Trirhinopholis styaniBoulenger 1899: 159

G. A. Boulenger described Plagiopholis styani in 1899, based on two specimens, BMNH 1946.1.14.11-12 (Boulenger 1899), but did not designate a holotype. Therefore, we designate the adult male specimen, BMNH 1946.1.14.11, as the lectotype and the specimen BMNH 1946.1.14.12 as the paralectotype due to it being a juvenile and the morphological characters not being stable.

BMNH 1946.1.14.11, adult male, collected by J. D. La Touche, from Kuatun (= Guadun Village, Wuyishan City), Fukien (= Fujian Province), China (Fig. 3).

BMNH 1946.1.14.12, juvenile female, with the same collection information as the lectotype.

(n = 9): CIB 10194 (field number 64I6669, adult male), FJNU 3625004 (adult male), FJNU SR (adult male), FJNU 3625005 (adult female), FJNU 3525002 (adult female), FJNU3625006 (adult female) from Wuyishan City, Fujian Province, China; WYS 2024001 (adult male) from Mt. Huanggang (the main peak of the Wuyi Mountains), Yanshan County, Jiangxi Province, China; ZJFRR 202207001 (subadult male) from Anji County, Zhejiang Province, China; and ZJFRR 202407001 (adult female) from Lin’an District, Zhejiang Province, China.

Plagiopholis styani can be distinguished from other species within Plagiopholis by the combination of the following morphological characters: (1) no loreal (rarely 1), posterior nasal forming a suture with the single preocular; (2) two postoculars; (3) temporals 2+2 (rarely 2+1); (4) 6 (rarely 7) supralabials, the 3^rd and 4^th entering the eye; (5) 6 (rarely 5) infralabials; (6) maxillary teeth 20–21 in males and around 18 in females; (7) dorsal scale smooth and rows 15-15-15; (8) ventral scales 114–121 in males and 116–122 in females; (9) anal entire; (10) subcaudal scales, 26–30 in males and 23–28 in females; (11) longer tail 42.36–51 mm (mean 45.42 mm) in males and shorter tail 32.6–44.15 mm (mean 40.59 mm) in females; (12) higher TAL/SVL ratio of 0.15–0.17 (mean 0.16) in males and smaller TAL/SVL ratio of 0.12–0.15 (mean 0.14) in females; (13) hemipenis half divided, bilaterally symmetrical, the truncus smooth throughout with almost no spinules; (14) a wide, black-brown blotch is present on the neck, often pentagonal, sub-annular, or sagittate-shaped, across 3–6 dorsal scale rows; (15) dorsal surface of the body is brick-red, with some dorsal scale edges black, forming a faint reticulated pattern.

Measurements and scalation. An adult male with TL 347 mm (SVL 303 mm and TAL 44 mm); tail relatively short, TAL/TL ratio of 0.13, and TAL/SVL ratio of 0.15; body slender and small; head slightly distinct from the neck; HW 8.46 mm; HL 12.41 mm; eye moderate, ED 2.49 mm; rostrum triangular, wider than high, slightly visible from above; internasals and prefrontals paired; frontal haxagonal, longer than wide, a little shorter than the parietals; PRO 1/1, PO 2/2; loreal absent; TEMP 4, arranged in two rows (2+2/2+2); SL 6/6, the 3^rd and 4^th entering the eye, the 5^th and 6^th largest; 2 pairs of chin-shields; IL 5/6, the 1^st to 3^rd touching the first pair of chin shields. DSR 15-15-15, all smooth, including the outermost row on both sides; VS 114, SC 27, CP entire.

Coloration and pattern. The dorsal surface of the head is reddish-brown, scattered with small black spots. The labial scales are light yellow with black edges on both sides. A blackish-brown subpentagonal blotch is present on the neck, pointing toward the head, and the widest part across four dorsal scale rows. Dorsal surface of the body is reddish-brown, with some dorsal scale edges outlined in black, forming a subtle reticulated pattern. Ventral and subcaudal scales are light yellow, with small dark brown spots on both sides (Fig. 3).

Head scalation.SL 6/6 (rarely 7), IL 6 (rarely 5); loreal absent or rarely present (1/12 specimens); PRO 1, PO 2; TEMP 2+2, rarely 2+1 (1/12 specimens).

Dentition. Based on two male specimens (CIB 10194 and ZJFRR 202207001) and one female specimen (ZJFRR 202407001) examined in this study and Pope (1935). Maxillary teeth 20–21 in males and 18 in females, without diastema; except for the first 2–4 being smaller, the rest are nearly equal in size.

Body scalation.DSR 15-15-15, all smooth, including the outermost row; VS 114–121 in males, and 116–122 in females; SC 26–30 in males, and 23–28 in females; CP entire.

Coloration in life. In life, the dorsal surface of the head is deep reddish-brown, sometimes scattered with small black-brown spots. The labial scales are creamy-white with black edges on both sides. A wide black-brown blotch is present on the neck, often pentagonal or irregularly ring-shaped, across 4 to 6 dorsal scale rows. Dorsal surface of the body brick-red, with some dorsal scale edges black, forming a faint reticulated pattern. The ventral and subcaudal scales range from creamy white to pale yellow, sometimes with small dark brown spots along the sides (Fig. 4).

Coloration in preservation. In preservation, the coloration still resembles the specimen in life, except that the coloration of dorsum further deepens, and the background color of the venter becomes uniform pale yellow (Figs 3, 5).

Hemipenis. Description based on the everted left organ of the subadult male specimen ZJFRR 202207001 (Fig. 6). When fully everted, the hemipenis is Y-shaped, bilaterally symmetrical, extends to SC 8, and bifurcates at lower 4/7.

Truncus is smooth, with almost no visible spinules; the spine ornamentation starts at the base of each branch. On the asulcate surface, each branch has a well-developed, calcareous basal hook, and the upper part with dense, medium-sized spines. The spines gradually decrease in size distally, reaching nearly half of the branches, then merging into thorny calyces. The calyx area extends to the distal end of each branch. Sulcate surface is similar to the asulcate surface, except for the absence of basal hook. The sulcus spermaticus is divided, extending centrifugally to the tip of each branch, and the sulcus lips are visible and slightly raised.

See Table 3 and Figs 3–5 for the details. The longest known male specimen has a TL of 353 mm (WYS 2024001; SVL 302 mm, TAL 51 mm), and the longest known female has a TL of 406 mm (KIZ 20180002; SVL 359 mm, TAL 47 mm). Tail relatively short, TAL/SVL ratio of 0.15–0.17 (mean 0.16) in males and 0.12–0.15 (mean 0.14) in females.

Plagiopholis styani have been found in Lin’an District of Hangzhou City, Anji County of Huzhou City in Zhejiang Province, and the Wuyi Mountains at the border of Fujian and Jiangxi Provinces (data corrected based on the results of the present study). According to the morphological data recorded in literature, distribution records from southern Anhui and northern Jiangxi Province might also belong to this species (Pope 1934; Chen 1991; Zhao et al. 1998; Zhao 2006). These regions are part of the mid-subtropical monsoon climate zone, characterized by distinct four seasons, warm and humid conditions (Zheng et al. 2013). This species is oviparous and distributed within an altitude range of 520–1359 m. They are commonly found in mid-subtropical evergreen broad-leaved forests and bamboo forests in this altitude range.

Pope (1929, 1935) documented the Fukien (now Fujian Province) population (Plagiopholis styani) to be found in the high forests and bamboo groves in the region of the type locality, which is consistent with our findings. He documented the stomachs of three Chungan Hsien (now Wuyishan City) specimens (P. styani), which contained a lot of gritty dirt suggestive of an earthworm diet. In addition, he recorded a female containing seven eggs, one of which measured 15.5 × 5.5 mm. Based on this observation, the species is inferred to be oviparous.

QHU 2024009, adult male, collected by the team of Lifang Peng in August 2024 from Xinshao County, Shaoyang City, Hunan Province, China (27.4533°N, 111.7558°E, 530 m a.s.l.)

QHU 2024010 (adult female), QHU 2024011 (adult male), with the same collected information as the holotype; QHU 2024044 (adult female) collected by the team of Lifang Peng in October 2024, from the same locality as the holotype.

QHU 2024012, juvenile, an offspring hatched in captivity on 16 September, 2024, from the egg laid by the female specimen QHU 2024010.

Plagiopholis acuta sp. nov. can be differentiated from other taxa in the genus Plagiopholis by the following combination: (1) no loreal, posterior nasal forming a suture with the single preocular; (2) two postoculars; (3) temporals 2+2 (rarely 2+1); (4) 6 (rarely 7) supralabials, the 3^rd and 4^th entering the eye; (5) 6 (rarely 5) infralabials; (6) maxillary teeth 17–18 in males and 15–17 in females; (7) dorsal scale smooth and rows 15-15-15; (8) ventral scales 114–115 in males and 115–121 in females; (9) anal entire; (10) subcaudal scales 29–30 in males and 24–25 in females; (11) longer tail length of 51–52 mm (mean 51.5 mm) in males and shorter tail length of 25–43 mm (mean 34 mm) in females; (12) larger TAL/SVL ratio of 0.18–0.19 (mean 0.19) in males and lower TAL/SVL ratio of 0.11–0.14 (mean 0.12) in females; (13) hemipenis half divided, bilaterally symmetrical, the truncus spinulate throughout; (14) a wide, black-brown blotch is present on the neck, often pentagonal or sub-annular shaped, across 4–6 dorsal scale rows; (15) dorsal surface of the body is brick-red, with some dorsal scale edges black, forming a faint reticulated pattern.

Measurements and scalation. An adult male with TL 313 mm (SVL 262 mm and TAL 51 mm); tail relatively short, TAL/TL ratio of 0.16, and TAL/SVL ratio of 0.19; body slender and small; head slightly distinct from the neck, HL 12.8 mm, HW 7.8 mm, HH 6.0 mm; eye moderate, ED 2.12 mm; rostrum triangular, wider than high, slightly visible from above; snout relatively short, ESD 3.7 mm; internasals and prefrontals paired; frontal haxagonal, longer than wide, a little shorter than the parietals; PRO 1/1, PO 2/2; loreal absent; TEMP 4, arranged in two rows (2+2/2+2); SL 6/6, the 3^rd and 4^th entering the eye, the 5^th and 6^th largest; chin shields in 2 pairs, the anterior pair longer than posterior one and in contact with the mental; IL 6/6, the 1^st to 3^rd touching the first pair of chin shields. DSR 15-15-15, all smooth, including the outermost row on both sides; VS 115, SC 30, CP entire.

Dentition. Maxillary teeth 18/17, without diastema; except for the first 3 being smaller, the rest are nearly equal in size.

Coloration in life. In life, dorsal surface of the head is deep reddish-brown, with small black-brown spots scattered along the sides; the supralabials are creamy-white with black edges on both sides. Ventral surface of head cream-white, the edges of partial scales have irregular black-brown patches. A wide, black-brown collar is present on the neck, across 5 dorsal scale rows. Dorsal surface of the body is brick-red, with some dorsal scale edges black, forming a faint reticulated pattern. The ventral and subcaudal scales uniform pale yellow (Fig. 7).

Coloration in preservation. In preservation, the coloration still resembles the specimen in life, except that the coloration of dorsum further deepens, and the ventral surface of head becomes pale yellow (Figs 8, 9).

The main morphological characters of Plagiopholis acuta sp. nov. are listed in Table 4. The paratypes exhibit a similar morphological pattern as the holotype, but there is a pronounced sexual dimorphism: compared to females, the examined males have a significantly longer tail, TAL/TL ratio of 0.15–0.16 and TAL/SVL ratio of 0.18–0.19 (vs. TAL/TL ratio of 0.10–0.12 and TAL/SVL ratio of 0.11–0.14 in females), fewer ventrals (114–115 vs. 115–121 in females), more subcaudals (29–30 vs. 24–25 in females), and more maxillary teeth (17–18 vs. 15–17 in females).

Hemipenis. Description based on the everted left organ of the male paratype QHU 2024011 (Fig. 10). When fully everted, the hemipenis is Y-shaped, bilaterally symmetrical, extends to SC 7, and bifurcates at lower 4/7. Both sulcate and asulcate surfaces densely ornamented with spinules, spines, or calyxes. Truncus spinulate throughout, branches divided into spine and calyx areas; a well-developed, calcareous basal hook is present on the asulcate surface of each branch.

Spiny area characterized by medium-size, dense spines, gradually decrease in size distally, reaching nearly half of the branches, then merging into thorny calyces. The calyx area extends to the distal end of each branch. The sulcus spermaticus is divided, extending centrifugally to the tip of each branch, and the sulcus lips are visible and slightly raised.

The specific epithet “acuta” means “sharp,” referring to the large, pointed basal hooks on the hemipenis of this species. Based on the type locality, Xinshao County, Hunan Province, we suggest “Hunan Mountain Snake” as its English common name and “湖南颈斑蛇 (Hú Nán Jǐng Bān Shé)” as its Chinese common name.

Specimen QHU 2024010 was a gravid female collected on August 7, 2024, and then kept in captivity under laboratory conditions at Qinghai University. It was housed in a 60 cm × 40 cm reptile terrarium with a substrate layer of approximately 3 cm of coarse peat soil. Humidity was maintained by providing a water dish and regular misting. On 11 August, the female laid five eggs, with an average length of 21 mm. The eggs were incubated in a moist mixture of coarse vermiculite and perlite at a constant temperature of 27 °C. After five days of incubation, candling with a flashlight revealed clear signs of embryonic development in only two eggs, while the remaining three showed no visible signs of development. After one week, the undeveloped eggs became moldy and shriveled. During the remainder of the incubation period, one of the two fertilized eggs failed to develop successfully. On September 16, the other one hatched (specimen QHU 2024012), measuring approximately 92 mm in length and weighing 1 g. However, it exhibited some deformities and morphological abnormalities (Table 4, Fig. 11). The first shedding occurred on September 25, and then the snake died on September 28. Based on the feeding and incubation results, we infer that this species may be better adapted to mid-elevations below 2000 m and relatively cool environments. The observed deformities may have been caused by high-altitude conditions or elevated temperatures during transportation and incubation.

Plagiopholis acuta sp. nov. has currently only been discovered in Xinshao County of Shaoyang City, Hunan Province. This area belongs to the mid-subtropical monsoon humid climate zone, characterized by distinct four seasons, abundant sunlight and heat, plentiful rainfall, with the rainy season coinciding with the hot season (Zheng et al. 2013). This new species is most probably distributed within mid-subtropical evergreen broad-leaved forests at an altitude of 530 m.

Plagiopholis acuta sp. nov. can be separated from P. styani as follows: (1) fewer maxillary teeth, 17–18 in males and 15–17 in females (vs. 20–21 in males and 18 in females); (2) tail relatively longer in males, and shorter in females (TAL/SVL ratio of 0.18–0.19, mean 0.19 vs. 0.15–0.17, mean 0.16 in males, and 0.11–0.14, mean 0.12 vs. 0.12–0.15, mean 0.14 in females); and (3) The truncus of the hemipenis spinulate throughout (vs. smooth). A detailed comparison between the new subspecies and its congeners is summarized in Table 5.

Plagiopholis acuta sp. nov. can be easily separated from P. blakewayi, P. nuchalis, and P. delacouri by having TEMP 2+2 (vs. 0/1+1 in P. blakewayi, 1+2 in P. nuchalis and P. delacouri) and different body coloration. Moreover, it can also be separated from P. blakewayi due to the more supralabials (6 vs. 5); and from P. nuchalis due to the fewer maxillary teeth in females (15–17 vs. 20–22) and the absence of the loreal (vs. loreal present).

Trirhinopholis styani – (part: population from Sichuan) Stejneger (1925);

Plagiopholis styani – (part: population from Sichuan) Ding and Zheng (1974).

CIB 119370 (field number EMS 202021), adult male, collected by the team of Bo Cai on 10 August 2020, from Longdong Village, Huangwan Town, Emeishan City, Sichuan Province, China (29.5798°N, 103.2937°E, 1324 m a.s.l.).

Males (n = 14): CIB 10176 (field number 562244), CIB 10177 (field number 55494), CIB 10180 (field number 505119), CIB 10181 (field number 561771), CIB 10184 (field number 561773), CIB 10186 (field number 562156), CIB 10189 (field number 562153), CIB 10192 (field number 562171), CIB 10193 (field number 12014) were collected from Mt. Emei in Emeishan City, Sichuan Province, China; CIB 10188 (field number 715013) was collected from Hongya County. Meishan City, Sichuan Province, China; QHU 2024006 from Mt. Qingcheng, Dujiangyan City, Sichuan Province, China; CIB XMG 2023001, Dujiangyan Panda Valley, Chengdu Research Base of Giant Panda Breeding, Chengdu City, Sichuan Province, China; QHU 2024007 was collected from Anziping Village, Wenchuan County, Aba Tibetan and Qiang Autonomous Prefecture, Sichuan Province, China. Females (n = 9): CIB 10178 (field number 562182), CIB 10179 (field number 562180), CIB 10182 (field number 562266), CIB 10185 (field number 561772), CIB 10187 (field number 561775), CIB 10191 (field number 562297), FMAEM were collected from Mt. Emei in Emeishan City, Sichuan Province, China; CIB CB25DY01, a roadkilled male, was collected from Dayi County in the Chengdu Section of Giant Panda National Park, Sichuan Province, China; SYS r002035 was collected from Baima Mountain Forest Farm, Bijie City, Guizhou Province. Juveniles (N = 2): CIB CB2024EMS001 was collected from Mt. Emei in Emeishan City, Sichuan Province, China; CIB 10183 (field number 595027) was collected from Hongya County, Meishan City, Sichuan Province, China.

Plagiopholis pluvialis sp. nov. can be diagnosed from other Plagiopholis species by the following unique combination of characters: (1) no loreal (rarely 1), posterior nasal forming a suture with the single preocular; (2) two postoculars; (3) temporals 2+2 (rarely 2+1); (4) 6 supralabials, the 3^rd and 4^th entering the eye; (5) 6 infralabials; (6) maxillary teeth 18 in males; (7) dorsal scale smooth and rows 15-15-15; (8) ventral scales 102–110 in males and 114–122 in females; (9) anal entire; (10) subcaudal scales 26–31 in males and 16–26 in females; (11) longer tail length 40–59 mm (mean 49 mm) in males, and shorter tail length 34–47 mm (mean 40 mm) in females; (12) larger TAL/SVL ratio of 0.14–0.21 (mean 0.18) in males and lower TAL/SVL ratio of 0.12–0.15 (mean 0.14) in females; (13) hemipenis half divided, bilaterally symmetrical, the truncus spinulate throughout; (14) a relatively narrow, black-brown blotch is present on the neck, often inverted “V” shaped, across 2–3 dorsal scale rows; (15) dorsal surface of the body is olive, with some dorsal scale edges yellow and black, forming a reticulated pattern.

Measurements and scalation. An adult male with TL 322 mm (SVL 282 mm and TAL 40 mm); tail relatively short, TAL/TL ratio of 0.12, and TAL/SVL ratio of 0.14; body slender and small; head slightly distinct from the neck, HL 11.5 mm, HW 7.6 mm, HH 5.7 mm; eye moderate, ED 2.3 mm; rostrum triangular, wider than high, slightly visible from above; snout relatively short, ESD 4.1 mm; internasals and prefrontals paired; frontal hexagonal, longer than wide, a little shorter than the parietals; PRO 1/1, PO 2/2; loreal absent; TEMP 4, arranged in two rows (2+2/2+2); SL 6/6, the 3^rd and 4^th entering the eye, the 6^th largest; chin shields in 2 pairs, the anterior pair longer than posterior one and in contact with the mental; IL 6/6, the 1^st to 3^rd touching the first pair of chin shields. DSR 15-15-15, all smooth, including the outermost row on both sides; VS 108, SC 27, CP entire.

Dentition. Maxillary teeth 18, without diastema; except for the first 3 being smaller, the rest are nearly equal in size.

Coloration in life. In life, dorsal surface of the head is olive, with small black-brown spots scattered along the sides; the supralabials are wheat with broad black edges on both sides. Ventral surface of head wheat, partial scales have irregular black-brown patches. A relatively narrow, black-brown collar is present on the neck, inverted “V” shaped, across 2–3 dorsal scale rows. Dorsal surface of the body is olive, with some dorsal scale edges gold and black, forming a faint reticulated pattern. The ventral and subcaudal scales uniform wheat. (Fig. 12).

Coloration in preservation. In preservation, the coloration still resembles the specimen in life (Figs 13, 14).

The main morphological characters of Plagiopholis pluvialis sp. nov. are listed in Table 6. All known specimens exhibit a similar morphological pattern to the holotype, but there is pronounced sexual dimorphism: compared to females, the examined males have a significantly long tail, TAL/TL ratio of 0.12–0.18 and TAL/SVL ratio of 0.14–0.21 (vs. TAL/TL ratio of 0.11–0.13 and TAL/SVL ratio of 0.12–0.15 in females), fewer ventrals (102–110 vs. 114–122 in females), more subcaudals (26–31 vs. 16–26). Most specimens have temporals 2+2 on each side, but the left of CIB10182 (field number 562266) is 2+1. Most specimens loreal absence but the female specimen USNM 65505 in Stejneger (1925) has one loreal on each side.

