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<document id="39DE7747C46E51585F283683101BBC99" 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="0E2487E3FF90FFBEFF0E3577FD97FE4F" 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="Epictinae Hedges, Adalsteinsson, &amp; Branch, New Subfamily" docType="treatment" docVersion="3" lastPageNumber="15" masterDocId="F21DFF9BFF9CFFB0FF993047FF95FFD5" masterDocTitle="Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squamata)" masterLastPageNumber="50" masterPageNumber="1" pageNumber="13" updateTime="1699087815032" updateUser="plazi" zenodo-license-document="CLOSED">
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<mods:title id="A4801E54D4458F018591258EB0F53869">Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squamata)</mods:title>
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<mods:namePart id="B3F98B12FA51206EF1DE305AB23FFF7E">Adalsteinsson, Solny A.</mods:namePart>
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<mods:namePart id="DED9D4EFCB84E9C5F11CBF5C473D2007">Branch, William R.</mods:namePart>
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<mods:namePart id="F226F574513D72F6384721598EE9C836">Trape, Sébastien</mods:namePart>
<mods:affiliation id="4B6E06F605AE4099B3082099568D1DFF">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="A6AF95ED021428C20999E4A8F7F67957">Vitt, Laurie J.</mods:namePart>
<mods:affiliation id="E9F3206DA1FB9D5A57253153A362752E">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="DCD4929A58D6E4CC79458813C5622314">Hedges, S. Blair</mods:namePart>
<mods:affiliation id="28D99A154C4133151D7AFB4D6CD45D12">Department of Biology, 208 Mueller Lab, Pennsylvania State University, University Park, PA 16802 - 5301 USA.</mods:affiliation>
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<paragraph id="863236F5FF90FFBCFF0E3577FB99FA9F" blockId="12.[151,1036,1328,1354]" box="[151,1036,1328,1354]" pageId="12" pageNumber="13">
<heading id="DD7A8199FF90FFBCFF0E3577FB99FA9F" bold="true" box="[151,1036,1328,1354]" fontSize="11" level="1" pageId="12" pageNumber="13" reason="1">
<emphasis id="B4F9EAE7FF90FFBCFF0E3577FB99FA9F" bold="true" box="[151,1036,1328,1354]" pageId="12" pageNumber="13">
Subfamily
<taxonomicName id="418D4D76FF90FFBCFE863577FCD5FA9F" authority="Hedges, Adalsteinsson, &amp; Branch" authorityName="Hedges, Adalsteinsson, &amp; Branch" box="[287,832,1328,1354]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="subFamily" status="subfam. nov." subFamily="Epictinae">Epictinae Hedges, Adalsteinsson, &amp; Branch</taxonomicName>
,
<taxonomicNameLabel id="AFCA579CFF90FFBCFCD43577FB99FA9F" box="[845,1036,1328,1354]" pageId="12" pageNumber="13" rank="subFamily">New Subfamily</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="CE97657EFF90FFBCFF0E353EFDBEFA45" box="[151,555,1401,1425]" pageId="12" pageNumber="13" type="type_taxon">
<paragraph id="863236F5FF90FFBCFF0E353EFDBEFA45" blockId="12.[151,555,1401,1425]" box="[151,555,1401,1425]" pageId="12" pageNumber="13">
<emphasis id="B4F9EAE7FF90FFBCFF0E353EFE88FA44" bold="true" box="[151,285,1401,1425]" pageId="12" pageNumber="13">
<typeStatus id="59368857FF90FFBCFF0E353EFF44FA44" box="[151,209,1401,1425]" pageId="12" pageNumber="13">Type</typeStatus>
genus.
