treatments-xml/data/37/55/34/375534006E3E558FB0B7515FC0FE43A0.xml

<document ID-DOI="http://dx.doi.org/10.3897/zookeys.935.37138" ID-GBIF-Dataset="1f378201-712a-46c0-bbe6-8e5e578c61dd" ID-PMC="PMC7256073" ID-Pensoft-Pub="1313-2970-935-121" ID-Pensoft-UUID="33263B29FD6E5F15BCFC61C058095F9C" ID-PubMed="32508505" ID-ZooBank="4D41FCF4A59544F095EFCF38D0FC127C" ModsDocID="1313-2970-935-121" checkinTime="1590112769381" checkinUser="pensoft" docAuthor="Bernardes, Marta, Le, Minh Duc, Nguyen, Truong Quang, Pham, Cuong The, Pham, Anh Van, Nguyen, Tao Thien, Roedder, Dennis, Bonkowski, Michael &amp; Ziegler, Thomas" docDate="2020" docId="375534006E3E558FB0B7515FC0FE43A0" docLanguage="en" docName="ZooKeys 935: 121-164" docOrigin="ZooKeys 935" docSource="http://dx.doi.org/10.3897/zookeys.935.37138" docTitle="Tylototriton sparreboomi M. Bernardes, M. D. Le, T. Q. Nguyen, C. T. Pham, A. V. Pham, T. T. Nguyen &amp; T. Ziegler 2020, sp. nov." docType="treatment" docUuid="4599D131-7C89-4D62-B43A-15E24C6473B9" docUuidSource="ZooBank" docVersion="5" id="33263B29FD6E5F15BCFC61C058095F9C" lastPageNumber="121" masterDocId="33263B29FD6E5F15BCFC61C058095F9C" masterDocTitle="Integrative taxonomy reveals three new taxa within the Tylototriton asperrimus complex (Caudata, Salamandridae) from Vietnam" masterLastPageNumber="164" masterPageNumber="121" pageNumber="121" updateTime="1668168868404" updateUser="ExternalLinkService">
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<mods:title>Integrative taxonomy reveals three new taxa within the Tylototriton asperrimus complex (Caudata, Salamandridae) from Vietnam</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bernardes, Marta</mods:namePart>
<mods:affiliation>Cologne Zoo, Riehler Str. 173, 50735 Cologne, Germany &amp; Terrestrial Ecology, Institute of Zoology, University of Cologne, Zuelpicher Str. 47 b, 50674 Cologne, Germany</mods:affiliation>
<mods:nameIdentifier type="email">mrtbernardes@gmail.com</mods:nameIdentifier>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Le, Minh Duc</mods:namePart>
<mods:affiliation>Faculty of Environmental Sciences, University of Science, Vietnam National University, Hanoi, 334 Nguyen Trai Road, Hanoi, Vietnam &amp; Central Institute for Natural Resources and Environmental Studies, Vietnam National University, Hanoi, 19 Le Thanh Tong, Hanoi, Vietnam &amp; Department of Herpetology, American Museum of Natural History, Central Park West at 79 th Street, New York, New York 10024, USA</mods:affiliation>
</mods:name>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Nguyen, Truong Quang</mods:namePart>
<mods:affiliation>Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, 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:nameIdentifier type="orcid">https://orcid.org/0000-0002-6601-0880</mods:nameIdentifier>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Pham, Cuong The</mods:namePart>
<mods:affiliation>Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, 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>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Pham, Anh Van</mods:namePart>
<mods:affiliation>Faculty of Natural Science and Technology, Tay Bac University, Quyet Tam Ward, Son La City, Son La Province, Vietnam</mods:affiliation>
</mods:name>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Nguyen, Tao Thien</mods:namePart>
<mods:affiliation>Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Hanoi, Vietnam &amp; Vietnam National Museum of Nature, 18 Hoang Quoc Viet St., Hanoi, Vietnam</mods:affiliation>
</mods:name>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Roedder, Dennis</mods:namePart>
<mods:affiliation>Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany</mods:affiliation>
</mods:name>
<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Bonkowski, Michael</mods:namePart>
