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<document ID-DOI="10.11646/zootaxa.5249.5.1" ID-ISSN="1175-5326" ID-Zenodo-Dep="7701357" ID-ZooBank="F146B808-9D5B-477F-9E73-09A8DFDBFA31" approvalRequired="12" approvalRequired_for_textStreams="12" checkinTime="1678097781678" checkinUser="plazi" docAuthor="Moyle, Peter B., Buckmaster, Nicholas &amp; Su, Yingxin" docDate="2023" docId="03D1EC51DE0BFF873FFFFC0BCCC9F8F8" docLanguage="en" docName="zootaxa.5249.5.1.pdf" docOrigin="Zootaxa 5249 (5)" docSource="http://dx.doi.org/10.11646/zootaxa.5249.5.1" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Rhinichthys klamathensis subsp. goyatoka Moyle &amp; Buckmaster &amp; Su 2023, new subspecies" docType="treatment" docVersion="2" lastPageNumber="532" masterDocId="FFE89429DE15FF983F68FF81CD38FF89" masterDocTitle="Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA" masterLastPageNumber="539" masterPageNumber="501" pageNumber="531" updateTime="1678098333465" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title>Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Moyle, Peter B.</mods:namePart>
<mods:affiliation>Center for Watershed Sciences &amp; Department of Wildlife, Fish and Conservation Biology, University of California, 425 LaRue Road, Davis CA 95626 USA.</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Buckmaster, Nicholas</mods:namePart>
<mods:affiliation>California Department of Fish &amp; Wildlife, 787 Main Street, Bishop, CA 93514 USA. Nick. buckmaster @ wildlife. ca. gov Department of Animal Science, University of California, Davis, 1 Shields Avenue, Davis CA 95616 USA. syxsu @ ucdavis. edu Corresponding author. pbmoyle @ ucdavis. edu; https: // orcid. org / 0000 - 0002 - 4825 - 4865</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Su, Yingxin</mods:namePart>
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<mods:date>2023</mods:date>
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<mods:number>2023-03-06</mods:number>
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<mods:identifier type="DOI">10.11646/zootaxa.5249.5.1</mods:identifier>
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<subSubSection box="[151,1085,906,933]" pageId="30" pageNumber="531" type="nomenclature">
<paragraph blockId="30.[151,1085,906,969]" box="[151,1085,906,933]" pageId="30" pageNumber="531">
<heading bold="true" box="[151,1085,906,933]" fontSize="11" level="1" pageId="30" pageNumber="531" reason="1">
<emphasis bold="true" box="[151,1085,906,933]" pageId="30" pageNumber="531">
<taxonomicName authority="Moyle &amp; Buckmaster &amp; Su, 2023" authorityName="Moyle &amp; Buckmaster &amp; Su" authorityYear="2023" box="[151,581,906,933]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="30" pageNumber="531" phylum="Chordata" rank="subSpecies" species="klamathensis" status="new subspecies" subSpecies="goyatoka">
<emphasis bold="true" box="[151,581,906,933]" italics="true" pageId="30" pageNumber="531">Rhinichthys klamathensis goyatoka</emphasis>
</taxonomicName>
,
<taxonomicNameLabel box="[596,787,906,933]" pageId="30" pageNumber="531" rank="subSpecies">new subspecies</taxonomicNameLabel>
. Warner Speckled Dace
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[151,236,943,969]" pageId="30" pageNumber="531" type="description">
<paragraph blockId="30.[151,1085,906,969]" box="[151,236,943,969]" pageId="30" pageNumber="531">
<figureCitation box="[151,236,943,969]" captionStart="FIGURE 4" captionStartId="14.[152,255,724,748]" captionTargetBox="[149,1438,181,700]" captionTargetId="figure-21@14.[149,1438,181,700]" captionTargetPageId="14" captionText="FIGURE 4. Holotypes of new taxa of Rhinichthys described in this paper, A. Santa Ana Speckled Dace (R. gabrieleno WFB 3498. B. Long Valley Speckled Dace (R. nevadensis caldera) WFB 5000, C. Warner Speckled Dace (R. klamathensis goyatoka) WFB 122-10-44, D. Sacramento Speckled Dace (R. klamathensis achomawi) WFB 3171." figureDoi="http://doi.org/10.5281/zenodo.7701365" httpUri="https://zenodo.org/record/7701365/files/figure.png" pageId="30" pageNumber="531">Fig. 4C</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="30" pageNumber="531" type="reference_group">
