Abstract
Lycodongammiei (Blanford, 1878), a rarely encountered species of Asian snake, is characterized by ambiguous systematics and biology. Based on a sole specimen of L.gammiei rediscovered in southeastern Xizang, China, we conduct a detailed morphological examination and description, and investigate the systematic position of this species. Morphologically, the newly collected specimen is closely aligned with specimens previously described. Mitochondrial DNA-based phylogenetic analyses reveal that L.gammiei constitutes an independent evolutionary lineage, forming a clade with L.fasciatus (Anderson, 1879), L.gongshan Vogel & Luo, 2011, L.butleri Boulenger, 1900, and L.cavernicolus Grismer, Quah, Anuar, Muin, Wood & Nor, 2014. The closest genetic distance between L.gammiei and its congeners was 10.2%. The discovery of L.gammiei in Medog County, China, signifies an eastward expansion of its known geographical distribution.
Key words: Himalayas, phylogeny, Qinghai-Xizang Plateau, snake
Introduction
Exploring the boundaries of geographic distribution and systematic position of species is crucial for understanding their evolutionary origins and diversification and for devising appropriate conservation strategies. Despite considerable progress in recent years, many species remain poorly known and explored. This is particularly evident for some snake species due to their rarity and cryptic habitats.
Lycodongammiei (Blanford, 1878), a rare non-venomous snake species within the family Colubridae, was initially described as Ophitesgammiei based on a single specimen collected from Darjeeling, West Bengal, India (Blanford 1878). Subsequently, it was reclassified into the genus Lycodon (Boulenger 1890) or Dinodon (Wall 1923; Smith 1943), identifying it as Lycodongammiei. Wall (1911) compared L.gammiei and L.fasciatus (Anderson, 1879), and he synonymized L.fasciatus with L.gammiei. However, Wall (1923) later revised this view, recognizing its distinctiveness and validity of L.fasciatus. Mahendra (1984) proposed that L.gammiei was a color variety of L.septentrionalis (Gunther, 1875), while this synonymy was not accepted by all authors. Since its initial description, L.gammiei has been found in southeastern Xizang, China (Agarwal et al. 2010) and in Bhutan (Wangyal 2013). To date, however, few specimens of the species have been collected, and no genetic data have been reported.
In 2023, we collected a living specimen of L.gammiei in Medog County, southeastern Xizang, China. The rediscovery of this species in Xizang not only extends this species’ geographic distribution but also allows the exploration of its systematic position through molecular data.
Materials and methods
Morphological examination
The specimen deposited at Yibin University (YBU 230088) was collected in Beibeng Town, Medog County, southeastern Xizang, China (29°14′02″N, 95°10′38″E) (Fig. 1) on 14 August 2023 at an elevation of 1,431 m by Xiaoqi Mi. The snake was found on a tree near a road at 23:30 hours. Characters relating to scalation, color pattern, and body proportions were recorded from the preserved specimen in laboratory. Snout–vent length (SVL) and tail length (TL) were measured using a meter ruler to the nearest 0.5 centimeter, while all remaining measurements were taken using digital calipers to the nearest millimeter. Symmetric mensural head characters were taken on the right side unless unavailable (e.g. damaged), while meristic characters were recorded on both sides and reported in left/right order.
Figure 1.
Map showing currently known localities of Lycodongammiei.
Comparative data of other specimens of this species were taken from the literature (Blanford 1878; Mistry et al. 2007; Chettri and Bhupathy 2009; Wangyal 2013).
Molecular phylogeny
Genomic DNA was extracted from the liver tissue of the newly collected specimen using an Animal Genomic DNA Purification Kit (TIANGEN Bio-tech Co., Ltd, Beijing, China). Subsequently, a fragment of the mitochondrial gene cytochrome b (cyt b) was amplified using primers H14919 (5′-AACCACCGTTGTTATTCAACT-3′) and L16064 (5′-CTTTGGTTTACAAGAACAATGCTTTA-3′) (Burbrink et al. 2000). The polymerase chain reaction (PCR) products were purified and sequenced in both directions by Sangon Biotech Co., Ltd (Chengdu, China). The obtained sequences were manually edited using SeqMan in Lasergene v. 7.1 (DNASTAR, USA), and aligned using the ClustalW algorithm with default parameters in MEGA v. 7.0 (Kumar et al. 2016), followed by a visual inspection for minor manual adjustments. The DNA sequences were translated into amino acid sequences using MEGA v. 7.0 to verify sequence quality and detect any unexpected stop codons (Kumar et al. 2016). Furthermore, 80 additional sequences were downloaded from GenBank (Table 1).
Table 1.
Detail information for the samples used in this study.
| No. | Species | Voucher Number | Locality | GenBank No. |
|---|---|---|---|---|
| 1 | Lycodonalbofuscus | LSUHC 3867 | – | KX660500 |
| 2 | Lycodonalbofuscus | LSUHC 4588 | – | KX660501 |
| 3 | Lycodonalcalai | KU 327847 | Barangay San Antonio, Batanes Province, Philippines | KC010344 |
| 4 | Lycodonalcalai | KU 327848 | Municipality of Sabtang, Batanes, Philippines | KC010345 |
| 5 | Lycodonanakradaya | SIEZC 20247 | Song Giang River, Khanh Hoa Province, Vietnam | OM674283 |
| 6 | Lycodonanakradaya | SIEZC 20248 | Song Giang River, Khanh Hoa Province, Vietnam | OM674284 |
| 7 | Lycodonaulicus | KU 315378 | Tablas Island, Romblon Province, Philippines | KC010350 |
| 8 | Lycodonaulicus | PNM 7705 | Leyte Island, Leyte province, Philippines | KC010349 |
| 9 | Lycodonbanksi | VNUF R2015.20 | Khammouane, Laos | MH669272 |
| 10 | Lycodonbibonius | KU 304589 | Cagayan, Philippines | KC010351 |
| 11 | Lycodonbutleri | LSUHC 8365 | Perak, Malaysia | KJ607892 |
| 12 | Lycodonbutleri | LSUHC 9137 | Perak, Malaysia | KJ607891 |
| 13 | Lycodoncapucinus | – | – | MK844525 |
| 14 | Lycodoncapucinus | MVZ 291703 | Timor | MK844522 |
| 15 | Lycodoncapucinus | MVZ 291704 | Timor | MK844523 |
| 16 | Lycodoncathaya | SYS r001542 | Longsheng County, Guangxi, China | MT602075 |
| 17 | Lycodoncathaya | SYS r001630 | Longsheng County, Guangxi, China | MT602076 |
| 18 | Lycodoncavernicolus | LSUHC 10500 | Perlis, Malaysia | KJ607890 |
| 19 | Lycodoncavernicolus | LSUHC 9985 | Perlis, Malaysia | KJ607889 |
| 20 | Lycodoncf.flavozonatus | KIZ 032400 | Zayu, Xizang, China | MW199792 |
| 21 | Lycodonchapaensis | KIZ 27593 | Tengchong, Yunnan, China | MW353741 |
| 22 | Lycodonchapaensis | KIZ 35013 | Lushui, Yunnan, China | MW353742 |
| 23 | Lycodonchrysoprateros | KU 307720 | Cagayan, Philippines | KC010360 |
| 24 | Lycodondeccanensis | – | Tumkur District, Karnataka, India | MW006487 |
| 25 | Lycodondeccanensis | NCBS NRC AA0010 | Karnataka, India | MW006486 |
| 26 | Lycodondumerilii | KU 305168 | Dinagat Island, Philippines | KC010362 |
| 27 | Lycodondumerilii | KU 319989 | Mindanao Island, Agusan del Sur Province, Philippines | KC010361 |
