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<document id="BD5C151575E6FF98684620CC5A15C093" ID-CLB-Dataset="34335" ID-DOI="http://dx.doi.org/10.3897/zookeys.244.3950" ID-GBIF-Dataset="4ed119c9-f77a-4521-a117-841f749ba9c0" ID-PMC="PMC3520105" ID-Pensoft-Pub="1313-2970-244-91" ID-PubMed="23275746" ModsDocAuthor="" ModsDocDate="2012" ModsDocID="1313-2970-244-91" ModsDocOrigin="ZooKeys 244" ModsDocTitle="A new species of blunt-headed vine snake (Colubridae, Imantodes) from the Chocó region of Ecuador" checkinTime="1451248523380" checkinUser="pensoft" docAuthor="Torres-Carvajal, Omar, Yanez-Munoz, Mario H., Quirola, Diego, Eric N. Smith, &amp; Almendariz, Ana" docDate="2012" docId="00CB79DAC4704A709F94482EC62B4F20" docLanguage="en" docName="ZooKeys 244: 91-110" docOrigin="ZooKeys 244" docSource="http://dx.doi.org/10.3897/zookeys.244.3950" docTitle="Imantodes chocoensis Torres-Carvajal, Yanez-Munoz, Quirola, Smith &amp; Almendariz, 2012, sp. n." docType="treatment" docVersion="5" lastPageNumber="99" masterDocId="2A34FF2B8737240AFFFDE623C1264B51" masterDocTitle="A new species of blunt-headed vine snake (Colubridae, Imantodes) from the Choco region of Ecuador" masterLastPageNumber="110" masterPageNumber="91" pageNumber="93" updateTime="1732940178040" updateUser="ExternalLinkService">
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<mods:title id="BCF4DCA63EDDC8B82ACAE4145BB82507">A new species of blunt-headed vine snake (Colubridae, Imantodes) from the Choco region of Ecuador</mods:title>
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<mods:namePart id="DC5F606E66FEB3AD662A6FBC52C1B253">Torres-Carvajal, Omar</mods:namePart>
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<mods:namePart id="A324113340728D7CD3D3155816B4930C">Yanez-Munoz, Mario H.</mods:namePart>
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<mods:namePart id="519F253EB86954AF1904521ACA773C17">Quirola, Diego</mods:namePart>
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<mods:namePart id="D61DF402EDD2DEEF33F380420787DFC6">Eric N. Smith,</mods:namePart>
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<mods:namePart id="DDA79822F8345FD18F305743F86B4FC4">Almendariz, Ana</mods:namePart>
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<treatment id="00CB79DAC4704A709F94482EC62B4F20" ID-GBIF-Taxon="152038051" LSID="urn:lsid:plazi:treatment:00CB79DAC4704A709F94482EC62B4F20" httpUri="http://treatment.plazi.org/id/00CB79DAC4704A709F94482EC62B4F20" lastPageId="8" lastPageNumber="99" pageId="2" pageNumber="93" scope_family="Colubridae">
<subSubSection id="7D675ABEC8D0D0C0C905DFA7D80FA1FF" pageId="2" pageNumber="93" type="nomenclature">
<paragraph id="526ADBAFEA6B50E09804B3B9611C584A" pageId="2" pageNumber="93">
<taxonomicName id="B9307A0269C22D4962E6B7147B78256D" ID-CoL="3PHNH" LSID="Proposed standard English name: Chocoan blunt-headed vine snakes" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="chocoensis">
<pageBreakToken id="ABCD9B3E6A5671EA4A97EDEEE9BFB010" pageId="2" pageNumber="93" start="start">Imantodes</pageBreakToken>
chocoensis
</taxonomicName>
<taxonomicNameLabel id="7806608BF436F094B0786A4EF9561FD0" pageId="2" pageNumber="93">sp. n.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="8D16CB18D6D2851B095D8B0923E31D3F" pageId="2" pageNumber="93" type="holotype">
<paragraph id="ACD85240804B47F2D8592BCF98D83079" pageId="2" pageNumber="93">Holotype.</paragraph>
<paragraph id="750299E246EE8A129F86748A624DF730" pageId="2" pageNumber="93">
- QCAZ 7984 (Figs. 1, 2), an adult male from 4 km N Durango,
<geoCoordinate id="6DC4349C5510ECDFD1665688138927C4" direction="north" orientation="latitude" precision="5" value="1.0283">1.0283°N</geoCoordinate>
,
<geoCoordinate id="7D7FC8EDD34560004B5858ABC159C394" direction="west" orientation="longitude" precision="5" value="-78.595">78.5950°W</geoCoordinate>
(DD), 253 m, Provincia Esmeraldas, Ecuador, collected on 24 April 2007 by E. Carrillo-Ponce, I. G. Tapia, and E. E.Tapia.
