410 lines
45 KiB
XML
410 lines
45 KiB
XML
<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 & Su, Yingxin" docDate="2023" docId="03D1EC51DE0DFF813FFFFF16C873F980" 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" docType="treatment" docVersion="2" lastPageNumber="526" masterDocId="FFE89429DE15FF983F68FF81CD38FF89" masterDocTitle="Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA" masterLastPageNumber="539" masterPageNumber="501" pageNumber="525" 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:namePart>Moyle, Peter B.</mods:namePart>
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<mods:affiliation>Center for Watershed Sciences & 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>
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<mods:affiliation>California Department of Fish & 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:namePart>Su, Yingxin</mods:namePart>
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<mods:nameIdentifier type="email">pbmoyle@ucdavis.edu</mods:nameIdentifier>
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<mods:title>Zootaxa</mods:title>
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<mods:part>
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<mods:date>2023</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2023-03-06</mods:number>
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<mods:detail type="volume">
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<mods:number>5249</mods:number>
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<mods:number>5</mods:number>
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<mods:identifier type="ISSN">1175-5326</mods:identifier>
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<taxonomicName authority="(Evermann and Meek 1898)" baseAuthorityName="Evermann and Meek" baseAuthorityYear="1898" box="[151,823,151,178]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="species" species="klamathensis">
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<emphasis bold="true" box="[151,465,151,178]" italics="true" pageId="24" pageNumber="525">Rhinichthys klamathensis</emphasis>
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(
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<bibRefCitation author="Evermann, B. W. & Meek, S. B." box="[482,814,151,177]" pageId="24" pageNumber="525" pagination="15 - 84" refId="ref27663" refString="Evermann, B. W. & 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>
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)
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</taxonomicName>
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, Western Speckled Dace
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</emphasis>
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</heading>
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</paragraph>
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<figureCitation box="[151,220,187,213]" 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="24" pageNumber="525">Fig. 6</figureCitation>
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.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="24" pageNumber="525" type="reference_group">
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<paragraph blockId="24.[151,1436,256,372]" box="[151,842,256,280]" pageId="24" pageNumber="525">
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<treatmentCitation author="Evermann, B. W. & Meek, S. B." box="[151,670,256,280]" page="74" pageId="24" pageNumber="525" year="1898">
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<taxonomicName authority="Evermann and Meek 1898: 74" authorityName="Evermann and Meek" authorityPageNumber="74" authorityYear="1898" box="[151,670,256,280]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="species" species="klamathensis">
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<emphasis box="[151,360,256,280]" italics="true" pageId="24" pageNumber="525">Agosia klamathensis</emphasis>
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<bibRefCitation author="Evermann, B. W. & Meek, S. B." box="[367,670,256,280]" pageId="24" pageNumber="525" pagination="15 - 84" refId="ref27663" refString="Evermann, B. W. & 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 1898:74</bibRefCitation>
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</taxonomicName>
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;
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<treatmentCitation author="Snyder, J. O." box="[680,842,256,280]" page="98" pageId="24" pageNumber="525" year="1908">
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<bibRefCitation author="Snyder, J. O." box="[680,842,256,280]" pageId="24" pageNumber="525" 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>
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<paragraph blockId="24.[151,1436,256,372]" box="[151,684,286,310]" pageId="24" pageNumber="525">
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<treatmentCitation author="Evermann, B. W. & Clark, H. W." box="[151,684,286,310]" page="55" pageId="24" pageNumber="525" year="1931">
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<taxonomicName authority="Evermann and Clark 1931: 55" authorityName="Evermann and Clark" authorityPageNumber="55" authorityYear="1931" box="[151,684,286,310]" class="Actinopterygii" family="Cyprinidae" genus="Apocope" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="species" species="klamathensis">
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<emphasis box="[151,378,286,310]" italics="true" pageId="24" pageNumber="525">Apocope klamathensis</emphasis>
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<bibRefCitation author="Evermann, B. W. & Clark, H. W." box="[385,684,286,310]" pageId="24" pageNumber="525" pagination="1 - 67" refId="ref27727" refString="Evermann, B. W. & 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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<treatmentCitationGroup pageId="24" pageNumber="525">
