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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

Kai Wang Zhong-Bin Yu Gernot Vogel Jing Che

Kai Wang, Zhong-Bin Yu, Gernot Vogel, Jing Che. 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. Zoological Research, 2021, 42(1): 62-86. doi: 10.24272/j.issn.2095-8137.2020.286
Citation: Kai Wang, Zhong-Bin Yu, Gernot Vogel, Jing Che. 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. Zoological Research, 2021, 42(1): 62-86. doi: 10.24272/j.issn.2095-8137.2020.286

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

Funds: This research is supported by the Second Tibetan Plateau Scientific Expedition and Research (STEP) program (2019QZKK0501), Biodiversity Survey and Assessment Project of the Ministry of Ecology and Environment (2019HJ2096001006), China's Biodiversity Observation Network (Sino-BON), Southeast Asia Biodiversity Research Institute, CAS (Y4ZK111B01: 2017CASSEABRIQG002), and the Animal Branch of the Germplasm Bank of Wild Species, CAS (Large Research Infrastructure Funding) to J.C. and NSF GRFP 2017216966 to K.W.
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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。
    #Authors contributed equally to this work
  • 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

    GenusSpeciesVoucher No.LocalityGenBank accession No.
    Lycodonchapaensis comb. nov.KIZ 035013Lushui, Nujiang Prefecture, Yunnan, ChinaMW353742
    Lycodonchapaensis comb. nov.KIZ 038282Fugong, Nujiang Prefecture, Yunnan, ChinaMW353744
    Lycodonchapaensis comb. nov.KIZ 035113Dulongjiang, Nujiang Prefecture, Yunnan, ChinaMW353743
    Lycodonchapaensis comb. nov.KIZ 027593Tengchong, Baoshan, Yunnan, ChinaMW353741
    Lycodonchapaensis comb. nov.KIZ 034331Xichou, Wenshan Prefecture, Yunnan, ChinaMW353739
    Lycodonchapaensis comb. nov.KIZ 006753Mengzi, Honghe Prefecture, Yunnan, ChinaMW353737
    Lycodonchapaensis comb. nov.KIZ 049166Daweishan, Honghe Prefecture, Yunnan, ChinaMW353738
    Lycodonchapaensis comb. nov.KIZ 047084Jingdong, Puer, Yunnan, ChinaMW353740
    LycodongongshanKIZ 035112Dulongjiang, Nujiang Prefecture, Yunnan, ChinaMW353748
    LycodongongshanKIZ 035114Dulongjiang, Nujiang Prefecture, Yunnan, ChinaMW353749
    Lycodongongshan/Yunlong Nature Reserve, Dali, Yunnan, ChinaMW353747
    Lycodonobvelatus sp. nov.KIZ 040146Panzhihua, Sichuan, ChinaMW353745
    LycodonseptentrionalisCIB 117521Medog, Nyinchi Prefecture, Tibet, ChinaMW353736
    Lycodonserratus sp. nov.KIZ 038335Deqin, Yunnan, ChinaMW353746
    BoigacynodonKU324614Negros Occidental, PhilippinesKC010340
    DasypeltisatraCAS 201641Kabale, UgandaAF471065
    LycodonalbofuscusUSMHC 1457KX822584
    LycodonalcalaiKU327848Municipality of Sabtang, Batanes, PhilippinesKC010345
    LycodonbanksiVNUF R.2015.20Khammouane, LaosMH669272
    Lycodonbibonius KU304589Cagayan, PhilippinesKC010351
    LycodonbutleriLSUHC 9137Perak, MalaysiaKJ607891
    LycodonbutleriLSUHC 8365Perak, MalaysiaKJ607892
    LycodoncapucinusMVZ 291704MK844523
    LycodoncapucinusMVZ 291703MK844522
    LycodoncathayaSYS r001542Longsheng County, Guangxi, ChinaMT602075
    LycodoncavernicolusLSUHC 10500Perlis, MalaysiaKJ607890
    LycodoncavernicolusLSUHC 9985Perlis, MalaysiaKJ607889
    Lycodonchapaensis comb nov. (septentrionalis)Yunnan, ChinaMK201305
    LycodonchrysopraterosKU 307720Cagayan, PhilippinesKC010360
    LycodondumeriliiPNM7751KC010363
    LycodondumeriliiKU 305168KC010362
    LycodoneffraenisLSUHC 9670KC010376
    LycodoneffraenisKU 328526Karome, Nakhon Si Thammarat, ThailandKC010364
    LycodonfasciatusCAS 234875Mindat, Chin State, MyanmarKC010365
    LycodonfasciatusCAS 234957Mindat, Chin State, MyanmarKC010366
