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RANGE EXTENSIONS AND NEW RECORDS FROM ALASKA AND BRITISH COLUMBIA FOR TWO SKATES, BATHYRAJA SPINOSISSIMA AND BATHYRAJA MICROTRACHYS.

The skates (Rajiformes) of the eastern North Pacific Ocean and Bering Sea have been the subject of recent taxonomic and distributional studies, leading to the descriptions of new species (Stevenson and others 2004; Orr and others 2011) and the identification of other species newly documented in the region (Hoff 2002; Stevenson and Orr 2005; Stevenson and others 2007). Five genera of skates in 2 families (Arhynchobatidae and Rajidae) are known in the eastern North Pacific. Fifteen species in 4 genera have been recognized from Alaska (Stevenson and others 2007; Orr and others 2011; Ishihara and others 2012; Table 1), and morphological and molecular variation among specimens collected from the eastern North Pacific and Bering Sea suggest the presence of additional undescribed species in the area (Spies and others 2006, 2011; Stevenson and others 2007). Just to the south of Alaska, surveys of the ichthyofauna of British Columbia and in the Salish Sea have identified 10 species of skates in 4 genera (Ebert 2003; Pietsch and Orr 2015; King and others in press; Table 1).

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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Fisheries Science Center, Resource Ecology and Fisheries Management Division, 7600 Sand Point Way NE, Seattle, WA 98115 USA (IS); Fisheries and Oceans Canada, Pacific Biological Station, 3190 Hammond Bay Rd, Nanaimo, BC V9T6N7 Canada (JAB). Submitted 9 October 1018, accepted 17 November 2018. Corresponding Editor: Robert Hoffman.
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
Article Type:Report
Geographic Code:1U9AK
Date:Mar 22, 2019
Words:5783
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