Research Article |
Corresponding author: Ying-Yong Wang ( wangyy@mail.sysu.edu.cn ) Academic editor: Annemarie Ohler
© 2020 Jian Wang, Zhi-Tong Lyu, Shuo Qi, Zhao-Chi Zeng, Wen-Xiang Zhang, Long-Shan Lu, 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:
Wang J, Lyu Z-T, Qi S, Zeng Z-C, Zhang W-X, Lu L-S, Wang Y-Y (2020) Two new Leptobrachella species (Anura, Megophryidae) from the Yunnan-Guizhou Plateau, southwestern China. ZooKeys 995: 97-125. https://doi.org/10.3897/zookeys.995.55939
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Two new toad species of the genus Leptobrachella are described from the Yunnan-Guizhou Plateau of China, based on the combination of molecular and morphological data. The description of Leptobrachella aspera Wang, Lyu, Qi & Wang, sp. nov. from Huanglianshan Nature Reserve represents the thirteenth Leptobrachella species known from Yunnan Province, and the description of Leptobrachella dorsospina Wang, Lyu, Qi & Wang, sp. nov. from Yushe Forest Park represents the sixth Leptobrachella species known from Guizhou Province. These new discoveries further emphasize the extremely high diversity of the Leptobrachella toads in these regions.
Leptobrachella aspera sp. nov., Leptobrachella dorsospina sp. nov., molecular phylogeny, morphology, taxonomy
The generic classifications within the family Megophryidae Bonaparte, 1850 have always been controversial. For example, recent comprehensive approaches have produced different taxonomic schemes for the genus Megophrys sensu lato Kuhl and Van Hasselt 1822 (
During recent field surveys in the Yunnan-Guizhou Plateau of southwestern China, a number of megophrid specimens were collected from Yushe Forest Park in western Guizhou (Fig.
Collection sites. Site 1-Yushe Forest Park, Shuicheng County, Guizhou Province, the type locality of Leptobrachella dorsospina sp. nov.; site 2-Huanglianshan Nature Reserve, Lyuchun County, Yunnan Province, the type locality of L. aspera sp. nov.; site 3-Xiaoqiaogou Nature Reserve, Xichou County, Yunnan Province, the type locality of L. feii; and site 4-Daweishan Nature Reserve, Pingbian County, Yunnan Province, another distribution locality of L. feii.
For the molecular analyses, a total of 80 sequences (nine muscle tissue samples was sequenced and 71 sequences obtained from GenBank) were used, including five sequences of the undescribed species from Guizhou, four sequences of the undescribed species from Yunnan, 69 sequences of 66 recognized congeners, and two out-group sequences of Oreolalax rhodostigmatus Hu & Fei, 1979 and Leptobrachium tengchongensis Yang & Huang, 2019, respectively (Table
DNA was extracted from muscle tissue using a DNA extraction kit from Tiangen Biotech (Beijing) Co., Ltd. The mitochondrial gene 16S ribosomal RNA gene (16S rRNA) fragment from each sample was sequenced. Fragments were amplified using the primer pairs L3975 (5'-CGCCTGTTTACCAAAAACAT-3') and H4551 (5'-CCGGTCTGAACTCAGATCACGT-3') (
Collection localities, voucher data and GenBank numbers (16S rRNA) for all samples used in this study.
ID | Ingroup | Collection Locality | Voucher No. | GenBank No. |
---|---|---|---|---|
1 | Leptobrachella aspera sp. nov. | Huanglianshan Nature Reserve, Lyuchun, Yunnan, China |
|
MW046199 |
2 | Leptobrachella aspera sp. nov. | Huanglianshan Nature Reserve, Lyuchun, Yunnan, China |
|
MW046200 |
3 | Leptobrachella aspera sp. nov. | Huanglianshan Nature Reserve, Lyuchun, Yunnan, China |
|
MW046201 |
4 | Leptobrachella aspera sp. nov. | Huanglianshan Nature Reserve, Lyuchun, Yunnan, China |
|
MW046202 |
5 | Leptobrachella dorsospina sp. nov. | Yushe Forest Park, Shuicheng, Guizhou, China |
|
MW046194 |
6 | Leptobrachella dorsospina sp. nov. | Yushe Forest Park, Shuicheng, Guizhou, China |
|
MW046195 |
7 | Leptobrachella dorsospina sp. nov. | Yushe Forest Park, Shuicheng, Guizhou, China |
|
MW046196 |
8 | Leptobrachella dorsospina sp. nov. | Yushe Forest Park, Shuicheng, Guizhou, China |
|
MW046197 |
9 | Leptobrachella dorsospina sp. nov. | Yushe Forest Park, Shuicheng, Guizhou, China |
|
MW046198 |
10 | Leptobrachella feii | Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ032625 | MT302635 |
11 | Leptobrachella feii | Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ048894 | MT302634 |
12 | Leptobrachella feii | Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ048972 | MT302636 |
13 | Leptobrachella feii | Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ048973 | MT302637 |
14 | Leptobrachella aerea | U Bo, Quang Binh, Vietnam | ZFMK 86362 | JN848409 |
15 | Leptobrachella alpina | Huangcaoling, Jingdong, Yunnan, China | KIZ046816 | MH055866 |
16 | Leptobrachella applebyi | Ngoc Linh, Kon Tum, Vietnam | AMS R 173778 | KR018108 |
17 | Leptobrachella arayai | Mesilau, Sabah, Malaysia | BORNEENSIS 22931 | AB847558 |
18 | Leptobrachella ardens | Kon Ka Kinh, Gia Lai, Vietnam | AMS R 176463 | KR018110 |
19 | Leptobrachella bidoupensis | Hon Giao, Lam Dong, Vietnam | NCSM 77321 | HQ902883 |
20 | Leptobrachella bijie | Zhaozishan Nature Reserve, Guizhou, China |
|
MK414539 |
21 | Leptobrachella botsfordi | Fansipan, Lao Cai, Vietnam | AMS R 176540 | MH055953 |
22 | Leptobrachella bourreti | Lao Cai, Vietnam | AMS R 177673 | KR018124 |
23 | Leptobrachella chishuiensis | Chishui, Guizhou Province, China | CIBCS20190518047 | MT117053 |
24 | Leptobrachella crocea | Kon Tum, Vietnam | AMS R 173740 | MH055954 |
25 | Leptobrachella dringi | Gunung Mulu National Park, Sarawak, Malaysia | NMBE1056372 | KJ831298 |
26 | Leptobrachella eos | Long Nai, Phongsaly, Laos | MNHN 2004.0274 | JN848452 |
27 | Leptobrachella firthi | Ngoc Linh Nature Reserve, Kon Tum, Vietnam | AMS R 176524 | JQ739206 |
28 | Leptobrachella flaviglandulosa | Xiaoqiaogou Nature Reserve, Yunnan, China | KIZ032626 | MT302633 |
29 | Leptobrachella fritinniens | Base Camp of Mulu NP, Sarawak, Malaysia | KUHE 55371 | AB847557 |
30 | Leptobrachella fuliginosa | Phetchaburi, Thailand | KUHE 20174 | LC201987 |
31 | Leptobrachella gracilis | Camp 1 of Gunung Mulu NP, Sarawak, Malaysia | NMBE1056364 | KJ831300 |
32 | Leptobrachella hamidi | Bukit Lanjak, Malaysia | KUHE 17545 | AB969286 |
33 | Leptobrachella heteropus | Larut, Malaysia | KUHE 15486 | LC202005 |
34 | Leptobrachella isos | Gia Lai, Vietnam | AMS R 176480 | KT824769 |
35 | Leptobrachella kajangensis | Tioman, Malaysia | LSUHC 4431 | LC202001 |
36 | Leptobrachella kalonensis | Song Luy, Binh Thuan, Vietnam | AMNH A191762 | KR018115 |
37 | Leptobrachella kecil | Cameron, Malaysia | KUHE 52440 | LC202004 |
38 | Leptobrachella khasiorum | Meghalaya, India | SDBDU 2009.329 | KY022303 |
39 | Leptobrachella laui | Tai Mo Shan, Hongkong, China |
|
KM014546 |
40 | Leptobrachella liui | Guadun, Mt. Wuyi, Fujian, China |
|
MH605574 |
41 | Leptobrachella macrops | Phu Yen, Vietnam | PYU DTD-508 | MG787991 |
42 | Leptobrachella maculosa | Phuoc Binh, Ninh Thuan, Vietnam | ZFMK 96600 | KR018120 |
43 | Leptobrachella mangshanensis | Mangshan Nature Reserve, Hunan, China | MSZTC201701 | MG132196 |
44 | Leptobrachella maoershanensis | Maoershan Nature Reserve, Guangxi, China | KIZ019385 | KY986930 |
45 | Leptobrachella marmorata | Annah Rais, Padawan, Malaysia | KUHE 53192 | AB969287 |
46 | Leptobrachella maura | Kinabalu, Malaysia | SP 21450 | AB847559 |
47 | Leptobrachella melanoleuca | Srat Thani, Thailand | KUHE 19719 | LC201990 |
48 | Leptobrachella melica | Virachey, Ratanakiri, Cambodia | MVZ 258197 | HM133599 |
49 | Leptobrachella minima | Changdao, Thailand | KUHE 23733 | LC201980 |
50 | Leptobrachella nahangensis | Na Hang Nature Reserve, Tuyen Quang, Vietnam | ROM 7035 | MH055853 |
51 | Leptobrachella namdongensis | Thanh Hoa, Vietnam | VNUF A.2017.95 | MK965390 |
52 | Leptobrachella niveimontis | Daxueshan Nature Reserve, Yunnan, China | KIZ015734 | MT302618 |
53 | Leptobrachella nyx | Malipo, Yunnan, China | ROM 35606 | MH055814 |
54 | Leptobrachella oshanensis | Mt. Emei, Sichuan, China |
|
KM014810 |
55 | Leptobrachella pallida | Gia Rich, Lam Dong, Vietnam | UNS00510 | KR018112 |
56 | Leptobrachella pelodytoides | Tam Dao, Vinh Phu, Vietnam | MVZ 223642 | AY236798 |
57 | Leptobrachella petrops | Tuyen Quang, Vietnam | VNMN:2016 A.06 | KY459998 |
58 | Leptobrachella picta | Gunung Kinabalu National Park, Sabah, Malaysia | UNIMAS 8705 | KJ831295 |
59 | Leptobrachella pluvialis | Sa Pa, Lao Cai, Vietnam | MNHN: 1999.5675 | JN848391 |
60 | Leptobrachella puhoatensis | Nghe An, Vietnam | AMS R184852 | KY849588 |
61 | Leptobrachella purpura | Yingjiang, Yunnan, China |
|
MG520355 |
62 | Leptobrachella purpuraventra | Wujing Nature Reserve, Guizhou, China |
|
MK414518 |
63 | Leptobrachella pyrrhops | Loc Bac, Lam Dong, Vietnam | ZMMU ABV-00176 | KP017576 |
64 | Leptobrachella rowleyae | Son Tra, Da Nang, Vietnam | ITBCZ 4113 | MG682549 |
65 | Leptobrachella sabahmontana | Mahua, Crocker, Malaysia | BORNEENSIS 12454 | AB847550 |
66 | Leptobrachella shangsiensis | Shiwandashan, Guangxi, China | NHMG1401032 | MK095460 |
67 | Leptobrachella sola | Terengganu, Malaysia | KUHE 52342 | LC202011 |
68 | Leptobrachella suiyangensis | Suiyang, Guizhou, China | GZNU20180606002 | MK829648 |
69 | Leptobrachella sungi | Bac Giang, Vietnam | ZMMU-NAP-02269 | MH055859 |
70 | Leptobrachella tadungensis | Dak Nong, Vietnam | UNS00517 | KR018122 |
71 | Leptobrachella tengchongensis | Tengchong, Yunnan, China |
|
KU589209 |
72 | Leptobrachella tuberosa | Kon Ka Kinh National Park, Gia Lai, Vietnam | ZMMU-NAP-02275 | MH055959 |
73 | Leptobrachella ventripunctata | Xishuangbanna, Yunnan, China |
|
KM014811 |
74 | Leptobrachella wuhuangmontis | Mt. Wuhuang, Pubei, Guangxi, China |
|
MH605577 |
75 | Leptobrachella wulingensis | Tianzishan Nature Reserve, Hunan, China | CSUFT 200 | MT530317 |
76 | Leptobrachella yingjiangensis | Yingjiang, Yunnan, China |
|
MG520350 |
77 | Leptobrachella yunkaiensis | Yunkaishan Nature Reserve, Guangdong, China |
|
MH605584 |
78 | Leptobrachella zhangyapingi | Chiang Mai, Thailand | KIZ07258 | MH055864 |
79 | Leptobrachium tengchongense | Tengchong, Yunnan, China |
|
KX066876 |
80 | Oreolalax rhodostigmatus | Da Fang, Guizhou, China |
|
EF397248 |
Sequences were aligned in Clustal X 2.0 (
Measurements followed
SVL snout-vent length (from tip of snout to vent);
HDL head length (from tip of snout to rear of jaws);
HDW head width (head width at commissure of jaws);
SNT snout length (from tip of snout to anterior corner of eye);
EYE eye diameter (diameter of exposed portion of eyeball);
IOD interorbital distance (minimum distance between upper eyelids);
IND internasal distance (distance between nares);
TMP tympanum diameter (horizontal diameter of tympanum);
TEY tympanum-eye distance (distance from anterior edge of tympanum to posterior corner of eye);
TIB tibia length (distance from knee to heel);
ML manus length (distance from tip of third digit to proximal edge of inner palmar tubercle);
PL pes length (distance from tip of fourth toe to proximal edge of the inner metatarsal tubercle);
LAHL length of lower arm and hand (distance from tip of the third finger to elbow);
HLL hindlimb length (distance from tip of fourth toe to vent).
Sex was determined by the presence of internal vocal sac openings, and the presence of eggs in abdomen seen via external inspection.
All specimens were fixed in 10% buffered formalin and later transferred to 70% ethanol for preservation, and deposited at the Museum of Biology, Sun Yat-sen University (
Comparative morphological data of Leptobrachella species were obtained from examination of museum specimens (see Appendix
Data sources of the 82 currently known species of the genus Leptobrachella.
ID | Leptobrachella species | Literature |
---|---|---|
1 | L. aerea (Rowley, Stuart, Richards, Phimmachak & Sivongxay, 2010) |
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2 | L. alpina (Fei, Ye & Li, 1990) |
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3 | L. applebyi (Rowley & Cao, 2009) |
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4 | L. arayai (Matsui, 1997) |
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5 | L. ardens (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
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6 | L. baluensis Smith, 1931 |
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7 | L. bijie Wang, Li, Li, Chen & Wang, 2019 |
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8 | L. bidoupensis (Rowley, Le, Tran & Hoang, 2011) |
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9 | L. bondangensis Eto, Matsui, Hamidy, Munir & Iskandar, 2018 |
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10 | L. botsfordi (Rowley, Dau & Nguyen, 2013) |
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11 | L. bourreti (Dubois, 1983) |
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12 | L. brevicrus Dring, 1983 |
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13 | L. crocea (Rowley, Hoang, Le, Dau & Cao, 2010) |
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14 | L. chishuiensis Li, Liu, Wei & Wang, 2020 |
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15 | L. dringi (Dubois, 1987) |
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16 | L. eos (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011) |
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17 | L. feii Chen, Yuan & Che, 2020 |
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18 | L. firthi (Rowley, Hoang, Dau, Le & Cao, 2012) |
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19 | L. fritinniens (Dehling & Matsui, 2013) |
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20 | L. fuliginosa (Matsui, 2006) |
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21 | L. flaviglandulosa Chen, Wang & Che, 2020 |
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22 | L. fusca Eto, Matsui, Hamidy, Munir & Iskandar, 2018 |
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23 | L. gracilis (Günther, 1872) |
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24 | L. hamidi (Matsui, 1997) |
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25 | L. heteropus (Boulenger, 1900) |
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26 | L. isos (Rowley, Stuart, Neang, Hoang, Dau, Nguyen & Emmett, 2015) |
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27 | L. itiokai Eto, Matsui & Nishikawa, 2016 |
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28 | L. juliandringi Eto, Matsui & Nishikawa, 2015 |
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29 | L. kajangensis (Grismer, Grismer & Youmans, 2004) |
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30 | L. kalonensis (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
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31 | L. kecil (Matsui, Belabut, Ahmad & Yong, 2009) |
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32 | L. khasiorum (Das, Tron, Rangad & Hooroo, 2010) |
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33 | L. lateralis (Anderson, 1871) |
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34 | L. laui (Sung, Yang & Wang, 2014) |
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35 | L. liui (Fei & Ye, 1990) |
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36 | L. macrops (Duong, Do, Ngo, Nguyen & Poyarkov, 2018) |
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37 | L. maculosa (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
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38 | L. mangshanensis (Hou, Zhang, Hu, Li, Shi, Chen, Mo & Wang, 2018) |
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39 | L. maoershanensis (Yuan, Sun, Chen, Rowley & Che, 2017) |
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40 | L. marmorata (Matsui, Zainudin & Nishikawa, 2014) |
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41 | L. maura (Inger, Lakim, Biun & Yambun, 1997) |
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42 | L. melanoleuca (Matsui, 2006) |
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43 | L. melica (Rowley, Stuart, Neang & Emmett, 2010) |
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44 | L. minima (Taylor, 1962) |
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45 | L. mjobergi Smith, 1925 |
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46 | L. nahangensis (Lathrop, Murphy, Orlov & Ho, 1998) |
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47 | L. natunae (Günther, 1895) | Günther 1895 |
48 | L. namdongensis Hoang, Nguyen, Luu, Nguyen & Jiang, 2019 |
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49 | L. neangi Stuart & Rowley, 2020 |
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50 | L. niveimontis Chen, Poyarkov, Yuan & Che, 2020 |
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51 | L. nokrekensis (Mathew & Sen, 2010) |
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52 | L. nyx (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011) |
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53 | L. oshanensis (Liu, 1950) |
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54 | L. pallida (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
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55 | L. palmata Inger & Stuebing, 1992 | Inger and Stuebing 1992 |
56 | L. parva Dring, 1983 |
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57 | L. pelodytoides (Boulenger, 1893) |
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58 | L. petrops (Rowley, Dau, Hoang, Le, Cutajar & Nguyen, 2017) |
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59 | L. pictua (Malkmus, 1992) |
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60 | L. platycephala (Dehling, 2012) |
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61 | L. pluvialis (Ohler, Marquis, Swan & Grosjean, 2000) |
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62 | L. puhoatensis (Rowley, Dau & Cao, 2017) |
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63 | L. purpura (Yang, Zeng & Wang, 2018) |
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64 | L. purpuraventra Wang, Li, Li, Chen & Wang, 2019 |
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65 | L. pyrrhops (Poyarkov, Rowley, Gogoleva, Vassilieva, Galoyan & Orlov, 2015) |
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66 | L. rowleyae (Nguyen, Poyarkov, Le, Vo, Ninh, Duong, Murphy & Sang, 2018) |
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67 | L. sabahmontana (Matsui, Nishikawa & Yambun, 2014) |
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68 | L. serasanae Dring, 1983 |
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69 | L. shangsiensis Chen, Liao, Zhou & Mo, 2019 |
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70 | L. sola (Matsui, 2006) |
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71 | L. suiyangensis Luo, Xiao, Gao & Zhou, 2020 |
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72 | L. sungi (Lathrop, Murphy, Orlov & Ho, 1998) |
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73 | L. tadungensis (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
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74 | L. tamdil (Sengupta, Sailo, Lalremsanga, Das & Das, 2010) |
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75 | L. tengchongensis (Yang, Wang, Chen & Rao, 2016) |
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76 | L. tuberosa (Inger, Orlov & Darevsky, 1999) |
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77 | L. ventripunctata (Fei, Ye & Li, 1990) |
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78 | L. wuhuangmontis Wang, Yang & Wang, 2018 |
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79 | L. wulingensis Qian, Xia, Cao, Xiao & Yang, 2020 | Qian et al. in publication |
80 | L. yingjiangensis (Yang, Zeng & Wang, 2018) |
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81 | L. yunkaiensis Wang, Li, Lyu & Wang, 2018 |
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82 | L. zhangyapingi (Jiang, Yan, Suwannapoom, Chomdej & Che, 2013) |
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The BI analyses are shown in Fig.
As shown by the phylogenetic result, Leptobrachella samples from Huanglianshan Nature Reserve are clustered in a distinct and robust monophyletic lineage with strong support (BPP 1.00). This lineage forms the sister taxon to L. feii occurring in Xiaoqiaogou Nature Reserve (BPP 1.00). The genetic distances between these two lineages are 3.0–3.4%, which is significantly larger than that among other recognized species (e.g., p-distance 2.6% between L. liui and L. mangshanensis). Detailed morphological examination also reveals a combination of characters that distinguish the specimens of the unnamed lineage from L. feii and other known congeners (see taxonomic comparison below). Therefore, based on the molecular and morphological differences, the population from Huanglianshan Nature Reserve is proposed as a new species, Leptobrachella aspera sp. nov.
Samples of the other unnamed lineage from Yushe Forest Park, cluster in another distinct and robust monophyletic lineage with strong support (BPP 1.00). This lineage is close to several species occurring in southwestern China, but its specific placement remains unresolved due to the insufficient support values. The smallest genetic distance between this lineage and another congener is 3.5% (vs. L. purpuraventra), which is significantly larger than that between other recognized species (e.g., p-distance 2.6% between L. liui and L. mangshanensis). Detailed morphological examination also reveals a combination of characteristics distinguishing the specimens of this lineage from all known congeners (see taxonomic comparison below). Therefore, based on the molecular and morphological differences, the population from Yushe Forest Park is proposed as a new species, Leptobrachella dorsospina sp. nov.
Comparisons of selected diagnostic characters for the new species described herein and congeners occurring north of the Kra Isthmus (modified from Rowley et al. 2017;
Leptobrachella species | Male SVL (mm) | Black spots on flanks | Toe webbing | Toe fringes | Ventral coloration | Dorsal skin texture |
---|---|---|---|---|---|---|
L. aspera sp. nov. | 22.4 | Present | Rudimentary | Narrow | Creamy white with distinct dark patches on chest and abdomen | Rough with dense conical granules, tubercles, and glandular folds |
L. dorsospina sp. nov. | 28.7–30.5 | Present | Rudimentary | Narrow | Greyish white with black spots and orange pigmentations | Rough with dense conical granules, tubercles, glandular folds, and conical spines |
L. aerea | 25.1–28.9 | Absent | Rudimentary | Wide | Near immaculate creamy white, brown speckles on margins | Finely tuberculate |
L. alpina | 24.0–26.4 | Present | Rudimentary | Wide in males | Creamy-white with dark spots | Relatively smooth, some with small warts |
L. applebyi | 19.6–22.3 | Present | Rudimentary | Absent | Reddish brown with white speckles | Smooth |
L. ardens | 21.3–24.7 | Present | Absent | Absent | Reddish brown with white speckles | Smooth, finely shagreened |
L. bidoupensis | 18.5–25.4 | Present | Rudimentary | Weak | Reddish brown with white speckles | Smooth |
L. bijie | 29.0–30.4 | Present | Rudimentary | Narrow | White with distinct nebulous greyish speckles on chest and ventrolateral flanks | Shagreened and granular |
L. botsfordi | 29.1–32.6 | Absent | Rudimentary | Narrow | Reddish brown with white speckles | Shagreened |
L. bourreti | 28.0–36.2 | Present | Rudimentary | Weak | Creamy white | Relatively smooth, some with small warts |
L. crocea | 22.2–27.3 | Absent | Rudimentary | Absent | Bright orange | Highly tuberculate |
L. chishuiensis | 30.8–33.4 | Present | Rudimentary | Narrow | White with distinct nebulous greyish speckles on chest and ventrolateral flanks | Shagreened and granular |
L. eos | 33.1–34.7 | Absent | Rudimentary | Wide | Creamy white | Shagreened |
L. feii | 21.5–22.8 | Present | Rudimentary | Narrow | Creamy white with black blotches | Shagreened with small tubercles and ridge |
L. firthi | 26.4–29.2 | Absent | Rudimentary | Wide in males | Creamy white | Shagreened with fine tubercles |
L. flaviglandulosa | 23.0–27.0 | Present | Poorly developed | Narrow | Whitish with black speckles on margins | Shagreened with yellowish- brown tubercles |
L. fuliginosa | 28.2–30.0 | Present | Rudimentary | Weak | White with brown dusting | Nearly smooth with few tubercles |
L. isos | 23.7–27.9 | Absent | Rudimentary | Wide in males | Creamy white with white dusting on margins | Mostly smooth, females more tuberculate |
L. kalonensis | 25.8–30.6 | Present | Absent | Absent | Pale, speckled brown | Smooth |
L. khasiorum | 24.5–27.3 | Present | Rudimentary | Wide | Creamy white | Isolated, scattered tubercles |
L. laui | 24.8–26.7 | Present | Rudimentary | Wide | Creamy white with dark brown dusting on margins | Round granular tubercles |
L. liui | 23.0–28.7 | Present | Rudimentary | Wide | Creamy white with dark brown spots on chest and margins | Round granular tubercles with glandular folds |
L. lateralis | 26.9–28.3 | Present | Rudimentary | Absent | Creamy white | Roughly granular |
L. macrops | 28.0–29.3 | Present | Rudimentary | Absent | Greyish violet with white speckles | Roughly granular with larger tubercles |
L. maculosa | 24.2–26.6 | Present | Absent | Absent | Brown with few white speckles | Mostly smooth |
L. mangshanensis | 22.2–27.8 | Present | Rudimentary | Weak | White speckles on throat and belly | Nearly smooth |
L. maoershanensis | 25.2–30.4 | Present | Rudimentary | Narrow | Creamy white chest and belly with irregular black spots | With longitudinal folds |
L. melica | 19.5–22.7 | Present | Rudimentary | Absent | Reddish brown with white speckles | Smooth |
L. minima | 25.7–31.4 | Present | Rudimentary | Absent | Creamy white | Smooth |
L. nahangensis | 40.8 | Present | Rudimentary | Absent | Creamy white with light speckles on throat and chest | Smooth |
L. niveimontis | 22.5–23.6 | Present | Rudimentary | Narrow | Marbling with black speckles | Relatively smooth with small tubercles |
L. nokrekensis | 26.0–33.0 | Present | Rudimentary | Unknown | Creamy white | Tubercles and longitudinal folds |
L. nyx | 26.7–32.6 | Present | Rudimentary | Absent | Creamy white with brown margins | Rounded tubercles |
L. namdongensis | 30.9 | Present | Rudimentary | Absent | Immaculate white, chest and belly with dark specking on outer margins | Low, round tubercles, more dense in posterior part of the back |
L. neangi | - | Present | Weak (in females) | Absent (in females) | Light purplish gray with dark brown mottling on throat |
Small, irregular bumps and ridges |
L. oshanensis | 26.6–30.7 | Present | Absent | Absent | Whitish with no markings or only small, light grey spots | Smooth with few glandular ridges |
L. pallida | 24.5–27.7 | Absent | Absent | Absent | Reddish brown with white speckles | Tuberculate |
L. pelodytoides | 27.5–32.3 | Present | Wide | Narrow | Whitish | Small, smooth warts |
L. petrops | 23.6–27.6 | Absent | Absent | Narrow | Immaculate creamy white | Highly tuberculate |
L. pluvialis | 21.3–22.3 | Present | Rudimentary | Absent | Dirty white with dark brown marbling | Smooth, flattened tubercles on flanks |
L. puhoatensis | 24.2–28.1 | Present | Rudimentary | Narrow | Reddish brown with white dusting | With longitudinal skin ridges |
L. purpura | 25.0–27.5 | Present | Rudimentary | Wide | Dull white with indistinct grey dusting | Shagreen with small tubercles |
L. purpuraventra | 27.3–29.8 | Present | Rudimentary | Narrow | Grey purple with distinct nebulous greyish speckles on chest and ventrolateral flanks | Shagreened with granules |
L. pyrrhops | 30.8–34.3 | Present | Rudimentary | Absent | Reddish brown with white speckles | Slightly shagreened |
L. rowleyae | 23.4–25.4 | Present | Absent | Absent | Pinkish milk-white to light brown chest and belly with numerous white speckles | Smooth with numerous tiny tubercles |
L. suiyangensis | 28.7–29.7 | Present | Rudimentary | Narrow | Yellowish creamy-white with marble texture chest and belly or with irregular light brown speckles | Shagreen with small granules |
L. sungi | 48.3–52.7 | Absent or small | Wide | Weak | White | Granular |
L. tadungensis | 23.3–28.2 | Present | Absent | Absent | Reddish brown with white speckles | Smooth |
L. tamdil | 32.3 | Present | Wide | Wide | White | Weakly tuberculate |
L. tengchongensis | 23.9–26.0 | Present | Rudimentary | Narrow | White with dark brown blotches | Shagreened with small tubercles |
L. tuberosa | 24.4–29.5 | Absent | Rudimentary | Absent | White with small grey spots/streaks | Highly tuberculate |
L. ventripunctata | 25.5–28.0 | Present | Rudimentary | Absent | Chest and belly with dark brown spots | Longitudinal skin ridges |
L. wuhuangmontis | 25.6–30.0 | Present | Rudimentary | Narrow | Greyish white mixed by tiny white and black dots | Rough, scattered with dense conical tubercles |
L. wulingensis | 22.7–30.5 | Present | Rudimentary | Narrow | Translucent creamy white, with distinct or indistinct brown speckles at margins | Shagreened with sparse large warts, some with longitudinal ridges |
L. yingjiangensis | 25.7–27.6 | Present | Rudimentary | Wide | Creamy white with dark brown flecks on chest and margins | Shagreened with small tubercles |
L. yunkaiensis | 25.9–29.3 | Present | Rudimentary | Wide | Belly pink with distinct or indistinct speckles | Shagreened with short skin ridges and raised warts |
L. zhangyapingi | 45.8–52.5 | Absent | Rudimentary | Wide | Creamy-white with brown margins | Mostly smooth with distinct tubercles |
Holotype.
Paratypes
(N = 3). Three adult females,
(1) Small size (SVL 22.4 mm in a single adult male, 25.0–26.4 in three adult females), (2) dorsal skin rough, with dense conical granules, tubercles and glandular folds, (3) iris bicolored, amber on upper half and silver on lower half, (4) tympanum distinctly discernible, distinct black supratympanic line present, (5) absence of webbing and lateral fringes on fingers, toes with rudimentary webbing and narrow lateral fringes both in males and females, (6) longitudinal ridges under toes not interrupted at the articulations, (7) relative finger lengths I < IV < II < III, relative toe length I < II < V < III < IV, (8) heels just meeting, tibia-tarsal articulation reaches the region between middle of eye to anterior corner of eye, (9) dorsum greyish brown to yellowish brown grounding, with small light orange granules and distinct darker brown markings scattered with irregular light orange or greyish white pigmentations, (10) flanks with several enlarged dark patches with light yellowish green margin, (11) ventral surface creamy white, with distinct regular dark patches on chest and abdomen.
From the 26 known congeners of the genus Leptobrachella occurring south of the Kra Isthmus, the presence of supra-axillary and ventrolateral glands, can easily distinguish L. aspera sp. nov. from L. arayai, L. dringi, L. fritinniens, L. gracilis, L. hamidi, L. heteropus, L. kajangensis, L. kecil, L. marmorata, L. melanoleuca, L. maura, L. picta, L. platycephala, L. sabahmontana and L. sola, all of which lack the supra-axillary and ventrolateral glands; and by the significantly larger body size, SVL 22.4 mm in a single male, L. aspera sp. nov. differs from the smaller L. baluensis (14.9–15.9 mm in males), L. brevicrus (17.1–17.8 mm in males), L. bondangensis (17.8 mm in male), L. fusca (16.3 mm in male), L. itiokai (15.2–16.7 mm in males), L. juliandringi (17.0–17.2 mm in males), L. mjobergi (15.7–19.0 mm in males), L. natunae (17.6 mm in one adult male), L. parva (15.0–16.9 mm in males), L. palmata (14.4–16.8 mm in males), and L. serasanae (16.9 mm in female).
Leptobrachella aspera sp. nov. is recovered as a sister taxon to L. feii in the phylogenetic tree (Fig.
While Leptobrachella pluvialis is distributed in the same mountain range on the Vietnamese side and possesses similar body size (
For the remaining 54 members of the genus Leptobrachella, in having SVL 22.4 mm in a single male, L. aspera sp. nov. differs from the larger L. aerea (25.1–28.9 in males), L. alpina (24.0–28.9 mm in males), L. bijie (29.0–30.4 mm in males), L. botsfordi (29.1–32.6 mm in males), L. bourreti (28.0–36.2 mm in males), L. chishuiensis (30.8–33.4 in males), L. eos (33.1–34.7 mm in males), L. firthi (26.4–29.2 mm in males), L. flaviglandulosa (23.0–27.0 mm in males), L. fuliginosa (28.2–30.0 mm in males), L. isos (23.7–27.9 mm in males), L. kalonensis (25.8–30.6 mm in males), L. khasiorum (24.5–27.3 mm in males), L. laui (24.8–26.7 mm in males), L. lateralis (26.9–28.3 mm in males), L. macrops (28.0–29.3 mm in males), L. maculosa (24.2–26.6 mm in males), L. minima (25.7–31.4 mm in males), L. nahangensis (40.8 mm in male), L. nokrekensis (26.0–33.0 mm in males), L. nyx (26.7–32.6 mm in males), L. neangi (30.9 mm in male), L. namdongensis (30.9 mm in male), L. oshanensis (26.6–30.7 mm in males), L. pallida (24.5–27.7 mm in males), L. pelodytoides (27.5–32.3 mm in males), L. petrops (23.6–27.6 mm in males), L. puhoatensis (24.2–28.1 mm in males), L. purpura (25.0–27.5 mm in males), L. purpuraventra (27.3–29.8 mm in males), L. pyrrhops (30.8–34.3 mm in males), L. rowleyae (23.4–25.4 mm in males), L. suiyangensis (28.7–29.7 mm in males), L. sungi (48.3–52.7 mm in males), L. tadungensis (23.3–28.2 mm in males), L. tamdil (32.3 mm in male), L. tengchongensis (23.9–26.0 mm in males), L. tuberosa (24.4–29.5 mm in males), L. ventripunctata (25.5–28.0 mm in males), L. wuhuangmontis (25.6–30.0 mm in males), L. yingjiangensis (25.7–27.6 mm in males), L. yunkaiensis (25.9–29.3 mm in males), and L. zhangyapingi (45.8–52.5 mm in males). By presence of black spots on flanks, the new species can be distinguished from L. crocea, versus absence of black spots on flanks; by rudimentary webbing on toes, the new species can be distinguished from L. ardens, versus absence of webbing on toes; by narrow lateral fringes on toes, the new species can be distinguished from L. applebyi, L. ardens, L. crocea, and L. melica, all having no lateral fringes on toes, and from L. liui, having wide lateral fringes on toes; by the creamy white ventral coloration and distinct regular dark patches on the chest and abdomen, the new species can be distinguished from L. applebyi, L. ardens, L. bidoupensis, and L. melica, all having reddish brown ventral coloration with white specks, from L. crocea, having bright orange ventral coloration, from L. mangshanensis, lacking dark skin patches on the chest and abdomen, from L. liui, having creamy white ventral coloration with dark brown spots on the chest and margins, and from L. niveimontis, having marbling ventral coloration with black speckling; by rough dorsal skin with dense conical granules, tubercles and glandular folds, the new species can be distinguished from L. applebyi, L. ardens, L. bidoupensis, L. mangshanensis, L. melica, and L. niveimontis, all having smooth dorsal skin.
Adult male. Body size small, SVL 22.4 mm. Head length slightly larger than head width, HDW/HDL 0.99; snout slightly protruding, projecting slightly beyond margin of the lower jaw; nostril closer to snout than eye; canthus rostralis gently rounded; loreal region slightly concave; interorbital space flat, internarial distance greater than interorbital distance, IND/IOD 1.07; pineal ocellus absent; pupil vertical; snout length longer than eye diameter, SNT/EYE 1.26; tympanum distinct, rounded, and slightly concave, diameter smaller than that of the eye and larger than tympanum-eye distance, TMP/EYE 0.52 and TEY/TMP 0.44; upper margin of tympanum in contact with supratympanic ridge; distinct black supratympanic line present; vomerine teeth absent; vocal sac openings slit-like, paired, located posterolaterally on floor of mouth in close proximity to the margins of the mandible; tongue deeply notched posteriorly; supratympanic ridge distinct, extending from posterior corner of eye to supra-axillary gland.
Tips of fingers rounded, slightly swollen; relative finger lengths I < IV < II < III; nuptial pad absent; subarticular tubercles absent; large, rounded inner palmar tubercle distinctly separated from small, rounded outer palmar tubercle; webbing and lateral fringes on fingers absent. Tips of toes rounded, slightly swollen; relative toe length I < II < V < III < IV; subarticular tubercles absent; distinct longitudinal dermal ridges present under the 3rd to 5th toes, not interrupted; large, oval inner metatarsal tubercle present, outer metatarsal tubercle absent; toes webbing rudimentary; narrow lateral fringes present on all toes. Tibia 47% of snout-vent length; tibiotarsal articulation reaching to anterior corner of eye; heels slightly overlapping when thighs are appressed at right angles with respect to body.
Dorsal skin rough, with dense conical granules, tubercles and glandular folds; ventral skin smooth; sparse tiny tubercles present on surface of chest; pectoral gland and femoral gland oval; the size of pectoral glands almost equal to tips of fingers and femoral glands; femoral gland situated on posteroventral surface of thigh, closer to knee than to vent; supra-axillary glands raised. Ventrolateral glands distinctly visible, raised, forming an incomplete line.
Dorsum greyish brown with small light orange granules, distinct darker brown markings scattered with irregular light orange and greyish white pigmentations. A dark brown inverted triangular pattern between the anterior corners of the eyes in connection with a dark brown W-shaped marking in the interorbital region, which is also connected to a W-shaped marking between the axillae. Tympanum dark brown. Small light orange granules present on dorsum of body and limbs; a dark brown blotch under the eye; transverse dark brown bars present on dorsal surface of limbs and digits; distinct dark brown patches with light yellowish green margin on flanks from groin to axilla; elbow and upper arms with distinct coppery orange coloration.
Ventral surface of throat, chest, and belly creamy white; presence of distinct nebulous greyish speckles present on throat, and distinct dark patches on chest and abdomen; ventral surface of limbs greyish purple, scattered with greyish white spots and small patches. Supra-axillary gland coppery orange; femoral, pectoral, and ventrolateral glands greyish white. Iris bicolored, amber on upper half and silver on lower half.
(Fig.
Measurements and body proportions are listed in Table
Voucher |
|
|
|
|
---|---|---|---|---|
Sex | Male | Female | Female | Female |
SVL | 22.4 | 25.3 | 25.0 | 26.4 |
HDL | 8.1 | 9.5 | 9.5 | 9.6 |
HDW | 8.0 | 9.3 | 9.2 | 9.0 |
SNT | 3.7 | 3.8 | 3.8 | 3.4 |
IND | 2.5 | 2.3 | 2.7 | 2.7 |
IOD | 2.3 | 2.5 | 2.5 | 2.5 |
EYE | 2.9 | 3.2 | 3.2 | 3.1 |
TMP | 1.5 | 1.8 | 1.9 | 1.6 |
TEY | 0.7 | 1.0 | 1.0 | 0.8 |
ML | 5.9 | 7.0 | 6.6 | 6.3 |
LAHL | 11.2 | 13.5 | 12.7 | 12.6 |
PL | 10.1 | 11.7 | 10.2 | 11.1 |
TIB | 10.6 | 12.4 | 11.9 | 11.9 |
HLL | 34.4 | 41.5 | 40.4 | 39.1 |
HDL/SVL | 0.36 | 0.37 | 0.38 | 0.36 |
HDW/SVL | 0.36 | 0.37 | 0.37 | 0.34 |
HDW/HDL | 0.99 | 0.98 | 0.97 | 0.94 |
SNT/HDL | 0.16 | 0.15 | 0.15 | 0.13 |
IND/HDW | 0.31 | 0.25 | 0.29 | 0.30 |
IOD/HDW | 0.29 | 0.27 | 0.27 | 0.28 |
IND/IOD | 1.07 | 0.91 | 1.08 | 1.09 |
EYE/HDL | 0.36 | 0.34 | 0.34 | 0.32 |
TMP/EYE | 0.52 | 0.56 | 0.60 | 0.51 |
ML/SVL | 0.26 | 0.28 | 0.26 | 0.24 |
LAHL/SVL | 0.50 | 0.53 | 0.51 | 0.48 |
PL/SVL | 0.45 | 0.46 | 0.41 | 0.42 |
TIB/SVL | 0.47 | 0.49 | 0.48 | 0.45 |
HLL/SVL | 1.53 | 1.64 | 1.61 | 1.48 |
The specific epithet, aspera, is a Latin adjective which means rough, in reference to the dorsal skin texture of the new species. According to its type locality, we suggest its English common name as “Huanglianshan Leaf Litter Toad”, and the Chinese name “Huang Lian Shan Zhang Tu Chan (黄连山掌突蟾)”.
Currently, Leptobrachella aspera sp. nov. is known only from its type locality Huanglianshan Nature Reserve, near the border between China and Vietnam. The new species was found along a drainage ditch of a mountainous road. The road was surrounded by broad-leaved forest at an altitude ca. 1930 m and not close to any hillstreams. Males were not heard calling during the field survey from 31 May to 1 June 2019.
Holotype.
Paratypes
(N = 6). An adult male,
(1) Small size (SVL 28.7–30.5 mm in two adult males, 32.1–39.8 mm in five adult females), (2) dorsal skin rough, with dense conical granules, tubercles, glandular folds and conical spines, (3) iris bicolored, light orange on upper half and silver on lower half, (4) tympanum distinctly discernible, distinct black supratympanic line present, (5) absence of webbing and lateral fringes on fingers, toes with rudimentary webbing and narrow lateral fringes both in males and females, (6) longitudinal ridges under toes interrupted at the articulations, (7) relative finger lengths II = IV < I < III, relative toe length I < II < V < III < IV, (8) heels slightly overlapping, tibia-tarsal articulation reaches forward to the posterior corners of eyes, (9) dorsum greyish brown to dark brown grounding, with distinct darker brown markings and scattered with irregular light greyish brown pigmentations and yellowish brown spots, (10) flanks with several enlarged dark patches positioned longitudinally in two rows, (11) ventral surface greyish white with black spots and orange pigmentations.
Compared with the 26 known congeners of the genus Leptobrachella occurring south of the Kra Isthmus, L. dorsospina sp. nov. can be easily distinguished by the presence of supra-axillary and ventrolateral glands, from L. arayai, L. dringi, L. fritinniens, L. gracilis, L. hamidi, L. heteropus, L. kajangensis, L. kecil, L. marmorata, L. melanoleuca, L. maura, L. picta, L. platycephala, L. sabahmontana and L. sola, all of which are lacking the supra-axillary and ventrolateral glands; and by the significantly larger body size, SVL 28.7–30.5 mm in two adult male, L. dorsospina sp. nov. differs from the smaller L. baluensis (14.9–15.9 mm in males), L. brevicrus (17.1–17.8 mm in males), L. bondangensis (17.8 mm in male), L. fusca (16.3 mm in male), L. itiokai (15.2–16.7 mm in males), L. juliandringi (17.0–17.2 mm in males), L. mjobergi (15.7–19.0 mm in males), L. natunae (17.6 mm in one adult male), L. parva (15.0–16.9 mm in males), L. palmata (14.4–16.8 mm in males), and L. serasanae (16.9 mm in female).
Leptobrachella dorsospina sp. nov. can be easily distinguished from Leptobrachella aspera sp. nov. by having distinctly larger body size, SVL 28.7–30.5 mm in males, 32.1–39.8 mm in females (vs. SVL 22.4 mm in male, 25.0–26.4 in females); conical spines on dorsal skin present (vs. absent); black spots on flanks in one row (vs. black spots on flanks in two rows); ventral skin greyish white with black spots and orange pigmentations (vs. ventral skin creamy white with distinct dark patches on chest and abdomen); longitudinal ridges under toes interrupted at the articulations (longitudinal ridges under toes not interrupted at the articulations).
For the remaining 56 members of the genus Leptobrachella, in having SVL 28.7–30.5 mm in two males, L. dorsospina sp. nov. differs from the larger L. eos (33.1–34.7 mm in males), L. nahangensis (40.8 mm in male), L. sungi (48.3–52.7 mm in males), L. tamdil (32.3 mm in male), and L. zhangyapingi (45.8–52.5 mm in males); and from the smaller L. alpina (24.0–26.4 mm in males), L. applebyi (19.6–22.3 mm in males), L. ardens (21.3–24.7 mm in males), L. bidoupensis (18.5–25.4 mm in males), L. crocea (22.2–27.3 mm in males), L. feii (21.5–22.8 mm in males), L. flaviglandulosa (23.0–27.0 mm in males), L. isos (23.7–27.9 mm in males), L. khasiorum (24.5–27.3 mm in males), L. laui (24.8–26.7 mm in males), L. maculosa (24.2–26.6 mm in males), L. mangshanensis (22.2–27.8 mm in males), L. melica (19.5–22.7 mm in males), L. niveimontis (22.5–23.6 mm in males), L. pallida (24.5–27.7 mm in males), L. petrops (23.6–27.6 mm in males), L. pluvialis (21.3–22.3 mm in males), L. puhoatensis (24.2–28.1 mm in males), L. purpura (25.0–27.5 mm in males), L. rowleyae (23.4–25.4 mm in males), L. tadungensis (23.3–28.2 mm in males), L. tengchongensis (23.9–26.0 mm in males), L. ventripunctata (25.5–28.0 mm in males), and L. yingjiangensis (25.7–27.6 mm in males). By having black spots on the flanks, L. dorsospina sp. nov. can be distinguished from L. aerea, L. botsfordi, L. firthi, and L. tuberosa, all of which lack black spots on the flanks. By having rough dorsal skin with conical spines, the new species can be distinguished from L. bijie, L. chishuiensis, L. liui, L. maoershanensis, L. pyrrhops, L. purpuraventra, L. suiyangensis, L. wuhuangmontis, L. wulingensis, and L. yunkaiensis (dorsal skin lacking spines); and from L. bourreti, L. fuliginosa, L. kalonensis, L. minima, L. oshanensis, and L. pelodytoides (dorsal skin smooth). By having narrow lateral fringes on the toes, the new species can be distinguished from L. lateralis, L. macrops, L. nyx, L. pyrrhops, L. namdongensis and L. neangi, all of which lack lateral fringes on the toes. The new species can be separated from the remaining L. nokrekensis by having greyish white ventral coloration with black patches and orange pigmentations (vs. creamy white), and having dense short glandular folds on the dorsal surface (vs. only a few glandular folds on the dorsal surface).
Adult male. Body size rather small, SVL 30.5 mm. Head length slightly larger than head width, HDW/HDL 0.99; snout slightly protruding, projecting slightly beyond margin of the lower jaw; nostril closer to snout than eye; canthus rostralis gently rounded; loreal region slightly concave; interorbital space flat, internarial distance smaller than interorbital distance, IND/IOD 0.91; pineal ocellus absent; vertical pupil; snout length larger than eye diameter, SNT/EYE 1.29; tympanum distinct, rounded, and slightly concave, diameter smaller than that of the eye and larger than tympanum-eye distance, TMP/EYE 0.43 and TEY/TMP 0.50; upper margin of tympanum in contact with supratympanic ridge; distinct black supratympanic line present; vomerine teeth absent; vocal sac openings slit-like, paired, located posterolaterally on floor of mouth in close proximity to the margins of the mandible; tongue deeply notched posteriorly; supratympanic ridge distinct, extending from posterior corner of eye to supra-axillary gland.
Tips of fingers rounded, slightly swollen; relative finger lengths II = IV < I < III; nuptial pad absent; subarticular tubercles absent; large, rounded inner palmar tubercle distinctly separated from small, rounded outer palmar tubercle; absence of webbing and lateral fringes on fingers. Tips of toes rounded, slightly swollen; relative toe length I < II < V < III < IV; subarticular tubercles absent; distinct longitudinal dermal ridges present under the 3rd to 5th toes, interrupted; large, oval inner metatarsal tubercle present, outer metatarsal tubercle absent; toes webbing rudimentary; narrow lateral fringes present on all toes. Tibia 44% of snout-vent length; tibiotarsal articulation reaches to posterior corner of eye; heels slightly overlapping when thighs are appressed at right angles with respect to body.
Dorsal skin rough, with dense conical granules, tubercles, glandular folds and conical spines; ventral skin smooth; pectoral gland and femoral gland oval; the size of pectoral glands almost equal to tips of fingers and femoral glands; femoral gland situated on posteroventral surface of thigh, closer to knee than to vent; supra-axillary glands raised. Ventrolateral glands distinctly visible, raised, forming an incomplete line.
Dorsum greyish brown with distinct darker brown markings on sides and scattered with irregular light greyish brown pigmentations and yellowish brown spots. An indistinct, darker brown inverted triangular pattern between anterior corners of the eyes, connected to an indistinct dark brown W-shaped marking between the axillae. Dense translucent spines present on dorsal skin of body and limbs. Upper 2/3 of the tympanum dark brown, lower 1/3 light orange, scattered with tiny coppery orange spots. Small greyish white and light brown granules present on the dorsum of the body and limbs; a dark brown vertical bar under the eye; transverse dark brown bars on the dorsal surface of the limbs and digits; distinct dark brown patches on the flanks, from groin to axilla; elbow and upper arms with distinct light orange coloration.
Ventral surface of throat, chest, and belly greyish white; throat with light brown speckles, chest, and abdomen with distinct dark patches; ventral surface of limbs dark grey, scattered with greyish white spots and small patches. Supra-axillary gland light orange; femoral, pectoral, and ventrolateral glands greyish white. Iris bicolored, light orange on upper half and silver on lower half.
(Fig.
Measurements and body proportions are listed in Table
Measurements, and body proportions of Leptobrachella dorsospina sp. nov.
Voucher |
|
|
|
|
|
|
|
---|---|---|---|---|---|---|---|
Sex | Male | Male | Female | Female | Female | Female | Female |
SVL | 28.7 | 30.5 | 36.1 | 37.3 | 39.8 | 32.1 | 33.8 |
HDL | 10.3 | 10.8 | 12.8 | 12.4 | 12.9 | 11.3 | 12.0 |
HDW | 10.6 | 10.7 | 11.9 | 12.9 | 13.2 | 11.8 | 12.2 |
SNT | 4.5 | 4.2 | 5.1 | 5.4 | 5.8 | 5.0 | 4.8 |
IND | 3.1 | 3.2 | 3.6 | 3.9 | 4.0 | 3.7 | 3.4 |
IOD | 3.4 | 2.9 | 3.5 | 3.4 | 3.3 | 3.0 | 2.9 |
EYE | 3.5 | 3.7 | 3.9 | 3.7 | 4.3 | 4.2 | 3.8 |
TMP | 1.7 | 1.6 | 2.3 | 2.3 | 2.6 | 2.1 | 2.1 |
TEY | 1.1 | 0.8 | 1.3 | 1.4 | 1.5 | 1.2 | 1.1 |
ML | 7.4 | 7.3 | 8.8 | 7.7 | 9.1 | 7.8 | 7.6 |
LAHL | 14.1 | 14.2 | 17.1 | 16.8 | 17.5 | 16.2 | 15.9 |
PL | 12.1 | 12.8 | 14.9 | 14.5 | 15.5 | 13.9 | 13.6 |
TIB | 13.5 | 13.4 | 15.5 | 16.3 | 16.6 | 14.9 | 14.5 |
HLL | 41.7 | 42.7 | 49.1 | 49.9 | 52.9 | 46.8 | 48.0 |
HDL/SVL | 0.36 | 0.35 | 0.35 | 0.33 | 0.32 | 0.35 | 0.36 |
HDW/SVL | 0.37 | 0.35 | 0.33 | 0.35 | 0.33 | 0.37 | 0.36 |
HDW/HDL | 1.03 | 0.99 | 0.93 | 1.04 | 1.02 | 1.04 | 1.02 |
SNT/HDL | 0.44 | 0.39 | 0.40 | 0.44 | 0.45 | 0.44 | 0.40 |
IND/HDW | 0.29 | 0.30 | 0.30 | 0.30 | 0.30 | 0.31 | 0.28 |
IOD/HDW | 0.32 | 0.27 | 0.29 | 0.26 | 0.25 | 0.25 | 0.24 |
EYE/HDL | 0.34 | 0.34 | 0.30 | 0.30 | 0.33 | 0.37 | 0.32 |
TMP/EYE | 0.49 | 0.43 | 0.59 | 0.62 | 0.60 | 0.50 | 0.55 |
ML/SVL | 0.26 | 0.24 | 0.24 | 0.21 | 0.23 | 0.24 | 0.22 |
LAHL/SVL | 0.49 | 0.47 | 0.47 | 0.45 | 0.44 | 0.50 | 0.47 |
PL/SVL | 0.42 | 0.42 | 0.41 | 0.39 | 0.39 | 0.43 | 0.40 |
TIB/SVL | 0.47 | 0.44 | 0.43 | 0.44 | 0.42 | 0.46 | 0.43 |
HLL/SVL | 1.45 | 1.40 | 1.36 | 1.34 | 1.33 | 1.46 | 1.42 |
The specific epithet, dorsospina, is in reference to the conical spines on the dorsal surface of body in the new species. According to its type locality, we suggest its English common name as “Shuicheng Leaf Litter Toad”, and the Chinese name “Shui Cheng Zhang Tu Chan (水城掌突蟾)”.
Currently, Leptobrachella dorsospina sp. nov. is known only from its type locality, Yushe Forest Park, which is near the border between Guizhou and Yunnan. The new species was found on the surface of fallen leaves by the clear-water rocky hill-stream in well-preserved montane evergreen broadleaf forest (ca. 2100 m a.s.l.). Males were not heard calling.
In the phylogenetic tree, the Leptobrachella pelodytoides (voucher number: MVZ 223642) sample from Tam Dao, northern Vietnam is clustered together with the topotypic L. ventripunctata (voucher number:
Yunnan and Guizhou are both largely within the species-rich Dian freshwater zoogeographical dominion (
We are grateful to Prof. Tao Thien Nguyen, Prof. Annemarie Ohler, and an anonymous reviewer for providing constructive comments that greatly improved the quality of the manuscript. We would like to thank Yu-Long Li, Run-Lin Li and Hong-Hui Chen for their help in the field work, and Yao Li for her help in the lab work.
Specimens examined
Leptobrachella alpina (n = 6): China: Yunnan Province: Jingdong County: Mt. Wuliang:
Leptobrachella bijie (n = 8): China: Guizhou: Bijie City:
Leptobrachella laui (n = 26): China: Hong Kong:
Leptobrachella liui (n = 18): China: Fujian Province: Mt. Wuyi:
Leptobrachella mangshanensis (n = 5): China: Guangdong: Nanling Nature Reserve:
Leptobrachella purpuraventra (n = 15): China: Guizhou: Bijie City:
Leptobrachella tengchongensis (n = 6): China: Yunnan Province: Baoshan City: Mt. Gaoligong:
Leptobrachella wuhuangmontis (n = 12): China: Guangxi Province: Pubei County: Mt. Wuhuang:
Leptobrachella yunkaiensis (n = 8): China: Guangdong Province: Maoming City: Dawuling Forest Station:
Table S1. Pairwise distances based on 16S gene among all sample used in this study
Data type: phylogenetic
Explanation note: Genetic distances among all Leptobrachella samples.