Add updates up until 2025-04-15 14:13:42

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<mods:title id="09DCC081ED4EB0C1B58CE56B83877ACE">The song of the Doto (Nudibranchia: Dotidae): newly documented diversity from the coast of California</mods:title>
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<mods:affiliation id="2629B2343CF7AFDAF2B0FEF9FDBDBACF">Department of Invertebrate Zoology and Geology, California Academy of Sciences, San Francisco, CA 94118, U. S. A. &amp; Department of Biological Sciences, Northern Arizona University, 617 S. Beaver St., Flagstaff, AZ 86011, U. S. A. &amp; Center for Ecosystem Science and Society, Northern Arizona University, 617 S. Beaver St., Flagstaff, AZ 86011</mods:affiliation>
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<mods:affiliation id="ED410479B1E3E5DB2C3A9A827ACC99AB">Department of Invertebrate Zoology and Geology, California Academy of Sciences, San Francisco, CA 94118, U. S. A.</mods:affiliation>
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Gosliner &amp; Adayapalam
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(
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,
<figureCitation id="131CB69EFFA2E97FD22F177DB717F8F1" box="[305,341,1829,1855]" captionStart="FIGURE 3" captionStartId="15.[151,250,1943,1967]" captionTargetBox="[222,1369,183,1917]" captionTargetId="figure-22@15.[212,1376,181,1920]" captionTargetPageId="15" captionText="FIGURE 3. Cerata. A. Doto urak sp. nov., paratype, CASIZ 184511. B. Doto columbiana ODonoghue, 1921, CASIZ 189302. C. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 2.5 mm." figureDoi="http://doi.org/10.5281/zenodo.15215502" httpUri="https://zenodo.org/record/15215502/files/figure.png" pageId="11" pageNumber="564">3A</figureCitation>
,
<figureCitation id="131CB69EFFA2E97FD27F177DB733F8F1" box="[353,369,1829,1855]" captionStart="FIGURE 4" captionStartId="16.[151,250,1169,1193]" captionTargetBox="[151,1436,181,1145]" captionTargetId="figure-38@16.[151,1436,181,1145]" captionTargetPageId="16" captionText="FIGURE 4. Doto urak sp. nov., scanning electron micrographs of radula. AC. Paratype, CASIZ 217599. AB. Detail of radular teeth. C. Large section of radular ribbon. D. Paratype, CASIZ 184511. Radular ribbon." figureDoi="http://doi.org/10.5281/zenodo.15215504" httpUri="https://zenodo.org/record/15215504/files/figure.png" pageId="11" pageNumber="564">4</figureCitation>
,
<figureCitation id="131CB69EFFA2E97FD263177DB7E2F88E" box="[381,416,1829,1856]" captionStart="FIGURE 5" captionStartId="18.[151,250,1908,1932]" captionTargetBox="[247,1337,182,1881]" captionTargetId="figure-38@18.[247,1340,181,1884]" captionTargetPageId="18" captionText="FIGURE 5. Reproductive system. A. Doto urak sp. nov., paratypes, composite of 184511 and 217599, scale = 2.0 mm. B. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 1.5 mm. Abbreviations: am = ampulla, ej = ejaculatory portion of vas deferens, fgm = female gland mass, p = penial sac, pr = prostatic portion of vas deferens, rs = receptaculum seminis, v = vagina." figureDoi="http://doi.org/10.5281/zenodo.15215506" httpUri="https://zenodo.org/record/15215506/files/figure.png" pageId="11" pageNumber="564">5A</figureCitation>
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, lower left photo, misidentification;
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, upper photo, misidentification;
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, middle photo, misidentification.
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<emphasis id="B9537609FFA2E97FD38917FFB688F870" box="[151,202,1959,1982]" italics="true" pageId="11" pageNumber="564">Doto</emphasis>
sp.
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form A—
<bibRefCitation id="EFB6D7EAFFA2E97FD24817FFB7B2F871" author="Goddard, J. H. R." box="[342,496,1959,1983]" pageId="11" pageNumber="564" pagination="43 - 54" refId="ref13129" refString="Goddard, J. H. R. (1996) Lecithotrophic development in Doto amyra (Nudibranchia: Dendronotacea), with a review of developmental mode in the genus. Veliger, 39 (1), 43-54." type="journal article" year="1996">Goddard, 1996</bibRefCitation>
;
<bibRefCitation id="EFB6D7EAFFA2E97FD2E517FFB54CF871" author="Shipman, C. &amp; Gosliner, T." box="[507,782,1959,1983]" pageId="11" pageNumber="564" pagination="57 - 101" refId="ref15905" refString="Shipman, C. &amp; Gosliner, T. (2015) Molecular and morphological systematics of Doto Oken, 1851 (Gastropoda: Heterobranchia) with descriptions of five new species and a new genus. Zootaxa, 3973 (1) 57-101. https://doi.org/10.11646/zootaxa.3973.1.2" type="journal article" year="2015">Shipman &amp; Gosliner, 2015</bibRefCitation>
;
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Behrens
<emphasis id="B9537609FFA2E97FD06F17FFB5E7F871" box="[881,933,1959,1983]" italics="true" pageId="11" pageNumber="564">et al.</emphasis>
, 2022: 112
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, lower left photo.
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<collectingRegion id="49E364F9FFA5E978D7D010CFB3DAFF7F" box="[1230,1432,151,177]" pageId="12" pageNumber="565">San Pedro Harbor</collectingRegion>
,
<date id="FF998CDBFFA5E978D38910E3B72DFF1B" box="[151,367,187,213]" pageId="12" pageNumber="565" value="2009-12-12">
<collectingDate id="EFDD7533FFA5E978D38910E3B72DFF1B" box="[151,367,187,213]" pageId="12" pageNumber="565" value="2009-12-12">12 December 2009</collectingDate>
</date>
,
<collectorName id="26D2CFCDFFA5E978D26410E4B7A5FF1B" box="[378,487,187,215]" pageId="12" pageNumber="565">Á. Valdés</collectorName>
collector.
</materialsCitation>
</paragraph>
<paragraph id="8B98AA1BFFA5E978D3D91087B7B2FE63" blockId="12.[151,1437,150,2013]" pageId="12" pageNumber="565">
<materialsCitation id="3B4FA046FFA5E978D3D91087B736FED3" collectingDate="1984-04-24" collectionCode="CASIZ" collectorName="Jeff Goddard" county="Drake's Estero Marin County" latitude="35.964554" location="Drake's Estero Marin County" longLatPrecision="1" longitude="122.00185" municipality="Ellwood" pageId="12" pageNumber="565" specimenCode="CASIZ 056222" specimenCount="15" stateProvince="CA" typeStatus="paratype">
<emphasis id="B9537609FFA5E978D3D91087B77DFF37" bold="true" box="[199,319,223,249]" pageId="12" pageNumber="565">
<typeStatus id="549C14B9FFA5E978D3D91087B77DFF37" box="[199,319,223,249]" pageId="12" pageNumber="565" type="paratype">Paratypes</typeStatus>
</emphasis>
:
<specimenCode id="DB810260FFA5E978D24E1087B443FF37" box="[336,513,222,249]" collectionCode="CASIZ" pageId="12" pageNumber="565">CASIZ 056222</specimenCode>
,
<specimenCount id="9D216192FFA5E978D1111086B4EEFF37" box="[527,684,222,249]" count="15" pageId="12" pageNumber="565" type="generic">15 specimens</specimenCount>
, mouth of
<collectingCounty id="62F9D297FFA5E978D0351087B23CFF37" box="[811,1150,223,249]" pageId="12" pageNumber="565">Drakes Estero Marin County</collectingCounty>
,
<collectingRegion id="49E364F9FFA5E978D7951087B2F7FF37" box="[1163,1205,223,249]" pageId="12" pageNumber="565">CA</collectingRegion>
,
<date id="FF998CDBFFA5E978D7DC1087B327FF37" box="[1218,1381,223,249]" pageId="12" pageNumber="565" value="1984-04-24">
<collectingDate id="EFDD7533FFA5E978D7DC1087B327FF37" box="[1218,1381,223,249]" pageId="12" pageNumber="565" value="1984-04-24">24 April 1984</collectingDate>
</date>
,
<collectorName id="26D2CFCDFFA5E978D66D1087B6BEFED3" pageId="12" pageNumber="565">Jeff Goddard</collectorName>
, collector.
</materialsCitation>
<materialsCitation id="3B4FA046FFA5E978D29E115BB27AFE8F" collectingDate="2010-03-15" collectionCode="CASIZ" collectorName="Jeff Goddard" county="Santa Barbara County" latitude="35.964554" location="Gaviota" longLatPrecision="1" longitude="122.00185" municipality="Ellwood" pageId="12" pageNumber="565" specimenCode="CASIZ 184511" specimenCount="7" stateProvince="California" typeStatus="paratype">
<specimenCode id="DB810260FFA5E978D29E115BB471FED3" box="[384,563,258,285]" collectionCode="CASIZ" pageId="12" pageNumber="565">CASIZ 184511</specimenCode>
,
<specimenCount id="9D216192FFA5E978D15D115BB497FED3" box="[579,725,259,285]" count="7" pageId="12" pageNumber="565" type="generic">7 specimens</specimenCount>
, one dissected,
<collectingMunicipality id="6BFC3061FFA5E978D08D115BB5B0FED3" box="[915,1010,259,285]" pageId="12" pageNumber="565">Ellwood</collectingMunicipality>
boulder field, between
<location id="8EF8FCC0FFA5E978D611115BB32AFED3" LSID="urn:lsid:plazi:treatment:038E1B0DFFA2E97AD3891685B5C7FDD7:8EF8FCC0FFA5E978D611115BB32AFED3" box="[1295,1384,259,285]" county="Santa Barbara County" latitude="35.964554" longLatPrecision="1" longitude="122.00185" municipality="Ellwood" name="Gaviota" pageId="12" pageNumber="565" stateProvince="California">Gaviota</location>
and
<location id="8EF8FCC0FFA5E978D389117FB6A6FE8F" LSID="urn:lsid:plazi:treatment:038E1B0DFFA2E97AD3891685B5C7FDD7:8EF8FCC0FFA5E978D389117FB6A6FE8F" box="[151,228,295,321]" county="Santa Barbara County" latitude="35.964554" longLatPrecision="1" longitude="122.00185" municipality="Ellwood" name="Goleta" pageId="12" pageNumber="565" stateProvince="California">Goleta</location>
,
<collectingCounty id="62F9D297FFA5E978D3EF117FB7AEFE8F" box="[241,492,295,321]" pageId="12" pageNumber="565">Santa Barbara County</collectingCounty>
,
<collectingRegion id="49E364F9FFA5E978D2E9117FB429FE8F" box="[503,619,295,321]" country="United States of America" name="California" pageId="12" pageNumber="565">California</collectingRegion>
,
<date id="FF998CDBFFA5E978D166117EB56AFE8F" box="[632,808,294,321]" pageId="12" pageNumber="565" value="2010-03-15">
<collectingDate id="EFDD7533FFA5E978D166117EB56AFE8F" box="[632,808,294,321]" pageId="12" pageNumber="565" value="2010-03-15">15 March 2010</collectingDate>
</date>
,
<collectorName id="26D2CFCDFFA5E978D02A117FB58AFE8F" box="[820,968,295,321]" pageId="12" pageNumber="565">Jeff Goddard</collectorName>
collector.
</materialsCitation>
<materialsCitation id="3B4FA046FFA5E978D721117FB3DFFEAB" collectingDate="2017-08-18" collectionCode="CASIZ" collectorName="Brenna Green" county="Drake's Estero Marin County" latitude="35.964554" location="Santa Cruz" longLatPrecision="1" longitude="122.00185" municipality="Ellwood" pageId="12" pageNumber="565" specimenCode="CASIZ 217592" specimenCount="1" stateProvince="CA" typeStatus="paratype">
<specimenCode id="DB810260FFA5E978D721117FB2ADFE8F" box="[1087,1263,294,321]" collectionCode="CASIZ" pageId="12" pageNumber="565">CASIZ 217592</specimenCode>
,
<specimenCount id="9D216192FFA5E978D7E5117FB3DBFE8F" box="[1275,1433,295,321]" count="1" pageId="12" pageNumber="565" type="generic">one specimen</specimenCount>
,
<location id="8EF8FCC0FFA5E978D3891113B756FEAB" LSID="urn:lsid:plazi:treatment:038E1B0DFFA2E97AD3891685B5C7FDD7:8EF8FCC0FFA5E978D3891113B756FEAB" box="[151,276,331,357]" county="Drake's Estero Marin County" latitude="35.964554" longLatPrecision="1" longitude="122.00185" municipality="Ellwood" name="Santa Cruz" pageId="12" pageNumber="565" stateProvince="CA">Santa Cruz</location>
boat harbor
<geoCoordinate id="EE13CCDCFFA5E978D2B81112B47EFEAB" box="[422,572,330,357]" degrees="35.964552" direction="north" orientation="latitude" pageId="12" pageNumber="565" precision="1" value="35.964554">35.964552°N</geoCoordinate>
<geoCoordinate id="EE13CCDCFFA5E978D14D1113B4B4FEAB" box="[595,758,331,357]" degrees="122.001849" direction="east" orientation="longitude" pageId="12" pageNumber="565" precision="1" value="122.00185">122.001849°E</geoCoordinate>
, Santa Cruz,
<collectingRegion id="49E364F9FFA5E978D0911113B5FBFEAB" box="[911,953,331,357]" pageId="12" pageNumber="565">CA</collectingRegion>
,
<date id="FF998CDBFFA5E978D0DB1113B23EFEAB" box="[965,1148,331,357]" pageId="12" pageNumber="565" value="2017-08-18">
<collectingDate id="EFDD7533FFA5E978D0DB1113B23EFEAB" box="[965,1148,331,357]" pageId="12" pageNumber="565" value="2017-08-18">18 August 2017</collectingDate>
</date>
,
<collectorName id="26D2CFCDFFA5E978D7961113B36AFEAB" box="[1160,1320,331,357]" pageId="12" pageNumber="565">Brenna Green</collectorName>
, collector.
</materialsCitation>
<materialsCitation id="3B4FA046FFA5E978D3891137B7A9FE63" collectingDate="2017-08-18" collectionCode="CASIZ" collectorName="Brenna Green" county="Drake's Estero Marin County" latitude="35.964554" location="Santa Cruz" longLatPrecision="1" longitude="122.00185" municipality="Ellwood" pageId="12" pageNumber="565" specimenCode="CASIZ 217599" specimenCount="1" stateProvince="CA" typeStatus="paratype">
<specimenCode id="DB810260FFA5E978D3891137B705FE47" box="[151,327,366,393]" collectionCode="CASIZ" pageId="12" pageNumber="565">CASIZ 217599</specimenCode>
,
<specimenCount id="9D216192FFA5E978D24D1137B7B3FE47" box="[339,497,367,393]" count="1" pageId="12" pageNumber="565" type="generic">one specimen</specimenCount>
,
<location id="8EF8FCC0FFA5E978D2E21137B43BFE47" LSID="urn:lsid:plazi:treatment:038E1B0DFFA2E97AD3891685B5C7FDD7:8EF8FCC0FFA5E978D2E21137B43BFE47" box="[508,633,367,393]" county="Drake's Estero Marin County" latitude="35.964554" longLatPrecision="1" longitude="122.00185" municipality="Ellwood" name="Santa Cruz" pageId="12" pageNumber="565" stateProvince="CA">Santa Cruz</location>
boat harbor
<geoCoordinate id="EE13CCDCFFA5E978D0141136B5E3FE47" box="[778,929,366,393]" degrees="35.964552" direction="north" orientation="latitude" pageId="12" pageNumber="565" precision="1" value="35.964554">35.964552°N</geoCoordinate>
<geoCoordinate id="EE13CCDCFFA5E978D0A61137B219FE47" box="[952,1115,367,393]" degrees="122.001849" direction="east" orientation="longitude" pageId="12" pageNumber="565" precision="1" value="122.00185">122.001849°E</geoCoordinate>
, Santa Cruz,
<collectingRegion id="49E364F9FFA5E978D7ED1137B35FFE47" box="[1267,1309,367,393]" pageId="12" pageNumber="565">CA</collectingRegion>
,
<date id="FF998CDBFFA5E978D6371137B690FE63" pageId="12" pageNumber="565" value="2017-08-18">
<collectingDate id="EFDD7533FFA5E978D6371137B690FE63" pageId="12" pageNumber="565" value="2017-08-18">18 August 2017</collectingDate>
</date>
,
<collectorName id="26D2CFCDFFA5E978D3C311CBB73EFE63" box="[221,380,403,429]" pageId="12" pageNumber="565">Brenna Green</collectorName>
, collector
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFA5E978D3D911EFB5A1FA9B" pageId="12" pageNumber="565" type="description">
<paragraph id="8B98AA1BFFA5E978D3D911EFB379FC93" blockId="12.[151,1437,150,2013]" pageId="12" pageNumber="565">
<emphasis id="B9537609FFA5E978D3D911EFB785FE1F" bold="true" box="[199,455,439,465]" pageId="12" pageNumber="565">External morphology</emphasis>
(
<figureCitation id="131CB69EFFA5E978D2CB11EFB410FE1F" box="[469,594,439,465]" captionStart="FIGURE 2" captionStartId="13.[151,250,1784,1808]" captionTargetBox="[151,1436,273,1761]" captionTargetId="figure-65@13.[151,1436,273,1761]" captionTargetPageId="13" captionText="FIGURE 2. Living animals. A. Doto urak sp. nov., holotype, CASIZ 182040, San Pedro, California, photo by T. M. Gosliner. B. Doto urak sp. nov., paratype, CASIZ 217592, Santa Cruz, California, photo by T. M. Gosliner. C. Doto urak sp. nov., showing egg mass, Hazard Canyon, San Luis Obispo County, California, photo by Jeff Goddard. D. Doto urak sp. nov., paratypes, CASIZ 056222, Drakes Estero, Marin County, California, photo by Jeff Goddard. E. Doto kwakwak sp. nov., holotype, CASIZ 234657, La Jolla, California, photo by Chloe and Trevor Van Loon. F. Doto kwakwak sp. nov., living animal, Anacapa Island, Ventura County, California, photo by Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.15215500" httpUri="https://zenodo.org/record/15215500/files/figure.png" pageId="12" pageNumber="565">Fig. 2AD</figureCitation>
). The living animals are
<quantity id="4CDF07FEFFA5E978D07D11EFB5FDFE1F" box="[867,959,439,465]" metricMagnitude="-3" metricUnit="m" metricValue="5.5" metricValueMax="7.0" metricValueMin="4.0" pageId="12" pageNumber="565" unit="mm" value="5.5" valueMax="7.0" valueMin="4.0">47 mm</quantity>
in length and the preserved
<typeStatus id="549C14B9FFA5E978D7E611EFB318FE1F" box="[1272,1370,439,465]" pageId="12" pageNumber="565" type="holotype">holotype</typeStatus>
is
<quantity id="4CDF07FEFFA5E978D66711EEB684FE3B" metricMagnitude="-3" metricUnit="m" metricValue="4.5" pageId="12" pageNumber="565" unit="mm" value="4.5">4.5 mm</quantity>
. The body color is translucent white with cream colored ovotestis visible through the body wall. There are a few to many small opaque white spots visible on the tips of the rhinophores not extending to the sheath or the head. The rhinophores extend just slightly above the sheath, which is well elevated from the notum. The apical margin of the rhinophore sheath is flared and of uniform width around the entire margin or wider anteriorly, and a few opaque white spots are occasionally present along the sheath margin. The digestive gland within each ceras ranges from a peach salmon to vibrant pinkish red. There are 47 pairs of cerata with around 15 tubercles on each ceras. The well-spaced tubercles are arranged in about 35 loosely organized whorls and are ovoid in shape. Each tubercle has an opaque white glandular apex. In some individuals (
<figureCitation id="131CB69EFFA5E978D03B128FB539FD3F" box="[805,891,727,753]" captionStart="FIGURE 2" captionStartId="13.[151,250,1784,1808]" captionTargetBox="[151,1436,273,1761]" captionTargetId="figure-65@13.[151,1436,273,1761]" captionTargetPageId="13" captionText="FIGURE 2. Living animals. A. Doto urak sp. nov., holotype, CASIZ 182040, San Pedro, California, photo by T. M. Gosliner. B. Doto urak sp. nov., paratype, CASIZ 217592, Santa Cruz, California, photo by T. M. Gosliner. C. Doto urak sp. nov., showing egg mass, Hazard Canyon, San Luis Obispo County, California, photo by Jeff Goddard. D. Doto urak sp. nov., paratypes, CASIZ 056222, Drakes Estero, Marin County, California, photo by Jeff Goddard. E. Doto kwakwak sp. nov., holotype, CASIZ 234657, La Jolla, California, photo by Chloe and Trevor Van Loon. F. Doto kwakwak sp. nov., living animal, Anacapa Island, Ventura County, California, photo by Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.15215500" httpUri="https://zenodo.org/record/15215500/files/figure.png" pageId="12" pageNumber="565">Fig. 2C</figureCitation>
) cerata have been lost and are regenerating. The pseudobranch (
<figureCitation id="131CB69EFFA5E978D25C12A3B7D8FCDB" box="[322,410,763,789]" captionStart="FIGURE 3" captionStartId="15.[151,250,1943,1967]" captionTargetBox="[222,1369,183,1917]" captionTargetId="figure-22@15.[212,1376,181,1920]" captionTargetPageId="15" captionText="FIGURE 3. Cerata. A. Doto urak sp. nov., paratype, CASIZ 184511. B. Doto columbiana ODonoghue, 1921, CASIZ 189302. C. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 2.5 mm." figureDoi="http://doi.org/10.5281/zenodo.15215502" httpUri="https://zenodo.org/record/15215502/files/figure.png" pageId="12" pageNumber="565">Fig. 3A</figureCitation>
) has up to five palmately lobed branches that are situated near the base of each ceras on the inner side. The anus is located on the right side between the first and second cerata. The genital opening is situated under the first ceras on the right side. The head is rounded with short, rounded extensions visible laterally.
</paragraph>
<paragraph id="8B98AA1BFFA5E978D3D9133FB2F2FBFB" blockId="12.[151,1437,150,2013]" pageId="12" pageNumber="565">
<emphasis id="B9537609FFA5E978D3D9133FB7D0FC4F" bold="true" box="[199,402,871,897]" pageId="12" pageNumber="565">Buccal armature</emphasis>
(
<figureCitation id="131CB69EFFA5E978D2BD133FB7A9FC4F" box="[419,491,871,897]" captionStart="FIGURE 4" captionStartId="16.[151,250,1169,1193]" captionTargetBox="[151,1436,181,1145]" captionTargetId="figure-38@16.[151,1436,181,1145]" captionTargetPageId="16" captionText="FIGURE 4. Doto urak sp. nov., scanning electron micrographs of radula. AC. Paratype, CASIZ 217599. AB. Detail of radular teeth. C. Large section of radular ribbon. D. Paratype, CASIZ 184511. Radular ribbon." figureDoi="http://doi.org/10.5281/zenodo.15215504" httpUri="https://zenodo.org/record/15215504/files/figure.png" pageId="12" pageNumber="565">Fig. 4</figureCitation>
). The buccal mass is small and muscular. The jaws are thin and membranous. The radula is elongate and narrow, consisting of a ribbon of at least 75 teeth. The asymmetrical teeth are simply arched posteriorly with a series of anterior denticles. Each tooth has an elongate central cusp that extends more anteriorly than the lateral denticles. The first two denticles on either side of the central cusp are offset, giving the tooth an asymmetrical appearance. There are two to three pairs of elongate denticles on either side of the central cusp. On the outer side of the tooth there are an additional 12 smaller denticles on either side of the tooth.
