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<mods:title id="8306E39F205E1CA5ACAB59C6E1C0725E">Cryptic Species of Freshwater Sculpin (Cottidae: Cottus) in California, USA</mods:title>
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<mods:namePart id="902B0465A2BDB32BEA49983EC5447D95">Moyle, Peter B.</mods:namePart>
<mods:affiliation id="E4F43D8CB6918732C4ECA5F2ACCDEBEB">Center for Watershed Sciences, 425 La Rue Road, University of California, Davis, California 95616</mods:affiliation>
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<mods:namePart id="936D086A99E9FB71271BFBFFE3D7E3F8">Campbell, Matthew A.</mods:namePart>
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<mods:affiliation id="9B1D0EE104DB9841C58D24B2BCF2C72D">Genomic Variation Laboratory, 2403 Meyer Hall, Department of Animal Science, University of California, Davis, California 95616. https: // orcid. org / 0000 - 0002 - 5826 - 0329</mods:affiliation>
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<treatment id="C14A87D6FFCCFFF82792FE1E518DF80F" ID-DOI="http://doi.org/10.5281/zenodo.6655463" ID-GBIF-Taxon="196079302" ID-Zenodo-Dep="6655463" LSID="urn:lsid:plazi:treatment:C14A87D6FFCCFFF82792FE1E518DF80F" httpUri="http://treatment.plazi.org/id/C14A87D6FFCCFFF82792FE1E518DF80F" lastPageNumber="516" pageId="15" pageNumber="516">
<subSubSection id="01F9654BFFCCFFF82792FE1E537FFDF7" box="[151,788,499,526]" pageId="15" pageNumber="516" type="nomenclature">
<paragraph id="495C36C0FFCCFFF82792FE1E537FFDF7" blockId="15.[151,788,499,562]" box="[151,788,499,526]" pageId="15" pageNumber="516">
<heading id="121481ACFFCCFFF82792FE1E537FFDF7" bold="true" box="[151,788,499,526]" fontSize="11" level="1" pageId="15" pageNumber="516" reason="1">
<emphasis id="7B97EAD2FFCCFFF82792FE1E537FFDF7" bold="true" box="[151,788,499,526]" pageId="15" pageNumber="516">
<taxonomicName id="8EE34D43FFCCFFF82792FE1E526BFDF7" ID-CoL="YYH6" authority="(Girard 1854)" baseAuthorityName="Girard" baseAuthorityYear="1854" box="[151,512,499,526]" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="gulosus">
<emphasis id="7B97EAD2FFCCFFF82792FE1E5121FDF7" bold="true" box="[151,330,499,526]" italics="true" pageId="15" pageNumber="516">Cottus gulosus</emphasis>
(
<bibRefCitation id="2D724B31FFCCFFF8265EFE1E519CFDF7" author="Girard, C." box="[347,503,500,526]" pageId="15" pageNumber="516" pagination="129 - 165" refId="ref15987" refString="Girard, C. (1854) Descriptions of new fishes collected by Dr. A. L. Heermann, naturalist attached to the survey of the Pacific railroad route, under Lieut. R. S. Williamson, U. S. A. Proceedings of the Academy of Natural Sciences of Philadelphia, 7, 129 - 165." type="journal article" year="1854">Girard 1854</bibRefCitation>
)
</taxonomicName>
, Inland Riffle Sculpin
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="01F9654BFFCCFFF82792FDF25307FAFA" pageId="15" pageNumber="516" type="description">
<paragraph id="495C36C0FFCCFFF82792FDF25080FDCB" blockId="15.[151,788,499,562]" box="[151,235,536,562]" pageId="15" pageNumber="516">
(