</paragraph>
</subSubSection>
<subSubSection id="B35DBBAA2DA2291C0C455408EDE71E54" pageId="2" pageNumber="93" type="paratypes (6)">
<paragraph id="95B2324BC71263935EB9528CBE78594D" pageId="2" pageNumber="93">Paratypes (6).</paragraph>
<paragraph id="E4B6B35512375D3DAD5CE4D48FAA6E3E" pageId="2" pageNumber="93">
- ECUADOR: Provincia Carchi: DHMECN 6753,
<normalizedToken id="F345F57B0E71B6AC14AE6E00E796CF16" originalValue="Río">Rio</normalizedToken>
San Juan,
<geoCoordinate id="C86CBD1170208E5F6953A941773FC6CE" direction="north" orientation="latitude" precision="5" value="1.1858">1.1858°N</geoCoordinate>
,
<geoCoordinate id="0FB4FB7702F735F6739A69A83FD43844" direction="west" orientation="longitude" precision="5" value="-78.5006">78.5006°W</geoCoordinate>
(DD), 243 m, collected on 12 September 2009 by M.
<normalizedToken id="C4FB32B4347B915C8B8009A6D31022AE" originalValue="Yánez-Muñoz">Yanez-Munoz</normalizedToken>
, L. Oyagata, and M. Altamirano; DHMECN 6757, Sendero Awa,
<geoCoordinate id="E3E8C03266F8CD0DDCC9E499AE99FCA8" direction="north" orientation="latitude" precision="5" value="1.1643">1.1643°N</geoCoordinate>
,
<geoCoordinate id="2647538C897245D8887ABEA46F3944B5" direction="west" orientation="longitude" precision="5" value="-78.5071">78.5071°W</geoCoordinate>
(DD), 257 m, collected on 16 September 2009 by M.
<normalizedToken id="097AE214A302EA440C6CF67677238EEC" originalValue="Yánez-Muñoz">Yanez-Munoz</normalizedToken>
, L. Oyagata, and M. Altamirano. Provincia Esmeraldas: UTA R-60205, San Lorenzo-Santa Rita,
<geoCoordinate id="A5F3C3A7DF572E61385174376E820E25" direction="north" orientation="latitude" precision="5" value="1.0321">1.0321°N</geoCoordinate>
,
<geoCoordinate id="BE238A3376E9E9D2A48BB23F72121480" direction="west" orientation="longitude" precision="5" value="-78.7138">78.7138°W</geoCoordinate>
(DD), 115 m, collected on 21 March 2008 by M. Alcoser, R. Betancourt, P. Loaiza L., L. Oyagata, S.
<normalizedToken id="C483EF0D536318FE4BE894F2C8E78E30" originalValue="Ramírez">Ramirez</normalizedToken>
J., J. W. Streicher, C. Tobar, and E. N. Smith; QCAZ 7978, same collection data as holotype; QCAZ 10185, 4 km W Alto Tambo,
<geoCoordinate id="1A434B341AA4000404F4565F33B5CEB9" direction="north" orientation="latitude" precision="1" value="0.91241">0.91241°N</geoCoordinate>
,
<geoCoordinate id="050619E63A57C0700F5723B2557D56A8" direction="west" orientation="longitude" precision="5" value="-78.5809">78.5809°W</geoCoordinate>
(DD), collected on 18 December 2009 by S. Poe, L. Gray, and I. Latella; QCAZ 10710, Playa de Oro, Estero Pote and Estero Angostura, lower part of Cotacachi Cayapas Ecological Reserve,
<geoCoordinate id="E776F0AA8F74B569B064792DC386574F" direction="north" orientation="latitude" precision="5" value="0.8285">0.8285°N</geoCoordinate>
,
<geoCoordinate id="DCEB490EB18EA4B30405968958775EF8" direction="west" orientation="longitude" precision="5" value="-78.722">78.7220°W</geoCoordinate>
(DD), collected on 27 November 1994 by E. Toral-Contreras, V. Ortiz, and F. Nogales.
