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<document ID-DOI="10.11646/zootaxa.5258.4.1" ID-ISSN="1175-5326" ID-Zenodo-Dep="7784322" ID-ZooBank="73F90E50-168F-4441-8B68-8DDFFF8E17D4" checkinTime="1680164509918" checkinUser="plazi" docAuthor="Fedler, Matthew T., Enge, Kevin M. &amp; Moler, Paul E." docDate="2023" docId="03BB87B1FF9E0F3443D5FDC9FD2DFD0B" docLanguage="en" docName="zootaxa.5258.4.1.pdf" docOrigin="Zootaxa 5258 (4)" docSource="http://dx.doi.org/10.11646/zootaxa.5258.4.1" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Siren nettingi" docType="treatment" docVersion="3" lastPageNumber="369" masterDocId="FF82FFC9FF8F0F264342FFEAFFDAFF8F" masterDocTitle="Unraveling Siren (Caudata: Sirenidae) systematics and description of a small seepage specialist" masterLastPageNumber="378" masterPageNumber="351" pageNumber="368" updateTime="1680189431920" updateUser="diego" zenodo-license-document="CLOSED">
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<mods:title>Unraveling Siren (Caudata: Sirenidae) systematics and description of a small seepage specialist</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Fedler, Matthew T.</mods:namePart>
<mods:nameIdentifier type="ORCID">0000-0002-4200-2341</mods:nameIdentifier>
<mods:affiliation>Fish and Wildlife Research Institute, Florida Fish and WildlifeConservation Commission, 1105 SW Williston Road, Gainesville, FL, USA. matthew. fedler @ MyFWC. com; https: // orcid. org / 0000 - 0002 - 4200 - 2341</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Enge, Kevin M.</mods:namePart>
<mods:nameIdentifier type="ORCID">0009-0006-2404-5088</mods:nameIdentifier>
<mods:affiliation>Fish and Wildlife Research Institute, Florida Fish and WildlifeConservation Commission, 1105 SW Williston Road, Gainesville, FL, USA. matthew. fedler @ MyFWC. com; https: // orcid. org / 0000 - 0002 - 4200 - 2341 &amp; kevin. enge @ MyFWC. com; https: // orcid. org / 0009 - 0006 - 2404 - 5088</mods:affiliation>
</mods:name>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Moler, Paul E.</mods:namePart>
<mods:nameIdentifier type="ORCID">0009-0007-3185-8703</mods:nameIdentifier>
<mods:affiliation>Fish and Wildlife Research Institute, Florida Fish and WildlifeConservation Commission, 1105 SW Williston Road, Gainesville, FL, USA. matthew. fedler @ MyFWC. com; https: // orcid. org / 0000 - 0002 - 4200 - 2341 &amp; paul. moler @ MyFWC. com; https: // orcid. org / 0009 - 0007 - 3185 - 8703</mods:affiliation>
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<mods:title>Zootaxa</mods:title>
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<mods:date>2023</mods:date>
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<mods:number>2023-03-30</mods:number>
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<mods:number>5258</mods:number>
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<mods:identifier type="DOI">10.11646/zootaxa.5258.4.1</mods:identifier>
<mods:identifier type="ISSN">1175-5326</mods:identifier>
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<subSubSection box="[151,479,546,573]" pageId="17" pageNumber="368" type="nomenclature">
<paragraph blockId="17.[151,505,546,645]" box="[151,479,546,573]" pageId="17" pageNumber="368">
<heading bold="true" box="[151,479,546,573]" fontSize="11" level="1" pageId="17" pageNumber="368" reason="1">
<taxonomicName ID-CoL="4XR2M" authority="(Goin, 1942)" baseAuthorityName="Goin" baseAuthorityYear="1942" box="[151,479,546,573]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis bold="true" box="[151,479,546,573]" pageId="17" pageNumber="368">
<emphasis bold="true" box="[151,317,547,573]" italics="true" pageId="17" pageNumber="368">Siren nettingi</emphasis>
(
<bibRefCitation author="Goin, C. J." box="[332,470,546,572]" pageId="17" pageNumber="368" pagination="211 - 217" refId="ref13779" refString="Goin, C. J. (1942) Description of a new race of Siren intermedia Le Conte. Annals of the Carnegie Museum, 29, 211 - 217. https: // doi. org / 10.5962 / p. 215158" type="journal article" year="1942">Goin, 1942</bibRefCitation>
)
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="17" pageNumber="368" type="description">
<paragraph blockId="17.[151,505,546,645]" box="[151,324,583,609]" pageId="17" pageNumber="368">
(