<figureCitation id="D1D82A45FFCCFFF827A5FDF2508BFDCB" box="[160,224,536,562]" captionStart="FIGURE 1" captionStartId="2.[151,250,1758,1782]" captionTargetBox="[238,1348,181,1734]" captionTargetId="figure-24@2.[238,1348,181,1734]" captionTargetPageId="2" captionText="FIGURE 1. Inland (A, B) and Coastal (C, D) Riffle Sculpins from the UC Davis Museum of Wildlife and Fish Biology. (A) San Joaquin Riffle Sculpin, Cottus gulosus gulosus. WFB-277-08-09. Kaweah River, Tulare County, California (68 mm SL, 82 mm TL). Coll. Larry Brown, September 10, 1985. Holotype is same as for C. gulosus (USNM 291). (B) Sacramento Riffle Sculpin, Cottus gulosus wintu. Holotype WFB-3464. North Fork Feather River, Butte County, California (78 mm SL, 96 mm TL). Coll. Jason Baumsteiger, October 5, 2017. (C) Coastal Riffle Sculpin, Cottus ohlone. Holotype. WFB-3402.Also, holotype for Ohlone Sculpin, C. o. ohlone. Guadalupe River, Santa Clara County, California (65 mm SL, 79 mm TL). Coll: J. J. Smith, November 18, 1986. (D) Pomo Riffle Sculpin, Cottus ohlone pomo. Holotype. WFB-3396. Pieta Creek near mouth on Russian River, Mendocino County, California (90 mm SL, 105 mm TL). Coll. P Moyle, J. Baumsteiger, August 2, 2017." figureDoi="http://doi.org/10.5281/zenodo.6655475" httpUri="https://zenodo.org/record/6655475/files/figure.png" pageId="15" pageNumber="516">Fig. 1</figureCitation>
)
</paragraph>
<paragraph id="495C36C0FFCCFFF82792FD8A5394FBB7" blockId="15.[151,1437,608,2039]" pageId="15" pageNumber="516">
Originally described by Charles Girard as
<taxonomicName id="8EE34D43FFCCFFF82569FD8A5359FD83" authorityName="Girard" authorityYear="1854" box="[620,818,608,634]" class="Actinopterygii" family="Cottidae" genus="Cottopsis" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="gulosus">
<emphasis id="7B97EAD2FFCCFFF82569FD8A5359FD83" box="[620,818,608,634]" italics="true" pageId="15" pageNumber="516">Cottopsis gulosus</emphasis>
</taxonomicName>
from the San Joaquin River, as follows: “Largest specimens a little over three inches {
<quantity id="8E1B9B25FFCCFFF826FEFD6E5221FD66" box="[507,586,644,671]" metricMagnitude="-1" metricUnit="m" metricValue="7.6" pageId="15" pageNumber="516" unit="cm" value="76.0">76 cm</quantity>
} in total length; of which the head forms the third, the caudal fin excepted. Preopercle provided with two small spines, such as may be observed in several species of
<taxonomicName id="8EE34D43FFCCFFF82390FD4254B5FD3B" box="[1173,1246,680,706]" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="genus">
<emphasis id="7B97EAD2FFCCFFF82390FD4254B5FD3B" box="[1173,1246,680,706]" italics="true" pageId="15" pageNumber="516">Cottus</emphasis>
</taxonomicName>
proper, the head otherwise is smooth; mouth proportionately large; posterior extremity of upper maxillary reaching a vertical line, which would pass behind the pupil. A space of five twentieths of an inch exists between the origin of the first dorsal and the occiput. Second dorsal connected with the first by a low membrane proceeding from the last spiny ray. Tips of posterior soft rays not quite reaching the base of caudal. Anal, about as high as second dorsal, but shorter. Caudal, well developed and posteriorly rounded. Origin of ventrals midway between the tip of lower jaw and the anus. Pectorals broad and large, its longest rays extending posteriorly as far as the fifth ray of second dorsal. D IX. 18. A 13. C 3. 1. 5. 4. 1. 2. V I. 4. P 15. Lateral line uninterrupted from thoracic region to base of caudal. Abdomen beset with minute prickles; skin elsewhere smooth. Ground color reddish brown; head and dorsal region spotted with black. Dorsal, caudal and pectorals barred with black; first dorsal provided posteriorly with an elongated black spot. Anal and ventrals unicolor. (
