Corresponding author: Ying-Yong Wang (
Academic editor: A. Crottini
Recent phylogenetic analysis encompassing multilocus nuclear-gene and matrilineal mtDNA genealogy has revealed a series of cryptic species of the subgenus
Lyu Z-T, Li Y-Q, Zeng Z-C, Zhao J, Liu Z-L, Guo G-X, Wang Y-Y (2020) Four new species of Asian horned toads (Anura, Megophryidae,
The Asian horned toad genus
During our herpetological surveys in the hilly areas among Guangdong, Guangxi and Hunan, southern China (Fig.
Map showing the collecting location of the new species.
Localities, voucher information, and GenBank numbers for all samples used in this study.
ID | Species | Localities | Voucher number | 16s | co1 |
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China: Guangxi: Huaping Nature Reserve |
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China: Guangxi: Huaping Nature Reserve |
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China: Guangxi: Huaping Nature Reserve |
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China: Guangxi: Huaping Nature Reserve |
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China: Guangdong: Shimentai Nature Reserve |
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China: Guangdong: Shimentai Nature Reserve |
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China: Guangdong: Shimentai Nature Reserve |
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China: Guangdong: Shimentai Nature Reserve |
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China: Hunan: Mt Yangming |
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China: Hunan: Mt Yangming |
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China: Hunan: Mt Yangming |
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China: Hunan: Mt Yangming |
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China: Hunan: Mt Yangming |
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China: Hunan: Mt Yangming |
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China: Hunan: Mt Yangming |
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China: Hunan: Mt Yangming |
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China: Hunan: Mt Yangming |
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China: Guangdong: Heishiding Nature Reserve |
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China: Chongqing: Baolong Town | KIZ 019216 |
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China: Yunnan: Mt. Jizu | KIZ 019441 |
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China: Sichuan: Mt. Wawu |
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China: Fujian: Mt. Wuyi |
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China: Hong Kong |
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China: Hunan: Badagongshan Nature Reserve |
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China: Hunan: Taoyuandong Nature Reserve |
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China: Yunnan: Mt. Dawei | KIZ 048997 |
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China: Guangdong: Mt. Yinping |
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Vietnam: Lao Cai: Sa Pa | VNMN 2018.01 |
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Vietnam: Lao Cai: Sa Pa | VNMN 07034 |
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China: Anhui: Mt. Huangshan |
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China: Guangdong: Nan’ao Island |
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China: Guizhou: Kuankuoshui Nature Reserve |
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China: Yunnan: Mt. Wuliang |
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China: Jiangxi: Mt. Jinggang |
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China: Jiangxi: Mt. Jiulian |
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China: Fujian: Mt. Wuyi |
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China: Guizhou: Mt. Leigong | SYSa002213 |
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China: Guizhou: Libo Country | 20150813001 |
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China: Hunan: Taoyuandong Nature Reserve |
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China: Zhejiang: Lishui City | WYF00169 |
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China: Sichuan: Mt. Qingcheng |
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China: Hunan: Mt. Mufu |
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China: Guangdong: Mt. Nankun |
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China: Guangdong: Nanling Nature Reserve |
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China: Guangdong: Heishiding Nature Reserve |
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China: Fujian: Mt. Wuyi | WUYI2015101 |
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China: Sichuan: Mt. Emei |
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Vietnam: Thanh Hoa: Pu Hu Nature Reserve | KIZ 011650 |
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Vietnam: Lao Cai: Sa Pa | VNMN 2017.002 |
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China: Hunan: Badagongshan Nature Reserve |
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China: Hunan: Nanshan Forest Park | HNNU 18NS01 |
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China: Guizhou: Mt. Leigong |
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China: Hunan: Badagongshan Nature Reserve |
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China: Jiangxi: Mt. Wugong |
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China: Yunnan: Mt. Wuliang |
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China: Hubei: Shennongjia Nature Reserve |
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China: Zhejiang: Xianju County |
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China: Yunnan: Mt. Gaoligong |
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China: Guangdong: Mt. Sanyue |
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DNA sequences were aligned by the Clustal W algorithm with default parameters (Thompson et al. 1997) and trimmed with the gaps partially deleted in MEGA 6 (Tamura et al. 2013). Two gene segments, 632 base pairs (
External measurements were made for the unnamed specimens with digital calipers (Neiko 01407A Stainless Steel 6-Inch Digital Caliper, USA) to the nearest 0.1 mm. Mean and standard deviation (SD) were calculated in R 3.3.2 (R Core Team 2016). These measurements were as follows:
Sex was determined by secondary sexual characters, i.e., the presence of vocal sac, nuptial pads/spines in males (Fei et al. 2016).
