中国白环蛇属Lycodon H. Boie in Fitzinger, 1827分类厘定,记两新种并恢复及提升白链蛇沙坝亚种Dinodon septentrionale chapaense Angel, Bourret, 1933
doi: 10.24272/j.issn.2095-8137.2020.286
Contribution to the taxonomy of the genus Lycodon H. Boie in Fitzinger, 1827 (Reptilia: Squamata: Colubridae) in China, with description of two new species and resurrection and elevation of Dinodon septentrionale chapaense Angel, Bourret, 1933
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摘要: 虽然近年来白环蛇属Lycodon分类研究进展迅速，但属内部分种组的分类问题仍有待解决，特别是在我国分布的种组及物种。一方面而言，国内仍有很多区域未开展过系统的爬行动物多样性调查；另一方面，已调查的区域仍有大量已知物种的存疑纪录亟需进一步确认。本研究结合形态学和分子生物学证据，对中国白环蛇属部分现存分类问题进行了梳理：其中，首次于云南西北部金沙江干热河谷记录白环蛇属物种，并将其描述为一新种，即锯纹白环蛇L. serratus sp. nov. ；同时于四川攀枝花市区公园内采集到原记录的双全白环蛇L. fasciatus，经对比确定其实际代表了双全白环蛇种组一新种，即隐士白环蛇L. obvelatus sp. nov. 。此外，厘定了白链蛇L. septentrionalis在中国的分布情况，恢复白链蛇沙坝亚种Dinodon septentrionale chapaense有效性，并将其提升至种级，即沙坝白环蛇L. chapaensis comb. nov.，同时将2019年于越南北部描述的南东白环蛇L. namdongensis定为沙坝白环蛇的次定同物异名；而恢复、提升后的沙坝白环蛇也代表了我国蛇类的新纪录。最后，本文在已发表文献资料的基础上，纠正了双全白环蛇和黑背白环蛇L. ruhstrati的部分误定和因此产生的错误分布纪录，指出了中国白环蛇属分类仍存的部分问题，并更新了我国白环蛇属已知20种的分类检索表及分布情况。Abstract: While considerable progress has been made in the taxonomic studies of the genus Lycodon in Asia, questions remain to be clarified regarding the taxonomy of certain groups, particularly those containing species in China. Not only do many regions in China remain overlooked by herpetologists, resulting in the possibility of undiscovered new species, but the surveyed areas also have suspicious records of recognized congeners that require taxonomic confirmations. Combining both morphological and genetic data, we tackle these outstanding issues in the taxonomy of Lycodon in China. In particular, we discover two new species of Lycodon: one from the previously neglected hot-dry valley in the northern Hengduan Mountain Region close to Tibet, and another recluse and cryptic species from the L. fasciatus complex in the downtown park of a major city in southern Sichuan Province. Additionally, we clarify the distribution of L. septentrionalis in China and resurrect and elevate its junior synonym subspecies, Dinodon septentrionale chapaense, as a full, valid species, and we synonymize the recently described L. namdongensis to the resurrected L. chapaensis comb. nov .. Lycodon chapaensis comb. nov . thus represents a new national record of reptilian fauna of China. Lastly, based on literature review, we also correct some of the erroneous records of L. fasciatus and L. ruhstrati in China, point out remaining taxonomic issues of the genus for future research, and update the dichotomous key and distribution of the 20 species of Lycodon currently recorded from China.
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Key words:
- Erroneous records /
- Guangdong /
- Hunan /
- Misidentification /
- Serpentes /
- Sichuan /
- Wolf Snake /
- Yunnan
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Figure 1. Distributions of focal members of the genus Lycodon in Southwest China and nearby countries for this paper
Different shapes represent different species, and numbers indicate different localities where the species have been recorded (details see Appendix I). Number 1 always indicates the type locality of that given species. Bifurcating white lines indicate sympatric distribution of two species at the same site. Lycodon serratus sp. nov. (brown star): Geyading, Deqin County, Yunnan, China. Lycodon obvelatus sp. nov. (red star): Panzhihua City, Panzhihua, Sichuan, China. Lycodon chapaensis comb. nov. (triangle): the orange triangle indicates the type locality of its junior synonym, L. namdongensis at Quan Son, Thanh Hoa Province, Vietnam; gray triangle indicates remaining localities of the species. Lycodon fasciatus sensu stricto (circle); L. gongshan (square); L. multizonatus (trapezoid); L. septentrionalis sensu stricto (pentagon); and L. synaptor (hexagon).