<paragraph blockId="30.[151,681,1011,1158]" box="[151,582,1011,1035]" pageId="30" pageNumber="531">
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<treatmentCitation author="Snyder, J. O." box="[151,582,1011,1035]" page="98" pageId="30" pageNumber="531" year="1908">
<taxonomicName authority="Snyder 1908: 98" authorityName="Snyder" authorityPageNumber="98" authorityYear="1908" box="[151,582,1011,1035]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="30" pageNumber="531" phylum="Chordata" rank="subSpecies" species="nubila" subSpecies="carringtoni">
<emphasis box="[151,414,1011,1035]" italics="true" pageId="30" pageNumber="531">Agosia nubila carringtoni</emphasis>
<bibRefCitation author="Snyder, J. O." box="[421,582,1011,1035]" pageId="30" pageNumber="531" pagination="69 - 102" refId="ref31713" refString="Snyder, J. O. (1908) Relationships of the fish fauna of the lakes of Southeastern Oregon. U. S. Bureau of Fisheries Bulletin, 27, 69 - 102." type="journal article" year="1908">Snyder 1908:98</bibRefCitation>
</taxonomicName>
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</paragraph>
<paragraph blockId="30.[151,681,1011,1158]" box="[151,482,1042,1066]" pageId="30" pageNumber="531">
<treatmentCitationGroup box="[151,482,1042,1066]" pageId="30" pageNumber="531">
<treatmentCitation author="Rutter, C." box="[151,482,1042,1066]" page="139" pageId="30" pageNumber="531" year="1908">
<taxonomicName authority="Rutter 1908: 139" authorityName="Rutter" authorityPageNumber="139" authorityYear="1908" box="[151,482,1042,1066]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="30" pageNumber="531" phylum="Chordata" rank="species" species="robusta">
<emphasis box="[151,304,1042,1066]" italics="true" pageId="30" pageNumber="531">Agosia robusta</emphasis>
<bibRefCitation author="Rutter, C." box="[311,482,1042,1066]" pageId="30" pageNumber="531" pagination="105 - 152" refId="ref30694" refString="Rutter, C. (1908) The fishes of the Sacramento-San Joaquin basin, with a study of their distribution and variation. United States Bureau of Fisheries Bulletin, 27, 105 - 152. [1907]" type="journal article" year="1908">Rutter 1908: 139</bibRefCitation>
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</paragraph>
<paragraph blockId="30.[151,681,1011,1158]" box="[151,667,1072,1096]" pageId="30" pageNumber="531">
<treatmentCitationGroup box="[151,667,1072,1096]" pageId="30" pageNumber="531">
<treatmentCitation author="Evermann, B. W. &amp; Clark, H. W." box="[151,667,1072,1096]" page="55" pageId="30" pageNumber="531" year="1931">
<taxonomicName authority="Evermann and Clark 1931: 55" authorityName="Evermann and Clark" authorityPageNumber="55" authorityYear="1931" box="[151,667,1072,1096]" class="Actinopterygii" family="Cyprinidae" genus="Apocope" kingdom="Animalia" order="Cypriniformes" pageId="30" pageNumber="531" phylum="Chordata" rank="species" species="carringtoni">
<emphasis box="[151,361,1073,1096]" italics="true" pageId="30" pageNumber="531">Apocope carringtoni</emphasis>
<bibRefCitation author="Evermann, B. W. &amp; Clark, H. W." box="[368,667,1072,1096]" pageId="30" pageNumber="531" pagination="1 - 67" refId="ref27727" refString="Evermann, B. W. &amp; Clark, H. W. (1931) A distributional list of freshwater fishes known to occur in California. Division of Fish and Game, Fish Bulletin, 35, 1 - 67." type="journal article" year="1931">Evermann and Clark 1931:55</bibRefCitation>
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</paragraph>
<paragraph blockId="30.[151,681,1011,1158]" box="[151,681,1103,1127]" pageId="30" pageNumber="531">
<treatmentCitationGroup box="[151,681,1103,1127]" pageId="30" pageNumber="531">
<emphasis box="[151,417,1103,1127]" italics="true" pageId="30" pageNumber="531">
<taxonomicName ID-CoL="4S7V3" baseAuthorityName="Miller" baseAuthorityYear="1973" box="[151,353,1103,1127]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="30" pageNumber="531" phylum="Chordata" rank="species" species="osculus">Rhinichthys osculus</taxonomicName>
subsp
</emphasis>
.
