179 lines
23 KiB
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179 lines
23 KiB
XML
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<mods:title id="83F9B5A4AAFC2F2832D01AE1A5E32928">A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex</mods:title>
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<treatment id="03816E226F00CC0DFF7FDF29FC65532C" ID-DOI="http://doi.org/10.5281/zenodo.5936937" ID-GBIF-Taxon="154529796" ID-Zenodo-Dep="5936937" LSID="urn:lsid:plazi:treatment:03816E226F00CC0DFF7FDF29FC65532C" httpUri="http://treatment.plazi.org/id/03816E226F00CC0DFF7FDF29FC65532C" lastPageId="3" lastPageNumber="224" pageId="2" pageNumber="223">
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<paragraph id="8B97DF346F00CC0CFF7FDF29FE3D550B" blockId="2.[151,484,1304,1330]" box="[151,484,1304,1330]" pageId="2" pageNumber="223">
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<heading id="D0DF68586F00CC0CFF7FDF29FE3D550B" bold="true" box="[151,484,1304,1330]" fontSize="11" level="1" pageId="2" pageNumber="223" reason="1">
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<emphasis id="B95C03266F00CC0CFF7FDF29FE3D550B" bold="true" box="[151,484,1304,1330]" pageId="2" pageNumber="223">
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<taxonomicName id="4C28A4B76F00CC0CFF7FDF29FF2D550B" ID-CoL="5BJH" ID-ENA="71764" box="[151,244,1304,1330]" class="Actinopterygii" family="Cyprinidae" genus="Lavinia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="2" pageNumber="223" phylum="Chordata" rank="genus">
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<emphasis id="B95C03266F00CC0CFF7FDF29FF2D550B" bold="true" box="[151,244,1304,1330]" italics="true" pageId="2" pageNumber="223">Lavinia</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="4C28A4B76F00CC0CFED9DF29FE3D550B" authorityName="Snyder" authorityYear="1913" box="[305,484,1304,1330]" class="Actinopterygii" family="Cyprinidae" genus="Hesperoleucus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="2" pageNumber="223" phylum="Chordata" rank="genus">
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<emphasis id="B95C03266F00CC0CFED9DF29FE3D550B" bold="true" box="[305,484,1304,1330]" italics="true" pageId="2" pageNumber="223">Hesperoleucus</emphasis>
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</taxonomicName>
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</emphasis>
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</heading>
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</paragraph>
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<paragraph id="8B97DF346F00CC0CFF7FDF6EFF3956DB" blockId="2.[151,1437,1374,2014]" pageId="2" pageNumber="223">
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One conundrum within the Hitch–Roach species complex is uncertainty regarding whether species should be placed in one genus or two genera. While the complex was originally divided between
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<taxonomicName id="4C28A4B76F00CC0CFB8CDFB2FB6055A3" box="[1124,1209,1411,1434]" class="Actinopterygii" family="Cyprinidae" genus="Lavinia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="2" pageNumber="223" phylum="Chordata" rank="genus">
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<emphasis id="B95C03266F00CC0CFB8CDFB2FB6055A3" box="[1124,1209,1411,1434]" italics="true" pageId="2" pageNumber="223">Lavinia</emphasis>
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</taxonomicName>
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and
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<emphasis id="B95C03266F00CC0CFB07DFB2FA4555A2" box="[1263,1436,1411,1435]" italics="true" pageId="2" pageNumber="223">
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<taxonomicName id="4C28A4B76F00CC0CFB07DFB2FA4E55A2" authorityName="Snyder" authorityYear="1913" box="[1263,1431,1411,1435]" class="Actinopterygii" family="Cyprinidae" genus="Hesperoleucus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="2" pageNumber="223" phylum="Chordata" rank="genus">Hesperoleucus</taxonomicName>
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,
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</emphasis>
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multiple examples of hybridization between the genera suggest they are more closely related than separate genera designations would normally indicate. For example, CA Roach are known to hybridize with Hitch in the Pajaro and Salinas rivers in Monterey Bay drainages (
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<bibRefCitation id="EFB9A2C56F00CC0CFD7ADFDFFCF8563E" author="Miller, R. R." box="[658,801,1518,1543]" pageId="2" pageNumber="223" pagination="197 - 204" refId="ref14142" refString="Miller, R. R. (1945) The status of Lavinia ardesiaca, a cyprinid fish from the Pajaro-Salinas River basin, California. Copeia, 1945, 197 - 204." type="journal article" year="1945">Miller 1945</bibRefCitation>
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), in Coyote, Alameda, and Walnut Creeks in the San Francisco Bay region (
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<bibRefCitation id="EFB9A2C56F00CC0CFE75DC23FDF35612" author="Miller, R. R." box="[413,554,1554,1579]" pageId="2" pageNumber="223" pagination="20 - 29" refId="ref14176" refString="Miller, R. R. (1963) Synonymy, characters, and variation of Gila crassicauda, a rare Californian minnow, with an account of its hybridization with Lavinia exilicauda. California Fish and Game, 49, 20 - 29." type="journal article" year="1963">Miller 1963</bibRefCitation>
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;
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<bibRefCitation id="EFB9A2C56F00CC0CFDD1DC23FD665612" author="Leidy, R. A." box="[569,703,1554,1579]" pageId="2" pageNumber="223" pagination="1 - 175" refId="ref13995" refString="Leidy, R. A. (1984) Distribution and ecology of stream fishes in the San Francisco Bay drainage, Hilgardia, 52 (8), 1 - 175." type="journal article" year="1984">Leidy 1984</bibRefCitation>
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,
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<bibRefCitation id="EFB9A2C56F00CC0CFD25DC22FCD15612" author="Leidy, R. A." box="[717,776,1555,1579]" pageId="2" pageNumber="223" refId="ref14028" refString="Leidy, R. A. (2007) Ecology, Assemblage Structure, Distribution, and Status of Fishes in Streams Tributary to the San Francisco Estuary, California. San Francisco Estuary Institute, San Francisco. 194 pp." type="book" year="2007">2007</bibRefCitation>
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, pers. comm.), and in Sacramento-San Joaquin drainages (
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<bibRefCitation id="EFB9A2C56F00CC0CFF76DC07FEBE5676" author="Avise, J. C. & Smith, J. J. & Ayala, F. J." box="[158,359,1590,1615]" pageId="2" pageNumber="223" pagination="411 - 426" refId="ref12641" refString="Avise, J. C., Smith, J. J. & Ayala, F. J. (1975) Adaptive differentiation with little genic change between two native California minnows. Evolution, 29, 411 - 426." type="journal article" year="1975">
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Avise
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<emphasis id="B95C03266F00CC0CFF00DC09FEFA5676" box="[232,291,1591,1615]" italics="true" pageId="2" pageNumber="223">et al.</emphasis>
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1975
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</bibRefCitation>
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;
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<bibRefCitation id="EFB9A2C56F00CC0CFE9CDC09FE215676" author="Jones, W. J." box="[372,504,1591,1615]" pageId="2" pageNumber="223" refId="ref13755" refString="Jones, W. J. (2001) DNA Sequence Divergence and Speciation in Two California Minnows (Cyprinidae: Lavinia exilicauda and L. symmetricus). PhD Dissertation, University of California, Santa Cruz. Ca. 150 pp." type="book" year="2001">Jones 2001</bibRefCitation>
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). Hybrids were fertile in the Pajaro River;
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<bibRefCitation id="EFB9A2C56F00CC0CFC0EDC07FB625676" author="Avise, J. C. & Smith, J. J. & Ayala, F. J." box="[998,1211,1590,1615]" pageId="2" pageNumber="223" pagination="411 - 426" refId="ref12641" refString="Avise, J. C., Smith, J. J. & Ayala, F. J. (1975) Adaptive differentiation with little genic change between two native California minnows. Evolution, 29, 411 - 426." type="journal article" year="1975">
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Avise
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<emphasis id="B95C03266F00CC0CFBC6DC09FBB15676" box="[1070,1128,1591,1615]" italics="true" pageId="2" pageNumber="223">et al.</emphasis>
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(1975)
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</bibRefCitation>
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found 8% of Hitch examined to be F
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<subScript id="17ACDD716F00CC0CFE8BDC5BFEB25641" attach="left" box="[355,363,1642,1656]" fontSize="6" pageId="2" pageNumber="223">1</subScript>
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hybrids and 5% to be backcrossed. Since it is unusual for two genera to be interfertile and because other endemic species in
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<collectingRegion id="49EC11D66F00CC0CFDC9DCB0FD4A56A3" box="[545,659,1665,1690]" country="United States of America" name="California" pageId="2" pageNumber="223">California</collectingRegion>
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do not appear to exhibit this behavior (
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<bibRefCitation id="EFB9A2C56F00CC0CFB8BDCB0FAF756A3" author="Avise, J. C. & Smith, J. J. & Ayala, F. J." box="[1123,1326,1665,1690]" pageId="2" pageNumber="223" pagination="411 - 426" refId="ref12641" refString="Avise, J. C., Smith, J. J. & Ayala, F. J. (1975) Adaptive differentiation with little genic change between two native California minnows. Evolution, 29, 411 - 426." type="journal article" year="1975">
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Avise
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<emphasis id="B95C03266F00CC0CFB46DCB3FB3056A0" box="[1198,1257,1665,1689]" italics="true" pageId="2" pageNumber="223">et al.</emphasis>
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1975
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</bibRefCitation>
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;
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<bibRefCitation id="EFB9A2C56F00CC0CFAD5DCB0FEC35687" author="Avise, J. C. & Ayala, F. J." pageId="2" pageNumber="223" pagination="46 - 58" refId="ref12603" refString="Avise, J. C. & Ayala, F. J. (1976) Genetic differentiation in speciose versus depauperate phylads: evidence from the California minnows. Evolution, 30, 46 - 58." type="journal article" year="1976">Avise & Ayala 1976</bibRefCitation>
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;
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<bibRefCitation id="EFB9A2C56F00CC0CFECFDC94FD965687" author="Moyle, P. B. & Massingill, M." box="[295,591,1701,1726]" pageId="2" pageNumber="223" pagination="196 - 198" refId="ref14567" refString="Moyle, P. B. & Massingill, M. (1981) Hybridization between hitch, Lavinia exilicauda, and Sacramento blackfish, Orthodon microlepidotus, in San Luis Reservoir, California. California Fish and Game, 67, 196 - 198." type="journal article" year="1981">Moyle & Massingill 1981</bibRefCitation>
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;
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<bibRefCitation id="EFB9A2C56F00CC0CFDB3DC94FD315687" author="Moyle, P. B." box="[603,744,1701,1726]" pageId="2" pageNumber="223" refId="ref14221" refString="Moyle, P. B. (2002) Inland Fishes of California. University of California Press, Berkeley, California. 502 pp." type="book" year="2002">Moyle 2002</bibRefCitation>
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), it has been proposed these two species belonged in a single genus.
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</paragraph>
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</subSubSection>
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<subSubSection id="C3328CBF6F00CC0DFF2FDCDCFC65532C" lastPageId="3" lastPageNumber="224" pageId="2" pageNumber="223" type="description">
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<paragraph id="8B97DF346F00CC0DFF2FDCDCFDD050CE" blockId="2.[151,1437,1374,2014]" lastBlockId="3.[151,1437,151,789]" lastPageId="3" lastPageNumber="224" pageId="2" pageNumber="223">
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<bibRefCitation id="EFB9A2C56F00CC0CFF2FDCDCFE0B573F" author="Aguilar, A. & Jones, W. J." box="[199,466,1773,1798]" pageId="2" pageNumber="223" pagination="373 - 381" refId="ref12469" refString="Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381." type="journal article" year="2009">Aguilar & Jones (2009)</bibRefCitation>
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found the two genera are closely related mtDNA lineages that could be considered to be one genus.
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<bibRefCitation id="EFB9A2C56F00CC0CFEACDD20FD855713" author="Schonhuth, S. & Hillis, D. M. & Neely, D. A. & Lozano-Vilano, L. & Perdices, A. & Mayden, R. L." box="[324,604,1809,1834]" pageId="2" pageNumber="223" pagination="427 - 446" refId="ref15257" refString="Schonhuth, S., Hillis, D. M., Neely, D. A., Lozano-Vilano, L., Perdices, A. & Mayden, R. L. (2012) Phylogeny, diversity, and species delimitation of the North American round-nosed minnows (Teleostei: Dionda), as inferred from mitochondrial and nuclear DNA sequences. Molecular Phylogenetics and Evolution, 62, 427 - 446." type="journal article" year="2012">
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Schönhuth
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<emphasis id="B95C03266F00CC0CFE2FDD23FDDA5710" box="[455,515,1809,1833]" italics="true" pageId="2" pageNumber="223">et al.</emphasis>
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(2012)
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</bibRefCitation>
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, however, noted that while their mtDNA analyses supported this close relationship, their limited nuclear DNA results did not. Our recent study (
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<bibRefCitation id="EFB9A2C56F00CC0CFC16DD04FAC35777" author="Baumsteiger, J. & Moyle, P. B. & Aguilar, A. & O'Rourke, S. M. & Miller, M. R." box="[1022,1306,1845,1870]" pageId="2" pageNumber="223" pagination="0189417" refId="ref12892" refString="Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417." type="journal article" year="2017">
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Baumsteiger
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<emphasis id="B95C03266F00CC0CFB70DD07FB0C5774" box="[1176,1237,1845,1869]" italics="true" pageId="2" pageNumber="223">et al.</emphasis>
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2017
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</bibRefCitation>
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) found the strongest genomic signal coincided with splitting samples into two genera. We discovered, however, that samples from the Pit River (Northern Roach) were a confounding factor, having mixed historical ancestry with both genera. It was estimated in one analysis that approximately 80% of the Northern Roach genome clustered with Hitch whereas only 20% clustered with CA Roach. But all subsequent analyses found Northern Roach to be a distinct species, on par with currently identified species. Thus Northern Roach appears to be the result of an ancient hybrid speciation event (
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||
<bibRefCitation id="EFB9A2C56F01CC0DFE89DA8DFE2950ED" author="Mallet, J." box="[353,496,188,213]" pageId="3" pageNumber="224" pagination="279" refId="ref14123" refString="Mallet, J. (2007) Hybrid speciation. Nature, 446 (7133), 279." type="journal article" year="2007">Mallet 2007</bibRefCitation>
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) between ancestral Hitch and CA Roach but it is currently genomically different enough to warrant species status.
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</paragraph>
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<paragraph id="8B97DF346F01CC0DFF2FDB34FC65532C" blockId="3.[151,1437,151,789]" pageId="3" pageNumber="224">
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It is our conclusion that despite hybridization between individuals in the different genera, the distinctiveness of each genus is still strongly maintained and prevalent throughout the genomes interrogated. If results were more admixed, the argument for a single genus would be more plausible. We recognize admixture between genera is uncommon but note that not all mutations lead to post-zygotic barriers (
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||
<bibRefCitation id="EFB9A2C56F01CC0DFBE0DB5EFB2951B1" author="Abbott, R. & Albach, D. & Ansell, S. & Arntzen, J. W. & Baird, S. J. & Bierne, N. & Boughman, J. & Brelsford, A. & Buerkle, C. A. & Buggs, R. & Butlin, R. K." box="[1032,1264,367,392]" pageId="3" pageNumber="224" pagination="229 - 246" refId="ref12389" refString="Abbott, R., Albach, D., Ansell, S., Arntzen, J. W., Baird, S. J., Bierne, N., Boughman, J., Brelsford, A., Buerkle, C. A., Buggs, R. & Butlin, R. K. (2013) Hybridization and speciation. Journal of Evolutionary Biology, 26, 229 - 246." type="journal article" year="2013">
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Abbott
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||
<emphasis id="B95C03266F01CC0DFB8DDB40FB7C51B1" box="[1125,1189,368,392]" italics="true" pageId="3" pageNumber="224">et al.</emphasis>
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2013
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</bibRefCitation>
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||
). Additionally, hybridization has been proposed between Roach/Hitch and other members of the subfamily
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<taxonomicName id="4C28A4B76F01CC0DFB4DDBA5FAF25194" box="[1189,1323,404,429]" class="Actinopterygii" family="Cyprinidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="224" phylum="Chordata" rank="subFamily" subFamily="Leuciscinae">Leuciscinae</taxonomicName>
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including Speckled Dace (
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<emphasis id="B95C03266F01CC0DFEB0DB89FD9C51E9" box="[344,581,440,464]" italics="true" pageId="3" pageNumber="224">
|
||
<taxonomicName id="4C28A4B76F01CC0DFEB0DB89FDE451E9" baseAuthorityName="Girard" baseAuthorityYear="1856" box="[344,573,440,464]" class="Actinopterygii" family="Cyprinidae" genus="Rhinichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="224" phylum="Chordata" rank="species" species="osculus">Rhinichthys osculus</taxonomicName>
