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<document id="0A06D6C4A79EAF7C68717AC36371FC21" ID-ISSN="1175-5326" IM.materialsCitations_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1629634115413" checkinUser="felipe" docAuthor="Adalsteinsson, Solny A., Branch, William R., Trape, Sébastien, Vitt, Laurie J. &amp; Hedges, S. Blair" docDate="2009" docId="0E2487E3FF89FFA5FF0E30D0FEF5FC8A" docLanguage="en" docName="zt02244p050.pdf" docOrigin="Zootaxa 2244" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Tetracheilostomina Hedges, Adalsteinsson, &amp; Branch, New Subtribe" docType="treatment" docVersion="3" lastPageNumber="22" masterDocId="F21DFF9BFF9CFFB0FF993047FF95FFD5" masterDocTitle="Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squamata)" masterLastPageNumber="50" masterPageNumber="1" pageNumber="22" updateTime="1699087815032" updateUser="plazi" zenodo-license-document="CLOSED">
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<mods:title id="3AE753CBF3E17503861A58D1024D7605">Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squamata)</mods:title>
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<mods:namePart id="74EB4371A7D23818FB06796C03145C94">Adalsteinsson, Solny A.</mods:namePart>
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<mods:namePart id="CFFF8D99DCE21BD94911CDDD92E5E7AF">Branch, William R.</mods:namePart>
<mods:affiliation id="2BCDC82D68894EFF18CD8E4263A266B0">Bayworld, P. O. Box 13147, Humewood 6013, South Africa</mods:affiliation>
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<mods:namePart id="9031CFC86D29C2267755656D78981101">Trape, Sébastien</mods:namePart>
<mods:affiliation id="AAE6B4058CA91EAFF872B372FD5CA6CA">Laboratoire ECOLAG, UMR 5119, Université Montpellier II, cc 093, Place E. Bataillon, 34095 Montpellier Cedex 5, France</mods:affiliation>
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<mods:namePart id="AF83729DD2A2D008FC6BA655DFF2A533">Vitt, Laurie J.</mods:namePart>
<mods:affiliation id="7274A93579A32F5289EAC688A4C7ED2F">Sam Noble Oklahoma Museum of Natural History and Zoology Department, 2401 Chautauqua Avenue, Norman, OK 73072, USA Corresponding author. E-mail: sbh 1 @ psu. edu</mods:affiliation>
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<mods:namePart id="6515E25CE03B70987D27EFFCD1F449EF">Hedges, S. Blair</mods:namePart>
<mods:affiliation id="7DBE732760B8DE2CD23E9D7CBA687D96">Department of Biology, 208 Mueller Lab, Pennsylvania State University, University Park, PA 16802 - 5301 USA.</mods:affiliation>
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<mods:date id="A07F89E5F229E4C06DC47BF767D6DBA4">2009</mods:date>
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<subSubSection id="CE97657EFF89FFA5FF0E30D0FBCBFF64" box="[151,1118,151,177]" pageId="21" pageNumber="22" type="nomenclature">
<paragraph id="863236F5FF89FFA5FF0E30D0FBCBFF64" blockId="21.[151,1118,151,177]" box="[151,1118,151,177]" pageId="21" pageNumber="22">
<heading id="DD7A8199FF89FFA5FF0E30D0FBCBFF64" bold="true" box="[151,1118,151,177]" fontSize="11" level="1" pageId="21" pageNumber="22" reason="1">
<emphasis id="B4F9EAE7FF89FFA5FF0E30D0FBCBFF64" bold="true" box="[151,1118,151,177]" pageId="21" pageNumber="22">
Subtribe
<taxonomicName id="418D4D76FF89FFA5FE9230D0FC30FF64" authority="Hedges, Adalsteinsson, &amp; Branch" authorityName="Hedges, Adalsteinsson, &amp; Branch" box="[267,933,151,177]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="22" phylum="Chordata" rank="subTribe" status="subtrib. nov." subTribe="Tetracheilostomina">Tetracheilostomina Hedges, Adalsteinsson, &amp; Branch</taxonomicName>
,
<taxonomicNameLabel id="AFCA579CFF89FFA5FC2B30D0FBCBFF64" box="[946,1118,151,177]" pageId="21" pageNumber="22" rank="subTribe">New Subtribe</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="CE97657EFF89FFA5FF0E30A6FD13FF2D" box="[151,646,225,249]" pageId="21" pageNumber="22" type="type_taxon">
<paragraph id="863236F5FF89FFA5FF0E30A6FD13FF2D" blockId="21.[151,646,225,249]" box="[151,646,225,249]" pageId="21" pageNumber="22">
<emphasis id="B4F9EAE7FF89FFA5FF0E30A6FE88FF2C" bold="true" box="[151,285,225,249]" pageId="21" pageNumber="22">
<typeStatus id="59368857FF89FFA5FF0E30A6FF44FF2C" box="[151,209,225,249]" pageId="21" pageNumber="22">Type</typeStatus>
genus.