<bibRefCitation id="2D724B31FFCCFFF8250EFBFA52FCFBD2" author="Girard, C." box="[523,663,1040,1067]" pageId="15" pageNumber="516" pagination="129 - 165" refId="ref15987" refString="Girard, C. (1854) Descriptions of new fishes collected by Dr. A. L. Heermann, naturalist attached to the survey of the Pacific railroad route, under Lieut. R. S. Williamson, U. S. A. Proceedings of the Academy of Natural Sciences of Philadelphia, 7, 129 - 165." type="journal article" year="1854">Girard 1854</bibRefCitation>
; p.130). This description still fits the species complex today, although larger samples show more variation in fin ray counts and characters (
<tableCitation id="0461037BFFCCFFF8249FFBDE5385FBB7" box="[922,1006,1076,1102]" captionStart="TABLE 1" captionStartId="4.[150,238,542,568]" captionTargetBox="[165,1966,655,1605]" captionTargetPageId="4" captionText="TABLE 1. Meristic values for Riffle Sculpin samples from the UC Davis Museum of Wildlife and Fish and the California Academy of Sciences, San Francisco. Cottus asper and C.pitensis data is presented for comparison.? means could not be determined. Locations include the year collected. See Table 2 for more precise location information. Sculpins at locations with * were C. asper originally identified as C. gulosus" httpUri="http://table.plazi.org/id/1D9C6648FFC7FFF32793FDF4530EFD79" pageId="15" pageNumber="516" tableUuid="1D9C6648FFC7FFF32793FDF4530EFD79">Table 1</tableCitation>
).
</paragraph>
<paragraph id="495C36C0FFCCFFF827C2FBB25307FAFA" blockId="15.[151,1437,608,2039]" pageId="15" pageNumber="516">Characteristics of Riffle Sculpins used in this study (n = 23) are as follows: anal-fin rays (1215), dorsal spines (610), dorsal rays (1419), dorsal fins usually joined, black spot on dorsal (yes), palatine teeth present (variable), mouth vs eye (variable but maxilla mostly reaches mid-eye), chin pores (02), pelvic fins spines + rays (1+23), pelvics touch vent when depressed (variable but mostly not), pectoral fin rays (1316), preopercular spines (12), lateral line completeness variable, and lateral line pores (2638).</paragraph>
</subSubSection>
<subSubSection id="01F9654BFFCCFFF827C2FAE1529EFAB2" pageId="15" pageNumber="516" type="materials_examined">
<paragraph id="495C36C0FFCCFFF827C2FAE1529EFAB2" blockId="15.[151,1437,608,2039]" pageId="15" pageNumber="516">
<materialsCitation id="F98B3C9DFFCCFFF827C2FAE1529EFAB2" ID-GBIF-Occurrence="3815677303" collectionCode="USNM" collectorName="A. L. Heermann &amp; Charles" location="San Joaquin River" pageId="15" pageNumber="516" specimenCode="USNM 291" specimenCount="1" stateProvince="California" typeStatus="holotype">
<emphasis id="7B97EAD2FFCCFFF827C2FAE1515DFADF" bold="true" box="[199,310,1291,1318]" italics="true" pageId="15" pageNumber="516">
<typeStatus id="96588862FFCCFFF827C2FAE15159FADF" box="[199,306,1291,1318]" pageId="15" pageNumber="516" type="holotype">Holotype</typeStatus>
.
