190 lines
23 KiB
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190 lines
23 KiB
XML
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<mods:title id="3EC245BF585703E55611853115949248">Phylogenetic relationships and biogeographical history of the genus Rhinoclemmys Fitzinger, 1835 and the monophyly of the turtle family Geoemydidae (Testudines: Testudinoidea)</mods:title>
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<mods:namePart id="3045F4F898CFB2AB31405866E38F668D">Le, Minh</mods:namePart>
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<mods:namePart id="03F917822BEE88E26DC7659D3CAD18CC">Mccord, William P.</mods:namePart>
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<mods:date id="198B91E94810336945C4C774217FD14F">2008</mods:date>
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<subSubSection id="04C26546FF93FFE21D6D02DEFAFEFF14" box="[849,1378,198,220]" pageId="12" pageNumber="763" type="nomenclature">
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<paragraph id="4C6736CDFF93FFE21D6D02DEFAFEFF14" blockId="12.[849,1378,198,220]" box="[849,1378,198,220]" pageId="12" pageNumber="763">
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<heading id="172F81A1FF93FFE21D6D02DEFAFEFF14" box="[849,1378,198,220]" centered="true" fontSize="9" level="2" pageId="12" pageNumber="763" reason="2">
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RELATIONSHIP BETWEEN
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<taxonomicName id="8BD84D4EFF93FFE21A4802DFFAFEFF14" authority="AND" box="[1140,1378,199,220]" class="Reptilia" family="Geoemydidae" genus="Rhinoclemmys" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21A4802DFFAB6FF15" box="[1140,1322,199,220]" italics="true" pageId="12" pageNumber="763">RHINOCLEMMYS</emphasis>
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AND
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</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="04C26546FF93FFE31D5202F1FEBCFC91" lastPageId="13" lastPageNumber="764" pageId="12" pageNumber="763" type="description">
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<paragraph id="4C6736CDFF93FFE21D5202F1FB69FED5" blockId="12.[878,1349,233,283]" pageId="12" pageNumber="763">
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OTHER GENERA AND THE MONOPHYLY OF THE FAMILY
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<taxonomicName id="8BD84D4EFF93FFE21A6C031EFB69FED5" baseAuthorityName="Theobald" baseAuthorityYear="1868" box="[1104,1269,262,283]" class="Reptilia" family="Geoemydidae" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="family">GEOEMYDIDAE</taxonomicName>
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</paragraph>
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<paragraph id="4C6736CDFF93FFE21D1A0358FA12FBFA" blockId="12.[806,1422,320,1903]" pageId="12" pageNumber="763">
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The results from the present study strongly support the monophyly of the family
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<taxonomicName id="8BD84D4EFF93FFE21A4A0347FA93FEBA" baseAuthorityName="Theobald" baseAuthorityYear="1868" box="[1142,1295,351,373]" class="Reptilia" family="Geoemydidae" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="family">Geoemydidae</taxonomicName>
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(BP = 85% in MP, BP = 71% in ML; PP = 98%) inclusive of
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<taxonomicName id="8BD84D4EFF93FFE21B670365FC3EFE7E" class="Reptilia" family="Geoemydidae" genus="Rhinoclemmys" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21B670365FC3EFE7E" italics="true" pageId="12" pageNumber="763">Rhinoclemmys</emphasis>
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</taxonomicName>
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. This is the first broad-sampling molecular analysis that strongly supports the monophyly of the family with regard to testudinids, and also supports the subfamilial status of
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<taxonomicName id="8BD84D4EFF93FFE21A6603E0FA9CFDC2" box="[1114,1280,504,525]" class="Reptilia" family="Geoemydidae" genus="Rhinoclemmys" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21A6603E0FA9CFDC2" box="[1114,1280,504,525]" italics="true" pageId="12" pageNumber="763">Rhinoclemmys</emphasis>
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</taxonomicName>
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. We herein propose to raise this genus to subfamily rank with the name of Rhinoclemminae. Morphologically, all species of the genus share at least two synapomorphies, the absence of lateral keels throughout their life (
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<bibRefCitation id="28494B3CFF93FFE21B00006AFC5DFD68" author="Claude J & Tong H" pageId="12" pageNumber="763" pagination="3 - 45" refId="ref9652" refString="Claude J, Tong H. 2004. Early Eocene testudinoid turtles from Saint Papoul, France, with comments on the early evolution of modern Testudinoidea. Oryctos 5: 3 - 45." type="journal article" year="2004">Claude & Tong, 2004</bibRefCitation>
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) and the shape of the upper triturating surface (our pers. observ.). Examination of
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<specimenCount id="5ADEFD44FF93FFE21B1A00B7FCFFFD2C" pageId="12" pageNumber="763" type="generic">60 specimens</specimenCount>
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of all species in this genus and
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<specimenCount id="5ADEFD44FF93FFE21ACD00D6FA11FD2B" box="[1265,1421,718,740]" pageId="12" pageNumber="763" type="generic">63 specimens</specimenCount>
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of other species (see supplementary Appendix S2) belonging to other major clades of the family reveals that the upper triturating surface in this group is different from that of other geoemydid species in that it is narrower in the anterior portion and expanded in the posterior potion. In addition, the upper triturating surface has a minimal lingual ridge on the inner rim.
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<bibRefCitation id="28494B3CFF93FFE21D5C01DBFC74FC16" author="Carr JL" box="[864,1000,963,985]" pageId="12" pageNumber="763" refId="ref9469" refString="Carr JL. 1991. Phylogenetic analysis of the neotropical turtle genus Rhinoclemmys Fitzinger (Testudines: Emydidae). DPhil. Thesis, Southern Illinois University." type="book" year="1991">Carr (1991)</bibRefCitation>
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proposed other synapomorphies for this group, but these characters either vary among other geoemydids or could not be checked because they are either karyotypic or biochemical characters.
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</paragraph>
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<paragraph id="4C6736CDFF93FFE21D030625FA8CF919" blockId="12.[806,1422,320,1903]" pageId="12" pageNumber="763">
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Although the other major clade of geoemydids, exclusive of
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<taxonomicName id="8BD84D4EFF93FFE21D890644FBC7FBBE" box="[949,1115,1116,1137]" class="Reptilia" family="Geoemydidae" genus="Rhinoclemmys" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21D890644FBC7FBBE" box="[949,1115,1116,1137]" italics="true" pageId="12" pageNumber="763">Rhinoclemmys</emphasis>
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</taxonomicName>
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, is not strongly supported by the MP analysis, it is consistently recovered in all of our analyses and received strong support from the Bayesian and ML analyses. Biogeographically, it is a distinct clade containing mostly Asian taxa. Within this clade, our MP results show the same topology as the one recovered by
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<bibRefCitation id="28494B3CFF93FFE21AA6070BFA16FAE6" author="Diesmos AC & Parham JF & Stuart BL & Brown RM" box="[1178,1418,1299,1321]" pageId="12" pageNumber="763" pagination="31 - 41" refId="ref9795" refString="Diesmos AC, Parham JF, Stuart BL, Brown RM. 2005. The phylogenetic position of the recently rediscovered Philippine forest turtle (Bataguridae: Heosemys leytensis). Proceedings of the California Academy of Sciences 56: 31 - 41." type="journal article" year="2005">
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Diesmos
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<emphasis id="7EACEADFFF93FFE21B3D070CFAA5FAE7" box="[1281,1337,1299,1321]" italics="true" pageId="12" pageNumber="763">et al.</emphasis>
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(2005)
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</bibRefCitation>
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. However, several basal nodes have significantly higher BP values, such as the clade consisting of
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<taxonomicName id="8BD84D4EFF93FFE21D1A0768FCF0FA4A" box="[806,876,1392,1413]" class="Reptilia" family="Geoemydidae" genus="Cuora" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21D1A0768FCF0FA4A" box="[806,876,1392,1413]" italics="true" pageId="12" pageNumber="763">Cuora</emphasis>
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</taxonomicName>
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+
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<taxonomicName id="8BD84D4EFF93FFE21DB70768FB9EFA4A" box="[907,1026,1392,1413]" class="Reptilia" family="Geoemydidae" genus="Mauremys" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21DB70768FB9EFA4A" box="[907,1026,1392,1413]" italics="true" pageId="12" pageNumber="763">Mauremys</emphasis>
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</taxonomicName>
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+
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<taxonomicName id="8BD84D4EFF93FFE21A1D0777FB16FA4B" authorityName="Bell" authorityYear="1834" box="[1057,1162,1391,1412]" class="Reptilia" family="Geoemydidae" genus="Cyclemys" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21A1D0777FB16FA4B" box="[1057,1162,1391,1412]" italics="true" pageId="12" pageNumber="763">Cyclemys</emphasis>
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</taxonomicName>
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+