</paragraph>
<paragraph id="8B98AA1BFFA5E978D3D91466B5A1FA9B" blockId="12.[151,1437,150,2013]" pageId="12" pageNumber="565">
<emphasis id="B9537609FFA5E978D3D91466B782FB97" bold="true" box="[199,448,1086,1113]" pageId="12" pageNumber="565">Reproductive system</emphasis>
(
<figureCitation id="131CB69EFFA5E978D2CE1467B468FB97" box="[464,554,1086,1113]" captionStart="FIGURE 5" captionStartId="18.[151,250,1908,1932]" captionTargetBox="[247,1337,182,1881]" captionTargetId="figure-38@18.[247,1340,181,1884]" captionTargetPageId="18" captionText="FIGURE 5. Reproductive system. A. Doto urak sp. nov., paratypes, composite of 184511 and 217599, scale = 2.0 mm. B. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 1.5 mm. Abbreviations: am = ampulla, ej = ejaculatory portion of vas deferens, fgm = female gland mass, p = penial sac, pr = prostatic portion of vas deferens, rs = receptaculum seminis, v = vagina." figureDoi="http://doi.org/10.5281/zenodo.15215506" httpUri="https://zenodo.org/record/15215506/files/figure.png" pageId="12" pageNumber="565">Fig. 5A</figureCitation>
). The reproductive system is androdiaulic. The pre-ampullary duct is short and expands into the wide, ovoid ampulla. The ampulla again narrows distally and divides into the oviduct and vas deferens. The oviduct widens into a serial receptaculum seminis that narrows into an elongate convoluted vagina that terminates at the female gonopore adjacent to the penis. The vas deferens widens abruptly into a curved, thickened prostatic portion and narrows into a thin, undulating ejaculatory portion. The distal end of vas deferens enters the large, straight penial sac, which terminates adjacent to the opening of the vagina. The female glands are not well differentiated from each other, but the mucous gland comprises the bulk of the organs and terminates adjacent to the opening of the vagina and penis and has a separate aperture.
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFA5E978D3D91507B4F2FA0F" pageId="12" pageNumber="565" type="etymology">
<paragraph id="8B98AA1BFFA5E978D3D91507B4F2FA0F" blockId="12.[151,1437,150,2013]" pageId="12" pageNumber="565">
<emphasis id="B9537609FFA5E978D3D91507B70AFAB7" bold="true" box="[199,328,1375,1401]" pageId="12" pageNumber="565">Etymology</emphasis>
. The species name urak comes from the southern Ohlone word for salmon, indicating the salmon pink to orange color of the cerata which is a distinctive feature of this species. The Ohlone people were the original inhabitants of much of the range of this species.
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFA5E978D3D91592B2C5FA2B" box="[199,1159,1482,1509]" pageId="12" pageNumber="565" type="distribution">
<paragraph id="8B98AA1BFFA5E978D3D91592B2C5FA2B" blockId="12.[151,1437,150,2013]" box="[199,1159,1482,1509]" pageId="12" pageNumber="565">
<emphasis id="B9537609FFA5E978D3D91592B71BFA2A" bold="true" box="[199,345,1482,1508]" pageId="12" pageNumber="565">Distribution</emphasis>
. Known from San Diego County to Humboldt County,
<collectingRegion id="49E364F9FFA5E978D0D61593B27EFA2B" box="[968,1084,1483,1509]" country="United States of America" name="California" pageId="12" pageNumber="565">California</collectingRegion>
,
<collectingCountry id="F330EA8BFFA5E978D7561593B2C6FA2B" box="[1096,1156,1483,1509]" name="United States of America" pageId="12" pageNumber="565">USA</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFA5E97AD3D915B7B5C7FDD7" lastPageId="14" lastPageNumber="567" pageId="12" pageNumber="565" type="discussion">
<paragraph id="8B98AA1BFFA5E979D3D915B7B728FF37" blockId="12.[151,1437,150,2013]" lastBlockId="13.[151,1437,151,249]" lastPageId="13" lastPageNumber="566" pageId="12" pageNumber="565">
<emphasis id="B9537609FFA5E978D3D915B7B776F9C7" bold="true" box="[199,308,1519,1545]" pageId="12" pageNumber="565">Remarks</emphasis>
.
<taxonomicName id="4C27D198FFA5E978D25D15B7B7F7F9C7" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[323,437,1519,1545]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D25D15B7B7F7F9C7" box="[323,437,1519,1545]" italics="true" pageId="12" pageNumber="565">Doto urak</emphasis>
</taxonomicName>
is sister to
<taxonomicName id="4C27D198FFA5E978D12615B7B4E0F9C6" box="[568,674,1519,1544]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D12615B7B4E0F9C6" box="[568,674,1519,1544]" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
, and both are sister to
<taxonomicName id="4C27D198FFA5E978D0BA15B7B209F9C7" box="[932,1099,1519,1545]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFA5E978D0BA15B7B209F9C7" box="[932,1099,1519,1545]" italics="true" pageId="12" pageNumber="565">D. columbiana</emphasis>
</taxonomicName>
.
<taxonomicName id="4C27D198FFA5E978D74715B7B28EF9C7" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1113,1228,1519,1545]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D74715B7B28EF9C7" box="[1113,1228,1519,1545]" italics="true" pageId="12" pageNumber="565">Doto urak</emphasis>
</taxonomicName>
is externally most similar to
<taxonomicName id="4C27D198FFA5E978D213164BB73BF9E2" box="[269,377,1555,1580]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D213164BB73BF9E2" box="[269,377,1555,1580]" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
and has been confused with this species previously (D.
<bibRefCitation id="EFB6D7EAFFA5E978D71F164BB2E5F9E3" author="Behrens, D." box="[1025,1191,1555,1581]" pageId="12" pageNumber="565" refId="ref12513" refString="Behrens, D. (1980) Pacific Coast Nudibranchs, a Guide to the Opisthobranchs of the Northeastern Pacific. Sea Challengers, Los Osos, California, 112 pp." type="book" year="1980">Behrens, 1980</bibRefCitation>
,
<bibRefCitation id="EFB6D7EAFFA5E978D7AA164BB2ADF9E3" author="Behrens, D." box="[1204,1263,1555,1581]" pageId="12" pageNumber="565" refId="ref12545" refString="Behrens, D. (1991) Pacific Coast Nudibranchs: A Guide to the Opisthobranchs, Alaska to Baja California. 2 nd Edition. Sea Challengers, Monterey, California, 107 pp." type="book" year="1991">1991</bibRefCitation>
; D.
<bibRefCitation id="EFB6D7EAFFA5E978D63C164BB723F99F" author="Behrens, D. &amp; Hermosillo, A." pageId="12" pageNumber="565" refId="ref12581" refString="Behrens, D. &amp; Hermosillo, A. (2005) Eastern Pacific Nudibranchs, a Guide to the Opisthobranchs from Alaska to Central America. 3 rd Edition. Sea Challengers, Monterey, California, 137 pp." type="book" year="2005">Behrens &amp; Hermosillo, 2005</bibRefCitation>
). Both species lack dark pigment on the body.
<taxonomicName id="4C27D198FFA5E978D06A166FB5ACF99E" authorityName="Er.Marcus" authorityYear="1961" box="[884,1006,1591,1616]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyr">
<emphasis id="B9537609FFA5E978D06A166FB5ACF99E" box="[884,1006,1591,1616]" italics="true" pageId="12" pageNumber="565">Doto amyr</emphasis>
</taxonomicName>
a is distinctly longer than
<taxonomicName id="4C27D198FFA5E978D60B166FB329F99F" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1301,1387,1591,1617]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D60B166FB329F99F" box="[1301,1387,1591,1617]" italics="true" pageId="12" pageNumber="565">D. urak</emphasis>
</taxonomicName>
and can reach
<quantity id="4CDF07FEFFA5E978D2171603B71AF9BB" box="[265,344,1627,1653]" metricMagnitude="-2" metricUnit="m" metricValue="1.2" pageId="12" pageNumber="565" unit="mm" value="12.0">12 mm</quantity>
in length. The rhinophores on most
<taxonomicName id="4C27D198FFA5E978D1F01603B51AF9BA" box="[750,856,1627,1652]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D1F01603B51AF9BA" box="[750,856,1627,1652]" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
specimens are proportionately shorter than those of
<taxonomicName id="4C27D198FFA5E978D3891627B6AEF957" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[151,236,1663,1689]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D3891627B6AEF957" box="[151,236,1663,1689]" italics="true" pageId="12" pageNumber="565">D. urak</emphasis>
</taxonomicName>
. The rhinophore sheaths of
<taxonomicName id="4C27D198FFA5E978D13B1627B439F957" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[549,635,1663,1689]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D13B1627B439F957" box="[549,635,1663,1689]" italics="true" pageId="12" pageNumber="565">D. urak</emphasis>
</taxonomicName>
are wide and more flared than those of
<taxonomicName id="4C27D198FFA5E978D7291627B2E3F956" box="[1079,1185,1663,1688]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D7291627B2E3F956" box="[1079,1185,1663,1688]" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
. The ceratal cores and digestive glands of
<taxonomicName id="4C27D198FFA5E978D27016FBB795F972" box="[366,471,1699,1724]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D27016FBB795F972" box="[366,471,1699,1724]" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
are a much lighter white/beige color, while those of
<taxonomicName id="4C27D198FFA5E978D70716FBB224F973" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1049,1126,1699,1725]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D70716FBB224F973" box="[1049,1126,1699,1725]" italics="true" pageId="12" pageNumber="565">D urak</emphasis>
</taxonomicName>
are generally a salmon pink to orange or dark rust/red color. Additionally,
<taxonomicName id="4C27D198FFA5E978D1BD169FB4BBF92F" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[675,761,1735,1761]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D1BD169FB4BBF92F" box="[675,761,1735,1761]" italics="true" pageId="12" pageNumber="565">D. urak</emphasis>
</taxonomicName>
has ceratal tubercles with opaque white glands that are not evident in
<taxonomicName id="4C27D198FFA5E978D21016B3B73AF8CA" box="[270,376,1771,1796]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D21016B3B73AF8CA" box="[270,376,1771,1796]" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
. In
<taxonomicName id="4C27D198FFA5E978D2BA16B3B7B8F8CB" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[420,506,1771,1797]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D2BA16B3B7B8F8CB" box="[420,506,1771,1797]" italics="true" pageId="12" pageNumber="565">D. urak</emphasis>
</taxonomicName>
, the ceratal tubercles are more widely spaced than in
<taxonomicName id="4C27D198FFA5E978D74A16B3B2FDF8CA" box="[1108,1215,1771,1796]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D74A16B3B2FDF8CA" box="[1108,1215,1771,1796]" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
. Also, the tubercles in
<taxonomicName id="4C27D198FFA5E978D3A81757B74FF8E7" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[182,269,1807,1833]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D3A81757B74FF8E7" box="[182,269,1807,1833]" italics="true" pageId="12" pageNumber="565">D. urak</emphasis>
</taxonomicName>
are more rounded and regular in shape than the flatter ovoid tubercles of
<taxonomicName id="4C27D198FFA5E978D77A1757B293F8E6" box="[1124,1233,1807,1832]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D77A1757B293F8E6" box="[1124,1233,1807,1832]" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
. The cerata of
<taxonomicName id="4C27D198FFA5E978D69F1757B69DF882" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D69F1757B69DF882" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
are usually longer, with some exceptions noted (
<bibRefCitation id="EFB6D7EAFFA5E978D1E1176BB5EBF883" author="Agarwal, R. G." box="[767,937,1843,1869]" pageId="12" pageNumber="565" refId="ref12341" refString="Agarwal, R. G. (2019) Hammerhead Doto (Doto amyra). iNaturalist. Available from: https://www.inaturalist.org/observations/19824870 (accessed 27 June 2024)" type="url" year="2019">Agarwal, 2019</bibRefCitation>
) and also have more rows of tubercles. In
<taxonomicName id="4C27D198FFA5E978D69F176BB69DF8BE" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D69F176BB69DF8BE" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
, the pseudobranch extends most of the length of the ceras with a few short lateral branches (
<bibRefCitation id="EFB6D7EAFFA5E978D7F3170FB3DAF8BF" author="Marcus, Er." box="[1261,1432,1879,1905]" pageId="12" pageNumber="565" pagination="1 - 85" refId="ref14444" refString="Marcus, Er. (1961 a) Opisthobranch mollusks from California. The Veliger, 3, 1-85, pls. 1 - 10." type="journal article" year="1961">Marcus, 1961a</bibRefCitation>
, figs. 131, 132, 136) whereas
<taxonomicName id="4C27D198FFA5E978D2FB1723B479F85B" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[485,571,1915,1941]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA5E978D2FB1723B479F85B" box="[485,571,1915,1941]" italics="true" pageId="12" pageNumber="565">D. urak</emphasis>
</taxonomicName>
has a palmately branched pseudobranch that is found basally on each ceras (
<figureCitation id="131CB69EFFA5E978D38117C7B6BBF877" box="[159,249,1951,1977]" captionStart="FIGURE 3" captionStartId="15.[151,250,1943,1967]" captionTargetBox="[222,1369,183,1917]" captionTargetId="figure-22@15.[212,1376,181,1920]" captionTargetPageId="15" captionText="FIGURE 3. Cerata. A. Doto urak sp. nov., paratype, CASIZ 184511. B. Doto columbiana ODonoghue, 1921, CASIZ 189302. C. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 2.5 mm." figureDoi="http://doi.org/10.5281/zenodo.15215502" httpUri="https://zenodo.org/record/15215502/files/figure.png" pageId="12" pageNumber="565">Fig. 3A</figureCitation>
). Previous literature on both species have indicated that
<taxonomicName id="4C27D198FFA5E978D07117C7B598F876" box="[879,986,1951,1976]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="565" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA5E978D07117C7B598F876" box="[879,986,1951,1976]" italics="true" pageId="12" pageNumber="565">D. amyra</emphasis>
</taxonomicName>
is more transparent than its counterpart, but some images suggest otherwise (
<bibRefCitation id="EFB6D7EAFFA5E978D15E179BB54AF813" author="McDonald, G." box="[576,776,1987,2013]" pageId="12" pageNumber="565" refId="ref14731" refString="McDonald, G. (2017) Mollusca: Gastropoda: Opisthobranchia: Nudibranchia: Dendronotina. In: Intertidal Invertebrates of the Monterey Bay Area. Available from: https://people.ucsc.edu/~mcduck/DreamWeaver/Mollusca/Nudibranchs/Dendronotina.html (accessed 30 August 2024)" type="url" year="2017">
<collectingCountry id="F330EA8BFFA5E978D15E179BB4FEF813" box="[576,700,1987,2013]" name="Heard Island and McDonald Islands" pageId="12" pageNumber="565">McDonald</collectingCountry>
, 2017
</bibRefCitation>
). The most significant difference is in the developmental biology of the two species:
<taxonomicName id="4C27D198FFA4E979D2D010CFB402FF7F" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[462,576,151,177]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="566" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA4E979D2D010CFB402FF7F" box="[462,576,151,177]" italics="true" pageId="13" pageNumber="566">Doto urak</emphasis>
</taxonomicName>
lays eggs averaging 76 µm in diameter and has planktotrophic larvae, while
<taxonomicName id="4C27D198FFA4E979D38910E3B740FF1A" box="[151,258,187,212]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="566" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA4E979D38910E3B740FF1A" box="[151,258,187,212]" italics="true" pageId="13" pageNumber="566">D. amyra</emphasis>
</taxonomicName>
lays eggs averaging 152 µm in diameter and has lecithotrophic larvae (J. H. R.
<bibRefCitation id="EFB6D7EAFFA4E979D79F10E3B36FFF1B" author="Goddard, J. H. R." box="[1153,1325,187,213]" pageId="13" pageNumber="566" pagination="43 - 54" refId="ref13129" refString="Goddard, J. H. R. (1996) Lecithotrophic development in Doto amyra (Nudibranchia: Dendronotacea), with a review of developmental mode in the genus. Veliger, 39 (1), 43-54." type="journal article" year="1996">Goddard, 1996</bibRefCitation>
;
<bibRefCitation id="EFB6D7EAFFA4E979D62710E3B71CFF37" author="Shipman, C. &amp; Gosliner, T." pageId="13" pageNumber="566" pagination="57 - 101" refId="ref15905" refString="Shipman, C. &amp; Gosliner, T. (2015) Molecular and morphological systematics of Doto Oken, 1851 (Gastropoda: Heterobranchia) with descriptions of five new species and a new genus. Zootaxa, 3973 (1) 57-101. https://doi.org/10.11646/zootaxa.3973.1.2" type="journal article" year="2015">Shipman &amp; Gosliner, 2015</bibRefCitation>
).
</paragraph>
<caption id="DF58FA93FFA4E979D38916A0B4E8F80A" ID-DOI="http://doi.org/10.5281/zenodo.15215500" ID-Zenodo-Dep="15215500" httpUri="https://zenodo.org/record/15215500/files/figure.png" pageId="13" pageNumber="566" startId="13.[151,250,1784,1808]" targetBox="[151,1436,273,1761]" targetPageId="13" targetType="figure">
<paragraph id="8B98AA1BFFA4E979D38916A0B4E8F80A" blockId="13.[151,1437,1784,1989]" pageId="13" pageNumber="566">
<emphasis id="B9537609FFA4E979D38916A0B74EF8DE" bold="true" box="[151,268,1784,1808]" pageId="13" pageNumber="566">FIGURE 2</emphasis>
. Living animals.
<emphasis id="B9537609FFA4E979D2A416A0B78EF8DE" bold="true" box="[442,460,1784,1808]" pageId="13" pageNumber="566">A</emphasis>
.
<taxonomicName id="4C27D198FFA4E979D2C616A1B47FF8DE" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[472,573,1784,1808]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="566" phylum="Mollusca" rank="species" species="urak" status="sp. nov.">
<emphasis id="B9537609FFA4E979D2C616A1B47FF8DE" box="[472,573,1784,1808]" italics="true" pageId="13" pageNumber="566">Doto urak</emphasis>
</taxonomicName>
<emphasis id="B9537609FFA4E979D15D16A1B4D0F8DF" bold="true" box="[579,658,1785,1809]" pageId="13" pageNumber="566">
<taxonomicNameLabel id="A260CB72FFA4E979D15D16A1B4D0F8DF" box="[579,658,1785,1809]" pageId="13" pageNumber="566" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, holotype, CASIZ 182040, San Pedro, California, photo by T. M. Gosliner.
<emphasis id="B9537609FFA4E979D69B16A1B3D4F8DF" bold="true" box="[1413,1430,1785,1809]" pageId="13" pageNumber="566">B</emphasis>
.
<taxonomicName id="4C27D198FFA4E979D3891745B6BCF8FA" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[151,254,1820,1844]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="566" phylum="Mollusca" rank="species" species="urak" status="sp. nov.">
<emphasis id="B9537609FFA4E979D3891745B6BCF8FA" box="[151,254,1820,1844]" italics="true" pageId="13" pageNumber="566">Doto urak</emphasis>
</taxonomicName>
<emphasis id="B9537609FFA4E979D21B1745B717F8FB" bold="true" box="[261,341,1821,1845]" pageId="13" pageNumber="566">
<taxonomicNameLabel id="A260CB72FFA4E979D21B1745B717F8FB" box="[261,341,1821,1845]" pageId="13" pageNumber="566" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, paratype, CASIZ 217592, Santa Cruz, California, photo by T. M. Gosliner.
<emphasis id="B9537609FFA4E979D7421744B22CF8FA" bold="true" box="[1116,1134,1820,1844]" pageId="13" pageNumber="566">C</emphasis>
.
<taxonomicName id="4C27D198FFA4E979D7651745B2A0F8FA" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1147,1250,1820,1844]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="566" phylum="Mollusca" rank="species" species="urak" status="sp. nov.">
<emphasis id="B9537609FFA4E979D7651745B2A0F8FA" box="[1147,1250,1820,1844]" italics="true" pageId="13" pageNumber="566">Doto urak</emphasis>
</taxonomicName>
<emphasis id="B9537609FFA4E979D7F71745B37BF8FB" bold="true" box="[1257,1337,1821,1845]" pageId="13" pageNumber="566">
<taxonomicNameLabel id="A260CB72FFA4E979D7F71745B37BF8FB" box="[1257,1337,1821,1845]" pageId="13" pageNumber="566" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, showing egg mass, Hazard Canyon, San Luis Obispo County, California, photo by Jeff Goddard.
<emphasis id="B9537609FFA4E979D7221719B20CF897" bold="true" box="[1084,1102,1857,1881]" pageId="13" pageNumber="566">D</emphasis>
.
<taxonomicName id="4C27D198FFA4E979D7401719B28AF896" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1118,1224,1856,1880]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="566" phylum="Mollusca" rank="species" species="urak" status="sp. nov.">
<emphasis id="B9537609FFA4E979D7401719B28AF896" box="[1118,1224,1856,1880]" italics="true" pageId="13" pageNumber="566">Doto urak</emphasis>
</taxonomicName>
<emphasis id="B9537609FFA4E979D7CF1719B367F897" bold="true" box="[1233,1317,1857,1881]" pageId="13" pageNumber="566">
<taxonomicNameLabel id="A260CB72FFA4E979D7CF1719B367F897" box="[1233,1317,1857,1881]" pageId="13" pageNumber="566" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, paratypes, CASIZ 056222, Drakes Estero, Marin County, California, photo by Jeff Goddard.
<emphasis id="B9537609FFA4E979D0C4173DB5A9F8B3" bold="true" box="[986,1003,1893,1917]" pageId="13" pageNumber="566">E</emphasis>
.
<taxonomicName id="4C27D198FFA4E979D0E9173DB2CEF8B2" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1015,1164,1892,1916]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="566" phylum="Mollusca" rank="species" species="kwakwak" status="sp. nov.">
<emphasis id="B9537609FFA4E979D0E9173DB2CEF8B2" box="[1015,1164,1892,1916]" italics="true" pageId="13" pageNumber="566">Doto kwakwak</emphasis>
</taxonomicName>
<emphasis id="B9537609FFA4E979D78C173DB2A0F8B3" bold="true" box="[1170,1250,1893,1917]" pageId="13" pageNumber="566">
<taxonomicNameLabel id="A260CB72FFA4E979D78C173DB2A0F8B3" box="[1170,1250,1893,1917]" pageId="13" pageNumber="566" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, holotype, CASIZ 234657, La Jolla, California, photo by Chloe and Trevor Van Loon.
<emphasis id="B9537609FFA4E979D05317D1B51EF86F" bold="true" box="[845,860,1929,1953]" pageId="13" pageNumber="566">F</emphasis>
.
