A new rock-dwelling gecko of the subgenus Japonigekko (Squamata, Gekkonidae, Gekko) from northwestern Sichuan Province, China

A new rock-dwelling gecko of the subgenus Japonigekko (Squamata, Gekkonidae, Gekko) from northwestern Sichuan Province, China Yuhao Xu1, Shun Ma2,3, Bo Cai2, Shuo Qi4, Tomoya Matsukoji5, Nikolay A. Poyarkov6, Fanyue Sun1, Shiyang Weng7, Tianxuan Gu8, Kaidi Tian9, Di Zhang9, Jianping Jiang2, Lifang Peng1 1 State Key Laboratory of Plateau Ecology and Agriculture, Qinghai University, Xining 810016, Qinghai, China 2 China-Croatia Belt and Road Joint Laboratory on Biodiversity and Ecosystem Services, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610213, Sichuan, China 3 University of Chinese Academy of Science, Beijing 100049, China 4 School of Life Sciences / School of Ecology, Sun Yat-sen University, Guangzhou 510275, Guangdong, China 5 Graduate School of Science and Life Technology, University of Tsukuba, Ibaraki 305-8572, Japan 6 Department of Vertebrate Zoology, Lomonosov Moscow State University, Leninskiye Gory, GSP–1, Moscow 119234, Russia 7 Institute of Plateau Biology of Xizang Autonomous Region, Lhasa 850008, Xizang, China 8 Longfor U City Community, Chongqing 401331, China 9 Yellow River Water Resources Protection Institute, Zhengzhou 450004, Henan, China https://zoobank.org/CCCD4D14-5256-40AC-B382-A16E665FD39A Corresponding authors: Lifang Peng ([email protected]); Jianping Jiang ([email protected]) Academic editor: Justin Bernstein ♦ Received 17 October 2025 ♦ Accepted 3 November 2025 ♦ Published 18 November 2025 Abstract We describe a new rock-dwelling species of the genus Gekko (subgenus Japonigekko) from Heishui County, Aba Tibetan and Qiang Autonomous Prefecture, Sichuan Province, China, based on both morphological and molecular (1,574 bp from the mitochondrial 16S rRNA and ND2 genes) evidence. Morphologically, Gekko tesselatus sp. nov. is characterized by its moderate body size; two (rarely one) enlarged postmentals; flattened dorsal tubercles extending from the posterior head through the neck to the anterior portion of the tail, arranged in 12–15 rows at midbody; 130–157 ventral scales between the mental and the cloacal slit; 98–106 midbody scale rows; 31–39 ventral scale rows; subdigital lamellae numbering 7–10 on finger I, 9–12 on finger IV, 7–9 on toe I, and 9–12 on toe IV; absence of webbing; 6–8 precloacal pores in males and absence in females; one (rarely two) postcloacal tubercle on each side; and a distinctive dorsal coloration. Phylogenetically, the new species forms a distinct clade within the subgenus Japonigekko, showing uncorrected 16S sequence divergences of at least 8.2% from its closest relative, G. liboensis, and at least 8.9% from all other congeners, as well as ND2 divergences of at least 14.3% from G. fengshanensis and at least 14.9% from other species in the subgenus. The discovery of this new species raises the number of Japonigekko species recorded in China to 26 and in Sichuan Province to seven. Key Words Gekko tesselatus sp. nov., molecular phylogeny, morphological characters, taxonomy Introduction The genus Gekko Laurenti, 1768, represents a taxonomically diverse group within the family Gekkonidae Gray, 1825. Currently, 93 valid species are recognized within the genus, and its members are widely distributed across eastern and southeastern Asia, as well as parts of northwestern Oceania and Melanesia (Ma et al. 2024; Cao et al. 2025; Huang et al. 2025; Pauwels et al. 2025; Uetz et al. 2025). Due to the high morphological similarity among species, which complicates diagnosis and hampers the identification of cryptic lineages, the taxonomy of the genus Gekko remained unreZoosyst. Evol. 101 (4) 2025, 2221–2242|DOI 10.3897/zse.101.175246 Copyright Xu, Y. et al. 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. zse.pensoft.net Xu, Y. et al.: A new Gekko species 2222 solved for long period of time (Zhao and Adler 1993; Zhao et al. 1999; Kluge 1993, 2001; Rösler et al. 2005, 2011). In recent years, however, the use of integrative taxonomic approaches that combine detailed morphological assessments with molecular phylogenetic analyses has clarified many previously ambiguous species boundaries and gradually revealed the true extent of diversity within the genus (Brown et al. 2009; Rösler et al. 2011; Luu et al. 2015; Wood et al. 2020; Lyu et al. 2021; Xia et al. 2023; Huang et al. 2025). According to the classification proposed by Wood et al. (2020), the genus is currently divided into seven subgenera: Gekko s. str., Japonigekko Wood, Guo, Travers, Su, Olson, Bauer, Grismer, Siler, Moyle, Andersen & Brown, Ptychozoon Kuhl & Van Hasselt, Rhacogekko Wood, Guo, Travers, Su, Olson, Bauer, Grismer, Siler, Moyle, Andersen & Brown, Lomatodactylus van der Hoeven, Balawangekko Wood, Guo, Travers, Su, Olson, Bauer, Grismer, Siler, Moyle, Andersen & Brown, and Archipelagekko Wood, Guo, Travers, Su, Olson, Bauer, Grismer, Siler, Moyle, Andersen & Brown. Among these, Japonigekko is the most diverse group, comprising 38 species, 25 of which occur in China (Cao et al. 2025; Huang et al. 2025; Zhou et al. 2025b). Members of Japonigekko are characterized by the following diagnostic features: moderate body size; nares usually in contact with the rostral; 2–4 nasals (including nasorostrals, supranasals, and postnasals); 0–21 rows of dorsal tubercles; 0–32 precloacal pores; 1–4 postcloacal tubercles; and lateral folds lacking tubercles (Wood et al. 2020; Hou et al. 2021; Ma et al. 2024; Wang et al. 2024; Zhou et al. 2025a; Uetz et al. 2025). Located in western China, Sichuan Province encompasses several major geomorphological regions, including the Qinghai–Xizang Plateau, the Hengduan Mountains, the Yunnan–Guizhou Plateau, the Qinba Mountains, and the Sichuan Basin. Its complex geological structure and highly diverse topography have given rise to a complex and varied climate. Together with significant geographical barriers and a mosaic of microhabitats, these conditions have driven pronounced regional differentiation of species. Consequently, Sichuan supports exceptionally high levels of biodiversity and is recognized as one of China’s most important reservoirs of genetic resources. During a field survey conducted in the summer of 2025, we collected a series of Gekko (Japonigekko) specimens from a dry-hot valley in Heishui County, Aba Tibetan and Qiang Autonomous Prefecture, Sichuan Province, China (Fig. 1). However, detailed morphological comparisons and mitochondrial DNA analyses revealed that these individuals differ markedly from all known Gekko species. Based on this evidence, we herein describe this population as a new species. Material and methods Sampling Field surveys were conducted in the dry-hot valley of Heishui County, Sichuan Province, China, in the summer of 2025. Geographic coordinates and elevation were recorded using the TwoStep Outdoor Assistant v7.9.13 (Shenzhen 2bulu Information Technology Co., Ltd., Shenzhen, China). After being photographed, all newly collected specimens were humanely euthanized using a lethal injection of 0.7% tricaine methanesulfonate (MS222) solution, then fixed in 95% ethanol for one day and transferred to 75% ethanol for long-term preservation. Fresh liver tissue was extracted, immediately preserved in 95% ethanol, and stored at –20 °C. The specimens were deposited at Qinghai