Print
A new species of the genus Takydromus (Squamata, Lacertidae) from southwestern Guangdong, China
expand article infoJian Wang, Zhi-Tong Lyu, Chen-Yu Yang, Yu-Long Li§, Ying-Yong Wang§
‡ Sun Yat-sen University, Guangzhou, China
§ Sun Yat-sen Universit, Guangzhou, China
Open Access

Abstract

A new species, Takydromus yunkaiensis J. Wang, Lyu, & Y.Y. Wang, sp. nov. is described based on a series of specimens collected from the Yunkaishan Nature Reserve located in the southern Yunkai Mountains, western Guangdong Province, China. The new species is a sister taxon to T. intermedius with a genetic divergence of 8.0–8.5% in the mitochondrial cytochrome b gene, and differs from all known congeners by a combination of the following morphological characters: (1) body size moderate, SVL 37.8–56.0 mm in males, 42.6–60.8 mm in females; (2) dorsal ground color brown; ventral surface green to yellow-green, but light blue-green on chin and throat, posteriorly green in adult males; (3) dorsolateral lines paired, strikingly yellowish-white bordered by black above and below, invisible or indistinct in juveniles and adult females; (4) flanks of body blackish brown with light brown marks in adult males; (5) presence of four pairs of chin-shields; (6) four supraoculars on each side; (7) presence of a row of supracilary granules that separate supracilaries from supraoculars; (8) two postnasals; (9) enlarged dorsal scales in six longitudinal rows on trunk of body, with strong keel; (10) enlarged ventral scales in six longitudinal rows, strongly keeled in males, smooth but outermost rows weakly keeled in females; (11) enlarged and keeled lateral scales in a row above ventrals; (12) femoral pores 2–3 on each side; (13) subdigital lamellae 20–23 under the fourth finger, 23–30 under the fourth toe; and (14) the first 2–3 subdigital lamellae under the fourth toe divided. The discovery of Takydromus yunkaiensis sp. nov. brings the total number of species of this genus to 24, of which nine occur in mainland China.

Keywords

grass lizard, southern China, species diversity, taxonomy, Takydromus yunkaiensis sp. nov.

Introduction

The Asian grass lizard genus Takydromus Daudin, 1802 currently contains 23 recognized species, widely distributed in the East Asian islands (Ryukyu Archipelago, Taiwan) and recorded from the Russian far east, extending southward across the Chinese mainland, Indochina, northeastern India, Borneo, the Natuna Islands, Sumatra, Bangka, and Java (Wang et al. 2017; Uetz et al. 2019). Eight species are recorded in mainland China: T. albomaculosus Wang, Gong, Liu & Wang, 2017, T. amurensis Peters, 1881, T. intermedius Stejneger, 1924, T. kuehnei van Denburgh, 1909, T. septentrionalis Günther, 1864, T. sexlineatus Daudin, 1802, T. sylvaticus Pope, 1928, and T. wolteri Fischer, 1885 (Zhao et al. 1999; Cai et al. 2011; Wang et al. 2017). In addition, T. formosanus Boulenger, 1894, T. hsuehshanensis Lin & Cheng, 1981, T. luyeanus Lue & Lin, 2008, T. sauteri Van Dengurgh, 1909, T. stejnegeri van Denburgh, 1912, and T. viridipunctatus Lue & Lin, 2008 are endemic to Taiwan Island; T. dorsalis Stejneger, 1904, T. smaragdinus Boulenger, 1887, T. tachydromoides Schlegel, 1838, and T. toyamai Takeda & Ota, 1996 are known only from Japan. Finally, T. hani Chou, Nguyen & Pauwels, 2001 and T. madaensis Bobrov, 2013 are only recorded from Vietnam while T. khasiensis Boulenger, 1917 and T. sikkimensis Günther, 1888 are only recorded from India.

Previous studies have revealed the very high biodiversity level of the genus Takydromus in southern China, for which the species diversity is just below that of Taiwan Island (Lue and Lin 2008; Wang et al. 2017). During repeated field surveys in the Yunkai Mountains, located in southwestern Guangdong Province (Fig. 1), a number of lacertid specimens were collected that could be assigned to the genus Takydromus by a combination of diagnostic characters defined by Arnold et al. (2007) and Zhao et al. (1999): (1) body slender with an extra-long tail, tail length usually more than two times larger than snout-vent length, (2) dorsal scales enlarged and keeled, ventral scales enlarged, keeled or smooth, (3) scales on flanks small and granular, (4) lateral teeth tricuspid, (5) temporal scales usually keeled, (6) 0–5 femoral pores on each side. Close examination of the external morphology and subsequent molecular analyses revealed that these specimens from Yunkaishan Nature Reserve, Guangdong Province, represented a distinct taxon. They are described below as a new species.

Figure 1. 

The type locality of Takydromus yunkaiensis sp. nov., Yunkaishan Nature Reserve.

Materials and methods

Sampling

Samples sequenced for molecular analyses were obtained from two specimens of the undescribed Takydromus species from Yunkaishan Nature Reserve, Guangdong Province; the paratype specimen (SYS r001292) of T. albomaculosus; two specimens of T. amurensis; four specimens of T. intermedius including a topotypic specimen (SYS r001602) from Mt. Emei, Sichuan; four specimens of T. kuehnei; three specimens of T. septentrionalis; two specimens of T. sexlineatus; one specimen of T. sylvaticus; and two specimens of T. wolteri, all freshly collected from China. Additional 14 sequences of T. dorsalis, T. formosanus, T. hsuehshanensis, T. sauteri, T. smaragdinus, T. stejnegeri, T. tachydromoides, and T. toyamai were obtained from GenBank and three sequences of Eremias persica Blanford, 1875, E. strauchi Kessler, 1878, and E. velox (Pallas, 1771) also from GenBank were used as the out-groups. Details of samples sequenced for mitochondrial cytochrome b gene and their associated GenBank accession numbers are listed in Table 1.

Localities, voucher information, and GenBank accession numbers (mitochondrial cytochrome b gene) for all specimens/sequences used in this study.

