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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="03D1EC51DE0CFF833FFFF9DBC811F81C" 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. klamathensis Moyle, Buckmaster &amp; Su, 2023, new combination" docType="treatment" docVersion="2" lastPageNumber="528" masterDocId="FFE89429DE15FF983F68FF81CD38FF89" masterDocTitle="Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA" masterLastPageNumber="539" masterPageNumber="501" pageNumber="526" updateTime="1678098333465" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:titleInfo>
<mods:title>Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<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>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<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>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Su, Yingxin</mods:namePart>
<mods:nameIdentifier type="ORCID">0000-0002-4825-4865</mods:nameIdentifier>
<mods:nameIdentifier type="email">pbmoyle@ucdavis.edu</mods:nameIdentifier>
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<mods:title>Zootaxa</mods:title>
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<mods:part>
<mods:date>2023</mods:date>
<mods:detail type="pubDate">
<mods:number>2023-03-06</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>5249</mods:number>
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<mods:detail type="issue">
<mods:number>5</mods:number>
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<mods:extent unit="page">
<mods:start>501</mods:start>
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<mods:url>http://dx.doi.org/10.11646/zootaxa.5249.5.1</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.11646/zootaxa.5249.5.1</mods:identifier>
<mods:identifier type="ISSN">1175-5326</mods:identifier>
<mods:identifier type="Zenodo-Dep">7701357</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.7701341" ID-Zenodo-Dep="7701341" LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C" httpUri="http://treatment.plazi.org/id/03D1EC51DE0CFF833FFFF9DBC811F81C" lastPageId="27" lastPageNumber="528" pageId="25" pageNumber="526">
<subSubSection box="[151,1183,1626,1653]" pageId="25" pageNumber="526" type="nomenclature">
<paragraph blockId="25.[151,1183,1626,1689]" box="[151,1183,1626,1653]" pageId="25" pageNumber="526">
<heading bold="true" box="[151,1183,1626,1653]" fontSize="11" level="1" pageId="25" pageNumber="526" reason="1">
<emphasis bold="true" box="[151,1183,1626,1653]" pageId="25" pageNumber="526">
<taxonomicName authorityName="Moyle &amp; Buckmaster &amp; Su" authorityYear="2023" baseAuthorityName="Moyle &amp; Buckmaster &amp; Su" baseAuthorityYear="2023" box="[151,632,1626,1653]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="subSpecies" species="klamathensis" status="new combination" subSpecies="klamathensis">
<emphasis bold="true" box="[151,632,1626,1653]" italics="true" pageId="25" pageNumber="526">Rhinichthys klamathensis klamathensis</emphasis>
</taxonomicName>
,
<taxonomicNameLabel box="[647,863,1626,1653]" pageId="25" pageNumber="526" rank="species">new combination</taxonomicNameLabel>
, Klamath Speckled Dace,
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="25" pageNumber="526" type="description">
<paragraph blockId="25.[151,1183,1626,1689]" box="[151,217,1663,1689]" pageId="25" pageNumber="526">
<figureCitation box="[151,217,1663,1689]" captionStart="FIGURE 6" captionStartId="26.[152,255,1615,1639]" captionTargetBox="[154,1434,275,1588]" captionTargetId="figure-59@26.[154,1434,272,1592]" captionTargetPageId="26" captionText="FIGURE 6. Top. Holotype, Lahontan Speckled Dace, from Rutter (1903). Middle, Long Valley Speckled Dace from Whitmore Marsh, 2014, showing life colors. Photo by Jacob Katz. Bottom: Western Speckled Dace, type specimen, from Evermann and Meek (1898)." figureDoi="http://doi.org/10.5281/zenodo.7701369" httpUri="https://zenodo.org/record/7701369/files/figure.png" pageId="25" pageNumber="526">Fig. 6</figureCitation>
</paragraph>
<paragraph blockId="25.[151,822,1731,1755]" box="[151,822,1731,1755]" pageId="25" pageNumber="526">
<emphasis bold="true" box="[151,265,1731,1755]" pageId="25" pageNumber="526">Synonymy</emphasis>
. Same as for Western Speckled Dace,
<taxonomicName box="[656,816,1731,1755]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="klamathensis">
<emphasis box="[656,816,1731,1755]" italics="true" pageId="25" pageNumber="526">R. klamathensis</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection box="[151,1124,1801,1828]" pageId="25" pageNumber="526" type="materials_examined">
<paragraph blockId="25.[151,1437,1801,2008]" box="[151,1124,1801,1828]" pageId="25" pageNumber="526">
<materialsCitation box="[151,1117,1801,1828]" location="Western Speckled Dace" pageId="25" pageNumber="526" specimenCount="2" typeStatus="holotype">
<emphasis bold="true" box="[151,437,1801,1827]" pageId="25" pageNumber="526">
<typeStatus box="[151,260,1801,1827]" pageId="25" pageNumber="526">Holotype</typeStatus>
and
<typeStatus box="[319,437,1801,1827]" pageId="25" pageNumber="526">paratypes</typeStatus>
</emphasis>
are the same as for
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0CFF813DFDF88BCE99F8AD" box="[661,929,1801,1828]" name="Western Speckled Dace" pageId="25" pageNumber="526">Western Speckled Dace</location>
,
<taxonomicName box="[940,1117,1801,1827]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="klamathensis">
<emphasis box="[940,1117,1801,1827]" italics="true" pageId="25" pageNumber="526">R. klamathensis</emphasis>
</taxonomicName>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="25" pageNumber="526" type="diagnosis">
<paragraph blockId="25.[151,1437,1801,2008]" pageId="25" pageNumber="526">
<emphasis bold="true" box="[199,314,1837,1863]" pageId="25" pageNumber="526">Diagnosis</emphasis>
.
<materialsCitation county="Western Speckled Dace" location="River Basin" municipality="Klamath River" pageId="25" pageNumber="526" specimenCount="1" stateProvince="Oregon and California">
Same as
<collectingCounty box="[433,704,1837,1864]" pageId="25" pageNumber="526">Western Speckled Dace</collectingCounty>
,
<taxonomicName box="[718,897,1837,1863]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="klamathensis">
<emphasis box="[718,897,1837,1863]" italics="true" pageId="25" pageNumber="526">R. klamathensis</emphasis>
</taxonomicName>
; the Speckled Dace lineage that is abundant in the streams, lakes, and other aquatic habitats in the
<collectingMunicipality box="[740,907,1873,1899]" pageId="25" pageNumber="526">
Klamath
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0CFF813C25F8D0CEB3F8E2" box="[845,907,1873,1899]" county="Western Speckled Dace" municipality="Klamath River" name="River Basin" pageId="25" pageNumber="526" stateProvince="Oregon and California">River</location>
</collectingMunicipality>
Basin in
<collectingRegion box="[1018,1275,1873,1899]" isEnumeration="true" pageId="25" pageNumber="526">Oregon and California</collectingRegion>
, including the
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0CFF813FFFF8F4CC14F806" box="[151,300,1909,1935]" county="Western Speckled Dace" municipality="Klamath River" name="Trinity River" pageId="25" pageNumber="526" stateProvince="Oregon and California">Trinity River</location>
in
<collectingRegion box="[341,458,1909,1935]" country="United States of America" name="California" pageId="25" pageNumber="526">California</collectingRegion>
. Distinguished statistically by having somewhat smaller scales (scales in lateral line, 6878; mean 73, n =49) than the other two subspecies (Sacramento, 5483, mean 70, n=123; Warner, 6871, mean 69, n=58). Otherwise, it is not distinguishable from other Speckled Dace except by genomics and distribution
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="26" pageNumber="527" type="description">
<paragraph blockId="26.[151,1437,150,249]" pageId="26" pageNumber="527">
<emphasis bold="true" box="[199,345,150,176]" pageId="26" pageNumber="527">Description.</emphasis>
<bibRefCitation author="Evermann, B. W. &amp; Meek, S. B." box="[351,665,150,177]" pageId="26" pageNumber="527" pagination="15 - 84" refId="ref27663" refString="Evermann, B. W. &amp; Meek, S. B. (1898) A report upon salmon investigations in the Columbia River and elsewhere on the Pacific Coast in 1896. Bulletin of the U. S. Fish Commission, 17, 15 - 84. https: // doi. org / 10.5962 / bhl. title. 35601" type="journal article" year="1898">Evermann and Meek (1898)</bibRefCitation>
described this cryptic subspecies as
<taxonomicName authorityName="Evermann and Meek" authorityYear="1898" box="[1075,1305,150,177]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="26" pageNumber="527" phylum="Chordata" rank="species" species="klamathensis">
<emphasis box="[1075,1305,150,177]" italics="true" pageId="26" pageNumber="527">Agosia klamathensis</emphasis>
</taxonomicName>
. Therefore, the description quoted in the Western Speckled Dace account is also for this subspecies, and it applies to the other two subspecies as well.
</paragraph>
</subSubSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7701369" ID-Zenodo-Dep="7701369" httpUri="https://zenodo.org/record/7701369/files/figure.png" pageId="26" pageNumber="527" startId="26.[152,255,1615,1639]" targetBox="[154,1434,275,1588]" targetPageId="26">
<paragraph blockId="26.[151,1437,1615,1712]" pageId="26" pageNumber="527">
<emphasis bold="true" box="[152,279,1615,1640]" pageId="26" pageNumber="527">FIGURE 6.</emphasis>
Top. Holotype, Lahontan Speckled Dace, from
<bibRefCitation author="Rutter, C." box="[761,900,1616,1640]" pageId="26" pageNumber="527" pagination="145 - 148" refId="ref30655" refString="Rutter, C. (1903) Notes on fishes from streams and lakes of northeastern California not tributary to the Sacramento basin. United States Fish Commission Bulletin, 22, 145 - 148. [1902]" type="journal article" year="1903">Rutter (1903)</bibRefCitation>
. Middle, Long Valley Speckled Dace from Whitmore Marsh, 2014, showing life colors. Photo by Jacob Katz. Bottom: Western Speckled Dace, type specimen, from
<bibRefCitation author="Evermann, B. W. &amp; Meek, S. B." pageId="26" pageNumber="527" pagination="15 - 84" refId="ref27663" refString="Evermann, B. W. &amp; Meek, S. B. (1898) A report upon salmon investigations in the Columbia River and elsewhere on the Pacific Coast in 1896. Bulletin of the U. S. Fish Commission, 17, 15 - 84. https: // doi. org / 10.5962 / bhl. title. 35601" type="journal article" year="1898">Evermann and Meek (1898)</bibRefCitation>
.
