Research Article |
Corresponding author: Ying-Yong Wang ( wangyy@mail.sysu.edu.cn ) Academic editor: Angelica Crottini
© 2020 Zhi-Tong Lyu, Yuan-Qiu Li, Zhao-Chi Zeng, Jian Zhao, Zu-Yao Liu, Guo-Xin Guo, Ying-Yong Wang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lyu Z-T, Li Y-Q, Zeng Z-C, Zhao J, Liu Z-Y, Guo G-X, Wang Y-Y (2020) Four new species of Asian horned toads (Anura, Megophryidae, Megophrys) from southern China. ZooKeys 942: 105-140. https://doi.org/10.3897/zookeys.942.47983
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Recent phylogenetic analysis encompassing multilocus nuclear-gene and matrilineal mtDNA genealogy has revealed a series of cryptic species of the subgenus Panophrys within genus Megophrys from southern and eastern China. This study demonstrates that the Panophrys specimens from the hilly areas among Guangdong, Guangxi and Hunan can be morphologically distinguished from all recognized congeners, thereby providing additional supports for the recognitions of four new species of Panophrys, namely Megophrys (Panophrys) mirabilis Lyu, Wang & Zhao, sp. nov. from northeastern Guangxi, Megophrys (Panophrys) shimentaina Lyu, Liu & Wang, sp. nov. from northern Guangdong, and Megophrys (Panophrys) xiangnanensis Lyu, Zeng & Wang, sp. nov. and Megophrys (Panophrys) yangmingensis Lyu, Zeng & Wang, sp. nov. from southern Hunan. The descriptions of these species take the number of Megophrys species to 101, 46 of which belong to the subgenus Panophrys.
cryptic species, diversity, morphology, Panophrys, taxonomy
The Asian horned toad genus Megophrys Kuhl & Van Hasselt, 1822 within the family Megophryidae Bonaparte, 1850, is a typical representative for Oriental fauna, spreading throughout southern China, southern and eastern Himalayas, across Indochina to islands of the Sunda Shelf and the Philippines (
During our herpetological surveys in the hilly areas among Guangdong, Guangxi and Hunan, southern China (Fig.
Taxonomic system. The higher systematics of Asian horned toads has been in intensive debates for decades (
Phylogeny. Two mitochondrial genes, namely partial 16S ribosomal RNA gene (16S) and partial cytochrome C oxidase 1 gene (CO1), were used for phylogenetic analysis. All sequences were attained from GenBank, encompassing 17 samples of the unnamed species (originally submitted by
Localities, voucher information, and GenBank numbers for all samples used in this study.
ID | Species | Localities | Voucher number | 16s | co1 |
---|---|---|---|---|---|
Megophrys (Panophrys) | |||||
1 | M. (Pa.) mirabilis sp. nov. | China: Guangxi: Huaping Nature Reserve |
|
MH406669 | MH406109 |
2 | M. (Pa.) mirabilis sp. nov. | China: Guangxi: Huaping Nature Reserve |
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MH406670 | MH406110 |
3 | M. (Pa.) mirabilis sp. nov. | China: Guangxi: Huaping Nature Reserve |
|
MH406681 | MH406127 |
4 | M. (Pa.) mirabilis sp. nov. | China: Guangxi: Huaping Nature Reserve |
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MH406724 | MH406176 |
5 | M. (Pa.) shimentaina sp. nov. | China: Guangdong: Shimentai Nature Reserve |
|
MH406655 | MH406092 |
6 | M. (Pa.) shimentaina sp. nov. | China: Guangdong: Shimentai Nature Reserve |
|
MH406656 | MH406093 |
7 | M. (Pa.) shimentaina sp. nov. | China: Guangdong: Shimentai Nature Reserve |
|
MH406787 | MH406249 |
8 | M. (Pa.) shimentaina sp. nov. | China: Guangdong: Shimentai Nature Reserve |
|
MH406788 | MH406250 |
9 | M. (Pa.) xiangnanensis sp. nov. | China: Hunan: Mt Yangming |
|
MH406713 | MH406165 |
10 | M. (Pa.) xiangnanensis sp. nov. | China: Hunan: Mt Yangming |
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MH406714 | MH406166 |
11 | M. (Pa.) xiangnanensis sp. nov. | China: Hunan: Mt Yangming |
|
MH406715 | MH406167 |
12 | M. (Pa.) xiangnanensis sp. nov. | China: Hunan: Mt Yangming |
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MH406717 | MH406169 |
13 | M. (Pa.) xiangnanensis sp. nov. | China: Hunan: Mt Yangming |
|
MH406718 | MH406170 |
14 | M. (Pa.) yangmingensis sp. nov. | China: Hunan: Mt Yangming |
|
MH406716 | MH406168 |
15 | M. (Pa.) yangmingensis sp. nov. | China: Hunan: Mt Yangming |
|
MH406719 | MH406171 |
16 | M. (Pa.) yangmingensis sp. nov. | China: Hunan: Mt Yangming |
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MH406720 | MH406172 |
17 | M. (Pa.) yangmingensis sp. nov. | China: Hunan: Mt Yangming |
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MH406721 | MH406173 |
18 | M. (Pa.) acuta | China: Guangdong: Heishiding Nature Reserve |
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KJ579119 | MH406122 |
19 | M. (Pa.) baolongensis | China: Chongqing: Baolong Town | KIZ 019216 | KX811813 | KX812093 |
20 | M. (Pa.) binchuanensis | China: Yunnan: Mt. Jizu | KIZ 019441 | KX811849 | KX812112 |
21 | M. (Pa.) binlingensis | China: Sichuan: Mt. Wawu |
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MH406892 | MH406354 |
22 | M. (Pa.) boettgeri | China: Fujian: Mt. Wuyi |
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MF667878 | MH406247 |
23 | M. (Pa.) brachykolos | China: Hong Kong |
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KJ560403 | MH406120 |
24 | M. (Pa.) caudoprocta | China: Hunan: Badagongshan Nature Reserve |
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MH406795 | MH406257 |
25 | M. (Pa.) cheni | China: Hunan: Taoyuandong Nature Reserve |
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KJ560398 | MH406098 |
26 | M. (Pa.) daweimontis | China: Yunnan: Mt. Dawei | KIZ 048997 | KX811867 | KX812125 |
27 | M. (Pa.) dongguanensis | China: Guangdong: Mt. Yinping |
|
MH406647 | MH406083 |
28 | M. (Pa.) fansipanensis | Vietnam: Lao Cai: Sa Pa | VNMN 2018.01 | MH514886 | / |
29 | M. (Pa.) hoanglienensis | Vietnam: Lao Cai: Sa Pa | VNMN 07034 | MH514890 | / |
30 | M. (Pa.) huangshanensis | China: Anhui: Mt. Huangshan |
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MF667883 | MH406161 |
31 | M. (Pa.) insularis | China: Guangdong: Nan’ao Island |
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MF667887 | MF667924 |
32 | M. (Pa.) jiangi | China: Guizhou: Kuankuoshui Nature Reserve |
|
MN107743 | MN107748 |
33 | M. (Pa.) jingdongensis | China: Yunnan: Mt. Wuliang |
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MH406773 | MH406232 |
34 | M. (Pa.) jinggangensis | China: Jiangxi: Mt. Jinggang |
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MH406780 | MH406239 |
35 | M. (Pa.) jiulianensis | China: Jiangxi: Mt. Jiulian |
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MH406791 | MH406253 |
36 | M. (Pa.) kuatunensis | China: Fujian: Mt. Wuyi |
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MF667881 | MH406206 |
37 | M. (Pa.) leishanensis | China: Guizhou: Mt. Leigong | SYSa002213 | MH406673 | MH406113 |
38 | M. (Pa.) liboensis | China: Guizhou: Libo Country | 20150813001 | MF285253 | / |
39 | M. (Pa.) lini | China: Hunan: Taoyuandong Nature Reserve |
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MF667874 | MH406135 |
40 | M. (Pa.) lishuiensis | China: Zhejiang: Lishui City | WYF00169 | KY021418 | / |
41 | M. (Pa.) minor | China: Sichuan: Mt. Qingcheng |
|
MF667862 | MH406194 |
42 | M. (Pa.) mufumontana | China: Hunan: Mt. Mufu |
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MK524104 | MK524135 |
43 | M. (Pa.) nankunensis | China: Guangdong: Mt. Nankun |
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MH406822 | MH406284 |
44 | M. (Pa.) nanlingensis | China: Guangdong: Nanling Nature Reserve |
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MH406646 | MH406082 |
45 | M. (Pa.) obesa | China: Guangdong: Heishiding Nature Reserve |
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KJ579121 | MH406123 |
46 | M. (Pa.) ombrophila | China: Fujian: Mt. Wuyi | WUYI2015101 | KX856397 | / |
47 | M. (Pa.) omeimontis | China: Sichuan: Mt. Emei |
|
MH406887 | MH406349 |
48 | M. (Pa.) palpebralespinosa | Vietnam: Thanh Hoa: Pu Hu Nature Reserve | KIZ 011650 | KX811889 | KX812138 |
49 | M. (Pa.) rubrimera | Vietnam: Lao Cai: Sa Pa | VNMN 2017.002 | MF536420 | / |
50 | M. (Pa.) sangzhiensis | China: Hunan: Badagongshan Nature Reserve |
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MH406797 | MH406259 |
51 | M. (Pa.) shunhuangensis | China: Hunan: Nanshan Forest Park | HNNU 18NS01 | MK836023 | MK977594 |
52 | M. (Pa.) spinata | China: Guizhou: Mt. Leigong |
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MH406675 | MH406115 |
53 | M. (Pa.) tuberogranulatus | China: Hunan: Badagongshan Nature Reserve |
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MH406801 | MH406263 |
54 | M. (Pa.) wugongensis | China: Jiangxi: Mt. Wugong |
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MH406853 | MH406315 |
55 | M. (Pa.) wuliangshanensis | China: Yunnan: Mt. Wuliang |
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MH406771 | MH406230 |
56 | M. (Pa.) wushanensis | China: Hubei: Shennongjia Nature Reserve |
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MH406732 | MH406184 |
57 | M. (Pa.) xianjuensis | China: Zhejiang: Xianju County |
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MN563753 | MN563769 |
Megophrys (Xenophrys) | |||||
58 | M. (X.) glandulosa | China: Yunnan: Mt. Gaoligong |
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MH406755 | MH406214 |
59 | M. (X.) mangshanensis | China: Guangdong: Mt. Sanyue |
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MH406666 | MH406106 |
DNA sequences were aligned by the Clustal W algorithm with default parameters (Thompson et al. 1997) and trimmed with the gaps partially deleted in MEGA 6 (Tamura et al. 2013). Two gene segments, 632 base pairs (bp) of CO1 and 541 bp of16S, were concatenated seriatim into a 1173-bp sequence, and were further tested in jmodeltest v2.1.2 with Akaike and Bayesian information criteria, all resulting the best-fitting nucleotide substitution models of GTR+I+G. Sequenced data was analyzed using Bayesian inference (BI) in MrBayes 3.2.4 (Ronquist et al. 2012). Two independent runs were conducted in a BI analysis, each of which was performed for 10,000,000 generations and sampled every 1000 generations with the first 25% samples were discarded as burn-in, resulting a potential scale reduction factor (PSRF) of < 0.005. Mean genetic distances of 16S gene between and within species were calculated in MEGA 6 using the uncorrected p-distance model.
Bioacoustics. Advertisement calls of the unnamed species were recorded in the field by a SONY PCM-D50 digital sound recorder. The sound files in wave format were sampled at 48 kHz with 24 bits in depth. Raven pro 1.5 (Cornell Lab of Ornithology, 2003–2014) was used to output the spectrograms and to measure interrelated parameters with Fast Fourier transform of 256 points and a 50% overlap. The following measurements were performed: call/note duration (the difference between begin time and end time for a selected call/note), notes per call, inter-note intervals (the difference between end time for a selected note and begin time for the next selected note), peak frequency (the frequency at which peak power occurs within the selected call), high frequency (the highest frequency of the selected call), low frequency (the lowest frequency of the selected call), bandwidth 90% (the difference between the 5% and 95% frequencies of a selected call).
Morphology. Thirty-six unnamed specimens from the hilly areas among Guangdong, Guangxi and Hunan, southern China were examined, 17 of which have been used in the phylogenetic analysis. All examined specimens were fixed in 10% buffered formalin and later transferred to 70% ethanol. All studied specimens are deposited in The Museum of Biology, Sun Yat-sen University (
External measurements were made for the unnamed specimens with digital calipers (Neiko 01407A Stainless Steel 6-Inch Digital Caliper, USA) to the nearest 0.1 mm. Mean and standard deviation (SD) were calculated in R 3.3.2 (R Core Team 2016). These measurements were as follows:
ED eye diameter (from the anterior corner of the eye to posterior corner of the eye);
FTL foot length (from distal end of shank to the tip of digit IV);
HDL head length (from tip of snout to the articulation of the jaw);
HDW head width (head width at the commissure of the jaws);
HND hand length (from the proximal border of the outer palmar tubercle to the tip of digit III);
IND internasal distance (distance between nares);
IOD interorbital distance (minimum distance between upper eyelids);
RAD radio-ulna length (from the flexed elbow to the proximal border of the outer palmar tubercle);
SNT snout length (from tip of snout to the anterior corner of the eye);
SVL snout-vent length (from tip of snout to posterior margin of vent);
TD tympanum diameter (horizontal diameter of tympanum);
TED tympanum-eye distance (from anterior edge of tympanum to posterior corner of the eye);
TIB tibial length (from the outer surface of the flexed knee to the heel).
Sex was determined by secondary sexual characters, i.e., the presence of vocal sac, nuptial pads/spines in males (Fei et al. 2016).
Morphological characters of all 42 recognized congeners of subgenus Panophrys for comparisons were based on the examination of museum specimens listed in Appendix I and on information available in the literature (Table
Literature for morphological characters of 42 recognized species of Megophrys (Panophrys).
ID | Species | References |
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1 | M. (Pa.) acuta Wang, Li & Jin, 2014 |
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2 | M. (Pa.) baolongensis Ye, Fei & Xie, 2007 |
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3 | M. (Pa.) binchuanensis Ye & Fei, 1995 |
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4 | M. (Pa.) binlingensis Jiang, Fei & Ye, 2009 |
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5 | M. (Pa.) boettgeri (Boulenger, 1899) |
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6 | M. (Pa.) brachykolos Inger & Romer, 1961 |
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7 | M. (Pa.) caudoprocta Shen, 1994 |
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8 | M. (Pa.) cheni (Wang & Liu, 2014) |
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9 | M. (Pa.) daweimontis Rao & Yang, 1997 |
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10 | M. (Pa.) dongguanensis Wang & Wang |
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11 | M. (Pa.) fansipanensis Tapley, Cutajar, Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Portway, Luong & Rowley, 2018 |
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12 | M. (Pa.) hoanglienensis Tapley, Cutajar, Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Portway, Luong & Rowley, 2018 |
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13 | M. (Pa.) huangshanensis Fei & Ye, 2005 |
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14 | M. (Pa.) insularis (Wang, Liu, Lyu, Zeng & Wang, 2017) |
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15 | M. (Pa.) jiangi Liu, Li, Wei, Xu, Cheng, Wang & Wu, 2020 |
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16 | M. (Pa.) jingdongensis Fei & Ye, 1983 |
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17 | M. (Pa.) jinggangensis (Wang, 2012) |
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18 | M. (Pa.) jiulianensis Wang, Zeng, Lyu & Wang |
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19 | M. (Pa.) kuatunensis Pope, 1929 |
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20 | M. (Pa.) latidactyla Orlov, Poyarkov & Nguyen, 2015 |
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21 | M. (Pa.) leishanensis Li, Xu, Liu, Jiang, Wei & Wang, 2018 |
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22 | M. (Pa.) liboensis (Zhang, Li, Xiao, Li, Pan, Wang, Zhang & Zhou, 2017) |
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23 | M. (Pa.) lini (Wang & Yang, 2014) |
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24 | M. (Pa.) lishuiensis (Wang, Liu & Jiang, 2017) |
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25 | M. (Pa.) minor Stejneger, 1926 |
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26 | M. (Pa.) mufumontana Wang, Lyu & Wang |
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27 | M. (Pa.) nankunensis Wang, Zeng & Wang |
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28 | M. (Pa.) nanlingensis Lyu, Wang, Liu & Wang |
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29 | M. (Pa.) obesa Wang, Li & Zhao, 2014 |
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30 | M. (Pa.) ombrophila Messenger & Dahn, 2019 |
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31 | M. (Pa.) omeimontis Liu, 1950 |
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32 | M. (Pa.) palpebralespinosa Bourret, 1937 |
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33 | M. (Pa.) robrimera Tapley, Cutajar, Mahony, Chung, Dau, Nguyen, Luong & Rowley, 2017 |
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34 | M. (Pa.) sangzhiensis Jiang, Ye & Fei, 2008 |
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35 | M. (Pa.) shuichengensis Tian & Sun, 1995 |
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36 | M. (Pa.) shunhuangensis Wang, Deng, Liu, Wu & Liu, 2019 |
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37 | M. (Pa.) spinata Liu & Hu, 1973 |
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38 | M. (Pa.) tuberogranulatus Shen, Mo & Li, 2010 |
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39 | M. (Pa.) wugongensis Wang, Lyu & Wang |
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40 | M. (Pa.) wuliangshanensis Ye & Fei, 1995 |
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41 | M. (Pa.) wushanensis Ye & Fei, 1995 |
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42 | M. (Pa.) xianjuensis Wang, Wu, Peng, Shi, Lu & Wu, 2020 |
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The BI phylogenetic result is shown in Fig.
The unnamed samples from Huaping Nature Reserve, Guangxi (samples ID 1–4 in Table
The samples from Shimentai Nature Reserve, Guangxi (samples ID 5–8 in Table
The other samples from Mt Yangming, Hunan (samples ID 9–13 in Table
Mean p-distance of 16S gene among the Megophrys (Panophrys) species used in this study.
ID | Species | 1– 4 | 5– 8 | 9– 13 | 14–17 | 18 | 19 | 20 | 21 | 22 | 23 | 24 | 25 | 26 | 27 | 28 | 29 | 30 | 31 | 32 | 33 | 34 | 35 | 36 | 37 | 38 | 39 | 40 | 41 | 42 | 43 | 44 | 45 | 46 | 47 | 48 | 49 | 50 | 51 | 52 | 53 | 54 | 55 | 56 | 57 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1– 4 | M. (Pa.) mirabilis sp. nov. | 0.0 | |||||||||||||||||||||||||||||||||||||||||||
5– 8 | M. (Pa.) shimentaina sp. nov. | 3.8 | 0.0 | ||||||||||||||||||||||||||||||||||||||||||
9– 13 | M. (Pa.) xiangnanensis sp. nov. | 3.9 | 4.1 | 0.0 | |||||||||||||||||||||||||||||||||||||||||
14–17 | M. (Pa.) yangmingensis sp. nov. | 3.4 | 3.4 | 4.3 | 0.3 | ||||||||||||||||||||||||||||||||||||||||
18 | M. (Pa.) acuta | 7.3 | 8.8 | 7.3 | 7.3 | / | |||||||||||||||||||||||||||||||||||||||
19 | M. (Pa.) baolongensis | 3.6 | 4.0 | 4.4 | 2.9 | 7.3 | / | ||||||||||||||||||||||||||||||||||||||
20 | M. (Pa.) binchuanensis | 4.8 | 5.4 | 4.8 | 4.9 | 8.1 | 4.4 | / | |||||||||||||||||||||||||||||||||||||
21 | M. (Pa.) binlingensis | 4.9 | 5.1 | 5.0 | 4.3 | 8.9 | 3.8 | 2.5 | / | ||||||||||||||||||||||||||||||||||||
22 | M. (Pa.) boettgeri | 2.4 | 3.4 | 3.9 | 2.1 | 7.0 | 2.7 | 4.0 | 3.7 | / | |||||||||||||||||||||||||||||||||||
23 | M. (Pa.) brachykolos | 5.4 | 5.5 | 3.9 | 5.2 | 8.1 | 4.6 | 5.3 | 6.5 | 4.7 | / | ||||||||||||||||||||||||||||||||||
24 | M. (Pa.) caudoprocta | 3.5 | 4.3 | 5.0 | 3.2 | 7.3 | 2.5 | 4.4 | 3.7 | 2.4 | 5.8 | / | |||||||||||||||||||||||||||||||||
25 | M. (Pa.) cheni | 3.5 | 4.0 | 3.0 | 3.5 | 7.8 | 2.7 | 3.8 | 4.3 | 2.6 | 4.3 | 3.9 | / | ||||||||||||||||||||||||||||||||
26 | M. (Pa.) daweimontis | 4.4 | 5.4 | 4.2 | 4.4 | 7.0 | 4.0 | 2.8 | 2.9 | 3.4 | 6.3 | 4.0 | 3.6 | / | |||||||||||||||||||||||||||||||
27 | M. (Pa.) dongguanensis | 5.2 | 4.7 | 4.3 | 4.9 | 8.1 | 4.0 | 5.3 | 5.2 | 4.1 | 5.8 | 5.0 | 2.8 | 4.6 | / | ||||||||||||||||||||||||||||||
28 | M. (Pa.) fansipanensis | 5.1 | 5.0 | 4.7 | 4.6 | 10.5 | 4.0 | 4.6 | 4.2 | 4.2 | 5.5 | 4.6 | 4.4 | 4.6 | 5.7 | / | |||||||||||||||||||||||||||||
29 | M. (Pa.) hoanglienensis | 4.6 | 5.0 | 4.7 | 4.0 | 8.4 | 3.8 | 3.2 | 2.9 | 3.6 | 5.7 | 4.0 | 4.0 | 3.6 | 4.9 | 2.5 | / | ||||||||||||||||||||||||||||
30 | M. (Pa.) huangshanensis | 2.8 | 3.8 | 4.3 | 2.6 | 7.0 | 3.0 | 4.4 | 4.1 | 0.7 | 5.0 | 2.8 | 3.0 | 3.6 | 4.5 | 4.6 | 4.0 | / | |||||||||||||||||||||||||||
31 | M. (Pa.) insularis | 5.2 | 4.5 | 4.1 | 4.5 | 9.2 | 4.2 | 5.7 | 5.2 | 4.5 | 5.6 | 5.2 | 2.8 | 5.5 | 4.3 | 4.5 | 4.7 | 4.9 | / | ||||||||||||||||||||||||||
32 | M. (Pa.) jiangi | 7.3 | 8.3 | 7.9 | 7.5 | 12.0 | 7.6 | 8.4 | 8.7 | 7.1 | 8.6 | 8.1 | 7.7 | 8.2 | 8.8 | 8.0 | 7.8 | 7.9 | 8.6 | / | |||||||||||||||||||||||||
33 | M. (Pa.) jingdongensis | 4.5 | 5.1 | 4.9 | 4.7 | 8.6 | 4.2 | 2.9 | 2.6 | 4.1 | 6.7 | 3.7 | 4.1 | 2.5 | 6.0 | 4.0 | 4.0 | 4.5 | 5.8 | 8.6 | / | ||||||||||||||||||||||||
34 | M. (Pa.) jinggangensis | 3.5 | 4.9 | 3.0 | 3.7 | 7.3 | 3.6 | 4.2 | 4.1 | 3.2 | 5.0 | 4.3 | 3.0 | 4.0 | 5.0 | 4.6 | 4.4 | 3.6 | 4.1 | 7.0 | 4.3 | / | |||||||||||||||||||||||
35 | M. (Pa.) jiulianensis | 2.8 | 3.8 | 4.3 | 2.8 | 8.1 | 2.9 | 4.9 | 4.3 | 2.2 | 5.2 | 3.0 | 3.5 | 4.8 | 4.5 | 3.8 | 3.2 | 2.6 | 4.5 | 8.1 | 5.0 | 3.9 | / | ||||||||||||||||||||||
36 | M. (Pa.) kuatunensis | 3.7 | 3.8 | 3.4 | 3.6 | 8.1 | 3.0 | 4.6 | 4.7 | 2.8 | 4.1 | 3.7 | 2.4 | 4.0 | 2.8 | 4.4 | 4.2 | 3.2 | 3.4 | 8.1 | 4.9 | 3.7 | 3.6 | / | |||||||||||||||||||||
37 | M. (Pa.) leishanensis | 2.4 | 3.6 | 4.1 | 2.6 | 7.3 | 2.5 | 3.4 | 3.4 | 1.7 | 4.9 | 2.8 | 3.0 | 3.4 | 4.7 | 3.6 | 3.4 | 2.1 | 4.7 | 6.6 | 3.5 | 3.4 | 2.4 | 3.4 | / | ||||||||||||||||||||
38 | M. (Pa.) liboensis | 3.4 | 3.8 | 4.7 | 3.2 | 7.8 | 3.2 | 4.9 | 4.9 | 1.7 | 4.6 | 3.2 | 3.5 | 4.4 | 4.7 | 4.7 | 4.2 | 2.1 | 5.0 | 7.7 | 5.2 | 4.3 | 3.0 | 3.0 | 2.8 | / | |||||||||||||||||||
39 | M. (Pa.) lini | 4.1 | 4.7 | 2.2 | 4.5 | 7.0 | 4.7 | 4.9 | 5.2 | 3.4 | 3.9 | 4.7 | 3.0 | 4.4 | 4.1 | 5.5 | 5.3 | 3.7 | 4.3 | 8.1 | 5.0 | 3.6 | 4.5 | 3.2 | 3.9 | 4.3 | / | ||||||||||||||||||
40 | M. (Pa.) lishuiensis | 5.0 | 6.9 | 5.8 | 5.0 | 8.4 | 4.5 | 6.3 | 6.3 | 3.8 | 6.8 | 5.3 | 4.8 | 6.0 | 5.8 | 6.5 | 5.3 | 4.8 | 5.3 | 10.6 | 7.8 | 6.0 | 4.8 | 4.5 | 4.5 | 4.5 | 6.0 | / | |||||||||||||||||
41 | M. (Pa.) minor | 5.6 | 6.2 | 6.2 | 5.1 | 9.5 | 6.1 | 5.5 | 5.6 | 4.5 | 6.4 | 5.6 | 5.4 | 5.1 | 6.4 | 5.5 | 5.1 | 5.1 | 7.1 | 7.0 | 5.8 | 5.4 | 6.0 | 5.4 | 5.1 | 5.4 | 6.2 | 7.6 | / | ||||||||||||||||
42 | M. (Pa.) mufumontana | 3.8 | 4.2 | 4.7 | 2.6 | 8.6 | 3.2 | 4.6 | 4.3 | 2.4 | 5.4 | 3.2 | 3.6 | 4.4 | 5.1 | 4.5 | 4.0 | 3.0 | 4.5 | 7.2 | 4.7 | 3.4 | 3.2 | 3.4 | 2.6 | 3.4 | 4.9 | 5.0 | 5.5 | / | |||||||||||||||
43 | M. (Pa.) nankunensis | 4.5 | 4.3 | 4.1 | 4.3 | 8.9 | 3.2 | 5.5 | 5.0 | 3.9 | 5.4 | 4.3 | 2.2 | 4.6 | 2.6 | 5.3 | 5.1 | 4.3 | 4.1 | 8.6 | 5.2 | 4.5 | 4.3 | 3.0 | 4.1 | 4.7 | 4.3 | 5.8 | 6.7 | 4.5 | / | ||||||||||||||
44 | M. (Pa.) nanlingensis | 4.9 | 5.8 | 3.4 | 4.9 | 7.8 | 5.3 | 5.7 | 5.2 | 4.3 | 5.0 | 6.0 | 3.9 | 5.3 | 4.9 | 5.5 | 5.3 | 4.7 | 4.5 | 8.5 | 6.0 | 4.7 | 5.0 | 4.5 | 4.5 | 5.8 | 3.6 | 6.3 | 6.0 | 5.3 | 5.0 | / | |||||||||||||
45 | M. (Pa.) obesa | 6.2 | 7.1 | 5.4 | 6.8 | 8.4 | 6.8 | 7.8 | 8.1 | 5.1 | 7.8 | 7.6 | 3.2 | 5.9 | 5.1 | 7.6 | 7.6 | 5.7 | 5.4 | 11.7 | 7.0 | 6.5 | 6.8 | 4.6 | 5.9 | 6.5 | 5.1 | 5.7 | 9.2 | 7.6 | 4.9 | 6.5 | / | ||||||||||||
46 | M. (Pa.) ombrophila | 5.2 | 5.8 | 5.0 | 5.2 | 9.7 | 4.9 | 5.9 | 6.2 | 4.5 | 6.0 | 6.0 | 3.2 | 5.3 | 4.7 | 5.5 | 5.5 | 4.9 | 4.5 | 8.4 | 6.0 | 5.2 | 5.2 | 4.7 | 4.3 | 5.6 | 5.2 | 4.8 | 7.1 | 5.4 | 4.7 | 6.0 | 4.1 | / | |||||||||||
47 | M. (Pa.) omeimontis | 4.7 | 5.1 | 3.9 | 4.5 | 9.2 | 4.6 | 2.7 | 2.4 | 3.9 | 5.8 | 4.3 | 3.9 | 2.7 | 5.0 | 3.6 | 3.0 | 4.3 | 5.2 | 9.0 | 2.4 | 4.5 | 4.5 | 4.3 | 3.6 | 4.7 | 4.5 | 6.1 | 5.6 | 4.5 | 4.7 | 5.2 | 6.8 | 5.4 | / | ||||||||||
48 | M. (Pa.) palpebralespinosa | 5.7 | 6.7 | 5.5 | 6.1 | 8.4 | 4.9 | 3.6 | 4.0 | 4.9 | 6.6 | 4.8 | 5.1 | 3.6 | 6.3 | 6.1 | 5.1 | 4.9 | 7.0 | 9.5 | 4.4 | 5.7 | 6.3 | 5.1 | 4.4 | 5.9 | 4.7 | 7.3 | 5.7 | 5.9 | 6.1 | 6.3 | 9.2 | 6.8 | 4.0 | / | |||||||||
49 | M. (Pa.) rubrimera | 4.7 | 5.4 | 5.1 | 4.2 | 9.7 | 4.2 | 2.5 | 2.7 | 2.8 | 5.9 | 4.2 | 3.8 | 2.3 | 5.5 | 4.2 | 3.6 | 3.6 | 5.7 | 8.2 | 2.3 | 4.2 | 4.2 | 4.2 | 3.2 | 4.2 | 5.3 | 6.5 | 5.1 | 3.8 | 4.9 | 5.7 | 7.3 | 5.5 | 2.5 | 4.4 | / | ||||||||
50 | M. (Pa.) sangzhiensis | 5.2 | 5.6 | 5.2 | 4.9 | 9.2 | 4.4 | 3.4 | 1.7 | 4.3 | 7.1 | 4.3 | 4.5 | 3.0 | 5.4 | 4.2 | 3.6 | 4.7 | 5.2 | 9.4 | 3.5 | 4.9 | 4.9 | 5.2 | 3.9 | 5.4 | 5.4 | 6.5 | 6.2 | 5.1 | 5.0 | 5.4 | 7.8 | 6.2 | 3.7 | 5.3 | 3.6 | / | |||||||
51 | M. (Pa.) shunhuangensis | 1.5 | 3.4 | 3.4 | 2.4 | 5.9 | 2.5 | 4.0 | 4.3 | 1.7 | 4.1 | 3.0 | 2.6 | 4.0 | 4.3 | 4.0 | 3.4 | 2.1 | 4.1 | 6.8 | 4.5 | 3.0 | 2.2 | 2.6 | 1.7 | 2.0 | 3.4 | 3.8 | 4.9 | 2.6 | 3.5 | 4.1 | 5.7 | 4.5 | 4.1 | 5.3 | 4.0 | 4.7 | / | ||||||
52 | M. (Pa.) spinata | 4.9 | 5.1 | 4.7 | 4.5 | 9.2 | 4.4 | 3.2 | 1.1 | 3.9 | 6.7 | 4.3 | 4.5 | 3.2 | 5.4 | 4.2 | 3.2 | 3.9 | 5.6 | 9.4 | 3.4 | 4.9 | 4.5 | 5.2 | 3.5 | 5.0 | 4.7 | 6.5 | 5.8 | 4.9 | 5.4 | 5.0 | 7.8 | 6.2 | 2.8 | 4.6 | 3.4 | 1.7 | 4.3 | / | |||||
53 | M. (Pa.) tuberogranulatus | 2.6 | 3.4 | 3.7 | 2.2 | 7.0 | 1.7 | 3.6 | 3.0 | 1.7 | 4.7 | 2.4 | 2.2 | 3.4 | 3.9 | 4.0 | 3.4 | 2.1 | 3.9 | 7.7 | 3.4 | 2.6 | 2.4 | 2.6 | 1.9 | 2.8 | 3.9 | 4.3 | 5.1 | 2.6 | 3.0 | 4.5 | 5.1 | 4.3 | 3.9 | 4.8 | 3.2 | 3.2 | 2.1 | 3.5 | / | ||||
54 | M. (Pa.) wugongensis | 3.9 | 4.2 | 3.9 | 3.9 | 8.9 | 4.2 | 4.6 | 4.5 | 3.6 | 6.0 | 4.7 | 2.8 | 4.0 | 4.3 | 4.2 | 4.0 | 3.7 | 4.5 | 8.1 | 4.3 | 3.6 | 3.4 | 4.3 | 3.7 | 4.7 | 4.5 | 6.0 | 5.3 | 4.9 | 4.3 | 4.5 | 5.7 | 4.9 | 4.5 | 6.5 | 4.2 | 4.5 | 3.7 | 4.1 | 3.4 | / | |||
55 | M. (Pa.) wuliangshanensis | 3.9 | 5.5 | 4.7 | 4.9 | 9.2 | 4.6 | 3.2 | 3.4 | 4.1 | 6.3 | 4.5 | 4.5 | 2.9 | 5.8 | 4.7 | 4.6 | 4.5 | 5.8 | 8.3 | 3.0 | 4.7 | 4.7 | 4.3 | 3.7 | 4.9 | 5.4 | 6.5 | 5.4 | 4.7 | 5.2 | 6.2 | 7.0 | 5.8 | 3.0 | 4.6 | 2.8 | 4.3 | 3.7 | 3.7 | 3.7 | 4.9 | / | ||
56 | M. (Pa.) wushanensis | 3.6 | 3.6 | 3.6 | 2.8 | 8.1 | 2.1 | 4.0 | 3.9 | 2.6 | 4.9 | 3.4 | 3.2 | 3.6 | 3.9 | 3.8 | 3.6 | 3.0 | 3.6 | 7.7 | 4.3 | 3.2 | 3.0 | 3.2 | 2.4 | 3.4 | 4.5 | 4.0 | 6.0 | 3.2 | 3.7 | 5.0 | 6.0 | 4.3 | 3.7 | 5.0 | 3.8 | 4.5 | 2.6 | 4.5 | 1.7 | 3.9 | 3.9 | / | |
57 | M. (Pa.) xianjuensis | 3.6 | 4.4 | 4.0 | 3.2 | 7.3 | 3.2 | 4.0 | 4.4 | 2.3 | 4.9 | 3.6 | 3.0 | 3.6 | 4.2 | 4.8 | 3.8 | 3.0 | 4.2 | 7.6 | 4.4 | 4.0 | 3.4 | 3.0 | 2.7 | 3.2 | 4.6 | 2.8 | 5.5 | 3.4 | 4.0 | 5.1 | 5.1 | 4.6 | 3.8 | 5.1 | 3.8 | 5.3 | 2.9 | 5.0 | 2.9 | 4.8 | 4.4 | 3.2 | / |
Diagnostic characters separating all 46 species of the Megophrys (Panophrys).
ID | Species | SVL in males (in mm) | SVL in females (in mm) | Horn-like tubercle at upper eyelid: slightly large (2), small (1) | Vomerine teeth: present (1), or absent (0) | Tongue: notched (1), or not notched (0) | Lateral fringes on toes: wide (2), narrow (1), or lacking (0) | Webs on toes: more than one-fourth (2), rudimentary (1), or lacking (0) | TD/ED | TIB/SVL |
---|---|---|---|---|---|---|---|---|---|---|
1 | M. (Pa.) mirabilis sp. nov. | 55.8–61.4 | 68.5–74.8 | 2 | 0 | 0 | 1 | 1 | 0.49–0.63 | 0.45–0.47 |
2 | M. (Pa.) shimentaina sp. nov. | 28.0–30.6 | / | 1 | 1 | 0 | 1 | 1 | 0.57–0.66 | 0.44–0.53 |
3 | M. (Pa.) xiangnanensis sp. nov. | 38.6–42.0 | 44.4 | 1 | 0 | 0 | 2 | 1 | 0.38–0.49 | 0.41–0.46 |
4 | M. (Pa.) yangmingensis sp. nov. | 33.2–37.1 | 45.2 | 1 | 0 | 0 | 1 | 1 | 0.42–0.50 | 0.44–0.51 |
5 | M. (Pa.) acuta | 27.1–33.0 | 28.1–33.6 | 2 | 0 | 0 | 1 | 1 | 0.57–0.71 | 0.38–0.45 |
6 | M. (Pa.) baolongensis | 42.0–45.0 | / | 1 | 0 | 1 | 0 | 0 | 0.41 | 0.46 |
7 | M. (Pa.) binchuanensis | 32.0–36.0 | 40.2–42.5 | 1 | 0 | 1 or 0 | 2 | 1 | 0.33–0.50 | 0.46–0.48 |
8 | M. (Pa.) binlingensis | 45.1–51.0 | / | 1 | 0 | 1 | / | 1 | 0.47–0.52 | 0.52–0.53 |
9 | M. (Pa.) boettgeri | 34.5–37.8 | 39.7–46.8 | 1 | 0 | 1 | 2 | 1 | 0.40–0.67 | 0.45–0.49 |
10 | M. (Pa.) brachykolos | 33.7–39.3 | 33.9–45.9 | 1 | 0 | 0 | 0 | 1 | > 0.50 | 0.37–0.42 |
11 | M. (Pa.) caudoprocta | 81.3 | / | 2 | 1 | 0 | / | 1 | 0.5 | 0.51 |
12 | M. (Pa.) cheni | 26.2–29.5 | 31.8–34.1 | 1 | 0 | 1 | 2 | 1 | 0.41–0.54 | 0.50–0.54 |
13 | M. (Pa.) daweimontis | 34.0–37.0 | 40.0–46.0 | 1 | 1 | / | 0 | 0 | / | 0.54 |
14 | M. (Pa.) dongguanensis | 30.2–39.3 | / | 1 | 1 | 0 | 0 | 1 | 0.42–0.60 | 0.41–0.46 |
15 | M. (Pa.) fansipanensis | 30.9–44.3 | 41.7–42.5 | 1 | 1 | 1 | 0 | 0 | 0.53–0.80 | 0.49–0.59 |
16 | M. (Pa.) hoanglienensis | 37.4–47.6 | 59.6 | 1 | 1 | 1 | 0 | 0 | 0.54–0.75 | 0.44–0.63 |
17 | M. (Pa.) huangshanensis | 36.0–41.6 | 44.2 | 1 | 0 | 1 | 0 | 0 | <0.50 | 0.42–0.45 |
18 | M. (Pa.) insularis | 36.8–41.2 | 47.1 | 1 | 1 | 1 | 0 | 1 | 0.46–0.57 | 0.40–0.43 |
19 | M. (Pa.) jiangi | 34.4–39.2 | 39.5–40.4 | 1 | 0 | 0 | 0 | 1 | / | / |
20 | M. (Pa.) jingdongensis | 53.0–56.5 | 63.5 | 1 | 1 | 1 | 2 | 2 | / | 0.58–0.59 |
21 | M. (Pa.) jinggangensis | 35.1–36.7 | 38.4–41.6 | 2 | 1 | 0 | 1 | 1 | 0.73–0.88 | 0.47–0.50 |
22 | M. (Pa.) jiulianensis | 30.4–33.9 | 34.1–37.5 | 1 | 1 | 1 | 0 | 1 | 0.50–0.59 | 0.44–0.48 |
23 | M. (Pa.) kuatunensis | 26.2–29.6 | 37.4 | 1 | 0 | 1 | 1 | 0 | 0.44 | 0.38–0.48 |
24 | M. (Pa.) latidactyla | 38.9 | / | 2 | 1 | 0 | 2 | 1 | 0.85 | 0.52 |
25 | M. (Pa.) leishanensis | 30.4–38.7 | 42.3 | 1 | 0 | 0 | 0 | 1 | / | / |
26 | M. (Pa.) liboensis | 60.5–67.7 | 60.8–70.6 | 2 | 1 | 1 | 2 | 1 | 0.48–0.78 | 0.44–0.61 |
27 | M. (Pa.) lini | 34.1–39.7 | 37.0–39.9 | 1 | 0 | 0 | 2 | 1 | 0.40–0.60 | 0.46–0.53 |
28 | M. (Pa.) lishuiensis | 30.7–34.7 | 36.9–40.4 | 1 | 0 | 0 | 0 | 0 | / | / |
29 | M. (Pa.) minor | 34.5–41.2 | / | 1 | 0 | 1 | 0 | 1 | 0.8–0.83 | 0.46–0.48 |
30 | M. (Pa.) mufumontana | 30.1–30.8 | 36.3 | 1 | 0 | 0 | 1 | 1 | 0.51–0.58 | 0.47–0.53 |
31 | M. (Pa.) nankunensis | 29.9–34.9 | 39.4–41.9 | 1 | 1 | 0 | 0 | 1 | 0.43–0.61 | 0.35–0.42 |
32 | M. (Pa.) nanlingensis | 30.5–37.3 | / | 1 | 1 | 1 | 1 | 1 | 0.43–0.57 | 0.45–0.51 |
33 | M. (Pa.) obesa | 35.6 | 37.5–41.2 | 1 | 0 | 0 | 0 | 1 | 0.51–0.66 | 0.41–0.47 |
34 | M. (Pa.) ombrophila | 27.4–34.5 | 32.8–35.0 | 1 | 0 | 0 | 0 | 0 | 0.53–0.69 | 0.33–0.41 |
35 | M. (Pa.) omeimontis | 56.0–59.5 | 68.0–72.5 | 1 | 1 | 1 | 1 | 1 | / | 0.52–0.56 |
36 | M. (Pa.) palpebralespinosa | 36.2–38.0 | / | 2 | 1 | 0 | 2 | 2 | / | 0.55 |
37 | M. (Pa.) rubrimera | 26.7–30.5 | / | 1 | 1 | 1 | 1 | 0 | 0.58–0.76 | 0.48–0.56 |
38 | M. (Pa.) sangzhiensis | 54.7 | / | 1 | 1 | 1 | 1 | 1 | 0.62 | 0.59 |
39 | M. (Pa.) shuichengensis | 102.0–118.3 | 99.8–115.6 | 2 | 0 | 1 | 2 | 2 | 0.67 | 0.43–0.47 |
40 | M. (Pa.) shunhuangensis | 30.3–33.7 | 37.6 | 1 | 0 | 0 | 0 | 1 | 0.40–0.63 | 0.50–0.55 |
41 | M. (Pa.) spinata | 47.2–54.4 | 54.0–55.0 | 1 | 0 | 1 | 2 | 2 | 0.43 | 0.56–0.58 |
42 | M. (Pa.) tuberogranulatus | 33.2–39.0 | 50.5 | 1 | 0 | 0 | 0 | 1 | 0.5 | 0.45–0.51 |
43 | M. (Pa.) wugongensis | 31.0–34.1 | 38.5–42.8 | 1 | 0 | 0 | 0 | 1 | 0.45–0.53 | 0.37–0.44 |
44 | M. (Pa.) wuliangshanensis | 27.3–31.6 | 41.3 | 1 | 0 | 1 or 0 | 0 | 0 | 0.5 | 0.50–0.51 |
45 | M. (Pa.) wushanensis | 30.4–35.5 | 38.4 | 1 | 0 | 0 | 0 (in female), 2 (in male) | 1 | 0.5 | 0.47–0.48 |
46 | M. (Pa.) xianjuensis | 31.0–36.3 | 41.6 | 1 | 0 | 0 | 1 | 1 | 0.48–0.60 | 0.40–0.50 |
Megophrys sp25 (
Holotype.
Paratypes. Three adult specimens from the same locality as the holotype: male
The specific epithet mirabilis means marvelous, referring to its distinctive habitus and color pattern of this species within the subgenus Panophrys.
(1) Body size relatively large, SVL 55.8–61.4 mm (N = 2) in adult males and SVL 68.5–74.8 (N = 2) mm in adult females; (2) snout rounded in dorsal view; (3) internasal distance smaller than interorbital distance; (4) tympanum clear, moderate size, TD/ED 0.49–0.63; (5) absence of vomerine ridge and vomerine teeth; (6) tongue small, majorly attached to the mandible, free margin small and rounded, not notched behind; (7) hindlimbs slender, heels overlapping and tibio-tarsal articulation reaching forward at the central eye; (8) fingers with distinct lateral fringes, presence of indistinct subarticular tubercles at the bases; (9) toes with distinct lateral fringes and rudiment of webs, presence of indistinct subarticular tubercles at the bases; (10) presence of slightly large horn-like tubercle at the edge of upper eyelid; (11) dorsal skin smooth with granules, (12) skin on flanks flabby, with spiny tubercles; (13) supratympanic fold distinct, with dense tubercles, forming an extremely swollen large shoulder gland above insertion of arm; (14) grayish brown above, tinged with blue in males, but dorsum of head and body reddish brown in females; (15) ventral surface of throat and chest with grayish blue latticed patches and black spots in males, but with orange latticed patches and black spots in females; (16) presence of underdeveloped nuptial pads on the dorsal surface of the first finger in adult males.
Megophrys (Panophrys) mirabilis sp. nov. can be easily distinguished from all recognized congeners, by having a small tongue, majorly attached to the mandible, flank skin flabby with spiny tubercles, and supratympanic fold with dense tubercles forming an extremely swollen large shoulder gland above insertion of arm.
Further, detailed comparative data of Megophrys (Panophrys) mirabilis sp. nov. with 42 recognized congeners of Panophrys are given in Table
Five Panophrys species were previously recorded from the hilly areas among Guangdong, Guangxi, and Hunan, namely Megophrys (Panophrys) acuta, M. (Pa.) brachykolos, M. (Pa.) nanlingensis, M. (Pa.) obesa, and M. (Pa.) shunhuangensis. M. (Pa.) mirabilis sp. nov. differs from M. (Pa.) acuta by the larger body size, SVL 55.8–61.4 mm in males and 68.5–74.8 mm in females (vs. 27.1–33.0 mm in males and 28.1–33.6 in females), snout rounded in dorsal view (vs. strongly remarkably pointed), fingers with distinct lateral fringes (vs. absent), and overlapping heels (vs. not meeting). M. (Pa.) mirabilis sp. nov. differs from M. (Pa.) brachykolos by the larger body size, SVL 55.8–61.4 mm in males and 68.5–74.8 mm in females (vs. 33.7–39.3 mm in males and 33.9–45.9 in females), slightly large horn-like tubercle at upper eyelid (vs. small), fingers and toes with distinct lateral fringes (vs. all absent), overlapping heels (vs. not meeting). M. (Pa.) mirabilis sp. nov. differs from M. (Pa.) nanlingensis by the larger body size, SVL 55.8–61.4 mm in males (vs. 30.5–37.3 mm), slightly large horn-like tubercle at upper eyelid (vs. small), absence of vomerine ridge and vomerine teeth (vs. both present), tongue not notched behind (vs. notched), and fingers with distinct lateral fringes (vs. absent). M. (Pa.) mirabilis sp. nov. differs from M. (Pa.) obesa by larger body size, SVL 55.8–61.4 mm in males and 68.5–74.8 mm in females (vs. 35.6 mm in male and 37.5–41.2 in females), slightly large horn-like tubercle at upper eyelid (vs. small), absence of vomerine ridge (vs. present), fingers and toes with distinct lateral fringes (vs. all absent), and overlapping heels (vs. not meeting). M. (Pa.) mirabilis sp. nov. differs from M. (Pa.) shunhuangensis by larger body size, SVL 55.8–61.4 mm in males and 68.5–74.8 mm in females (vs. 30.3–33.7 mm in males and 37.6 in female), slightly large horn-like tubercle at upper eyelid (vs. small), and fingers and toes with lateral fringes (vs. all absent).
With a large body size, SVL 55.8–61.4 mm in adult males and 68.5–74.8 mm in adult females, Megophrys (Panophrys) mirabilis sp. nov. is significantly different from 30 congeners whose SVL < 50 mm in males or < 60 mm in females, namely M. (Pa.) baolongensis, M. (Pa.) binchuanensis, M. (Pa.) boettgeri, M. (Pa.) cheni, M. (Pa.) daweimontis, M. (Pa.) dongguanensis, M. (Pa.) fansipanensis, M. (Pa.) hoanglienensis, M. (Pa.) huangshanensis, M. (Pa.) insularis, M. (Pa.) jiangi, M. (Pa.) jinggangensis, M. (Pa.) jiulianensis, M. (Pa.) kuatunensis, M. (Pa.) latidactyla, M. (Pa.) leishanensis, M. (Pa.) lini, M. (Pa.) lishuiensis, M. (Pa.) minor, M. (Pa.) mufumontana, M. (Pa.) nankunensis, M. (Pa.) ombrophila, M. (Pa.) palpebralespinosa, M. (Pa.) rubrimera, M. (Pa.) spinata, M. (Pa.) tuberogranulatus, M. (Pa.) wugongensis, M. (Pa.) wuliangshanensis, M. (Pa.) wushanensis, and M. (Pa.) xianjuensis.
Megophrys (Panophrys) mirabilis sp. nov. can be further distinguished from the remaining seven congeners by the following characteristics: SVL 55.8–61.4 mm in adult males and 68.5–74.8 mm in adult females [vs. SVL 45.1–51.0 mm in adult males in M. (Pa.) binlingensis; vs. SVL 81.3 mm in adult male in M. (Pa.) caudoprocta; vs. SVL 63.5 mm in adult female in M. (Pa.) jingdongensis; vs. SVL 102.0–118.3 mm in adult males and 99.8–115.6 mm in adult females in M. (Pa.) shuichengensis]; slightly large horn-like tubercle at upper eyelid [vs. small in M. (Pa.) binlingensis, M. (Pa.) jingdongensis, M. (Pa.) omeimontis, and M. (Pa.) sangzhiensis]; vomerine teeth absent [vs. present in M. (Pa.) caudoprocta, M. (Pa.) jingdongensis, M. (Pa.) liboensis, M. (Pa.) omeimontis, and M. (Pa.) sangzhiensis]; tongue not notched behind [vs. notched in M. (Pa.) binlingensis, M. (Pa.) jingdongensis, M. (Pa.) liboensis, M. (Pa.) omeimontis, M. (Pa.) sangzhiensis, and M. (Pa.) shuichengensis]; lateral fringes on toes narrow [vs. wide in M. (Pa.) jingdongensis, M. (Pa.) liboensis, and M. (Pa.) shuichengensis]; rudimentary webs on toes [vs. more than one-fourth webs in M. (Pa.) jingdongensis and M. (Pa.) shuichengensis].
Adult male. Body size large, SVL 61.4 mm; head width slightly larger than head length, HDW/HDL 1.02; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes large, ED 0.31 of HDL, pupil vertical; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance smaller than interorbital distance; tympanum clear, TD/ED 0.49; large ovoid choanae at the base of the maxilla; absence of vomerine ridge and vomerine teeth; tongue small, majority attached at the mouth, margin rounded, not notched behind; absence of vocal sac.
Radio-ulna length 0.26 of SVL and hand 0.28 of SVL; hand without webs, fingers with distinct lateral fringes, relative finger length II < I < IV < III; tips of fingers slightly dilated, round; one indistinct subarticular tubercle at the bases of each finger; metacarpal tubercles indistinct, the inner one observably enlarged and the outer one smaller; presence of underdeveloped nuptial pad on the dorsal surface of the first finger, without nuptial spines. Hindlimbs slender, tibio-tarsal articulation reaching forward at the central eye when hindlimb is stretched along the side of the body; heels overlapping when the flexed hindlimbs are held at right angles to the body axis; tibia length 0.47 of SVL and foot length 0.71 of SVL; relative toe length I < II < V < III < IV; tips of toes round and slightly dilated; toes with narrow lateral fringes and rudiment of webs; one indistinct subarticular tubercle at the bases of each toe; inner metatarsal tubercle long ovoid and the outer one absent.
Dorsal skin smooth with sparse granules; flanks flabby with spiny tubercles; distinct supratympanic fold curving postero-ventrally from posterior corner of eye to a level above insertion of arm; small tubercles arranged from above the nostril, along the canthus rostralis, edge of upper eyelid and supratympanic fold, to the posterior margin of temporal region; a distinct horn-like prominent tubercle on the edge of upper eyelid; a discontinuous X-shaped ridge with several short ridges on two sides on the back; transverse skin ridges on the dorsal shank and thigh; ventral surface smooth; several tubercles on posterior hindlimbs; small pectoral gland closer to axilla; a single large femoral gland on rear of thigh.
Grayish brown above in life; an dark interorbital triangle with light colored center and edge; a dark X-shaped making with light edge on the central of dorsum; dark brown transverse bands on forearms and hindlimbs; supratympanic fold light gray; dark vertical band below the eye; iris grayish brown; ventral surface grayish white; throat and chest with grayish blue latticed patches and black spots; ventral hands and feet grayish white, tips of digits creamy white, metacarpal tubercle and metatarsal tubercle grayish white; pectoral gland and femoral gland white.
Measurement data of type series are listed in Table
Currently, Megophrys (Panophrys) mirabilis sp. nov. is only known from Huaping Nature Reserve, northeastern Guangxi. The individuals were found on shrubbery branches near trail paths between elevations of 1300–1330 m a.s.l. from June to September. Males were not calling when found, but the collected female specimens bear mature yellowish oocytes. Tadpoles have not been found and ecological information remains unknown.
Measurements (in mm) of the type series of Megophrys (Panophrys) mirabilis sp. nov., * for the holotype.
|
|
|
|
|
---|---|---|---|---|
Sex | Male | Male | Female | Female |
SVL | 61.4 | 55.8 | 74.8 | 68.5 |
HDL | 21.4 | 18.8 | 23.7 | 22.6 |
HDW | 21.8 | 18.8 | 23.9 | 22.4 |
SNT | 7.8 | 7.1 | 9.0 | 8.8 |
IND | 6.7 | 5.9 | 7.5 | 6.8 |
IOD | 7.2 | 6.5 | 8.1 | 7.6 |
ED | 6.7 | 5.9 | 8.1 | 6.8 |
TD | 3.3 | 3.2 | 4.3 | 4.3 |
TED | 3.3 | 3.2 | 4.2 | 3.7 |
HND | 17.3 | 15.3 | 20.2 | 19.5 |
RAD | 15.9 | 13.9 | 18.3 | 17.6 |
FTL | 43.7 | 37.8 | 48.8 | 43.2 |
TIB | 28.9 | 26.3 | 33.8 | 30.5 |
Megophrys sp29 (
Holotype.
Paratypes. Eleven adult males from the same locality as the holotype:
The specific epithet shimentaina refers to its type locality, Shimentai Nature Reserve.
Shimentai Horned Toad (in English) / Shí Mén Taí Jiăo Chán (石门台角蟾in Chinese)
(1) Body size small, SVL 28.0–30.6 (28.9 ± 0.9, N = 12) mm in adult males; (2) snout rounded in dorsal view; (3) tympanum clear, TD/ED 0.57–0.66; (4) presence of weak vomerine ridge and vomerine teeth; (5) margin of tongue rounded, not notched behind; (6) hindlimbs slender, heels overlapping and tibio-tarsal articulation reaching forward between tympanum to anterior corner of eye; (7) tibia 0.44–0.53 of SVL and foot 0.62–0.76 of SVL; (8) fingers with narrow lateral fringes, presence of indistinct subarticular tubercles at the bases; (9) toes with narrow lateral fringes and rudiment of webs, absence of subarticular tubercle; (10) presence of a small horn-like tubercle at the edge of upper eyelid; (11) presence of tiny, barely visible, black to dark brown spines on the whole dorsal skin, flanks, dorsal limbs, the region around cloaca, and rear of hindlimbs; (12) dorsal skin rough, a discontinuous “/ \”-shaped ridge with two discontinuous dorsolateral ridges on two sides on the back; (13) several large warts on the flanks; (14) supratympanic fold distinct and white, with tiny spines; (15) light brown above, a dark brown stripe on each upper eyelid; (16) single subgular vocal sac in males; (17) weak nuptial pads with serried olive nuptial spines, on the dorsal surface of the first and second fingers in adult males.
Comparative data of Megophrys (Panophrys) shimentaina sp. nov. with M. (Pa.) mirabilis sp. nov. and 42 recognized congeners of Panophrys are given in Table
Megophrys (Panophrys) shimentaina sp. nov. differs from M. (Pa.) mirabilis sp. nov. by the smaller body size, SVL 28.0–30.6 mm in males (vs. 55.8–61.4 mm in males), small horn-like tubercle at upper eyelid (vs. slightly large), presence of vomerine teeth (vs. absent), the presence of tiny spines on the whole dorsal skin, flanks, dorsal limbs, the region around cloaca, and rear of hindlimbs (vs. such spines absent), presence of vocal sac in males (vs. absent), and presence of nuptial spines in males (vs. absent).
Compared with the five Panophrys species previously recorded from the hilly areas among Guangdong, Guangxi, and Hunan, Megophrys (Panophrys) shimentaina sp. nov. differs from M. (Pa.) acuta by the small horn-like tubercle at upper eyelid (vs. slightly large), snout rounded in dorsal view (vs. strongly remarkably pointed), presence of vomerine teeth (vs. absent), presence of tiny spines on the whole dorsal skin, flanks, dorsal limbs, the region around cloaca, and rear of hindlimbs (vs. such spines absent), and overlapping heels (vs. not meeting). M. (Pa.) shimentaina sp. nov. differs from M. (Pa.) brachykolos by the smaller body size SVL 28.0–30.6 mm in males (vs. 33.7–39.3 mm in males), presence of vomerine teeth (vs. absent), presence of tiny spines on the whole dorsal skin, flanks, dorsal limbs, the region around cloaca, and rear of hindlimbs (vs. such spines absent), narrow lateral fringes on toes (vs. absent), and overlapping heels (vs. not meeting). M. (Pa.) shimentaina sp. nov. differs from M. (Pa.) nanlingensis by the presence of tiny spines on the whole dorsal skin, flanks, dorsal limbs, the region around cloaca, and rear of hindlimbs (vs. such spines absent), and tongue not notched behind (vs. notched). M. (Pa.) shimentaina sp. nov. differs from M. (Pa.) obesa by the smaller body size SVL 28.0–30.6 mm in males (vs. 35.6 mm in single male), presence of vomerine teeth (vs. absent), presence of tiny spines on the whole dorsal skin, flanks, dorsal limbs, the region around cloaca, and rear of hindlimbs (vs. such spines absent), narrow lateral fringes on toes (vs. absent), and overlapping heels (vs. not meeting). M. (Pa.) shimentaina sp. nov. differs from M. (Pa.) shunhuangensis by the presence of vomerine teeth (vs. absent), tibio-tarsal articulation reaching forward between tympanum to anterior corner of eye (vs. at the eye), and the presence of tiny spines on the whole dorsal skin, flanks, dorsal limbs, the region around cloaca, and rear of hindlimbs (vs. such spines absent).
With a small body size, SVL 28.0–30.6 mm in adult males, Megophrys (Panophrys) shimentaina sp. nov. is significantly different from 15 congeners whose SVL > 35 mm in males, namely M. (Pa.) baolongensis, M. (Pa.) binlingensis, M. (Pa.) caudoprocta, M. (Pa.) hoanglienensis, M. (Pa.) huangshanensis, M. (Pa.) insularis, M. (Pa.) jingdongensis, M. (Pa.) jinggangensis, M. (Pa.) latidactyla, M. (Pa.) liboensis, M. (Pa.) omeimontis, M. (Pa.) palpebralespinosa, M. (Pa.) sangzhiensis, M. (Pa.) shuichengensis, and M. (Pa.) spinata.
Megophrys (Panophrys) shimentaina sp. nov. can be further distinguished from the remaining 22 congeners by the following characteristics: vomerine teeth present [vs. absent in M. (Pa.) binchuanensis, M. (Pa.) boettgeri, M. (Pa.) cheni, M. (Pa.) jiangi, M. (Pa.) kuatunensis, M. (Pa.) leishanensis, M. (Pa.) lini, M. (Pa.) lishuiensis, M. (Pa.) minor, M. (Pa.) mufumontana, M. (Pa.) ombrophila, M. (Pa.) tuberogranulatus, M. (Pa.) wugongensis, M. (Pa.) wuliangshanensis, M. (Pa.) wushanensis, and M. (Pa.) xianjuensis]; tongue not notched behind [vs. notched in M. (Pa.) cheni, M. (Pa.) boettgeri, M. (Pa.) fansipanensis, M. (Pa.) jiulianensis, M. (Pa.) kuatunensis, M. (Pa.) minor, and M. (Pa.) rubrimera]; lateral fringes on toes narrow [vs. absent in M. (Pa.) daweimontis, M. (Pa.) dongguanensis, M. (Pa.) fansipanensis, M. (Pa.) jiangi, M. (Pa.) jiulianensis, M. (Pa.) leishanensis, M. (Pa.) lishuiensis, M. (Pa.) minor, M. (Pa.) nankunensis, M. (Pa.) ombrophila, M. (Pa.) tuberogranulatus, M. (Pa.) wugongensis, and M. (Pa.) wuliangshanensis; wide in M. (Pa.) binchuanensis, M. (Pa.) boettgeri, M. (Pa.) cheni, and M. (Pa.) lini; vs. absent in females while wide in males in M. (Pa.) wushanensis]; rudimentary webs on toes [vs. lacking webs in M. (Pa.) daweimontis, M. (Pa.) fansipanensis, M. (Pa.) kuatunensis, M. (Pa.) lishuiensis, M. (Pa.) ombrophila, M. (Pa.) rubrimera, and M. (Pa.) wuliangshanensis].
Adult male. Body size small, SVL 28.4 mm; head width slightly smaller than head length, HDW/HDL 0.95; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes large, ED 0.33 of HDL, pupil vertical; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum clear, in medium size, TD/ED 0.61; large ovoid choanae at the base of the maxilla; presence of weak vomerine ridge and vomerine teeth; margin of tongue rounded, not notched behind; presence of a single subgular vocal sac, a pair of slit-like openings at posterior of jaw.
Radio-ulna length 0.22 of SVL and hand 0.26 of SVL; hand without webs, fingers with narrow lateral fringes, relative finger length I ≈ II < IV < III; tips of fingers slightly dilated, round; one indistinct subarticular tubercle at the bases of each finger; inner metacarpal tubercle observably enlarged and the outer one smaller; nuptial pads with serried olive nuptial spines on the dorsal surface of the first and second fingers. Hindlimbs slender, tibio-tarsal articulation reaching forward to the posterior corner of eye when hindlimb is stretched along the side of the body; heels overlapping when the flexed hindlimbs are held at right angles to the body axis; tibia length 0.47 of SVL and foot length 0.67 of SVL; relative toe length I < II < V < III < IV; tips of toes round and slightly dilated; toes with distinct lateral fringes and rudiment of webs, without subarticular tubercle; inner metatarsal tubercle long ovoid and the outer one absent.
Dorsal skin rough; numerous granules densely arranged on the top of head, loreal region, lips, temporal region, dorsal body, flanks and dorsal limbs; several tubercles on upper eyelid, including a horn-like prominent tubercle on the edge; all granules and tubercles bearing tiny, barely visible spines; clear supratympanic fold with tiny spines, curving postero-ventrally from posterior corner of eye to a level above insertion of arm; tubercles and granules forming discontinuous “/ \”-shaped ridge and two discontinuous dorsolateral ridges on two sides at the central back; large tubercles and warts on the flanks; ventral surface smooth; several granules bearing black spines on the region around cloaca and rear of hindlimbs; small pectoral gland closer to axilla; a single large femoral gland on rear of thigh.
Light brown above in life; a dark brown stripe on dorsal surface of each eye; narrow dark brown transverse bands on forearms and hindlimbs; supratympanic fold white; dark vertical band below the eye; iris reddish brown; all spines black or dark brown; ventral surface pale; throat flesh color; scarlet spots on the chest; a large white blotch on the belly; a pair of lateroventral longitudinal broad black stripes with several white tubercles on two sides; ventral limbs flesh color with white spots; ventral hands and ventral feet brown, tips of digits pale brown; metacarpal tubercle and metatarsal tubercle reddish; pectoral gland and femoral gland white.
Measurement data of type series are listed in Table
Currently, Megophrys (Panophrys) shimentaina sp. nov. is known only from Shimentai Nature Reserve, northern Guangdong. This toad is uncommon in its distribution areas. All individuals were found from two slowly flowing mountain streams between elevations of 210–500 m a.s.l. Males call on plant leaves from April to August, suggesting their breeding season corresponds to this period. Females and tadpoles have not been found.
The advertisement calls of Megophrys (Panophrys) shimentaina sp. nov. were recorded from four males at 18–20 °C air temperature on 27 April 2016. Thirty calls with 96 notes are measured and the spectrograms are shown in Fig.
Measurements (in mm) of the type series of Megophrys (Panophrys) shimentaina sp. nov., * for the holotype.
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Sex | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male |
SVL | 28.4 | 28.5 | 28.1 | 30.6 | 29.0 | 29.2 | 28.8 | 28.0 | 30.4 | 28.0 | 29.3 | 28.7 |
HDL | 10.0 | 10.1 | 9.9 | 10.4 | 10.1 | 10.1 | 10.0 | 9.9 | 10.5 | 10.7 | 10.1 | 10.1 |
HDW | 9.6 | 9.5 | 9.5 | 10.0 | 9.6 | 9.7 | 9.5 | 9.8 | 10.0 | 10.3 | 9.8 | 9.9 |
SNT | 3.3 | 3.4 | 3.3 | 3.5 | 3.3 | 3.4 | 3.4 | 3.3 | 3.4 | 3.2 | 3.4 | 3.4 |
IND | 3.0 | 3.0 | 3.0 | 3.1 | 3.1 | 3.1 | 3.0 | 3.1 | 3.2 | 2.8 | 3.1 | 3.0 |
IOD | 2.6 | 2.8 | 2.8 | 2.9 | 2.8 | 2.8 | 2.7 | 2.8 | 2.9 | 3.1 | 2.8 | 2.6 |
ED | 3.3 | 3.2 | 3.3 | 3.4 | 3.3 | 3.3 | 3.3 | 3.4 | 3.4 | 3.4 | 3.4 | 3.4 |
TD | 2.0 | 1.8 | 1.9 | 2.2 | 2.0 | 1.9 | 2.0 | 1.9 | 2.1 | 2.1 | 2.2 | 2.1 |
TED | 1.6 | 1.6 | 1.5 | 1.6 | 1.7 | 1.5 | 1.5 | 1.4 | 1.6 | 1.5 | 1.5 | 1.5 |
HND | 7.4 | 7.2 | 7.3 | 7.5 | 7.5 | 7.4 | 7.3 | 6.8 | 7.2 | 7.8 | 7.1 | 7.5 |
RAD | 6.2 | 6.1 | 6.2 | 6.4 | 6.2 | 6.2 | 6.0 | 5.5 | 6.1 | 6.3 | 6.0 | 6.4 |
FTL | 19.1 | 19.9 | 19.0 | 20.5 | 19.4 | 19.3 | 19.3 | 17.9 | 18.7 | 21.4 | 19.4 | 20.3 |
TIB | 13.5 | 14.2 | 13.2 | 14.9 | 13.4 | 14.3 | 13.6 | 12.8 | 13.3 | 14.9 | 13.4 | 14.6 |
Megophrys sp2 (
Eleven adult specimens, female
The specific epithet xiangnanensis is an adjective derived from Chinese Pinyin Xiāng Nán, which means southern Hunan, for the distribution area of this species.
Southern Hunan Horned Toad (in English) / Xiāng Nán Jiăo Chán (湘南角蟾 in Chinese)
(1) Moderate body size, SVL 38.6–42.0 mm (40.3 ± 1.3, N = 11) in adult males and SVL 44.4 mm in adult female; (2) snout rounded in dorsal view; (3) tympanum clear, TD/ED 0.38–0.49; (4) presence of weak vomerine ridge, absence of vomerine teeth; (5) margin of tongue rounded, not notched behind; (6) hindlimbs slender, heels just meeting and tibio-tarsal articulation reaching forward between eye and tympanum; (7) tibia 0.41–0.46 of SVL and foot 0.57–0.62 of SVL; (8) fingers without lateral fringes, presence of distinct subarticular tubercles at the bases; (9) toes with relatively wide lateral fringes and rudiment of webs, presence of distinct subarticular tubercles at the bases; (10) presence of small horn-like tubercle at the edge of upper eyelid; (11) dorsal skin smooth with sparse granules, a discontinuous X-shaped ridge with two discontinuous dorsolateral ridges on two side on the back; (12) sparse tubercles on the flanks; (13) supratympanic fold light colored; (14) single subgular vocal sac in males; (15) presence of nuptial pads on the dorsal surface of the first and second fingers in adult males.
Comparative data of Megophrys (Panophrys) xiangnanensis sp. nov. with M. (Pa.) mirabilis sp. nov., M. (Pa.) shimentaina sp. nov., and 42 recognized congeners of Panophrys are given in Table
Megophrys (Panophrys) xiangnanensis sp. nov. differs from M. (Pa.) mirabilis sp. nov. by the smaller body size, SVL 38.6–42.0 mm in males and 44.4 mm in single female (vs. 55.8–61.4 mm in males and 68.5–74.8 in females), small horn-like tubercle at upper eyelid (vs. slightly large), wide lateral fringes on toes (vs. narrow), heels just meeting (vs. overlapping), presence of vocal sac in males (vs. absent), and presence of nuptial spines in males (vs. absent).
Megophrys (Panophrys) xiangnanensis sp. nov. differs from M. (Pa.) shimentaina sp. nov. by the larger body size, SVL 38.6–42.0 mm in males (vs. 28.0–30.6 mm in males), absence of vomerine teeth (vs. present), wide lateral fringes on toes (vs. narrow), and heels just meeting (vs. overlapping).
Compared with the five Panophrys species previously recorded from the hilly areas among Guangdong, Guangxi and Hunan, Megophrys (Panophrys) xiangnanensis sp. nov. differs from M. (Pa.) acuta by the larger body size, SVL 38.6–42.0 mm in males and 44.4 mm in single female (vs. 27.1–33.0 mm in males and 28.1–33.6 mm in females), small horn-like tubercle at upper eyelid (vs. slightly large), snout rounded in dorsal view (vs. strongly remarkably pointed), wide lateral fringes on toes (vs. narrow), and heels just meeting (vs. not meeting). M. (Pa.) xiangnanensis sp. nov. differs from M. (Pa.) brachykolos by the wide lateral fringes on toes (vs. absent), and heels just meeting (vs. not meeting). M. (Pa.) xiangnanensis sp. nov. differs from M. (Pa.) nanlingensis by the larger body size, SVL 38.6–42.0 mm in males (vs. 30.5–37.3 mm in males), absence of vomerine teeth (vs. present), tongue not notched behind (vs. notched), wide lateral fringes on toes (vs. narrow) , and heels just meeting (vs. overlapping). M. (Pa.) xiangnanensis sp. nov. differs from M. (Pa.) obesa by the larger body size, SVL 38.6–42.0 mm in males and 44.4 mm in single female (vs. 35.6 mm in single male and 37.5–41.2 mm in females), wide lateral fringes on toes (vs. absent), and heels just meeting (vs. not meeting). M. (Pa.) xiangnanensis sp. nov. differs from M. (Pa.) shunhuangensis by the larger body size, SVL 38.6–42.0 mm in males and 44.4 mm in single female (vs. 30.3–33.7 mm in males and 37.6 in female), wide lateral fringes on toes (vs. absent), and heels just meeting (vs. overlapping).
With a moderate body size, SVL 38.6–42.0 mm in adult males, Megophrys (Panophrys) xiangnanensis sp. nov. is significantly different from 18 congeners whose SVL< 35 mm or > 45 mm in males, namely M. (Pa.) binlingensis, M. (Pa.) caudoprocta, M. (Pa.) cheni, M. (Pa.) jingdongensis, M. (Pa.) jiulianensis, M. (Pa.) kuatunensis, M. (Pa.) liboensis, M. (Pa.) lishuiensis, M. (Pa.) mufumontana, M. (Pa.) nankunensis, M. (Pa.) ombrophila, M. (Pa.) omeimontis, M. (Pa.) rubrimera, M. (Pa.) sangzhiensis, M. (Pa.) shuichengensis, M. (Pa.) spinata, M. (Pa.) wugongensis, and M. (Pa.) wuliangshanensis.
Megophrys (Panophrys) xiangnanensis sp. nov. can be further distinguished from the remaining 19 congeners by the following characteristics: small horn-like tubercle at upper eyelid [vs. slightly large in M. (Pa.) jinggangensis, M. (Pa.) latidactyla, and M. (Pa.) palpebralespinosa]; vomerine teeth absent [vs. present in M. (Pa.) daweimontis, M. (Pa.) dongguanensis, M. (Pa.) fansipanensis, M. (Pa.) hoanglienensis, M. (Pa.) insularis, M. (Pa.) jinggangensis, M. (Pa.) latidactyla, and M. (Pa.) palpebralespinosa]; tongue not notched behind [vs. notched in M. (Pa.) baolongensis, M. (Pa.) boettgeri, M. (Pa.) fansipanensis, M. (Pa.) hoanglienensis, M. (Pa.) huangshanensis, M. (Pa.) insularis, and M. (Pa.) minor]; lateral fringes on toes wide [vs. absent in M. (Pa.) baolongensis, M. (Pa.) daweimontis, M. (Pa.) dongguanensis, M. (Pa.) fansipanensis, M. (Pa.) hoanglienensis, M. (Pa.) huangshanensis, M. (Pa.) insularis, M. (Pa.) jiangi, M. (Pa.) leishanensis, M. (Pa.) minor, and M. (Pa.) tuberogranulatu; vs. narrow in M. (Pa.) jinggangensis and M. (Pa.) xianjuensis; vs. absent in females while wide in males in M. (Pa.) wushanensis]; rudimentary webs on toes [vs. more than one-fourth webs in M. (Pa.) palpebralespinosa; vs. lacking webs in M. (Pa.) baolongensis, M. (Pa.) daweimontis, M. (Pa.) fansipanensis, M. (Pa.) hoanglienensis, and M. (Pa.) huangshanensis].
Adult male. Moderate body size, SVL 40.9 mm; head width slightly larger than head length, HDW/HDL 1.02; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes large, ED 0.41 of HDL, pupil vertical; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum clear, TD/ED 0.44; large ovoid choanae at the base of the maxilla; presence of weak vomerine ridge, absence of vomerine teeth; margin of tongue rounded, not notched behind; presence of a single subgular vocal sac, a pair of slit-like openings at posterior of jaw.
Radio-ulna length 0.22 of SVL and hand 0.23 of SVL; hand without webs, fingers without lateral fringes, relative finger length I < II < IV < III; tips of fingers slightly dilated, round; one distinct subarticular tubercle at the bases of each finger; inner metacarpal tubercle observably enlarged and the outer one smaller; a single nuptial pad on the dorsal surface of the first and second fingers. Hindlimbs slender, tibio-tarsal articulation reaching forward between eye and tympanum when hindlimb is stretched along the side of the body; heels just meeting when the flexed hindlimbs are held at right angles to the body axis; tibia length 0.42 of SVL and foot length 0.58 of SVL; relative toe length I < II < V < III < IV; tips of toes round and slightly dilated; toes with relatively wide lateral fringes and rudiment of webs; one distinct subarticular tubercle at the bases of each toe; inner metatarsal tubercle long ovoid and the outer one absent.
Dorsal skin smooth with sparse granules; sparse tubercles on the flanks; a horn-like prominent tubercle on the edge; clear supratympanic fold curving postero-ventrally from posterior corner of eye to a level above insertion of arm; a discontinuous X-shaped ridge and two discontinuous dorsolateral ridges on two sides at the central back; sparse tubercles on the dorsal shank and thigh; ventral surface smooth; several tubercles on posterior hindlimbs; small pectoral gland closer to axilla; a single large femoral gland on rear of thigh.
Yellowish brown above in life; a dark interorbital triangle with light colored center and edge; a dark X-shaped making with light edge on the central of dorsum; dark brown transverse bands on forearms and hindlimbs; supratympanic fold light colored; dark vertical band below the eye; iris light brown with net-like stripes; throat and anterior chest reddish gray; a longitudinal stripe on the throat; a large white blotch with scarlet spots on the belly; one pair of lateroventral longitudinal broad reddish stripes on two sides; ventral limbs flesh color; ventral hands purplish, tips of fingers pale-grey, metacarpal tubercle reddish; ventral feet purplish brown, tips of fingers pale grey, metatarsal tubercle reddish; pectoral gland and femoral gland white.
Measurement data of type series are listed in Table
Megophrys (Panophrys) xiangnanensis sp. nov. is currently known only from Mt Yangming, southwestern Hunan. This toad inhabits areas near slowly flowing mountain streams surrounded by moist subtropical secondary evergreen broadleaf forests between elevations of 900–1400 m a.s.l. Males call from May to July, and during this time the males bear nuptial pads. Only one female individual was found, and tadpoles and other ecological information remain unknown.
The advertisement calls of Megophrys (Panophrys) xiangnanensis sp. nov. were recorded from the Holotype at 16 °C air temperature on 12 June 2014. Four calls with 98 notes are measured and the spectrograms are shown in Fig.
Measurements (in mm) of the type series of Megophrys (Panophrys) xiangnanensis sp. nov., * for the holotype.
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Sex | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Female |
SVL | 40.9 | 38.7 | 39.0 | 40.2 | 38.6 | 40.5 | 41.7 | 41.5 | 42.0 | 41.0 | 39.1 | 44.4 |
HDL | 13.2 | 12.6 | 12.3 | 13.1 | 12.9 | 13.0 | 13.1 | 13.2 | 13.4 | 13.0 | 12.8 | 14.0 |
HDW | 13.5 | 12.5 | 12.6 | 13.3 | 12.8 | 13.2 | 13.1 | 13.2 | 14.0 | 13.1 | 13.0 | 14.3 |
SNT | 4.5 | 4.4 | 4.3 | 4.4 | 4.3 | 4.6 | 4.2 | 4.5 | 4.7 | 4.6 | 4.3 | 5.0 |
IND | 4.5 | 4.2 | 4.2 | 4.5 | 4.3 | 4.0 | 4.4 | 4.6 | 4.5 | 4.5 | 4.3 | 4.5 |
IOD | 3.7 | 3.6 | 4.0 | 4.0 | 3.8 | 3.7 | 3.7 | 4.0 | 3.8 | 3.9 | 3.8 | 44.3 |
ED | 5.4 | 4.6 | 5.0 | 5.1 | 4.8 | 5.0 | 5.0 | 5.1 | 5.1 | 5.0 | 5.1 | 5.5 |
TD | 2.4 | 2.1 | 2.0 | 2.4 | 2.1 | 1.9 | 2.2 | 2.1 | 2.5 | 2.2 | 2.2 | 2.7 |
TED | 2.3 | 2.4 | 2.0 | 2.3 | 2.1 | 2.9 | 2.3 | 2.5 | 2.1 | 2.5 | 2.2 | 2.4 |
HND | 9.3 | 9.0 | 8.8 | 9.0 | 8.9 | 10.3 | 9.3 | 8.9 | 9.0 | 9.3 | 9.2 | 9.8 |
RAD | 9.0 | 8.7 | 8.8 | 8.8 | 8.9 | 9.8 | 9.3 | 8.9 | 9.0 | 9.3 | 9.2 | 9.8 |
FTL | 23.9 | 23.0 | 24.3 | 22.9 | 23.1 | 24.8 | 24.2 | 24.3 | 24.3 | 23.5 | 23.8 | 27.6 |
TIB | 17.0 | 17.9 | 17.8 | 17.0 | 17.9 | 18.2 | 18.2 | 18.2 | 17.3 | 17.4 | 17.7 | 19.1 |
Megophrys sp28 (
Seven adult specimens from the same locality as the holotype: female
The specific epithet yangmingensis refers to its type locality, Mt Yangming.
(1) Body size small, SVL 33.2–37.1 mm (35.3 ± 1.4, N = 7) in adult males and SVL 45.2 mm in adult female; (2) snout rounded in dorsal view; (3) tympanum clear, TD/ED 0.42–0.50; (4) presence of weak vomerine ridge, absence of vomerine teeth; (5) margin of tongue rounded, not notched behind; (6) hindlimbs slender, heels overlapping and tibio-tarsal articulation reaching forward at the anterior corner of the eye; (7) tibia 0.47–0.51 of SVL and foot 0.64–0.69 of SVL in males, while tibia 0.44 of SVL and foot 0.51 of SVL in female; (8) fingers without lateral fringes, presence of distinct subarticular tubercles at the bases; (9) toes with lateral fringes and rudiment of webs, presence of subarticular tubercles at the bases; (10) presence of small horn-like tubercle at the edge of upper eyelid; (11) dorsal skin rough with sparse granules, a discontinuous X-shaped ridge with two discontinuous dorsolateral ridges on two side on the back; (12) sparse tubercles on the flanks; (13) orange-brown or light brown above, a dark interorbital triangle with light colored center and edge, a dark X-shaped making with light edge on the central of dorsum; (14) single subgular vocal sac in males; (15) presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males.
Comparative data of Megophrys (Panophrys) yangmingensis sp. nov. with M. (Pa.) mirabilis sp. nov., M. (Pa.) shimentaina sp. nov., M. (Pa.) xiangnanensis sp. nov., and 42 recognized congeners of Panophrys are given in Table
Megophrys (Panophrys) yangmingensis sp. nov. differs from M. (Pa.) mirabilis sp. nov. by the smaller body size, SVL 33.2–37.1 mm in males and 45.2 mm in single female (vs. 55.8–61.4 mm in males and 68.5–74.8 in females), small horn-like tubercle at upper eyelid (vs. slightly large), presence of vocal sac in males (vs. absent), and presence of nuptial spines in adult males (vs. absent).
Megophrys (Panophrys) yangmingensis sp. nov. differs from M. (Pa.) shimentaina sp. nov. by the larger body size, SVL 33.2–37.1 mm in males (vs. 28.0–30.6 mm in males), absence of vomerine teeth (vs. present), and absence of tiny spines on the whole dorsal skin, flanks, dorsal limbs, the region around cloaca, and rear of hindlimbs (vs. such spines present).
Megophrys (Panophrys) yangmingensis sp. nov. differs from M. (Pa.) xiangnanensis sp. nov. by the smaller body size, SVL 33.2–37.1 mm in males (vs. 38.6–42.0), heels overlapping (vs. just meeting), tibio-tarsal articulation reaching forward at the anterior corner of the eye (vs. between eye and tympanum), and narrow lateral fringes on toes (vs. wide).
Compared with the five Panophrys species previously recorded from the hilly areas among Guangdong, Guangxi and Hunan, Megophrys (Panophrys) yangmingensis sp. nov. differs from M. (Pa.) acuta by the larger body size, SVL 33.2–37.1 mm in males and 45.2 mm in single female (vs. 27.1–33.0 mm in males and 28.1–33.6 mm in females), small horn-like tubercle at upper eyelid (vs. slightly large), snout rounded in dorsal view (vs. strongly remarkably pointed), and heels overlapping (vs. not meeting). M. (Pa.) yangmingensis sp. nov. differs from M. (Pa.) brachykolos by the narrow lateral fringes on toes (vs. absent), and heels overlapping (vs. not meeting). M. (Pa.) yangmingensis sp. nov. differs from M. (Pa.) nanlingensis by the absence of vomerine teeth (vs. present), and tongue not notched behind (vs. notched). M. (Pa.) yangmingensis sp. nov. differs from M. (Pa.) obesa by the narrow lateral fringes on toes (vs. absent), and heels overlapping (vs. not meeting). M. (Pa.) yangmingensis sp. nov. differs from M. (Pa.) shunhuangensis sp. nov. by the presence of vomerine ridge (vs. absence).
With a small body size, SVL 33.2–37.1 mm in adult males, Megophrys (Panophrys) yangmingensis sp. nov. is significantly different from nine congeners whose SVL > 40 mm in males, namely M. (Pa.) baolongensis, M. (Pa.) binlingensis, M. (Pa.) caudoprocta, M. (Pa.) jingdongensis, M. (Pa.) liboensis, M. (Pa.) omeimontis, M. (Pa.) sangzhiensis, M. (Pa.) shuichengensis, and M. (Pa.) spinata.
Megophrys (Panophrys) yangmingensis sp. nov. can be further distinguished from the remaining 28 congeners by the following characteristics: small horn-like tubercle at upper eyelid [vs. slightly large in M. (Pa.) jinggangensis, M. (Pa.) latidactyla, and M. (Pa.) palpebralespinosa]; vomerine teeth absent [vs. present in M. (Pa.) daweimontis, M. (Pa.) dongguanensis, M. (Pa.) fansipanensis, M. (Pa.) hoanglienensis, M. (Pa.) insularis, M. (Pa.) jinggangensis, M. (Pa.) jiulianensis, M. (Pa.) latidactyla, M. (Pa.) nankunensis, M. (Pa.) palpebralespinosa, and M. (Pa.) rubrimera]; tongue not notched behind [vs. notched in M. (Pa.) cheni, M. (Pa.) boettgeri, M. (Pa.) fansipanensis, M. (Pa.) hoanglienensis, M. (Pa.) huangshanensis, M. (Pa.) insularis, M. (Pa.) jiulianensis, M. (Pa.) kuatunensis, M. (Pa.) minor, and M. (Pa.) rubrimera]; lateral fringes on toes narrow [vs. absent in M. (Pa.) daweimontis, M. (Pa.) dongguanensis, M. (Pa.) fansipanensis, M. (Pa.) hoanglienensis, M. (Pa.) huangshanensis, M. (Pa.) insularis, M. (Pa.) jiangi, M. (Pa.) jiulianensis, M. (Pa.) leishanensis, M. (Pa.) lishuiensis, M. (Pa.) minor, M. (Pa.) nankunensis, M. (Pa.) ombrophila, M. (Pa.) tuberogranulatus, M. (Pa.) wugongensis, and M. (Pa.) wuliangshanensis; vs. wide in M. (Pa.) binchuanensis, M. (Pa.) boettgeri, M. (Pa.) cheni, M. (Pa.) latidactyla, M. (Pa.) lini, and M. (Pa.) palpebralespinosa; vs. absent in females while wide in males in M. (Pa.) wushanensis]; rudimentary webs on toes [vs. more than one-fourth webs in M. (Pa.) palpebralespinosa; vs. lacking webs in M. (Pa.) daweimontis, M. (Pa.) fansipanensis, M. (Pa.) hoanglienensis, M. (Pa.) huangshanensis, M. (Pa.) kuatunensis, M. (Pa.) lishuiensis, M. (Pa.) ombrophila, M. (Pa.) rubrimera, and M. (Pa.) wuliangshanensis]; tympanum clear with distinct edge [vs. upper 1/4 of tympanum concealed by supratympanic fold in M. (Pa.) mufumontana]; tibio-tarsal articulation reaching forward at the anterior corner of the eye [vs. between tympanum and eye in M. (Pa.) xianjuensis].
Adult male. Body size moderate, SVL 35.1 mm; head width slightly larger than head length, HDW/HDL 1.01; snout rounded in dorsal view, projecting, protruding well beyond margin of lower jaw; top of head flat; eyes large, ED 0.43 of HDL, pupil vertical; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum clear, TD/ED 0.43; large ovoid choanae at the base of the maxilla; presence of weak vomerine ridge, absence of vomerine teeth; margin of tongue rounded, not notched behind; presence of a single subgular vocal sac, a pair of slit-like openings at posterior of jaw.
Radio-ulna length 0.24 of SVL and hand 0.23 of SVL; hand without webs, fingers without lateral fringes, relative finger length II < I < IV < III; tips of fingers slightly dilated, round; one distinct subarticular tubercle at the bases of each finger; inner metacarpal tubercle observably enlarged and the outer one smaller; villiform black nuptial spines on the dorsal surface of the first and second fingers. Hindlimbs slender, tibio-tarsal articulation reaching forward at the anterior corner of eye when hindlimb is stretched along the side of the body; heels overlapping when the flexed hindlimbs are held at right angles to the body axis; tibia length 0.51 of SVL and foot length 0.67 of SVL; relative toe length I < II < V < III < IV; tips of toes round and slightly dilated; toes with lateral fringes and rudiment of webs; one subarticular tubercle at the bases of each toe; inner metatarsal tubercle long ovoid and the outer one absent.
Dorsal skin rough with sparse granules; sparse tubercles on the flanks and hindlimbs; several tubercles on upper eyelid, including a horn-like prominent tubercle on the edge; clear supratympanic fold curving postero-ventrally from posterior corner of eye to a level above insertion of arm; a discontinuous X-shaped ridge and two discontinuous dorsolateral ridges on two sides at the central back; four transverse skin ridges on the dorsal shank and thigh; ventral surface smooth; several granules on posterior hindlimbs; small pectoral gland closer to axilla; a single large femoral gland on rear of thigh.
Orange-brown above in life; a triangular making with light edge between eyes; a dark X-shaped making with light edge on the central of dorsum; supratympanic fold light brown; dark vertical band below the eye; iris orange-brown; throat and anterior chest purplish brown; belly dark gray with a large white blotch on the central; ventral limbs purplish; ventral hands reddish brown with dark stripes, tips of fingers pale-grey, metacarpal tubercle reddish; ventral feet purplish, tips of fingers pale-grey, metatarsal tubercle reddish; pectoral gland and femoral gland white.
Measurement data of type series are listed in Table
Currently, Megophrys (Panophrys) yangmingensis sp. nov. is only known from Mt Yangming, southwestern Hunan. This toad inhabits near flowing mountain streams over 1300 m a.s.l. Males call from early June to early September. Males found in early June bear well developed nuptial spines, while the spines are absent in males found in early September, suggesting the breeding season of this toad is before September. Only one female was found, and tadpoles and more ecological information remain unknown.
The advertisement calls of Megophrys (Panophrys) yangmingensis sp. nov. were recorded from the Holotype at 16 °C air temperature on 12 June 2014. Five calls with 160 notes are measured and the spectrograms are shown in Fig.
Measurements (in mm) of the type series of Megophrys (Panophrys) yangmingensis sp. nov., * for the holotype.
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Sex | Male | Male | Male | Male | Male | Male | Male | Female |
SVL | 35.1 | 34.5 | 36.6 | 33.2 | 37.1 | 36.4 | 34.5 | 45.2 |
HDL | 11.3 | 11.6 | 11.7 | 11.2 | 11.9 | 11.5 | 11.2 | 13.6 |
HDW | 11.4 | 11.9 | 11.7 | 11.1 | 11.8 | 11.5 | 11.3 | 13.5 |
SNT | 4.0 | 4.4 | 4.2 | 3.8 | 4.4 | 4.3 | 4.2 | 4.8 |
IND | 3.9 | 3.6 | 4.0 | 3.8 | 3.9 | 3.8 | 3.8 | 4.2 |
IOD | 3.4 | 3.8 | 3.8 | 3.4 | 3.8 | 3.6 | 3.7 | 4.4 |
ED | 4.9 | 4.7 | 4.6 | 4.8 | 4.7 | 5.0 | 4.7 | 5.6 |
TD | 2.1 | 2.2 | 2.3 | 2.2 | 2.2 | 2.1 | 2.3 | 2.8 |
TED | 1.5 | 1.4 | 1.4 | 1.4 | 2.1 | 1.7 | 1.6 | 2.1 |
HND | 8.0 | 8.2 | 8.2 | 7.0 | 8.2 | 8.2 | 7.9 | 10.1 |
RAD | 8.5 | 8.2 | 8.2 | 7.0 | 9.2 | 8.4 | 8.0 | 9.9 |
FTL | 24.2 | 23.3 | 23.4 | 21.1 | 24.1 | 24.1 | 23.2 | 23.0 |
TIB | 17.8 | 17.2 | 17.3 | 15.5 | 17.3 | 17.6 | 17.0 | 19.9 |
The phylogenetic analysis encompassing multilocus nuclear-gene and matrilineal mtDNA genealogy (
The genus Panophrys was established by
We thank to Yong-You Zhao, Run-Lin Li, Jian Wang, Yu-Long Li, Tian-Du Zhang, Jiang-Mo Zhang, Hai-Long He, Si-Yu Zhang, and Hong-Hui Chen for their help in the fieldwork. This work was supported by the Project of Comprehensive Scientific Survey of Luoxiao Mountains Region of Ministry of Science and Technology of PR China (No. 2013FY111500), the Specimen Platform of Ministry of Science and Technology of PR China, teaching specimens sub-platform (No.2005DKA21403-JK), the Biodiversity Conservation Programme of Ministry of Ecology and Environment of PR China, and the Project of Animal Diversity Survey and Monitoring System Construction of Guangdong Shimentai National Nature Reserve.
Specimens examined
Megophrys (Panophrys) acuta (10): China: Guangdong: Fengkai: Heishiding Nature Reserve (type locality):
Megophrys (Panophrys) binlingensis (2): China: Sichuan: Hongya: Mt. Wawu (type locality):
Megophrys (Panophrys) boettgeri (16): China: Fujian: Wuyishan: Mt. Wuyi (type locality):
Megophrys (Panophrys) brachykolos (2): China: Hong Kong (type locality):
Megophrys (Panophrys) caudoprocta (3): China: Hunan: Sangzhi: Badagongshan Nature Reserve (type locality):
Megophrys (Panophrys) cheni (19): China: Jiangxi: Jinggangshan: Mt. Jinggang (type locality):
Megophrys (Panophrys) huangshanensis (13): China: Anhui: Huangshan: Mt. Huangshan (type locality):
Megophrys (Panophrys) jingdongensis (14): China: Yunnan: Jingdong: Mt. Wuliang (type locality):
Megophrys (Panophrys) jinggangensis (11): China: Jiangxi: Jinggangshan: Mt. Jinggang (type locality):
Megophrys (Panophrys) kuatunensis (3): China: Fujian: Wuyishan: Guadun:
Megophrys (Panophrys) lini (27): China: Hunan: Yanling: Taoyuandong Nature Reserve (type locality):
Megophrys (Panophrys) minor (5): China: Sichuan: Dujiangyan: Mt. Qingcheng (type locality):
Megophrys (Panophrys) obesa (4): China: Guangdong: Fengkai: Heishiding Nature Reserve (type locality):
Megophrys (Panophrys) omeimontis (11): China: Sichuan: Emeishan: Mt. Emei (type locality):
Megophrys (Panophrys) sangzhiensis (6): China: Hunan: Sangzhi: Badagongshan Nature Reserve (type locality):
Megophrys (Panophrys) spinata (2): China: Guizhou: Leishan: Mt. Leigong (type locality):
Megophrys (Panophrys) tuberogranulatus (1): China: Hunan: Sangzhi: Badagongshan Nature Reserve (type locality):
Megophrys (Panophrys) wushanensis (5): China: Hubei: Shennongjia: Shennongjia Nature Reserve:
Megophrys (Panophrys) wuliangshanensis (5): China: Yunnan: Jingdong: Mt. Wuliang (type locality):
Megophrys (Xenophrys) glandulosa (13): China: Yunnan: Jingdong: Mt. Wuliang (type locality):
Megophrys (Xenophrys) cf. major (3): China: Yunnan: Mengla: Zhushihe:
Megophrys (Xenophrys) mangshanensis (10): China: Guangdong: Ruyuan: Nanling Nature Reserve:
Megophrys (Xenophrys) medogensis (3): China: Xizang: Motuo: Beibeng (type locality):
Megophrys (Xenophrys) pachyproctus (1): China: Xizang: Motuo: Beibeng (type locality):