls. XIII: 5, XX: 31; Koehler, Vaney, 1905: 72–74, pl. XII: 10; Clark, 1920: 141; Grieg, 1921: 10–11, pl. III: 1–2. Benthodytes janthina von Marenzeller, 1882 — Hérouard, 1902: 30; Hérouard, 1923: 103. LOCALITIES: “Swedish Deep–Sea Expedition” St. 313, 357, 387 (Madsen, 1953); “Ingolf”, St. 20; “Princesse Alice II”, St. 2111; “Michael Sars”, St. 35, 53; “Hirondelle”, St. 248; “Talisman” and “Travailleur”, 32°19′– 34°46′N, 36°11′– 38°04′W, 3175– 3243 m (Perrier, 1902); Porcupine Abyssal Plain (Billett, 1991); numerous “Discovery” stations in the North-East Alantic (Billett, 1988); “G.O. Sars”, MAR-ECO cruise, St. 40-367, 42-368, 46-372, 50373, 52-374, 54-377. DISTRIBUTION: cosmopolitan. DEPTH RANGE: 1873–4700 m. Benthodytes sanguinolenta Théel, 1882 Benthodytes sanguinolenta Théel, 1882a: 104–105, Pls. XXIII, XL: 4–5, XLII: 6; Ludwig, 1894: 53–60, Pl. I: 1–8; Koehler, Vaney, 1905: 72; Clark, 1913: 233; Ohshima, 1915: 245; Clark, 1920: 142; Clark, 1923a: 162; Clark, 1923b: 420; Heding, 1940: 367; Hansen, 1956: 44– 45; Hansen, 1975: 94–96; Pls. III–VI, IX: 6–7, XII: 4–5; Carney, Carey, 1976: 69; Pawson, 1982: 129–145; Bluhm, Gebruk, 1999: 175, Fig. 3D; Gebruk, 2008: 50, 51; Rogacheva et al., 2009: 463–464, fig. 2. LOCALITIES: “G.O. Sars”, MAR-ECO cruise, St. 40/367, 54/377, 64/381, 66/383, 68/384, 74/387 (Gebruk, 2008); “James Cook” ECOMAR, St. JC037/ 19 (Rogacheva et al., 2013). DISTRIBUTION: cosmopolitan. DEPTH RANGE: 768–7250 m. Benthodytes gosarsi Gebruk, 2008 Benthodytes gosarsi Gebruk, 2008: 49–52, figs. 1A, 2, 3. LOCALITIES: “G.O. Sars”, MAR-ECO cruise, St. 40/367, 54/377 (Gebruk, 2008); BIOICE St. 2862, 3572; “James Cook” ECOMAR, Sts. JC011/ 23, JC011/75, JC011/101, JC037/15, JC037/19, JC037/27, JC037/61, JC037/67, JC037/70; “James Cook”, St. JC036/04, JC36/21 (Rogacheva et al., 2013). DISTRIBUTION: Atlantic: Mid-Atlantic Ridge from Azores to Iceland and Northeast Atlantic, Whittard Canyon (Rogacheva et al., 2013). DEPTH RANGE: 2967–3670 m. Genus Psychropotes Théel 1882 TYPE SPECIES: Psychropotes longicauda Théel, 1882. COMPOSITION: 10 species. DISTRIBUTION: Atlantic — 3, Indian — 2, Antarctic — 1, Pacific — 2, cosmopolitan — 2 species. Psychropotes semperiana Théel, 1882 Psychropotes semperiana Théel, 1882a: 100–101, pl. XXXIV: 10–11; Hansen, 1975: 102–105, figs. 41–42. Psychropotes kerhervei Hérouard, 1902: 27–30, pl. IV: 1– 9. Euphronides kerhervei — Hérouard, 1923: 104, pl. III: 4–5; Deichman, 1940: 202–203, pl.XXXV: 9–12; Madsen, 1953: 161–163, fig. 9. LOCALITIES: “Princesse Alice”, St. 749, “Princesse Alice II”, St. 1306, 2111; Porcupine Abyssal Plain (Billett, 1991); BIOGAS (1972–1974) “Jean Charcot“, St.2. DISTRIBUTION: Atlantic and the western part of Indian Ocean. DEPTH RANGE: 1433–5600 m. Psychropotes depressa (Théel, 1882) Euphronides depressa — Théel, 1882a: 93–96, pls. XXVI, XXX: 5–6, XL: 7, XLVI: 4; Ohshima, 1915: 244– 245, fig. 1; Ohshima, 1916–1919, with three figures. Psychropotes depressa — Hansen, 1975: 106–111, figs. 43–44; Gebruk, 2008: 50, 51; Rogacheva et al., 2013: 599, fig. 17f, g. Euphronides depressa var. minor Théel, 1886b: 2. Euphronides cornuta Verrill, 1884: 217; Verrill, 1885: 518, 538, figs. 32–33; Deichmann, 1930: 127–128; Heding, 1940: 368. Euphronides tanneri — Ludwig, 1894: 39–44, pls. III: 7, IV, V: 17–19. Euphronides auriculata Perrier, 1896: 901–902; Perrier, 1902: 434–438, pls. XIII: 1–2, XX: 12–13; Grieg, 1921: 8–9. Euphronides violacea Perrier, 1896: 902; Perrier, 1902: 438–441, pl. XX: 14; Deichmann, 1930: 128–129; Deichmann, 1940: 201–202; Heding, 1942a: 15–16; Madsen, 1947: 16; Deichmann, 1954: 384. Euphronides talismani Perrier, 1896: 902; Deep-sea fauna of European seas. Holothuroidea Perrier, 1902: 441–444, pl. XX: 15; Hérouard, 1902: 30– 31, pl. II: 19–22; Deichmann, 1930: 129; Heding, 1942a: 15, fig. 15. Benthodytes assimilis Théel, 1886b: 2–3. LOCALITIES: “Ingolf”, St. 11, 18, “M. Sars”, St. 25A; “Princesse Alice”, St. 673; “Skagerak” (Madsen, 1947); “Talisman” and “Travailleur”, St. 129, St.38, (Perrier, 1902); Porcupine Abyssal Plain (Billett, 1991); BIOGAS (1972–1974) “Jean Charcot“, St. 2, 3; numerous “Discovery” stations (Billett, 1988); “G.O. Sars”, MAR-ECO cruise, Sts. 40/ 367, 42/368, 46/372, 66/383, 72/386 (Gebruk, 2008). BIOICE, St., 2854, 2855, 3070, 3073, 3074, 3075, 3077, 3172, 3571, 3572, 3574 (unpublished); “James Cook” ECOMAR, Sts. JC011/23, JC011/101, St. JC037/15, St. JC037/19, JC037/27, JC037/61, JC037/67, JC037/70 (Rogacheva et al., 2013). DISTRIBUTION: cosmopolitan. DEPTH RANGE: 957–4060 m. Psychropotes longicauda Théel, 1882 Psychropotes longicauda Théel, 1882a: 96–98, pls. XXVII: 1, XXVIII, XXXV: 13–17, XXXVII: 10; Agatep, 1967b: 67, pl. XI: 1–7; Hansen, 1975: 115–126, figs. 49– 54; Billett et al., 1985: 405, figs. 1, 4; Walker et al., 1987: 277–282; Gebruk, 1993: 240, fig. 6: 4–5; Bluhm, Gebruk, 1999: 173–174; Wigham et al., 2003: 409–441; Gebruk, 2008: 50, 51; Rogacheva et al., 2009: 473–474, fig. 7. Psychropotes longicauda var. monstrosa Théel, 1882a: 98–99, pls. XXIX: 2, XXX, XXXIX: 1. Psychropotes longicauda var. Fusco-purpurea Théel, 1882a: 99, pls. XXIX: 1, XXXV: 11. Psychropotes longicauda var. antarctica Vaney, 1908: 419–420. Psychropotes buglossa E. Perrier, 1886: 283, fig. 200; R. Perrier, 1902: 445–453, fig. 7, pls. XIII: 3–4, XX: 16–28; Hérouard, 1923: 105– 108, pls. I: 32, VI: 2. Psychropotes raripes Ludwig, 1894: 48–51, pl. IV: 1–16; Ohshima, 1915: 244; Ohshima, 1916–1919: with one figure; Clark, 1920: 140–141, pl. I: 1. Psychropotes dubiosa Ludwig, 1894: 52–53, pl. II: 5– 7. Psychropotes grimaldii Hérouard, 1896: 167, fig. 2; Hérouard, 1902: 25–27, pl. III: 1–2. Psychropotes fucata Perrier, 1896: 902; Perrier, 1902: 453–455, pl. XX: 29–30. Psychropotes laticauda Vaney, 1908: 420–422, pl. II: 14, 24. Psychropotes brucei Vaney, 1908: 422–423, pls. I: 13, II: 21–22, III: 41–42. Euphronides dyscrita Clark, 1920: 139, pl. II: 3. Nectothuria translucida Belyaev, Vinogradov, 1969: 711–716, figs. 1–4. ?Psychropotes longicaudata — Carney, Carey, 1982: 597–607. LOCALITIES: “Princesse Alice”, St. 527, “Princesse Alice II”, 2948, 2964, 2986; “Talisman” and “Travailleur”, St. 135; localities without station numbers: 30°09′– 44°29′N, 15°52′–23°37′W, 2110–5005 m (Perrier, 1902); BIOGAS (1972–1974) “Jean Charcot”, Sts. 2, 3, 4, 5; Porcupine Abyssal Plain (Billett, 1991); “G.O. Sars”, MAR-ECO cruise, Sts. 40/367, 54/377 (Gebruk, 2008). DISTRIBUTION: cosmopolitan. DEPTH RANGE: 2210–5173 m. Family Elpidiidae Genus Peniagone Théel, 1882 TYPE SPECIES: Peniagone wyvillii Théel, 1882. 161 COMPOSITION: about 20 species. DISTRIBUTION: widely distributed in the world ocean, except for the Arctic. Highest species diversity is found in the Pacific and the Antarctic. Peniagone azorica von Marenzeller, 1892 Peniagone azorica von Marenzeller, 1892: 64; von Marenzeller, 1893b: 12–13, pls. I: 4, II: 5; Hérouard, 1902: 42–43, pl. IV: 21–26; Hérouard, 1923: 87–88; Grieg, 1921: 8, fig. 4; Heding, 1942a: 20; Hansen, 1975: 138– 142 (partim: fig. 63, 5–9); Gebruk, 1990: 110–111, fig. 45 (partim); Rogacheva et al., 2013: 603–605, figs. 9, 12h, i, 17j, 19d. LOCALITIES: “Hirondelle”, St. 248; “PrincesseAlice”, St. 527, “Princesse Alice II”, 2990, 3006; “Ingolf”, Sts. 11, 41; “Michael Sars”, St. 88; BIOGAS (1972–1974) “Jean Charcot“, Sts. 1, 2, 4; other numerous localities in the North-East Atlantic (Gage et al., 1985; Tyler et al., 1985); “G.O. Sars” MARECO cruise, St. 50/373; BIOICE, St. 734, Sample Nr 2862, 2863, 3169, 3170, 3172; “James Cook” ECOMAR, Sts. JC011/101, JC011/106, JC011/111, JC037/ 79, JC048/24 Dive 165 (Rogacheva et al., 2013). DISTRIBUTION: Reliable records in the central and eastern North Atlantic (Rogacheva et al., 2013). DEPTH RANGE: 1385–4020 m. Peniagone porcella Perrier, 1896 Peniagone porcella Perrier, 1896: 901; Perrier, 1902: 426–429, pls. XIII: 7–9, XIX: 13–23; Madsen, 1953: 155– 156, fig. 4; Hansen, 1975: 134; Gebruk, 1990: 97–98, fig. 37: 1–9. LOCALITIES: “ Talisman” St. 134, 42°19′N, 23°36′W, 4060 m (Perrier, 1902). DISTRIBUTION: Atlantic, North Indian Ocean and Antarctic. DEPTH RANGE: 3400–5044 m. Peniagone islandica Deichmann, 1930 Peniagone islandica Deichmann, 1930: 137; Heding, 1942a: 20–21, fig. 19; Hansen, 1975: 150; Gebruk, 1990: 95–96, fig. 35. REMARKS: may be a synonym of P. azorica (Rogacheva et al., 2013). LOCALITIES: “Ingolf”, St. 18; “James Cook” ECOMAR, Sts. JC011/23, JC037/15, JC037/19, JC037/27, JC037/79, JC048/43 Dive 174, JC048/54 Dive 179 (Rogacheva et al., 2013). DISTRIBUTION: Northern Mid-Atlantic Ridge, from Charlie-Gibbs Fracture Zone Area to Iceland. DEPTH RANGE: 2137–2758 m. Peniagone diaphana (Théel, 1882) Scotoanassa diaphana Théel, 1882a: 55–56, pls. IX: 3–5, XXXV: 18, XLIV: 9. Peniagone diaphana — Hansen, 1975: 153–155, fig. 71; Gebruk, 1990: 91–93, fig. 33. Scotoanassa translucida Hérouard, 1899: 71–72, fig. 3; Hérouard, 1902: 43–45, pls. III: 4–6, VI: 17–20; Hérouard, 1923: 88–90, pls. III: 7–8, IV: 4; Madsen, 1953: 158–159, fig. 6. 