A new species of Bush frog (Anura, Rhacophoridae, Raorchestes) from southeastern Yunnan, China Author Huang, Junkai Key Laboratory of Ecology of Rare and Endangered Species and Environmental Protection (Guangxi Normal University), Ministry of Education, Guilin 541004, China Author Liu, Xiao Long Guangxi Key Laboratory of Rare and Endangered Animal Ecology, Guangxi Normal University, Guilin 541006, Guangxi, China Author Du, Lingyun https://orcid.org/0000-0002-5761-4017 Key Laboratory of Ecology of Rare and Endangered Species and Environmental Protection (Guangxi Normal University), Ministry of Education, Guilin 541004, China Author Bernstein, Justin M. https://orcid.org/0000-0002-5249-3340 Key Laboratory for Conserving Wildlife with Small Populations in Yunnan, Southwest Forestry University, Kunming 650224, Yunnan, China Author Liu, Shuo https://orcid.org/0000-0001-7825-3006 Department of Biological Sciences, Rutgers University Newark, 195 University Ave Newark, NJ 07102, USA Author Yang, Yun Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650223, China Author Yu, Guohua Key Laboratory of Ecology of Rare and Endangered Species and Environmental Protection (Guangxi Normal University), Ministry of Education, Guilin 541004, China yugh2018@126.com Author Wu, Zhengjun Key Laboratory of Ecology of Rare and Endangered Species and Environmental Protection (Guangxi Normal University), Ministry of Education, Guilin 541004, China wu_zhengjun@aliyun.com text ZooKeys 2023 2023-02-28 1151 47 65 http://dx.doi.org/10.3897/zookeys.1151.95616 journal article http://dx.doi.org/10.3897/zookeys.1151.95616 1313-2970-1151-47 4AACB0A62800465DA10FE69B4CEE3E05 58DA6C4192BB58FAB832DC3540943D93 Raorchestes malipoensis sp. nov. Fig. 4 Pseudophilautus gryllus " Pseudophilautus gryllus " ( Li et al. 2009 ). Raorchestes gryllus " Raorchestes gryllus " ( Biju et al. 2010 ). Holotype. GXNU 000339, adult male, collected from Malipo County, Yunnan Province ( 23.182°N , 104.78°E , elevation 1496 m) on 22 July 2020 by Shuo Liu. Paratypes. SWFU 3110, SWFU 3113, SWFU 3114, SWFU 3116, GXNU 000338, GXNU 000341 (six adult males), SWFU 3111, SWFU 3112, GXNU 000340, GXNU 000342 (four adult females), collected at the same locality as the holotype on 22 July 2020 by Xiaolong Liu and Shuo Liu. Diagnosis. The genus Raorchestes is a group of small frogs, diagnosed primarily on the basis of an adult snout-vent length between 15 and 45 mm; vomerine teeth absent; large gular pouch transparent while calling; nocturnally active; direct development without free-swimming tadpoles in all species for which the development is known ( Biju et al. 2010 ). Although the mode of development in the new species remains unknown, R. malipoensis sp. nov. is placed in the genus Raorchestes due to the combination of following characters: small body size, vomerine teeth absent, single translucent external subgular vocal sac present, and tips of all fingers and toes expanded into discs with circum-marginal grooves. The new species is distinguished from geographically and molecularly relevant congeners by the following combination of characters: (1) very small body size (males SVL 14.6-17.7 mm, n = 7; females SVL 18.3-19.3 mm, n = 4); (2) head wider than long; (3) tympanum small, supratympanic fold distinct; (4) tips of all fingers and toes yellow; (5) webbing formula (I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V); (6) inner and outer metacarpal tubercle indistinct; (7) heels not meeting when limbs held at right angles to body; (8) tibiotarsal articulation reaching anterior border of eye when hindlimb is stretched alongside of body; (9) iris golden brown; (10) nuptial pad small and milky white; (11) inner metatarsal tubercle rounded, outer metatarsal tubercle absent; (12) fingers and toes having lateral dermal fringe; and (13) interorbital distance larger than eye horizontal diameter. Description of the holotype. Adult male (Fig. 4 ), body size small (SVL 17.7 mm); head wider than long (HL 6.4 mm; HW 6.8 mm); top of head