RANGE EXTENSIONS AND NEW RECORDS FROM ALASKA AND BRITISH COLUMBIA FOR TWO SKATES, BATHYRAJA SPINOSISSIMA AND BATHYRAJA MICROTRACHYS.
Recent surveys of the continental slope in the eastern North Pacific and Bering Sea have broadened our understanding of the ichthyofauna in the region. Unusual specimens collected on the eastern Bering Sea slope have resulted in the descriptions of new species of fishes (Stevenson and Anderson 2005; Stevenson and Orr 2006; Baldwin and Orr 2010; Orr 2012) and the identification of others as new to Alaska or to the Bering Sea or both (Hoff 2002; Orr and Stevenson 2005; Orr and others 2005; Kenaley and Orr 2006; Maslenikov and others 2013). Specimens collected in recent surveys of the continental slope of British Columbia (Hanke and others 2015) have also resulted in additions to the Canadian ichthyofauna (Hanke and Roias 2012; Hanke and others 2014, 2015; Milkova and others 2016; King and others in press). Among these unusual specimens, an unidentified juvenile skate of the genus Bathyraja was collected in the eastern Bering Sea and later identified as B. spinosissima. This specimen represents the 1st record of the species from the Bering Sea and Alaska. In addition, a large adult skate collected off British Columbia, initially identified as B. abyssicola based on morphology and then as B. spinicauda based on DNA barcoding (Steinke and others 2009, supplement), was subsequently reidentified as B. spinosissima, a new record also for British Columbia. Additionally, we discovered 5 specimens of B. microtrachys, initially identified as B. violacea, from collections off the west coast of Vancouver Island, which represent the 1st records of the species from Canada.
Bathyraja spinosissima (Beebe and Tee-Van 1941), the Pacific White Skate, is a rare arhynchobatid skate known from only a few specimens around the world, ranging from the type locality off Cocos Island, Costa Rica, and the Galapagos Ridge to Waldport, Oregon, in the eastern Pacific (Beebe 1926; Provost and others 2015; Last and others 2016; Eschmeyer and others 2017; Salinas-de-Leon and others 2018). It has also been reported from the Sea of Okhotsk off Kamchatka (Dudnik and Dolganov 1992; Dolganov 1999; Dolganov and Tuponogov 1999; Sheiko and Fedorov 2000; Parin 2001; Fedorov and others 2003; Grigorov and Orlov 2013; Parin and others 2014; Ebert and others 2017) and the northern Kuril Islands (Pitruk and Fedorov 1990) in the western Pacific, although these records may represent a different species (Ebert 2003; Provost and others 2015; Last and others 2016). Living at depths of 800 to 2938 m, B. spinosissima is thought to be one of the deepest-dwelling skates worldwide (Ebert 2003; Salinas-de-Leon and others 2018), second only to B. microtrachys. Its preference for these great depths, which lie outside the range of most benthic survey efforts, combined with its preference for rocky habitat (Provost and others 2015), may explain why it is so rarely encountered.
Bathyraja microtrachys (Osburn and Nichols 1916), the Fine-spined Skate, is thought to be the deepest-dwelling skate, commonly found living below 2000 m to a maximum depth of 3300 m (Ebert and others 2017). It ranges from off central Baja California to Washington (Ebert 2003; Love and others 2005; Carlisle and others 2015; Last and others 2016; Ebert and others 2017). As for B. spinosissima, few specimens have been collected.
Methods of measuring and counting for this study follow Orr and others (2011). Specimens were either frozen or fixed in 10% formalin at sea and stored in 70% ethanol. Before whole specimens were preserved, tissues were taken for genetic analysis, preserved in 95% ethanol, and stored at--8O[degrees]C. Newly collected specimens are archived at the University of Washington, Burke Museum of Natural History and Culture, Fish Collection (UW) and the Royal British Columbia Museum (RBCM). The holotype of B. spinosissima is archived at the California Academy of Sciences (CAS). Measurements and counts of the holotype were recorded from the original description or taken from photographs and radiographs provided by CAS, available at https://www.calacademy.org/scientists/ichthyology-collections. Sequence data from the cytochrome c oxidase subunit I (COI) region were obtained following the procedures of Spies and others (2006) and were deposited in the Barcode of Life database (BOLD) and Genbank.