Morphological characters of all 42 recognized congeners of subgenus
Literature for morphological characters of 42 recognized species of
ID | Species | References |
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The
Phylogenetic tree of
The unnamed samples from Huaping Nature Reserve, Guangxi (samples ID 1–4 in Table
The samples from Shimentai Nature Reserve, Guangxi (samples ID 5–8 in Table
The other samples from Mt Yangming, Hunan (samples ID 9–13 in Table
Mean
ID | Species | 1– 4 | 5– 8 | 9– 13 | 14–17 | 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 |
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0.0 | ||||||||||||||||||||||||||||||||||||||||||||
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3.8 | 0.0 | |||||||||||||||||||||||||||||||||||||||||||
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3.9 | 4.1 | 0.0 | ||||||||||||||||||||||||||||||||||||||||||
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3.4 | 3.4 | 4.3 | 0.3 | |||||||||||||||||||||||||||||||||||||||||
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7.3 | 8.8 | 7.3 | 7.3 | / | |||||||||||||||||||||||||||||||||||||||
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3.6 | 4.0 | 4.4 | 2.9 | 7.3 | / | ||||||||||||||||||||||||||||||||||||||
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4.8 | 5.4 | 4.8 | 4.9 | 8.1 | 4.4 | / | |||||||||||||||||||||||||||||||||||||
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4.9 | 5.1 | 5.0 | 4.3 | 8.9 | 3.8 | 2.5 | / | ||||||||||||||||||||||||||||||||||||
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2.4 | 3.4 | 3.9 | 2.1 | 7.0 | 2.7 | 4.0 | 3.7 | / | |||||||||||||||||||||||||||||||||||
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5.4 | 5.5 | 3.9 | 5.2 | 8.1 | 4.6 | 5.3 | 6.5 | 4.7 | / | ||||||||||||||||||||||||||||||||||
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3.5 | 4.3 | 5.0 | 3.2 | 7.3 | 2.5 | 4.4 | 3.7 | 2.4 | 5.8 | / | |||||||||||||||||||||||||||||||||
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3.5 | 4.0 | 3.0 | 3.5 | 7.8 | 2.7 | 3.8 | 4.3 | 2.6 | 4.3 | 3.9 | / | ||||||||||||||||||||||||||||||||
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4.4 | 5.4 | 4.2 | 4.4 | 7.0 | 4.0 | 2.8 | 2.9 | 3.4 | 6.3 | 4.0 | 3.6 | / | |||||||||||||||||||||||||||||||
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5.2 | 4.7 | 4.3 | 4.9 | 8.1 | 4.0 | 5.3 | 5.2 | 4.1 | 5.8 | 5.0 | 2.8 | 4.6 | / | ||||||||||||||||||||||||||||||
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5.1 | 5.0 | 4.7 | 4.6 | 10.5 | 4.0 | 4.6 | 4.2 | 4.2 | 5.5 | 4.6 | 4.4 | 4.6 | 5.7 | / | |||||||||||||||||||||||||||||
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4.6 | 5.0 | 4.7 | 4.0 | 8.4 | 3.8 | 3.2 | 2.9 | 3.6 | 5.7 | 4.0 | 4.0 | 3.6 | 4.9 | 2.5 | / | ||||||||||||||||||||||||||||
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2.8 | 3.8 | 4.3 | 2.6 | 7.0 | 3.0 | 4.4 | 4.1 | 0.7 | 5.0 | 2.8 | 3.0 | 3.6 | 4.5 | 4.6 | 4.0 | / | |||||||||||||||||||||||||||
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5.2 | 4.5 | 4.1 | 4.5 | 9.2 | 4.2 | 5.7 | 5.2 | 4.5 | 5.6 | 5.2 | 2.8 | 5.5 | 4.3 | 4.5 | 4.7 | 4.9 | / | ||||||||||||||||||||||||||
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7.3 | 8.3 | 7.9 | 7.5 | 12.0 | 7.6 | 8.4 | 8.7 | 7.1 | 8.6 | 8.1 | 7.7 | 8.2 | 8.8 | 8.0 | 7.8 | 7.9 | 8.6 | / | |||||||||||||||||||||||||
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4.5 | 5.1 | 4.9 | 4.7 | 8.6 | 4.2 | 2.9 | 2.6 | 4.1 | 6.7 | 3.7 | 4.1 | 2.5 | 6.0 | 4.0 | 4.0 | 4.5 | 5.8 | 8.6 | / | ||||||||||||||||||||||||
