Corresponding author: Ying-Yong Wang (
Academic editor: Angelica Crottini
Two new species of the genus
Wang J, Li Y-L, Li Y, Chen H-H, Zeng Y-J, Shen J-M, Wang Y-Y (2019) Morphology, molecular genetics, and acoustics reveal two new species of the genus
The Asian leaf litter toad genus
During recent field surveys in northwestern Guizhou Province of China in 2018, a number of specimens were collected from Zhaozishan Nature Reserve and Wujing Nature Reserve in Qixingguan District of Bijie City, respectively (Figure
Collection localities of the two new
For molecular analyses, a total of 71 sequences (23 muscle tissues were sequenced and 48 sequences downloaded from GenBank) from 32 species of the genus
Localities and voucher data for all specimens used in this study.
ID | Species | Locality | voucher No. | GenBank No. |
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16S rRNA | ||||
1 | China: Wujing Nature Reserve, Bijie City, Guizhou |
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2 | China: Wujing Nature Reserve, Bijie City, Guizhou |
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3 | China: Wujing Nature Reserve, Bijie City, Guizhou |
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4 | China: Wujing Nature Reserve, Bijie City, Guizhou |
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5 | China: Wujing Nature Reserve, Bijie City, Guizhou |
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6 | China: Wujing Nature Reserve, Bijie City, Guizhou |
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7 | China: Wujing Nature Reserve, Bijie City, Guizhou |
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8 | China: Wujing Nature Reserve, Bijie City, Guizhou |
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9 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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10 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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11 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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12 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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13 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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14 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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15 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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16 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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17 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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18 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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19 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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20 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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21 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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22 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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23 | China: Zhaozishan Nature Reserve, Bijie City, Guizhou |
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24 |
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Vietnam: Quang Binh | RH60165 |
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25 |
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Vietnam: Kon Tum | AMS R 173778 |
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26 |
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Vietnam: Kon Tum | AMS R 173635 |
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27 |
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Vietnam: Lam Dong | AMS R 173133 |
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28 |
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Vietnam: Lam Dong | NCSM 77321 |
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29 |
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Vietnam: Lao Cai | AMS R 177673 |
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30 |
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Lao: Phongsaly | MNHN: 2004.0278 |
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31 |
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Vietnam: Kon Tum | AMS R 176524 |
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32 |
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Malaysia: Borneo | KUHE55371 |
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33 |
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Malaysia: Borneo | KUHE55624 |
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34 |
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Malaysia: Borneo | KUHE17545 |
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35 |
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Malaysia: Peninsula | KUHE15487 |
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36 |
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Vietnam: Gia Lai | VNMN A 2015.4 / AMS R 176480 |
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37 |
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China: San zhoutian, Shenzhen |
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38 |
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China: Mt. Wutong, Shenzhen |
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39 |
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China: Mt. Wuyi, Fujian |
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40 |
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China: Mt. Huanggang, Jiangxi |
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41 |
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China: Mangshan, Hunan | MSZTC201702 |
