Careproctus rastrinus Gilbert & Burke 1912

Orr, James Wilder, Kai, Yoshiaki & Nakabo, Tetsuji, 2015, Snailfishes of the Careproctus rastrinus complex (Liparidae): redescriptions of seven species in the North Pacific Ocean region, with the description of a new species from the Beaufort Sea, Zootaxa 4018 (3), pp. 301-348 : 307-316

Careproctus rastrinus Gilbert & Burke 1912

English common name: Salmon Snailfish Japanese common name: Sake-bikunin

Figures 1 A, 5A, 6; Tables 1–3

Careproctus rastrinus Gilbert & Burke 1912:362 , pl. 43, fig. 2. Type locality: Sea of Okhotsk, 48°36'10"N, 145°17'30"E.— Burke 1930:136, figs. 57–58 (description, key).— Soldatov & Lindberg 1930:24 (key).— Chapman & DeLacy 1934:3 (comparisons).— Taranetz 1937:137 (Sea of Okhotsk, key).— Okada & Matsubara 1938:346, pl. 85-2 [Sea of Okhotsk, Sakhalin; plate modified from Gilbert & Burke (1912)].— Schmidt 1950:203 (brief description, Russia).— Böhlke 1953:136 (type catalog).— Matsubara 1955:1193 (southern Sea of Okhotsk).— Kato 1956: 329 (Sea of Japan, list, = C. trachysoma ?).— Ueno 1971:97 (off Sakhalin).— Quast & Hall 1972:29 (Alaska).— Fedorov 1973:66 (Bering Sea).—Kido 1984:339, pl. 305-D (brief description, Sea of Okhotsk).— Lindberg & Krasyukova 1987:446 (Sea of Okhotsk and Sea of Japan, in part?).— Allen & Smith 1988:67 (Alaska, “pink snailfish”, in part).— Kido 1988:217, fig. 48 (description, phylogenetics, in part).— Pitruk 1990:38 (Sea of Okhotsk).— Tsuda 1990:513 (Sea of Japan, Sea of Okhotsk, in part).— Robins et al. 1991:44 (list, based on Allen & Smith 1988).—Nakabo 1993:586 (Sea of Japan, Sea of Okhotsk, and Pacific coast of northern Japan, in key, in part).— Amaoka et al. 1995:211 (Sea of Japan, Sea of Okhotsk, and Pacific coast of northern Japan, in part).— Shinohara et al. 1996:178 (Pacific Japan, = C. pellucidus ?).— Kido 1997:245 (Sea of Japan, Sea of Okhotsk, and Pacific coast of northern Japan, in part).—Nakabo 2000:672 (Sea of Japan, Sea of Okhotsk, and Pacific coast of northern Japan, in key, in part).— Sheiko & Fedorov 2000:32 (northern Kuril Is. and Kamchatka, in part).— Mecklenburg et al. 2002:611 (Alaska, illustration, in key, = C. scottae , in part).—Nakabo 2002:672 (Sea of Japan, Sea of Okhotsk, and Pacific coast of northern Japan, in key, in part).— Youn 2002:278 (list, = C. trachysoma ?).— Fedorov et al. 2003:99 (Sea of Okhotsk).— Maeda & Tsutsui 2003:495 (off Hokkaido, list, in part).— Chernova et al. 2004:14 (checklist).— Nelson et al. 2004:124 (list, = C. scottae ).— Chernova 2005b:S7 (comparisons).—Love et al. 2005:103 (checklist, = C. scottae in part).— Orlov 2005:141 (Kamchatka, Kuril Islands, in part).— Kitagawa et al. 2008:77 (Sea of Japan, Sea of Okhotsk, and Pacific coast of northern Japan, in part, photo = C. pellucidus ?).— Shinohara et al. 2009:720 (Pacific Japan, = C. pellucidus ?).— Kai et al. 2011a:143 (genetics, morphology, phylogenetics, as “OKH1”).— Kai et al. 2011b:368 (fig. 1a, phylogenetics, as “OKH1”).— Honma 2013 (Sea of Japan).— Nakabo & Kai 2013:1213 (northern Japan to Gulf of Alaska, in key, in part).—Page et al. 2013:127 (list, = C. scottae ).— Shinohara et al. 2014:256 (Sea of Japan, = C. trachysoma ?).

Holotype. USNM 73331, 243.3 mm, Russia, southern Sea of Okhotsk, off Cape Patience, Sakhalin Island, 48.6028°N, 145.2917°E, Albatross station 5026, 217 m depth, 28 September 1906.

Paratypes. SU 22371, 147.8 mm, Russia, southern Sea of Okhotsk, off Cape Patience, Sakhalin Island, 48.5417°N, 145.1458°E, Albatross station 5021, 114 m depth, 27 September 1906.

