Lycodon neomaculatus, Nguyen & Lee & Pauwels & Kennedy-Gold & Poyarkov & David & Vogel, 2024

Nguyen, Tan Van, Lee, Justin L., Pauwels, Olivier S. G., Kennedy-Gold, Stevie R., Poyarkov, Nikolay A., David, Patrick & Vogel, Gernot, 2024, The nomenclatural status of “ Anoplophallus maculatus ” Cope, 1895, its consequences on the systematics of Lycodon subcinctus auctorum, and the description of a new species (Squamata: Colubridae), Zootaxa 5519 (4), pp. 487-537 : 494-508

Lycodon neomaculatus sp. nov.

Indochinese Banded Wolf Snake

( Table 1 ; Figs. 1 , 3 , 5 , 6 , 7 (A, B), 8, 12; Appendix Tables S1 , S2, S6)

Chresonymy

Lycodon subcinctus (non Lycodon subcinctus Boie, 1827 )— Smith (1923: 202); Stejneger (1925: 90–91, in part); Smith (1943: 258, in part); Deuve (1970: 126, in part); Zhao et al. (1998: 183–184, in part); Zhao (2006: 218, in part); Wogan et al. (2008: 87); Nguyen et al. (2009: 321; in part); Geissler et al. (2011: 13); Siler et al. (2013: in part); Wallach et al. (2014: 396, in part); Chan-ard et al. (2015: in part); Vassilieva et al. (2016: 140–141, in part); Wang et al. (2020: 93, 108); Le et al. (2021: 210–201, in part); David et al. (2023: 328–335, in part); Uetz et al. (2024, page “ Lycodon subcinctus ”, in part).

Ophites subcinctus — Zhao & Adler (1993: 247, in part).

Lycodon subcinctus subcinctus — Taylor (1965: 739, in part).

Lycodon (Lycodon) subcinctus — Poyarkov et al. (2023: 341, in part).

Lycodon cf. subcinctus — Zhang et al. (2024: 81–94).

“ Anoplophallus maculatus ” (not a validly described taxon)— Cope (1895).

Lycodon subcinctus maculatus — Leviton (1955: 199, 201).

Lycodon maculatus — Liu et al. (2023), Uetz et al. (2024, page “ Lycodon maculatus ”, in part).

Holotype: MCZ R-175968 (adult male) from Chek Lap Kok Island , Hong Kong Special Administrative Region, China collected on 25 June 1990 by S.J. Karsen.

Paratypes (n = 8):— China. MCZ R-175967 (adult female) from Pak Ngan Village , Mui Wo Town, Lantau Island, Hong Kong SAR. collected on 29 June 1990 by S.J. Karsen — Myanmar. CAS 235846 (adult male) from Hepu Stream, Hepu Village , Indawgyi WS, Moenyin Township, Kachin State, Myanmar (25.0931666667N, 96.4001944444E; altitude 254 m asl.) collected on 14 May 2003 by Wogan G.O.U., Wilkinson J.A., Vindum J.V., Win H. & Thin T; CAS 229726 (adult female) from East of Thaye Chaung Town , Thaye Chaung Township, Dawei District, Tanintharyi Region (13.8671111111N, 98.2836944444E; altitude 45 m asl.) collected on 19 January 2003 by Win H., Tun H., Lwin K.S., Shein A.K. & Naing Z.M — Vietnam. DTU 552 (adult male) from Gung Re Commune , Di Linh District, Lam Dong Province (11.467058°N, 108.069084°E; altitude 1000 m asl.) collected on 5 October 2020 by T.V. Nguyen; DTU 325 (adult female) from Ia Pech Commune , Ia Glai District, Gia Lai Province (13.885446°N, 107.828661°E; altitude 585 m asl.) collected on 15 July 2021 by T.V. Nguyen; NHMW 39658 :1 (adult female) from Tam Dao NP , Vinh Phuc Province — Cambodia. RBINS 2738 (adult female, formerly under RBINS 18563 ) from Sambour District , Kratie Province (13.01917N, 105.92712E; altitude 55 m) collected on 13 May 2018.— Thailand. USNM 164427 (adult male) from Chiang Mai Province, Thailand (ca. 18.7903N, 98.9972E) collected in 1962.

Additional material (n = 12): China. MCZ R-176514–15, R-177133 (three juveniles, sex unknown) from Hong Kong SAR.— Myanmar. CAS 245971 (one juvenile, sex unknow) collected from the Tanintharyi Nature Reserve along a pipeline road near the vicinity of Khotama military camp, Yebyu Township, Dawei District, Tanintharyi Region— Vietnam. SIFASV 98 (one adult female, released) from Tan Thai Commune, Dai Tu District, Thai Nguyen Province; SIFASV 97 (one adult female, released) from Ba Na-Nui Chua NR, Da Nang City; SIFASV 99 (one adult female, released) from Pu Mat NP, Nghe An Province; SIFASV 101 (one adult female, released) from Pu Mat NP, Nghe An Province; SIFASV 102 (adult female, released) from Nui Chua NP, Ninh Thuan Province; USNM 166987 (one juvenile, sex unknown) from Bien Hoa City, Dong Nai Province; SIFASV 96 (one juvenile, sex unknown, released) from Phu Quoc NP, Kien Giang Province — Thailand. FMNH 180149 (one juvenile, male) from Nakhon Ratchasima Province collected on 6 April 1969 by W. R. Heyer; RBINS 16990 (one juvenile, sex unknown) from Sakaerat Biosphere Reserve Trail, Nakhon Ratchasima Province.

Referred material from the literature (n = 24).— Vietnam. IEBR 4757 (one adult female) from Na Hang NR, Tuyen Quang Province (see details in Le et al. 2021); IEBR A.2010.42 (one adult male), ZFMK 91899 & IEBR A.2010.43 (two juveniles, sex unknown) from Cat Tien NP, Dong Nai Province (see details in Geissler et al. 2011)— China. NHMUK 1937.2.1.4 (one juvenile, sex unknown) from Qionghai City, Hainan Province (see details in Smith 1923); AMNH R-27755 (one adult female) from Nada Town, Danzhou City, Hainan Province (see details in Pope 1935); No. 58 (one adult male) & No. 10 (one juvenile, sex unknown) from Guangxi Province, No. 732 (one adult male) & No. 197, No. 655122, No. 524 (three adult females) from Hainan Province, No. 452 & No. 2701 (two juveniles, sex unknown) from Fujian Province (see details in Zhao et al. 1998); KIZ 2023031 (one adult male) from Mengla County, Xishuangbanna Dai Autonomous Prefecture, Yunnan Province, KIZ 2023034 (one adult male) from Jinghong County, Xishuangbanna Dai Autonomous Prefecture, Yunnan Province, KIZ 2009061301 (one adult male) from Tian Town, Hechi City, Guangxi Province, KIZ 20190902 (one adult female) from Huangjiang Town, Hechi City, Guangxi Province, KIZ 2023032 (one adult male) from Luohu Town, Shenzhen City, Guangdong Province, KIZ 2023029 (one adult female) from Huangpu Town, Guangzhou City, Guangdong Province, KIZ 2023044 (one juvenile, sex unknown) from Xiegang Town, Dongguan City, Guangdong Province, CIB 9820 & CIB 78124 (two adult females) from Sanya City, Hainan Province, KIZ 2023030 (adult female) from Hanjiang Town, Putian City, Fujian Province (see details in Liu et al. 2023); KIZ 039729 (adult female) from Mengla County, Xishuangbanna Dai Autonomous Prefecture, Yunnan Province (see details in Zhang et al. 2024).

Etymology. The species nomen is composed of the prefix neo -, drawn from the classical Greek adjective νἐος (α, ν) (neos, nea, neon), meaning “new”, and the Latin adjective maculatus (- a, - um), meaning “spotted” or “blotched”, by allusion to the dorsal pattern of this species. We selected this species nomen, “the new Lycodon maculatus ”, because it describes both the long use of the combination in the literature following Cope’s (1895) misidentification, and our own correction in the present paper, from “ Lycodon maculatus ” sensu Cope (1895) to our own definition of this species.

Diagnosis. A species of the genus Lycodon characterized by the following characters: body size moderate, maximum total length up to 900 mm, slender; no preocular, both loreal and prefrontal entering orbit; dorsal scale rows 17-17-15, with the outermost rows smooth and the remaining rows keeled or feebly keeled; ventral scales 187– 208; subcaudals 68–84, paired; supralabials 8, with the 3 rd –5 th (rarely 3 rd –6 th) contacting the orbit; postoculars 2/2; temporals usually 1+2; cloacal plate divided; dorsum black or black gray, dark brown or gray brown posteriorly; 5–10 white cross-bands on the body in adults (often less in larger specimens), widely separated and relatively narrow (about 4–7 vertebral dorsal scales long); in adults, dorsal bands darken along the anterior edges of each dorsal scale, producing a speckled, reticulated or maculated pattern; large adult specimens lack body and tail bands along the posterior portion of the dorsum but maintain at least a few whitish bands anteriorly; in juveniles, 19–27 plain white body and tail bands; venter plain, cream or light gray, occasionally a mid-dorsal line present along the underside of the tail (data from Smith 1923; Pope 1935; Zhao et al. 1998; Geissler et al. 2011; Le et al. 2021; Liu et al. 2023; Zhang et al. 2024; this study).

Description of the holotype ( Fig . 5). Specimen MCZ R- 175968 in excellent condition, no incisions present along ventral surface, both hemipenes everted, but right organ more fully prepared than left. Body slender; tail long, gradually tapering to a blunt and enlarged terminal scute; head oblong shaped, longer than wide (HeadW/HeadL 0.57), slightly flattened, moderately distinct from the neck; snout elongate, projecting over the lower jaw, rounded in dorsal profile, truncate and slightly depressed in lateral profile with a weak canthus rostralis; nostrils large, positioned dorso-laterally, round in shape; eyes medium with an elliptical and slightly vertical pupil.

Measurements. SVL 456 mm, TaL 129 mm, TL 585 mm (ratio TaL/TL 0.221), HeadL 16.2 mm, HeadW 9.3 mm, SnL 5.2 mm, SnW 4.1 mm, IOD 5.3 mm, EyeD 2.5 mm, FrontalL 3.8 mm, FrontalW 3.6 mm.

Body scalation. Dorsal scale rows 17-17-15; dorsal scale rows smooth anteriorly, remaining scale rows feebly keeled at midbody and posteriorly, except for the outermost row, which is always smooth; scales on the vertebral row not enlarged; no apical pits; 192 ventral scales, laterally angulate; 72 subcaudals, paired and laterally angulate; total body scales 265; subcaudal ratio 27.17%; cloacal plate divided.

Head scalation. Rostral wider than high, barely visible from above, posterior suture not projecting into internasal region, forming a straight angle when viewed in dorsal profile; nasal 1.5 times longer than high, partially divided by a small suture above the nostril; posterior half of nasal subpentagonal, slightly larger than anterior nasal; nasal surrounded by the first two supralabials, rostral, internasal, prefrontal and loreal; internasals paired, anterior sutures slightly concave in dorsal profile, in contact with rostral, nasal, and prefrontal; each internasal 1.7 times wider than long; prefrontals paired, approximately 2.2 times longer than internasals, subrectangular, each scale 1.6 times shorter than frontal; each prefrontal 1.1 times wider than long, in contact with internasal, nasal, loreal, eye, and frontal; supraoculars paired, subrectangular shaped, 1.5 times longer than wide, in contact with prefrontal, posterior suture of each scale 1.9 times wider than anterior suture; frontal small, hexagonal, shield shaped, slightly longer than wide (ratio FrontalL/FrontalW 1.05), tapering posteriorly; anterior suture of frontal projecting past eye sockets in dorsal profile; angle formed by the posterior vertex of frontal slightly acute; parietals paired, suture 1.2 times longer than wide and 1.1 times longer than frontal, entire length of each parietal 1.7 times longer than wide; loreal 1/1, subrectangular, 1.4 times longer than high, in contact with eye; preocular absent; subocular absent; postoculars 2/2, uppermost scale approximately 1.5 times larger than lowermost; temporals 1+2; supralabials 8/8, first and second in contact with nasal, second and third in contact with loreal, third and fifth in contact with eye, sixth supralabial largest; infralabials 9/9, first pair in broad contact, first to fifth in contact with the anterior pair of chin shields; mental subtriangular in ventral profile, 2.1 times wider than long; anterior chin shields 1.4 times longer than posterior shields; posterior chin shields in narrow contact medially, separated by skin tissue along the mental groove.

Coloration in preservation. After 34 years of preservation, dorsal surface of body black anteriorly, gradually transitioning to dark brown by midbody and pale brown posteriorly, lighter along flanks; seven white body bands present, the first six with a conspicuous dark brown or black spot on the anterior end of each dorsal scale, forming a speckled pattern across every band, and the seventh band more subdued with dark brown pigment engulfing all of the dorsal scales; remaining posterior portion of the dorsum plain; tail tip cream. Dorsal surface of head dark brown from the anterior half of the frontal and supraocular scales to the snout, including the rostral, nasal, loreal and first two supralabials; remaining dorsal portion of head covered with pale brown marbling stretching from the posterior half of the supraoculars to the end of the parietals, curving in a slightly concave manner across the occipital region; within the marbling a small dark-brown spot present medially along the parietal suture; eye black, pupil white; remaining six supralabials cream; mental, first pair of infralabials and anterior edge of anterior chin shields dark brown; remaining underside of head white, immaculate. Ventral surface of head, body, tail white, and immaculate. Coloration in life not recorded.

General description and variation ( Table 1 ; Appendix Table S1 ; Figs. 1 , 5 , 6 , 7A, B , 8 ). Based on 47 specimens (12 males, 22 females, and 13 juveniles). The maximum length is 900 mm (SVL 740 mm; TaL 160 mm; based on an adult female specimen [no. 197] reported in Zhao et al. 1998: 184). The maximum length in males is 810 mm (SVL 651 mm, TaL 159 mm; specimen IEBR A.2010.42; Geissler et al. 2011: 13); ratio TaL/TL 0.178 –0.221 (mean = 0.201 ± 0.012, n = 42), without sexual dimorphism. Most specimens greatly resemble the holotype. Body slender; tail resembling the holotype, gradually tapering to a blunt and enlarged terminal scute; head oblong shaped, longer than wide, subrectangular, widest posteriorly and slightly flattened, distinct from neck; snout elongate, rounded dorsally, truncate and slightly depressed laterally, weak canthus rostralis; nostrils always pointed dorsolaterally, large, oval shaped, dividing the nasal scale anteriorly and occasionally posteriorly as well; eyes medium with an elliptical and vertical pupil.

Body scalation. Dorsal scales in 17-17-15 rows; all dorsal scale rows smooth anteriorly, first to fifth outermost rows smooth at midbody and posteriorly, remaining scales keeled; scales of the outermost dorsal scale row slightly enlarged; ventrals 187–208 (mean = 198.98 ± 4.90; n = 57), without sexual dimorphism, laterally angulate; subcaudals 68–84 (mean = 75.07±4.65, n = 30), without sexual dimorphism, all paired, laterally angulate; total body scales 260–290 (mean = 274.36 ± 7.46, n = 47); subcaudal ratio 25.09–30.21% (mean = 27.48 ± 1.26, n = 42); cloacal plate divided.

Head scalation. Arrangement of cephalic scales complete, including two internasals, two prefrontals, two supraoculars, one frontal, and two parietals, with all except the frontal arranged in pairs. Rostral wider than high, visible from above, posterior suture not projecting into internasal region or separating the internasals only a short distance medially; angle formed by the posterior suture of the rostral scale and the internasals straight or broadly obtuse angled; nasal partially divided or completely divided into two distinct scales by a suture above or below the nostril; anterior half subrectangular, posterior half pentagonal and distinctly larger; internasals subrectangular, not narrowing anteriorly, wider than long, separate from loreal; prefrontals subrectangular, larger than internasals, in broad contact with loreal, in contact with orbit; supraoculars subrectangular, posterior end of each scale slightly wider than anterior end; frontal hexagonal or pentagonal, shield shaped; angle formed by the posterior vertex of frontal scale acute angled; anterior edge of frontal scale positioned in front of eye sockets; parietals paired, each scale and suture larger and longer than frontal; loreal longer than high, subrectangular, in broad contact with nasal and prefrontal; loreal in contact with orbit; supralabials normally 8/8, 1 st –2 nd supralabials in contact with nasals, 2 nd –3 rd supralabials in contact with loreal, 3 rd –5 th (45/ 49 specimens) or rarely 3 rd –6 th (4/ 49 specimens) entering orbit, 6 th supralabial largest; preocular absent; postoculars two, uppermost scalelargest; temporals 1+2 (rarely 1+1); infralabials 9/9 (42/ 49 specimens), rarely 8/8 (7/ 49 specimens), 2 nd –3 rd infralabials small, first pair larger and in contact with each other; 1 st –4 th (40/ 49 specimens) or rarely 1 st –5 th (9/ 49 specimens) infralabials in contact with anterior chin shields, 5 th and 6 th infralabials greatly enlarged; mental triangular, small; anterior chin shields as long as or slightly longer than posterior chin shields, paired.

