Research Article |
Corresponding author: Ying-Yong Wang ( wangyy@mail.sysu.edu.cn ) Academic editor: Thomas Ziegler
© 2020 Jian Wang, Shuo Qi, Zhi-Tong Lyu, Zhao-Chi Zeng, Ying-Yong Wang.
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:
Wang J, Qi S, Lyu Z-T, Zeng Z-C, Wang Y-Y (2020) A new species of the genus Lycodon (Serpentes, Colubridae) from Guangxi, China. ZooKeys 954: 85-108. https://doi.org/10.3897/zookeys.954.53432
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A new species of colubrid snake, Lycodon cathaya sp. nov., is described based on two adult male specimens collected from Huaping Nature Reserve, Guangxi, southern China. In a phylogenetic analyses, the new species is shown to be a sister taxon to the clade composed of L. futsingensis and L. namdongensis with low statistical support, and can be distinguished from all known congeners by the significant genetic divergence in the mitochondrial cytochrome b gene fragment (p-distance ≥ 7.9%), and morphologically by the following combination of characters: (1) dorsal scales in 17–17–15 rows, smooth throughout; (2) supralabials eight, third to fifth in contact with eye, infralabials nine; (3) ventral scales 199–200 (plus two preventral scales), subcaudals 78; (4) loreal single, elongated, in contact with eye or not, not in contact with internasals; (5) a single preocular not in contact with frontal, supraocular in contact with prefrontal, two postoculars; (6) maxillary teeth 10 (4+2+2+2); (7) two anterior temporals, three posterior temporals; (8) precloacal plate entire; (9) ground color from head to tail brownish black, with 31–35 dusty rose bands on body trunk, 13–16 on tail; (10) bands in 1–2 vertebral scales broad in minimum width; (11) bands separate ground color into brownish black ellipse patches arranged in a row along the top of body and tail; (12) elliptical patches in 3–6 scales of the vertebral row in maximum width; (13) ventral surface of body with wide brownish black strip, margined with a pair of continuous narrow greyish white ventrolateral lines. With the description of the new species, 64 congeners are currently known in the genus Lycodon, with 16 species occurring in China.
Colubrinae, Guangxi, Lycodon cathaya sp. nov., morphology, phylogeny, taxonomy
The colubrid genus Lycodon Boie, 1827 currently comprises 63 known species, and is distributed widely throughout the Middle East to Southeast Asia, as well as to the Indo-Australian Archipelago (
During recent herpetological surveys in Guangxi, southern China, two colubrid snake specimens were collected from Huaping Nature Reserve (Fig.
Morphological examinations were performed based on two specimens collected from Huaping Nature Reserve, Guangxi, China. All specimens were fixed in 10 % buffered formalin and later transferred to 70 % ethanol for permanent preservation, and deposited in the Museum of Biology, Sun Yat-sen University (
Morphological descriptions followed
ED eye horizontal diameter;
HL head length (from tip of snout to posterior margin of the mandible);
HW maximum head width;
SVL snout-vent length (from tip of snout to posterior margin of cloacal plate);
TaL tail length (from posterior margin of cloacal plate to tip of tail);
TL total length (from tip of snout to tip of tail).
Scalation features and their abbreviations are as follows: dorsal scale rows (DSR) counted at one head length behind head, at midbody, and at one head length before vent, respectively; supralabials (SPL); numbers of supralabials in contact with the eye (SPL-E); infralabials (IFL); chin shields (CS); numbers of infralabials in contact with the anterior chin shield (IFL-aCS); number of infralabials in contact with the posterior chin shield (IFL-pCS); preoculars (PrO); postoculars (PtO); loreal (LoR); loreal in contact with the eye or not (L-E); anterior temporals (aTMP); posterior temporals (pTMP); preventral scales (PrV); ventral scales (V); precloacal plate (PrC); subcaudals (SC); and body scale surface (BSC). Sex was determined by dissection or by the presence/absence of everted hemipenis. The number of maxillary teeth (MT) were counted by carefully dissecting the gums of the right maxilla under the stereo microscope. The light bands on the body and tail were counted on one side; hardly visible or incomplete bands were counted as one band; obviously fused bands were counted as two bands. The collar band on the neck was not included in counts and bands covering the cloacal plate were regarded as body bands.