Additional material examined. A total of 35 specimens, not including the types above, 72.0– 385.6 mm SL. See “Non-type material examined below.”

Diagnosis. Careproctus rastrinus is distinguished from all other species of Careproctus by the combination of cyt b and 16S rRNA sequences (“OKH1” of Kai et al. 2011a, b; Table 1), an anteriorly robust body covered by cactus-like prickles, the presence of a postorbital pore, a moderate-sized pelvic disc, a lower pectoral-fin lobe longer than upper lobe, a pale peritoneum, and a speckled gray stomach. It is most similar to light colored C. trachysoma of the western Pacific, from which it can be distinguished by the color of its peritoneum and stomach (pale peritoneum and speckled gray stomach vs. speckled pale and dark in C. trachysoma ), and its deeper and longer head and anterior body. It is also similar to the eastern Pacific C. scottae , from which it can be distinguished by its higher counts of caudal vertebrae (51–56 in C. rastrinus vs. 48–52 in C. scottae ) and anal-fin rays (51–56 vs. 47–53), speckled gray stomach (pale in C. scottae ), smaller pelvic disc (9.9–26.8 vs. 13.9–26.5 % HL), and shorter nasal tube (0.9–4.7 vs. 3.2–6.4 % HL), and to C. phasma from which it can be distinguished by the postorbital pore (present in C. rastrinus vs. absent in C. phasma ), smaller pelvic disc (9.9–26.8 vs. 20.1–35.9 % HL), longer upper jaw (36.6–55.1 vs. 31.4–48.1 % HL), shorter nasal tube (0.9–4.7 vs. 2.9–5.7% HL), speckled gray stomach (pale in C. phasma ), higher counts of dorsal-fin rays (57–63 vs. 50–56), higher counts of anal-fin rays (51–56 vs. 43–49), and higher counts of vertebrae (62–66 vs. 55–60). It is further distinguished from C. spectrum by its higher counts of dorsal-fin rays (57–63 vs. 52–54), smaller pelvic disc (9.9–26.8 vs. 22.5–23.1 % HL), and smaller orbit (16.7– 33.3 vs. 33.9–34.4 % HL).

Description. Body heavy and deep anteriorly, tapering strongly posteriorly, strongly compressed; depth at pectoral-fin base 90.6–145.7 (126.3) % HL. Head large, 20.2–32.7 (26.2) % SL, and robust, dorsal profile rounded from nape to snout. Snout blunt, slightly projecting anterior to lower jaw. Mouth terminal, small, horizontal; upper jaw 36.6–55.1 (43.8) % HL, maxilla extending to anterior part of orbit or mid-orbit, oral cleft extending to anterior rim of orbit; mandible 45.1–60.9 % HL. Premaxillary tooth plates matching mandibular tooth plates. Premaxillary and mandibular teeth simple with weak shoulders in 22–46 oblique rows of 7–14 teeth forming narrow bands. Diastema absent at symphysis of upper and lower jaws. Orbit 16.7–33.2 (21.8) % HL, dorsal margin well below dorsal contour of head, suborbital depth to upper jaw 11.1–19.8 (13.5) % HL, to lower jaw 26.1–36.6 (34.2) % HL; pupil round. Interorbital space broad, fleshy distance 31.8–54.4 (43.1) % HL, bony distance 17.8–32.1 (32.1) % HL, strongly convex. Snout typically much longer than orbit, 93.1–257.3 (176.0) % OL, 30.6–46.1 (38.3) % HL. Nostril single, with well-developed tube at level with lower rim of orbit; nostril tube length 3.7–18.3 % OL.

Pores of cephalic lateralis system of moderate size, pore pattern 2-6-7-2, chin pores paired. Interorbital pore absent.

Gill opening small, 19.7–49.4 (29.6) % HL, upper margin at or just above level of dorsal rim of orbit, extending ventrally to just above the upper pectoral-fin ray to pectoral-fin ray 1–5 (ray 4). Opercular flap rounded to slightly angular (rounded). Gill rakers 9–12 (Tables 2–3), short, blunt.

Dorsal-fin rays 57–63 (60; Tables 2–3), anterior dorsal lobe absent, anterior rays buried in tissue, tips of more posterior rays not exserted. Anteriormost dorsal-fin pterygiophore inserted between neural spines 3 and 4, rayless or bearing a single small or rudimentary ray (between 3 and 4, rayless). Predorsal length 23.3–33.1 (30.4) % SL. Anal-fin rays 51–56 (52; Tables 2–3), one or two anal-fin pterygiophores anterior to first haemal spine (one), each bearing a single ray, tips of all rays slightly exserted. Anal-fin origin below vertebrae 13–14 (caudal vertebrae 2–3), preanal length 33.4–48.7 (36.8) % SL.

