Description of six new species of the subgenus Panophrys within the genus Megophrys (Anura, Megophryidae) from southeastern China based on molecular and morphological data Author Wang, Jian Author Lyu, Zhi-Tong Author Liu, Zu-Yao Author Liao, Cheng-Kai Author Zeng, Zhao-Chi Author Zhao, Jian Author Li, Yu-Long Author Wang, Ying-Yong text ZooKeys 2019 851 113 164 http://dx.doi.org/10.3897/zookeys.851.29107 journal article http://dx.doi.org/10.3897/zookeys.851.29107 1313-2970-851-113 F0AB358B811E42EEB25A5CF24658E6CA Megophrys (Panophrys) wugongensis J. Wang, Lyu & Y.Y. Wang sp. nov. Fig. 9, Table 8 Holotype. SYS a002625, adult male, collected by Guo-Ling Chen and Jian Zhao on 9 May 2014 from Yangshimu Scenic Area ( 27°34'47.93"N , 114°15'7.34"E ; 550 m a.s.l.), Pingxiang City, Jiangxi Province, China. Paratypes (three males & nine females). Adult males, SYS a004777/CIB110011, SYS a004796, 4800, collected by Zhi-Tong Lyu and Ying-Yong Wang on 23 May 2016, and adult females, SYS a002610-2611, collected by Guo-Ling Chen and Jian Zhao on 8 May 2014, SYS a004797-4799, 4801-4804, collected by Zhi-Tong Lyu and Ying-Yong Wang on 23 May 2016, from Wugongshan Scenic Area ( 27°34'3.94"N , 114°10'28.38"E ; 1050-1080 m a.s.l.), Anfu County, Ji'an City, Jiangxi Province, China. Diagnosis. (1) Body size small, SVL 31.0-34.1 mm in four adult males and body size moderate, SVL 38.5-42.8 mm in nine adult females; (2) tympanum distinct, slightly convex, moderate-sized, TD/ED ratio 0.47-0.52; (3) vomerine teeth absent; (4) margin of tongue not notched posteriorly; (5) hindlimbs short, heels not meeting, tibia-tarsal articulation reaching forward to the region between posterior corner of eye and posterior margin of tympanum; (6) TIB/SVL ratio 0.39-0.44, FTL/SVL ratio 0.56-0.64; (7) fingers without lateral fringes, presence of a subarticular tubercle at the bases of each finger, relative finger lengths II <I = IV <III; (8) toes with rudimentary webbing at their bases and without lateral fringes, subarticular tubercles only present at the base of each toe; (9) numerous granules present on dorsal surface of body, several large tubercles present on surface of flanks and dorsal surface of limbs; (10) presence of a small horn-like tubercle at the edge of the upper eyelid; (11) supratympanic fold distinct, whitish; (12) yellowish brown or reddish brown dorsally, with an incomplete dark triangular marking between eyes and an X-shaped marking on back of trunk; (13) ventral surface greyish brown, ventral surface of abdomen with creamy white nebulous patches and black spots; (14) males with a single subgular vocal sac; (15) gravid females bear creamy yellow oocytes. Comparisons. Comparative data of Megophrys wugongensis sp. nov. with M. dongduanensis sp. nov., M. nankunensis sp. nov., M. jiulianensis sp. nov., Megophrys nanlingensis sp. nov., M. feii and the 33 recognized members of Megophrys s.l. allocated to the subgenus Panophrys are listed in Table 5. Megophrys wugongensis sp. nov. differs from M. dongguanensis sp. nov., M. nankunensis sp. nov., M. jiulianensis sp. nov. and M. nanlingensis sp. nov. by a combination of following characters: vomerine teeth absent (vs. vomerine teeth present), tongue not notched posteriorly (vs. tongue notched in M. jiulianensis sp. nov. and M. nanlingensis sp. nov.), absence of lateral fringes on toes (vs. presence of narrow lateral fringes on toes in M. nanlingensis sp. nov.), heels not meeting when the flexed hindlimbs are held at right angles to the body axis (vs. heels overlapping in M. jiulianensis sp. nov. and M. nanlingensis sp. nov.), absence of black spines on dorsal skin (vs. present in M. jiulianensis sp. nov.), relative finger lengths II <I = IV <III (vs. II <I <IV <III in M. nankunensis sp. nov., M. jiulianensis sp. nov. and M. nanlingensis sp. nov.), ventral surface with creamy white nebulous patches (vs. absence of such patched on ventral surface in M. dongguanensis sp. nov. and M. nankunensis sp. nov.). With the smaller body size, SVL 31.0-34.1 mm in males and 38.5-42.8 mm in females, Megophrys wugongensis sp. nov. differs from the 13 members with larger SVL values: M. baolongensis (42.0-45.0 mm in males), M. binlingensis (45.1-51.0 mm in males), M. caudoprocta (81.3 mm in single male), M. hoanglienensis (37.4-47.6 mm in males, 59.6 mm in single female), M. jingdongensis (53.0-56.5 mm in males 63.5 in single female), M. latidactyla (38.9 mm in single male), M. liboensis (34.7-67.7 mm in males, 60.8-70.6 mm in females), M. omeimontis (56.0-59.5 mm in males, 68.0-72.5 mm in females), M. palpebralespinosa (36.2-38.0 mm in males), M. sangzhiensis (54.7 mm in single male), M. shuichengensis (102.0-118.3 mm in males, 99.8-115.6 mm in females), M. spinata (47.2-54.4 mm in males, 54.0-55.0 mm in females), and M. tuberogranulatus (33.2-39.6 mm in males, 50.5 mm in single female). Megophrys wugongensis sp. nov. differs from 12 species occurring in eastern and southern China ( M. acuta , M. brachykolos , M. boettgeri , M. cheni , M. huangshanensis , M. insularis , M. jinggangensis , M. kuatunensis , M. lini , M. lishuiensis , M. obesa and M. ombrophila ) by the following combination of characters: vomerine teeth absent (vs. present in M. insularis and M. jinggangensis ), tongue not notched posteriorly (vs. tongue notched in M. boettgeri , M. huangshanensis , M. kuatunensis and M. insularis ), toes without lateral fringes (vs. laterals fringes on toes narrow in M. acuta , M. kuatunensis and M. jinggangensis ; wide in M. boettgeri , M. cheni and M. lini ), toes with rudimentary webbing (vs. toes without webbing in M. huangshanensis , M. lishuiensis and M. ombrophila ), hindlimbs short, with heels not meeting when the flexed hindlimbs are held at right angles to the body axis (vs. hindlimbs comparatively longer, with heels overlapping in M. boettgeri , M. cheni , M. kuatunensis , M. jinggangensis and M. lini ), relative finger lengths II <I = IV <III (vs. I <II ≤ IV <III in M. acuta and M. ombrophila ; IV <II <I <III in M. brachykolos ; I <II = IV <III in M. lishuiensis ; I <II ≤ IV <III in M. obesa ), males bearing nuptial pads with nuptial spines during breeding season (vs. nuptials absence in adult males of M. acuta ), ventral surface with creamy white nebulous patches (vs. absence of such patched in M. brachykolos and M. obesa ). Megophrys nanlingensis sp. nov. differs from the remaining eight members of the Megophrys s.l. allocated to the subgenus Panophrys which share a moderate or small body size, by a combination of following characters: horn-like tubercle small at edge of the upper eyelid (vs. horn-like tubercle indistinct or absent in M. binchuanensis , M. minor , M. wuliangshanensis and M. wushanensis ), absence of vomerine teeth (vs. present in M. daweimontis , M. fansipanensis and M. rubrimera ), tongue not notched posteriorly (vs. tongue notched in M. minor , M. fansipanensis and M. rubrimera ), toes without lateral fringes (vs. lateral fringes wide in M. binchuanensis , M. wushanensis (in males); narrow in M. rubrimera ), toes with rudimentary webbing (vs. toes without webbing in M. daweimontis , M. fansipanensis , M. rubrimera and M. wuliangshanensis ), heels not meeting when the flexed hindlimbs are held at right angles to the body axis (vs. heels overlapping in M. minor and M. wuliangshanensis ), heels not meeting when the flexed hindlimbs are held at right angles to the body axis (heels meeting in M. binchuanensis ; heels meeting or overlapping in M. minor and M. wushanensis ; heels overlapping in M. leishanensis , and M. wuliangshanensis ). Megophrys wugongensis sp. nov. further differs from M. feii , for which molecular data are lacking and cannot be allocated to any subgenus base on morphology only ( Yang et al. 2018 ) by the larger body size, SVL 31.0-34.1 mm in males and 38.5-42.8 mm in females (VS. 24.3-25.1 mm in males, 28.2-28.9 mm in females), presence of nuptial pad with nuptial spines in males during breeding season (vs. absent), absence of lateral fringes on toes (vs. moderate or wide). Description of holotype. Adult male. Habitus small, SVL 31.0 mm; head width slightly larger than head length, HDW/HWL 1.03; snout rounded in dorsal view, tip of snout slightly sharpened, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eye large, ED/HDL 0.41; nostril oblique ovoid; pupil vertical; canthus rostralis well developed; loreal region vertical; internasal distance slightly larger than interorbital distance; tympanum distinct, moderate-sized, TD/ED 0.47; large ovoid choanae at the base of the maxilla; weak vomerine ridge present, vomerine teeth absent; margin of tongue not notched posteriorly; internal vocal slits present near the rear of the lower mandible. RAD/SVL 0.24, HND/SVL 0.22; absence of lateral fringes and webbing on fingers, relative finger lengths II <I = IV <III; tip of finger rounded, slightly swollen; presence of a distinct subarticular tubercle on the base of each finger; outer metacarpal tubercles indistinct, inner metacarpal tubercles distinct and observably enlarged. Hindlimbs short, tibio-tarsal articulation reaching forward the posterior corner of eye when hindlimb is stretched along the side of the body; heels not meeting when the flexed hindlimbs are held at right angles to the body axis; TIB/SVL 0.43 and FTL/SVL 0.61; relative toe lengths I <II <V <III <IV; tips of toes round and slightly dilated; presence of rudimentary webbing on toes but absence of lateral fringes and tarsal folds; presence of a subarticular tubercle only at the bases of each toes; presence of a long ovoid inner metatarsal tubercle and absence of outer metatarsal tubercle. Dorsal skin texture rough with dense granules, some of which forming an X-shaped skin ridge on center of trunk; surface of flanks with large tubercles; presence of a small horn-like tubercle at the edge of eye lid; distinct supratympanic fold curving posteroventrally from posterior corner of eye to a level above insertion of arm; superior margin of tympanum in connect with supratympanic fold; ventral skin texture smooth with granules on the surface of abdomen; pectoral gland large, closer to axilla; single large femoral gland on rear of thigh. Measurements of holotype (in mm). SVL 30.8, HDL 11.9, HDW 11.7, SNT 3.5, IND 3.0, IOD 2.8, ED 3.5, TD 1.8, TED 1.7, HND 8.5, RAD 7.2, FTL 21.8, TIB 15.1 Coloration of holotype in life. (Fig. 9 A-C ) Dorsal surface reddish brown, with a distinct and dark triangular marking with yellow edges between eyes. Hindlimb with broad black transverse bands. A dark brown vertical band below the eye. Canthus rostralis and supratympanic fold white. Horn-like tubercle at the edge of the upper eyelid yellow. Surface of throat and chest dark brown, with scarlet marbling, posterior region of abdomen white. Ventral surface of limbs brown with white spots and patches. Ventral surface of hand and foot brown, inner and outer metatarsal tubercles and inner metacarpal tubercle pink. Pectoral and femoral glands white. Iris reddish brown. Figure 9. Megophrys wugongensis sp. nov. in life: A-C SYS a002625, male holotype D SYS a002610, female paratype E, F SYS a002611, female paratype. Coloration of holotype in preservative. Dorsum dark brown, markings on dorsal surface became indistinct, transverse bands on limbs became dark grey and became more distinct . Surface of throat and chest light brown, posterior region of abdomen light yellow, ventral surface of limbs light brown, inner and outer metatarsal tubercles and inner metacarpal tubercle light yellow, all marbling, colored spots and patches absent. Variation. Measurement data of type series are listed in Table 8. All paratypes are very similar to holotype in morphology and color pattern. However, dorsal surface yellowish brown in female paratypes SYS a004798, 4801, 4804, markings on dorsal skin indistinct in male paratypes SYS a004777/CIB110011 and SYS a004796, and female paratypes SYS a002610 (Fig. 9D), 4797, 4799, presence of a rectangle marking on central back of trunk in the female paratype SYS a002611 (Fig. 9 E-F ). Table 8. Measurements (in mm; minimum-maximum, mean +/- SD) of the type series of Megophrys wugongensis sp. nov.

Species	Megophrys wugongensis sp. nov.
	Males (n = 4)	Females (n = 9)
SVL
HDL
HDW
SNT
IND
IOD
ED
TD
TED
HND
RAD
FTL
TIB
HDLSVL
HDWSVL
HDWHDL
SNTHDL
SNTSVL
INDHDW
IODHDW
EDHDL
EDSVL
TDED
TEDTD
HNDSVL
RADSVL
TIBSVL
FTLSVL

Etymology. The specific epithet "wugongensis" is in reference to the type locality of the new species in the Wugong Mountains. We propose the common English name "Wugongshan Horned Toad" and Chinese name "Wu Gong Shan Jiao Chan ( 武功山角蟾)" . Distribution and habits. Currently, Megophrys wugongensis sp. nov. is known from the type locality, Yangshimu Scenic Area, Pingxiang City, Jiangxi Province at approximate 550 m a.s.l., Wugongshan Scenic Area, Ji'an City, Jiangxi Province at approximate 1050 -1080 m a.s.l., all located in the Luoxiao Mountains in eastern China. All specimens were collected on leaf litter near a stream in the bamboo forest, males were not heard calling. In consideration of the invisible nuptial pad and nuptial spines in all male specimens and the undeveloped fallopian tubes in all female specimens, the breeding season of M. wugongensis sp. nov. still remains unknown. Tadpoles were not observed. Megophrys wugongensis sp. nov. is sympatric with M. jinggangensis in all localities.