Hemipenis. Description based on the everted left organ of the male paratypes QHU 2024006 and QHU 2024007 (Fig. 15). When fully everted, the hemipenis is Y-shaped, bilaterally symmetrical, extends to SC 7 in QHU 2024007 and SC 8 in QHU 2024006, bifurcates at about lower 5/9. Both sulcate and asulcate surfaces densely ornamented with spinules, spines, or calyxes. Truncus spinulate throughout, branches divided into spine and calyx areas; a well-developed, calcareous basal hook is present on the asulcate surface of each branch.

Spine area characterized by medium-sized, dense spines, gradually decrease in size distally, reaching nearly half of the branches, then merging into thorny calyces. The calyx area extends to the distal end of each branch. The sulcus spermaticus is divided, extending centrifugally to the tip of each branch, and the sulcus lips are visible and slightly raised.

The Latin specific epithet pluvialis means ‘related to rain’ or ‘rainy,’ indicating that the species inhabits particularly moist environments, especially within the West China Rain Zone in China. It is suggested that the English name for this species be “Western China Mountain Snake” and the Chinese name be “华西颈斑蛇 (huá xī jǐng bān shé)”.

This species is known to be distributed in mountainous areas at altitudes of 744–1900 m at several localities, including Wenchuan County in Aba Tibetan and Qiang Autonomous Prefecture; Anzhou District in Mianyang City; Dayi County and Dujiangyan City in Chengdu City; Hongya County in Meishan City; and Emeishan City in Leshan City, all in Sichuan Province, China; as well as Qixingguan District in Bijie City and Mt. Fanjing in Tongren City, both in Guizhou Province, China (Stejneger 1925; Pope 1929, 1935; Zhao et al. 1998; Zhao 2006; Zhong et al. 2015; Liu et al. 2025, and this study). The distribution areas fall within the mid-subtropical monsoon climate zone characterized by mixed evergreen and deciduous broad-leaved forests, where four distinct seasons are observed, with hot and rainy summers and mild, less rainy winters (Zheng et al. 2013). Particularly the snake’s main distribution area lies within the West China Rain Zone in Sichuan, a unique geographical region known for its significant rainfall due to the convergence of various climatic fronts, making it one of the wettest places in China.

Pope (1929, 1935) documented that the stomachs of two specimens from Mt. Emei had eaten an earthworm apiece and found 5, 6, and 11 eggs, respectively, in three Mt. Emei females. One of the eggs in the clutch of 5 measured 19 × 11 mm and contained a very small embryo, while the female with 11 eggs was collected in July. This new species is also certainly oviparous.

Plagiopholis pluvialis sp. nov. differs from P. styani and P. acuta sp. nov. by having the following combined characteristics: (1) a relatively narrow, black-brown blotch is present on the neck, often inverted “V” shaped, across 2–3 dorsal scale rows (vs. a wide, black-brown blotch is present on the neck, often pentagonal or irregularly semiring-shaped, across 4 to 6 dorsal scale rows in P. styani and P. acuta sp. nov.); (2) the truncus of the hemipenis spinulate throughout (vs. smooth in P. styani); (3) maxillary teeth 18 in males (vs. 20–21 in P. styani in males); (4) fewer ventral scales 102–110 in males (vs. 114–115 in P. acuta sp. nov. and 114–121 in P. styani in males); (5) the dorsal surface of the body is olive, with some dorsal scale edges gold and black (vs. brick-red, with some dorsal scale edges black in P. styani and P. acuta sp. nov.); (Fig. 16). A detailed comparison between the new species and its congeners is summarized in Table 5.

Plagiopholis pluvialis sp. nov. can be easily separated from P. blakewayi, P. nuchalis, and P. delacouri by having TEMP 2+2 (vs. 0/1+1 in P. blakewayi, 1+2 in P. nuchalis and P. delacouri) and significantly different body coloration. Moreover, it can also be separated from P. blakewayi due to the more supralabials (6 vs. 5).