taxonID	type	description	language	source
C1391E19FF943758FC7BFEC0FAC4B79D.taxon	description	Notes: Our phylogenetic analysis supports the monophyly of the Hippasterinae as a discrete subgroup within the Goniasteridae. Several characters support hippasterines as members of the Goniasteridae, including a heavily calcified body wall, abactinal plates arranged into discrete primary-radial series and smaller inter-radial, secondary plates, papulae present radially but absent inter-radially, two series of clearly delineated and heavily calcified marginal plates, actinal plates arranged into distinct chevron-shaped rows as well as suckered tube feet in biserial rows. Our assessment disagrees with Fisher’s (1940) conclusion that the Hippasterinae is ‘ superfluous. ’ Fisher based his conclusion on the ‘ intermediate’ morphological characters observed in Cladaster. Mah (2006) supported Cladaster as the sister group to the Circeaster lineage; however, it is possible that either the Circeaster and Hippasterine lineages are closely related or that Fisher (1940) referred to different Cladaster spp. than those utilized in Mah (2006). In either case, the data presented here support the Hippasterinae as a discrete group that is recognized herein. Diagnosis: Pulpy tissue present. Abactinal plates with spiny-granular or angular accessory fringe. Abactinal plates tightly articulated. Superomarginal and inferomarginal plates, wide to quadrate with large, prominent spines in most taxa. Pedicellariae enlarged, abundant and often on raised base. Marginal plates facing laterally. Disk strongly swollen in most.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF943754FC30FA39FE00B7D6.taxon	materials_examined	Type species: Gilbertaster anacanthus Fisher, 1906. Included species: G. anacanthus Fisher, 1906; G. caribaea (Verrill, 1899). Diagnosis: Arms triangular, broad to tapering, narrow (Fig. 3 A, E). Disk weakly swollen (Fig. 3 A, E). Tissue thick, pulpy covering abactinal, marginal, actinal plates. Abactinal plates low, polygonal covered by one to 12, closely articulated angular granules (Fig. 3 A – C, E – G) elongate to round in length, flattened, forming smooth to rough surface texture. Fasciolar grooves shallow. Secondary plates present between abactinal plates (Fig. 3 C). Abactinal plates with angular accessories (Fig. 3 A – C). Pedicellariae large, bivalved with smooth valves, the length of one to two plates and abundant on abactinal surface (Fig. 3 B, C). Marginal plates, 50 – 70 per inter-radius (armtip to armtip), squarish in outline with rounded edges, completely covered by angular granules similar to those on abactinal surface. Variable surfaces smooth (on G. anacanthus) to roughened (on G. caribaea). Pedicellariae large, bivalve (Fig. 3 C) similar to those on abactinal surface on marginal plate surface, often bisecting the width of the plate. Spines absent from superomarginal and inferomarginal plate series (Fig. 3 A, C, E). Granules, densely arranged, covering superomarginal and inferomarginal plate series complete (Fig. 3 A, B, C, E). Superomarginal and inferomarginal plates quadrate at inter-radius (Fig. 3 C, E). Fasciolar grooves on marginal and actinal surfaces absent. Fringe of accessories on marginal plates poorly differentiated. Superomarginal plates forming prominent dorsolateral fringe (Fig. 3 A, E). Actinal plates covered by one to 15 flattened, polygonal, angular granules. Large bivalve pedicellariae similar to those on abactinal, marginal plate surfaces abundant on actinal plates. Actinal plates with granules, but lacking large spines or spinelets (Fig. 3 D, G). Pedicellariae, bivalved, present on plate series at perpendicular angle (Fig. 3 A – G), adjacent to ambulacral furrow. Pedicellariae, flat-tong shaped with serrated blades present on actinal plates (Fig. 3 D, F). Pedicellariae very abundant, present on raised bases (Fig. 3 A – G). Furrow spines two to four (usually three) blunt, thickened spines, horizontally flattened (G. anacanthus) to triangular / quadrate in cross-section (G. caribaea) (Fig. 3 D, G). Subambulacral spines, one to four, blunt, flattened (Fig. 3 D, G). Round to quadrate (G. anacanthus) to triangular in cross-section (G. caribaea). Pedicellariae, bivalved, enlarged on first adambulacral (similar to others) replacing subambulacral spination (Fig. 3 D, G) and sometimes replacing furrow spination. Subambulacral spines smaller in size, more abundant (Fig. 3 D, G). Furrow spines round in cross-section, not compressed. Oral plates covered by ten to 20 flattened, angular, closely articulated granules (Fig. 3 D, G), sometimes with enlarged bivalve pedicellariae. Oral plate furrow spines, typically five, flattened to oval in cross-section. Oral region concave (Fig. 3 D, G).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF943754FC30FA39FE00B7D6.taxon	description	Fisher, 1906: 1063; A. M. Clark, 1993: 223; H. E. S. Clark & McKnight, 2001: 49 (as Gilbertaster anacanthus) McKnight, 1973: 192; A. M. Clark, 1993: 223 (as Gilbertaster brodiei) Occurrence: Hawaiian Islands, Palau to New Zealand. 277 – 868 m. Material examined: HOLOTYPE: USNM 21168, Malae Point, Hawaii, 20 ° 6 ′ N, 155 ° 59 ′ W, 463 – 699 m, coll. USFC Albatross, 11. vii. 1902 [1 wet spec. R = 6.5, r = 2.2]; CASIZ 159079, off Kona, 19 ° 38 ′ N, 156 ° 2 ′ W, 670.0 m, coll. Sandra Brooke & Michelle Wood on R / V Pisces V, 14. xii. 2001 (1 wet spec. R = 7.3, r = 2.2); CASIZ 159080, Off Kona, 19 ° 38 ′ N, 156 ° 2 ′ W, 868.0 m, coll. Sandra Brooke & Michelle Wood on R / V Pisces V, 14. xii. 2001 (1 wet spec. R = 5.2, r = 2.2); CRECH 129, Mutremdu, Palau, 7 ° 16 ′ N, 134 ° 31 ′ W, 277.3 m (910 ft), coll. P. Colin aboard Deep Worker submersible, 6. iii. 2001 (1 wet spec. R = 9.9, r = 3.2). Description: Arms five. Disk broad, arms long, narrow (R: r = 2.26 – 3.0), distinct from disk. Abactinal plates tightly abutted, covered by granules, one to six, round to irregularly polygonal to oblong to elongate in outline, forming angular fringe around each (Fig. 3 E, F). Carinal series distinct (Fig. 3 F). Granules slightly convex to rounded but low and close to disk surface (Fig. 3 E, F). Periphery of plate surrounded by four to 12 elongate to roundedpolygonal granules that sit in low concavities on each plate. Plates larger proximally but becoming smaller distally at contact with superomarginal border. Madreporite pentagonal, with well-developed sulci. Secondary plates present but few. Pedicellariae bivalve large (~ 1.0 – 1.5 mm in length) and present in irregular cluster all across the abactinal disk surface. Apparently more common on disk but also present in less abundance on distal arm surface. Papulae present radially, absent inter-radially. Marginal plates quadrate in shape, 18 – 22 in number (superomarginals and inferomarginals identical in number and appearance), largely flat but weakly convex and covered with granules, 20 – 70, flush, close-set, convex granules, forming a weakly expressed border around the disk periphery. Marginal plate surface more flattened inter-radially becoming more rounded and more convex distally. Granular covering angular, crowded but other major ornamentation (i. e. spines, etc.) absent from marginal plate surface. Granules form weakly differentiated periphery around marginal plate surface. Marginal fascioles absent (Fig. 3 E, F). Superomarginal plate surface with dorsol facing (Fig. 3 E), especially on distal arm segment. Pedicellariae, bivalved, uncommonly present on marginal plate surface. Actinal plates, very tightly articulated, quadrate to rounded in shape, forming three distinct chevrons on actinal intermediate surface (Fig. 3 G). Granules present, four to 16 in number, closely abutted and polygonal-oblong to irregular, rounded with low convex appearance in shape (Fig. 3 G). Granular cover on actinals flush with those on adjacent inferomarginal plate series. Actinal plate series adjacent to adambulacral plate series each with one enlarged bivalved pedicellariae, decreasing in size proximally to distally, each one flanked by a granular ring, four to 12, enlarged, quadrate (Fig. 3 G). Pedicellariae, bivalve, enlarged (~ 1.0 mm in length), present on several actinal plates, each surrounded by ring of enlarged quadrate granules. Adambulacral plates quadrate. First adambulacral with giant pedicellariae, bivalved, smooth on each inter-radius, flush with furrow margin, extending the whole length of the plate (Fig. 3 G). Furrow spines, two or three, horizontally flattened, oval in cross-section, becoming reduced to a granule distally. Some adambulacral plates with spines, subequal and very small. These latter spines are observed in conjunction with subambulacral pedicellariae. Subambulacral armature varies. Plates covered by granules, four to six, prismatic to quadrate in crosssection in irregular rows (Fig. 3 G). Small granular row present between adambulacrals and proximal actinal plate series. Other plates with large bivalve pedicellariae. Oral plates slightly convex with furrow spines, five to six triangular in cross-section, largest proximally. Oral plate surface covered by granules similar to others, large, polygonal, flattened, five pairs on each plate. Colour in life is dark orange to yellow. Fisher (1906) notes one specimen as ‘ Dull yellow on dorsal surface, brightest on marginal plates, central part of dorsal area with a brownish cast. Actinal surface a pale Naples yellow with a brownish suggestion’. Habitat description: This species has been observed in situ by the Hawaiian Undersea Research Laboratory (HURL) as solitary individuals on soft substratum among round basaltic rocks (C. Kelley, HURL pers. comm., 2009).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF943754FC30FA39FE00B7D6.taxon	description	Verrill, 1899: 174, pl. 28; Halpern, 1970 a: 190; Clark & Downey, 1992: 246 (as Hippasteria caribaea) Occurrence: Known from Cumberland Island, Georgia, Savannah Banks, Stetson Banks, off Jacksonville, FL, and off the coast of West End, Grand Bahama. 500 – 805 m. Material examined: HOLOTYPE: USNM 18425, Cumberland Island, Georgia, North Atlantic Ocean, 30 ° 58 ′ N, 79 ° 38 ′ W, 538 m, coll. USFC Albatross, st. 4041, 5. v. 1886 (1 dry spec. R = 1.8, r = 1.0); USNM 1126236 Jacksonville Lithoherms, 30 ° 31 ′ N, 79 ° 39 ′ W, 553 m, coll. K. J. Sulak, JSL 4683, 10. vi. 2004 (1 dry spec. R = 7.7, r = 3.2); USNM 1124498, Savannah Banks, 31 ° 44 ′ N, 79 ° 05 ′ W, 500 m, coll. S. W. Ross, JSL 4687 12. vi. 2004 (1 wet spec. R = 5.9, r = 2.8); USNM 1126239 Savannah Banks, 31 ° 46 ′ N, 79 ° 12 ′ W, 509 m, coll. C. Morrison, JSL 4905, 30. x. 2005 (1 wet spec. R = 13.3, r = 5.1); USNM 1126238 Stetson Banks, 31 ° 50 ′ N, 77 ° 36 ′ W, 694 m, coll. T. Casazza, JSL 4699, 18. vi. 2004 (1 wet spec. R = 4.3, r = 2.6); MCZ 3806, Florida, off Jacksonville, 796 – 805 m (435 – 440 fathoms), coll. Atlantis St. 3782 (1 dry spec. R = 10.2, r = 3.4). NSU no #. Off coast of West End, Grand Bahama. 27 ° 04 ′ N, 79 ° 19 ′ W, 604 m. JSL II 3698, coll. C. Messing. (1 wet spec. R = 5.0, r = 2.3). Description: Arms five. Disk broad, arms short (R: r = 2.1 – 2.4) distinct from disk (Fig. 3 A). Abactinal surface covered by coarse granules, densely packed with no plate surface visible. Granules forming continuous cover, nearly contiguous with superomarginal plate series (Fig. 3 A, B). Plates, each with one (exceptionally two or more) large round, tubercular granule (s), surrounded by three to six smaller rounded coarse granules (Fig. 3 C). Abactinal plates closely abutting (Fig. 3 B) in adult specimens, somewhat less so in smaller individuals (when R = 4 – 5 cm). Papulae single, distributed over most of abactinal surface but absent from narrow triangular area adjacent to contact with superomarginal plate series. Madreporite round with well-developed sulcae, surrounded by 12 – 15 plates. Pedicellariae large (0.8 – 2.0 mm in length), bivalve equivalent to three to six granules in relative length evenly distributed over abactinal surface (Fig. 3 B, C). Marginal plates elongate [length (L)> width (W)], largest inter-radially, becoming more equal in size and smaller distally. Marginal plates 40 – 50 per interradius (counted from armtip to armtip) covered by densely packed coarse granules (20 – 60 per plate) similar to those on abactinal surface. Granules more evenly spaced, less dense in smaller individuals (R = 4 – 5 cm). Plate surface not visible on inter-radial plates with smooth, bare patches present near arm terminus. Granule shape varies from round, hemispherical to polygonal to more oblong (Fig. 3 C). Superomarginal granules prominent, strongly convex, not forming even surface with other granules (Fig. 3 C). Granules on inferomarginal plates more polygonal, forming close pavement. Pedicellariae absent from marginals on paratype (smaller specimen) but present on holotype (larger specimen) where they bisect the entire width of superomarginal and inferomarginal plates. Distinct groove present around inferomarginal plate contact with actinal intermediate plate surface. Actinal plates forming irregular chevron-like pattern. Actinal plates adjacent to adambulacral plates all with enlarged bivalved pedicellariae equal to length of plate on which it sits. Approximately six to seven granules flank each valve of these pedicellariae. Largest centrally becoming smallest near the ends of each pedicellariae. Other actinal intermediate plate chevron series more irregular with approximately 60 % of plates bearing a large bivalve pedicellariae (Fig. 3 D). Holotype with nearly all plates bearing a large bivalve pedicellariae. Remaining plates covered by four to 14 (mean of nine) granules. One enlarged, convex granule, flanked by four to seven smaller polygonal granules, varying in size. Pedicellariae present closest to mouth and tube foot furrows becoming almost completely absent on plates adjacent to inferomarginal plate series. Furrow spines, thick, triangular to round-oblong rectangle in cross-section, two to three per adambulacral plate with fewer spines on distal plates (Fig. 3 D). One large thick (3 – 4 ¥ thickness of furrow spines), angular, subambulacral spine, present immediately behind furrow spines. This large subambulacral spine flanked by one to two smaller spines, triangular in cross-section, roughly half the height of the large subambulacral (Fig. 3 D). Several thickened, blunt granules, polygonal-triagonal in crosssection, five to nine adjacent to the subambulacral spine, similar in size to those on actinal surface. Typically, one enlarged, round granule present adjacent to subambulacrals with other granules irregularly trailing off in size. Distinct groove between adambulacral plates and first adjacent actinal intermediate plate (which bears the large bivalve pedicellariae). Enlarged triangular spines two, present on oral plate surface projecting into mouth, adjacent to four thick, polygonal spines on the side of each oral plate (Fig. 3 D). Oral plate covered by two enlarged subambulacral spines, round to triangular in cross-section, three to four times as thick as the adjacent furrow spines. Smaller, lower polygonal granules, five to seven present on oral plate surface adjacent to enlarged pedicellariae adjacent to the mouth. Colour in life is yellow-orange. Biology: Hippasteria (= Gilbertaster) caribaea was measured for reflectance in a bioluminescence study presented by Johnson (2005). Habitat description: Specimens provided by M. Nizinski were observed as solitary individuals collected on soft substrata. No other organisms were observed at the collection site.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF983754FEB8FA18FE46B40A.taxon	description	Description: As for species.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF983756FED2F9C7FD7CB197.taxon	description	FIGURE 4 A – H Occurrence: Known only from Savannah Banks and off the coast of Jacksonville, FL. 252 – 501 m. Material examined: HOLOTYPE: USNM 1124468, Savannah Banks, 31 ° 42 ′ N, 79 ° 07 ′ W, 498 m, coll. T. Casazza, JSL 4902, 26. x. 2005 (1 wet spec. R = 9.6, r = 4.7). PARATYPE-USNM E 15539, off coast of Jacksonville, FL, North Atlantic Ocean. 