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<document ID-DOI="http://dx.doi.org/10.3897/vz.72.e82201" ID-Pensoft-Pub="2625-8498-72-371" ID-Pensoft-UUID="0F0F6B209F615C299100DE481CEB5C04" ID-ZooBank="E1F61203222C418AA8AC100CE72604A5" ModsDocID="2625-8498-72-371" checkinTime="1655292402210" checkinUser="pensoft" docAuthor="Nguyen, Anh The, Duong, Tang Van, Wood Jr., Perry L. &amp; Grismer, L. Lee" docDate="2022" docId="BAFEBE2DBE005E6896DED88121A55C92" docLanguage="en" docName="VertZool 72: 371-384" docOrigin="Vertebrate Zoology 72" docPubDate="2022-06-14" docSource="http://dx.doi.org/10.3897/vz.72.e82201" docTitle="Lycodon anakradaya Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer 2022, sp. nov." docType="treatment" docUuid="FF1EA2E5-923B-4FA6-BF92-BECBEF7B824E" docUuidSource="ZooBank" docVersion="2" id="0F0F6B209F615C299100DE481CEB5C04" lastPageNumber="371" masterDocId="0F0F6B209F615C299100DE481CEB5C04" masterDocTitle="Two new syntopic species of wolf snakes (genus Lycodon H. Boie in Fitzinger, 1826) from an imperiled ecosystem in the Song Giang River Valley of southern Vietnam (Squamata: Colubridae)" masterLastPageNumber="384" masterPageNumber="371" pageNumber="371" updateTime="1655292716029" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Two new syntopic species of wolf snakes (genus Lycodon H. Boie in Fitzinger, 1826) from an imperiled ecosystem in the Song Giang River Valley of southern Vietnam (Squamata: Colubridae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Nguyen, Anh The</mods:namePart>
<mods:affiliation>Vietnambirds Foundation, No 94, Street 64, District 6, Ho Chi Minh city, Vietnam</mods:affiliation>
<mods:nameIdentifier type="email">vietnamwildlife2012@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Duong, Tang Van</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0001-9943-8843</mods:nameIdentifier>
<mods:affiliation>Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, Hanoi, Vietnam &amp; Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Hanoi, Vietnam</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wood Jr., Perry L.</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0003-3767-5274</mods:nameIdentifier>
<mods:affiliation>Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, Michigan, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Grismer, L. Lee</mods:namePart>
<mods:affiliation>Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92505, USA &amp; Department of Herpetology, San Diego Natural History Museum, PO Box 121390, San Diego, California, 92112, USA</mods:affiliation>
<mods:nameIdentifier type="email">lgrismer@lasierra.edu</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>Vertebrate Zoology</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2022</mods:date>
<mods:detail type="pubDate">
<mods:number>2022-06-14</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>72</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>371</mods:start>
<mods:end>384</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location>
<mods:url>http://dx.doi.org/10.3897/vz.72.e82201</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/vz.72.e82201</mods:identifier>
<mods:identifier type="Pensoft-Pub">2625-8498-72-371</mods:identifier>
<mods:identifier type="ZooBank">E1F61203222C418AA8AC100CE72604A5</mods:identifier>
<mods:identifier type="Pensoft-UUID">0F0F6B209F615C299100DE481CEB5C04</mods:identifier>
</mods:mods>
<subSection pageId="0" pageNumber="371" type="results">
<treatment LSID="urn:lsid:zoobank.org:act:FF1EA2E5-923B-4FA6-BF92-BECBEF7B824E" httpUri="http://treatment.plazi.org/id/BAFEBE2DBE005E6896DED88121A55C92" lastPageNumber="371" pageId="0" pageNumber="371">
<subSubSection pageId="0" pageNumber="371" type="nomenclature">
<paragraph pageId="0" pageNumber="371">
<taxonomicName LSID="http://zoobank.org/FF1EA2E5-923B-4FA6-BF92-BECBEF7B824E" authority="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer, 2022" authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya" status="sp. nov.">Lycodon anakradaya</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="371">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="description">
<paragraph pageId="0" pageNumber="371">
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. A. Holotype of Lycodon anakardaya sp. nov. SIEZC 20247. B Holotype of Lycodon truongi sp. nov. SIEZC 20248. C Paratype of Lycodon ankardaya sp. nov. SIEZC 20248. Photographs by Anh The Nguyen." figureDoi="10.3897/vz.72.e82201.figure4" httpUri="https://binary.pensoft.net/fig/696042" pageId="0" pageNumber="371">Figures 4</figureCitation>
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Holotype of Lycodon anakradaya sp. nov. SIEZC 20247 from the Song Giang River Valley Khanh Trung Commune, Khanh Vinh District, Khanh Hoa Province, Vietnam. A Dorsal view of head. B Gular region. C Lateral view of head. D Lateral view of a portion of the body. E Dorsal view of body. F Ventral view of body. Photographs by Anh The Nguyen." figureDoi="10.3897/vz.72.e82201.figure5" httpUri="https://binary.pensoft.net/fig/696043" pageId="0" pageNumber="371">, 5</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="suggested common name">
<paragraph pageId="0" pageNumber="371">Suggested common name.</paragraph>
<paragraph pageId="0" pageNumber="371">
Rhade wolf snake - Rắn khuyết