162 A.V. Gebruk et al. LOCALITIES: “Princesse-Alice”, St. 749, 753, “Princesse Alice II”, 1558, 2983, 2997; “Vityaz”, St. 7943; “Meteor”, 1988/6 MOC1-11: B5, B8, B9; BIOGAS (1972–1974) “Jean Charcot“, Sts. 1, 2, 3, 4, 5, 6; also numerous stations of “Discovery” and “Challenger” in the North-East Atlantic (Billett et al., 1985; Gage et al., 1985; Billett, 1991); “G.O. Sars”, MAR-ECO cruise, St. 68/384. DISTRIBUTION: throughout the North Atlantic, common in the North Pacific, Antarctic. DEPTH RANGE: 1529–5600 m. Peniagone longipapillata Gebruk, 2008 Peniagone longipapillata Gebruk, 2008: 56–59, figs. 1B, 9, 10; Rogacheva et al., 2013: 606–608, figs. 12k–m, 17o, 18f, g, p. LOCALITIES: “G.O. Sars”, MAR-ECO cruise, St. 40/367, 46/372, 50/373, 52/374, 68/384, 72/386; “James Cook” ECOMAR, Sts. JC011/111, JC048/ 16 Dive 162, JC048/24 Dive 165. DISTRIBUTION: North Atlantic, Mid-Atlantic Ridge from the Azores to the Charlie-Gibbs Fracture Zone, and Northeast Atlantic (Porcupine Seabight and Whittard Canyon) [Rogacheva et al., 2013]. DEPTH RANGE: 2398–3036 m. Peniagone marecoi Gebruk, 2008 Peniagone marecoi Gebruk, 2008: 54–56, figs. 7, 8. LOCALITIES: “G.O. Sars”, MAR-ECO cruise, Sts. 42/368, 46/372, 50/373, 52/374, 54/377, 72/ 386. DISTRIBUTION: Atlantic, Mid-Atlantic Ridge from the Azores to the Charlie-Gibbs Fracture Zone. DEPTH RANGE: 1771–3509 m. Peniagone purpurea (Théel, 1882) Elpidia purpurea Théel, 1882a: 21–23, pls. VII: 4–6, XXXIII: 13–14, XLIV: 6. Peniagone purpurea — Hansen, 1975: 151–152; Gebruk, 1990: 111–113, fig. 46. Elpidia ambigua Théel, 1882a: 27–28, pl. XXXIII: 6. Peniagone lacinora Agatep, 1967: 53–55, pl. III: 1–9. Peniagone vexillum Perrier, 1902: 429, pls. XII: 6, XIX: 24–25.?Peniagone ferruginea Grieg, 1921: 7–8, fig. 3, pl. I: 4–6. LOCALITIES: “Talisman”, St. 134, 42°19′N, 25°36′W, 4060 m (Perrier, 1896). DISTRIBUTION: Antarctic, Atlantic, West Pacific. DEPTH RANGE: 2800–5610 m. DEPTH RANGE: 2600–2750 m. Genus Achlyonice Théel, 1879 TYPE SPECIES: Achlyonice ecalcarea Théel, 1879. COMPOSITION: 5 species (Gebruk, 1990, 1997). DISTRIBUTION: West Pacific — 3, central Atlantic — 1, cosmopolitan — 1 species. Achlyonice myriamae Gebruk, 1997 Achlyonice myriamae Gebruk, 1997: 214–215, fig. 2. LOCALITIES: BIOGAS III CV 26). DISTRIBUTION: known only from type locality. DEPTH: 2822 m. Genus Amperima Pawson, 1965 TYPE SPECIES: Periamma roseum Perrier, 1896. COMPOSITION: 8 species. DISTRIBUTION: Pacific — 2, Antarctic — 2, Atlantic — 1, cosmopolitan — 3 species. Amperima rosea (Perrier, 1896) Periamma roseum Perrier, 1896: 901; Perrier, 1902: 419–423, pls. XIII: 10–12, XX: 1–11; Hérouard, 1923: 91–94. Amperima rosea — Hansen, 1975: 158–159, fig, 74; Gebruk, 1990: 140–141, fig.61: 3–6. LOCALITIES: “Talisman” (Perrier, 1902): 42°19′–44°29′N, 15°52′– 23°36′W, 4060–5005 m; 44°20′N, 19°31′W, 4255 m; 44°29′N, 15°52′W, 5005 m; “Princesse-Alice II”, St. 2994; Porcupine Abyssal Plain (Billett, 1991; Wigham et al., 2003); BIOGAS (1972–1974) “Jean Charcot“, Sts. 1, 2, 3, 5, 6; “G.O. Sars”, MAR-ECO cruise, St. 50/373. DISTRIBUTION: North Atlantic, North-West Pacific. DEPTH RANGE: 4060–5740 m. Amperima furcata (Herouard, 1899) Kolga furcata Hérouard, 1899: 171, fig. 2; Hérouard, 1902: 40–41, pls. III: 7, VI: 4–10, VIII: 17. Amperima furcata — Hansen, 1975: 159, fig. 75; Gebruk, 1990: 141– 142, fig. 61: 7–9; Rogacheva et al., 2013: 600–601, figs. 7a–e, 17k, 18m, n, 19c. Pariamma furcata — Hérouard, 1923: 91. Peniagone coccinea Rogacheva et al., 2013: 608– 610, figs. 13, 14, 18h, i, o, 19e, in situ video record (online suppl.). LOCALITIES: “Princesse-Alice II”, St. 2990; “G.O. Sars”, MAR-ECO cruise, St. 40/367 (Gebruk, 2008); “James Cook” ECOMAR, Sts. JC011/23, jc037/15, JC048/43 Dive 174, JC048/54 Dive 179 (Rogacheva et al., 2013). DISTRIBUTION: North Atlantic, North Pacific. DEPTH RANGE: 1846–3015 m. LOCALITIES: “James Cook” ECOMAR, Sts. JC037/15, JC037/19, JC037/27, JC048/38 Dive 172, JC048/43, Dive 174, JC048/54, Dive 179. DISTRIBUTION: Mid-Atlantic Ridge, CharlieGibbs Fracture Zone Area. TYPE SPECIES: Kolga ludwigi von Marenzeller, 1893. COMPOSITION: 4 species. Peniagone coccinea Rogacheva et Gebruk in Rogacheva et al., 2013 Genus Penilpidia Gebruk, 1988 Deep-sea fauna of European seas. Holothuroidea DISTRIBUTION: Atlantic — 2 species, Mediterranean — 1, Pacific — 1. Penilpidia midatlantica Gebruk, 2008 Penilpidia midatlantica Gebruk, 2008: 52–56, figs. 4–6; Rogacheva et al., 2013: 610–611. Locality: “G.O. Sars”, MAR-ECO cruise, St. 46/ 368; “James Cook” ECOMAR, St. JC037/19. DISTRIBUTION: Atlantic, Mid-Atlantic Ridge. DEPTH RANGE: 2063–2750 m. Genus Kolga Danielssen et Koren, 1879 TYPE SPECIES: Kolga hyalina Danielssen et Koren, 1879. COMPOSITION: 1 (probably 3) species. DISTRIBUTION: Cosmopolitan, 652–6235 m. Kolga hyalina Danielssen et Koren, 1879 Kolga hyalina Danielssen, Koren, 1879: 83–106, pl. I, II; Danielssen, Koren, 1882: 3–20, pls. I–III, Théel, 1882a: 39; Ludwig, 1898: 12; Ludwig, 1901: 140; Michailovskij, 1903:4; Mortensen, 1927: 27; Deichmann, 1930: 132–133; Mortensen, 1932: 43, fig. 5; Djakonov, 1933: 135, fig. 69c; Heding, 1942a: 19, textfig. 18; Gorbunov, 1946: 47; Gurjanova, 1957: 361; Koltun, 1964: 46, 47, 53, 62, 68–69, 76, 77, 78; Baranova, 1964: 368; Agatep, 1967a: 140 (partim); Hansen, 1975: 170, figs. 86, 95, 2–3, pl. 9: 8, pl. 12: 11; Gebruk, 1990: 121, fig. 51; Hansen, Madsen, 1994: 88, fig. 56, 57, map 22; Rogacheva, 2007: 384–391, figs. 11–13; Rogacheva, 2012: 1185– 1186, figs. 2–3; non Kolga hyalina — Billett, Hansen, 1982: 799, fig. 1–8; Gage et al., 1985: 200; Harvey et al., 1988: 185; non Kolga nana — Théel, 1882a: 39–42, pls. 2, 34, 33 (1–2), 34 (5), 36 (26), 42 (5, 8); non Elpidia nana Théel, 1879: 15–16, figs. 20–22. LOCALITIES: “Vøringen”, Norwegian North– Atlantic Expedition (1876–1878), St. 295, 303, 353 (Danielssen, Koren, 1879, 1882); “Ingolf” (1895–1896), St. 20, 36, 37, 112 (Heding, 1942a); Second Test Cruise on the Icebreaker “Ermak” (1899), St. 31 (Michailovskij, 1903); Swedish Zoological Polar Expedition (1898), St. 26, 27; Swedish Zoological Polar Expedition (1900), St. 13 (unpublished data); I High Latitude Arctic Expedition on the Ice Steamer “Sadko” (1935), St. 3/16 (Rogacheva, 2007), 6, 10 (det. G.P. Gorbunov, unpublished), 59 (Gorbunov, 1946); III High Latitude Arctic Expedition on the Ice Steamer “Sadko” (1937–1938), St. 97, 100 (Gorbunov, 1946); Drifting Station “Severnyi Polus–3” [“North Pole”–3] (1954–1955), St. 18 (Gurjanova, 1957; Koltun, 1964); Drifting Station “Severnyi Polus–4” [“North Pole”–4] 1st shift (1954– 1955), St. 4, 7, 8 (Gurjanova, 1957; Koltun, 1964); Drifting Station “Severnyi Polus–4” [“North Pole”– 4] 2nd shift (1955), St. 2, 4 (Rogacheva, 2007), 3 (det. T. S. Saveljeva, unpublished); Drifting Station “Severnyi Polus–5” [“North Pole”–5] (1955), St. 1, 2, 4 (Rogacheva, 2007, original identification by T.S. Saveljeva); High Latitude Oceanographic Ex- 163 pedition on the Ice Steamer “Lena” (1955), St. 35, 37 (Koltun, 1964; Baranova, 1964); High Latitude Oceanographic Expedition on the Ice Steamer “Ob” (1956), St. 6, 22, 45 (Koltun, 1964); “Sevastopol’”, St. 1709 (Rogacheva, 2007); Arctic Research Laboratory Ice Station 2, St. 380 (Agatep, 1967a); Drifting Station “Severnyi Polus–22” [“North Pole”–22] (1976–1980), St. 58; Drifting Station “Severnyi Polus–23” [“North Pole”–23] (1977), St. 4 (Rogacheva, 2007, original identification by G.M. Belyaev); “Håkon Mosby” St. 82.11.21.2, 82.11.21.4, 83.06.10.1, 83.06.11., St. 84.03.17.1, 84.03.17.2, 86.06.12.1, 86.07.24.1, 86.07.26.1, 86.07.28.1, 87.06.14.1 (Rogacheva, 2007); “Polarstern” ARK IX/4 cruise (1993): St. 32, 50, 54 (Rogacheva, 2007, original identification by A.V. Smirnov); “Polarstern” ARK XI/1 cruise (1995), St. 44-GKG(bio1), 49-GKG(bio1) (det. A.V. Smirnov, unpublished); “Polarstern” ARK XV/ 1 cruise (1999), Dive 4 [as Irpa abyssicola in Gebruk et