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<treatmentCitation author="Shapovalov, L. & Dill, W. A." box="[151,828,317,341]" page="386" pageId="24" pageNumber="525" year="1950">
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<taxonomicName authority="Shapovalov and Dill 1950: 386" authorityName="Shapovalov and Dill" authorityPageNumber="386" authorityYear="1950" box="[151,828,317,341]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="subSpecies" species="osculus" subSpecies="klamathensis">
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<emphasis box="[151,497,317,341]" italics="true" pageId="24" pageNumber="525">Rhinichthys osculus klamathensis</emphasis>
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<bibRefCitation author="Shapovalov, L. & Dill, W. A." box="[507,828,317,341]" pageId="24" pageNumber="525" pagination="382 - 391" refId="ref31318" refString="Shapovalov, L. & Dill, W. A. (1950) A check list of the fresh-water and anadromous fishes of California. California Fish and Game, 36, 382 - 391" type="journal article" year="1950">Shapovalov and Dill 1950:386</bibRefCitation>
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;
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<treatmentCitation author="Hubbs, C. L. & Follett, W. I. & Dempster, L. J." box="[842,1066,317,341]" page="12" pageId="24" pageNumber="525" year="1979">
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<bibRefCitation author="Hubbs, C. L. & Follett, W. I. & Dempster, L. J." box="[842,1066,317,341]" pageId="24" pageNumber="525" pagination="1 - 54" refId="ref28578" refString="Hubbs, C. L., Follett, W. I. & Dempster, L. J. (1979) List of the fishes of California. Occasional Papers of the California Academy of Sciences, 133, 1 - 54." type="journal article" year="1979">
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Hubbs
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<emphasis box="[917,971,317,341]" italics="true" pageId="24" pageNumber="525">et al.</emphasis>
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1979:12
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;
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<treatmentCitation author="Shapovalov, L. & Cordone, A. J. & Dill, W. A." box="[1079,1357,317,341]" page="25" pageId="24" pageNumber="525" year="1981">
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<bibRefCitation author="Shapovalov, L. & Cordone, A. J. & Dill, W. A." box="[1079,1357,317,341]" pageId="24" pageNumber="525" pagination="4 - 38" refId="ref31398" refString="Shapovalov, L., Cordone, A. J. & Dill, W. A. (1981) A list of the freshwater and anadromous fishes of California. California Fish and Game, 67, 4 - 38." type="journal article" year="1981">
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||
Shapovalov
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<emphasis box="[1208,1262,317,341]" italics="true" pageId="24" pageNumber="525">et al.</emphasis>
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||
1981:25
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</bibRefCitation>
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</treatmentCitation>
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;
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||
<treatmentCitation author="Moyle, P. B." page="161" pageId="24" pageNumber="525" year="2002">
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<bibRefCitation author="Moyle, P. B." pageId="24" pageNumber="525" 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 2002:161</bibRefCitation>
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</treatmentCitation>
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;
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<treatmentCitation author="Markle, D. F." box="[308,471,347,372]" page="50" pageId="24" pageNumber="525" year="2016">
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<bibRefCitation author="Markle, D. F." box="[308,471,347,372]" pageId="24" pageNumber="525" 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>
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<subSubSection pageId="24" pageNumber="525" type="materials_examined">
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<paragraph blockId="24.[151,1437,418,2028]" pageId="24" pageNumber="525">
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<materialsCitation collectingDate="1896-11-03" collectionCode="R" country="United States of America" county="Seth Meek" location="Pelican Bay" municipality="Upper Klamath Lake" pageId="24" pageNumber="525" specimenCode="USNM 48225" specimenCount="1" stateProvince="Oregon" typeStatus="holotype">
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<emphasis bold="true" box="[151,260,418,444]" pageId="24" pageNumber="525">
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<typeStatus box="[151,260,418,444]" pageId="24" pageNumber="525">Holotype</typeStatus>
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</emphasis>
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||
:
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||
<specimenCode box="[274,430,418,445]" 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="24" pageNumber="525" type="Museum">USNM 48225</specimenCode>
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(
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SL) as
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<taxonomicName authority="Evermann and Meek 1898" authorityName="Evermann and Meek" authorityYear="1898" box="[605,1136,418,444]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="species" species="klamathensis">
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<emphasis box="[605,834,418,444]" italics="true" pageId="24" pageNumber="525">Agosia klamathensis</emphasis>
|
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<bibRefCitation author="Evermann, B. W. & Meek, S. B." box="[840,1136,418,444]" pageId="24" pageNumber="525" pagination="15 - 84" refId="ref27663" refString="Evermann, B. W. & 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>
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</taxonomicName>
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||
.
