A new Leptobrachella species (Anura, Megophryidae) from South China, with comments on the taxonomic status of L. chishuiensis and L. purpurus Author Wang, Jian https://orcid.org/0000-0003-4249-7767 State Key Laboratory of Biocontrol, Sun Yat-sen University, Guangzhou 510275, China & The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China Author Qi, Shuo https://orcid.org/0000-0002-2924-6093 State Key Laboratory of Biocontrol, Sun Yat-sen University, Guangzhou 510275, China Author Dai, Ke-Yuan School of Ecology, Sun Yat-sen University, Guangzhou 510275, China Author Lyu, Zhi-Tong https://orcid.org/0000-0002-4891-3859 State Key Laboratory of Biocontrol, Sun Yat-sen University, Guangzhou 510275, China Author Zeng, Zhao-Chi State Key Laboratory of Biocontrol, Sun Yat-sen University, Guangzhou 510275, China & The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China Author Chen, Hong-Hui State Key Laboratory of Biocontrol, Sun Yat-sen University, Guangzhou 510275, China & The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China Author Li, Yuan-Qiu School of Ecology, Sun Yat-sen University, Guangzhou 510275, China Author Zhao, Yong-You School of Ecology, Sun Yat-sen University, Guangzhou 510275, China Author Wang, Yun-Ze Shuanghuamu Biological Technology Co., Ltd, Shenzhen 518000, China Author Wang, Ying-Yong State Key Laboratory of Biocontrol, Sun Yat-sen University, Guangzhou 510275, China wangyy@mail.sysu.edu.cn text Zoosystematics and Evolution 2022 2022-06-03 98 1 165 180 http://dx.doi.org/10.3897/zse.98.73162 journal article http://dx.doi.org/10.3897/zse.98.73162 1860-0743-1-165 EE314DBE46504C50B1D2953FDD47F876 FAA9F860168D569CAF6376719661F100 Leptobrachella shimentaina J. Wang, Z.-T. Lyu & Y.-Y. Wang sp. nov. Shimentai Leaf Litter Toad (in English) / Shi Men Tai Zhang Tu Chan (石门台掌突蟾in Chinese) Figs 3 , 4 , 5 , 6 Chresonymy. " Leptolalax sp.6" ( Chen et al. 2018 ). Type material. Holotype ♂. SYS a007684, from Jiuzhou Yizhan ( 24°23'38.01"N , 113°06'35.38"E ; ca. 300 m a.s.l.), Shimentai Nature Reserve, Yingde City, Guangdong Province, China, collected by Jian Wang and Yong-You Zhao on 23 April 2019. Figure 3. Morphological features of the holotype of Leptobrachella shimentaina sp. nov., SYS a007684 in life: A. Dorsal view; B. Dorsolateral view; C. Ventral view; D. Character of eye and temporal region; E. Ventral view of hand; F. Ventral view of foot. Paratypes (n = 7). 5 ♂: SYS a007683/ CIB116079, same collection place as holotype; SYS a007685-7686, collected by Yu-Long Li and Hong-Hui Chen on 23 April 2019 from Shijiao ( 24°26'35.31"N , 113°09'21.55"E ; ca 400 m a.s.l.), Shimentai Nature Reserve; SYS a007687, collected by Jian Wang and Yong-You Zhao on 24 April 2019 from Longtankeng ( 24°26'31.38"N , 113°15'28.81"E ; ca 310 m a.s.l.), Shimentai Nature Reserve; SYS a008329, collected by Jian Wang on 27 April 2020 from Luokeng Nature Reserve ( 24°31'42.3"N , 113°15'54.3"E ; ca 590 m a.s.l.), Shaoguan City, Guangdong Province, China. 2 ♀: SYS a004711-4712, collected by Ying-Yong Wang, Jian Wang, and Zhi-Tong Lyu on 27 April 2016 from Hengshitang ( 24°26'14.92"N , 113°17'50.48"E ; ca 380 m a.s.l.), Shimentai Nature Reserve. Etymology. The specific epithet " shimentaina " is an adjective derived from "shimentai" , referring to the type locality of the new species, Shimentai Nature Reserve, " ina " is used as a feminine suffix which indicates the relationship of position. Diagnosis. (1) small body size [SVL 26.4-28.9 mm in six adult males, SVL 30.1 and 30.7 mm in two adult females], (2) iris bicolored, upper half coppery orange and lower half silver, (3) tympanum distinct, (4) supratympanic line black, (5) fingers unwebbed, with distinct lateral fringes in males [absence in females], (6) toes with rudimentary webbing, lateral fringes wide in males [narrow in females], (7) longitudinal ridges under toes continuous, with constrictions at interphalangeal articulations, (8) heels slightly overlapping when adpressed, tibial-tarsal articulation reaching mid-orbit, (9) relative lengths of fingers I = II = IV <III, and toe I <II <III = V <IV, (10) dorsal surface shagreened and granular, lacking enlarged warts, with some granules forming short longitudinal folds, (11) dorsum grayish brown to yellowish brown, with small light orange granules and distinct darker brown scattered markings with irregular light orange pigmentation, (12) flanks with several dark spots, (13) ventral surface grayish pink, with distinct hazy brown speckling on chest and ventrolateral flanks, (14) densely-distributed, small, raised pectoral and abdominal tubercles present; (15) dorsal surfaces of forelimbs and digits with dark transverse bars. Description of holotype. Adult male. Body size small, SVL in 28.6 mm. Head length slightly larger than head width, HDW/HDL 0.91; snout slightly protruding, projecting slightly beyond margin of lower jaw; nostril closer to snout than eye; canthus rostralis gently rounded; loreal region slightly concave; interorbital space flat, internarial distance larger than interorbital distance, IND/IOD 1.10; pineal ocellus absent; pupil vertical; snout longer than eye diameter, SNT/EYE 1.19; tympanum distinct, rounded, diameter smaller than that of eye and larger than tympanum-eye distance, TMP/EYE 0.41, TEY/TMP 0.48; upper margin of tympanum in contact with supratympanic ridge; vomerine teeth absent; a single vocal sac; vocal sac openings slit-like, paired, located posterolaterally on floor of mouth, close to margins of 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 = II = IV <III; nuptial pad absent; subarticular tubercles absent; inner palmar tubercle large, rounded, distinctly separated from outer palmar tubercle; outer palmar tubercle small, rounded; fingers lacking interdigital webbing, with distinct lateral fringes. Tips of toes rounded, slightly swollen; relative toe length I <II <III = V <IV; longitudinal ridges under toes continuous, with constrictions at interphalangeal articulations; inner metatarsal tubercle large, oval; outer metatarsal tubercle absent; interdigital webbing between toes rudimentary; wide lateral fringes present on all toes. Tibia 47% of snout-vent length; tibiotarsal articulation reaches to middle of eye; heels slightly overlapping when thighs are appressed at right angles with respect to body. Dorsal surface shagreened and granular, lacking enlarged tubercles or warts, some granules forming short longitudinal folds; ventral skin smooth, densely-distributed small raised scapular and abdominal tubercles present; pectoral and femoral glands oval; both larger in diameter than tip of fingers; femoral glands larger in diameter than tip of toes; femoral gland situated on posteroventral surface of thigh, closer to knee than to vent; supra-axillary gland raised. Ventrolateral gland distinctly visible, forming an longitudinal uncontinuous series. Coloration of holotype in life. Dorsal background color yellowish-brown, with small orange granules, distinct dark brown markings, and rounded spots, and scattered with irregular light grayish-brown pigmentation. A dark brown inverted triangular marking present between anterior corners of eyes, connecting to dark brown W-shaped marking in interorbital region. This W-shaped marking is in connection to the other W-shaped marking on occipital region. Tympanum dark brown, lower margin grayish yellow. Supratympanic line black. Dorsal surfaces of body and limbs with small orange granules; a pair of dark brown vertical bars present under eyes; transverse dark brown bars on dorsal surfaces of distal limbs and digits; distinct dark brown blotches on flanks; surfaces of elbows and upper arms coppery orange, without dark bars. Surface of throat, chest, and belly grayish pink, with distinct hazy brown speckling on chest and ventrolateral flanks, without black spots; surface of chin and lower lip brown with grayish white spots and patches; ventral surface of limbs brown with pink hues. Supra-axillary gland coppery orange, pectoral glands grayish white, femoral glands coppery orange, ventrolateral glands brown. Iris bicolored, upper half coppery orange, lower half silver. Coloration of holotype in preservative (Fig. 4 ). Dorsal background color grayish brown, scattered with irregular light grayish-brown pigmentation. All markings, bars and spots become more distinct. Tympanum dark brown, lower margin gray. Figure 4. The holotype of Leptobrachella shimentaina sp. nov., SYS a007684 in preservative. Ventral surface yellowish brown; speckling on chest and ventrolateral flanks become more distinct; ventral surface of limbs dark brown. Supra-axillary, pectoral, and ventrolateral glands greyish yellow. Variation. Measurements and body proportions are listed in Table 2 . All male paratypes from Shimentai Nature Reserve (SYS a007683/ CIB116079, SYS a007685 (Fig. 5B ), 7686 (Fig. 5C ), 7687) show very similar morphological characters to holotype SYS a007684; paratype SYS a008329 (Fig. 6 ) from Luokeng Nature Reserve shows a darker coloration on dorsum, and brighter coloration on ventrum. Moreover, female paratypes, SYS a004711 (Fig. 5A ) and SYS a004712 show lighter background coloration and more obvious dark brown marking orange tubercles on back. Table 2. Measurements (minimum-maximum (mean +/- SD); in mm), and body proportions of Leptobrachella shimentaina sp. nov.

