Boulenophrys nebulosa Shen & Li, 2025

Shen, Deming, Zhou, Haiying, Wang, Chaoyang, Xu, Jun, Xu, Shi, Zhong, Yifeng, Wang, Zhenyu, Cai, Hao, Wu, Yahui, Wei, Meicai & Li, Yankuo, 2025, A new species of Boulenophrys (Anura, Megophryidae) from northern Jiangxi, China, Zoosystematics and Evolution 101 (4), pp. 2251-2266 : 2251-2266

Boulenophrys nebulosa Shen & Li sp. nov.

Fig. 3 , Table 2

Type materials.

Holotype. • JXNU 21042833 (Fig. 3 A – F ), adult male, collected by Deming Shen, Haiying Zhou, Yifeng Zhong, Chaoyang Wang, Hao Cai, Jun Xu and Shi Xu on 28 April 2021 from Mt. Lushan ( 29°32.56'N, 115°57.58'E; ca. 970 m a. s. l.), Lushan City, Jiujiang, Jiangxi, China .

Paratypes. ( n = 6) • JXNU 21042832 (Fig. 4 A – C ), JXNU 21042834 , adult males, and JXNU 21042810 , adult female, collected by Deming Shen, Haiying Zhou, Yifeng Zhong, Chaoyang Wang, Hao Cai, Jun Xu and Shi Xu on 28 April 2021 from Mt. Lushan ( 29°31.87'N, 115°55.87'E; ca. 660 m a. s. l.) ; • JXNU 21061501 (Fig. 4 D – F ), JXNU 21061502 , JXNU 21061504 , adult females, collected by same collectors on 15 June 2021 from the type locality ( 29°32.56'N, 115°57.58'E; ca. 970 m a. s. l.) .

Other material examined.

• One tadpole ( JXNU 21110702 ; Fig. 5 ) collected by Haiying Zhou on 2 November 2021 .

Etymology.

The specific epithet nebulosa (feminine of Latin nebulosus) refers to the frequently mist-shrouded montane forests above 600 m on Mt. Lushan, Jiangxi Province, China. This name not only reflects the species’ habitat preference but also alludes to the region’s famous “Cloud & Fog Tea” (云雾茶, yún wù chá) speciality. The English vernacular name “Yunwu Horned Toad” is here proposed, with the corresponding Chinese name “云雾角蟾” (Yúnwù Jiǎochán).

Diagnosis.

This species can be distinguished from its congeners by the combination of following morphological characteristics: (1) adult males, SVL 31.8–32.3 mm (n = 3); adult females, SVL 35.0– 37.3 mm (n = 4); (2) canthus rostralis well developed, tongue not notched posteriorly; (3) tympanum distinct; (4) vomerine ridges weak; vomerine teeth present; (5) outer margin of the upper eyelid bears a small, prominent horn-like tubercle; supratympanic fold distinct, narrow, curving posteroventrally to above the arm; (6) relative finger lengths I <II <IV <III; indistinct subarticular tubercle on base of each finger; (7) heels meeting when hindlimbs folded; tibiotarsal articulation reaching center of eye when leg stretched forward; (8) toes with narrow fringes and only rudimentary webbing; (9) rough dorsal skin, a V-shaped, X-shaped or Y-shaped ridge is present on central back of trunk, dorsal surface bears short, irregular and weak ridges, with scattered tubercles and granules, dorsolateral ridges absent; several tubercles on flanks and dorsal hindlimbs; ventral skin smooth with granules; (10) dorsal surface yellowish brown; hollow brown triangular marking between eyes; brown V-shaped, X-shaped, or Y-shaped marking on central back of trunk; dorsal limbs and digits with dark brown transverse bands; and (11) dense nuptial spines on dorsal bases of fingers I and II in breeding adult males, subgular vocal sac present in males.

Description of holotype.

JXNU 21042833 , adult male, body size small, SVL 32.2 mm; head width slightly smaller than head length, HDW / HDL 0.91; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes large, ED 0.35 of HDL, pupil vertical, ovoid in natural state; nostril obliquely ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance larger than interorbital distance; tympanic region oblique, tympanum moderate in size, margin clear, upper margin in contact with supratympanic fold, TD / ED 0.41; large ovoid choanae at base of maxilla; vomerine ridge weak, vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched distally; presence of subgular vocal sac.

Forearm length 0.24 of SVL, hand 0.22 of SVL; webbing absent between fingers, lateral fringes absent, relative finger length I <II <IV <III; tips of fingers slightly dilated, round; indistinct subarticular tubercles present; inner metacarpal tubercle observably enlarged, outer one indistinct; single nuptial pad bearing nuptial spines present on dorsal surface of first and second fingers, respectively.

Hindlimbs short, heels meeting when flexed hindlimbs are held at right angles to the body axis; tibiotarsal articulation reaching the center of the eye when the hindlimb is stretched forward along the body; tibiofibular length 0.45 of SVL, foot length 0.39 of SVL; relative toe length I <II <V <III <IV; tips of toes rounded and slightly dilated; toes with narrow lateral fringes and rudimentary webbing; subarticular tubercles indistinct at the base of each toe; inner metatarsal tubercle long and ovoid; outer metatarsal tubercle absent.

Dorsal skin smooth with granules; a discontinuous Y-shaped ridge present on the central dorsum of trunk, discontinuous longitudinal ridges present laterally; several rounded tubercles present on the flanks and dorsal surfaces of the hindlimbs; a prominent, small horn-like tubercle located on the upper eyelid; supratympanic fold distinct, extending from the posterior corner of the eye to above the insertion of the forelimb; tympanum distinct; ventral skin smooth to weakly granular, with spinose tubercles surrounding the cloacal region; pectoral gland small and distinct, located closer to the axilla; a single large femoral gland present on the posterior surface of the thigh, situated approximately midway between the knee and the cloaca.

Coloration of holotype.

In life, dorsum yellowish brown, with a hollow brown triangular interorbital marking between eyes and a distinct Y-shaped brown marking at the center of the dorsum, surrounded by indistinct brown netlike pattern; dorsal surfaces of limbs yellowish brown, each bearing several brown transverse bars; axillary region and adjacent lateral flank light yellowish brown; tubercles on the upper eyelids orange; a vertical brown stripe present below the eye, extending from the lower margin of the eye to the upper lip; ventral surface of throat and chest greyish brown, bearing irregular dark brown and white blotches; ventral surfaces of limbs, manus and pes purple, with white blotches; abdominal surface greyish white, scattered with creamy white and orange spots; inner and outer metatarsal tubercles and inner metacarpal tubercle pale purple; pectoral white, digits and femoral glands greyish white; iris light yellowish brown.

In preservative, the dorsum fades from yellowish brown to greyish, while brown markings, including a Y-shaped interscapular marking, a triangular interorbital marking and transverse bands on the dorsal surfaces of the forelimbs and hindlimbs, darken to greyish brown and become more conspicuous. The ventral surfaces fade to greyish white, making the original white spots and blotches more prominent.

Variation.

Morphometric data are given in Table 2 . All paratypes conform to the holotype in key diagnostic characters but differ in dorsal patterning, ranging from specimens with an indistinct brown netlike pattern as in the holotype to those bearing only an X-shaped interscapular marking and a triangular interorbital marking (e. g., JXNU 21042832 ). Female paratypes lack nuptial pads and keratinized spines, which are present in males.

Tadpole description.

The tadpole was confirmed as Boulenophrys nebulosa sp. nov. by molecular phylogenetic analyses. The following tadpole description is based on the specimen JXNU 21110702 at Gosner Stage 29 (Fig. 5 ). Body depressed and longitudinally oval; TOL 28.0 mm; body width longer than body height, BW / BH 1.3; eyes dorsolateral, pupils round; nostril near eyes; spiracle on the left side of the body and distinct; tail approximately 2.3 times as long as snout-vent length; tail height greater than body height, TAH / BH 1.2; dorsal fin arising just behind tail base, highest near mid-length, tapering gradually to bluntly pointed tip; oral disk terminal; lips expanded, directed upward, forming umbelliform structure; lateral corners of oral disc fold medially, turning upward when not fully extended.

In preservative, the background color of the head and trunk fades from brown to dark brown, interspersed with dark-brown chromatophores, and becomes progressively paler toward the tail tip. Areas lacking white pigmentation contain fewer melanocytes. The belly bears dense white spots, especially concentrated on the chest, while the lateral surfaces of the tail are covered with yellowish-brown spots. BH 2.7; BW 3.6; IOD 2.5; MW 1.6; SL 1.8; SS 3.2; SVL 8.5; TAH 3.3; TAL 19.5; TBW 1.4; TOL 28.0.

Advertisement call.

Call descriptions are based on the vocalizations of the male holotype JXNU 21042833 . Spectrograms and waveforms are shown in Fig. 6 . A total of 13 call groups were analyzed. Each call had an average duration of 120.3 ± 7.4 ms (110.3–135.6 ms) and an average inter-call interval of 487.6 ± 48.8 ms (415.9–573.4 ms). Calls were repeated at a mean rate of 2.05 calls / s (constant repetition rate). Each call consisted of 31.7 ± 2.1 pulses (28–35 pulses). The dominant frequency averaged 3.70 ± 0.08 kHz (3.59–3.84 kHz).

Comparisons.

Morphological comparisons between the proposed new species and other recognized species of the genus Boulenophrys are provided (Table 3 ). Boulenophrys nebulosa sp. nov. is phylogenetically most closely related to B. mufumontana , forming a well-supported sister clade. However, the new species can be readily distinguished from B. mufumontana by the following combination of characters: 1) relative finger lengths I <II <IV <III (vs. II = IV <I <III); 2) indistinct subarticular tubercle on base of each finger (vs. distinct subarticular tubercle); 3) larger male body size, SVL 31.8–32.3 mm (vs. 30.1–30.8 mm); 4) heels just meeting when hindlimbs folded (vs. heels overlapping); 5) tibiotarsal articulation reaching the center of eye when leg stretched forward (vs. reaching region between posterior margin of tympanum and center of eye); 6) vomerine teeth present (vs. vomerine teeth absent).

The new species differs from B. boettgeri by the following combination of characters: 1) smaller male body size, SVL 31.8–32.3 mm (vs. 34.5–37.8 mm); 2) vomerine teeth present (vs. vomerine teeth absent); 3) tongue not notched (vs. tongue notched); 4) fringes on toes narrow (vs. fringes on toes wide); 5) indistinct subarticular tubercle on base of each finger (vs. distinct subarticular tubercle).

Additionally, Boulenophrys nebulosa sp. nov. can be distinguished from the following congeners by its heels just meeting when flexed hindlimbs held at right angles to body axis: B. anlongensis , B. baishanzuensis , B. binlingensis , B. congjiangensis , B. changyangensis , B. cheni , B. chishuiensis , B. dalaolingensis , B. dupanglingensis , B. jiangi , B. jingdongensis , B. jinggangensis , B. jiulianensis , B. leishanensis , B. liboensis , B. lini , B. lushanensis , B. mirabilis , B. nanlingensis , B. omeimontis , B. palpebralespinosa , B. qianbeiensis , B. sangzhiensis , B. sanmingensis , B. shimentaina , B. shunhuangensis , B. spinata , B. tongboensis , B. wuliangshanensis , B. xianjuensis , B. yangmingensis , and B. yezhongensis (vs. heels overlapping); and from B. acuta , B. brachykolos , B. daoji , B. daxuemontis , B. dongguanensis , B. fengshunensis , B. gutu , B. gaolanensis , B. hengshanensis , B. hungtai , B. insularis , B. lichun , B. nankunensis , B. obesa , B. ombrophila , B. pepe , B. puningensis , B. wugongensis , and B. wushanensis (vs. heels not meeting).

Boulenophrys nebulosa sp. nov. can be distinguished from the following congeners by its fringes on toes narrow: B. angka , B. baolongensis , B. caobangensis , B. daweimontis , B. fansipanensis , B. frigida , B. hoanglienensis , B. lishuiensis , B. minor , B. tuberogranulatus , B. xuefengmontis , B. yaoshanensis , B. yingdeensis , B. yunkaiensis , and B. huangniushiensis (vs. fringes on toes absent); and from B. binchuanensis , B. fanjingmontis , B. shuichengensis , and B. xiangnanensis (vs. fringes on toes wide).

For the remaining congeners, Boulenophrys nebulosa sp. nov. can be distinguished from B. elongata and B. rubrimera by its rudimentary webbing on toes (vs. lacking webbing); from B. caudoprocta by its horn-like tubercle small at upper eyelid (vs. elongated as triangular appendage); from B. daiyunensis by its vomerine teeth present (vs. vomerine teeth absent); and from B. lushuiensis by its tongue not notched (vs. tongue notched).

Distribution and habits.

Boulenophrys nebulosa sp. nov. is currently known only from its type locality, Mt. Lushan, Jiujiang City, Jiangxi Province, China. The species occurs at elevations of 600–1,200 m, primarily in mixed coniferous and broadleaved forests (Fig. 7 ). Individuals are typically found on the forest floor or under leaf litter near montane streams, as well as along moist roadside drainage channels. Calling males were observed from May to July, all of which were observed calling beneath accumulations of coniferous leaf litter. Females were occasionally observed in drainage ditches lacking flowing water. Tadpoles are observed throughout the year and coexist with the tadpoles of Quasipaa spinosa and B. lushanensis in some aquatic habitats.

Figures

Figure 3. Holotype of Boulenophrys nebulosa sp. nov. (JXNU 21042833) in life: A. Lateral view; B. Dorsal view; C. Ventral view; D. Volar and dorsal views of left hand; E. Plantar view of left foot; F. Edge of tongue. Photos by Deming Shen.

Figure 4. Paratypes of Boulenophrys nebulosa sp. nov. in life. Male paratype (JXNU 21042832): A. Lateral view; B. Dorsal view; C. Ventral view. Female paratype (JXNU 21061501): D. Lateral view; E. Dorsal view; F. Ventral view. Photos by Deming Shen.

Figure 5. Tadpole specimen of Boulenophrys nebulosa sp. nov. (JXNU 21110702) in preservative: A. Lateral view; B. Ventral view; C. Dorsal view. Photos by Deming Shen.

Figure 6. Advertisement call of Boulenophrys nebulosa sp. nov. male holotype (JXNU 21042833): A. Waveform showing five call groups; B. Spectrogram corresponding to the waveform.

Figure 7. Habitat of Boulenophrys nebulosa sp. nov. A. Landscape of mixed coniferous and broadleaved forests at the type locality; B. Microhabitat showing female paratype (JXNU 21061502) perching on needle litter (white box). Photos by Deming Shen.