LINNEAN SOCIETY ZOOLOGICAL Journals _ Linnean Society Zoological Journal of the Linnean Society, 2010, 160, 266?301. With 11 figures Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores, including one new genus and three new species CHRISTOPHER MAH1*, MARTHA NIZINSKI2 and LONNY LUNDSTEN3 ^Smithsonian Institution - Invertebrate Zoology, MRC-163, PO Box 37012 National Museum of Natural History Washington District of Columbia 20013-7012, USA 2National Marine Fisheries Service - National Systematics Lab, MRC-153, PO BOX 37012, National Museum of Natural History, Washington, District of Columbia, USA 20013-7012 ^Monterey Bay Aquarium Research Institute - Video Lab, 7700 Sandholdt Road Moss Landing, CoZi/brmw, 95039 (7SA Received 23 June 2009; accepted for publication 8 October 2009 The Hippasterinae is a subfamily within the Goniasteridae, consisting of five genera and 26 species, which occur in cold-water settings ranging from subtidal to abyssal depths. All known genera were included in a cladistic analysis resulting in two most parsimonious trees, supporting the Hippasterinae as monophyletic. Our review supports Sthenaster emmae gen. et sp. nov. as a new genus and species from the tropical Atlantic and two new Evoplosoma species, Evoplosoma claguei sp. nov. and Evoplosoma voratus sp. nov. from seamounts in the North Pacific. Hippasteria caribaea is reassigned to the genus Gilbertaster, which previously contained a single Pacific species. Our analysis supports Evoplosoma as a derived deep water lineage relative to its continental-shelf, shallow water sister taxa. The genus Hippasteria contains approximately 15 widely distributed, but similar-looking species, which occur in the northern and southern hemispheres. Except for Gilbertaster, at least one species in each genus has been observed or is inferred to prey on deep-sea corals, suggesting that this lineage is important to the conservation of deep-sea coral habitats. The Hippasterinae shares several morphological similarities with Circeaster and Calliaster, suggesting that they may be related. ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301. doi: 10.HH/j.1096-3642.2010.00638.x ADDITIONAL KEYWORDS: Arctic ? deep-sea coral ? deep-sea echinoderm ? Evoplosoma ? feeding biology ? Hippasteria ? onshore-offshore ? phylogeny ? taxonomy. INTRODUCTION Deep-sea or cold-water coral ecosystems have received worldwide attention (e.g. Roberts & Hirsh- field, 2004). Exploration of these unique, productive regions has increased recently, leading to an improved understanding of and appreciation for these structure-forming habitats along continental slopes and other deep-sea settings (Hourigan etal., 2007). Corresponding author. E-mail: mahch@si.edu For example, the slope region off the south-eastern US may have one of the most extensive deep coral areas in US waters (Hain & Corcoran, 2004). Amongst the most prominent members of deep-sea coral reefs are echinoderms (e.g. Chave & Malahoff, 1998; Krieger & Wing, 2002), especially members of the Asteroidea, also known as sea stars or starfish. Ecological interactions between asteroids and colonial cnidarians, such as corals have been heavily studied (e.g. Birkeland & Lucas, 1990) and available data suggest that asteroids are ecologically important in these settings (e.g. Birkeland, 1974). Conservation of coral habitats begins with an understanding of 266 ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 267 community structure and function of associated fauna, including identification and inventory of the taxa that compose that fauna. Surveys of deep-sea coral habitats in the Atlantic and Pacific have led to the collection and observation of several asteroids, especially those in the Gonias- teridae, which have been observed in association with deep-sea corals. These asteroids belong primarily to the Hippasterinae, a small but discrete subfamily known primarily from cold-water settings. New data, presented herein, demonstrate predation by nearly all members of this group on multiple species of deep-sea coral. In the Atlantic, recent collections (S. W. Ross and M. Nizinski, unpubl. data) at multiple sites off the south-eastern coast of the US have revealed, to date, many specimens of asteroids, representing 12 fami- lies, 18 genera, and 18 species (M. Nizinski, unpubl. data), associated with deep-sea coral habitats. This includes two forms of hippasterines from the south- eastern US, including one species that was consistent with prior descriptions of 'Hippasteria' caribaea as summarized by Verrill (1899) and a second species from the Georgia coast that did not fit into any known goniasterid genera. In the North Pacific, surveys by the Monterey Bay Aquarium Research Institute (MBARI) from sea- mounts (Lundsten et al., 2009) have also resulted in collections of several hippasterine asteroids, from deep-sea coral assemblages. Numerous collected specimens included Hippasteria and the rarely encountered Cryptopeltaster, and Evoplosoma. Col- lected Evoplosoma specimens were unknown from this region of the North Pacific and could not be reconciled with any known species. Motivated by discovery of this new material in addition to new in situ observations of hippasterines preying on deep-sea corals, we present (1) descrip- tions of one new genus and three new species of previously undescribed hippasterine sea stars; (2) a review and phylogenetic revision of the Hippasteri- nae; and (3) discussion of corallivorous behaviour in the Goniasteridae. Because of the complex nature of relationships amongst species in Hippasteria, a com- prehensive assessment of species in this genus was beyond the scope of the current study. However, a full summary of all known hippasterine species is provided as a starting point to encourage future studies. TAXONOMIC BACKGROUND Hippasteria Gray, 1840 and related taxa are amongst the first described asteroids (with pre-Linnean desig- nation Pentaceros planus Linck, 1733). The subfamily Hippasterinae was designated by Verrill (1899) to include Hippasteria and Cladaster. Fisher (1905, 1906) subsequently described two new genera, Cryp- topeltaster and Evoplosoma, respectively, which he included in the Hippasterinae (Fisher, 1911). Fisher (1906) described Gilbertaster and suggested it was an 'aberrant member' of the Hippasterinae but never included Gilbertaster in this subfamily with Cryp- topeltaster and Evoplosoma. Fisher (1911, 1940) later removed Cladaster from the Hippasterinae. Mah (2006) determined that Cladaster had closer affinities with Circeaster based on morphology, thus providing support for removal of Cladaster from the Hippasteri- nae. The Hippasterinae has not been widely recog- nized in recent monographs and classifications (e.g. Clark & Downey, 1992; Clark & McKnight, 2001). Fisher (1940) described the subfamily Hippasterinae as 'superfluous', based on perceived phylogenetic con- fluence (Cladaster) with the Goniasteridae. However, Spencer & Wright (1966) retained usage of the Hip- pasterinae and the grouping has persisted within the Goniasteridae (e.g. Clark & Downey, 1992). Dons (1937) recognized two subgenera, Hippasteria (Euhip- pasteria), with large, subquadrate or rectangular marginal plates and Hippasteria (Nehippasteria), which had more oval-shaped plates. These concepts, however, have never been tested. Although revisions of local faunas have been performed (Clark & Downey, 1992; Clark & McKnight, 2001; McKnight, 2006), a full evaluation of this group and its members has not been attempted. Hippasteria, the type genus for the group, includes approximately 18 nominal living species distributed largely in cold-water settings, including high-latitude and deep-water habitats (Clark, 1993). Several of the species and subspecies, especially those associated with Hippasteria phrygiana and Hippasteria spinosa, are questionable and often show weakly defined species boundaries. Many of these species are widely distributed but may be morphologically indistinguish- able. For example, the South Pacific?New Zealand Hippasteria trojana has been synonymized with the Atlantic H. phrygiana (Clark in H.E.S. Clark & McK- night, 2001). Surprising amounts of cryptic species diversity have been encountered in widespread species with highly similar to identical external mor- phology (e.g. Vogler etaL, 2008), suggesting that several of the species boundaries in Hippasteria, are unlikely to be resolved solely on morphological data. Cryptopeltaster includes two nominal species, occur- ring in moderately deep-water (-200?700 m) along the west coast of North and South America. Evoplo- soma includes four nominal species and occurs in deeper water (700?2000 m) in the Indian, Pacific, and Atlantic ocean basins (Clark, 1993). Gilbertaster occurs only in deep-water in the central and south Pacific (Clark & McKnight, 2001). ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 268 C. MAH ETAL. MATERIAL AND METHODS In addition to new material provided by M. Nizinski and L. Lundsten (see map in Fig. 1 for collection localities), additional specimens from the Smithso- nian Institution (USNM) and the Museum of Com- parative Zoology (MCZ) were examined. External features and variation in accessory structures were primary sources of character data observed mostly from dried specimens, although wet specimens were also examined. Mediaster aequalis (Stimpson, 1857) and Peltaster placenta (Miiller & Troschel, 1842) were selected as outgroups based on preliminary morphological data supporting these taxa as stemward relative to Hip- pasteria (Mah, 2005a). Cladaster was also selected owing to its past classification history within the Hippasterinae. Ingroup taxa were chosen based on their taxonomic inclusion in the Hippasterinae following Fisher (1911) and Spencer & Wright (1966). Gilbertaster was included based on the close morphological similarity mentioned by Fisher (1906) and partial support from preliminary morphological data (Mah, 2005b). Hip- pasteria includes a large number of morphologically similar species, some of which occur across wide dis- tances (e.g. northern vs. southern hemispheres) and a full revision was beyond the scope of the project. Hippasteria species chosen for inclusion were repre- sentative of the diversity of body plans observed within the genus. Evoplosoma was represented by two Atlantic species in addition to two undescribed Pacific species. Nearly every available species of Evoplosoma was studied from type or previously pub- lished specimen material. Type material for two of the known species (Evoplosoma forcipifera and Evoplo- soma augusti) is missing. METHODS Phylogenetic analysis utilized morphological data from 46 characters scored from 12 ingroup taxa (see Appendices 1 and 2). Data were entered into Mac- Clade, exported to PAUP*4.0bl0 (Swofford, 2003) and analysed using the Branch-and-Bound search option. Characters were treated as unordered and with equal weight. Branch support was calculated for Bremer and bootstrap values (Fig. 2). Both were calculated using Central California, USA N I O Evoplosoma claguei A, Evoplosoma voratus Contours (500 m) Figure 1. Collection locality maps. A, tropical Atlantic localities for Sthenaster and Gilbertaster caribaea; B, North-east Pacific localities for Evoplosoma spp. nov. ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 269 57% OG: Mediaster aequalis ? OG: Cladaster analogus - OG: Peltasterplacenta 1.9-2.5-2.6-2.11-4.3 7.6 89% 1.5-2.1- 2.2-2.12- 3.3-4.1 1.8- 2.4- 3.4- 4.3- 2.3- 2.7- 3.5- 7.5 1.1-1.6- 1.10-2.5 2.6-7.1- 7.2-7.4- 7.7-8.1- 8.2 SG% 1.6-1.9-1.1 2.11-8.3 1.3-12- 4.5-16- 7.1 7.3 Hippasteria californica Hippasterias heathi 3s' 12-7.5 Hippasteria spinosa Hippasteria phrygiana 1.5-1.6-1.11-1.12-2.1-2.2- 2.3-2.4-2.9-2.10-7.4 * Evoplosoma virgo 7.2- 7.7 1.8-3.1 65% 1.2-1.6- 2.3-2.4- 2.5-2.6- 2.8-2.9- 2.10-3.1 2.7- 3.5 65?/ 1.5 -* Evoplosoma claguei n. sp. 3.4-7.5 -%Evoplsoma voratus n. sp. 2.3-2.4-4.1 -#- Evoplosoma scorpio -Sthenaster emmae n. gen. n. sp. 1.5-2.1-2.2- 3.3-4.1-4.4 -#? Cryptopeltaster lepidonotus 1.2-1.4-2.8-2.9-2.10 3.2-7.4-7.7-8.1-8.2-8? 3.1; 5 changes 100% "a 1.7 "Hippasteria" caribaea Gilbertaster anacanthus Gilbertaster Figure 2. Tree results: topology and support values. In the other equally parsimonious tree topology the positions of Evoplosoma virgo and Evoplosoma scorpio are transposed. Otherwise the tree is identical. Bootstrap values are present as % in bold vs. Bremer values which are oversized numbers that sit below the bootstrap values. Decimal numbers represent apomorphies on each node for each member of ingroup taxa correspond to terms in Appendix 2. OG, outgroup. PAUP. Bremer was calculated based on the addition of additional steps to the most parsimonious tree until branches collapsed into polytomies. Bootstrap values were calculated from 200 replicates using random stepwise addition. RESULTS AND CLASSIFICATION The analysis resulted in two most parsimonious trees each with 111 steps and a consistency index of 0.6036 and a homoplasy index of 0.3964. Each tree displayed essentially identical topologies (Fig. 2), differing only in the positions of Evoplosoma virgo and Evoplosoma scorpio (discussed further below). The Hippasterinae (Fig. 2) are strongly supp- orted (bootstrap = 99%) including a monophyletic Hippasteria (bootstrap = 85%), and Gilbertaster (boot- strap = 100%), Gilbertaster + Cryptopeltaster (boot- strap = 62%), and Evoplosoma (bootstrap = 66%). The Hippasterinae was separated from the outgroup taxa, including Cladaster, which was previously included amongst the Hippasterinae. Cryptopeltaster and Sth- enaster are both monotypic taxa and although some variation within the species is present, none war- ranted further testing within the context of the phy- togeny. Bootstrap support was absent for the Evoplosoma plus Hippasteria branch as well as for Hippasteria itself. Gilbertaster was relatively well supported within the Hippasterinae as the sister taxon to the mono- typic Cryptopeltaster as part of the sister clade to the other hippasterines. This relationship loosely sup- ports statements expressed by Fisher (1906: 1065) that 'Gilbertaster appears to be allied to some of the ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 270 C. MAH ETAL. Hippasterine, particularly to Cryptopeltaster . . .' as well as his general perception that Gilbertaster was 'intermediate between the Goniasterinae and Hippas- terinae . . .' and 'might be considered ... an aberrant member of the Hippasterinae'. Hippasteria caribaea is strongly supported as sister taxon to the Pacific Gilbertaster anacanthus (boot- strap = 100%). Based on these results and the number of shared characters, Hippasteria caribaea is moved to the genus Gilbertaster. Two species are now included in Gilbertaster, which extends the range of this genus to include the western Atlantic. Hippasteria has a fairly distinctive suite of diag- nostic characters in addition to strong bootstrap support (Fig. 1). Hippasteria californica is supported as the sister group to a polytomy including Hippas- teria heathi, Hippasteria spinosa, and Hippasteria phrygiana. Hippasteria californica shares at least one character, oval to rounded marginal plate shape, with Hippasteria insignis and Hippasteria phrygiana strongylactis, which are either synonymous or subspe- cies of H. phrygiana. Hippasteria insignis was included in a separate subgenus Hippasteria (Nehip- pasteria) (Dons, 1937) but has since been synony- mized with Hippasteria (Clark & Downey, 1992). Some species, such as Hippasteria falklandica and Hippasteria tasmanica have been described as having more elongate marginals then those observed in H. phrygiana or H. spinosa, suggesting that further species may be considered for inclusion in H. (Nehip- pasteria). Thus, further taxon and character sampling may provide the potential for discovering additional diversity within Hippasteria. All four species of Evoplosoma included in this analysis were supported together on a single branch (Fig. 2) but taxonomic sampling remains incomplete. Information for almost all of the species of Evoplo- soma was based on single specimens but all species seem to display a relatively consistent suite of mor- phological characters. The original description of the Indian Ocean species, E. augusti by Koehler (1909) (specimen could not be located for this study), sug- gests that this species may show greater divergence from the Atlantic and Pacific species and may actually demonstrate a morphology intermediate to other hippasterines. SYSTEMATICS Notes on terminology: Sections entitled Diagnosis are intended for summary descriptions for genera as opposed to sections entitled Description, which are intended for full species descriptions. Monotypic taxa did not warrant a diagnosis separate from specific description. A full summary of morphological termi- nology and an identification key is in Appendix 3. Appendix 4 includes a full list of new genera and species as well as taxonomic changes included below. FAMILY GONIASTERIDAE FORBES, 1841 SUBFAMILY HIPPASTERINAE VERRILL, 1899 Verrill, 1899: 174; Fisher, 1906: 1165; 1910: 223; Spencer & Wright, 1966: U58 Notes: Our phylogenetic analysis supports the mono- phyly 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) con- clusion that the Hippasterinae is 'superfluous.' Fisher based his conclusion on the 'intermediate' morphologi- cal characters observed in Cladaster. Mah (2006) sup- ported Cladaster as the sister group to the Circeaster lineage; however, it is possible that either the Cir- ceaster 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. Supermarginal and infero- marginal plates, wide to quadrate with large, promi- nent spines in most taxa. Pedicellariae enlarged, abundant and often on raised base. Marginal plates facing laterally. Disk strongly swollen in most. GILBERTASTER FISHER, 1906 Fisher, 1906: 1062; McKnight, 1973: 192; Mah, 1998: 66; H.E.S. Clark & McKnight, 2001: 49 Type species: Gilbertaster anacanthus Fisher, 1906. Included species: G. anacanthus Fisher, 1906; G. car- ibaea (Verrill, 1899). Diagnosis: Arms triangular, broad to tapering, narrow (Fig. 3A, E). Disk weakly swollen (Fig. 3A, E). Tissue thick, pulpy covering abactinal, marginal, actinal plates. Abactinal plates low, polygonal covered by one to 12, closely articulated angular granules ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 271 Figure 3. Gilbertaster caribaea comb. nov. USNM 1126236. A, abactinal surface; B, close-up abactinal surface; C, dorsolateral surface showing superomarginals; D, oral region. Gilbertaster anacanthus holotype USNM 021168; E, abactinal surface; F, close-up abactinal surface; G, oral region. Scale bars: A, C, E = 5 mm; B, F, G = 2 mm. (Fig. 3A- C, E-G) elongate to round in length, flat- tened, forming smooth to rough surface texture. Fas- ciolar grooves shallow. Secondary plates present between abactinal plates (Fig. 3C). Abactinal plates with angular accessories (Fig. 3A?C). Pedicellariae large, bivalved with smooth valves, the length of one to two plates and abundant on abactinal surface (Fig. 3B, C). Marginal plates, 50?70 per inter-radius (armtip to armtip), squarish in outline with rounded edges, com- pletely covered by angular granules similar to those on abactinal surface. Variable surfaces smooth (on G. anacanthus) to roughened (on G. caribaea). Pedicel- lariae large, bivalve (Fig. 3C) similar to those on abactinal surface on marginal plate surface, often bisecting the width of the plate. Spines absent from supermarginal and inferomarginal plate series (Fig. 3A, C, E). Granules, densely arranged, covering supermarginal and inferomarginal plate series com- plete (Fig. 3A, B, C, E). Superomarginal and infero- ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 272 C. MAH ETAL. marginal plates quadrate at inter-radius (Fig. 3C, E). Fasciolar