University (QHU), the Museum of Biology, Sun Yat-sen University (SYS), and the Chengdu Institute of Biology (CIB), Chinese Academy of Sciences. All sampling procedures involving live geckos complied with the Wildlife Protection Law of China and were approved by the Institutional Ethics Committee of Qinghai University (Protocol No. PJ202501-89). Molecular phylogeny Since all specimens were collected from the same locality and showed consistent morphological traits, we randomly selected 10 individuals for DNA sequencing. Total genomic DNA was extracted from preserved liver tissue using the QIAamp DNA Mini Kit (QIAGEN, Changsheng Biotechnology Co., Ltd.). Two mitochondrial gene fragments, 16S ribosomal RNA (16S) and NADH dehydrogenase subunit 2 (ND2), were amplified via polymerase chain reaction (PCR) following the protocols outlined by Wang et al. (2024). The following primer pairs were used: L3975 (5’-CGCCTGTTTACCAAAAACAT-3’) and H4551 (5’-CCGGTCTGAACTCAGATCACGT-3’) for 16S (Simon et al. 1994), and rMet-3L (5’-ATACCCCGACAATGTTGG-3’) and rAla-1H (5’-GCCTTAGCTTAATTAAAGTG-3’) for ND2 (Jonniaux and Kumazawa 2008). The PCR products were sequenced by Shanghai Map Biotech Co., Ltd. Raw sequences were assembled using SeqMan in the DNASTAR software package (Burland 2000). Newly generated sequences have been submitted to the DDBJ; accession numbers are listed in Table 1. Apart from the 20 newly obtained sequences, we included 80 additional sequences representing 48 individuals from 28 species of the subgenus Japonigekko (Table 1). Gekko gecko (Linnaeus, 1758) and G. reevesii (Gray, 1831) were selected as outgroups, following the approach of Wang et al. (2024). A concatenated dataset of the two gene fragments (16S + ND2, 1,574 bp in total) was aligned using MUSCLE (Edgar 2004) in MEGA X (Kumar et al. 2018). Uncorrected pairwise distances (p-distances) between closely related species were also calculated in MEGA X. The Maximum Likelihood (ML) analysis was conducted in IQ-TREE v1.6.12 (Nguyen et al. 2015), partitioned by gene, with both 16S and ND2 fragments independently selected under the bestfit model GTR+F+I+G4 by ModelFinder (Kalyaanamoorthy et al. 2017) implemented in PhyloSuite v1.2.3 (Zhang et al. 2020), according to the Bayesian Information Criterion (BIC) (Kalyaanamoorthy et al. 2017). Zoosyst. Evol. 101 (4) 2025, 2221–2242 zse.pensoft.net 2223 Table 1. NCBI/DDBJ accession numbers, localities, and voucher information for all specimens used in this study. * indicates DDBJ accession number. No. Species Localities Voucher ID. 16S ND2 Reference 1G. tesselatus sp. nov. Heishui, Sichuan, China CIB 119371 LC899308*LC899318* This study 2G. tesselatus sp. nov. Heishui, Sichuan, China CIB 119372 LC899309*LC899319* This study 3G. tesselatus sp. nov. Heishui, Sichuan, China CIB 119373 LC899310*LC899320* This study 4G. tesselatus sp. nov. Heishui, Sichuan, China CIB 119374 LC899311*LC899321* This study 5G. tesselatus sp. nov. Heishui, Sichuan, China CIB 119375 LC899312*LC899322* This study 6G. tesselatus sp. nov. Heishui, Sichuan, China QHU R2025022 LC899313*LC899323* This study 7G. tesselatus sp. nov. Heishui, Sichuan, China QHU R2025023 LC899314*LC899324* This study 8G. tesselatus sp. nov. Heishui, Sichuan, China QHU R2025024 LC899315*LC899325* This study 9G. tesselatus sp. nov. Heishui, Sichuan, China QHU R2025025 LC899316*LC899326* This study 10 G. tesselatus sp. nov. Heishui, Sichuan, China QHU R2025026 LC899317*LC899327* This study 11 G. adleri Jingxi, Guangxi, China SYS r001400 MW451654 OR902178 Lyu et al. (2021); Wang et al. (2024) 12 G. adleri Cao Bang, Vietam IEBR A.2012.24 KC700623 – Nguyen et al. (2013) 13 G. alpinus Batang, Sichuan, China CIB 121663 PQ255983 PQ303501 Ma et al. (2024) 14 G. alpinus Mangkang, Xizang, China CIB 121656 PQ255976 PQ303494 Ma et al. (2024) 15 G. auriverrucosus Yuncheng, Shanxi, China NNU Z20050801.004 – JN019062 Rösler et al. (2011) 16 G. bonkowskii Khammouane, Laos VFU R.2014.10 – KT266818 Luu et al. 2015 17 G. chinensis Hong Kong, China SYS r001211 MW451644 OR902183 Lyu et al. (2021); Wang et al. (2024) 18 G. chinensis Shenzhen, Guangdong, China SYS r001085 MW451632 OR902184 Lyu et al. (2021); Wang et al. (2024) 19 G. cib Hejiang, Sichuan, China SYS r001489 MW451655 OR902165 Lyu et al. (2021); Wang et al. (2024) 20 G. cib Emeishan, Sichuan, China SYS r000708 MW451629 OR902166 Lyu et al. (2021); Wang et al. (2024) 21 G. fengshanensis Fengshan, Guangxi, China NHMG 240713 PV652773 PV657377 Huang et al. (2025) 22 G. fengshanensis Fengshan, Guangxi, China NHMG 240714 PV652774 PV657378 Huang et al. (2025) 23 G. hokouensis Wuyishan, Fujian, China SYS r001290 MW451647 OR902173 Lyu et al. (2021); Wang et al. (2024) 24 G. hokouensis Nanchang, Jiangxi, China SYS r001311 MW451648 OR902172 Lyu et al. (2021); Wang et al. (2024) 25 G. ichangensis Yichang, Hubei, China SWU 0011602 PQ587503 – Cao et al. (2025) 26 G. ichangensis Yichang, Hubei, China SWU 0011603 PQ587504 – Cao et al. (2025) 27 G. japonicus Wuyishan, Fujian, China SYS r000672 MW451628 OR902176 Lyu et al. (2021); Wang et al. (2024) 28 G. japonicus Lushan, Jiangxi, China SYS r001317 MW451649 OR902177 Lyu et al. (2021); Wang et al. (2024) 29 G. jinjiangensis Deqin, Yunan, China CIB 5334220088 PQ255987 MT449432 Hou et al. (2021); Ma et al. (2024) 30 G. jinjiangensis Derong, Sichuan, China CIB 5133380017 – MT449437 Hou et al. (2021) 31 G. kaiyai Xinxian, Henan, China AHUXXBH01 OQ780318 – Zhang et al. (2023) 32 G. kaiyai Shangcheng, Henan, China AHUJGTBH01 OQ780322 – Zhang et al. (2023) 33 G. khunkhamensis Khammouane, Laos VNUF R.2021.23 – OL416111 Sitthivong et al. (2021) 34 G. kwangsiensis Wuming, Guangxi, China SYS r001195 MW451642 OR902175 Lyu et al. (2021); Wang et al. (2024) 35 G. kwangsiensis Wuming, Guangxi, China SYS r001194 MW451641 OR902174 Lyu et al. (2021); Wang et al. (2024) 36 G. liboensis Libo, Guizhou, China SYS r002876 PV652777 PV657381 Huang et al. (2025) 37 G. liboensis Libo, Guizhou, China SYS r002877 PV652778 PV657382 Huang et al. (2025) 38 G. melli Dongguan, Guangdong, China SYS r001742 MW451661 OR902169 Lyu et al. (2021); Wang et al. (2024) 39 G. melli Puning, Guangdong, China SYS r001702 MW451660 OR902170 Lyu et al. (2021); Wang et al. (2024) 40 G. nadenensis Khammouane, Laos ZFMK 98741 – KY421618 Luu et al. (2017) 41 G. palmatus Zhaoqing, Guangdong, China SYS r002797 OR903156 OR902179 Wang et al. (2024) 42 G. palmatus Napo, Guangxi, China SYS r001185 MW451637 OR902182 Lyu et al. (2021); Wang et al. (2024) 43 G. paucituberculatus Baise, Guangxi, China SYS r002806 OR903154 OR902163 Wang et al. (2024) 44 G. paucituberculatus Baise, Guangxi, China SYS r002806 OR903155 OR902164 Wang et al. (2024) 45 G. prep Changjiang, Hainan, China HN2024R038 PV368865 PV389916 Zhou et al. (2025a) zse.pensoft.net Xu, Y. et al.: A new Gekko species 2224 No. Species Localities Voucher ID. 16S ND2 Reference 46 G. prep Changjiang, Hainan, China HN2024R039 PV368866 PV389915 Zhou et al. (2025a) 47 G. scientiadventura Quang Binh, Vietnam IEBR A.2014.7 – KP205392 Luu et al. (2014) 48 G. sengchanthavongi Khammouane, Laos VFU R2014.14 – KT266816 Luu et al. (2015) 49 G. similignum Wuzhishan, Hainan, China SYS r001597 MW451658 OR902185 Lyu et al. (2021); Wang et al. (2024) 50 G. similignum Wuzhishan, Hainan, China SYS r001598 MW451659 OR902186 Lyu et al. (2021); Wang et al. (2024) 51 G. scabridus Yanbian, Sichuan, China