Figure 2. Phylogenetic trees of the genus Lycodon inferred by Bayesian analyses (BI) based on 1 117 bp of mitochondrial gene cyt b
Both bootstrap supports (BS) and Bayesian posterior probabilities (BPP) are indicated on each of the corresponding node. “–” indicates a differential topology between ML and BI results. Support values for strongly supported (BS≥70, BPP≥0.95) intraspecific nodes were omitted, except few cases where the nodes are specifically referred in text.
Figure 3. Comparisons between true Lycodon septentrionalis (A, B); L. namdongensis (C), and Yunnan specimen of “D. septentrionalis” (D–G)
A, B: Closeup and dorsolateral overview of a un-vouchered individual from Medog, Tibet, China; C: Holotype of L. namdongensis (VNUF R. 2017.23) from Quan Son, Thanh Hoa, Vietnam; D: Un-vouchered individual from Daweishan Nature Reserve, Honghe Prefecture, Yunnan; E: Vouchered adult male from Yongping County, Dali, Yunnan, China (KIZ 035594); F: Vouchered adult male from Tengchong County, Baoshan, Yunnan (KIZ 027593); G: Hemipenis of KIZ 027593 after preservation. Photos of L. namdongensis taken from Luu et al. (2019), remaining photos by Chao Wu, Kai Wang, Shaobin Hou, Weiliang Xie, and Zhongbin Yu.
Figure 4. The dorsolateral view (1) and ventral view (2) of the holotype of Lycodon serratus sp. nov. (KIZ 038335) (A) and L. obvelatus sp. nov. in life (KIZ 040146) (B) (Photos by Wen-Jie Dong and Kai Wang)
Figure 5. Comparisons of (1) dorsal overview, (2) ventral overview, (3) lateral head, (4) dorsal head, (5) ventral head, and (6) dorsum close-up among Lycodon serratus sp. nov. (KIZ 038335; holotype) (A), L. obvelatus sp. nov. (KIZ 040146; holotype) (B), L. multizonatus (KIZ 01623; topotype) (C), L. fasciatus (KIZ 74II0262) (D), L. gongshan (KIZ 730034; holotype) (E), “L. septentrionalis” (KIZ 035594; from Dali, Yunnan, China) (F), L. chapaensis comb. nov. (MNHN-RA-1933.0011, holotype; from Chapa, Tonkin, Vietnam) (G), and L. septentrionalis (CIB 117521; from Medog, Tibet, China) (H) (Photos of the holotype of L. chapaensis comb. nov. are obtained from the website of Muséum National d’Histore Naturelle, remaining photos by Zhong-Bin Yu and Jin-Long Ren)
Figure 6. The habitats at the type locality of Lycodon serratus sp. nov. near Geyading Village, Deqin County, Yunnan Province, China (A) and L. obvelatus sp. nov. in Panzhihua City Park, Panzhihua, Sichuan, China (B) (Photos by Zhong-Bin Yu and Ben-Fu Miao)
Figure 7. Photos of live Lycodon aulicus-capucinus complex from China
A: From Hong Kong, China; B: From Yingjiang County, Yunnan, China. Photos by Jin-Long Ren and Franco Leung Ka Wah.