Bathyraja spinosissima (Beebe and Tee-Van 1941) Pacific White Skate; Figures 1-3
Description of New Specimen from Bering Sea.--UW 157183, 225 mm TL without caudal filament, 255 mm with caudal filament, male, 56.1039[degrees]N, l69.2300[degrees]W, 1066 m depth, 30 June 2010, F/V Vesteraalen, cruise 2010-01, haul 114, collected by GR Hoff and S Kotwicki, BOLD accession number NPSKT002-18, GenBank MH427012 (Fig. 1). Morphometric and meristic data are presented in Table 2.
Neonate with vestige of yolk sac. Disc wider than long, margins of pectoral fins broadly rounded; anterior lobe of pelvic fins slender, pointed; tail longer than disc; snout soft, flexible, especially near tip, internal supporting rostral cartilage slender and weak; teeth small, numerous, 32 rows in upper jaw; interorbital space wide, slightly convex; thorns absent from disc; tail thorns 29, interdorsal thorns 2; dorsal and ventral surface of head and body uniformly covered with small denticles; 2 small, well-separated dorsal fins, about equal in height and length; predorsal vertebrae between pelvic girdle and origin of first dorsal fin about 86.
In life, dorsal surface uniform dark plum gray (Fig. lA); ventral surface dark brown with patches of white around mouth, gill slits, and cloaca, white band at base of tail; clasper tips white; caudal filament with dark dorsal ridge, pale to translucent ventrally (Fig. 1B). In preservation, dorsal surface uniform dark; ventral surface dark with light areas around mouth, gill slits, and cloaca, light band at base of tail; clasper tips light.
Description of New Specimen from British Columbia. --RBCM 16121, 1461 mm TL, 48.5940[degrees]N, l26.9008[degrees]W, 1951 m depth, 18 October 2005, F/V Frosti, haul 20, benthic trawl, collected by J Boutillier, BOLD accession number TZFPB359-05 (Fig. 2). Morphometric and meristic data are presented in Table 2.
Adult male with well-developed claspers. Disc wider than long, margins of pectoral fins broadly rounded; anterior lobes of pelvic fins slender, pointed; disc longer than tail; snout soft, flexible, especially near tip, internal supporting rostral cartilage slender and weak; teeth small, numerous, 28 rows in upper jaw; interorbital space wide, flat to slightly concave; thorns absent from disc; tail thorns 29, interdorsal thorns 0; dorsal and ventral surface of head and body uniformly covered with small denticles; dorsal fins small, well separated, about equal in height and length.
In life, dorsum uniform dark brownish gray, slightly darker at pectoral and pelvic fin margin (Fig. 2A); ventrum dark brownish gray, margin of gill openings, pectoral and pelvic fins slightly darker, with patches of white around the mouth, cloaca, and clasper tips. In preservation, dorsum uniform dark brown, slightly darker at pectoral-and pelvic-fin margins (Fig. 2B); ventrum dark brown, margin of gill openings and pectoral and pelvic fins slightly darker, with patches of white around the mouth, cloaca, and clasper tips (Fig. 2C).
Sequence Data.--The specimen from British Columbia (RBCM 16121, previously as RBCM 006-034-024) was originally identified as Bathyraja abyssicola, the only other species of skate known from British Columbia waters with denticles on the ventral surface, but is 1.6% divergent from B. abyssicola sequences in BOLD (MFC328-08, TZFPB311-05, TZFPB421-05). The identification was later changed to B. spinicauda because the sequence is only 0.6% divergent from numerous sequences labeled as B. spinicauda in BOLD (Steinke and others 2009, supplement; GH, pers. obs.; GBGC10833-10861, RNEZ001-010), even though B. spinicauda is known only from the North Atlantic and Barents Sea. The COI sequence obtained for RBCM 16121 is 100% identical (652/652 bases) to the COI sequence obtained for the specimen from the Bering Sea (UW 157183). There are no previously documented reference sequences in BOLD or GenBank for B. spinosissima with which to compare these sequences.
Discussion.--The holotype of B. spinosissima (CAS-SU 46500; Fig. 3) is a neonate that was collected enclosed in its egg case, but the egg case has since been lost (Ebert and Davis 2007). It is very similar to the specimen reported here from the Bering Sea. Nearly all proportional measurements and meristic counts are close to or identical to those of the holotype (Table 2). Slight differences from published descriptions are the presence of 2 interdorsal thorns (rather than their absence; Ebert 2003) and a higher count of about 86 to 90 predorsal vertebrae (vs. 82; Ebert 2003).
Bathyraja spinosissima is similar to the Deepsea Skate, B. abyssicola, of the North Pacific in having the ventral surface covered with denticles, forming a shagreen-like surface. However, it differs from B. abyssicola in lacking nuchal and middorsal thorns, which are both present and relatively strong in B. abyssicola, and in having more slender and pointed anterior pelvic-fin lobes, a narrower interspiracular width in preorbital snout length of 0.8-1.3 (Muler and Lea 1972; new data) (vs. 1.4-2.7 in B. abyssicola; Miller and Lea 1972; Zorzi and Anderson 1988; Stevenson and Orr 2005), and a wider interorbital width of 5.8-6.6 % TL (Table 2) vs. 3.8-4.6 % TL in B. abyssicola (Stevenson and Orr 2005). It can also be distinguished from other skates in the area by its lack of scapular thorns and in having a uniform salty gray coloration both dorsally and ventrally in adults, as well as a ventral surface covered with denticles (Ebert 2003).
Batliyraja microtrachys (Osburn and Nichols 1916) Fine-spined Skate; Figure 4
Description of New Specimens from British Columbia.--Five specimens (RBCM 005-0137-001, -002, -003, -004 [Fig. 4], -005); 670-695 mm TL, SW of Tofino, Vancouver Island, 48.5830[degrees]N, l26.8838[degrees]W, 1939-1951 m depth, 10 October 2005, F/V Frosti, haul 20, benthic trawl, collected by J Boutillier. BOLD accession numbers are TZFPB352-05, TZFPB353-05, TZFPB355-05, duplicated records at RBCM. Morphometric TZFPB351-05, and TZFPB354-05, respectively, and meristic data are presented in Table 3; 1 Catalog numbers in BOLD differ from the specimen (RBCM 005-0137-001) with a truncated correct numbers presented here because of tail was not included.
All adult males with fully developed claspers. Disc wider than long, anterior margin moderately to strongly scalloped, apices of pectoral fins moderately rounded; anterior lobes of pelvic fins broad; disc shorter than tail; snout soft, flexible, especially near tip, internal supporting rostral cartilage slender and weak; teeth small, numerous, 21-28 rows in upper jaw; interorbital space flat to slightly concave; thorns absent from disc; tail thorns 19-24, interdorsal thorns 0; dorsal surface of head and body uniformly covered with small denticles, ventral surface naked; dorsal fins small, well separated, about equal in height and length.
In life and preservation, dorsal surface uniform light to darker brown (Fig. 4AB); ventral surface of body mostly dark (Fig. 4C), central part of body and anterior lobes of pelvic fins white, with scattered darker mottling, ventral surface of pectoral fins from their bases to tips dark to lighter brown, area around cloaca to base of tail and posterior lobes of pelvic fins typically dark, claspers dark except for light tips; tail entirely dark in 3 specimens, with lighter mottling in 2 specimens.
Sequence Data.--The specimens from British Columbia were originally identified as Bathyraja violacea. However, sequence data are 3.3% divergent from the B. violacea sequence in the GenBank and BOLD databases (DQ665298.2/GBGC1479-06), which was taken from a specimen (UW 113488) identified and sequenced as B. violacea by Spies and others (2006) and Stevenson and others (2007). This sequence is the only publicly available data from a confirmed specimen of B. violacea.
Discussion.--Other similar skates known from deep waters (>2OOO m) of the region include Amblyraja badia (Garman), Bathyraja abyssicola, B. trachura (Gilbert), and now B. spinosissima. All except B. trachura and B. spinosissima have thorns on the disc, and B. abyssicola and B. spinosissima have denticles on the ventrum, unlike B. microtrachys. In B. trachura, the tail is longer than the disc length, larger denticles are present on the disc and especially the tail, and the dorsal surface is darker brown to black--all characters that differ from B. microtrachys.
Presumably because of the absence of thorns on the disc combined with its brown dorsal coloration, Steinke and others (2009, supplement) and Coulson and others (2011) misidentified these specimens as B. violacea, and Coulson and others (2011) questioned the identifications of B. violacea by Spies and others (2006) and Stevenson and others (2007). However, Suvorov (1935), author of the original description of B. violacea, described the species (as Raja violacea) from the Sea of Okhotsk based on 2 specimens, captured at depths of 47 m and 100 m. The English summary of this description includes the phrase "Ventral side white" (Suvorov, 1935:433). The specimen (UW 113488) reported by Spies and others (2006) and Stevenson and others (2007) was collected in the western Aleutian Islands within 2000 km of the type locality, at a depth of 261 m, and has a white ventrum. In contrast, the 5 specimens labeled "Bathyraja violacea" by Steinke and others (2009, supplement) and Coulson and others (2011) were collected some 9000 km from the type locality, at a depth of 1945 m, and have a heavily pigmented ventrum with a distinct white central band, as described for B. microtrachys by Last and others (2016). In addition, the specimen identified by Spies and others (2006) and Stevenson and others (2007) matches the original and other descriptions of B. violacea in all other particulars, including most significantly in lacking a pseudosiphon (Ishihara and Ishiyama 1985; Stevenson and others 2004). The 5 specimens from British Columbia are males with a clearly defined pseudosiphon. These specimens also have a COI sequence 3.3% divergent from the B. violacea from the Aleutian Islands. Thus, we agree with the assessment of Coulson and others (2011) that "a possible misidentification" lies at the root of this discrepancy, but the misidentification was theirs. The specimen from the Aleutian Islands (UW 113488) represents B. violacea, and based on our examination of the specimens, the RBCM material from British Columbia represents B. microtrachys.
Acknowledgments.--We thank K Maslenikov and L Tornabene for collections support, G McFarlane for discussions, and T Pietsch, N Laman, and M Love for their critical reviews of the manuscript. This project was funded in part by the North Pacific Research Board, grant number 1509.
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NOAA, National Marine Fisheries Service, Alaska Fisheries Science Center, Resource Assessment and Conservation Engineering Division, 7600 Sand Point Way NE, Seattle, WA 98115 USA (JWO, DES, GRH); firstname.lastname@example.org; Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia, BC V8W 9W2 Canada (GH); NOAA, National Marine Fisheries Service, Alaska
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TABLE 1. Skates (Rajiformes) reported from waters of Alaska and British Columbia, supported by archived specimens. UBC = University of British Columbia, Beatty Biodiversity Museum. Species Common name Geographic range Amblyraja badia Broad Skate Arctic Ocean, Bering Sea to Central America, western Pacific Bathyraja Deepsea Skate Bering Sea to southern abyssicola California, western Pacific B. aleutica Aleutian Skate Bering Sea to northern California, western Pacific B. interrupta Bering Skate Bering Sea to Washington B. kincaidii Sandpaper Skate British Columbia to California B. lindbergi Commander Skate Bering Sea to British Columbia, western Pacific B. maculata Whiteblotched Skate Bering Sea to Gulf of Alaska, western Pacific B. mariposa Butterfly Skate central Aleutian Islands B. microtrachys Fine-spined Skate British Columbia to California B. minispinosa Whitebrow Skate Bering Sea to Gulf of Alaska, western Pacific B. panthera Leopard Skate western Aleutian Islands B. parmifera Alaska Skate Bering Sea to British Columbia, western Pacific B. spinosissima Pacific White Skate Bering Sea to California, western Pacific B. taranetzi Mud Skate Bering Sea, Aleutian Islands, western Pacific B. trachura Roughtail Skate Bering Sea to Baja California, western Pacific B. violacea Okhotsk Skate western Aleutian Islands, western Pacific Beringraja Big Skate Bering Sea to Baja California binoculata Raja rhina Longnose Skate Bering Sea to Baja and Gulf of California Species Depth range (m) Citation Amblyraja badia 1061-2322 Stevenson and others (2007) Bathyraja 362-2904 Stevenson and others (2007) abyssicola B. aleutica 15-1602 Stevenson and others (2007) B. interrupta 26-1050 Stevenson and others (2007); Pietsch and Orr (2015) B. kincaidii 18-500 Pietsch and Orr (2015) B. lindbergi 126-1193 Stevenson and others (2007); King and others (in press) B. maculata 73-1193 Stevenson and others (2007) B. mariposa 90-448 Stevenson and others (2004, 2007) B. microtrachys 2000-3300 this publication B. minispinosa 150-1420 Stevenson and others (2007) B. panthera 48-396 Orr and others (2011) B. parmifera 17-392 Orr and others (2011); UBC 13720 B. spinosissima 800-2938 this publication B. taranetzi 58-1054 Stevenson and others (2007) B. trachura 213-2550 Stevenson and others (2007) B. violacea 124-510 Stevenson and others (2007) Beringraja 16-402 Stevenson and binoculata others (2007); Pietsch and Orr (2015) Raja rhina 9-1294 Stevenson and others (2007); Pietsch and Orr (2015) TABLE 2. Measurements in millimeters (mm), proportions of total length (TL) with or without caudal filament (cf), and merist!c counts for the holotype of Bathyraja spinosissima and new specimens from the eastern Bering Sea and British Columbia. Values for the