<mods:affiliation>Terrestrial Ecology, Institute of Zoology, University of Cologne, Zuelpicher Str. 47 b, 50674 Cologne, Germany</mods:affiliation>
</mods:name>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Ziegler, Thomas</mods:namePart>
<mods:affiliation>Cologne Zoo, Riehler Str. 173, 50735 Cologne, Germany &amp; Terrestrial Ecology, Institute of Zoology, University of Cologne, Zuelpicher Str. 47 b, 50674 Cologne, Germany</mods:affiliation>
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<mods:date>2020</mods:date>
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<mods:number>935</mods:number>
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<treatment ID-GBIF-Taxon="164248502" LSID="urn:lsid:zoobank.org:act:4599D131-7C89-4D62-B43A-15E24C6473B9" httpUri="http://treatment.plazi.org/id/375534006E3E558FB0B7515FC0FE43A0" lastPageNumber="121" pageId="0" pageNumber="121">
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<paragraph pageId="0" pageNumber="121">
<taxonomicName LSID="37553400-6E3E-558F-B0B7-515FC0FE43A0" authority="M. Bernardes, M. D. Le, T. Q. Nguyen, C. T. Pham, A. V. Pham, T. T. Nguyen &amp; T. Ziegler" authorityName="M. Bernardes, M. D. Le, T. Q. Nguyen, C. T. Pham, A. V. Pham, T. T. Nguyen &amp; T. Ziegler" authorityYear="2020" class="Amphibia" family="Salamandridae" genus="Tylototriton" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tylototriton sparreboomi" order="Caudata" pageId="0" pageNumber="121" phylum="Chordata" rank="species" species="sparreboomi" status="sp. nov.">Tylototriton sparreboomi M. Bernardes, M. D. Le, T. Q. Nguyen, C. T. Pham, A. V. Pham, T.T. Nguyen &amp; T. Ziegler</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="121">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 9" captionStartId="F10" captionText="Figure 9. Dorsal and ventral views of the new taxa in life. Top: Tylototriton pasmansi sp. nov. (holotype); Center: Tylototriton pasmansi obsti ssp. nov. (holotype); Bottom: Tylototriton sparreboomi sp. nov. Photographs: C. T. Pham (upper row) and A. V. Pham (central and lower rows)." figureDoi="10.3897/zookeys.935.37138.figure9" httpUri="https://binary.pensoft.net/fig/412605" pageId="0" pageNumber="121">Figures 9</figureCitation>
<figureCitation captionStart="Figure 11" captionStartId="F13" captionText="Figure 11. Tylototriton sparreboomi sp. nov. (holotype). In sequence: dorsal view; ventral view; lateral view; and detail of dorsal view of the head. Photographs T. Ziegler." figureDoi="10.3897/zookeys.935.37138.figure11" httpUri="https://binary.pensoft.net/fig/412607" pageId="0" pageNumber="121">, 11</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="reference_group">
<paragraph pageId="0" pageNumber="121">
<taxonomicName authorityName="Anderson" authorityYear="1871" class="Amphibia" family="Salamandridae" genus="Tylototriton" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tylototriton" order="Caudata" pageId="0" pageNumber="121" phylum="Chordata" rank="genus">Tylototriton</taxonomicName>
taxon 3 (this study).
</paragraph>
<paragraph pageId="0" pageNumber="121">
<taxonomicName genus="T." higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="T." pageId="0" pageNumber="121" rank="genus">T.</taxonomicName>
sp.: 
<bibRefCitation DOI="https://doi.org/10.1038/srep44443" author="Laking, AE" journalOrPublisher="Scientific Reports" pageId="0" pageNumber="121" pagination="1 - 5" refId="B25" refString="Laking, AE, Ngo, HN, Pasmans, F, Martel, A, Nguyen, TT, 2017. Batrachochytrium salamandrivorans is the predominant chytrid fungus in Vietnamese salamanders. Scientific Reports 7 (44443): 1 - 5, DOI: https://doi.org/10.1038/srep44443" title="Batrachochytrium salamandrivorans is the predominant chytrid fungus in Vietnamese salamanders." url="https://doi.org/10.1038/srep44443" volume="7" year="2017">Laking et al. 2017</bibRefCitation>
, page 2.