al. (2003)]; “Healy”, St. 11, 13 (Rogacheva, 2007). REMARKS. Kolga hyalina was regarded as cosmopolitan for many years. Evidence from recent revision of Arctic elpidiids suggests that this species is confined to the Arctic deep-sea biogeographical subregion (Rogacheva, 2007). DISTRIBUTION: confirmed as common in the Arctic Basin and the Norwegian Sea north of the Faroe-Iceland Ridge. DEPTH RANGE: 659–4106 m. Kolga nana (Théel, 1979) Elpidia nana Théel, 1879: 15–16, figs. 20–22. Kolga nana — Théel, 1882a: 39–42, pls. 2, 34, 33 (1–2), 34 (5), 36 (26), 42 (5, 8); Rogacheva, 2012: 1186–1190, figs. 4– 7. Kolga hyalina — Hansen, 1975: 170–171 (partim); Billett, Hansen, 1982: 804–806, Figs. 2 (1–9), 4–6; Gage et al., 1985: 200; Harvey et al., 1988: 185; Gebruk, 1990: 121–122 (partim); Kolga sp. — Gebruk, 2008: 50, 51, 52, 58, fig. 1c. LOCALITIES: “Discovery”, St. 7711#62, 7711#85, 9756#9, 9756#14 10113#1, 10114#1, 10115#1, “Challenger”, St. 50603#1, 50604#1, 50605#1, (Billett, Hansen, 1982); “Challenger” (1980–1983), St. ES 172 (Gage et al., 1985), ES 266 (Harvey et al., 1988); “G.O. Sars”, MAR-ECO cruise, St.64-381 (Gebruk, 2008); BIOICE, St. 3176, 3571. DISTRIBUTION: North Atlantic, Antarctic and Subantartic. DEPTH RANGE: 1484–6235 m (Rogacheva, 2012). Genus Ellipinion Hérouard, 1923 TYPE SPECIES: Scotoplanes delagei Hérouard, 1896. COMPOSITION: 9 species. DISTRIBUTION: Atlantic — 2, Pacific — 3, Atlantic and Pacific — 1, Indian — 1, Antarctic — 1, cosmopolitan — 1. 164 A.V. Gebruk et al. Ellipinion delagei (Hérouard, 1896) Elpidia heckeri Baranova, 1989 Scotoplanes delagei Hérouard, 1896: 167–168, fig. 3. Ellipinion delagei — Gebruk, 1990: 133, fig. 58 (list); Rogacheva et al., 2013: 601, figs. 8, 17n, 19f. Elpidia heckeri Baranova, 1989: 218–222, figs. 1–3; Smirnov, Smirnov, 2006: 102–103 fig. 13 (partim); Rogacheva, 2007: 376–378, figs. 3, 4. Elpidia glacialis — Danielssen, Koren, 1882: 80 (partim); Heding, 1942a: 16– 19 (partim); Gorbunov, 1946: 47, 98 (partim); Baranova, 1964: 367–368; Koltun, 1964: 46, 47, 68–69, 76; Bluhm, 1999: 10, 11. Elpidia glacialis glacialis — Agatep, 1967a: 135–139, fig. 1a–d; Hansen, 1975: 176–178, figs. 90, 91 (partim). LOCALITIES: “James Cook” ECOMAR, Sts. JC037/15, JC037/19, JC048/24 Dive 165 (Rogacheva et al., 2013). DISTRIBUTION: North Atlantic. DEPTH RANGE: 1165–2750 m. Ellipinion alani Rogacheva et Gebruk in Rogacheva et al., 2013 Ellipinion alani Rogacheva et al., 2013: 601–603, figs. 8, 17n, 19f. LOCALITIES: “James Cook” ECOMAR, Sts. JC048/24 Dive 165, JC048/43 Dive 174 (Rogacheva et al., 2013). DISTRIBUTION: Known from its type locality on the Northern Mid-Atlantic Ridge, Charlie-Gibbs Fracture Zone area. DEPTH RANGE: 2398–2620 m. Genus Elpidia Théel, 1876 TYPE SPECIES: Elpidia glacialis Théel, 1876. COMPOSITION: 22 species. DISTRIBUTION: cosmopolitan, especially common in trenches, most diversity in the Pacific (Belyaev, 1975). Elpidia gracilis Belyaev, 1975 Elpidia gracilis Belyaev, 1975: 266–267, fig. 6; Gebruk, 1993: 235–236, fig. 3(1–5); Thandar, 1999: 392– 396, figs. 11, 15e. LOCALITIES: “Discovery”, Sts. 9754#3, 9753, 9756#14, “Challenger”, Sts. 50602#2, 50603#1, 50604#1 (Budaeva, Rogacheva, 2013). REMARKS: Perier described the species Tutela echinata based on specimens collected off Morocco (Perrier, 1896). In the description details on body and ossicle morphology and illustrations were lacking. Later Perrier (1902) assigned Tutela echinata to Elpidia glacialis. Belyaev (1971) considered this species as Elpidia sp. 2 since insufficiency of description. According to Belyaev, the species name «echinata» implies ossicles with long vertical apophyses similar to those in E. gracilis. If the two species are conspecific, the name E. echinata has a priority. E. gracilis is the only species in the genus reliably known from the north and equatorial Atlantic. There are two records of Elpidia in the Atlantic that may be related to E. gracilis (E. echinata): south of Canaries (Heding, 1940 as E. glacialis) and in the Romanche Trench (Belyaev, 1971 as Elpidia sp. 3). DISTRIBUTION: South Atlantic (off the South Orkney islands), Weddell Sea, North-East Atlantic (Porcupine Seabight and Porcupine Abyssal Plain). DEPTH RANGE: 1484–6145 m, in the North Atlantic up to 4000 m. LOCALITIES: “Vøringen”, Norwegian NorthAtlantic Expedition (1876–1878), St. 53, 295, 353; “Ingolf”, St. 113; “Ermak”, St. 14/50, 34/60; “Sadko”, St. 10/32, 59, 100; Drifting station “Severnyi Polus –2”, St. 9; “Fedor Litke”, Sts. 35, 37; Drifting station “Severnyi Polus –4/(2)” [“North Pole”–4/ (2)], St. 2, 3, 4; Drifting station “Severnyi Polus –5” [“North Pole”–5], St. 5; “Ob’”, St. 45; Drifting station “ARLIS II”, St. 29; “NORBI”, St. 10; Drifting station “Severnyi Polus–22” [“North Pole”–22], St. 14, 21, 27, 58; “Alaid”, Sts. 30.4, 30.5; “Håkon Mosby”, Sts. 81.08.14.5, 82.11.24.1, 83.06.11.1, 85.01.10.1; “Polarstern”, ARKTIS IX/4, Sts. 32, 54; “Polarstern”, ARKTIS XI/1, St. 23a; “Polarstern”, ARKTIS XV/1, dive 3; “Polarstern”, ARKTIS XVII/ 1, St.81-1; “Polarstern”, ARKTIS XVIII/1a, St. 6242, 62-60; “Healy”, St. 6, 15 (Rogacheva, 2007); Swedish Zoological Polar Expedition (1898), St. 26, 27; Swedish Zoological Polar Expedition (1900), 72Ú50’N 3Ú8’W (station number unknown) and St. 29 (unpublished data); BIOICE, St. 3203, 3204, 3211, 3213, 3214, 3649 (Budaeva, Rogacheva, 2013 and unpublished data). DISTRIBUTION: Arctic Basin and the Norwegian Sea north of the Faroe-Iceland Ridge. DEPTH RANGE: 1700–5550 m. Elpidia belyaevi Rogacheva, 2007 Elpidia belyaevi Rogacheva, 2007: 378–382, figs. 6– 9. Elpidia glacialis — Danielssen, Koren, 1882: 80 (partim). Mortensen, 1932: 41–43, pl. 1 figs. 4–5; Heding, 1942a: 16–19, tables I, II (partim); Gorbunov, 1946: 47, 98 (partim); Koltun, 1964: 77; Piepenburg et al., 1996: 440, 444; Vinogradova et al., 1996: 203, 204, 206. Elpidia glacialis glacialis — Hansen, 1975: 176–178, figs. 90 1– 3, 91 3–8. Elpidia sp.1 — Belyaev, 1971: 356–357. Elpidia heckeri — Smirnov, Smirnov, 2006: 102–103 fig. 13 (partim). LOCALITIES: “Vøringen”, Norwegian NorthAtlantic Expedition (1876–1878), St. 40; “Michael Sars” St. 9; “Sadko”, St. 98; Drifting station “Severnyi Polus–3” [“North Pole”–3], St. 6; “Håkon Mosby”, Sts. 81.06.07.3, 81.08.15.4, 85.01.11.3, 87.06.13.2; “Akademik Mstislav Keldysh”, St. 2079, 2091, 2679, 2702, 2737, 3153, 3177, 3287, 3291, 3318; “Polarstern”, ARKTIS VIII/2, St. 108; BIOICE, Sts. 3204, 3207, 3210, 3230, 3637, 3638, 3648, 3649 (Rogacheva, 2007, and unpublished data). REMARKS: Both Elpidia heckeri and E. belyaevi are often confused with Elpidia glacialis, an Deep-sea fauna of European seas. Holothuroidea upper bathyal and sublittoral species occurring at depths of 70–610 m in the Kara and north–eastern Barents Seas (Rogacheva, 2007) and at 250 m off Franz Josef Fjord in East Greenland (unpublished data). DISTRIBUTION: Arctic basin, Norwegian Sea north of the Faroe-Iceland Ridge, Baffin Bay. DEPTH RANGE: 610–2222 m. Genus Psychroplanes Gebruk, 1988 TYPE SPECIES: Elpidia rigida Théel, 1882. COMPOSITION: 4 species. DISTRIBUTION: circumtropical and in part in mid latitudes. Atlantic — 1, Indian — 2, Pacific — 3 species. Psychroplanes obsoleta (Herouard, 1899) Kolga obsoleta Hérouard, 1899: 170, fig. 1; Hérouard, 1902: 41–42, pls. VI: 11–15, VIII: 16, 18. Psychroplanes obsoleta — Gebruk, 1990: 82–84, fig. 29. Kolga foliacea Hérouard, 1912: 5–6, figs. 3–4. Peniagone foliacea — Hérouard, 1923: 86–87, pls. I: 31, IX: 1–2. Peniagone nybelini — Madsen, 1953: 157–158, fig. 5. Peniagone obsoleta Hansen, 1975: 134–135, fig. 56. LOCALITIES: “Princesse-Alice”, St. 753, “Princesse-Alice II”, St. 1306; “Vityaz”, St. 7943. DISTRIBUTION: Atlantic and Pacific (one locality east of the Japanese Trench). DEPTH RANGE: 4275–6096 m. Family Pelagothuriidae Genus Enypniastes Théel, 1882 TYPE SPECIES: Enypniastes eximia Théel, 1882. COMPOSITION: 1 species. DISTRIBUTION: cosmopolitan (Gebruk, 1989). Enypniastes eximia (Théel, 1882) Enypniastes eximia Théel, 1882a: 56–57, pl. VIII: 6– 7 (full synonymy in Gebruk, 1989). LOCALITIES: “Meteor”, 1988/6 MOC1-11: B7; “Discovery”, St. 9756#6 (Gebruk et al., 1997); numerous other localities in the North-East Atlantic (Billett et al., 1985); “James Cook” ECOMAR, St. JC048/43 Dive 174 (Rogacheva et al., 2013). DISTRIBUTION: cosmopolitan, in the NorthEast Atlantic very common (Billett et al., 1985; Gebruk, 1989). DEPTH RANGE: from surface to 5433 m. Family Deimatidae Genus Deima