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<document id="84881E271D7FCC4256AC70AC9DFD27BA" ID-DOI="10.5281/zenodo.187018" ID-GBIF-Dataset="35d81ccb-1856-4852-a5b6-d9058b3dbc0b" ID-ISSN="1175-5326" ID-Zenodo-Dep="187018" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461060371161" checkinUser="plazi" docAuthor="Hedges, Blair, Couloux, Arnaud &amp; Vidal, Nicolas" docDate="2009" docId="674E87DDF755FFF0FF36960FC910BC97" docLanguage="en" docName="zt02067p028.pdf" docOrigin="Zootaxa 2067" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Caraiba Zaher et al. 2009" docType="treatment" docVersion="7" lastPageNumber="14" masterDocId="9B77FFA5F758FFFDFFA1950DC830B939" masterDocTitle="Molecular phylogeny, classification, and biogeography of West Indian racer snakes of the Tribe Alsophiini (Squamata, Dipsadidae, Xenodontinae)" masterLastPageNumber="28" masterPageNumber="1" pageNumber="14" updateTime="1698582155577" updateUser="plazi">
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<mods:title id="1506CBA87F6F766DFC650D6B35CA3BD6">Molecular phylogeny, classification, and biogeography of West Indian racer snakes of the Tribe Alsophiini (Squamata, Dipsadidae, Xenodontinae)</mods:title>
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<mods:namePart id="6FBA0A5906FB6CD5C4BA32B67266CE99">Hedges, Blair</mods:namePart>
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<mods:namePart id="F6EA1F21C3EE9BB8C2A39DC4B63EA0AC">Couloux, Arnaud</mods:namePart>
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<mods:namePart id="265BEE74D4802D4C3C3ED1E35D8696D7">Vidal, Nicolas</mods:namePart>
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<mods:date id="C2D719B8342EED1F292F96EEF00EA96E">2009</mods:date>
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<treatment id="674E87DDF755FFF0FF36960FC910BC97" ID-DOI="http://doi.org/10.5281/zenodo.6220085" ID-GBIF-Taxon="119614015" ID-Zenodo-Dep="6220085" LSID="urn:lsid:plazi:treatment:674E87DDF755FFF0FF36960FC910BC97" httpUri="http://treatment.plazi.org/id/674E87DDF755FFF0FF36960FC910BC97" lastPageNumber="14" pageId="13" pageNumber="14">
<subSubSection id="A7FD6540F755FFF0FF36960FCA00BA25" box="[151,560,770,796]" pageId="13" pageNumber="14" type="nomenclature">
<paragraph id="EF5836CBF755FFF0FF36960FCA00BA25" blockId="13.[151,560,770,796]" box="[151,560,770,796]" pageId="13" pageNumber="14">
<heading id="B41081A7F755FFF0FF36960FCA00BA25" bold="true" box="[151,560,770,796]" fontSize="11" level="1" pageId="13" pageNumber="14" reason="1">
<emphasis id="DD93EAD9F755FFF0FF36960FCA00BA25" bold="true" box="[151,560,770,796]" pageId="13" pageNumber="14">
Genus
<taxonomicName id="28E74D48F755FFF0FF4F960FCA00BA25" authority="Zaher et al., 2009" authorityName="Zaher et al." authorityYear="2009" box="[238,560,770,796]" class="Reptilia" family="Colubridae" genus="Caraiba" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F755FFF0FF4F960FC960BA25" bold="true" box="[238,336,770,796]" italics="true" pageId="13" pageNumber="14">Caraiba</emphasis>
<bibRefCitation id="8B764B3AF755FFF0FEF6960FCA00BA25" author="Zaher" box="[343,560,770,796]" pageId="13" pageNumber="26" refString="Zaher, H., Grazziotin, F. G., Cadle, J. E., Murphy, R. W., Moura-Leite, J. C. &amp; Bonatto, S. L. (2009) Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South America xenodontines: a revised classification and descriptions of new taxa. Papeis Avulsos de Zoologia, 49,115 - 153." type="book" year="2009">Zaher et al., 2009</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="A7FD6540F755FFF0FF369642CB47BA53" box="[151,887,847,874]" pageId="13" pageNumber="14" type="materials_examined">