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3.5 | 4.9 | 3.0 | 3.7 | 7.3 | 3.6 | 4.2 | 4.1 | 3.2 | 5.0 | 4.3 | 3.0 | 4.0 | 5.0 | 4.6 | 4.4 | 3.6 | 4.1 | 7.0 | 4.3 | / | |||||||||||||||||||||||
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2.8 | 3.8 | 4.3 | 2.8 | 8.1 | 2.9 | 4.9 | 4.3 | 2.2 | 5.2 | 3.0 | 3.5 | 4.8 | 4.5 | 3.8 | 3.2 | 2.6 | 4.5 | 8.1 | 5.0 | 3.9 | / | ||||||||||||||||||||||
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3.7 | 3.8 | 3.4 | 3.6 | 8.1 | 3.0 | 4.6 | 4.7 | 2.8 | 4.1 | 3.7 | 2.4 | 4.0 | 2.8 | 4.4 | 4.2 | 3.2 | 3.4 | 8.1 | 4.9 | 3.7 | 3.6 | / | |||||||||||||||||||||
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2.4 | 3.6 | 4.1 | 2.6 | 7.3 | 2.5 | 3.4 | 3.4 | 1.7 | 4.9 | 2.8 | 3.0 | 3.4 | 4.7 | 3.6 | 3.4 | 2.1 | 4.7 | 6.6 | 3.5 | 3.4 | 2.4 | 3.4 | / | ||||||||||||||||||||
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3.4 | 3.8 | 4.7 | 3.2 | 7.8 | 3.2 | 4.9 | 4.9 | 1.7 | 4.6 | 3.2 | 3.5 | 4.4 | 4.7 | 4.7 | 4.2 | 2.1 | 5.0 | 7.7 | 5.2 | 4.3 | 3.0 | 3.0 | 2.8 | / | |||||||||||||||||||
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4.1 | 4.7 | 2.2 | 4.5 | 7.0 | 4.7 | 4.9 | 5.2 | 3.4 | 3.9 | 4.7 | 3.0 | 4.4 | 4.1 | 5.5 | 5.3 | 3.7 | 4.3 | 8.1 | 5.0 | 3.6 | 4.5 | 3.2 | 3.9 | 4.3 | / | ||||||||||||||||||
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5.0 | 6.9 | 5.8 | 5.0 | 8.4 | 4.5 | 6.3 | 6.3 | 3.8 | 6.8 | 5.3 | 4.8 | 6.0 | 5.8 | 6.5 | 5.3 | 4.8 | 5.3 | 10.6 | 7.8 | 6.0 | 4.8 | 4.5 | 4.5 | 4.5 | 6.0 | / | |||||||||||||||||
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5.6 | 6.2 | 6.2 | 5.1 | 9.5 | 6.1 | 5.5 | 5.6 | 4.5 | 6.4 | 5.6 | 5.4 | 5.1 | 6.4 | 5.5 | 5.1 | 5.1 | 7.1 | 7.0 | 5.8 | 5.4 | 6.0 | 5.4 | 5.1 | 5.4 | 6.2 | 7.6 | / | ||||||||||||||||
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3.8 | 4.2 | 4.7 | 2.6 | 8.6 | 3.2 | 4.6 | 4.3 | 2.4 | 5.4 | 3.2 | 3.6 | 4.4 | 5.1 | 4.5 | 4.0 | 3.0 | 4.5 | 7.2 | 4.7 | 3.4 | 3.2 | 3.4 | 2.6 | 3.4 | 4.9 | 5.0 | 5.5 | / | |||||||||||||||
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4.5 | 4.3 | 4.1 | 4.3 | 8.9 | 3.2 | 5.5 | 5.0 | 3.9 | 5.4 | 4.3 | 2.2 | 4.6 | 2.6 | 5.3 | 5.1 | 4.3 | 4.1 | 8.6 | 5.2 | 4.5 | 4.3 | 3.0 | 4.1 | 4.7 | 4.3 | 5.8 | 6.7 | 4.5 | / | ||||||||||||||
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4.9 | 5.8 | 3.4 | 4.9 | 7.8 | 5.3 | 5.7 | 5.2 | 4.3 | 5.0 | 6.0 | 3.9 | 5.3 | 4.9 | 5.5 | 5.3 | 4.7 | 4.5 | 8.5 | 6.0 | 4.7 | 5.0 | 4.5 | 4.5 | 5.8 | 3.6 | 6.3 | 6.0 | 5.3 | 5.0 | / | |||||||||||||
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6.2 | 7.1 | 5.4 | 6.8 | 8.4 | 6.8 | 7.8 | 8.1 | 5.1 | 7.8 | 7.6 | 3.2 | 5.9 | 5.1 | 7.6 | 7.6 | 5.7 | 5.4 | 11.7 | 7.0 | 6.5 | 6.8 | 4.6 | 5.9 | 6.5 | 5.1 | 5.7 | 9.2 | 7.6 | 4.9 | 6.5 | / | ||||||||||||
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5.2 | 5.8 | 5.0 | 5.2 | 9.7 | 4.9 | 5.9 | 6.2 | 4.5 | 6.0 | 6.0 | 3.2 | 5.3 | 4.7 | 5.5 | 5.5 | 4.9 | 4.5 | 8.4 | 6.0 | 5.2 | 5.2 | 4.7 | 4.3 | 5.6 | 5.2 | 4.8 | 7.1 | 5.4 | 4.7 | 6.0 | 4.1 | / | |||||||||||
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4.7 | 5.1 | 3.9 | 4.5 | 9.2 | 4.6 | 2.7 | 2.4 | 3.9 | 5.8 | 4.3 | 3.9 | 2.7 | 5.0 | 3.6 | 3.0 | 4.3 | 5.2 | 9.0 | 2.4 | 4.5 | 4.5 | 4.3 | 3.6 | 4.7 | 4.5 | 6.1 | 5.6 | 4.5 | 4.7 | 5.2 | 6.8 | 5.4 | / | ||||||||||
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5.7 | 6.7 | 5.5 | 6.1 | 8.4 | 4.9 | 3.6 | 4.0 | 4.9 | 6.6 | 4.8 | 5.1 | 3.6 | 6.3 | 6.1 | 5.1 | 4.9 | 7.0 | 9.5 | 4.4 | 5.7 | 6.3 | 5.1 | 4.4 | 5.9 | 4.7 | 7.3 | 5.7 | 5.9 | 6.1 | 6.3 | 9.2 | 6.8 | 4.0 | / | |||||||||
