Poraniomorpha hispida M. Sars 1872

Poraniomorpha hispida (M. Sars 1872) Figure 8 A–F Goniaster hispidus M. Sars in G. O. Sars 1872: 28; M. Sars 1877: 72 –75, pl. 8, figs. 24–26; Storm 1878: 253 –254; 1879: 20; 1881: 90. Asterina borealis Verrill 1878: 213; 1879: 14; 1885: pl. 18, fig. 46 Porania spinulosa Verrill 1880: 202 –203; 1882: 218 Poraniomorpha rosea Danielssen & Koren 1881: 189 –192; 1884: 67–70, pl. 10, figs 8–14; Sluiter 1895: 59; Grieg 1896: 12; Petersen & Levinsen 1900: 56; Ludwig 1900: 459 –460; Grieg 1902: 21 –22, pl. 1, figs 1,2; Norman 1903: 408. Porania borealis Verrill 1882: 218 ? Rhegaster murrayi Sladen 1883: 156, pl. 26, figs. 2–7; 1889: 368–371; Bell 1893: 80 Pentagonaster hispidus Danielssen & Koren 1884: 58 –59, pl. 15, fig. 6; Grieg 1895: 6; 1898: 24. Poraniomorpha spinulosa Verrill 1885: 542; 1895: 139. Poraniomorpha borealis Verrill 1885: 551; 1895: 139–140; A.H. Clark 1949: 373 (?) Lasiaster hispidus Sladen 1889: 374; Norman 1893: 347; Pfeffer 1894: 118; Verrill 1899: 198; Ludwig 1900: 460; Grieg 1902: 22 –24, pl. figs. 3, 4; Michailovski 1903: 486; Süssbach & Breckner 1911: 219 –220. Lasiaster villosus Sladen 1889: 372, pl. 58, figs 7–10 Poraniomorpha hispida Østergren 1904: 615; Koehler 1909: 100 –101, pl. 2, fig. 8, pl. 11, fig. 7, pl.23, fig. 7; Mortensen 1912: 258; 1914a: 332–333, Koehler 1924: 157 –159, pl. 5, fig. 9; Grieg 1927: 129 –133, figs. 1–6; Mortensen 1927: 92; Djakonov 1946: 163 –169 (pt.); 1950a: 58 (in key), 59 (1968: 50); Einarsson 1948: 11; Franz et al. 1981: 405, 406 [?Non P. hispida: Gallo 1937 and other references from Portugal = P. (Culcitopsis) borealis]; Haedrich & Maunder 1984: 40 –42; Anisimova & Cochrane 2003: 121. Poraniomorpha hispida var. rosea Østergren 1904: 615; Grieg 1907: 42; Mortensen 1912: 258; 1927: 92, fig. 53; Djakonov 1950: 59 (1968:50). Poraniomorpha (Lasiaster) hispidus Grieg 1907: 40 –45, figs 5, 6 Poraniomorpha (Poraniomorpha) hispida hispida A.M. Clark 1984: 33 –34, 40, figs. 8b, 9A, B, 11D–F; Clark & Downey 1992: 216. Poraniomorpha (Poraniomorpha) hispida rosea A.M. Clark 1984: 34, figs 8c, 11B,C; Dilman 2008: 139. Poraniomorpha hispida rosea Harvey et al. 1988: 163 Synonymy of Poraniomorpha hispida rosea. Harvey et al. (1988: 163) and Clark and Downey (1992: 214) outline the distinctions between the subspecies P. hispida hispida and P. hispida rosea. Separation of Poraniomorpha hispida rosea from P. h i s p i da was based primarily on whether the body shape showed a more distinctly stellate shape with more angular interbrachial arcs and triangular rays, and its occurrence in the Norwegian Basin along the Norwegian Trench and down the Rockall Trough to the Bay of Biscay at 290 to 1400 m depth. Clark and Downey (1992: 214) outline the R/r ratio as ranging between 1.7 to 2.2 with a mean of 2.0 among specimens with R> 2.5 cm. The morphological differences are not universal for this species across the Atlantic as records of deeper water Poraniomorpha hispida from other localities do not show this variation. Poraniomorpha hispida USNM specimens, E 34636 and E34637, show R/r of 1.5 to 1.6 (at R=3.4 and 3.2) from796 and 1990–2020 m, respectively. USNM 12008 shows R/r of 1.4 from 814 m, which does little to support the association of P. hispida rosea with depth. Dilman (2008) reported P. hispida rosea from the Mid-Atlantic ridge at 966–1019 m but with a less stellate shape [R/r ratio of 1.5 (1.8 cm / 1.2 cm)]. Other widely occurring cold-water asteroids (e.g., H. phrygiana) show significant morphological variation (Foltz et al., 2013; Mah et al. 2014). Morphological variation in this subspecies appears to be relatively minor, perhaps attributed to growth or environmental/hydrodynamic influence (e.g. Hayne & Palmer 2013). For these reasons, I have entered P. hispida rosea into synonymy of P. hispida. Occurrence. Norwegian fjords to western Sweden. South from the Arctic Ocean, Newfoundland, south to South Carolina, including Virginia and North Carolina. 100– 2020 m. Description. Body pentagonal (Fig. 8 A) to stellate in outline, (R/r =1.2 to 2.5), arms triangular with rounded tips. Interradial arcs nearly straight to weakly curved. Body thick with flattened actinal surface (Fig. 8 B). Clark and Downey (1992) note R up to 5.5 cm. Body surface covered by a discrete, soft layer of skin over plates. Abactinal surface tumescent. Plates vary in outline from rod-like to irregularly rounded and lobate, tightly arranged into a dense reticulation, centered around papular pores. Round plates articulated with rod-like plates. Spaces between abactinal plates separated by thickened skin. Papular regions more widely present proximally on disk, disappearing distally, with abactinal plates forming a closely arranged mosaic near armtip. More lobate, irregularly shaped, paired plates present bisecting each interradial region forming confluence with marginal plates. Abactinal plates present, bisecting each interradii in nearly all individuals examined (Figs. 8 A, E). Madreporite irregularly round to polygonal, flanked by three or more plates and/or papular regions. Sulci abundant, Madreporite located on interradial bisector. Papulae pores, one to 20, typically in clusters of four to 15, present between reticulated openings throughout abactinal surface (Fig. 8 A, E). Single pores tend to occur more distally adjacent to armtip. Larger, more confluent regions with more numerous pores present more proximally, closer to disk center. Papulae absent from regions where bisecting interradial abactinal plates are present. Abactinal, marginal, actinal surfaces covered by a continuous cover of fine, papilliform spinules which obscure boundaries between plates. Spinules on abactinal and marginal surface densely arranged but becoming more widely dispersed on actinal intermediate surface. Pedicellariae not observed. Marginal plates wide, blocky, weakly convex, approximately 32 to 34 per interradius (from R=2.5 to 5.5) (Fig. 7 C). Marginal plate surface covered by continuous layer of papilliform spinules, obscuring boundaries between plates (Fig. 8 C). Skin layer covering surface between superomarginal and inferomarginal plates. Intermarginal papulae present, one to six, occurring on tissue filled spaces at the contact zones between superomarginals and between superomarginal and inferomarginals (Fig. 8 C). Intermarginal papulae absent from spaces between inferomarginal plates. Intermarginal papulae greatest in number interradially but gradually decreasing and disappearing distally (two to six plates away from arm tip). Superomarginal and inferomarginal plates, similar in size, articulated with one another 1:1. Interradial bisector converges with superomarginal plates at center of interradius. Spines, other prominent features absent from superomarginal plate surface. Inferomarginals form discrete fringe around actinolateral edge. Actinolateral edge of inferomarginal surface composed of seven to 20 flattened, pointed spines which stand apart from continuous papilliform spinules present elsewhere. Second cluster of spines, one to ten present on actinal facing surface of inferomarginals locally becoming confluent with actinolateral cluster. Inferomarginal plates form prominent actinal border Actinal surface flattened, formed from three to four transverse actinal series, extending from adambulacral to adambulacral (Fig. 8 B). Longest, most complete series is most distal, with each series becoming shorter and more irregular as they draw proximally toward the mouth. Spination on actinal surface varies from smaller spinelets to larger spines, which are roughly twice as long as those on marginal and abactinal surface. Spinelets of either size are widely and evenly distributed. Actinal plates imbricate, irregularly rounded to polygonal in outline, but boundaries obscured by spination. Patch of one to 20 large, at least two to three times as long as surface spinules, with pointed tips present on the surface of each actinal plate. Patches with fewest numbers of spines present most proximally, closest to mouth. Actinal regions grooved, forming corresponding fasciolar channels between actinals and inferomarginals. Adambulacral plate surface sharply convex, with rounded edges, tissue present between plates (Fig. 8 D). Furrow spines two, perpendicular to subambulacrals, arranged in parallel along tube foot furrow, each spine round in cross-section with blunt, pointed tip. Subambulacral spines, arranged transversely, up to five per cluster, also round in cross-section with pointed tips, all approximately similar in size. Oral plates with up to five (three to four) furrow spines, the apical one thicker than the oral plate furrow spines and paired with the matching spine of the other oral plate (Fig. 8 D). Up to three to five conical, pointed spines present on each oral plate surface (six to 10 total). Color in life is “rose red”, light to dark orange on abactinal surface, lighter orange on actinal surface. Material examined. North Atlantic. MNHN, no #. South of Newfoundland, 47º23.5’N, 57º48.0’W to 47º23.2’N, 57º54.9’W, 210– 216 m. Coll. ERHAPS 851, Sta. L85. 20 Feb 1985 (1 dry spec. R=2.9, r=2.0); USNM E7940 south of Flemish Cap, Newfoundland North Atlantic Ocean. 47°08’N 44°52’W, Coll. R/V. Atlantis. (1 dry spec. R=5.5, r=3.7); USNME 20648 SE of Casco Bay, North Atlantic Ocean, 43°33’N 69°40’W, 135 m, coll. R/V. Albatross IV (2 dry specs. R=4.7, r=2.9; R=5.6, r=3.3); USNM E34636 Cape Fear, North Carolina, 33º05’ 44”N, 76º 24’ 40”W, 796 m. coll. 22 May 1985 (1 dry spec. R=3.4, r=2.2); USNM E34637 Off Charleston Bump, South Carolina, North Atlantic, 32º10’ 14”N 76º42’50”W, 1990–2020 m. Coll. R/V Gyre, Sept. 18, 1985. (1 dry spec. R=3.2, r=2.0); USNM E38679 East of Virginia Beach, North Atlantic. 36º41’30”N, 74º37’ 24”W, 690– 832 m. Coll. R/V Columbus Iselin, June 1973. (1 dry spec. R=4.6, r=3.2); USNM 6148 Hudson Canyon, North Atlantic Ocean. 39°58’ 35”, 71°00’ 30’W, 360 m, Coll. R/V Albatross (1 dry spec. R=2.1, r=1.2); USNM 120

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