Revision of the snailfish genus Allocareproctus Pitruk & Fedorov (Teleostei: Liparidae), with descriptions of four new species from the Aleutian Islands. Author James Wilder Orr Author Morgan Scott Busby text Zootaxa 2006 1173 1 37 http://www.zoobank.org/urn:lsid:zoobank.org:pub:BCE33E6B-BA3F-4E35-8EE4-9F34B9358943 journal article z01173p001 Allocareproctus jordani (Burke 1930) Cherry Snailfish (Figures 1-2, 4-7, 12-13; Tables 1-4) Careproctus gilberti Jordan & Thompson 1914 :282, pl. 34, figs. 1, 1a (original description from Sagami Bay, junior primary homonym of C. gilberti Burke 1912 ). Careproctus jordani Burke 1930 :192 (replacement name, redescription); Okada & Matsubara 1938:347 (checklist and keys); Matsubara 1955:1194 (checklist and keys); Kido 1984:339, pl. 365 (brief description); Kido 1988:197, pl. 36 (review, phylogenetics of Liparidae); Pitruk 1990:37 (review, phylogenetics of Liparidae); Nakabo 2000:669, 2002:669 (checklist and keys). Allocareproctus jordani Pitruk & Fedorov 1993:17, figs. 1-5 ( new genus , redescription); Sheiko & Fedorov 2000:31 (possible synonym of “Allocareproctus” pycnosoma ); Orr & Busby 2001:56 (in part; compared to Prognatholiparis ptychomandibularis ); Chernova et al. 2004:3 (checklist). Material examined FMNH 57514 , 130.0 mm, holotype , Misaki, Sagami Bay , Japan ; UW 112293 , 1 (142 mm), 52.3687ºN , 171.2406ºW , 323 m depth, 31 May 2000 , M/V Dominator, cruise 2000- 01, haul 51 , J.W. Orr ; UW 112286 , 1(139.3 mm), 52.0532ºN , 171.8035ºW , 331 m depth, 6 August 2002 , F/V Sea Storm, cruise 2002-01, haul 209 ; UW 112284 , 1(150 mm), Kuril Islands , 16 September 2000 , haul 93, A.M. Orlov ; UW 112287 , 1(173 mm), Kuril Islands , 22 Sep 2000, haul 114, 47.9667ºN , 154.6167ºE , 390 m depth, 22 September 2000 , A.M. Orlov ; UW 112282 , 11(74-86 mm), 52.331ºN , 172.7470ºW , 441 m depth, 23 July 2002 , F/ V Sea Storm, cruise 2002-01, haul 156 , J.W. Orr ; UW 112285 , 1(105 mm), 52.3217ºN , 172.7447 ºW , 458 m depth, 16 June 1994 , F/V Pacific Knight, cruise 1994-01, haul 54 ; UW 112283 , 14(53.3-85.2 mm; 74.2 mm cleared and stained), 52.3405ºN , 172.7455ºW , 441 m depth, 31 May 2002 , F/V Morning Star, cruise 2002-01, haul 32 , J.W. Orr ; UW 112280 , 2(130-137.7 mm), 52.3732ºN , 171.3548ºW , 324 m depth, 10 August 2002 , F/V Sea Storm, cruise 2002-01, haul 225 , R.C. Harrison ; UW 112291 , 2(63.2-132 mm), 52.3687ºN , 171.2406ºW , 323 m depth, 31 May 2002 , M/V Dominator, cruise 2000-01, haul 51 , benthic bag , J.W. Orr ; UW 112288 , 2(126.5-136.5 mm), 56.1354ºN , 169.4400ºW , 631 m depth, 15 July 2002 , F/V Morning Star, cruise 2002-02, haul 119 ; ZIN 49891 , 1(149.5 mm), Kuril Is., Shimushir I. , 47.5ºN , 153ºE , 350 m depth, A. A. Balanov ; UW 112290 , 1(47 mm), 52.2103ºN , 172.2066ºW , 359 m depth, 4 June 2000 , M/V Dominator, cruise 2000-01, haul 74 , benthic bag , J.W. Orr ; UW 112281 , 2(69.2-79 mm), 52.2099ºN , 172.2057ºW , 348 m depth, 23 July 2002 , F/V Sea Storm, cruise 2002-01, haul 157 , J.W. Orr ; UW 112279 , 7(54-104.3 mm), 52.3282ºN , 172.7468ºW , 444 m depth, 4 June 2000 , F/ V Vesteraalen, cruise 2000-01, haul 67 , W.C. Flerx ; UW 112289 , 2(54-104.3 mm), 52.3184ºN , 172.7453ºW , 455 m depth, 22 June 1997 , F/V Vesteraalen, cruise 1997-01, haul 58 , W.C. Flerx ; UW 112292 , 1(145 mm), 52.2754ºN , 170.5991ºW , 235 m depth, 14 June 2004 , F/V Sea Storm, cruise 2004-01, haul 34 , J.W. Orr ; UW 113693 , 1(150 mm), 52.2102ºN , 172.2060ºW , 341 m depth, 23 June 2004 , F/V Sea Storm, cruise 2004-01, haul 73 , J.W. Orr ; UW 113697 , 20(51-117 mm), 52.3255ºN , 172.7466ºW , 450 m depth, 19 June 2004 , F/V Sea Storm, cruise 2004-01, haul 60 , benthic bag , J.W. Orr ; UW 113685 , 1(128 mm), 51.4514ºN , 178.6024ºE , 382 m depth, 21 July 2004 , F/V Gladiator, cruise 2004-01, haul 174 , R.N. Clark ; UW 113686 , 2(118-125 mm), 56.1430ºN , 169.4567ºW , 625 m depth, 20 June 2004 , F/V Northwest Explorer, cruise 2004-01, haul 57 , D.E. Stevenson ; UW 25139 , 1(145 mm), southern Bering Sea , May 1981 ; UW 113687 , 1(125 mm), 56.1913ºN , 169.4864ºW , 478 m depth, 29 July 2004 , F/V Northwest Explorer, cruise 2004-01, haul 158 , J.W. Orr . Diagnosis Teeth simple or with weak shoulders (Fig. 2A); nasal pore 1 with pigmented papilla; peritoneum black; orobuccal valve with single finger-like projection (Fig. 12A); orobuccal cavity pale; gill rakers 8-12, short, blunt; interorbital papilla present or absent; pyloric caeca on right side; body red; iris silver gray. Description Body depth at pectoral-fin base 13.0-19.4 (15.0)%, at center of pelvic disk 15.4-21.9 (15.4)%, at anal-fin origin 17.6-23.7 (18.9)%. Predorsal length 24.4-30.1 (27.9)%. Preanal length 37.4-44.5 (43.9)%. Head large, width 11.8-19.2 (14.6)%, length 22.9-27.5 (25.8)%. Interorbital width 4.5-8.3 (5.2)%. Snout 6.0-9.1 (8.1)%. Mouth small, maxilla length 8.3-13.0 (11.3)%, extending to anterior portion of orbit. Teeth simple, often with weak shoulders (Fig. 2A), in a band of 7-11 oblique rows of 5-9 teeth per row. Orbit large, diameter 5.6-7.3 (6.6)%. TABLE 3. Proportional morphometric characters in percent standard length and meristic characters for specimens of Allocareproctus jordani and selected type material of new species used for statistical analyses. Range, with mean +/- standard deviation below in parentheses. Differences were considered significant at P <0.01. "X" = significant difference between one or more species pairs. "NS" = nonsignificant. "-" indicates the character was not analyzed because it failed to meet statistical assumptions.
