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Research Article
A new species of karst-adapted gecko (Squamata, Gekkonidae, Gekko) from Guangxi, southern China
expand article infoZhong Huang‡, Hao-Tian Wang§, Shuo Qi§, Han-Ming Song§, Yong Huang|, Ying-Yong Wang§, Yun-Ming Mo‡
‡ Natural History Museum of Guangxi, Nanning, China
§ Sun Yat-sen University, Guangzhou, China
| Guangxi University of Chinese Medicine, Nanning, China

Corresponding author: Ying-Yong Wang ( wangyy@mail.sysu.edu.cn )

Corresponding author: Yun-Ming Mo ( moyunming@163.com )

Academic editor: Minh Duc Le
© 2025 Zhong Huang, Hao-Tian Wang, Shuo Qi, Han-Ming Song, Yong Huang, Ying-Yong Wang, Yun-Ming Mo.
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: Huang Z, Wang H-T, Qi S, Song H-M, Huang Y, Wang Y-Y, Mo Y-M (2025) A new species of karst-adapted gecko (Squamata, Gekkonidae, Gekko) from Guangxi, southern China. ZooKeys 1245: 289-310. https://doi.org/10.3897/zookeys.1245.153769
ZooBank: urn:lsid:zoobank.org:pub:4B3EEDA4-CCA2-4EBB-9B22-B2CF3057C6A1
Open Access

Abstract

A new species of the genus Gekko Laurenti, 1768, Gekko fengshanensis sp. nov., is described based on six specimens from Fengshan County, Hechi City, Guangxi Zhuang Autonomous Region, China. It is placed into the subgenus Japonigekko based on morphological and molecular phylogenetic analyses, and distinguished from consubgeners of Japonigekko by a combination of morphological characters in body size, cephalic proportions, and pholidosis features. Molecular phylogenetic analyses using mitochondrial 16S and ND2 sequences reveal that G. fengshanensis sp. nov. forms a sister relationship with G. kwangsiensis, collectively forming a clade with G. liboensis and G. paucituberculatus endemic to southern China’s karst ecosystems. This discovery increases the number of recognized Gekko species in the South China Karst to five, underscoring the role that fragmented karst landscapes play in driving speciation and maintaining high levels of biodiversity in this unique ecosystem.

Key words:

Gekko fengshanensis sp. nov., G. liboensis, Guangxi, integrative taxonomy, Japonigekko, South China Karst

Introduction

The genus Gekko Laurenti, 1768, currently containing 92 known species, is a widely distributed group of nocturnal gekkonid lizards, mainly distributing in throughout plains and plateaus across temperate and tropical Asia and the western islands of Pacific Ocean (Cao et al. 2025; Pauwels et al. 2025; Uetz et al. 2025). According to recent phylogenetic studies based on genomic data, the genus is currently divided into seven subgenera, namely Archipelagekko, Balawangekko, Gekko, Japonigekko, Lomatodactylus, Ptychozoon, and Rhacogekko (Wood et al. 2020). Among these subgenera, Japonigekko exhibits the highest diversity and contains the greatest number of species in China. Its distribution spans across the country, from Liaoning in the northeast to Xizang in the west, and down to Hainan in the south, thriving in a variety of habitats (Qiu et al. 2023; Ma et al. 2024). Notably, Guangxi is home to the largest number of species within this subgenus.

Guangxi’s karst topography, exemplified by iconic formations such as the Leye Giant Sinkhole Cluster and the Guilin Tower Karst, forms an integral component of the South China Karst, a UNESCO World Heritage Site. This lithological mosaic has facilitated the radiation of specialized herpetofauna, with recent decades witnessing the discovery of multiple karst-obligate species including the frog Odorrana lipuensis Mo, Chen, Wu, Zhang & Zhou, 2015, the geckos Gekko kwangsiensis Yang, 2015 and G. paucituberculatus Wang, Qi, Zhou & Wang, 2024, and other saxicolous taxa. In 2024, during two field surveys in the karst forests of northwest Guangxi, six Gekko individuals were collected. Morphologically, these specimens clearly belong to subgenus Japonigekko, characterized by relatively moderate size; nares in contact with rostral; the presence of dorsal tubercle rows and precloacal pores; and lacking tubercles on ventrolateral folds. Phylogenetic analysis places them in a distinct evolutionary lineage, separate from closely related congeners within the subgenus. This suggests they represent a previously undescribed species, distinguished by both molecular and morphological differences. In light of these findings, we provide a detailed description of the new species herein.

Materials and methods

Specimens and morphology

Six specimens of Gekko fengshanensis sp. nov. were collected from Fengshan County, Hechi City, Guangxi Zhuang Autonomous Region, China in 2024 (Fig. 1). The specimens were euthanized and then fixed in 10% buffered formalin, later transferred to 75% ethanol and deposited in the Natural History Museum of Guangxi (NHMG), Nanning and Sun Yat-sen University (SYS), Guangzhou, China. Liver tissue samples were preserved in 95% ethanol for molecular analysis of four specimens (NHMG 240713/SYS r002885, NHMG 240714/SYS r002886, NHMG 202408005, 006). Two specimens of G. liboensis (SYS r002876, 77) were also collected and sequenced.

Figure 1. 

Localities of Gekko fengshanensis sp. nov., G. kwangsiensis, G. liboensis, and G. paucituberculatus. 1 Fengcheng Town, Fengshan County, Hechi City, Guangxi; 2 Wuming District, Nanning City, Guangxi; 3 Maolan National Nature Reserve, Libo County, Guizhou Province; 4 Tianyang District, Baise City, Guangxi. “?” indicates the unconfirmed locality of G. fengshanensis sp. nov.

Measurements were taken with digital calipers (Deli DL91200 Digital Vernier Caliper) to the nearest 0.1 mm on the right side of the body, and scalation features were counted under a binocular scope (Leica EZ4 HD). Bilateral scale counts are given as left/right. External measurements, meristic traits and their abbreviations followed Wang et al. (2024). They are snout-vent length (SVL, from tip of snout to anterior margin of cloaca) ; tail length (TaL, from posterior margin of cloaca to tip of tail) ; axillia-groin length (>AG, distance between axilla and groin) ; head length (HL, from tip of snout to posterior margin of ear opening) ; head width (HW, at the angle of the jaws) ; head height (HH, from the top of the head posterior to the eyes to the bottom of the lower jaw) ; snout length (SNT, from snout tip to anterior corner of eye) ; maximum eye diameter (ED) ; maximum ear opening diameter (EOD) ; maximum rostral width (RW) ; maximum rostral height (RH) ; maximum mental length (ML). Meristic characters are nasals (N, nasorostrals, supranasals and postnasals) ; intersupranasals (I, scales between supranasals, in contact with rostral ; supralabials and infralabials (SPL and IFL, number of scales from commissure of jaw to the rostral /mental scale) ; interorbitals (IO, number of scales in a line between anterior margins of eyes) ; preorbitals (PO, number of scales in a line from nostril to anterior margin of the eye) ; postmentals (PM, scales bordering the mental) ; gulars bordering the postmentals (GP) ; dorsal tubercle rows at midbody (DTR) ; granules surrounding dorsal tubercles (GSDT) ; scales in a line from mental to the front of cloacal slit (SMC) ; ventral scale rows at midbody between ventrolateral folds (V) ; scale rows at midbody (SR, including ventral scales) ; subdigital lamellae of entire first finger (LF1) ; subdigital lamellae of entire fourth finger (LF4) ; subdigital lamellae of entire first toe (LT1 ; subdigital lamellae of entire fourth toe (LT4) ; precloacal pores (PP) ; postcloacal tubercles (PAT) ; transverse dorsal scale rows in the middle of the third caudal whorl (S3W). The information on character states of other Japonigekko species was obtained from the literature (Table 1).

Table 1.
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Literature and authorities for species of Japonigekko morphological characters used in this study.

