Bioacoustics of Gracixalus guangdongensis

Description of holotype . Adult male, dorsoventrally compressed; head width almost equal to head length (HDW:HDL 1.01); snout triangularly pointed in dorsal view, projecting beyond margin of the lower jaw; canthus rostralis distinct and rounded; loreal region oblique and concave; nostrils oval, significantly protuberant, closer to tip of snout than to the eye; interorbital region flat, interorbital distance slightly broader than internasal distance (IOD:IND 1.03); eye large, horizontal diameter slightly smaller than snout length (ED:SL 0.83); pupil horizontal; tympanum distinct, tympanic rim weakly developed; tympanum diameter smaller than half of eye diameter (TYD:ED 0.41); supratympanic fold distinct, extending from the posterior corner of the eye above insertion of arm; vomerine teeth absent; tongue deeply notched behind; a single subgular vocal sac present. Forelimb moderately robust; forearm and hand relatively long (FAHL:SVL 0.48), hand significantly longer than forearm (HAL:SVL 0.46); fingers dorsoventrally compressed, webbing rudimentary, with narrow lateral fringes on outer edge of all fingers; relative finger length I <II <IV <III; tips of all fingers with well expanded discs with distinct transverse circum-marginal grooves; disc of third finger large, its width significantly broader than width of distal phalanx of finger III (FDW III:FPW III 2.1), and slightly larger than tympanum diameter (FDW III:TYD 1.24); subarticular tubercles markedly elevated and prominent, rounded, one on finger I and II, two on finger III and IV; supernumerary tubercles present; prepollex distinct, oval; a nuptial pad with barely visible minute granules present on the first finger; nuptial pad on dorsolateral surface of first finger strongly dilated, extending from hand base to level of subarticular tubercle; several tubercles on palmar, the outer palmar tubercle largest. Hindlimb long (TIBL:SVL 0.47, FTL:SVL 0.69); tibio-tarsal articulation reaching the middle of the eye when hindlimb adpressed along the side of the body; heels slightly overlapping when the flexed hindlimbs are held at right angles to the body axis; toes moderately long, relative toe lengths I <II <III <V <IV; tips of toes with expanded discs with distinct transverse circum-marginal grooves; discs of toes smaller than those of fingers; subarticular tubercles distinct, rounded, one on finger I and II, two on finger III and V, three on finger IV; webbing formula I (2), (2½) II (1½), (3) III (2), (3) IV (3), (2) V; sole smooth with small tubercles; inner metatarsal tubercle elongated, ellipsoid; outer metatarsal tubercle absent; inner tarsal fold absent. Skin of dorsal surface of head, body and limbs rough, sparsely scattered with tubercles; temporal region and corner of the mouth densely covered with large tubercles; pectoral and subgular skin smooth with barely visible granules; belly granular; anterior surface of hindlimbs smooth, posterior and ventral surface of hindlimbs with tubercles, relatively smooth; tarsal fold absent. Measurements of holotype (in mm). SVL 30.2, HDL 10.7, HDW 10.8, SL 5.1, IND 3.5, IOD 3.6, ED 4.2, TYD 1.7, TED 0.8, HAL 9.8, FAHL 14.6, TIBL 14.1, FTL 20.7, FDW III 2.1, FPW III 1.0, TDW IV 1.9, TPW IV 1.2. Coloration of holotype in life. Dorsal surface brown, with an inverse Y-shaped, dark-brown marking, starting at the interorbital region, bifurcating into two branches on shoulder, extending posteriorly; dorsal surface of fore- and hindlimbs with dark brown transverse bands; several faint, small, black blotches scattered on ventrolateral region of flanks and on ventral surface; ventral surface of hindlimbs, throat and chest creamy white, surface of belly and forelimbs light brown; iris brown, pupil black. Coloration of holotype in preservative. Color of dorsal surface dark grey; ventral surface of head and body greyish white; the Y-shaped marking and transverse black bands, blotches on the ventrolateral region of the flanks and the ventral surface are greyish black, and more distinct than in life. Variation. The external morphology and color pattern of all paratypes corresponds to that of the holotype , except for the following characters: background color of dorsal surface of head, body and limbs is beige in SYS a004671 (the single juvenile), SYS a004672 and SYS a004686 (vs. brown in holotype SYS a005724); adult males collected in the breeding season (SYS a004672–4673, 4685, 4687–4689, 4983, 4902–4904, 5724, 5750–5751, 5773–5775) possess nuptial pads on the dorsal surface of first finger, but the single specimen collected in the nonbreading season (SYS a005241) has barely visible nuptial pads; specimens from MNK and NNR have a smaller body size than specimens from DWL, measurement and body proportions of the type series specimens of G. guangdongensis sp. nov. are given in Table 3. Comparisons. Gracixalus guangdongensis sp. nov. was assigned with strong node supporting values ( 0.82 in BI and 86% in ML) to Clade I in our phylogenetic tree containing G. jinxiuensis , G. jinggangensis , G. sapaensis , G. nonggangensis and G. waza . The new species differs from G. jinxiuensis by the relatively larger body size, SVL 26.1–34.7 mm in adult males, 34.9–35.4 mm in adult females (vs. 24.2–26.3 mm in males, 28–29.2 mm in females of G. jinxiuensis ), more developed webbing (formula of webbing of the fourth toes in (3) IV (3)) (vs. (3½) IV (3½) in G. jinxiuensis ), ventral surface of hand, foot and palm smooth with sparse small tubercles (vs. rough with dense large tubercles in G. jinxiuensis ), tibio-tarsal articulation reaching the middle of the eye (vs. reaching the posterior corner of eye the in G. jinxiuensis ), and having heels overlapping when the flexed hindlimbs are held at right angles to the body axis (vs. just meeting in G. jinxiuensis ); from G. jinggangensis by the relatively larger body size in females, SVL 34.9–35.4 mm (vs. 31.6 mm in G. jinggangensis ), presence of nuptial pad on the first finger of male (vs. on first and second fingers of male G. jinggangensis ), and heels overlapping when the flexed hindlimbs are held at right angles to the body axis (vs. just meeting in G. jinggangensis ); from G. sapaensis by the presence of a distinct tympanum (vs. indistinct in G. sapaensis ), presence of a single vocal sac (vs. paired vocal sac in in G. sapaensis ), brown to beige background color of the dorsal surface of head, body and limbs in life (vs. greyish to reddish brown in G. sapaensis ), creamy white to light brown ventral