    LycodonfasciatusSYS r001654Xishuangbanna, Yunnan, ChinaMK201559
    LycodonfasciatusKIZ 014125Xishuangbanna, Yunnan, ChinaMK201557
    LycodonflavozonatusSYSr000640Huangganshan, Jiangxi, ChinaMK201413
    LycodonflavozonatusHS15101Huangshan, Anhui, ChinaMK201312
    LycodongongshanGP3548Lingcang, Yunnan, ChinaKP901026
    LycodongongshanGP3547Lingcang, Yunnan, ChinaKP901025
    LycodongongshanGP3546Lingcang, Yunnan, ChinaKP901024
    LycodongongshanGP3516Lingcang, Yunnan, ChinaKP901022
    LycodonjaraCAS 235387Putao, Kachin, MyanmarKC010367
    LycodonlaoensisKU 328529Karome, Nakhon Si Thammarat, ThailandKC010371
    LycodonlaoensisFMNH 258659Salavan, LaosKC010368
    Lycodonliuchengchaoi (fasciatus)GP 2094Nanling, Guangdong, ChinaKC733201
    Lycodonliuchengchaoi (fasciatus)GP 2097Chebaling, Guangdong, ChinaKC733202
    LycodonliuchengchaoiJK 201704Ningshan, Shaanxi, ChinaMK201563
    LycodonliuchengchaoiSYS r001654Shennongjia, Hubei, ChinaMK201580
    LycodonliuchengchaoiDL 14315KF732928
    LycodonmeridionalisVNUF R.2017.123Thanh Hoa, VietnamMH669270
    LycodonmeridionalisVNUF R.2012.4Bac Kan, VietnamMH669271
    LycodonmultizonatusSyS r002411Baishuijiang National Nature Reserve, Longnan, Gansu, ChinaMT625863
    LycodonmultizonatusKIZ 01623Luding, Sichuan, ChinaKF732926
    Lycodonmultizonatus (liuchengchaoi)HS 11252Sanjiazhai, Yunnan, ChinaMK201303
    Lycodonchapaensis comb. nov. (namdongensis)VNUF R. 2017.23Thanh Hoa, VietnamMK585007
    LycodonpictusZFMK93747Cao Bang, VietnamMN395830
    LycodonpictusZFMK93746Cao Bang, VietnamMN395829
    LycodonrufozonatusLSUMZ 44977AF471063
    LycodonrufozonatusGP133Tongjiang, Sichuan, ChinaKC733194
    LycodonruhstratiGP2243Ruyuan, Guangdong, ChinaKC733208
    LycodonruhstratiGP2049Chebaling, Guangdong, ChinaKC733200
    LycodonruhstratiGP991Gongcheng, Guangxi, ChinaKC733197
    LycodonruhstratiGP285Junlian, Sichuan, ChinaKC733195
    LycodonruhstratiSYSr001631Huangping, Guangxi, ChinaMK201538
    LycodonruhstratiSYSr001555Yangmeiao, Guangxi, ChinaMK201521
    LycodonruhstratiSYSr001309Jiulianshan, Jiangxi, ChinaMK201473
    LycodonruhstratiSYSr001275Jiangshi, Fujian, ChinaMK201467
    LycodonruhstratiHS12069Chebaling, Guangdong, ChinaMK201310
    LycodonruhstratiDL12678KF732925
    LycodonsealeiKU327571Palawan, PhilippinesKC010384
    LycodonsealeiKU309447Palawan, PhilippinesKC010385
    LycodonsemicarinatusRyukyu Archipelago, JapanAB008539
    LycodonsidikiMZB.Ophi.5980KX822583
    LycodonstormiJAM7487KC010380
    LycodonstriatusFBRC_DNA205MK089444
    LycodonsubcinctusMVZ291678IndonesiaMK844529
    LycodonsubcinctusMVZ291679IndonesiaMK844530
    LycodonsubcinctusMVZ291680IndonesiaMK844531
    LycodonsubcinctusMVZ291681IndonesiaMK844532
    LycodonsubcinctusMVZ291682IndonesiaMK844533
    LycodonsubcinctusMVZ291683IndonesiaMK844534
    LycodonsubcinctusMVZ291684IndonesiaMK844535
    LycodonsubcinctusMVZ291685IndonesiaMK844536
    LycodonsubcinctusLSUHC5016Pahang, West MalaysiaKC010382
    LycodonsynaptorGP2188Yunnan, ChinaKC733204
    LycodonsynaptorSYS r001775Darongshan, Guangxi, ChinaMK201582
    LycodonsynaptorSYS r001800Dawuling, Guangdong, ChinaMK201581
    LycodonsynaptorHS13002Honghe, Yunnan, ChinaMK201309
    LycodonsynaptorHS12087Kunming, Yunnan, ChinaMK201308
    LycodonsynaptorHS11006Mengzi, Yunnan, ChinaMK201304
    LycodonsynaptorGP3515Lingcang, Yunnan, ChinaKP901021
    LycodonsynaptorGP3288Maandi, Yunnan, ChinaKP901020
    LycodonsynaptorGP3270Daweishan, Yunnan, ChinaKP901019
    LycodonsynaptorGP3545Yunnan, ChinaKP901023
    LycodonzawiCAS 239944Kyaukpyu, Rakhine State, MyanmarKC010386
    LycodonzawiCAS 210323Thabakesay, Saging, MyanmarAF471040
    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.
    下载: 导出CSV

    Table  2.   Uncorrected genetic distance (%) based on 1 117 bp fragment of cyt b among selected members of the genus Lycodon

    1234567891011
    1L. serratus sp. nov.
    2L. obvelatus sp. nov.11.9
    3L. septentrionalis14.314.2
    4L. namdongensis14.814.96.6
    5L. septentrionalis13.7–15.214.3–15.05.7–7.40.1–1.70–1.8
    6L. gongshan10.1–10.611.3–12.312.5–14.612.5–14.511.3–14.70–2.2
    7L. fasciatus9.9–11.113.3–13.612.4–14.112.9–14.112.4–14.45.7–8.00–2.4
    8L. pictus8.3–8.412.2–12.313.5–13.713.012.5–13.310.5–11.410.7–12.00.6
    9L. liuchengchaoi5.2–5.411.1–11.512.6–14.213.2–14.512.9–14.68.1–10.59.4–11.77.9–8.40–0.2
    10L. multizonatus3.6–4.011.3–12.012.5–14.013.1–14.312.8–14.68.2–11.29.4–11.78.2–8.95.4–6.40.5–2.2
    11L. synaptor10.9–12.68.6–9.512.1–13.812.7–14.111.4–14.28.5–12.110.1–12.09.4–11.110.5–12.510.9–12.80–3.0
    The diagonal values are intraspecific genetic distances for the species with multiple available specimens.
    下载: 导出CSV

    Table  3.   Comparisons between holotype of Dinodon septentrionale chapaense, holotype of Lycodon namdongensis, true L. septentrionalis from Tibet, and “L. septentrionalis” from Yunnan Province

    SpeciesLycodon septentrionalisDinodon septentrionale chapaenseL. namdongensisL. septentrionalis
    Voucher No.CIB 117521, CIB M20150607MNHN-RA-1933.0011 (holotype)VNUF R. 2017.23 (holotype)KIZ 006753, 027593, 035594, 038282, 034331
    Sample size2 (1)115
    ToL1 1871 051+723691–1 114
    SVL945–955890575564–915
    TaL242161 (incomplete)148127–199
    TaL/ToL20.4%20.5%17.1%–18.4% (average 17.6%)
    DSRH17171717
    DSRM17171717
    DSRV15151515
    SL8 (2-3-3)8 (2-3-3)8 (2-3-3)7 or 8 (2-3-3, 3-2-3, or 2-2-3)
    IL9 (4)9 (5)10 (5)8–10 (first 4 or 5 in contact with anterior chin shield)
    PrO1111
    PtO2222
    LoR1111
    LoR-ENoNoNoNo
    TMP2+32+32+32+2 or 2+3
    VEN207–212224218200–225
    SC7856+8574–84
    Cloacal PlateEntireEntireEntireEntire
    MT81211 or 12
    NCBAbsenceAbsenceAbsenceAbsence
    NDB33–35282323–37
    NTB19111411–14
    BSTKeeledSmoothSmoothSmooth 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.
    下载: 导出CSV

    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)

    SpeciesLycodon serratus
    sp. nov.
    L. obvelatus sp. nov.L. multizonatus L. gongshanL. fasciatusL. liuchengchaoi
    SexMMMMFMFMF
    Sample size1 (holotype)1 (holotype)422132 (1)5 (3)
    ToL628551440–520928–963753–1003696418–613747389–615
    SVL480447350–428691–740589–798553341–493595–676309–481
    TaL14810490–96223–237164–20714371–12015280–138
    TaL/ToL23.6%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%
    DSRH191717171717171717
    DSRM171717171717171717
    DSRV151515151515151515
    SL8 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)
    IL10 (5)8 (4 or 5)8 (4)9 (4)9 (4)9 (5)9 (4) or 9 (5)8 (4)8 (4)
    PrO111111111
    PtO222222222
    LoR111111111
    LoR-EYesYesYesYesYesYesYes/No (*)YesYes
    TMP2+22+2 or 2+32+32+2 or 2+32+22+22+2 or 2+31+2 or 2+21+2, 2+2, or 2+3
    VEN198199191–195210–212209–215198205–211202–204200–228
    SC847663–7594–9692(n=1)8466–9568–69+75–81
    Cloacal plateDividedEntireDividedEntireEntireEntireEntireDividedDivided
    MT12 (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 adultsPresencePresencePresenceAbsenceAbsenceAbsenceAbsencePresencePresence
    NDB663155–6237 or 3832–363431–3740–4533–45
    NTB261311–1915 or 1613 (n=1)1611–1710–1511–13
    BSTSmoothSmoothKeeledKeeledKeeledKeeledKeeledKeeledKeeled
    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.
    下载: 导出CSV
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  • 收稿日期:  2020-09-29
  • 录用日期:  2020-12-17
  • 网络出版日期:  2020-12-18
  • 刊出日期:  2021-01-18

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