Research Article |
Corresponding author: Shuo Liu ( liushuo@mail.kiz.ac.cn ) Corresponding author: Dingqi Rao ( raodq@mail.kiz.ac.cn ) Academic editor: Thomas Ziegler
© 2022 Shuo Liu, Dingqi Rao.
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:
Liu S, Rao D (2022) A new species of Cyrtodactylus Gray, 1827 (Squamata, Gekkonidae) from southwestern Yunnan, China. ZooKeys 1084: 83-100. https://doi.org/10.3897/zookeys.1084.72868
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A new species of the Cyrtodactylus chauquangensis species group is described based on four specimens collected from the karst formations of Menglian County, Puer City, Yunnan Province, China. The new species can be separated from all other congeners by having a unique combination of morphological characters: a medium-sized body; ventrolateral folds present with interspersed small tubercles; seven precloacal pores in a continuous series in males, absent in females; enlarged femoral scales and femoral pores absent; two postcloacal tubercles on each side; and one or two rows of enlarged subcaudals. Genetically, the new species most closely related to C. wayakonei and the uncorrected sequence divergences of the ND2 gene and its flanking tRNAs between the new species and investigated congeners range from 7.2% to 18.4%.
Bent-toed gecko, Cyrtodactylus chauquangensis group, Menglian County, taxonomy
Cyrtodactylus Gray, 1827 is the most speciose and ecologically diverse gekkotan genus with more than 300 recognized species so far (
The Cyrtodactylus chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, 2007 species group, previously C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler, 2010 species group, is a karst ecotype species group, which is distributed in northern Indochina, ranging from northern Thailand and Laos to northwestern and central Vietnam, and to Yunnan Province in southern China (
During our fieldwork in southern Yunnan Province, China, in 2021, some specimens of Cyrtodactylus were collected from the karst formations of Menglian County, Puer City. Morphological and molecular phylogenetic analyses revealed that the new collection belonged to an unnamed species of the C. chauquangensis species group. We herein describe it as a new species.
Specimens were collected by hand. Photographs were taken to document the color pattern of specimens in life prior to their euthanization. Liver tissues were stored in 99% ethanol and specimens were preserved in 75% ethanol. Specimens were deposited at Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences (
Measurements were taken with digital calipers to the nearest 0.1 mm. Bilateral scale counts were given as left/right. The methodology of measurements and meristic counts followed
AG axilla to groin distance;
DTR dorsal tubercle rows, number of dorsal, longitudinal rows of tubercles at midbody between ventrolateral folds;
ED ear diameter, greatest diameter of ear;
EE eye orbit to ear distance, from posterior corner of eye orbit to anterior margin of ear opening;
EFS enlarged femoral scales, number of enlarged femoral scale beneath each thigh;
ForeaL forearm length, from base of palm to elbow;
FP femoral pores;
GSDT granular scales surrounding dorsal midbody tubercles;
HH maximum head height, from occiput to underside of jaws;
HL head length, from tip of snout to posterior margin of retroarticular of lower jaw;
HW maximum head width;
I postrostrals or internasals;
IFL infralabials;
IND internarial distance, measured between inner borders of nostrils;
IOD interorbital distance, measured across narrowest point of frontal bone;
LF4 subdigital lamellae under the fourth finger;
LT4 subdigital lamellae under the fourth toe;
ML mental length;
MW mental width;
OD greatest diameter of orbit;
PAT postcloacal tubercles, number of tubercles on each side of postcloacal region;
PM postmentals, scales bordering mental shield, except infralabials;
PP precloacal pores;
PVT paravertebral tubercles, counted in a single paravertebral row from the level of the forelimb insertions to the level of the hind limb insertion;
RH rostral height;
RW rostral width;
SE snout to eye distance, from tip of snout to anterior corner of eye orbit;
SL shank length, from the base of heel to the knee;
SPL supralabials;
SVL snout–vent length, from tip of snout to anterior margin of cloaca;
TaL tail length, from posterior margin of cloaca to tip of tail;
V longitudinal ventral scale rows, counted across the belly between the ventrolateral folds at midbody.
Morphological comparisons were based on the original descriptions of each species (
Molecular data were generated for three specimens collected from Menglian County, Puer City, Yunnan Province, China, and available sequences of the Cyrtodactylus chauquangensis species group were obtained from GenBank; the new sequences have been deposited on GenBank under the accessions OM296042–OM296044. Cyrtodactylus dattkyaikensis Grismer, Wood, Quah, Grismer, Thura, Oaks & Lin, 2020 and C. sinyineensis Grismer, Wood Jr, Thura, Zin, Quah, Murdoch, Grismer, Lin, Kyaw & Lwin, 2017 were used as the outgroups according to
Sequences were aligned using ClustalW (
The obtained sequence alignment is 1397 bp in length. The sequences of the three specimens collected from Menglian County, Yunnan, China, were nested within the Cyrtodactylus chauquangensis species group and represented a distinct clade sister to C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler, 2010 with strong support (Fig.
Mean uncorrected pairwise genetic distances (%) among the species of the Cyrtodactylus chauquangensis species group and outgroups based on the ND2 gene and its flanking tRNAs.
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 Cyrtodactylus menglianensis sp. nov. | |||||||||||||||||||||||
2 Cyrtodactylus auribalteatus | 12.2 | ||||||||||||||||||||||
3 Cyrtodactylus bichnganae | 18.4 | 18.3 | |||||||||||||||||||||
4 Cyrtodactylus bobrovi | 15.0 | 13.9 | 19.7 | ||||||||||||||||||||
5 Cyrtodactylus chauquangensis | 12.6 | 13.3 | 18.1 | 8.6 | |||||||||||||||||||
6 Cyrtodactylus cucphuongensis | 15.0 | 14.5 | 19.9 | 7.9 | 8.4 | ||||||||||||||||||
7 Cyrtodactylus doisuthep | 15.5 | 13.6 | 16.6 | 15.9 | 14.4 | 15.9 | |||||||||||||||||
8 Cyrtodactylus dumnuii | 11.4 | 11.9 | 17.0 | 13.7 | 12.3 | 14.4 | 14.3 | ||||||||||||||||
9 Cyrtodactylus erythrops | 13.9 | 13.8 | 16.7 | 14.8 | 13.5 | 14.7 | 11.0 | 13.4 | |||||||||||||||
10 Cyrtodactylus gulinqingensis | 14.3 | 13.2 | 18.1 | 13.8 | 14.0 | 14.0 | 14.1 | 12.9 | 13.8 | ||||||||||||||
11 Cyrtodactylus houaphanensis | 14.8 | 14.6 | 19.4 | 6.5 | 9.0 | 7.5 | 15.5 | 14.2 | 14.9 | 14.1 | |||||||||||||
12 Cyrtodactylus huongsonensis | 14.3 | 13.4 | 17.7 | 14.3 | 12.5 | 14.3 | 14.7 | 13.9 | 14.3 | 12.4 | 14.7 | ||||||||||||
13 Cyrtodactylus ngoiensis | 13.1 | 13.2 | 18.2 | 11.1 | 10.5 | 10.7 | 14.9 | 12.0 | 14.2 | 13.1 | 11.3 | 13.1 | |||||||||||
14 Cyrtodactylus otai | 15.2 | 14.6 | 19.1 | 3.6 | 9.1 | 8.4 | 16.3 | 15.6 | 16.4 | 15.6 | 6.8 | 14.7 | 12.2 | ||||||||||
15 Cyrtodactylus puhuensis | 14.2 | 13.4 | 18.9 | 5.7 | 8.0 | 7.1 | 14.7 | 12.9 | 14.2 | 13.6 | 2.8 | 13.8 | 10.5 | 6.2 | |||||||||
16 Cyrtodactylus soni | 13.4 | 13.0 | 18.2 | 14.3 | 13.6 | 14.4 | 14.2 | 13.1 | 14.2 | 13.0 | 15.3 | 6.7 | 14.0 | 14.7 | 14.2 | ||||||||
17 Cyrtodactylus sonlaensis | 17.2 | 16.2 | 19.4 | 17.5 | 16.8 | 18.1 | 16.2 | 16.8 | 17.3 | 14.8 | 18.0 | 15.0 | 16.2 | 17.7 | 18.0 | 15.2 | |||||||
18 Cyrtodactylus spelaeus | 14.6 | 13.9 | 18.3 | 10.0 | 9.2 | 10.4 | 15.7 | 13.4 | 15.0 | 13.9 | 10.4 | 14.3 | 11.1 | 11.3 | 9.1 | 14.3 | 17.7 | ||||||
19 Cyrtodactylus taybacensis | 16.8 | 15.6 | 9.3 | 17.1 | 15.5 | 17.3 | 15.7 | 14.9 | 16.4 | 16.3 | 17.3 | 16.1 | 16.6 | 18.3 | 16.7 | 15.6 | 18.9 | 16.5 | |||||
20 Cyrtodactylus vilaphongi | 14.1 | 13.4 | 17.8 | 8.1 | 7.3 | 8.2 | 14.2 | 13.3 | 14.0 | 13.5 | 8.2 | 14.2 | 9.5 | 9.1 | 7.0 | 13.5 | 16.9 | 9.6 | 16.5 | ||||
21 Cyrtodactylus wayakonei | 7.2 | 13.1 | 18.0 | 15.5 | 13.1 | 15.5 | 16.3 | 12.7 | 15.6 | 15.3 | 14.7 | 15.1 | 12.2 | 15.4 | 14.2 | 13.9 | 16.4 | 15.2 | 17.5 | 13.7 | |||
22 Cyrtodactylus zhenkangensis | 10.7 | 12.0 | 18.4 | 14.1 | 13.2 | 13.8 | 15.5 | 11.8 | 14.0 | 12.9 | 13.9 | 13.4 | 13.2 | 15.5 | 13.2 | 13.7 | 17.3 | 14.0 | 15.8 | 13.6 | 11.9 | ||
23 Cyrtodactylus dattkyaikensis | 18.3 | 17.2 | 21.4 | 18.8 | 18.0 | 19.4 | 18.0 | 17.0 | 16.8 | 16.8 | 19.9 | 17.2 | 18.0 | 21.5 | 18.8 | 17.9 | 21.8 | 19.3 | 18.4 | 17.8 | 19.9 | 17.9 | |
24 Cyrtodactylus sinyineensis | 18.8 | 17.8 | 18.9 | 18.7 | 17.5 | 18.1 | 18.2 | 17.0 | 18.7 | 18.9 | 18.9 | 19.1 | 17.5 | 20.9 | 18.1 | 18.7 | 21.9 | 18.9 | 17.9 | 18.1 | 19.4 | 18.4 | 13.1 |
Bayesian phylogram of the Cyrtodactylus chauquangensis species group inferred from the ND2 gene and its flanking tRNAs. Numbers before slashes indicate Bayesian posterior probabilities and numbers after slashes indicate the ML ultrafast bootstrap. The symbol “–” represents the value below 0.90/90.
Holotype. KIZ20210713, adult male, collected on 18 July 2021 by Shuo Liu from Menglian County, Puer City, Yunnan Province, China (22°20'11"N, 99°34'29"E, 980 m elevation).
Paratypes. KIZ20210714 and KIZ20210716, two adult females; KIZ20210715, adult male; all collected on 19 July 2021 by Shuo Liu from the same locality as the holotype.
The specific epithet refers to Menglian County, the locality where the new species was found. We propose “Menglian Bent-toed Gecko” for the common English name and “孟连裸趾虎” (Mèng Lián Luǒ Zhǐ Hǔ) for the common Chinese name of the new species.
Medium body size (SVL 67.7–78.1 mm in adults); ventrolateral folds present with interspersed small tubercles; seven precloacal pores in a continuous series in males, absent in females; femoral scales not enlarged; femoral pores absent; two postcloacal tubercles on each side; 17–22 lamellae under finger IV, 21–23 lamellae under toe IV; one or two rows of subcaudals enlarged; dark postocular streak and nuchal loop absent; six or seven dark irregular dorsal bands between limb insertions, most bands discontinuous.
Adult male, SVL 77.8 mm; head distinguished from neck, moderately long (HL/SVL 0.28), relatively widened (HW/HL 0.76), slightly depressed (HH/HL 0.45); two supranasals separated by one internasal; nares oval, surrounded by supranasal, rostral, first supralabial, and three postnasals; loreal region concave; snout long (SE/HL 0.43), round anteriorly, longer than diameter of orbit (OD/SE 0.63); snout scales small, round, granular, larger than those in frontal and parietal regions; eye large (OD/HL 0.27), pupils vertical; upper eyelid fringe with spinous scales; ear opening oval, small (ED/HL 0.09); rostral wider than high (RH/RW 0.58), medially divided dorsally by a suture, reaching to approximately half down rostral, in contact with first supralabial and nostrils laterally, and supranasals and internasal dorsally; mental triangular, narrower than rostral (MW/RW 0.82), slightly wider than high (ML/MW 0.94); two postmentals, enlarged, in contact posteriorly, bordered by mental anteromedially, first infralabial anterolaterally, two enlarged chin scales posterolaterally, and small chin scales posteriorly; 10/12 supralabials; 9/9 infralabials.