ID Species Locality Voucher GenBank Number
1 Takydromus yunkaiensis sp. nov. China: Guangdong: Gaozhou: Xianrendong Scenic Area SYS r001513 MN239954
2 Takydromus yunkaiensis sp. nov. China: Guangdong: Gaozhou: Xianrendong Scenic Area SYS r001514 MN239955
3 Takydromus albomaculosus China: Guangdong: Ruyuan: Tianjingshan Forestry Station SYS r001292 (paratype) MF631870
4 Takydromus amurensis China: Liaoning: Fushun: Nanzamu County: Mt. Langya SYS r001890 MN239956
5 Takydromus amurensis China: Liaoning: Fushun: Nanzamu County: Mt. Langya SYS r001891 MN239957
6 Takydromus dorsalis Japan AY248460
7 Takydromus dorsalis Japan AY248461
8 Takydromus formosanus China: Taiwan Island AY248458
9 Takydromus formosanus China: Taiwan Island AY248459
10 Takydromus hsuehshanensis China: Taiwan Island AY248482
11 Takydromus hsuehshanensis China: Taiwan Island AY248483
12 Takydromus intermedius China: Sichuan: Mt. Emei (type locality) SYS r001602 (topotype) MN239958
13 Takydromus intermedius China: Guizhou: Libo: Maolan Nature Reserve SYS r000856 MN239959
14 Takydromus intermedius China: Guangxi: Hechi: Jiuwanshan Nature Reserve SYS r001553 MN239960
15 Takydromus intermedius China: Guangxi: Hechi: Cenwanglaoshan Nature Reserve SYS r001741 MN239961
16 Takydromus kuehnei China: Taiwan Island: Xinzhu County SYS r001797 MN239962
17 Takydromus kuehnei China: Taiwan Island: Taipei SYS r001798 MN239963
18 Takydromus kuehnei China: Jiangxi: Longnan: Jiulianshan Nature Reserve SYS r001268 MN239964
19 Takydromus kuehnei China: Zhaoqing: Fengkai: Heishiding Nature Reserve SYS r001338 MN239965
20 Takydromus sauteri China: Taiwan Island AY248465
21 Takydromus sauteri China: Taiwan Island AY248466
22 Takydromus septentrionalis China: Zhejiang: Lishui: Jingning County: Makeng Village SYS r000912 MN239966
23 Takydromus septentrionalis China: Zhejiang: Wenzhou: Chashan County SYSr001886 MN239967
24 Takydromus septentrionalis China: Jiangsu: Xiaotangshan SYSr001882 MN239968
25 Takydromus sexlineatus China: Zhaoqing: Fengkai: Heishiding Nature Reserve SYS r001335 MN239969
26 Takydromus sexlineatus China: Zhaoqing: Fengkai: Heishiding Nature Reserve SYS r001336 MN239970
27 Takydromus smaragdinus Japan: Akashima LC066078
28 Takydromus stejnegeri China: Taiwan Island AY248473
29 Takydromus stejnegeri China: Taiwan Island AY248474
30 Takydromus sylvaticus China: Fujian: Shaowu: Longhu Forestry Station SYS r001276 MN239971
31 Takydromus tachydromoides Japan: Nagasaki – (topotype) LC066067
32 Takydromus tachydromoides Japan: Nagasaki – (topotype) LC066068
33 Takydromus toyamai Japan AY248480
34 Takydromus wolteri China: Anhui: Mt. Langya SYSr001888 MN239972
35 Takydromus wolteri China: Anhui: Mt. Langya SYSr001889 MN239973
36 Eremias persica Iran FJ416286
37 Eremias strauchi Iran: Yengeje: Neyshabur-Khorasan Razavi KJ468076
38 Eremias velox Iran: Jajarm area-Northern Khorasan KJ468081

All specimens were fixed in 10 % buffered formalin and later transferred to 70% ethanol for preservation, and deposited at the Museum of Biology, Sun Yat-sen University (SYS); liver tissue samples were separately preserved in 95% ethanol for molecular studies.

DNA Extraction, PCR and sequencing

DNA was extracted from liver tissue using a standard phenol-chloroform extraction protocol (Sambrook et al. 1989). The fragment of mitochondrial cytochrome b gene was PCR amplified and sequenced using the primers L14919 5’-AACCACCGTTGTTATTCAACT-3’ and H16064 5’-CTTTGGTTTACAAGAACAATGCTTTA-3’ (Burbrink et al. 2000). PCR amplifications were performed in a 20 µL reaction volume with the following cycling conditions: an initial denaturing step at 95 °C for five min.; 35 cycles of denaturing at 95 °C for 40 s, annealing at 53 °C for 40 s, and extending at 72 °C for one min., and a final extending step of 72 °C for 10 min. PCR products were purified with spin columns. The purified products were sequenced with both forward and reverse primers using a BigDye Terminator Cycle Sequencing Kit (ThermoFisher Scientific, Waltham, MA) according to the manufacturer’s guidelines. The products were sequenced on an ABI Prism 3730 automated DNA sequencer (Shanghai Majorbio Bio-pharm Technology Co., Ltd).

Phylogenetic analyses

Sequence alignments were first conducted using Clustal X 2.0 (Thompson et al. 1997), with default parameters and the alignment being checked and manually revised, if necessary. The data were tested in jmodeltest v2.1.2 with Akaike and Bayesian information criteria, resulting in the best-fitting nucleotide substitution models of GTR + I + G. Phylogenetic relationships were reconstructed using Maximum Likelihood (ML) as implemented in RaxmlGUI 1.3 (Silvestro and Michalak 2012), and Bayesian Inference (BI) using MrBayes 3.12 (Ronquist et al. 2012). For ML analysis, we used the rapid-bootstrapping algorithm (1000 replicates) with the thorough ML search option. Bootstrap values less than 60 were collapsed. For BI analysis, two independent runs with four Markov Chain Monte Carlo simulations were performed for ten million iterations and sampled every 1000th iteration. The first 25 % of samples were discarded as burn-in. Convergence of the Markov Chain Monte Carlo simulations was assessed using Tracer v.1.4 (http://tree.bio.ed.ac.uk/software/tracer/). We also calculated pairwise sequence divergence based on uncorrected p-distance implemented in MEGA 6 (Tamura et al. 2013).

Morphometrics

Measurements of all specimens were taken with a digital caliper to the nearest 0.1 mm. Abbreviations of measurements followed the convention of Lue and Lin (2008):

ALL arm-leg length (from insertion of the forelimb to insertion of hindlimb);

HH head height (measured at the highest point);

HL head length (from tip of snout to anterior margin of ear opening with claw);

HLL hindlimb length (from groin to tip of fourth toe);

HW head width (measured at the broadest point);

LTL length of fourth toe excluding claw;

RUL radius-ulna length;

SAL snout-arm length (from tip of snout to anterior insertion margin of forelimb);

SEL snout-eye length (from tip of snout to anterior margin of eye);

SKL skull length (from tip of snout to posterior margin of occipital);

SVL snout-vent length (from tip of snout to anterior margin of cloaca);

TaL tail length (from cloaca to tip of tail);

TFL tibia-fibula length.

Moreover, 20 external morphological characters were examined from the specimens listed in Appendix 1. Modified abbreviations of these characters followed Arnold (1997), Lue and Lin (2008) and Wang et al. (2017) as follows:

ADSR anterior dorsal scale rows, distinctly enlarged and keeled scales on anterior dorsum, counted transversely at position of forelimbs;

CS chin-shields;

CSR caudal scale rows, counted around the tail in the position of the 11th to 13th subcaudal scales;

ESRF enlarged and keeled lateral scales in longitudinal row(s) above ventrals on lower flanks;

FP femoral pores;

IFL infralabials;

LDSN dorsal scale numbers, counted longitudinally from posterior margin of occipital to posterior margin of hind limbs;

MBSR scales in a transverse row at mid-body, including ventrals;

MDSR transverse dorsal scale rows at mid-body;

PDSR posterior dorsal scale rows, counted transversely at the position of hind limbs;

SDLF-IV subdigital lamellae under fourth finger;

SDLT-IV subdigital lamellae under fourth toe;

SPC supraciliary;

SPL supralabials;

SPO supraocular;

SPT supratemporals;

SSRF small flat and granular scales in a transverse row on flank at mid-body;

TSRF enlarged and keeled scale rows above ventrals on flank;

VN ventral scale numbers, counted longitudinally from the posterior margin of collars to the anterior margin of precloacal scales;

VR ventral scale rows, counted transversely at mid-body.