<normalizedToken originalValue="Ê">E</normalizedToken>
<normalizedToken originalValue="đê">đe</normalizedToken>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="holotype">
<paragraph pageId="0" pageNumber="371">Holotype.</paragraph>
<paragraph pageId="0" pageNumber="371">
Adult male (SIEZC 20247) collected on 21 December 2020 by Anh The Nguyen from Song Giang River Valley (
<geoCoordinate degrees="12.37079" direction="north" orientation="latitude" precision="1" value="12.37079">12.37079°N</geoCoordinate>
,
<geoCoordinate degrees="108.83643" direction="east" orientation="longitude" precision="1" value="108.83643">108.83643°E</geoCoordinate>
; at elevation 493 m a.s.l.), Khanh Trung Commune, Khanh Vinh District, Khanh Hoa Province, Vietnam.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="paratype">
<paragraph pageId="0" pageNumber="371">Paratype.</paragraph>
<paragraph pageId="0" pageNumber="371">
Juvenile (SIEZC 20248) collected on 21 August 2020 by Anh The Nguyen from Song Giang River Valley (
<geoCoordinate degrees="12.37079" direction="north" orientation="latitude" precision="1" value="12.37079">12.37079°N</geoCoordinate>
,
<geoCoordinate degrees="108.83643" direction="east" orientation="longitude" precision="1" value="108.83643">108.83643°E</geoCoordinate>
; at elevation 580 m a.s.l.), Khanh Trung Commune, Khanh Vinh District, Khanh Hoa Province, Vietnam.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="diagnosis">
<paragraph pageId="0" pageNumber="371">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="371">
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
is separated from all other species in the
<taxonomicName genus="L." lsidName="L. rufozonatus" pageId="0" pageNumber="371" rank="species" species="rufozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. rufozonatus</emphasis>
</taxonomicName>
complex by having the combination of a maximum SVL length of 790 mm; TaL 190 mm; 17 or 18-17-15 dorsal scale rows; 12 maxillary teeth; eight supralabials with the 3rd-5th contacting the eye; nine or 10 infralabials; one preocular; two postoculars; an elongate loreal not contacting the eye; two anterior temporals; three posterior temporals; 223-232 ventral scales; 87 paired subcaudal scales; a divided precloacal plate; five keeled vertebral scale rows from midbody to vent; uniform dark-brown to black adult head pattern; no wide light-colored nuchal-occipital collar; dark-brown/black dorsal ground color; 12 wide, orangish body bands; 5-7 orangish caudal bands; orangish dorsal bands on ventrolateral section of body as opposed to a reticulated pattern; and anterior one-half of venter orangish grading into a dark-brown posteriorly. These characters are scored across all species of the
<taxonomicName genus="L." lsidName="L. rufozonatus" pageId="0" pageNumber="371" rank="species" species="rufozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. rufozonatus</emphasis>
</taxonomicName>
complex in Table
<tableCitation captionStart="Table 2" captionStartId="T2" captionText="Table 2. Diagnostic characters of Lycodon truongi sp. nov. and L. anakradaya sp. nov. that differentiate them from each other and all other species of the L. rufizonatus complex. Green cells denote characters that separate L. truongi sp. nov. and L. anakradaya sp. nov. from the other species. Orange cells denote characters separating L. anakradaya sp. nov. from all other species. Blue cells denote characters separating L. truongi sp. nov. from all other species. Data for previously described species come from Jansen et al (2019), Luu et al (2018, 2019), Wang et al (2020, 2021), and Song (2021)." httpUri="http://table.plazi.org/id/66AA650EF548A8AD967E5BC514FBCE0F" pageId="0" pageNumber="371" tableUuid="66AA650EF548A8AD967E5BC514FBCE0F">2</tableCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="description">
<paragraph pageId="0" pageNumber="371">Description of holotype</paragraph>
<paragraph pageId="0" pageNumber="371">
<emphasis bold="true" pageId="0" pageNumber="371">
(Figs
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. A. Holotype of Lycodon anakardaya sp. nov. SIEZC 20247. B Holotype of Lycodon truongi sp. nov. SIEZC 20248. C Paratype of Lycodon ankardaya sp. nov. SIEZC 20248. Photographs by Anh The Nguyen." figureDoi="10.3897/vz.72.e82201.figure4" httpUri="https://binary.pensoft.net/fig/696042" pageId="0" pageNumber="371">4</figureCitation>
,
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Holotype of Lycodon anakradaya sp. nov. SIEZC 20247 from the Song Giang River Valley Khanh Trung Commune, Khanh Vinh District, Khanh Hoa Province, Vietnam. A Dorsal view of head. B Gular region. C Lateral view of head. D Lateral view of a portion of the body. E Dorsal view of body. F Ventral view of body. Photographs by Anh The Nguyen." figureDoi="10.3897/vz.72.e82201.figure5" httpUri="https://binary.pensoft.net/fig/696043" pageId="0" pageNumber="371">5</figureCitation>
; Table
<tableCitation captionStart="Table 3" captionStartId="T3" captionText="Table 3. Meristic, morphometric, and discrete color pattern and scale morphology characters of the type specimens of Lycodon truongi sp. nov. and L. anakradaya sp. nov. All measurements are in millimeters." httpUri="http://table.plazi.org/id/E330F51DC6F2A4C981FECB4D7BB131BB" pageId="0" pageNumber="371" tableUuid="E330F51DC6F2A4C981FECB4D7BB131BB">3</tableCitation>
).