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||
<date box="[1145,1355,418,444]" pageId="24" pageNumber="525" value="1896-11-03">
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||
<collectingDate box="[1145,1355,418,444]" pageId="24" pageNumber="525" value="1896-11-03">November 3, 1896</collectingDate>
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</date>
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||
, mouth of small creek flowing into
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||
,
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||
<collectingMunicipality box="[608,848,454,480]" pageId="24" pageNumber="525">Upper Klamath Lake</collectingMunicipality>
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||
,
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||
<collectingRegion box="[859,946,454,480]" country="United States of America" name="Oregon" pageId="24" pageNumber="525">Oregon</collectingRegion>
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||
,
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||
,
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<collectingCounty box="[1026,1147,454,480]" pageId="24" pageNumber="525">Seth Meek</collectingCounty>
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||
and A.
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||
<collectionCode box="[1227,1249,454,480]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="24" pageNumber="525">R</collectionCode>
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. Alexander.
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||
</materialsCitation>
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</paragraph>
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<paragraph blockId="24.[151,1437,418,2028]" box="[199,1370,490,517]" pageId="24" pageNumber="525">
|
||
<materialsCitation box="[199,1367,490,517]" collectionCode="USNM" pageId="24" pageNumber="525" specimenCode="USNM 48231" specimenCount="77" typeStatus="paratype">
|
||
<emphasis bold="true" box="[199,319,490,516]" pageId="24" pageNumber="525">
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||
<typeStatus box="[199,319,490,516]" pageId="24" pageNumber="525">Paratypes</typeStatus>
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||
</emphasis>
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||
.
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||
<specimenCode box="[333,491,490,516]" 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="24" pageNumber="525" type="Museum">USNM 48231</specimenCode>
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||
(
|
||
<specimenCount box="[506,660,490,516]" pageId="24" pageNumber="525" type="generic">39 specimens</specimenCount>
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||
) and 125012 (
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||
<specimenCount box="[822,980,490,516]" pageId="24" pageNumber="525" type="generic">38 specimens</specimenCount>
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||
). Same date and place as
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||
<typeStatus box="[1265,1367,490,516]" pageId="24" pageNumber="525">holotype</typeStatus>
|
||
</materialsCitation>
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||
.
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||
</paragraph>
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||
</subSubSection>
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||
<subSubSection pageId="24" pageNumber="525" type="diagnosis">
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||
<paragraph blockId="24.[151,1437,418,2028]" pageId="24" pageNumber="525">
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||
<emphasis bold="true" box="[199,322,525,551]" pageId="24" pageNumber="525">Diagnosis.</emphasis>
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A cryptic species readily recognized as Speckled Dace by small adult size (
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||
<quantity box="[1229,1330,526,552]" metricMagnitude="-2" metricUnit="m" metricValue="8.5" metricValueMax="11.0" metricValueMin="6.0" pageId="24" pageNumber="525" unit="cm" value="8.5" valueMax="11.0" valueMin="6.0">6–11 cm</quantity>
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||
SL) and distinctive body shape. Dark blotches on sides merge into a line of dark pigmentation below eye, to snout. Meristics are in
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<tableCitation box="[217,300,598,624]" captionStart="TABLE 3" captionStartId="29.[152,245,151,175]" captionText="TABLE 3. Mean values of meristics of Amargosa Speckled Dace (from Amargosa River, Ash Meadows, and Owens Valley) compared to Long Valley and Lahontan Speckled Dace, from Sada et al. (1993). LLS, lateral line scales; LLP, lateral line scales with pores, PFR, pectoral-fin rays; PEFR, pelvic; Vert, vertebrae; Barbels, percent with maxillary barbels; STB canal, percent with a complete supratemporal canal; Frenum, percent with complete frenum. All counts are mean numbers." pageId="24" pageNumber="525">Table 3</tableCitation>
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. The best way to characterize this abundant fish is as a distinct Speckled Dace lineage found throughout the Klamath, Sacramento, and Warner watersheds with three sub-lineages that cannot be readily distinguished from one another.
