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Research Article
A new mountain pitviper of the genus Ovophis Burger in Hoge & Romano-Hoge, 1981 (Serpentes, Viperidae) from Yunnan, China
expand article infoXian-Chun Qiu‡, Jin-Ze Wang‡, Zu-Yao Xia§, Zhong-Wen Jiang|, Yan Zeng, Nan Wang|, Pi-Peng Li‡, Jing-Song Shi#
‡ Shenyang Normal University, Shenyang, China
§ University of California, Davis, United States of America
| Beijing Forestry University, Beijing, China
¶ Institute of Zoology, Chinese Academy of Sciences, Beijing, China
# Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China

Corresponding author: Nan Wang ( wangnan761227@bjfu.edu.cn )

Corresponding author: Pi-Peng Li ( 104466606@qq.com )

Corresponding author: Jing-Song Shi ( shijingsong0827@163.com )

Academic editor: Minh Duc Le
© 2024 Xian-Chun Qiu, Jin-Ze Wang, Zu-Yao Xia, Zhong-Wen Jiang, Yan Zeng, Nan Wang, Pi-Peng Li, Jing-Song Shi.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Qiu X-C, Wang J-Z, Xia Z-Y, Jiang Z-W, Zeng Y, Wang N, Li P-P, Shi J-S (2024) A new mountain pitviper of the genus Ovophis Burger in Hoge & Romano-Hoge, 1981 (Serpentes, Viperidae) from Yunnan, China. ZooKeys 1203: 173-187. https://doi.org/10.3897/zookeys.1203.119218
ZooBank: urn:lsid:zoobank.org:pub:86CD1FAF-8A41-41D2-BF2E-B28EBC805789
Open Access

Abstract

Based on a molecular phylogenetic analysis and morphological comparison, a new species of mountain pitviper, Ovophis jenkinsi sp. nov., is described. The new species was collected in Yingjiang County, Yunnan Province, China. It can be distinguished from congeneric species by the following characters: (1) internasals in contact or separated by one small scale; (2) second supralabial entire and bordering the loreal pit; (3) dorsal scales in 23 (25)–21 (23, 25)–19 (17, 21) rows; (4) 134–142 ventrals; (5) 40–52 pairs of subcaudals; (6) third supralabial larger than fourth in all examined specimens of Ovophis jenkinsi sp. nov.; (7) deep orange-brown or dark brownish-grey markings on dorsal head surface; (8) background color of dorsal surface deep orange-brown or dark brownish-grey; (9) both sides of dorsum display dark brown trapezoidal patches; (10) scattered small white spots on dorsal surface of tail.

Key words

Morphology, Ovophis jenkinsi sp. nov., snake, taxonomy, Yingjiang County

Introduction

The subfamily Crotalinae (pitvipers) is the largest group of family Viperidae, with 294 species in 23 genera, and widely distributed in Asia and America (Speybroeck et al. 2016; Uetz et al. 2024). The mountain pitviper (Ovophis) is a group of medium-sized venomous snakes that are mainly distributed through eastern Asia, the southern Himalayas, and the northern Indochina Peninsula (Che et al. 2020). Within the genus, the distribution of O. monticola (Günther, 1864), O. makazayazaya (Takahashi, 1922), O. tonkinensis (Bourret, 1934), and O. zayuensis (Jiang, 1977) in China was supported and defined preliminarily by Malhotra et al. (2011) by molecular phylogenetic analyses (12S, 16S, cytb, and ND4) and comparative morphology. Subsequently, Zeng et al. (2023) revised the molecular phylogeny (cytb, ND4, BACH1, c-mos, NT3, and Rag1) with additional specimens. The result restricted the distribution of O. monticola to Zhangmu County (southern Xizang) in China, demarcated the populations that are distributed through Sichuan–Yunnan in the west to Taiwan in the east as O. makazayazaya, and introduced O. malhotrae Zeng et al., 2023 as a new species representing the southern Yunnan population with its holotype description presented in non-paginationed supporting documents. The distribution of O. malhotrae is currently recorded only in Jinping and Pingbian, Yunnan and Lao Cai, Vietnam. MtDNA phylogenetic inference of the genus Ovophis and partial species of the rest of family Viperidae indicated when “O.” okinavensis (Boulenger, 1892) is included, genus Ovophis is polyphyletic, while “O.” okinavensis sistering Trimeresurus gracilis Oshima, 1920 (Malhotra and Thorpe 2000, 2004; Shi et al. 2021). Although the epithet change of “O.” okinavensis has not yet been declared, this species is no longer included in genus Ovophis in some recent taxonomic studies (Malhotra et al. 2011; Zeng et al. 2023). Currently, therefore, the genus Ovophis includes six species: O. monticola, O. convictus (Stoliczka, 1870), O. makazayazaya, O. tonkinensis, O. zayuensis, and O. malhotrae (Malhotra et al. 2011; Zeng et al. 2023; Uetz et al. 2024).

In 2018 and 2023, five specimens of genus Ovophis were collected in Yingjiang County, Yunnan. With applied comparative morphology and molecular phylogenetic analysis, these specimens were revealed as distinct from the other Ovophis species. Thus, we described here this new population as a new species.

Materials and methods

Sampling

Five specimens (IOZ 002679, IOZ 002680, YJ201801, YJ201802, and YJ201803) were collected by Zhong-Wen Jiang and Xian-Chun Qiu in October 2018 and 2023 from Tongbiguan Township, Yingjiang County, Yunnan Province, China. After euthanasia, liver tissues of specimens IOZ 002679 and IOZ 002680 were extracted and preserved in 95% ethanol for molecular analyses. All specimens were fixed in 10% buffered formalin and then transferred to 75% ethanol for permanent preservation. The specimens IOZ 002679 and IOZ 002680 are deposited in the Institute of Zoology, Chinese Academy of Sciences (IOZ, Beijing, China). The specimens YJ201801, YJ201802, and YJ201803 are deposited in Beijing Forestry University (BFU, Beijing, China).

Morphometrics

Morphological descriptions are accorded to Zhao (2006). Abbreviations are accorded to Darko et al. (2022). A total of 24 morphological characters were examined, including 11 mensural characters and 13 scalation characters. Morphological measurements were taken with digital calipers (Guanglu 111N-101V, accuracy 0.03 mm, Guanglu Digital Instruments, Guilin) to the nearest 0.1 mm. Abbreviations are as follows: SVL snout–vent length (distance from tip of snout to posterior margin of cloacal plate); TAL tail length (distance from posterior margin of cloacal plate to tip of tail); TL total length (distance from tip of snout to tip of tail); HL head length (distance from tip of rostral to posterior end of jaw); HW head width (maximum width of head); HH head height (maximum height between dorsal and ventral surfaces of head); ED eye diameter (horizontal eye diameter); IOD interorbital distance (distance between the top margin of eyes); IN internarial distance (distance between nostrils); RH maximum rostral height; RW maximum rostral width; LOR loreal; PRO preoculars; PO postoculars; SBO suboculars; ATEM anterior temporals; PTMP posterior temporals; SL supralabials; IL infralabials; CS chin shields; DSR dorsal scale rows (counted at one head length behind head, at midbody, and one head length before vent); PRV preventral scales (elongated scales situtated beneath the head before the ventrals); VS ventral scales (elongated scales situtated beneath the body between neck and vent); SC subcaudal scales.

Other morphological characters of Ovophis species were obtained from Zhao et al. (1998), Li et al. (2010), Neang et al. (2011), Sharma et al. (2013), Che et al. (2020), Guo et al. (2021), Huang (2021), and Zeng et al. (2023).

Phylogenetic analyses

Four mtDNA sequences are specifically amplified in this study: 12S rRNA using primers 12SFPhe and 12SRVal (Knight and Mindell 1993); 16S rRNA using primers 16sFL and 16sRH (Palumbi et al. 1991); cytochrome b (cytb) using primers L14910 and H16064 (Burbrink et al. 2000); and NADH dehydrogenase subunit 4 (ND4) using primers ND4F and LEUR (Arévalo et al. 1994). The standard PCR protocol is performed in 20 μl of reactant with at least 20 ng of template DNA and 10 pmol of primers. The PCR conditions: initial denaturation for 3 min at 94 °C, followed by 35 cycles, denaturation at 94 °C for 30 s, 30 s of annealing at different temperatures (52 °C for 12S, 50 °C for 16S, 56 °C for ND4, and 48 °C for cytb), and then elongation at 72 °C for 60 s, then finalized with elongation step of 10 min at 72 °C. Sequencing was conducted by Beijing Tianyi Huiyuan Bio-tech Co., Ltd. Sequence data were uploaded to GenBank (Table 1).

Table 1.
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Samples and sequences used for phylogenetic analysis in this study.

Species Locality Voucher GenBank accession number
12S 16S cytb ND4
Ovophis jenkinsi sp. nov. Yingjiang, Yunnan, China IOZ 002679 PP574250 PP574252 PP171456 PP171459
O. jenkinsi sp. nov. Yingjiang, Yunnan, China IOZ 002680 PP574249 PP574251 PP171455 PP171458
O. monticola Gandaki, Nepal ZMB 70216 HQ325260 HQ325078 HQ325138 HQ325199
O. monticola Gandaki, Nepal ZMB 70218 HQ325253 HQ325071 HQ325131 HQ325192
O. convictus Cameron Highlands, Pahang, Malaysia AM B628 HQ325264 HQ325082 HQ325141 –
O. convictus Pulau Langkawi, Malaysia AM B629 HQ325265 HQ325083 HQ325142 –
O. convictus Cameron Highlands, Pahang, Malaysia AM B580 – – HQ325129 HQ325190
O. malhotrae Yunnan, China GP 2041 – – OP441841 OP441784
O. malhotrae Jinping, Yunnan, China GP 2053 – – OP441842 OP441785
O. malhotrae Lao Cai, Vietnam ROM 39381 HQ325283 HQ325102 HQ325160 HQ325218
O. zayuensis Bomi, Xizang, China GP 713 – – OP441890 OP441833
O. zayuensis Chayu, Xizang, China GP 1505 – – OP441892 OP441836
O. makazayazaya Huili, Sichuan, China GP 21 – – OP441856 OP441798
O. makazayazaya Luquan, Yunnan, China KIZ 02143 – – OP441860 OP441802
O. makazayazaya Weixi, Yunnan, China YPX 53011 – – OP441861 OP441803
O. tonkinensis Maoming, Guangdong, China GP 1665 – – OP441876 OP441818
O. tonkinensis Xuan Son, Phu Tho, Vietnam KIZ 011602 – – OP441880 OP441822
Vipera berus Jilin, China – – – MF945570 MF945570

Note: The missing data are marked as “–”.

Corresponding homologous sequences of Ovophis species were obtained from GenBank, and the sequences of Vipera berus (Linnaeus, 1758) were used as outgroup in the phylogenetic analysis (Zeng et al. 2023). DNA nucleotide sequences were aligned in MEGA 6 (Tamura et al. 2013) with Clustal W algorithm, default parameters (Thompson et al. 1997). PartitionFinder 2.1.1 (Lanfear et al. 2012) was used to test the best partitioning scheme. Pairwise sequence divergence (uncorrected p-distances) was calculated using MEGA 6.

Bayesian inference was performed using MrBayes 3.1.2 (Ronquist et al. 2011). All searches consisted of three heated chains and a single cold chain. Three independent iterations each comprising two runs of 100 million generations, sampled every 10,000 generations, and parameter estimates were plotted against generation. The first 25% of the samples were discarded as burn-in, resulting in a potential scale reduction factor (PSRF) of < 0.005. A maximum-likelihood analysis was run with RaxML (Silvestro and Michalak 2012), and a majority rule consensus tree was calculated with 1,000 bootstrap replicates.

Results

Molecular phylogeny

A total of 2,703 aligned base pairs were obtained including 441 bp from 12S, 465 bp from 16S, 1,110 bp from cytb, and 687 bp from ND4. With respect to the different evolutionary characters of each molecular marker, the dataset was split into five partitions by genes and codon positions as recommended by PartitionFinder 2.1.1 (Table 2). The topological structures of the maximum-likelihood (ML) and Bayesian-inference (BI) trees are generally consistent (Fig. 1). All Ovophis samples are divided into seven clades. The cladistic relationship within the group of samples from Yingjiang was resolved with strong support (1.00/100 for BI and ML). The Yingjiang group is sister to the O. monticola group with decent support (1.00/99 for BI and ML), and the group of “Yingjiang”+O. monticola clustered with O. convictus, and formed a larger clade sister to the O. malhotrae+O. zayuensis lineage.

Figure 1. 

Bayesian-inference tree of Ovophis species inferred from the combined fragments of 12S, 16S, cytb, and ND4. Bayesian posterior probabilities/bootstrap support values for the clades are shown adjacent to the node.

Table 2.
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Partitions and their molecular evolution models selected by PartitionFinder 2.1.1.

Partitions Locus Length (bp) Models
Partition 1 12S, cytb pos 1, ND4 pos 3 1,040 TVM+I+G
Partition 2 16S 465 GTR+I
Partition 3 ND4 pos 1,
cytb pos 2		 599 TRN+I
Partition 4 cytb pos 3 370 TIM+G
Partition 5 ND4 pos 2 229 TIM+G

TVM: transversional substitution model; GTR: General Time-Reversible model; TRN: Tamura-Nei; TIM: transitional substitution model; +I: proportion of invariable sites; +G: rate heterogeneity.

The uncorrected p-distance based on cytb gene between the specimens IOZ 002679, IOZ 002680 from Yingjiang, Yunnan and O. monticola is 6.2–6.5%, equivalent to those among other recognized species, such as O. malhotrae vs. O. zayuensis (6.0–6.7%) (Table 3). Thus, the molecular phylogeny supports the validity of Ovophis jenkinsi sp. nov.

Table 3.
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Uncorrected p-distance among the sequences based on the cytb gene fragments of Ovophis species in this study.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
1 Ovophis jenkinsi sp. nov. IOZ 002679
2 O. jenkinsi sp. nov. IOZ 002680 0.003
3 O. monticola ZMB 70216 0.065 0.062
4 O. monticola ZMB 70218 0.065 0.062 0.000
5 O. convictus B580 0.097 0.097 0.106 0.106
6 O. convictus B628 0.118 0.118 0.125 0.125 0.041
7 O. convictus B629 0.095 0.095 0.101 0.101 0.006 0.038
8 O. makazayazaya GP21 0.102 0.102 0.111 0.111 0.111 0.110 0.105
9 O. makazayazaya KIZ02143 0.098 0.098 0.115 0.115 0.117 0.115 0.111 0.007
10 O. makazayazaya YPX53011 0.100 0.104 0.128 0.128 0.113 0.099 0.107 0.038 0.037
11 O. malhotrae GP2041 0.102 0.102 0.108 0.108 0.092 0.094 0.086 0.081 0.083 0.084
12 O. malhotrae GP2053 0.102 0.102 0.108 0.108 0.092 0.094 0.086 0.081 0.083 0.084 0.000
13 O. malhotrae ROM 39381 0.110 0.110 0.104 0.104 0.095 0.097 0.090 0.079 0.084 0.086 0.007 0.007
14 O. tonkinensis GP1665 0.105 0.101 0.119 0.119 0.096 0.096 0.094 0.095 0.100 0.109 0.081 0.081 0.083
15 O. tonkinensis KIZ011602 0.107 0.103 0.120 0.120 0.098 0.102 0.099 0.098 0.103 0.113 0.084 0.084 0.086 0.011
16 O. zayuensis GP713 0.085 0.085 0.110 0.110 0.090 0.092 0.088 0.085 0.085 0.094 0.060 0.060 0.062 0.089 0.088
17 O. zayuensis GP1505 0.088 0.088 0.106 0.106 0.092 0.094 0.090 0.087 0.090 0.095 0.065 0.065 0.067 0.094 0.094 0.019

Taxonomic account

Viperidae Oppel, 1811

Ovophis jenkinsi sp. nov.

https://zoobank.org/45FF4F16-3F01-4ADC-8DA7-24E84E3B810D

Type material

Holotype. IOZ 002679, an adult male (Figs 2, 3) from Tongbiguan Township, Yingjiang County, Yunnan Province, China (24°36′33″N, 97°39′29″E; 1,343 m a.s.l.) (Fig. 4). It was collected near the road by Zhong-Wen Jiang and Xian-Chun Qiu.

Figure 2. 

Holotype of Ovophis jenkinsi sp. nov. (IOZ 002679) in life.

Figure 3. 

Head of the holotype of Ovophis jenkinsi sp. nov. (IOZ 002679) A lateral (right) view B lateral (left) view C dorsal view D ventral view.

Figure 4. 

Habitat of Ovophis jenkinsi sp. nov. at the type locality in Tongbiguan Township, Yingjiang County, Yunnan Province, China A microhabitat, photographed by Sheng-Chao Shi B microhabitat, photographed by Guo-Wei Mo C, D macrohabitats, photographed by Xiao-Jun Gu.

Paratype. IOZ 002680 and YJ201801, adult females from Tongbiguan Township, Yingjiang County, Yunnan Province, China (24°35′04″N, 97°41′13″E; 1,321 m a.s.l.) collected by Zhong-Wen Jiang and Xian-Chun Qiu in October 2023 and 2018; juveniles YJ201802 and YJ201803 from the same locality collected by Zhong-Wen Jiang in October 2018.

Etymology

The specific epithet of the new species is dedicated to Robert “Hank” William Garfield Jenkins AM (August 1947–September 2023), a herpetologist and former chairman of the CITES Animals Committee from Australia, with a passion for snakes, especially pitvipers, and helped China, along with many Asian countries, complete snake census, conservation, and management projects. We suggest the common name “Jenkins’ mountain pitviper” in English and “yíng jiāng lào tiě tóu shé” (盈江烙铁头蛇) in Chinese.

Diagnosis

Ovophis jenkinsi sp. nov. can be distinguished by the following combination of morphological characters: (1) internasals in contact or separated by one small scale; (2) second supralabial entire and bordering the loreal pit; (3) dorsal scales in 23 (25)–21 (23, 25)–19 (17, 21) rows; (4) 134–142 ventrals; (5) 40–52 pairs of subcaudals; (6) third supralabial larger than fourth in all examined specimens of Ovophis jenkinsi sp. nov.; (7) deep orange-brown or dark brownish-grey markings on dorsal head surface; (8) background color of dorsal surface deep orange-brown or dark brownish-grey; (9) both sides of dorsum display dark brown trapezoidal patches; (10) scattered small white spots on dorsal surface of tail.

Description of holotype

Adult male; body stout and robust, medium-sized, tail slender, TL 515.9 mm (SVL 421.0 mm, TAL 94.9 mm, TAL/TL: 0.23); head triangular in dorsal view, moderately distinct from neck, longer than width, HL 26.6 mm, HW 18.6 mm (HW/HL: 0.70). Snout blunt and rounded, rostral trapezoidal, broader than high, RW 4.6 mm, RH 3.5 mm (RW/RH: 1.31; RW/HW: 0.25), upper edge visible from dorsum; eye small, ED 2.7 mm (ED/HL 0.10), pupil vertical; nostril subcircular, located in the middle of nasal; nasal divided into two scales by nostril; two internasals, elliptical, separated anteriorly by a small scale and bordered by the upper edge of rostral, connected posteriorly; loreal single; two preoculars, in contact with eye posteriorly; two postoculars, upper one in contact with the lower edge of supraocular; subocular single and elongate, respectively separated by two small scales from the third, fourth and fifth supralabials; supraocular single, the largest scales on the dorsal surface of head, separated by 7–8 scales; supralabials eight, first and second in contact with nasal, second entire and bordering the loreal pit, third larger than fourth; 11 infralabials on left (seventh and eighth infralabials bipartitioned relative to right), 10 infralabials on right, first pair in broad contact with each other, first to third in contact with chin shields; mental triangular; one pair of chin shields, meeting in midline, the right one slightly larger than the left; dorsal scales in 25–21–19 rows, bluntly keeled, except outer row; 134 ventrals, excluding six preventrals; subcaudal scales 49, paired, excluding tail tip; cloacal plate entire.

Coloration in life

Dorsal head surface black, with deep-orange blotches; a deep orange marking resembling an open pair of surgical scissors exists on the front of neck; a deep-orange stripe exists from the upper postocular to the anterior nape, the stripe demarcated from black dorsal head at top, gradually transitioning to black at bottom, approximately one scale row in width behind orbit of eye, after three scales, approximately two scales rows in width, enlarge to 3–4 scales rows in width on the posterior of head. Lateral head surface black, tiny white and vermilion spots exist on the surface of scales near snout; an irregular stripe extends from subocular to the fifth and sixth infralabials, the outermost ring of vermilion, subtle, second ring of white, obvious; the stripe splits in two at fifth and sixth supralabials, one extending backward through seventh, eighth supralabials and the last two infralabials, the other extending downward through seventh and eighth infralabials (left) and seventh infralabial (right), converging at the outer row of dorsal scales; similar markings exist on the third supralabial and third to fifth infralabials. Background color of ventral head surface deep orange, mixed with irregular white blotches with vermilion edges. Pupil black; iris deep orange mixed with white and black.

Background color of dorsal surface deep orange, with 18 connected or disconnected dark brown patches on both sides of body and three similar spots on anterior section of tail visible from dorsum; dorsal blotches predominantly trapezoidal, approximately 2–6 scales in length, and 4–5 scales rows in width, mottled with a few deep orange tiny spots on most dark brown patches; two clusters of lateral dark brown patches exist under each dorsal dark brown patch, each patch covers 2–3 dorsal scales and separated from ventral scales by 2–3 rows of dorsolateral scales. Posterior section of tail pink, 21 tiny spots exist on the dorsal surface, spots white with brown edges, no more than a scale in size. Mixed cream and tan on ventral surface of anterior tail and body, clean pink on posterior section of tail.

Intraspecific variation

Morphometric data are summarized in Table 4. Dorsal head surface of each paratype specimen has different approximately symmetrical markings respectively. Internasals are separated by one scale, and the dorsal background color is dark brownish-grey in paratypes IOZ 002680 and YJ201801. Light greyish-brown on background color of dorsal body in juveniles YJ201802 and YJ201803. Third to tenth subcaudals unpaired in YJ201803. The patches on dorsal body are mostly rectangular in IOZ 002680.

Table 4.
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Scalation data and measurements (in mm) of Ovophis jenkinsi sp. nov.

IOZ 002679 Holotype IOZ 002680 Paratype YJ201801 Paratype YJ201802 Paratype YJ201803 Paratype
Sex Male Female Female Juvenile Juvenile
TL 515.9 402.3 690.0 261.0 279.0
TAL 94.9 66.9 91.1 43.8 45.0
HL 26.6 26.0 38.3 16.2 16.8
HW 18.6 20.6 28.8 11.7 11.2
HW/HL 0.70 0.79 0.75 0.72 0.67
PRO 2 2 2 2 2
SBO 1 3 2 2 2
PO 2 3 2 2 3
SL 8/8 9/9 8/10 8/– 8/8
IL 11/10 12/12 11/11 11/11 11/11
DSR 25–21–19 25–23–19 25–25–21 23–21–19 23–23–17
VS 134 142 138 134 135
SC 49 40 40 50 52

Note: the missing data are marked as “–”.

Comparisons

Ovophis jenkinsi sp. nov. can be distinct from other congeneric species by the following characters (Table 5): internasals in contact or separated by one small scale (vs internasals separated by two small scales in O. malhotrae); second supralabial entire and bordering the loreal pit (vs second supralabial bordering the loreal pit or separated by a loreal in O. makazayazaya, O. tonkinensis, and O. zayuensis); dorsal scales in 23 (25)–21 (23, 25)–19 (17, 21) rows (vs dorsal scales 23 (21)–23 (21)–19 in O. monticola, 25 (27, 29)–23 (21–25)–19 (21) in O. tonkinensis, 27–23–19 in O. malhotrae and 25 (27)–23–19 (17) in O. zayuensis); 134–142 ventrals (vs 141–172 ventrals in O. monticola, 145 in O. malhotrae and 160–177 in O. zayuensis); subcaudal scales in pairs (vs unpaired in O. tonkinensis and O. zayuensis); 40–52 pairs of subcaudals (vs 17–31 pairs in O. convictus); the third supralabial being larger than fourth (vs fourth larger than third supralabial in O. makazayazaya and O. tonkinensis); deep orange-brown or dark brownish-grey markings on dorsal head surface (vs no markings on dorsal head surface in O. convictus and O. tonkinensis); background color of dorsal surface deep orange-brown or dark brownish-grey (vs yellowish-brown or dark-grey in O. makazayazaya, yellowish-brown in O. monticola and O. tonkinensis, reddish-brown or brown in O. zayuensis); both sides of dorsum display dark brown trapezoidal patches (vs mostly rectangular patches in O. monticola); adults HW/HL 0.70–0.79 (vs 0.64–0.65 in O. monticola); scattered small white spots on dorsal surface of tail (vs continuous small white spots on dorsal surface of tail in O. tonkinensis and O. malhotrae).