Théel, 1879 TYPE SPECIES: Deima validum Théel, 1879. COMPOSITION: 1 species with 2 subspecies. DISTRIBUTION: cosmopolitan. Deima validum validum Théel, 1879 Deima validum Théel, 1879: 5, figs. 36–38 ; Théel, 1882a: 68–70, pls. XVIII, XIX, XXXI: 4–9, XXXVI: 4, 165 XXXVII: 8, XLIII: 7, XLIV: 13, XLVI: 5 ; Sluiter, 1901b: 60. Deima validum validum — Hansen, 1967: 488–490, fig. 5; Rogacheva et al., 2013: 595, fig. 18e. Deima fastosum Théel, 1879: 5–6, figs. 1–3; Théel, 1882a: 71– 73, pls. XX, XXI: 1, XXXI: 10–13, XXXV: 7–10, XXXVI: 7, XXXVII: 3, XLIII: 2–3, 5, XLVI: 8. Deima blakei Théel, 1886b: 1–2, figs. 1–2; Koehler, Vaney, 1905: 55–57, pl. XI: 13–15; Hérouard, 1923: 40–41, pls. V: 7, VI: 5; Deichmann, 1930: 115–116, pls. X: 7–11, XI: 1–3; Deichmann, 1940: 198–199. Deima atlanticum Hérouard, 1898: 88–89, figs. 1–2; Hérouard, 1902: 32–35, pls. III: 3, IV: 18, V: 1–5, VIII: 26–29, Grieg, 1921: 4, pl. I: 2–3. Deima mosaicum Ohshima, 1915: 233–234, Ohshima, 1916–1919. LOCALITIES: “Princesse Alice”, St. 753, “Princesse Alice II”, St. 3006; BIOGAS (1972–1974), “Jean Charcot“, St. 2, 3, 4, 5 (Sibuet, 1977); “Challenger” (1981–1982), 51216#1, 51414#2, 51415#1, 51608#1; “G.O. Sars” MAR-ECO cruise, St. 40-367 (Gebruk, 2008); “James Cook” ECOMAR, Sts. JC011/23, JC048/36 Dive 171, JC048/43 Dive 174 (Rogacheva et al., 2013). DISTRIBUTION: cosmopolitan. DEPTH RANGE: 724–4820 m. In the NorthEast Atlantic: 2779–4360 m. Genus Oneirophanta Théel, 1879 TYPE SPECIES: Oneirophanta mutabilis Théel, 1879. COMPOSITION: 3 species. DISTRIBUTION: Pacific — 1, Indian — 1, cosmopolitan — 1 species. Oneirophanta mutabilis mutabilis Théel, 1879 Oneirophanta mutabilis Théel, 1879: 6–7, figs. 4–6; Théel, 1882a: 62–68, pls. XXI: 2, XXII, XXXI: 1–3, XXXVI: 1–2, 8–11, XXXVII: 4, 13, XXXVIII: 11–12, XL: 1–3, XLI: 1–2, 4, XLII: 9, XLIII: 1,6, XLV, XLVI: 6–7; Perrier, 1902: 374–380, pl. XVIII: 10–15; Clark, 1913: 232; Grieg, 1921: 5, pl. II: 1–2; Hérouard, 1923: 39–40, pls. IV: 10, V: 3–4; Ekman, 1927: 364–366, figs. 1–2; Agatep, 1967b: 63–65, pl. X: 1–7. Oneirophanta mutabilis mutabilis — Hansen, 1967: 485–488, figs. 3–4. Oneirophanta alternata Perrier, 1900: 117–118; Perrier, 1902: 380–386, pls. XIV: 3–4, XVIII: 16–22. Oneirophanta alternata var. talismani Perrier, 1902: 386–388, fig. 6. LOCALITIES: “Princesse Alice II”, St. 2986; “Travailleur” and “Talisman” St. 133, 134, 135, 137, 138, 139 (Perrier, 1902); “M. Sars”, St. 10 (Grieg, 1921); “Discovery” St. 12930#26, 12930#46, 12930#64, 12930#78, 13078#29, 13078#37, 13200#09, 13925#1, 14317#1; “Challenger” St. 54301#06 (Witbaard et al., 2003); BIOGAS (1972– 1974) “Jean Charcot“, St. 1, 2, 3, 4, 5; INCAL WS 04, CP 10, CP 11 (Massin, 1984); “G.O. Sars”, MARECO cruise, St. 40-367, 46/372 (Gebruk, 2008). DISTRIBUTION: cosmopolitan; in the NorthEast Atlantic very common. DEPTH RANGE: 3200–6000 m. 166 A.V. Gebruk et al. Order Aspidochirotida Family Synallactidae1 Genus Bathyplotes Östergren, 1896 TYPE SPECIES: Bathyplotes natans (M. Sars, 1868). COMPOSITION: ca. 20 species. DISTRIBUTION: Pacific — 7; Indo–Pacific — 2; Indian Ocean — 5; Caribbean — 1; North Atlantic — 1; Antarctic — 1; cosmopolitan — 2 species. Bathyplotes natans (M. Sars, 1868) Holothuria natans M. Sars, 1868: 20. Bathyplotes natans — Östergren, 1896: 352–353, pl. 18: 27–35; Östergren, 1902: 6; Ludwig, 1901: 137; Grieg, 1921: 7; Mortensen, 1927: 384–385, textfigs. 228: 2, 229; Deichmann, 1930: 100–102, pl. 9:1,–2, 9; Deichmann, 1954: 386; Heding, 1942a: 10–12, textfigs. 10, 11: 1–10, textfig. 12: 1–2; Pawson, 1963b: 90–94, pl. 7: 1–7; Pawson, 1965: 16– 18, fig. 4; Gage et al., 1985: 194; Harvey et al., 1988: 183; Miller, Pawson, 1990: 4; Madsen, Hansen, 1994: 79–82, figs.48–50, map 20; Rowe, Gates, 1995: 328; Liao, 1997: 73–74, fig. 39. Stichopus natans — G.O. Sars, 1872: 30; M. Sars, 1877: 58, pl. 7: 18–41; Théel, 1886a: 193; Bell, 1892: 51. Stichopus pourtalesii Théel, 1886a: 4. Stichopus tizardi Théel, 1882b: 696; Bell, 1892: 51; Koehler, 1895: 48–50, textfigs. 13–14. Bathyplotes tizardi — Östergren, 1896: 354; Ludwig, 1901: 138, pl. 12: 3–4, pl. 18: 19; Perrier, 1902: 350; Mitsukuri, 1912: 35–39, textfig. 8; Grieg, 1921: 4. Bathyplotes pourtalesii — Mortensen, 1927; Deichmann, 1930: 102, pl. 9: 3–7; Deichmann, 1940: 186, pl. 31: 34; Deichmann, 1954: 386; Rowe, Gates, 1995: 328. Bathyplotes fallax Östergren, 1896: 355, pl. 18: 44. Herpysidia reptans Perrier, 1898: 247– 248. Bathyplotes reptans — Perrier, 1902: 352–358, pl. 12: 3–4, pl. 18: 1–9; Mortensen, 1927: 384. Bathyplotes assimilis Koehler, Vaney, 1905: 25–26, pl. 3: 3, pl. 10: 1– 3. Bathyplotes papillosus — Koehler, Vaney, 1905: 28– 29, pl. 10: 21–24. Non Bathyplotes papillosa Koehler, Vaney, 1905: 2 (list). Bathyplotes patagiatus Fisher, 1907: 688–690, pl. 72: 1, 1a–k. Bathyplotes östergreni Ohshima, 1915: 225–226, pl. 8: 3a–d. Bathyplotes heterostylides Heding, 1942a: 12–13, textfig. 12: 3–5, textfig. 13: 1– 15. Bathyplotes bipartitus Hérouard, 1923: 34–36. LOCALITIES: “Talisman”, St. 42 (Perrier, 1902); “James Cook” ECOMAR, Sts. JC037/19 (Rogacheva et al., 2013). DISTRIBUTION: essentially cosmopolitan. In the East Atlantic known from the Lofoten Islands (Norway) to Cap Vert. Reported from the eastern and northern Gulf of Mexico; the Rockall Trough (Gage et al., 1985). Also recorded from Wanganella Bank (New Zealand), Tasman Sea, East China Sea (Liao, 1997) and Japan (Ohta, 1983). DEPTH RANGE: 193–2750 m. Genus Benthothuria Perrier, 1898 TYPE SPECIES: Benthothuria funebris Perrier, 1898. 1 Synonymy after Solís-Marín, 2003. COMPOSITION: 5 species. DISTRIBUTION: Indian Ocean — 3; Indo– Pacific — 1; Atlantic — 1 species. Benthothuria funebris Perrier, 1898 Benthothuria funebris Perrier, 1898: 1665; Perrier, 1899: 248; Perrier, 1902: 365–371; Mortensen, 1927: 378; Deichmann, 1930: 91; Heding, 1940: 363–364; Heding, 1942a: 6; Gage et al., 1985: 194–195; Harvey et al., 1988: 183. Localities (not complete): “Ingolf”, St. 36; “Discovery”, St. 9640; St. 13914#1; “Challenger” (1973– 1985), St. ES 27, ES 28, AT 119, AT 121, ES 147, ES 152, ES 169, SWT 12, SWT 27, OTSB 51001, OTSB 51301 (Gage et al., 1985); ES 204, AT 284, ES 285, AT 286, OTSB3/85/5 (Harvey et al., 1985); “G.O. Sars”, MAR-ECO cruise, Sts. 40/367, 52/ 374, 54/377, 64/381, 66/383, 74/387 (Gebruk, 2008); “James Cook” ECOMAR, Sts. JC011/23, JC011/ 75, JC011/101, JC037/15, JC037/19, JC037/27, JC037/61 (Rogacheva et al., 2013). DISTRIBUTION: off North-West Africa and South-West Greenland; Rockall Trough, North-East Atlantic (Harvey et al., 1988); off Moroccan coast (specimens collected by the “Talisman”); off Sudan (Red Sea) and Senegal (Deichmann, 1930); NorthWest Atlantic (Heding, 1942a). DEPTH RANGE: 782–3757 m. It appears that this species occurs deeper in more northerly latitudes in the North-East Atlantic (2700–3757 m) than off North-West Africa (782–1230 m) (Billett, 1991). Genus Paroriza Hérouard, 1902 TYPE SPECIES: Paroriza prouhoi Hérouard, 1902 by monotypy. COMPOSITION: 4 species. DISTRIBUTION: Pacific — 2, Atlantic — 2 species. Paroriza pallens (Koehler, 1895) Stichopus pallens Koehler, 1895: 50–52, textfig. 15. Paroriza pallens — Clark, 1922: 46 (list). Type locality: “Caudan”, St. 15. LOCALITIES: BIOGAS (1972–1974) “Jean Charcot”, St. 2 (Sibuet, 1977); “G.O. Sars”, MARECO cruise, St. 66/384 (Gebruk, 2008). DISTRIBUTION: North-East Atlantic; northern Spain north to the Porcupine Seabight. DEPTH RANGE: 1300–2900 m (Koehler, 1896; Khripounoff, Sibuet, 1980). Paroriza prouhoi Hérouard, 1902 Paroriza prouhoi Hérouard, 1902: 24–25, pl. 7: 1–2, pl. 8: 30; Hérouard, 1923: 29–30, pl. 2: 3–4; Perrier, 1901: 323; Mortensen, 1927: 386; Deichmann, 1930: 112–113. Mesothuria expectans — Perrier, 1899: 245; Perrier, 1902: 317–320; Deichmann, 1930: 92, Tortonese, 1949: 11 (list); Madsen, 1953: 168 (list). Deep-sea fauna of European seas. Holothuroidea LOCALITIES: “Princesse Alice”, St. 753 (Hérouard, 1902); “Princesse-Alice II”, St. 2964 (Hérouard, 1923); “Challenger”, St. 50811#1 (SolísMarín, 2003); “Talisman”, St. 136; BIOGAS (1972– 1974) “Jean Charcot”, St. 2 (Sibuet, 1977); “Discovery”, St. 14317#1 (Wigham et al., 2003). DISTRIBUTION: North-East Atlantic, off the Azores, the Bay of Biscay and the Porcupine Seabight; Mediterranean Sea. DEPTH RANGE: 4080–4880 m. Genus Paelopatides Théel, 1886 TYPE SPECIES: Paelopatides confundens Théel, 1886. COMPOSITION: 19 species (see Solís-Marín, 2003). DISTRIBUTION: Pacific — 6, Atlantic — 3, Indian Ocean — 8, Indonesia — 2 species. Paelopatides atlantica Hérouard, 1902 Pelopatides atlantica Hérouard, 1902: 16, pl. 1: 15; Hérouard, 1923: 16; Mortensen, 1927: 388 (key); Deichmann, 1930: 106. LOCALITIES: known only from the type locality, “Princesse Alice”, St. 527, off the Azores. DEPTH RANGE: 4020 m. Paelopatides gigantea (Verrill, 1884) Benthodytes gigantea Verrill, 1884: 216; Verrill, 1885: 538, pl. 11: 31 a–b; Grieg, 1921: 9, textfigs. 5–6. Paelopatides gigantea — Deichmann, 1930: 104–106 (partim); Miller, Pawson, 1990: 5. Paelopatides agassizi Théel, 1886a: 3. Paelopatides grisea — Perrier, 1902: 381; Mortensen, 1927: 388. LOCALITIES: “Discovery” St. 9775#3; BIOGAS (1972–1974) “Jean Charcot” St. 5, 6. DISTRIBUTION: North Atlantic Ocean (Deichmann, 1930; Pérez et al., 1984). DEPTH RANGE: 2454–2653 m. Paelopatides grisea Perrier, 1898 Paelopatides grisea Perrier, 1898: 1665; Perrier, 1899: 248; Perrier, 1902: 361–365; Mortensen, 1927: 388; Heding, 1940: 351; Billett et al., 1985: 407; Gage et al., 1985: 195–196; Harvey et al., 1988: 183; Miller, Pawson, 1990: 5; Gebruk, 2008: 50, 51; Rogacheva et al., 2013: 593, fig. 18a. Paelopatides gigantea — Deichmann, 1930: 104–106 (partim); Sibuet, 1977: 554; Miller, Pawson, 1990: 35. LOCALITIES: “Talisman”, St. 134; “Discovery” St. 9775#3 (Billett et al., 1985); “G.O. Sars”, MAR-ECO cruise, Sts. 66/383, 68/384, 72/386 (Gebruk, 2008); “James Cook” ECOMAR, Sts. JC011/ 23, JC011/101, JC011/106, JC011/111, JC037/15, JC037/19, JC037/61, JC048/06 Dive 159, JC048/15 Dive 161 (Rogacheva et al., 2013). DISTRIBUTION: North Atlantic, Caribbean. DEPTH RANGE: 1695–4060 m. 167 Genus Molpadiodemas Heding, 1935 sensu O’Loughlin et Ahearn, 2005 TYPE SPECIES: Molpadiodemas acaudum Heding, 1935, junior synonym of Pseudostichopus atlanticus Perrier, 1898, according to O’Loughlin, 2002. COMPOSITION: 16 species. DISTRIBUTION: Pacific — 3, South Pacific and Antarctic — 1, Antarctic — 3, Atlantic — 4, Indonesia — 1, cosmopolitan — 4 species. Molpadiodemas atlanticus (Perrier, 1898) Pseudostichopus atlanticus Perrier, 1898: 165; Perrier, 1899: 246–247; Perrier, 1902: 333–337, pl. 17: 15–20; Mortensen, 1927: 386–387; Deichmann, 1930: 87–88; O’Loughlin, 2002: 315. Molpadiodemas atlanticus — Heding, 1940: 353–359; O’Loughlin, Ahearn, 2005: 153, fig. 3a, b, 4a–d, 6a, b. Meseres atlanticus — O’Loughlin, 1998: 497; Thandar, 1999: 376–379, fig. 4. Molpadiodemas acaudum — Heding, 1935: 78–80, Pl. 6, fig. 1, 2; Heding, 1940: 354–355, 357; Deichmann, 1940: 209, 211; Heding, 1942a: 4–5. Meseres acaudum — O’Loughlin, 1998: 497. LOCALITIES: Talisman, St. 134, off the Azores (type locality); “Ingolf”, St. 37, (type locality of Molpadiodemas acaudum, Heding, 1935), West European Basin, 4780–4795 m (O’Loughlin, Ahearn 2005). DISTRIBUTION: North Atlantic, South-East Atlantic, Pacific. DEPTH RANGE: 2610–4795 m. Molpadiodemas depressus (Hérouard, 1902) Pseudostichopus depressus Hérouard, 1902: 15–16, pl. 2: 15–18, Mortensen, 1927: 387, Deichmann, 1930: 88. Molpadiodemas depressus — O’Loughlin, Ahearn, 2005: 158–159, fig. 3h, 4i–l. Pseudostichopus (Pseudostichopus) depressus — Heding, 1940: 359 (key). LOCALITIES: “Princesse Alice”, St. 753, between Portugal and Azores; West European Basin, 4426–4435 m (O’Loughlin, Ahearn 2005). North Atlantic, 40°N, 20°W to 20°N, 75°W, 2995–4360 m (Madsen, 1953). DISTRIBUTION: North and South Atlantic Ocean; West European Basin, Sargasso Sea, off Bahamas and West Indies. DEPTH RANGE: 1353–5690 m. Molpadiodemas involutus (Sluiter, 1901) Meseres involutus Sluiter, 1901a: 11–12; Sluiter, 1901b: 49–50, pl. 8, fig. 6; Perrier, 1902: 359; O’Loughlin, 2002: 306, fig. 2e, tables 1, 3, 4. Molpadiodemas involutus — O’Loughlin, Ahearn, 2005: 160–161, fig. 3m–o, 4u–x. Pseudostichopus globigerinae Hérouard, 1923: 23–25, pl. 4: 6; Mortensen, 1927: 386, 388; Deichmann, 1930: 87, 90; Sibuet, 1977: 554; Gebruk, 2008: 50, 51. Pseudostichopus (Pseudostichopus) globigerinae — Heding, 1940: 353, 357; Imaoka, 1978: table 1–1; Thandar, 1992: 167. Meseres globigerinae — O’Loughlin, 2002: 305. Pseudostichopus (Pseudostichopus) dilatorbis Imaoka, 1978: 378–380, 384, fig. 1b–e, tabl. 1–1. Pseudostichopus villo- 168 A.V. Gebruk et al. sus — Hansen, 1956: 47–48 (partim, non Pseudostichopus villosus Théel, 1886). LOCALITIES: “Chain”, [USNM 1005340], West European Basin, 4426–4435 m (O’Loughlin, Ahearn, 2005); BIOGAS (1972–1974) “Jean Charcot”, St. 1, 2, 4 (Sibuet, 1977); BIOICE, St. 3070, 3073, 3169 (unpublished data). DISTRIBUTION: North and South Atlantic, including Antarctic waters (Scotia Sea), North Pacific, East China Sea, Indonesia and Tasman Sea. DEPTH RANGE: 400–5801 m. Molpadiodemas villosus (Théel, 1886) Pseudostichopus villosus Théel, 1886a: 170–171, Hérouard, 1896: 164 (distribution list), Hérouard, 1902: 11–13, pl. 2: 1–3, pl. 7: 3, Hérouard, 1923: 23, Vaney, 1908: 407– 408, Grieg, 1921: 4, Hérouard, 1923: 23, Mortensen, 1927: 387, 388, Deichmann, 1930: 89, Heding, 1940: 353–360, Hansen, 1956: 47–48. Molpadiodemas villosus — O’Loughlin, Ahearn, 2005: 164–165, tabl. 3, figs. 2a, e, 7d– f, 8q–t, 12a–d. Pseudostichopus villosus var. violaceus Théel, 1886a: 172, pl. 10: 6b. Meseres villosus — O’Loughlin, 1998: 497; O’Loughlin, 2002: 313, figs. 3a–b. Type locality: “Challenger”, St. 156, 62°26′S, 95°44′E, 3594 m, 26–02–1874. LOCALITIES: “Princesse Alice II”, St. 1306 (Hérouard, 1923); “Michael Sars”, St. 53 (Grieg, 1921); “Discovery”, St. 9638#2, 10115#1, 11908#44, 11908#68, 12930#37, 12930#46, 12930#60, 12930#64, 12930#78, 13078#29, 13078#31, 13078#37, 13200#9, 13200#27, 13200#35, 13200#60, 13200#88, 13200#99, 13368#23, 13368#51, 13369#1, 13627#10, 13627#23, 13907#1, 13925#1; “Challenger” St. 50514#1, 50515#1, 50711#1, 50812#2, 50910#1, 51414#1, 52216#8, 52403#25, 52701#42, 53201#1, 53201#24, 53201#28, 53205#3,54301#6, 54301#8, 54901#2, 54901#5, 54901#7, 54902#1, 54903#1; “Meteor” St. 52602#1 (Solís-Marín, 2003). DISTRIBUTION: Bay of Biscay, Mediterranean Sea (Tortonese, 1949); North-West Atlantic (Solís–Marín, 2003); North-East Atlantic (Deichmann 1930, referred as P. atlanticus); Caribbean Sea, (Deichmann 1930, referred as P. atlanticus); South Atlantic, African coast (Thandar, 1999). The Pacific Ocean records need validation. DEPTH RANGE: 896–7000 m (Deichmann, 1930; Hansen, 1956). Molpadiodemas violaceus (Théel, 1886) Pseudostichopus villosus var. violaceus Théel, 1886: 172, pl. 10, fig.6b. Molpadiodemas violaceus — O’Loughlin, Ahearn, 2005: 165, figs. 1e, i, 2f, 7g–i, 8u–x (list); Rogacheva et al., 2013: 592. Pseudostichopus villosus Théel, 1886: 170–171 (partim — syntypes of Pseudostichopus villosus from HMS “Challenger” Sts. 61, 147, 325. LOCALITIES: “James Cook”, St. JC011/75 (Rogacheva et al., 2013). DISTRIBUTION: Antarctic and Subantarctic, North Atlantic and South Pacific. DEPTH RANGE: 2196–6354 m, in the North Atlantic 2605–5212 m (Rogacheva et al., 2013). Genus Pseudostichopus Théel, 1886 sensu O’Loughlin et Ahearn, 2005 TYPE SPECIES: Pseudostichopus mollis Théel, 1886 (subsequent designation by Fisher, 1907). COMPOSITION: 11 species. DISTRIBUTION: North-East Atlantic — 2, Pacific — 4, Indian Ocean — 1, Antarctic — 1, cosmopolitan — 3 species. Pseudostichopus aemulatus Solís-Marín et Billet, 2004 in Solís-Marín et al., 2004 Pseudostichopus aemulatus Solís-Marín et al., 2004: 1079–1081, fig. 1A–I; O’Loughlin, Ahearn, 2005: 169– 170, Fig. 11a, b, 12m; Pseudostichopus sp. — Billett et al., 2001: 336. LOCALITIES: “Challenger”, St. 52701#42, 54901/5, 54901/7, 54901/9, 54903/1, 54905/1; “Discovery”, St. 12930#46, 12930#78, 13078#29, 13627#10. DISTRIBUTION: North-East Atlantic, Porcupine Abyssal Plain. DEPTH RANGE: 4350–4850 m. Pseudostichopus peripatus (Sluiter, 1901) sensu O’Loughlin et Ahearn, 2005 Meseres peripatus Sluiter, 1901a: 10–11; Sluiter, 1901b: 48–49, pl. 5 fig. 5, pl. 8 fig.7; Perrier, 1902: 359. Pseudostichopus peripatus — O’Loughlin, Ahearn, 2005: 174–175, Figs. 1f, 10f–h, 11i–1, 12g, h. Pseudostichopus occultatus — Hérouard, 1902: 14–15, pl. 2 figs 4–14 (partim, illustrated; non Pseudostichopus occultatus Marenzeller, 1893). Pseudostichopus occultatus var. plicatus Koehler, Vaney, 1905: 9–10, pl. 3 fig. 8, pl. 9 figs 1– 3; Heding, 1940: 353 (non Pseudostichopus occultatus Marenzeller, 1893). Plicastichopus plicatus — Heding, 1940: 354–356; Heding, 1942a: 6. Pseudostichopus propinquus Fisher, 1907: 691–693, pl. 71 fig. 3, pl. 72 fig. 2, pl. 73 fig. 3, pl. 74 fig. 