Cottus schitsuumsh, Lemoine, Michael, Young, Michael K., Mckelvey, Kevin S., Eby, Lisa, Pilgrim, Kristine L. & Schwartz, Michael K., 2014
publication ID |
https://doi.org/ 10.11646/zootaxa.3755.3.3 |
publication LSID |
lsid:zoobank.org:pub:5147B3DB-9071-408B-A8D1-B3575ED5806E |
DOI |
https://doi.org/10.5281/zenodo.6137307 |
persistent identifier |
https://treatment.plazi.org/id/B07387BF-FFD9-FF92-FF17-C24DFB4BFFB2 |
treatment provided by |
Plazi |
scientific name |
Cottus schitsuumsh |
status |
sp. nov. |
Cottus schitsuumsh View in CoL , new species
Cedar Sculpin
Holotype. UW 151670, male, 91 mm SL, haplotype group 1 ( JX311895 View Materials ), East Fork Steamboat Creek, a tributary of the Coeur d’Alene River at West Fork Road ( USFS Road 409) (47.71821°N, 116.20010°W), Shoshone County, Idaho, 29 September 2011.
Paratypes. UW 151671, 2 males and 1 female, 57–73 mm SL, haplotype group 1 ( JX311896 View Materials ), CIDA 101,703, 2 males and 1 female, 72–99 mm SL, collected with the holotype.
Additional materials (nontypes, examined morphologically and genetically). Idaho, Shoshone County: (5, 66– 81 mm SL, haplotype group 2, JX311919 View Materials ) Bird Creek (47.27235°N, 115.60885°W), 29 September 2011; (5, 68–80 SL, haplotype group 1, JX311898 View Materials ) Leiberg Creek (47.72100°N, 116.38335°W), 6 September 2011; (5, 64– 88 mm SL, haplotype group 2, JX311921 View Materials ) Red Ives Creek (47.06016°N, 115.33448°W), 29 September 2011; (5, 69– 96 mm SL, haplotype group 1, JX311902 View Materials ) West Fork Steamboat Creek (47.71718°N, 116.20380°W), 7 September 2011. Tissue samples (and extracted DNA) are archived at the Wildlife Genetics Laboratory, Rocky Mountain Research Station, Missoula, MT.
Additional materials (nontypes, examined morphologically). Idaho, Benewah County: (2, 50– 70 mm SL, CIDA 68425) Renfro Creek (47.16278°N, 116.40462°W), 11 July 1996. Idaho, Shoshone County: (5, 69– 84 mm SL) Beaver Creek (47.08260°N, 115.35738°W), 29 September 2011; (1, 62 mm SL, CIDA 69113) Buckskin Creek (47.99148°N, 116.22247°W), 24 July 1997; (2, 71– 86 mm SL, CIDA 68475) Eagle Creek (47.64813°N, 115.91920°W), 23 August 1996; (5, 71– 83 mm SL) East Fork Steamboat (47.71821°N, 116.20010°W), 29 September 2011; (5, 72– 79 mm SL) Leiberg Creek (47.72100°N, 116.38335°W), 6 September 2011; (10, 44– 95 mm SL) Steamboat Creek (47.69685°N, 116.167867°W), 18 October 2011; (4, 70– 97 mm SL) West Fork Steamboat Creek (47.71718°N, 116.2038°W), 18 October 2011; (1, 73 mm SL, UW 152969) West Fork Steamboat Creek (47.71718°N, 116.2038°W), 7 September 2011. Montana, Mineral County: (5, 48– 62 mm SL) Fish Creek (46.951°N, 114.67359°W), 12 October 2011; (5, 50– 76 mm SL) Fish Creek (46.96693°N, 114.65292°W), 12 October 2011; (5, 45– 67 mm SL) St. Regis River (47.43131°N, 115.74357°W), 15 October 2011; (5, 48– 86 mm SL) Trout Creek (47.09768°N, 114.90475°W), 12 October 2011; (5, 51– 67 mm SL) Trout Creek (47.13611°N, 114.85036°W), 12 October 2011; (5, 51– 91 mm SL) Trout Creek (47.14311°N, 114.83298°W), 12 October 2011; (6, 49– 83 mm SL) Twomile Creek (47.295958°N, 115.171311°W), 12 October 2011. Montana, Sanders County: (8, 53– 77 mm SL) Prospect Creek (47.57452°N, 115.39381°W), 15 October 2011.
Additional materials (nontypes, examined genetically). Haplotype group 1. Idaho, Kootenai County: (2, JX282597 View Materials ) 4th of July Creek (47.58208°N, 116.46721°W); (2, JX311897 View Materials ) Hayden Creek (47.82378°N, 116.65348°W). Idaho, Shoshone County: (2, JX311893 View Materials ) Bumblebee Creek (47.63522°N, 116.27645°W); (2, JX311899 View Materials ) Moon Creek (47.55719°N, 116.03062°W); (2, JX311900 View Materials ) Rampike Creek (47.81965°N, 115.96189°W); (2, JX311901 View Materials ) West Fork Eagle Creek (47.67772°N, 115.88087°W). Montana, Mineral County: (2, JX311903 View Materials ) Big Creek (47.37580°N, 115.39283°W). (1, JX311904 View Materials ) Burdette Creek (46.81331°N, 114.62097°W). (1, JX311905 View Materials ) Burdette Creek (46.83436°N, 114.60488°W). (2, JX311907 View Materials ) East Fork Packer Creek (47.43260°N, 115.49248°W). (2, JX311908 View Materials ) Savenac Creek (47.39438°N, 115.39210°W). (1, JX311909 View Materials ) Savenac Creek (47.43467°N, 115.39532°W). (1, JX311910 View Materials ) Savenac Creek (47.46251°N, 115.38812°W). (2, JX311911 View Materials ) St. Regis River (47.41834°N, 115.62450°W). (1, JX311912 View Materials ) St. Regis River (47.43131°N, 115.74357°W). (1, JX311913 View Materials ) St. Regis River (47.43675°N, 115.68220°W). (2, JX311914 View Materials ) Straight Creek (46.91092°N, 114.81761°W). (2, JX311915 View Materials ) Twelvemile Creek (47.37603°N, 115.25982°W). (1, JX311915 View Materials ) Twelvemile Creek (47.42681°N, 115.24120°W). Montana, Sanders County: (2, JX311917 View Materials ) Wilkes Creek (47.55313°N, 115.42111°W). Haplotype group 2. Idaho, Shoshone County: (2, JX282598 View Materials ) Beaver Creek (47.08314°N, 115.35790°W). (2, JX311918 View Materials ) Bird Creek (47.27229°N 115.60893°W). (2, JX311920 View Materials ) North Fork St. Joe River (47.35768°N, 115.73658°W). (2, JX311922 View Materials ) Simmons Creek (47.13888°N, 115.39229°W). (2, JX311923 View Materials ) St. Joe River (47.04652°N, 115.18094°W). Haplotype group 3. Idaho, Shoshone County: (2, JX282599 View Materials ) Bechtel Creek (46.99509°N, 116.28479°W). Tissue samples (and extracted DNA) are archived at the Wildlife Genetics Laboratory, Rocky Mountain Research Station, Missoula, MT. See Young et al. (2013; table S1) for additional materials used in this analysis.
Morphological diagnosis. Cottus schitsuumsh is diagnosed from all other Cottus in the upper Columbia River basin by the lack of a second preopercular spine, palatine tooth patches smaller and separate from the vomerine tooth patch, and axillary prickling.
Cottus schitsuumsh is distinguished from C. cognatus , C. confusus , C. hubbsi , and C. rhotheus by the lack of a second preopercular spine. Cottus schitsuumsh is distinguished from C. cf. bairdii and C. beldingii by the presence of axillary prickling. In addition, C. schitsuumsh has palatine tooth patches smaller than the vomerine tooth patch and the patches are separated. Cottus cf. bairdii has palatine tooth patches equal to or larger in size than the vomerine tooth patch. Cottus beldingii generally lacks palatine teeth.
Molecular diagnosis. Haplotypes of C. schitsuumsh differed from those of all other members of the genus Cottus (represented in GenBank or in this study) at three positions ( Table 1 View TABLE 1 ). Additionally, they differed from all other North American Cottus haplotypes at an additional position. Nucleotides at two other positions were diagnostic for individual haplotypes.
1Based on C. reinii mitome (GenBank accession AP004442 View Materials ; Miya et al. 2003).
2Relative to all Cottus haplotypes in GenBank (n = 104 from 283 accessions) and from Cottus collected throughout the upper Columbia and Missouri River basins ( Young et al. 2013) and in this study (n = 44).
Description. Cottus schitsuumsh is a moderately sized species of Cottus rarely exceeding 90 mm SL with anal fin rays 14 (rarely 13 or 15), the final posterior anal-fin ray commonly branched; dorsal-fin spines 8 (rarely 7 or 9); dorsal-fin rays 17–20 (usually 18–19) with the posterior dorsal-fin ray branched; pectoral-fin rays 13–14 (usually 14); and pelvic-fin rays 4 (rarely 3) and of similar length. There are 2 medial-chin pores, 10–12 preoperculomandibular pores, and 2 postmaxillary pores (rarely 0). Dorsal fins are generally not connected but occasionally have a slight connection at the base. Prickling is consistent and restricted to a trapezoidal area, widest in the axillary region and narrowest along the lateral line under the beginning of the second dorsal fin. Palatine teeth are present in two or fewer rows and sometimes on only one side. The palatine tooth patches are smaller than and separated from the vomerine tooth patch and generally equal if present on both sides. The lateral line is incomplete, ending below the latter half of the second dorsal fin. One preopercular spine was observed among all specimens. Preopercular spines are greatly reduced and skin-covered. The mouth width/head width fraction ranges from ½ to 2/3.
Morphometrics of the holotype (91 mm SL, male) and 3 paratypes (57–73 mm SL, 2 males and 1 female) as a percentage of SL (mean and range): caudal peduncle depth (8.8, 8.1–10.6); caudal peduncle length (13.9, 12.3– 17.4); head length (19.7, 16.8–23.4); head width (26.8, 21.6–34.3); lateral line length (59.5, 51.3–70.6); snout length (9.0, 6.5–17.4); mouth width (14.2, 12.9–15.9); interpelvic width (3.5, 2.3–4.6); longest pectoral ray (24.3, 21.2–33.7), longest pelvic ray (18.5, 15.1–24.4).
Mottling of C. schitsuumsh is variable and consists of at least two forms. One has black-brown mottling on a brown-olive background mainly above the lateral line that stops under the soft dorsal fin ( Fig. 4 View FIGURE 4 ), and weak saddling is occasionally present (Fig. 5). The pectoral fins have faint black-brown contours running vertically. The other form lacks mottling or spotting over the body and the background color can be light brown, rust orange, or yellow. Both types have a dark blotchy marking near the base of the caudal rays and the first dorsal fin is generally dorsally darkened. Both forms are sympatric in the upper Spokane River basin, Idaho, but the mottled form has only been observed in Montana.
Comparisons. Cottus schitsuumsh tended to have more anal fin rays (mode, 14 rays) than the other species (modes, 12 or 13 rays) among our samples ( Table 2 View TABLE 2 ) and relative to other sculpins in this region ( Wydoski and Whitney 2003). Pelvic ray counts are similar among the observed Cottus species, but C. cognatus commonly had a greatly reduced fourth pelvic-fin ray (cf. Wydoski & Whitney 2003). Cottus rhotheus has more pectoral rays than C. schitsuumsh . Counts of medial-chin pores did not differ among species, although C. schitsuumsh tended to have more preoperculomandibular pores than other species and more postmaxillary pores than C. cf. bairdii , C. beldingii , and C. cognatus ( Table 3 View TABLE 3 ). Cottus schitsuumsh can be further distinguished from C. cognatus and C. beldingii by the presence of palatine teeth ( Table 4 View TABLE 4 ). The palatine tooth patch of C. schitsuumsh consists of two or fewer rows, may occur on only one side, and is about ½ the size of the vomerine tooth patch. The palatine tooth patch is similar to or larger than the vomerine tooth patch in C. rhotheus , C. hubbsi , and C. cf. bairdii . Cottus schitsuumsh also differs from C. hubbsi and C. rhotheus by having an incomplete lateral line.
Originally confused with C. confusus ( Bailey & Bond 1963) , C. schitsuumsh can be distinguished from it by having only one preopercular spine that is small (length less than half the eye diameter) and skin-covered ( Fig. 6 View FIGURE 6 ). Most specimens of C. schitsuumsh also lack lateral-line pores on the caudal peduncle ( Fig. 7 View FIGURE 7 ). Pores along the lateral line of C. confusus , usually end beneath the latter half of the second dorsal fin and reappear at the end of the caudal peduncle near the origin of the caudal rays.
1Fourth ray in the pelvic fin of C. cognatus View in CoL reduced (ca. ½ of other rays; cf. Wydoski & Whitney 2003).
Most morphometric features separated C. schitsuumsh View in CoL from one or more of the Cottus View in CoL species. Caudalpeduncle depth, head length, mouth width, and interpelvic width statistically differed more often between C. schitsuumsh View in CoL and the other species than did other measures ( Table 5 View TABLE 5 ). Frequency distributions of these features emphasized these differences ( Tables 6 View TABLE 6 , 7 View TABLE 7 ). With respect to C. confusus View in CoL , C. schitsuumsh View in CoL differed by having a shorter lateral line, wider mouth, longer snout, and narrower interpelvic width.
Standardized head length Distribution and ecology. Cottus schitsuumsh View in CoL is distributed in all streams sampled in the upper Coeur d’Alene and St. Joe River drainages of the Spokane River basin above Post Falls near Post Falls, Idaho, and sporadically in the middle Clark Fork River basin between Fish Creek and Prospect Creek, approximately between Tarkio and Thompson Falls, Montana ( Fig. 1 View FIGURE 1 ). This species is common to abundant in cool to cold tributaries with cobble and gravel bottoms. Cottus schitsuumsh View in CoL is sympatric with Westslope Cutthroat Trout, Oncorhynchus clarkii lewisi View in CoL , and Bull Trout, Salvelinus confluentus View in CoL , throughout its distribution, with C. rhotheus View in CoL in Idaho, and with C. cognatus View in CoL in Montana. No effort was undertaken to find this species in Lake Coeur d’Alene or in the river main stems, but haplotype differences between fish from the Coeur d’Alene and St. Joe River basins imply that lineages in these basins are isolated and diverging.
The distribution in Idaho is consistent with that of other terrestrial and aquatic fauna e.g., Idaho Giant Salamander, Plethodon idahoensis View in CoL , that persisted immediately south of the extent of Cordilleran glaciation in the Northern Rocky Mountain Refugium ( Carstens et al. 2005; Shafer et al. 2010). The disjunct distribution in the Clark Fork River basin in western Montana is puzzling (although known for some time; Hendricks 1997). There are no known peri- or post-glacial hydrological connections between these basins because all mountain divides are more than 150 m above the glacial Lake Missoula highstand ( Smith 2006). Because sculpins were a common and legal baitfish in Montana until this century, it is plausible that populations in western Montana resulted from human-assisted translocation. Regardless of their means of arrival, the Montana populations have haplotypes similar to those in the Coeur d’Alene River basin and probably originated from that watershed.
Etymology. The word schitsu’umsh (pronounced s-CHEET-sue-umsh) is Americanist phonetic notation of the spoken word used by the first peoples who inhabited portions of northern Idaho and western Montana, now recognized as the Coeur d’Alene Tribe. In modern usage it means “those who were found here” and refers to the people of this tribe. The current distribution of C. schitsuumsh overlaps with most of the historical homeland of this tribe. The common name refers to a common riparian tree in this area, Western Redcedar, Thuja plicata .