<treatmentCitation author="Williams, J. E. &amp; Stern, M. A. &amp; Munhall, A. V. &amp; Anderson, G. A." box="[429,681,1103,1127]" page="243" pageId="30" pageNumber="531" year="1990">
<bibRefCitation author="Williams, J. E. &amp; Stern, M. A. &amp; Munhall, A. V. &amp; Anderson, G. A." box="[429,681,1103,1127]" pageId="30" pageNumber="531" pagination="243 - 248" refId="ref32785" refString="Williams, J. E., Stern, M. A., Munhall, A. V. &amp;. Anderson, G. A. (1990) Conservation status of threatened fishes in the Warner Basin, Oregon. Great Basin Naturalist, 50 (3), 243 - 248. [http: // www. jstor. org / stable / 41712597]" type="journal article" year="1990">
Williams
<emphasis box="[528,579,1103,1127]" italics="true" pageId="30" pageNumber="531">et al.</emphasis>
1990:243
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</paragraph>
<paragraph blockId="30.[151,681,1011,1158]" box="[151,660,1134,1158]" pageId="30" pageNumber="531">
<treatmentCitationGroup box="[151,660,1134,1158]" pageId="30" pageNumber="531">
<treatmentCitation author="Markle, D. F." box="[151,660,1134,1158]" page="50" pageId="30" pageNumber="531" year="2016">
<taxonomicName authority="Markle 2016: 50" authorityName="Markle" authorityPageNumber="50" authorityYear="2016" box="[151,660,1134,1158]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="30" pageNumber="531" phylum="Chordata" rank="subSpecies" species="osculus" subSpecies="klamathensis">
<emphasis box="[151,491,1134,1158]" italics="true" pageId="30" pageNumber="531">Rhinichthys osculus klamathensis</emphasis>
<bibRefCitation author="Markle, D. F." box="[498,660,1134,1158]" pageId="30" pageNumber="531" refId="ref29323" refString="Markle, D. F. (2016) A Guide to the Freshwater Fishes of Oregon. Oregon State University Press, Corvallis, Oregon, 140 pp." type="book" year="2016">Markle 2016:50</bibRefCitation>
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</paragraph>
</subSubSection>
<subSubSection pageId="30" pageNumber="531" type="materials_examined">
<paragraph blockId="30.[151,1437,1204,2023]" pageId="30" pageNumber="531">
<materialsCitation collectingDate="1988-11-02" collectorName="Jack E. Williams." county="Lake County" location="Twelve Mile Creek" pageId="30" pageNumber="531" specimenCount="1" stateProvince="Oregon" typeStatus="holotype">
<emphasis bold="true" box="[151,260,1204,1230]" pageId="30" pageNumber="531">
<typeStatus box="[151,260,1204,1230]" pageId="30" pageNumber="531">Holotype</typeStatus>
</emphasis>
: WFB-122-10-44 (
<figureCitation box="[479,576,1204,1230]" captionStart="FIGURE 1" captionStartId="2.[152,255,1650,1674]" captionTargetBox="[156,1431,489,1626]" captionTargetId="figure-162@2.[156,1431,489,1626]" captionTargetPageId="2" captionText="FIGURE 1. Top: Lahontan Speckled Dace (ca. 70 mm TL) and cover habitat in Taylor Creek, California, a tributary to Lake Tahoe. The body shape, stripe on head, and speckling are characteristic of live fish in many populations from Mexico to Canada that have been considered to be just one species, Speckled Dace (Rhinichthys osculus). Photo by T. L. Taylor, July 17, 1991. Bottom: Preserved specimen of Sacramento Speckled Dace, collected by John Otterbein Snyder from Alameda Creek, California in 1898, showing basic external anatomy. Note the presence of tiny maxillary barbels on head, an example of a distinctive characteristic of some populations. Photo by Jon Fong, California Academy of Sciences, of specimen SU 16172." figureDoi="http://doi.org/10.5281/zenodo.7701359" httpUri="https://zenodo.org/record/7701359/files/figure.png" pageId="30" pageNumber="531">Figure 1</figureCitation>
).
<quantity box="[602,683,1204,1231]" metricMagnitude="-2" metricUnit="m" metricValue="5.9" pageId="30" pageNumber="531" unit="mm" value="59.0">59 mm</quantity>
SL.
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0BFF873FFFFC0BCCC9F8F8:8EA70B9CDE0BFF863D8CFB35CEFBFB47" box="[740,963,1204,1230]" county="Lake County" name="Twelve Mile Creek" pageId="30" pageNumber="531" stateProvince="Oregon">Twelve Mile Creek</location>
,
<collectingCounty box="[975,1124,1204,1230]" pageId="30" pageNumber="531">Lake County</collectingCounty>
,
<collectingRegion box="[1137,1225,1204,1230]" country="United States of America" name="Oregon" pageId="30" pageNumber="531">Oregon</collectingRegion>
.
<collectorName box="[1237,1437,1204,1231]" pageId="30" pageNumber="531">Jack E. Williams.</collectorName>
<date box="[151,363,1240,1266]" pageId="30" pageNumber="531" value="1988-11-02">
<collectingDate box="[151,363,1240,1266]" pageId="30" pageNumber="531" value="1988-11-02">November 2, 1988</collectingDate>
</date>
.