</emphasis>
Head flattened slightly sloping anteriorly, distinct from the neck, HL 27.8 mm, HW 18.0 mm, HH 11.8 mm, snout somewhat elongate, SnL 8.2 mm, EN 5.2 mm; nostril oval, large, in the middle of the nasal, IND 6.0 mm; eye moderately sized, ED 4.0 mm, with a vertically elliptic pupil; rostral triangular, hardly visible from above; nasal vertically divided by a furrow along posterior margin of nostril; two square internasals in wide, medial contact, and in contact with two large, square prefrontals posteriorly; followed by a single, azygous, subpentagonal frontal, nearly as wide as long; two large, elongate parietals, contacted laterally by upper anterior and posterior temporals and a larger paraparietal; 1/1 wide, elongate supraocular; 1/1 small preocular, contacting third supralabial; 2/2 postoculars, upper slightly larger than lower; 1/1 elongate loreal not contacting eye, in contact with second and third supralabials ventrally, the prefrontal dorsally, preocular posteriorly, the nasal anteriorly; 8/8 supralabials, second to sixth higher than wide; first and second supralabials in contact with nasal; third, fourth, and fifth, supralabials contacting eye; sixth and seventh supralabials largest; two elongate anterior temporals, lower temporal largest; three square posterior temporals of similar size; 10/10 infralabials; first pair of infralabials contact medially forming a deep, medial groove; first five infralabials in contact with first pair of chinshields; similarly sized anterior and posterior pair of elongate chinshields, bearing deep, medial grooves contiguous with groove separating first pair of infralabials; 12 maxillary teeth.
</paragraph>
<paragraph pageId="0" pageNumber="371">Body elongate, somewhat laterally compressed; SVL 790 mm; TaL 190 mm; ToL 980 mm. 225 ventrals, 87 paired, subcaudals; cloacal shield divided; dorsal scales in 18-17-15 rows with three keeled vertebral rows; vertebral scale row not enlarged; no apical pits.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="coloration in life">
<paragraph pageId="0" pageNumber="371">Coloration in life</paragraph>
<paragraph pageId="0" pageNumber="371">
<emphasis bold="true" pageId="0" pageNumber="371">
(Figs
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. A. Holotype of Lycodon anakardaya sp. nov. SIEZC 20247. B Holotype of Lycodon truongi sp. nov. SIEZC 20248. C Paratype of Lycodon ankardaya sp. nov. SIEZC 20248. Photographs by Anh The Nguyen." figureDoi="10.3897/vz.72.e82201.figure4" httpUri="https://binary.pensoft.net/fig/696042" pageId="0" pageNumber="371">4</figureCitation>
,
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Holotype of Lycodon anakradaya sp. nov. SIEZC 20247 from the Song Giang River Valley Khanh Trung Commune, Khanh Vinh District, Khanh Hoa Province, Vietnam. A Dorsal view of head. B Gular region. C Lateral view of head. D Lateral view of a portion of the body. E Dorsal view of body. F Ventral view of body. Photographs by Anh The Nguyen." figureDoi="10.3897/vz.72.e82201.figure5" httpUri="https://binary.pensoft.net/fig/696043" pageId="0" pageNumber="371">5</figureCitation>
).
