taxonID	type	description	language	source
494A87B9FF834D192A0E11DCFE5E2CA7.taxon	description	(Fig. 1)	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF834D192A0E11DCFE5E2CA7.taxon	materials_examined	HOLOTYPE. — Philippines. MUSORSTOM 3, stn CP 121, 12 ° 08 ’ S, 121 ° 17 ’ E, 73 - 84 m, 3. VI. 1985 (MNHN EcCh 186). PARATYPES. — Philippines. MUSORSTOM 3 (no data) (MNHN EcCh 185). New Caledonia. SMIB 5, stn DW 100, 23 ° 23.9 ’ S, 168 ° 05.4 ’ E, 80 - 120 m, 14. IX. 1989 (MNHN EcCs 10234). ETYMOLOGY. — From the Greek titan, one of a family of mythological giants that ruled the earth until overthrown by the Olympian gods; now, more broadly, one of gigantic size or power, a reference to the great thickness of the arms of this species relative to those of other comatulids. DISTRIBUTION. — Known only from the Philippines and New Caledonia in 73 - 84 m (possibly to 120 m). The specimen lacking data was found in a jar with a label for RV Coriolis stn 116. However, the depth at this station was 804 - 812 m and the accompanying specimens included Asterometra longicirra (Carpenter, 1888), typical of 200 - 300 m, and Comatella nigra (Carpenter, 1888), characteristic of <50 m. The correct label is thus considered lost. RV Coriolis occupied all 54 MUSORSTOM 3 stations in the vicinity of Mindoro Island. DIAGNOSIS. — A species of Comactinia reaching extremely large size, with proximal arms bearing strong alternating articular tubercles and reaching a diameter of 5 mm; cirri up to XXXVI in number, up to 31.4 mm in length with 22 segments; proximal segments of proximal pinnules smooth and cylindrical or rounded rhombic; combs tapering to a point, present to P 5 in large specimens. DESCRIPTION Centrodorsal discoidal, up to 10.0 mm across; polar area flat or slightly concave, smooth, up to 7.6 mm across. Polar area of small specimen with crescentic traces of obsolete sockets. Cirri of large specimens XXVII-XXXVI (up to X regenerating), 16 - 22, maximum length 31.4 mm, in crowded single and partly double marginal row. Proximal cirrals cylindrical; c 1 short; following segments increasing in length to c 5 - 7; L / W of longest cirral 1.0; following cirrals gradually shorter and slightly compressed with L / W becoming 0.8, sometimes wider than proximal cirrals. Distal 2 - 3 cirrals preceding penultimate slightly narrower and more elongated, L / W 1.0 - 1.1; antepenultimate distally polished; penultimate cirral smaller than preceding, L / W 0.9 - 1.1, with erect conical opposing spine; claw curved, longer than penultimate cirral. Small specimen with cirri XXII, 14 - 15, up to 13.2 mm long; c 4 longest, L / W 1.4; antepenultimate cirral with a weak aboral tubercle. Radials visible in interradial angles or completely hidden by centrodorsal. Single tip of basal ray possibly visible under edge of centrodorsal in one interradius of one specimen. IBr 2 short, apposed laterally and flattened aborally, joined by close synarthry (possibly cryptosynarthry). Ibr 1 oblong with diverging lateral margins, partly hidden by centrodorsal in the two large specimens; W / L c. 5.0 in small specimen. Ibr 2 (axil) pentagonal with short diverging lateral margins, or triangular; W / L 1.9 - 3.0. Lateral aboral margins of brachitaxes ossicles and exterior lateral margins of proximal few arm ossicles thickened and covered with superficial pale reticulated stereom. Articulations between brachitaxes ossicles and proximal brachials also sometimes covered with pale yellowish tissue that obscures articulations. Arms 10; anterior arms longer; small specimen with anterior ray length 145 mm and posterior 65 mm. Large paratype with posterior ray length 137 mm; anterior rays broken at 140 mm. Arms of holotype broken at 110 mm or less; longest arm fragment tapering rapidly near broken tip as in posterior arms of small specimen. An apparently anterior arm (remaining ray length 100 mm) is 2.5 mm across at its broken tip. A comparison with the small specimen suggests that the holotype’s intact anterior ray length may have reached 300 mm. Arm ossicles (except brr 1 - 2 and slender distalmost ossicles) with convex lateral margins; middle brachials with thickened distal margins, the combination making each ossicle appear swollen. Brachials from brr 2 - 5 to brr 9 - 13 (br 2 