</paragraph>
</caption>
<subSubSection lastPageId="27" lastPageNumber="528" pageId="26" pageNumber="527" type="distribution">
<paragraph blockId="26.[151,1437,1758,1928]" pageId="26" pageNumber="527">
<emphasis bold="true" box="[199,345,1758,1784]" pageId="26" pageNumber="527">Distribution</emphasis>
. The Klamath Speckled Dace is found throughout the Klamath Basin in
<collectingRegion box="[1175,1259,1758,1784]" country="United States of America" name="Oregon" pageId="26" pageNumber="527">Oregon</collectingRegion>
and
<collectingRegion box="[1316,1432,1758,1784]" country="United States of America" name="California" pageId="26" pageNumber="527">California</collectingRegion>
, including the Klamath and Trinity rivers and tributaries, as well as in the Rogue River to the north (Wiesenfeld
<emphasis italics="true" pageId="26" pageNumber="527">et al.</emphasis>
2017). It is also widespread in the upper Klamath Basin, including Upper Klamath Lake. The Speckled Dace in the Eel River, to the south of the Klamath Basin, is an introduced population of
<emphasis box="[1053,1104,1866,1892]" italics="true" pageId="26" pageNumber="527">R. k.</emphasis>
<taxonomicName authority="(Kinziger et al. 2011)" baseAuthorityName="Kinziger" baseAuthorityYear="2011" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="26" pageNumber="527" phylum="Chordata" rank="species" species="klamathensis">
<emphasis box="[1111,1257,1866,1892]" italics="true" pageId="26" pageNumber="527">klamathensis</emphasis>
(
<bibRefCitation author="Kinziger, A. P. &amp; Nakamoto, R. J. &amp; Anderson, E. C. &amp; Harvey, B. C." pageId="26" pageNumber="527" pagination="73 - 84" refId="ref29117" refString="Kinziger, A. P., Nakamoto, R. J., Anderson, E. C. &amp; Harvey, B. C. (2011) Small founding number and low genetic diversity in an introduced species exhibiting limited invasion success (Speckled Dace, Rhinichthys osculus). Ecology and Evolution, 1 (1), 73 - 84. https: // doi. org / 10.1002 / ece 3.8" type="journal article" year="2011">
Kinziger
<emphasis box="[1380,1437,1866,1892]" italics="true" pageId="26" pageNumber="527">et al.</emphasis>
2011
</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph blockId="26.[151,1436,1974,2036]" lastBlockId="27.[151,1437,151,1941]" lastPageId="27" lastPageNumber="528" pageId="26" pageNumber="527">
<emphasis bold="true" box="[199,477,1974,2000]" pageId="26" pageNumber="527">Geology/zoogeography.</emphasis>
The geologic history of the Klamath region is complex. The Snake River, now a tributary to the Columbia River, was once a major river that originated in the
<collectingRegion box="[1019,1162,2010,2036]" country="United States of America" name="Idaho" pageId="26" pageNumber="527">Idaho region</collectingRegion>
and flowed to the ocean via the ancestral Klamath River during the Pliocene (
<bibRefCitation author="Minckley, W. L. &amp; Hendrickson, D. A. &amp; Bond, C. E." box="[768,1014,151,177]" pageId="27" pageNumber="528" pagination="519 - 614" refId="ref29499" refString="Minckley, W. L., Hendrickson, D. A. &amp; Bond, C. E. (1986) Geography of western North America freshwater fishes: Description and relationship to intercontinental tectonism. In: Hocutt, C. H. &amp; Wiley, E. O. (Eds.), The Zoogeography of North American Freshwater Fishes. New York, John Wiley and Sons, pp. 519 - 614." type="book chapter" year="1986">
Minckley
<emphasis box="[887,946,151,177]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
1986
</bibRefCitation>
,
<bibRefCitation author="Smith, G. R. &amp; Chow, J. &amp; Unmack, P. J. &amp; Markle, D. F. &amp; Dowling, T. E." box="[1028,1226,151,177]" pageId="27" pageNumber="528" 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="[1104,1156,151,177]" italics="true" pageId="27" pageNumber="528">et al</emphasis>
2017
</bibRefCitation>
). When the Snake broke through to the Klamath, it carried with it the lake fauna of the Great Basin, which gave rise to the presentday endemic, freshwater-dispersing fish fauna (12 species) of the upper Klamath basin (
<bibRefCitation author="Moyle, P. B." box="[1142,1285,223,249]" pageId="27" pageNumber="528" refId="ref29646" refString="Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp." type="book" year="2002">Moyle 2002</bibRefCitation>
), presumably including Speckled Dace.
<bibRefCitation author="Pfrender, M. E. &amp; Hicks, J. &amp; Lynch, M." box="[455,758,259,285]" pageId="27" pageNumber="528" pagination="490 - 502" refId="ref30505" refString="Pfrender, M. E., Hicks, J. &amp; Lynch, M. (2004) Biogeographic patterns and current distribution of molecular-genetic variation among populations of Speckled Dace, Rhinichthys osculus (Girard). Molecular Phylogenetics and Evolution, 30 (3), 490 - 502. https: // doi. org / 10.1016 / S 1055 - 7903 (03) 00242 - 2" type="journal article" year="2004">
Pfrender
<emphasis box="[562,621,259,285]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
(2004:498)
</bibRefCitation>
, using a molecular clock based on mtDNA from Speckled Dace from the major river basins of
<collectingRegion box="[573,661,295,321]" country="United States of America" name="Oregon" pageId="27" pageNumber="528">Oregon</collectingRegion>
, speculated that “…the levels of sequence divergence in
<taxonomicName baseAuthorityName="Miller" baseAuthorityYear="1973" box="[1318,1436,295,321]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="27" pageNumber="528" phylum="Chordata" rank="species" species="osculus">
<emphasis box="[1318,1436,295,321]" italics="true" pageId="27" pageNumber="528">R. osculus</emphasis>
</taxonomicName>
among these major basins are more consistent with a Pliocene or Miocene sundering of gene flow between major basins. Miocene isolation of these river systems is substantially earlier than has previously been suggested.” They thought such ancient divergence could explain the genetic diversity found in dace within the Klamath basin, a diversity that was also documented by Wiesenfeld
<emphasis box="[738,797,439,465]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
(2017). See also the geology discussion under Western Speckled Dace.
</paragraph>
<paragraph blockId="27.[151,1437,151,1941]" pageId="27" pageNumber="528">
Geographically, Klamath Speckled Dace co-occur with other fish species endemic to the Klamath watershed. However, the endemic fishes in the upper watershed (above Klamath Falls) are different from those in the lower river and it appears that dace from the two regions diverge as well, a situation noted also for Marbled Sculpin,
<taxonomicName authority="(Daniels and Moyle 1984)" baseAuthorityName="Daniels and Moyle" baseAuthorityYear="1984" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="27" pageNumber="528" phylum="Chordata" rank="species" species="klamathensis">
<emphasis italics="true" pageId="27" pageNumber="528">Cottus klamathensis</emphasis>
(
<bibRefCitation author="Daniels, R. A. &amp; Moyle, P. B." box="[312,594,619,645]" pageId="27" pageNumber="528" pagination="949 - 959" refId="ref27398" refString="Daniels, R. A. &amp; Moyle, P. B. (1984) Geographic variation and a taxonomic reappraisal of the Marbled Sculpin, Cottus klamathensis. Copeia, 1984, 949 - 959. https: // doi. org / 10.2307 / 1445339" type="journal article" year="1984">Daniels and Moyle 1984</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph blockId="27.[151,1437,151,1941]" pageId="27" pageNumber="528">
<emphasis bold="true" box="[199,427,654,680]" pageId="27" pageNumber="528">Genetics/genomics.</emphasis>
<materialsCitation collectionCode="BS" county="Sacramento" location="Goose Lake" municipality="Pit River" pageId="27" pageNumber="528" specimenCount="1" stateProvince="California and Oregon">
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="[686,860,655,681]" pageId="27" pageNumber="528" 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="[722,779,655,681]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
(2022)
</bibRefCitation>
unveiled the three lineages within
<taxonomicName baseAuthorityName="Daniels and Moyle" baseAuthorityYear="1984" box="[1252,1429,655,681]" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="27" pageNumber="528" phylum="Chordata" rank="species" species="klamathensis">
<emphasis box="[1252,1269,655,680]" italics="true" pageId="27" pageNumber="528">R</emphasis>
.
<emphasis box="[1283,1429,655,681]" italics="true" pageId="27" pageNumber="528">klamathensis</emphasis>
</taxonomicName>
, as discussed. Within the Klamath drainage, Wiesenfeld
<emphasis box="[762,816,691,717]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
(2017:8), using analyses of microsatellites and mtDNA, found that The KlamathTrinity Basin Speckled Dace were resolved as nearly monophyletic [bootstrap (
<collectionCode box="[1339,1375,727,753]" pageId="27" pageNumber="528">BS</collectionCode>
) 77], and exhibited a sister group relationship with nearby basins in
<collectingRegion box="[862,1116,763,789]" isEnumeration="true" pageId="27" pageNumber="528">California and Oregon</collectingRegion>
(
<collectingCounty box="[1132,1269,763,789]" pageId="27" pageNumber="528">Sacramento</collectingCounty>
,
<collectingMunicipality box="[1281,1384,763,789]" pageId="27" pageNumber="528">Pit River</collectingMunicipality>
, and
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0EFF833FFFFC9ECC1BFCB0" box="[151,291,799,825]" county="Sacramento" municipality="Pit River" name="Goose Lake" pageId="27" pageNumber="528" stateProvince="California and Oregon">Goose Lake</location>
).
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0EFF833E5EFC9ECF37FCB0" box="[310,527,799,825]" county="Sacramento" municipality="Pit River" name="The Klamath River" pageId="27" pageNumber="528" stateProvince="California and Oregon">The Klamath River</location>
and
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0EFF833D2EFC9ECFE1FCB0" box="[582,729,799,825]" county="Sacramento" municipality="Pit River" name="Trinity River" pageId="27" pageNumber="528" stateProvince="California and Oregon">Trinity River</location>
populations, however, were found to be sufficiently distinct to suggest the two rivers were once isolated from one another, with their connection being fairly recent.
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0EFF833A60FCC2C80CFCD4" box="[1288,1332,835,861]" county="Sacramento" municipality="Pit River" name="The" pageId="27" pageNumber="528" stateProvince="California and Oregon">The</location>
presence of apparent hybrids in the region where the two rivers meet today also supports this conclusion (
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0EFF833BB3FCE6C863FC08" box="[1243,1371,871,897]" county="Sacramento" municipality="Pit River" name="Wiesenfeld" pageId="27" pageNumber="528" stateProvince="California and Oregon">Wiesenfeld</location>
<emphasis box="[1379,1437,871,897]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
2017). A genetically distinct population is also found in
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0EFF833C6DFC0ACEABFC2C" box="[773,915,907,933]" county="Sacramento" municipality="Pit River" name="Jenny Creek" pageId="27" pageNumber="528" stateProvince="California and Oregon">Jenny Creek</location>
, a tributary to the
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0EFF833B33FC0AC9C7FC2C" box="[1115,1279,907,933]" county="Sacramento" municipality="Pit River" name="Klamath River" pageId="27" pageNumber="528" stateProvince="California and Oregon">Klamath River</location>
that is largely in
<collectingRegion box="[180,264,943,969]" country="United States of America" name="Oregon" pageId="27" pageNumber="528">Oregon</collectingRegion>
(
<location LSID="urn:lsid:plazi:treatment:03D1EC51DE0CFF833FFFF9DBC811F81C:8EA70B9CDE0EFF833E7EFC2ECCAEFC40" box="[278,406,943,969]" county="Sacramento" municipality="Pit River" name="Wiesenfeld" pageId="27" pageNumber="528" stateProvince="California and Oregon">Wiesenfeld</location>
<emphasis box="[414,470,943,969]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
2017)
</materialsCitation>
.
</paragraph>
<paragraph blockId="27.[151,1437,151,1941]" pageId="27" pageNumber="528">
<bibRefCitation author="Pfrender, M. E. &amp; Hicks, J. &amp; Lynch, M." box="[199,433,979,1005]" pageId="27" pageNumber="528" pagination="490 - 502" refId="ref30505" refString="Pfrender, M. E., Hicks, J. &amp; Lynch, M. (2004) Biogeographic patterns and current distribution of molecular-genetic variation among populations of Speckled Dace, Rhinichthys osculus (Girard). Molecular Phylogenetics and Evolution, 30 (3), 490 - 502. https: // doi. org / 10.1016 / S 1055 - 7903 (03) 00242 - 2" type="journal article" year="2004">
Pfrender
<emphasis box="[303,352,979,1005]" italics="true" pageId="27" pageNumber="528">et al</emphasis>
(2004)
</bibRefCitation>
used mtDNA to compare dace from the upper Klamath and Rogue rivers with those from two tributaries to the Columbia River in
<collectingRegion box="[625,713,1015,1041]" country="United States of America" name="Oregon" pageId="27" pageNumber="528">Oregon</collectingRegion>
. They found species-level differences between dace in the two great watersheds, with estimated divergence times of 3.9 to 9.6 million years ago. Within the upper Klamath basin, Pferender
<emphasis box="[267,323,1087,1113]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
(2004) found considerable genetic structure but of more recent origin (&lt;1 million years), including Jenny Creek as a distinct population.
</paragraph>
<paragraph blockId="27.[151,1437,151,1941]" pageId="27" pageNumber="528">
Overall, analysis of mtDNA data of dace from their entire range showed that dace from the Klamath, Sacramento, and Warner basins share much of the same lineage; together they form a separate lineage within the Northwest clade of
<bibRefCitation author="Smith, G. R. &amp; Chow, J. &amp; Unmack, P. J. &amp; Markle, D. F. &amp; Dowling, T. E." box="[182,396,1231,1257]" pageId="27" pageNumber="528" 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="[255,312,1231,1257]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
(2017)
</bibRefCitation>
.
</paragraph>
<paragraph blockId="27.[151,1437,151,1941]" pageId="27" pageNumber="528">
<emphasis bold="true" box="[199,264,1267,1293]" pageId="27" pageNumber="528">Note.</emphasis>
The Klamath Speckled Dace was originally described as a species that differed from other Speckled Dace by having finer scales (
<bibRefCitation author="Evermann, B. W. &amp; Meek, S. B." box="[480,789,1303,1329]" pageId="27" pageNumber="528" pagination="15 - 84" refId="ref27663" refString="Evermann, B. W. &amp; Meek, S. B. (1898) A report upon salmon investigations in the Columbia River and elsewhere on the Pacific Coast in 1896. Bulletin of the U. S. Fish Commission, 17, 15 - 84. https: // doi. org / 10.5962 / bhl. title. 35601" type="journal article" year="1898">Evermann and Meek 1898</bibRefCitation>
). Subsequently, it retained its identity as a subspecies,
<emphasis italics="true" pageId="27" pageNumber="528">
<taxonomicName authorityName="Shapovalov and Dill" authorityYear="1950" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="27" pageNumber="528" phylum="Chordata" rank="subSpecies" species="osculus" subSpecies="klamathensis">R. osculus klamathensis</taxonomicName>
.