Table 1. Samples and their Genbank accession Nos. in the present study
Genus Species Voucher No. Locality GenBank accession No. Lycodon chapaensis comb. nov. KIZ 035013 Lushui, Nujiang Prefecture, Yunnan, China MW353742 Lycodon chapaensis comb. nov. KIZ 038282 Fugong, Nujiang Prefecture, Yunnan, China MW353744 Lycodon chapaensis comb. nov. KIZ 035113 Dulongjiang, Nujiang Prefecture, Yunnan, China MW353743 Lycodon chapaensis comb. nov. KIZ 027593 Tengchong, Baoshan, Yunnan, China MW353741 Lycodon chapaensis comb. nov. KIZ 034331 Xichou, Wenshan Prefecture, Yunnan, China MW353739 Lycodon chapaensis comb. nov. KIZ 006753 Mengzi, Honghe Prefecture, Yunnan, China MW353737 Lycodon chapaensis comb. nov. KIZ 049166 Daweishan, Honghe Prefecture, Yunnan, China MW353738 Lycodon chapaensis comb. nov. KIZ 047084 Jingdong, Puer, Yunnan, China MW353740 Lycodon gongshan KIZ 035112 Dulongjiang, Nujiang Prefecture, Yunnan, China MW353748 Lycodon gongshan KIZ 035114 Dulongjiang, Nujiang Prefecture, Yunnan, China MW353749 Lycodon gongshan / Yunlong Nature Reserve, Dali, Yunnan, China MW353747 Lycodon obvelatus sp. nov. KIZ 040146 Panzhihua, Sichuan, China MW353745 Lycodon septentrionalis CIB 117521 Medog, Nyinchi Prefecture, Tibet, China MW353736 Lycodon serratus sp. nov. KIZ 038335 Deqin, Yunnan, China MW353746 Boiga cynodon KU324614 Negros Occidental, Philippines KC010340 Dasypeltis atra CAS 201641 Kabale, Uganda AF471065 Lycodon albofuscus USMHC 1457 – KX822584 Lycodon alcalai KU327848 Municipality of Sabtang, Batanes, Philippines KC010345 Lycodon banksi VNUF R.2015.20 Khammouane, Laos MH669272 Lycodon bibonius KU304589 Cagayan, Philippines KC010351 Lycodon butleri LSUHC 9137 Perak, Malaysia KJ607891 Lycodon butleri LSUHC 8365 Perak, Malaysia KJ607892 Lycodon capucinus MVZ 291704 – MK844523 Lycodon capucinus MVZ 291703 – MK844522 Lycodon cathaya SYS r001542 Longsheng County, Guangxi, China MT602075 Lycodon cavernicolus LSUHC 10500 Perlis, Malaysia KJ607890 Lycodon cavernicolus LSUHC 9985 Perlis, Malaysia KJ607889 Lycodon chapaensis comb nov. (septentrionalis) – Yunnan, China MK201305 Lycodon chrysoprateros KU 307720 Cagayan, Philippines KC010360 Lycodon dumerilii PNM7751 – KC010363 Lycodon dumerilii KU 305168 – KC010362 Lycodon effraenis LSUHC 9670 – KC010376 Lycodon effraenis KU 328526 Karome, Nakhon Si Thammarat, Thailand KC010364 Lycodon fasciatus CAS 234875 Mindat, Chin State, Myanmar KC010365 Lycodon fasciatus CAS 234957 Mindat, Chin State, Myanmar KC010366 Lycodon fasciatus SYS r001654 Xishuangbanna, Yunnan, China MK201559 Lycodon fasciatus KIZ 014125 Xishuangbanna, Yunnan, China MK201557 Lycodon flavozonatus SYSr000640 Huangganshan, Jiangxi, China MK201413 Lycodon flavozonatus HS15101 Huangshan, Anhui, China MK201312 Lycodon gongshan GP3548 Lingcang, Yunnan, China KP901026 Lycodon gongshan GP3547 Lingcang, Yunnan, China KP901025 Lycodon gongshan GP3546 Lingcang, Yunnan, China KP901024 Lycodon gongshan GP3516 Lingcang, Yunnan, China KP901022 Lycodon jara CAS 235387 Putao, Kachin, Myanmar KC010367 Lycodon laoensis KU 328529 Karome, Nakhon Si Thammarat, Thailand KC010371 Lycodon laoensis FMNH 258659 Salavan, Laos KC010368 Lycodon liuchengchaoi (fasciatus) GP 2094 