Comparative morphological data were obtained from the literature for Takydromus albomaculosus (Wang et al. 2017), T. hani (Chou et al. 2001), T. viridipunctatus, and T. luyeanus (Lue and Lin 2008), T. sikkimensis (Bhupathy et al. 2009), T. madaensis (Bobrov 2013), T. sylvaticus (Pope 1928, 1929; Yang and Wang 2010), T. smaragdinus and T. toyamai (Takeda and Ota 1996); T. kuehnei (van Denburgh 1909; Arnold 1997; Norval et al. 2012; Wang et al. 2017), T. intermedius (Stejneger 1924; Wang et al. 2017), T. amurensis, T. dorsalis, T. formosanus, T. hsuehshanensis, T. sauteri, T. stejnegeri, and T. tachydromoides (Takeda and Ota 1996; Lue and Lin 2008), T. sexlineatus, T. wolteri, T. septentrionalis, and T. khasiensis (Arnold 1997; Zhao et al. 1999). All examined specimens are listed in Appendix I.

Results

BI and ML phylogenetic trees were constructed based on DNA sequences of the mitochondrial cytochrome b gene with a total length of 1074 -bp. The two analyses resulted in essentially identical topologies and are integrated in Figure 2, in which the Bayesian posterior probabilities (BPP) > 0.75 and the bootstrap supports (BS) for ML analysis > 60 were retained. The specimens from Yunkaishan Nature Reserve grouped in a strongly supported clade (BPP 1.00 and BS 98) with small divergence (p-distance 0.2 %), forming the sister taxon to Takydromus intermedius with strong support (BPP 1.00 and BS 92) and significant divergences (p-distance 8.0–8.5 %), and then to T. dorsalis, T. sylvaticus, and T. albomaculosus (BPP 1.00 and BS 95), indicating that the population from Yunkaishan Nature Reserve represents a separate evolutionary lineage.

Figure 2. 

Bayesian Inference and Maximum Likelihood phylogenies. The Bayesian posterior probabilities (BPP) > 0.75 and the bootstrap supports for Maximum Likelihood analysis (BS) > 60 were retained.

Morphologically, the unnamed specimens can be clearly distinguished from its congeners by the following characters: (1) body size moderate, SVL 37.8–56.0 mm in males, 42.6–60.8 mm in females; (2) dorsal ground color brown; ventral surface green to yellow-green, but light blue-green on chin and throat, posteriorly green in adult males; (3) dorsolateral lines paired, strikingly yellowish-white bordered by black above and below, invisible or indistinct in juveniles and adult females; (4) flanks of body blackish brown with light brown marks in adult males; (5) the presence of four pairs of chin-shields; (6) four supraoculars on each side; (7) presence of a row of supracilary granules that separate supracilaries from supraoculars; (8) two postnasals; (9) enlarged dorsal scales in six longitudinal rows on trunk of body, with strong keel; (10) enlarged ventral scales in six longitudinal rows, strongly keeled in males, smooth but outermost rows weakly keeled in females; (11) enlarged and keeled lateral scales in a row above ventrals; (12) femoral pores 2–3 on each side; (13) subdigital lamellae 20–23 under the fourth finger, 23–30 under the fourth toe; and (14) the first 2–3 subdigital lamellae under the fourth toe divided.

Based on the comprehensive evidence of molecular and morphological analyses, we hereby describe these specimens from Yunkaishan Nature Reserve as a new species, Takydromus yunkaiensis sp. nov. Now, the genus Takydromus contains 24 species, nine of which are recorded from mainland China.

Takydromus yunkaiensis J. Wang, Lyu & Y.Y. Wang, sp. nov.

Fig. 3

Material

Holotype. SYS r001580, adult male, collected by Jian Wang on 16 August 2016 from Dawuling Forestry Station (22°16'32.90"N, 111°11'42.87"E; 1500 m a.s.l.), Yunkaishan National Nature Reserve, Xinyi City, Guangdong Province, China.

Paratypes. Three adult males, collected by Ying-Yong Wang, Jian Wang, Zhi-Tong Lyu and Zhao-Chi Zeng: SYS r001439, 1442 on 15 and 16 April 2016, SYS r001684 on 17 April 2017, all from Dawuling Forestry Station (1200–1500 m a.s.l.). Six adult females: SYS r001513 and SYS r001514 collected by Jian Wang on 9 July 2016 from Xianrendong Scenic Area (22°165'45.99"N, 111°13'16.35"E; 1000 m a.s.l.), Yunkaishan National Nature Reserve, Xinyi City, Guangdong Province; SYS r001434 collected by Jian Wang and Zhi-Tong Lyu on 14 April 2016, SYS r001507 collected by Jian Wang on 28 June 2016, SYS r001581 collected by Jian Wang on 16 August 2016, and SYS r001901 collected by Jian Wang and Hong-Hui Chen on 10 April 2018, all from Dawuling Forestry Station (1200–1500 m a.s.l.).

Figure 3. 

Morphological features of the adult male holotype SYS r001580 of Takydromus yunkaiensis sp. nov. in life. A Habitus view and close-up of flank B close-up of dorsal body C close-up of ventral body D–G close-up of head scales.

Etymology

The specific epithet, yunkaiensis, is in reference to the type locality of the new species. We propose the standard name “Yunkai grass lizard” and the Chinese name “Yun Kai Cao Xi (云开草蜥)”.

Diagnosis

(1) body size moderate, SVL 37.8–56.0 mm in males, 42.6–60.8 mm in females; (2) dorsal ground color brown; ventral surface green to yellow-green, but light blue-green on ventral head and neck, posteriorly green in adult males; (3) dorsolateral lines paired, strikingly yellowish-white bordered by black above and below, invisible or indistinct i n juveniles and adult females; (4) flanks of body blackish brown with light brown marks in adult males; (5) the presence of four pairs of chin-shields; (6) four supraoculars on each side; (7) presence of a row of supracilary granules that separate supracilaries from supraoculars; (8) two postnasals; (9) enlarged dorsal scales with strong keel in six longitudinal rows on trunk of body; (10) enlarged ventral scales in six longitudinal rows, strongly keeled in males, smooth but outermost rows weakly keeled in females; (11) enlarged and keeled lateral scales in a row above ventrals; (12) femoral pores 2–3 on each side; (13) subdigital lamellae 20–23 under the fourth finger, 23–30 under the fourth toe; and (14) the first 2–3 subdigital lamellae under the fourth toe divided.

Comparisons

In this study we only compare the new species with the other 22 recognized species, excluding Takydromus haughtonianus, which is currently an uncertain species and poorly known (Jerdan 1870; Arnold 1997). Measurements, body proportions, and scale counts of the new species are listed in Tables 3 and 4; comparative data of the new species and nine other recognized members of the genus Takydromus occurring on the Chinese mainland are listed in Tables 5 and 6.

Uncorrected p-distances among Takydromus species based on mitochondrial cytochrome b gene. Takydromus yunkaiensis sp. nov. (1)–(2); T. albomaculosus (3); T. amurensis (4)–(5); T. dorsalis (6)–(7); T. formosanus (8)–(9); T. hsuehshanensis (10)–(11); T. intermedius (12)–(15); T. kuehnei (16)–(19); T. sauteri (20)–(21); T. septentrionalis (22)–(24); T. sexlineatus (25)–(26); T. smaragdinus (27); T. stejnegeri (28)–(29); T. sylvaticus (30); T. tachydromoides (31)–(32); T. toyamai (33); T. wolteri (34)–(35).

SpecNo. (1)–(2) (3) (4)–(5) (6)–(7) (8)–(9) (10)–(11) (12)–(15) (16)–(19) (20)–(21) (22)–(24) (25)–(26) (27) (28)–(29) (30) (31)–(32) (33) (34)–(35)
(1)–(2) 0.2
(3) 16.5–17.0
(4)–(5) 21.9–22.3 24.5 0
(6)–(7) 13.5–14.4 18.1–18.2 23.2–23.6 1.2
(8)–(9) 25.9–26.1 25.9–26.1 23.0–23.5 24.5–25.2 1.3
(10)–(11) 21.9–22.2 23.4–23.6 23.6–23.8 22.6–23.3 15.8–16.2 0.1
(12)–(15) 8.0–8.5 16.3–16.9 20.3–21.3 13.9–15.0 23.1–23.9 20.8–21.6 0.4–1.6
(16)–(19) 21.5–22.8 24.2–25.2 25.6–25.9 23.0–24.2 22.3–23.6 20.2–22.5 24.2–25.4 0–5.4
(20)–(21) 23.5 25.5–25.6 22.3–22.5 22.3–22.7 23.2–23.6 23.2–23.6 22.1–23.1 23.9–25.3 0.4
(22)–(24) 21.1–22.1 24.3–24.7 20.2–22.2 24.2–24.8 17.8–20.0 15.8–17.5 20.2–21.3 21.4–23.4 24.1–24.8 0–0.6
(25)–(26) 21.0–21.8 25.9 27.6 23.3 25.7–25.9 25.5–25.7 23.4–23.8 24.2–25.0 26.6–26.7 22.1–23.9 0
(27) 18.9–19.7 21.7 19.8 19.3–19.7 24.2–24.6 21.2–21.4 19.0–19.4 23.4–24.0 21.0–21.2 22.9–23.7 26.3 -
(28)–(29) 23.6–24.3 24.4–24.6 20.1–20.3 23.6–24.3 17.2–18.2 16.2–16.8 21.7–23.0 20.1–21.8 25.2–25.6 10.3–12.4 24.3–25.2 21.2–21.7 2.2
(30) 13.3–13.4 18.5 21.5 13.4–13.8 25.6–25.8 20.1–20.3 12.5–12.8 22.2–23.8 22.7–22.9 19.9–20.6 23.7 18.7 20.6–21.2 -
(31)–(32) 21.8–22.2 23.7 22.5 23.0–23.3 26.6–26.8 23.9–24.1 22.6–23.0 23.7–24.4 21.7 22.4–22.7 29.2 21.6 24.0–24.2 22.2 0
(33) 21.7–21.9 25.2 19.6 21.5–21.9 19.1–19.9 15.7–15.9 20.9–21.9 22.2–23.2 22.7–23.2 9.5–10.9 21.6 21.9 11.7–12.6 20.5 22.8 /
(34)–(35) 22.8–23.3 23.9–24.1 21.6 23.1–24.2 10.6–11.1 12.7–12.9 22.8–23.5 23.3–24.1 21.8–22.6 17.2–17.8 25.9–26.0 21.9–22.1 17.4–17.6 21.7–21.8 24.3–24.4 18.1 0.1

Measurements and body proportions of type series of Takydromus yunkaiensis sp. nov.

Voucher SYS r SYS r SYS r SYS r SYS r SYS r SYS r SYS r SYS r SYS r
Number 001439 001442 001580 001684 001434 001507 001513 001514 001581 001901
Sex
SVL 37.8 42.1 43.0 56.0 42.6 60.8 52.5 47.6 49.9 51.9
TaL 100.4 112.0 111.3 155.0 111.5 155.3 143.5 75.7 (broken tail) 148.3 156.7
HL 9.6 10.5 11.1 14.8 10.4 16.3 13.8 13.0 12.0 13.8
HW 6.4 6.6 6.7 8.1 6.5 7.6 6.9 6.3 7.4 7.7
HH 4.6 5.1 5.2 6.0 5.3 6.3 5.3 5.1 5.3 5.5
SKL 10.1 10.9 11.0 14.5 11.0 15.6 12.0 11.3 12.5 13.3
SEL 4.6 4.8 5.0 6.4 4.3 6.8 5.4 5.4 5.6 6.3
ALL 17.7 19.8 19.7 28.3 21.3 30.9 25.0 23.4 25.3 26.6
SAL 14.6 16.6 18.0 21.6 17.7 23.9 20.0 18.5 19.3 20.0
RUL 4.1 5.6 5.6 6.5 4.6 6.9 5.9 5.8 6.5 6.0
HLL 20.0 25.2 25.4 28.3 21.7 29.4 28.1 24.4 28.4 26.9
TFL 4.9 6.3 6.3 7.5 5.3 8.4 7.9 5.9 8.0 7.4
LTL 5.3 7.7 7.9 10.0 7.2 10.2 9.3 8.4 9.2 9.4
TaL/SVL 2.66 2.66 2.59 2.77 2.62 2.55 2.73 1.59 2.97 3.02
HL/SVL 0.25 0.25 0.26 0.26 0.24 0.27 0.26 0.27 0.24 0.27
HL/HW 1.50 1.58 1.66 1.83 1.61 2.14 2.01 2.06 1.63 1.79
SKL/HL 1.05 1.04 0.99 0.98 1.06 0.96 0.87 0.87 1.04 0.96
SEL/HL 0.48 0.46 0.45 0.43 0.41 0.42 0.39 0.41 0.47 0.46
ALL/SVL 0.47 0.47 0.46 0.51 0.50 0.51 0.48 0.49 0.51 0.51
SAL/SVL 0.39 0.39 0.42 0.39 0.42 0.39 0.38 0.39 0.39 0.39
RUL/SVL 0.11 0.13 0.13 0.12 0.11 0.11 0.11 0.12 0.13 0.12
HLL/SVL 0.53 0.60 0.59 0.51 0.51 0.48 0.53 0.51 0.57 0.52
TFL/SVL 0.13 0.15 0.15 0.13 0.12 0.14 0.15 0.12 0.16 0.14
LTL/SVL 0.14 0.18 0.18 0.18 0.17 0.17 0.18 0.18 0.19 0.18
HLL/ALL 1.13 1.27 1.29 1.00 1.02 0.95 0.13 1.04 1.12 1.01

Scale counts of type series of Takydromus yunkaiensis sp. nov.

Voucher SYS r SYS r SYS r SYS r SYS r SYS r SYS r SYS r SYS r SYS r
No. 001434 001439 001442 001507 001513 001514 001580 001581 001684 001901
CS 4 4 4 4 4 4 4 4 4 4
FP 3 3 3 3 2 2 3 3 3 3
SPL 7 7 7 7 7 7 6 7/6 7 6
IFL 7 7 7 7 7 6 6 6/7 7 6
SPO 4 4 4 4 4 4 4 4 4 4
SPC 4 4 4 3 4 4 4/2 4 4 4
SPT 3/4 4/3 4/3 4 4/3 4/3 3 3/4 3/4 3
ADSR 9 10 9 9 9 9 9 9 9 9
PDSR 7 7 7 7 7 7 7 7 7 7
MDSR 8 8 8 8 8 8 7 8 7 7
LDSN 47 49 51 47 49 47 47 51 47 47
MBSR 40 44 42 40 46 41 42 41 46 41
SSRF 13/13 17/13 12/16 13/13 16/16 13/14 14/15 14/13 17/16 14/14
VR 6 6 6 6 6 6 6 6 6 6
VN 25 26 25 25 26 25 24 26 25 27
ESRF 1 1 1 1 1 1 1 1 1 1
CSR 13 13 12 10 12 13 12 13 13 12
SDLF-4 20 20 22 23 23 21 21 23 20 21
SDLT-4 27 23 26 28 30 27 26 28 27 25

Selected body proportions of Takydromus yunkaiensis sp. nov. (data of the female paratype SYS r001514 with a broken tail is not included), and its morphologically most similar species T. intermedius and T. kuehnei; data obtained from Wang et al. (2017).