</emphasis>
<specimenCode id="19459EBBFFCCFFF82638FAE651AAFADF" box="[317,449,1292,1318]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="15" pageNumber="516" type="Museum">USNM 291</specimenCode>
,
<taxonomicName id="8EE34D43FFCCFFF826C9FAE652F8FADF" authorityName="Girard" authorityYear="1854" box="[460,659,1292,1318]" class="Actinopterygii" family="Cottidae" genus="Cottopsis" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="gulosus">
<emphasis id="7B97EAD2FFCCFFF826C9FAE652F8FADF" box="[460,659,1292,1318]" italics="true" pageId="15" pageNumber="516">Cottopsis gulosus</emphasis>
</taxonomicName>
from
<location id="4C3C601BFFCCFFF825D3FAE653CEFADF" LSID="urn:lsid:plazi:treatment:C14A87D6FFCCFFF82792FE1E518DF80F:4C3C601BFFCCFFF825D3FAE653CEFADF" box="[726,933,1292,1318]" name="San Joaquin River" pageId="15" pageNumber="516" stateProvince="California">San Joaquin River</location>
,
<collectingRegion id="8B27F822FFCCFFF824B5FAE65449FADF" box="[944,1058,1292,1318]" country="United States of America" name="California" pageId="15" pageNumber="516">California</collectingRegion>
(
<specimenCount id="5FE5FD49FFCCFFF82335FAE654B0FADE" box="[1072,1243,1292,1319]" pageId="15" pageNumber="516" type="generic">two specimens</specimenCount>
), Collected by
<collectorName id="E4165316FFCCFFF82284FAE65140FAB2" pageId="15" pageNumber="516">A. L. Heermann</collectorName>
and described by
<collectorName id="E4165316FFCCFFF826F2FADA5226FAB3" box="[503,589,1328,1354]" pageId="15" pageNumber="516">Charles</collectorName>
<bibRefCitation id="2D724B31FFCCFFF82556FADA529AFAB2" author="Girard, C." box="[595,753,1328,1355]" pageId="15" pageNumber="516" pagination="129 - 165" refId="ref15987" refString="Girard, C. (1854) Descriptions of new fishes collected by Dr. A. L. Heermann, naturalist attached to the survey of the Pacific railroad route, under Lieut. R. S. Williamson, U. S. A. Proceedings of the Academy of Natural Sciences of Philadelphia, 7, 129 - 165." type="journal article" year="1854">Girard (1854)</bibRefCitation>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="01F9654BFFCCFFF827C2FABE52FBFA23" pageId="15" pageNumber="516" type="diagnosis">
<paragraph id="495C36C0FFCCFFF827C2FABE52FBFA23" blockId="15.[151,1437,608,2039]" pageId="15" pageNumber="516">
<emphasis id="7B97EAD2FFCCFFF827C2FABE5151FA97" bold="true" box="[199,314,1364,1390]" italics="true" pageId="15" pageNumber="516">Diagnosis</emphasis>
. It is distinguished from other members of the
<taxonomicName id="8EE34D43FFCCFFF8243BFABE53D8FA97" authorityName="Girard" authorityYear="1854" box="[830,947,1364,1390]" class="Actinopterygii" family="Cottidae" genus="Cottopsis" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="gulosus">
<emphasis id="7B97EAD2FFCCFFF8243BFABE53D8FA97" box="[830,947,1364,1390]" italics="true" pageId="15" pageNumber="516">C. gulosus</emphasis>
</taxonomicName>
complex as a distinct lineage determined by genomic studies and its limited geographic distribution (