1, pl. 76 fig. 3; Imaoka, 1978: 382. Pseudostichopus (Trachostichopus) propinquus — Heding, 1940: 357; Imaoka, 1978: table 1–1; Imaoka, 1990: 148, 152. Meseres propinquus — O’Loughlin, 2002: 309. Pseudostichopus aleutianus Ohshima, 1915: 228, pl. 8 figs 5a–c; Imaoka, 1978: 380. Pseudostichopus (Trachostichopus) aleutianus — Heding, 1940: 353–359; Imaoka, 1978, table 1–2. Pseudostichopus unguiculatus — Ohshima, 1915: 230–231, pl. 8 figs 7a–c; Imaoka, 1978: 384; Rowe, 1995: 285. Pseudostichopus (Pseudostichopus) unguiculatus — Heding, 1940: 353–360; Imaoka, 1978: table 1–1; Imaoka, 1990: 152; Thandar, 1992: 167. Pseudostichopus marenzelleri Hérouard, 1923: 25; Mortensen, 1927: 287–288; Deichmann, 1930: 90; Gebruk, 2008: 50, 51. Pseudostichopus (Pseudostichopus) marenzelleri — Heding, 1940: 353–359; Imaoka, 1978: table 1–1; Thandar, 1992: 167. Pseudostichopus lapidus Hérouard, 1923: 26–28, pl. 4 fig. 5; Mortensen, 1927: 387; Deich- Deep-sea fauna of European seas. Holothuroidea mann, 1930: 90. Pseudostichopus (Pseudostichopus) lapidus — Heding, 1940: 353–360. Plicastichopus ingolfi Heding, 1942a: 5–6, figs 4–5, pl. 1 figs 4–5. Meseres ingolfi — Rowe, 1995: 285. Pseudostichopus (Trachostichopus) tuberculatus — Imaoka, 1990: 149–152, pl. p. 149, fig. P. 15. LOCALITIES: Princesse Alice”, St. 527 (type locality of Pseudostichopus lapidus Hérouard, 1923), St. 650 (type locality of Pseudostichopus marenzelleri Hérouard, 1923), “Ingolf”, St. 18, (type locality of Plicastichopus ingolfi Heding, 1942a); “G.O. Sars”, MAR-ECO cruise, Sts. 68/384, 72/386; “James Cook” ECOMAR, Sts. JC011/23; JC011/75; JC011/ 101; JC011/106; JC037/15; JC037/19; JC037/27; JC037/61; JC037/67; JC037/70, JC048/24 Dive 165 (Rogacheva et al., 2013). DISTRIBUTION: Mediterranean, North and South Atlantic Ocean, Indo-Pacific Region, North and South Pacific Ocean, Scotia Sea, Antarctic Ocean, Ross Sea, Weddell Sea. DEPTH RANGE: 134–5453 m. Scotothuria Hansen, 1978 TYPE SPECIES: Scotothuria herringi Hansen, 1978. COMPOSITION: 1 species. Scotothuria herringi Hansen, 1978 Scotothuria herringi Hansen, 1978: 34–37, figs. 1– 9; Billett et al., 1985: 406–407, fig. 5; Miller, Pawson, 1990: 4. LOCALITIES: “Discovery”, St. 9022#1, 9801#90, 10651#1, 10651#2, 11121#22, 11121#23 (Billett et al., 1985). REMARKS: Benthopelagic species; swims using udulating movements of ventro-lateral fringe. DISTRIBUTION: Numerous records in the East Atlantic in pelagic trawls from 20 to 3900 m above seafloor. The only record in the benthic trawl at the Galathea St. 238 in the Indian Ocean off Kenia. DEPTH RANGE: 1250–4980 m. Genus Synallactes Ludwig, 1893 TYPE SPECIES: Synallactes alexandri Ludwig, 1893. COMPOSITION: 22 species. DISTRIBUTION: Pacific — 11, Atlantic — 6, Indian Ocean — 4, Antarctic — 1 species. Synallactes crucifera Perrier, 1898 Synallactes crucifera Perrier, 1898: 1665; Perrier, 1899: 247; Perrier, 1902: 339–345, pl. 12: 5–6, pl. 17: 21– 35; Mortensen, 1927: 378 (key), textfig. 224, fig. 9; Deichmann, 1930: 106 (passim); Deichmann, 1940: 186, pl. 31: 5–6; Gebruk, 2008: 50, 51. LOCALITIES: “Talisman”, St. 40. DISTRIBUTION: North-East Atlantic Ocean, coast of Morocco; Caribbean Sea, off Venezuela; 169 Mid-Atlantic Ridge, south of the Charlie-Gibbs Fracture Zone. DEPTH RANGE: 2160–2340 m. Synallactes longipapillata Sibuet, 1978 Synallactes longipapillata Sibuet, 1978: 311–318, pls. 1–3. LOCALITIES: BIOGAS (1972–1974) “Jean Charcot”, St. 2, DS 40. Known only from the type locality. DISTRIBUTION: North-East Atlantic, Bay of Biscay. DEPTH RANGE: 3345 m. Family Mesothuriidae Genus Mesothuria Ludwig, 1894 TYPE SPECIES: Mesothuria multiples Ludwig, 1894. COMPOSITION: 27 species (Solis-Marin, 2003; Gebruk et al., 2012). DISTRIBUTION: Pacific — 9; Indian Ocean — 6; Atlantic — 6: Caribbean — 3; Antarctic to North Atlantic — 1; Antarctic — 1; cosmopolitan — 1 species. Mesothuria bifurcata Hérouard, 1901 Mesothuria bifurcata Hérouard, 1901: 40; Hérouard, 1906: 4–6, pl. 2: 3; Jangoux, Massin, 1986: 84 (list); O’Loughlin et al., 1994: 553–554; O’Loughlin, 2002: 313, 315; Gebruk et al., 2012: 283–284, fig. 5. Mesothuria (Mesothuria) bifurcata — Heding, 1940: 333; Heding, 1942a: 8, text fig. 7, figs. 1–6. LOCALITIES: “Ingolf”, St. 18 (Heding, 1942a); BIOICE, St. 2914, 3077, 3572, 3574 (unpublished data). DISTRIBUTION: Antarctic species. One record in the North Atlantic. DEPTH RANGE: 320–2337 m (Heding, 1942a; O’Loughlin, 2002). Mesothuria cathedralis Heding, 1940 Mesothuria (Allantis) cathedralis — Heding, 1940: 338–340, textfig. 5; Mesothuria (Penichrothuria) cathedralis, — Heding, 1942a: 8–9, textfig. 8, figs. 1–5; non Mesothuria (Allantis) candelabra — Heding, 1940: 334– 335, textfig. 3, figs. 1–6; Mesothuria cathedralis, Gage et al., 1985: 196; Gebruk, 2008: 50, 51; Gebruk, 2012: 284, 286–289, fig. 7. LOCALITIES: “Ingolf”, St. 18; “Challenger”, St. ES 10; “G.O. Sars”, MAR-ECO cruise, St. 72/ 386 (Gebruk 2008); “James Cook”, Sts. JC011/17, JC011/23, JC011/75, JC011/101, JC037/15, JC037/ 19, JC037/27, JC037/27, JC037/70, JC048/16 Dive 162 (Rogacheva et al., 2013). DISTRIBUTION: Atlantic; Gulf of Guinea, off Cape Bojador (Morocco), Irminger Basin off southern Greenland and Gulf of Mexico. 170 A.V. Gebruk et al. Mesothuria milleri Gebruk et Solís-Marín, 2012 in Gebruk et al., 2012 Mesothuria milleri Gebruk et al., 2012: 274–283, fig. 4. Holothuria verrilli (Théel, 1886a) — Marenzeller, 1893b: 7–9, pl. 1: 2, pl. 2: 2. Mesothuria verrilli (Théel, 1886) — Östergren, 1896: 345; Perrier, 1902: 307–312, pl. 16: 22–31; Hérouard, 1923: 10–13; Mortensen, 1927: 381–382, fig. 224: 4–5; Grieg, 1921: 4. LOCALITIES: numerous throughout east Atlantic, e.g. “Talisman”, Sts. 75, 129, 134, 135, 136; BIOICE, Sts. 2861, 3070, 3572. DISTRIBUTION: This species is widely distributed in the North-Eastern North Atlantic: off northwest Africa, the Canary Islands (Perrier, 1902; Grieg, 1932) the Azores (Hérouard, 1902; 1923; Perrier, 1902), the Bay of Biscay (Koehler, 1896; Perrier, 1902) the Porcupine Seabight (Mortensen, 1927), Goban Spur, Rockall Trough (Harvey et al., 1988), off British Isles, the Azores, the Canary Islands and Morocco (Perez et al., 1984). DEPTH RANGE: 550–4255 m (Perrier, 1902). Perrier was uncertain about two records from deeper than 4000 m. Excluding these records, the lower limit is 3018 m (Hérouard, 1923). In the North–East Atlantic this species was found to be most abundant between 1430 and 1530 m (Billett, 1988). Mesothuria intestinalis (Ascanius, 1805) Koehler, 1927; Sibuet, 1977) and the western Atlantic (Deichmann, 1930). The species may occur in the Gulf of Mexico (Deichmann, 1954). DEPTH RANGE: 18–4255 m (Tortonese 1949). The shallowest records come only from cold waters off Norway. Mesothuria maroccana Perrier, 1899 Mesothuria maroccana Perrier, 1899: 245; Perrier, 1902: 312–317, pl. 16: 32–35; Hérouard, 1923: 17; Deichmann, 1930: 97, pl. 7: 2–7; Deichmann, 1940: 191; Deichmann, 1954: 385; Grieg, 1921: 4; Hansen, 1956: 46, fig. 14a; Gebruk, 2008: 50, 51; Gebruk et al., 2012: 301– 303, figs. 9a, b; Rogacheva et al., 2013: 592, fig. 17d. Holothuria intestinalis var. verrilli — Hérouard, 1896: 163. Mesothuria murrayi var. grandipes Hérouard, 1923: 15, pl. 4: 7–9. Mesothuria (Mesothuria) maroccana — Heding, 1940: 333, Heding, 1942a: 8. LOCALITIES: “Talisman”, Sts. 35, 39 (Perrier, 1902); “Ingolf”, Sts. 18, 65 (Heding, 1942a); “Michael Sars”, St. 88 (Grieg, 1921); BIOGAS (1972– 1974) “Jean Charcot”, Sts. 1, 2, 6 (Sibuet, 1977); “G.O. Sars”, MAR-ECO cruise, Sts. 40/367, 42/ 368, 50/373, 64/381 (Gebruk, 2008); “Challenger”, St. 50518 #1, “Discovery”, St. 10106 #1 (unpublished); “James Cook” ECOMAR, Sts. JC011/17, JC011/75, JC037/15, JC037/19, JC037/27, JC048/ 40 Dive 173 (Rogacheva et al., 2013). DISTRIBUTION: Caribbean Sea, Gulf of Mexico, North Atlantic. DEPTH RANGE: 700–3120 m. Holothuria intestinalis Ascanius, 1805: 5, pl. 45; Marenzeller, 1893a: 15; Marenzeller, 1895: 21; Ludwig, 1893: 174; Théel, 1886: 209; Théel, 1901: 1–38, pls. 1–2: 1–19, textfigs. 