Panophrys congjiangensis, Luo & Wang & Wang & Lu & Wang & Deng & Zhou, 2021

Luo, Tao, Wang, Yali, Wang, Siwei, Lu, Xueli, Wang, Weifeng, Deng, Huaiqing & Zhou, Jiang, 2021, A species of the genus Panophrys (Anura, Megophryidae) from southeastern Guizhou Province, China, ZooKeys 1047, pp. 27-60 : 27

Panophrys congjiangensis sp. nov.

, Figs 4 , 5 , 6

Type material.

Holotype. GZNU20200706010 (Figs 4 , 5 ), adult male, collected by Tao Luo on 6 May 2020 in the Yueliangshan Nature Reserve, Congjiang County, Guizhou Province, China (25.614417°N, 108.410076°E; ca. 730 m a.s.l.).

Paratypes. Nineteen adult specimens (15 males and 4 females) from the same locality. Eleven males (GZNU20200706001-06009, GZNU20200706012-06013) collected with the holotype on 6 July 2020 by Tao Luo, Xueli Lu, and Weifeng Wang. One female (GZNU20200706011) collected with the holotype by Tao Luo. Three males (GZNU200707001-07003) collected on 7 July 2020 by Tao Luo. Three females (GZNU20200706004, GZNU20200706005, and GZNU20200706006) collected on 7 July 2020 by Tao Luo.

Etymology.

The specific epithet " Panophrys congjiangensis " refers to the holotype locality, which is Yueliangshan Nature Reserve, located in Congjiang County, Guizhou Province, China. We propose the English common name "Congjiang Horned Toad" and the Chinese common name "Cong Jiang Jiao Chan ( 从江角蟾)”.

Differential diagnosis.

Panophrys congjiangensis sp. nov. is assigned to the genus Panophrys based on molecular phylogenetic analyses and the following characteristics, which are diagnostic for this genus: (1) snout shield-like; (2) snout projecting beyond the lower jaw; (3) tympanum distinct (4) canthus rostralis distinct; (5) chest gland small and round, closer to axilla than to midventral line; (6) femoral gland on rear of thigh; (7) vertical pupils (Fei et al. 2006; Fei and Ye 2016; Su et al. 2020).

Panophrys congjiangensis sp. nov. is distinguished from its congeners by a combination of the following characters: (1) medium body size (SVL: 28.6-33.4 mm in males and 38.4-40.2 mm in females); (2) single small horn-like tubercle at edge of each upper eyelid; (3) tympanum distinctly visible (TD/ED ratio 0.47-0.66); (4) vomerine teeth absent; (5) tongue not notched behind; (6) a narrow and unobvious lateral fringe on toes; (7) relative finger lengths II <I <V <III; (8) rudimentary webs on toes; (9) hindlimbs slender, heels overlapping when thighs are positioned at right angles to body; (10) two metacarpal tubercles on palm, with inner metatarsal tubercle long and oval-shaped; (11) tibiotarsal articulation reaching nostril when leg is stretched forward; (12) dorsal skin rough, with numerous orange-red granules, ventral surface smooth; (13) single internal subgular vocal sac present in males; (14) in breeding males, weak gray-black nuptial pads with black nuptial spines present on dorsal surfaces of bases of first and second fingers.

Description of holotype.

GZNU20200706010 (Figs 4 , 5 ), adult male. Medium body size, SVL 33.4 mm; head length slightly larger than head width (HDL/HDW ratio 1.02); snout short, rounded and projecting beyond the lower jaw in dorsal view, longer than eye diameter (SNT/ED ratio 1.11); nostril rounded, distinct, and closer to the tip of the snout than to the eye (SNT/NED ratio 1.83); internasal distance greater than interorbital distance (IND/IOD ratio 1.19); internasal distance greater than upper eyelid width (IND/UEW ratio 1.28); region vertical and concave; canthus rostralis well-developed; top of head slightly concave in dorsal view; a small horn-like tubercle at the edge of the upper eyelid; eyes large, slightly protuberant in dorsal view, eye diameter 34% of head length, pupils vertical (Fig. 4H ); tympanum distinct, tympanum diameter less than eye diameter (TD/ED ratio 0.63); vomerine ridges and vomerine teeth absent; tongue is melon seed-shaped and not notched behind (Fig. 5E ).