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42 |
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China: Mangshan, Hunan | MSZTC201703 |
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43 |
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Malaysia: Borneo | KUHE53227 |
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44 |
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Malaysia: Borneo | SP21450 |
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45 |
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Vietnam: Phu Yen Prov. | ZMMU-A5823 |
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46 |
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China: Maoershan, Guangxi | KIZ019386 |
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47 |
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Cambodia: Ratanakiri | MVZ258198 |
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48 |
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Thailand: Chiangmai | / |
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49 |
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/ | ROM26828 |
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50 |
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China: Sichuan | SYSa001830 |
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51 |
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Vietnam: Lam Dong | UNS00511 |
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52 |
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Malaysia: Borneo | UNIMAS 8705 |
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53 |
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Vietnam: Lao Cai | MNHN:1999.5675 |
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54 |
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China: Yingjiang, Yunnan |
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55 |
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Vietnam: Lam Dong | ZMMU A-5208 |
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56 |
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Vietnam: Lam Dong | ZMMU A-4873 (ABV-00213) |
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57 |
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Malaysia: Borneo | BORNEENSIS 12632 |
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58 |
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China: Tengchong County, Yunnan |
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59 |
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China: Tengchong County, Yunnan |
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60 |
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Laos: Phongsaly | MNHN 2005.0116 |
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61 |
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China: Zhushihe, Xishuangbanna, Yunnan |
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62 |
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China: Yingjiang, Yunnan |
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63 |
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China: Dawuling Forest Station, Maoming City, Guangdong |
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64 |
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China: Dawuling Forest Station, Maoming City, Guangdong |
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65 |
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China: Mt. Wuhuang, Pubei County, Guangxi |
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66 |
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China: Mt. Wuhuang, Pubei County, Guangxi |
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67 |
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Thailand: Chiang Mai | KJ-2013 |
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68 |
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Vietnam: Lao Cai | AMS R 171623 |
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69 |
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/ | MVZ234658 |
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70 |
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/ | / |
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71 |
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Vietnam: Kon Tum | AMS R173870 |
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All specimens were fixed in 10 % buffered formalin and later transferred to 70 % ethanol for preservation, and deposited at the Museum of Biology,
DNA was extracted from muscle tissue using a DNA extraction kit from Tiangen Biotech (Beijing) Co., Ltd. The mitochondrial gene 16S ribosomal RNA gene (16S rRNA) fragment from each sample was sequenced. Fragments were amplified using primer pairs L3975 (5’-CGCCTGTTTACCAAAAACAT-3’) and H4551 (5’-CCGGTCTGAACTCAGATCACGT-3’) (
Sequences were first aligned in Clustal X 2.0 (
Measurements followed
Sex was determined by direct observation of calling in life, the presence of internal vocal sac openings, and the presence of eggs in abdomen seen via external inspection. Comparative morphological data of
References for morphological characters for congeners of the genus
ID | Literature obtained | |
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Duong et al. 2018 | |
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Inger and Stuebing 1992 | |
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Wang et al. 2018 | |
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Wang et al. 2018 | |
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We compared the advertisement calls from three localities. One was in Wujing Nature Reserve and two were in Zhaozishan Nature Reserve. Advertisement calls were recorded between 20:00–24:00 h on 2–6 July 2018, using a Sony PCM-D100 digital sound recorder held within 20 cm of the calling individuals. The ambient temperature of the type locality was obtained using a Volt TP-2200 Humidity & Temperature Logger. The sound files in wave format were sampled at 44.1 kHz with sampling depth 24 bits. Praat 6.0.27 (
Mean and standard deviation (
Bayesian inference (BI) and maximum likelihood (
Bayesian inference tree of
Uncorrected p-distances among
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0 | |||||||||
3.9–4.2 | 0.0–0.3 | ||||||||
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9.7 | 10.5–10.9 | - | ||||||
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13.5 | 14.7–15.1 | 14.7 | 0 | |||||
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17 | 17.8–18.2 | 15.9 | 10.6 | 0 | ||||
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3.2 | 5.6–5.9 | 10.9 | 14.6 | 17.4 | - | |||
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5.6 | 7.3–7.7 | 12 | 14.7 | 15.4 | 4.2 | - | ||
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14.6 | 14.2–14.6 | 13.1 | 16.6 | 18.7 | 13.5 | 13.8 | - | |
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18.9 | 19.3–19.7 | 16.1 | 18.9 | 16.5 | 18.5 | 17.2 | 18.5 | - |
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22 | 23.3–23.8 | 20.7 | 18.1 | 21.8 | 21.5 | 22.8 | 24.1 | 13.1 |
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18.6 | 20.8–21.2 | 17.4 | 14.9 | 18.1 | 19.4 | 16.5 | 19 | 8.7 |
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21 | 23.2–23.7 | 18.1 | 17 | 18.9 | 21.5 | 21.4 | 22.6 | 19.6 |
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13.5 | 15.1–15.5 | 13.5 | 16.2 | 14.6 | 12 | 13.5 | 12.8 | 18.9 |
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11.3 | 10.9–11.2 | 11.3 | 16.7 | 17.4 | 10.9 | 10.2 | 14.7 | 18.9 |
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8.3 | 9.0–9.4 | 9.4 | 15.4 | 14.6 | 8.7 | 8 | 13.1 | 17.7 |
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9.7 | 10.5–10.8 | 10.5 | 16.2 | 15.7 | 10.1 | 9.4 | 15.1 | 19.3 |
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15.7 | 17.7–18.1 | 15.7 | 13 | 17.7 | 16.1 | 14.9 | 17.3 | 9.4 |
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16.5 | 17.0–17.4 | 17 | 17 | 17.7 | 18.2 | 17.4 | 18.5 | 10.8 |
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15.4 | 17.9–18.3 | 14.2 | 12 | 9.8 | 15.8 | 15 | 17.8 | 16.9 |
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9.8 | 12.8–13.2 | 8.7 | 16.2 | 14.6 | 10.9 | 10.9 | 17.1 | 18.9 |
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11.9 | 14.3–14.7 | 11.7 | 6.3 | 10.3 | 13.8 | 15.1 | 16.6 | 16.9 |
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11.2 | 11.2–11.6 | 6.2 | 15 | 17 | 11.2 | 12 | 14.2 | 18.5 |
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9 | 10.8–11.2 | 5.9 | 13.5 | 15 | 9.4 | 9.7 | 11.6 | 18.1 |
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4.9 | 5.6–5.9 | 10.5 | 14.2 | 18.6 | 3.9 | 5.9 | 13.4 | 18.1 |
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16.1 | 17.3–17.8 | 14.7 | 11.2 | 6.6 | 17.7 | 14.9 | 19 | 16.1 |
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18.5 | 19.7–20.2 | 17.3 | 16.1 | 17.7 | 18.1 | 17.2 | 17.3 | 5.6 |
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9.7 | 11.9–12.3 | 5.2 | 14.6 | 15.4 | 10.1 | 11.2 | 14.2 | 18.4 |
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4.3 | 5.2–5.6 | 10.1 | 13.9 | 14.6 | 3.9 | 3.2 | 13 | 16 |
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10.4–10.8 | 10.8–12.3 | 5.6 | 16.2–16.6 | 17.9–18.3 | 11.6–12.7 | 11.9–13.1 | 11.6–11.9 | 16.9–17.6 |
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10.9 | 12.0–12.4 | 12.4 | 15.6 | 13.9 | 10.9 | 9.4 | 16.2 | 18.1 |
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10.5–10.8 | 12.0–12.7 | 11.7–12 | 17.5 | 16.5–16.9 | 10.1–10.5 | 10.1–10.5 | 16.2 | 20.2–20.6 |
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13.1 | 14.3–14.7 | 8 | 16 | 15.4 | 11.2 | 12 | 13.9 | 19 |
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11.7 | 12.0–12.4 | 10.3 | 15.5 | 16.2 | 11.3 | 10.1 | 13.1 | 19.8 |
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- | ||||||||
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12.8 | - | |||||||
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21.8 | 18.5 | - | ||||||
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23.3 | 17.7 | 22.3 | - | |||||
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22.4 | 18.6 | 22.8 | 15.5 | 0 | ||||
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24.9 | 19.5 | 21.5 | 13.2 | 0.6 | 0 | |||
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24.7 | 21.3 | 22.7 | 14.3 | 5.6 | 2.2 | 0 | ||
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12.4 | 5.3 | 18.4 | 17.7 | 17.3 | 16.1 | 17.7 | - | |
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12 | 10.2 | 19.5 | 16.5 | 19.5 | 174 | 19.5 | 9.4 | - |
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20.7 | 16.5 | 21.4 | 15.4 | 16.6 | 14.6 | 14.9 | 14.9 | 17 |
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24.3 | 20.4 | 21.9 | 15.1 | 7.7 | 6.3 | 6.2 | 17.7 | 19.5 |
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14.9 | 16.6 | 17.7 | 16.7 | 17.6 | 16.7 | 17.9 | 13.4 | 15 |
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21.5 | 19.5 | 19.4 | 14.3 | 9.8 | 9.4 | 9.7 | 16.9 | 17.8 |
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23.7 | 17.3 | 18.1 | 13.1 | 9.8 | 8.3 | 9.7 | 15.7 | 17.3 |
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20.2 | 19.4 | 22.8 | 12.7 | 8.7 | 8.3 | 9 | 17.3 | 17.7 |
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19.7 | 16.9 | 20.5 | 18.3 | 15.4 | 15.3 | 15.7 | 15.3 | 16.5 |
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15 | - | |||||||
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11.3 | 16.3 | - | ||||||
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16.1 | 9.4 | 14.6 | - | |||||
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16.1 | 8.1 | 12.8 | 8 | - | ||||
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16.6 | 11.3 | 14.2 | 9.4 | 9.7 | - | |||
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9 | 15.4 | 12.1 | 15 | 15.8 | 16.1 | - | ||
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16.5 | 19.3 | 16.1 | 18 | 18.1 | 18.4 | 16.9 | - | |
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16.1 | 7.3 | 14.7 | 7.2 | 5.9 | 10.5 | 14.6 | 18.4 | - |
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14.6 | 10.1 | 14.3 | 11.6 | 9.4 | 5.6 | 14.5 | 17.3 | 9.7 |
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8.3–8.7 | 14.9–15.3 | 12.4–12.8 | 16.1–16.5 | 15.7–16.1 | 16.5–16.9 | 7.6–8.0 | 16.9–17.3 | 15.7–16.1 |
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15.3 | 19.9 | 13.7 | 16.9 | 18.9 | 17.3 | 14.9 | 5.2 | 19.3 |
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15.8 | 10.9 | 13.9 | 9.4 | 9.7 | 8.7 | 15.8 | 16.4 | 10.8 |
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17.8–18.1 | 9.4–9.8 | 15.1–15.4 | 6.2–7.2 | 6.2–6.6 | 10.9–12.0 | 16.6–17.0 | 17.6–18.4 | 6.9–7.2 |
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15.7–16.5 | 0.3 | |||||||
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16.9 | 16.1–16.4 | - | ||||||
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8.7 | 16.1–16.5 | 16.5 | 0 | |||||
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10.8–11.9 | 16.1–16.2 | 16.5–16.6 | 16.5–17.3 | 9.4–10.5 | 0.9 | |||
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9.4 | 13.9–14.3 | 18.6 | 9.1 | 12.7 | 13.1 | - | ||
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10.1–10.5 | 16.2–16.5 | 16.6–17.0 | 21.1–21.5 | 12.4–12.7 | 11.6–11.9 | 0.3 | ||
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12.4 | 16.2 | 16.6 | 19 | 13.9 | 9.8 | 10.9–15.6 | 0 | |
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9.4 | 18.2 | 18.7 | 19 | 9.5 | 10.9 | 11.3 | 12.4 | - |
Calling from nine male individuals were measured, respectively. They were recorded in Jinjiazhai Village (two males), Baimashan Forest Station (three males), and Qingshan Village (four males) at an ambient temperature approximately of 18.8 °C, 19.3 °C, and 18.6 °C, respectively. The result of hierarchical clustering analysis was consistent with the molecular result (Figure
Hierarchical clustering of advertisement calls of
Measurements (mean ± standard deviation) of 11 acoustic properties for
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1 | A (n=20) | 111.6±3.0 | 119.1±8.0 | 30.8±27.3 | 4806.2±135.7 | 361.7±53.0 | 4±0 | 6±1 | 87.2±2.4 | 58.2±3.5 | 27.8±3.4 | 25.6±2.8 |
B (n=14) | 189.9±13.6 | 193.6±36.3 | 41.4±20.6 | 4835.7±196.6 | 319.9±148.9 | 4±1 | 20±2 | 69.3±9.6 | 42.6±11.2 | 24.7±5.6 | 122.5±11.4 | ||
2 | A (n=24) | 110.7±5.0 | 128.5±13.7 | 57.8±23.1 | 4679.9±65.6 | 236.9±85.2 | 4±0 | 4±0 | 85.8±3.3 | 51.6±4.7 | 32.4±4.8 | 26.7±3.8 | |
B (n=18) | 188±10.9 | 196.2±39.3 | 54.1±24.0 | 4679.9±66.0 | 306.2±73.7 | 3±0.75 | 19.5±1 | 57.3±30.1 | 43.1±9.1 | 25.3±5.9 | 119.7±7.4 | ||
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3 | A (n=20) | 93.7±6.1 | 102.6±13.2 | 41.3±23.4 | 4823.4±0.0 | 180.9±38.5 | 3±0 | 4±1 | 70.5±3.1 | 45.0±4.2 | 24.5±2.8 | 24.2±6.0 |
B (n=21) | 192.2±13.0 | 174.5±51.1 | 50.4±20.1 | 4675.8±61.7 | 328.1±51.8 | 2±0 | 17±1 | 55.6±9.2 | 37.5±8.6 | 16.9±4.5 | 137.7±8.8 | ||
4 | A (n=20) | 90.3±2.0 | 90.8±5.6 | 47.2±9.9 | 4823.4±0.0 | 344.5±0.0 | 3±0 | 4±0.25 | 69.4±2.0 | 39.4±2.2 | 28.2±1.1 | 22.8±2.4 | |
B (n=25) | 191.8±11.9 | 174.5±51.1 | 60.6±16.6 | 4823.4±0.0 | 186.1±47.7 | 3±1 | 16±2 | 63.9±4.1 | 40.4±6.7 | 22.1±6.2 | 129.4±11.6 | ||
5 | A (n=20) | 91.7±2.2 | 117.2±31.2 | 52.5±9.7 | 4720.1±86.6 | 198.1±63.1 | 3±0 | 4±0 | 67.4±3.4 | 40.3±4.7 | 25.2±3.7 | 26.2±3.4 | |
B (n=11) | 144.8±31.5 | 217.8±64.9 | 27.2±7.1 | 4745.1±89.9 | 219.3±80.4 | 2±0.5 | 13±3.5 | 53.5±6.7 | 33.5±8.0 | 18.3±6.4 | 93.3±25.9 | ||
Summary | A (n=104) | 100.0±10.4 | 112.3±21.3 | 46.3±22.0 | 4767.1±97.3 | 263.4±92.8 | 3±1 | 4±1 | 76.4±9.1 | 47.1±8.1 | 27.8±4.4 | 25.2±4.1 | |
B (n=89) | 185.0±21.7 | 182.7±47.9 | 49.7±21.4 | 4751.8±115.6 | 269.0±100.4 | 3±1 | 17±3 | 60.2±15.7 | 39.8±9.0 | 21.4±6.5 | 123.8±18.3 | ||
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6 | A (n=26) | 103.1±2.3 | 109.0±3.3 | 28.0±31.1 | 5068.6±86.8 | 344.5±0.0 | 2±0 | 3±0 | 87.8±2.1 | 70.1±7.8 | 16.4±7.2 | 16.6±0.8 |
B (n=21) | 221.0±14.0 | 235.6±45.1 | 38.1±20.5 | 5036.7±92.9 | 278.9±85.7 | 2±0 | 17±2 | 82.2±3.9 | 63.0±5.2 | 17.7±5.3 | 140.4±12.8 | ||
7 | A (n=25) | 98.8±6.4 | 122.2±23.4 | 19.5±18.9 | 4823.4±0.0 | 172.3±0.0 | 3±1 | 4±1 | 76.4±3.7 | 55.9±4.9 | 19.3±4.6 | 23.5±6.5 | |
B (n=20) | 206.8±10.1 | 245.2±79.4 | 56.6±20 | 4780.4±76.5 | 206.7±70.7 | 2±1 | 20±3 | 68.4±10.2 | 48.6±9.3 | 18.4±5.7 | 139.7±9.9 | ||
8 | A (n=22) | 102.6±7.5 | 112.9±8.2 | 71.4±15.4 | 4909.6±88.2 | 172.3±0.0 | 3±0 | 4±1 | 85.3±5.2 | 61.3±5.9 | 22.5±1.6 | 18.9±3.2 | |
B (n=28) | 253.0±19.0 | 225.6±61.1 | 71.5±34.9 | 4835.7±45.2 | 319.9±61.4 | 2±0.25 | 23±2.25 | 69.7±11.4 | 51.7±12.6 | 16.5±7.3 | 184.9±19.0 | ||
9 | A (n=33) | 99.4±3.2 | 101.9±6.4 | 34.4±29.1 | 4823.4±0 | 302.8±74.9 | 2±1 | 2±1 | 83.0±2.9.0 | 60.1±6.6 | 21.7±6.6 | 17.7±1.9 | |
B (n=25) | 264.0±12.5 | 166.5±44.5 | 64.6±23.5 | 4823.4±70.3 | 227.4±82.0 | 2±1 | 23±2 | 83.6±7.2 | 61.1±7.8 | 21.2±6.1 | 181.7±11.6 | ||
Summary | A (n=106) | 100.8±5.4 | 110.7±14.5 | 37.0±31.0 | 4901.4±116.8 | 255.2±86.4 | 3±1 | 3±2 | 83.1±5.4 | 61.8±8.2 | 20.0±6.0 | 19.0±4.5 | |
B (n=94) | 239.0±27.0 | 216.3±65.4 | 59.0±28.7 | 4865.6±117.7 | 262.1±86.5 | 2±1 | 21.5±4 | 75.9±11.1 | 56.1±11.0 | 18.4±6.4 | 164.5±25.8 |
* Median instead of mean, inter-quartile range instead of
All advertisement calls contain two notes, each of which consists of repeated pulses (Figure
Different call types (
Combining morphological, molecular genetics, and acoustic evidence, we herein describe these specimens as two new species.
Seven adult males,
(1) small size (
General aspect in life:
Comparative morphological data of
Compared with the 26 known congeners of the genus
Selected diagnostic characters for species described herein and species in the genus
Species | Male |
Black spots on flanks | Toes webbing | Fringes on toes | Ventral coloration | Dorsal skin texture |
---|---|---|---|---|---|---|
29.0–30.4 | Yes | Rudimentary | Narrow | White with distinct nebulous greyish speckling on chest and ventrolateral flanks | Shagreened and granular | |
27.3–29.8 | Yes | Rudimentary | Narrow | Grey purple with distinct nebulous greyish speckling on chest and ventrolateral flanks | Shagreened and granular | |
|
25.1–28.9 | No | Rudimentary | Wide | Near immaculate creamy white, brown specking on margins | Finely tuberculate |
|
24.0–26.4 | Yes | Rudimentary | Wide in males | Creamy-white with dark spots | Relatively smooth, some with small warts |
|
19.6–22.3 | Yes | Rudimentary | No | Reddish brown with white speckling | Smooth |
|
21.3–24.7 | Yes | No | No | Reddish brown with white speckling | Smooth- finely shagreened |
|
18.5–25.4 | Yes | Rudimentary | Weak | Reddish brown with white speckling | Smooth |
|
29.1–32.6 | No | Rudimentary | Narrow | Reddish brown with white speckling | Shagreened |
|
28.0–36.2 | Yes | Rudimentary | Weak | Creamy white | Relatively smooth, some with small warts |
|
22.2–27.3 | No | Rudimentary | No | Bright orange | Highly tuberculate |
|
33.1–34.7 | No | Rudimentary | Wide | Creamy white | Shagreened |
|
26.4–29.2 | No | Rudimentary | Wide in males | Creamy white | Shagreened with fine tubercles |
|
28.2–30.0 | Yes | Rudimentary | Weak | White with brown dusting | Nearly smooth, few tubercles |
|
23.7–27.9 | No | Rudimentary | Wide in males | Creamy white with white dusting on margins | Mostly smooth, females more tuberculate |
|
25.8–30.6 | Yes | No | No | Pale, speckled brown | Smooth |
|
24.5–27.3 | Yes | Rudimentary | Wide | Creamy white | Isolated, scattered tubercles |
|
26.9–28.3 | Yes | Rudimentary | No | Creamy white | Roughly granular |
|
24.8–26.7 | Yes | Rudimentary | Wide | Creamy white with dark brown dusting on margins | Round granular tubercles |
|
23.0-28.7 | Yes | Rudimentary | Wide | Creamy white with dark brown spots on chest and margins | Round granular tubercles with glandular folds |
|
28.0–29.3 | Yes | Rudimentary | No | Greyish-violet with white speckling | Roughly granular with larger tubercles |
|
24.2–26.6 | Yes | No | No | Brown, less white speckling | Mostly smooth |
|
25.2–30.4 | Yes | Rudimentary | Narrow | Creamy white chest and belly with irregular black spots | Longitudinal folds |
|
22.22–27.76 | Yes | Rudimentary | Weak | White speckles on throat and belly | Nearly smooth |
|
19.5–22.7 | Yes | Rudimentary | No | Reddish brown with white speckling | Smooth |
|
25.7–31.4 | Yes | Rudimentary | No | Creamy white | Smooth |
|
40.8 | Yes | Rudimentary | No | Creamy white with light specking on throat and chest | Smooth |
|
26.0–33.0 | Yes | Rudimentary | unknown | Creamy white | Tubercles and longitudinal folds |
|
26.7–32.6 | Yes | Rudimentary | No | Creamy white with white with brown margins | Rounded tubercles |
|
26.6–30.7 | Yes | No | No | Whitish with no markings or only small, light grey spots | Smooth with few glandular ridges |
|
24.5–27.7 | No | No | No | Reddish brown with white speckling | Tuberculate |
|
27.5–32.3 | Yes | Wide | Narrow | Whitish | Small, smooth warts |
|
23.6–27.6 | No | No | Narrow | Immaculate creamy white | Highly tuberculate |
|
21.3–22.3 | Yes | Rudimentary | No | Dirty white with dark brown marbling | Smooth, flattened tubercles on flanks |
|
24.2–28.1 | Yes | Rudimentary | Narrow | Reddish brown with white dusting | Longitudinal skin ridges |
|
25.0–27.5 | Yes | Rudimentary | Wide | Dull white with indistinct grey dusting | Shagreen with small tubercles |
|
30.8–34.3 | Yes | Rudimentary | No | Reddish brown with white speckling | Slightly shagreened |
|
23.4–25.4 | Yes | No | No | Pinkish milk-white to light brown chest and belly with numerous white speckles | Smooth with numerous tiny tubercles |
|
48.3–52.7 | No or small | Wide | Weak | White | Granular |
|
23.3–28.2 | Yes | No | No | Reddish brown with white speckling | Smooth |
|
32.3 | Yes | Wide | Wide | White | Weakly tuberculate |
|
23.9–26.0 | Yes | Rudimentary | Narrow | White with dark brown blotches | Shagreened with small tubercles |
|
24.4–29.5 | No | Rudimentary | No | White with small grey spots/streaks | Highly tuberculate |
|
25.5–28.0 | Yes | Rudimentary | No | Chest and belly with dark brown spots | Longitudinal skin ridges |
|
25.6–30.0 | Yes | Rudimentary | Narrow | Greyish white mixed by tiny white and black dots | Rough, scattered with dense conical tubercles |
|
25.7–27.6 | Yes | Rudimentary | Wide | Creamy white with dark brown flecks on chest and margins | Shagreened with small tubercles |
|
25.9–29.3 | Yes | Rudimentary | Wide | Belly pink with distinct or indistinct speckling | Shagreened with short skin ridges and raised warts |
|
45.8–52.5 | No | Rudimentary | Wide | Creamy-white with white with brown margins | Mostly smooth with distinct tubercles |
For the remaining 45 members of the genus
In having black spots on flanks, the new species differs from
Adult male. Body size small,
Tips of fingers rounded, slightly swollen; relative finger lengths I = II = IV < III; nuptial pad absent; subarticular tubercles absent; a large, rounded inner palmar tubercle distinctly separated from small, round outer palmar tubercle; absence of webbing and lateral fringes on fingers. Tips of toes like fingers; relative toe length I < II < V = III < IV; subarticular tubercles absent; distinct dermal ridges present under the 3rd to 5th toes, not interrupted; large, oval inner metatarsal tubercle present, outer metatarsal tubercle absent; toes webbing rudimentary; narrow lateral fringes present on all toes. Tibia 47% of snout-vent length; tibiotarsal articulation reaches to middle of eye; heels just meeting each other when thighs are appressed at right angles with respect to body.
Dorsal surface shagreened and granular, lacking enlarge tubercles or warts, some of the granules forming short longitudinal folds; ventral skin smooth; dense tiny conical spines present on surface of chest; pectoral gland and femoral gland oval; pectoral glands greater than tips of fingers and femoral glands; femoral gland situated on posteroventral surface of thigh, closer to knee than to vent; supra-axillary gland raised. Ventrolateral gland distinctly visible, forming an incomplete line.
Dorsum greyish-brown grounding, with small reddish granules, distinct darker brown markings and rounded spots and scattered with irregular light orange pigmentation. A dark brown inverted triangular pattern between anterior corner of eyes, in connected to the dark brown W-shaped marking on interorbital region, and the W-shaped marking in connected to the other W-shaped marking between axillae. Tympanum brown. Small light orange granules present on dorsum of body and limb; a dark brown vertical bar under the eye; transverse dark brown bars on dorsal surface of limbs; distinct dark brown blotches on flanks from groin to axilla, longitudinally in two rows; elbow and upper arms with dark bars and distinct coppery orange coloration; fingers and toes with distinct dark bars.
Ventral surface of throat, chest, and belly white, presence of distinct nebulous greyish speckling on chest and ventrolateral flanks; ventral surface of limbs grey purple. Supra-axillary gland coppery orange; femoral, pectoral and ventrolateral glands greyish white. Iris bicolored, coppery orange on upper half and silver on lower half.
Dorsum of body and limbs dark brown; transverse bars on limbs become more distinct; dark brown patterns, markings and spots on back become indistinct, orange pigmentations become greyish white. Ventral surface of body and limbs greyish white, nebulous speckling on chest and flanks balck brown. Supra-axillary, femoral, pectoral and ventrolateral glands greyish white.
Measurements and body proportions were listed in Table
Measurements (minimum–maximum (mean ±
|
|
||
|
29.0–30.4 (29.7 ± 0.6) |
|
43.0–45.5 (43.7 ± 0.8) |
|
10.0–10.6 (10.2 ± 0.2) |
|
1.02–1.05 (1.04 ± 0.01) |
|
9.5–10.2 (9.8 ± 0.3) |
|
0.33–0.35 (0.34 ± 0.01) |
|
4.0–4.7 (4.3 ± 0.3) |
|
0.40–0.44 (0.42 ± 0.02) |
|
3.6–4.1 (3.8 ± 0.2) |
|
1.11–1.15 (1.13 ± 0.02) |
|
2.8–3.4 (3.1 ± 0.2) |
|
1.85–1.95 (1.89 ± 0.04) |
|
2.8–3.4 (3.1 ± 0.2) |
|
1 |
|
1.9–2.2 (2.0 ± 0.1) |
|
0.51–0.54 (0.53 ± 0.01) |
|
0.9–1.1 (1.0 ± 0.1) |
|
0.45–0.53 (0.48 ± 0.02) |
|
13.5–14.4 (13.8 ± 0.3) |
|
0.45–0.47 (0.47 ± 0.01) |
|
7.4–8.3 (7.8 ± 0.3) |
|
0.47–0.49 (0.48 ± 0.01) |
|
13.0–13.8 (13.3 ± 0.2) |
|
1.45–1.50 (1.47 ± 0.02) |
|
14.0–14.8 (14.3 ± 0.3) |
|
0.31–0.32 (0.31 ± 0.01) |
The specific epithet
Currently,
The habitat of
A single adult female,
(1) small size (
General aspect in life:
Comparative morphological data of
In the phylogenetic trees (Figure
Compared with the 26 known congeners of the genus
For the remaining 45 members of the genus
In having black spots on flanks, the new species differs from
Adult male. Body size small,
Tips of fingers rounded, slightly swollen; relative finger lengths I = II = IV < III; nuptial pad absent; subarticular tubercles absent; a large, rounded inner palmar tubercle distinctly separated from small, round outer palmar tubercle; absence of webbing and lateral fringes on fingers. Tips of toes like fingers; relative toe length I < II < V < III < IV; subarticular tubercles absent; distinct dermal ridges present under the 3rd to 5th toes, not interrupted; large, oval inner metatarsal tubercle present, outer metatarsal tubercle absent; toes webbing rudimentary; narrow lateral fringes present on all toes. Tibia 45% of snout-vent length; tibiotarsal articulation reaches to middle of eye; heels just meeting each other when thighs are appressed at right angles with respect to body.
Dorsal surface shagreened and granular, lacking enlarge tubercles or warts, some of the granules forming short longitudinal folds; ventral skin smooth; dense tiny conical spines present on surface of chest and extending to anterior region of abdomen; pectoral gland and femoral gland oval; pectoral glands greater than tips of fingers and femoral glands; femoral gland situated on posteroventral surface of thigh, closer to knee than to vent; supra-axillary gland raised. Ventrolateral gland distinctly visible, forming an incomplete line.
Dorsum dark purple brown grounding, with small light orange granules, distinct darker brown markings and rounded spots and scattered with irregular light orange pigmentations. A dark brown V-shaped pattern between anterior corner of eyes, in connected to the dark brown W-shaped marking on interorbital region, and the W-shaped marking in connected to the other W-shaped marking between axillae. Tympanum brown. A dark brown vertical bar under the eye; transverse dark brown bars on dorsal surface of limbs; distinct dark brown blotches on flanks from groin to axilla, longitudinally in two rows; elbow and upper arms with dark bars and distinct coppery orange coloration; fingers and toes with distinct dark bars.
Ventral surface grey purple, with distinct nebulous greyish speckling scattered with white spots on chest and ventrolateral flanks. Supra-axillary gland coppery orange with dark brown speckling; femoral, pectoral and ventrolateral glands greyish white. Iris bicolored, coppery orange on upper half and silver on lower half.
Dorsum of body and limbs dark brown; transverse bars on limbs become more distinct; dark brown patterns, markings and spots on back become indistinct, orange pigmentations become greyish white. Ventral surface of body and limbs greyish white, nebulous speckling on chest and flanks balck brown. Supra-axillary, femoral, pectoral and ventrolateral glands greyish white.
Measurements and body proportions were listed in Table
Measurements (minimum–maximum (mean ±
Population | A | B | A + B | |||
---|---|---|---|---|---|---|
SEX | Males | Female | Males | Females | Males | Females |
(n = 7) | (n = 1) | (n = 4) | (n = 3) | (n = 11) | (n = 4) | |
|
27.3–29.6 | 35.3 | 28.3–29.8 | 33.0–34.5 |
|
|
(28.6 ± 0.7) | (29.3 ± 0.6) | (33.5 ± 0.7) |
|
|
||
|
9.6–10.2 | 12 | 9.7–10.3 | 11.0–11.7 |
|
|
(9.9 ± 0.2) | (10.1 ± 0.2) | (11.3 ± 0.3) |
|
|
||
|
9.3–9.7 | 11.5 | 9.6–9.8 | 10.5–11.3 |
|
|
(9.5 ± 0.1) | (9.8 ± 0.1) | (10.9 ± 0.3) |
|
|
||
|
3.5–4.0 | 4.6 | 3.8–4.1 | 4.2–4.4 |
|
|
(3.8 ± 0.1) | (4.0 ± 0.1) | (4.3 ± 0.1) |
|
|
||
|
3.1–3.5 | 3.8 | 3.3–3.6 | 3.7–3.9 |
|
|
(3.3 ± 0.2) | (3.5 ± 0.1) | (3.8 ± 0.1) |
|
|
||
|
2.6–3.2 | 3.5 | 3.0–3.2 | 3.2–3.3 |
|
|
(2.9 ± 0.2) | (3.1 ± 0.1) | (3.2 ± 0.1) |
|
|
||
|
2.7–3.5 | 3.6 | 3.2–3.3 | 3.3–3.5 |
|
|
(3.0 ± 0.3) | (3.0 ± 0.1) | (3.4 ± 0.1) |
|
|
||
|
1.7–1.9 | 2.1 | 1.8–1.9 | 2.0–2.1 |
|
|
(1.8 ± 0.1) | (1.8 ± 0.1) | (2.0 ± 0.1) |
|
|
||
|
1.2–1.3 | 1.3 | 1.1–1.2 | 1.2–1.3 |
|
|
(1.3 ± 0.1) | (1.1 ± 0.1) | (1.2 ± 0.1) |
|
|
||
|
12.5–13.3 | 15.5 | 13.2–14.0 | 14.6–15.4 |
|
|
(12.8 ± 0.3) | (13.6 ± 0.4) | (15.0 ± 0.3) |
|
|
||
|
7.0–7.7 | 7.8 | 7.5–7.6 | 7.7–8.0 |
|
|
(7.3 ± 0.2) | (7.5 ± 0.1) | (7.9 ± 0.1) |
|
|
||
|
12.1–12.7 | 14.8 | 12.6–13.2 | 13.7–14.7 |
|
|
(12.4 ± 0.2) | (13.0 ± 0.2) | (14.2 ± 0.4) |
|
|
||
|
12.6–13.8 | 15.5 | 13.4–14.0 | 14.7–15.7 |
|
|
(13.2 ± 0.4) | (13.6 ± 0.2) | (15.1 ± 0.4) |
|
|
||
|
39.0–42.7 | 47.8 | 40.1–44.6 | 46.0–47.3 |
|
|
(40.4 ± 1.4) | (43.2 ± 1.8) | (46.8 ± 0.6) |
|
|
||
|
1.01–1.05 | 1.04 | 1.01–1.06 | 1.04–1.05 |
|
|
(1.04 ± 0.01) | (1.04 ± 0.02) | (1.04 ± 0.01) |
|
|
||
|
0.33–0.35 | 0.34 | 0.34–0.35 | 0.33–0.34 |
|
|
(0.34 ± 0.02) | (0.34 ± 0.01) | (0.34 ± 0.01) |
|
|
||
|
0.36–0.39 | 0.38 | 0.39–0.40 | 0.38 |
|
|
(0.38 ± 0.01) | (0.39 ± 0.01) |
|
||||
|
1.12–1.19 | 1.21 | 1.11–1.15 | 1.11–1.16 |
|
|
(1.14 ± 0.02) | (1.14 ± 0.02) | (1.13 ± 0.02) |
|
|
||
|
1.82–1.89 | 1.81 | 1.83–1.94 | 1.85–1.90 |
|
|
(1.86 ± 0.03) | (1.89 ± 0.04) | (1.87 ± 0.02) |
|
|
||
|
0.53–0.55 | 0.55 | 0.51–0.55 | 0.53–0.54 |
|
|
(0.54 ± 0.01) | (0.53 ± 0.01) | (0.53 ± 0.01) |
|
|
||
|
1.03–1.09 | 1.03 | 1.03–1.10 | 1.03–1.06 |
|
|
(1.06 ± 0.02) | (1.07 ± 0.03) | (1.05 ± 0.01) |
|
|
||
|
0.67–0.76 | 0.62 | 0.61–0.67 | 0.60–0.62 |
|
|
(0.71 ± 0.03) | (0.64 ± 0.02) | (0.61 ± 0.01) |
|
|
||
|
0.44–0.46 | 0.44 | 0.45–0.47 | 0.44–0.46 |
|
|
(0.45 ± 0.01) | (0.47 ± 0.01) | (0.45 ± 0.01) |
|
|
||
|
0.45–0.47 | 0.44 | 0.45–0.47 | 0.44–0.46 |
|
|
(0.46 ± 0.01) | (0.46 ± 0.01) | (0.45 ± 0.01) |
|
|
||
|
1.36–1.46 | 1.35 | 1.42–1.51 | 1.37–1.42 |
|
|
(1.41 ± 0.03) | (1.47 ± 0.04) | (1.39 ± 0.02) |
|
|
||
|
0.31–0.32 | 0.32 | 0.31–0.33 | 0.32–0.33 |
|
|
(0.31 ± 0.01) | (0.32 ± 0.01) | (0.32 ± 0.01) |
|
|
General aspect in life of the female paratypes of
The specific epithet
Currently,
The habitat of
General aspect observed in the field of
The discoveries of
Studies of the taxonomy and phylogeny of
We would like to thank Mr Yong-You Zhao and Jia-He Li for their help in the field work. This work was supported by the Project of the specimen platform of China (2005DKA21403-JK), teaching specimen sub-platform to Y.Y. Wang.