Research Article |
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Corresponding author: Yunhe Wu ( yunhe2009@163.com ) Corresponding author: Shuo Liu ( liushuo@mail.kiz.ac.cn ) Academic editor: Ben Wielstra
© 2024 Dongru Zhang, Yunhe Wu, Changsheng Zuo, Fawang Yin, Shuo Liu.
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:
Zhang D, Wu Y, Zuo C, Yin F, Liu S (2024) First description of the female of Cyrtodactylus dianxiensis Liu & Rao, 2021, with extended diagnosis of this species (Squamata, Gekkonidae). Herpetozoa 37: 65-72. https://doi.org/10.3897/herpetozoa.37.e119492
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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.
bent-toed gecko, China, morphology, Yunnan
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 (
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 (
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.
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 (
The adult female specimen preserved in 75% ethanol was measured with digital calipers to the nearest 0.1 mm. Morphological terminology followed
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 (
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 (
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 (
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.
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 |
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.
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.
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.
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.
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.
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.
The female specimen agrees well with the original description (
The holotype of C. dianxiensis was collected at an altitude of 1170 m and the paratype was collected at an altitude of 1200 m (
Cyrtodactylus dianxiensis was previously known based on only two adult males and one juvenile (
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).