<paragraph id="EF5836CBF755FFF0FF369642CB47BA53" blockId="13.[151,1437,847,1454]" box="[151,887,847,874]" pageId="13" pageNumber="14">
<emphasis id="DD93EAD9F755FFF0FF369642C909BA50" bold="true" box="[151,313,847,873]" pageId="13" pageNumber="14">
<typeStatus id="305C8869F755FFF0FF369642C8E4BA50" box="[151,212,847,873]" pageId="13" pageNumber="14">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="28E74D48F755FFF0FEE1965DCB42BA53" authority="Reinhardt &amp; Lutken, 1862:214" authorityName="Reinhardt &amp; Lutken" authorityPageNumber="214" authorityYear="1862" box="[320,882,848,874]" class="Reptilia" family="Colubridae" genus="Liophis" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="andreae">
<emphasis id="DD93EAD9F755FFF0FEE1965DC9CFBA50" box="[320,511,848,873]" italics="true" pageId="13" pageNumber="14">Liophis andreae</emphasis>
<bibRefCitation id="8B764B3AF755FFF0FDA7965DCB0EBA53" author="Reinhardt" box="[518,830,848,874]" pageId="13" pageNumber="25" refString="Reinhardt, J. &amp; Lutken, C. F. (1862) Bidrag tii det vestindiske Origes og navnligen tii de dansk-vestindiske Oers Herpetologie. Vidensk. Meddel. Naturhist. For. Kjobenhavn, 153 - 291." type="book chapter" year="1862">Reinhardt &amp; Lütken, 1862</bibRefCitation>
:214
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="A7FD6540F755FFF0FF67967BCCA0BD6B" pageId="13" pageNumber="14" type="diagnosis">
<paragraph id="EF5836CBF755FFF0FF67967BCA4BBD12" blockId="13.[151,1437,847,1454]" pageId="13" pageNumber="14">
<emphasis id="DD93EAD9F755FFF0FF67967BC976BAA9" bold="true" box="[198,326,886,912]" pageId="13" pageNumber="14">Diagnosis.</emphasis>
The species in this genus has 17 midbody scale rows, 131157 ventrals, 90120 subcaudals, 1 apical scale pit, 8 upper labials, 9 lower labials, 2023 total maxillary teeth, and 2528 dentary teeth (
<tableCitation id="A2650370F755FFF0FA929690CDBDBA8E" box="[1331,1421,925,951]" captionStart="TABLE 2" captionStartId="11.[151,242,977,1001]" captionText="TABLE 2. Morphological variation in the genera of alsophiine snakes (SVL = snout-vent length), from Barbour &amp;" pageId="13" pageNumber="14">Table 2</tableCitation>
).
<taxonomicName id="28E74D48F755FFF0FF3696C9C8CCBAE4" box="[151,252,964,989]" class="Reptilia" family="Colubridae" genus="Caraiba" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F755FFF0FF3696C9C8CCBAE4" box="[151,252,964,989]" italics="true" pageId="13" pageNumber="14">Caraiba</emphasis>
</taxonomicName>
differs in at least one of these characters from all other alsophiine genera except
<taxonomicName id="28E74D48F755FFF0FB4196C9CDA4BAE4" box="[1248,1428,964,989]" class="Reptilia" family="Colubridae" genus="Hypsirhynchus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F755FFF0FB4196C9CDA4BAE4" box="[1248,1428,964,989]" italics="true" pageId="13" pageNumber="14">Hypsirhynchus</emphasis>
</taxonomicName>
. From that genus it differs in having a hemipene with enlarged papillate body calyces in the basal region and medial surface of the lobes (
<bibRefCitation id="8B764B3AF755FFF0FE43911CCA5EBD12" author="Zaher" box="[482,622,1041,1067]" pageId="13" pageNumber="26" refString="Zaher, H. (1999) Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History, 240, 1 - 168." type="journal article" year="1999">Zaher 1999</bibRefCitation>
).
</paragraph>
<paragraph id="EF5836CBF755FFF0FF67913ACCA0BD6B" blockId="13.[151,1437,847,1454]" box="[198,1168,1079,1106]" pageId="13" pageNumber="14">
<emphasis id="DD93EAD9F755FFF0FF67913AC903BD68" bold="true" box="[198,307,1079,1105]" pageId="13" pageNumber="14">Content.</emphasis>
One species (six species + subspecies) is included in the genus (
<tableCitation id="A2650370F755FFF0FB869135CC4EBD6B" box="[1063,1150,1080,1106]" captionStart="TABLE 1" captionStartId="7.[151,241,151,175]" captionText="TABLE 1. Classification of West Indian xenodontine snakes, Tribe Alsophiini. The arrangement used in this study is compared with that in previous classifications (Maglio 1970; Schwartz &amp; Henderson 1991; Powell et al. 1996)." pageId="13" pageNumber="14">Table 1</tableCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="A7FD6540F755FFF0FF679153CCA9BD41" box="[198,1177,1118,1144]" pageId="13" pageNumber="14" type="distribution">
<paragraph id="EF5836CBF755FFF0FF679153CCA9BD41" blockId="13.[151,1437,847,1454]" box="[198,1177,1118,1144]" pageId="13" pageNumber="14">
<emphasis id="DD93EAD9F755FFF0FF679153C957BD41" bold="true" box="[198,359,1118,1144]" pageId="13" pageNumber="14">Distribution.</emphasis>
The genus is distributed on
<collectingCountry id="97F0765BF755FFF0FD139153CAC3BD41" box="[690,755,1118,1144]" name="Cuba" pageId="13" pageNumber="14">Cuba</collectingCountry>
, including
<collectingCountry id="97F0765BF755FFF0FCD49153CBAFBD41" box="[885,927,1118,1144]" name="Iceland" pageId="13" pageNumber="14">Isla</collectingCountry>
de Juventud (
<figureCitation id="77DC2A4EF755FFF0FBE29153CCBBBD41" box="[1091,1163,1118,1144]" captionStart="FIGURE 2" captionStartId="11.[151,255,804,828]" captionTargetBox="[279,1307,205,780]" captionTargetId="figure@11.[279,1307,194,780]" captionTargetPageId="11" captionText="FIGURE 2. Map showing the West Indies (including the southern tip of Florida above, Central America to the west, and the northern edge of South America below) and the distributions of the genera of the Tribe Alsophiini, Subfamily Xenodontinae, Family Dipsadidae." httpUri="https://zenodo.org/record/187020/files/figure.png" pageId="13" pageNumber="14">Fig. 2</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="A7FD6540F755FFF0FF679188C910BC97" pageId="13" pageNumber="14" type="discussion">
<paragraph id="EF5836CBF755FFF0FF679188C910BC97" blockId="13.[151,1437,847,1454]" pageId="13" pageNumber="14">
<emphasis id="DD93EAD9F755FFF0FF679188C970BDA6" bold="true" box="[198,320,1157,1183]" pageId="13" pageNumber="14">Remarks.</emphasis>
The single species of
<taxonomicName id="28E74D48F755FFF0FDF49188CA87BDA7" box="[597,695,1157,1182]" class="Reptilia" family="Colubridae" genus="Caraiba" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F755FFF0FDF49188CA87BDA7" box="[597,695,1157,1182]" italics="true" pageId="13" pageNumber="14">Caraiba</emphasis>
</taxonomicName>
is a moderate-sized species of racer, occurring sympatrically with smaller and larger genera of racers on
<collectingCountry id="97F0765BF755FFF0FD3991A1CAEABDFF" box="[664,730,1196,1222]" name="Cuba" pageId="13" pageNumber="14">Cuba</collectingCountry>
. The finding here (
<figureCitation id="77DC2A4EF755FFF0FC1A91A1CC3DBDFF" box="[955,1037,1196,1222]" captionStart="FIGURE 1. A" captionStartId="10.[151,255,1833,1857]" captionTargetBox="[379,1213,185,1804]" captionTargetId="figure@10.[379,1215,185,1807]" captionTargetPageId="10" captionText="FIGURE 1. A phylogeny of Alsophiini. Bayesian tree obtained from the combined data set of six genes (RAG 2, 12 S &amp; 16 S rRNA, cytochrome b, ND 2 and ND 4; 3387 sites). Alsophis portoricensis portoricensis and A. p. anegadae have identical sequences at all genes sampled, and therefore the latter taxon is not shown. Values are ML bootstrap values above 70 % followed by Bayesian posterior probabilities above 90 %. The generic taxonomy in this tree reflects usage prior to this study and shows paraphyly and polyphyly of Alsophis (blue), Arrhyton (purple), and Antillophis (orange). See Table 1 and Figure 4 for the new classification proposed here." httpUri="https://zenodo.org/record/187019/files/figure.png" pageId="13" pageNumber="14">Figs. 1</figureCitation>
and
<figureCitation id="77DC2A4EF755FFF0FBE791A1CC66BDFF" box="[1094,1110,1196,1222]" captionStart="FIGURE 4. A" captionStartId="15.[151,255,1783,1807]" captionTargetBox="[169,1429,197,1741]" captionTargetId="figure@15.[169,1429,197,1745]" captionTargetPageId="15" captionText="FIGURE 4. A timetree of Alsophiini. Divergence times and credibility / confidence intervals are shown in Table 3. Plei = Pleistocene. The generic taxonomy in this tree reflects the new classification proposed here and detailed in Table 1." httpUri="https://zenodo.org/record/187022/files/figure.png" pageId="13" pageNumber="14">4</figureCitation>
) that “
<taxonomicName id="28E74D48F755FFF0FB0691A1CD17BDFC" box="[1191,1319,1196,1221]" class="Reptilia" family="Colubridae" genus="Antillophis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F755FFF0FB0691A1CD17BDFC" box="[1191,1319,1196,1221]" italics="true" pageId="13" pageNumber="14">Antillophis</emphasis>
</taxonomicName>
<taxonomicName id="28E74D48F755FFF0FA9C91A1C8DDBDD5" authority="(Cuba)" baseAuthorityName="Cuba" class="Reptilia" family="Colubridae" genus="Liophis" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="andreae">
<emphasis id="DD93EAD9F755FFF0FA9C91A1CDACBDFC" box="[1341,1436,1196,1221]" italics="true" pageId="13" pageNumber="14">andreae</emphasis>
(
<collectingCountry id="97F0765BF755FFF0FF3E91DFC8D4BDD5" box="[159,228,1234,1260]" name="Cuba" pageId="13" pageNumber="14">Cuba</collectingCountry>
)
</taxonomicName>
groups with large Cuban racers of another genus (see below) and that “
<taxonomicName id="28E74D48F755FFF0FBDA91DECD35BDD5" box="[1147,1285,1235,1260]" class="Reptilia" family="Colubridae" genus="Antillophis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F755FFF0FBDA91DECD35BDD5" box="[1147,1285,1235,1260]" italics="true" pageId="13" pageNumber="14">Antillophis</emphasis>
</taxonomicName>
<taxonomicName id="28E74D48F755FFF0FABA91DEC91BBC2A" authority="Hispaniola" authorityName="Hispaniola" class="Reptilia" family="Colubridae" genus="Antillophis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="parvifrons">
<emphasis id="DD93EAD9F755FFF0FABA91DECDADBDD5" box="[1307,1437,1235,1260]" italics="true" pageId="13" pageNumber="14">parvifrons</emphasis>
(Hispaniola)
</taxonomicName>
groups with Hispaniolan species (
<taxonomicName id="28E74D48F755FFF0FD6791F4CB47BC2B" box="[710,887,1273,1298]" class="Reptilia" family="Colubridae" genus="Hypsirhynchus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F755FFF0FD6791F4CB47BC2B" box="[710,887,1273,1298]" italics="true" pageId="13" pageNumber="14">Hypsirhynchus</emphasis>
</taxonomicName>
) is also supported by morphology: the former species (
<collectingCountry id="97F0765BF755FFF0FF5F902DC970BC03" box="[254,320,1312,1338]" name="Cuba" pageId="13" pageNumber="14">Cuba</collectingCountry>
) have 17 midbody scale rows whereas the latter species (Hispaniola) have 19 midbody scale rows.
<bibRefCitation id="8B764B3AF755FFF0FF47904BC9A3BC59" author="Myers" box="[230,403,1350,1376]" pageId="13" pageNumber="25" refString="Myers, C. W. (1973) A new genus for Andean snakes related to Lygophis boursieri and a new species (Colubridae). American Museum Novitates, 2522, 1 - 37." type="journal article" year="1973">Myers (1973)</bibRefCitation>
also had reservations about Maglios (1970) recognition of
<taxonomicName id="28E74D48F755FFF0FB36904ACD13BC59" box="[1175,1315,1351,1376]" class="Reptilia" family="Colubridae" genus="Antillophis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F755FFF0FB36904ACD13BC59" box="[1175,1315,1351,1376]" italics="true" pageId="13" pageNumber="14">Antillophis</emphasis>
</taxonomicName>
based on morphology.
<bibRefCitation id="8B764B3AF755FFF0FE9D9060CA16BCBE" box="[316,550,1389,1415]" pageId="13" pageNumber="26" refString="Zaher, H., Grazziotin, F. G., Cadle, J. E., Murphy, R. W., Moura-Leite, J. C. &amp; Bonatto, S. L. (2009) Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South America xenodontines: a revised classification and descriptions of new taxa. Papeis Avulsos de Zoologia, 49,115 - 153." type="book">Zaher et al. (2009)</bibRefCitation>
described
<taxonomicName id="28E74D48F755FFF0FD0F9060CB24BCBF" box="[686,788,1389,1414]" class="Reptilia" family="Colubridae" genus="Caraiba" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F755FFF0FD0F9060CB24BCBF" box="[686,788,1389,1414]" italics="true" pageId="13" pageNumber="14">Caraiba</emphasis>
</taxonomicName>
for the single species, based only on its hemipenial differences.
</paragraph>
</subSubSection>
</treatment>
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