Research Article |
Corresponding author: Bin Wang ( wangbin@cib.ac.cn ) Academic editor: Angelica Crottini
© 2023 Jing Liu, Shengchao Shi, Shize Li, Mengfei Zhang, Sunjun Xiang, Gang Wei, 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:
Liu J, Shi S, Li S, Zhang M, Xiang S, Wei G, Wang B (2023) A new Asian leaf litter toad of the genus Leptobrachella (Amphibia, Anura, Megophryidae) from central south China. ZooKeys 1149: 103-134. https://doi.org/10.3897/zookeys.1149.85895
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A new species of the Asian leaf litter toad genus Leptobrachella from central south China is described. Molecular phylogenetic analyses, based on mitochondrial 16S rRNA and nuclear RAG1 gene sequences indicated the new species as an independent clade in the genus. The new species could be distinguished from its congeners by a combination of the following characters: body of medium size (SVL 29.2–34.2 mm in 15 adult males and 34.4–43.1 mm in seven adult females); distinct black spots present on flanks; toes rudimentary webbed, with wide lateral fringes; ventral belly white with distinct nebulous brown speckling on ventrolateral flanks; skin on dorsum shagreened with fine tiny granules or short ridges; iris copper above, silver below; heels overlapped when thighs are positioned at right angles to the body; tibia-tarsal articulation reaches the middle eye; dorsal surface of tadpole semi-transparent light brown, spots on tail absent, keratodont row formula I: 3+3/2+2: I; call series basically consist of repeated long calls, at dominant frequency (5093 ± 412 Hz).
China, molecular phylogenetic analyses, morphology, new species, taxonomy
The Asian leaf litter toads of the genus Leptobrachella Smith, 1925 (Anura, Megophryidae) are widely distributed from southern China west to north-eastern India and Myanmar, through mainland Indochina to peninsular Malaysia and the island of Borneo (
In recent years, we carried out a series of biodiversity surveys in Hunan and Guizhou Provinces, China and collected some specimens of Leptobrachella. Morphological comparisons, molecular phylogenetic analyses and bioacoustic comparisons consistently indicated these specimens as an undescribed species. We describe it herein as a new species.
A total of 22 specimens of the new species (Suppl. material
A total of 15 samples of the new species were used for molecular analyses (Table
Information for samples used in molecular phylogenetic analyses in this study.
ID | Species | Locality | Voucher number | GenBank accession number | |
---|---|---|---|---|---|
16S | RAG1 | ||||
1 | Leptobrachella dong sp. nov. | Tongdao County, Hunan Province, China |
|
OP764529 | / |
2 | Leptobrachella dong sp. nov. | Tongdao County, Hunan Province, China |
|
OP764530 | / |
3 | Leptobrachella dong sp. nov. | Tongdao County, Hunan Province, China |
|
OP764531 | OP776448 |
4 | Leptobrachella dong sp. nov. | Tongdao County, Hunan Province, China |
|
OP764532 | / |
5 | Leptobrachella dong sp. nov. | Congjiang County, Guizhou Province, China |
|
OP764533 | / |
6 | Leptobrachella dong sp. nov. | Congjiang County, Guizhou Province, China |
|
OP764534 | OP776439 |
7 | Leptobrachella dong sp. nov. | Congjiang County, Guizhou Province, China |
|
OP764535 | OP776441 |
8 | Leptobrachella dong sp. nov. | Congjiang County, Guizhou Province, China |
|
OP764536 | OP776440 |
9 | Leptobrachella dong sp. nov. | Congjiang County, Guizhou Province, China |
|
OP764537 | OP776449 |
10 | Leptobrachella dong sp. nov. | Suining County, Hunan Province, China |
|
OP764538 | OP776442 |
11 | Leptobrachella dong sp. nov. | Suining County, Hunan Province, China |
|
OP764539 | OP776443 |
12 | Leptobrachella dong sp. nov. | Suining County, Hunan Province, China |
|
OP764540 | OP776444 |
13 | Leptobrachella dong sp. nov. | Suining County, Hunan Province, China |
|
OP764541 | OP776445 |
14 | Leptobrachella dong sp. nov. | Suining County, Hunan Province, China |
|
OP764542 | OP776446 |
15 | Leptobrachella dong sp. nov. | Suining County, Hunan Province, China |
|
OP764543 | OP776447 |
16 | L. graminicola | Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010904 | MZ224651 | / |
17 | L. graminicola | Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010905 | MZ224648 | / |
18 | L. graminicola | Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010912 | MZ224647 | / |
19 | L. graminicola | Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010908 | MZ224653 | / |
20 | L. graminicola | Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010910 | MZ224655 | / |
21 | L. graminicola | Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010909 | MZ224649 | / |
22 | L. yeae | Linggongli, Mount Emei, Sichuan Province, China | CIBEMLGL19052104 | MT957006 | MT975979 |
23 | L. yeae | Mount Emei, Sichuan Province, China | SYS a001830 | KM014810 | / |
24 | L. yeae | Mount Emei, Sichuan Province, China | CIBEM1867 | / | MT975978 |
25 | L. bourreti | Mount Pu Ta Leng, Lao Cai, Vietnam | VNMN 010916 | MZ209167 | / |
26 | L. bourreti | Bat Xat District, Lao Cai, Vietnam | ZMMU-A5636-02280 | MH055872 | / |
27 | L. bourreti | Sapa, Lao Cai Province, Vietnam | AMS R 177673 | KR018124 | / |
28 | L. bourreti | Ky Quan San, Lao Cai, Vietnam | AMS R 188515 | MZ208835 | / |
29 | L. bourreti | Sapa, Lao Cai, Vietnam | 1999.566 | KR827860 | / |
30 | L. chishuiensis | Chishui National Nature Reserve, Chishui City, Guizhou Province, China | CIBCS20190518042 | MT117054 | / |
31 | L. cf. oshanensis | Changning County, Sichuan Province, China | CIB20050095 | KC460337 | / |
32 | L. bijie | Zhaozishan Nature Reserve, Bijie City, Guizhou Province, China | SYS a007314 | MK414533 | / |
33 | L. jinshaensis | Lengshuihe Nature Reserve, Jinsha County, Guizhou Province, China |
|
MT814014 | / |
34 | L. suiyangensis | Huoqiuba Nature Reserve, Suiyang County, Guizhou Province, China | GZNU20180606005 | MK829649 | OL800396 |
35 | L. suiyangensis | Huoqiuba Nature Reserve, Suiyang County, Guizhou Province, China | GZNU20180606002 | / | OL800395 |
36 | L. purpuraventra | Wujing Nature Reserve, Bijie City, Guizhou Province, China | SYS a007081 | MK414517 | / |
37 | L. niveimontis | Daxueshan Nature Reserve, Yunnan Province, China | KIZ015734 | MT302618 | / |
38 | L. wulingensis | Tianquanshan Forest Park, Zhangjiajie, Hunan Province, China | CSUFT 177 | MT530315 | / |
39 | L. cf. oshanensis | Nanchuan District, Chongqing City, China | ZYC799 | AY526215 | / |
40 | L. dorsospina | Yushe Forest Park, Shuicheng County, Guizhou Province, China | SYS a004961 | MW046194 | / |
41 | L. alpina | Caiyanghe, Yunnan Province, China | KIZ049024 | MH055867 | MH056093 |
42 | L. purpura | Yingjiang, Yunnan Province, China | SYS a006530 | MG520354 | / |
43 | L. eos | Boun Tay, Phongsaly, Laos | NCSM 80551 | MH055887 | / |
44 | L. oshanensis | Baoguosi, Mount Emei, Sichuan Province, China | CIBEMS20190421BGS1 | MT957023 | MT975988 |
45 | L. oshanensis | Shengshuige, Mount Emei, Sichuan Province, China | CIBEMS20190422SSG1-4 | MT975985 | |
46 | L. murphyi | Doi Inthanon, Chiang Mai, Thailand | KIZ034039 | MZ710519 | / |
47 | L. yunyangensis | Qiyaoshan Nature Reserve, Yunyang County, Chongqing, China | GZNU20210622001 | OL800364 | OL800393 |
48 | L. yunyangensis | Qiyaoshan Nature Reserve, Yunyang County, Chongqing, China | GZNU20210622002 | / | OL800394 |
49 | L. tengchongensis | Gaoligong Shan, Yunnan Province, China | SYS a004598 | KU589209 | / |
50 | L. khasiorum | Khasi Hills, Meghalaya, India | SDBDU 2009.329 | KY022303 | KY022348 |
51 | L. tamdil | Mizoram, India | MZMU2224 | MW665130 | / |
52 | L. yingjiangensis | Yingjiang County, Yunnan Province, China | SYS a006533 | MG520350 | / |
53 | L. petrops | Ba Vi National Park, Ha Tay, Vietnam | ROM 13483 | MH055901 | MH056092 |
54 | L. puhoatensis | Pu Hu, Thanh Hoa, Vietnam | VNMN:2016 A.23 | KY849587 | / |
55 | L. namdongensis | Thanh Hoa Province, Vietnam | VNUF A.2017.37 | MK965389 | / |
56 | L. liui | Wuyi Shan City, Fujian Province, China | SYS a001597 | KM014547 | / |
57 | L. mangshanensis | Mangshan, Hunan Province, China | MSZTC201701 | MG132196 | / |
58 | L. verrucosa | Lianshan Bijiashan Nature Reserve, Guangdong, China | GEP a059 | OP279589 | / |
59 | L. shimentaina | Shimentai Nature Reserve, Guangdong, China | SYS a004712 | MH055926 | / |
60 | L. bashaensis | Basha Nature Reserve, Congjiang County, Guizhou Province, China | GIB196403 | MW136294 | / |
61 | L. maoershanensis | Mao’er Shan, Guangxi Province, China | KIZ07614 | MH055927 | MH056099 |
62 | L. laui | Shenzhen City, Guangdong Province, China | SYS a002450 | MH055904 | / |
63 | L. yunkaiensis | Dawuling Forest Station, Maoming City, Guangdong Province, China | SYS a004663 | MH605584 | / |
64 | L. flaviglandulosa | Xiaoqiaogou Nature Reserve, Yunnan Province, China | KIZ016072 | MH055934 | MH056098 |
65 | L. firthi | Quang Nam Province, Vietnam | AMS R 171714 | JQ739203 | / |
66 | L. isos | Gia Lai, Vietnam | AMS R 176469 | KT824767 | / |
67 | L. sungi | Tam Dao, Vinh Phuc, Vietnam | ROM 20236 | MH055858 | MH056104 |
68 | L. zhangyapingi | Chiang Mai, Thailand | KIZ07258 | MH055864 | MH056102 |
69 | L. aspera | Huanglianshan Nature Reserve, Lyuchun, Yunnan, China | SYS a007743 | MW046199 | / |
70 | L. feii | Xiaoqiaogou Nature Reserve, Yunnan Province, China | KIZ048894 | MT302634 | / |
71 | L. pelodytoides | Tam Dao, Vinh Phu, Vietnam | ROM18282 | EF397244 | / |
72 | L. ventripunctata | Wenlong, Yunnan Province, China | KIZ013621 | MH055824 | MH056090 |
73 | L. aerea | Vilabuly, Savannakhet, Laos | NCSM 76038 | MH055809 | / |
74 | L. minima | Doi Phu Fa, Nan, Thailand | KIZ024317 | MH055852 | MH056091 |
75 | L. shiwandashanensis | Fangcheng City, Guangxi Province, China | NNU202103146 | MZ326691 | / |
76 | L. wuhuangmontis | Pubei County, Guangxi Province, China | SYS a003485 | MH605577 | / |
77 | L. damingshanensis | Guangxi Province, China | NNU202103281 | MZ145229 | / |
78 | L. nyx | Ha Giang, Vietnam | ROM 36692 | MH055816 | / |
79 | L. nahangensis | Na Hang Nature Reserve, Tuyen Quang, Vietnam | ROM 7035 | MH055853 | / |
80 | L. pluvialis | Fansipan, Lao Cai, Vietnam | ROM 30685 | MH055843 | / |
81 | L. shangsiensis | Guangxi Province, China | NHMG1401032 | MK095460 | / |
82 | L. pallida | Lam Dong, Vietnam | UNS00511 | KU530190 | / |
83 | L. kalonensis | Binh Thuan Province, Vietnam | AMNH A191762 | KR018115 | / |
84 | L. bidoupensis | Bidoup-Nui Ba National Park, Lam Dong, Vietnam | ZMMU-A-4797-01454 | MH055945 | MH056110 |
85 | L. tadungensis | Dak Nong Province, Vietnam | UNS00515 | KR018121 | / |
86 | L. maculosa | Ninh Thuan Province, Vietnam | AMS R 177660 | KR018119 | / |
87 | L. pyrrhops | Loc Bac, Lam Dong, Vietnam | ZMMU-A-4873-00158 | MH055950 | MH056109 |
88 | L. macrops | Phu Yen, Vietnam | ZMMU-A5823 | MG787993 | / |
89 | L. melica | Ratanakiri, Cambodia | MVZ258198 | HM133600 | / |
90 | L. rowleyae | Da Nang City, Vietnam | ITBCZ2783 | MG682552 | / |
91 | L. applebyi | Phong Dien Nature Reserve, Thua Thien-Hue, Vietnam | KIZ010701 | MH055947 | MH056105 |
92 | L. ardens | Kon Ka Kinh National Park, Gia Lai, Vietnam | ZMMU-NAP-06099 | MH055949 | MH056108 |
93 | L. crocea | Thua Thien-Hue, Vietnam | ZMMU-NAP-02274 | MH055955 | MH056114 |
94 | L. tuberosa | Kon Ka Kinh National Park, Gia Lai, Vietnam | ZMMU-NAP-02275 | MH055959 | MH056111 |
95 | L. botsfordi | Fansipan, Lao Cai, Vietnam | AMS R 176540 | MH055952 | MH056088 |
96 | L. fuliginosa | Phetchaburi, Thailand | KUHE 20197 | LC201988 | / |
97 | L. melanoleuca | Kapoe, Ranong, Thailand | KIZ018031 | MH055967 | MH056115 |
98 | L. neangi | Veal Veng District, Pursat Province, Cambodia | CBC 1624 | MT644613 | / |
99 | L. heteropus | Larut, Perak, Malaysia | KUHE15487 | AB530453 | / |
100 | L. sola | Gunung Stong, Kelantan, Malaysia | KU RMB20973 | MH055973 | MH056119 |
101 | L. kecil | Cameron, Malaysia | KUHE 52440 | LC202004 | / |
102 | L. kajangensis | Tioman, Malaysia | LSUHC 4431 | LC202001 | / |
103 | L. dringi | Gunung Mulu, Malaysia | KUHE:55610 | AB847553 | / |
104 | L. sabahmontana | Borneo, Malaysia | BORNEENSIS 12632 | AB847551 | / |
105 | L. picta | Borneo, Malaysia | UNIMAS 8705 | KJ831295 | / |
106 | L. fritinniens | Danum Valley Field Center, Sabah, Malaysia | FMNH 244800 | MH055971 | MH056118 |
107 | L. arayai | Liwagu, Kinabalu, Malaysia | BORNEEISIS 22931 | AB847558 | / |
108 | L. hamidi | Bukit Lanjan, Selangor, Malaysia | KUHE17545 | AB969286 | / |
109 | L. marmorata | Borneo, Malaysia | KUHE53227 | AB969289 | / |
110 | L. gracilis | Bukit Kana, Sarawak, Malaysia | FMNH 273682 | MH055972 | MH056117 |
111 | L. maura | Borneo, Malaysia | SP 21450 | AB847559 | / |
112 | L. baluensis | Tambunan, Sabah, Borneo, Malaysia | SP 21604 | LC056792 | / |
113 | L. parva | Mulu National Park, Sarawak, Malaysia | KUHE:55308 | LC056791 | MH056121 |
114 | L. brevicrus | Gunung Mulu National Park, Sarawak, Malaysia | UNIMAS 8957 | KJ831303 | / |
115 | L. itiokai | Mulu National Park, Sarawak, Malaysia | KUHE:5589 | LC137805 | MH056120 |
116 | L. mjobergi | Gading NP, Sarawak, Borneo, Malaysia | KUHE:47872 | LC056787 | / |
117 | L. juliandringi | Mulu NP, Sarawak, Borneo, Malaysia | KUHE 55333 | LC056780 | / |
118 | Megophrys glandulosa | Yunnan Province, China | KIZ048439 | KX811762 | MH056125 |
119 | Leptobrachium huashen | Yunnan Province, China | KIZ049025 | KX811931 | MH056122 |
Total DNA was extracted using a standard phenol-chloroform extraction protocol (
Sequences were assembled and aligned using the Clustalw module in BioEdit v. 7.0.9.0 (
All 22 specimens of the new taxon were measured. The terminology and methods followed
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 the fourth toe);
HDL head length (distance from the tip of the snout to the articulation of jaw);
HDW head width (greatest width between the left and right articulations of jaw);
HLL hind-limb length (distance from tip of fourth toe to vent);
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);
ML manus length (distance from tip of third digit to proximal edge of inner palmar tubercle);
SL snout length (distance from the tip of the snout to the anterior corner of the eye);
TL tibia length (distance from knee to tarsus);
TYD maximal tympanum diameter;
UEW upper eyelid width (greatest width of the upper eyelid margins measured perpendicular to the anterior-posterior axis).
One tadpole specimen of the undescribed species was measured. Nineteen morphometric characters were measured for the tadpole:
BH maximum body height;
BL body length, from tip of snout to conjunction of body and tail;
BW maximum body width;
ED eye diameter (distance from the anterior corner to the posterior corner of the eye);
IND internasal distance (minimum distance between the inner margins of the external nares);
KRF keratodont row formula;
LF maximum height of lower tail fin;
NE distance between nostril and eye;
ODW oral disc width;
PP interpupilar distance;
RN rostro-narial distance;
SN snout length, from tip of snout to the anterior corner of eye;
SS distance from tip of snout to opening of spiracle;
SU distance from snout to beginning of upper tail fin;
TAL tail length;
TMH maximum tail muscle height;
TMW maximum tail muscle width;
UF maximum height of upper tail fin;
TH maximum tail height.
The new taxon was also compared with all other congeners of Leptobrachella, based on morphological characters. Comparative morphological data were obtained from literature (Table
References for morphological characters for congeners of the genus Leptobrachella.
ID | Species | Literature reviewed |
---|---|---|
1 | L. aerea (Rowley, Stuart, Richards, Phimmachak & Sivongxay, 2010) |
|
2 | L. alpina (Fei, Ye & Li, 1990) |
|
3 | L. applebyi (Rowley & Cao, 2009) |
|
4 | L. arayai (Matsui, 1997) |
|
5 | L. ardens (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
|
6 | L. aspera Wang, Lyu, Qi & Wang, 2020 |
|
7 | L. baluensis Smith, 1931 |
|
8 | L. bashaensis Lyu, Dai, Wei, He, Yuan, Shi, Zhou, Ean, Kuang, Guo, Wei & Yuan, 2020 |
|
9 | L. bidoupensis (Rowley, Le, Tran & Hoang, 2011) |
|
10 | L. bijie Wang, Li, Li, Chen & Wang, 2019 |
|
11 | L. bondangensis Eto, Matsui, Hamidy, Munir & Iskandar, 2018 |
|
12 | L. botsfordi (Rowley, Dau & Nguyen, 2013) |
|
13 | L. bourreti (Dubois, 1983) |
|
14 | L. brevicrus Dring, 1983 |
|
15 | L. chishuiensis (Li, Liu, Wei & Wang, 2020) |
|
16 | L. crocea (Rowley, Hoang, Le, Dau & Cao, 2010) |
|
17 | L. damingshanensis Chen, Yu, Cheng, Meng, Wei, Zhou & Lu, 2021 |
|
18 | L. dorsospina Wang, Lyu, Qi & Wang, 2020 |
|
19 | L. dringi (Dubois, 1987) |
|
20 | L. eos (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011) |
|
21 | L. feii (Chen, Yuan & Che, 2020) |
|
22 | L. firthi (Rowley, Hoang, Dau, Le & Cao, 2012) |
|
23 | L. flaviglandulosa (Chen, Wang & Che, 2020) |
|
24 | L. fritinniens (Dehling & Matsui, 2013) |
|
25 | L. fuliginosa (Matsui, 2006) |
|
26 | L. fusca Eto, Matsui, Hamidy, Munir & Iskandar, 2018 |
|
27 | L. gracilis (Günther, 1872) |
|
28 | L. graminicola Nguyen, Tapley, Nguyen, Luong & Rowley, 2021 |
|
29 | L. hamidi (Matsui, 1997) |
|
30 | L. heteropus (Boulenger, 1900) |
|
31 | L. isos (Rowley, Stuart, Neang, Hoang, Dau, Nguyen & Emmett, 2015) |
|
32 | L. itiokai Eto, Matsui & Nishikawa, 2016 |
|
33 | L. jinshaensis Cheng, Shi, Li, Liu, Li & Wang, 2021 |
|
34 | L. juliandringi Eto, Matsui & Nishikawa, 2015 |
|
35 | L. kajangensis (Grismer, Grismer & Youmans, 2004) |
|
36 | L. kalonensis (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
|
37 | L. kecil (Matsui, Belabut, Ahmad & Yong, 2009) |
|
38 | L. khasiorum (Das, Tron, Rangad & Hooroo, 2010) |
|
39 | L. lateralis (Anderson, 1871) |
|
40 | L. laui (Sung, Yang & Wang, 2014) |
|
41 | L. liui (Fei & Ye, 1990) |
|
42 | L. macrops (Duong, Do, Ngo, Nguyen & Poyarkov, 2018) |
|
43 | L. maculosa (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
|
44 | L. mangshanensis (Hou, Zhang, Hu, Li, Shi, Chen, Mo & Wang, 2018) |
|
45 | L. maoershanensis (Yuan, Sun, Chen, Rowley & Che, 2017) |
|
46 | L. marmorata (Matsui, Zainudin & Nishikawa, 2014) |
|
47 | L. maura (Inger, Lakim, Biun & Yambun, 1997) |
|
48 | L. melanoleuca (Matsui, 2006) |
|
49 | L. melica (Rowley, Stuart, Neang & Emmett, 2010) |
|
50 | L. minima (Taylor, 1962) |
|
51 | L. mjobergi (Smith, 1925) |
|
52 | L. murphyi Chen, Suwannapoom, Wu, Poyarkov, Xu, Pawangkhanant & Che, 2021 |
|
53 | L. namdongensis (Hoang, Nguyen, Luu, Nguyen & Jiang, 2019) |
|
54 | L. nahangensis (Lathrop, Murphy, Orlov & Ho, 1998) |
|
55 | L. natunae (Günther, 1895) |
|
56 | L. neangi Stuart & Rowley, 2020 |
|
57 | L. niveimontis (Chen, Poyarkov, Yuan & Che, 2020) |
|
58 | L. nokrekensis (Mathew & Sen, 2010) | |
59 | L. nyx (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011) |
|
60 | L. oshanensis (Liu, 1950) |
|
61 | L. pallida (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
|
62 | L. palmata Inger & Stuebing, 1992 | Inger and Stuebing (1992) |
63 | L. parva Dring, 1983 |
|
64 | L. pelodytoides (Boulenger, 1893) |
|
65 | L. petrops (Rowley, Dau, Hoang, Le, Cutajar & Nguyen, 2017) |
|
66 | L. picta (Malkmus, 1992) |
|
67 | L. pingbianensis (Rao, Hui, Zhu & Ma, 2022) | |
68 | L. platycephala (Dehling, 2012) |
|
69 | L. pluvialis (Ohler, Marquis, Swan & Grosjean, 2000) |
|
70 | L. puhoatensis (Rowley, Dau & Cao, 2017) |
|
71 | L. purpuraventra Wang, Li, Li, Chen & Wang, 2019 |
|
72 | L. purpurus (Yang, Zeng & Wang, 2018) |
|
73 | L. pyrrhops (Poyarkov, Rowley, Gogoleva, Vassilieva, Galoyan & Orlov, 2015) |
|
74 | L. rowleyae (Nguyen, Poyarkov, Le, Vo, Ninh, Duong, Murphy & Sang, 2018) |
|
75 | L. sabahmontana (Matsui, Nishikawa & Yambun, 2014) |
|
76 | L. serasanae Dring, 1983 |
|
77 | L. shangsiensis Chen, Liao, Zhou & Mo, 2019 |
|
78 | L. shimentaina Wang, Lyu & Wang, 2022 |
|
79 | L. shiwandashanensis Chen, Peng, Pan, Liao, Liu & Huang, 2021 |
|
80 | L. sola (Matsui, 2006) |
|
81 | L. sungi (Lathrop, Murphy, Orlov & Ho, 1998) |
|
82 | L. suiyangensis (Luo, Xiao, Gao & Zhou, 2020) |
|
83 | L. tadungensis (Rowley, Tran, Le, Dau, Peloso, Nguyen, Hoang, Nguyen & Ziegler, 2016) |
|
84 | L. tamdil (Sengupta, Sailo, Lalremsanga, Das & Das, 2010) |
|
85 | L. tengchongensis (Yang, Wang, Chen & Rao, 2016) |
|
86 | L. tuberosa (Inger, Orlov & Darevsky, 1999) |
|
87 | L. ventripunctata (Fei, Ye & Li, 1990) |
|
88 | L. verrucosa Wang, Zeng, Lin & Li, 2022 |
|
89 | L. wuhuangmontis Wang, Yang & Wang, 2018 |
|
90 | L. wulingensis Qian, Xiao, Cao, Xiao & Yang, 2020 |
|
91 | L. yeae Shi, Hou, Song, Jiang & Wang, 2021 |
|
92 | L. yingjiangensis (Yang, Zeng & Wang, 2018) |
|
93 | L. yunkaiensis Wang, Li, Lyu & Wang, 2018 |
|
94 | L. yunyangensis Luo, Deng & Zhou, 2022 |
|
95 | L. zhangyapingi (Jiang, Yan, Suwannapoom, Chomdej & Che, 2013) |
|
The advertisement calls of the new taxon were recorded from specimens
The aligned sequence matrix of 16S and RAG1 gene contained 498 bps and 888 bps, respectively. ML and BI analyses, based on the 16S gene matrix, resulted in essentially identical topologies (Fig.
Phylogenetic trees of the genus Leptobrachella and a haplotype network constructed based on RAG1 gene sequences A maximum Likelihood (ML) tree reconstructed, based on mitochondrial 16S gene sequences B maximum Likelihood tree reconstructed, based on nuclear RAG1 gene sequences. Bayesian posterior probabilities (BPP) from BI analyses/ bootstrap supports (BS) from ML analyses are listed beside the nodes. The symbol “-” represents a value below 0.50/50. For information of samples 1–119, refer to Table
The undescribed species could be identified from its congeners in a series of morphological and bioacoustics characters. For the detailed demonstration, based on morphological and bioacoustics comparisons, see the following section describing the new species.
Molecular phylogenetic analyses, morphological comparisons and bioacoustics analyses indicated that the specimens from Tongdao and Suining County, Hunan Province and Congjiang County, Guizhou Province, China represent an undescribed species which is described as follows.
Holotype.
Paratypes. Four adult males
Leptobrachella dong sp. nov. is assigned to the genus Leptobrachella, based on molecular data and the following morphological characters: medium size, rounded finger tips, the presence of an elevated inner palmar tubercle not continuous to the thumb, presence of macroglands on body (including supra-axillary, pectoral and femoral glands), vomerine teeth absent, tubercles on eyelids and anterior tip of snout with vertical white bar (
Leptobrachella dong sp. nov. could be distinguished from its congeners by a combination of the following characters: (1) body of medium size (SVL 29.2–32.0 mm in 15 adult males and 37.4–43.1 mm in seven adult females); (2) distinct black spots present on flanks; toes rudimentary webbed, with wide lateral fringes; (3) ventral belly white with distinct nebulous brown speckling on ventrolateral flanks; (4) skin on dorsum shagreened with fine tiny granules or short ridges; (5) heels overlapped when thighs are positioned at right angles to the body; (6) tibia-tarsal articulation reaches the middle eye; (7) dorsal surface of tadpole semi-transparent light brown, spots on tail absent, keratodont row formula I: 3+3/2+2: I; (8) calls with two types, at dominant frequency (5.1 ± 0.4 kHz).
Adult male. SVL in 32.0 mm. Head width almost equal with head length slightly (HDW / HDL 1.03); snout rounded in both ventral view and lateral view, projecting slightly beyond margin of the lower jaw; nostril closer to snout than eye; loreal region oblique; canthus rostralis indistinct; eyes large (ED / HDL 0.40), eye diameter slightly longer than snout length (ED / SL 1.07), eyes notably protuberant in both dorsal and lateral views, pupil vertical; tympanum distinct, rounded, tympanum diameter smaller than eye (TYD / ED 0.38), upper margin of tympanum in contact with supratympanic ridge; vomerine teeth absent; tongue notched behind; supratympanic ridge distinct, extending from posterior corner of eye to supra-axillary gland.
Fore-limb relatively long (LAL / SVL 0.46), fingers long and slender (ML / SVL 0.25), webbing absent, lateral fringes on fingers narrow; relative finger lengths II < I < IV < III; tips of fingers rounded and slightly swollen; subarticular tubercles absent on fingers, inner metacarpal tubercle large and rounded, separated from the smaller, round outer metacarpal; supra-axillary glands oval.
Hind-limb relatively long (HLL / SVL 1.53), heels overlapping when the tibias perpendicular to the body axis; tibio-tarsal articulation of adpressed limb reaching middle of eye, tibia length about half of snout-vent length (TL / SVL 0.49); relative toe length: I < II < V < III < IV; toe tips rounded and slightly swollen; rudimentary webbing present between all five toes; wide lateral fringes present on all toes; dermal ridges under fourth toes interrupted; subarticular tubercles distinct under the base of II, III and IV toe; inner metatarsal tubercle oval and distinct, outer metatarsal tubercle absent (Fig.
Dorsal skin relatively smooth with small tubercles and short folds; supra-axillary gland distinct and yellowish; pectoral gland small and indistinct; round femoral glands present and protuberant on rear of thigh, closer to knee than to vent; femoral adipose glands distinct, attached to inner side of skin on posterior ventral surface of thigh; ventral skin smooth; ventrolateral glands forming a distinct white line on flanks.
In life, dorsal surface of head and trunk yellowish-brown, with distinct olive reverse-triangle dark markings between eyes connecting to a dark W-shaped marking between axillae that are fringed with greyish-white colour; elbow to upper arm distinctly yellowish-orange in colour on the dorsum; four transverse black bars present on dorsal surface of thighs and three on dorsal surface of lower arm; one dark blotch between nostril and eyes on loreal region and a dark blotch under the eye; supratympanic ridge reddish and a large black marking under supratympanic ridge; distinct dark blotches on flanks from groin to axilla, longitudinally in two rows; ventral surfaces light coloured; throat and ventral arms pinkish with cream speckling on margins; chest and belly cream white, on the lateral belly with dense brown speckling; ventral hind-limbs pinkish with sparse white glands; upper iris copper, lower iris silver.
Dorsum of body and limbs fading to brown copper; transverse bars on limbs becoming more distinct. Ventral surface of body and limbs fading to cream white. Supra-axillary, femoral and pectoral glands fading to cream yellow.
Measurements and basic statistics of adult specimens are presented in Suppl. materials
(Fig.
Advertisement calls of Leptobrachella dong sp. nov. A, B waveform and sonogram of the first call type (type A) over 2 seconds of the paratype
Compared with the 26 known congeners occurring south of the Isthmus of Kra, Leptobrachella dong sp. nov. could be distinguished from them by several characters: by having supra-axillary and ventrolateral glands, the new species differs from L. arayai, L. dringi, L. fritinniens, L. gracilis, L. hamidi, L. heteropus, L. kajangensis, L. kecil, L. marmorata, L. maura, L. melanoleuca, L. picta, L. platycephala, L. sabahmontana, and L. sola (vs. absent in the latter); by having rounded fingertips and moderate body size (29.2–34.2 mm in 15 adult males and 34.4–43.1 mm in seven adult females), the new species differs from the following species with pointed fingertips and smaller body size: L. baluensis (14.9–15.9 mm in males), L. bondangensis (17.8 mm in male), L. brevicrus (17.1–17.8 mm in males), 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. palmata (14.4–16.8 mm in males), L. parva (15.0–16.9 mm in males) and L. serasanae (16.9 mm in female).
Leptobrachella dong sp. nov. could also be identified from 65 known Leptobrachella species occurring north of the Isthmus of Kra by some characters (see Table
Basic statistics for the measurements of Leptobrachella dong sp. nov. Units in mm. See abbreviations for characters in the Materials and methods section.
Measurement | Male (n = 15) | Female (n = 7) | ||
---|---|---|---|---|
Ranging | Mean ± SD | Ranging | Mean ± SD | |
SVL | 29.2–34.2 | 30.9 ± 1.4 | 34.4–43.1 | 39.4 ± 2.8 |
HDL | 9.8–11.7 | 10.5 ± 0.6 | 11.8–13.8 | 12.7 ± 0.7 |
HDW | 9.8–12.2 | 11.0 ± 0.7 | 12.4–14.5 | 13.7 ± 0.8 |
SL | 4.0–5.0 | 4.5 ± 0.3 | 4.9–6.3 | 5.6 ± 0.4 |
IND | 2.8–4.4 | 3.3 ± 0.4 | 3.4–4.7 | 4.3 ± 0.5 |
IOD | 2.3–3.9 | 3.2 ± 0.5 | 3.4–4.1 | 3.6 ± 0.2 |
UEW | 2.6–3.4 | 2.9 ± 0.2 | 2.9–4.3 | 3.6 ± 0.5 |
ED | 3.6–4.4 | 4.1 ± 0.2 | 4.2–5.3 | 4.8 ± 0.4 |
TYD | 1.5–2.2 | 1.7 ± 0.2 | 1.6–3.0 | 2.5 ± 0.4 |
LAL | 13.5–15.5 | 14.6 ± 0.5 | 16.8–18.9 | 17.9 ± 0.9 |
ML | 7.2–8.6 | 7.9 ± 0.4 | 8.6–10.5 | 9.6 ± 0.7 |
TL | 14.5–16.4 | 15.2 ± 0.7 | 17.4–19.6 | 18.3 ± 0.8 |
FL | 13.5–15.5 | 14.5 ± 0.6 | 17.4–19.4 | 17.9 ± 1.0 |
HLL | 41.3–51.6 | 47.6 ± 3.1 | 54.6–62.5 | 58.4 ± 2.9 |
Diagnosis characters on morphology of Leptobrachella dong sp. nov. from other congeners.
ID | Species | Male SVL (mm) | Female SVL (mm) | Black spots on flanks | Toes webbing | Fringes on toes | Ventral colouration | Dorsal skin texture |
---|---|---|---|---|---|---|---|---|
1 | Leptobrachella dong sp. nov. | 29.2–32.0 | 34.4–43.1 | Yes | Rudimentary | Wide | White with distinct nebulous brown speckling on ventrolateral flanks | Shagreened with fine tubercles |
2 | L. aerea | 25.1–28.9 | 27.1–38.6 | No | Rudimentary | Wide | Near immaculate creamy-white, brown speckling on margins | Finely tuberculate |
3 | L. alpina | 24.0–26.4 | 31.7–32.5 | Yes | Rudimentary | Wide in males | Creamy-white with dark spots | Relatively smooth, some with small warts |
4 | L. applebyi | 19.6–22.3 | 21.7 | Yes | Rudimentary | No | Reddish-brown with white speckling | Smooth |
5 | L. ardens | 21.3–24.7 | 25.4 | Yes | No | No | Reddish-brown with white speckling | Smooth-finely shagreened |
6 | L. aspera | 22.4 | 25.0–26.4 | Yes | Rudimentary | Narrow | Creamy-white with distinct dark patches on chest and abdomen | Rough with dense conical granules, tubercles and glandular folds |
7 | L. bashaensis | 22.9–25.6 | 27.1 | Yes | Rudimentary | Narrow | Creamy-white chest and off-white belly with irregular black spots | Dorsal skin slightly shagreened with small tubercles and irregular brown stripes |
8 | L. bidoupensis | 23.6–24.6 | 29.2–29.4 | Yes | Rudimentary | Weak | Reddish-brown with white speckling | Smooth |
9 | L. bijie | 29.0–30.4 | / | Yes | Rudimentary | Narrow | White with distinct nebulous greyish speckling on chest and ventrolateral flanks | Shagreened and granular |
10 | L. botsfordi | 29.1–32.6 | 30.0–31.8 | No | Rudimentary | Narrow | Reddish-brown with white speckling | Shagreened |
11 | L. bourreti | 27.4–36.2 | 39.5–45.0 | Yes | Rudimentary | Weak | Creamy-white | Relatively smooth, some with small warts |
12 | L. chishuiensis | 30.8–33.4 | 34.2 | Yes | Rudimentary | Narrow | White with distinct nebulous greyish speckling on chest and ventrolateral flanks | Shagreened and granular |
13 | L. crocea | 22.2–27.3 | / | No | Rudimentary | No | Bright orange | Highly tuberculate |
14 | L. damingshanensis | 33.6–34.4 | / | Yes | Rudimentary | Narrow | Creamy-white ventral surface with small, creamy-white glands on throat, chest and belly, becoming more concentrated near lateral margin | Rough dorsal skin with sparse jacinth tubercles and some short longitudinal ridges |
15 | L. dorsospina | 28.7–30.5 | 32.1–39.8 | Yes | Rudimentary | Narrow | Greyish-white with black spots and orange pigmentations | Rough with dense conical granules, tubercles, glandular folds and conical spines |
16 | L. eos | 33.1–34.7 | 40.7 | No | Rudimentary | Wide | Creamy-white | Shagreened |
17 | L. feii | 21.5–22.8 | 25.7 | Yes | Rudimentary | Narrow | Creamy-white with black blotches | Shagreened with small tubercles and ridge |
18 | L. firthi | 26.4–29.2 | 25.7–36.9 | No | Rudimentary | Wide in males | Creamy-white | Shagreened with fine tubercles |
19 | L. flaviglandulosa | 23.0–27.0 | 29.3 | Yes | Poorly developed | Narrow | Whitish, black speckling on margins | Shagreened with yellowish-brown tubercles |
20 | L. fuliginosa | 28.2–30.0 | / | Yes | Rudimentary | Weak | White with brown dusting | Nearly smooth, few tubercles |
21 | L. graminicola | 23.1–24.6 | 28.6–32.9 | No | Rudimentary | Wide | White with brown spots | Smooth, with many tubercles |
22 | L. isos | 23.7–27.9 | 28.6–31.5 | No | Rudimentary | Wide in males | Creamy-white with white dusting on margins | Mostly smooth, females more tuberculate |
23 | L. jinshaensis | 29.7–31.2 | / | Yes | No | Narrow | Cream yellow, presence of distinct nebulous greyish speckling on flanks | Shagreened and granular |
24 | L. kalonensis | 25.8–30.6 | 28.9–30.6 | Yes | No | No | Pale, speckled brown | Smooth |
25 | L. khasiorum | 24.5–27.3 | 31.2–33.4 | Yes | Rudimentary | Wide | Creamy white | Isolated, scattered tubercles |
26 | L. lateralis | 26.9–28.3 | 36.6 | Yes | Rudimentary | No | Creamy white | Roughly granular |
27 | L. laui | 24.8–26.7 | 28.1 | Yes | Rudimentary | Wide | Creamy-white with dark brown dusting on margins | Round granular tubercles |
28 | L. liui | 23.0–28.7 | 24.5–27.8 | Yes | Rudimentary | Wide | Creamy-white with dark brown spots on chest and margins | Round granular tubercles with glandular folds |
29 | L. macrops | 28.0–29.3 | 30.3 | Yes | Rudimentary | No | Greyish-violet with white speckling | Roughly granular with larger tubercles |
30 | L. maculosa | 24.2–26.6 | 27 | Yes | No | No | Brown, less white speckling | Mostly smooth |
31 | L. mangshanensis | 22.2–27.8 | 30.2 | Yes | Rudimentary | Weak | White speckles on throat and belly | Nearly smooth |
32 | L. maoershanensis | 25.2–30.4 | 29.1 | Yes | Rudimentary | Narrow | Creamy-white chest and belly with irregular black spots | Longitudinal folds |
33 | L. melica | 19.5–22.8 | / | Yes | Rudimentary | No | Reddish-brown with white speckling | Smooth |
34 | L. minima | 25.7–31.4 | 31.6–37.3 | Yes | Rudimentary | No | Creamy-white | Smooth |
35 | L. murphyi | 23.2–24.9 | 29.3–32.1 | Yes | Rudimentary | Wide | Creamy-white belly with small black spots on the margin | Shagreened with reddish tubercles and folds |
36 | L. nahangensis | 40.8 | / | Yes | Rudimentary | No | Creamy-white with light specking on throat and chest | Smooth |
37 | L. namdongensis | 30.9 | 32.1–35.3 | Yes | Rudimentary | No | Creamy-white with brown dusting on margins | Finely tuberculate |
38 | L. neangi | / | 35.4–36.3 | Yes | Rudimentary (in females) | absent (in females) | Light purplish-grey with dark brown mottling on throat | Small, irregular bumps and ridges |
39 | L. niveimontis | 22.5–23.6 | 28.5–28.7 | Yes | Rudimentary | No | Ventral sides marbled with distinct irregular black speckling | Skin on dorsum scattered with fine reddish tubercles |
40 | L. nokrekensis | 26.0–33.0 | 34.0–35.0 | Yes | Rudimentary | unknown | White with distinct nebulous greyish speckling on chest and ventrolateral flanks | Tubercles and longitudinal folds |
41 | L. nyx | 26.7–32.6 | 37.0–41.0 | Yes | Rudimentary | No | Creamy-white with white and brown margins | Rounded tubercles |
42 | L. oshanensis | 26.6–30.7 | 31.6 | Yes | No | No | Whitish with no markings or only small, light grey spots | Smooth with few glandular ridges |
43 | L. pallida | 24.5–27.7 | / | No | No | No | Reddish-brown with white speckling | Tuberculate |
44 | L. pelodytoides | 27.5–32.3 | / | Yes | Wide | Narrow | Whitish | Small, smooth warts |
45 | L. petrops | 23.6–27.6 | 30.3–47.0 | No | No | Narrow | Immaculate creamy white | Highly tuberculate |
46 | L. pluvialis | 21.3–22.3 | 25.5–33.5 | Yes | Rudimentary | No | Dirty white with dark brown marbling | Smooth, flattened tubercles on flanks |
47 | L. puhoatensis | 24.2–28.1 | 27.3–31.5 | Yes | Rudimentary | Narrow | Reddish-brown with white dusting | Longitudinal skin ridges |
48 | L. purpurus | 25.0–27.5 | / | Yes | Rudimentary | Wide | Dull white with indistinct grey dusting | Shagreen with small tubercles |
49 | L. purpuraventra | 27.3–29.8 | 33.0–35.3 | Yes | Rudimentary | Narrow | Grey purple with distinct nebulous greyish speckling on chest and ventrolateral flanks | Shagreened and granular |
50 | L. pyrrhops | 30.3–33.9 | 30.8–34.3 | Yes | Rudimentary | No | Reddish-brown with white speckling | Slightly shagreened |
51 | L. rowleyae | 23.4–25.4 | 27.0–27.8 | Yes | No | No | Pinkish milk-white to light brown chest and belly with numerous white speckles | Smooth with numerous tiny tubercles |
52 | L. shangsiensis | 24.9–29.4 | 30.8–35.9 | Yes | Rudimentary | Narrow | ventral surface yellowish-creamy-white with marble texture | Smooth |
53 | L. shiwandashanensis | 26.8–29.7 | / | Yes | No | No | Creamy-white ventral surface with small, creamy-white glands on throat, chest and belly, becoming more concentrated near lateral margin | Shagreened with small raised tubercles and ridges |
54 | L. sungi | 48.3–52.7 | 56.7–58.9 | No or small | Wide | Weak | White | Granular |
55 | L. suiyangensis | 28.7–29.7 | 30.5–33.5 | Yes | Rudimentary | Narrow | Yellowish-creamy-white with marble texture chest and belly or with irregular light brown speckling | Shagreen with small granules |
56 | L. tadungensis | 23.3–28.2 | 32.1 | Yes | No | No | Reddish-brown with white speckling | Smooth |
57 | L. tamdil | 32.3 | 32.3 | Yes | Wide | Wide | White | Weakly tuberculate |
58 | L. tengchongensis | 23.9–26.0 | 28.8–28.9 | Yes | Rudimentary | Narrow | White with dark brown blotches | Shagreened with small tubercles |
59 | L. tuberosa | 24.4–29.5 | 30.2 | No | Rudimentary | No | White with small grey spots/streaks | Highly tuberculate |
60 | L. ventripunctata | 23.7–27.7 | 31.5–35.0 | Yes | Rudimentary | No | Chest and belly with dark brown spots | Longitudinal skin ridges |
61 | L. wuhuangmontis | 25.6–30.0 | 33.0–36.0 | Yes | Rudimentary | Narrow | Greyish-white mixed by tiny white and black dots | Rough, scattered with dense conical tubercles |
62 | L. wulingensis | 24.5–32.8 | 29.9–38.5 | Yes | Rudimentary | Narrow | Creamy white, with distinct or indistinct brown speckling at margins | Shagreened with sparse large warts, sometimes with longitudinal ridges |
63 | L. yeae | 25.8–32.6 | 33.7–34.1 | Yes | Rudimentary | Narrow | Ventral belly cream white with variable brown specking | Dorsum relatively smooth with fine tiny granules or short ridges |
64 | L. yingjiangensis | 25.7–27.6 | / | Yes | Rudimentary | Wide | Creamy-white with dark brown flecks on chest and margins | Shagreened with small tubercles |
65 | L. yunkaiensis | 25.9–29.3 | 34.0–35.3 | Yes | Rudimentary | Wide | Belly pink with distinct or indistinct speckling | Shagreened with short skin ridges and raised warts |
66 | L. zhangyapingi | 45.8–52.5 | / | No | Rudimentary | Wide | Creamy-white with white and brown | Mostly smooth with distinct tubercles |
67 | L. pingbianensis | 28 | 30 | Yes | Rudimentary | Unknown | Chest and belly with dark brown spots | Smooth |
68 | L. shimentaina | 26.4–8.9 | 30.1–30.7 | Yes | Rudimentary | Wide in males | Greyish-pink with distinct hazy brown speckling on chest and ventrolateral flanks | Round granular tubercles with glandular folds |
69 | L. verrucosa | 23.2–25.9 | / | Yes | Rudimentary | Narrow | Creamy-white with greyish-white and dark brown spots | Shagreened with numerous conical tubercles |
70 | L. yunyangensis | 28.3–30.6 | / | Yes | Rudimentary | Narrow | Light greyish-creamy-white, interspersed with light brown spots | Rough with sparse large warts, with short longitudinal ridges |
By having medium size of body (SVL 29.2–34.2 mm in males) Leptobrachella dong sp. nov. differs from the smaller males L. aerea (25.1–28.9 mm), L. alpina (24.0–26.4 mm), L. applebyi (19.6–22.3 mm), L. ardens (21.3–24.7 mm), L. aspera (22.4 mm), L. bashaensis (22.9–25.6 mm), L. bidoupensis (23.6–24.6), L. crocea (22.2–27.3 mm), L. feii (21.5–22.8 mm), L. flaviglandulosa (23.0–27.0 mm), L. isos (23.7–27.9 mm), L. graminicola (23.1–24.6 mm), L. khasiorum (24.5–27.3 mm), L. lateralis (26.9–28.3 mm), L. laui (24.8–26.7 mm), L. liui (24.8–26.7 mm), L. maculosa (24.2–26.6 mm), L. mangshanensis (22.22–27.76 mm), L. maura (26.1 mm), L. melica (19.5–22.8 mm), L. murphyi (23.2–24.9 mm), L. niveimontis (22.5–23.6 mm), L. pallida (24.5–27.7 mm), L. petrops (23.6–27.6 mm), L. pluvialis (21.3–22.3 mm), L. puhoatensis (24.2–28.1 mm), L. pyrrhops (25.0–27.5 mm), L. rowleyae (23.4–25.4 mm), L. tadungensis (23.3–28.2 mm), L. tengchongensis (23.9–26.0 mm), L. ventripunctata (23.7–27.7 mm) and L. yingjiangensis (25.7–27.6 mm) and differs from the larger in males L. nahangensis (40.8 mm), L. platycephala (35.1 mm), L. sungi (48.3–52.7 mm in males) and L. zhangyapingi (45.8–52.5 mm).
By having a larger size of body (SVL 34.4–43.1 mm in females), Leptobrachella dong sp. nov. differs from the smaller females L. alpina (31.7–32.5 mm), L. applebyi (21.7 mm), L. ardens (25.4 mm), L. aspera (25.0–26.4 mm), L. bashaensis (27.1), L. botsfordi (30.0–31.8 mm), L. graminicola (28.6–32.9 mm), L. isos (28.6–31.5 mm), L. kalonensis (28.9–30.6 mm), L. khasiorum (31.2–33.4 mm), L. liui (24.5–27.8 mm), L. macrops (30.3 mm), L. maculosa (27 mm), L. mangshanensis (30.2 mm), L. maoershanensis (29.1 mm), L. murphyi (29.3–32.1 mm), L. macrops (30.3 mm), L. maoershanensis (29.1 mm), L. niveimontis (28.5–28.7 mm), L. oshanensis (31.6 mm), L. pluvialis (25.5–33.5 mm), L. puhoatensis (27.3–31.5 mm), L. rowleyae (27.0–27.8 mm), L. shimentaina (30.1–30.7 mm), L. suiyangensis (30.5–33.5 mm), L. tadungensis (32.1 mm), L. tamdil (32.3 mm), L. tengchongensis (28.8–28.9 mm) and L. tuberosa (30.2 mm).
By having black spots on flanks, Leptobrachella dong sp. nov. differs from L. aerea, L. botsfordi, L. crocea, L. eos, L. firthi, L. isos, L. pallida, L. petrops, L. tuberosa and L. zhangyapingi (vs. lacking distinct black spots on the flanks in the latter).
By having rudimentary webbing, Leptobrachella dong sp. nov. differs from L. ardens, L. jinshaensis, L. kalonensis, L. maculosa, L. oshanensis, L. pallida, L. petrops, L. rowleyae, L. shiwandashanensis and L. tadungensis (vs. absent webbing in the latter).
By having wide fringes on toes, Leptobrachella dong sp. nov. differs from L. applebyi, L. ardens, L. aspera, L. bashaensis, L. bidoupensis, L. bijie, L. botsfordi, L. bourreti, L. chishuiensis, L. crocea, L. damingshanensis, L. dorsospina, L. feii, L. flaviglandulosa, L. fuliginosa, L. jinshaensis, L. kalonensis, L. lateralis, L. macrops, L. maculosa, L. mangshanensis, L. melica, L. minima, L. nahangensis, L. namdongensis, L. niveimontis, L. nyx, L. oshanensis, L. pallida, L. pelodytoides, L. petrops, L. pluvialis, L. puhoatensis, L. purpuraventra, L. pyrrhops, L. rowleyae, L. shangsiensis, L. shiwandashanensis, L. sungi, L. tengchongensis, L. tuberosa, L. ventripunctata, L. verrucosa, L. wuhuangmontis, L. wulingensis, L. yeae and L. yunyangensis (vs. fringes on toes narrow or absent in the latter).
By having dorsal surface shagreened with fine tubercles, Leptobrachella dong sp. nov. differs from L. applebyi, L. bidoupensis, L. kalonensis, L. melica, L. minima, L. nahangensis, L. pingbianensis, L. shangsiensis and L. tadungensis, all of which have the dorsum smooth and L. bourreti (dorsum smooth with small warts), L. fuliginosa (dorsum smooth with fine tubercles), L. liui (dorsum with round tubercles), L. macrops (dorsum roughly granular with large tubercles), L. maoershanensis (dorsum shagreened with tubercles), L. minima (dorsum smooth), L. neangi (dorsum with small, irregular bumps and ridges), L. nyx (dorsum with round tubercles), L. nokrekensis (dorsum tubercles and longitudinal folds), L. pelodytoides (dorsum with small, smooth warts), L. tamdil (dorsum weakly tuberculate, with low, oval tubercles), L. tuberosa (dorsum highly tuberculate), L. yunkaiensis (dorsum with raised warts) and L. wuhuangmontis (dorsum rough with conical tubercles).
The advertisement calls of Leptobrachella dong sp. nov. (Results and Fig.
In mitochondrial DNA trees, Leptobrachella dong sp. nov. was clustered as an independent clade and sister to a clade comprising of L. graminicola and L. yeae. The new species differs from L. graminicola by the following characters: body size larger with SVL 29.2–34.2 mm in adult males and 34.4–43.1 mm in adult females (vs. 23.1–24.6 mm in adult males and 28.6–32.9 mm in adult females); black spots on flanks present (vs. absent); ventral surface white with distinct nebulous brown speckling on ventrolateral flanks (vs. white with brown spots); dorsal surface shagreened with fine tubercles (vs. smooth, with many tubercles); and tibiotarsal articulation reaching to middle of eye (vs. anterior edge of eye). The new species differs from L. yeae by having wide fringes on toes (vs. narrow); dorsal surface shagreened with fine tubercles (vs. relatively smooth with fine tiny granules or short ridges); and males with a pair of subgular internal vocal sacs (vs. internal single subgular vocal sac).
(in mm). Description based on sequenced tadpole
Adult males with a pair of subgular vocal sacs (Fig.
Habitats of Leptobrachella dong sp. nov. A landscape of the type locality Tongdao County, Hunan Province, China B a torrent mountain stream in the type locality (insert holotype
Leptobrachella dong sp. nov. has been found in three localities: Tongdao County and Suining County, Hunan Province and Congjiang County, Guizhou Province, China. Elevations recorded range from 620 m to 1200 m. Population from the Tongdao County inhabited a torrent stream covered by evergreen shrubs and the new species always found on the stones (Fig.
This specific name “dong” refers to the Dong people, as the new species distributed in the concentrated area of Dong people. We suggest its English common name “Dong leaf litter toads” and Chinese name “Dong Zhang Tu Chan (侗掌突蟾)”.
South-western China was proposed as a biodiversity hotspot (
This new species was found in three localities and the largest geographical distance between the localities is over 200 km. However, in this study, phylogenetic analyses, based on mitochondrial DNA, suggested the three populations as the same species and different from its congeners on a series of morphological characters. This perhaps indicated that the species have a widespread distribution. Further surveys are needed to evaluate the population status of the species.
We are grateful for the kind help from Dong people, especially Zhen Shi and her families in Shangdong Village, Tongdao County, Hunan Province. We are grateful for the help of Fu Shu and Keji Guo of Central South Inventory and Planning Institute of National Forestry and Grassland Administration, Jingbo Long of Huangsang National Nature Reserve and Yongjian QIN of Forestry and Grassland Administration of Suining County for their help in fieldwork in Suining County, Hunan Province.
This work was supported by the Projects from the National Natural Science Foundation of China (Nos. 32270498, 31960099, 32260136, and 32070426); West Light Foundation of The Chinese Academy of Sciences (Grant No. 2021XBZG_XBQNXZ_A_006); Special funds of Guiyang Bureau of Science and Technology for supporting the Guiyang College (GYKJ-GYU[2021]); Guizhou Provincial Science and Technology Projects (Nos. [2020]1Y083, ZK[2022]540 and [2020]4Y029); Forestry Science and Technology Research Project of Guizhou Forestry Department (No. [2020]13, J[2018]03 and [2022]41); Zunyi Bureau of Science and Technology (No. HZ[2020]319); Guizhou Provincial Department of Education Youth Science and Technology Talents Growth Project (Nos. KY[2018]470, KY[2018]452 and KY[2020]234); Science Research Project of Moutai Institute (No.myk2019009); and Biodiversity Survey of Suining County, Hunan Province.
Measurements of adult specimen of Leptobrachella dong sp. nov.
Data type: table (excel document)
Explanation note: Morphological data. Units in mm. See abbreviations for characters in the Materials and methods section.
Uncorrected p-distance between Leptobrachella species on the 16S rRNA gene
Data type: table (excel document)
Explanation note: Genetic distance. The values below 3% are coloured in red.
Call data
Data type: table (excel document)
Explanation note: Call measurements (mean ± standard deviation) of Leptobrachella dong sp. nov.