Nanling, Guangdong, China KC733201 Lycodon liuchengchaoi (fasciatus) GP 2097 Chebaling, Guangdong, China KC733202 Lycodon liuchengchaoi JK 201704 Ningshan, Shaanxi, China MK201563 Lycodon liuchengchaoi SYS r001654 Shennongjia, Hubei, China MK201580 Lycodon liuchengchaoi DL 14315 – KF732928 Lycodon meridionalis VNUF R.2017.123 Thanh Hoa, Vietnam MH669270 Lycodon meridionalis VNUF R.2012.4 Bac Kan, Vietnam MH669271 Lycodon multizonatus SyS r002411 Baishuijiang National Nature Reserve, Longnan, Gansu, China MT625863 Lycodon multizonatus KIZ 01623 Luding, Sichuan, China KF732926 Lycodon multizonatus (liuchengchaoi) HS 11252 Sanjiazhai, Yunnan, China MK201303 Lycodon chapaensis comb. nov. (namdongensis) VNUF R. 2017.23 Thanh Hoa, Vietnam MK585007 Lycodon pictus ZFMK93747 Cao Bang, Vietnam MN395830 Lycodon pictus ZFMK93746 Cao Bang, Vietnam MN395829 Lycodon rufozonatus LSUMZ 44977 – AF471063 Lycodon rufozonatus GP133 Tongjiang, Sichuan, China KC733194 Lycodon ruhstrati GP2243 Ruyuan, Guangdong, China KC733208 Lycodon ruhstrati GP2049 Chebaling, Guangdong, China KC733200 Lycodon ruhstrati GP991 Gongcheng, Guangxi, China KC733197 Lycodon ruhstrati GP285 Junlian, Sichuan, China KC733195 Lycodon ruhstrati SYSr001631 Huangping, Guangxi, China MK201538 Lycodon ruhstrati SYSr001555 Yangmeiao, Guangxi, China MK201521 Lycodon ruhstrati SYSr001309 Jiulianshan, Jiangxi, China MK201473 Lycodon ruhstrati SYSr001275 Jiangshi, Fujian, China MK201467 Lycodon ruhstrati HS12069 Chebaling, Guangdong, China MK201310 Lycodon ruhstrati DL12678 – KF732925 Lycodon sealei KU327571 Palawan, Philippines KC010384 Lycodon sealei KU309447 Palawan, Philippines KC010385 Lycodon semicarinatus – Ryukyu Archipelago, Japan AB008539 Lycodon sidiki MZB.Ophi.5980 – KX822583 Lycodon stormi JAM7487 – KC010380 Lycodon striatus FBRC_DNA205 – MK089444 Lycodon subcinctus MVZ291678 Indonesia MK844529 Lycodon subcinctus MVZ291679 Indonesia MK844530 Lycodon subcinctus MVZ291680 Indonesia MK844531 Lycodon subcinctus MVZ291681 Indonesia MK844532 Lycodon subcinctus MVZ291682 Indonesia MK844533 Lycodon subcinctus MVZ291683 Indonesia MK844534 Lycodon subcinctus MVZ291684 Indonesia MK844535 Lycodon subcinctus MVZ291685 Indonesia MK844536 Lycodon subcinctus LSUHC5016 Pahang, West Malaysia KC010382 Lycodon synaptor GP2188 Yunnan, China KC733204 Lycodon synaptor SYS r001775 Darongshan, Guangxi, China MK201582 Lycodon synaptor SYS r001800 Dawuling, Guangdong, China MK201581 Lycodon synaptor HS13002 Honghe, Yunnan, China MK201309 Lycodon synaptor HS12087 Kunming, Yunnan, China MK201308 Lycodon synaptor HS11006 Mengzi, Yunnan, China MK201304 Lycodon synaptor GP3515 Lingcang, Yunnan, China KP901021 Lycodon synaptor GP3288 Maandi, Yunnan, China KP901020 Lycodon synaptor GP3270 Daweishan, Yunnan, China KP901019 Lycodon synaptor GP3545 Yunnan, China KP901023 Lycodon zawi CAS 239944 Kyaukpyu, Rakhine State, Myanmar KC010386 Lycodon zawi CAS 210323 Thabakesay, Saging, Myanmar AF471040 New sequences are indicated in bold. Species name in parentheses indicates the previous synonym or misidentified species name used for the given sequence. “/” indicates non-voucher data, “–” indicates that information is not available or could not be located. 
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Table 2. Uncorrected genetic distance (%) based on 1 117 bp fragment of cyt b among selected members of the genus Lycodon
1 2 3 4 5 6 7 8 9 10 11 1 L. serratus sp. nov. − 2 L. obvelatus sp. nov. 11.9 − 3 L. septentrionalis 14.3 14.2 − 4 L. namdongensis 14.8 14.9 6.6 − 5 “L. septentrionalis” 13.7–15.2 14.3–15.0 5.7–7.4 0.1–1.7 0–1.8 6 L. gongshan 10.1–10.6 11.3–12.3 12.5–14.6 12.5–14.5 11.3–14.7 0–2.2 7 L. fasciatus 9.9–11.1 13.3–13.6 12.4–14.1 12.9–14.1 12.4–14.4 5.7–8.0 0–2.4 8 L. pictus 8.3–8.4 12.2–12.3 13.5–13.7 13.0 12.5–13.3 10.5–11.4 10.7–12.0 0.6 9 L. liuchengchaoi 5.2–5.4 11.1–11.5 12.6–14.2 13.2–14.5 12.9–14.6 8.1–10.5 9.4–11.7 7.9–8.4 0–0.2 10 L. multizonatus 3.6–4.0 11.3–12.0 12.5–14.0 13.1–14.3 12.8–14.6 8.2–11.2 9.4–11.7 8.2–8.9 5.4–6.4 0.5–2.2 11 L. synaptor 10.9–12.6 8.6–9.5 12.1–13.8 12.7–14.1 11.4–14.2 8.5–12.1 10.1–12.0 9.4–11.1 10.5–12.5 10.9–12.8 0–3.0 The diagonal values are intraspecific genetic distances for the species with multiple available specimens.
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Table 3. Comparisons between holotype of Dinodon septentrionale chapaense, holotype of Lycodon namdongensis, true L. septentrionalis from Tibet, and “L. septentrionalis” from Yunnan Province
Species Lycodon septentrionalis Dinodon septentrionale chapaense L. namdongensis “L. septentrionalis” Voucher No. CIB 117521, CIB M20150607 MNHN-RA-1933.0011 (holotype) VNUF R. 2017.23 (holotype) KIZ 006753, 027593, 035594, 038282, 034331 Sample size 2 (1) 1 1 5 ToL 1 187 1 051+ 723 691–1 114 SVL 945–955 890 575 564–915 TaL 242 161 (incomplete) 148 127–199 TaL/ToL 20.4% – 20.5% 17.1%–18.4% (average 17.6%) DSRH 17 17 17 17 DSRM 17 17 17 17 DSRV 15 15 15 15 SL 8 (2-3-3) 8 (2-3-3) 8 (2-3-3) 7 or 8 (2-3-3, 3-2-3, or 2-2-3) IL 9 (4) 9 (5) 10 (5) 8–10 (first 4 or 5 in contact with anterior chin shield) PrO 1 1 1 1 PtO 2 2 2 2 LoR 1 1 1 1 LoR-E No No No No TMP 2+3 2+3 2+3 2+2 or 2+3 VEN 207–212 224 218 200–225 SC 78 56+ 85 74–84 Cloacal Plate Entire Entire Entire Entire MT 8 – 12 11 or 12 NCB Absence Absence Absence Absence NDB 33–35 28 23 23–37 NTB 19 11 14 11–14 BST Keeled Smooth Smooth Smooth or only posterior vertebral row feebly keeled Abbreviations are explained in methods. “–” indicates data not available. The number in parentheses for the Sample size row indicate differential sample size for total length and tail length due to the incomplete tail of one of the specimen (CIB M20150607). Except the holotype of L. namdongensis, all remaining specimens were examined in person by authors.
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Table 4. Comparison of key morphological characters between the holotypes of Lycodon serratus sp. nov., L. obvelatus sp. nov., and morphologically similar congeners that are also found in the Hengduan Mountain Region (i.e., L. gongshan, L. fasciatus, L. multizonatus, and L. liuchengchaoi)
Species Lycodon serratus
sp. nov.L. obvelatus sp. nov. L. multizonatus L. gongshan L. fasciatus L. liuchengchaoi Sex M M M M F M F M F Sample size 1 (holotype) 1 (holotype) 4 2 2 1 3 2 (1) 5 (3) ToL 628 551 440–520 928–963 753–1003 696 418–613 747 389–615 SVL 480 447 350–428 691–740 589–798 553 341–493 595–676 309–481 TaL 148 104 90–96 223–237 164–207 143 71–120 152 80–138 TaL/ToL 23.6% 18.9% 17.7%–20.5% 23.1%–23.2% 21.8% (n=1) 20.5% 17.0%–19.6% 20.3% 20.6%–24.8% DSRH 19 17 17 17 17 17 17 17 17 DSRM 17 17 17 17 17 17 17 17 17 DSRV 15 15 15 15 15 15 15 15 15 SL 8 or 9 (2-3-3 or 2-4-3) 8 (2-3-3) 8 (2-3-3) 8 (2-3-3) 8 (2-3-3) 8 (2-3-3) 8 (2-3-3) 7 (2-3-2) or 8 (2-3-3) 8 (2-3-3) IL 10 (5) 8 (4 or 5) 8 (4) 9 (4) 9 (4) 9 (5) 9 (4) or 9 (5) 8 (4) 8 (4) PrO 1 1 1 1 1 1 1 1 1 PtO 2 2 2 2 2 2 2 2 2 LoR 1 1 1 1 1 1 1 1 1 LoR-E Yes Yes Yes Yes Yes Yes Yes/No (*) Yes Yes TMP 2+2 2+2 or 2+3 2+3 2+2 or 2+3 2+2 2+2 2+2 or 2+3 1+2 or 2+2 1+2, 2+2, or 2+3 VEN 198 199 191–195 210–212 209–215 198 205–211 202–204 200–228 SC 84 76 63–75 94–96 92(n=1) 84 66–95 68–69+ 75–81 Cloacal plate Divided Entire Divided Entire Entire Entire Entire Divided Divided MT 12 (6-1-1-4 or 6-1-2-3) 11 (7-1-1-2) 11 (no distint gap) (n=2) 10 (7-1-2) or 11 (7-2-2) 11 (5-3-3) (n=1) 12 (8-2-2) 12 (8-2-2) 8 – CB in adults Presence Presence Presence Absence Absence Absence Absence Presence Presence NDB 66 31 55–62 37 or 38 32–36 34 31–37 40–45 33–45 NTB 26 13 11–19 15 or 16 13 (n=1) 16 11–17 10–15 11–13 BST Smooth Smooth Keeled Keeled Keeled Keeled Keeled Keeled Keeled Abbreviations can be found in methods. “–” indicates not available due to incomplete tail. “*” indicates only loreals of a single specimen (KIZ 75I473) do not enter orbit. The number in parentheses for the Sample Size row indicates differential sample size for total length and tail length for L. liuchengchaoi. Data for the female of L. gongshan and for all L. liuchengchaoi were obtained from literature (L. gongshan: Vogel& Luo, 2011; L. liuchengchaoi: Peng et al., 2014, 2017, 2018; Zhang et al., 2011b; Zhang et al., 2019). M: Male; F: Female.
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