</emphasis>
Dorsal ground color of head, body and tail dark-brown to nearly uniformly black; body bearing 13 wide orangish body bands extending to ventral scales, seven wide orangish caudal bands incompletely encircling tail; anterior half of venter orangish with dark-brown mottling grading into a uniform dark-brown venter and subcaudal region.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="variation">
<paragraph pageId="0" pageNumber="371">Variation</paragraph>
<paragraph pageId="0" pageNumber="371">
<emphasis bold="true" pageId="0" pageNumber="371">
(Figs
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. A. Holotype of Lycodon anakardaya sp. nov. SIEZC 20247. B Holotype of Lycodon truongi sp. nov. SIEZC 20248. C Paratype of Lycodon ankardaya sp. nov. SIEZC 20248. Photographs by Anh The Nguyen." figureDoi="10.3897/vz.72.e82201.figure4" httpUri="https://binary.pensoft.net/fig/696042" pageId="0" pageNumber="371">4</figureCitation>
-
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Paratype of Lycodon anakradaya sp. nov. SIEZC 20248 from the Song Giang River Valley Khanh Trung Commune, Khanh Vinh District, Khanh Hoa Province, Vietnam. A Dorsal view of head. B Gular region. C Lateral view of head. D Dorsal view of body. E Lateral view of body. Photographs by Anh The Nguyen." figureDoi="10.3897/vz.72.e82201.figure6" httpUri="https://binary.pensoft.net/fig/696044" pageId="0" pageNumber="371">6</figureCitation>
; Table
<tableCitation captionStart="Table 3" captionStartId="T3" captionText="Table 3. Meristic, morphometric, and discrete color pattern and scale morphology characters of the type specimens of Lycodon truongi sp. nov. and L. anakradaya sp. nov. All measurements are in millimeters." httpUri="http://table.plazi.org/id/E330F51DC6F2A4C981FECB4D7BB131BB" pageId="0" pageNumber="371" tableUuid="E330F51DC6F2A4C981FECB4D7BB131BB">3</tableCitation>
).
</emphasis>
The paratype SIEZC 20248 resembles the holotype in overall color patter except the dorsal and caudal bands are white and the head plates are edged in white. Presumably the 12 dorsal and five caudal bands become orangish and the head will become unicolor dark-brown/black with increasing SVL. The paratype lacks keeled vertebral scales which also may become keeled with increasing SVL. Meristic differences are listed in Table
<tableCitation captionStart="Table 3" captionStartId="T3" captionText="Table 3. Meristic, morphometric, and discrete color pattern and scale morphology characters of the type specimens of Lycodon truongi sp. nov. and L. anakradaya sp. nov. All measurements are in millimeters." httpUri="http://table.plazi.org/id/E330F51DC6F2A4C981FECB4D7BB131BB" pageId="0" pageNumber="371" tableUuid="E330F51DC6F2A4C981FECB4D7BB131BB">3</tableCitation>
.
</paragraph>
<caption doi="10.3897/vz.72.e82201.figure5" httpUri="https://binary.pensoft.net/fig/696043" pageId="0" pageNumber="371" start="Figure 5" startId="F5">
<paragraph pageId="0" pageNumber="371">
<emphasis bold="true" pageId="0" pageNumber="371">Figure 5.</emphasis>
Holotype of
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
SIEZC 20247 from the Song Giang River Valley Khanh Trung Commune, Khanh Vinh District, Khanh Hoa Province, Vietnam.
<emphasis bold="true" pageId="0" pageNumber="371">A</emphasis>
Dorsal view of head.
<emphasis bold="true" pageId="0" pageNumber="371">B</emphasis>
Gular region.
<emphasis bold="true" pageId="0" pageNumber="371">C</emphasis>
Lateral view of head.
<emphasis bold="true" pageId="0" pageNumber="371">D</emphasis>
Lateral view of a portion of the body.
<emphasis bold="true" pageId="0" pageNumber="371">E</emphasis>
Dorsal view of body.
<emphasis bold="true" pageId="0" pageNumber="371">F</emphasis>
Ventral view of body. Photographs by Anh The Nguyen.
</paragraph>
</caption>
<caption doi="10.3897/vz.72.e82201.figure6" httpUri="https://binary.pensoft.net/fig/696044" pageId="0" pageNumber="371" start="Figure 6" startId="F6">
<paragraph pageId="0" pageNumber="371">
<emphasis bold="true" pageId="0" pageNumber="371">Figure 6.</emphasis>
Paratype of
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
SIEZC 20248 from the Song Giang River Valley Khanh Trung Commune, Khanh Vinh District, Khanh Hoa Province, Vietnam.
<emphasis bold="true" pageId="0" pageNumber="371">A</emphasis>
Dorsal view of head.
<emphasis bold="true" pageId="0" pageNumber="371">B</emphasis>
Gular region.