<figureCitation box="[159,265,583,609]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURE 3" captionStart-3="FIGURE 4" captionStart-4="FIGURE 5" captionStart-5="FIGURE 6" captionStartId-0="4.[152,255,1973,1997]" captionStartId-1="7.[152,255,1329,1353]" captionStartId-2="8.[152,255,965,989]" captionStartId-3="8.[152,255,1825,1849]" captionStartId-4="9.[152,255,1430,1454]" captionStartId-5="11.[152,255,998,1022]" captionTargetBox-0="[209,1391,453,1942]" captionTargetBox-1="[151,1436,561,1306]" captionTargetBox-2="[163,1422,193,936]" captionTargetBox-3="[166,1421,1058,1794]" captionTargetBox-4="[163,1425,426,1398]" captionTargetBox-5="[177,1403,190,969]" captionTargetId-0="figure-156@4.[197,1391,453,1950]" captionTargetId-1="figure-206@7.[151,1436,561,1306]" captionTargetId-2="figure-20@8.[151,1436,181,941]" captionTargetId-3="figure-53@8.[158,1429,1050,1801]" captionTargetId-4="figure-127@9.[154,1434,417,1406]" captionTargetId-5="figure-18@11.[165,1423,181,974]" captionTargetPageId-0="4" captionTargetPageId-1="7" captionTargetPageId-2="8" captionTargetPageId-3="8" captionTargetPageId-4="9" captionTargetPageId-5="11" captionText-0="FIGURE 1. Bayesian probability tree of concatenated 16S and ND4 regions inferred using BEAST2. Individual specimen labels include voucher ID or GenBank number if no voucher, state, county, and lineage." captionText-1="FIGURE 2. Median joining networks of Siren haplotypes for RAG1 (left) and NCX1 (right) regions made using PopART." captionText-2="FIGURE 3. Box plots with points colored by sex showing distribution of tail proportions of Siren lineages. Outliers are noted with red triangles on the associated box plot." captionText-3="FIGURE 4. Box plots with points representing specimens per showing distribution of costal groove counts of Siren lineages." captionText-4="FIGURE 5. Box plots of the distributions of snout-vent length and the five size-corrected linear measurements compared using linear discriminant analysis." captionText-5="FIGURE 6. Results from our linear discriminant function analysis based on the five size-corrected linear measurements taken from 396 Siren specimens. Except for S. reticulata, each lineage failed to create meaningful separation." figureDoi-0="http://doi.org/10.5281/zenodo.7784326" figureDoi-1="http://doi.org/10.5281/zenodo.7784328" figureDoi-2="http://doi.org/10.5281/zenodo.7784330" figureDoi-3="http://doi.org/10.5281/zenodo.7784332" figureDoi-4="http://doi.org/10.5281/zenodo.7784334" figureDoi-5="http://doi.org/10.5281/zenodo.7784336" httpUri-0="https://zenodo.org/record/7784326/files/figure.png" httpUri-1="https://zenodo.org/record/7784328/files/figure.png" httpUri-2="https://zenodo.org/record/7784330/files/figure.png" httpUri-3="https://zenodo.org/record/7784332/files/figure.png" httpUri-4="https://zenodo.org/record/7784334/files/figure.png" httpUri-5="https://zenodo.org/record/7784336/files/figure.png" pageId="17" pageNumber="368">Figs. 16</figureCitation>
&amp;
<figureCitation box="[301,316,583,609]" captionStart="FIGURE 9" captionStartId="18.[152,255,1509,1533]" captionTargetBox="[151,1436,668,1486]" captionTargetId="figure-242@18.[151,1436,668,1486]" captionTargetPageId="18" captionText="FIGURE 9. Distribution map of specimens assigned to Siren intermedia (orange circles) and S. nettingi (blue circles). The red line is the eastern boundary of the Mobile drainage from EDNA Derived Watersheds for Major Named Rivers (https://edna. usgs.gov/watersheds/kml_index.htm). The Mobile drainage lacks S. intermedia records and is the easternmost drainage where S. nettingi occurs. Our westernmost records of S. intermedia are from the Escambia drainage." figureDoi="http://doi.org/10.5281/zenodo.7784342" httpUri="https://zenodo.org/record/7784342/files/figure.png" pageId="17" pageNumber="368">9</figureCitation>
)
</paragraph>
<paragraph blockId="17.[151,505,546,645]" box="[151,505,618,645]" pageId="17" pageNumber="368">
<emphasis bold="true" box="[151,341,618,645]" pageId="17" pageNumber="368">Common name.</emphasis>
Western
<taxonomicName box="[446,505,619,645]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="genus">Siren</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="17" pageNumber="368" type="materials_examined">
<paragraph blockId="17.[151,1437,690,2013]" pageId="17" pageNumber="368">
<materialsCitation collectingDate="1928-05" country="United States of America" county="Lawrence Co." location="Imboden" pageId="17" pageNumber="368" specimenCount="1" stateProvince="Arkansas" typeStatus="holotype">
<emphasis bold="true" box="[151,267,691,717]" pageId="17" pageNumber="368">
<typeStatus box="[151,262,691,717]" pageId="17" pageNumber="368">Holotype</typeStatus>
.