|
||
)
|
||
</emphasis>
|
||
, Arroyo Chub (
|
||
<taxonomicName id="4C28A4B76F01CC0DFCEEDB89FC4C51F6" baseAuthorityName="Eigenmann & Eigenmann" baseAuthorityYear="1890" box="[774,917,440,464]" class="Actinopterygii" family="Cyprinidae" genus="Gila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="224" phylum="Chordata" rank="species" species="orcuttii">
|
||
<emphasis id="B95C03266F01CC0DFCEEDB89FC4C51F6" box="[774,917,440,464]" italics="true" pageId="3" pageNumber="224">Gila orcuttii</emphasis>
|
||
</taxonomicName>
|
||
), and the now extinct Thicktail Chub (
|
||
<taxonomicName id="4C28A4B76F01CC0DFA82DB89FEC751CD" baseAuthorityName="Baird & Girard" baseAuthorityYear="1854" class="Actinopterygii" family="Cyprinidae" genus="Gila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="224" phylum="Chordata" rank="species" species="crassicauda">
|
||
<emphasis id="B95C03266F01CC0DFA82DB89FEC751CD" italics="true" pageId="3" pageNumber="224">Gila crassicauda</emphasis>
|
||
</taxonomicName>
|
||
) (
|
||
<bibRefCitation id="EFB9A2C56F01CC0DFEA8DBEDFD4C51CD" author="Greenfield, D. W. & Deckert, G. D." box="[320,661,476,501]" pageId="3" pageNumber="224" pagination="417 - 427" refId="ref13577" refString="Greenfield, D. W. & Deckert, G. D. (1973). Introgressive hybridization between Gila orcutti and Hesperoleucus symmetricus (Pisces: Cyprinidae) in the Cuyama River Basin, California: II. Ecological aspects. Copeia, 1973, 417 - 427." type="journal article" year="1973">Greenfield & Deckert 1973</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFB9A2C56F01CC0DFD43DBEDFC8C51CD" author="Hopkirk, J. D." box="[683,853,476,501]" pageId="3" pageNumber="224" pagination="1 - 160" refId="ref13678" refString="Hopkirk, J. D. (1973) Endemism in fishes of the Clear Lake region of central California. University of California Publications in Zoology, 96, 1 - 160." type="journal article" year="1973">Hopkirk 1973</bibRefCitation>
|
||
). Thus the frequency of hybridization suggests reproductive barriers are poorly developed in the endemic minnows of
|
||
<collectingRegion id="49EC11D66F01CC0DFC39DBCEFB9F5221" box="[977,1094,511,536]" country="United States of America" name="California" pageId="3" pageNumber="224">California</collectingRegion>
|
||
. Secondly, although Northern Roach represent an ancient hybridization event and cluster primarily with individuals from the
|
||
<taxonomicName id="4C28A4B76F01CC0DFB12D814FA965205" box="[1274,1359,549,572]" class="Actinopterygii" family="Cyprinidae" genus="Lavinia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="224" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95C03266F01CC0DFB12D814FA965205" box="[1274,1359,549,572]" italics="true" pageId="3" pageNumber="224">Lavinia</emphasis>
|
||
</taxonomicName>
|
||
genus (~80% but only in the admixture analysis), we cannot definitively say they are
|
||
<taxonomicName id="4C28A4B76F01CC0DFBAED879FB425266" box="[1094,1179,584,607]" class="Actinopterygii" family="Cyprinidae" genus="Lavinia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="224" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95C03266F01CC0DFBAED879FB425266" box="[1094,1179,584,607]" italics="true" pageId="3" pageNumber="224">Lavinia</emphasis>
|
||
</taxonomicName>
|
||
. Northern Roach still strongly resemble CA Roach in appearance (see below), but this may simply be the result of natural selection for the CA Roach morphotype. Future phylogenetic analyses between all genera within the family may provide the solution but it remains, as yet, unavailable. Thus, without universal agreement between genomic analyses or improved criteria for taxonomically delineating hybrids, we default to the original genus assignment by
|
||
<bibRefCitation id="EFB9A2C56F01CC0DFAA5D8E6FF3C532D" author="Snyder, J. O." pageId="3" pageNumber="224" pagination="49 - 72" refId="ref15647" refString="Snyder, J. O. (1913) The fishes of the streams tributary to Monterey Bay, California. Bulletin, U. S. Bureau of Fisheries, 32, 49 - 72." type="journal article" year="1913">Snyder (1913)</bibRefCitation>
|
||
. Therefore, the species of
|
||
<taxonomicName id="4C28A4B76F01CC0DFDE3D8CDFD69532D" authorityName="Snyder" authorityYear="1913" box="[523,688,764,788]" class="Actinopterygii" family="Cyprinidae" genus="Hesperoleucus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="224" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95C03266F01CC0DFDE3D8CDFD69532D" box="[523,688,764,788]" italics="true" pageId="3" pageNumber="224">Hesperoleucus</emphasis>
|
||
</taxonomicName>
|
||
are as described below.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |