Research Article
Print
Research Article
Description of a new karst-adapted species of the subgenus Japonigekko (Squamata: Gekkonidae: Gekko) from Guangxi, southern China
expand article infoHao-Tian Wang‡, Shuo Qi‡, Dan-Yang Zhou§, Ying-Yong Wang‡
‡ Sun Yat-sen University, Guangzhou, China
§ Unaffiliated, Zhejiang, China

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

Academic editor: Uwe Fritz
© 2024 Hao-Tian Wang, Shuo Qi, Dan-Yang Zhou, Ying-Yong Wang.
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: Wang H-T, Qi S, Zhou D-Y, Wang Y-Y (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
ZooBank: urn:lsid:zoobank.org:pub:BCDCB720-4442-4AFC-97CE-6F2DAD451FF0
Open Access

Abstract

A new species of the genus Gekko Laurenti, 1768, Gekko paucituberculatus sp. nov., is described here, based on two specimens from Tianyang District, Baise City, Guangxi Zhuang Autonomous Region, China. It was placed in subgenus Japonigekko on the basis of morphological and molecular phylogenetic analysis and can be distinguished from all congeners in this subgenus by significant divergences in the mitochondrial 16S and ND2 genes and by a combination of the following morphological characters: Moderate body size, SVL 77.2 mm in the adult male and SVL 85.9 mm in the adult female; tubercles only present along dorsolateral trunk and absent on other regions; fingers and toes with weak webbing; continuous precloacal pores 12 in the male, absent in the female; a single postcloacal tubercle on each side; a light-coloured vertebral line from nape to tip of tail; dorsum greyish-brown, with 7–8 dirty-white bands between nape and sacrum. Meanwhile, the distribution of G. palmatus in China has been confirmed as occurring in Guangxi and Guangdong Provinces. This study brings the total species of the subgenus Japonigekko in China to 19.

Key words

Gekko paucituberculatus sp. nov., Gekko palmatus, karst, new national record, taxonomy

Introduction

The genus Gekko Laurenti, 1768, currently containing 88 recognised species assigned to seven subgenera, is a widely distributed group of nocturnal lizards (Wood et al. 2020; Uetz et al. 2023; Zhang et al. 2023). Amongst these subgenera, G. (Japonigekko) is the most diverse group with many karst-dwellers. At present, there are three described G. (Japonigekko) species-adapted karst ecosystems in China, namely G. adleri Nguyen et al., 2013 and G. kwangsiensis Yang, 2015 from Guangxi and G. liboensis Zhou & Li, 1982 from Guizhou and Guangxi (Nguyen et al. 2013; Jono et al. 2015; Yang 2015). During the field survey in the karst forest of western Guangxi, southern China (Fig. 1), two gekkonid individuals were collected. Morphologically, the specimens possess the common characteristics of subgenus G. (Japonigekko), but are apparently distinct from all known congeners by the following characters: Moderate body size, tubercles only present along dorsolateral trunk, continuous precloacal pores 12 in the male, a single postcloacal tubercle on each side and a light-coloured vertebral line from nape to tip of tail. Phylogenetic studies reveal that these specimens belong to a separated taxon most closely related to G. kwangsiensis. Hence, we consider it as a new species and describe it below.

Figure 1. 

Localities of Gekko paucituberculatus sp. nov. and G. palmatus. 1 Tianyang District, Baise City, Guangxi; 2 Napo County, Baise City, Guangxi; 3 Nonggang Nature Reserve, Chongzuo City, Guangxi; 4 Mt. Dinghu, Zhaoqing City, Guangdong.

Materials and Methods

Specimens and morphology

Two specimens were collected from Tianyang District, Baise City, Guangxi Zhuang Autonomous Region, China on 3 August 2023. The specimens were euthanised and then fixed in 10% buffered formalin, later being transferred to 75% ethanol and deposited in the Museum of Biology, Sun Yat-sen University (SYS), Guangzhou, China. Liver tissue samples were preserved in 95% ethanol for molecular analysis.

Measurements were taken with digital callipers (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 follow Lyu et al. (2021) and Grismer et al. (2022). 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, maximum head length from tip of snout to posterior margin of ear opening); head width (HW, measured 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 width (MW); maximum mental length (ML); 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 corners of eyes); preorbitals (PO, number of scales in a line from nostril to anterior corner of the eye); postmentals (PM, scales bordering the mental); gulars bordering the postmentals (GP); dorsal tubercle rows at mid-body (DTR); granules surrounding dorsal tubercles (GSDT); scales in a line from mental to the front of cloacal slit (SMC); ventral scale rows at mid-body (V); scale rows at mid-body (SR, including ventral scales); subdigital lamellae under entire first finger (LF1); subdigital lamellae under entire fourth finger (LF4); subdigital lamellae under entire first toe (LT1); subdigital lamellae under entire fourth toe (LT4); precloacal pores (PP); postcloacal tubercles (PAT); dorsal scale rows in the middle of the third caudal whorl (S3W).

In addition, the specimens were compared with other Gekko (Japonigekko) congeners on the basis of descriptions in literature (Zhou et al. 1982; Song 1985; Ota et al. 1995; Toda et al. 2008; Phung and Ziegler 2011; Rösler et al. 2011; Nguyen et al. 2013; Luu et al. 2014, 2015, 2017; Jono et al. 2015; Ngo et al. 2015; Yang 2015; Lin and Yao 2016; Hou et al. 2021; Lyu et al. 2021; Sitthivong et al. 2021; Zhang et al. 2023).

Phylogenetic sampling and analyses

A total of 24 samples of subgenus Gekko (Japonigekko) were used for molecular analysis in this study. All samples were attained from euthanasia specimens and then preserved in 95% ethanol and stored at -40°C. Furthermore, 22 sequences were obtained from GenBank and incorporated into our dataset for phylogenetic analysis. G. gecko and G. reevesii, belonging to subgenus G. (Gekko), were used to root the tree, based on Rösler et al. (2011) and Lyu et al. (2021). Detailed information of these samples is given in Table 1.

Table 1.
Download as
CSV
XLSX

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

ID Species Localities Voucher ID GenBank accession numbers References
16S ND2
Ingroups
1 Gekko paucituberculatus sp. nov. China: Guangxi: Baise, Tianyang SYS r002806 OR903154 OR902163 This study
2 Gekko paucituberculatus sp. nov. China: Guangxi: Baise, Tianyang SYS r002807 OR903155 OR902164 This study
3 G. cib China: Sichuan: Hejiang SYS r001489 MW451655 OR902165 Lyu et al. (2021), this study
4 G. cib China: Sichuan: Mt. Emei SYS r000708 MW451629 OR902166 Lyu et al. (2021), this study
5 G. subpalmatus China: Zhejiang: Fenghua SYS r001762 MW451662 OR902167 Lyu et al. (2021), this study
6 G. subpalmatus China: Zhejiang: Fenghua SYS r001767 MW451663 OR902168 Lyu et al. (2021), this study
7 G. melli China: Guangdong: Dongyuan SYS r001742 MW451661 OR902169 Lyu et al. (2021), this study
8 G. melli China: Guangdong: Puning SYS r001702 MW451660 OR902170 Lyu et al. (2021), this study
9 G. swinhonis China: Hebei: Zunhua SYS r001814 MW451666 OR902171 Lyu et al. (2021), this study
10 G. hokouensis China: Jiangxi: Mt. Meiling SYS r001311 MW451648 OR902172 Lyu et al. (2021), this study
11 G. hokouensis China: Fujian: Mt. Wuyi SYS r001290 MW451647 OR902173 Lyu et al. (2021) this study
12 G. kwangsiensis China: Guangxi: Wuming SYS r001194 MW451641 OR902174 Lyu et al. (2021), this study
13 G. kwangsiensis China: Guangxi: Wuming SYS r001195 MW451642 OR902175 Lyu et al. (2021) this study
14 G. japonicus China: Fujian: Mt. Wuyi SYS r000672 MW451628 OR902176 Lyu et al. (2021), this study
15 G. japonicus China: Jiangxi: Lushan SYS r001317 MW451649 OR902177 Lyu et al. (2021), this study
16 G. adleri Vietnam: Cao Bang IEBR A.2012.24 KC700623 — Nguyen et al. (2013)
17 G. adleri China: Guangxi: Jingxi SYS r001400 MW451654 OR902178 Lyu et al. (2021), this study
18 G. palmatus China: Guangdong: Mt.Dinghu SYS r002797 OR903156 OR902179 This study
19 G. palmatus China: Guangdong: Mt.Dinghu SYS r002804 OR903157 OR902180 This study
20 G. palmatus Vietnam: Lang Son IEBR 2474 KC710234 — Nguyen et al. (2013)
21 G. palmatus Vietnam: Lang Son IEBR 3620 KC710238 — Nguyen et al. (2013)
22 G. palmatus Vietnam: Bac Giang IEBR A.0807 KC710233 — Nguyen et al. (2013)
23 G. palmatus China: Guangxi: Nonggang Nature Reserve SYS r001192 MW451639 OR902181 Lyu et al. (2021), this study
24 G. palmatus China: Guangxi: Napo SYS r001185 MW451637 OR902182 Lyu et al. (2021), this study
25 G. chinensis China: Hong Kong SYS r001211 MW451644 OR902183 Lyu et al. (2021), this study
26 G. chinensis China: Guangdong: Shenzhen SYS r001085 MW451632 OR902184 Lyu et al. (2021), this study
27 G. similignum China: Hainan: Mt. Wuzhi SYS r001597 MW451658 OR902185 Lyu et al. (2021), this study
28 G. similignum China: Hainan: Mt. Wuzhi SYS r001598 MW451659 OR902186 Lyu et al. (2021), this study
29 G. auriverrucosus China: Shanxi: Yuncheng NNU Z 20050716.004 — JN019062 Rösler et al. (2011)
30 G. bonkowskii Laos: Khammouane VFU R.2014.10 — KT266818 Luu et al. (2015)
31 G. khunkhamensis Laos: Khammouane VNUF R.2021.23 — OL416111 Sitthivong et al. (2021)
32 G. khunkhamensis Laos: Khammouane VNUF R.2021.02 — OL416109 Sitthivong et al. (2021)
33 G. jinjiangensis China: Yunnan: Deqin CIB5334220115 — MT449431 Hou et al. (2021)
34 G. jinjiangensis China: Sichuan: Derong CIB5133380017 — MT449437 Hou et al. (2021)
35 G. scabridus China: Sichuan:Yanbian CIBYN201909199 — MT449429 Hou et al. (2021)
36 G. scabridus China: Sichuan:Yanbian CIBYN201909200 — MT449430 Hou et al. (2021)
37 G. scientiadventura Vietnam: Quang Binh IEBR A.2014.7 — KP205392 Luu et al. (2014)
38 G. scientiadventura Laos: Khammouane VFU2014.2 — KP205394 Luu et al. (2014)
39 G. sengchanthavongi Laos: Khammouane VFU R2014.14 — KT266816 Luu et al. (2015)
40 G. sengchanthavongi Laos: Khammouane IEBR A.2015.33 — KT266817 Luu et al. (2015)
41 G. thakhekensis Laos: Khammouane:Thakhek IEBR A.2014.6 — KP205396 Luu et al. (2014)
42 G. thakhekensis Laos: Khammouane:Thakhek VFU R.2014.9 — KP205397 Luu et al. (2014)
43 G. truongi Vietnam: Khanh Hoa IEBR A.2011.1 — KP205398 Luu et al. (2014)
44 G. nadenensis Laos: Khammouane ZFMK 98741 — KY421618 Luu et al. (2017)
Outgroups
45 G. gecko China: Guangxi: Nanning N/A AY282753 AY282753 Zhou et al. (2006)
46 G. reevesii China: Guangdong: Mt. Yinping SYS r000796 MW451630 OR902187 Lyu et al. (2021), this study

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 the partial 16S ribosomal RNA gene (16S) and partial NADH dehydrogenase subunit 2 gene (ND2) were amplified. The primers used for two genes are listed in Table 2. PCR amplifications were processed with the following cycling conditions: 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.

Table 2.
Download as
CSV
XLSX

Primers used in this study.

Gene Primer Sequence Reference
16S L3975 5’-CGCCTGTTTACCAAAAACAT-3’ Simon et al. (1994)
H4551 5’-CCGGTCTGAACTCAGATCACGT-3’
ND2 rMet-3L 5’- ATACCCCGACAATGTTGG-3’ Jonniaux and Kumazawa (2008)
rAla-1H 5’- GCCTTAGCTTAATTAAAGTG-3’

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. ND2 was further partitioned by codon position and the best fit models for all partitions was GTR+I+G. Sequenced data were analysed 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 amongst and within species using the complete deletion option which removes missing data and gaps (Tamura et al. 2021).

Results

The aligned dataset contained a total of 1578 nucleotide base pairs (bp), with 566 bp for 16S and 1012 bp for ND2. The BI and ML analyses resulted in essentially identical topologies (Fig. 2). The mean uncorrected p distances, based on 16S and ND2 used in this study, are given in Tables 3 and 4.

Table 3.
Download as
CSV
XLSX

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

Species 1 2 3 4 5 6 7 8 9 10 11 12
1 G. paucituberculatus sp. nov. 0.00
2 G. cib 9.76 0.00
3 G. subpalmatus 13.15 6.37 0.00
4 G. melli 10.96 4.68 6.87 0.60
5 G. swinhonis 13.94 10.16 12.15 10.76 —
6 G. hokouensis 9.56 10.46 12.35 10.56 12.85 0.40
7 G. kwangsiensis 8.96 11.45 12.25 11.45 14.44 10.66 0.40
8 G. japonicus 12.35 11.55 11.95 11.75 14.14 10.76 11.85 0.00
9 G. adleri 11.75 11.55 12.75 10.76 14.14 12.15 12.35 13.35 0.40
10 G. palmatus 10.70 10.90 12.29 10.70 14.17 11.60 11.81 12.89 2.59 0.34
11 G. chinensis 10.26 10.26 11.65 10.46 13.84 11.25 10.96 11.45 3.49 3.09 0.20
12 G. similignum 10.96 10.96 11.75 10.96 14.14 11.16 11.85 11.16 3.78 3.39 1.29 0.00
Table 4.
Download as
CSV
XLSX

Uncorrected p distances (%) of the ND2 gene amongst species of Gekko (Japonigekko) used in this study.

Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
1 G. paucituberculatus sp. nov. 0.00
2 G. cib 19.02 0.00
3 G. subpalmatus 20.45 17.38 0.00
4 G. melli 21.06 17.59 14.52 2.66
5 G. swinhonis 21.47 21.27 20.25 20.45 —
6 G. hokouensis 18.92 22.09 20.76 20.86 19.94 1.84
7 G. kwangsiensis 15.54 18.61 22.09 21.68 19.84 20.86 0.41
8 G. japonicus 19.84 23.11 21.68 21.27 20.86 21.06 22.29 0.00
9 G. adleri 22.09 23.72 23.72 20.14 22.70 21.88 21.06 23.52 —
10 G. palmatus 21.32 24.18 25.10 23.16 22.75 21.37 21.47 22.96 6.44 0.82
11 G. chinensis 22.70 23.62 23.21 21.88 23.01 22.29 21.68 22.60 14.72 15.39 0.82
12 G. similignum 22.39 24.64 23.82 22.49 22.39 22.80 21.68 23.01 14.83 15.18 4.19 0.20
13 G. auriverrucosus 18.61 19.22 20.04 21.57 18.81 20.14 21.68 19.02 24.13 23.57 25.36 26.69 —
14 G. bonkowskii 18.81 19.22 20.86 21.78 21.68 19.63 21.37 20.45 24.54 23.01 20.96 22.39 21.88 —
15 G. khunkhamensis 21.47 20.86 22.49 23.52 23.52 23.93 21.47 22.49 25.77 25.66 24.23 24.44 24.13 15.24 0.41
16 G. jinjiangensis 17.79 21.37 20.55 20.04 20.76 19.43 19.33 17.89 20.86 20.86 22.09 21.68 20.65 18.61 21.37 2.45
17 G. scabridus 17.79 21.57 20.86 19.94 20.35 17.48 20.25 18.00 19.02 19.73 20.76 20.76 19.53 20.04 22.09 10.53 1.23
18 G. scientiadventura 19.02 21.17 21.37 21.78 22.80 21.47 20.55 21.98 24.34 23.62 22.24 22.60 20.86 12.99 14.42 18.40 18.71 3.68
19 G. sengchanthavongi 19.22 21.68 21.27 22.09 21.88 22.90 20.86 21.68 23.93 23.57 21.98 22.60 20.65 13.91 15.54 19.53 19.02 9.92 0.00
20 G. thakhekensis 17.79 20.45 20.65 20.76 23.52 21.06 20.65 20.14 21.78 22.44 20.25 20.65 21.17 6.95 15.95 19.02 19.33 12.37 12.88 0.61
21 G. truongi 20.86 21.68 21.47 20.45 23.72 21.98 22.19 22.29 19.84 21.32 20.04 20.35 24.54 22.09 22.09 20.35 18.40 21.88 21.88 20.76 —
22 G. nadenensis 18.61 20.86 19.43 20.76 21.68 22.29 20.55 21.27 23.52 22.96 21.57 22.39 21.06 6.95 13.70 18.00 19.63 13.39 12.07 6.95 22.29 —
Figure 2. 

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

All Gekko (Japonigekko) samples used in this study formed a strongly-supported monophyletic lineage (BPP = 1.00; BS = 100). However, intrageneric relationships within this lineage remained unclear due to low intrageneric nodal supports. Two samples from Baise City, Guangxi were placed in subgenus G. (Japonigekko), which clustered together with strong support (BPP = 1.00; BS = 100) and exhibited low intra-lineage genetic differentiation (0%). This unnamed lineage is the sister taxon to G. kwangsiensis from Wuming County, Guangxi and these together formed the sister group to G. hokouensis Pope, 1928. The lowest genetic distance between the unnamed lineage and a known Gekko species were, on average, 8.96% in 16S and 15.54% in ND2, as compared to G. kwangsiensis, although we did not use genetic divergences to make species delimitation, as it served as a reference only. The newly-collected specimens showed recognisable morphological differences from all known congeners (see Taxonomic account below). Hence, the unnamed population from Baise City is considered as an undefined species.

In addition, two samples from Mt. Dinghu, Guangdong grouped together with other G. palmatus Boulenger, 1907 samples from Vietnam and Guangxi, China and exhibit a robust monophyletic lineage with strong support (BPP = 1.00; BS = 100) and low intrapopulational genetic differentiation (0.34% in 16S and 0.82% in ND2). Morphologically, the characters of the specimens from Guangdong mostly agree with the original description and re-description of G. palmatus Boulenger, 1907 (Ota et al. 1995). Combined with results of Lyu et al. (2021), we regard G. palmatus as representing a new national record from China, occurring in Guangdong and Guangxi.

Taxonomic account

Gekko paucituberculatus sp. nov.

https://zoobank.org/C2814152-01A1-4558-892B-9D538ED1FE9D
Figures 3, 4 and 5

Holotype

SYS r002806 (Figs 3A and 4), adult male, from Tianyang District (23°42′55″N, 106°59′14″E; 120 m a.s.l.), Baise City, Guangxi Zhuang Autonomous Region, China, collected on 3 August 2023 by Dan-Yang Zhou.

Figure 3. 

Type specimens of Gekko paucituberculatus sp. nov. in life. A Holotype SYSr002806, adult male; B paratype SYSr002807, adult female. Photos by Dan-Yang Zhou.

Figure 4. 

Morphological features of the adult male holotype SYS r002806 of Gekko paucituberculatus 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 dorsal scalation at mid-body; I ventral tail. Photos by Hao-Tian Wang and Han-Ming Song.

Paratype

SYS r002807 (Figs 3B and 5), adult female, data identical to the holotype.

Diagnosis

Gekko paucituberculatus sp. nov. is distinguished from all congeners in the subgenus G. (Japonigekko) by a combination of the following morphological characters: (1) moderate body size, SVL 77.2 mm in the adult male and SVL 85.9 mm in the adult female; (2) nares in contact with rostral, internasal absent; (3) two enlarged postmentals; (4) tubercles flattened, only present along dorsolateral trunk and absent on other regions; (5) ventral scales between mental and cloacal slit 189–192; (6) mid-body scale rows 136–140; (7) ventral scale rows 42–44; (8) subdigital lamellae on first fingers 10–11, on fourth fingers 12–13, on first toes 11, on fourth toes 11–13 and fingers and toes webbing weakly developed; (9) continuous precloacal pores 12 in the male, absent in the female; (10) a single postcloacal tubercle on each side; (11) a light-coloured vertebral line from nape to tip of tail; (12) dorsum greyish-brown, with 7–8 dirty-white bands between nape and sacrum.

Etymology

The specific name paucituberculatus means few tubercles in Latin and refers to its tubercles being fewer than other congeners. According to its type locality, we suggest the common name as “Baise gecko” in English and Chinese formal name as “bǎi sè bì hǔ” (百色壁虎).

Description of holotype

Adult male, moderate size, SVL 77.2 mm; head depressed (HH/HL 0.38), length longer than width (HL/HW 1.24), distinct from neck; snout rounded at tip, elongate (SNT/HL 0.44), larger than eye (SNT/ED 1.69); rostral regular rectangular, nearly twice as wide as high (RW/RH 1.94) and wider than mental (RW/MW 1.22), with mid-dorsal notch approximately one third; nares oval, rounded by rostral, first supralabial, supranasal and two enlarged nasals posteriorly; internasals absent; preorbitals 18/18, preorbital region deeply concave; eye large (ED/HL 0.26), pupil vertical, margins crenulated; interorbital scales between anterior corners of eyes 37; ear opening elliptical, obliquely orientated, moderate in size (EOD/ED 0.40); mental pentagonal, wider than long (MW/ML 1.42); postmentals two, hexagonal and enlarged, twice as long as wide, touching mental and first infralabial on both sides and six gular scales posteriorly; supralabials 11/11; infralabials 10/10; tubercles absent on dorsal head, granulars on anteriodorsal head larger than those on posterior.

Body slender, elongate (AG/SVL 0.41); dorsals smooth, round or oval, granular and juxtaposed; tubercles flattened, only present on dorsal lateral surface, two rows on each side, surrounded by eight dorsal scales, absent on other regions; lateral fold present, without tubercles; ventrals distinctly larger than dorsal scales, smooth, imbricate and largest in middle of belly; ventral scale rows at mid-body 42; scale rows around mid-body 140; ventral scales in a row between mental and cloacal slit 189; precloacal scales enlarged, but no enlarged scales on thighs; precloacal pores 12, in a continuous row; postcloacal tubercle 1/1, large.

Fore- and hind-limbs well-developed; tubercles absent on dorsal 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, under manus 11-11-12-13-10 (left) and 11-11-12-13-11 (right), under pes 11-12-12-13-13 (left) and 11-11-12-13-12 (right); relative length fingers and toes I < II < V< III < IV.

Original tail (broken when capturing), longer than body (TaL 89.3 mm, TaL/SVL 1.16); distinctly swollen at base, oval in section; dorsal scales small, flat, smooth; caudal whorl distinct, 10 dorsal scale rows in the middle of the third one; subcaudals enlarged, arranged in a longitudinal row.

Colouration of holotype

In life, the dorsal regions of head and body are greyish-brown in colour, with scattered white spots on the anterior of head. An inverted U-shaped marking is present on the occipital region and there are seven regularly arranged, dirty-white bands between the nape and sacrum. The first band, located at the nape, extends forwards and backwards to the posterior corners of eye and the second band, respectively. A light-coloured vertebral line is present from the nape to the tail terminal. Some light spots are visible on the lateral sides. Limbs are light brown with many indistinct pale marks. Dorsal tail is black with seven bands, from dirty white to pure white in colour towards terminal. The ventral surface of the holotype is light flesh-coloured. The body colour becomes darker after capture.

In preservative, the dorsal ground colour of head, body and limbs is greyish-black; ventral surface fading to greyish-white.

Morphological variation

Measurements and scale counts of two individuals are shown in Tables 5 and 6 and the female paratype is shown in Figure 5.

Table 5.
Download as
CSV
XLSX

Measurements (in mm) and body proportions of the type series of Gekko paucituberculatus sp. nov. See Materials and Methods section for abbreviations. “*” regenerated tail.

Holotype SYS r002806 Paratype SYS r002807
Sex ♂ ♀
SVL 77.2 85.9
TaL 89.3 83.9*
AG 31.9 38.8
HL 20.0 21.2
HW 16.1 17.4
HH 7.6 8.4
SNT 8.8 9.8
ED 5.2 5.7
EOD 2.1 2.3
RH 1.7 2.0
RW 3.3 3.6
MW 2.7 2.8
ML 1.9 2.2
TaL/SVL 1.16 0.98
AG/SVL 0.41 0.45
HL/SVL 0.26 0.25
HL/HW 1.24 1.22
HH/HL 0.38 0.40
SNT/HL 0.44 0.46
SNT/ED 1.69 1.72
ED/HL 0.26 0.27
EOD/ED 0.40 0.40
RW/RH 1.94 1.80
RW/MW 1.22 1.29
MW/ML 1.42 1.27
Table 6.
Download as
CSV
XLSX

Scalation features of the type series of Gekko paucituberculatus sp. nov. Bilateral scale counts are given as left/right.

Holotype SYS r002806 Paratype SYS r002807
N 3/3 3/3
I 0 0
SPL 11/11 11/11
IFL 10/10 9/10
IO 37 37
PO 18/18 15/14
PM 2 2
GP 6 4
DTR 4 4
GSDT 8 8
SMC 189 192
SR 140 136
V 42 44
LF1 11/11 10/10
LF4 13/13 12/12
LT1 11/11 11/11
LT4 13/13 12/11
PP 12 —
PAT 1/1 1/1
S3W 10 9
Figure 5. 

Morphological features of the adult female paratype SYS r002807 of Gekko paucituberculatus sp. nov. A Dorsal view of body; B ventral view of body; C dorsal head; D ventral head; E right hand; F right foot; G dorsal scalation at mid-body. Photos by Hao-Tian Wang and Han-Ming Song.

Precloacal pores are absent in the female. In the male, the postcloacal tubercle is significantly larger than in the female. The paratype specimen exhibits eight light bands between the nape and sacrum.

Comparisons

Gekko paucituberculatus sp. nov. is compared with all 32 recognised species within the subgenus G. (Japonigekko).

The new species can be easily distinguished from the following 13 congeners by the presence of tubercles on the dorsolateral trunk: Absence of tubercles in G. aaronbaueri Tri et al., 2015, G. bonkowskii Luu et al., 2015, G. cib Lyu et al., 2021, G. guishanicus Lin & Yao, 2016, G. khunkhamensis Sitthivong et al., 2021, G. melli Vogt, 1922, G. nadenensis Luu et al., 2017, G. scientiadventura Rösler et al., 2004, G. sengchanthavongi Luu et al., 2015, G. subpalmatus Günther, 1864, G. tawaensis Okada, 1956, G. thakhekensis Luu et al., 2014, and G. truongi Phung & Ziegler, 2011.

The new species can be easily distinguished from the following 12 congeners by having 12 precloacal pores in the male: Gekko adleri (17–21), G. canhi Rösler et al., 2010 (5), G. chinensis Gray, 1842 (17–27), G. jinjiangensis Hou et al., 2021 (4–5), G. palmatus (23–30), G. shibatai Toda, Sengoku, Hikida & Ota, 2008 (0), G. similignum Smith, 1923 (17), G. taibaiensis Song, 1985 (4–6), G. vertebralis Toda, Sengoku, Hikida & Ota, 2008 (0), G. vietnamensis Sang, 2010 (0), G. wenxianensis Zhou & Wang, 2008 (6–8) and G. yakuensi Matsui & Okada, 1968 (6–8).

The new species can be easily distinguished from the following three congeners by having four dorsal tubercle rows: Gekko auriverrucosus Zhou & Liu, 1982 (16–20), G. hokouensis (12–18), G. kaiya Zhang et al., 2023 (11–18), G. scabridus Liu & Zhou, 1982 (17–21).

For the remaining congeners, by having a single postcloacal tubercles, the new species differs from G. japonicus (Schlegel, 1836) (2–4) and G. swinhonis Günther, 1864 (2–3).

The new species is most similar to G. kwangsiensis and G. liboensis, which are also from karst areas in Guangxi, but it differs from the former by the following characters: Less dorsal tubercle rows (4 vs. 9–11); more interorbital scales (37 vs. 29–31); more precloacal pores in male (12 vs. 9–10); fewer subdigital lamellae on fourth toes (11–13 vs. 13–18); fewer, but broader bands between nape and sacrum (7–8 vs. 9–10).

Furthermore, the new species differs from G. liboensis by tubercles only present along the dorsolateral surface not on other regions (vs. present from occipital region to tail base) and having regular and broad bands between nape and sacrum (vs. irregular and thin bands with many scattered round-shaped spots).

Distribution and ecology

Currently, Gekko paucituberculatus sp. nov. is limited to Tianyang District, Baise City, in Guangxi Zhuang Autonomous Region of China. The new gecko species is a rock-dwelling specialist. Both of the two individuals were discovered on rocks near the entrance to a limestone karst cave at night.

Discussion

Gekko palmatus was initially described by G. A. Boulenger in 1907 (as Gecko palmatus), based on a single female specimen from the mountains of Man Son, Vietnam (Boulenger 1907). Poyarkov et al. (2023) cited Nguyen et al. (2009) who argued that G. palmatus occurs in China, but it was not recognised due to lack of evidence (Wang et al. 2020; Cai et al. 2022). Both this study and Lyu et al. (2021) confirmed distributions of G. palmatus in Napo County and Nonggang Natural Reserve in Guangxi and Mt. Dinghu in Guangdong of China. Regarding morphology, the specimens from Guangdong share the diagnoses with G. palmatus: Nostril in contact with rostral; internasals 0–2, much smaller than supranasal when present; tubercles present on dorsum of body, lacking on forelimb and thigh; webs well developed; male preanal pores 24–27; cloacal spur single; one pair of dark, roundish or somewhat elongated spots in occipital region; smaller, but more distinct dark spot in nuchal region; light broken mid-dorsal stripe evident on body (Ota et al. 1995).

The discoveries of Gekko paucituberculatus sp. nov. and G. palmatus bring the number of known species in the subgenus G. (Japonigekko) in China to 19, with seven in Guangxi, the provincial district with the highest diversity. Four of them are karst dwellers: Gekko adleri, G. kwangsiensis, G. liboensis and G. paucituberculatus.

The multiple landforms (e.g. peak cluster, depression, cave and peak forest) in the karst region have led to the formation of fragmented and unique microhabitats, resulting in a comparatively significant biodiversity and the presence of numerous endemic species (Clements et al. 2006; Grismer et al. 2021). The globally-concentrated karst is primarily found in south-central Europe, eastern North America and southern China (Li and Xiong 2021). Karstic landscapes from southern China are considered one of the most representative regions of tropical-subtropical karst formation (Xiong et al. 2008). In the last decade, researchers have discovered several narrow-ranged species of amphibian and reptile that are specifically adapted to the karst ecosystems in this region (Mo et al. 2015; Qi et al. 2020; Luo et al. 2021; Agung et al. 2022; Lin et al. 2022). However, these studies have been limited to only very recent and unconnected survey areas, leaving much of the region unexplored. Additionally, the slow soil formation rate of limestone in karst creates a thin soil layer that is prone to soil erosion and rocky desertification (Yang 1990). These features render the karst ecosystem highly vulnerable and challenging to restore after damage. Consequently, conducting a thorough and comprehensive survey of amphibians and reptiles in this expansive karst region is imperative to assess their diversity and the hazards they encounter. This will enable the development of effective conservation strategies to prevent further loss of unidentified species.

Acknowledgements

We are very grateful to Shuo-Rong He for providing literature on karst landscapes; to Han-Ming Song for his help to the photos; to Uwe Fritz and three reviewers for their constructive comments on this manuscript. This work was supported by DFGP Project of Fauna of Guangdong-202115.

References

  • Agung AP, Chornelia A, Grismer LL, Grismer JL, Quah ESH, Lu JM, Tomlinson KW, Hughes AC (2022) Description of two new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from karst landscapes in Yunnan, China, highlights complex conservation needs. Zoological Research 43: 767–786. https://doi.org/10.24272/j.issn.2095-8137.2022.105
  • Cai B, Ji X, Wang Y, Rao D, Huang S, Wang Y, Song Z, Jiang J (2022) An annotated list of lizards (Sauria: Squamata) recorded from the People’s Republic of China. Asian Herpetological Research 13: 64–74. https://10.16373/j.cnki.ahr.210020
  • Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: More models, new heuristics and parallel computing. Nature Methods 9: 772–772. https://doi.org/10.1038/nmeth.2109
  • Grismer LL, Pinto LD, Quah ESH, Anuar S, Cota M, McGuire JA, Iskandar DT, Wood Jr PL, Grismer JL (2022) Phylogenetic and multivariate analyses of Gekko smithii Gray, 1842 recover a new species from Peninsular Malaysia and support the resurrection of G. albomaculatus (Giebel, 1861) from Sumatra. Vertebrate Zoology 72: 47–80. https://doi.org/10.3897/vz.72.e77702
  • 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: 1–15. https://doi.org/10.3390/d13050183
  • 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: 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
  • 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
  • Li L, Xiong KN (2021) Study on peak cluster-depression rocky desertification landscape evolution and human activity-influence in South of China. European Journal of Remote Sensing 54: 309–317. https://doi.org/10.1080/22797254.2020.1777588
  • Lin SS, Li YH, Su HL, Yi H, Pan Z, Sun YJ, Zeng ZC, Wang J (2022) Discovery of a new limestone karst-restricted odorous frog from northern Guangdong, China (Anura, Ranidae, Odorrana). ZooKeys 1120: 47–66. https://doi.org/10.3897/zookeys.1120.87067
  • 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 [in Chinese]. https://doi.org/10.6532/JNTM.2016.69(2).01
  • Luo T, Wang S, Xiao N, Wang Y, Zhou J (2021) A new species of odorous frog genus Odorrana (Anura, Ranidae) from southern Guizhou Province, China. Asian Herpetological Research 12: 381–398.
  • 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: 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: 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: 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: 236–258. https://doi.org/10.11646/zootaxa.4951.2.2
  • 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: 501–518. https://doi.org/10.11646/zootaxa.3652.5.1
  • Nguyen SV, Ho CT, Nguyen TQ (2009) Herpetofauna of Vietnam. Edition Chimaira, Frankfurt am Main, 687 pp.
  • Qi S, Wang J, Grismer LL, Chen H, Lyu Z, Wang Y (2020) The stoor hobbit of Guangdong: Goniurosaurus gollum sp. nov., a cave-dwelling Leopard Gecko (Squamata, Eublepharidae) from South China. ZooKeys 991: 137–153. https://doi.org/10.3897/zookeys.991.54935
  • 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: 181–196. https://doi.org/10.1080/03946975.1995.10539278
  • Poyarkov NA, Nguyen TV, Popov ES, Geissler P, Pawangkhanant P, Neang T, Suwannapoom C, Ananjeva NB, Orlov NL (2023) Recent progress in taxonomic studies, biogeographic analysis, and revised checklist of reptiles in Indochina. Russian Journal of Herpetology 30: 255–476. https://doi.org/10.30906/1026-2296-2023-30-5-255-476
  • 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: 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: 1–50. 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: 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: 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].
  • 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: 452–466. https://doi.org/10.1643/CH-06-281
  • Wang K, Ren JL, Chen HM, Lyu ZT, Guo XG, Jiang K, Chen JM, Li JT, Guo P, Wang YY, Che J (2020) The updated checklists of amphibians and reptiles of China. Biodiversity Science 28: 189–218 [in Chinese].
  • Wood 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
  • Xiong KN, Xiao SZ, Liu ZQ, Chen PD (2008) Comparative analysis on world natural heritage value of south China karst. Engineering Sciences 10: 17–28 [in Chinese].
  • Yang MD (1990) On the fragility of karst environment. Yunnan Geographic Environment Research 2: 21–29 [in Chinese].
  • 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: 3796. https://doi.org/10.3390/ani13243796
  • 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].
  • Zhou KY, Li H, Han DM, Bauer AM, Feng J (2006) The complete mitochondrial genome of Gekko gecko (Reptilia: Gekkonidae) and support for the monophyly of Sauria including Amphisbaenia. Molecular Phylogenetics and Evolution 40: 887–892. https://doi.org/10.1016/j.ympev.2006.04.003
login to comment