1–12; Bell, 1892: 48–49, pl. 5: 3; Hérouard, 1896: 163. Mesothuria intestinalis — Östergren, 1896: 347–351, pl. 18: 1–26; Östergren, 1902: 6–7; Perrier, 1902: 304–307, text figs. 1–2, pl. 16: 19–21; Ludwig, 1901: 139; Théel, 1902: 4–34, pl. 1–2: 1–19 and 12 text figs.; Hérouard, 1923: 10, pl. 5: 5–6; Mortensen, 1927: 381, text fig. 225, 228: 3; Koehler, 1927: 240, pl. 15: 3 (partim); Deichmann, 1930: 94–95, pl. 6: 9–10; Deichmann, 1954: 385–386; Tortonese, 1949: 13 (list); Tortonese, 1965: 69–70, text fig. 26; Sibuet, 1974a: 795; Harvey et al., 1988: 184; Madsen, Hansen, 1994: 76–79, figs. 46–47, map 29; Massin, 1996: 43; Gebruk et al., 2012: 291–300, figs. 1, 9c–d. Fistularia mollis Sars, 1835: 40. Thyonidium scabrum Sars, 1868: 19–20. Holothuria verrilli — Marenzeller, 1893b: 7–9, pl. 1: 2, pl. 2: 2. Allantis intestinalis var. verrilli — Hérouard, 1902: 18– 21, pl. 1: 3–6 (partim); Allantis intestinalis — Heding, 1942a: 7, text fig. 6. Non Allantis intestinalis var. verrilli — Hérouard, 1902 (= Mesothuria verrilli). Mesothuria (Allantis) intestinalis — Heding, 1942a: 7, textfig. 6, figs. 1–7; Panning, 1952: 123–125, figs. 1–3. Mesothuria verrilli — Tortonese, 1952: 228; Tortonese, 1961, pl. 1, text fig. 1. Mesothuria triradiata Heding, 1942b: 217–218, textfig. 1, figs. 1–6. Holothuria lactea Théel, 1886a: 6–7; Théel, 1886b: 183–184, pl. 9: 15. Zygothuria lactea — Gebruk et al., 2012: 310–321, figs. 13, 14. Mesothuria lactea (Théel) — Sluiter, 1901a: 25; Hérouard, 1902: 21–23, pl. 1: 17–19; Hérouard, 1923: 13–15, pl. 4: 1–3; Mortensen, 1927: 382– 383 (partim), fig. 227. Mesothuria (Zygothuria) lactea (Théel) — Heding, 1940: 340–341, fig. 7. Mesothuria (Zygothuria) lactea lactea (Théel) — Heding, 1942a: 9– 10, fig. 9. Mesothuria lactea (Théel) — Perrier, 1902: 322–327 (partim), pl. 17: 1–6; Deichmann, 1930: 108– 111, pl. 8: 8–9; Deichmann, 1940: 190–191; Deichmann, 1954: 386. DISTRIBUTION: widely distributed in the North-East Atlantic (Harvey et al., 1988) from off North-West Africa (Hérouard, 1923) to the coast of Norway, although nowhere in the deep sea does it appear to be particularly common. A few specimens are known from the Mediterranean (Perrier, 1902; DISTRIBUTION: Cosmopolitan species. However, some old records may not be reliable because most authors did not recognize the species oxysclera and these two species could have been confused. This is especially likely with the records from the Gulf of Mexico and the Caribbean (Deichmann, Genus Zygothuria Perrier, 1898 TYPE SPECIES: Zygothuria lactea (Théel, 1886) designated by Hérouard (1902). COMPOSITION: 6 species (Solis-Marin, 2003; Gebruk et al., 2012). DISTRIBUTION: Atlantic — 3, Pacific — 2, cosmopolitan — 1 species. Zygothuria lactea (Théel, 1886) Deep-sea fauna of European seas. Holothuroidea 1930) where Z. oxysclera occurs. Numerous records from the eastern North-East Atlantic, also known from the North-West Atlantic, off West Africa, SouthEast Atlantic, Indo-Malayan archipelago and New Zealand waters (Gebruk et al., 2012). DEPTH RANGE: reliable bathymetric range from 694 m (Sluiter, 1901a) to 2102 m (Hérouard, 1902). M. lactea var. spinosa (Heding, 1940), recorded from 5108 m, differs significantly from Z. lactea both in the shape and the size of ossicles and should probably be assigned to Z. candelabri (Hérouard, 1923). Another deep record, from 4400 m (Sibuet, 1977), was not supported by morphological details and hence is not reliable. In the Porcupine Seabight the species was most abundant between 1430 and 1930 m (for detailed distribution, see Billett, 1988). Zygothuria candelabri (Hérouard, 1923) Mesothuria candelabri Hérouard, 1923: 17–19, pl. 1: 1–10; Madsen, 1953: 153. Non Mesothuria (Allantis) candelabri — Heding, 1940 [= Mesothuria (Penichrothuria) cathedralis]. Zygothuria candelabri — Deichmann, 1930: 111; Gebruk et al., 2012: 322–325, fig. 16. REMARKS: This species was described from some fragments. The tables are indicated to have relatively few holes in the skin and excessively long diverging spines on top of the spire, with a few small teeth scattered along the sides of the spines. It is likely identical with Z. lactea var. spinosa and Z. thomsoni (Gebruk et al., 2012). Some authors have mis-spelt “candelabri” as “candelabra”. LOCALITIES: “Princesse Alice”, St. 2986, Bay of Biscay; Swedish Deep-Sea Expedition, St. 387; Porcupine Seabight, Porcupine Abyssal Plain (Billett, 1991). DISTRIBUTION: North-East Atlantic Ocean. DEPTH RANGE: 3890–4870 m. Order Dendrochirotida Family Cucumariidae Genus Staurocucumis Ekman, 1927 TYPE SPECIES: Cucumaria liouvillei Vaney, 1914. COMPOSITION: 6 species. DISTRIBUTION: Atlantic Ocean — 1, Antarctic — 4, cosmopolitan — 1 species. Staurocucumis abyssorum (Théel, 1886) Cucumaria abyssorum Théel, 1886a: 66–67, pl. 4, fig. 6, pl. 16, fig. 6; von Marenzeller, 1893b: 14; Ludwig, 1894: 122–125, pl. 9, figs. 28–29, pl. 13, figs. 1–5; Grieg, 1921: 11, text-fig. 9; Ludwig, Heding, 1935: 179; Cherbonnier, 1941: 93–96, 101, fig. 1, 3j, n, o, p. Staurocucumis abyssorum — Ekman, 1927: 385–387; Clark, Deichmann, 1936: 566; Hansen, 1988: 302–303, fig. 1; Rogacheva et al., 2013: 590, fig. 17a. Abyssocucumis abyssorum — Heding, 1942a: 33–35, figs. 34–36; Gage et al., 1985: 191. Cucumaria abyssorum var. grandis Théel, 1886a: 67–68, pl. 5 fig.1. Cucumaria abyssorum var. 171 hyalina — Théel, 1886a: 68–69, pl. 4 fig. 7. Cucumaria sluiteri — Ohshima, 1915: 263, pl. 10, fig. 21a, b. Cucumaria ingolfi Deichmann in Mortensen, 1927: 396. Staurocucumis ingolfi — Clark, Deichmann, 1936: 567. Cucumaria albatrossi Cherbonnier, 1941: 96–101, 103, fig. 2, 3a–I, k–m. LOCALITIES: “Hirondelle”, St. 248, (von Marenzeller, 1893b); “M. Sars”, St. 88 (Grieg, 1921); “Challenger”, SWT 15, (Gage et al., 1985); “G.O. Sars”, MAR-ECO cruise, Sts. 52/374, 54/377 (Gebruk, 2008); “James Cook” ECOMAR, Sts. JC011/ 17, JC011/23, JC037/15, JC37/019, JC37/027 (Rogacheva et al., 2013). DISTRIBUTION: cosmopolitan (Atlantic, Pacific, Indian Oceans and Antarctic). DEPTH RANGE: 869–4025 m, Ekman (1927) referred to a juvenile specimen from 385 m in the Antarctic, but Hansen (1975) believed that it might have been misidentified. Genus Echinocucumis M. Sars, 18592 TYPE SPECIES: Echinocucumis typica M. Sars, 1859 (=Eupyrgus hispidus Barrett, 1857) by monotypy. COMPOSITION: 7 species. DISTRIBUTION: Atlantic — 2, Atlantic and Pacific — 1, Pacific — 2, Indian Ocean — 1, Antarctic — 1 species. Echinocucumis hispida (Barrett, 1857) Eupyrgus hispidus Barrett, 1857: 46, pl. 4 fig 1 a, b. Echinocucumis hispida — Mortensen, 1927: 404, figs. 242–1i, 243; Deichmann, 1930: 150, pl. 18 fig. 9; Ludwig, Heding, 1935: 167; Heding, 1942a: 29–31, figs. 31, 32; Panning, 1949: 454; Pawson, 1965: 8–10, fig. 2; Sibuet, 1977: 554; Gage et al., 1985: 194; Thandar, 1999: 370– 373, fig. 2. Echinocucumis typica M. Sars, 1861: 102, pl. 10 figs. 11–20, pl. 11 figs. 1–17; Théel, 1886a: 118–119; Théel, 1886b: 9, fig. 3; Hérouard, 1923: 118–127, pl. VII figs. 7, 10. Cucumaria typica Ludwig, 1901: 149. TYPE LOCALITY: Nordland (western Norway, Lofoten area). LOCALITIES: “Princesse Alice II”, St. 1116 (Hérouard, 1923); BIOGAS, St. 4 (Sibuet, 1977). DISTRIBUTION: Atlantic, off east coast of New Zealand. DEPTH RANGE: 50–3257 m. Family Ypsilothuriidae Genus Ypsilothuria E. Perrier, 1886 TYPE SPECIES: Ypsilothuria talismani E. Perrier, 1886. COMPOSITION: 2 species DISTRIBUTION: Atlantic (Y. talismani); Atlantic and Pacific (Y. bitentaculata). 2 Smirnov (2012: 822) transferred this genus from family Ypsilothuriidae into family Cucumariidae. 172 A.V. Gebruk et al. Ypsilothuria talismani talismani E. Perrier, 1886 Ypsilothuria talismani Perrier E., 1886: 286, fig.294; Perrier R., 1902: 518, textfig. 12, Pl. XII figs. 9–10. Ypsilothuria talismani talismani — Heding, 1942a: 26– 27, textfig. 24, textfig. 25, 5, 6, textfig. 26, 1, 3, textfig. 27, 3, textfig. 28, 1, 2, textfig. 29; Gebruk, 2008: 50, 51. Echinocucumis typica var. abyssalis — Koehler, 1896: 118, fig. 22. LOCALITIES: “Travailleur” (1881) Dragage 1 (Perrier, 1902); “G.O. Sars”, MAR-ECO cruise, Sts.: 54377, 64381, 66383 (Gebruk, 2008); BIOICE, Sts.: 2854, 2863, 3169, 3170, 3172, 3176 (unpublished data). DISTRIBUTION: North Atlantic. DEPTH RANGE: 480–3527 m. Ypsilothuria bitentaculata attenuata E. Perrier, 1886 Ypsilothuria attenuata Perrier E., 1886: 285, fig. 203; Perrier R., 1902: 522, textfig. 13. Ypsilothuria bitentaculata attenuata — Heding, 1942a: 28, pl. 2 figs. 1–10; textfig. 25 1–4, 9–10, textfig. 26 4–7, textfig. 27 2, 5, textfig. 