Three new species of the genus Boulenophrys (Anura, Megophryidae) from southern China Author Qi, Shuo 0000-0002-2924-6093 State Key Laboratory of Biocontrol / The Museum of Biology, School of Ecology / School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China & qishuo 1992 @ outlook. com; https: // orcid. org / 0000 - 0002 - 2924 - 6093 & These authors contributed equally to this work qishuo1992@outlook.com Author Lyu, Zhi-Tong 0000-0002-4891-3859 State Key Laboratory of Biocontrol / The Museum of Biology, School of Ecology / School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China & lvzht @ foxmail. com; https: // orcid. org / 0000 - 0002 - 4891 - 3859 & These authors contributed equally to this work lvzht@foxmail.com Author Wang, Jian 0000-0003-4249-7767 State Key Laboratory of Biocontrol / The Museum of Biology, School of Ecology / School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China & wangj- 1994 @ outlook. com; https: // orcid. org / 0000 - 0003 - 4249 - 7767 wangj-1994@outlook.com Author Mo, Yun-Ming Natural History Museum of Guangxi, Nanning, Guangxi 530012, China. moyunming @ 163. com Author Zeng, Zhao-Chi 0000-0003-4054-8399 State Key Laboratory of Biocontrol / The Museum of Biology, School of Ecology / School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China & zhaochizeng @ outlook. com; https: // orcid. org / 0000 - 0003 - 4054 - 8399 zhaochizeng@outlook.com Author Zeng, Yang-Jin Guangdong Shimentai National Nature Reserve, Qingyuan, Guangdong 513000, China Author Dai, Ke-Yuan Guangdong Shimentai National Nature Reserve, Qingyuan, Guangdong 513000, China Author Li, Yuan-Qiu Guangdong Shimentai National Nature Reserve, Qingyuan, Guangdong 513000, China Author Grismer, L. Lee 0000-0001-8422-3698 Herpetology Laboratory, Department of Biology, La Sierra University, Riverside, California 92515, USA & Department of Herpetology, San Diego Natural History Museum, PO Box 121390, San Diego, California, 92112, USA & lgrismer @ lasierra. edu; https: // orcid. org / 0000 - 0001 - 8422 - 3698 lgrismer@lasierra.edu Author Wang, Ying-Yong 0000-0003-2024-6641 State Key Laboratory of Biocontrol / The Museum of Biology, School of Ecology / School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China & wangyy @ mail. sysu. edu. cn; https: // orcid. org / 0000 - 0003 - 2024 - 6641 wangyy@mail.sysu.edu.cn text Zootaxa 2021 2021-12-02 5072 5 401 438 journal article 3143 10.11646/zootaxa.5072.5.1 5758176c-96f6-43b9-b9a9-b2fac894dca8 1175-5326 5748979 B23A6B99-6D47-469F-8A4F-8AD41D77151A Boulenophrys yingdeensis sp. nov. Qi, Lyu, Wang & Wang Yingde Horned Toad / ying de jiao chan (Ȓ德fflDz) Figures 4 , 5 , 7B Chresonymy. Megophrys sp 4 (SYS a002100, 4721, 5447)— Liu et al . 2018 Holotype . SYS a002099 ( Figures 4A , 7B ), adult male, collected on 26 April 2013 by Run-Lin Li from Shimentai Nature Reserve ( 24.4435°N , 113.3034°E ; ca 357 m a.s.l. ), Yingde City , Guangdong Province , China . Paratypes . Eight adult specimens from the same locality as the holotype: female SYS a001563 collected on 23 April 2012 by Run-Lin Li; female SYS a004721 collected on 29 April 2016 by Ying-Yong Wang, Jian Wang and Zhi-Tong Lyu; female SYS a005447 and male SYS a005449 collected on 19 August 2016 by Zhi-Tong Lyu; male SYS a007114/ CIB 116084 collected on 20 June 2018 by Hong-Hui Chen, Jia-He Li and Jian Wang; male SYS a007405 and two females SYS a007406 and SYS a007407 collected on 20 August 2018 by Jian Wang . Etymology. The specific epithet yingdeensis refers to its type locality, Yingde City in northern Guangdong . Diagnosis. (1) Small body size, SVL 33.2–35.3 mm (34.2 ± 1.0, N = 4) in adult males and SVL 36.3–45.8 mm (40.7 ± 4.2, N = 5) in adult females; (2) snout rounded in dorsal view; (3) tympanum boundary clear, ED/TD 1.65– 1.95 in males, 1.48–2.09 in females; (4) vomerine ridge prominent bearing vomerine teeth; (5) margin of tongue rounded, not notched behind; (6) hindlimbs slender, heels overlapping or just meeting and tibio-tarsal articulation reaching forward between tympanum to posterior corner of eye; (7) tibia 0.46–0.48 of SVL and foot 0.61–0.67 of SVL in males, while tibia 0.44–0.46 of SVL and foot 0.61–0.66 of SVL in females; (8) toes without lateral fringes and with only rudimentary webbing; (9) presence of small, horn-like tubercle at the edge of upper eyelid; (10) dorsal skin smooth, a discontinuous “Y” or “X”-shaped ridge on the mid-dorsum, two discontinuous dorsolateral ridges on two side on the dorsum; (11) skin of flanks smooth with small conical tubercles; (12) single subgular vocal sac in males; (13) presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males. Comparisons. Comparative data of Boulenophrys yingdeensis sp. nov. from B. yaoshanensis sp. nov. and the other recognized members of the genus Boulenophrys are listed in Table 3 . Having a smaller body size with SVL 33.2–35.3 mm in males, Boulenophrys yingdeensis sp. nov. differs from seven congeners whose SVL> 50 mm in males, including B. caudoprocta ( 81.3 mm in single male), B. jingdongensis (53.0– 56.5 mm in males), B. liboensis ( 60.5–67.7 mm in males), B. mirabilis ( 55.8–61.4 mm in males), B. omeimontis (56.0– 59.5 mm in males), B. sangzhiensis ( 54.7 mm in single male), and B. shuichengensis (102.0– 118.3 mm in males). Having relatively longer shanks with the heels overlapping or meeting when the flexed hindlimbs are held at right angles to the body axis, Boulenophrys yingdeensis sp. nov. can be easily distinguished from the following nine congeners, B. acuta , B. brachykolos , B. daoji , B. dongguanensis , B. insularis , B. nankunensis , B. obesa , B. ombrophila , and B. wugongensis (vs. all of which have relatively shorter shanks with the heels not meeting). Having vomerine teeth, Boulenophrys yingdeensis sp. nov. differs from B. acuta , B. angka , B. anlongensis , B. baishanzuensis , B. baolongensis , B. binchuanensis , B. binlingensis , B. boettgeri , B. brachykolos , B. caobangensis , B. congjiangensis , B. cheni , B. chishuiensis , B. daoji , B. huangshanensis , B. jiangi , B. kuatunensis , B. leishanensis , B. lini , B. lishuiensis , B. lushuiensis , B. minor , B. mirabilis , B. mufumontana , B. obesa , B. ombrophila , B. sanmingensis , B. shuichengensis , B. shunhuangensis , B. spinata , B. tuberogranulatus , B. wugongensis , B. wuliangshanensis , B. wushanensis , B. xiangnanensis , B. xianjuensis , B. yangmingensis , and B. yaoshanensis sp. nov. (vs. absence of vomerine teeth in these species). Having an unnotched tongue, Boulenophrys yingdeensis sp. nov. can be distinguished from B. baolongensis , B. binlingensis , B. boettgeri , B. cheni , B. hoanglienensis , B. huangshanensis , B. insularis , B. jingdongensis , B. jiulianensis , B. kuatunensis , B. liboensis , B. lushuiensis , B. minor , B. nanlingensis , B. omeimontis , B. qianbeiensis , B. sangzhiensis , B. sanmingensis , B. shuichengensis B. spinata , and B. tongboensis (vs. tongue notched posteriorly in these species). Lacking lateral fringes on toes, Boulenophrys yingdeensis sp. nov. differs from B . acuta , B. anlongensis , B . baishanzuensis , B. binchuanensis , B. boettgeri , B. congjiangensis , B. cheni , B. daiyunensis , B. daoji , B. jingdongensis , B. jinggangensis , B. liboensis , B. lini , B. lushuiensis , B. mirabilis , B. mufumontana , B. nanlingensis , B. omeimontis , B. palpebralespinosa , B. qianbeiensis , B. rubrimera , B. sangzhiensis , B. sanmingensis , B. shimentaina , B. shuichengensis , B. spinata , B. xiangnanensis , B. xianjuensis , and B. yangmingensis (vs. presence of lateral fringes on toes in these species); and from B. wushanensis (vs. presence of wide lateral fringes on toes in males while lacking in females). Having rudimentary webbing on toes, Boulenophrys yingdeensis sp. nov. differs from B. baishanzuensis , B. baolongensis , B. daweimontis , B. fansipanensis , B. frigida , B. huangshanensis , B. kuatunensis , B. lishuiensis , B. ombrophila , B. rubrimera , B. tongboensis , and B. wuliangshanensis (vs. absence of webbing on toes in these species); and from B. jingdongensis , B. palpebralespinosa , B. qianbeiensis , B. shuichengensis , and B. spinata (vs. presence of well-developed webbing on toes in these species). Description of holotype . Adult male. small body size, SVL 35.3 mm ; head width slightly larger than head length, HWD/HDL 1.07; 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 0.30 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 boundary clear, ED/TD 1.68; large ovoid choanae at the base of the maxilla; vomerine ridge prominent, vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched behind; presence of a single subgular vocal sac, a pair of slit-like openings at posterior of jaw. FIGURE 4. Boulenophrys yingdeensis sp. nov. in life. A: holotype (SYS a002099), male; B: paratype (SYS a004721), female; 1: dorsolateral view; 2: ventral view; 3: hand; 4: foot. Radio-ulnar length 0.20 of SVL and hand 0.26 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; inner metacarpal tubercle observably enlarged and the outer one smaller; a single nuptial pad on the dorsal surface of the first and second fingers. Hindlimbs slender, tibio-tarsal articulation reaching forward the posterior corner of the eye when hindlimb is stretched along the side of the body; heels overlapping when the flexed hindlimbs are held at right angles to the body axis; tibia length 0.48 of SVL and foot length 0.67 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 webs; ventromedial callous pad on toe I is round, on others are long strip shape; inner metatarsal tubercle long ovoid and the outer one absent. TABLE 5. Measurements (in mm) and ratios of the type series of Boulenophrys yingdeensis sp. nov . , * indicates the holotype.

SYS a002099*	SYS a005449	SYS a007114/ CIB 116084	SYS a007405	SYS a001563	SYS a004721	SYS a005447	SYS a007406	SYS a007407
Sex	Male	Male	Male	Male	Female	Female	Female	Female	Female
SVL	35.3	33.4	33.2	34.8	45.8	37.4	39.9	36.3	44.2
HDL	12.4	11.3	11.1	11.9	15.2	13.3	12.9	11.7	14.0
HDW	13.3	11.6	11.9	11.6	15.8	14.2	13.1	12.2	14.5
SNT	4.1	3.8	4.1	4.0	4.7	4.8	4.4	4.2	4.8
IND	4.1	3.7	4.0	3.8	4.3	4.4	4.3	3.8	4.2
IOD	3.6	3.0	3.3	3.3	4.0	3.5	3.8	3.4	3.9
ED	3.7	3.9	3.9	3.8	4.9	4.6	3.7	3.6	4.2
TD	2.2	2.1	2.0	2.3	3.0	2.2	2.5	2.0	2.7
TED	2.3	2.1	1.9	2.2	2.4	1.9	2.5	2.1	2.6
HND	9.2	7.2	8.3	8.7	11.3	9.0	9.6	9.1	10.2
RAD	6.9	7.1	7.4	6.8	9.2	8.4	7.7	7.8	8.9
TIB	16.8	15.2	15.9	16.2	20.9	16.9	17.5	16.6	19.6
FTL	23.8	20.3	21.9	22.1	30.1	22.9	25.5	24.1	27.6
ED/TD	1.68	1.86	1.95	1.65	1.63	2.09	1.48	1.80	1.56
TIB/SVL	0.48	0.46	0.48	0.47	0.46	0.45	0.44	0.46	0.44
FTL/SVL	0.67	0.61	0.66	0.64	0.66	0.61	0.64	0.66	0.62

FIGURE 5. Color variation of Boulenophrys yingdeensis sp. nov. from the type locality. These individuals were captured, photographed, and finally released. Skin of dorsum bearing small granules; skin of flanks smooth with small conical tubercles; a small, horn-like prominent tubercle on the edge of upper eyelid; clear supratympanic fold curving posteroventrally from posterior corner of eye to a level above insertion of arm; a discontinuous “X” shaped ridge on the mid-dorsum and two discontinuous dorsolateral ridges on two sides at the mid-back; sparse tubercles on the dorsal shank and thigh; ventral surface smooth; several tubercles on posterior hindlimbs; small pectoral gland closer to axilla; a single femoral positioned subequally distant from knees and cloaca on posterior surface of each thigh. Coloration of holotype in life. Dorsal surface of body olive-brown with an inverted brown triangular marking between eyes; an “X” shaped marking on the mid-dorsum. Forearms and hindlimbs with brown transverse bands. Supratympanic fold with a continuous white line; a dark vertical band below the eye, from the inferior margin of the eye to the upper lip. Ventral surface of throat and chest grayish brown with brown patches, an indistinct longitudinal stripe on surface of throat; a pair of dark brown longitudinal stripes scattered on surface of lateroventral flanks; ventral surface of body white with brown patches; ventral surface of limbs grayish brown with dark brown spots and blotches; ventral surfaces of hands and ventral surfaces of feet brown, tips of digits pale-brown; metacarpal tubercle and metatarsal tubercle reddish. Pectoral gland and femoral gland white. Iris yellowish brown. Coloration of holotype in preservative. Olive-brown faded to greyish brown dorsally. Triangular marking between eyes, “X” shaped marking on the mid-dorsum, transverse bands on dorsal forearms and hind limbs became indistinct. Color of ventral surface faded to greyish white all bands and spots became indistinct. Variation and sexual dimorphism. Measurement data of type series are listed in Table 5 . Females ( SVL 36.3–45.8 mm ) are larger than males ( SVL 33.2–35.3 mm ). Presence of villiform black nuptial spines on the dorsal surface of the first and second fingers in adult males. Body coloration is quite variable, light brown, reddish brown, olive brown or dark brown dorsally ( Figure 5 ) . Distribution and ecology. Currently, Boulenophrys yingdeensis sp. nov. is known only from Shimentai Nature Reserve, Guangdong , China , and sympatric with B. shimentaina . All individuals were found in evergreen secondary forest, near lowland streams and nearbyleaf litter at elevations between 300– 400 m . Males perch and call on plant leaves from April to June, suggesting their breeding season corresponds to this period. Females were found on the forest floor and tadpoles were not observed.