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<taxonomicName id="8BD84D4EFF93FFE21A940768FA8BFA4A" box="[1192,1303,1392,1413]" class="Reptilia" family="Geoemydidae" genus="Heosemys" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21A940768FA8BFA4A" box="[1192,1303,1392,1413]" italics="true" pageId="12" pageNumber="763">Heosemys</emphasis>
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</taxonomicName>
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+
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<taxonomicName id="8BD84D4EFF93FFE21B090777FA17FA4B" authorityName="Gray" authorityYear="1870" box="[1333,1419,1391,1412]" class="Reptilia" family="Geoemydidae" genus="Sacalia" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21B090777FA17FA4B" box="[1333,1419,1391,1412]" italics="true" pageId="12" pageNumber="763">Sacalia</emphasis>
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</taxonomicName>
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(BP = 90% vs. 57%) and the clade of
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<taxonomicName id="8BD84D4EFF93FFE21B330796FAE4FA6C" authorityName="Bell" authorityYear="1834" box="[1295,1400,1422,1443]" class="Reptilia" family="Geoemydidae" genus="Cyclemys" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21B330796FAE4FA6C" box="[1295,1400,1422,1443]" italics="true" pageId="12" pageNumber="763">Cyclemys</emphasis>
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</taxonomicName>
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+
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<taxonomicName id="8BD84D4EFF93FFE21D1A07B5FC09FA0D" box="[806,917,1453,1474]" class="Reptilia" family="Geoemydidae" genus="Heosemys" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21D1A07B5FC09FA0D" box="[806,917,1453,1474]" italics="true" pageId="12" pageNumber="763">Heosemys</emphasis>
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</taxonomicName>
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+
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<taxonomicName id="8BD84D4EFF93FFE21D8C07B4FB9AFA0E" authorityName="Gray" authorityYear="1870" box="[944,1030,1452,1473]" class="Reptilia" family="Geoemydidae" genus="Sacalia" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21D8C07B4FB9AFA0E" box="[944,1030,1452,1473]" italics="true" pageId="12" pageNumber="763">Sacalia</emphasis>
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</taxonomicName>
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(BP = 97% vs. 74%). The topology resulting from our Bayesian and ML analyses for this major clade is the same as one proposed by
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<bibRefCitation id="28494B3CFF93FFE21B0307F2FC2EF9D1" author="Spinks PQ & Shaffer HB & Iverson JB & McCord WP" pageId="12" pageNumber="763" pagination="164 - 182" refId="ref11701" refString="Spinks PQ, Shaffer HB, Iverson JB, McCord WP. 2004. Phylogenetic hypotheses for the turtle family Geoemydidae. Molecular Phylogenetics and Evolution 32: 164 - 182." type="journal article" year="2004">
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Spinks
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<emphasis id="7EACEADFFF93FFE21D1A0411FCC2F9D2" box="[806,862,1544,1566]" italics="true" pageId="12" pageNumber="763">et al.</emphasis>
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(2004)
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</bibRefCitation>
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, but the support values in all nodes are generally higher. The most significant discrepancy between our MP and the Bayesian and ML analyses is the position of
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<taxonomicName id="8BD84D4EFF93FFE21DF5047CFBA2F9B6" authorityName="Gray" authorityYear="1834" box="[969,1086,1636,1657]" class="Reptilia" family="Geoemydidae" genus="Geoemyda" kingdom="Animalia" order="Testudines" pageId="12" pageNumber="763" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF93FFE21DF5047CFBA2F9B6" box="[969,1086,1636,1657]" italics="true" pageId="12" pageNumber="763">Geoemyda</emphasis>
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</taxonomicName>
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. It is likely, however, that this problem could be eliminated by increasing taxon sampling. We are currently investigating this problem using more taxa and more molecular data.