relatively flat; snout rounded in profile, projecting beyond lower jaw; snout length almost equal to interorbital distance at narrowest point (SL 2.6 mm; IOS 2.6 mm); the canthus rostralis rounded, loreal region slightly concave; tympanum small (TD 1.5 mm); internarial distance wider than maximum width of upper eyelid (INS 2.1 mm; UEW 1.3 mm); nostril slightly closer to tip of snout than to anterior corner of eyes; tongue pyriform, with a deep notch at posterior tip; vomerine teeth absent; pineal ocellus absent; eyes moderately large (EHD 2.6 mm) and protruding, pupil horizontal; supratympanic fold distinct, from posterior corner of eye to above insertion of arm. Figure 4. Holotype (GXNU 000339) of Raorchestes malipoensis sp. nov. in life. Forelimbs fairly robust (FAHL 8.2 mm); relative finger lengths: I <II <IV <III, tips of all four fingers expanded into discs with circum-marginal grooves; all fingers with lateral dermal fringes on both sides; subarticular tubercles distinct, rounded; supernumerary tubercles absent; no webbing between fingers; inner and outer metacarpal tubercle indistinct; nuptial pad is small and milky white on dorsal surface of the first finger. Foot long and relatively robust (TFL 11 mm), longer than tibia length (TBL 8.8 mm); relative toe lengths: I <II <V <III <IV; tips of toes with discs having circum-marginal grooves, toe discs smaller than finger discs; all toes with lateral dermal fringes on both sides; subarticular tubercles distinct, rounded; supernumerary tubercles absent; webbing formula (I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V); inner metatarsal tubercle rounded, outer metatarsal tubercle absent. Dorsal surfaces of head, body, forelimbs, thighs, and tibia rough with small granules; upper eyelid with several small granules; throat, chest, and ventral surfaces of forelimbs smooth; abdomen, ventral side of thigh, and area around vent with granules; dorsolateral folds absent. Coloration of holotype in life. For coloration of the holotype in life see Fig. 4 . Dorsal surface beige, with pale brown band between eyes; dorsal surface with a dark brown X-shaped marking; pale brown interorbital rectangle between eyes; upper and lower lips with white and black dots; supratympanic fold pale brown; iris golden brown; dorsal parts of arms and legs with dark brown crossbars that align; crotch with a distinct black patch bordering large creamy white plaque below the black patch near the groin; dorsal thigh beige with one brown crossbar when leg is bent in resting position; ventral surface body and beige, and area around vent with small black spots; discs of fingers and toes yellow. Coloration in alcohol. After preservation in alcohol, the general pattern did not change. Dorsal color changed to grayish brown, the blotches or spots blackish brown, discs on the fingers become pale gray similar to the body color, ventral side become whiter (Fig. 5 ). Figure 5. Holotype (GXNU 000339) of Raorchestes malipoensis sp. nov. in preservative, showing A dorsal view B ventral view C ventral view of hand D ventral view of foot. Etymology. The specific epithet is named for the type locality, Malipo County, Yunnan Province, China. We suggest "Malipo Bush Frog" as its English common name, and "Ma Li Po Guan Shu Wa ( 麻栗坡灌树蛙)" as its Chinese common name. Distribution. Currently known from the type locality, Malipo County (Fig. 1 ), Yunnan Province, China and Pac Ban, Tuyen Quang, in north of Vietnam. Variation. The measurements are given in Table 1 . GXNU 000338 has large black spots on dorsal side and GXNU000342 has distinctly darker ground color on dorsal side. Comparisons. Rather than comparing R. malipoensis sp. nov. to all known Raorchestes , we focus on our morphological comparison with phylogenetically closely related taxa and species without genetic data in adjacent countries (Table 5 ). Table 5. Comparison of R. malipoensis sp. nov. with phylogenetically closely related taxa or those with no genetic data in surrounding countries. "-" means unknown.