T. spp. yunkaiensis sp. nov. intermedius kuehnei
Sex ♂ (N = 4) ♀ (N = 5) ♂ (N = 1) ♀ (N = 3) ♂ (N = 2) ♀ (N = 1)
TaL/SVL 2.59–2.77 2.55–3.02 2.22 2.54–3.25 3.07–3.08 -
(2.67±0.07) (2.78±0.21) (2.90±0.36) (3.08)
HL/SVL 0.25–0.26 0.24–0.27 0.25 0.22–0.25 0.24–0.25 0.22
(0.26±0.01) (0.26±0.01) (0.24±0.02) (0.25)
HL/HW 1.50–1.83 1.61–2.14 1.80 1.68–1.72 1.69–1.83 1.84
(1.64±1.14) (1.87±0.23) (1.70±0.02) (1.76)
SKL/HL 0.98–1.05 0.87–1.06 1.04 1.02–1.05 1.02–1.03 1.05
(1.01±0.04) (0.96±0.08) (1.04±0.02) (1.02)
SEL/HL 0.43–0.48 0.39–0.47 0.46 0.46–0.48 0.45–0.46 0.49
(0.46±0.02) (0.43±0.03) (0.47±0.01) (0.46)
ALL/SVL 0.46–0.51 0.48–0.51 0.55 0.53–0.55 0.52–0.53 0.58
(0.48±0.02) (0.50±0.01) (0.54±0.01) (0.53)
SAL/SVL 0.39–0.42 0.38–0.42 0.39 0.37–0.42 0.38–0.39 0.36
(0.40±0.02) (0.39±0.01) (0.40±0.03) (0.38)
RUL/SVL 0.11–0.13 0.11–0.13 0.16 0.12–0.14 0.13–0.14 0.11
(0.12±0.01) (0.12±0.01) (0.14±0.01) (0.13)
HLL/SVL 0.51–0.60 0.48–0.57 0.55 0.48–0.53 0.54–0.57 0.49
(0.56±0.05) (0.52±0.03) (0.51±0.03) (0.56)
TFL/SVL 0.13–0.15 0.12–0.16 0.17 0.14–0.16 0.14–0.16 0.13
(0.14±0.01) (0.14±0.01) (0.15±0.01) (0.15)
LTL/SVL 0.14–0.18 0.17–0.19 0.19 0.16–0.20 0.20 0.19
(0.17±0.02) (0.18±0.01) (0.19±0.02)
HLL/ALL 1.00–1.29 0.95–1.13 0.99 0.89–1.01 1.03–1.09 0.85
(1.17±0.14) (1.04±0.07) (0.96±0.06) (1.06)

Selected scale counts of the nine species of the genus Takydromus recorded from the Chinese mainland, modified from Wang et al. (2017); differences are marked in bold.

Takydromus yunkaiensis sp. nov. albomaculosus amurensis wolteri septentrionalis sexlineatus intermedius kuehnei sylvaticus
Species (N = 10) (N = 2) (N = 2) (N = 1) (N = 25) (N = 5) (N = 8) (N = 5) (N = 3)
CS 4 4 4 4 3 3 4–5 4 (rare 3*) 4
FP 2–3 3–4 4 1 1 1 2–3 3–5 3
SPL 6–7 6–7 5–7 7 5–8 5–6 6–7 6–7 5–7
IFL 6–7 6–7 6–7 6–7 5–6 4–5 5–7 5–6 5–7
SPO 4 3 (rare 4#) 4 4 4 (rare 3) 3 4 4 4
SPC 4 (rarely 2, 3) 4–6 4 4 3–5 3 4–5 4 4–5
SPT 3–4 3 2–3 3 1–4 2–3 2–5 3–4 2–4
ADSR 9–10 6 7–8 9 6–8 6 6–8 5–7 /
PDSR 7 6 6–7 7 4–6 4 6 6 9–10
MDSR 7–8 7 7–8 8 5–6 4 7–8 6–7 11–14
LDSN 47–51 52–53 46 56 37–46 34–35 36–46 42–47 67–81
MBSR 40–46 42–43 33–38 36 34–42 28–33 40–44 39–44 45–47
SSRF 12–17 13–14 5–9 10 7–11 6–8 12–15 13–16 13
VR 6 6 8 8 8 8 6 6 6
VN 24–27 23–26 27 30 25–29 26–27 21–24 27–29 26–29
ESRF 1 1 1–3 3 2–3 2–3 1 0–1 0
CSR 10–13 12 16–18 16 12–14 14 12 12–13 12
SDLF-4 20–23 23–24 18–19 17 18–22 13–16 20–21 18–20 21–22
SDLT-4 23–30 29–30 24–25 22–23 23–28 19 26–27 23–24 27–28

In our phylogenetic tree, Takydromus yunkaiensis sp. nov. is a sister taxon to T. intermedius, from which it differs by having two postnasals (only one in T. intermedius), having a pair of strikingly yellowish-white dorsolateral lines in adult males (vs. always absent or indistinct in T. intermedius), flanks of body blackish brown with light brown spots in adult males (vs. pure brown without spots in T. intermedius), ADSR 9–10, PDSR 7 (vs. ADSR 6–8, PDSR 6 in T. intermedius).

Morphologically, Takydromus yunkaiensis sp. nov. is most similar to T. kuehnei (Fig. 4). The new species can be distinguished from T. kuehnei by having a pair of strikingly yellowish-white dorsolateral lines in adult males (vs. absent or dorsolateral stripes blurred, pale brown only present in old individuals in T. kuehnei); surface of ventrals green (vs. surface of ventrals white or light yellow in T. kuehnei), ADSR 9–10, PDSR 7 (vs. ADSR 5–7, PDSR 6 in T. kuehnei); TaL/SVL 2.59–2.77 in males (vs. tail relatively longer, TaL/SVL 3.07–3.08 in T. kuehnei); relatively shorter trunk (arm-leg length), ALL/SVL 0.46–0.51 in males, 0.48–0.51 in females (vs. relatively larger arm-leg length, ALL/SVL 0.52–0.53 in males and 0.58 in female of T. kuehnei).

Figure 4. 

Sexual dimorphism in color patterns. A Male paratype of Takydromus yunkaiensis sp. nov. (SYS r001439) B female paratype of T. yunkaiensis sp. nov. (SYS r001901) C male topotype of T. intermedius (SYS r001601) from Mt. Emei, China D female topotype of T. intermedius (SYS r001602) from Mt. Emei, China E male T. kuehnei (SYS r001268) from Jiulianshan Nature Reserve, China F female topotype of T. kuehnei (SYS r001798) from Taiwan Island, China.

From the remaining six Takydromus species which occur on mainland China (T. albomaculosus, T. amurensis, T. wolteri, T. septentrionalis, T. sexlineatus, and T. sylvaticus), Takydromus yunkaiensis sp. nov. can be distinguished by having dense mottles on flanks in males (vs. several particular white round spots on the flanks in T. albomaculosus; white ocellus bordered by black edges in males of T. sexlineatus); dorsum brown (vs. dorsum green in T. sylvaticus); four pairs of chin-shields (vs. three in T. septentrionalis and T. sexlineatus); two or three pairs of femoral pores (vs. only one in T. wolteri, T. septentrionalis and T. sexlineatus; four in T. amurensis); IFL 6–7 (vs. 4–5 in T. sexlineatus); SPO 4 (vs. three in T. sexlineatus); ADSR 9–10 (vs. six in T. albomaculosus and T. sexlineatus; 7–8 in T. amurensis; 6–8 in T. septentrionalis); PDSR 7 (vs. six in T. albomaculosus; 4–6 in T. septentrionalis; four in T. sexlineatus; 9–10 in T. sylvaticus); MDSR 7–8 (vs. 5–6 in T. septentrionalis; 4 in T. sexlineatus; 11–14 in T. sylvaticus); LDSN 47–51 (vs. 56 in T. wolteri; 67–81 in T. sylvaticus); ESRF 1 (vs. three in T. wolteri; 2–3 in T. septentrionalis and T. sexlineatus; none in T. sylvaticus).

Takydromus yunkaiensis sp. nov. differs from T. formosanus, T. hsuehshanensis, T. luyeanus, T. sauteri, T. stejnegeri, and T. viridipunctatus, which only occurred in Taiwan Island of China, by having four pairs of chin-shields (vs. three pairs in T. formosanus, T. viridipunctatus, T. luyeanus, T. hsuehshanensis and T. stejnegeri); FP 2–3 pairs (vs. only one in T. sauteri and T. stejnegeri); VR 6 (vs. eight in T. formosanus, T. stejnegeri , T. viridipunctatus and T. luyeanus); ventrals keeled (vs. ventrals smooth in T. hsuehshanensis); mottles on flanks in males (vs. absent in males of T. formosanus, T. sauteri and T. stejnegeri); surface of ventrals green (vs. surface of ventrals white in T. formosanus, T. hsuehshanensis, and T. sauteri); rostral and nostril separated (vs. rostral touching nostril in T. sauteri); dorsum brown (vs. dorsum green in T. sauteri).

Takydromus yunkaiensis sp. nov. differs from T. dorsalis, T. smaragdinus, T. tachydromoides, and T. toyamai, which only occur in Japan, by having a brown dorsum (vs. green dorsum in T. dorsalis, T. smaragdinus, and T. toyamai); dorsal scales large, in longitudinal rows (vs. dorsal scales small, not in obvious longitudinal rows in T. dorsalis); FP 2–3 pairs (vs. only one in T. smaragdinus and T. toyamai); ventrals keeled (vs. smooth in T. tachydromoides); VR 6 (vs. 8 in T. tachydromoides and T. toyamai); CS 4 pairs (vs. 3 in T. smaragdinus and T. toyamai).

Takydromus yunkaiensis sp. nov. differs from the remaining four members, T. hani and T. madaensis from Vietnam, T. khasiensis and T. sikkimensis from India, by having the dorsum brown (vs. dorsum green in T. hani); VR 6 (vs. 8 in T. hani and T. khasiensis; VR 12 in T. sikkimensis); CS 4 pairs (vs. 3 in T. khasiensis and T. sikkimensis); FP 2–3 pairs (vs. FP 6–8 in T. hani); loreals 2 (vs. 3 in T. madaensis); SPO 4 (vs. 3 in T. madaensis); SDLT-4 23–30 (vs. SDLT-4 17 in T. madaensis).

Description of holotype

Adult male. Body size slightly small, SVL 43.0 mm; trunk of body short, ALL 19.7 mm, 46 % of SVL; head slightly long, HL 11.1, HW 6.7 mm, HH 5.2 mm, HL 26 % of SVL; skull length larger than head length, SKL 11.9 mm; snout moderately long, SEL in 5.0 mm, SEL 45 % of HL. Rostral large, pentagonal, visible in dorsal view, in contact with the first supralabials posteriorly on both sides, and supranasals dorsolaterally; nostril surrounded by a supranasal, two postnasals and the first supralabial on each side; one supranasal on each side, large, in contact with each other dorso-medially, separating rostral from frontonasal, and in contact with the upper postnasal posteriorly, not in contact with the anterior loreal; postnasals two, both in contact with the anterior loreal posteriorly, the upper one in contact with supranasal dorsolaterally, with frontonasal dorsally, the lower one in contact with the first supralabial ventrally; supralabials six on each side, the fifth one largest, under the eye; two loreals on each side, anterior one smaller than posterior one; posterior loreal in contact with anteriormost supraocular and anteriormost supraciliary scale posteriorly; four supraoculars on each side, the posteriormost one much smaller than others; supraciliaries four on left side, the second one longest; supraciliaries two on right side, the first one longest; supracilary granules arranged in a row, separated supracilaries from supraoculars; frontonasal large, smooth, hexagonal, separated from frontal by a pair of prefrontals; prefrontals two, weakly keeled, in contact with each other medially, with frontal and anterior two supraoculars posteriorly, with loreals laterally, respectively; a single frontal hexagonal, weakly keeled, in contact with second and third supraoculars laterally, with frontoparietals posteriorly; frontoparietals two, pentagonal, in contact with each other medially, with parietal and interparietal posteriorly, respectively; interparietal diamond, surrounded by two frontoparietals, two parietals and the single occipital; parietal pit located in the central of interparietal, distinctly visible; parietals two, large, weakly keeled, slightly in contact with each other medially; a single occipital between two parietals; temporal scales granular, slightly keeled; supratemporals three on each side, keeled, anteriormost one largest, longer than total length of posterior two; mental large, semielliptical; infralabials six on each side; four pairs of chin-shields, anterior two pairs in contact with each other medially, posterior two pairs separated from each other by gular scales; following gular scales gradually increasing in size, keeled, and become imbricated; enlarged, strongly keeled median gular scales extending anteriorly to the line joined posterior edges of ears; collars clear, composed of scales in ten rows pointed backwards, and forming a free serration; enlarged, imbricated dorsal scales on body with strong keel oriented posteriorly that form continuous ridges, extending anteriorly beyond forelimbs on to the nape, in nine rows in position of forelimbs, seven rows in position of hindlimbs; seven rows at mid-body, including a much smaller and discontinuous central row; longitudinal dorsal scales (LDSN) 47; ventrals in six rows, imbricate, strongly keeled and pointed posteriorly; enlarged and keeled lateral scales in a row above ventrals; longitudinal ventral scales (VN) 24; small flat and granular scales in a transverse row on flank at mid-body (SSRF) 14 on left side and 15 on right side, including a row of scales (enlarged and keeled, shorter than ventrals) adjoining the ventrals; four rows of scales on lower flanks reduced, flattened, keeled; nine rows of small granular scales on upper flanks on left side and ten on right side; a discontinuous row of scales adjoining outermost dorsal scale row reduced, flattened, keeled; a total of 42 scales (MBSR) in a transverse row in mid-body region; a single precloacal entire, enlarged, surrounded by eight continuous moderately sized scales anteriorly and laterally; three femoral pores on each side.

Forelimbs moderately long, RUL 5.6 mm, 13% of SVL; scales on anterior and dorsal surfaces of upper arm enlarged, keeled, rhomboid, imbricate, in seven rows; scales on ventral surface of upper arm granular, homogeneous in size; scales on upper insertion of upper arm granular; scales on dorsal surface of forearm keeled, heterogeneous in size, extending to wrist; dorsal scales on hand slightly keeled; scales on palm granular; dorsal scales on fingers in a row, smooth; subdigital lamellae under fingers I–V respectively (left/right) 9/9 (3 entire + 1 divided + 1 entire), 12/12 (6 entire + 5 divided + 1 entire), 16/16 (10 entire + 5 divided + 1 entire), 22/22 (15 entire + 6 divided + 1 entire), 13/13 (6 entire + 6 decided + 1 entire); relative lengths of adpressed fingers I < V < II < III < IV; hindlimbs slender and long, fourth toe reaching the posterior edge of insertion of upper arm when hindlimb adpressed along the side of the body; HLL 25.4 mm, 59% of SVL, 129% of ALL; TFL 6.3 mm, 15% of SVL; LTL 7.9 mm, 18% of SVL; three rows of large smooth scales running beneath thigh with traces of a fourth row; two rows of enlarged keeled scales and one rows of small keeled scales on dorsal surface of thigh; granular scales homogeneous in size on rear of thigh; internal tibial scale of row one formed by enlarged and smooth tibial scale; dorsal tibial scale flat, keeled, heterogeneous in size, extending to dorsal surface of foot; scales on sole of the foot granular; dorsal scales on toes in a row, smooth; subdigital lamellae under toes I–V respectively (left/right) 9/9 (2 entire + 6 divided + 1 entire), 13/14 (7 entire + 5/6 divided + 1 entire), 18/21 (11 entire + 6 divided + 1 entire), 26/26 (2 divided + 17 entire + 6 divided + 1 entire), 18/18 (2 divided + 7 entire + 7 divided + 1 entire); basal two subdigital lamellae of toe IV and V divided; relative lengths of adpressed toes I < II < V < III < IV.

Tail original, TaL 111.3 mm, TaL/SVL ratio 259%, SVL/TaL ratio 39 %, with strongly keeled scales in 15 rows at base (fifth subcaudal scale), in 13 rows in position of the 13th to 15th subcaudal scales (CSR); paired vertebral series of large scales on tail extending on to hind body.

Coloration of holotype in life

Dorsal surface of head, body, limbs, and tail bright brown, with a pair of strikingly yellowish-white dorsolateral lines bordered by black above and below, each beginning from the posterior margin of the most last supratemporal, running along outermost dorsal scale row, posteriorly extending to the forepart of the tail; flanks of body blackish brown with light brown marks; a pair of orange ventrolateral lines beginning from axilla, running along lower part of flanks, posteriorly extending to the groin; labial series, mental, chin-shields, granular scales on throat, collars light blue-green, posteriorly yellowish green from chest, venter, until to subcaudal region; ventral surface of limbs brown, tinged with green.

Coloration of holotype in preservative

Dorsal surface of head, body, limbs and tail brown; labial series, mental, chin-shields, granular scales on throat, ventral surface of body and tail pale blue; mottles on flanks blurry, color of mottles on flanks faded; ventral surface of limbs beige; dorsolateral stripes greyish white with black-brown edges at the inner sides; color of ventrolateral stripes faded, greyish white.

Variations and sexual dimorphism

Measurements, body proportions, and scale counts of the type series of Takydromus yunkaiensis sp. nov. are listed in Tables 2 and 4.

In the holotype SYS r001580, there are four supraciliaries on left side and two on right side, the first one longest on right side, the second supraciliary longest on left side (vs. four supraciliaries on each side, and the second one longest in the paratypes SYS r001439, 1440, 1442, 1513, 1514, 1581, 1684, 1901; three supraciliaries on each sides, and the second one longest in the paratype SYS r001507); prefrontal in contact with the anterior two supraoculars posteriorly in the holotype (vs. prefrontal only in contact with the first supraocular posteriorly on the right side of the paratype SYS r001439); three pairs of femoral pores in the holotype (vs. only two pairs present in the paratypes SYS r001513, 1514); tail relatively longer in two of the female paratypes, TaL/SVL 2.97 in SYS r001581 and 3.02 in SYS r001901 (vs. TaL/SVL 2.59 in the holotype).

Takydromus yunkaiensis sp. nov. exhibits noticeable sexual dimorphism:

(1) enlarged ventral scales strongly keeled in males (vs. smooth but outermost rows weakly keeled in females);

(2) dorsolateral lines strikingly yellowish-white bordered by black above and below (vs. invisible or indistinct in adult females, also in juveniles);

(3) a pair of orange ventrolateral lines present on lower flanks (vs. invisible in females, also in juveniles);

(4) flanks of body blackish brown with light brown marks in adult males (vs. absent in females).

Distribution and habits

Currently, Takydromus yunkaiensis sp. nov. is known only from its type locality of Dawuling Forestry Station, adjacent Xianrendong Scenic Area located in the southern Yunkai Mountains in western Guangdong Province, China (Fig. 1).

The diurnal species was found to be very active in daytime and rapidly escapes when disturbed, and is usually observed resting on fern leaves at night. The surrounding environment was covered by well-preserved montane evergreen broad-leaved forest or mixed forest (Fig. 5) at altitudes of 900–1600 m.

Figure 5. 

Habitat of Takydromus yunkaiensis sp. nov. in Yunkaishan Nature Reserve.

Discussion

The description of Takydromus yunkaiensis sp. nov. brings the total number of species of this genus to 24, nine of which occur in mainland China. As noted, six species were recorded from Guangdong Province: T. albomaculosus, T. kuehnei, T. septentrionalis, T. sexlineatus, T. sylvaticus, and Takydromus yunkaiensis sp. nov., which further support the very high biodiversity level of the genus in southern China (Zhao et al. 1999; Lue and Lin 2008; Yang and Wang 2010; Wang et al. 2017).

Most of the early descriptions of Takydromus species only listed relatively limited diagnostic characteristics, resulting in considerable challenges in field identification of the species, and causing ambiguities in taxonomy. Moreover, in recent years, a number of new or cryptic species were discovered and described from southern mainland China and Taiwan Island (Lin et al. 2002; Lue and Lin 2008; Wang et al. 2017). These discoveries confirm the substantially underestimated species diversity within the tropical genus Takydromus, and more field research is required to increase the knowledge of the diversity.

Located in the western Guangdong Province, the Yunkai Mountains have gradually been recognized for its unique biodiversity. During herpetological surveys during the last several years, we have discovered a number of new species including some cryptic species, as well as providing new regional records of amphibians and reptiles (Yang et al. 2011; Lyu et al. 2018; Wang et al. 2018a; Wang et al. 2018b; Lyu et al. 2019), suggesting that future herpetological exploration will likely continue to yield new discoveries from the region.

Acknowledgements

We would like to thank Can-Rong Lin from Guangdong Lingnan Vocational and Technical College, Chun-Peng Guo from Chengdu Institute of Biology, the Chinese Academy of Sciences, Hai-Long He, Run-Lin Li, Hong-Hui Chen, Yao Li, Chao-Yu Lin, Zu-Yao Liu, and Zhao-Chi Zeng from the Museum of Biology, Sun Yat-sen University, for their help in the field work. This work was supported by the Scientific Expedition of Biological Resources of Yunkai Mountains in Guangdong Province (No. 2018B030320001) to Wen-Bo Liao and the Specimen Platform of Ministry of Science and Technology, P.R. China, teaching specimens sub-platform (No.2005DKA21403JK) to Ying-Yong Wang.

References

  • Arnold EN, Arribas O, Carranza S (2007) Systematics of the Palaearctic and Oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. Zootaxa 1430(1): 1–86. https://doi.org/10.11646/zootaxa.1430.1.1
  • Bhupathy S, Chettri B, Bauer AM (2009) Rediscovery and revalidation of Takydromus sikkimensis (Günther, 1888) (Squamata: Lacertidae) from Sikkim, India. Journal of Herpetology 43(2): 267–274. https://doi.org/10.1670/08-136R1.1
  • Bobrov VV (2013) A new species of grass lizard (Reptilia, Sauria, Lacertidae, Takydromus) from southern Vietnam. Current Studies in Herpetology 13: 97–100. [in Russian]
  • Cai Y, Yan J, Xu XF, Lin ZH, Ji X (2011) Mitochondrial DNA phylogeography reveals a west-east division of the northern grass lizard (Takydromus septentrionalis) endemic to China. Journal of Zoological Systematics and Evolutionary Research 50: 137–144. https://doi.org/10.1111/j.1439-0469.2012.00655.x
  • Chou WH, Nguyen TQ, Pauwels OSG (2001) A new species of Takydromus (Reptilia: Lacertidae) from Vietnam. Herpetologica 57: 497–508.
  • Jerdon TC (1870) Notes on Indian Herpetology. Proceedings of the Asiatic Society of Bengal 1870: 66–85. [T. haughtonianus, new species, pp. 72]
  • Lin SM, Chen CLA, Lue KY (2002) Molecular phylogeny and biogeography of the Grass Lizards Genus Takydromus (Reptilia: Lacertidae) of East Asia. Molecular Phylogenetics and Evolution 22(2): 276–288. https://doi.org/10.1006/mpev.2001.1059
  • Lyu ZT, Huang LS, Wang J, Li YQ, Chen HH, Qi S, Wang YY (2019) Description of two cryptic species of the Amolops ricketti group (Anura, Ranidae) from southeastern China. ZooKeys 812: 133–156. https://doi.org/10.3897/zookeys.812.29956
  • Lyu ZT, Wu J, Wang J, Sung YH, Liu ZY, Zeng ZC, Wang X, Li YY, Wang YY (2018) A new species of Amolops (Anura: Ranidae) from southwestern Guangdong, China. Zootaxa 4418(6): 562–576. https://doi.org/10.11646/zootaxa.4418.6.4
  • Norval G, Mao JJ, Goldberg SR (2012) Filling the gaps: additional notes on the reproduction of the Kühne’s grass lizard (Takydromus kuehnei van Denburgh, 1909; Squamata: Lacertidae) from southwestern Taiwan. Herpetological Conservation and Biology 7(3): 383–390. https://doi.org/10.2984/65.3.383
  • Pope CH (1928) Seven new reptiles from Fukien Province, China. American Museum Novitates 320: 1–6.
  • Pope CH (1929) Notes on reptiles from Fukien and other Chinese provinces. Bulletin of the American Museum of Natural History 58: 335–487.
  • Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. https://doi.org/10.1093/sysbio/sys029
  • Sambrook J, Fritsch EF, Maniatis T (1989) Molecular Cloning: A Laboratory Manual. Cold Spring Harbor Laboratory Press, New York, 125 pp.
  • Stejneger LH (1924) Herpetological novelties from China. Occasional Papers of the Boston Society of Natural History 5: 119–121.
  • Takeda N, Ota H (1996) Description of a new species of Takydromus from the Ryukyu Archipelago, Japan, and a taxonomic redefinition of T. smaragdinus Boulenger 1887 (Reptilia: Lacertidae). Herpetologica 52: 77–88.
  • Tamura K, Stecher G, Peterson D, Filipski A, Kumar S (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular biology and evolution 30(12): 2725–2729. https://doi.org/10.1093/molbev/mst197
  • Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG (1997) The CLUSTAL_X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 22: 4673–4680. https://doi.org/10.1093/nar/25.24.4876
  • Van Denburgh J (1909) New and previously unrecorded species of reptiles and amphibians from the island of Formosa. Proceedings of the California Academy of Sciences 4: 49–56.
  • Wang J, Yang JH, Li Yao, Lyu ZT, Zeng ZC, Liu ZY, Ye YH, Wang YY (2018a) Morphology and molecular genetics reveal two new Leptobrachella species in southern China (Anura, Megophryidae). ZooKeys 776: 105–137. https://doi.org/10.3897/zookeys.776.22925
  • Wang J, Zeng ZC, Lyu ZT, Liu ZY, Wang YY (2018b) Description of a new species of Gracixalus (Amphibia: Anura: Rhacophoridae) from Guangdong Province, southeastern China. Zootaxa 4420(2): 251–269. https://doi.org/10.11646/zootaxa.4420.2.7
  • Wang YY, Gong SP, Liu P, Wang X (2017) A new species of the genus Takydromus (Squamata: Lacertidae) from Tianjingshan Forestry Station, northern Guangdong, China. Zootaxa 4338(3): 441–458. https://doi.org/10.11646/zootaxa.4338.3.2
  • Yang JH, Wang YY (2010) Range extension of Takydromus sylvaticus (Pope, 1928) with notes on morphological variation and sexual dimorphism. Herpetology Notes 3: 279–283.
  • Yang JH, Wang YY, Zhang B, Lau MWN, Chow WH (2011) Revision of the diagnostic characters of Opisthotropis maculosa Stuart & Chuaynkern, 2007 with notes on its distribution and variation, and a key to the genus Opisthotropis (Squamata: Natricidae). Zootaxa 2785: 61–68. https://doi.org/10.11646/zootaxa.2785.1.4
  • Zhao EM, Zhao KT, Zhou KY (1999) Fauna Sinica, Reptilia. Vol. 2. Squamata. Science Press, Beijing, 394 pp.

Appendix 1

Examined specimens

Takydromus albomaculosus (N = 2): Chia: Guangdong Province: Tianjingshan Forest Station: SYS r001292, 1624.

Takydromus amurensis (N = 2): China: Heilongjiang Province: SYS r001635; Suifenhe City: SYS r001647.

Takydromus wolteri (N = 1): China: Heilongjiang Province: SYS r001636.

Takydromus septentrionalis (N = 25): China: Jiangxi Province: Mt. Sanqing: SYS r000179; Wuyuan County: Mt. Dazhang: SYS r000644, 653–655; Guixi City: Yangjifeng Nature Reserve: SYS r000115, 0133, 0135, 0147; Yanshan County: Wuyishan Nature Reserve: SYS r000642; Guangfeng County: Tongboshan Nature Reserve: SYS r000656, 0471, 0472, 0741, 0742, 0772; Jinggangshan City: Mt. Jinggang: SYS r000282, 1307; Zhejiang Province: Jingning County, Dongkeng: SYS r000912; Fujian Province: Wuyishan City: Sangang Village: SYS r000667, 0676, 0678; Guangdong Province: Ruyuan County: Tianjingshan Forestry Station: SYS r000929, 0930; unknown locality: SYS r000168.

Takydromus sexlineatus (N = 5): China: Guangdong Province: Fengkai County: Heishiding Nature Reserve: SYS r001335, 1336, 1337, 1552; Guangxi Zhuang Autonomous Region: Shangsi County: Shiwandashan Forest Park: SYS r000127.

Takydromus intermedius (N = 8): China: Sichuan Province: Mt. Emei: SYS r001601, 1602, CIB 3745, 3750; Guizhou Province: Libo County: Maolan Nature Reserve: SYS r000856; Hunan Province: Sangzhi County: Badagongshan Nature Reserve: SYS r001330, 1331; Guangxi Zhuang Autonomous Region: Hechi City: Jiuwanshan Nature Reserve: SYS r001553.

Takydromus sylvaticus (N = 3): China: Jiangxi Province: Guixi City: Yangjingfeng Nature Reserve: SYS r000159, 0184; Fujian Province: Shaowu City: Longhu Forestry Station: SYS r001276.

Takydromus kuehnei (N = 5): China: Jiangxi Province: Longnan County: Jiulianshan Nature Reserve: SYS r001268; Guangdong Province: Fengkai County: Heishiding Nature Reserve: SYS r000119, 0132, 1338; Renhua County: Huangshakeng Village: SYS r000206.