Biodiversity Data Journal : Taxonomy & Inventories
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Taxonomy & Inventories
Confirmation of Tylototriton ziegleri Nishikawa, Matsui & Nguyen, 2013 in China, with discussion on the relationship between T. verrucosus Anderson, 1871 and T. panwaensis Grismer, Wood, Quah, Thura, Espinoza & Murdoch, 2019 (Caudata, Salamandridae)
expand article infoShuo Liu, Mian Hou§, Dingqi Rao|
‡ Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China
§ College of Continuing (Online) Education, Sichuan Normal University, Chengdu, China
| Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China
Open Access

Abstract

Background

The distribution of the Ziegler’s Crocodile Newt Tylototriton ziegleri Nishikawa, Matsui & Nguyen, 2013 in China has been controversial. This species was originally recorded uncertainly from Guangxi Autonomous Region, China. Subsequently, this species was recorded from Yunnan and Guangdong provinces, China. Thereafter, the record from Guangdong was denied and the record from Yunnan was questioned.

New information

Two specimens of Tylototriton Anderson, 1871 were collected from Wenshan Prefecture, Yunnan Province, China, in 2020. Phylogenetically, the sequences of these two specimens clustered with the sequences of T. ziegleri (including the holotype) from its type locality with strong support, and morphologically agree well with the original description of T. ziegleri. We confirm the record of T. ziegleri in China and present detailed collection site and morphological description of the specimens from China. In addition, we found that T. panwaensis Grismer, Wood, Quah, Thura, Espinoza & Murdoch, 2019 may be the synonym of T. verrucosus Anderson, 1871. We discussed the relationship between T. verrucosus and T. panwaensis.

Keywords

morphology, ND2, newt, phylogeny, Yunnan

Introduction

Tylototriton ziegleri Nishikawa, Matsui & Nguyen, 2013 is a member of the T. asperrimus species group, that was described from northern Vietnam (Nishikawa et al. 2013). This species was previously reported in China by Hernandez (2016) from Jingxi County, Guangxi Autonomous Region; however, he also mentioned that the population in Jingxi may not be the true T. ziegleri. Subsequently, this species was reported from Malipo County, Yunnan Province, based on only molecular data of a single specimen without morphological description and detailed collection information (Ye et al. 2017). Thereafter, this species was reported from Guangdong Province, China by Li et al. 2020. Lyu et al. (2021) found that the record of T. ziegleri from Guangdong by Li et al. (2020) actually represented a new species at that time and described it as T. sini Lyu, Wang, Zeng, Zhou, Qi, Wan & Wang, 2021. Therefore, Lyu et al. (2021) removed the record of T. ziegleri from the herpetofauna of Guangdong. In addition, Lyu et al. (2021) questioned the record of T. ziegleri from Yunnan.

Tylototriton verrucosus Anderson, 1871 is the type species of the genus Tylototriton. However, the current taxonomy of the true T. verrucosus is problematic because Anderson (1871) neither specified an exact type locality nor mentioned which specimens the original description of T. verrucosus was based on. Tylototriton panwaensis Grismer, Wood, Quah, Thura, Espinoza & Murdoch, 2019 is another species of Tylototriton closely resembling T. verrucosus and this species was described from Kachin State, north-eastern Myanmar, a location close to the border between Myanmar and western Yunnan of China.

During our field surveys in south-eastern Yunnan, China, in 2020, two specimens of Tylototriton were collected from Malipo County, Wenshan Prefecture. Detailed morphological comparisons and molecular analysis indicated that these specimens belong to T. ziegleri. Therefore, we confirm the distribution of T. ziegleri in China and provide a detailed description of these specimens herein. In addition, we collected some specimens of T. panwaensis in 2019 from western Yunnan, China, which is probably the type locality of T. verrucosus. Therefore, we also discuss the relationship between T. verrucosus and T. panwaensis herein.

Materials and methods

Sampling

Specimens were collected, humanely euthanised and then fixed in 75% ethanol for permanent storage. Photographs were taken to document the colour pattern in life prior to euthanasia. Liver tissue samples were preserved in 99% ethanol for molecular analysis. All specimens were deposited at Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences (KIZ).