</emphasis>
Carnegie Museum 7580. Adult female collected in
<date box="[854,969,691,717]" pageId="17" pageNumber="368" value="1928-05">
<collectingDate box="[854,969,691,717]" pageId="17" pageNumber="368" value="1928-05">May 1928</collectingDate>
</date>
from
<location LSID="urn:lsid:plazi:treatment:03BB87B1FF9E0F3443D5FDC9FD2DFD0B:8ECD607CFF9E0F37474DFD59FBAFFD42" box="[1039,1141,691,717]" country="United States of America" county="Lawrence Co." name="Imboden" pageId="17" pageNumber="368" stateProvince="Arkansas">Imboden</location>
,
<collectingCounty box="[1154,1314,691,717]" pageId="17" pageNumber="368">Lawrence Co.</collectingCounty>
,
<collectingRegion box="[1325,1432,691,717]" country="United States of America" name="Arkansas" pageId="17" pageNumber="368">Arkansas</collectingRegion>
,
<collectingCountry box="[151,205,727,753]" name="United States of America" pageId="17" pageNumber="368">USA</collectingCountry>
(
<bibRefCitation author="Goin, C. J." box="[220,345,727,753]" pageId="17" pageNumber="368" pagination="211 - 217" refId="ref13779" refString="Goin, C. J. (1942) Description of a new race of Siren intermedia Le Conte. Annals of the Carnegie Museum, 29, 211 - 217. https: // doi. org / 10.5962 / p. 215158" type="journal article" year="1942">Goin 1942</bibRefCitation>
).
</materialsCitation>
</paragraph>
<paragraph blockId="17.[151,1437,690,2013]" box="[199,525,763,789]" pageId="17" pageNumber="368">
<emphasis bold="true" box="[199,326,763,789]" pageId="17" pageNumber="368">
<typeStatus box="[199,321,763,789]" pageId="17" pageNumber="368">Paratypes</typeStatus>
.
</emphasis>
See
<bibRefCitation author="Goin, C. J." box="[380,521,763,789]" pageId="17" pageNumber="368" pagination="211 - 217" refId="ref13779" refString="Goin, C. J. (1942) Description of a new race of Siren intermedia Le Conte. Annals of the Carnegie Museum, 29, 211 - 217. https: // doi. org / 10.5962 / p. 215158" type="journal article" year="1942">Goin (1942)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="17" pageNumber="368" type="diagnosis">
<paragraph blockId="17.[151,1437,690,2013]" pageId="17" pageNumber="368">
<emphasis bold="true" box="[199,322,798,824]" pageId="17" pageNumber="368">Diagnosis.</emphasis>
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[330,483,799,825]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[330,483,799,825]" italics="true" pageId="17" pageNumber="368">Siren nettingi</emphasis>
</taxonomicName>
has typical
<taxonomicName box="[622,680,799,825]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="genus">
<emphasis box="[622,680,799,825]" italics="true" pageId="17" pageNumber="368">Siren</emphasis>
</taxonomicName>
characteristics: external gills with three fimbriate gill stalks, three associated gill slits, four toes on the forelimbs, lack of pelvic girdle and hindlimbs, and a thin, pigment-bearing mucus layer that overlies the keratinized skin.
<bibRefCitation author="Goin, C. J." box="[696,838,871,897]" pageId="17" pageNumber="368" pagination="211 - 217" refId="ref13779" refString="Goin, C. J. (1942) Description of a new race of Siren intermedia Le Conte. Annals of the Carnegie Museum, 29, 211 - 217. https: // doi. org / 10.5962 / p. 215158" type="journal article" year="1942">Goin (1942)</bibRefCitation>
distinguished
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[1008,1125,871,897]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[1008,1125,871,897]" italics="true" pageId="17" pageNumber="368">S. nettingi</emphasis>
</taxonomicName>
from
<taxonomicName baseAuthorityName="Noble &amp; Marshall" baseAuthorityYear="1932" box="[1200,1353,871,897]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="intermedia">
<emphasis box="[1200,1353,871,897]" italics="true" pageId="17" pageNumber="368">S. intermedia</emphasis>
</taxonomicName>
by the presence of well-defined light spots on the sides and venter and also specifying that these were not the bar-like streaks found in
<taxonomicName box="[342,472,943,969]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="lacertina">
<emphasis box="[342,472,943,969]" italics="true" pageId="17" pageNumber="368">S. lacertina</emphasis>
</taxonomicName>
. Additionally,
<bibRefCitation author="Goin, C. J." box="[642,775,943,969]" pageId="17" pageNumber="368" pagination="211 - 217" refId="ref13779" refString="Goin, C. J. (1942) Description of a new race of Siren intermedia Le Conte. Annals of the Carnegie Museum, 29, 211 - 217. https: // doi. org / 10.5962 / p. 215158" type="journal article" year="1942">Goin (1942</bibRefCitation>
, p. 212) stated that
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[1001,1117,943,969]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[1001,1117,943,969]" italics="true" pageId="17" pageNumber="368">S. nettingi</emphasis>
</taxonomicName>
has “about two more costal grooves (usually
<quantity box="[349,409,979,1005]" metricMagnitude="-1" metricUnit="m" metricValue="8.382" pageId="17" pageNumber="368" unit="in" value="33.0">33 in</quantity>
<taxonomicName baseAuthorityName="Noble &amp; Marshall" baseAuthorityYear="1932" box="[418,539,979,1005]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="intermedia">
<emphasis box="[418,539,979,1005]" italics="true" pageId="17" pageNumber="368">intermedia</emphasis>
</taxonomicName>
,
<quantity box="[556,616,978,1005]" metricMagnitude="-1" metricUnit="m" metricValue="8.89" pageId="17" pageNumber="368" unit="in" value="35.0">35 in</quantity>
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[625,711,979,1004]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[625,711,979,1004]" italics="true" pageId="17" pageNumber="368">nettingi</emphasis>
</taxonomicName>
),” a value slightly overlapping but less than the average for
<taxonomicName class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="lacertina">
<emphasis box="[1415,1429,979,1005]" italics="true" pageId="17" pageNumber="368">S</emphasis>
.