grooves on marginal and actinal surfaces absent. Fringe of accessories on marginal plates poorly differentiated. Superomarginal plates forming prominent dorsolateral fringe (Fig. 3A, E). Actinal plates covered by one to 15 flattened, polygonal, angular granules. Large bivalve pedicel- lariae similar to those on abactinal, marginal plate surfaces abundant on actinal plates. Actinal plates with granules, but lacking large spines or spinelets (Fig. 3D, G). Pedicellariae, bivalved, present on plate series at perpendicular angle (Fig. 3A?G), adjacent to ambulacral furrow. Pedicellariae, flat-tong shaped with serrated blades present on actinal plates (Fig. 3D, F). Pedicellariae very abundant, present on raised bases (Fig. 3A-G). Furrow spines two to four (usually three) blunt, thickened spines, horizontally flattened (G. anacan- thus) to triangular/quadrate in cross-section (G. car- ibaea) (Fig. 3D, G). Subambulacral spines, one to four, blunt, flattened (Fig. 3D, G). Round to quadrate (G. anacanthus) to triangular in cross-section (G. carib- aea). Pedicellariae, bivalved, enlarged on first adam- bulacral (similar to others) replacing subambulacral spination (Fig. 3D, G) and sometimes replacing furrow spination. Subambulacral spines smaller in size, more abundant (Fig. 3D, G). Furrow spines round in cross-section, not compressed. Oral plates covered by ten to 20 flattened, angular, closely articulated granules (Fig. 3D, G), sometimes with enlarged bivalve pedicellariae. Oral plate furrow spines, typically five, flattened to oval in cross-section. Oral region concave (Fig. 3D, G). GlLBERTASTER ANACANTHUS FlSHER, 1906 FIGURE 3E-G Fisher, 1906: 1063; A.M. Clark, 1993: 223; H.E.S. Clark & McKnight, 2001: 49 (as Gilbertaster ana- canthus) McKnight, 1973: 192; A.M. Clark, 1993: 223 (as Gil- bertaster 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, ll.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 gran- ules, one to six, round to irregularly polygonal to oblong to elongate in outline, forming angular fringe around each (Fig. 3E, F). Carinal series distinct (Fig. 3F). Granules slightly convex to rounded but low and close to disk surface (Fig. 3E, F). Periphery of plate surrounded by four to 12 elongate to rounded- polygonal 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 iden- tical 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 ornamen- tation (i.e. spines, etc.) absent from marginal plate surface. Granules form weakly differentiated periph- ery around marginal plate surface. Marginal fascioles absent (Fig. 3E, F). Superomarginal plate surface with dorsol facing (Fig. 3E), 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. 3G). Granules present, four to 16 in number, closely abutted and polygonal-oblong to irregular, rounded with low convex appearance in shape (Fig. 3G). Granular cover on actinals flush with those on adjacent inferomar- ginal 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. 3G). 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 ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 273 the whole length of the plate (Fig. 3G). 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 cross- section in irregular rows (Fig. 3G). 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). GlLBERTASTER CARIBAEA (VERRILL, 1899) COMB. NOV. FIGURE 3A-D Verrill, 1899: 174, pi. 28; Halpern, 1970a: 190; Clark & Downey, 1992: 246 (as Hippasteria caribaea) Occurrence: Known from Cumberland Island, Georgia, Savannah Banks, Stetson Banks, off Jack- sonville, FL, and off the coast of West End, Grand Bahama. 500-805 m. Material examined: HOLOTYPE: USNM 18425, Cum- berland 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, l0.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. A = 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. 3A). Abactinal surface covered by coarse granules, densely packed with no plate surface visible. Gran- ules forming continuous cover, nearly contiguous with supermarginal plate series (Fig. 3A, B). Plates, each with one (exceptionally two or more) large round, tubercular granule(s), surrounded by three to six smaller rounded coarse granules (Fig. 3C). Abactinal plates closely abutting (Fig. 3B) in adult specimens, somewhat less so in smaller individuals (when R = 4- 5 cm). Papulae single, distributed over most of abac- tinal surface but absent from narrow triangular area adjacent to contact with superomarginal plate series. Madreporite round with well-developed sulcae, sur- rounded 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. 3B, C). Marginal plates elongate [length (L) > width (W)], largest inter-radially, becoming more equal in size and smaller distally. Marginal plates 40?50 per inter- radius (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, hemi- spherical to polygonal to more oblong (Fig. 3C). Superomarginal granules prominent, strongly convex, not forming even surface with other granules (Fig. 3C). 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 pedicel- lariae. Largest centrally becoming smallest near the ends of each pedicellariae. Other actinal intermediate plate chevron series more irregular with approxi- mately 60% of plates bearing a large bivalve pedicel- lariae (Fig. 3D). Holotype with nearly all plates bearing a large bivalve pedicellariae. Remaining plates covered by four to 14 (mean of nine) granules. ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 274 C. MAH ETAL. 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 adambu- lacral plate with fewer spines on distal plates (Fig. 3D). One large thick (3^ x thickness of furrow spines), angular, subambulacral spine, present imme- diately behind furrow spines. This large subambulac- ral spine flanked by one to two smaller spines, triangular in cross-section, roughly half the height of the large subambulacral (Fig. 3D). Several thick- ened, blunt granules, polygonal-triagonal in cross- section, five to nine adjacent to the subambulacral spine, similar in size to those on actinal surface. Typically, one enlarged, round granule present adja- cent to subambulacrals with other granules irregu- larly trailing off in size. Distinct groove between adambulacral plates and first adjacent actinal inter- mediate 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. 3D). Oral plate covered by two enlarged subam- bulacral 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. Niz- inski were observed as solitary individuals collected on soft substrata. No other organisms were observed at the collection site. STHENASTER GEN. NOV. Description: As for species. STHENASTER EMMAE SP. NOV. FIGURE 4A-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 E15539, off coast of Jack- sonville, 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. 4A, B). Abactinal plates round to polygonal to oblong in outline with numerous interspersed secondary plates (Fig. 4F) embedded in a thick, pulpy tissue forming heavily thickened abactinal body wall (Fig. 4A). Plates with high-aspect; mound-like in shape, forming deep but open channels between plates (Fig. 4A, E). Abactinal plates are covered with two to ten blunt to conical spine-like granules, usually forming a periph- ery 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 pedi- cellariae present, each with nine to 12 interlocking teeth per valve (Fig. 4E). Pedicellariae located cen- trally on plate surface varying in size from one-third to three-quarters of plate diameter, flanked by spine- like 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 inter- radius (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. 4H). Superomarginals facing laterally, correspond 1:1 to inferomarginals along most of series with one to two plates irregularly offset, possibly because of sublethal predation. Supermarginal 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. 4H). One to two relatively small clam-shell like pedicellariae with interlocking teeth present on nearly every marginal plate but regularly present, adjacent to inferomar- ginal 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. ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 275 Figure 4. Sthenaster emmae gen. et sp. nov. USNM 1124468. A, abactinal surface; B, actinal surface; C, oral region; D, actinal-inferomarginal contact; E, close-up of abactinal surface showing pedicellariae; F, close-up of abactinal plates on armtip; G, adambulacral furrow spines; H, marginal plates ? lateral facing of supermarginal plates. Scale bars: A, B = 10 mm; C-H = 5 mm. 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 supermar- ginal 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 supero- marginals present on inferomarginals facing ven- trally. Terminal plate round, bulbous. Actinal intermediate plates, similar in size, shape to abactinal plates with well-developed fasciolar chan- nels running between plates (Fig. 4C, D). Actinal plate series adjacent to adambulacral plate series with large pedicellariae nearly equal to size of plate on which it sits (Fig. 4C, D), teeth poorly developed to absent relative to those on abactinal plates. Pedicel- lariae 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. Pedi- cellariae on each actinal plate surrounded by 20?35 sharp spinelets, some round, some triagonal to polygonal in cross-section (Fig. 4C, 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 cross- section. Oral cavity sunken (Fig. 4B, C). Adambulacral furrow spines blunt tipped, three to four per plate, triangular to oblong ovate in cross- section in weakly convex series (Fig. 4G). 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 cross- section (Fig. 4G). Subambulacral spine series far- thest away from furrow spines shortest, with central ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 276 C. MAH ETAL. spines only slightly less thick than furrow spines with smallest spines located on ends of second sub- ambulacral series. Distinct fasciolar channel sepa- rates 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. AC). 