Table 5.
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Morphological comparison of Ovophis species.

DSR VS SC Does the 2nd SL border the loreal pit 3rd and 4th SL Dosal head surface Dorsal background color Dorsal patches White spots on dorsal surface of tail
Ovophis jenkinsi sp. nov. 23 (25)–21 (23, 25)–19 (17, 21) 134–142 40–52, paired Yes 3rd > 4th Patterned Deep orange-brown or dark brownish-grey Mostly trapezoidal Scattered
O. monticola 23 (21)–23 (21)–19 141–172 37–58, paired Yes 3rd > 4th Patterned Yellowish-brown Mostly rectangular Scattered
O. convictus 25–25–18 136 17–31, paired Yes 3rd > 4th Unpatterned Yellowish-brown Mostly rectangular Scattered
O. makazayazaya 25 (27, 29)–23 (25, 21)–19 (21) 131–159 34–52, paired Yes or no 3rd < 4th Patterned or unpatterned Yellowish-brown or dark-grey Rectangular or irregular patches Scattered
O. malhotrae 27–23–19 145 47, paired Yes 3rd > 4th Patterned Dark-brown Mostly rectangular Continuous
O. tonkinensis 25 (27, 29)–23 (21–25)–19 (21) 128–134 39–49, unpaired Yes or no 3rd < 4th Unpatterned Yellowish-brown Rectangular or irregular patches Continuous
O. zayuensis 25 (27)–23–19 (17) 160–177 43–64, unpaired Yes or no 3rd > 4th Patterned or unpatterned Reddish-brown or brown Mostly trapezoidal and triangular No visible white spots

Distribution and ecology

Ovophis jenkinsi sp. nov. is currently known only from Yingjiang County, Yunnan Province, China. It was found in the tropical montane rainforest at an altitude of around 1,300 m. Overlapping herpetofauna includes Lycodon chapaensis (Angel & Bourret, 1933), Trimeresurus popeiorum Smith, 1937, Pseudocalotes jingpo Xu et al., 2024, and other species (Xu et al. 2024). The new species reaches activity peak in autumn and is active nocturnally during light rain or high humidity, at temperatures around 15–22 °C (Fig. 5). The type specimens were collected at night in October. When threatened, these snakes inflate their bodies to make themselves appear larger and strike quickly. The specimen IOZ 002680 had released odour from the cloacal scent glands when captured. We are currently unsure of the feeding habit of O. jenkinsi sp. nov. in the wild. They fed on juvenile mice (Mus musculus Linnaeus, 1758) in our captivity observations. Therefore, we presume the species prey on small mammals in the wild.

Figure 5. 

Ovophis jenkinsi sp. nov. and its microhabitat. Photographed by Zhong-Wen Jiang in Yingjiang, Yunnan.

Discussion

The phylogenetic topology in this study supports Zeng et al. (2023): Ovophis malhotrae sistering O. zayuensis (Zeng et al. 2023). However, the phylogenetic topology in this study differs from previous publications (Malhotra et al. 2011; Zeng et al. 2023). In this study, O. tonkinensis clustered with all other congeners (excluding “O.” okinavensis), while it clustered with O. makazayazaya in Malhotra et al. (2011) and Zeng et al. (2023). Thus, introducing a larger quantity of genetic dataset is suggested when conducting further phylogenetic studies of genus Ovophis.

In recent years, new snake species have been discovered constantly near the Yunnan border (Jiang et al. 2020; Chen et al. 2021; Hou et al. 2021; Lee 2021; Liu et al. 2021; Shi et al. 2022; Ma et al. 2023). Zeng et al. (2023) described the new species O. malhotrae based on specimens from southern Yunnan, and its molecular systematic position indicated that several populations from Vietnam and Laos may refer to cryptic species of the genus Ovophis. Thus, snake diversity in this region may have been underestimated in previous studies.

According to field surveys and recent publications, we identified updated distribution sites of Ovophis species in China (Zhang et al. 2011; Che et al. 2020; Zeng et al. 2023; Uetz et al. 2024) (Fig. 6). In Yunnan, O. makazayazaya is widely distributed in most parts of the province, O. zayuensis in the Gaoligong Mountain region of western Yunnan, and O. malhotrae in Jinping and Pingbian in southeastern Yunnan (Wang et al. 2022; Zeng et al. 2023). In some areas, there may be overlapping distributions of O. makazayazaya and other congeners. The specimens of “Trimersurus monticola” from Hotha, Longchuan County, Yunnan Province, cited by Anderson (1878a), displayed intra-specific polymorphism: the second supralabial completed or divided from the anterior of loreal pit, consistent with the character of O. makazayazaya (Guo et al. 2021). Further examinations through photos of the specimen that was recorded by Anderson (1878b) were conducted and showed that it displays 10 supralabials, the second divided from the loreal pit and fourth larger than third, suggesting taxonomic placement under O. makazayazaya. However, since Longchuan County is adjacent to Yingjiang County, and in specimens from Hotha, Longchuan, the second supralabial also reaches the loreal pit, they should belong to O. jenkinsi sp. nov. In addition, Yingjiang County, Yunnan Province is located on the border of China and is adjacent to Myanmar; thus, the new species may also be distributed in Myanmar.

Figure 6. 

The type locality of Ovophis jenkinsi sp. nov. and some distribution sites of Ovophis species in China.

The new species is morphologically most similar to O. monticola, but can be distinguished by morphological characters such as wider head, fewer ventrals, trapezoidal patches on dorsal body, and deep orange-brown or dark brownish-grey dorsal surface rather than yellowish brown. In the specimens examined, the new species has a maximum TL of 690 mm (specimen YJ201801), while O. monticola appears to be larger, with a maximum TL of 1,300 mm (Sharma et al. 2013). We will collect more specimens of O. jenkinsi sp. nov. in the future to supplement the morphological data. Molecular phylogenetic analyses show O. jenkinsi sp. nov. is genetically differentiated from O. malhotrae (p-distance 10.2–11.0). Currently, only the holotype of O. malhotrae is described in supporting documents, the intraspecific variation of this species is unclear. It is, therefore, suggested that additional samples from along the Yunnan border and adjacent areas would enrich the current morphological dataset of O. jenkinsi sp. nov., O. malhotrae, and other Ovophis species and support further biodiversity discoveries.

Acknowledgements

We thank Shuo Qi for helping in the research method. We thank Sheng-Chao Shi, Guo-Wei Mo, and Xiao-Jun Gu for providing habitat photos for this study. We thank Dehong Forestry and Grassland Administration and Tongbiguan Natural Reserve for their support in specimen collection.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by National Natural Science Foundation of China (NSFC 42202014, Jingsong Shi); The National Key R&D Program of China (nos. 2022YFC2601200, 2023YFC2604904, Ming Bai); the National Science & Technology Fundamental Resources Investigation Program of China (nos. 2022FY100500, 2023FY100301, Ming Bai); the Second Tibetan Plateau Scientific Expedition and Research Program (2019QZKK0705, Tao Deng and Xi-Jun Ni).

Author contributions

Xian-Chun Qiu: Writing original draft. Jin-Ze Wang: Participating in field surveys; writing original draft. Zu-Yao Xia: Searched for references needed for this thesis; reviewed and revised this thesis. Zhong-Wen Jiang: Participating in the field survey. Yan Zeng: production of distribution maps and the literature search. Nan Wang: Reviewed and revised this thesis. Pi-Peng Li: Supervised the writing of this thesis; reviewed and revised this thesis. Jing-Song Shi: Methodology, funding and review.

Author ORCIDs

Xian-Chun Qiu https://orcid.org/0009-0001-5776-711X

Jing-Song Shi https://orcid.org/0000-0001-9168-1734

Data availability

All of the data that support the findings of this study are available in the main text.

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