<taxonomicName id="4C27D198FFA4E979D07417D1B242F86E" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[874,1024,1928,1952]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="566" phylum="Mollusca" rank="species" species="kwakwak" status="sp. nov.">
<emphasis id="B9537609FFA4E979D07417D1B242F86E" box="[874,1024,1928,1952]" italics="true" pageId="13" pageNumber="566">Doto kwakwak</emphasis>
</taxonomicName>
<emphasis id="B9537609FFA4E979D71917D1B21AF86F" bold="true" box="[1031,1112,1929,1953]" pageId="13" pageNumber="566">
<taxonomicNameLabel id="A260CB72FFA4E979D71917D1B21AF86F" box="[1031,1112,1929,1953]" pageId="13" pageNumber="566" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, living animal, Anacapa Island, Ventura County, California, photo by Scott Johnson.
</paragraph>
</caption>
<paragraph id="8B98AA1BFFA7E97AD3D910CFB4DDFE47" blockId="14.[151,1437,151,537]" pageId="14" pageNumber="567">
Internally, the radular teeth of most of the northeastern Pacific
<taxonomicName id="4C27D198FFA7E97AD06510CFB24BFF7F" box="[891,1033,151,177]" pageId="14" pageNumber="567">
<emphasis id="B9537609FFA7E97AD06510CFB5F1FF7E" box="[891,947,151,176]" italics="true" pageId="14" pageNumber="567">Doto</emphasis>
species
</taxonomicName>
are very similar in shape and appear to all have a certain degree of asymmetry (
<bibRefCitation id="EFB6D7EAFFA7E97AD16810E3B560FF1B" author="Marcus, Er." box="[630,802,187,213]" pageId="14" pageNumber="567" pagination="1 - 85" refId="ref14444" refString="Marcus, Er. (1961 a) Opisthobranch mollusks from California. The Veliger, 3, 1-85, pls. 1 - 10." type="journal article" year="1961">Marcus, 1961a</bibRefCitation>
). The reproductive systems are also similar in form with a serial receptaculum seminis (
<bibRefCitation id="EFB6D7EAFFA7E97AD2E41087B4E5FF37" author="Marcus, Er." box="[506,679,223,249]" pageId="14" pageNumber="567" pagination="1 - 85" refId="ref14444" refString="Marcus, Er. (1961 a) Opisthobranch mollusks from California. The Veliger, 3, 1-85, pls. 1 - 10." type="journal article" year="1961">Marcus, 1961a</bibRefCitation>
), but do appear to differ in several details of their anatomy. In
<taxonomicName id="4C27D198FFA7E97AD69F1087B688FED3" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA7E97AD69F1087B688FED3" italics="true" pageId="14" pageNumber="567">D. urak</emphasis>
</taxonomicName>
, the penial sac is more elongate than that of
<taxonomicName id="4C27D198FFA7E97AD1D8115BB573FED2" box="[710,817,259,284]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA7E97AD1D8115BB573FED2" box="[710,817,259,284]" italics="true" pageId="14" pageNumber="567">D. amyra</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFB6D7EAFFA7E97AD05E115BB5AEFED3" author="Marcus, Er." box="[832,1004,259,285]" pageId="14" pageNumber="567" pagination="1 - 85" refId="ref14444" refString="Marcus, Er. (1961 a) Opisthobranch mollusks from California. The Veliger, 3, 1-85, pls. 1 - 10." type="journal article" year="1961">Marcus, 1961a</bibRefCitation>
: figs. 134, 138 as
<taxonomicName id="4C27D198FFA7E97AD7A4115BB355FED2" authorityName="Marcus" authorityYear="1961" box="[1210,1303,259,284]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="wara">
<emphasis id="B9537609FFA7E97AD7A4115BB355FED2" box="[1210,1303,259,284]" italics="true" pageId="14" pageNumber="567">D. wara</emphasis>
</taxonomicName>
, fig. 146 as
<taxonomicName id="4C27D198FFA7E97AD389117FB6BDFE8F" authorityName="Marcus" authorityYear="1961" box="[151,255,295,321]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="ganda">
<emphasis id="B9537609FFA7E97AD389117FB6BDFE8F" box="[151,255,295,321]" italics="true" pageId="14" pageNumber="567">D. ganda</emphasis>
</taxonomicName>
), while the prostate is not as convoluted as in
<taxonomicName id="4C27D198FFA7E97AD01B117FB532FE8E" box="[773,880,295,320]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA7E97AD01B117FB532FE8E" box="[773,880,295,320]" italics="true" pageId="14" pageNumber="567">D. amyra</emphasis>
</taxonomicName>
. Also, there is a distinct muscular sphincter at the distal end of the vagina in
<taxonomicName id="4C27D198FFA7E97AD2A71113B41FFEAB" box="[441,605,331,357]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFA7E97AD2A71113B41FFEAB" box="[441,605,331,357]" italics="true" pageId="14" pageNumber="567">D. columbiana</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFB6D7EAFFA7E97AD1751113B557FEAB" author="Marcus, Er." box="[619,789,331,357]" pageId="14" pageNumber="567" pagination="1 - 85" refId="ref14444" refString="Marcus, Er. (1961 a) Opisthobranch mollusks from California. The Veliger, 3, 1-85, pls. 1 - 10." type="journal article" year="1961">Marcus, 1961a</bibRefCitation>
: fig. 129) and
<taxonomicName id="4C27D198FFA7E97AD0AC1113B29AFEAB" authority="(Marcus, 1961 a)" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[946,1240,331,357]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA7E97AD0AC1113B259FEAA" box="[946,1051,331,356]" italics="true" pageId="14" pageNumber="567">D. amyra</emphasis>
(
<bibRefCitation id="EFB6D7EAFFA7E97AD7361113B292FEAB" author="Marcus, Er." box="[1064,1232,331,357]" pageId="14" pageNumber="567" pagination="1 - 85" refId="ref14444" refString="Marcus, Er. (1961 a) Opisthobranch mollusks from California. The Veliger, 3, 1-85, pls. 1 - 10." type="journal article" year="1961">Marcus, 1961a</bibRefCitation>
)
</taxonomicName>
that is apparently absent in
<taxonomicName id="4C27D198FFA7E97AD21C1137B71AFE47" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[258,344,367,393]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA7E97AD21C1137B71AFE47" box="[258,344,367,393]" italics="true" pageId="14" pageNumber="567">D. urak</emphasis>
</taxonomicName>
and
<taxonomicName id="4C27D198FFA7E97AD2901137B4D8FE47" authority="(Marcus, 1961 a)" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[398,666,367,393]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFA7E97AD2901137B79AFE47" box="[398,472,367,393]" italics="true" pageId="14" pageNumber="567">D. kya</emphasis>
(
<bibRefCitation id="EFB6D7EAFFA7E97AD2F81137B4D0FE47" author="Marcus, Er." box="[486,658,367,393]" pageId="14" pageNumber="567" pagination="1 - 85" refId="ref14444" refString="Marcus, Er. (1961 a) Opisthobranch mollusks from California. The Veliger, 3, 1-85, pls. 1 - 10." type="journal article" year="1961">Marcus, 1961a</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="8B98AA1BFFA7E97AD3D911CBB5C7FDD7" blockId="14.[151,1437,151,537]" pageId="14" pageNumber="567">
<taxonomicName id="4C27D198FFA7E97AD3D911CBB778FE63" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[199,314,403,429]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA7E97AD3D911CBB778FE63" box="[199,314,403,429]" italics="true" pageId="14" pageNumber="567">Doto urak</emphasis>
</taxonomicName>
is molecularly distinct (BI = 1, ML = 100) and is a single distinct species according to each species delimitation method undertaken.
<taxonomicName id="4C27D198FFA7E97AD11511EFB422FE1F" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[523,608,439,465]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA7E97AD11511EFB422FE1F" box="[523,608,439,465]" italics="true" pageId="14" pageNumber="567">D. urak</emphasis>
</taxonomicName>
has an intraspecific variation
<emphasis id="B9537609FFA7E97AD0AC11EFB582FE1E" box="[946,960,439,464]" italics="true" pageId="14" pageNumber="567">p</emphasis>
-distance of 0.001.90%. The interspecific
<emphasis id="B9537609FFA7E97AD3891183B6E7FE3A" box="[151,165,475,500]" italics="true" pageId="14" pageNumber="567">p</emphasis>
-distances between
<taxonomicName id="4C27D198FFA7E97AD2971183B7A0FE3B" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[393,482,475,501]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFA7E97AD2971183B7A0FE3B" box="[393,482,475,501]" italics="true" pageId="14" pageNumber="567">D. urak</emphasis>
</taxonomicName>
and the closest sister species are: 4.956.69% to
<taxonomicName id="4C27D198FFA7E97AD72F1183B2FEFE3A" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[1073,1212,475,500]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFA7E97AD72F1183B2FEFE3A" box="[1073,1212,475,500]" italics="true" pageId="14" pageNumber="567">Doto amyra</emphasis>
</taxonomicName>
, 7.909.09% to
<taxonomicName id="4C27D198FFA7E97AD69F1183B759FDD7" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFA7E97AD69F1183B759FDD7" italics="true" pageId="14" pageNumber="567">D. columbiana</emphasis>
</taxonomicName>
, 10.4912.68% to
<taxonomicName id="4C27D198FFA7E97AD2EE11A7B478FDD7" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[496,570,511,537]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFA7E97AD2EE11A7B478FDD7" box="[496,570,511,537]" italics="true" pageId="14" pageNumber="567">D. kya</emphasis>
</taxonomicName>
, and 6.297.54% to
<emphasis id="B9537609FFA7E97AD03C11A7B57CFDD6" box="[802,830,511,536]" italics="true" pageId="14" pageNumber="567">D.</emphasis>
sp. B.
</paragraph>
</subSubSection>
</treatment>
</document>

View file

@ -0,0 +1,604 @@
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<taxonomicName id="4C27D198FFA7E97AD3891233B4C3FD4A" ID-CoL="KTVQQ" authority="Gosliner &amp; Adayapalam" authorityName="Gosliner &amp; Adayapalam" authorityYear="2025" box="[151,641,618,645]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="kwakwak" status="sp. nov.">
<emphasis id="B9537609FFA7E97AD3891233B70BFD4B" bold="true" box="[151,329,618,645]" italics="true" pageId="14" pageNumber="567">Doto kwakwak</emphasis>
Gosliner &amp; Adayapalam
</taxonomicName>
<taxonomicNameLabel id="A260CB72FFA7E97AD1971233B4A4FD4B" box="[649,742,619,645]" pageId="14" pageNumber="567" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFA7E97AD38912D7B7DEFD03" pageId="14" pageNumber="567" type="description">
<paragraph id="8B98AA1BFFA7E97AD38912D7B597FD67" blockId="14.[151,981,618,717]" box="[151,981,655,681]" pageId="14" pageNumber="567">
<uri id="FFB6A619FFA7E97AD38912D7B597FD67" box="[151,981,655,681]" pageId="14" pageNumber="567">
urn:lsid:zoobank.org:act:
<uuid id="FF8190CEFFA7E97AD2B212D7B597FD67" box="[428,981,655,681]" pageId="14" pageNumber="567">F10B38D1-ECA4-4BB6-BF37-9ECB962834CB</uuid>
</uri>
</paragraph>
<paragraph id="8B98AA1BFFA7E97AD38912EBB7DEFD03" blockId="14.[151,981,618,717]" box="[151,412,690,717]" pageId="14" pageNumber="567">
(
<figureCitation id="131CB69EFFA7E97AD38112EBB75EFD03" box="[159,284,691,717]" captionStart="FIGURE 2" captionStartId="13.[151,250,1784,1808]" captionTargetBox="[151,1436,273,1761]" captionTargetId="figure-65@13.[151,1436,273,1761]" captionTargetPageId="13" captionText="FIGURE 2. Living animals. A. Doto urak sp. nov., holotype, CASIZ 182040, San Pedro, California, photo by T. M. Gosliner. B. Doto urak sp. nov., paratype, CASIZ 217592, Santa Cruz, California, photo by T. M. Gosliner. C. Doto urak sp. nov., showing egg mass, Hazard Canyon, San Luis Obispo County, California, photo by Jeff Goddard. D. Doto urak sp. nov., paratypes, CASIZ 056222, Drakes Estero, Marin County, California, photo by Jeff Goddard. E. Doto kwakwak sp. nov., holotype, CASIZ 234657, La Jolla, California, photo by Chloe and Trevor Van Loon. F. Doto kwakwak sp. nov., living animal, Anacapa Island, Ventura County, California, photo by Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.15215500" httpUri="https://zenodo.org/record/15215500/files/figure.png" pageId="14" pageNumber="567">Figs. 2EF</figureCitation>
,
<figureCitation id="131CB69EFFA7E97AD23912EBB708FD03" box="[295,330,691,717]" captionStart="FIGURE 3" captionStartId="15.[151,250,1943,1967]" captionTargetBox="[222,1369,183,1917]" captionTargetId="figure-22@15.[212,1376,181,1920]" captionTargetPageId="15" captionText="FIGURE 3. Cerata. A. Doto urak sp. nov., paratype, CASIZ 184511. B. Doto columbiana ODonoghue, 1921, CASIZ 189302. C. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 2.5 mm." figureDoi="http://doi.org/10.5281/zenodo.15215502" httpUri="https://zenodo.org/record/15215502/files/figure.png" pageId="14" pageNumber="567">3C</figureCitation>
,
<figureCitation id="131CB69EFFA7E97AD24812EAB73AFD03" box="[342,376,690,717]" captionStart="FIGURE 5" captionStartId="18.[151,250,1908,1932]" captionTargetBox="[247,1337,182,1881]" captionTargetId="figure-38@18.[247,1340,181,1884]" captionTargetPageId="18" captionText="FIGURE 5. Reproductive system. A. Doto urak sp. nov., paratypes, composite of 184511 and 217599, scale = 2.0 mm. B. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 1.5 mm. Abbreviations: am = ampulla, ej = ejaculatory portion of vas deferens, fgm = female gland mass, p = penial sac, pr = prostatic portion of vas deferens, rs = receptaculum seminis, v = vagina." figureDoi="http://doi.org/10.5281/zenodo.15215506" httpUri="https://zenodo.org/record/15215506/files/figure.png" pageId="14" pageNumber="567">5B</figureCitation>
,
<figureCitation id="131CB69EFFA7E97AD29B12EBB7D6FD03" box="[389,404,691,717]" captionStart="FIGURE 6" captionStartId="19.[151,250,1184,1208]" captionTargetBox="[151,1436,181,1160]" captionTargetId="figure-22@19.[151,1436,181,1160]" captionTargetPageId="19" captionText="FIGURE 6. Doto kwakwak sp. nov., holotype, CASIZ 234657, scanning electron micrographs of radula. A. Segments of the radular ribbon. BC. Dorsolateral view of radular teeth. D. Lateral view of radular teeth." figureDoi="http://doi.org/10.5281/zenodo.15215508" httpUri="https://zenodo.org/record/15215508/files/figure.png" pageId="14" pageNumber="567">6</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFA7E97AD38912A0B4EDFCC1" box="[151,687,759,783]" pageId="14" pageNumber="567" type="reference_group">
<paragraph id="8B98AA1BFFA7E97AD38912A0B4EDFCC1" blockId="14.[151,687,759,783]" box="[151,687,759,783]" pageId="14" pageNumber="567">
<treatmentCitationGroup id="AB378D35FFA7E97AD38912A0B4EDFCC1" box="[151,687,759,783]" pageId="14" pageNumber="567">
<taxonomicName id="4C27D198FFA7E97AD38912A0B6BDFCC1" box="[151,255,759,783]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="567" phylum="Mollusca" rank="species" species="undefined-4">
<emphasis id="B9537609FFA7E97AD38912A0B688FCC1" box="[151,202,760,783]" italics="true" pageId="14" pageNumber="567">Doto</emphasis>
sp. 4
</taxonomicName>
<treatmentCitation id="0A868C0AFFA7E97AD20412AFB45BFCC1" author="Behrens, D. W. &amp; Fletcher, K. &amp; Jensen, G." box="[282,537,759,783]" page="114" pageId="14" pageNumber="567" year="2022">
<bibRefCitation id="EFB6D7EAFFA7E97AD20412AFB45BFCC1" author="Behrens, D. W. &amp; Fletcher, K. &amp; Jensen, G." box="[282,537,759,783]" pageId="14" pageNumber="567" refId="ref12622" refString="Behrens, D. W., Fletcher, K. &amp; Jensen, G. (2022) Nudibranchs &amp; Sea Slugs of the Eastern Pacific. MolaMarine, Bremerton, Washington, 170 pp." type="book" year="2022">
Behrens
<emphasis id="B9537609FFA7E97AD26D12A0B7E4FCC1" box="[371,422,759,783]" italics="true" pageId="14" pageNumber="567">et al.</emphasis>
, 2022: 114
</bibRefCitation>
</treatmentCitation>
, lower photos.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFA7E97AD3891365B5A0FCB5" pageId="14" pageNumber="567" type="materials_examined">
<paragraph id="8B98AA1BFFA7E97AD3891365B5A0FCB5" blockId="14.[151,1437,829,2008]" pageId="14" pageNumber="567">
<emphasis id="B9537609FFA7E97AD3891365B705FC99" bold="true" box="[151,327,829,855]" pageId="14" pageNumber="567">Type material.</emphasis>
<materialsCitation id="3B4FA046FFA7E97AD24E1365B5A0FCB5" collectingDate="2022-02-27" collectionCode="CASIZ" collectorName="Chloe &amp; Trevor Van Loon" latitude="32.843075" location="Cuvier Park" longLatPrecision="1" longitude="117.28189" municipality="La Jolla" pageId="14" pageNumber="567" specimenCode="CASIZ 234657" specimenCount="1" stateProvince="California" typeStatus="holotype">
<emphasis id="B9537609FFA7E97AD24E1365B7FFFC99" bold="true" box="[336,445,829,855]" pageId="14" pageNumber="567">
<typeStatus id="549C14B9FFA7E97AD24E1365B7FFFC99" box="[336,445,829,855]" pageId="14" pageNumber="567" type="holotype">Holotype</typeStatus>
</emphasis>
.
<specimenCode id="DB810260FFA7E97AD2D21365B43FFC96" box="[460,637,829,856]" collectionCode="CASIZ" pageId="14" pageNumber="567">CASIZ 234657</specimenCode>
,
<quantity id="4CDF07FEFFA7E97AD1941365B4A0FC96" box="[650,738,829,856]" metricMagnitude="-3" metricUnit="m" metricValue="4.5" pageId="14" pageNumber="567" unit="mm" value="4.5">4.5 mm</quantity>
preserved, off
<location id="8EF8FCC0FFA7E97AD08E1365B25AFC99" LSID="urn:lsid:plazi:treatment:038E1B0DFFA7E965D3891233B745FD2B:8EF8FCC0FFA7E97AD08E1365B25AFC99" box="[912,1048,829,855]" latitude="32.843075" longLatPrecision="1" longitude="117.28189" municipality="La Jolla" name="Cuvier Park" pageId="14" pageNumber="567" stateProvince="California">Cuvier Park</location>
<geoCoordinate id="EE13CCDCFFA7E97AD73F1365B2F5FC99" box="[1057,1207,829,855]" degrees="32.843073" direction="north" orientation="latitude" pageId="14" pageNumber="567" precision="1" value="32.843075">32.843073°N</geoCoordinate>
<geoCoordinate id="EE13CCDCFFA7E97AD7D11365B333FC99" box="[1231,1393,829,855]" degrees="117.281888" direction="east" orientation="longitude" pageId="14" pageNumber="567" precision="1" value="117.28189">117.281888°E</geoCoordinate>
,
<collectingMunicipality id="6BFC3061FFA7E97AD6611366B68DFCB5" pageId="14" pageNumber="567">La Jolla</collectingMunicipality>
,
<collectingRegion id="49E364F9FFA7E97AD3C41339B70CFCB5" box="[218,334,865,891]" country="United States of America" name="California" pageId="14" pageNumber="567">California</collectingRegion>
,
<date id="FF998CDBFFA7E97AD2441339B466FCB5" box="[346,548,865,891]" pageId="14" pageNumber="567" value="2022-02-27">
<collectingDate id="EFDD7533FFA7E97AD2441339B466FCB5" box="[346,548,865,891]" pageId="14" pageNumber="567" value="2022-02-27">27 February 2022</collectingDate>
</date>
,
<collectorName id="26D2CFCDFFA7E97AD1311339B430FCB5" box="[559,626,865,891]" pageId="14" pageNumber="567">Chloe</collectorName>
and
<collectorName id="26D2CFCDFFA7E97AD1B9133AB525FCB2" box="[679,871,866,892]" pageId="14" pageNumber="567">Trevor Van Loon</collectorName>
collectors.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFA7E97AD3D913DDB4FBF8A2" pageId="14" pageNumber="567" type="description">
<paragraph id="8B98AA1BFFA7E97AD3D913DDB455FA59" blockId="14.[151,1437,829,2008]" pageId="14" pageNumber="567">
<emphasis id="B9537609FFA7E97AD3D913DDB785FC51" bold="true" box="[199,455,901,927]" pageId="14" pageNumber="567">External morphology</emphasis>
(
<figureCitation id="131CB69EFFA7E97AD2C813DDB40EFC51" box="[470,588,901,927]" captionStart="FIGURE 2" captionStartId="13.[151,250,1784,1808]" captionTargetBox="[151,1436,273,1761]" captionTargetId="figure-65@13.[151,1436,273,1761]" captionTargetPageId="13" captionText="FIGURE 2. Living animals. A. Doto urak sp. nov., holotype, CASIZ 182040, San Pedro, California, photo by T. M. Gosliner. B. Doto urak sp. nov., paratype, CASIZ 217592, Santa Cruz, California, photo by T. M. Gosliner. C. Doto urak sp. nov., showing egg mass, Hazard Canyon, San Luis Obispo County, California, photo by Jeff Goddard. D. Doto urak sp. nov., paratypes, CASIZ 056222, Drakes Estero, Marin County, California, photo by Jeff Goddard. E. Doto kwakwak sp. nov., holotype, CASIZ 234657, La Jolla, California, photo by Chloe and Trevor Van Loon. F. Doto kwakwak sp. nov., living animal, Anacapa Island, Ventura County, California, photo by Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.15215500" httpUri="https://zenodo.org/record/15215500/files/figure.png" pageId="14" pageNumber="567">Fig. 2EF</figureCitation>
). The living animal has a translucent white body color with a cream colored ovotestis visible through the body. There are small opaque white spots visible along the margins of the foot, along the margin of the rhinophore sheath, and on the entire length of the rhinophore. The dorsal notum and sides of the body have numerous scattered ovoid saffron yellowish orange spots, some of which have irregular borders and are slightly raised. The digestive gland within each ceras is light brownish with opaque white glands present within the tubercles on the cerata. There are 7 pairs of cerata with 4 whorls of elongate tubercles with bulbous tips. The size of the tubercles varies, but in general the more basal ones are smaller with the largest being the single terminal tubercle. The cerata often lack pseudobranchs but in one large ceras the pseudobranch consisted of a single linear axis without lateral branches. The rhinophoral sheaths are well-elevated from the notum with an extension on the anterior side. The head is broader than the rest of the body with a pair of curved lobes situated laterally. The anal papilla is situated dorsally, posterior to the second right ceras and genital openings on right side of body, immediately ventral to the first ceras. On the head of
<specimenCount id="9D216192FFA7E97AD2A1154AB419FAE5" box="[447,603,1297,1324]" count="1" pageId="14" pageNumber="567" type="generic">one specimen</specimenCount>
, photographed from off the east end of Santa Cruz Island, there is a brown patch of what appears to be a subcutaneous pigmentation (
<bibRefCitation id="EFB6D7EAFFA7E97AD03B156DB5EBFA81" author="Klug, D." box="[805,937,1333,1359]" pageId="14" pageNumber="567" refId="ref14109" refString="Klug, D. (2019) Genus Doto. iNaturalist. Available from: https://www.inaturalist.org/observations/35397064 (accessed 27 June 2024)" type="url" year="2019">Klug, 2019</bibRefCitation>
). An additional specimen, from off the Palos Verdes Peninsula and included in
<bibRefCitation id="EFB6D7EAFFA7E97AD10A1501B540FABD" author="Behrens, D. W. &amp; Fletcher, K. &amp; Jensen, G." box="[532,770,1369,1395]" pageId="14" pageNumber="567" refId="ref12622" refString="Behrens, D. W., Fletcher, K. &amp; Jensen, G. (2022) Nudibranchs &amp; Sea Slugs of the Eastern Pacific. MolaMarine, Bremerton, Washington, 170 pp." type="book" year="2022">
Behrens
<emphasis id="B9537609FFA7E97AD1691502B4F2FABD" box="[631,688,1369,1395]" italics="true" pageId="14" pageNumber="567">et al.</emphasis>
(2022)
</bibRefCitation>
has subcutaneous black streaks extending from the head to near the posterior end of the body.