surface (vs. venter pink in G. sapaensis ), and rough dorsal surface (vs. nearly smooth in G. sapaensis ); from G. nonggangensis by the more developed webbing (formula of webbing of the fourth toes in (3) IV (3)) (vs. (3½) IV (3½) in G. nonggangensis ), ventral surface of hand, foot and palm smooth with sparse small tubercles (vs. rough with dense large tubercles in G. nonggangensis ), presence of nuptial pad on the first finger of males (vs. absent in G. nonggangensis ), presence of rudimentary finger webbing (vs. finger webbing absent in G. nonggangensis ), tibio-tarsal articulation reaching the middle of the eye (vs. reaching the tip of the snout in G. nonggangensis ), and brown to beige background color of the dorsal surface of the head, body and limbs in life (vs. yellowish olive in G. nonggangensis ); from G. waza by the relatively small body size in adult females, SVL 34.9–35.4 mm (vs. 37.6 mm in G. waza ), brown to beige background color of the dorsal surface of the head, body and limbs in life (vs. greyish green to moss-green in G. waza ), and rough dorsal skin with tubercles (vs. nearly smooth dorsal skin in G. waza ). Gracixalus guangdongensis sp. nov. can also be easily distinguished from the six members of Clade II ( Fig 2 ). It differs from G. seesom by the relatively larger body size, SVL 26.1–34.7 mm in adult males, 34.9–35.4 mm in adult females (vs. 21.6–23 mm in males and 23.2–25.4 mm in female G. seesom ), presence of nuptial pads on the first finger of males (vs. absence in male G. seesom ), and rough dorsal skin with tubercles (vs. nearly smooth dorsal skin in G. seesom ); from G. quangi by the relatively larger body size, SVL 26.1–34.7 mm in adult males, 34.9–35.4 mm in adult females (vs. 21.4–22.9 mm in males and 26.8–27.3 mm in females of G. quangi ), absence of spines on the upper eyelid (vs. presence in G. quangi ), absence of a tibiotarsal projection (vs. presence in G. quangi ), and brown to beige background color of the dorsal surface of the head, body and limbs in life (vs. greenish to brownish green in G. quangi ); from G. gracilipes by the brown to beige background color of dorsal surface of head, body and limbs and the presence of inverse Y-shaped dark brown marking on back in life (vs. transparent green with Xshaped brown markings in G. gracilipes ), absence of a white patch under the eye to the tympanum (vs. presence in G. gracilipes ), and absence of spines on upper eyelid (vs. presence in G. gracilipes ); from G. supercornutus by the brown to beige background color of the dorsal surface (vs. yellowish green in G. supercornutus ), the absence of a white patch extending from under the eye to the tympanum (present in G. supercornutus ), absence of spines on upper eyelid (vs. present in G. supercornutus ), absence of a tibiotarsal projection (vs. present in G. supercornutus ); from G. quyeti by having distinct supratympanic fold (vs. indistinct in G. quyeti ), brown to beige background color of dorsal surface of head, body and limbs (vs. brown to moss-green in G. quyeti ); from G. lumarius by the relatively smaller body size, SVL 26.1–34.7 mm in adult males, 34.9–35.4 mm in adult females (vs. 38.9–41.6 mm in males and 36.3 mm in female of G. lumarius ), distinct tympanum and supratympanic fold (vs. indistinct in G. lumarius ), skin on dorsal surface of head, body rough, sparsely scattered with brown tubercles (vs. dorsum with distinctive dense network of white conical tubercles in adult males in G. lumarius ), venter creamy white to greyish white (vs. venter pink in G. lumarius ), single vocal sac in males (vs. a pair of vocal sacs in G. lumarius ). The new species also differs from the two congeners for which comparative molecular material is not available. The new species differs from Gracixalus megdogensis by the presence of rudimentary finger webbing (vs. absence of finger webbing in G. megdogensis ), tibio-tarsal articulation reaching the middle of the eye (vs. reaching the anterior corner of the eye in G. megdogensis ), rough dorsal skin with tubercles (vs. smooth dorsal skin in G. megdogensis ), and brown to beige background color of dorsal surface of head, body and limbs in life (vs. green in G. megdogensis ); from G. carinensis by the relatively smaller body size in females, SVL 34.9–35.4 mm (vs. 38 mm in G. carinensis ), more developed webbing (webbing formula of the fourth toe (3) IV (3)) (vs. webbing formula (2½) IV (2) in G. carinensis ), and snout length slightly longer than diameter of eye (SL> ED) (vs. snout length shorter than diameter of eye (SL <ED) in G. carinensis ). Gracixalus guangdongensis sp. nov. further differs from Kurixalus ananjevae (Matsui & Orlov) , which is morphologically and likely molecularly similar to the Gracixalus species of Clade II (i.e. Clade I in this study) according to Rowley et al . 2011 , by the smaller body size in females, SVL 34.9–35.4 mm (vs. 43.0 mm in female K. ananjevae ), dark grey dorsum in preservation (vs. pinkish gray in preservation in K. ananjevae ), snout length slightly longer than diameter of eye (SL> ED) (vs. snout length equal to or shorter than diameter of eye (SL ≤ ED) in K. ananjevae ), and more developed webbing (webbing formula of the fourth toes is (3) IV (3)) (vs. (3) IV (2½) in K. ananjevae ). Gracixalus guangdongensis sp. nov. further differs from the small- and medium-sized rhacophorids occurred in southern China and northern Vietnam : the new species differs from Liuixalus calcarius Milto, Poyarkov, Orlov & Nguyen , L. feii Yang, Rao & Wang , L. jinxiuensis Li, Mo, Jiang, Xie & Jiang , L. hainanus (Liu and Wu) , L. ocellatus (Liu & Hu) , L. romeri (Smith) , L. shiwandashan Li, Mo, Jiang, Xie & Jiang , Raorchestes longchuanensis (Yang and Li) and Ra. menglaensis (Kou) by the significantly larger body size, SVL 26.1–34.7 mm in adult males, 34.9–35.4 mm in adult females (vs. SVL values not-overlaping and less than 21.0 mm in males and 22.0 mm in females) and the more developed webbing on toes (vs. very poorly-developed webbing on the toes); from Philautus abditus Inger, Orlov & Darevsky by the presence of nuptial pads in males and a distinct tympanum (vs. nuptial pads absent and tympanum completely hidden under skin); from Buergeria oxycephala (Boulenger) by the significantly smaller body size and the absence of vomerine teeth (vs. SVL 34–38 in males and 60–68 in females, vomerine teeth present); from Thelerdema albopunctatum (Liu & Hu), T. annae Nguyen, Pham, Nguyen, Ngo & Ziegler, T. bicolor (Bourret), T. corticale (Boulenger), T. gordoni