<figureCitation id="D1D82A45FFCCFFF82417FA92530BFA6B" box="[786,864,1400,1426]" captionStart="FIGURE 1" captionStartId="2.[151,250,1758,1782]" captionTargetBox="[238,1348,181,1734]" captionTargetId="figure-24@2.[238,1348,181,1734]" captionTargetPageId="2" captionText="FIGURE 1. Inland (A, B) and Coastal (C, D) Riffle Sculpins from the UC Davis Museum of Wildlife and Fish Biology. (A) San Joaquin Riffle Sculpin, Cottus gulosus gulosus. WFB-277-08-09. Kaweah River, Tulare County, California (68 mm SL, 82 mm TL). Coll. Larry Brown, September 10, 1985. Holotype is same as for C. gulosus (USNM 291). (B) Sacramento Riffle Sculpin, Cottus gulosus wintu. Holotype WFB-3464. North Fork Feather River, Butte County, California (78 mm SL, 96 mm TL). Coll. Jason Baumsteiger, October 5, 2017. (C) Coastal Riffle Sculpin, Cottus ohlone. Holotype. WFB-3402.Also, holotype for Ohlone Sculpin, C. o. ohlone. Guadalupe River, Santa Clara County, California (65 mm SL, 79 mm TL). Coll: J. J. Smith, November 18, 1986. (D) Pomo Riffle Sculpin, Cottus ohlone pomo. Holotype. WFB-3396. Pieta Creek near mouth on Russian River, Mendocino County, California (90 mm SL, 105 mm TL). Coll. P Moyle, J. Baumsteiger, August 2, 2017." figureDoi="http://doi.org/10.5281/zenodo.6655475" httpUri="https://zenodo.org/record/6655475/files/figure.png" pageId="15" pageNumber="516">Figs. 1</figureCitation>
,
<figureCitation id="D1D82A45FFCCFFF8246BFA925315FA6B" box="[878,894,1400,1426]" captionStart="FIGURE 3" captionStartId="10.[151,250,678,702]" captionTargetBox="[157,1420,181,654]" captionTargetId="figure-24@10.[156,1431,181,654]" captionTargetPageId="10" captionText="FIGURE 3. (A) Distribution of Cottus pitensis in northeastern California (light green). This distribution is mostly in streams of the Pit River basin in California but extends into Oregon tributaries of Goose Lake (upper right corner of map). (B) Drawing of C. pitensis holotype from Bailey and Bond (1963), collected from the North Fork Pit River, Modoc County, CA." figureDoi="http://doi.org/10.5281/zenodo.6655479" httpUri="https://zenodo.org/record/6655479/files/figure.png" pageId="15" pageNumber="516">3</figureCitation>
). In streams, it can co-occur with
<taxonomicName id="8EE34D43FFCCFFF823FAFA925535FA6B" box="[1279,1374,1400,1426]" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="asper">
<emphasis id="7B97EAD2FFCCFFF823FAFA925535FA6B" box="[1279,1374,1400,1426]" italics="true" pageId="15" pageNumber="516">C. asper</emphasis>
</taxonomicName>
from which it can be most easily separated by its short anal fin (1215 rays vs. 1619 for
<taxonomicName id="8EE34D43FFCCFFF82346FA7654CAFA4F" box="[1091,1185,1436,1462]" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="asper">
<emphasis id="7B97EAD2FFCCFFF82346FA7654CAFA4F" box="[1091,1185,1436,1462]" italics="true" pageId="15" pageNumber="516">C. asper</emphasis>
</taxonomicName>
) and short pelvic fins, which do not touch the vent when depressed.
</paragraph>
</subSubSection>
<subSubSection id="01F9654BFFCCFFF827C2FA0951D6F857" pageId="15" pageNumber="516" type="distribution">
<paragraph id="495C36C0FFCCFFF827C2FA0951D6F857" blockId="15.[151,1437,608,2039]" pageId="15" pageNumber="516">
<emphasis id="7B97EAD2FFCCFFF827C2FA095138FA07" bold="true" box="[199,339,1507,1534]" italics="true" pageId="15" pageNumber="516">Distribution</emphasis>
.
<taxonomicName id="8EE34D43FFCCFFF82665FA0E51BDFA07" authorityName="Girard" authorityYear="1854" box="[352,470,1508,1534]" class="Actinopterygii" family="Cottidae" genus="Cottopsis" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="gulosus">
<emphasis id="7B97EAD2FFCCFFF82665FA0E51BDFA07" box="[352,470,1508,1534]" italics="true" pageId="15" pageNumber="516">C. gulosus</emphasis>
</taxonomicName>
is the name historically assigned to all populations in the species complex as described in this paper, including
<taxonomicName id="8EE34D43FFCCFFF826ADF9E2524AF9DB" authorityName=", Pit Sculpin Bailey and Bond" authorityYear="1963" box="[424,545,1544,1570]" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="pitensis">
<emphasis id="7B97EAD2FFCCFFF826ADF9E2524AF9DB" box="[424,545,1544,1570]" italics="true" pageId="15" pageNumber="516">C. pitensis</emphasis>
</taxonomicName>
. Our study shows that the name should be restricted to (a) populations found in streams in the western Sierra
<collectingRegion id="8B27F822FFCCFFF82501F9C65237F9BF" box="[516,604,1580,1606]" country="United States of America" name="Nevada" pageId="15" pageNumber="516">Nevada</collectingRegion>
, on the east side of the southern Central Valley, from the American River watershed in the north to the Kern River watershed in the south (
<figureCitation id="D1D82A45FFCCFFF8247BF9BA53AFF993" box="[894,964,1616,1642]" captionStart="FIGURE 2" captionStartId="9.[151,250,1810,1834]" captionTargetBox="[174,1414,181,1786]" captionTargetId="figure-17@9.[174,1414,181,1786]" captionTargetPageId="9" captionText="FIGURE 2. Distribution of Riffle Sculpin taxa in California, described in the text. Pit Sculpin (Cottus pitensis) is found in the northeast (light green). Inland Riffle Sculpin (Cottus gulosus) is found mainly in streams of the Central Valley (blue/orange), while Coastal Riffle Sculpin (Cottus ohlone) is confined to coastal and South San Francisco Bay watersheds (yellow/pink)." figureDoi="http://doi.org/10.5281/zenodo.6655477" httpUri="https://zenodo.org/record/6655477/files/figure.png" pageId="15" pageNumber="516">Fig. 2</figureCitation>
), including the San Joaquin River and its tributaries and (b) populations in the Sacramento River and streams tributary to it, except the Pit River.
<bibRefCitation id="2D724B31FFCCFFF8222CF99E5139F94B" author="Baumsteiger, J. &amp; Kinziger, A. P. &amp; Reid., S. B. &amp; Aguilar, A." pageId="15" pageNumber="516" pagination="2602 - 2618" refId="ref15005" refString="Baumsteiger, J., Kinziger, A. P., Reid., S. B. &amp; Aguilar, A. (2014) Complex phylogeography and historical hybridization between sister taxa of freshwater sculpin (Cottus). Molecular Ecology, 23, 2602 - 2618. https: // doi. org / 10.1111 / mec. 12758" type="journal article" year="2014">
Baumsteiger
<emphasis id="7B97EAD2FFCCFFF827C1F973509CF94B" box="[196,247,1688,1714]" italics="true" pageId="15" pageNumber="516">et al</emphasis>
. (2014)
</bibRefCitation>
used mitochondrial and nuclear DNA to examine a large sample (n = 872) of sculpins assigned to the
<taxonomicName id="8EE34D43FFCCFFF827DAF956513DF92F" authorityName="Girard" authorityYear="1854" box="[223,342,1724,1750]" class="Actinopterygii" family="Cottidae" genus="Cottopsis" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="gulosus">
<emphasis id="7B97EAD2FFCCFFF827DAF956513DF92F" box="[223,342,1724,1750]" italics="true" pageId="15" pageNumber="516">C. gulosus</emphasis>
</taxonomicName>
species complex. They found a distinct, genetically-based separation of
<taxonomicName id="8EE34D43FFCCFFF8238FF956556AF92F" authorityName="Girard" authorityYear="1854" box="[1162,1281,1724,1750]" class="Actinopterygii" family="Cottidae" genus="Cottopsis" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="gulosus">