<emphasis bold="true" pageId="0" pageNumber="371">C</emphasis>
Lateral view of head.
<emphasis bold="true" pageId="0" pageNumber="371">D</emphasis>
Dorsal view of body.
<emphasis bold="true" pageId="0" pageNumber="371">E</emphasis>
Lateral view of body. Photographs by Anh The Nguyen.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="etymology">
<paragraph pageId="0" pageNumber="371">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="371">
The specific epithet &quot;
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon anakradaya</emphasis>
</taxonomicName>
&quot; is given in a reference to the Ede people (&quot;
<emphasis italics="true" pageId="0" pageNumber="371">Anak Radaya</emphasis>
&quot; in Ede language), an Austronesian ethnic group living in the upland forested areas of southern Vietnam including those that surround the Song Giang River valley.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="natural history">
<paragraph pageId="0" pageNumber="371">Natural history</paragraph>
<paragraph pageId="0" pageNumber="371">
<emphasis bold="true" pageId="0" pageNumber="371">
(Fig.
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Riparian habitat of a tributary of the Song Giang River where Lycodon truongi sp. nov. and L. anakardaya sp. nov. were collected. Photographs by Anh The Nguyen." figureDoi="10.3897/vz.72.e82201.figure7" httpUri="https://binary.pensoft.net/fig/696045" pageId="0" pageNumber="371">7</figureCitation>
).
</emphasis>
The holotype was observed between 18:30-22:30 h swimming across a small, clear stream with a sandy bottom in an effort to escape detection. Once crossing the stream, it climbed up a steep river bank. The stream is at 600 m elevation and bordered by moss-covered granite boulders less than 4 m wide beneath a canopy of broad-leaved evergreen forest. The stream supports many small fishes (
<taxonomicName baseAuthorityName="Valenciennes" baseAuthorityYear="1842" class="Actinopterygii" family="Cyprinidae" genus="Poropuntius" kingdom="Animalia" lsidName="Poropuntius deauratus" order="Cypriniformes" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="deauratus">Poropuntius cf. deauratus</taxonomicName>
,
<taxonomicName baseAuthorityName="Hamilton" baseAuthorityYear="1822" class="Actinopterygii" family="Channidae" genus="Channa" kingdom="Animalia" lsidName="Channa gachua" order="Perciformes" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="gachua">Channa cf. gachua</taxonomicName>
,
<taxonomicName class="Actinopterygii" family="Nemacheilidae" genus="Schistura" kingdom="Animalia" lsidName="Schistura" order="Cypriniformes" pageId="0" pageNumber="371" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="371">Schistura</emphasis>
</taxonomicName>
sp.) and various species of amphibians (
<taxonomicName authorityName="Tschudi" authorityYear="1838" class="Amphibia" family="Ranidae" genus="Hylarana" kingdom="Animalia" lsidName="Hylarana" order="Anura" pageId="0" pageNumber="371" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="371">Hylarana</emphasis>
</taxonomicName>
sp.,
<taxonomicName authorityName="Fei, Ye &amp; Huang" authorityYear="1990" class="Amphibia" family="Ranidae" genus="Odorrana" kingdom="Animalia" lsidName="Odorrana" order="Anura" pageId="0" pageNumber="371" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="371">Odorrana</emphasis>
</taxonomicName>
sp.). The juvenile paratype of
<taxonomicName genus="L." lsidName="L. anakradaya" pageId="0" pageNumber="371" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
was found on a branch 30 cm above the surface of the water approximately 1 km upstream from the type locality.
</paragraph>
<caption doi="10.3897/vz.72.e82201.figure7" httpUri="https://binary.pensoft.net/fig/696045" pageId="0" pageNumber="371" start="Figure 7" startId="F7">
<paragraph pageId="0" pageNumber="371">
<emphasis bold="true" pageId="0" pageNumber="371">Figure 7.</emphasis>
Riparian habitat of a tributary of the Song Giang River where
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon truongi" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="truongi">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon truongi</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
and
<taxonomicName genus="L." lsidName="L. anakardaya" pageId="0" pageNumber="371" rank="species" species="anakardaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. anakardaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
were collected. Photographs by Anh The Nguyen.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="371" type="comparisons">
<paragraph pageId="0" pageNumber="371">Comparisons</paragraph>
<paragraph pageId="0" pageNumber="371">
<emphasis bold="true" pageId="0" pageNumber="371">
(Table
<tableCitation captionStart="Table 2" captionStartId="T2" captionText="Table 2. Diagnostic characters of Lycodon truongi sp. nov. and L. anakradaya sp. nov. that differentiate them from each other and all other species of the L. rufizonatus complex. Green cells denote characters that separate L. truongi sp. nov. and L. anakradaya sp. nov. from the other species. Orange cells denote characters separating L. anakradaya sp. nov. from all other species. Blue cells denote characters separating L. truongi sp. nov. from all other species. Data for previously described species come from Jansen et al (2019), Luu et al (2018, 2019), Wang et al (2020, 2021), and Song (2021)." httpUri="http://table.plazi.org/id/66AA650EF548A8AD967E5BC514FBCE0F" pageId="0" pageNumber="371" tableUuid="66AA650EF548A8AD967E5BC514FBCE0F">2</tableCitation>
).
</emphasis>
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
could not be separated statistically from other species of the
<taxonomicName genus="L." lsidName="L. rufozonatus" pageId="0" pageNumber="371" rank="species" species="rufozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. rufozonatus</emphasis>
</taxonomicName>
complex because of its small sample size (n=2). Therefore, the comparisons below are based on discrete differences some of which are color pattern characters tradionally used to separate species (Jansen et al. (2019),
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4586.2.3" author="Luu, VQ" journalOrPublisher="Tropical Zoology" pageId="0" pageNumber="371" refId="B17" refString="Luu, VQ, Bonkowski, M, Nguyen, TQ, Le, MD, Calame, T, Ziegler, T, 2018. A new species of Lycodon Boie, 1826 (Serpentes: Colubridae) from central Laos. Revue Suisse de Zoologie 125: 263-276. https://doi.org/10.11646/zootaxa.4586.2.3" title="A new species of Lycodon Boie, 1826 (Serpentes: Colubridae) from central Laos. Revue Suisse de Zoologie 125: 263 - 276." url="https://doi.org/10.11646/zootaxa.4586.2.3" year="2018">Luu et al. (2018</bibRefCitation>
,
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4586.2.3" author="Luu, VQ" journalOrPublisher="Tropical Zoology" pageId="0" pageNumber="371" refId="B18" refString="Luu, VQ, Ziegler, T, Van, HN, Le, MD, Hoang, TT, 2019. A new species of Lycodon Boie, 1826 (Serpentes: Colubridae) from Thanh Hoa Province, Vietnam. Zootaxa 4586: 261-277. https://doi.org/10.11646/zootaxa.4586.2.3" title="A new species of Lycodon Boie, 1826 (Serpentes: Colubridae) from Thanh Hoa Province, Vietnam. Zootaxa 4586: 261 - 277." url="https://doi.org/10.11646/zootaxa.4586.2.3" year="2019">2019</bibRefCitation>
),
<bibRefCitation DOI="https://doi.org/10.3897/zookeys.954.53432" author="Wang, J" journalOrPublisher="Russian Journal of Herpetology" pageId="0" pageNumber="371" refId="B31" refString="Wang, J, Qi, S, Lyu, ZT, Zeng, ZC, Wang, YY, 2020. A new species of the genus Lycodon (Serpentes, Colubridae) from Guangxi, China. ZooKeys 954: 85-108. https://doi.org/10.3897/zookeys.954.53432" title="A new species of the genus Lycodon (Serpentes, Colubridae) from Guangxi, China. ZooKeys 954: 85 - 108." url="https://doi.org/10.3897/zookeys.954.53432" year="2020">Wang et al. (2020</bibRefCitation>
,
<bibRefCitation DOI="https://doi.org/10.3897/zookeys.954.53432" author="Wang, K" journalOrPublisher="Russian Journal of Herpetology" pageId="0" pageNumber="371" refId="B32" refString="Wang, K, Yu, Z-B, Vogel, G, Che, J, 2021. Contribution to the taxonomy of the genus Lycodon H. Boie in Fitzinger, 1827 (Reptilia: Squamata: Colubridae) in China, with description of two new species and resurrection and elevation of Dinodon septentrionale chapaense Angel, Bourret, 1933. Zoological Research 42: 62-86. https://doi.org/10.3897/zookeys.954.53432" title="Contribution to the taxonomy of the genus Lycodon H. Boie in Fitzinger, 1827 (Reptilia: Squamata: Colubridae) in China, with description of two new species and resurrection and elevation of Dinodon septentrionale chapaense Angel, Bourret, 1933. Zoological Research 42: 62 - 86." url="https://doi.org/10.3897/zookeys.954.53432" year="2021">2021</bibRefCitation>
), and
<bibRefCitation author="Song, H" journalOrPublisher="Zoologica Scripta" pageId="0" pageNumber="371" refId="B26" refString="Song, H, 2021. Sinoophis - Atlas of Snakes in China. (in press) 636 pages." title="Sinoophis - Atlas of Snakes in China. (in press) 636 pages." year="2021">Song (2021)</bibRefCitation>
. Differences between
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
and
<taxonomicName genus="L." lsidName="L. truongi" pageId="0" pageNumber="371" rank="species" species="truongi">
<emphasis italics="true" pageId="0" pageNumber="371">L. truongi</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
are discussed above.