Taylor, T. lateriticum Bain, Nguyen & Doan and T. rhododiscus (Liu &Hu) by the absence of skin ridges and large warts on dorum (vs. presence of irregular skin ridges and large warts on dorsum in T. bicolor , T. corticale , T. gordoni ; and presence of irregular skin ridges on dorsum in T. albopunctatum and T. rhododiscus ), presence of a single vocal sac (vs. paired vocal sac in T. albopunctatum , and absent in T. annae , T. lateriticum and T. rhododiscus ), and the absence of vomerine teeth (vs. vomerine teeth present in T. bicolor and T. corticale ); from Kurixalus bisacculus (Taylor) by the presence of a single vocal sac in males and fingers with rudiment of web (vs. paired vocal sac in males and fingers without web), from K. lenquanensis Yu, Wang, Hou, Rao & Yang and K. odontotarsus (Ye and Fei) by the absence of vomerine teeth and absence of conical tubercles on posterolateral surface of forearms and tibia (vs. vomerine teeth present, distinct conical tubercles on posterolateral surface of forearms and tibia present); from Chiromantis doriae (Boulenger) by having rough dorsal skin with tubercles and a “Y-shaped” marking on the back (vs. smooth dorsal skin with granules and five longitudinal stripes on the back); from Feihyla fuhua Fei, Ye & Jiang , F. palpebralis (Smith) and F. vittata (Boulenger) by the single vocal sac in males (vs. paired vocal sac in males); from Polypedates braueri (Vogt) , Po. impresus Yang , Po. megacephalus Hallowell and Po. mutus (Smith) by the absence of vomerine teeth and presence of rudiment of web on fingers (vs. vomerine teeth present and webbing on fingers absent); from Rhacophorus dorsoviridis Bourret , Rh. hoanglienensis Orlov, Lathrop, Murphy & Ho , Rh. laoshan Mo, Jiang, Xie & Ohler , Rh. larissae Ostroshabov, Orlov & Nguyen , Rh. leucofasciatus Liu & Hu , Rh. minimus Rao, Wilkinson & Liu , Rh. nigropunctatus Liu, Hu & Yan , Rh. pinglongensis Mo, Chen, Liao & Zhou , Rh. puerensis (He) , Rh. rhodopus Liu & Hu , Rh. yaoshanensis Liu & Hu and Rh. yinggelingensis Chou, Lau , and Chan by the absence of vomerine teeth (vs. vomerine teeth present), rudimentary webbing on the fingers (vs. welldeveloped webbing on the fingers in Rh. leucofasciatus and Rh. rhodopus ), heels overlapping when the flexed hindlimbs are held at right angles to the body axis (vs. heels not meeting in Rh. dorsoviridis and Rh. nigropunctatus ; heels just meeting in Rh. yaoshanensis and Rh. yinggelingensis ), presence of nuptial pads on the first finger (vs. presence of nuptial pads on first and second fingers in Rh. puerensis ), absence of tibiotarsal projection (vs. tibiotarsal projection present in Rh. hoanglienensis and Rh. larissae ). Advertisement calls. The advertisement calls of Gracixalus guangdongensis sp. nov. , are compared to all species nested in Clade I of our phylogenetic analyses for which acoustic data is available, G. jinggangensis and G. nonggangensis . All advertisement calls of the four male G . guangdongensis sp. nov. recorded (two from DWL, one from MNK and one from NNR) are similar in structure, composed of an introductory note followed by a single ‘click’ note ( Fig. 4 , A–D). The call of the new species therefore differs from that of G. jinggangensis (which has an introductory note followed by two or three click notes, Fig. 4E ) and G. nonggangensis (which has an introductory note followed by 39–71 click notes, Fig. 4F ). Moreover, the call of G. guangdongensis sp. nov. differs from G. jinggangensis in the duration of the introductory note (0.40 ± 0.05 vs. 0.20 ± 0.02 in two click notes form and 0.16 ± 0.01 in three click notes form). Measurements of the advertisement calls of the three species are listed in Table 4 .

Ecology and behavior. The new species appears to be restricted to bamboo forest with evergreen broad-leaf forest ( Fig. 5 ), at elevations of approximately 600 m at MNK, 1000–1600 m at DWL, and 1000–1200 m at NNR and MNR. The single juvenile (SYS a004671) from DWL was collected from a shrub ( Litsea sp.) adjacent to a stream, 50 m away from bamboo forest. Num. DWL MNK NNR ALL SEX Female (n=2) Male (n=9) Male (n=3) Male (n=5) Male (n=17) SVL 34.9̅35.4 (35.2) 28.7̅34.7 (30.9 ± 2.0) 26.4̅28.0 (28.0 ± 1.7) 28.5̅33.1 (30.0 ± 1.7) 26.4̅34.7 (30.1 ± 2.1) HDL 12.0̅12.3 (12.2) 10.0̅11.9 (10.9 ± 0.7) 9.0̅10.2 (9.7 ± 0.6) 10.0̅11.4 (10.6 ± 0.5) 9.0̅11.9(10.6 ± 0.7) HDW 12.2̅12.4 (12.3) 10.0̅12.0 (10.9 ± 0.7) 9.3̅10.6 (10.0 ± 0.7) 10.1̅11.7(10.7 ± 0.6) 9.3̅12.0(10.7 ± 0.7) SL 5.2̅5.3 (5.3) 4.1̅5.3 (4.8 ± 0.3) 3.7̅4.5 (4.1 ± 0.4) 4.6̅5.1 (4.8 ± 0.2) 3.7̅5.3 (4.7 ± 0.4) IND 3.8̅4.1 (4.0) 3.0̅3.7 (3.4 ± 0.2) 3.0̅3.4 (3.2 ± 0.2) 3.2̅3.6 (3.3 ± 0.2) 3.0̅3.7 (3.4 ± 0.2) IOD 3.9̅4.0 (4.0) 3.2̅4.0 (3.6 ± 0.2) 3.3̅3.6 (3.4 ± 0.2) 3.3̅3.8 (3.5 ± 0.2) 3.2̅4.0 (3.6 ± 0.2) ED 4.7̅4.8 (4.8) 3.7̅4.6 (4.1 ± 0.3) 3.4̅4.0 (3.7 ± 0.3) 3.8̅4.5 (4.1 ± 0.3) 3.4̅4.6 (4.1 ± 0.3) TYD 1.7̅1.8 (1.8) 1.5̅1.7 (1.6 ± 0.1) 1.4̅1.5 (1.5 ± 0.1) 1.5̅1.8 (1.6 ± 0.1) 1.4̅1.8 (1.6 ± 0.1) TED 0.8̅0.9 (0.9) 0.8̅0.9 (0.9 ± 0.1) 0.9 0.8̅0.9(0.9 ± 0.1) 0.8̅0.9(0.9 ± 0.1) FAHL 16.3̅17.0 (16.7) 13.1̅15.9 (14.9 ± 1.0) 12.9̅14.9 (13.8 ± 1.0) 14.5̅16.1 (15.1 ± 0.6) 12.9̅16.1 (14.7 ± 1.0) HAL 10.2̅11.3 (10.8) 8.7̅10.6 (9.6 ± 0.6) 8.4̅9.5 (8.8 ± 0.6) 8.9̅10.6 (9.4 ± 0.7) 8.4̅10.6 (9.4 ± 0.7) TIBL 14.8̅16.8 (15.8) 14.1̅16.0 (14.9 ± 0.7) 13.0̅14.1 (13.5 ± 0.5) 14.0̅15.3 (14.8 ± 0.5) 13.0̅16.0 (14.6 ± 0.8) FTL 22.6̅23.6 (23.1) 19.0̅22.0 (20.6 ± 1.1) 17.5̅20.0 (18.5 ± 1.3) 20.0̅21.3 (20.7 ± 0.6) 17.5̅22.0 (20.3 ± 1.3) FDW III 1.8 1.5̅2.1 (2.0 ± 0.2) 1.2̅1.8 (1.4 ± 0.2) 1.6̅1.9 (1.7 ± 0.1) 1.2̅2.1 (1.8 ± 0.3) FPW III 0.8̅0.9 (0.9) 0.7̅1.0(0.9 ± 0.1) 0.6̅0.8(0.7 ± 0.1) 0.8̅0.9(0.8 ± 0.1) 0.6̅1.0(0.9 ± 0.1) TDW IV 1.5̅1.8 (1.7) 1.5̅1.9 (1.7 ± 0.1) 1.0̅1.6 (1.2 ± 0.3) 1.2̅1.6 (1.4 ± 0.2) 1.0̅1.9 (1.5 ± 0.3) TPW IV 1.0̅1.1 (1.1) 1.0̅1.2 (1.1 ± 0.1) 0.7̅1.0 (0.8 ± 0.2) 0.7̅1.0 (0.9 ± 0.1) 0.7̅1.2 (1.0 ± 0.2) HDL:SVL 0.34̅0.35 (0.35) 0.34̅0.37 (0.35 ± 0.01) 0.34̅0.35 (0.35 ± 0.01) 0.34̅0.37 (0.35 ± 0.01) 0.34̅0.37 (0.35 ± 0.01) HDW:SVL 0.35 0.34̅0.37 (0.35 ± 0.01) 0.35̅0.37 (0.36 ± 0.01) 0.35̅0.37 (0.36 ± 0.01) 0.34̅0.37 (0.36 ± 0.01) HDW:HDL 1.01̅1.02 (1.01) 0.99̅1.01 (1.00 ± 0.01) 1.03̅1.05 (1.04 ± 0.01) 0.99̅1.03 (1.02 ± 0.02) 0.99̅1.05 (1.01 ± 0.02) SL:HDL 0.43 0.41̅0.48 (0.44 ± 0.02) 0.41̅0.44 (0.43 ± 0.01) 0.43̅0.48 (0.46 ± 0.02) 0.41̅0.48 (0.44 ± 0.02) IND:HDW 0.31̅0.33 (0.32) 0.30̅0.33 (0.32 ± 0.01) 0.32 0.31̅0.32 (0.31 ± 0.01) 0.30̅0.33 (0.31 ± 0.01) IOD:HDW 0.31̅0.33 (0.32) 0.31̅0.36 (0.33 ± 0.02) 0.33̅0.35 (0.34 ± 0.01) 0.32̅0.34 (0.33 ± 0.01) 0.31̅0.36 (0.33 ± 0.01) ED:SL 