Dentition. Based on four specimens, namely CAS 229726 (paratype), CAS 235846 (paratype), CAS 245971 (paratype), and FMNH 180149 (additional voucher specimen). Total of 12–14 maxillary teeth on each upper jaw, with 8–9 small anterior teeth + a wide diastema, longer than the largest teeth + 5–6 moderately enlarged posterior teeth, without gap, last one very large. The maxilla is strongly arched and distinctly bent inwards anteriorly.

Hemipenial morphology. Based on three specimens (MCZ R-175968 (holotype) see Fig. 5F ; CAS 235846 (paratype) see Fig. 6F , and USMN 7339 fide Cope 1895). The hemipenis is elongate and unilobed. The proximal half of the hemipenis is poorly ornamented with approximately three or four rows of irregularly arranged spines present in asulcate and sulcate profile. The distal half is calyculate, encircled by small, longitudinally arranged flounces each with a single dense row of small spinose calyces. The apical region of the hemipenis is entirely nude. The sulcus spermaticus is simple and arranged centripetally, sulcus lips indistinct.

Coloration ( Figs. 1 , 5 , 6 , 7A–B , 8 ). Dorsal surface black or black gray anteriorly, gradually transitioning to gray brown or pale brown at midbody and posteriorly; lighter brown along flanks; 5–10 white or cream colored body bands in adults, widely separated, about 4–7 dorsal scales long on the vertebral row, slightly expanded ventrolaterally, 7–10 dorsal scales long above the ventral plates; the first body band starts at the level of the 15 st –20 th ventral scale, and the first two to four cross-bands are separated from one another by 10–16 vertebral dorsal scales; in adults, the dorsal bands become progressively more heavily speckled with dark brown along the anterior portion of each dorsal scale until the band becomes completely subdued with dark pigment and appears indistinct relative to the rest of the dorsal surface; the posteriormost bands along the body and tail darken first, and frequently only 4–7 of the anteriormost pale bands are visible in large specimens; remaining posterior portion of the dorsum is plain in adults; tail tip is cream. Dorsal surface of the head blackish-brown or black; in some specimens the posterior half of the head beyond the frontal scale is slightly marbled with shades of brown, but in larger individuals the marbling is restricted to the posterior half of the parietals and the adjacent occipital scales; snout dark brown, with dark pigment normally invading the first two supralabials and ventral surface around the mental scale; supralabials cream or white, especially in their lower half; chin and throat plain white, sometimes pale yellow, with occasional dark brown spots on the first few infralabials; eye brown in life, pupil black. Venter white, cream or pale yellow, with a dark brown, diffuse spot of the anterior edge of each ventral scale; progressively, the ventral scales become more strongly speckled or tinted with brown at midline; in some specimens the entire ventral surface is plain cream or white. The underside of the tail is plain white but in some specimens is dark grey with white or pale yellow marbling.

Juvenile specimens are more strongly patterned than adults and are usually black dorsally with 11–19 white, uniform bands on the body, generally visible up to the vent, and 7–12 bands on the tail. The head is largely as black along the dorsal surface and on the sides of the snout, however the posterior portion of the head has a white facial collar present across the entirety of the parietals, the posterior half of the frontal, most of the temporals and the remaining occipital region and nape; supralabials white or cream, occasionally with dark gray mottling near the snout and eyes. The ventral surface is uniform cream or white.

Comparisons. Lycodon neomaculatus sp. nov. can be easily distinguished from all other known Lycodon species (except L. subcinctus , L. sealei , L. sidiki Wostl, Hamidy, Kurniawan & Smith ) by the lack of a preocular, and by having both the loreal and prefrontal scales enter the orbit. We summarize the main characters separating Lycodon neomaculatus sp. nov. from L. subcinctus and L. sealei in Table 1 .

Lycodon neomaculatus sp. nov. differs from L. subcinctus by having: a lower number of ventrals (VEN 187– 208 [mean = 198.98 ± 4.90] vs. 201–224 [mean = 212.59 ± 5.50]), a lower number of total body scales (TOTAL 260–290 [mean = 274.36 ± 7.46] vs. 274–311 [mean = 289.81 ± 7.29]), relative size of the largest supralabial scale (sixth supralabial largest vs. seventh supralabial largest), and by the pattern of pigmentation on the white bands in adults (white bands darkening along the anterior edge of each dorsal scale, creating a speckled, reticulate or ‘maculate’ patterning vs. white bands darkening along the entire dorsal scale, pattern more smeared, patched or ‘piebald’ in appearance; see Figs. 7 , 9 ). In addition, some adult L. subcinctus (and L. sealei ) specimens may become completely melanistic, whereas all adult L. neomaculatus sp. nov. we have examined retain light pigment along the dorsal bands and are never fully melanistic.

Lycodon neomaculatus sp. nov. differs from L. sealei by having a higher relative tail length (TaL/TL 0.178 – 0.221 [mean = 0.201 ± 0.012] vs. 0.154 –0.184 [mean = 0.168 ± 0.009]), a higher number of subcaudals (SC 68–84 [mean = 75.07 ± 4.65] vs. 59–69 [mean = 63.58 ± 5.53]), a higher subcaudal ratio (SCR 25.09–30.21 [mean = 27.48 ± 1.26] vs. 22.61–25.27 [mean = 23.70 ± 0.08]), relative size of the largest supralabial scale (sixth supralabial largest vs. seventh supralabial largest), and a higher number of dorsal body bands in juvenile specimens (BB+TB 19–27 vs. 8–12; see Fig. 10 ). The same color pattern characteristics that differentiate L. neomaculatus sp. nov. from L. subcinctus s. str. also apply to L. sealei . Moreover, the geographic ranges of both species are separated by that of L. subcinctus s. str. (see Figs. 11 , 12 ).

In addition, L. neomaculatus sp. nov. differs from L. sidiki by having the prefrontal enter the orbit (vs. not entering); temporals 1+2 (vs. 2+3); cloacal plate divided (vs. undivided); body in adults with pale and wide body bands (vs. dark and narrow crossbars) (see Wostl et al. 2017 and Nguyen et al. 2024).

Distribution ( Fig. 12 ). Based on our definition of Lycodon neomaculatus sp. nov. we establish its range as follows: China (southern), Myanmar (northern and peninsular), Vietnam (throughout), Laos (throughout), Cambodia (throughout), and Thailand (except peninsular). According to our data and an examination of records from the literature, L. neomaculatus sp. nov. inhabits the following provinces/states:— China, south-east: Hong Kong SAR.; Fujian, Guangdong, Fujian, Guangxi, Yunnan, and Hainan provinces;— Myanmar: northern and peninsular: Kachin State; Mandalay, and Tanintharyi Regions);— Vietnam: Ha Noi capital; Da Nang City; Bac Kan, Tuyen Quang, Thai Nguyen, Vinh Phuc, Lai Chau, Son La, Thanh Hoa, Ninh Binh, Nghe An, Quang Tri, Thua Thien-Hue, Ha Tinh, Quang Nam, Kon Tum, Quang Ngai, Gia Lai, Dak Lak, Phu Yen, Khanh Hoa, Ninh Thuan, Lam Dong, Dak Nong, Binh Phuoc, Dong Nai, Ba Ria-Vung Tau, Tay Ninh, and Kien Giang provinces;— Laos: Bokeo, Luang Prabang, Vientiane, Bolikhamxay, Khammouane, and Champasak provinces;— Cambodia: Siem Reap, Kratie, Mondulkiri, Koh Kong, Sihanoukville, and Kep provinces;— Thailand: Chiang Rai, Chiang Mai, Phrae, Phitsanulok, Loei, Nong Khai, Udon Thani, Kalasin, Nakhon Ratchasima, Nakhon Nayok, Sa Kaeo, Chachoengsao, Rayong, Chanthaburi, Uthai Thani, Phetchaburi, and Prachuap Khiri Khan provinces (based on Pope 1935; Zhao et al. 1998; Le et al. 2021; David et al. 2023; Liu et al. 2023; examined specimens;records from iNaturalist).

Natural history. Lycodon neomaculatus sp. nov. inhabits regions covered with primary and secondary rainforests, tropical evergreen forests and adjacent lowlands between 150–800 m asl. ( Pope 1935; Liu et al. 2023; this study). This species may also be found in disturbed moist forests, thickets, tangled vegetation, and near human habitats such as shrublands, plantations, rice fields, abandoned structures, edges of cultivated areas, and villages ( Zhao 2006; David et al. 2023; Liu et al. 2023; this study). This species is chiefly nocturnal (active between 1930– 2330 hrs) but can also be active in the early morning. It is mainly terrestrial and is frequently found crawling on the forest floor or climbing in the forest understory. During the day, it retreats under stones, stumps, pieces of wood, and vegetation. This species can also be arboreal, sometimes ascending to the crowns of large trees, such as rubber and coffee trees (T.V. Nguyen, pers. obs.). There have been no reports of its reproduction, but like other Lycodon , it is presumably oviparous. Lycodon neomaculatus sp. nov. is known to eat amphibians and lizards specifically skinks like Eutropis multifasciata (Kuhl) (see Pope, 1935), Ateuchosaurus chinensis Gray (see https://www.inaturalist.org/ observations/14752734), Tropidophorus microlepis Günther (https://www.inaturalist.org/observations/214567395), and possibly small snakes and small birds.

Conservation status. According to the IUCN Red List, both Lycodon subcinctus and L. sealei are listed as Species of Least Concern (LC) due to their wide geographic distributions and ecological adaptability, presence in several large, protected areas, and slight adaptability to habitat degradation ( Kim & Schoppe 2023; Lilley et al. 2023). Lycodon neomaculatus sp. nov. is also distributed across in China and Indochina and inhabits several national parks and other protected areas but does not appear to be abundant wherever it occurs. No major threats are known for this species; however, some local populations could be threatened by deforestation and other forms of habitat loss. Many species of wolf snakes, including Lycodon davisonii Blanford , L. laoensis Günther , and Lycodon neomaculatus sp. nov. resemble venomous elapid snakes, such as kraits in the genus Bungarus (including Bungarus candidus (Linnaeus) , B. multicinctus Blyth , B. sagittatus Aksornneam, Rujirawan, Yodthong, Sung & Aowphol , B. suzhenae Chen, Shi, Vogel, Ding & Shi , and B. wanghaotingi Pope ) and often live in areas where humans co-occur. Consequently, wolf snakes, although non-venomous, are occasionally the victims of misidentifications and are needlessly killed. L. neomaculatus sp. nov. is not protected by national or international regulations and is not listed in CITES Appendices. Given this information, we suggest Lycodon neomaculatus sp. nov. should be considered Least Concern (LC) following the IUCN’s Red List categories ( IUCN Standards and Petitions Committee 2024).

Figures

FIGURE 1. Colubrid snake species involved in the case of Anoplophallus maculatus Cope, 1895.—USNM 7339, previously considered the holotype of “Lycodon maculatus” reported by Liu et al. (2023): A—General dorsal view; B—General ventral view; C—Dorsalview ofthe head;D—Lateralview of thehead,leftside; E—Livespecimen of Leptodeira maculata (= Megalops maculatus) from Minatitlán, Colima, Mexico; F—Live specimen of Lycodon subannulatus from Penang Island, Malaysia, of which Nymphophidium maculatum is a junior synonym. Images not to scale. Photos by T. Hsu (A–D), C. Grünwald (E), J. L. Lee (F).

FIGURE 3. Bayesian phylogenetic tree of Lycodon inferred from the mitochondrial cytb sequences. Numbers before slashes indicate Bayesian posterior probabilities and numbers after slashes indicate ML bootstrap supports. Nodes filled in with a black circle indicate strong support by both the ML and BI analyses, whereas nodes filled in with a white circle indicate strong support only in the BI analysis. Inset photo of Lycodon neomaculatus sp. nov. taken by A.M. Bragin.

FIGURE 5. Lycodon neomaculatus sp. nov. in preservative—MCZ R-175968 (holotype, adult male) A—General dorsal view; B—General ventral view; C—Lateral view of the head, right side; D—Lateral view of the head, left side; E—Dorsal view of the head; F—Ventral view of the tail with the everted hemipenes. Images not to scale. Photos by M. Gage and J. Martinez.

FIGURE 6. Lycodon neomaculatus sp. nov. in preservative—specimen CAS 235846 (paratype, adult male) A—General dorsal view; B—General ventral view; C—dorsal view of the head; D—Ventral view of the head; E—Lateral view of the head, right side; F—Ventral view of the tail with the everted hemipenes. Photos by J.L. Lee.

FIGURE 7. Differences in dorsal body band coloration in preservative between Lycodon neomaculatus sp. nov. (A, B) and L. subcinctus (C, D). A—FMNH 180149 from Nakhon Ratchasima, Thailand; B—CAS 229729 from Tanintharyi, Myanmar; C—FMNH 229855 from Selangor, Malaysia; D—FMNH 183772 from Selangor, Malaysia. Images not to scale. Photos by J.L. Lee.

FIGURE 8. Lycodon neomaculatus sp. nov. in life (not collected, sex not determined): A, B—Hong Kong, China (adult, subadult; respectively); C—Qiongzhong, Hainan, China (adult); D—Xishuangbanna, Yunnan, China; E—Pu Mat NP, Nghe An, Vietnam; F—Ma Da NR, Dong Nai, Vietnam (adult); G—Cat Tien NP, Dong Nai, Vietnam (subadult); H—Loc Bac, Lam Dong, Vietnam (adult); I—Phuoc Binh NP, Ninh Thuan, Vietnam (adult); J—Hinhurp, Vientiane, Laos (subadult); K—Kompong Seila, Seihanuk, Cambodia (adult); L—Chai Prakarn, Chiang Mai, Thailand (subadult); M—Mae Yom NP, Phrae, Thailand (adult); N—Wang Nam Khiao, Nakhon Ratchasima, Thailand (adult); O—Kaeng Krachan NP, Phetchaburi, Thailand (subadult). Photos by:A. Tomaszek (A, B); C.X. Liao (C); Z.H. Ma (D); P.S. Hamilton (E); H.X.N. Nguyen (F); A.M. Bragin (G, I); N.A. Poyarkov (H); T. Calame (J); T. Samnang (K); A. Kaosung (L); R. Grassby-Lewis (M, N); A. Pierce (O).

FIGURE 9. Lycodon subcinctus in life (not collected, sex not determined): A—Kathu, Phuket, Thailand (adult); B—Hala Bala WS, Narathiwat, Thailand (subadult); C—Khao Phanom Bencha NP, Krabi, Thailand (subadult); D—Khao Sok NP, Surat Thani, Thailand (adult); E—Baling, Kedah, Malaysia (adult); F—Fraser’s Hill, Pahang, Malaysia (subadult); G—West Malaysia (adult); H—Upper Seletar Reservoir, Singapore (adult); I—Padang, West Sumatra, Indonesia (adult); J—Bekasi, Java, Indonesia (adult); K—Jembrana, Bali, Indonesia (adult); L—Alor, East Nusa Tenggara, Indonesia (subadult). Photos by: A. Tomaszek (A, K); R. Grassby-Lewis (B); R. Jaihan (C, D), P. Pawangkhanant (E, I); G. Vogel (G), R. Kittrell (F); R. Tann (H); A.R. Mudzakir (J); C. Trainor (L).

FIGURE 10. Lycodon sealei in life (not collected, sex not determined): A—Puerto Princesca, Palawan, Philippines (adult); B—Taytay, Palawan, Philippines (subadult); C—Balabac, Palawan, Philippines (adult); D—Ranau, Sabah, Malaysia (subadult); E—Marudi, Sarawak, Malaysia (subadult); F—Sandakan, Sabah, Malaysia (subadult); G—Kuching, Sarawak, Malaysia (adult); H—Petra Jaya, Kuching, Sarawak, Malaysia (adult); I—West Kotawaringin, Central Kalimantan, Indonesia (adult). Photos by: R. Grassby-Lewis (A); A. Sonlee (B); C. Supsup (C); R. Monday (D); C.C. Lee (E, H); S. Lee (F); L. Platania (G); H. Kurniawan (I).

FIGURE 11. Distribution ranges of Lycodon sealei, L. subcinctus, and L. cf. subcinctus (Nicobar Island). Notes: numbers indicate different localities where the species have been recorded (see Appendix Table S5 for the details of localities).

FIGURE 12. Distribution ranges of Lycodon neomaculatus sp. nov. Numbers indicate various localities where the species have been recorded (see Appendix Table S6 for the details of localities).