</paragraph>
<paragraph pageId="0" pageNumber="121">
<taxonomicName higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="T. verrucosus" pageId="0" pageNumber="121" rank="species" species="verrucosus">T. verrucosus</taxonomicName>
: 
<bibRefCitation author="Orlov, N" journalOrPublisher="Russian Journal of Herpetology" pageId="0" pageNumber="121" pagination="81 - 104" refId="B36" refString="Orlov, N, Murphy, RW, Ananjeva, NB, Ryabov, SA, Cuc, HT, 2002. Herpetofauna of Vietnam, a checklist. Part I Amphibia. Russian Journal of Herpetology 9 (2): 81 - 104" title="Herpetofauna of Vietnam, a checklist. Part I Amphibia." volume="9" year="2002">Orlov et al. 2002</bibRefCitation>
, page 101; 
<bibRefCitation author="van Dijk, PP" journalOrPublisher="Nucleic Acids Research" pageId="0" pageNumber="121" refId="B57" refString="van Dijk, PP, Wogan, G, Lau, MWN, Dutta, S, Shrestha, TK, Roy, D, Truong, NQ, 2009. Tylototritonverrucosus. IUCN Red List Threatened Species 2009 e.T59487A11934912." title="Tylototritonverrucosus. IUCN Red List Threatened Species 2009 e. T 59487 A 11934912." year="2009">van Dijk et al. 2009</bibRefCitation>
, page 1; 
<bibRefCitation author="Nguyen, TQ" journalOrPublisher="Vietnam Journal of Biotechnology" pageId="0" pageNumber="121" pagination="325 - 333" refId="B30" refString="Nguyen, TQ, Nguyen, VS, Ho, LT, Le, QK, Nguyen, TT, 2009. Phylogenetic relationships and taxonomic review of the family Salamandridae (Amphibia: Caudata) from Vietnam (in Vietnamese). Vietnam Journal of Biotechnology 7 (3): 325 - 333" title="Phylogenetic relationships and taxonomic review of the family Salamandridae (Amphibia: Caudata) from Vietnam (in Vietnamese)." volume="7" year="2009">Nguyen et al. 2009</bibRefCitation>
, page 329.
</paragraph>
<paragraph pageId="0" pageNumber="121">
<taxonomicName higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="T. asperrimus" pageId="0" pageNumber="121" rank="species" species="asperrimus">T. cf. asperrimus</taxonomicName>
&quot;North Vietnam&quot;: 
<bibRefCitation author="Hernandez, A" journalOrPublisher="Bulletin de la Societe Herpetologique de France" pageId="0" pageNumber="121" pagination="79 - 82" refId="B15" refString="Hernandez, A, 2018. Ecological affinities of Tylototriton asperrimus Unterstein, 1930 (Amphibia, Caudata: Salamandridae) at Bainiu, Dayaoshan, Guangxi Province, China with an emphasis on its cryptic diversity. Bulletin de la Societe Herpetologique de France 166 (2018): 79 - 82" title="Ecological affinities of Tylototriton asperrimus Unterstein, 1930 (Amphibia, Caudata: Salamandridae) at Bainiu, Dayaoshan, Guangxi Province, China with an emphasis on its cryptic diversity." volume="166" year="2018">Hernandez 2018</bibRefCitation>
, page 80.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="holotype">
<paragraph pageId="0" pageNumber="121">Holotype.</paragraph>
<paragraph pageId="0" pageNumber="121">IEBR 4476, adult male, collected in Sa De Phin Commune, Sin Ho District, Lai Chau Province, Vietnam, at an elevation of 1670 m a.s.l., in May 2015 by A. V. Pham and M. A. Vang.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="paratypes">
<paragraph pageId="0" pageNumber="121">Paratypes.</paragraph>
<paragraph pageId="0" pageNumber="121">Two adult males, same data as the holotype: IEBR 4477 and TBU 10; two adult males, collector unknown: IEBR 4478 and IEBR 4479.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="etymology">
<paragraph pageId="0" pageNumber="121">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="121">
The specific epithet is dedicated to late Prof. Dr. Max Sparreboom, who has made great contributions to the understanding of 
<taxonomicName lsidName="" order="Urodela" pageId="0" pageNumber="121" rank="order">Urodela</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="diagnosis">
<paragraph pageId="0" pageNumber="121">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="121">The new species is distinguished from other species of the genus by the following combination of characters: head longer than wide; snout truncate in dorsal view; tips of fingers reaching nostril when foreleg is laid forward; skin tubercles on ventral side shaped like transverse wrinkles; rib nodules distinct and round; vertebral ridge segmented, high and relatively wide; relatively wide distance between the eyes; and gular and labial folds present.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="description">
<paragraph pageId="0" pageNumber="121">Description of holotype.</paragraph>
<paragraph pageId="0" pageNumber="121">Habitus stout; head broader than body, longer than wide, depressed and slightly oblique in profile; snout wider than long (IN&gt; EN), truncate in dorsal view, rounded in profile and protruding beyond lower jaw; nostrils close to snout tip and not visible from above; labial fold slightly evident; dorsolateral bony ridges on head prominent, wide, moderately protruding, from above eye to above anterior end of parotoid, posterior ends slightly scrolled inside; middorsal ridge on head almost indistinct; parotoids enlarged, projecting backwards; ventral skin smoother than dorsal skin, with tubercles shaped like transverse wrinkles; gular fold weak; glandular vertebral ridge high, wide, smooth and segmented extending from top of head to base of tail, separated from middorsal ridge; number of trunk vertebrae 13; rib nodules distinct and roundish, the third anterior rib nodule on right side is located below the second nodule and the fourth nodule seems to not be associated with the fourth vertebra, nodules appear knob-like anteriorly, becoming smaller posteriorly; tips of fore- and hind limbs overlap when adpressed along body; tips of fingers reaching nostril when foreleg laid forward; and tail laterally compressed, thin and tip acuminated.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="color of holotype">
<paragraph pageId="0" pageNumber="121">Color of holotype.</paragraph>
<paragraph pageId="0" pageNumber="121">
In preservative, with an overall faded dark brown coloration, with faded yellow markings on vent, ventral margin of tail, tips of fingers and toes, and part of palms. For color in life see Fig. 
<figureCitation captionStart="Figure 9" captionStartId="F10" captionText="Figure 9. Dorsal and ventral views of the new taxa in life. Top: Tylototriton pasmansi sp. nov. (holotype); Center: Tylototriton pasmansi obsti ssp. nov. (holotype); Bottom: Tylototriton sparreboomi sp. nov. Photographs: C. T. Pham (upper row) and A. V. Pham (central and lower rows)." figureDoi="10.3897/zookeys.935.37138.figure9" httpUri="https://binary.pensoft.net/fig/412605" pageId="0" pageNumber="121">9</figureCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="description">
<paragraph pageId="0" pageNumber="121">Measurements of holotype (in mm).</paragraph>
<paragraph pageId="0" pageNumber="121">SVL 68.71; MHW 17.60; HW 9.85; HL 19.95; PL 10.18; PH 6.12; EL 3.43; EN 4.43; IN 6.26; IE 9.04; LJL 10.74; UEL 4.74; HUM 9.27; RAD 15.86; FEM 8.77; TIB 16.03; TL 59.70; TH 8.57; ClL 9.66; WVr 2.37; L5W 3.04; AG 30.99; and TkL 42.72.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="variation">
<paragraph pageId="0" pageNumber="121">Variation.</paragraph>
<paragraph pageId="0" pageNumber="121">
