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Deep-sea ophiuroids from western Mexico have been documented since 1899, but mostly by non-Mexican expeditions. TALUD is a Mexican project designed to study the deep-sea fauna from Mexico. As part of it, the present contribution provides for the first time detailed information about the taxonomy and distribution, as well as images, of deep-sea ophiuroids from western Mexico, representing a useful taxonomic identification tool for Ophiuroidea from the eastern Pacific. A total of 38 species of ophiuroid (35 identified at species level, one at genus level, and two as confer) were collected from 83 stations located at 123‒2,309 m depth. At a regional scale, several new species records are presented: six for western Baja California, five for western Baja California Sur, two for the Gulf of California, four for Jalisco, 14 for Colima, and five for Guerrero. Geographic (six) and bathymetric (seven) distribution ranges of species are extended. An updated list of deep-sea ophiuroids (61 species) from western Mexico is provided together with an identification key to species collected during this survey. In addition, nine records of deep-sea ophiuroids for the Mexican Pacific are unproven, doubtful or invalid. We corroborated the presence of Ophiacantha eurypoma and Ophiacantha pacifica in the study area and added Amphiura gymnogastra as the first record for Mexico. We propose to transfer Ophiacantha phragma to Ophiolimna on account of it having both granules and spines on the dorsal disc and striated arm plates. TALUD cruises collected 57 % of the total deep-sea species known to occur off Mexico, making it the most important survey of deep-sea Ophiuroidea carried out in western Mexico to date.

Amphiura carchara H.L. Clark, 1911 Fig. 14G‒L Amphiura carchara H.L. Clark, 1911: 142‒143, fig. 55.— Martynov 2010: figs. 15m, 31f. Pandelia carchara. Fell 1962: 10.— Kyte 1969a: 1737. Material examined. 31 individuals at three stations. TALUD XVIB, Sta. 22, 1 ind. (ICML-EMU-11149-A) and 4 ind. (ICML-EMU-11149-B); Sta. 26, 1 ind. (ICML-EMU-11149-C); Sta. 27, 10 ind. (ICML-EMU-11150) and 15 ind. (ICML-EMU-11667). Comparative material. Holotype: USNM 25594. Paratypes, 158 ind.: MCZ OPH-3140, MCZ OPH-3141, MCZ OPH-3370, USNM 25993, USNM 26053, USNM 26114, USNM 26115, USNM 26213, USNM 26615, USNM 27017, USNM 27018, USNM 27054, USNM 32377, USNM 33639 (Supplementary file 2). Description (ICML-EMU-11667). DD = 7 mm. Disc rounded. Dorsal disc covered by imbricated scales. Primary plates oval, separated from each other by scales. RS triangular with rounded edges, in contact only distally, the rest separated by scales (Fig. 14G). Ventral interradii covered by integument, distally with a fold of scales (Fig. 14H). OSh broader than long, bell-shaped with rounded edges, a distal lobe. Madreporite not evident. AdSh triangular, wider distally, meeting in front of OSh or separated. Jaws bearing three oral papillae at each side; AdShSp elongated, pointed; BSc elongated, pointed; IPa quadrangular with rounded edges, strong. vT elongated with straight edge (Fig. 14I). Arms slender. DAP broader than long, fan-shaped with rounded edges, contiguous (Fig. 14J). VAP longer than broad, pentagonal, slightly separated from each other. LAP with three ArSp, small (approximately 1.5 arm segment in length), pointed, middle ArSp stronger than rest. Tentacle pores with one oval TSc (Fig. 14K). Color pattern dorsally and ventrally beige-whitish (ethanol preservation) (Fig. 14G‒L). Habitat and distribution. Bering Sea, Gulf of Alaska, Washington, USA, and northwestern Mexico (H.L. Clark 1911; Granja-Fernández et al. 2015); 568‒ 3,611 m depth, sandy and muddy substrates (H.L. Clark 1911; Maluf 1988). The material examined was collected in northwestern Baja California; 982‒ 1,560 m depth. Remarks. Examination of the types and the material available herein showed that the morphology in Amphiura carchara is very constant. The only variation observed during this study was the presence of round and separated primary plates in specimens with a DD 6 mm., Published as part of Granja-Fernández, Rebeca, Hendrickx, Michel E., Rangel-Solís, Pedro Diego & López-Pérez, Andrés, 2023, Deep-sea Ophiuroidea (Echinodermata) collected during the TALUD cruises in western Mexico, pp. 1-71 in Zootaxa 5259 (1) on pages 48-50, DOI: 10.11646/zootaxa.5259.1.1, http://zenodo.org/record/7794984, {"references":["Clark, H. L. (1911) North Pacific Ophiurans in the collection of the United States National Museum. U. S. National Museum Bulletin, 75, 1 - 302. https: // doi. org / 10.5479 / si. 03629236.75.1","Martynov, A. (2010) Reassessment of the classification of the Ophiuroidea (Echinodermata), based on morphological characters. I. General character evaluation and delineation of the families Ophiomyxidae and Ophiacanthidae. Zootaxa, 2697 (1), 1 - 154. https: // doi. org / 10.11646 / zootaxa. 2697.1.1","Fell, B. H. (1962) A revision of the major genera of Amphiuroid Ophiuroidea. Transactions of the Royal Society of New Zealand, Zoology, 2 (2), 1 - 26.","Kyte, M. A. (1969 a) A synopsis and key to the recent Ophiuroidea of Washington State and Southern British Columbia. Journal of the Fisheries Research Board of Canada, 26 (7), 1727 - 1741. https: // doi. org / 10.1139 / f 69 - 160","Granja-Fernandez, R., Herrero-Perezrul, M. D., Lopez-Perez, R. A., Hernandez-Morales, A. & Rangel-Solis, P. D. (2015) A literature review of the Ophiuroidea (Echinodermata) from the Pacific coast of Mexico. Revista de Biologia Tropical, 63 (2), 37 - 47.","Maluf, L. Y. (1988) Composition and distribution of the central Eastern Pacific Echinoderms. Natural History Museum of Los Angeles County, Technical Report, 2, 1 - 306."]}

Ophiocnida californica Ziesenhenne, 1940 Fig. 17G‒L Ophiocnida californica Ziesenhenne, 1940: 25‒27, pl. 5, figs. 4‒6.— Granja-Fernández et al. 2014: 109. Material examined. Seven individuals at one station. TALUD XIV, Sta. 32, 3 ind. (ICML-EMU-11174), 3 ind. (ICML-EMU-11175), and 1 ind. (ICML-EMU-11678). Comparative material. Holotype: LACM 615.1. Paratypes, 32 ind.: ICML-UNAM 3.76.0, LACM 615.2, LACM 615.3, LACM 615.4, LACM 615.5, MCZ OPH-7032, USNM E8007 (Supplementary file 2). Description (ICML-EMU-11175). DD = 6 mm. Disc circular. Dorsal disc covered by imbricated scales and scattered small spines mostly around the margin of the disc. Primary plates rounded, widely separated from each other. RS almost bar-like, separated by elongated scales for half their length (Fig. 17G). Ventral interradii covered by imbricated scales smaller than dorsal disc (Fig. 17H). OSh broader than long, spearhead-shaped. Madreporite slightly larger than OSh. AdSh trapezoidal, meeting in front of OSh. Jaws bearing three oral papillae at each side; AdShSp slightly serrated, rectangular, the largest; 2AdShSp pointed; IPa quadrangular-rounded. vT quadrangular (Fig. 17I). Arms slender. DAP twice as wide as long, triangular, some fragmented in 2‒3 pieces, contiguous (Fig. 17J). VAP slightly broader than long, pentagonal, contiguous. LAP with up to three arm spines, small (approximately one arm segment in length), blunt-tip, middle the longest, dorsalmost the shortest. Tentacle pores with two oval TSc (Fig. 17K). Color pattern dorsally and ventrally light brown (ethanol preservation) (Fig. 17G‒L). Habitat and distribution. Gulf of California, Mexico; 5‒302 m depth, coral, algae, sandy, muddy, and rocky substrates (Ziesenhenne 1940; pers. obs.). The material examined was collected only in one locality in the Gulf of California; 123 m depth. Remarks. In some specimens of O. californica collected during this survey, the central primary plate is always evident, contrary to the radial primary plates, which are not always conspicuous. In addition, the fragmentation of the dorsal arm plates is not always very evident, the serration of the AdShSp is sometimes conspicuous and sometimes not. Finally, the ventral arm plates are usually triangular. Other Ophiocnida species have scattered spines on the dorsal disc as in O. californica; however, they differ in having all oral papillae rounded and small (vs. AdShSp slightly serrated and rectangular in O. californica), non-fragmented dorsal arm plates (vs. fragmented), and tentacle scales arranged perpendicularly (vs. arranged in parallel) (e.g., Gondim et al. 2013; Granja-Fernández et al. 2014). Due to these differences, a revision of the generic placement of O. californica is needed., Published as part of Granja-Fernández, Rebeca, Hendrickx, Michel E., Rangel-Solís, Pedro Diego & López-Pérez, Andrés, 2023, Deep-sea Ophiuroidea (Echinodermata) collected during the TALUD cruises in western Mexico, pp. 1-71 in Zootaxa 5259 (1) on page 57, DOI: 10.11646/zootaxa.5259.1.1, http://zenodo.org/record/7794984, {"references":["Ziesenhenne, F. C. (1940) New Ophiurans of the Allan Hancock Pacific Expeditions. Allan Hancock Pacific Expeditions, 8 (2), 9 - 52.","Granja-Fernandez, R., Herrero-Perezrul, M. D., Lopez-Perez, R. A., Hernandez, L., Rodriguez-Zaragoza, F. A., Jones, R. W. & Pineda-Lopez, R. (2014) Ophiuroidea (Echinodermata) from coral reefs in the Mexican Pacific. ZooKeys, 406, 101 - 145. https: // doi. org / 10.3897 / zookeys. 406.6306","Gondim, A. I., Alonso, C., Dias, T. L. P., Manso, C. L. C. & Christoffersen, M. L. (2013) A taxonomic guide to the brittle-stars (Echinodermata, Ophiuroidea) from the State of Paraiba continental shelf, Northeastern Brazil. ZooKeys, 307, 45 - 96. https: // doi. org / 10.3897 / zookeys. 307.4673"]}

Amphipholis pugetana (Lyman, 1860) Fig. 16G‒L Amphiura pugetana Lyman, 1860: 193‒194. Amphipholis pugetana. H.L. Clark 1911: 166‒167.— Nielsen 1932: 281, 288‒290.— Fell 1962: 13, 24.— Lambert & Austin 2007: 102‒104, figs. 55‒56.— Boolotian & Leighton 1966: 5, 9, fig. 9.— Hendler 1996: 140‒142, fig. 7.9. Axiognatus pugetana. Kyte 1969a: 1728, 1735, 1737. See Hendler (1996) for other synonymous records. Material examined. 132 individuals at five stations. TALUD X, Sta. 4, 1 ind. (ICML-EMU-11162-A) and 16 ind. (ICML-EMU-11162-B). TALUD XIV, Sta. 15, 57 ind. (ICML-EMU-11163), 5 ind. (ICML-EMU-11164), and 28 ind. (ICML-EMU-11164-A); Sta. 20, 5 ind. (ICML-EMU-11164-B). TALUD XV, Sta. 20, 1 ind. (ICML-EMU-11165) and 2 ind. (ICML-EMU-12994); Sta. 23, 17 ind. (ICML-EMU-11166). Comparative material. None available. According to Lyman (1860), the type series of A. pugetana was deposited at the USNM, but this material has not been located in this collection or in any other collection (i.e., LACM, MCZ, NHMD; pers. comm. Tom Schiøtte & Penny Benson, 2016) and is probably lost. Description (ICML-EMU-11163). DD = 4 mm. Disc pentagonal. Dorsal disc covered by imbricated scales. Primary plates not evident. RS elongated half-circular, contiguous but with 1‒2 scales inserted proximally (Fig. 16G). Ventral interradii covered by scales smaller than dorsal disc (Fig. 16H). OSh as broad as long, spearhead-shaped. Madreporite slightly larger than OSh. AdSh triangular, meeting in front of OSh. Jaws bearing three oral papillae at each side; AdShSp twice as wide as high, rounded rectangular, the largest; 2AdShSp rounded; IPa quadrangular. vT quadrangular (Fig. 16I). Arms slender. DAP fan-shaped, as long as wide, separated (Fig. 16J). VAP longer than broad, pentagonal, slightly separated from each other. LAP with up to three ArSp, elongated (approximately two arm segments in length), pointed, middle one often much longer and swollen at the tip. Tentacle pores with two TSc, abradial larger (Fig. 16K). Color pattern dorsally and ventrally beige-whitish (ethanol preservation) (Fig. 16G‒L). Habitat and distribution. Alaska, British Columbia, Canada, Washington, and California, USA, and Mexico; 15‒ 871 m depth, possibly to 1,620 m (H.L. Clark 1911; Hendler 1996; Lambert & Austin 2007; Granja-Fernández et al. 2015, 2021; See Remarks), buried in sand and mud (Maluf 1988). The material examined was collected off western Baja California Sur and in the northern Gulf of California; 410‒871 m depth. Some specimens were attached to a log. Remarks. We could not compare our material with the type material (presumably lost), but its morphology is consistent with previous species descriptions (e.g., Lyman 1860; Hendler 1996). Previously recorded from shallow water and as deep as 604 m, and possibly to 1,620 m (Hendler 1996). Our finding of the species at 871 m depth corresponds to the deepest confirmed record to date., Published as part of Granja-Fernández, Rebeca, Hendrickx, Michel E., Rangel-Solís, Pedro Diego & López-Pérez, Andrés, 2023, Deep-sea Ophiuroidea (Echinodermata) collected during the TALUD cruises in western Mexico, pp. 1-71 in Zootaxa 5259 (1) on page 52, DOI: 10.11646/zootaxa.5259.1.1, http://zenodo.org/record/7794984, {"references":["Lyman, T. (1860) Descriptions of New Ophiuridae, belonging to the Smithsonian Institution and to the Museum of Comparative Zoology at Cambridge. Proceedings of the Boston Society of Natural History, 7, 193 - 204 + 252 - 262. https: // doi. org / 10.5962 / bhl. part. 4822","Clark, H. L. (1911) North Pacific Ophiurans in the collection of the United States National Museum. U. S. National Museum Bulletin, 75, 1 - 302. https: // doi. org / 10.5479 / si. 03629236.75.1","Nielsen, E. (1932) Ophiurans from the Gulf of Panama, California and the Strait of Georgia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 91, 241 - 346.","Fell, B. H. (1962) A revision of the major genera of Amphiuroid Ophiuroidea. Transactions of the Royal Society of New Zealand, Zoology, 2 (2), 1 - 26.","Lambert, P. & Austin, W. (2007) Brittle stars, sea urchins and feather stars of British Columbia, Southeast Alaska and Puget Sound. Royal BC Museum Handbook, Victoria, 150 pp.","Boolotian, R. A. & Leighton, D. (1966) A key to the species of Ophiuroidea (brittle stars) of the Santa Monica Bay and adjacent areas. Los Angeles County Museum Contributions in Science, 93, 1 - 20. https: // doi. org / 10.5962 / p. 241083","Hendler, G. (1996) Taxonomic atlas of the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel. Class Ophiuroidea. Santa Barbara Museum of Natural History, 14, 113 - 179.","Kyte, M. A. (1969 a) A synopsis and key to the recent Ophiuroidea of Washington State and Southern British Columbia. Journal of the Fisheries Research Board of Canada, 26 (7), 1727 - 1741. https: // doi. org / 10.1139 / f 69 - 160","Granja-Fernandez, R., Herrero-Perezrul, M. D., Lopez-Perez, R. A., Hernandez-Morales, A. & Rangel-Solis, P. D. (2015) A literature review of the Ophiuroidea (Echinodermata) from the Pacific coast of Mexico. Revista de Biologia Tropical, 63 (2), 37 - 47.","Granja-Fernandez, R., Maya-Alvarado, B., Cupul-Magana, A. L., Rodriguez-Troncoso, A. P., Solis-Marin, F. A. & Sotelo-Casas, R. S. (2021) Echinoderms (Echinodermata) from the Central Mexican Pacific. Revista de Biologia Tropical, 69 (S 1), 219 - 253. https: // doi. org / 10.15517 / rbt. v 69 iSuppl. 1.46356","Maluf, L. Y. (1988) Composition and distribution of the central Eastern Pacific Echinoderms. Natural History Museum of Los Angeles County, Technical Report, 2, 1 - 306."]}

Amphiura (Amphiura) diomedeae Lütken & Mortensen, 1899 Fig. 15G‒L Amphiura diomedeae Lütken & Mortensen, 1899: 151‒152, pl. 12, figs. 1‒7.— H.L. Clark 1911: 140‒141; 1917: 436‒437.— Koehler 1922: 158‒159, pl. 67, figs. 7‒10.— Hendler 1996: 147‒149, fig. 7.12.— Lambert & Boutillier 2011: 53. Amphiura (Amphiura) diomedeae. Cherbonnier & Guille 1978: 35‒36, fig. 10. See Hendler (1996) for other synonymous records. Material examined. 40 individuals at 10 stations. TALUD IV, Sta. 25, 4 ind. (ICML-EMU-11151-A). TALUD VIII, Sta. 15, 1 ind. (ICML-EMU-11151-B). TALUD XII, Sta. 13, 4 ind. (ICML-EMU-11152); Sta. 25, 1 ind. (ICML-EMU-11696); Sta. 26, 3 ind. (ICML-EMU-11157). TALUD XIV, Sta. 15, 1 ind. (ICML-EMU-11665). TALUD XVIB, Sta. 1, 1 ind. (ICML-EMU-11153); Sta. 5, 1 ind. (ICML-EMU-11156-A); Sta. 22, 6 ind. (ICML-EMU-11154) and 3 ind. (ICML-EMU-11156-B); Sta. 27, 15 ind. (ICML-EMU-11155). Comparative material. Syntypes, 9 ind.: MCZ OPH-1305, MCZ OPH-1379 (Supplementary file 2). Description (ICML-EMU-11155). DD = 12 mm. Disc interradially incised. Dorsal disc covered by imbricated scales, center scales larger. Primary plates oval, separated from each other by scales. RS triangular, in contact or slightly in contact only distally, the rest separated by scales (Fig. 15G). Ventral interradii covered by imbricated scales, smaller than dorsal scales (Fig. 15H). OSh broader than long, diamond-shaped with rounded edges. Madreporite larger than OSh, with about five hydropores. AdSh scalene triangular, separated. Jaws bearing four oral papillae at each side; AdShSp and 2AdShSp rounded, short; BSc pointed, short; IPa rounded; a gap between BSc and IPa. vT quadrangular (Fig. 15I). Arms slender. DAP broader than long, widely fan-shaped, contiguous (Fig. 15J). VAP almost as broad as long, pentagonal, slightly separated from each other. LAP with up to three ArSp, elongated (almost two arm segments in length), pointed, middle the longest, dorsalmost the shortest and most slender. Tentacle pores with two oval TSc (Fig. 15K). Color pattern dorsally and ventrally light brown (ethanol preservation) (Fig. 15G‒L). Habitat and distribution. Sri Lanka, Maldives, Philippines Islands, Japan, Tahiti, USA, Mexico, Costa Rica, Panama, Ecuador, and Peru (Lütken & Mortensen 1899; H.L. Clark 1911, 1917; Koehler 1922; Hendler 1996; Stöhr & O’Hara 2021); 71‒ 3,310 m depth (Hendler 1996; Stöhr & O’Hara 2021), sandy and rocky substrates (Lütken & Mortensen 1899). The material examined was collected from off western Baja California, in the Gulf of California, and off Colima and Guerrero; 772‒ 2,054 m depth. Remarks. Despite its wide geographical distribution, Amphiura (Amphiura) diomedae had previously been recorded in Mexico only in the Gulf of California (Granja-Fernández et al. 2015). The new records off western Baja California, Colima, and Guerrero indicates a much wider distribution range in western Mexico than previously thought., Published as part of Granja-Fernández, Rebeca, Hendrickx, Michel E., Rangel-Solís, Pedro Diego & López-Pérez, Andrés, 2023, Deep-sea Ophiuroidea (Echinodermata) collected during the TALUD cruises in western Mexico, pp. 1-71 in Zootaxa 5259 (1) on pages 50-51, DOI: 10.11646/zootaxa.5259.1.1, http://zenodo.org/record/7794984, {"references":["Lu ¨ tken, C. F. &. Mortensen, T. (1899) The Ophiuridae. Memoirs of the Museum of Comparative Zoology at Harvard College, 23 (2), 93 - 208.","Clark, H. L. (1911) North Pacific Ophiurans in the collection of the United States National Museum. U. S. National Museum Bulletin, 75, 1 - 302. https: // doi. org / 10.5479 / si. 03629236.75.1","Clark, H. L. (1917) Reports on the scientific results of the expedition to the Eastern Tropical Pacific in charge of Alexander Agassiz, by the U. S. Fish Commission Steamer \" Albatross \" from October, 1904, to March, 1905, Lieut. Commander L. M. Garrett U. S. N., Commanding. XXX. Ophiuroidea. Bulletin of the Museum of Comparative Zoology at Harvard College, 61 (12), 429 - 453.","Koehler, R. (1922) Ophiurans of the Philippine Seas and adjacent waters. Bulletin of the United States National Museum, 100, 1 - 486. https: // doi. org / 10.5962 / bhl. title. 32917","Hendler, G. (1996) Taxonomic atlas of the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel. Class Ophiuroidea. Santa Barbara Museum of Natural History, 14, 113 - 179.","Lambert, P. & Boutillier, J. (2011) Deep-sea Echinodermata of British Columbia, Canada. Canadian Technical Report of Fisheries and Aquatic Sciences, 2929, 1 - 140.","Cherbonnier G. & Guille, A. (1978) Faune de Madagascar: Ophiurides. Vol. 48. Museum national d'Histoire naturelle, Paris, 272 pp.","Stohr, S. & O'Hara, T. D. (2021) Deep-sea Ophiuroidea (Echinodermata) from the Danish Galathea II Expedition, 1950 - 52, with taxonomic revisions. Zootaxa, 4963 (3), 505 - 529. https: // doi. org / 10.11646 / zootaxa. 4963.3.6","Granja-Fernandez, R., Herrero-Perezrul, M. D., Lopez-Perez, R. A., Hernandez-Morales, A. & Rangel-Solis, P. D. (2015) A literature review of the Ophiuroidea (Echinodermata) from the Pacific coast of Mexico. Revista de Biologia Tropical, 63 (2), 37 - 47."]}

