Corresponding author: Bin Wang (
Academic editor: A. Crottini
A new species of the Asian leaf litter toad genus
Liu J, Shi S, Li S, Zhang M, Xiang S, Wei G, Wang B (2023) A new Asian leaf litter toad of the genus
The Asian leaf litter toads of the genus
In recent years, we carried out a series of biodiversity surveys in Hunan and Guizhou Provinces, China and collected some specimens of
A total of 22 specimens of the new species (Suppl. material
Sampling localities in this study. Localities 1–3 were all in China: 1, Tongdao County, Hunan Province; 2, Suining County, Hunan Province; 3, Congjiang County, Guizhou Province, China.
A total of 15 samples of the new species were used for molecular analyses (Table
Information for samples used in molecular phylogenetic analyses in this study.
ID | Species | Locality | Voucher number | GenBank accession number | |
---|---|---|---|---|---|
16S |
|
||||
1 | Tongdao County, Hunan Province, China |
|
/ | ||
2 | Tongdao County, Hunan Province, China |
|
/ | ||
3 | Tongdao County, Hunan Province, China |
|
|
||
4 | Tongdao County, Hunan Province, China |
|
/ | ||
5 | Congjiang County, Guizhou Province, China |
|
/ | ||
6 | Congjiang County, Guizhou Province, China |
|
|
||
7 | Congjiang County, Guizhou Province, China |
|
|
||
8 | Congjiang County, Guizhou Province, China |
|
|
||
9 | Congjiang County, Guizhou Province, China |
|
|
||
10 | Suining County, Hunan Province, China |
|
|
||
11 | Suining County, Hunan Province, China |
|
|
||
12 | Suining County, Hunan Province, China |
|
|
||
13 | Suining County, Hunan Province, China |
|
|
||
14 | Suining County, Hunan Province, China |
|
|
||
15 | Suining County, Hunan Province, China |
|
|
||
16 |
|
Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010904 |
|
/ |
17 |
|
Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010905 |
|
/ |
18 |
|
Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010912 |
|
/ |
19 |
|
Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010908 |
|
/ |
20 |
|
Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010910 |
|
/ |
21 |
|
Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010909 |
|
/ |
22 |
|
Linggongli, Mount Emei, Sichuan Province, China | CIBEMLGL19052104 |
|
|
23 |
|
Mount Emei, Sichuan Province, China | SYS a001830 |
|
/ |
24 |
|
Mount Emei, Sichuan Province, China | CIBEM1867 | / |
|
25 |
|
Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010916 |
|
/ |
26 |
|
Bat Xat District, Lao Cai, Vietnam | ZMMU-A5636-02280 |
|
/ |
27 |
|
Sapa, Lao Cai Province, Vietnam | AMS R 177673 |
|
/ |
28 |
|
Ky Quan San, Lao Cai, Vietnam | AMS R 188515 |
|
/ |
29 |
|
Sapa, Lao Cai, Vietnam | 1999.566 |
|
/ |
30 |
|
Chishui National Nature Reserve, Chishui City, Guizhou Province, China | CIBCS20190518042 |
|
/ |
31 |
|
Changning County, Sichuan Province, China | CIB20050095 |
|
/ |
32 |
|
Zhaozishan Nature Reserve, Bijie City, Guizhou Province, China | SYS a007314 |
|
/ |
33 |
|
Lengshuihe Nature Reserve, Jinsha County, Guizhou Province, China |
|
/ | |
34 |
|
Huoqiuba Nature Reserve, Suiyang County, Guizhou Province, China | GZNU20180606005 |
|
|
35 |
|
Huoqiuba Nature Reserve, Suiyang County, Guizhou Province, China | GZNU20180606002 | / |
|
36 |
|
Wujing Nature Reserve, Bijie City, Guizhou Province, China | SYS a007081 |
|
/ |
37 |
|
Daxueshan Nature Reserve, Yunnan Province, China | KIZ015734 |
|
/ |
38 |
|
Tianquanshan Forest Park, Zhangjiajie, Hunan Province, China | CSUFT 177 |
|
/ |
39 |
|
Nanchuan District, Chongqing City, China | ZYC799 |
|
/ |
40 |
|
Yushe Forest Park, Shuicheng County, Guizhou Province, China | SYS a004961 |
|
/ |
41 |
|
Caiyanghe, Yunnan Province, China | KIZ049024 |
|
|
42 |
|
Yingjiang, Yunnan Province, China | SYS a006530 |
|
/ |
43 |
|
Boun Tay, Phongsaly, Laos | NCSM 80551 |
|
/ |
44 |
|
Baoguosi, Mount Emei, Sichuan Province, China | CIBEMS20190421BGS1 |
|
|
45 |
|
Shengshuige, Mount Emei, Sichuan Province, China | CIBEMS20190422SSG1-4 |
|
|
46 |
|
Doi Inthanon, Chiang Mai, Thailand | KIZ034039 |
|
/ |
47 |
|
Qiyaoshan Nature Reserve, Yunyang County, Chongqing, China | GZNU20210622001 |
|
|
48 |
|
Qiyaoshan Nature Reserve, Yunyang County, Chongqing, China | GZNU20210622002 | / |
|
49 |
|
Gaoligong Shan, Yunnan Province, China | SYS a004598 |
|
/ |
50 |
|
Khasi Hills, Meghalaya, India | SDBDU 2009.329 |
|
|
51 |
|
Mizoram, India | MZMU2224 |
|
/ |
52 |
|
Yingjiang County, Yunnan Province, China | SYS a006533 |
|
/ |
53 |
|
Ba Vi National Park, Ha Tay, Vietnam | ROM 13483 |
|
|
54 |
|
Pu Hu, Thanh Hoa, Vietnam | VNMN:2016 A.23 |
|
/ |
55 |
|
Thanh Hoa Province, Vietnam | VNUF A.2017.37 |
|
/ |
56 |
|
Wuyi Shan City, Fujian Province, China | SYS a001597 |
|
/ |
57 |
|
Mangshan, Hunan Province, China | MSZTC201701 |
|
/ |
58 |
|
Lianshan Bijiashan Nature Reserve, Guangdong, China | GEP a059 |
|
/ |
59 |
|
Shimentai Nature Reserve, Guangdong, China | SYS a004712 |
|
/ |
60 |
|
Basha Nature Reserve, Congjiang County, Guizhou Province, China | GIB196403 |
|
/ |
61 |
|
Mao’er Shan, Guangxi Province, China | KIZ07614 |
|
|
62 |
|
Shenzhen City, Guangdong Province, China | SYS a002450 |
|
/ |
63 |
|
Dawuling Forest Station, Maoming City, Guangdong Province, China | SYS a004663 |
|
/ |
64 |
|
Xiaoqiaogou Nature Reserve, Yunnan Province, China | KIZ016072 |
|
|
65 |
|
Quang Nam Province, Vietnam | AMS R 171714 |
|
/ |
66 |
|
Gia Lai, Vietnam | AMS R 176469 |
|
/ |
67 |
|
Tam Dao, Vinh Phuc, Vietnam | ROM 20236 |
|
|
68 |
|
Chiang Mai, Thailand | KIZ07258 |
|
|
69 |
|
Huanglianshan Nature Reserve, Lyuchun, Yunnan, China | SYS a007743 |
|
/ |
70 |
|
Xiaoqiaogou Nature Reserve, Yunnan Province, China | KIZ048894 |
|
/ |
71 |
|
Tam Dao, Vinh Phu, Vietnam | ROM18282 |
|
/ |
72 |
|
Wenlong, Yunnan Province, China | KIZ013621 |
|
|
73 |
|
Vilabuly, Savannakhet, Laos | NCSM 76038 |
|
/ |
74 |
|
Doi Phu Fa, Nan, Thailand | KIZ024317 |
|
|
75 |
|
Fangcheng City, Guangxi Province, China | NNU202103146 |
|
/ |
76 |
|
Pubei County, Guangxi Province, China | SYS a003485 |
|
/ |
77 |
|
Guangxi Province, China | NNU202103281 |
|
/ |
78 |
|
Ha Giang, Vietnam | ROM 36692 |
|
/ |
79 |
|
Na Hang Nature Reserve, Tuyen Quang, Vietnam | ROM 7035 |
|
/ |
80 |
|
Fansipan, Lao Cai, Vietnam | ROM 30685 |
|
/ |
81 |
|
Guangxi Province, China | NHMG1401032 |
|
/ |
82 |
|
Lam Dong, Vietnam | UNS00511 |
|
/ |
83 |
|
Binh Thuan Province, Vietnam | AMNH A191762 |
|
/ |
84 |
|
Bidoup-Nui Ba National Park, Lam Dong, Vietnam | ZMMU-A-4797-01454 |
|
|
85 |
|
Dak Nong Province, Vietnam | UNS00515 |
|
/ |
86 |
|
Ninh Thuan Province, Vietnam | AMS R 177660 |
|
/ |
87 |
|
Loc Bac, Lam Dong, Vietnam | ZMMU-A-4873-00158 |
|
|
88 |
|
Phu Yen, Vietnam | ZMMU-A5823 |
|
/ |
89 |
|
Ratanakiri, Cambodia | MVZ258198 |
|
/ |
90 |
|
Da Nang City, Vietnam | ITBCZ2783 |
|
/ |
91 |
|
Phong Dien Nature Reserve, Thua Thien-Hue, Vietnam | KIZ010701 |
|
|
92 |
|
Kon Ka Kinh National Park, Gia Lai, Vietnam | ZMMU-NAP-06099 |
|
|
93 |
|
Thua Thien-Hue, Vietnam | ZMMU-NAP-02274 |
|
|
94 |
|
Kon Ka Kinh National Park, Gia Lai, Vietnam | ZMMU-NAP-02275 |
|
|
95 |
|
Fansipan, Lao Cai, Vietnam | AMS R 176540 |
|
|
96 |
|
Phetchaburi, Thailand | KUHE 20197 |
|
/ |
97 |
|
Kapoe, Ranong, Thailand | KIZ018031 |
|
|
98 |
|
Veal Veng District, Pursat Province, Cambodia | CBC 1624 |
|
/ |
99 |
|
Larut, Perak, Malaysia | KUHE15487 |
|
/ |
100 |
|
Gunung Stong, Kelantan, Malaysia | KU RMB20973 |
|
|
101 |
|
Cameron, Malaysia | KUHE 52440 |
|
/ |
102 |
|
Tioman, Malaysia | LSUHC 4431 |
|
/ |
103 |
|
Gunung Mulu, Malaysia | KUHE:55610 |
|
/ |
104 |
|
Borneo, Malaysia | BORNEENSIS 12632 |
|
/ |
105 |
|
Borneo, Malaysia | UNIMAS 8705 |
|
/ |
106 |
|
Danum Valley Field Center, Sabah, Malaysia | FMNH 244800 |
|
|
107 |
|
Liwagu, Kinabalu, Malaysia | BORNEEISIS 22931 |
|
/ |
108 |
|
Bukit Lanjan, Selangor, Malaysia | KUHE17545 |
|
/ |
109 |
|
Borneo, Malaysia | KUHE53227 |
|
/ |
110 |
|
Bukit Kana, Sarawak, Malaysia | FMNH 273682 |
|
|
111 |
|
Borneo, Malaysia | SP 21450 |
|
/ |
112 |
|
Tambunan, Sabah, Borneo, Malaysia | SP 21604 |
|
/ |
113 |
|
Mulu National Park, Sarawak, Malaysia | KUHE:55308 |
|
|
114 |
|
Gunung Mulu National Park, Sarawak, Malaysia | UNIMAS 8957 |
|
/ |
115 |
|
Mulu National Park, Sarawak, Malaysia | KUHE:5589 |
|
|
116 |
|
Gading NP, Sarawak, Borneo, Malaysia | KUHE:47872 |
|
/ |
117 |
|
Mulu NP, Sarawak, Borneo, Malaysia | KUHE 55333 |
|
/ |
118 |
|
Yunnan Province, China | KIZ048439 |
|
|
119 |
|
Yunnan Province, China | KIZ049025 |
|
|
Total DNA was extracted using a standard phenol-chloroform extraction protocol (
Sequences were assembled and aligned using the Clustalw module in BioEdit v. 7.0.9.0 (
All 22 specimens of the new taxon were measured. The terminology and methods followed
One tadpole specimen of the undescribed species was measured. Nineteen morphometric characters were measured for the tadpole:
The new taxon was also compared with all other congeners of
References for morphological characters for congeners of the genus
ID | Species | Literature reviewed |
---|---|---|
1 |
|
|
2 |
|
|
3 |
|
|
4 |
|
|
5 |
|
|
6 |
|
|
7 | ||
8 |
|
|
9 |
|
|
10 |
|
|
11 |
|
|
12 |
|
|
13 | ||
14 | ||
15 |
|
|
16 |
|
|
17 |
|
|
18 |
|
|
19 | ||
20 |
|
|
21 |
|
|
22 |
|
|
23 |
|
|
24 |
|
|