</paragraph>
<paragraph id="8B98AA1BFFA7E97AD3D915F9B5B6F985" blockId="14.[151,1437,829,2008]" pageId="14" pageNumber="567">
<emphasis id="B9537609FFA7E97AD3D915F9B7D0FA75" bold="true" box="[199,402,1441,1467]" pageId="14" pageNumber="567">Buccal armature</emphasis>
(
<figureCitation id="131CB69EFFA7E97AD2BD15F9B7A8FA75" box="[419,490,1441,1467]" captionStart="FIGURE 6" captionStartId="19.[151,250,1184,1208]" captionTargetBox="[151,1436,181,1160]" captionTargetId="figure-22@19.[151,1436,181,1160]" captionTargetPageId="19" captionText="FIGURE 6. Doto kwakwak sp. nov., holotype, CASIZ 234657, scanning electron micrographs of radula. A. Segments of the radular ribbon. BC. Dorsolateral view of radular teeth. D. Lateral view of radular teeth." figureDoi="http://doi.org/10.5281/zenodo.15215508" httpUri="https://zenodo.org/record/15215508/files/figure.png" pageId="14" pageNumber="567">Fig. 6</figureCitation>
). The buccal mass is small and muscular. The jaws are thin and membranous. The radula is elongate and narrow, consisting of a ribbon of at least 69 teeth. The teeth are simply arched posteriorly with a series of anterior denticles. Each tooth has an elongate central cusp that extends more anteriorly than the lateral denticles. There are one to two pairs of elongate denticles on either side of the central cusp. On the outer side of the tooth there are an additional 12 smaller denticles on either side of the tooth.
</paragraph>
<paragraph id="8B98AA1BFFA7E97AD3D9160DB4FBF8A2" blockId="14.[151,1437,829,2008]" pageId="14" pageNumber="567">
<emphasis id="B9537609FFA7E97AD3D9160DB783F9A1" bold="true" box="[199,449,1621,1647]" pageId="14" pageNumber="567">Reproductive system</emphasis>
(
<figureCitation id="131CB69EFFA7E97AD2CE160DB46AF9BE" box="[464,552,1621,1648]" captionStart="FIGURE 5" captionStartId="18.[151,250,1908,1932]" captionTargetBox="[247,1337,182,1881]" captionTargetId="figure-38@18.[247,1340,181,1884]" captionTargetPageId="18" captionText="FIGURE 5. Reproductive system. A. Doto urak sp. nov., paratypes, composite of 184511 and 217599, scale = 2.0 mm. B. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 1.5 mm. Abbreviations: am = ampulla, ej = ejaculatory portion of vas deferens, fgm = female gland mass, p = penial sac, pr = prostatic portion of vas deferens, rs = receptaculum seminis, v = vagina." figureDoi="http://doi.org/10.5281/zenodo.15215506" httpUri="https://zenodo.org/record/15215506/files/figure.png" pageId="14" pageNumber="567">Fig. 5B</figureCitation>
). The reproductive system is androdiaulic. The pre-ampullary duct is short and expands into the wide, ovoid ampulla. The ampulla again narrows distally and divides into the oviduct and vas deferens. The oviduct widens into a serial receptaculum seminis that narrows into an elongate curved vagina that terminates at the female gonopore adjacent to the penis. The vas deferens widens into a curved, thickened prostatic portion and narrows into a thin, curved ejaculatory portion. The distal end of vas deferens enters the thick, curved penial sac, which terminates adjacent to the opening of the vagina. The female glands are not well differentiated from each other, but the mucous gland comprises the bulk of the organs and terminates adjacent to the opening of the vagina and penis and has a separate aperture.
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFA7E97AD3D9172DB5F3F819" pageId="14" pageNumber="567" type="etymology">
<paragraph id="8B98AA1BFFA7E97AD3D9172DB5F3F819" blockId="14.[151,1437,829,2008]" pageId="14" pageNumber="567">
<emphasis id="B9537609FFA7E97AD3D9172DB70AF841" bold="true" box="[199,328,1909,1935]" pageId="14" pageNumber="567">Etymology</emphasis>
. The species epithet comes from the Kumeyaay indigenous name kwakwak, meaning yellow, signifying the yellow-orange spots on the notum that distinguish this species. The Kumeyaay are the original inhabitants of the region around the
<typeStatus id="549C14B9FFA7E97AD13317E6B41FF816" box="[557,605,1982,2008]" pageId="14" pageNumber="567">type</typeStatus>
locality in San Diego County.
</paragraph>
</subSubSection>
<caption id="DF58FA93FFA6E97BD38917CFB514F81D" ID-DOI="http://doi.org/10.5281/zenodo.15215502" ID-Zenodo-Dep="15215502" httpUri="https://zenodo.org/record/15215502/files/figure.png" pageId="15" pageNumber="568" startId="15.[151,250,1943,1967]" targetBox="[222,1369,183,1917]" targetPageId="15" targetType="figure">
<paragraph id="8B98AA1BFFA6E97BD38917CFB514F81D" blockId="15.[151,1437,1943,2004]" pageId="15" pageNumber="568">
<emphasis id="B9537609FFA6E97BD38917CFB74FF861" bold="true" box="[151,269,1943,1967]" pageId="15" pageNumber="568">FIGURE 3</emphasis>
. Cerata.
<emphasis id="B9537609FFA6E97BD27917CFB73BF861" bold="true" box="[359,377,1943,1967]" pageId="15" pageNumber="568">A</emphasis>
.
<taxonomicName id="4C27D198FFA6E97BD29B17C0B7AEF87E" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[389,492,1944,1968]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="568" phylum="Mollusca" rank="species" species="urak" status="sp. nov.">
<emphasis id="B9537609FFA6E97BD29B17C0B7AEF87E" box="[389,492,1944,1968]" italics="true" pageId="15" pageNumber="568">Doto urak</emphasis>
</taxonomicName>
<emphasis id="B9537609FFA6E97BD2EC17C0B400F87E" bold="true" box="[498,578,1944,1968]" pageId="15" pageNumber="568">
<taxonomicNameLabel id="A260CB72FFA6E97BD2EC17C0B400F87E" box="[498,578,1944,1968]" pageId="15" pageNumber="568" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, paratype, CASIZ 184511.
<emphasis id="B9537609FFA6E97BD04617C0B52BF87E" bold="true" box="[856,873,1944,1968]" pageId="15" pageNumber="568">B</emphasis>
.
<taxonomicName id="4C27D198FFA6E97BD06817C0B2B3F87E" authority="O'Donoghue, 1921" authorityName="O'Donoghue" authorityYear="1921" box="[886,1265,1944,1968]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="568" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFA6E97BD06817C0B264F87E" box="[886,1062,1944,1968]" italics="true" pageId="15" pageNumber="568">Doto columbiana</emphasis>
<bibRefCitation id="EFB6D7EAFFA6E97BD73217C0B2B3F87E" author="O'Donoghue, C. H." box="[1068,1265,1944,1968]" pageId="15" pageNumber="568" pagination="147 - 209" refId="ref15121" refString="O'Donoghue, C. H. (1921) Nudibranchiate Mollusca from the Vancouver Island region. Transactions of the Royal Canadian Institute, 13 (1), 147-209, pls. VII - XII (I - V)." type="journal article" year="1921">ODonoghue, 1921</bibRefCitation>
</taxonomicName>
, CASIZ 189302.
<emphasis id="B9537609FFA6E97BD38917E3B6EBF81D" bold="true" box="[151,169,1979,2003]" pageId="15" pageNumber="568">C</emphasis>
.
<taxonomicName id="4C27D198FFA6E97BD3A817E4B70EF81A" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[182,332,1980,2004]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="15" pageNumber="568" phylum="Mollusca" rank="species" species="kwakwak" status="sp. nov.">
<emphasis id="B9537609FFA6E97BD3A817E4B70EF81A" box="[182,332,1980,2004]" italics="true" pageId="15" pageNumber="568">Doto kwakwak</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A260CB72FFA6E97BD24C17E4B7DFF81D" box="[338,413,1980,2003]" pageId="15" pageNumber="568" rank="species">sp. nov</taxonomicNameLabel>
, holotype, CASIZ 234657, scale = 2.5 mm.
</paragraph>
</caption>
<caption id="DF58FA93FFB9E964D38914C9B217FB03" ID-DOI="http://doi.org/10.5281/zenodo.15215504" ID-Zenodo-Dep="15215504" httpUri="https://zenodo.org/record/15215504/files/figure.png" pageId="16" pageNumber="569" startId="16.[151,250,1169,1193]" targetBox="[151,1436,181,1145]" targetPageId="16" targetType="figure">
<paragraph id="8B98AA1BFFB9E964D38914C9B217FB03" blockId="16.[151,1436,1169,1229]" pageId="16" pageNumber="569">
<emphasis id="B9537609FFB9E964D38914C9B752FB67" bold="true" box="[151,272,1169,1193]" pageId="16" pageNumber="569">FIGURE 4</emphasis>
.
<taxonomicName id="4C27D198FFB9E964D20114CAB7CBFB67" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[287,393,1169,1193]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="urak" status="sp. nov.">
<emphasis id="B9537609FFB9E964D20114CAB7CBFB67" box="[287,393,1169,1193]" italics="true" pageId="16" pageNumber="569">Doto urak</emphasis>
</taxonomicName>
<emphasis id="B9537609FFB9E964D28D14C9B7A4FB67" bold="true" box="[403,486,1169,1193]" pageId="16" pageNumber="569">
<taxonomicNameLabel id="A260CB72FFB9E964D28D14C9B7A4FB67" box="[403,486,1169,1193]" pageId="16" pageNumber="569" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, scanning electron micrographs of radula.
<emphasis id="B9537609FFB9E964D0B714C9B598FB67" bold="true" box="[937,986,1169,1193]" pageId="16" pageNumber="569">AC</emphasis>
. Paratype, CASIZ 217599.
<emphasis id="B9537609FFB9E964D61F14C9B372FB67" bold="true" box="[1281,1328,1169,1193]" pageId="16" pageNumber="569">AB</emphasis>
. Detail of radular teeth.
<emphasis id="B9537609FFB9E964D23D14EDB777FB03" bold="true" box="[291,309,1205,1229]" pageId="16" pageNumber="569">C</emphasis>
. Large section of radular ribbon.
<emphasis id="B9537609FFB9E964D19714EDB4D9FB03" bold="true" box="[649,667,1205,1229]" pageId="16" pageNumber="569">D</emphasis>
. Paratype, CASIZ 184511. Radular ribbon.
</paragraph>
</caption>
<subSubSection id="C33DF990FFB9E964D3D91549B2DAFA80" pageId="16" pageNumber="569" type="distribution">
<paragraph id="8B98AA1BFFB9E964D3D91549B2DAFA80" blockId="16.[151,1437,1297,2027]" pageId="16" pageNumber="569">
<emphasis id="B9537609FFB9E964D3D91549B71BFAE5" bold="true" box="[199,345,1297,1323]" pageId="16" pageNumber="569">Distribution</emphasis>
. Known intertidally from La Jolla, and subtidally from off the Palos Verdes Peninsula,
<collectingCountry id="F330EA8BFFB9E964D6331549B317FAE5" box="[1325,1365,1297,1323]" name="Iceland" pageId="16" pageNumber="569">Isla</collectingCountry>
Vista, and Anacapa and Santa Cruz Islands,
<collectingRegion id="49E364F9FFB9E964D123156CB4F3FA80" box="[573,689,1332,1358]" country="United States of America" name="California" pageId="16" pageNumber="569">California</collectingRegion>
,
<collectingCountry id="F330EA8BFFB9E964D1A3156CB4B1FA80" box="[701,755,1332,1358]" name="United States of America" pageId="16" pageNumber="569">USA</collectingCountry>
(
<bibRefCitation id="EFB6D7EAFFB9E964D01C156CB5A5FA80" author="Behrens, D. W. &amp; Fletcher, K. &amp; Jensen, G." box="[770,999,1332,1359]" pageId="16" pageNumber="569" refId="ref12622" refString="Behrens, D. W., Fletcher, K. &amp; Jensen, G. (2022) Nudibranchs &amp; Sea Slugs of the Eastern Pacific. MolaMarine, Bremerton, Washington, 170 pp." type="book" year="2022">
Behrens
<emphasis id="B9537609FFB9E964D078156DB5DDFA80" box="[870,927,1332,1358]" italics="true" pageId="16" pageNumber="569">et al.</emphasis>
, 2022
</bibRefCitation>
, present study).
</paragraph>
</subSubSection>
<subSubSection id="C33DF990FFB9E965D3D9150FB745FD2B" lastPageId="17" lastPageNumber="570" pageId="16" pageNumber="569" type="discussion">
<paragraph id="8B98AA1BFFB9E964D3D9150FB31FFA30" blockId="16.[151,1437,1297,2027]" pageId="16" pageNumber="569">
<emphasis id="B9537609FFB9E964D3D9150FB776FABF" bold="true" box="[199,308,1367,1393]" pageId="16" pageNumber="569">Remarks</emphasis>
. This species is known only from a few photographs and the only documented specimen collected is the
<typeStatus id="549C14B9FFB9E964D3DC1522B765FA5A" box="[194,295,1402,1428]" pageId="16" pageNumber="569" type="holotype">holotype</typeStatus>
, from La Jolla, CA (
<figureCitation id="131CB69EFFB9E964D10D1522B42BFA5B" box="[531,617,1402,1429]" captionStart="FIGURE 2" captionStartId="13.[151,250,1784,1808]" captionTargetBox="[151,1436,273,1761]" captionTargetId="figure-65@13.[151,1436,273,1761]" captionTargetPageId="13" captionText="FIGURE 2. Living animals. A. Doto urak sp. nov., holotype, CASIZ 182040, San Pedro, California, photo by T. M. Gosliner. B. Doto urak sp. nov., paratype, CASIZ 217592, Santa Cruz, California, photo by T. M. Gosliner. C. Doto urak sp. nov., showing egg mass, Hazard Canyon, San Luis Obispo County, California, photo by Jeff Goddard. D. Doto urak sp. nov., paratypes, CASIZ 056222, Drakes Estero, Marin County, California, photo by Jeff Goddard. E. Doto kwakwak sp. nov., holotype, CASIZ 234657, La Jolla, California, photo by Chloe and Trevor Van Loon. F. Doto kwakwak sp. nov., living animal, Anacapa Island, Ventura County, California, photo by Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.15215500" httpUri="https://zenodo.org/record/15215500/files/figure.png" pageId="16" pageNumber="569">Fig. 2E</figureCitation>
) (
<bibRefCitation id="EFB6D7EAFFB9E964D19F1523B57DFA5B" author="Van Loon, C. &amp; Van Loon, T." box="[641,831,1403,1429]" pageId="16" pageNumber="569" refId="ref16181" refString="Van Loon, C. &amp; Van Loon, T. (2022) Genus Doto. iNaturalist. Available from: https://www.inaturalist.org/observations/107604567 (accessed 27 June 2024)" type="url" year="2022">Van Loon, 2022</bibRefCitation>
). Six other specimens have been photographed: three from Anacapa Island in the Channel Islands by
<bibRefCitation id="EFB6D7EAFFB9E964D1A815C6B527FA76" author="Johnson, S." box="[694,869,1438,1464]" pageId="16" pageNumber="569" refId="ref13663" refString="Johnson, S. (2018) Genus Doto. iNaturalist. Available from: https://www.inaturalist.org/observations/108620551 (accessed 27 June 2024)" type="url" year="2018">Johnson (2018)</bibRefCitation>
(
<figureCitation id="131CB69EFFB9E964D06815C6B589FA76" box="[886,971,1438,1464]" captionStart="FIGURE 2" captionStartId="13.[151,250,1784,1808]" captionTargetBox="[151,1436,273,1761]" captionTargetId="figure-65@13.[151,1436,273,1761]" captionTargetPageId="13" captionText="FIGURE 2. Living animals. A. Doto urak sp. nov., holotype, CASIZ 182040, San Pedro, California, photo by T. M. Gosliner. B. Doto urak sp. nov., paratype, CASIZ 217592, Santa Cruz, California, photo by T. M. Gosliner. C. Doto urak sp. nov., showing egg mass, Hazard Canyon, San Luis Obispo County, California, photo by Jeff Goddard. D. Doto urak sp. nov., paratypes, CASIZ 056222, Drakes Estero, Marin County, California, photo by Jeff Goddard. E. Doto kwakwak sp. nov., holotype, CASIZ 234657, La Jolla, California, photo by Chloe and Trevor Van Loon. F. Doto kwakwak sp. nov., living animal, Anacapa Island, Ventura County, California, photo by Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.15215500" httpUri="https://zenodo.org/record/15215500/files/figure.png" pageId="16" pageNumber="569">Fig. 2F</figureCitation>
) and
<bibRefCitation id="EFB6D7EAFFB9E964D71215C6B2D2FA76" author="Klug, D." box="[1036,1168,1438,1464]" pageId="16" pageNumber="569" pagination="14 - 17" refId="ref14069" refString="Klug, D. (2017) nudibranch 14 Jan 14-17 [Photo]. Available from: https://www.flickr.com/photos/diverdoug/32277377371 (accessed 27 June 2024)" type="book chapter" year="2017">Klug (2017</bibRefCitation>
,
<bibRefCitation id="EFB6D7EAFFB9E964D78315C6B298FA76" author="Klug, D." box="[1181,1242,1438,1464]" pageId="16" pageNumber="569" refId="ref14142" refString="Klug, D. (2020) Genus Doto. iNaturalist. Available from: https://www.inaturalist.org/observations/59394208 (accessed 27 June 2024)" type="url" year="2020">2020</bibRefCitation>
); one from Santa Cruz Island in the Channel Islands (
<bibRefCitation id="EFB6D7EAFFB9E964D1281599B4FEFA15" author="Klug, D." box="[566,700,1473,1499]" pageId="16" pageNumber="569" refId="ref14109" refString="Klug, D. (2019) Genus Doto. iNaturalist. Available from: https://www.inaturalist.org/observations/35397064 (accessed 27 June 2024)" type="url" year="2019">Klug, 2019</bibRefCitation>
); one by Margaret Webb from off Haggertys, Palos Verdes, CA (
<bibRefCitation id="EFB6D7EAFFB9E964D38115BCB7C4FA30" author="Behrens, D. W. &amp; Fletcher, K. &amp; Jensen, G." box="[159,390,1508,1535]" pageId="16" pageNumber="569" refId="ref12622" refString="Behrens, D. W., Fletcher, K. &amp; Jensen, G. (2022) Nudibranchs &amp; Sea Slugs of the Eastern Pacific. MolaMarine, Bremerton, Washington, 170 pp." type="book" year="2022">
Behrens
<emphasis id="B9537609FFB9E964D21D15BDB77EFA30" box="[259,316,1508,1534]" italics="true" pageId="16" pageNumber="569">et al.</emphasis>
, 2022
</bibRefCitation>
); and one from the shallow subtidal off Coal Oil Point, Isla Vista, CA by
<bibRefCitation id="EFB6D7EAFFB9E964D7A715BDB31BFA30" author="Mazza, M." box="[1209,1369,1508,1535]" pageId="16" pageNumber="569" refId="ref14667" refString="Mazza, M. (2019). Genus Doto. iNaturalist. Available from: https://www.inaturalist.org/observations/20974121 (accessed 27 June 2024)" type="url" year="2019">Mazza (2019)</bibRefCitation>
.