</paragraph>
</subSubSection>
<subSubSection id="174F7466E115E4369A6DB24E8AB83F22" pageId="2" pageNumber="93" type="diagnosis">
<paragraph id="DB154E1E2A8A82F823764C365CFE9F6C" pageId="2" pageNumber="93">Diagnosis.</paragraph>
<paragraph id="40D6444E8A58CE1FBAAADD047DF07311" pageId="2" pageNumber="93">
<taxonomicName id="1A81893B33E627C63BB6F14318B4C0AB" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
differs from all other known congeners in lacking a loreal scale. It can be further distinguished from its sister species (see Phylogenetic relationships)
<taxonomicName id="CD07F70E1D84B590897C31DFD340D75F" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes lentiferus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="lentiferus">Imantodes lentiferus</taxonomicName>
by having 17 longitudinal rows of dorsal scales at midbody and at nearly one head length anterior to the cloaca (15 in
<taxonomicName id="3157133735ABA65AF6E0A3F3EF15ED8C" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes lentiferus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="lentiferus">Imantodes lentiferus</taxonomicName>
), more ventrals (t = 7.27, P &lt;0.001), more subcaudals (t = -4.31, P &lt;0.001), more postoculars (2-3, mean = 2.43
<normalizedToken id="10A45710395C6BC122552FE0A45442A6" originalValue="±">+/-</normalizedToken>
0.51; 1-2, mean = 1.81
<normalizedToken id="DE265DEE2798ACA26E905AE95B7D0110" originalValue="±">+/-</normalizedToken>
0.39 in
<taxonomicName id="E64AEDE0A46B8138CB13B32B48AB1F61" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes lentiferus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="lentiferus">Imantodes lentiferus</taxonomicName>
), more infralabials (12-15, mean = 13.21
<normalizedToken id="8CB2F343A833948788855C3A7C5BA783" originalValue="±">+/-</normalizedToken>
0.80; 9-12, mean = 10.68
<normalizedToken id="86BFFE1142BDE2D4F372DFEE28875B01" originalValue="±">+/-</normalizedToken>
0.60 in
<taxonomicName id="38860A7D8E157AD4A650D3B24A8E6A38" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes lentiferus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="lentiferus">Imantodes lentiferus</taxonomicName>
), and smaller dark blotches on dorsum (Fig. 3). Among other species of
<taxonomicName id="054FBE620C140E12CC05C8BF293981CA" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes" pageId="2" pageNumber="93" phylum="Chordata" rank="genus">Imantodes</taxonomicName>
known from Ecuador, the new species differs further from
<taxonomicName id="3D56B27B50FE5511977EDCB1FC969C3E" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes inornatus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="inornatus">Imantodes inornatus</taxonomicName>
(N = 2-3) in having more ventrals (t = 6.74, P &lt;0.001), more subcaudals (t = -5.05, P = 0.002), more infralabials (9-11, mean 10.00
<normalizedToken id="697CE11A6020A59BD8029670F489FE4F" originalValue="±">+/-</normalizedToken>
0.89 in
<taxonomicName id="EC70750F2CD0DF809FF597B98C4191FF" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes inornatus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="inornatus">Imantodes inornatus</taxonomicName>
), a longer head (head length/width 1.54-1.71, mean = 1.63
<normalizedToken id="15E87C0D3D5C76EFD2C4A72205FF324C" originalValue="±">+/-</normalizedToken>
0.07 in
<taxonomicName id="EB4D489A666B03415125C224E375EE71" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n.; 1.29-1.61, mean = 1.45
<normalizedToken id="D2EDCF973001DF18A1FA93118E572D1C" originalValue="±">+/-</normalizedToken>
0.16 in
<taxonomicName id="A01C9CBA9F3AE466FD41BF3101B2CE75" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes inornatus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="inornatus">Imantodes inornatus</taxonomicName>
), and dark blotches on dorsum (dark spots and flecks in
<taxonomicName id="7321DE35B612B180AD6E8D7C3D13A9C2" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes inornatus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="inornatus">Imantodes inornatus</taxonomicName>
; Fig. 3). The new species can also be distinguished from
<taxonomicName id="DC478B09DE45D14335A8F9FC2B5E640E" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes cenchoa" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="cenchoa">Imantodes cenchoa</taxonomicName>
by having a single anal scale (vrs. two), fewer ventrals (t = 7.73, P &lt;0.001), fewer subcaudals (t = -4.04, P &lt;0.001), more infralabials (7-12, mean 9.92
<normalizedToken id="9921F2EF51E9423DB18276E1F8ECA0EE" originalValue="±">+/-</normalizedToken>
0.85 in
<taxonomicName id="2E9CB79ADFE7FE77511CD29B26BF4315" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes cenchoa" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="cenchoa">Imantodes cenchoa</taxonomicName>
), and dorsal dark blotches that include two or fewer vertebral scales and do not extend laterally onto ventrals (blotches are larger in
<taxonomicName id="86CC71EC36488F4ACBF9255DC342A66A" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes cenchoa" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="cenchoa">Imantodes cenchoa</taxonomicName>
and extend onto lateral tips of ventrals; Fig. 3). Scale counts and measurements of species of
<taxonomicName id="8186AFFC01A464EBFFB2D917C9F89E22" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes" pageId="2" pageNumber="93" phylum="Chordata" rank="genus">Imantodes</taxonomicName>
from Ecuador are presented in Table 2.