25 |
|
|
26 |
|
|
27 | ||
28 |
|
|
29 |
|
|
30 |
|
|
31 |
|
|
32 |
|
|
33 |
|
|
34 |
|
|
35 |
|
|
36 |
|
|
37 |
|
|
38 |
|
|
39 | ||
40 |
|
|
41 | ||
42 |
|
|
43 |
|
|
44 |
|
|
45 |
|
|
46 |
|
|
47 |
|
|
48 |
|
|
49 |
|
|
50 | ||
51 |
|
|
52 |
|
|
53 |
|
|
54 |
|
|
55 |
|
|
56 |
|
|
57 |
|
|
58 |
|
|
59 |
|
|
60 |
|
|
61 |
|
|
62 | Inger and Stuebing (1992) | |
63 |
|
|
64 | ||
65 |
|
|
66 |
|
|
67 |
|
|
68 |
|
|
69 | ||
70 |
|
|
71 |
|
|
72 |
|
|
73 |
|
|
74 |
|
|
75 |
|
|
76 |
|
|
77 |
|
|
78 |
|
|
79 |
|
|
80 |
|
|
81 |
|
|
82 |
|
|
83 |
|
|
84 |
|
|
85 |
|
|
86 |
|
|
87 |
|
|
88 |
|
|
89 |
|
|
90 |
|
|
91 |
|
|
92 |
|
|
93 |
|
|
94 |
|
|
95 |
|
The advertisement calls of the new taxon were recorded from specimens
The aligned sequence matrix of 16S and
Phylogenetic trees of the genus
The undescribed species could be identified from its congeners in a series of morphological and bioacoustics characters. For the detailed demonstration, based on morphological and bioacoustics comparisons, see the following section describing the new species.
Molecular phylogenetic analyses, morphological comparisons and bioacoustics analyses indicated that the specimens from Tongdao and Suining County, Hunan Province and Congjiang County, Guizhou Province, China represent an undescribed species which is described as follows.
The holotype specimen
Photos of the holotype
Adult male. SVL in 32.0 mm. Head width almost equal with head length slightly (
Fore-limb relatively long (
Hind-limb relatively long (
Dorsal skin relatively smooth with small tubercles and short folds; supra-axillary gland distinct and yellowish; pectoral gland small and indistinct; round femoral glands present and protuberant on rear of thigh, closer to knee than to vent; femoral adipose glands distinct, attached to inner side of skin on posterior ventral surface of thigh; ventral skin smooth; ventrolateral glands forming a distinct white line on flanks.
In life, dorsal surface of head and trunk yellowish-brown, with distinct olive reverse-triangle dark markings between eyes connecting to a dark W-shaped marking between axillae that are fringed with greyish-white colour; elbow to upper arm distinctly yellowish-orange in colour on the dorsum; four transverse black bars present on dorsal surface of thighs and three on dorsal surface of lower arm; one dark blotch between nostril and eyes on loreal region and a dark blotch under the eye; supratympanic ridge reddish and a large black marking under supratympanic ridge; distinct dark blotches on flanks from groin to axilla, longitudinally in two rows; ventral surfaces light coloured; throat and ventral arms pinkish with cream speckling on margins; chest and belly cream white, on the lateral belly with dense brown speckling; ventral hind-limbs pinkish with sparse white glands; upper iris copper, lower iris silver.
Dorsum of body and limbs fading to brown copper; transverse bars on limbs becoming more distinct. Ventral surface of body and limbs fading to cream white. Supra-axillary, femoral and pectoral glands fading to cream yellow.
Measurements and basic statistics of adult specimens are presented in Suppl. materials
Colour variation in
(Fig.
Advertisement calls of
Compared with the 26 known congeners occurring south of the Isthmus of Kra,
Basic statistics for the measurements of
Measurement | Male (n = 15) | Female (n = 7) | ||
---|---|---|---|---|
Ranging | Mean ± SD | Ranging | Mean ± SD | |
SVL | 29.2–34.2 | 30.9 ± 1.4 | 34.4–43.1 | 39.4 ± 2.8 |
|
9.8–11.7 | 10.5 ± 0.6 | 11.8–13.8 | 12.7 ± 0.7 |
|
9.8–12.2 | 11.0 ± 0.7 | 12.4–14.5 | 13.7 ± 0.8 |
|
4.0–5.0 | 4.5 ± 0.3 | 4.9–6.3 | 5.6 ± 0.4 |
|
2.8–4.4 | 3.3 ± 0.4 | 3.4–4.7 | 4.3 ± 0.5 |
|
2.3–3.9 | 3.2 ± 0.5 | 3.4–4.1 | 3.6 ± 0.2 |
|
2.6–3.4 | 2.9 ± 0.2 | 2.9–4.3 | 3.6 ± 0.5 |
|
3.6–4.4 | 4.1 ± 0.2 | 4.2–5.3 | 4.8 ± 0.4 |
|
1.5–2.2 | 1.7 ± 0.2 | 1.6–3.0 | 2.5 ± 0.4 |
|
13.5–15.5 | 14.6 ± 0.5 | 16.8–18.9 | 17.9 ± 0.9 |
|
7.2–8.6 | 7.9 ± 0.4 | 8.6–10.5 | 9.6 ± 0.7 |
|
14.5–16.4 | 15.2 ± 0.7 | 17.4–19.6 | 18.3 ± 0.8 |
|
13.5–15.5 | 14.5 ± 0.6 | 17.4–19.4 | 17.9 ± 1.0 |
|
41.3–51.6 | 47.6 ± 3.1 | 54.6–62.5 | 58.4 ± 2.9 |
Diagnosis characters on morphology of
ID | Species | Male SVL (mm) | Female SVL (mm) | Black spots on flanks | Toes webbing | Fringes on toes | Ventral colouration | Dorsal skin texture |
---|---|---|---|---|---|---|---|---|
1 | 29.2–32.0 | 34.4–43.1 | Yes | Rudimentary | Wide | White with distinct nebulous brown speckling on ventrolateral flanks | Shagreened with fine tubercles | |
2 |
|
25.1–28.9 | 27.1–38.6 | No | Rudimentary | Wide | Near immaculate creamy-white, brown speckling on margins | Finely tuberculate |
3 |
|
24.0–26.4 | 31.7–32.5 | Yes | Rudimentary | Wide in males | Creamy-white with dark spots | Relatively smooth, some with small warts |
4 |
|
19.6–22.3 | 21.7 | Yes | Rudimentary | No | Reddish-brown with white speckling | Smooth |
5 |
|
21.3–24.7 | 25.4 | Yes | No | No | Reddish-brown with white speckling | Smooth-finely shagreened |
6 |
|
22.4 | 25.0–26.4 | Yes | Rudimentary | Narrow | Creamy-white with distinct dark patches on chest and abdomen | Rough with dense conical granules, tubercles and glandular folds |
7 |
|
22.9–25.6 | 27.1 | Yes | Rudimentary | Narrow | Creamy-white chest and off-white belly with irregular black spots | Dorsal skin slightly shagreened with small tubercles and irregular brown stripes |
8 |
|
23.6–24.6 | 29.2–29.4 | Yes | Rudimentary | Weak | Reddish-brown with white speckling | Smooth |
9 |
|
29.0–30.4 | / | Yes | Rudimentary | Narrow | White with distinct nebulous greyish speckling on chest and ventrolateral flanks | Shagreened and granular |
10 |
|
29.1–32.6 | 30.0–31.8 | No | Rudimentary | Narrow | Reddish-brown with white speckling | Shagreened |
11 |
|
27.4–36.2 | 39.5–45.0 | Yes | Rudimentary | Weak | Creamy-white | Relatively smooth, some with small warts |
12 |
|
30.8–33.4 | 34.2 | Yes | Rudimentary | Narrow | White with distinct nebulous greyish speckling on chest and ventrolateral flanks | Shagreened and granular |
13 |
|
22.2–27.3 | / | No | Rudimentary | No | Bright orange | Highly tuberculate |
14 |
|
33.6–34.4 | / | Yes | Rudimentary | Narrow | Creamy-white ventral surface with small, creamy-white glands on throat, chest and belly, becoming more concentrated near lateral margin | Rough dorsal skin with sparse jacinth tubercles and some short longitudinal ridges |
15 |
|
28.7–30.5 | 32.1–39.8 | Yes | Rudimentary | Narrow | Greyish-white with black spots and orange pigmentations | Rough with dense conical granules, tubercles, glandular folds and conical spines |
16 |
|
33.1–34.7 | 40.7 | No | Rudimentary | Wide | Creamy-white | Shagreened |
17 |
|
21.5–22.8 | 25.7 | Yes | Rudimentary | Narrow | Creamy-white with black blotches | Shagreened with small tubercles and ridge |
18 |
|
26.4–29.2 | 25.7–36.9 | No | Rudimentary | Wide in males | Creamy-white | Shagreened with fine tubercles |
19 |
|
23.0–27.0 | 29.3 | Yes | Poorly developed | Narrow | Whitish, black speckling on margins | Shagreened with yellowish-brown tubercles |
20 |
|
28.2–30.0 | / | Yes | Rudimentary | Weak | White with brown dusting | Nearly smooth, few tubercles |
21 |
|
23.1–24.6 | 28.6–32.9 | No | Rudimentary | Wide | White with brown spots | Smooth, with many tubercles |
22 |
|
23.7–27.9 | 28.6–31.5 | No | Rudimentary | Wide in males | Creamy-white with white dusting on margins | Mostly smooth, females more tuberculate |
23 |
|
29.7–31.2 | / | Yes | No | Narrow | Cream yellow, presence of distinct nebulous greyish speckling on flanks | Shagreened and granular |
24 |
|
25.8–30.6 | 28.9–30.6 | Yes | No | No | Pale, speckled brown | Smooth |
25 |
|
24.5–27.3 | 31.2–33.4 | Yes | Rudimentary | Wide | Creamy white | Isolated, scattered tubercles |
26 |
|
26.9–28.3 | 36.6 | Yes | Rudimentary | No | Creamy white | Roughly granular |
27 |
|
24.8–26.7 | 28.1 | Yes | Rudimentary | Wide | Creamy-white with dark brown dusting on margins | Round granular tubercles |
28 |
|
23.0–28.7 | 24.5–27.8 | Yes | Rudimentary | Wide | Creamy-white with dark brown spots on chest and margins | Round granular tubercles with glandular folds |
29 |
|
28.0–29.3 | 30.3 | Yes | Rudimentary | No | Greyish-violet with white speckling | Roughly granular with larger tubercles |
30 |
|
24.2–26.6 | 27 | Yes | No | No | Brown, less white speckling | Mostly smooth |
31 |
|
22.2–27.8 | 30.2 | Yes | Rudimentary | Weak | White speckles on throat and belly | Nearly smooth |
32 |
|
25.2–30.4 | 29.1 | Yes | Rudimentary | Narrow | Creamy-white chest and belly with irregular black spots | Longitudinal folds |
33 |
|
19.5–22.8 | / | Yes | Rudimentary | No | Reddish-brown with white speckling | Smooth |
34 |
|
25.7–31.4 | 31.6–37.3 | Yes | Rudimentary | No | Creamy-white | Smooth |
35 |
|
23.2–24.9 | 29.3–32.1 | Yes | Rudimentary | Wide | Creamy-white belly with small black spots on the margin | Shagreened with reddish tubercles and folds |
36 |
|
40.8 | / | Yes | Rudimentary | No | Creamy-white with light specking on throat and chest | Smooth |
37 |
|
30.9 | 32.1–35.3 | Yes | Rudimentary | No | Creamy-white with brown dusting on margins | Finely tuberculate |
38 |
|
/ | 35.4–36.3 | Yes | Rudimentary (in females) | absent (in females) | Light purplish-grey with dark brown mottling on throat | Small, irregular bumps and ridges |
39 |
|
22.5–23.6 | 28.5–28.7 | Yes | Rudimentary | No | Ventral sides marbled with distinct irregular black speckling | Skin on dorsum scattered with fine reddish tubercles |
40 |
|
26.0–33.0 | 34.0–35.0 | Yes | Rudimentary | unknown | White with distinct nebulous greyish speckling on chest and ventrolateral flanks | Tubercles and longitudinal folds |
41 |
|
26.7–32.6 | 37.0–41.0 | Yes | Rudimentary | No | Creamy-white with white and brown margins | Rounded tubercles |
42 |
|
26.6–30.7 | 31.6 | Yes | No | No | Whitish with no markings or only small, light grey spots | Smooth with few glandular ridges |
43 |
|
24.5–27.7 | / | No | No | No | Reddish-brown with white speckling | Tuberculate |
44 |
|
27.5–32.3 | / | Yes | Wide | Narrow | Whitish | Small, smooth warts |
45 |
|
23.6–27.6 | 30.3–47.0 | No | No | Narrow | Immaculate creamy white | Highly tuberculate |
46 |
|
21.3–22.3 | 25.5–33.5 | Yes | Rudimentary | No | Dirty white with dark brown marbling | Smooth, flattened tubercles on flanks |
47 |
|
24.2–28.1 | 27.3–31.5 | Yes | Rudimentary | Narrow | Reddish-brown with white dusting | Longitudinal skin ridges |
48 |
|
25.0–27.5 | / | Yes | Rudimentary | Wide | Dull white with indistinct grey dusting | Shagreen with small tubercles |
49 |
|
27.3–29.8 | 33.0–35.3 | Yes | Rudimentary | Narrow | Grey purple with distinct nebulous greyish speckling on chest and ventrolateral flanks | Shagreened and granular |
50 |
|
30.3–33.9 | 30.8–34.3 | Yes | Rudimentary | No | Reddish-brown with white speckling | Slightly shagreened |
51 |
|
23.4–25.4 | 27.0–27.8 | Yes | No | No | Pinkish milk-white to light brown chest and belly with numerous white speckles | Smooth with numerous tiny tubercles |
52 |
|
24.9–29.4 | 30.8–35.9 | Yes | Rudimentary | Narrow | ventral surface yellowish-creamy-white with marble texture | Smooth |
53 |
|
26.8–29.7 | / | Yes | No | No | Creamy-white ventral surface with small, creamy-white glands on throat, chest and belly, becoming more concentrated near lateral margin | Shagreened with small raised tubercles and ridges |
54 |
|
48.3–52.7 | 56.7–58.9 | No or small | Wide | Weak | White | Granular |
55 |
|
28.7–29.7 | 30.5–33.5 | Yes | Rudimentary | Narrow | Yellowish-creamy-white with marble texture chest and belly or with irregular light brown speckling | Shagreen with small granules |
56 |
|
23.3–28.2 | 32.1 | Yes | No | No | Reddish-brown with white speckling | Smooth |
57 |
|
32.3 | 32.3 | Yes | Wide | Wide | White | Weakly tuberculate |
58 |
|
23.9–26.0 | 28.8–28.9 | Yes | Rudimentary | Narrow | White with dark brown blotches | Shagreened with small tubercles |
59 |
|
24.4–29.5 | 30.2 | No | Rudimentary | No | White with small grey spots/streaks | Highly tuberculate |
60 |
|
23.7–27.7 | 31.5–35.0 | Yes | Rudimentary | No | Chest and belly with dark brown spots | Longitudinal skin ridges |
61 |
|
25.6–30.0 | 33.0–36.0 | Yes | Rudimentary | Narrow | Greyish-white mixed by tiny white and black dots | Rough, scattered with dense conical tubercles |
62 |
|
24.5–32.8 | 29.9–38.5 | Yes | Rudimentary | Narrow | Creamy white, with distinct or indistinct brown speckling at margins | Shagreened with sparse large warts, sometimes with longitudinal ridges |
63 |
|
25.8–32.6 | 33.7–34.1 | Yes | Rudimentary | Narrow | Ventral belly cream white with variable brown specking | Dorsum relatively smooth with fine tiny granules or short ridges |
64 |
|
25.7–27.6 | / | Yes | Rudimentary | Wide | Creamy-white with dark brown flecks on chest and margins | Shagreened with small tubercles |
65 |
|
25.9–29.3 | 34.0–35.3 | Yes | Rudimentary | Wide | Belly pink with distinct or indistinct speckling | Shagreened with short skin ridges and raised warts |
66 |
|
45.8–52.5 | / | No | Rudimentary | Wide | Creamy-white with white and brown | Mostly smooth with distinct tubercles |
67 |
|
28 | 30 | Yes | Rudimentary | Unknown | Chest and belly with dark brown spots | Smooth |
68 |
|
26.4–8.9 | 30.1–30.7 | Yes | Rudimentary | Wide in males | Greyish-pink with distinct hazy brown speckling on chest and ventrolateral flanks | Round granular tubercles with glandular folds |
69 |
|
23.2–25.9 | / | Yes | Rudimentary | Narrow | Creamy-white with greyish-white and dark brown spots | Shagreened with numerous conical tubercles |
70 |
|
28.3–30.6 | / | Yes | Rudimentary | Narrow | Light greyish-creamy-white, interspersed with light brown spots | Rough with sparse large warts, with short longitudinal ridges |
By having medium size of body (SVL 29.2–34.2 mm in males)
By having a larger size of body (SVL 34.4–43.1 mm in females),
By having black spots on flanks,
By having rudimentary webbing,
By having wide fringes on toes,
By having dorsal surface shagreened with fine tubercles,
The advertisement calls of
In mitochondrial DNA trees,
The tadpole
Adult males with a pair of subgular vocal sacs (Fig.
Habitats of
This specific name “dong” refers to the Dong people, as the new species distributed in the concentrated area of Dong people. We suggest its English common name “Dong leaf litter toads” and Chinese name “Dong Zhang Tu Chan (侗掌突蟾)”.
South-western China was proposed as a biodiversity hotspot (
This new species was found in three localities and the largest geographical distance between the localities is over 200 km. However, in this study, phylogenetic analyses, based on mitochondrial DNA, suggested the three populations as the same species and different from its congeners on a series of morphological characters. This perhaps indicated that the species have a widespread distribution. Further surveys are needed to evaluate the population status of the species.
We are grateful for the kind help from Dong people, especially Zhen Shi and her families in Shangdong Village, Tongdao County, Hunan Province. We are grateful for the help of Fu Shu and Keji Guo of Central South Inventory and Planning Institute of National Forestry and Grassland Administration, Jingbo Long of Huangsang National Nature Reserve and Yongjian QIN of Forestry and Grassland Administration of Suining County for their help in fieldwork in Suining County, Hunan Province.
This work was supported by the Projects from the National Natural Science Foundation of China (Nos. 32270498, 31960099, 32260136, and 32070426); West Light Foundation of The Chinese Academy of Sciences (Grant No. 2021XBZG_XBQNXZ_A_006); Special funds of Guiyang Bureau of Science and Technology for supporting the Guiyang College (GYKJ-GYU[2021]); Guizhou Provincial Science and Technology Projects (Nos. [2020]1Y083, ZK[2022]540 and [2020]4Y029); Forestry Science and Technology Research Project of Guizhou Forestry Department (No. [2020]13, J[2018]03 and [2022]41); Zunyi Bureau of Science and Technology (No. HZ[2020]319); Guizhou Provincial Department of Education Youth Science and Technology Talents Growth Project (Nos. KY[2018]470, KY[2018]452 and KY[2020]234); Science Research Project of Moutai Institute (No.myk2019009); and Biodiversity Survey of Suining County, Hunan Province.
Measurements of adult specimen of
table (excel document)
Morphological data. Units in mm. See abbreviations for characters in the Materials and methods section.
Uncorrected p-distance between
table (excel document)
Genetic distance. The values below 3% are coloured in red.
Call data
table (excel document)
Call measurements (mean ± standard deviation) of