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||
</paragraph>
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||
</subSubSection>
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||
<subSubSection pageId="24" pageNumber="525" type="description">
|
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<paragraph blockId="24.[151,1437,418,2028]" pageId="24" pageNumber="525">
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||
<emphasis bold="true" box="[199,337,705,731]" pageId="24" pageNumber="525">Description</emphasis>
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||
. According to
|
||
<bibRefCitation author="Evermann, B. W. & Meek, S. B." box="[506,888,706,733]" pageId="24" pageNumber="525" pagination="15 - 84" refId="ref27663" refString="Evermann, B. W. & 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 (1898:74-75</bibRefCitation>
|
||
,
|
||
<figureCitation box="[900,967,706,732]" 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="24" pageNumber="525">Fig. 6</figureCitation>
|
||
)
|
||
<emphasis box="[983,999,706,732]" italics="true" pageId="24" pageNumber="525">“</emphasis>
|
||
the fish from [Upper Klamath Lake]… possess certain characters which serve to distinguish it from
|
||
<taxonomicName box="[849,954,742,768]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="species" species="nubila">
|
||
<emphasis box="[849,954,742,768]" italics="true" pageId="24" pageNumber="525">A. nubila</emphasis>
|
||
</taxonomicName>
|
||
. While the differences are slight, they are plainly evident and must be recognized, and we therefore describe it as a new species (p. 74). Head 4 [times into standard length]; depth 4.5; eye
|
||
<quantity box="[513,577,814,841]" metricMagnitude="-1" metricUnit="m" metricValue="1.143" pageId="24" pageNumber="525" unit="in" value="4.5">4.5 in</quantity>
|
||
head; D[orsal rays]i.8; A [anal rays] i,7; [lateral line] scales about 14–78–10 (average,
|
||
<quantity box="[262,319,850,876]" metricMagnitude="0" metricUnit="m" metricValue="1.8542" pageId="24" pageNumber="525" unit="in" value="73.0">73 in</quantity>
|
||
<specimenCount box="[325,484,850,876]" pageId="24" pageNumber="525" type="generic">49 specimens</specimenCount>
|
||
).”
|
||
</paragraph>
|
||
<paragraph blockId="24.[151,1437,418,2028]" pageId="24" pageNumber="525">
|
||
Body robust, subcylindrical, back somewhat elevated; snout rather long, mouth inferior, little oblique, the lower jaw included; maxillary not reaching front of orbit; upper lip without frenum; barbels present but small. Lateral line incomplete, interrupted in many places, about 30 pores developed. Origin of dorsal fin midway between front of pupil and base of caudal fin; pectorals rather short, reaching about three-fourth the distance to ventrals; ventrals reaching vent; anal[fin] large, its longest ray
|
||
<quantity box="[655,719,1030,1057]" metricMagnitude="-2" metricUnit="m" metricValue="3.81" pageId="24" pageNumber="525" unit="in" value="1.5">1.5 in</quantity>
|
||
head.
|
||
</paragraph>
|
||
<paragraph blockId="24.[151,1437,418,2028]" pageId="24" pageNumber="525">
|
||
Color in alcohol, olivaceous, mottled and blotched with darker on back and sides; under parts pale; an obscure pale streak from eye to base of caudal fin, below which is a broad dark band; dorsals, pectorals, caudal dusky; other fins plain; a black blotch at base of caudal. This form is distinguished from
|
||
<taxonomicName box="[924,1075,1138,1164]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="species" species="nubila">
|
||
<emphasis box="[924,1075,1138,1164]" italics="true" pageId="24" pageNumber="525">Agosia nubila</emphasis>
|
||
</taxonomicName>
|
||
chiefly by the smaller size of the scales.
|
||
<taxonomicName box="[230,329,1174,1200]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="species" species="nubila">
|
||
<emphasis box="[230,329,1174,1200]" italics="true" pageId="24" pageNumber="525">A. nubila</emphasis>
|
||
</taxonomicName>
|
||
has 49–66 lateral line scales (mean 57, n=73), while
|
||
<taxonomicName authorityName="Evermann and Meek" authorityYear="1898" box="[909,1081,1174,1200]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="species" species="klamathensis">
|
||
<emphasis box="[909,1081,1174,1200]" italics="true" pageId="24" pageNumber="525">A. klamathensis</emphasis>
|
||
</taxonomicName>
|
||
has 68–78
|
||
<emphasis box="[1205,1307,1174,1200]" italics="true" pageId="24" pageNumber="525">(mean 73</emphasis>
|
||
,
|
||
<emphasis box="[1321,1395,1174,1200]" italics="true" pageId="24" pageNumber="525">n =49)</emphasis>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="24.[151,1437,418,2028]" pageId="24" pageNumber="525">
|
||
Overall, there are only minor (statistical) differences among the three subspecies from this basic description. However,
|
||
<bibRefCitation author="Markle, D. F." box="[270,435,1246,1272]" pageId="24" pageNumber="525" 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>