0.90̅0.91 (0.91) 0.80̅0.92 (0.87 ± 0.04) 0.89̅0.92 (0.90 ± 0.01) 0.79̅0.90 (0.85 ± 0.05) 0.79̅0.92 (0.87 ± 0.04) ED:HDL 0.39 0.37̅0.39 (0.38 ± 0.01) 0.38̅0.39 (0.39 ± 0.01) 0.37̅0.39 (0.39 ± 0.01) 0.37̅0.39 (0.38 ± 0.01) TYD:HDL 0.14̅0.15 (0.15) 0.14̅0.16 (0.15 ± 0.01) 0.14̅0.16 (0.15 ± 0.01) 0.14̅0.16 (0.15 ± 0.01) 0.14̅0.16 (0.15 ± 0.01) FAHL:SVL 0.47̅0.48 (0.48) 0.46̅0.51 (0.48 ± 0.02) 0.48̅0.50 (0.49 ± 0.01) 0.49̅0.51 (0.50 ± 0.01) 0.46̅0.51 (0.49 ± 0.02) HAL:SVL 0.29̅0.32 (0.31) 0.30̅0.33 (0.31 ± 0.01) 0.30̅0.32 (0.31 ± 0.01) 0.30̅0.32 (0.31 ± 0.01) 0.30̅0.33 (0.31 ± 0.01) TIBL:SVL 0.42̅0.47 (0.45) 0.45̅0.51 (0.48 ± 0.02) 0.46̅0.51 (0.48 ± 0.02) 0.46̅0.51 (0.49 ± 0.02) 0.45̅0.51 (0.49 ± 0.02) FTL:SVL 0.65̅0.67 (0.66) 0.63̅0.71 (0.67 ± 0.03) 0.63̅0.68 (0.66 ± 0.03) 0.64̅0.71 (0.69 ± 0.03) 0.63̅0.71 (0.67 ± 0.03) FDW III:FPW III 2.00̅2.25 (2.13) 2.00̅2.14 (2.07 ± 0.06) 2.07̅2.19 (2.14 ± 0.07) 2.07̅2.25 (2.14 ± 0.06) 2.00̅2.25 (2.10 ± 0.07) TDW IV:TPW IV 1.50̅1.64 (1.57) 1.50̅1.58 (1.53 ± 0.03) 1.55̅1.63 (1.58 ± 0.04) 1.58̅1.64 (1.61 ± 0.02) 1.50̅1.64 (1.56 ± 0.05) Species Voucher number Temperature (Û ) n Number of notes Call duration (s) Peak frequency (kHz) Frequency range (kHz) Duration of the introductory note (s) Duration of the click note (s) Interval between the introductory note and the first click note (s) DWL 1 SYS a004672 17 Û 27 2 0.65 ± 0.02 2.4 2.0–2.8 0.47 ± 0.02 0.04 ± 0.01 (n=27) 0.14 ± 0.01 DWL 2 SYS a004685 18 Û 10 2 0.50 ± 0.02 2.6 2.3–2.9 0.36 ± 0.02 0.04 ± 0.01 (n=10) 0.10 ± 0.01 G.guangdongensis sp.nov. MNK SYS a004902 21 Û 11 2 0.50 ± 0.02 2.8 2.3–3.0 0.37 ± 0.02 0.05 ± 0.01 (n=11) 0.08 ± 0.01 NNR SYS a005750 18 Û 24 2 0.51 ± 0.02 4 2.0–2.8 0.36 ± 0.02 0.05 ± 0.01 (n=24) 0.11 ± 0.01 ALL / / 72 2 0.56 ± 0.07 2.5 ± 0.1 2.0–3.0 0.40 ± 0.05 0.05 ± 0.01 (n=72) 0.11 ± 0.02 10 3 0.50 ± 0.02 2.6 2.0–3.1 0.20 ± 0.02 0.06 ± 0.01 (n=20) 0.09 ± 0.01 G.jinggangensis SYS a004811 17.6 Û 0.05 ± 0.01 10 4 0.60 ± 0.01 2.6 2.0–3.1 0.16 ± 0.01 (n=30) 0.08 ± 0.01 G.nonggangensis SYS a005867 17.0 Û 12 40–72 15.06 ± 4.17 2.2 1.6–2.7 0.49 ± 0.05 0.12 ± 0.04 (n=266) 0.19 ± 0.03 According to our observations, the main breeding season of the DWL population begins in April and lasts at least three months, as males with distinct nuptial pads and calling males were observed at these times. A small number of males were also heard calling in August. The female collected in April (SYS a004686) has a distinct fat body and visible fallopian tubes, while a female collected in June (SYS a004984) has barely visible fallopian tubes. The male holotype was found calling on a bamboo stem about 1.7 m above the ground on 9 April 2017 at DWL ( Fig. 3A ), at the same time, two individuals were found sitting on another bamboo pole (male above and female below) about 2 m above the ground ( Fig. 6A ). Eggs with jelly coating were discovered after splitting apparently air-tight bamboo joints on 28 June 2016 . On 17 April 2017 at DWL, eggs were found on the walls of an openended bamboo joint half full of water where a female individual was found inside the night before ( Fig. 6 , B & C). Eggs and embryos were also found in another water-filled bamboo joint the next day ( Fig. 6D ). Other details of the ecology and behavior of the new species remains unknown.