ID Species References
1 Gekko aaronbaueri Ngo, Thai, Phimvohan, David & Teynié, 2015 Ngo et al. (2015)
2 G. adleri Nguyen, Wang, Yang, Lehmann, Le, Ziegler & Bonkowski, 2013 Nguyen et al. (2013)
3 G. alpinus Ma, Shi, Shen, Chang & Jiang, 2024 Ma et al. (2024)
4 G. auriverrucosus Zhou & Liu, 1982 Zhou et al. (1982)
5 G. bonkowskii Luu, Calame, Nguyen, Le & Ziegler, 2015 Luu et al. (2015)
6 G. canhi Rösler, Nguyen, Van Doan, Ho, Nguyen & Ziegler, 2010 Rösler et al. (2010)
7 G. chinensis (Gray, 1842) Ota et al. (1995)
8 G. cib Lyu, Lin, Ren, Jiang, Zhang, Qi & Wang, 2021 Lyu et al. (2021)
9 G. guishanicus Lin & Yao, 2016 Lin and Yao (2016)
10 G. hokouensis Pope, 1928 Zhou et al. (1982)
11 G. ichangensis Cao, Sucharitakul, Tie, Suwannapoom, Yan & Chomdej, 2025 Cao et al. (2025)
12 G. japonicus (Schlegel, 1836) Zhou et al. (1982)
13 G. jinjiangensis Hou, Shi, Wang, Shu, Zheng, Qi, Liu, Jiang & Xie, 2021 Hou et al. (2021)
14 G. kaiyai Zhang, Wu & Zhang, 2023 Zhang et al. (2023)
15 G. khunkhamensis Sitthivong, Lo, Nguyen, Ngo, Khotpathoom, Le, Ziegler & Luu, 2021 Sitthivong et al. (2021)
16 G. kwangsiensis Yang, 2015 Yang (2015)
17 G. liboensis Zhou, Liu & Li, 1982 This study
18 G. melli (Vogt, 1922) Lyu et al. (2021)
19 G. nadenensis Luu, Nguyen, Le, Bonkowski & Ziegler, 2017 Luu et al. (2017)
20 G. palmatus Boulenger, 1907 Ota et al. (1995); Song et al. (2024)
21 G. paucituberculatus Wang, Qi, Zhou & Wang, 2024 Wang et al. (2024)
22 G. scabridus Liu & Zhou, 1982 Zhou et al. (1982)
23 G. scientiadventura Rösler, Ziegler, Vu, Herrmann & Böhme, 2004 Rösler et al. (2004)
24 G. sengchanthavongi Luu, Calame, Nguyen, Le & Ziegler, 2015 Luu et al. (2015)
25 G. shibatai Toda, Sengoku, Hikida & Ota, 2008 Toda et al. (2008)
26 G. similignum Smith, 1923 Ota et al. (1995)
27 G. subpalmatus (Günther, 1864) Lyu et al. (2021)
28 G. swinhonis Günther, 1864 Rösler et al. (2011)
29 G. taibaiensis Song, 1985 Song (1985)
30 G. tawaensis Okada, 1956 Rösler et al. (2011)
31 G. thakhekensis Luu, Calame, Nguyen, Le, Bonkowski & Ziegler, 2014 Luu et al. (2014)
32 G. truongi Phung & Ziegler, 2011 Phung and Ziegler (2011)
33 G. vertebralis Toda, Sengoku, Hikida & Ota, 2008 Toda et al. (2008)
34 G. vietnamensis Sang, 2010 Sang (2010)
35 G. wenxianensis Zhou & Wang, 2008 Zhou and Wang (2008)
36 G. yakuensis Matsui & Okada, 1968 Rösler et al. (2011)

The morphometric measurements were statistically analyzed using R v. 4.4.2 (R Core Team, 2024). 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 each species. One-way analysis of variance (ANOVA) was conducted with statistically similar variances (p > 0.05 in the Levene’s test) and performed on a dataset coded for species to examine statistically significant mean differences (p < 0.05) among characters using the car R package. Character means showing significant differences were subjected to a Tukey HSD test to determine which pairs of species differed significantly for those specific characters. Principal component analysis (PCA) was performed to cluster the morphometrics except SVL related to each species using GroupStruct R package (Chan and Grismer 2022). Multiple Factor Analysis (MFA) is a multivariate approach for integrating multiple sets of quantitative traits into a single analysis (Pagès 2015). Each dataset is first analyzed separately using PCA and then standardized by dividing all values by the square root of the first eigenvalue. The standardized datasets are then combined for a global PCA. This ensures that all trait sets contribute equally to the overall variation (Grismer et al. 2024).

To evaluate morphological differences among species or populations, we performed a non-parametric permutation multivariate analysis of variance (PERMANOVA) from the vegan package in R (Oksanen et al. 2020) on the PCA scores and the first five MFA dimensions. This test, based on Euclidean distances and 50,000 permutations, assesses whether the group centroids and dispersions differ significantly in multivariate space. A significant adjusted p-value (< 0.05) indicates clear separation among groups. Given the number of individuals available for closely related species, we performed ANOVA comparisons only with Gekko kwangsiensis, while in the PCA and MFA, we include all three (G. kwangsiensis, G. liboensis, and G. paucituberculatus) for visualization. The raw data are given in Suppl. material 1.

Phylogenetic sampling and analyses

Twelve new sequences and 51 sequences from GenBank were used for molecular analysis in this study. All newly collected tissue samples were obtained from euthanized specimens and then preserved in 95% ethanol and stored at -40 °C. Gekko gecko (Linnaeus, 1758) and G. reevesii (Gray, 1831) belonging to subgenus Japonigekko were used to root the tree based on Rösler et al. (2011) and Lyu et al. (2021). Detail information of these samples is given in Table 2.

Table 2.
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Localities, voucher information, and GenBank accession numbers for all samples used in this study.

ID Species Locality Voucher ID 16S ND2 Reference
1 Gekko fengshanensis sp. nov. China: Guangxi: Hechi: Fengshan NHMG 240713/SYS r002885 PV652773 PV657377 This study
2 Gekko fengshanensis sp. nov. China: Guangxi: Hechi: Fengshan NHMG 240714/SYS r002886 PV652774 PV657378 This study
3 Gekko fengshanensis sp. nov. China: Guangxi: Hechi: Fengshan NHMG 202408005 PV652775 PV657379 This study
4 Gekko fengshanensis sp. nov. China: Guangxi: Hechi: Fengshan NHMG 202408006 PV652776 PV657380 This study
5 G. adleri China: Guangxi: Jingxi SYS r001400 MW451654 OR902178 Lyu et al. (2021); Wang et al. (2024)
6 G. alpinus China: Xizang: Mangkang CIB 121656 PQ255976 PQ303494 Ma et al. (2024)
7 G. auriverrucosus China: Shanxi: Yuncheng NNU Z 20050716.004 — JN019062 Rösler et al. (2011)
8 G. bonkowskii Laos: Khammouane VFU R.2014.10 — KT266818 Luu et al. (2015)
9 G. chinensis China: Hong Kong SYS r001211 MW451644 OR902183 Lyu et al. (2021); Wang et al. (2024)
10 G. cib China: Sichuan: Hejiang SYS r001489 MW451655 OR902165 Lyu et al. (2021); Wang et al. (2024)
11 G. hokouensis China: Jiangxi: Mt. Meiling SYS r001311 MW451648 OR902172 Lyu et al. (2021); Wang et al. (2024)
12 G. hokouensis China: Fujian: Mt. Wuyi SYS r001290 MW451647 OR902173 Lyu et al. (2021); Wang et al. (2024)
13 G. japonicus China: Fujian: Mt. Wuyi SYS r000672 MW451628 OR902176 Lyu et al. (2021); Wang et al. (2024)
14 G. japonicus China: Jiangxi: Lushan SYS r001317 MW451649 OR902177 Lyu et al. (2021); Wang et al. (2024)
15 G. jinjiangensis China: Yunnan: Deqin CIB 5334220088 — MT449431 Hou et al. (2021)
16 G. kaiyai China: Henan: Xinyang: Xinxian AHUXXBH01 OQ780318 — Zhang et al. (2023)
17 G. kaiyai China: Henan: Xinyang: Xinxian AHUXXBH02 OQ780319 — Zhang et al. (2023)
18 G. khunkhamensis Laos: Khammouane VNUF R.2021.23 — OL416111 Sitthivong et al. (2021)
19 G. kwangsiensis China: Guangxi: Wuming SYS r001194 MW451641 OR902174 Lyu et al. (2021); Wang et al. (2024)
20 G. kwangsiensis China: Guangxi: Wuming SYS r001195 MW451642 OR902175 Lyu et al. (2021); Wang et al. (2024)
21 G. liboensis China: Guizhou: Libo: Maolan SYS r002876 PV652777 PV657381 This study
22 G. liboensis China: Guizhou: Libo: Maolan SYS r002877 PV652778 PV657382 This study
23 G. melli China: Guangdong: Dongyuan SYS r001742 MW451661 OR902169 Lyu et al. (2021); Wang et al. (2024)
24 G. nadenensis Laos: Khammouane ZFMK 98741 — KY421618 Luu et al. (2017)
25 G. palmatus China: Guangdong: Mt.Dinghu SYS r002797 OR903156 OR902179 Wang et al. (2024)
26 G. paucituberculatus China: Guangxi: Baise: Tianyang SYS r002806 OR903154 OR902163 Wang et al. (2024)
27 G. paucituberculatus China: Guangxi: Baise: Tianyang SYS r002807 OR903155 OR902164 Wang et al. (2024)
28 G. scabridus China: Sichuan: Yanbian CIB YN201909199 PQ255992 MT449429 Hou et al. (2021); Ma et al. (2024)
29 G. scientiadventura Vietnam: Quang Binh IEBR A.2014.7 — KP205392 Luu et al. (2014)
30 G. sengchanthavongi Laos: Khammouane VFU R2014.14 — KT266816 Luu et al. (2015)
31 G. similignum China: Hainan: Mt. Wuzhi SYS r001597 MW451658 OR902185 Lyu et al. (2021); Wang et al. (2024)
32 G. subpalmatus China: Zhejiang: Fenghua SYS r001762 MW451662 OR902167 Lyu et al. (2021); Wang et al. (2024)
33 G. swinhonis China: Hebei: Zunhua SYS r001814 MW451666 OR902171 Lyu et al. (2021); Wang et al. (2024)
34 G. thakhekensis Laos: Khammouane: Thakhek IEBR A.2014.6 — KP205396 Luu et al. (2014)
35 G. truongi Vietnam: Khanh Hoa IEBR A.2011.1 — KP205398 Luu et al. (2014)
36 G. gecko China: Guangxi: Nanning N/A AY282753 AY282753 Zhou et al. (2006)
37 G. reevesii China: Guangdong: Mt. Yinping SYS r000796 MW451630 OR902187 Lyu et al. (2021); Wang et al. (2024)

Genomic DNA was extracted from liver tissue using a DNA extraction kit (Tiangen Biotech Co., Ltd, Beijing). Two fragments of the mitochondrial genes that encode partial 16S ribosomal RNA gene (16S) and partial NADH dehydrogenase subunit 2 gene (ND2) were amplified. Primers used for two genes are obtained from Simon et al. (1994) and Jonniaux and Kumazawa (2008). PCR amplifications were processed with the cycling conditions that initial denaturing step at 95 °C for 4 min and 5 min (16S and ND2, respectively), 35 cycles of denaturing at 95 °C for 40 s, annealing at 53 °C for 34 s (16S) and 55 °C for 40 s (ND2), extending at 72 °C for 60 s, and a final extending step at 72 °C for 10 min. PCR products were purified with spin columns and then sequenced with a forward primer using BigDye Terminator Cycle Sequencing Kit (Applied Biosystems, Waltham, MA, USA). Sequencing was performed on an ABI Prism 3730 automated DNA sequencer by Wuhan Tianyi Huiyuan Bioscience and Technology Inc.

DNA sequences were aligned by the MUSCLE algorithm with default parameters (Edgar 2004). PartitionFinder2 was used to determine the best partitioning scheme (Lanfear et al. 2017)and jModelTest v. 2.1.2 was used to determine the best fitting nucleotide substitution models (Darriba et al. 2012), resulting in the partitions by gene and ND2 was further partitioned by codon position and the best fit models for all partitions as GTR + I + G. Sequenced data were analyzed using Bayesian Inference (BI) in MrBayes 3.2.4 (Ronquist et al. 2012) and Maximum Likelihood (ML) in RaxmlGUI 1.3 (Silvestro and Michalak 2012). Two independent runs were conducted in the BI analysis with 2,000,000 generations each and sampled every 1000 generations with the first 25% of samples discarded as burn-in, resulting in a potential scale reduction factor (PSRF) of < 0.005. In the ML analysis, a bootstrap consensus tree inferred from 1000 replicates was generated. MEGA11 was used to calculate uncorrected pairwise sequence divergence for 16S and ND2 among and within species using the complete deletion option which removes missing data and gaps (Tamura et al. 2021).

Results

Morphological analyses

The results of one-way ANOVA of morphometrics (Table 3) show that the Japonigekko populations from Fengshan County is significantly different from Gekko kwangsiensis, especially in the characteristics of HL, ED, SNT, MW, and ML. In PCA analysis, the extracted components PC1, PC2, PC3, and PC4 eigenvectors account for 64.25%, 12.92%, 8.62%, and 6.10% of the variance, respectively, or 91.89% cumulatively. As illustrated in the scatter plots of PC1 and PC2 (Fig. 2), samples from Fengshan County cluster together and do not overlap with other species. However, the PERMANOVA analyses indicated that the centroid locations of the Fengshan population in the PCA are statistically not significantly different from G. kwangsiensis (Table 4). In the MFA, the Fengshan population and G. kwangsiensis showed slight overlap in morphospace (Fig. 3A) but were significantly different based on PERMANOVA (adjusted p-vaule < 0.05, Table 4). Dimension 1, which accounted for 28.6% of the total variation, was primarily influenced by morphometric traits and contributed most to the separation of species along the primary axis (Fig. 3B). In contrast, Dimensions 2–4 were dominated by meristic data, highlighting their role in further resolving interspecific differences beyond the major morphometric patterns.

Table 3.
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Morphometric comparisons based on the morphometric measurements of Gekko fengshanensis sp. nov. (n = 6) and G. kwangsiensis (n = 6). * p-values < 0.05, ** p-values < 0.01, *** p-values < 0.001.

G. fengshanensis sp. nov. G. kwangsiensis F values p-values
SVL 54.1–79.9 53.8–69.7 0.66876 0.432537
65.95 ± 9.45 61.92 ± 6.49
AG 24.7–34.2 23.4–32.6 0.366798 0.558249
29.78 ± 3.86 27.62 ± 3.28
HL 14.1–20.3 14.2–19.1 18.57999 0.001536**
16.90 ± 2.37 16.77 ± 1.79
HW 11.6–16.0 11.2–14.4 1.963985 0.191352
14.00 ± 1.77 12.90 ± 1.28
HH 5.5–9.1 6.0–7.7 2.043348 0.183361
7.12 ± 1.35 7.03 ± 0.81
ED 3.8–5.7 3.5–4.7 11.42863 0.006995**
4.73 ± 0.65 4.22 ± 0.49
SNT 6.0–8.8 6.3–8.4 8.377919 0.015984*
7.55 ± 1.06 7.47 ± 0.85
RW 2.5–3.2 2.3–3.0 3.082883 0.109644
2.73 ± 0.27 2.76 ± 0.30
RH 1.1–1.6 1.1–1.6 2.014796 0.186184
1.37 ± 0.18 1.35 ± 0.16
MW 1.6–2.3 1.9–2.4 8.324792 0.016238*
2.02 ± 0.29 2.20 ± 0.20
ML 1.0–1.4 1.3–1.7 27.29398 0.000387***
1.15 ± 0.14 1.43 ± 0.18
Table 4.
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PERMANOVA summary statistics for the centroid placement between all species pairs from the PCA and MFA analyses. Bold fonts denote insignificant adjusted p-values.

Species pairs F. Model R2 p-value adjusted p-value
PCA statistics
G. fengshanensis sp. nov. vs kwangsiensis 1.9464875 0.162933875 0.133037339 0.798224036
G. fengshanensis sp. nov. vs liboensis 0.191965061 0.031002284 0.892857143 1
G. fengshanensis sp. nov. vs paucituberculatus 10.68575178 0.640411767 0.035714286 0.214285714
G. kwangsiensis vs liboensis 1.418195136 0.191177923 0.392857143 1
G. kwangsiensis vs paucituberculatus 17.91796582 0.749142546 0.035714286 0.214285714
G. liboensis vs paucituberculatus 3.879400985 0.659829291 0.333333333 1
MFA statistics
G. fengshanensis sp. nov. vs kwangsiensis 5.722068474 0.363951377 0.001939961 0.011639767
G. fengshanensis sp. nov. vs liboensis 14.22442629 0.703329038 0.035714286 0.214285714
G. fengshanensis sp. nov. vs paucituberculatus 9.340062225 0.608867297 0.035714286 0.214285714
G. kwangsiensis vs paucituberculatus 8.838729531 0.595652715 0.035714286 0.214285714
G. kwangsiensis vs liboensis 13.81080264 0.697134936 0.035714286 0.214285714
G. liboensis vs paucituberculatus 17.9312 0.899655 0.333333 1
Figure 2. 

Scatter plot of PC1 and PC2 of Principal Component Analysis based on the morphometric measurements.

Figure 3. 

A. MFA of Gekko fengshanensis sp. nov., G. kwangsiensis, G. liboensis, and G. paucituberculatus; B. Percent contributions of each data type to the inertia of dimensions 1–4 of the MFA. Percentage values on the bar graphs are the amounts of inertia for their respective dimensions.

Phylogenetic analyses

The aligned dataset contained a total of 1578 nucleotide base pairs (bp), with 559 bp for 16S and 1015 bp for ND2. The BI and ML analyses resulted in essentially identical topologies (BI topology with ML bootstrap values in Fig. 4). The mean uncorrected p distances based on 16S and ND2 used in this study are given in Tables 5, 6. In both analyses, the newly collected samples from Fengshan County consistently formed a strongly supported (BPP = 1.00; BS = 100) monophyletic lineage within the subgenus Japonigekko. Moreover, the phylogenetic tree revealed that this lineage is sister to Gekko kwangsiensis with moderate support (BPP = 0.90; BS = 80), and together they are sister to G. liboensis. Finally, the three aforementioned taxa, together with G. paucituberculatus, formed a strongly supported endemic clade that exclusively inhabits the South China Karst. Based on uncorrected p distances, the new samples exhibited 8.20% (G. liboensis) to 10.32% (G. paucituberculatus) divergence in the 16S and 14.11% (G. paucituberculatus) to 16.36% (G. kwangsiensis) divergence in the ND2 gene compared to the other three species.

Table 5.
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Uncorrected p distances (%) of the 16S gene amongst species of Japonigekko used in this study.

Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
1 G. fengshanensis sp. nov. 0.00
2 G. adleri 15.08 —
3 G. alpinus 12.96 14.29 —
4 G. chinensis 15.08 3.70 12.43 —
5 G. cib 12.70 14.81 9.52 12.96 —
6 G. hokouensis 13.10 15.21 13.10 13.62 12.57 0.26
7 G. japonicus 13.49 15.08 10.85 14.02 12.70 12.83 0.00
8 G. kaiyai 12.96 14.02 13.76 12.70 11.90 6.75 13.23 0.00
9 G. kwangsiensis 8.73 14.81 12.83 13.36 12.57 12.17 13.89 11.38 0.53
10 G. liboensis 8.20 13.76 11.64 12.43 11.64 12.83 11.90 11.90 9.52 0.00
11 G. melli 12.17 12.96 9.26 12.17 5.29 12.30 12.70 11.64 12.83 10.85 —
12 G. palmatus 16.14 2.65 13.23 3.44 14.02 14.15 14.81 13.23 14.02 13.76 12.70 —
13 G. paucituberculatus 10.32 15.34 11.11 13.76 12.17 11.77 14.55 12.17 10.32 7.41 12.96 14.29 0.00
14 G. scabridus 12.17 13.49 5.56 12.70 10.05 10.98 10.85 11.90 13.36 11.64 10.85 12.70 10.32 —
15 G. similignum 16.40 3.97 13.23 1.32 14.29 13.89 13.76 12.96 14.68 12.96 12.96 3.70 15.08 12.96 —
16 G. subpalmatus 14.55 15.08 10.32 14.02 6.61 14.15 14.02 12.43 12.83 13.76 6.08 14.29 15.34 12.43 14.81 —
17 G. swinhonis 15.61 16.67 14.02 15.61 11.64 14.15 14.29 13.76 15.74 13.49 12.17 16.14 15.34 14.55 15.87 13.23
Table 6.
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Uncorrected p distances (%) of the ND2 gene amongst species of Japonigekko used in this study.0

Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
1 G. fengshanensis sp. nov. 0.00
2 G. adleri 21.27 —
3 G. alpinus 19.43 21.27 —
4 G. auriverrucosus 20.04 24.13 20.86 —
5 G. bonkowskii 19.63 24.54 19.22 21.88 —
6 G. chinensis 22.09 14.52 21.47 25.15 20.86 —
7 G. cib 20.86 23.72 22.29 19.22 19.22 23.31 —
8 G. hokouensis 19.73 21.88 18.51 20.14 19.63 22.09 22.09 1.84
9 G. japonicus 21.06 23.52 19.22 19.02 20.45 22.29 23.11 21.06 0.00
10 G. jinjiangensis 18.40 21.27 8.38 21.06 19.22 22.09 21.27 20.35 18.40 —
11 G. khunkhamensis 22.29 25.97 21.47 24.34 15.34 24.34 21.06 24.13 22.70 21.27 —
12 G. kwangsiensis 16.36 21.06 21.37 21.68 21.37 21.47 18.61 20.86 22.29 19.33 21.57 0.41
13 G. liboensis 15.44 21.98 20.45 20.35 19.12 22.80 20.76 16.97 21.57 19.84 21.57 17.28 0.20
14 G. melli 23.11 20.86 20.86 22.29 22.09 22.29 17.59 21.27 21.68 20.86 23.52 21.68 21.17 —
15 G. nadenensis 20.65 23.52 19.22 21.06 6.95 21.27 20.86 22.29 21.27 18.61 13.91 20.55 18.51 21.47 —
16 G. palmatus 22.29 6.54 22.09 23.93 23.31 15.54 24.34 21.47 22.90 21.27 26.18 21.27 21.78 23.72 23.31 —
17 G. paucituberculatus 14.11 22.09 17.59 18.61 18.81 22.70 19.02 18.92 19.84 18.20 21.68 15.54 15.85 21.27 18.61 21.27 0.00
18 G. scabridus 16.16 18.61 11.86 19.43 19.84 20.45 21.27 17.28 17.79 10.84 21.88 19.84 17.48 20.45 19.43 19.63 18.00 —
19 G. scientiadventura 19.63 24.34 18.81 21.06 13.70 22.70 21.47 21.68 22.09 18.61 14.72 20.86 20.14 22.09 13.91 23.52 19.02 17.79 —
20 G. sengchanthavongi 20.04 23.93 20.45 20.65 13.91 21.68 21.68 22.90 21.68 19.84 15.75 20.86 19.73 22.49 12.07 23.72 19.22 18.81 10.43 —
21 G. similignum 22.49 14.72 22.49 26.58 22.49 4.09 24.74 22.70 22.90 22.09 24.74 21.68 22.80 23.11 22.49 15.34 22.49 20.45 22.90 22.70 —
22 G. subpalmatus 22.49 23.72 21.27 20.04 20.86 22.90 17.38 20.76 21.68 21.27 22.70 22.09 19.73 15.13 19.43 25.15 20.45 20.45 21.68 21.27 23.93 —
23 G. swinhonis 20.86 22.70 21.47 18.81 21.68 22.90 21.27 19.94 20.86 21.06 23.52 19.84 20.14 20.65 21.68 22.90 21.47 20.04 23.11 21.88 22.29 20.25 —
24 G. thakhekensis 19.63 21.68 19.43 21.06 7.16 19.63 20.45 20.96 20.04 19.63 15.95 20.35 18.51 21.06 6.95 22.70 17.79 19.02 13.09 12.88 20.45 20.45 23.72 —
25 G. truongi 23.31 19.84 20.86 24.54 22.09 19.84 21.68 21.98 22.29 21.06 22.29 22.19 21.37 20.86 22.29 21.47 20.86 17.79 22.09 21.88 20.45 21.47 23.72 20.45
Figure 4. 

Bayesian inference tree inferred from16S and ND2 genes. Numbers before slash indicate Bayesian posterior probabilities (BPP) and numbers after slash are bootstrap support (BS).

In summary, when integrated with the morphological data, the phylogenetic results provide compelling evidence that the observed genetic differences of the new population represent a separate evolutionary lineage. Consequently, the data robustly support the recognition of this taxon as a new species of the subgenus Japonigekko.

Taxonomic account

Gekko fengshanensis sp. nov.

https://zoobank.org/755C0A3B-57DC-4DE8-9B00-141BA36D54C4
Figs 5, 6

Type material.

Holotype. • NHMG 202408004 (Figs 5A, 6), adult male, from Fenghuang Village, Fengcheng Town, Fengshan County (24°31'59.25"N, 107°6'45.02"E; 680 m a.s.l.), Hechi City, Guangxi Zhuang Autonomous Region, China, collected on 5 August 2024 by Zhong Huang, Xiao-Wen Liao, Ben-Ze Huang and Yun-Ming Mo. Paratypes. • NHMG 202408005–007 (Fig. 5B, C), adult males, data identical to the holotype: NHMG 240713/SYS r002885, adult female, and NHMG 240714/SYS r002886, subadult male, from Xinglong Village, Fengcheng Town, Fengshan County (24°31'0.87"N, 107°4'23.97"E; 650 m a.s.l.), collected on 11 July 2024 by the same collector.

Figure 5. 

Type specimens of Gekko fengshanensis sp. nov. in life. A. Holotype NHMG 202408004, adult male; B. Paratype NHMG 202408005, adult male; C. Paratype NHMG 202408007, adult male. Photos by ZH.

Figure 6. 

Morphological features of the adult male holotype NHMG 202408004 of Gekko fengshanensis sp. nov.; A. Dorsal view of body; B. Ventral view of body; C. Dorsal head; D. Ventral head; E. Precloacal pores; F. Left hand; G. Left foot; H. Ventral tail. Photos by HMS.

Diagnosis.

Gekko fengshanensis sp. nov. is assigned to the subgenus Japonigekko and distinguished from congeners by the following combination of characters: (1) moderate body size, SVL 60.0–79.9 mm in the adult male and SVL 62.2 in the adult female; (2) nares in contact with rostral, internasal absent; (3) enlarged postmentals two; (4) tubercles flattened, present from the region behind the eyes along the neck to the tail base, 8–11 rows at midbody ; (5) ventral scales between mental and cloacal slit 193–213; (6) midbody scale rows 149–161; (7) ventral scale rows 40–49; (8) subdigital lamellae on first fingers 11–13, on fourth fingers 12–16, on first toes 12–14, on fourth toes 13–15,and fingers and toes webbing weakly developed; (9) continuous precloacal pores 9–12 in males, absent in the female; (10) a single postcloacal tubercle on each side.

Description of holotype.

Adult male, moderate size, SVL 73.8 mm; head depressed (HH/HL 0.47), length longer than width (HL/HW 1.20), distinct from neck; snout rounded anteriorly, elongate (SNT/HL 0.42), larger than eye (SNT/ED 1.65); rostral regular rectangular, nearly twice as wide as high (RW/RH 1.93), and wider than the width mental (RW/MW 1.32); nares oval, bordered by rostral, first supralabial, supranasal, and two enlarged nasals posteriorly; internasals absent; preorbitals 18/19, preorbital region deeply concave; eye large (ED/HL 0.27), vertical pupil with crenulated margins; interorbital scales between anterior margins of eyes 27; ear opening elliptical, obliquely oriented, moderate in size (EOD/ED 0.33); mental pentagonal, wider than long (MW/ML 1.83); postmentals two, hexagonal and enlarged, twice as long as wide, touching mental and first infralabial on both sides and five gular scales posteriorly; supralabials 11/12; infralabials 11/10; tubercles present on the region behind the eyes, granular scales on anterodorsal region of head larger than those on posterior region.

Body slender, elongate (AG/SVL 0.46); dorsals smooth, round to oval, granular and juxtaposed; tubercles flattened, from postorbital region along the neck to tail base, nine rows at midbody, surrounded by 10 dorsal scales; ventrolateral fold present, without tubercles; ventrals distinctly larger than dorsals, smooth, imbricate, and largest in middle of belly; ventral scale rows at midbody 43; scale rows around midbody 149; ventral scales in a row between mental and cloacal slit 206; precloacal scales enlarged, but no enlarged scales on thighs; precloacal pores 12, in a continuous row across midline; postcloacal tubercle 1/1, large.

Fore- and hindlimbs well-developed; tubercles absent on dorsal surface of limbs; digits moderately dilated; II–IV fingers and toes clawed; claws depressed laterally, extending beyond terminal lamellae; webbing on fingers and toes weakly developed; subdigital lamellae undivided, manus for 12-11-13-15-12 (left) and 10-11-12-12-12 (right), pes for 13-11-14-13-11 (left) and 13-13-13-14-11 (right); relative length fingers and toes I < II < V < III < IV.

Original tail longer than body (TaL 87.3 mm, TaL/SVL 1.18); distinctly swollen at base; dorsal scales small, flat, smooth; caudal whorls distinct, 10 dorsal scale rows in the middle of the third one; subcaudals transversely enlarged.

Coloration of holotype.

In life, the dorsal regions of the head and body are light reddish-brown, with scattered white spots on the snout and posterior orbit. The iris is yellow-green with vermiform markings, and the pupil is dark brownish black. Seven irregularly shaped light patches are arranged along the ridge between the nape and the sacrum, with one or two rows of smaller spots parallelly arranged on each side. An intermittent light-colored vertebral line with black edges extends from the nape to the base of the tail. The anterior part of the tail exhibits a sharp contrast in color, which gradually fades and blends towards the posterior end. The ventral surface is lightly flesh-colored. In preservative, the dorsal ground color of the head, body, and limbs turns greyish black, while the ventral surface fades to greyish-white.

Morphological variation.

Measurements and scale counts of six individuals are shown in Table 7. Precloacal pores are absent in the female. In males, the postcloacal tubercle is significantly larger than in the female. Except for those with a broken or regenerated tail, all paratypes have tails with a clear distinction between light and dark areas. NHMG 202408005 has a dark-brown “()”-shaped marking on the occipital region, and some of the dorsal patches combine with the spots on both sides to form a slightly larger, irregular patch (Fig. 5B). The light spots on the body of NHMG 202408007 are almost uniform in size, hollow, and resemble leopard spots (Fig. 5C). In terms of meristic traits, the precloacal pores of NHMG 202408007 are discontinuous, with four on the left side and five on the right side (Fig. 7A). The postmentals of NHMG 202408005 are slightly enlarged, and even smaller than the scales immediately following them (Fig. 7B).

Table 7.
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Measurements (in mm), body proportions, and scalation features of the type series of Gekko fengshanensis sp. nov. See Materials and Methods section for abbreviations. “*” regenerated tail; “—” unavailable data. Bilateral scale counts are given as left/right.

Holotype Paratypes
Voucher Number NHMG 202408004 NHMG 202408005 NHMG 202408006 NHMG 202408007 NHMG 240713/SYS r002885 NHMG 240714/SYS r002886
Sex Male Male Male Male Female Subadult male
SVL 73.8 60.0 65.7 79.9 62.2 54.1
TaL 87.3 67.1 — 74.3* 75.7 —
AG 34.1 26.9 30.4 34.2 28.4 24.7
HL 19.2 15.7 16.5 20.3 15.6 14.1
HW 16.0 12.9 14.3 16 13.2 11.6
HH 9.1 6.5 6.6 8.4 6.6 5.5
SNT 8.6 7.1 7.8 8.8 7.0 6.0
ED 5.2 4.5 4.6 5.7 4.6 3.8
EOD 1.7 1.3 1.6 1.9 1.7 1.2
RH 1.5 1.3 1.4 1.6 1.3 1.1
RW 2.9 2.7 2.6 3.2 2.5 2.5
MW 2.2 2.2 2.1 2.3 1.7 1.6
ML 1.2 1.1 1.1 1.4 1.1 1.0
TaL/SVL 1.18 1.12 — 0.93 1.22 —
AG/SVL 0.46 0.45 0.46 0.43 0.46 0.46
HL/SVL 0.26 0.26 0.25 0.25 0.25 0.26
HL/HW 1.20 1.22 1.15 1.27 1.18 1.22
HH/HL 0.47 0.41 0.40 0.41 0.42 0.39
SNT/HL 0.45 0.45 0.47 0.43 0.45 0.43
SNT/ED 1.65 1.58 1.70 1.54 1.52 1.58
ED/HL 0.27 0.29 0.28 0.28 0.29 0.27
EOD/ED 0.33 0.29 0.35 0.33 0.37 0.32
RW/RH 1.93 2.08 1.86 2.00 1.92 2.27
RW/MW 1.32 1.23 1.24 1.39 1.47 1.56
MW/ML 1.83 2.00 1.91 1.64 1.55 1.60
N 3/3 3/3 3/3 3/3 3/3 3/3
I 0 0 0 0 0 0
SPL 11/12 12/12 13/12 9/11 11/11 11/11
IFL 11/10 11/10 11/11 10/10 11/10 13/11
IO 27 26 24 22 25 25
PO 18/19 17/17 15/16 16/16 15/16 18/18
PM 2 2 2 2 2 2
GP 5 3 6 6 4 5
DTR 9 8 11 8 11 10
GSDT 10 9 9 9 9 9
SMC 206 197 205 213 198 193
SR 149 158 153 157 161 156
V 43 49 40 44 44 48
LF1 12/10 12/12 13/12 12/13 12/12 11/11
LF4 12/12 12/12 14/15 13/12 15/14 16/16
LT1 13/13 12/12 12/13 14/12 14/14 13/14
LT4 14/13 13/13 13/13 14/14 14/15 13/13
PP 12 11 11 9 — 12
PAT 1/1 1/1 1/1 1/1 1/1 1/1
S3W 10 9 9 11 10 10
Table 8.
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Scalation features comparisons among Gekko fengshanensis sp. nov., G. kwangsiensis, G. liboensis and G. paucituberculatus; differences are marked in bold.

Species G. fengshanensis sp. nov. G. kwangsiensis G. liboensis G. paucituberculatus
n = 6 n = 6 n = 2 n = 2
Max SVL (mm) 79.9 69.7 79.7 85.9
N 3 3 3 3
I 0 0 or 1 0 0
SPL 9–13 10–13 11 11
IFL 10–13 11–13 9–11 9–10
IO 22–27 29–31 32–35 37
PO 15–19 18–20 17–18 14–18
PM 2 2 2 2
GP 3–6 4–6 4–6 4–6
DTR 9–11 9–11 9–10 4
GSDT 9–10 8–10 9–10 8
SMC 197–213 185–208 183–195 189–192
SR 149–161 143–156 131–140 136–142
V 40–49 41–45 38–41 42–44
LF1 11–13 10–13 12–13 10–11
LF4 12–16 12–14 14–17 12–13
LT1 12–14 11–13 12–13 11
LT4 13–15 14–18 14–15 11–13
PP 9–12 9–10 9 12
PAT 1 1 1 1
Figure 7. 

Morphological variation of Gekko fengshanensis sp. nov. A. Paratype NHMG 202408007, note the discontinuous precloacal pores; B. Paratype NHMG 202408007, note the postmentals. Photos by HMS.

Etymology.

The specific epithet fengshanensis refers to Fengshan County, the type locality in Guangxi Zhuang Autonomous Region, China. The common name “Fengshan gecko” (English) and formal Chinese name “凤山壁虎” (fèng shān bì hǔ) are proposed.

Comparisons.

The molecular analyses indicated that Gekko fengshanensis sp. nov. is sister to G. kwangsiensis, and together they form a clade with G. liboensis and G. paucituberculatus, to which it is also morphologically similar. Morphological comparisons and analyses revealed their differences (Tables 3, 8). The new species differs from G. kwangsiensis in mean values of HL, ED, SNT, MW, ML (Table 3) and by having fewer interorbitals (22–27 vs 29–31); differs from G. liboensis by having fewer interorbitals (22–27 vs 32–35), more scales in a line from mental to the front of cloacal slit (197–213 vs 183–195), and more scale rows at midbody (149–161 vs 131–140); differs from G. paucituberculatus by having smaller maximum SVL (79.9 vs 85.9), fewer interorbitals (22–27 vs 37), more dorsal tubercle rows at midbody (9–11 vs 4), more scales in a line from mental to the front of cloacal slit (197–213 vs 189–192), more scale rows at midbody (149–161 vs 136–142), and coloration pattern (irregular light patches with lateral spots vs dirty-white transverse bands between the nape and sacrum).

For the remaining congeners, the new species differs from the following 13 congeners by the presence of tubercles on dorsolateral trunk: the absence of tubercles in Gekko aaronbaueri, G. bonkowskii, G. cib, G. guishanicus, G. khunkhamensis, G. melli, G. nadenensis, G. scientiadventura, G. sengchanthavongi, G. subpalmatus, G. tawaensis, G. thakhekensis, and G. truongi; differs from the following 13 congeners by having 9–12 precloacal pores in males: Gekko adleri (17–21), G. alpinus (4–7), G. canhi (5), G. chinensis (17–27), G. jinjiangensis (4–5), G. palmatus (23–30), G. shibatai (0), G. similignum (17), G. taibaiensis (4–6), G. vertebralis (0), G. vietnamensis (0), G. wenxianensis (6–8) and G. yakuensis (6–8); differs from G. kaiyai by the absence of tubercles on limbs (vs present); differs from G. hokouensis by having more scale rows at midbody (149–161 vs 119–130); differs from G. auriverrucosus and G. scabridus by having fewer dorsal tubercle rows (8–11 vs 16–20 and 17–21, respectively); differs from G. ichangensis, G. japonicus and G. swinhonis by having a single postcloacal tubercles (vs 3, 2–4 and 2 or 3, respectively).

Distribution and ecology.

Currently, Gekko fengshanensis sp. nov. is known only from Fengshan County, Hechi City, Guangxi Zhuang Autonomous Region, China. All six individuals were discovered at night on the walls of artificial buildings located near the karst forests (Fig. 8A). Other amphibian and reptile species co-occurring with it include Kurixalus hainanus (Zhao, Wang, & Shi, 2005), Pseudocalotes microlepis (Boulenger, 1888), and Sinomicrurus peinani Liu, Yan, Hou, Wang, Nguyen, Murphy, Che, & Guo, 2020.

Figure 8. 

A. Habitats of Gekko fengshanensis sp. nov.; B. The egg of Gekko fengshanensis sp. nov. Photos by ZH.

At the type locality, one milky-white, elliptical egg measuring ~1.5 cm in diameter was found in a crevice of a house (Fig. 8B). It is presumed to belong to this species, suggesting that the breeding season likely occurs in July and August.

Additional specimens examined.

Gekko liboensis (n = 2): SYS r002876 (adult male) and SYS r002877 (subadult female), from Weng’ang Station, Maolan National Nature Reserve, Libo County, Guizhou Province, China, collected on 24 July 2024 by Hao-Tian Wang and Ying-Yong Wang.

Gekko paucituberculatus (n = 2): SYS r002806 (adult male) and SYS r002807 (adult female), from Tianyang District, Baise City, Guangxi Zhuang Autonomous Region, China, collected on 3 August 2023 by Dan-Yang Zhou.

The GenBank accession numbers for the four specimens listed above can be found in Table 2, and the corresponding morphological data are provided in Suppl. material 1. These specimens are deposited in the Sun Yat-sen University (SYS), Guangzhou, China.

Discussion

The discovery of Gekko fengshanensis sp. nov. represents a significant addition to the biodiversity of the South China Karst, bringing the total number of recognized Gekko species inhabiting this unique karst ecosystem to five. Molecular phylogenetic analyses revealed that these species (Gekko fengshanensis sp. nov., G. kwangsiensis, G. liboensis, and G. paucituberculatus), excluding G. adleri, form a monophyletic lineage, indicating their shared evolutionary origin within karst habitats. This finding strongly supports the hypothesis that the fragmented nature of karst landscapes (e.g., isolated peak clusters, depressions, and canyons) has limited or only historical gene flow among populations, driving allopatric speciation (Chung et al. 2014; Grismer et al. 2020, 2021). The three species — G. fengshanensis sp. nov. (mid- to high-elevation canyons), G. kwangsiensis (isolated Daming Mountain, Youjiang River basin), and G. paucituberculatus (low-elevation limestone in the Baise Basin) — occupy ecologically distinct habitats across their largely allopatric ranges. Their distributions are confined to the western side of the Hongshui River, which, together with the Longjiang River, forms a biogeographic barrier that isolates them from G. liboensis populations to the east (Fig. 1). Dispersal among these species is likely constrained by intervening non-karst terrain, with the dual-river system coinciding with species boundaries and potentially contributing to the reinforcement of genetic divergence. The karst region’s complex landforms have played a crucial role in shaping biodiversity by creating fragmented and unique microhabitats. These diverse landscapes have fostered a high degree of endemism and species richness (Clements et al. 2006; Grismer et al. 2021). Grismer et al. (2021) further confirmed that karst landscapes actively drive species diversification within Cyrtodactylus, rather than merely serving as “imperiled arks of biodiversity” that act as refugia for relic species. This pattern aligns with the diversification of Gekko species in the South China Karst, where geographic isolation and habitat specialization have been key drivers of speciation.

Gekko liboensis was originally described by Zhou et al. (1982) based on a single female specimen from Libo County, Guizhou Province. However, its taxonomic validity has long been debated due to its extreme rarity, with only four specimens ever recorded (Zhao et al. 1999). Zhao and Adler (1993), Günther (1994), and Kluge (2001) considered it a junior synonym of G. hokouensis, whereas Kluge (1993), Bauer (1994), Matsui and Ota (1995), Zhao et al. (1999), and Rösler et al. (2011) recognized it as a valid species. Based on newly collected specimens from Libo, Guizhou (the type locality of G. liboensis), and Guangxi, Jono et al. (2015) employed morphometric methods to confirm the validity of G. liboensis, while also extending its known distribution to Guangxi. In the present study, we further validate its taxonomic status based on the molecular evidence of topotypes newly collected for this study. The G. liboensis specimens reported from Guangxi by Jono et al. (2015) were recorded in Nalao (24°25'N, 107°22'E), but both the locality name and its coordinates are imprecise. There are several villages named Nalao or with similar names in this vicinity (the “?” symbol in Fig. 1), specifically Nalang Village, Sanshi Town, Donglan County (24°24'35.95"N, 107°21'20.83"E); Nalao Village, Wuzuan Town, Donglan County (24°21'33.01"N, 107°16'49.26"E); and Nalao Village, Xishan Town, Bama Yao Autonomous County (24°14'13.12"N, 107°18'07.02"E). All these localities lie on the western side of the Hongshui River and are in proximity to the type locality of G. fengshanensis sp. nov. (~35 km). Given the striking morphological similarities between the two species, the specimens identified as G. liboensis by Jono et al. (2015) from Guangxi likely represent misidentifications of G. fengshanensis sp. nov. Consequently, the reported occurrence of G. liboensis in Guangxi remains questionable and necessitates further verification.

Acknowledgements

We are very grateful to Xiao-Wen Liao and Ben-Ze Huang for their help to fieldwork. We would like to express our sincere gratitude to Dr. Larry L. Grismer for conducting additional analyses on our dataset and kindly granting us permission to incorporate these results into our manuscript. Their valuable contribution has significantly enhanced the rigor and quality of this study. We also thank the editor Dr. Minh Duc Le and another reviewer Dr. Vinh Quang Luu for their insightful comments and suggestions, which greatly improved the manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

This work was supported by Wildlife Theme 2-Amphibian Reptiles of the Southwest Karst National Park (Guangxi) Comprehensive Scientific Investigation Project (LKWT-2023-095), Science and Technology Planning Projects of Guangdong Province (2021B1212110002) and Guangxi Natural Science Foundation (Grant No. 2023GXNSFDA026065).

Author contributions

All authors have contributed equally.

Author ORCIDs

Zhong Huang https://orcid.org/0009-0001-7825-4778

Hao-Tian Wang https://orcid.org/0000-0002-2415-3234

Shuo Qi https://orcid.org/0000-0002-2924-6093

Han-Ming Song https://orcid.org/0009-0005-8244-1567

Data availability

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

References

  • Cao J, Sucharitakul P, Tie MH, Suwannapoom C, Yan F, Chomdej S (2025) A new species of the genus Gekko Laurenti, 1768 (Squamata: Gekkonidae) from Hubei, China. Asian Herpetological Research 16(1): 110–121. https://doi.org/10.3724/ahr.2095-0357.2024.0046
  • Chung K-F, Leong W-C, Rubite RR, Repin R, Kiew R, Liu Y, Peng C-I (2014) Phylogenetic analyses of sect. Coelocentrum and allied limestone species of China shed light on the evolution of Sino-Vietnamese karst flora. Botanical Studies 55(1): 1. https://doi.org/10.1186/1999-3110-55-1
  • Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: More models, new heuristics and parallel computing. Nature Methods 9(8): 772–772. https://doi.org/10.1038/nmeth.2109
  • Edgar RC (2004) MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32(5): 1792–1797. https://doi.org/10.1093/nar/gkh340
  • Grismer LL, Wood Jr P, Le MD, Quah E, Grismer J (2020) Evolution of habitat preference in 243 species of Bent-toed geckos (Genus Cyrtodactylus Gray, 1827) with a discussion of karst habitat conservation. Ecology and Evolution 10(24): 13717–13730. https://doi.org/10.1002/ece3.6961
  • Grismer LL, Wood Jr PL, Poyarkov N, Le M, Karunarathna S, Chomdej S, Suwannapoom C, Qi S, Liu S, Che J, Quah E, Kraus F, Oliver P, Riyanto A, Pauwels O, Grismer JL (2021) Karstic Landscapes Are Foci of Species Diversity in the World’s Third-Largest Vertebrate Genus Cyrtodactylus Gray, 1827 (Reptilia: Squamata; Gekkonidae). Diversity 13(5): 1–15. https://doi.org/10.3390/d13050183
  • Grismer LL, Aowphol A, Grismer JL, Aksornneam A, Quah ESH, Murdoch ML, Gregory JJ, Nguyen E, Kaatz A, Bringsøe H, Rujirawan A (2024) A new species of the Cyrtodactylus chauquangensis group (Squamata, Gekkonidae) from the borderlands of extreme northern Thailand. ZooKeys 1203: 211–238. https://doi.org/10.3897/zookeys.1203.122758
  • Günther R (1994) Eine neue Art der Gattung Gekko (Reptilia, Squamata, Gekkonidae) aus dem Süden Vietnams. Zoologischer Anzeiger 233: 57–67. [in Germany]
  • Hou YM, Shi SC, Wang G, Shu GC, Zheng PY, Qi Y, Liu GH, Jiang JP, Xie F (2021) A New species of the Gekko japonicus group (Squamata: Gekkonidae) from southwest China. Asian Herpetological Research 12: 36–48. https://doi.org/10.16373/j.cnki.ahr.200064
  • Jonniaux P, Kumazawa Y (2008) Molecular phylogenetic and dating analyses using mitochondrial DNA sequences of eyelid geckos (Squamata: Eublepharidae). Gene 407(1-2): 105–115. https://doi.org/10.1016/j.gene.2007.09.023
  • Jono T, Ding L, Kaito T, Tang Y, Toda M (2015) Reevaluation of the taxonomic status of a poorly known gecko, Gekko liboensis (Reptilia: Squamata). Asian Herpetological Research 6: 229–236. https://doi.org/10.16373/j.cnki.ahr.150009
  • Kluge AG (1993) . Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego, 245 pp.
  • Kluge AG (2001) Gekkotan lizard taxonomy. Hamadryad 26: 1–209.
  • Lanfear R, Frandsen PB, Wright AM, Senfeld T, Calcott B (2017) PartitionFinder 2: New methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution 34: 772–773. https://doi.org/10.1093/molbev/msw260
  • Lin JT, Yao CJ (2016) A new species of the genus Gekko (Squamata: Sauria: Gekkonidae) from Guishan Isle, Yilan County, Taiwan. Journal of the National Taiwan Museum 69: 1–14. https://doi.org/10.6532/JNTM.2016.69(2).01 [in Chinese]
  • Luu VQ, Calame T, Nguyen TQ, Le MD, Bonkowski M, Ziegler T (2014) A new species of the Gekko japonicus group (Squamata: Gekkonidae) from central Laos. Zootaxa 3895(1): 73–88. https://doi.org/10.11646/zootaxa.3895.1.4
  • Luu VQ, Calame T, Nguyen TQ, Le MD, Ziegler T (2015) Morphological and molecular review of the Gekko diversity of Laos with descriptions of three new species. Zootaxa 3986(3): 279–306. https://doi.org/10.11646/zootaxa.3986.3.2
  • Luu VQ, Nguyen TQ, Le MD, Bonkowski M, Ziegler T (2017) A new karst dwelling species of the Gekko japonicus group (Squamata: Gekkonidae) from central Laos. Zootaxa 4263(1): 179–193. https://doi.org/10.11646/zootaxa.4263.1.10
  • Lyu ZT, Lin CY, Ren JL, Jiang K, Zhang YP, Qi S, Wang J (2021) Review of the Gekko (Japonigekko) subpalmatus complex (Squamata, Sauria, Gekkonidae), with description of a new species from China. Zootaxa 4951(2): 236–258. https://doi.org/10.11646/zootaxa.4951.2.2
  • Ma S, Shi S, Shen C, Chang L, 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
  • Matsui M, Ota H (1995) On Chinese herpetology. Herpetologica 51: 234–250.
  • Mo YM, Chen WC, Wu HY, Zhang W, Zhou SC (2015) A new species of Odorrana inhabiting complete darkness in a karst cave in Guangxi, China. Asian Herpetological Research 6: 11–17.
  • Nguyen TQ, Wang YY, Yang JH, Lehmann T, Le MD, Ziegler T, Bonkowski M (2013) A new species of the Gekko japonicus group (Squamata: Sauria: Gekkonidae) from the border region between China and Vietnam. Zootaxa 3652(5): 501–518. https://doi.org/10.11646/zootaxa.3652.5.1
  • Oksanen J, Blanchet FG, Friendly M, Kindt R, Legendre P, McGlinn D, Minchin PR, O’Hara RB, Simpson GL, Solymos P, Stevens MHH, Szoecs E, Wagner H (2020) Package ‘vegan’. Version 2.5–7. https://cran.r-project.org/web/packages/vegan/
  • Ota H, Lau MW, Weidenhöfer T, Yasukawa Y, Bogadek A (1995) Taxonomic review of the geckos allied to Gekko chinensis Gray 1842 (Gekkonidae Reptilia) from China and Vietnam. Tropical Zoology 8(1): 181–196. https://doi.org/10.1080/03946975.1995.10539278
  • Pauwels OSG, Meesook W, Donbundit N, Jindamad T, Topai N, Sumontha M (2025) Gekko (Gekko) shiva, a new limestone-dwelling gecko from Sa Kaeo Province, eastern Thailand (Squamata, Gekkonidae). Zootaxa 5588(2): 305–322. https://doi.org/10.11646/zootaxa.5588.2.6
  • Qiu XC, Zhou SB, Qi S, Wang XP, Wang HT, Wang JZ, Shi JS, Li PP (2023) The Updated Checklist and Zoogeographic Division of the Reptilian Fauna of Liaoning Province. Dongwuxue Zazhi 58(4): 523–539. https://doi.org/10.13859/j.cjz.202304006 [in Chinese]
  • R Core Team (2024) . R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/
  • Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029
  • Rösler H, Bauer AM, Heinicke MP, Greenbaum E, Jackman T, Nguyen TQ, Ziegler T (2011) Phylogeny, taxonomy, and zoogeography of the genus Gekko Laurenti, 1768 with the revalidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae). Zootaxa 2989(1): 1–50. https://doi.org/10.11646/zootaxa.2989.1.1
  • Simon C, Frati F, Beckenbach A, Crespi B, Liu H, Flook P (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87(6): 651–701. https://doi.org/10.1093/aesa/87.6.651
  • Sitthivong S, Lo OV, Nguyen TQ, Ngo HT, Khotpathoom T, Le MD, Ziegler T, Luu VQ (2021) A new species of the Gekko japonicus group (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa 5082(6): 553–571. https://doi.org/10.11646/zootaxa.5082.6.3
  • Song MT (1985) A new species of Gekko from Shaanxi. Acta Herpetologica Sinica 4: 329–330. [in Chinese]
  • Song HM, Wang HT, Lin ZH, Zheng SS, Zhou ZE, Gong YN, Liu ZF, Li YY, Qi S, Wang YY (2024) Eight New Records of Amphibians and Reptiles in Guangdong Province, China. Dongwuxue Zazhi 59(5): 658–686. https://doi.org/10.13859/j.cjz.202424014 [in Chinese]
  • Tamura K, Stecher G, Kumar S (2021) MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Battistuzzi FU (Ed.). Molecular Biology and Evolution 38: 3022–3027. https://doi.org/10.1093/molbev/msab120
  • Toda M, Sengoku S, Hikida T, Ota H (2008) Description of two new species of the genus Gekko (Squamata: Gekkonidae) from the Tokara and Amami Island Groups in the Ryukyu Archipelago, Japan. Copeia 2008(2): 452–466. https://doi.org/10.1643/CH-06-281
  • Wang HT, Qi S, Zhou DY, Wang YY (2024) Description of a new karst-adapted species of the subgenus Japonigekko (Squamata: Gekkonidae: Gekko) from Guangxi, southern China. Vertebrate Zoology 74: 121–132. https://doi.org/10.3897/vz.74.e113899
  • Wood Jr PL, Guo X, Travers SL, Su Y, Olson KV, Bauer AM, Grismer LL, Siler CD, Moyle RG, Andersen MJ, Brown RM (2020) Parachute geckos free fall into synonymy: Gekko phylogeny, and a new subgeneric classification, inferred from thousands of ultraconserved elements. Molecular Phylogenetics and Evolution 146: 106731. https://doi.org/10.1016/j.ympev.2020.106731
  • Zhang CW, Wu AF, Cai B, Wang LR, Pang DP, Ma HH, Yu L, Li XY, Huang H, Zeng L, Li L, Yan J, Li P, Zhang BW (2023) A New Species of the Genus Gekko (Squamata: Sauria: Gekkonidae) from the Dabie Mountains, China. Animals 13(24): 3796. https://doi.org/10.3390/ani13243796
  • Zhao E, Adler K (1993) Herpetology of China. Oxford, Ohio, 522 pp[+48plates].
  • Zhao E, Zhao K, Zhou K (1999) Fauna Sinica Reptilia, Vol. 2: Squamata, Lacertilia. Science Press, Beijing, [xi +] 394 pp. [in Chinese]
  • Zhou KY, Liu YZ, Li DJ (1982) Three new species of Gekko and remarks on Gekko hokouensis (Lacertiformes, Gekkonidae). Acta Zootaxonomica Sinica 7: 438–446. [in Chinese]

Zhong Huang and Hao-Tian Wang contributed equally to this work.

Supplementary material

Supplementary material 1 

Raw morphological data used in PCA and MFA analyses

Zhong Huang, Hao-Tian Wang, Shuo Qi, Han-Ming Song, Yong Huang, Ying-Yong Wang, Yun-Ming Mo

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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