Morphological characteristics

Morphological terminology followed Nishikawa et al. (2013). Measurements were taken with an electronic vernier caliper to the nearest 0.1 mm. Morphometric characters included: SVL: snout-vent length, from tip of snout to anterior tip of vent; HL: head length, from tip of snout to wrinkle of throat; HW: head width, measured at the angle anterior to the parotid gland; MXHW: maximum head width, measured at widest point; SL: snout length, from tip of snout to anterior tip of upper eyelid; LJL: lower jaw length, from tip of lower jaw to jaw angle; ENL: eyelid-nostril length, minimum distance between eyelid and nostril; IND: internarial distance, minimum distance between the external nares; IOD: interorbital distance, minimum distance between upper eyelids; UEW: upper eyelid width, greatest width of upper eyelid; UEL: upper eyelid length, greatest length of upper eyelid; OL: orbit length, maximum length of orbit; AGD: axilla-groin distance, minimum distance between axilla and groin; TRL: trunk length, from wrinkle of throat to anterior tip of vent; TAL: tail length, from anterior tip of vent to tail tip; VL: vent length, from anterior to posterior tip of vent; BTAW: basal tail width, tail width measured at root of tail; MTAW: medial tail width, tail width measured at middle; BTAH: basal tail height, tail height measured at base of tail; MXTAH: maximum tail height, tail height measured at highest point; MTAH: medial tail height, tail height measured at middle; FLL: fore-limb length, distance from axilla to tip of longest finger; HLL: hind-limb length, 2FL: second finger length; 3FL: third finger length; 3TL: third toe length; and 5TL: fifth toe length.

Molecular analysis

Genomic DNA was extracted from liver tissue samples preserved in 99% ethanol using a standard phenol-chloroform extraction protocol (Sambrook et al. 1989). A fragment of the NADH dehydrogenase subunit two (ND2) was amplified for the newly-collected specimens. Methods for amplification and sequencing of the DNA fragment are the same as Lyu et al. (2021). All new sequences were deposited in GenBank. Homologous and outgroup sequences were obtained from GenBank (Table 1).

Table 1.

Localities, voucher information and GenBank accession numbers for all samples used in this study.