Pectoral fin deeply notched, with 32–39 (37) rays (Tables 2–3). Upper lobe 54.5–75.7 (56.5) % HL, with 24– 31 (29) rays extending well beyond anus to or near anal-fin origin, shorter than lower lobe, dorsalmost rays lengthening to rays 8–10, more ventral rays gradually shortening to shortest ray of notch. Lower lobe elongate, 52.6–141.9 (101.0) % HL, with 7–11 rays (8), extending beyond anus to or near anal-fin origin; dorsal rays gradually lengthening to elongate rays 7–8, ventral rays gradually shortening to ventralmost ray near pectoral symphysis. Tips of rays in dorsal lobe 0–25% free of membrane, rays of lower lobe more strongly exserted up to 70% free. Notch strong, rays in notch slightly more widely spaced than rays of lobes, more widely spaced ventrally. Uppermost pectoral-fin ray level with region between ventral rim of orbit and cleft. Insertion of lowermost pectoral-fin ray below mid-orbit. Proximal pectoral radials four (3+1), robust: radials 1–2 notched and hour-glass shaped; radial 3 a rounded square, slightly notched; radial 4 small, round ( Fig. 5 A). Interradial fenestrae three, extending between scapula and proximal radials 1–3: fenestrae between the scapula and radials 1 and 2 elliptical, generally elongate in the sagittal plane, fenestra between radials 2 and 3 oval. Scapula broadly T-shaped with robust distally broadened helve; coracoid with broad triangular head and broad helve, angled slightly anteriorly. Distal radials present at base of rays 2–29, ventralmost at level of proximal radial 4, dorsalmost ray and more ventral rays articulating directly with pectoral cartilage.

Pelvic disc small, length 9.9–26.8 (10.6) % HL, round, about as long as wide, width 11.0–25.8 (12.2) % HL, anterior lobe weakly developed, slightly cupped, distance from snout to pelvic disc 9.6–14.6 (12.3) % SL. Anus at level about even with posterior rim of orbit, close behind pelvic disc; distance from snout to anus 13.7–26.1 (16.3) % SL, 54.0–82.6 % HL.

Principal caudal-fin rays 9–10, dorsal procurrent rays 1–2, ventral procurrent rays 0–2 (1–2 + 4–5/5 + 0–2) (? + 4/5 +?). Caudal fin 30.8–54.0 (37.4) % HL. Membrane of posterior dorsal-fin rays attached to caudal fin at shorter distance than anal-fin rays: dorsal-fin rays attached to caudal fin 35.1–66.5 (49.4) % CL; anal-fin rays, 38.6–71.1 (57.1) % CL. Depth at base of caudal fin 12.0–21.3 (18.5) % CL.

Skin relatively thick, thick gelatinous layer beneath skin, cactus-like prickles uniformly covering body, in most dense region about 13 prickles in orbit length. Pyloric caeca 21–34, length about 27–53 (36.2) % HL, center-left side of visceral cavity.

Vertebrae 62–66 (64), precaudal 10–12 (10), caudal 51–56 (54; Tables 2–3). Pleural ribs 2 or 3, anteriormost small when 3, others long and slender, present on vertebrae 8–9 or 9–10 or 10–11 (9–10).

Coloration. Body and fins orangish pink and white in life ( Fig. 1 A); head, dorsum from nape to caudal fin, and ventrum at anal-fin origin to caudal fin orangish pink; isthmus, base of pectoral fin, and body posterior of gill slit to anal-fin origin lighter; area above belly silvery white (crystalline guanine) with orangish-pink highlights becoming obsolete at about a quarter to half the anal-fin length; base and lower lobe of pectoral fin white; dorsal margin and distal portion of fin orangish pink; dorsal half of eye dark, silvery gray ventrally. Body and fins pale in preservation; base of fins beneath skin with pigment, showing faint line between fins and body. Peritoneum pale, speckled gray; orobranchial cavity pale; stomach gray, intestines pale to darkly mottled, pyloric caeca pale to darkly mottled, and urogenital papilla pale.

TABLE 2. Proportional morphometric and meristic characters of species of the Careproctus rastrinus species complex. Morphometric data are given in percent SL (standard length) and presented as the range, followed by the mean +/- standard deviation (SD).