CIB YN201909199 PQ255992 MT449429 Hou et al. (2021); Ma et al. (2024) 52 G. scabridus Yanbian, Sichuan, China CIB YN201909200 – MT449430 Hou et al. (2021) 53 G. subpalmatus Fenghua, Zhejiang, China SYS r001762 MW451662 OR902167 Lyu et al. (2021); Wang et al. (2024) 54 G. subpalmatus Fenghua, Zhejiang, China SYS r001767 MW451663 OR902168 Lyu et al. (2021); Wang et al. (2024) 55 G. swinhonis Zunhua, Hebei, China SYS r001814 MW451666 OR902171 Lyu et al. (2021); Wang et al. (2024) 56 G. swinhonis Zunhua, Hebei, China SYS r001815 MW451667 – Lyu et al. (2021) 57 G. thakhekensis Thakhek, Khammouane, Laos IEBR A.2014.6 – KP205396 Luu et al. (2014) 58 G. truongi Khanh Hoa, Vietnam IEBR A.2011.1 – KP205398 Luu et al. (2014) Out group 59 G. gecko Nanning, Guangxi, China N/A AY282753 AY282753 Zhou et al. (2006) 60 G. reevesii Mt. Yinping, Guangdong, China SYS r000796 MW451630 OR902187 Lyu et al. (2021); Wang et al. (2024) Nodes were regarded as well supported when ultrafast bootstrap values (UFB) ≥ 95% and SH-like approximate likelihood ratio test values (SH-aLRT) ≥ 80% (Stephane et al. 2010; Hoang et al. 2018). The phylogenetic tree was visualized using FigTree v1.4.4 (Rambaut 2018). Morphological examination Comparisons with other congeners of the subgenus Japonigekko were based on available literature: Stejneger (1907); Zhou et al. (1982); Song (1985); Ota et al. (1995); Zhao et al. (1999); Goris and Maeda (2004); Rösler et al. (2005, 2011); Toda et al. (2008); Zhou and Wang (2008); Phung and Ziegler (2011); Nguyen et al. (2013); Luu et al. (2014, 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); Ma et al. (2024); Wang et al. (2024); Cao et al. (2025); Huang et al. (2025); Zhou et al. (2025a, b). Morphological descriptions followed the methodology outlined in Lyu et al. (2021) and Grismer et al. (2022), with certain abbreviations revised according to the standards set following the abbreviations list in Darko et al. (2022). Three morphometric characters were measured with Deli digital calipers (No. 90150B) to the nearest 0.1 mm: SVL = snout– vent length, measured from the tip of the snout to the posterior edge of the vent; TAL = tail length, measured from the posterior margin of the vent to the tip of the tail; AGD = axilla–groin distance, defined as the distance between the posterior edge of the forelimb insertion and the anterior edge of the hindlimb insertion, with both limbs inserted perpendicularly to the body wall. All other measurements were taken using Mitutoyo digital calipers (CD-15AX) to the nearest 0.01 mm under a Leica stereomicroscope (EZ4): HL = head length, measured from the tip of the snout to the posterior margin of the ear opening; HW = head width, measured at the angle of the jaws; HH = head height, measured from the top of the head posterior to the eyes to the bottom of the lower jaw; ESD = eye-to-snout distance/snouth length, measured from the tip of the snout to the anterior corner of the eye; ED = eye diameter, measured as the greatest distance from the anterior to the posterior corner of the eye; EL = maximum ear opening diameter; RW = maximum rostral width; RH = maximum rostral height; MW = maximum mental width; and ML = maximum mental length. Scalation features and their abbreviations were as follows: NS = number of nasals, including nasorostrals, supranasals, and postnasals; INS = internasals, number of scales between supranasals, in contact with the rostral; SL = supralabials; IL = infralabials; IOS = interorbitals, number of scales in a line between the anterior corners of the eyes; PO = preorbitals, number of scales in a line from the nostril to the anterior margin of the eye; PM = postmentals, number of scales bordering the mental; GP = gulars bordering the postmentals; MBSR = midbody scale rows; VS = ventral scale rows at midbody; SMC = scales in a line from the mental to the front of the cloacal slit; DTR = dorsal tubercle rows at midbody; GSDT = granules surrounding dorsal tubercles, counted by randomly selecting 10 tubercles on the trunk and recording the number of granules surrounding each; LF1 = number of the subdigital lamellae beneath the entire first finger; LF4 = number of the subdigital lamellae beneath the entire fourth finger; LT1 = number of the subdigital lamellae beneath the entire first toe; LT4 = number of the subdigital lamellae beneath the entire fourth toe; PP = number of the precloacal pores; PAT = number of the postcloacal tubercles. Zoosyst. Evol. 101 (4) 2025, 2221–2242 zse.pensoft.net 2225 Figure 1. Map showing the locality of Gekko tesselatus sp. nov. The star indicates the type locality, and circles denote other known distribution sites. In addition, we recorded the presence or absence of webbing (Web); the presence of tubercles on the forelimbs (Fore tubercles), hindlimbs (Hind tubercles), and tail (Tail tubercles); the presence of enlarged subcaudal scales; and the types of dorsal coloration patterns. Other abbreviations are as follows: Is.: Island; Mt.: Mountain; Mts.: Mountains; NP: National Park; NR: Nature Reserve; WS: Wildlife Sanctuary; asl.: above sea level. Results The lengths of the 16S and ND2 fragments in the final alignment comprised 563 base pairs (bp) and 1,011 bp, respectively. In the ML analysis (Fig. 2), all Gekko (Japonigekko) samples included in this study formed a strongly supported monophyletic lineage (SH = 100 / UFB = 100). However, relationships among species within the subgenus remain incompletely resolved due to low nodal support at interspecific levels. The newly collected specimens from Heishui County, Sichuan Province, China, constituted a well-supported monophyletic lineage within Japonigekko (SH = 100 / UFB = 100), which was recovered as sister to a clade comprising G. fengshanensis Huang, Wang, Qi, Song, Huang, Wang & Mo, 2025, G. liboensis Zhou, Liu & Li, 1982, and G. kwangsiensis Yang, 2015, although with relatively low support (SH = 80 / UFB = 65). Finally, these four taxa, along with G. paucituberculatus Wang, Qi, Zhou & Wang, 2024, formed a well-supported clade (SH = 100 / UFB = 100), whose members exhibit specialized ecological adaptations to rocky environments. Uncorrected p-distances for the 16S and ND2 gene fragments are summarized in Tables 2 and 3, respectively. Interspecific genetic distances within Japonigekko ranged from 3.5% between G. chinensis (Gray, 1842) and G. similignum Smith, 1923 to 16.8% between G. adleri Nguyen, Wang, Yang, Lehmann, Le, Ziegler & Bonkowski, 2013 and G. swinhonis Günther, 1864 for the 16S gene (Table 2); and from 5.4% between G. chinensis and G. similignum to 26.7% between G. prep Zhou, Chen, Liu, Li, Qi, Li & Peng, 2025 and G. swinhonis Günther, 1864 for the ND2 gene (Table 3). The Heishui population displayed substantial genetic divergence from all congeners, ranging from 8.2% (vs. G. liboensis) to 16.0% (vs. G. swinhonis) for the 16S gene and from 14.3% (vs. G. fengshanensis) to 25.2% (vs. G. melli (Vogt, 1922)) for the ND2 gene. In light of the well-supported monophyly, substantial genetic divergence, and distinct morphological characters that consistently differentiate the Gekko population from Sichuan Province from all known congeners (see Comparisons below), we herein describe the Gekko sp. from Heishui County, Sichuan Province, China, as a new species. zse.pensoft.net Xu, Y. et al.: A new Gekko species 2226 Table 2. Uncorrected p-distances (%) of the 16S gene among species of Japonigekko used in this study. NO Species 1 2 3 4 5 6 7 8 9 10 11 1G. tesselatus sp. nov. 0.0–0.8 2G. adleri 13.6–14.2 0.7 3G. alpinus 10.7–11.4 13.8–14.8 0.5 4G. chinensis 12.8–13.5 5.6–5.8 11.8–12.6 0.2 5G. cib 11.9–12.6 14.9 9.4–9.5 13.5–13.7 0.0 6G. fengshanensis 9.1–9.7 14.6–15.5 11.2–11.9 14.6–14.9 13.7–14.0 0.6 7G. hokouensis 11.3–11.9 14.2–14.4 12.0–13.1 12.8–13.2 13.2–13.4 12.6–13.1 0.4 8G. ichangensis 12.1–13.0 15.4 9.6–10.5 15.1–15.3 12.4–12.5 14.8–15.0 14.0–14.3 0.0 9G. japonicus 13.4–13.8 15.1–15.6 10.5–11.4 14.4–14.8 13.6–14.0 14.3–14.6 13.7–14.1 7.7–7.8 0.2 10 G. jinjiangensis 9.5–9.8 13.0–13.4 4.0–4.4 11.3–11.5 10.2 12.0–12.2 10.0–10.2 9.9–10.0 10.8 – 11 G. kaiyai 12.7–13.6 14.1–15.4 12.4–13.5 13.1–14.2 12.6–13.6 13.9–14.0 8.8–9.5 13.7–14.9 14.2–15.2 10.9–11.2 1.2 12 G. kwangsiensis 10.2–10.6 14.7 11.5–12.5 13.4–13.7 14.9–15.1 10.3–10.7 13.2–13.7 15.3–15.5 15.3–15.6 11.4–11.5 13.2–13.5 13 G. liboensis 8.2–9.2 14.2–15.2 11.3–12.4 13.8–14.2 13.3–13.5 9.0–9.4 14.6–14.7 14.4–14.6 13.3–13.7 12.4–12.6 14.3–15.2 14 G. melli 11.0–11.8 13.6–14.2 8.6–8.9 13.1–13.5 7.8–8.0 12.6–13.3 13.4–13.6 12.6 14.5–14.7 9.6–9.8 13.4–14.0 15 G. palmatus 13.1–13.5 2.5–2.9 13.2–14.0 3.8–4.4 13.7–14.1 15.3–15.6 12.6–13.2 15.4–15.8 13.7–14.1 12.3–12.7 13.6–14.0 16 G. paucituberculatus 8.9–9.3 14.0 10.5–11.2 12.5–12.7 12.9 10.7–10.9 11.7 16.0–16.1 15.4–15.5 10.2 12.9 17 G. prep 13.0–13.8 6.0–6.8 12.0–12.4 3.5–3.7 15.0–15.1 15.6–15.7 14.6–14.8 15.3–15.5 14.6–14.8 11.9–12.0 13.1–13.8 18 G. similignum 12.6–13.2 6.4 12.2–13.0 1.9–2.0 15.2 15.4–15.5 13.4–13.6 16.0–16.1 14.8–15.0 11.3 13.9–14.8 19 G. scabridus 11.0–11.2 13.5–13.9 4.7–4.9 12.2–12.4 9.7 11.5 11.1–11.3 11.5 11.1 5.4 11.2–11.5 20 G. subpalmatus 14.3–14.9 16.0 9.8–9.9 15.0–15.1 9.6 13.9–14.4 14.9–15.3 14.0 14.8–15.0 12.0 14.6–15.0 21 G. swinhonis 15.8–16.0 16.2–16.8 13.0–14.4 15.0–15.4 12.4–13.0 16.1–16.8 14.3–15.1 14.9–15.5 15.7–16.1 14.1–14.5 15.4–16.4 NO. Species 12 13 14 15 16 17 18 19 20 21 12 G. kwangsiensis 0.4 13 G. liboensis 11.9–12.1 0.2 14 G. melli 14.4–14.8 13.5–13.9 0.9 15 G. palmatus 13.5–13.7 14.1–14.3 12.4–13.0 0.4 16 G. paucituberculatus 12.0–12.3 9.6–9.8 13.1–13.3 12.0–12.2 0.0 17 G. prep 13.6–13.8 15.0–15.2 13.7–13.9 4.2–4.6 14.0 0.2 18 G. similignum 14.2–14.3 14.2–14.4 13.7–14.3 4.6–5.0 14.0 4.3 0.0 19 G. scabridus 12.1–12.4 12.2 10.0–10.2 12.6–13.1 10.6 12.6–12.7 12.4 – 20 G. subpalmatus 13.4–13.6 15.5–15.7 8.8–9.0 14.1–14.5 15.1 14.8 15.7 12.0 – 21 G. swinhonis 16.0–16.3 15.1–15.5 14.3–14.8 14.6–15.0 15.8–16.2 15.4–16.0 15.4–15.9 14.9–15.1 13.9–14.4 1.1 Zoosyst. Evol. 101 (4) 2025, 2221–2242 zse.pensoft.net 2227 Table 3. Uncorrected p-distances (%) of the ND2 gene among species of Japonigekko used in this study. NO. Species 1 2 3 4 5 6 7 8 9 10 11 12 13 1G. tesselatus sp. nov. 0.0–1.0 2G. adleri 23.8–25.0 – 3G. alpinus 19.6–20.9 23.4–23.6 1.6 4G. auriverrucosus 19.6–20.5 24.8 21.5–22.2 – 5G. bonkowskii 20.3–20.7 24.6 19.0–19.4 21.5 – 6G. chinensis 22.5–24.5 14.4–14.6 21.9–22.7 24.3–24.5 21.2–21.4 0.4 7G. cib 22.1–22.7 26.1 22.2–22.8 20.7 19.4 24.0–24.3 0.0 8G. fengshanensis 14.3–14.9 23.4 20.0–20.3 22.1 19.6 23.7 21.7 0.0 9G. hokouensis 19.7–21.2 24.8–25.2 19.9–21.6 21.0–21.5 19.0–20.3 24.2–25.0 23.0–23.6 20.6–20.9 1.3 10 G. japonicus 20.3–21.8 26.1 19.6–20.6 20.9 19.6 24.9–25.2 22.9 21.4 22.5–22.7 0.0 11 G. jinjiangensis 16.9–17.2 20.7–21.3 7.1–8.4 19.9–20.5 17.9–19.0 21.1–22.4 20.6–20.7 17.0–18.3 18.0–20.4 16.9–17.2 2.4 12 G. khunkhamensis 22.4–22.8 26.0 21.3–21.7 24.3 15.2 24.6–24.8 21.1 22.2 23.8–24.2 22.6 20.9–21.7 – 13 G. kwangsiensis 16.6–17.4 24.1–24.2 21.8–22.7 22.4–22.5 20.9–21.0 23.0–23.3 20.7–20.9 15.9–16.1 21.4–21.8 22.7–22.8 19.1–19.4 21.5–21.7 0.4 14 G. liboensis 16.4–16.7 23.4–23.5 20.8–21.7 22.1–22.2 19.4–19.6 24.7–24.9 21.4–21.5 17.0–17.1 19.6–20.2 22.8–22.9 19.3–20.0 21.5–21.7 18.1–18.4 15 G. melli 23.4–25.2 24.5–25.1 21.9–23.0 22.6–23.5 21.2–21.8 24.7–25.2 18.9–19.0 24.1–24.3 23.6–24.6 23.6–23.8 19.3–21.1 23.4 23.3–23.8 16 G. nadenensis 21.6–22.0 23.3 18.6–19.4 20.2 6.9 21.2–21.8 20.3 20.5 21.2–22.2 19.9 16.9–18.1 13.8 20.5–20.7 17 G. palmatus 22.3–24.2 6.5 22.4–22.6 23.6–23.8 23.1–23.6 14.3–14.4 25.9–26.1 23.1–23.2 23.7–24.4 24.9–25.0 20.7–21.1 25.6–26.2 23.4–23.5 18 G. paucituberculatus 14.9–16.6 25.3–25.4 19.4–20.4 21.2–21.3 19.2 25.4–25.5 21.0–21.1 17.4–17.5 21.0–21.8 21.7–21.8 17.2–17.6 21.5 19.0–19.3 19 G. prep 23.1–24.7 15.9–16.0 24.6–25.6 26.1–26.3 21.6–22.2 7.4–8.0 24.8–24.9 25.0 25.6–26.0 26.4–26.7 22.6–23.5 25.4–25.6 24.2–24.4 20 G. scientiadventura 20.1–20.5 24.4 18.4–18.8 20.4 13.8 22.5 20.5 19.6 21.4–21.6 21.0 17.7 14.8 20.9 21 G. sengchanthavongi 19.4 23.8 20.0–20.2 20.0 14.2 22.2–22.7 21.6 20.1 22.0–22.9 20.9 19.0–19.2 15.7 21.4 22 G. similignum 22.5–24.0 15.0–15.1 22.6–23.3 25.3–25.4 22.9–23.1 5.4–5.5 25.3–25.4 24.0–24.1 24.6–25.1 25.0–25.1 21.3–22.2 24.8–25.0 23.5–23.6 23 G. scabridus 17.4–17.8 18.3–19.1 11.5–12.6 19.2–19.7 19.9–20.3 20.2–20.7 21.1–21.7 16.7 17.0–17.4 17.2–17.6 9.6–11.5 21.7–22.6 20.2–20.9 24 G. subpalmatus 21.3–22.4 25.0 21.7–22.2 21.6 20.5 24.1–24.4 17.9 21.7 23.3–23.4 23.1 19.1–20.2 22.6 22.2 25 G. swinhonis 20.2–20.9 25.1 22.6–22.7 20.7 21.2 25.7–25.8 21.9 20.7 22.0–22.5 21.9 20.4 23.6 21.8–21.9 26 G. thakhekensis 20.9–21.2 21.8 19.0–19.6 20.6 6.7 19.9–20.5 20.3 19.4 20.5–20.9 19.0 18.1–19.2 15.9 20.1–20.3 27 G. truongi 22.9 20.5 20.2–20.3 24.3 22.0 20.9–21.2 21.8 23.6 22.3 21.8 19.6–19.9 22.2 22.7–22.9 NO. Species 14 15 16 17 18 19 20 21 22 23 24 25 26 14 G. liboensis 0.1 15 G. melli 23.9–24.5 2.7 16 G. nadenensis 18.8–19.0 20.1–21.4 – 17 G. palmatus 23.2–23.4 25.2–25.9 22.5–23.2 1.2 18 G. paucituberculatus 17.8–18.0 25.0–25.3 18.6 24.4–24.5 0.1 19 G. prep 26.2–26.3 26.4–27.0 22.3–22.9 15.2–15.4 26.4–26.5 0.4 20 G. scientiadventura 20.3–20.5 21.2–21.6 13.4 22.9–23.5 18.6 23.8–24.4 – 21 G. sengchanthavongi 19.7–19.9 21.8–22.5 11.9 23.5 19.0 22.7–23.3 10.4 – 22 G. similignum 24.5–24.7 25.8–26.4 22.5–22.7 13.9–14.5 24.9–25.0 8.5–9.0 22.9–23.1 22.5–22.7 0.1 23 G. scabridus 16.9–18.3 19.3–21.3 19.2–19.6 19.7–20.2 17.8 21.9–22.4 17.1–18.8 18.4–19.2 20.2–21.1 1.5 24 G. subpalmatus 21.4–21.5 17.8–18.4 18.6 25.3–25.5 22.4–22.5 25.5–25.6 20.3 20.5 25.0–25.1 20.0–21.1 0.0 25 G. swinhonis 21.9–22.0 23.1–23.4 21.4 24.0–24.3 23.7–23.8 26.5–26.7 22.5 22.2 25.5–25.6 19.6–20.2 21.8 – 26 G. thakhekensis 18.6–18.8 19.7–20.7 6.7 22.2–22.9 18.1 21.6–22.2 13.0 13.0 20.9–21.0 19.0–19.4 19.9 23.1 – 27 G. truongi 21.8–22.0 20.7–21.4 22.0 21.6–21.8 21.2 22.2–22.7 22.2 21.8 21.6–21.8 18.2–19.7 21.4 24.0 20.3 zse.pensoft.net Xu, Y. et al.: A new Gekko species 2228 Taxonomic account Family Gekkonidae Oppel, 1811 Genus Gekko Laurenti, 1768 Subgenus Japonigekko Wood, Guo, Travers, Su, Olson, Bauer, Grismer, Siler, Moyle, Andersen & Brown, 2020 Gekko tesselatus Xu, Ma, Cai, Qi, Matsukoji, Poyarkov, Sun, Jiang & Peng, sp. nov. https://zoobank.org/6373BF5E-F05B-4248-8B7D-0E50B1E10BE3 Figs 3–10, Tables 4–6 Type material. Holotype • CIB 119371 (Field no. LFR 2025142), adult male, from Se’ergu Town, Heishui County, Aba Tibetan and Qiang Autonomous Prefecture, Sichuan Province, China (31.9531°N, 103.3971°E; 1,847 m asl.), collected by Yuhao Xu, Fanyue Sun, and Jiaxiang Wu on 3 August 2025. Paratypes (N = 13, all from Sichuan Province, China) • QHU R2025022 (Field no. LFR 2025143, adult female) • QHU R2025023 (Field no. LFR 2025144, adult male) • QHU R2025024 (Field no. LFR 2025145, adult female), with the same collecting information as the holotype • QHU R2025025 (Field no. LFR 2025146, adult male), from Longba Town, Heishui County (31.9855°N, 103.3503°E; 2,047 m asl.), collected by Yuhao Xu, Fanyue Sun and Jiaxiang Wu on August 04, 2025 • QHU R2025026 (Field no. LFR 2025147, subadult male), from Se’ergu Town, Heishui County (31.9290°N, 103.4211°E; 1,827 m asl.), collected by Yuhao Xu, Fanyue Sun and Jiaxiang Wu on August 4, 2025 • SYS r003029 (Field no. LFR 2025148, adult male) • SYS r003030 (Field no. LFR 2025149, adult female), from Se’ergu Town, Heishui County (31.9260°N, 103.44324°E; 1,886 m asl.), collected by Yuhao Xu, Fanyue Sun and Jiaxiang Wu on August 4, 2025 • SYS r003031 (Field no. LFR 2025150, adult female) • SYS r003032 (Field no. LFR 2025151, subadult female), from the same locality as the holotype, collected by Yuhao Xu, Fanyue Sun and Jiaxiang Wu on August 4, 2025 • CIB 119372 (adult female) • CIB 119373 (adult female) • CIB 119374 (adult female) • CIB 119375 (subadult female), from Se’ergu Town, Heishui County (31.9650°N, 103.3966°E; 1,938 m asl.), collected by Bo Cai in July, 2025. Etymology. The specific name “tesselatus” is a Latinized adjective in the nominative singular (masculine gender), derived from “tessella” (a small square tile), and means “reticulated” or “checkerboard-like.” It refers to the characteristic dorsal pattern of the new species, which consists of a series of irregular dark transverse bands that are faintly interrupted medially by a narrow vertebral line and, laterally, are partly fragmented and partly interconnected, forming an irregular, checkerboard-like reticulated pattern across the dorsum. For the common names, we suggest “Checkered Gecko” in English and “Bān Wén Bì Hǔ” (斑纹壁虎) in Chinese. Diagnosis. Gekko tesselatus sp. nov. can be diagnosed from other Japonigekko species by the following unique combination of characters: (1) a moderate body size (SVL reaches up to 64.9 mm in males and 72.9 mm in females); (2) nares in contact with the rostral, internasal 0 or 1; (3) two enlarged postmentals, rarely one; (4) flattened dorsal tubercles present from the posterior head through the neck to the anterior portion of the tail, arranged in 12–15 rows CIB 119371 QHU R2025023 QHU R2025025 QHU R2025022 QHU R2025024 QHU R2025026 CIB 119374 CIB 119373 CIB 119375 CIB 119372 G. fengshanensis NHMG 240713 G. fengshanensis NHMG 240714 G. liboensis SYS r002876 G. melli SYS r001742 G. kwangsiensis SYS r001194 G. subpalmatus SYS r001762 G. alpinus CIB 121663 G. paucituberculatus SYS r002806 G. hokouensis SYS r001209 G. kaiyai AHUXXBH01 G. liboensis SYS r002877 G. kwangsiensis SYS r001195 G. paucituberculatus SYS r002807 G. hokouensis SYS r001311 G. kaiyai AHUJGTBH01 G. melli SYS r001702 G. subpalmatus SYS r001767 G. cib SYS r001489 G. swinhonis SYS r001814 G. auriverrucosus NNU Z20050801.004 G. cib SYS r000708 G. swinhonis SYS r001815 G. alpinus CIB 121656 G. jinjiangensis CIB 5334220088 G. jinjiangensis CIB 5133380017 G. japonicus SYS r000672 G. khunkhamensis VNUF R.2021.23 G. nadenensis ZFMK 98741 G. bonkowskii VFU R.2014.10 G. scabridus CIB YN201909199 G. ichangensis SWU 0011602 G. thakhekensis IEBR A.2014.6 G. scabridus CIB YN201909200 G. ichangensis SWU 0011603 G. japonicus SYS r001317 G. scientiadventura IEBR A.2014.7 G. sengchanthavongi VNUF R.2014.14 G. chinensis SYS r001211 G. similignum SYS r001597 G. prep HN2024R038 G. adleri SYS r001400 G. chinensis SYS r001085 G. similignum SYS r001598 G. prep HN2024R039 G. adleri IEBR A.2012.24 G. gecko N/A G. palmatus SYS r002797 G. truongi IEBR A.2011.1 G. reevesii SYS r000796 G. palmatus SYS r001185 100/100 80/69 55/60 77/64 100/100 95/96 98/100 89/98 100/100 –/76 76/74 76/74 83/77 84/82 84/82 98/100 100/100 99/100 89/97 87/90 –/81 100/100 91/97 –/– 100/100 100/100 99/100 100/100 66/84 0.06 Gekko tesselatus sp. nov. Gekko (Japonigekko G. tesselatus sp. nov. ) Gekko (Gekko)100/100 Figure 2. Maximum likelihood (ML) tree of Japonigekko inferred from 16S and ND2 genes. Node support values are presented as SH-like approximate likelihood ratio test (SH)/ultrafast bootstrap approximation (UFB); those lower than 50 are shown as “–”. Photograph on thumbnail by YHX. Zoosyst. Evol. 101 (4) 2025, 2221–2242 zse.pensoft.net 2229 at midbody; (5) 130–157 ventral scales between the mental and the cloacal slit; (6) 98–106 midbody scale rows; (7) 31–39 ventral scale rows; (8) subdigital lamellae 7–10 on finger I, 9–12 on finger IV, 7–9 on toe I, and 9–12 on toe IV; (9) webbing absent; (10) 6–8 precloacal pores in males, absent in females; (11) one postcloacal tubercle on each side, rarely two; (12) in life, the dorsum is predominantly greyish brown, with a series of irregular dark transverse bands that are faintly interrupted medially by a narrow vertebral stripe and laterally fragmented and interconnected, forming an irregular, checkerboard-like reticulated pattern. Description of the holotype. (Fig. 3). An adult male specimen with original tail. Size medium, SVL 61.4 mm; body slender and trunk relatively elongate, AGD 29.0 mm, AGD/SVL ratio 0.47; tail slightly longer than body, distinctly swollen at base, oval in section, TAL 72.3 mm, TAL/SVL ratio 1.19. Head depressed, noticeably longer than wide, and clearly separated from the neck, HL 17.09 mm, HW 12.31 mm, HH 5.00 mm, HH/HL ratio 0.29, HL/HW ratio 1.39. Snout obtuse, round anteriorly, ESD 7.14 mm, ESD/ HL ratio 0.42. Eye large, ED 3.89 mm, ED/HL ratio 0.23; pupil vertical, featuring crenulated edges. Ear opening small, approximately elliptical, obliquely oriented, EL 1.12 mm, EL/ED ratio 0.29. Rostral approximately rectangular in shape, wider than the high, RW 2.31 mm, RH 1.46 mm, RW/RH ratio 1.58. Nares oval, rounded by the rostral, 1st supralabial, one distinctly enlarged supranasal, and three slightly enlarged nasals posteriorly; internasals absent. Preorbitals 18/17, preorbital region deeply concave; interorbital scales between the anterior corners of eyes 24; supralabials 10/10; infralabials 9/9. Mental pentagonal, wider than long, MW 1.80, ML 1.42, MW/ML ratio 1.27; mental narrower than the rostral, RW/MW ratio 1.28; postmentals two, enlarged, twice as long as wide, touching the mental and the first infralabial on both sides and three gular scales posteriorly. Tubercles present on the dorsal surface of the head behind the eyes; granular scales on the anterodorsal region of the head are larger than those on the posterior region. Dorsal scales smooth, round to oval, granular, and juxtaposed; tubercles flattened, irregularly arranged, extending from the posterior part of the head to the anterior one-third of the tail, forming approximately 14 rows at midbody, surrounded by 8–9 dorsal scales. Ventrolateral fold present, without tubercles; ventrals slightly larger than dorsals, smooth, imbricate, and largest in middle of belly; ventral scale rows at midbody 36; scale rows around midbody 106; ventral scales in a row between mental and cloacal slit 138; precloacal scales enlarged, but no enlarged scales on thighs; precloacal pores 6 (Fig. 4A), situated at the distal of each scale, the tips of the two central scales are slightly inclined laterally, separating the precloacal pores at the midline into two groups (3/3); postcloacal tubercle 1/1 (Fig. 5A), large, surrounded by several smaller tubercles delineated by shallow grooves. Dorsal scales of tail flat and smooth, irregular in size, with a few tubercles present on the anterior one-third of the tail; subcaudals small at the base, distinctly enlarged beyond the swollen portion, and arranged in a longitudinal row. Forelimbs and hindlimbs well developed; dorsal surface of forelimbs without tubercles, but dorsal surface of hindlimbs bearing tubercles; no webbing between the fingers and toes; digits moderately dilated; fingers and toes II–IV clawed; claws laterally compressed, extending beyond the terminal lamellae; no webbing between fingers and toes; subdigital lamellae undivided, under manus 7-9-10-11-9 (left) / 7-8-9-11-9 (right), and under pes 8-8-10-11-8 (left) / 8-8-11-11-10 (right); relative length of fingers and toes I < II < V < III < IV. Coloration of the holotype in life. (Fig. 6). In life, the dorsal ground color of the head and body is light greyish-brown, scattered with small dark brown to yellowish-brown spots. A faint W-shaped dark marking extending anteriorly from the occipital region to the posterior corners of the eyes. On the lateral sides of the head, a dark longitudinal stripe running from the posteroventral corner of the eye to the ear opening. Eight irregular dark transverse bands between the nape and the sacrum. The dark bands faintly divided along the midline by a narrow, light-colored vertebral line running intermittently from the nape to the tail tip. Laterally, the bands partly fragmented and interconnected, forming an irregular reticulated pattern reminiscent of a checkerboard. All dark markings and bands consist of a mixture of yellowish-brown and dark brown spots. The dorsal surfaces of the limbs bearing irregular dark transverse or reticulated markings. The tail light greyish-brown at the base, gradually becoming paler posteriorly, and bearing 12 dark transverse bands; the tail tip dark brown. The head and ventral surfaces of the body uniformly creamy white, while the ventral surface of the tail pale yellowish-white. After capture, the overall body color becomes paler, turning yellowish-brown, and the dark markings on the dorsum become blurred. However, after being kept in captivity for several days, the color pattern gradually returns to its original state, even when housed in a white container. Coloration of the holotype in preservation. (Fig. 3). In preservative, the dorsal ground color of the head, body, and limbs turns grey, with the yellowish-brown spots on the dorsum disappearing, while the dark brown markings remained distinct; the ventral surface faded to greyish-white. Variation. The main morphological characters of Gekko tesselatus sp. nov. are summarized in Table 4. The species exhibits no evident sexual dimorphism in morphometric ratios, scale counts, or coloration. The longest known male measures SVL 64.9 mm, with TAL 72.9 mm (QHU R2025025); and the female measures SVL 65.3 mm, with TAL 65.1+ mm (QHU R2025022); TAL/SVL ratio 1.07– 1.23. Head depressed, noticeably longer than wide, and clearly separated from the neck, HL/HW ratio 1.16–1.47. Snout obtuse, round anteriorly, ESD/HL ratio 0.38–0.43. Eye large; pupil vertical, featuring crenulated edges. Ear opening small, approximately elliptical, obliquely oriented, EL/ED ratio 0.19–0.40, and EL/HL ratio 0.04–0.08. zse.pensoft.net Xu, Y. et al.: A new Gekko species 2236 Figure 8. Preserved specimen of the female paratype of Gekko tesselatus sp. nov. (QHU R2025024). A. Dorsal view; B. Ventral view; C. dorsal view of head; D. Lateral view of head; E. Ventral view of head; F. Ventral view of hand; G. Ventral view of foot; H. Dorsal view of midbody. Photographs by YHX. Scale bars: 5 mm. more, it can be distinguished from G. paucituberculatus by (1) MBSR 98–106 (vs. 136–142), (2) VS 31–39 (vs. 42–44), (3) SMC 130–157 (vs. 189–192), (4) DTR 12–15 (vs. 4), (5) PP 6–8 (vs. 12), and (6) IOS 24–28 (vs. 37). The main characteristics that distinguish Gekko tesselatus sp. nov. from other species of the subgenus Japonigekko are summarized in Table 6. By having 6–8 precloacal pores in males, the new species can be easily distinguished from G. aaronbaueri Tri, Thai, Phimvohan, David & Teynié, 2015, G. adleri Nguyen, Wang, Yang, Lehmann, Le, Ziegler & Bonkowski, 2013, G. canhi canhi Rösler, Nguyen, Van Doan, Ho, Nguyen & Ziegler, 2010, G. chinensis, G. jinjiangensis Hou, Shi, Wang, Shu, Zheng, Qi, Liu, Jiang & Xie, 2021, G. kaiyai Zhang, Wu & Zhang, 2023, G. khunkhamensis Sitthivong, Lo, Nguyen, Ngo, Khotpathoom, Le, Ziegler & Luu, 2021, G. liui Zhou, Zhou, Wang, Li, Zhang, Li, Shen, Liu & Rao, 2025, G. melli, G. palmatus Boulenger, Zoosyst. Evol. 101 (4) 2025, 2221–2242 zse.pensoft.net 2237 Figure 9. Habitat and field observations of Gekko tesselatus sp. nov. A. Macrohabitat of the new species in Se’ergu Town, Heishui County, Sichuan, China; B. Microhabitat of the new species; C. Gekko tesselatus sp. nov. in life, in situ; D. The sympatric Lycodon multizonatus. Photographs by YHX (A–C) and TXG (D). 1907, G. prep, G. scabridus Liu & Zhou, 1982, G. scientiadventura Rösler, Ziegler, Vu, Herrmann & Böhme, 2004, G. sengchanthavongi Luu, Calame, Nguyen, Le & Ziegler, 2015, G. shibatai Toda, Sengoku, Hikida & Ota, 2008, G. similignum, G. tawaensis Okada, 1956, G. thakhekensis Luu, Calame, Nguyen, Le, Bonkowski & Ziegler, 2014; G. truongi Phung & Ziegler, 2011; and G. vertebralis Toda, Sengoku, Hikida & Ota, 2008 (vs. 3–4 in G. aaronbaueri, 17–21 in G. adleri, 5 in G. canhi, 17–27 in G. chinensis, 4–5 in G. jinjiangensis, 9–12 in G. kaiyai, 0 in G. khunkhamensis, 20–23 in G. liui, 9–11 in G. melli, 23–30 in G. palmatus, 24 in G. prep, 10–15 in G. scabridus, 23–30 in G. scientiadventura, 4–5 in G. sengchanthavongi, 0–3 in G. shibatai, 17 in G. similignum, 0 in G. tawaensis, 1–5 in G. thakhekensis, 10–11 in G. truongi, and 0–1 in G. vertebralis). By having MBSR 98–106, Gekko tesselatus sp. nov. can be distinguished from G. bonkowskii Luu, Calame, Nguyen, Le & Ziegler, 2015; G. cib Lyu, Lin, Ren, Jiang, Zhang, Qi & Wang, 2021; G. hokouensis Pope, 1928; G. ichangensis Cao, Sucharitakul, Tie, Suwannapoom, Yan & Chomdej, 2025; G. japonicus (Duméril & Bibron, 1836); G. nadenensis Luu, Nguyen, Le, Bonkowski & Ziegler, 2017; and G. subpalmatus (Günther, 1864) (vs. 117 in G. bonkowskii, 128–149 in G. cib, 119–130 in G. hokouensis, 127–145 in G. ichangensis, 130–144 in G. japonicus, and 129–156 in G. nadenensis). By having DTR 12–15, Gekko tesselatus sp. nov. can be distinguished from G. auriverrucosus Zhou & Liu, 1982, G. guishanicus Lin & Yao, 2016, G. swinhonis, and G. wenxianensis Zhou & Wang, 2008 (vs. 16–20 in G. auriverrucosus, 0 in G. guishanicus, 6–8 in G. swinhonis, and 10 in G. wenxianensis). Moreover, by having LT4 9–12, Gekko tesselatus sp. nov. can be distinguished from G. alpinus Ma, Shi, Shen, Chang & Jiang, 2024, G. taibaiensis Song, 1985, and G. yakuensis Matsui & Okada, 1968 (vs. 13–15 in G. alpinus, 7–8 in G. taibaiensis, and 15 in G. yakuensis). Discussion Molecular phylogenetic analyses recovered Gekko tesselatus sp. nov. as a strongly supported monophyletic lineage (SH = 100 / UFB = 100) within the subgenus Japonigekko. This lineage is nested within a well-supported clade (SH = 100 / UFB = 100) that also includes G. fengshanensis, G. kwangsiensis, G. liboensis, and G. paucituberculatus, together forming a distinct evolutionary assemblage. The close genetic relationships among zse.pensoft.net Xu, Y. et al.: A new Gekko species 2238 Table 5. Comparison of morphological characters among Gekko tesselatus sp. nov., G. fengshanensis, G. kwangsiensis, G. liboensis, and G. paucituberculatus; differences are shown in bold. Species G. tesselatus sp. nov. G. fengshanensis G. kwangsiensis G. liboensis G. paucituberculatus N = 14 N = 6 N = 6 N = 2 N = 2 SVLmax (mm) 65.3 79.9 69.7 79.7 85.9 NS 3 or 4 3 3 3 3 INS 0 or 1 0 0 or 1 0 0 SL 10 or 11 9–13 10–13 11 11 IL 9–12 10–13 11–13 9–11 9–10 IOS 22–26 22–27 29–31 32–35 37 PO 12–15 15–19 18–20 17–18 14–18 PM 2 (rarely 1) 2 2 2 2 GP 3–6 3–6 4–6 4–6 4–6 DTR 12–15 9–11 9–11 9–10 4 GSDT 7–9 9–10 8–10 9–10 8 SMC 130–157 197–213 185–208 183–195 189–192 MBSR 98–106 149–161 143–156 131–140 136–142 VS 31–39 40–49 41–45 38–41 42–44 LF1 7–10 11–13 10–13 12–13 10–11 LF4 9–12 12–16 12–14 14–17 12–13 LT1 7–9 12–14 11–13 12–13 11 LT4 9–12 13–15 14–18 14–15 11–13 PP 6–8 9–12 9–10 9 12 PAT 1 (rarely 2) 1 1 1 1 Web Absent Present, weakly Present, weakly Present, weakly Pesent, weakly Fore tubercles Absent Absent Absent Absent Absent Hind tubercles Present Absent Absent Absent Absent Tail tubercles Present Absent Present – Absent Sources This study Huang et al. (2025) Yang et al. (2015) Huang et al. (2025) Wang et al. (2024) these taxa suggest a potentially shared evolutionary origin. However, despite its phylogenetic affinity with the karst-dwelling species, G. tesselatus sp. nov. exhibits markedly different ecological and geographic characteristics. The new species inhabits highly weathered rocky slopes within hot–dry valleys of northwestern Sichuan Province, in contrast to its congeners, which are confined to warm limestone karst forests in southern China. These contrasting habitat preferences suggest that environmental heterogeneity has likely played a significant role in driving divergence within this clade. Geographically, Gekko tesselatus sp. nov. is widely separated from its closest relatives. The nearest known rock-dwelling species, G. liboensis, occurs nearly 900 km away. It is therefore plausible that additional undescribed rock-dwelling species of Japonigekko exist in these underexplored areas. This observation underscores the need for more comprehensive and systematic fieldwork in southwestern and central China to improve our understanding of species diversity, distribution patterns, and evolutionary history within the subgenus Japonigekko. The discovery of Gekko tesselatus sp. nov. raises the total number of species in the subgenus Japonigekko to 39, with 26 recorded from China, including seven species currently known from Sichuan Province: Gekko tesselatus sp. nov., G. alpinus, G. chinensis (record questionable), G. cib, G. japonicus, G. jinjiangensis, and G. scabridus (Cai et al. 2018; Hou et al. 2021; Lyu et al. 2021; Ma et al. 2024). With the exception of the newly described species, all of the above mentioned taxa exhibit varying degrees of adaptability to human-modified environments, such as buildings, bridges, or concrete embankments. In contrast, field observations indicate that G. tesselatus sp. nov. is almost never found within human settlements. Even at sites adjacent to villages or towns, individuals were only observed on nearby undeveloped rocky slopes, suggesting that the species may be more sensitive to human disturbance than its congeners. Given the current lack of data on population size, trends, and distribution range, we recommend that Gekko tesselatus sp. nov. be preliminarily classified as Data Deficient (DD) under the IUCN Red List criteria. Further field surveys in adjacent montane regions are essential to better understand its conservation status and to assess whether the species may qualify for listing under a threatened category in the future. Zoosyst. Evol. 101 (4) 2025, 2221–2242 zse.pensoft.net 2239 Table 6. Morphological characters of Gekko (Japonigekko); bold fonts indicate differences from the new species; Y = “Yes” or present, N = “No” or absent; differences are shown in bold for clarity. No. Species SVLmax SL IL IOS DTR SMC MBSR VS LT 1 LT4 Web Fore tubercles Hind tubercles Tail tubercles PP 1G. tesselatus sp. nov. 65.3 10 or 11 9–12 22–26 12–15 130–157 98–106 31–39 7–9 9–12 N N Y Y 6–8 2G. aaronbaueri 80 13–14 10–11 34–37 0 – 98–104 39–43 14–17 14–16 Y NN N 3–4 3G. adleri 75.3 10–15 9–13 27–36 7–11 168–190 123–144 35–44 11–14 11–15 YN Y Y 17–21 4G. alpinus 74.16 9–13 8–10 22–28 12–17 158–189 92–114 32–39 8–11 13–15 NYY Y 4–7 5G. auriverrucosus 69 9–11 9–11 26–27 16–20 – – – 6–8 6–8 N YY Y 8–11 6G. bonkowskii 69.2 12–14 10–11 49–50 0 154–169 117 37–40 11–13 15 Y NN N 6 7G. canhi 99.2 14 10–12 49–50 11–12 168–170 205–227 49–51 13–16 14–17 N N Y N5 8G. chinensis 72 10–14 9–13 35–48 10 156–167 118–140 37–39 8–10 9–12 YN Y Y 17–27 9G. cib 66.4 10–12 10–14 28–36 0 171–196 128–149 37–45 9–13 9–17 YNN N 7–9 10 G. fengshanensis 79.9 9–13 10–13 22–27 9–11 197–213 149–161 40–49 12–14 13–15 Y NN N 9–12 11 G. guishanicus 64 – – – 0– – – 8–10 8–10 YNN N 6–8 12 G. hokouensis 70 10–14 8–11 30–33 12–18 153–174 119–130 36–43 8–11 15–18 N N NY 5–9 13 G. ichangensis 66.5 7–11 7–11 27–36 10–12 174–195 127–145 39–47 12–15 13–18 Y Y Y Y 5–8 14 G. japonicus 74 9–13 8–13 32–35 9–14 169–188 130–144 39–44 10–12 14–16 NYY Y 4–9 15 G. jinjiangensis 61.6 7–10 6–9 20–24 12–16 146–169 111–149 31–47 8–11 11–15 N YY Y 4–5 16 G. kaiyai 64.99 9–12 9–13 22–33 11–18 157–209 99–121 30–43 8–9 7–11 N YY Y 9–12 17 G. khunkhamensis 75.2 9–10 9–10 31–32 0 181–185 127–138 42–45 13–14 14–15 Y NN N 0 18 G. kwangsiensis 69.7 10–12 11–13 29–31 9–11 185–208 143–156 41–45 11–13 13–18 Y NNY9–11 19 G. liboensis 79.7 11 9–11 32–35 9–10 183–195 131–140 38–41 12–13 14–15 Y NN–9 20 G. liui 56.67 11–13 11–12 37–42 7–8 140–148 103–118 34–39 10–11 12–13 YNNY20–23 21 G. melli 80.3 10–13 9–12 34–40 0 171–192 148–160 44–46 10–12 11–14 YNN N 9–11 22 G. nadenensis 77.1 12–14 10–12 28–30 0 175–185 123–140 38–40 13–15 14–16 Y NN N 6 23 G. palmatus 79.7 11–15 9–13 27–36 4–12 160–191 116–147 36–47 10–13 10–16 YNNY23–30 24 G. paucituberculatus 85.9 11 9–10 37 4 189–192 136–140 42–44 11 11–13 YNN N 12 25 G. prep 75.7 14 11 37–38 12 144–152 122–126 38–40 11–12 12–13 YNNY24 26 G. scientiadventura 73 12–14 9–13 41–51 0 118–140 139–143 38–48 12–15 14–17 Y NNY23–30 27 G. scabridus 64 9–11 9–11 30 17–21 – – – 6–9 7–9 N YY Y 10–15 28 G. sengchanthavongi 77.3 8–10 6–7 28–32 0 175–184 120–135 35–43 11–14 13–17 YNN N 4–5 29 G. shibatai 70.9 10–13 10–14 37–52 5–14 – 114–134 –11 or 12 9–16 N N NY0–3 30 G. similignum 58.9 12–14 11 46–48 11 –144–153 –11–13 12–14 YNNY17 31 G. subpalmatus 65.8 8–12 7–12 28–37 0 144–190 129–156 39–46 9–12 11–14 YNN N 5–9 32 G. swinhonis 66 7–12 7–11 23–24 6–8 – – 40 6–9 6–9 N YY – 7–9 33 G. taibaiensis 69 9–10 8–10 28 – – – – 6–7 7–8 Y – – – 4–6 34 G. tawaensis 71 15 13 –0– – – 10 12 N N N N 0 35 G. thakhekensis 79.2 12–14 10–11 22–26 0 165–174 110–116 32–40 11–13 14–15 Y NN N 1–5 36 G. truongi 95.9 13–15 11–13 45–48 0 160–172 131–143 35–36 11–13 15–17 N N N N 10–11 37 G. vertebralis 69.2 10–15 10–15 35–50 2–12 – 112–139 10–12 – 9–17 N N N N 0–1 38 G. wenxianensis 59 12 11 – 10 – – 42–44 6 9 N N Y – 6–8 39 G. yakuensis 72 12–13 9–13 – – – – – 10 15 N N NY 6–8 zse.pensoft.net Xu, Y. et al.: A new Gekko species 2240 Acknowledgments We would like to express our gratitude to Mr. Jiaxiang Wu (China), Danyang Zhou (China), and Maozhou Xu (China) for their help with fieldwork and photography. We also thank Mr. Tierui Zhang (AHU, China), Hanming Song (SYS, China), and Haotian Wang (HAINNU, China) for assistance with data analysis and manuscript polishing. We are grateful to the anonymous reviewer for constructive comments on an earlier version of the manuscript. This research was funded by the National Natural Science Foundation of China [32301325], the Open Project of the State Key Laboratory of Plateau Ecology and Agriculture, Qinghai University [2025-KF-02], the China Biodiversity Observation Networks (Sino BON–Amphibian & Reptile), and the Russian Science Foundation (RSF grant Nº 22-1400037-P, supporting the work of Dr. Nikolay A. Poyarkov). References Brown RM, Oliveros CH, Siler CD, Diesmos AC (2009) Phylogeny of Gekko from the Northern Philippines, and description of a new species from Calayan Island. Journal of Herpetology 43(4): 620–635. https://doi.org/10.1670/08-207.1 Burland TG (2000) DNASTAR’s Lasergene sequence analysis software. Methods in Molecular Biology (Clifton, N.J.) 132: 71–91. https://doi.org/10.1385/1-59259-192-2:71 Cai B, Lyu K, Chen YY, Li JT, Wang YZ, Gu HJ, Gu XD (2018) The distributional list of amphibians and reptiles in Sichuan Province, China. Science Data Bank 3(1). [in Chinese with English abstract]. https://doi.org/10.11922/sciencedb.524 Cao J, Sucharitakul P, Tie M, 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 Figure 10. Comparison of Gekko tesselatus sp. nov. and four closely related species in life, in situ. A. Gekko tesselatus sp. nov., unvouchered adult male individual from Se’ergu, Heishui, Sichuan, China; B. Gekko tesselatus sp. nov., unvouchered juvenile individual from Se’ergu, Heishui, Sichuan, China; C. G. liboensis, unvouchered individual from Libo, Guizhou, China; D. G. fengshanensis, NHMG 202408007, adult male from Fengshan, Guangxi, China; E. G. kwangsiensis, unvouchered adult female individual from Nanning, Guangxi, China; F. G. paucituberculatus, unvouchered individual from Baise, Guangxi, China. Photographs by YHX (A, B), Dan-Yang Zhou (C, F), reproduced from Huang et al. (2025) (D), and Yang (2015) (E). Zoosyst. Evol. 101 (4) 2025, 2221–2242 zse.pensoft.net 2241 Darko YA, Voss O, Uetz P (2022) A dictionary of abbreviations used in reptile descriptions. Zootaxa 5219(5): 421–432. https://doi.org/10.11646/ zootaxa.5219.5.2 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 Goris RC, Maeda N (2004) Guide to the Amphibians and Reptiles of Japan. Florida, Malabar: Krieger Publishing Company, 176–178. 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 Hoang DT, Chernomor O, Haeseler AV, Minh BQ, Vinh LS (2018) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35(2): 518–522. https://doi.org/10.1093/ molbev/msx281 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 Huang Z, Wang HT, Qi S, Song HM, Huang Y, Wang YY, Mo YM (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 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 Kalyaanamoorthy S, Minh BQ, Wong TKF, Haeseler AV, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589. https://doi.org/10.1038/nmeth.4285 Kluge AG (1993). Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego, 245 pp. Kluge AG (2001) Gekkotan lizard taxonomy. Hamadryad 26: 1–209. Kumar S, Stecher G, Li M, Knyaz C, Tamura K (2018) MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution 35(6): 1547–1549. https://doi. org/10.1093/molbev/msy096 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 SC, Shen C, Chang LM, Jiang JP (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 Ngo VT, Pham TH, Phimvohan A, David P, Teynié A (2015) Gekko aaronbaueri, a new gecko (Squamata: Gekkonidae) from central Laos. Zootaxa 3914(2): 144–156. https://doi.org/10.11646/zootaxa.3914.2.4 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 Nguyen LT, Schmidt HA, Haeseler AV, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274. https://doi.org/10.1093/molbev/msu300 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 Phung TM, Ziegler T (2011) Another new Gekko species (Squamata: Gekkonidae) from southern Vietnam. Zootaxa 3129(1): 51–61. https://doi.org/10.11646/zootaxa.3129.1.4 Rambaut A (2018) FigTree v1.4.4. http://tree.bio.ed.ac.uk/software/figtree/ [accessed 10 April 2025] Rösler H, Ziegler T, Vu NT, Herrmann HW, Böhme W (2005) “2004”) A new lizard of the genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from the Phong Nha-Ke Bang National Park, Quang Binh Province, Vietnam. Bonner Zoologische Beiträge 53: 135–148. 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] zse.pensoft.net Xu, Y. et al.: A new Gekko species 2242 Stejneger L (1907) Herpetology of Japan and adjacent territory. Bulletin – United States National Museum 58(58): 165–169. https://doi. org/10.5479/si.03629236.58.i Stephane G, Jean-franc OD, Vincent L, Maria A, Wim H, And OG (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology 59(3): 307–321. https://doi.org/10.1093/sysbio/syq010 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 Uetz P, Freed P, Aguilar R, Reyes F, Hošek J (2025) The Reptile Database. http://www.reptile-database.org [accessed 15 October 2025] 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 Xia LJ, Cai FN, Chen SS, Cai Y, Zhou KY, Yan J, Li P (2023) Phylogenetic analysis and genetic structure of Schlegel’s Japanese gecko (Gekko japonicus) from China based on mitochondrial DNA sequences. Genes 14(1): 18. https://doi.org/10.3390/genes14010018 Yang JH (2015) A new species of the genus Gekko Laurenti (Squamata: Sauria: Gekkonidae) from Guangxi, China. Zootaxa 3936(2): 287– 295. https://doi.org/10.11646/zootaxa.3936.2.9 Zhang D, Gao FL, Jakovlić I, Zou H, Zhang J, Li WX, Wang GT (2020) PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20(1): 348–355. https://doi.org/10.1111/1755-0998.13096 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 (Basel) 13(24): 3796. https://doi. org/10.3390/ani13243796 Zhao E, Adler K (1993) Herpetology of China. Oxford, Ohio, 522 pp. [+48 plates] Zhao E, Zhao K, Zhou K (1999) Fauna Sinica Reptilia, Vol. 2: Squamata, Lacertilia. Science Press, Beijing, [xi +] 394 pp. [in Chinese] Zhou K, Wang Q (2008) New species of Gekko (Squamata: Sauria: Gekkonidae) from China: morphological and molecular evidence. Zootaxa 1778(1): 59–68. https://doi.org/10.11646/zootaxa.1778.1.5 Zhou KY, Liu YZ, Li DJ (1982) Three new species of Gekko and remarks on Gekko hokouensis (Lacertiformes, Gekkonidae). Dong Wu Fen Lei Xue Bao 7: 438–446. [in Chinese] Zhou KY, Li HD, Han DM, Bauer AM, Feng JY (2006) The complete mitochondrial genome of Gekko gecko (Reptilia: Gekkonidae) and support for the monophyly of Sauria including Amphisbaenia. Molecular Phylogenetics and Evolution 40(3): 887–892. https://doi. org/10.1016/j.ympev.2006.04.003 Zhou RB, Chen KZ, Liu QM, Li XJ, Qi XM, Li JL, Peng XP (2025a) Description of a new species of the subgenus Japonigekko (Squamata: Gekkonidae: Gekko) from Hainan, China. Zootaxa 5633(3): 528–540. https://doi.org/10.11646/zootaxa.5633.3.6 Zhou RB, Zhou HX, Wang HS, Li J, Zhang DR, Li XY, Shen ZQ, Liu S, Rao DQ (2025b) Taxonomic review of Gekko (Japonigekko) species in Hainan Province and description of a new species. Asian Herpetological Research 16(2): 247–259. https://doi.org/10.3724/ahr.20950357.2024.0055