Corresponding author: Ying-Yong Wang (
Academic editor: A. Ohler
Two new toad species of the genus
Wang J, Lyu Z-T, Qi S, Zeng Z-C, Zhang W-X, Lu L-S, Wang Y-Y(2020) Two new
The generic classifications within the family
During recent field surveys in the Yunnan-Guizhou Plateau of southwestern China, a number of megophrid specimens were collected from Yushe Forest Park in western Guizhou (Fig.
Collection sites. Site 1-Yushe Forest Park, Shuicheng County, Guizhou Province, the type locality of
For the molecular analyses, a total of 80 sequences (nine muscle tissue samples was sequenced and 71 sequences obtained from GenBank) were used, including five sequences of the undescribed species from Guizhou, four sequences of the undescribed species from Yunnan, 69 sequences of 66 recognized congeners, and two out-group sequences of
DNA was extracted from muscle tissue using a DNA extraction kit from Tiangen Biotech (Beijing) Co., Ltd. The mitochondrial gene 16S ribosomal RNA gene (
Collection localities, voucher data and GenBank numbers (
ID | Ingroup | Collection Locality | Voucher No. | GenBank No. |
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Huanglianshan Nature Reserve, Lyuchun, Yunnan, China |
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Huanglianshan Nature Reserve, Lyuchun, Yunnan, China |
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Huanglianshan Nature Reserve, Lyuchun, Yunnan, China |
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Huanglianshan Nature Reserve, Lyuchun, Yunnan, China |
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Yushe Forest Park, Shuicheng, Guizhou, China |
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Yushe Forest Park, Shuicheng, Guizhou, China |
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Yushe Forest Park, Shuicheng, Guizhou, China |
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Yushe Forest Park, Shuicheng, Guizhou, China |
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Yushe Forest Park, Shuicheng, Guizhou, China |
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Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ032625 |
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Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ048894 |
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Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ048972 |
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Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ048973 |
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U Bo, Quang Binh, Vietnam | ZFMK 86362 |
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Huangcaoling, Jingdong, Yunnan, China | KIZ046816 |
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Ngoc Linh, Kon Tum, Vietnam | AMS R 173778 |
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Mesilau, Sabah, Malaysia | BORNEENSIS 22931 |
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Kon Ka Kinh, Gia Lai, Vietnam | AMS R 176463 |
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Hon Giao, Lam Dong, Vietnam | NCSM 77321 |
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Zhaozishan Nature Reserve, Guizhou, China |
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Fansipan, Lao Cai, Vietnam | AMS R 176540 |
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Lao Cai, Vietnam | AMS R 177673 |
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Chishui, Guizhou Province, China | CIBCS20190518047 |
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Kon Tum, Vietnam | AMS R 173740 |
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Gunung Mulu National Park, Sarawak, Malaysia | NMBE1056372 |
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Long Nai, Phongsaly, Laos | MNHN 2004.0274 |
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Ngoc Linh Nature Reserve, Kon Tum, Vietnam | AMS R 176524 |
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Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ032626 |
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Base Camp of Mulu NP, Sarawak, Malaysia | KUHE 55371 |
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Phetchaburi, Thailand | KUHE 20174 |
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Camp 1 of Gunung Mulu NP, Sarawak, Malaysia | NMBE1056364 |
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Bukit Lanjak, Malaysia | KUHE 17545 |
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Larut, Malaysia | KUHE 15486 |
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Gia Lai, Vietnam | AMS R 176480 |
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Tioman, Malaysia | LSUHC 4431 |
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Song Luy, Binh Thuan, Vietnam | AMNH A191762 |
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Cameron, Malaysia | KUHE 52440 |
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Meghalaya, India | SDBDU 2009.329 |
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Tai Mo Shan, Hongkong, China |
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Guadun, Mt. Wuyi, Fujian, China |
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Phu Yen, Vietnam | PYU DTD-508 |
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Phuoc Binh, Ninh Thuan, Vietnam | ZFMK 96600 |
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Mangshan Nature Reserve, Hunan, China | MSZTC201701 |
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Maoershan Nature Reserve, Guangxi, China | KIZ019385 |
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Annah Rais, Padawan, Malaysia | KUHE 53192 |
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Kinabalu, Malaysia | SP 21450 |
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Srat Thani, Thailand | KUHE 19719 |
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Virachey, Ratanakiri, Cambodia | MVZ 258197 |
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Changdao, Thailand | KUHE 23733 |
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Na Hang Nature Reserve, Tuyen Quang, Vietnam | ROM 7035 |
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Thanh Hoa, Vietnam | VNUF A.2017.95 |
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Daxueshan Nature Reserve, Yunnan, China | KIZ015734 |
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Malipo, Yunnan, China | ROM 35606 |
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Mt. Emei, Sichuan, China |
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Gia Rich, Lam Dong, Vietnam | UNS00510 |
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Tam Dao, Vinh Phu, Vietnam | MVZ 223642 |
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Tuyen Quang, Vietnam | VNMN:2016 A.06 |
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Gunung Kinabalu National Park, Sabah, Malaysia | UNIMAS 8705 |
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Sa Pa, Lao Cai, Vietnam | MNHN: 1999.5675 |
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Nghe An, Vietnam | AMS R184852 |
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Yingjiang, Yunnan, China |
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Wujing Nature Reserve, Guizhou, China |
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Loc Bac, Lam Dong, Vietnam | ZMMU ABV-00176 |
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Son Tra, Da Nang, Vietnam | ITBCZ 4113 |
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Mahua, Crocker, Malaysia | BORNEENSIS 12454 |
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Shiwandashan, Guangxi, China | NHMG1401032 |
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Terengganu, Malaysia | KUHE 52342 |
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Suiyang, Guizhou, China | GZNU20180606002 |
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Bac Giang, Vietnam | ZMMU-NAP-02269 |
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Dak Nong, Vietnam | UNS00517 |
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Tengchong, Yunnan, China |
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Kon Ka Kinh National Park, Gia Lai, Vietnam | ZMMU-NAP-02275 |
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Xishuangbanna, Yunnan, China |
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Mt. Wuhuang, Pubei, Guangxi, China |
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Tianzishan Nature Reserve, Hunan, China | CSUFT 200 |
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Yingjiang, Yunnan, China |
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Yunkaishan Nature Reserve, Guangdong, China |
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Chiang Mai, Thailand | KIZ07258 |
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Tengchong, Yunnan, China |
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Da Fang, Guizhou, China |
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Sequences were aligned in Clustal X 2.0 (
Measurements followed
Sex was determined by the presence of internal vocal sac openings, and the presence of eggs in abdomen seen via external inspection.
All specimens were fixed in 10% buffered formalin and later transferred to 70% ethanol for preservation, and deposited at the
Comparative morphological data of
Data sources of the 82 currently known species of the genus
ID | Literature | |
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Günther 1895 | |
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Inger and Stuebing 1992 | |
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Qian et al. in publication | |
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The
Bayesian Inference tree. The Bayesian posterior probabilities (
As shown by the phylogenetic result,
Samples of the other unnamed lineage from Yushe Forest Park, cluster in another distinct and robust monophyletic lineage with strong support (
Comparisons of selected diagnostic characters for the new species described herein and congeners occurring north of the Kra Isthmus (modified from Rowley et al. 2017;
Male |
Black spots on flanks | Toe webbing | Toe fringes | Ventral coloration | Dorsal skin texture | |
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22.4 | Present | Rudimentary | Narrow | Creamy white with distinct dark patches on chest and abdomen | Rough with dense conical granules, tubercles, and glandular folds | |
28.7–30.5 | Present | Rudimentary | Narrow | Greyish white with black spots and orange pigmentations | Rough with dense conical granules, tubercles, glandular folds, and conical spines | |
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25.1–28.9 | Absent | Rudimentary | Wide | Near immaculate creamy white, brown speckles on margins | Finely tuberculate |
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24.0–26.4 | Present | Rudimentary | Wide in males | Creamy-white with dark spots | Relatively smooth, some with small warts |
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19.6–22.3 | Present | Rudimentary | Absent | Reddish brown with white speckles | Smooth |
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21.3–24.7 | Present | Absent | Absent | Reddish brown with white speckles | Smooth, finely shagreened |
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18.5–25.4 | Present | Rudimentary | Weak | Reddish brown with white speckles | Smooth |
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29.0–30.4 | Present | Rudimentary | Narrow | White with distinct nebulous greyish speckles on chest and ventrolateral flanks | Shagreened and granular |
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29.1–32.6 | Absent | Rudimentary | Narrow | Reddish brown with white speckles | Shagreened |
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28.0–36.2 | Present | Rudimentary | Weak | Creamy white | Relatively smooth, some with small warts |
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22.2–27.3 | Absent | Rudimentary | Absent | Bright orange | Highly tuberculate |
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30.8–33.4 | Present | Rudimentary | Narrow | White with distinct nebulous greyish speckles on chest and ventrolateral flanks | Shagreened and granular |
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33.1–34.7 | Absent | Rudimentary | Wide | Creamy white | Shagreened |
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21.5–22.8 | Present | Rudimentary | Narrow | Creamy white with black blotches | Shagreened with small tubercles |
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26.4–29.2 | Absent | Rudimentary | Wide in males | Creamy white | Shagreened with fine tubercles |
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23.0–27.0 | Present | Poorly developed | Narrow | Whitish with black speckles on margins | Shagreened with yellowish- |
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28.2–30.0 | Present | Rudimentary | Weak | White with brown dusting | Nearly smooth with few tubercles |
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23.7–27.9 | Absent | Rudimentary | Wide in males | Creamy white with white dusting on margins | Mostly smooth, females more tuberculate |
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25.8–30.6 | Present | Absent | Absent | Pale, speckled brown | Smooth |
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24.5–27.3 | Present | Rudimentary | Wide | Creamy white | Isolated, scattered tubercles |
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24.8–26.7 | Present | Rudimentary | Wide | Creamy white with dark brown dusting on margins | Round granular tubercles |
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23.0–28.7 | Present | Rudimentary | Wide | Creamy white with dark brown spots on chest and margins | Round granular tubercles with glandular folds |
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26.9–28.3 | Present | Rudimentary | Absent | Creamy white | Roughly granular |
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28.0–29.3 | Present | Rudimentary | Absent | Greyish violet with white speckles | Roughly granular with larger tubercles |
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24.2–26.6 | Present | Absent | Absent | Brown with few white speckles | Mostly smooth |
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22.2–27.8 | Present | Rudimentary | Weak | White speckles on throat and belly | Nearly smooth |
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25.2–30.4 | Present | Rudimentary | Narrow | Creamy white chest and belly with irregular black spots | With longitudinal folds |
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19.5–22.7 | Present | Rudimentary | Absent | Reddish brown with white speckles | Smooth |
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25.7–31.4 | Present | Rudimentary | Absent | Creamy white | Smooth |
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40.8 | Present | Rudimentary | Absent | Creamy white with light speckles on throat and chest | Smooth |
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22.5–23.6 | Present | Rudimentary | Narrow | Marbling with black speckles | Relatively smooth with small tubercles |
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26.0–33.0 | Present | Rudimentary | Unknown | Creamy white | Tubercles and longitudinal folds |
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26.7–32.6 | Present | Rudimentary | Absent | Creamy white with brown margins | Rounded tubercles |
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30.9 | Present | Rudimentary | Absent | Immaculate white, chest and belly with dark specking on outer margins | Low, round tubercles, more dense |
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- | Present | Weak (in females) | Absent (in females) | Light purplish gray with dark |
Small, irregular bumps and ridges |
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26.6–30.7 | Present | Absent | Absent | Whitish with no markings or only small, light grey spots | Smooth with few glandular ridges |
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24.5–27.7 | Absent | Absent | Absent | Reddish brown with white speckles | Tuberculate |
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27.5–32.3 | Present | Wide | Narrow | Whitish | Small, smooth warts |
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23.6–27.6 | Absent | Absent | Narrow | Immaculate creamy white | Highly tuberculate |
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21.3–22.3 | Present | Rudimentary | Absent | Dirty white with dark brown marbling | Smooth, flattened tubercles on flanks |
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24.2–28.1 | Present | Rudimentary | Narrow | Reddish brown with white dusting | With longitudinal skin ridges |
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25.0–27.5 | Present | Rudimentary | Wide | Dull white with indistinct grey dusting | Shagreen with small tubercles |
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27.3–29.8 | Present | Rudimentary | Narrow | Grey purple with distinct nebulous greyish speckles on chest and ventrolateral flanks | Shagreened with granules |
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30.8–34.3 | Present | Rudimentary | Absent | Reddish brown with white speckles | Slightly shagreened |
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23.4–25.4 | Present | Absent | Absent | Pinkish milk-white to light brown chest and belly with numerous white speckles | Smooth with numerous tiny tubercles |
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28.7–29.7 | Present | Rudimentary | Narrow | Yellowish creamy-white with marble texture chest and belly or with irregular light brown speckles | Shagreen with small granules |
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48.3–52.7 | Absent or small | Wide | Weak | White | Granular |
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23.3–28.2 | Present | Absent | Absent | Reddish brown with white speckles | Smooth |
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32.3 | Present | Wide | Wide | White | Weakly tuberculate |
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23.9–26.0 | Present | Rudimentary | Narrow | White with dark brown blotches | Shagreened with small tubercles |
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24.4–29.5 | Absent | Rudimentary | Absent | White with small grey spots/streaks | Highly tuberculate |
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25.5–28.0 | Present | Rudimentary | Absent | Chest and belly with dark brown spots | Longitudinal skin ridges |
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25.6–30.0 | Present | Rudimentary | Narrow | Greyish white mixed by tiny white and black dots | Rough, scattered with dense conical tubercles |
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22.7–30.5 | Present | Rudimentary | Narrow | Translucent creamy white, with distinct or indistinct brown speckles at margins | Shagreened with sparse large warts, some with longitudinal ridges |
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25.7–27.6 | Present | Rudimentary | Wide | Creamy white with dark brown flecks on chest and margins | Shagreened with small tubercles |
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25.9–29.3 | Present | Rudimentary | Wide | Belly pink with distinct or indistinct speckles | Shagreened with short skin ridges and raised warts |
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45.8–52.5 | Absent | Rudimentary | Wide | Creamy-white with brown margins | Mostly smooth with distinct tubercles |
Morphological features in life.
(1) Small size (
From the 26 known congeners of the genus
While
For the remaining 54 members of the genus
Adult male. Body size small,
Tips of fingers rounded, slightly swollen; relative finger lengths I < IV < II < III; nuptial pad absent; subarticular tubercles absent; large, rounded inner palmar tubercle distinctly separated from small, rounded outer palmar tubercle; webbing and lateral fringes on fingers absent. Tips of toes rounded, slightly swollen; relative toe length I < II < V < III < IV; subarticular tubercles absent; distinct longitudinal 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 reaching to anterior corner of eye; heels slightly overlapping when thighs are appressed at right angles with respect to body.
Dorsal skin rough, with dense conical granules, tubercles and glandular folds; ventral skin smooth; sparse tiny tubercles present on surface of chest; pectoral gland and femoral gland oval; the size of pectoral glands almost equal to tips of fingers and femoral glands; femoral gland situated on posteroventral surface of thigh, closer to knee than to vent; supra-axillary glands raised. Ventrolateral glands distinctly visible, raised, forming an incomplete line.
Dorsum greyish brown with small light orange granules, distinct darker brown markings scattered with irregular light orange and greyish white pigmentations. A dark brown inverted triangular pattern between the anterior corners of the eyes in connection with a dark brown W-shaped marking in the interorbital region, which is also connected to a W-shaped marking between the axillae. Tympanum dark brown. Small light orange granules present on dorsum of body and limbs; a dark brown blotch under the eye; transverse dark brown bars present on dorsal surface of limbs and digits; distinct dark brown patches with light yellowish green margin on flanks from groin to axilla; elbow and upper arms with distinct coppery orange coloration.
Ventral surface of throat, chest, and belly creamy white; presence of distinct nebulous greyish speckles present on throat, and distinct dark patches on chest and abdomen; ventral surface of limbs greyish purple, scattered with greyish white spots and small patches. Supra-axillary gland coppery orange; femoral, pectoral, and ventrolateral glands greyish white. Iris bicolored, amber on upper half and silver on lower half.
Morphological features in preserved specimens of
Measurements and body proportions are listed in Table
Measurements and body proportions of
Voucher | ||||
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Male | Female | Female | Female |
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22.4 | 25.3 | 25.0 | 26.4 |
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8.1 | 9.5 | 9.5 | 9.6 |
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8.0 | 9.3 | 9.2 | 9.0 |
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3.7 | 3.8 | 3.8 | 3.4 |
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2.5 | 2.3 | 2.7 | 2.7 |
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2.3 | 2.5 | 2.5 | 2.5 |
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2.9 | 3.2 | 3.2 | 3.1 |
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1.5 | 1.8 | 1.9 | 1.6 |
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0.7 | 1.0 | 1.0 | 0.8 |
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5.9 | 7.0 | 6.6 | 6.3 |
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11.2 | 13.5 | 12.7 | 12.6 |
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10.1 | 11.7 | 10.2 | 11.1 |
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10.6 | 12.4 | 11.9 | 11.9 |
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34.4 | 41.5 | 40.4 | 39.1 |
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0.36 | 0.37 | 0.38 | 0.36 |
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0.36 | 0.37 | 0.37 | 0.34 |
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0.99 | 0.98 | 0.97 | 0.94 |
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0.16 | 0.15 | 0.15 | 0.13 |
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0.31 | 0.25 | 0.29 | 0.30 |
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0.29 | 0.27 | 0.27 | 0.28 |
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1.07 | 0.91 | 1.08 | 1.09 |
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0.36 | 0.34 | 0.34 | 0.32 |
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0.52 | 0.56 | 0.60 | 0.51 |
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0.26 | 0.28 | 0.26 | 0.24 |
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0.50 | 0.53 | 0.51 | 0.48 |
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0.45 | 0.46 | 0.41 | 0.42 |
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0.47 | 0.49 | 0.48 | 0.45 |
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1.53 | 1.64 | 1.61 | 1.48 |
The specific epithet,
Currently,
Morphological features in life.
(1) Small size (
Compared with the 26 known congeners of the genus
For the remaining 56 members of the genus
Adult male. Body size rather small,
Tips of fingers rounded, slightly swollen; relative finger lengths II = IV < I < III; nuptial pad absent; subarticular tubercles absent; large, rounded inner palmar tubercle distinctly separated from small, rounded outer palmar tubercle; absence of webbing and lateral fringes on fingers. Tips of toes rounded, slightly swollen; relative toe length I < II < V < III < IV; subarticular tubercles absent; distinct longitudinal dermal ridges present under the 3rd to 5th toes, interrupted; large, oval inner metatarsal tubercle present, outer metatarsal tubercle absent; toes webbing rudimentary; narrow lateral fringes present on all toes. Tibia 44% of snout-vent length; tibiotarsal articulation reaches to posterior corner of eye; heels slightly overlapping when thighs are appressed at right angles with respect to body.
Dorsal skin rough, with dense conical granules, tubercles, glandular folds and conical spines; ventral skin smooth; pectoral gland and femoral gland oval; the size of pectoral glands almost equal to tips of fingers and femoral glands; femoral gland situated on posteroventral surface of thigh, closer to knee than to vent; supra-axillary glands raised. Ventrolateral glands distinctly visible, raised, forming an incomplete line.
Dorsum greyish brown with distinct darker brown markings on sides and scattered with irregular light greyish brown pigmentations and yellowish brown spots. An indistinct, darker brown inverted triangular pattern between anterior corners of the eyes, connected to an indistinct dark brown W-shaped marking between the axillae. Dense translucent spines present on dorsal skin of body and limbs. Upper 2/3 of the tympanum dark brown, lower 1/3 light orange, scattered with tiny coppery orange spots. Small greyish white and light brown granules present on the dorsum of the body and limbs; a dark brown vertical bar under the eye; transverse dark brown bars on the dorsal surface of the limbs and digits; distinct dark brown patches on the flanks, from groin to axilla; elbow and upper arms with distinct light orange coloration.
Ventral surface of throat, chest, and belly greyish white; throat with light brown speckles, chest, and abdomen with distinct dark patches; ventral surface of limbs dark grey, scattered with greyish white spots and small patches. Supra-axillary gland light orange; femoral, pectoral, and ventrolateral glands greyish white. Iris bicolored, light orange on upper half and silver on lower half.
Measurements and body proportions are listed in Table
Measurements, and body proportions of
Voucher | |||||||
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Male | Male | Female | Female | Female | Female | Female |
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28.7 | 30.5 | 36.1 | 37.3 | 39.8 | 32.1 | 33.8 |
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10.3 | 10.8 | 12.8 | 12.4 | 12.9 | 11.3 | 12.0 |
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10.6 | 10.7 | 11.9 | 12.9 | 13.2 | 11.8 | 12.2 |
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4.5 | 4.2 | 5.1 | 5.4 | 5.8 | 5.0 | 4.8 |
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3.1 | 3.2 | 3.6 | 3.9 | 4.0 | 3.7 | 3.4 |
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3.4 | 2.9 | 3.5 | 3.4 | 3.3 | 3.0 | 2.9 |
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3.5 | 3.7 | 3.9 | 3.7 | 4.3 | 4.2 | 3.8 |
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1.7 | 1.6 | 2.3 | 2.3 | 2.6 | 2.1 | 2.1 |
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1.1 | 0.8 | 1.3 | 1.4 | 1.5 | 1.2 | 1.1 |
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7.4 | 7.3 | 8.8 | 7.7 | 9.1 | 7.8 | 7.6 |
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14.1 | 14.2 | 17.1 | 16.8 | 17.5 | 16.2 | 15.9 |
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12.1 | 12.8 | 14.9 | 14.5 | 15.5 | 13.9 | 13.6 |
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13.5 | 13.4 | 15.5 | 16.3 | 16.6 | 14.9 | 14.5 |
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41.7 | 42.7 | 49.1 | 49.9 | 52.9 | 46.8 | 48.0 |
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0.36 | 0.35 | 0.35 | 0.33 | 0.32 | 0.35 | 0.36 |
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0.37 | 0.35 | 0.33 | 0.35 | 0.33 | 0.37 | 0.36 |
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1.03 | 0.99 | 0.93 | 1.04 | 1.02 | 1.04 | 1.02 |
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0.44 | 0.39 | 0.40 | 0.44 | 0.45 | 0.44 | 0.40 |
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0.29 | 0.30 | 0.30 | 0.30 | 0.30 | 0.31 | 0.28 |
|
0.32 | 0.27 | 0.29 | 0.26 | 0.25 | 0.25 | 0.24 |
|
0.34 | 0.34 | 0.30 | 0.30 | 0.33 | 0.37 | 0.32 |
|
0.49 | 0.43 | 0.59 | 0.62 | 0.60 | 0.50 | 0.55 |
|
0.26 | 0.24 | 0.24 | 0.21 | 0.23 | 0.24 | 0.22 |
|
0.49 | 0.47 | 0.47 | 0.45 | 0.44 | 0.50 | 0.47 |
|
0.42 | 0.42 | 0.41 | 0.39 | 0.39 | 0.43 | 0.40 |
|
0.47 | 0.44 | 0.43 | 0.44 | 0.42 | 0.46 | 0.43 |
|
1.45 | 1.40 | 1.36 | 1.34 | 1.33 | 1.46 | 1.42 |
The specific epithet,
Currently,
In the phylogenetic tree, the
Yunnan and Guizhou are both largely within the species-rich Dian freshwater zoogeographical dominion (
We are grateful to Prof. Tao Thien Nguyen, Prof. Annemarie Ohler, and an anonymous reviewer for providing constructive comments that greatly improved the quality of the manuscript. We would like to thank Yu-Long Li, Run-Lin Li and Hong-Hui Chen for their help in the field work, and Yao Li for her help in the lab work.
Table S1. Pairwise distances based on 16S gene among all sample used in this study
phylogenetic
Genetic distances among all