</paragraph>
<paragraph id="8B98AA1BFFB9E965D3D91650B248FEF0" blockId="16.[151,1437,1297,2027]" lastBlockId="17.[151,1437,151,741]" lastPageId="17" lastPageNumber="570" pageId="16" pageNumber="569">
In our phylogeny,
<taxonomicName id="4C27D198FFB9E964D2881650B47FF9EF" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[406,573,1543,1569]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB9E964D2881650B47FF9EF" box="[406,573,1543,1569]" italics="true" pageId="16" pageNumber="569">Doto kwakwak</emphasis>
</taxonomicName>
and
<taxonomicName id="4C27D198FFB9E964D16A1650B499F9EF" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[628,731,1543,1569]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFB9E964D16A1650B499F9EF" box="[628,731,1543,1569]" italics="true" pageId="16" pageNumber="569">Doto kya</emphasis>
</taxonomicName>
are sister to most of the other members of the Atlantic/eastern Pacific clade (
<figureCitation id="131CB69EFFB9E964D22B1672B734F98B" box="[309,374,1578,1605]" captionStart="FIGURE 1" captionStartId="11.[151,250,1044,1068]" captionTargetBox="[153,1432,184,1019]" captionTargetId="figure-22@11.[151,1436,181,1020]" captionTargetPageId="11" captionText="FIGURE 1. Bayesian phylogenetic tree of Doto estimated from the three genes (COI + 16S + H3) concatenated data. Numbers above branches refer to BI posterior probabilities (PP), while numbers below branches refer to ML non-parametric bootstrapping values (BS). Relationships not recovered during ML analysis are indicated by dashes. Bars on the right of the tree indicate the different species delimitation methods. ABGD, ASAP, and GMYC are based in the COI dataset and the bPTP in the three-gene (COI + 16S + H3) concatenated dataset." figureDoi="http://doi.org/10.5281/zenodo.15215498" httpUri="https://zenodo.org/record/15215498/files/figure.png" pageId="16" pageNumber="569">Fig. 1</figureCitation>
). Of the species found along the Pacific coast of the
<collectingCountry id="F330EA8BFFB9E964D0A11672B217F98B" box="[959,1109,1578,1605]" name="United States of America" pageId="16" pageNumber="569">United States</collectingCountry>
and
<collectingCountry id="F330EA8BFFB9E964D7971672B2A2F98A" box="[1161,1248,1578,1604]" name="Canada" pageId="16" pageNumber="569">Canada</collectingCountry>
, it can be readily distinguished externally by the presence of yellowish-orange spots on the body. Similarly,
<taxonomicName id="4C27D198FFB9E964D76D1616B2B8F9A6" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1139,1274,1614,1640]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB9E964D76D1616B2B8F9A6" box="[1139,1274,1614,1640]" italics="true" pageId="16" pageNumber="569">D. kwakwak</emphasis>
</taxonomicName>
, like
<taxonomicName id="4C27D198FFB9E964D62B1616B3DFF9A9" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[1333,1437,1614,1639]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFB9E964D62B1616B3DFF9A9" box="[1333,1437,1614,1639]" italics="true" pageId="16" pageNumber="569">D. amyra</emphasis>
</taxonomicName>
and
<taxonomicName id="4C27D198FFB9E964D3D6162AB75CF945" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[200,286,1649,1675]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFB9E964D3D6162AB75CF945" box="[200,286,1649,1675]" italics="true" pageId="16" pageNumber="569">D. urak</emphasis>
</taxonomicName>
, generally lacks the scattered brown pigment on the body that is found in
<taxonomicName id="4C27D198FFB9E964D775162AB2F7F945" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[1131,1205,1649,1675]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFB9E964D775162AB2F7F945" box="[1131,1205,1649,1675]" italics="true" pageId="16" pageNumber="569">D. kya</emphasis>
</taxonomicName>
and
<taxonomicName id="4C27D198FFB9E964D7F0162AB3D7F945" authorityName="O'Donoghue" authorityYear="1921" box="[1262,1429,1649,1675]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFB9E964D7F0162AB3D7F945" box="[1262,1429,1649,1675]" italics="true" pageId="16" pageNumber="569">D. columbiana</emphasis>
</taxonomicName>
, but some specimens may have subcutaneous dark pigment (
<bibRefCitation id="EFB6D7EAFFB9E964D02616CCB5FEF960" author="Klug, D." box="[824,956,1684,1710]" pageId="16" pageNumber="569" refId="ref14109" refString="Klug, D. (2019) Genus Doto. iNaturalist. Available from: https://www.inaturalist.org/observations/35397064 (accessed 27 June 2024)" type="url" year="2019">Klug, 2019</bibRefCitation>
). In
<taxonomicName id="4C27D198FFB9E964D0F016CDB21BF960" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[1006,1113,1685,1710]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFB9E964D0F016CDB21BF960" box="[1006,1113,1685,1710]" italics="true" pageId="16" pageNumber="569">D. amyra</emphasis>
</taxonomicName>
, the lateral extensions of the foot are elongate, whereas, they are much shorter in
<taxonomicName id="4C27D198FFB9E964D1FB16E0B5CEF91F" authorityName="O'Donoghue" authorityYear="1921" box="[741,908,1719,1745]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFB9E964D1FB16E0B5CEF91F" box="[741,908,1719,1745]" italics="true" pageId="16" pageNumber="569">D. columbiana</emphasis>
</taxonomicName>
,
<taxonomicName id="4C27D198FFB9E964D08416E0B5A6F91F" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[922,996,1719,1745]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFB9E964D08416E0B5A6F91F" box="[922,996,1719,1745]" italics="true" pageId="16" pageNumber="569">D. kya</emphasis>
</taxonomicName>
,
<taxonomicName id="4C27D198FFB9E964D0EC16E0B334F91F" authority="Marcus &amp; Marcus, 1967" authorityName="Marcus &amp; Marcus" authorityYear="1967" box="[1010,1398,1719,1746]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="lancei">
<emphasis id="B9537609FFB9E964D0EC16E0B21AF91F" box="[1010,1112,1719,1745]" italics="true" pageId="16" pageNumber="569">D. lancei</emphasis>
<bibRefCitation id="EFB6D7EAFFB9E964D77E16E0B334F91F" author="Marcus, Ev. &amp; Marcus, Er." box="[1120,1398,1719,1746]" pageId="16" pageNumber="569" pagination="1 - 256" refId="ref14535" refString="Marcus, Ev. &amp; Marcus, Er. (1967) American opisthobranch mollusks Part I, Tropical American opisthobranchs. Part II, Opisthobranchs from the Gulf of California. Studies in Tropical Oceanography, 6 (1 - 2), 1-256." type="journal article" year="1967">Marcus &amp; Marcus, 1967</bibRefCitation>
</taxonomicName>
,
<emphasis id="B9537609FFB9E964D69F16E0B693F93A" italics="true" pageId="16" pageNumber="569">
<taxonomicName id="4C27D198FFB9E964D69F16E0B68FF93A" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="urak">D. urak</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C27D198FFB9E964D2191683B7CDF93A" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[263,399,1754,1780]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB9E964D2191683B7CDF93A" box="[263,399,1754,1780]" italics="true" pageId="16" pageNumber="569">D. kwakwak</emphasis>
</taxonomicName>
. In
<taxonomicName id="4C27D198FFB9E964D2A41683B44CF93A" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[442,526,1754,1780]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFB9E964D2A41683B44CF93A" box="[442,526,1754,1780]" italics="true" pageId="16" pageNumber="569">D. urak</emphasis>
</taxonomicName>
and
<taxonomicName id="4C27D198FFB9E964D15A1683B48FF93A" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[580,717,1754,1780]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB9E964D15A1683B48FF93A" box="[580,717,1754,1780]" italics="true" pageId="16" pageNumber="569">D. kwakwak</emphasis>
</taxonomicName>
, the rhinophore sheaths are flared apically while they are largely cylindrical in
<taxonomicName id="4C27D198FFB9E964D22A16A6B7E2F8D9" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[308,416,1790,1815]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFB9E964D22A16A6B7E2F8D9" box="[308,416,1790,1815]" italics="true" pageId="16" pageNumber="569">D. amyra</emphasis>
</taxonomicName>
,
<taxonomicName id="4C27D198FFB9E964D2B016A6B421F8D6" authorityName="O'Donoghue" authorityYear="1921" box="[430,611,1790,1816]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="columbaiana">
<emphasis id="B9537609FFB9E964D2B016A6B421F8D6" box="[430,611,1790,1816]" italics="true" pageId="16" pageNumber="569">D. columbaiana</emphasis>
</taxonomicName>
,
<emphasis id="B9537609FFB9E964D16C16A6B481F8D6" box="[626,707,1790,1816]" italics="true" pageId="16" pageNumber="569">
<taxonomicName id="4C27D198FFB9E964D16C16A6B4FDF8D6" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[626,703,1790,1816]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kya">D. kya</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C27D198FFB9E964D1E516A6B520F8D6" authorityName="Marcus &amp; Marcus" authorityYear="1967" box="[763,866,1790,1816]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="lancei">
<emphasis id="B9537609FFB9E964D1E516A6B520F8D6" box="[763,866,1790,1816]" italics="true" pageId="16" pageNumber="569">D. lancei</emphasis>
</taxonomicName>
. The ceratal tubercles of
<taxonomicName id="4C27D198FFB9E964D79D16A6B34FF8D6" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1155,1293,1790,1816]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB9E964D79D16A6B34FF8D6" box="[1155,1293,1790,1816]" italics="true" pageId="16" pageNumber="569">D. kwakwak</emphasis>
</taxonomicName>
,
<taxonomicName id="4C27D198FFB9E964D60216A6B330F8D6" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1308,1394,1790,1816]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFB9E964D60216A6B330F8D6" box="[1308,1394,1790,1816]" italics="true" pageId="16" pageNumber="569">D. urak</emphasis>
</taxonomicName>
,
<emphasis id="B9537609FFB9E964D69F16A6B6A0F8F5" italics="true" pageId="16" pageNumber="569">
<taxonomicName id="4C27D198FFB9E964D69F16A6B69CF8F5" authorityName="Marcus &amp; Marcus" authorityYear="1967" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="lancei">D. lancei</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C27D198FFB9E964D205177AB724F8F5" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[283,358,1825,1851]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFB9E964D205177AB724F8F5" box="[283,358,1825,1851]" italics="true" pageId="16" pageNumber="569">D. kya</emphasis>
</taxonomicName>
are spherical to elongate apically, while those of
<taxonomicName id="4C27D198FFB9E964D087177AB202F8F5" authorityName="O'Donoghue" authorityYear="1921" box="[921,1088,1825,1851]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFB9E964D087177AB202F8F5" box="[921,1088,1825,1851]" italics="true" pageId="16" pageNumber="569">D. columbiana</emphasis>
</taxonomicName>
and
<taxonomicName id="4C27D198FFB9E964D767177AB2A6F8F5" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[1145,1252,1826,1851]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFB9E964D767177AB2A6F8F5" box="[1145,1252,1826,1851]" italics="true" pageId="16" pageNumber="569">D. amyra</emphasis>
</taxonomicName>
are much lower to flattened or irregular in shape (
<figureCitation id="131CB69EFFB9E964D106171CB433F890" box="[536,625,1860,1886]" captionStart="FIGURE 3" captionStartId="15.[151,250,1943,1967]" captionTargetBox="[222,1369,183,1917]" captionTargetId="figure-22@15.[212,1376,181,1920]" captionTargetPageId="15" captionText="FIGURE 3. Cerata. A. Doto urak sp. nov., paratype, CASIZ 184511. B. Doto columbiana ODonoghue, 1921, CASIZ 189302. C. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 2.5 mm." figureDoi="http://doi.org/10.5281/zenodo.15215502" httpUri="https://zenodo.org/record/15215502/files/figure.png" pageId="16" pageNumber="569">Fig. 3B</figureCitation>
). Also, the opaque white glandular apices of the ceratal tubercles of
<taxonomicName id="4C27D198FFB9E964D69F171DB6BCF84F" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB9E964D69F171DB6BCF84F" italics="true" pageId="16" pageNumber="569">D. kwakwak</emphasis>
</taxonomicName>
,
<emphasis id="B9537609FFB9E964D2101730B72EF84F" box="[270,364,1895,1921]" italics="true" pageId="16" pageNumber="569">
<taxonomicName id="4C27D198FFB9E964D2101730B72AF84F" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[270,360,1895,1921]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="urak">D. urak</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C27D198FFB9E964D2B91730B7B1F84F" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[423,499,1895,1921]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFB9E964D2B91730B7B1F84F" box="[423,499,1895,1921]" italics="true" pageId="16" pageNumber="569">D. kya</emphasis>
</taxonomicName>
differ from those of
<taxonomicName id="4C27D198FFB9E964D1F91730B5CDF84F" authorityName="O'Donoghue" authorityYear="1921" box="[743,911,1895,1921]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFB9E964D1F91730B5CDF84F" box="[743,911,1895,1921]" italics="true" pageId="16" pageNumber="569">D. columbiana</emphasis>
</taxonomicName>
,
<emphasis id="B9537609FFB9E964D0811730B24DF84F" box="[927,1039,1895,1921]" italics="true" pageId="16" pageNumber="569">
<taxonomicName id="4C27D198FFB9E964D0811730B249F84F" authorityName="Marcus &amp; Marcus" authorityYear="1967" box="[927,1035,1895,1921]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="lancei">D. lancei</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C27D198FFB9E964D7541730B2F4F84F" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[1098,1206,1896,1921]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFB9E964D7541730B2F4F84F" box="[1098,1206,1896,1921]" italics="true" pageId="16" pageNumber="569">D. amyra</emphasis>
</taxonomicName>
, which lack opaque white on the tubercle apices.
<taxonomicName id="4C27D198FFB9E964D2FF17D3B4C5F86A" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[481,647,1930,1956]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB9E964D2FF17D3B4C5F86A" box="[481,647,1930,1956]" italics="true" pageId="16" pageNumber="569">Doto kwakwak</emphasis>
</taxonomicName>
has many tubercles of different sizes shapes (some bulbous and some longer), in contrast to
<taxonomicName id="4C27D198FFB9E964D28F17F6B7BEF809" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[401,508,1966,1991]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFB9E964D28F17F6B7BEF809" box="[401,508,1966,1991]" italics="true" pageId="16" pageNumber="569">D. amyra</emphasis>
</taxonomicName>
,
<taxonomicName id="4C27D198FFB9E964D11717F6B41DF806" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[521,607,1966,1992]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFB9E964D11717F6B41DF806" box="[521,607,1966,1992]" italics="true" pageId="16" pageNumber="569">D. urak</emphasis>
</taxonomicName>
,
<taxonomicName id="4C27D198FFB9E964D17017F6B496F806" authorityName="Marcus &amp; Marcus" authorityYear="1967" box="[622,724,1966,1992]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="lancei">
<emphasis id="B9537609FFB9E964D17017F6B496F806" box="[622,724,1966,1992]" italics="true" pageId="16" pageNumber="569">D. lancei</emphasis>
</taxonomicName>
,
<emphasis id="B9537609FFB9E964D1FD17F6B5D3F806" box="[739,913,1966,1992]" italics="true" pageId="16" pageNumber="569">
<taxonomicName id="4C27D198FFB9E964D1FD17F6B5CFF806" authorityName="O'Donoghue" authorityYear="1921" box="[739,909,1966,1992]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="columbiana">D. columbiana</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C27D198FFB9E964D0D617F6B250F806" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[968,1042,1966,1992]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFB9E964D0D617F6B250F806" box="[968,1042,1966,1992]" italics="true" pageId="16" pageNumber="569">D. kya</emphasis>
</taxonomicName>
, which have uniformly tuberculate cerata. The cerata or
<taxonomicName id="4C27D198FFB9E964D296178AB451F825" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[392,531,2001,2027]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB9E964D296178AB451F825" box="[392,531,2001,2027]" italics="true" pageId="16" pageNumber="569">D. kwakwak</emphasis>
</taxonomicName>
are elongate like those of
<taxonomicName id="4C27D198FFB9E964D05D178AB5EDF825" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[835,943,2002,2027]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFB9E964D05D178AB5EDF825" box="[835,943,2002,2027]" italics="true" pageId="16" pageNumber="569">D. amyra</emphasis>
</taxonomicName>
,
<taxonomicName id="4C27D198FFB9E964D0A0178AB24BF825" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[958,1033,2001,2027]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFB9E964D0A0178AB24BF825" box="[958,1033,2001,2027]" italics="true" pageId="16" pageNumber="569">D. kya</emphasis>
</taxonomicName>
,
<emphasis id="B9537609FFB9E964D706178AB2C5F825" box="[1048,1159,2001,2027]" italics="true" pageId="16" pageNumber="569">
<taxonomicName id="4C27D198FFB9E964D706178AB2C1F825" authorityName="Marcus &amp; Marcus" authorityYear="1967" box="[1048,1155,2001,2027]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="lancei">D. lancei</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C27D198FFB9E964D7DF178AB32BF825" authorityName="O'Donoghue" authorityYear="1921" box="[1217,1385,2001,2027]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="16" pageNumber="569" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFB9E964D7DF178AB32BF825" box="[1217,1385,2001,2027]" italics="true" pageId="16" pageNumber="569">D. columbiana</emphasis>
</taxonomicName>
, but tend to get longer towards the posterior end of the body. The cerata of
<taxonomicName id="4C27D198FFB8E965D0A910CFB24FFF7F" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[951,1037,151,177]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFB8E965D0A910CFB24FFF7F" box="[951,1037,151,177]" italics="true" pageId="17" pageNumber="570">D. urak</emphasis>
</taxonomicName>
are shorter and more rounded. The digestive gland within the ceratal core of
<taxonomicName id="4C27D198FFB8E965D17410E3B4B6FF1A" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[618,756,186,212]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB8E965D17410E3B4B6FF1A" box="[618,756,186,212]" italics="true" pageId="17" pageNumber="570">D. kwakwak</emphasis>
</taxonomicName>
appears to be a much more vibrant rust/red color, than in
<taxonomicName id="4C27D198FFB8E965D69F10E3B6FCFF39" baseAuthorityName="Marcus" baseAuthorityYear="1961" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFB8E965D69F10E3B6FCFF39" italics="true" pageId="17" pageNumber="570">D. kya</emphasis>
</taxonomicName>
,
<emphasis id="B9537609FFB8E965D3D51086B77AFF39" box="[203,312,221,247]" italics="true" pageId="17" pageNumber="570">
<taxonomicName id="4C27D198FFB8E965D3D51086B776FF39" authorityName="Marcus &amp; Marcus" authorityYear="1967" box="[203,308,221,247]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="lancei">D. lancei</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C27D198FFB8E965D2701086B451FF39" authorityName="O'Donoghue" authorityYear="1921" box="[366,531,221,247]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFB8E965D2701086B451FF39" box="[366,531,221,247]" italics="true" pageId="17" pageNumber="570">D. columbiana</emphasis>
</taxonomicName>
, which have beige to light orange cores. In
<taxonomicName id="4C27D198FFB8E965D0E71086B2C0FF39" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1017,1154,221,247]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB8E965D0E71086B2C0FF39" box="[1017,1154,221,247]" italics="true" pageId="17" pageNumber="570">D. kwakwak</emphasis>
</taxonomicName>
, a pseudobranch is either absent from the inner side of the cerata or if present is only an unbranched vertical line. In all other eastern Pacific species of
<taxonomicName id="4C27D198FFB8E965D213117CB707FEF3" authorityName="Oken" authorityYear="1815" box="[269,325,292,317]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="genus">
<emphasis id="B9537609FFB8E965D213117CB707FEF3" box="[269,325,292,317]" italics="true" pageId="17" pageNumber="570">Doto</emphasis>
</taxonomicName>
a well-developed, branched pseudobranch is present (
<figureCitation id="131CB69EFFB8E965D0BA117BB5BEFEF0" box="[932,1020,291,318]" captionStart="FIGURE 3" captionStartId="15.[151,250,1943,1967]" captionTargetBox="[222,1369,183,1917]" captionTargetId="figure-22@15.[212,1376,181,1920]" captionTargetPageId="15" captionText="FIGURE 3. Cerata. A. Doto urak sp. nov., paratype, CASIZ 184511. B. Doto columbiana ODonoghue, 1921, CASIZ 189302. C. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 2.5 mm." figureDoi="http://doi.org/10.5281/zenodo.15215502" httpUri="https://zenodo.org/record/15215502/files/figure.png" pageId="17" pageNumber="570">Fig. 3B</figureCitation>
).
</paragraph>
<paragraph id="8B98AA1BFFB8E965D3D9111FB7FFFDDF" blockId="17.[151,1437,151,741]" pageId="17" pageNumber="570">
As stated in the remarks regarding
<taxonomicName id="4C27D198FFB8E965D145111FB4F1FEAF" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[603,691,327,353]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFB8E965D145111FB4F1FEAF" box="[603,691,327,353]" italics="true" pageId="17" pageNumber="570">D. urak</emphasis>
</taxonomicName>
, the species of
<taxonomicName id="4C27D198FFB8E965D076111FB5E2FEAE" authorityName="Oken" authorityYear="1815" box="[872,928,327,352]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="genus">
<emphasis id="B9537609FFB8E965D076111FB5E2FEAE" box="[872,928,327,352]" italics="true" pageId="17" pageNumber="570">Doto</emphasis>
</taxonomicName>
from the Pacific coast of North America all have similar radular tooth morphology, and there is nothing distinctive about the radular teeth of
<taxonomicName id="4C27D198FFB8E965D7C41133B321FE4A" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1242,1379,362,388]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB8E965D7C41133B321FE4A" box="[1242,1379,362,388]" italics="true" pageId="17" pageNumber="570">D. kwakwak</emphasis>
</taxonomicName>
. The reproductive system of
<taxonomicName id="4C27D198FFB8E965D2BE11D6B468FE69" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[416,554,397,423]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB8E965D2BE11D6B468FE69" box="[416,554,397,423]" italics="true" pageId="17" pageNumber="570">D. kwakwak</emphasis>
</taxonomicName>
is similar to that of
<taxonomicName id="4C27D198FFB8E965D01111D6B5F7FE69" authorityName="O'Donoghue" authorityYear="1921" box="[783,949,397,423]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFB8E965D01111D6B5F7FE69" box="[783,949,397,423]" italics="true" pageId="17" pageNumber="570">D. columbiana</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFB6D7EAFFB8E965D0DB11D5B233FE69" author="Marcus, Er." box="[965,1137,397,423]" pageId="17" pageNumber="570" pagination="1 - 85" refId="ref14444" refString="Marcus, Er. (1961 a) Opisthobranch mollusks from California. The Veliger, 3, 1-85, pls. 1 - 10." type="journal article" year="1961">Marcus, 1961a</bibRefCitation>
: fig. 129) with a relatively short, curved ejaculatory segment and a relatively short penial sac. However, the vaginal duct of
<taxonomicName id="4C27D198FFB8E965D7EA11E9B3DEFE04" authorityName="O'Donoghue" authorityYear="1921" box="[1268,1436,432,458]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFB8E965D7EA11E9B3DEFE04" box="[1268,1436,432,458]" italics="true" pageId="17" pageNumber="570">D. columbiana</emphasis>
</taxonomicName>
has a distinct sphincter as in
<taxonomicName id="4C27D198FFB8E965D2C0118CB40BFE23" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[478,585,468,493]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFB8E965D2C0118CB40BFE23" box="[478,585,468,493]" italics="true" pageId="17" pageNumber="570">D. amyra</emphasis>
</taxonomicName>
, in contrast to that of
<taxonomicName id="4C27D198FFB8E965D05D118CB582FE23" box="[835,960,467,493]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kwakwa">
<emphasis id="B9537609FFB8E965D05D118CB582FE23" box="[835,960,467,493]" italics="true" pageId="17" pageNumber="570">D. kwakwa</emphasis>
</taxonomicName>
k (
<figureCitation id="131CB69EFFB8E965D0C1118BB275FE20" box="[991,1079,467,494]" captionStart="FIGURE 5" captionStartId="18.[151,250,1908,1932]" captionTargetBox="[247,1337,182,1881]" captionTargetId="figure-38@18.[247,1340,181,1884]" captionTargetPageId="18" captionText="FIGURE 5. Reproductive system. A. Doto urak sp. nov., paratypes, composite of 184511 and 217599, scale = 2.0 mm. B. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 1.5 mm. Abbreviations: am = ampulla, ej = ejaculatory portion of vas deferens, fgm = female gland mass, p = penial sac, pr = prostatic portion of vas deferens, rs = receptaculum seminis, v = vagina." figureDoi="http://doi.org/10.5281/zenodo.15215506" httpUri="https://zenodo.org/record/15215506/files/figure.png" pageId="17" pageNumber="570">Fig. 5B</figureCitation>
),
<taxonomicName id="4C27D198FFB8E965D752118CB2E0FE23" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[1100,1186,467,493]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFB8E965D752118CB2E0FE23" box="[1100,1186,467,493]" italics="true" pageId="17" pageNumber="570">D. urak</emphasis>
</taxonomicName>
(
<figureCitation id="131CB69EFFB8E965D7AD118BB34FFE20" box="[1203,1293,467,494]" captionStart="FIGURE 5" captionStartId="18.[151,250,1908,1932]" captionTargetBox="[247,1337,182,1881]" captionTargetId="figure-38@18.[247,1340,181,1884]" captionTargetPageId="18" captionText="FIGURE 5. Reproductive system. A. Doto urak sp. nov., paratypes, composite of 184511 and 217599, scale = 2.0 mm. B. Doto kwakwak sp. nov, holotype, CASIZ 234657, scale = 1.5 mm. Abbreviations: am = ampulla, ej = ejaculatory portion of vas deferens, fgm = female gland mass, p = penial sac, pr = prostatic portion of vas deferens, rs = receptaculum seminis, v = vagina." figureDoi="http://doi.org/10.5281/zenodo.15215506" httpUri="https://zenodo.org/record/15215506/files/figure.png" pageId="17" pageNumber="570">Fig. 5A</figureCitation>
), and
<taxonomicName id="4C27D198FFB8E965D64C118CB3DEFE23" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[1362,1436,467,493]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kya">
<emphasis id="B9537609FFB8E965D64C118CB3DEFE23" box="[1362,1436,467,493]" italics="true" pageId="17" pageNumber="570">D. kya</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFB6D7EAFFB8E965D38111AFB709FDDF" author="Marcus, Er." box="[159,331,503,529]" pageId="17" pageNumber="570" pagination="1 - 85" refId="ref14444" refString="Marcus, Er. (1961 a) Opisthobranch mollusks from California. The Veliger, 3, 1-85, pls. 1 - 10." type="journal article" year="1961">Marcus, 1961a</bibRefCitation>
: fig. 140).
</paragraph>
<paragraph id="8B98AA1BFFB8E965D3D91243B5EBFD50" blockId="17.[151,1437,151,741]" pageId="17" pageNumber="570">
<taxonomicName id="4C27D198FFB8E965D3D91243B732FDFA" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[199,368,538,564]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB8E965D3D91243B732FDFA" box="[199,368,538,564]" italics="true" pageId="17" pageNumber="570">Doto kwakwak</emphasis>
</taxonomicName>
is molecularly distinct and is a single distinct species according to each species delimitation method undertaken.
<taxonomicName id="4C27D198FFB8E965D2631266B444FD99" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[381,518,573,599]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kwakwak">
<emphasis id="B9537609FFB8E965D2631266B444FD99" box="[381,518,573,599]" italics="true" pageId="17" pageNumber="570">D. kwakwak</emphasis>
</taxonomicName>
interspecific
<emphasis id="B9537609FFB8E965D1811266B4EFFD99" box="[671,685,574,599]" italics="true" pageId="17" pageNumber="570">p</emphasis>
-distances to the closest species are: 12.5112.99% to
<taxonomicName id="4C27D198FFB8E965D6101266B3D4FD99" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[1294,1430,574,599]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="amyra">
<emphasis id="B9537609FFB8E965D6101266B3D4FD99" box="[1294,1430,574,599]" italics="true" pageId="17" pageNumber="570">Doto amyra</emphasis>
</taxonomicName>
, 14.27% to
<taxonomicName id="4C27D198FFB8E965D2061239B783FDB4" authorityName="O'Donoghue" authorityYear="1921" box="[280,449,608,634]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="columbiana">
<emphasis id="B9537609FFB8E965D2061239B783FDB4" box="[280,449,608,634]" italics="true" pageId="17" pageNumber="570">D. columbiana</emphasis>
</taxonomicName>
, 14.0914.46% to
<emphasis id="B9537609FFB8E965D1BE1239B4B1FDB4" box="[672,755,608,634]" italics="true" pageId="17" pageNumber="570">
<taxonomicName id="4C27D198FFB8E965D1BE1239B4ADFDB4" baseAuthorityName="Marcus" baseAuthorityYear="1961" box="[672,751,608,634]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="kya">D. kya</taxonomicName>
,
</emphasis>
13.5414.64% to
<taxonomicName id="4C27D198FFB8E965D0D41239B276FDB4" authorityName="Marcus &amp; Marcus" authorityYear="1967" box="[970,1076,608,634]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="lancei">
<emphasis id="B9537609FFB8E965D0D41239B276FDB4" box="[970,1076,608,634]" italics="true" pageId="17" pageNumber="570">D. lancei</emphasis>
</taxonomicName>
, 12.4712.84% from
<emphasis id="B9537609FFB8E965D62D1239B30DFDB4" box="[1331,1359,609,634]" italics="true" pageId="17" pageNumber="570">D.</emphasis>
sp. B, 14.09% to
<emphasis id="B9537609FFB8E965D20F12DCB76FFD53" box="[273,301,644,669]" italics="true" pageId="17" pageNumber="570">D.</emphasis>
sp. C, 14.83% to
<emphasis id="B9537609FFB8E965D2EB12DCB453FD53" box="[501,529,644,669]" italics="true" pageId="17" pageNumber="570">D.</emphasis>
sp. I, and 13.9014.93% to
<taxonomicName id="4C27D198FFB8E965D05212DCB5E3FD53" authorityName="Adayapalam &amp; Bonomo &amp; Shipman &amp; Gosliner" authorityYear="2025" box="[844,929,643,669]" class="Gastropoda" family="Dotidae" genus="Doto" kingdom="Animalia" order="Nudibranchia" pageId="17" pageNumber="570" phylum="Mollusca" rank="species" species="urak">
<emphasis id="B9537609FFB8E965D05212DCB5E3FD53" box="[844,929,643,669]" italics="true" pageId="17" pageNumber="570">D. urak</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B98AA1BFFB8E965D3D912FFB745FD2B" blockId="17.[151,1437,151,741]" pageId="17" pageNumber="570">Little is known about the natural history of this species and its larval biology and development mode remains unknown.</paragraph>
</subSubSection>
</treatment>
</document>

View file

@ -1,137 +0,0 @@
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<mods:title id="20296118104234F8E7B71F9D75D017F4">New species and records of Sericinae from India and Nepal (Coleoptera: Scarabaeidae)</mods:title>
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<emphasis id="B943E75BA079FF96C0AE8C87F85DC3D0" bold="true" box="[151,586,151,178]" pageId="10" pageNumber="381">
<emphasis id="B943E75BA079FF96C0AE8C87FB75C3D0" bold="true" box="[151,354,151,178]" italics="true" pageId="10" pageNumber="381">Maladera affinis</emphasis>
(
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)
</emphasis>
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</heading>
</paragraph>
</subSubSection>
<subSubSection id="C32D68C2A079FF96C0AE8CCFFB14C294" pageId="10" pageNumber="381" type="materials_examined">
<paragraph id="8B883B49A079FF96C0AE8CCFFF4EC398" blockId="10.[151,1369,223,250]" box="[151,1369,223,250]" pageId="10" pageNumber="381">
<emphasis id="B943E75BA079FF96C0AE8CCFFB6EC398" bold="true" box="[151,377,223,249]" pageId="10" pageNumber="381">Material examined</emphasis>
.
<materialsCitation id="3B5F3114A079FF96C1BE8CCFFF42C398" box="[391,1365,223,250]" collectingDate="1956-06-15" collectionCode="ZMUH" country="India" county="Nr" location="Timli" municipality="Exp" pageId="10" pageNumber="381" specimenCount="1" specimenCount-male="1">
<specimenCount id="9D31F0C0A079FF96C1BE8CCFFBA6C39B" box="[391,433,223,250]" count="1" pageId="10" pageNumber="381" type="male">1 ♂</specimenCount>
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,
<location id="8EE86D92A079FF96C2298CCFF87FC398" LSID="urn:lsid:plazi:treatment:039E8A5FA079FF96C0AE8C87FB14C294:8EE86D92A079FF96C2298CCFF87FC398" box="[528,616,223,249]" country="India" county="Nr" municipality="Exp" name="Siwalik" pageId="10" pageNumber="381">Siwalik</location>
,
<date id="FF891D89A079FF96C24D8CCFF8E1C39B" box="[628,758,223,250]" pageId="10" pageNumber="381" value="1956-06-15">
<collectingDate id="EFCDE461A079FF96C24D8CCFF8E1C39B" box="[628,758,223,250]" pageId="10" pageNumber="381" value="1956-06-15">15.VI.1956</collectingDate>
</date>
, D.
<collectingCountry id="F3207BD9A079FF96C31A8CCFF97CC398" box="[803,875,223,249]" name="India" pageId="10" pageNumber="381">Indien</collectingCountry>
<collectingMunicipality id="6BECA133A079FF96C34B8CF0F9B5C39B" box="[882,930,224,250]" pageId="10" pageNumber="381">Exp</collectingMunicipality>
,
<collectingCounty id="62E943C5A079FF96C3948CF0F9DCC39B" box="[941,971,224,250]" pageId="10" pageNumber="381">Nr</collectingCounty>
1284, Eing. Nr, 1, 1956” (
<collectionCode id="ED26A38CA079FF96C4CD8CCFFF5AC398" box="[1268,1357,223,249]" country="Viet Nam" httpUri="http://grbio.org/cool/82c5-17ij" name="Zoological Museum, University of Hanoi" pageId="10" pageNumber="381" type="">ZMUH</collectionCode>
)
</materialsCitation>
.
</paragraph>
<paragraph id="8B883B49A079FF96C0AE8D7DF82CC2EB" blockId="10.[151,571,365,394]" box="[151,571,365,394]" pageId="10" pageNumber="381">
<taxonomicName id="4C3740CAA079FF96C0AE8D7DF82CC2EB" authority="(Moser, 1915)" baseAuthorityName="Moser" baseAuthorityYear="1915" box="[151,571,365,394]" class="Insecta" family="Scarabaeidae" genus="Maladera" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="381" phylum="Arthropoda" rank="species" species="basalis">
<emphasis id="B943E75BA079FF96C0AE8D7DF82CC2EB" bold="true" box="[151,571,365,394]" pageId="10" pageNumber="381">
<emphasis id="B943E75BA079FF96C0AE8D7DFB6DC2EB" bold="true" box="[151,378,365,394]" italics="true" pageId="10" pageNumber="381">Maladera basalis</emphasis>
(Moser, 1915)
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<emphasis id="B943E75BA079FF96C0AE8DA7FB91C2B0" bold="true" box="[151,390,439,465]" pageId="10" pageNumber="381">Material examined.</emphasis>
<materialsCitation id="3B5F3114A079FF96C1AA8DA7FAE9C294" collectingDate="1955-11-02" collectionCode="ZMUH" country="India" location="Fanvelbei" pageId="10" pageNumber="381" specimenCount="2" specimenCount-female="1" specimenCount-male="1">
<specimenCount id="9D31F0C0A079FF96C1AA8DA7FBD2C2B3" box="[403,453,439,466]" count="1" pageId="10" pageNumber="381" type="male">1 ♂</specimenCount>
,
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<location id="8EE86D92A079FF96C2248DA7F886C2B0" LSID="urn:lsid:plazi:treatment:039E8A5FA079FF96C0AE8C87FB14C294:8EE86D92A079FF96C2248DA7F886C2B0" box="[541,657,439,465]" country="India" name="Fanvelbei" pageId="10" pageNumber="381">Fanvelbei</location>
,
<collectingCountry id="F3207BD9A079FF96C2988DA7F916C2B0" box="[673,769,439,465]" name="India" pageId="10" pageNumber="381">Bombay</collectingCountry>
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<collectingDate id="EFCDE461A079FF96C32B8DA7F983C2B3" box="[786,916,439,466]" pageId="10" pageNumber="381" value="1955-11-02">02.XI.1955</collectingDate>
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, D.
<collectingCountry id="F3207BD9A079FF96C3F48DA7FE02C2B0" box="[973,1045,439,465]" name="India" pageId="10" pageNumber="381">Indien</collectingCountry>
Exp.Nr 533, Eing. Nr, 1, 1956” (
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)
</materialsCitation>
.
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View file

@ -0,0 +1,108 @@
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<emphasis id="B943E75BA079FF96C0AE8C87F85DC3D0" bold="true" box="[151,586,151,178]" pageId="10" pageNumber="381">
<emphasis id="B943E75BA079FF96C0AE8C87FB75C3D0" bold="true" box="[151,354,151,178]" italics="true" pageId="10" pageNumber="381">Maladera affinis</emphasis>
(
<bibRefCitation id="EFA646B8A079FF96C1488C87F856C3D0" author="Blanchard, C." box="[369,577,151,177]" pageId="10" pageNumber="381" pagination="1 - 128" refId="ref10257" refString="Blanchard, C. (1850) Ier Famille - Scarabaeidae. In: Milne-Edwards, H., Blanchard, C. E. &amp; Lucas, H. (Eds.), Catalogue de la collection entomologique du Museum d'Histoire Naturelle de Paris. Classe des insectes. Ordre des Coleopteres. Tome I. Premiere livraison. Gide et Baudry, Paris, pp. 1-128." type="book chapter" year="1850">Blanchard, 1850</bibRefCitation>
)
</emphasis>
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<emphasis id="B943E75BA079FF96C0AE8CCFFB6EC398" bold="true" box="[151,377,223,249]" pageId="10" pageNumber="381">Material examined</emphasis>
.
<materialsCitation id="3B5F3114A079FF96C1BE8CCFFF42C398" box="[391,1365,223,250]" collectingDate="1956-06-15" collectionCode="ZMUH" country="India" county="Nr" location="Timli" municipality="Exp" pageId="10" pageNumber="381" specimenCount="1" specimenCount-male="1">
<specimenCount id="9D31F0C0A079FF96C1BE8CCFFBA6C39B" box="[391,433,223,250]" count="1" pageId="10" pageNumber="381" type="male">1 ♂</specimenCount>
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,
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, D.
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,
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1284, Eing. Nr, 1, 1956” (
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</materialsCitation>
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View file

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<taxonomicName id="4C3740CAA079FF96C0AE8D7DF82CC2EB" ID-CoL="7ZP8J" authority="(Moser, 1915)" baseAuthorityName="Moser" baseAuthorityYear="1915" box="[151,571,365,394]" class="Insecta" family="Scarabaeidae" genus="Maladera" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="381" phylum="Arthropoda" rank="species" species="basalis">
<emphasis id="B943E75BA079FF96C0AE8D7DF82CC2EB" bold="true" box="[151,571,365,394]" pageId="10" pageNumber="381">
<emphasis id="B943E75BA079FF96C0AE8D7DFB6DC2EB" bold="true" box="[151,378,365,394]" italics="true" pageId="10" pageNumber="381">Maladera basalis</emphasis>
(Moser, 1915)
</emphasis>
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="C32D68C2A079FF96C0AE8DA7FB14C294" pageId="10" pageNumber="381" type="materials_examined">
<paragraph id="8B883B49A079FF96C0AE8DA7FB14C294" blockId="10.[151,1436,439,501]" pageId="10" pageNumber="381">
<emphasis id="B943E75BA079FF96C0AE8DA7FB91C2B0" bold="true" box="[151,390,439,465]" pageId="10" pageNumber="381">Material examined.</emphasis>
<materialsCitation id="3B5F3114A079FF96C1AA8DA7FAE9C294" collectingDate="1955-11-02" collectionCode="ZMUH" country="India" location="Fanvelbei" pageId="10" pageNumber="381" specimenCount="2" specimenCount-female="1" specimenCount-male="1">
<specimenCount id="9D31F0C0A079FF96C1AA8DA7FBD2C2B3" box="[403,453,439,466]" count="1" pageId="10" pageNumber="381" type="male">1 ♂</specimenCount>
,
<specimenCount id="9D31F0C0A079FF96C1EF8DA7F811C2B3" box="[470,518,439,466]" count="1" pageId="10" pageNumber="381" type="female">1 ♀</specimenCount>
<location id="8EE86D92A079FF96C2248DA7F886C2B0" LSID="urn:lsid:plazi:treatment:039E8A5FA079FF96C0AE8D7DFB14C294:8EE86D92A079FF96C2248DA7F886C2B0" box="[541,657,439,465]" country="India" name="Fanvelbei" pageId="10" pageNumber="381">Fanvelbei</location>
,
<collectingCountry id="F3207BD9A079FF96C2988DA7F916C2B0" box="[673,769,439,465]" name="India" pageId="10" pageNumber="381">Bombay</collectingCountry>
,
<date id="FF891D89A079FF96C32B8DA7F983C2B3" box="[786,916,439,466]" pageId="10" pageNumber="381" value="1955-11-02">
<collectingDate id="EFCDE461A079FF96C32B8DA7F983C2B3" box="[786,916,439,466]" pageId="10" pageNumber="381" value="1955-11-02">02.XI.1955</collectingDate>
</date>
, D.
<collectingCountry id="F3207BD9A079FF96C3F48DA7FE02C2B0" box="[973,1045,439,465]" name="India" pageId="10" pageNumber="381">Indien</collectingCountry>
Exp.Nr 533, Eing. Nr, 1, 1956” (
<collectionCode id="ED26A38CA079FF96C0A68DCBFAE4C294" box="[159,243,475,501]" country="Viet Nam" httpUri="http://grbio.org/cool/82c5-17ij" name="Zoological Museum, University of Hanoi" pageId="10" pageNumber="381">ZMUH</collectionCode>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>

View file

@ -0,0 +1,175 @@
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<paragraph id="8BC136B9151CFFEEFF23FF7DFD78FF6F" blockId="11.[151,655,151,178]" box="[151,655,151,178]" pageId="11" pageNumber="182">
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Key to described species of
<taxonomicName id="4C7E4D3A151CFFEEFE5AFF7DFD78FF6F" ID-CoL="7FSN" authorityName="Shaw" authorityYear="1985" box="[494,655,151,178]" class="Insecta" family="Braconidae" genus="Sericobracon" kingdom="Animalia" order="Hymenoptera" pageId="11" pageNumber="182" phylum="Arthropoda" rank="genus">
<emphasis id="B90AEAAB151CFFEEFE5AFF7DFD78FF6F" bold="true" box="[494,655,151,178]" italics="true" pageId="11" pageNumber="182">Sericobracon</emphasis>
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<subSubSection id="C3646532151CFFEEFF23FF36FA6BFD8F" pageId="11" pageNumber="182" type="key">
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<paragraph id="8BC136B9151CFFEEFF23FF36FA6BFF2C" blockId="11.[151,1437,220,594]" box="[151,1436,220,242]" pageId="11" pageNumber="182">
<keyLead id="308F958C151CFFEEFF23FF36FA6BFF2C" box="[151,1436,220,242]" pageId="11" pageNumber="182">
1. Mesopleuron mostly smooth (as in
<figureCitation id="13452A3C151CFFEEFD81FF36FD77FF2F" box="[565,640,220,242]" captionStart="FIGURE 5" captionStartId="8.[152,255,1904,1928]" captionTargetBox="[152,1435,561,1881]" captionTargetId="figure-189@8.[152,1435,561,1881]" captionTargetPageId="8" captionText="FIGURE 5. Sericobracon puravida sp. nov. Holotype, female. A) Habitus, lateral view; B) head, frontal view; C) head and mesosoma, dorsal view; D) mesosoma, lateral view; metasoma, E) dorsal view; E) fore wing." figureDoi="http://doi.org/10.5281/zenodo.15216365" httpUri="https://zenodo.org/record/15216365/files/figure.png" pageId="11" pageNumber="182">Fig. 5D</figureCitation>
)................................................................ 2
</keyLead>
</paragraph>
<paragraph id="8BC136B9151CFFEEFF23FF10FA6BFED2" blockId="11.[151,1437,220,594]" box="[151,1436,249,271]" pageId="11" pageNumber="182">
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- Mesopleuron mostly coriaceous (as in
<figureCitation id="13452A3C151CFFEEFDE1FF13FD69FED2" box="[597,670,249,271]" captionStart="FIGURE 3" captionStartId="5.[152,255,1060,1084]" captionTargetBox="[151,1435,181,1036]" captionTargetId="figure-15@5.[151,1435,181,1036]" captionTargetPageId="5" captionText="FIGURE 3. Sericobracon evansi Shaw. Holotype, female. A) Mesosoma and metasoma, lateral view; B) mesosoma, lateral view; C) mesosoma, dorsal view; D) metasoma, dorsal view. Museum of Comparative Zoology, Harvard University; © President and Fellows of Harvard College." figureDoi="http://doi.org/10.5281/zenodo.15216361" httpUri="https://zenodo.org/record/15216361/files/figure.png" pageId="11" pageNumber="182">Fig. 3B</figureCitation>
)............................................................. 3
</keyLead>
</paragraph>
</keyStep>
<keyStep id="308A2E1C151CFFEEFF23FEFDFA6BFE62" pageId="11" pageNumber="182">
<paragraph id="8BC136B9151CFFEEFF23FEFDFA6BFEBA" blockId="11.[151,1437,220,594]" pageId="11" pageNumber="182">
<keyLead id="308F958C151CFFEEFF23FEFDFA6BFEBA" pageId="11" pageNumber="182">
2. Second metasomal tergite with posterior border of longitudinally striate area broadly “V” shaped and shortest medially (
<figureCitation id="13452A3C151CFFEEFAC1FEFDFEF7FE94" captionStart="FIGURE 4" captionStartId="6.[152,255,1679,1703]" captionTargetBox="[177,1411,525,1655]" captionTargetId="figure-166@6.[177,1411,525,1655]" captionTargetPageId="6" captionText="FIGURE 4. Sericobracon paulmarshi sp. nov. Holotype, female.A) Habitus, lateral view; B) head and mesosoma, dorsal view; C) head, frontal view; D) head and mesosoma, lateral view; D) metasoma, dorsal view." figureDoi="http://doi.org/10.5281/zenodo.15216363" httpUri="https://zenodo.org/record/15216363/files/figure.png" pageId="11" pageNumber="182">Fig. 4E</figureCitation>
); notauli becoming obscure before end of mesoscutum in a median longitudinally striate area (
<figureCitation id="13452A3C151CFFEEFB3FFEDEFB22FE94" box="[1163,1237,308,330]" captionStart="FIGURE 4" captionStartId="6.[152,255,1679,1703]" captionTargetBox="[177,1411,525,1655]" captionTargetId="figure-166@6.[177,1411,525,1655]" captionTargetPageId="6" captionText="FIGURE 4. Sericobracon paulmarshi sp. nov. Holotype, female.A) Habitus, lateral view; B) head and mesosoma, dorsal view; C) head, frontal view; D) head and mesosoma, lateral view; D) metasoma, dorsal view." figureDoi="http://doi.org/10.5281/zenodo.15216363" httpUri="https://zenodo.org/record/15216363/files/figure.png" pageId="11" pageNumber="182">Fig. 4B</figureCitation>
).................................................................................................