</paragraph>
<caption id="7793227309557A624CA0EA6DB639C54A" ID-Table-UUID="7793227309557A624CA0EA6DB639C54A" httpUri="http://table.plazi.org/id/7793227309557A624CA0EA6DB639C54A" pageId="2" pageNumber="93">
<paragraph id="9EDCB61B0104407E26B72CC68ED3BD4D" pageId="2" pageNumber="93">
Table 2.. Scale counts and measurements of species of
<taxonomicName id="29670A8CAEB258CCDFCA5A8FBF4506EC" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes" pageId="2" pageNumber="93" phylum="Chordata" rank="genus">Imantodes</taxonomicName>
from Ecuador. Range (first line) and mean
<normalizedToken id="9768F2DA291E81F20AA8B32A7C00A7F5" originalValue="±">+/-</normalizedToken>
SD (second line) are presented when appropriate. Sample size is presented in parentheses if different from that in heading.
</paragraph>
</caption>
<paragraph id="9F49D7F314E182B6296D453CE7C49672" pageId="2" pageNumber="93">
<table id="9A86D0EE1C084C2231228DAF3ED9AF53" pageId="2" pageNumber="93">
<tr id="3BE43F28B54A518CFE66D47A6B22FCB7" pageId="2" pageNumber="93">
<th id="186D0C0C36A5ABA68F28D1D80F50900F" colspan="1" pageId="2" pageNumber="93" rowspan="1">Character</th>
<th id="1028C9CAE64CBB4DFB9CEC55BC51D0D2" colspan="1" pageId="2" pageNumber="93" rowspan="1">
<taxonomicName id="B362538F578263FB70D22675D982DF40" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes cenchoa" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="cenchoa">Imantodes cenchoa</taxonomicName>
N = 42
</th>
<th id="1B6EB74774BB7124940D504D370CF3DE" colspan="1" pageId="2" pageNumber="93" rowspan="1">
<taxonomicName id="5A490FD264B93971E99801976B0DDC3C" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes inornatus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="inornatus">Imantodes inornatus</taxonomicName>
N = 6
</th>
<th id="036D960BD4AAA79AA8A348BE739ACBF6" colspan="1" pageId="2" pageNumber="93" rowspan="1">
<taxonomicName id="A116AC47BBF39D34568446E1D5DBF9FC" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes lentiferus" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="lentiferus">Imantodes lentiferus</taxonomicName>
N = 30
</th>
<th id="780AB7F039E7BF44997EEC9482627632" colspan="1" pageId="2" pageNumber="93" rowspan="1">
<taxonomicName id="74E683DB6739DE0A00B24B1A4C48BAD9" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="2" pageNumber="93" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. N = 7
</th>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection id="4A1F0DC8208674672F173D5D7232BDC7" lastPageId="5" lastPageNumber="96" pageId="2" pageNumber="93" type="description">
<paragraph id="3131805A45EFF4BC73778D56BBA74864" pageId="2" pageNumber="93">Description of holotype.</paragraph>
<paragraph id="A2EE8E5FFDEAB54B9935432497E2BB04" pageId="2" pageNumber="93">Male (Figs. 1, 2); SVL = 66.30 mm; tail length = 30.40 mm; head width = 7.98 mm; head length = 13.26 mm; head height = 5.37 mm.</paragraph>
<paragraph id="4BFEADB6052358A7873812C2ADDB6EBE" pageId="3" pageNumber="94">
<pageBreakToken id="031B7B9189391E8EA284B53C4F041FCA" pageId="3" pageNumber="94" start="start">Short</pageBreakToken>
, blunt head 1.7 times longer than broad and 2.5 times longer than deep; head abruptly distinct from neck, three times wider than thinnest part of neck and also slightly wider than greatest width of body; eye large and protuberant, occupying 27% of length of head, with elliptical pupil visible from anterior, lateral, dorsal, and ventral aspects; rostral 1.6 times wider than high, concave in anterior view, and narrowly visible from above; paired prefrontals extending anteroventrally to level of center of eye, each in contact with its mate and with frontal, supraocular, preocular, nasal, and internasal; frontal pentagonal, 1.6 times longer than wide (greatest width), and about 1.2 times longer than distance from its anterior edge to tip of snout; supraocular anteriorly narrow and posteriorly nearly as wide as greatest frontal width; broad parietals, about 1.3 times longer than wide; interparietal suture 1.2 times longer than length of frontal, and 1.4 times longer than distance from frontal to tip of snout; nasal plate single, centrally pierced by large naris (0.79 mm in diameter), in contact with rostral anteriorly, internasal dorsally, prefrontal posterodorsally, preocular posteriorly, and first and second supralabials ventrally; loreal absent; one large and high preocular; two postoculars (an extra tiny scale on left side ventrally), the lower somewhat less than half the size of the upper; temporals 2+2+3; supralabials 9, first and second in contact with nasal, fourth in contact with preocular, and fourth to seventh bordering the orbit; infralabials 13, with first six in contact with anterior genial, and sixth to eighth touching posterior genial; first pair of infralabials in contact medially behind mental; anterior and posterior genials nearly equal in length; gular scales with posterolateral apical pit.
</paragraph>
<paragraph id="0941FDD478B84F0C45422D7451B6E8B2" lastPageId="4" lastPageNumber="95" pageId="3" pageNumber="94">
Body higher than wide, rounded ventrolaterally; dorsal scales smooth, juxtaposed or subimbricate; dorsal body scales in 17 rows throughout; scales of vertebral row
<pageBreakToken id="F8D32E7FFD40FCF64F450576A85CAB57" pageId="4" pageNumber="95" start="start">conspicuously</pageBreakToken>
enlarged, 2.5 times wider than adjacent dorsals, with concave posterior margins; ventrals 242; anal plate single; subcaudals 161.
</paragraph>
<paragraph id="08B87E3014E59CBA14F4D1634A4604CA" pageId="4" pageNumber="95">Color in preservative of holotype (Figs 1, 2). Dorsal background light brown, with a longitudinal series of 63 dark brown middorsal blotches from head to cloaca; dark middorsal blotches longer anteriorly, 2-3 vertebral scales long, than posteriorly, 1-2 vertebral scales long, and extending laterally 1-3 (anteriorly) or more (posteriorly) dorsal scale rows, but never reaching ventral scales; each dark middorsal blotch irregularly bordered anteriorly and posteriorly by thin cream line; ventral aspect of body yellowish cream with dark brown spots and flecks; ventral aspect of tail yellowish cream with spots concentrating midventrally; dorsal surface of head light brown with several dark brown spots and two short dark stripes extending from posterior aspect of parietals to a point just posterior to head; ventral surface of head whitish cream.</paragraph>
<paragraph id="88643ABB7FC12C8231BB84F8433528F5" lastPageId="5" lastPageNumber="96" pageId="4" pageNumber="95">
Hemipenes (Fig. 4). The right hemipenis of the paratype DHMECN 6753 of
<taxonomicName id="B9DA06D276BBEAA902E16898FEA04323" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="4" pageNumber="95" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
was removed, fully everted and expanded (Fig. 4). The organ is bulbous and relatively long, 11.2 mm in length, and when adpressed to the outside of the tail it extends from the cloaca to the sixth subcaudal scale. The organ is longer than wide (width 46% of length), unilobed, symmetrical, calyculate, capitate, and arched towards the sulcal side. The sulcus spermaticus is simple, linear, semicentripetal, and thin, demarcated by thick bordering tissue at the base, particularly at the anterior border, and ending on the surface of the capitulum facing medially. The capitulum is ornamented with papillated calyces, spinulated proximally. The capitulum, approximately 45% the length of the hemipenis, is slightly demarcated by a groove, more prominent on the sulcal side and joining the sulcus spematicus. In the asulcate side the base of the capitulum has more prominent spines. Truncus covered by large spines, on the sulcate
<pageBreakToken id="6EA4CFC3E082D787CFD8BAFC23A33850" pageId="5" pageNumber="96" start="start">and</pageBreakToken>
asulcate and surfaces, 23 in total, 13 to the right of the sulcus spermaticus and 10 to the left, and has a few rows of small spines, two at the base on the asulcate side and two to three rows just right of the sulcus spermaticus.
</paragraph>
<caption id="0CBAEE4CF9B67F91E5014C6C96ACE850" pageId="5" pageNumber="96">
<paragraph id="E0605F7CA553CACA25E4A08B75D24AD1" pageId="5" pageNumber="96">
Figure 1. Holotype of
<taxonomicName id="DFF742B2DA40CE89DBD863C124EC0528" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. in dorsal (left) and ventral (right) views. Photographs by OTC.
</paragraph>
</caption>
<caption id="46C309A9E31CDE97C627412FBE4E55FA" pageId="5" pageNumber="96">
<paragraph id="ABA30797653E5A9BB09F663D7D4873AD" pageId="5" pageNumber="96">
Figure 2. Head of holotype of
<taxonomicName id="78C04C6863E4F4DE159740B8F74EBABE" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. in dorsal (top), lateral (middle) and ventral (bottom) views. Photographs by OTC.
</paragraph>
</caption>
<caption id="E92EF902B49C6F63906F448438D848DB" pageId="5" pageNumber="96">
<paragraph id="F17F79AA99DC0C4E257CC5EB1AFCE1DC" pageId="5" pageNumber="96">
Figure 3. Body segments of species of
<taxonomicName id="B84F536D78D22EA6A46D357066AD3B89" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes" pageId="5" pageNumber="96" phylum="Chordata" rank="genus">Imantodes</taxonomicName>
from Ecuador in dorsal (left) and lateral (right) views. A
<taxonomicName id="83C271AFDF68E86AAA214A459AC20D38" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. (DHMECN 6753, paratype) B
<taxonomicName id="40F020BC4F7572F006F2E84C699C803F" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes lentiferus" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="lentiferus">Imantodes lentiferus</taxonomicName>
(DHMECN 8345) C
<taxonomicName id="0A53227E842CCEA88E76CEB04392DE36" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes cenchoa" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="cenchoa">Imantodes cenchoa</taxonomicName>
(DHMECN 7826) D
<taxonomicName id="52168A3826B9072695B25C1083059A13" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes inornatus" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="inornatus">Imantodes inornatus</taxonomicName>
(DHMECN 5661). Photographs by MYM.
</paragraph>
</caption>
<caption id="956C4C602712D888A4C1F53F0F301113" pageId="5" pageNumber="96">
<paragraph id="BDCBAF872E00B32920EEDB988AA4A080" pageId="5" pageNumber="96">
Figure 4. Right hemipenis of
<taxonomicName id="95230B59DD575AB0F4457FE7627EED40" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. (DHMECN 6753, paratype) in sulcal (A), asulcal (B), and lateral (C) views D close-up of distal end showing spines interrupted by sulci. Photographs by MYM.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="F20982ACD3659D6C77969EE053517A43" pageId="5" pageNumber="96" type="variation">
<paragraph id="80D2A5A5A694E10523EBB43838C2EABC" pageId="5" pageNumber="96">Variation.</paragraph>
<paragraph id="4EA628F84E42818BD076913D3CD48D34" pageId="5" pageNumber="96">
Intraspecific variation in scale counts and measurements in
<taxonomicName id="7B821F2EADC83FC581EE241ABBF40C6D" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. is presented in Table 2. Color in life of paratypes UTA R-60205 and DHMECN 6753 (Fig. 5) is similar to color in preservative of holotype; iris copper brown. Middorsal blotches from head to cloaca vary between 55-66; one specimen (UTA R-60205) had one middorsal blotch covering five vertebral scales.
</paragraph>
<caption id="B837610D36BC3BEB1063BC9872A9631E" pageId="5" pageNumber="96">
<paragraph id="75531208D76039A028855B85CA066F7B" pageId="5" pageNumber="96">
Figure 5. Paratypes of
<taxonomicName id="E8036DD0AA4D2A506D5FD9A557CAC39A" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. UTA R-60205 (top), DHMECN 6753 (bottom). Photographs by ENS and MYM.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="49EA4BEC1C473AD5BF8AF33BE72F8FA8" lastPageId="7" lastPageNumber="98" pageId="5" pageNumber="96" type="distribution">
<paragraph id="F8E7066B7D4FE179EEC985239F7CC2ED" pageId="5" pageNumber="96">Distribution and ecology.</paragraph>
<paragraph id="5BD7B0AE31E7A9485B2888831D35C0AE" lastPageId="7" lastPageNumber="98" pageId="5" pageNumber="96">
<taxonomicName id="B142CB3FF985B7B74A819A10931745DB" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
inhabits Chocoan rainforests on the Pacific coast in northern Ecuador (Fig. 6). It occurs in lowland evergreen forest (
<bibRefCitation id="2D02E1BEFF3D499FF78219E247B45C95" pageId="5" pageNumber="96">
<normalizedToken id="A6B6E30A76EFAC71EA115D91DAEAC2AF" originalValue="Cerón">Ceron</normalizedToken>
et al. 1999
</bibRefCitation>
) at elevations of 115-260 m in the provinces of Carchi and Esmeraldas. This new species has been collected in sympatry with
<taxonomicName id="09DEBFE5A9A3A48B9F1930E404D07A86" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes cenchoa" pageId="5" pageNumber="96" phylum="Chordata" rank="species" species="cenchoa">Imantodes cenchoa</taxonomicName>
in Esmeraldas,
<pageBreakToken id="74142A9B5ED9395C116048D44D6D270F" pageId="6" pageNumber="97" start="start">and</pageBreakToken>
most likely also shares its distribution with
<taxonomicName id="71DC4A21B1DBF53E8AF8C3D06F86C648" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes inornatus" pageId="6" pageNumber="97" phylum="Chordata" rank="species" species="inornatus">Imantodes inornatus</taxonomicName>
. Other colubrid snakes collected in Tobar Donoso (Carchi) are
<taxonomicName id="2EDE324FADF2FDF5DBD60089C68B65D2" class="Squamata" family="Colubridae" genus="Chironius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chironius grandisquamis" pageId="6" pageNumber="97" phylum="Chordata" rank="species" species="grandisquamis">Chironius grandisquamis</taxonomicName>
,