</emphasis>
<taxonomicName id="418D4D76FF90FFBCFEBD353DFDB2FA45" authority="Gray, 1845: 139" authorityName="Gray" authorityPageNumber="139" authorityYear="1845" box="[292,551,1401,1425]" class="Reptilia" family="Leptotyphlopidae" genus="Epictia" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF90FFBCFEBD353DFEE4FA44" box="[292,369,1402,1425]" italics="true" pageId="12" pageNumber="13">Epictia</emphasis>
<bibRefCitation id="E21C4B04FF90FFBCFEEE353EFDB2FA45" author="Gray" box="[375,551,1401,1424]" firstAuthor="Gray" page="139" pageId="12" pageNumber="13" refId="ref24039" refString="Gray, J. E. (1845) Catalogue of the Specimens of Lizards in the Collections of the British Museum. British Museum, London, 289 pp." type="book" year="1845">Gray, 1845: 139</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF90FFBCFF0E3587FBAAF917" pageId="12" pageNumber="13" type="diagnosis">
<paragraph id="863236F5FF90FFBCFF0E3587FB9DF94E" blockId="12.[151,1436,1472,2001]" pageId="12" pageNumber="13">
<emphasis id="B4F9EAE7FF90FFBCFF0E3587FE8DFA0F" bold="true" box="[151,280,1472,1498]" pageId="12" pageNumber="13">Diagnosis.</emphasis>
Compared with other subfamilies, members of this subfamily tend to have short, thick tails, and the fewest subcaudal scales: relative tail length is 2.111.5% total length versus 4.118.9% in the
<taxonomicName id="418D4D76FF90FFBCFF0E364AFEFAF9F2" baseAuthorityName="Hedges" baseAuthorityYear="2008" box="[151,367,1549,1575]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="subFamily" subFamily="Leptotyphlopinae">Leptotyphlopinae</taxonomicName>
; tail shape is 1.36.1 versus 3.211.7; and subcaudals number 830 versus
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the
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(Table 2;
<figureCitation id="1EB62A70FF90FFBCFE713673FDA4F99B" box="[488,561,1588,1614]" captionStart="FIGURE 5" captionStartId="15.[151,259,1207,1231]" captionTargetBox="[172,1399,199,1182]" captionTargetId="figure-276@15.[172,1416,199,1183]" captionTargetPageId="15" captionText="FIGURE 5. Histograms showing differences in scalation and proportions among taxa of leptotyphlopid snakes, assembled from descriptions of species (cited in Table 1) and earlier summaries (e.g., Broadley &amp; Broadley 1999; Broadley &amp; Wallach 2007; Wallach 1998). (A) Subcaudal scales in Epictinae (red, left) and Leptotyphlopinae (blue, right). (B) Relative tail length (tail length/ total length x 100) in Epictinae (red, left) and Leptotyphlopinae (blue, right). (C) Tail shape (tail length/ tail width at midtail) in Epictinae (red, left) and Leptotyphlopinae (blue, right). (D) Middorsal scales in two genera of Leptotyphlopinae: Leptotyphlops (blue, left) and Myriopholis (red, right). For each panel, frequency is on the Y-axis. In panels B and C, continuous numbers were rounded to integers before binning, and therefore bins are whole numbers as indicated." pageId="12" pageNumber="13">Fig. 5</figureCitation>
). All leptotyphlopids with more than two supralabials and more than 14 midbody scale rows are in this subfamily. The support for this group was 44% BP and 0% PP for the fourgene tree (
<figureCitation id="1EB62A70FF90FFBCFE8836C6FEC2F94E" box="[273,343,1665,1691]" 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="12" pageNumber="13">Fig. 3</figureCitation>
) and 94% BP and 100% PP for the nine-gene tree (
<figureCitation id="1EB62A70FF90FFBCFC2B36C6FC6EF94E" box="[946,1019,1665,1691]" 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="12" pageNumber="13">Fig. 4</figureCitation>
).
</paragraph>
<paragraph id="863236F5FF90FFBCFF5C36EFFBAAF917" blockId="12.[151,1436,1472,2001]" box="[197,1087,1704,1730]" pageId="12" pageNumber="13">
<emphasis id="B4F9EAE7FF90FFBCFF5C36EFFEA4F917" bold="true" box="[197,305,1704,1730]" pageId="12" pageNumber="13">Content.</emphasis>
Two tribes, three subtribes, eight genera, and 62 species (
<tableCitation id="CB0F034EFF90FFBCFC4F36EFFBBBF917" box="[982,1070,1704,1730]" 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="12" pageNumber="13">Table 1</tableCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF90FFBCFF5C3689FF59F804" pageId="12" pageNumber="13" type="distribution">
<paragraph id="863236F5FF90FFBCFF5C3689FF59F804" blockId="12.[151,1436,1472,2001]" pageId="12" pageNumber="13">
<emphasis id="B4F9EAE7FF90FFBCFF5C3689FEF0F93D" bold="true" box="[197,357,1742,1768]" pageId="12" pageNumber="13">Distribution.</emphasis>
The subfamily is distributed in the New World and in equatorial Africa. In the New World it ranges from North America (California, Utah, and Kansas) south through Middle and South America (exclusive of the high Andes) to
<collectingCountry id="FE9A7665FF90FFBCFD84375BFD11F8E3" box="[541,644,1820,1846]" name="Uruguay" pageId="12" pageNumber="13">Uruguay</collectingCountry>
and
<collectingCountry id="FE9A7665FF90FFBCFD27375BFCA6F8E3" box="[702,819,1820,1846]" name="Argentina" pageId="12" pageNumber="13">Argentina</collectingCountry>
on the Atlantic side. It also occurs on San
<collectingCountry id="FE9A7665FF90FFBCFAAA375BFF4BF889" name="Salvador Island" pageId="12" pageNumber="13">Salvador Island</collectingCountry>
(
<collectingCountry id="FE9A7665FF90FFBCFF773705FECAF889" box="[238,351,1858,1884]" name="Bahamas" pageId="12" pageNumber="13">Bahamas</collectingCountry>
), Hispaniola, the Lesser Antilles, Cozumel Island (
<collectingCountry id="FE9A7665FF90FFBCFC283705FB85F889" box="[945,1040,1858,1884]" name="Mexico" pageId="12" pageNumber="13">Mexico</collectingCountry>
), Islas de Bahia and Swan Islands (
<collectingCountry id="FE9A7665FF90FFBCFF06372EFE82F856" box="[159,279,1897,1923]" name="Honduras" pageId="12" pageNumber="13">Honduras</collectingCountry>
),
<collectingRegion id="4449F817FF90FFBCFEB3372EFE24F856" box="[298,433,1897,1923]" country="Colombia" name="Providencia y Santa Catalina" pageId="12" pageNumber="13">San Andres</collectingRegion>
and Providencia Islands (
<collectingCountry id="FE9A7665FF90FFBCFD47372EFCCDF856" box="[734,856,1897,1923]" name="Colombia" pageId="12" pageNumber="13">Colombia</collectingCountry>
),
<collectingCountry id="FE9A7665FF90FFBCFCF2372EFC5EF856" box="[875,971,1897,1923]" name="Bonaire, Sint Eustatius and Saba" pageId="12" pageNumber="13">Bonaire</collectingCountry>
, Margarita Islands, and
<collectingCountry id="FE9A7665FF90FFBCFB7B372EFADFF856" box="[1250,1354,1897,1923]" name="Trinidad and Tobago" pageId="12" pageNumber="13">Trinidad</collectingCountry>
. It also occurs in equatorial Africa, from southern
<collectingCountry id="FE9A7665FF90FFBCFD0137D7FD6CF87F" box="[664,761,1936,1962]" name="Senegal" pageId="12" pageNumber="13">Senegal</collectingCountry>
,
<taxonomicName id="418D4D76FF90FFBCFC9F37D7FCCAF87F" box="[774,863,1936,1962]" class="Reptilia" family="Leptotyphlopidae" genus="Guinea" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="46" phylum="Chordata" rank="genus">Guinea</taxonomicName>
, and Bioko Island in the west to
<collectingCountry id="FE9A7665FF90FFBCFB7237D7FAC5F87F" box="[1259,1360,1936,1962]" name="Ethiopia" pageId="12" pageNumber="13">Ethiopia</collectingCountry>
in the east.
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF92FFBEFF5C30D0FD97FE4F" pageId="14" pageNumber="15" type="discussion">
<paragraph id="863236F5FF92FFBEFF5C30D0FD97FE4F" blockId="14.[151,1436,151,410]" pageId="14" pageNumber="15">
<emphasis id="B4F9EAE7FF92FFBEFF5C30D0FED4FF64" bold="true" box="[197,321,151,177]" pageId="14" pageNumber="15">Remarks.</emphasis>
The inclusion of six African species (all but one from West Africa) in this otherwise New World group (
<tableCitation id="CB0F034EFF92FFBEFEA330F9FE07FF0D" box="[314,402,190,216]" 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="14" pageNumber="15">Table 1</tableCitation>
;
<figureCitation id="1EB62A70FF92FFBEFE3830F9FD9AFF0D" box="[417,527,190,216]" captionStart-0="FIGURE 3" captionStart-1="FIGURE 4" captionStartId-0="6.[151,255,1684,1708]" captionStartId-1="7.[151,260,1282,1306]" captionTargetBox-0="[181,1406,188,1658]" captionTargetBox-1="[252,1334,313,1258]" captionTargetId-0="figure-173@6.[179,1407,184,1660]" captionTargetId-1="figure-366@7.[252,1335,310,1258]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionText-0="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." captionText-1="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="14" pageNumber="15">Figs. 34</figureCitation>
) was surprising, and was not found in morphological analyses of visceral and other data (
<bibRefCitation id="E21C4B04FF92FFBEFEB830A2FE5AFF2A" author="Wallach" box="[289,463,229,255]" firstAuthor="Wallach" pageId="14" pageNumber="15" refId="ref28986" refString="Wallach, V. (1998) The Visceral Anatomy of Blindsnakes and Wormsnakes and its Systematic Implications (Serpentes: Anomalepididae, Typhlopidae, Leptotyphlopidae). Ph. D. Dissertation. Northeastern University, Boston, 291 pp." type="book" year="1998">Wallach 1998</bibRefCitation>
). Nonetheless, the unusually high scale row count (16) of
<taxonomicName id="418D4D76FF92FFBEFB0030A2FABCFF2B" box="[1177,1321,229,254]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF92FFBEFB0030A2FABCFF2B" box="[1177,1321,229,254]" italics="true" pageId="14" pageNumber="15">Rhinoleptus</emphasis>
</taxonomicName>
has been recorded in two other New World genera in this subfamily,
<taxonomicName id="418D4D76FF92FFBEFCC7314BFC52FEF0" box="[862,967,268,293]" class="Reptilia" family="Leptotyphlopidae" genus="Mitophis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF92FFBEFCC7314BFC52FEF0" box="[862,967,268,293]" italics="true" pageId="14" pageNumber="15">Mitophis</emphasis>
</taxonomicName>
n. gen and
<taxonomicName id="418D4D76FF92FFBEFBCD314BFAB5FEF0" authorityName="Jan" authorityYear="1861" box="[1108,1312,268,293]" class="Reptilia" family="Leptotyphlopidae" genus="Tetracheilostoma" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF92FFBEFBCD314BFAB5FEF0" box="[1108,1312,268,293]" italics="true" pageId="14" pageNumber="15">Tetracheilostoma</emphasis>
</taxonomicName>
(Table 2). Also, the West African members of
<taxonomicName id="418D4D76FF92FFBEFDD93175FD3AFE99" box="[576,687,306,332]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Epictinae">Epictinae</taxonomicName>
have relatively short and thick tails, low subcaudal counts, and high supralabial counts as in New World
<taxonomicName id="418D4D76FF92FFBEFD3B311EFC8DFEA6" box="[674,792,345,371]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Epictinae">Epictinae</taxonomicName>
but in contrast to other Old World leptotyphlopids (Subfamily
<taxonomicName id="418D4D76FF92FFBEFEBB31C7FE63FE4F" baseAuthorityName="Hedges" baseAuthorityYear="2008" box="[290,502,384,410]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="subFamily" subFamily="Leptotyphlopinae">Leptotyphlopinae</taxonomicName>
).
</paragraph>
</subSubSection>
</treatment>
</document>