30 ° 31 ′ N, 80 ° 05 ′ W, 252 m (1 dry spec. R = 9.3, r = 4.4); PARATYPE- USNM 1124469, Savannah Banks, 31 ° 44 ′ N, 79 ° 05 ′ W, 501 m, coll. C. Caddigan, JSL 4900, 22. x. 2005 (1 dry spec. R = ~ 9.2, r = 4.8). Etymology: Genus is derived from sthenos, Greek for ‘ strength’ and – aster for ‘ star. ’ Specific epithet is named after Dr Emma Bullock in honour of her contributions to the geochemistry of meteorites and asteroids. Description: Arms five, triangular in outline. Disk large, broad, swollen, especially in wet specimen. Body stout, thick (Fig. 4 A, B). Abactinal plates round to polygonal to oblong in outline with numerous interspersed secondary plates (Fig. 4 F) embedded in a thick, pulpy tissue forming heavily thickened abactinal body wall (Fig. 4 A). Plates with high-aspect; mound-like in shape, forming deep but open channels between plates (Fig. 4 A, E). Abactinal plates are covered with two to ten blunt to conical spine-like granules, usually forming a periphery around plate edge with only one or two granules / spines centrally located. Some plates bear enlarged conical, spine-like granules, surrounded by smaller blunt spinelets. Large abactinal clam-shell like pedicellariae present, each with nine to 12 interlocking teeth per valve (Fig. 4 E). Pedicellariae located centrally on plate surface varying in size from one-third to three-quarters of plate diameter, flanked by spinelike granules. Pedicellariae occurring unevenly over abactinal surface, becoming densely concentrated in some areas but absent from others. Madreporite sunken, bordered by seven to eight abactinal plates. Papulae, small, four to six, interspersed between plates. Marginal plates wide (W> L), 42 – 45 per interradius (from armtip to armtip), which become smaller and with more equivocal dimensions distally adjacent to terminal. Large spines absent from marginal plates series. Fasciolar channels present between marginal plates, relatively deep (Fig. 4 H). Superomarginals facing laterally, correspond 1: 1 to inferomarginals along most of series with one to two plates irregularly offset, possibly because of sublethal predation. Superomarginal plate surface convex, bare except for 20 – 35 widely spaced, sharp, conical spinelets, which are most densely concentrated ventrally on plate at contact with inferomarginal plate (Fig. 4 H). One to two relatively small clam-shell like pedicellariae with interlocking teeth present on nearly every marginal plate but regularly present, adjacent to inferomarginal contact, irregularly present closer to contact with abactinal surface. Periphery of each plate with 15 – 40 small evenly spaced conical spinelets, which occur more densely at contact with inferomarginals. Inferomarginal plates with ventral facing and more densely covered by ten to 70 irregularly sized conical to blunt spinelets. Higher number of larger, conical, more pointed spinelets present closer to superomarginal contact. Higher numbers of spinelets present inter-radially decreasing distally corresponding with smaller inferomarginal plate size. Pedicellariae, one to four, typically two, identical to the type on superomarginals present on inferomarginals facing ventrally. Terminal plate round, bulbous. Actinal intermediate plates, similar in size, shape to abactinal plates with well-developed fasciolar channels running between plates (Fig. 4 C, D). Actinal plate series adjacent to adambulacral plate series with large pedicellariae nearly equal to size of plate on which it sits (Fig. 4 C, D), teeth poorly developed to absent relative to those on abactinal plates. Pedicellariae on actinal series adjacent to adambulacral plates occur with less frequency distally along the arm with some smaller actinal plates adjacent to armtip with pedicellariae absent, bearing only three to six spinelets. Actinal pedicellariae occur most heavily adjacent to mouth, becoming less common to absent adjacent to inferomarginal plate contact. Pedicellariae on each actinal plate surrounded by 20 – 35 sharp spinelets, some round, some triagonal to polygonal in cross-section (Fig. 4 C, D). Actinal plates adjacent to inferomarginal plate series covered with four to 30 sharp, irregularly sized, conical granular spines. Some with polygonal to triangular in crosssection. Oral cavity sunken (Fig. 4 B, C). Adambulacral furrow spines blunt tipped, three to four per plate, triangular to oblong ovate in crosssection in weakly convex series (Fig. 4 G). Distinct grooves between adambulacral plates. Central spines longest and thickest with spines shortest on ends. Subambulacral spines three in two series. Subambulacral spine series adjacent to furrow spines somewhat shorter than furrow spines but comparable in thickness, round to oval in crosssection (Fig. 4 G). Subambulacral spine series farthest away from furrow spines shortest, with central spines only slightly less thick than furrow spines with smallest spines located on ends of second subambulacral series. Distinct fasciolar channel separates adambulacral plates from actinal intermediate plates. Oral plates with four to six furrow spines. Oral plate surface with three to four spines per plate (six to eight total on paired oral plates) with one to three spines projecting into oral opening (Fig. 4 C). Spines triangular to flattened triangular in cross-section, most are comparable in length to furrow spines but sometimes with one or two that are similar to granular spines. Colour in life is orange. Habitat description: The holotype was collected on hard substrata covered by the gorgonian Eunicella modesta (Verrill, 1883). Other unidentified gorgonians, the scleractinian Lophelia pertusa, sponges, and coral rubble were observed at the collection site. The paratype was collected from Savannah Banks on hard substrata where various gorgonians, sponges, scleractinian corals, and coral rubble were also present.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF9A3750FEEFFC39FC93B7D6.taxon	description	Codoceo & Andrade, 1981: 379 (as Criptopeltaster) Type specimen: NEOTYPE: USNM E 33356, south of Santa Cruz Island, Channel Islands, California, 33 ° 55 ′ 30 ′ N, 119 ° 41 ′ 30 ′ W, 486 m, coll. USFC Albatross, 7. ii. 1889. Included species: Cryptopeltaster lepidonotus Fisher, 1905 (Cryptopeltaster philipii is now a synonym of C. lepidonotus). Diagnosis, distribution, and characters: As per species.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF9A3750FEEFFC39FC93B7D6.taxon	description	Ludwig, 1905: 138 (as Hippasteria pacifica) Fisher, 1905: 311; 1911: 237; Lambert, 1978 a: 9; Maluf, 1988: 34, 118; Clark, 1992: 251 (as C. lepidonotus) Codoceo & Andrade, 1981: 379 (as C. philippii) Pawson & Ahearn, 2001: 42 (as Cryptopeltaster cf. lepidonotus) Occurrence: Chile to Aleutian Islands (Alaska), including records from Rodriguez Seamount, Santa Cruz, California and British Columbia 188 – 1244 m. Material examined: CASIZ 108628, Monterey, CA off Point Sur, 914.0 m (500 fms), coll. M. Eric Anderson, 7. vi. 1977 (1 wet spec. R = 4.4, r = 1.7); CASIZ 11828, Oregon, off the coast, 47 ° 15 ′ N, 124 ° 53 ′ W, 188 – 216 m, coll. Roger N. Clark aboard R / V Miller Freeman, 22. x. 1996 (1 dry spec. R = 13.1, r = 5.2); USNM 1129943, Rodriguez Seamount, 34 ° 2 ′ N, 121 ° 4 ′ W, 667.3 m, coll. D. Clague, on board ROV Tiburon, 29. iv. 2004 (1 wet spec. R = 9.6, r = 6.1); USNM E 47396, Washington, north-west of Grays Harbor, 47 ° 10 ′ N, 124 ° 57 ′ W, 195 – 242 m, coll. R. N. Clark on board R / V Miller Freeman, 22. x. 1996 (2 dry specs. R = 9.8, r = 3.8; R = 10.8, r = 4.8); USNM E 51296, North of Seymour Island, Galapagos Islands, 00 ° 21 ′ S, 90 ° 15 ′ W, 599 m (1964 ft), coll. C. Baldwin & J. McCosker, Johnson Sea Link II, 26. vii. 1998 (1 wet spec. R = 8.1, r = 3.5). Description: R: r = 2.3 – 2.6, arms triangular, disk broad. Abactinal surface covered by large, coarse, flat, angular granules, densely abutting around spines and pedicellariae. Abactinal plates largest proximally becoming smaller distally adjacent to contact with superomarginal plate series. Spines conical, present, large, numerous on abactinal surface with granules forming flattened, angular skirt around each spine base. Spines or pointed granules present on nearly every abactinal plate, especially those on radial regions, but are nearly absent distally on regions adjacent to superomarginal series. Pedicellariae large (length equivalent to about seven to nine granules), bivalved. Secondary plates present sometimes covered by granules, one or two, similar to others. Marginal plates, 40 – 55 per inter-radius (from terminal to terminal), each covered with granules, densely arranged polygonal, quadrate to angular in shape. Number of marginal plates increases as adult size increases. Granules number 20 – 30 around each marginal plate periphery forming convex contact with abactinal and actinal surfaces. Granules on central marginal plate surfaces number 20 – 40. Granules, smooth, angular in outline, flattened to convex and often with a pointed tip, distributed evenly throughout surface. Spines, one to three (typically one), short, conical to tubercular present at lower end of each superomarginal plate adjacent to contact with inferomarginal plates. Spines present on inferomarginals inter-radially, becoming lower and more tubercular distally along arms. Granules becoming more flush with others distally on arms. Actinal intermediate areas covered with similar flattened, closely abutting, angular granules, almost all with spines or tubercles. Spines, conical single and prominent, on each actinal intermediate plate number highest proximal to mouth. Spines, smaller and disappearing distally (adjacent to inferomarginal plate contact). Large bivalve pedicellariae (length about 3.0 mm each) in a distinct linear series adjacent to adambulacral plate series, each surrounded by 11 – 15 angular granules. Approximately five to seven chevrons of actinal plates per inter-radius. Adambulacral plates primarily occupied by two to three (primarily three) furrow spines per plate, but a large bivalve or trivalve pedicellariae will replace these spines on the first postoral adambulacral plate and irregularly if infrequently on the furrow spines. Furrow spines thick, club-shaped and round in cross-section. Each paired oral plate with six to eight angular granules along median axis but with four to six granules covering remaining oral plate surface. Furrow spines, three to four on each oral plate. Spine, thickened, oblong in cross-section on the surface of each oral plate facing into the mouth. Holotype: The original holotype for this species has been lost (C. G. Ahearn, pers. comm., 2007). A neotype (USNM 33356) from part of Fisher’s original voucher series, collected near to the original type locality is herein designated as its replacement. Codoceo & Andrade (1981) were the last authors to refer to the holotype of C. lepidonotus. Synonymy of C. philippii: A new Chilean species, Cryptopeltaster philippii was described by Codoceo & Andrade (1981) who distinguished C. philippii from C. lepidonotus on the basis of fewer pedicellariae on the body surface, an undivided madreporite, and fewer supero- and inferomarginal plates per interradius. Cryptopeltaster from the Galapagos (USNM E 51296) corresponds to this description (Pawson & Ahearn, 2001). These characters fail to differentiate between these two species and strongly support the synonymy of C. philippii into C. lepidonotus. Based on the greater number of specimens available, it is determined that the characters vary across the range of the genus and amongst differently sized individuals. Pedicellariae number is variable across different individuals and does little to differentiate between any two specimens. The madreporite was atypically divided by a seam in the holotype (Fisher, 1911: pl. 47, fig. 1). Other specimens clearly show this to be unique to that specimen making this character individually variable and unhelpful as a diagnostic character. Finally, the number of marginal plates in Cryptopeltaster increases in larger specimens. The Galapagos specimen (USNM E 51296) has approximately 42 marginal plates but is smaller (R = 8.1 cm) than specimens collected farther north. A small (R = 4.4 cm) specimen from off Point Sur, California, which otherwise corresponds to the description of C. lepidonotus, also had 42 marginal plates. Hippasteria pacifica Ludwig, 1905 from Mexico was synonymized with C. lepidonotus by Fisher (1911) and was represented by a smaller specimen (R = 4.8 cm) with approximately 40 marginal plates. Codoceo & Andrade (1981) did not include the size of the holotype, which is now apparently lost or unavailable (requests for material from the Museo Nacional de Historia Nautral in Santiago, Chile have gone unanswered). ‘ Cryptopeltaster lepidonotus ’ is misidentified in Imaoka et al. (1991). The species pictured in their monograph features the oval marginal plates characteristic of H. californica and other ‘ Nehippasteria ’ type hippasterines but absent in Cryptopeltaster. The polygonal granules, enlarged pedicellariae, and furrow spine replacement by pedicellariae are also absent from the specimen figured in their monograph.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF9C3750FECEFA02FC68B136.taxon	materials_examined	Type species: Evoplosoma forcipifera Fisher, 1906 Diagnosis: Body strongly swollen, arm narrow. Tissue with pulpy texture covers abactinal plates (seen more clearly in wet specimens). Abactinal fasciolar grooves shallow. Secondary plates present. Abactinal plates, flat and platform-like. Carinal series poorly distinguished. Abactinal plates tightly articulated. Large spines or spinelets on abactinal, superomarginal, inferomarginal plate, and actinal series. Accessories on superomarginal and inferomarginal surface widely spaced. Superomarginal plate series with lateral facing. Superomarginal and inferomarginal plates at interradius quadrate in shape. Spinelets present on both marginal plate series. Fasciolar grooves between marginal plate series shallow. Accessories around superomarginal and inferomarginal plate edge poorly differentiated. Actinal fasciolar grooves shallow. Actinal spinelets present. Furrow spines compressed, angular in cross-section, pedicellariae with serrated valves. Included species: E. augusti Koehler, 1909; Evoplosoma claguei sp. nov.; E. forcipifera Fisher, 1906; E. scorpio Downey, 1982; Evoplosoma timorensis Aziz & Jangoux, 1985 a; E. virgo Downey, 1982; Evoplosoma voratus sp. nov.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF9C3751FC59FCA0FB54B647.taxon	description	FIGURE 7 A – F Occurrence: Known from CoAxial Cone and Rodriguez Seamount and off Islas Tres Marias in the North Pacific. 730 – 2405.6 m. Material examined: HOLOTYPE: USNM 1124507. Rodriguez Seamount 33 ° 57 ′ N, 121 ° 8 ′ W, 1842.8 m, coll. ROV Tiburon, Sta. T- 629, A 8, 14. x. 2003 (1 wet spec. R = 9.9, r = 2.6 cm). PARATYPES: SIO E 2440, Islas Tres Marias, Mexico. 21 ° 52 ′ N, 106 ° 12 ′ W, 730 m, coll. R. Wisher, 18. v. 1959 (1 wet spec. R = 8.3, r = 2.6). USNM 1136366 CoAxial Cone Seamount, 46 ° 30 ′ N, - 129 ° 35 ′, 2405.6 m. Coll. ROV Doc Ricketts, Sta. D 77 - A 3. 30 Aug 2009 (1 wet spec. R = 11.1, r = 2.7 cm). Etymology: This species is named after Dr David Clague, MBARI geologist and primary investigator of the cruise on which this species was collected. Description: Disk pentagonal, with wide inter-radial almost straight lateral sides between arms. Arms elongate and triangular in outline, distinctly set off from the disk appearing to be attached at the disk corners. Abactinal surface inflated, swollen in life. Abactinal plates rounded polygonal to completely round surrounded by 15 – 28 small blunt angular accessory granules forming plate periphery. Each plate with one prominent accessory that varies in shape from spherical tubercle to short, pointed granule to large conical (bullet-shaped) spine. Smaller specimens (R = 8.3) lack well-developed spines but have several pointed, bullet-shaped tubercles. Largest spines are about 2.0 mm in length and appear consistent in size across the abactinal surface. Plates bearing large spines, especially those on the arms, are typically bare aside from accessory granules, but smaller plates may be completely covered by smaller accessory granules. Pulpy membrane present over surface, obscuring granulation and plate surface near periphery of disk at contact with marginal plates. Madreporite convex, swollen with shallow sulci present on surface. Abactinal surface on arms is sharply distinct from plates on disk (Fig. 7 C). Abactinal plates larger, approximately six plates across from superomarginal to superomarginal, narrowing to one to two plates at distal end of arm. Each plate with a large, pointed conical spine per plate. Spine-bearing plates are bare except for periphery of 20 – 30 pointed granules on each plate (Fig. 7 C). Spaces between abactinal arm plates covered by flattened, quadrate granules. Pulpy membrane present but not as strongly expressed and difficult to ascertain on dry specimens. Marginal plate series face laterally, 58 – 60 plates per inter-radius (armtip to armtip) in the larger specimen (R = 9.9) and 50 – 54 plates in the smaller specimen (R = 8.3). Quadrate with rounded edges, plates wider distally (W> L) becoming more equivocal [L = W] inter-radially. Plates slightly convex, bare except for spines, and one to four tubercular granules present infrequently. Superomarginals slightly offset relative to inferomarginals. One to four large, conical to chiselshaped spines per plate. Spines (three to four) and tubercles (one to five) present with higher numbers inter-radially. Spine number decreasing distally to one to two spines distally. More weakly developed spines on smaller specimen, with one to five bullet-shaped granules / tubercles present on plate surface. Single spines on inferomarginal plates form a distinct linear fringe. Pointed granules, 55 – 65 total, form periphery on marginal plates with approximately ten per side in contact with other marginal plates, approximately 15 – 18 per contact with abactinal surface. Terminal plate smooth but with three conical spines, two on distalmost tip, one on abactinal surface. Actinal plate, chevron-like pattern is irregular with actinal plate series adjacent to adambulacral series very elongate, approximately three to four times the length of those in the centre of the actinal intermediate areas and angular in shape. Periphery of these plates covered by 13 – 40 quadrate granules, approximately 20 per side. Approximately 24 per inter-radius with 12 per side. Actinal plates restricted to disk, do not extend onto arms. Remaining actinal plates on disk approximately 25 per inter-radius, circular to irregular in shape, and size but becoming smaller adjacent to contact with inferomarginal plate series. Actinal plates with one to four large, conical spines and / or short, tubercular granules one each plate. Accessory number varies with plate size. Actinals plates elongate with up to four spines and / or granules whereas smaller, circular plates with single spine. Adambulacral plates elongate. Furrow spines four to six. Six proximally and decreasing in number distally. Nine to ten furrow spines on first adambulacral plate. Furrow spines flattened, paddle like to triangular in cross-section many with roughened, worn down ragged tips. Degree of wear varies from spine to spine but seems more pronounced on proximal spines. Subambulacral ornament composed of a single large clam-shell to paddle-shaped pedicellariae proximal on the adambulacral and an extremely thickened, large spine, many with pronounced club-like to almost lobate head with worn tip sometimes with pronounced striations. Ambulacral and subambulacral series flanked by ten to 15 round, hemispherical granules varying in size. Largest adjacent to subambulacrals, becoming smaller and flatter adjacent to actinal inter-radial regions. Oral plates with four thick oral spines (quadrate in cross-section) projecting into mouth (two per plate) and four to five on surface of oral plate at apex of inter-radius. These latter spines are thick and round to quadrate in cross-section. Spines have worn tips with striations. Region between the oral plates and remainder of the actinal intermediate plates is covered by flattened, round granules, densely packed, similar to those adjacent to the subambulacral plates. Colour in life was orange-reddish.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF9D3752FC6EFB73FACCB1E3.taxon	description	FIGURE 8 A – E Occurrence: Known from Davidson Seamount, 2669.9 m. Material examined: HOLOTYPE: USNM 1124506, Davidson Seamount, 35 ° 37 ′ N, 122 ° 49 ′ W, 2669.9 m, coll. J. Barry, L. Lundsten, Sta. T- 947 - A 10, 2. ii. 2006 (1 wet spec. R = 8.4, r = 2.7). Etymology: The epithet for this species refers to the Latin voratus which means to ‘ greedily devour’ in reference to its observed feeding habits on deep-sea coral. Description: Arms tapering, slender but gradually extending from disk. Disk, swollen more pentagonal with relatively lateral to curved inter-radial arcs. Abactinal surface with weakly developed pulpy surface with abactinal surface on arms appearing more pulpy. Abactinal plates round to polygonal becoming smaller and less round distally near contact with superomarginal plate series but larger distally, especially on arms. One to four plates across distance of arm between superomarginal plate series. Abactinal plate surfaces mostly bare with short, stubby conical tubercles. Plates with tubercles not very abundant, only about four to six per arm base on disk. Each plate with ten to 30 peripheral rounded granules that are small and rounded to quadrate in crosssection. Region adjacent to madreporite (about 0.75 ¥ 1.0 cm area) covered with densely packed, large, round granules about twice the size of granules on the abactinal surface and embedded in the pulpy tissue present on the body wall. Madreporite is ovalshaped with deeply etched sulci. Peripheral granules surround madreporite with some granules two to three times as large as others. One plate with two to four large granules on one end of madreporite. Paddle-shaped pedicellariae uncommonly scattered over surface, approximately one or two for every five to seven plates. Marginal plates, bare with a single large, conical spine on most plates, arranged in linear series becoming smaller to absent on distalmost six or nine superomarginal plates. Several spines, especially along the arms, with four to six short, spiny granules at the base of each spine that are absent distally. Spines continue along all inferomarginal plates up to practically the terminal plate. Both marginal plate series covered by skin, which is difficult to observe in dry specimens. Pedicellariae small, tong-like present irregularly on superomarginals and inferomarginals, especially inter-radially. Peripheral granules rounded, 40 – 60 per plate, including ~ ten per side adjacent to other marginal plate and ~ 25 per side adjacent to abactinal and actinal surface. Superomarginal and inferomarginal offset from one another and each series with different overall dimensions. Superomarginals number 44 – 45 from armtip to armtip, more elongate (L> W), becoming almost ovalate. Inferomarginals number 54 – 55 from armtip to armtip, more quadrate (L = W) overall. Superomarginal plate contact with abactinal surface, strongly convex, ovallike especially along the base and along the arm. Inferomarginal with more gently curved contact with actinal intermediate plates. Actinal intermediate plates in approximately three to four chevron-like patterns but in very jumbled, irregular, unclear series present only on disk, ending at arm base. Plates quadrate to irregular in shape and bare with no spination or surface accessories with eight to 25 peripheral granules save for irregularly occurring paddle-like pedicellariae. Pedicellariae, one to six, present in actinal intermediate region. Adambulacral plates elongate, with typically one (exceptionally two) large paddle-like to rectangular pedicellariae sitting on plate always closest to mouth adjacent to enlarged, thickened subambulacral spine that sits distal to mouth. Approximately six to eight large granules on periphery of adambulacral plates in contact with actinal intermediate plates. Subambulacral spines up to two to three times the thickness of furrow spines with blunt to conical tips. Pedicellariae begin to occur on first adambulacral, becoming less regular on distal adambulacral plates. Furrow spines, compressed, quadrate to oval in cross-section, four to six in number, five on average, with middle spines tallest, shortest spines on the ends. Furrow spine tips darkened brown sometimes roughened and jagged. First adambulacral with six to seven furrow spines, significantly thicker with more jagged tips than other furrow spines. Oral region concave with approximately 13 – 15 compressed furrow spines on each oral plate. Furrow spines, flattened and blade-like, one to two present at plate apex, the largest of which pairs off with its twin on the other oral plate and forms a closely articulated array of spines over the mouth. Oral plate surfaces appear bare save for soft, pulpy tissue covering approximately eight to 12 plates through which two to four short granules emerge. Colour in life of the holotype was deep-brick red, which remained almost two weeks following preservation in 75 % ethanol. Observed in situ apparently feeding on Trissopathes pseudotristicha (Cladopathidae, Antipatharia).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF9E3752FC7AFBC6FAF1B780.taxon	description	Occurrence: Indian Ocean-Laccadive Sea (6 ° 31 ′ N, 79 ° 38 ′ E). 733 m (401 fms). Comments: The type specimen of this species could not be located at the Museum national d’Historie naturelle, Paris, France and inquiries to the Calcutta Museum went unanswered. It is presumed lost.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF9E3753FC77FA25FC46B57B.taxon	description	Occurrence: Hawaiian Islands, east of Kauai Island, 48 ° S 15 ′ W. 929 – 1247 m (682 – 508 fms). Comments: The specimen image of E. forcipifera from Trego (2008) lacks abactinal spination and shows quadrate marginals abutting along the midradius, a character absent from Evoplosoma, indicating that the specimen described in this paper is misidentified, invalidating Trego’s (2008) range extension for this species. The holotype for this species is missing from the USNM collections (Ahearn, 1995).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF9E3753FC77FA25FC46B57B.taxon	description	Downey, 1981: 561; Gage et al., 1983: 280; Clark & Downey, 1992: 242; Clark, 1993: 253. Occurrence: South-west Rockall Trough to northeastern European Basin (48.5 ° N, 10 ° W) extended herein to off Delaware 38 ° 45 ′ N, 72 ° 40 ′ W. 1600 – 2105 m. Material examined: USNM E 50539, off Delaware, North Atlantic Ocean, 38 ° 45 ′ N, 72 ° 40 ′ W, 2105 m, coll. Lamont-Doherty Geological Observatory, DSRV Alvin, 15. vii. 1981 (1 dry spec. R = 5.5, r = 1.8); USNM E 23623, off Ireland, North Atlantic, 55 ° 12 ′ N, 15 ° 50 ′ W, 1900 m, coll. J. Gage, R / V Challenger (1 dry spec. R = 8.2, r = 2.4).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF80374DFF40F980FD23B4DE.taxon	description	Occurrence: East Timor Region, Malaysia, 8 ° 50.2 ’ S, 127 ° 2 ′ E. 883 m. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF80374DFF40F980FD23B4DE.taxon	description	Downey, 1982: 772; Clark, 1993: 253 Occurrence: Gulf of Mexico, 2056 m. Material examined: HOLOTYPE: USNM E 24285, Gulf of Mexico, North Atlantic Ocean, 26 ° 08 ′ N, 92 ° 43 ′ W, 2056 m (1124 fms), coll. W. Pequegnat, 30. vii. 1971 (1 dry spec. R = 11.5 cm, r = 3.3 cm).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF81374DFEA0F8EDFC00B57B.taxon	description	Occurrence: Off Java (Indonesia). 2186 m. Comments: This species was unidentified in Fujikura et al. (2008) but can be placed within the genus Evoplosoma based on its arm and disk shape as well as apparent surface spination.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF82374FFEB2FB62FE64B4E4.taxon	materials_examined	Type species: Asterias phrygiana Parelius, 1768 (with synonym Hippasteria europaea Gray, 1840 as well as all four nominal species included by Gray as synonyms with H. phrygiana). Diagnosis: Disk strongly swollen, Arms relatively broad and short. Body often pentagonal to weakly stellate. Tissue with pulpy texture covers abactinal plates. Shallow fasciolar grooves present. Secondary plates present. Abactinal plates, tightly articulated, polygonal to irregular in outline, flat and platform-like in shape. Carinal series are poorly distinguished. Abactinal spinelets (sometimes granular) forming fringe around abactinal plates. Spines, large, conical, granules often present on abactinal plates. Large spines present on superomarginal and inferomarginal plates of most taxa. Superomarginal and inferomarginal plates bare, quadrate to rounded in outline at inter-radii with no other accessories other than large spines. Spinelets present on marginal plates. Shallow fasciolar grooves present between marginal plates. Marginal accessories (granules, spinelets, etc.) differentiated into a fringe on superomarginal and inferomarginal plates. Superomarginal plates dorsal-facing in most taxa, Actinal fasciolar grooves shallow. Large actinal spines and spinelets present. Subambulacral spines large (and thus few in number). Furrow spines large, blunt, and round in cross-section, usually few in number. Enlarged bivalve pedicellariae on raised bases present on body surface. Included species: * (asterisk denotes fossil taxon) Hippasteria antiqua Fell, 1956; H. californica Fisher, 1905; Hippasteria colossa Djakonov, 1950; Hippasteria derjungini Djakonov, 1950; H. falklandica Fisher, 1940; H. heathi Fisher, 1905; Hippasteria imperialis Goto, 1914; Hippasteria kurilensis Fisher, 1911; Hippasteria leiopelta Fisher, 1910; Hippasteria mammifera Djakonov, 1950; Hippasteria nozawai Goto, 1914; Hippasteria pedicellaris Djakonov, 1950; H. phrygiana phrygiana (Parelius, 1768), H. phrygiana argentinensis Bernasconi, 1961; H. phrygiana capensis Mortensen, 1933; H. spinosa Verrill, 1909; H. strongylactis Clark, 1926; H. tasmanica McKnight, 2006. Comments: Hippasteria, the best known genus of the hippasterines, is encountered on shallow, near-shore to deep-sea settings worldwide and includes approximately 19 nominal taxa, many of which are similar in appearance and have overlapping characteristics. Some taxa occur in a continuous distribution over a wide geographical range but show relatively conservative overall morphology. Hippasteria phrygiana, for example, occurs in the North Atlantic off Europe and North America, as well as off South America, South Africa, New Zealand, and in the Southern Ocean (Clark & Downey, 1992; Clark & McKnight, 2001). The morphologically similar H. spinosa was described by Lambert (2000) as having pelagic, lecithotrophic larvae. If it were to be found that other Hippasteria spp. had lecithotrophic larvae, then long distance dispersal could explain the observed broad geographical ranges. Some species, such as H. imperalis and / or H. nozawai may actually represent very distinct taxa from the H. spinosa and H. phrygiana complexes, but are poorly known and require further study. Further issues pertaining to Hippasteria species complexes are included below in the Discussion. Biology: Other ecological observations have included interactions with hagfish, which exploit prey captured, but not consumed, by H. phrygiana (Auster & Barber, 2006) and associations with fish assemblages observed at human-made structures (Love & York, 2005). Hippasteria has been described as the host to a number of commensals and parasites, including polynoid polychaetes, in H. californica (Pettibone, 1969), the ascothoracid crustacean Dendrogaster in H. phrygiana (Hamel & Mercier 1994), and the parasitic chlorophyte Coccomyxa astericola in H. phrygiana (Mortensen & Rosenvinge, 1933). Biochemical compounds have been extracted from H. phrygiana (Burnell, Apsimon & Gilgan, 1986; Levina et al., 2005) and gonad ultrastructure has been studied by Walker (1979).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF833748FC84F936FE48B38A.taxon	description	Occurrence: Bering Sea, 238 – 250 m. Comments: This is another of the possible synonyms that compose the H. spinosa complex in the Aleutian / North Pacific region. Characters that distinguish this species are either variable or plesiomorphic to H. spinosa. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF843748FF2AFE27FE48B111.taxon	description	Occurrence: Okhotsk Sea, off the northeast coast of Sakhalin. 192 m. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF843748FF1FFCBFFE48B6BE.taxon	description	Occurrence: Falkland Islands (= Islas Malvinas), northern Argentina, Marion and Prince Edward Islands region to south of Tasmania, approx. 49 ° S, 150 ° E. 225 – 1148 m. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF843748FEFDFB13FD34B425.taxon	description	Occurrence: Alaska, Gulf of Alaska. 377 – 454 m. Material examined: HOLOTYPE: USNM 22338, Guard Island, Behm Canal, Alexander Archipelago, Alaska, 55 ° N, 131 ° W, 377 – 454 m, coll. USFC Albatross, 9. vii. 1903 (1 dry spec. R = 7.8, r = 3.9).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF843748FF09F988FBFFB2CF.taxon	description	Grigg et al., 1987: 387 (as Hippasteria spinosa) Occurrence: Sagami Bay / Tosa Bay region, Kii Strait, southern Japan, Hawaiian Islands. 245 – 600 m. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF843748FC7CFEEFFBFFB138.taxon	description	Occurrence: Okhotsk Sea, off the southernmost point of Kamchatka and in the vicinity of Simushir Island (Kuriles), Petrel Bank, Aleutian Islands. 165 – 600 m. Comments: This species was distinguished from others in the genus based on long, conical abactinal and marginal spines by Djakonov (1950). This was originally a subspecies designated by Fisher (1911) and later raised to a species by Djakonov (1950). This is also a likely synonym within the H. spinosa complex. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF843748FC79FC9EFBFFB6B6.taxon	description	Fisher, 1910: 553; 1911: 227; Djakonov, 1950: 56 (1968: 47); 1952: 413. Occurrence: Southern Bering Sea, Okhotsk Sea, Tartar Strait, and Aniva Bay off the south-east coast of Kamchatka. 35 – 418 m. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF843748FC82FB24FBFFB483.taxon	description	Occurrence: Okhotsk Sea. 97 m. Comments: This species was described by Djakonov (1950) as a separate species based on extremely swollen and hemispherical marginal and abactinal plates. This character is plesiomorphic to H. spinosa (and to a certain extent, H. phrygiana), which undermines the distinctiveness of this species and further lends support to its synonymy with H. spinosa. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF843749FC55F92BFE24B2CF.taxon	description	Occurrence: Hokkaido, northern Japan. (No depth information available.) Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF85374AFF48FEE8FE57B116.taxon	description	Occurrence: Okhotsk Sea. At ‘ inconsiderable depths and above-zero water temperatures’ according to Djakonov (1968: 46). Comments: Characteristics used to distinguish this species include two to three adambulacral spines, one subambulacral spine with a flattened pedicellariae, and close-set granules on the abactinal plates. However, these characters are also observed in H. spinosa suggesting that this species may also be a synonym or part of the H. spinosa species complex. Material examined: None. HIPPASTERIA PHRYGIANA (PARELIUS, 1768) HIPPASTERIA PHRYGIANA PHRYGIANA (PARELIUS, 1768) FIGURE 6 A – D Linck, 1733: 21 (as Pentaceros planus) Parelius, 1768: 425 [1770: 349] (as Asterias phrygiana) Lamarck, 1816: 555 [non A. equestris Retzius, 1805] (as Asterias equestris) Gray in Johnston, 1836: 146 (as Asterias johnstoni) Gray, 1840: 279, 1866: 9 (as Hippasteria europaea, H. johnstoni, H. cornuta) Gray, 1840: 279; 1866: 9; Perrier, 1875: 270 [1876: 65], Danielssen & Koren, 1881: 268; Sladen, 1883: 159; Perrier, 1888: 764; Sladen, 1889: 341; Koehler, 1909: 88, 1924: 179 (as Hippasteria plana) Forbes, 1841: 125 (as Goniaster equestris) Müller & Troschel, 1842: 52; Duben & Koren, 1846: 246 (as Astrogonium phrygianum) Forbes, 1843: 280 (as Goniaster abbensis) Barrett, 1857: 47 (as Astrogonium aculeatum) Norman, 1865: 128 (as Goniaster phrygiana) Dons, 1937: 17 (as Hippasteria [Euhippasteria] phrygiana and Hippasteria [Nehippasteria] insignis) Perrier, 1891: K 128; A. M. Koehler, 1926: 107; Clark, 1962: 22 (as Hippasteria hyadesi) Verrill, 1874: 413; 1885: 542, Ganong, 1893: 56; Grieg, 1895: 6; Verrill, 1895: 137, 1899: 148; Döderlein, 1900: 218; Hartlaub, 1900: 191; Ludwig, 1900: 457; Whiteaves, 1901: 50; Grieg, 1902: 21; Pearcey, 1902: 308; Simpson, 1903: 40; H. L. Clark, 1905: 1; Grieg, 1905: 4; Nordgaard, 1905: 160, 235; Grieg, 1907: 28, 32; Sussbach & Breckner, 1911: 215; Grieg, 1912: 6; 1913: 115; 1917: 8; 1921: 6; H. L. Clark, 1923: 270; Mortensen, 1927: 88, 1933: 245; Haubold, 1933: 200; A. H. Clark, 1949: 373; Djakonov, 1950 (tr. in 1968): 53; Blacker, 1957: 18, 45; Buchanan, 1966: 25; Wolff, 1968: 82, Walker, 1978: 361; Codoceo & Andrade 1978: 156; Franz, Worley & Merrill, 1981: 406, 415; O’Connor & Tyndall, 1986: 96; Moore et al., 2004: 246 (as Hippasteria phrygiana) Fell, 1958: 11, pl. 1, figs A, G; 1959: 136, fig. 21; 1960: 61, pls. 2, 3; 1962: 33; McKnight, 1967: 300; H. E. S. Clark, 1970: 3; A. M. Clark, 1993: 259; Rowe & Gates, 1995: 65; Koslow & Gowlett-Holmes, 1998: 44 (as Hippasteria trojana) Occurrence: Arctic, North Atlantic: South to Cape Cod in the west, including Bear Seamount. To the Kattegat, northern Scotland, and northernmost Ireland in the east. Norwegian coast south to the Kattegat, south-western part of the Barents Sea, Kola Bay. South Pacific: Chile to Magellan Strait to the Marion and Prince Edward Island region to Chatham Island, east of New Zealand, Campbell Plateau, seamounts off southern Tasmania. Southern Ocean: Lavoisier Island, Antarctic Peninsula. Depth is highly variable: 20 – 1275 m and varies by region. Most records for New Zealand are over 500 m. Comments: Southern hemisphere Hippasteria species have been scrutinized since the early 20 th century (e. g. Koehler, 1926, Clark, 1962) because of their strong morphological similarities with the northern Atlantic Hippasteria phrygiana. The New Zealand – South Pacific H. trojana was synonymized with H. phrygiana by H. E. S. Clark in Clark & McKnight (2001). The sub-Antarctic Hippasteria hyadesi was synonymized with H. plana (= H. phyrgiana) by Koehler (1926) but was thought to have had an incorrectly identified type locality by Clark (1962). Stampanato & Jangoux (2004) and Branch et al. (1993) regarded H. hyadesi as a valid species. However, examination of the specimens in the USNM collections from the South Pacific and Antarctic Peninsula (near the type locality) shows few to no morphological distinctions, suggesting that the original conclusion made by Koehler (1926) was correct. Magellanic-Antarctic forms of Hippasteria phrygiana may eventually be supported as cryptic species distinct from those in the northern hemisphere but based on specimens available for study, external morphology has limited usefulness and other characters will need to be considered. Material examined: North Atlantic: USNM E 46610, Browns Bank, Nova Scotia, 42 ° 34 ′ N, 65 ° 44 ′ W, 93 m, coll. R / V Albatross IV, 15. x. 1965 (1 dry spec. R = 7.3, r = 3.5); USNM E 46606, south-east of Cape Elizabeth, Gulf of Maine, Maine. 43 ° 31 ′ N, 69 ° 49 ′ W, 97 m. Coll. R / V Albatross IV, 14. vii. 1965 (1 dry spec. R = 6.2, r = 3.7). USNM 5236 off Eastern Point, Gloucester Harbor, Massachusetts. 42 ° 30 ′ N, 70 ° 38 ′ W, 78.6 m (43 fms). Coll. R / V Speedwell (1 spec. R = 9.2, r = 5.0). USNM E 46611 Browns Bank, Nova Scotia, Canada. 42 ° 45 ′ N, 65 ° 7 ′ W. 97.0 m. Coll. R / V Albatross IV, 15. x. 1965 (1 dry spec. R = 1.7, r = 1.0). USNM E 46613 western part of Georges Bank, Massachusetts. 41 ° 24 ′ N, 68 ° 25 ′ W, 68 m. Coll. R / V Albatross IV, 28. x. 1965 (1 dry spec. R = 2.2, r = 1.7). South Pacific: USNM E 13586, north-west of Amundsen Sea, Southeast Pacific Basin. 54 ° 49 ′ S, 129 ° 48 ′ W, 549 m. Coll. R / V Eltanin. (3 dry specs. R = 3.5, r = 2.0; R = 8.2, r = 4.6; R = 9.56, r = 5.1). USNM 1121154. Magallanes y Antartica, Chile. 53 ° 39 ′ S, 70 ° 55 ′ W, 82 m. Coll. R / V Hero. (1 dry spec. R = ~ 7.8, r = 4.6). Southern Ocean. USNM E 43921, Lavoisier Island, Biscoe Islands, Antarctic Peninsula. 66 ° 20 ’ S, 67 ° 47 ′ W, 325 m. Coll. J. Tyler, 25. iii. 1959. (1 dry spec. R = 6.7, r = 3.1). USNM 1082740. 67 ° 23 ’ S to 67 ° 24 ’ S, 180 ° 00 ′ W to 179 ° 58 ′ W, 595 – 516 m. Coll. R / V Eltanin. (2 dry specs. R = 9.2, r = 4.0; R = ~ 11.0, r = 5.5).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF86374AFF1DFCB3FE48B650.taxon	description	Occurrence: Northern Argentina. 108 – 162 m. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF86374AFEF8FB75FE48B42A.taxon	description	Mortensen, 1933: 245; A. M. Clark, 1952: 170, 196 (as Hippasteria phrygiana var. capensis) A. M. Clark & Courtman-Stock, 1976: 63; Clark & Downey, 1992: 249 (as Hippasteria phrygiana capensis) Occurrence: South Africa, 310 – 980 m. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF86374AFF3DF97FFBFFB2CF.taxon	description	Occurrence: South Africa. 320 – 980 m. Material examined: None.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF86374AFC60FEEDFBD9B7D0.taxon	description	Occurrence: Southern California, British Columbia, to Bering Sea. Okhotsk Sea off Cape Elizavety, and near the Kurile Islands. 49 – 1170 m. Comments: This species forms an extensive species complex throughout the Aleutian and North Pacific sub-Arctic region similar in several ways to Leptasterias and other wide ranging high-latitude species. This extensive variation is likely to encompass all of the species described by Djakonov (1950 translated in Dyakonov, 1968) from the Okhotsk Sea and adjoining regions, including H. mammifera, H. pedicellaris, H. colossa, and H. kurilensis. These are mentioned here for future discussion but type specimens have not been examined. Material examined: USNM 32470, west of San Nicolas Island, 33 ° 13 ′ N, 120 ° 4 ′ W, 825 m, coll. USFC Albatross, 26. iv. 1911. (1 dry spec. R = 0.9, r = 0.4). USNM 33352 south-west of Cape Flattery, Washington. 