Lycodon gongshan Vogel & Luo, 2011

Nguyen, Tan Van, Lee, Justin L., Jiang, Ke, Ding, Li, Chit, May Thu, Poyarkov, Nikolay A. & Vogel, Gernot, 2025, A new species of wolf snake Lycodon Fitzinger, 1826 from China and Myanmar (Squamata: Colubridae), and new data on Lycodon gongshan Vogel & Luo, 2011, Zootaxa 5621 (1), pp. 1-51 : 21-26

Redescription of Lycodon gongshan Vogel & Luo, 2011

( Figs. 5G–I , 6E–F , 7C , 8 , Appendix Fig. S2 ; Table 1 , Appendix Table S4)

Referred specimens (n = 13). China: Sichuan Province — CIB DL.055 (subadult male; formerly R20140901 in Chen et al. 2018) from the Daheishan Forest Park , Panzhihua City (26.640800ºN, 101.695200ºE; altitude 2173 meters asl.) collected by L. Ding ; CIB PZH.2017.09.01 (adult male) from Hongbao Miao and Yi Township , Yanbian County, Panzhihua City (27.125400ºN, 101.491900ºE; altitude 1517 meters asl.) ; Xizang AR—CIB 7557 (adult male) from Chawalong Township , Zayu County (28.485833ºN, 98.455833ºE; altitude 1981 meters asl.) ; Yunnan Province — CAS 241963 (adult male; field no. GLGS-2992) ca. 3 km N of hotel ( Kongdang ) on Dulong Road , Kongdang Village, Gongshan County (27.896833ºN, 98.338667ºE; altitude 1540 meters asl.) collected on 6 Jul 2005 by D.-Q. Rao, M.W. Zhang, J.V. Vindum, and J.A. Wilkinson ; CAS 242669 (subadult female) from N of Kongdang, Kongdang-Gougshan road, Dulong Valley , Gongshan County (27.881250ºN, 98.342889ºE; altitude 1600 meters asl.) collected on 7 Sep 2007 by M.W. Zhang, J.V. Vindum and J.A. Wilkinson; CIB YN.2019.09.189 and CIB YN.2019.09.286 (both adult males) from Luzhang Town, Lushui City (25.951389ºN, 98.768611ºE; altitude 2006 meters asl.) collected by S.-C. Shi (see details in Huang et al. 2021) ; HNU 200505002 (adult female) from Xiaoheishan, Longjiang Township , Longling County (Baoshan City) collected by Y. Hengmei, L. Hongbin and G. Keji (paratype) ; HNU 200609001 (adult male) from Dulongjiang Township , Gongshan County collected by Q. Hou (paratype) ; KIZ 730034 (adult male) , KIZ 35112 (adult female) and KIZ 730008 (immature male) from “ Bapo , Gongshan , China ” [= Dulongjiang Township , Gongshan County], collectors unknown (holotype and paratypes, respectively; see details in Vogel and Luo 2011) . Myanmar: Kachin State — CAS 245470 (subadult male; field no. CAS-MHS-29011) from Lukpwi Village , Chipwe Township , Myitkyina District, Kachin State (25.701333ºN, 98.336083ºE; altitude 1845 meters asl.) collected on 1 Aug 2009 by M. Hlaing, S.L. Oo, Z.H. Aung, K.S. Lwin, and Y.M. Win .

Revised diagnosis. A large-sized Lycodon with a maximum total length of 1,005 mm (longest specimen without broken tail 963 mm); relative tail length 0.21–0.27; dorsal scale rows 17–17–15, first 5–9 upper dorsal scale rows keeled at midbody and posteriorly, remaining rows smooth; ventrals 203–216; subcaudals 79–96, paired; cloacal plate undivided; supralabials usually eight (rarely nine) with third to fifth scales contacting eye; infralabials usually eight (rarely nine), the first four (rarely five) in contact with anterior pair of chin shields; preocular 1/1, blocking prefrontals from contacting eye; loreal usually in narrow contact with eye; postoculars 2/2; temporals 2+2 (rarely 2+3). Dorsum variable, typical morph black in life with 26–46 orange brown or reddish brown light crossbands, 9–16 tail crossbands; first dorsal band starting between ventrals 4–9, 3–6 ventral scales long at base and 1–2 vertebral dorsal scales long; head usually plain black dorsally, occasionally indistinct light brown pigment present on supralabials below eye; underside of head black along chin; throat usually also black but occasionally plain white; venter with discrete light and dark crossbands throughout, spotting rarely present. Second color morph (known only from Zayu County, Xizang AR) olive brown dorsally with frequent yellow or light brown mottling along edges of dorsal scales and 67 indistinct shaped black body crossbands and 28 black tail crossbands; head dark brown dorsally, supralabials beige, underside of head black along chin but throat light cream; ventral surface with array of irregular and subdued dark crossbands and midventral spotting (based on Vogel & Luo 2011; Chen et al. 2018; Huang et al. 2021; and this study).

General description and variation. Based on thirteen specimens (including the type series; holotype + three paratypes). The longest specimen is an adult female (KIZ 35112; SVL 798 mm, TaL 207 mm but partially broken). The longest male specimen is the holotype (KIZ 730034; SVL 740 mm, TaL 223 mm), which also happens to be the longest specimen with a complete tail. Ratio TaL/TL 0.21–0.27 (x̄ = 0.23±0.02; n = 10). Body elongate, tail gracile, terminal scute blunt ended. Head ovate in adults, more oblong in juveniles, distinct from neck, temporal and parietal region same height as rest of head. Snout obtuse, slightly depressed in lateral profile, no canthus rostralis. Nostrils large, rounded, positioned medially within nasal. Eyes moderate, pupil elliptical. No sexual dimorphism evident in body or head proportions.

Body scalation. Dorsal scales in 17-17-15 rows. Most specimens have entirely keeled scales except for the outermost five rows (which are smooth) at midbody and posteriorly. However , a specimen from Kachin State, Myanmar ( CAS 245470 ) has four smooth rows at midbody and two posteriorly, and one specimen from Panzhihua, Sichuan Province ( CIB DL.055) has three smooth rows posteriorly. Ventral scales 203–216 (x̄ = 210.7±3.4; n = 13), with distinct keeling along edges of each scale, less distinct in one of the Panzhihua specimens ( CIB DL.055). Subcaudals 79–96 (x̄ = 88.4±6.5; n = 10), paired. Total body scales 286–312 (x̄ = 300.1±8.9; n = 10), subcaudal ratio 0.27–0.31 (x̄ = 0.29±0.01; n = 10). Sexual dimorphism absent in all body scale counts.

Dentition. In two specimens (CAS 241963 and 245470), maxillary tooth counts range from 9–11. There are four or five anterior teeth that gradually increase in size, followed by one larger tooth and four to six smaller teeth, each separated by a diastema.

Description of hemipenis. Partially everted hemipenis of CAS 241963 is short ( Fig. 7C ), weakly bilobed, spinose. Images of the hemipenes of CIB 07557 in Huang et al. (2021) depict a weakly bilobed organ with each lobe slightly bulbous in shape. Minute spinous calyces present at the base of organ in both asulcate and sulcate views. Medially, two or three rows of slightly enlarged spines present, hook-like in shape, less prevalent on the sulcate side of organ and largely continuous with base. Remaining distal two-thirds of organ covered with small spinous and papillate shaped calyces, equally sized. Sulcus spermaticus simple, arranged centripetally. In CIB 07557 the apex is less spinose along the crotch near each lobe in sulcate view. Sulcus lips nude, broad, distinctly raised in CAS 241963 (see Fig. 7C ), but less pronounced in CIB 07557.

Head scalation. Rostral wider than high, barely visible from above. Posterior suture of rostral bordering internasals obtuse angled, ‘gull-wing’ shaped (~146–163º). Nasal scale vertically divided by indistinct suture above and below nostril, in contact with internasal, loreal, prefrontal, rostral and first two supralabials. Anterior portion of nasal smaller than posterior, each segment higher than long and subpentagonal. Internasals paired, rectangular, anterior edges concave, entire scale 1.1–1.9 times wider than long, 1.3–2.5 times wider than medial suture; each internasal contacting rostral, nasal and prefrontal. Prefrontals paired, subrectangular, area 2.6–3.6 times larger than internasals, entire scale 0.8–1.1 time longer than wide. Medial prefrontal suture usually shorter than prefrontal width (0.8–1.0 times longer than wide), roughly equal (1.0 times) in CAS 241963 and CAS 242669 ; each prefrontal contacting internasals, nasals, loreal, supraoculars, and frontal. Supraoculars paired, subrectangular, 1.6–2.0 times longer than wide, posterior end 1.3–1.9 times wider than anterior end. Frontal small, hexagonal, shield shaped, 1.1– 1.2 times longer than wide, 1.6–2.1 times longer than prefrontal suture. Anterior frontal edge positioned past eyes in all specimens; posterior vertex of frontal bordering parietals usually acute angled (~80–96º). Parietals paired, entire scale 1.8–2.3 times longer than wide, medial parietal suture 1.2–1.7 times longer than wide, its length 0.8–1.2 times as long as entire length of frontal. Anterior edge of parietal straightened, its suture with frontal and supraoculars forming an obtuse angle (~122–133º) that is, usually pointed laterally, though the angle is posterolaterally directed in CAS 242669 and KIZ 730038 . Parietals surrounded by 10–12 total scales behind upper postoculars. Loreal 1/1, subrectangular, longer than high, in narrow contact with eye on either side but blocked from contacting eye in both specimens from Panzhihua ( CIB DL.055 and PZH. 2017.09.01). Preocular 1/1, vertical, subrectangular. No presubocular or subocular. Postoculars 2/2, vertical, subrectangular, each scale approximately equal in size. Temporals normally 2+2, but 2+ 3 in three of the paratypes ( KIZ 730008 , HNU 200609001 and HNU 200505002 ), and 2+2/2+ 3 in the holotype ( KIZ 730034 ), lowermost anterior temporal largest. A single paraparietal scale present behind the posterior temporal row in all specimens (except for CAS 245470 , in which the scale is fused with the uppermost posterior temporal). Each paraparietal surrounded by 6–8 scales (6/ 6 in CAS 242669 , CIB DL.055 and PZH. 2017.09.01; 6/ 7 in CAS 245470 ; 7/ 7 in 241963; 6/ 8 in KIZ 730034 ; 8/ 8 in HNU 200609001 and KIZ 730008 ) with 3–5 scales located below them (3/ 3 in HNU 200609001 and KIZ 730008 ; 4/ 4 in CAS 241963 , CIB DL.055, CIB PZH. 2017.09.01 and HNU 2005.05.002; 5/ 5 in remaining specimens). Supralabials 8/8, first and second contacting nasal, second and third contacting loreal, and third to fifth contacting eye; sixth supralabial normally largest, but sixth and seventh scales occasionally equal in size; first supralabial smallest. Infralabials normally 8/8 (8/ 9 in CAS 245470 ; 9/8 CAS 241963 ; 9/ 9 in CIB 7557 ); infralabials 1–4 in contact with the anterior chin shields (except in CAS 245470 , which has 4/5 infralabials in contact), 5–6 contacting posterior chin shields. Mental subtriangular in ventral profile, wider than long. Anterior chin shields slightly longer or equal in size relative to posterior chin shields; 1–3 gular scales present behind posterior chin shields; usually one or two preventral scales present.

Coloration in preservation: Two color morphs are now known to exist in this species ( Huang et al. 2021). The first and typical morphotype is described first. In these specimens, the dorsal ground color is black in both juveniles and adults with 26–46 light body crossbands (x̄ = 35.9±5.6; n = 12) and 9–16 tail crossbands (x̄ = 13.1±2.8; n = 10). Anterior light body crossbands plain white anteriorly, occasionally orange brown or light brown medially, each narrower than dark bands and broadening ventrolaterally. First light crossband starts at ventral scale 4.0–9.0 (x̄ = 6.10±1.52; n = 12), 3.0–6.0 ventral scales long at base (x̄ = 3.90±0.97; n = 12) and 1.0–2.0 vertebral dorsal scales long (x̄ = 1.14±0.32; n = 12). Remaining light crossbands orange brown or light brown with plain white edges, 2.0–3.0 vertebral dorsal scales long at midbody. These crossbands may be plain white or tan in some juvenile and subadult specimens (CAS 242669 and KIZ 730038). Usually, the light crossbands are longer than dark crosbands after the 17 th to 22 nd band (CAS 241963 and KIZ 730034 and 730038), but the length of light crossbands are equal to dark bands in two specimens (in CAS 245470 and CAS 242669) and narrower than dark bands in both specimens from Panzhihua (CIB DL.055, CIB PZH. 2017.09.01). In all cases, the light crossbands broaden slightly in lateral profile and do not cause the dark crossbands to narrow significantly. Color of head black in all specimens with a broad nuchal collar present in juveniles and subadults starting from the posterior vertex of frontal and posterior edge of parietals, extending across the temporals and paraparietals, and continuing through to the last one to two supralabials/infralabials until reaching the gular region (e.g., CIB DL.055 and KIZ 730038). Color of nuchal collar plain white in juveniles, but mottled with tan or brown in subadult specimens (CAS 245470 and 242669, CIB DL.055). In adults, nuchal collar reduced to a weak white band along the lateral portion of the nape (e.g., CAS 241963), or denigrated to a small white line along the edges of the occipital scales behind the parietal (CIB YN.2019.09.189 and PZH. 2017.09.01). Bottom halves of supralabials with indistinct light brown or cream spots, usually present on scales below the eye, but never on the first supralabial. Underside of head white, black across mental, first 4–5 infralabials and first pair of anterior chin shields. Gular scales and preventrals mottled with black spots but mottling absent in CAS 241963 and 245470. Ventral surface black with plain white crossbands. First dark ventral bands usually distinct, covering 3–5 ventral scales (except in CAS 245470, where first dark ventral bands do not enter ventral surface). Remaining dark bands distinct, rectangular, 1–3 ventral scales wide, light bands 1.5–4.0 scales wide, occasional black spotting present posteriorly (e.g., in CAS 241963). Arrangement and color of bands on underside of tail same as body; occasionally, a black midventral stripe may intersect the dark crossbands (CIB YN.2019.09.189).

In the second color morph, the arrangement of the body and tail crossbands is essentially reversed. The dorsum is olive brown with 67 irregularly shaped black body crossbands and 28 crossbands on the tail. Edges of each dorsal scale with frequent yellow or light brown mottling creating a striated pattern across the body and crossbands. Head dark brown dorsally, supralabials beige, underside of head black along chin, but throat light cream. The ventral surface is plain white with an array of irregular and subdued dark gray crossbands extending from the dorsal surface. Midventrally, there are frequent black or dark gray spots that become more prominent posteriorly and continue onto the tail (description adapted from English translation of Huang et al. 2021).

Coloration in life (see Fig. 5G–I ). In life, dorsal ground color black. Light crossbands in adult specimens similar as in preservative but tend to be bolder and more vibrant. In the typical morph, light crossbands white anteriorly, with orange or brown pigment vertebrally. Light crossbands orange, orange brown, or brown at midbody and posteriorly, their edges yellow or white. Photographed adults have an entirely black head except for remnants of the nuchal collar and spotting across the infralabials and supralabials, with the infralabials and supralabial spots in CIB YN.2019.09.286 vibrant yellow. Eyes jet black with the pupil indistinct from the iris. Ventral underside white or tan, with darker crossbands usually black, but occasionally dark gray. Live coloration of the second morph similar in preservative, but more vibrant olive brown (based on Huang et al. 2021). In another specimen resembling the second morph (iNaturalist obs. 191251388, see Fig. 5I ), the dorsum is completely engulfed with light olive and yellow speckling, making it difficult to decipher the dark crossbands across the body. In the same specimen, the supralabial region of the head is black with light pigment restricted to the infralabials and supralabials immediately below the eye, similar to the color condition observed in the typical morph. Guo et al. (2015) also provided a photograph of a specimen they identified as L. gongshan but did not provide any morphological data. This specimen resembles the typical morph of L. gongshan by bearing a black dorsum with orange light crossbands but has a much more prominent white nuchal collar that extends across most of the head. Moreover, the dark crossbands in the photographed specimen become more irregularly shaped, and posteriorly are interrupted from connecting along the flanks by the light crossbands. The photographed specimen presumably makes up one of the sequenced samples Guo et al. (2015) presented in their study, but we refrain from including them in our updated species diagnosis owing to a lack of additional data presented by the authors.

Comparisons.Here, we update the comparisons between L. gongshan and L. fasciatus ,which are morphologically similar to one another. Comparisons with L. latifasciatus sp. nov. are provided in the description of that species and comparisons with other L. fasciatus group members are provided in Table 2 . With the acquisition of new specimens, some features now overlap between L. gongshan and L. fasciatus , but most are still significantly different (p <0.05). A few characters that went unnoticed by past authors may also differentiate each species. However, subcaudal counts and relative tail length can no longer be considered sexually dimorphic in L. gongshan , as unpaired t -tests recovered no statistical differences in each character between males and females (p> 0.05 in both cases).

On average, specimens of L. gongshan have a longer relative tail length than L. fasciatus (0.21–0.27 [x̄ = 0.230±0.019; n = 11] versus 0.19–0.22 [x̄ = 0.206±0.008; n = 46]; p <0.00001), a slightly higher number of ventral scales (203–216 [x̄ = 210.7±3.4; n = 13] versus 197–213 [x̄ = 206.1±4.6; n = 58]; p = 0.00269), a higher number of subcaudals (79–96 [x̄ = 88.4±6.5; n = 11] versus 74–90 [x̄ = 81.8±3.6; n = 49]; p = 0.00002), a higher number of total body scales (286–312 [300.1±8.9; n = 11] versus 278–302 [x̄ = 288.9±5.7; n = 49]; p <0.00001), and a slightly higher subcaudal ratio (0.27–0.31 [x̄ = 0.294±0.014; n = 11] versus 0.26–0.30 [x̄ = 0.283±0.011; n = 49]; p = 0.00870). In addition, L. gongshan almost always has eight infralabials and four scales contacting the anterior chin shields on at least one side of the head, whereas L. fasciatus usually have nine or eight infralabials and five scales contacting the anterior chin shields on either side of the head (p <0.00001 in both cases). The position of the first light crossband starts earlier in L. gongshan than L. fasciatus (above ventral 4.0–9.0 [x̄ = 6.10±1.52; n = 10] versus ventral 5.0–18.0 [x̄ = 10.91±3.37; n = 55]). The length of the first light crossband is also narrower at its base (3.0–6.0 ventrals [x̄ = 3.90±0.97; n = 10] versus 4.0–14.0 ventrals [x̄ = 7.69±2.56; n = 55]) and narrower vertebrally (1.0–2.0 scales in length [x̄ = 1.14±0.32; n = 11] versus 1.0–3.5 scales [x̄ = 2.29±0.69; n = 55]). All typical L. gongshan lack conspicuous supralabial spots or have indistinct and subdued light pigment on the supralabials when present, whereas a majority of L. fasciatus have conspicuous light pigment on most of the supralabials. Furthermore, in L. gongshan (including the second morphotype) the underside of the head is black with dark pigment engulfing the entire first three or four infralabials, anterior chin shields and throat, whereas in L. fasciatus dark pigment is usually restricted to the first few infralabials. Moreover, adult L. gongshan are larger in body size than L. fasciatus (max SVL 798 mm [x̄ = 606.6±126.6; n = 13] versus 680 mm [x̄ = 471.7±92.0; n = 50]; p = 0.00037). The olive brown and irregular crossbanded pattern of the second Lycodon gongshan color morph (sensu Huang et al. 2021) from Zayu County, Xizang AR, China is easily distinguishable from L. fasciatus and should not pose an issue for identification.

Distribution. Lycodon gongshan is known from multiple localities across Southwestern China and extreme Northeastern Myanmar, with most observations concentrated along the Hengduan Mountains and the Three Parallel Rivers region, although records now exist outside of these regions ( Fig. 1 , Appendix Table S4). In Yunnan Province, China, it is known from multiple locations in Nujiang Prefecture ( Gongshan County, Lushui City ) and Baoshan Prefecture (Longling County). Guo et al. (2015) also recorded specimens from Lincang City further east in Southwestern Yunnan Province and Wang et al. (2021) reported an unvouchered genetic sample from the Yunlong NR in Yunlong County, Dali Prefecture. Chen et al. (2018) recorded two specimens from Sichuan Province in Hongbao Village and Daheishan National Forest in Panzhihua City. Moreover, the newly reported color morph reported in Huang et al. (2021) extends the range of this species further north into Zayu County, Xizang Autonomous Region. A specimen of L. gongshan (CAS 245470) is also reported from Kachin State, Myanmar, constituting the first documented occurrence in the country, a distance of ~ 52.6 km SW of the nearest locality in Lushui City, Yunnan Province, China.

Natural history. Specimens of Lycodon gongshan were found at mid elevations between 1540–2173 meters in a variety of habitats. Most specimens were located in subtropical evergreen forests and montane broadleaf forests but appear to tolerate more disturbed forested habitats and agricultural edges as well ( Chen et al. 2018; Cai & Ding 2024). The specimen recorded by Huang et al. (2021) from Zayu County was found in rocky, semi-arid steppe associated with dry river canyons of the Nujiang (Salween) River. Based on field observations, this species seems to be primarily nocturnal throughout its range. Specimens from Gongshan County (CAS 241963, 242669) were found crawling through vegetation or rocky slopes along roadsides during the evening in humid conditions (19–23ºC air temperature; 78–91% humidity). Unfortunately, whether the rock substrate was limestone or granite was not recorded. The Myanmar specimen (CAS 245470) was also recorded in the evening (2053 hrs local time) during humid conditons (75.6ºF, 70% humidity). Another specimen from Lushui City was found dead on the road in the evening ( Huang et al. 2021), and both specimens from the Panzhihua City region were collected at night near roadsides ( Chen et al. 2018) prior to rain. Two lizard eggs were disgorged from the stomach of CAS 241963, and Huang et al. (2021) noted that a specimen from Lushui City was found foraging in vegetation along the sides of a creek in pursuit of a skink ( Sphenomorphus indicus (Gray)) . Nothing else is known about the natural history and basic biology of this species, however it is presumed to be oviparous like other Lycodon .

Conservation status. The most recent IUCN Red List assessment of Lycodon gongshan ( Cai & Ding 2024) lists this species as “Least Concern” and did not identify any major threats across its range. The distribution of L. gongshan has significantly expanded since its original description and is now known to occur within several protected areas (including the Gongshan and Yulong NRs). Additionally, this species seems to be tolerant of a wide variety of habitats, including those that may undergo some levels of human disturbance. Nevertheless, large scale deforestation in parts of its range (such as Myanmar) may negatively impact populations.

Remarks. The first records of L. gongshan from Sichuan Province derive from Chen et al. (2018), who reported two specimens from the surroundings of Panzhihua City (one from Daheishan Forest Park and another from Hongbao Village). Wang et al. (2021) considered the presence of L. gongshan within this region to be inconclusive and pointed out discrepancies between the morphological data in Chen et al. (2018) ’s table and the individuals photographed in the same paper. We were able to re-examine the specimens from Panzhihua City that were photographed by Chen et al. (2018: Fig. 1 ) and agree that they are L. gongshan (see Table 1 ). DNA sequences obtained from one of the two individuals (CIB PZH. 2017.09.01) adds further validation to this identification, since the sample was recovered deep in the same clade as other L. gongshan ( Fig. 2 ). Still, practically all our scale and crossband counts of the Panzhihua City specimens differ from the table in Chen et al. (2018), confirming Wang et al. (2021) ’s observations. Compared to Table 1 of Chen et al. (2018) (their data in parentheses), our respective measurements of CIB DL.055 and CIB PZH. 2017.09.01 were 377 and 783 mm for total length (versus 473.2 and 826.3 mm), 102 and 176 mm for tail length (versus 100.5 and 172.6 mm), 210 in each specimen for ventrals (versus 211 and 213), 84 and 80 for subcaudals (versus 88 and 86), 27+10 and 26+9 for dorsal + tail bands (versus 37+13 and 36+13), and 9 and 8 for the relative ventral scale position of the first band (versus both ventral 7). The discrepancies in tail length are trivial, and the ventral scale and first band counts might reflect different counting methods since Chen et al. (2018) did not specify what procedures they used to score these characters. However, the differences between our counts for total length, subcaudals and body+tail bands are more significant, and it is uncertain as to why the values in Chen et al. (2018) are so different from ours. One possibility is that Chen et al. (2018) may have added the number of body and tail bands together by mistake because their values are similar to the summed version of our body + tail band counts (e.g., 37 and 35, respectively). However, we have repeated the counts we originally obtained for the Panzhihua specimens and found no match between our results and those from Chen et al. (2018), and therefore report our own data throughout this study. Irrespective of the morphological discrepancies found within Chen et al. (2018), there is no doubt that the two specimens originally reported by these authors represent L. gongshan , thus confirming the presence of this species in Sichuan Province.