Remarks. Although originally assigned to C. confusus , a broad-scale genetic biodiversity inventory of fishes in the northern Rocky Mountains ( Young et al. 2013) indicated, based on sequence differences among two mtDNA regions, that C. schitsuumsh was not closely related to that species. In the present study, the more geographically comprehensive sample of fishes subjected to genetic analysis coupled with the morphological assessment confirmed that interpretation. In a broader context, however, these findings support previous assertions that the taxonomy of sculpins in North America is far from resolved ( Neely 2003; Kinziger et al. 2005; April et al. 2011). Paraphyly and polyphyly among a number of taxa are evident in phylogenetic trees based on COI ( Fig. 2 View FIGURE 2 ). Although partly attributable to a lack of resolution resulting from the limited number of nucleotides in this region ( Pollock et al. 2002), the results are similar to those observed for other mtDNA regions that included additional taxa and more nucleotides ( Kinziger et al. 2005; Young et al. 2013). In addition, the large genetic differences between clades within other species of Cottus in the western U.S., such as C. beldingii and C. confusus , suggest that additional taxa remain to be described (also see April et al. 2011).
The ambiguous morphological distinctions among sculpin species ( Jenkins & Burkhead 1994; McPhail 2007) argue for the use of genetic information for species discovery and delineation ( Baldwin & Weigt 2012). Moreover, some of the variation in diagnostic traits within species ascribed to geography may instead reflect the existence of cryptic taxa ( Bickford et al. 2007). As has been suggested for other groups of species, an integrative taxonomy relying on morphology and genes will be necessary to understand the details of the systematic relations and boundaries of individual taxa among members of the genus Cottus .
Comparative material for morphological and genetic analyses (GenBank accession numbers for COI haplotypes). Cottus beldingii : (4, 45– 54 mm SL, JX311884 View Materials ) North Fork Asotin Creek, Asotin County, WA (46.25856°N, 117.29962°W), 9 August 2011. Cottus cognatus : (5, 65– 81 mm SL, JX311885 View Materials ) Marten Creek, Sanders County, MT (47.89233°N 115.82088°W), 20 September 2011; (5, 44– 74 mm SL, JX311886 View Materials , JX311887 View Materials ) Skalkaho Creek, Ravalli County, MT (46.18128°N, 114.07984°W), 8 January 2012. Cottus cf. bairdii (Rocky Mountain Sculpin): (5, 60– 91 mm SL, JX311882 View Materials ) Elk Creek, Missoula County, MT (46.92707°N, 113.43934°W), 23 August 2011. Cottus confusus : (5, 77– 89 mm SL, JX311888 View Materials ) Little North Fork Clearwater River, Shoshone County, ID (47.06579°N, 115.85287°W), 19 September 2011; (2, 74– 89 mm SL, pending) Pole Creek, Blaine County, ID (43.90974°N, 114.76370°W), 7 November 2012; (2, 59– 68 mm SL, pending) Salmon River, Blaine County, ID (43.88382°N, 114.76370°W), 7 November 2012; (2, 67– 90 mm SL, pending) Smiley Creek, Blaine County, ID (43.89650°N, 114.79853°W), 7 November 2012. Cottus hubbsi : (1, 112 mm SL, JX311892 View Materials ) Dragoon Creek, Spokane County, WA (47.88619°N 117.38400°W), 16 August 2011. Cottus rhotheus : (1, 71 mm SL, JX311890 View Materials ) Dragoon Creek, Spokane County, WA (47.88619°N 117.38400°W), 16 August 2011; (5, 59– 84 mm SL, JX311891 View Materials ) Spokane River, Spokane County, WA (47.66060°N, 117.43559°W), 12 September 2011.
Comparative material for morphological analyses. Cottus beldingii : (2, 56– 57 mm SL, CIDA 68169) Big Bear Creek, Latah County, ID (46.63927°N, 116.65417°W), 1 July 1995; (4, 57– 68 mm SL, CIDA 68443) Clear Creek, Clearwater County, ID (46.04655°N, 115.79120°W) 18 July 1996; (2, 57– 58 mm SL, CIDA 68110) East Fork Potlatch River, Clearwater County, ID (46.82873°N, 116.38967°W), 25 July 1969; (2, 52– 60 mm SL, CIDA 68369) Glade Creek, Idaho County, ID (46.22338°N, 115.52972°W), 18 July 1996; (2, 51– 58 mm SL, CIDA 68309) Lochsa River, Idaho County, ID (46.46113°N, 115.04775°W), 15 August 1969; (4, 39– 64 mm SL, CIDA 68192) Middle Fork Clearwater River, Idaho