</emphasis>
<taxonomicName id="418D4D76FF89FFA5FEBD30A6FD17FF2D" authority="Jan, 1861: 191" authorityName="Jan" authorityPageNumber="191" authorityYear="1861" box="[292,642,225,248]" class="Reptilia" family="Leptotyphlopidae" genus="Tetracheilostoma" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF89FFA5FEBD30A6FE49FF2D" box="[292,476,225,248]" italics="true" pageId="21" pageNumber="22">Tetracheilostoma</emphasis>
<bibRefCitation id="E21C4B04FF89FFA5FE7A30A6FD17FF2D" author="Jan" box="[483,642,225,248]" firstAuthor="Jan" page="191" pageId="21" pageNumber="22" pagination="178 - 199" refId="ref25178" refString="Jan, G. (1861) Note sulla famiglia dei tiflopidi sui loro generi e sulle specie del genere Stenostoma. Archivio Per La Zoologia, L'Anatomia E La Fisiologia, 1, 178 - 199." type="journal article" year="1861">Jan, 1861: 191</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF89FFA5FF0E3160FCF0FE63" pageId="21" pageNumber="22" type="diagnosis">
<paragraph id="863236F5FF89FFA5FF0E3160FA09FE5A" blockId="21.[151,1436,295,863]" pageId="21" pageNumber="22">
<emphasis id="B4F9EAE7FF89FFA5FF0E3160FE82FE94" bold="true" box="[151,279,295,321]" pageId="21" pageNumber="22">Diagnosis.</emphasis>
<taxonomicName id="418D4D76FF89FFA5FE84316FFE68FE97" box="[285,509,296,322]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="2" phylum="Chordata" rank="subTribe" status="subtrib. nov." subTribe="Tetracheilostomina">Tetracheilostomina</taxonomicName>
is distinguished from the other two subtribes of
<taxonomicName id="418D4D76FF89FFA5FBA8316FFB1BFE97" box="[1073,1166,296,322]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="22" phylum="Chordata" rank="tribe" tribe="Epictini">Epictini</taxonomicName>
by usually having four supralabials (two in Epictina and
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<taxonomicName id="418D4D76FF89FFA5FD133109FD74FEBD" box="[650,737,334,360]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="2" phylum="Chordata" rank="subTribe" status="subtrib. nov." subTribe="Renina">Renina</taxonomicName>
) (Table 2). The support for this group was 100% BP and 100% PP for the four-gene tree (
<figureCitation id="1EB62A70FF89FFA5FD963132FDC3FE5A" box="[527,598,373,399]" captionStart="FIGURE 3" captionStartId="6.[151,255,1684,1708]" captionTargetBox="[181,1406,188,1658]" captionTargetId="figure-173@6.[179,1407,184,1660]" captionTargetPageId="6" captionText="FIGURE 3. A phylogeny of leptotyphlopid snakes based on sequences of four mitochondrial genes (12S rRNA, tRNA- Valine, 16S rRNA, and cytochrome b). Maximum likelihood tree of 91 samples and 2,971 sites. Values are ML bootstrap values followed by Bayesian posterior probabilities. Outgroups are not shown, but included Typhlopidae (Ramphotyphlops), Boidae (Boa), Pythonidae (Python), and Elapidae (Dendroaspis and Naja). The generic taxonomy in this tree reflects usage prior to this study. See Table 1 and Figure 12 for the new classification proposed here." pageId="21" pageNumber="22">Fig. 3</figureCitation>
); only one of the two genera was included in the nine-gene tree (
<figureCitation id="1EB62A70FF89FFA5FADF3132FA1BFE5A" box="[1350,1422,373,399]" captionStart="FIGURE 4" captionStartId="7.[151,260,1282,1306]" captionTargetBox="[252,1334,313,1258]" captionTargetId="figure-366@7.[252,1335,310,1258]" captionTargetPageId="7" captionText="FIGURE 4. A phylogeny of leptotyphlopid snakes based on sequences of nine genes: five nuclear genes nine (amelogenin, BDNF, C-mos, NT3, and RAG1) and four mitochondrial and nuclear genes (12S rRNA, tRNA-Valine, 16S rRNA, and cytochrome b). Maximum likelihood tree obtained from the nine-gene data set (24 species; 5,563 sites). Values are ML bootstrap values followed by Bayesian posterior probabilities. Outgroups are not shown, but included Typhlopidae (Ramphotyphlops), Boidae (Boa), Pythonidae (Python), and Elapidae (Dendroaspis and Naja). The generic taxonomy in this tree reflects usage prior to this study. See Table 1 and Figure 12 for the new classification proposed here." pageId="21" pageNumber="22">Fig. 4</figureCitation>
).
