Corresponding authors: Zhi-Tong Lyu (
Academic editor Raffael Ernst
The recognition for the floating frogs’ genus
The floating frog subfamily
Different generic affiliations for the congeners of
Species original nomen | Authorship |
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Subclade in this study | References for morphological characters |
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Gravenhorst, 1829 |
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I: |
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Gray, 1831 |
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I: |
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Inger, Boeadi & Taufik, 1996 |
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II | |
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Matsui, Nishikawa, Eto, Hamidy, Hossman & Fukuyama, 2021 |
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II | |
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Chen, Peng, Liu, Huang, Liao & Mo, 2022 |
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III | This study |
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Boulenger, 1896 |
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IV: |
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Taylor, 1922 |
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IV: |
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Peters, 1877 |
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V: |
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Günther, 1858 |
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VI: |
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Taylor & Elbel, 1958 |
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VIII: |
Taylor & Elbel 1958; |
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Köhler, Vargas, Than & Thammachoti, 2021 |
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VIII: |
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Trageser, Al-Razi, Maria, Nobel, Asaduzzaman & Rahman, 2021 |
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VIII: |
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Lyu & Wang sp. nov. |
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VIII: |
This study |
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Peters, 1867 |
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VIII: |
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Smith, 1927 |
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Mertens, 1927 |
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Smith, 1927 |
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Iskandar, Arifin & Rachmanasah, 2011 |
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As the earliest introduced genus and also the type genus of
Despite the continuous controversy on the generic recognitions, taxonomic studies on species level of
In southeastern China where includes the subdivisions of Guangdong, Hong Kong, Macao, Fujian, and Hainan, two
Thirty-two newly collected samples of
Localities, vouchers, and GenBank accession numbers for all samples used in this study.
ID | Species | Voucher | Locality | 16s |
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1 |
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Alive individual | Indonesia: Java |
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2 |
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JBS 5405 | Myanmar: Mandalay: Na Htoe Gyi |
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3 |
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USNM 520376 | Myanmar: Sagaing: Chatthin |
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4 |
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SMF 103817 | Myanmar: Naypyidaw: near Yamethin |
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5 |
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CAS 213254 | Myanmar: Yangon: Hlaw Ga Park |
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6 |
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SMF 103815 | Myanmar: Yangon: near Taw Hlan village |
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7 |
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USNM 586924 | Myanmar: Tanintharyi: Yeybu village |
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8 |
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MNHN 0086Y | Thailand: Prachuap Khiri Khan: Huay Yang NP |
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9 |
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SMF GK7721 | Thailand: Rayong: near Rayong |
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10 |
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MNHN 2003.0327 | Cambodia: Pouthisat: Pneum Kravanh |
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11 |
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MNHN T2814 | Laos: Louangphrabang: Luang Prabang |
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12 | China: Guangdong: Mt Danxia |
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13 | China: Guangdong: Yingde |
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14 | China: Guangdong: Yingde |
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15 | China: Guangdong: Yingde |
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16 | China: Guangdong: Yingde |
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17 | China: Guangdong: Yingde |
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18 | China: Guangdong: Yingde |
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19 | China: Guangdong: Yingde |
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20 | China: Guangxi: Guiping |
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21 | China: Guangdong: Shenzhen |
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22 | China: Guangdong: Shenzhen |
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23 | China: Guangdong: Shenzhen |
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24 | China: Guangdong: Shenzhen |
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25 | China: Guangdong: Shenzhen |
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26 | China: Guangdong: Shenzhen |
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27 | China: Guangdong: Xinyi |
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28 | China: Guangdong: Xinyi |
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29 | China: Guangdong: Xinyi |
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30 | China: Guangdong: Zhanjiang |
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31 | China: Guangdong: Zhanjiang |
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32 | China: Guangdong: Zhuhai |
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33 | China: Guangdong: Zhuhai |
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34 | China: Hainan: Changjiang |
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35 | China: Hainan: Changjiang |
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36 | China: Hainan: Mt Diaoluo |
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37 | Not given | Malaysia: Kuala Lumpur |
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38 | USNM 586941 | Myanmar: Tanintharyi, Yeybu village |
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39 | USNM 586942 | Myanmar: Tanintharyi, Yeybu village |
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40 | CUMZ PT2634 | Thailand: Bangkok: Bangkok |
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41 | KUHE 19790 | Thailand: Chachoengso |
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42 | SMF GK7349 | Thailand: Chonburi: Ya Teng Homestay |
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43 | MNHN P324 | Thailand: Phangnga: Sa Nang Manora |
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44 | Alive idividual | Thailand: Ranong |
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45 | FMNH 266020 | Thailand: Sa Kaeo |
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46 | CUMZ PT2754 | Thailand: Songkhla: Wang Pha |
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47 | CUMZ PT2755 | Thailand: Songkhla: Wang Pha |
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48 | FMNH 268335 | Thailand: Surat Thani |
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49 | FMNH 268805 | Thailand: Krabi |
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50 | SMF GK7713 | Thailand: Trat: Ko Kut Resort |
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51 | SMF GK7695 | Thailand: Trat: Trat |
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52 | China: Yunnan: Jinghong |
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53 | China: Yunnan: Jinghong |
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54 | China: Yunnan: Mengla |
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55 | China: Yunnan: Mengla |
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56 | China: Yunnan: Mengla |
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57 | China: Yunnan: Menglian |
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58 | China: Yunnan: Menglian |
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59 | SCUM H020 | China |
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60 | MNHN 2004.0357 | Laos: Phongsali: Buon Tai |
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61 | CUMZ PT0167 | Thailand: Chaiyaphume: Ban Na Si Nuan |
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62 | CUMZ PT1543 | Thailand: Chiang Mai: Huai Hong Khrai |
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63 | CUMZ PT1544 | Thailand: Chiang Mai: Huai Hong Khrai |
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64 | MNHN 0119Y | Thailand: Phetchabun: Thung Salaeng Luang |
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65 | MNHN 0026Y | Thailand: Uttaradit: Nam Pad district |
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66 | JBS 19932 | Myanmar: Yangon: Mingalardon |
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67 | USNM 587386 | Myanmar: Yangon: Mingalardon |
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68 | USNM 587389 | Myanmar: Yangon: Mingalardon |
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69 |
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JnUZool-A0719 | Bangladesh: Chattogram |
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70 |
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JnUZool-A0819 | Bangladesh: Chattogram |
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71 |
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JnUZool-A0919 | Bangladesh: Chattogram |
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72 |
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JnUZool-A1019 | Bangladesh: Chattogram |
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73 |
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JnUZool-A1117 | Bangladesh: Chattogram |
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74 |
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MNHN 712D | Laos: Vientiane: Viangchan |
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75 |
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FMNH 255134 | Laos: Champasak: Mounlapamok |
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76 |
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NKMA 2196-15 | Thailand: Kalasin |
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77 |
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SMF GK7855 | Thailand: Nakhon Phanom: Ban Kan Luang |
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78 |
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SMF GK7396 | Thailand: Roi Et: near Ban Sa At Na Di |
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79 |
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SMF GK7532 | Thailand: Sakon Nakhon: Ban Phaeng Yai |
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80 |
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SMF GK7533 | Thailand: Sakon Nakhon: Ban Phaeng Yai |
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81 |
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FMNH 261789 | Cambodia: Koh Kong |
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82 |
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USNM 587105 | Myanmar: Bago: Dawei |
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83 |
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USNM 587107 | Myanmar: Bago: Dawei |
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84 |
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SMF 103800 | Myanmar: Magwe: near Taungdwingyi |
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85 |
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SMF 103840 | Myanmar: Yangon: East Yangon University |
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86 |
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SMF 103841 | Myanmar: Yangon: East Yangon University |
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87 |
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USNM 587402 | Myanmar: Yangon: Mingalardon |
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88 |
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NNU 202103284 | China: Guangxi: Mt Shiwandashan |
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89 |
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NNU 202103285 | China: Guangxi: Mt Shiwandashan |
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90 |
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NNU 202103320 | China: Guangxi: Mt Shiwandashan |
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91 |
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NNU 202103321 | China: Guangxi: Mt Shiwandashan |
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92 |
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KUHE 17327 | Malaysia: Sarawak: Matang |
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93 |
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KUHE 17327 | Malaysia: Sarawak: Matang |
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94 |
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KUHE 17327 | Malaysia: Sarawak: Matang |
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95 | NMBE 1064176 | Malaysia: Sarawak: Kubah NP |
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96 | NMBE 1065363 | Malaysia: Sarawak: Gunung Penrissen |
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97 | NMBE 1061694 | Malaysia: Sarawak: Gunung Murud |
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98 | NMBE 1066057 | Malaysia: Sarawak: Payeh Maga |
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99 | NMBE 1069835 | Malaysia: Sarawak: Usun Apau |
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100 | NMBE 1074010 | Malaysia: Sarawak: Batang Ai |
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101 | NMBE 1072307 | Malaysia: Sabah: Danum Valley Conservation Area |
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102 | FMNH 234895 | Malaysia: Sabah: Sipitang |
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103 | NMBE1072456 | Malaysia: Sabah: Tawau Hills NP |
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104 | NMBE 1056418 | Malaysia: Sarawak: Gunung Mulu NP |
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105 | KU 310493 | Philippinen: Eastern Samar: Taft |
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106 | FMNH 259486 | Philippinen: Kalinga: Balbalasang |
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107 | KU 306652 | Philippinen: Negros Oriental: Valencia |
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108 | PNM 7446 | Philippinen: Quezon: Lao |
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109 | KU 302276 | Philippinen: Romblon: Magdiwang |
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110 | EMD 424 | Philippines: Agusan Del Norte |
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111 | PNM ACD2011 | Philippines: Isabela |
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112 | KU 302322 | Philippines: Oriental Mindoro |
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113 | KU 308966 | Philippines: Palawan: Irawan |
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114 | PNM ACD5414 | Philippines: South Cotobato |
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115 | KU 306301 | Philippines: Western Samar |
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116 | KU 314471 | Philippines: Zamboanga: Pasonanca |
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117 | USNM 586929 | Myanmar: Tanintharyi: Yeybu village |
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118 | USNM 586928 | Myanmar: Tanintharyi: Yeybu village |
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119 | MZB Amph 16392 | Indonesia: Java |
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120 | MZB RMB2134 | Indonesia: Java |
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121 | Not given | Indonesia: Sumatra: Bukit Barisan Selatan NP |
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122 | Not given | Indonesia: Sumatra: Hutan Harapan |
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123 | Not given | Indonesia: Sumatra: Hutan Harapan |
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124 | FRIM 1136 | Malaysia: Pahang: Bukit Rengit |
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125 | FMNH 267890 | Malaysia: Sarawak: Bintulu |
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126 | FMNH 269753 | Malaysia: Sarawak: Binyo-Penyilam |
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127 | FRIM 1132 | Malaysia: Selangor: Kepong |
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128 | NMBE 1072541 | Malaysia: Sabah: Danum Valley Conservation Area |
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129 | FMNH 235605 | Malaysia: Sabah: Kota Marudu |
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130 | FMNH 242747 | Malaysia: Sabah: Sipitang |
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131 | FMNH273695 | Malaysia: Sarawak: Bintulu |
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132 | NMBE 1064771 | Malaysia: Sarawak: Gunung Mulu NP |
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133 | NMBE 1073926 | Malaysia: Sarawak: Payeh Maga |
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134 | Not given | Malaysia: Sarawak: Pelagus NP |
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135 | Not given | Malaysia: Sarawak: Pelagus NP |
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136 |
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KU 321225 | Philippines: Mindanao: Zamboanga: Pasonanca |
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137 |
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KU 321226 | Philippines: Mindanao: Zamboanga: Pasonanca |
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138 |
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KU 321227 | Philippines: Mindanao: Zamboanga: Pasonanca |
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139 |
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USNM 587302 | Myanmar: Mon: Kyaikhtiyo Wildlife Sanctuary |
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140 |
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USNM 586921 | Myanmar: Tanintharyi: Yeybu village |
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141 |
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China: Guangdong: Mt Danxia |
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142 |
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China: Tibet: Jiacha |
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143 |
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China: Guangxi: Mt Dayao |
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For phylogenetic analyses, 108 additional sequences of floating frogs species were obtained from GenBank and incorporated into our dataset (Table
Thirty-one newly collected samples of
External measurements were made on the examined specimens, using digital calipers (Neiko 01407A Stainless Steel 6-Inch Digital Caliper) to the nearest 0.1 mm, including snout–vent length (
The
As illustrated in the phylogenetics, all samples of genus
All samples previously recorded as
Phylogenetic trees of the genus
Map showing the floating frogs
All samples previously recorded as
The inner relationships for the other subclades of
In this work, it is the first time to integrate the molecular data of floating frogs published in recent years (e.g.
Particularly, since there are distinct morphological differences among some of these subclades (especially between the “
The species diversity of floating frogs is dramatically underestimated as indicated in our analysis. Deep divergences are presented among the samples currently identified as
Within subclade I (“
Within subclade VIII (“
Seven adult males and three adult females. Males
The specific name
“Chinese floating frog” in English / “中国浮蛙 (zhōng guó fú wā)” in Chinese.
(1) Body stocky, size small,
Based on the examined specimens (n=10). All specimens were similar in morphology. The measurements are given in Table
Body stocky, small-sized,
Measurements (in mm) of the examined specimens of
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Sex | Male | Male | Male | Male | Male | Male | Male | Female | Female | Female |
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24.2 | 24.7 | 27.5 | 24.3 | 26.2 | 27.0 | 24.8 | 32.2 | 32.0 | 31.5 |
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9.1 | 9.4 | 8.9 | 8.8 | 9.4 | 9.4 | 8.8 | 10.4 | 10.4 | 10.1 |
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8.8 | 9.6 | 9.7 | 8.6 | 9.4 | 9.3 | 8.6 | 11.0 | 11.4 | 10.0 |
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2.7 | 3.1 | 3.3 | 2.8 | 3.3 | 3.1 | 3.2 | 4.2 | 4.0 | 3.7 |
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1.5 | 1.3 | 1.6 | 1.4 | 1.4 | 1.5 | 1.2 | 1.6 | 1.5 | 1.6 |
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1.5 | 1.2 | 1.5 | 1.3 | 1.4 | 1.4 | 1.1 | 1.5 | 1.4 | 1.4 |
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3.2 | 2.8 | 2.7 | 2.7 | 2.8 | 2.9 | 2.6 | 3.1 | 3.0 | 3.2 |
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7.7 | 7.5 | 7.8 | 7.1 | 7.3 | 7.4 | 6.6 | 8.2 | 8.8 | 8.5 |
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4.0 | 4.5 | 3.6 | 4.1 | 4.4 | 4.0 | 3.9 | 4.9 | 5.1 | 4.9 |
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19.3 | 18.4 | 18.4 | 18.0 | 18.8 | 18.8 | 17.5 | 21.3 | 21.5 | 21.6 |
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11.3 | 11.4 | 11.2 | 11.1 | 11.7 | 11.5 | 10.9 | 13.4 | 13.6 | 12.8 |
Forelimbs short, lower arm 13–18% (n=10) of
Hind limbs robust, tibia 41–47% (n=8) of
Dorsal surface relatively rough, transverse wrinkles and dense tubercles on the dorsum, head, flanks, and limbs; small granules on the dorsal rears of hands and tarsi; not bearing spinules on the dorsal skin; supratympanic fold distinct, raised and granulated, extending from the posterior corner of the eye, curved on the temporal region, to the previous shoulder; dorsolateral fold absent. Ventral surface with large flattened tubercles, denser on the throat and thighs; dense granules on the ventral feet and tarsi.
In life (Fig.
In preservative (Fig.
Morphological features of
Morphological features of
Male with a single subgular vocal sac; in breeding season, a single, light grey nuptial pad on the dorsal surface of finger I, nuptial spinules invisible. Males (
This frog inhabits natural or artificial ponds and paddy fields in plain areas. They quickly dive underwater after being disturbed during the daytime, while become relatively insensitive at night. Males call in the water surface or waterside grass from dusk to dawn, more active during the rain. The breeding season is from April to August (this study;
This species was previously reported as common and widespread species in southeastern China under the nomen
The type specimen of
Moreover, there were two historic species currently regarded as synonymies of
Five adult males and five adult females. Male
The specific name
“Lingnan floating frog” in English / “岭南浮蛙 (lǐng nán fú wā)” in Chinese.
(1) Body stocky, size small,
For the remaining 13 congeners,
Morphological features of the adult male holotype
Forelimbs short, lower arm 15% of
Hind limbs robust, tibia 44% of
Dorsal surface relatively rough, large tubercles scattering on skin of dorsum, flanks, and dorsal limbs, not bearing spinules on the dorsal skin; a faint fold across head between orbits; supratympanic fold distinct, raised, extending from the posterior corner of the eye, slightly curved on the temporal region, to the previous shoulder; dorsolateral fold absent. Ventral surface with large flattened tubercles; a fold across breast; dense granules on the ventral tarsi.
In life (Fig.
In preservative, dorsum light gray; black speckles on dorsum and transverse bars on limbs light brown; mid-dorsal stripe grayish white and more distinct; nuptial pad light gray, slightly transparent; ventral surface grayish white; mottling on throat gray white.
The measurements of the type series are given in Table
Color variation of
Measurements (in mm) of the examined specimens of
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Sex | Male | Male | Male | Male | Male | Male | Male | Male | Female | Female | Female |
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22.1 | 21.6 | 21.4 | 21.5 | 19.9 | 20.2 | 21.8 | 21.7 | 26.8 | 28.5 | 28.8 |
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7.8 | 7.1 | 7.5 | 7.5 | 6.6 | 6.6 | 7.1 | 7.8 | 9.0 | 9.5 | 9.0 |
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7.4 | 6.7 | 7.2 | 7.2 | 6.5 | 6.5 | 6.6 | 7.5 | 8.9 | 9.3 | 9.0 |
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2.9 | 2.7 | 2.5 | 2.6 | 2.3 | 2.3 | 2.9 | 2.9 | 3.0 | 3.5 | 3.3 |
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1.9 | 1.8 | 1.6 | 1.6 | 1.5 | 1.6 | 1.7 | 1.8 | 2.1 | 2.2 | 2.3 |
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1.5 | 1.4 | 1.1 | 1.2 | 1.0 | 1.1 | 1.3 | 1.3 | 1.7 | 1.8 | 1.9 |
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2.6 | 2.5 | 2.5 | 2.5 | 2.3 | 2.3 | 2.5 | 2.7 | 3.3 | 3.4 | 3.0 |
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5.4 | 5.3 | 5.3 | 5.3 | 4.7 | 4.6 | 5.5 | 5.4 | 5.6 | 6.8 | 6.1 |
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3.7 | 3.5 | 3.2 | 3.4 | 3.5 | 3.7 | 3.4 | 3.3 | 4.2 | 4.4 | 4.8 |
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14.1 | 14.4 | 13.4 | 14.2 | 12.9 | 13.8 | 14.7 | 14.1 | 17.9 | 19.2 | 19.5 |
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9.8 | 9.7 | 9.4 | 9.8 | 8.6 | 9.2 | 9.5 | 9.5 | 11.7 | 11.4 | 11.8 |
Male with a single subgular vocal sac; in breeding season, a single, light yellow, swollen, and granular nuptial pad on the dorsal surface of finger I, nuptial spinules invisible. Males (
This frog inhabits natural or artificial ponds and paddy fields in hilly regions. Males call in the water surface or waterside grass from dusk to dawn. The breeding season is from May to August (this study;
We thank Guangdong Yunkaishan National Nature Reserve, Guangdong Nanling National Nature Reserve, Guangdong Shimentai National Nature Reserve, Guangdong Danxiashan National Nature Reserve, Qing Du, Wei-Liang Xie, Run-Lin Li, Hai-Long He, Yong-You Zhao, and Hong-Hui Chen, for their help in the fieldwork, lab work, and manuscript preparation. We thank Gunther Köhler for his useful comments on our manuscript. This research was supported by the DFGP Project of Fauna of Guangdong-202115, the National Animal Collection Resource Center, China, the Project of Animal Diversity Survey and Monitoring System Construction of Guangdong Shimentai National Nature Reserve, and the Project of Survey of Terrestrial Vertebrate Diversity in Guangdong Danxiashan National Nature Reserve.
Table S1