| 28 | Lycodondumerilii | PNM 7751 | Leyte Island, Leyte Province, Philippines | KC010363 |
| 29 | Lycodoneffraenis | KU 328526 | Karome, Nakhon Si Thammarat, Thailand | KC010364 |
| 30 | Lycodoneffraenis | LSUHC 9670 | Kedah, West Malaysia | KC010376 |
| 31 | Lycodonfasciatus | CHS 837 | Yunnan, China | MK201559 |
| 32 | Lycodonfasciatus | KIZ 46120 | Himalayan region in China | MW111468 |
| 33 | Lycodonflavicollis | – | Devarayanadurga, Karnataka, India | MW006488 |
| 34 | Lycodonflavicollis | AIWC 081 | India | MZ029434 |
| 35 | Lycodonflavozonatus | KIZ 023279 | Xizang, China | MW199789 |
| 36 | Lycodonflavozonatus | KIZ 07067 | Xizang, China | MW199790 |
| 37 | Lycodonfutsingensis | CHS 670 | Nankunshan, Guangdong, China | MK201463 |
| 38 | Lycodonfutsingensis | CHS 751 | Guangdong, China | MK201504 |
| 39 | Lycodongammiei | YBU 230088 | Medog, Xizang, China | OR842906 |
| 40 | Lycodongongshan | GP 3547 | Lingcang,Yunnan, China | KP901025 |
| 41 | Lycodongongshan | GP 3548 | Lingcang,Yunnan, China | KP901026 |
| 42 | Lycodonjara | CAS 235387 | Putao, Kachin, Myanmar | KC010367 |
| 43 | Lycodonlaoensis | FMNH 258659 | Salavan, Laos | KC010368 |
| 44 | Lycodonlaoensis | LSUHC 8481 | O’Lakmeas, Pursat Province, Cambodia | KC010370 |
| 45 | Lycodonliuchengchaoi | CHS 158 | Sanjiazhai, Yunnan, China | MK201303 |
| 46 | Lycodonliuchengchaoi | CHS 843 | Ningshan, Shaanxi, China | MK201563 |
| 47 | Lycodonliuchengchaoi | CHS 873 | Shennongjia, Hubei, China | MK201580 |
| 48 | Lycodonmackinnoni | ADR 197 | Dhobighat, BWLS, Mussoorie, Uttarakhand | MW862977 |
| 49 | Lycodonmeridionalis | CHS 870 | Hechi, Guangxi, China | MK201578 |
| 50 | Lycodonmeridionalis | VNUF R2012.4 | Bac Kan, Vietnam | MH669271 |
| 51 | Lycodonmeridionalis | VNUF R2017.123 | Thanh Hoa Province, Vietnam | MH669270 |
| 52 | Lycodonmuelleri | DLSUD 031 | Luzon Island, Cavite Province, Philippines | KC010373 |
| 53 | Lycodonmuelleri | KU 313891 | Luzon Island, Camarines Norte Province, Philippines | KC010375 |
| 54 | Lycodonmuelleri | KU 323384 | Luzon Island, Aurora Province, Philippines | KC010374 |
| 55 | Lycodonnamdongensis | VNUF R2017.23 | ThanhHoa, Vietnam | MK585007 |
| 56 | Lycodonobvelatus | KIZ 040146 | Panzhihua, Sichuan, China | MW353745 |
| 57 | Lycodonpictus | CIB 115609 | Longzhou, Guangxi, China | MT845095 |
| 58 | Lycodonpictus | VNMN 011227 | Ha Lang, Cao Bang, Vietnam, | MT845094 |
| 59 | Lycodonrosozonatus | CHS 794 | Jianfengling, Hainan, China | MK201531 |
| 60 | Lycodonrufozonatus | CHS 601 | Huangshan, Anhui, China | MK201427 |
| 61 | Lycodonrufozonatus | CHS 710 | Yingpanxu, Hunan, China | MK201482 |
| 62 | Lycodonruhstrati | CHS 776 | Guangxi, China | MK201521 |
| 63 | Lycodonruhstrati | CHS 803 | Huaping, Guangxi, China | MK201538 |
| 64 | Lycodonsemicarinatus | KUZJPN 28044 | – | LC640371 |
| 65 | Lycodonseptentrionalis | CHS 162 | Yunnan, China | MK201305 |
| 66 | Lycodonseptentrionalis | KIZ 46117 | Xizang, China | MW199801 |
| 67 | Lycodonserratus | KIZ 038335 | Deqin, Yunnan, China | MW353746 |
| 68 | Lycodonstormi | JAM 7487 | Air Terjun Moramo, Sulawesi, Indonesia | KC010380 |
| 69 | Lycodonstriatus | – | Savandurga, Karnataka, India | MW006489 |
| 70 | Lycodonstriatus | CUHC 10368 | Pakistan | OQ282988 |