holotype were taken from the original description, counted from the radiograph and photo, or measured from the photo (in bold). Missing entries for the holotype were not available. RBCM 16121 was not radiographed; vertebral counts are not known. UW 157183 CAS-SU 46500 mm % TL w/o cf %TL w/cf mm %TL w/cf Total length 225 - - - - Total length 255 - - 248 - + caudal filament Disk width 146 64.89 57.25 145 58.47 Disk length 117 52.00 45.88 115 46.37 Snout to end of 118 52.44 46.27 125 50.40 pelvic fin length Snout to maximum 72 32.00 28.24 72 29.03 disk width Dorsal snout length 27 12.00 10.59 23 9.27 Head length 40.6 18.04 15.92 30 12.10 Orbit length 8.6 3.82 3.37 8 3.23 Eyeball length 6.8 3.02 2.67 7 2.82 Orbit to spiracle 12.1 5.38 4.75 12 4.84 length Spiracle length 4.6 2.04 1.80 5 2.02 Interorbital width 14.8 6.58 5.80 14 5.65 IOW (outer orbital 28.6 12.71 11.22 23 9.27 margin) Interspiracular 21.6 9.60 8.47 25 10.08 width Pretail length 104 46.22 40.78 103 41.53 Tail length to D1 92 40.89 36.08 82 33.06 origin Tail length to D2 106 47.11 41.57 92 37.10 origin Interdorsal length 4.8 2.13 1.88 3 1.21 Tail length 125 55.56 49.02 135 54.44 Caudal-fin length - - - - - Ventral snout length 22.6 10.04 8.86 27 10.89 Prenasal length 21.5 9.56 8.43 20 8.06 Snout to gill slit 59 26.22 23.14 48 19.35 5 length Snout to gill slit 48.5 21.56 19.02 47 18.95 1 length Mouth width 26.2 11.64 10.27 21 8.47 Internarial distance 20.3 9.02 7.96 23 9.27 Nasal curtain length 6.9 3.07 2.71 7 2.82 Width of 1st gill slit 3.2 1.42 1.25 3 1.21 Width of 5th gill slit 2.7 1.20 1.06 2 0.81 Distance between 45.8 20.36 17.96 44 17.74 1 st gill slits Distance between 32.8 14.58 12.86 30 12.10 5th gill slits Anterior pelvic-fin 19.4 8.62 7.61 17 6.85 lobe length Posterior pelvic-fin 27.8 12.36 10.90 25 10.08 lobe length Tail width at axil 9.5 4.22 3.73 13 5.24 Dorsal-fin 1 length 6.4 2.84 2.51 6 2.42 Dorsal-fin 1 height 3.1 1.38 1.22 - - Dorsal-fin 2 length 6.6 2.93 2.59 7 2.82 Dorsal-fin 2 height 3.9 1.73 1.53 - - Dorsal-fin 1 to 28.3 12.58 11.10 39 15.73 caudal tip Dorsal-fin 2 to 16.9 7.51 6.63 - - caudal tip Teeth 32 - - 30 - Pseudobranchial 17 - - - - folds Predorsal vertebrae ~86 - - ~90 - Vertebrae (total) 107 - - 112 - Tail thorns 29 - - 25 - Interdorsal thorns 2 - - 0 - Total thorns 31 - - 25 - RBCM 16121 mm % of TL Total length 1461 - Total length 1461 - + caudal filament Disk width 906 62.01 Disk length 773 52.90 Snout to end of 824 56.39 pelvic fin length Snout to maximum 487 33.33 disk width Dorsal snout length 165 11.29 Head length 275 18.82 Orbit length 43 2.94 Eyeball length 23 1.57 Orbit to spiracle 65 4.44 length Spiracle length 37 2.53 Interorbital width 151 10.33 IOW (outer orbital 172 11.77 margin) Interspiracular 150 10.26 width Pretail length 735 50.30 Tail length to D1 530 36.27 origin Tail length to D2 585 40.04 origin Interdorsal length 7 0.47 Tail length 618 42.29 Caudal-fin length 26 1.77 Ventral snout length 147 10.06 Prenasal length 145 9.24 Snout to gill slit 400 27.37 5 length Snout to gill slit 315 21.56 1 length Mouth width 175 11.97 Internarial distance 142 9.71 Nasal curtain length 29 1.98 Width of 1st gill slit 32 2.19 Width of 5th gill slit 22 1.50 Distance between 294 20.12 1 st gill slits Distance between 202 13.82 5th gill slits Anterior pelvic-fin 40 2.73 lobe length Posterior pelvic-fin 96 6.57 lobe length Tail width at axil 64 4.38 Dorsal-fin 1 length 58 3.96 Dorsal-fin 1 height 18 1.23 Dorsal-fin 2 length 61 4.17 Dorsal-fin 2 height 20 1.36 Dorsal-fin 1 to 141 3.16 caudal tip Dorsal-fin 2 to 82 5.61 caudal tip Teeth 28 - Pseudobranchial 18 - folds Predorsal vertebrae - - Vertebrae (total) - - Tail thorns 29 - Interdorsal thorns 0 - Total thorns 29 - TABLE 3. Proportional measurements and meristic counts for new specimens of Bathyraja microtrachys from British Columbia. Morphometric data are given in percent TL (total length) and presented as the range, followed by the mean +/- standard deviation (SD). Category n Range Mean [+ or -] SD Total length 4 670-695 680 [+ or -] 10.8 Disk width 4 55.6-59.3 56.8 [+ or -] 1.7 Disk length 4 48.9-50.8 50.1 [+ or -] 0.9 Snout to 4 29.6-33.1 31.2 [+ or -] 1.8 maximum disk width Dorsal snout 4 12.6-13.3 12.9 [+ or -] 0.3 length Head length 4 18.5-20.1 19.3 [+ or -] 0.6 Orbit length 4 2.4-3.0 2.8 [+ or -] 0.2 Eyeball length 4 1.6-2.2 2.0 [+ or -] 0.3 Orbit to spiracle 4 4.2-4.8 4.5 [+ or -] 0.2 length Spiracle length 4 2.0-2.4 2.2 [+ or -] 0.2 Interorbital width 4 4.1-5.5 4.7 [+ or -] 0.6 Interorbital width 4 5.9-6.7 6.3 [+ or -] 0.4 (outer orbital margin) Interspiracular 4 6.6-7.7 7.1 [+ or -] 0.5 width Pretail length 4 45.9-18.8 47.5 [+ or -] 1.2 Tail length to 4 38.8-12.2 40.9 [+ or -] 1.5 D1 origin Tail length to 4 44.6-47.1 45.9 [+ or -] 1.1 D2 origin Interdorsal 4 0.9-1.4 1.0 [+ or -] 0.2 length Tail length 4 51.3-54.1 53.0 [+ or -] 1.2 Caudal-fin 4 1.9-4.5 3.2 [+ or -] 1.1 length Ventral snout 4 11.1-12.2 11.8 [+ or -] 0.5 length Prenasal length 4 10.2-11 10.7 [+ or -] 0.3 Snout to gill 4 25.9-27.2 26.7 [+ or -] 0.6 slit 5 length Mouth width 4 9.3-9.8 9.5 [+ or -] 0.3 lnternarial 4 7.5-8.8 8.3 [+ or -] 0.6 distance Nasal curtain 4 3.4-4.1 3.7 [+ or -] 0.4 length Width of 1st 4 1.3-1.3 1.3 [+ or -] 0 gill slit Width of 5th 4 0.9-1.0 0.9 [+ or -] 0.1 gill slit Distance between 4 14-15.6 14.6 [+ or -] 0.7 1st gill slits Distance between 4 10.1-10.9 10.4 [+ or -] 0.4 5th gill slits Anterior pelvic-fin 4 9.2-9.7 9.4 [+ or -] 0.2 lobe length Posterior pelvic-fin 4 15.6-16.2 15.9 [+ or -] 0.2 lobe length Tail width at axil 4 4.1-5.0 4.6 [+ or -] 0.4 Dorsal-fin 1 length 4 3.3-4.9 3.9 [+ or -] 0.7 Dorsal-fin 1 height 4 1.6-2.2 2.0 [+ or -] 0.3 Dorsal-fin 2 length 4 3.3-4.1 3.7 [+ or -] 0.4 Dorsal-fin 2 height 4 1.9-2.4 2.1 [+ or -] 0.2 Dorsal-fin 1 to 4 9.9-14.5 11.9 [+ or -] 1.9 caudal tip Dorsal-fin 2 to 4 5.5-9.1 7.1 [+ or -] 1.5 caudal tip Teeth 4 21-28 23.8 [+ or -] 3.1 Tail thorns 4 19-24 21.3 [+ or -] 2.2 Interdorsal thorns 4 0 0 Pseudobranchial 4 12-17 13.8 [+ or -] 2.4 folds Total thorns 4 19-24 21.3 [+ or -] 2.2
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|Title Annotation:||NORTHWESTERN NATURALIST|
|Author:||Orr, James W.; Stevenson, Duane E.; Hanke, Gavin; Spies, Ingrjd B.; Boutillier, James A.; Hoff, Gera|
|Publication:||Northwestern Naturalist: A Journal of Vertebrate Biology|
|Date:||Mar 22, 2019|
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