<taxonomicName id="4C7E4D3A151CFFEEFB70FEB8FABEFEBB" authorityName="Zaldívar-Riverón &amp; Castañeda-Osorio &amp; Shaw" authorityYear="2025" box="[1220,1353,337,359]" class="Insecta" family="Braconidae" genus="Sericobracon" kingdom="Animalia" order="Hymenoptera" pageId="11" pageNumber="182" phylum="Arthropoda" rank="species" species="paulmarshi" status="sp. nov.">
<emphasis id="B90AEAAB151CFFEEFB70FEB8FABEFEBB" box="[1220,1353,337,359]" italics="true" pageId="11" pageNumber="182">S. paulmarshi</emphasis>
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<taxonomicNameLabel id="A23957D0151CFFEEFAE4FEB8FA6BFEBA" box="[1360,1436,338,359]" pageId="11" pageNumber="182" rank="species">sp. nov.</taxonomicNameLabel>
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<paragraph id="8BC136B9151CFFEEFF23FE85FA6BFE62" blockId="11.[151,1437,220,594]" pageId="11" pageNumber="182">
<keyLead id="308F958C151CFFEEFF23FE85FA6BFE62" pageId="11" pageNumber="182">
- Second metasomal tergite with posterior border of longitudinally striate area linear, running straight across the segment, not “V” shaped (
<figureCitation id="13452A3C151CFFEEFEEAFE66FE5FFE7F" box="[350,424,396,418]" captionStart="FIGURE 5" captionStartId="8.[152,255,1904,1928]" captionTargetBox="[152,1435,561,1881]" captionTargetId="figure-189@8.[152,1435,561,1881]" captionTargetPageId="8" captionText="FIGURE 5. Sericobracon puravida sp. nov. Holotype, female. A) Habitus, lateral view; B) head, frontal view; C) head and mesosoma, dorsal view; D) mesosoma, lateral view; metasoma, E) dorsal view; E) fore wing." figureDoi="http://doi.org/10.5281/zenodo.15216365" httpUri="https://zenodo.org/record/15216365/files/figure.png" pageId="11" pageNumber="182">Fig. 5E</figureCitation>
); notauli becoming obscure before end of mesoscutum in a slightly rugose area (
<figureCitation id="13452A3C151CFFEEFB1BFE66FB0FFE7F" box="[1199,1272,396,418]" captionStart="FIGURE 5" captionStartId="8.[152,255,1904,1928]" captionTargetBox="[152,1435,561,1881]" captionTargetId="figure-189@8.[152,1435,561,1881]" captionTargetPageId="8" captionText="FIGURE 5. Sericobracon puravida sp. nov. Holotype, female. A) Habitus, lateral view; B) head, frontal view; C) head and mesosoma, dorsal view; D) mesosoma, lateral view; metasoma, E) dorsal view; E) fore wing." figureDoi="http://doi.org/10.5281/zenodo.15216365" httpUri="https://zenodo.org/record/15216365/files/figure.png" pageId="11" pageNumber="182">Fig. 5C</figureCitation>
)................................................................................................
<taxonomicName id="4C7E4D3A151CFFEEFB6FFE40FABDFE63" authorityName="Zaldívar-Riverón &amp; Castañeda-Osorio &amp; Shaw" authorityYear="2025" box="[1243,1354,425,447]" class="Insecta" family="Braconidae" genus="Sericobracon" kingdom="Animalia" order="Hymenoptera" pageId="11" pageNumber="182" phylum="Arthropoda" rank="species" species="puravida" status="sp. nov.">
<emphasis id="B90AEAAB151CFFEEFB6FFE40FABDFE63" box="[1243,1354,425,447]" italics="true" pageId="11" pageNumber="182">S. puravida</emphasis>
</taxonomicName>
<emphasis id="B90AEAAB151CFFEEFAE4FE40FA6BFE62" bold="true" box="[1360,1436,426,447]" pageId="11" pageNumber="182">
<taxonomicNameLabel id="A23957D0151CFFEEFAE4FE40FA6BFE62" box="[1360,1436,426,447]" pageId="11" pageNumber="182" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</keyLead>
</paragraph>
</keyStep>
<keyStep id="308A2E1C151CFFEEFF23FE2DFA6BFE24" pageId="11" pageNumber="182">
<paragraph id="8BC136B9151CFFEEFF23FE2DFA6BFE01" blockId="11.[151,1437,220,594]" box="[151,1436,455,477]" pageId="11" pageNumber="182">
<keyLead id="308F958C151CFFEEFF23FE2DFA6BFE01" box="[151,1436,455,477]" pageId="11" pageNumber="182">
3. Propleuron, pronotum and most part of mesopleuron whitish yellow to honey yellow (
<figureCitation id="13452A3C151CFFEEFBA8FE2DFB90FE00" box="[1052,1127,455,477]" captionStart="FIGURE 6" captionStartId="10.[152,255,1444,1468]" captionTargetBox="[152,1435,181,1420]" captionTargetId="figure-27@10.[152,1435,181,1420]" captionTargetPageId="10" captionText="FIGURE 6. Sericobracon zunigai sp. nov. Holotype, female. A) Habitus, lateral view; B) head, frontal view; C) head and mesosoma, dorsal view; D) mesosoma, lateral view; E) metasoma, dorsal view; F) fore wing." figureDoi="http://doi.org/10.5281/zenodo.15216367" httpUri="https://zenodo.org/record/15216367/files/figure.png" pageId="11" pageNumber="182">Fig. 6D</figureCitation>
).........
<taxonomicName id="4C7E4D3A151CFFEEFB5FFE2DFABDFE01" authorityName="Zaldívar-Riverón &amp; Castañeda-Osorio &amp; Shaw" authorityYear="2025" box="[1259,1354,455,476]" class="Insecta" family="Braconidae" genus="Sericobracon" kingdom="Animalia" order="Hymenoptera" pageId="11" pageNumber="182" phylum="Arthropoda" rank="species" species="zunigai" status="sp. nov.">
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<taxonomicNameLabel id="A23957D0151CFFEEFAE4FE2DFA6BFE01" box="[1360,1436,455,476]" pageId="11" pageNumber="182" rank="species">sp. nov.</taxonomicNameLabel>
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<keyLead id="308F958C151CFFEEFF23FE0FFA6BFE24" box="[151,1436,484,506]" pageId="11" pageNumber="182">
- Propleuron, pronotum and most part of mesopleuron mostly dark brown or black (as in
<figureCitation id="13452A3C151CFFEEFB9EFE0EFB84FE24" box="[1066,1139,484,506]" captionStart="FIGURE 3" captionStartId="5.[152,255,1060,1084]" captionTargetBox="[151,1435,181,1036]" captionTargetId="figure-15@5.[151,1435,181,1036]" captionTargetPageId="5" captionText="FIGURE 3. Sericobracon evansi Shaw. Holotype, female. A) Mesosoma and metasoma, lateral view; B) mesosoma, lateral view; C) mesosoma, dorsal view; D) metasoma, dorsal view. Museum of Comparative Zoology, Harvard University; © President and Fellows of Harvard College." figureDoi="http://doi.org/10.5281/zenodo.15216361" httpUri="https://zenodo.org/record/15216361/files/figure.png" pageId="11" pageNumber="182">Fig. 3B</figureCitation>
)...................... 4
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4. First metasomal tergite longitudinally striate; metapleuron mostly strongly rugose-areolate (
<figureCitation id="13452A3C151CFFEEFBE0FDEBFB68FDCA" box="[1108,1183,513,535]" captionStart="FIGURE 2" captionStartId="4.[152,255,1921,1945]" captionTargetBox="[152,1436,956,1896]" captionTargetId="figure-234@4.[152,1436,956,1896]" captionTargetPageId="4" captionText="FIGURE 2. Sericobracon arimaensis Shaw. Holotype, female. A) Habitus, lateral view; B) head, frontal view; C) head and mesosoma, dorsal view; D) mesosoma, lateral view. Museum of Comparative Zoology, Harvard University; © President and Fellows of Harvard College." figureDoi="http://doi.org/10.5281/zenodo.15216359" httpUri="https://zenodo.org/record/15216359/files/figure.png" pageId="11" pageNumber="182">Fig. 2D</figureCitation>
)...
<taxonomicName id="4C7E4D3A151CFFEEFB54FDE8FA6BFDCA" ID-CoL="6YJ37" authority="Shaw" authorityName="Shaw" box="[1248,1436,513,535]" class="Insecta" family="Braconidae" genus="Sericobracon" kingdom="Animalia" order="Hymenoptera" pageId="11" pageNumber="182" phylum="Arthropoda" rank="species" species="arimaensis">
<emphasis id="B90AEAAB151CFFEEFB54FDE8FA96FDCA" box="[1248,1377,514,535]" italics="true" pageId="11" pageNumber="182">S. arimaensis</emphasis>
Shaw
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<keyLead id="308F958C151CFFEEFF23FDF5FA6BFD8F" pageId="11" pageNumber="182">
- First metasomal tergite smooth to slightly longitudinally striate (
<figureCitation id="13452A3C151CFFEEFCE1FDF5FC54FDE9" box="[853,931,543,565]" captionStart="FIGURE 3" captionStartId="5.[152,255,1060,1084]" captionTargetBox="[151,1435,181,1036]" captionTargetId="figure-15@5.[151,1435,181,1036]" captionTargetPageId="5" captionText="FIGURE 3. Sericobracon evansi Shaw. Holotype, female. A) Mesosoma and metasoma, lateral view; B) mesosoma, lateral view; C) mesosoma, dorsal view; D) metasoma, dorsal view. Museum of Comparative Zoology, Harvard University; © President and Fellows of Harvard College." figureDoi="http://doi.org/10.5281/zenodo.15216361" httpUri="https://zenodo.org/record/15216361/files/figure.png" pageId="11" pageNumber="182">Fig. 3D</figureCitation>
); metapleuron coriaceous-slightly rugose (
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)......................................................................................
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<emphasis id="B90AEAAB151CFFEEFAB8FDD6FA96FD8F" box="[1292,1377,572,594]" italics="true" pageId="11" pageNumber="182">S. evansi</emphasis>
Shaw
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The raw data obtained from the overview study comprised the allozyme genotypes from
@ -72,7 +74,7 @@ of
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representing 45 populations overall. These data are summarized in
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<tableCitation id="D25D031FFFDCFFF1FE056CD52C5996AC" box="[432,518,1761,1783]" captionStart="Table 2" captionStartId="7.[147,211,601,622]" captionTargetBox="[147,1848,776,1493]" captionText="Table 2. Allozyme frequencies at 45 loci for all previously recognized forms of Chelodina examined in the overview study. Two additional loci, Hb and Gp, could not be scored across all taxa but were included in various Stage 2 analyses. Sample sizes for each OTU are presented in parentheses, with the first value representing the number included in the overview study and any second value representing the number subsequently examined in the Stage 2 analyses. Where a locus is polymorphic within a population, the frequency of each common allozyme is given as a percentage; the frequency of the rarest allozyme can be calculated by subtraction. Taxon abbreviations are given in Table 1" httpUri="http://table.plazi.org/id/CBA0662CFFDDFFF0FF26686D2C9592B8" pageId="6" pageNumber="407" tableDoi="http://doi.org/10.5281/zenodo.15223995" tableUuid="CBA0662CFFDDFFF0FF26686D2C9592B8">Table 2</tableCitation>
as allele frequencies for each diagnosable taxon (including the morphologically diagnosable
<taxonomicName id="58DF4D27FFDCFFF1FED06D2B2FAD976F" box="[357,498,1823,1844]" class="Reptilia" family="Chelidae" genus="Chelodina" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="407" phylum="Chordata" rank="species" species="reimanni">
<emphasis id="ADABEAB6FFDCFFF1FED06D2B2FAD976F" box="[357,498,1823,1844]" italics="true" pageId="6" pageNumber="407">C. reimanni</emphasis>
@ -82,7 +84,7 @@ plus the various geographical/putative forms of the
<emphasis id="ADABEAB6FFDCFFF1FE466D0A2C009708" box="[499,607,1854,1875]" italics="true" pageId="6" pageNumber="407">C. rugosa</emphasis>
</taxonomicName>
group).
<tableCitation id="D25D031FFFDCFFF1FD0F6D092EC0972A" captionStart="Table 3" captionStartId="9.[159,223,196,217]" captionTargetPageId="9" captionText="Table 3. Subsets of taxa and allozyme loci examined in the Stage 2 analysis. Species abbreviations are as for Table 1" pageId="6" pageNumber="407">Table 3</tableCitation>
<tableCitation id="D25D031FFFDCFFF1FD0F6D092EC0972A" captionStart="Table 3" captionStartId="9.[159,223,196,217]" captionTargetPageId="9" captionText="Table 3. Subsets of taxa and allozyme loci examined in the Stage 2 analysis. Species abbreviations are as for Table 1" httpUri="http://table.plazi.org/id/CBA0662CFFD3FFFEFF2A6AF02BDD9082" pageId="6" pageNumber="407" tableDoi="http://doi.org/10.5281/zenodo.15223997" tableUuid="CBA0662CFFD3FFFEFF2A6AF02BDD9082">Table 3</tableCitation>
gives details of the subgroups that were subjected to further screening in Stage 2 of the analysis. The genotypic data for both the overview and various Stage 2 analyses are not presented due to considerations of space, but are available from the authors upon request.
</paragraph>
<paragraph id="9F6036A4FFDCFFF1FCF46B6B2DF39255" blockId="6.[809,1425,197,1906]" pageId="6" pageNumber="407">It was clear from the outset that four individuals in the overview study and one in the Stage 2 analyses had hybrid origins. These specimens were omitted from the species boundary analyses. The analysis of their hybrid origins is presented separately, later in the results.</paragraph>
@ -120,7 +122,7 @@ and
<emphasis id="ADABEAB6FFDCFFF1FC0E697D2A2C9305" box="[955,1139,841,862]" italics="true" pageId="6" pageNumber="407">C. steindachneri</emphasis>
</taxonomicName>
, the two taxa which were sampled from multiple localities across their entire range, were examined for a suite of diagnostic and polymorphic loci in Stage 2 (
<tableCitation id="D25D031FFFDCFFF1FBD869912A9D93E0" box="[1133,1218,933,955]" captionStart="Table 3" captionStartId="9.[159,223,196,217]" captionTargetPageId="9" captionText="Table 3. Subsets of taxa and allozyme loci examined in the Stage 2 analysis. Species abbreviations are as for Table 1" pageId="6" pageNumber="407">Table 3</tableCitation>
<tableCitation id="D25D031FFFDCFFF1FBD869912A9D93E0" box="[1133,1218,933,955]" captionStart="Table 3" captionStartId="9.[159,223,196,217]" captionTargetPageId="9" captionText="Table 3. Subsets of taxa and allozyme loci examined in the Stage 2 analysis. Species abbreviations are as for Table 1" httpUri="http://table.plazi.org/id/CBA0662CFFD3FFFEFF2A6AF02BDD9082" pageId="6" pageNumber="407" tableDoi="http://doi.org/10.5281/zenodo.15223997" tableUuid="CBA0662CFFD3FFFEFF2A6AF02BDD9082">Table 3</tableCitation>
). The genetic profiles of these additional animals confirmed both their provisional identification and the utility of the diagnostic loci employed. Of the two taxa, only
<taxonomicName id="58DF4D27FFDCFFF1FAA06E352BCE944C" box="[1301,1425,1025,1047]" class="Reptilia" family="Chelidae" genus="Chelodina" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="407" phylum="Chordata" rank="species" species="expansa">
<emphasis id="ADABEAB6FFDCFFF1FAA06E352BCE944C" box="[1301,1425,1025,1047]" italics="true" pageId="6" pageNumber="407">C. expansa</emphasis>
@ -141,7 +143,7 @@ and
<emphasis id="ADABEAB6FFDCFFF1FB886F212A839571" box="[1085,1244,1301,1322]" italics="true" pageId="6" pageNumber="407">C. siebenrocki</emphasis>
</taxonomicName>
split into three interim diagnosable taxa that did not correspond with existing species boundaries. These three interim taxa were combined as one subgroup for further detailed analysis using all available specimens (
<tableCitation id="D25D031FFFDCFFF1FB5A6FA42B1995FD" box="[1263,1350,1424,1446]" captionStart="Table 3" captionStartId="9.[159,223,196,217]" captionTargetPageId="9" captionText="Table 3. Subsets of taxa and allozyme loci examined in the Stage 2 analysis. Species abbreviations are as for Table 1" pageId="6" pageNumber="407">Table 3</tableCitation>
<tableCitation id="D25D031FFFDCFFF1FB5A6FA42B1995FD" box="[1263,1350,1424,1446]" captionStart="Table 3" captionStartId="9.[159,223,196,217]" captionTargetPageId="9" captionText="Table 3. Subsets of taxa and allozyme loci examined in the Stage 2 analysis. Species abbreviations are as for Table 1" httpUri="http://table.plazi.org/id/CBA0662CFFD3FFFEFF2A6AF02BDD9082" pageId="6" pageNumber="407" tableDoi="http://doi.org/10.5281/zenodo.15223997" tableUuid="CBA0662CFFD3FFFEFF2A6AF02BDD9082">Table 3</tableCitation>
). Similarly, forms currently assigned to
<taxonomicName id="58DF4D27FFDCFFF1FB046F9B2B35959F" box="[1201,1386,1455,1476]" class="Reptilia" family="Chelidae" genus="Chelodina" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="407" phylum="Chordata" rank="species" species="novaeguineae">
<emphasis id="ADABEAB6FFDCFFF1FB046F9B2B35959F" box="[1201,1386,1455,1476]" italics="true" pageId="6" pageNumber="407">C. novaeguineae</emphasis>
@ -163,12 +165,12 @@ and
<emphasis id="ADABEAB6FFDCFFF1FB496FF92BCE95B9" box="[1276,1425,1485,1506]" italics="true" pageId="6" pageNumber="407">C. longicollis</emphasis>
</taxonomicName>
were combined as a separate subgroup (
<tableCitation id="D25D031FFFDCFFF1FB426FD82B119659" box="[1271,1358,1516,1538]" captionStart="Table 3" captionStartId="9.[159,223,196,217]" captionTargetPageId="9" captionText="Table 3. Subsets of taxa and allozyme loci examined in the Stage 2 analysis. Species abbreviations are as for Table 1" pageId="6" pageNumber="407">Table 3</tableCitation>
<tableCitation id="D25D031FFFDCFFF1FB426FD82B119659" box="[1271,1358,1516,1538]" captionStart="Table 3" captionStartId="9.[159,223,196,217]" captionTargetPageId="9" captionText="Table 3. Subsets of taxa and allozyme loci examined in the Stage 2 analysis. Species abbreviations are as for Table 1" httpUri="http://table.plazi.org/id/CBA0662CFFD3FFFEFF2A6AF02BDD9082" pageId="6" pageNumber="407" tableDoi="http://doi.org/10.5281/zenodo.15223997" tableUuid="CBA0662CFFD3FFFEFF2A6AF02BDD9082">Table 3</tableCitation>
).
</paragraph>
<paragraph id="9F6036A4FFDCFFF1FCF46C3F2BD5976F" blockId="6.[809,1425,197,1906]" pageId="6" pageNumber="407">
A summary of the number of fixed differences between taxa that were diagnosable at one or more loci is provided in
<tableCitation id="D25D031FFFDCFFF1FC646C7C2A769606" box="[977,1065,1608,1630]" captionStart="Table 4" captionStartId="10.[145,209,196,217]" captionTargetPageId="10" captionText="Table 4. Raw fixed differences (lower matrix) and Rogers D (upper matrix) for all forms of Chelodina that were diagnosable at one or more loci in our sample of populations. Species abbreviations are as for Table 1" pageId="6" pageNumber="407">Table 4</tableCitation>
<tableCitation id="D25D031FFFDCFFF1FC646C7C2A769606" box="[977,1065,1608,1630]" captionStart="Table 4" captionStartId="10.[145,209,196,217]" captionTargetPageId="10" captionText="Table 4. Raw fixed differences (lower matrix) and Rogers D (upper matrix) for all forms of Chelodina that were diagnosable at one or more loci in our sample of populations. Species abbreviations are as for Table 1" httpUri="http://table.plazi.org/id/CBA0662CFFD0FFFDFF246AF02AC790AD" pageId="6" pageNumber="407" tableDoi="http://doi.org/10.5281/zenodo.15223999" tableUuid="CBA0662CFFD0FFFDFF246AF02AC790AD">Table 4</tableCitation>
. This table was constructed by integrating the overview analysis and the various subgroup analyses on the assumption that the monomorphic loci from the overview analysis that were not further analysed would have remained largely monomorphic (they certainly would have failed to become in any way diagnostic) had they been subject to greater scrutiny in the subgroup analyses.
</paragraph>
<paragraph id="9F6036A4FFDCFFFEFCF46D092B019428" blockId="6.[809,1425,197,1906]" lastBlockId="9.[823,1439,657,1599]" lastPageId="9" lastPageNumber="410" pageId="6" pageNumber="407">
@ -286,7 +288,7 @@ Thomson
</materialsCitation>
.
