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Research Article
First description of the female of Cyrtodactylus dianxiensis Liu & Rao, 2021, with extended diagnosis of this species (Squamata, Gekkonidae)
expand article infoDongru Zhang§, Yunhe Wu, Changsheng Zuo|, Fawang Yin|, Shuo Liu
‡ Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China
§ Liupanshui Normal University, Liupanshui, China
| Yingjiang sub-office, Tongbiguan Provincial Natural Reserve Management and Protection Bureau, Yingjiang, China
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Abstract

Cyrtodactylus dianxiensis Liu & Rao, 2021 was originally described based on only two adult male specimens from Tongbiguan Nature Reserve, Dehong Autonomous Prefecture, western Yunnan, China. So far, no information on the females of this species is available. During comprehensive herpetofaunal investigations in 2022, one female specimen of C. dianxiensis was collected from Tongbiguan Nature Reserve. The female specimen agrees well with the original description of C. dianxiensis, and also shows some slight differences in coloration. This study reported the female specimen of this species for the first time, and provided a description and photos of the female specimen; meanwhile, we extended the diagnosis of this species.

Key Words

bent-toed gecko, China, morphology, Yunnan

Introduction

Cyrtodactylus Gray, 1827 (bent-toed geckos) is the most speciose and ecologically diverse gekkonid genus, with more than 350 recognized species distributed from the Western Himalayas through southeast Asia to the Western Pacific (Wood et al. 2012; Uetz et al. 2024). Most species of Cyrtodactylus are karst-adapted or granite-adapted species (Grismer et al. 2020). With the in-depth investigation of karst areas, the underestimated biodiversity of Cyrtodactylus has been gradually revealed (Grismer et al. 2018; Liu et al. 2021, 2023).

So far, ten species of Cyrtodactylus have been recognized in China, and all are distributed in southwestern China, namely Yunnan Province and Tibet Autonomous Region. Four of them were recorded in Tibet: C. tibetanus (Boulenger, 1905), C. cayuensis Li, 2007, C. zhaoermii Shi & Zhao, 2010 and C. kamengensis Mirza, Bhosale, Thackeray, Phansalkar, Sawant, Gowande, Patel & Kamengensis, 2022. The other six were recorded in Yunnan: C. dianxiensis Liu & Rao, 2021, C. gulinqingensis Liu, Li, Hou, Orlov & Ananjeva, 2021, C. hekouensis Zhang, Liu, Bernstein, Wang & Yuan, 2021, C. zhenkangensis Liu & Rao, 2021, C. menglianensis Liu & Rao, 2022 and C. caixitaoi Liu, Rao, Hou, Wang & Ananjeva, 2023, among which, C. dianxiensis belong to the C. khasiensis species group, and the remaining belong to the C. chauquangensis species group.

Cyrtodactylus dianxiensis is similar to C. khasiensis and previously the species have been confused with one another (Liu and Rao 2021; Wang et al. 2022). Liu and Rao (2021) describe the population of C. khasiensis distributed in Western Yunnan as a new species, C. dianxiensis. Cyrtodactylus dianxiensis, currently the only forest-dwelling Cyrtodactylus species recorded in Yunnan Province, China, was described based on one adult male holotype (KIZL2019044) from Tongbiguan Township, Yingjiang County, and one adult male paratype (KIZ2003170) from Longchuan County, both in Tongbiguan Nature Reserve, Dehong Autonomous Prefecture, western Yunnan, China (Liu and Rao 2021); no female information about this species was available. Identification verification of C. khasiensis voucher specimens deposited in museums could potentially yield additional specimens of C. dianxiensis. However, we have not yet been given the opportunity to examine the relevant specimens in other museums.

During a comprehensive herpetofaunal survey of Tongbiguan Nature Reserve in 2022, one female specimen of Cyrtodactylus was collected from Tongbiguan Nature Reserve, Xueli Village, Taiping Town, Yingjiang County, Dehong Prefecture, Yunnan Province, China (Fig. 1). Morphological and molecular phylogenetic analyses revealed that the specimen is C. dianxiensis. Here, we provide morphological data for the newly collected female specimen of C. dianxiensis, and extend the diagnosis of this species.

Figure 1. 