Taxa

Locality

Voucher

Accession

Tylototriton anguliceps

Vietnam: Dien Bien: Muong Nhe

VNMN A20143

LC017832

Tylototriton anhuiensis

China: Anhui: Yuexi

AHU-13-EE-006

KY321413

Tylototriton asperrimus

China: Guangxi: Jinxiu

CIB GX20080714

KY800819

Tylototriton broadoridgus

China: Hunan: Sangzhi

CIB200084

KY800837

Tylototriton dabienicus

China: Anhui: Shangcheng

HNNU 1004-026

KY800869

Tylototriton daloushanensis

China: Guizhou: Suiyang

CIBWG200600019

KY800817

Tylototriton hainanensis

China: Hainan: Mt Diaoluo

CIB 20081048

KC147817

Tylototriton himalayanus

Nepal: Mechi: Illam

CIB 201406287

KT765210

Tylototriton kachinorum

Myanmar: Kachin: Indawgyi

ZMMU A5953

MK097273

Tylototriton kweichowensis

China: Guizhou: Shuicheng

SYS a004967

OK539843

Tylototriton liuyangensis

China: Hunan: Liuyang

CSUFT 20100108

KJ205598

Tylototriton lizhengchangi

China: Hunan: Yizhang

KUHE 42317

AB769533

Tylototriton maolanensis

China: Guizhou: Libo

CIBML20180427001

MK820699

Tylototriton ngarsuensis

Myanmar: Shan: Taunggyi

LSUHC 13763

MH836584

Tylototriton notialis

Laos: Khammouan: Boualapha

FMNH HERP 271120

HM462061

Tylototriton panhai

Thailand: Loei: Phu Luang WS

PL009

AB830736

Tylototriton panwaensis

Myanmar: Kachin: Myitkyina

CAS 245418

KT304279

Tylototriton panwaensis

China: Yunnan: Tengchong

KIZ 040240

ON159332

Tylototriton panwaensis

China: Yunnan: Tengchong

KIZ 040242

ON159331

Tylototriton panwaensis

China: Yunnan: Tengchong

KIZ 040243

ON159330

Tylototriton panwaensis

China: Yunnan: Tengchong

KIZ 040246

ON159329

Tylototriton panwaensis

China: Yunnan: Tengchong

KIZ 040247

ON159328

Tylototriton panwaensis

China: Yunnan: Yingjiang

KIZ20190701

ON159327

Tylototriton panwaensis

China: Yunnan: Yingjiang

KIZ20190702

ON159326

Tylototriton panwaensis

China: Yunnan: Yingjiang

KIZ20190703

ON159325

Tylototriton panwaensis

China: Yunnan: Yingjiang

KIZ20190704

ON159324

Tylototriton pasmansi

Vietnam: Phu Tho: Tan Son

IEBR 4467

MT210167

Tylototriton phukhaensis

Thailand: Nan: Doi Phu Kha NP

CUMZ A-7718

MN912574

Tylototriton podichthys

Laos: Luang Phabang: Phoukhoun

NCSM 77725

KT304295

Tylototriton pseudoverrucosus

China: Sichuan: Ningnan

CIB WCG2012003

KY800861

Tylototriton pulcherrima

China: Yunnan: Lüchun

CIB TY040

KY800890

Tylototriton shanjing

China: Yunnan: Jingdong

KIZ 201306102

KY800858

Tylototriton shanorum

Myanmar: Shan: Taunggyi

CAS 230933

AB922822

Tylototriton sini

China: Guangdong: Mt Yunkai

SYS a008354

OK539836

Tylototriton sparreboomi

Sin Ho, Lai Chau, Vietnam

IEBR 4476

MT210162

Tylototriton taliangensis

China: Sichuan: Liangshan

CAS 195126

DQ517853

Tylototriton thaiorum

Vietnam: Nghe An: Pu Hoat NR

ZMMU A-7577

MW883478

Tylototriton umphangensis

Thailand: Tak: Umphang WS

CUMZ-A-8243

OK092618

Tylototriton uyenoi

Thailand: Chiang Mai: Doi Suthep

KUHE 19147

AB830733

Tylototriton verrucosus

China: Yunnan: Longchuan

CIB TSHS1

KY800847

Tylototriton vietnamensis

Vietnam: Bac Giang: Son Dong

IEBR 3243

HM770088

Tylototriton wenxianensis

China: Gansu: Wenxian

CIB 20090527

KC147813

Tylototriton yangi

China: Yunnan: Pingbian

KUHE 42282

KY800887

Tylototriton ziegleri

Vietnam: Ha Giang: Quan Ba

VNMN 3390

AB769539

Tylototriton ziegleri

Vietnam: Ha Giang: Quan Ba

KUHE 55077

AB769540

Tylototriton ziegleri

Vietnam: Ha Giang: Quan Ba

KUHE 55078

AB769541

Tylototriton ziegleri

Vietnam: Ha Giang: Quan Ba

VNUH HG.081

KY800871

Tylototriton ziegleri

Vietnam: Ha Giang: Quan Ba

VNUH HG.082

KY800870

Tylototriton ziegleri

China: Yunnan: Malipo

KIZ20210504

ON159334

Tylototriton ziegleri

China: Yunnan: Malipo

KIZ20210505

ON159333

Tylototriton cf. ziegleri

Vietnam: Cao Bang: Bao Lac

VNMN 3389

AB769542

Tylototriton cf. ziegleri

Vietnam: Cao Bang: Quang Thanh

ROM 35330

DQ517856

Tylototriton cf. ziegleri

Vietnam: Cao Bang: Quang Thanh

ROM 35364

HM462056

Echinotriton chinhaiensis

China: Zhejiang: Ningbo

TP26195

EU880315

Echinotriton andersoni

Japan: Kagoshima: Tokunoshima

MVZ 232187

EU880314

Phylogenetic analyses

Sequences were aligned using MUSCLE 3.6 (Edgar 2004) with default parameters. Average genetic distances were calculated in MEGA 11 (Tamura et al. 2021) using the uncorrected p-distance model. The best substitution model GTR+F+I+G4 was selected using the Akaike Information Criterion (AIC) in ModelFinder (Kalyaanamoorthy et al. 2017). Bayesian Inference was performed in MrBayes 3.2.7 (Ronquist et al. 2012). Two runs were performed simultaneously with four Markov chains, the chains were run for 10,000,000 generations and sampled every 1,000 generations, the first 25% of the initial samples was discarded as burn-in after the standard deviation of split frequencies of the two runs was less than a value of 0.01. Maximum Likelihood analysis was performed in IQ-TREE 1.6.12 (Nguyen et al. 2015). The ultrafast bootstrap approximation algorithm was used via 1,000 bootstrap pseudoreplicates.

Taxon treatment

Tylototriton ziegleri Nishikawa, Matsui & Nguyen, 2013

Materials   Download as CSV 
  1. scientificName:
    Tylototriton ziegleri Nishikawa, Matsui & Nguyen, 2013
    ; family:
    Salamandridae
    ; country:
    China
    ; stateProvince:
    Yunnan
    ; locality:
    Zhongzhai Village, Xiajinchang Township, Malipo County, Wenshan Prefecture
    ; verbatimElevation:
    1750 m
    ; verbatimCoordinates:
    23°7′8″N 104°50′6″E
    ; eventRemarks:
    collected by Shuo Liu on 10 May 2020
    ; individualCount:
    1
    ; sex:
    male
    ; lifeStage:
    adult
    ; catalogNumber:
    KIZ20210504
    ; basisOfRecord:
    preserved specimen
  2. scientificName:
    Tylototriton ziegleri Nishikawa, Matsui & Nguyen, 2013
    ; family:
    Salamandridae
    ; country:
    China
    ; stateProvince:
    Yunnan
    ; locality:
    Zhongzhai Village, Xiajinchang Township, Malipo County, Wenshan Prefecture
    ; verbatimElevation:
    1750 m
    ; verbatimCoordinates:
    23°7′8″N 104°50′6″E
    ; eventRemarks:
    collected by Shuo Liu on 10 May 2020
    ; individualCount:
    1
    ; sex:
    male
    ; lifeStage:
    adult
    ; catalogNumber:
    KIZ20210505
    ; basisOfRecord:
    preserved specimen

Description the specimens from China: 

Morphometric data are provided in Table 2. Body moderately stout, medium in size (SVL 58.2–60.8 mm, TAL 68.0–69.7 mm). Head width almost equal to head length (HW/HL 0.96–1.05); head nearly hexagonal in shape in dorsal view, depressed, gently sloping in profile. Snout short, truncate, slightly beyond lower jaw. Nostril on anterior margin of snout, located notably closer to snout tip than to eye. Tongue oval, not notched distally, attached to mouth floor, but free laterally; vomerine tooth series in an inverted V-shape, converging anteriorly, but not reaching choanae. Labial fold absent; gular fold present, but weak; parotoids distinct, projecting posteriorly; costal folds absent. Dorsolateral supratemporal bony ridges on head protruding, beginning at the anterior corner of orbit continuing to anterior end of parotoid, posterior ends curved inside; mid-dorsal bony ridge on head short. Vertebral mid-dorsal ridge distinctly protruding, segmented, forming a row of tubercles, running from occiput region to the base of tail, separated from mid-dorsal bony ridge on head by a small gap. Rib nodules distinct, forming knob-like warts, relatively small, arranged in two longitudinal series on dorsolateral surfaces of dorsum from axilla to base of tail, counting 15–17 nodules on each side of body; rib nodules in the middle largest and decreasing anteriorly and posteriorly (Fig. 1).

Table 2.

Measurements (in mm) of the specimens of Tylototriton ziegleri from China.

KIZ20210504

KIZ20210505

SVL

58.2

60.8

HL

15.3

15.7

HW

16.0

15.1

MXHW

16.0

15.2

SL

5.8

6.3

LJL

12.7

12.6

ENL

3.7

3.8

IND

5.4

6.1

IOD

8.6

8.6

UEW

1.6

1.6

UEL

3.3

3.2

OL

4.2

4.2

AGD

28.9

31.6

TRL

43.2

45.8

TAL

69.7

68.0

VL

7.1

5.9

BTAW

5.5

6.0

MTAW

2.7

3.0

BTAH

7.4

7.2

MXTAH

7.4

7.3

MTAH

6.9

7.0

FLL

19.5

21.8

HLL

22.5

24.0

2FL

3.7

4.2

3FL

4.0

4.5

3TL

5.5

5.3

5TL

2.3

2.4

Figure 1.  

Tylototriton ziegleri from China in preservative. Dorsal view (left) and ventral view (right).

Limbs slender, tips of fore-limbs and hind-limbs overlapping when adpressed towards each other along body; fingers and toes free of webbing; relative finger lengths III >II > I ≥ IV, relative toe lengths III > IV > II > I ≥ V. Tail long, TAL/SVL 1.12–1.20; laterally compressed along entire length, tapering posteriorly, tip pointed, dorsal fin more distinct posteriorly, ventral ridge smooth.

Skin rough with fine granules, dense on dorsum and ventrum, but small and sparse on throat. Cloacal region slightly swollen, vent as a longitudinal slit, vent edges with numerous small transverse folds.

Colouration in life: 

Dorsum almost uniformly black; venter slightly lighter than dorsum; bony ridges on head and vertebral ridge black, rib nodules black or reddish, only tips or most regions of fingers and toes, vent and ventral ridge of tail orange (Fig. 2).

Figure 2.  

Tylototriton ziegleri from China in life: A-C the adult male KIZ20210504; D-F the adult male KIZ20210505.

Ecological notes: 

The specimens were collected in a small stream in the forest at night, the water in the stream was shallow, and both sides of the stream were covered with vegetation. No eggs or larvae were found.

Distribution: 

This species was recorded from Ha Giang and Cao Bang provinces, northern Vietnam; Malipo County, Yunnan Province and Jingxi County, Guangxi Autonomous Region, China (Fig. 3). However, the populations in Cao Bang and Jingxi may not represent the true Tylototriton ziegleri (see Discussion section).

Figure 3.  

Map showing the type locality of Tylototriton ziegleri in Ha Giang, Vietnam (1); the collection site of T. ziegleri in Malipo, Yunnan, China (2); the distributions of T. cf. ziegleri in Cao Bang, Vietnam (3 and 4); the distribution of T. cf. ziegleri in Jingxi, Guangxi, China (5); the type locality of T. verrucosus designated by Nussbaum et al. (1995) in Longchuan, Yunnan, China (6); the type locality of T. panwaensis in Kachin, Myanmar (7); the collection site of T. panwaensis in Tengchong, Yunnan, China (8); and the collection site of T. panwaensis in Yingjiang, Yunnan, China (9).

Analysis

Morphologically, the specimens from Malipo County, Wenshan Prefecture, Yunnan, China, agree well with the original description of Tylototriton ziegleri: medium body size; skin rough with fine granules; bony ridges on head distinct; vertebral ridge prominent and segmented; rib nodules prominent; tips of fore-limbs and hind-limbs overlapping when adpressed along body; tail thin; dorsum uniform blackish; finger and toe tips, vent and ventral ridge of tail orange.

The obtained sequence alignment is 1044 bp in length. BI and ML analyses showed basically consistent topology (Fig. 4). The two specimens collected from Malipo County, Wenshan Prefecture, Yunnan, China, were homogeneous and clustered with Tylototriton ziegleri (including the holotype) from Ha Giang Province, Vietnam, with strong support. The genetic distance (uncorrected p-distance) between the specimens from China and T. ziegleri (including the holotype) from Ha Giang, Vietnam was only 0.1% (Suppl. material 1).

Figure 4.  

Bayesian Inference tree, based on mitochondrial ND2 sequences. Numbers before slashes indicate Bayesian posterior probabilities (values below 0.9 are not shown) and numbers after slashes indicate bootstrap support for Maximum Likelihood analyses (values below 90 are not shown).

Combining the results of morphological and molecular analysis, we determined that the specimens from Malipo County, Wenshan Prefecture, Yunnan, China, belong to Tylototriton ziegleri.

Discussion

In the phylogenetic analyses in Nishikawa et al. (2013) and in this study, Tylototriton ziegleri formed two strongly-supported lineages, one (including the holotype) from Ha Giang Province, Vietnam and the other from Cao Bang Province, Vietnam and there is a relatively large genetic divergence (2.6%) between the two lineages, more than that (2.1%) between T. verrucosus and T. pulcherrima and close to that (2.7%) between T. taliangensis and T. pseudoverrucosus. Therefore, we regard the lineage containing the holotype as T. ziegleri sensu stricto and the other lineage as Tylototriton cf. ziegleri. The lineage of Tylototriton cf. ziegleri may, therefore, represent a cryptic new species of Tylototriton. Our newly-collected specimens from China clustered with the lineage of T. ziegleri sensu stricto and showed negligible genetic divergence (0.1%) with this lineage, confirming T. ziegleri sensu stricto. Ziegler et al. (2018) reported the longevity of T. ziegleri and extended the diagnosis of this species, based on specimens collected from Cao Bang, Vietnam; according to their result of molecular identification, these specimens from Cao Bang should be assigned to to Tylototriton cf. ziegleri rather than T. ziegleri sensu stricto. As for the population in Jingxi, Guangxi, China, record by Hernandez (2016), judging from the geographical location, we consider that this population should also belong to Tylototriton cf. ziegleri.

In the original description of Tylototriton verrucosus, Anderson (1871) described this species as uniform blackish-brown, but not orange-patterned. Usually, the uniform blackish-brown colour of these salamanders in life remains constant in preservative and the orange colour of these salamanders in life fades to light yellow or white, but the pattern remains distinct in preservative. However, according to Nussbaum et al. (1995) and Chanda et al. (2000), all the syntypes of T. verrucosus are orange-patterned. This has always made people wonder why the syntypes were inconsistent with the original description by Anderson (1871). Anderson (1871) did not specify a precise type locality, but merely mentioned some valleys in western Yunnan, including Nantin, Momien, Hotha, Ponsee and Nampoung. That is to say, any of these valleys may be the type locality of T. verrucosus. “Momien” generally refers to Tengchong City, Yunnan, China, nowadays; “Hotha” generally refers to Husa Township, Longchuan County, Yunnan, China, nowadays; “Ponsee” generally refers to Xueli Village, Yingjiang County, Yunnan, China, nowadays; “Nampoung” generally refers to Nabang Town, Yingjiang County, Yunnan, China, nowadays; and “Nantin” is uncertain. Nussbaum et al. (1995) considered that the syntypes of T. verrucosus are inconsistent with the original description of T. verrucosus and, as the specimens from Husa agree with the original description of T. verrucosus, they designated a neotype (KIZ 74II0061 VI.16) of T. verrucosus from Gongwa Village, Longchuan County, Yunnan Province, China. However, Fei et al. (2006) questioned the validity of this neotype. Since the syntypes of T. verrucosus are extant (Sclater 1892, Chanda et al. 2000), one of the syntypes should be selected as the lectotype instead of designating a neotype and this designation contravenes the requirements of Art. 74.7 of the Code (ICZN, 1999). Therefore, we agree with Fei et al. (2006) that the designation of the neotype (KIZ 74II0061 VI.16) is unvalid.

We collected some specimens from western Tengchong City and neighbouring north-western Yingjiang County, Yunnan, China, in 2019; these specimens were uniform blackish-brown or uniform brown in life, that is to say, not only the specimens from Husa agree with the original description of T. verrucosus, but also the specimens from western Tengchong and north-western Yingjiang agree with the original description of T. verrucosus. Therefore, Tengchong or Yingjiang cannot be ruled out as the type locality of T. verrucosus. Interestingly, the black surface skins of the ones who underwent poor preservation fell off and the body colour turned to orange-patterned, while others are still uniform blackish-brown or uniform brown (Fig. 5). The colour of some of these specimens in preservative became inconsistent with that in life; this seems to be similar to the case of the syntypes of T. verrucosus. Nevertheless, as we do not know the previous preservation status of the syntypes of T. verrucosus, we cannot determine whether the colours of these syntypes have changed. More interestingly, the specimens from western Tengchong and north-western Yingjiang were assigned to T. panwaensis in the molecular analysis.

Figure 5.  

The specimens of Tylototriton panwaensis from western Yunnan, China, in preservative.

Although the population in Husa and the population in western Tengchong are both in agreement with the original description of Tylototriton verrucosus, the two populations belong to two different species. Since it is impossible to tell whether Hotha or Momien is the type locality of T. verrucosus, it is impossible to identify whether the population in Husa or in western Tengchong is the true T. verrucosus. If Hotha is the type locality of T. verrucosus, then the population in Husa is the true T. verrucosus and T. panwaensis is a valid species and also distributed in China; and if Momien is the type locality of T. verrucosus, then the population in western Tengchong may be the true T. verrucosus, T. panwaensis may be the synonym of T. verrucosus and the population in Husa may remain an unnamed species. We can only make this speculation at present, as more research is needed to solve this problem.

Acknowledgements

Thank to my colleagues for their help and advice. Thanks also to the editors and reviewers for improving 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) and the project of Ministry of Ecology and Environment of China: Investigation and assessment of amphibians and reptiles in Yunnan.

References

Supplementary material

Suppl. material 1: Mean uncorrected p-distances (%), based on mitochondrial ND2 sequences. 
Authors:  Shuo Liu, Miao Hou, Dingqi Rao
Data type:  Table