to brr 5 - 7 in small specimen) with well developed alternating articular swellings. br 1 oblong (slightly longer exteriorly in small specimen), united interiorly; W / L 3.0 - 4.4. br 2 longer exteriorly and with P 1 articulation clearly visible in aboral view; W / L 2.6 - 3.1. Exterior lateral margins of both br 1 and br 2 thickened, apposed and flattened against ossicles of adjacent rays; midaboral surface of brr 1 - 2 with low synarthrial swelling. br 3 + 4 short, oblong; syzygial articulation usually sinusoidal; W / L 2.3 - C 3.0; diameter 2.5 - 4.0 mm. One to several following brachials short, oblong; W / L 2.8 - 3.8. Following few brachials cuneate, becoming triangular with smooth, thickened distal margins concave in aboral view (with fine short tooth-like spines in the small specimen). Middle brachials short, triangular, with distal margins as in preceding brachials; W / L 2.8 - 4.1. Some arms of larger specimens distinctly wider in middle than at base, up to 5.0 mm across. Brachials becoming cuneate distally, proportionally not as short as middle brachials; lateral margins not as convex; distal margins neither concave nor thickened; W / L 1.5 - 2.5. First syzygy at br 3 + 4 (br 4 + 5 on one arm of large paratype); second chiefly at br 11 + 12, but varying from br 8 + 9 to br 24 + 25. Following interval widely variable with no uniform pattern; chiefly 4 - 5 (rarely 3, 7 or 8) in small specimen. In the holotype, second syzygies between br 8 + 9 and br 12 + 13 chiefly followed by two to five intervals of 4 - 7 muscular articulations each, with intervals of chiefly 8 - 9 thereafter. In the large paratype, two to three intervals following second syzygies at br 10 + 11 or br 11 + 12 consist of four or five articulations and intervals of 6 - 10 thereafter. Intervals following a second syzygy at br 23 + 24 or br 24 + 25 are chiefly 11 - 12; arm with br 4 + 5 followed by intervals of 17, 17, 8 and 12 muscular articulations. Proximal pinnules decreasing in length from P 1 to P 4 or P 5, robust proximally, slender and flagellate distally. P 1 of up to 52 segments, 33.4 mm long, with up to 13 low, separated, nonconfluent teeth with flat or rounded apical profiles; fully developed teeth often slightly displaced toward one lateral margin; two or three distal pinnulars with diminishing teeth, tapering to a terminal pinnular that lacks a tooth. Proximal pinnulars cylindrical (rounded rhombic in the small specimen), wider than long; middle segments cylindrical and about as long as wide; distal segments preceding comb slightly constricted with L / W 1.5 - 1.7. Large specimens with all segments smooth. P 1 of small specimen with 32 segments and six teeth, 16.1 mm long, with numerous very short spines on lateral margins of pinnulars facing arm tip. P 2 similar to P 1, up to 51 segments, 32.4 mm long, with up to 16 teeth; the longest segments preceding the comb shorter than in P 1, L / W 1.2. P 2 distinctly shorter than P 1 in small specimen. P 3 similar to P 2 but not quite as wide across the base, 44 segments, 26.9 mm, 15 teeth; proximal teeth short and flat-topped; more distal teeth rounded and proportionally taller than on preceding pinnules; distal three pinnulars tapering to point. P 3 on small specimen much shorter and less robust than P 2, 18 segments, 8.4 mm, six teeth (the first rudimentary). P 4 up to 36 segments, 21.5 mm, with 13 teeth (not including two rudimentary initial teeth); small specimen with P 4 of 12 segments, 6.8 mm long, no comb and with distal pinnulars compressed and longer than wide. P 5 with or without comb in large specimens; comb-bearing P 5 of up to 28 segments, 16.5 mm, with eight teeth (including tapering final three pinnulars); P 5 without comb of up to 23 segments, 16.5 mm; proximal pinnulars short and cylindrical as in preceding pinnules; middle segments squarish; distal segments longer than wide, compressed and with fine spines on lateral margin facing arm tip. Middle pinnules longer than those immediately following oral pinnules, of up to 33 segments, 20.3 mm; basal few pinnulars short, following segments squarish with distal adambulacral margins thickened; distal segments slender and elongated, L / W to 2.4. Distal pinnules more slender than middle pinnules, up to 26 segments, 16.1 mm; pinnulars beyond basal few longer than wide; distal pinnulars in small specimen with L / W up to 3.0. Mouth marginal; anus central; tegmen naked. Color of rays (preserved) tan or pinkish tan with short bands of darker faded purple. Cirri and pinnules tan. One large specimen with distal cirrals dark brown; the other with the proximal parts of the rays faded purple. The small specimen is faded pink. DISCUSSION At a maximum diameter of 5 mm, C. titan n. sp. has thicker, more massive arms than any other comasterid. The arms actually appear wider because the distance from the left side of one brachial to the right side of the succeeding brachial is slightly greater than the diameter of each brachial. The species is placed in Comactinia because it shares with the other two members of the genus – C. meridionalis (Agassiz, 1865) and C. echinoptera (Müller, 1841) – the following diagnostic features: 10 arms; IBr 2 series joined by synarthry; first brachial syzygy at br 3 + 4; mouth excentric, and oral pinnule combs consisting of nonconfluent, often off-center, single teeth flattened along the pinnule axis. C. titan n. sp. also shares with C. meridionalis terminal oral pinnulars that taper to a point, and short cirrals of similar length, almost completely lacking any aboral ornamentation. The small specimen of C. titan n. sp. shares with some specimens of C. echinoptera a weak, rudimentary aboral spine on the antepenultimate cirral (Messing 1978; Messing & Dearborn 1990). Apart from its longer cirri with more numerous segments and strongly developed alternating articular tubercles, both of which may derive simply from its much greater size, C. titan n. sp. differs from the other two in having the proximal segments of the oral pinnules smooth and cylindrical or rounded rhombic. In C. meridionalis, these ossicles are spinose, rhombic and sometimes strongly projecting; in C. echinoptera, the basal pinnulars of the oral pinnules bear rounded keels. With the addition of C. titan n. sp., Comactinia becomes the only comasterid genus known from the tropical western regions of both Atlantic and Pacific oceans. Antedon de Freminville, 1811 (family Antedonidae Norman, 1865) is the only other comatulid genus known from shelf depths in both regions, but it is far more widespread, also occurring in European and West African waters. The possibility exists that C. titan n. sp. is convergent rather than congeneric with the two Atlantic species, especially given the relatively generalized features of the genus (i. e., 10 arms, excentric mouth, IBr 2 with a synarthry, and first syzygy at br 3 + 4). However, C. titan n. sp. exhibits no characteristics that place it outside the genus as currently construed.	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF874D152A6414DEFF002926.taxon	description	(Fig. 2)	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF874D152A6414DEFF002926.taxon	materials_examined	HOLOTYPE. — Vanuatu. MUSORSTOM 8, stn DW 1071, 15 ° 37 ’ S, 167 ° 16 ’ E, 180 - 191 m, 4. X. 1994 (MNHN EcCh 189). PARATYPES. — Vanuatu. MUSORSTOM 8, stn DW 1021, 17 ° 43 ’ S, 168 ° 37 ’ E, 124 - 130 m, 28. IX. 1994 (MNHN EcCh 188); stn DW 1086, 15 ° 37 ’ S, 167 ° 16 ’ E, 182 - 215 m, 5. X. 1994 (MNHN EcCh 187); stn DW 1071, 15 ° 37 ’ S, 167 ° 16 ’ E, 180 - 191 m, 4. X. 1994 (MNHN EcCh 190).	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF874D152A6414DEFF002926.taxon	etymology	ETYMOLOGY. — From the Latin squarrosus meaning “ rough with stiff scales ” (Brown 1956), a reference to the scaly roughened appearance in naked-eye view of arm ossicles from brr 6 - 10 to brr 20 - 25.	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF874D152A6414DEFF002926.taxon	distribution	DISTRIBUTION. — Known only from Vanuatu in 130 to about 180 m (possibly 215 m). Two specimens identified as Capillaster multiradiatus (Linnaeus, 1758) by A. H. Clark (1931) from the Kei Islands, Indonesia, in 20 - 85 m, may represent this species (see below).	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF874D152A6414DEFF002926.taxon	diagnosis	DIAGNOSIS. — A small species of Capillaster (ray length up to c. 100 mm) with up to 20 arms; cirri with up to 30 segments, 21.0 mm long; brachials brr 6 - 10 through brr 20 - 25 triangular, distinctly wider than those preceding, swollen and with strongly thickened spinose distal margins, so that the proximal half of the arm appears distinctly roughened, scaly and up to 1.5 times wider than the arm base in naked-eye view; pinnule combs present to about P 16 - 19.	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF874D152A6414DEFF002926.taxon	description	DESCRIPTION Centrodorsal discoidal, often with sloping sides, 4.0 - 5.4 mm across; D / H 3.4 - 4.0. Polar area faintly depressed or convex; larger specimens with five faint interradial swellings radiating from around a shallow central depression. Cirri XX-XXVIII, 23 - 30, up to 21.0 mm, in single or partly double crowded marginal row. Larger specimens have fewer cirri; c 1 short; following cirrals increasing in length to c 5; c 4 with L / W 0.8 - 1.1; c 5 longest, transitional, with oral margin slightly concave and with slightly raised dentate distal aboral margin, L / W 1.3 - 1.8. Following cirrals polished, becoming shorter, compressed, squarish by c 9, shorter than wide thereafter with L / W 0.8 - 0.9. Cirrus slightly tapered distally; distal 2 - 3 cirrals preceding claw A sometimes with L / W 1.0. Raised, dentate aboral distal margin of c 5 - 6 becoming strong curved midaboral spine flanked by one to three pairs of smaller spines by c 7 - 9; flanking spines usually reduced to single pair on distal cirrals, but sometimes disappearing beyond about c 20; strong midaboral spine becoming erect on distal cirrals. Opposing spine erect and sharp; penultimate cirral squarish, smaller than preceding cirrals, without flanking spines; claw longer than penultimate cirral. Distal margin of radials just visible beyond centrodorsal margin, more visible in interradial angles; distal margin concave, slightly concave in smaller specimens. Brachitaxes aborally convex, with or without weak midaboral synarthrial swellings. Ibr 1 short, oblong or shallow U-shaped in aboral view, united and sometimes slightly inflated laterally, with slightly diverging lateral margins; W / L 4.0 - 5.6 (MNHN EcCh 190 with lateral margins united only proximally and converging distolaterally; W / L 3.9). Ibr 2 (axil) almost triangular (lateral margins extremely short), apposed or separated laterally; W / L 1.8 - 2.4. IIBr series chiefly 4 (3 + 4); IIbr 1 short, oblong, united interiorly, sometimes with a few small midaboral spines; IIbr 2 cuneate or almost triangular, longer exteriorly, with a strongly thickened and finely spinose aboral distal margin, and with W / L 2.3 - 3.1. IIbr 3 + 4 pentagonal, the distal angle thickened and finely spinose; W / L 1.8 - 1.9. MNHN EcCh 188 bearing two IIBr 2 on one ray, resembling IBr 2 but smaller; MNHN EcCh 189 with 7 IIBr 4 (3 + 4) and 3 IIBr 3 (2 + 3). Arms 12 - 20; ray length 85 - 102 mm. Some arms sometimes distinctly broader at the base than others. Distal aboral margins of brachials thickened and finely spinose. On arms arising from IBr series, br 1 and br 2 both longer exteriorly; br 1 united interiorly, with W / L 2.9 - 3.2; br 2 longer than br 1, with W / L 2.5; br 3 + 4 oblong, W / L 2.2 - 2.5 and 2.1 - 2.5 mm across. On arms arising from IIBr series, br 1 oblong or cuneate, united interiorly, with W / L 1.4 - 1.9; br 2 + 3 oblong, with W / L 1.7 - 2.1 and 1.8 - 2.4 mm across. br 4 and br 5 usually oblong and no wider than br 2 + 3, sometimes slightly cuneate and wider; W / L 2.0 - 2.6. Following brachials cuneate, becoming distinctly wider, swollen and almost triangular with strongly thickened, spinose distal margins and distinctly darker articulations by brr 6 - 10; W / L 2.3 - 2.6. Widest brachials 1.2 - 1.3 times diameter of arm base, but up to 1.5 times when measured across the combined width of two successive brachials. Brachials gradually diminishing in width beyond about br 20, reduced to same diameter as arm base by about brr 25 - 28; the combination of swollen brachials and dark narrow articulations giving the proximal half of the arm a distinctly roughened, scaly appearance in naked-eye view. Following middle brachials smaller, narrower, less swollen, not as strongly cuneate, with weaker distal thickening; W / L 2.2 - 2.6. Distal brachials short, weakly cuneate; distal margins finely spinose but only slightly thickened; W / L 1.5 - 2.1. Brachials near arm tip almost oblong or squarish. Syzygies on arms arising from IBr series at br 3 + 4, br 11 + 12 to br 14 + 15, and at intervals of 4 - 6 muscular articulations thereafter. On arms arising from IIBr series, syzygies at br 2 + 3, br 10 + 11 to br 20 + 21, and at intervals of 5 - 8 (rarely 9) thereafter. Some arms tend to have two longer intervals (7 - 9) following the second syzygy and shorter intervals (5 - 6) thereafter. P II of up to 33 segments, 14.2 mm, with nine tall triangular teeth. Proximal several pinnulars wider than long with expanded, spinose distal rims; middle segments with L / W 1.8 - 1.9, only slightly shorter than proximal pinnulars, but much narrower and centrally constricted, with distal spines disappearing by mid-pinnule and only a single strong, conical, lateral spine remaining on distal segments preceding comb. Several pinnulars just preceding comb shorter and wider, with L / W 1.6. P 1 sometimes similar to P II, of up to 34 segments, 11 teeth, 11.8 mm long; sometimes slenderer and much shorter, with proximal pinnulars as long as wide by the fourth segment and middle segments with L / W 2.0 - 2.1. P 2 of up to 30 segments, 13 teeth, 6.6 mm. Following several pinnules similarly short but with more elongated middle segments. P 7 up to 22 segments, nine teeth, 6.1 mm; following pinnules gradually becoming longer. Combs present to about P 16 - 19. Middle pinnules of up to 24 segments, nine A. H. Clark (1931) placed the type of C. borneteeth, 7.3 mm; proximal few pinnulars (some- ensis (Grube, 1875) as a synonym of C. multitimes excepting the short first) with expanded, radiatus rather than C. tenuicirrus despite its having spinose distal rims; middle pinnulars with cluster cirrals longer than broad from the fifth onward. of spines on side facing arm tip, L / W 1.8; pinnu- Capillaster squarrosus n. sp. most closely resemlars preceding comb with two or three strong lat- bles C. multiradiatus but differs consistently as eral spines arranged in a transverse row. Distal follows. C. squarrosus n. sp. has a distinctive series pinnules slender, up to 22 segments, 7.4 mm; of swollen triangular proximal brachials (Fig. 2 F, G) pinnulars beyond basal two longer than wide, whereas the same ossicles in C. multiradiatus are centrally constricted and with one or a few lateral short and resemble the following brachials spines, L / W 2.4 - 2.7. (Fig. 2 A). The cirri in C. squarrosus n. sp. consist Disk naked, anus central, mouth excentric. Anal of more cirrals at a shorter length than in C. multicone and surrounding interambulacral area with radiatus. Finally, pinnule combs occur as far as small, scattered papillae in one specimen. P 4 - 10 (rarely P 15) in C. multiradiatus and as far as P 16 - 19 in C. squarrosus n. sp. Two specimens DISCUSSION briefly described by A. H. Clark (1931: 188, Hoggett & Rowe (1986) include seven species in 189; probably in the Zoologisk Museum, Capillaster. C. macrobrachius (Hartlaub, 1890) Copenhagen, but Clark listed no catalogue numbears a small stellate centrodorsal lacking cirri, bers) as C. multiradiatus may represent C. squar- and C. asterias A. H. Clark, 1931 bears a few cirri rosus n. sp., although both were collected in composed of elongated segments that taper to a shallower water than the specimens described fine tip. The remaining species have more than above. One (Danish Kei Islands Expedition, 10 cirri with distal cirrals shorter than proximal stn 40, 25. IV. 1922, 25 m, T. Mortensen coll.) cirrals. Of these, C. sentosus (Carpenter, 1888) has 19 “ unusually stout ” arms that “ increase in and C. gracilicirrus A. H. Clark, 1912 bear more width to about the twentieth brachial, where they than 40 (often more than 60) arms, while the are twice as broad as at the base ”; cirri of 29 - remaining three (C. mariae (A. H. Clark, 1907), 30 segments, 22 mm long, and pinnule combs C. multiradiatus (Linnaeus, 1758) and present to near the arm tips, though “ becoming C. tenuicirrus A. H. Clark, 1912) rarely have more and more reduced ”. The other (stn 53, more than 30. C. mariae differs from C. multira- 8. V. 1922, 85 m) has 15 arms (140 mm; longer diatus in having perfectly smooth brachitaxes than in C. squarrosus n. sp.), and cirri of 31 - (division series) and a mottled disk lacking any 32 segments, 25 - 30 mm long. Its largest cirri are calcareous nodules. However, the distinctions similar to those of several C. sentosus examined, among several of these species remain unclear. but the latter have many more arms (33 - 80). A. H. Clark (1931) mentions five specimens ofLikewise, a specimen of C. sentosus with cirri C. multiradiatus (out of 214) with 40 - 43 arms similar to those of C. squarrosus n. sp. also has and seven specimens of C. sentosus (out of 32) many more arms (60). with fewer than 37 arms. Although cirri are typically ≤ 25 mm long with ≤ 25 segments in the former and greater than this in the latter, he also Genus Cenolia A. H. Clark, 1916 refers to individuals of C. multiradiatus with cirri of up to 27 segments or up to 30 mm long. No TYPE SPECIES. — Comatula trichoptera Müller, 1846. other features are known to distinguish the two. DIAGNOSIS. — A genus of Comasteridae with IBr 2 C. gracilicirrus differs from C. sentosus, and joined by synarthry; first brachial syzygy br 3 + 4 at on all C. tenuicirrus differs from C. multiradiatus, only of two ossicles occurring irregularly; P on br 1 2; pinnule arms; brachitaxes beyond IBr usually 4 (3 + 4), any series in having distal cirrals slightly longer than broad combs usually present as far as P 4, sometimes to P 19, rather than broader than long. However, consisting of pairs of straight teeth of more or less equal size, each confluent with the pinnular margin; teeth of a pair sometimes joined to form a transverse bar; aboral surface of disk not heavily plated; mouth excentric (emended from Rowe et al. 1986).	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF874D152A6414DEFF002926.taxon	discussion	REMARKS Cenolia amezianeae n. sp., described below, differs from all species previously assigned to Cenolia in having combs present on pinnules distal to P 4. Instead, combs occur to between P 8 and P 19, although one or two more proximal pinnules sometimes lack one. It also differs in having usually rod-shaped calcareous nodules on the disk and proximal arm ambulacra, although tegminal deposits vary within the genus, and these may be similar to the small spines reported around the anal cone in C. tasmaniae (A. H. Clark, 1918). C. amezianeae n. sp. shares with the other five members of the genus all other diagnostic features.	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF8B4D10284B177EFF272CA7.taxon	description	(Fig. 3)	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
494A87B9FF8B4D10284B177EFF272CA7.taxon	materials_examined	HOLOTYPE. — South New Caledonia. CHALCAL 2, stn CH 05, 24 ° 44 ’ S, 168 ° 08 ’ E, 223 m, 27. X. 1986 (MNHN EcCh 191). PARATYPES. — South New Caledonia. CHALCAL 2, stn CH 05, 24 ° 44 ’ S, 168 ° 08 ’ E, 223 m, 27. X. 1986 (MNHN EcCh 192, 1). Vanuatu. MUSORSTOM 8, stn CP 970, 20 ° 19 ’ S, 169 ° 53 ’ E, 252 - 310 m, 21. IX. 1994 (MNHN EcCs 10235); stn CP 1018, 17 ° 53 ’ S, 168 ° 25 ’ E, 300 - 301 m, 27. IX. 1994 (MNHN EcCs 10236, 1). ETYMOLOGY. — Named for Nadia Améziane, Curator of Echinoderms at the Muséum national d’Histoire naturelle, Paris, who has contributed substantially to our understanding of crinoid systematics and through whose good offices this collection was made available for study. DISTRIBUTION. — Known only from off southern New Caledonia and Vanuatu in 223 to 300 m. C. amezianeae n. sp. occurs in more tropical waters than other Cenolia, but the gap between its southernmost record and the northernmost previous records is only a few degrees of latitude. Rowe & Gates (1995) recorded Cenolia spanoschistum (H. L. Clark, 1916) and C. glebosus Rowe, Hoggett, Birtles & Vail, 1986, from southern Queensland; McKnight (1977) record- ed C. trichoptera (Müller, 1846) (probably C. spanoschistum according to Rowe et al. 1986) from Norfolk Island in 310 m. DIAGNOSIS. — A species of Cenolia with stout cirri of up to 17 short segments; distal cirrals each bearing a transverse aboral ridge; up to 20 arms; pinnule combs occurring as far as P 8 to P 19, though one or two more proximal pinnules may lack a comb, and with numerous chiefly rod-shaped calcareous nodules on anal interambulacral area, along ambulacra of proximal arms and, sparsely, along genital pinnule ambulacra. DESCRIPTION Centrodorsal