Body slender (AG/SVL 0.39), ventrolateral folds slightly developed with interspersed small tubercles; dorsal scales granular; dorsal tubercles round and weakly keeled, four or five times larger than the size of adjoining scales, conical, present on neck, back and tail base, each surrounded by 10 granular scales, in 19 irregular longitudinal rows at the midbody, 26 paravertebral tubercles; gular region with homogenous smooth scales; ventral scales smooth, larger than those of dorsum, round, subimbricate, largest posteriorly, in 29 longitudinal rows at midbody; precloacal groove absent; a patch of precloacal scales significantly enlarged; seven precloacal pores in a continuous series, the two on the edge round, the one in the middle pitted, others horizontally elongated; femoral scales not enlarged; femoral pore absent.
Fore and hind limbs moderately slender (ForeaL/SVL 0.17, SL/SVL 0.20); dorsal surface of forelimbs covered by a few weakly developed tubercles; interdigital webbing absent; lamellae under finger IV 22/21, under toe IV 23/22; relative length of fingers I < II < V < III < IV, relative length of toes I < II < III < V < IV.
Tail regenerated (TaL 60.8 mm); 2/2 postcloacal tubercles; dorsal tail base with tubercles; subcaudals smooth, enlarged but arranged irregularly.
Dorsal ground color brownish yellow; dorsal surface of head with irregular brown blotches with black edges, largest at occiput; nuchal loop absent; dorsum with many irregular brownish black blotches with black edges, forming eight transverse discontinuous bands faintly, one on the neck, one between hind limbs, and six between fore and hind limb insertions; dorsal surfaces of limbs with brown bands with black edges; a brown band with black edge on dorsal tail base, dorsal surface of regenerated tail greyish black; ventral surface of head, limbs, and body greyish white; ventral surface of regenerated tail grey; iris bronze.
The color pattern very much resembles that in life. Brownish yellow dorsal ground color turned to greyish white, the brown blotches and bands with black edges remained; ventral surface faded to pale white; iris became white.
The paratypes resemble the holotype except that the female KIZ20210714 has a longer regenerated tail, and the female KIZ20210716 has a longer original tail with one or two rows of subcaudals enlarged, and they both have no precloacal pores; the male KIZ20210715 has a smaller body size and much shorter regenerated tail; other morphometric and meristic differences are presented in Table
Measurements (mm) and meristic data for the type series of Cyrtodactylus menglianensis sp. nov. Abbreviations defined in Materials and methods. “*” represents regenerated tail and “#” represents original tail.
KIZ20210713 Holotype | KIZ20210714 Paratype | KIZ20210715 Paratype | KIZ20210716 Paratype | |
---|---|---|---|---|
Male | Female | Male | Female | |
SVL | 77.8 | 78.1 | 69.1 | 67.7 |
TaL | 60.8* | 62.9* | 33.2* | 70.6# |
HH | 9.7 | 9.7 | 8.3 | 8.7 |
HL | 21.7 | 21.8 | 19.4 | 19.2 |
HW | 16.4 | 15.8 | 14.8 | 14.5 |
OD | 5.9 | 6.9 | 5.1 | 5.3 |
SE | 9.3 | 9.2 | 8.2 | 8.3 |
EE | 6.3 | 5.9 | 5.6 | 5.6 |
IND | 3.1 | 3.1 | 2.9 | 2.6 |
IOD | 3.3 | 3.5 | 2.8 | 2.7 |
ED | 1.9 | 1.3 | 1.3 | 1.2 |
AG | 30.5 | 34.6 | 27.8 | 28.4 |
ForeaL | 13.0 | 12.9 | 11.4 | 10.6 |
SL | 15.7 | 15.6 | 14.0 | 13.1 |
RW | 3.8 | 3.4 | 2.9 | 3.0 |
RH | 2.2 | 2.1 | 1.9 | 1.8 |
MW | 3.1 | 3.2 | 2.9 | 3.8 |
ML | 2.9 | 2.6 | 2.6 | 2.0 |
SPL | 10/12 | 9/10 | 8/9 | 10/9 |
IFL | 9/9 | 9/9 | 9/8 | 9/7 |
I | 1 | 1 | 1 | 1 |
PM | 2 | 2 | 2 | 2 |
GSDT | 10 | 10 | 10 | 10 |
DTR | 19 | 21 | 20 | 18 |
PVT | 26 | 27 | 25 | 29 |
V | 29 | 26 | 26 | 28 |
EFS | 0 | 0 | 0 | 0 |
PP | 7 | 0 | 7 | 0 |
FP | 0 | 0 | 0 | 0 |
PAT | 2/2 | 2/2 | 2/2 | 2/2 |
LF4 | 22/21 | 19/18 | 18/17 | 21/22 |
LT4 | 23/22 | 22/22 | 22/21 | 23/23 |
All specimens were collected at night on large stones or cliffs of the karst formations in a park. The surrounding habitats was karst forest, there is a plank road and a river nearby.
Cyrtodactylus menglianensis sp. nov. is distinguishable from all other members of the C. chauquangensis species group by a unique combination of morphological characters. Cyrtodactylus menglianensis sp. nov. differs from C. auribalteatus Sumontha, Panitvong & Deein, 2010; C. bichnganae Ngo & Grismer, 2010; C. doisuthep Kunya, Panmongkol, Pauwels, Sumontha, Meewasana, Bunkhwamdi & Dangsri, 2014; C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya, 2010; C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya, 2009; C. gulinqingensis Liu, Li, Hou, Orlov & Ananjeva, 2021; C. hekouensis Zhang, Liu, Bernstein, Wang & Yuan, 2021; C. huongsonensis Luu, Nguyen, Do & Ziegler, 2011; C. ngoiensis Schneider, Luu, Sitthivong, Teynié, Le, Nguyen & Ziegler, 2020; C. soni Le, Nguyen, Le & Ziegler, 2016; C. sonlaensis Nguyen, Pham, Ziegler, Ngo & Le, 2017; and C. zhenkangensis Liu & Rao, 2021 by not having enlarged femoral scales and femoral pores (vs having enlarged femoral scales and femoral pores).
Cyrtodactylus menglianensis sp. nov. differs from C. puhuensis Nguyen, Yang, Le, Nguyen, Orlov, Hoang, Nguyen, Jin, Rao, Hoang, Che, Murphy & Zhang, 2014 and C. taybacensis Pham, Le, Ngo, Ziegler & Nguyen, 2019 by not having enlarged femoral scales (vs having enlarged femoral scales). In addition, C. menglianensis sp. nov. differs from C. puhuensis by having more precloacal pores in males (seven vs five) and differs from C. taybacensis by having fewer precloacal pores in males (seven vs 11–13).
Cyrtodactylus menglianensis sp. nov. differs from C. cucphuongensis Ngo & Chan, 2011 by having precloacal pores in males (vs not having precloacal pores in males).
Cyrtodactylus menglianensis sp. nov. differs from C. bobrovi Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, 2015; C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, 2007; C. houaphanensis Schneider, Luu, Sitthivong, Teynié, Le, Nguyen & Ziegler, 2020; C. otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, 2015; C. spelaeus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov, 2014; and C. vilaphongi Schneider, Nguyen, Le, Nophaseud, Bonkowski & Ziegler, 2014 by not having dark postocular streak and nuchal loop (vs having very obvious dark postocular streak and not obvious nuchal loop).
Cyrtodactylus menglianensis sp. nov. differs from C. martini Ngo, 2011 by not having enlarged femoral scales (vs having indistinctly enlarged femoral scales), having fewer longitudinal ventral scale rows (26–29 vs 39–43), having more precloacal pores in males (seven vs four), and having more white rings on the original tail (10 vs 7).
Cyrtodactylus menglianensis sp. nov. differs from C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler, 2010 by having fewer longitudinal ventral scale rows (26–29 vs 31–35), not having precloacal pores in females (vs having precloacal pores in females), and having more white rings on the original tail (10 vs 6).
The new species was found in a park just beside the county seat. There is a plank road along the limestone cliffs in the park, and there are many lamps on the limestone cliffs along the plank road (Fig.
There are still many karst landforms in southern Yunnan which have not been surveyed in detail. Additional cryptic new species of Cyrtodactylus are likely to be found in these areas. It is necessary to strengthen the protection of these karst landforms and to survey these areas.
We thank to our workmates for their help and advice. We also thank the editors and reviewers for their comments on the manuscript. This work was supported by Science-Technology Basic Condition Platform from the Ministry of Science and Technology of the People’s Republic of China (grant no. 2005DKA21402), National Natural Science Foundation Project: Investigation and Classificatory and Phylogenetic Studies on the Lizards of Gekkonidae of China (grant no. NSFC-31970404), and the project of Ministry of Ecology and Environment of China: Investigation and assessment of amphibians and reptiles in southern Yunnan.