A new species of gekkonid, Hemiphyllodactylus gengmaensis sp. nov., is described based on six specimens from Gengma Dai and Wa Autonomous County, Yunnan, China. The new species can be distinguished from its congeners by a significant genetic divergence of greater than 9.7% in the mitochondrial ND2 gene and a combination of the following characters: a maximum SVL of 43.24mm; 8 or 9 chin scales; six circumnasal scales; 2 or 3 internasal scales; 9–11 supralabial scales; 8 or 9 infralabial scales; 11–18 dorsal scales; 8–10 ventral scales; a manual lamellar formula of 5–5–5–4 or 5–6–5–4 and a pedal lamellar formula of 5–5–6–5; 20–25 precloacal and femoral pore-bearing scales contiguous in males; dark postorbital stripes or striping on body; dark dorsal transverse blotches present; and a brown postsacral mark bearing anteriorly projecting arms. The discovery of this new species brings the number of Hemiphyllodactylus species in China to 15.

Gengma Dai and Wa Autonomous County, Hemiphyllodactylus gengmaensis sp. nov., integrative taxonomy, molecular phylogeny, slender gecko

Species of the genus Hemiphyllodactylus Bleeker, 1860 are small nocturnal geckos (SVL < 63 mm) distributed in South Asia, Southeast Asia, South China, and the Indo-Pacific region (Zug 2010; Grismer et al. 2013, 2018a; Agarwal et al. 2019; Eliades et al. 2019; Agung et al. 2021, 2022). Furthermore, all Hemiphyllodactylus are well camouflaged, occur in low densities, are forest-dwelling, and have small populations (Zug 2010; Grismer et al. 2013, 2018a; Agarwal et al. 2019; Eliades et al. 2019; Agung et al. 2021, 2022). Hence, Hemiphyllodactylus was considered to a low-diversity taxon (Zug 2010) until Grismer et al. (2013) revealed its high diversity using integrative taxonomy. In the following decade, the number of species within the group has increased from 12 to 54 (Uetz et al. 2023).

In China, Hemiphyllodactylus species have also been overlooked. The first species of slender geckos known from China is Gehyra yunnanensis (Boulenger, 1903), then “Smith revised its taxonomic status, placing it in the genus Hemiphyllodactylus” (Zug 2010). Zhou et al. (1981) recognized three subspecies of H. yunnanensis based on the digital lamellae patterns of specimens collected in Yunnan, Guizhou, and Guangxi Zhuang Autonomous Region, China: H. y. dushanensis Zhou & Liu, H. y. jinpingensis Zhou & Liu, and H. y. longlingensis Zhou & Liu. However, for the subsequent 32 years, most of Chinese slender geckos were regarded as simply H. yunnanensis, until Grismer et al. (2013) elevated these three subspecies to full species rank. Subsequently, nine additional species were incrementally recorded from China: H. zugi Nguyen, Lehmann, Le Duc, Duong, Bonkowski & Ziegler, 2013; H. changningensis Guo, Zhou, Yan & Li, 2015; H. huishuiensis Yan, Lin, Guo, Li & Zhou, 2016; H. hongkongensis Sung, Lee, Ng, Zhang & Yang, 2018; H. zayuensis Jiang, Wang & Che, 2020; H. dupanglingensis Zhang, Qian & Yang, 2020; H. zhutangxiangensis Agung, Grismer, Grismer, Quah, Chornelia, Lu & Hughes, 2021; H. simaoensis Agung, Chornelia, Grismer, Grismer, Quah, Lu, Tomlinson & Hughes, 2022; and H. yanshanensis Agung, Chornelia, Grismer, Grismer, Quah, Lu, Tomlinson & Hughes, 2022. The fourteenth species, H. typus Bleeker, 1860, as a widely distributed species, its actual coordinate data for distribution in China is unavailable (Agung et al. 2021; Uetz et al. 2023). According to recent genus-wide molecular phylogenetic studies, all Hemiphyllodactylus species in China belong to the typus group, and they are divided into four clades (Grismer et al. 2013, 2014a, 2014b, 2017, 2018a, 2020a, 2020b; Ngo et al. 2014; Agung et al. 2021, 2022): both clade 3 (H. longlingensis, H. zhutangxiangensis, H. zayuensis, and H. changningensis) and clade 4 (H. jinpingensis and H. simaoensis) of Agung et al. (2022) colonized China from western Indochina, and both clade 6 (H. dushanensis, H. hongkongensis, H. dupanglingensis, H. zugi, H. huishuiensis, and H. yanshanensis) and clade 7 (H. yunnanensis) of Agung et al. (2022) colonized China from eastern Indochina.

During our herpetological survey in Banxing Village, Gengma Dai and Wa Autonomous County, Yunnan, China, we collected six specimens belonging to the genus Hemiphyllodactylus. These specimens are distinguished from known species of Hemiphyllodactylus based on molecular and morphological data. Therefore, we describe them as a new species below.

Our results of ML and BI analyses were similar to those obtained by Agung et al. (2021): the specimens from Gengma County were recovered as members of Clade 3 in both ML and BI analyses (Figs 2, 3), which includes Hemiphyllodactylus longlingensis, H. zalonicus, H. changningensis, and H. zhutangxiangensis. The mean percentage of uncorrected pairwise distance between the Gengma County specimens and H. changningensis is 9.7% (Table 2). Furthermore, the new species also could be distinguished from its congeners by body proportions, CN, VS, Lamellar formulae hands and feet II–V, SL1T and total number of femoroprecloacal pores. Therefore, we describe them here as a new species.

Figure 2. 

Maximum-likelihood consensus tree based on 1038 bp mitochondrial ND2 gene. Numbers by the nodes indicate ML bootstrap support values.

Figure 3. 

Phylogenetic tree by Bayesian inference based on 1038 bp mitochondrial ND2 gene. Numbers by the nodes indicate posterior probability values of the BI.

Taxonomic account

 Hemiphyllodactylus gengmaensis sp. nov.

https://zoobank.org/16760DC6-5331-4742-A5FE-FD3F6E42DAC8

Figs 4, 5

Material

Holotype. 2014002302, adult female, collected by Hong Hui on 15 May 2014 from Banxing Village, Gengma Dai and Wa Autonomous County, Yunnan, China (23.517°N, 98.925°E, at an elevation of 664 m).

Figure 4. 

Hemiphyllodactylus gengmaensis sp. nov. A holotype, 2014002302, dorsal view in alcohol B holotype, 2014002302, ventral view in alcohol C paratype, 2014002301, dorsal view in alcohol.

Paratypes. 2014002297, 2014002298, 2014002299, adult females, 2014002300, 20140022301, adult males, collected at the same locality as the holotype on 15 May 2014.

Figure 5. 

Paratype (2014002301) of Hemiphyllodactylus gengmaensis sp. nov. A, B lateral views of head, red lines indicate IL and SL, respectively C ventral view, red lines indicate chin scales D ventral view, red lines indicate femoroprecloacal pores.

Diagnosis

Hemiphyllodactylus gengmaensis sp. nov. can be distinguished from its congeners by the combination of the following characters: maximum SVL of 43.24 mm; 8–9 chin scales; enlarged postmentals; 6 circumnasal scales; 2–3 internasal scales; 9–11 supralabial scales; 8–9 infralabial scales; 11–18 dorsal scales; 8–10 ventral scales; a manual lamellar formula of 5–5–5–4 or 5–6–5–4 and a pedal lamellar formula of 5–5–6–5; 20–25 precloacal and femoral pore-bearing scales contiguous in males. dark postorbital stripes or striping on body; dark dorsal transverse blotches; and a brown postsacral mark bearing anteriorly projecting arms.

Description of holotype

Adult female, SVL 38.52 mm; head triangular in dorsal profile, depressed, distinct from neck (HL 10.80 mm; HW 7.36 mm); lores flat; snout short (SnEye 3.94 mm; SnEye/HL 36%), narrow (SnW 1.78 mm; SnW/HW 24%); eye large (ED 2.12 mm; ED/HL 20%); rostral scale wider than high, bordered posteriorly by two large supranasals and three internasals (IS); nares bordered anteriorly by rostral scale, ventrally by first and second supralabial scale, dorsally by supranasal scale, posteriorly by three postnasals; supralabials square, 10/9 (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, eight chin scales touching internal edges of infralabials from juncture of the second and the third 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 45%), dorsoventrally compressed; dorsal body scales small, granular, 18 dorsal scales at midbody contained within one eye diameter; ventral body scales smooth and flat, much larger than dorsal scales, subimbricate, 10 ventral scales at midbody contained within one eye diameter; granular scales on the limbs; finger I is vestigial, clawless, and with rectangular subdigital lamellae, while fingers II–V are well developed; the proximal subdigital lamellae are undivided and rectangular, while the distal subdigital lamellae are divided, angular, U-shaped, except for the terminal lamellae, which are rounded and undivided; the forefoot and hindfoot digital formulae are unidentifiable; white cloacal spur present, one on each side. Tail length (TL/SVL = 73%), with dorsal scales on the tail larger than those on the body and head, but smaller than the subcaudals. The ventral scales are large and flat.

Coloration in ethanol

The dorsal surface of head and body is light brown; dark brown stripes extend from the posterior corner of the eye socket to the neck; the back is covered with irregular, dark-brown stripes that interconnect to form a net-like pattern; the dorsal surfaces of the limbs are brown, with irregular, dark-brown stripes; the tail is brown, with several dark-brown, transverse stripes; the regenerated tail is gray and without transverse stripes; the ventral surfaces of the head and limbs are cream-grey.

Variation

Variation of mensural and meristic data are presented in Table 3. Dark dorsal transverse blotches on the body of this species are relatively small, with those of two specimens (2014002302, 2014002300) being indistinct and fragmented. Furthermore, females are slightly larger than males. The postsacral mark, bearing anteriorly projecting arms, of one individual (2014002300) is indistinct, possibly due to prolonged preservation.

Table 3.

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Mensural (in mm), meristic, color pattern, and proportions of the type series of Hemiphyllodactylus gengmaensis sp. nov. (–) = data unavailable. (*) = regenerated tail.

Character	Holotype	Paratypes
	2014002302	2014002300	2014002299	2014002297	2014002298	2014002301
Sex	Female	Male	Female	Female	Female	Male
SVL	38.52	35.24	36.5	40.86	43.24	39.45
TL	28.22	–	9.18*	–	–	15.34*
TrunkL	17.46	17.54	17.6	21.6	23.22	18.82
HL	10.80	10.80	10.24	11.24	11.14	11.24
HW	7.36	6.94	6.88	7.86	8.2	7.74
ED	2.12	2.56	2.16	2.62	2.6	2.34
SnEye	3.94	3.88	4.02	4.5	4.52	4.3
NarEye	2.78	2.74	2.96	3.04	3.32	2.72
SnW	1.78	1.78	1.6	1.84	2.72	1.7
TrunkL/SVL	0.45	0.50	0.48	0.53	0.54	0.48
HL/SVL	0.28	0.31	0.28	0.28	0.26	0.28
HW/SVL	0.19	0.20	0.19	0.19	0.19	0.20
HW/HL	0.68	0.64	0.67	0.70	0.74	0.69
SnEye/HL	0.36	0.36	0.39	0.40	0.41	0.38
NarEye/HL	0.26	0.25	0.29	0.27	0.30	0.24
ED/HL	0.20	0.24	0.21	0.23	0.23	0.21
SnW/HL	0.16	0.16	0.16	0.16	0.24	0.15
ED/NarEye	0.76	0.93	0.73	0.86	0.78	0.86
Snw/HW	0.24	0.26	0.23	0.23	0.33	0.22
Chin	8	9	8	9	9	8
CN	6	6	6	6	6	6
IS	3	2	2	3	3	2
SL (left/right)	10/9	10/10	10/11	10/9	10/9	10/10
IL (left/right)	9/9	9/9	8/8	9/9	9/9	9/9
VS	10	9	9	8	8	9
DS	18	14	11	15	15	16
Lamellar formulae hands II–V (left)	–	–	–	–	5–5–5–4	–
Lamellar formulae hands II–V (right)	–	–	–	–	5–6–5–4	–
Lamellar formulae foot II–V (left)	–	–	–	–	5–5–6–5	–
Lamellar formulae foot II–V (right)	–	–	–	–	5–5–6–5	–
SL1F	–	–	–	–	5	–
SL1T	–	–	–	–	6	–
Precloacal and femoral pore series separate (1) or continuous (0)	–	0	–	–	–	0
Total femoroprecloacal pores	0	20	0	0	0	25
CloacS on each side	1	1	1	1	2	1
Subcaudals enlarged, plate–like	No	No	No	No	No	No
Dark postorbital stripe	Yes	Yes	Yes	Yes	Yes	Yes
Dorsolateral light–colored spots on trunk	No	No	No	No	No	No
Dark dorsolateral stripe on trunk	No	No	No	No	No	No
Dark ventrolateral stripe on trunk	No	No	No	No	No	No
Dark dorsal transverse blotches	Indistinct	Indistinct	Yes	Yes	Yes	Yes
Dark reticulate pattern on dorsum	Indistinct	Yes	Yes	Indistinct	Yes	Yes
Postsacral marking anteriorly projecting arms	Yes	indistinct	Yes	Yes	Yes	Yes

Distribution

This species is currently known to be distributed at the type locality Banxing Village, Gengma Dai and Wa Autonomous County of Yunnan Province in China (Fig. 1).

Natural History

Hemiphyllodactylus gengmaensis sp. nov. was found at an elevation of 664 m a.s.l., around 21:00. The specimens were found on a restaurant’s wall, which was rough and with crevices. When illuminated with a flashlight, the animals quickly crawled into the crevices.

Etymology

The scientific name “gengmaensis” is derived from its type locality Gengma Dai and Wa Autonomous County in Yunnan province. We suggest Gengma Slender Gecko in English and “耿马半叶趾虎 (Gěng Mǎ Bàn Yè Zhǐ Hǔ)” in Chinese.

Comparisons

We compared the morphology of the new species against its closely related species, specifically species from clade 3, as inferred from the phylogeny we constructed (Table 4). In terms of body proportions, H. gengmaensis sp. nov. has a longer head which separates it from H. longlingensis, H. zalonicus, H. changningensis, and H. zhutangxiangensis (HL/SVL = 0.26–0.31 versus 0.22–0.24, 0.22–0.23, 0.22–0.25, 0.17–0.20, respectively); greater SnW distance (SnW/HW = 0.22–0.33 versus 0.15–0.18, 0.21, 0.16–0.20, 0.16–0.21, respectively); the new species has a shorter SnEye distance which separates it from H. longlingensis, H. changningensis, and H. zhutangxiangensis (SnEye/HL = 0.36–0.41 versus 0.42–0.45, 0.41–0.49, 0.53–0.60, respectively); shorter head width (HW/HL = 0.64–0.74 versus 0.75–0.80, 0.72–0.80, 1.03–1.13, respectively); it has a shorter NarEye compared to H. changningensis and H. zhutangxiangensis (NarEye/HL = 0.24–0.30 versus 0.30–0.37, 0.39–0.4, respectively); smaller eyes compared to H. zalonicus and H. zhutangxiangensis (ED/HL = 0.20–0.24 versus 0.23–0.30, 0.30–0.36, respectively). In terms of scalation, the new species has more CN can be distinguished from H. longlingensis, H. zalonicus, H. changningensis, and H. zhutangxiangensis (CN = 6 versus 4–5, 5, 3–4, 5, respectively); more VS compared to H. longlingensis, H. changningensis, and H. zhutangxiangensis (VS = 8–10 versus 6–7, 6–8, 5–7, respectively); the new species has more femoroprecloacal pores which separates it from H. zalonicus, H. changningensis, and H. zhutangxiangensis (20–25 versus 16–20, 19–22, 20–23, respectively). For the lamellar and coloration, the new species differs from H. longlingensis, H. zalonicus, and H. changningensis by having more lamellae on the hand (5–5(6)–5–4 versus 3–4–4–4(3), 3–3(4)–3(4)–3(4) and 3–3(4)–3(4)–3, respectively); differs from H. zalonicus, H. changningensis, and H. zhutangxiangensis by having more lamellae on first fingers (SL1T = 6 versus 4, 3–4, 4–5, respectively). Furthermore, H. gengmaensis sp. nov. has dark transverse blotches on the dorsum, which H. zalonicus does not have. The new species has a dark, reticulate dorsal pattern, which H. zalonicus and H. zhutangxiangensis do not have. It has a postsacral marking with anteriorly projecting arms, which is absent in H. zalonicus and H. changningensis.

Table 4.

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Diagnostic characters separating Hemiphyllodactylus gengmaensis sp. nov. from other nominal taxa of Hemiphyllodactylus within clade 3 of Agung et al. (2022). (—) = data unavailable. Mensural characters are in mm. Data for H. zalonicus and H. longlingensis were obtained from Grismer et al. (2020a). Data for H. changningensis were obtained from Guo et al. (2015). Data for H. zhutangxiangensis were obtained from Agung et al. (2021).

Character	H. gengmaensis sp. nov.	H. longlingensis	H. zalonicus	H. changningensis	H. zhutangxiangensis
Max SVL	43.24	45.7	37.7	43.8	44.42
n	6	–	2	10	13
TrunkL	17.46–23.22	–	18.1–18.9	17.4–22.5	16.1–23.1
HL	10.8–11.24	–	8.4–8.5	8.2–10.1	6.2–7.6
HW	6.88–8.2	–	5.7–5.8	6.1–7.5	6.5–8.2
ED	2.12–2.62	–	2.0–2.6	1.7–2.3	2.1–2.7
SnEye	3.88–4.52	–	3.4–3.5	3.5–4.5	3.4–4.4
NarEye	2.72–3.32	–	2.4–2.7	2.8–3.4	2.6–3.2
SnW	1.6–2.72	–	1.2	1.1–1.4	1.3–1.6
TrunkL/SVL	0.45–0.54	0.47–0.52	0.49–0.50	0.46–0.51	0.48–0.52
HL/SVL	0.26–0.31	0.22–0.24	0.22–0.23	0.22–0.25	0.17–0.20
HW/SVL	0.19–0.2	0.17–0.19	0.15	0.17–0.18	0.18–0.20
HW/HL	0.64–0.74	0.75–0.80	0.68	0.72–0.80	1.03–1.13
SnEye/HL	0.36–0.41	0.42–0.45	0.40–0.42	0.41–0.49	0.53–0.60
NarEye/HL	0.24–0.30	0.29–0.34	0.28–0.31	0.30–0.37	0.39–0.44
ED/HL	0.20–0.24	0.22–0.25	0.23–0.30	0.21–0.25	0.30–0.36
SnW/HL	0.15–0.24	0.12–0.14	0.14	0.12–0.16	0.19–0.22
ED/NarEye	0.73–0.94	0.66–0.82	0.74–1.08	0.61–0.77	0.70–0.91
Snw/HW	0.22–0.33	0.15–0.18	0.21	0.16–0.20	0.16–0.21
Chin	8–9	7–9	8–10	7–8	7–9
CN	6	4–5	5	3–4	5
IS	2–3	1–3	3–4	2–3	2–4
SL	9–11	9–10	10	8–11	8–11
IL	8–9	8–10	8–9	8–10	8–11
VS	8–10	6–7	9–10	6–8	5–7
DS	11–18	10–14	17–18	11–15	11–15
Lamellar formulae hands II–V	5–5(6)–5–4	3–4–4–4(3)	3–3–3–3	3–3(4)–3(4)–3	(3–5)–(4–6)–(4 or 5)–(4 or 5)
		4–4–4(5)–4
Lamellar formulae feet II–V	5–5–6–5	4–4(5)–4(5)–4	3–4–4–4	3–3–3–3	(4 or 5)–(4 or 5)–(4–6)–(4 or 5)
				3–4–4–4
SL1F	5	4–5	3	3–4	4–5
SL1T	6	4–6	4	3–4	4–5
Precloacal and femoral pore series separate (1) or continuous (0)	0	0	0	0	0
Total femoroprecloacal pores	20–25	16–27	16–20	19–22	20–23
CloacS on each side	1 or 2	1 or 2	1	1 or 2	1 or 2
Subcaudals enlarged, plate–like	No	No	No	No	No
Dark postorbital stripe	Yes	Yes	Yes	Yes	Yes
Dorsolateral light–colored spots on trunk	No	No	No	No	No
Dark dorsolateral stripe on trunk	No	No	No	No	No
Dark ventrolateral stripe on trunk	No	No	No	No	No
Dark dorsal transverse blotches	Yes or indistinct	Variable	No	Yes	Yes
Dark reticulate pattern on dorsum	Yes or indistinct	Variable	No	Yes	No
Postsacral marking anteriorly projecting arms	Yes or indistinct	Pale colored	Indistinct or not	No	Fork-like, dark colored

Our research supports the recognition of the Hemiphyllodactylus gengmaensis sp. nov. as a new species, belonging to clade 3 of Agung et al. (2021). It is sister taxa to H. longlingensis, H. zalonicus, H. changningensis, and H. zhutangxiangensis. Except for H. zalonicus, all species of clade 3 occur in China. Considering the 292 km of unexplored area between H. zalonicus and its closest distance species (H. longlingensis), there may be numerous undescribed species in northern Myanmar and Dehong Dai and Jingpo Autonomous Prefecture, Yunnan Province, China. The 56 km distance between H. gengmaensis sp. nov. and H. changningensis, along with recent new species discoveries (Grismer et al. 2013; Guo et al. 2015; Che et al. 2020; Agung et al. 2021, 2022), suggests a high diversity of this genus within Yunnan. The border area of Dali Bai Autonomous Prefecture, Pu’er City, and Lincang City in southern Yunnan Province could be a key area for future research.

Our study increases the number of recognized species in the Hemiphyllodactylus in China to 15. Apart from H. typus, H. dupanglingensis, and H. hongkongensis, all other species in the genus Hemiphyllodactylus are montane species. Hemiphyllodactylus gengmaensis sp. nov. was found at an elevation of 664 m a.s.l. The discovery of the new species may represent an intermediate elevation type given by the known ranges, suggesting that species in the Hemiphyllodactylus may have a wide distribution range in southern China, spanning elevations from 120 m (Sung et al. 2018) to 2,169 m (unpublished data). Additionally, Grismer et al. (2018b) discovered 12 new gecko species within two weeks in a single study of karsts in Myanmar, with similar climatic and habitat conditions likely to exist in southern China. Moreover, some populations previously considered as H. yunnanensis in China have also been described as new species (Deng et al. 1998; Shi et al. 2011; Che et al. 2020; Zhang B et al. 2020). Therefore, it is possible that there are still numerous undiscovered cryptic species in southern China.

We thank Hong Hui for assistance in the field. We also thank the editors and reviews for their comments on the manuscript.