</paragraph>
<paragraph id="863236F5FF89FFA5FF5C31DCFCF0FE63" blockId="21.[151,1436,295,863]" box="[197,869,411,438]" pageId="21" pageNumber="22">
<emphasis id="B4F9EAE7FF89FFA5FF5C31DCFEA4FE60" bold="true" box="[197,305,411,437]" pageId="21" pageNumber="22">Content.</emphasis>
Two genera and seven species (
<tableCitation id="CB0F034EFF89FFA5FD3131DBFC94FE63" box="[680,769,412,438]" captionStart="TABLE 1" captionStartId="9.[151,240,151,175]" captionText="TABLE 1. Classification of snakes of the Family Leptotyphlopidae. The arrangement used in this study is compared with that in previous classifications (e.g., McDiarmid et al. 1999; Uetz et al. 2009). Abbreviations for geographic regions are: AR (Arabia), CAF (Central Africa), EAF (East Africa), MAM (Middle America), NAM (North America), SAF (South Africa), SAM (South America), SOC (Socotra Island), SWA (Southwest Asia), WAF (West Africa), and WI (West Indies). Species in bold were sampled in the molecular analyses. Undescribed species used in this study are not listed." pageId="21" pageNumber="22">Table 1</tableCitation>
;
<figureCitation id="1EB62A70FF89FFA5FC8931DBFCC2FE63" box="[784,855,412,438]" captionStart="FIGURE 7" captionStartId="20.[151,255,1611,1635]" captionTargetBox="[151,1436,197,1579]" captionTargetId="figure-117@20.[151,1436,197,1579]" captionTargetPageId="20" captionText="FIGURE 7. Representatives of the snake Family Leptotyphlopidae from the New World (continued)." pageId="21" pageNumber="22">Fig. 7</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF89FFA5FF5C3185FB13FDD6" pageId="21" pageNumber="22" type="distribution">
<paragraph id="863236F5FF89FFA5FF5C3185FB13FDD6" blockId="21.[151,1436,295,863]" pageId="21" pageNumber="22">
<emphasis id="B4F9EAE7FF89FFA5FF5C3185FEF9FE09" bold="true" box="[197,364,450,476]" pageId="21" pageNumber="22">Distribution.</emphasis>
<taxonomicName id="418D4D76FF89FFA5FEEC3185FDCAFE09" box="[373,607,450,476]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="2" phylum="Chordata" rank="subTribe" status="subtrib. nov." subTribe="Tetracheilostomina">Tetracheilostomina</taxonomicName>
is distributed in the West Indies: on the island of Hispaniola in the Greater Antilles, and on
<collectingCountry id="FE9A7665FF89FFA5FE2131AEFDA8FDD6" box="[440,573,489,515]" name="Martinique" pageId="21" pageNumber="22">Martinique</collectingCountry>
,
<collectingCountry id="FE9A7665FF89FFA5FDD031AEFD47FDD6" box="[585,722,489,515]" name="Saint Lucia" pageId="21" pageNumber="22">Saint Lucia</collectingCountry>
, and
<collectingCountry id="FE9A7665FF89FFA5FC9631AEFCEBFDD6" box="[783,894,489,515]" name="Barbados" pageId="21" pageNumber="22">Barbados</collectingCountry>
in the Lesser Antilles.