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
differs from all other species of the
<taxonomicName genus="L." lsidName="L. rufozonatus" pageId="0" pageNumber="371" rank="species" species="rufozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. rufozonatus</emphasis>
</taxonomicName>
complex by having 12 or 13 light-colored dorsal bands as opposed to 19-115 collectively in the remaining species. It differs further from all other species in that the body bands are orangish as opposed to white, reddish, yellow, or light-rose in adults.
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
differs from
<taxonomicName genus="L." lsidName="L. chapaensis" pageId="0" pageNumber="371" rank="species" species="chapaensis">
<emphasis italics="true" pageId="0" pageNumber="371">L. chapaensis</emphasis>
</taxonomicName>
and
<taxonomicName genus="L." lsidName="L. septentrionalis" pageId="0" pageNumber="371" rank="species" species="septentrionalis">
<emphasis italics="true" pageId="0" pageNumber="371">L. septentrionalis</emphasis>
</taxonomicName>
in the third, fourth, and fifth supralabials contacting the eye as opposed to the fourth and fifth in the former and second or third in the latter. Having 5-7 light-colored caudal bands also separates it from all other species which collectively have 13-35 caudal bands.
<taxonomicName authorityName="Nguyen &amp; Duong &amp; Wood Jr. &amp; Grismer" authorityYear="2022" class="Reptilia" family="Colubridae" genus="Lycodon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lycodon anakradaya" order="Squamata" pageId="0" pageNumber="371" phylum="Chordata" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">Lycodon anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
differs from
<taxonomicName genus="L." lsidName="L. cathaya" pageId="0" pageNumber="371" rank="species" species="cathaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. cathaya</emphasis>
</taxonomicName>
by have 12 maxillary teeth as opposed to 10 and from
<taxonomicName genus="L." lsidName="L. septentrionalis" pageId="0" pageNumber="371" rank="species" species="septentrionalis">
<emphasis italics="true" pageId="0" pageNumber="371">L. septentrionalis</emphasis>
</taxonomicName>
which has eight. The divided cloacal shield differentiates
<taxonomicName genus="L." lsidName="L. anakradaya" pageId="0" pageNumber="371" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
from
<emphasis italics="true" pageId="0" pageNumber="371">
<taxonomicName genus="L." lsidName="L. banksi" pageId="0" pageNumber="371" rank="species" species="banksi">L. banksi</taxonomicName>
.
<taxonomicName genus="L." lsidName="L. cathaya" pageId="0" pageNumber="371" rank="species" species="cathaya">L. cathaya</taxonomicName>
</emphasis>
,
<taxonomicName genus="L." lsidName="L. chapaensis" pageId="0" pageNumber="371" rank="species" species="chapaensis">
<emphasis italics="true" pageId="0" pageNumber="371">L. chapaensis</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. rufozonatus" pageId="0" pageNumber="371" rank="species" species="rufozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. rufozonatus</emphasis>
</taxonomicName>
, and
<taxonomicName genus="L." lsidName="L. septentrionalis" pageId="0" pageNumber="371" rank="species" species="septentrionalis">
<emphasis italics="true" pageId="0" pageNumber="371">L. septentrionalis</emphasis>
</taxonomicName>
. Having keeled dorsal scales as adults differentiates it from
<taxonomicName genus="L." lsidName="L. cathaya" pageId="0" pageNumber="371" rank="species" species="cathaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. cathaya</emphasis>
</taxonomicName>
and
<taxonomicName genus="L." lsidName="L. futsingensis" pageId="0" pageNumber="371" rank="species" species="futsingensis">
<emphasis italics="true" pageId="0" pageNumber="371">L. futsingensis</emphasis>
</taxonomicName>
. It is further separated from
<taxonomicName genus="L." lsidName="L. meridionalis" pageId="0" pageNumber="371" rank="species" species="meridionalis">
<emphasis italics="true" pageId="0" pageNumber="371">L. meridionalis</emphasis>
</taxonomicName>
by having 223-232 ventral scales versus 199-212 in
<taxonomicName genus="L." lsidName="L. cathaya" pageId="0" pageNumber="371" rank="species" species="cathaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. cathaya</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. futsingensis" pageId="0" pageNumber="371" rank="species" species="futsingensis">
<emphasis italics="true" pageId="0" pageNumber="371">L. futsingensis</emphasis>
</taxonomicName>
, and
<taxonomicName genus="L." lsidName="L. septentrionalis" pageId="0" pageNumber="371" rank="species" species="septentrionalis">
<emphasis italics="true" pageId="0" pageNumber="371">L. septentrionalis</emphasis>
</taxonomicName>
and from
<taxonomicName genus="L." lsidName="L. banksi" pageId="0" pageNumber="371" rank="species" species="banksi">