TBU 10 (in worse preserved condition) presents rib-nodules thinner than holotype, glandular vertebral ridge more tubercular, and tail tip slightly rounded. The remaining characters were similar to the holotype in morphology. For detailed measurements see Table 
<tableCitation captionStart="Table 4" captionStartId="T4" captionText="Table 4. Morphological measurements between the Tylototriton males from Son La and Thanh Hoa provinces (taxon 1), from Phu Tho and Hoa Binh provinces (taxon 2), and from Lai Chau Province (taxon 3). Measures as absolute values (in mm) and ratios of characters to snout vent length (% SVL). Values are presented as mean + / - standard deviation above minimum and maximum ranges, and for abbreviations see Materials and methods." httpUri="http://table.plazi.org/id/9846492791068CD2D89D2E0F9846C65D" pageId="0" pageNumber="121" tableUuid="9846492791068CD2D89D2E0F9846C65D">4</tableCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="comparisons">
<paragraph pageId="0" pageNumber="121">Comparisons.</paragraph>
<paragraph pageId="0" pageNumber="121">
<taxonomicName authorityName="M. Bernardes, M. D. Le, T. Q. Nguyen, C. T. Pham, A. V. Pham, T. T. Nguyen &amp; T. Ziegler" authorityYear="2020" class="Amphibia" family="Salamandridae" genus="Tylototriton" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tylototriton sparreboomi" order="Caudata" pageId="0" pageNumber="121" phylum="Chordata" rank="species" species="sparreboomi">
<emphasis italics="true" pageId="0" pageNumber="121">Tylototriton sparreboomi</emphasis>
</taxonomicName>
sp. nov. differs from other related species of 
<taxonomicName authorityName="Anderson" authorityYear="1871" class="Amphibia" family="Salamandridae" genus="Tylototriton" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tylototriton" order="Caudata" pageId="0" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="121">Tylototriton</emphasis>
</taxonomicName>
as follows: from 
<taxonomicName lsidName="T. anhuiensis" pageId="0" pageNumber="121" rank="species" species="anhuiensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. anhuiensis</emphasis>
</taxonomicName>
by distinctly separated rib nodules (versus continuous nodule-like warts in 
<taxonomicName lsidName="T. anhuiensis" pageId="0" pageNumber="121" rank="species" species="anhuiensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. anhuiensis</emphasis>
</taxonomicName>
); from 
<taxonomicName lsidName="T. asperrimus" pageId="0" pageNumber="121" rank="species" species="asperrimus">
<emphasis italics="true" pageId="0" pageNumber="121">T. asperrimus</emphasis>
</taxonomicName>
by a head longer than wide (versus wider than long in 
<taxonomicName lsidName="T. asperrimus" pageId="0" pageNumber="121" rank="species" species="asperrimus">
<emphasis italics="true" pageId="0" pageNumber="121">T. asperrimus</emphasis>
</taxonomicName>
according to 
<bibRefCitation DOI="https://doi.org/10.5358/hsj.32.34" author="Nishikawa, K" journalOrPublisher="Current Herpetology" pageId="0" pageNumber="121" pagination="34 - 49" refId="B32" refString="Nishikawa, K, Matsui, M, Nguyen, TT, 2013b. A new species of Tylototriton from Northern Vietnam (Amphibia: Urodela: Salamandridae). Current Herpetology 32 (1): 34 - 49, DOI: https://doi.org/10.5358/hsj.32.34" title="A new species of Tylototriton from Northern Vietnam (Amphibia: Urodela: Salamandridae)." url="https://doi.org/10.5358/hsj.32.34" volume="32" year="2013 b">Nishikawa et al. 2013b</bibRefCitation>
; 
<bibRefCitation DOI="https://doi.org/10.1163/9789004285620" author="Sparreboom, M" journalOrPublisher="Knnv Publishing, Zeist, Netherlands" pageId="0" pageNumber="121" refId="B50" refString="Sparreboom, M, 2014. Salamanders of the Old World: the salamanders of Europe, Asia and northern Africa. Knnv Publishing, Zeist, Netherlands, DOI: https://doi.org/10.1163/9789004285620" title="Salamanders of the Old World: the salamanders of Europe, Asia and northern Africa." url="https://doi.org/10.1163/9789004285620" year="2014">Sparreboom 2014</bibRefCitation>
; 
<bibRefCitation author="Hernandez, A" journalOrPublisher="Edition Chimaira, Frankfurt" pageId="0" pageNumber="121" refId="B14" refString="Hernandez, A, 2016. Crocodile Newts, The Primitive Salamandridae from Asia (Genera Echinotriton and Tylototriton). Edition Chimaira, Frankfurt" title="Crocodile Newts, The Primitive Salamandridae from Asia (Genera Echinotriton and Tylototriton)." year="2016">Hernandez 2016</bibRefCitation>
), however the female holotype shows similar head proportions-see Discussion); from 
<taxonomicName lsidName="T. broadoridgus" pageId="0" pageNumber="121" rank="species" species="broadoridgus">
<emphasis italics="true" pageId="0" pageNumber="121">T. broadoridgus</emphasis>
</taxonomicName>
by head slightly longer than wide (versus equally long and wide), presence of skin on ventral side shaped like transverse wrinkles (versus covered with round shaped tubercles, like the dorsal side), distinctly separated rib nodules (versus continuous nodule-like warts), and narrower vertebral ridge (versus wider vertebral ridge in 
<taxonomicName lsidName="T. broadoridgus" pageId="0" pageNumber="121" rank="species" species="broadoridgus">
<emphasis italics="true" pageId="0" pageNumber="121">T. broadoridgus</emphasis>
</taxonomicName>
); from 
<taxonomicName lsidName="T. hainanensis" pageId="0" pageNumber="121" rank="species" species="hainanensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. hainanensis</emphasis>
</taxonomicName>
by a head longer than wide (versus much wider than long), tips of fingers reaching nostril (versus eyes) when foreleg is laid forward, and a snout truncate in dorsal view (versus rounded in 
<taxonomicName lsidName="T. hainanensis" pageId="0" pageNumber="121" rank="species" species="hainanensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. hainanensis</emphasis>
</taxonomicName>
); from 
<taxonomicName lsidName="T. liuyangensis" pageId="0" pageNumber="121" rank="species" species="liuyangensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. liuyangensis</emphasis>
</taxonomicName>
by a wider (versus shorter) distance between eyes, distinctly separated rib nodules (versus continuous nodule-like warts), and lateral skin shaped like transverse wrinkles (versus covered by warts in 
<taxonomicName lsidName="T. liuyangensis" pageId="0" pageNumber="121" rank="species" species="liuyangensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. liuyangensis</emphasis>
</taxonomicName>
); from 
<taxonomicName lsidName="T. notialis" pageId="0" pageNumber="121" rank="species" species="notialis">
<emphasis italics="true" pageId="0" pageNumber="121">T. notialis</emphasis>
</taxonomicName>
by a broader (versus narrower) head, longer (versus shorter) hind-limbs, and higher tail (versus thinner tail in 
<taxonomicName lsidName="T. notialis" pageId="0" pageNumber="121" rank="species" species="notialis">
<emphasis italics="true" pageId="0" pageNumber="121">T. notialis</emphasis>
</taxonomicName>
); from 
<taxonomicName lsidName="T. panhai" pageId="0" pageNumber="121" rank="species" species="panhai">
<emphasis italics="true" pageId="0" pageNumber="121">T. panhai</emphasis>
</taxonomicName>
by wider (versus shorter) distance between the eyes, presence (versus absence) of labial fold, and overall dorsal coloration mostly dark (versus with presence of characteristic dorsal colorful markings in 
<taxonomicName lsidName="T. panhai" pageId="0" pageNumber="121" rank="species" species="panhai">
<emphasis italics="true" pageId="0" pageNumber="121">T. panhai</emphasis>
</taxonomicName>
); from 
<taxonomicName lsidName="T. pasmansi" pageId="0" pageNumber="121" rank="species" species="pasmansi">
<emphasis italics="true" pageId="0" pageNumber="121">T. pasmansi</emphasis>
</taxonomicName>
sensu lato by a narrower (versus wider) head, slightly wider (versus slightly narrower) distance between the eyes, tips of fingers reaching nostril (versus eye) when foreleg laid forward, longer (versus shorter) humerus length, and slightly enlarged round-like rib nodules (versus slightly smaller, pointy to rounded rib nodules in 
<taxonomicName lsidName="T. pasmansi" pageId="0" pageNumber="121" rank="species" species="pasmansi">
<emphasis italics="true" pageId="0" pageNumber="121">T. pasmansi</emphasis>
</taxonomicName>
sensu lato); from 
<taxonomicName lsidName="T. p. subsp. pasmansi" pageId="0" pageNumber="121" rank="subSpecies" species="p." subSpecies="pasmansi">
<emphasis italics="true" pageId="0" pageNumber="121">T. p. pasmansi</emphasis>
</taxonomicName>
by a longer (versus shorter) length between eye and nostril and wider (versus narrower) vertebral ridge in 
<taxonomicName lsidName="T. p. subsp. pasmansi" pageId="0" pageNumber="121" rank="subSpecies" species="p." subSpecies="pasmansi">
<emphasis italics="true" pageId="0" pageNumber="121">T. p. pasmansi</emphasis>
</taxonomicName>
; from 
<taxonomicName lsidName="T. pasmansi subsp. obsti" pageId="0" pageNumber="121" rank="subSpecies" species="pasmansi" subSpecies="obsti">
<emphasis italics="true" pageId="0" pageNumber="121">T. pasmansi obsti</emphasis>
</taxonomicName>
by a longer (versus shorter) femur length; from 
<taxonomicName lsidName="T. vietnamensis" pageId="0" pageNumber="121" rank="species" species="vietnamensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. vietnamensis</emphasis>
</taxonomicName>
by a moderately stout (versus slender) habitus, presence (versus absence) of gular fold, and round (versus slightly flattened) rib nodules and high vertebral ridge (versus low vertebral ridge in 
<taxonomicName lsidName="T. vietnamensis" pageId="0" pageNumber="121" rank="species" species="vietnamensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. vietnamensis</emphasis>
</taxonomicName>
); from 
<taxonomicName lsidName="T. wenxianensis" pageId="0" pageNumber="121" rank="species" species="wenxianensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. wenxianensis</emphasis>
</taxonomicName>
by a truncate (versus more rounded) snout in dorsal view, wider (versus narrower) distance between the eyes, distinctly separated rib nodules (versus continuous nodule-like warts), smoother (versus extremely rough) skin on ventral side shaped like transverse wrinkles (versus rounded shaped and uniform to dorsal side), and colored marking on ventral slit (versus black colored ventral slit in 
<taxonomicName lsidName="T. wenxianensis" pageId="0" pageNumber="121" rank="species" species="wenxianensis">
<emphasis italics="true" pageId="0" pageNumber="121">T. wenxianensis</emphasis>
</taxonomicName>
); from 
<taxonomicName lsidName="T. ziegleri" pageId="0" pageNumber="121" rank="species" species="ziegleri">
<emphasis italics="true" pageId="0" pageNumber="121">T. ziegleri</emphasis>
</taxonomicName>
by a head longer than wide (versus wider than long), rounded but smaller (versus enlarged knob-like) rib nodules, and distinctly segmented vertebral ridge (versus even more segmented vertebral ridge in 
<taxonomicName lsidName="T. ziegleri" pageId="0" pageNumber="121" rank="species" species="ziegleri">
<emphasis italics="true" pageId="0" pageNumber="121">T. ziegleri</emphasis>
</taxonomicName>
).
</paragraph>
<caption doi="10.3897/zookeys.935.37138.figure11" httpUri="https://binary.pensoft.net/fig/412607" pageId="0" pageNumber="121" start="Figure 11" startId="F13">
<paragraph pageId="0" pageNumber="121">
<emphasis bold="true" pageId="0" pageNumber="121">Figure 11.</emphasis>
<taxonomicName authorityName="M. Bernardes, M. D. Le, T. Q. Nguyen, C. T. Pham, A. V. Pham, T. T. Nguyen &amp; T. Ziegler" authorityYear="2020" class="Amphibia" family="Salamandridae" genus="Tylototriton" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tylototriton sparreboomi" order="Caudata" pageId="0" pageNumber="121" phylum="Chordata" rank="species" species="sparreboomi">
<emphasis italics="true" pageId="0" pageNumber="121">Tylototriton sparreboomi</emphasis>
</taxonomicName>
sp. nov. (holotype). In sequence: dorsal view; ventral view; lateral view; and detail of dorsal view of the head. Photographs T. Ziegler.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="distribution">
<paragraph pageId="0" pageNumber="121">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="121">
Known only from the type locality in Lai Chau Province, northern Vietnam (Fig. 
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Distribution map of the new populations of Tylototriton from North Vietnam, based on the following symbols: square (taxon 3, this study) the population from Sin Ho District, Lai Chau Province; diamond (taxon 1, this study) the upper one identifies the population from Van Ho District, Son La Province, and the lower one identifies the population from Thuong Xuan District, Thanh Hoa Province; triangle (taxon 2, this study) the upper one identifies the population from Tan Son District, Phu Tho Province, the middle one identifies the population from Da Bac District, Hoa Binh Province, and the lower one identifies the population from Lac Son District, Hoa Binh Province. The two populations identified by the circles represent T. asperrimus sensu stricto from China. High resolution remote sensing land cover information was extracted from &quot; GLAD-UMD and SERVIR-Mekong, Natural annual tree canopy structure and surface water dynamics products, 2017 &quot; (lower left panel). Bioclimatic variables (right side) were extracted from remote sensing data provided by Deblauwe et al. (2016)." figureDoi="10.3897/zookeys.935.37138.figure2" httpUri="https://binary.pensoft.net/fig/412499" pageId="0" pageNumber="121">2</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="121" type="natural history">
<paragraph pageId="0" pageNumber="121">Natural history.</paragraph>
<paragraph pageId="0" pageNumber="121">
Specimens were found in water between 9:00 and 16:30 h in ponds. The surrounding habitat was secondary forest of large, medium and small hardwoods mixed with shrubs and vines. Air temperature at the sites was 23 to 27 °C and relative humidity was 80 to 85%. Based on remote sensing information, the species occurs at sites with an annual mean temperature of 19.3 °C, ranging from 11.6 to 26.1 °C during the year. Annual precipitation is about 1843.7 mm with yearly variations from 9.6 to 421.6 mm. Further bioclimatic information is provided in Table 
<tableCitation captionStart="Table 6" captionStartId="T6" captionText="Table 6. Bioclimatic conditions at the species records. Abbreviations: Annual Mean Temperature BIO 1, Mean Diurnal Range BIO 2, Isothermality BIO 3, Temperature Seasonality BIO 4, Max Temperature of Warmest Month BIO 5, Min Temperature of Coldest Month BIO 6, Temperature Annual Range BIO 7, Mean Temperature of Wettest Quarter BIO 8, Mean Temperature of Driest Quarter BIO 9, Mean Temperature of Warmest Quarter BIO 10, Mean Temperature of Coldest Quarter BIO 11, Annual Precipitation BIO 12, Precipitation of Wettest Month BIO 13, Precipitation of Driest Month BIO 14, Precipitation Seasonality BIO 15, Precipitation of Wettest Quarter BIO 16, Precipitation of Driest Quarter BIO 17, Precipitation of Warmest Quarter BIO 18, and Precipitation of Coldest Quarter BIO 19." httpUri="http://table.plazi.org/id/BB144C8D4CD7D39BB267CAA03AB15FE3" pageId="0" pageNumber="121" tableUuid="BB144C8D4CD7D39BB267CAA03AB15FE3">6</tableCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>