<taxonomicName id="4F9B35E43567567A2089F7F2C385A888" class="Squamata" family="Colubridae" genus="Clelia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Clelia clelia" pageId="6" pageNumber="97" phylum="Chordata" rank="species" species="clelia">Clelia clelia</taxonomicName>
,
<taxonomicName id="31C9F1EEA86628F092EE019C8FF32DF1" class="Squamata" family="Colubridae" genus="Dendrophidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dendrophidion clarkii" pageId="6" pageNumber="97" phylum="Chordata" rank="species" species="clarkii">Dendrophidion clarkii</taxonomicName>
,
<taxonomicName id="3EA53E404911B2E637628B926059129A" class="Squamata" family="Colubridae" genus="Leptophis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptophis ahaetulla" pageId="6" pageNumber="97" phylum="Chordata" rank="species" species="ahaetulla">Leptophis ahaetulla</taxonomicName>
,
<taxonomicName id="B43A700611CC10FA6B57D2D1CB876637" class="Squamata" family="Colubridae" genus="Mastigodryas" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mastigodryas" pageId="6" pageNumber="97" phylum="Chordata" rank="genus">Mastigodryas</taxonomicName>
sp.,
<taxonomicName id="8A989B63F2BD66BE9A70E95ABB8FB498" class="Squamata" family="Colubridae" genus="Ninia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ninia atrata" pageId="6" pageNumber="97" phylum="Chordata" rank="species" species="atrata">Ninia atrata</taxonomicName>
,
<taxonomicName id="5A02C125AB7B20059F0696CACD75FE7F" class="Squamata" family="Colubridae" genus="Oxyrhopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Oxyrhopus petola" pageId="6" pageNumber="97" phylum="Chordata" rank="species" species="petola">Oxyrhopus petola</taxonomicName>
,
<taxonomicName id="670F1592B5F648709321E0933488BF38" genus="Pseustes" lsidName="Pseustes shropshirei" pageId="7" pageNumber="98" rank="species" species="shropshirei">
<pageBreakToken id="39C1CD47F793366016E946C2767FDC12" pageId="7" pageNumber="98" start="start">Pseustes</pageBreakToken>
shropshirei
</taxonomicName>
,
<taxonomicName id="648C1D252D750631518B025C953B9AA9" class="Squamata" family="Colubridae" genus="Sibon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sibon nebulatus" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="nebulatus">Sibon nebulatus</taxonomicName>
,
<taxonomicName id="60555A5A4907A2D0318CD24C7B46FAFE" class="Squamata" family="Colubridae" genus="Synophis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Synophis bicolor" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="bicolor">Synophis bicolor</taxonomicName>
,
<taxonomicName id="A263677C31AB589EB19A23E7C004126D" class="Squamata" family="Colubridae" genus="Tantilla" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tantilla melanocephala" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="melanocephala">Tantilla melanocephala</taxonomicName>
, and
<taxonomicName id="5145DBB3A0AA2FAD26C5A4061894446F" class="Squamata" family="Colubridae" genus="Xenodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Xenodon rabdocephalus" order="Tetraodontiformes" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="rabdocephalus">Xenodon rabdocephalus</taxonomicName>
. The known localities of
<taxonomicName id="3DA690FE0C4E8A4EDD3885B00A6C7633" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
lie in close proximity to the Ecuador-Colombia border and we expect for it to be found in neighboring Colombia.
</paragraph>
<caption id="04258687D737C4F7885C0B5AB4A49A0C" pageId="7" pageNumber="98">
<paragraph id="7D0A440F38DFD4744B771FB9C4F42FB0" pageId="7" pageNumber="98">
Figure 6. Distribution of
<taxonomicName id="DF062B6994581F765FB70B2FED6D964F" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. (circles) and its sister species
<taxonomicName id="39D4650D323710729CBA92C26FDE4D7A" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes lentiferus" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="lentiferus">Imantodes lentiferus</taxonomicName>
(squares) in Ecuador.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="2DDD54C092B71CD068820DAE93EF81E0" pageId="7" pageNumber="98" type="etymology">
<paragraph id="05C4C349CD103E77D3E23E871CF682E1" pageId="7" pageNumber="98">Etymology.</paragraph>
<paragraph id="96EF2A44B1001CA4E85CFCF31478FDEF" pageId="7" pageNumber="98">
The specific epithet chocoensis is an adjective derived from
<normalizedToken id="3DC97EE1A5F4E33978580327F312AB67" originalValue="Chocó">Choco</normalizedToken>
, the very humid tropical region comprising the Pacific coast of northern Ecuador, Colombia and Panama (
<bibRefCitation id="8DFDCF009F8E611A6C75FCF47E611CCA" author="Morrone, JJ" journalOrPublisher="Caldasia" pageId="9" pageNumber="100" pagination="43 - 6" title="Toward a cladistic model for the Caribbean subregion: delimitation of areas of endemism." volume="23" year="2001">Morrone 2001</bibRefCitation>
). This region is part of the 274,597 km2
<normalizedToken id="5A57B081D6BEBE53EF6579B0FFA1552E" originalValue="Tumbes-Chocó-Magdalena">Tumbes-Choco-Magdalena</normalizedToken>
hotspot as defined by Conservation International, which includes more than 320 species of reptiles.