Amphiura gymnogastra Lütken & Mortensen, 1899 Fig. 15A‒F Amphiura gymnogastra Lütken & Mortensen, 1899: 145‒146, pl. 9, figs. 6‒8.— H.L. Clark 1917: 437. Hemilepis gymnogastra. Fell 1962: 10, 22. Material examined. Four individuals at two stations. TALUD XII, Sta. 13, 3 ind. (ICML-EMU-11148-A); Sta. 26, 1 ind. (ICML-EMU-11148-B). Comparative material. Syntypes, 5 ind.: MCZ OPH-1321, USNM 19526 (Supplementary file 2). Description (ICML-EMU-11148-A). DD = 6.6 mm. Disc malformed. Dorsal disc covered by imbricated scales, those surrounding RS larger, integument distally. Primary plates not evident. RS triangular with rounded edges, prominent distal end, in contact only distally, the rest separated by scales (Fig. 15A). Ventral interradii covered by scale-less integument (Fig. 15B). OSh broader than long, spearhead-shaped with strongly convex proximal and lateral edges and distal lobe. Madreporite rounder and larger than OSh. AdSh scalene triangular, separated. Jaws bearing three oral papillae at each side; AdShSp scale-like round; BSc slender, pointed; IPa quadrangular with rounded edges. vT quadrangular (Fig. 15C). Arms slender. DAP broader than long, oval, contiguous (Fig. 15D). VAP longer than broad, pentagonal, contiguous. LAP with 5‒6 ArSp, small (approximately 1.2 arm segment in length), conical with blunt tip; ventralmost the longest, dorsalmost the thinnest and shortest. Tentacle pores with two oval TSc, adradial longer (Fig. 15E). Color pattern in dorsally and ventrally beige-whitish (ethanol preservation) (Fig. 15A‒F). Habitat and distribution. Mexico, Panama, and the Galapagos Islands; 549‒ 2,323 m depth, hard and muddy substrates (Lütken & Mortensen 1899; H.L. Clark 1911; Maluf 1988; see Remarks). The material examined was collected off Colima and Guerrero; 1,199 ‒1,956 m depth. Remarks. Specimens of Amphiura gymnogastra tend to lose their disc and oral papillae very easily, thus making correct identification very complicated. This species can have a dorsal disc covered with imbricated scales or with scales and small areas near the margin of the disc with only integument.Areas with integument in the dorsal disc can result from the pressure by the ventral disc on the dorsal side during the sampling process. Amphiura gymnogastra has been previously recorded off Panama and the Galapagos Islands (Lütken & Mortensen 1899; H.L. Clark 1911), and records off central and southern Mexico represent a significant extension of its northernmost distribution limit to 18º33′27″N, 104º28′21″W. Available material of A. gymnogastra is currently limited to 13 specimens: 1) the type material, which is not in good condition (MCZ OPH-1321, two specimens; USNM 19526, three specimens), 2) one lot from the Galapagos Islands (USNM E669, four specimens), and 3) two lots from the TALUD cruises (ICML -EMU-11148-A, three specimens; ICML-EMU-11148-B, one specimen).

Dougaloplus sp. Fig. 17A‒F Material examined. 24 individuals at eight stations. TALUD IV, Sta. 19, 1 ind. (ICML-EMU-11158-A); Sta. 25, 1 ind. (ICML-EMU-11158-B). TALUD VII, Sta. 19, 9 ind. (ICML-EMU-9014-A) and 1 ind. (ICML-EMU-9014-B). TALUD VIII, Sta. 3, 1 ind. (ICML-EMU-11159-A); Sta. 20, 4 ind. (ICML-EMU-11159-B) and 4 ind. (ICML-EMU-11670). TALUD IX, Sta. 4, 1 ind. (ICML-EMU-11160-A); Sta. 21B, 1 ind. (ICML-EMU-11160-B). TALUD XII, Sta. 13, 1 ind. (ICML-EMU-11161). Comparative material. Amphilimna pentacantha H.L. Clark, 1911 (junior synonym of Dougaloplus amphacanthus (McClendon, 1909)), holotype: USNM 25614. Paratypes, 10 ind.: MCZ OPH- 3172, MCZ OPH- 3384. Amphioplus hexacanthus H.L. Clark, 1911 (junior synonym of Dougaloplus amphacanthus (McClendon, 1909)), syntype, 1 ind.: MCZ OPH- 3169. Amphiura notacantha Lütken & Mortensen, 1899 (original name of Dougaloplus notacanthus (Lütken & Mortensen, 1899), holotype: USNM 19543 (Supplementary file 2). Description (ICML-EMU-9014-A). DD = 7 mm. Disc malformed. Dorsal disc covered by imbricated scales, largest in the center, and by pointed spines on interradius and RS. Primary plates not evident. RS bar-like, elongated and narrow, meeting distally, separated by scales except distally (Fig. 17A). Ventral interradii covered by scales smaller than dorsal disc (Fig. 17B). OSh broader than long, rounded pentagonal, distal edge concave. Madreporite slightly larger than OSh. AdSh scalene triangular, separated. Jaws bearing four oral papillae at each side; AdShSp rounded, the shortest; 2AdShSp pointed; BSc pointed; IPa quadrangular-rounded. vT quadrangular (Fig. 17C). Arms slender. DAP broader than long, widely oval, separated from each other (Fig. 17D). VAP longer than broad, pentagonal, slightly separated from each other. Up to five ArSp, small (approximately one arm segment in length), pointed, ventralmost the longest, dorsalmost the shortest. Tentacle pores with one round TSc (Fig. 17E). Color pattern dorsally and ventrally beige-whitish (ethanol preservation) (Fig. 17A‒F). Habitat and distribution. The material examined was collected in the southern Gulf of California and off Guerrero; 778‒ 1,395 m depth. Remarks. The genus Dougaloplus is represented in eastern tropical Pacific by three species: Dougaloplus amphacanthus (McClendon, 1909), Dougaloplus gastracanthus (Lütken & Mortensen, 1899), and Dougaloplus notacanthus (Lütken & Mortensen, 1899). Compared with the material of Dougaloplus sp. examined, D. amphacanthus is characterized by a dorsal disc densely covered with spines, two tentacle scales, and a bifurcated tip on the arm spines. Dougaloplus sp. is close to D. gastracanthus and D. notacanthus (which are very similar in appearance), but these two species differ from our material in having less spines on the dorsal disc, and the elongated and pointed oral papillae. Information and material of D. gastracanthus and D. notacanthus are restricted to the original description of D. gastracanthus (Lütken & Mortensen 1899) and the damaged holotype of D. notacanthus; additional, non-type material of these two species is not available. More specimens of these species are needed to elucidate whether Dougaloplus sp. belongs to either D. gastracanthus or D. notacanthus, or if it belongs to a new species., Published as part of Granja-Fernández, Rebeca, Hendrickx, Michel E., Rangel-Solís, Pedro Diego & López-Pérez, Andrés, 2023, Deep-sea Ophiuroidea (Echinodermata) collected during the TALUD cruises in western Mexico, pp. 1-71 in Zootaxa 5259 (1) on page 55, DOI: 10.11646/zootaxa.5259.1.1, http://zenodo.org/record/7794984, {"references":["Clark, H. L. (1911) North Pacific Ophiurans in the collection of the United States National Museum. U. S. National Museum Bulletin, 75, 1 - 302. https: // doi. org / 10.5479 / si. 03629236.75.1","McClendon, J. F. (1909) The ophiurans of the San Diego region. University of California Publications in Zoology, 6 (3), 33 - 64.","Lu ¨ tken, C. F. &. Mortensen, T. (1899) The Ophiuridae. Memoirs of the Museum of Comparative Zoology at Harvard College, 23 (2), 93 - 208."]}

Genus Dougaloplus A.M. Clark, 1970 Species of Dougaloplus reported in the eastern Pacific were originally described in the genera Amphiura and Ophiocnida (Lütken & Mortensen 1899; McClendon 1909) and subsequently transferred to Amphiacantha, a preoccupied genus, by Matsumoto (1917). Consequently, A.M. Clark (1970) proposed the replacement name Dougaloplus for the genus. Some authors have described a few characteristics of Dougaloplus (e.g., disc covered with scales and spines, 1‒2 tentacle scales, 3‒6 arm spines; Matsumoto 1917; A.M. Clark 1970); however, a proper diagnosis of the genus is lacking so far., Published as part of Granja-Fernández, Rebeca, Hendrickx, Michel E., Rangel-Solís, Pedro Diego & López-Pérez, Andrés, 2023, Deep-sea Ophiuroidea (Echinodermata) collected during the TALUD cruises in western Mexico, pp. 1-71 in Zootaxa 5259 (1) on page 55, DOI: 10.11646/zootaxa.5259.1.1, http://zenodo.org/record/7794984, {"references":["Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History) Zoology, 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085","Lu ¨ tken, C. F. &. Mortensen, T. (1899) The Ophiuridae. Memoirs of the Museum of Comparative Zoology at Harvard College, 23 (2), 93 - 208.","McClendon, J. F. (1909) The ophiurans of the San Diego region. University of California Publications in Zoology, 6 (3), 33 - 64.","Matsumoto, H. (1917) A monograph of Japanese Ophiuroidea, arranged according to a new classification. Journal of the College of Science, Imperial University, Tokyo, 32 (2), 1 - 408."]}

Amphiura arcystata H.L. Clark, 1911 Fig. 14A‒F Amphiura acrystata H.L. Clark, 1911: 145‒148, fig. 58 (lapsus calamus for arcystata) Amphiura arcystata. H.L. Clark 1915: 224; 1940: 336; 1970: 10‒11.— Nielsen 1932: 264.— Ziesenhenne 1937: 223.— Boolotian & Leighton 1966: 4, 8, fig. 7.— Hendler 1996: 145‒147. Hemilepis arcystata. Fell 1962: 10, 19. See Hendler (1996) for other synonymous records. Material examined. One individual at one station. TALUD XV, Sta. 23, 1 ind. (ICML-EMU-12995). Comparative material. Holotype: USNM 25623. Paratypes, 53 ind.: MCZ OPH-3135, MCZ OPH-3136, MCZ OPH-3137, MCZ OPH-3138, MCZ OPH-3368, MCZ OPH-3369, USNM 25834, USNM 25837, USNM 26078, USNM 26130, USNM 26188, USNM 26197, USNM 26609, USNM 27094, USNM 27124, USNM 33638 (Supplementary file 2). Description (ICML-EMU-12995). DD = 5.3 mm. Disc strongly shrinked due to preservation. Dorsal disc covered mostly by integument, imbricated scales around RS. Primary plates not evident. RS very conspicuous, thicker distally, in contact only distally, the rest separated by scales (Fig. 14A). Ventral interradii covered by scaleless integument (Fig. 14B). OSh broader than long, oval, with distal lobe. Madreporite larger than OSh. AdSh triangular, separated interradially and radially. Jaws bearing three oral papillae at each side; AdShSp elongated when present; BSc elongated, pointed; IPa rounded. vT quadrangular (Fig. 14C). Arms slender, elongated. DAP broader than long, oval, meeting (Fig. 14D). VAP a little broader than long, almost quadrangular, meeting. LAP with five ArSp, small (approximately one arm segment in length), pointed, ventralmost the longest and dorsalmost the shortest. Tentacle pores with two oval TSc (Fig. 14E). Color pattern in dorsal and ventral disc and arms beigewhitish (ethanol preservation) (Fig. 14A‒F). Habitat and distribution. Japan, California, USA, Mexico, Costa Rica, Colombia, and the Galapagos Islands (H.L. Clark 1911; Maluf & Brusca 2005); 60‒848 m depth, muddy and sandy substrates (H.L. Clark 1911; Ziesenhenne 1937; Hendler 1996). The material examined was collected off western Baja California Sur; 625 m depth. Remarks. During the TALUD cruises, only one specimen of Amphiura arcystata was collected and it is smaller than the holotype (USNM 25623, DD = 10 mm). The specimen differs in the number of arm spines (5 vs. 6‒7 in the holotype) and the thickness of the radial shields (thicker in the smaller specimen). Juvenile or small individuals of A. arcystata are known to significantly differ morphologically from adults, and it has been recorded that this species shows a great morphological variability regardless of size (H.L. Clark 1911; Hendler 1996). In this respect, the revision of the type material evidenced two morphologies (one from Japan ‒USNM 26130, MCZ OPH-3137 and the other from California‒USNM 25623, MCZ OPH-3135, MCZ OPH-3369) featuring differences in the presenceabsence of scales and integument in dorsal disc, the shape of the radial shields, oral shields, dorsal arm plates, ventral arm plates, and arm spines. The unique specimen collected during this survey agrees with the holotype from California. Japan and California morphologies probably correspond to different species, but more morphological and molecular evidence is needed. Due to the significant morphological variability observed here, we recommend reviewing the material of A. arcystata from Central and South America (Costa Rica, Colombia, and the Galapagos Islands). In Mexico, it has previously been collected off western Baja California, in the Gulf of California, and off Guerrero (Granja-Fernández et al. 2015). The current record is a first for the west coast of Baja California Sur., Published as part of Granja-Fernández, Rebeca, Hendrickx, Michel E., Rangel-Solís, Pedro Diego & López-Pérez, Andrés, 2023, Deep-sea Ophiuroidea (Echinodermata) collected during the TALUD cruises in western Mexico, pp. 1-71 in Zootaxa 5259 (1) on pages 47-48, DOI: 10.11646/zootaxa.5259.1.1, http://zenodo.org/record/7794984, {"references":["Clark, H. L. (1911) North Pacific Ophiurans in the collection of the United States National Museum. U. S. National Museum Bulletin, 75, 1 - 302. https: // doi. org / 10.5479 / si. 03629236.75.1","Clark, H. L. (1915) Catalogue of recent ophiurans. Memoirs of the Museum of Comparative Zoology, 25 (4), 163 - 376.","Clark, H. L. (1940) Eastern Pacific Expeditions of the New York Zoological Society. XXI. Notes on Echinoderms from the West Coast of Central America. Zoologica, 25 (22), 331 - 352. https: // doi. org / 10.5962 / p. 184708","Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History) Zoology, 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085","Nielsen, E. (1932) Ophiurans from the Gulf of Panama, California and the Strait of Georgia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 91, 241 - 346.","Ziesenhenne, F. C. (1937) The Templeton Crocker Expedition. X. Echinoderms from the West Coast of Lower California, the Gulf of California and Clarion Island. Zoologica, 22 (15), 209 - 239. https: // doi. org / 10.5962 / p. 184686","Boolotian, R. A. & Leighton, D. (1966) A key to the species of Ophiuroidea (brittle stars) of the Santa Monica Bay and adjacent areas. Los Angeles County Museum Contributions in Science, 93, 1 - 20. https: // doi. org / 10.5962 / p. 241083","Hendler, G. (1996) Taxonomic atlas of the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel. Class Ophiuroidea. Santa Barbara Museum of Natural History, 14, 113 - 179.","Fell, B. H. (1962) A revision of the major genera of Amphiuroid Ophiuroidea. Transactions of the Royal Society of New Zealand, Zoology, 2 (2), 1 - 26.","Maluf, L. Y. & Brusca, R. C. (2005) Chapter 18. Echinodermata. In: Hendrickx, M. E., Brusca, R. C. & Findley, L. T. (Eds.), A Distributional checklist of the macrofauna of the Gulf of California, Mexico. Arizona-Sonora Desert Museum, Tucson, Arizona, pp. 327 - 343.","Granja-Fernandez, R., Herrero-Perezrul, M. D., Lopez-Perez, R. A., Hernandez-Morales, A. & Rangel-Solis, P. D. (2015) A literature review of the Ophiuroidea (Echinodermata) from the Pacific coast of Mexico. Revista de Biologia Tropical, 63 (2), 37 - 47."]}

Amphiura remota sp. nov. Fig. 13E–J, 14A–Q http://zoobank.org/ urn:lsid:zoobank.org:act: 2758B368-7570-4981-A052-D6A951012506 Type material. MD50 DC167, north of Île Saint-Paul, 38° 24.43´S, 77° 28.64´E, 1430–1600m, Charcot dredge, 24/7/1986, MNHN IE.2019.4748 (holotype), MNHN IE.2009.1557 (16 paratypes). Other material examined. MD 50 DC34, MNHN IE.2009.1535 (2). MD 50 DC55, MNHN IE.2009.1537 (1). MD 50 DC137, MNHN IE.2009.1552 (1), MNHN IE.2009.1551 (3). Holotype description. Disc is 4.4 mm dd, petaloid, indented interradially, covered in overlapping scales, 15–16 scales from disc centre to margin, some primary scales are evident (0.3 long, 0.4 mm wide), separate, scales in disc centre 0.18 to 0.3 mm wide, becoming abruptly smaller and denser near the disc margin and outside of the radial shields (0.125 –0.175 mm wide). A slender triangular scale (0.86 mm long) separates the radial shields (Fig. 13F). Scales have microscopic glassy beads, marginal scales have finely denticulate margins (Fig. 13H). Radial shields are long (0.3 mm wide and 1.1 mm long), contiguous distally but separated proximally by the long triangular and a few smaller rounded scales, rounded distally, tapered for the proximal 2/3 of their length and terminating in a sharp angle. Ventral disc surfaces (Fig. 13G) have similar scales to dorsal margin, becoming less overlapped near oral shield. Genital slits are open along the first 3 arm segments, bordered by a prominent adradial genital scale that persists proximally until the 2nd set of arm spines, where it is followed along the slit by a series of stout plates until the distal projection of the oral plates to the rear of the oral shield. Jaws are wider than long, slit open and teeth form a buccal funnel (sensu Hendler 2018). Oral shield spear-headshaped, as wide as long, with a small distal lobe. Madreporite is larger than other shields, longer than wide with a prominent wide distal lobe. Adoral shields are slightly separated interradially, with a concave proximal margin, rounded lateral angles and extending distally to border the genital slit; exposed surface of the oral plate rectangular, is separated distally. Four teeth are visible, upper ones rounded, lower ones truncate and longer, all bordered by imperforate stereom. Infradental papillae are pentagonal, square at their base and pointed proximally, a small conical buccal scale adjacent to each infradental papilla on the oral plate, tall flask-shaped to conical adoral shield spine, 2 times as long as wide, sitting proximoventral to the large exposed 2nd oral tentacle pore. DAPs are fan-shaped then transversely ovoid, 1.3 times wider than long, with the glassy distal margin broadly convex and the proximal side produced into a very obtuse angle, separate. First VAPs are hourglass-shaped, with concave sides formed by the rounded edge of the adoral shields, a little longer than wide, with an obtuse proximal angle and truncate distal edge, contiguous with the 2nd VAP which is 1.25 times longer than wide with a straight proximal margin, divergent proximolateral sides, concave lateral margins around the large tentacle pores, and a rounded distal edge. Succeeding plates are similar but becoming progressively narrower and more tapered proximally, so that they are 2 times as long as wide by the 10th segment and only narrowly contiguous with neighbouring plates. Two arm spines are on the first segments, thereafter 3 arise from the lateral side of the arm. Spines are long and rounded, gradually tapering to a fine rounded tip, composed of sculptured stereom and fine thorns, the 2 ventral ones are longest, to 0.55 mm, the dorsal one is more slender and 0.6 times shorter. One or two middle arm spines have a diminutive thorn at the tip. Two small tentacle scales occur at right angles to each other around each tentacle pore, one on the LAP and a slightly smaller one on the VAP, which do not completely cover the pore. Colour (dry): white. Paratype variations. Other specimens from stn DC167 measure 3.5 to 7.5 mm dd. One has two oral tentacle scales on one jaw. On some specimens, the adoral shield spine is very tall and slender after the swollen base, up to 4 times as tall as wide. On smaller specimens the primary plates are relatively large. All specimens have a white or pale-grey disc and white arms. A non-type specimen from stn DC55 is four-rayed (Fig. 13I) and has notable distallydirected thorns on the tips of middle arm spines (Fig. 13J). Paratype ossicle description. Proximal (Fig. 14A) and median (Fig. 14B–C) LAPs are two times higher than long, strongly arched, relatively thin, with strongly protruding ventral portion; dorsal edge conspicuously angular; outer surface with moderately finely meshed stereom with trabecular intersections transformed into tiny tubercles; outer proximal edge with a central area of more finely meshed stereom. Spine articulations (Fig. 14D) are freestanding in the centre of the distal edge, on the same level as the outer surface stereom, moderately large, composed of parallel and completely separated lobes, the dorsal one relatively slender and slightly arched and the ventral one shorter and thicker, encompassing a moderately large muscle opening and a slightly smaller nerve opening. Inner side of LAP has three round, moderately well-defined knobs, a large one close to the centre of the inner proximal edge, a second smaller knob in the dorso-proximal corner of the inner side of the LAP, and a third smaller knob between the large one and the distal edge of the LAP; single small perforation on the ventro-distal edge of the third small knob; tentacle notch large but very shallow and inconspicuous. Vertebrae (Fig. 14F–J) are small, delicate, with relatively small, rounded muscle fossae on the distal and proximal faces; articulations very small, typically zygospondylous, with a large zygosphene on the distal face; dorso-distal muscle fossae deeply carved into the lateral side of the vertebra, with converging dorso-distal tips strongly elongated and pointed, almost projecting beyond zygocondyles; very large, deep ventral furrow, with large, moderately well-defined podial basins. Oral plates (Fig. 14M) are nearly as high as long, with adradial muscle fossa transformed into a vertical spoonshaped depression lining the entire lateral edge of articulation area. Dental plates (Fig. 14N) are undivided, rounded, with single row of large, tooth sockets surrounded by separate, strongly protruding knobs; dorsalmost tooth socket very large, perforating dental plate; second dorsalmost tooth socket smaller, penetrating dental plate but with a vertical septum. Abradial genital plates (Fig. 14O) are slightly shorter than the adradial one (Fig. 14P), very slender, bar-like, bent, without ridges or groves and with a small, concave adradio-distal tip. Radial shields (Fig. 14Q) have an irregular, rounded triangular outline, slightly oblique, with a more finely-meshed stereom in the middle part of the outer surface. A proximal dorsal arm spine (Fig. 14E), dorsal arm plate (Fig. 14K), and ventral arm plate (Fig. 14L) are also figured. Distribution. SPA (1015–2200 m). Remarks. This species shares many morphological features with the Amphiura joubini -group of species, that includes A. joubini Koehler, 1912, A. joubini abyssalis Hertz, 1927a, Amphioplus acutus Mortensen, 1936 from the Southern Ocean; Amphiura lanceolata Lyman, 1879, Amphioplus ctenacantha Baker, 1977, Amphioplus (Unioplus) cipus Baker, 1977 from New Zealand; and Amphiura otteri Ljungman, 1872 from the North Atlantic. They all form a monophyletic clade on the phylogenies in O’Hara et al. (2017) and Christodoulou et al. (2019). They share discs with long divergent radial shields, fine disc scales, spear-head-shaped oral shields, one to two long spiniform adoral shield spines, long pointed arm spines often with a terminal thorn, and two small tentacle scales. Here, all these species are reassigned to Amphiura pending a revision of this large genus. The fact that these species are spread over Amphiura, Amphioplus, Amphioplus (Unioplus) and (formerly) Amphiodia shows the artificiality of these genera (O’Hara et al. 2017). Species in the joubini -group have been referred to Amphioplus or Amphiodia on the basis that some specimens can have two distal oral papillae. However, the presence of two distal oral papillae is homoplasic, and in the case of joubini-group can be interpreted as paired adoral shield spines rather than one being a superficial oral tentacle spine (sensu Clark 1970). The new species differs from the other species in having only three arm spines, whereas all the other species have four to eight. Other differences include the bihamulate (two lateral terminal thorns) arm spines on A. ctenacantha, the notably denticulate disc scales on O. joubini, and the naked ventral side of the disc on A. lanceolata, A. ctenacantha and O. joubini abyssalis. O. joubini is partially scaled ventrally, with few scales next to the oral shields. Amphioplus acutus is a variant of O. joubini that has a third tiny oral papilla near the end of the jaw slit. The species limits of these species need to be determined. However, the morphology of the MD 50 specimens do not conform to any of these existing species. Etymology. Named from “remota”, a female Latin adjective that means distant or far away, in reference to the presence of this species in the middle of the south Indian Ocean., Published as part of O'Hara, Timothy D. & Thuy, Ben, 2022, Biogeography and taxonomy of Ophiuroidea (Echinodermata) from the Îles Saint- Paul and Amsterdam in the southern Indian Ocean, pp. 1-49 in Zootaxa 5124 (1) on pages 35-38, DOI: 10.11646/zootaxa.5124.1.1, http://zenodo.org/record/6404674, {"references":["Hendler, G. (2018) Armed to the teeth: a new paradigm for the buccal skeleton of brittle stars (Echinodermata: Ophiuroidea). Contributions in Science, 526, 189 - 311. https: // doi. org / 10.5962 / p. 324539","Koehler, R. (1912) Echinodermes (Asteries, Ophiures et Echinides). In: Deuxieme Expedition Antarctique Francaise (1908 - 10), Paris, pp. 1 - 272, 16 pls. https: // doi. org / 10.5962 / bhl. title. 85347","Hertz, M. (1927 a) Die Ophiuroiden der Deutschen Sudpolar-Expedition 1901 - 1903. Deutsche Sudpolar-Expedition, 19, 1 - 56, pls. 1 - 9.","Mortensen, T. (1936) Echinoidea and Ophiuroidea. Discovery Reports, 12, 199 - 348, 9 pls. https: // doi. org / 10.5962 / bhl. part. 8051","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 University, 6, 17 - 83, pls. 11 - 19.","Baker, A. N. (1977) Some deep-sea Ophiuroidea from New Zealand. Records of the National Museum of New Zealand, 1, 149 - 160.","Ljungman, A. V. (1872) Forteckning ofver uti Vestindien af Dr. A. Goes samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af Kongl. Vetenskaps-Akademiens Forhandlingar. Stockholm, 28, 615 - 657.","O'Hara, T. D., Hugall, A. F., Thuy, B., Stohr, S. & Martynov, A. V. (2017) Restructuring higher taxonomy using broad-scale phylogenomics: the living Ophiuroidea. Molecular Phylogenetics and Evolution, 107, 415 - 430. https: // doi. org / 10.1016 / j. ympev. 2016.12.006","Christodoulou, M., O'Hara, T. D., Hugall, A. F. & Arbizu, P. M. (2019) Dark Ophiuroid Biodiversity in a Prospective Abyssal Mine Field. Current Biology, 29 (e 3), 3909 - 3912.","Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History), Zoology, 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085"]}