<emphasis italics="true" pageId="0" pageNumber="371">L. banksi</emphasis>
</taxonomicName>
with 241ventral scales. Its uniform black head pattern differentiates it from
<taxonomicName genus="L." lsidName="L. futsingensis" pageId="0" pageNumber="371" rank="species" species="futsingensis">
<emphasis italics="true" pageId="0" pageNumber="371">L. futsingensis</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. flavozonatus" pageId="0" pageNumber="371" rank="species" species="flavozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. flavozonatus</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. meridionalis" pageId="0" pageNumber="371" rank="species" species="meridionalis">
<emphasis italics="true" pageId="0" pageNumber="371">L. meridionalis</emphasis>
</taxonomicName>
, and
<taxonomicName genus="L." lsidName="L. rufozonatus" pageId="0" pageNumber="371" rank="species" species="rufozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. rufozonatus</emphasis>
</taxonomicName>
. It is separated from
<taxonomicName genus="L." lsidName="L. banksi" pageId="0" pageNumber="371" rank="species" species="banksi">
<emphasis italics="true" pageId="0" pageNumber="371">L. banksi</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. cathaya" pageId="0" pageNumber="371" rank="species" species="cathaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. cathaya</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. flavozonatus" pageId="0" pageNumber="371" rank="species" species="flavozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. flavozonatus</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. meridonalis" pageId="0" pageNumber="371" rank="species" species="meridonalis">
<emphasis italics="true" pageId="0" pageNumber="371">L. meridonalis</emphasis>
</taxonomicName>
, and
<taxonomicName genus="L." lsidName="L. rufozonatus" pageId="0" pageNumber="371" rank="species" species="rufozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. rufozonatus</emphasis>
</taxonomicName>
by having the ventral portion of the orangish body bands forming the pattern on the ventrolateral side of the body as opposed to a light-colored reticulum. It differs from
<taxonomicName genus="L." lsidName="L. banksi" pageId="0" pageNumber="371" rank="species" species="banksi">
<emphasis italics="true" pageId="0" pageNumber="371">L. banksi</emphasis>
</taxonomicName>
by not having an orangish brown or uniform grey-colored venter. It differs further from
<taxonomicName genus="L." lsidName="L. cathaya" pageId="0" pageNumber="371" rank="species" species="cathaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. cathaya</emphasis>
</taxonomicName>
and
<taxonomicName genus="L." lsidName="L. futsingnesis" pageId="0" pageNumber="371" rank="species" species="futsingnesis">
<emphasis italics="true" pageId="0" pageNumber="371">L. futsingnesis</emphasis>
</taxonomicName>
by not having a wide light-colored nuchal-occipital collar. Having wide light-colored body bands separates
<taxonomicName genus="L." lsidName="L. anakradaya" pageId="0" pageNumber="371" rank="species" species="anakradaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. anakradaya</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="371">sp. nov.</emphasis>
from
<taxonomicName genus="L." lsidName="L. banksi" pageId="0" pageNumber="371" rank="species" species="banksi">
<emphasis italics="true" pageId="0" pageNumber="371">L. banksi</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. cathaya" pageId="0" pageNumber="371" rank="species" species="cathaya">
<emphasis italics="true" pageId="0" pageNumber="371">L. cathaya</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. chapaensis" pageId="0" pageNumber="371" rank="species" species="chapaensis">
<emphasis italics="true" pageId="0" pageNumber="371">L. chapaensis</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. flavozonatus" pageId="0" pageNumber="371" rank="species" species="flavozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. flavozonatus</emphasis>
</taxonomicName>
,
<taxonomicName genus="L." lsidName="L. meridonalis" pageId="0" pageNumber="371" rank="species" species="meridonalis">
<emphasis italics="true" pageId="0" pageNumber="371">L. meridonalis</emphasis>
</taxonomicName>
, and
<taxonomicName genus="L." lsidName="L. rufozonatus" pageId="0" pageNumber="371" rank="species" species="rufozonatus">
<emphasis italics="true" pageId="0" pageNumber="371">L. rufozonatus</emphasis>
</taxonomicName>
and
<taxonomicName genus="L." lsidName="L. septentrionalis" pageId="0" pageNumber="371" rank="species" species="septentrionalis">
<emphasis italics="true" pageId="0" pageNumber="371">L. septentrionalis</emphasis>
</taxonomicName>
which have narrow body bands.
</paragraph>
</subSubSection>
</treatment>
</subSection>
</document>
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