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<document id="4B9267609DF0D079A4D7416463A884C5" ID-CLB-Dataset="305726" ID-DOI="10.1016/j.jcz.2023.01.003" ID-GBIF-Dataset="76d25374-8cb8-4c1f-a9f2-9db8726e2855" ID-ISSN="1873-2674" ID-Zenodo-Dep="10375602" IM.bibliography_approvedBy="juliana" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="juliana" IM.metadata_approvedBy="juliana" IM.tables_approvedBy="juliana" IM.taxonomicNames_approvedBy="juliana" IM.treatmentCitations_approvedBy="felipe" IM.treatments_approvedBy="juliana" checkinTime="1702545563571" checkinUser="felipe" docAuthor="Entiauspe-Neto, Omar M., Franco, Francisco L., Koch, Claudia, Tiutenko, Arthur, Wingert, Juliana M. &amp; Borges-Martins, Márcio" docDate="2023" docId="03DCB451FF91FFC4B11EFC5AE64BFBF8" docLanguage="en" docName="ZoolAnz.303.10-25.pdf" docOrigin="Zoologischer Anzeiger 303" docSource="http://dx.doi.org/10.1016/j.jcz.2023.01.003" docStyle="DocumentStyle:32483D6D135DE2FF927BA3F1400E0FFD.1:ZoolAnz.2022-.journal_article" docStyleId="32483D6D135DE2FF927BA3F1400E0FFD" docStyleName="ZoolAnz.2022-.journal_article" docStyleVersion="1" docTitle="Liotyphlops sousai Santos &amp; Reis 2018" docType="treatment" docVersion="5" lastPageNumber="17" masterDocId="FFE5CC29FF95FFC3B22CFFF8E304FFC7" masterDocTitle="More than meets no eyes: Taxonomic status of a Liotyphlops Peters, 1881 (Serpentes: Anomalepididae) blindsnake from the Atlantic Rainforest" masterLastPageNumber="25" masterPageNumber="10" pageNumber="14" updateTime="1732951271683" updateUser="ExternalLinkService">
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<mods:title id="9EBADBF01EED24FAEDAEA8499DEB744A">More than meets no eyes: Taxonomic status of a Liotyphlops Peters, 1881 (Serpentes: Anomalepididae) blindsnake from the Atlantic Rainforest</mods:title>
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<mods:namePart id="7668180DB183FAD45F9F16835B55E648">Entiauspe-Neto, Omar M.</mods:namePart>
<mods:affiliation id="212814D1724BECD00AA54C0F88DD977F">Laborat´orio de Coleç ˜ oes Zool´ogicas, Instituto Butantan, 05503 - 90, Av. Vital Brazil, 1500, Butanta ˜, S ˜ ao Paulo, SP, Brazil &amp; Programa de P´os-Graduaç ˜ ao em Biologia Animal, Departamento de Zoologia, Instituto de Biociˆencias, Universidade Federal do Rio Grande do Sul, Avenida Bento Gonçalves, 91501 - 970, Porto Alegre, RS, Brazil</mods:affiliation>
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<mods:namePart id="C8B0440AA08BCD2728CE1BDFC7218254">Franco, Francisco L.</mods:namePart>
<mods:affiliation id="4E13BABD46FA7C98F548615C2FA4CF65">Laborat´orio de Coleç ˜ oes Zool´ogicas, Instituto Butantan, 05503 - 90, Av. Vital Brazil, 1500, Butanta ˜, S ˜ ao Paulo, SP, Brazil</mods:affiliation>
<mods:nameIdentifier id="7E9DE80C7D9ED72F77C198A852B1A50A" type="email">francisco.franco@butantan.gov.br</mods:nameIdentifier>
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<mods:namePart id="CA20B7279E2494501B3560E9DFDBCA5A">Koch, Claudia</mods:namePart>
<mods:affiliation id="C4E11748EE11934F8CE8B193F359E3E4">Leibniz Institute for the Analysis of Biodiversity Change - Museum Koenig, Adenauerallee 160, 53113, Bonn, Germany</mods:affiliation>
<mods:nameIdentifier id="CA9359DCCDF0744E31FA1216E3B53AFA" type="email">C.Koch@leibniz-zfmk.de</mods:nameIdentifier>
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<mods:namePart id="CF63FE29C959B63E2D18668D618BEE7A">Tiutenko, Arthur</mods:namePart>
<mods:affiliation id="0A3C7CC095C85C1E541D0DEF06D9A4C9">Friedrich-Alexander-Universit ¨ at Erlangen-Nürnberg, Schlossplatz 4, 91054, Erlangen, Germany</mods:affiliation>
<mods:nameIdentifier id="A96312D15A4AAF816F03D0E4E6671047" type="email">arthur@tiutenko.de</mods:nameIdentifier>
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<mods:namePart id="2074CA69614539ABD9972B60F16AA483">Wingert, Juliana M.</mods:namePart>
<mods:affiliation id="CEB8F530BEC679E7CA5B07D2E4499EC8">Laborat´orio de Coleç ˜ oes Zool´ogicas, Instituto Butantan, 05503 - 90, Av. Vital Brazil, 1500, Butanta ˜, S ˜ ao Paulo, SP, Brazil</mods:affiliation>
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<mods:namePart id="C189EB10505C4F42884DEF629502DED3">Borges-Martins, Márcio</mods:namePart>
<mods:affiliation id="D10615C9346E2C8C73C63C1B43084ADF">Laboratorio de Coleçoes Zoologicas, Instituto Butantan, 05503 - 90, Av. Vital Brazil, 1500, Butanta, Sao Paulo, SP, Brazil</mods:affiliation>
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<mods:date id="D1153B00244A68714F096F84BA54483B">2023</mods:date>
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<treatment id="03DCB451FF91FFC4B11EFC5AE64BFBF8" ID-GBIF-Taxon="248133381" LSID="urn:lsid:plazi:treatment:03DCB451FF91FFC4B11EFC5AE64BFBF8" httpUri="http://treatment.plazi.org/id/03DCB451FF91FFC4B11EFC5AE64BFBF8" lastPageId="7" lastPageNumber="17" pageId="4" pageNumber="14" scope_family="Anomalepididae">
<subSubSection id="C36F56CCFF91FFC7B11EFC5AE763FC72" box="[818,1127,930,949]" pageId="4" pageNumber="14" type="nomenclature">
<paragraph id="8BCA0547FF91FFC7B11EFC5AE763FC72" blockId="4.[818,1127,930,949]" box="[818,1127,930,949]" pageId="4" pageNumber="14">
<heading id="D082B22BFF91FFC7B11EFC5AE763FC72" box="[818,1127,930,949]" fontSize="36" level="2" pageId="4" pageNumber="14" reason="3">
<emphasis id="B901D955FF91FFC7B11EFC5AE763FC72" bold="true" box="[818,1127,930,949]" italics="true" pageId="4" pageNumber="14">
3.2. Identity of
<taxonomicName id="4C757EC4FF91FFC7B1E9FC5AE763FC72" ID-CoL="88KBD" authority="Santos &amp; Reis, 2018" authorityName="Santos &amp; Reis" authorityYear="2018" box="[965,1127,930,949]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">Liotyphlops sousai</taxonomicName>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="C36F56CCFF91FFC4B17DFC23E64BFBF8" lastPageId="7" lastPageNumber="17" pageId="4" pageNumber="14" type="description">
<paragraph id="8BCA0547FF91FFC7B17DFC23E71FFA23" blockId="4.[818,1488,986,1982]" pageId="4" pageNumber="14">
In the description of
<taxonomicName id="4C757EC4FF91FFC7B63AFC23E767FC2A" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1046,1123,986,1006]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B63AFC23E767FC2A" bold="true" box="[1046,1123,986,1006]" italics="true" pageId="4" pageNumber="14">L. sousai</emphasis>
</taxonomicName>
, no information is given regarding the sex of the type specimen (
<bibRefCitation id="EFE478B6FF91FFC7B609FC0EE60BFBCE" author="Santos, F. J. M. &amp; Reis, R. E." box="[1061,1295,1014,1034]" pageId="4" pageNumber="14" pagination="507 - 514" refId="ref15631" refString="Santos, F. J. M., Reis, R. E., 2018. Two new blind snake species of the genus Liotyphlops (Serpentes: Anomalepididae), from central and SouthSouth Brazil. Copeia 106, 507 - 514." type="journal article" year="2018">Santos &amp; Reis, 2018:507</bibRefCitation>
). We were unable to visually verify the presence of an hemipenis in the
<typeStatus id="54CEBBE5FF91FFC7B720FBEAE65AFBE2" box="[1292,1374,1042,1061]" pageId="4" pageNumber="14" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF91FFC7B750FBEAE6CDFBE2" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1404,1481,1042,1061]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B750FBEAE6CDFBE2" bold="true" box="[1404,1481,1042,1061]" italics="true" pageId="4" pageNumber="14">L. sousai</emphasis>
</taxonomicName>
. Examination of its scale counts (particularly, the number of subcaudals and ventrals to subcaudals ratio) and manual probing also corroborate that the
<typeStatus id="54CEBBE5FF91FFC7B153FB9EE0D5FBBE" box="[895,977,1126,1145]" pageId="4" pageNumber="14" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF91FFC7B1C1FB9EE73DFBBE" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1005,1081,1126,1145]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B1C1FB9EE73DFBBE" bold="true" box="[1005,1081,1126,1145]" italics="true" pageId="4" pageNumber="14">L. sousai</emphasis>
</taxonomicName>
is actually a female specimen. According to the diagnosis of
<bibRefCitation id="EFE478B6FF91FFC7B1E0FB7AE790FB52" author="Santos, F. J. M. &amp; Reis, R. E." box="[972,1172,1154,1173]" pageId="4" pageNumber="14" pagination="507 - 514" refId="ref15631" refString="Santos, F. J. M., Reis, R. E., 2018. Two new blind snake species of the genus Liotyphlops (Serpentes: Anomalepididae), from central and SouthSouth Brazil. Copeia 106, 507 - 514." type="journal article" year="2018">Santos &amp; Reis (2018)</bibRefCitation>
,
<taxonomicName id="4C757EC4FF91FFC7B68CFB7AE7E9FB52" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1184,1261,1154,1173]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B68CFB7AE7E9FB52" bold="true" box="[1184,1261,1154,1173]" italics="true" pageId="4" pageNumber="14">L. sousai</emphasis>
</taxonomicName>
“
<emphasis id="B901D955FF91FFC7B6D1FB7AE05BFB77" bold="true" italics="true" pageId="4" pageNumber="14">is distinguished from all other</emphasis>
<taxonomicName id="4C757EC4FF91FFC7B14BFB66E0DCFB76" authorityName="Peters" authorityYear="1881" box="[871,984,1182,1201]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="genus" subOrder="Serpentes">Liotyphlops</taxonomicName>
,
<emphasis id="B901D955FF91FFC7B1CFFB66E7C6FB76" bold="true" box="[995,1218,1181,1201]" italics="true" pageId="4" pageNumber="14">
except
<taxonomicName id="4C757EC4FF91FFC7B60FFB65E7C6FB76" baseAuthorityName="Cope" baseAuthorityYear="1864" box="[1059,1218,1181,1201]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="anops" subOrder="Serpentes">Liotyphlops anops</taxonomicName>
</emphasis>
,
<taxonomicName id="4C757EC4FF91FFC7B6FDFB65E682FB77" authorityName="Dixon and Kofron" authorityYear="1983" box="[1233,1414,1181,1200]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="argaleus" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B6FDFB65E682FB77" bold="true" box="[1233,1414,1181,1200]" italics="true" pageId="4" pageNumber="14">Liotyphlops argaleus</emphasis>
</taxonomicName>
,
<emphasis id="B901D955FF91FFC7B7B9FB65E077FB0B" bold="true" italics="true" pageId="4" pageNumber="14">
and
<taxonomicName id="4C757EC4FF91FFC7B792FB66E077FB0B" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="trefauti" subOrder="Serpentes">L. trefauti</taxonomicName>
</emphasis>
,
<emphasis id="B901D955FF91FFC7B152FB41E635FB2F" bold="true" italics="true" pageId="4" pageNumber="14">by having four scales contacting the posterior edge of the prefrontal (vs. three scales contacting posterior edge of prefrontal)</emphasis>
(
<bibRefCitation id="EFE478B6FF91FFC7B76EFB2EE08BFAC3" author="Santos, F. J. M. &amp; Reis, R. E." pageId="4" pageNumber="14" pagination="507 - 514" refId="ref15631" refString="Santos, F. J. M., Reis, R. E., 2018. Two new blind snake species of the genus Liotyphlops (Serpentes: Anomalepididae), from central and SouthSouth Brazil. Copeia 106, 507 - 514." type="journal article" year="2018">Santos &amp; Reis, 2018:507</bibRefCitation>
). According to the authors, the contact of four scales to the posterior edge of the prefrontal appears to be the sole diagnostic character of
<taxonomicName id="4C757EC4FF91FFC7B151FAD1E0CCFAFB" authorityName="Santos &amp; Reis" authorityYear="2018" box="[893,968,1321,1340]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B151FAD1E08CFAFB" bold="true" box="[893,904,1321,1340]" italics="true" pageId="4" pageNumber="14">L</emphasis>
.
<emphasis id="B901D955FF91FFC7B1BEFAD1E0CCFAFB" bold="true" box="[914,968,1321,1340]" italics="true" pageId="4" pageNumber="14">sousai</emphasis>
</taxonomicName>
against
<taxonomicName id="4C757EC4FF91FFC7B63AFAD1E754FAFB" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1046,1104,1321,1340]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B63AFAD1E725FAFB" bold="true" box="[1046,1057,1321,1340]" italics="true" pageId="4" pageNumber="14">L</emphasis>
.
<emphasis id="B901D955FF91FFC7B607FAD1E754FAFB" bold="true" box="[1067,1104,1321,1340]" italics="true" pageId="4" pageNumber="14">beui</emphasis>
</taxonomicName>
. As mentioned before, individual contact states among head scales in
<taxonomicName id="4C757EC4FF91FFC7B668FABDE7ADFA9F" authorityName="Peters" authorityYear="1881" box="[1092,1193,1349,1368]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="genus" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B668FABDE7ADFA9F" bold="true" box="[1092,1193,1349,1368]" italics="true" pageId="4" pageNumber="14">Liotyphlops</emphasis>
</taxonomicName>
are considered as highly variable, and other authors suggest not describing new species using small series due to this (
<bibRefCitation id="EFE478B6FF91FFC7B1CDFA85E7C3FA57" author="Dixon, J. R. &amp; Kofron, C. P." box="[993,1223,1405,1424]" pageId="4" pageNumber="14" pagination="241 - 264" refId="ref13425" refString="Dixon, J. R., Kofron, C. P., 1983. The central and SouthSouth American Anomalepid snakes of the genus Liotyphlops. Amphibia-Reptilia 4, 241 - 264." type="journal article" year="1983">Dixon and Kofron, 1983</bibRefCitation>
). The “
<emphasis id="B901D955FF91FFC7B723FA84E73FFA6C" bold="true" italics="true" pageId="4" pageNumber="14">four scales contacting the posterior edge of prefrontal</emphasis>
” are defined by
<bibRefCitation id="EFE478B6FF91FFC7B6FCFA61E6B3FA6B" author="Dixon, J. R. &amp; Kofron, C. P." box="[1232,1463,1433,1452]" pageId="4" pageNumber="14" pagination="241 - 264" refId="ref13425" refString="Dixon, J. R., Kofron, C. P., 1983. The central and SouthSouth American Anomalepid snakes of the genus Liotyphlops. Amphibia-Reptilia 4, 241 - 264." type="journal article" year="1983">Dixon and Kofron (1983)</bibRefCitation>
as being postfrontals, and regarded as variable to their size and number in all species of
<taxonomicName id="4C757EC4FF91FFC7B19DFA28E712FA24" authorityName="Peters" authorityYear="1881" box="[945,1046,1488,1507]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="genus" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B19DFA28E712FA24" bold="true" box="[945,1046,1488,1507]" italics="true" pageId="4" pageNumber="14">Liotyphlops</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BCA0547FF91FFC7B17DFA14E606F904" blockId="4.[818,1488,986,1982]" pageId="4" pageNumber="14">
Closer examination of the
<typeStatus id="54CEBBE5FF91FFC7B663FA14E7A5FA38" box="[1103,1185,1516,1535]" pageId="4" pageNumber="14" type="holotype">holotype</typeStatus>
(
<specimenCode id="DBD3AD3CFF91FFC7B683FA14E628FA38" box="[1199,1324,1516,1535]" collectionCode="UFRGS" country="Brazil" httpUri="http://grbio.org/cool/3q02-t412" name="Universidade Federale do Rio Grande do Sul" pageId="4" pageNumber="14">UFRGS 6274</specimenCode>
) reveals that the contact of four postfrontal scales to the posterior edge of prefrontal is actually absent, with a seemingly aberrant ocular scale, that produces a proximal ridge, which was misinterpreted as a projection of a postfrontal scale (Supplementary
<figureCitation id="134E19C2FF91FFC7B62FF9A4E73EF9A8" box="[1027,1082,1628,1647]" captionStart="Fig" captionStartId="4.[284,314,623,640]" captionTargetBox="[336,1220,148,595]" captionTargetPageId="4" captionText="Fig. 2. Principal Component Analysis (PCA) for females (left) and males (right) of the six OTUs analyzed in this work." figureDoi="http://doi.org/10.5281/zenodo.10375606" httpUri="https://zenodo.org/record/10375606/files/figure.png" pageId="4" pageNumber="14">Fig. 2</figureCitation>
); the left side of the head on the holotype also has three postfrontal scales contacting the prefrontal. Therefore, the diagnostic of four postfrontal scales in contact with the prefrontal scale is considered herein as unreliable and erroneous.
</paragraph>
<paragraph id="8BCA0547FF91FFC6B17DF933E145F89C" blockId="4.[818,1488,986,1982]" lastBlockId="5.[100,770,1529,1967]" lastPageId="5" lastPageNumber="15" pageId="4" pageNumber="14">
Of the
<typeStatus id="54CEBBE5FF91FFC7B1BEF933E0E0F919" box="[914,996,1739,1758]" pageId="4" pageNumber="14" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF91FFC7B62FF934E754F919" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1027,1104,1739,1759]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B62FF934E754F919" bold="true" box="[1027,1104,1739,1759]" italics="true" pageId="4" pageNumber="14">L. sousai</emphasis>
</taxonomicName>
, there are CT scan images of the skull in
<bibRefCitation id="EFE478B6FF91FFC7B11EF91FE0FEF93D" author="Santos, F. J. M. &amp; Reis, R. E." box="[818,1018,1767,1787]" pageId="4" pageNumber="14" pagination="507 - 514" refId="ref15631" refString="Santos, F. J. M., Reis, R. E., 2018. Two new blind snake species of the genus Liotyphlops (Serpentes: Anomalepididae), from central and SouthSouth Brazil. Copeia 106, 507 - 514." type="journal article" year="2018">Santos &amp; Reis (2018)</bibRefCitation>
. No description or raw data were made available on request to the authors. Therefore, we can only compare the skulls from the
<specimenCount id="9D73CECEFF91FFC7B1ABF8E7E71CF8F5" box="[903,1048,1823,1842]" count="3" pageId="4" pageNumber="14" type="generic" typeStatus="paratype">three paratypes</specimenCount>
of
<taxonomicName id="4C757EC4FF91FFC7B619F8E7E775F8F5" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1077,1137,1823,1842]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B619F8E7E775F8F5" bold="true" box="[1077,1137,1823,1842]" italics="true" pageId="4" pageNumber="14">L. beui</emphasis>
</taxonomicName>
with the images from the
<typeStatus id="54CEBBE5FF91FFC7B749F8E7E6B3F8F5" box="[1381,1463,1823,1842]" pageId="4" pageNumber="14" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF91FFC7B11EF8C3E07AF889" authorityName="Santos &amp; Reis" authorityYear="2018" box="[818,894,1851,1870]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B11EF8C3E07AF889" bold="true" box="[818,894,1851,1870]" italics="true" pageId="4" pageNumber="14">L. sousai</emphasis>
</taxonomicName>
presented in
<bibRefCitation id="EFE478B6FF91FFC7B1D6F8C3E7BAF889" author="Santos, F. J. M. &amp; Reis, R. E." box="[1018,1214,1851,1871]" pageId="4" pageNumber="14" pagination="507 - 514" refId="ref15631" refString="Santos, F. J. M., Reis, R. E., 2018. Two new blind snake species of the genus Liotyphlops (Serpentes: Anomalepididae), from central and SouthSouth Brazil. Copeia 106, 507 - 514." type="journal article" year="2018">Santos &amp; Reis (2018)</bibRefCitation>
, drawing limited conclusions regarding similarities and differences. One of the most striking differences in the skull of the
<typeStatus id="54CEBBE5FF91FFC7B63DF88BE767F841" box="[1041,1123,1907,1926]" pageId="4" pageNumber="14" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF91FFC7B6ACF88BE7C9F841" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1152,1229,1907,1926]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B6ACF88BE7C9F841" bold="true" box="[1152,1229,1907,1926]" italics="true" pageId="4" pageNumber="14">L. sousai</emphasis>
</taxonomicName>
compared to that of
<emphasis id="B901D955FF91FFC7B7A2F88BE6D4F842" bold="true" box="[1422,1488,1906,1926]" italics="true" pageId="4" pageNumber="14">
<taxonomicName id="4C757EC4FF91FFC7B7A2F88BE6CFF842" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1422,1483,1906,1926]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">L. beui</taxonomicName>
,
</emphasis>
is the fact that there is still a rather large fontanelle between the parietal bones of
<taxonomicName id="4C757EC4FF91FFC7B1A9F853E0D5F879" authorityName="Santos &amp; Reis" authorityYear="2018" box="[901,977,1963,1982]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="14" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF91FFC7B1A9F853E0D5F879" bold="true" box="[901,977,1963,1982]" italics="true" pageId="4" pageNumber="14">L. sousai</emphasis>
</taxonomicName>
and there are also rather large distances between most of the other cranial bones. Although possible to constitute an artifact of the CT scan method, these are typical features for a juvenile blind snake in which the ossification of the skull is not yet completed due to ontogenetic variation of dermal bones (
<bibRefCitation id="EFE478B6FF90FFC6B391F9B5E196F9A7" author="Cundall, D. &amp; Irish, F." box="[445,658,1613,1632]" pageId="5" pageNumber="15" pagination="349 - 692" refId="ref13371" refString="Cundall, D., Irish, F., 2008. The snake skull; pp. 349 - 692. In: Gans, C., Parsons, T. S. (Eds.), Biology of the Reptilia, vol. 20. Academic Press, London." type="book chapter" year="2008">Cundall &amp; Irish, 2008</bibRefCitation>
). Since the
<typeStatus id="54CEBBE5FF90FFC6B248F991E3BBF9BB" box="[100,191,1641,1660]" pageId="5" pageNumber="15" type="paratype">paratypes</typeStatus>
of
<taxonomicName id="4C757EC4FF90FFC6B2CFF991E226F9BC" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[227,290,1640,1660]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B2CFF991E226F9BC" bold="true" box="[227,290,1640,1660]" italics="true" pageId="5" pageNumber="15">L. beui</emphasis>
</taxonomicName>
are adult specimens, we cannot compare aspects related to bone distances and sizes of foramina between these specimens and the
<typeStatus id="54CEBBE5FF90FFC6B29AF958E20CF974" box="[182,264,1696,1715]" pageId="5" pageNumber="15" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF90FFC6B307F959E27EF974" authorityName="Santos &amp; Reis" authorityYear="2018" box="[299,378,1696,1716]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B307F959E27EF974" bold="true" box="[299,378,1696,1716]" italics="true" pageId="5" pageNumber="15">L. sousai</emphasis>
</taxonomicName>
. As a further difference, in
<taxonomicName id="4C757EC4FF90FFC6B0A0F959E1DFF974" authorityName="Santos &amp; Reis" authorityYear="2018" box="[652,731,1696,1716]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B0A0F959E1DFF974" bold="true" box="[652,731,1696,1716]" italics="true" pageId="5" pageNumber="15">L. sousai</emphasis>
</taxonomicName>
the anterior surface of the nasal is not rugose thickened, as the anterior end of the nasal does not appear pointed in the dorsal view. Instead, it exhibits foramina on both sides of the midline. As similarities of
<taxonomicName id="4C757EC4FF90FFC6B098F90CE005F8C0" authorityName="Santos &amp; Reis" authorityYear="2018" box="[692,769,1780,1799]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B098F90CE005F8C0" bold="true" box="[692,769,1780,1799]" italics="true" pageId="5" pageNumber="15">L. sousai</emphasis>
</taxonomicName>
and
<taxonomicName id="4C757EC4FF90FFC6B2BDF8E8E3D5F8E4" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[145,209,1808,1827]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B2BDF8E8E3D5F8E4" bold="true" box="[145,209,1808,1827]" italics="true" pageId="5" pageNumber="15">L. beui</emphasis>
</taxonomicName>
and shared differences to
<taxonomicName id="4C757EC4FF90FFC6B3F1F8E8E14BF8E4" baseAuthorityName="Peters" baseAuthorityYear="1858" box="[477,591,1808,1827]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="albirostris" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B3F1F8E8E14BF8E4" bold="true" box="[477,591,1808,1827]" italics="true" pageId="5" pageNumber="15">L. albirostris</emphasis>
</taxonomicName>
can be noted the absence of a supraoccipital bone, as well as a more pronounced and longer projection of the lateral flange of the nasal.
</paragraph>
<caption id="DF0A55CFFF90FFC6B248FA74E0FCFA17" ID-DOI="http://doi.org/10.5281/zenodo.10375610" ID-Zenodo-Dep="10375610" httpUri="https://zenodo.org/record/10375610/files/figure.png" pageId="5" pageNumber="15" startId="5.[100,130,1420,1437]" targetBox="[302,1286,149,1392]" targetPageId="5" targetType="figure">
<paragraph id="8BCA0547FF90FFC6B248FA74E0FCFA17" blockId="5.[100,1487,1420,1488]" pageId="5" pageNumber="15">
<emphasis id="B901D955FF90FFC6B248FA74E39AFA5A" bold="true" box="[100,158,1420,1437]" pageId="5" pageNumber="15">Fig. 3.</emphasis>
Phylogenetic inferences and uncorrected
<emphasis id="B901D955FF90FFC6B02AFA74E10BFA5B" bold="true" box="[518,527,1420,1436]" italics="true" pageId="5" pageNumber="15">p</emphasis>
-distances for Scolecophidia terminals, based on the small subunit ribosomal RNA (16S rRNA) gene fragment. A) Maximum likelihood (RAxML) phylogenetic tree inference and geographic distribution of evaluated samples (white circles =
<taxonomicName id="4C757EC4FF90FFC6B6B2FA5EE7D1FA71" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1182,1237,1445,1462]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B6B2FA5EE7D1FA71" bold="true" box="[1182,1237,1445,1462]" italics="true" pageId="5" pageNumber="15">L. beui</emphasis>
</taxonomicName>
literature records; black and white circles =
<taxonomicName id="4C757EC4FF90FFC6B2C0FA47E226FA17" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[236,290,1471,1488]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B2C0FA47E226FA17" bold="true" box="[236,290,1471,1488]" italics="true" pageId="5" pageNumber="15">L. beui</emphasis>
</taxonomicName>
examined records); B) Pairwise uncorrected
<emphasis id="B901D955FF90FFC6B089FA47E1AAFA08" bold="true" box="[677,686,1471,1487]" italics="true" pageId="5" pageNumber="15">p</emphasis>
-distances for Scolecophidia terminals.
</paragraph>
</caption>
<paragraph id="8BCA0547FF90FFC4B2A8F89CE209FA4F" blockId="5.[100,770,1529,1967]" lastBlockId="7.[100,770,1173,1611]" lastPageId="7" lastPageNumber="17" pageId="5" pageNumber="15">
The only noteworthy difference in
<taxonomicName id="4C757EC4FF90FFC6B3CDF89CE136F8B0" authorityName="Santos &amp; Reis" authorityYear="2018" box="[481,562,1892,1911]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B3CDF89CE136F8B0" bold="true" box="[481,562,1892,1911]" italics="true" pageId="5" pageNumber="15">L. sousai</emphasis>
</taxonomicName>
is the presence of a supratemporal bone, which is absent in the analyzed
<typeStatus id="54CEBBE5FF90FFC6B07CF878E1AFF854" box="[592,683,1920,1939]" pageId="5" pageNumber="15" type="paratype">paratypes</typeStatus>
of
<taxonomicName id="4C757EC4FF90FFC6B0EBF878E006F855" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[711,770,1919,1939]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B0EBF878E006F855" bold="true" box="[711,770,1919,1939]" italics="true" pageId="5" pageNumber="15">L. beui</emphasis>
</taxonomicName>
and present in the
<typeStatus id="54CEBBE5FF90FFC6B334F863E26EF869" box="[280,362,1947,1966]" pageId="5" pageNumber="15" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF90FFC6B3A7F864E2EEF869" authorityName="Boulenger" authorityYear="1896" box="[395,490,1947,1967]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="ternetzii" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B3A7F864E2EEF869" bold="true" box="[395,490,1947,1967]" italics="true" pageId="5" pageNumber="15">L. ternetzii</emphasis>
</taxonomicName>
. According to Santos (2018), this would represent an intraspecifically variable character, considering that of
<specimenCount id="9D73CECEFF90FFC6B15BF9EDE700F9EF" box="[887,1028,1557,1576]" count="4" pageId="5" pageNumber="15" type="generic">four specimens</specimenCount>
of
<taxonomicName id="4C757EC4FF90FFC6B60FF9EDE764F9EF" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1059,1120,1557,1576]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF90FFC6B60FF9EDE764F9EF" bold="true" box="[1059,1120,1557,1576]" italics="true" pageId="5" pageNumber="15">L. beui</emphasis>
</taxonomicName>
examined by this author (MCP 10853, MCP 16362, MCZ-R 16702, MCZ-R 17842), the
<specimenCount id="9D73CECEFF90FFC6B73AF9C9E69BF983" box="[1302,1439,1585,1604]" count="2" pageId="5" pageNumber="15" type="generic" typeStatus="paratype">two paratypes</specimenCount>
also lacked the supratemporal bone, whereas the other
<specimenCount id="9D73CECEFF90FFC6B72BF9B5E68BF9A7" box="[1287,1423,1613,1632]" count="2" pageId="5" pageNumber="15" type="generic">two specimens</specimenCount>
had it. Therefore, together with our additionally examined
<typeStatus id="54CEBBE5FF90FFC6B701F991E684F9BB" box="[1325,1408,1641,1660]" pageId="5" pageNumber="15" type="paratype">paratype</typeStatus>
(BMNH 1946.1.11.12), this bone was absent in
<specimenCount id="9D73CECEFF90FFC6B685F97DE644F95F" box="[1193,1344,1669,1688]" count="3" pageId="5" pageNumber="15" type="generic">three specimens</specimenCount>
and present in two.
<bibRefCitation id="EFE478B6FF90FFC6B148F959E0DFF974" author="Rage, J. C." box="[868,987,1696,1716]" pageId="5" pageNumber="15" refId="ref15468" refString="Rage, J. C., 1984. Serpentes. Handbuch der Palaontologie, part 11. G. Fischer Verlag, Stuttgart." type="book" year="1984">Rage (1984)</bibRefCitation>
describes the supratemporal as a thin and minute, rudimentary bone in
<taxonomicName id="4C757EC4FF90FFC6B1D4F944E796F908" authorityName="Taylor" authorityYear="1939" box="[1016,1170,1724,1743]" class="Squamata" family="Anomalepididae" kingdom="Animalia" pageId="5" pageNumber="15" phylum="Chordata" rank="family">Anomalepididae</taxonomicName>
and
<taxonomicName id="4C757EC4FF90FFC6B693F944E662F908" authorityName="Stejneger" authorityYear="1892" box="[1215,1382,1724,1743]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="family" subOrder="Serpentes">Leptotyphlopidae</taxonomicName>
, lacking its functional articulation with the skull. In
<taxonomicName id="4C757EC4FF90FFC6B69CF920E6CBF92C" authority=", Kley (2006)" authorityName="Kley" authorityYear="2006" box="[1200,1487,1752,1771]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="15" phylum="Chordata" rank="family" subOrder="Serpentes">
Leptotyphlopidae,
<bibRefCitation id="EFE478B6FF90FFC6B74CF920E6CBF92C" author="Kley, N. J." box="[1376,1487,1752,1771]" pageId="5" pageNumber="15" pagination="494 - 515" refId="ref14365" refString="Kley, N. J., 2006. Morphology of the lower jaw and suspensorium in the Texas blindsnake, Leptotyphlops dulcis (Scolecophidia: Leptotyphlopidae). J. Morphol. 267, 494 - 515." type="journal article" year="2006">Kley (2006)</bibRefCitation>
</taxonomicName>
reports a non-ossified ligament that is displaced between the lateral surface of the exoccipital and dorsomedial aspect of the proximal quadrate head, in accordance with the interpretation of
<bibRefCitation id="EFE478B6FF90FFC6B77FF8D4E6CDF8F8" author="Rage, J. C." box="[1363,1481,1836,1855]" pageId="5" pageNumber="15" refId="ref15468" refString="Rage, J. C., 1984. Serpentes. Handbuch der Palaontologie, part 11. G. Fischer Verlag, Stuttgart." type="book" year="1984">Rage (1984)</bibRefCitation>
, homologous to the supratemporal bone.
<bibRefCitation id="EFE478B6FF90FFC6B697F8B0E629F89C" author="Kley, N. J." box="[1211,1325,1864,1883]" pageId="5" pageNumber="15" pagination="494 - 515" refId="ref14365" refString="Kley, N. J., 2006. Morphology of the lower jaw and suspensorium in the Texas blindsnake, Leptotyphlops dulcis (Scolecophidia: Leptotyphlopidae). J. Morphol. 267, 494 - 515." type="journal article" year="2006">Kley (2006)</bibRefCitation>
raises the possibility of the transformation of dermal bones into ligamentous tissues for the supratemporal, which could be prone to ontogenetic and intraspecific variation, and also would not be visible in the osteological analyses, requiring histological analyses for their visualization. Furthermore,
<bibRefCitation id="EFE478B6FF93FFC5B248F87EE235F85E" author="Rieppel, O. &amp; Kley, N. J. &amp; Maisano, J. A." box="[100,305,1926,1945]" pageId="6" pageNumber="16" pagination="536 - 557" refId="ref15527" refString="Rieppel, O., Kley, N. J., Maisano, J. A., 2009. Morphology of the skull of the white-nosed blindsnake, Liotyphlops albirostris (Scolecophidia: Anomalepididae). J. Morphol. 270, 536 - 557." type="journal article" year="2009">Rieppel et al. (2009)</bibRefCitation>
reports different degrees in ossification for the supratemporal in anomalepidids, ranging from the much reduced supratemporal of
<taxonomicName id="4C757EC4FF93FFC5B1CCF891E654F8BA" authority="(Schlegel, 1839)" baseAuthorityName="Schlegel" baseAuthorityYear="1839" box="[992,1360,1897,1917]" class="Squamata" family="Anomalepididae" genus="Typhlophis" kingdom="Animalia" pageId="6" pageNumber="16" phylum="Chordata" rank="species" species="squamosus">
<emphasis id="B901D955FF93FFC5B1CCF891E7ADF8BA" bold="true" box="[992,1193,1897,1917]" italics="true" pageId="6" pageNumber="16">Typhlophis squamosus</emphasis>
(Schlegel, 1839)
</taxonomicName>
to absent in
<taxonomicName id="4C757EC4FF93FFC5B11EF87DE773F85E" authority="Taylor, 1939" authorityName="Taylor" authorityYear="1939" box="[818,1143,1925,1945]" class="Squamata" family="Anomalepididae" genus="Anomalepis" kingdom="Animalia" pageId="6" pageNumber="16" phylum="Chordata" rank="species" species="aspinosus">
<emphasis id="B901D955FF93FFC5B11EF87DE0F7F85F" bold="true" box="[818,1011,1925,1944]" italics="true" pageId="6" pageNumber="16">Anomalepis aspinosus</emphasis>
Taylor, 1939
</taxonomicName>
.
<bibRefCitation id="EFE478B6FF93FFC5B6ADF87EE7F2F85E" author="Palci, A." box="[1153,1270,1926,1945]" pageId="6" pageNumber="16" refId="ref14992" refString="Palci, A., 2014. On the Origin and Evolution of the Ophidia. Doctoral dissertation. University of Alberta]. Era Library University of Alberta." type="book" year="2014">Palci (2014)</bibRefCitation>
also reports a variable presence of supratemporal bones in the genus
<taxonomicName id="4C757EC4FF93FFC5B728F859E66FF873" authorityName="Jan" authorityYear="1860" box="[1284,1387,1953,1972]" class="Squamata" family="Anomalepididae" genus="Anomalepis" kingdom="Animalia" pageId="6" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B901D955FF93FFC5B728F859E66FF873" bold="true" box="[1284,1387,1953,1972]" italics="true" pageId="6" pageNumber="16">Anomalepis</emphasis>
</taxonomicName>
, that was previously reported as lacking a supratemporal bone (
<bibRefCitation id="EFE478B6FF92FFC4B05BFB6DE3D9FB03" author="McDowell, S. B. &amp; Bogert, C. M." pageId="7" pageNumber="17" pagination="1 - 142" refId="ref14682" refString="McDowell, S. B., Bogert, C. M., 1954. The systematic position of Lanthanotus and the affinities of the anguinomorphan lizards. Bull. Am. Mus. Nat. Hist. 105, 1 - 142." type="journal article" year="1954">McDowell and Bogert, 1954</bibRefCitation>
;
<bibRefCitation id="EFE478B6FF92FFC4B2C4FB4AE255FB03" author="Haas, G." box="[232,337,1201,1221]" pageId="7" pageNumber="17" pagination="63 - 139" refId="ref14101" refString="Haas, G., 1968. Anatomical observations on the head of Anomalepis aspinosus (Typhlopidae, Ophidia). Acta Zool. 49, 63 - 139." type="journal article" year="1968">Haas, 1968</bibRefCitation>
;
<bibRefCitation id="EFE478B6FF92FFC4B377FB49E114FB03" author="Rieppel, O. &amp; Kley, N. J. &amp; Maisano, J. A." box="[347,528,1201,1221]" pageId="7" pageNumber="17" pagination="536 - 557" refId="ref15527" refString="Rieppel, O., Kley, N. J., Maisano, J. A., 2009. Morphology of the skull of the white-nosed blindsnake, Liotyphlops albirostris (Scolecophidia: Anomalepididae). J. Morphol. 270, 536 - 557." type="journal article" year="2009">Rieppel et al., 2009</bibRefCitation>
).
<bibRefCitation id="EFE478B6FF92FFC4B00FFB49E1ABFB03" author="Rieppel, O." box="[547,687,1201,1220]" pageId="7" pageNumber="17" pagination="140 - 150" refId="ref15493" refString="Rieppel, O., 1979. A cladistic classification of primitive snakes based on skull structure. J. Zool. Syst. Evol. Res. 17, 140 - 150." type="journal article" year="1979">Rieppel (1979)</bibRefCitation>
suggests that neoteny, paedomorphosis in ontogenetic ossification, can be attributable for the disappearance of the lateral wing of the basisphenoid in Scolecophidia. Therefore, we find it likely that the visualization of the supratemporal bone in scolecophids, possibly related to different degrees in ossification, demineralization due to exposure to formalin, or even visualization technique artifacts, is prone to intraspecific variation.
</paragraph>
<caption id="DF0A55CFFF93FFC5B623FF6DE69EFE35" ID-DOI="http://doi.org/10.5281/zenodo.10375612" ID-Zenodo-Dep="10375612" httpUri="https://zenodo.org/record/10375612/files/figure.png" pageId="6" pageNumber="16" startId="6.[1039,1069,149,166]" targetBox="[100,1015,148,707]" targetPageId="6" targetType="figure">
<paragraph id="8BCA0547FF93FFC5B623FF6DE69EFE35" blockId="6.[1039,1488,149,498]" pageId="6" pageNumber="16">
<emphasis id="B901D955FF93FFC5B623FF6DE74AFF61" bold="true" box="[1039,1102,149,166]" pageId="6" pageNumber="16">Fig. 4.</emphasis>
Maximum clade credibility Bayesian tree, based on the small subunit ribosomal RNA (16S rRNA) gene fragment, highlighting the divergence time of terminals. Bars represent the 95% Highest Posterior Density (HPD) interval for divergence dates. Scale indicates million years ago (Myr). Full topology and posterior probability support of nodes are provided as supplementary material. Analyses of species delimitation are: Bayesian Poisson Tree Process (bPTP); Poisson Tree Process (PTP); Assemble Species by Automatic Partitioning (ASAP); Generalized Mixed Yule Coalescent (GMYC). Inset photograph:
<taxonomicName id="4C757EC4FF93FFC5B67DFE3FE611FE1F" baseAuthorityName="Garman" baseAuthorityYear="1883" box="[1105,1301,455,472]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" isUncertain="true" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="16" phylum="Chordata" rank="species" species="wilderi" subOrder="Serpentes">
<emphasis id="B901D955FF93FFC5B67DFE3FE7A8FE1F" bold="true" box="[1105,1196,455,472]" italics="true" pageId="6" pageNumber="16">Liotyphlops</emphasis>
aff.
<emphasis id="B901D955FF93FFC5B6F3FE3FE611FE1F" bold="true" box="[1247,1301,455,472]" italics="true" pageId="6" pageNumber="16">wilderi</emphasis>
</taxonomicName>
from Minas Gerais, Brazil. Photograph credits: Henrique C. Costa.
</paragraph>
</caption>
<caption id="DF0A55CFFF93FFC5B248F8EEE03DF886" ID-DOI="http://doi.org/10.5281/zenodo.10375614" ID-Zenodo-Dep="10375614" httpUri="https://zenodo.org/record/10375614/files/figure.png" pageId="6" pageNumber="16" startId="6.[100,130,1814,1831]" targetBox="[344,1252,840,1780]" targetPageId="6" targetType="figure">
<paragraph id="8BCA0547FF93FFC5B248F8EEE03DF886" blockId="6.[100,1487,1813,1857]" pageId="6" pageNumber="16">
<emphasis id="B901D955FF93FFC5B248F8EEE399F8E1" bold="true" box="[100,157,1814,1831]" pageId="6" pageNumber="16">Fig. 5.</emphasis>
Head scalation of
<taxonomicName id="4C757EC4FF93FFC5B311F8EEE2B9F8E0" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[317,445,1814,1831]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="16" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF93FFC5B311F8EEE2B9F8E0" bold="true" box="[317,445,1814,1831]" italics="true" pageId="6" pageNumber="16">Liotyphlops beui</emphasis>
</taxonomicName>
(A, MCZ-R 17842, paratype) from Instituto Butantan, S˜ao Paulo, S˜ao Paulo state, southeastern Brazil, and
<taxonomicName id="4C757EC4FF93FFC5B712F8EEE6CBF8E0" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1342,1487,1814,1831]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="16" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF93FFC5B712F8EEE6CBF8E0" bold="true" box="[1342,1487,1814,1831]" italics="true" pageId="6" pageNumber="16">Liotyphlops sousai</emphasis>
</taxonomicName>
(B, UFRGS 6274, holotype), from Passos Maia, Santa Catarina state, southern Brazil.
</paragraph>
</caption>
<caption id="DF0A55CFFF92FFC4B248FC0DE3B6FBAB" ID-DOI="http://doi.org/10.5281/zenodo.10375618" ID-Zenodo-Dep="10375618" httpUri="https://zenodo.org/record/10375618/files/figure.png" pageId="7" pageNumber="17" startId="7.[100,130,1013,1030]" targetBox="[118,747,149,962]" targetPageId="7" targetType="figure">
<paragraph id="8BCA0547FF92FFC4B248FC0DE3B6FBAB" blockId="7.[100,771,1013,1132]" pageId="7" pageNumber="17">
<emphasis id="B901D955FF92FFC4B248FC0DE3A0FBC1" bold="true" box="[100,164,1013,1030]" pageId="7" pageNumber="17">Fig. 6.</emphasis>
Specimens of
<taxonomicName id="4C757EC4FF92FFC4B302FC0DE28DFBC1" authorityName="Peters" authorityYear="1881" box="[302,393,1013,1030]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="genus" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B302FC0DE28DFBC1" bold="true" box="[302,393,1013,1030]" italics="true" pageId="7" pageNumber="17">Liotyphlops</emphasis>
</taxonomicName>
from Passos Maia, Santa Catarina state, southern Brazil. A) Dorsal and ventral views of UFRGS 6274, juvenile female holotype of
<taxonomicName id="4C757EC4FF92FFC4B2FDFBD0E261FBFE" authorityName="Santos &amp; Reis" authorityYear="2018" box="[209,357,1064,1081]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B2FDFBD0E261FBFE" bold="true" box="[209,357,1064,1081]" italics="true" pageId="7" pageNumber="17">Liotyphlops sousai</emphasis>
</taxonomicName>
; B) Dorsal and ventral views of UFRGS 6275, juvenile male identified as
<taxonomicName id="4C757EC4FF92FFC4B364FBBAE2CDFB94" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[328,457,1090,1107]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B364FBBAE2CDFB94" bold="true" box="[328,457,1090,1107]" italics="true" pageId="7" pageNumber="17">Liotyphlops beui</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFE478B6FF92FFC4B3C4FBBAE1AFFB94" author="Santos, F. J. M. &amp; Reis, R. E." box="[488,683,1090,1107]" pageId="7" pageNumber="17" pagination="507 - 514" refId="ref15631" refString="Santos, F. J. M., Reis, R. E., 2018. Two new blind snake species of the genus Liotyphlops (Serpentes: Anomalepididae), from central and SouthSouth Brazil. Copeia 106, 507 - 514." type="journal article" year="2018">Santos and Reis (2018)</bibRefCitation>
. Scale bar = 5 mm.
</paragraph>
</caption>
<paragraph id="8BCA0547FF92FFC4B2A8FA69E0F5FE31" blockId="7.[100,770,1173,1611]" lastBlockId="7.[818,1488,147,1088]" pageId="7" pageNumber="17">
Our final alignment totalized 490 base pairs of the 16S rRNA partial gene fragment, for 24 Scolecophidia terminals and two outgroups (Supplementary File 1). Our maximum likelihood phylogenetic inference recovers a final tree topology with a score InL = 2336.628103, supporting the monophyly of the genera
<taxonomicName id="4C757EC4FF92FFC4B3C4F9F8E149F9D4" authorityName="Peters" authorityYear="1881" box="[488,589,1536,1555]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="genus" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B3C4F9F8E149F9D4" bold="true" box="[488,589,1536,1555]" italics="true" pageId="7" pageNumber="17">Liotyphlops</emphasis>
</taxonomicName>
,
<taxonomicName id="4C757EC4FF92FFC4B076F9F8E1ADF9D4" box="[602,681,1536,1555]" class="Squamata" family="Typhlopidae" genus="Typhlops" kingdom="Animalia" pageId="7" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="B901D955FF92FFC4B076F9F8E1ADF9D4" bold="true" box="[602,681,1536,1555]" italics="true" pageId="7" pageNumber="17">Typhlops</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C757EC4FF92FFC4B0F3F9F8E380F9E8" authorityName="Gray" authorityYear="1845" class="Squamata" family="Leptotyphlopidae" genus="Epictia" kingdom="Animalia" pageId="7" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="B901D955FF92FFC4B0F3F9F8E380F9E8" bold="true" italics="true" pageId="7" pageNumber="17">Epictia</emphasis>
</taxonomicName>
, with a strongly supported clade (bootstrap support
<emphasis id="B901D955FF92FFC4B045F9E4E17DF9E8" box="[617,633,1564,1583]" italics="true" pageId="7" pageNumber="17">&gt;</emphasis>
90) containing
<taxonomicName id="4C757EC4FF92FFC4B248F9C0E3B6F98C" authorityName="Santos &amp; Reis" authorityYear="2018" box="[100,178,1592,1611]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B248F9C0E3B6F98C" bold="true" box="[100,178,1592,1611]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
(UFRGS 6274) from Santa Catarina and
<taxonomicName id="4C757EC4FF92FFC4B017F9C0E17CF98C" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[571,632,1592,1611]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="24" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
L.
<emphasis id="B901D955FF92FFC4B07FF9C0E17CF98C" bold="true" box="[595,632,1592,1611]" italics="true" pageId="7" pageNumber="17">beui</emphasis>
</taxonomicName>
(KR815891.1, UFRGS 6494) from S˜ao Paulo and
<collectingRegion id="49B1CBA5FF92FFC4B6A5FF6CE639FF60" box="[1161,1341,148,167]" country="Brazil" name="Rio Grande do Sul" pageId="7" pageNumber="17">Rio Grande do Sul</collectingRegion>
(
<figureCitation id="134E19C2FF92FFC4B762FF6CE69CFF60" box="[1358,1432,148,167]" captionStart="Fig" captionStartId="5.[100,130,1420,1437]" captionTargetBox="[302,1286,149,1392]" captionTargetId="figure-376@5.[301,1287,148,1393]" captionTargetPageId="5" captionText="Fig. 3. Phylogenetic inferences and uncorrected p-distances for Scolecophidia terminals, based on the small subunit ribosomal RNA (16S rRNA) gene fragment. A) Maximum likelihood (RAxML) phylogenetic tree inference and geographic distribution of evaluated samples (white circles = L. beui literature records; black and white circles = L. beui examined records); B) Pairwise uncorrected p-distances for Scolecophidia terminals." figureDoi="http://doi.org/10.5281/zenodo.10375610" httpUri="https://zenodo.org/record/10375610/files/figure.png" pageId="7" pageNumber="17">Fig. 3A</figureCitation>
). The observed uncorrected
<emphasis id="B901D955FF92FFC4B624FF48E716FF04" bold="true" box="[1032,1042,176,195]" italics="true" pageId="7" pageNumber="17">p</emphasis>
-distances also provide no support for genetic divergence among these taxa, ranging from 0 to 1% (
<figureCitation id="134E19C2FF92FFC4B70CFF33E660FF18" box="[1312,1380,203,223]" captionStart="Fig" captionStartId="5.[100,130,1420,1437]" captionTargetBox="[302,1286,149,1392]" captionTargetId="figure-376@5.[301,1287,148,1393]" captionTargetPageId="5" captionText="Fig. 3. Phylogenetic inferences and uncorrected p-distances for Scolecophidia terminals, based on the small subunit ribosomal RNA (16S rRNA) gene fragment. A) Maximum likelihood (RAxML) phylogenetic tree inference and geographic distribution of evaluated samples (white circles = L. beui literature records; black and white circles = L. beui examined records); B) Pairwise uncorrected p-distances for Scolecophidia terminals." figureDoi="http://doi.org/10.5281/zenodo.10375610" httpUri="https://zenodo.org/record/10375610/files/figure.png" pageId="7" pageNumber="17">Fig. 3B</figureCitation>
). Similarly, intraspecific genetic variation for
<taxonomicName id="4C757EC4FF92FFC4B647FF1FE6CBFF3D" authority="(Richmond, 1955)" baseAuthorityName="Richmond" baseAuthorityYear="1955" box="[1131,1487,231,250]" class="Squamata" family="Typhlopidae" genus="Typhlops" kingdom="Animalia" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="biminiensis">
<emphasis id="B901D955FF92FFC4B647FF1FE624FF3D" bold="true" box="[1131,1312,231,250]" italics="true" pageId="7" pageNumber="17">Typhlops biminiensis</emphasis>
(Richmond, 1955)
</taxonomicName>
are also recovered as ranging from 0 to 1%, in
<taxonomicName id="4C757EC4FF92FFC4B6D6FEFBE066FEF5" authority="Klauber, 1939" authorityName="Klauber" authorityYear="1939" class="Squamata" family="Leptotyphlopidae" genus="Epictia" kingdom="Animalia" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="tenella">
<emphasis id="B901D955FF92FFC4B6D6FEFBE67CFED1" bold="true" box="[1274,1400,259,278]" italics="true" pageId="7" pageNumber="17">Epictia tenella</emphasis>
Klauber, 1939
</taxonomicName>
as 0%, and in
<taxonomicName id="4C757EC4FF92FFC4B1C6FEE7E7E6FEF5" authority="(Peters, 1858)" baseAuthorityName="Peters" baseAuthorityYear="1858" box="[1002,1250,287,307]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="albirostris" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B1C6FEE7E75CFEF5" bold="true" box="[1002,1112,287,306]" italics="true" pageId="7" pageNumber="17">L. albirostris</emphasis>
(
<bibRefCitation id="EFE478B6FF92FFC4B64AFED8E7DEFEF5" author="Peters, W. C. H." box="[1126,1242,287,307]" pageId="7" pageNumber="17" refId="ref15251" refString="Peters, W. C. H., 1858. Vier neue amerikanische Schlangen aus der Familie der Typhlopinen und daruber einige vorlaufige ¨ Mittheilungen. Monatsber. k ¨ onigl. Akad. Wiss. Berlin. 1857, 402, 1857." type="journal volume" year="1858">Peters, 1858</bibRefCitation>
)
</taxonomicName>
as ranging from 0 to 2%. Our ASAP, GMYC, PTP, and bPTP species delimitation analyses also support a single evolutionary entity between
<taxonomicName id="4C757EC4FF92FFC4B6CEFEAFE634FEAD" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1250,1328,343,362]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B6CEFEAFE634FEAD" bold="true" box="[1250,1328,343,362]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
and
<taxonomicName id="4C757EC4FF92FFC4B74EFEAFE69BFEAD" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1378,1439,343,362]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B74EFEAFE69BFEAD" bold="true" box="[1378,1439,343,362]" italics="true" pageId="7" pageNumber="17">L. beui</emphasis>
</taxonomicName>
. Our Bayesian inference also recovers a single clade with
<taxonomicName id="4C757EC4FF92FFC4B739FE8BE665FE41" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1301,1377,371,390]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B739FE8BE665FE41" bold="true" box="[1301,1377,371,390]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
and
<taxonomicName id="4C757EC4FF92FFC4B7A3FE8BE6CEFE42" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1423,1482,370,390]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B7A3FE8BE6CEFE42" bold="true" box="[1423,1482,370,390]" italics="true" pageId="7" pageNumber="17">L. beui</emphasis>
</taxonomicName>
, with terminals likely diverging during the early Pleistocene, an interspecific divergence similar to what is observed in
<taxonomicName id="4C757EC4FF92FFC4B721FE53E678FE7A" baseAuthorityName="Peters" baseAuthorityYear="1858" box="[1293,1404,426,446]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="albirostris" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B721FE53E678FE7A" bold="true" box="[1293,1404,426,446]" italics="true" pageId="7" pageNumber="17">L. albirostris</emphasis>
</taxonomicName>
(
<figureCitation id="134E19C2FF92FFC4B7A7FE53E6C5FE79" box="[1419,1473,427,446]" captionStart="Fig" captionStartId="6.[1039,1069,149,166]" captionTargetBox="[100,1015,148,707]" captionTargetPageId="6" captionText="Fig. 4. Maximum clade credibility Bayesian tree, based on the small subunit ribosomal RNA (16S rRNA) gene fragment, highlighting the divergence time of terminals. Bars represent the 95% Highest Posterior Density (HPD) interval for divergence dates. Scale indicates million years ago (Myr). Full topology and posterior probability support of nodes are provided as supplementary material. Analyses of species delimitation are: Bayesian Poisson Tree Process (bPTP); Poisson Tree Process (PTP); Assemble Species by Automatic Partitioning (ASAP); Generalized Mixed Yule Coalescent (GMYC). Inset photograph: Liotyphlops aff. wilderi from Minas Gerais, Brazil. Photograph credits: Henrique C. Costa." figureDoi="http://doi.org/10.5281/zenodo.10375612" httpUri="https://zenodo.org/record/10375612/files/figure.png" pageId="7" pageNumber="17">Fig. 4</figureCitation>
). These results consistently support a conspecific relationship between
<taxonomicName id="4C757EC4FF92FFC4B11EFE1BE07BFE32" authorityName="Santos &amp; Reis" authorityYear="2018" box="[818,895,482,502]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B11EFE1BE07BFE32" bold="true" box="[818,895,482,502]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
and
<taxonomicName id="4C757EC4FF92FFC4B183FE1BE0EFFE32" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[943,1003,482,502]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B183FE1BE0EFFE32" bold="true" box="[943,1003,482,502]" italics="true" pageId="7" pageNumber="17">L. beui</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BCA0547FF92FFC4B17DFE06E7E3FCEF" blockId="7.[818,1488,147,1088]" pageId="7" pageNumber="17">
Another issue is raised by the sympatry of
<taxonomicName id="4C757EC4FF92FFC4B72DFE07E656FDD6" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1281,1362,510,530]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B72DFE07E656FDD6" bold="true" box="[1281,1362,510,530]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
and
<taxonomicName id="4C757EC4FF92FFC4B7A6FE07E6CEFDD6" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1418,1482,510,530]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B7A6FE07E6CEFDD6" bold="true" box="[1418,1482,510,530]" italics="true" pageId="7" pageNumber="17">L. beui</emphasis>
</taxonomicName>
. Although there is no explicit mention to this in the work of
<bibRefCitation id="EFE478B6FF92FFC4B77DFDE2E076FD8E" author="Santos, F. J. M. &amp; Reis, R. E." pageId="7" pageNumber="17" pagination="507 - 514" refId="ref15631" refString="Santos, F. J. M., Reis, R. E., 2018. Two new blind snake species of the genus Liotyphlops (Serpentes: Anomalepididae), from central and SouthSouth Brazil. Copeia 106, 507 - 514." type="journal article" year="2018">Santos &amp; Reis (2018)</bibRefCitation>
, another specimen of
<taxonomicName id="4C757EC4FF92FFC4B668FDCEE7ADFD8E" authorityName="Peters" authorityYear="1881" box="[1092,1193,566,585]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="genus" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B668FDCEE7ADFD8E" bold="true" box="[1092,1193,566,585]" italics="true" pageId="7" pageNumber="17">Liotyphlops</emphasis>
</taxonomicName>
was also found in the same locality as the type of
<taxonomicName id="4C757EC4FF92FFC4B1D2FDAAE74AFDA2" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1022,1102,594,613]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B1D2FDAAE74AFDA2" bold="true" box="[1022,1102,594,613]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
, the Usina Hidrel´etrica Passos Maia, at Passos Maia municipality, Santa Catarina,
<collectingCountry id="F36245D7FF92FFC4B6FCFD96E60DFD46" box="[1232,1289,622,641]" name="Brazil" pageId="7" pageNumber="17">Brazil</collectingCountry>
, as indicated by the appendix of examined material. The single mentioned specimen (UFRGS 6275), also from Usina Hidrel´etrica Passos Maia, was identified by
<bibRefCitation id="EFE478B6FF92FFC4B11EFD3AE70DFD12" author="Santos, F. J. M. &amp; Reis, R. E." box="[818,1033,705,725]" pageId="7" pageNumber="17" pagination="507 - 514" refId="ref15631" refString="Santos, F. J. M., Reis, R. E., 2018. Two new blind snake species of the genus Liotyphlops (Serpentes: Anomalepididae), from central and SouthSouth Brazil. Copeia 106, 507 - 514." type="journal article" year="2018">Santos &amp; Reis (2018)</bibRefCitation>
as
<taxonomicName id="4C757EC4FF92FFC4B61FFD3AE771FD13" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1075,1141,705,725]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B61FFD3AE73AFD12" bold="true" box="[1075,1086,706,725]" italics="true" pageId="7" pageNumber="17">L</emphasis>
.
<emphasis id="B901D955FF92FFC4B67CFD39E771FD13" bold="true" box="[1104,1141,705,724]" italics="true" pageId="7" pageNumber="17">beui</emphasis>
</taxonomicName>
. Further examination in the same collection revealed additional specimens from the same locality, and that these would represent a series (UFRGS 6272, 6273, 6274 [
<typeStatus id="54CEBBE5FF92FFC4B755FD01E6CBFCCB" box="[1401,1487,761,780]" pageId="7" pageNumber="17" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF92FFC4B167FCEEE09CFCEF" authorityName="Santos &amp; Reis" authorityYear="2018" box="[843,920,789,809]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B167FCEEE09CFCEF" bold="true" box="[843,920,789,809]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
], 6275) collected on the same day.
</paragraph>
<paragraph id="8BCA0547FF92FFC4B17DFCC9E64BFBF8" blockId="7.[818,1488,147,1088]" pageId="7" pageNumber="17">
Direct examination of the series of
<taxonomicName id="4C757EC4FF92FFC4B6B4FCC9E7F9FC83" authorityName="Peters" authorityYear="1881" box="[1176,1277,817,836]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="genus" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B6B4FCC9E7F9FC83" bold="true" box="[1176,1277,817,836]" italics="true" pageId="7" pageNumber="17">Liotyphlops</emphasis>
</taxonomicName>
specimens from Usina Hidrel´etrica Passos Maia (UFRGS 6272, 6273, 6274 [
<typeStatus id="54CEBBE5FF92FFC4B775FCB5E6ABFCA7" box="[1369,1455,845,864]" pageId="7" pageNumber="17" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF92FFC4B11EFC91E087FCBB" authorityName="Santos &amp; Reis" authorityYear="2018" box="[818,899,873,892]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B11EFC91E087FCBB" bold="true" box="[818,899,873,892]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
], 6275) reveals no consistent character for its interspecific diagnosis. A comparison of size and coloration between the
<typeStatus id="54CEBBE5FF92FFC4B748FC7DE6B2FC5F" box="[1380,1462,901,920]" pageId="7" pageNumber="17" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C757EC4FF92FFC4B11EFC59E079FC73" authorityName="Santos &amp; Reis" authorityYear="2018" box="[818,893,929,948]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B11EFC59E079FC73" bold="true" box="[818,893,929,948]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
(UFRGS 6274) and a sympatric
<taxonomicName id="4C757EC4FF92FFC4B688FC59E7DAFC74" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1188,1246,928,948]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B688FC59E7DAFC74" bold="true" box="[1188,1246,928,948]" italics="true" pageId="7" pageNumber="17">L. beui</emphasis>
</taxonomicName>
(UFRGS 6275) (
<figureCitation id="134E19C2FF92FFC4B759FC59E6C3FC73" box="[1397,1479,929,948]" captionStart-0="Fig" captionStart-1="Fig" captionStartId-0="6.[100,130,1814,1831]" captionStartId-1="7.[100,130,1013,1030]" captionTargetBox-0="[344,1252,840,1780]" captionTargetBox-1="[118,747,149,962]" captionTargetId-0="figure-285@6.[335,1253,793,1786]" captionTargetId-1="figure-707@7.[106,767,148,986]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionText-0="Fig. 5. Head scalation of Liotyphlops beui (A, MCZ-R 17842, paratype) from Instituto Butantan, S˜ao Paulo, S˜ao Paulo state, southeastern Brazil, and Liotyphlops sousai (B, UFRGS 6274, holotype), from Passos Maia, Santa Catarina state, southern Brazil." captionText-1="Fig. 6. Specimens of Liotyphlops from Passos Maia, Santa Catarina state, southern Brazil. A) Dorsal and ventral views of UFRGS 6274, juvenile female holotype of Liotyphlops sousai; B) Dorsal and ventral views of UFRGS 6275, juvenile male identified as Liotyphlops beui by Santos and Reis (2018). Scale bar = 5 mm." figureDoi-0="http://doi.org/10.5281/zenodo.10375614" figureDoi-1="http://doi.org/10.5281/zenodo.10375618" httpUri-0="https://zenodo.org/record/10375614/files/figure.png" httpUri-1="https://zenodo.org/record/10375618/files/figure.png" pageId="7" pageNumber="17">Figs. 5–6</figureCitation>
) reveals identical phenotypes. Furthermore, a comparison between morphological diagnostic characters reveals a complete overlap in pholidosis between
<taxonomicName id="4C757EC4FF92FFC4B1C5FC0DE731FBC0" authorityName="Santos &amp; Reis" authorityYear="2018" box="[1001,1077,1012,1032]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B1C5FC0DE731FBC0" bold="true" box="[1001,1077,1012,1032]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
and
<taxonomicName id="4C757EC4FF92FFC4B64DFC0DE798FBC0" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1121,1180,1012,1032]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B64DFC0DE798FBC0" bold="true" box="[1121,1180,1012,1032]" italics="true" pageId="7" pageNumber="17">L. beui</emphasis>
</taxonomicName>
(
<tableCitation id="C6F730FCFF92FFC4B684FC0CE7E9FBCF" box="[1192,1261,1012,1032]" captionStart="Table 1" captionStartId="7.[100,150,1671,1687]" captionTargetPageId="7" captionText="Table 1 Selected diagnostic characters for Liotyphlops species from the Atlantic Rainforest, sampled in this study (n = number of specimens with scale counts, * = junior synonym of Liotyphlops beui, ** =species with marginal records in Atlantic Rainforest). For raw values, refer to Supplementary Table 1." httpUri="http://table.plazi.org/id/DF0A55CFFF92FFC4B248F97FE7E3F90D" pageId="7" pageNumber="17" tableUuid="DF0A55CFFF92FFC4B248F97FE7E3F90D">Table 1</tableCitation>
). Therefore, we propose to allocate
<taxonomicName id="4C757EC4FF92FFC4B1B1FBE9E0E8FBE4" authorityName="Santos &amp; Reis" authorityYear="2018" box="[925,1004,1040,1060]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="sousai" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B1B1FBE9E0E8FBE4" bold="true" box="[925,1004,1040,1060]" italics="true" pageId="7" pageNumber="17">L. sousai</emphasis>
</taxonomicName>
as a junior synonym of
<taxonomicName id="4C757EC4FF92FFC4B6F4FBE9E611FBE4" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1240,1301,1040,1060]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B6F4FBE9E611FBE4" bold="true" box="[1240,1301,1040,1060]" italics="true" pageId="7" pageNumber="17">L. beui</emphasis>
</taxonomicName>
. We also provide a redescription for a topotypical specimen of
<taxonomicName id="4C757EC4FF92FFC4B6E1FBD4E60DFBF8" baseAuthorityName="Amaral" baseAuthorityYear="1924" box="[1229,1289,1068,1087]" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="beui" subOrder="Serpentes">
<emphasis id="B901D955FF92FFC4B6E1FBD4E60DFBF8" bold="true" box="[1229,1289,1068,1087]" italics="true" pageId="7" pageNumber="17">L. beui</emphasis>
</taxonomicName>
below.
</paragraph>
</subSubSection>
</treatment>
</document>
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