C. rastrinus C. trachysoma C. scottae C. acanthodes

n Range N Range n Range n Range

(Mean ± SD) (Mean ± SD) (Mean ± SD) (Mean ± SD) Standard length 37 62.3–385.6 35 101.6–264.8 118 44.6–253.2 22 66.9–104.7 Morphometrics

Head length 37 20.2–32.7 35 19.3–26.3 114 21.7–32 22 24.2–27

(25.2 ± 2.3) (22.4 ± 1.7) (26.3 ± 1.8) (25.6 ± 0.7)

Head width 31 7.7–17.1 32 8.8–14.8 53 7.9–16.8 13 11.9–16.7

(12.3 ± 2.3) (11.7 ± 1.6) (12.4 ± 2.1) (14.4 ± 1.6)

Greatest body depth 35 22.7–35.9 34 20.8–31.1 106 20.6–41 22 21.1–28.2

(29.3 ± 3.2) (25.9 ± 2.6) (30.9 ± 4) (25.1 ± 1.8)

Body depth at anal-fin origin 8 23.3–31 5 22.7–24.3 15 18.9–35.5 7 20.5–26.5

(27.2 ± 2.7) (23.5 ± 0.8) (28.7 ± 4.1) (23.1 ± 1.9)

Dorsal-fin origin to anal-fin origin 7 27–33.7 5 25.8–30 13 26.8–37 7 22.6–31.6

(30.4 ± 2.9) (27.9 ± 1.8) (31.7 ± 2.9) (26.9 ± 3.4)

Body depth at pectoral-fin base 36 22.7–35.4 33 18.7–30.7 105 18.4–41 22 21.1–28.2

(28.7 ± 3.3) (25 ± 2.9) (30.8 ± 4.1) (25 ± 1.8)

Body depth at pelvic fin 7 15.3–23.8 5 12.2–18 12 16.3–22.6 7 16.4–20.5

(19.6 ± 2.9) (15.2 ± 2.1) (19.6 ± 2) (19.2 ± 1.4)

Snout length 36 7.2–11.6 35 6.2–9.5 105 7.2–13.4 22 6.9–10

(9.3 ± 1) (8 ± 0.7) (9.9 ± 1.3) (8.9 ± 0.9)

Orbit length 36 3.7–9.2 35 4–6.4 112 4–9.4 22 6.1–8.5

(5.8 ± 1.2) (5.4 ± 0.6) (6.2 ± 1) (6.9 ± 0.7)

Interorbital width (bony) 19 4.8–9 5 4.9–6.4 18 5–8.6 10 3.7–7.5

(6.5 ± 1.1) (5.8 ± 0.6) (6.7 ± 1) (6.3 ± 1.5)

Interorbital width (fleshy) 36 8.4–14 35 7.8–13.1 101 5.5–15.1 22 11–19.3

(11.5 ± 1.4) (11.1 ± 1) (10.7 ± 1.9) (13.9 ± 2.1)

Suborbital depth to upper jaw 19 2.7–5 9 2.5–3.5 15 2.3–4.6 10 1.6–4.8

(4 ± 0.6) (3 ± 0.4) (3.6 ± 0.7) (3.7 ± 1) Suborbital depth to lower jaw 19 6.4–10.3 9 5.8–7.4 15 6.9–9.1 10 5.1–9

(8.4 ± 0.9) (6.4 ± 0.6) (8.1 ± 0.7) (7.4 ± 1.3)

Mouth width 19 8.9–15.4 5 8.7–11.5 15 9.4–14 7 10.8–15.6

(12.2 ± 1.8) (10.2 ± 1.1) (12.1 ± 1.3) (13.5 ± 1.8)

Maxilla length 31 9.2–13.6 32 7.7–11.7 105 9–14.1 13 10.4–12.1

(11.6 ± 1.1) (9.8 ± 0.9) (11.1 ± 1) (11.2 ± 0.5)

Mandible length 19 10.6–16.2 9 9.4–11.4 15 11.8–14.9 10 10.9–13.2

(13.6 ± 1.3) (10.2 ± 0.6) (12.9 ± 0.9) (12.1 ± 0.7)

slit length 30 4.6–12 33 4.7–9.6 103 4.7–11.2 21 4.6–11

(8.3 ± 1.8) (7.4 ± 1.1) (8.3 ± 1.3) (7.4 ± 1.3)

Pectoral fin length 29 12.3–18.5 34 13.3–18.8 103 12.9–21.2 19 12.7–19.2

(15.8 ± 1.6) (16.1 ± 1.3) (16.3 ± 1.7) (16.5 ± 1.6)

Pectoral-fin lower lobe length 29 14.8–34.6 35 18.5–28.4 101 15.7–34.7 17 8.9–15.8

(23.8 ± 4) (24.5 ± 2.4) (23 ± 4.3) (12.3 ± 2)

Notch ray length 18 3.3–8.5 10 5.8–9.2 16 3.8–8.1 7 4–7.5

(5.3 ± 1.3) (7.7 ± 1.3) (6 ± 1.2) (5.8 ± 1.4)

……continued on the next page TABLE 2. (Continued)

C. rastrinus C. trachysoma C. scottae C. acanthodes

n Range N Range n Range n Range

(Mean ± SD) (Mean ± SD) (Mean ± SD) (Mean ± SD) Predorsal length 36 23.3–33.1 35 21.9–30.4 100 22.8–34.8 21 26.3–32.6

(27.8 ± 2.4) (25.7 ± 1.8) (28.9 ± 2.3) (29.3 ± 1.6) Preanal length 36 33.4–48.7 35 31.5–44.4 99 30.5–49.5 21 33.4–45.2

(41.1 ± 3.6) (37.6 ± 3.5) (39.6 ± 3.8) (39.2 ± 3.1) to pelvic disc length 19 9.6–14.6 5 10–12.4 15 9.9–15.8 10 11.7–14.9

(12.4 ± 1.3) (10.9 ± 1.1) (11.8 ± 1.4) (13.2 ± 1) to anus length 29 13.7–26.1 34 13.7–19.9 104 13.1–26.2 22 16.8–22.5

(17.2 ± 2.7) (16.5 ± 1.6) (18.8 ± 2.8) (19.9 ± 1.5) Pelvic disc length 37 2.3–7.3 34 2.5–4.2 115 3.5–6.8 22 4.6–7.3

(4 ± 1.1) (3.3 ± 0.4) (4.8 ± 0.7) (5.7 ± 0.6) Pelvic disc width 31 2.6–7.2 18 2.3–3.9 96 2.8–6.4 22 4.2–7.6

(3.8 ± 1) (3.2 ± 0.4) (4.6 ± 0.6) (5.9 ± 0.9) disc to anus length 19 0.1–3.6 9 1.2–3.8 15 0.6–2.8 10 0.1–1.9

(1.9 ± 0.7) (2.3 ± 0.8) (1.6 ± 0.7) (1 ± 0.6) to anal fin length 19 17.2–30.8 5 20.3–30.6 15 16.3–27.8 10 19.1–27.8

(24.4 ± 2.7) (27.2 ± 4.1) (23.3 ± 3.4) (22.3 ± 2.7) Caudal fin length 30 8.4–14.1 14 8.8–11.3 72 9–13.4 15 9.2–12.1

(10.7 ± 1.3) (10.3 ± 0.7) (10.7 ± 1.1) (10.7 ± 0.9) Dorsal-fin attachment to caudal fin length 30 3.4–7.4 13 4–6.2 71 3–7.6 14 2.1–5.1

(5.4 ± 1) (5.2 ± 0.6) (5.7 ± 0.9) (3.6 ± 0.8) Anal-fin attachment to caudal fin length 29 3.8–7.6 12 3–6.7 71 4.3–8.4 14 3.3–6.3

(5.9 ± 0.9) (5.4 ± 1.1) (6.2 ± 0.8) (4.4 ± 0.9) Caudal base depth 19 1.2–2.2 5 1.1–1.7 15 1.4–2.7 10 0.7–1.9

(1.7 ± 0.3) (1.4 ± 0.2) (1.8 ± 0.3) (1.5 ± 0.4) tube length 15 0.2–1.2 15 0.3–1 10 0.8–1.5 7 0.5–1.6

(0.6 ± 0.3) (0.7 ± 0.2) (1 ± 0.2) (0.9 ± 0.4)

Meristics

Dorsal-fin rays 31 57–63 35 57–62 124 54–61 21 52–55

(59.6 ± 1.3) (59.5 ± 1.3) (56.9 ± 1.2) (53.6 ± 1.1) Anal-fin rays 31 51–56 34 51–57 124 47–53 22 44–49

(52.5 ± 1.3) (53.6 ± 1.4) (50.0 ± 1.2) (47.2 ± 1.1) Pectoral-fin rays 31 32–39 35 31–37 124 31–39 21 29–36

(35.3 ± 2.0) (33.7 ± 1.6) (33.8 ± 1.9) (32.2 ± 1.9)

Pectoral-fin lower lobe rays 31 7–11 34 7–10 124 7–12 19 6–9

(8.2 ± 1.1) (8.7 ± 1) (8.5 ± 0.8) (7.1 ± 0.9) rakers 24 9–12 21 8–11 120 8– 13 6 7–10

(10.8 ± 1.0) (9.2 ± 1) (10.1 ± 0.9) (9.2 ± 1.2) Principal caudal-fin rays 30 9–10 22 8–11 105 8– 10 9 9–11

(9.1 ± 0.4) (9.2 ± 0.8) (9 ± 0.3) (10.1 ± 1.1) Precaudal vertebrae 36 10–12 29 10–12 116 10–13 22 9–11

(10.7 ± 0.6) (10.9 ± 0.5) (10.7 ± 0.5) (9.8 ± 0.7) Caudal vertebrae 36 51–56 29 51–56 113 48–52 22 47–51