<emphasis id="7B97EAD2FFCCFFF8238FF956556AF92F" box="[1162,1281,1724,1750]" italics="true" pageId="15" pageNumber="516">C. gulosus</emphasis>
</taxonomicName>
from streams flowing into the southern Central Valley, from similar sculpins in streams flowing into the northern Central Valley (Sacramento Valley). These populations here are treated here as new subspecies. The two river systems join in the tidal Sacramento-San Joaquin Delta. A similar break was found in the distribution of the Sacramento Riffle Sculpin lineage at the mouth of the Pit River (now under Shasta Reservoir); sculpins above this break are genetically Pit Sculpins while those below the break are a subspecies that shows evidence of ancient hybridization with
<taxonomicName id="8EE34D43FFCCFFF82222F89A51D2F857" authority="(Baumsteiger et al. 2014)" baseAuthorityName="Baumsteiger" baseAuthorityYear="2014" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="pitensis">
<emphasis id="7B97EAD2FFCCFFF82222F89A55F6F873" box="[1319,1437,1904,1930]" italics="true" pageId="15" pageNumber="516">C. pitensis</emphasis>
(
<bibRefCitation id="2D724B31FFCCFFF8279AF87E51DAF857" author="Baumsteiger, J. &amp; Kinziger, A. P. &amp; Reid., S. B. &amp; Aguilar, A." box="[159,433,1940,1967]" pageId="15" pageNumber="516" pagination="2602 - 2618" refId="ref15005" refString="Baumsteiger, J., Kinziger, A. P., Reid., S. B. &amp; Aguilar, A. (2014) Complex phylogeography and historical hybridization between sister taxa of freshwater sculpin (Cottus). Molecular Ecology, 23, 2602 - 2618. https: // doi. org / 10.1111 / mec. 12758" type="journal article" year="2014">
Baumsteiger
<emphasis id="7B97EAD2FFCCFFF82630F87F510DF857" box="[309,358,1940,1966]" italics="true" pageId="15" pageNumber="516">et al</emphasis>
. 2014
</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="01F9654BFFCCFFF827C2F85D518DF80F" pageId="15" pageNumber="516" type="etymology">
<paragraph id="495C36C0FFCCFFF827C2F85D518DF80F" blockId="15.[151,1437,608,2039]" pageId="15" pageNumber="516">
<emphasis id="7B97EAD2FFCCFFF827C2F85D51F3F82B" box="[199,408,1975,2002]" italics="true" pageId="15" pageNumber="516">
<emphasis id="7B97EAD2FFCCFFF827C2F85D5123F82B" bold="true" box="[199,328,1975,2002]" italics="true" pageId="15" pageNumber="516">Etymology.</emphasis>
<taxonomicName id="8EE34D43FFCCFFF8264AF85251F3F82B" box="[335,408,1976,2002]" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="genus">Cottus</taxonomicName>
</emphasis>
was the Roman name for European sculpin, while
<taxonomicName id="8EE34D43FFCCFFF824D4F852544CF82B" authorityName="Girard" authorityYear="1854" box="[977,1063,1976,2002]" class="Actinopterygii" family="Cottidae" genus="Cottopsis" kingdom="Animalia" order="Scorpaeniformes" pageId="15" pageNumber="516" phylum="Chordata" rank="species" species="gulosus">
<emphasis id="7B97EAD2FFCCFFF824D4F852544CF82B" box="[977,1063,1976,2002]" italics="true" pageId="15" pageNumber="516">gulosus</emphasis>
</taxonomicName>
roughly translates as big mouth or gluttonous (
<bibRefCitation id="2D724B31FFCCFFF8264EF83651B1F80F" author="Moyle, P. B." box="[331,474,2012,2038]" pageId="15" pageNumber="516" refId="ref17225" refString="Moyle, P. B. (2002) Inland Fishes of California. Revised and Expanded. University of California Press, Berkeley, 517 pp." type="book" year="2002">
<collectingRegion id="8B27F822FFCCFFF8264EF83651FDF80F" box="[331,406,2012,2038]" country="United Kingdom" name="Moyle" pageId="15" pageNumber="516">Moyle</collectingRegion>
2002
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>