County, ID (46.14052°N, 115.69819°W), 23July 1969; (1, 46 mm SL) North Fork Asotin Creek, Asotin County, WA (44.25856°N, 117.29962°W), 9 August 2011; (4, 49– 63 mm SL, CIDA 68389) North Fork Clearwater River, Clearwater County, ID (46.81125°N, 115.20330°W), 20 August 1969; (4, 40– 64 mm SL, CIDA 81824) South Fork Clearwater River, Idaho County, ID (46.03107°N, 115.97420°W), 20 August 2003; (2, 48– 55 mm SL) Swauk Creek, Kittatas County, WA (47.17556°N, 120.72944°W), 9 June 1996; (6, 42– 75 mm SL) Tucannon River, Columbia County, WA (46.21694°N, 117.71111°W), 1 August 1993; (2, 35– 68 mm SL, CIDA 69562) Weitas Creek, Clearwater County, ID (46.53143°N, 115.42108°W), 4 August 1998. Cottus cf. bairdii (Rocky Mountain Sculpin): (5, 84– 99 mm SL) Elk Creek, Missoula County, MT (46.92707°N, 113.43934°W), 23 August 2011. Cottus cognatus : (10, 32– 81 mm SL) Blodgett Creek, Ravalli County, MT (46.28756°N, 114.16837°W), 3 August 2012; (2, 51– 54 mm SL, CIDA 70563) Burton Creek, Boundary County, ID (48.772227°N, 116.46445°W), 26 July 2001; (2, 31– 40 mm SL, CIDA 69961) Kalispell Creek, Bonner County, ID (48.57615°N, 116.95607°W), 18 July 2000; (10, 59– 88 mm SL) Overwhich Creek, Ravalli County, MT (45.66498°N, 114.28005°W), 21 August 2012; (3, 50– 56 mm SL, CIDA 68609) Pack River, Bonner County, ID (48.47588°N, 116.55390°W), 12 October 1972; (5, 48– 71 mm SL) Petty Creek, Mineral County, MT (46.91728°N, 114.45675°W), 12 October 2011; (5, 52– 82 mm SL) Petty Creek, Mineral County, MT (46.99196°N, 114.44762°W), 12 October 2011; (5, 48– 76 mm SL) Petty Creek, Mineral County, MT (46.93116°N, 114.44568°W), 12 October 2011; (2, 48– 53 mm SL) Skalkaho Creek, Ravalli County, MT (46.18128°N, 114.07984°W), 8 January 2012; (10, 66- 77 mm SL) Sleeping Child Creek, Ravalli County, MT (46.11015°N, 114.00677°W), 10 August 2012; (3, 32– 81 mm SL) Sullivan Creek, Pend Oreille County, WA (48.86333°N, 117.30500°W), 20 August 1996; (6, 42– 71 mm SL) Tonata Creek, Ferry County, WA (48.83308°N, 118.80858°W), 1 September 1996. Cottus confusus : (3, 69– 77 mm SL) Alpine Creek, Blaine County, ID (43.89740°N, 114.91070°W), 15 August 2011; (1, 47 mm SL) American River, Yakima County, WA (46.92611°N, 121.36389°W), 1 September 1993; (3, 35– 70 mm SL) Entiat River, Chelan County, WA (47.92528°N, 120.51167°W), 1 October 1996; (2, 31– 56 mm SL) Frenchman Creek, Blaine County, ID (43.87108°N, 114.77263°W), 1 August 1998; (3, 54– 64 mm SL) Lost River, Okanogan County, WA (48.65556°N, 120.50444°W), 31 August 1999; (3, 52– 77 mm SL, CIDA 69441) Pole Creek, Blaine County, ID (43.91063°N, 114.74935°W), 26 August 1998; (3, 47– 70 mm SL, CIDA 69831) Salmon River, Blaine County, ID (43.83368°N, 114.75885°W), 26 July 1990; (5, 32– 60 mm SL, CIDA 70993) Salmon River, Blaine County, ID (43.85745°N, 114.75448°W), 3 August 2003; (3, 54– 74 mm SL) Salmon River, Blaine County, ID (43.88382°N, 114.76370°W), 7 November 2012; (7, 33– 78 mm SL, CIDA 69443) Smiley Creek, Blaine County, ID (43.82638°N, 114.82743°W), 26 July 1998; (3, 33– 78 mm SL) Smiley Creek, Blaine County, ID (43.89650°N, 114.79853°W), 7 November 2012; (2, 50 mm SL, CIDA 69503) Vat Creek, Blaine County, ID (43.97637°N, 114.84807°W), 8 August 1998. Cottus hubbsi : (4, 57– 68 mm SL) Chewuch River, Okanogan County, WA (48.56472°N, 120.17667°W), 31 August 1999; (2, 44– 53 mm SL, CIDA 68934) Siegel Creek, Idaho County, ID (45.78730°N, 115.36688°W), 11 July 1997; (5, 35– 73 mm SL) Similkameen River, Okanogan County, WA (48.95667°N, 119.64583°W), 21 July 1998; (3, 53– 63 mm SL, CIDA 69699) Skull Creek, Clearwater County, ID (46.87165°N, 115.38710°W), 9 September 1999; (3, 67– 103 mm SL, CIDA 68494) Selway River, Idaho County, ID (45.73492°N, 114.75885°W), 12 July 1970; (1, 68 mm SL) West Fork Sanpoil River, Okanogan County, WA (48.51417°N, 118.88889°W), 23 August 1999. Cottus rhotheus : (5, 67– 79 mm SL) Ramskull Creek, Benewah County, ID (47.08918°N, 116.63813°W), 11 August 2011; (5, 63– 94 mm SL) Spokane River, Spokane County, WA (47.66060°N, 117.43559°W), 12 September 2011; (10, 41– 90 mm SL, JX311891 View Materials ) Spokane River, Spokane County, WA (47.69878°N, 117.04451°W), 12 September 2011.
