Stylaster campylecus (Fisher, 1938)

Cairns, Stephen D. & Lindner, Alberto, 2011, A Revision of the Stylasteridae (Cnidaria, Hydrozoa, Filifera) from Alaska and Adjacent Waters, ZooKeys 158, pp. 1-88 : 36-40

Stylaster campylecus (Fisher, 1938) Figs 17 A–D18A–M

Allopora campyleca Fisher, 1938: 505-506, pls. 34, 36.- Broch 1942: 101.- Boschma 1957: 19-20.-Not Lowenstam 1964: 382 (probably Stylaster parageus parageus ).

Allopora polyorchis Fisher, 1938: 503-505, pl. 35, fig. 1, pl. 37, 38.- Naumov 1960: 581.- Boschma 1957: 27.

Allopora campyleca tylota Fisher, 1938: 509-510, pl. 41, figs 2-2e.- Naumov 1960: 580.

Allopora moseleyana Fisher, 1938: 512-514, pl. 49, fig. 2, pl. 51 (not pl. 50), pl. 53, fig. 1 (not forma leptostyla).- Boschma 1957: 23.-? Naumov 1960: 577.

Allopora campyleca campyleca .- Naumov 1960: 578.

Stylaster campylecus .- Cairns 1983b: 430.-Not Cairns and Macintrye 1992: 100 (probably Stylaster parageus parageus ).- Brooke and Stone 2007: 523, 524, fig. 2C.-Not Lindner et al. 2008: 3, Table S1 (= Stylaster brochi ).

Stylaster moseleyanus .- Cairns 1983b: 429.- Wing and Barnard 2004: 11, 28, fig. 30.- Heifetz et al. 2005: 124137 (listed).- Stone and Shotwell 2007: 108 (listed).- Jamieson et al. 2007: 224 (listed).

Stylaster polyorchis .- Cairns 1983b: 430.- Cairns and Macintrye 1992: 100-101 (mineralogy).- Heifetz 2002: 22 (listed).- Wing and Barnard 2004: 11, 28.- Heifetz et al. 2005: 134, 137 (listed).- Stone and Shotwell 2007: 108 (listed).- Jamieson et al. 2007: 224 (listed).-Not Lindner et al. 2008: 3, Table S1 (= Stylaster brochi ).

Stylaster campylecus tylotus .- Cairns 1983b: 430.- Wing and Barnard 2004: 27 (listed).- Heifetz et al. 2005: 134 (listed).- Stone and Shotwell 2007: 108 (listed)-Jameison et al. 2007: 224 (listed).

Stylaster campylecus campylecus .- Wing and Barnard 2004: 11, 27, fig. 32.- Heifetz et al. 2005: 134 (listed).- Stone and Shotwell 2007: 108 (listed)- Jamieson et al. 2007: 224 (listed).

Type material.

Allopora campyleca : Holotype: Alb-3480, 1 dry male colony, now in two pieces, length 16 cm, and SEM stub 1524-25, 1545, USNM 42870 (Fig. 17A). Paratypes: Alb-3480, 7 dry colonies and 7 in alcohol, and SEM stub 1526, USNM 43767, 52262, 52265, 58099, and CAS 28297; Alb-2852 (missing in 2011); Alb-2858, 3 dry colonies (not considered conspecific with Stylaster campyleca); Alb-3599, 2 dry colonies (1 male is Stylaster campylecus, 1 female is Stylaster brochi), USNM 76814; Alb-4230, 1 dry male colony (not considered to be conspecific with campyleca); Alb-4302, 2 female dry colonies, USNM 76816. Type locality.Alb-3480: 52°06'N, 171°45'W (Amukta Pass, Aleutian Islands), 518 m.

Allopora polyorchis : Holotype: Alb-3480, 1 large dry colony now in two pieces, and SEM stubs 1527-28, USNM 43266 (Fig. 17B). Paratypes: Alb-3480: about 15 dry branch fragments, USNM 76540-76542, CAS 28293. Type locality.Alb-3480: 52°06'N, 171°45'W (Amukta Pass, Aleutian Islands), 518 m.

Allopora campyleca tylota: Syntypes: Alb-4781, 4 dry female colonies, the largest 15 cm in height, and SEM stub 1539, USNM 43263, and 12 dry male branches, and SEM stub 1540, USNM 86000 (Fig. 17C). Type locality.Alb-4781: 52°14'30"N, 174°13'E (off Agattu I., Near Islands, Aleutians), 882 m.

Allopora moseleyana : Holotype: Alb-4781: 1 dry male colony now in two pieces, largest 13 cm in length, USNM 42869 (Fig. 17D). Paratypes: Alb-4781, 1 dry male colony, USNM 86003; Alb-3480, 3 female and 1 male dry colonies, and SEM stub 1519, USNM 76556, 76557; Alb-3480, 2 dry female and 1 male colonies (considered as Stylaster leptostylus ), USNM 76555. Type locality.Alb-4781: 52°14'30"N, 174°13'E (off Agattu I., Near Islands, Aleutians), 882 m.

Material examined.

Types of Allopora campyleca , Allopora polyorchis , Allopora campyleca tylota , and Allopora moseleyana ; Alb-3480, numerous branch fragments (topotypic), USNM 76565; Alaskan Leader-35, 53°01'42"N, 170°05'59"W, 200-400 m, 4 Jun 2000, 1 female, USNM 1122510; Alaskan Leader 64, 53°07'N, 166°53'42"W, 324-766 m, 21 Jun 2002, 1 female, USNM 1122484; Alb-4781, type locality, 1 female, USNM 76700; Ballyhoo, 54°47'20"N, 178°46'53"E, 254 m, 5 Jun 2000, 1 female, USNM 1122535; Ballyhoo, 54°42.44'N, 178°46.83'E, 250 m, 5 Jun 2000, 1 female, AB00-0013; Delta, 51°50'55"N, 179°49'26"W, 125 m, 17 Jul 2002, 1 indet., USNM 1123463; Delta Alfa, western Aleutian Islands, depth unknown, 1961, 1 female, USNM 52447; Dominator 971-181, 51°27'43"N, 176°38'20"E, 384 m, 27 Jul 1997, 2 female, 2 male, USNM 1123363; Jason 2095-7-7, 51°48.693'N, 173°49.965'W, 843 m, 25 Jul 2004, 1 male, AB09-0013; Jason 2102-7-8, 51°30.56'N, 177°55.36'W, 824 m, 2 Aug 2004, 6 male, AB09-0012; Jason 2101-9-1, 51°31.43'N, 177°57.5'W, 494 m, 2 Aug 2004, 1 male, AB09-0015; Jason 2103 –15– 1, 51°50.968'N, 179°51.007'E, 678 m, 1 male, AB08-0037; Jason 2104-1-5, 51°43.83'N, 179°36.0'W, 1011 m, 5 Aug 2004, 1 female, 1 male, AB08-0038 and AB09-0016; MF 833-38, 51°42'54"N, 178°51'30"W, 582 m, 3 Aug 1983, 1 male in alcohol, USNM 77055; MF 833-47, 51°55'36"N, 176°52'48"W, depth unknown, 5 Aug 1983, 1 female, USNM 96529; North Pacific, 52°04'22"N, 176°58'01"E, 384 m, 20 Oct 2000, 1 male, USNM 1122539; Ocean Olympic, 52°09'06"N, 176°09'48"E, 384 m, 2000, 1 male, USNM 1122536; Pacific Knight 941-49, 52°46'N, 171°45'W, 0-340 m, 14 Jun 1994, 1 female, USNM 96528; Pacific Knight 941-75, 52°31'N, 173°30'W, 0-213 m, 20 Jun 1994, 1 male, USNM 96527; Pacific Knight 941-204, 53°06'N, 171°42'E, 0-455 m, 31 Jul 1994, 1 female, 1 male, USNM 96249 and 1123534; Patricia Lee, 52°21'09"N, 179°32'52"E, 375 m, 2000, 1 female, USNM 1122500; Sea Storm 90, 51°36'30"N, 177°10'48"W, 202-217 m, 4 Jul 2002, 1 male in alcohol, USNM 1123012; Sea Storm 91, 51°40'51"N, 177°10'49"W, 82 m, 4 Jul 2002, 1 male, USNM 1123284; Sea Storm 92, 51°33'34"N, 177°36'59"W, 367 m, 4 Jul 2002, 1 female in alcohol, 1 male, USNM 1123028-29; Sea Storm 107, 52°10'28"N, 175°14'14"E, 214 m, 8 Jul 2002, 1 female, 2 male in alcohol, USNM 1123001, -05, -31; Sea Storm 108, 52°11'32"N, 175°17'E, 208 m, 8 Jul 2002, 1 female, 1 male, both in alcohol, USNM 1122998 and 1123041; Sea Storm 109, 52°17'16"N, 175°20'56"E, 238 m, 8 Jul 2002, 2 indet. in alcohol, USNM 1122997 and 1123536; Sea Storm 111, 52°16'20"N, 175°59'13"E, 137 m, 9 Jul 2002, 1 male, 1123034; Sea Storm 112, 52°15'12"N, 176°00'42"E, 137 m, 9 Jul 2002, 1 indet., USNM 1122999; Sea Storm 134, 52°14'38"N, 176°01'25"E, 138 m, 15 Jul 2002, 1 indet. in alcohol, USNM 1123033; Sea Storm 138, 52°13'50"N, 175°14'31"E, 146 m, 16 Jul 2002, 1 female, 1 male, USNM 1122769 and 1122965; Sea Storm 148, 52°28'16"N, 173°19'10"W, 194 m, 21 Jul 2002, 2 male, USNM 1122748 and 1122752; Sea Storm 150, 52°30'47"N, 173°29'W, 213 m, 21 Jul 2002, 1 female, USNM 1122758; Sea Storm 151, 52°33'40"N, 173°19'10"W, 203 m, 21 Jul 2002, 1 female, 1 male, USNM 1122755-6; Sea Storm 155, 52°38'43"N, 172°16'23"W, 393-401 m, 22 Jul 2002, 2 indet., USNM 1122761, -64; Shishaldin, 54°07'N, 179°45'E, 366 m, 20 Feb 2000, 1 female, USNM 1122529; Shishaldin, 54°24'03"N, 179°36'11"E, 313 m, 25 Feb 2000, 1 female, USNM 1122532; Shishaldin, 53°56'54"N, 179°49'31"E, 318 m, 14 mar 2000, 1 indet., USNM 1122568; Chew, coll., 51°32'N, 179°15'E, 218-289 m, 1 Sep 1968, 3 female, USNM 76573 and 96532; Haaga, coll. 53, 51°24;12:N, 177°37'20"W, 12 Jun 2000, 1 female, USNM 1122515; Haaga, coll., 54, 51°45;48"N, 178°09'47"W, 441-791 m, 10 Jun 2000, 1 female, 1 male, USNM 1122496 and AB02-30; McCloskey, coll., 52°03'26"N, 179°12'20"E, 475 m, 27 Apr 2000, 2 indet., USNM 1122449 and 1122461; McCloskey, coll., 52°17'45"N, 179°28'57"E, 28 May 2000, 1 female, USNM 1122964; McCloskey, coll., 51°16'46"N, 178°56'01"E, 373 m 21 May 2000, 2 indet., USNM 1122463; Miller, coll., 51°31'09"N, 176°53'14"W, 375 m, 2000, 1 female, USNM 1122498; Myer, coll., 52°15'N, 173°30'W, 208-274 m, 20 Nov 1973, 4 female, USNM 76567; Slear, coll., 52°07'54"N, 177°14'46"E, 238 m, 8 Dec 2000, 1 female, USNM 1122453; Slear, coll., 51°54'04"N, 179°17'28"E, 567 m, 9 Dec 2000, 2 female, USNM 1122450, -51; Stone, coll., 51°51.133'N, 179°50.9'E, 630 m, 8 Aug 2005, 1 female, AB05-0055; Tierny, coll., Cross Sound, Alexander Archipelago, 334 m, 30 Jul 1953, 1 indet., USNM 76569;

Description.

Corallum essentially uniplanar or multiplanar, with occasional short side branches oriented perpendicular to flabellum. Branches anastomose only in larger colonies, the largest colony being the holotype of Stylaster polyorchis , which is 28 cm tall and 35 cm broad, having a massive, dense basal branch diameter of 3.9 cm (Fig. 17B). Distal branches circular to slightly flattened in cross section; larger-diameter branches often rectangular in cross section, the longer axis of the rectangle oriented perpendicular to plane of flabellum. Commensal spionid worm tubes not present. Coenosteum reticular-granular in texture, the coenosteal strips 50-55 µm in width, the slits being a discontinuous series of elongate pores about 15 µm in width; strips covered by small granules. Although generally reticulate in texture, coenosteal strips parallel and straight on exsert abaxial side of each cyclosystem (Fig. 18A). In some specimens the coenosteum is porcellaneous (polyorchis form, Fig. 18I), whereas in others (typical form, Fig. 18 G–H) it appears to be more porous. Coenosteum white, pale orange, and pale pink.

Cyclosystems linearly arranged on both edges of branches, as well as on anterior face, and occasionally on posterior face. Those on branch edges often closely spaced, sometimes directly adjacent to one another (Fig. 18C) or even coalescent. Cyclosystems circular, elliptical, or irregular in shape, 1.0-1.3 mm in diameter, slightly flared, and standing slightly exsert from branch; gastropores circular, 0.40-0.45 mm in diameter. Gastropore tubes cylindrical and usually slightly curved (Fig. 18M) on distal branches, such that gastrostyle tip is difficult to see; ring palisade often absent or only poorly developed. Gastrostyles lanceolate, about 0.5 mm in height, and occupying only lower quarter of gastropore tube; H:D about 2.3-2.5. Gastrostyle bears spines up to 0.4 mm in length.

Dactylotomes 0.10-0.11 mm in width; dactylostyles inconspicuous. Range of dactylopores per cyclosystem is 7-17 (n = 50, average = 11.94 (σ = 2.4), mode = 10). Supernumerary dactylopores absent. Pseudosepta slender and fairly uniform in width, 0. 11-0.13 mm in width, an adcauline diastema of about twice pseudoseptal width sometimes present.

Female ampullae (Fig. 18L) large, smooth, superficial hemispheres 1.0-1.1 mm in diameter, often having a lateral efferent pore about 0.25 mm in diameter. Male ampullae (Fig. 18 A–B) superficial swellings 0.4-0.5 mm in diameter, often clustered on anterior face.

Remarks.

Fisher (1938: 505) discussed the similarities of Stylaster campylecus and Stylaster polyorchis , but concluded they were different species based on six minor differences: Stylaster polyorchis had smaller cyclosystems, straighter gastropore tubes, no ridges on the inside of the gastropore tube, often linked cyclosystems, a ring palisade, and wrinkled female ampullae. Based on more detailed SEM observations on many more specimens, these differences are considered to be intraspecific variation, the holotype of Stylaster campylecus itself having some linked cyclosystems and wrinkled female ampullae. The ring palisade is absent in most coralla, but in some specimens is weakly developed. Stylaster campylecus tylotus was also compared to but differentiated from typical Stylaster campylecus by Fisher (1938) in having a more delicate corallum, a larger ring palisade, a stouter gastrostyle, larger male ampullae, and more dactylopores per cyclosystem, but closer examinations also shows all these character to be within the range of variation of typical Stylaster campylecus (see Table 2). Likewise, no significant differences could be found between Stylaster moseleyanus and Stylaster campylecus , and thus these four taxa are considered to be the same. Interestingly, types of three of the four taxa were collected at Albatross station 3480, prompting Fisher (1938: 514) to remark about "the extraordinary number of species and subspecies dredged at station 3480," considering the possibility that hybridism might be taking place. Stylaster campylecus and Stylaster brochi are the two most commonly collected stylasterids in the Aleutian Islands. Stylaster campylecus can be distinguished from other Alaskan species in Stylaster (Group B) by its slightly flared cyclosystems and the linearity of the coenosteal strips near the cyclosystems (Table 2). The corallum was found to be 100% aragonitic according to Cairns and Macintrye (1992). Of 94 colonies examined, 44 are female, 39 male, and 11 indeterminate, resulting in a fairly equal sex ratio.

Distribution.

Known from throughout the Aleutian Islands from Agattu Island to Unalaska, including Petrel and Bowers Banks, two disjunct records in Alexander Archipelago; 82-1011 m, but most records from 150-500 m.