</paragraph>
<caption id="CBA0662CFFDDFFF0FF26686D2C9592B8" pageId="7" pageNumber="408" startId="7.[147,211,601,622]" targetBox="[147,1848,776,1493]" targetIsTable="true" targetPageId="7">
<caption id="CBA0662CFFDDFFF0FF26686D2C9592B8" ID-DOI="http://doi.org/10.5281/zenodo.15223995" ID-Table-UUID="CBA0662CFFDDFFF0FF26686D2C9592B8" ID-Zenodo-Dep="15223995" httpUri="http://table.plazi.org/id/CBA0662CFFDDFFF0FF26686D2C9592B8" pageId="7" pageNumber="408" startId="7.[147,211,601,622]" targetBox="[147,1848,776,1493]" targetIsTable="true" targetPageId="7">
<paragraph id="9F6036A4FFDDFFF0FF26686D2C9592B8" blockId="7.[147,1851,601,739]" pageId="7" pageNumber="408">
<emphasis id="ADABEAB6FFDDFFF0FF26686D2EB19235" bold="true" box="[147,238,601,622]" pageId="7" pageNumber="408">Table 2.</emphasis>
Allozyme frequencies at 45 loci for all previously recognized forms of
@ -1293,7 +1295,7 @@ and
</tr>
</table>
</paragraph>
<caption id="CBA0662CFFD3FFFEFF2A6AF02BDD9082" box="[159,1410,196,217]" pageId="9" pageNumber="410" startId="9.[159,223,196,217]" targetBox="[159,1436,254,551]" targetIsTable="true" targetPageId="9">
<caption id="CBA0662CFFD3FFFEFF2A6AF02BDD9082" ID-DOI="http://doi.org/10.5281/zenodo.15223997" ID-Table-UUID="CBA0662CFFD3FFFEFF2A6AF02BDD9082" ID-Zenodo-Dep="15223997" box="[159,1410,196,217]" httpUri="http://table.plazi.org/id/CBA0662CFFD3FFFEFF2A6AF02BDD9082" pageId="9" pageNumber="410" startId="9.[159,223,196,217]" targetBox="[159,1436,254,551]" targetIsTable="true" targetPageId="9">
<paragraph id="9F6036A4FFD3FFFEFF2A6AF02BDD9082" blockId="9.[159,1410,196,217]" box="[159,1410,196,217]" pageId="9" pageNumber="410">
<emphasis id="ADABEAB6FFD3FFFEFF2A6AF02EA59082" bold="true" box="[159,250,196,217]" pageId="9" pageNumber="410">Table 3.</emphasis>
Subsets of taxa and allozyme loci examined in the Stage 2 analysis. Species abbreviations are as for Table 1
@ -1535,7 +1537,7 @@ and
<emphasis id="ADABEAB6FFD3FFFEFC456EC32AD69550" box="[1008,1161,1270,1292]" italics="true" pageId="9" pageNumber="410">C. longicollis</emphasis>
</taxonomicName>
(
<figureCitation id="07E42A21FFD3FFFEFB2B6EC22AB89556" box="[1182,1255,1270,1293]" captionStart="Figure 4" captionStartId="10.[809,888,1502,1523]" captionTargetBox="[812,1425,901,1475]" captionTargetId="figure-386@10.[814,1419,927,1475]" captionTargetPageId="10" captionText="Figure 4. Principal components analysis applied to a matrix of Rogers D genetic distances between all individuals of Chelodina longicollis (O), C. mccordi (Δ), C. pritchardi (•), C. novaeguineae (New Guinea form) (), C. novaeguineae (Australian form) (O) and C. reimanni (Z). Two individuals which are hybrids between C. longicollis and C. novaeguineae (Australian form) are included (Ɨ). Five groups are evident. Chelodina longicollis, C. mccordi and C. pritchardi are well differentiated. Chelodina novaeguineae (New Guinea form) and C. reimanni are undifferentiated and may represent a single taxon (but see text). The Australian and New Guinea forms of C. novaeguineae are clearly differentiated, but only one difference has moved to fixation." pageId="9" pageNumber="410">Fig. 4</figureCitation>
<figureCitation id="07E42A21FFD3FFFEFB2B6EC22AB89556" box="[1182,1255,1270,1293]" captionStart="Figure 4" captionStartId="10.[809,888,1502,1523]" captionTargetBox="[812,1425,901,1475]" captionTargetId="figure-386@10.[814,1419,927,1475]" captionTargetPageId="10" captionText="Figure 4. Principal components analysis applied to a matrix of Rogers D genetic distances between all individuals of Chelodina longicollis (O), C. mccordi (Δ), C. pritchardi (•), C. novaeguineae (New Guinea form) (), C. novaeguineae (Australian form) (O) and C. reimanni (Z). Two individuals which are hybrids between C. longicollis and C. novaeguineae (Australian form) are included (Ɨ). Five groups are evident. Chelodina longicollis, C. mccordi and C. pritchardi are well differentiated. Chelodina novaeguineae (New Guinea form) and C. reimanni are undifferentiated and may represent a single taxon (but see text). The Australian and New Guinea forms of C. novaeguineae are clearly differentiated, but only one difference has moved to fixation." figureDoi="http://doi.org/10.5281/zenodo.15223895" httpUri="https://zenodo.org/record/15223895/files/figure.png" pageId="9" pageNumber="410">Fig. 4</figureCitation>
). Each differed from the other by 68 fixed differences and so represent clear diagnosable taxa. Two additional groupings corresponded to the Australian
<taxonomicName id="58DF4D27FFD3FFFEFB006F672B2C9533" box="[1205,1395,1363,1384]" class="Reptilia" family="Chelidae" genus="Chelodina" kingdom="Animalia" order="Testudines" pageId="9" pageNumber="410" phylum="Chordata" rank="species" species="novaeguineae">
<emphasis id="ADABEAB6FFD3FFFEFB006F672B2C9533" box="[1205,1395,1363,1384]" italics="true" pageId="9" pageNumber="410">C. novaeguineae</emphasis>
@ -1610,7 +1612,7 @@ and
<emphasis id="ADABEAB6FFD1FFFCFF2A6B6B2F08912F" box="[159,343,351,372]" italics="true" pageId="11" pageNumber="412">C. novaeguineae</emphasis>
</taxonomicName>
(
<tableCitation id="D25D031FFFD1FFFCFED06B6A2FE5912F" box="[357,442,350,372]" captionStart="Table 5" captionStartId="12.[145,209,196,217]" captionTargetBox="[145,1423,489,1918]" captionTargetPageId="12" captionText="Table 5. Parental allele frequencies and hybrid genotypes at the 21 loci where the percentage difference between any two parental taxa is greater than 50%. Each of the two alleles displayed by a hybrid animal is indicated with a cross (+). Parental profiles for Clo and Cno have largely been taken from the overview study (Table 2; data for Hb from Stage 2) whereas those for Cru (combining the NT and Qld populations) and Cbu (NT populations only) were obtained from the Stage 2 analyses. None of the 24 loci not presented show genotypes inconsistent with a hybrid origin. An asterisk indicates that the presence of the null allele Acycf has been assumed, given the observed electrophoretic phenotype (the same mobility as allozyme e but a much-reduced activity). Superscripts are used to indicate which loci are diagnostic for the various parental taxa: AClo vs. Cno; BCno vs. Cru; CCru vs. Cbu (or effectively diagnostic if Cru is also represented by the NT populations only). (na = genotype was not determined)" pageId="11" pageNumber="412">Table 5</tableCitation>
<tableCitation id="D25D031FFFD1FFFCFED06B6A2FE5912F" box="[357,442,350,372]" captionStart="Table 5" captionStartId="12.[145,209,196,217]" captionTargetBox="[145,1423,489,1918]" captionTargetPageId="12" captionText="Table 5. Parental allele frequencies and hybrid genotypes at the 21 loci where the percentage difference between any two parental taxa is greater than 50%. Each of the two alleles displayed by a hybrid animal is indicated with a cross (+). Parental profiles for Clo and Cno have largely been taken from the overview study (Table 2; data for Hb from Stage 2) whereas those for Cru (combining the NT and Qld populations) and Cbu (NT populations only) were obtained from the Stage 2 analyses. None of the 24 loci not presented show genotypes inconsistent with a hybrid origin. An asterisk indicates that the presence of the null allele Acycf has been assumed, given the observed electrophoretic phenotype (the same mobility as allozyme e but a much-reduced activity). Superscripts are used to indicate which loci are diagnostic for the various parental taxa: AClo vs. Cno; BCno vs. Cru; CCru vs. Cbu (or effectively diagnostic if Cru is also represented by the NT populations only). (na = genotype was not determined)" httpUri="http://table.plazi.org/id/CBA0662CFFD6FFFBFF246AF02CA49198" pageId="11" pageNumber="412" tableDoi="http://doi.org/10.5281/zenodo.15224001" tableUuid="CBA0662CFFD6FFFBFF246AF02CA49198">Table 5</tableCitation>
). Both animals displayed the heterozygous genotypes expected for a
<taxonomicName id="58DF4D27FFD1FFFCFDEB6B4A2CAD91C9" box="[606,754,381,403]" class="Reptilia" family="Chelidae" genus="Chelodina" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="412" phylum="Chordata" rank="species" species="longicollis">
<emphasis id="ADABEAB6FFD1FFFCFDEB6B4A2CAD91C9" box="[606,754,381,403]" italics="true" pageId="11" pageNumber="412">C. longicollis</emphasis>
@ -1646,10 +1648,10 @@ and Australian
<emphasis id="ADABEAB6FFD1FFFCFE2468FB2C1692BF" box="[401,585,719,740]" italics="true" pageId="11" pageNumber="412">C. novaeguineae</emphasis>
</taxonomicName>
clusters (
<figureCitation id="07E42A21FFD1FFFCFD0268FA2CA592BE" box="[695,762,718,741]" captionStart="Figure 4" captionStartId="10.[809,888,1502,1523]" captionTargetBox="[812,1425,901,1475]" captionTargetId="figure-386@10.[814,1419,927,1475]" captionTargetPageId="10" captionText="Figure 4. Principal components analysis applied to a matrix of Rogers D genetic distances between all individuals of Chelodina longicollis (O), C. mccordi (Δ), C. pritchardi (•), C. novaeguineae (New Guinea form) (), C. novaeguineae (Australian form) (O) and C. reimanni (Z). Two individuals which are hybrids between C. longicollis and C. novaeguineae (Australian form) are included (Ɨ). Five groups are evident. Chelodina longicollis, C. mccordi and C. pritchardi are well differentiated. Chelodina novaeguineae (New Guinea form) and C. reimanni are undifferentiated and may represent a single taxon (but see text). The Australian and New Guinea forms of C. novaeguineae are clearly differentiated, but only one difference has moved to fixation." pageId="11" pageNumber="412">Fig. 4</figureCitation>
<figureCitation id="07E42A21FFD1FFFCFD0268FA2CA592BE" box="[695,762,718,741]" captionStart="Figure 4" captionStartId="10.[809,888,1502,1523]" captionTargetBox="[812,1425,901,1475]" captionTargetId="figure-386@10.[814,1419,927,1475]" captionTargetPageId="10" captionText="Figure 4. Principal components analysis applied to a matrix of Rogers D genetic distances between all individuals of Chelodina longicollis (O), C. mccordi (Δ), C. pritchardi (•), C. novaeguineae (New Guinea form) (), C. novaeguineae (Australian form) (O) and C. reimanni (Z). Two individuals which are hybrids between C. longicollis and C. novaeguineae (Australian form) are included (Ɨ). Five groups are evident. Chelodina longicollis, C. mccordi and C. pritchardi are well differentiated. Chelodina novaeguineae (New Guinea form) and C. reimanni are undifferentiated and may represent a single taxon (but see text). The Australian and New Guinea forms of C. novaeguineae are clearly differentiated, but only one difference has moved to fixation." figureDoi="http://doi.org/10.5281/zenodo.15223895" httpUri="https://zenodo.org/record/15223895/files/figure.png" pageId="11" pageNumber="412">Fig. 4</figureCitation>
).
</paragraph>
<caption id="CBA0662CFFD0FFFDFF246AF02AC790AD" pageId="10" pageNumber="411" startId="10.[145,209,196,217]" targetBox="[145,1424,284,715]" targetIsTable="true" targetPageId="10">
<caption id="CBA0662CFFD0FFFDFF246AF02AC790AD" ID-DOI="http://doi.org/10.5281/zenodo.15223999" ID-Table-UUID="CBA0662CFFD0FFFDFF246AF02AC790AD" ID-Zenodo-Dep="15223999" httpUri="http://table.plazi.org/id/CBA0662CFFD0FFFDFF246AF02AC790AD" pageId="10" pageNumber="411" startId="10.[145,209,196,217]" targetBox="[145,1424,284,715]" targetIsTable="true" targetPageId="10">
<paragraph id="9F6036A4FFD0FFFDFF246AF02AC790AD" blockId="10.[145,1424,196,246]" pageId="10" pageNumber="411">
<emphasis id="ADABEAB6FFD0FFFDFF246AF02EB29082" bold="true" box="[145,237,196,217]" pageId="10" pageNumber="411">Table 4.</emphasis>
Raw fixed differences (lower matrix) and Rogers D (upper matrix) for all forms of
@ -1914,7 +1916,7 @@ are undifferentiated, and also probably represent a single taxon. The Northern T
represents the third group. These three groups differ each by only one fixed difference.
</paragraph>
</caption>
<caption id="CBA0662CFFD0FFFDFC9C6FEA2A38972A" pageId="10" pageNumber="411" startId="10.[809,888,1502,1523]" targetBox="[812,1425,901,1475]" targetPageId="10">
<caption id="CBA0662CFFD0FFFDFC9C6FEA2A38972A" ID-DOI="http://doi.org/10.5281/zenodo.15223895" ID-Zenodo-Dep="15223895" httpUri="https://zenodo.org/record/15223895/files/figure.png" pageId="10" pageNumber="411" startId="10.[809,888,1502,1523]" targetBox="[812,1425,901,1475]" targetPageId="10">
<paragraph id="9F6036A4FFD0FFFDFC9C6FEA2A38972A" blockId="10.[809,1425,1502,1905]" pageId="10" pageNumber="411">
<emphasis id="ADABEAB6FFD0FFFDFC9C6FEA2DC595A8" bold="true" box="[809,922,1502,1523]" pageId="10" pageNumber="411">Figure 4.</emphasis>
Principal components analysis applied to a matrix of Rogers D genetic distances between all individuals of
@ -2024,7 +2026,7 @@ and
, as it displays the heterozygous genotypes expected for an F
<subScript id="035B34E1FFD1FFFCFD4A6EB12D5894C8" attach="right" box="[767,775,1157,1171]" fontSize="6" pageId="11" pageNumber="412">1</subScript>
hybrid at all of the 14 loci that display absolute or effective fixed differences between the parental species, and no genotypic inconsistencies at any other locus (
<tableCitation id="D25D031FFFD1FFFCFF5D6EC22F609557" box="[232,319,1270,1292]" captionStart="Table 5" captionStartId="12.[145,209,196,217]" captionTargetBox="[145,1423,489,1918]" captionTargetPageId="12" captionText="Table 5. Parental allele frequencies and hybrid genotypes at the 21 loci where the percentage difference between any two parental taxa is greater than 50%. Each of the two alleles displayed by a hybrid animal is indicated with a cross (+). Parental profiles for Clo and Cno have largely been taken from the overview study (Table 2; data for Hb from Stage 2) whereas those for Cru (combining the NT and Qld populations) and Cbu (NT populations only) were obtained from the Stage 2 analyses. None of the 24 loci not presented show genotypes inconsistent with a hybrid origin. An asterisk indicates that the presence of the null allele Acycf has been assumed, given the observed electrophoretic phenotype (the same mobility as allozyme e but a much-reduced activity). Superscripts are used to indicate which loci are diagnostic for the various parental taxa: AClo vs. Cno; BCno vs. Cru; CCru vs. Cbu (or effectively diagnostic if Cru is also represented by the NT populations only). (na = genotype was not determined)" pageId="11" pageNumber="412">Table 5</tableCitation>
<tableCitation id="D25D031FFFD1FFFCFF5D6EC22F609557" box="[232,319,1270,1292]" captionStart="Table 5" captionStartId="12.[145,209,196,217]" captionTargetBox="[145,1423,489,1918]" captionTargetPageId="12" captionText="Table 5. Parental allele frequencies and hybrid genotypes at the 21 loci where the percentage difference between any two parental taxa is greater than 50%. Each of the two alleles displayed by a hybrid animal is indicated with a cross (+). Parental profiles for Clo and Cno have largely been taken from the overview study (Table 2; data for Hb from Stage 2) whereas those for Cru (combining the NT and Qld populations) and Cbu (NT populations only) were obtained from the Stage 2 analyses. None of the 24 loci not presented show genotypes inconsistent with a hybrid origin. An asterisk indicates that the presence of the null allele Acycf has been assumed, given the observed electrophoretic phenotype (the same mobility as allozyme e but a much-reduced activity). Superscripts are used to indicate which loci are diagnostic for the various parental taxa: AClo vs. Cno; BCno vs. Cru; CCru vs. Cbu (or effectively diagnostic if Cru is also represented by the NT populations only). (na = genotype was not determined)" httpUri="http://table.plazi.org/id/CBA0662CFFD6FFFBFF246AF02CA49198" pageId="11" pageNumber="412" tableDoi="http://doi.org/10.5281/zenodo.15224001" tableUuid="CBA0662CFFD6FFFBFF246AF02CA49198">Table 5</tableCitation>
).
</paragraph>
<paragraph id="9F6036A4FFD1FFFCFF0D6F212DD091E9" blockId="11.[159,775,197,1906]" lastBlockId="11.[823,1439,197,1170]" pageId="11" pageNumber="412">
@ -2077,7 +2079,7 @@ genotype at seven loci and an expected
<emphasis id="ADABEAB6FFD1FFFCFD5B6D0A2F67972A" italics="true" pageId="11" pageNumber="412">C. novaeguineae</emphasis>
</taxonomicName>
genotype at the remaining seven loci (
<tableCitation id="D25D031FFFD1FFFCFC8B6AF12DCB9080" box="[830,916,197,219]" captionStart="Table 5" captionStartId="12.[145,209,196,217]" captionTargetBox="[145,1423,489,1918]" captionTargetPageId="12" captionText="Table 5. Parental allele frequencies and hybrid genotypes at the 21 loci where the percentage difference between any two parental taxa is greater than 50%. Each of the two alleles displayed by a hybrid animal is indicated with a cross (+). Parental profiles for Clo and Cno have largely been taken from the overview study (Table 2; data for Hb from Stage 2) whereas those for Cru (combining the NT and Qld populations) and Cbu (NT populations only) were obtained from the Stage 2 analyses. None of the 24 loci not presented show genotypes inconsistent with a hybrid origin. An asterisk indicates that the presence of the null allele Acycf has been assumed, given the observed electrophoretic phenotype (the same mobility as allozyme e but a much-reduced activity). Superscripts are used to indicate which loci are diagnostic for the various parental taxa: AClo vs. Cno; BCno vs. Cru; CCru vs. Cbu (or effectively diagnostic if Cru is also represented by the NT populations only). (na = genotype was not determined)" pageId="11" pageNumber="412">Table 5</tableCitation>
<tableCitation id="D25D031FFFD1FFFCFC8B6AF12DCB9080" box="[830,916,197,219]" captionStart="Table 5" captionStartId="12.[145,209,196,217]" captionTargetBox="[145,1423,489,1918]" captionTargetPageId="12" captionText="Table 5. Parental allele frequencies and hybrid genotypes at the 21 loci where the percentage difference between any two parental taxa is greater than 50%. Each of the two alleles displayed by a hybrid animal is indicated with a cross (+). Parental profiles for Clo and Cno have largely been taken from the overview study (Table 2; data for Hb from Stage 2) whereas those for Cru (combining the NT and Qld populations) and Cbu (NT populations only) were obtained from the Stage 2 analyses. None of the 24 loci not presented show genotypes inconsistent with a hybrid origin. An asterisk indicates that the presence of the null allele Acycf has been assumed, given the observed electrophoretic phenotype (the same mobility as allozyme e but a much-reduced activity). Superscripts are used to indicate which loci are diagnostic for the various parental taxa: AClo vs. Cno; BCno vs. Cru; CCru vs. Cbu (or effectively diagnostic if Cru is also represented by the NT populations only). (na = genotype was not determined)" httpUri="http://table.plazi.org/id/CBA0662CFFD6FFFBFF246AF02CA49198" pageId="11" pageNumber="412" tableDoi="http://doi.org/10.5281/zenodo.15224001" tableUuid="CBA0662CFFD6FFFBFF246AF02CA49198">Table 5</tableCitation>
). The genotypes at the other 31 loci were also in line with the hypothesis that this individual has resulted from a genetic cross between an F
<subScript id="035B34E1FFD1FFFCFAD06B382B329141" attach="left" box="[1381,1389,268,282]" fontSize="6" pageId="11" pageNumber="412">1</subScript>
(
@ -2126,7 +2128,7 @@ and
, and the allozyme profile of this individual is consistent with it being an F
<subScript id="035B34E1FFD1FFFCFA8A68EC2B1892BD" attach="left" box="[1343,1351,728,742]" fontSize="6" pageId="11" pageNumber="412">1</subScript>
hybrid between the two species (
<tableCitation id="D25D031FFFD1FFFCFBED68D92AF29358" box="[1112,1197,749,771]" captionStart="Table 5" captionStartId="12.[145,209,196,217]" captionTargetBox="[145,1423,489,1918]" captionTargetPageId="12" captionText="Table 5. Parental allele frequencies and hybrid genotypes at the 21 loci where the percentage difference between any two parental taxa is greater than 50%. Each of the two alleles displayed by a hybrid animal is indicated with a cross (+). Parental profiles for Clo and Cno have largely been taken from the overview study (Table 2; data for Hb from Stage 2) whereas those for Cru (combining the NT and Qld populations) and Cbu (NT populations only) were obtained from the Stage 2 analyses. None of the 24 loci not presented show genotypes inconsistent with a hybrid origin. An asterisk indicates that the presence of the null allele Acycf has been assumed, given the observed electrophoretic phenotype (the same mobility as allozyme e but a much-reduced activity). Superscripts are used to indicate which loci are diagnostic for the various parental taxa: AClo vs. Cno; BCno vs. Cru; CCru vs. Cbu (or effectively diagnostic if Cru is also represented by the NT populations only). (na = genotype was not determined)" pageId="11" pageNumber="412">Table 5</tableCitation>
<tableCitation id="D25D031FFFD1FFFCFBED68D92AF29358" box="[1112,1197,749,771]" captionStart="Table 5" captionStartId="12.[145,209,196,217]" captionTargetBox="[145,1423,489,1918]" captionTargetPageId="12" captionText="Table 5. Parental allele frequencies and hybrid genotypes at the 21 loci where the percentage difference between any two parental taxa is greater than 50%. Each of the two alleles displayed by a hybrid animal is indicated with a cross (+). Parental profiles for Clo and Cno have largely been taken from the overview study (Table 2; data for Hb from Stage 2) whereas those for Cru (combining the NT and Qld populations) and Cbu (NT populations only) were obtained from the Stage 2 analyses. None of the 24 loci not presented show genotypes inconsistent with a hybrid origin. An asterisk indicates that the presence of the null allele Acycf has been assumed, given the observed electrophoretic phenotype (the same mobility as allozyme e but a much-reduced activity). Superscripts are used to indicate which loci are diagnostic for the various parental taxa: AClo vs. Cno; BCno vs. Cru; CCru vs. Cbu (or effectively diagnostic if Cru is also represented by the NT populations only). (na = genotype was not determined)" httpUri="http://table.plazi.org/id/CBA0662CFFD6FFFBFF246AF02CA49198" pageId="11" pageNumber="412" tableDoi="http://doi.org/10.5281/zenodo.15224001" tableUuid="CBA0662CFFD6FFFBFF246AF02CA49198">Table 5</tableCitation>
). The individual concerned displayed the heterozygous genotype expected for an F
<subScript id="035B34E1FFD1FFFCFC2269002DC09319" attach="left" box="[919,927,820,834]" fontSize="6" pageId="11" pageNumber="412">1</subScript>
at the diagnostic locus
@ -2145,7 +2147,7 @@ and
shows a neighbour-joining tree applied to the matrix of Rogers D distances (
<bibRefCitation id="FB4E4B55FFD1FFFCFB286F212B6D9570" author="Rogers JS" box="[1181,1330,1301,1323]" pageId="11" pageNumber="412" pagination="145 - 153" refId="ref16788" refString="Rogers JS. 1972. Measures of genetic similarity and genetic distance. Studies in Genetics VII. University of Texas Publication No. 7213: 145 - 153." type="book chapter" year="1972">Rogers, 1972</bibRefCitation>
) of
<tableCitation id="D25D031FFFD1FFFCFAD56F212D179512" captionStart="Table 4" captionStartId="10.[145,209,196,217]" captionTargetPageId="10" captionText="Table 4. Raw fixed differences (lower matrix) and Rogers D (upper matrix) for all forms of Chelodina that were diagnosable at one or more loci in our sample of populations. Species abbreviations are as for Table 1" pageId="11" pageNumber="412">Table 4</tableCitation>
<tableCitation id="D25D031FFFD1FFFCFAD56F212D179512" captionStart="Table 4" captionStartId="10.[145,209,196,217]" captionTargetPageId="10" captionText="Table 4. Raw fixed differences (lower matrix) and Rogers D (upper matrix) for all forms of Chelodina that were diagnosable at one or more loci in our sample of populations. Species abbreviations are as for Table 1" httpUri="http://table.plazi.org/id/CBA0662CFFD0FFFDFF246AF02AC790AD" pageId="11" pageNumber="412" tableDoi="http://doi.org/10.5281/zenodo.15223999" tableUuid="CBA0662CFFD0FFFDFF246AF02AC790AD">Table 4</tableCitation>
. It did not differ topologically from the maximum likelihood FitchMargoliash or in any substantial way from trees generated using percentage fixed differences as the distance measure. The tree is shown as an unrooted network because without an outgroup for this study, it was not possible to root the tree using data presented here. However, a number of previous molecular studies that have included
<taxonomicName id="58DF4D27FFD1FFFCFA9F6C3E2BC09644" box="[1322,1439,1546,1567]" class="Reptilia" family="Chelidae" genus="Chelodina" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="412" phylum="Chordata" rank="genus">
<emphasis id="ADABEAB6FFD1FFFCFA9F6C3E2BC09644" box="[1322,1439,1546,1567]" italics="true" pageId="11" pageNumber="412">Chelodina</emphasis>

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