A. jordani A. tanix A. unangas A. kallaion A. ungak Significance
n 34 4 30 30 30
Standard length (mm) 53.3-173.0 45.8-77.6 40.3-129.7 67.8-195 55-129.3
Head length 22.2-27.5 (24.6±1.1) 26.3-29.7 (27.8±1.4) 25.1-28.2 (26.5±0.8) 21.7-24.3 (23.2±0.7) 23.9-27.3 (25.8±0.8) X
Head width 11.8-19.2 (14.9±1.7) 12.4-16.8 (14.5±1.8) 12.7-18.8 (14.9±1.4) 12.4-19.6 (14.0±1.6) 11.8-20.4 (14.6±1.6) -
Depth at anal-fin origin 17.6-23.7 (20.2±1.4) 23.3-25.3 (24.2±0.9) 18.8-26.5 (22.7±1.8) 14.0-22.0 (18.4±1.7) 18.2-25.2 (21.6±1.7) X
Depth from dorsal-fin to anal-fin origin 21.6-29.3 (25.9±1.8) 27.3-30.8 (29.2±1.5) 22.4-31.3 (27.2±1.8) 22.1-31.0 (26.4±2.2) 24.1-30.0 (26.8±1.6) X
Depth at pectoral-fin base 13.0-19.4 (16.4±1.7) 17.8-23.0 (20.6±2.1) 14.2-22.6 (18.5±1.8) 11.8-18.6 (15.0±1.5) 15.0-21.0 (17.7±1.6) X
Depth at pelvic disk 15.4-23.3 (18.8±1.8) 19.5-24.5 (21.6±2.1) 18.9-24.6 (21.2±1.5) 13.8-21.2 (17.5±1.7) 16.6-22.9 (20.0±1.7) X
Snout length 6.0-9.1 (7.2±0.8) 7.7-9.2 (8.4±0.7) 6.5-9.2 (7.9±0.6) 5.5-8.8 (7.3±0.7) 6.8-9.3 (7.8±0.6) X
Orbit length 5.6-7.3 (6.5±0.5) 6.4-7.1 (6.8±0.3) 5.0-7.8 (6.6±0.6) 5.3-7.3 (6.3±0.5) 5.2-7.0 (6.3±0.4) X
Interorbital width 4.5-8.3 (6.4±1.1) 5.5-6.4 (5.9±0.4) 5.1-8.2 (6.4±0.6) 4.3-7.7 (5.9±1.0) 4.9-8.0 (6.5±0.6) NS
Suborbital depth to oral cleft 2.1-3.8 (3±0.5) 3.5-4.7 (4.1±0.6) 1.7-4.8 (3.3±0.7) 1.9-4.4 (2.9±0.5) 2.2-4.5 (3.1±0.5) X
Suborbital depth to mandible 5.1-7.9 (6.5±0.7) 7.7-8.3 (8.0±0.3) 5.9-7.9 (6.9±0.6) 4.6-7.2 (5.9±0.6) 5.0-8.4 (6.5±0.8) X
Mouth width 11.2-15.8 (13.5±1.2) 11.8-14.2 (13.4±1.1) 11.1-15.8 (13.1±1.1) 4.6-15.4 (12.8±1.8) 10.6-15.8 (12.9±1.3) -
Maxilla length 8.3-13.0 (10.0±1.0) 10.8-12 (11.5±0.5) 8.4-12.0 (10.8±0.8) 7.5-11.3 (9.9±0.8) 9.0-12.1 (10.2±0.6) -
Mandible length 7.4-14.2 (10.8±1.3) 10.7-12.0 (11.5±0.6) 8.2-13.0 (11.4±0.9) 9.9-12.1 (11.0±0.5) 10.0-12.3 (11.3±0.6) -
Gill-slit length 3.7-8.7 (5.3±1.0) 4.9-7.1 (6.3±1.0) 4.7-7.1 (6.0±0.6) 4.6-6.7 (5.6±0.6) 4.9-7.8 (6.3±0.7) -
to be continued TABLE 3 (continued).