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 granu- lar 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 gorgon- ians, 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, sclerac- tinian corals, and coral rubble were also present. CRYPTOPELTASTER FISHER, 1905 Fisher, 1905: 311; 1911: 237; Spencer & Wright, 1966: U58; A.M. Clark, 1993: 251. 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.H.1889. Included species: Cryptopeltaster lepidonotus Fisher, 1905 {Cryptopeltaster philipii is now a synonym of C. lepidonotus). Diagnosis, distribution, and characters: As per species. CRYPTOPELTASTER LEPIDONOTUS FISHER, 1905 FIGURE 5A-E Ludwig, 1905: 138 (as Hippasteria pacifica) Fisher, 1905: 311; 1911: 237; Lambert, 1978a: 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 fins), coll. M. Eric Anderson, 7.vi.l977 (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. # = 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. # = 9.6, r = 6.1); USNM E47396, 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. # = 9.8, r = 3.8; R = 10.8, r = 4.8); USNM E51296, 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. # = 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 supermarginal 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 supermarginal 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 ter- minal 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 through- out surface. Spines, one to three (typically one), short, conical to tubercular present at lower end of each supermarginal plate adjacent to contact with infero- marginal 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 ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 277 i'l^Mf^ Figure 5. Cryptopeltaster lepidonotus neotype USNM 33356. A, abactinal surface; B, close-up of pedicellariae and large angular granules; C, adambulacral furrow spines and bivalve pedicellariae. USNM E47396; D, lateral view showing supermarginal plate series; E, actinal surface showing furrow spines, pedicellariae, and actinal spines and granules. Scale bars: A, D, E = 5 mm; B = 4 mm; C = 3 mm. 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 ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 278 C. MAH ETAL. the body surface, an undivided madreporite, and fewer supero- and inferomarginal plates per inter- radius. Cryptopeltaster from the Galapagos (USNM E51296) 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: pi. 47, fig. 1). Other speci- mens 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 speci- men (USNM E51296) 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 corre- sponds to the description of C. lepidonotus, also had 42 marginal plates. Hippasteria paciflca Ludwig, 1905 from Mexico was synonymized with C. lepidono- tus by Fisher (1911) and was represented by a smaller specimen (R = 4.8 cm) with approximately 40 mar- ginal plates. Codoceo & Andrade (1981) did not include the size of the holotype, which is now appar- ently 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 charac- teristic 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. EVOPLOSOMA FISHER, 1906 Fisher, 1906: 1065; Koehler, 1909: 96; Spencer & Wright, 1966: U58; A.M. Clark, 1993: 253. 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 fas- ciolar grooves shallow. Secondary plates present. Abactinal plates, fiat and platform-like. Carinal series poorly distinguished. Abactinal plates tightly articu- lated. Large spines or spinelets on abactinal, super- marginal, inferomarginal plate, and actinal series. Accessories on supermarginal and inferomarginal surface widely spaced. Superomarginal plate series with lateral facing. Supermarginal and inferomarginal plates at inter- radius quadrate in shape. Spinelets present on both marginal plate series. Fasciolar grooves between mar- ginal plate series shallow. Accessories around supero- marginal 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; Evoplo- soma claguei sp. nov.; E. forcipifera Fisher, 1906; E. scorpio Downey, 1982; Evoplosoma timorensis Aziz & Jangoux, 1985a; E. virgo Downey, 1982; Evoplosoma voratus sp. nov. EVOPLOSOMA CLAGUEI SP. NOV. FIGURE 7A-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, A8, 14.X.2003 (1 wet spec. R = 9.9, r = 2.6 cm). PARATYPES: SIO E2440, 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. D77- A3. 30 Aug 2009 (1 wet spec. R = 11.1, r = 2.7 cm). Etymology: This species is named after Or 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 con- ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 279 sistent 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. 7C). Abac- tinal plates larger, approximately six plates across from supermarginal to supermarginal, 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. 7C). 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 chisel- shaped 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 interme- diate areas and angular in shape. Periphery of these plates covered by 13^10 quadrate granules, approxi- mately 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 dis- tally. Nine to ten furrow spines on first adambulacral plate. Furrow spines flattened, paddle like to trian- gular 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 pro- nounced striations. Ambulacral and subambulacral series flanked by ten to 15 round, hemispherical gran- ules varying in size. Largest adjacent to subambulac- rals, 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. EVOPLOSOMA VORATUS SP. NOV. FIGURE 8A-E Occurrence: Known from Davidson Seamount, 2669.9 Material examined: HOLOTYPE: USNM 1124506, Davidson Seamount, 35?37'N, 122?49'W, 2669.9 m, coll. J. Barry, L. Lundsten, Sta. T-947-A10, 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 supermarginal plate series but larger distally, especially on arms. One to four plates across distance ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 280 C. MAH ETAL. of arm between superomarginal plate series. Abacti- nal plate surfaces mostly bare with short, stubby conical tubercles. Plates with tubercles not very abun- dant, only about four to six per arm base on disk. Each plate with ten to 30 peripheral rounded gran- ules that are small and rounded to quadrate in cross- section. Region adjacent to madreporite (about 0.75 x 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 oval- shaped 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 becom- ing 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. Superomar- ginal and inferomarginal offset from one another and each series with different overall dimensions. Supero- marginals number 44?45 from armtip to armtip, more elongate (L > W), becoming almost ovalate. Infero- marginals number 54?55 from armtip to armtip, more quadrate (L = W) overall. Superomarginal plate contact with abactinal surface, strongly convex, oval- like 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. Subambulac- ral 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 com- pressed 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 preser- vation in 75% ethanol. Observed in situ apparently feeding on Trissopathes pseudotristicha (Cladopathidae, Antipatharia). EVOPLOSOMA AUGUSTI KOEHLER, 1909 Koehler, 1909: 96; Clark, 1993: 253 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. EVOPLOSOMA FORCIPIFERA FlSHER, 1906 Fisher, 1906: 1065; Clark, 1993: 253 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 ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 281 Figure 6. Evoplosoma scorpio USNM E23623. A, abactinal surface of disk and arms; B, actinal surface proximal to mouth; C, furrow spines and pedicellariae. USNM E50539; D, actinal intermediate surface region, showing adambulacral and furrow spination; E, adambulacral and furrow spination along actinal arm surface. Scale bars: A = 3 mm; B?E = 2 mm. species. The holotype for this species is missing from the USNM collections (Ahearn, 1995). EVOPLOSOMA SCORPIO DOWNEY, 1981 FIGURE 6A-E Downey, 1981: 561; Gage etal, 1983: 280; Clark & Downey, 1992: 242; Clark, 1993: 253. Occurrence: South-west Rockall Trough to north- eastern European Basin (48.5?N, 10?W) extended herein to off Delaware 38?45'N, 72?40'W. 1600- 2105 m. Material examined: USNM E50539, 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 E23623, 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). ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 282 C. MAH ETAL. Figure 7. Evoplosoma claguei sp. nov. USNM 1124507. Holotype. A, abactinal surface; B, close-up showing abactinal plates, spines, and marginal series; C, abactinal plates on arm surface; D, actinal surface; E, close-up of furrow spines and adambulacral armature; F, In situ observation of Evoplosoma (cf. E. claguei) feeding on Keratoisis from Rodriguez seamount (dive 662-02, at 1842 m). Scale bars: A, D = 5mm; B, C, E = 3 mm; F = 5 cm. EVOPLOSOMA VIRGO DOWNEY, 1982 FIGURE 9A-D Downey, 1982: 772; Clark, 1993: 253 Occurrence: Gulf of Mexico, 2056 m. Material examined: HOLOTYPE: USNM E24285, Material examined: None. Gulf of Mexico, North Atlantic Ocean, 26?08'N, ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 EVOPLOSOMA TIMORENSIS Aziz & JANGOUX, 1985 Aziz & Jangoux, 1985a: 263; Clark, 1993: 253 Occurrence: East Timor Region, Malaysia, 8?50.2'S, 127?2'E. 883 m. NEW TAXA & PHYLOGENY OF HIPPASTERINAE 283 u ? * * . y* jMjfa^j ^ 4 r3 -<*---*%M .. fc., ?"?*3I ^/^ Figure 8. Evoplosoma voratus sp. nov. USNM 1124506. A, abactinal surface; B, close-up of abactinal plates and madreporite; C, lateral view of marginal plates; D, oral surface; E, close-up of furrow spines an adambulacral armature; F, in situ video of E. voratus feeding on Trissopathes pseudotristicha on Davidson Seamount. Two circular dots (lights) are 29 cm. Scale bars: A, B, D, E = 3 mm; C = 6 mm. 92?43'W, 2056 m (1124 fms), coll. W. Pequegnat, Occurrence: Off Java (Indonesia). 2186 m. 30.vii.1971 (1 dry spec. R = 11.5 cm, r = 3.3 cm). EVOPLOSOMA SP. 1 Fujikura, Okutani & Maruyama, 2008: 272 (as Goni- asteridae gen. et sp. fig. 22.19) 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. ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 284 C. MAH ETAL. Figure 9. Evoplosoma virgo USNM E24285 HOLOTYPE. A, abactinal surface; B, oral surface showing furrow and adambulacral armature; C, close-up of abactinal granules; D, dorsolateral view showing abactinal plates (AB), super- marginal (SM), and inferomarginal (IM) plate series. Scale bars: A = 20 cm; B, C = 2 mm; D = 5 mm. HIPPASTERIA GRAY, 1840 Gray, 1840: 279; 1866: 9; Perrier, 1875: 271; (1876: 86); Sladen, 1889: 341; Fisher, 1911: 223; Verrill, 1914: 300; Koehler, 1924: 178; Mortensen, 1927: 88; Dons, 1937: 17; Mortensen, 1940: 125; Djakonov, 1950: 51 (1968: 42); Bernasconi, 1963: 15; Bernasconi, 1964: 253; Halpern, 1970a: 183; A.M. Clark & Courtman-Stock, 1976: 63; Clark & Downey, 1992: 246; Clark & McKnight, 2001: 54. Type species: Asterias phrygiana Parelius, 1768 (with synonym Hippasteria europaea Gray, 1840 as well as all four nominal species included by Gray as syn- onyms 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, polygo- nal to irregular in outline, flat and platform-like in shape. Carinal series are poorly distinguished. Abac- tinal spinelets (sometimes granular) forming fringe around abactinal plates. Spines, large, conical, gran- ules often present on abactinal plates. Large spines present on supermarginal and inf- eromarginal 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 supero- marginal 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 ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 285 derjungini Djakonov, 1950; H. falklandica Fisher, 1940; H. heathi Fisher, 1905; Hippasteria imperialis Goto, 1914; Hippasteria kurilensis Fisher, 1911; Hippasteria leiopelta Fisher, 1910; Hippasteria mam- mifera 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. strongy- lactis 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 approxi- mately 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 conser- vative 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 geographi- cal 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 haghsh, which exploit prey cap- tured, 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). 'HIPPASTERIA' ANTIQUA FELL, 1956 Fell, 1956: 11. Occurrence: Fossil taxon. From Senonian (upper Cre- taceous) sediments in north Canterbury, New Zealand. Comments: Based on Fell's (1956) description of this species, relatively few characters were readily observ- able from the available specimen. Characters used by Fell (1956) in the description of this specimen that placed it in Hippasteria include larger primary plates with a radial series that are circular in shape in addition to numerous smaller secondary plates scat- tered throughout and quadrangular marginal plates. These characters are plesiomorphic for the Hippas- terinae and are present in stemward taxa, such as Sthenaster, and some Hippasteria spp. Further examination of the fossil and comparison with living taxa is required. HIPPASTERIA CALIFORNICA FISHER, 1905 FIGURE 6E-G Fisher, 1905: 310; 1911: 233; HL Clark, 1913: 194; Alton, 1966: 1702; Lambert, 1978b: 62; Maluf, 1988: 34, 118 (as H. californica) Imaoka etal., 1991: 54 (as Cryptopeltaster lepidonotus) Occurrence: California, Washington to British Colum- bia, and Hokkaido, Japan 110?1820 m. Material examined: USNM 33354, San Miguel Island, Channel Islands, California. 34?15'N, 120?36'W, 519 m, coll. USFC Albatross, 5.1.1889. (1 dry spec. R = 8.2, r = 4.3); USNM E10413, south-west of mouth of Columbia River, Oregon. 45?52'N, 124? 52'W, 823 m. Coll. R/V Commando, 13.V.1963 (1 dry spec. R = 6.3, r = 3.2). USNM E 10414 south-west of mouth of Columbia River, Oregon. 45?54'N, 124?55'W 823 m. Coll. R/V. Commando, 10.xii.1961. (1 dry spec. R = 8.7, r = 3.3). USNM E 10443, south-west of mouth of Columbia River, Oregon. 45?54'N, 124?55'W, 823 m. Coll. R/V Commando 10.xii.1961 (1 dry spec. R = 4.2, r = 22). USNM E 10441 south-west of Columbia River, Oregon. 46?2'N, 124?57'W, 915 m. Coll. R/V Com- mando 30.V.1964. (1 dry spec. R = 1.9, r = 1.0). USNM E10442 south-west of mouth of Columbia River, Oregon. 45?55'N, 124?54'W, 732 m. Coll. R/V Com- mando 5.ix.l964. (1 dry spec. R = 2.9, r = 1.5). HIPPASTERIA COLOSSA DJAKONOV, 1950 Djakonov, 1950: 55 (1968: 47); 1952: 411; Baranova, 1957: 162 Occurrence: Bering Sea, 238?250 m. ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 286 C. MAH ETAL. 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. HlPPASTERIA DERJUNGINI DJAKONOV, 1950 (Hippasteria derjungini Djakonov, 1949: 22 is a nomen nudum, insufficiently characterized) Djakonov, 1950: 55 (1968: 46); 1952: 412 Occurrence: Okhotsk Sea, off the northeast coast of Sakhalin. 192 m. Material examined: None. HIPPASTERIA FALKLANDICA FISHER, 1940 Fisher, 1940: 125; Bernasconi, 1973: 287; Clark & Downey, 1992: 247; Stampanato & Jangoux, 2004: 6; McKnight, 2006: 97 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. HIPPASTERIA KURILENSIS FISHER, 1911 Fisher, 1911: 226 (as H. spinosa kurilensis) Gish, 2007: 39 (as H. kurilensis) 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. HIPPASTERIA LEIOPELTA FISHER, 1910 (with formae aculeata and longimana Djakonov, 1950) 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. Material examined: None. HIPPASTERIA HEATHI FISHER, 1905 Fisher, 1905: 319; 1911: 231; Krieger & Wing, 2002: 86. Occurrence: Alaska, Gulf of Alaska. 377^54 m. Material examined: HOLOTYPE: USNM 22338, Guard Island, Behm Canal, Alexander Archipelago, Alaska, 55?N, 131?W, 377-454 m, coll. USFC Alba- tross, 9.vii.l903 (1 dry spec. R = 7.8, r = 3.9). HIPPASTERIA IMPERIALIS GOTO, 1914 Goto, 1914: 338; Hayashi, 1952: 338; 1973: 6, 15; Imaoka etal., 1990: 50; Mah, 1998a: 67; Fujikura et al., 2008: 272 (as H. imperialis) Grigg etal., 1987: 387 (as Hippasteria spinosa) HIPPASTERIA MAMMIFERA DJAKONOV, 1950 Djakonov, 1950: 54 (1968: 45), 1952: 409. 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 under- mines the distinctiveness of this species and further lends support to its synonymy with H. spinosa. Material examined: None. HIPPASTERIA NOZAWAI GOTO, 1914 Goto, 1914: 344. Occurrence: Sagami Bay/Tosa Bay region, Kii Strait, southern Japan, Hawaiian Islands. 245?600 m. Occurrence: Hokkaido, northern Japan. (No depth information available.) ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 287 Material examined: None. HlPPASTERIA PEDICELLARIS DJAKONOV, 1950 Djakonov, 1950: 1950: 54 (1968: 45); 1952: 410. 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. HlPPASTERIA PHRYGIANA (PARELIUS, 1768) HlPPASTERIA PHRYGIANA PHRYGIANA (PARELIUS, 1768) FIGURE 6A-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 piano) Forbes, 1841: 125 (as Goniaster equestris) Midler & 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: K128; 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; Doder- lein, 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, pi. 1, figs A, G; 1959: 136, fig. 21; 1960: 61, pis. 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 Katte- gat, 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 20th 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 synony- mized 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 charac- ters 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 E46606, 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 ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 288 C. MAH ETAL. (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 E46613 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 E13586, north-west of Amundsen Sea, South- east 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 E43921, 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). HlPPASTERIA PHRYGIANA ARGENTINENSIS BERNASCONI, 1961 Bernasconi, 1961: 1; Clark & Downey, 1992: 248 Occurrence: Northern Argentina. 108?162 m. Material examined: None. HlPPASTERIA PHRYGIANA CAPENSIS MORTENSEN, 1933 H.L. Clark, 1923: 270; 1926: 13 (as Hippasteria phry- giana) Mortensen, 1933: 245; A.M. Clark, 1952: 170, 196 (as Hippasteria phrygiana var. capensis) AM. Clark & Courtman-Stock, 1976: 63; Clark & Downey, 1992: 249 (as Hippasteria phrygiana capensis) Occurrence: South Africa, 310?980 m. Material examined: None. Material examined: None. HIPPASTERIA SPINOSA VERRILL, 1909 Verrill, 1909: 63; Fisher, 1911: 224; HL Clark, 1913: 194; Goto, 1914: 349; Verrill, 1914: 301; Djakonov, 1950: 53 (1968: 45); Baranova, 1957: 162; Alton, 1966: 1702; Carey, 1972: 38, 39; Lambert, 1978a: 4, 14; Maluf, 1988: 34, 118; Clark, 1993: 259; Lambert, 2000: 64. 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 Leptast- erias 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 Alba- tross, 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 l.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 ll.viii.1980. (1 dry spec. R = 7.1, r = 3.5); USNM E10504. South-west of mouth of Columbia River, Oregon. 46?8'N 124?30'W, 137 m. Coll. R/V Cobb 14.1.1964. (1 dry spec. # = 12.8, r = 7.1). HIPPASTERIA TASMANICA MCKNIGHT, 2006 McKnight, 2006: 97 HIPPASTERIA PHRYGIANA STRONGYLACTIS H.L. CLARK, 1926 H.L. Clark, 1926: 13; A.M. Clark, 1952: 196; A.M. Clark & Courtman-Stock, 1976: 63; Clark & Downey, 1992: 249 Occurrence: South Africa. 320?980 m. Occurrence: South Tasman Rise, south of Tasmania, Australia. 935-1058 m. Material examined: None. OUTGROUP TAXA: Cladaster analogous. USNM E38569. 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 ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 289 Islands, California. 33?17'N, 118?24'W, 86.0 m. Coll. USFC Albatross, 8.iv.l897. (1 dry spec. R = 6.6, r = 3.3). Peltaster placenta. USNM E12587 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). DISCUSSION COMPARISONS WITH OTHER TAXA A full summary of taxonomic changes is listed in Appendix 4. Sthenaster's overall appearance is inter- mediate between Evoplosoma and Hippasteria, pos- sessing the toothed pedicellariae and sharp spinelets of the former in conjunction with the pedicellariae, short arms, broad disk, and massive appearance of the latter. The presence of sharp spinelets and toothed pedicellariae is not observed in other gonias- terid genera with the exception of Lydiaster and certain species in its sister genus, Circeaster (Mah, 2006). Circeaster has been observed sitting on coral skeletons, possibly feeding (Mah, 2006) suggesting that it may also share corallivorous habits with the Hippasterinae. Also similar to hippasterines are Calliaster and those Goniasteridae related to Calliaster, such as Mabahissaster and Milteliphaster (Aziz & Jangoux, 1985b), which share the large concial spines, present on the abactinal, marginal, and actinal surface, bare marginal plates, and toothed pedicellariae. Chave & Malahoff (1998) observed Calliaster pedicellaris feeding on a deep sea gorgonian in the Hawaiian Islands region. Thus, three groups, the Hippasteri- nae, the Circeaster lineage, and the Calliaster 'group' share similar sets of morphological characters with preliminary evidence suggesting coral or cnidarian predation occurring nominally in each group. It is unclear if these characters are support for these dif- ferent taxa as a single lineage, or are perhaps corre- lated with corallivory, occurring in parallel amongst different goniasterid lineages that have come to exploit an identical food source. Cladaster was not supported as a member of the Hippasterinae (Fig. 2). However, its perceived similar- ity to hippasterines as suggested by Fisher (1911) and its sister group relationship to the Circeaster clade (Mah, 2006) suggest that it may occupy an intermedi- ate or close phylogenetic position to these two taxa. FOSSILS, BATHYMETRIC, AND BIOGEOGRAPHICAL PATTERNS Bathymetric relationships mapped onto the tree (Fig. 11) support an 'onshore-offshore' hypothesis as outlined by Jablonski & Bottjer (1988, 1990). The more derived Evoplosoma is present in deep water relative to hippasterines, such as Gilbertaster and Cryptopeltaster, which occur at relatively shallower depths. This pattern is observed in several other asteroid taxa including the Brisingida (Mah, 1998b), the Zoroasteridae (Mah, 2007a) as well as gonias- terids in the Pentagonasterinae (Mah, 2007b). The fossil 'Hippasteria' antiqua Fell, 1956 supports the possibility that the hippasterines have occupied a wide geographical range in the past. Morphological characters, such as body shape, quadrate marginals, and the larger primary plates, are similar to more basal hippasterines such as Cryptopeltaster and/or Sthenaster, suggesting close relationships with these taxa. Given the geographical disparity between these taxa (west coast of North America, south-eastern United States) and H. antiqua (New Zealand), extinc- tion of widely distributed hippasterines may provide one possible explanation for the distribution observed within the Hippasterinae. In contrast, there is some suggestion that 'offshore- onshore' relationships may be present within the genus Hippasteria. All available proposed sister taxa to Hippasteria occur in relatively deep water (> 200 m) but several Hippasteria species occur at depths as shallow as 20 m (e.g. H. phrygiana, H. spinosa). This suggests that these shallow species may be more derived relative to those occurring at deeper depths, such as H. californica. Similar patterns have been observed in other goniasterids (e.g. Mah, 2005b). Several species, such as H. colossa, H. kurilensis, and H. derjungi, from the North Pacific and Arctic regions probably represent a species complex with ties to the North Atlantic H. phrygiana. This rela- tionship is consistent with relationships presented herein and with previous hypotheses (e.g. Nesis, 1961). Close affinities on the tree between H. spinosa and H. phrygiana may reflect a close bio- geographical relationship reflecting glaciation- related, relatively recent diversification and gene flow via the Arctic, similar to that observed in other asteroids, such as Leptasterias (Foltz etal., 2007) and Asterias (Wares, 2001). Franz etal. (1981) speculated on nominal relationships amongst several Atlantic-Arctic-Pacific taxa, including Cera- master, Leptychaster, and Pseudarchaster. More problematic is the assessment of Clark & McKnight (2001) that H. phrygiana occurs both in the North Atlantic and the South Pacific in and around New Zealand. This could possibly be a wide-ranging taxon with conserved morphology and genetic differ- entiation. It could alternatively be representative of cryptic species similar to what was observed amongst populations of the deep-sea Zoroaster fulgens (Howell etal, 2004). The sister group relationship (Figs 1, 11) between the tropical Atlantic G. caribea and the Pacific G. ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 290 C. MAH ETAL. Figure 10. Hippasteria phrygiana USNM E46606.A, abactinal view; B, actinal view; C, lateral view; D, H. phrygiana USNM 5235. Actinal surface showing bivalve pedicellariae; E, Hippasteria californica USNM E10413. Abactinal plates internal view; F, H. californica USNM 33354 abactinal surface pedicellariae; G, USNM E10413. Lateral view showing marginal plate series. Scale bars: A?C, G = 5 mm; D, E = 2 mm; F = 1 mm. anacanthus (Hawaii to New Zealand) suggests former continuity between the two Gilbertaster populations, perhaps before the closure of the Tethys Seaway (Smith, Smith & Funnell, 1994) by the formation of the Panamanian isthmus. This relationship has also been observed in the deep-sea goniasterid Circeaster between the tropical Atlantic Circeaster americanus and the Hawaiian Circeaster pullus (Mah, 2006). The sister group relationship between Gilbertaster and the Pacific Cryptopeltaster would suggest a Pacific ancestor for Gilbertaster. THE HIPPASTERINAE AND CORALLIVOROUS GONIASTERIDAE Deep-sea corals form habitats that are host to a variety of associated fauna, including fishes and invertebrates. The importance of these habitats to fishes has been investigated in a number of regions (Huesbo et al., 2002; Costello et al., 2005; Stone, 2006; Ross & Quattrini, 2007). However, habitat utilization by invertebrates is not fully understood. Several recent investigations conducted in both the eastern ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 291 OG: Mediaster aequalis OG: Cladaster analogus OG: Peltaster placenta Hippasteria californica Washington to S. California 110-2000 m Hippasterias heathi Alaska, Aleutian Islands 377-454 m Hippasteria spinosa Aleutian Islands to California 10-512 m Hippasteria phrygiana North & South Atlantic/Pacific 10-860 m a NW Gulf of Mexico, 2060-2105 m Rodriguez Seamount & NE Pacific, 730-1842 m Davidson Seamount NE Pacific, 2669 m N. Atlantic, Rockall Trough, 1600-1900 m tS3 I" ?Sthenaster emmae n. gen. n. sp. S. Carolina, tropical Atlantic 110-2000 m - Cryptopeltaster lepidonotus Aleutian Islands to Chile, 360-1244 m 5 changes Gilbertaster caribaea W. Tropical Atlantic 360-1244 m Gilbertaster anacanthus Hawaii to New Zealand 462-913 m Gilbertaster Figure 11. Biogeographical and bathymetric ranges mapped onto the phylogenetic tree. Evoplosoma occurs in a distinctly deeper depth range (indicated by the shaded box) than any of the other hippasterines. Sampled Evoplosoma spp. were Atlantic and Pacific (Evoplosoma includes additional species from the tropical Indian and Pacific Oceans) but all occur in this depth range. OG, outgroup. and western North Atlantic involving quantitative surveys of mega- and macroinvertebrates report diversity to be extremely high (Jensen & Frederiksen, 1992; Mortensen etal, 1995, Jonsson etal, 2004, Reed etal, 2005, 2006; Henry & Roberts, 2007; Roberts etal., 2008). These recent surveys, however, often report only a fraction of the diversity associated with these habitats because sampling methods uti- lized target only a portion (either size- or taxon- specific) 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 collec- tions 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 ecologi- cal interpretations. Several papers provide vague entries (unidentified asteroid, Mortensen etal., 1995; Asteroidea, unid. sp., Reed, Weaver & Pomponi, 2006) or have identified individuals only to the generic level (Mortensen etal., 1995; Jonsson etal., 2004). Few report species-level identifications (e.g. Jensen & Fre- deriksen, 1992; Henry & Roberts, 2007). Much remains to be learned about the megafauna associated with deep-sea coral habitats. Large, promi- nent animals such as the Asteroidea have been shown to have ecologically complex relationships with cni- darians 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 deep- sea coral. Subsequent observations have now shown that at least one species of every genus included in Hippasterinae (Goniasteridae), except for Gilber- ts) 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 292 C. MAH ETAL. taster, 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 impe- rialis 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 etal., 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 sea- mounts 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 dis- tinctive 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 hippas- terines. If Sthenaster is indeed a predator on gorgo- nians, 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 shallow- water 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 nudi- branch Armina sp. (Lambert, 2000). Additionally, H. spinosa elicits an escape response in the sea anemone Stomphia sp. (Lambert, 2000). Similar to its north- west Pacific congener, H. phrygiana has been reported to incite swimming behaviour in the Atlantic sea anemone, Stomphia coccinea (Miiller, 1776) (Robson, 1961). Hippasteria phrygiana preys upon Metridium senile (Linne, 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, mol- luscs, 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 non- hippasterine 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, 1970a, 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 investiga- tion 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 phytog- eny could be important particularly with regard to management and conservation of these ecosystems. ACKNOWLEDGEMENTS The authors of this study are indebted to Cynthia Gust Abeam (deceased), collection manager for the NMNH Echinodermata Collection, and Cheryl Ames (UNCW, National Systematics Laboratory, NMFS) for curatorial assistance. Steve Cairns, curator of Coelenterates, NMNH identified the gorgonian spi- cules. Dave Pawson, Paul Greenhall, and Linda Ward (NMNH) provided additional logistical and technical support. Mary Catherine Boyett (MCZ) prepared the loan of type material from MCZ. David Clague and James Barry (MBARI) made recent collections avail- able. Support for sampling off the south-eastern coast of the US was provided by grants from NOAA Office of Ocean Exploration (S.W Ross, lead Investigator). The science party, the crew of the R/V Seward Johnson, and the pilots of the Johnson Sea Link are greatly acknowledged for assistance with collection of western Atlantic material. We thank the crew of the R/V Western Flyer and the pilots of the ROV Tiburon for their assistance with Pacific collections as well as the David and Lucile Packard Foundation for support of MBARI. Michael Carlson prepared the map for Atlantic collections in Figure 1. We also thank Chris- topher Kelley, Hawaiian Undersea Research Labora- tory, for making in situ observations available to us. Daniel Blake, University of Illinois at Urbana Cham- paign, Andy Gale, University of Portsmouth, and an anonymous reviewer provided useful comments that improved the manuscript. C. 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Disseiiation Abstracts 18 (2): 707-708. ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 297 Taxon/Node APPENDIX 1 DATA MATRIX 11111111112 22222222233333333334444444 12 34567890123456789012 3456789012 3456789012 3456 OG: Mediaster aequalis OG: Peltaster placenta OG: Cladaster analogus Hippasteria californica Hippasteria heathi Hippasteria spinosa Hippasteria phrygiana Sthenaster emmae Evoplosoma virgo Evoplosoma scorpio Evoplosoma claguei Evoplosoma voratus Cryptopeltaster lepidonotus Gilbertaster caribaea Gilbertaster anacanthus 000000 011100 010100 111111 111111 111111 111111 131103 110103 110111 110111 110121 111112 121202 121202 001000 101000 001000 110100 110100 110100 110100 111100 101111 111100 101100 101100 102112 102112 002112 000000000 000000010 001200010 111122110 111122110 111122110 111122110 002200131 000011111 112211110 111111010 111111010 110011010 000011021 000011021 01000 01010 01010 00110 00010 00010 00010 11030 11110 OHIO OHIO OHIO 02010 12021 12021 000000 000000 000101 111100 111100 111100 111100 011001 011111 111211 000111 010111 100100 000000 000000 00001 00000 00000 00011 00011 00011 00011 00011 01100 01100 01100 01100 10011 00011 00011 000000000 000000001 100000001 011111101 010011101 010011101 010011101 011011101 101001102 111001102 110001102 111001102 010011111 020002010 020002010 APPENDIX 2 2.1. CHARACTER LIST 2.2. 1.1. Pulpy tissue in body wall. (0) absent, (1) present (Figs 7A, 8A). 2.3. 1.2. Abactinal fasciolar channels. (0) channels well developed, (1) channels shallow, (2) channels absent, (3) channels wide (Fig. 4A, E, H). 2.4. 1.3. Secondary plates. (0) absent, (1) present (Fig. 10E). 1.4. Abactinal plate morphology. (0) tabular, (1) flat 2.5. and platform like (Fig. 4A, E), (2) low (Fig. 3C, E). 1.5. Abactinal primary structure, large. (0) absent 2.6. (Fig. 8B), (1) big spine (Figs 6A, 7C, 10A, C), (2) tubercle present. 2.7. 1.6. Accessory fringe on abactinal plates. (0) granu- lar fringe, (1) spiny fringe (Fig. 10F), (2) 2.8. angular fringe (Fig. 5B, D), (3) fringe absent. 1.7. Carinal series. (0) clearly distinguished, (1) poorly distinguished (Fig. 4A). 2.9. 1.8. Abactinal spinelets. (0) absent, (1) present. (Figs 4E, 9C, 10A, C) 1.9. Large, coarse granules. (0) absent, (1) smaller, 2.10. (2) larger (Figs 3B, C, F, 5B,D). 1.10. Abactinal plate articulation. (0) moderately well articulated, (1) tightly articulated (Fig. 4F). 2.11. 1.11. Surface granulation flush between granules. (0) not forming flush surface, (1) forms flush 2.12. surface with other granules. (Fig. 3B, C, E) 1.12 Granular distribution. (0) incomplete cover to absent, (1) complete dense granular cover 3.1. (Fig. 9C), (2) coarse granules, widely spaced. Large superomarginal spines. (0) absent, (1) present. (Figs 5D, 6A, D, 10A-C) Large inferomarginal spines. (0) absent, (1) present. (Figs 5D, 6A, D, 10B-C) Superomarginal surface. (0) granules form com- plete cover (Fig. 9C), (1) bare to partial cover, (2) widely spaced. (Fig. 10A, B, C, G) Inferomarginal surface. (0) granules form com- plete cover (Fig. 9C), (1) bare to partial cover (Fig. 10B, C), (2) widely spaced. (Fig. 10B, C, G). Superomarginal plate shape at inter-radius. (0) wide (Fig. 4H), (1) quadrate (Fig. 10A-C), (2) rounded (Fig. 10G). Inferomarginal plate shape at inter-radius, wide, (1) quadrate, (2) rounded (Fig. 10G). Spinelets on marginal plates. (0) absent, present (Figs 4H, D, 90, 10G). Fasciolar grooves between marginal plates. (0) well developed, (1) shallow (Fig. 10C, G) absent, (3) wide (Fig. 4C). Differentiated superomarginal present (Fig. 10G), (1) poorly differentiated (Fig. 3C). Differentiated inferomarginal present (Fig. 10G), (1) poorly (Fig. 3C). Surface granule size. (0) absent (Fig. 9D), (2) larger (Fig. 3C). Marginal plate series calcification. (0) moderate to heavily calcified (Fig. 10A-C), (1) lightly developed (Fig. 9A). Actinal fasciolar groove. (0) well developed, (1) shallow, (2) absent, (3) wide (Fig. 4C, D). (0) (1) (2) fringe. (0) fringe. (0) differentiated (1) smaller ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 298 C. MAH ETAL. 3.2. Large pedicellariae on first adambulacral. (0) 7.1. absent, (1) present (Fig. 3D, G). 3.3. Large actinal spines. (0) absent, (1) present 7.2. (Fig. 61)). 3.4. Actinal granules. (0) present (Fig. 3D, G), (1) 7.3. absent. 3.5. Actinal spinelets. (0) absent, (1) present 7.4. (Figs 4C, D; 6D; 9B). 4.1. Subambulacral spines: abundance-size relation- 7.5. ship. (0) several, smaller, (1) fewer larger, (2) many (Figs 6B, C, 8D,E). 7.6. 4.2. Furrow spines in cross-section. (0) round 7.7. (Fig. 5C), (1) angular (Fig. 6C). 4.3. Furrow spines compressed. (0) absent, (1) 8.1. present (Fig. 6B, C). 4.4. Pedicellariae replacement of furrow spines. (0) all furrow spines present (Fig. 61)), 8.2. (1) pedicellariae replaces furrow spines (Fig. 5C, E). 8.3. 4.5. Furrow spines with rough tips. (0) smooth, (1) rough (Fig. 6B, C). 8.4. 4.6. Subambulacral composition. (0) spines only, (1) spines plus pedicellariae (Fig. 7E). Enlarged pedicellariae. (0) absent, (1) present (Fig. 3C, 0, G, 5B, D, E). Bivalve pedicellariae. (0) absent, (1) present (Figs 3C, D, G, 5B, D, E). Flat-paddle like pedicellariae. (0) absent, (1) present (Fig. 10F). Pedicellariae abundance. (0) absent to few, (1) abundant, (2) very abundant (Fig. 3C). Serrated teeth on pedicellariae. (0) absent, (1) present (Figs 4F, 6C). Triangular pedicellariae. (0) absent, (1) present. Pedicellariae on raised base. (0) absent, (1) present (Fig. 4E). Supermarginal plate facing on abactinal surface. (0) dorsal facing, (1) lateral facing (Fig. 5A, 0), (2) dorsolateral facing. Body-disk swollen. (0) weakly so, (1) strongly so (Fig. 4A, B). Granules, large & flattened. (0) absent, (1) present (Figs 5C, 0). Arm dimensions. (0) moderate to narrow, (1) broad to short (Fig. 4A, B), (2) narrow and tapering. APPENDIX 3 ARTIFICIAL KEY TO INCLUDED TAXA (INCLUDING TERMINOLOGY) Pulpy tissue covering abactinal plates. Abactinal plates tightly articulated with secondary plates. Fasciolar grooves between plates often shallow to absent. Body coelom swollen, thick. Large primary conical spines, coarse granules and/or pedicellariae (equivalent to diameter of plate base) present on abactinal, marginal, and actinal plate surface. Bivalve pedicellariae are common. Marginal plates quadrate to ovalate, often bare but surrounded by peripheral accessories. Furrow spines often thickened, large and relatively few in number (two to four in most). Adults often large with heavy skeletons, body shape is frequently stellate but smaller individuals can be weakly pentagonal (but never pentagonal as adults) Hippasterinae (proceed to 0/0a) Pulpy tissue absent. Abactinal plates not tightly articulated. Secondary plates present or absent. Paxillae or tabulae present or absent, often with well-developed fasciolar grooves present or absent. Large conical spines and pedicellariae may be present but not on all plate surfaces and not generally observed together. Marginal plates variable but may be bare to completely covered by granules. Furrow spines may number from few (two to three) to very high (over ten) per plate. Body skeleton more dorsoventrally flattened. Marginal plates abutting over midradius present or absent. Body shape can be completely pentagonal or stellate as adults Other Goniasteridae (0) Spines and/or spinelets absent from abactinal, marginal, and actinal plate surface (Fig. 3A, E). Secondary plates absent. Marginals never bare ? always covered by closely articulated, blunt, flattened granules (Fig. 3C). Large, bivalve pedicellariae, abundant, flush with surface (Fig. 3A, C, D, G). First adambulacral with enlarged bivalve pedicellariae on entire length of plate (Fig. 3D, G), superceding the furrow spines. Two to three furrow spines per plate Gilbertaster Abactinal plates with one to five smaller plates enclosed by peripheral plates (Fig. 3E, F). Multiple, smaller subambulacral spines present (Fig. 3G). R : r = 2.68. Tropical Pacific: Hawaii, New Zealand. Gilbertaster anacanthus Fisher, 1906 (Fig. 3E-G) Abactinal plates dominated by pedicellariae, small tubercles, and ruffled processes (Fig. 3A, B). None enclosed by peripheral plates. Single triangular wedge-like subambulacral spine present (Fig. 3D). R : r = 2.5. Tropical Atlantic: off Florida/Carolinian coast Gilbertaster caribaea (Verrill, 1899) (Fig. 3A?D) (0a) Spinelets or large, prominent conical spines present on most abactinal, marginal, and actinal plate surfaces (e.g. Figs 6A, D, E, 10B, C). Secondary plates present (Fig. 10E). Marginals completely bare (Fig. 10B, C) or sometimes partially covered by spinelets (Fig. 6D) or granules. Up to ten furrow spines per plate, but often fewer than four, often thick and round in cross-section but sometimes compressed (Fig. 6C) (1) ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 299 (1) Abactinal, marginal, actinal plates covered by small, conical spinelets (Fig. 4A, B, C, D, E, H). No single large spines present (Fig. 4A, B). Marginal plates at inter-radius wide in outline (Fig. 4H) Sthenaster emmae (Fig. 4A?H) (la) Abactinal, marginal, actinal plates may or may not be covered by large conical spines (Figs 6A, 7A, C, 10A, C). Spinelets present or absent. Marginal plates at inter-radius equidistant quadrate to ovaloid in shape (2) (2) Tight fitting, angular granules cover abactinal (Fig. 5B, D), marginal (Fig. 5D), actinal plates (Fig. 5E). Prominent spines on marginal plates (Fig. 5D). Large pedicellariae adjacent to adambulacral plates (Fig. 5E) Cryptopeltaster lepidonotus (Fig. 5A?E) (2a) Tight fitting, angular granules absent. Large pedicellariae present or absent (3) (3) Marginal plates large, heavily calcified, facing dorsally (Fig. 10A) or are visible dorsolaterally. Large bivalve pedicellariae present with smooth blades (Fig. 10D). Spines typically blunt or conical ((Figs 10A?C). Furrow spines enlarged, rounded in cross-section (Fig. 10B). Body wall thick, swollen Hippasteria (Fig. 10A?G) (3a) Marginal plates more lightly calcified, facing laterally (Figs 6A, 7A). Smaller palmate tong-like pedicellariae, with teeth on blades (Fig. 6C). Sharp spines and spinelets on abactinal, actinal surface (Fig. 6A, D). Furrow spines small and numerous, often angular to quadrate in cross-section with furrows (Figs 6B, C, 9B). Body wall relatively thin Evoplosoma (Figs 6?9) (4) Spinelets present on marginal plates, sometimes in abundance (5) (4a) Spinelets absent from marginal plates, which are largely bare of accessories (Figs 7B, 8C), except for pedicellariae and blunt granules (8) (5) Abactinal, marginal, and actinal plate surfaces covered by densely packed, spiny granules (Fig. 9A, C). Larger, prominent abactinal, marginal, and actinal intermediate spines absent E. virgo Downey, 1982 (Fig. 9A?D) (5a) Abactinal, marginal, and actinal plate surfaces not covered by densely packed granules. Large, prominent abactinal, marginal and actinal intermediate spines present (6) (6) Overall disk large, gradually tapering into arms. Adults (R = 11.0 cm) with 70?90 marginal plates per inter- radius. Known only from Indian Ocean E. augusti Koehler, 1909 (6a) Overall disk smaller relative to arms, which are tapering and more sharply set off from disk. Adults (R < 8.0 cm) with 60?66 marginal plates per inter-radius. North Atlantic and Hawaii (7) (7) Furrow spines five to seven, mostly six or seven on adambulacrals proximal to mouth. Spinelets on marginal plates cover surface of plate (Fig. 6D). Approximately 60?66 superomarginals per inter-radius (approximately 30-33 from midline to tip). North Atlantic E. scorpio Downey, 1981 (Fig. 6A-E) (7a) Furrow spines three to five (six on first, five on second, three or four on rest). Spinelets do not evenly or completely cover surface of supermarginal plate (conical or pointed granules may be clustered, however). Approximately 30 superomarginals per inter-radius (approximately 15 from midline to tip). Hawaiian Islands (based on the size of the specimen described, this is possibly a smaller/sub-adult specimen. Larger adult specimens may show character variation.) E. forcipifera Fisher, 1906 (8) Abactinal, marginal, actinal surface covered by prominent cover of thickened, blunt spines evenly distributed over body. Furrow spines five to six. Superomarginal plates per inter-radius -50?70 Indo-Malaysian region E. timorensis Aziz & Jangoux, 1985 (8a) Body surface covered by short, round bullet-like tubercles or prominent, conical spines (Fig. 7A). Superomarginal plates per radius per inter-radius -58?60. Rodriguez Seamount & Islas Tres Marias (Mexico). North Pacific region E. claguei sp. nov. (Fig. 7A-F) (8b) No prominent conical spines on abactinal plates (although tubercles are present). Abactinal plates are mostly bare of larger, primary spines or tubercles (Fig. 8B). Superomarginal plates per radius per inter-radius -44?45 (greater number of smaller-sized inferomarginals = 54?55 per inter-radius). Superomarginals larger, fewer than infero- marginals. More rounded, ovalate in shape (Fig. 8C) Davidson Seamount, North Pacific region E. voratus sp. nov. (Fig. 8A-F) TERMINOLOGY USED IN KEY Abactinal - The surface above the superomarginal series that includes the primary circlet, carinal, and dorsal surface plate series, as well as accessories and primaries that sit on those plates. Accessories - Smaller structures, relative to the pri- maries, such as spinelets and granules, that are mounted on or adorn a plate surface. Accessories are primarily mounted on other plates and are smaller than the plate they adorn and as such, occur in large numbers compared to primaries, described below. Actinal ? The surface below the inferomarginal series that includes the mouth, actinal, oral, and adambu- lacral plate series, as well as all accessories and primaries, which sit on these plate series. Bivalve pedicellariae - Pedicellariae with two low valves inset in a depression in a plate, often resem- bling a bivalve mollusc. ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 300 C. MAH ETAL. Carinal plate series - Abactinal plates in series along midradial line along abactinal arm surface, often flanked along arm's length by adjacent lateral series. Fasciolar channels - Channels between two rows of plates lined with ciliated epithelium. Fasciolar grooves are most easily distinguished between tall, shaft-like plates, but occur on lower mound-like plates as well. Channels are created between any two series of acces- sories (spinelets, granules, etc.) or plates. Furrow spines ? Series of spines present on furrow margin of adambulacral plate. Granules - A small round, hemispherical or irregular grain of calcite present on the plate surface. (Granules are frequently lost during preservation or rubbed off specimens over time.) Inferomarginal plate - A plate of the lower of the two marginal series. Marginal plates - A plate of one of the two horizontal series usually defining the ambitus extending to the terminal plate. Facing of the ambitus may be directed dors ally, ventrally, or laterally, depending on the size and shape of the marginal plate series, which varies from taxon to taxon. Marginals can be paired or offset and are often, but not always, equal in number. Pedicellariae - Small pincer-like organs on the body surface, which are variously modified in shape and number of components. Individual pieces are known as blades or valves. Pointed granule - A small rounded granule with a sharp or pointed tip. "R" and "r" ? measurements of the radius as defined from the center of the disk, usually by the anus. "R" is the distance from the center of the disk to the armtip whereas "r" is the distance from the center of the disk to the interradius. The ratio of R:r is often a means of determining overall shape of the body (with 1:1 being pentagonal). Secondary plates - Smaller plates (less than half to one quarter of other plate size) found between abactinal. Spinelets-SimilaT to a pointed granule but more conical at the base with a sharp or pointed tip. Spines - A prominent, externally expressed conical or pointed shaft-like primary structure, usually articu- lated on the surface of a plate. Superomarginal plates - A plate of the upper marginal series. Tong-like pedicellariae (aka alveolar pedicellariae) - Pedicellariae with two slender, or toothed valves, usually opened out and lying in alveolus of matching shape. Tubercle - Swollen, enlarged, blunt, and rounded cal- cified primary structure, parallel to a spine, but spheri- cal to club-like in shape ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301 NEW TAXA & PHYLOGENY OF HIPPASTERINAE 301 APPENDIX 4 SUMMARY LIST OF GENERA, SPECIES, AND SUMMARY NOTES Genus Species Notes Cryptopeltaster Cryptopeltaster Evoplosoma Evoplosoma Evoplosoma Evoplosoma Evoplosoma Evoplosoma Evoplosoma Gilbertaster Gilbertaster Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Hippasteria Sthenaster lepidonotus philippii augusti claguei scorpio timorensis virgo voratus sp. 1 anacanthus caribaea antiqua californica colossa derjungini falklandica heathi imperialis kurilensis leiopelta mammifera nozawai pedicellaris phrygiana argentinensis phrygiana phrygiana phrygiana capensis phrygiana strongylactis spinosa tasmanica Range extended, neotype designated. Synonymized with Cryptopeltaster lepidonotus Type may be lost. New species. Range extended. New species. From Fujikura etal., 2008. From off Java (Indonesia). Range extended to Palau. Species placed into Gilbertaster. Range extended. Cretaceous fossil. Probably not true Hippasteria Part of H. spinosa complex Part of H. spinosa complex Part of H. spinosa complex Part of H. spinosa complex Part of H. spinosa complex Part of H. phrygiana complex Part of H. phrygiana complex Part of H. phrygiana complex Part of H. phrygiana complex Species complex in North Pacific New genus, new species ? 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 266?301