O'Hara, Timothy D., Thuy, Ben (2022): Biogeography and taxonomy of Ophiuroidea (Echinodermata) from the Îles Saint- Paul and Amsterdam in the southern Indian Ocean. Zootaxa 5124 (1): 1-49, DOI: https://doi.org/10.11646/zootaxa.5124.1.1

Amphipholis squamata (Delle Chiaje 1828) Asteria squamata Delle Chiaje, 1828: 74. Amphipholis squamata.— Mortensen 1933a: 364–365.—Clark, A.M. 1976: 258.— Olbers et al. 2019: 217–218, fig. 214–215. Ophiactis minor Döderlein, 1910: 253, pl. 5(3–3a) [see Mortensen 1933b]. Amphipholis minor.— Hertz 1927a: 35. Material examined. MD 50 DC2, MNHN IE.2009.1522 (2). MD 50 WP4, MNHN IE.2009.1523 (1). MD 50 DC8, MNHN IE.2009.1524 (5). MD 50 DC44, MNHN IE.2009.1525 (16). MD 50 DC59, MNHN IE.2009.1526 (4). MD 50 DC71, MNHN IE.2009.1527 (2). MD 50 DC79, MNHN IE.2009.1528 (2). MD 50 DC105, MNHN IE.2009.1529 (1). MD 50 DC114, MNHN IE.2009.1530 (1). MD 50 CP149, MNHN IE.2009.1531 (9). MD 50 DC157, MNHN IE.2009.1532 (21). MD 50 DC167, MNHN IE.2009.1533 (1). Distribution. Arctic (29–2369 m), NW Atlantic (0–1962 m), NE Atlantic (0–1560 m), NW Pacific (0–413 m), NE Pacific (0–933 m), W Atlantic (0–353 m), E Atlantic (0–741 m), W Indian (0–750 m), E Indo-W Pacific (0–694 m), E Pacific (0–46 m), S America (0–134 m), S Africa (0–1600 m), S Australia (0–841 m), New Zealand (2–1059 m). SPA (3–1600 m). Remarks. Amphipholis squamata is a polyphyletic (O’Hara et al. 2017) assemblage of numerous genetic lineages (Boissin et al. 2008). Only synonyms relevant to the SPA are listed above. The wide bathymetric range of the MD50 samples suggests that several lineages may be present in the SPA. Hertz (1927a) named her specimens A. minor after a South African lineage, however, the diagnostic characters used by Döderlein (1910) to distinguish this species were inadequate (Mortensen 1933a). The MD50 material is small and damaged., Published as part of O'Hara, Timothy D. & Thuy, Ben, 2022, Biogeography and taxonomy of Ophiuroidea (Echinodermata) from the Îles Saint- Paul and Amsterdam in the southern Indian Ocean, pp. 1-49 in Zootaxa 5124 (1) on pages 38-39, DOI: 10.11646/zootaxa.5124.1.1, http://zenodo.org/record/6404674, {"references":["Delle Chiaje, S. (1828) Memoire sulla Storia e Notomia degli Animali senza vertebre del Regno di Napoli. Vol. 3. Societa Tipografica, Napoli, 232 pp.","Mortensen, T. (1933 a) Echinoderms of South Africa (Asteroidea: Ophiuroidea). Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 93, 215 - 400, pls. 8 - 19.","Olbers, J. M., Griffiths, C. L., O'Hara, T. D. & Samyn, Y. (2019) Field guide to the brittle and basket stars (Echinodermata: Ophiuroidea) of South Africa. Brussels: Royal Belgium Institute of Natural Sciences.","Doderlein, L. (1910) Asteroidea, Ophiuroidea, Echinoidea. In: Schultze, L. (Ed.), Forschungsreise im westlichen Sudafrica 4 (1). Denkschriften der Medizinisch-Naturwissenschaftlichen Gesellschaft zu Jena, 16, pp. 245 - 258, pls. 4 - 5.","Mortensen, T. (1933 b) The echinoderms of St. Helena. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i KObenhavn, 93, 401 - 472.","Hertz, M. (1927 a) Die Ophiuroiden der Deutschen Sudpolar-Expedition 1901 - 1903. Deutsche Sudpolar-Expedition, 19, 1 - 56, pls. 1 - 9.","O'Hara, T. D., Hugall, A. F., Thuy, B., Stohr, S. & Martynov, A. V. (2017) Restructuring higher taxonomy using broad-scale phylogenomics: the living Ophiuroidea. Molecular Phylogenetics and Evolution, 107, 415 - 430. https: // doi. org / 10.1016 / j. ympev. 2016.12.006","Boissin, E., Feral, J. P. & Chenuil, A. (2008) Defining reproductively isolated units in a cryptic and syntopic species complex using mitochondrial and nuclear markers: the brooding brittle star, Amphipholis squamata (Ophiuroidea). Molecular Ecology, 17, 1732 - 1744. https: // doi. org / 10.1111 / j. 1365 - 294 X. 2007.03652. x"]}

Amphiura brevispina Marktanner-Turneretscher, 1887 Amphiura brevispina Marktanner-Turneretscher, 1887: 301, pl. 12(14–15).— Koehler 1907b: 299. Amphiura capensis.—Clark, A.M. 1976: 258–259 [Non Amphiura (Amphiura) capensis Ljungman, 1867]. Distribution. SPA (0–2 m). Remarks. A.M. Clark (1976) identified small amphiurids collected from the coastal zone around Île Saint-Paul and Île Amsterdam as the South African species Amphiura capensis, apparently unaware that an earlier species A. brevispina Marktanner-Turneretscher, 1887 had been described from Île Saint-Paul on the Novara expedition in 1857 and later collected by the La Dive expedition in 1872 (Koehler 1907b). The two species are very similar, having oblong proximally divergent radial shields, 1/6 dd in length, small disc scales, often sparser near the oral shields, DAPs slightly wider than long, one small tentacle scale, spear-head-shaped oral shields and a small conical adoral shield spine (distal oral papilla). There are generally 4–5 small arm spines at 3–5 mm dd, the middle ones slightly flattened and truncate. Large 11 mm dd specimens of A. capensis from South Africa can have up to 7 arm spines (Olbers et al. 2019). Amphiura capensis broods its young(Mortensen1933a)and presumably A.brevispina does as well.Consequently, the common ancestor of A. capensis-brevispina presumably rafted to SPA in kelp holdfasts or on wood. Interestingly, A. capensis has also been collected from the remote island of Tristan da Cunha in the South Atlantic (Mortensen 1941). Unfortunately, we do not have DNA data for these populations. However, phylogeographic data is available for the benthic egg-laying asteroid Parvulastra exigua (Lamarck) with a similar distribution (Hart et al. 2006; Waters & Roy 2004). Network analyses of COI sequences showed that Parvulastra specimens from Amsterdam Island were phylogenetically distinct from clades on the South African and Australian coasts (Hart et al. 2006; Waters & Roy 2004). Presumably, long distance rafting is an infrequent event allowing the island populations to diverge from their African source populations. Until there is genetic data for this Amphiura, the conservative approach is taken here of recognising A. brevispina as an endemic species of the SPA islands. Except for the next species, the numerous other Amphiura specimens (, Published as part of O'Hara, Timothy D. & Thuy, Ben, 2022, Biogeography and taxonomy of Ophiuroidea (Echinodermata) from the Îles Saint- Paul and Amsterdam in the southern Indian Ocean, pp. 1-49 in Zootaxa 5124 (1) on page 35, DOI: 10.11646/zootaxa.5124.1.1, http://zenodo.org/record/6404674, {"references":["Marktanner-Turneretscher, G. (1887) Beschreibung neuer Ophiuriden und Bemerkungen zu bekannten. Annalen des Naturhistorischen Museums in Wien, 2, 291 - 316 pls. 12 - 13. https: // doi. org / 10.5962 / bhl. part. 29076","Koehler, R. (1907 b) Revision de la collection des Ophiures du Museum d'Histoire Naturelle Paris. Bulletin Scientique de la France et de la Belgique, 41, 279 - 351, pls. 10 - 14.","Ljungman, A. V. (1867) Ophiuroidea viventia hucusque cognita enumerat. Ofversigt af Kongliga Svenska Vetenskaps-Akademiens Forhandlingar Stockholm, 23, 303 - 336.","Clark, A. M. (1976) Asterozoa from Amsterdam and St Paul Islands, southern Indian Ocean. Bulletin of the British Museum (Natural History), 30, 247 - 261, pls. 1 - 2. https: // doi. org / 10.5962 / bhl. part. 2384","Olbers, J. M., Griffiths, C. L., O'Hara, T. D. & Samyn, Y. (2019) Field guide to the brittle and basket stars (Echinodermata: Ophiuroidea) of South Africa. Brussels: Royal Belgium Institute of Natural Sciences.","Mortensen, T. (1933 a) Echinoderms of South Africa (Asteroidea: Ophiuroidea). Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 93, 215 - 400, pls. 8 - 19.","Mortensen, T. (1941) Echinoderms of Tristan da Cunha. Results of the Norwegian Scientific Expedition to Tristan da Cunha 1937 - 1938, 7, 1 - 10, pl. 1.","Hart, M. W., Keever, C. C., Dartnall, A. J. & Byrne, M. (2006) Morphological and genetic variation indicate cryptic species within Lamarck's little sea star, Parvulastra (= Patiriella) exigua. Biological Bulletin, 210, 158 - 167. https: // doi. org / 10.2307 / 4134604","Waters, J. M. & Roy, M. S. (2004) Out of Africa: The Slow Train to Australasia. Systematic Biology, 53, 18 - 24. https: // doi. org / 10.1080 / 10635150490264671"]}

Family Amphiuridae Ljungman, 1867 Type genus Amphiura Forbes, 1843. Other genera Acrocnida Gisl��n, 1926; Amphichondrius Nielsen, 1932; Amphicontus Hill, 1940; Amphigyptis Nielsen, 1932; Amphiodia Verrill, 1899; Amphiomya H.L. Clark, 1939; Amphioncus H.L. Clark, 1939; Amphioplus Verrill, 1899; Amphipholis Ljungman, 1866; Amphipholizona H.L. Clark, 1915; Amphistigma H.L. Clark, 1938; Dougaloplus A.M. Clark, 1970; Microphiopholis Turner, 1985; Nannophiura Mortensen, 1933; Nudamphiura Tommasi, 1965; Ophiocentrus Ljungman, 1867; Ophiocnida Lyman, 1865; Ophiodaphne Koehler, 1930; Ophionephthys L��tken, 1869; Ophiophragmus Lyman, 1865; Ophiosphaera Brock, 1888; Ophiostigma L��tken, 1856; Paracrocnida Mortensen, 1940; Paramphichondrius Guille & Wolff, 1984; Paramphiura Koehler, 1895; Triplodia Turner & Hallan, 2011. Description ARM. Arm shape unbranched; length 3���4�� or more than 4�� disc diameter; covering without granules/spines/tubercles. Arm combs absent. Dorsal arm plate present. Accessory dorsal arm plate absent. Ventral arm plate present. Accessory ventral arm plate absent. Girdle of tiny hooks on the arms absent. Spines all similar in shape. Arm spine cross section round. The ratio of thickness of distal ⅓ of arm spine to maximum thickness (in a given cross-sectional plane) 0.6���0.9, or 0.9���1. Arm spines tapering (conical), cylindrical, or pointed; mainly lateral; predominantly parallel to arm axis (adpressed). Longest arm spines between half and one, or between one and two segments long. Tentacle scales present or absent (Ophiophragmus); pores only proximally visible along the arm; nearly as long as wide; shorter than the spine nearby; cross section oval. Position of lateral arm plate (LAP) only lateral. Dorsal and/or ventral edge(s) of LAPs without constriction. Ventral position of LAP not projecting ventro-proximalwards. Ventro-distal tip of LAP not projecting ventralwards. Arm spine articulation without dorsal and ventral lobes, or with dorsal and ventral lobes; without sigmoidal fold; arranged over entire distal LAP edge; ventralwards increasing in size. Distance between spine articulations dorsalwards increasing.. Nerve and muscle openings separated by small ridge if at all. Nerve opening approximately as large as muscle opening. DORSAL DISC. Disc pentagonal to circular; flat (about same level as arms) to high (above arm level). Integument not obscuring plates. Dorsal disc without granules; with obvious primary plates or with concealed primary plates; without scales; without spines or with spines, granules and tubercles in some genera (O���Hara et al. 2018). Dorsal disc scale/plate without tubercles. Radial shield present; without granules/spines; length between one third and half of the disc radius; scalene (oblique) triangular, or isosceles (mirror-symmetric) triangular to pear-shaped, or bar-like. VENTRAL DISC. Gonads restricted to the disc. Ventral interradii without granules/spines or with spines. Scales on ventral interradii covering the whole area of interradius or absent. The number of scales on H line of an interradius about 14���21. Ventral teeth single. Infradental papillae on oral plate present. Buccal scales on OP present. Accessory oral papillae on OP absent. Lateral oral papillae on OP absent. Adoral shield spine present. Lyman���s ossicle present. Oral shield covering less than one third of interradius; much wider than long or about as long as wide (Amphipholis). Madreporite one; with one pore or with numerous pores. Adoral shield separated or meeting in front of the oral shield. Abradial genital plate not twisted. Number of genital slits one per bursa. Genital slit longer than half the length of an interradius. Genital papillae absent. Dental plate dorsal half widest, entire (single plate). Dental sockets slit-shaped to heart-shaped, equal to or more that 50% of the width of dental plate. At least one socket on DP as perforation with septum. Teeth block-like (with square tip). Teeth on DP arranged as single column. VERTEBRAE. Vertebrae zygospondylous; beyond segment 5 with short keel; without oral bridge., Published as part of Goharimanesh, Mona, St��hr, Sabine, Mirshamsi, Omid, Ghassemzadeh, Fereshteh & Adriaens, Dominique, 2021, Interactive identification key to all brittle star families (Echinodermata; Ophiuroidea) leads to revised morphological descriptions, pp. 1-63 in European Journal of Taxonomy 766 (1) on pages 44-45, DOI: 10.5852/ejt.2021.766.1483, http://zenodo.org/record/5514587, {"references":["O'Hara T. D., Stohr S., Hugall A. F., Thuy B. & Martynov A. 2018. Morphological diagnoses of higher taxa in Ophiuroidea (Echinodermata) in support of a new classification. European Journal of Taxonomy 416: 1 - 35. https: // doi. org / 10.5852 / ejt. 2018.416"]}

Ophiuroidea is the largest class among extant echinoderms, with over 2000 described species assigned to 33 families. Here, the first identification key to the recently revised classification was developed, and revised morphological descriptions were derived from it, expanding the previous short diagnoses. The key was built by analyzing internal and external skeletal characters of predominantly the type species of each family, including at least two mutually exclusive attributes per family. Various numeric and multistate characters were used to create a traditional as well as an interactive key using the DELTA and Xper software programs­. Illustrations (SEM and digital photos) are included in the key to facilitate the assessment of character states by users. Not only is it the first identification key to the families, according to the recently proposed new classification and the examined species, but this interactive key also assists users in understanding the family level taxonomy of brittle stars. The interactive key allows new characters and states to be added, when more species will have been analyzed, without the need to reconfigure the complete key (as may be necessary with conventional keys).

Silax abditus (A.M Clark, 1970) Triodia abdita A.M Clark, 1970: 67���69. Triplodia abdita ��� Turner & Hallan 2011: 7���8. non Amphioplus abditus (Verrill, 1871): 132., Published as part of St��hr, Sabine & O'Hara, Timothy D., 2021, Deep-sea Ophiuroidea (Echinodermata) from the Danish Galathea II Expedition 1950 - 52, with taxonomic revisions, pp. 505-529 in Zootaxa 4963 (3) on page 525, DOI: 10.11646/zootaxa.4963.3.6, http://zenodo.org/record/4704429, {"references":["Turner, R. L. & Hallan, J. K. (2011) Triplodia, replacement name for Triodia A. M. Clark, 1970 (Ophiuroidea: Amphiuridae), non Hubner, 1820 (Lepidoptera: Hepialidae). Proceedings of the Biological Society of Washington, 124, 7 - 8. https: // doi. org / 10.2988 / 09 - 32.1"]}

Genus Silax Fell, 1962 Triplodia Turner & Hallan, 2011: 7���8. Triodia A.M. Clark, 1970: 67. Amphioplus Verrill, 1899 [part]: 314. Amphioplus Verrill, 1899 ��� Paterson 1985: 92., Published as part of St��hr, Sabine & O'Hara, Timothy D., 2021, Deep-sea Ophiuroidea (Echinodermata) from the Danish Galathea II Expedition 1950 - 52, with taxonomic revisions, pp. 505-529 in Zootaxa 4963 (3) on page 524, DOI: 10.11646/zootaxa.4963.3.6, http://zenodo.org/record/4704429, {"references":["Fell, H. B. (1962) A revision of the major genera of amphiurid Ophiuroidea. Transactions of the Royal Society of New Zealand, Zoology, 2, 1 - 26.","Turner, R. L. & Hallan, J. K. (2011) Triplodia, replacement name for Triodia A. M. Clark, 1970 (Ophiuroidea: Amphiuridae), non Hubner, 1820 (Lepidoptera: Hepialidae). Proceedings of the Biological Society of Washington, 124, 7 - 8. https: // doi. org / 10.2988 / 09 - 32.1","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."]}

Amphiura (Fellaria) heptacantha (Mortensen, 1940) Ophionephthys heptacantha Mortensen, 1940: 81. Amphiura (Fellaria) heptacantha— A. M. Clark 1970: 18. Habitat: Sand, 10 m (Mortensen 1940). Persian Gulf and Gulf of Oman: Chabahar Bay (17) (ibid.). Indian Ocean: Records from other areas are doubtful., Published as part of Fatemi, Yaser & Stöhr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 97, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History), 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085"]}

Published records on the Ophiuroidea fauna of the Persian Gulf and Gulf of Oman are scattered in difficult to access journals and books. This study presents a compilation of all published records, complemented with data from new samples. Distribution, habitat and depth in the study area, as well as known Indian Ocean distributions, are included. The taxonomic status of all species was evaluated, critical comments were added as applicable, and several previous records were reassigned to other species. Ophiocoma erinaceus was removed from the fauna of the Persian Gulf and Gulf of Oman, because the only published record was instead Ophiocoma schoenleinii. Previous studies assumed that 46 species of brittle star were known from the study area, but only 38 species were confirmed by this re-assessment, including two new records (Ophiothrix (Ophiothrix) foveolata and Ophiocomella sexradia). Diagnostic features are supplied for difficult to distinguish species. Five species (Macrophiothrix elongata, Amphiura fasciata, Amphiura (Ophiopeltis) hexactis, Amphioplus echinulatus, and Amphioplus seminudus) are so far worldwide known only from the Persian Gulf area. A hexamerous, fissiparous species of Ophiothela that does not concur with any known species was found and may represent a new species. All newly collected species are illustrated with photographs.