Figures

FIGURE 1. Distribution ranges of Lycodon latifasciatus sp. nov., L. fasciatus, and L. gongshan. Notes: numbers indicate different localities where the species have been recorded (see Appendix Table S4 for the details of localities).

FIGURE 2. Maximum-likelihood phylogenetic tree of Lycodon inferred from two mitochondrial genes (16S RNA, CytB) and one nuclear gene (Rag1); (A) Inset version of ML phylogeny with (B) showing a pruned version of same tree showing relationships within the Lycodon fasciatus group, and (C) showing a haplotype network based on phased Rag1 sequences of Lycodon. Numbers between nodes denote Ultrafast Bootstrap (UF BS), Shimodaira-Hasegawa approximate likelihood ratio test value (SH-aLRT), and Bayesian posterior probability values (BI PP), respectively. Nodes filled in with a black circle indicate strong support by both analyses, whereas nodes filled in with a white circle indicate strong support by only one of the ML analysis (either SH-aLRT or UF BS). Inset photo of Lycodon latifasciatus sp. nov. taken by N. A. Poyarkov.

FIGURE 5. Photographs of live Lycodon latifasciatus sp. nov., L. fasciatus, and L. gongshan.—(A–C) L. latifasciatus sp. nov. from Myanmar and China: (A) ZMMU Re-16728 (paratype, adult male) from Ban Mauk District, Sagaing Region, Myanmar; (B) ZMMU Re-16729 (immature male), same locality as ZMMU Re-16728; (C) CIB DL.2015.25.52 (paratype, adult male) from Medog County, Xizang AR, China.—(D–F) L. fasciatus specimens from China and India: (D) uncollected specimen (adult, sex unknown) from Dehong, Yunnan Province, China; (E) uncollected specimen (adult, sex unknown) from Daweishan NR, Pingbian County, Yunnan Province; (F) uncollected specimen (adult, sex unknown) from Aizawl, Mizoram State, India.—(G–I) L. gongshan in China: (G) uncollected specimen (adult, sex unknown) from Tengchong, Baoshan County, Yunnan Province; (H) CIB YN201909286 (adult male) from Lushui, Nuijang County, Yunnan; (I) uncollected specimen (adult, sex unknown) from Zayu County, Xizang AR (iNaturalist obs. 191251388). Photos by N. A. Poyarkov (A), P. Pawangkhanant (B), L. Ding (C, D, G), H. R. Gao (E), P. Shinde (F), S. C. Shi (H), and K. Ouyang (I).

FIGURE 6. Morphological comparisons between Lycodon latifasciatus sp. nov., L. fasciatus, and L. gongshan showing differences in head shape and coloration between adult and subadult specimens in right lateral (1), dorsal (2), and ventral (3) views, respectively. (A–B) L. latifasciatus sp. nov., (A) CAS 245367 (adult male) and (B) CIB DL.053 (adult female). (C–D) L. fasciatus, (C) FMNH 270716 (adult male), (D) CAS 245498 (subadult female). (E–F) L. gongshan, (E) CAS 241963 (adult male), (F) CAS 245470 (subadult male). Photos by J. L. Lee (A, C–F) and G. Vogel (B).

FIGURE 7. Hemipenial morphology of Lycodon latifasciatus sp. nov., L. fasciatus, and L. gongshan, showing asulcate (1), sulcate (2), and apical views (3). (A) L. latifasciatus sp. nov. CAS 245367 (holotype), partially everted organ; (B) L. fasciatus (FMNH 270716), completely everted organ; (C) L. gongshan (CAS 241963), partially everted organ. Photos by J. L. Lee.

FIGURE 8. New country record of Lycodon gongshan from Chipwi Township, Myitkyina District, Kachin State, Myanmar (CAS 245470; subadult male). (A) Dorsal and (B) ventral views of specimen; (C) right lateral, (D) left lateral (E) dorsal, and (F) ventral views of head. Photos by J. L. Lee.