| 71 | Lycodonstriatus | CUHC 11257 | – | OQ282989 |
| 72 | Lycodonstriatus | CUHC 9457 | – | OQ282987 |
| 73 | Lycodonsubcinctus | CHS 734 | Guangdong, China | MK201493 |
| 74 | Lycodonsubcinctus | CHS 797 | Diaoluoshan Mountain, Hainan, China | MK201534 |
| 75 | Lycodonsynaptor | GP 3515 | Lingcang, Yunnan, China | KP901021 |
| 76 | Lycodonsynaptor | KIZ 046953 | Xizang, China | MW199805 |
| 77 | Lycodontruongi | SIEZC 20249 | Song Giang River, Khanh Hoa Province, Vietnam | OM674282 |
| 78 | Lycodonzawi | CAS 210323 | Thabakesay, Saging, Myanmar | AF471040 |
| 79 | Lycodonzawi | CAS 239944 | Kyaukpyu, RakhineState, Myanmar | KC010386 |
| 80 | Lycodonzayuensis | GP 7327 | Zayu, Xizang, China | OP434398 |
| 81 | Lycodonzayuensis | GP 7329 | Zayu, Xizang, China | OP434399 |
Both Bayesian-inference (BI) and maximum-likelihood (ML) analyses were executed for the final dataset. Prior to analyses, the best-fit model of nucleotide substitution was selected for each partition (codon position) using Akaike Information Criterion (AIC) implemented in PartitionFinder (Lanfear et al. 2012). The BI analyses were conducted using MrBayes v. 3.2.2 (Ronquist et al. 2012). Searches consisted of three independent runs, each involving four Markov chains (three heated chains and one cold chain), with 10 million generations, sampling every 2,000 generations and with 25% of initial samples discarded as burn-in. Convergence was determined via effective sample size (ESS > 200) and likelihood plots against time using Tracer v. 1.7 (Rambaut et al. 2018). The resulting trees were combined to determine the posterior probabilities (PP) for each node based on a 50% majority-rule consensus tree. The ML trees were constructed in IQ-tree (Lam-Tung et al. 2015) using the GTRCAT model and the same partitioning scheme. In total, 1,000 Ultrafast bootstraps (UFB) topological replicates were performed for branch support assessment. Boigacynodon (Boie, 1827) was selected as the outgroup following previous research (Guo et al. 2013).
Uncorrected genetic distance (p-distance) was calculated in MEGA v. 7.0 (Kumar et al. 2016).
Results
Morphological description
Female, SVL 698 mm and TL 223 mm. Body elongated; head rather flattened; snout blunt. Rostral large, trapezoid; internasals much broader than long; prefrontals 3.0 mm in length, distinctly wider than long, extending beyond both sides and touching preocular and loreal; frontal peltate, 4.6 mm in length and 4.1 mm in width; parietals subrectangular, 7.9 mm in length and 4.2 mm in width. Nasals large, nostril located anteriorly and opening backward; loreal scale 1, long, nearly rectangular, failing to touch eye; preocular 1, postoculars 2; temporals 2+2+3. Supralabials 8, 1st small, 3rd, 4th, and 5th entering orbit, 6th highest, 7th largest; infralabials 10, first pair in contact, 1st to 5th in contact with anterior chin shields. Chin shield pairs 2, elongate, anterior pair slightly larger than latter pair. Dorsal scales 17-17-15 rows, scales weakly keeled, except for outermost several rows; scales reduced from 17 to 15 at 143rd ventral position. Ventrals 228 (+ 1 preventral); cloacal plate entire; subcaudals 106, paired, dorsal scales of the tail reduced from 6 to 4 at 16th subcaudal position.
Head black, with yellow spots or short lines on some shields. Large, yellow spots on each side of posterior part of head. Conspicuous yellow collar on neck. Supralabials and anterior infralabials light yellow with dusky margins. Body surrounded by alternating dusky and light-yellow rings with very irregular, crooked margins. Yellow rings on body totaling 43, first pale ring clear above, anterior dark patch not continuous across throat, remaining rings encircling body. Lower part of head and neck light yellow. On belly, across anterior part of body, dark rings only about half as broad as light-yellow rings, less difference above, dark rings near head much broader above than white rings. Yellow rings on tail totaling 21 (Fig. 2). Preserved specimen somewhat faded, with no yellow visible (Fig. 3).
Figure 2.
General view of the studied specimen (YBU 230088) in life and its microhabitat a big tree trunk (by XQ Mi).
Figure 3.
Views of the studied specimen (YBU 230088) in preservation. General dorsal (A) and ventral (B) views of specimen, dorsal (C), ventral (D) and lateral (E) views of head (by P Guo).
Molecular phylogeny
In total, 1,047 bp of sequence data from 84 samples were aligned, with the generated novel sequence deposited in GenBank (Table 1). No deletions, insertions, or stop codons were detected, indicating that unintentional amplification of pseudogenes was unlikely (Zhang and Hewitt 1996). The best-fit evolutionary models of the data were: GTR+I+G for the first codon position, HKY+I+G for the second codon position, and GTR+G for the third codon position.
The mtDNA-based BI and ML analyses depicted relatively consistent topologies, with slight disagreement in several shallow nodes (Fig. 4). Both analyses indicated that all putative species of Lycodon formed a highly supported lineage (100 PP and 84% UFB). The newly collected specimen formed a clade with L.fasciatus, L.gongshan Vogel & Luo, 2011, L.butleri Boulenger, 1900, and L.cavernicolus Grismer, Quah, Anuar, Muin, Wood & Nor, 2014 with high support (100 PP and 97% UFB). Nevertheless, it occupied a basal position in relation to this clade and did not exhibit monophyly with any individual member. Uncorrected p-distances among the species within this clade ranged from 7.2% (L.gongshan and L.fasciatus) to 12.9% (L.gammiei and L.cavernicolus), while genetic distances between L.gammiei and its congeners within this clade ranged from 10.2% to 12.9% (data not shown).
Figure 4.
Bayesian 50% majority-rule consensus tree of Lycodon inferred from cyt b sequences analyzed using models detailed in the text. Posterior probabilities from BI analysis (>0.50) and Ultrafast bootstraps from ML analysis (>50%) are given adjacent to respective nodes for major nodes. Branch support indices are not given for most nodes to preserve clarity.
Discussion
Lycodongammiei is an exceedingly rare species, with a global record of only approximately 10 specimens. The majority of these are from Sikkim and West Bengal, India (Mistry et al. 2007; Chettri and Bhupathy 2009), with only two specimens reported in Cona County, Xizang, China (originally recorded in Eaglenest Wildlife Sanctuary, India) (Mistry et al. 2007) and Bhutan (Wangyal 2013), respectively. Based on the record by Mistry et al. (2007), Luo et al. (2010) recognized the existence of this species in China, although this recognition has been overlooked in subsequent publications (Wallach et al. 2014; Wang et al. 2020; Uetz et al. 2024). The discovery of this species in Medog County, Xizang, China, not only confirms its presence in China but also indicates a further eastward extension of its distribution.
Morphologically, the newly collected specimen shares most characters with the other conspecific specimens (Blanford 1878; Mistry et al. 2007; Chettri and Bhupathy 2009), including eight supralabials (3rd to 5th touching eye, 6th largest), single loreal, 2+3 temporals, one preocular, two postoculars, two genial pairs, cloacal plate entire, and dorsal scales in 17-17-15 rows. However, the new specimen has a greater number of ventral scales (228+1) than all previously reported specimens (205–220) (Mistry et al. 2007; Chettri and Bhupathy 2009).
The taxonomic status of L.gammiei has a controversial history. Although previously misidentified as both L.fasciatus (Wall 1911) and L.septentrionalis (Mahendra 1984), Mistry et al. (2007) later clarified its distinct status and validity based on morphological comparisons. In the current study, we present the first genetic data pertaining to this species. Notably, mtDNA-based phylogenetic analyses indicated that L.gammiei formed a highly supported monophyly with a clade containing L.fasciatus but was not the closest congener to L.fasciatus within this assemblage (Fig. 4). Lycodongammiei shows a greater genetic distance from L.septentrionalis than from L.fasciatus, further affirming its validity and unique taxonomic position. The closer genetic affinity of L.gammiei with the clade encompassing L.fasciatus aligns with their geographical closeness along the southern slopes of the Himalayas.
Lycodonzayuensis Jiang, Wang, Jin & Che, 2020 coexists with L.gammiei in southeastern Xizang, China (Che et al. 2020; Lyu et al. 2022). Both species exhibit similarities in external morphology, including dorsal scales in 17-17-15 rows, eight supralabials, one preocular, and two postoculars. However, the two species are genetically divergent (Fig. 2), and L.gammiei can be easily distinguished from L.zayuensis by its broader and fewer yellow body cross-bands (30–43 vs 88–93) (Blanford 1878; Lyu et al. 2022).
Citation
Shu F, Lyu B, Guo K, Zhang T, Mi X, Li L, Wu Y, Guo P (2024) Rediscovery of Lycodon gammiei (Blanford, 1878) (Serpentes, Colubridae) in Xizang, China, with comments on its systematic position. ZooKeys 1200: 65–74. https://doi.org/10.3897/zookeys.1200.117260
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Funding
This study was supported by the Second Tibetan Plateau Scientific Expedition and Research (STEP) Program (2019QZKK05010105), National Natural Science Foundation of China (32000308 and 32370486), and Yarlung Zangbo Grand Canyon National Nature Reserve Research and Monitor Program (Linzhi Forestry and Grassland Bureau)
Author contributions
Conceptualization: XM. Formal analysis: BL. Methodology: TZ. Resources: KG. Software: YW. Supervision: PG. Validation: LL. Writing – original draft: FS.
Author ORCIDs
Fu Shu https://orcid.org/0000-0002-6082-8112
Bing Lyu https://orcid.org/0000-0001-5594-1543
Keji Guo https://orcid.org/0000-0001-7508-1173
Tong Zhang https://orcid.org/0009-0009-1492-585X
Xiaoqi Mi https://orcid.org/0000-0003-1744-3855
Li Li https://orcid.org/0009-0007-4149-6662
Yayong Wu https://orcid.org/0000-0003-2752-4085
Data availability
All of the data that support the findings of this study are available in the main text.
References
- Agarwal I, Mistry VK, Athreya R. (2010) A preliminary checklist of the reptiles of Eaglenest Wildlife Sanctuary, West Kameng district, Arunachal Pradesh, India. Russian Journal of Herpetology 17(2): 81–93. [Google Scholar]
- Blanford WT. (1878) Notes on some Reptilia from the Himalayas and Burma. Part II. Journal of the Asiatic Society of Bengal (Natural History) 47(3): 125–131. [Google Scholar]
- Burbrink FT, Lawson R, Slowinski JB. (2000) Mitochondrial DNA phylogeography of the polytypic north American rat snake (Elapheobsoleta): A critique of the subspecies concept. Evolution 54(6): 2107–2118. 10.1111/j.0014-3820.2000.tb01253.x [DOI] [PubMed] [Google Scholar]
- Che J, Jiang K, Yan F, Zhang YP. (2020) Amphibians and Reptiles in Tibet – Diversity and Evolution. Science Press, Beijing, 691 pp. [Google Scholar]
- Chettri B, Bhupathy S. (2009) Occurrence of Dinodongammiei (Blanford, 1878) in Sikkim, Eastern Himalaya, India. Journal of Threatened Taxa 1(1): 60–61. 10.11609/JoTT.o1960.60-1 [DOI] [Google Scholar]
- Guo P, Zhang L, Liu Q, Li C, Pyron RA, Jiang K, Burbrink FT. (2013) Lycodon and Dinodon: One genus or two? Evidence from molecular phylogenetics and morphological comparisons. Molecular Phylogenetics and Evolution 68(1): 144–149. 10.1016/j.ympev.2013.03.008 [DOI] [PubMed] [Google Scholar]
- Kumar S, Stecher G, Tamura K. (2016) MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33(7): 1870–1874. 10.1093/molbev/msw054 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Lam-Tung N, Schmidt HA, Arndt VH, Quang MB. (2015) IQ-tree: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution (1): 268–274. 10.1093/molbev/msu300 [DOI] [PMC free article] [PubMed]
- Lanfear R, Calcott B, Ho SYW, Guindon S. (2012) PartitionFinder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29(6): 1695–1701. 10.1093/molbev/mss020 [DOI] [PubMed] [Google Scholar]
- Luo J, Ryabov SA, Luo Y, Gao HY, Luo ZR, Hu XC. (2010) Classification and distribution of the genus Dinodon. Sichuan Journal of Zoology 29(4): 579–582. [Google Scholar]
- Lyu B, Li QL, Li K, Li L, Shu F, Wu YY, Guo P. (2022) Expanded morphological description of the recently described Lycodonzayuensis (Serpentes: Colubridae). Zootaxa 5213(2): 159–168. 10.11646/zootaxa.5213.2.4 [DOI] [PubMed] [Google Scholar]
- Mahendra BC. (1984) Handbook of the snakes of India, Ceylon, Burma, Bangladesh, and Pakistan. Annals of Zoology, 22B, 412 pp. [Google Scholar]
- Mistry V, Vogel G, Tillack F. (2007) Rediscovery of Dinodongammiei (Blanford 1878) (Serpentes, Colubridae), with description of its validity. Hamadryad 31(2): 265–273. [Google Scholar]
- Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA. (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67(5): 901–904. 10.1093/sysbio/syy032 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Ronquist F, Teslenko M, Mark PVD, Ayres D, Darling A, Hohna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP. (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. 10.1093/sysbio/sys029 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Smith MA. (1943) The Fauna of British India, Ceylon and Burma, Including the Whole of the Indo-Chinese Sub-region. Reptilia and Amphibia. 3 (Serpentes). Taylor and Francis, London, 583 pp. [Google Scholar]
- Uetz P, Freed P, Aguilar R, Reyes F, Kudera J, Hošek J. (2024) The Reptile Database. http://www.reptile-database.org [Accessed on Jan. 2024]
- Wall F. (1923) A hand-list of snakes of the Indian empire. Part II. The Journal of the Bombay Natural History Society 29(3): 598–632. [Google Scholar]
- Wallach V, Williams KL, Boundy J. (2014) Snakes of the World: A Catalogue of Living and Extinct Species. Taylor and Francis, CRC Press, 1237 pp. 10.1201/b16901 [DOI] [Google Scholar]
- Wang K, Ren JL, Chen HM, Lyu ZT, Guo XG, Jiang K, Chen JM, Li JT, Guo P, Wang YY, Che J. (2020) The updated checklists of amphibians and reptiles of China. Biodiversity Science 28(2): 189–218. 10.17520/biods.2019238 [DOI] [Google Scholar]
- Wangyal JT. (2013) New records of reptiles and amphibians from Bhutan. Journal of Threatened Taxa 5(13): 4774–4783. 10.11609/JoTT.o3539.4774-83 [DOI] [Google Scholar]
- Zhang DX, Hewitt GM. (1996) Nuclear integrations: Challenges for mitochondrial DNA markers. Trends in Ecology & Evolution 11: 247–251. 10.1016/0169-5347(96)10031-8 [DOI] [PubMed] [Google Scholar]
Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Data Availability Statement
All of the data that support the findings of this study are available in the main text.