<emphasis box="[151,252,1015,1041]" italics="true" pageId="17" pageNumber="368">lacertina</emphasis>
</taxonomicName>
. See comments below for possible reasons for variation in data.
</paragraph>
<paragraph blockId="17.[151,1437,690,2013]" pageId="17" pageNumber="368">
In areas of sympatry, the costal groove range of
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[750,865,1051,1077]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[750,865,1051,1077]" italics="true" pageId="17" pageNumber="368">S. nettingi</emphasis>
</taxonomicName>
overlaps that of
<emphasis box="[1057,1391,1051,1077]" italics="true" pageId="17" pageNumber="368">
<taxonomicName authorityName="Fedler &amp; Enge &amp; Moler" authorityYear="2023" box="[1057,1228,1051,1077]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="sphagnicola">S. sphagnicola</taxonomicName>
.
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[1239,1391,1051,1077]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">Siren nettingi</taxonomicName>
</emphasis>
has 3234 costal grooves, with two outliers of 31 and 36 (
<figureCitation box="[770,840,1087,1113]" captionStart="FIGURE 4" captionStartId="8.[152,255,1825,1849]" captionTargetBox="[166,1421,1058,1794]" captionTargetId="figure-53@8.[158,1429,1050,1801]" captionTargetPageId="8" captionText="FIGURE 4. Box plots with points representing specimens per showing distribution of costal groove counts of Siren lineages." figureDoi="http://doi.org/10.5281/zenodo.7784332" httpUri="https://zenodo.org/record/7784332/files/figure.png" pageId="17" pageNumber="368">Fig. 4</figureCitation>
).
<bibRefCitation author="Goin, C. J." box="[862,1001,1087,1113]" pageId="17" pageNumber="368" pagination="211 - 217" refId="ref13779" refString="Goin, C. J. (1942) Description of a new race of Siren intermedia Le Conte. Annals of the Carnegie Museum, 29, 211 - 217. https: // doi. org / 10.5962 / p. 215158" type="journal article" year="1942">Goin (1942)</bibRefCitation>
gave a slightly higher range of 3337 costal grooves, but we infer that he included the axial groove or terminated counting grooves at a point posterior to our stopping point.
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[372,524,1159,1185]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[372,524,1159,1185]" italics="true" pageId="17" pageNumber="368">Siren nettingi</emphasis>
</taxonomicName>
specimens similar in size to
<taxonomicName authorityName="Fedler &amp; Enge &amp; Moler" authorityYear="2023" box="[850,1016,1159,1185]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="sphagnicola">
<emphasis box="[850,1016,1159,1185]" italics="true" pageId="17" pageNumber="368">S. sphagnicola</emphasis>
</taxonomicName>
can be distinguished by the presence of a solid yellow labial stripe and rostral patch, whereas
<taxonomicName authorityName="Fedler &amp; Enge &amp; Moler" authorityYear="2023" box="[786,952,1195,1221]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="sphagnicola">
<emphasis box="[786,952,1195,1221]" italics="true" pageId="17" pageNumber="368">S. sphagnicola</emphasis>
</taxonomicName>
has, at most, several beige spots where the labial stripe occurs in other species. In addition, bodies of small
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[870,984,1231,1257]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[870,984,1231,1257]" italics="true" pageId="17" pageNumber="368">S. nettingi</emphasis>
</taxonomicName>
often have a green or yellowish hue and gold flecking, both of with are lacking in
<taxonomicName authorityName="Fedler &amp; Enge &amp; Moler" authorityYear="2023" box="[617,782,1267,1293]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="sphagnicola">
<emphasis box="[617,782,1267,1293]" italics="true" pageId="17" pageNumber="368">S. sphagnicola</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection pageId="17" pageNumber="368" type="discussion">
<paragraph blockId="17.[151,1437,690,2013]" pageId="17" pageNumber="368">
<emphasis bold="true" box="[199,335,1302,1328]" pageId="17" pageNumber="368">Comments.</emphasis>
This species occupies a large geographic area, and we analyzed relatively few specimens, mainly from
<collectingRegion box="[212,313,1339,1365]" country="United States of America" name="Alabama" pageId="17" pageNumber="368">Alabama</collectingRegion>
and
<collectingRegion box="[369,479,1339,1365]" country="United States of America" name="Louisiana" pageId="17" pageNumber="368">Louisiana</collectingRegion>
(
<figureCitation box="[496,565,1339,1365]" captionStart="FIGURE 9" captionStartId="18.[152,255,1509,1533]" captionTargetBox="[151,1436,668,1486]" captionTargetId="figure-242@18.[151,1436,668,1486]" captionTargetPageId="18" captionText="FIGURE 9. Distribution map of specimens assigned to Siren intermedia (orange circles) and S. nettingi (blue circles). The red line is the eastern boundary of the Mobile drainage from EDNA Derived Watersheds for Major Named Rivers (https://edna. usgs.gov/watersheds/kml_index.htm). The Mobile drainage lacks S. intermedia records and is the easternmost drainage where S. nettingi occurs. Our westernmost records of S. intermedia are from the Escambia drainage." figureDoi="http://doi.org/10.5281/zenodo.7784342" httpUri="https://zenodo.org/record/7784342/files/figure.png" pageId="17" pageNumber="368">Fig. 9</figureCitation>
). Our information may not represent the full extent of variation found in this species.
<bibRefCitation author="Reinhard, S. &amp; Voitel, S. &amp; Kupfer, A." box="[246,493,1375,1401]" pageId="17" pageNumber="368" pagination="1 - 5" refId="ref15937" refString="Reinhard, S., Voitel, S. and Kupfer, A. (2013) External fertilisation and paternal care in the paedomorphic salamander Siren intermedia Barnes, 1826 (Urodela: Sirenidae). Zoologischer Anzeiger-A Journal of Comparative Zoology, 253 (1), 1 - 5. https: // doi. org / 10.1016 / j. jcz. 2013.06.002" type="journal article" year="2013">
Reinhard
<emphasis box="[356,412,1375,1401]" italics="true" pageId="17" pageNumber="368">et al.</emphasis>
(2013)
</bibRefCitation>
provided photographs of larvae and mature adults of this species.
</paragraph>
<paragraph blockId="17.[151,1437,690,2013]" pageId="17" pageNumber="368">
After examining the
<typeStatus box="[430,531,1411,1437]" pageId="17" pageNumber="368">holotype</typeStatus>
, we concluded that the well-defined, light spots on the venter referenced by
<bibRefCitation author="Goin, C. J." pageId="17" pageNumber="368" pagination="211 - 217" refId="ref13779" refString="Goin, C. J. (1942) Description of a new race of Siren intermedia Le Conte. Annals of the Carnegie Museum, 29, 211 - 217. https: // doi. org / 10.5962 / p. 215158" type="journal article" year="1942">Goin (1942)</bibRefCitation>
represent sensory pores rather than chromatophores. Chromatophores on the
<typeStatus box="[1115,1213,1447,1473]" pageId="17" pageNumber="368">holotype</typeStatus>
are neither defined nor extensive in coverage. Regardless of this distinction, neither the distinct, light-colored sensory pores nor the yellow to yellow-green flecking of chromatophores is unique to
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[879,993,1519,1545]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[879,993,1519,1545]" italics="true" pageId="17" pageNumber="368">S. nettingi</emphasis>
</taxonomicName>
. These pores are frequently obfuscated by the slime layer but become more visible when this is removed. This slime layer frequently sloughs off of poorly preserved or frozen specimens.
