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<mods:title id="523927285600CF906B340A7CED9AAD59">Uca (Petruca), a new subgenus for the rock fiddler crab Uca panamensis (Stimpson, 1859) from Central America, with comments on some species of the American broad-fronted subgenera</mods:title>
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<emphasis id="DEBCEAD6FFD970701EDCEB55FFA17307" bold="true" box="[151,200,295,321]" italics="true" pageId="15" pageNumber="486">Uca</emphasis>
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<emphasis id="DEBCEAD6FFD970701E92EB55FE5E7307" bold="true" box="[217,311,295,321]" italics="true" pageId="15" pageNumber="486">Petruca</emphasis>
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<emphasis id="DEBCEAD6FFD970701F03EB55FEBE7307" bold="true" box="[328,471,295,321]" italics="true" pageId="15" pageNumber="486">panamensis</emphasis>
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(
<figureCitation id="74F32A41FFD970701ED4EB38FE637325" box="[159,266,330,355]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURE 3" captionStart-3="FIGURE 4" captionStart-4="FIGURE 5" captionStartId-0="6.[151,250,1911,1933]" captionStartId-1="7.[151,250,1925,1947]" captionStartId-2="8.[151,250,1887,1909]" captionStartId-3="9.[151,250,1917,1939]" captionStartId-4="10.[151,250,1927,1949]" captionTargetBox-0="[163,1423,215,1877]" captionTargetBox-1="[159,1425,203,1891]" captionTargetBox-2="[161,1423,207,1857]" captionTargetBox-3="[159,1422,203,1887]" captionTargetBox-4="[159,1427,205,1895]" captionTargetId-0="figure@6.[151,1436,193,1890]" captionTargetId-1="figure@7.[151,1436,193,1903]" captionTargetId-2="figure@8.[151,1436,193,1866]" captionTargetId-3="figure@9.[151,1436,193,1896]" captionTargetId-4="figure@10.[151,1436,193,1906]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionTargetPageId-2="8" captionTargetPageId-3="9" captionTargetPageId-4="10" captionText-0="FIGURE 1. Uca panamensis (neotype, USNM 1294205, CW 16.8 mm). A, dorsal view; B, boss on carapace behind right dorsolateral margin (arrowed); C, ventral view. Scale, B = 2 mm." captionText-1="FIGURE 2. Uca panamensis (neotype, USNM 1294205, CW 16.8 mm). A, orbital region; B, inner side of orbital region, showing the tubercle of orbital armature (arrowed); C, frontal view; D, front; E, third maxillipeds. Scale, A, E = 2 mm; B = 1 mm; C = 10 mm; D = 5 mm." captionText-2="FIGURE 3. Major and minor chelipeds of Uca panamensis (neotype, USNM 1294205, CW 16.8 mm). A, B, outer and inner views of major cheliped; C, D, outer and inner views of minor cheliped. Scale, C, D = 2 mm." captionText-3="FIGURE 4. Uca panamensis, right G 1 (neotype, USNM 1294205, CW 16.8 mm). A, dorsal view; B, dorsal view of terminal segment; C, D, ventral view of terminal segment. Scale, A = 2 mm; B, C = 0.5 mm." captionText-4="FIGURE 5. Urocardiac ossicle of Uca panamensis. A, B, neotype, USNM 1294205 (16.8 mm); C, ZRC (CW 12.6 mm, ♂); D, ZRC (CW 13.7 mm, ♀). A, C, D, ventral view; B, lateroventral view. Scale, A, B = 1 mm; C, D = 0.5 mm." httpUri-0="https://zenodo.org/record/235574/files/figure.png" httpUri-1="https://zenodo.org/record/235575/files/figure.png" httpUri-2="https://zenodo.org/record/235576/files/figure.png" httpUri-3="https://zenodo.org/record/235577/files/figure.png" httpUri-4="https://zenodo.org/record/235578/files/figure.png" pageId="15" pageNumber="486">Figs. 15</figureCitation>
,
<figureCitation id="74F32A41FFD970701F5DEB39FE4D7325" box="[278,292,331,355]" captionStart="FIGURE 8" captionStartId="13.[151,250,1719,1741]" captionTargetBox="[163,1425,205,1689]" captionTargetId="figure@13.[151,1436,193,1698]" captionTargetPageId="13" captionText="FIGURE 8. Uca panamensis ♂ (A, B) and its habitat (C). Photographs taken in Panamá. B, courtesy of P. Backwell." httpUri="https://zenodo.org/record/235581/files/figure.png" pageId="15" pageNumber="486">8</figureCitation>
)
</paragraph>
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<emphasis id="DEBCEAD6FFD970701EDCEBE0FEE973EE" box="[151,384,402,424]" italics="true" pageId="15" pageNumber="486">Gelasimus panamensis</emphasis>
<treatmentCitation id="6D6910D5FFD970701FCDEBE0FD2773EE" author="Stimpson" box="[390,590,402,424]" page="63" pageId="15" pageNumber="486" year="1859">
<bibRefCitation id="88594B35FFD970701FCDEBE0FD4173EE" author="Stimpson" box="[390,552,402,424]" pageId="15" pageNumber="486" refString="Stimpson, W. (1859) Notes on North American Crustacea, No. 1. Annals of the Lyceum of Natural History of New York, 7 (4 - 5), 49 - 93, pl. I." type="journal article" year="1859">Stimpson, 1859</bibRefCitation>
: 63
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;
<treatmentCitation id="6D6910D5FFD970701C13EBE0FC6A73EE" author="Smith" box="[600,771,402,424]" page="139" pageId="15" pageNumber="486" year="1870">
<bibRefCitation id="88594B35FFD970701C13EBE0FDB873EE" author="Smith" box="[600,721,402,424]" pageId="15" pageNumber="486" refString="Smith, S. I. (1870) Notes on American Crustacea. No. 1. Ocypodoidea. Transactions of the Connecticut Academy of Arts and Science, 2, 113 - 176, pls. 2 - 5." type="journal article" year="1870">Smith 1870</bibRefCitation>
: 139
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, pl. 4(5);
<treatmentCitation id="6D6910D5FFD970701D2DEBE0FB4673EE" author="Kingsley" box="[870,1071,402,424]" page="150" pageId="15" pageNumber="486" year="1880">
<bibRefCitation id="88594B35FFD970701D2DEBE0FC9473EE" author="Kingsley" box="[870,1021,402,424]" pageId="15" pageNumber="486" refString="Kingsley, J. S. (1880) Carcinological notes, no. II. - Revision of the Gelasimi. Proceedings of the Academy of Natural Sciences of Philadelphia, 1880, 135 - 155." type="proceedings paper" year="1880">Kingsley 1880</bibRefCitation>
: 150
</treatmentCitation>
, fig. 10(24);
<treatmentCitation id="6D6910D5FFD970701AF8EBE0FA3173EE" author="Cano" box="[1203,1368,402,424]" page="235" pageId="15" pageNumber="486" year="1889">
<bibRefCitation id="88594B35FFD970701AF8EBE0FA4C73EE" author="Cano" box="[1203,1317,402,424]" pageId="15" pageNumber="486" refString="Cano, G. (1889) Crostacei brachiuri ed anomuri raccolti nel viaggio della &quot; Vettor Pisani &quot; intorno al globo. Bollettino della Societa di Naturalisti in Napoli, 3, 79 - 106, 169 - 268." type="journal article" year="1889">Cano 1889</bibRefCitation>
: 235
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.
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</paragraph>
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<treatmentCitationGroup id="CCD811EAFFD970701EDCEBC0FED073A0" pageId="15" pageNumber="486">
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<emphasis id="DEBCEAD6FFD970701EDCEBC0FE287381" box="[151,321,434,455]" italics="true" pageId="15" pageNumber="486">Uca panamensis</emphasis>
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<treatmentCitation id="6D6910D5FFD970701F20EBC3FD7B7381" author="Nobili" box="[363,530,433,455]" page="49" pageId="15" pageNumber="486" year="1901">
<bibRefCitation id="88594B35FFD970701F20EBC3FE837381" author="Nobili" box="[363,490,433,455]" pageId="15" pageNumber="486" refString="Nobili, G. (1901) Viaggio del Dr. Enrico Festa nella Republica dell'Ecuador e regione vicine. Decapodi e Stomatopodi. Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Universit di Torino, 16 (415), 1 - 58." type="journal article" year="1901">Nobili 1901</bibRefCitation>
: 49
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;
<treatmentCitation id="6D6910D5FFD970701C55EBC3FD8F7381" author="Rathbun" box="[542,742,433,455]" page="412" pageId="15" pageNumber="486" year="1918">
<bibRefCitation id="88594B35FFD970701C55EBC3FDDB7381" author="Rathbun" box="[542,690,433,455]" pageId="15" pageNumber="486" refString="Rathbun, M. J. (1918) The grapsoid crabs of America. Bulletin of the United States National Museum, 97, 1 - 461, pls. 1 - 161. http: // dx. doi. org / 10.5479 / si. 03629236.97. i" type="journal article" year="1918">Rathbun 1918</bibRefCitation>
: 412
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, pl. 149;
<treatmentCitation id="6D6910D5FFD970701D00EBC3FC937381" author="Crane" box="[843,1018,433,455]" page="204" pageId="15" pageNumber="486" year="1941">
<bibRefCitation id="88594B35FFD970701D00EBC3FCAC7381" author="Crane" box="[843,965,433,455]" pageId="15" pageNumber="486" refString="Crane, J. (1941) Crabs of the genus Uca from the west coast of Central America. Eastern Pacific Expeditions of the New York Zoological Society. XXVI. Zoologica, 26 (19), 145 - 208." type="journal article" year="1941">Crane 1941</bibRefCitation>
: 204
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, test-fig.
<date id="98761004FFD970701A17EBC3FB177381" box="[1116,1150,433,455]" pageId="15" pageNumber="486">4X</date>
;
<treatmentCitation id="6D6910D5FFD970701AC1EBC3FA437381" author="Garth" box="[1162,1322,433,455]" page="60" pageId="15" pageNumber="486" year="1948">
<bibRefCitation id="88594B35FFD970701AC1EBC3FA6B7381" author="Garth" box="[1162,1282,433,455]" pageId="15" pageNumber="486" refString="Garth, J. S. (1948) The Brachyura of the &quot; Askoy &quot; Expedition, with remarks on carcinological collecting in the Panama Bight. Bulletin of the American Museum of Natural History, 92, 1 - 66." type="journal article" year="1948">Garth 1948</bibRefCitation>
: 60
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;
<treatmentCitation id="6D6910D5FFD970701B7CEBC3FE1A73A0" author="Takeda" page="179" pageId="15" pageNumber="486" year="2003">
<bibRefCitation id="88594B35FFD970701B7CEBC3FE2873A0" author="Takeda" pageId="15" pageNumber="486" refString="Takeda, S. &amp; Murai, M. (2003) Morphological and behavioural adaptations to the rocky substrate by the fiddler crab Uca panamensis (Stimpson, 1859): preference for feeding substratum and feeding mechanism. Journal of Experimental Marine Biology and Ecology, 287, 179 - 191. http: // dx. doi. org / 10.1016 / S 0022 - 0981 (02) 00550 - 6" type="journal article" year="2003">Takeda &amp; Murai 2003</bibRefCitation>
: 179
</treatmentCitation>
, fig. 1.
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<taxonomicName id="2BC84D47FFD970701EDCEB82FECE7043" ID-CoL="7Z6P4" box="[151,423,494,517]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="panamensis" subGenus="Minuca">
<emphasis id="DEBCEAD6FFD970701EDCEB82FFA87043" box="[151,193,496,517]" italics="true" pageId="15" pageNumber="486">Uca</emphasis>
(
<emphasis id="DEBCEAD6FFD970701E99EB82FE767043" box="[210,287,496,517]" italics="true" pageId="15" pageNumber="486">Minuca</emphasis>
)
<emphasis id="DEBCEAD6FFD970701F7BEB82FECE7043" box="[304,423,496,517]" italics="true" pageId="15" pageNumber="486">panamensis</emphasis>
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<treatmentCitation id="6D6910D5FFD970701F99EB9DFD197042" author="Bott" box="[466,624,494,516]" page="162" pageId="15" pageNumber="486" year="1954">
<bibRefCitation id="88594B35FFD970701F99EB9DFD557042" author="Bott" box="[466,572,494,516]" pageId="15" pageNumber="486" refString="Bott, R. (1954) Dekapoden (Crustacea) aus El Salvador. 1. Winkerkrabben (Uca). Senckenbergiana Biologica, 35 (3 / 4), 155 - 180." type="journal article" year="1954">Bott 1954</bibRefCitation>
: 162
</treatmentCitation>
, fig. 4, pl. 15(4);
<treatmentCitation id="6D6910D5FFD970701D66EB9CFCB57042" author="Crane" box="[813,988,494,516]" page="158" pageId="15" pageNumber="486" year="1975">
<bibRefCitation id="88594B35FFD970701D66EB9CFCCE7042" author="Crane" box="[813,935,494,516]" pageId="15" pageNumber="486" refString="Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp." type="book" year="1975">Crane 1975</bibRefCitation>
: 158
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, pl. 22AD, figs. 46H, 66G;
<treatmentCitation id="6D6910D5FFD970701B44EB9CFE467065" author="Prahl" page="147" pageId="15" pageNumber="486" year="1979">
<bibRefCitation id="88594B35FFD970701B44EB9CFF947065" author="Prahl" pageId="15" pageNumber="486" refString="Prahl, H. von &amp; Guhl, F. (1979) Notas sobre la biologia de Uca (Minuca) panamensis (Stimpson, 1859) (Crustacea, Ocipodidae) y su distribucion en Colombia. Anales del Instituto de Investigaciones Marinas de Punta Betin, 11, 147 - 152." type="journal article" year="1979">Prahl &amp; Guhl 1979</bibRefCitation>
: 147
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, figs. 12.
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</paragraph>
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<emphasis id="DEBCEAD6FFD970701EDCE85FFFA87004" box="[151,193,557,578]" italics="true" pageId="15" pageNumber="486">Uca</emphasis>
(
<emphasis id="DEBCEAD6FFD970701E9BE85FFE4B7004" box="[208,290,557,578]" italics="true" pageId="15" pageNumber="486">Leptuca</emphasis>
)
<emphasis id="DEBCEAD6FFD970701F7BE85FFEC17004" box="[304,424,557,578]" italics="true" pageId="15" pageNumber="486">panamensis</emphasis>
</taxonomicName>
<treatmentCitation id="6D6910D5FFD970701F86E85EFDCE7004" author="Rosenberg" box="[461,679,556,578]" page="848" pageId="15" pageNumber="486" year="2001">
<bibRefCitation id="88594B35FFD970701F86E85EFD1C7004" author="Rosenberg" box="[461,629,556,578]" pageId="15" pageNumber="486" refString="Rosenberg, M. S. (2001) The systematics and taxonomy of fiddler crabs: a phylogeny of the genus Uca. Journal of Crustacean Biology, 21, 839 - 869. http: // dx. doi. org / 10.1163 / 20021975 - 99990176" type="journal article" year="2001">Rosenberg 2001</bibRefCitation>
: 848
</treatmentCitation>
, 852; Beinlich &amp; von
<treatmentCitation id="6D6910D5FFD970701DC3E85FFB457004" author="Hagen" box="[904,1068,556,578]" page="22" pageId="15" pageNumber="486" year="2006">
<bibRefCitation id="88594B35FFD970701DC3E85FFB6E7004" author="Hagen" box="[904,1031,556,578]" pageId="15" pageNumber="486" refString="Beinlich, B. &amp; von Hagen, H. O. (2006) Materials for a more stable subdivision of the genus Uca Leach. Zoologische Mededelingen, 80, 9 - 32." type="journal article" year="2006">Hagen 2006</bibRefCitation>
: 22
</treatmentCitation>
, 26, fig. 7j; Ng
<emphasis id="DEBCEAD6FFD970701A80E85FFB967004" box="[1227,1279,556,578]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2008: 241.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="A4D2654FFFD970701EDCE81CFDFE717D" pageId="15" pageNumber="486" type="materials_examined">
<paragraph id="EC7736C4FFD970701EDCE81CFC837151" blockId="15.[151,1437,622,2018]" pageId="15" pageNumber="486">
<emphasis id="DEBCEAD6FFD970701EDCE81CFEEC70C1" bold="true" box="[151,389,622,647]" pageId="15" pageNumber="486">Material examined.</emphasis>
<typeStatus id="33738866FFD970701FDAE81DFE9970C1" box="[401,496,623,647]" pageId="15" pageNumber="486" type="neotype">Neotype</typeStatus>
for
<taxonomicName id="2BC84D47FFD970701C62E81CFC4670C0" box="[553,815,622,647]" class="Malacostraca" family="Ocypodidae" genus="Gelasimus" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="panamensis">
<emphasis id="DEBCEAD6FFD970701C62E81CFC4670C0" box="[553,815,622,647]" italics="true" pageId="15" pageNumber="486">Gelasimus panamensis</emphasis>
</taxonomicName>
,
<specimenCount id="FACEFD4DFFD970701D09E81DFC1870C0" box="[834,881,623,647]" pageId="15" pageNumber="486" type="generic">1 ♂</specimenCount>
(
<quantity id="2B309B21FFD970701DCDE81CFC9D70C1" box="[902,1012,622,647]" metricMagnitude="-2" metricUnit="m" metricValue="1.68" pageId="15" pageNumber="486" unit="mm" value="16.8">16.8 mm</quantity>
),
<collectionCode id="8AD9AE01FFD970701A47E81DFB3470C1" LSID="urn:lsid:biocol.org:col:34871" box="[1036,1117,623,647]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="15" pageNumber="486">USNM</collectionCode>
1294205 (transferred from NCHUZOOL 13581), Culebra Island,
<collectingCountry id="94DF7654FFD970701C06E8E1FDC070ED" box="[589,681,659,683]" name="Panama" pageId="15" pageNumber="486">Panamá</collectingCountry>
, coll. J. Christy, 2008;
<specimenCount id="FACEFD4DFFD970701DFBE8E1FCB570EC" box="[944,988,659,683]" pageId="15" pageNumber="486" type="generic">1 ♂</specimenCount>
(13.0 mm), NCHUZOOL 13581, same data as
<typeStatus id="33738866FFD970701EA0E8C5FE217089" box="[235,328,695,719]" pageId="15" pageNumber="486" type="neotype">neotype</typeStatus>
;
<specimenCount id="FACEFD4DFFD970701F18E8C5FE147088" box="[339,381,695,719]" pageId="15" pageNumber="486" type="generic">1 ♂</specimenCount>
(
<quantity id="2B309B21FFD970701FC6E8C4FE9F7089" box="[397,502,694,719]" metricMagnitude="-2" metricUnit="m" metricValue="1.26" pageId="15" pageNumber="486" unit="mm" value="12.6">12.6 mm</quantity>
),
<collectionCode id="8AD9AE01FFD970701C42E8C5FD567089" box="[521,575,695,719]" httpUri="http://grbio.org/cool/16z4-1tbx" name="Zoological Reference Collection, National University of Singapore" pageId="15" pageNumber="486">ZRC</collectionCode>
2012.0126, Playa San Juanillo,
<collectingCountry id="94DF7654FFD970701DE1E8C5FB4E7088" box="[938,1063,694,719]" name="Costa Rica" pageId="15" pageNumber="486">Costa Rica</collectingCountry>
, coll. Z. Jaafar,
<date id="98761004FFD970701A91E8C4FAFE7089" box="[1242,1431,694,719]" pageId="15" pageNumber="486" value="2011-07-25" valueMax="2011-07-27" valueMin="2011-07-25">2527 July 2011</date>
; 4 ♂♂ (
<quantity id="2B309B21FFD970701EA2E8A8FEFB70B5" box="[233,402,730,755]" metricMagnitude="-2" metricUnit="m" metricValue="1.31" metricValueMax="1.51" metricValueMin="1.1099999999999999" pageId="15" pageNumber="486" unit="mm" value="13.1" valueMax="15.1" valueMin="11.1">11.115.1 mm</quantity>
), 2 ♀♀ (11.7,
<quantity id="2B309B21FFD970701C71E8A9FDCD70B5" box="[570,676,731,755]" metricMagnitude="-2" metricUnit="m" metricValue="1.3699999999999999" pageId="15" pageNumber="486" unit="mm" value="13.7">13.7 mm</quantity>
),
<collectionCode id="8AD9AE01FFD970701CF2E8A9FD9B70B5" box="[697,754,731,755]" httpUri="http://grbio.org/cool/16z4-1tbx" name="Zoological Reference Collection, National University of Singapore" pageId="15" pageNumber="486">ZRC</collectionCode>
,
<specimenCount id="FACEFD4DFFD970701CB4E8A9FC4270B4" box="[767,811,731,755]" pageId="15" pageNumber="486" type="generic">1 ♂</specimenCount>
(
<quantity id="2B309B21FFD970701D77E8A8FCC170B5" box="[828,936,730,755]" metricMagnitude="-2" metricUnit="m" metricValue="1.64" pageId="15" pageNumber="486" unit="mm" value="16.4">16.4 mm</quantity>
), 1 ♀ (
<quantity id="2B309B21FFD970701DB2E8A8FB0D70B5" box="[1017,1124,730,755]" metricMagnitude="-2" metricUnit="m" metricValue="1.68" pageId="15" pageNumber="486" unit="mm" value="16.8">16.8 mm</quantity>
), NCHUZOOL 14753, San Juanillo rocky shore, Ostional,
<collectingCountry id="94DF7654FFD970701FBEE88DFD1B7151" box="[501,626,766,791]" name="Costa Rica" pageId="15" pageNumber="486">Costa Rica</collectingCountry>
, coll. Z. Jaafar,
<date id="98761004FFD970701D6FE88CFCC87151" box="[804,929,766,791]" pageId="15" pageNumber="486" value="1926-07-29">2629 July</date>
, 2011.
</paragraph>
<paragraph id="EC7736C4FFD970701E8CE950FDFE717D" blockId="15.[151,1437,622,2018]" box="[199,663,802,827]" pageId="15" pageNumber="486">
<emphasis id="DEBCEAD6FFD970701E8CE950FEBE717D" bold="true" box="[199,471,802,827]" pageId="15" pageNumber="486">Comparative material.</emphasis>
See Appendix 1.
</paragraph>
</subSubSection>
<subSubSection id="A4D2654FFFD9706D1E8CE934FF8A717E" lastPageId="18" lastPageNumber="489" pageId="15" pageNumber="486" type="discussion">
<paragraph id="EC7736C4FFD970701E8CE934FD927639" blockId="15.[151,1437,622,2018]" pageId="15" pageNumber="486">
<emphasis id="DEBCEAD6FFD970701E8CE934FE527119" bold="true" box="[199,315,838,863]" pageId="15" pageNumber="486">Remarks.</emphasis>
<taxonomicName id="2BC84D47FFD970701F0EE934FD237118" box="[325,586,838,863]" class="Malacostraca" family="Ocypodidae" genus="Gelasimus" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="panamensis">
<emphasis id="DEBCEAD6FFD970701F0EE934FD237118" box="[325,586,838,863]" italics="true" pageId="15" pageNumber="486">Gelasimus panamensis</emphasis>
</taxonomicName>
was described by
<bibRefCitation id="88594B35FFD970701D61E934FC807119" author="Stimpson" box="[810,1001,838,863]" pageId="15" pageNumber="486" refString="Stimpson, W. (1859) Notes on North American Crustacea, No. 1. Annals of the Lyceum of Natural History of New York, 7 (4 - 5), 49 - 93, pl. I." type="journal article" year="1859">Stimpson (1859)</bibRefCitation>
from three specimens, all apparently immature, from an undisclosed site in
<collectingCountry id="94DF7654FFD970701C13E919FDDD71C5" box="[600,692,875,899]" name="Panama" pageId="15" pageNumber="486">Panamá</collectingCountry>
. No figures were provided. In any case, the
<typeStatus id="33738866FFD970701AF6E919FB9171C5" box="[1213,1272,875,899]" pageId="15" pageNumber="486">types</typeStatus>
are no longer extant. Like most of his material, they were almost certainly lost in the great Chicago Fire (see
<bibRefCitation id="88594B35FFD970701A81E9FDFA3871E1" author="Evans" box="[1226,1361,910,935]" pageId="15" pageNumber="486" refString="Evans, A. C. (1967) Syntypes of Decapoda described by William Stimpson and James Dana in the collections of the British Museum (Natural History). Journal of Natural History, 1, 399 - 411. http: // dx. doi. org / 10.1080 / 00222936700770391" type="journal article" year="1967">Evans 1967</bibRefCitation>
;
<bibRefCitation id="88594B35FFD970701B15E9FCFE35718D" author="Deiss" pageId="15" pageNumber="486" refString="Deiss, W. A. &amp; Manning, R. B. (1981) The fate of the invertebrate collections of the North Pacific Exploring Expedition, 1853 - 1856. In: Wheeler, A. &amp; Price, J. H. (Eds), History in the Service of Systematics. Society for the Bibliography of Natural History, London, pp. 79 - 85." type="book chapter" year="1981">Deiss &amp; Manning 1981</bibRefCitation>
;
<bibRefCitation id="88594B35FFD970701F22E9C0FD78718D" author="Manning" box="[361,529,946,971]" pageId="15" pageNumber="486" refString="Manning, R. B. (1993) The scientific contributions of William Stimpson, an early American naturalist and taxonomist. Crustacean Issues, 8, 109 - 117." type="journal article" year="1993">Manning 1993</bibRefCitation>
;
<bibRefCitation id="88594B35FFD970701C55E9C0FD8C718D" author="Vasile" box="[542,741,946,971]" pageId="15" pageNumber="486" refString="Vasile, R. S., Manning, R. B. &amp; Lemaitre, R. (2005) William Stimpson's Journal from the North Pacific Exploring Expedition, 1853 - 1856. Crustacean Research Special, 5, 1 - 220." type="journal article" year="2005">
Vasile
<emphasis id="DEBCEAD6FFD970701C21E9C6FDCA718D" box="[618,675,946,971]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2005
</bibRefCitation>
;
<bibRefCitation id="88594B35FFD970701CB9E9C0FC92718D" author="Manning" box="[754,1019,946,971]" pageId="15" pageNumber="486" refString="Manning, R. B. &amp; Reed, K. J. (2006) Decapod crustaceans deposited in the Zoological Museum of Copenhagen by William Stimpson in 1859. Raffles Bulletin of Zoology, 54, 283 - 293." type="journal article" year="2006">Manning &amp; Reed 2006</bibRefCitation>
). Although the species has been well treated by many authors, notably
<bibRefCitation id="88594B35FFD970701C6FE9A5FDD671A9" author="Crane" box="[548,703,982,1007]" pageId="15" pageNumber="486" refString="Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp." type="book" year="1975">Crane (1975)</bibRefCitation>
, since a new subgenus is established here for the species, the designation of a
<typeStatus id="33738866FFD970701F2AE989FED37655" box="[353,442,1019,1043]" pageId="15" pageNumber="486" type="neotype">neotype</typeStatus>
for
<taxonomicName id="2BC84D47FFD970701FB9E988FCD47655" authority="Stimpson, 1859" authorityName="Stimpson" authorityYear="1859" box="[498,957,1018,1043]" class="Malacostraca" family="Ocypodidae" genus="Gelasimus" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="panamensis">
<emphasis id="DEBCEAD6FFD970701FB9E988FD907655" box="[498,761,1018,1043]" italics="true" pageId="15" pageNumber="486">Gelasimus panamensis</emphasis>
<bibRefCitation id="88594B35FFD970701D4EE988FCD47655" author="Stimpson" box="[773,957,1018,1043]" pageId="15" pageNumber="486" refString="Stimpson, W. (1859) Notes on North American Crustacea, No. 1. Annals of the Lyceum of Natural History of New York, 7 (4 - 5), 49 - 93, pl. I." type="journal article" year="1859">Stimpson, 1859</bibRefCitation>
</taxonomicName>
, is important for longer term taxonomic stability. This is especially important given that there is now active research on
<taxonomicName id="2BC84D47FFD970701A50EE6DFB237670" box="[1051,1098,1055,1078]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFD970701A50EE6DFB237670" box="[1051,1098,1055,1078]" italics="true" pageId="15" pageNumber="486">Uca</emphasis>
</taxonomicName>
. To this effect, we here select an adult male with CW of
<quantity id="2B309B21FFD970701F8FEE30FD40761C" box="[452,553,1090,1115]" metricMagnitude="-2" metricUnit="m" metricValue="1.68" pageId="15" pageNumber="486" unit="mm" value="16.8">16.8 mm</quantity>
collected from
<collectingCountry id="94DF7654FFD970701C90EE31FC5D761D" box="[731,820,1091,1115]" name="Panama" pageId="15" pageNumber="486">Panamá</collectingCountry>
as the
<typeStatus id="33738866FFD970701DCEEE31FCB7761D" box="[901,990,1091,1115]" pageId="15" pageNumber="486" type="neotype">neotype</typeStatus>
of the species. It is a recently collected specimen (USNM 1294205) that also has genetic data.
</paragraph>
<paragraph id="EC7736C4FFD970701E8CEEF8FAE87770" blockId="15.[151,1437,622,2018]" pageId="15" pageNumber="486">
<emphasis id="DEBCEAD6FFD970701E8CEEF8FD4276E5" bold="true" box="[199,555,1162,1187]" pageId="15" pageNumber="486">DNA analyses and discussion.</emphasis>
A 589 bp segment of the 16S, 658 bp segment of COI and 689 bp segment of 28S, from 55 species of fiddler crabs, were amplified and aligned (
<tableCitation id="A14A037FFFD970701DCBEEDCFCBA7681" box="[896,979,1198,1223]" captionStart="TABLE 1" captionStartId="2.[151,232,157,178]" captionText="TABLE 1. Haplotypes of 16 S rDNA, COI and 28 S rDNA for the subgenus Petruca subgen. nov. and other related subgenera used in this study. See “ Material and methods ” for abbreviations of institutions. *, see “ DNA analyses and discussion ”." pageId="15" pageNumber="486">Table 1</tableCitation>
). The phylogenetic tree of the combined markers was reconstructed from the
<collectingCountry id="94DF7654FFD970701C7DEEA1FD3B76AC" box="[566,594,1235,1258]" name="Belgium" pageId="15" pageNumber="486">BI</collectingCountry>
analysis, with the support values from two methods of ML analyses (
<figureCitation id="74F32A41FFD970701B24EEA0FFCE7754" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="15" pageNumber="486">Fig. 9</figureCitation>
). To estimate the divergence time, the substitution rate of 1.17% /10 6 year for the combined 16S and COI of marine sesarmids (cf.
<bibRefCitation id="88594B35FFD970701FC5EF6FFD1A7770" author="Schubart" box="[398,627,1309,1334]" pageId="15" pageNumber="486" refString="Schubart, C. D., Diesel, R. &amp; Hedges, S. B. (1998) Rapid evolution to terrestrial life in Jamaican crabs. Nature, 393, 363 - 365. http: // dx. doi. org / 10.1038 / 30724" type="journal article" year="1998">
Schubart
<emphasis id="DEBCEAD6FFD970701FB2EF6CFD587770" box="[505,561,1309,1334]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
1998
</bibRefCitation>
;
<bibRefCitation id="88594B35FFD970701C35EF6FFC1C7770" author="Ragionieri" box="[638,885,1309,1334]" pageId="15" pageNumber="486" refString="Ragionieri, L., Fratini, S, Vannini, M. &amp; Schubart, C. D. (2009) Phylogenetic and morphometric differentiation reveal geographic radiation and pseudo-cryptic speciation in a mangrove crab from the Indo-West Pacific. Molecular Phylogenetics and Evolution, 52, 825 - 834." type="journal article" year="2009">
Ragionieri
<emphasis id="DEBCEAD6FFD970701CB7EF6CFC5D7770" box="[764,820,1309,1334]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2009
</bibRefCitation>
) was applied to the p-distance between clades.
</paragraph>
<paragraph id="EC7736C4FFD970701E8CEF33FA7477E4" blockId="15.[151,1437,622,2018]" pageId="15" pageNumber="486">
The relationships of the IWP subgenera (
<figureCitation id="74F32A41FFD970701CDCEF33FDB2771C" box="[663,731,1345,1370]" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="15" pageNumber="486">Fig. 9</figureCitation>
) generally agree with
<bibRefCitation id="88594B35FFD970701D92EF33FBC7771C" box="[985,1198,1345,1370]" pageId="15" pageNumber="486" refString="Shih, H. - T., Ng, P. K. L. &amp; Liu, M. - Y. (2013 b) Systematics of the Indo-West Pacific broad-fronted fiddler crabs (Crustacea: Ocypodidae: genus Uca). Raffles Bulletin of Zoology, 61, 641 - 649." type="journal article">
Shih
<emphasis id="DEBCEAD6FFD970701A5FEF30FB27771C" box="[1044,1102,1345,1370]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
(2013b)
</bibRefCitation>
and Shih (2015) and are not repeated here. The American broad-fronted species form a highly-supported main clade by three methods, with three distinct clades corresponding to three subgenera with high support at least by the
<collectingCountry id="94DF7654FFD970701AD4EFF8FBD277E7" box="[1183,1211,1418,1441]" name="Belgium" pageId="15" pageNumber="486">BI</collectingCountry>
method.
</paragraph>
<paragraph id="EC7736C4FFD970701E8CEFDFFDF17448" blockId="15.[151,1437,622,2018]" pageId="15" pageNumber="486">
The subgenus
<emphasis id="DEBCEAD6FFD970701F27EFDCFEAC7783" box="[364,453,1454,1477]" italics="true" pageId="15" pageNumber="486">Petruca</emphasis>
<emphasis id="DEBCEAD6FFD970701F85EFDFFD0A7780" bold="true" box="[462,611,1453,1478]" pageId="15" pageNumber="486">subgen. nov.</emphasis>
diverged earlier than the others at about 14 million years ago (mya) (pdistance of the combined 16S and COI = 16.45%). Both
<taxonomicName id="2BC84D47FFD970701D59EFA0FC0177AF" box="[786,872,1490,1513]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFD970701D59EFA0FC0177AF" box="[786,872,1490,1513]" italics="true" pageId="15" pageNumber="486">Minuca</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFD970701DD5EFA0FC9177AF" box="[926,1016,1490,1513]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFD970701DD5EFA0FC9177AF" box="[926,1016,1490,1513]" italics="true" pageId="15" pageNumber="486">Leptuca</emphasis>
</taxonomicName>
are monophyletic, are closely related and diverged at 9 mya (p-distance = 10.63%).
</paragraph>
<paragraph id="EC7736C4FFD970701E8CEC6BFA2C7514" blockId="15.[151,1437,622,2018]" pageId="15" pageNumber="486">
The six specimens of
<taxonomicName id="2BC84D47FFD970701F98EC68FD167477" box="[467,639,1562,1585]" class="Malacostraca" family="Ocypodidae" genus="Gelasimus" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="panamensis">
<emphasis id="DEBCEAD6FFD970701F98EC68FD167477" box="[467,639,1562,1585]" italics="true" pageId="15" pageNumber="486">U. panamensis</emphasis>
</taxonomicName>
from the Pacific coasts of
<collectingCountry id="94DF7654FFD970701D84EC68FB267474" box="[975,1103,1561,1586]" name="Costa Rica" pageId="15" pageNumber="486">Costa Rica</collectingCountry>
and
<collectingCountry id="94DF7654FFD970701AD9EC68FB827474" box="[1170,1259,1562,1586]" name="Panama" pageId="15" pageNumber="486">Panamá</collectingCountry>
form a highly supported clade supporting the establishment of
<emphasis id="DEBCEAD6FFD970701C87EC4CFC4C7413" box="[716,805,1598,1621]" italics="true" pageId="15" pageNumber="486">Petruca</emphasis>
<emphasis id="DEBCEAD6FFD970701D64EC4FFCAF7410" bold="true" box="[815,966,1597,1622]" pageId="15" pageNumber="486">subgen. nov.</emphasis>
Its intraspecific K2P divergence COI is average 1.73% (0.332.81%), which is lower than
<taxonomicName id="2BC84D47FFD970701C84EC10FC37743C" box="[719,862,1633,1658]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="splendida">
<emphasis id="DEBCEAD6FFD970701C84EC10FC37743C" box="[719,862,1633,1658]" italics="true" pageId="15" pageNumber="486">U. splendida</emphasis>
</taxonomicName>
(1.86%; cf. Shih
<emphasis id="DEBCEAD6FFD970701A6FEC10FB35743C" box="[1060,1116,1633,1658]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2012), but higher than other reported fiddler crabs. For example, the largest intraspecific divergences are 1.23% in
<taxonomicName id="2BC84D47FFD970701A38ECF4FA6B74D8" box="[1139,1282,1669,1694]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="jocelynae">
<emphasis id="DEBCEAD6FFD970701A38ECF4FA6B74D8" box="[1139,1282,1669,1694]" italics="true" pageId="15" pageNumber="486">U. jocelynae</emphasis>
</taxonomicName>
(recalculated from
<bibRefCitation id="88594B35FFD970701E9CECDBFEFC7484" author="Shih" box="[215,405,1705,1730]" pageId="15" pageNumber="486" refString="Shih, H. - T., Naruse, T. &amp; Ng, P. K. L. (2010) Uca jocelynae sp. nov., a new species of fiddler crab (Crustacea: Brachyura: Ocypodidae) from the Western Pacific. Zootaxa, 2337, 47 - 62." type="journal article" year="2010">
Shih
<emphasis id="DEBCEAD6FFD970701F5FECD8FE267484" box="[276,335,1705,1730]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2010
</bibRefCitation>
); 0.98% in
<taxonomicName id="2BC84D47FFD970701C57ECD8FCEE7484" authority="Shih et al. 2009" authorityName="Shih et al." authorityYear="2009" box="[540,903,1705,1730]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="annulipes">
<emphasis id="DEBCEAD6FFD970701C57ECD8FDC77484" box="[540,686,1705,1730]" italics="true" pageId="15" pageNumber="486">U. annulipes</emphasis>
(
<bibRefCitation id="88594B35FFD970701CF4ECDBFC167484" author="Shih" box="[703,895,1705,1730]" pageId="15" pageNumber="486" refString="Shih, H. - T., Kamrani, E., Davie, P. J. F. &amp; Liu, M. - Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U." type="book" year="2009">
Shih
<emphasis id="DEBCEAD6FFD970701CB5ECD8FC507484" box="[766,825,1705,1730]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2009
</bibRefCitation>
)
</taxonomicName>
; about 0.83% in
<taxonomicName id="2BC84D47FFD970701A05ECD8FBAF7487" box="[1102,1222,1706,1729]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="pugnax">
<emphasis id="DEBCEAD6FFD970701A05ECD8FBAF7487" box="[1102,1222,1706,1729]" italics="true" pageId="15" pageNumber="486">U. pugnax</emphasis>
</taxonomicName>
(cf.
<bibRefCitation id="88594B35FFD970701AB4ECDBFFBD74A0" author="Sanford" pageId="15" pageNumber="486" refString="Sanford, E., Holzman, S. B., Haney, R. A., Rand, D. M. &amp; Bertness, M. D. (2006) Larval tolerance, gene flow, and the northern geographic range limit of fiddler crabs. Ecology, 87, 2882 - 2894. http: // dx. doi. org / 10.1890 / 0012 - 9658 (2006) 87 [2882: LTGFAT] 2.0. CO; 2" type="journal article" year="2006">
Sanford
<emphasis id="DEBCEAD6FFD970701B2AECD8FAF57484" box="[1377,1436,1705,1730]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2006
</bibRefCitation>
); 0.46% in
<taxonomicName id="2BC84D47FFD970701F18ECBCFEB274A0" box="[339,475,1741,1766]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="sindensis">
<emphasis id="DEBCEAD6FFD970701F18ECBCFEB274A0" box="[339,475,1741,1766]" italics="true" pageId="15" pageNumber="486">U. sindensis</emphasis>
</taxonomicName>
(recalculated from Shih
<emphasis id="DEBCEAD6FFD970701CBAECBCFC4074A0" box="[753,809,1741,1766]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2015); about 0.32% in
<taxonomicName id="2BC84D47FFD970701A7BECBCFBA574A3" box="[1072,1228,1742,1765]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="maracoani">
<emphasis id="DEBCEAD6FFD970701A7BECBCFBA574A3" box="[1072,1228,1742,1765]" italics="true" pageId="15" pageNumber="486">U. maracoani</emphasis>
</taxonomicName>
(cf.
<bibRefCitation id="88594B35FFD970701AB4ECBFFFBD754C" author="Wieman" pageId="15" pageNumber="486" refString="Wieman, A C., Berendzen, P. B., Hampton, K. R., Jang, J., Hopkins, M. J., Jurgenson, J., McNamara, J. C. &amp; Thurman, C. L. (2014) A panmictic fiddler crab from the coast of Brazil? Impact of divergent ocean currents and larval dispersal potential on genetic and morphological variation in Uca maracoani. Marine Biology, 161, 173 - 185." type="journal article" year="2014">
Wieman
<emphasis id="DEBCEAD6FFD970701B2FECBCFAF574A0" box="[1380,1436,1741,1766]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2014
</bibRefCitation>
); and about 0.13% in
<taxonomicName id="2BC84D47FFD970701F87EC80FD34754C" box="[460,605,1777,1802]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="annulipes">
<emphasis id="DEBCEAD6FFD970701F87EC80FD34754C" box="[460,605,1777,1802]" italics="true" pageId="15" pageNumber="486">U. annulipes</emphasis>
</taxonomicName>
(=
<taxonomicName id="2BC84D47FFD970701CCDEC80FC2A754C" box="[646,835,1777,1802]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="annulipes">
<emphasis id="DEBCEAD6FFD970701CCDEC80FDC8754F" box="[646,673,1778,1801]" italics="true" pageId="15" pageNumber="486">U.</emphasis>
aff.
<emphasis id="DEBCEAD6FFD970701C9DEC83FC2A754C" box="[726,835,1777,1802]" italics="true" pageId="15" pageNumber="486">annulipes</emphasis>
</taxonomicName>
, see
<bibRefCitation id="88594B35FFD970701D35EC83FB2B754C" author="Shih" box="[894,1090,1777,1802]" pageId="15" pageNumber="486" refString="Shih, H. - T., Ng, P. K. L. &amp; Liu, M. - Y. (2013 b) Systematics of the Indo-West Pacific broad-fronted fiddler crabs (Crustacea: Ocypodidae: genus Uca). Raffles Bulletin of Zoology, 61, 641 - 649." type="journal article" year="2013" yearSuffix="b">
Shih
<emphasis id="DEBCEAD6FFD970701DF1EC80FC9A754C" box="[954,1011,1777,1802]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2013b
</bibRefCitation>
) from East Africa (
<bibRefCitation id="88594B35FFD970701B6BEC83FFBD7568" author="Silva" pageId="15" pageNumber="486" refString="Silva, I. C., Mesquita, N. &amp; Paula, J. (2010) Lack of population structure in the fiddler crab Uca annulipes along an East African latitudinal gradient: genetic and morphometric evidence. Marine Biology, 157, 1113 - 1126." type="journal article" year="2010">
Silva
<emphasis id="DEBCEAD6FFD970701B28EC80FAF5754C" box="[1379,1436,1777,1802]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2010
</bibRefCitation>
). The high genetic variability within a species may be explained by past historical or present barriers (e.g., ocean currents and salinity) affecting their larval dispersal (see
<bibRefCitation id="88594B35FFD970701D17ED48FB1E7514" author="Lopez-Duarte" box="[860,1143,1849,1874]" pageId="15" pageNumber="486" refString="Lopez-Duarte, P. C., Christy, J. H. &amp; Tankersley, R. A. (2011) A behavioral mechanism for dispersal in fiddler crab larvae (genus Uca) varies with adult habitat, not phylogeny. Limnology and Oceanography, 56, 1879 - 1892. http: // dx. doi. org / 10.4319 / lo. 2011.56.5.1879" type="journal article" year="2011">
López-Duarte
<emphasis id="DEBCEAD6FFD970701DB5ED48FB5E7514" box="[1022,1079,1849,1874]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2011
</bibRefCitation>
; Shih
<emphasis id="DEBCEAD6FFD970701AF6ED48FB9C7514" box="[1213,1269,1849,1874]" italics="true" pageId="15" pageNumber="486">et al.</emphasis>
2015).