Morphological characters of recognized Lycodon species were obtained from examination of museum specimens (see Appendix
Selected morphological characters of Lycodon species for comparison (after
Lycodon species | cathaya sp. nov. | albofuscus | alcalai | anamallensis | aulicus | banksi | bibonius | butleri | capucinus | cardamomensis | carinatus |
DSR | 17–17–15 | ?–17–? | 19–17–15 | 17–17–15 | 17–17–15 | 17–17–15 | 19–17–15 | ?–17/19–? | 17–17–15 | 19–17–15 | 17/19–19–17 |
MT | 10 | 12 | 11–13 | ? | ? | ? | 11–14 | ? | 15 | 10–12 | ? |
SPL | 8 | 8 | 9 | 9 | 8–10 | 8 | 7–9 | 8–9 | 9–10 | 9–10 | 8–9 |
SPL-E | 3rd–5th | 3rd–5th | 4th–5th | 3rd–5th | 3rd–5th | 3rd–5th | 3rd/4th–5th | 3rd–5th | 3rd–5th | 3rd–5th | 3rd–5th |
IFL | 9 | ? | 10 | 10–11 | 10–11 | 10 | 9–10 | 9–10 | 9–10 | 9–10 | ? |
PrO | 1 | 1 | 2 | 1 | 1 | 1 | 2 | 1 | 1 | 1 | 1 |
PtO | 2 | 2 | 3 | 2–3 | 2 | 2 | 2–3 | 2 | 2 | 2–3 | 2 |
Loreal | 1 | 1 | 1 | 1–2 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
L-E | yes/no | no | no | ? | no | no | no | yes | no | no | no |
aTMP | 2 | 2 | 2 | 2 | 2 | 2 | 2+3 | 2 | 2 | 2 | 2 |
pTMP | 3 | 2 | 3 | 3+4 | 3 | 3 | 2+3+3 | 2 | 3 | 2–3 | 2–3 |
V | 199–200 | 241 | 203–207 | 174–204 | 180–215 | 241 | 204–212 | 220–227 | 182–211 | 215–228 | 185–202 |
SC | 78 | 155–208 | 108–126 | 60–74 | 57–78 | 26 (broken tail) | 110–120 | 81–96 | 59–74 | 87–93 | 51–64 |
PrC | entire | divided | entire | entire /divided | divided | entire | entire | entire | divided | entire | entire |
BSC | smooth | keeled | smooth | smooth | smooth and glossy | smooth (six central DSR of posterior 1/3 feebly keeled) | smooth | keeled | weakly keeled | weakly keeled | strongly keeled |
Lycodon species | cavernicolus | chrysoprateros | davidi | davisonii | dumerilii | effraenis | fasciatus | fausti | ferroni | flavicollis | flavomaculatus |
DSR | 17–17–15 | 19–17–15 | 17–17–15 | ?–13–? | 19–17–15 | ?–17–? | 17–17–15 | 19–17–15 | 19–17–15 | 17–17–15 | 17–17–15 |
MT | ? | 11–13 | 11 | ? | 13–15 | ? | 11 | 13 | 12 | ? | ? |
SPL | 9–10 | 9 | 8 | 7 | 11–13 | 9 | 8 | 9 | 10 | 9 | 9 |
SPL-E | 4th–6th | 3rd–5th | 3rd–5th | 3rd–4th | 4th–5th | 3rd–5th | 3rd–5th | 4th–5th | 4th–6th | 3rd–5th | 3rd–5th |
IFL | 10–11 | 10 | 10 | 8 | 9–10 | 10–11 | 8–10 | 9–10 | 10 | 11 | 10 |
PrO | 1 | 2 | 1 | 1 | 1–2 | 1 | 1 | 2 | 2 | 1 | 1 |
PtO | 2 | 2–3 | 2 | 2–3 | 2 | 2–3 | 2 | 3 | 2 | 2 | 2 |
Loreal | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 |
L-E | yes | no | no | yes | yes/no | no LoR | yes | ? | no | no | no |
aTMP | 2–3 | 2+3+4 | 2 | 1–2 | 2 | 2 | 2 | 2 | 2 | 2–3 | 1–2 |
pTMP | 3–4 | 2+3+4 | 2–3 | 2 | 3 | 2–3 | 2 | 2–3 | 3+4 | 3 (rarely 2) | 3 (rarely 2) |
V | 232–245 | 186–194 | 224 | 233–265 | 195–221 | 215–228 | 182–225 | 207–215 | 203 | 210–224 | 165–183 |
SC | 92–113 | 111–117 | 99 | 90–108 | 111–120 | 72–99 | 65–94 | 135–148 | 109 | 65–72 | 53–63 |
PrC | entire | entire | entire | entire | entire | entire | entire | entire | entire | divided | divided |
BSC | the 8 medial rows weakly keeled | smooth | middorsal scale rows slightly keeled | smooth | ? | smooth | keeled | smooth | smooth | smooth with single apical pit | smooth |
Lycodon species | flavozonatus | futsingensis | gammiei | gibsonae | gongshan | gracilis | hypsirhinoides | jara | kundui | laoensis | liuchengchaoi |
DSR | 17–17–15 | 17–16/17–15 | 17–17/19–15 | 17–17–15 | 17–17–15 | ?–15–? | 17–17–15 | 17–17–15 | 15–15–15 | 17–17–15 | 17–17–15 |
MT | 13 | 12–15 | ? | 13 | ? | 9 | ? | ? | ? | ? | 8–9 |
SPL | 8 | 7–8 | 7–9 | 8 | 8 | 8 | 9 | 8–9 | 7 | 9–10 | 7–8 |
SPL-E | 3rd–5th | 2–4; 3/4–5; 4–6 | 3rd–4th/5th | 3rd–4th/5th | 3rd–5th | 3rd–4th | 3rd–5th | 3rd–5th | 3rd–4th | 3rd–5th | 3rd–5th |
IFL | 10 | 9–11 | ? | 10 | 8 | ? | 10 | ? | ? | 10 | 7–9 |
PrO | 1 | 1 | 1 | 1 | 1 | 2 | 1 | 1 | ? | 1 | 1 |
PtO | 2 | 2–3 | 1–2 | 2 | 2 | 2 | 2 | 2 | 2 | 2–3 | 2 |
Loreal | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
LoR-E | no | no | no | yes | yes | yes | no | no | no | no | yes |
aTMP | 2 | 1–2 | 2 or irregular | 2 | 2 | 2 | 2 | 1–2 | 1 | 2 | 1–3 |
pTMP | 2–3 | 2–3 | 2 or irregular | 3 | 2–3 | 3 | 3 | 2–3 | 2 | 3 | 1–3 |
V | 211–221 | 193–208 | 205–220 | 223–226 | 210–216 | 234 | 188–210 | 167–188 | 186 | 163–192 | 190–228 |
SC | 80–88 | 72–87 | 98–111 | 91–92 | 92–96 | 81–83 | 61–75 | 52–74 | 70 | 60–76 | 68–75 |
PrC | entire /divided | entire | entire | entire | entire | entire | divided | divided | entire /divided | divided | divided |
BSC | the 7 medial rows feebly keeled | smooth | the 9 medial rows keeled | upper 3 or 4 rows keeled | the 7–13 medial rows keeled | keeled | smooth | smooth | smooth | smooth | feebly keeled in median rows |
Lycodon species | mackinnoni | meridionalis | muelleri | multifasciatus | multizonatus | namdongensis | nympha | ophiophagus | orientalis | paucifasciatus | philippinus |
DSR | 17–17–15 | 17–17–15 | 19–17–15 | 17–17–? | 17–17–15 | 17–17–15 | 13–13–13 | 17–17–15 | ?–17–? | 19–17/19–15 | ?–15–? |
MT | ? | 11 | 14–15 | ? | 10–11 | 12 | 8–10 | 11–13 | 10–11 | 11–12 | 8 |
SPL | 7–8 | 8 | 9 | ? | 7–8 | 8 | 6–8 | 8 | 8 | 8 | 7 |
SPL-E | 3rd–5th | 3rd–5th | 4th–5th | ? | 3rd–5th | 3rd–5th | 3rd–4th | 3rd–5th | 3rd–5th | 3rd–5th | 3rd–4th |
IFL | 8 | 10 | 10 | ? | 7–8 | 10 | ? | 10 | ? | 10 | 7 |
PrO | 1 | 1 | 1–2 | ? | 0–1 | 1–2 | 1–2 | 1 | 0 | 1 | 0–1 |
PtO | 2 | 2 | 2–3 | ? | 2 | 3 | 2 | 2 | 2 | 2 | 2–3 |
Loreal | 0–1 | 1 | 1 | ? | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
L-E | no | no | no | no | yes | no | yes | no | yes | no | yes |
aTMP | 1–2 | 2 | 2 | ? | 1–2 | 2 | 2 | 2 | 2 | 2 | 2 |
pTMP | 2–3 | 3 | 3+4 | ? | 2–3 | 2 | 2–3 | 3 | 3 | 3 | 3 |
V | 163–187 | 227–240 | 205–213 | 229–237 | 190–195 | 218 | 200–243 | 211–212 | 200–208 | 219–222 | 216–225 |
SC | 48–56 | 96–106 | 112–117 | 106–119 | 68–75 | 85 | 65–88 | 87–90 | 68–74 | 90–92 | 87–99 |
PrC | divided | divided | entire | ? | divided | entire | divided | entire | divided | entire | entire |
BSC | smooth | the 10–12 medial rows feebly keeled | ? | keeled | smooth | smooth | keeled | smooth | scales with a very faint keel along their anterior half | the 3–5 medial rows distinctly keeled | smooth |
Lycodon species | pictus | rosozonatus | rufozonatus | ruhstrati ruhstrati | ruhstrati abditus | sealei | semicarinatus | septentrionalis | sidiki | solivagus | stormi |
DSR | 17–17–15 | 19–19–15/17 | 17/19–17–15 | 17–17–15 | ?–17–? | ?–17–? | 17–17–15 | 17–17–15 | 19–17–15 | ?–19–? | |
MT | 13–14 | 12–13 | 11–13 | ? | 11–13 | ? | ? | 7 | 7 | 11–13 | ? |
SPL | 8 | 8 | 8 | 8 | ? | 8 | 8 | 8 | 9 | 8 | |
SPL-E | 3rd–5th | ? | 3rd–5th | 3rd–5th | ? | 3rd–5th | 3rd–5th | 3rd–5th | 4th–5th | 3rd–4th | |
IFL | 10 | ? | 9–10 | 9–10 | 9–11 | ? | ? | 7–8 | 9–10 | 10 | ? |
PrO | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 2 | 1 | |
PtO | 2 | 2 | 2 | 1–2 | 2 | ? | 2 | 2 | 2 | 2–3 | 2 |
Loreal | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
L-E | yes | no | no | no | yes | no | no | yes | no | no | |
aTMP | 2 | 2 | 2 | 1–2 | ? | 2 | 2 | 2 | 2 | 1 | |
pTMP | 3 | 3 | 3 | 2–3 | ? | 3 | 3 | 2 | 3 | 3 | |
V | 212–218 | 221–234 | 184–225 | 212–228 | 197–229 | ? | 211–234 | 202–224 | 195 | 198–203 | 217 |
SC | 90–91 | ? | 53–98 | 97–114 | 90–103 | ? | 65–105 | 83–104 | 85 | 112–115 | 75 |
PrC | entire | ? | entire | entire | divided | entire | entire | divided | entire | entire | |
BSC | smooth | weakly keeled | feebly keeled in the posterior body part | the 7–13 medial rows distinctly keeled | the 5 medial rows distinctly keeled | ? | keeled along anterior half (4 outer rows smooth) | the 7/9 medial rows feebly keeled | keeled | smooth | smooth |
Lycodon species | striatus | subannulatus | subcinctus | striatus | subannulatus | subcinctus | synaptor | tessellatus | tiwarii | travancoricus | tristrigatus |
DSR | 17–17–15 | 15–15–15 | 17–17–15 | 17–17–15 | 15–15–15 | 17–17–15 | 15/17–17–15 | 17–17–15 | ?–17–15 | 17–17–15 | ?–15–? |
MT | ? | 8–10 | 8–14 | ? | 8–10 | 8–14 | 10 | ? | ? | ? | 8–10 |
SPL | 9 | 7 | 8 | 9 | 7 | 8 | 8 | 8–9 | ? | 9 | 7 |
SPL-E | 3rd–5th | 3rd–4th | 3rd–5th/6th | 3rd–5th | 3rd–4th | 3rd–5th/6th | 3rd–5th | 4th–5th | ? | 3rd–5th | 3rd–4th |
IFL | 11 | 8 | 7–8 | 11 | 8 | 7–8 | 8 | ? | ? | ? | ? |
PrO | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | ? | 1 | 0 |
PtO | 2 | 2 | 2–3 | 2 | 2 | 2–3 | 2 | 2 | ? | 2 | 2 |
Loreal | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | ? | 1 | 1 |
L-E | no | yes | yes | no | yes | yes | no | no | ? | no | yes |
aTMP | 2 (rarely 1) | 2 | 1 | 2 (rarely 1) | 2 | 1 | 2 | 2 | ? | 2–3 | 2 |
pTMP | 3 (rarely 2) | 2 | 2 | 3 (rarely 2) | 2 | 2 | 2 | 2–3 | ? | 3 | 2–3 |
V | 153–178 | 225–244 | 190–230 | 153–178 | 225–244 | 190–230 | 201–203 | 222–232 | 218–237 | 176–206 | 224 |
SC | 42–66 | 93–111 | 60–91 | 42–66 | 93–111 | 60–91 | 68–69 | 56 | 61–102 | 64–76 | 86 |
PrC | divided | entire | entire /divided | divided | entire | entire /divided | entire | divided | divided | entire | entire |
BSC | smooth | keeled | feebly keeled | smooth | keeled | feebly keeled | the 6–7 medial rows keeled | smooth | ? | smooth | keeled |
Lycodon species | striatus | subannulatus | subcinctus | synaptor | tessellatus | tiwarii | travancoricus | tristrigatus | zawi | zoosvictoriae | |
DSR | 17–17–15 | 15–15–15 | 17–17–15 | 15/17–17–15 | 17–17–15 | ?–17–15 | 17–17–15 | ?–15–? | 17–17–15 | 17–17–15 | |
MT | ? | 8–10 | 8–14 | 10 | ? | ? | ? | 8–10 | 12 | 9 | |
SPL | 9 | 7 | 8 | 8 | 8–9 | ? | 9 | 7 | 8–9 | 8 | |
SPL-E | 3rd–5th | 3rd–4th | 3rd–5th/6th | 3rd–5th | 4th–5th | ? | 3rd–5th | 3rd–4th | 3rd–5th | 3rd/4th–5th | |
IFL | 11 | 8 | 7–8 | 8 | ? | ? | ? | ? | 9–10 | 10 | |
PrO | 1 | 1 | 0 | 1 | 1 | ? | 1 | 0 | 1 | 1–2 | |
PtO | 2 | 2 | 2–3 | 2 | 2 | ? | 2 | 2 | 1–2 | 2 | |
Loreal | 1 | 1 | 1 | 1 | 1 | ? | 1 | 1 | 1 | 1 | |
L-E | no | yes | yes | no | no | ? | no | yes | no | no | |
aTMP | 2 (rarely 1) | 2 | 1 | 2 | 2 | ? | 2–3 | 2 | 2–3 | 2 | |
pTMP | 3 (rarely 2) | 2 | 2 | 2 | 2–3 | ? | 3 | 2–3 | 3–4 | 2 | |
V | 153–178 | 225–244 | 190–230 | 201–203 | 222–232 | 218–237 | 176–206 | 224 | 179–207 | 213 | |
SC | 42–66 | 93–111 | 60–91 | 68–69 | 56 | 61–102 | 64–76 | 86 | 45–75 | 85 | |
PrC | divided | entire | entire /divided | entire | divided | divided | entire | entire | divided | entire | |
BSC | smooth | keeled | feebly keeled | the 6–7 medial rows keeled | smooth | ? | smooth | keeled | smooth | weakly keeled |
For molecular analysis, a total of 20 samples was used, encompassing 18 samples from eight known Lycodon species (one sample of L. fasciatus, two samples of L. flavozonatus, four samples of L. futsingensis, two samples of L. liuchengchaoi, one sample of L. multizonatus, two samples of L. rufozonatus, four samples of L. ruhstrati, and two samples of L. subcinctus) and two samples of the unnamed species. Tissue samples were taken prior to fixation, and preserved in 99 % alcohol and stored at -40 °C.
Genomic DNA was extracted from muscle or liver tissue samples, using a DNA extraction kit from Tiangen Biotech (Beijing) Co., Ltd. A fragment of the mitochondrial cytochrome b (CYTB) gene was amplified using the primer pair L14910 (5’–GACCTGTGATMTGAAAACCAYCGTTGT-3’) and H16064 (5’– CTTTGGTTTACAAGAACAATGCTTTA-3’) following
Twenty sequences from 12 known Lycodon species and two out-group sequences Boiga cynodon (Boie, 1872) and Dasypeltis atra Sternfeld, 1912, following
Localities, voucher information, and GenBank numbers for all samples used in this study.
Lycodon species | Voucher No. | Collection locality | GenBank No. | References |
---|---|---|---|---|
(1) Lycodon cathaya sp. nov. |
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China: Huaping National NR, Longsheng County, Guangxi | MT602075 | This study |
(2) Lycodon cathaya sp. nov. |
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China: Huaping National NR, Longsheng County, Guangxi | MT602076 | This study |
(3) L. banksi | VNUF R.2015.20 | Laos: Khammouane Province | MH669272 |
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(4) L. butleri | LSUHC:8365 | Malaysia: Bukit Larut, Perak | KJ607892 |
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(5) L. butleri | LSUHC:9137 | Malaysia: Bukit Larut, Perak | KJ607891 |
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(6) L. cavernicolus | LSUHC 9985 | Malaysia: Perlis | KJ607889 |
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(7) L. cavernicolus | LSUHC 10500 | Malaysia: Perlis | KJ607890 |
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(8) L. fasciatus | CAS 234875 | Myanmar: Chin State | KC010365 |
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(9) L. fasciatus | CAS 234957 | Myanmar: Chin State | KC010366 |
|
(10) L. fasciatus |
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China: Ruili City, Yunnan | MT625862 | This study |
(11) L. flavozonatus |
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China: Bamianshan National NR, Guidong County, Hunan | MT625850 | This study |
(12) L. flavozonatus |
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China: Bamianshan National NR, Guidong County, Hunan | MT625851 | This study |
(13) L. futsingensis |
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China: Mt. Nankun, Huizhou City, Guangdong | MT625847 | This study |
(14) L. futsingensis |
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China: Shimentai National NR, Yingde City, Guangdong | MT625853 | This study |
(15) L. futsingensis |
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China: Gaoping Provincial NR, Renhua County, Guangdong | MT625857 | This study |
(16) L. futsingensis |
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China: Gaoping Provincial NR, Renhua County, Guangdong | MT625861 | This study |
(17) L. gongshan | GP 3516 | China: Lincang City, Yunnan | KP901022 | Guo et al. 2015 |
(18) L. gongshan | GP 3546 | China: Lincang City, Yunnan | KP901024 | Guo et al. 2015 |
(19) L. laoensis | FMNH 258659 | Laos: Salavan Province | KC010368 |
|
(20) L. laoensis | LSUHC 8481 | Cambodia: Pursat Province | KC010370 |
|
(21) L. liuchengchaoi |
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China: Shennongjia National NR, Hubei | MT625855 | This study |
(22) L. liuchengchaoi |
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China: Shennongjia National NR, Hubei | MT625856 | This study |
(23) L. namdongensis | VNUF R.2017.23 | Vietnam: Nam Dong Nature Reserve, Thanh Hoa | MK585007 |
|
(24) L. meridionalis | VNUF R.2017.54 | Vietnam: Ninh Binh | MH669268 |
|
(25) L. meridionalis | VNUF R.2017.88 | Vietnam: Ninh Binh | MH669269 |
|
(26) L. multizonatus | KIZ01623 | China: Luding County, Sichuan | KF732926 |
|
(27) L. multizonatus |
|
China: Baishuijiang National NR, Longnan City, Gansu | MT625863 | This study |
(28) L. pictus | ZFMK93746 | Vietnam: Ha Lang District, Cao Bang | MN395829 |
|
(29) L. pictus | ZFMK93747 | Vietnam: Ha Lang District, Cao Bang | MN395830 |
|
(30) L. rufozonatus |
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China: Mt. Tiantai, Zhejiang | MT625858 | This study |
(31) L. rufozonatus |
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China: Yangjifeng National NR, Guixi City, Jiangxi | MT625860 | This study |
(32) L. ruhstrati |
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China: Shaowu Jiangshi Provincial NR, Nanping City, Fujian | MT625848 | This study |
(33) L. ruhstrati |
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China: Jiulianshan National NR, Longnan County, Jiangxi | MT625849 | This study |
(34) L. ruhstrati |
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China: Bamianshan National NR, Guidong County, Hunan | MT625852 | This study |
(35) L. ruhstrati |
|
China: Huaping National NR, Longsheng County, Guangxi | MT625854 | This study |
(36) L. semicarinatus | N/A | Japan: Ryukyu Archipelago | AB008539 | Kumazawa et al. 1996 |
(37) L. subcinctus |
|
China: Neilingding Island, Shenzhen City, Guangdong | MT625846 | This study |
(38) L. subcinctus |
|
China: Shimentai National NR, Yingde City, Guangdong | MT625859 | This study |
(39) L. synaptor | GP 3515 | China: Lincang City, Yunnan | KP901021 | Guo et al. 2015 |
(40) L. synaptor | GP 3545 | China: Lincang City, Yunnan | KP901023 | Guo et al. 2015 |
Outgroups | ||||
(41) Boiga cynodon | KU 324614 | Philippines: Negros Occidental | KC010340 |
|
(42) Dasypeltis atra | CAS 201641 | Uganda: Kabale district | AF471065 | Lawson et al. 2005 |
The CYTB nucleotide sequence matrix contained 1050 characters without insertion deletions. The MP and BI analyses produced essentially identical topologies, which were integrated in Fig.
Uncorrected p-distances among Lycodon species based on partial mitochondrial CYTB gene.
ID | Lycodon species | 1–2 | 3 | 4–5 | 6–7 | 8–10 | 11–12 | 13–16 | 17–18 | 19–20 | 21–22 | 23 | 24–25 | 26–27 | 28–29 | 30–31 | 32–35 | 36 | 37–38 | 39–40 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1–2 | Lycodon cathaya sp. nov. | 0 | ||||||||||||||||||
3 | L. banksi | 9.6 | – | |||||||||||||||||
4–5 | L. butleri | 17.3 | 20.2 | 0 | ||||||||||||||||
6–7 | L. cavernicolus | 17 | 18.7 | 9.6 | 0 | |||||||||||||||
8–10 | L. fasciatus | 12–13.7 | 14.6–16.3 | 10.4–11.5 | 9.8–10.7 | 0.7–1.8 | ||||||||||||||
11–12 | L. flavozonatus | 9.3 | 10.1 | 18 | 17.4 | 14.2–14.6 | 0 | |||||||||||||
13–16 | L. futsingensis | 8.9 | 9.5 | 16.9 | 17.1 | 14.8–15.4 | 9.2 | 0 | ||||||||||||
17–18 | L. gongshan | 14.5–14.7 | 14.9–15.1 | 8.9–9.1 | 7.6–7.7 | 7.1–8.5 | 14.1–14.3 | 14.1–14.3 | 0.1 | |||||||||||
19–20 | L. laoensis | 16.6–17.4 | 17–17.2 | 20.1–21.3 | 17.6–19.2 | 16.3–17.9 | 16.6–17.2 | 17.8–18.8 | 15.4–17 | 0.2 | ||||||||||
21–22 | L. liuchengchaoi | 16.3–16.5 | 17–17.2 | 13.7–13.8 | 13.4–13.5 | 12.3–13 | 16.3–16.5 | 14.6–14.8 | 10.1–10.5 | 18.7–20.1 | 0.1 | |||||||||
23 | L. namdongensis | 7.9 | 8.8 | 17.1 | 16.5 | 14.2–15.2 | 8 | 6.8 | 14.3–14.5 | 17.2–18.2 | 15.7–15.9 | – | ||||||||
24–25 | L. meridionalis | 7.9 | 9.6 | 17.2 | 17.4 | 13.1–13.6 | 2.7 | 8.5 | 13.7–13.9 | 15.9–16.5 | 15.3–15.5 | 8.1 | 0 | |||||||
26–27 | L. multizonatus | 14.8–15.1 | 16–16.7 | 14.2–14.5 | 15.6–15.8 | 12.7–13 | 16.1–16.5 | 14.6–15.4 | 11.9–12.1 | 18–19.4 | 6.7–7.1 | 15.2–15.9 | 15.3 | 1.6 | ||||||
28–29 | L. pictus | 14.3–14.7 | 15.7–15.9 | 14.2–14.8 | 15.3–16 | 12.8–13.6 | 13.8–14.2 | 14.9 | 12–12.5 | 17.6–18.6 | 9.5–9.7 | 14.1–14.5 | 14 | 10–10.4 | 0.6 | |||||
30–31 | L. rufozonatus | 10.7–11.2 | 12.2–12.7 | 17.1 | 17.9–18.6 | 15.2–15.9 | 8.9–9.4 | 10.1–11 | 14.5–14.8 | 17.7–18.7 | 15.9–16.7 | 10.4–10.6 | 9.4–9.6 | 15.2–15.7 | 14–15.1 | 2 | ||||
32–35 | L. ruhstrati | 14–14.4 | 15.9–16.3 | 13.4–13.6 | 12.9–13.1 | 12.2–13.2 | 13.8–14 | 15.1–15.3 | 9.7–10.3 | 16–17.6 | 11.5–11.9 | 14.9–15.3 | 13.4–13.6 | 10.6–11.1 | 10.8–11.4 | 14–14.6 | 0–0.3 | |||
36 | L. semicarinatus | 11.2 | 12.2 | 17.7 | 18.9 | 15.2–15.5 | 11.8 | 12.8 | 15.1–15.3 | 18.1–18.3 | 16.4–16.6 | 12.3 | 11.2 | 15.6 | 15.9–16.1 | 10.5–10.9 | 15–15.4 | – | ||
37–38 | L. subcinctus | 15.8 | 17.3 | 18.4 | 16.5 | 16.2–16.8 | 15.5 | 16.2 | 16.1–16.3 | 15.7–16.5 | 17.9–18.1 | 16.1 | 15.9 | 17.9–18.1 | 16.1–17.1 | 16.4–17 | 13.6–13.8 | 17.9 | 0–0.3 | |
39–40 | L. synaptor | 16.6 | 18 | 15.4 | 13 | 12.8–13 | 15.3 | 15.4 | 11.5–11.6 | 18.9–19.5 | 14–14.2 | 15.4 | 15.1 | 14.3–14.7 | 12.2–12.4 | 14.1–14.5 | 11.1–11.2 | 17.1–17.6 | 17.6 | 0 |
The phylogenetic topologies are very similar to those recovered by previous study (
Moreover, it is noteworthy that the unnamed Lycodon possesses significant morphological differences that can be easily distinguished from all other congeners (see below). Therefore, based on the combination of molecular and morphological data, we describe the unnamed population from Huaping Nature Reserve, Guangxi, southern China as a new species, Lycodon cathaya sp. nov.
General aspects in life and close-ups of body scales of A Lycodon cathaya sp. nov. (
The specific name cathaya is a noun referring to the monotypic botanic genus Cathaya Chun & Kuang, 1958. The single species C. argyrophylla Chun & Kuang, 1958 is an endangered relict plant, and was firstly discovered from Huaping Nature Reserve by the investigation team of Sun Yat-sen University. In memory of the predecessors and their contributions on the taxonomy of Chinese flora and fauna, we denominate this new snake species from Huaping Nature Reserve as Lycodon cathaya sp. nov. Its common name is suggested as “Huaping wolf snake” in English and “Hua Ping Bai Huan She (花坪白环蛇)” in Chinese.
Lycodon cathaya sp. nov. can be differentiated from its congeners by the combination of the following morphological characters: (1) dorsal scales in 17–17–15 rows, smooth throughout; (2) supralabials eight, third to fifth in contact with eye, infralabials 9; (3) ventral scales 199–200 (plus two preventral scales), subcaudals 78; (4) loreal single, elongated, in contact with eye or not, not in contact with internasals; (5) a single preocular not in contact with frontal, supraocular in contact with prefrontal, two postoculars; (6) maxillary teeth 10 (4+2+2+2); (7) two anterior temporals, three posterior temporals; (8) precloacal plate entire; (9) ground color from head to tail brownish black, with 31–35 dusty rose bands on body trunk, 13–16 on tail; (10) bands in 1–2 vertebral scales broad in minimum width; (11) bands separate ground color into brownish black ellipse patches, similar arrangement in a row along the top of body and tail; (12) elliptical patches in 3–6 scales of the vertebral row in maximum width; (13) ventral surface of body with a wide brownish black strip, margined with a pair of continuous narrow greyish white ventrolateral lines.
The detailed comparisons among all Lycodon congeners are given in Table
In our phylogenetic tree (Fig.
Lycodon cathaya sp. nov. can be further distinguished from L. ruhstrati (Figs
Lycodon cathaya sp. nov. can be significantly distinguished from L. albofuscus, L. banksi, L. butleri, L. capucinus, L. cardamomensis, L. carinatus, L. cavernicolus, L. davidi, L. fasciatus, L. flavozonatus, L. gammiei, L. gibsonae, L. gongshan, L. gracilis, L. liuchengchaoi, L. meridionalis, L. multifasciatus, L. nympha, L. orientalis, L. paucifasciatus, L. rosozonatus, L. semicarinatus, L. septentrionalis, L. sidiki, L. subannulatus, L. subcinctus, L. synaptor, L. tristrigatus and L. zoosvictoriae by its smooth dorsal scales (vs. dorsal body with keeled scales). By having dorsal scales in 17–17–15 rows, Lycodon cathaya sp. nov. can be easily distinguished from L. alcalai (DSR 19–17–15), L. bibonius (DSR 19–17–15), L. chrysoprateros (DSR 19–17–15), L. davisonii (DSR ?–13–?), L. dumerilii (DSR 19–17–15), L. fausti (DSR 19–17–15), L. ferroni (DSR ?–13–?), L. kundui (DSR 15–15–15), L. muelleri (DSR 19–17–15), L. philippinus (DSR ?–15–?), L. solivagus (DSR 19–17–15) and L. stormi (DSR ?–19–?). From the remaining 18 congeners, Lycodon cathaya sp. nov. can be easily distinguished from L. ophiophagus, L. pictus, and L. zawi by having fewer maxillary teeth; from L. anamallensis, L. effraenis, L. flavicollis, L. flavomaculatus, L. hypsirhinoides, L. laoensis, L. striatus, and L. travancoricus by having fewer supralabials; from L. anamallensis, L. aulicus, L. effraenis, L. flavicollis, L. flavomaculatus, L. hypsirhinoides, L. laoensis, L. multizonatus, L. ophiophagus, L. pictus, and L. striatus by having fewer infralabials and from L. mackinnoni by having more infralabials; from L. aulicus, L. flavicollis, L. flavomaculatus, L. hypsirhinoides, L. jara, L. laoensis, L. mackinnoni, L. multizonatus, L. sealei, L. striatus, L. tessellatus, and L. tiwarii by having an entire precloacal plate (vs. precloacal plate divided); from L. jara, L. mackinnoni, and L. striatus by having more ventrals and from L. pictus, L. tessellatus, and L. tiwarii by having fewer ventrals; from L. anamallensis, L. flavicollis, L. hypsirhinoides, L. jara, L. laoensis, L. flavomaculatus, L. mackinnoni, L. multizonatus, L. striatus, L. tessellatus and L. zawi by having more subcaudals and from L. ophiophagus and L. pictus by having fewer subcaudals; from L. effraenis and L. sealei by the presence of a single loreal (vs. loreal absent).
Adult male. Body slender, TL 562.5 mm (SVL 451.4 mm, TaL 111.1 mm, TaL/TL ratio 0.198); dorsal scales in 17–17–15 rows, smooth throughout, the vertebral scales not enlarged; head elongate, moderately distinct from neck, rather flattened, longer than wide, and narrow anteriorly, HL 17.2 mm, HW 11.1 mm (HW/HL ratio 0.643); eye large, ED 2.2 mm, pupil vertically elliptic; rostral triangular, much broader than high, barely visible from above; nostril lateral, located in the middle of nasal; nasal divided into two scales by nostril; two internasals, anteriorly rounded, almost as wide as high, bordered by two large, pentagonal prefrontals posteriorly; a single enlarged hexagonal frontal, narrowed posteriorly; parietals paired, longer than wide, in contact with each other medially, with upper anterior and posterior temporals, paraparietal laterally and four nuchal scales posteriorly; paraparietal slightly elongate, nearly rectangular; one elongated loreal on each side, in contact with eye, not in contact with internasals; one preocular located above loreal, in contact with eye and supraocular posteriorly, with prefrontal anteriorly, and not in contact with frontal; two postoculars, almost equal in length, upper one in contact with eye anteriorly, with supraocular and parietal, and with upper temporal posteriorly, lower one in contact with eye anteriorly, with anterior temporals posteriorly, and with fifth and sixth supralabials below; eight supralabials on each side, first and second in contact with nasal, third to fifth entering orbit; nine infralabials on each side, first pair in broad contact with each other, first to fourth in contact with anterior pair of chin shields, fourth to fifth in contact with posterior chin shields; two pairs of chin shields, elongate, anterior pair larger, second pair meeting in midline; two anterior temporals, almost equal in size, three posterior temporals, upper one smallest, lower one largest; 199 ventrals plus two preventrals; 78 pairs of subcaudals, excluding tail tip; precloacal plate entire.
10 (4+2+2+2) maxillary teeth on both sides, four small anterior teeth, enlarged posteriorly; two noticeably enlarged snag shaped teeth (second largest); two moderately enlarged teeth; two moderately enlarged kukri liked teeth (the anterior one larger, both with posterior cutting edges). Diastemas present between the above-mentioned maxillary teeth groups.
Hemipenis elongated, apex not fully everted after injection of formalin. Truncus bulbous, lower 1/3 smooth without spines, spine ornamentation starting at upper part with somewhat enlarged, medium sized spines. Apex with dense microspines. Sulcus spermaticus stretches to base of apex. Apex not fully everted, ending somewhat widened with an oblique opening, with microspines inside.
In life (Figs
Measurements, body proportions and scale counts of the two specimens are listed in Table
Measurements, scale counts, and body proportions of Lycodon cathaya sp. nov.
Voucher number | ||
---|---|---|
Character | 1542 | 1630 |
Age | adult | adult |
Sex | male | male |
SVL | 451.4 | 730.1 |
TaL | 111.1 | 180.5 |
TL | 562.5 | 910.6 |
TaL/TL | 0.198 | 0.198 |
HL | 17.2 | 23.3 |
HW | 11.1 | 14.6 |
HW/HL | 0.643 | 0.627 |
ED | 2.2 | 3.0 |
DSR | 17–17–15 | 17–17–15 |
SpL | 8 | 8 |
IfL | 9 | 9 |
IFL-1CS | 1st–4th | 1st–4th |
IFL-2CS | 4th–5th | 4th–5th |
CS | 2 | 2 |
V | 199 | 200 |
Sc | 78 | 78 |
S-V Bands | 35 | 31 |
TaL Bands | 16 | 13 |
MT | 10 | 10 |
Currently, Lycodon cathaya sp. nov. is only known from its type locality, Huaping Nature Reserve (Fig.
The description of Lycodon cathaya brings the total species number of this genus to 64, 16 of which occur in China. The new discovery further emphasizes the very high diversity level of the genus Lycodon (
The Huaping Nature Reserve is located in the hilly region among Guangxi, Hunan, and Guizhou. Thus, the new species is expected to occur in southwestern Hunan and southeastern Guizhou. The area within the jurisdiction of Huaping Nature Reserve has been well valued and protected by relevant local departments, with a considerable amount of research and investigation efforts having been conducted. However, further research on the true distribution, population sizes and trends, habitat conditions and conservation actions are urgently needed in the potential distribution areas outside the jurisdiction of Huaping Nature Reserve. Moreover, since the rapid and notable developments on the knowledge about the Chinese herpetofauna, the hilly regions in southern China have received more attention and a number of new species have been discovered in the recent years (
We thank Guangxi Huaping National Nature Reserve, Shi-Shi Lin, Si-Yu Zhang, Jia-He Li, and Chun-Peng Guo for their help in the field work, Yao Li and Chao-Yu Lin for their help in the lab work, and Jin-Long Ren for his kind support regarding the hemipenes description. This work was supported by the Project of Comprehensive Scientific Survey of Luoxiao Mountains Region of Ministry of Science and Technology, China (No. 2013FY111500), the Project of Scientific Investigation on the Amphibian, Reptilian and Avian Animals in Jiangxi Jiulianshan National Nature Reserve, the Project of Animal Diversity Survey and Monitoring System Construction of Guangdong Shimentai National Nature Reserve, the Project of Survey of Terrestrial Vertebrate Diversity in Guangdong Danxiashan National Nature Reserve.
Examined specimens
Lycodon flavozonatus
(N = 11): Nanling Nature Reserve, Guangdong, China:
Lycodon futsingensis
(N = 9): Nanling Nature Reserve, Guangdong, China:
Lycodon liuchengchaoi
(N = 1): Shimentai Nature Reserve, Guangdong, China:
Lycodon meridionalis
(N = 5): Heishiding Nature Reserve, Guangdong, China:
Lycodon rosozonatus
(N = 2): Jianfengling, Hainan, China:
Lycodon rufozonatus
(N = 3): Mt. Jinggang, Jiangxi, China:
Lycodon ruhstrati
(N = 7): Mt. Jiulian, Jiangxi, China:
Lycodon subcinctus
(N = 13): Sun Yet-sen University, Zhuhai, Guangdong, China: