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Research Article
Discovery of a new species of the subgenus Japonigekko (Squamata, Gekkonidae, Gekko) from the Hengduan Mountains, southwestern China: the best Japonigekko mountaineer
expand article infoShun Ma‡§, Sheng-Chao Shi‡|, Cheng Shen‡, Li-Ming Chang‡, Jian-Ping Jiang‡§
‡ Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu, China
§ University of Chinese Academy of Science, Beijing, China
| Jianghan University, Wuhan, China
¶ Mangkang Biodiversity and Ecological Station, Xizang Ecological Safety Monitor Network, Changdu, China

Corresponding author: Jian-Ping Jiang ( jiangjp@cib.ac.cn )

Academic editor: Thomas Ziegler
© 2024 Shun Ma, Sheng-Chao Shi, Cheng Shen, Li-Ming Chang, Jian-Ping Jiang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Ma S, Shi S-C, Shen C, Chang L-M, Jiang J-P (2024) Discovery of a new species of the subgenus Japonigekko (Squamata, Gekkonidae, Gekko) from the Hengduan Mountains, southwestern China: the best Japonigekko mountaineer. ZooKeys 1215: 289-309. https://doi.org/10.3897/zookeys.1215.125043
ZooBank: urn:lsid:zoobank.org:pub:5EAB81EF-326D-4884-B046-5ECEE3DAEC26
Open Access

Abstract

A new Gekko (subgenus Japonigekko) species, Gekko alpinus sp. nov., is described from the Jinsha River Basin in southwestern China, between the border of Mangkang County, Xizang Autonomous Region and Batang County, Sichuan Province, according to the integrative taxonomic results combining molecular data and morphological characters obtained from the type series comprising 11 specimens. Our molecular phylogeny inferred from the mitochondrial 16S and ND2 gene fragments indicated that this new species is most closely related to Gekko jinjiangensis, but a considerable amount of genetic divergence exists between them (p-distance: 3.6%-4.1% (16S) and 7.1%–9.1% (ND2)). The new species can be distinguished from its congeners via a combination of series morphological characters. The discovery of this new species marks the highest altitudinal range (2400 to 2542 m a.s.l.) recorded for the subgenus Japonigekko and also represents a new provincial record for the genus in Xizang Autonomous Region.

Key words

Gekko alpinus sp. nov., Gekko jinjiangensis, Gekkonidae, molecular phylogeny, morphological characters, new provincial genus record

Introduction

The gekkonid genus Gekko Laurenti, 1768 is widely distributed across eastern and southeastern Asia, Northwest Oceania, and Melanesia. Currently, this genus contains 88 known gecko species and has been divided into seven subgenera: Archipelagekko, Balawangekko, Gekko, Japonigekko, Lomatodactylus, Ptychozoon, and Rhacogekko (Wood et al. 2020; Wang et al. 2024; Uetz et al. 2024). Amongst these, Japonigekko comprises 33 species, accounting for one-third of the species within the genus. Members of this subgenus are distributed across East Asia, with the majority found in China (20/33), of which the members present the following characters: size moderate; nare usually touching rostral; nasals two or three; dorsal tubercles 0–21 rows; precloacal pores 0–32; postcloacal tubercles 1–4; lateral folds without tubercles (Wood et al. 2020; Hou et al. 2021; Wang et al. 2024; Uetz et al. 2024).

During our field work in the Jinsha River Basin, along the border between Mangkang County of Xizang Autonomous Region and Batang County of Sichuan Province, China, a series of Gekko (Japonigekko) specimens was collected (Fig. 1). This discovery marks the first recorded occurrence of genus Gekko in Xizang Autonomous Region (Che et al. 2020). Phylogenetic analysis revealed a significant genetic differentiation between these specimens and their closest relative, G. jinjiangensis Hou, Shi, Wang, Jiang & Xie, 2021. Upon closer examination, we found that these specimens are morphologically distinct from G. jinjiangensis by having a relatively narrower head, more supralabials and infralabials, more interorbitals and dorsal tubercle rows at the midbody in females, fewer scales in a line from the mental to the front of the cloacal slit, and fewer scale rows at the mid-body. Hence, we describe these specimens as a new species.

Figure 1. 

Distribution of Gekko alpinus sp. nov. and its sister taxon G. jinjiangensis.

Materials and methods

Specimen preparation

A total of 11 specimens, two specimens (one adult male and one adult female) from Zhubalong Village, Mangkang County, Xizang Autonomous Region collected in July, 2022, and nine specimens (three adult males, four adult females and two subadult females) from Zhubalong Village, Batang County, Sichuan Province in June, 2020 (Fig. 1), were preserved in 75% ethanol and deposited in Chengdu Institute of Biology, Chinese Academy of Sciences (CIB, CAS), with their liver tissue samples separately preserved in 95% ethanol for molecular analyses.

Molecular data and phylogenetic analysis

Total genomic DNA was extracted by Vazyme FastPure Blood/Cell/Tissue/Bacteria DNA Isolation Mini Kit (Vazyme Biotech Co., Ltd, Nanjing, China) from the liver tissue samples of each specimen. Two mitochondrial gene fragments of partial 16S ribosomal RNA gene (16S) and partial NADH dehydrogenase subunit 2 gene (ND2) were respectively amplified by primers L3975 (5’-CGCCTGTTTACCAAAAACAT-3’) and H4551 (5’-CCGGTCTGAACTCAGATCACGT-3’) for 16S (Simon et al. 1994), and rMet-3L (5’-ATACCCCGACAATGTTGG-3’) and rAla-1H (5’-GCCTTAGCTTAATTAAAGTG-3’) for ND2 (Jonniaux and Kumazawa 2008). The polymerase chain reaction (PCR) was performed in 25 μl reactant with the following cycling conditions: first an initial denaturing step at 95 °C for 5 min; then 35 cycles of denaturing at 95 °C for 40 s, annealing at 53 °C for 40 s and extending at 72 °C for 60 s; last a final extending step at 72 °C for 10 min. PCR products were sequenced by Beijing Qingke New Industry Biotechnology Co., Ltd.

For our phylogenetic analysis, DNA sequences of 49 specimens were used (Table 1), amongst which ND2 (of specimens No. 1–11, 49) and 16S (of specimens No. 1–11, 13–17, 44, 49) were sequenced in this study and others were obtained from GenBank. Gehyra mutilata (Wiegmann, 1834) (No. 49) was used as the outgroup (Pyron et al. 2013). All 16S (569 bp) and ND2 (1011 bp) sequences were input in MEGA11 (Tamura et al. 2021) respectively and aligned by MUSCLE (Edgar 2004). Then we calculated the uncorrected pairwise distances (p-distance) for each data matrix in MEGA11. Then the concatenation sequences (1580 bp) were prepared for the phylogenetic analysis. The Maximum Likelihood (ML) analysis was performed in IQ-TREE 1.6.12 (Nguyen et al. 2015) based on the best-fit model TIM2+F+I+G4 for the 1st and 3rd codons of ND2 and all three codons of 16S and HKY+F+I+G4 for the 2nd codon of ND2 were computed by ModelFinder for IQ-Tree in PhyloSuite 1.2.3 according to Bayesian Information Criterion (BIC) (Kalyaanamoorthy et al. 2017; Zhang et al. 2020). Ultrafast Bootstrap Approximation (UFB) nodal support was assessed via using ten thousand ultrafast bootstrap replicates and when the value (UFB, %) is ≥ 95, it would be considered as significantly supported (Hoang et al. 2018). The single branch tests were conducted by SH-like approximate likelihood ratio test (SH-aLRT) by 1000 replicates and when the nodal support (SH, %) is ≥ 80, it would also be considered well supported (Stephane et al. 2010). The Bayesian Inference (BI) analysis was conducted via MrBayes 3.2.1 (Ronquist et al. 2012) under the best-fit model GTR+F+I+G4 for the 1st and 3rd codons of ND2 and all three codons of 16S and HKY+F+I+G4 for the 2nd codon of ND2, which was calculated according to BIC as well by ModelFinder for MrBayes in PhyloSuite 1.2.3. The BI analysis program worked through two independent runs with a four-chain run calculated for 20 million generations using the Markov Chain Monte Carlo (MCMC), sampling every 1000 with the first 25% of samples discarded as burn-in and resulting in a potential scale reduction factor (PSRF) of < 0.005. The nodal support Bayesian posterior probabilities (BI, %) ≥ 95 were considered significantly supported.

Table 1.
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Information and references for 16S and ND2 used in this study.

No. Species Localities Voucher ID. 16S GenBank Accession No. ND2 GenBank Accession No. Reference
1 Gekko alpinus sp. nov. Mangkang, Xizang, China CIB 121656 PQ255976 PQ303494 This study
2 CIB 121657 PQ255977 PQ303495
3 Batang, Sichuan, China CIB 121658 PQ255978 PQ303496
4 CIB 121659 PQ255979 PQ303497
5 CIB 121660 PQ255980 PQ303498
6 CIB 121661 PQ255981 PQ303499
7 CIB 121662 PQ255982 PQ303500
8 CIB 121663 PQ255983 PQ303501
9 CIB 121664 PQ255984 PQ303502
10 CIB 121665 PQ255985 PQ303503
11 CIB 121666 PQ255986 PQ303504
12 G. jinjiangensis Deqin, Yunan, China CIB 5334220115 – MT449431 Hou et al. 2021, this study
13 CIB 5334220088 PQ255987 MT449432
14 CIB 5334220089 PQ255988 MT449433
15 CIB 5334220090 PQ255989 MT449434
16 CIB 5334220100 PQ255990 MT449435
17 CIB 5334220114 PQ255991 MT449436
18 Derong, Sichuan, China CIB 5133380017 – MT449437
19 CIB 5133380019 – MT449438
20 CIB 5133380021 – MT449439
21 CIB 5133380024 – MT449440
22 CIB 5133380025 – MT449441
23 CIB 5133380026 – MT449442
24 CIB 5133380047 – MT449443
25 G. adleri Jingxi, Guangxi, China SYS r001400 MW451654 OR902178 Lyu et al. 2021; Wang et al. 2024
26 G. auriverrucosus Yuncheng, Shanxi, China NNU Z20050801.004 – JN019062 Rösler et al. 2011
27 G. bonkowskii Khammouane, Laos VFU R.2014.10 – KT266818 Luu et al. 2015
28 G. chinensis Hong Kong, China SYS r001211 MW451644 OR902183 Lyu et al. 2021; Wang et al. 2024
29 G. cib Chengdu, Sichuan, China AMB 6567 – JN019063 Rösler et al. 2011
30 G. cib Hejiang, Sichuan, China SYS r001489 MW451655 OR902165 Lyu et al. 2021; Wang et al. 2024
31 G. hokouensis Jinzhai, Anhui, China NNU Z20050902.001 – JN019060 Rösler et al. 2011
32 G. hokouensis Wuyishan, Fujian, China SYS r001290 MW451647 OR902173 Lyu et al. 2021; Wang et al. 2024
33 G. japonicus Zhoushan, Zhejiang, China NNU Z20050801.004 – JN019059 Rösler et al. 2011
34 G. japonicus Wuyishan, Fujian, China SYS r000672 MW451628 OR902176 Lyu et al. 2021; Wang et al. 2024
35 G. khunkhamensis Khammouane, Laos VNUF R.2021.23 – OL416111 Sitthivong et al. 2021
36 G. kwangsiensis Wuming, Guangxi, China SYSr 001195 MW451642 OR902175 Lyu et al. 2021; Wang et al. 2024
37 G. melli Dongguan, Guangdong, China SYS r001742 MW451661 OR902169 Lyu et al. 2021; Wang et al. 2024
38 G. nadenensis Khammouane, Laos ZFMK 98741 – KY421618 Luu et al. 2017
39 G. palmatus Zhaoqing, Guangdong, China SYS r002797 OR903156 OR902179 Wang et al. 2024
40 G. paucituberculatus Baise, Guangxi, China SYS r002806 OR903154 OR902163 Wang et al. 2024
41 G. scientiadventura Quang Binh, Vietnam IEBR A.2014.7 – KP205392 Luu et al. 2014
42 G. sengchanthavongi Khammouane, Laos VFU R2014.14 – KT266816 Luu et al. 2015
43 G. similignum Wuzhishan, Hainan, China SYS r001597 MW451658 OR902185 Lyu et al. 2021; Wang et al. 2024
44 G. scabridus Yanbian, Sichuan, China CIB YN201909199 PQ255992 MT449429 Hou et al. 2021; this study
45 G. subpalmatus Fenghua, Zhejiang, China SYS r001762 MW451662 OR902167 Lyu et al. 2021; Wang et al. 2024
46 G. swinhonis Zunhua, Hebei, China SYS r001814 MW451666 OR902171 Lyu et al. 2021; Wang et al. 2024
47 G. thakhekensis Thakhek, Khammouane, Laos IEBR A.2014.6 – KP205396 Luu et al. 2014
48 G. truongi Khanh Hoa, Vietnam IEBR A.2011.1 – KP205398 Luu et al. 2014
49 Gehyra mutilata Xishuangbanna, Yunnan, China CIB R201711 PQ255993 PQ303505 this study

Morphological comparisons and statistical analysis

Morphological data were obtained from the 11 Gekko alpinus sp. nov. (four males, four adult females, and three subadult females). The terminology and methods of mensural characters and meristic features followed Zhao et al. (1999) and Rösler et al. (2011). Bilateral morphological characters measurements and scale counts were given as left/right.

The mensural characters were measured to the nearest 0.01 mm using a Deli Caliper (DL92150): (1) snout-vent length (SVL: from tip of snout to anterior margin of cloaca); (2) tail length (TaL: from posterior margin of cloaca to tip of tail); (3) axilla-groin distance (AGD: distance between axilla and groin); (4) head length (HL: maximum head length from tip of snout to posterior margin of auricular opening); (5) head width (HW: maximum head width measured at the angle of the jaws); (6) head height (HH: maximum head height from the top of the head posterior to the eyes to the bottom of the lower jaw); (7) snout length (SL: from snout tip to anterior corner of eye); (8) eye-ear distance (EED: distance between posterior margin of eye to posterior margin of ear opening) (9) maximum eye diameter (ED); (10) maximum ear opening diameter (EOD); (11) maximum rostral width (RW); (12) maximum rostral height (RH); (13) maximum mental width (MW); (14) maximum mental length (ML); (15) forelimb length (FlL: length from the base of the palm to the elbow); (16) hindlimb length (HlL: distance from the base of heel to the knee).

All mensural characters except for TaL, FIL, and HIL, which were lacking for G. jinjiangensis, were statistically analyzed using R v. 4.3.2, and sexes were separated for subsequent comparisons among the samples due to sexual dimorphism within geckos. For analyses, all measurements were ln-transformed to normalize and reduce the variance, and then scaled to remove allometric effects of body size using the following equation: Xa = Xln–β ∙ (SVLln–SVLm), where Xa = adjusted value; Xln = ln-transformed measurements; β = unstandardized regression coefficient for each species; SVLln = ln-transformed SVL; and SVLm = overall average SVLln of all samples. This project was performed under GroupStruct R package (Thorpe 1975, 1976, 1983; Reist 1985; Lleonart et al. 2000; Chan and Grismer 2021). Principal component analysis (PCA) was performed to cluster the morphometrics except SVL, TaL, FIL, HIL related to each species using prcomp R function and factoextra R package.

The meristic features were taken as the followings: (1) supralabials (SPL: number of scales from commissure of jaw to the rostral scale); (2) infralabials (IFL: number of scales from commissure of jaw to the mental scale); (3) interorbitals (IO: number of scales in a line between anterior corners of eyes); (4) postmentals (PM: scales bordering the mental); (5) dorsal tubercles row at midbody (DTR); (6) scales in a line from mental to the front of cloacal slit (SMC); (7) scale rows at midbody (SR); (8) ventral scales at midbody from one ventrolateral fold to the other (V); (9) subdigital lamellae under entire first finger (LF1); (10) subdigital lamellae under entire fourth finger (LF4); (11) subdigital lamellae under entire first toe (LT1); (12) subdigital lamellae under entire fourth toe (LT4); (13) precloacal pores (PP); (14) postcloacal tubercles (PAT).

One-way analysis of variance (ANOVA) test was used to evaluate significant differences in the mensural and meristic characteristics between the newly collected specimens and G. jinjiangensis, with significant different variances (p-values < 0.05 in the Levene’s test) using the aov R function.

Morphological information of G. jinjiangensis were obtained from Hou et al. (2021), and for other species, morphological data were taken from the literature (Stejneger 1907; Zhou et al. 1982; Song 1985; Zhao et al. 1999; Goris and Maeda 2004; Rösler et al. 2005, 2011; Toda et al. 2008; Zhou and Wang 2008; Phung and Ziegler 2011; Nguyen et al. 2013; Luu et al. 2014; Ngo et al. 2015; Yang 2015; Luu et al. 2015, 2017; Hou et al. 2021; Lyu et al. 2021; Sitthivong et al. 2021; Zhang et al. 2023; Wang et al. 2024).

Results

Phylogenetic analysis

The new Gekko (Japonigekko) alpinus sp. nov. specimens formed a well-supported sister lineage (SH 100/UFB 100/BI 100) to G. jinjiangensis (SH 98/UFB 100/BI 100) with considerable evolutionary differentiation (Fig. 2). The uncorrected pairwise divergences amongst some species of the subgenus Japonigekko studied in this work inferred from the mitochondrial 16S/ND2 gene fragments range from 2.2% (G. chinensis (Gray, 1842) vs G. similignum Smith, 1923) / 5.4% (G. chinensis vs G. similignum) to 18.4% (G. chinensis vs G. swinhonis Günther, 1864, and G. similignum vs G. swinhonis) / 26.5% (G. melli (Vogt, 1922) vs G. similignum), while the genetic distances amongst Gekko alpinus sp. nov. with its congeners range from 3.6% (vs G. jinjiangensis) to 14.0% (vs G. swinhonis) for 16S and 7.1% (vs G. jinjiangensis) to 24.1% (vs G. similignum) for ND2 (Tables 2, 3), indicating that Gekko alpinus sp. nov. have distinct interspecific genetic differentiation from its congeners. Based on the molecular results, these Gekko alpinus sp. nov. are supported as representing a new taxon.

Figure 2. 

Maximum Likelihood tree topology of Japonigekko inferred from the concatenated 16S and ND2 gene fragments (1580 bp). The support values of each node present on the tree: SH / UFB / BI (the ones lower than 50 are displayed as “-”). The ID numbers of Gekko alpinus sp. nov. are noted in red.

Table 2.
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Uncorrected p-distance (%) of some species in the subgenus Japonigekko based on the partial mitochondrial 16S gene sequences. Numbers refer to specimens listed in Table 1.

Species 1–11 13–17 25 28 30 32 34 36 37 39 40 43 44 45
1–11 Gekko alpinus sp. nov. 0–0.5
13–17 G. jinjiangensis 3.6–4.1 0–0.6
25 G. adleri 12.6–13.5 12.9–13.3
28 G. chinensis 11.7–12.4 11.4–11.9 7.2
30 G. cib 8.9–9.4 9.2–9.8 13.5 11.9
32 G. hokouensis 11.3–12.2 9.8–10.5 13.6 12.9 13.3
34 G. japonicus 9.8–11.0 9.8–10.5 15.3 14.3 13.7 11.1
36 G. kwangsiensis 11.7–12.8 10.8–11.5 15.6 15.0 11.9 11.9 14.5
37 G. melli 8.9–9.4 9.8–10.4 14.2 13.8 6.0 12.1 13.0 15.0
39 G. palmatus 11.9–12.9 12.5–12.8 3.5 5.0 12.7 14.2 12.8 12.9 10.9
40 G. paucituberculatus 9.8–10.6 9.8–10.5 13.2 12.1 12.0 11.1 13.0 9.8 11.9 14.4
43 G. similignum 11.9–12.7 11.6–12.1 7.4 2.2 12.9 13.6 14.5 16.2 14.2 5.2 13.2
44 G. scabridus 4.7–5.5 4.7–5.4 12.8 12.1 9.7 10.2 10.4 12.4 10.3 11.7 9.7 12.8
45 G. subpalmatus 9.5–9.9 10.9–11.3 16.3 15.9 7.1 13.2 14.1 15.1 10.1 12.8 13.7 16.7 11.3
46 G. swinhonis 13.2–14.0 14.0–14.6 17.2 18.4 11.7 13.4 15.3 17.4 15.3 14.6 14.9 18.4 14.6 15.7
Table 3.
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Uncorrected p-distance (%) of some species in the subgenus Japonigekko based on the partial mitochondrial ND2 gene sequences. Numbers refer to specimens listed in Table 1.

Species 1–11 12–24 25 26 27 28 29–30 31–32 33–34 35 36 37 38 39 40 41 42 43 44 45 46 47 48
1–11 Gekko alpinus sp. nov. 0–2.9
12–24 G. jinjiangensis 7.1–9.1 0–3.5
25 G. adleri 22.8–23.8 20.3–21.6
26 G. auriverrucosus 21.2–22.9 20.1–20.8 24.9
27 G. bonkowskii 18.8–19.4 18.0–19.1 24.6 21.7
28 G. chinensis 21.8–23.3 20.7–22.2 14.5 24.4 21.2
29–30 G. cib 22.6–23.7 20.3–20.9 26.3 20.7 19.3 24.2 0
31–32 G. hokouensis 20.7–22.3 18.7–21.1 25.4–25.5 21.5–21.7 20.3 25.0 23.7–23.8 0.3
33–34 G. japonicus 19.8–21.2 17.0–18.1 26.2–26.4 21.0–21.2 19.7–19.9 25.0–25.2 22.8–22.9 22.6–23.0 0.1
35 G. khunkhamensis 21.3–21.7 20.9–21.7 26.0 24.3 15.2 24.6 21.1 24.2 22.6–22.8
36 G. kwangsiensis 22.0–23.5 19.2–19.6 24.3 22.5 21.0 23.2 20.6 21.7–21.8 22.8–23.1 21.5
37 G. melli 23.1–23.6 20.1–21.1 25.3 23.5 21.8 25.3 19.0 24.7 23.8–24.0 23.4 23.8
38 G. nadenensis 18.4–19.4 17.1–18.4 23.3 20.4 6.9 21.2 20.3 21.8–22.1 20.1–20.5 13.8 20.6 21.4
39 G. palmatus 21.4–22.8 21.1–21.6 6.5 23.9 23.6 14.5 26.3 24.6–24.7 25.0–25.2 26.2 23.6 26.1 23.3
40 G. paucituberculatus 20.0–21.7 17.5–18.3 25.5 21.2 19.1 25.6 21.0 21.7 21.6–21.8 21.5 18.9 25.3 18.6 24.6
41 G. scientiadventura 18.4–18.8 17.8–18.6 24.4 20.6 13.9 22.5 20.5 21.4–21.6 21.1–21.2 14.8 21.0 21.6 13.5 23.5 18.6
42 G. sengchanthavongi 19.8–20.3 19.1–19.7 23.8 20.2 14.3 22.1 21.6 22.5–22.9 21.0–21.3 15.7 21.6 22.5 12.0 23.5 18.9 10.5
43 G. similignum 22.8–24.1 21.1–22.6 15.0 25.4 23.1 5.4 25.5 25.1 25.1–25.2 25.0 23.7 26.5 22.7 14.4 25.1 23.1 22.7
44 G. scabridus 11.2–11.8 10.1–10.6 18.5 19.1 19.9 20.3 21.3 17.2 17.2–17.5 21.7 20.1 20.7 19.1 19.2 17.9 17.6 18.4 20.3
45 G. subpalmatus 21.9–23.1 19.2–20.7 25.1 21.8 20.6 24.2 18.0 23.3–23.5 23.3–23.5 22.6 22.3 18.5 18.8 25.6 22.4 20.5 20.6 25.2 20.0
46 G. swinhonis 22.0–23.4 19.4–20.7 25.3 20.8 21.2 25.9 22.0 22.4–22.6 21.9–22.0 23.6 21.8 23.6 21.4 24.5 23.9 22.5 22.1 25.6 19.8 21.7
47 G. thakhekensis 18.8–19.6 18.2–19.5 21.8 20.8 6.8 19.9 20.3 20.8 19.1–19.4 15.9 20.5 20.6 6.8 22.9 18.0 13.1 13.1 21.0 18.9 20.1 23.1
48 G. truongi 20.2–20.6 19.7–20.8 20.5 24.5 22.1 20.8 21.8 22.3 22.0–22.2 22.2 23.1 21.4 22.1 21.8 21.2 22.3 22.0 21.8 18.2 21.6 24.0 20.5

Morphological analysis

Morphological characters of 11 Gekko (Japonigekko) alpinus sp. nov. specimens are presented in Table 4, which can be easily distinguished from all other known congeners (Table 5). By tubercles existing on dorsal body, forelimbs, hindlimbs, and tails, Gekko alpinus sp. nov. can be distinguished from the following 26 species: G. aaronbaueri Tri, Thai, Phimvohan, David & Teynié, 2015; G. adleri Nguyen, Wang, Yang, Lehmann, Le, Ziegler & Bonkowski, 2013; G. bonkowskii Luu, Calame, Nguyen, Le & Ziegler, 2015; G. canhi Rösler, Nguyen, Van, Doan, Ho, Nguyen & Ziegler, 2010; G. chinensis; G. cib Lyu, Lin, Ren, Jiang, Zhang, Qi & Wang, 2021; G. guishanicus Lin & Yao, 2016; G. hokouensis Pope, 1928; G. khunkhamensis Sitthivong, Lo, Nguyen, Ngo, Khotpathoom, Le, Ziegler & Luu, 2021; G. kwangsiensis Yang, 2015; G. liboensis Zhou, Liu & Li, 1982; G. melli; G. nadenensis Luu, Nguyen, Le, Bonkowski & Ziegler, 2017; G. palmatus Boulenger, 1907; G. paucituberculatus Wang, Qi, Zhou & Wang, 2024; G. scientiadventura Rösler, Ziegler, Vu, Herrmann & Böhme, 2004; G. sengchanthavongi Luu, Calame, Nguyen, Le & Ziegler, 2015; G. shibatai Toda, Sengoku, Hikida & Ota, 2008; G. similignum; G. subpalmatus (Günther, 1864); G. tawaensis Okada, 1956; G. thakhekensis Luu, Calame, Nguyen, Le, Bonkowski & Ziegler, 2014; G. truongi Phung & Ziegler, 2011; G. vertebralis Toda, Sengoku, Hikida & Ota, 2008; G. wenxianensis Zhou & Wang, 2008; G. yakuensis Matsui & Okada, 1968. By having 4–7 precloacal pores in the male, Gekko alpinus sp. nov. differs from G. kaiyai Zhang, Wu & Zhang, 2023 (9–12) and G. scabridus Liu & Zhou, 1982 (10–15). By having 13–15 subdigital lamellae on fourth toes, Gekko alpinus sp. nov. is different from G. swinhonis Günther, 1864 (6–9) and G. taibaiensis Song, 1985 (7–8). Gekko alpinus sp. nov. can be differed from G. japonicus (Schlegel, 1836) by having fewer interorbitals (IO 22–28 vs 32–35), fewer scale rows at midbody (SR 92–114 vs 130–144) and fewer ventral scales at midbody (32–39 vs 39–44).

Table 4.
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The measurements (in mm) and meristic characters of the type series of Gekko alpinus sp. nov. (“H/P” = holotype and paratype respectively; “F/M” = the gender female and male respectively; “#” = subadult; “*” = the length of regenerated tail; “-” = data unavailable; “+” = tail is broken).

ID CIB 121663 CIB 121661 CIB 121664 CIB 121658 CIB 121659 CIB 121662 CIB 121666 CIB 121660 CIB 121665 CIB 121656 CIB 121657 Range Mean ± SD
Type H P P P P P P P P P P
Sex M M M F F F F F# F# M F#
SVL 74.16 68.28 65.68 66.76 59.02 66.22 64.34 51.58 50.56 56.44 50.74 50.56–74.16 61.25 ± 7.67
TaL 68.02 54.52* 72.06 59.10 59.16* 61.18* 49.22* 59.32 20.28(+) 6.88(+) 39.46* 68.02–72.06 70.04 ± 2.02
AGD 27.70 32.34 30.62 34.78 21.16 29.88 31.44 18.62 24.46 24.24 23.72 18.62–34.78 27.18 ± 4.86
HL 18.44 17.82 17.22 20.80 16.12 17.42 15.84 15.24 14.92 17.24 19.92 14.92–20.80 17.36 ± 1.76
HW 13.26 12.62 12.68 13.24 11.42 12.56 12.56 10.34 10.28 12.22 10.18 10.18–13.26 11.94 ± 1.13
HH 6.78 6.74 6.58 5.76 5.14 5.38 5.72 5.12 4.34 7.10 5.44 4.34–7.10 5.83 ± 0.82
SL 7.62/7.68 7.12/7.22 6.98/7.16 7.42/7.64 6.72/6.72 7.06/7.16 6.90/6.92 5.82/5.86 5.72/5.74 6.90/6.96 5.78/5.80 5.72–7.68 6.77 ± 0.66
EED 5.72/5.66 6.26/6.30 5.62/5.52 5.64/5.72 5.02/5.04 5.78/5.72 5.36/5.46 4.24/4.38 4.64/4.62 5.76/5.76 4.42/4.42 4.24–6.30 5.32 ± 0.61
ED 4.36/4.26 3.96/3.98 3.84/3.76 3.94/3.96 3.36/3.38 3.98/3.94 3.26/3.28 2.96/3.02 2.90/2.94 4.24/4.22 3.38/3.40 2.90–4.36 3.65 ± 0.46
EOD 1.18/1.12 1.06/1.08 0.74/0.86 1.02/0.98 1.24/1.16 1.14/1.26 0.86/0.88 0.94/0.96 0.86/0.82 1.24/1.24 0.64/0.70 0.64–1.26 1.00 ± 0.18
RW 2.12 3.02 2.08 2.52 3.04 2.08 2.92 1.62 1.86 1.56 2.40 1.56–3.04 2.29 ± 0.51
RH 1.22 1.42 1.12 1.32 1.44 0.82 1.40 0.84 1.12 1.18 1.18 0.82–1.44 1.19 ± 0.20
MW 2.22 2.04 1.66 1.82 1.64 1.42 1.60 1.38 1.86 1.62 1.42 1.38–2.22 1.70 ± 0.25
ML 1.62 1.54 1.52 1.48 1.32 1.28 1.52 1.18 1.28 1.82 1.62 1.18–1.82 1.47 ± 0.18
FIL 6.86/6.70 7.52/7.34 7.58/7.64 7.58/7.74 6.82/6.88 7.42/7.56 6.42/6.36 5.82/5.98 6.32/6.02 7.98/8.00 6.14/6.16 5.82–8.00 6.95 ± 0.69
HIL 8.56/8.48 8.62/8.84 8.72/8.62 8.60/8.58 9.06/9.02 8.72/8.84 7.42/7.64 6.44/6.48 7.34/7.38 7.18/7.22 6.92/7.04 6.44–9.06 7.99 ± 0.85
SPL 11/11 10/9 10/10 9/9 10/10 11/12 10/10 10/11 10/10 13/11 10/10 9–13 10.32 ± 0.92
IFL 10/10 8/8 9/9 10/10 9/9 9/9 9/9 9/9 .10/9 9/10 9/9 8–10 9.18 ± 0.57
IO 27 23 25 24 28 23 28 22 25 24 25 22–28 25.00 ± 2.00
PM 2 2 2 2 2 2 2 2 2 2 2 2 2.00 ± 0
DTR 15 16 12 17 17 17 15 14 15 17 17 12–17 15.64 ± 1.55
SMC 165 175 173 158 162 162 179 163 169 189 181 158–189 170.55 ± 9.25
SR 114 109 112 106 101 101 104 101 92 98 95 92–114 103.00 ± 6.54
V 33 36 35 34 34 38 36 32 36 38 39 32–39 35.55 ± 2.10
LF1 10/10 10/11 10/10 11/11 10/10 11/10 10/9 9/9 9/9 8/9 9/9 8–11 9.45 ± 1.34
LT1 11/10 10/10 11/11 10/10 10/10 11/10 10/10 -/9 10/10 9/8 10/9 8–11 9.95 ± 0.72
LF4 13/13 13/12 14/13 12/13 14/14 12/13 12/13 13/13 14/12 12/12 13/13 12–14 12.86 ± 0.69
LT4 15/14 14/15 14/12 14/14 15/15 14/14 14/14 -/15 15/15 13/14 13/14 12–15 14.14 ± 0.77
PP 7 6 4 0 0 0 0 0 0 5 0 4–7 5.50 ± 1.12
PAT 1/1 1/1 1/1 2/2 1/1 2/2 1/1 1/1 1/1 2/2 1/1 1–2 1.30 ± 0.46
Table 5.
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Morphological characters of Japonigekko (“*” = species distributed in China; “-” = data unavailable; bold = difference between the new species).

No. Species SVLmax SPL IFL IO DTR SMC SR V LT1 LT4 Web Fore tubercles Hind tubercles Tail tubercles PP
1* Gekko alpinus sp. nov. 74.16 9–13 8–10 22–28 12–17 158–189 92–114 32–39 8–11 13–15 0 1 1 1 4–7
2 G. aaronbaueri 80 13–14 10–11 34–37 0 – 98–104 39–43 14–17 14–16 – 0 0 0 3–4
3* G. adleri 75.3 10–15 9–13 27–36 7–11 168–190 123–144 35–44 11–14 11–15 1 0 1 1 17–21
4* G. auriverrucosus 69 9–11 9–11 26–27 16–20 – – – 6–8 6–8 0 1 1 1 8–11
5 G. bonkowskii 69.2 12–14 10–11 49–50 0 154–169 117 37–40 11–13 15 1 0 0 0 6
6 G. canhi 99.2 14 10–12 49–50 11–12 168–170 205–227 49–51 13–16 14–17 0 0 1 0 5
7* G. chinensis 72 10–14 9–13 35–48 10 156–167 118–140 37–39 8–10 9–12 1 0 1 1 17–27
8* G. cib 66.4 10–12 10–14 28–36 0 171–196 128–149 37–45 9–13 9–17 1 0 0 0 7–9
9* G. guishanicus 64 – – – 0 – – – 8–10 8–10 0 0 0 0 6–8
10* G. hokouensis 70 10–14 8–11 30–33 12–18 153–174 119–130 36–43 8–11 15–18 0 0 0 1 5–9
11* G. japonicus 74 9–13 8–13 32–35 9–14 169–188 130–144 39–44 10–12 14–16 0 1 1 1 4–9
12* G. jinjiangensis 61.6 7–10 6–9 20–24 12–16 146–169 111–149 31–47 8–11 11–15 0 1 1 1 4–5
13* G. kaiyai 64.99 9–12 9–13 22–33 11–18 157–209 99–121 30–43 8–9 7–11 0 1 1 1 9–12
14 G. khunkhamensis 75.2 9–10 9–10 31–32 0 181–185 127–138 42–45 13–14 14–15 1 0 0 0 0
15* G. kwangsiensis 69.7 10–12 11–13 29–31 9–11 185–208 143–156 41–45 11–13 13–18 1 0 0 1 9–11
16* G. liboensis 85 12 11 40 10 – – – 8 9 0 0 0 – –
17* G. melli 80.3 10–13 9–12 34–40 0 171–192 148–160 44–46 10–12 11–14 1 0 0 0 9–11
18 G. nadenensis 77.1 12–14 10–12 28–30 0 175–185 123–140 38–40 13–15 14–16 1 0 0 0 6
19* G. palmatus 79.7 11–15 9–13 27–36 4–12 160–191 116–147 36–47 10–13 10–16 1 0 0 1 23–30
20* G. paucituberculatus 85.9 11 9–10 37 4 189–192 136–140 42–44 11 11–13 0 0 0 0 12
21 G. scientiadventura 73 12–14 9–13 41–51 0 118–140 139–143 38–48 12–15 14–17 1 0 0 1 23–30
22* G. scabridus 64 9–11 9–11 30 17–21 – – – 6–9 7–9 0 1 1 1 10–15
23 G. sengchanthavongi 77.3 8–10 6–7 28–32 0 175–184 120–135 35–43 11–14 13–17 1 0 0 0 4–5
24 G. shibatai 70.9 10–13 10–14 37–52 5–14 – 114–134 – – 9–16 0 0 0 1 0–3
25* G. similignum 58.9 12–14 11 46–48 11 – 144–153 – 11–13 12–14 1 0 0 1 17
26* G. subpalmatus 65.8 8–12 7–12 28–37 0 144–190 129–156 39–46 9–12 11–14 1 0 0 0 5–9
27* G. swinhonis 66 7–12 7–11 23–24 6–8 – – 40 6–9 6–9 0 1 1 – 7–9
28* G. taibaiensis 69 9–10 8–10 28 – – – – 6–7 7–8 – – – – 4–6
29 G. tawaensis 71 15 13 – 0 – – – 10 12 0 0 0 0 0
30 G. thakhekensis 79.2 12–14 10–11 22–26 0 165–174 110–116 32–40 11–13 14–15 1 0 0 0 1–5
31 G. truongi 95.9 13–15 11–13 45–48 0 160–172 131–143 35–36 11–13 15–17 0 0 0 0 10–11
32 G. vertebralis 69.2 10–15 10–15 35–50 2–12 – 112–139 – – 9–17 0 0 0 0 0–1
33* G. wenxianensis 59 12 11 – 10 – – 42–44 6 9 0 0 1 – 6–8
34 G. yakuensis 72 12–13 9–13 – – – – – 10 15 0 0 0 1 6–8

The results of the ANOVA indicated that Gekko alpinus sp. nov. is significantly different from its sister taxon G. jinjiangensis (Table 6) in the following characters: (1) male: HL, HW, SL, ED, SPL, IFL, SMC, SR, LF4; (2) female: AGD, HL, HW, HH, SL, EOD, SPL, IFL, IO, DTR, SMC, SR, LF4, and LT4. In the PCA analysis (Table 7), the first four principal components explained 97.96% of the total variation in the males, where PC1, PC2, PC3, and PC4 eigenvectors accounted for 45.77%, 20.84%, 17.51%, and 13.86% of the total variance respectively, and similarly, the first four principal components occupied a considerable proportion in the females, 76.82% of the total, whereas the PC1, PC2, PC3, and PC4 eigenvectors accounted for 33.42%, 18.13%, 15.47%, and 9.80% of the total variance respectively. As illustrated in the scatter plots of PC1 and PC2 (Fig. 3), regardless of sex, the samples of each species cluster together and do not overlap with each other. The results of the ANOVA and PCA indicated that Gekko alpinus sp. nov. was significantly different from the closely related G. jinjiangensis. As for morphological comparisons, Gekko alpinus sp. nov. can be distinguished from G. jinjiangensis by (1) relatively narrower head (HW/HL 0.72 ± 0.01 vs 0.83 ± 0.01 in males and 0.68 ± 0.08 vs 0.81 ± 0.05 in females); (2) more supralabials (SPL 10.63 ± 1.16 vs 8.00 ± 0.50 in males and 10.14 ± 0.74 vs 8.89 ± 0.50 in females) and infralabials (IFL 9.13 ± 0.83 vs 6.63 ± 0.48 in males and 9.21 ± 0.41 vs 7.33 ± 0.70 in females); (3) more interorbitals in females (IO 25 ± 2.14 vs 21.33 ± 1.63); (4) more dorsal tubercles row at midbody in females (DTR 16.00 ± 1.20 vs 13.67 ± 1.15); (5) fewer scales in a line from mental to the front of cloacal slit (SMC 175.50 ± 8.65 vs 155.25 ± 5.63 in males and 167.71 ± 8.34 vs 158.89 ± 6.61 in females); and fewer scale rows at midbody (SR 108.25 ± 6.18 vs 131.75 ± 10.03 in males and 100.00 ± 4.54 vs 123.78 ± 10.20 in females).

Figure 3. 

Principal component analysis performed for Gekko alpinus sp. nov. and G. jinjiangensis based on 12 commonly used morphological traits (except SVL, Tal, FIL, HIL). Numbers inside the brackets indicate the percentages of the total variance explained by each axis.

Table 6.
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Morphological comparisons of Gekko alpinus sp. nov. with G. jinjiangensis. “–” = data unavailable, “*” = p-values < 0.05, “**” = p-values < 0.01, “***” = p-values < 0.001.

Gekko alpinus sp. nov. n = 4 G. jinjiangensis n = 4 p-values Gekko alpinus sp. nov. n = 7 G. jinjiangensis n = 9 p-values
Range Mean ± SD (males) Range Mean ± SD (males) Range Mean ± SD (females) Range Mean ± SD (females)
SVL 56.44–74.16 66.14 ± 6.39 50.2–61.6 56.25 ± 4.26 0.0672 50.56–66.76 58.46 ± 6.90 54.6–61.5 56.57 ± 2.02 0.477
AGD 24.24–32.34 28.73 ± 3.08 22.1–24.7 23.83 ± 1.04 0.0547 18.62–34.78 26.29 ± 5.43 20.2–26 24.29 ± 1.65 0.0262*
HL 17.22–18.44 17.68 ± 0.50 12.1–15.5 14.03 ± 1.24 0.0000*** 14.92–20.80 17.18 ± 2.15 12.2–15.1 13.59 ± 0.89 0.0001***
HW 12.22–13.26 12.70 ± 0.37 11.1–12.9 11.65 ± 1.00 0.0047** 10.18–13.24 11.51 ± 1.19 9.2–12.3 11.01 ± 0.87 0.0082**
HH 6.58–7.10 6.80 ± 0.19 6.4–7 6.68 ± 0.22 0.144 4.34–5.76 5.27 ± 0.45 4.9–6.8 6.06 ± 0.52 0.0197*
SL 6.90–7.68 7.21 ± 0.29 5.4–7 5.98 ± 0.63 0.0012** 5.72–7.64 6.52 ± 0.68 5.2–6.2 5.70 ± 0.41 0.0009***
EED 5.52–6.30 5.83 ± 0.26 5.6–6.1 5.40 ± 0.53 0.143 4.24–5.78 5.03 ± 0.55 3.8–5.7 5.03 ± 0.54 0.722
ED 3.76–4.36 4.08 ± 0.18 3.3–3.8 3.58 ± 0.19 0.0021** 2.90–3.98 3.41 ± 0.38 3–3.9 3.36 ± 0.27 0.386
EOD 0.74–1.24 1.07 ± 0.16 0.6–1.3 0.85 ± 0.29 0.0833 0.64–1.26 0.96 ± 0.18 0.4–1 0.72 ± 0.20 0.0046**
RW 1.56–3.02 2.20 ± 0.53 1.7–2.1 1.95 ± 0.17 0.828 1.62–3.04 2.35 ± 0.49 1.9–2.5 2.18 ± 0.17 0.146
RH 1.12–1.42 1.24 ± 0.11 1.1–1.2 1.15 ± 0.05 0.339 0.82–1.44 1.16 ± 0.23 1–1.6 1.23 ± 0.20 0.616
MW 1.62–2.22 1.89 ± 0.25 1.6–1.9 1.75 ± 0.11 0.631 1.38–1.86 1.59 ± 0.18 1.1–2.1 1.51 ± 0.28 0.319
ML 1.52–1.82 1.63 ± 0.12 1.4–1.8 1.68 ± 0.16 0.995 1.18–1.62 1.38 ± 0.15 1.2–2 1.54 ± 0.24 0.202
FIL 6.70–8.00 7.45 ± 0.47 – – 5.82–7.74 6.66 ± 0.65 – –
HIL 7.18–8.84 8.28 ± 0.66 – – 6.44–9.06 7.82 ± 0.92 – –
SPL 9–13 10.63 ± 1.16 7–10 8.00 ± 0.50 0.0001*** 9–12 10.14 ± 0.74 8–10 8.89 ± 0.50 0.0000***
IFL 8–10 9.13 ± 0.83 6–7 6.63 ± 0.48 0.0000*** 9–10 9.21 ± 0.41 6–9 7.33 ± 0.70 0.0000***
IO 23–27 25.00 ± 1.63 21–24 22.50 ± 1.12 0.0803 22–28 25 ± 2.14 20–24 21.33 ± 1.63 0.0027**
DTR 12–17 15.00 ± 1.87 13–16 14.25 ± 1.30 0.589 14–17 16.00 ± 1.20 12–15 13.67 ± 1.15 0.0024**
SMC 165–189 175.50 ± 8.65 146–161 155.25 ± 5.63 0.0145* 158–181 167.71 ± 8.34 146–169 158.89 ± 6.61 0.0444*
SR 98–114 108.25 ± 6.18 124–149 131.75 ± 10.03 0.0136* 92–106 100.00 ± 4.54 111–142 123.78 ± 10.20 0.0001***
V 33–38 35.50 ± 1.80 35–47 39.08 ± 4.70 0.14 32–39 35.57 ± 2.26 31–47 38.44 ± 4.52 0.172
LF1 8–11 9.75 ± 0.88 8–10 8.85 ± 0.66 0.0901 6–11 9.29 ± 1.53 8–10 8.94 ± 0.66 0.412
LT1 8–11 10.00 ± 0.99 8–10 9.31 ± 0.46 0.334 9–11 9.92 ± 0.47 8–10 9.17 ± 0.53 0.0003
LF4 12–14 12.75 ± 0.70 10–13 11.92 ± 0.83 0.0154* 12–14 12.93 ± 0.61 11–14 11.94 ± 0.83 0.0020**
LT4 12–15 13.88 ± 0.64 12–14 13.23 ± 0.89 0.067 13–15 14.31 ± 0.61 11–14 13.17 ± 0.85 0.0001***
PP 4–7 5.50 ± 1.12 4–5 4.50 ± 0.43 0.121 – – –
PAT 1–2 1.25 ± 0.45 1–2 1.25 ± 0.43 1 1–2 1.29 ± 0.45 1–2 1.11 ± 0.33 0.222
Table 7.
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Variable loadings with the first four principal components of Gekko alpinus sp. nov. and G. jinjiangensis, with morphometric characters corrected.

Mensural characteristics Male Female
PC1 PC2 PC3 PC4 PC1 PC2 PC3 PC4
AGD 0.2599 -0.4656 -0.0058 -0.0808 -0.1946 0.1391 -0.3604 0.5002
HL 0.4186 0.0068 0.0178 0.1257 -0.4724 0.0931 -0.0008 0.1661
HW 0.4042 0.1511 0.1207 -0.0109 -0.3806 -0.0953 -0.0450 -0.1095
HH 0.1810 -0.1023 -0.0092 0.6899 0.1077 -0.5288 0.2804 0.1429
SL 0.3534 -0.2054 0.2237 0.2483 -0.4333 0.0069 0.1578 -0.1221
EED 0.3066 0.2964 0.1377 -0.3658 -0.2573 -0.4521 0.3332 0.0167
ED 0.4202 0.0147 -0.0935 -0.0302 -0.2948 -0.0683 0.1355 0.2130
EOD 0.3339 0.2310 -0.1955 -0.2986 -0.3289 0.3158 0.1295 0.1447
RW -0.0120 -0.3874 -0.4658 -0.3179 -0.2810 -0.3568 -0.2907 -0.0266
RH 0.2125 -0.1708 -0.5392 0.0107 -0.0212 -0.4456 -0.4152 -0.2448
MW -0.0851 0.2069 -0.5609 0.3128 -0.1462 0.0650 -0.5343 -0.3276
ML 0.0396 0.5866 -0.2162 0.1305 0.1808 -0.2107 -0.2756 0.6636
Standard deviation 2.3435 1.5812 1.4494 1.2894 2.0027 1.4749 1.3626 1.0844
Percentage of total variance 45.766 20.836 17.506 13.855 33.424 18.128 15.472 9.799
Cumulative percentage 45.766 66.602 84.108 97.963 33.424 51.552 67.024 76.823

Taxonomic account

Gekko alpinus sp. nov.

https://zoobank.org/CA4785F4-B8D1-4F62-AAF2-B8EB9CC093A3
Figs 4, 5, 6, 7

Type materials

Holotype. • CIB 121663 (Figs 4, 5), an adult male, collected 26 June 2020 (29.615722°N, 99.02285°E; 2542 m a.s.l.), from Zhubalong Village, Batang County, Ganzi Zang Autonomous Prefecture, Sichuan Province, China by Sheng-Chao Shi, Cheng Shen, Xian-Guang Guo, and Jian-Ping Jiang. Paratypes. • One adult male: CIB 121661, four adult females: CIB 121658–60, and CIB 121664, and one subadult female: CIB 121662, with the same collection information as the holotype • One adult male: CIB 121665 and one subadult female: CIB QZ088 collected 29 June 2020 (29.731613°N, 99.002355°E; 2494 m a.s.l.), with the same collection locality and collectors’ information • One adult male: CIB 121656 and one subadult female: CIB 121657 (Fig. 6) collected 7 July 2022 (29.758142°N, 99.005975°E; 2400 m a.s.l.), from Zhubalong Village, Mangkang County, Changdu City, Xizang Autonomous Region, China by Cheng Shen, Li-Ming Chang, and Qun-De Zhang.

Figure 4. 

Holotype (CIB 121663, adult male) of Gekko alpinus sp. nov. A dorsal view of body B ventral view of body. Photographs by S-C Shi.

Figure 5. 

Holotype (CIB 121663, adult male) of Gekko alpinus sp. nov. A right lateral view of head B dorsal view of head C ventral view of body D dorsal view of middle body E ventral view of precloacal region F ventral view of left hand G ventral view of left foot. Photographs by S-C Shi.

Figure 6. 

Paratype (CIB 121657, subadult female) of Gekko alpinus sp. nov. A dorsal view of body B ventral view of body C dorsal view of head D ventral view of head E dorsal view of middle body F ventral view of precloacal region. Photos by S Ma.

Diagnosis

(1) body size moderate, SVL 56.44–74.16 mm in adults; (2) head relatively narrow, HW/HL 0.51–0.79; (3) midbody scale rows 92–114, 98–114 in males and 92–106 in females; (4) interorbital scales between anterior corners of the eyes 22–28; (5) ventral scale rows 32–39; (6) tubercles present on dorsal body, forelimbs, hindlimbs and tails; (7) precloacal pores 4–7 in males and absent in the females; (8) subdigital lamellae on first finger 8–11, on fourth finger 12–14, on first toe 8–11, on fourth toe 12–15, no webbing between the fingers and toes; (9) ventral scales between mental and cloacal slit 158–189; (10) nares in contact with rostral; (11) postcloacal tubercles one or two; (12) dorsal surface of body with six or seven large dark taupe bands between nape and sacrum.

Description of holotype

(Figs 3, 4) An adult male, moderate size, SVL 74.16 mm; body slender and trunk relatively elongate (AGD/SVL 0.37); tail little broken at end, TaL 68.02 mm, slightly shorter than SVL.

Head depressed (HH/HL 0.37), length longer than width (HL/HW), distinct from neck. Snout rounded at top, elongate (SL/HL 0.41/0.42), larger than eye (SL/ED 1.75/1.80); rostral irregular polygon, wider than high (RW/RH 1.74) and slightly narrower than mental (RW/MW 0.95); rostral groove absent; rostral in contact with nostril, first supralabial and nasorostral; nares oval, touching rostral, first supralabial three nasals (nasorostral, supranasal, postnasal); one small internasal; snout region medially concave; preorbitals 12/12, preorbital region deeply concave; eye large (ED/HL 0.24/0.23), pupil vertical with crenulated margins; interorbital scales between anterior corners of eyes 27; ear opening oval, obliquely oriented, much smaller than eye (EOD/ED = 0.27/0.26); mental pentagon, width more than length (MW/ML = 0.73); two enlarged postmentals, hexagonal, twice as long as wide; postmentals in contact with mental and first infralabials anteriorly and five gular scales posteriorly; supralabials 11/11; infralabials 10/10; tubercles absent on dorsal head, granulars on anteriodorsal head larger than those on posterior.

Dorsal scales on body smooth, round or oval, granular, juxtaposed; dorsal tubercles 3–4 times the size of dorsal scales, smooth, round to oval, convex, surrounded by 8–10 dorsal scales; dorsal tubercles extending from occiput region to base of tail; tubercles in 15 regular rows at midbody; ventrolateral fold weakly developed, without tubercles; ventrals distinctly larger than dorsal scales, smooth, imbricate and largest in middle of belly; ventral scale rows at midbody 33; scale rows around mid-body 114; ventral scales in a row between mental and cloacal slit 165; precloacal scales enlarged, but no enlarged scales on thighs; precloacal pores seven, in a continuous row.

Forelimbs and hindlimbs well developed, tubercles on fore and hind limbs are present, moderately long, slender; forearm and tibia moderately long, forearm shorter than tibia; digits moderately expanded, both first finger and first toe, clawless, others remaining digits clawed; webbing on fingers and toes absent; subdigital lamellae unnotched and undivided: 10/10-9/10-11/10-13/13-11/11 (manus) and 11/10-11/10-12/12-15/14-13/13 (pes). Relative length of fingers: IV > III > V > II > I; relative length of toes: IV > III > V > II > I.

Tail oval in section, swollen at base, gradually tapering; postcloacal tubercle 1/1, obviously large on tail base side; dorsal scales small, flat, smooth, with dorsal tubercles at the tail base dorsum; ventral scales much larger than dorsal, smooth, and imbricated, with enlarged subcaudal plates arranged into a longitudinal row formed ~ 1/6 TaL distance from the cloaca.

Coloration of holotype in life

(Figs 4, 5). Dorsal surfaces of head, neck and body dark taupe, irregularly scattered with some pale grey threads or blotches, alternatively ornamented with eight large pale grey and seven dark taupe wide bands from neck to the sacrum; an indistinct pale-colored vertebral line is present from the nape to the tail base; dorsal surfaces of limbs, also dark taupe, mottled with small and pale blotches; dorsal tail dark taupe, alternatively ornamented with nine large pale grey and nine larger dark taupe bands, mottled at the ends; ventral skin creamy white, mosaiced with small taupe pigments.

Coloration of holotype in preservative

The coloration pattern of the specimen mostly faded. Dorsal surfaces of head, neck, and body black taupe, but compared to the living status, much wider body area irregularly creamy white, and still alternatively ornamented with eight large creamy white and seven black taupe wide bands from neck to the sacrum. A creamy white vertebral line extends from the nape to the tail terminal; dorsal surfaces of limbs creamy white, mottled with small taupe blotches; dorsal tail dark taupe, alternatively decorated with nine large creamy white and nine larger taupe bands, and mottled towards the end. Ventral skin creamy white, mosaicked with small taupe pigments, though some areas turned creamy yellow due to prolonged alcohol storage.

Variation

All paratypes are very similar to the holotype. Variation of the mensural characters and meristic features among individuals of the type series are presented in Table 4.

Distribution and habits

Gekko alpinus sp. nov. is currently known only from the Jinsha River Basin between the border of Mangkang County, Xizang Autonomous Region and Batang County, Sichuan Province, China, at elevations ranging from 2400 to 2542 meters above sea level. This new species is nocturnal and inhabits shrubs or dry rocky cliffs in the arid Jinsha River valley, as well as on building walls (Fig. 7). Ants, discovered in the gut of one specimen, are among the recorded food choices of this species.

Figure 7. 

Habitats of Gekko alpinus sp. nov. A macrohabitat: Jinsha River dry-hot valley in Zhubalong Village at the border between Batang County, Sichuan Province and Mangkang County, Xizang Autonomous Region B microhabitat: house walls C one individual found on the dry rocky cliffs D one individual found in the rock crevices on cliff E one individual found on a house wall. Photos by S-C Shi.

Etymology

The specific name alpinus is derived from Latin, alpinus, -a, -um, meaning from Alpēs (“the Alps”) + -īnus, of or pertaining to the Alps, alpine. This refers to the “great high mountains”, referring to not only its distribution range in the great high Hengduan Mountains, but also the highest distribution elevation for all currently known Japonigekko species. The suggested common English name is “Alpine Gecko” and the Chinese name is “高山壁虎” (Gāo Shān Bì Hŭ).

Discussion

The discovery of Gekko alpinus sp. nov. raises the total species number of the genus Gekko to 89, in the subgenus Japonigekko to 34, and within this subgenus in China to 21, including six species distributed in Sichuan Province (Gekko alpinus sp. nov., G. chinensis, G. cib, G. japonicus, G. jinjiangensis, and G. scabridus). Additionally, this is the only Gekko species recorded in Xizang Autonomous Region, marking a new provincial record of this genus in Xizang (Li et al. 2010; Cai et al. 2018; Che et al. 2020; Hou et al. 2021; Lyu et al. 2021).

Hou et al. (2021) reported that the elevation range of G. jinjiangensis as 2000 m to 2476 m a.s.l. However, the type series of G. jinjiangensis was only found from 2045 m to 2114 m a.s.l. The 2476 m of G. jinjiangensis record originally pertains to a Gekko population in Batang County, which is actually Gekko alpinus sp. nov., as described in this study. Consequently, we revise the elevation range of G. jinjiangensis to 2045–2114 m a.s.l., while Gekko alpinus sp. nov. is distributed between 2400 m to 2542 m a.s.l., making it the highest-altitude Japonigekko species currently recognized. Future surveys are recommended to assess the population status of this new species.

The dry-hot valley of Jinsha River in the Hengduan Mountain features habitat heterogeneity and diverse topographic complexity, which supports a variety of reptile species and promotes rapid species evolutionary changes. This is particularly evident in species of Diploderma Hallowell, 1861 (Squamata, Agamidae) (Wang et al. 2020; Cai et al. 2022). The discovery of Gekko alpinus sp. nov. also highlights the previously underestimated reptile diversity of this area. The Gekko alpinus sp. nov. populations found on each side of the Jinsha River do not exhibit significant genetic differentiation (uncorrected p-distance among No 1, 2, 6, 7 of 16S/ND2: 0–0.2%/0–0.2%) (Tables 2, 3), similar to Diploderma batangense (Li, Deng, Wu & Wang, 2001) (uncorrected p-distance of ND2: 0–0.4%), implying that the Jinsha River of Hengduan Mountain in Batang and Mangkang do not pose a significant geographical isolation barrier for local reptiles (Wang et al. 2020). To gain a deeper understanding of the reptile diversity patterns and evolutionary histories of Jinsha River Basin in Hengduan Mountain, future field surveys and comprehensive multidimensional analyses are essential.

Acknowledgements

We send sincere thanks herein to Xian-Guang Guo and Qun-De Zhang (Chengdu Institute of Biology, Chinese Academy of Sciences) for their help in the field survey. We greatly thank Jun-Da Zhao from Seattle University for carefully polishing the manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by the Second Tibetan Plateau Scientific Expedition and Research Program (STEP: 2019QZKK05010503) and China Biodiversity Observation Networks (Sino BON – Amphibian & Reptile).

Author contributions

Conceptualization: SM, SCS, JPJ. Data curation: SCS, SM. Formal analysis: SM. Funding acquisition: JPJ. Investigation: SM, JPJ, SCS, CS, LMC. Methodology: JPJ, SM. Project administration: JPJ. Resources: LMC, CS, JPJ. Software: SM. Supervision: JPJ. Validation: JPJ, SCS. Visualization: SM. Writing - original draft: SM. Writing - review and editing: SM, SCS, JPJ, LMC, CS.

Author ORCIDs

Shun Ma https://orcid.org/0009-0003-8611-4550

Sheng-Chao Shi https://orcid.org/0000-0003-2337-6572

Li-Ming Chang https://orcid.org/0000-0002-2411-6978

Jian-Ping Jiang https://orcid.org/0000-0002-1051-7797

Data availability

All of the data that support the findings of this study are available in the main text.

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