</paragraph>
<paragraph id="EC7736C4FFD9706F1E8CED2FFDF47293" blockId="15.[151,1437,622,2018]" lastBlockId="16.[151,1437,151,1977]" lastPageId="16" lastPageNumber="487" pageId="15" pageNumber="486">
Within the
<taxonomicName id="2BC84D47FFD970701F02ED2CFEF77533" box="[329,414,1886,1909]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFD970701F02ED2CFEF77533" box="[329,414,1886,1909]" italics="true" pageId="15" pageNumber="486">Minuca</emphasis>
</taxonomicName>
clade,
<taxonomicName id="2BC84D47FFD970701FB8ED2DFDE07530" box="[499,649,1885,1910]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="brevifrons">
<emphasis id="DEBCEAD6FFD970701FB8ED2DFDE07530" box="[499,649,1885,1910]" italics="true" pageId="15" pageNumber="486">U. brevifrons</emphasis>
</taxonomicName>
diverged earliest at 8 mya (p-distance = 9.27%), and is sister to the remaining species of this subgenus. The habitat of
<taxonomicName id="2BC84D47FFD970701C9AEDF0FC0C75DC" box="[721,869,1921,1946]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="15" pageNumber="486" phylum="Arthropoda" rank="species" species="brevifrons">
<emphasis id="DEBCEAD6FFD970701C9AEDF0FC0C75DC" box="[721,869,1921,1946]" italics="true" pageId="15" pageNumber="486">U. brevifrons</emphasis>
</taxonomicName>
is unusual. This species lives on the muddy banks of freshwater and brackish streams, sometimes even in wet forest several meters from the nearest stream (
<bibRefCitation id="88594B35FFD970701B13EDD4FFB875A4" author="Crane" pageId="15" pageNumber="486" refString="Crane, J. (1941) Crabs of the genus Uca from the west coast of Central America. Eastern Pacific Expeditions of the New York Zoological Society. XXVI. Zoologica, 26 (19), 145 - 208." type="journal article" year="1941">Crane 1941</bibRefCitation>
,
<bibRefCitation id="88594B35FFD970701E95EDBBFE7375A4" author="Crane" box="[222,282,1993,2018]" pageId="15" pageNumber="486" refString="Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp." type="book" year="1975">1975</bibRefCitation>
). Perhaps the simpler form of urocardiac ossicles of gastric mill present in this species (
<figureCitation id="74F32A41FFD970701AB5EDBBFA2B75A4" box="[1278,1346,1993,2018]" captionStart="FIGURE 6" captionStartId="11.[151,250,1886,1908]" captionTargetBox="[173,1413,207,1861]" captionTargetId="figure@11.[163,1423,193,1868]" captionTargetPageId="11" captionText="FIGURE 6. Urocardiac ossicle of the subgenus Minuca species. A, Uca brevifrons (ZRC 2012.0126, CW 22.1, ♂); B, U. cf. virens (NCHUZOOL 13946, CW 17.0 mm, ♂); C, U. victoriana (NCHUZOOL 13945, CW 12.2 mm, ♀); D, U. pugnax (NCHUZOOL 13941, CW 15.2 mm, ♂); E, U. burgersi (NCHUZOOL 13956, CW 13.0 mm); F, U. rapax (NCHUZOOL 13942, CW 23.4 mm, ♂); G, U. mordax (NCHUZOOL 13940, CW 18.0 mm, ♀); H, U. minax (NCHUZOOL 13957, CW 23.2 mm, ♂); I, U. vocator (NCHUZOOL 13948, CW 21.9 mm, ♀). Scale = 1 mm." httpUri="https://zenodo.org/record/235579/files/figure.png" pageId="15" pageNumber="486">Fig. 6</figureCitation>
A) is an ancestral character.
<bibRefCitation id="88594B35FFC6706F1F3DEAEAFD6572F6" author="Crane" box="[374,524,151,176]" pageId="16" pageNumber="487" refString="Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp." type="book" year="1975">Crane (1975)</bibRefCitation>
treated
<taxonomicName id="2BC84D47FFC6706F1C23EAEBFD9572F6" box="[616,764,151,176]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="brevifrons">
<emphasis id="DEBCEAD6FFC6706F1C23EAEBFD9572F6" box="[616,764,151,176]" italics="true" pageId="16" pageNumber="487">U. brevifrons</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1D7FEAEBFCF472F6" box="[820,925,153,176]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="minax">
<emphasis id="DEBCEAD6FFC6706F1D7FEAEBFCF472F6" box="[820,925,153,176]" italics="true" pageId="16" pageNumber="487">U. minax</emphasis>
</taxonomicName>
as the same “superspecies [
<taxonomicName id="2BC84D47FFC6706F1A90EAEBFA7672F6" box="[1243,1311,153,176]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="minax">
<emphasis id="DEBCEAD6FFC6706F1A90EAEBFA7672F6" box="[1243,1311,153,176]" italics="true" pageId="16" pageNumber="487">minax</emphasis>
</taxonomicName>
]” but their close relationship is not supported genetically.
</paragraph>
<paragraph id="EC7736C4FFC6706F1E8CEAADFE4D7323" blockId="16.[151,1437,151,1977]" pageId="16" pageNumber="487">
The taxonomy of
<taxonomicName id="2BC84D47FFC6706F1FDEEA93FC2A72BE" authority="Salmon &amp; Atsaides, 1968" authorityName="Salmon &amp; Atsaides" authorityYear="1968" box="[405,835,223,248]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="virens">
<emphasis id="DEBCEAD6FFC6706F1FDEEA93FD7972BE" box="[405,528,225,248]" italics="true" pageId="16" pageNumber="487">Uca virens</emphasis>
<bibRefCitation id="88594B35FFC6706F1C51EAADFC2A72BE" author="Salmon" box="[538,835,223,248]" pageId="16" pageNumber="487" refString="Salmon, M. &amp; Atsaides, S. P. (1968) Behavioral, morphological and ecological evidence for two new species of fiddler crabs (genus Uca) from the Gulf coast of the United States. Proceedings of the Biological Society of Washington, 81, 275 - 290." type="journal article" year="1968">Salmon &amp; Atsaides, 1968</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="2BC84D47FFC6706F1DC9EA93FA0072BE" authority="Salmon &amp; Atsaides, 1968" authorityName="Salmon &amp; Atsaides" authorityYear="1968" box="[898,1385,223,248]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="longisignalis">
<emphasis id="DEBCEAD6FFC6706F1DC9EA93FB5E72BE" box="[898,1079,223,248]" italics="true" pageId="16" pageNumber="487">U. longisignalis</emphasis>
<bibRefCitation id="88594B35FFC6706F1A0BEAADFA0072BE" author="Salmon" box="[1088,1385,223,248]" pageId="16" pageNumber="487" refString="Salmon, M. &amp; Atsaides, S. P. (1968) Behavioral, morphological and ecological evidence for two new species of fiddler crabs (genus Uca) from the Gulf coast of the United States. Proceedings of the Biological Society of Washington, 81, 275 - 290." type="journal article" year="1968">Salmon &amp; Atsaides, 1968</bibRefCitation>
</taxonomicName>
, has been debated since the publication of the description of the two species (
<bibRefCitation id="88594B35FFC6706F1D83EB76FB81735B" author="Salmon" box="[968,1256,260,285]" pageId="16" pageNumber="487" refString="Salmon, M. &amp; Atsaides, S. P. (1968) Behavioral, morphological and ecological evidence for two new species of fiddler crabs (genus Uca) from the Gulf coast of the United States. Proceedings of the Biological Society of Washington, 81, 275 - 290." type="journal article" year="1968">Salmon &amp; Atsaides 1968</bibRefCitation>
). By comparing the
<typeStatus id="33738866FFC6706F1E8BEB55FE447306" box="[192,301,295,320]" pageId="16" pageNumber="487" type="holotype">holotypes</typeStatus>
of the above two species, von
<bibRefCitation id="88594B35FFC6706F1CCDEB5AFC497306" author="Hagen" box="[646,800,296,320]" pageId="16" pageNumber="487" refString="Von Hagen, H. - O. (1980) A key to the &quot; x-species &quot; of North American fiddler crabs (genus Uca). Zoologische Mededelingen, Leiden, 55 (8), 87 - 96." type="journal article" year="1980">Hagen (1980)</bibRefCitation>
confirmed them as synonyms of
<taxonomicName id="2BC84D47FFC6706F1ADFEB5BFB9E7306" box="[1172,1271,297,320]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="rapax">
<emphasis id="DEBCEAD6FFC6706F1ADFEB5BFB9E7306" box="[1172,1271,297,320]" italics="true" pageId="16" pageNumber="487">U. rapax</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1B65EB5BFAFD7306" box="[1326,1428,297,320]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="minax">
<emphasis id="DEBCEAD6FFC6706F1B65EB5BFAFD7306" box="[1326,1428,297,320]" italics="true" pageId="16" pageNumber="487">U. minax</emphasis>
</taxonomicName>
, respectively.
</paragraph>
<paragraph id="EC7736C4FFC6706F1E8CEB1DFB2973B2" blockId="16.[151,1437,151,1977]" pageId="16" pageNumber="487">
The synonymization of
<taxonomicName id="2BC84D47FFC6706F1F92EB03FD2873CE" box="[473,577,369,392]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="virens">
<emphasis id="DEBCEAD6FFC6706F1F92EB03FD2873CE" box="[473,577,369,392]" italics="true" pageId="16" pageNumber="487">U. virens</emphasis>
</taxonomicName>
as
<taxonomicName id="2BC84D47FFC6706F1C20EB03FDB973CE" box="[619,720,369,392]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="rapax">
<emphasis id="DEBCEAD6FFC6706F1C20EB03FDB973CE" box="[619,720,369,392]" italics="true" pageId="16" pageNumber="487">U. rapax</emphasis>
</taxonomicName>
has been followed by later studies (e.g.
<bibRefCitation id="88594B35FFC6706F1AEDEB1DFFBA73EA" author="Barnwell" pageId="16" pageNumber="487" refString="Barnwell, F. H. &amp; Thurman, C. L. (1984) Taxonomy and biogeography of the fiddler crabs (Ocypodidae: genus Uca) of the Atlantic and Gulf coasts of eastern North America. Zoological Journal of the Linnean Society, 81, 23 - 87." type="journal article" year="1984">Barnwell &amp; Thurman 1984</bibRefCitation>
; Beinlich &amp; von
<bibRefCitation id="88594B35FFC6706F1FEEEBE7FD5173EB" author="Hagen" box="[421,568,404,429]" pageId="16" pageNumber="487" refString="Beinlich, B. &amp; von Hagen, H. O. (2006) Materials for a more stable subdivision of the genus Uca Leach. Zoologische Mededelingen, 80, 9 - 32." type="journal article" year="2006">Hagen 2006</bibRefCitation>
). Although there are several studies still using the name of “
<taxonomicName id="2BC84D47FFC6706F1B5AEBE7FAE673EA" box="[1297,1423,405,428]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="virens">
<emphasis id="DEBCEAD6FFC6706F1B5AEBE7FAE673EA" box="[1297,1423,405,428]" italics="true" pageId="16" pageNumber="487">Uca virens</emphasis>
</taxonomicName>
” (
<bibRefCitation id="88594B35FFC6706F1ED5EBC5FE327396" author="Rosenberg" box="[158,347,439,464]" pageId="16" pageNumber="487" refString="Rosenberg, M. S. (2001) The systematics and taxonomy of fiddler crabs: a phylogeny of the genus Uca. Journal of Crustacean Biology, 21, 839 - 869. http: // dx. doi. org / 10.1163 / 20021975 - 99990176" type="journal article" year="2001">Rosenberg 2001</bibRefCitation>
;
<bibRefCitation id="88594B35FFC6706F1F2DEBCAFD247396" author="Swanson" box="[358,589,439,464]" pageId="16" pageNumber="487" refString="Swanson, B. O., George, M. N., Anderson, S. P. &amp; Christy, J. H. (2013) Evolutionary variation in the mechanics of fiddler crab claws. BMC Evolutionary Biology, 13, 137. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 13 - 137" type="journal article" year="2013">
Swanson
<emphasis id="DEBCEAD6FFC6706F1F98EBCBFD627396" box="[467,523,439,464]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
2013
</bibRefCitation>
;
<bibRefCitation id="88594B35FFC6706F1C11EBC5FCF27396" author="Perez-Mozqueda" box="[602,923,439,464]" pageId="16" pageNumber="487" refString="Perez-Mozqueda, L. L., Del Castillo-Falconi, V. &amp; Luis Bortolini-Rosales, J. (2014) Registros adicionales del genero Uca (Brachyura: Ocypodidae) en la laguna de Tamiahua, Veracruz, Mexico. [Additional records of the genus Uca (Brachyura: Ocypodidae) in Tamiahua Lagoon, Veracruz, Mexico.] Revista Mexicana De Biodiversidad, 85, 969 - 971. http: // dx. doi. org / 10.7550 / rmb. 36600" type="journal article" year="2014">
Pérez-Mozqueda
<emphasis id="DEBCEAD6FFC6706F1D54EBCBFC317396" box="[799,856,439,464]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
2014
</bibRefCitation>
; del Castillo
<emphasis id="DEBCEAD6FFC6706F1A65EBCBFB0E7396" box="[1070,1127,439,464]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
2015), the identification of the species in these studies will need to be reconfirmed (C. Thurman, pers. comm.).