</paragraph>
</subSubSection>
<subSubSection id="0C3799774FF3F6A4A37E40CACBF3E38C" lastPageId="8" lastPageNumber="99" pageId="7" pageNumber="98" type="phylogenetic relationships">
<paragraph id="BAEFD457F5CA2DEECDCB026522616B89" pageId="7" pageNumber="98">Phylogenetic relationships.</paragraph>
<paragraph id="B9D6431F7D6605AF6E67216CCA2CD804" lastPageId="8" lastPageNumber="99" pageId="7" pageNumber="98">
Selected models of evolution for sampled fragments of ND4 and cyt-b genes were HKY+I+G and TPM2uf+I+G, respectively. The resulting 50% majority rule consensus tree (Fig. 7) supports strongly (PP=1) a sister taxon relationship between
<taxonomicName id="0D433B52D35F13249B3462DC5C2121B9" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. and
<taxonomicName id="7176ADBE6C56C7EBDC840F3481201639" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes lentiferus" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="lentiferus">Imantodes lentiferus</taxonomicName>
, as well as the exclusivity (
<bibRefCitation id="B7C34CD181EC12206124BC3C5981281A" author="Rieppel, O" journalOrPublisher="Journal of Zoological Systematics and Evolutionary Research" pageId="9" pageNumber="100" pagination="1 - 8" title="Species monophyly." url="10.1111/j.14390469.2009.00545.x" volume="48" year="2010">Rieppel 2010</bibRefCitation>
) of both species. Similarly,
<taxonomicName id="12BC3D0FB9EB93E08BDC4F5CE144ECFD" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes inornatus" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="inornatus">Imantodes inornatus</taxonomicName>
and
<taxonomicName id="F645FC8A2CA7EF131F608585E5DE6666" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes cenchoa" pageId="7" pageNumber="98" phylum="Chordata" rank="species" species="cenchoa">Imantodes cenchoa</taxonomicName>
are recovered as
<pageBreakToken id="70A96E7FFF89E2908DBA8DB662CF726A" pageId="8" pageNumber="99" start="start">exclusive</pageBreakToken>
clades with strong support (PP=1). Noteworthy the
<taxonomicName id="219437FF08E035963023830B59DB0130" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes cenchoa" pageId="8" pageNumber="99" phylum="Chordata" rank="species" species="cenchoa">Imantodes cenchoa</taxonomicName>
clade includes samples from Guatemala, Costa Rica, Panama, Brasil, and Colombia, Ecuador and Peru on both sides of the Andes. In agreement with previous hypotheses,
<taxonomicName id="340DDAA831D3AE936C54D3687C05EC41" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes gemmistratus" pageId="8" pageNumber="99" phylum="Chordata" rank="species" species="gemmistratus">Imantodes gemmistratus</taxonomicName>
is paraphyletic; of three samples included in this study, one from Guatemala is sister to the
<taxonomicName id="005C3A7C8B9192460C4ADFB2382F8CB9" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes cenchoa" pageId="8" pageNumber="99" phylum="Chordata" rank="species" species="cenchoa">Imantodes cenchoa</taxonomicName>
clade with strong support (PP=1), whereas the other two, from Mexico, are weakly supported as sister to the
<taxonomicName id="3AEA7BC74CB7682F4D7A62A104806D3C" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes chocoensis" pageId="8" pageNumber="99" phylum="Chordata" rank="species" species="chocoensis">Imantodes chocoensis</taxonomicName>
sp. n. and
<taxonomicName id="D51166122BE2015A6C806B26F76BB4CE" class="Squamata" family="Colubridae" genus="Imantodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Imantodes lentiferus" pageId="8" pageNumber="99" phylum="Chordata" rank="species" species="lentiferus">Imantodes lentiferus</taxonomicName>
clade.
</paragraph>
<caption id="F4E1B93BCC7CFE74B23BF75B057A2771" pageId="8" pageNumber="99">
<paragraph id="A8DC218C32B9524047647D1878C4D9A2" pageId="8" pageNumber="99">Figure 7. Majority rule (50%) consensus tree of 18,000 trees obtained from a Bayesian analysis of two mitochondrial genes (cyt-b, ND4) and 29 specimens. Asterisks correspond to posterior probability values&gt; 0.99. Voucher numbers followed by country of collection are indicated for each terminal. E: east of the Andes, W: west of the Andes.</paragraph>
</caption>
</subSubSection>
</treatment>
</document>