48 ° 17 ′ N 124 ° 52 ′ W, 70 m. Coll. USFC Albatross 24. ix. 1888 (1 dry spec. R = 8.2, r = 4.3); USNM 39833 Heceta Bank, Oregon, 43 ° 58 ′ N, 124 ° 36 ′ W, 170 m. Coll. USFC Albatross 1. ix. 1889. (1 dry spec. R = 9.2, r = 4.7). USNM 47600 Umnak Island, Islands of Four Mountains, Aleutian Islands, Alaska. 53 ° 3 ′ N, 169 ° 57 ′ W, 146 m. Coll. R / V Harvester 11. viii. 1980. (1 dry spec. R = 7.1, r = 3.5); USNM E 10504. South-west of mouth of Columbia River, Oregon. 46 ° 8 ′ N 124 ° 30 ′ W, 137 m. Coll. R / V Cobb 14. i. 1964. (1 dry spec. R = 12.8, r = 7.1).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF86374BFC83FA05FE41B361.taxon	description	Occurrence: South Tasman Rise, south of Tasmania, Australia. 935 – 1058 m. Material examined: None. OUTGROUP TAXA: Cladaster analogous. USNM E 38569. Shag Rocks, Scotia Sea. 53 ° 20 ’ S, 42 ° 42 ′ W, 417 – 514 m. Coll. R / V Siedlecki, USARP. 29. xi. 1986. (1 dry spec. R = 3.2, r = 1.8). Mediaster aequalis. USNM 33275. Dakins Cove, Santa Catalina Island, Channel Islands, California. 33 ° 17 ′ N, 118 ° 24 ′ W, 86.0 m. Coll. USFC Albatross, 8. iv. 1897. (1 dry spec. R = 6.6, r = 3.3). Peltaster placenta. USNM E 12587 west of Barbuda, Antigua, Caribbean Sea. 17 ° 38 ′ N, 62 ° 16 ′ W, 329 – 338 m (180 – 185 fms). Coll. R / V Oregon 19. v. 1967. (1 dry spec. R = 7.2, r = 5.4).	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
C1391E19FF883746FC8DF9DEFA0BB12B.taxon	description	OG: Mediaster aequalis Hippasteria californica OG: Cladaster analogus Washington to S. California OG: Peltaster placenta 110 - 2000 m Hippasteria Hippasterias 377 Alaska - 454, m Aleutian spinosa heathi Islands Hippasteria Aleutian Islands to California 10 - 512 m Hippasteria phrygiana North & South Atlantic / Pacific 10 - 860 m Evoplsoma virgo NW Gulf of Mexico, 2060 - 2105 m Evoplsoma NE Davidson Rodriguez Evoplosoma Pacific Seamount, voratus Seamount 730 claguei - 1842 n. sp m & n .. sp. Evoplosoma NE Pacific, 2669 m Evoplsoma scorpio N. Atlantic, Rockall Trough, 1600 - 1900 m Sthenaster emmae n. gen. n. sp. S. Carolina, tropical Atlantic 110 - 2000 m Cryptopeltaster lepidonotus Aleutian Islands to Chile, 360 - 1244 m Gilbertaster caribaea W. Tropical Atlantic 360 - 1244 m Gilbertaster 5 changes Gilbertaster anacanthus Hawaii to New Zealand 462 - 913 m and western North Atlantic involving quantitative surveys of mega- and macroinvertebrates report diversity to be extremely high (Jensen & Frederiksen, 1992; Mortensen et al., 1995, Jonsson et al., 2004, Reed et al., 2005, 2006; Henry & Roberts, 2007; Roberts et al., 2008). These recent surveys, however, often report only a fraction of the diversity associated with these habitats because sampling methods utilized target only a portion (either size- or taxonspecific) of the associated fauna, the interest of the investigators is limited to a few taxonomic groups, or the overall goals of the project are such that not all associated fauna are collected. As a result, many megafaunal taxa are often over-looked during collections or are under-sampled. Asteroids are known from many deep-sea coral study sites but observations of these taxa have lacked taxonomic precision, precluding more precise ecological interpretations. Several papers provide vague entries (unidentified asteroid, Mortensen et al., 1995; Asteroidea, unid. sp., Reed, Weaver & Pomponi, 2006) or have identified individuals only to the generic level (Mortensen et al., 1995; Jonsson et al., 2004). Few report species-level identifications (e. g. Jensen & Frederiksen, 1992; Henry & Roberts, 2007). Much remains to be learned about the megafauna associated with deep-sea coral habitats. Large, prominent animals such as the Asteroidea have been shown to have ecologically complex relationships with cnidarians from shallow marine habitats (e. g. Birkeland, 1974; Birkeland & Lucas, 1990) and seem to be important to those living in deep-sea habitats as well (e. g. Chave & Malahoff, 1998; Krieger & Wing, 2002). The in situ observations of Krieger & Wing (2002) documented Hippasteria as a main predator of deepsea coral. Subsequent observations have now shown that at least one species of every genus included in Hippasterinae (Goniasteridae), except for Gilbertaster, has been reported to feed on deep-sea corals, mainly gorgonians, but also alcyonaceans, antipatharians, and other cnidarian species. Krieger & Wing (2002) reported H. heathi, in addition to other species, as the main predator on the gorgonian Primnoa sp. in the Gulf of Alaska. Hippasteria imperialis has been observed feeding on isidid corals in the Hawaiian Islands, off Kona (C. Mah, unpubl. observ.). Japanese researchers using submersibles have also observed an unidentified species of Evoplosoma feeding on deep-sea coral in Java (Fujikura et al., 2008). In the Pacific, submersibles from the Monterey Bay Aquarium Research Institute have documented several instances of hippasterines feeding on deep-sea cnidarians, including gorgonians (Paragorgia), bamboo corals (Keratoisis and Lepidisis) and sea whips (Halipteris) by Cryptopeltaster, Evoplosoma, and Hippasteria at Rodriguez and Davidson seamounts off the west coast of California. Video observations of Sthenaster show the holotype hunched over colonies of the co-occurring gorgonian Eunicella modesta (Verrill 1883). Gut contents of the holotype of Sthenaster (USNM 1124468) included distinctive spicules belonging to the E. modesta (Verrill, 1883). These observations support the hypothesis of corallivory in Sthenaster, and suggests that dietary preferences would be similar to other known hippasterines. If Sthenaster is indeed a predator on gorgonians, this would be the first account of hippasterine predation on gorgonians in the Atlantic. It is unclear how specific nutritional preferences are within the Hippasterinae. Feeding in the shallowwater Hippasteria suggests a feeding preference for many types of cnidarians but not to the exclusion of other food sources. The north-west Pacific species Hippasteria spinosa has been reported as a predator on the sea pen Ptilosarcus (Mauzey, Birkeland & Dayton, 1968; Birkeland, 1974), the white-plumed sea anemone Metridium sp., the zoanthid Epizoanthus scotinus (Wood, 1958), the tunicate Metandrocarpa sp., the polychaete Nereis sp., and eggs of the nudibranch Armina sp. (Lambert, 2000). Additionally, H. spinosa elicits an escape response in the sea anemone Stomphia sp. (Lambert, 2000). Similar to its northwest Pacific congener, H. phrygiana has been reported to incite swimming behaviour in the Atlantic sea anemone, Stomphia coccinea (Müller, 1776) (Robson, 1961). Hippasteria phrygiana preys upon Metridium senile (Linné, 1761) in Maine (Harris, 1991) and has been reported feeding on ‘ cnidarians’ (Mercier & Hamel, 2008). Stomach contents from H. phrygiana have indicated that echinoderms, polychaetes, molluscs, and sediment are also incorporated in the diet of this species. In contrast, stomach contents from the deep-sea H. californica suggest that it is primarily a sediment / detrital feeder (Carey, 1972). Sediments were also found in the gut of H. spinosa (Birkeland, 1974). Other examples of presumed corallivory in nonhippasterine members of the Goniasteridae include the Hawaiian Calliaster pedicellaris (Fisher, 1906) (Chave & Malahoff, 1998) and Circeaster pullus (Mah, 2006), and the Atlantic Plinthaster dentatus (Perrier, 1884) (Halpern, 1970 a, b) and Tessellaster nobilis (Clark, 1941). This work represents a first step to further our understanding of the major taxonomic concepts within the neglected Hippasterinae. Future efforts would be best directed towards a complete investigation of Hippasteria, which occurs worldwide, is the most speciose of the Hippasterinae, and is one of the most frequently encountered asteroids observed feeding on deep-sea corals. Questions relating to dietary preferences / restrictions correlated to phylogeny could be important particularly with regard to management and conservation of these ecosystems.	en	Mah, Christopher, Nizinski, Martha, Lundsten, Lonny (2010): Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society 160 (2): 266-301, DOI: 10.1111/j.1096-3642.2010.00638.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00638.x