</paragraph>
<paragraph id="EC7736C4FFC6706F1E8CEB8DFBB6708B" blockId="16.[151,1437,151,1977]" pageId="16" pageNumber="487">
Based on the allozyme study of hemolymph amylases (
<bibRefCitation id="88594B35FFC6706F1D1BEB8DFBF9705E" author="Barnwell" box="[848,1168,511,536]" pageId="16" pageNumber="487" refString="Barnwell, F. H. &amp; Thurman, C. L. (1984) Taxonomy and biogeography of the fiddler crabs (Ocypodidae: genus Uca) of the Atlantic and Gulf coasts of eastern North America. Zoological Journal of the Linnean Society, 81, 23 - 87." type="journal article" year="1984">Barnwell &amp; Thurman 1984</bibRefCitation>
) and esterase-1 and -2 (
<bibRefCitation id="88594B35FFC6706F1ED5E856FEC0707A" author="Salmon" box="[158,425,548,573]" pageId="16" pageNumber="487" refString="Salmon, M. &amp; Kettler, M. K. (1987) The importance of behavioral and biochemical differences between fiddler crab taxa, with special reference to Uca rapax (Smith) and U. virens (Salmon and Atsaides). Contributions in Marine Science, 30, 63 - 76." type="journal article" year="1987">Salmon &amp; Kettler 1987</bibRefCitation>
), it is apparent that there are two “forms” of
<taxonomicName id="2BC84D47FFC6706F1DD5E857FB68707A" box="[926,1025,549,572]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="rapax">
<emphasis id="DEBCEAD6FFC6706F1DD5E857FB68707A" box="[926,1025,549,572]" italics="true" pageId="16" pageNumber="487">U. rapax</emphasis>
</taxonomicName>
, corresponding to the distribution of
<taxonomicName id="2BC84D47FFC6706F1EDCE83BFF947026" box="[151,253,585,608]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="virens">
<emphasis id="DEBCEAD6FFC6706F1EDCE83BFF947026" box="[151,253,585,608]" italics="true" pageId="16" pageNumber="487">U. virens</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1F79E83BFEFC7026" box="[306,405,585,608]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="rapax">
<emphasis id="DEBCEAD6FFC6706F1F79E83BFEFC7026" box="[306,405,585,608]" italics="true" pageId="16" pageNumber="487">U. rapax</emphasis>
</taxonomicName>
. In our study based on 16S, COI and 28S markers (
<figureCitation id="74F32A41FFC6706F1D9FE835FB717026" box="[980,1048,583,608]" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="16" pageNumber="487">Fig. 9</figureCitation>
), there is a clade sister to
<taxonomicName id="2BC84D47FFC6706F1B72E83BFAF57026" box="[1337,1436,585,608]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="rapax">
<emphasis id="DEBCEAD6FFC6706F1B72E83BFAF57026" box="[1337,1436,585,608]" italics="true" pageId="16" pageNumber="487">U. rapax</emphasis>
</taxonomicName>
with similar morphology. Before additional studies are done to confirm whether it is actually
<taxonomicName id="2BC84D47FFC6706F1AFAE81FFA7F70C2" box="[1201,1302,621,644]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="virens">
<emphasis id="DEBCEAD6FFC6706F1AFAE81FFA7F70C2" box="[1201,1302,621,644]" italics="true" pageId="16" pageNumber="487">U. virens</emphasis>
</taxonomicName>
, we use the name “
<taxonomicName id="2BC84D47FFC6706F1EADE8E3FE0770EE" box="[230,366,655,680]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="virens">
<emphasis id="DEBCEAD6FFC6706F1EADE8E3FF9370EE" box="[230,250,657,680]" italics="true" pageId="16" pageNumber="487">U</emphasis>
. cf.
<emphasis id="DEBCEAD6FFC6706F1F60E8E3FE0770EE" box="[299,366,657,680]" italics="true" pageId="16" pageNumber="487">virens</emphasis>
</taxonomicName>
” for convenience. The two species diverged at 2 mya (p-distance = 2.32%), although
<bibRefCitation id="88594B35FFC6706F1B61E8FDFE5F708A" author="Salmon" pageId="16" pageNumber="487" refString="Salmon, M. &amp; Kettler, M. K. (1987) The importance of behavioral and biochemical differences between fiddler crab taxa, with special reference to Uca rapax (Smith) and U. virens (Salmon and Atsaides). Contributions in Marine Science, 30, 63 - 76." type="journal article" year="1987">Salmon &amp; Kettler (1987)</bibRefCitation>
estimated the divergence time is 2050 thousand years based on the allozyme data.
</paragraph>
<paragraph id="EC7736C4FFC6706F1E8CE8A5FA7A711A" blockId="16.[151,1437,151,1977]" pageId="16" pageNumber="487">
While previous studies considered the morphological characters of
<taxonomicName id="2BC84D47FFC6706F1DFDE8ABFB7270B6" box="[950,1051,729,752]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="virens">
<emphasis id="DEBCEAD6FFC6706F1DFDE8ABFB7270B6" box="[950,1051,729,752]" italics="true" pageId="16" pageNumber="487">U. virens</emphasis>
</taxonomicName>
were within the range of
<taxonomicName id="2BC84D47FFC6706F1B72E8ABFAF570B6" box="[1337,1436,729,752]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="rapax">
<emphasis id="DEBCEAD6FFC6706F1B72E8ABFAF570B6" box="[1337,1436,729,752]" italics="true" pageId="16" pageNumber="487">U. rapax</emphasis>
</taxonomicName>
(
<bibRefCitation id="88594B35FFC6706F1ED5E88EFE427152" author="Hagen" box="[158,299,764,788]" pageId="16" pageNumber="487" refString="Von Hagen, H. - O. (1980) A key to the &quot; x-species &quot; of North American fiddler crabs (genus Uca). Zoologische Mededelingen, Leiden, 55 (8), 87 - 96." type="journal article" year="1980">Hagen 1980</bibRefCitation>
;
<bibRefCitation id="88594B35FFC6706F1F7CE88EFD037152" author="Barnwell" box="[311,618,764,789]" pageId="16" pageNumber="487" refString="Barnwell, F. H. &amp; Thurman, C. L. (1984) Taxonomy and biogeography of the fiddler crabs (Ocypodidae: genus Uca) of the Atlantic and Gulf coasts of eastern North America. Zoological Journal of the Linnean Society, 81, 23 - 87." type="journal article" year="1984">Barnwell &amp; Thurman 1984</bibRefCitation>
), their gastric mill structures show differences in the posterior tooth plate of urocardiac ossicles, with 8 ridges in
<taxonomicName id="2BC84D47FFC6706F1C2AE953FDAE717E" box="[609,711,801,824]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="rapax">
<emphasis id="DEBCEAD6FFC6706F1C2AE953FDAE717E" box="[609,711,801,824]" italics="true" pageId="16" pageNumber="487">U. rapax</emphasis>
</taxonomicName>
(
<figureCitation id="74F32A41FFC6706F1C91E96DFC76717E" box="[730,799,799,824]" captionStart="FIGURE 6" captionStartId="11.[151,250,1886,1908]" captionTargetBox="[173,1413,207,1861]" captionTargetId="figure@11.[163,1423,193,1868]" captionTargetPageId="11" captionText="FIGURE 6. Urocardiac ossicle of the subgenus Minuca species. A, Uca brevifrons (ZRC 2012.0126, CW 22.1, ♂); B, U. cf. virens (NCHUZOOL 13946, CW 17.0 mm, ♂); C, U. victoriana (NCHUZOOL 13945, CW 12.2 mm, ♀); D, U. pugnax (NCHUZOOL 13941, CW 15.2 mm, ♂); E, U. burgersi (NCHUZOOL 13956, CW 13.0 mm); F, U. rapax (NCHUZOOL 13942, CW 23.4 mm, ♂); G, U. mordax (NCHUZOOL 13940, CW 18.0 mm, ♀); H, U. minax (NCHUZOOL 13957, CW 23.2 mm, ♂); I, U. vocator (NCHUZOOL 13948, CW 21.9 mm, ♀). Scale = 1 mm." httpUri="https://zenodo.org/record/235579/files/figure.png" pageId="16" pageNumber="487">Fig. 6</figureCitation>
F) and 6 ridges in
<taxonomicName id="2BC84D47FFC6706F1DB1E953FB0B717E" box="[1018,1122,801,824]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="virens">
<emphasis id="DEBCEAD6FFC6706F1DB1E953FB0B717E" box="[1018,1122,801,824]" italics="true" pageId="16" pageNumber="487">U. virens</emphasis>
</taxonomicName>
(
<figureCitation id="74F32A41FFC6706F1A3FE96DFBD2717E" box="[1140,1211,799,824]" captionStart="FIGURE 6" captionStartId="11.[151,250,1886,1908]" captionTargetBox="[173,1413,207,1861]" captionTargetId="figure@11.[163,1423,193,1868]" captionTargetPageId="11" captionText="FIGURE 6. Urocardiac ossicle of the subgenus Minuca species. A, Uca brevifrons (ZRC 2012.0126, CW 22.1, ♂); B, U. cf. virens (NCHUZOOL 13946, CW 17.0 mm, ♂); C, U. victoriana (NCHUZOOL 13945, CW 12.2 mm, ♀); D, U. pugnax (NCHUZOOL 13941, CW 15.2 mm, ♂); E, U. burgersi (NCHUZOOL 13956, CW 13.0 mm); F, U. rapax (NCHUZOOL 13942, CW 23.4 mm, ♂); G, U. mordax (NCHUZOOL 13940, CW 18.0 mm, ♀); H, U. minax (NCHUZOOL 13957, CW 23.2 mm, ♂); I, U. vocator (NCHUZOOL 13948, CW 21.9 mm, ♀). Scale = 1 mm." httpUri="https://zenodo.org/record/235579/files/figure.png" pageId="16" pageNumber="487">Fig. 6</figureCitation>
B). More studies of morphological characters, especially the gastric mills, are necessary to clarify the identity of
<taxonomicName id="2BC84D47FFC6706F1AEDE937FA65711A" box="[1190,1292,837,860]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="virens">
<emphasis id="DEBCEAD6FFC6706F1AEDE937FA65711A" box="[1190,1292,837,860]" italics="true" pageId="16" pageNumber="487">U. virens</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="EC7736C4FFC6706F1E8CE915FE237683" blockId="16.[151,1437,151,1977]" pageId="16" pageNumber="487">
Another confused species is
<taxonomicName id="2BC84D47FFC6706F1C43E91BFDD271C6" box="[520,699,871,896]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="longisignalis">
<emphasis id="DEBCEAD6FFC6706F1C43E91BFDD271C6" box="[520,699,871,896]" italics="true" pageId="16" pageNumber="487">U. longisignalis</emphasis>
</taxonomicName>
, which has been synonymized as
<taxonomicName id="2BC84D47FFC6706F1A73E91BFBF671C6" box="[1080,1183,873,896]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="minax">
<emphasis id="DEBCEAD6FFC6706F1A73E91BFBF671C6" box="[1080,1183,873,896]" italics="true" pageId="16" pageNumber="487">U. minax</emphasis>
</taxonomicName>
by von
<bibRefCitation id="88594B35FFC6706F1AB1E91AFAF171C6" author="Hagen" box="[1274,1432,872,896]" pageId="16" pageNumber="487" refString="Von Hagen, H. - O. (1980) A key to the &quot; x-species &quot; of North American fiddler crabs (genus Uca). Zoologische Mededelingen, Leiden, 55 (8), 87 - 96." type="journal article" year="1980">Hagen (1980)</bibRefCitation>
.
<bibRefCitation id="88594B35FFC6706F1EDCE9FEFE3871E2" author="Thurman" box="[151,337,908,933]" pageId="16" pageNumber="487" refString="Thurman, C. L. (1982) On the distinctness of the fiddler crabs Uca minax (Leconte) and Uca longisignalis Salmon &amp; Atsaides in their region of Sympatry (Decapoda Brachyura, Ocypodidae). Crustaceana, 43, 37 - 50." type="journal article" year="1982">Thurman (1982)</bibRefCitation>
considered it is a valid species because there are substantial morphological differences between the two species, a decision that has been followed by other studies thereafter. Genetically,
<bibRefCitation id="88594B35FFC6706F1AD0E9DDFAF5718E" author="Felder" box="[1179,1436,943,968]" pageId="16" pageNumber="487" refString="Felder, D. L. &amp; Staton, J. L. (1994) Genetic differentiation in trans-Floridian species complexes of Sesarma and Uca (Decapoda: Brachyura). Journal of Crustacean Biology, 14, 191 - 209. http: // dx. doi. org / 10.2307 / 1548900" type="journal article" year="1994">Felder &amp; Saton (1994)</bibRefCitation>
showed that the allozyme evidence did not support the hypothesis that
<taxonomicName id="2BC84D47FFC6706F1DFFE9A7FB0E71AB" box="[948,1127,980,1005]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="longisignalis">
<emphasis id="DEBCEAD6FFC6706F1DFFE9A7FB0E71AB" box="[948,1127,980,1005]" italics="true" pageId="16" pageNumber="487">U. longisignalis</emphasis>
</taxonomicName>
is different from
<taxonomicName id="2BC84D47FFC6706F1B65E9A7FAFD71AA" box="[1326,1428,981,1004]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="minax">
<emphasis id="DEBCEAD6FFC6706F1B65E9A7FAFD71AA" box="[1326,1428,981,1004]" italics="true" pageId="16" pageNumber="487">U. minax</emphasis>
</taxonomicName>
.
<bibRefCitation id="88594B35FFC6706F1EDCE985FED37656" box="[151,442,1015,1040]" pageId="16" pageNumber="487" refString="Landstorfer, R. B., Schubart, C. D. &amp; Felder, D. L. (2008) Barcoding gene COI fails to distinguish between two fiddler crabs (Brachyura: Ocypodidae: Uca) across their entire range of geographic overlap. Annual Meeting, The Crustacean Society, Galveston, Texas." type="book">
Landstorfer
<emphasis id="DEBCEAD6FFC6706F1F6DE98BFE0A7656" box="[294,355,1015,1040]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
(2008)
</bibRefCitation>
and
<bibRefCitation id="88594B35FFC6706F1FB0E985FC697656" box="[507,768,1015,1040]" pageId="16" pageNumber="487" refString="Warwick, A. R., Hopkins, M. J., Berendzen, P. B. &amp; Thurman, C. L. (2009) Morphological, physiological, and genetic variation in the red-jointed fiddler crab, Uca minax (Le Conte). Annual Meeting, Society for Integrative and Comparative Biology, Boston, MA, USA" type="book">
Warwick
<emphasis id="DEBCEAD6FFC6706F1C27E98BFDC37656" box="[620,682,1015,1040]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
(2009)
</bibRefCitation>
also reported the failure to separate the two species by mitochondrial COI or nuclear internal transcribed spacer (ITS-1) markers. We included in our study specimens of the two species with typical characters of each taxon (mainly based on with or without pubescence ventrally; cf.
<bibRefCitation id="88594B35FFC6706F1EDCEE16FEA7763A" author="Barnwell" box="[151,462,1124,1149]" pageId="16" pageNumber="487" refString="Barnwell, F. H. &amp; Thurman, C. L. (1984) Taxonomy and biogeography of the fiddler crabs (Ocypodidae: genus Uca) of the Atlantic and Gulf coasts of eastern North America. Zoological Journal of the Linnean Society, 81, 23 - 87." type="journal article" year="1984">Barnwell &amp; Thurman 1984</bibRefCitation>
), but the molecular data only shows one clade (
<figureCitation id="74F32A41FFC6706F1DA1EE16FB46763A" box="[1002,1071,1124,1149]" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="16" pageNumber="487">Fig. 9</figureCitation>
), therefore agreeing with earlier genetic studies. Further studies, with more complete sampling, will be necessary to clarify the taxonomy of the
<taxonomicName id="2BC84D47FFC6706F1BCAEEFBFFB57682" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="minax">
<emphasis id="DEBCEAD6FFC6706F1BCAEEFBFFB57682" italics="true" pageId="16" pageNumber="487">U. minax</emphasis>
</taxonomicName>
complex.
</paragraph>
<paragraph id="EC7736C4FFC6706F1E8CEEBDFDF17712" blockId="16.[151,1437,151,1977]" pageId="16" pageNumber="487">
The close relationship of
<taxonomicName id="2BC84D47FFC6706F1FAAEEA3FD0876AE" box="[481,609,1231,1256]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="burgersi">
<emphasis id="DEBCEAD6FFC6706F1FAAEEA3FD0876AE" box="[481,609,1231,1256]" italics="true" pageId="16" pageNumber="487">U. burgersi</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1CDEEEA3FC6576AE" box="[661,780,1231,1256]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="mordax">
<emphasis id="DEBCEAD6FFC6706F1CDEEEA3FC6576AE" box="[661,780,1231,1256]" italics="true" pageId="16" pageNumber="487">U. mordax</emphasis>
</taxonomicName>
suggested by
<bibRefCitation id="88594B35FFC6706F1DE2EEBDFBA976AE" author="Salmon" box="[937,1216,1231,1256]" pageId="16" pageNumber="487" refString="Salmon, M. &amp; Kettler, M. K. (1987) The importance of behavioral and biochemical differences between fiddler crab taxa, with special reference to Uca rapax (Smith) and U. virens (Salmon and Atsaides). Contributions in Marine Science, 30, 63 - 76." type="journal article" year="1987">Salmon &amp; Kettler (1987)</bibRefCitation>
is highly supported (
<figureCitation id="74F32A41FFC6706F1ED4EE86FF8A774A" box="[159,227,1268,1293]" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="16" pageNumber="487">Fig. 9</figureCitation>
). Within the
<taxonomicName id="2BC84D47FFC6706F1F3FEE87FEA6774A" box="[372,463,1269,1292]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC6706F1F3FEE87FEA6774A" box="[372,463,1269,1292]" italics="true" pageId="16" pageNumber="487">Leptuca</emphasis>
</taxonomicName>
clade,
<taxonomicName id="2BC84D47FFC6706F1C56EE87FDF7774A" box="[541,670,1269,1292]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="panacea">
<emphasis id="DEBCEAD6FFC6706F1C56EE87FDF7774A" box="[541,670,1269,1292]" italics="true" pageId="16" pageNumber="487">U. panacea</emphasis>
</taxonomicName>
, and
<taxonomicName id="2BC84D47FFC6706F1C90EE87FC0D774B" box="[731,868,1268,1293]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="pugilator">
<emphasis id="DEBCEAD6FFC6706F1C90EE87FC0D774B" box="[731,868,1268,1293]" italics="true" pageId="16" pageNumber="487">U. pugilator</emphasis>
</taxonomicName>
are closely related, with the divergence time at 2.8 mya (p-distance = 3.24%), which supports the original hypothesis that both species have diverged recently (
<bibRefCitation id="88594B35FFC6706F1B1BEF65FE257712" author="Novak" pageId="16" pageNumber="487" refString="Novak, A. &amp; Salmon, M. (1974) Uca panacea, a new species of fiddler crab from the Gulf coast of the United States. Proceedings of the Biological Society of Washington, 87, 313 - 326." type="journal article" year="1974">Novak &amp; Salmon 1974</bibRefCitation>
;
<bibRefCitation id="88594B35FFC6706F1F1CEF4EFDE27712" author="Barnwell" box="[343,651,1340,1365]" pageId="16" pageNumber="487" refString="Barnwell, F. H. &amp; Thurman, C. L. (1984) Taxonomy and biogeography of the fiddler crabs (Ocypodidae: genus Uca) of the Atlantic and Gulf coasts of eastern North America. Zoological Journal of the Linnean Society, 81, 23 - 87." type="journal article" year="1984">Barnwell &amp; Thurman 1984</bibRefCitation>
).
</paragraph>
<paragraph id="EC7736C4FFC6706F1E8CEF2DFB04744E" blockId="16.[151,1437,151,1977]" pageId="16" pageNumber="487">
The supposed close relationship of
<taxonomicName id="2BC84D47FFC6706F1C18EF13FDBF773E" box="[595,726,1377,1400]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="speciosa">
<emphasis id="DEBCEAD6FFC6706F1C18EF13FDBF773E" box="[595,726,1377,1400]" italics="true" pageId="16" pageNumber="487">U. speciosa</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1D46EF13FCCE773E" box="[781,935,1377,1400]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="spinicarpa">
<emphasis id="DEBCEAD6FFC6706F1D46EF13FCCE773E" box="[781,935,1377,1400]" italics="true" pageId="16" pageNumber="487">U. spinicarpa</emphasis>
</taxonomicName>
(
<bibRefCitation id="88594B35FFC6706F1DFEEF12FB55773E" author="Crane" box="[949,1084,1375,1400]" pageId="16" pageNumber="487" refString="Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp." type="book" year="1975">Crane 1975</bibRefCitation>
;
<bibRefCitation id="88594B35FFC6706F1A03EF2DFA10773E" author="Barnwell" box="[1096,1401,1375,1400]" pageId="16" pageNumber="487" refString="Barnwell, F. H. &amp; Thurman, C. L. (1984) Taxonomy and biogeography of the fiddler crabs (Ocypodidae: genus Uca) of the Atlantic and Gulf coasts of eastern North America. Zoological Journal of the Linnean Society, 81, 23 - 87." type="journal article" year="1984">Barnwell &amp; Thurman 1984</bibRefCitation>
) is not supported in this study (
<figureCitation id="74F32A41FFC6706F1F85EFF6FD7B77DA" box="[462,530,1412,1437]" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="16" pageNumber="487">Fig. 9</figureCitation>
). Our study, however, agrees well with the allozyme studies (
<bibRefCitation id="88594B35FFC6706F1A8BEFF6FAF177DA" author="Salmon" box="[1216,1432,1412,1437]" pageId="16" pageNumber="487" refString="Salmon, M., Ferris, S. D., Johnston, D., Hyatt, G. &amp; Whitt, G. S. (1979) Behavioral and biochemical evidence for species distinctiveness in the fiddler crabs, Uca speciosa and U. spinicarpa. Evolution, 33, 182 - 191. http: // dx. doi. org / 10.2307 / 2407375" type="journal article" year="1979">
Salmon
<emphasis id="DEBCEAD6FFC6706F1B56EFF7FA3F77DB" box="[1309,1366,1412,1437]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
1979
</bibRefCitation>
;
<bibRefCitation id="88594B35FFC6706F1EDCEFD5FEF57786" author="Salmon" box="[151,412,1447,1472]" pageId="16" pageNumber="487" refString="Salmon, M. &amp; Kettler, M. K. (1987) The importance of behavioral and biochemical differences between fiddler crab taxa, with special reference to Uca rapax (Smith) and U. virens (Salmon and Atsaides). Contributions in Marine Science, 30, 63 - 76." type="journal article" year="1987">Salmon &amp; Kettler 1987</bibRefCitation>
) that the genetic distance of two species is large.
<taxonomicName id="2BC84D47FFC6706F1D8BEFDBFB3F7786" box="[960,1110,1449,1472]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="speciosa">
<emphasis id="DEBCEAD6FFC6706F1D8BEFDBFB3F7786" box="[960,1110,1449,1472]" italics="true" pageId="16" pageNumber="487">Uca speciosa</emphasis>
</taxonomicName>
is distinct from other species genetically, although it forms a main clade, with
<taxonomicName id="2BC84D47FFC6706F1CF4EFBFFC3B77A3" box="[703,850,1484,1509]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="dorotheae">
<emphasis id="DEBCEAD6FFC6706F1CF4EFBFFC3B77A3" box="[703,850,1484,1509]" italics="true" pageId="16" pageNumber="487">U. dorotheae</emphasis>
</taxonomicName>
,
<taxonomicName id="2BC84D47FFC6706F1D29EFBFFB6E77A3" box="[866,1031,1484,1509]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="deichmanni">
<emphasis id="DEBCEAD6FFC6706F1D29EFBFFB6E77A3" box="[866,1031,1484,1509]" italics="true" pageId="16" pageNumber="487">U. deichmanni</emphasis>
</taxonomicName>
, and
<taxonomicName id="2BC84D47FFC6706F1A0DEFBFFBB677A3" box="[1094,1247,1484,1509]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="cumulanta">
<emphasis id="DEBCEAD6FFC6706F1A0DEFBFFBB677A3" box="[1094,1247,1484,1509]" italics="true" pageId="16" pageNumber="487">U. cumulanta</emphasis>
</taxonomicName>
.
<taxonomicName id="2BC84D47FFC6706F1AA5EFBFFAF577A2" box="[1262,1436,1485,1508]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="spinicarpa">
<emphasis id="DEBCEAD6FFC6706F1AA5EFBFFAF577A2" box="[1262,1436,1485,1508]" italics="true" pageId="16" pageNumber="487">Uca spinicarpa</emphasis>
</taxonomicName>
is, however, closely related to the species pair,
<taxonomicName id="2BC84D47FFC6706F1CEFEF83FC7F744E" box="[676,790,1519,1544]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="thayeri">
<emphasis id="DEBCEAD6FFC6706F1CEFEF83FC7F744E" box="[676,790,1519,1544]" italics="true" pageId="16" pageNumber="487">U. thayeri</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1D07EF83FCB4744E" box="[844,989,1519,1544]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC6706F1D07EF83FCB4744E" box="[844,989,1519,1544]" italics="true" pageId="16" pageNumber="487">U. umbratila</emphasis>
</taxonomicName>
(see below).
</paragraph>
<paragraph id="EC7736C4FFC6706F1E8CEC66FCB075FE" blockId="16.[151,1437,151,1977]" pageId="16" pageNumber="487">
Cranes (1975) subgenus
<emphasis id="DEBCEAD6FFC6706F1FA3EC66FD3F746B" box="[488,598,1556,1581]" italics="true" pageId="16" pageNumber="487">Boboruca</emphasis>
(=
<taxonomicName id="2BC84D47FFC6706F1CCBEC66FC09746A" authority="Bott, 1973" authorityName="Bott" authorityYear="1973" box="[640,864,1556,1581]" class="Malacostraca" family="Ocypodidae" genus="Planuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC6706F1CCBEC66FDB4746B" box="[640,733,1556,1581]" italics="true" pageId="16" pageNumber="487">Planuca</emphasis>
<bibRefCitation id="88594B35FFC6706F1CADEC67FC09746A" author="Bott" box="[742,864,1556,1580]" pageId="16" pageNumber="487" refString="Bott, R. (1973) Die verwandtschaftlichen Beziehungen der Uca Arten. Senckenbergiana Biologica, 54, 315 - 325." type="journal article" year="1973">Bott, 1973</bibRefCitation>
</taxonomicName>
), established for
<taxonomicName id="2BC84D47FFC6706F1A68EC67FBFF746B" box="[1059,1174,1556,1581]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="thayeri">
<emphasis id="DEBCEAD6FFC6706F1A68EC67FBFF746B" box="[1059,1174,1556,1581]" italics="true" pageId="16" pageNumber="487">U. thayeri</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1A84EC67FA08746B" box="[1231,1377,1556,1581]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC6706F1A84EC67FA08746B" box="[1231,1377,1556,1581]" italics="true" pageId="16" pageNumber="487">U. umbratila</emphasis>
</taxonomicName>
, was synonymized with
<taxonomicName id="2BC84D47FFC6706F1F38EC4BFEA07416" box="[371,457,1593,1616]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC6706F1F38EC4BFEA07416" box="[371,457,1593,1616]" italics="true" pageId="16" pageNumber="487">Minuca</emphasis>
</taxonomicName>
by Albrecht &amp; von
<bibRefCitation id="88594B35FFC6706F1CF0EC4AFC357416" author="Hagen" box="[699,860,1592,1616]" pageId="16" pageNumber="487" refString="Albrecht, H. &amp; von Hagen, H. O. (1981) Differential weighting of electrophoretic data in crayfish and fiddler crabs (Decapoda: Astacidae and Ocypodidae). Comparative Biochemistry and Physiology, 70 B, 393 - 399. http: // dx. doi. org / 10.1016 / 0305 - 0491 (81) 90272 - 8" type="journal article" year="1981">Hagen (1981)</bibRefCitation>
, based on combined allozyme and morphological characters, and this has been followed by all later authors (e.g., von
<bibRefCitation id="88594B35FFC6706F1D93EC2FFB007432" author="Hagen" box="[984,1129,1628,1653]" pageId="16" pageNumber="487" refString="Von Hagen, H. - O. (1987) Morphologie und Winkbalz einer neuen Uca - Art (Crustacea, Brachyura) aus dem Staat Espirito Santo (Brasilien). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 84, 81 - 94." type="journal article" year="1987">Hagen 1987</bibRefCitation>
;
<bibRefCitation id="88594B35FFC6706F1A30EC2EFAFE7432" author="Salmon" box="[1147,1431,1628,1653]" pageId="16" pageNumber="487" refString="Salmon, M. &amp; Zucker, N. (1988) Interpreting differences in the reproductive behaviour of fiddler crabs (genus Uca). In: Chelazzi, G. &amp; Vannini, M. (Eds.), Behavioral Adaptation to Intertidal Life. Plenum Press, New York, pp. 387 - 407. http: // dx. doi. org / 10.1007 / 978 - 1 - 4899 - 3737 - 7 _ 26" type="book chapter" year="1988">Salmon &amp; Zucker 1988</bibRefCitation>
;
<bibRefCitation id="88594B35FFC6706F1EDCEC0DFE3974DE" author="Rosenberg" box="[151,336,1663,1688]" pageId="16" pageNumber="487" refString="Rosenberg, M. S. (2001) The systematics and taxonomy of fiddler crabs: a phylogeny of the genus Uca. Journal of Crustacean Biology, 21, 839 - 869. http: // dx. doi. org / 10.1163 / 20021975 - 99990176" type="journal article" year="2001">Rosenberg 2001</bibRefCitation>
; Beinlich &amp; von
<bibRefCitation id="88594B35FFC6706F1C5BECF2FDF574DE" author="Hagen" box="[528,668,1663,1688]" pageId="16" pageNumber="487" refString="Beinlich, B. &amp; von Hagen, H. O. (2006) Materials for a more stable subdivision of the genus Uca Leach. Zoologische Mededelingen, 80, 9 - 32." type="journal article" year="2006">Hagen 2006</bibRefCitation>
; Ng
<emphasis id="DEBCEAD6FFC6706F1C9BECF3FC6074DE" box="[720,777,1663,1688]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
2008). In this study,
<taxonomicName id="2BC84D47FFC6706F1DBDECF3FB0E74DE" box="[1014,1127,1663,1688]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="thayeri">
<emphasis id="DEBCEAD6FFC6706F1DBDECF3FB0E74DE" box="[1014,1127,1663,1688]" italics="true" pageId="16" pageNumber="487">U. thayeri</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1AD5ECF3FA4674DE" box="[1182,1327,1663,1688]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC6706F1AD5ECF3FA4674DE" box="[1182,1327,1663,1688]" italics="true" pageId="16" pageNumber="487">U. umbratila</emphasis>
</taxonomicName>
came out clearly inside the
<taxonomicName id="2BC84D47FFC6706F1F17ECD7FEDF74FA" box="[348,438,1701,1724]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC6706F1F17ECD7FEDF74FA" box="[348,438,1701,1724]" italics="true" pageId="16" pageNumber="487">Leptuca</emphasis>
</taxonomicName>
clade (
<figureCitation id="74F32A41FFC6706F1C43ECD6FD2574FA" box="[520,588,1700,1725]" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="16" pageNumber="487">Fig. 9</figureCitation>
). In the studies of Levinton
<emphasis id="DEBCEAD6FFC6706F1DCCECD7FCD674FB" box="[903,959,1700,1725]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
(1996) and Sturmbauer
<emphasis id="DEBCEAD6FFC6706F1A84ECD7FA6E74FB" box="[1231,1287,1700,1725]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
(1996) using the 16S marker, the position of
<taxonomicName id="2BC84D47FFC6706F1C4AECBBFDFA74A6" box="[513,659,1735,1760]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC6706F1C4AECBBFDFA74A6" box="[513,659,1735,1760]" italics="true" pageId="16" pageNumber="487">U. umbratila</emphasis>
</taxonomicName>
is within a “derived American clade” composed of a mixture of the subgenera
<taxonomicName id="2BC84D47FFC6706F1F5BEC9FFE0F7542" box="[272,358,1773,1796]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC6706F1F5BEC9FFE0F7542" box="[272,358,1773,1796]" italics="true" pageId="16" pageNumber="487">Minuca</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1FD6EC9FFE9E7542" box="[413,503,1773,1796]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC6706F1FD6EC9FFE9E7542" box="[413,503,1773,1796]" italics="true" pageId="16" pageNumber="487">Leptuca</emphasis>
</taxonomicName>
.
<taxonomicName id="2BC84D47FFC6706F1C4DEC9FFDC27543" box="[518,683,1772,1797]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC6706F1C4DEC9FFDC27543" box="[518,683,1772,1797]" italics="true" pageId="16" pageNumber="487">Uca umbratila</emphasis>
</taxonomicName>
is nevertheless located within
<taxonomicName id="2BC84D47FFC6706F1A4DEC9FFB357542" box="[1030,1116,1773,1796]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC6706F1A4DEC9FFB357542" box="[1030,1116,1773,1796]" italics="true" pageId="16" pageNumber="487">Minuca</emphasis>
</taxonomicName>
in Landstorfer &amp; Schubart's (2010) study, although their sequences of COI (
<accessionNumber id="F39BAB27FFC6706F1CF8ED62FC47756E" box="[691,814,1808,1832]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/FN430708" pageId="16" pageNumber="487">FN430708</accessionNumber>
) and 28S (
<accessionNumber id="F39BAB27FFC6706F1DE1ED62FB4C756E" box="[938,1061,1808,1832]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/FN430719" pageId="16" pageNumber="487">FN430719</accessionNumber>
) agree well with
<taxonomicName id="2BC84D47FFC6706F1AA7ED63FA17756E" box="[1260,1406,1807,1832]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC6706F1AA7ED63FA17756E" box="[1260,1406,1807,1832]" italics="true" pageId="16" pageNumber="487">U. umbratila</emphasis>
</taxonomicName>
in our study. The different conclusions are probably caused by the inclusion of 16S marker, as well as the number of species used in ingroups and outgroups. Hampton
<emphasis id="DEBCEAD6FFC6706F1CA1ED2BFC4E7536" box="[746,807,1879,1904]" italics="true" pageId="16" pageNumber="487">et al.</emphasis>
(2014) have pointed out the unsolved relationship of
<emphasis id="DEBCEAD6FFC6706F1EDCED0EFE6F75D3" box="[151,262,1916,1941]" italics="true" pageId="16" pageNumber="487">Boboruca</emphasis>
from the above studies. Our study gives genetic support for including
<taxonomicName id="2BC84D47FFC6706F1A56ED0FFBE775D3" box="[1053,1166,1916,1941]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="thayeri">
<emphasis id="DEBCEAD6FFC6706F1A56ED0FFBE775D3" box="[1053,1166,1916,1941]" italics="true" pageId="16" pageNumber="487">U. thayeri</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC6706F1A8EED0FFA3C75D3" box="[1221,1365,1916,1941]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC6706F1A8EED0FFA3C75D3" box="[1221,1365,1916,1941]" italics="true" pageId="16" pageNumber="487">U. umbratila</emphasis>
</taxonomicName>
in the subgenus
<taxonomicName id="2BC84D47FFC6706F1F4DEDD3FE0975FE" box="[262,352,1953,1976]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC6706F1F4DEDD3FE0975FE" box="[262,352,1953,1976]" italics="true" pageId="16" pageNumber="487">Leptuca</emphasis>
</taxonomicName>
(
<figureCitation id="74F32A41FFC6706F1F24EDD2FEDD75FE" box="[367,436,1952,1977]" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="16" pageNumber="487">Fig. 9</figureCitation>
), although their morphology is closer to
<taxonomicName id="2BC84D47FFC6706F1D37EDD3FCB875FE" box="[892,977,1953,1976]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="16" pageNumber="487" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC6706F1D37EDD3FCB875FE" box="[892,977,1953,1976]" italics="true" pageId="16" pageNumber="487">Minuca</emphasis>
</taxonomicName>
.
</paragraph>
<caption id="B8B7664CFFC7706E1EDCED06FEFA75B6" httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="17" pageNumber="488" targetBox="[160,1392,213,1872]" targetPageId="17">
<paragraph id="EC7736C4FFC7706E1EDCED06FEFA75B6" blockId="17.[151,1436,1908,2032]" pageId="17" pageNumber="488">
<emphasis id="DEBCEAD6FFC7706E1EDCED06FE6475CF" bold="true" box="[151,269,1908,1930]" pageId="17" pageNumber="488">FIGURE 9</emphasis>
. A Bayesian inference (BI) tree of
<emphasis id="DEBCEAD6FFC7706E1C3CED07FDAE75CC" box="[631,711,1909,1930]" italics="true" pageId="17" pageNumber="488">Petruca</emphasis>
<emphasis id="DEBCEAD6FFC7706E1C85ED06FC3C75CF" bold="true" box="[718,853,1908,1929]" pageId="17" pageNumber="488">subgen. nov.</emphasis>
and other related subgenera, based on the combined 16S rDNA, cytochrome oxidase subunit I genes (COI) and 28S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML* from RAxML and ML** from GARLI.
</paragraph>
</caption>
<paragraph id="EC7736C4FFC4706D1E8CEAEAFDBE7306" blockId="18.[151,1437,151,1149]" pageId="18" pageNumber="489">
One of the main characters placing these two species in
<taxonomicName id="2BC84D47FFC4706D1D0AEAEBFCFE72F6" box="[833,919,153,176]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1D0AEAEBFCFE72F6" box="[833,919,153,176]" italics="true" pageId="18" pageNumber="489">Minuca</emphasis>
</taxonomicName>
is the presence of two posterolateral striae on each side of carapace (cf. Beinlich &amp; von
<bibRefCitation id="88594B35FFC4706D1C36EACFFC637293" author="Hagen" box="[637,778,188,213]" pageId="18" pageNumber="489" refString="Beinlich, B. &amp; von Hagen, H. O. (2006) Materials for a more stable subdivision of the genus Uca Leach. Zoologische Mededelingen, 80, 9 - 32." type="journal article" year="2006">Hagen 2006</bibRefCitation>
;
<bibRefCitation id="88594B35FFC4706D1D5CEACFFCDD7292" author="Bezerra" box="[791,948,188,212]" pageId="18" pageNumber="489" refString="Bezerra, L. E. A. (2012) The fiddler crabs (Crustacea: Brachyura: Ocypodidae: genus Uca) of the South Atlantic Ocean. Nauplius, 20, 203 - 246. http: // dx. doi. org / 10.1590 / S 0104 - 64972012000200011" type="journal article" year="2012">Bezerra 2012</bibRefCitation>
). This character, however, can be variable. While there are two striae on each side of the carapace in
<taxonomicName id="2BC84D47FFC4706D1D7DEA93FCC372BE" box="[822,938,223,248]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="thayeri">
<emphasis id="DEBCEAD6FFC4706D1D7DEA93FCC372BE" box="[822,938,223,248]" italics="true" pageId="18" pageNumber="489">U. thayeri</emphasis>
</taxonomicName>
,
<taxonomicName id="2BC84D47FFC4706D1DF1EA93FB2572BE" box="[954,1100,223,248]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC4706D1DF1EA93FB2572BE" box="[954,1100,223,248]" italics="true" pageId="18" pageNumber="489">U. umbratila</emphasis>
</taxonomicName>
only has one stria (cf.
<bibRefCitation id="88594B35FFC4706D1B12EA92FFBA735A" author="Crane" pageId="18" pageNumber="489" refString="Crane, J. (1941) Crabs of the genus Uca from the west coast of Central America. Eastern Pacific Expeditions of the New York Zoological Society. XXVI. Zoologica, 26 (19), 145 - 208." type="journal article" year="1941">Crane 1941</bibRefCitation>
: text-fig. 7). This has not been discussed before. The available data also implies the morphological similarity between the two species and
<taxonomicName id="2BC84D47FFC4706D1F97EB5BFD5B7306" box="[476,562,297,320]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1F97EB5BFD5B7306" box="[476,562,297,320]" italics="true" pageId="18" pageNumber="489">Minuca</emphasis>
</taxonomicName>
is convergent.
</paragraph>
<paragraph id="EC7736C4FFC4706D1E8CEB3EFF8A717E" blockId="18.[151,1437,151,1149]" pageId="18" pageNumber="489">
The present study shows that
<taxonomicName id="2BC84D47FFC4706D1C58EB3FFDED7323" box="[531,644,332,357]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="thayeri">
<emphasis id="DEBCEAD6FFC4706D1C58EB3FFDED7323" box="[531,644,332,357]" italics="true" pageId="18" pageNumber="489">U. thayeri</emphasis>
</taxonomicName>
,
<taxonomicName id="2BC84D47FFC4706D1CD9EB3FFC4B7323" box="[658,802,332,357]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC4706D1CD9EB3FFC4B7323" box="[658,802,332,357]" italics="true" pageId="18" pageNumber="489">U. umbratila</emphasis>
</taxonomicName>
, and
<taxonomicName id="2BC84D47FFC4706D1D2BEB3FFC937322" box="[864,1018,333,356]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="spinicarpa">
<emphasis id="DEBCEAD6FFC4706D1D2BEB3FFC937322" box="[864,1018,333,356]" italics="true" pageId="18" pageNumber="489">U. spinicarpa</emphasis>
</taxonomicName>
form a highly supported clade which may be the result of allopatric speciation. Whereas
<taxonomicName id="2BC84D47FFC4706D1C97EB03FC2773CE" box="[732,846,367,392]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="thayeri">
<emphasis id="DEBCEAD6FFC4706D1C97EB03FC2773CE" box="[732,846,367,392]" italics="true" pageId="18" pageNumber="489">U. thayeri</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC4706D1DCDEB03FB7173CE" box="[902,1048,367,392]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="umbratila">
<emphasis id="DEBCEAD6FFC4706D1DCDEB03FB7173CE" box="[902,1048,367,392]" italics="true" pageId="18" pageNumber="489">U. umbratila</emphasis>
</taxonomicName>
diverged at 3.7 mya (p-distance = 4.34%), a well-known species pair separated after the closure of the Isthmus of
<collectingCountry id="94DF7654FFC4706D1A6EEBE6FB1773EA" box="[1061,1150,404,428]" name="Panama" pageId="18" pageNumber="489">Panamá</collectingCountry>
around 3.1 million years ago (reviewed by
<bibRefCitation id="88594B35FFC4706D1F2FEBC5FD247396" author="Schubart" box="[356,589,439,464]" pageId="18" pageNumber="489" refString="Schubart, C. D., Diesel, R. &amp; Hedges, S. B. (1998) Rapid evolution to terrestrial life in Jamaican crabs. Nature, 393, 363 - 365. http: // dx. doi. org / 10.1038 / 30724" type="journal article" year="1998">
Schubart
<emphasis id="DEBCEAD6FFC4706D1F9BEBCBFD637396" box="[464,522,439,464]" italics="true" pageId="18" pageNumber="489">et al.</emphasis>
1998
</bibRefCitation>
;
<bibRefCitation id="88594B35FFC4706D1C11EBC5FD9B7396" author="Lessios" box="[602,754,439,464]" pageId="18" pageNumber="489" refString="Lessios, H. A. (2008) The Great American Schism: Divergence of marine organisms after the rise of the Central American Isthmus. Annual Review of Ecology and Systematics, 39, 63 - 91. http: // dx. doi. org / 10.1146 / annurev. ecolsys. 38.091206.095815" type="journal article" year="2008">Lessios 2008</bibRefCitation>
),
<taxonomicName id="2BC84D47FFC4706D1D4CEBCBFCCA7396" box="[775,931,441,464]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="spinicarpa">
<emphasis id="DEBCEAD6FFC4706D1D4CEBCBFCCA7396" box="[775,931,441,464]" italics="true" pageId="18" pageNumber="489">U. spinicarpa</emphasis>
</taxonomicName>
is endemic to the Gulf of
<collectingCountry id="94DF7654FFC4706D1A99EBC5FA4E7396" box="[1234,1319,439,464]" name="Mexico" pageId="18" pageNumber="489">Mexico</collectingCountry>
in a more northern distribution, and it diverged from the other two species at 6.3 mya (p-distance = 7.38%).
<taxonomicName id="2BC84D47FFC4706D1B26EBAFFE59705E" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="subcylindrica">
<emphasis id="DEBCEAD6FFC4706D1B26EBAFFE59705E" italics="true" pageId="18" pageNumber="489">Uca subcylindrica</emphasis>
</taxonomicName>
is endemic along the border between Texas and northern
<collectingCountry id="94DF7654FFC4706D1DFAEB8DFB6F705E" box="[945,1030,511,536]" name="Mexico" pageId="18" pageNumber="489">Mexico</collectingCountry>
in the Gulf of
<collectingCountry id="94DF7654FFC4706D1AE2EB8DFA68705E" box="[1193,1281,511,536]" name="Mexico" pageId="18" pageNumber="489">Mexico</collectingCountry>
, and has been considered as an anomalous form, with unusually rounded body, striking form of G1 and hypersaline lagoonal habitat, without close alliance to other species (
<bibRefCitation id="88594B35FFC4706D1CF2E83AFC2B7026" author="Crane" box="[697,834,583,608]" pageId="18" pageNumber="489" refString="Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp." type="book" year="1975">Crane 1975</bibRefCitation>
;
<bibRefCitation id="88594B35FFC4706D1D1BE835FBE27026" author="Barnwell" box="[848,1163,583,608]" pageId="18" pageNumber="489" refString="Barnwell, F. H. &amp; Thurman, C. L. (1984) Taxonomy and biogeography of the fiddler crabs (Ocypodidae: genus Uca) of the Atlantic and Gulf coasts of eastern North America. Zoological Journal of the Linnean Society, 81, 23 - 87." type="journal article" year="1984">Barnwell &amp; Thurman 1984</bibRefCitation>
;
<bibRefCitation id="88594B35FFC4706D1AD3E835FA2D7026" author="Thurman" box="[1176,1348,583,608]" pageId="18" pageNumber="489" refString="Thurman, C. L. (1987) Fiddler crabs (genus Uca) of eastern Mexico (Decapoda, Brachyura, Ocypodidae). Crustaceana, 53, 94 - 105. http: // dx. doi. org / 10.1163 / 156854087 X 00664" type="journal article" year="1987">Thurman 1987</bibRefCitation>
).
<bibRefCitation id="88594B35FFC4706D1B12E83AFF8870C3" author="Crane" pageId="18" pageNumber="489" refString="Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp." type="book" year="1975">Crane (1975)</bibRefCitation>
placed it in
<taxonomicName id="2BC84D47FFC4706D1F26E81FFEAB70C2" box="[365,450,621,644]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1F26E81FFEAB70C2" box="[365,450,621,644]" italics="true" pageId="18" pageNumber="489">Minuca</emphasis>
</taxonomicName>
, but noted with reservation that it was close to
<taxonomicName id="2BC84D47FFC4706D1D92E81FFB5A70C2" box="[985,1075,621,644]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1D92E81FFB5A70C2" box="[985,1075,621,644]" italics="true" pageId="18" pageNumber="489">Leptuca</emphasis>
</taxonomicName>
. Because some characters of
<taxonomicName id="2BC84D47FFC4706D1BCAE81FFE5970EE" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="subcylindrica">
<emphasis id="DEBCEAD6FFC4706D1BCAE81FFE5970EE" italics="true" pageId="18" pageNumber="489">U. subcylindrica</emphasis>
</taxonomicName>
are shared with several members of
<taxonomicName id="2BC84D47FFC4706D1CA9E8E3FBF370EE" authority="Barnwell &amp; Thurman (1984)" authorityName="Barnwell &amp; Thurman" authorityYear="1984" box="[738,1178,655,680]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1CA9E8E3FC5570EE" box="[738,828,657,680]" italics="true" pageId="18" pageNumber="489">Leptuca</emphasis>
,
<bibRefCitation id="88594B35FFC4706D1D06E8FDFBF370EE" author="Barnwell" box="[845,1178,655,680]" pageId="18" pageNumber="489" refString="Barnwell, F. H. &amp; Thurman, C. L. (1984) Taxonomy and biogeography of the fiddler crabs (Ocypodidae: genus Uca) of the Atlantic and Gulf coasts of eastern North America. Zoological Journal of the Linnean Society, 81, 23 - 87." type="journal article" year="1984">Barnwell &amp; Thurman (1984)</bibRefCitation>
</taxonomicName>
considered it was not suitable to retain in the homogeneous
<taxonomicName id="2BC84D47FFC4706D1C09E8C7FDFE708A" box="[578,663,693,716]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1C09E8C7FDFE708A" box="[578,663,693,716]" italics="true" pageId="18" pageNumber="489">Minuca</emphasis>
</taxonomicName>
and transferred it to
<taxonomicName id="2BC84D47FFC4706D1DC8E8C7FCB4708A" box="[899,989,693,716]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1DC8E8C7FCB4708A" box="[899,989,693,716]" italics="true" pageId="18" pageNumber="489">Leptuca</emphasis>
</taxonomicName>
. Their action is supported by our study, as
<taxonomicName id="2BC84D47FFC4706D1EF1E8ABFE1370B6" box="[186,378,727,752]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="subcylindrica">
<emphasis id="DEBCEAD6FFC4706D1EF1E8ABFE1370B6" box="[186,378,727,752]" italics="true" pageId="18" pageNumber="489">U. subcylindrica</emphasis>
</taxonomicName>
forms a distinct clade from other species of this subgenus (
<figureCitation id="74F32A41FFC4706D1A03E8A5FBF870B6" box="[1096,1169,727,752]" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="18" pageNumber="489">Fig. 9</figureCitation>
). The simpler form of urocardiac ossicles of the gastric mill (
<figureCitation id="74F32A41FFC4706D1C10E88EFDC87152" box="[603,673,764,789]" captionStart="FIGURE 7" captionStartId="12.[151,250,1882,1904]" captionTargetBox="[176,1408,203,1846]" captionTargetId="figure@12.[165,1421,193,1861]" captionTargetPageId="12" captionText="FIGURE 7. Urocardiac ossicle of the subgenus Leptuca species. A, Uca subcylindrica (NCHUZOOL 13952, CW 15.2 mm, ♂); B, U. terpsichores (NCHUZOOL 13582, CW 9.9 mm, ♂); C, U. uruguayensis (NCHUZOOL 13577, CW 14.1 mm, ♂); D, U. spinicarpa (NCHUZOOL 13947, CW 13.3 mm, ♀); E, U. panacea (NCHUZOOL 13950, CW 14.8 mm, ♂); F, U. pugilator (NCHUZOOL 13586, CW 15.9 mm, ♂); G, U. speciosa (NCHUZOOL 13951, CW 12.6 mm, ♂); H, U. umbratila (NCHUZOOL 13579, CW 16.2 mm, ♂); I, U. thayeri (NCHUZOOL 13953, CW 19.0 mm, ♂). Scale = 1 mm." httpUri="https://zenodo.org/record/235580/files/figure.png" pageId="18" pageNumber="489">Fig. 7</figureCitation>
A) may also be a reflection of the early split of this species from others.
</paragraph>
</subSubSection>
<subSubSection id="A4D2654FFFC4706D1E8CE937FB1C763B" pageId="18" pageNumber="489" type="reference_group">
<paragraph id="EC7736C4FFC4706D1E8CE937FB1C763B" blockId="18.[151,1437,151,1149]" pageId="18" pageNumber="489">
<taxonomicName id="2BC84D47FFC4706D1E8CE937FEEA711A" ID-CoL="7Z6P4" box="[199,387,837,860]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="panamensis">
<emphasis id="DEBCEAD6FFC4706D1E8CE937FEEA711A" box="[199,387,837,860]" italics="true" pageId="18" pageNumber="489">Uca panamensis</emphasis>
</taxonomicName>
was placed in
<taxonomicName id="2BC84D47FFC4706D1C7FE937FDE3711A" ID-CoL="7PP9B" box="[564,650,837,860]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1C7FE937FDE3711A" box="[564,650,837,860]" italics="true" pageId="18" pageNumber="489">Minuca</emphasis>
</taxonomicName>
mainly by the characters of the front of the carapace and male first gonopod (
<bibRefCitation id="88594B35FFC4706D1F46E91AFEEE71C6" author="Bott" box="[269,391,871,896]" pageId="18" pageNumber="489" refString="Bott, R. (1954) Dekapoden (Crustacea) aus El Salvador. 1. Winkerkrabben (Uca). Senckenbergiana Biologica, 35 (3 / 4), 155 - 180." type="journal article" year="1954">Bott 1954</bibRefCitation>
,
<bibRefCitation id="88594B35FFC4706D1FDCE91AFEBB71C6" author="Bott" box="[407,466,872,896]" pageId="18" pageNumber="489" refString="Bott, R. (1973) Die verwandtschaftlichen Beziehungen der Uca Arten. Senckenbergiana Biologica, 54, 315 - 325." type="journal article" year="1973">1973</bibRefCitation>
;
<bibRefCitation id="88594B35FFC4706D1FAFE91AFD0671C6" author="Crane" box="[484,623,871,896]" pageId="18" pageNumber="489" refString="Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp." type="book" year="1975">Crane 1975</bibRefCitation>
). Based on a morphological analysis,
<bibRefCitation id="88594B35FFC4706D1A72E915FA6171C6" author="Rosenberg" box="[1081,1288,871,896]" pageId="18" pageNumber="489" refString="Rosenberg, M. S. (2001) The systematics and taxonomy of fiddler crabs: a phylogeny of the genus Uca. Journal of Crustacean Biology, 21, 839 - 869. http: // dx. doi. org / 10.1163 / 20021975 - 99990176" type="journal article" year="2001">Rosenberg (2001)</bibRefCitation>
placed it in
<taxonomicName id="2BC84D47FFC4706D1EDCE9FFFF9B71E2" ID-CoL="7PMPX" box="[151,242,909,932]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1EDCE9FFFF9B71E2" box="[151,242,909,932]" italics="true" pageId="18" pageNumber="489">Leptuca</emphasis>
</taxonomicName>
along with
<taxonomicName id="2BC84D47FFC4706D1F3DE9FFFD4871E3" ID-CoL="7DDTR" box="[374,545,908,933]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="terpsichores">
<emphasis id="DEBCEAD6FFC4706D1F3DE9FFFD4871E3" box="[374,545,908,933]" italics="true" pageId="18" pageNumber="489">U. terpsichores</emphasis>
</taxonomicName>
,
<taxonomicName id="2BC84D47FFC4706D1C7BE9FFFDBC71E3" ID-CoL="7DDQW" box="[560,725,908,933]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="deichmanni">
<emphasis id="DEBCEAD6FFC4706D1C7BE9FFFDBC71E3" box="[560,725,908,933]" italics="true" pageId="18" pageNumber="489">U. deichmanni</emphasis>
</taxonomicName>
,
<taxonomicName id="2BC84D47FFC4706D1CA8E9FFFCE671E3" ID-CoL="7DDR3" box="[739,911,908,933]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="leptodactyla">
<emphasis id="DEBCEAD6FFC4706D1CA8E9FFFCE671E3" box="[739,911,908,933]" italics="true" pageId="18" pageNumber="489">U. leptodactyla</emphasis>
</taxonomicName>
, and
<taxonomicName id="2BC84D47FFC4706D1D87E9FFFB3671E3" ID-CoL="84HGH" box="[972,1119,908,933]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="dorotheae">
<emphasis id="DEBCEAD6FFC4706D1D87E9FFFB3671E3" box="[972,1119,908,933]" italics="true" pageId="18" pageNumber="489">U. dorotheae</emphasis>
</taxonomicName>
. The many unusual features of
<taxonomicName id="2BC84D47FFC4706D1EFEE9C3FE35718E" ID-CoL="7Z6P4" box="[181,348,945,968]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="panamensis">
<emphasis id="DEBCEAD6FFC4706D1EFEE9C3FE35718E" box="[181,348,945,968]" italics="true" pageId="18" pageNumber="489">U. panamensis</emphasis>
</taxonomicName>
(e.g., flatter carapace, stiff setal brushes on the tips of the small chelae, the posterior extension of
<taxonomicName id="2BC84D47FFC4706D1EDCE9A6FFB171AB" ID-CoL="84GD3" box="[151,216,980,1005]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="major">major</taxonomicName>
manus, preference for rocky habitats etc.) suggest this classification is incorrect. The present use of new morphological characters (viz. the orbital armature at the inner corner of orbital floor, simple form of urocardiac ossicles of gastric mill, large and bent inner process of the G1) as well as molecular evidence using mitochondrial and nuclear markers (
<figureCitation id="74F32A41FFC4706D1FD8EE4DFEB3761E" box="[403,474,1087,1112]" captionStart="FIGURE 9. A" captionStartId="17.[151,250,1908,1930]" captionTargetBox="[160,1392,213,1872]" captionTargetId="figure@17.[151,1436,193,1886]" captionTargetPageId="17" captionText="FIGURE 9. A Bayesian inference (BI) tree of Petruca subgen. nov. and other related subgenera, based on the combined 16 S rDNA, cytochrome oxidase subunit I genes (COI) and 28 S rDNA. See Table 1 for details of the specimens. Probability values at the nodes represent support values for BI, as well as two methods of maximum likelihood (ML), with ML * from RAxML and ML ** from GARLI." httpUri="https://zenodo.org/record/235582/files/figure.png" pageId="18" pageNumber="489">Fig. 9</figureCitation>
), support the hypothesis that
<taxonomicName id="2BC84D47FFC4706D1D7FEE33FCB4761E" ID-CoL="7Z6P4" box="[820,989,1089,1112]" class="Malacostraca" family="Ocypodidae" genus="Uca" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="species" species="panamensis">
<emphasis id="DEBCEAD6FFC4706D1D7FEE33FCB4761E" box="[820,989,1089,1112]" italics="true" pageId="18" pageNumber="489">U. panamensis</emphasis>
</taxonomicName>
belongs to its own subgenus (
<emphasis id="DEBCEAD6FFC4706D1B08EE33FAF5761E" box="[1347,1436,1089,1112]" italics="true" pageId="18" pageNumber="489">Petruca</emphasis>
<emphasis id="DEBCEAD6FFC4706D1EDCEE16FE43763B" bold="true" box="[151,298,1124,1149]" pageId="18" pageNumber="489">subgen. nov.</emphasis>
), and diverged from other American
<taxonomicName id="2BC84D47FFC4706D1C87EE17FC4B763A" ID-CoL="7PP9B" box="[716,802,1125,1148]" class="Malacostraca" family="Ocypodidae" genus="Minuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1C87EE17FC4B763A" box="[716,802,1125,1148]" italics="true" pageId="18" pageNumber="489">Minuca</emphasis>
</taxonomicName>
and
<taxonomicName id="2BC84D47FFC4706D1D13EE17FCDA763A" ID-CoL="7PMPX" box="[856,947,1125,1148]" class="Malacostraca" family="Ocypodidae" genus="Leptuca" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="18" pageNumber="489" phylum="Arthropoda" rank="genus">
<emphasis id="DEBCEAD6FFC4706D1D13EE17FCDA763A" box="[856,947,1125,1148]" italics="true" pageId="18" pageNumber="489">Leptuca</emphasis>
</taxonomicName>
at about 14 mya.
</paragraph>
</subSubSection>
</treatment>
</document>