Species	R. malipoensis sp. nov.	R. menglaensis	R. parvulus	R. dulongensis	R. hillisi	R. huanglianshan	R. cangyuanensis	R. ghatei	R. rezakhani	R. annandalii	R. bombayensis	R. tuberohumerus	R. longchuanensis	R. andersoni
SVL of adult males (in mm)	14.6-17.7, n = 7	16.6-21.6, n = 14	-	15.0-19.0, n = 3	14.5-17.7, n = 3	17.0-19.6, n = 11	16.1-19.0 mm, n = 3	19.1-25.5, n = 9	18.8-19.0 mm, n = 4	-	30 mm, n = -	17.4-18.2 mm, n = 3	21.4-23.9 mm, n = 5	13.5-24.0 mm, n = 2
SVL of adult females (in mm)	18.3-19.3, n = 4	18.9-20.5, n = 2	23.6, n = 1	-	17.5, n = 1	21.5, n = 1	-	15.4-29.8, n = 13	-	17.0 mm, n = 1	-	-	-	-
SVL of adult (in mm)	14.6-19.3, n = 11	16.6-21.6, n = 18	23.6, n = 1	15.0-19.0, n = 3	14.5-17.7, n = 4	17.0-21.5, n = 4	16.1-19.0 mm, n = 3	15.4-29.8, n = 22	18.8-19.0 mm, n = 4	17.0 mm, n = 1	30 mm, n = -	17.4-18.2 mm, n = 3	21.4-23.9 mm, n = 5	13.5-24.0 mm, n = 2
IOS/EHD	IOS> EHD, or IOS = EHD	IOS> EHD	IOS <EHD	IOS <EHD	IOS <EHD	IOS> EHD, or IOS=EHD	IOS <EHD	-	IOS <EHD	IOS> EHD, or IOS = EHD	-	IOS> EHD, or IOS = EHD	IOS> EHD	IOS <EHD
HDW/HDL	HDW> HDL	HDW <HDL	HDW> HDL	HDW <HDL	HDW <HDL	HDW> HDL	HDW> HDL	HDW> HDL	HDW> HDL	HDW <HDL	-	HDW> HDL	HDW ≈ HDL	HDW> HDL
Tympanum	Distinct	Indistinct	Distinct	Distinct	Distinct	Distinct	Indistinct	Indistinct	Indistinct	Distinct	Indistinct	Indistinct	Distinct	Distinct
Nuptial pad	Small and milky white	white nuptial pad	-	Absent	Present	Present	Reddish nuptial pad	Absent	Absent	-	-	-	Present	-
Toe web	I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V	II 1 - 2 III 1 - 21/2 IV 21/2 - 1 V	Webbing present, medium	Rudimentary web	II 1 - 2 III 1-21/2 IV 21/2 - 1 V	II 1 - 2 III 1 - 2 - IV 2 - 1 V	Rudimentary web	I 2 - 2 II 2 - 21/2 III 2 - 3 IV 21/2 - 2 V	I2 - 2 II 13/4 - 2 III 11/2 - 3 IV 23/4 - 2 V	Rudimentary web	1/3 webbing	Rudimentary web	1/4 webbing	1/3 webbing
Lateral dermal fringe	Present	Absent	-	-	Present	Absent	Present	Present	Absent	Present	-	-	-	-
Disc color	Yellow	Not orange in life	-	Greyish or orange	-	Orange	Orange	-	Reddish or whitish	-	-	-	Reddish, orange, or whitish	orange
Inner metacarpal tubercle	Indistinct	Present	Present	Present	Indistinct	Indistinct	-	-	Absent	Present	-	-	Present	Present
Outer metacarpal tubercle	Indistinct	Present	Present	Present	Indistinct	Indistinct	-	-	Absent	Present	-	-	Present	Present
Inner metatarsal tubercle	Round	Present	Present	Round	Round	Round	Round	Round	Absent	Absent	-	Present	Present	Present
Outer metatarsal tubercle	Absent	Present	Absent	Absent	Absent	Absent	Absent	Absent	Absent	Absent	-	Absent	Absent	Absent
Relative toe lengths	I <II <V <III <IV	III ≈ V, or V> III	I <II <V <III <IV	I <II <V <III <IV	I<II<III<V<IV	I<II<III<V<IV	I<II<V<III<IV	I<II<V=III<IV	I <II <V <III <IV	I <II <V = III <IV	-	I <II ≤ V <III <IV	III ≈ V	I <II <III = V <IV
Range	Malipo, Yunnan, China and the north of Vietnam	Mengla, Yunnan, China	Indochina Peninsula and peninsular Malaysia	Gongshan, Yunnan, China	Menghai, Yunnan, China	Lvchun, Yunnan, China	Cangyuan, Yunnan, China	Western Ghats, India	Northeastern Bangladesh	Himalayas and northeastern India	Western Ghats, India	Western Ghats, India	Yunnan, China and Lai Chau,Vietnam	India, North Myanmar, Tibet and Yunnan, China

The new species differs from R. menglaensis by 1) tubercles absent along the outer side of the forearm and foot; (2) head wider than long; (3) tympanum distinct (TD 1.1-1.6 mm, n = 11); (4) webbing formula (I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V); (5) lateral dermal fringe present (6) inner and outer metacarpal tubercle indistinct; (7) outer metatarsal tubercle absent; and (8) relative toe lengths: I <II <V <III <IV (vs. a series of tubercles along the outer side of the forearm and foot; head length and head width are approximately the same; tympanum indistinct; webbing formula (II 1 - 2 III 1 - 21/2 IV 21/2-1 V); lateral dermal fringe present; inner and outer metatarsal tubercle present; outer metatarsal tubercle present; relative toe lengths: III ≈ V, or V> III). The new species differs from R. parvulus by (1) smaller female body size (females 18.3-19.3 mm, n = 4); (2) interorbital distance larger than eye horizontal diameter; and (3) inner and outer metacarpal tubercle indistinct; (vs. female 23.6 mm, n = 1; interorbital distance smaller than eye horizontal diameter; inner and outer metacarpal tubercle present). The new species differs from R. dulongensis by (1) head wider than long; (2) interorbital distance larger than eye horizontal diameter; (3) nuptial pad present; (4) yellow disc; and (5) inner and outer metacarpal tubercle indistinct (vs. head smaller than long; interorbital distance smaller than eye horizontal diameter; nuptial pad absent; greyish or orange disc; inner and outer metacarpal tubercle indistinct present). The new species differs from R. hillisi by (1) larger female body size (females 18.3-19.3 mm, n = 4); (2) head wider than long; (3) interorbital distance larger than eye horizontal diameter; (4) webbing formula (I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V); and (5) and relative toe lengths: I <II <V <III <IV (vs. female 17.5 mm, n = 1; head longer than wider; interorbital distance smaller than eye horizontal diameter; webbing formula (II 1-2 III 1-21/2 IV 21/2-1 V); relative toe lengths: I <II <III <V <IV). The new species differs from R. huanglianshan by (1) smaller female body size (females18.3-19.3 mm, n = 4); (2) lateral dermal fringe present; (3) yellow disc; (4) webbing formula (II 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V); and (5) relative toe lengths: I <II <V <III <IV (vs. female 21.5 mm, n = 1; lateral dermal fringe absent; orange disc; fingers and toes lacking lateral dermal fringe; webbing formula (II 1-2 III 1 - 2 - IV 2 - 1 V); relative toe lengths: I <II <III <V <IV). The new species differs from R. cangyuanensis by (1) interorbital distance larger than eye horizontal diameter; (2) nuptial pad small and milky white; and (3) yellow discs (vs. interorbital distance smaller than eye horizontal diameter; reddish nuptial pad at the base of first finger; orange disc). The new species differs from R. ghatei by (1) smaller body size (males 14.6-17.7 mm, n = 7; females18.3-19.3 mm, n = 4); (2) tympanum distinct (TD 1.1-1.6 mm, n = 11); (3) nuptial pad present; (4) webbing formula (I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V); and (5) relative toe lengths: I <II <V <III <IV (vs. males 19.1-25.5 mm, n = 9; females 15.4-29.8 mm, n = 13; tympanum indistinct; nuptial pad absent; webbing formula (I 2 - 2 II 2 - 21/2 III 2- 3 IV 21/2 - 2 V); relative toe lengths: I <II <V = III <IV). The new species differs from R. rezakhani by (1) smaller male body size (males 14.6-17.7 mm, n = 7); (2) interorbital distance larger than eye horizontal diameter; (3) tympanum distinct (TD 1.1-1.6 mm, n = 11); (4) nuptial pad present; (5) lateral dermal fringe present; (6) yellow disc; (7) inner and outer metacarpal tubercle indistinct; (8) inner metatarsal tubercle round; and (9) webbing formula (I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V) (vs. males 18.8-19.0 mm; interorbital distance smaller than eye horizontal diameter; tympanum indistinct; nuptial pad absent; lateral dermal fringe absent; reddish or whitish; inner and outer metacarpal tubercle absent; inner metatarsal tubercle absent; webbing formula (I2 - 2 II 13/4 - 2 III 11/2 - 3 IV 23/4 - 2 V). The new species differs from R. annandalii by (1) head wider than long; and (2) relative toe lengths: I <II <V <III <IV (vs. head longer than wide; relative toe lengths: I <II <V = III <IV). The new species differs from R. bombayensis by (1) smaller body size (males 14.6-17.7 mm, n = 7; females 18.3-19.3 mm, n = 4); (2) tympanum distinct (TD 1.1-1.6 mm, n = 11); and (3) webbing formula (I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V) (vs. 30 mm, n = 1; tympanum indistinct; 1/3 webbing between toes). The new species differs from R. tuberohumerus by (1) tympanum distinct (TD 1.1-1.6 mm, n = 11); and (2) relative toe lengths: I <II <V <III <IV (vs. tympanum indistinct; relative toe lengths: I <II ≤ V <III <IV). The new species differs from R. longchuanensis by (1) smaller male body size (males 14.6-17.7 mm, n = 7); (2) webbing formula (I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V); and (3) yellow disc (vs. males 21.4-23.9 mm, n = 5; 1/4 webbing between toes; reddish, orange, or whitish disc). The new species differs from R. andersoni by (1) interorbital distance larger than eye horizontal diameter; (2) webbing formula (I 2 - 2 II 2 - 2 III 2 - 3 IV 3 - 2 V); (3) yellow disc; and (4) relative toe lengths: I <II <V <III <IV (vs. interorbital distance smaller than eye horizontal diameter; 1/3 webbing between toes; orange disc; relative toe lengths: I <II <III = V <IV).