Abstract
Most of the taxonomically reliable internal and microstructural characters (e.g. jaws, dental plate, genital plates, vertebrae) of the recent Ophiuroidea are studied using SEM on a broad comparative basis for the first time, including examination of the arm spine articulation shape in 178 species from 105 genera and 16 families encompassing all major ophiuroid generic diversity. Numerous taxonomic contradictions caused by “over-applying” of external characters to traditional ophiuroid systematics are found and analyzed. Among newly applied microstructural characters, the shape of the arm spine articulations is found to be of great importance for ophiuroid taxonomy at all levels, from order to species. An identification key of the ophiuroid families based exclusively on the shape of the arm spine articulations is presented. Major genera of Ophiacanthidae were studied in order to delineate this family. The group of taxa, traditionally known as the ophiacanthid subfamily Ophiotominae (Paterson, 1985) that was apparently intermediate between Ophiomyxidae and Ophiacanthidae, including the genera Amphilimna Verrill, 1899, Ophiocymbium Lyman, 1880, Ophiodaces Koehler, 1922, Ophiodelos Koehler, 1930, Ophiolimna Verrill, 1899, Ophiologimus H.L. Clark, 1911, Ophiomedea Koehler, 1906, Ophiophrura H.L. Clark, 1911, Ophiopristis Verrill, 1899, Ophioprium H.L. Clark, 1915, Ophiosparte Koehler, 1922, Ophiotoma Lyman, 1883, Ophiotrema Koehler, 1896 was studied in detail using most of available type specimens. In order to study interspecific variability and usefulness as a taxonomic marker of the arm spine articulations, four new species of the apparently ophiotomin genus Ophiocymbium are described: O. antarcticus sp. nov., O. ninae sp. nov., O. tanyae sp. nov. and O. rarispinum sp. nov. A new genus and species, which has affinities to Ophiotominae, Ophioplexa condita gen. et sp. nov. is described. It is demonstrated that many of the genera traditionally included in the subfamily Ophiotominae, e.g. the genera Ophiocymbium, Ophiologimus, Ophiophrura, Ophioprium and Ophioplexa condita gen. et sp. nov., belong to the family Ophiomyxidae instead of Ophiacanthidae. Another apparently intermediate taxon, Ophiorupta discrepans (Koehler, 1922) comb. nov. is also considered as an ophiomyxid. Several further genera with disputed taxonomic placement, e.g. Amphilimna, Ophiopsila, Ophiolimna, Ophioconis, were studied especially and their revised placement is proposed. The following genera are exluded from the family Ophiacanthidae: Amphilimna, Ophiocymbium, Opiodaces, Ophiodelos, Ophiologimus, Ophiophrura, Ophioprium and Ophiosparte. The previously proposed paraphyly of the family Ophiacanthidae (Smith et al., 1995) was to a great extent caused by including a number of genera from distantly related families. The relationship between extinct Oegophiurida and recent ophiuroids was analyzed. A remarkable similarity between arm spine articulations of some Paleozoic oegophiurids and the recent ophiomyxid Ophioscolex glacialis Müller & Troschel, 1842 was discovered. Oegophiurid groove spines are suggested to be homologous with the tentacle scales of the remaining Ophiuroidea. It is suggested that the family Ophiomyxidae thus may be related to some crown Oegophiurida that had already acquired fused vertebrae. The higher ophiuroid taxonomy, based on the genital plate patterns, is critically analyzed in the light of the present data. It is suggested that instead of earlier proposed numerous ophiuroid subgroups most ophiuroid families are closely related. It is suggested, that most of the ophiuroid families (includes Ophiomyxidae, Ophiacanthidae, Ophiodermatidae, Ophiocomidae, Ophionereididae, Ophiochitonidae, Amphilepididae, Amphiuridae, Ophiactidae, Ophiolepididae, Hemieuryalidae, Ophiotrichidae) form a compact group with numerous intermediate taxa even between apparently very different families, whereas the family Ophiuridae and the traditional order Euryalida are more distantly related to the rest of Ophiuroidea. An appropriate name for this higher ophiuroid group will be suggested after a detailed analysis of other ophiuroid groups, which will be made in further publications of this series.
References
Ayres, W.O. (1851) Account of the structure of the Ophiuridae, and presented a description and drawings of a new species belonging to the genus Ophiolepis. Proceedings of the Boston Society of Natural History 4, 133–135.
Baker, A.N. (1979) Some Ophiuroidea from the Tasman Sea and adjacent waters. New Zealand Journal of Zoology 6, 21–51.
Baker, A.N. & Devaney, D.M. (1981) New records of Ophiuroidea (Echinodermata) from Southern Australia, including new species of Ophiacantha and Ophionereis. Transactions of the Royal Society of Southern Australia, 155–178.
Bartsch, I. (1982) Ophiuroidea (Echinodermata) from the Patagonian shelf. Mitteilungen aus dem hamburgischen zoologischen Museum und Institut 79, 211–250.
Bartsch, I. (1983) Ophiambix meteoris n. sp., ein neuer Schlangenstern aus der Iberische Tiefsee. Spixiana 6, 97–100.
Bartsch, I. (1987) Notes on Ophiuroidea (Echinodermata) from the northeastern Atlantic Ocean. I. Ophiacanthidae. Spixiana 10, 115–130.
Bell, F.J. (1892) A contributions to the classification of ophiuroids, with description of some new and little known forms. Proceedings of the Zoological Society of London, 175–183.
Bell, F.J. (1894) On the echinoderms collected during the voyage of H.M.S. Penguin and by H.M.S. Egeria, when surveying Macclesfield Bank. Proceedings of the Zoological Society of London, 392–413.
Bell, F.J. (1902) Echinoderms. In: Report on the collections of natural history made in the Antarctic regions during the voyage of the “Southern Cross”, 214–220.
Belyaev, G.M. & Litvinova, N.M. (1972) New genera and species of deep-sea Ophiuroidea. Bulletin of the Moscow Natural History Society 77, 5–20.
Belyaev, G.M. & Litvinova, N.M. (1976) New and rare species of the deep water Ophiuroidea from Pacific and Indian oceans. Proceeding of P.P. Shirshov Institute of Oceanology 99, 126–139.
Bernasconi, I. & D’Agostino, M.M. (1975) Equinodermos antarticos. III. Ofiuroideos. 1). Ofiuroideos del extremo norte de la Peninsula Antartica. Revista Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’. Hidrobiologia 4, 80–133.
Brock, J. (1888) Die Ophiuroidea-fauna des indischen Archipels. Zeitschrift wissentschaftliche Zoologie, 47, 465–439.
Bruzelius, N. (1805) Dissertatio sistens species cognitas asteriarum, quamr. sub praesidio D.M.And.J. Retzii, pro laurea modeste exhibet Nicolaus Bruzelius. Lund, pp. 1–37.
Byrne, M. (1994) Ophiuroidea. In Harrison F. W., Chia F. S., (Eds), Microscopic anatomy of invertebrates. Volume 14: Echinodermata. New York: Wiley-Liss, Chichester etc., 247–343.
Byrne, M. & Hendler, G. (1988) Arm structure of the ophiomyxid brittlestars (Echinodermata: Ophiuroidea: Ophiomyxidae). In Burke R. D., Mladenov P. V., Lambert P., Parsley R. L. (Eds)., Echinoderm biology. Proceedings of the Sixth International Echinoderm Conference, Victoria, 23–28 August 1987. Rotterdam: Balkema.
Cherbonnier, G. & Guille, A. (1972) Redescription et position systématique de l’Ophiure Ophiosphaera insignis Brock. Bulletin du Musum National d'Histoire Naturelle Paris, 3e serie, Zoologie 24, 280–282.
Cherbonnier, G. & Guille, A. (1978) Faune de Madagascar. 48 Echinoderms: Ophiures. Paris, Editions du C.N.R.S.
Clark, A.M. (1953) A revision of the genus Ophionereis (Echinodermata: Ophiuroidea). Proceedings of the Zoological Society of London 123, 65–94.
Clark, A.M. (1965) Japanese and other ophiuroids from the collections of the Munich Museum. Bulletin of the British Museum (Natural History), Zoology 13, 37–71.
Clark, A.M. (1970) Notes on the family Amphiuridae. Bulletin of the British Museum (Natural History), Zoology Series 19, 3–81.
Clark, A.M. (1974) Notes on some echinoderms from southern Africa. Bulletin of the British Museum of Natural History (Zoology) 26, 423–487.
Clark, A.M. & Courtman-Stock, J. (1976) The Echinoderms of Southern Africa. British Museum (Natural History) 766, 1–277.
Clark, A.M. & Rowe, F.W.E. (1971) Ophiuroidea. Monograph of shallow-water Indo-West Pacific Echinoderms. British Museum, London.
Clark, H.L. (1901) The echinoderms of Porto Rico. Bulletin of the United States Fisheries Commission 20, 231–263.
Clark, H.L. (1909) Scientific results of the trawling expedition of H.M.C.S. Thetis. Echinodermata. Memoirs of the Australian Museum, 4, 519–564.
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.
Clark, H.L. (1914) Growth changes in brittle stars. Publications of the Carnegie Institution 5, 93–126.
Clark, H.L. (1915) Catalogue of recent Ophiurans based on collections of the Museum of Comparative Zoology. Memoirs of the Museum of Comparative Zoology, Harvard 25, 165–376.
Clark, H.L. (1919) The distribution of the littoral Echinoderms of the west Indies. Papers Department Marine Biology Carnegie Institution Washington. (Publication 281) 13, 47–74.
Clark, H.L. (1925) Marine zoology of tropical central Pacific: Echinoderms other than sea-stars. Bulletin of the Bernice Pauahi Bishop Museum 27, 89–111.
Clark, H.L. (1933) A handbook of the littoral echinoderms of Porto Rico and the other West Indian Islands. Scientific Survey of Porto Rico and the Virgin Islands 16, 1–147.
Clark, H.L. (1938) Echinoderms from Australia. An account of collections made in 1929 and 1932. Memoirs of the Museum of Comparative Zoology, Harvard University 55, 1–596.
Clark, H.L. (1939) Ophiuroidea. Scientific Reports. The John Murray Expedition 6, 29–136.
Clark, H.L. (1941) Reports on the scientific results of the Atlantis expeditions to the West Indies under the joint auspices of the University of Havana and Harvard University. The echinoderms (other than holothurians). Memorias de la Sociedad cubana de Historia Natural "Felipe Poey" 15, 1–154.
Corliss, J.B., Dymond, J., Gordo, L.I., Edmond, J.M., von Herzen, R.P., Ballard, R.D., Green, K., Williams, D., Bainbridge, A., Crane, K. & van Andel, T.H.(1979) Submarine thermal hot springs on the Galapagos Rift. Science, 203, 1073–1083.
Danielssen, C. & Koren, J. (1877) Fra den norske Nordhavs Expedition. Nyt Magazin for Natur- videnskaberne, 32, 39 p.
Dearborn, J.H, Hendler, G. & Edwards, K.C. (1996) The diet of Ophiosparte gigas (Echinodermata: Ophiuroidea) along the Antarctic Peninsula, with comments on its taxonomic status. Polar Biology 16, 309–320.
Delle Chiaje, S. (1828) Memorie sulla storia e notomia degli animali senza vertebre del regno di Napoli 3, 1–232.
Devaney, D.M. (1970) Studies on ophiocomid brittlestars. I. A new genus (Clarkoma) of Ophiocominae with a re-evaluation of the genus Ophiocoma. Smithsonian Contributions to Zoology 51, 1–41.
Devaney, D.M. (1974) Shallow-water echinoderms from British Honduras, with a description of a new species of Ophiocoma (Ophiuroidea). Bulletin Marine Sciences 24, 123–164.
Düben, M.W. & Koren, J. (1846) Ofversigt af Skandinaviens Echinodermer. Kungliga Svenska Wetenskapsakademiens Handlingar 1844, 229–328.
Duncan, P.M. (1879) On some Ophiuroidea from the Korean Seas. Journal of the Linnean Society 14, 445–482.
Duncan, P.M. (1887) On the Ophiuridae of the Mergui Archipelago, collected for the Trustees of the Indian Museum, Calcutta by Dr. J. Anderson. Journal of the Linnean Society 21, 805–106.
Dyakonov, A.M. (1949) Opredelitel iglokozhikh dalnevostochnykh morei [Identification keys of Echinodermata of Far-Eastern seas of USSR]. Izvestia TINRO, 30, 4–127.
Dyakonov, A.M. (1954) Ophiury (Zmeekhvostki) Morei SSSR [Ophiurans of the seas of USSR]. Opredeliteli po faune SSSR, Volume 55. Zoological Institute, Moskva - Leningrad (In Russian).
Fell, H.B. (1952) Echinoderms from southern New Zealand. Zoology Publications from Victoria University College, Wellington 18, 1–37.
Fell, H.B. (1960) Synoptic keys to the genera of Ophiuroidea. Zoological Publications of the Victoria University Wellington 24, 1–40.
Fell, H.B. (1961) The fauna of the Ross Sea. Part 1. Ophiuroidea. New Zealand Department of Scientific and Industrial Research Bulletin 142, 9–79.
Fell, H.B. (1962) Evidence for the validity of Matsumoto’s classification of Ophiuroidea. Publications of the Seto Marine Biological Laboratory 10, 145–152.
Forbes, E. (1839) On the Asteriadae of the Irish Sea. Memoires Wernerian Society Edinburgh 8, 114–129.
Forbes, E. (1843) On the Radiata of the Eastern Mediterranean. Part I., Ophiuridae. Transactions of the Linnean Society of London 19, 143–153.
Forbes, E. (1852) Notes on animals of the class Echinodermata collected by Dr. Sutherland in Assisatnce Bay. Sutherland’s Journal of a voyage to Baffin’s Bay 2, 214–216.
Fujita, T., Ishida, Y., & Irimura, S. (1997) Ophiuroids collected from Deep waters of Suruga Bay, Central Japan. National Science Museum Monograph, No. 12, 257–268.
Fujita, T., Ishida, Y., Kato, T. & Irimura, S. (2004) Ophiuroids (Echinodermata) collected from the Oki islands in the Sea of Japan. Bulletin Natural Science Museum Tokyo, Ser. A., 30, 191–218.
Fujita, T. & Irimura, S. (2005) Ophiuroids (Echinodermata) collected by R/V Yoko–Maru off Southwestern Japan in the East China Sea. Deep-Sea fauna and pollutants in Nansei Islands. National Science Museum Monographs, No. 29, 357–384.
Fujita, T & Hendler, G. (2001) Description of a new species of Astrophiura (Echinodermata: Ophiuroidea) from Tosa Bay, Japan, and several of its remarkable anatomical characteristics. In Fujita, T, Saito H, Takeda M, eds. Deep-Sea fauna and pollutants in Tosa Bay. National Science Museum Monographs 20, 263–281.
Gonzalo, J. & Alarcón, C. (1968) Contribucion al conocimento de los ofiuroides chilenos. Gayana, 14, 3–64.
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 Memoires 16, 67–87.
Hendler, G. (1978) Development of Amphioplus abditus (Verrill) (Echinodermata: Ophiuroidea). II. Description and discussion of ophiuroid skeletal ontogeny and homologies. Biological Bulletin, Marine Biological Laboratory, Woods Hole, 154, 79–95.
Hendler, G. (1988) Ophiuroid skeleton ontogeny reveals homologies among skeletal plates of adults: a study of Amphiura filiformis, Amphiura stimpsonii and Ophiophragmus filograneus (Echinodermata). Biological Bulletin, Marine Biological Laboratory, Woods Hole, 174, 20–29.
Hendler, G. & Miller, J.E. (1984) Ophioderma devaneyi and Ophioderma ensiferum, new brittle-star species from the western Atlantic (Echinodermata: Ophiuroidea). Proceedings of the Bbiological Society of Washington, 97, 442–461.
Hendler, G. & Miller, J.E. (1991) Swimming ophiuroids - real and imagined. In Yanigasawa, T., Yasumasu, I., Oguro, C., Suzuki, N. and Motokawa, T. (Eds.), Biology of Echinodermata, 179–190.
Hendler, G., Miller, J.E. Pawson, D.L. & Kier, P.M. (1995) Sea stars, sea urchins and allies.Washington & London: Smithsonian Institution Press.
Hendler, G. & Turner, R.L. (1987) Two new species of Ophiolepis (Echinodermata: Ophiuroidea) from the Caribbean Sea and Gulf of Mexico: with notes on ecology, reproduction, and morphology. Contributions in Science, Natural History Museum Los Angeles County, 395, 1–14.
Hertz, M. (1927a) Die Ophiuroiden der deutscher Tiefsee-Expedition. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer Valdivia 22 1898–1899, 3, 59–122.
Hertz, M. (1927b) Die Ophiuroiden der Deutschen Sudpolar-Expedition 1901–1903. Deutsche Sudpolar-Expedition 1901–1903, 19, 1–56.
Hess, H. (1962) Mikropaläontologische Untersuchungen an Ophiuren. Eclogae Geologicae Helvetiae 55, 595–608.
Hotchkiss, F.H.C. (1977) Ophiuroid Ophiocanops (Echinodermata) not a living fossil. Journal Natural History 11, 377–380.
Hotchkiss, F.H.C. (1980) The early growth stage of a Devonian ophiuroid and its bearing on echinoderm phylogeny. Journal of Natural History 14, 91–96.
Hotchkiss, F.H.C. (1982) Ophiuroidea (Echinodermata) from Carrie Bow Cay, Belize. In: Rützler, K., MacIntyre, I. G. (Eds): The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, I: Structure and Communities. - Smithsonian Contributions to the Marine Sciences, 12, 387–412.
Hunter, R.L. (2007) Morphological cladistic analysis of Ophiurolepis Matsumoto, 1915 (Ophiurida: Ophiuridae) from the Southern Ocean. Zootaxa, 1401, 33–51.
Hyman, L.H. (1955) The Invertebrates: Echinodermata: The Coelomate Bilateria. MacGraw-Hill Book Company: New York, Toronto and London, Jaekel O. M. J.
Irimura, S. (1969) Supplemental report of Dr. Murakami’s paper on the ophiurans of Amakusa, Kyushu. Publications from the Amakusa Marine Biological Laboratory, 2, 37–48.
Irimura, S. (1981) Ophiurans from Tanabe Bay and its vicinity, with the description of a new species of Ophiocentrus. Publications of the Seto Marine Biological Laboratory, 26, 15–49.
Irimura, S. (1982) The brittle-stars of Sagami Bay. Biological Laboratory, Imperial Household, Japan.
Irimura, S. (1988) Ossicles of the stomach wall of Ophiuroidea and their taxonomic significance. In: Burke R.D., Mladenov P.V., Lambert P., Parsley R.L. (Eds), Echinoderm biology. Proceedings Sixth International Echinoderm Conference, Victoria, 23–28 August 1987. Rotterdam: Balkema.
Irimura, S. (1993) Ophiostriatus sexradiatus, a new species of Ophiuroidea from the North Mariana Islands. Bulletin from the National Science Museum, 19, 161–164.
Irimura, S. & Fujita, T. (2003) Intraspecific variation of vertebral ossicle morphology. In: Féral J.P., David B. (Eds), Echinoderm research 2001. Swets, Zeitlinger & Lisse, 161–166.
Irimura, S., Kubodera, T. & Ishida, Y. (1995) Catalogue of the specimens of the class Ophiuroidea (Echinodermata) donated by Dr. Seiichi Irimura in the National Science Museum. National Science Museum, Tokyo.
Jagt, J.W.M. (2000) Late Cretaceous-Early Palaeogene echinoderms and the K/T boundary in the southeast Netherlands and northeast Belgium – Part 3: Ophiuroids. Scripta Geologica 121, 1–45.
Jagt, J.W.M & Kutscher, M. (1998) Late Cretaceous ophiuroids from Germany and the Netherlands: An update. In: Mooi R., Telford M. (Eds), Echinoderms: San Francisco. Proceedings of the Ninth International Echinoderm Conference, San Francisco, 5–9 August 1996. Rotterdam: Balkema, 371–376.
Kesling, R.V. (1969) A new brittle-star from the middle Devonian Arkona shale of Ontario. Contributions from the Museum of Paleontology, University of Michigan 23, 37–51.
Kesling, R.V. (1970) Drepanaster wrighti, a new species of the brittle-star from the middle Devonian Arkona Shale of Ontario. Contributions from the Museum of Paleontology, University of Michigan 23, 73–79.
Kesling, R.V. (1972) Strataster devonicus, a new brittle-star with unusual preservation from the middle Devonian silica formation of Ohio. Contributions from the Museum of Paleontology, University of Michigan 24, 9–15.
Kesling, R.V. & Vasseur, D. (1971) Strataster ohionensis, a new Early Mississippian brittle-star, and the paleoecology of its community. Contributions of the Museum of Paleontology, University of Michigan 23, 305–341.
Koehler. R. (1896a) Resultats scientifiques de la compagne du Caudan dans le Golfe de Gascogne. Echinoderms. Annales de l’Universit de Lyon 26, 33–122.
Koehler, R. (1896b) Note prliminaire sur les ophiures recueillies pendant les campagnes de la Princesse Alice. Mémoires de la Société Zoologique de France, 9, 241–253.
Koehler, R. (1898) Echinides et Ophiures provenant des compagnes du yacht l’Hirondelle. Resultats des Campagnes Scientifiques accomplies sur son yacht par Albert Ier Prince souverain deMonaco 12, 1–78.
Koehler, R. (1901) Echinides et ophiures. Resultats du Voyage du S.Y. Belgica en 1897–1898–1899, Rapports Scientifiques, Zoologie, 3–42.
Koehler, R. (1904a) Ophiures de l’expdition du Siboga. I. Ophiures de mer profonde. Siboga Expeditie Monograph, 45a, 1–176.
Koehler, R. (1904b) Ophiures nouvelles ou peu connues. Mémoires de la Sociét Zoologique de France 17, 54–119.
Koehler, R. (1905) Ophiures littorales. Siboga-Expeditie Monograph, 45, 1–142,
Koehler, R. (1906). Description des Ophiures nouvelle recueillies par le Travailleur et Talisman pendant les campagnes 1880, 1881, 1882 et 1883. Mémoires de la Sociét Zoologique de France, 19, 5–34.
Koehler, R. (1907a) Ophiures. Expeditions Scientifiques du Travailleur et du Talisman 7, 245–311.
Koehler, R. (1907b) Revision de la collection des ophiures du Muséum d’Histoire Naturelle de Paris. Bulletin scientifiques de la France et de la Belgique 44, 279–351.
Koehler, R. (1914) A contribution to the study of Ophiurans of the United States National Museum. Bulletin of the United States National Museum 84, 1–173.
Koehler, R. (1922a) Contributions to the biology of the Philippines Archipelago and adjacent regions. Ophiurans of the Philippines seas and adjacent waters. Bulletin of the United States National Museum 100, 1–486.
Koehler, R. (1922b) Ophiuroidea. Scientific Reports Australasian Antarctic Expedition 1911–1914 8, 1–98.
Koehler, R. (1930) Ophiures recueillies par le Docteur Th. Mortensen dans les Mers d’Australie et dans l’Archipel Malais. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 89, 1–295.
Kutscher, M. & Jagt, J.W.M. (2000) Early Maastrichtian ophiuroids from Rügen (northeast Germany) and Møn (Denmark). Scripta Geologica 121, 45–107.
Lamarck, J.B. de (1816) Histoire naturelle des animaux sans vertèbres. Vol. 2. Verdière, Paris.
Lambert, P. & Austin, W.C. (2007). Brittle stars, sea urchins and feather stars of British Columbia, Southeast Alaska and Puget Sound. Royal BC handbook. Victoria, Canada. 150pp.
Leach, W. E. (1819) Descriptions des nouvelles espèces d'Animaux découvertes par le vaisseau Isabelle dans un voyage au pôle boréal. Journal de Physique, de Chimie et d'Histoire Naturelle 88, 462–467.
LeClair, E. (1996) Arm joint articulation in the ophiuran brittle-stars (Echinodermata: Ophiuroidea): a morphometric analysis of ontogenetic, serial, and interspecific variation. Journal of Zoology, London 240, 245–275.
Liao, Y. (1983) On Amphilimna polyacantha sp. nov. and the systematic position of A. multispina Koehler (Ophiuroidea). Chinese Journal of Oceanology and Limnology 1, 177–182.
Liao, Y. (1989) Two new species of the genus Amphilimna (Echinodermata: Ophiuroidea) from southern China. Chinese Journal of Oceanology and Limnology 7, 339–344.
Linnaeus, C. (1758) Systema naturae per regna tria naturae secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Holmiae, pp. 1–824.
Linnaeus, C. (1767) Systema Naturae, seu per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Holmiae, Laurentii Salvii, pp. 533–1327.
Litvinova, N.M. (1975) Ophiuroids of the Caribbean and Gulf of Mexico collected during 14th cruise of the R/V “Akademik Kurchatov. Proceeding of the P.P. Shirshov Institute of Oceanology 100, 196–204.
Litvinova, N.M. (1981) Brittle-stars (Ophiuroidea). Proceedings of the P.P. Shirshov Institute of Oceanology, 113–131.
Litvinova, N.M. (1984) A remarkable new species of the genus Ophiopyrgus (Echinodermata, Ophiuroidea), with comments on this genus. Zoologicheskii Zhurnal 58, 1585–1588.
Litvinova, N.M. (1989a) Ecological interpretation of structural peculiarities of arms in brittle stars 1. Variability of the arm vertebrae shape. Zoologicheskii Zhurnal 68, 97–105.
Litvinova, N.M. (1989b) Ecological interpretation of structural characteristics of arms in brittle stars. 2. Variability and functional morphology of vertebral joints in arms. Zoologicheskii Zhurnal 68, 41–55.
Litvinova, N.M. (1994) The life forms of Ophiuroidea (based on the morphological structures of their arms), In: David, B., Guille, A., Féral J.-P., Roux M. (Eds), Echinoderms through Time: Proceedings of the eighth International Echinoderm Conference, 6–10 September 1993, Dijon (France). Balkema, Rotterdam, 449–454.
Litvinova, N.M. (1996) Ecological interpretation of structural characteristics of brittlestars arms. 3. Arm forms and external skeletons (integuments). Zoologicheskii Zhurnal 75, 1676–1689.
Ljungman, A.V. (1867) Ophiuroidea viventia huc usque cognita enumerat. Öfversigt af Kongl. Vetenskaps-Akademiens Förhandlingar, Stockholm 23, 303–336.
Ljungman, A.V. (1872) Förteckning öfver uti Vestindien af Dr. A. Goës samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Öfversigt af Kongelige Vetenskaps-Akademiens Förhandlingar, Stockholm 28, 615–657.
Lonsdale, P. (1977) Clustering of suspension-feeding macrobenthos near abyssal hydrothermal vents at oceanic spreading centers. Deep-Sea Research, 24, 857–863.
Loriol, P.de. (1893) Echinoderms de la Baie d’Amboine. Revue Suisse de Zoologie, 1, 359–426.
Ludwig, H. (1901) 3 Klasse, Ophiuroidea, Schlangensterne. In: Bronn H.G. (Ed), Klassen und Ordnungen des Thier-Reichs, Band 2, Abtheilung 3, Buch 3, Leipzig: C.F. Winter’sche Verlagshandlung, 745–966.
Lütken, C.F. (1855) Bidrag til Kundskab om Slangestjernerne. I. Forelöbig Oversigt over Grönlandshavets Ophiurer. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 95–104.
Lütken, C. F. (1858) Systematisk oversigt over Gronlands echinodemer. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn, 1–7, 1–109
Lütken, C.F. (1859) Additamenta ad historiam Ophiuridarum. Beskrivelser af nye eller hidtil kun ufuldstaendigt kjendte Arter af Slangestjerner. Andem Afdeling. Kongelige Danske Vedenskabernes Selskab Skrifter Naturvidenskabelig of Mathematisk Afdeling, 5, 179–271.
Lütken, C.F. (1869) Additamenta ad historiam Ophiuridarum. 3. Beskrivende og kritiske Bidrag til Kundskab an Slangestjernerne. Kongelige Dansk Videnskabernes. Selskabs Skrifter. Kjøbenhavn 5, 22–109.
Lütken, C.F. (1872) Ophiuridarum novarum vel minus cognitarum descriptiones nonnullae. Berkrivelser af vogle rye eller mindre bekjendte Slangestjerner. Med nogle Bemaerkninger om Selvdelingen hos Straaldyrene. Oversigt over det Kongelige Danske Videnskabernes Selskabs Forhandlinger, 1872, 75–158
Lütken, C.F. & Mortensen, T. (1899) 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 U.S. Fish Commission steamer Albatross, during 1891, Lieut. Commander Z.L. Tanner, U.S.N., commanding. XXV. The Ophiuridae. Memoirs of the Museum of Comparative Zoology, Harvard University 23, 93–208.
Lyman, T. (1869) Preliminary report on the Ophiuridae and Astrophytidae dredged in deep water between Cuba and the Florida Reef by L. E. Pourtales. Bulletin Museum of Comparative Zoology Harvard Colledge 1, 309–354.
Lyman, T. (1875) Zoological results of the Hassler Expedition. II. Ophiuridae and Astrophytidae, including those dredged by the late Dr. William Stimpson. Ilustrated Catalogue Museum Comparative Zoology, Harvard College, 8, 1–34.
Lyman, T. (1878) Ophiuridae and Astrophytidae of the exploring voyage of H.M.S. Challenger, under Prof. Sir Wyville Thomson, F.R.S. Part 1. Bulletin of the Museum of Comparative Zoology, Harvard 5, 65–168.
Lyman, T. (1879) Ophiuridae and Astrophytidae of the exploring voyage of H.M.S. Challenger under Prof. Sir Wyville Thomson, F.R.S. Part 2. Bulletin of the Museum of Comparative Zoology, Harvard 6, 17–83.
Lyman, T. (1880) A structural feature, hitherto unknown among Echinodermata, found in deep-sea ophiurans. Anniversary Memoirs of the Boston Society of Natural History, 3–12.
Lyman, T. (1882) Report on the Ophiuroidea. Report on the Scientific Results of the Voyage of H.M.S. Challenger, Zoology 5(14), 1–386.
Lyman, T. (1883) Report on the Ophiuroidea. Bulletin of the Museum of Comparative Zoölogy at Harvard College 10, 227–287.
Madsen, F.J. (1967) Ophiuroidea. BANZ Antarctic Research Expedition 1929–1931, Reports. Series B (Zoology and Botany) 9, 123–145.
Madsen, F.J. (1970) West African ophiuroids. Atlantide Report, 11, 151–243.
Madsen, F.J. (1983) A review of the Ophioleucinae stat. rev. (Echinodermata, Ophiuroidea) with the erection of a new genus, Ophiostriatus. Steenstrupia 9, 29–69.
Martens, E. von (1867) Über vier neue Schlangensterne, Ophiuren. Monatsberichte der Königlichen Preussischen Akademie der Wissenschaften, 345–348.
Martynov, A.V. (2010) Structure of the arm spine articulation ridges as a basis for taxonomy of Ophiuroidea (a preliminary report). In: Harris L., ed, Proceedings of the Twelfth International Echinoderm Conference, Durham, NH, 6–12 August 2006. Rotterdam: Balkema, 233–239.
Martynov, A.V. & Litvinova, N.M. (2008) Deep-water Ophiuroidea of the northern Atlantic with descriptions of three new species and taxonomic remarks on certain genera and species. Marine Biology Research 4, 76–111.
Matsumoto, H. (1915) A new classification of the Ophiuroidea. Proceedings of the Academy of Natural Sciences Philadelphia 67, 43–92.
Matsumoto, H. (1917) A monograph of Japanese Ophiuroidea, arranged according to a new classification. Journal of the College Science, Imperial University Tokyo 38, 1–408.
McKnight, D.G. (2003) New brittle-stars (Echinodermata: Ophiuroidea) from New Zealand waters. Zootaxa 352, 1–36.
Medeiros-Bergen, D.E. (1996) On the stereom microstructure of ophiuroid teeth. Ophelia 25, 211–222.
Mortensen, T. (1925) Echinodermes du Maroc et de Mauritanie. Bulletin de la Societe des Science Naturelle du Maroc, 5, 178–187.
Mortensen, T. (1927) Handbook of the Echinoderms of the British Isles. London: Oxford University Press.
Mortensen, T. (1932) Papers from Dr. Th. Mortensen’s Pacific Expedition 1914–1916. LX. On an extraordinary ophiurid, Ophiocanops fugiens Koehler. Videnskabelige Meddellelser fra Dansk naturhistorisk Forening 93, 1–21.
Mortensen, T. (1933a). Ophiuroidea. Danish Ingolf Expedition 4, 5–121.
Mortensen, T. (1933b). Echinoderms of South Africa. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 93, 215–400.
Mortensen, T. (1933c) On an extraordinary ophiurid, Ophiocanops fugiens Koehler. With remarks on Astrogymnotes, Ophiopteron, and on an albino Ophiocoma. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening 93, 1–22.
Mortensen, T. (1933d) Biological observations on ophiurids, with descriptions of two new genera and four species. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening 93, 171–194.
Mortensen, T. (1936) Echinoidea and Ophiuroidea. Discovery Reports, 12, 199–348.
Müller, J., Troschel, F.H. (1842) System der Asteriden. 1. Asteridae. 2. Ophiuridae. Braunschweig.
Müller, J., Troschel, F.H. (1844) Beschreibung neuer Asteriden. Archiv für Naturgeschichte 10 (1-2), 178–185.
Müller, O. F. (1789) Zoologica Danica sev animalium Daniae et Norvegiae rariorum ac minus notorum descriptiones et historia, ed. 3, vol. 3, 1–71.
Murakami, S. (1942) Ophiurans of Izu, Japan. Journal of the Department of Agriculture, Kyûshu Imperial University, 7, 1–36.
Murakami, S. (1944) Report on the Ophiurans from off Ogasawara Islands and from off the Yaeyama group, Nippon. Journal of the Department of Agriculture, Kyûshu Imperial University, 7, 235–257.
Murakami, S. (1963) The dental and oral plates of Ophiuroidea. Transactions of the Royal Society of New Zealand, Zoology 4, 1–48.
Norman A.M. (1876) The Valarous Expedition. Proceedings of the Royal Society, 25, 202–215.
O’Hara, T.D. & Stöhr, S. (2006) Deep water Ophiuroidea (Echinodermata) of New Caledonia: Ophiacanthidae and Hemieuryalidae. In Richer De Forges, B. & Justine, J.-L. (Eds), Tropical Deep-Sea Benthos. Volume 24. Mémoires du Muséum national d’Histoire Naturelle 193, 33–141.
Paterson, G.L.J. (1980) A new abyssal genus of the family Ophiuridae (Echinodermata: Ophiuroidea). Bulletin of the British Museum (Natural History) Zoology. 38, 211–218.
Paterson, G.L.J. (1985) The deep-sea Ophiuroidea of the North Atlantic Ocean. Bulletin of the British Museum (Natural History), Zoology Series 49, 1–162.
Paterson, G.L.J & Baker, A. (1988) A revision of the genus Ophiambix (Echinodermata: Ophiuroidea) including the description of a new species. Journal of Natural History 22, 1579–1590.
Pearse, V.B., Pearse, J.S., Hendler, G. & Byrne, M. (1998) An accessible population of Ophiocanops, off NE Sulawesi, Indonesia. In: Mooi R. & Telford M. (Eds), Echinoderms: San Francisco, , Balkema, Rotterdam, 413–418.
Perrier, E. (1891) Traité de Zoologie. Librairie F Savy, FI Paris.
Rowe, F.W.E., Pawson, D.L. (1977) A catalogue of echinoderm type-specimens in the Australian Museum, Sydney. Records of the Australian Museum 30, 337–364.
Sars, M. (1857) Bidrag til kundshaben om Middelhavest Littoral-Fauna. Nyt Magazin for Naturvidenskaberne, 9, 110–164.
Sars, M. (1861) Oversigt af Norges Echinodermer. Christiania (Norway).
Sars, G.O. (1871) Nye Echinodermer fra den Norske kyst. Videnskabelige Selskabs Forhandlingar 1871, 1–31.
Schoener, A. (1967) Two new species of Amphitarsus (Ophiuroidea) from the western North Atlantic. Breviora, 269, 1–9.
Shackleton, J.D. (2005) Skeletal homologies, phylogeny and classification of the earliest asterozoan echinoderms. Journal of Systematic Palaeontology 3, 29–114.
Smirnov, I.S. (1977) New species of the genus Ophiomisidium (Ophiuroidea) from subantarctic water of Indian Ocean. New species and genus of marine invertebrates. Explorations of the fauna of the seas, 21, 105–108.
Smith, A.B., Paterson, G.L.J. & Lafay, B. (1995) Ophiuroid phylogeny and higher taxonomy: morphological, molecular and palaentological perspectives. Zoological Journal of the Linnean Society 114, 213–243.
Spencer, W.K. & Wright, C.W. (1966) Asterozoans. In: Moore R.C. (Ed) Treatise on Invertebrate Paleontology. Lawrence and the Geological Society of America, New York: University of Kansas Press, U4–107.
Stauber, M. & Märkel, K. (1988) Comparative morphology of muscle-skeleton attachments in the Echinodermata. Zoomorphology 108, 137–148.
Stewart, B. (2000) Anatomical features of the euryalid snake star Astrobrachion constrictum (Ophiuroidea: Asteroschematidae). Invertebrate Biology 119, 222–233.
Stöhr, S. (2003) A new fissiparous amphiurid brittlestar (Echinodermata: Ophiuroidea) from southwest of Iceland. Sarsia 88, 373–378.
Stöhr, S. (2005) Who’s who among baby brittle stars (Echinodermata: Ophiuroidea): postmetamorphic development of some North Atlantic forms. Zoological Journal of the Linnean Society 143, 543–576.
Stöhr, S., Conand, C. & Boissin E. (2008) Brittle stars (Echinodermata: Ophiuroidea) from La Réunion and the systematic position of Ophiocanops Koehler, 1922. Zoological Journal of the Linnean Society, 153, 545–560.
Stöhr, S. & O'Hara, T. (2003) Deep-sea ophiuroids of New Caledonia - a preliminary report. In: Féral, J.-P. & David, B. (Eds): Echinoderm Research 2001 - Proceedings of the 6th European Conference on Echinoderms, Banyuls-sur- Mer, France, 3–7 Sep 2001, 49–52. Lisse: Swets & Zeitlinger.
Stöhr, S. & Segonzac, M. (2005) Deep-sea ophiuroids (Echinodermata) from reducing and non-reducing environments in the North Atlantic Ocean. Journal of the Marine Biology Association of United Kingdom 85, 383–402.
Stöhr, S. & Segonzac, M. (2006a) Two new genera and species of ophiuroid (Echinodermata) from hydrothermal vents in the East Pacific. Species Diversity, 11, 7–32.
Stöhr, S. & Segonzac, M. (2006b) Ophiuroidea. In: Debruyres, D., Segonzac, M. & Bright M. (Eds), Handbook of Deep-sea Hydrothermal Vent Fauna. 2nd completely revised edition. Denisia 18, 481–485.
Studer, T. (1876) Über Echinodermen aus dem antarktischen Meere und zwei neue Seeigel von den Papua-Inseln, gesammelt auf der Reise S.M.S. Gazelle um die Erde. Monatsberichte der Königlichen Preussischen Akademie der Wissenschaften. Berlin, 452–465.
Studer, T. (1882) Übersicht über die Ophiuriden, welche während der Reise S.M.S. Gazelle um die Erde 1874–1876 gesammelt wurden. Abhandlungen der Preussischen Akademie der Wissenschaften zu Berlin 1, 1–37.
Sumida, P.Y.G., Tyler, P.A., Gage, J.D. & Norrevang, A. (1998) Postlarval development in shallow and deep-sea ophiuroids (Echinodermata: Ophiuroidea) of the NE Atlantic Ocean. Zoological Journal of the Linnean Society 124, 267–300.
Thomas, L.P. (1966) A revision of the tropical American species of Amphipholis (Echinodermata: Ophiuroidea). Bulletin of Marine Science 16, 827–833.
Thomas, L.P. (1967) The systematic position of Amphilimna (Echinodermata; Ophiuroidea). Proceedings of the Biological Society of Washington 80, 123–130.
Thomas, L.P. (1973) Western Atlantic brittle stars of the genus Ophionereis. Bulletin of Marine Science, 23, 585–599.
Thomas, L.P. (1975) The ophiacanthid genus Amphilimna (Ophiuroidea, Echinodermata). Proceedings of the Biological Society of Washington 88, 127–140.
Thomas, L.P. & Schoener, A. (1972) Growth changes in Amphilimna olivacea (Lyman) and the systematic status of Amphitarsus spinifer Schoener. Breviora, Museum of Comparative Zoology 387, 2–9.
Tommasi, L.R. (1967) Sobre dois Amphiuridae de fauna marinha do sul do Brazil. Contribuições Avulsas do Instituto Oceanográfico, Universidade de São Paulo, Série Oceanografia Biológica 12, 1–5.
Tommasi, L.R. (1968) Amphiuridae de la Bahia de Valparaiso. Revista de Biologia Marina, 13, 71–78.
Tommasi, L.R. (1970) Os ofiuróides recentes do Brasil e de regiões vizinhas. Contribuições Avulsas do Instituto Oceanográfico, Universidade de São Paulo, Série Oceanografia Biológica, 20, 1–146.
Tortonese, E. (1965) Fauna d'Italia. Echinodermata. Edizioni Calderini Bologna; Officine Grafiche Calderini. Bologna, Italy.
Turner, R.L. & Heyman, R.M. (1995) Rediagnosis of the brittlestar genus Ophiosyzygus and notes on its type species O. disacanthus (Echinodermata: Ophiuroidea: Ophiomyxidae) based on the type specimens from Japanese waters and new material from the Gulf of Mexico. Proceedings of the Biological Society of Washington 108, 292–297.
Tyler, P. A., Paterson, G. J. L., Sibuet, M., Guille, A., Murtons, B.J. and Segonzac, M. (1995) A new genus of ophiuroid (Echinodermata: Ophiuroidea) from hydrothermal mounds along the Mid-Atlantic Ridge. Journal of the Marine Biological Association of the United Kingdom, 75, 977–986.
Vadon, C. (1990) Ophiozonella novaecaledoniae n. sp. (Ophiuroidea: Echinodermata): description, ontogeny and phyletic position. Journal of Natural History, 24, 165–179.
Verrill, A.E. (1869) On new and imperfectly known echinoderms and corals. Proceedings of the Boston Society of Natural History, 12, 381–391.
Verrill, A.E. (1899) Report on the Ophiuroidea; collected by the Bahama Expedition from the University of Iowa in 1893. Bulletin from the Laboratories of Natural History of the State University of Iowa 5, 1–86.
Webb, P.M. & Tyler, P.A. (1985). Post-larval development of the common north-west European brittle stars Ophiura ophiura, O. albida and Acrocnida brachiata (Echinodermata: Ophiuroidea). Marine Biology 89, 281–292.
Wilkie, I.C. (1980) The systematic position of Ophiocomina Koehler and a reconsideration of certain interfamilial relationship within the Ophiuroidea. In: Jangoux M (ed). Proceedings of the European Colloquium in Echinoderms, Brussels, 3–8 September 1979, Rotterdam: Balkema, 151–157.
Ziesenhenne, F.C. (1955) A review of the genus Ophioderma M. & T. Essays in the natural sciences in honour of Captain Alan Hancock, 185–201.