(53.1 ± 1.3) (54.2 ± 1.2) (50.4 ± 0.9) (48.5 ± 1.2) vertebrae 31 62–66 29 62–67 123 59–64 22 56–61

(63.9 ± 1.2) (65.1 ± 1.2) (61.1 ± 1.0) (58.4 ± 1.3) TABLE 2. (Continued)

C. pellucidus C. phasma C. spectrum C. lerikimae

n Range n Range n Range n Range

(Mean ± SD) (Mean ± SD) (Mean ± SD) (Mean ± SD) Standard length 34 92.6–285.5 58 54.3–272 3 68.2–91 27 50.5–159 Morphometrics

Head length 34 21.2–28.3 58 22.5–29.7 2 26.6–31.8 27 22.9–29.8

(23.8 ± 1.7) (26.1 ± 1.7) (29.2 ± 31.8) (27.1 ± 1.7)

Head width 26 8.5–15.1 32 7.3–19.3 2 14.3–17.1 25 9–17.5

(12.3 ± 2) (12.8 ± 2.6) (15.7 ± 17.1) (13 ± 2.4) Greatest body depth 34 22.7–35.3 46 20.4–34.1 3 24.7–28.5 24 21.4–31.6

(28.4 ± 2.8) (28.6 ± 3.5) (26.7 ± 28.5) (26.1 ± 2.2) Body depth at anal-fin origin 9 23.5–29.6 5 21.2–28.3 3 23–28.5 25 15.1–26.2

(26.5 ± 2.6) (23.8 ± 2.8) (26.1 ± 28.5) (20.6 ± 3) Dorsal- to anal-fin origin 9 26.5–35 5 25.2–36.4 1 28.0 25 20.4–31.8

(29.1 ± 2.7) (30.9 ± 4.1) (26.7 ± 3.2) Body depth at pectoral-fin base 34 22.7–35.3 46 20.4–34.1 1 24.7 25 20.8–29

(28.4 ± 2.9) (28.6 ± 3.5) (25.3 ± 2) Body depth at pelvic fin 9 16.5–20.1 5 17.7–26.4 1 20.3 25 15.3–21.6

(18 ± 1.3) (21.9 ± 3.9) (18 ± 1.5) Snout length 34 6.7–11.3 47 7.1–11.3 2 7.5–10.2 25 7.4–10.6

(9 ± 1) (9.5 ± 1) (8.8 ± 10.2) (9 ± 1)

Orbit length 34 5–9.6 55 4.4–9 3 9–10.9 26 5.4–8.5

(7.7 ± 1.1) (6.1 ± 1) (10.1 ± 10.9) (6.9 ± 0.8) Interorbital width (bony) 17 4.8–7.9 12 3.7–8.6 1 5.5 26 4.5–8.2

(6.2 ± 0.9) (6.1 ± 1.3) (6.2 ± 1.1) Interorbital width (fleshy) 34 9.5–14.5 44 5.4–13.4 2 8.2–12.2 26 7.6–15.9

(12 ± 1.3) (9.7 ± 2.2) (10.2 ± 12.2) (12 ± 2.2)

Suborbital depth to upper jaw 17 2.6–4.2 11 3.2–4.8 1 2.5 25 3–5.1

(3.2 ± 0.5) (3.9 ± 0.5) (4.1 ± 0.5) Suborbital depth to lower jaw 17 6.6–9.4 11 6.6–9.9 1 9.0 25 6.8–10.4

(7.6 ± 0.8) (8.1 ± 0.8) (8.6 ± 0.9)

Mouth width 17 8.9–14 11 9.6–16.3 1 15.4 25 8–18.2

(11.1 ± 1.3) (12.8 ± 2.2) (12 ± 2.3) Maxilla length 26 9.7–12.8 45 8.1–12.2 2 11.9–14.2 25 10.4–14.1

(11.3 ± 0.7) (10.3 ± 1) (13 ± 14.2) (12.4 ± 1) Mandible length 17 10.3–15.4 12 7.1–15.3 1 14.4 25 11.9–17.8

(12.7 ± 1.4) (12.2 ± 2.1) (14.5 ± 1.4)

slit length 31 5.2–10.7 46 3.4–10.2 1 4.7 25 4.6–10

(8 ± 1.5) (6.9 ± 1.4) (7.5 ± 1.3) Pectoral fin length 33 11.1–17.5 47 13.7–22.9 1 17.4 26 13–21.5

(15.3 ± 1.4) (18.1 ± 2) (17.5 ± 2.1) Pectoral-fin lower lobe length 33 14.2–29 47 17.6–34.2 1 9.1 26 15.8–25

(22.5 ± 3.6) (26.2 ± 3.9) (19.8 ± 2.6) Notch ray length 13 4.7–9.8 11 4.8–10.3 1 4.0 24 3.6–9.9

(6.5 ± 1.6) (7.9 ± 1.6) (6.4 ± 1.5)

……continued on the next page TABLE 2. (Continued)

C. pellucidus C. phasma C. spectrum C. lerikimae

n Range n Range n Range n Range

(Mean ± SD) (Mean ± SD) (Mean ± SD) (Mean ± SD) Predorsal length 34 25–35.1 48 24–32.8 1 24.7 25 25.2–32.2

(28.9 ± 2.1) (28.6 ± 2.2) (28.8 ± 1.8) Preanal length 34 32.9–46.5 47 31.6–51.8 1 39.1 25 36.3–48.2

(41.3 ± 3.1) (41.5 ± 3.7) (42 ± 3.2) to pelvic disc length 18 8.7–14.4 11 9.3–15.7 1 13.7 25 11–16

(12 ± 1.5) (11.3 ± 1.8) (13.1 ± 1.4) to anus length 33 14.1–21.2 45 13.7–27.8 1 20.3 25 14.7–25.2

(16.8 ± 1.6) (18.5 ± 3.3) (19.7 ± 2.4) Pelvic disc length 34 2.2–4.1 58 4.9–9.6 2 6.2–7.1 27 5–7.4

(3.3 ± 0.5) (6.8 ± 1) (6.6 ± 7.1) (6.2 ± 0.6) Pelvic disc width 34 2.2–3.9 42 4.8–7.8 2 6–6.5 25 4.2–6.8

(3.1 ± 0.4) (6.3 ± 0.8) (6.3 ± 6.5) (5.5 ± 0.7) disc to anus length 18 0.7–3.2 11 0.4–1.9 1 1.1 26 0.1–2.6

(1.6 ± 0.7) (1 ± 0.4) (1.3 ± 0.7) to anal fin length 18 17.8–33.1 12 20.8–30.6 1 22.9 25 20.9–32.1

(26.9 ± 4.4) (26.8 ± 3.5) (25.4 ± 3) Caudal fin length 32 10–13.1 37 8.5–14.7 25 8.7–13.5

(11.4 ± 0.8) (11.7 ± 1.4) (11.4 ± 1.3) Dorsal fin attachment to caudal fin length 32 4–7.9 34 3.2–8.4 22 2.3–6.5

(5.8 ± 0.9) (5.9 ± 0.9) (4.4 ± 1.2) Anal-fin attachment to caudal fin length 32 4.1–8.1 34 3.9–8.2 21 3–6.9

(6.2 ± 1) (6.2 ± 1) (5.3 ± 1) Caudal base depth 18 1.3–2 11 1.3–2.4 1 2 26 1.1–2

(1.5 ± 0.2) (1.7 ± 0.3) (1.5 ± 0.2) tube length 10 0.2–0.7 11 0.7–1.5 17 0.1–1.1

(0.4 ± 0.2) (1.1 ± 0.3) (0.7 ± 0.2)

Meristics

Dorsal-fin rays 34 51–60 56 50–56 2 52–54 27 56–59

(55 ± 1.8) (53.1 ± 1.2) (53 ± 1.4) (57.6 ± 1) Anal-fin rays 34 46–53 56 43–49 2 47 27 48–51

(49.1 ± 1.5) (45.8 ± 1.3) (50.2 ± 0.9) Pectoral-fin rays 34 30–38 57 27–35 2 32–32 26 28–36

(33.6 ± 1.9) (30.7 ± 1.7) (32 ± 0) (31 ± 2.2) Pectoral-fin lower lobe rays 32 6–10 54 6– 12 1 8 26 7–10

(8.5 ± 1) (7.8 ± 0.9) (8.1 ± 0.7) rakers 26 9–13 51 6–10 2 8–9 24 6–10

(10.6 ± 0.9) (8.8 ± 1.1) (8.5 ± 0.7) (8.1 ± 1)

Caudal-fin rays 29 9–11 51 8– 11 1 9 21 9–9

(9.2 ± 0.5) (9 ± 0.4) (9 ± 0) Precaudal vertebrae 33 9–11 56 10–12 1 10 27 10–12

(9.9 ± 0.3) (11.1 ± 0.5) (10.9 ± 0.5) Caudal vertebrae 33 47–53 56 44–50 1 49 27 49–52

(49.5 ± 1.2) (46.6 ± 1.3) (51.1 ± 0.8) vertebrae 33 57–63 56 55–60 1 59 27 59–63

(59.5 ± 1.3) (57.7 ± 1.2) (62 ± 1) Dorsal-fin rays

50 51 52 53 54 55 56 57 58 59 60 61 62 63 n. rastrinus 1 4 11 8 6 1 31. trachysoma 4 3 8 16 6 1 38. scottae 1 12 36 42 19 10 2 1 123. acanthodes 4 8 5 6 23. pellucidus 1 1 3 10 6 7 4 1 1 34. phasma 2 2 10 23 12 7 1 57. spectrum 1 1 2. lerikimae 4 9 9 5 1 28

Anal-fin rays

43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 n. rastrinus 7 12 5 6 1 31. trachysoma 2 4 12 10 4 2 1 36. scottae 3 13 24 38 37 6 2 123. acanthodes 1 2 12 6 3 24. pellucidus 1 3 5 16 5 2 2 34. phasma 3 7 8 22 12 4 1 57. spectrum 2 2. lerikimae 2 2 12 12 28

Pectoral-fin rays

27 28 29 30 31 32 33 34 35 36 37 38 39 n

. rastrinus 1 5 4 9 1 5 4 2 31

. trachysoma 1 5 2 11 8 4 6 1 38

. scottae 18 19 13 30 19 19 2 1 2 123. acanthodes 2 3 2 4 8 3 1 1 24

. pellucidus 2 1 8 5 8 5 2 2 1 34

. phasma 1 5 10 9 16 8 7 1 1 58

. spectrum 2 2

. lerikimae 3 8 2 3 4 4 1 1 1 27

…… continued on the next page Vertebrae

55 56 57 58 59 60 61 62 63 64 65 66 67 n rastrinus 3 9 11 5 3 31 trachysoma 2 6 9 10 2 29 scottae 6 21 56 30 8 1 122 acanthodes 1 3 13 3 2 2 24 pellucidus 2 4 11 13 1 2 33 phasma 4 4 14 18 15 2 57 spectrum 1 1 lerikimae 1 1 4 13 9 1 29

Gill rakers

6 7 8 9 10 11 12 13 n

rastrinus 4 4 10 6 24

trachysoma 6 8 4 3 21

scottae 2 25 56 26 9 1 119

acanthodes 1 2 3 6

pellucidus 2 12 8 3 1 26

phasma 1 5 15 16 15 52

spectrum 1 1 2

lerikimae 1 6 8 9 1 25

Life history. Largest specimen examined was a 385.6 mm (FAKU 131691). The only ripe female with yolked eggs was 210 mm; males 146.5–285 mm were ripe with large testes. At least two sizes of eggs were present in the ripe female: yolked eggs were about 3.1 mm in diameter and white eggs had diameters of 0.5–1.5 mm.

Distribution. Careproctus rastrinus has been collected from the Sea of Okhotsk, from west of Kamchatka south to eastern Sakhalin Island and northeastern Hokkaido Island ( Fig. 6 ). Collection depths range from 114 to 217 m.

Etymology. The specific epithet is derived from the Latin rastrum, meaning “rake”, likely a reference to the dense covering of cactus-like prickles on the types.

Remarks. Prior to our examination of type material and the availability of tissues suitable for genetic analysis, C. rastrinus had been recognized as one of the most common liparids in the Bering Sea and Aleutian Islands. Herein, we recognize C. rastrinus to be limited in geographic range to the Sea of Okhotsk, replaced in the eastern Bering Sea and Aleutian Islands by the similar species C. scottae . Records of C. rastrinus from the Pacific Ocean east of the Kuril Islands into the western Bering Sea (e.g., Orlov 2005) require verification.

We reidentified one paratype of C. rastrinus as C. acanthodes . It was among the smallest collected but was readily identified as C. acanthodes by counts of vertebrae, the large size of the pelvic disc, and short lower pectoral-fin lobe.

Figures

FIGURE 1. Selected western Pacific species of the Careproctus rastrinus species complex: A) C. rastrinus, FAKU 131687, 320.7 mm; B) C. trachysoma, FAKU 130800, 214.4 mm, light morphotype; C) C. trachysoma, FAKU 131402, 264.8 mm, dark morphotype.

FIGURE 5. Pectoral girdles of species of the Careproctus rastrinus species complex: A) C. rastrinus, HUMZ 60772, 149.2 mm; B) C. trachysoma, FAKU 131179, 229.2 mm; C) C. scottae, UW 155110, 165 mm; D) C. acanthodes, FAKU 135569, 79.2 mm; E) C. pellucidus, FAKU 135317, 210.0 mm; F) C. phasma, UW 117934, 191 mm; G) C. lerikimae n. sp., paratype, UW 154841. Scale bar = 5 mm. c = coracoid, f = interradial fenestra, pr = proximal radial, dr = distal radial, s = scapula. Shaded areas represent cartilage.

FIGURE 6. Distribution of species of the Careproctus rastrinus species complex: C. rastrinus (□), C. trachysoma (●), C. scottae (○), C. acanthodes (◊), C. pellucidus (▲), C. phasma (■), C. spectrum (∆), and C. lerikimae n. sp. (♦), based on material examined. Each symbol may represent more than one capture.