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4.7 | 5.4 | 5.1 | 4.2 | 9.7 | 4.2 | 2.5 | 2.7 | 2.8 | 5.9 | 4.2 | 3.8 | 2.3 | 5.5 | 4.2 | 3.6 | 3.6 | 5.7 | 8.2 | 2.3 | 4.2 | 4.2 | 4.2 | 3.2 | 4.2 | 5.3 | 6.5 | 5.1 | 3.8 | 4.9 | 5.7 | 7.3 | 5.5 | 2.5 | 4.4 | / | ||||||||
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5.2 | 5.6 | 5.2 | 4.9 | 9.2 | 4.4 | 3.4 | 1.7 | 4.3 | 7.1 | 4.3 | 4.5 | 3.0 | 5.4 | 4.2 | 3.6 | 4.7 | 5.2 | 9.4 | 3.5 | 4.9 | 4.9 | 5.2 | 3.9 | 5.4 | 5.4 | 6.5 | 6.2 | 5.1 | 5.0 | 5.4 | 7.8 | 6.2 | 3.7 | 5.3 | 3.6 | / | |||||||
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1.5 | 3.4 | 3.4 | 2.4 | 5.9 | 2.5 | 4.0 | 4.3 | 1.7 | 4.1 | 3.0 | 2.6 | 4.0 | 4.3 | 4.0 | 3.4 | 2.1 | 4.1 | 6.8 | 4.5 | 3.0 | 2.2 | 2.6 | 1.7 | 2.0 | 3.4 | 3.8 | 4.9 | 2.6 | 3.5 | 4.1 | 5.7 | 4.5 | 4.1 | 5.3 | 4.0 | 4.7 | / | ||||||
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4.9 | 5.1 | 4.7 | 4.5 | 9.2 | 4.4 | 3.2 | 1.1 | 3.9 | 6.7 | 4.3 | 4.5 | 3.2 | 5.4 | 4.2 | 3.2 | 3.9 | 5.6 | 9.4 | 3.4 | 4.9 | 4.5 | 5.2 | 3.5 | 5.0 | 4.7 | 6.5 | 5.8 | 4.9 | 5.4 | 5.0 | 7.8 | 6.2 | 2.8 | 4.6 | 3.4 | 1.7 | 4.3 | / | |||||
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2.6 | 3.4 | 3.7 | 2.2 | 7.0 | 1.7 | 3.6 | 3.0 | 1.7 | 4.7 | 2.4 | 2.2 | 3.4 | 3.9 | 4.0 | 3.4 | 2.1 | 3.9 | 7.7 | 3.4 | 2.6 | 2.4 | 2.6 | 1.9 | 2.8 | 3.9 | 4.3 | 5.1 | 2.6 | 3.0 | 4.5 | 5.1 | 4.3 | 3.9 | 4.8 | 3.2 | 3.2 | 2.1 | 3.5 | / | ||||
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3.9 | 4.2 | 3.9 | 3.9 | 8.9 | 4.2 | 4.6 | 4.5 | 3.6 | 6.0 | 4.7 | 2.8 | 4.0 | 4.3 | 4.2 | 4.0 | 3.7 | 4.5 | 8.1 | 4.3 | 3.6 | 3.4 | 4.3 | 3.7 | 4.7 | 4.5 | 6.0 | 5.3 | 4.9 | 4.3 | 4.5 | 5.7 | 4.9 | 4.5 | 6.5 | 4.2 | 4.5 | 3.7 | 4.1 | 3.4 | / | |||
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3.9 | 5.5 | 4.7 | 4.9 | 9.2 | 4.6 | 3.2 | 3.4 | 4.1 | 6.3 | 4.5 | 4.5 | 2.9 | 5.8 | 4.7 | 4.6 | 4.5 | 5.8 | 8.3 | 3.0 | 4.7 | 4.7 | 4.3 | 3.7 | 4.9 | 5.4 | 6.5 | 5.4 | 4.7 | 5.2 | 6.2 | 7.0 | 5.8 | 3.0 | 4.6 | 2.8 | 4.3 | 3.7 | 3.7 | 3.7 | 4.9 | / | ||
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3.6 | 3.6 | 3.6 | 2.8 | 8.1 | 2.1 | 4.0 | 3.9 | 2.6 | 4.9 | 3.4 | 3.2 | 3.6 | 3.9 | 3.8 | 3.6 | 3.0 | 3.6 | 7.7 | 4.3 | 3.2 | 3.0 | 3.2 | 2.4 | 3.4 | 4.5 | 4.0 | 6.0 | 3.2 | 3.7 | 5.0 | 6.0 | 4.3 | 3.7 | 5.0 | 3.8 | 4.5 | 2.6 | 4.5 | 1.7 | 3.9 | 3.9 | / | |
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3.6 | 4.4 | 4.0 | 3.2 | 7.3 | 3.2 | 4.0 | 4.4 | 2.3 | 4.9 | 3.6 | 3.0 | 3.6 | 4.2 | 4.8 | 3.8 | 3.0 | 4.2 | 7.6 | 4.4 | 4.0 | 3.4 | 3.0 | 2.7 | 3.2 | 4.6 | 2.8 | 5.5 | 3.4 | 4.0 | 5.1 | 5.1 | 4.6 | 3.8 | 5.1 | 3.8 | 5.3 | 2.9 | 5.0 | 2.9 | 4.8 | 4.4 | 3.2 | / |
Diagnostic characters separating all 46 species of the
ID | Species | Horn-like tubercle at upper eyelid: slightly large (2), small (1) | Vomerine teeth: present (1), or absent (0) | Tongue: notched (1), or not notched (0) | Lateral fringes on toes: wide (2), narrow (1), or lacking (0) | Webs on toes: more than one-fourth (2), rudimentary (1), or lacking (0) | ||||
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55.8–61.4 | 68.5–74.8 | 2 | 0 | 0 | 1 | 1 | 0.49–0.63 | 0.45–0.47 | |
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28.0–30.6 | / | 1 | 1 | 0 | 1 | 1 | 0.57–0.66 | 0.44–0.53 | |
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38.6–42.0 | 44.4 | 1 | 0 | 0 | 2 | 1 | 0.38–0.49 | 0.41–0.46 | |
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33.2–37.1 | 45.2 | 1 | 0 | 0 | 1 | 1 | 0.42–0.50 | 0.44–0.51 | |
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27.1–33.0 | 28.1–33.6 | 2 | 0 | 0 | 1 | 1 | 0.57–0.71 | 0.38–0.45 |
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42.0–45.0 | / | 1 | 0 | 1 | 0 | 0 | 0.41 | 0.46 |
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32.0–36.0 | 40.2–42.5 | 1 | 0 | 1 or 0 | 2 | 1 | 0.33–0.50 | 0.46–0.48 |
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45.1–51.0 | / | 1 | 0 | 1 | / | 1 | 0.47–0.52 | 0.52–0.53 |
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34.5–37.8 | 39.7–46.8 | 1 | 0 | 1 | 2 | 1 | 0.40–0.67 | 0.45–0.49 |
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33.7–39.3 | 33.9–45.9 | 1 | 0 | 0 | 0 | 1 | > 0.50 | 0.37–0.42 |
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81.3 | / | 2 | 1 | 0 | / | 1 | 0.5 | 0.51 |
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26.2–29.5 | 31.8–34.1 | 1 | 0 | 1 | 2 | 1 | 0.41–0.54 | 0.50–0.54 |
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34.0–37.0 | 40.0–46.0 | 1 | 1 | / | 0 | 0 | / | 0.54 |
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30.2–39.3 | / | 1 | 1 | 0 | 0 | 1 | 0.42–0.60 | 0.41–0.46 |
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30.9–44.3 | 41.7–42.5 | 1 | 1 | 1 | 0 | 0 | 0.53–0.80 | 0.49–0.59 |
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37.4–47.6 | 59.6 | 1 | 1 | 1 | 0 | 0 | 0.54–0.75 | 0.44–0.63 |
|
|
36.0–41.6 | 44.2 | 1 | 0 | 1 | 0 | 0 | <0.50 | 0.42–0.45 |
|
|
36.8–41.2 | 47.1 | 1 | 1 | 1 | 0 | 1 | 0.46–0.57 | 0.40–0.43 |
|
|
34.4–39.2 | 39.5–40.4 | 1 | 0 | 0 | 0 | 1 | / | / |
|
|
53.0–56.5 | 63.5 | 1 | 1 | 1 | 2 | 2 | / | 0.58–0.59 |
|
|
35.1–36.7 | 38.4–41.6 | 2 | 1 | 0 | 1 | 1 | 0.73–0.88 | 0.47–0.50 |
|
|
30.4–33.9 | 34.1–37.5 | 1 | 1 | 1 | 0 | 1 | 0.50–0.59 | 0.44–0.48 |
|
|
26.2–29.6 | 37.4 | 1 | 0 | 1 | 1 | 0 | 0.44 | 0.38–0.48 |
|
|
38.9 | / | 2 | 1 | 0 | 2 | 1 | 0.85 | 0.52 |
|
|
30.4–38.7 | 42.3 | 1 | 0 | 0 | 0 | 1 | / | / |
|
|
60.5–67.7 | 60.8–70.6 | 2 | 1 | 1 | 2 | 1 | 0.48–0.78 | 0.44–0.61 |
|
|
34.1–39.7 | 37.0–39.9 | 1 | 0 | 0 | 2 | 1 | 0.40–0.60 | 0.46–0.53 |
|
|
30.7–34.7 | 36.9–40.4 | 1 | 0 | 0 | 0 | 0 | / | / |
|
|
34.5–41.2 | / | 1 | 0 | 1 | 0 | 1 | 0.8–0.83 | 0.46–0.48 |
|
|
30.1–30.8 | 36.3 | 1 | 0 | 0 | 1 | 1 | 0.51–0.58 | 0.47–0.53 |
|
|
29.9–34.9 | 39.4–41.9 | 1 | 1 | 0 | 0 | 1 | 0.43–0.61 | 0.35–0.42 |
|
|
30.5–37.3 | / | 1 | 1 | 1 | 1 | 1 | 0.43–0.57 | 0.45–0.51 |
|
|
35.6 | 37.5–41.2 | 1 | 0 | 0 | 0 | 1 | 0.51–0.66 | 0.41–0.47 |
|
|
27.4–34.5 | 32.8–35.0 | 1 | 0 | 0 | 0 | 0 | 0.53–0.69 | 0.33–0.41 |
|
|
56.0–59.5 | 68.0–72.5 | 1 | 1 | 1 | 1 | 1 | / | 0.52–0.56 |
|
|
36.2–38.0 | / | 2 | 1 | 0 | 2 | 2 | / | 0.55 |
|
|
26.7–30.5 | / | 1 | 1 | 1 | 1 | 0 | 0.58–0.76 | 0.48–0.56 |
|
|
54.7 | / | 1 | 1 | 1 | 1 | 1 | 0.62 | 0.59 |
|
|
102.0–118.3 | 99.8–115.6 | 2 | 0 | 1 | 2 | 2 | 0.67 | 0.43–0.47 |
|
|
30.3–33.7 | 37.6 | 1 | 0 | 0 | 0 | 1 | 0.40–0.63 | 0.50–0.55 |
|
|
47.2–54.4 | 54.0–55.0 | 1 | 0 | 1 | 2 | 2 | 0.43 | 0.56–0.58 |
|
|
33.2–39.0 | 50.5 | 1 | 0 | 0 | 0 | 1 | 0.5 | 0.45–0.51 |
|
|
31.0–34.1 | 38.5–42.8 | 1 | 0 | 0 | 0 | 1 | 0.45–0.53 | 0.37–0.44 |
|
|
27.3–31.6 | 41.3 | 1 | 0 | 1 or 0 | 0 | 0 | 0.5 | 0.50–0.51 |
|
|
30.4–35.5 | 38.4 | 1 | 0 | 0 | 0 (in female), 2 (in male) | 1 | 0.5 | 0.47–0.48 |
|
|
31.0–36.3 | 41.6 | 1 | 0 | 0 | 1 | 1 | 0.48–0.60 | 0.40–0.50 |
The specific epithet
Huaping Horned Toad (in English) / Huā Píng Jiăo Chán (花坪角蟾 in Chinese)
Adult male holotype
(1) Body size relatively large,
Further, detailed comparative data of
Five
The holotype specimen of each new species in preservative
With a large body size,
Adult male. Body size large,
Radio-ulna length 0.26 of
Dorsal skin smooth with sparse granules; flanks flabby with spiny tubercles; distinct supratympanic fold curving postero-ventrally from posterior corner of eye to a level above insertion of arm; small tubercles arranged from above the nostril, along the canthus rostralis, edge of upper eyelid and supratympanic fold, to the posterior margin of temporal region; a distinct horn-like prominent tubercle on the edge of upper eyelid; a discontinuous X-shaped ridge with several short ridges on two sides on the back; transverse skin ridges on the dorsal shank and thigh; ventral surface smooth; several tubercles on posterior hindlimbs; small pectoral gland closer to axilla; a single large femoral gland on rear of thigh.
Grayish brown above in life; an dark interorbital triangle with light colored center and edge; a dark X-shaped making with light edge on the central of dorsum; dark brown transverse bands on forearms and hindlimbs; supratympanic fold light gray; dark vertical band below the eye; iris grayish brown; ventral surface grayish white; throat and chest with grayish blue latticed patches and black spots; ventral hands and feet grayish white, tips of digits creamy white, metacarpal tubercle and metatarsal tubercle grayish white; pectoral gland and femoral gland white.
Measurement data of type series are listed in Table
Currently,
Measurements (in mm) of the type series of
|
Male | Male | Female | Female |
|
61.4 | 55.8 | 74.8 | 68.5 |
|
21.4 | 18.8 | 23.7 | 22.6 |
|
21.8 | 18.8 | 23.9 | 22.4 |
|
7.8 | 7.1 | 9.0 | 8.8 |
|
6.7 | 5.9 | 7.5 | 6.8 |
|
7.2 | 6.5 | 8.1 | 7.6 |
|
6.7 | 5.9 | 8.1 | 6.8 |
|
3.3 | 3.2 | 4.3 | 4.3 |
|
3.3 | 3.2 | 4.2 | 3.7 |
|
17.3 | 15.3 | 20.2 | 19.5 |
|
15.9 | 13.9 | 18.3 | 17.6 |
|
43.7 | 37.8 | 48.8 | 43.2 |
|
28.9 | 26.3 | 33.8 | 30.5 |
The specific epithet
Shimentai Horned Toad (in English) / Shí Mén Taí Jiăo Chán (石门台角蟾in Chinese)
(1) Body size small,
Adult male holotype
Comparative data of
Compared with the five
With a small body size,
Advertisement calls spectrograms
Adult male. Body size small,
Radio-ulna length 0.22 of
Dorsal skin rough; numerous granules densely arranged on the top of head, loreal region, lips, temporal region, dorsal body, flanks and dorsal limbs; several tubercles on upper eyelid, including a horn-like prominent tubercle on the edge; all granules and tubercles bearing tiny, barely visible spines; clear supratympanic fold with tiny spines, curving postero-ventrally from posterior corner of eye to a level above insertion of arm; tubercles and granules forming discontinuous “/ \”-shaped ridge and two discontinuous dorsolateral ridges on two sides at the central back; large tubercles and warts on the flanks; ventral surface smooth; several granules bearing black spines on the region around cloaca and rear of hindlimbs; small pectoral gland closer to axilla; a single large femoral gland on rear of thigh.
Light brown above in life; a dark brown stripe on dorsal surface of each eye; narrow dark brown transverse bands on forearms and hindlimbs; supratympanic fold white; dark vertical band below the eye; iris reddish brown; all spines black or dark brown; ventral surface pale; throat flesh color; scarlet spots on the chest; a large white blotch on the belly; a pair of lateroventral longitudinal broad black stripes with several white tubercles on two sides; ventral limbs flesh color with white spots; ventral hands and ventral feet brown, tips of digits pale brown; metacarpal tubercle and metatarsal tubercle reddish; pectoral gland and femoral gland white.
Measurement data of type series are listed in Table
Currently,
The advertisement calls of
Measurements (in mm) of the type series of
|
|
|
|
|
|
|
|
|
|
|
||
|
Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male |
|
28.4 | 28.5 | 28.1 | 30.6 | 29.0 | 29.2 | 28.8 | 28.0 | 30.4 | 28.0 | 29.3 | 28.7 |
|
10.0 | 10.1 | 9.9 | 10.4 | 10.1 | 10.1 | 10.0 | 9.9 | 10.5 | 10.7 | 10.1 | 10.1 |
|
9.6 | 9.5 | 9.5 | 10.0 | 9.6 | 9.7 | 9.5 | 9.8 | 10.0 | 10.3 | 9.8 | 9.9 |
|
3.3 | 3.4 | 3.3 | 3.5 | 3.3 | 3.4 | 3.4 | 3.3 | 3.4 | 3.2 | 3.4 | 3.4 |
|
3.0 | 3.0 | 3.0 | 3.1 | 3.1 | 3.1 | 3.0 | 3.1 | 3.2 | 2.8 | 3.1 | 3.0 |
|
2.6 | 2.8 | 2.8 | 2.9 | 2.8 | 2.8 | 2.7 | 2.8 | 2.9 | 3.1 | 2.8 | 2.6 |
|
3.3 | 3.2 | 3.3 | 3.4 | 3.3 | 3.3 | 3.3 | 3.4 | 3.4 | 3.4 | 3.4 | 3.4 |
|
2.0 | 1.8 | 1.9 | 2.2 | 2.0 | 1.9 | 2.0 | 1.9 | 2.1 | 2.1 | 2.2 | 2.1 |
|
1.6 | 1.6 | 1.5 | 1.6 | 1.7 | 1.5 | 1.5 | 1.4 | 1.6 | 1.5 | 1.5 | 1.5 |
|
7.4 | 7.2 | 7.3 | 7.5 | 7.5 | 7.4 | 7.3 | 6.8 | 7.2 | 7.8 | 7.1 | 7.5 |
|
6.2 | 6.1 | 6.2 | 6.4 | 6.2 | 6.2 | 6.0 | 5.5 | 6.1 | 6.3 | 6.0 | 6.4 |
|
19.1 | 19.9 | 19.0 | 20.5 | 19.4 | 19.3 | 19.3 | 17.9 | 18.7 | 21.4 | 19.4 | 20.3 |
|
13.5 | 14.2 | 13.2 | 14.9 | 13.4 | 14.3 | 13.6 | 12.8 | 13.3 | 14.9 | 13.4 | 14.6 |
Eleven adult specimens, female
The specific epithet
Southern Hunan Horned Toad (in English) / Xiāng Nán Jiăo Chán (湘南角蟾 in Chinese)
(1) Moderate body size,
Adult male holotype
Comparative data of
Compared with the five
With a moderate body size,
Adult male. Moderate body size,
Radio-ulna length 0.22 of
Dorsal skin smooth with sparse granules; sparse tubercles on the flanks; a horn-like prominent tubercle on the edge; clear supratympanic fold curving postero-ventrally from posterior corner of eye to a level above insertion of arm; a discontinuous X-shaped ridge and two discontinuous dorsolateral ridges on two sides at the central back; sparse tubercles on the dorsal shank and thigh; ventral surface smooth; several tubercles on posterior hindlimbs; small pectoral gland closer to axilla; a single large femoral gland on rear of thigh.
Yellowish brown above in life; a dark interorbital triangle with light colored center and edge; a dark X-shaped making with light edge on the central of dorsum; dark brown transverse bands on forearms and hindlimbs; supratympanic fold light colored; dark vertical band below the eye; iris light brown with net-like stripes; throat and anterior chest reddish gray; a longitudinal stripe on the throat; a large white blotch with scarlet spots on the belly; one pair of lateroventral longitudinal broad reddish stripes on two sides; ventral limbs flesh color; ventral hands purplish, tips of fingers pale-grey, metacarpal tubercle reddish; ventral feet purplish brown, tips of fingers pale grey, metatarsal tubercle reddish; pectoral gland and femoral gland white.
Measurement data of type series are listed in Table
The advertisement calls of
Measurements (in mm) of the type series of
|
|
|
|
|
|
|
|
|
|
|
||
|
Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Female |
|
40.9 | 38.7 | 39.0 | 40.2 | 38.6 | 40.5 | 41.7 | 41.5 | 42.0 | 41.0 | 39.1 | 44.4 |
|
13.2 | 12.6 | 12.3 | 13.1 | 12.9 | 13.0 | 13.1 | 13.2 | 13.4 | 13.0 | 12.8 | 14.0 |
|
13.5 | 12.5 | 12.6 | 13.3 | 12.8 | 13.2 | 13.1 | 13.2 | 14.0 | 13.1 | 13.0 | 14.3 |
|
4.5 | 4.4 | 4.3 | 4.4 | 4.3 | 4.6 | 4.2 | 4.5 | 4.7 | 4.6 | 4.3 | 5.0 |
|
4.5 | 4.2 | 4.2 | 4.5 | 4.3 | 4.0 | 4.4 | 4.6 | 4.5 | 4.5 | 4.3 | 4.5 |
|
3.7 | 3.6 | 4.0 | 4.0 | 3.8 | 3.7 | 3.7 | 4.0 | 3.8 | 3.9 | 3.8 | 44.3 |
|
5.4 | 4.6 | 5.0 | 5.1 | 4.8 | 5.0 | 5.0 | 5.1 | 5.1 | 5.0 | 5.1 | 5.5 |
|
2.4 | 2.1 | 2.0 | 2.4 | 2.1 | 1.9 | 2.2 | 2.1 | 2.5 | 2.2 | 2.2 | 2.7 |
|
2.3 | 2.4 | 2.0 | 2.3 | 2.1 | 2.9 | 2.3 | 2.5 | 2.1 | 2.5 | 2.2 | 2.4 |
|
9.3 | 9.0 | 8.8 | 9.0 | 8.9 | 10.3 | 9.3 | 8.9 | 9.0 | 9.3 | 9.2 | 9.8 |
|
9.0 | 8.7 | 8.8 | 8.8 | 8.9 | 9.8 | 9.3 | 8.9 | 9.0 | 9.3 | 9.2 | 9.8 |
|
23.9 | 23.0 | 24.3 | 22.9 | 23.1 | 24.8 | 24.2 | 24.3 | 24.3 | 23.5 | 23.8 | 27.6 |
|
17.0 | 17.9 | 17.8 | 17.0 | 17.9 | 18.2 | 18.2 | 18.2 | 17.3 | 17.4 | 17.7 | 19.1 |
Seven adult specimens from the same locality as the holotype: female
The specific epithet
Mt Yangming Horned Toad (in English) / Yáng Míng Shān Jiăo Chán (阳明山角蟾in Chinese)
Adult male holotype
(1) Body size small,
Comparative data of
Compared with the five
With a small body size,
Adult male. Body size moderate,
Radio-ulna length 0.24 of
Dorsal skin rough with sparse granules; sparse tubercles on the flanks and hindlimbs; several tubercles on upper eyelid, including a horn-like prominent tubercle on the edge; clear supratympanic fold curving postero-ventrally from posterior corner of eye to a level above insertion of arm; a discontinuous X-shaped ridge and two discontinuous dorsolateral ridges on two sides at the central back; four transverse skin ridges on the dorsal shank and thigh; ventral surface smooth; several granules on posterior hindlimbs; small pectoral gland closer to axilla; a single large femoral gland on rear of thigh.
Orange-brown above in life; a triangular making with light edge between eyes; a dark X-shaped making with light edge on the central of dorsum; supratympanic fold light brown; dark vertical band below the eye; iris orange-brown; throat and anterior chest purplish brown; belly dark gray with a large white blotch on the central; ventral limbs purplish; ventral hands reddish brown with dark stripes, tips of fingers pale-grey, metacarpal tubercle reddish; ventral feet purplish, tips of fingers pale-grey, metatarsal tubercle reddish; pectoral gland and femoral gland white.
Measurement data of type series are listed in Table
Currently,
The advertisement calls of
Measurements (in mm) of the type series of
|
Male | Male | Male | Male | Male | Male | Male | Female |
|
35.1 | 34.5 | 36.6 | 33.2 | 37.1 | 36.4 | 34.5 | 45.2 |
|
11.3 | 11.6 | 11.7 | 11.2 | 11.9 | 11.5 | 11.2 | 13.6 |
|
11.4 | 11.9 | 11.7 | 11.1 | 11.8 | 11.5 | 11.3 | 13.5 |
|
4.0 | 4.4 | 4.2 | 3.8 | 4.4 | 4.3 | 4.2 | 4.8 |
|
3.9 | 3.6 | 4.0 | 3.8 | 3.9 | 3.8 | 3.8 | 4.2 |
|
3.4 | 3.8 | 3.8 | 3.4 | 3.8 | 3.6 | 3.7 | 4.4 |
|
4.9 | 4.7 | 4.6 | 4.8 | 4.7 | 5.0 | 4.7 | 5.6 |
|
2.1 | 2.2 | 2.3 | 2.2 | 2.2 | 2.1 | 2.3 | 2.8 |
|
1.5 | 1.4 | 1.4 | 1.4 | 2.1 | 1.7 | 1.6 | 2.1 |
|
8.0 | 8.2 | 8.2 | 7.0 | 8.2 | 8.2 | 7.9 | 10.1 |
|
8.5 | 8.2 | 8.2 | 7.0 | 9.2 | 8.4 | 8.0 | 9.9 |
|
24.2 | 23.3 | 23.4 | 21.1 | 24.1 | 24.1 | 23.2 | 23.0 |
|
17.8 | 17.2 | 17.3 | 15.5 | 17.3 | 17.6 | 17.0 | 19.9 |
The phylogenetic analysis encompassing multilocus nuclear-gene and matrilineal mtDNA genealogy (
The genus
We thank to Yong-You Zhao, Run-Lin Li, Jian Wang, Yu-Long Li, Tian-Du Zhang, Jiang-Mo Zhang, Hai-Long He, Si-Yu Zhang, and Hong-Hui Chen for their help in the fieldwork. This work was supported by the Project of Comprehensive Scientific Survey of Luoxiao Mountains Region of Ministry of Science and Technology of PR China (No. 2013FY111500), the Specimen Platform of Ministry of Science and Technology of PR China, teaching specimens sub-platform (No.2005DKA21403-JK), the Biodiversity Conservation Programme of Ministry of Ecology and Environment of PR China, and the Project of Animal Diversity Survey and Monitoring System Construction of Guangdong Shimentai National Nature Reserve.