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</paragraph>
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<paragraph id="4C6736CDFF93FFE31D0304C7FEBCFC91" blockId="12.[806,1422,320,1903]" lastBlockId="13.[164,779,198,862]" lastPageId="13" lastPageNumber="764" pageId="12" pageNumber="763">
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In terms of the synapomorphy of this family, we agree with
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<bibRefCitation id="28494B3CFF93FFE21D8604E5FB19F8DC" author="Hirayama R" box="[954,1157,1789,1811]" pageId="12" pageNumber="763" pagination="141 - 157" refId="ref10415" refString="Hirayama R. 1984. Cladistic analysis of batagurine turtles (Batagurinae: Emydidae: Testudinidae); a preliminary result. Studia Geologica Salmanticensia 1: 141 - 157." type="journal article" year="1984">Hirayama (1984)</bibRefCitation>
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and
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<bibRefCitation id="28494B3CFF93FFE21AED04E5FCF2F8FD" author="Yasukawa Y & Hirayama R & Hikida T" pageId="12" pageNumber="763" pagination="105 - 133" refId="ref12148" refString="Yasukawa Y, Hirayama R, Hikida T. 2001. Phylogenetic relationships of geoemydine turtles (Reptilia: Bataguridae). Current Herpetology 20: 105 - 133." type="journal article" year="2001">
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Yasukawa
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<emphasis id="7EACEADFFF93FFE21B6904E6FA11F8DD" box="[1365,1421,1789,1811]" italics="true" pageId="12" pageNumber="763">et al.</emphasis>
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(2001)
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</bibRefCitation>
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that the presence of inguinal and axillary musk duct foramina is the character uniting all geoemydids. This hypothesis has been criticised as musk duct foramina are also present in other groups of turtles (
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<bibRefCitation id="28494B3CFF92FFE31F3D02FDFE39FF35" author="Waagen GN" box="[257,421,229,250]" pageId="13" pageNumber="764" refId="ref11994" refString="Waagen GN. 1972. Musk glands in recent turtles. MS Thesis, University of Utah." type="book" year="1972">Waagen, 1972</bibRefCitation>
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;
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<bibRefCitation id="28494B3CFF92FFE31F8F02FCFC9AFF35" author="Ehrenfeld JG & Ehrenfeld DW" box="[435,774,228,250]" pageId="13" pageNumber="764" pagination="305 - 314" refId="ref9980" refString="Ehrenfeld JG, Ehrenfeld DW. 1973. Externally secreting glands of freshwater and sea turtles. Copeia 1973: 305 - 314." type="journal article" year="1973">Ehrenfeld & Ehrenfeld, 1973</bibRefCitation>
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;
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<bibRefCitation id="28494B3CFF92FFE31E98031BFE52FED7" author="Gaffney E & Meylan PA" box="[164,462,259,281]" pageId="13" pageNumber="764" pagination="157 - 219" refId="ref10188" refString="Gaffney E, Meylan PA. 1988. A phylogeny of turtles. In: Benton MJ, ed. The phylogeny and classification of the tetrapods, vol. 1: Amphibian, reptiles, birds systematics association special volume no 35 A. Oxford: Clarendon Press, 157 - 219." type="book chapter" year="1988">Gaffney & Meylan, 1988</bibRefCitation>
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;
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<bibRefCitation id="28494B3CFF92FFE31FDC031BFC9AFED7" author="Weldon PJ & Gaffney ES" box="[480,774,259,281]" pageId="13" pageNumber="764" pagination="556 - 557" refId="ref12051" refString="Weldon PJ, Gaffney ES. 1998. An ancient integumentary gland in turtles. Naturwissenschaften 85: 556 - 557." type="journal article" year="1998">Weldon & Gaffney, 1998</bibRefCitation>
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;
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<bibRefCitation id="28494B3CFF92FFE31E98033AFE0FFEF7" author="Joyce WJ & Bell CJ" box="[164,403,290,312]" pageId="13" pageNumber="764" pagination="53 - 109" refId="ref10763" refString="Joyce WJ, Bell CJ. 2004. A review of the comparative morphology of testudinoid turtles (Reptilia: Testudines). Asiatic Herpetological Research 10: 53 - 109." type="journal article" year="2004">Joyce & Bell, 2004</bibRefCitation>
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). Nevertheless, the musk duct positions and formation can be used to differentiate geoemydids from emydids and testudinids. Within testudinoids, emydids tend to have only one pair of musk duct foramina in the axillary region and testudinids do not possess these foramina. Geoemydids commonly have two pairs, in the axillary and inguinal buttresses. The genus
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<taxonomicName id="8BD84D4EFF92FFE31F9603E0FD95FDC2" box="[426,521,504,525]" class="Reptilia" family="Geoemydidae" genus="Morenia" kingdom="Animalia" order="Testudines" pageId="13" pageNumber="764" phylum="Chordata" rank="genus">
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<emphasis id="7EACEADFFF92FFE31F9603E0FD95FDC2" box="[426,521,504,525]" italics="true" pageId="13" pageNumber="764">Morenia</emphasis>
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</taxonomicName>
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used to be considered the only geoemydid genus that does not possess musk duct foramina (
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<bibRefCitation id="28494B3CFF92FFE31F6B002DFE67FD85" author="Waagen GN" box="[343,507,565,586]" pageId="13" pageNumber="764" refId="ref11994" refString="Waagen GN. 1972. Musk glands in recent turtles. MS Thesis, University of Utah." type="book" year="1972">Waagen, 1972</bibRefCitation>
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;
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<bibRefCitation id="28494B3CFF92FFE31C34002DFC9AFD85" author="Yasukawa Y & Hirayama R & Hikida T" box="[520,774,565,587]" pageId="13" pageNumber="764" pagination="105 - 133" refId="ref12148" refString="Yasukawa Y, Hirayama R, Hikida T. 2001. Phylogenetic relationships of geoemydine turtles (Reptilia: Bataguridae). Current Herpetology 20: 105 - 133." type="journal article" year="2001">
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Yasukawa
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<emphasis id="7EACEADFFF92FFE31CB9002EFD21FD85" box="[645,701,565,587]" italics="true" pageId="13" pageNumber="764">et al.</emphasis>
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, 2001
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</bibRefCitation>
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;
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<bibRefCitation id="28494B3CFF92FFE31E98004CFEE1FDA5" author="Joyce WJ & Bell CJ" box="[164,381,596,618]" pageId="13" pageNumber="764" pagination="53 - 109" refId="ref10763" refString="Joyce WJ, Bell CJ. 2004. A review of the comparative morphology of testudinoid turtles (Reptilia: Testudines). Asiatic Herpetological Research 10: 53 - 109." type="journal article" year="2004">Joyce & Bell, 2004</bibRefCitation>
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), but our observation of a
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<taxonomicName id="8BD84D4EFF92FFE31C90004CFF67FD48" baseAuthorityName="Dumeril & Bibron" baseAuthorityYear="1835" class="Reptilia" family="Geoemydidae" genus="Morenia" kingdom="Animalia" order="Testudines" pageId="13" pageNumber="764" phylum="Chordata" rank="species" species="ocellata">
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<emphasis id="7EACEADFFF92FFE31C90004CFF67FD48" italics="true" pageId="13" pageNumber="764">Morenia ocellata</emphasis>
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</taxonomicName>
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specimen (see supplementary Appendix S2) revealed that they do have two pairs of small musk duct foramina as do other geoemydids.
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<bibRefCitation id="28494B3CFF92FFE31CAB00B7FEB0FD2B" author="Yasukawa Y & Hirayama R & Hikida T" pageId="13" pageNumber="764" pagination="105 - 133" refId="ref12148" refString="Yasukawa Y, Hirayama R, Hikida T. 2001. Phylogenetic relationships of geoemydine turtles (Reptilia: Bataguridae). Current Herpetology 20: 105 - 133." type="journal article" year="2001">
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Yasukawa
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<emphasis id="7EACEADFFF92FFE31E9800D7FF40FD2C" box="[164,220,718,740]" italics="true" pageId="13" pageNumber="764">et al.</emphasis>
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(2001)
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</bibRefCitation>
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also proposed that expanded iliac blades distinguish the family
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<taxonomicName id="8BD84D4EFF92FFE31F8600F5FDCFFCCC" baseAuthorityName="Theobald" baseAuthorityYear="1868" box="[442,595,749,771]" class="Reptilia" family="Geoemydidae" kingdom="Animalia" order="Testudines" pageId="13" pageNumber="764" phylum="Chordata" rank="family">Geoemydidae</taxonomicName>
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from all other families, but we were unable to check this character due to the rarity of complete iliac blades in examined specimens.
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |