Boulenophrys yezhongensis, Liu & Feng & Shen & Li & Cheng & Wei & Wang & Su, 2025

Liu, Jing, Feng, Chao-Bo, Shen, Tuo, Li, Shi-Ze, Cheng, Yanlin, Wei, Gang, Wang, Bin & Su, Haijun, 2025, A new species of the genus Boulenophrys (Anura, Megophryidae) from Guizhou, China, Herpetozoa 38, pp. 117-136 : 117-136

publication ID	https://doi.org/10.3897/herpetozoa.38.e155430
publication LSID	lsid:zoobank.org:pub:9FDADB27-63C0-41C2-9A42-2E19DDB20F55
DOI	https://doi.org/10.5281/zenodo.15660084
persistent identifier	https://treatment.plazi.org/id/37E38F0A-0C88-50B2-89B9-2C44B39ED79D
treatment provided by	Herpetozoa by Pensoft
scientific name	Boulenophrys yezhongensis
status	sp. nov.

Treatment

Boulenophrys yezhongensis sp. nov.

Type material.

Holotype • CIB SC 20240531005 View Materials (Figs 4 View Figure 4 , 5 View Figure 5 ), adult male, from Yezhong Nature Reserve , Shuicheng County, Guizhou Province, China (26.163426°N, 104.880083°E, ca. 2022 m a. s. l.), collected by Jing Liu on 31 May 2024. GoogleMaps

Paratypes • Seven adult specimens (4 males and 3 females) from the same place as holotype. Four males ( CIB SC 20240531001 – 4) collected on 31 May 2024 by Shize Li and Chaobo Feng GoogleMaps . • Three females ( CIB SC 20240531006 –8) collected on 31 May 2024 by Jing Liu and Shize Li GoogleMaps .

Diagnosis.

Boulenophrys yezhongensis sp. nov. is assigned to the genus Boulenophrys based on molecular phylogenetic analyses and the following generic diagnostic characters: snout shield-like; projecting beyond the lower jaw; canthus rostralis distinct; chest glands small and round, closer to the axilla than to midventral line; femoral glands on rear part of thigh; vertical pupils; and the presence of subarticular tubercles at the base of each finger ( Fei et al. 2016; Lyu et al. 2023).

Boulenophrys yezhongensis sp. nov. can be distinguished from its congeners by a combination of the following morphological characters: body size moderate (SVL 41.2–46.2 mm in adult males and 51.8–58.6 mm in adult females); a small horn-like tubercle at edge of each upper eyelid; vomerine teeth absent; tongue not notched posteriorly; tympanum distinctly visible, round; two metacarpal tubercles on hand; toes with rudimentary webbing and narrow lateral fringes; heels overlapping when thighs are positioned at right angles to the body; tibiotarsal articulation reaching the level to the middle of eye when leg stretched forward; a single internal subgular vocal sac in adult males; in breeding males, nuptial pads with black nuptial spines on the dorsal bases of the first and second fingers.

Description of holotype.

CIB SC 20240531005 (Figs 4 View Figure 4 , 5 View Figure 5 ). SVL 44.4 mm; head width greater than head length (HDW / HDL ratio about 1.2); snout obtusely pointed, protruding well beyond the margin of the lower jaw in ventral view; loreal region vertical and concave; canthus rostralis well-developed; top of head flat in dorsal view; eye large, eye diameter 40.0 % of head length; pupils vertical; nostril orientated laterally, closer to snout than eye; tympanum distinct, TYP / EYE ratio 0.6; vomerine ridges present and vomerine teeth absent; margin of tongue smooth, not notched behind.

Forelimbs slender and comparatively short relative to those of congeneric species; length of lower arm and hand 44.4 % of SVL; fingers robust; relative finger lengths: II <IV <I <III; digit tips globular, without lateral fringes; distinct subarticular tubercles at finger bases; two prominent metacarpal tubercles, oval-shaped, the inner larger than the outer.

Hindlimbs slender (HLL / SVL 1.47); heels overlapping when thighs at right angles to body, tibiotarsal articulation reaching mid-eye when leg stretched forward; tibia longer than thigh; relative toe lengths: I <II <V <III <IV; toes round, slightly dilated; subarticular tubercles at toe base; toes with rudimentary webbing, narrow and barely visible lateral fringe; inner metatarsal tubercle oval-shaped; outer metatarsal tubercle absent.

Dorsal skin rough, with numerous brownish-yellow granules; several large warts scattered on flanks; tubercles on the dorsum forming a weak V-shaped ridge; two discontinuous dorsolateral parallel ridges on either side of the V-shaped ridges; tubercles on flanks, dorsal thighs, and tibias; supratympanic fold distinct; white granules with black hard spines on temporal fold region and adjacent areas.

Ventral surface smooth with numerous white granules; pectoral glands indistinct; femoral glands on rear of thighs; inner thighs and shanks with white granules, each tipped with single black hard spine.

Coloration of holotype in life.

(Fig. 4 View Figure 4 ). Inverted triangular brown speckle between eyes; dorsal V-shaped ridges dotted with brown speckles; dorsal surfaces of the thigh and shank characterized by four prominent dark brown transverse bands; upper and lower lips with alternating dark brown and white vertical bars; belly light orange-red; large white blotch and small grey blotch at center, small white blotches and large black patches on sides (from discontinuous line); posterior limb venters orange, thickly covered with numerous granules; palms, soles, and digit tips purple-grey; femoral glands pure white; iris copper-brown; throat and anterior chest light purple-brown.

Coloration of holotype in preservation.

(Fig. 5 View Figure 5 ). Dorsal surface faded to brown; inverted triangular brown speckle between eyes and V-shaped ridges on dorsum indistinct; throat and anterior chest light brownish-tan, abdomen pale orange-red with central white spot, lateral small white speckles interspersed with large black patches; ventral hindlimbs with white tubercles, males with black hardened spines.

Variation.

In CIB SC 20240531002 (Fig. 6 A View Figure 6 ): Overall color relatively dark; indistinct transverse stripes between thighs and shanks; black spines on white granules sparse, color not prominent. In CIB SC 20240531004 (Fig. 6 B View Figure 6 ): Dorsal parallel ridges in reticular pattern. In CIB SC 20240531006 (Fig. 6 C View Figure 6 ), tympanic membrane small, color indistinct; dorsal parallel ridges obscure, with only one ridge discernible. In CIB SC 20240531008 (Fig. 6 D View Figure 6 ): dorsum dark brownish; markings abundant and broad; wide transverse stripes on thighs and shanks; dorsal wart granules small, several large ones on flanks.

Secondary sexual characters.

Adult males (SVL 41.2–46.2 mm) are smaller than adult females (SVL 51.8–58.6 mm). Adult males have a single subgular vocal sac. In breeding males, there are dark black nuptial pads at the bases of the first and second fingers, and these pads are covered with fine, dense black nuptial spines. Moreover, white granules are present on the inner thighs, around the anus, the temporal fold region, and these granules are adorned with black hard spines. In contrast, females possess only white granules.

Comparisons.

(Suppl. material 4) Boulenophrys yezhongensis sp. nov. is phylogenetically closest to B. fanjingmontis , B. qianbeiensis , B. sangzhiensis , and B. spinata (Fig. 7 View Figure 7 ). The new species is easily distinguished from B. fanjingmontis by its slightly smaller body size: in adult males, the SVL is 41.2–46.2 mm (vs. 58.2 – 63.6 mm in the latter), and in adult females, the SVL is 51.8–58.6 mm (vs. 62.8–72.2 mm in the latter); tongue not notched posterior (vs. tongue feebly notched behind in the latter); having narrow lateral fringes on toes (vs. wide lateral fringes in the latter); breeding males have dark black nuptial pads at the bases of the first and second fingers, and these pads are covered with fine and dense black nuptial spines (vs. obvious black nuptial spines, spines on nuptial pads are larger and sparser in the latter); the values of morphological characteristics such as SVL, IND, IOD, LW, HLL, THL, TL, TW, TFL, FL, TYD, and IFE are significantly smaller in the new species than those in B. fanjingmontis (Table 3 View Table 3 ).

The new species differs from B. qianbeiensis by the following characters: having a slightly smaller body size, in adult males SVL 41.2–46.2 mm (vs. 49.3–58.2 mm in B. qianbeiensis ), in adult females SVL 51.8–58.6 mm (vs. 65.2 mm in B. qianbeiensis ); Breeding males have dark black nuptial pads at the bases of the first and second fingers, and these pads are covered with fine and dense black nuptial spines (vs. obvious black nuptial spines, spines on nuptial pads are larger and sparser in the latter); vomerine ridges separated and weak, vomerine teeth absent (vs. vomerine ridges and vomerine teeth present and distinct in the latter); tongue not notched posterior (vs. tongue feebly notched behind in the latter); toes with rudimentary webbing (vs. toes one quarter webbed in the latter); having narrow lateral fringes on toes (vs. wide lateral fringes in the latter); tibiotarsal articulation reaching the middle of eye when leg stretched forward (vs. reaching the level between tympanum and eye in the latter); the values of morphological characteristics such as SVL, HDW, NED, NSD, IND, IOD, UEW, LW, HLL, THL, TL, TW, TFL and FL are significant smaller in the new species than those in B. qianbeiensis (Table 3 View Table 3 ).

The new species differs from B. sangzhiensis by the following characters: having a slightly smaller body size, in adult males SVL 41.2–46.2 mm (vs. 55.7–62.2 mm in the latter), in adult females SVL 51.8–58.6 mm (vs. 62.2–76.3 mm in the latter); breeding males have dark black nuptial pads at the bases of the first and second fingers, and these pads are covered with fine and dense black nuptial spines (vs. obvious black nuptial spines, spines on nuptial pads are larger and sparser in the latter); tongue not notched posterior (vs. tongue feebly notched behind in the latter); the values of morphological characteristics such as SVL, HDW, NSD, IND, IOD, LW, HLL, THL, TL, TW, TFL, FL, TYD and IFE are significant smaller in the new species than those in B. sangzhiensis (Table 3 View Table 3 ).

The new species differs from B. spinata by the following characters: having a slightly smaller body size, in adult males SVL 41.2–46.2 mm (vs. 47.2–54.4 mm in the latter), in adult females the SVL is not significant different; tongue not notched posterior (vs. tongue feebly notched behind in the latter); toes with rudimentary webbing (vs. toes one quarter webbed in the latter); having narrow lateral fringes on toes (vs. wide lateral fringes in the latter); breeding males have dark black nuptial pads at the base of the first and second fingers, and these pads are covered with fine and dense black nuptial spines (vs. B. spinata has obvious black nuptial spines, and the spines on its nuptial pads are larger and sparser); the values of morphological characteristics such as SVL, TYD, IAE, NED, NSD, IND, IOD, LW, HLL, THL, TL, TW, TFL and FL are significant smaller in the new species than those in B. spinata (Table 3 View Table 3 ).

By having medium body size, Boulenophrys yezhongensis sp. nov. differs from B. acuta , B. angka , B. baishanzuensis , B. binchuanensis , B. boettgeri , B. cheni , B. congjiangensis , B. daiyunensis , B. daoji , B. daweimontis , B. elongata , B. frigida , B. gaolanensis , B. hungtai , B. jinggangensis , B. jiulianensis , B. kuatunensis , B. lichun , B. lishuiensis , B. lushuiensis , B. mufumontana , B. nankunensis , B. obesa , B. ombrophila , B. pepe , B. puningensis , B. rubrimera , B. sanmingensis , B. shimentaina , B. shunhuangensis , B. tongboensis , B. wugongensis , B. wuliangshanensis , B. wushanensis , B. xianjuensis , B. yangmingensis and B. yingdeensis (minimum SVL> 41.0 mm in the new species vs. maximum SVL <37.0 mm in the latter), and differs from B. caudoprocta , B. fanjingmontis , B. jingdongensis , B. liboensis , B. mirabilis , B. omeimontis , B. qianbeiensis , B. sangzhiensis and B. shuichengensis (maximum SVL <46.0 mm in the new species vs. minimum SVL> 50.0 mm in the latter).

By vomerine teeth being absent, Boulenophrys yezhongensis sp. nov. differs from B. brachykolos , B. caudoprocta , B. daiyunensis , B. daweimontis , B. dongguanensis , B. elongata , B. fansipanensis , B. fengshunensis , B. frigida , B. hoanglienensis , B. insularis , B. jingdongensis , B. jinggangensis , B. jiulianensis , B. liboensis , B. nankunensis , B. nanlingensis , B. omeimontis , B. palpebralespinosa , B. puningensis , B. qianbeiensis , B. rubrimera , B. shimentaina , B. tongboensis , and B. yingdeensis (vs. present in the latter).

By the vomerine ridges present, Boulenophrys yezhongensis sp. nov. differs from B. baolongensis , B. binchuanensis , B. boettgeri , B. chishuiensis , B. congjiangensis , B. daoji , B. dupanglingensis , B. gaolanensis , B. hengshanensis , B. hungtai , B. leishanensis , B. lichun , B. lishuiensis , B. minor , B. mirabilis , B. ombrophila , B. sanmingensis , B. shunhuangensis , B. tuberogranulatus , B. wuliangshanensis , B. wushanensis , and B. xuefengmontis (vs. absent in the latter).

By a small horn-like tubercle present at the edge of each upper eyelid, Boulenophrys yezhongensis sp. nov. differs from B. acuta , B. caudoprocta , B. jinggangensis , B. liboensis , B. mirabilis , and B. palpebralespinosa (vs. having a prominent and elongated tubercle in the latter).

By tongue not notched posteriorly, Boulenophrys yezhongensis sp. nov. differs from B. baolongensis , B. binlingensis , B. boettgeri , B. cheni , B. fanjingmontis , B. insularis , B. jingdongensis , B. jiulianensis , B. kuatunensis , B. liboensis , B. lushuiensis , B. minor , B. nanlingensis , B. omeimontis , B. pepe , B. qianbeiensis , B. sangzhiensis , B. sanmingensis , B. shuichengensis , B. spinata , and B. tongboensis (vs. tongue feebly notched behind in the latter).

By toes with narrow lateral fringes, Boulenophrys yezhongensis sp. nov. differs from B. angka , B. baolongensis , B. brachykolos , B. caobangensis , B. chishuiensis , B. daweimontis , B. dongguanensis , B. dupanglingensis , B. fansipanensis , B. fengshunensis , B. frigida , B. gaolanensis , B. hengshanensis , B. hoanglienensis , B. hungtai , B. insularis , B. jiangi , B. jiulianensis , B. leishanensis , B. lichun , B. lishuiensis , B. major , B. nankunensis , B. obesa , B. ombrophila , B. pepe , B. puningensis , B. shunhuangensis , B. tongboensis , B. tuberogranulata , B. wugongensis , B. wuliangshanensis , B. xuefengmontis , B. yaoshanensis , B. yingdeensis , and B. yunkaiensis (vs. lacking lateral fringes on toes). differs from B. binchuanensis , B. boettgeri , B. cheni , B. fanjingmontis , B. jingdongensis , B. liboensis , B. lini , B. palpebralespinosa , B. qianbeiensis , B. rubrimera , B. sanmingensis , B. shuichengensis , B. spinata , and B. xiangnanensis (vs. having lateral wide fringes on toes in the latter).

By toes with rudimentary webbing, Boulenophrys yezhongensis sp. nov. differs from B. baishanzuensis , B. baolongensis , B. daweimontis , B. elongata , B. fansipanensis , B. frigida , B. gaolanensis , B. hengshanensis , B. hungtai , B. kuatunensis , B. lichun , B. lishuiensis , B. ombrophila , B. pepe , B. rubrimera , B. shunhuangensis , B. tongboensis , B. wuliangshanensis , and B. xuefengmontis (vs. toes lacking webbing in the latter) and differs from B. jingdongensis , B. palpebralespinosa , B. qianbeiensis , B. shuichengensis , B. spinata , and B. wushanensis (vs. toes with one-fourth in the latter).

By heels overlapping when thighs are positioned at right angles to the body, Boulenophrys yezhongensis sp. nov. differs from B. acuta , B. brachykolos , B. daoji , B. dongguanensis , B. fengshunensis , B. gaolanensis , B. hengshanensis , B. hungtai , B. insularis , B. kuatunensis , B. lichun , B. nankunensis , B. obesa , B. ombrophila , B. pepe , B. puningensis , B. shuichengensis , and B. wugongensis (vs. not meeting in the latter). differs from B. binchuanensis , B. elongata , B. minor , B. xiangnanensis , and B. xuefengmontis (vs. just meeting in the latter).

By tibiotarsal articulation reaching to the level to the middle of eye when leg stretched forward, Boulenophrys yezhongensis sp. nov. differs from B. cheni , B. congjiangensis , B. daweimontis , B. lini , B. obesa , B. shunhuangensis and B. yangmingensis (vs. reaching the anterior corner of the eye or beyond eye or nostril and tip of snout in the latter); and differs from B. brachykolos , B. chishuiensis , B. daoji , B. dongguanensis , B. fengshunensis , B. hungtai , B. insularis , B. jiangi , B. kuatunensis , B. lishuiensis , B. mufumontana , B. nankunensis , B. puningensis , B. qianbeiensis , B. shimentaina , B. xiangnanensis , B. xuefengmontis , B. yingdeensis and B. yunkaiensis (vs. reaching the level between tympanum and eye or reach the tympanum or be even closer in the latter).

By having an internal single subgular vocal sac in males, Boulenophrys yezhongensis sp. nov. differs from B. caudoprocta and B. shuichengensis (vs. vocal sac absent in the latter).

Distribution and habitats.

Boulenophrys yezhongensis sp. nov. is known from the type locality, Yezhong Nature Reserve, Shuicheng County, Guizhou Province, China, at elevations of approximately 2022 m. The individuals of the new species were frequently found on stones in the streams or gorges surrounded by evergreen broadleaved forest (Fig. 8 View Figure 8 ), and two sympatric amphibian species, i. e., Rana chaochiaoensis and Quasipaa boulengeri , were found.

Etymology.

The specific epithet yezhongensis refers to the type locality of the species. We propose the common English name “ Yezhong Horned Toad ” and the Chinese name “ Yě Zhōng Jiǎo Chán (野钟角蟾) ” for this species.