Forelimbs slender and comparatively short, the length of lower arm and hand 44.01% of SVL; fingers slender, relative finger lengths: II <I <IV <III; tips of fingers slightly dilated, round, without lateral fringes; one distinct subarticular tubercle at the base of each finger; two metacarpal tubercles on the palm; prominent, the outer one long and thin, the inner one oval-shaped, inner metacarpal tubercles longer than outer metacarpal tubercles (IPTL/OPT ratio 1.13).

Hindlimbs slender (HLL/SVL ratio 1.80); heels slightly overlapping when thighs are positioned at right angles to the body; tibiotarsal articulation reaching the nostril when leg stretched forward; foot length less than tibia length (FL/TL ratio 0.90); relative toe lengths I <II <V <III <IV; tips of toes round and slightly dilated; toes with narrow and unobvious lateral fringes and rudiment webs; one subarticular tubercle at the base of each toe; inner metatarsal tubercle long oval-shaped and the outer one absent.

Dorsal skin rough with numerous orange-red granules; several large warts scattered on flanks and dorsal limbs; several tubercles on upper eyelid, including a small horn-like prominent tubercle on the edge (Fig. 4H ); supratympanic fold distinct; tubercles on the dorsum forming a discontinuous X-shaped ridge, the V-shaped ridges disconnected; two discontinuous dorsolateral parallel ridges on either side of the X-shaped ridges; an inverted triangular brown speckle between two upper eyelids; four transverse skin ridges on the dorsal shank and thigh; ventral surface smooth; chest with small, round glands, closer to the axilla than to midventral line; femoral glands on rear of thigh; numerous white granules on ventral surface of thigh; posterior end of body distinctly protruding, forming an arc-shaped swelling above anal region.

Coloration of holotype in life

(Fig. 4 ). Dorsal surfaces of head and trunk brownish gray; triangular marking with light edge between eyes; dark X-shaped marking with light edge on central dorsum; supratympanic fold light brown; four dark brown transverse bands on dorsal surfaces of thigh and shank; 2-4 dark brown and white vertical bars on lower and upper lip; dark vertical band below eye; iris copper-brown; throat and anterior chest light purple-brown; belly light orange-red with large white blotch and small grey blotch in belly center, and small white blotches and large black patches on belly sides, forming a discontinuous line; ventral surfaces of forelimbs purplish brown; some white spots on the ventral surfaces of hindlimbs; palms orange-red with a small black-brown blotch; ventral surfaces of first and second toes orange-red, ventral surfaces of remaining three toes black-brown; soles black-brown; tips of digits grey-white; pectoral and femoral glands white.

Preserved holotype coloration

(Fig. 5 ). After preservation in ethanol, dorsal surfaces light brownish grey; dorsal surface of head dark gray; X-shaped ridges on dorsum indistinct and transverse bands on limbs and digits distinct, coloration lighter; throat dark black-brown; chest light black-brown; belly light gray-white with large black-brown blotches on sides and a small gray-brown blotch in center; posterior ventral body surface, inner thigh, and upper part of tibia milky yellow; palms and metatarsal tubercle milky yellow with a small gray-brown blotch; ventral surfaces of soles and toes dark black-brown; inner metatarsal tubercle milky yellow.

Variations.

Measurements of the type series are shown in Tables 2 , 4 . Females (SVL 39.3 ± 0.7 mm, N = 4) had larger bodies than males (SVL 31.2 ± 1.4 mm, N = 15). In life, the diagnostic morphological characters of all paratypes were identical to those of the holotype. However, coloration and stripe patterns differed among individuals (Fig. 6 ). For example, male GZNU20200706007 (Fig. 6A ) had a brown-black back and a black-brown belly with some large white patches, as well as two V-shaped markings that were virtually connected. This specimen also had warts on both sides of the body, forming a transverse skin ridge that almost connected to the second V-shaped marking. In contrast, male GZNU20200706008 (Fig. 6B ) had a large black spot between the upper eyelids. The throat and anterior belly of this specimen were purple-brownish, while the belly was light milky yellow, with two large black blotches and a small white blotch on the body sides. In specimens GZNU20200706009 and GZNU20200706012 (Fig. 6C, E ), the warts on both sides of the body formed transverse skin ridges connected to the second V-shaped marking and extending behind the tympanum; three white small blotches were present on the body sides. In specimens GZNU20200706013 and GZNU20200706012 (Fig. 6D, E ), the back was light reddish brown.

Advertisement call.

The call description is based on recordings of the holotype GZNU 20200706010 (Fig. 7 ) from the bamboo forest near the streamlet. The ambient air temperature during the recording was 25.3 °C. Each call contains 9-14 syllables (mean 11.60 ± 2.07, N = 5). The call consists of a few strophes, each 2.41-3.43 s in duration (mean 2.75 ± 0.46, N = 4). Each syllable has a duration of 0.05-0.09 s (mean 0.07 ± 0.06, N = 58). The interval between syllables has a duration of 0.10-0.31 s (mean 0.167 ± 0.042, N = 53).

Sexual dimorphism.

Adult males (SVL 28.6-33.4 mm) smaller than adult females (SVL 38.4-40.2 mm). Adult males with single internal subgular vocal sac (Fig. 4A ). Breeding males with grey-black nuptial pads with obvious black nuptial spines on dorsal surfaces of bases of first and second fingers.

Comparisons.

Comparative data of Panophrys congjiangensis sp. nov. with 59 recognized congeners of Panophrys are given in Suppl. material 4: Table S4.

By having small body size, SVL 30.4-34.1 mm in males, Panophrys congjiangensis sp. nov. differs from P. baolongensis (42.0-45.0 in males), P. binlingensis (45.1-51.0 in males), P. boettgeri (34.5-37.8 in males), P. caobangensis (34.9-38.9 in males), P. caudoprocta (81.3 in single male), P. hoanglienensis (37.4-47.6 in males), P. huangshanensis (36.0-41.6 in males), P. insularis (36.8-41.2 in males), P. jingdongensis (53.0-56.5 in males), P. mirabilis (55.8-61.4 in males), P. obesa (35.6 in single male), P. palpebralespinosa (36.2-38.0 in males), P. sangzhiensis (54.7 in single male), and P. xiangnanensis (38.6-42.0 in males). By having larger body size, SVL 30.4-34.1 mm in males, Panophrys congjiangensis sp. nov. differs from P. cheni (26.2-29.5 in males), P. daiyunensis (27.6-28.7 in males), P. kuatunensis (26.2-29.6 in males), P. sanmingensis (27.0-29.5 in males), P. yeae (23 in single male), and P. zhoui (23.8-29.1 in males). By having small body size, SVL 38.9-40.2 mm in females, Panophrys congjiangensis sp. nov. differs from P. fansipanensis (41.7-42.5 in females), P. minor (42.0-48.2 in females), P. tuberogranulata (50.5 in single female), P. wuliangshanensis (41.3 in single female), P. xianjuensis (41.6 in single female), and P. yangmingensis (45.2 in single female).

Nine Panophrys species were previously recorded from the Guizhou Province, namely P. anlongensis , P. chishuiensis , P. jiangi , P. leishanensis , P. liboensis , P. omeimontis , P. shuichengensis , P. spinata , and P. qianbeiensis . Panophrys congjiangensis sp. nov. differs from P. anlongensis by having small body size, SVL 30.4-34.1 mm in males and 38.9-40.2 mm in females (vs. 40.0-45.5 mm in males and 48.9-51.2 in females), tibiotarsal articulation reaching the nostril when leg stretched forward (vs. reaching to the level of mid-eye). Panophrys congjiangensis sp. nov. differs from P. chishuiensis by having small body size, SVL 30.4-34.1 mm in males and 38.9-40.2 mm in females (vs. 43.4-44.1 mm in males and 44.8-49.8 in females), rudimentary webs on toes (vs. lacking webs), subarticular tubercles present on each toes (vs. absent), tibiotarsal articulation reaching the nostril when leg stretched forward (vs. reaching the level between tympanum and eye). Panophrys congjiangensis sp. nov. differs from P. jiangi by having slightly small body size, SVL 30.4-34.1 mm in males (vs. 34.4-39.2 mm in males), relative finger lengths are II <I <V <III (vs. I <II <V <III), tibiotarsal articulation reaching the nostril when leg stretched forward (vs. reaching forward to the region between tympanum and eye). Panophrys congjiangensis sp. nov. differs from P. liboensis , P. omeimontis , P. qianbeiensis , P. shuichengensis , and P. spinata by having small body size, SVL 30.4-34.1 mm in males and 38.9-40.2 mm in females (vs. SVL>40 mm in males in P. liboensis , P. omeimontis , P. qianbeiensis , P. shuichengensis , and P. spinata ; vs. SVL>50 mm in females in P. liboensis , P. omeimontis , P. shuichengensis , and P. spinata ), small horn-like tubercle at the edge of each upper eyelid (vs. slightly large in P. liboensis and P. shuichengensis ; absence in P. qianbeiensis ), absence of vomerine teeth (vs. present in P. liboensis , P. omeimontis , and P. qianbeiensis ), tongue not notched behind (vs. notched in P. liboensis , P. omeimontis , P. qianbeiensis , P. shuichengensis , and P. spinata ), lateral fringes on toes narrow and unobvious (vs. wide in P. liboensis , P. qianbeiensis , P. shuichengensis , and P. spinata ), rudimentary webs on toes (vs. more than one-fourth webs in P. qianbeiensis , P. shuichengensis , and P. spinata ), subarticular tubercles present on each toes (vs. absent in P. liboensis and P. shuichengensis ), tibiotarsal articulation reaching the nostril when leg stretched forward (vs. reaching to ocular region in P. liboensis , P. omeimontis , P. shuichengensis , and P. spinata ; reaching to the level between tympanum and eye in P. qianbeiensis ). Panophrys congjiangensis sp. nov. differs from P. leishanensis by having slightly small body size, SVL 38.9-40.2 mm in females (vs. 42.3 in single female), having narrow and unobvious lateral fringes on toes (vs. lacking), tibiotarsal articulation reaching the nostril when leg stretched forward (vs. reaching between tympanum to eye). The mean SVL of male Panophrys congjiangensis sp. nov. was significantly greater than that of P. leishanensis . In addition, the ratios of HDL, HDW, SNT, LAHL, LW, THL, TFL, HLL, and TW to SVL were all significantly greater in male Panophrys congjiangensis than in male P. leishanensis (all p -values <0.05; Table 4 ). Panophrys congjiangensis sp. nov. also differs from P. leishanensis by having one call 9-14 syllables (vs. calls of P. leishanensis , which are 12-14 syllables long), shorter call intervals between syllables (0.167 ± 0.042 s, N = 53 in the new species vs. 0.409 ± 0.075 s, N = 36 in P. leishanensis ), and shorter call syllables (0.07 ± 0.06 s, N = 58 in the new species vs. 0.105 ± 0.003 s, N = 37 in P. leishanensis ).

From the remaining 24 species occurring in Panophrys , Panophrys congjiangensis sp. nov. can be distinguished by the absence of vomerine teeth (vs. present in P. daweimontis , P. dongguanensis , P. frigida , P. jinggangensis , P. jiulianensis , P. nankunensis , P. nanlingensis , P. rubrimera , P. shimentaina , and P. tongboensis ), by the unnotched tongue (vs. tongue notched in P. binchuanensis , P. cheni , P. kuatunensis , and P. lushuiensis ), by the small horn-like tubercle at edge of upper eyelid (vs. slightly large in P. acuta ), by the absence of lateral fringes on toes (vs. lacking in P. angka , P. brachykolos , P. lishuiensis , P. ombrophila , P. shunhuangensis , and P. wugongensis ; vs. wide in P. lini ; vs. lacking in males in P. wushanensis , wide in females in P. wushanensis ), by the subarticular tubercles present (vs. absent in P. baishanzuensis and P. mufumontana ), tibiotarsal articulation reaching the nostril when leg stretched forward (vs. reaching to ocular region in P. acuta , P. baishanzuensis , P. binchuanensis , P. jiulianensis , P. lini , P. nanlingensis , P. ombrophila , and P. wushanensis ; vs. reaching to the level between tympanum and eye in P. angka , P. dongguanensis , P. kuatunensis , P. lishuiensis , and P. nankunensis ; vs. reaching to the level between eye and snout in P. cheni , P. daweimontis , and P. shunhuangensis ; vs. reaching to the level behind the eye in P. brachykolos , P. mufumontana , P. shimentaina , and P. wugongensis ; vs. reaching to the level at center of tympanum P. daoji ).

Distribution and ecology.

Panophrys congjiangensis sp. nov. is only known from the type locality, Yueliangshan Nature Reserve, Congjiang County, Guizhou Province, China at elevations of 1142-1206 m. Individuals of the new species were frequently found in bamboo forests, grasses, and shrubberies near streams. Plants in the type locality predominantly fall into the families Urticaceae , Gramineae , Cyperaceae , Rosaceae , Dryopteridaceae , Polygonaceae , Aquifoliaceae , and Fagaceae . In the Yueliang Mountains, Panophrys congjiangensis sp. nov. is sympatric with Pachytriton inexpectatus Nishikawa, Jiang, Matsui & Mo, 2010; Amolops sinensis Lyu, Wang & Wang, 2019; Nidirana leishanensis Li, Wei, Xu, Cui, Fei, Jiang, Liu & Wang, 2019; Hylarana latouchii (Boulenger, 1899); Quasipaa boulengeri ( Günther, 1889); Hyla annectans (Jerdon, 1870); Opisthotropis zhaoermii Ren, Wang, Jiang, Guo & Li, 2017; Trimeresurus stejnegeri (Schmidt, 1925); and Rhabdophis tigrinus (Boie, 1826). These species were often found in the same streams as Panophrys congjiangensis sp. nov.

Figures

Figure 4. Morphological features of the live adult male holotype GZNU 20200706010 of Panophrys congjiangensis sp. nov. A single subgular vocal sac B dorsal view C dorsolateral view D ventral view E ventral view of hand F ventral view of foot G dorsal view of hand (1 indicates villiform gray-black nuptial spines) H iris. A was photographed at about 9 p. m., and B to H during the day, respectively.

Figure 5. Morphological features of the preserved adult male holotype GZNU 20200706010 of Panophrys congjiangensis sp. nov. A dorsal view B ventral view C lateral view D view of oral cavity E tongue F ventral view of hand G ventral view of foot.

Figure 6. Paratypes of Panophrys congjiangensis sp. nov. in life A GZNU 20200706007, adult male B GZNU 20200706008, adult male C GZNU 20200706009, adult male D GZNU 2020706013, adult male E GZNU 20200706012, adult female. So, the images were all taken at 8 am.

Figure 7. Visualization of advertisement calls of Panophrys congjiangensis sp. nov. A waveform showing 12 notes of one call B waveform showing one note C sonogram showing 12 notes of one call D sonogram showing one note.