|
||
used lateral line scale number as the principal character to separate Western Speckled Dace from other dace taxa in
|
||
<collectingRegion box="[482,570,1282,1308]" country="United States of America" name="Oregon" pageId="24" pageNumber="525">Oregon</collectingRegion>
|
||
. In contrast,
|
||
<bibRefCitation author="Smith, G. R. & Chow, J. & Unmack, P. J. & Markle, D. F. & Dowling, T. E." box="[713,925,1282,1308]" pageId="24" pageNumber="525" pagination="1 - 83" refId="ref31441" refString="Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. & 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="[787,843,1282,1308]" italics="true" pageId="24" pageNumber="525">et al.</emphasis>
|
||
(2017)
|
||
</bibRefCitation>
|
||
could not readily distinguish lineages despite multivariate analyses of combined meristic, morphological, mtDNA, and fossil data from the entire region in which Speckled Dace
|
||
<emphasis box="[325,439,1354,1380]" italics="true" pageId="24" pageNumber="525">sensu lato</emphasis>
|
||
occur.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="25" lastPageNumber="526" pageId="24" pageNumber="525" type="distribution">
|
||
<paragraph blockId="24.[151,1437,418,2028]" pageId="24" pageNumber="525">
|
||
<emphasis bold="true" box="[199,345,1389,1415]" pageId="24" pageNumber="525">Distribution</emphasis>
|
||
. The Western Speckled Dace is found throughout the Klamath and Sacramento watersheds, including streams as far south as the Santa Maria River in southern
|
||
<collectingRegion box="[900,1017,1426,1452]" country="United States of America" name="California" pageId="24" pageNumber="525">California</collectingRegion>
|
||
. Its distribution in the Warner basin is mostly in springs and small cool-water streams. Only the headwaters of Twelvemile Creek are in
|
||
<collectingRegion box="[1228,1342,1462,1488]" country="United States of America" name="California" pageId="24" pageNumber="525">California</collectingRegion>
|
||
(Modoc County). Before the introduction of non-native game fishes, it probably had a wider distribution within the basin. See the subspecies accounts for more detailed distributional information.
|
||
</paragraph>
|
||
<paragraph blockId="24.[151,1437,418,2028]" pageId="24" pageNumber="525">
|
||
<emphasis bold="true" box="[199,369,1569,1596]" pageId="24" pageNumber="525">Note: Geology</emphasis>
|
||
. The Klamath River has not been connected to the ancestral Sacramento River system since the end of the Pliocene (
|
||
<emphasis box="[389,423,1607,1632]" italics="true" pageId="24" pageNumber="525">ca.</emphasis>
|
||
three mya); however, extensive deformation (
|
||
<emphasis box="[949,982,1607,1632]" italics="true" pageId="24" pageNumber="525">e.g</emphasis>
|
||
. down-faulting of the Klamath Graben) and vulcanism (both from the Cascade-arc and Medicine Lake volcanic fields) occurred almost continuously in the northern Sacramento and southern Klamath Basins (
|
||
<bibRefCitation author="Colman, S. M. & Platt Bradbury, J. & Rosenbaum, J. G." box="[832,1063,1678,1704]" pageId="24" pageNumber="525" pagination="129 - 138" refId="ref27068" refString="Colman, S. M., Platt Bradbury, J. & Rosenbaum, J. G. (2004) Paleolimnology and paleoclimate studies in Upper Klamath Lake, Oregon. Journal of Paleolimnology, 31 (2), 129 - 138. https: // doi. org / 10.1023 / B: JOPL. 0000019235.72107.92" type="journal article" year="2004">
|
||
Colman
|
||
<emphasis box="[933,992,1678,1704]" italics="true" pageId="24" pageNumber="525">et al.</emphasis>
|
||
2004
|
||
</bibRefCitation>
|
||
). It is probable that this activity led to repeated drainage captures of between the two basins and allowed for intermittent gene flow among dace populations. Inter-basin connectivity presumably was less frequent between Klamath/Sacramento streams and those in the Warner Basin. The development of the Great Basin faulting and extension belt of down-faulting resulted in creation of the Warner Range (
|
||
<emphasis box="[537,571,1823,1848]" italics="true" pageId="24" pageNumber="525">ca.</emphasis>
|
||
3 mya) which resulted in a permanent topographic separation of the basins sometime in the late-Pleistocene (1.0–0.1 mya) (
|
||
<bibRefCitation author="Egger, A. E. & Miller, E. L." box="[694,892,1858,1884]" pageId="24" pageNumber="525" pagination="756 - 773" refId="ref27599" refString="Egger, A. E. & Miller, E. L. (2011) Evolution of the northwestern margin of the Basin and Range: The geology and extensional history of the Warner Range and environs, northeastern California. Geosphere, 7 (3), 756 - 773. https: // doi. org / 10.1130 / GES 00620.1" type="journal article" year="2011">
|
||
Egger
|
||
<emphasis box="[769,826,1858,1884]" italics="true" pageId="24" pageNumber="525">et al.</emphasis>
|
||
2011
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="24.[151,1437,418,2028]" lastBlockId="25.[151,1437,151,1545]" lastPageId="25" lastPageNumber="526" pageId="24" pageNumber="525">
|
||