</paragraph>
<paragraph blockId="17.[151,1437,690,2013]" pageId="17" pageNumber="368">
Validity of the Rio Grande
<taxonomicName box="[500,559,1627,1653]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="genus">Siren</taxonomicName>
(
<taxonomicName authority="Goin, 1957" authorityName="Goin" authorityYear="1957" box="[574,937,1626,1653]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="subSpecies" species="intermedia" subSpecies="texana">
<emphasis box="[574,804,1627,1653]" italics="true" pageId="17" pageNumber="368">S. intermedia texana</emphasis>
<bibRefCitation author="Goin, C. J." box="[810,937,1626,1653]" pageId="17" pageNumber="368" pagination="37 - 42" refId="ref13823" refString="Goin, C. J. (1957) Description of a new salamander of the genus Siren from the Rio Grande. Herpetologica, 13 (1), 37 - 42. https: // doi. org / 10.2307 / 1439392" type="journal article" year="1957">Goin, 1957</bibRefCitation>
</taxonomicName>
) as a taxonomic unit distinct from
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[1323,1437,1627,1653]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[1323,1437,1627,1653]" italics="true" pageId="17" pageNumber="368">S. nettingi</emphasis>
</taxonomicName>
has been questioned in recent decades. Our analyses suggest that
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[886,1001,1663,1689]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[886,1001,1663,1689]" italics="true" pageId="17" pageNumber="368">S. nettingi</emphasis>
</taxonomicName>
and “texana” are either closely related sister taxa or ecomorphs of the same species adapted to habitats present in different regions. Further investigation of this topic is needed, but we treated “texana” as a distinct unit for morphological analyses. Furthermore,
<emphasis box="[1291,1333,1735,1761]" italics="true" pageId="17" pageNumber="368">S. i.</emphasis>
“texana” is no longer recognized by
<bibRefCitation author="Highton, R. &amp; Bonett, R. M. &amp; Jockusch, E. L." box="[468,714,1771,1797]" pageId="17" pageNumber="368" pagination="18 - 30" refId="ref14319" refString="Highton, R., Bonett, R. M. &amp; Jockusch, E. L. (2017) Caudata salamanders. In: Crother, B. I. (Ed.), Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with Comments Regarding Confidence in our Understanding. Society for the Study of Amphibians and Reptiles Herpetological Circular No. 43. 8 th Edition. Society for the Study of Amphibians and Reptiles, s. n., pp. 18 - 30." type="book chapter" year="2017">
Highton
<emphasis box="[569,628,1771,1797]" italics="true" pageId="17" pageNumber="368">et al.</emphasis>
(2017)
</bibRefCitation>
, who accepted the erroneous designation by
<bibRefCitation author="Flores-Villela, O. &amp; Brandon, R. A." pageId="17" pageNumber="368" pagination="289 - 291" refId="ref13630" refString="Flores-Villela, O. &amp; Brandon, R. A. (1992) Siren lacertina (Amphibia: Caudata) in northeastern Mexico and southern Texas. Annals of the Carnegie Museum, 61, 289 - 291. https: // doi. org / 10.5962 / p. 226656" type="journal article" year="1992">Flores-Villela and Brandon (1992)</bibRefCitation>
that the large siren from southern Texas and
<collectingCountry box="[834,920,1807,1833]" name="Mexico" pageId="17" pageNumber="368">Mexico</collectingCountry>
is
<taxonomicName authority="(LaFortune 2015)" baseAuthorityName="LaFortune" baseAuthorityYear="2015" box="[952,1290,1806,1833]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="lacertina">
<emphasis box="[952,1081,1807,1833]" italics="true" pageId="17" pageNumber="368">S. lacertina</emphasis>
(
<bibRefCitation author="LaFortune, T. C." box="[1096,1282,1806,1833]" pageId="17" pageNumber="368" refId="ref14606" refString="LaFortune, T. C. (2015) Species Identification and Habitat Assessment of the South Texas Siren. M. S. Thesis, University of Texas at Brownsville, Brownsville, Texas, 91 pp." type="book" year="2015">LaFortune 2015</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph blockId="17.[151,1437,690,2013]" pageId="17" pageNumber="368">
<emphasis bold="true" box="[199,254,1842,1868]" pageId="17" pageNumber="368">Size.</emphasis>
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[260,410,1843,1869]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[260,410,1843,1869]" italics="true" pageId="17" pageNumber="368">Siren nettingi</emphasis>
</taxonomicName>
has a reported maximum TL of
<quantity box="[762,853,1843,1869]" metricMagnitude="-1" metricUnit="m" metricValue="6.859999999999999" pageId="17" pageNumber="368" unit="mm" value="686.0">686 mm</quantity>
for the larger “texana” form found in southern Texas and
<collectingCountry box="[198,286,1879,1905]" name="Mexico" pageId="17" pageNumber="368">Mexico</collectingCountry>
, whereas the smaller form found in the rest of its range has a reported maximum TL of
<quantity box="[1251,1343,1878,1905]" metricMagnitude="-1" metricUnit="m" metricValue="5.02" pageId="17" pageNumber="368" unit="mm" value="502.0">502 mm</quantity>
(
<bibRefCitation author="Martof, B. S." pageId="17" pageNumber="368" pagination="1 - 3" refId="ref14975" refString="Martof, B. S. (1973 a) Siren intermedia Le Conte. Lesser Siren. Catalogue of American Amphibians and Reptiles, 127, 1 - 3." type="journal article" year="1973">Martof 1973a</bibRefCitation>
).
<bibRefCitation author="Flores-Villela, O. &amp; Brandon, R. A." box="[243,631,1915,1941]" pageId="17" pageNumber="368" pagination="289 - 291" refId="ref13630" refString="Flores-Villela, O. &amp; Brandon, R. A. (1992) Siren lacertina (Amphibia: Caudata) in northeastern Mexico and southern Texas. Annals of the Carnegie Museum, 61, 289 - 291. https: // doi. org / 10.5962 / p. 226656" type="journal article" year="1992">Flores-Villela and Brandon (1992)</bibRefCitation>
cited
<bibRefCitation author="Goin, C. J." box="[701,839,1914,1941]" pageId="17" pageNumber="368" pagination="37 - 42" refId="ref13823" refString="Goin, C. J. (1957) Description of a new salamander of the genus Siren from the Rio Grande. Herpetologica, 13 (1), 37 - 42. https: // doi. org / 10.2307 / 1439392" type="journal article" year="1957">Goin (1957)</bibRefCitation>
for total lengths given by
<bibRefCitation author="Martof, B. S." box="[1138,1302,1915,1941]" pageId="17" pageNumber="368" pagination="1 - 3" refId="ref14975" refString="Martof, B. S. (1973 a) Siren intermedia Le Conte. Lesser Siren. Catalogue of American Amphibians and Reptiles, 127, 1 - 3." type="journal article" year="1973">Martof (1973a</bibRefCitation>
and
<bibRefCitation author="Martof, B. S." box="[1357,1428,1915,1941]" pageId="17" pageNumber="368" pagination="1 - 2" refId="ref15006" refString="Martof, B. S. (1973 b) Siren lacertina Linnaeus. Greater Siren. Catalogue of American Amphibians and Reptiles, 128, 1 - 2." type="journal article" year="1973">1973b</bibRefCitation>
) and incorrectly reported the TL as the SVL for both the “texana” form of
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[966,1080,1951,1977]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[966,1080,1951,1977]" italics="true" pageId="17" pageNumber="368">S. nettingi</emphasis>
</taxonomicName>
(then
<taxonomicName authorityName="Goin" authorityYear="1957" box="[1150,1380,1951,1977]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="subSpecies" species="intermedia" subSpecies="texana">
<emphasis box="[1150,1380,1951,1977]" italics="true" pageId="17" pageNumber="368">S. intermedia texana</emphasis>
</taxonomicName>
) and
<taxonomicName baseAuthorityName="LaFortune" baseAuthorityYear="2015" box="[151,280,1987,2013]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="368" phylum="Chordata" rank="species" species="lacertina">
<emphasis box="[151,280,1987,2013]" italics="true" pageId="17" pageNumber="368">S. lacertina</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="369" type="distribution">
<paragraph blockId="18.[151,1436,150,644]" pageId="18" pageNumber="369">
<emphasis bold="true" box="[199,353,150,176]" pageId="18" pageNumber="369">Distribution.</emphasis>
Based on our examination of museum specimens and genetic analyses,
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[1177,1292,151,177]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[1177,1292,151,177]" italics="true" pageId="18" pageNumber="369">S. nettingi</emphasis>
</taxonomicName>
occurs from the Mobile Bay drainage westward, and native populations do not occur in any Florida drainage (
<figureCitation box="[1253,1322,186,212]" captionStart="FIGURE 9" captionStartId="18.[152,255,1509,1533]" captionTargetBox="[151,1436,668,1486]" captionTargetId="figure-242@18.[151,1436,668,1486]" captionTargetPageId="18" captionText="FIGURE 9. Distribution map of specimens assigned to Siren intermedia (orange circles) and S. nettingi (blue circles). The red line is the eastern boundary of the Mobile drainage from EDNA Derived Watersheds for Major Named Rivers (https://edna. usgs.gov/watersheds/kml_index.htm). The Mobile drainage lacks S. intermedia records and is the easternmost drainage where S. nettingi occurs. Our westernmost records of S. intermedia are from the Escambia drainage." figureDoi="http://doi.org/10.5281/zenodo.7784342" httpUri="https://zenodo.org/record/7784342/files/figure.png" pageId="18" pageNumber="369">Fig. 9</figureCitation>
). A single specimen that sequenced as
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[471,586,223,249]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[471,586,223,249]" italics="true" pageId="18" pageNumber="369">S. nettingi</emphasis>
</taxonomicName>
was collected from Tatess Hell State Forest, Franklin Co., Florida, but we suspect this specimen was translocated as live fish bait. Based on phenotypes and costal groove counts,
<bibRefCitation author="Goin, C. J." box="[1299,1436,258,284]" pageId="18" pageNumber="369" pagination="211 - 217" refId="ref13779" refString="Goin, C. J. (1942) Description of a new race of Siren intermedia Le Conte. Annals of the Carnegie Museum, 29, 211 - 217. https: // doi. org / 10.5962 / p. 215158" type="journal article" year="1942">Goin (1942)</bibRefCitation>
suggested that Florida Parishes of Louisiana represented the break between the then-recognized subspecies