discoidal, usually with small, triangular, orally-directed interradial processes, 7.9 - 10.3 mm across; D / H 3.4 - 4.1. Polar area with faint interradial swellings radiating from around shallow central depression. Apical margins of cirrus sockets project above polar area in one specimen. Cirri XXXII-XLI, 14 - 17, up to 15.6 mm long, in crowded double marginal row. c 1 short; following segments increasing in length to c 5 - 6, which have L / W 1.1 - 1.4; following segments progressively shorter, becoming as wide as long by c 7 - 8 and shorter thereafter. c 6 - 7 with distal aboral margin thickened, appearing rounded triangular in lateral view, increasing slightly in strength and moving to middle of segment in side view on distal cirrals, becoming a sharp, transverse ridge on 1 - 2 cirrals preceding penultimate on some cirri. Penultimate cirral smaller than preceding; opposing spine weak or distinct, conical or rounded, occasionally transversely chisel-shaped. Claw longer than preceding segment; tip bluntly rounded. Radials completely hidden by centrodorsal or just visible in interradial angles. IBr 2 short, aborally flattened; Ibr 1 very short, oblong, united laterally, partly hidden by centrodorsal; Ibr 2 (axil) triangular or with very short diverging lateral margins, apposed or just separated laterally, with W / L 2.2 - 2.3. IIBr 4 (3 + 4) present on at least some rays of all specimens (one IIBr 2 on MNHN EcCh 192). IIbr 1 short, oblong, united interiorly; W / L 3.3 - 3.4. IIbr 2 triangular (longer exteriorly), with a strong, distolateral, alternating articular tubercle; W / L 3.0 - 3.9. IIbr 3 + 4 pentagonal with slightly diverging lateral margins and W / L 1.5 - 1.75; each ossicle with a distinct transverse groove across the aboral surface. Specimen MNHN EcCs 10235 bearing six rays: four rays normal; two remaining rays with short IBr 2 crowded together and with adjacent margins closely united and irregular. Arms 14 - 20, mostly broken. (Arm numbers in specimens with rays broken at their bases are estimated by comparing the diameters of terminal remaining ossicles with those of IIbr 2 and br 2). Specimen MNHN EcCh 192 with an almost intact posterior arm (ray length 85 mm); its longest anterior ray broken at 98 mm. Proximal several brachials with strong, rounded, alternating articular swellings. br 1 oblong or weakly cuneate and longer exteriorly, interiorly united proximally (adjacent interior distolateral margins diverging from each other so that br 2 are separated interiorly), with W / L 2.3 - 2.7; br 2 longer exteriorly, with W / L 3.0 - 3.7; br 3 + 4 oblong, with W / L 2.0 - 2.5 and diameter 1.9 - 2.6 mm (on arms arising from IBr 2, W / L 3.2; diameter 2.8 - 3.1 mm); br 3 with a shallow groove running transversely across the aboral surface; brr 5 - 6 oblong, with W / L 2.3 - 2.8. Following brachials becoming strongly cuneate, slightly wider than arm base, and almost triangular by br 9, with W / L 2.7; brachials brr 8 - 10 and following with thickened distal aboral margins. Middle brachials triangular with strongly thickened and very finely spinulose distal aboral margins; W / L 2.3 - 2.7. Distal brachials triangular or strongly cuneate with distal aboral margins finely spinose; thickening of distal aboral margins reduced beyond mid-arm and disappearing distally. First arm syzygy at br 3 + 4; second chiefly from br 11 + 12 to br 14 + 15 (rarely br 7 + 8 to br 17 + 18); following interval chiefly 4 - 6 muscular articulations (rarely 3, 7, 9). P II of up to 52 segments, 17.8 mm long, with 7 - 10 comb teeth; all pinnulars rhombic and wider than long; proximal two teeth developed as transverse filled arches (the transverse bars of Rowe et al. 1986); following teeth erect, confluent, separated and paired; distal three pinnulars diminishing to a blunt tip. P 1 of up to 50 segments, 17.6 mm, with 10 - 12 teeth; comb as in P II but with up to 4 proximal transversely oriented teeth and up to 4 terminal pinnulars tapering to a point. Several comb-bearing pinnulars in a row sometimes having the teeth on one side weaker than on the other; a single confluent tooth rarely developed on the opposite face of a pinnular bearing a normal pair, producing a bipectinate profile (these two features occurring on any oral pinnule). P 2 of up to 34 segments, 10.3 mm, 14 teeth; comb-bearing pinnulars on this and succeeding pinnules flat and transversely broadened so that the teeth of a pair are widely separated and sometimes connected by a low narrow ridge. P 3 the shortest pinnule, bearing a gonad, of up to 26 segments, 7.8 mm, 12 - 15 teeth; third to sixth or seventh pinnulars of this and following pinnules strongly projecting laterally, giving proximal pinnule a strongly serrate profile on the side facing the arm tip. P 4 of up to 28 segments, 10.0 mm, 17 teeth. Following pinnules similar; combs present to between P 14 and P 19 (only to P 8 or P 9 in MNHN EcCs 10236), but one or two more proximal pinnules sometimes without comb. Middle pinnules of up to 30 segments, 8.8 - 11.5 mm, 13 - 14 teeth. Distal pinnules slender; proximal and middle segments with distal spinose rim; no strong lateral projections; pinnulars following the third longer than wide, with L / W up to 1.6. Anus central; mouth marginal; numerous chiefly rod-shaped calcareous nodules on large inflated anal interambulacral area, along ambulacra of proximal arms and with a few along ambulacra of genital pinnules. Nodules also round, irregular, club-shaped or slender conical, and sometimes concentrated on anal tube. DISCUSSION Cenolia amezianeae n. sp. shares with C. spanoschistum transverse aboral ridges (or bars, Rowe et al. 1986) on the distal cirrals and no more than 20 arms. However, C. spanoschistum is apparently a smaller, less robust species with a centrodorsal reaching only 5 mm across (A. H. Clark 1931). Specimens with centrodorsals 3 - 4 mm across (AM J 11075, TAS 212) have mature gonads, 20 arms, ray length 40 - 70 mm, and longest cirral (c 4) with L / W up to 3.0. Unlike C. amezianeae n. sp., the brachitaxes in C. spanoschistum (and its synonym C. novaezelandiae (A. H. Clark, 1918 )) are smooth and lack both alternating articular tubercles and transverse grooves. The holotype of Cenolia benhami (A. H. Clark, 1916) (USNM 38684) has a large centrodorsal (9.5 mm across) similar to that of C. amezianeae n. sp., but has LXVII cirri up to 23 mm long with 24 - 27 segments. Although it has more arms (28) than C. amezianeae n. sp., the ossicles of the brachitaxes are proportionally more elongated (a feature expected of smaller specimens); W / L of IIbr 3 + 4 is 1.4 in C. benhami and 1.8 in C. amezianeae n. sp. Brachitaxes ossicles are smoothly rounded as in C. spanoschistum. Also unlike C. amezianeae n. sp., the distal cirrals in C. benhami bear a curved transverse aboral ridge (that may appear weakly forked in aboral view) or tubercle. The opposing spine is conical or weak. Cenolia tasmaniae (A. H. Clark, 1918) differs from C. amezianeae n. sp. in having more arms (up to 37) at a smaller size; distal cirrals with a small pointed aboral tubercle or rounded hump; brachitaxes without alternating articular tubercles or transverse grooves, and combs only to P 2 or P 3. A. H. Clark (1931) refers to brachitaxes ossicles as everted and finely spinous in C. tasmaniae. Rowe et al. (1986) note that small spines (perhaps similar to the rods in C. amezianeae n. sp.?) usually occur around the anal cone. However, AM J 14791, a specimen of C. tasmaniae similar in size to the holotype of that species (centrodorsal diameter 4.0 mm; cirrus length 9.0 mm), has smooth brachitaxes and a naked disk. Cenolia trichoptera and C. glebosus both bear up to 40 arms. The former has combs as far as P 3 with 6 - 27 teeth consisting chiefly of filled arches (= transverse bars; see Messing 2001: fig. 5 d), sometimes with some distal paired teeth. A large sexually mature specimen (AM J 14408; centrodorsal diameter 7.1 mm; cirrus length 13.0 mm; 36 arms; ray length 125 mm) has smooth brachitaxes with weak alternating articular tubercles on the exterior sides of the IIBr series and on the articulation between br 2 and br 3. The comb-bearing pinnulars are not as wide and the projections on the proximal pinnulars are not as strong as in C. amezianeae n. sp. C. glebosus differs from all other Cenolia in having hemispherical or globular reticulate calcareous nodules on the disk.	en	Messing, Charles G. (2003): Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema 25 (1): 149-162, DOI: 10.5281/zenodo.5402202