30; Gage et al., 1985: 192; Harvey et al., 1988: 183; Alvà, 1991: 459–460; Massin, 1996: 44–46, fig. 1A–G, fig. 2A–B. Ypsilothuria bitentaculata — Thandar, 1999: 373–376, fig. 3, 14D–F. Sphaerothuria bitentaculata — Deichmann, 1930: 152, pl. 19 figs. 4–5. Echinocucumis typica — Clark, 1923: 418 (non Echinocucumis typica M. Sars, 1859 = E. hispida Barrett, 1857). Ypsilothuria talismani — Mortensen, 1932: 49; Tyler, Gage, 1983: 609–616 (non Ypsilothuria talismani E. Perrier, 1886) LOCALITIES: “Challenger” (1973–1985): Sts. ES 06, ES 08, ES 10, ES 12, ES 27, ES 34, ES 52, ES 54, ES 55, ES 56, ES 57, ES 59, ES 111, ES 118, AT 119, AT 121, ES 122, ES 129, AT 130, ES 137, ES 140, ES 141, ES 143, AT 144, ES 147, SBC 150, AT 151, ES 152, AT 153, AT 154, ES 164, AT 167, ES 169, AT 171, ES 172, AT 175, ES 176, AT 177, ES 180, AT 181, ES 184, ES 185 (Gage et al., 1985; Harvey et al., 1988), AT 186 (Gage et al., 1985), ES 190 (Gage et al., 1985; Harvey et al., 1988), AT 191, AT 195, ES 197, AT 198 (Gage et al., 1985), ES 200 (Gage et al., 1985; Harvey et al., 1988), AT 201, ES 202 (Gage et al., 1985), ES 204 (Gage et al., 1985; Harvey et al., 1988), ES 218, ES 231, ES 232, AT 233, ES 244, AT 245, ES 266, AT 267, AT 271, AT 273, AT 282, ES 283, ES 285, AT 286, AT 288, ES 289 (Tyler, Gage, 1983; Gage et al., 1985; Harvey et al., 1988). DISTRIBUTION: possibly cosmopolitan. DEPTH RANGE: 375–3231 m. DISTRIBUTION: 3 species known from the Atlantic, others from the Indo-Pacific, Antarctic and Arctic. REMARKS. In the list below the shallow-water species Molpadia borealis M. Sars, 1859, is not included, although it was mentioned from the depth ca. 2000 m by Harvey et al. (1988) (“Challenger” St. AT107A). Molpadia musculus Risso, 1826 Molpadia musculus Risso, 1826: 293; Clark, 1908: 165 (complete list of references till 1907); Ohshima, 1915: 250; Hérouard, 1923: 123, p. 132, pl. 5 fig. 1; Deichmann, 1930: 198, pl. 23, figs. 4–7; Heding, 1931b: 279; Deichmann, 1940: 225, pl. 40 figs. 1–15; Deichmann, 1947: 342; Deichmann, 1954: 405; Djakonov at al., 1958: 376; Cherbonnier, 1965: 17, pl. 7 figs. i–q, pl. 8 figs. a–j; Pawson, 1965: 11, fig. 3 1, 4–6; Tortonese, 1965: 98, fig. 42; Pawson, 1977: 100, fig. 1–3, 4a–e, map 1 (complete list of references); Pawson et al., 2001: 317–318, fig. 2a– c. Ankyroderma musculus — Perrier, 1902: 529–533, pl. 22 figs. 16–22. Molpadia violacea — Studer, 1876: 464; Pawson, 1963a: 15–16, pl. 3, figs. 4–8; Pawson, 1965: 12– 13. Trochostoma violaceum Théel, 1886a: 42, pl. 2, fig. 6; Lampert, 1889: 842; Perrier, 1905: 65. Haplodactyla violacea — Heding, 1931b: 280. Eumolpadia violacea — Heding, 1935: 42, textfig. 8 7–10, pl. 5 fig. 10, pl. 7 fig. 3, pl. 8 fig. 4; Ludwig, Heding, 1935: 144–145, textfig. 11. Ankyroderma danielsseni Théel, 1886a: 39, pl. 2 fig. 6. Ankyroderma loricatum Perrier, 1898: 1666; Perrier, 1902: 535, pl. 22 figs. 23–28; Hérouard, 1923: 133. Eumolpadia asaphes Heding, 1935: 42–44, textfig. 9, pl. 5 fig. 9, pl. 7 fig. 2. LOCALITIES: “Princesse Alice”, St. 515 (Hérouard, 1902, 1923); “Talisman”, St. 97 (Perrier, 1902); “Ingolf”, St. 36 (Heding, 1935); “G.O. Sars”, MAR-ECO cruise, Sts. 54/377, 64/381, 66/383 (Gebruk, 2008); BIOICE, St. 3172 (unpublished data); “James Cook” ECOMAR, Sts. JC011/17, JC011/ 101, JC011/111, JC037/15, JC037/19, JC037/27, JC037/61 (Rogacheva et al., 2013). DISTRIBUTION: cosmopolitan. DEPTH RANGE: 35–5205 m (Pawson, 1977). Molpadia blakei (Théel, 1886) Order Molpadiida Family Molpadiidae Genus Molpadia (Cuvier, 1817) Risso, 1826 Trochostoma blakei Théel, 1886b: 16, pl. 1 fig. 8; Perrier, 1902: 525, pl. 22 fig. 3–6. Molpadia blakei — Clark, 1908: 33, 168; Deichmann, 1930: 196–197, pl. 22 fig. 19–23; Deichmann, 1940: 224, pl. 38 fig. 6–8; Sibuet, 1977: 554; Khripunov, Sibuet, 1980: 187; Gage et al., 1985: 205; Tyler et al., 1987: 388; Harvey et al., 1988: 191; Pawson et al., 2001: 320–322, fig. 3. Trochostoma blakei var. excentrica Hérouard, 1923: 136–137, pl. 9 figs. 3–9. Trochostoma angulatum Hérouard, 1923: 136, pl. 3 fig. 6, pl. 8 fig. 3. Trochostoma grossularia Hérouard, 1923: 137–139, pl. 9 figs. 13–16, 21–32. Paratrochostoma spiniferum Heding, 1935: 72–76, fig. 21, pl. 4 figs. 13–14, pl. 5 figs. 20–21. TYPE SPECIES: Molpadia musculus Risso, 1826. COMPOSITION: about 58 nominal species. LOCALITIES: “Princesse Alice II”, St. 3006 (Hérouard, 1923 as Trochostoma blakei var. excen- Deep-sea fauna of European seas. Holothuroidea trica); “Princesse Alice, St. 757, “Princesse Alice II”, St. 2994 (Hérouard, 1923 as Trochostoma grossularia); “Challenger” St.: ES 143, AT 144, SBC 150, AT 151, ES 152, AT 153, AT 167, AT 171, AT 175, ES 176, AT 177, AT 181, ES 184, ES 185, AT 191, AT 195, ES 197, AT 198, ES 200, AT 201, ES 202, ES 207 (Gage et al., 1985), ES 283, AT 288, 3/ 85/5 OTSB (Harvey et al., 1988); “Discovery” St. 9638#2, “Challenger” St. 50515, 50711, 50811 (Tyler et al., 1987); Porcupine Abyssal Plain, 2470–4795 m (Billett, 1991); BIOGAS (1972–1974) “Jean Charcot” St. 2-6 (Sibuet, 1977); BIOICE, St. 2863, 3176 (unpublished data). DISTRIBUTION: North Atlantic (Hérouard, 1923; Heding, 1935; Sibuet, 1977; Gage et al., 1985; Harvey et al., 1988; Billett, 1991), Gulf of Mexico and off east Coast of Brazil (Pawson et al., 2001). DEPTH RANGE: 1727–5270 m. 173 Hedingia albicans (Théel, 1886) Deichmann, 1938 Trochostoma albicans Théel, 1886a: 44, pl.11, fig. 3; Perrier, 1902: 526–528, pl. 22, figs. 7–8; Koehler, Vaney, 1905: 89–90, pl. 13, figs. 9–10. Hedingia albicans — Deichmann, 1938: 112; Deichmann, 1940: 216–217; Harvey et al., 1988: 191–192; Pawson et al., 2001: 324–325, fig. 4C; Bohn, 1985: 33. Caudina albicans — Clark, 1908: 37, 174–175, pl. 10 fig. 12; Heding, 1931b: 283. Haplodactyla albicans — Heding, 1935: 65–67, fig. 18, 19 (erroneously 21), pl. 4 fig. 9, pl. 5 fig. 17, pl. 8 fig. 10. Trochostoma albicans var. glabra Théel, 1886a: 46. Caudina arenata var. armata Théel, 1886b: 17 [nec Caudina arenata (Gould, 1841)]. LOCALITIES: Porcupine Abyssal Plain, 1430– 2790 m (Billett, 1991). DISTRIBUTION: North Atlantic, Mediterranean, Indian Ocean, New Zealand. DEPTH RANGE: 494–3200 m. Genus Cherbonniera Sibuet, 1974 TYPE SPECIES: Cherbonniera utriculus Sibuet, 1974. COMPOSITION: monotypic. DISTRIBUTION: North Atlantic. Cherbonniera utriculus Sibuet, 1974 Cherbonniera utriculus Sibuet, 1974b: 1443–1445, pl. 1; Sibuet, 1977: 554; Gage et al., 1985: 205; Tyler et al., 1987: 385; Harvey et al., 1988: 191; Pawson et al., 2001: 315–317, fig. 1A–D. LOCALITIES: POLYGAS and BIOGAS 4 (1972 and 1974), DS 20, DS 21, DS 25, DS 51, DS 55, DS 56, DS 59; “Challenger”: St. ES 02, ES 06, ES 08, ES 10, ES 27, ES 28, ES 34, ES 55, ES 56, ES 57, SBC 58, ES 59, ES 111, ES 118, AT 119, ES 129, ES 135, ES 137, ES 140, ES 143, ES 147, SBC 150, ES 164, ES 169, ES 172, ES 180, ES 185, SBC 188, ES 190, ES 204, SBC 205, ES 207 (Gage et al. 1985), SBC 174,ES 231, ES 283, ES 285 (Harvey et al., 1988); “Discovery“ 9576#14, “Discovery“ St. 50604#1, 10114#1, 10115#1 (Tyler et al., 1987); Porcupine Abyssal Plain, 2820–4140 m (Billett, 1991); BIOGAS Sts. 2, 3, 6 (Sibuet, 1977). DISTRIBUTION: Atlantic Ocean: Bay of Biscay (Sibuet, 1974b) Rockall Trough and adjacent areas (Gage et al., 1985; Harvey et al., 1988); Porcupine Abyssal Plain (Billett, 1991), off New York, east of Cape Hatteras, and north-east of the Falkland Islands (Pawson et al., 2001). DEPTH RANGE: 2039–5223 m. Family Caudinidae Genus Hedingia Deichmann, 1938 TYPE SPECIES: Trochostoma albicans Théel, 1886. COMPOSITION: 6 species (Deichmann, 1938). DISTRIBUTION: Atlantic — 1, Pacific — 4, cosmopolitan — 1 species. Family Gephyrothuriidae Genus Gephyrothuria Koehler et Vaney, 1905 TYPE SPECIES: Gephyrothuria alcocki Koehler et Vaney, 1905. COMPOSITION: monotypic. DISTRIBUTION: cosmopolitan. Gephyrothuria alcocki Koehler et Vaney, 1905 Gephyrothuria alcocki Koehler, Vaney, 1905: 78–80, pl. 5, fig. 6–8; Clark, 1908: 22, 186; Hérouard, 1923: 33; Deichmann, 1930: 202; Heding, 1935: 78; Deichmann, 1940: 209–211; Heding, 1940: 358; Hansen, 1956: 48; O’Loughlin, 1998: 495–496, fig. 1; Rogacheva et al., 2013: 613–614, figs. 16c, d. 19l. Himasthlephora glauca — Clark, 1908: 22, 40–41, 185 pl. 13, fig. 1–4; Heding, 1935: 78. Gephyrothuria glauca — Hérouard, 1923: 33; Deichmann, 1930: 202–203; Deichmann, 1940: 209–211; Heding, 1940: 358; Hansen, 1956: 48. Gephyrothuria europeensis — Hérouard, 1923: 30–33, pl. IX: 10a, b; Deichmann, 1940: 209–211; Heding, 1940: 358; Sibuet, 1977: 554. REMARKS: Species Gephyrothuria europeensis Hérouard, 1923 was synonymized with Gephyrothuria alcocki Koehler et Vaney, 1905 by O’Loughlin (1998). LOCALITIES: “Princesse-Alice II”, St. 2990 (type locality of Gephyrothuria europeensis); “James Cook” ECOMAR, Sts. 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