A. jordani A. tanix A. unangas A. kallaion A. ungak Significance
n 34 4 30 30 30
Upper pectoral-fin length 15.7-19.7 (17.9±0.9) 18.0-20.7 (19.2±1.2) 15.4-20.1 (18.3±1.0) 15.8-20.1 (17.7±1.1) 16.0-20.0 (17.8±0.9) NS
Lower pectoral-fin length 12.0-19.0 (15.7±1.6) 16.6-19.9 (17.8±1.4) 13.4-19.5 (16.6±1.5) 13.3-20.2 (15.7±1.4) 14.4-19.8 (16.6±1.4) -
Pectoral-fin notch-ray length 5.3-11.2 (7.6±1.4) 9.0-10.0 (9.5±0.4) 4.9-14.2 (8.4±1.7) 4.6-9.5 (7.4±1.2) 5.8-10.5 (8.0±1.3) -
Predorsal length 24.4-30.1 (27.0±1.4) 29.0-34.5 (32.0±2.3) 26.5-33.3 (29.6±1.3) 22.7-28.6 (25.8±1.3) 25.9-29.7 (27.8±1.1) X
Preanal length 37.4-44.5 (41.0±1.7) 40.6-48.0 (45.0±3.3) 37.4-46.4 (42.1±2.3) 38.8-47.0 (42.9±1.8) 38.4-47.3 (42.2±2) NS
Snout to pelvic disk length 12.4-16.9 (15.3±1.0) 16.8-20.1 (18.5±1.4) 14.3-20 (16.9±1.4) 12.5-16.2 (14.1±0.9) 14.8-24.1 (17.2±2) -
Snout to anus length 25.5-32.9 (29.2±1.5) 30.3-36.9 (33.8±2.8) 28.3-33.8 (31.2±1.5) 24.9-31.6 (28.1±1.5) 25.6-34.7 (29.8±1.8) X
Mandible to pelvic disk length 10.7-14.4 (12.9±0.9) 13.4-16.4 (14.6±1.3) 12.7-17.1 (14.6±1.1) 10.9-13.9 (12.2±0.7) 12.6-21.4 (15.1±1.7) -
Mandible to anus length 24.8-31.2 (27.6±1.6) 28.5-33.6 (30.8±2.1) 15.1-33.7 (29±3.7) 17.6-30.2 (26.5±2.3) 25.3-33.1 (28.3±1.8) -
Pelvic disk length 8.5-11.1 (9.7±0.6) 9.9-11.6 (10.7±0.7) 7.81-1.4 (9.8±0.7) 8.0-9.7 (8.7±0.4) 8.6-11.8 (9.9±0.8) X
Pelvic disk width 7.9-10.7 (9.0±0.6) 9.0-11.1 (9.7±0.9) 7.4-10.7 (8.9±0.8) 7.1-8.8 (8.0±0.4) 7.1-11.1 (9.2±0.9) X
Pelvic disk to anus length 2.6-7.0 (5.0±1.0) 4.7-7.6 (5.9±1.3) 1.7-8.3 (5.3±1.2) 2.7-9.4 (6.1±1.5) 1.5-5.5 (3.4±1.0) X
Anus to anal-fin length 9.0-16.9 (12.1±1.8) 10.6-15.2 (13.2±2.0) 5.7-15.8 (11.3±2.1) 11.6-18.9 (15.7±1.8) 6.7-18.8 (13.0±2.2) -
Caudal-fin length 11.7-15.6 (14.3±0.9) 14.6-17.4 (15.8±1.2) 11.7-17.0 (14.3±1.2) 12.8-17.1 (15.2±0.9) 13.0-17.1 (14.8±0.9) -
Length of dorsal-fin connection to caudal fin 1.8-5.3 (3.4±0.8) 3.3-6.6 (5.0±1.4) 2.5-5.0 (3.8±0.6) 2.0-4.4 (2.9±0.6) 2.3-5.8 (4.0±0.9) X
Length of anal-fin connection to caudal fin 2.6-5.3 (3.9±0.7) 4.1-6.9 (5.7±1.4) 3.0-5.3 (4.3±0.7) 2.6-5.1 (3.9±0.6) 3.0-5.9 (4.8±0.7) X
Caudal-fin base depth 3.1-4.8 (3.6±0.4) 3.7-5.2 (4.1±0.7) 2.4-5.2 (3.7±0.6) 2.7-4.6 (3.6±0.4) 3.3-5.2 (4.3±0.5) -
to be continued TABLE 3 (continued).