Sampling sites of Cyrtodactylus dianxiensis: solid black star, the locality of the holotype; solid black triangle, the locality of the paratype; solid green circle, the locality of the newly collected female specimen.

Materials and methods

Sampling

The specimen was sampled by hand at night. Sex was further determined based on whether the ventral side of the tail base is swollen. After euthanasia, liver tissues were taken and preserved in 95% alcohol. The specimen was then directly preserved in 75% ethanol without formalin fixation. The voucher specimen was deposited in Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences (KIZ).

Morphological analyses

The adult female specimen preserved in 75% ethanol was measured with digital calipers to the nearest 0.1 mm. Morphological terminology followed Liu and Rao (2021). Measurements included the following: snout-vent length (SVL), from tip of snout to anterior margin of cloaca ; trunk length (TRL), axilla to groin distance ; body width (BW), maximum width of body ; tail length (TL), from posterior margin of cloaca to tip of tail ; tail width (TW), maximum width of tail ; head length (HL), from tip of snout to posterior margin of jaw ; head width (HW), maximum width of head ; head height (HH), from occiput to underside of jaws ; forearm length (FL), from the base of the palm to the elbow ; crus length (CL), from the base of heel to the knee ; eye diameter (ED), greatest diameter of eye ; nostril to eye distance (NE), from nostril to anterior corner of eye orbit ; snout to eye distance (SE), from tip of snout to anterior corner of eye orbit ; orbit to ear distance (EE), from posterior corner of eye orbit to anterior margin of ear opening ; ear length (EL), greatest diameter of ear ; internarial distance (IN), measured between inner borders of nostrils ; interorbital distance (IO), measured across narrowest point of frontal bone. The counting of the following characteristics also followed Liu and Rao (2021): dorsal tubercle rows (DTR) ; ventral scale rows (MVSR) ; paravertebral tubercles (PVT) ; supralabials (SL) ; infralabials (IL) ; precloacal pores (PcP) ; precloacofemoral pores (PcFP) ; femoral pores on each thigh (FP) ; subdigital lamellae under the fourth finger (LF4) and under the fourth toe (LT4).

DNA extraction, PCR amplification, and sequencing

Total genomic DNA was extracted from tissue samples preserved in 95% ethanol. The tissue sample was then digested using proteinase K, and subsequently purified using DNeasy Tissue Kit (QIAGEN). A fragment of NADH Dehydrogenase subunit 2 (ND2) was amplified and sequenced using the primers L4437b and H5934 (Macey et al. 1997). The experiment protocols are the same as Liu and Rao (2021). New sequence was assembled and edited using SEQMAN in Lasergene 7.1 (DNASTAR Inc., Madison, WI, USA).

Molecular analysis

Phylogenetic relationships within the C. khasiensis species group were inferred from ND2. The homologous sequences of the C. khasiensis species group, and the outgroups species C. slowinskii, were downloaded from GenBank (Clark et al. 2016) (Table 1). Sequences were aligned using MUSCLE 3.6 (Edgar 2004), then checked by eye for accuracy and trimmed to minimize missing characters in MEGA 6.0.6 (Tamura et al. 2013).

Table 1.

Locality, voucher ID, and GenBank accession (ND2) for all samples used in this study. * denotes the holotype of Cyrtodactylus dianxiensis.

Species Locality Voucher Accession
Cyrtodactylus aaronbaueri Mizoram, India MZMU 2015 MW596520
Cyrtodactylus agarwali Meghalaya, India MZMU 2158 MW596515
Cyrtodactylus arunachalensis Arunachal Pradesh, India BNHS 2777 MT341522
Cyrtodactylus aunglini Mandalay, Myanmar LSUHC 13948 MH764589
Cyrtodactylus ayeyarwadyensis Ayeyarwady, Myanmar CAS 212459 JX440526
Cyrtodactylus bapme Meghalaya, India BNHS 2756 MW367435
Cyrtodactylus bengkhuaiai Mizoram, India MZMU 1985 MW596516
Cyrtodactylus chrysopylos Shan State, Myanmar LSUHC 13937 MH764604
Cyrtodactylus dianxiensis Yunnan, China KIZ059201 MW971927
Cyrtodactylus dianxiensis Yunnan, China KIZL2019044* MW971926
Cyrtodactylus dianxiensis Yunnan, China KIZ2022159 PP394340
Cyrtodactylus exercitus Meghalaya, India MZMU 2545 OK247679
Cyrtodactylus gansi Chin State, Myanmar CAS 222412 JX440537
Cyrtodactylus guwahatiensis Assam, India BNHS 2146 KM255194
Cyrtodactylus jaintiaensis Meghalaya, India BNHS 2248 KM255195
Cyrtodactylus karsticola Meghalaya, India MZMU 2156 MW596513
Cyrtodactylus kazirangaensis Assam, India BNHS 2147 KM255170
Cyrtodactylus khasiensis Meghalaya, India BNHS 2249 KM255188
Cyrtodactylus lungleiensis Mizoram, India MZMU 2428 MZ645742
Cyrtodactylus montanus Tripura, India BNHS 2231 KM255200
Cyrtodactylus mombergi Kachin State, Myanmar LSUHC 14734 MN059875
Cyrtodactylus nagalandensis Nagaland, India BNHS 2253 KM255199
Cyrtodactylus namtiram Manipur, India BNHS 2751 MW367433
Cyrtodactylus ngopensis Mizoram, India MZMU 2360 OM912605
Cyrtodactylus septentrionalis Assam, India BNHS 1989 MH971164
Cyrtodactylus siahaensis Mizoram, India MZMU 2445 OK247677
Cyrtodactylus tripuraensis Tripura, India BNHS 2238 KM255183
Cyrtodactylus urbanus Assam, India VR/ERS/ZSI/688 MN911174
Cyrtodactylus vairengtensis Mizoram, India MZMU 2903 OP874800
Cyrtodactylus slowinskii Sagaing, Myanmar CAS 210205 JX440559

Phylogenetic reconstruction was conducted using Bayesian inference (BI) and maximum likelihood (ML) methods based on the ND2 gene. The best-fit substitution model was selected under the Bayesian Information Criterion by the program MODELFINDER (Kalyaanamoorthy et al. 2017) implemented in IQ-TREE 1.6.12 (Nguyen et al. 2015). Bayesian inference analyses were performed in MRBAYES 3.2.7 (Ronquist et al. 2012) based on the model GTR+F+I+G4. Two independent runs were initiated, each with four simultaneous Markov Chain Monte Carlo (MCMC) chains for one million generations and sampled every 100 generations with a burn-in of 25%. The convergence was examined with an average standard deviation of split frequencies less than 0.01 and effective sample size (ESS) values greater than 200 in TRACER 1.5 (Rambaut and Drummond 2009). Maximum likelihood analyses were conducted using IQ-TREE 1.6.12 (Nguyen et al. 2015), based on the model TIM+F+I+G4. Nodal support was estimated by 1,000 bootstrap replicates using the ultrafast bootstrap feature. Pairwise divergences were calculated using uncorrected p-distances implemented in MEGA 6.0.6 (Tamura et al. 2013).

Results

Bayesian inference and ML trees showed consistent topology. The newly collected female specimen clustered with the specimens (including the holotype) of C. dianxiensis with strong support by both BI and ML (BI/ML=1/100, Fig. 2). The genetic distance (uncorrected p-distance) between the newly collected female specimen and the specimens (including the holotype) of C. dianxiensis was only 0.9% (Table 2).

Table 2.

Uncorrected pairwise divergence (%) between Cyrtodactylus khasiensis species group members based on ND2 sequences.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
1 C. aaronbaueri
2 C. agarwali 24.3
3 C. arunachalensis 23.4 20.6
4 C. aunglini 19.6 20.8 24.2
5 C. ayeyarwadyensis 21.9 14.8 20.2 18.2
6 C. bapme 22.9 7.2 20.8 20.7 14.2
7 C. bengkhuaiai 12.0 23.6 20.9 18.3 19.9 21.9
8 C. chrysopylos 21.7 21.4 23.1 18.5 18.9 21.9 19.2
9 C. dianxiensis 21.2 20.3 21.9 16.6 18.8 20.4 21.0 18.8
10 KIZ2022159 21.0 20.6 21.8 19.8 21.7 20.7 21.0 22.0 0.9
11 C. exercitus 22.5 11.8 18.4 21.7 16.2 13.6 22.5 21.9 21.3 21.4
12 C. gansi 19.2 21.0 22.0 15.8 16.9 20.9 18.2 17.6 15.5 18.4 21.1
13 C. guwahatiensis 22.6 12.5 20.3 21.6 15.4 13.6 22.1 22.9 22.4 22.7 4.5 21.1
14 C. jaintiaensis 19.2 23.1 22.9 19.1 21.8 22.5 17.3 22.0 19.1 19.3 21.4 19.0 22.4
15 C. karsticola 22.9 6.4 19.1 19.2 12.6 6.7 23.8 21.1 20.0 20.1 13.4 21.1 13.1 22.0
16 C. kazirangaensis 24.7 20.0 21.5 22.5 16.7 19.6 22.9 23.7 21.0 21.3 18.3 21.7 19.8 23.0 17.1
17 C. khasiensis 22.7 17.2 21.0 21.5 15.6 16.9 19.5 22.1 21.9 22.6 17.0 20.0 16.9 21.5 15.2 18.5
18 C. lungleiensis 12.1 21.6 19.9 18.3 19.2 20.7 7.2 20.4 20.3 19.7 20.2 18.5 21.1 18.0 22.0 21.8 18.8
19 C. mombergi 20.0 20.4 20.4 17.4 17.3 19.9 20.4 19.2 17.2 20.3 20.4 15.9 21.8 19.7 19.6 21.0 20.3 18.4
20 C. montanus 10.3 22.9 22.6 19.6 20.0 20.5 12.1 21.5 19.5 20.1 20.0 19.6 21.9 18.8 22.0 23.3 21.7 10.9 19.6
21 C. nagalandensis 21.5 24.8 22.3 19.8 21.7 23.8 19.2 22.7 19.0 19.4 22.6 18.0 23.6 19.0 22.0 23.1 22.0 19.1 20.6 19.6
22 C. namtiram 15.7 22.6 22.5 20.4 20.9 22.3 14.2 20.7 21.9 22.0 23.3 20.3 23.3 20.0 24.1 23.7 21.9 13.9 19.5 15.4 19.4
23 C. ngopensis 9.9 23.6 22.0 18.3 20.7 21.1 11.2 20.3 20.2 19.9 21.0 18.4 21.6 17.8 22.7 24.2 22.2 11.5 19.6 10.5 19.7 14.8
24 C. septentrionalis 21.2 11.1 20.6 20.8 14.6 11.6 20.5 21.7 20.3 21.0 10.9 19.8 11.3 21.4 10.7 18.7 15.7 19.1 20.6 19.6 23.6 21.5 20.1
25 C. siahaensis 11.4 22.9 20.0 18.7 20.1 20.9 8.6 20.8 19.3 19.1 21.8 18.9 21.1 17.5 23.4 22.5 19.4 6.1 19.1 11.0 17.2 14.2 10.3 19.3
26 C. tripuraensis 21.8 13.2 18.4 18.8 7.5 12.3 18.4 19.6 18.9 19.8 14.7 16.7 12.9 18.6 13.1 14.5 12.5 19.1 17.1 18.2 19.0 21.1 18.8 11.5 18.6
27 C. urbanus 23.8 15.6 20.8 21.3 13.7 15.1 21.5 21.5 20.8 21.1 14.9 20.0 15.2 21.8 14.3 17.3 14.2 20.1 20.5 21.8 22.9 21.6 21.4 13.4 20.1 11.5
28 C. vairengtensis 4.0 24.4 24.2 18.9 21.1 22.2 11.2 21.1 20.8 20.5 21.9 18.5 22.2 18.8 22.3 24.5 22.8 11.7 20.4 10.2 20.1 15.9 10.3 20.5 10.6 20.0 22.3
Figure 2. 

Bayesian phylogram of the Cyrtodactylus khasiensis species group inferred from ND2 sequences. Numbers before slashes indicate Bayesian posterior probabilities and numbers after slashes indicate the ML ultrafast bootstrap. Values below 0.90/90 are not shown.

Cyrtodactylus dianxiensis Liu & Rao, 2021

Figs 3, 4, Table 3

Specimen examined

KIZ2022159, adult female, collected on 13 September 2022 by Shuo Liu from Tongbiguan Nature Reserve, Xueli Village, Taiping Town, Yingjiang County, Dehong Prefecture, Yunnan Province, China (24°26'35"N, 97°33'1"E; at an elevation of 380 m) (Fig. 1).

Figure 3. 

The female specimen (KIZ2022159) of Cyrtodactylus dianxiensis in life. A. Dorsal view; B. Close-up dorsal view of the head; C. Close-up dorsolateral view of the head.

Description of the female specimen

SVL 75.0 mm; head relatively large (HL/SVL 0.26, HW/HL 0.68), depressed (HH/HL 0.45), distinct from neck; loreal and interorbital region concave, canthus rostralis slightly swollen; snout moderately long (SE/HL 0.40); eye large (ED/HL 0.24); pupil vertical with crenulated margins; ear opening oval, obliquely oriented (EL/HL 0.08); rostral with midrostral suture dorsally; two large supranasals, separated by one small internasal; dorsal head scales heterogeneous, rounded, granules; mental triangular with a pair of enlarged postmentals followed by some gradually decreasing chin-shields; scales on other region of the ventral head almost homogeneous, small, rounded, granules; supralabials seven to midorbital position on both sides, nine to angle of the jaw on right side and ten to angle of the jaw on the left side; infralabials nine on right side and seven on left side.

Figure 4. 

The female specimen (KIZ2022159) of Cyrtodactylus dianxiensis in preservative. A. Dorsal view; B. Ventral view; C. Close-up view of the precloacal and femoral regions.

Body slender (TRL/SVL 0.45); dorsal scales heterogeneous, primarily small rounded granules, intermixed with irregularly arranged large circular tubercles, tubercles on nape and occipital region smaller than those on dorsum; ventrolateral folds present; ventral scales larger than dorsal; enlarged femoral scales and femoral pores absent; precloacal scales enlarged, precloacal pores absent; no precloacal groove; cloacal spurs indistinct.

Table 3.

Measurements (mm) and meristic data for the female specimen of Cyrtodactylus dianxiensis. See Materials and methods for abbreviations.

KIZ2022159 KIZ2022159
SVL 75.0 EE 5.9
TRL 33.8 EL 1.6
BW 11.6 IN 2.5
TL 73.7 IO 2.6
TW 6.8 FP 0
HL 19.3 PcP 0
HW 13.2 MVSR 35
HH 8.6 PVT 32
FL 11.5 DTR 17
CL 13.3 SL (R/L) 9(7)/10(7)
ED 4.6 IL (R/L) 9/8
NE 6.1 LF4 17/16
SE 7.8 LT4 20/20

Limbs slender (FL/SVL 0.15, CL/SVL 0.18); digits strongly inflected at each joint, all bearing robust, recurved claws; relative length of digits: I<II<V<III<IV (manus) and I<II<III<V<IV (pes); scales on dorsal forelimbs heterogeneous, granules, with small tubercles interspersed, scales on dorsal hind limbs heterogeneous, granular, with large conical tubercles interspersed; ventral scales of limbs almost homogeneous, granular, smaller than those on ventral body.

Original tail broken but exists, subequal to body length (TL/SVL 0.98); dorsal tail scales heterogeneous, with small tubercles interspersed; two rows of subcaudal scales slightly enlarged.

Coloration in life

Dorsal surface of head almost uniform brownish gray; upper lip brownish gray with many white spots, an indistinct black postorbital streak that extends backwards above ear opening on each side; a disconnected W-shaped black stripe on the occiput; dorsal surface of body brownish gray with many short black and white streaks roughly forming longitudinal rows; dorsal surface of limbs gray with indistinct white bands and spots; dorsal surface of tail grayish black with ten white bands; ventral surfaces of head, body, and limbs white, ventral surface of tail checkered with brownish gray and white; iris bronze with dark reticulations, edge of pupil orange red.

Extended diagnosis

Body size moderate (SVL 73.8–79.9 mm in adults); 9–12 supralabials; 8–11 infralabials; 17–19 longitudinal rows of rounded, conical dorsal tubercles; 31–32 paravertebral tubercles; 35–41 ventral scales between ventrolateral folds; no precloacal groove; femoral scales not enlarged, no femoral pores; 7–8 precloacals in males, no precloacal in female; 16–17 total subdigital lamellae beneath finger IV, 19–20 total subdigital lamellae beneath toe IV; subcaudal scales not transversely enlarged or two median rows slightly enlarged. Dorsum with light and dark blotches roughly forming longitudinal markings; a W-shaped black stripe present on occipital region; tail with 8–10 alternating dark and light bands; iris blueish gray or bronze with orange red edge.

Discussion

The female specimen agrees well with the original description (Liu and Rao 2021) of C. dianxiensis in measurements and meristic counts, however, there are some slight differences in coloration between them. The female specimen has a relatively lighter body color; the dorsal head of the holotype of C. dianxiensis is mixed with brown and grayish white, while it is almost uniform brownish gray in the female specimen; the iris of the holotype is blueish gray with no orange red edge, while the iris is bronze with orange red edge in the female specimen. Therefore, in this species, the coloration of dorsal head is not always mottled, but may also be uniform; in addition, the iris of this species has two color types, blueish gray and bronze with orange red edge.

The holotype of C. dianxiensis was collected at an altitude of 1170 m and the paratype was collected at an altitude of 1200 m (Liu and Rao 2021). However, the female specimen was collected at a much lower altitude. This record extends the lowest elevational distribution of this species to 380 m.

Cyrtodactylus dianxiensis was previously known based on only two adult males and one juvenile (Liu and Rao 2021). This study reported the female specimen of this species for the first time, and provided a detailed description and photos of the female of this species. However, we have only collected one female specimen of this species, and it is unknown whether some differences between male and female are permanent. In addition, the range and population status of this species will need to be determined by collecting more specimens in the future.

Acknowledgements

Thanks to the editors and reviewers for their work on the manuscript. We also would like to thank Yunnan key laboratory of biodiversity and ecological conservation of Gaoligong Mountain for its support. This study was supported by the Science-Technology Basic Condition Platform from the Ministry of Science and Technology of the People’s Republic of China (No. 2005DKA21402) and Yunnan Applied Basic Research Projects (No. 202301AT070312).

References

  • Grismer LL, Wood Jr. PL, Le MD, Quah ESH, Grismer JL (2020) Evolution of habitat preference in 243 species of Bent-toed geckos (Genus Cyrtodactylus Gray, 1827) with a discussion of karst habitat conservation. Ecology and Evolution 10: 13717–13730. https://doi.org/10.1002/ece3.6961
  • Grismer LL, Wood Jr PL, Thura MK, Zin T, Quah ESH, Murdoch ML, Grismer MS, Lin A, Kyaw H, Lwin N (2018) Twelve new species of Cyrtodactylus Gray (Squamata: Gekkonidae) from isolated limestone habitats in east-central and southern Myanmar demonstrate high localized diversity and unprecedented microendemism. Zoological Journal of the Linnean Society 182: 862–959. https://doi.org/10.1093/zoolinnean/zlx057
  • Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS (2017) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589. https://doi.org/10.1038/nmeth.4285
  • Liu S, Rao DQ, Hou M, Wang QY, Ananjeva NB (2023) A new species of Cyrtodactylus Gray, 1827 (Squamata, Gekkonidae), previously confused with C. wayakonei Nguyen, Kingsada, Rösler, Auer et Ziegler, 2010. Russian Journal of Herpetology 30: 529–538. https://doi.org/10.30906/1026-2296-2023-30-6-529-538
  • Macey JR, Larson A, Ananjeva NB, Fang Z, Papenfuss TJ (1997) Two novel gene orders and the role of light-strand replication in rearrangement of the vertebrate mitochondrial genome. Molecular Biology and Evolution 14: 91–104. https://doi.org/10.1093/oxfordjournals.molbev.a025706
  • Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32: 268–274. https://doi.org/10.1093/molbev/msu300
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Hohna 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
  • Tamura K, Stecher G, Peterson D, Filipski A, Kumar S (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution 30: 2725–2729. https://doi.org/10.1093/molbev/mst197
  • Wang K, Lyu ZT, Wang J, Qi S, Che J (2022) The updated checklist and zoogeographic division of the reptilian fauna of Yunnan Province, China. Biodiversity Science 30: 21326. https://doi.org/10.17520/biods.2021326
  • Wood PL, Heinicke MP, Jackman TR, Bauer AM (2012) Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of diversification. Molecular Phylogenetics and Evolution 65: 992–1003. https://doi.org/10.1016/j.ympev.2012.08.025
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