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF89FFA5FF5C3248FEF5FC8A" pageId="21" pageNumber="22" type="discussion">
<paragraph id="863236F5FF89FFA5FF5C3248FEF5FC8A" blockId="21.[151,1436,295,863]" pageId="21" pageNumber="22">
<emphasis id="B4F9EAE7FF89FFA5FF5C3248FED6FDFC" bold="true" box="[197,323,527,553]" pageId="21" pageNumber="22">Remarks.</emphasis>
<taxonomicName id="418D4D76FF89FFA5FED53257FDA2FDFF" box="[332,567,528,554]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="2" phylum="Chordata" rank="subTribe" status="subtrib. nov." subTribe="Tetracheilostomina">Tetracheilostomina</taxonomicName>
includes species in the former &quot;
<taxonomicName id="418D4D76FF89FFA5FC553257FBD0FDFC" authorityName="Mcdiarmid, Campbell &amp; Toure" authorityYear="1999" box="[972,1093,528,553]" class="Reptilia" family="Leptotyphlopidae" genus="Leptotyphlops" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="bilineatus">
<emphasis id="B4F9EAE7FF89FFA5FC553257FBD0FDFC" box="[972,1093,528,553]" italics="true" pageId="21" pageNumber="22">bilineatus</emphasis>
</taxonomicName>
Group&quot; of &quot;
<taxonomicName id="418D4D76FF89FFA5FB783257FA1AFDFC" authorityName="Fitzinger" authorityYear="1843" box="[1249,1423,528,553]" class="Reptilia" family="Leptotyphlopidae" genus="Leptotyphlops" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF89FFA5FB783257FA1AFDFC" box="[1249,1423,528,553]" italics="true" pageId="21" pageNumber="22">Leptotyphlops</emphasis>
</taxonomicName>
&quot; (
<bibRefCitation id="E21C4B04FF89FFA5FF063271FED1FD85" author="Hedges" box="[159,324,566,592]" firstAuthor="Hedges" pageId="21" pageNumber="22" pagination="1 - 30" refId="ref24581" refString="Hedges, S. B. (2008) At the lower limit of size in snakes: two new species of threadsnakes (Squamata: Leptotyphlopidae: Leptotyphlops) from the Lesser Antilles. Zootaxa, 1841, 1 - 30." type="journal article" year="2008">Hedges 2008</bibRefCitation>
;
<bibRefCitation id="E21C4B04FF89FFA5FECB3271FE6EFD85" author="Thomas" box="[338,507,566,592]" firstAuthor="Thomas" pageId="21" pageNumber="22" pagination="1 - 12" refId="ref27909" refString="Thomas, R. (1965) The genus Leptotyphlops in the West Indies with the description of a new species from Hispaniola (Serpentes, Leptotyphlopidae). Breviora, Museum of Comparative Zoology, 1 - 12." type="book chapter" year="1965">Thomas 1965</bibRefCitation>
;
<bibRefCitation id="E21C4B04FF89FFA5FD903271FD6FFD85" author="Thomas" box="[521,762,566,592]" etAl="et al." firstAuthor="Thomas" pageId="21" pageNumber="22" pagination="204 - 220" refId="ref27949" refString="Thomas, R., McDiarmid, R. W. &amp; Thompson, F. G. (1985) Three new species of thread snakes (Serpentes, Leptotyphlopidae) from Hispaniola. Proceedings of the Biological Society of Washington, 98, 204 - 220." type="journal article" year="1985">
Thomas
<emphasis id="B4F9EAE7FF89FFA5FDEB3270FD25FD85" box="[626,688,567,592]" italics="true" pageId="21" pageNumber="22">et al.</emphasis>
1985
</bibRefCitation>
). The high number (four) of supralabials is rare among leptotyphlopids, otherwise occurring only in
<taxonomicName id="418D4D76FF89FFA5FD57321AFCF7FDA3" box="[718,866,605,630]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF89FFA5FD57321AFCF7FDA3" box="[718,866,605,630]" italics="true" pageId="21" pageNumber="22">Rhinoleptus</emphasis>
</taxonomicName>
. As a unifying character for this West Indian radiation it is further supported by the molecular phylogeny (
<figureCitation id="1EB62A70FF89FFA5FC0932C3FC49FD4B" box="[912,988,644,670]" captionStart="FIGURE 3" captionStartId="6.[151,255,1684,1708]" captionTargetBox="[181,1406,188,1658]" captionTargetId="figure-173@6.[179,1407,184,1660]" captionTargetPageId="6" captionText="FIGURE 3. A phylogeny of leptotyphlopid snakes based on sequences of four mitochondrial genes (12S rRNA, tRNA- Valine, 16S rRNA, and cytochrome b). Maximum likelihood tree of 91 samples and 2,971 sites. Values are ML bootstrap values followed by Bayesian posterior probabilities. Outgroups are not shown, but included Typhlopidae (Ramphotyphlops), Boidae (Boa), Pythonidae (Python), and Elapidae (Dendroaspis and Naja). The generic taxonomy in this tree reflects usage prior to this study. See Table 1 and Figure 12 for the new classification proposed here." pageId="21" pageNumber="22">Fig. 3</figureCitation>
). However, the included species are considerably divergent in other scale characters, body size, and coloration. The species from Hispaniola have a high number of middorsal scales, are thin, and pale brown or pink in color. In contrast, the Lesser Antillean species have a low number of middorsals, are stout, and dark brown in color with dull yellowish stripes. The molecular phylogeny supports the distinction of these two groups of species and we recognize them here at the generic level.
</paragraph>
</subSubSection>
</treatment>
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