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<document id="0B787F8874E14592C5350B93576C1DA0" 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="674E87DDF74BFFEEFF3694C0CD1EBD1E" 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="Arrhyton Gunther 1858" docType="treatment" docVersion="7" lastPageNumber="20" 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="20" updateTime="1698582155577" updateUser="plazi">
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<mods:title id="3A9EDA868D05F3F1F5B594685708420E">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="1F027E919CA80C7840B81E2A1C98111B">Hedges, Blair</mods:namePart>
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<mods:namePart id="F4B5EC3490D4B39CA80BAB4FEDE6DEFD">Couloux, Arnaud</mods:namePart>
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<mods:namePart id="8AD98855E3C2491FDEA8400331455B20">Vidal, Nicolas</mods:namePart>
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<mods:date id="A8503259CB80DB9A119F546D59F38049">2009</mods:date>
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<treatment id="674E87DDF74BFFEEFF3694C0CD1EBD1E" ID-DOI="http://doi.org/10.5281/zenodo.6220101" ID-GBIF-Taxon="119614019" ID-Zenodo-Dep="6220101" LSID="urn:lsid:plazi:treatment:674E87DDF74BFFEEFF3694C0CD1EBD1E" httpUri="http://treatment.plazi.org/id/674E87DDF74BFFEEFF3694C0CD1EBD1E" lastPageNumber="20" pageId="19" pageNumber="20">
<subSubSection id="A7FD6540F74BFFEEFF3694C0CA26B8DE" box="[151,534,461,487]" pageId="19" pageNumber="20" type="nomenclature">
<paragraph id="EF5836CBF74BFFEEFF3694C0CA26B8DE" blockId="19.[151,534,461,487]" box="[151,534,461,487]" pageId="19" pageNumber="20">
<heading id="B41081A7F74BFFEEFF3694C0CA26B8DE" bold="true" box="[151,534,461,487]" fontSize="11" level="1" pageId="19" pageNumber="20" reason="1">
<emphasis id="DD93EAD9F74BFFEEFF3694C0CA26B8DE" bold="true" box="[151,534,461,487]" pageId="19" pageNumber="20">
Genus
<taxonomicName id="28E74D48F74BFFEEFF4C94C0CA26B8DE" authority="Gunther, 1858" authorityName="Gunther" authorityYear="1858" box="[237,534,461,487]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F74BFFEEFF4C94C0C96CB8DE" bold="true" box="[237,348,461,487]" italics="true" pageId="19" pageNumber="20">Arrhyton</emphasis>
<bibRefCitation id="8B764B3AF74BFFEEFEC594C0CA26B8DE" author="Gunther" box="[356,534,461,487]" pageId="19" pageNumber="24" refString="Gunther, A. (1858) Catalogue of colubrine snakes of the British Museum. British Museum, London, 281 pp." type="book" year="1858">Günther, 1858</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="A7FD6540F74BFFEEFF36971BCB21BB09" box="[151,785,534,560]" pageId="19" pageNumber="20" type="materials_examined">
<paragraph id="EF5836CBF74BFFEEFF36971BCB21BB09" blockId="19.[151,1437,534,1063]" box="[151,785,534,560]" pageId="19" pageNumber="20">
<emphasis id="DD93EAD9F74BFFEEFF36971BC909BB09" bold="true" box="[151,313,534,560]" pageId="19" pageNumber="20">
<typeStatus id="305C8869F74BFFEEFF36971BC8E4BB09" box="[151,212,534,560]" pageId="19" pageNumber="20">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="28E74D48F74BFFEEFEE1971ACB39BB09" authority="Gunther, 1858:244" authorityName="Gunther" authorityPageNumber="244" authorityYear="1858" box="[320,777,534,560]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="taeniatum">
<emphasis id="DD93EAD9F74BFFEEFEE1971ACA16BB09" box="[320,550,535,560]" italics="true" pageId="19" pageNumber="20">Arrhyton taeniatum</emphasis>
<bibRefCitation id="8B764B3AF74BFFEEFD8C971BCAE7BB09" author="Gunther" box="[557,727,534,560]" pageId="19" pageNumber="24" refString="Gunther, A. (1858) Catalogue of colubrine snakes of the British Museum. British Museum, London, 281 pp." type="book" year="1858">Günther, 1858</bibRefCitation>
:244
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="A7FD6540F74BFFEEFF649730CB48BB47" pageId="19" pageNumber="20" type="diagnosis">
<paragraph id="EF5836CBF74BFFEEFF649730CB75BB6E" blockId="19.[151,1437,534,1063]" box="[197,837,573,599]" pageId="19" pageNumber="20">
<emphasis id="DD93EAD9F74BFFEEFF649730C974BB6E" bold="true" box="[197,324,573,599]" pageId="19" pageNumber="20">Diagnosis.</emphasis>
See diagnosis for the subtribe Arrhytonina.
</paragraph>
<paragraph id="EF5836CBF74BFFEEFF64976ECB48BB47" blockId="19.[151,1437,534,1063]" box="[197,888,611,638]" pageId="19" pageNumber="20">
<emphasis id="DD93EAD9F74BFFEEFF64976EC901BB44" bold="true" box="[197,305,611,637]" pageId="19" pageNumber="20">Content.</emphasis>
Eight species are included in the genus (
<tableCitation id="A2650370F74BFFEEFCAE9769CB57BB47" box="[783,871,612,638]" 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="19" pageNumber="20">Table 1</tableCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="A7FD6540F74BFFEEFF649787CB88BB9D" box="[197,952,650,676]" pageId="19" pageNumber="20" type="distribution">
<paragraph id="EF5836CBF74BFFEEFF649787CB88BB9D" blockId="19.[151,1437,534,1063]" box="[197,952,650,676]" pageId="19" pageNumber="20">
<emphasis id="DD93EAD9F74BFFEEFF649787C955BB9D" bold="true" box="[197,357,650,676]" pageId="19" pageNumber="20">Distribution.</emphasis>
The genus is distributed throughout
<collectingCountry id="97F0765BF74BFFEEFCB59787CB62BB9D" box="[788,850,650,676]" name="Cuba" pageId="19" pageNumber="20">Cuba</collectingCountry>
(
<figureCitation id="77DC2A4EF74BFFEEFCC39787CB9ABB9D" box="[866,938,650,676]" 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="19" pageNumber="20">Fig. 2</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="A7FD6540F74BFFEEFF6497BDCD1EBD1E" pageId="19" pageNumber="20" type="discussion">
<paragraph id="EF5836CBF74BFFEEFF6497BDCD1EBD1E" blockId="19.[151,1437,534,1063]" pageId="19" pageNumber="20">
<emphasis id="DD93EAD9F74BFFEEFF6497BDC975BBF3" bold="true" box="[197,325,688,714]" pageId="19" pageNumber="20">Remarks.</emphasis>
Species of
<taxonomicName id="28E74D48F74BFFEEFE7B97BCCA7ABBF3" box="[474,586,689,714]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F74BFFEEFE7B97BCCA7ABBF3" box="[474,586,689,714]" italics="true" pageId="19" pageNumber="20">Arrhyton</emphasis>
</taxonomicName>
(
<figureCitation id="77DC2A4EF74BFFEEFDFC97BCCA97BBF2" box="[605,679,689,715]" captionStart="FIGURE 3" captionStartId="14.[151,261,1777,1801]" captionTargetBox="[174,1412,190,1725]" captionTargetId="figure@14.[151,1436,166,1753]" captionTargetPageId="14" captionText="FIGURE 3. Representatives of genera of the snake Tribe Alsophiini (Dipsadidae: Xenonodontinae). Subtribe Alsophiina: Alsophis manselli (Woodlands Spring, Montserrat), Borikenophis portoricensis (1.5 km W. Playa de Tamarindo, Puerto Rico), Cubophis cantherigerus (2.0 km W Viñales, Pinar del Rio, Cuba), Hypsirhynchus ferox (Barahona, Barahona, Dominican Republic), Ialtris dorsalis (3 km N Bois Sec, GrandAnse, Haiti), and Magliophis stahli (1.9 km NE Vista Alegre, Puerto Rico). Subtribe Arrhytonina: Arrhyton taeniatum (0.2 km WE Windmill Beach, Guantanamo Bay Naval Station, Cuba). Subtribe Uromacerina: Uromacer oxyrhynchus (4.4 km W Canada Honda, La Altagracia, Dominican Republic). Photos by S. B. Hedges." httpUri="https://zenodo.org/record/187021/files/figure.png" pageId="19" pageNumber="20">Fig. 3</figureCitation>
G) are small (
<quantity id="281F9B2EF74BFFEEFCF997BCCBF4BBF2" box="[856,964,689,715]" metricMagnitude="-1" metricUnit="m" metricValue="4.4799999999999995" pageId="19" pageNumber="20" unit="mm" value="448.0">448 mm</quantity>
, maximum SVL) racers. An informal classification of Cuban
<taxonomicName id="28E74D48F74BFFEEFE1397D5CA2DBBC8" box="[434,541,728,753]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F74BFFEEFE1397D5CA2DBBC8" box="[434,541,728,753]" italics="true" pageId="19" pageNumber="20">Arrhyton</emphasis>
</taxonomicName>
was based on color pattern and scalation, especially ventral and subcaudal counts (
<bibRefCitation id="8B764B3AF74BFFEEFF5397F3CA17BA21" author="Schwartz" box="[242,551,766,792]" pageId="19" pageNumber="26" refString="Schwartz, A. &amp; Garrido, O. H. (1981) A review of the Cuban members of the genus Arrhyton (Reptilia, Serpentes, Colubridae). Annals of the Carnegie Museum of Natural History, 50, 207 - 230." type="journal article" year="1981">Schwartz &amp; Garrido 1981</bibRefCitation>
;
<bibRefCitation id="8B764B3AF74BFFEEFD9297F3CB61BA21" author="Hedges" box="[563,849,766,792]" pageId="19" pageNumber="24" refString="Hedges, S. B. &amp; Garrido, O. H. (1992) Cuban snakes of the genus Arrhyton: two new species and a reconsideration of A. redimitum Cope. Herpetologica, 48, 168 - 177." type="journal article" year="1992">Hedges &amp; Garrido 1992</bibRefCitation>
). The most distinct species, morphologically, is
<taxonomicName id="28E74D48F74BFFEEFA2297F2C93CBA07" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="taeniatum">
<emphasis id="DD93EAD9F74BFFEEFA2297F2C93CBA07" italics="true" pageId="19" pageNumber="20">A. taeniatum</emphasis>
</taxonomicName>
, because it lacks a loreal scale, has an enlarged rostral (presumably related to semi-fossorial habits), and has a boldly striped pattern. It was placed in its own species group (
<taxonomicName id="28E74D48F74BFFEEFC549641CC5BBA5C" box="[1013,1131,844,869]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="taeniatum">
<emphasis id="DD93EAD9F74BFFEEFC549641CC5BBA5C" box="[1013,1131,844,869]" italics="true" pageId="19" pageNumber="20">taeniatum</emphasis>
</taxonomicName>
Group) and the sequence evidence (
<figureCitation id="77DC2A4EF74BFFEEFEB1967FC952BAB5" box="[272,354,882,908]" 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="19" pageNumber="20">Figs. 1</figureCitation>
and
<figureCitation id="77DC2A4EF74BFFEEFE3C967FC99EBAB5" box="[413,430,882,908]" 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="19" pageNumber="20">4</figureCitation>
), although missing
<taxonomicName id="28E74D48F74BFFEEFD36967FCB10BAB2" box="[663,800,882,907]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="ainictum">
<emphasis id="DD93EAD9F74BFFEEFD36967FCB10BAB2" box="[663,800,882,907]" italics="true" pageId="19" pageNumber="20">A. ainictum</emphasis>
</taxonomicName>
, shows it to be the sister group of all other species of
<taxonomicName id="28E74D48F74BFFEEFF369694C930BA8B" box="[151,256,921,946]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="DD93EAD9F74BFFEEFF369694C930BA8B" box="[151,256,921,946]" italics="true" pageId="19" pageNumber="20">Arrhyton</emphasis>
</taxonomicName>
. The three species having high ventral scale counts and long tails (
<taxonomicName id="28E74D48F74BFFEEFBB19694CC93BA8B" box="[1040,1187,921,946]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="dolichura">
<emphasis id="DD93EAD9F74BFFEEFBB19694CC93BA8B" box="[1040,1187,921,946]" italics="true" pageId="19" pageNumber="20">A. dolichura</emphasis>
</taxonomicName>
,
<taxonomicName id="28E74D48F74BFFEEFB109694CD73BA8B" box="[1201,1347,921,946]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="procerum">
<emphasis id="DD93EAD9F74BFFEEFB109694CD73BA8B" box="[1201,1347,921,946]" italics="true" pageId="19" pageNumber="20">A. procerum</emphasis>
</taxonomicName>
, and
<taxonomicName id="28E74D48F74BFFEEFA229694C919BAE0" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="tanyplectum">
<emphasis id="DD93EAD9F74BFFEEFA229694C919BAE0" italics="true" pageId="19" pageNumber="20">A. tanyplectum</emphasis>
</taxonomicName>
), were placed together in the
<taxonomicName id="28E74D48F74BFFEEFD3396CDCB36BAE0" box="[658,774,960,985]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="dolichura">
<emphasis id="DD93EAD9F74BFFEEFD3396CDCB36BAE0" box="[658,774,960,985]" italics="true" pageId="19" pageNumber="20">dolichura</emphasis>
</taxonomicName>
Group, also supported by DNA evidence (
<figureCitation id="77DC2A4EF74BFFEEFAAD96CDCD63BAE3" box="[1292,1363,960,986]" 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="19" pageNumber="20">Fig. 1</figureCitation>
). The remaining species, with lower ventral counts and relatively short tails, were placed in a separate group, the
<taxonomicName id="28E74D48F74BFFEEFF369100C8C7BD1F" box="[151,247,1037,1062]" class="Reptilia" family="Colubridae" genus="Arrhyton" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="vittatum">
<emphasis id="DD93EAD9F74BFFEEFF369100C8C7BD1F" box="[151,247,1037,1062]" italics="true" pageId="19" pageNumber="20">vittatum</emphasis>
</taxonomicName>
Group, though this is resolved (without maximal support) as paraphyletic in our phylogeny.
</paragraph>
</subSubSection>
</treatment>
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