Morphology, molecular genetics, and acoustics reveal two new species of the genus Leptobrachella from northwestern Guizhou Province, China (Anura, Megophryidae)
Author
Wang, Jian
Author
Li, Yu-Long
Author
Li, Yao
Author
Chen, Hong-Hui
Author
Zeng, Ya-Jun
Author
Jing-Min Shen,
Author
Wang, Ying-Yong
text
ZooKeys
2019
848
119
154
http://dx.doi.org/10.3897/zookeys.848.29181
journal article
http://dx.doi.org/10.3897/zookeys.848.29181
1313-2970-848-119
F688F9C1BD6E4986BD3E9C219C4DF585
F688F9C1BD6E4986BD3E9C219C4DF585
Leptobrachella bijie J. Wang, Y.L. Li, Y. Li, H.H. Chen & Y.Y. Wang
sp. nov.
Figure 5
Holotype.
SYS a007316, adult male, collected by Jian Wang (JW hereafter) and Yulong Li (YLL hereafter) on 6 July 2018 from Qingshan Village (
27°39'24"N
,
105°23'14"E
; 1670 m a.s.l.) in Zhaozishan Nature Reserve, Linkou Town, Qixingguan District, Bijie City, Guizhou Province, China.
Paratypes.
Seven adult males, SYS a007313/ CIB 110002, SYS a007314-7315, 7317-7320, collected by Honghiu Chen (HHC hereafter), Yongyou Zhao (YYZ hereafter) and Jiahe Li (JHL), the same collection data as the holotype.
Diagnosis.
(1) small size (SVL 29.0-30.4 mm in eight adult males), (2) dorsal skin shagreened, some of the granules forming longitudinal short skin ridges, (3) iris bicolored, coppery orange on upper half and silver on lower half, (4) tympanum distinctly discernible, slightly concave, distinct black supratympanic line present, (5) internasal distance equal to interorbital distance, (6) supra-axillary, femoral, pectoral and ventrolateral glands distinctly visible, (7) absence of webbing and lateral fringes on fingers, toes with rudimentary webbing and narrow lateral fringes, (8) longitudinal ridges under toes not interrupted at the articulations, (9) relative finger lengths I = II = IV <III, relative toe length I <II <V = III <IV, (10) heels just meeting, tibia-tarsal articulation reaches the region between middle of eye to anterior corner of eye, (11) dorsal surface shagreened and granular, lacking enlarge tubercles or warts, some of the granules forming short longitudinal folds, (12) dorsum greyish-brown grounding, with small light orange granules, distinct darker brown markings scattered with irregular light orange pigmentations, (13) flanks with several dark blotches, longitudinally in two rows, (14) ventral surface white, with distinct nebulous greyish speckling on chest and ventrolateral flanks, (15) dorsal limbs including fingers and toes with dark bars, and (16) dense tiny conical spines present on surface of chest in males during breeding season.
Figure 5. General aspect in life:
A-D
SYS a007316, the male holotype of
Leptobrachella bijie
sp. nov. E the male paratype SYS a007313 F the male paratype SYS a007317.
Comparisons.
Comparative morphological data of
Leptobrachella bijie
sp. nov. and 45 recognized
Leptobrachella
species occurring north of the Isthmus of Kra were listed in Table 5.
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. bijie
sp. nov. can be easily 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 lacking supra-axillary and ventrolateral glands; and by the significantly larger body size, SVL 29.0-30.4 mm in males,
L. bijie
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),
L. serasanae
(16.9 mm in female), and
Dring's
(1983)
Leptobrachella
sp. 3
"baluensis"
(15.0-16.0 mm in males).
Table 5. Selected diagnostic characters for species described herein and species in the genus
Leptobrachella
occurring north of the Isthmus of Kra (modified from
Rowley et al. 2017
;
Yuan et al. 2017
;
Yang et al. 2018
; Wang et al. 2018).
| Species |
Male
SVL
(mm)
|
Black spots on flanks |
Toes webbing |
Fringes on toes |
Ventral coloration |
Dorsal skin texture |
|
L. bijie
|
|
L. purpuraventra
|
|
L. aerea
|
|
L. alpinus
|
|
L. applebyi
|
|
L. ardens
|
|
L. bidoupensis
|
|
L. botsfordi
|
|
L. bourreti
|
|
L. crocea
|
|
L. eos
|
|
L. firthi
|
|
L. fuliginosa
|
|
L. isos
|
|
L. kalonensis
|
|
L. khasiorum
|
|
L. lateralis
|
|
L. laui
|
|
L. liui
|
|
L. macrops
|
|
L. maculosa
|
|
L. maoershanensis
|
|
L. mangshanensis
|
|
L. melica
|
|
L. minima
|
|
L. nahangensis
|
|
L. nokrekensis
|
|
L. nyx
|
|
L. oshanensis
|
|
L. pallida
|
|
L. pelodytoides
|
|
L. petrops
|
|
L. pluvialis
|
|
L. puhoatensis
|
|
L. purpura
|
|
L. pyrrhops
|
|
L. rowleyae
|
|
L. sungi
|
|
L. tadungensis
|
|
L. tamdil
|
|
L. tengchongensis
|
|
L. tuberosa
|
|
L. ventripunctata
|
|
L. wuhuangmontis
|
|
L. yingjiangensis
|
|
L. yunkaiensis
|
|
L. zhangyapingi
|
For the remaining 45 members of the genus
Leptobrachella
, having SVL of 29.0-30.4 mm in males,
L. bijie
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. alpinus
(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. isos
(23.7-27.9 mm in males),
L. khasiorum
(24.5-27.3 mm in males),
L. lateralis
(26.9-28.3 mm in males),
L. laui
(24.8-26.7 mm in males),
L. maculosa
(24.2-26.6 mm in males),
L. melica
(19.5-22.7 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).
In having black spots on flanks, the new species differs from
L. aerea
,
L. botsfordi
,
L. firthi
, and
L. tuberosa
, all of which lacking distinct black spots on the flanks; by having rudimentary webbing on toes, the new species differs from
L. kalonensis
and
L. oshanensis
, both of which lacking webbing on toes, and from
L. pelodytoides
, which bears wide webbing on toes; by having narrow lateral fringes on toes, the new species differs from
L. aerea
,
L. firthi
,
L. liui
, and
L. yunkaiensis
, all of which having wide lateral fringes on toes, from
L. bourreti
and
L. fuliginosa
, both of which having weak lateral fringes on toes, and from
L. kalonensis
,
L. macrops
,
L. minima
,
L. nyx
,
L. oshanensis
,
L. pyrrhops
, and
L. tuberosa
, all of which lacking lateral fringes on toes; by having dorsal surface shagreened and granular, lacking enlarge tubercles or warts, the new species differs from
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. nyx
(dorsum with round tubercles),
L. pelodytoides
(dorsum with small, smooth warts),
L. tuberosa
(dorsum hingly tuberculate),
L. yunkaiensis
(dorsum with raised warts), and
L. wuhuangmontis
(dorsum rough with conical tubercles); by having ventral surface white with distinct nebulous greyish speckling on chest and flanks, the new species differs from
L. botsfordi
and
L. pyrrhops
, (ventral reddish brown with white speckling),
L. maoershanensis
(belly with irregular black spots); by having tiny spines on surface of chest in males during breeding season, the new species differs from all male specimens collected in breeding season of
L. liui
,
L. oshanensis
,
L. yunkaiensis
, and
L. wuhuangmontis
, all of which are lacking such spines.
Description of holotype.
Adult male. Body size small, SVL in 29.3 mm. Head length slightly larger than head width, HDL/HDW 1.03; 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 equal to interorbital distance, IND/IOD 1.00; pineal ocellus absent; vertical pupil; snout length larger than eye diameter, SNT/EYE 1.11; tympanum distinct, rounded, and slightly concave, diameter smaller than that of the eye and larger than tympanum-eye distance, TMP/EYE 0.53 and TEY/TMP 0.47; upper margin of tympanum incontact 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 behind; 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; a large, rounded inner palmar tubercle distinctly separated from small, round outer palmar tubercle; absence of webbing and lateral fringes on fingers. Tips of toes like fingers; relative toe length I <II <V = III <IV; subarticular tubercles absent; distinct 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 reaches to middle of eye; heels just meeting each other when thighs are appressed at right angles with respect to body.
Dorsal surface shagreened and granular, lacking enlarge tubercles or warts, some of the granules forming short longitudinal folds; ventral skin smooth; dense tiny conical spines present on surface of chest; pectoral gland and femoral gland oval; pectoral glands greater than tips of fingers and femoral glands; femoral gland situated on posteroventral surface of thigh, closer to knee than to vent; supra-axillary gland raised. Ventrolateral gland distinctly visible, forming an incomplete line.
Measurements of holotype (in mm).
SVL 29.2, HDL 10.0, HDW 9.7, SNT 4.0, EYE 3.6, IOD 3.0, IND 3.0, TMP 1.9, TEY 0.9, TIB 13. 8, ML 7.8, PL 13.2, LAHL 14.1, HLL 43.3.
Coloration of holotype in life.
Dorsum greyish-brown grounding, with small reddish granules, distinct darker brown markings and rounded spots and scattered with irregular light orange pigmentation. A dark brown inverted triangular pattern between anterior corner of eyes, in connected to the dark brown W-shaped marking on interorbital region, and the W-shaped marking in connected to the other W-shaped marking between axillae. Tympanum brown. Small light orange granules present on dorsum of body and limb; a dark brown vertical bar under the eye; transverse dark brown bars on dorsal surface of limbs; distinct dark brown blotches on flanks from groin to axilla, longitudinally in two rows; elbow and upper arms with dark bars and distinct coppery orange coloration; fingers and toes with distinct dark bars.
Ventral surface of throat, chest, and belly white, presence of distinct nebulous greyish speckling on chest and ventrolateral flanks; ventral surface of limbs grey purple. Supra-axillary gland coppery orange; femoral, pectoral and ventrolateral glands greyish white. Iris bicolored, coppery orange on upper half and silver on lower half.
Coloration of holotype in preservative.
Dorsum of body and limbs dark brown; transverse bars on limbs become more distinct; dark brown patterns, markings and spots on back become indistinct, orange pigmentations become greyish white. Ventral surface of body and limbs greyish white, nebulous speckling on chest and flanks balck brown. Supra-axillary, femoral, pectoral and ventrolateral glands greyish white.
Variations.
Measurements and body proportions were listed in Table 6. All paratypes match the overall characters of the holotype except that: coloration of tympanum brown in the holotype SYS a007316 (vs. black in paratypes SYS a007313/ CIB 110002 (Figure 5E), SYS a007315, 7317 (Figure 5F)); heels just meeting, tibia-tarsal articulation reaching the middle of eye in the holotype (vs. heels slightly overlapping in paratypes SYS a007315, 7317, 7319-7320; tibia-tarsal articulation reaching the anterior corner of eye in paratypes SYS a007315, 7317, 7319); W-shaped marking on interorbital region in connected to the other W-shaped marking between axillae in the holotype (vs. such markings not in connected with each other in paratypes SYS a007313/ CIB 110002, SYS a007320); a dark brown inverted triangular pattern between anterior corner of eyes in the holotype (vs. a V-shaped pattern between anterior corner of eyes instead in paratype SYS a007317, 7320); relatively larger black spots on flanks (vs. black spots distinctly small in paratypes SYS a007313/ CIB 110002, SYS a007317).
Table 6. Measurements (
minimum-maximum
(mean
+/-
SD); in mm), and body proportions of
Leptobrachella bijie
sp. nov. from Qingshan Village of Zhaozishan Nature Reserve.
| SEX |
Males (n = 8) |
| SVL |
HLL |
| HDL |
HDL/HDW |
| HDW |
HDL/SVL |
| SNT |
SNT/HDL |
| EYE |
SNT/EYE |
| IOD |
EYE/TMP |
| IND |
IND/IOD |
| TMP |
TMP/EYE |
| TEY |
TEY/TMP |
| TIB |
TIB/SVL |
| ML |
LAHL/SVL |
| PL |
HLL/SVL |
| LAHL |
TIB/HLL |
Etymology.
The specific epithet bijie is in reference to the type locality, Qingshan Village in Bijie City of Guizohu Province, China. For the common name, we suggest "Bijie Leaf Litter Toad", and for the Chinese name "Bi Jie Zhang Tu Chan (
毕节掌突蟾)"
.
Distribution and habits.
Currently,
Leptobrachella bijie
sp. nov. is known only from its type locality Qingshan Village in Zhaozishan Nature Reserve, Linkou County, Qixingguan District, Bijie City, Guizhou Province, China (Figure 1). The new species was found along a clear-water rocky stream (ca. 2 m in width and ca. 20-30 cm in depth; 1670-1750 m a.s.l.) in karst landforms. The stream was surrounded by broad-leaved forest at an altitude below 1700 m, and by coniferous forest at an altitude above 1700 m (Figure 6, 1700 m a.s.l.). On 6 July 2018 at 22:00-23:30 P.M., a large number of males were found calling on leaves of plants (Figure 10A), and some were found calling perching on the rocks or under rocks by the side of the stream.
Figure 6. The habitat of
Leptobrachella bijie
sp. nov. in Qingshan Village of Zhaozishan Nature Reserve in Guizhou Province.