Amphiodia sp. Habitat: sediment, subtidal (Attaran-Fariman & Beygmoradi 2016). Persian Gulf and Gulf of Oman: Tis (16) (ibid.). Indian Ocean: n/a Remarks: As discussed below, the specimens erroneously reported as Amphipholis squamata by Attaran-Fariman & Beygmoradi (2016) belong in the genus Amphiodia instead. The specimen figured by them has a fully scaled disc, as is found in A. microplax, with a sharp edge that possibly bears spiny processes that are typical for A. microplax and A. obtecta (Stöhr et al. 2010). However, the figures are not detailed enough to allow assignment of the specimen to a species., Published as part of Fatemi, Yaser & Stöhr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 98, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Attaran-Fariman, G. & Beygmoradi, A. (2016) Short communication: The first report of Amphipholis squamata (Delle Chiaje, 1829) (Echinodermata: Ophiuroidea) from Chabahar Bay - northern Oman Sea. Iranian Journal of Fisheries Sciences, 15, 1254 - 1261.","Stohr, S., Cinar, M. E. & Dagli, E. (2010) Brittle stars (Echinodermata: Ophiuroidea) from the southern coast of Turkey (eastern Mediterranean): new records and revision of Amphiodia obtecta Mortensen, 1940. Zootaxa, 2483 (1), 45 - 57. https: // doi. org / 10.11646 / zootaxa. 2483.1.4"]}

Amphipholis squamata (Delle Chiaje, 1828) Asterias squamata Delle Chiaje, 1828: 74, pl. 34, fig. 1. Amphipholis squamata — Ljungman 1872: 633. Material examined: Gulf of Oman, Iran, Tis, intertidal, hand collected, January 2019, collector Yaser Fatemi, 1 spm [not kept]. Habitat: Intertidal shell and gravel bed, under a stone (present study), grass bed and subtidal sand, 10–34 m (Price 1981; present study) (Fig. 2E). Persian Gulf and Gulf of Oman: Tarut Bay (25), Jinna Island (21), Tis (16), Bahrain (ibid.). Indian Ocean: Southern Africa (Clark & Courtman-Stock 1976), West Indian Ocean (Clark & Rowe 1971), Gulf of Mannar (James 1985), Pakistan (Kazmi & Tahera 2003), Mascarene Isl. (Boissin et al. 2016), Scattered Isl. (Conand et al. 2016). Remarks: A record from Chabahar Bay by Attaran-Fariman & Beygmoradi (2016) is an error and omitted here. Their images show an animal with narrow, sunken in radial shields (Amphipholis has short, half-circle shaped ones), bell-shaped overlapping dorsal arm plates (separated in A. squamata), small narrow tentacle scales (larger oval in A. squamata), widely separated infradental papillae (close together in A. squamata), and the distal oral papilla is only slightly widened, not as wide as in A. squamata. In addition, according to the scale bar on the images, the animal had a disc diameter of 8 mm, which is larger than A. squamata (maximum 5 mm), and they reported the arms to be 6 cm long, whereas A. squamata has about 2 cm long arms. The characters fit with Amphiodia, but the images do not show the necessary details for species identification. Consequently, the material found by the present study is the first record of A. squamata from the Gulf of Oman. Amphipholis squamata has a circumtropical distribution, but it is most likely a species complex (Sponer & Roy 2002; Boissin et al. 2015)., Published as part of Fatemi, Yaser & Stöhr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 99, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Delle Chiaje, S. (1828) Memorie sulla storia e anotomia degli animali senza vertebre del Regno di Napoli. Vol. 3. Stamperia della Societa Tipografica, Naples, 232 pp.","Ljungman, A. V. (1872) Forteckning ofver uti Vestindien af Dr A. Goes samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af Kungliga Vetenskapsakademiens Forhandlingar 1871, 6, 615 - 658.","Price, A. R. G. (1981) Studies on the echinoderm fauna of the western Arabian Gulf. Journal of Natural History, 15, 1 - 15. https: // doi. org / 10.1080 / 00222938100770011","Clark, A. M. & Courtman-Stock, J. (1976) The echinoderms of Southern Africa. British Museum, London, 277 pp.","Clark, A. M. & Rowe, F. W. E. (1971) Monograph of Shallow-water Indo-west Pacific Echinoderms. Trustees of the British Museum (Natural History), London, 234 pp.","James, D. B. (1985) Echinoderm fauna of the proposed National Marine Park in the Gulf of Mannar. Proceeding of Symposium Endangered Marine Animals and Marine Parks, 1, 403 - 406.","Kazmi, Q. & Tahera, Q. (2003) A new record of Amphipholis squamata (Delle Chiaje, 1828) (Echinodermata: Ophiuroidea) in Pakistan waters. Pakistan Journal of marine science, 12, 91 - 92.","Boissin, E., Hoareau, T. B., Paulay, G. & Bruggemann, J. H. (2016) Shallow-water reef ophiuroids (Echinodermata: Ophiuroidea) of Reunion (Mascarene Islands), with biogeographic considerations. Zootaxa, 4098 (2), 273 - 297. https: // doi. org / 10.11646 / zootaxa. 4098.2.4","Conand, C., Mulochau, T., Stohr, S., Eleaume, M. & Chabanet, P. (2016) Inventory of echinoderms in the Iles Eparses (Europa, Glorieuses, Juan de Nova), Mozambique Channel, France. Acta Oecologica, 72, 53 - 61. https: // doi. org / 10.1016 / j. actao. 2015.06.007","Attaran-Fariman, G. & Beygmoradi, A. (2016) Short communication: The first report of Amphipholis squamata (Delle Chiaje, 1829) (Echinodermata: Ophiuroidea) from Chabahar Bay - northern Oman Sea. Iranian Journal of Fisheries Sciences, 15, 1254 - 1261.","Sponer, R. & Roy, M. S. (2002) Phylogeographic analysis of the brooding brittle star Amphipholis squamata (Echinodermata) along the coast of New Zealand reveals high cryptic variation and cryptic dispersal potential. Evolution, 56, 1954 - 1967.","Boissin, E., Egea, E., Feral, J. P. & Chenuil, A. (2015) Contrasting population genetic structures in Amphipholis squamata, a complex of brooding, self-reproducing sister species sharing life history traits. Marine Ecology Progress Series, 539, 165 - 177. https: // doi. org / 10.3354 / meps 11480"]}

Amphiura (Amphiura) velox Koehler, 1910 Amphiura velox Koehler, 1910: 292, pl. 15, fig. 6, pl. 16, fig. 5. Habitat: Corals, stones, sand, coral-gravel and shells, 12���33 m (Mortensen 1940). Persian Gulf and Gulf of Oman: Kharg Island (1), Stiffe���s Bank (28) (ibid.). Indian Ocean: Western Indian Ocean and Red Sea (Marsh & Marshall 1983), Madagascar (Cherbonnier & Guille 1978)., Published as part of Fatemi, Yaser & St��hr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 96, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Koehler, R. (1910) Asteries et Ophiures des iles Aru et Kei. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 33, 265 - 295.","Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Marsh, L. M. & Marshall, J. I. (1983) Some Aspects of the Zoogeography of Northwestern Australian Echinoderms (Other Than Holothurians). Bulletin of Marine Science, 33, 671 - 687.","Cherbonnier, G. & Guille, A. (1978) 48 Echinodermes: Ophiurides. Centre National de la Recherche Scientifique, Paris, 272 pp."]}

Amphioplus echinulatus Mortensen, 1940 Amphioplus echinulatus Mortensen, 1940: 97. Habitat: Gravel and shells. 38���60 m (Mortensen 1940). Persian Gulf and Gulf of Oman: Tunb Island (18) (ibid.). Indian Ocean: Appears not to have been found elsewhere., Published as part of Fatemi, Yaser & St��hr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 98, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112."]}

Amphioplus (Lymanella) laevis (Lyman, 1874) Amphiura laevis Lyman, 1874: 229, pl. 4, figs 18–21. Amphioplus (Lymanella) laevis —A. M. Clark 1970: 52. Habitat: Clay, 31–70 m (Mortensen 1940). Persian Gulf and Gulf of Oman: Jask (15), Hengam Island (9), Kuh-Mobarak (14) (ibid.). Indian Ocean: Red Sea (A. M. Clark & Rowe 1971), Madagascar (Cherbonnier & Guille 1978). Possibly a recent migrant in the Mediterranean Sea (Streftaris et al. 2005)., Published as part of Fatemi, Yaser & Stöhr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 98, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Lyman, T. (1874) Ophiuridae and Astrophytidae, new and old. Bulletin of the Museum of Comparative Zoology at Harvard College, Cambridge, 3, 221 - 272. Available from: https: // biodiversitylibrary. org / page / 6705490 (Accessed 12 Dec. 2019)","Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History), 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085","Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Clark, A. M. & Rowe, F. W. E. (1971) Monograph of Shallow-water Indo-west Pacific Echinoderms. Trustees of the British Museum (Natural History), London, 234 pp.","Cherbonnier, G. & Guille, A. (1978) 48 Echinodermes: Ophiurides. Centre National de la Recherche Scientifique, Paris, 272 pp.","Streftaris, N., Zenetos, A. & Papathanassiou, E. (2005) Globalisation in Marine Ecosystems: The Story of Non-Indigenous Marine Species Across European Seas. Oceanography and Marine Biology, 43, 419 - 453. https: // doi. org / 10.1201 / 9781420037449"]}

Amphiodia (Amphispina) microplax Burfield, 1924 Amphiodia microplax Burfield, 1924: 146. Amphiodia (Amphispina) microplax ���A. M. Clark 1970: 25. Habitat: Clay and sand, 12���69 m (Mortensen 1940). Persian Gulf and Gulf of Oman: Khoor Mousa (30), Strait of Hormuz (33) (ibid.). Indian Ocean: Red Sea (Clark & Rowe 1971), South Africa (Clark 1974), Madagascar (Cherbonnier & Guille 1978). Remarks: Amphiodia (A.) microplax and A. (A.) obtecta are difficult to distinguish and may have been confused (Mortensen 1940; St��hr et al. 2010). They differ in the ventral disc, which lacks scales in A. obtecta but is fully scaled in A. microplax. Both have spiny processes distal to the radial shields and marginal dorsal disc scales with spiny extensions (St��hr et al. 2010), which places them in the subgenus Amphispina Nielsen, 1932 as defined by A. M. Clark (1970)., Published as part of Fatemi, Yaser & St��hr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 97, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Burfield, S. T. (1924) XV. - A new species and a new variety of Ophiuroid, with notes on a collection of Ophiuroidea from the Sudanese Red Sea. Annals and Magazine of Natural History, Series 9, 13 (73), 144 - 154. https: // doi. org / 10.1080 / 00222932408633015","Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History), 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085","Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Clark, A. M. & Rowe, F. W. E. (1971) Monograph of Shallow-water Indo-west Pacific Echinoderms. Trustees of the British Museum (Natural History), London, 234 pp.","Clark, A. M. (1974) Notes on some echinoderms from southern Africa. Bulletin of the British Museum (Natural History), 26, 423 - 487. https: // doi. org / 10.5962 / bhl. part. 210","Cherbonnier, G. & Guille, A. (1978) 48 Echinodermes: Ophiurides. Centre National de la Recherche Scientifique, Paris, 272 pp.","Stohr, S., Cinar, M. E. & Dagli, E. (2010) Brittle stars (Echinodermata: Ophiuroidea) from the southern coast of Turkey (eastern Mediterranean): new records and revision of Amphiodia obtecta Mortensen, 1940. Zootaxa, 2483 (1), 45 - 57. https: // doi. org / 10.11646 / zootaxa. 2483.1.4","Nielsen, E. (1932) Papers from Dr. Th. Mortensen's Pacific Expedition 1914 - 16. LIX. Ophiurans from the Gulf of Panama, California, and the Strait of Georgia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 91, 241 - 346."]}

Paracrocnida persica Mortensen, 1940 Paracrocnida persica Mortensen, 1940: 86, pl. 1, fig. 8, textfig. 14. Habitat: Sand, gravel and subtidal mud, 0���5 m (Mortensen 1940; Price 1981, 1983; Jones 1986). Persian Gulf and Gulf of Oman: Bahrain, Kuwait, Tarut Bay (25) (ibid.). Indian Ocean: Red Sea (Clark & Rowe 1971), Mozambique (Clark 1980)., Published as part of Fatemi, Yaser & St��hr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 97, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Price, A. R. G. (1981) Studies on the echinoderm fauna of the western Arabian Gulf. Journal of Natural History, 15, 1 - 15. https: // doi. org / 10.1080 / 00222938100770011","Price, A. R. G. (1983) Echinoderms of Saudi Arabia: Echinoderms of the Arabian Gulf Coast of Saudi Arabia. In: Wittmer, W. & Buttiker, W. (Eds.), Fauna of Saudi Arabia. Karger Libri, Basel, pp. 28 - 108.","Jones, D. A. (1986) A field guide to the sea shores of Kuwait and the Arabian Gulf. University of Kuwait, Kuwait City, 204 pp.","Clark, A. M. & Rowe, F. W. E. (1971) Monograph of Shallow-water Indo-west Pacific Echinoderms. Trustees of the British Museum (Natural History), London, 234 pp.","Clark, A. M. (1980) Some Ophiuroidea from the Seychelles Islands and Inhaca, Mozambique (Echinodermata). Revue de Zoologie Africaine, 94, 534 - 558."]}

Amphiura (Ophiopeltis) tenuis (H. L. Clark, 1938) Ophionephthys tenuis H. L. Clark, 1938: 241. Ophionephthys iranica Mortensen, 1940: 84. Amphiura (Ophiopeltis) iranica —A. M. Clark 1970: 18. Amphiura (Ophiopeltis) tenuis — Clark & Rowe 1971: 96. Habitat: Subtidal mud, clay with sand, 7–12 m (Mortensen 1940). Persian Gulf and Gulf of Oman: Jask (15), Asalouyeh (3) (ibid.). Indian Ocean: Madagascar (Cherbonnier & Guille 1978), Karachi (Clark & Rowe 1971)., Published as part of Fatemi, Yaser & Stöhr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on pages 96-97, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Clark, H. L. (1938) Echinoderms from Australia. An account of collections made in 1929 and 1932. Memoirs of the Museum of Comparative Zoology, at Harvard College, Cambridge, Mass., 55, 1 - 596, pls. 1 - 28. https: // doi. org / 10.5962 / bhl. title. 49349","Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History), 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085","Clark, A. M. & Rowe, F. W. E. (1971) Monograph of Shallow-water Indo-west Pacific Echinoderms. Trustees of the British Museum (Natural History), London, 234 pp.","Cherbonnier, G. & Guille, A. (1978) 48 Echinodermes: Ophiurides. Centre National de la Recherche Scientifique, Paris, 272 pp."]}

Amphiodia (Amphispina) obtecta Mortensen, 1940 Amphiodia obtecta Mortensen, 1940: 88. Amphiodia (Amphispina) obtecta ��� St��hr et al. 2010: 46. Habitat: Brown clay, soft gray sand and mud, 8���34 m (Mortensen 1940; Price 1981). Persian Gulf and Gulf of Oman: Kharg Island (1), Bushehr (2), Farur Island (8), Hengam Island (9), Strait of Hormuz (33), Kuh-Mobarak (14), Jinna Island (21) and Tarut Bay (25) (ibid.). Indian Ocean: Appears not to have been found in the Indian Ocean but A. obtecta is a recent migrant in the Mediterranean Sea (St��hr et al. 2010)., Published as part of Fatemi, Yaser & St��hr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 97, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Stohr, S., Cinar, M. E. & Dagli, E. (2010) Brittle stars (Echinodermata: Ophiuroidea) from the southern coast of Turkey (eastern Mediterranean): new records and revision of Amphiodia obtecta Mortensen, 1940. Zootaxa, 2483 (1), 45 - 57. https: // doi. org / 10.11646 / zootaxa. 2483.1.4","Price, A. R. G. (1981) Studies on the echinoderm fauna of the western Arabian Gulf. Journal of Natural History, 15, 1 - 15. https: // doi. org / 10.1080 / 00222938100770011"]}

Amphiura crispa Mortensen, 1940 Amphiura crispa Mortensen, 1940: 77, fig. 9. Habitat: Corals, stones, sand, 5���12.5 m (Mortensen 1940; Price 1981). Persian Gulf and Gulf of Oman: Kharg Island (1), Dhahran (27), As Saffaniya (32) (ibid.). Indian Ocean: Madagascar (Cherbonnier & Guille 1978), R��union Island (Guille & Ribes 1981)., Published as part of Fatemi, Yaser & St��hr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 96, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Price, A. R. G. (1981) Studies on the echinoderm fauna of the western Arabian Gulf. Journal of Natural History, 15, 1 - 15. https: // doi. org / 10.1080 / 00222938100770011","Cherbonnier, G. & Guille, A. (1978) 48 Echinodermes: Ophiurides. Centre National de la Recherche Scientifique, Paris, 272 pp.","Guille, A. & Ribes, S. (1981) Echinodermes associes aux Scleractiniaires d'un recif frangeant de l'ile de La Reunion (ocean Indien). Bulletin Museum national d'Histoire naturelle, 4 e series, 3, 73 - 92."]}

Amphioplus seminudus Mortensen, 1940 Amphioplus seminudus Mortensen, 1940: 91, pl. 1, fig. 9. Habitat: Clay and sandy-clay, 6���17 m (Mortensen 1940; Price 1981). Persian Gulf and Gulf of Oman: Bushehr (2), Chiru (6), Hormuz Island (12), Jask (15), Tarut Bay (25), Manifa (29), Strait of Hormuz (33) (ibid.). Indian Ocean: Records from other areas could not be confirmed., Published as part of Fatemi, Yaser & St��hr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 98, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Price, A. R. G. (1981) Studies on the echinoderm fauna of the western Arabian Gulf. Journal of Natural History, 15, 1 - 15. https: // doi. org / 10.1080 / 00222938100770011"]}

Amphiura (Amphiura) sp. Habitat: Rock tidal flat, mixed rock and sand beach, 0 m (Price 1981). Persian Gulf and Gulf of Oman: Juaymah (23), Tarut Bay (25) (ibid.). Indian Ocean: n/a Remarks: The two unidentified species of Amphiura may belong to known species, because Price (1981) said that they were poorly preserved, which may have made them unrecognizable. If the material can be found, they need to be re-examined and identification with molecular methods should be tried (e.g. barcoding)., Published as part of Fatemi, Yaser & St��hr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 96, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Price, A. R. G. (1981) Studies on the echinoderm fauna of the western Arabian Gulf. Journal of Natural History, 15, 1 - 15. https: // doi. org / 10.1080 / 00222938100770011"]}

Amphiura fasciata Mortensen, 1940 Amphiura fasciata Mortensen, 1940: 76, fig. 8. Habitat: Clay, grass bed, 2 m (Mortensen 1940; Price 1981, 1983). Persian Gulf: Bushehr (2), Tarut Bay (25), Bahrain (ibid.). Indian Ocean: Appears not to have been found elsewhere., Published as part of Fatemi, Yaser & St��hr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on pages 95-96, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Price, A. R. G. (1981) Studies on the echinoderm fauna of the western Arabian Gulf. Journal of Natural History, 15, 1 - 15. https: // doi. org / 10.1080 / 00222938100770011","Price, A. R. G. (1983) Echinoderms of Saudi Arabia: Echinoderms of the Arabian Gulf Coast of Saudi Arabia. In: Wittmer, W. & Buttiker, W. (Eds.), Fauna of Saudi Arabia. Karger Libri, Basel, pp. 28 - 108."]}