Vocher	SYS a007683	SYS a007684	SYS a007685	SYS a007686	SYS a007687	SYS a008329	Range	Vocher	SYS a004711	SYS a004712	Range
Sex	Male	Male	Male	Male	Male	Male	Males (n = 6)	Sex	Female	Female	Females (n = 2)
SVL	28.9	28.6	28.4	28.7	28.4	26.4	26.4-28.9 (28.2 +/- 0.9)	SVL	30.1	30.7	30.1-30.7 (30.4 +/- 0.4)
HDL	11.7	10.7	10.8	10.9	10.8	9.8	9.8-11.7 (10.8 +/- 0.6)	HDL	10.1	10.4	10.1-10.4 (10.3 +/- 0.2)
HDW	9.7	9.7	9.7	9.7	9.7	9.2	9.2-9.7 (9.6 +/- 0.2)	HDW	9.7	9.9	9.7-9.9 (9.8 +/- 0.2)
SNT	4.3	4.2	4.1	4.3	4.2	3.3	3.3-4.3 (4.1 +/- 0.4)	SNT	4.0	4.3	4.0-4.3 (4.1 +/- 0.2)
IND	2.9	2.7	2.9	2.8	2.7	2.9	2.7-2.9 (2.8 +/- 0.1)	IND	2.7	2.8	2.7-2.8 (2.7 +/- 0.1)
IOD	2.6	2.5	2.5	2.5	2.5	2.6	2.5-2.6 (2.5 +/- 0.1)	IOD	2.7	2.8	2.7-2.8 (2.7 +/- 0.0)
EYE	3.5	3.5	3.4	3.4	3.4	3.3	3.3-3.5 (3.4 +/- 0.1)	EYE	3.5	3.6	3.5-3.6 (3.5 +/- 0.1)
TMP	1.5	1.5	1.5	1.5	1.5	1.5	1.5-1.5 (1.5 +/- 0.0)	TMP	1.5	1.8	1.5-1.8 (1.7 +/- 0.2)
TEY	0.7	0.7	0.7	0.7	0.7	0.6	0.6-0.7 (0.7 +/- 0.0)	TEY	0.6	0.9	0.6-0.9 (0.8 +/- 0.2)
ML	7.4	7.2	7.2	7.4	7.3	6.9	6.9-7.4 (7.2 +/- 0.2)	ML	7.2	7.8	7.2-7.8 (7.5 +/- 0.4)
LAHL	14.4	14.0	14.4	14.5	14.4	12.5	12.5-14.5 (14.0 +/- 0.8)	LAHL	14.1	15.0	14.1-15.0 (14.6 +/- 0.6)
PL	12.6	12.6	13.0	12.2	12.6	11.9	11.9-13.0 (12.5 +/- 0.4)	PL	13.1	12.5	12.5-13.1 (12.8 +/- 0.4)
TIB	14.2	13.3	13.3	13.3	13.4	12.4	12.4-14.2 (13.3 +/- 0.6)	TIB	14.1	14.6	14.1-14.6 (14.3 +/- 0.4)
HLL	45.1	44.5	44.5	45.1	44.9	39.61	39.6-45.1 (43.9 +/- 2.1)	HLL	44.4	45.6	44.4-45.6 (45.0 +/- 0.9)
HDL/SVL	0.40	0.37	0.38	0.38	0.38	0.37	0.37-0.40 (0.38 +/- 0.01)	HDL/SVL	0.34	0.34	0.34-0.34 (0.34 +/- 0.00)
HDW/SVL	0.34	0.34	0.34	0.34	0.34	0.35	0.34-0.35 (0.34 +/- 0.00)	HDW/SVL	0.32	0.32	0.32-0.32 (0.32 +/- 0.00)
HDW/HDL	0.83	0.91	0.90	0.89	0.90	0.94	0.83-0.94 (0.90 +/- 0.03)	HDW/HDL	0.95	0.95	0.95-0.95 (0.95 +/- 0.00)
SNT/HDL	0.37	0.39	0.38	0.39	0.39	0.34	0.34-0.39 (0.38 +/- 0.02)	SNT/HDL	0.39	0.41	0.39-0.41 (0.40 +/- 0.02)
IND/HDW	0.30	0.28	0.29	0.28	0.28	0.31	0.28-0.31 (0.29 +/- 0.01)	IND/HDW	0.28	0.28	0.28-0.28 (0.28 +/- 0.01)
IOD/HDW	0.27	0.25	0.26	0.25	0.25	0.29	0.25-0.29 (0.26 +/- 0.01)	IOD/HDW	0.28	0.28	0.28-0.28 (0.28 +/- 0.00)
EYE/HDL	0.30	0.33	0.31	0.31	0.32	0.34	0.30-0.34 (0.32 +/- 0.01)	EYE/HDL	0.34	0.35	0.34-0.35 (0.34 +/- 0.00)
TMP/EYE	0.42	0.41	0.43	0.43	0.43	0.45	0.41-0.45 (0.43 +/- 0.01)	TMP/EYE	0.44	0.50	0.44-0.50 (0.47 +/- 0.04)
ML/SVL	0.25	0.25	0.25	0.26	0.26	0.26	0.25-0.26 (0.26 +/- 0.00)	ML/SVL	0.24	0.25	0.24-0.25 (0.25 +/- 0.01)
LAHL/SVL	0.50	0.49	0.51	0.50	0.51	0.47	0.47-0.51 (0.50 +/- 0.01)	LAHL/SVL	0.47	0.49	0.47-0.49 (0.48 +/- 0.01)
PL/SVL	0.43	0.44	0.46	0.42	0.44	0.45	0.42-0.46 (0.44 +/- 0.01)	PL/SVL	0.43	0.41	0.41-0.43 (0.42 +/- 0.02)
TIB/SVL	0.49	0.47	0.47	0.46	0.47	0.47	0.46-0.49 (0.47 +/- 0.01)	TIB/SVL	0.47	0.47	0.47-0.47 (0.47 +/- 0.01)
HLL/SVL	1.56	1.55	1.57	1.57	1.58	1.50	1.50-1.58 (1.55 +/- 0.03)	HLL/SVL	1.47	1.48	1.47-1.48 (1.48 +/- 0.01)

Figure 5. Morphological features of the paratypes of Leptobrachella shimentaina sp. nov.: A1. A female (SYS a004711 in situ; ventral view of the same specimen's hand [A2] and foot [A3]; B. Male SYS a007685 in situ; C. Male SYS a007686 in situ. Figure 6. Morphological features of a paraype (SYS a007687) of Leptobrachella shimentaina sp. nov. in life. Distribution and ecology. Leptobrachella shimentaina sp. nov. is known from its type locality Shimentai Nature Reserve, Yingde City, and the adjacent Luokeng Nature Reserve, Shaoguan City, Guangdong, China (Fig. 1 ; ca. 300-600 m a.s.l.). In the Hengshitang Protection Station of the Shimentai Nature Reserve, the new species is sympatric with L. mangshanensis . The new species inhabits primitive clear-water rocky streams surrounded by broad-leaved forest in karst landforms or granite landforms. From April to June, males were observed calling while perched on the rocks or under rocks in flowing streams, and females possessed mature oocytes. Comparisons. In our phylogenetic analyses (Fig. 2 , Table 3 ) Leptobrachella shimentaina sp. nov. clusters with L. bashaensis , L. laui , L. liui , L. mangshanensis , L. maoershanensis , and L. yunkaiensis , thus, these six species constitute the most important comparisons, for the purpose of diagnosis and recognition of the new species. In having wide fringes along lateral surfaces of toes in males, it can be distinguished from L. bashaensis , L. mangshanensis and L. maoershanensis (vs. lateral fringes narrow); by ventral coloration, grayish pink with distinct hazy brown speckling on chest and ventrolateral flanks; it can be distinguished from L. bashaensis (vs. creamy-white chest and belly with irregular black spots), L. laui (vs. creamy white with dark brown dusting on ventrolateral flanks), L. liui (vs. creamy white with dark brown spots on chest and ventrolateral flanks), L. maoershanensis (vs. creamy white chest and belly with irregular black spots), and L. yunkaiensis (vs. belly pink with distinct or indistinct speckling); by having dorsal skin with round granular tubercles and glandular folds, the new species can be further distinguished from L. laui (vs. dorsal skin lacking glandular folds), L. mangshanensis (vs. dorsal skin almost smooth with tiny transparent spines), and L. yunkaiensis (vs. dorsal skin shagreened with short skin ridges and raised warts); by dense small raised tubercles present on surface of chest and both sides of abdomen; it can be further distinguished from L. mangshanensis , L. liui , L. yunkaiensis , L. maoershanensis and L. laui (vs. ventral surface smooth); by having longitudinal ridges under toes continuous, with constrictions at interphalangeal articulations, it can be distinguished from L. laui (vs. longitudinal ridges without constrictions at interphalangeal articulations), L. liui (vs. longitudinal ridges interrupted at interphalangeal articulations), L. maoershanensis (vs. longitudinal ridges indistinct and not interrupted at interphalangeal articulations), and L. yunkaiensis (vs. longitudinal ridges without constrictions at interphalangeal articulations). Table 3. Comparisons of selected diagnostic characters for the new species and its phylogenetically related congeners.

Leptobrachella spp.	Male SVL (mm)	Toes webbing	Fringes on toes	Longitudinal ridges under toes (interrupt + / not interrupt -)	Ventral coloration	Dorsal skin texture
L. shimentaina sp. nov.	26.4-28.9	Rudimentary	Wide in males	-	Grayish pink with distinct hazy brown speckling on chest and ventrolateral flanks	Round granular tubercles with glandular folds
L. laui	24.8-26.7	Rudimentary	Wide	-	Creamy white with dark brown dusting on ventrolateral flanks	Round granular tubercles
L. liui	23.0-28.7	Rudimentary	Wide	+	Gray white to creamy white with dark brown spots on chest and ventrolateral flanks	Shagreened with numerous small tubercles, round warts and sparse short skin ridges
L. mangshanensis	22.2-27.8	Rudimentary	Narrow	-	Creamy white belly, scattered with white speckles	Almost smooth with tiny transparent spines, small tubercles and sparse short skin ridges
L. maoershanensis	25.2-30.4	Rudimentary	Narrow	-	Creamy white chest and belly with irregular black spots	With longitudinal folds
L. bashaensis	22.9-25.6	Rudimentary	Narrow	-	Creamy-white chest and belly with irregular black spots	Slightly shagreened with small tubercles
L. yunkaiensis	25.9-29.3	Rudimentary	Wide	-	Belly pink with distinct or indistinct speckles	Shagreened with short skin ridges and raised warts

Compared with the 26 known congeners of the genus Leptobrachella occurring south of the Isthmus of Kra, by the presence of supra-axillary and ventrolateral glands, L. shimentaina sp. nov. can easily be distinguished 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 supra-axillary and ventrolateral glands; and by its distinctly larger male body size, SVL 26.4-28.9 mm, L. shimentaina sp. nov. differs from the smaller L. baluensis (14.9-15.9), L. brevicrus (17.1-17.8), L. bondangensis (17.8), L. fusca (16.3), L. itiokai (15.2-16.7s), L. juliandringi (17.0-17.2), L. mjobergi (15.7-19.0), L. natunae (17.6), L. parva (15.0-16.9), and L. palmata (14.4-16.8); the female of L. serasanae (16.9), is also smaller than L. shimentaina sp. nov. (females 30.1-30.7). For the remaining 60 members of the genus Leptobrachella , males L. shimentaina sp. nov. (SVL 26.4-28.9 mm) differs from males of the larger L. chishuiensis (30.8-33.4), L. damingshanensis (33.6-34.4), L. eos (33.1-34.7), L. graminicola (23.1-24.6), L. neangi (30.9), L. nahangensis (40.8), L. pyrrhops (30.8-34.3), L. sungi (48.3-52.7), L. tamdil (32.3), and L. zhangyapingi (45.8-52.5); and from the smaller L. aspera (22.4), L. applebyi (19.6-22.3), L. ardens (21.3-24.7), L. bidoupensis (18.5-25.4), L. feii (21.5-22.8), L. melica (19.5-22.7), L. murphyi (23.2-24.9), L. niveimontis (22.5-23.6), L. pluvialis (21.3-22.3), and L. rowleyae (23.4-25.4). Through its possession of toes with rudimentary webbing and with wide lateral fringes in males, the new species can be distinguished from L. bijie , L. dorsospina , L. flaviglandulosa , L. jinshaensis , L. purpuraventra , L. puhoatensis , L. shangsiensis , L. suiyangensis , L. tengchongensis , L. wuhuangmontis and L. yeae (lateral fringes narrow), L. bourreti , L. fuliginosa (lateral fringes weak), L. jinshaensis , L. kalonensis , L. maculosa , L. oshanensis , L. shiwandashan , L. tadungensis , L. ventripunctata (no webbing or lateral fringes), L. lateralis , L. namdongensis , L. macrops , L. minima , L. nyx (no lateral fringes), and L. pelodytoides (extensive webbing and narrow lateral fringes), and additionally from L. alpina , L. khasiorum , L. nokrekensis , L. yingjiangensis (ventral coloration creamy white) and L. purpurus (ventral coloration dull white) by having greyish pink ventral coloration. By having black spots on flanks, it further differs from L. aerea , L. botsfordi , L. crocea , L. firthi , L. isos , L. pallida , L. petrops , and L. tuberosa , all of which lack black spots on the flanks.