A. jordani A. tanix A. unangas A. kallaion A. ungak Significance
n 34 4 30 30 30
Meristics -
Dorsal-fin rays 41-45 (42.9±1.0) 41-42 (41.3±0.5) 42-45 (43.1±0.9) 42-45 (43.1±0.9) 39-42 (40.8±0.8) X
Anal-fin rays 34-38 (36.1±0.8) 34-36 (35.3±1.0) 36-39 (37.1±0.8) 34-37 (35.5±0.7) 33-35 (34.4±0.6) X
Pectoral-fin rays 34-39 (36.1±1.2) 38-39 (38.3±0.5) 38-42 (39.4±1.3) 36-40 (37.9±1.1) 35-40 (37.4±1.5) X
Principal caudal-fin rays 11-13 (12.0±0.6) 12-13 (12.3±0.5) 11-12 (11.9±0.3) 11-14 (12.1±0.6) 12-13 (12.1±0.3) -
Abdominal vertebrae 10-12 (11.1±0.3) 9-10 (9.8±0.5) 10-11 (10.2±0.4) 12-12 (12.0±0.0) 10-11 (10.4±0.5) -
Caudal vertebrae 35-38 (36.6±0.9) 35-37 (36.3±1.0) 36-40 (37.6±1.0) 33-37 (35.6±0.9) 34-36 (35.1±0.6) X
Gill rakers on first arch 8-12 (10.2±0.9) 9-10 (9.75±0.5) 8-12 (9.3±1.0) 14-18 (16.3±1.1) 7-11 (9.0±1.1) X
Papillae present on many pores: present on nasal pore 1, absent from nasal pore 2; present on maxillary pores 4-6, often present on pores 2-3 (4-6); present on preoperculomandibular pores 5-6, often present also on pores 4 and 7 (6-7); present on suprabranchial pores 1-2 (Fig. 4B). Interorbital papilla present or absent (present in holotype). Cephalic free neuromasts profuse and relatively large, scattered evenly over interorbit and nape. Gill opening small, 3.7-8.7 (5.3)%, entirely above pectoral fin or extending to pectoral-fin ray 2. Gill rakers on anterior arch 8-12 (ca. 10, gill slit of holotype not dissected), blunt and stout. Central projection of orobuccal valve a moderately elongate single lobe (Fig. 12A). Dorsal-fin rays 41-45 (44), tips of anterior 4-8 rays projecting from fin membrane, anteriormost rays about 50% free from membrane, succeeding rays less so; posteriormost ray attached membranously to dorsalmost caudal-fin ray for 1.8-5.3 (2.7)%. Anal fin with 33-38 (38) rays, posteriormost ray membranously attached to ventralmost caudal-fin ray for 2.6-5.3 (3.2)%. Two anal-fin pterygiophores and associated rays anterior to first haemal spine. Pectoral-fin rays 33-39 (38) in two lobes separated by a shallow notch, 8-10 (9) rays in lower lobe. Pectoral-fin rays in notch slightly more widely spaced than rays of lobes. Dorsalmost ray at level of ventral rim of orbit. Upper lobe rounded, extending to anal-fin origin, length 15.7-19.7 (16.7)%, with ray 6 longest; length of shortest notch ray 5.3-9.5 ( 6.3)%; length of lower lobe 12.6-17.7 (15.2)%, extending between posterior margin of pelvic disk and anus, with ray 2 longest, rays 3-10 shortening ventrally. Pelvic disk large, length 8.5-11.1 (9.8)%, width 7.9-10.7 (8.9)%. Distance from disk to anus 2.6-7.0 (5.8)%, about 25-70 (59.0)% disk length (DL), distance from anus to analfin origin 9.0-16.9 (11.7)%, about 80-175 (119.7)% DL. Urogenital papilla conical, short, when protruding about 11.5 (not protruding in holotype)% pelvic disk length, unpigmented. Pyloric caeca about 23 (holotype not dissected), on right side of body, finger-like, 25-50% HL. Caudal fin slightly rounded, length 11.7-15.6 (13.9)%, depth at hypural plate 3.1- 4.8 (4.2)%, with principal rays 11-13 (12), dorsal principal rays 5-6 (6), ventral principal rays 6-7 (6). Dorsal procurrent rays 1-3 (2), borne on epural and posteriormost neural spine; ventral procurrent ray one, borne on expanded