As the result of these active geologic processes, each of the three major subspecies of
|
||
<taxonomicName authorityName="Evermann and Meek" authorityYear="1898" box="[1215,1395,1894,1920]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="24" pageNumber="525" phylum="Chordata" rank="species" species="klamathensis">
|
||
<emphasis box="[1215,1395,1894,1920]" italics="true" pageId="24" pageNumber="525">R. klamathensis</emphasis>
|
||
</taxonomicName>
|
||
, as discussed here, is endemic to a single large watershed, plus some coastal watersheds in the case of Sacramento Speckled Dace. They share these waters with other endemic fishes. Dace ultimately colonized these three watersheds from the ancient Columbia River Basin during a series of volcanic and other geologic events that caused the predecessor of the upper Snake River to flow into the Pacific Ocean via the Klamath-Sacramento region, apparently during the Pliocene period. Fish fossils from ancient deposits in the
|
||
<collectingRegion box="[914,1059,187,213]" country="United States of America" name="Idaho" pageId="25" pageNumber="526">Idaho region</collectingRegion>
|
||
, that predate the rise of the Sierra
|
||
<collectingRegion box="[151,237,223,249]" country="United States of America" name="Nevada" pageId="25" pageNumber="526">Nevada</collectingRegion>
|
||
and coastal mountain ranges, indicate a source of the predecessors of the highly distinctive endemic fish fauna of the lower Sacramento-San Joaquin Watershed (
|
||
<bibRefCitation author="Minckley, W. L. & Hendrickson, D. A. & Bond, C. E." box="[787,1030,259,285]" pageId="25" pageNumber="526" pagination="519 - 614" refId="ref29499" refString="Minckley, W. L., Hendrickson, D. A. & Bond, C. E. (1986) Geography of western North America freshwater fishes: Description and relationship to intercontinental tectonism. In: Hocutt, C. H. & 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="[905,962,259,285]" italics="true" pageId="25" pageNumber="526">et al.</emphasis>
|
||
1986
|
||
</bibRefCitation>
|
||
). It is likely that the Speckled Dace was not part of those early colonization events but arrived later, during a period when volcanic activity caused the upper Pit River to switch from flowing into the Sacramento River to flowing into the Klamath instead, which had its own endemic fish fauna. Some Klamath fishes (
|
||
<emphasis box="[735,775,367,392]" italics="true" pageId="25" pageNumber="526">e.g.</emphasis>
|
||
<taxonomicName box="[784,911,367,393]" class="Actinopterygii" family="Cottidae" genus="Cottus" kingdom="Animalia" order="Scorpaeniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="undetermined">
|
||
<emphasis box="[784,857,367,393]" italics="true" pageId="25" pageNumber="526">Cottus</emphasis>
|
||
spp.
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Girard" baseAuthorityYear="1856" box="[927,1133,367,393]" class="Actinopterygii" family="Cyprinidae" genus="Siphatales" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="bicolor">
|
||
<emphasis box="[927,1133,367,393]" italics="true" pageId="25" pageNumber="526">Siphatales bicolor</emphasis>
|
||
</taxonomicName>
|
||
) were able to colonize the upper Pit watershed as a result. The ancestral Pit River then broke through the volcanic dike barrier and rejoined the Sacramento River system, bringing in its water members of the Klamath fish fauna. Alternately,
|
||
<bibRefCitation author="Smith, G. R. & Chow, J. & Unmack, P. J. & Markle, D. F. & Dowling, T. E." box="[1226,1436,439,465]" pageId="25" pageNumber="526" pagination="1 - 83" refId="ref31441" refString="Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. & 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="[1299,1355,439,465]" italics="true" pageId="25" pageNumber="526">et al.</emphasis>
|
||
(2017)
|
||
</bibRefCitation>
|
||
suggest colonization of the Sacramento system could have come from a pass through the southern Sierra
|
||
<collectingRegion box="[1351,1437,475,501]" country="United States of America" name="Nevada" pageId="25" pageNumber="526">Nevada</collectingRegion>
|
||
that apparently was open 3.2 million years ago or earlier. However, the genomic similarities between Sacramento and Klamath daces and between them and Warner Speckled Dace are presumably the result of their ancestors colonizing the region during a long geologically active period in the Klamath-Pit region. The genomic study of
|
||
<bibRefCitation author="Su, Y. & Moyle, P. B. & Campbell, M. A. & Finger, A. J. & O'Rourke, S. & Baumsteiger, J. & Miller, M. R." pageId="25" pageNumber="526" pagination="695 - 710" refId="ref32073" refString="Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & 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="[151,207,619,645]" italics="true" pageId="25" pageNumber="526">et al.</emphasis>
|
||
(2022)
|
||
</bibRefCitation>
|
||
supports this view (
|
||
<figureCitation box="[514,611,619,645]" captionStart="FIGURE 2" captionStartId="11.[152,255,1729,1753]" captionTargetBox="[225,1366,187,1692]" captionTargetId="figure-18@11.[221,1366,181,1705]" captionTargetPageId="11" captionText="FIGURE 2. Phylogenetic relationships among Speckled Dace populations in this study, modified from Su et al (2022). The molecular phylogeny was generated by an ML coalescent model in SVDQuartets with 100 bootstraps, using RAD sequencing data. Numbers on each lineage refer to the percentage of bootstrap runs (out of 100) that produced the same result. Details can be found in Su et al. (2022). The lineage names refer to major drainages from which our Speckled Dace samples were collected from California, as well as from major basins outside of California. We did not examine the latter findings in detail; they probably represent multiple undescribed species/subspecies. The capital letters refer to lineages identified as species in this paper. A. Desert Speckled Dace (R. nevadensis), B. Western Speckled Dace (R. klamathensis), C. Santa Ana Speckled Dace (R. gabrielino). Tui Chub (Siphatales bicolor) and Relict Dace (Relictus solitarius) were outgroups used to root the phylogeny because they are both western cyprinids distantly related to Rhinichthys (Schönhuth et al. 2012)." figureDoi="http://doi.org/10.5281/zenodo.7701361" httpUri="https://zenodo.org/record/7701361/files/figure.png" pageId="25" pageNumber="526">Figure 2</figureCitation>
|
||
) and that the Western Speckled Dace merits full species status.
|
||
</paragraph>
|
||
<paragraph blockId="25.[151,1437,151,1545]" pageId="25" pageNumber="526">
|
||
<emphasis bold="true" box="[199,365,654,681]" pageId="25" pageNumber="526">Note: History</emphasis>
|
||
. The three lineages that make up this species have different taxonomic histories (
|
||
<bibRefCitation author="Evermann, B. W. & Clark, H. W." pageId="25" pageNumber="526" pagination="1 - 67" refId="ref27727" refString="Evermann, B. W. & 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</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Moyle, P. B." box="[338,479,691,717]" pageId="25" pageNumber="526" 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 was recognized in 1898 as a species, and it retained its identity even when subsumed as a subspecies under
|
||
<emphasis box="[730,852,727,753]" italics="true" pageId="25" pageNumber="526">
|
||
<taxonomicName baseAuthorityName="Miller" baseAuthorityYear="1973" box="[730,847,727,753]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="osculus">R. osculus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The Sacramento Speckled Dace was largely ignored as a taxon despite its abundance and wide distribution. Presumably early workers such as Cloudsley Rutter, Charles Gilbert, and John Otterbein Snyder saw that it could not be readily distinguished from other Speckled Dace and accepted by default the use of an improper name (some variant of
|
||
<taxonomicName authorityName=": Snyder" authorityYear="1913" box="[888,1100,835,861]" class="Actinopterygii" family="Cyprinidae" genus="Agosia" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="carringtoni">
|
||
<emphasis box="[888,1100,835,861]" italics="true" pageId="25" pageNumber="526">Agosia carringtoni</emphasis>
|
||
</taxonomicName>
|
||
) or else treated it as Lahontan Speckled Dace. The Warner Speckled Dace came to our attention because of controversy over the status of the population in remote Foskett Spring, which modern genetic analysis showed to be part of the Warner Speckled Dace lineage described in this paper.
|
||
<bibRefCitation author="Su, Y. & Moyle, P. B. & Campbell, M. A. & Finger, A. J. & O'Rourke, S. & Baumsteiger, J. & Miller, M. R." box="[502,675,943,969]" pageId="25" pageNumber="526" pagination="695 - 710" refId="ref32073" refString="Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & 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="[538,594,943,969]" italics="true" pageId="25" pageNumber="526">et al.</emphasis>
|
||
(2022)
|
||
</bibRefCitation>
|
||
show that, based on genomics, the three lineages are similar enough so that together they can be treated as a single species,
|
||
<emphasis box="[767,1060,979,1005]" italics="true" pageId="25" pageNumber="526">
|
||
<taxonomicName box="[767,1056,979,1005]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="klamathensis">Rhinichthys klamathensis</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="25.[151,1437,151,1545]" pageId="25" pageNumber="526">
|
||
Nevertheless, all three populations are isolated from one another, with long independent histories. The early history of each basin includes one or more periods of ancient interconnectedness which allowed Speckled Dace to colonize all three basins. The three lineages co-occur with other endemic fishes in their respective basins. The endemic fishes in each basin have close relatives in the other basins (
|
||
<emphasis box="[949,982,1123,1148]" italics="true" pageId="25" pageNumber="526">e.g</emphasis>
|
||
.
|
||
<emphasis box="[998,1299,1123,1149]" italics="true" pageId="25" pageNumber="526">
|
||
<taxonomicName box="[998,1294,1123,1149]" class="Actinopterygii" family="Cyprinidae" genus="Siphatales" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="subSpecies" species="bicolor" subSpecies="subspp">Siphatales bicolor subspp</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
), as well as species unique to each basin.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="25" pageNumber="526" type="etymology">
|
||
<paragraph blockId="25.[151,1437,151,1545]" pageId="25" pageNumber="526">
|
||
<emphasis bold="true" box="[199,334,1195,1221]" pageId="25" pageNumber="526">Etymology.</emphasis>
|
||
The name ‘Western Speckled Dace’ (
|
||
<bibRefCitation author="Markle, D. F." box="[741,888,1195,1221]" pageId="25" pageNumber="526" 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>
|
||
,
|
||
<bibRefCitation author="Smith, G. R. & Chow, J. & Unmack, P. J. & Markle, D. F. & Dowling, T. E." box="[897,1085,1195,1221]" pageId="25" pageNumber="526" pagination="1 - 83" refId="ref31441" refString="Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. & 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="[968,1022,1195,1221]" italics="true" pageId="25" pageNumber="526">et al.</emphasis>
|
||
2017
|
||
</bibRefCitation>
|
||
) covers three lineages: Klamath Speckled Dace (
|
||
<emphasis box="[334,537,1231,1257]" italics="true" pageId="25" pageNumber="526">
|
||
R. k.
|
||
<taxonomicName box="[391,537,1231,1257]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="klamathensis">klamathensis</taxonomicName>
|
||
</emphasis>
|
||
), Sacramento Speckled Dace (
|
||
<emphasis box="[881,1053,1231,1257]" italics="true" pageId="25" pageNumber="526">R. k. achomawi</emphasis>
|
||
) and Warner Speckled Dace (
|
||
<emphasis italics="true" pageId="25" pageNumber="526">R. k. goyatoka</emphasis>
|
||
). Only the Klamath Speckled Dace has been previously named, so
|
||
<taxonomicName box="[1021,1199,1267,1293]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" kingdom="Animalia" order="Cypriniformes" pageId="25" pageNumber="526" phylum="Chordata" rank="species" species="klamathensis">
|
||
<emphasis box="[1021,1199,1267,1293]" italics="true" pageId="25" pageNumber="526">R. klamathensis</emphasis>
|
||
</taxonomicName>
|
||
becomes the species epithet for three subspecies. Klamath is an anglicized version of a name for the native peoples inhabiting the upper Klamath Lake region in
|
||
<collectingRegion box="[425,509,1339,1365]" country="United States of America" name="Oregon" pageId="25" pageNumber="526">Oregon</collectingRegion>
|
||
(
|
||
<bibRefCitation author="Gudde, E. G. & Bright, W." box="[523,793,1339,1365]" pageId="25" pageNumber="526" refId="ref28383" refString="Gudde, E. G. & Bright, W. (1998) California Place Names. University of California Press, Berkeley, California, xxviii + 467 pp. https: // doi. org / 10.1525 / 9780520920545" type="book" year="1998">Gudde and Bright 1998</bibRefCitation>
|
||
). See subspecies accounts for further explanations.
|
||
</paragraph>
|
||
<paragraph blockId="25.[151,1437,151,1545]" pageId="25" pageNumber="526">
|
||
<emphasis bold="true" box="[199,439,1374,1400]" pageId="25" pageNumber="526">Conservation Status</emphasis>
|
||
. Sacramento and Klamath Speckled Dace are wide-spread and abundant in the large basins they inhabit. Warner Speckled Dace, including the population in Foskett Spring, are limited by the low availability of water in the desert Warner Basin. Their populations for now seem reasonably secure, although the basin has been invaded by non-native species. The Foskett Spring population was listed in 1985 as a Threatened species under the federal Endangered Species Act but it was delisted in 2019, following implementation of the recovery plan.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |