Research Article |
Corresponding author: Bin Wang ( wangbin@cib.ac.cn ) Academic editor: Annemarie Ohler
© 2020 Shi-Ze Li, Ning-Ning Lu, Jing Liu, Bin 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:
Li S-Z, Lu N-N, Liu J, Wang B (2020) Description of a new Megophrys Kuhl & Van Hasselt, 1822 (Anura, Megophryidae) from Guizhou Province, China. ZooKeys 986: 101-126. https://doi.org/10.3897/zookeys.986.57119
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A new species of the genus Megophrys is described from Guizhou Province, China. Molecular phylogenetic analyses based on mitochondrial DNA indicated the new species as a clade clustered into the Megophrys clade. The new species can be distinguished from its congeners by a combination of the following characters: body size moderate (SVL 40.0–45.5 mm in males and 48.9–51.2 mm in females); vomerine teeth absent; tongue not notched behind; tympanum distinctly visible, oval; a small horn-like tubercle at the edge of each upper eyelid; two metacarpal tubercles in hand; toes with rudimentary webbing; heels overlapping when thighs are positioned at right angles to the body; tibiotarsal articulation reaching the level of mid-eye when leg stretched forward; in breeding males, an internal single subgular vocal sac present and brownish nuptial pads, made up of black nuptial spines, present on the dorsal base of the first two fingers.
Molecular phylogenetic analysis, morphology, new species, taxonomy
The Asian horned toad Megophrys Kuhl & Van Hasselt, 1822 (Anura: Megophryidae Bonaparte, 1850) is widely distributed in eastern and central China, throughout southeastern Asia, and extending to the islands of the Sunda Shelf and the Philippines (
During field surveys in Anlong County, Guizhou Province, China, we collected eight Megophrys specimens. Molecular phylogenetic analyses and morphological comparisons supported it as an undescribed species and it is described herein as a new species.
Three adult females and five adult males of the undescribed species were collected from Anlong County, Guizhou Province, China (Fig.
Measurements of the adult specimens of Megophrys anlongensis sp. nov. Units given in mm. See abbreviations for the morphological characters in Materials and methods section.
Voucher | Sex | SVL | HDL | HDW | SL | IND | IOD | UEW | ED | TYD | LAL | LW | HLL | THL | TL | TW | TFL | FL |
CIBAL20190531021 | ♂ | 45.5 | 12.3 | 15.9 | 6.0 | 5.6 | 4.1 | 4.0 | 4.7 | 3.1 | 19.9 | 5.0 | 70.8 | 18.8 | 22.5 | 5.2 | 33.1 | 22.7 |
CIBAL20190531019 | ♂ | 41.1 | 12.4 | 14.0 | 5.0 | 4.5 | 3.2 | 4.3 | 4.8 | 2.7 | 18.8 | 4.3 | 65.4 | 17.7 | 22.3 | 5.1 | 29.4 | 19.9 |
CIBAL20190531017 | ♂ | 42.5 | 11.5 | 14.4 | 5.1 | 4.7 | 4.1 | 3.7 | 4.4 | 3.1 | 19.7 | 4.2 | 67.7 | 20.2 | 22.1 | 5.5 | 31.1 | 21.5 |
CIBAL20190531020 | ♂ | 42.5 | 11.6 | 14.5 | 6.0 | 5.2 | 3.4 | 4.3 | 5.1 | 3.5 | 19.4 | 4.5 | 63.9 | 19.9 | 20.9 | 5.2 | 29.2 | 19.0 |
CIBAL20190531018 | ♂ | 40.0 | 13.0 | 14.5 | 5.6 | 4.7 | 4.0 | 3.8 | 4.6 | 2.7 | 19.1 | 3.8 | 65.2 | 19.7 | 21.2 | 5.0 | 29.6 | 20.0 |
Range of males | 40.0–45.5 | 11.5–13.0 | 14.0–15.9 | 5.0–6.0 | 4.5–5.6 | 3.2–4.1 | 3.7–4.3 | 4.4–5.1 | 2.7–3.5 | 18.8–19.9 | 3.8–5.0 | 63.9–70.8 | 17.7–20.2 | 20.9–22.5 | 5.0–5.5 | 29.2–33.1 | 19.0–22.7 | |
Mean ± SD of males | 42.3 ± 2.04 | 12.1 ± 0.62 | 14.6 ± 0.71 | 5.5 ± 0.48 | 4.9 ± 0.45 | 3.8 ± 0.41 | 4.0 ± 0.28 | 4.7 ± 0.25 | 3.0 ± 0.36 | 19.4 ± 0.44 | 4.34 ± 0.44 | 66.6 ± 2.70 | 19.3 ± 1.03 | 21.8 ± 0.73 | 5.2 ± 0.18 | 30.3 ± 1.64 | 20.6 ± 1.47 | |
CIBAL20190531022 | ♀ | 51.2 | 12.9 | 17.4 | 6.5 | 5.7 | 4.2 | 4.7 | 4.9 | 3.3 | 23.6 | 4.0 | 83.7 | 26.0 | 27.5 | 6.0 | 38.6 | 25.6 |
CIBAL20190811015 | ♀ | 48.9 | 13.1 | 16.1 | 5.5 | 5.1 | 3.1 | 4.1 | 4.9 | 2.9 | 23.8 | 3.7 | 83.4 | 24.7 | 26.2 | 5.3 | 38.0 | 26.0 |
CIBAL20190811014 | ♀ | 49.4 | 13.2 | 16.5 | 6.0 | 5.3 | 4.0 | 4.5 | 5.1 | 3.1 | 24.5 | 3.3 | 88.2 | 25.4 | 28.0 | 5.0 | 40.4 | 20.5 |
Range of females | 48.9–51.2 | 12.9–13.2 | 16.0–17.0 | 5.5–6.5 | 5.1–5.7 | 3.1–4.2 | 4.1–4.7 | 4.9–5.1 | 2.9–3.3 | 23.6–24.5 | 3.3–4.0 | 83.4–88.2 | 24.7–26.0 | 26.2–28.0 | 5.0–6.0 | 38.0–40.4 | 20.5–26.0 | |
Mean ± SD of females | 49.8 ±1.21 | 13.1 ±0.17 | 16.7 ±0.66 | 6.0 ±0.50 | 5.4 ±0.31 | 3.8 ±0.59 | 4.4 ±0.31 | 5.0 ±0.12 | 3.1 ±0.21 | 24.0 ±0.48 | 3.7 ±0.35 | 85.1 ±2.68 | 25.4 ±0.64 | 27.2 ±0.92 | 5.4 ±0.51 | 39.0 ±1.24 | 24.0 ±3.06 |
Six specimens of the undescribed species were included in the molecular analyses (Table
For molecular analyses, the available sequence data for congeners of Megophrys were downloaded from GenBank (Table
Information for samples used in molecular phylogenetic analyses in this study.
ID | Species | Voucher number | Locality | GenBank accession number | |
---|---|---|---|---|---|
16S | COI | ||||
1 | Megophrys anlongensis sp. nov. | CIBAL20190531018 | Anlong County, Guizhou, China | MT823184 | MT823261 |
2 | CIBAL20190531017 | Anlong County, Guizhou, China | MT823185 | MT823262 | |
3 | CIBAL20190531022 | Anlong County, Guizhou, China | MT823186 | MT823263 | |
4 | CIBAL20190811014 | Anlong County, Guizhou, China | MT823187 | MT823264 | |
5 | CIBAL20190811015 | Anlong County, Guizhou, China | MT823188 | MT823265 | |
6 | CIBAL20190531019 | Anlong County, Guizhou, China | MT823189 | MT823266 | |
7 | Megophrys nankunensis | SYS a004498 | Nankun Shan, Guangdong, China | MK524108 | MK524139 |
8 | Megophrys dongguanensis | SYS a001972 | Yinping Shan, Guangdong, China | MK524098 | MK524129 |
9 | Megophrys cheni | SYS a001427 | Jinggang Shan, Jiangxi, China | KJ560391 | – |
10 | Megophrys obesa | SYS a002272 | Heishiding Nature Reserve, Guangdong, China | KJ579122 | – |
11 | Megophrys ombrophila | KRM18 | Wuyishan, Fujian, China | KX856404 | – |
12 | Megophrys wugongensis | SYS a002610 | Wugongshan Scenic Area, Jiangxi, China | MK524114 | MK524145 |
13 | Megophrys lini | SYS a002370 | Suichuan, Jiangxi, China | KJ560412 | – |
14 | Megophrys xiangnanensis | SYS a002874 | Yangming Shan, Hunan, China | MH406713 | MH406165 |
15 | Megophrys nanlingensis | SYS a001959 | Nanling Nature Reserve, Guangdong, China | MK524111 | MK524142 |
16 | Megophrys kuatunensis | SYS a001579 | Wuyi Shan, Fujian, China | KJ560376 | – |
17 | Megophrys jinggangensis | KIZ07132 | Chashan Forest Farm, Jiangxi, China | KX811840 | KX812108 |
18 | Megophrys xianjuensis | CIBXJ190505 | Xianju, Zhejiang, China | MN563753 | MN563769 |
19 | Megophrys lishuiensis | WYF00169 | Lishui, Zhejiang, China | KY021418 | – |
20 | Megophrys huangshanensis | KIZ022004 | Huang Shan, Anhui, China | KX811821 | KX812107 |
21 | Megophrys boettgeri | Tissue ID: YPXJK033 | Wuyi Shan, Fujian, China | KX811814 | KX812104 |
22 | Megophrys liboensis | GNUG:20160408003 | Libo, Guizhou, China | MF285262 | – |
23 | Megophrys mufumontana | SYS a006391 | Mufu Shan, Hunan, China | MK524105 | MK524136 |
24 | Megophrys mirabilis | SYS a002192 | Huaping Nature Reserve, Guangxi, China | MH406669 | MH406109 |
25 | Megophrys shunhuangensis | HNNU16SH02 | Shunhuang Mountains, Hunan, China | MK836037 | – |
26 | Megophrys acuta | SYS a001957 | Heishiding Nature Reserve, Guangdong, China | KJ579118 | – |
27 | Megophrys leishanensis | CIBLS20171101001 | Leigong Shan, Guizhou, China | MK005310 | MK005306 |
28 | Megophrys shimentaina | SYS a002077 | Shimentai Nature Reserve Guangdong, China | MH406655 | MH406092 |
29 | Megophrys yangmingensis | SYS a002877 | Yangming Shan, Hunan, China | MH406716 | MH406168 |
30 | Megophrys jiulianensis | SYS a002107 | Jiulian Shan, Jiangxi, China | MK524099 | MK524130 |
31 | Megophrys wushanensis | KIZ045469 | Guangwu Shan, Sichuan, China | KX811838 | KX812094 |
32 | Megophrys baolongensis | KIZ019216 | Baolong, Chongqing, China | KX811813 | KX812093 |
33 | Megophrys tuberogranulata | Tissue ID: YPX10987 | Badagongshan Nature Reserve, Hunan, China | KX811823 | KX812095 |
34 | Megophrys binchuanensis | KIZ019441 | Jizu Shan, Yunnan, China | KX811849 | KX812112 |
35 | Megophrys sangzhiensis | SYSa004307 | Zhangjiajie, Hunan, China | MH406798 | MH406260 |
36 | Megophrys spinata | SYSa002227 | Leigong Shan, Guizhou, China | MH406676 | MH406116 |
37 | Megophrys binlingensis | SYSa005313 | Wawu Shan, Sichuan, China | MH406892 | MH406354 |
38 | Megophrys angka | KIZ040591 | Kiew Mae Pan nature trail, Chiang Mai, Thailand | MN508052 | – |
39 | Megophrys omeimontis | KIZ025765 | Emei Shan, Sichuan, China | KX811884 | KX812136 |
40 | Megophrys palpebralespinosa | KIZ011603 | Pu Hu Nature Reserve, Thanh Hoa, Vietnam | KX811888 | KX812137 |
41 | Megophrys jingdongensis | KIZ-LC0805067 | Huanglianshan National Nature Reserve, Yunnan, China | KX811872 | KX812131 |
42 | Megophrys daweimontis | KIZ048997 | Dawei Shan, Yunnan, China | KX811867 | KX812125 |
43 | Megophrys wuliangshanensis | KIZ046812 | Huangcaoling, Yunnan, China | KX811881 | KX812129 |
44 | Megophrys fansipanensis | VNMN 2018.01 | Lao Cai, Sa Pa, Vietnam | MH514886 | – |
45 | Megophrys hoanglienensis | VNMN 2018.02 | Lao Cai, Sa Pa, Vietnam | MH514889 | – |
46 | Megophrys minor | KIZ01939 | Qingcheng Shan, Sichuan, China | KX811896 | KX812145 |
47 | Megophrys jiangi | CIBKKS20180722006 | Kuankuosui Nature Reserve, Guizhou, China | MN107743 | MN107748 |
48 | Megophrys chishuiensis | CIBCS20190518031 | Chishui Nature Reserve, Guizhou, China | MN954707 | MN928958 |
49 | Megophrys brachykolos | ROM 16634 | Hong Kong, China | KX811897 | KX812150 |
50 | Megophrys elfina | ZMMU ABV-00454 | Bidoup Mountain, Lam Dong, Vietnam | KY425379 | – |
51 | Megophrys gerti | ITBCZ 1108 | Nui Chua National Park, Ninh Thuan, Vietnam | KX811917 | KX812161 |
52 | Megophrys synoria | FMNH 262778 | O’Reang, Mondolkiri, Cambodia | KY022198 | – |
53 | Megophrys microstoma | KIZ048799 | Xiaoqiaogou Nature Reserve, Yunnan, China | KX811914 | KX812156 |
54 | Megophrys hansi | KIZ010360 | Phong Dien Nature Reserve, Thua Thien Hue, Vietnam | KX811913 | KX812155 |
55 | Megophrys pachyproctus | KIZ010978 | Beibeng, Xizang, China | KX811908 | KX812153 |
56 | Megophrys baluensis | ZMH A13125 | Gunung Kinabalu National Park, Kogopan Trail, Malaysia | KJ831310 | – |
57 | Megophrys stejnegeri | KU 314303 | Pasonanca Natural Park, Zamboanga, Philippines | KX811922 | KX812052 |
58 | Megophrys ligayae | ZMMU NAP-05015 | Palawan, Philippines | KX811919 | KX812051 |
59 | Megophrys nasuta | KIZ019419 | Malaysia | KX811921 | KX812054 |
60 | Megophrys kobayashii | UNIMAS 8148 | Gunung Kinabalu National Park, Sabah, Malaysia | KJ831313 | – |
61 | Megophrys edwardinae | FMNH 273694 | Bintulu, Sarawak, Malaysia | KX811918 | KX812050 |
62 | Megophrys aceras | KIZ025467 | Khao Nan National Park, Nakhon Si Thammarat, Thailand | KX811925 | KX812159 |
63 | Megophrys zhangi | KIZ014278 | Zhangmu, Xizang, China | KX811765 | KX812084 |
64 | Megophrys sanu | K5198/ZSI11393 | – | KX894679 | – |
65 | Megophrys katabhako | ZSIA11799 | – | KX894669 | – |
66 | Megophrys periosa | BNHS 6061 | West Kameng dist., Arunachal Pradesh, India | KY022309 | MH647528 |
67 | Megophrys himalayana | SDBDU2009.75 | East Siang dist., Arunachal Pradesh, India | KY022311 | – |
68 | Megophrys glandulosa | KIZ048439 | Husa, Yunnan, China | KX811762 | KX812075 |
69 | Megophrys medogensis | KIZ06621 | Beibeng, Xizang, China | KX811767 | KX812082 |
70 | Megophrys flavipunctata | SDBDU2009.297 | East Khasi Hills dist., Meghalaya, India | KY022307 | MH647536 |
71 | Megophrys maosonensis | KIZ016045 | Xiaoqiaogou Nature Reserve, Yunnan, China | KX811780 | KX812080 |
72 | Megophrys mangshanensis | KIZ021786 | Nanling National Forest Park, Guangdong, China | KX811790 | KX812079 |
73 | Megophrys oreocrypta | BNHS 6046 | West Garo Hills dist., Meghalaya, India | KY022306 | – |
74 | Megophrys major | SYSa002961 | Zhushihe, Yunnan, China | MH406728 | MH406180 |
75 | Megophrys parva | SYSa003042 | Zhushihe, Yunnan, China | MH406737 | MH406189 |
76 | Megophrys auralensis | NCSM 79599 | Aural, Kampong Speu, Cambodia | KX811807 | – |
77 | Megophrys dringi | UNIMAS 8943 | Gunung Mulu National Park, Sarawak, Malaysia | KJ831317 | – |
78 | Megophrys gigantica | SYSa003933 | Wuliang shan, Yunnan, China | MH406775 | MH406235 |
79 | Megophrys shapingensis | KIZ014512 | Liziping Nature Reserve, Sichuan, China | KX811904 | KX812060 |
80 | Megophrys wawuensis | KIZ025799 | Wawu Shan, Sichuan, China | KX811902 | KX812062 |
81 | Megophrys nankiangensis |
|
Nanjiang, Sichuan, China | KX811900 | – |
82 | Megophrys lancip | MZB:Amp:22233 | – | KY679891 | – |
83 | Megophrys montana | LSUMZ 81916 | Sukabumi, Java, Indonesia | KX811927 | KX812163 |
84 | Megophrys popei | SYS a000589 | Naling Nature Reserve, Guangdong, China | KM504251 | – |
85 | Megophrys carinense | Tissue ID: YPX20455 | Dayao Shan, Guangxi, China | KX811811 | KX812057 |
86 | Megophrys feae | KIZ046706 | Huangcaoling, Yunnan, China | KX811810 | KX812056 |
87 | Megophrys chuannanensis | CIB20050081 | Hejiang, Sichuan, China | KM504261 | – |
88 | Megophrys intermedia | ZFMK 87596 | U Bo, Phong Nha-Ke Bang NP, Vietnam | HQ588950 | – |
89 | Leptobrachium boringii | Tissue ID: YPX37539 | Emei Shan, Sichuan, China | KX811930 | KX812164 |
90 | Leptobrachella oshanensis | KIZ025778 | Emei Shan, Sichuan, China | KX811928 | KX812166 |
All eight adult specimens of the undescribed species were measured (Table
ED eye diameter (distance from the anterior corner to the posterior corner of the eye);
FL foot length (distance from tarsus to the tip of fourth toe);
HDL head length (distance from the tip of the snout to the articulation of jaw);
HDW maximum head width (greatest width between the left and right articulations of jaw);
HLL hindlimb length (maximum length from the vent to the distal tip of the Toe IV);
IND internasal distance (minimum distance between the inner margins of the external nares);
IOD interorbital distance (minimum distance between the inner edges of the upper eyelids);
LAL length of lower arm and hand (distance from the elbow to the distal end of the Finger IV);
LW lower arm width (maximum width of the lower arm);
SVL snout-vent length (distance from the tip of the snout to the posterior edge of the vent);
SL snout length (distance from the tip of the snout to the anterior corner of the eye);
TFL length of foot and tarsus (distance from the tibiotarsal articulation to the distal end of the Toe IV);
THL thigh length (distance from vent to knee);
TL tibia length (distance from knee to tarsus);
TW maximal tibia width;
TYD maximal tympanum diameter;
UEW upper eyelid width (greatest width of the upper eyelid margins measured perpendicular to the anterior-posterior axis).
The undescribed species was also compared with all other congeners on morphology. Comparative data were obtained from related species as described in literature (Table
Bibliographic references for morphological characters for congeners of the genus Megophrys.
Species | Literature obtained |
---|---|
M. aceras Boulenger, 1903 |
|
M. acuta Wang, Li & Jin, 2014 |
|
M. ancrae Mahony, Teeling & Biju, 2013 |
|
M. angka Wu, Suwannapoom, Poyarkov, Chen, Pawangkhanant, Xu, Jin, Murphy & Che, 2019 |
|
M. auralensis Ohler, Swan & Daltry, 2002 |
|
M. baluensis (Boulenger, 1899) |
|
M. baolongensis Ye, Fei & Xie, 2007 |
|
M. binchuanensis Ye & Fei, 1995 |
|
M. binlingensis Jiang, Fei & Ye, 2009 |
|
M. boettgeri (Boulenger, 1899) |
|
M. brachykolos Inger & Romer, 1961 |
|
M. carinense (Boulenger, 1889) |
|
M. caobangensis Nguyen, Pham, Nguyen, Luong, and Ziegler, 2020 |
|
M. caudoprocta Shen, 1994 |
|
M. cheni (Wang & Liu, 2014) |
|
M. chishuiensis Xu, Li, Liu, Wei & Wang, 2020 |
|
M. chuannanensis (Fei, Ye & Huang, 2001) |
|
M. damrei Mahony, 2011 |
|
M. daweimontis Rao & Yang, 1997 |
|
M. dongguanensis Wang & Wang, 2019 |
|
M. dringi Inger, Stuebing & Tan, 1995 |
|
M. edwardinae Inger, 1989 |
|
M. elfina Poyarkov, Duong, Orlov, Gogoleva, Vassilieva, Nguyen, Nguyen, Nguyen, Che & Mahony, 2017 |
|
M. fansipanensis Tapley, Cutajar, Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Portway, Luong & Rowley, 2018 |
|
M. feae Boulenger, 1887 |
|
M. feii Yang, Wang & Wang, 2018 |
|
M. flavipunctata Mahony, Kamei, Teeling & Biju, 2018 |
|
M. gerti (Ohler, 2003) |
|
M. gigantica Liu, Hu & Yang, 1960 |
|
M. glandulosa Fei, Ye & Huang, 1990 |
|
M. hansi (Ohler, 2003) |
|
M. himalayana Mahony, Kamei, Teeling & Biju, 2018 |
|
M. hoanglienensis Tapley, Cutajar, Mahony, Nguyen, Dau, Luong, Le, Nguyen, Nguyen, Portway, Luong & Rowley, 2018 |
|
M. huangshanensis Fei & Ye, 2005 |
|
M. insularis (Wang, Liu, Lyu, Zeng & Wang, 2017) |
|
M. intermedia Smith, 1921 |
|
M. jiangi Liu, Li, Wei, Xu, Cheng, Wang & Wu, 2020 |
|
M. jingdongensis Fei & Ye, 1983 |
|
M. jinggangensis (Wang, 2012) |
|
M. jiulianensis Wang, Zeng, Lyu & Wang, 2019 |
|
M. kalimantanensis Munir, Hamidy, Matsui, Iskandar, Sidik & Shimada, 2019 |
|
M. kobayashii Malkmus & Matsui, 1997 |
|
M. koui Mahony, Foley, Biju & Teeling, 2017 |
|
M. kuatunensis Pope, 1929 |
|
M. lancip Munir, Hamidy, Farajallah & Smith, 2018 |
|
M. leishanensis Li, Xu, Liu, Jiang, Wei & Wang, 2018 |
|
M. lekaguli Stuart, Chuaynkern, Chan-ard & Inger, 2006 |
|
M. liboensis (Zhang, Li, Xiao, Li, Pan, Wang, Zhang & Zhou, 2017) |
|
M. ligayae Taylor, 1920 |
|
M. lini (Wang & Yang, 2014) |
|
M. lishuiensis (Wang, Liu & Jiang, 2017) |
|
M. longipes Boulenger, 1886 |
|
M. major Boulenger, 1908 |
|
M. mangshanensis Fei & Ye, 1990 |
|
M. maosonensis Bourret, 1937 |
|
M. medogensis Fei, Ye & Huang, 1983 |
|
M. megacephala Mahony, Sengupta, Kamei & Biju, 2011 |
|
M. microstoma (Boulenger, 1903) |
|
M. minor Stejneger, 1926 |
|
M. mirabilis Lyu, Wang & Zhao |
|
M. montana Kuhl & Van Hasselt, 1822 |
|
M. monticola (Günther, 1864) |
|
M. mufumontana Wang, Lyu & Wang, 2019 |
|
M. nankiangensis Liu & Hu, 1966 | Hu and Liu 1966 |
M. nankunensis Wang, Zeng &. Wang, 2019 |
|
M. nanlingensis Lyu, Wang, Liu & Wang, 2019 |
|
M. nasuta (Schlegel, 1858) |
|
M. obesa Wang, Li & Zhao, 2014 |
|
M. ombrophila Messenger & Dahn, 2019 |
|
M. omeimontis Liu, 1950 |
|
M. oreocrypta Mahony, Kamei, Teeling & Biju, 2018 |
|
M. oropedion Mahony, Teeling & Biju, 2013 |
|
M. orientalis Li, Lyu, Wang & Wang, 2020 |
|
M. pachyproctus Huang, 1981 |
|
M. palpebralespinosa Bourret, 1937 |
|
M. parallela Inger & Iskandar, 2005 |
|
M. parva (Boulenger, 1893) |
|
M. periosa Mahony, Kamei, Teeling & Biju, 2018 |
|
M. popei (Zhao, Yang, Chen, Chen & Wang, 2014) |
|
M. robusta Boulenger, 1908 |
|
M. rubrimera Tapley, Cutajar, Mahony, Chung, Dau, Nguyen, Luong & Rowley, 2017 |
|
M. sangzhiensis Jiang, Ye & Fei, 2008 |
|
M. serchhipii (Mathew & Sen, 2007) |
|
M. shapingensis Liu, 1950 |
|
M. shimentaina Lyu, Liu & Wang |
|
M. shuichengensis Tian & Sun, 1995 |
|
M. shunhuangensis Wang, Deng, Liu, Wu & Liu, 2019 |
|
M. spinata Liu & Hu, 1973 |
|
M. stejnegeri Taylor, 1920 |
|
M. synoria (Stuart, Sok & Neang, 2006) |
|
M. takensis Mahony, 2011 |
|
M. tuberogranulata Shen, Mo & Li, 2010 | Mo et al. 2012 |
M. vegrandis Mahony, Teeling, Biju, 2013 |
|
M. wawuensis Fei, Jiang & Zheng, 2001 |
|
M. wugongensis Wang, Lyu & Wang, 2019 |
|
M. wuliangshanensis Ye & Fei, 1995 |
|
M. wushanensis Ye & Fei, 1995 |
|
M. xianjuensis Wang, Wu, Peng, Shi, Lu & Wu, 2020 |
|
M. xiangnanensis Lyu, Zeng & Wang |
|
M. yangmingensis Lyu, Zeng & Wang |
|
M. zhangi Ye & Fei, 1992 |
|
M. zunhebotoensis (Mathew & Sen, 2007) |
|
The advertisement calls of the undescribed species were recorded from the holotype specimen CIBAL20190531018 in the field on 31 May 2019 in Anlong County, Guizhou Province, China. The advertisement call of the undescribed species was recorded in the stream at ambient air temperature of 18.5 °C and air humidity of 83%. SONY PCM-D50 digital sound recorder was used to record within 30 cm of the calling individual. The sound files in wave format were resampled at 48 kHz with sampling depth 24 bits. The sonograms and waveforms were generated by WaveSurfer software (
Aligned sequence matrix of 16S+COI contains 1104 bp. ML and BI trees of the mitochondrial DNA dataset presented almost consistent topology (Fig.
Genetic distances between samples of the undescribed species either on 16S or on COI genes were below 0.2% much lower than the interspecific genetic distance between recognised Megophrys species (Suppl. materials
The new species could be identified from its congeners in a series of morphological characters (Suppl. material
CIBAL20190531018 (Figs
Four adult males and three females from the same place as holotype collected by Shi-Ze Li and Jing Liu. CIBAL20190531017, CIBAL20190531019, CIBAL20190531021 and CIBAL20190531022 collected on 31 May 2019 by Jing Liu, and CIBAL20190811014 and CIBAL20190811015 collected by Shi-Ze Li on 11 August 2019.
Megophrys anlongensis sp. nov. is assigned to the genus Megophrys based on molecular phylogenetic analyses and the following generic diagnostic characters: snout shield-like, projecting beyond the lower jaw; canthus rostralis distinct; chest glands small and round, closer to the axilla than to midventral line; femoral glands on rear part of thigh; vertical pupils.
Megophrys anlongensis sp. nov. could be distinguished from its congeners by a combination of the following morphological characters: (1) body size moderate (SVL 40.0–45.5 mm in males and 48.9–51.2 mm in females); (2) vomerine teeth absent; (3) tongue not notched behind; (4) a small horn-like tubercle at the edge of each upper eyelid; (5) tympanum distinctly visible, oval; (6) two metacarpal tubercles on hand; (7) toes with rudimentary webbing; (8) heels overlapping when thighs are positioned at right angles to the body; (9) tibiotarsal articulation reaching the level of mid-eye when leg stretched forward; (10) an internal single subgular vocal sac in male; (11) in breeding males, brownish nuptial pads, made up of black nuptial spines, present on the dorsal base of the first two fingers.
(Figs
Forelimbs slender, the length of lower arm and hand 47.9% of SVL; fingers slender, relative finger lengths: I < II < V < III; tips of digits globular, without lateral fringes; subarticular tubercle distinct at the base of each finger; two metacarpal tubercles, prominent, oval-shaped, the inner one bigger than the outer one.
Hindlimbs slender; heels overlapping when thighs are positioned at right angles to the body; tibiotarsal articulation reaching the middle eye when leg stretched forward; tibia length longer than thigh length; relative toe lengths I < II < V < III < IV; tips of toes round, slightly dilated; subarticular tubercles present on each toes; toes with rudimentary webbing and narrow lateral fringe; inner metatarsal tubercle oval-shaped; outer metatarsal tubercle absent.
Dorsal skin rough, several large warts scattered on flanks; a small horn-like tubercle at the edge of each upper eyelid; tubercles on the dorsum forming a weak X-shaped ridge, two dorsolateral parallel ridges on either side of the X-shaped ridges; an inverted triangular brown speckle between two upper eyelids; several tubercles on the flanks and dorsal surface of thighs and tibias; supratympanic fold distinct.
Ventral surface smooth; numerous granules scattered on flanks; glands on chest indistinct; numerous white granules on outer thighs and posterior end of the body distinctly protruding and forming an arc-shaped swelling above the anal region.
(Fig.
(Fig.
In CIBAL20190531017 the inverted triangular brown speckle is connected to the X-shape ridge (Fig.
Colour variation in Megophrys anlongensis sp. nov. A dorsolateral view of the specimen CIBAL20190531017 B ventral view of the male specimen CIBAL20190531017 C dorsolateral view of the specimen CIBAL20190531022 D ventral view of the specimen CIBAL20190531022 E dorsolateral view of the specimen CIBAL20190811014 F ventral view of the specimen CIBAL20190811014.
The call description is based on recordings of the holotype CIBAL20190531018 (Fig.
Adult males have a single subgular vocal sac. In breeding males, brownish nuptial pads, made up of black nuptial spines, present on the dorsal bases of the first two fingers (Fig.
By body size medium, Megophrys anlongensis sp. nov. differs from M. aceras, M. acuta, M. angka, M. auralensis, M. binchuanensis, M. boettgeri, M. caobangensis, M. cheni, M. daweimontis, M. dringi, M. elfina, M. feii, M. gerti, M. jinggangensis, M. jiulianensis, M. kuatunensis, M. leishanensis, M. lishuiensis, M. microstoma, M. mufumontana, M. nankunensis, M. nanlingensis, M. obesa, M. ombrophila, M. oropedion, M. pachyproctus, M. palpebralespinosa, M. rubrimera, M. serchhipii, M. shimentaina, M. shunhuangensis, M. vegrandis, M. wugongensis, M. wuliangshanensis, M. wushanensis, M. xianjuensis, M. yangmingensis, M. zhangi, and M. zunhebotoensis (SVL > 40.0 mm in the new species vs. maximum SVL < 39.0 mm in the latter), and differs from M. carinense, M. caudoprocta, M. chuannanensis, M. damrei, M. feae, M. flavipunctata, M. gigantica, M. glandulosa, M. himalayana, M. kalimantanensis, M. kobayashii, M. lekaguli, M. ligayae, M. mangshanensis, M. medogensis, M. mirabilis, M. nasuta, M. omeimontis, M. orientalis, M. periosa, M. platyparietus, M. popei, M. sangzhiensis, M. shapingensis, and M. shuichengensis (maximum SVL < 52.0 mm in the new species vs. minimum SVL > 54.0 mm in the latter), and differs from M. edwardinae and M. monticola (SVL 48.9–51.2 mm in female in the new species vs. 69–82 mm in M. edwardinae and 40.5 mm in M. monticola).
By vomerine teeth absent, Megophrys anlongensis sp. nov. differs from M. ancrae, M. baluensis, M. carinense, M. caudoprocta, M. chuannanensis, M. damrei, M. daweimontis, M. dongguanensis, M. fansipanensis, M. feae, M. flavipunctata, M. glandulosa, M. himalayana, M. hoanglienensis, M. insularis, M. intermedia, M. jingdongensis, M. jinggangensis, M. jiulianensis, M. kalimantanensis, M. kobayashii, M. lancip, M. lekaguli, M. liboensis, M. ligayae, M. longipes, M. mangshanensis, M. maosonensis, M. medogensis, M. megacephala, M. montana, M. nankunensis, M. nanlingensis, M. nasuta, M. omeimontis, M. oreocrypta, M. orientalis, M. oropedion, M. pachyproctus, M. palpebralespinosa, M. parallela, M. parva, M. periosa, M. platyparietus, M. popei, M. robusta, M. rubrimera, M. serchhipii, M. shimentaina, M. stejnegeri, M. takensis, M. zhangi, and M. zunhebotoensis (vs. present in the latter).
By a small horn-like tubercle at the edge of each upper eyelid, Megophrys anlongensis sp. nov. differs from M. aceras, M. acuta, M. carinense, M. caudoprocta, M. chuannanensis, M. feae, M. gerti, M. hansi, M. intermedia, M. intermedia, M. jinggangensis, M. kalimantanensis, M. koui, M. lancip, M. liboensis, M. microstoma, M. montana, M. nasuta, M. orientalis, M. palpebralespinosa, M. platyparietus, M. popei, M. shuichengensis, M. stejnegeri, and M. synoria (vs. having a prominent and elongated tubercle in the latter).
By tongue not notched behind, Megophrys anlongensis sp. nov. differs from M. ancrae, M. baolongensis, M. binlingensis, M. boettgeri, M. carinense, M. cheni, M. chuannanensis, M. damrei, M. dringi, M. fansipanensis,M. feae, M. feii, M. flavipunctata, M. gerti, M. glandulosa, M. hoanglienensis, M. huangshanensis, M. insularis, M. jiulianensis. M. jingdongensis, M. kalimantanensis , M. kuatunensis, M. liboensis, M. mangshanensis, M. maosonensis, M. medogensis, M. minor, M. nankiangensis, M. nanlingensis, M. omeimontis, M. oropedion, M. pachyproctus, M. parallela, M. popei, M. robusta, M. sangzhiensis, M. shapingensis, M. shuichengensis, M. spinata, M. vegrandis, M. wawuensis, M. zhangi, and M. zunhebotoensis (vs. tongue notched behind in the latter).
By toes with narrow lateral fringes, Megophrys anlongensis sp. nov. differs from M. angka, M. baolongensis, M. brachykolos, M. caobangensis, M. chishuiensis, M. damrei, M. daweimontis, M. dongguanensis, M. fansipanensis, M. feae, M. himalayana, M. hoanglienensis, M. huangshanensis, M. insularis, M. jiangi, M. jiulianensis, M. kalimantanensis, M. koui, M. lekaguli, M. lishuiensis, M. major, M. mangshanensis, M. medogensis, M. megacephala, M. microstoma, M. minor, M. nankunensis, M. obesa, M. ombrophila, M. oreocrypta, M. oropedion, M. pachyproctus, M. parva, M. periosa, M. shunhuangensis, M. takensis, M. tuberogranulata, M. wawuensis, M. wugongensis, M. wuliangshanensis, and M. xianjuensis (vs. lacking lateral fringes on toes in the latter), and differs from M. binchuanensis, M. boettgeri, M. carinense, M. cheni, M. chuannanensis, M. dringi, M. feii, M. gigantica, M. glandulosa, M. intermedia, M. jingdongensis, M. liboensis, M. lini, M. orientalis, M. palpebralespinosa, M. platyparietus, M. shapingensis, M. shuichengensis, M. spinata, and M. xiangnanensis (vs. with wide lateral fringes in the latter).
By toes with rudimentary webbing, Megophrys anlongensis sp. nov. differs from M. brachykolos, M. carinense, M. flavipunctata, M. jingdongensis, M. jinggangensis, M. lini, M. major, M. palpebralespinosa, M. popei, M. shuichengensis, and M. spinata (vs. at least one-fourth webbed in the latter).
By heels overlapping when thighs are positioned at right angles to the body, Megophrys anlongensis sp. nov. differs from M. acuta, M. brachykolos, M. dongguanensis, M. huangshanensis, M. kuatunensis, M. nankunensis, M. obesa, M. ombrophila, and M. wugongensis (vs. not meeting in the latter).
By tibiotarsal articulation reaching to the level of mid-eye when leg stretched forward, Megophrys anlongensis sp. nov. differs from M. daweimontis, M. glandulosa, M. lini, M. major, M. medogensis, M. obesa, and M. sangzhiensis (vs. reaching the anterior corner of the eye or beyond eye or nostril or tip of snout in the latter), differs from M. mufumontana (vs. reaching tympanum in males and to the eye in females in the latter), and differs from M. chishuiensis (vs. reaching the level between tympanum and eye in the latter).
By having an internal single subgular vocal sac in male, Megophrys anlongensis sp. nov. differs from M. caudoprocta, M. shapingensis, and M. shuichengensis (vs. vocal sac absent in the latter).
Megophrys anlongensis sp. nov. is genetically closest to M. binchuanensis. The new species could be identified from M. binchuanensis distinctly by having a bigger body size (SVL 40.0–45.5 mm in males and 48.9–51.2 in females in the new species vs. SVL 32.0–36.0 mm in males and 40.2–42.5 mm in females in the latter), having narrow lateral fringes on toes (vs. wide in the latter), and heels overlapping when thighs are positioned at right angles to the body (vs. just meeting in the latter).
Megophrys anlongensis sp. nov. is known only from the type locality, Anlong County, Guizhou Province, China at elevations between 1400–1600 m. The individuals were frequently found near the streams surrounded by evergreen broadleaved forests (Fig.
The specific name anlongensis refers to the known distribution of this species, Anlong County, Guizhou Province, China. We propose the common English name “Anlong horned toad”, and Chinese name “An Long Jiao Chan” (安龙角蟾).
Southwestern China was proposed as biodiversity hotspot (
We are grateful to editors and reviewers for their working on the manuscript. This work was supported by National Natural Sciences Foundation of China (Grant Nos.: 32070426 and 31960099), Basic research project of science and technology department of Guizhou Province (Nos. [2020] 1Y083), Science and technology support project of science and technology department of Guizhou Provincial (No. [2020] 4Y029) and Guizhou Provincial Department of Education Youth Science and Technology Talents Growth Project (Nos. KY[2018]455 and KY[2018]468).
Table S1. Uncorrected p-distances between the Megophrys species based on the 16S gene sequences.
Data type: statistical data
Table S2. Uncorrected p-distances between the Megophrys species based on the COI gene sequences
Data type: statistical data
Table S3. Diagnostic characters separating the new species described in this study from other species of Megophrys
Data type: species data