<documentID-CLB-Dataset="3828"ID-DOI="http://dx.doi.org/10.3897/zookeys.1061.70420"ID-GBIF-Dataset="74154259-34a6-4153-93ce-447ac1a90a27"ID-PMC="PMC8505391"ID-Pensoft-Pub="1313-2970-1061-87"ID-Pensoft-UUID="AF38CEA80126576FA4C341BB14489336"ID-PubMed="34707454"ID-ZooBank="F2701BACD2E14F97A0BDDC8B09966247"ModsDocID="1313-2970-1061-87"checkinTime="1633431410146"checkinUser="pensoft"docAuthor="Shi, Jing-Song, Liu, Jin-Cheng, Giri, Rohit, Owens, John Benjamin, Santra, Vishal, Kuttalam, Sourish, Selvan, Melvin, Guo, Ke-Ji & Malhotra, Anita"docDate="2021"docId="01E6E10AADF759CAA589BD6DFEDF3CAA"docLanguage="en"docName="ZooKeys 1061: 87-108"docOrigin="ZooKeys 1061"docPubDate="2021-10-04"docSource="http://dx.doi.org/10.3897/zookeys.1061.70420"docTitle="Gloydius lipipengi Shi, Liu & Malhotra 2021, sp. nov."docType="treatment"docUuid="6DF30D06-937B-470B-AFE4-D4CABEAF7DAB"docUuidSource="ZooBank"docVersion="6"id="AF38CEA80126576FA4C341BB14489336"lastPageNumber="87"masterDocId="AF38CEA80126576FA4C341BB14489336"masterDocTitle="Molecular phylogenetic analysis of the genus Gloydius (Squamata, Viperidae, Crotalinae), with description of two new alpine species from Qinghai-Tibet Plateau, China"masterLastPageNumber="108"masterPageNumber="87"pageNumber="87"updateTime="1732846796274"updateUser="ExternalLinkService">
<mods:titleid="7644E26EFED47806FC8465FAB73C0F93">Molecular phylogenetic analysis of the genus Gloydius (Squamata, Viperidae, Crotalinae), with description of two new alpine species from Qinghai-Tibet Plateau, China</mods:title>
<mods:affiliationid="5D47E3FEC7D830380EA980D686468CA0">Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China & Institute of Herpetology, Shenyang Normal University, Shenyang 110034, China</mods:affiliation>
<mods:affiliationid="D432E46BB4FED05984151583B22B36E2">Molecular Ecology and Evolution at Bangor, School of Natural Sciences, Bangor University, Bangor, Gwynedd LL 57 2 UW, UK & Captive and Field Herpetology, Wales, UK</mods:affiliation>
<mods:affiliationid="7C908BE5FD33E624E6291F06EB9FF05C">Captive and Field Herpetology, Wales, UK & Society for Nature Conservation, Research and Community Engagement (CONCERN), Nalikul, Hooghly, West Bengal, India</mods:affiliation>
<mods:affiliationid="4162C6E6C7EF4EFD1831BD3B2B054A8D">Captive and Field Herpetology, Wales, UK & Society for Nature Conservation, Research and Community Engagement (CONCERN), Nalikul, Hooghly, West Bengal, India</mods:affiliation>
<mods:affiliationid="A04EE12B6B5D19806EB9E9C9A56EBEB7">Central South Inventory and Planning Institute of National Forestry and Grassland Administration, College of Life Sciences and Technology, Central South University of Forestry and Technology, Changsha, Hunan 410004, China</mods:affiliation>
<mods:affiliationid="310E845E9B7436ED16466C807673FE33">Molecular Ecology and Evolution at Bangor, School of Natural Sciences, Bangor University, Bangor, Gwynedd LL 57 2 UW, UK</mods:affiliation>
<taxonomicNameid="B8EEA176045D650B97AAAE82F63AC64C"ID-CoL="8SJNQ"LSID="http://zoobank.org/6DF30D06-937B-470B-AFE4-D4CABEAF7DAB"authority="Shi, Liu & Malhotra"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"higherTaxonomySource="treatment-meta"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi"status="sp. nov.">Gloydius lipipengi Shi, Liu & Malhotra</taxonomicName>
sixtieth birthday. Prof. Li has devoted himself to the study of the herpetological diversity of the Qinghai-Tibet Plateau. The senior author became an Asian pit viper enthusiast and professional herpetological researcher under his instruction. The common name of
<taxonomicNameid="5B0BBC474138C26F5D92486044B55323"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<taxonomicNameid="0A408CA1F5E3C387287E2E0E144BBC0E"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<figureCitationid="15B9104B8BFB5BA3519D4B456C276F72"captionStart="Figure 1"captionStartId="F1"captionText="Figure 1. Gloydius lipipengi sp. nov. (A, B IVPP OV 2720, holotype) and Gloydius swild sp. nov. (C IVPP OV 2725, holotype, D IVPP OV, 2726, paratype) in life, not to scale."figureDoi="10.3897/zookeys.1061.70420.figure1"httpUri="https://binary.pensoft.net/fig/595356"pageId="0"pageNumber="87">1</figureCitation>
<taxonomicNameid="F2796A169CF6CCDE79EFD2024C027E70"authorityName="Hoge & Romano-Hoge"authorityYear="1981"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius"order="Squamata"pageId="0"pageNumber="87"rank="genus">
<taxonomicNameid="BC1900F42DA57876B2906F2773FAA3FA"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
sp. nov. differs from other congeneric species in the following characteristics: i) third supralabial scale not touching the orbit; ii) a pair of prominent black markings on the occiput; iii) black-bordered greyish cheek stripe extending from the posterior margin of orbit (not separated by the postoculars) to the ventral surface of the neck; iv) black irregular annular crossbands on the mid-body; iv) two rows of black blotches on the ventral side; v) 23-21-15 circum-body scales; vi) 165 ventral scales; and vii) 46 subcaudal scales.
<tableCitationid="8F902D73720DE9F662F560BFEFFDAA5C"captionStart="Table 5"captionStartId="T5"captionText="Table 5. Brief morphological comparisons between Gloydius lipipengi sp. nov., G. swild sp. nov. and other congeneric species."httpUri="http://table.plazi.org/id/BD83D965C796E96243DDBDE40DE25ED9"pageId="0"pageNumber="87"tableUuid="BD83D965C796E96243DDBDE40DE25ED9">5</tableCitation>
<taxonomicNameid="349FD049EB3A8392EB01BD23360E8ED2"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<tdid="CD9E15B04CFD29267275F7CD3F2BEB78"colspan="1"pageId="0"pageNumber="87"rowspan="1">one triangular spot between the third and fourth supralabial</td>
<tdid="46243E06B231884A42623B6A3DBCF3FB"colspan="1"pageId="0"pageNumber="87"rowspan="1">reddish brown or light grey</td>
<tdid="29ED94D457E616C582BAD03632D9FC4B"colspan="1"pageId="0"pageNumber="87"rowspan="1">brown, reddish brown or dark grey small blotches or large black bordered patches</td>
<tdid="EA85B5FFC5BA768C011CBB57E7602888"colspan="1"pageId="0"pageNumber="87"rowspan="1">khaki or yellowish brown</td>
<tdid="EC44481DB12771387C49213672B98716"colspan="1"pageId="0"pageNumber="87"rowspan="1">regular large (or discrete small) scarlet or brownish yellow patches or stripes</td>
<tdid="390593DE4E9C17C4D5639039893D4FFD"colspan="1"pageId="0"pageNumber="87"rowspan="1">greenish brown, yellowish brown or nut-brown</td>
<tdid="0894B17CE8C4C4EEBAC4831FA2DC8C9C"colspan="1"pageId="0"pageNumber="87"rowspan="1">patches, transverse crossbands or four longitudinal zigzag strips</td>
<taxonomicNameid="175E51BE2460ED8CCB7C1ABBD751F5F7"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<taxonomicNameid="A717FB65E206DEFDE98F00B7E1778BEF"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<bibRefCitationid="4C482189ACCF8AAB3E1DA9209B31F7BF"DOI="https://doi.org/10.1670/18-126"author="Wang, K"journalOrPublisher="Journal of Herpetology"pageId="0"pageNumber="87"pagination="224 - 236"refId="B22"refString="Wang, K, Ren, J, Dong, W, Jiang, K, Shi, J, Siler, CD, Che, J, 2019. A New Species of Plateau Pit Viper (Reptilia: Serpentes: Gloydius) from the Upper Lancang (= Mekong) Valley in the Hengduan Mountain Region, Tibet, China. Journal of Herpetology 53 (3): 224 - 236, DOI: https://doi.org/10.1670/18-126"title="A New Species of Plateau Pit Viper (Reptilia: Serpentes: Gloydius) from the Upper Lancang (= Mekong) Valley in the Hengduan Mountain Region, Tibet, China."url="https://doi.org/10.1670/18-126"volume="53"year="2019">Wang et al. 2019</bibRefCitation>
by its greyish brown body colour (versus yellowish-brown body colour) and lacking a lateral white line on each lateral side (versus possessing a lateral white line on each side).
<taxonomicNameid="1BA3AB796271E05FC51A3EFD615F4F13"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
IVPP OV 2720, adult male, a slender pit viper with a total length of 628.2 mm (SVL 540.6 mm and TL 87.6 mm), preserved in 75% ethanol with its left hemipenes partially extruded (Figs
<figureCitationid="6B77533F87E3448CBB78F9A176F0E3A4"captionStart="Figure 1"captionStartId="F1"captionText="Figure 1. Gloydius lipipengi sp. nov. (A, B IVPP OV 2720, holotype) and Gloydius swild sp. nov. (C IVPP OV 2725, holotype, D IVPP OV, 2726, paratype) in life, not to scale."figureDoi="10.3897/zookeys.1061.70420.figure1"httpUri="https://binary.pensoft.net/fig/595356"pageId="0"pageNumber="87">1</figureCitation>
<figureCitationid="87FB7020CC6E083FB436C3E5D3ACB9A0"captionStart="Figure 2"captionStartId="F2"captionText="Figure 2. Holotype of Gloydius lipipengi sp. nov. (IVPP OV 2720) in preservative A dorsal view B ventral view."figureDoi="10.3897/zookeys.1061.70420.figure2"httpUri="https://binary.pensoft.net/fig/595357"pageId="0"pageNumber="87">2</figureCitation>
<taxonomicNameid="0014F648D28EA9391D9FF3669CD6A9B4"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<taxonomicNameid="E29807DE8383899762AC89856D920857"authorityName="Shi & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius swild"order="Squamata"pageId="0"pageNumber="87"rank="species"species="swild">
<taxonomicNameid="AC99A275D3E1C116B59AC165D36CFDD6"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<paragraphid="892C35CCE21B1FE6E29510A2518D3401"pageId="0"pageNumber="87">The head is slender and triangular shaped in dorsal view, distinct from the neck. Canthus rostralis are not distinct. The head is 25.2 mm in length, 13.2 mm in width and 8.2 mm in depth.</paragraph>
Rostral scale slightly up-turned, visible from dorsal view; nasal divided, anterior part larger; seven supralabials on both sides: second smallest, not reaching the pit; third highest, not touching the bottom of orbit (separated by one small subocular); fourth longest, not touching the orbit; three preoculars, two postoculars, inferior one touching the top of the third supralabials, forming the bottom margin of the orbit; two rows of temporals (2+4); infralabials 10 on left side while 11 on right, first pair in contact behind the mental; second, third and fourth pairs meet on the chin shield; chin shield is rhomboidal in shape, the posterior chin shield comprises two pairs of scales, forming the mental groove (Fig.
<figureCitationid="DA2F59E12AB3F8C940C4EC87CB155F4B"captionStart="Figure 3"captionStartId="F3"captionText="Figure 3. Head squamation of Gloydius lipipengi sp. nov. (Holotype, IVPP OV 2720: A lateral view B dorsal view C ventral view) and G. swild sp. nov. (Holotype, IVPP OV 2725: D lateral view E dorsal view F ventral view). Scale bar: 10 mm."figureDoi="10.3897/zookeys.1061.70420.figure3"httpUri="https://binary.pensoft.net/fig/595358"pageId="0"pageNumber="87">3A-C</figureCitation>
<taxonomicNameid="B27DBA29FA23E9E0A4DB56DE953DC9C5"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<paragraphid="6DFA0BCF1EECD96A10BBE27AA8C384EE"pageId="0"pageNumber="87">Dorsal scales in 23-21-15 rows (reducing from 19 to 18 posteriorly at ventral 94-96), keeled except for the first scale row bordering the ventral scales; ventral scales 165 (excluding four preventral scales); anal plate single; subcaudals 46, in pairs</paragraph>
Eye dark brown on the upper half while black on the bottom half, pupil black, vertical with light yellow margins; postorbital stripe wide, greyish brown and black bordered on the lower edge, extending from the posterior orbit to the ventral surface of the neck; supralabials and infralabials greyish brown, scattered with very small irregularly sized black blotches. One black triangular mark on the anterodorsal head, covering the caudomedial part of prefrontals. One bold black M-shaped mark on the dorsomedial head, covering the caudal part of lateral frontals, the lateral part of parietals, merged with the postorbital stripe at the largest temporal scale (but not covering the upper postorbital). The upper postorbital white while the top part of the bottom postorbital is black (covered by the postorbital stripe).
The body coloration is dark greyish brown, with two rows of irregular black annular crossbands on the mid-body, each covering 20 or more scales, separated by a gap of two row scale vertically, extending laterally to one or two dorsal scales from the ventrals. Ventral scales light grey, with two large black blotches on each side, clustered into two ventral stripes. The tip of tail is similar to the main body in coloration (Figs
<figureCitationid="71588003BDD5AF23752FC68CEEC8452F"captionStart="Figure 1"captionStartId="F1"captionText="Figure 1. Gloydius lipipengi sp. nov. (A, B IVPP OV 2720, holotype) and Gloydius swild sp. nov. (C IVPP OV 2725, holotype, D IVPP OV, 2726, paratype) in life, not to scale."figureDoi="10.3897/zookeys.1061.70420.figure1"httpUri="https://binary.pensoft.net/fig/595356"pageId="0"pageNumber="87">1</figureCitation>
<figureCitationid="69E8282C20A7F18C9B23D688332EA97C"captionStart="Figure 2"captionStartId="F2"captionText="Figure 2. Holotype of Gloydius lipipengi sp. nov. (IVPP OV 2720) in preservative A dorsal view B ventral view."figureDoi="10.3897/zookeys.1061.70420.figure2"httpUri="https://binary.pensoft.net/fig/595357"pageId="0"pageNumber="87">2</figureCitation>
The premaxilla has bifurcated transverse process on each side. The anterior margin of the premaxilla is blunt. The dorsal tip of the ascending process of premaxilla is triangular in lateral view, not reaching the anterior tip of nasals. The horizontal laminae of the nasals are scutiform in dorsal view. The septomaxillae have prominent dorsolateral processes, nearly meeting the horizontal laminae of the nasals.
The parietal is roughly T-shaped in dorsal view. The anterolateral part of the parietal bulges prominently laterally while the dorsoposterior part tapers medially. The postorbital processes of the parietal are prominent. The frontals are squared. The lateral margin of frontals concaved obviously on each side, forming the dorsal edge of orbit. The prefrontal has an elongate blunt lateral process, posterolaterally pointed. The lacrimal foramen perforates the medial lamina of the prefrontal. The prefrontal-frontal join surface is waved in dorsal view.
<paragraphid="5BAA8749BDA3AA5662E81CC1EA05024A"pageId="0"pageNumber="87">The postorbital is relatively small and cashew-shaped, the top of the postorbital does not reach the posterolateral end of frontal. The basisphenoid is spearhead in shape, narrow anteriorly and expanded laterally. The supraoccipital is longitudinally compressed, occupies almost two thirds the total width of the otic region.</paragraph>
The fang is relatively short and curved, roughly the same length of the maxilla, one third the length of ectopterygoid, attached with seven or eight replacement fangs on each side. The palatine bears three teeth. The ectopterygoid is flat and widened at the anterior part. The pterygoid is slender, the dentigerous process of pterygoid is straight, bearing 12/11 teeth (left/right), occupies almost half the total width of the pterygoid, the posterior portion of pterygoid is medially expanded.
The supratemporal is slender, has a lateral process, anterolaterally pointed, lies in front of the supratemporal-quadrate articulation. The quadrate is straight, slender, and enlarged on both ends. The mandible is slender and moderately curved. The prearticular crest of the compound bone is prominent while the surangular crest is slightly concaved. The dental bone bears 11/12 teeth (left/right); the dentary teeth are perpendicular to the dentary bone, decreasing in size at the third tooth. The posterior tip of ventral process of dentary extends farther posteriorly than the dorsal process.
but differ by the possession of longer and stronger spines, seven or eight subcaudals in length, and forked for two subcaudals. Small and stubby spines range from the basal to the distal side of the organ, without any conspicuously enlarged spines (versus 3-5 enlarged spines on the base in the
<bibRefCitationid="CDBB2BDF23E16A7397DBB60F164CBC81"DOI="https://doi.org/10.5358/hsj1972.14.1_23"author="Gloyd, HK"journalOrPublisher="Society for the Study of Amphibians and Reptiles, USA"pageId="0"pageNumber="87"refId="B7"refString="Gloyd, HK, 1990. Snakes of the Agkistrodon Complex: A Monographic Review. Society for the Study of Amphibians and Reptiles, USA, DOI: https://doi.org/10.5358/hsj1972.14.1_23"title="Snakes of the Agkistrodon Complex: A Monographic Review."url="https://doi.org/10.5358/hsj1972.14.1_23"year="1990">Gloyd 1990</bibRefCitation>
<taxonomicNameid="674C3EFF259BA2996BDDC00060CFB590"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<figureCitationid="26049CED3F5C6FA2A08857D069684774"captionStart="Figure 6"captionStartId="F6"captionText="Figure 6. Type localities of Gloydius lipipengi sp. nov. (red triangles) and G. swild sp. nov. (black triangles), with the collection localities of some other congeneric species."figureDoi="10.3897/zookeys.1061.70420.figure6"httpUri="https://binary.pensoft.net/fig/596192"pageId="0"pageNumber="87">6</figureCitation>
<figureCitationid="3192040AD6B5509373819F3334BF78E7"captionStart="Figure 7"captionStartId="F7"captionText="Figure 7. The habitat of Gloydius lipipengi sp. nov. (A Muza Village, Zaty, Tibet, type locality of G. lipipengi sp. nov. B the landscape of the Nujiang River, 15 km from the type locality) and Gloydius swild sp. nov. (C Heishui, Sichuan) A and B Photographs by Jin-Cheng Liu."figureDoi="10.3897/zookeys.1061.70420.figure7"httpUri="https://binary.pensoft.net/fig/595362"pageId="0"pageNumber="87">7</figureCitation>
<taxonomicNameid="5D5D3D46E5B0A3AB5A00B1E207D435A2"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<taxonomicNameid="F71D9AA4D49B69A252706AE1188ED13A"authorityName="Shi, Liu & Malhotra"authorityYear="2021"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius lipipengi"order="Squamata"pageId="0"pageNumber="87"rank="species"species="lipipengi">
<taxonomicNameid="1B6B32BCB8CA07A338F12C81A071FBA9"authorityName="Hoge & Romano-Hoge"authorityYear="1981"family="Viperidae"genus="Gloydius"kingdom="Animalia"lsidName="Gloydius"order="Squamata"pageId="0"pageNumber="87"rank="genus">
, based on 12S, 16S, ND4, and cytb sequences, with the maximum likelihood bootstrap supports (left, regular) and Bayesian posterior probabilities (right, italic) displayed on the nodes (those <50% are displayed as