</materialsCitation>
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[199,1108,1276,1303]" pageId="30" pageNumber="531">
<emphasis bold="true" box="[199,319,1276,1302]" pageId="30" pageNumber="531">
<typeStatus box="[199,319,1276,1302]" pageId="30" pageNumber="531">Paratypes</typeStatus>
</emphasis>
: WFB-122-10-42a—122-10-42j (n = 10
<emphasis bold="true" box="[773,789,1276,1302]" pageId="30" pageNumber="531">).</emphasis>
Same location and collector
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[199,563,1312,1338]" pageId="30" pageNumber="531">
<emphasis bold="true" box="[199,319,1312,1338]" pageId="30" pageNumber="531">Meristics:</emphasis>
<typeStatus box="[326,430,1312,1338]" pageId="30" pageNumber="531">Holotype</typeStatus>
(
<typeStatus box="[445,556,1312,1338]" pageId="30" pageNumber="531">paratypes</typeStatus>
)
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[206,692,1348,1375]" pageId="30" pageNumber="531">
<emphasis box="[206,359,1348,1374]" italics="true" pageId="30" pageNumber="531">Length (mm):</emphasis>
standard 59; fork 67; total 71
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" pageId="30" pageNumber="531">
<emphasis box="[206,423,1384,1410]" italics="true" pageId="30" pageNumber="531">Lateral line scales:</emphasis>
76 (6184). Lateral line incomplete in most individuals; counts include 23 scales beyond end of lateral line.
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[199,614,1456,1482]" pageId="30" pageNumber="531">
<emphasis box="[199,485,1456,1482]" italics="true" pageId="30" pageNumber="531">Scales above lateral line:</emphasis>
13 (1114)
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[199,600,1492,1518]" pageId="30" pageNumber="531">
<emphasis box="[199,485,1492,1518]" italics="true" pageId="30" pageNumber="531">Scales below lateral line:</emphasis>
11 (913)
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[199,871,1528,1555]" pageId="30" pageNumber="531">
<emphasis box="[199,380,1528,1554]" italics="true" pageId="30" pageNumber="531">Dorsal-fin rays:</emphasis>
8(8), counts include single unbranched ray.
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[199,851,1564,1591]" pageId="30" pageNumber="531">
<emphasis box="[199,346,1564,1590]" italics="true" pageId="30" pageNumber="531">Anal-fin rays</emphasis>
: 7 (7), counts include single unbranched ray.
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[199,896,1600,1627]" pageId="30" pageNumber="531">
<emphasis box="[199,390,1600,1626]" italics="true" pageId="30" pageNumber="531">Pectoral-fin rays</emphasis>
12 (1013), counts include unbranched rays.
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[199,733,1636,1663]" pageId="30" pageNumber="531">
<emphasis box="[199,269,1636,1662]" italics="true" pageId="30" pageNumber="531">Pelvic</emphasis>
7 (79), counts include unbranched rays.
</paragraph>
<paragraph blockId="30.[151,1437,1204,2023]" box="[199,514,1672,1698]" pageId="30" pageNumber="531">
<emphasis box="[199,376,1672,1698]" italics="true" pageId="30" pageNumber="531">Caudal-fin rays</emphasis>
19 (1920).
</paragraph>
</subSubSection>
<subSubSection pageId="30" pageNumber="531" type="diagnosis">
<paragraph blockId="30.[151,1437,1204,2023]" pageId="30" pageNumber="531">
<emphasis bold="true" box="[199,322,1708,1734]" pageId="30" pageNumber="531">Diagnosis.</emphasis>
Genetically distinct Speckled Dace endemic to the isolated Warner Basin in
<collectingRegion box="[1178,1262,1708,1734]" country="United States of America" name="Oregon" pageId="30" pageNumber="531">Oregon</collectingRegion>
and
<collectingRegion box="[1316,1433,1708,1734]" country="United States of America" name="California" pageId="30" pageNumber="531">California</collectingRegion>
. Relatively small adult size (&lt;
<quantity box="[506,581,1744,1771]" metricMagnitude="-1" metricUnit="m" metricValue="1.0" pageId="30" pageNumber="531" unit="cm" value="10.0">10 cm</quantity>
SL); classic Speckled Dace body shape: thick caudal peduncle, robust (subcylindrical) body, small fins, and bluntly pointed snout with subterminal mouth. Maxillary barbels and frenum usually present.
</paragraph>
</subSubSection>
<subSubSection lastPageId="31" lastPageNumber="532" pageId="30" pageNumber="531" type="description">
<paragraph blockId="30.[151,1437,1204,2023]" lastBlockId="31.[151,1437,150,1905]" lastPageId="31" lastPageNumber="532" pageId="30" pageNumber="531">
<emphasis bold="true" box="[199,345,1852,1878]" pageId="30" pageNumber="531">Description.</emphasis>
The description is the same as Sacramento Speckled Dace.
<bibRefCitation author="Snyder, J. O." box="[1042,1205,1852,1878]" pageId="30" pageNumber="531" pagination="69 - 102" refId="ref31713" refString="Snyder, J. O. (1908) Relationships of the fish fauna of the lakes of Southeastern Oregon. U. S. Bureau of Fisheries Bulletin, 27, 69 - 102." type="journal article" year="1908">Snyder (1908)</bibRefCitation>
examined Speckled Dace from throughout the arid basins of southeastern
<collectingRegion box="[769,853,1888,1914]" country="United States of America" name="Oregon" pageId="30" pageNumber="531">Oregon</collectingRegion>
and could find no distinguishing morphological or meristic features to separate Warner Speckled Dace from dace in other basins, not even lateral line scale numbers (which partially define Klamath Speckled Dace). For lateral line scales, Snyders counts were: Warner Basin 68 71(N=58), Sacramento, 6174 (N=143), and Klamath 6878 (n=18). He also noted that in Warner Speckled Dace, maxillary barbels were usually present (45 out of 53 fish examined). For comparison, they were present on 43/48 Klamath basin dace but only 9/68 for Sacramento basin dace. A frenum is usually present.
</paragraph>
</subSubSection>
<subSubSection pageId="31" pageNumber="532" type="distribution">
<paragraph blockId="31.[151,1437,150,1905]" pageId="31" pageNumber="532">
<emphasis bold="true" box="[199,345,222,248]" pageId="31" pageNumber="532">Distribution</emphasis>
. In a survey of fishes of the Warner Basin,
<bibRefCitation author="Williams, J. E. &amp; Stern, M. A. &amp; Munhall, A. V. &amp; Anderson, G. A." box="[880,1141,223,249]" pageId="31" pageNumber="532" pagination="243 - 248" refId="ref32785" refString="Williams, J. E., Stern, M. A., Munhall, A. V. &amp;. Anderson, G. A. (1990) Conservation status of threatened fishes in the Warner Basin, Oregon. Great Basin Naturalist, 50 (3), 243 - 248. [http: // www. jstor. org / stable / 41712597]" type="journal article" year="1990">
Williams
<emphasis box="[994,1055,223,249]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
(1990)
</bibRefCitation>
found Speckled Dace in Twelvemile Creek, Twentymile Creek, Deep Creek, and upper Honey Creek, plus the isolated Foskett Spring. Only Twelvemile Creek has headwaters (and Speckled Dace) in
<collectingRegion box="[827,944,295,321]" country="United States of America" name="California" pageId="31" pageNumber="532">California</collectingRegion>
. Foskett Spring is on the edge of Coleman Lake, on the southeast corner of the basin; the lake is dry most of the time. The mtDNA study by
<bibRefCitation author="Smith, G. R. &amp; Chow, J. &amp; Unmack, P. J. &amp; Markle, D. F. &amp; Dowling, T. E." box="[1227,1436,331,357]" pageId="31" pageNumber="532" pagination="1 - 83" refId="ref31441" refString="Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. &amp; Dowling, T. E. (2017) Evolution of the Rhinichthys osculus complex (Teleostei: Cyprinidae) in Western North America. Miscellaneous Publications Museum of Zoology University of Michigan, 204 (2), 1 - 83" type="journal article" year="2017">
Smith
<emphasis box="[1300,1356,331,357]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
(2017)
</bibRefCitation>
indicates Speckled Dace in the Warner Basin share a recent ancestry with those from the Chewaucan River to the north (which flows into Lake Abert,
<collectingRegion box="[581,667,403,429]" country="United States of America" name="Oregon" pageId="31" pageNumber="532">Oregon</collectingRegion>
) and with those in Wall Canyon Creek (
<collectingRegion box="[1141,1230,403,429]" country="United States of America" name="Nevada" pageId="31" pageNumber="532">Nevada</collectingRegion>
) to the southeast. Further genomic studies may show these populations could be included within Warner Speckled Dace.
</paragraph>
<paragraph blockId="31.[151,1437,150,1905]" pageId="31" pageNumber="532">
<emphasis bold="true" box="[199,303,474,500]" pageId="31" pageNumber="532">Geology.</emphasis>
The Warner Valley is one of a number of isolated watersheds within the northern Great Basin, which have a long complex history of occasional connectivity to big river systems, such as the Snake and the Klamath (
<bibRefCitation author="Smith, G. R. &amp; Dowling, T. E. &amp; Gobalet, K. T. &amp; Lugaski, T. &amp; Shiozawa, D. &amp; Evans, P." box="[159,356,547,573]" pageId="31" pageNumber="532" pagination="175 - 233" refId="ref31509" refString="Smith, G. R., Dowling, T. E., Gobalet, K. T., Lugaski, T., Shiozawa, D. &amp; Evans, P. (2002) Biogeography and timing rates of evolutionary events among Great Basin fishes. In: Hershler, R., Madsen, D. B. &amp; Curry, D. R. (Eds.), Great Basin Aquatic Systems History, Smithsonian Contributions to the Earth Sciences, 33, pp. 175 - 233." type="journal article" year="2002">
Smith
<emphasis box="[234,289,547,573]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
2002
</bibRefCitation>
). The onset of Great Basin faulting and uplift of the Warner Range (
<emphasis box="[1107,1141,547,572]" italics="true" pageId="31" pageNumber="532">ca.</emphasis>
3 mya) likely resulted in a permanent separation and isolation of the Warner Basin from the Sacramento and Klamath basins, sometime in the late-Pleistocene (~1.0 mya) (Eggar
<emphasis box="[553,610,619,645]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
2011). This isolation is reflected in other endemic fishes which co-occur with Warner Speckled Dace: Warner Sucker (
<taxonomicName authorityName="Snyder" authorityYear="1908" box="[656,930,655,681]" class="Actinopterygii" family="Catostomidae" genus="Catostomus" kingdom="Animalia" order="Cypriniformes" pageId="31" pageNumber="532" phylum="Chordata" rank="species" species="warnerensis">
<emphasis box="[656,930,655,681]" italics="true" pageId="31" pageNumber="532">Catostomus warnerensis</emphasis>
</taxonomicName>
), Great Basin Redband Trout (
<taxonomicName baseAuthorityName="Walbaum" baseAuthorityYear="1792" class="Actinopterygii" family="Salmonidae" genus="Oncorhynchus" kingdom="Animalia" order="Salmoniformes" pageId="31" pageNumber="532" phylum="Chordata" rank="species" species="mykiss">
<emphasis italics="true" pageId="31" pageNumber="532">Oncorhynchus mykiss</emphasis>
</taxonomicName>
subsp.) and Tui Chub (
<taxonomicName baseAuthorityName="Girard" baseAuthorityYear="1856" box="[495,701,691,717]" class="Actinopterygii" family="Cyprinidae" genus="Siphatales" kingdom="Animalia" order="Cypriniformes" pageId="31" pageNumber="532" phylum="Chordata" rank="species" species="bicolor">
<emphasis box="[495,701,691,717]" italics="true" pageId="31" pageNumber="532">Siphatales bicolor</emphasis>
</taxonomicName>
subsp.) (
<bibRefCitation author="Williams, J. E. &amp; Stern, M. A. &amp; Munhall, A. V. &amp; Anderson, G. A." box="[805,1041,691,717]" pageId="31" pageNumber="532" pagination="243 - 248" refId="ref32785" refString="Williams, J. E., Stern, M. A., Munhall, A. V. &amp;. Anderson, G. A. (1990) Conservation status of threatened fishes in the Warner Basin, Oregon. Great Basin Naturalist, 50 (3), 243 - 248. [http: // www. jstor. org / stable / 41712597]" type="journal article" year="1990">
Williams
<emphasis box="[917,974,691,717]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
1990
</bibRefCitation>
;
<bibRefCitation author="Markle, D. F." box="[1054,1204,691,717]" pageId="31" pageNumber="532" refId="ref29323" refString="Markle, D. F. (2016) A Guide to the Freshwater Fishes of Oregon. Oregon State University Press, Corvallis, Oregon, 140 pp." type="book" year="2016">Markle 2016</bibRefCitation>
). In fact, the Warner Valley is one of a number of isolated watersheds within the northern Great Basin that support endemic aquatic species (Hubbs
<emphasis box="[328,385,763,789]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
1978), so it is likely that Speckled Dace are endemic to other watersheds as well.
</paragraph>
<paragraph blockId="31.[151,1437,150,1905]" pageId="31" pageNumber="532">
<emphasis bold="true" box="[199,420,798,824]" pageId="31" pageNumber="532">Genetics/genomics</emphasis>
.
<bibRefCitation author="Ardren, W. R. &amp; Baumsteiger, J. &amp; Allen, C. S." box="[435,668,799,825]" pageId="31" pageNumber="532" pagination="1299 - 1315" refId="ref26452" refString="Ardren, W. R., Baumsteiger, J. &amp; Allen, C. S. (2010) Genetic analysis and uncertain taxonomic status of threatened Foskett Spring Speckled Dace. Conservation Genetics, 11 (4), 1299 - 1315. https: // doi. org / 10.1007 / s 10592 - 009 - 9959 - 0" type="journal article" year="2010">
Ardren
<emphasis box="[523,582,799,825]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
(2010)
</bibRefCitation>
, using mtDNA, suggested that Speckled Dace in the Warner Basin are different at the species level from dace in the neighboring Goose Lake Basin (Sacramento drainage).
<bibRefCitation author="Hoekzema, K. &amp; Sidlauskas, B. L." pageId="31" pageNumber="532" pagination="238 - 250" refId="ref28517" refString="Hoekzema, K. &amp; Sidlauskas, B. L. (2014) Molecular phylogenetics and microsatellite analysis reveal cryptic species of Speckled Dace (Cyprinidae: Rhinichthys osculus) in Oregon's Great Basin. Molecular Phylogenetics and Evolution, 77, 238 - 250. https: // doi. org / 10.1016 / j. ympev. 2014.04.027" type="journal article" year="2014">Hoekzema and Sidlauskas (2014)</bibRefCitation>
examined mtDNA from dace populations in the Warner Basin and in surrounding basins in
<collectingRegion box="[181,269,907,933]" country="United States of America" name="Oregon" pageId="31" pageNumber="532">Oregon</collectingRegion>
. They found a high level of genetic divergence in dace from the basins, dating to the Pliocene or early Pleistocene, which “…should likely be elevated to species-status once their full geographic extent is discovered, and their morphological diversity described (
<bibRefCitation author="Hoekzema, K. &amp; Sidlauskas, B. L." box="[649,1059,978,1005]" pageId="31" pageNumber="532" pagination="238 - 250" refId="ref28517" refString="Hoekzema, K. &amp; Sidlauskas, B. L. (2014) Molecular phylogenetics and microsatellite analysis reveal cryptic species of Speckled Dace (Cyprinidae: Rhinichthys osculus) in Oregon's Great Basin. Molecular Phylogenetics and Evolution, 77, 238 - 250. https: // doi. org / 10.1016 / j. ympev. 2014.04.027" type="journal article" year="2014">Hoekzema and Sidlauskas 2014:245</bibRefCitation>
). Distinct lineages included fishes from Goose Lake, Silver Lake, Lake Abert, and the Warner basin, including Foskett Spring. The mtDNA study of
<bibRefCitation author="Smith, G. R. &amp; Chow, J. &amp; Unmack, P. J. &amp; Markle, D. F. &amp; Dowling, T. E." box="[151,364,1051,1077]" pageId="31" pageNumber="532" pagination="1 - 83" refId="ref31441" refString="Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. &amp; Dowling, T. E. (2017) Evolution of the Rhinichthys osculus complex (Teleostei: Cyprinidae) in Western North America. Miscellaneous Publications Museum of Zoology University of Michigan, 204 (2), 1 - 83" type="journal article" year="2017">
Smith
<emphasis box="[225,282,1051,1077]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
(2017)
</bibRefCitation>
and the genomic study of
<bibRefCitation author="Su, Y. &amp; Moyle, P. B. &amp; Campbell, M. A. &amp; Finger, A. J. &amp; O'Rourke, S. &amp; Baumsteiger, J. &amp; Miller, M. R." box="[663,838,1051,1077]" pageId="31" pageNumber="532" pagination="695 - 710" refId="ref32073" refString="Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. &amp; Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388" type="journal article" year="2022">
Su
<emphasis box="[700,756,1051,1077]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
(2022)
</bibRefCitation>
both indicate that Warner Basin Speckled Dace are a distinct lineage, divergent from the Sacramento and Klamath lineages, at the subspecies or species level.
</paragraph>
</subSubSection>
<subSubSection pageId="31" pageNumber="532" type="etymology">
<paragraph blockId="31.[151,1437,150,1905]" pageId="31" pageNumber="532">
<emphasis bold="true" box="[199,334,1123,1149]" pageId="31" pageNumber="532">Etymology.</emphasis>
The name honors the Goyatöka people who, before the invasion of their homeland by Euro- Americans, lived on lands in the Warner Basin where Speckled Dace occupied the streams, lakes, and springs (
<bibRefCitation author="Dixon, R. B." box="[159,299,1195,1221]" pageId="31" pageNumber="532" pagination="208 - 220" refId="ref27487" refString="Dixon, R. B. (1908) Notes on the Achomawi and Atsugewii Indians of northern California. American Anthropologist, 10 (2), 208 - 220. https: // doi. org / 10.1525 / aa. 1908.10.2.02 a 00030" type="journal article" year="1908">Dixon 1908</bibRefCitation>
,
<bibRefCitation author="Stewart, O. C." box="[313,468,1195,1221]" pageId="31" pageNumber="532" pagination="127 - 149" refId="ref31973" refString="Stewart, O. C. (1939) The Northern Paiute Bands. Anthropological Records, 2 (3), 127 - 149." type="journal article" year="1939">Stewart 1939</bibRefCitation>
). The common name indicates the geographic location of the dace populations. The basin is named for William Warner, an army officer who died while mapping the region.
</paragraph>
<paragraph blockId="31.[151,1437,150,1905]" pageId="31" pageNumber="532">
<emphasis bold="true" box="[199,273,1267,1293]" pageId="31" pageNumber="532">Notes.</emphasis>
Our study supports the finding that the Warner Speckled Dace is a distinct lineage (subspecies) allied with Klamath and Sacramento Speckled Dace.
<bibRefCitation author="Ardren, W. R. &amp; Baumsteiger, J. &amp; Allen, C. S." box="[675,898,1303,1329]" pageId="31" pageNumber="532" pagination="1299 - 1315" refId="ref26452" refString="Ardren, W. R., Baumsteiger, J. &amp; Allen, C. S. (2010) Genetic analysis and uncertain taxonomic status of threatened Foskett Spring Speckled Dace. Conservation Genetics, 11 (4), 1299 - 1315. https: // doi. org / 10.1007 / s 10592 - 009 - 9959 - 0" type="journal article" year="2010">
Ardren
<emphasis box="[761,817,1303,1329]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
(2010)
</bibRefCitation>
and
<bibRefCitation author="Su, Y. &amp; Moyle, P. B. &amp; Campbell, M. A. &amp; Finger, A. J. &amp; O'Rourke, S. &amp; Baumsteiger, J. &amp; Miller, M. R." box="[952,1125,1303,1329]" pageId="31" pageNumber="532" pagination="695 - 710" refId="ref32073" refString="Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. &amp; Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388" type="journal article" year="2022">
Su
<emphasis box="[988,1044,1303,1329]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
(2022)
</bibRefCitation>
show that Warner Speckled Dace have evolutionary ties to the daces in the Klamath and Sacramento Basins. These Speckled Dace became a species of interest when dace in Foskett Spring, in an isolated part of the Warner Basin, were listed as a threatened species without any special taxonomic designation. Subsequent genetic analyses revealed Foskett Spring Speckled Dace could at best be considered a Distinct Population Segment of Warner Speckled Dace; these same analyses suggested that Speckled Dace throughout the Warner Basin constituted a distinct taxon (
<bibRefCitation author="Ardren, W. R. &amp; Baumsteiger, J. &amp; Allen, C. S." box="[1137,1349,1483,1509]" pageId="31" pageNumber="532" pagination="1299 - 1315" refId="ref26452" refString="Ardren, W. R., Baumsteiger, J. &amp; Allen, C. S. (2010) Genetic analysis and uncertain taxonomic status of threatened Foskett Spring Speckled Dace. Conservation Genetics, 11 (4), 1299 - 1315. https: // doi. org / 10.1007 / s 10592 - 009 - 9959 - 0" type="journal article" year="2010">
Ardren
<emphasis box="[1225,1282,1483,1509]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
2010
</bibRefCitation>
).
</paragraph>
<paragraph blockId="31.[151,1437,150,1905]" pageId="31" pageNumber="532">
<emphasis bold="true" box="[199,450,1518,1544]" pageId="31" pageNumber="532">Conservation Status.</emphasis>
Most of what is known about the Warner Speckled Dace and other fishes of the basin comes from surveys for (a) Warner Sucker, which was listed as threatened in 1985, (b) Foskett Spring Speckled Dace, which was also listed as a threatened species in 1985 (USFWS 2019, https://ecos.fws.gov/ecp/species/651), and (c) Great Basin Redband Trout. The Foskett Spring Speckled Dace was delisted in 2019 as the result of extensive habitat improvement, although it remains a conservation dependent species (USFWS 2019). Most of the permanent water in the basin inhabited by the Warner Sucker is dominated by non-native fishes, but a population of native Tui Chub is also present (
<bibRefCitation author="Williams, J. E. &amp; Stern, M. A. &amp; Munhall, A. V. &amp; Anderson, G. A." box="[446,684,1735,1761]" pageId="31" pageNumber="532" pagination="243 - 248" refId="ref32785" refString="Williams, J. E., Stern, M. A., Munhall, A. V. &amp;. Anderson, G. A. (1990) Conservation status of threatened fishes in the Warner Basin, Oregon. Great Basin Naturalist, 50 (3), 243 - 248. [http: // www. jstor. org / stable / 41712597]" type="journal article" year="1990">
Williams
<emphasis box="[558,616,1735,1761]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
1990
</bibRefCitation>
). The remaining Warner Speckled Dace populations (and those of Warner Sucker and Tui Chub) probably qualify for listing as Threatened under the federal ESA. Their decline is due to the combined problems of non-native predatory fishes occupying much of the aquatic habitat, inadequate management of grazing and diversions, and climate change/drought reducing flows and increasing temperatures in streams (
<bibRefCitation author="Williams, J. E. &amp; Stern, M. A. &amp; Munhall, A. V. &amp; Anderson, G. A." box="[251,485,1879,1905]" pageId="31" pageNumber="532" pagination="243 - 248" refId="ref32785" refString="Williams, J. E., Stern, M. A., Munhall, A. V. &amp;. Anderson, G. A. (1990) Conservation status of threatened fishes in the Warner Basin, Oregon. Great Basin Naturalist, 50 (3), 243 - 248. [http: // www. jstor. org / stable / 41712597]" type="journal article" year="1990">
Williams
<emphasis box="[362,418,1879,1905]" italics="true" pageId="31" pageNumber="532">et al.</emphasis>
1990
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>