Boulenophrys puningensis, Wang & Zeng & Lyu & Qi & Liu & Chen & Lu & Xiao & Lin & Chen & Wang, 2022

Wang, Jian, Zeng, Zhao-Chi, Lyu, Zhi-Tong, Qi, Shuo, Liu, Zu-Yao, Chen, Hong-Hui, Lu, Yu-Hong, Xiao, Hui-Wen, Lin, Can-Rong, Chen, Kai & Wang, Ying-Yong, 2022, Description of three new Boulenophrys species from eastern Guangdong, China, emphasizing the urgency of ecological conservation in this region (Anura, Megophryidae), Zootaxa 5099 (1), pp. 91-119 : 102-106

Boulenophrys puningensis sp. nov. Wang, Zeng, Lyu, Xiao & Wang

Puning Horned Toad (in English) / Pǔ Níng Jiǎo Chán (DṪDzdz in Chinese)

Figures 3–4

Holotype. SYS a005770, adult male, collected by Jian Wang on 24April 2017 from Longkeng Village (23°7'54.07"N, 115°51'5.28"E; ca. 120 m a.s.l.), Daping Town , Puning, Jieyang, Guangdong, China.

Paratypes (N=5). Adult male, SYS a006755/ CIB118526 , collected by Jian Wang, Can-Rong Lin and Hui-Wen Xiao on 14 February 2018; adult males SYS a007649, 7650 and adult females SYS a007647, 7648, collected by Jian Wang, Can-Rong Lin and Hui-Wen Xiao on 18 March 2019, all from the same stream as the holotype at elevations between 250–300 m.

Etymology. The specific epithet “ puningensis ” refers to the type locality of the new species in Puning. Three of the authors of this work (Jian Wang, Hui-Wen Xiao and Can-Rong Lin) chose this nomen in honor of their hometown.

Diagnosis. (1) Small body size, SVL 31.7–34.6 mm (33.0 ± 1.3, N = 4) in adult males and SVL 37.8–38.3 mm (N = 2) in adult females; (2) snout rounded in dorsal view; (3) tympanum large, TD/ED 0.68–0.71; (4) tympanic region smooth without granules or tubercles; (5) vomerine ridge and vomerine teeth present; (6) margin of tongue rounded, not notched distally; (7) hindlimbs short, heels not meeting and tibio-tarsal articulation reaching forward to the region between tympanum and eye; (8) a subarticular tubercle present at the base of each fingers; (9) toes without lateral fringes and with rudiment of webbing; (10) distinct enlarged tubercles on the surface of limbs, flanks, chest, belly and around the cloaca; (11) tips of the enlarged tubercles on posterior abdomen, ventral thighs and around the cloaca bearing tiny spines; (12) single subgular vocal sac in males; (13) nuptial pads with villiform black nuptial spines on the dorsal surface of the first and second fingers in breeding males.

Comparisons. Comparative data of Boulenophrys puningensis sp. nov. and the other recognized members of Boulenophrys are listed in Table 3 .

With a smaller body size, SVL 31.7–34.6 mm in adult males and SVL 37.8–38.3 mm in adult females, Boulenophrys puningensis sp. nov. differs from the eight congeners whose SVL ≥ 50 mm in adult males or females, including B. caudoprocta (81.3 mm in a single male), B. jingdongensis (53.0– 56.5 mm in males and 63.5 mm in a single female), B. liboensis (60.5–67.7 mm in males and 60.8–70.6 in females), B. mirabilis (55.8–61.4 mm in males and 68.5–74.8 mm in females), B. omeimontis (56.0– 59.5 mm in males and 68.0– 72.5 mm in females), B. sangzhiensis (54.7 mm in a single male), B. shuichengensis (102.0– 118.3 mm in males and 99.8–115.6 mm in females), and B. spinata (54.0–55.0 mm in females).

Boulenophrys puningensis sp. nov. shows the least genetic divergence from B. kuatunensis (mean p -distances 5.3 % in the 16S gene) and B. daiyunensis (mean p -distances 6.2 % in the 16S gene). However, the new species distinctively differs from these species by having relatively shorter shanks with the heels not meeting when the flexed hindlimbs are held at right angles to the body axis (vs. heels meeting or overlapping in B. daiyunensis ); having rudiment of webbing and no lateral fringes on toes (vs. lateral fringes narrow in B. daiyunensis ; webbing absent in B. kuatunensis ); having raised and enlarged tubercles with spines on their tips on surface of posterior abdomen, ventral thighs and around the cloaca (vs. such tubercles not enlarged and without spines in B. daiyunensis ; ventral surface smooth in B. kuatunensis ).

Boulenophrys puningensis sp. nov. is morphologically most similar to B. brachykolos , which is restricted to Hong Kong and Shenzhen, China ( Liu et al. 2018). The new species differs from B. brachykolos by having vomerine teeth (vs. absent in B. brachykolos ); lacking spines on the surface of the tympanic region (vs. having dense tiny spines on the surface of the tympanic region in B. brachykolos ); and having different relative finger length formula (I = II <IV <III in Banophrys puningensis sp. nov. vs. II <IV <I <III in B. brachykolos ).

In having relatively shorter shanks with heels that do not meet when the flexed hind limbs are held at right angles to the body axis, Boulenophrys puningensis sp. nov. can be easily distinguished from the following 32 congeners, i.e. B. angka , B. anlongensis , B. baishanzuensis , B. baolongensis , B. binchuanensis , B. binlingensis , B. boettgeri , B. congjiangensis , B. cheni , B. chishuiensis , B. jiangi , B. jinggangensis , B. jiulianensis , B. leishanensis , B. lini , B. minor , B. mufumontana , B. nanlingensis , B. palpebralespinosa , B. qianbeiensis , B. sanmingensis , B. shimentaina , B. shunhuangensis , B. tongboensis , B. wuliangshanensis , B. wushanensis , B. xiangnanensis , B. xianjuensis , B. yaoshanensis , B. yangmingensis , B. yingdeensis and B. yunkaiensis , all of which have relatively longer shanks with the heels meeting or overlapping.

By the presence of vomerine teeth, Boulenophrys puningensis sp. nov. differs from B. acuta , B. caobangensis , B. daoji , B. huangshanensis , B. lishuiensis , B. lushuiensis , B. obesa , B. ombrophila , B. tuberogranulatus , and B. wugongensis , all of which lack vomerine teeth. By having a rounded tongue margin that is not notched distally, Boulenophrys puningensis sp. nov. differs from B. hoanglienensis , and B. insularis , all of which have notched tongues. By the absence of lateral fringes on toes, Boulenophrys puningensis sp. nov. differs from B. rubrimera , which has narrow lateral fringes on toes. By the presence of rudimentary webbing on the toes, Boulenophrys puningensis sp. nov. differs from B. daweimontis , B. fansipanensis , and B. frigida , all of which lack webbing on the toes. Boulenophrys puningensis sp. nov. further differs from the remaining B. dongguanensis and B. nankunensis by having raised tubercles bearing spines on the surface of the posterior abdomen, ventral thighs, and around the cloaca (vs. absence of such tubercles and spines in B. dongguanensis and B. nankunensis ).

Description of holotype. Adult male. Body size small, SVL 34.6 mm. Head width slightly larger than head length, HWD/HDL 1.03; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes moderate in size, ED 0.36 of HDL, pupil vertical, near diamond-shaped; nostril oblique-ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanum large with an obvious margin, TD/ED 0.70; large ovoid choanae at base of maxilla; vomerine ridge and vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched distally; presence of single subgular vocal sac, and pair of slit-like openings at posterior of jaw.

Forearm (i.e., radioulna) length 0.21 of SVL and hand 0.23 of SVL; hand without webbing, fingers without lateral fringes, relative finger length I = II <IV <III; tips of fingers slightly dilated, round; subarticular tubercles present and distinct; inner metacarpal tubercle observably enlarged and outer one slightly smaller; single nuptial pad bearing nuptial spines present on dorsal surface of first and second fingers, respectively. Hindlimbs short, tibiotarsal articulation reaching forward to region between tympanum and eye when hindlimb stretched along body; heels not meeting when flexed hindlimbs held at right angles to body axis; crus (i.e., tibiofibular) length 0.38 of SVL and foot length 0.54 of SVL; relative toe length I <II <V <III <IV; tips of toes round and slightly dilated; toes without lateral fringes and with rudiment of webbing; subarticular present and distinct; inner metatarsal tubercle long ovoid and lacking outer metatarsal tubercle.

Dorsal skin rough and granular, with raised conical tubercles; sparse large tubercles on flanks; single horn-like prominent tubercle on edge of upper eyelid; obvious supratympanic fold curving posteroventrally from posterior corner of eye to level above insertion of arm; tympanic region smooth without granules or tubercles; dense tubercles on skin of upper lip, upper eyelid, mandibular articulation, loreal, temporal region excluding tympanum and surface around cloaca; a single discontinuous “V” shaped ridge present on occipital region; dense tubercles on shanks and thighs; ventral surface with dense raised tubercles; tubercles on surface of posterior abdomen, ventral surface of thighs and around cloaca bearing tiny spines on their tips; small pectoral gland closer to axilla; single femoral gland positioned on posterior surface of thigh at midpoint between knee and cloaca.

Coloration of holotype in life. Dorsal surface of body yellowish brown, with incomplete dark brown triangular marking between eyes. Two wide oblique black bands present on forearm. Dorsal surface of fingers and hindlimbs with dark grey transverse bands. Presence of vertical dark brown band below eye. Tubercles on edge of upper eyelid beige. Supratympanic fold light brown. Ventral surface dark grey, with black longitudinal band on surface of throat; surface of throat and chest mottled with orange patches. Tubercles on ventral surface of chest, belly, and thighs greyish white; spines on tips of tubercles on surface of posterior abdomen; ventral surface of thighs and around cloaca dark grey. Digits, inner and outer metacarpal tubercles and inner metatarsal tubercle greyish white. Pectoral glands and femoral glands beige, mottled with orange patches. Iris yellowish brown, with greyish white patches on upper and lower margin.

Coloration of holotype in preservative. Yellowish brown fades to greyish brown dorsally. Color of the triangular marking between eyes, oblique bands on forearms, patterns on ventral surface faded. Orange patches on surface of throat, chest; color of pectoral glands and femoral glands faded.

Variation. Mensural data of the type series are listed in Table 4 . Most of the paratypes are similar to the holotype in morphology and color pattern, except for the following: dorsal surface of body yellowish brown in the holotype (vs. dorsal surface of body light brown in the paratypes SYS a007647 ( Fig. 4C ) and SYS a007648 ( Fig. 4E ); ventral surface dark grey with orange patches (vs. ventral surface lacking bright patches in the paratypes SYS a007649 ( Fig. 4B ), SYS a007647 ( Fig. 4D ) and SYS a007648 ( Fig. 4F ); iris yellowish brown, with greyish white patches on its upper and lower margin in the holotype (vs. iris grey with beige and dark mottling in the paratype SYS a007649 ( Fig. 4A ); tubercles on posterior part of abdomen of the paratype SYS a007648 ( Fig. 4F ) are weakly developed. Females are distinctly larger than the males.

Distribution and natural history. Currently, Boulenophrys puningensis sp. nov. is only known from its type locality, Longkeng Village of Puning. It inhabits flowing montane streams and the nearby forest floor and leaf litter at elevations between 120– 300 m. Advertisement calls of males were heard from February until April. Males were found calling in rock crevices in the flowing streams. Tadpoles could be found in this period.

Figures

FIGURE 3. The holotype (SYS a005770) of Boulenophrys puningensis sp. nov. in life.

FIGURE 4. The paratypes of Boulenophrys puningensis sp. nov. in life. A–B: male paratype, SYS a007649; C–D: female paratype, SYS a007647; E–F: female paratype, SYS a007648.

Tables

Table 4. Measurements (in mm) of the type series of Boulenophrys puningensis sp. nov., * indicates the holotype.

	SYS a005770*	SYS a006755	SYS a007649	SYS a007650	SYS a007647	SYS a007648
Sex	Male	Male	Male	Male	Female	Female
SVL	34.6	31.7	32.3	33.2	37.8	38.3
HDL	12.1	11.0	11.8	11.2	12.1	13.2
HDW	12.5	12.1	11.9	12.2	13.0	14.1
SNT	4.5	4.2	4.3	4.4	4.7	4.8
IND	4.4	3.9	4.0	3.9	4.8	4.8
IOD	3.8	3.4	3.4	3.5	3.9	4.1
ED	3.7	3.6	3.7	3.7	3.8	4.1
TD	2.6	2.6	2.5	2.6	2.7	2.9
TED	1.5	1.5	1.5	1.5	1.5	1.7
HND	8.0	7.8	8.1	8.3	8.4	9.9
RAD	7.2	7.0	7.2	7.1	7.6	7.7
TIB	13.1	12.5	13.1	12.6	13.5	15.1
FTL	18.7	17.7	18.7	18.4	20.5	22.1