Leptobrachella wulingensis Qian & Xia & Cao & Xiao & Yang 2020, sp. nov.
Creators: name: Qian, Tian-Yu
name: Xia, Xin
name: Cao, Yue
name: Xiao, Neng-Wen
name: Yang, Dao-De
Publisher: name: Zenodo
Year: 2020
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Megophryidae
Leptobrachella
Leptobrachella wulingensis

description: Leptobrachella wulingensis sp. nov.
Figs. 3–6.
Holotype. CSUFT 201, adult female, collected by Tianyu Qian and Xin Xia from Tianzishan Nature Reserve, Zhangjiajie, Hunan Province, China (29°23′12.67″N, 110°27′29.6″E), elevation 979 m. on September 9, 2019.
Paratypes. CSUFT 177, adult female, CSUFT 178, subadult female and CSUFT 179, 180, two adult males, collected by Tianyu Qian and Xin Xia from Tianquanshan Forest Park, Zhangjiajie, Hunan Province, China (29°15′57.58″N, 110°12′13.38″E), elevation 414 m. on September 7, 2019; CSUFT 193, adult male and CSUFT 194, adult female, collected by Tianyu Qian and Xin Xia from Tianquanshan Forest Park (29°15′39.43″N, 110°13′29.47″E), elevation 530 m. on September 8, 2019; CSUFT 200, adult male, CSUFT 202–204, three subadult females, collected by Tianyu Qian and Xin Xia from the same location as holotype, on September 9, 2019; CSUFT 301, subadult female and CSUFT 302, 303, two subadult males, collected by Yue Cao and Xin Xia from the same location as the holotype, on August 6, 2019.
Etymology. The specific name wulingensis is derived from the Wuling Mountains, where the two collection localities occur. The suggested common name is the “Wuling leaf-litter toad” in English and “Wǔ Líng Zhǎng Tū Chán (ĒỄẠẊḇ)” in Chinese.
Diagnosis. Leptobrachella wulingensis sp. nov. can be distinguished from its congeners by having (1) medium body size (SVL 24.5–32.8 mm in four adult males, 29.9–38.5 mm in three adult females); (2) dorsum brown to reddish brown with indistinct markings; (3) dorsal skin shagreened with sparse, large warts, sometimes with longitudinal ridges; (4) ventral surface of body creamy white or translucent creamy white, with distinct or indistinct pale brown speckling on chest and margins, and marbled texture on belly; small tubercle-like white dots presents on chest and anterior region of abdomen in some specimens; (5) tympanum distinct, slightly concave; (6) flanks with small to moderate black spots, with glandular warts present among these spots; (7) absence of webs and lateral fringes on fingers, toes with rudimentary webbing and narrow lateral fringes; (8) distinct longitudinal ridges under toes and interrupted at the articulations; (9) tibia length 44%–49% of SVL in males; (10) iris bicolored with bright orange or golden upper half, fades to silver in lower half; (11) ventrolateral glands distinctly visible, forming an incomplete line; small supra-axillary and femoral glands present; pectoral glands indistinct; (12) tongue cordiform, notched behind; (13) and dense small white conical spines present on lateral and ventral surface of tarsus, surface of tibia-tarsal, inner-side surface of shank and around cloacal region.
Description of holotype. Adult female, medium in body size, SVL 31.1 mm; head length about equal to head width, HDL/HDW 1.03; snout rounded, snout length equal to eye diameter, SNT/EYE 1.00; nostril closer to snout than eye; canthus rostralis gently rounded; loreal region slightly concave; pupil vertical; vomerine teeth absent; tympanum rounded, slightly concave, upper margin in contact with supratympanic ridge; supratympanic fold from eye towards supra-axillary gland; tongue cordiform, notched behind.
Tips of fingers rounded, slightly swollen; relative finger lengths III> IV = II> I; subarticular tubercles absent; inner palmar tubercle large and rounded, connected with a smaller, round outer palmar tubercle; absence of webbing and lateral fringes on fingers; tips of toes rounded and thickened; relative toe length IV> III> V> II> I; subarticular tubercles absent, replaced by longitudinal dermal ridges, extending on phalanges and interrupted at the articulations; elongate, large oval inner metatarsal tubercle present; toes webbing rudimentary; narrow lateral fringes present on all toes. Tibia 47% of snout-vent length; tibiotarsal articulation reaches to middle of eye.
Dorsal surface shagreened, the region between eyes relatively smooth, dense large warts present on the posterior part of dorsum; upper eyelids and limbs with small tubercles; flanks with distinct glandular warts forming two rows; skin on ventral surfaces of trunk, head and forelimbs smooth; white conical spines present on lateral and ventral surface of tarsus, surface of tibia-tarsal, inner-side surface of shank and surface around cloacal region; pectoral gland elongate, and femoral gland oval; pectoral glands greater than femoral glands, PEC/FEM 1.08; femoral glands situated on the posteroventral surface of thigh, closer to knee than to vent; supra-axillary gland raised; ventrolateral gland line distinctly visible.
Coloration of holotype in life. Dorsal surface brown with a dark inverted triangular marking in the interorbital region and followed by a "W" shaped mark between axillae; a “Y” shape mark between the loreal region linked to the triangular marking between the eyes; three vertical bars present at the rostral-eye region; supratympanic line weak, lower edge black; granules present on dorsum, flanks and limbs having dark orange pigmentation in the center; moderate black spots presents on flanks; glandular warts on flanks white to orange; transverse bars presents on lower arms and legs, as well as fingers and toes; elbow and upper arms coppery orange.
Ventral surface of throat and limbs greyish-pink, chest greyish pink with creamy white pigmentation, belly creamy white but translucent with greyish-pink skin color; dense small tubercle-like white dots present on chest; marbled texture present on belly; vaguely visible brown flecks present on chest and margin of belly, close to the ventrolateral gland line; supra-axillary gland orange, ventrolateral glands whitish orange, femoral and pectoral glands white; iris bicolored, golden in the upper half, and fade to dark silver in lower half.
Coloration of holotype in preservative. The background color on the dorsum faded to dark grey; dorsalateral markings have become hardly visible; dark vertical bars, transverse bars and black spots distinct; the orange color on tubercles, glands, and elbow has faded to white-grey; ventral surfaces of body and limbs grey-white with dark pigmentations present at the edge of jaw and margin of belly, sparse pigmentations vaguely visible on the chin and chest; iris uniformly dark grey, the upper half and lower half are indistinguishable.
Measurements of holotype (in mm). SVL 31.1; HDL 10.8; HDW 10.5; SNT 4.2; EYE 4.2; IOD 3.4; TMP 1.7; TEY 1.1; TIB 14.5; ML 7.9; PL 13.1; PEC 1.4; FEM 1.3.
Comparison. Comparative morphological data of Leptobrachella wulingensis sp. nov., and 53 recognized Leptobrachella species occurring north of the Isthmus of Kra are listed in Table 3. Genetically, L. wulingensis sp. nov. is most closely related to L. alpina, L. bijie, L. bourreti, L. eos, L. oshanensis, L. purpuraventra, L. purpurus, and L. suiyangensis. The uncorrected 16S rRNA between L. wulingensis sp. nov. and above mentioned species are 3.6%–4.6% (L. alpina), 2.3%–2.9% (L. bourreti), 4.0% (L. eos), 3.9%–4.3% (L. oshanensis), 4.0% (L. purpuraventra), 4.3% (L. purpurus), and 4.3% (L. suiyangensis). Morphological characters can further distinguish them by the following:
L. wulingensis sp. nov. differs from the phylogenetically close congener, L. purpurus by having narrow lateral fringes on toes (vs. wide in purpurus), brown/reddish brown above in life (vs. purplish brown in purpurus), presence of longitudinal skin folds on dorsum (vs. absent in purpurus), supratympanic line weak (vs. distinct and black in purpurus), and dorsal tubercles orange in life (vs. reddish in purpurus).
L. wulingensis sp. nov. differs from the phylogenetically close congener, L. alpina by dorsum without white tiny flecks (vs. distinct white tiny flecks present on dorsum in alpina, see Yang et al., 2018: Fig. 6C), absence of distinct dark brown spots/blotches on belly (present in alpina, see Yang et al., 2018: Fig. 6D) in life, tibiotarsal articulation reaches to middle of the eye (vs. reaches to anterior corner of the eye in alpina), narrow lateral fringes on toes (vs. wide in males in alpina), and indistinct dorsolateral markings (vs. distinct in alpina).
L. wulingensis sp. nov. differs from the phylogenetically close congener, L. bourreti by having a relatively smaller body size (males SVL 24.5–32.8 mm vs. 28.0– 36.2 mm in bourreti; females SVL 29.9–38.5 mm vs. 41.8– 45.0 mm in bourreti), dermal ridges under toes distinct (vs. indistinct in bourreti), absence of dermal fringes on fingers (vs. present on Fingers II and III in bourreti), head longer than or as long as wide (vs. head wider than long in bourreti), indistinct dorsolateral markings (vs. distinct in bourreti), and indistinct supra-axillary glands in specimens reaching metamorphosis (see Fig. 6A) (vs. distinct in bourreti).
L. wulingensis sp. nov. differs from the phylogenetically close congener, L. eos by having narrow lateral fringes on toes (vs. wide in eos), presence of black spots on flanks (vs. absent in eos), large warts present on dorsal surface (vs. absent in eos), orange dorsal surface of elbow and upper arms in life (vs. reddish-brown in eos), and ventral surface creamy white with distinct/indistinct brown speckling in life (vs. throat with orange/yellow spots, chest and belly pearly-orange in eos).
L. wulingensis sp. nov. differs from the phylogenetically close congener, L. oshanensis by the presence of webbing and lateral fringes on toes (vs. absent in oshanensis), longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in oshanensis), large warts present on dorsal surface (vs. absent in oshanensis), supratympanic line weak (vs. distinct and black in oshanensis), pectoral gland lager than femoral gland (vs. reversed condition in oshanensis), femoral gland closer to knee than to vent (vs. reversed condition in oshanensis), and indistinct dorsolateral markings (vs. distinct in oshanensis).
L. wulingensis sp. nov. differs from the phylogenetically close congener, L. suiyangensis by having large warts present on dorsal surface (vs. absent in suiyangensis), brown/reddish brown above in life (vs. greyish brown in suiyangensis), supratympanic line weak (vs. deep and black in suiyangensis), indistinct dorsolateral markings (vs. distinct in suiyangensis), upper parts of iris bright orange/golden in life (vs. coppery in suiyangensis), and smaller tympanum (TMP/EYE ratio 0.32–0.52 vs. 0.42–1.06 in suiyangensis).
L. wulingensis sp. nov. differs from the phylogenetically close congener, L. purpuraventra and L. bijie by having large warts present on dorsal surface (vs. absent in purpuraventra and bijie), supratympanic line weak (vs. distinct and black in purpuraventra and bijie), ventral surface creamy white with/without brown speckling in life (vs. grey purple with nebulous greyish speckling in purpuraventra and white with nebulous greyish speckling in bijie), longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in purpuraventra and bijie), indistinct dorsolateral markings (vs. distinct in purpuraventra and bijie), and absence of tiny conical spines on surface of chest in males during breeding season (vs. present in purpuraventra and bijie).
Compared with the members from the L. applebyi species group. L. wulingensis sp. nov. differs by having a creamy white chest and belly in life (vs. reddish brown or greyish violet in all members, except rowleyae), presence of webbing and lateral fringes on toes (vs. absent in ardens, kalonensis, maculosa, pallida, rowleyae and tadungensis), bicolored iris (vs. uniform in ardens and tadunensis), distinct ventrolateral glandular line (vs. indistinct or absent in macrops, pyrrhops, rowleyae, tadungensis and melica), dorsal skin texture shagreened with sparse large warts (vs. mostly smooth in applebyi, bidoupensis, kalonensis, maculosa and melica), supratympanic line weak (vs. distinct and black in ardens, kalonensis, maculosa and tadungensis), upper parts of iris bright orange/golden in life (vs. reddish or copper in bidoupensis, maculosa and pallida), and lower parts of iris silver in life (vs. gold, golden green, or silver green in macrops, maculosa, pallida, pyrrhops and rowleyae); and further differs from L. applebyi by having a larger body size (males SVL 24.5–32.8 mm vs. 19.6–20.8 mm in applebyi) and the presence of dermal fringes on toes (vs. absent in applebyi); from L. bidoupensis by having a larger body size (males SVL 24.5–32.8 mm vs. 23.6–24.6 mm in bidoupensis) and the presence of glandular warts on flanks (vs. absent in bidoupensis); from L. macrops by the presence of longitudinal ridges on dorsum (vs. absent in macrops) and lateral fringes on toes (vs. absent in macrops); from L. melica by the absence of dermal fringes on fingers (vs. present in melica) and the presence of lateral fringes on toes (vs. absent in melica).
Compared with a congener occurring in Hunan province (L. mangshanensis), L. wulingensis sp. nov. differs by having narrow lateral fringes on toes (vs. weak in mangshanensis), distinct longitudinal ridges under toes interrupt- ed at the articulations (vs. longitudinal ridges indistinct and not interrupted at the articulations in mangshanensis), supratympanic line weak (vs. distinct and black in mangshanensis), tibiotarsal articulation reaches to middle of the eye (vs. reaches anterior margin of snout in mangshanensis), dorsal skin texture shagreened with sparse large warts (vs. mostly smooth in mangshanensis), and indistinct dorsolateral markings (vs. distinct in mangshanensis).
Compared with three congeners recently described by Chen et al. (2020) from Yunnan Province, L. wulingensis sp. nov. differs from L. niveimontis by having a creamy white chest and belly (vs. marbled with distinct irregular black speckling on bluish-white background in niveimontis), orange supra-axillary glands (vs. white in niveimontis), orange dorsal surface of elbow and upper arms (vs. reddish-brown in niveimontis) in life, and longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in niveimontis); from L. flaviglandulosa by having small white spines around cloacal region (vs. absent in flaviglandulosa), orange supra-axillary glands (vs. yellowish in flaviglandulosa), orange dorsal surface of elbow and upper arms (vs. yellowish in flaviglandulosa), greyish-pink ventral surface of throat and limbs (vs. dark brown in flaviglandulosa), and lacking yellow markings on dorsum in scapular region (vs. present in flaviglandulosa) in life; from L. feii by weak supratympanic line (vs. distinct and black in feii), absence of distinct black blotches scattered on chest and belly (vs. present in feii), longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in feii), small to moderate black spots on flanks (vs. large blotches in feii), and absence of nuptial spines on the dorsal surfaces of first and second fingers (vs. presence in males in feii).
For the other congeners known from China, L. wulingensis sp. nov. differs by having narrow lateral fringes on toes (vs. broad or wide in laui, liui, yingjiangensis and yunkaiensis; absent in nyx and ventripunctata), absence of lateral fringes on fingers (vs. present in laui and yingjiangensis), supratympanic line weak (vs. distinct and black in maoershanensis, shangsiensis, ventripunctata and wuhuangmontis), longitudinal ridges under toes interrupted at the articulations (vs. not interrupted in laui, maoershanensis, wuhuangmontis and yunkaiensis), indistinct dorsalateral markings (vs. distinct in liui, maoershanensis, pelodytoides, tengchongensis, ventripunctata, wuhuangmontis and yunkaiensis), absence of black spots or dark blotches on ventral surface (vs. present in maoershanensis, tengchongensis and ventripunctata), black spots on flanks small to moderate in size (vs. large in maoershanensis, pelodytoides and tengchongensis), distinct dermal ridges under toes (vs. indistinct in nyx and pelodytoides) and distinct ventrolateral glandular line (vs. indistinct in nyx and tengchongensis); and further differs from L. laui by the presence of longitudinal ridges on dorsum (vs. absent in laui); from L. liui by the absence of light dorsal markings in preservatives (vs. large, dark dorsal markings edged with light margins in liui, see Sung et al., 2014: Fig. 4B); from L. shangsiensis by having shagreened dorsal skin texture with sparse large warts (vs. smooth with numerous tiny tubercles in shangsiensis); from L. pelodytoides by having distinct dermal fringes under toes (vs. indistinct in pelodytoides); from L. tengchongensis by having bicolored iris (vs. uniform in tengchongensis); from L. ventripunctata by upper parts of iris bright orange/golden in life (vs. copper in ventripunctata) and lower parts of iris silver in life (vs. grey-brown in ventripunctata); from L. yingjiangensis by having orange supra-axillary glands in life (vs. brown in yingjiangensis) and the absence of white and dark brown mottling on ventral surface of thigh (vs. present in yingjiangensis); from L. yunkaiensis by having creamy white ventral surface (vs. pinkish in yunkaiensis) in life.
For the other 18 congeners occurring north of the Isthmus of Kra, L. wulingensis sp. nov. differs by having indistinct dorsalateral markings (vs. distinct in botsfordi, fuliginosa, khasiorum, lateralis, minima, nahangensis, nokrekensis, pluvialis, puhoatensis, sungi, tamdil and zhangyapingi), presence of black spots on flanks (vs. absent in aerea, botsfordi, firthi and tuberosa), creamy white ventral surface (vs. reddish brown or bright orange in botsfordi, puhoatensis and crocea) in life, dorsal skin texture shagreened with sparse large warts (vs. mostly smooth in fuliginosa, minima, nahangensis, pluvialis and zhangyapingi), presence of narrow lateral fringes on toes (vs. absent in crocea, minima, namdongensis and pluvialis; wide in males in firthi), supratympanic line weak (vs. distinct and black in khasiorum, lateralis, minima, nahangensis, namdongensis, nokrekensis and zhangyapingi), head longer or as long as wide (vs. head wider than long in crocea, firthi, khasiorum, lateralis, namdongensis, nokrekensis and tamdil), distinct ventrolateral glandular line (vs. indistinct or absent in crocea, petrops and sungi), distinct tympanum (vs. indistinct in crocea and tuberosa), bicolored iris (vs. uniform in aerea, nahangensis and sungi), upper parts of iris bright orange/golden in life (vs. reddish orange in fuliginosa and coppery in namdongensis) and lower parts of iris silver in life (vs. golden in fulignosa and namdongensis); and further differs from L. aerea by tibiotarsal articulation reaching to middle of the eye (vs. reaching to tip of snout in aerea); from L. botsfordi by having smaller femoral glands (0.7–1.4 mm, 2%–4% SVL vs. 2.4–4.3 mm, 7%–14% SVL in botsfordi); from L. firthi by the absence of dermal fringes on fingers (vs. most males with wide lateral dermal fringes on Finger II in firthi); from L. fuliginosa by the absence of brown network laterally on ventral surface (vs. present in fuliginosa); from L. khasiorum by having orange supra-axillary glands (vs. pinkish-red in khasiorum) in life; from L. minima by the canthus rostralis gently rounded (vs. angular in minima); from L. nahangensis by having smaller body size (males SVL 24.5–32.8 mm vs. 40.8 mm in nahangensis) and small to moderate spots on flanks (vs. large in nahangensis); from L. petrops by having rudimentary webbing on toes (vs. lacking in petrops); from L. pluvialis by having larger body size (males SVL 24.5–32.8 mm vs. 21.3–22.3 mm in pluvialis), relatively shorter hindlimbs (males TIB/SVL ratio 0.44–0.49 vs. 0.52–0.56 in pluvialis), tibiotarsal articulation reaches to middle of the eye (vs. reaches the nostril in pluvialis), creamy white ventral surface with distinct/indistinct brown speckling (vs. dirty white with dark brown marbling in pluvialis) in life, and presence of webbing on toes (vs. absent in pluvialis); from L. sungi by having smaller body size (males SVL 24.5–32.8 mm vs. 48.3–52.7 mm in sungi); from L. tuberosa by having a creamy white ventral surface with distinct/indistinct brown speckling (vs. white or pale creamy orange with blackish grey specks or streaks in tuberosa, see Rowley et al. 2010c: Fig. 2D) in life; from L. zhangyapingi by having smaller body size (males SVL 24.5–32.8 mm vs. 45.8–52.5 mm in zhangyapingi), ventral surface creamy white with distinct/indistinct brown speckling (vs. nearly immaculate white in zhangyapingi) in life, and relatively longer hindlimbs (males TIB/SVL ratio 0.44–0.49 vs. 0.41–0.43 in zhangyapingi).
Variation. Measurements of type series are shown in Table 4, and photographs of paratypes are shown in Figs. 4 and 5. Specimens show a large variation in body size. The smallest male measured 24.5 mm, whereas the largest male measured 32.8 mm. Size variation may be due to specimen age. Paratypes match most characters of the holotype, except males having indistinct slit-like paired vocal sac openings present posteriorly on floor of mouth. Specimens also varied in dorsal color, translucency of ventral surface, brightness of iris color, and size of black spots on flanks. Dorsal coloration varied from brown to reddish brown, except CSUFT 179 (adult male, Fig. 4A) being greenish brown in poor body condition. Individuals with larger sizes displayed less translucency on the white abdomen, and usually have distinct brown speckles on chest and lateral side of belly. The white abdomen in small individuals is usually translucent, and brown speckles are indistinct. For the bicolored iris, the upper part varies from bright orange to golden (Fig. 5), and a bit greenish in 1/5 upper part of CSUFT 203 (Fig. 5J). Due to the poor condition of CSUFT 179 (Fig. 5H), only the upper golden part is visible in the photo. The division of upper and lower part are unclear in CSUFT 200, CSUFT 204, CSUFT 202 and CSUFT 203 (Fig. 5F, G, I, and J), and also in the holotype CSUFT 201 (Fig. 3B), which are all from TNR populations. The division is clear in other specimens from TFP populations. More investigations are needed to determine if this is a subpopulation character or an effect of timing in which the specimens were photographed (we photographed individuals of the TFP population an hour after time of collection at night, while individuals of the TNR population were photographed during the day, 16 hours after time of collection). The black spots on flanks varied from small to moderate, and were indistinct in CSUFT 194. The pectoral glands were indistinct in all specimens except CSUFT 178 (subadult female), and became hardly visible in CSUFT 177, 193, 204 after preservation. Specimens collected in early August were smaller in body size compared to specimens collected in early September.
One gravid female (CSUFT 194) had large (diameter 1.60 mm), well-developed, yellowish-white oocytes in her ovaries; whereas the other two adult females (CSUFT 177 and 201) had smaller (0.95 mm and 0.37 mm respectively), less developed, yellowish-white oocytes. Females with elongate, translucent white ovaries, and lacking macroscopicly visible oocytes were considered subadults. Males with large testes and having vocal sac openings were considered as adults; others without vocal sac openings and having small, unformed testes were considered as subadults.
Distribution and habitat. Leptobrachella wulingensis sp. nov. is currently known from two localities in the Wuling Mountains. These are Tianquanshan Forest Park (TFP) and Tianzishan Nature Reserve (TNR), both in Zhangjiajie, Hunan Province, China. These two localities were separated by a straight line distance of approximately 27 km. All of the specimens were found at night in mixed forest between 414– 979 m. The new species was observed on rocks, stream banks, vegetation, or on soils adjacent to rocky streams. One female was gravid, indicating that the breeding season includes early September. No advertisement calls were detected and this may be because of the dry weather. A froglet and three adults were photographed in the TNR habitat (Fig. 6). We also found breeding adults of Megophrys tuberogranulatus Shen, Mo & Li (Mo et al. 2010) in sympatry in TNR during the August and September field surveys.
description: Published as part of Qian, Tian-Yu, Xia, Xin, Cao, Yue, Xiao, Neng-Wen & Yang, Dao-De, 2020, A new species of Leptobrachella (Anura: Megophryidae) Smith, 1925 from Wuling Mountains in Hunan Province, China, pp. 491-526 in Zootaxa 4816 (4) on pages 497-505, DOI: 10.11646/zootaxa.4816.4.4, http://zenodo.org/record/3954622