Amphiura richardi Koehler, 1896b Reports for the Azores: Amphiura richardi $ Koehler, 1896b: 245–246, 1909: 178–179, pl. 27, figs. 1–2; Mortensen 1927a: 209; García-Diez et al. 2005: 49. Type locality: Azores (38°26’00”N, 26°30’45”W). See: Koehler (1909); Paterson (1985: 88). Occurrence: Northeast Atlantic; a rare species known from the Bay of Biscay (Cherbonnier 1970) to the Azores (Koehler 1909) and? Canary Islands (Paterson 1985). Depth: 850– 1,494 m (Koehler 1909, Cherbonnier 1970); AZO: 1,165 –1,494 m (Koehler 1909). Habitat: soft to hard substrate (Koehler 1909). Remarks: based on Talisman and Travailleur historical reports (e.g. Koehler 1906b, Perrier 1894), Paterson (1985) record of A. richardi from the Moroccan waters is erroneous, likely due to mislabelling. We believe that Paterson examined the same specimen reported by Koehler (1896b) from Bay of Biscay (Travailleur, sta 39, 1881: 44°05’N, 7°05’26”W). In contrast, we could not trace the original material on which Paterson (1985) based his record from the Canaries., Published as part of Madeira, Patrícia, Kroh, Andreas, Cordeiro, Ricardo, De, António M., Martins, Frias & Ávila, Sérgio P., 2019, The Echinoderm Fauna of the Azores (NE Atlantic Ocean), pp. 1-231 in Zootaxa 4639 (1) on page 21, DOI: 10.11646/zootaxa.4639.1, http://zenodo.org/record/3342161, {"references":["Koehler, R. (1896 b) Note preliminaire sur les Ophiures des premieres campagnes de la \" Princesse Alice \". Memoires. Societe Zoologique de France, 9, 241 - 253.","Koehler, R. (1909) Echinodermes provenant des campagnes du yacht Princesse-Alice (Asteries, Ophiures, Echinides et Crinoides). Resultats des campagnes scientifiques accomplies sur son yacht par Albert Ier Prince Souverain de Monaco, 34, 1 - 317.","Mortensen, T. (1927 a) Handbook of the echinoderms of the British Isles. Oxford University Press, viii + 471 pp. https: // doi. org / 10.5962 / bhl. title. 6841","Garcia-Diez, C., Porteiro, F. M., Meirinho, A., Cardigos, F. & Tempera, F. (2005) Taxonomic review of selected invertebrate groups collected during the Campaigns of the Prince Albert I of Monaco in the Azorean waters. Arquipelago. Life and Marine Sciences, 22 A, 35 - 59.","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), 1 - 162.","Cherbonnier, G. (1970) Echinodermes recoltes par la \" Thalassa \" au large de cotes d'espagne et du Golfe de Gascogne (18 - 25 AOUT 1968). Bulletin Museum National Histoire Naturelle, Paris, 2 ª Serie, 41 (5), 1266 - 1277.","Koehler, R. (1906 b) Ophiures. Expeditions Scientifiques du Travailleur et du Talisman, 8, 245 - 311.","Perrier, M. E. (1894) Stellerides. Expeditions Scientifiques du Travailleur et du Talisman. Masson, Paris, 431 pp."]}

Amphiura (Amphiura) grandisquama Lyman, 1869 Reports for the Azores: Amphiura longispina $ Koehler, 1896a: 211–212, 1898: 52–53, pl. 9, figs. 45–46; Amphiura grandisquama Lyman, 1869 — $ Koehler 1907b: 301, 1909: 177–178; Koehler 1914b: 190–191; Mortensen 1927a: 209; Madsen 1970: 177–178, fig. 12; Paterson 1985: 87, fig. 33; García-Diez et al. 2005: 48. Type locality: off Tennessee Reef (Florida Strait). See: Lyman (1869: 334–337); Madsen (1970); Paterson (1985). Occurrence: cosmopolitan, in the Atlantic, Pacific, Indian Oceans and in the?Mediterranean Sea; in the west Atlantic from South Carolina to the Caribbean (Koehler 1914a), including Tristan da Cunha (Mortensen 1936); in the east Atlantic from Iceland to the Gulf of Guinea (Mortensen 1936), including the Azores (Koehler 1909), Madeira (Jesus & Abreu 1998), Canaries, Cape Verde (Koehler 1906b), Saint Helena (Mortensen 1933c), Meteor and Josephine seamounts (Bartsch 2008). The subspecies A. grandisquama natalensis Mortensen 1933b is restricted to southern Africa (Alva & Vadon 1989). Depth: (?2)18– 2,870 m (Mortensen 1936, Cherbonnier & Sibuet 1972); AZO: 599–880 m (Koehler 1909). Habitat: soft to hard substrates (Koehler 1909). Larval stage: brooder (embryos are kept in the bursae until the juvenile stage is reached; Bartsch 2008). Remarks: Koehler (1896a, 1898) described Amphiura longispina based on a single specimen collected by Hirondelle in the Azores (sta 242: 38°48’30”N, 27°58’46”W, 861 m). On the re-examination of A. grandisquama from the Caribbean, Koehler (1907b) confirmed both are conspecific. Additionally, the species Monamphiura apicula from the Mediterranean Sea (Tortonese 1965; Koukouras et al. 2007) has been considered conspecific with A. grandisquama by some authors (e.g., Madsen 1970; Paterson 1985; Bartsch 2008). If this is the case, the distribution range of the latter should be extended to the Mediterranean Sea and to depths as shallow as 2 m (Koukouras et al. 2007)., Published as part of Madeira, Patrícia, Kroh, Andreas, Cordeiro, Ricardo, De, António M., Martins, Frias & Ávila, Sérgio P., 2019, The Echinoderm Fauna of the Azores (NE Atlantic Ocean), pp. 1-231 in Zootaxa 4639 (1) on page 23, DOI: 10.11646/zootaxa.4639.1, http://zenodo.org/record/3342161, {"references":["Lyman, T. (1869) Preliminary report on the Ophiuridae and Astrophytidae dredged in deep water between Cuba and the Florida Reef, by L. F. de Pourtales. Bulletin of the Museum of Comparative Zo ˆ logy at Harvard College, 1 (10), 309 - 354.","Koehler, R. (1896 a) Note preliminaire sur les Ophiures recueillies pendant les campagnes de \" L'Hirondelle \". Memoires. Societe Zoologique de France, 9, 203 - 213.","Koehler, R. (1898) Echinides et Ophiures provenant des campagnes du yacht Hirondelle (Golfe de Gascogne, Acores, Terre- Neuve). Resultats des campagnes scientifiques accomplies sur son yacht par Albert Ier Prince Souverain de Monaco, 12, 1 - 78.","Koehler, R. (1907 b) Revision de la collection des Ophiures du Museum d'histoire Naturelle Paris. Bulletin Sciences de la France et de la Belgique, 41, 279 - 351.","Koehler, R. (1909) Echinodermes provenant des campagnes du yacht Princesse-Alice (Asteries, Ophiures, Echinides et Crinoides). Resultats des campagnes scientifiques accomplies sur son yacht par Albert Ier Prince Souverain de Monaco, 34, 1 - 317.","Koehler, R. (1914 b) Echinoderma I: Asteroidea, ophiuroidea et Echnoidea. In: Michaelsen, W. (Ed.), Beitrage zur Kenntnis der Meeresfauna Westafrikas. 1 (2). L. Friederichsen & Co., Hamburg, pp. 127 - 303.","Mortensen, T. (1927 a) Handbook of the echinoderms of the British Isles. Oxford University Press, viii + 471 pp. https: // doi. org / 10.5962 / bhl. title. 6841","Madsen, F. J. (1970) West African Ophiuroids. Atlantidae Report, 11, 151 - 243.","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), 1 - 162.","Garcia-Diez, C., Porteiro, F. M., Meirinho, A., Cardigos, F. & Tempera, F. (2005) Taxonomic review of selected invertebrate groups collected during the Campaigns of the Prince Albert I of Monaco in the Azorean waters. Arquipelago. Life and Marine Sciences, 22 A, 35 - 59.","Koehler, R. (1914 a) A Contribution to the Study of Ophiurans of the United States National Museum. Bulletin of the United States National Museum, 84, 1 - 173. https: // doi. org / 10.5479 / si. 03629236.84.1","Mortensen, T. (1936) Echinoidea and Ophiuroidea. Discovery Reports, 12, 199 - 348. https: // doi. org / 10.5962 / bhl. part. 8051","Jesus, D. & Abreu, A. D. (1998) Contribution to the knowledge of the soft bottom echinoderms of Madeira Island. Boletim do Museu Municipal do Funchal (Historia Natural), 50 (286), 59 - 69., 4 figs.","Koehler, R. (1906 b) Ophiures. Expeditions Scientifiques du Travailleur et du Talisman, 8, 245 - 311.","Mortensen, T. (1933 c) The echinoderms of St. Helena (Other than crinoids). Papers from Dr. Th. Mortensen's Pacific Expedition 1914 - 16. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening, 93, 401 - 473.","Bartsch, I. (2008) Notes on ophiuroids from the Great Meteor Seamount (Northeastern Atlantic). Spixiana, 31 (2), 233 - 239.","Mortensen, T. (1933 b) Echinoderms of South Africa (Asteroidea and Ophiuroidea). Papers from Dr. Th. Mortensen's Pacific Expedition 1914 - 16. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening, 93, 215 - 400.","Alva, V. & Vadon, C. (1989) Ophiuroids from the western coast of Africa (Namibia and Guinea-Bissau). Scientia Marina, 53, 827 - 845.","Cherbonnier, G. & Sibuet, M. (1972) Resultats scientifiques de la campagne Noratlante: Asterides et Ophiurides. Bulletin du Museum National D'histoire Naturelle, 102 (76), 1333 - 1394.","Tortonese, E. (1965) Fauna D'Italia-Echinodermata. Edizioni Calderini, Bologna, 422 pp.","Koukouras, A., Sinis, A. I., Bobori, D., Kazantzidis, S. & Kitsos, M. - S. (2007) The echinoderm (Deuterostomia) fauna of the Aegean Sea, and comparison with those of the neighbouring seas. Journal of Biological Research, 7, 67 - 92."]}

Madeira, Patrícia, Kroh, Andreas, Cordeiro, Ricardo, De, António M., Martins, Frias, Ávila, Sérgio P. (2019): The Echinoderm Fauna of the Azores (NE Atlantic Ocean). Zootaxa 4639 (1): 1-231, DOI: https://doi.org/10.11646/zootaxa.4639.1

Amphiura sarsi Ljungman, 1872 (Fig. 6), Published as part of Madeira, Patrícia, Kroh, Andreas, Cordeiro, Ricardo, De, António M., Martins, Frias & Ávila, Sérgio P., 2019, The Echinoderm Fauna of the Azores (NE Atlantic Ocean), pp. 1-231 in Zootaxa 4639 (1) on page 21, DOI: 10.11646/zootaxa.4639.1, http://zenodo.org/record/3342161, {"references":["Ljungman, A. V. (1872) F ˆ rteckning ˆ fver uti Vestindien af Dr A. Goes samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af Kongliga Vetenskaps-Akademiens F ˆ rhandlingar, 28 (5), 615 - 658. [1871]"]}

The ophiuroid species listed were collected by benthic faunal surveys from 2016 to 2018 in submarine caves of the Ryukyu Islands, southwestern Japan. The material includes 20 species from 8 families, Amphiuridae, Ophiocomidae, Ophiodermatidae, Ophiolepididae, Ophioleucidae, Ophiomyxidae, Ophionereididae and Ophiotrichidae. This is the first comprehensive report of submarine cave-dwelling ophiuroids from Japanese waters. Also provided is a redescription of a rare species, Dougaloplus echinatus (Ljungman, 1867), with a detailed description of its ossicle morphology. The species has previously been reported from Brazil, Australia, Eastern Indian Ocean to Southern China, in 11–118 m depth, and this is the first record of D. echinatus from Japanese waters. A brief discussion of the scarcity of submarine cave systems, and the need for their conservation, is provided.

Amphioplus aff. causatus Koehler, 1905 (Fig. 2) Material examined. Ie Island (RUMF-ZE-02036[1], RUMF-ZE-02037[1], RUMF-ZE-02038[1], RUMF-ZE- 02039[1]): ���entrance��� (RUMF-ZE-02036, RUMF-ZE-02037, RUMF-ZE-02038) and ���right hole��� (RUMF-ZE- 02039) of the ���Unnamed cave���, under coral rubble, approximately 20 m depth, 24 June 2017. Remarks. The present specimens agree well with the diagnostic characters of Amphioplus causatus Koehler, 1905 in having small and numerous disc scales, long and narrow radial shields, three oral papillae, two tentacle scales (Koehler, 1905). However, there is a difference in the morphology of the oral papillae: these are flat and polygonal in our specimens, differing from the original description of the type material which has pointed and triangular oral papillae. Further more intensive comparisons are required. Sampling locality of the specimens. Ie Island, Okinawa Island Group, southwestern Japan., Published as part of Okanishi, Masanori & Fujita, Yoshihisa, 2019, A comprehensive taxonomic list of brittle stars (Echinodermata: Ophiuroidea) from submarine caves of the Ryukyu Islands, southwestern Japan, with a description of a rare species, Dougaloplus echinatus (Amphiuridae), pp. 73-98 in Zootaxa 4571 (1) on pages 76-77, DOI: 10.11646/zootaxa.4571.1.5, http://zenodo.org/record/2605976, {"references":["Koehler, R. (1905) Ophiures de l'Expedition du Siboga. Part II. Ophiures de mer profonde. Siboga-Expeditie, 45 b, 1 - 142."]}

Dougaloplus echinatus (Ljungman, 1867) [New Japanese name: Toge-ana-suna-kumohitode] (Figs 3–7) Ophiophragmus echinatus Ljungman 1867: 316. Ophiocnida echinata ? (sic)— Lyman 1874: 230 –231, pl. 4, figs 22, 23. Ophiocnida echinata. Lyman 1882: 155.— Sluiter 1898: 306.— Koehler 1898: 68, pl. 5, fig. 47; 1900: pl. 17, fig. 25; 1905: 32; 1930: 113.— Verrill 1899: 317.— H. L. Clark 1915: 250; 1932: 204; 1946, 203. Amphioplus luctator Koehler 1922: 178 –179, pl. 68, figs. 1–3; 1930: 106. Dougaloplus echinatus. A. M. Clark 1970: 35 –36, fig. 7h–k.— A. M. Clark & Rowe 1971: 80 –81,100.— Gibbs et al. 1976: 123.— Kingston 1981: 129.— Liao & A. M. Clark 1995: 200 –202, fig. 97.— Rowe & Gates 1995: 353. Ophiostigma formosa Lütken 1872: 77, pls 1–2, fig. 5.— Koehler 1898: 77, 1905: 24, pl. 2, fig. 1. Material examined. Ie Island (RUMF-ZE-02040[1], RUMF-ZE-02041[1]): “entrance” of the “Unnamed cave”, under coral rubble, approximately 20 m depth, 24 June 2017. Discs of all specimens are separated from the bodies by being cast off upon collection. Description of external morphology (RUMF-ZE-02040). Disc. Five lobed, 10 mm in diameter (Figs 3; 4A), covered by scales, approximately 100–200 µm in length (Fig. 4B). Some disc scales bear spines, approximately 70–100 µm in height (Fig. 4B, C). Exposed part of radial shields narrow and long, approximately 1 mm long, 150 µm wide distally, and gradually tapering towards disc center (Fig. 4B), slightly in contact distally but separated towards disc center. On oral surface, adoral shields triangular, wider than long, approximately 650 µm in length, 200 µm in width on inner side, 350 µm in width on outer side, in contact with the first ventral arm plates (Fig. 4E). Oral plates invisible, obscured by skin (Fig. 4E). Oral shields oblong, longer than wide, approximately 650 µm in length and 430 µm in width, and slightly pointed on proximal side (Fig. 4E). All oral shields similar in size and shape, oral shield serving as madreporite unrecognizable in external view (Fig. 4E). Interradial oral surface also covered by imbricating scales, approximately 160–200 µm in length (Fig. 4F). Some scales on peripheral area bearing spines, approximately 150 µm in length (Fig. 4F). Shapes and length of genital slits unknown because discs of all examined specimens detached (Fig. 4D). Two or three oral papillae at each side of jaw (Fig. 4E), proximal two papillae square, the distal one 1.5 times as wide as the proximal papillae, and the distal most papillae smaller and flat (Fig. 4E). Two infradental papillae at the apex of each jaw, trapezoidal, slightly larger than middle larger oral papillae (Fig. 4E). Teeth forming vertical series l row aboral to the infradental papillae, large, wider than long (Fig. 4E). Second tentacle pore inside mouth angle (Fig. 4E). Arms. Five, approximately 135–180 mm long, 1.1 mm wide and 0.7 mm high, with an arched aboral surface and flattened oral surface at proximal portion. Arms tapering gradually toward distal tip and aboral surface is flattened (Fig. 3). On proximal to middle portion of the arm, ventral arm plates pentagonal with slightly convex edges, of equal length and width, in contact with each other (Fig. 5A). On distal portion of the arm, ventral arm plates becoming triangular and separated from each other (Fig. 5C). Dorsal arm plate fan-shaped, slightly larger than wide, contacted with each other on proximal portion (Fig. 5D). On middle portion of arm, dorsal arm plates becoming pentagonal, almost twice wider than long (Fig. 5D); toward the distal tip, the plates becoming triangular, almost as long as wide, separated from each other (Fig. 5E). Lateral arm plates small, widely separated by dorsal and ventral arm plates on proximal portion (Fig. 5A, B, D, F), in contact with each other on distal portion (Fig. 5C, E) on both oral and aboral sides. Three conical arm spines, of almost the same length as the corresponding arm segment on proximal to middle portion of arms (Fig. 5A, B, D, F), though few segments have four arm spines. On distal portion, the number of spines decreasing to two and becoming acute, spiniform (Fig. 5C, E). Two oval tentacle scales, almost the same sizes at each tentacle pore (Fig. 5A). Ossicle morphology (RUMF-ZE-02040). Dorsal arm plates trapezoid, wider than long on distal portion (Fig. 6A), and triangular with convex distal edge on proximal portion (Fig. 6B). Ventral arm plates pentagonal with slightly convex lateral side on proximal portion (Fig. 6C), and pentagonal with straight lateral side on distal portion (Fig. 6D). Lateral arm plates longer than wide, aboral edge straight and oral-distal edge concave (Fig. 6E, F). On inner side, 2 defined horizontally elevated structures on center part (Fig. 6E), proximal structure slightly longer than wide on proximal portion of arm. On distal portion, the structures almost of the same size (Fig. 6I). On external side, proximal edge convex with three equal-sized spine articulations on distal edge (Fig. 6F, G), composed of parallel, horizontal dorsal and ventral lobes (Fig. 6H). The lobes almost equal-sized on proximal portion of arm but on distal portion of arm, ventral lobe slightly shorter than dorsal lobe (Fig. 6K). Large tentacle notches opening on vantro-distal edge and end in center of lateral arm plates, where a single perforation is opening (Fig. 6E, I). Arm spines conical, thicker at oral side and thinner at aboral side the arms (Fig. 6L, M), and smaller, pointed on distal portion (Fig. 6N). The spines with serrate minute spinelets on lateral surfaces (Fig. 6L, M, N). Vertebrae with zygospondylus articulation (Fig. 7A, B, E, F), longitudinal keel on aboral side (Fig. 7C, G) and longitudinal oral groove on oral side (Fig. 7D, H). Two continuous holes on the aboral keel (Fig. 7C, G). Ambulacral groove open (Fig. 7D, H). Depression for tentacles located on lateral distal side of the vertebra (Fig. 7D, H). Color. In living, disc dull yellow, arms with red and white bands (Fig. 3). Distribution. Ie Island, Okinawa Island Group, southwestern Japan (this study); Orissa, India (Koehler, 1898); Gaspar Strait, Indonesia (Ljungman, 1867; Koehler, 1905, 1930, type locality); Singpore (Ljungman, 1867); Philippines (Lyman, 1874; H. L. Clark, 1915); Gulf of Thailand (Koehler, 1930); Pernambuco, Brazil (Sluiter, 1898). Great Barrier reef, Gillet Cay, northeastern Australia (H. L. Clark, 1932; Gibbs et al., 1976; Kingston, 1981); Southern China to Taiwan Strait (Lütken, 1872; Liao & A. M. Clark, 1995). Depth range 11– 118 m. Remarks. This species falls within the genus Dougaloplus by virtue of having disc scales that bear spines and the presence of tentacle scales, and identified as D. echinatus in having numerous disc spines on the aboral disc surface and on the interradial oral peripheral surface of the disc; three oral papillae on each jaw; and narrow, mostly separated radial shield (A. M. Clark, 1970). This is the first record of Dougaloplus echinatus and the second record of species of Dougaloplus from Japanese waters. The two specimens were collected at entrances of caves, not an anchialine environment. Therefore, this discovery may not indicate an anchialine environment affinity of D. echinatus., Published as part of Okanishi, Masanori & Fujita, Yoshihisa, 2019, A comprehensive taxonomic list of brittle stars (Echinodermata: Ophiuroidea) from submarine caves of the Ryukyu Islands, southwestern Japan, with a description of a rare species, Dougaloplus echinatus (Amphiuridae), pp. 73-98 in Zootaxa 4571 (1) on pages 77-79, DOI: 10.11646/zootaxa.4571.1.5, http://zenodo.org/record/2605976, {"references":["Ljungman, A. V. (1867) Ophiuroidea viventia huc usque cognita enumerat. In: Ofversigt af Kgl. Vetenskaps-Akademiens Forhandlingar. pp. 221 - 272.","Lyman, T. (1874) Ophiuridae and Astrophytidae new and old. Bulletin of the Museum of Comparative Zoology at Harvard College, 3, 221 - 272.","Lyman, T. (1882) Report on the Opiuroidea. Report of the Scientific Results of the voyage of H. M. S. Challenger 1873 - 76. Zoology, 5, 1 - 386.","Sluiter, C. P. (1898) On a probable periodical amputation of the disk-covering by some ophiuroids. Tijdschrift der Nederlandsche Dierkundige Vereeniging, 5.","Koehler, R. (1898) Echinoderms recueillis par l'Investigator dans l'Ocean Indien, II. Les Opiures littorales. Bulletin scientifique de la France et de la Belgique, 31, 55 - 126.","Verrill, A. E. (1899) North American Ophiuroidea. I. Revision of certain families and genera of West Indian ophiurans. Transactions of the Connecticut Academy Arts and Sciences 10, 301 - 371. https: // doi. org / 10.5962 / bhl. part. 7032","Clark, H. L. (1915) Catalogue of recent ophiurans: based on the colletion of the Museum of Comparative Zoology. Memorians of the Museum of Comparative Zoology at Havard College, 25, 163 - 376.","Koehler, R. (1922) Echinodermata: Ophiuroidea. Australasian Antarctic Expedition 1911 - 1914. Scientific Report, Series C, 8, 5 - 98.","Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History) Zoology, 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085","Clark, A. M. & Rowe, F. W. E. (1971) 690 Trustees of the British Museum (Natural History) Monograph of shallow-water Indo- West Pacific Echinoderms. British Museum (Naturak History), London.","Gibbs, P. E., Clark, A. M. & Clark, C. M. (1976) Echinodermas from the northern region of the Great Barrier Reef, Australia. Bulletin of the British Museum (Natural History) Zoology, 30, 101 - 144.","Kingston, S. C. (1981) The Swain Reefs Expedition: Ophiuroidea. Records of the Australian Museum, 33, 123 - 147. https: // doi. org / 10.3853 / j. 0067 - 1975.33.1980.277","Liao, Y. & Clark, A. M. (1995) The echinoderms of southern China. Science Press, Beijing, New York.","Rowe, F. W. E. & Gates, J. (1995) Echinodermata. In: A. Wellis (Ed), Zoological Catalogue of Australia. CSIRO, Melbourne, pp. 1 - 510.","Lutken, C. F. (1872) Ophiuridarum novarum vel minus cognitarum descriptiones nonnullae. ogle nye eller mindre bekjerdte Slangestjernerne beskrevne - Med nogle Bemaerkninger om Selvdelingen hos Straaldyrene -. Oversigt over det Kongelige Danske Videnskabernes Selska bs forhandlinger, 77, 75 - 158.","Koehler, R. (1905) Ophiures de l'Expedition du Siboga. Part II. Ophiures de mer profonde. Siboga-Expeditie, 45 b, 1 - 142.","Koehler, R. (1930) Ophiures recueillies par le Docteur Th. Mortensen dans les Mers d'Australie et dans l'Archipel Malais. Papers from Dr. Th. Mortensen's Pacific Expedition 1914 - 16. LIV. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening, 89, 1 - 295.","Clark, H. L. (1932) Echinodermata (other than Asteroidea). Great Barrier Reef Expedition, 1928 - 29 Scientific Reports, 4, 197 - 239."]}

The peristomial plates are skeletal components of the interbrachial frame (or mouth frame), which is located below the true mouth of ophiuroids. Whilst the peristomial plates were extensively described and used as diagnostic characters by some early workers, for the past 100 years they have been largely neglected as a taxonomic resource. In this investigation the peristomial plates of 48 species representing 21 families were examined directly, and information on a further 61 species, including representatives of another eight families, was obtained from the published literature. Observations were made with regard to fragmentation state, relative size and orientation of the peristomial plates. Although fragmentation state showed little consistency at any taxonomic level, relative size and orientation segregated a group of families comprising species with relatively small, inclined peristomial plates, viz. Ophiotrichidae, Ophiopholidae, Ophiactidae, Amphiuridae and Ophiocomidae, together with a single hemieuryalid species ? Ophioplocus januarii. The distribution of peristomial plate traits was strongly correlated with that of several other character states pertaining to the interbrachial frame. This supported the proposition that two major types of interbrachial frame are present in ophiuroids (designated ?A? and ?B?). Current phylogenies inferred from both morphological and molecular data imply that type B is derived and has evolved independently at least twice in the orders Amphilepidida and Ophiacanthida. This represents a remarkable example of evolutionary convergence. An analysis of the distribution of all interbrachial frame character states suggested that within the Amphilepidida paedomorphosis was probably responsible for the complete reversion of the interbrachial frame to the ancestral type A condition in two families (Ophiothamnidae and Amphilepididae) of suborder Gnathophiurina and possibly responsible for varying degrees of trait reversal in the four families of suborder Ophionereidina. Such paedomorphic events may have been associated with a secondary return to the deep-sea from shallow-sea environments. Fil: Wilkie, Iain C.. University of Glasgow; Reino Unido Fil: Brogger, Martin Ignacio. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Instituto de Biología de Organismos Marinos; Argentina

Amphiura kinbergi Ljungman, 1872 (Fig. 8) Type locality. Ubatuba, Brazil. Maximum size. dd up to 9 mm (Barboza & Borges 2005). Material examined. 4 specimens (dd: 3.5���6.4 mm) from subtidal: ZUEC OPH 2129, St. 34, 1 spm; ZUEC OPH 2212, St. 71, 1 spm; ZUEC OPH 2274, St. XXVI, 1 spm; ZUEC OPH 2341, St. 20H, 1 spm. Description. Disc: (dd: 3.5 mm) star-shaped, covered by a narrow border of small and imbricating scales around the radial shields. Other portions bear small scattered and transparent scales. Radial shields four times as long as wide, large (accounting for more than half of the disc radius) (Fig. 8A). Ventral interradius with numerous, small and transparent scales embedded in the skin. Bursal slits broad (Fig. 8B). Oral shields rounded diamondshaped, as long as wide, broadly rounded distally with an obtuse angle proximally. Madreporite well developed and with pores distally. Adoral shields triangular, wider distally, with deeply concave proximal sides and separated proximally. Two lateral oral papillae, distal larger and perpendicular to the jaw, buccal scale tapered and set deeper on jaw. A pair of blunt infradental papillae, larger than lateral oral papillae (Fig. 8C). Arms: dorsal arm plates trapezoid, straight proximally, rounded distally and contiguous (Fig. 8D,F). Ventral arm plates pentagonal, as long as wide, contiguous (Fig. 8E,G). Two small tentacle scales (about 1/5 of the length of the ventral arm plate) attached only at lateral arm plate (Fig. 8E). Seven arm spines at proximal segments, decreasing to six distally; second arm spine from ventral hatchet-shaped (Fig. 8E). Lateral arm plates (Fig. 8H,I): general outline: ventral portion projecting ventro-proximalwards; ventro-distal tip not projecting ventralwards. Outer surface ornamentation: trabecular intersections not protruding. Outer proximal edge: surface lined by discernible band of different stereom structure, restricted to central part; without spurs; central part not protruding; surface without horizontal striation. Spine articulations: on same level as remaining outer surface, ventralwards increasing in size; distance between spine articulations increasing dorsalwards. Lobes simply separated, dorsal lobe clearly larger than the ventral lobe; lobes parallel, dorsalmost bent, and oriented nearly horizontal; stereom massive; sigmoidal fold absent. Inner side, ridges and knobs: dominated by two separate central knobs; without additional dorsal structure on inner side; single large perforation on inner side. Vertebrae: zygospondylous of universal type and keeled. Large groove proximal side of vertebrae dorsally corresponding to distalwards projecting dorso-distal muscular fossae of distal side (Fig. 8J). Zygocondyles dorsalwards converging and zygosphene fused with pair of zygocondyles (Fig. 8K). Dorso-distal muscular fossae transformed distalwards almost projecting beyond zygocondyles (true keel) (Fig. 8L). Zygosphene not projecting beyond ventral edge of zygocondyles (Fig. 8M). Taxonomic comments. A. kinbergi has keeled vertebrae, which are similar to that present in A. princeps, but different from Hemipholis cordifera and Ophiothrix angulata. The dorsal projections and depressions connected by dorsal accessory muscles are more evident in H. cordifera and O. angulata than in A. kinbergi and A. princeps. The holotype of A. kinbergi was described by Ljungman (1872). However, due to the loss of this specimen, the species was re-described from a Brazilian collection sampled from Ubatuba, southeastern Brazil (Thomas 1965; Tommasi 1970). Remarks. It occurs in calcareous algae, mud, sand and rubble (Tommasi 1970; Manso et al. 2008; Lima et al. 2011). A. kinbergi was collected from sand (coarse and medium sand) and rubble bottom with van Veen grab (2 spms), dredge (1 spm), and multicorer (1 spm). Distribution. Tropical Atlantic (realm), Tropical Southwestern Atlantic (province): Northeastern and Eastern Brazil (Albuquerque & Guille 1991; Borges & Amaral 2005; Magalh��es et al. 2005; Manso et al. 2008; Lima et al. 2011; Gondim et al. 2013b). Temperate South America (realm), Warm Temperate Southwestern Atlantic (province): Southeastern Brazil (Thomas 1965; Borges & Amaral 2005). From 3 up to 300 m depth (Alvarado & Sol��s-Mar��n 2013). The present study samples occurred at depths ranging from 20 to 21.5 m. Selected references. Ljungman (1872): p. 643; Thomas (1965): p. 638, fig.1; Borges & Amaral (2005): p. 265, fig. a���d; Manso et al. (2008): p. 193, fig. 20a���c [as Amphiura kinbergi]; Fell (1962): p. 10 [as Hemilepis kinbergensis]; Tommasi (1970): p. 48 [as Amphiura (Hemilepis) kinbergi]; Albuquerque (1986): p. 72, fig. 13a���c [as Amphiura (Ophiolepis) kinbergi]., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Ara���� Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on pages 26-28, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Ljungman, A. V. (1872) Forteckning ofver uti Vestindien af Dr A. Goes samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af Kungliga Vetenskaps-Akademiens Forhandlingar, 28, 615 - 658.","Borges, M. & Amaral, A. C. Z. (2005) Classe Ophiuroidea. In: Amaral, A. C. Z., Rizzo, A. E. & Arruda, E. P. (Eds.), Manual de identificacao dos invertebrados marinhos da regiao sudeste-sul do Brasil. EdUSP, Sao Paulo, pp. 237 - 272.","Thomas, L. P. (1965) The rediscovery of Amphiura kinbergi Ljungman, 1871 (Ophiuroidea: Echinodermata). Bulletin of Marine Science, 15, 638 - 641.","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8, 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Lima, M. L. F., Correia, M. D., Sovierzoski, H. H. & Manso, C. L. C. (2011) New records of Ophiuroidea (Echinodermata) from shallow waters off Maceio, State of Alagoas, Brazil. Marine Biodiversity Records, 4, 1 - 10. https: // doi. org / 10.1017 / S 175526721100090 X","Albuquerque, M. N. & Guille, A. (1991) Ophiuroidea (Echinodermata) ao largo do Brasil: Banco dos Abrolhos, cadeia submarina Vitoria-Trindade e plataforma continental adjacente. Boletim do Museu Nacional, 353, 1 - 30.","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do Estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9, 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Gondim, A. I., Dias, T. L. P. & Christoffersen, M. L. (2013 b) Annotated checklist of Echinoderms from Maranhao and Piaui States, Northeastern Brazil. Check List, 9, 510 - 518. https: // doi. org / 10.15560 / 9.3.510","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Springer, Berlin / Heidelberg, 665 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Fell, H. B. (1962) A Revision of the Major Genera of Amphiurid Ophiuroidea. Transactions of the Royal Society of New Zealand Zoology, 2, 1 - 26.","Albuquerque, M. N. (1986) Ophiuroidea Gray, 1840 (Echinodermata) da plataforma continental do norte e nordeste brasileiro. Phd thesis, Universidade de Sao Paulo, Sao Paulo, 409 pp."]}

Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley, Borges, Michela (2018): Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Araçá Bay (Southeastern Brazil), with spatial distribution considerations. Zootaxa 4405 (1): 1-66, DOI: https://doi.org/10.11646/zootaxa.4405.1.1

Genus Microphiopholis Turner, 1985 Type taxon. Microphiopholis atra (Stimpson, 1852), originally described as Ophiolepis atra. Diagnosis. Disc covered by small scales, even smaller on ventral interradius. Two lateral oral papillae, distal larger and wider than the proximal. Dorsal and ventral arm plates form overlapping series. Two tentacle scales, the one attached to the ventral arm plate rather wide. Three to five arm spines, slender and pointed (Thomas 1966; Tommasi 1970; Turner 1985). Comments. Microphiopholis was suggested by Turner (1985) to replace Micropholis, formerly assigned by Thomas (1966). In accordance with International Commission on Zoological Nomenclature, a replacement name was required because more than a century earlier, Micropholis was used for an amphibian taxon (Turner 1985). Microphiopholis is distinguished from Amphipholis by the presence of much finer scales on the disc, which explains its name (Thomas 1966; Turner 1985). These authors suggest that only Microphiopholis has a perforation in the arm vertebrae. Because this feature was also observed in other species of amphiurids and ophiotrichids (Irimura & Fujita 2003), we do not consider this character in our diagnosis. Microphiopholis presently includes six species (Stöhr et al. 2016) with three of them recorded from Brazil (Barboza & Borges 2012): M. atra (Stimpson, 1852), M. gracillima (Stimpson, 1852) and M. subtilis (Ljungman, 1867)., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Araçá Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on page 35, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Turner, R. (1985) Microphiopholis, replacement name for Micropholis Thomas, 1966 (Ophiuroidea: Amphiuridae), non Huxley, 1859 (Amphibia: Dissorophidae). Proceedings of the Biological Society of Washington, 98, 1028 - 1029.","Stimpson, W. (1852) Descriptions of two new species of Ophiolepis from the southern coast of the United States. Proceedings of the Boston Society of Natural History, 4, 224 - 226.","Thomas, L. P. (1966) A revision of the tropical American species of Amphipholis (Echinodermata: Ophiuroidea). Bulletin of Marine Science, 16, 827 - 833.","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Irimura, S. & Fujita, T. (2003) Intraspecific variation of vertebral ossicle morphology in the Ophiuroidea. In: Feral, J. - P. & David, B. (Eds.), Echinoderm Research 2001. Swets & Zeitlinger, Lisse, pp. 161 - 166.","Stohr, S. & Martynov, A. (2016) Paedomorphosis as an Evolutionary Driving Force: Insights from Deep-Sea Brittle Stars. PLoS ONE, 11, e 0164562. https: // doi. org / 10.1371 / journal. pone. 0164562","Barboza, C. A. M. & Borges, M. (2012) A checklist of the extant species of ophiuroids (Echinodermata: Ophiuroidea) from Brazilian waters. Zootaxa, 3447, 1 - 21.","Ljungman, A. V. (1867) Ophiuroidea viventia huc usque cognita, enumerat. Ofversigt af Konglige Vetenskaps-Akademiens Forhandlingar, 23, 303 - 336."]}

Microphiopholis subtilis (Ljungman, 1867) (Fig. 13) Type locality. Rio de Janeiro, Brazil. Maximum size. dd up to 5.8 mm (present study). Material examined. 105 specimens (dd: 1.5–5.8 mm) from subtidal: ZUEC OPH 2120, St. 27, 7 spms; ZUEC OPH 2134, St. I, 6 spms; ZUEC OPH 2135, St. 26, 3 spms; ZUEC OPH 2136, St. 35, 1 spm; ZUEC OPH 2137, St. 31, 1 spm; ZUEC OPH 2138, St. 33, 1 spm; ZUEC OPH 2139, St. 30, 1 spm; ZUEC OPH 2140, St. 32, 1 spm; ZUEC OPH 2141, St. III, 5 spms; ZUEC OPH 2142, St. 36, 1 spm; ZUEC OPH 2143, St. 32, 1 spm; ZUEC OPH 2144, St. 37, 1 spm; ZUEC OPH 2145, St. 37, 1 spm; ZUEC OPH 2157, St. 65, 1 spm; ZUEC OPH 2159, St. 70, 1 spm; ZUEC OPH 2160, St. 74, 1 spm; ZUEC OPH 2161, St. 74, 1 spm; ZUEC OPH 2162, St. VII, 2 spms; ZUEC OPH 2163, St. 68, 1 spm; ZUEC OPH 2165, St. 64, 1 spm; ZUEC OPH 2167, St. 74, 1 spm; ZUEC OPH 2168, St. 73, 2 spms; ZUEC OPH 2169, St. 71, 1 spm; ZUEC OPH 2170, St. 74, 3 spms; ZUEC OPH 2171, St. 72, 2 spms; ZUEC OPH 2173, St. 63, 2 spms; ZUEC OPH 2174, St. 66, 1 spm; ZUEC OPH 2180, St. VIII, 1 spm; ZUEC OPH 2184, St. 69, 2 spms; ZUEC OPH 2255, St. XVII, 3 spm; ZUEC OPH 2256, St. XX, 2 spms; ZUEC OPH 2257, St. XV, 4 spms; ZUEC OPH 2259, St. 111, 1 spm; ZUEC OPH 2260, St. 103, 1 spm; ZUEC OPH 2262, St. XVIII, 1 spm; ZUEC OPH 2263, St. 101, 1 spm; ZUEC OPH 2264, St. 104, 1 spm; ZUEC OPH 2265, St. XV, 1 spm; ZUEC OPH 2281, St. XXVI, 3 spm; ZUEC OPH 2284, St. XXVII, 9 spms; ZUEC OPH 2287, St. 147, 1 spm; ZUEC OPH 2289, St. XXII, 1 spm; ZUEC OPH 2294, St. 137, 1 spm; ZUEC OPH 2295, St. 148, 1 spm; ZUEC OPH 2297, St. 139, 1 spm; ZUEC OPH 2299, St. 148, 1 spm; ZUEC OPH 2300, St. 137, 1 spm; ZUEC OPH 2301, St. 143, 1 spm; ZUEC OPH 2302, St. 141, 1 spm; ZUEC OPH 2303, St. 143, 1 spm; ZUEC OPH 2304, St. 144, 1 spm; ZUEC OPH 2333, St. 5H, 1 spm; ZUEC OPH 2334, St. 6H, 1 spm; ZUEC OPH 2335, St. 11H, 1 spm; ZUEC OPH 2337, St. 8H, 2 spms; ZUEC OPH 2342, St. 7H, 1 spm; ZUEC OPH 2343, St. 29H, 1 spm; ZUEC OPH 2344, St. 16H, 1 spm; ZUEC OPH 2345, St. 16H, 4 spms; ZUEC OPH 2349, St. 27H, 1 spm; ZUEC OPH 2350, St. 17H, 1 spm. Description. Disc: (dd: 3.7 mm) pentagonal, covered by numerous small, imbricating and slender scales, approximately 40 between the centrodorsal and the edge of the disc. Radial shields narrow, five times as long as wide, touching distally and separated proximally by two or three elongated scales (Fig. 13A). Ventral interradius covered with scales smaller than the dorsal and more imbricated. Bursal slits broad and long (Fig. 13B). Oral shields oval with slight lateroposterior indentations. Madreporite larger than other oral shields and more whitish. Adoral shields triangular, broadened distally, and tapered proximally. Two lateral oral papillae, distal twice as wide than the proximal. A pair of rectangular infradental papillae, similar to proximal oral papilla (Fig. 13C). Arms: long and delicate. Dorsal arm plates 1.5 times as wide as long, rounded distally and tapered proximally, with an apparent line in the middle of the plate entire length of the arm, not contiguous (Fig. 13D,F). Ventral arm plates pentagonal, twice as long as wide, straight distally and tapered proximally, contiguous (Fig. 13E,G). Two tentacle scales, the larger attached to the ventral arm plate and the smaller to the lateral arm plate. Three slender arm spines, tip tapered. Lateral arm plates (Fig. 13H,I): general outline: ventral portion projecting ventro-proximalwards; ventro-distal tip not projecting ventralwards. Outer surface ornamentation: trabecular intersections protruding to form knobs approximately the same size than stereom pores. Outer proximal edge: surface lined by discernible band of different stereom structure, restricted to central part; without spurs; central part not protruding; surface without horizontal striation. Spine articulations: on same level as remaining outer surface, middle spine articulation larger; distance between spine articulations equidistant. Lobes simply separated, ventral lobe clearly larger than the dorsal lobe; lobes parallel, straight, and oriented nearly horizontal; stereom massive; sigmoidal fold absent. Inner side, ridges and knobs: dominated by two separate central knobs (appear merged); without additional dorsal structure on inner side; single large perforation on inner side. Vertebrae: zygospondylous of universal type and non-keeled. Proximal side of vertebrae dorsally without large groove on the dorsal-distal muscular fossae (Fig. 13J). Zygocondyles dorsalwards converging and zygosphene fused with pair of zygocondyles (Fig. 13K). Tapering tip on dorsal and ventral groove, visible on the proximal and distal surfaces of the vertebrae (Fig. 13J,K). Dorso-distal muscular fossae transformed distalwards projecting but far from distal edge of zygocondyles and with a vertical oval hole through the ossicle, almost a fifth as long as the vertebra (Fig. 13L,M). Zygosphene projecting beyond ventral edge of zygocondyles with projecting part as long as zygocondyles (Fig. 13M). Taxonomic comments. This is the only species that was noted as having tapering tips visible on the proximal and distal surfaces of the vertebrae. This tapering tip is related to elevated structures involving the dorsal and ventral groove. M. subtilis is easily identified by the shape of radial shields and by the orange or reddish color on each arm segment in living or recently fixed specimens (Monteiro 1987). These features were observed in most specimens in addition to other characters that clearly differentiate them from other species, such as long arms and lateral arm plates that touch dorsally. M. subitilis is similar to M. gracillima, but the latter has four to five arm spines and radial shields contiguous for almost their entire length (Tommasi 1970). Remarks. It occurs in mud and rubble bottoms (Monteiro 1987), and has typical adaptations to this habitat, including very elongated arms, small disc, and well-developed tentacle scales which protect the pore from the sediment (Tommasi 1970). M. subitilis was collected from sand (very fine, fine and medium), silt (medium, coarse and very coarse), and rubble bottom with a dredge (41% of spms), van Veen grab (37% of spms) and multicorer (21% of spms). Distribution. Tropical Atlantic (realm), Tropical Northwestern Atlantic (province): Eastern Caribbean (Tommasi 1970); Tropical Southwestern Atlantic (province): Northeastern and Eastern Brazil (Tommasi & Aron 1988; Magalhães et al. 2005; Manso et al. 2008). Temperate South America (realm), Warm Temperate Southwestern Atlantic (province): Southeastern (Tommasi 1970; Manso & Absalão 1988; Pires-Vanin et al. 1997; Netto et al. 2005; Oliveira et al. 2010; Pires-Vanin et al. 2014). Common from 2 up to 50 m depth. The present study samples occurred at depths ranging from 8.5 to 23.5 m. Selected references. Ljungman (1867): p. 314; Fell (1962): p. 13 [as Amphipholis subtilis]; Lyman (1882): p. 146 [as Amphiura subtilis]; Thomas (1966): p. 831; Tommasi (1970): p. 40, fig. 41–42 [as Micropholis subtilis]; Monteiro (1987): p. 84, est. VII–f [as Microphiopholis subtilis]., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Araçá Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on pages 38-40, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Ljungman, A. V. (1867) Ophiuroidea viventia huc usque cognita, enumerat. Ofversigt af Konglige Vetenskaps-Akademiens Forhandlingar, 23, 303 - 336.","Monteiro, A. M. G. (1987) Ophiuroidea (Echinodermata) da regiao de Ubatuba (SP) - Aspectos morfologicos e ecologicos. Phd thesis, Universidade de Sao Paulo, Sao Paulo, 166 pp.","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Tommasi, L. R. & Aron, M. A. (1988) Equinodermes da plataforma continental do sudeste do Estado da Bahia. Relatorios Internos do Instituto Oceanografico, Universidade de Sao Paulo, 19, 1 - 6.","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do Estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9, 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8, 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Manso, C. L. C. & Absalao, R. S. (1988) Ophiuroidea: Situacao Pre Operacional nos sacos de Piraquara, regiao sob influencia da descarga da Central Nuclear Almirante Alvaro Alberto (CNAAA). Revista Brasileira de Biologia, 48, 75 - 82.","Pires-Vanin, A. M. S., Corbisier, T. N., Arasaki, E. & Moellmann, A. M. (1997) Composicao e distribuicao espaco-temporal da fauna bentica no Canal de Sao Sebastiao. Relatorios Tecnicos do Instituto Oceanografico, 41, 29 - 46.","Netto, L. F., Hadel, V. F. & Tiago, C. G. (2005) Echinodermata from Sao Sebastiao Channel (Sao Paulo, Brazil). Revista de Biologia Tropical, 53, 207 - 218.","Oliveira, J. P., Oliveira, J. & Manso, C. L. C. (2010) Inventario da colecao de equinodermos do LABIMAR, Campus Prof °. Alberto Carvalho, Universidade Federal de Sergipe. Scientia Plena, 6, 1 - 14.","Pires-Vanin, A. M. S., Muniz, P. & Bromberg, S. (2014) Inventory of the marine soft bottom macrofauna of Sao Sebastiao Channel, southeastern Brazilian continental shelf. Check List, 10, 795 - 807. https: // doi. org / 10.15560 / 10.4.795","Fell, H. B. (1962) A Revision of the Major Genera of Amphiurid Ophiuroidea. Transactions of the Royal Society of New Zealand Zoology, 2, 1 - 26.","Lyman, T. (1882) Report on the Ophiuroidea dredge, by HMS Challenger, during the years 1873 - 1876. Reports on the Zoology of HMS Challenger, 5, 1 - 386.","Thomas, L. P. (1966) A revision of the tropical American species of Amphipholis (Echinodermata: Ophiuroidea). Bulletin of Marine Science, 16, 827 - 833."]}

Amphipholis januarii Ljungman, 1866 (Fig. 14) Type locality. Rio de Janeiro, Brazil. Maximum size. dd up to 6 mm (Paim et al. 2015). Material examined. 20 specimens (dd: 2.4���5.5 mm) from subtidal: ZUEC OPH 2197, St. VII, 1 spm; ZUEC OPH 2210, St. 71, 1 spm; ZUEC OPH 2245, St. XIX, 1 spm; ZUEC OPH 2249, St. XIX, 1 spm; ZUEC OPH 2268, St. 145, 3 spms; ZUEC OPH 2272, St. XXVI, 6 spms; ZUEC OPH 2327, St. 9H, 2 spms; ZUEC OPH 2340, St. 20H, 2 spms; ZUEC OPH 2352, St. XXXIV, 3 spms. Description. Disc: (dd: 4.4 mm) pentagonal, covered by irregular and imbricated small scales, approximately 20 between the centrodorsal and the edge of the disc. Radial shields narrow and twice as long as wide, separated proximally by one or two scales (Fig. 14A). Ventral interradius covered by scales similar to the dorsal surface, but more imbricated. Bursal slits narrow and long (Fig. 14B). Oral shields as long as wide, proximal half constricted or much narrower and elongated, distal half much widened and the distal edge with a lobe. Madreporite larger and more whitish than other oral shields. Adoral shields broadened distally and widely separated proximally. Two lateral oral papillae, distal larger and widened. A pair of elongated infradental papillae, widely separated from each other, in almost lateral position (Fig. 14C). Arms: dorsal arm plates broadly oval, three times as wide as long and contiguous (Fig. 14D,F). Ventral arm plates pentagonal, 1.5 times as wide as long, with pointed proximal angle and straight distal edge, barely contiguous (Fig. 14E,G). Two tentacle scales, the larger inserted on the ventral arm plate and the smaller on the lateral arm plate. Four elongated and blunt arm spines, reduced to three at the end of the arms. In segments with four spines, the second ventral-most hatchet-shaped and smaller thorny denticles at all edges. At segments with three spines, middle one hatchet-shaped (Fig. 14D). Lateral arm plates (Fig. 14H,I): general outline: ventral portion projecting ventro-proximalwards; ventro-distal tip not projecting ventralwards. Outer surface ornamentation: trabecular intersections protruding to form knobs approximately the same size than stereom pores. Outer proximal edge: surface lined by discernible band of different stereom structure, restricted to central part; without spurs; central part not protruding; surface without horizontal striation. Spine articulation: on same level as remaining outer surface, middle spine articulations larger; distance between spine articulations equidistant. Lobes simply separated, dorsal lobe clearly larger than the ventral lobe; lobes parallel, bent, and oriented nearly horizontal; stereom massive sigmoidal fold absent. Inner side, ridges and knobs: dominated by two separate central knobs; without additional dorsal structure on inner side; single large perforation on inner side. Vertebrae: zygospondylous of universal type and non-keeled. Proximal side of vertebrae dorsally without large groove on the dorsal-distal muscular fossae (Fig. 14J). Zygocondyles dorsalwards converging and zygosphene fused with pair of zygocondyles (Fig. 14K). Dorso-distal muscular fossae transformed distalwards projecting but far from distal edge of zygocondyles (Fig. 14L). Zygosphene projecting beyond ventral edge of zygocondyles with projecting part longer than zygocondyles (Fig. 14M). Taxonomic comments. Adoral shields may be joined or separated proximally, depending on the size of the specimen, making this a variable morphological characteristic (Borges 2006). Only two of the analyzed specimens had the adoral shields united proximally. Approximately 80% of the specimens examined were without the dorsal part of the disc. Remarks. As A. januarii is common in shallow waters and easily recognizable, it can be used as a tool in monitoring coastal benthic communities (Barboza et al. 2015a). It is found in coral reefs, seagrass, bryozoans, muddy, rocky and sandy bottoms (Tommasi 1970; Hendler et al. 1995; Gondim et al. 2013a; Paim et al. 2015). A. januarii was collected from sand (medium sand) and rubble bottom with a dredge (70% of spms) and van Veen grab. Distribution. Temperate Northern Atlantic (realm), Warm Temperate Northwest Atlantic (province): Carolinian and Northern Gulf of Mexico (Hendler et al. 1995). Tropical Atlantic (realm), Tropical Northwestern Atlantic (province): Southern Gulf of Mexico to Eastern Caribbean (Pomory 2007; Alvarado et al. 2008; Miloslavich et al. 2010); Tropical Southwestern Atlantic (province): Northeastern and Eastern Brazil (Lima-Verde 1969; Avila-Pires 1983; Alves & Cerqueira 2000; Magalh��es et al. 2005; Gondim et al. 2008; Oliveira et al. 2010; Lima et al. 2011; Gondim et al. 2013a; Gondim et al. 2013b; Paim et al. 2015). Temperate South America (realm), Warm Temperate Southwestern Atlantic (province): Southeastern Brazil (Ljungman 1867; Tommasi 1970; Pires- Vanin et al. 1997; Netto et al. 2005; Pires-Vanin et al. 2014). From 0 to 311 m depth (Alvarado & Sol��s-Mar��n 2013). The present study samples occurred at depths ranging from 9 to 21.5 m. Selected references. Ljungman (1866): p. 165; Thomas (1966): p. 827; Tommasi (1967): p. 1, fig. 1; Tommasi (1970): p. 35, fig. 34,35; Monteiro (1987): p. 58, est. III d-f; Hendler et al. (1995): p. 161, fig. 78, 102 c���e; Borges & Amaral (2005): p. 260, fig. a���d; Manso et al. (2008): p. 190, fig. 18 a���d; Gondim et al. (2013a): p. 59, fig. 5 a���f; Paim et al. (2015): p. 3, fig. 2a���c [as Amphipholis januarii]; Thomas (1962): p. 657, fig. 11 [as Amphipholis pachybactra]., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Ara���� Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on pages 41-43, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Ljungman, A. V. (1866) Om nagra nya arter af Ophiurider. Ofversigt af Konglige Vetenskaps-Akademiens Forhandlingar, 1867, 163 - 166.","Paim, F. G., Guerrazzi, M. C. & Borges, M. (2015) Descriptive and illustrated diagnosis of Ophiuroidea fauna (Echinodermata) in the shallow waters of North-eastern Brazil. Marine Biodiversity Records, 8, 1 - 17. https: // doi. org / 10.1017 / S 1755267215000652","Borges, M. (2006) Taxonomia, distribuicao e biologia reprodutiva de Ophiuroidea (Echinodermata) das Regioes Sudeste e Sul do Brasil. Phd thesis, Universidade Estadual Paulista \" Julio de Mesquita Filho \", Rio Claro, 143 pp.","Barboza, C. A. M., Martins, C. C. & Lana, P. C. (2015 a) Dissecting the distribution of brittle stars along a sewage pollution gradient indicated by organic markers. Marine Pollution Bulletin, 100, 438 - 444. https: // doi. org / 10.1016 / j. marpolbul. 2015.08.008","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Hendler, G., Miller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea stars, sea urchins, and allies: echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp. https: // doi. org / 10.1017 / s 0025315400031568","Gondim, A. I., Alonso, C., Dias, T. L. P., Manso, C. L. C. & Christoffersen, M. L. (2013 a) A taxonomic guide to the brittle-stars (Echinodermata, Ophiuroidea) from the State of Paraiba continental shelf, Northeastern Brazil. ZooKeys, 307, 45 - 96. https: // doi. org / 10.3897 / zookeys. 307.4673","Pomory, C. M. (2007) Key to the common shallow-water brittle stars (Echinodermata: Ophiuroidea) of the Gulf of Mexico and Caribbean Sea. Caribbean Journal of Science, Special Publication, 10, 1 - 42.","Alvarado, J. J., Solis-Marin, F. A. & Ahearn, C. (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biologia Tropical, 56, 37 - 55.","Miloslavich, P., Diaz, J. M., Klein, E., Alvarado, J. J., Diaz, C., Gobin, J., Escobar-Briones, E., Cruz-Motta, J. J., Weil, E. & Cortes, J. (2010) Marine Biodiversity in the Caribbean: Regional Estimates and Distribution Patterns. PLoS ONE, 5, e 11916. https: // doi. org / 10.1371 / journal. pone. 0011916","Lima-Verde, J. S. (1969) Primeira contribuicao ao inventario dos equinodermas do nordeste brasileiro. Arquivos de Ciencias do Mar, 9, 9 - 13.","Avila-Pires, T. C. S. (1983) Contribuicao ao estudo do genero Echinaster Muller & Troschel, 1840 (Echinodermata: Asteroidea) no litoral brasileiro. Anais da Academia Brasileira de Ciencias, 55, 431 - 448.","Alves, O. F. S. & Cerqueira, W. R. P. (2000) Echinodermata das praias de Salvador (Bahia, Brasil). Revista Brasileira de Zoologia, 17, 543 - 553. https: // doi. org / 10.1590 / S 0101 - 81752000000200024","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do Estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9, 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Gondim, A. I., Lacouth, P., Alonso, C. & Manso, C. L. C. (2008) Echinodermata da Praia do Cabo Branco, Joao Pessoa, Paraiba, Brasil. Biota Neotropica, 8, 151 - 159. https: // doi. org / 10.1590 / S 1676 - 06032008000200016","Oliveira, J. P., Oliveira, J. & Manso, C. L. C. (2010) Inventario da colecao de equinodermos do LABIMAR, Campus Prof °. Alberto Carvalho, Universidade Federal de Sergipe. Scientia Plena, 6, 1 - 14.","Lima, M. L. F., Correia, M. D., Sovierzoski, H. H. & Manso, C. L. C. (2011) New records of Ophiuroidea (Echinodermata) from shallow waters off Maceio, State of Alagoas, Brazil. Marine Biodiversity Records, 4, 1 - 10. https: // doi. org / 10.1017 / S 175526721100090 X","Gondim, A. I., Dias, T. L. P. & Christoffersen, M. L. (2013 b) Annotated checklist of Echinoderms from Maranhao and Piaui States, Northeastern Brazil. Check List, 9, 510 - 518. https: // doi. org / 10.15560 / 9.3.510","Ljungman, A. V. (1867) Ophiuroidea viventia huc usque cognita, enumerat. Ofversigt af Konglige Vetenskaps-Akademiens Forhandlingar, 23, 303 - 336.","Netto, L. F., Hadel, V. F. & Tiago, C. G. (2005) Echinodermata from Sao Sebastiao Channel (Sao Paulo, Brazil). Revista de Biologia Tropical, 53, 207 - 218.","Pires-Vanin, A. M. S., Muniz, P. & Bromberg, S. (2014) Inventory of the marine soft bottom macrofauna of Sao Sebastiao Channel, southeastern Brazilian continental shelf. Check List, 10, 795 - 807. https: // doi. org / 10.15560 / 10.4.795","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Springer, Berlin / Heidelberg, 665 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Thomas, L. P. (1966) A revision of the tropical American species of Amphipholis (Echinodermata: Ophiuroidea). Bulletin of Marine Science, 16, 827 - 833.","Tommasi, L. R. (1967) Sobre dois Amphiuridae da fauna marinha do sul do Brasil. Contribuicoes do Instituto Oceanografico, Universidade de Sao Paulo, 12, 1 - 5.","Monteiro, A. M. G. (1987) Ophiuroidea (Echinodermata) da regiao de Ubatuba (SP) - Aspectos morfologicos e ecologicos. Phd thesis, Universidade de Sao Paulo, Sao Paulo, 166 pp.","Borges, M. & Amaral, A. C. Z. (2005) Classe Ophiuroidea. In: Amaral, A. C. Z., Rizzo, A. E. & Arruda, E. P. (Eds.), Manual de identificacao dos invertebrados marinhos da regiao sudeste-sul do Brasil. EdUSP, Sao Paulo, pp. 237 - 272.","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8, 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Thomas, L. P. (1962) The shallow water amphiurid brittle stars (Echinodermata, Ophiuroidea) of Florida. Bulletin of Marine Science, 12, 623 - 694."]}

Genus Amphipholis Ljungman, 1866 Type taxon. Amphipholis januarii Ljungman, 1866. Diagnosis. Disc with scales dorsally and ventrally, without spines. Radial shields contiguous for almost their entire length or separated proximally by up to four scales. Two lateral oral papillae, distal one much larger and widened. Two tentacle scales. Three to four arm spines, usually small and pointed (Ljungman 1867; Verrill 1899a; Thomas 1962; Clark 1970; Tommasi 1970; Albuquerque 1986). Comments. Amphipholis was described from Amphipholis januarii collected from Rio de Janeiro, Brazil (Ljungman 1866). In 1872, an identification key was constructed with a total of 27 species (Ljungman 1872). Amphipholis has undergone several revisions. One of these divided the genus into three groups according to the number of tentacle scales (Fell 1962). This proposal was not accepted by Thomas (1966), who recommended organizing into three genera, including Micropholis (now Microphiopholis) and Axiognathus. This classification was accepted by Tommasi (1970), but questioned by Clark (1970) due to the proximity between A. januarii and A. squamata, type taxon of genus Axiognathus and Amphipholis respectively. Currently, Axiognathus is invalid and accepted as Amphipholis (Stöhr et al. 2016). Amphipholis is similar to Amphiodia, but the main difference between them is the shape and size of the distal oral papilla. In Amphipholis it is large, greater than the proximal one, rectangular and wide, tending to cover the oral slit. In Amphiodia the distal lateral oral papilla is circular or triangular (Verrill 1899a). Currently, 26 species are accepted (Stöhr et al. 2016), two of which are recorded from Brazil (Barboza & Borges 2012): A. januarii Ljungman, 1866 and A. squamata (Delle Chiaje, 1828)., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Araçá Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on page 40, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Ljungman, A. V. (1866) Om nagra nya arter af Ophiurider. Ofversigt af Konglige Vetenskaps-Akademiens Forhandlingar, 1867, 163 - 166.","Ljungman, A. V. (1867) Ophiuroidea viventia huc usque cognita, enumerat. Ofversigt af Konglige Vetenskaps-Akademiens Forhandlingar, 23, 303 - 336.","Verrill, A. E. (1899 a) Report on the Ophiuroidea collected by the Bahama expedition in 1893. Bulletin from the Laboratories of Natural History of the State University of Iowa, 5, 1 - 88.","Thomas, L. P. (1962) The shallow water amphiurid brittle stars (Echinodermata, Ophiuroidea) of Florida. Bulletin of Marine Science, 12, 623 - 694.","Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History), Zoology, 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Albuquerque, M. N. (1986) Ophiuroidea Gray, 1840 (Echinodermata) da plataforma continental do norte e nordeste brasileiro. Phd thesis, Universidade de Sao Paulo, Sao Paulo, 409 pp.","Ljungman, A. V. (1872) Forteckning ofver uti Vestindien af Dr A. Goes samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af Kungliga Vetenskaps-Akademiens Forhandlingar, 28, 615 - 658.","Fell, H. B. (1962) A Revision of the Major Genera of Amphiurid Ophiuroidea. Transactions of the Royal Society of New Zealand Zoology, 2, 1 - 26.","Thomas, L. P. (1966) A revision of the tropical American species of Amphipholis (Echinodermata: Ophiuroidea). Bulletin of Marine Science, 16, 827 - 833.","Stohr, S. & Martynov, A. (2016) Paedomorphosis as an Evolutionary Driving Force: Insights from Deep-Sea Brittle Stars. PLoS ONE, 11, e 0164562. https: // doi. org / 10.1371 / journal. pone. 0164562","Barboza, C. A. M. & Borges, M. (2012) A checklist of the extant species of ophiuroids (Echinodermata: Ophiuroidea) from Brazilian waters. Zootaxa, 3447, 1 - 21.","Delle Chiaje, S. (1828) Memorie sulla storia e notomia degli animali senza vertebre del regno di Napoli. Napoli: Fratelli Fernandes, and Societa Tipografica, 1, 1 - 232. https: // doi. org / 10.5962 / bhl. title. 10021"]}

Ophiophragmus luetkeni (Ljungman, 1872) (Fig. 7) Type locality. Tortola, British Virgin Islands. Maximum size. dd up to 9 mm (Barboza & Borges 2005). Material examined. 2 specimens (dd: 5.7–9.1 mm) from intertidal: ZUEC OPH 2164, St. 48, 1 spm; ZUEC OPH 2346, St. 13H, 1 spm. Description. Disc: (dd: 5.7 mm) circular or pentagonal, covered by numerous small and imbricating scales, approximately 20 between the centrodorsal and the edge of the disc. Radial shields as long as wide, separated by three scales. Fence of tapered papillae at the edge of the disc interradius (Fig. 7A,C). Ventral interradius covered with scales smaller than the dorsal and strongly imbricated. Bursal slits broad (Fig. 7B). Oral shields spearheadshaped, distally rounded, proximal angle acute, with lateroposterior indentations (Fig. 7D). Madreporite larger than other oral shields, but with similar shape and with pores distally (Fig. 7E). Adoral shields triangular, wider distally and united proximally. Two lateral oral papillae, distal slightly larger than the proximal. A pair of infradental papillae, well developed and rectangular (Fig. 7D). Arms: dorsal arm plates semi-circular to rectangular, three times as wide as long, some fragmented, contiguous (Fig. 7F,H). Ventral arm plates pentagonal, twice as wide as long, distally straight and proximal angle acute, contiguous (Fig. 7G,I). Two tentacle scales on the proximal segments, one attached to ventral plate and one to lateral plate. Three pointed arm spines (Fig. 7G). Lateral arm plates (Fig. 7J,K): general outline: ventral portion projecting ventro-proximalwards; ventro-distal tip not projecting ventralwards. Outer surface ornamentation: trabecular intersections protruding to form knobs approximately the same size as stereom pores. Outer proximal edge: surface lined by discernible band of different stereom structure, restricted to central part; without spurs; central part not protruding; surface without horizontal striations. Spine articulations: on same level as remaining outer surface; sizes all similar; distance between spine articulations increasing dorsalwards. Lobes simply separated, dorsal lobe clearly larger than the ventral lobe; lobes parallel, dorsalmost bent, and oriented nearly horizontal; stereom massive; sigmoidal fold absent. Inner side, ridges and knobs: dominated by two separate central knobs; without dorsal structure on inner side; single large perforation on inner side. Vertebrae: zygospondylous of universal type and non-keeled. Proximal side of vertebrae dorsally without large groove on the dorsal-distal muscular fossae (Fig. 7L). Zygocondyles dorsalwards converging and zygosphene fused with pair of zygocondyles (Fig. 7M). Dorso-distal muscular fossae transformed distalwards projecting far from distal edge of zygocondyles (Fig. 7N). Zygosphene projecting beyond ventral edge of zygocondyles with projecting part longer than zygocondyles (Fig. 7O). Taxonomic comments. Only two specimens of O. luetkeni were collected, one with a regenerating disc (Fig. 7) and another without the disc. They were identified by comparing with descriptions in the literature (Thomas 1962; Tommasi 1970; Borges & Amaral 2005), and with specimens from ZUEC OPH (Ophiophragmus brachyactis, O. filograneus, O. pulcher and O. wurdemanii). O. brachyactis differs from O. luetkeni in having two or three terminal hooks on the second arm spines. O. filograneus differs in having a disc with papillae on the ventral surface and bluntly rounded marginal disc papillae. O. pulcher differs in having arms banded with green, and a green or red stripe the length of the dorsal arm surface and O. wurdemanii differs in having arms banded with dark brown or black. Remarks. Often only a few specimens of O. luetkeni are collected, possibly indicating it is not abundant (Borges & Amaral 2005). It occurs in muddy, sandy and silt bottoms (Clark 1918; Tommasi 1970; Borges & Amaral 2005; Manso et al. 2008). O. luetkeni was collected from very fine sand with corer. Distribution. Tropical Atlantic (realm), Tropical Southwestern Atlantic (province): Northeastern and Eastern Brazil (Manso et al. 2008; Correia & Sovierzoski 2013). Temperate South America (realm), Warm Temperate Southwestern Atlantic (province): Southeastern (Tommasi 1970; Manso & Absalão 1988; Pires-Vanin et al. 1997; Netto et al. 2005; Oliveira et al. 2010; Pires-Vanin et al. 2014). From intertidal to 50 m depth (Alvarado & Solís-Marín 2013). The present study samples were collected in the intertidal. Selected references. Ljungman (1872): p. 636 [as Amphipholis luetkeni]; Tommasi (1970): p. 31, fig. 28, 29; Monteiro (1987): p. 88, est.Ve,f; Borges & Amaral (2005): p. 269, fig. a–d [as Ophiophragmus lutkeni]; Manso et al. (2008): p. 193, fig. 19f,g [as Ophiophragmus luetkeni]., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Araçá Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on pages 24-26, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Ljungman, A. V. (1872) Forteckning ofver uti Vestindien af Dr A. Goes samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af Kungliga Vetenskaps-Akademiens Forhandlingar, 28, 615 - 658.","Borges, M. & Amaral, A. C. Z. (2005) Classe Ophiuroidea. In: Amaral, A. C. Z., Rizzo, A. E. & Arruda, E. P. (Eds.), Manual de identificacao dos invertebrados marinhos da regiao sudeste-sul do Brasil. EdUSP, Sao Paulo, pp. 237 - 272.","Thomas, L. P. (1962) The shallow water amphiurid brittle stars (Echinodermata, Ophiuroidea) of Florida. Bulletin of Marine Science, 12, 623 - 694.","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Clark, H. L. (1918) Brittle-stars, new and old. Bulletin of the Museum of Comparative Zoology at Harvard College, 62, 263 - 338.","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8, 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Correia, M. D. & Sovierzoski, H. H. (2013) Endemic Marine Benthic Invertebrates on the Brazilian Reef Ecosystems. Journal of Earth Science and Engineering, 3, 672 - 682.","Manso, C. L. C. & Absalao, R. S. (1988) Ophiuroidea: Situacao Pre Operacional nos sacos de Piraquara, regiao sob influencia da descarga da Central Nuclear Almirante Alvaro Alberto (CNAAA). Revista Brasileira de Biologia, 48, 75 - 82.","Pires-Vanin, A. M. S., Corbisier, T. N., Arasaki, E. & Moellmann, A. M. (1997) Composicao e distribuicao espaco-temporal da fauna bentica no Canal de Sao Sebastiao. Relatorios Tecnicos do Instituto Oceanografico, 41, 29 - 46.","Netto, L. F., Hadel, V. F. & Tiago, C. G. (2005) Echinodermata from Sao Sebastiao Channel (Sao Paulo, Brazil). Revista de Biologia Tropical, 53, 207 - 218.","Oliveira, J. P., Oliveira, J. & Manso, C. L. C. (2010) Inventario da colecao de equinodermos do LABIMAR, Campus Prof °. Alberto Carvalho, Universidade Federal de Sergipe. Scientia Plena, 6, 1 - 14.","Pires-Vanin, A. M. S., Muniz, P. & Bromberg, S. (2014) Inventory of the marine soft bottom macrofauna of Sao Sebastiao Channel, southeastern Brazilian continental shelf. Check List, 10, 795 - 807. https: // doi. org / 10.15560 / 10.4.795","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Springer, Berlin / Heidelberg, 665 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Monteiro, A. M. G. (1987) Ophiuroidea (Echinodermata) da regiao de Ubatuba (SP) - Aspectos morfologicos e ecologicos. Phd thesis, Universidade de Sao Paulo, Sao Paulo, 166 pp."]}

Genus Ophiophragmus Lyman, 1865 Type taxon. Ophiophragmus wurdemanii (Lyman, 1860), originally described as Amphiura wurdemanii. Diagnosis. Disc with scales dorsally and ventrally. Radial shields partially or entirely joined. Fence of papillae at the edge of the disc interradius. Two lateral oral papillae, distal slightly larger than the proximal. Dorsal arm plates usually as wide as long. One to three tentacle scales. Three arm spines (Lyman 1865; Thomas 1962; Tommasi 1970; Albuquerque 1986). Comments. The genus Ophiophragmus was proposed for species of Amphiura possessing a fence of papillae at the edge of the disc (Lyman 1865). This is the most helpful diagnostic feature for the genus (Clark 1918). However, other morphological characteristics must be examined as these structures can be confused with the uppermost series of plates on the dorsal interradial areas of some Amphiodia (Clark 1918). The shape of arm spines, oral shields, and dorsal arm plates are also regarded as diagnostic features for the genus (Clark 1918; Thomas 1962). Ophiophragmus presently includes 18 species (Stöhr et al. 2016) with seven being recorded from Brazil (Barboza & Borges 2012): O. brachyactis H.L. Clark, 1915, O. cubanus (A. H. Clark, 1917), O. filograneus (Lyman, 1875), O. luetkeni (Ljungman, 1871), O. pulcher H. L. Clark, 1918, O. septus (Lütken, 1859) and O. wurdemanii (Lyman, 1860)., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Araçá Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on page 24, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Lyman, T. (1865) Ophiuridae and Astrophytidae. University Press: Welch, Bigelow, & Company, Cambridge, 203 pp.","Lyman, T. (1860) Descriptions of new Ophiuridae, belonging to the Smithsonian Institution and to the Museum of Comparative Zoology at Cambridge. Proceedings of the Boston Society of Natural History, 7, 193 - 204. https: // doi. org / 10.5962 / bhl. part. 4822","Thomas, L. P. (1962) The shallow water amphiurid brittle stars (Echinodermata, Ophiuroidea) of Florida. Bulletin of Marine Science, 12, 623 - 694.","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Albuquerque, M. N. (1986) Ophiuroidea Gray, 1840 (Echinodermata) da plataforma continental do norte e nordeste brasileiro. Phd thesis, Universidade de Sao Paulo, Sao Paulo, 409 pp.","Clark, H. L. (1918) Brittle-stars, new and old. Bulletin of the Museum of Comparative Zoology at Harvard College, 62, 263 - 338.","Stohr, S. & Martynov, A. (2016) Paedomorphosis as an Evolutionary Driving Force: Insights from Deep-Sea Brittle Stars. PLoS ONE, 11, e 0164562. https: // doi. org / 10.1371 / journal. pone. 0164562","Barboza, C. A. M. & Borges, M. (2012) A checklist of the extant species of ophiuroids (Echinodermata: Ophiuroidea) from Brazilian waters. Zootaxa, 3447, 1 - 21.","Clark, H. L. (1915) Catalogue of Recent Ophiurans: Based on the Collection of the Museum of Comparative Zoology. Memoirs of the Museum of Comparative Zoology, 25, 165 - 376.","Lyman, T. (1875) Zoological results of the Hassler Expedition. II. Ophiuridae and Astrophytidae. Illustrated Catalogue of the Museum of Comparative Zoology, 8, 1 - 34.","Lutken, C. F. (1859) Additamenta ad historiam Ophiuridarum: Beskrivelser af nye eller hidtil kun ufuldstaendigt kjendte Arter af Slangestjerner. Kongelige Danske Videnskabernes Selskabs Skrifter, Copenhagen, 95 pp. [pp. 177 - 271]"]}

Genus Amphiura Forbes, 1843 Type taxon. Amphiura chiajei Forbes, 1843. Diagnosis. Disc with scales dorsally and ventrally. Radial shields as long as wide, completely separate or united distally. Two lateral oral papillae, buccal scale higher up on the jaw. None, one or two tentacle scales. Three to seven arm spines (Forbes 1843; Verrill 1899a; Thomas 1962; Tommasi 1970; Albuquerque 1986). Comments. Amphiura has undergone several taxonomic revisions since its original description by Forbes (1843). The most currently accepted classification split the group into four genera: Amphiura, Amphipholis, Amphioplus, and Amphiodia (Verrill 1899a). However, due to the large number of species, several authors have proposed the further splitting into four subgenera (Stöhr et al. 2016): Amphiura Forbes, 1843, Fellaria A. M. Clark, 1970, Ophiopeltis Dürben & Koren, 1846, and Ophionema Lütken, 1869. Amphiura comprises 208 species, 18 of which are recorded from Brazil (Barboza & Borges 2012): A. algida Koehler, 1911, A. (Amphiura) callida Albuquerque, Campos-Creasey & Guille, 2011, A. (Amphiura) complanata Ljungman, 1867, A. crassipes Ljungman, 1867, A. deichmani Tommasi, 1965, A. fibulata Koehler, 1913, A. flexuosa (Ljungman, 1867), A. (Ophionema) intricata (Lütken, 1859), A. iraciae Tommasi & Oliveira, 1976, A. joubini Koehler, 1912, A. kinbergi Ljungman, 1872, A. latispina Ljungman, 1867, A. muelleri Marktanner-Turneretscher, 1887, A. palmeri Ljungman, 1872, A. princeps Koehler, 1907, A. rosae Tommasi & Oliveira, 1976, A. semiermis Lütken, 1869 and A. stimpsoni Lütken, 1859., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Araçá Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on page 26, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Forbes, E. (1843) On the Radiata of the Eastern Mediterranean. Part I., Ophiuridae. Transactions of the Linnean society of London, 19, 143 - 153. https: // doi. org / 10.1111 / j. 1096 - 3642.1842. tb 00360. x","Verrill, A. E. (1899 a) Report on the Ophiuroidea collected by the Bahama expedition in 1893. Bulletin from the Laboratories of Natural History of the State University of Iowa, 5, 1 - 88.","Thomas, L. P. (1962) The shallow water amphiurid brittle stars (Echinodermata, Ophiuroidea) of Florida. Bulletin of Marine Science, 12, 623 - 694.","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Albuquerque, M. N. (1986) Ophiuroidea Gray, 1840 (Echinodermata) da plataforma continental do norte e nordeste brasileiro. Phd thesis, Universidade de Sao Paulo, Sao Paulo, 409 pp.","Stohr, S. & Martynov, A. (2016) Paedomorphosis as an Evolutionary Driving Force: Insights from Deep-Sea Brittle Stars. PLoS ONE, 11, e 0164562. https: // doi. org / 10.1371 / journal. pone. 0164562","Clark, A. M. (1970) Notes on the family Amphiuridae (Ophiuroidea). Bulletin of the British Museum (Natural History), Zoology, 19, 1 - 81. https: // doi. org / 10.5962 / bhl. part. 24085","Barboza, C. A. M. & Borges, M. (2012) A checklist of the extant species of ophiuroids (Echinodermata: Ophiuroidea) from Brazilian waters. Zootaxa, 3447, 1 - 21.","Koehler, R. (1911) Asteries, Ophiures, et Echinides de l'Expedition Antarctique Anglaise de 1907 - 1909. British Antarctic Expedition 1907 - 1909. Reports on the Scientific Investigations, 2 (Biology), 4, 25 - 66. https: // doi. org / 10.5962 / bhl. title. 12579","Ljungman, A. V. (1867) Ophiuroidea viventia huc usque cognita, enumerat. Ofversigt af Konglige Vetenskaps-Akademiens Forhandlingar, 23, 303 - 336.","Lutken, C. F. (1859) Additamenta ad historiam Ophiuridarum: Beskrivelser af nye eller hidtil kun ufuldstaendigt kjendte Arter af Slangestjerner. Kongelige Danske Videnskabernes Selskabs Skrifter, Copenhagen, 95 pp. [pp. 177 - 271]","Ljungman, A. V. (1872) Forteckning ofver uti Vestindien af Dr A. Goes samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af Kungliga Vetenskaps-Akademiens Forhandlingar, 28, 615 - 658.","Koehler, R. (1907) Revision de la collection des ophiures du Museum d'Histoire Naturelle de Paris. Bulletin Scientique de la France et de la Belgique, 41, 279 - 351."]}

Amphiodia pulchella (Lyman, 1869) (Fig. 10) Type locality. Florida, United States. Maximum size. dd up to 5.2 mm (Thomas 1962). Material examined. 17 specimens (dd: 1.7–4.7 mm) from subtidal: ZUEC OPH 2132, St. 34, 1 specimen (spm); ZUEC OPH 2176, St. XII, 1 spm; ZUEC OPH 2205, St. XII, 4 spms; ZUEC OPH 2214, St. 71, 1 spm; ZUEC OPH 2276, St. XXVI, 8 spms; ZUEC OPH 2286, St. 147, 1 spm; ZUEC OPH 2347, St. 20H, 1 spm. Description. Disc: (dd: 4 mm) circular with soft interradial depressions, covered by numerous small and imbricating scales, approximately 25 between the centrodorsal and the edge of the disc. Central primary plate and primary radial plates evident and larger than scales. Radial shields four times as long as wide, narrow and contiguous throughout and separated proximally by three small scales, one elongated wedge-shaped and the others semi-triangular (Fig. 10A). Ventral interradius covered by scales similar to the dorsal surface. Bursal slits long and wide (Fig. 10B). Oral shields twice as long as wide, narrow proximally, rounded edges distally and small lateroposterior re-entrances. Madreporite larger than other oral shields, more whitish and with pores at the distal margin. Adoral shields semi-triangular and separated proximally. Two lateral oral papillae, distal slightly larger and circular. One pair of elongated infradental papillae widely separated from each other (Fig. 10C). Arms: dorsal arm plates contiguous, twice as wide as long, proximal edge strongly convex, distal edge straight (Fig. 10D,F). Ventral arm plates pentagonal, as long as wide, with proximal edge pointed and distal slightly convex, first 4-5 contiguous, and others separated by lateral arm plates (Fig. 10E,G). One tentacle scale, half as long as ventral arm plate and attached to lateral plate (Fig. 10E). Three arm spines, middle one blunt, flattened dorsoventrally, tip hatchet-shaped. Lateral arm plates (Fig. 10H,I): general outline: ventral portion projecting ventro-proximalwards; ventro-distal tip not projecting ventralwards. Outer surface ornamentation: trabecular intersections protruding to form knobs larger than stereom pores on most of outer surface. Outer proximal edge: surface lined by discernible band of different stereom structure, restricted to central part; without spurs; central part not protruding; surface without horizontal striation. Spine articulation: on same level as remaining outer surface, sizes all similar; distance between spine articulation equidistant. Lobes simply separated, dorsal lobe clearly larger than the ventral lobe; lobes parallel, straight, and oriented nearly horizontal; stereom massive; sigmoidal fold absent. Inner side, ridges and knobs: inner side dominated by two separate central knobs; without additional dorsal structure on inner side; single large perforation on inner side. Vertebrae: zygospondylous of universal type and non-keeled. Proximal side of vertebrae dorsally without large groove on the dorsal-distal muscular fossae (Fig. 10J). Zygocondyles dorsalwards converging and zygosphene fused with pair of zygocondyles (Fig. 10K). Dorso-distal muscular fossae transformed distalwards projecting but far from distal edge of zygocondyles (Fig. 10L). Zygosphene projecting beyond ventral edge of zygocondyles with projecting part longer than zygocondyles (Fig. 10M). Taxonomic comments. This species was easily identified by its single tentacle scale (Koehler 1914), differentiating it from other Amphiodia species in the present collection. Evident primary plates were observed in specimens up to 4 mm of dd, however in the largest specimen these plates were almost imperceptible. The lateral arm plates are concave, “C” shaped causing them to touch dorsally and ventrally after the first 4–5 segments. This condition is only observed ventrally due to the shape of the ventral arm plate, which is pentagonal, unlike the dorsal surface where the dorsal arm plates cover the lateral arm plates. Remarks. It occurs in high densities (Hendler et al. 1995), often with the amphiurid Microphiopholis atra and the ophiactids Hemipholis cordifera and Ophiactis lymani (Tommasi 1970). A. pulchella was found with these species and with the amphiurids Amphiodia riisei, Amphipholis januarii, Amphiura kinbergi, Amphiura princeps and Microphiopholis subtilis. As with most amphiurids, A. pulchella extends two to four arms across the sediment surface, with its arm tips slightly raised to feed (Hendler et al. 1995). This species can be found on coral reefs, seagrass, muddy, rocky and sandy bottom (Alvarado & Solís-Marín 2013). A. pulchella was collected from sandy bottom (coarse and medium sand), medium silt, and rubble bottom with a dredge (70% of spms) and van Veen grab. Distribution. Tropical Atlantic (realm), Tropical Northwestern Atlantic (province): Floridian to Eastern Caribbean (Hendler et al. 1995; Pomory 2007; Miloslavich et al. 2010); Tropical Southwestern Atlantic (province): Northeastern and Eastern Brazil (Magalhães et al. 2005; Manso et al. 2008). Temperate South America (realm), Warm Temperate Southwestern Atlantic (province): Southeastern Brazil (Tommasi 1970; Manso & Absalão 1988; Manso 1989, 1993; Pires-Vanin et al. 1997; Netto et al. 2005; Pires-Vanin et al. 2014) and Uruguay – Buenos Aires Shelf (Alvarado & Solís-Marín 2013). Commonly found at shallow depths up to 80 m (Tommasi 1970; Manso et al. 2008), but there are records of up to 370 m (Alvarado & Solís-Marín 2013). The present study samples occurred at depths ranging from 19 to 22 m. Selected references. Lyman (1869): p. 337 [as Amphiura pulchella]; Clark (1915): p. 250; Fell (1962): p. 5; Thomas (1962): p. 641, fig. 5; Parslow & Clark (1963): p. 26; Tommasi (1970): p. 26, fig.18,19; Monteiro (1987): p. 41; Hendler et al. (1995): p. 153, fig. 72, 100J, K; Borges & Amaral (2005): p. 256, fig. a–e; Manso et al. (2008): p. 190, fig. 17 a–c [as Amphiodia pulchella]; Clark (1915): p. 249; Verrill (1899b): p. 313 [as Amphiodia repens]; Lyman (1875): p. 18, pl. III fig. 38–40 [as Amphiura repens]., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Araçá Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on pages 31-33, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Lyman, T. (1869) Preliminary Report on the Ophiuridae and Astrophytidae dredge in deep water between Cuba and the Florida Reef. Bulletin of the Museum of Comparative Zoology, 1, 309 - 354.","Thomas, L. P. (1962) The shallow water amphiurid brittle stars (Echinodermata, Ophiuroidea) of Florida. Bulletin of Marine Science, 12, 623 - 694.","Koehler, R. (1914) A contribution to the study of ophiurans of the United States National Museum. Government Printing Office, Washington, 173 pp.","Hendler, G., Miller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea stars, sea urchins, and allies: echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp. https: // doi. org / 10.1017 / s 0025315400031568","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Springer, Berlin / Heidelberg, 665 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Pomory, C. M. (2007) Key to the common shallow-water brittle stars (Echinodermata: Ophiuroidea) of the Gulf of Mexico and Caribbean Sea. Caribbean Journal of Science, Special Publication, 10, 1 - 42.","Miloslavich, P., Diaz, J. M., Klein, E., Alvarado, J. J., Diaz, C., Gobin, J., Escobar-Briones, E., Cruz-Motta, J. J., Weil, E. & Cortes, J. (2010) Marine Biodiversity in the Caribbean: Regional Estimates and Distribution Patterns. PLoS ONE, 5, e 11916. https: // doi. org / 10.1371 / journal. pone. 0011916","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do Estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9, 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8, 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Manso, C. L. C. & Absalao, R. S. (1988) Ophiuroidea: Situacao Pre Operacional nos sacos de Piraquara, regiao sob influencia da descarga da Central Nuclear Almirante Alvaro Alberto (CNAAA). Revista Brasileira de Biologia, 48, 75 - 82.","Manso, C. L. C. (1989) Os Echinodermata da plataforma continental interna entre Cabo Frio e Saquarema, Rio de Janeiro, Brasil. Revista Brasileira de Biologia, 49, 355 - 359.","Manso, C. L. C. (1993) Ofiuroides da Plataforma Brasileira. Parte II: Norte do Estado do Rio de Janeiro, Estado do Espirito Santo, Sul do Estado da Bahia e Bancos Royal Charlotte, Hostpur e Davis (Echinodermata: Ophiuroidea). Revista Brasileira de Biologia, 35, 189 - 195.","Pires-Vanin, A. M. S., Corbisier, T. N., Arasaki, E. & Moellmann, A. M. (1997) Composicao e distribuicao espaco-temporal da fauna bentica no Canal de Sao Sebastiao. Relatorios Tecnicos do Instituto Oceanografico, 41, 29 - 46.","Netto, L. F., Hadel, V. F. & Tiago, C. G. (2005) Echinodermata from Sao Sebastiao Channel (Sao Paulo, Brazil). Revista de Biologia Tropical, 53, 207 - 218.","Pires-Vanin, A. M. S., Muniz, P. & Bromberg, S. (2014) Inventory of the marine soft bottom macrofauna of Sao Sebastiao Channel, southeastern Brazilian continental shelf. Check List, 10, 795 - 807. https: // doi. org / 10.15560 / 10.4.795","Clark, H. L. (1915) Catalogue of Recent Ophiurans: Based on the Collection of the Museum of Comparative Zoology. Memoirs of the Museum of Comparative Zoology, 25, 165 - 376.","Fell, H. B. (1962) A Revision of the Major Genera of Amphiurid Ophiuroidea. Transactions of the Royal Society of New Zealand Zoology, 2, 1 - 26.","Parslow, R. E. & Clark, A. M. (1963) Ophiuroidea of the Lesser Antilles. Studies on the Fauna of Curacao and others Caribbean Islands, XV, 24 - 50.","Monteiro, A. M. G. (1987) Ophiuroidea (Echinodermata) da regiao de Ubatuba (SP) - Aspectos morfologicos e ecologicos. Phd thesis, Universidade de Sao Paulo, Sao Paulo, 166 pp.","Borges, M. & Amaral, A. C. Z. (2005) Classe Ophiuroidea. In: Amaral, A. C. Z., Rizzo, A. E. & Arruda, E. P. (Eds.), Manual de identificacao dos invertebrados marinhos da regiao sudeste-sul do Brasil. EdUSP, Sao Paulo, pp. 237 - 272.","Verrill, A. E. (1899 b) Revision of certain families and genera of West Indian Ophiurans. Transactions of the Connecticut Academy of Arts and Sciences, 10, 301 - 371. https: // doi. org / 10.5962 / bhl. part. 7032","Lyman, T. (1875) Zoological results of the Hassler Expedition. II. Ophiuridae and Astrophytidae. Illustrated Catalogue of the Museum of Comparative Zoology, 8, 1 - 34."]}