Comparative material for genetic analyses from GenBank. Cottus aleuticus : (5) EU523991 View Materials , EU523992 View Materials , HQ010070 View Materials , JN024987 View Materials , JN024988 View Materials . Cottus asper : (2) EU523994 View Materials , JQ354065 View Materials . Cottus aturi : (1) EF416966 View Materials . Cottus bairdii : (20) EF416967 View Materials , EU522459 View Materials , EU523998 View Materials , EU524490 View Materials , EU524496 View Materials , EU524506 View Materials , HQ557187 View Materials , HQ557189 View Materials , JN024992 View Materials , JN024996 View Materials , JN024997 View Materials , JN024999 View Materials , JN025004 View Materials , JN025006 View Materials , JN025010 View Materials , JN025011 View Materials , JN025012 View Materials , JN025013 View Materials , JN025022 View Materials , JN025023 View Materials . Cottus beldingii : (5) HQ557196 View Materials , HQ579024 View Materials , JN025027 View Materials , JN025029 View Materials , JN025034 View Materials . Cottus caeruleomentum : (2) JN025040 View Materials , JN025041 View Materials . Cottus carolinae : (8) JN025046 View Materials , JN025047 View Materials , JN025051 View Materials , JN025052 View Materials , JN025053 View Materials , JN025058 View Materials , JN025060 View Materials , JN025064 View Materials . Cottus cf. bairdii : JN025069 View Materials , JN025070 View Materials , JN025071 View Materials . Cottus cf. carolinae : (2) JN025072 View Materials , JN025073 View Materials . Cottus chattahoochee : (1) JN025077 View Materials . Cottus cognatus : (7) EU523999 View Materials , EU524511 View Materials , EU524517 View Materials , JN025085 View Materials , JN025090 View Materials , JN025091 View Materials , JN025099 View Materials . Cottus duranii : (1) EF416968 View Materials . Cottus girardi : (1) HQ557143 View Materials . Cottus gobio : (6) EF416970 View Materials , EF416971 View Materials , EF416973 View Materials , HQ960512 View Materials , HQ960869 View Materials , HQ960870 View Materials . Cottus gulosus : (2) JN025102 View Materials , JN025103 View Materials . Cottus hispaniolensis : (1) EF416977 View Materials . Cottus hubbsi : (1) JN025104 View Materials . Cottus hypselurus : (3) JN025105 View Materials , JN025106 View Materials , JN025110 View Materials . Cottus klamathensis : (1) JN025112 View Materials . Cottus leiopomus : (1) HQ971431 View Materials . Cottus perifretum : (1) EF416978 View Materials . Cottus perplexus : (3) EF416984 View Materials , JN025117 View Materials , JN025120 View Materials . Cottus pitensis : (1) JN025122 View Materials . Cottus poecilopus : (5) HQ536339 View Materials , HQ960874 View Materials , HQ960878 View Materials , HQ961093 View Materials , JN025128 View Materials . Cottus reinii : (1) AP004442 View Materials . Cottus rhenanus : (3) EF416985 View Materials , EF416986 View Materials , EF416988 View Materials . Cottus rhotheus : (3) EU524000 View Materials , HQ579026 View Materials , JN025131 View Materials . Cottus ricei : (4) EU522462 View Materials , EU524521 View Materials , HQ557327 View Materials , JN025134 View Materials . Cottus sibiricus : (1) EF416990 View Materials . Cottus tallapoosae : (6) JN025137 View Materials , JN025140 View Materials , JN025143 View Materials , JN025144 View Materials , JN025145 View Materials , JN025146 View Materials . Cottus tenuis : (2) HQ579025 View Materials , JN025147 View Materials . Leptocottus armatus : (1) FJ164712 View Materials .
Comparative material for genetic analyses from regional sampling (with GenBank accession numbers, if available). Cottus beldingii : (5, JX282576 View Materials – JX282580 View Materials , 2 pending) Feather Creek, Clearwater County, ID (46.91599°N, 116.40383°W), Potlatch River, Clearwater County, ID (46.91376°N, 116.39121°W); Goddard Creek, Idaho County, ID (46.10088°N, 115.55723°W); Musselshell Creek, Clearwater County, ID (46.35239°N, 115.75575°W); Lolo Creek, Clearwater County, ID (46.40720°N, 115.67415°W); Portneuf River, Bannock County, ID (42.70837°N, 112.21222°W); Pass Creek, Custer County, ID (43.93533°N, 113.44703°W). Cottus cf. bairdii : (5, JX282572 View Materials – JX282575 View Materials , JX311883 View Materials , 2 pending) Alice Creek, Lewis and Clark County, MT (47.10702°N, 112.4767°W); Colts Creek, Flathead County, MT (48.99259°N, 114.49092°W); Trout Creek, Lewis and Clark