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<document id="56A637F4D00C271B2839262AD43EA9EA" 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="0E2487E3FF84FFA8FF0E3224FBFCF894" 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="Rhinoleptus Orejas-Miranda, Roux-Esteve, and Guibe 1970" docType="treatment" docVersion="3" lastPageNumber="25" masterDocId="F21DFF9BFF9CFFB0FF993047FF95FFD5" masterDocTitle="Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squamata)" masterLastPageNumber="50" masterPageNumber="1" pageNumber="25" updateTime="1699087815032" updateUser="plazi" zenodo-license-document="CLOSED">
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<mods:title id="73C85F92955D322802BF0C535F25E0AF">Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squamata)</mods:title>
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<mods:namePart id="9815DBD1E6B8C6811842C6780AAAD333">Adalsteinsson, Solny A.</mods:namePart>
<mods:affiliation id="3F1A53AB62D2ADA855D6604B771D2EBB">Department of Biology, 208 Mueller Lab, Pennsylvania State University, University Park, PA 16802 - 5301 USA.</mods:affiliation>
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<mods:namePart id="DA4F3645FDAD5237BF667093335A5666">Branch, William R.</mods:namePart>
<mods:affiliation id="2331648A5409843F9839134B25798CCD">Bayworld, P. O. Box 13147, Humewood 6013, South Africa</mods:affiliation>
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<mods:namePart id="61FA7A6A3626D7F7048C85ADC78EFDB1">Trape, Sébastien</mods:namePart>
<mods:affiliation id="EB888F12B03DF2E6713C9AA712B3EB4B">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="605DFE08057F97FD099B2AAA08CDDCED">Vitt, Laurie J.</mods:namePart>
<mods:affiliation id="E10278CCF0CE621571C8B48FD5DB9D30">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="5DCB067C578F2069F543A1F349610856">Hedges, S. Blair</mods:namePart>
<mods:affiliation id="110A34E8C5CB4F17BC480002C6F43710">Department of Biology, 208 Mueller Lab, Pennsylvania State University, University Park, PA 16802 - 5301 USA.</mods:affiliation>
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<paragraph id="863236F5FF84FFA8FF0E3224FC4EFDA8" blockId="24.[151,987,611,638]" box="[151,987,611,638]" pageId="24" pageNumber="25">
<heading id="DD7A8199FF84FFA8FF0E3224FC4EFDA8" bold="true" box="[151,987,611,638]" fontSize="11" level="1" pageId="24" pageNumber="25" reason="1">
<emphasis id="B4F9EAE7FF84FFA8FF0E3224FC4EFDA8" bold="true" box="[151,987,611,638]" pageId="24" pageNumber="25">
Genus
<taxonomicName id="418D4D76FF84FFA8FF743223FC4EFDA8" authority="Orejas-Miranda, Roux-Esteve, and Guibe, 1970" authorityName="Orejas-Miranda, Roux-Esteve, and Guibe" authorityYear="1970" box="[237,987,611,638]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF84FFA8FF743223FE15FDAB" bold="true" box="[237,384,612,638]" italics="true" pageId="24" pageNumber="25">Rhinoleptus</emphasis>
<bibRefCitation id="E21C4B04FF84FFA8FE113224FC4EFDA8" author="Orejas-Miranda, Roux-Esteve, and Guibe" box="[392,987,611,637]" firstAuthor="Orejas-Miranda" pageId="24" pageNumber="25" pagination="1 - 4" refId="ref26293" refString="Orejas-Miranda, B. R., Roux-Esteve, M. R. &amp; Guibe, J. (1970) Un nouveau genre de Leptotyphlopides (Ophidia) Rhinoleptus koniagui (Villiers). Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo, 10, 1 - 4." type="journal article" year="1970">Orejas-Miranda, Roux-Estève, and Guibé, 1970</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF84FFA8FF0E32E1FCB0FD68" box="[151,805,678,702]" pageId="24" pageNumber="25" type="type_taxon">
<paragraph id="863236F5FF84FFA8FF0E32E1FCB0FD68" blockId="24.[151,805,678,702]" box="[151,805,678,702]" pageId="24" pageNumber="25">
<emphasis id="B4F9EAE7FF84FFA8FF0E32E1FEBEFD6B" bold="true" box="[151,299,678,702]" pageId="24" pageNumber="25">
<typeStatus id="59368857FF84FFA8FF0E32E1FF5AFD6B" box="[151,207,678,702]" pageId="24" pageNumber="25">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="418D4D76FF84FFA8FEA832E1FD12FD68" authority="Villers, 1956" authorityName="Villers" authorityYear="1956" box="[305,647,678,701]" class="Reptilia" family="Typhlopidae" genus="Typhlops" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="koniagui">
<emphasis id="B4F9EAE7FF84FFA8FEA832E1FE61FD68" box="[305,500,678,701]" italics="true" pageId="24" pageNumber="25">Typhlops koniagui</emphasis>
Villers, 1956
</taxonomicName>
, by monotypy.
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF84FFA8FF0E32AAFB9EFBC3" pageId="24" pageNumber="25" type="diagnosis">
<paragraph id="863236F5FF84FFA8FF0E32AAFB3FFC3A" blockId="24.[151,1436,749,1858]" pageId="24" pageNumber="25">
<emphasis id="B4F9EAE7FF84FFA8FF0E32AAFE82FCD2" bold="true" box="[151,279,749,775]" pageId="24" pageNumber="25">Diagnosis.</emphasis>
Species in this genus have 16 midbody scale rows, 14 midtail scale rows, 302546 middorsal scale rows, 2130 subcaudals, 24 supralabials, small anterior supralabials,
<quantity id="41759B10FF84FFA8FC6C3353FB00FCFB" box="[1013,1173,788,814]" metricMagnitude="-1" metricUnit="m" metricValue="3.1" metricValueMax="4.6" metricValueMin="1.6" pageId="24" pageNumber="25" unit="mm" value="310.0" valueMax="460.0" valueMin="160.0">160460 mm</quantity>
maximum adult total length, a body shape of 67160 (total length/width), a relative tail length of 3.710.0 %, a tail shape of 3.5, no striped pattern, a brown dorsum, and brown venter (Table 2). They are distinguished from the other genus in this tribe,
<taxonomicName id="418D4D76FF84FFA8FE8F33CFFEFAFC74" box="[278,367,904,929]" class="Reptilia" family="Leptotyphlopidae" genus="Guinea" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF84FFA8FE8F33CFFEFAFC74" box="[278,367,904,929]" italics="true" pageId="24" pageNumber="25">Guinea</emphasis>
</taxonomicName>
, by having 16 midbody scale rows (versus 14), 14 midtail rows (versus 12), 302546 middorsal rows (versus 173288), 2130 subcaudals (versus 616), and a body shape of 67160 (versus 24 69.2). Only one species was included in the molecular phylogenetic analyses (
<figureCitation id="1EB62A70FF84FFA8FBB33392FB09FC3A" box="[1066,1180,981,1007]" 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="24" pageNumber="25">Figs. 34</figureCitation>
).
</paragraph>
<paragraph id="863236F5FF84FFA8FF5C33BCFB9EFBC3" blockId="24.[151,1436,749,1858]" box="[197,1035,1019,1046]" pageId="24" pageNumber="25">
<emphasis id="B4F9EAE7FF84FFA8FF5C33BCFEA4FBC0" bold="true" box="[197,305,1019,1045]" pageId="24" pageNumber="25">Content.</emphasis>
Two species (
<tableCitation id="CB0F034EFF84FFA8FE4E33BBFDA5FBC3" box="[471,560,1020,1046]" 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="24" pageNumber="25">Table 1</tableCitation>
;
<figureCitation id="1EB62A70FF84FFA8FDA633BBFD10FBC3" box="[575,645,1020,1046]" captionStart="FIGURE 9" captionStartId="25.[151,257,1600,1624]" captionTargetBox="[151,1436,194,1576]" captionTargetId="figure-169@25.[151,1436,194,1576]" captionTargetPageId="25" captionText="FIGURE 9. Representatives of the snake Family Leptotyphlopidae from the Old World. (A) Guinea bicolor (Mali; Sikasso; Doussoudiana); photograph by Sébastien Trape. (B) Rhinoleptus koniagui (Senegal: Tambacounda; Ibel), preserved specimen from Sébastien Trape; photograph by S. Blair Hedges. (C) Myriopholis boueti (Sénégal; Dakar; Dakar); photograph by Sébastien Trape. (D) Myriopholis longicauda (South Africa: Northern Province; Limpopo); photograph by William R. Branch. (E) Leptotyphlops distanti (South Africa: Mpumalanga: near Middleburg); photograph by William R. Branch. (F) Leptotyphlops incognitus (South Africa: Mpumalanga: Komati River); photograph by William R. Branch." pageId="24" pageNumber="25">Fig. 9</figureCitation>
), although see &quot;Remarks&quot; below.
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF84FFA8FF5C3465FF51FB5F" pageId="24" pageNumber="25" type="distribution">
<paragraph id="863236F5FF84FFA8FF5C3465FF51FB5F" blockId="24.[151,1436,749,1858]" pageId="24" pageNumber="25">
<emphasis id="B4F9EAE7FF84FFA8FF5C3465FEFDFBE9" bold="true" box="[197,360,1058,1084]" pageId="24" pageNumber="25">Distribution.</emphasis>
<taxonomicName id="418D4D76FF84FFA8FEE83465FD95FBEE" authorityName="Orejas-Miranda, Roux-Esteve, and Guibe" authorityYear="1970" box="[369,512,1058,1083]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF84FFA8FEE83465FD95FBEE" box="[369,512,1058,1083]" italics="true" pageId="24" pageNumber="25">Rhinoleptus</emphasis>
</taxonomicName>
is distributed in West Africa (
<taxonomicName id="418D4D76FF84FFA8FCF63465FBFBFBEE" authorityName="Villiers" authorityYear="1956" box="[879,1134,1058,1083]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="koniagui">
<emphasis id="B4F9EAE7FF84FFA8FCF63465FBFBFBEE" box="[879,1134,1058,1083]" italics="true" pageId="24" pageNumber="25">Rhinoleptus koniagui</emphasis>
</taxonomicName>
), including
<collectingCountry id="FE9A7665FF84FFA8FA9A3465FAF1FBE9" box="[1283,1380,1058,1084]" name="Senegal" pageId="24" pageNumber="25">Senegal</collectingCountry>
, and
<taxonomicName id="418D4D76FF84FFA8FF0E340EFF65FBB6" box="[151,240,1097,1123]" class="Reptilia" family="Leptotyphlopidae" genus="Guinea" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="46" phylum="Chordata" rank="genus">Guinea</taxonomicName>
, and
<collectingCountry id="FE9A7665FF84FFA8FEA9340EFEFDFBB6" box="[304,360,1097,1123]" name="Serbia" pageId="24" pageNumber="25">Mali</collectingCountry>
(
<bibRefCitation id="E21C4B04FF84FFA8FEE1340EFDE7FBB6" author="Trape &amp; Mane" box="[376,626,1097,1123]" firstAuthor="Trape" pageId="24" pageNumber="25" refId="ref28237" refString="Trape, J. - F. &amp; Mane, Y. (2006) Guide des serpentes d'Afrique occidentale: savane et desert. IRD Editions, Paris, 226 pp." type="book" year="2006">Trape &amp; Mané 2006</bibRefCitation>
); and in East Africa (
<taxonomicName id="418D4D76FF84FFA8FCED340EFBF6FBB7" authorityName="EAF" baseAuthorityName="Broadley" baseAuthorityYear="1999" box="[884,1123,1097,1122]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="parkeri">
<emphasis id="B4F9EAE7FF84FFA8FCED340EFBF6FBB7" box="[884,1123,1097,1122]" italics="true" pageId="24" pageNumber="25">Rhinoleptus parkeri</emphasis>
</taxonomicName>
), including
<collectingCountry id="FE9A7665FF84FFA8FB6C340EFACFFBB6" box="[1269,1370,1097,1123]" name="Ethiopia" pageId="24" pageNumber="25">Ethiopia</collectingCountry>
(
<figureCitation id="1EB62A70FF84FFA8FAF5340EFF23FB5F" captionStart="FIGURE 11" captionStartId="29.[151,257,1371,1395]" captionTargetBox="[151,1428,180,1359]" captionTargetId="figure-270@29.[151,1436,174,1359]" captionTargetPageId="29" captionText="FIGURE 11. Distributions of genera of leptotyphlopid snakes in the Old World. (A) Guinea and Leptotyphlops. (B) Rhinoleptus. (C) Epacrophis and Namibiana. (D) Myriopholis." pageId="24" pageNumber="25">Fig. 11</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF84FFA8FF5C34D1FEEAFB2B" pageId="24" pageNumber="25" type="etymology">
<paragraph id="863236F5FF84FFA8FF5C34D1FEEAFB2B" blockId="24.[151,1436,749,1858]" pageId="24" pageNumber="25">
<emphasis id="B4F9EAE7FF84FFA8FF5C34D1FED8FB65" bold="true" box="[197,333,1174,1200]" pageId="24" pageNumber="25">Etymology</emphasis>
. The generic name is masculine and derived from the Greek noun
<emphasis id="B4F9EAE7FF84FFA8FBEE34D0FB56FB65" box="[1143,1219,1175,1200]" italics="true" pageId="24" pageNumber="25">rhinos</emphasis>
(nose) and Greek adjective
<emphasis id="B4F9EAE7FF84FFA8FE9234FAFEC7FB03" box="[267,338,1213,1238]" italics="true" pageId="24" pageNumber="25">leptos</emphasis>
(thin), in allusion to the unusual rostral scale of
<emphasis id="B4F9EAE7FF84FFA8FC3B34FAFB3FFB03" box="[930,1194,1213,1238]" italics="true" pageId="24" pageNumber="25">
<taxonomicName id="418D4D76FF84FFA8FC3B34FAFB30FB03" authorityName="Villiers" authorityYear="1956" box="[930,1189,1213,1238]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="koniagui">Rhinoleptus koniagui</taxonomicName>
,
</emphasis>
with its narrow and pointed anterior tip.
</paragraph>
</subSubSection>
<subSubSection id="CE97657EFF84FFA8FF5C354DFBFCF894" pageId="24" pageNumber="25" type="discussion">
<paragraph id="863236F5FF84FFA8FF5C354DFC18F955" blockId="24.[151,1436,749,1858]" pageId="24" pageNumber="25">
<emphasis id="B4F9EAE7FF84FFA8FF5C354DFEABFAF1" bold="true" box="[197,318,1290,1316]" pageId="24" pageNumber="25">Remarks.</emphasis>
We were unable to obtain a tissue sample of
<taxonomicName id="418D4D76FF84FFA8FCD6354CFBADFAF1" authorityName="EAF" baseAuthorityName="Broadley" baseAuthorityYear="1999" box="[847,1080,1291,1316]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="parkeri">
<emphasis id="B4F9EAE7FF84FFA8FCD6354CFBADFAF1" box="[847,1080,1291,1316]" italics="true" pageId="24" pageNumber="25">Rhinoleptus parkeri</emphasis>
</taxonomicName>
but assign it here to the genus
<taxonomicName id="418D4D76FF84FFA8FF0E3576FEB6FA9F" authorityName="Orejas-Miranda, Roux-Esteve, and Guibe" authorityYear="1970" box="[151,291,1329,1354]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF84FFA8FF0E3576FEB6FA9F" box="[151,291,1329,1354]" italics="true" pageId="24" pageNumber="25">Rhinoleptus</emphasis>
</taxonomicName>
because it shares with
<taxonomicName id="418D4D76FF84FFA8FDA93576FD22FA9F" authorityName="Villiers" authorityYear="1956" box="[560,695,1329,1354]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="koniagui">
<emphasis id="B4F9EAE7FF84FFA8FDA93576FD22FA9F" box="[560,695,1329,1354]" italics="true" pageId="24" pageNumber="25">R. koniagui</emphasis>
</taxonomicName>
a series of unique or rare traits in the family: an unusually high number of midbody scale rows (16) and midtail scale rows (14), parietals small or undifferentiated, and occipitals undifferentiated. In his description of
<taxonomicName id="418D4D76FF84FFA8FD603538FBA6FA4D" authority=", Broadley (1999)" authorityName=", Broadley" authorityYear="1999" box="[761,1075,1406,1432]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="parkeri">
<emphasis id="B4F9EAE7FF84FFA8FD603538FCC1FA4D" box="[761,852,1407,1432]" italics="true" pageId="24" pageNumber="25">parkeri</emphasis>
,
<bibRefCitation id="E21C4B04FF84FFA8FCFF3539FBA6FA4D" author="Broadley" box="[870,1075,1406,1432]" firstAuthor="Broadley" pageId="24" pageNumber="25" pagination="1 - 36" refId="ref22972" refString="Broadley, D. G. &amp; Broadley, S. (1999) A review of the African wormsnakes from south of latitude 12 ° S (Serpentes: Leptotyphlopidae). Syntarsus, 5, 1 - 36." type="journal article" year="1999">Broadley (1999)</bibRefCitation>
</taxonomicName>
considered these traits to be ancestral assuming that all other leptotyphlopids (apart from
<taxonomicName id="418D4D76FF84FFA8FC1E35E2FB81FA6B" authorityName="Villiers" authorityYear="1956" box="[903,1044,1445,1470]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="koniagui">
<emphasis id="B4F9EAE7FF84FFA8FC1E35E2FB81FA6B" box="[903,1044,1445,1470]" italics="true" pageId="24" pageNumber="25">R. koniagui</emphasis>
</taxonomicName>
) formed a monophyletic group.
<bibRefCitation id="E21C4B04FF84FFA8FF0E358BFED9FA33" author="Wallach" box="[151,332,1484,1510]" firstAuthor="Wallach" pageId="24" pageNumber="25" 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>
also found that
<taxonomicName id="418D4D76FF84FFA8FD90358BFDCAFA30" authorityName=", Broadley" authorityYear="1999" box="[521,607,1484,1509]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="parkeri">
<emphasis id="B4F9EAE7FF84FFA8FD90358BFDCAFA30" box="[521,607,1484,1509]" italics="true" pageId="24" pageNumber="25">parkeri</emphasis>
</taxonomicName>
branched early in the tree based largely on visceral characters, and the position of this species was discussed further by
<bibRefCitation id="E21C4B04FF84FFA8FD7435B5FBC2F9D9" author="Broadley and Wallach" box="[749,1111,1522,1548]" firstAuthor="Broadley" pageId="24" pageNumber="25" pagination="3 - 78" refId="ref23147" refString="Broadley, D. G. &amp; Wallach, V. (2007) A revision of the genus Leptotyphlops in northeastern Africa and southwestern Arabia (Serpentes: Leptotyphlopidae). Zootaxa, 3 - 78." type="book chapter" year="2007">Broadley and Wallach (2007)</bibRefCitation>
. However, considering the phylogenetic relationships obtained in our study (
<figureCitation id="1EB62A70FF84FFA8FC99365EFCE3F9E6" box="[768,886,1561,1587]" 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="24" pageNumber="25">Figs. 34</figureCitation>
) showing that
<taxonomicName id="418D4D76FF84FFA8FBA9365EFB57F9E7" authorityName="Orejas-Miranda, Roux-Esteve, and Guibe" authorityYear="1970" box="[1072,1218,1561,1586]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF84FFA8FBA9365EFB57F9E7" box="[1072,1218,1561,1586]" italics="true" pageId="24" pageNumber="25">Rhinoleptus</emphasis>
</taxonomicName>
is not the closest relative of all other leptotyphlopids, those characteristics of
<taxonomicName id="418D4D76FF84FFA8FCC53607FC46F98C" authorityName=", Broadley" authorityYear="1999" box="[860,979,1600,1625]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="parkeri">
<emphasis id="B4F9EAE7FF84FFA8FCC53607FC46F98C" box="[860,979,1600,1625]" italics="true" pageId="24" pageNumber="25">R. parkeri</emphasis>
</taxonomicName>
are now re-evaluated as being derived within
<taxonomicName id="418D4D76FF84FFA8FF703621FEE9F955" box="[233,380,1638,1664]" class="Reptilia" family="Leptotyphlopidae" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="tribe" tribe="Rhinoleptini">Rhinoleptini</taxonomicName>
rather than ancestral among leptotyphlopids.
</paragraph>
<paragraph id="863236F5FF84FFA8FF5C36CAFBFCF894" blockId="24.[151,1436,749,1858]" pageId="24" pageNumber="25">
The specimen of
<taxonomicName id="418D4D76FF84FFA8FE0336CAFDBEF973" authorityName="Orejas-Miranda, Roux-Esteve, and Guibe" authorityYear="1970" box="[410,555,1677,1702]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis id="B4F9EAE7FF84FFA8FE0336CAFDBEF973" box="[410,555,1677,1702]" italics="true" pageId="24" pageNumber="25">Rhinoleptus</emphasis>
</taxonomicName>
from West Africa sampled here (
<figureCitation id="1EB62A70FF84FFA8FC5E36CAFBB0F972" box="[967,1061,1677,1703]" captionStart="FIGURE 9" captionStartId="25.[151,257,1600,1624]" captionTargetBox="[151,1436,194,1576]" captionTargetId="figure-169@25.[151,1436,194,1576]" captionTargetPageId="25" captionText="FIGURE 9. Representatives of the snake Family Leptotyphlopidae from the Old World. (A) Guinea bicolor (Mali; Sikasso; Doussoudiana); photograph by Sébastien Trape. (B) Rhinoleptus koniagui (Senegal: Tambacounda; Ibel), preserved specimen from Sébastien Trape; photograph by S. Blair Hedges. (C) Myriopholis boueti (Sénégal; Dakar; Dakar); photograph by Sébastien Trape. (D) Myriopholis longicauda (South Africa: Northern Province; Limpopo); photograph by William R. Branch. (E) Leptotyphlops distanti (South Africa: Mpumalanga: near Middleburg); photograph by William R. Branch. (F) Leptotyphlops incognitus (South Africa: Mpumalanga: Komati River); photograph by William R. Branch." pageId="24" pageNumber="25">Fig. 9B</figureCitation>
) agrees in many respects with
<taxonomicName id="418D4D76FF84FFA8FF0E36F3FE04F918" authorityName="Villiers" authorityYear="1956" box="[151,401,1716,1741]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="koniagui">
<emphasis id="B4F9EAE7FF84FFA8FF0E36F3FE04F918" box="[151,401,1716,1741]" italics="true" pageId="24" pageNumber="25">Rhinoleptus koniagui</emphasis>
</taxonomicName>
(e.g., greatly enlarged rostral, 16 scale rows, oblique orientation of head scales,
<bibRefCitation id="E21C4B04FF84FFA8FADF36F3FF4CF921" author="Villiers" firstAuthor="Villiers" pageId="24" pageNumber="25" pagination="143 - 162" refId="ref28835" refString="Villiers, A. (1956) Le Parc National du Niokolo-Koba. V. Reptiles. Memoires de L'Institut Francais D'Afrique Noire, 48, 143 - 162." type="journal article" year="1956">Villiers 1956</bibRefCitation>
). However, it and some other specimens from
<collectingCountry id="FE9A7665FF84FFA8FC8E369DFCE2F921" box="[791,887,1754,1780]" name="Senegal" pageId="24" pageNumber="25">Senegal</collectingCountry>
lack the distinctive horn on the rostral of
<taxonomicName id="418D4D76FF84FFA8FA1B369CFF6AF8CF" authorityName="Villiers" authorityYear="1956" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="koniagui">
<emphasis id="B4F9EAE7FF84FFA8FA1B369CFF6AF8CF" italics="true" pageId="24" pageNumber="25">R. koniagui</emphasis>
</taxonomicName>
(Hedges and Trape, unpub. obs.). We conservatively refer it to
<taxonomicName id="418D4D76FF84FFA8FC673746FB69F8CF" authorityName="Villiers" authorityYear="1956" box="[1022,1276,1793,1818]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="koniagui">
<emphasis id="B4F9EAE7FF84FFA8FC673746FB69F8CF" box="[1022,1276,1793,1818]" italics="true" pageId="24" pageNumber="25">Rhinoleptus koniagui</emphasis>
</taxonomicName>
but note that additional material may signal the presence of an additional species of
<emphasis id="B4F9EAE7FF84FFA8FC4C376FFBFCF894" box="[981,1129,1832,1857]" italics="true" pageId="24" pageNumber="25">
<taxonomicName id="418D4D76FF84FFA8FC4C376FFBF0F894" authorityName="Orejas-Miranda, Roux-Esteve, and Guibe" authorityYear="1970" box="[981,1125,1832,1857]" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">Rhinoleptus</taxonomicName>
.
</emphasis>
</paragraph>
</subSubSection>
</treatment>
</document>