A new species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from Yunnan, China and its phylogenetic relationship to other congeners Author Agung, Ade Prasetyo 0000-0002-0954-2960 Landscape Ecology Group, Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Menglun, Mengla, Yunnan 666303, China. & University Chinese Academy of Sciences, Huairou, China. & prasetyo. ade 22 @ gmail. com; https: // orcid. org / 0000 - 0002 - 0954 - 2960 prasetyo.ade22@gmail.com Author Grismer, L. Lee 0000-0002-2369-6405 Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515, USA. Lee Kong Chian Natural History Museum, National University of Singapore, 2 Conservatory Drive, 117377 Singapore. & lgrismer @ lasierra. edu; https: // orcid. org / 0000 - 0002 - 2369 - 6405 lgrismer@lasierra.edu Author Grismer, Jesse L. 0000-0002-2542-5430 Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515, USA. Lee Kong Chian Natural History Museum, National University of Singapore, 2 Conservatory Drive, 117377 Singapore. & jgrismer @ lasierra. edu; https: // orcid. org / 0000 - 0002 - 2542 - 5430 jgrismer@lasierra.edu Author Quah, Evan S. H. 0000-0002-5357-1953 evanquah @ yahoo. com; https: // orcid. org / 0000 - 0002 - 5357 - 1953 evanquah@yahoo.com Author Chornelia, Ada 0000-0002-8541-5947 Landscape Ecology Group, Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Menglun, Mengla, Yunnan 666303, China. & chorneliaa @ gmail. com; https: // orcid. org / 0000 - 0002 - 8541 - 5947 chorneliaa@gmail.com Author Lu, Jianmei 0000-0001-5917-6084 Landscape Ecology Group, Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Menglun, Mengla, Yunnan 666303, China. & lujianmei @ xtbg. ac. cn; https: // orcid. org / 0000 - 0001 - 5917 - 6084 lujianmei@xtbg.ac.cn Author Hughes, Alice C. 0000-0002-4899-3158 Landscape Ecology Group, Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Menglun, Mengla, Yunnan 666303, China. & ach _ conservation 2 @ hotmail. com; https: // orcid. org / 0000 - 0002 - 4899 - 3158 ach_conservation2@hotmail.com text Zootaxa 2021 2021-05-31 4980 1 1 27 journal article 6103 10.11646/zootaxa.4980.1.1 f3fe1336-f9be-4f9d-84f6-8713b988de3a 1175-5326 4882802 F0BBBFB4-489E-42B0-9720-FD68ED2E6565 Hemiphyllodactylus zhutangxiangensis sp. nov. Zhutangxiang slender gecko Holotype . Adult male ( KIZ061157 ) collected on 21 August 2018 by Ade P. Agung , Ada Chornelia , Jianmei Lu, L. Lee Grismer, Jesse L. Grismer , Evan S.H. Quah , Brian Folt , and Myin Kyaw Thura from forested karst approximately 1 km northwest of Zhutangxiang town , Lancang Lahu Autonomous County , Yunnan Province , China ( 22.72345°N , 99.80087°E ; 1499 m in elevation). Paratypes . Four adult females ( KIZ061156 , KIZ061158–60 ) bear the same data as the holotype, two adult males ( KIZ061164 , KIZ061166 ) and six adult females ( KIZ061161–63 , KIZ061165 , KIZ061167–68 ) were collected on 26 June 2019 by Ade P. Agung , Jianmei Lu and Yang Zongbao from karst forest nearby Zhutangxiang town , Lancang Lahu Autonomous County , Yunnan Province , China ( 22.732306°N 99.798167°E ; 1492 m in elevation) . Diagnosis. Hemiphyllodactylus zhutangxiangensis sp. nov. can be distinguished from all congeners by having the unique combination of a maximum SVL of 44.42 mm ; 7–9 chin scales; enlarged postmentals; five circumnasal scales; 2–4 internasal scales; 8–11 supralabial scales; 8–11 infralabial scales; subdigital lamellae on fingers II–V (3–5)-(4–6)-(4 or 5)-(4 or 5); subdigital lamellae on toes II–V (4 or 5)-(4 or 5)-(4–6)-(4 or 5); dorsal scales 11–15; ventral scales 5–7; having pale-grey base color on body with various transverse blotched patterns on the dorsal side; a dark postorbital stripe extending to at least base of neck; no dorsolateral or ventrolateral stripe on trunk; postsacral marking bearing dark fork-like pattern with anteriorly projecting arms. FIGURE 2. Maximum Likelihood topology illustrating the phylogenetic relationships between Hemiphyllodactylus zhutangxiangensis sp. nov. (within clade 3) and other Hemiphyllodactylus species. Nodes with UFBoot ≥ 95 are considered highly supported. FIGURE 3. Bayesian Inference topology illustrating the phylogenetic relationships between Hemiphyllodactylus zhutangxiangensis sp. nov. (within clade 3) and other Hemiphyllodactylus species. Nodes with BPP ≥ 0.95 are considered highly supported. FIGURE 4. Mensural (7) and meristic (1) traits that differ significantly between H. changningensis and H. zhutangxiangensis sp. nov. Boxplots show raw data, but significance tests for mensural traits were conducted using adjusted values aligned to the grand mean of SVL across both species (see Methods). See Table 5 for significance levels of differences. FIGURE 5. Hemiphyllodactylus zhutangxiangensis sp. nov. : (A) holotype, KIZ061157, male in life; (B) paratype, KIZ061159, female, in life; (C) & (D) ventral and dorsal view of the holotype’s head in preserved condition, black circles indicates Chin and IS, respectively; (E) dorsal view of holotype (KIZ061157)—on the left, with three paratypes (KIZ061159, KIZ061156, KIZ061158), respectively. Photos: (A) & (B) by Evan S. H. Quah; (C), (D) & (E) by Ada Chornelia. TABLE 4 . The mean percentage of the uncorrected pairwise genetic distance (p-distance) between Hemiphyllodactylus species.

Species name (n)	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15
1. H. linnwayensis (2)	-
2. H. montawaensis (6)	11.8	-
3. H. ngwelwini (9)	9.1	10.4	-
4. H. tonywhitteni (5)	10.1	6.2	9.5	-
5. H. uga (1)	4.6	11.9	9.7	9.8	-
6. H. ywanganensis (2)	2.7	11.8	9.1	9.1	3.9	-
7. H. changningensis (4)	19.0	17.2	19.0	17.4	18.3	19.9	-
8. H. zalonicus (1)	18.5	17.0	17.8	17.6	17.4	18.4	13.9	-
9. H. longlingensis (4)	18.1	18.3	18.1	17.7	17.4	18.2	17.6	17.6	-
10. H. khlonglanensis (7)	15.9	16.4	16.4	16.1	16.2	16.6	16.8	16.8	19.1	-
11. H. jinpingensis (4)	11.0	11.7	11.1	11.9	12.6	11.5	16.7	16.8	17.8	15.7	-
12. H. chiangmaiensis (2)	14.3	13.5	13.6	13.0	12.7	13.9	18.6	18.8	18.4	16.2	14.0	-
13. H. kyaiktiyoensis (5)	19.2	18.7	19.1	18.6	18.8	19.8	19.8	20.1	20.8	18.7	18.9	18.2	-
14. H. pinlaungensis (14)	20.6	19.3	19.4	18.9	21.1	21.1	18.8	20.2	20.4	17.7	18.6	19.5	9.0	-
15. H. zwegabinensis (1)	18.9	20.1	19.5	19.0	18.3	19.4	18.4	21.7	19.8	18.1	19.4	17.6	15.8	16.9	-
16. H. zhutangxiangensis sp. nov. (13)	16.5	16.0	17.4	15.0	16.3	16.8	14.7	15.5	15.6	16.0	17.2	17.5	18.3	18.4	19.7

TABLE 5. Model choice and variance test results for pairwise comparisons of groups. Variance tests used were ANOVA for linear models and Chi-square tests for general linear models.

Trait	Selected model	Variance test probability
Mensural
SVL	gls	NS
TrunkL	lm	0.020
ED	lm	<0.001
HL	lm	<0.001
HW	lm	<0.001
NarEye	lm	0.018
SnEye	lm	0.020
SnW	lm	<0.001
Meristic traits
DS	glm	NS
VS	glm	NS
CloacS	glm	NS
CN	glm	0.051
IS	glm	NS
SL	glm	NS
IL	glm	NS
Chin	glm	NS
SL1F	glm	NS
SL1T	glm	NS

Description of holotype . Adult male, SVL 41.35 mm ; head triangular in dorsal profile, depressed, distinct from neck (HL 7.20 mm ; HW 7.57 mm ); lores flat; snout long (SnEye 3.95 mm ; SnEye/HL 55%), narrow (SnW 1.40 mm ; SnW/HW 16%); eye large (ED 2.12 mm ; ED/HL 30%); rostral scale wider than high, bordered posteriorly by two large supranasals and four internasals (IS); nares bordered anteriorly by rostral scale, ventrally by first supralabial scale, dorsally by supranasal scale, posteriorly by two postnasals; supralabials square, 9/9 (left/right), tapering from rostral to a point in line with posterior margin of orbit (SL); infralabials square, 9/9 (left/right), tapering from mental to a point in line with posterior margin of orbit (IL); scales on head small, rounded, largest on rostrum; mental triangular, bordered by first infralabials and posteriorly by two enlarged postmentals; each postmental bordered anterolaterally by first infralabial; eight chin scales touching internal edges of infralabials from juncture of the 2nd and the 3rd on left and right and mental scale ( Chin ); scales in gular region rounded, non-overlapping, becoming larger and more ovoid on venter. Robust body type and small, (TrunkL/SVL 48%), dorsoventrally compressed; dorsal body scales small, granular, 11 dorsal scales at midbody contained within one eye diameter; ventral body scales smooth and flat, much larger than dorsal scales, subimbricate, five ventral scales at midbody contained within one eye diameter; fore-limbs relatively short, covered dorsally with granular, subimbricate scales, smaller smooth scales ventrally; palmar scales flat, unevenly shaped, non-overlapping; finger I vestigial, clawless, fingers II–V welldeveloped; proximal subdigital lamellae undivided, rectangular; distal subdigital lamellae divided and undivided, angular, U-shaped, except terminal lamellae rounded, undivided; lamellar formula on fingers II–V (4)-(4)-(4)-(4) on left hand and (3)-(4)-(4)-(4) on right hand; claws on fingers II–V well-developed, unsheathed, strongly curved; hind limbs short, covered dorsally with granular, subimbricate scales, smaller smooth scales ventrally; plantar scales flat, unevenly shaped, non-overlapping; toe I vestigial, clawless, toes II–V well-developed; proximal subdigital lamellae undivided, rectangular; distal subdigital lamellae divided and undivided, angular, U-shaped, except terminal lamellae rounded, undivided; lamellar formula on toes II–V (4)-(4)-(4)-(4) on both feet; claws on toes II–V welldeveloped, unsheathed, strongly curved; cloacal spurs (CloacS) one on each side; tail long, original (TL 37.40 mm ; TL/SVL 90%), round in cross-section, dorsal scales on tail larger than on body and head, smaller than subcaudals, no plate-like subcaudal scales. Coloration in life ( Figure 5 ). All Hemiphyllodactylus are known to have the ability to considerably change the intensity and boldness of their coloration and pattern. The description below was taken when the holotype was photographed in the morning after it had been captured. The base color of the dorsal side of the head, body, and limbs are pale-grey and densely mottled with darker markings; top of head overlain with dark blotches; dark pre-and postorbital stripes extend from the external nares, through eye to just anterior of forelimb insertion on the body; postsacral marking bearing dark fork-like pattern with anteriorly projecting arms; limbs bearing irregularly shaped, diffuse, dark markings; tail bearing irregularly shaped, dark and light colored alternating markings; abdomen unicolor grey. Variation . Coloration of this species varies considerably ( Figure 5 ). Variation of mensural and meristic data are presented in Table 6 . Distribution. Hemiphyllodactylus zhutangxiangensis sp. nov. is at present only known from the type localities nearby Zhutangxiang town, Lancang Lahu Autonomous County, Yunnan Province , China ( Figure 7 ). FIGURE 6. Habitat of Hemiphyllodactylus zhutangxiangensis sp. nov. : (A) site where the specimens were found close to quarrying activities; (B) abandoned building where the specimens spotted; (C) view of karst hills near Zhutangxiang town, Lancang Lahu Autonomous County, Yunnan Province, China. Photos: (A) & (B) by Ada Chornelia; (C) by Jianmei Lu. TABLE 6. Mensural (in mm), meristic, color pattern, and proportions of the types series of Hemiphyllodactylus zhutangxiangensis sp. nov. from Zhutangxiang town, Lancang Lahu Autonomous County, Yunnan Province, China. (/) = data unavailable. (*) = regenerated tail. For abbreviations see Methods.

Character	Holotype	Paratypes
KIZ 061157	KIZ 061156	KIZ 061158	KIZ 061159	KIZ 061160	KIZ 061161	KIZ 061162	KIZ 061163	KIZ 061164	KIZ 061165	KIZ 061166	KIZ 061167	KIZ 061168
Sex	male	female	female	female	female	female	female	female	male	female	male	female	female
SVL	41.35	38.80	42.16	40.67	41.39	42.17	44.42	44.14	38.82	42.42	34.67	32.29	40.57
TL	37.40	36.90	33.91	31.98*	40.30	40.94	28.33*	37.17	34.74*	42.00	(/)	27.13*	29.19*
TrunkL	19.75	19.00	22.08	19.82	21.04	21.63	22.68	23.10	19.72	21.04	17.28	16.05	20.42
HL	7.20	7.00	7.29	6.79	7.04	7.39	7.64	7.47	7.08	7.46	6.31	6.22	7.44
HW	7.57	7.92	8.23	7.35	7.96	8.05	7.84	8.04	7.72	7.65	6.98	6.53	8.07
ED	2.12	2.34	2.40	2.42	2.46	2.27	2.70	2.40	2.29	2.42	2.20	2.05	2.48
SnEye	3.95	3.93	3.95	4.03	4.19	4.04	4.36	4.04	3.90	4.01	3.45	3.44	3.95
NarEye	3.04	2.74	2.98	2.67	3.01	3.20	3.20	3.01	3.08	3.03	2.63	2.58	3.08
SnW	1.40	1.36	1.62	1.48	1.55	1.61	1.42	1.39	1.39	1.50	1.25	1.35	1.41
TrunkL/SVL	0.48	0.49	0.52	0.49	0.51	0.51	0.51	0.52	0.51	0.50	0.50	0.50	0.50
HL/SVL	0.17	0.18	0.17	0.17	0.17	0.18	0.17	0.17	0.18	0.18	0.18	0.20	0.18
HW/SVL	0.18	0.20	0.20	0.18	0.19	0.19	0.18	0.18	0.20	0.18	0.20	0.20	0.20
HW/HL	1.05	1.13	1.13	1.08	1.13	1.09	1.03	1.08	1.09	1.03	1.11	1.05	1.09
SnEye/HL	0.55	0.56	0.54	0.59	0.60	0.55	0.57	0.54	0.55	0.54	0.55	0.55	0.53
NarEye/HL	0.42	0.39	0.41	0.39	0.43	0.43	0.42	0.40	0.44	0.41	0.42	0.42	0.41
ED/HL	0.30	0.33	0.33	0.36	0.35	0.31	0.35	0.32	0.32	0.32	0.35	0.33	0.33
SnW/HL	0.19	0.19	0.22	0.22	0.22	0.22	0.19	0.19	0.20	0.20	0.20	0.22	0.19
ED/NarEye	0.70	0.85	0.81	0.91	0.82	0.71	0.84	0.80	0.74	0.80	0.84	0.80	0.81
Snw/HW	0.16	0.17	0.20	0.20	0.20	0.20	0.18	0.17	0.18	0.20	0.18	0.21	0.18
Chin	8	8	9	7	7	8	8	9	9	9	8	8	8
CN	5	5	5	5	5	5	5	5	5	5	5	5	5
IS	4	4	3	2	3	3	3	3	3	3	3	3	2
SL (left/right)	9/9	8/9	9/10	10/10	10/9	9/10	10/9	9/9	10/10	10/11	8/9	9/9	10/9
IL (left/right)	9/9	8/9	8/10	9/11	9/8	10/10	9/10	10/9	9/10	10/11	9/8	10/9	9/9
VS	5	6	7	7	7	6	7	6	6	7	7	7	6

......continued on the next page TABLE 6. (Continued)

Character	Holotype	Paratypes
KIZ 061157	KIZ 061156	KIZ 061158	KIZ 061159	KIZ 061160	KIZ 061161	KIZ 061162	KIZ 061163	KIZ 061164	KIZ 061165	KIZ 061166	KIZ 061167	KIZ 061168
DS	11	13	15	13	14	13	14	13	12	12	14	14	14
Lamelar formulae hands II–V (left)	4-4-4-4	4-4-5-4	4-4-5-5	4-5-4-4	4-4-4-4	5-5-5-5	4-5-5-4	5-5-5-5	4-4-4-4	4-6-5-5	4-4-4-4	3-4-4-4	4-4-4-4
Lamelar formulae hands II–V (right)	3-4-4-4	4-4-5-4	4-5-5-5	4-5-4-4	4-4-4-4	5-5-5-5	4-5-5-4	5-5-5-5	4-4-4-4	4-6-5-5	4-4-4-4	3-4-4-4	4-4-4-4
Lamelar formulae foot II–V (left)	4-4-4-4	4-5-5-5	4-5-5-5	4-4-4-4	5-5-5-5	5-5-6-5	5-5-5-5	5-5-6-5	4-5-5-5	5-5-6-5	4-4-4-4	4-4-4-4	4-4-4-4
Lamelar formulae foot II–V (right)	4-4-4-4	4-5-5-5	4-5-5-5	4-4-4-4	5-5-5-5	5-5-5-5	5-5-5-5	5-5-5-5	4-5-5-5	5-5-6-5	4-4-4-4	4-4-4-4	4-4-4-4
SL1F	4	5	4	4	4	5	5	5	4 or 5	5	5	4	4
SL1T	5	4 or 5	4	4	4 or 5	5	5	5	4 or 5	5	5	4	4
Precloacal and femoral pore series separate (1) or continuous (0)	0	0	0	0	(/)	0	0	0	0	0	0	(/)	0
Total femoroprecloacal pores	23	0	0	0	(/)	0	0	0	20	0	22	(/)	0
CloacS on each side	1	2	1	2	1	1	1	1 or 2	1	1	1	1	1
Subcaudals enlarged, plate-like	No	No	No	No	No	No	No	No	No	No	No	No	No
Dark postorbital stripe	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes
Dorsolateral light-colored spots on trunk	No	No	No	No	No	No	No	No	No	No	No	No	No
Dark dorsolateral stripe on trunk	No	No	No	No	No	No	No	No	No	No	No	No	No
Dark ventrolateral stripe on trunk	No	No	No	No	No	No	No	No	No	No	No	No	No
Dark dorsal transverse blotches	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes
Dark reticulate pattern on dorsum	No	No	No	No	No	No	No	No	No	No	No	No	No
Postsacral marking bearing dark fork-like pattern anteriorly project- ing arms	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes

TABLE 7. Potentially diagnostic characters (in bold) separating the new species from other Hemiphyllodactylus within clade 3. (/) = data unavailable. (*) = regenerated tail. Mensural characters are in mm. For abbreviations see Methods. Data for H. zalonicus and H. longlingensis were obtained from Grismer et al. 2020b . Data for H. changningensis were obtained from Guo et al. 2015 .

Character	zalonicus	longlingensis	changningensis	zhutangxiangensis sp. nov.
Max SVL	37.7	45.7	43.8	44.42
n	2	(/)	10	13
TrunkL	18.1–18.9	(/)	17.4–22.5	16.1–23.1
ED	2.0–2.6	(/)	1.7–2.3	2.1–2.7
HL	8.4–8.5	(/)	8.2–10.1	6.2–7.6
HW	5.7–5.8	(/)	6.1–7.5	6.5–8.2
NarEye	2.4–2.7	(/)	2.8–3.4	2.6–3.2
SnEye	3.4–3.5	(/)	3.5–4.5	3.4–4.4
SnW	1.2	(/)	1.1–1.4	1.3–1.6
TrunkL/SVL	0.49–0.50	0.47–0.52	0.46–0.51	0.48–0.52
HL/SVL	0.22–0.23	0.22–0.24	0.22–0.25	0.17–0.20
HW/SVL	0.15	0.17–0.19	0.17–0.18	0.18–0.20
HW/HL	0.68	0.75–0.80	0.72–0.80	1.03–1.13
SnEye/HL	0.40–0.42	0.42–0.45	0.41–0.49	0.53–0.60
NarEye/HL	0.28–0.31	0.29–0.34	0.30–0.37	0.39–0.44
ED/HL	0.23–0.30	0.22–0.25	0.21–0.25	0.30–0.36
SnW/HL	0.14	0.12–0.14	0.12–0.16	0.19–0.22
ED/NarEye	0.74–1.08	0.66–0.82	0.61–0.77	0.70–0.91
SnW/HW	0.21	0.15–0.18	0.16–0.20	0.16–0.21
Chin	8–10	7–9	7–8	7–9
Postmentals distinctly enlarged	Yes	Yes	Yes	Yes
CN	5	4–5	3–4	5
IS	3–4	1–3	2–3	2–4
SL	10	9–10	8–11	8–11
IL	8–9	8–10	8–10	8–11
VS	9–10	6–7	6–8	5–7
DS	17–18	10–14	11–15	11–15

......continued on the next page TABLE 7. (Continued)

Character	zalonicus	longlingensis	changningensis	zhutangxiangensis sp. nov.
Lamellar formulae on hands (finger II–V)	3-3-3-3	3-4-4-4(3) 4-4-4(5)-4	3-3(4)-3(4)-3	(3–5)-(4–6)-(4 or 5)-(4 or 5)
Lamellar formulae on foot (finger II–V)	3-4-4-4	4-4(5)-4(5)-4	3-3-3-3 3-4-4-4	(4 or 5)-(4 or 5)-(4–6)-(4 or 5)
SL1F	3	4–5	3–4	4–5
SL1T	4	4–6	3–4	4–5
Precloacal and femoral pore series separate (1) or continuous (0)	0	0	0	0
Total femoroprecloacal pores	16–20	16–27	19–22	20–23
CloacS on each side	1	1 or 2	1 or 2	1 or 2
Subcaudals enlarged, plate-like	No	No	No	No
Dark postorbital stripe	Yes	Yes	Yes	Yes
Dorsolateral light-colored spots on trunk	No	No	No	No
Dark dorsolateral stripe on trunk	No	No	No	No
Dark ventrolateral stripe on trunk	No	No	No	No
Dark dorsal transverse blotches	No	Variable	No	Yes
Dark reticulate pattern on dorsum	No	Variable	Yes	No
Postsacral marking anteriorly projecting arms	Indistinct or No	Light-colored	No	Fork-like dark-colored

FIGURE 7 . Distribution map of H. zhutangxiangensis sp. nov. and several of its congeners from Clade 3, Clade 4, and Indochina clade. Natural history. The holotype was collected together with the four paratypes ( KIZ061156 , KIZ061158–60 ) in the evening on 21 August 2018 on the cement wall of an abandoned building which was surrounded by cement mining activities in a forested karst area approximately 1 km northwest of Zhutangxiang town ( Figure 6 ). The other eight paratypes ( KIZ061161–68 ) were collected together in the evening on 26 June 2019 approximately 1 km further north from the holotype’s locality, in another cement mine within an area of forested karst. The specimens were also collected when they were crawling on the cement wall of an abandoned building . Etymology. The specific epithet, zhutangxiangensis , refers to the name of the town Zhutangxiang in Lancang Lahu Autonomous County, Yunnan Province , China , close to where the specimens were found. Morphological comparisons. We statistically compared the new species with Hemiphyllodactylus changningensis in its sister lineage, in clade 3 (see results, Table 5 , and Figure 4 ). In addition, in terms of body ratios, H. zhutangxiangensis sp. nov. differs from H. changningensis , H. zalonicus , and H. longlingensis by having a shorter head (HL/SVL = 0.17–0.20 versus 0.22–0.25, 0.22–0.23, 0.22–0.24, respectively); wider head (HW/HL = 1.03–1.13 versus 0.72–0.80, 0.68, 0.75–0.80, respectively); greater SnEye distance (SnEye/HL = 0.53–0.60 versus 0.41–0.49, 0.40–0.42, 0.42–0.45, respectively); greater NarEye distance (NarEye/HL = 0.39–0.44 versus 0.30–0.37, 0.28–0.31, 0.29–0.34, respectively); larger eyes (ED/HL = 0.30–0.36 versus 0.21–0.25, 0.23–0.30, 0.22–0.25, respectively); a wider snout (SnW/HL = 0.19–0.22 versus 0.12–0.16, 0.14, 0.12–0.14, respectively). In terms of scalation, the new species differs from H. changningensis by having more CN (five versus three or four); differs from H. zalonicus by having fewer VS and DS (5–7 versus 9–10, and 11–15 versus 17–18, respectively). For the lamellar, femoroprecloacal pores, and coloration, the new species differs from H. changningensis and H. zalonicus by having more lamellae on the hand (3–5)-(4–6)-(4 or 5)-(4 or 5) versus (3)-(3 or 4)-(3 or 4)-(3), and (3)-(3)-(3)-(3), respectively); more lamellae on the foot (4 or 5)-(4 or 5)-(4–6)-(4 or 5) versus (3)-(3 or 4)-(3 or 4)-(3 or 4), and (3)-(4)-(4)-(4), respectively). Furthermore, H. zhutangxiangensis sp. nov. also differs from H. changningensis by having dark transverse blotches on the dorsum, while such blotching is absent in the later; the absence of a dark reticulate dorsal pattern, present on the later; and having a fork-like dark-colored postsacral marking, that is absent in the later. It also differs from H. zalonicus by having more femoroprecloacal pores (20–23 versus 16–20); having dark transverse dorsal blotches, while these are absent in the later; having a fork-like dark-colored postsacral marking, that is absent or indistinct in the later (see Table 7 for comparative values).