</emphasis>
This is not surprising given that the Klamath Basin has long been recognized as an isolated basin which supports high endemism in its fishes (
<bibRefCitation author="Moyle, P. B." box="[746,895,1375,1401]" pageId="27" pageNumber="528" refId="ref29646" refString="Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp." type="book" year="2002">Moyle 2002</bibRefCitation>
). The Klamath Speckled Dace has retained its identity through the regions complex geologic history, enduring the rise of mountain ranges, high levels of vulcanism, and invading interior rivers (
<bibRefCitation author="Minckley, W. L. &amp; Hendrickson, D. A. &amp; Bond, C. E." box="[634,892,1447,1473]" pageId="27" pageNumber="528" pagination="519 - 614" refId="ref29499" refString="Minckley, W. L., Hendrickson, D. A. &amp; Bond, C. E. (1986) Geography of western North America freshwater fishes: Description and relationship to intercontinental tectonism. In: Hocutt, C. H. &amp; Wiley, E. O. (Eds.), The Zoogeography of North American Freshwater Fishes. New York, John Wiley and Sons, pp. 519 - 614." type="book chapter" year="1986">
Minckley
<emphasis box="[757,819,1447,1473]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
1986
</bibRefCitation>
). This complex geologic history has kept the Klamath fish fauna isolated and distinct. In the upper Klamath basin, Speckled Dace live with other endemic fishes that had their ancestors in the Great Basin. Even in the lower Klamath River, which is too swift for the lake-adapted fishes of the upper basin, it co-occurs with the endemic Klamath Smallscale Sucker,
<taxonomicName authority="(Moyle 2002)" baseAuthorityName="Moyle" baseAuthorityYear="2002" class="Actinopterygii" family="Catostomidae" genus="Catostomus" kingdom="Animalia" order="Cypriniformes" pageId="27" pageNumber="528" phylum="Chordata" rank="species" species="rimiculus">
<emphasis italics="true" pageId="27" pageNumber="528">Catostomus rimiculus</emphasis>
(
<bibRefCitation author="Moyle, P. B." box="[280,431,1591,1617]" pageId="27" pageNumber="528" refId="ref29646" refString="Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp." type="book" year="2002">Moyle 2002</bibRefCitation>
)
</taxonomicName>
. Given the genetic structure that Wiesenfeld
<emphasis box="[987,1050,1591,1617]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
(2017) found in the basin, it is possible that dace populations from the upper and lower river should be treated as separate subspecies.
<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." pageId="27" pageNumber="528" 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="[1379,1436,1627,1653]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
(2022)
</bibRefCitation>
confirmed results from other genetic methods that the Klamath Speckled Dace is a distinct evolutionary lineage.
</paragraph>
</subSubSection>
<subSubSection pageId="27" pageNumber="528" type="etymology">
<paragraph blockId="27.[151,1437,151,1941]" pageId="27" pageNumber="528">
<emphasis bold="true" box="[199,334,1735,1761]" pageId="27" pageNumber="528">Etymology.</emphasis>
The Klamath Speckled Dace is named for the river system to which it is endemic. The river is named for the native peoples who lived (and still do) in the upper Klamath Basin.
</paragraph>
<paragraph blockId="27.[151,1437,151,1941]" pageId="27" pageNumber="528">
<emphasis bold="true" box="[199,448,1806,1832]" pageId="27" pageNumber="528">Conservation Status.</emphasis>
The Klamath Speckled Dace is widespread and abundant in streams and natural lakes in the Klamath Basin, including the Trinity River. The main concern is that some distinct populations, such as in Jenny Creek, may be lost as the waters are dammed and diverted. The genetic diversity of Speckled Dace populations within the river system is just beginning to be appreciated (
<bibRefCitation author="Pfrender, M. E. &amp; Hicks, J. &amp; Lynch, M." box="[813,1042,1915,1941]" pageId="27" pageNumber="528" pagination="490 - 502" refId="ref30505" refString="Pfrender, M. E., Hicks, J. &amp; Lynch, M. (2004) Biogeographic patterns and current distribution of molecular-genetic variation among populations of Speckled Dace, Rhinichthys osculus (Girard). Molecular Phylogenetics and Evolution, 30 (3), 490 - 502. https: // doi. org / 10.1016 / S 1055 - 7903 (03) 00242 - 2" type="journal article" year="2004">
Pfrender
<emphasis box="[919,975,1915,1941]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
2004
</bibRefCitation>
; Wiesenfeld
<emphasis box="[1185,1242,1915,1941]" italics="true" pageId="27" pageNumber="528">et al.</emphasis>
2017).
</paragraph>
</subSubSection>
</treatment>
</document>