<taxonomicName baseAuthorityName="Noble &amp; Marshall" baseAuthorityYear="1932" box="[1315,1436,294,320]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="intermedia">
<emphasis box="[1315,1436,294,320]" italics="true" pageId="18" pageNumber="369">intermedia</emphasis>
</taxonomicName>
and
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[198,284,331,356]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[198,284,331,356]" italics="true" pageId="18" pageNumber="369">nettingi</emphasis>
</taxonomicName>
. This boundary was later considered to be a hybridization zone that stretched into Mississippi (
<bibRefCitation author="Boyd, C. &amp; Vickers, D. H." pageId="18" pageNumber="369" pagination="202 - 205" refId="ref13278" refString="Boyd, C. &amp; Vickers, D. H. (1963) Distribution of some Mississippi amphibians and reptiles. Herpetologica, 19 (3), 202 - 205." type="journal article" year="1963">Boyd &amp; Vickers 1963</bibRefCitation>
) and then to the Mobile Bay drainage (
<bibRefCitation author="Caldwell, R. D. &amp; Howell, W. M." box="[749,1040,366,392]" pageId="18" pageNumber="369" pagination="310 - 311" refId="ref13311" refString="Caldwell, R. D. &amp; Howell, W. M. (1966) Siren intermedia nettingi from Alabama. Herpetologica 22 (4), 310 - 311." type="journal article" year="1966">Caldwell &amp; Howell 1966</bibRefCitation>
). We suspect part of this confusion resulted from examining
<taxonomicName authorityName="Fedler &amp; Enge &amp; Moler" authorityYear="2023" box="[440,608,402,429]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="sphagnicola">
<emphasis box="[440,608,402,429]" italics="true" pageId="18" pageNumber="369">S. sphagnicola</emphasis>
</taxonomicName>
and attributing them to
<emphasis box="[889,1064,402,429]" italics="true" pageId="18" pageNumber="369">
S. i.
<taxonomicName baseAuthorityName="Noble &amp; Marshall" baseAuthorityYear="1932" box="[943,1064,402,428]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="intermedia">intermedia</taxonomicName>
</emphasis>
because of their resemblance to grayish
<taxonomicName baseAuthorityName="Noble &amp; Marshall" baseAuthorityYear="1932" box="[240,388,438,465]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="intermedia">
<emphasis box="[240,388,438,465]" italics="true" pageId="18" pageNumber="369">S. intermedia</emphasis>
</taxonomicName>
that can be found farther east. Furthermore,
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[880,993,439,465]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[880,993,439,465]" italics="true" pageId="18" pageNumber="369">S. nettingi</emphasis>
</taxonomicName>
is currently thought to be the only
<taxonomicName pageId="18" pageNumber="369">
<emphasis box="[1378,1436,439,465]" italics="true" pageId="18" pageNumber="369">Siren</emphasis>
species
</taxonomicName>
that occurs from the Mississippi River west and south to
<collectingRegion box="[876,979,475,501]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="18" pageNumber="369">Veracruz</collectingRegion>
,
<collectingCountry box="[990,1079,474,500]" name="Mexico" pageId="18" pageNumber="369">Mexico</collectingCountry>
.
</paragraph>
<paragraph blockId="18.[151,1436,150,644]" pageId="18" pageNumber="369">
<emphasis bold="true" box="[199,395,510,536]" pageId="18" pageNumber="369">Common Name.</emphasis>
We suggest the common name Western
<taxonomicName box="[859,922,510,536]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="genus">Siren</taxonomicName>
. Except for the Mobile Bay drainage system, where
<taxonomicName box="[227,363,546,573]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="reticulata">
<emphasis box="[227,363,546,573]" italics="true" pageId="18" pageNumber="369">S. reticulata</emphasis>
</taxonomicName>
is present,
<taxonomicName baseAuthorityName="Goin" baseAuthorityYear="1942" box="[491,605,547,573]" class="Amphibia" family="Sirenidae" genus="Siren" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="369" phylum="Chordata" rank="species" species="nettingi">
<emphasis box="[491,605,547,573]" italics="true" pageId="18" pageNumber="369">S. nettingi</emphasis>
</taxonomicName>
represents the largest or only siren (from the
<collectingRegion box="[1116,1244,546,572]" country="United States of America" name="Mississippi" pageId="18" pageNumber="369">Mississippi</collectingRegion>
River westward) present throughout its range; thus, the “Lesser” moniker is misleading.
</paragraph>
<paragraph blockId="18.[151,1436,150,644]" box="[199,759,618,644]" pageId="18" pageNumber="369">
<emphasis bold="true" box="[199,454,618,644]" pageId="18" pageNumber="369">Specimens examined.</emphasis>
See Supplemental
<tableCitation box="[669,753,618,644]" captionStart="TABLE 1" captionStartId="2.[152,245,548,572]" captionTargetPageId="2" captionText="TABLE 1. Primers used for PCR and sequencing gene fragments." pageId="18" pageNumber="369">Table 1</tableCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>