posteriormost haemal spine. Vertebrae 46-49 (49), abdominal vertebrae 10-12 (11), caudal vertebrae 35-38 (38). Body in life pink to red, with dark speckling at origin of dorsal fin; iris silver gray. Color in alcohol pale, with scattered speckling on nasal and other cephalic papillae and at dorsal-fin origin. Peritoneum black; orobranchial cavity pale. Largest specimen examined a 173 mm female (UW 112287). Smallest female with yolked eggs 105 mm; smallest male with enlarged, swollen testes 74.5 mm.
Range Allocareproctus jordani ranges from the type locality in Sagami Bay, Japan, through the Kuril Islands, eastern Sea of Okhotsk, off the southern tip of Kamchatka, the Aleutian Islands from Attu to Amukta I., and Pribilof Canyon, at depths of 75-631 m (Fig. 7). FIGURE 7. Distribution of Allocareproctus jordani based on material examined, 76 specimens in 23 lots. Each symbol may represent more than one capture. Etymology The specific epithet is a patronym honoring David Starr Jordan (Burke 1930). Comparisons Allocareproctus jordani is most similar to two new species described below: A. unangas and A. ungak . Both species differ from A. jordani in possessing strongly trilobed teeth. In addition, although counts of median-fin rays, pectoral-fin rays, vertebrae, and gill rakers overlap in total ranges, several characters differ significantly (Tables 1-3). Modes of all these characters differentiate A. jordani from A. ungak , A. jordani having significantly higher counts of median-fin rays, caudal vertebrae, and gill rakers, and lower counts of pectoral-fin rays. In contrast, anal-fin ray, pectoral-fin ray, and caudal vertebrae counts are significantly higher in A. unangas , while gill-raker counts are lower than in A. jordani . The two other new species of Allocareproctus are readily distinguished from A. jordani by several characters. Allocareproctus kallaion differs externally in coloration, having a dark gray head, blotchy red body, and dark orobuccal cavity, unlike the uniform red and pale coloration of A. jordani . It also has a comb-like orobuccal valve (Fig. 12B), a higher count of larger gill rakers (Table 2), and a uniserial series of teeth in a comb-like row on the dentary. Counts of abdominal vertebrae are invariably 12 in A. kallaion versus typically 10-11 in A. jordani (3 of 55 specimens examined had 12 abdominal vertebrae; Table 2). The pale peritoneum, fewer cephalic pores with papillae, and strongly trilobed teeth distinguish A. tanix from A. jordani . Several morphometric characters also differ significantly between A. jordani and A. tanix , including a greater body depth, head length, suborbital depth to both oral cleft and mandible, predorsal length, snout-to-pelvic disk length, and caudal length in A. tanix . Remarks Pitruk & Fedorov (1993) redescribed A. jordani on the basis of material collected from the Kuril Islands. We examined new material from the region (UW 112287 and UW 112284) and confirmed the identity of one of their specimens as A. jordani (ZIN 49891). Other components of their redescription of the species, including meristics, tooth shape, distribution of papillae, coloration, and overall body shape, agree with our characterization of A. jordani and exclude from their account the new species described below.