County, MT (46.90852°N, 112.38711°W); Taylor Fork Gallatin River, Gallatin County, MT (45.04594°N, 111.37768°W); Highwood Creek, Choteau County, MT (47.47503°N, 110.61261°W); Thayer Creek, Beaverhead County, MT (45.20349°N, 113.29354°W); Mussigbrod Creek, Beaverhead County, MT (45.76858°N, 113.58538°W). Cottus cognatus : (8, JX282581 View Materials – JX282588 View Materials ) Beaver Creek, Missoula County, MT (47.38776°N, 113.66057°W); Little Thompson River, Sanders County, MT (47.69347°N, 114.81318°W); Lake Creek, Lincoln County, MT (48.03698°N, 115.49420°W); Kraft Creek, Missoula County, MT (47.45570°N, 113.75697°W); Twelvemile, Mineral County, MT (47.37603°N, 115.25982°W); Lower Twin Creek, Flathead County, MT (47.99185°N, 113.56328°W); Overwhich Creek, Ravalli County, MT (45.66399°N, 114.15168°W). Cottus confusus : (7, JX282589 View Materials – JX282594 View Materials , JX311889 View Materials , 9 pending) Crooked Fork Creek, Clearwater County, ID (46.66158°N, 114.67718°W); Fishing Creek, Clearwater County, ID (46.51996°N, 114.86661°W); Cayuse Creek, Clearwater County, ID (46.59984°N, 114.89314°W); French Creek, Clearwater County, ID (46.52659°N, 115.65508°W); South Fork Red River, Idaho County, ID (45.67204°N, 115.36070°W); Newsome Creek, Idaho County, ID (45.91054°N, 115.62920°W); Baldy Creek, Idaho County, ID (45.91278°N, 115.64021°W); Fallert Springs, Butte County, ID (43.91625°N, 112.12339°W); Wildhorse Creek, Custer County, ID (43.85320°N, 114.08758°W); Pass Creek, Clark County, ID (44.13444°N, 112.99285°W); Big Gulch, Custer County, ID (44.34647°N, 113.52006°W); Eighteenmile Creek, Lemhi County, ID (44.53546°N, 113.24102°W); Loon Creek, Custer County, ID (44.80694°N, 114.81070°W); Salmon River, Blaine County, ID (43.88382°N, 114.76370°W); Pole Creek, Blaine County, ID (43.90974°N, 114.75629°W); Smiley Creek, Blaine County, ID (43.89650°N, 114.79853°W). Cottus hubbsi : (1, pending) South Fork Rock Creek, Power County, ID (42.50190°N, 112.82991°W). Cottus rhotheus : (2, JX282595 View Materials – JX282596 View Materials ) East Fork Emerald Creek, Latah County, ID (47.02486°N, 116.32903°W); Little Sand Creek, Latah County, ID (46.92103°N, 116.62829°W).
TABLE 1. Diagnostic characters for Cottus schitsuumsh in the mtDNA cytochrome c oxidase subunit 1 (COI) region.
Position1 | Nucleotide | Diagnostic for2 |
---|---|---|
5577 | T | C. schitsuumsh haplotype 3 |
5664 | G | C. schitsuumsh in North America |
5685 | T | C. schitsuumsh haplotype 2 |
5868 | T | C. schitsuumsh |
6081 | T | C. schitsuumsh |
6120 | T | C. schitsuumsh |
TABLE 2. Range (mode) for counts of Cottus fin spines and rays examined in this study. Samples sizes are noted after each sampling location or species.
Species | Dorsal spines | Dorsal rays | Anal rays | Pelvic rays | Pectoral rays |
---|---|---|---|---|---|
Spokane River basin (n = 64) | 7–9 (8) | 17–19 (18) | 13–15 (14) | 3–4 (4) | 12–14 (14) |
Clark Fork River basin (n = 44) | 7–9 (8) | 17–19 (18) | 12–14 (14) | 3–4 (4) | 12–15 (14) |
C. beldingii (n = 39) | 7–8 (7) | 16–18 (17) | 11–13 (12) | 3–4 (4) | 13–15 (14) |
C. cf. bairdii (n = 10) | 6–8 (8) | 16–19 (17) | 12–14 (13) | 4 | 13–15 (14) |
C. cognatus (n = 73) | 7–9 (8) | 15–19 (18) | 10–13 (12) | 3–4 (4)1 | 12–14 (13) |
C. confusus (n = 49) | 7–9 (8) | 15–19 (17) | 10–14 (13) | 4 | 11–15 (13) |
C. hubbsi (n = 19) | 7–8 (8) | 15–18 (17) | 12–14 (13) | 4 | 13–16 (15) |
C. rhotheus (n = 26) | 7–9 (8) | 15–18 (16) | 11–14 (12) | 4 | 15–17 (16) |
TABLE 3. Range (mode) of pore counts for Cottus examined in this study.
Species Medial chin pores | Preoperculomandibular pores | Postmaxillary | pores |
---|---|---|---|
Spokane River basin (n = 64) (2) | 10–12 (12) | 0–2 (2) | |
Clark Fork River basin (n = 44) (2) | 10–12 (12) | 0–2 (2) | |
C. beldingii View in CoL (n = 39) (2) | 10–12 (11) | (2) | |
C. cf. bairdii View in CoL (n = 10) (2) | 10–11 (11) | 0–2 (1)1 | |
C. cognatus View in CoL (n = 73) (2) | 10–12 (10) | 0–1 (0) | |
C. confusus View in CoL (n = 49) (2) | 10–12 (11) | 1–2 (2) | |
C. hubbsi View in CoL (n = 19) (2) | 10–12 (11) | 1–2 (2) | |
C. rhotheus View in CoL (n = 26) (2) | 10–12 (11) | 1–2 (2) | |
1Postmaxillary pores for C. cf. bairdii View in CoL were consistently in a | lower position compared to that of other | species. |
TABLE 4. Additional meristic and morphological characters of Cottus examined in this study.
Species | Preopercular spines | Prickling | Palatine teeth | Lateral line pores on caudal peduncle |
---|---|---|---|---|
Spokane River basin (n = 64) | 1 | Consistent and restricted to a trapezoidal area, widest in the axillary region and narrowest along the lateral line | 1–2 rows on 1–2 sides | No (59), yes (5) |
Clark Fork River basin (n = 44) | 1 | Consistent and restricted to a trapezoidal area, widest in the axillary region and narrowest along the lateral line | 1–2 rows on 1–2 sides | No (40), yes (4) |
C. beldingii View in CoL (n = 39) | 1–2 (1) | Absent | Absent | No |
C. cf. bairdii View in CoL (n = 10) | 1–2 (2) | Generally absent; some prickles (1–7) can be present axial to the pectoral fin | ≥ 2 rows on both sides | No |
C. cognatus View in CoL (n = 73) | 2–3 (2) | Generally absent; occasionally small patches in the axillary region | Absent | No |
C. confusus View in CoL (n = 49) | 2–3 (2) | Consistent and restricted to a trapezoidal area, widest in the axillary region and narrowest along the lateral line | 1–2 rows on 1–2 sides | No (3), yes (46) |
C. hubbsi View in CoL (n = 19) | 3 | Large patch under pectoral fins, scattered elsewhere | ≥ 2 rows on both sides | Yes |
C. rhotheus View in CoL (n = 26) | 3–4 (3) | Covers most of the body | ≥ 2 rows on both sides | Yes |
TABLE 5. Mean values for 11 morphometric characteristics among Cottus examined in this study. Morphometric values were standardized by dividing by standard length, except mouth width was standardized by head width. An asterisk denotes a significant pairwise difference (ANCOVA, Scheffé test; P <0.05) between the pooled sample of C. schitsuumsh and another species.
Species | Caudal peduncle depth | Caudal peduncle length | Head length | Head width | Lateral line length | Snout length |
---|---|---|---|---|---|---|
Spokane River basin (n = 64) | 0.086 | 0.144 | 0.191 | 0.264 | 0.582 | 0.086 |
Clark Fork River basin (n = 44) | 0.088 | 0.144 | 0.187 | 0.265 | 0.584 | 0.086 |
C. beldingii View in CoL (n = 39) | 0.097* | 0.134* | 0.211* | 0.273 | 0.620* | 0.092 |
C. cf. bairdii View in CoL (n = 10) | 0.077* | 0.131 | 0.208* | 0.301* | 0.599 | 0.084 |
C. cognatus View in CoL (n = 73) | 0.078* | 0.165* | 0.221* | 0.257 | 0.565 | 0.076* |
C. confusus View in CoL (n = 49) | 0.084 | 0.157 | 0.202 | 0.265 | 0.667* | 0.078* |
C. hubbsi View in CoL (n = 19) | 0.081* | 0.131 | 0.245* | 0.299* | 0.711* | 0.095 |
C. rhotheus View in CoL (n = 26) | 0.062* | 0.134 | 0.213* | 0.304* | 0.678* | 0.109* |
continued. |
TABLE 6. Frequency distribution of head length divided by standard length (%) for Cottus examined in this study.
Species Spokane River basin (n = 64) | Mouth width 0.608 | Interpelvic width 0.034 | Longest pectoral ray 0.338 | Longest pelvic ray 0.180 | Longest dorsal ray 0.124 |
---|---|---|---|---|---|
Clark Fork River basin (n = 44) | 0.605 | 0.036 | 0.337 | 0.179 | 0.123 |
C. beldingii View in CoL (n = 39) | 0.509* | 0.042* | 0.282* | 0.167* | 0.116 |
C. cf. bairdii View in CoL (n = 10) | 0.522* | 0.043* | 0.325 | 0.191 | 0.106 |
C. cognatus View in CoL (n = 73) | 0.505* | 0.044* | 0.287* | 0.198* | 0.110* |
C. confusus View in CoL (n = 49) | 0.527* | 0.039* | 0.314 | 0.178 | 0.117 |
C. hubbsi View in CoL (n = 19) | 0.541* | 0.045* | 0.302 | 0.217* | 0.126 |
C. rhotheus View in CoL (n = 26) | 0.599 | 0.038 | 0.284* | 0.185 | 0.129 |
TABLE 7. Frequency distribution of interpelvic width divided by standard length (%) for Cottus examined in this study.
Standardized interpelvic width | ||||||||
---|---|---|---|---|---|---|---|---|
Species | 1.5 2.0 2.5 3.0 3.5 | 4.0 | 4.5 | 5.0 | 5.5 | 6.0 | 6.5 | 7.0 |
Spokane River basin (n = 64) | 2 1 5 18 16 | 19 | 2 | 1 | ||||
Clark Fork River basin (n = 44) | 2 1 10 15 | 12 | 2 | 2 | ||||
C. beldingii (n = 39) | 2 7 | 12 | 11 | 4 | 1 | 2 | ||
C. cf. bairdii (n = 10) | 4 | 4 | 2 | |||||
C. cognatus (n = 73) | 1 2 7 | 19 | 19 | 15 | 7 | 2 | 1 | |
C. confusus (n = 49) | 5 15 | 16 | 11 | 1 | 1 | |||
C. hubbsi (n = 19) | 4 | 3 | 6 | 2 | 2 | 1 | 1 | |
C. rhotheus (n = 26) | 4 2 6 | 8 | 5 | 1 |
No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.
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