Zoosyst. Evol. 101 (2) 2025, 437-447 | DOI 10.3897/zse.101.142297 Gee BERLIN A new species of Rhacophorus (Anura, Rhacophoridae) from Xizang, China, with a revision of the distribution of R. bipunctatus Shiyang Weng*”, Xiaolong Liu***, Jianchuan Li*, Guohua Yu°®, Junkai Huang? 1 Institute of Plateau Biology of Xizang, Lhasa, 850030, China 2 Integrative Science Center of Germplasm Creation in Western China (CHONGQING) Science City, Biological Science Research Center, Southwest University, Chongqing, 400715, China 3 Key Laboratory of Freshwater Fish Reproduction and Development (Ministry of Education), School of Life Sciences, Southwest University, Chongqing, 400715, China 4 Xizang Museum of Natural Science, Lhasa, 850011, China 5 Key Laboratory of Ecology of Rare and Endangered Species and Environmental Protection (Guangxi Normal University), Ministry of Education, Guilin, 541004, China 6 Guangxi Key Laboratory of Rare and Endangered Animal Ecology, College of Life Science, Guangxi Normal University, Guilin, 541004, China https://zoobank. org/F7C7A5DF-O0BBA-4EE1-B3F8-336F62197FAB Corresponding authors: Junkai Huang (1198094210@qq.com); Guohua Yu (yugh2018@126.com) Academic editor: Umilaela Arifin @ Received 20 November 2024 Accepted 29 January 2025 Published 24 February 2025 Abstract A new species of Rhacophorus (Anura, Rhacophoridae), Rhacophorus medogensis sp. nov., is described from Medog, Xizang Tibetan Autonomous Region, China. The new taxon can be distinguished from all phylogenetically closely related taxa (R. rhodopus and R. bipunctatus complexes) by the following combination of features: 1) body size medium (adult males SVL 31.6—38.7 mm, n= 17; adult females SVL 50.1-55.7 mm, n = 2); 2) pineal ocellus obvious; 3) toe webbing formula: 11—1I]1—1.5011—-11V1-1V; 4) irregularly shaped large black spots, white pattern in black spots on flanks; 5) snout pointed with appendage on tip; 6) tongue pyriform, with a deep notch at posterior tip; 7) tibiotarsal articulation reaching eye. In addition, we also discuss the distribution of R. bipunctatus, which is limited to northern India and central-western Myanmar, rather than the traditionally presumed range across South and Southeast Asia. Key Words Biodiversity, cryptic species, Rhacophorus medogensis sp. nov., taxonomy Introduction Khasi Hills in northern India (Bordoloi et al. 2007). The species has been reported from a broad distribu- The genus Rhacophorus Kuhl & Van Hasselt, 1822, currently includes 46 species and is widely distributed across the tropical and subtropical regions of southern Asia, from India to China and Southeast Asia (Am- phibiaChina 2024; AmphibiaWeb 2024; Frost 2024). Rhacophorus bipunctatus, commonly known as the Himalaya Flying Frog, was first described from the * These authors contributed equally to this work. tion across South and Southeast Asia, including Ban- gladesh, Cambodia, China, Laos, Malaysia, Myanmar, Thailand, and Vietnam (Frost 2024). However, the taxonomic relationship between R. bipunctatus and R. rhodopus has been a topic of considerable debate over recent decades (Inger et al. 1999; Frost et al. 2006; Bordoloi et al. 2007; Chan et al. 2018). Copyright Weng, S. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 438 Recent phylogenetic analyses have revealed the com- plexity in the relationships between Rhacophorus bi- punctatus and R. rhodopus. Chan et al. (2018) suggested that they belong to distinct species complexes, indicating potential cryptic diversity. The phylogenetic analyses of Che et al. (2020) showed that the “R. rhodopus” popula- tion from Medog, Xizang, is distantly related to the type locality of R. rhodopus and is morphologically more sim- ilar to R. bipunctatus (Mathew & Sen, 2010), classifying itas R. bipunctatus. However, Li et al. (2022) and Tang et al. (2024) showed that the “R. bipunctatus” from Medog is distantly related to R. bipunctatus from northern India; four species (R. rhodopus, R. napoensis, R. giongica, and R. kio) in China belong to the R. rhodopus and R. bipunc- tatus complexes. However, these studies did not include molecular and morphological data from the type locality of R. bipunctatus, leaving uncertainties regarding the true distribution and diversity within this complex. To address these gaps, this study incorporated sequences from near the type locality of R. bipunctatus (Mawblang, Cherapun- ji, Southern Khasi Hills, northern India) to reconstruct the phylogenetic relationships within the R. rhodopus and R. bipunctatus complexes. Based on both morphological and molecular evidence, we also describe a new species of Rhacophorus from Medog, Xizang. Materials and methods sampling Fieldwork was conducted at Medog, Xizang Tibetan Au- tonomous Region, China. Eight specimens were collect- ed in May 2024. The specimens were collected by hand and subsequently euthanized with a low concentration of clove oil solution following standard euthanasia proto- cols for amphibians (Leary et al. 2020). Liver or muscle tissues were taken from the specimens and preserved in 95% ethanol. While the specimens were fixed in 75% eth- anol. Voucher specimens SWU 0008599, SWU 0008600, SWU 0008601, SWU 0008602, SWU 0008603, SWU 0008604, SWU 0008699, and SWU 0008701 were de- posited at Southwest University (SWU). Morphology and morphometrics We measured all the voucher specimens. All the measure- ments were made with slide calipers to the nearest 0.1 mm. Morphological terminology followed Fei (1999) and Fei et al. (2009) and is listed in Table 1; the webbing formula fol- lowed Myers and Duellman (1982). Morphological mea- surements of all specimens are listed in Table 2. Compara- tive morphological data of the new species and congeneric Species were obtained from published literatures (Liu and Hu 1960; Ohler and Delorme 2006; Bordoloi et al. 2007; Fei et al. 2009, 2012: Chan and Grismer 2010; Rowley et al. 2012; Matsui et al. 2013; Li et al. 2022; Tang et al. 2024; Che et al. 2020). Sex of the specimens was determined by the presence of nuptial pads and vocal sac openings. zse.pensoft.net Weng, S. et al.: One new species of the genus Rhacophorus from China Table 1. Morphological characters used for adult individuals. Abbreviation Morphology SVL Snout-vent length HL Head length HW Head width Si Snout length IND Internarial distance lOD Interorbital distance UEW Width of upper eyelid ED Diameter of eye TD Diameter of tympanum DNE Distance from nostril to eye FHL Length of forearm and hand TL Tibia length el Length of foot and tarsus FL Foot length DNA sequencing and molecular analyses To construct a phylogeny for Rhacophorus rhodopus and R. bipunctatus complexes, we extracted total DNA from liver or muscle tissue using the Animal Tissue DNA Iso- lation Kit provided by Thermo Fisher Scientific. In this study, we chose five specimens (SWU 0008599, SWU 0008600, SWU 0008601, SWU 0008603, and SWU 0008699) to sequence three consecutive mitochondrial gene segments: partial 12S rRNA, tRNA”, and partial 16S rRNA. The primers used for polymerase chain re- action (PCR) amplification are detailed in Table 3. The PCR amplification process was carried out in a 50-uL reaction volume, following the reaction cycling settings below: an initial denaturing step at 95 °C for 4 min; 35 cycles of denaturing at 94 °C for 60 s, annealing at 51 °C for 60 s, and extending at 72 °C for 60 s; and a fi- nal extending step of 72 °C for 10 min. Sequencing was conducted using the corresponding PCR primers and an internal primer (Rhint: 5’-GACAGTGTAACCCTCGT- GAT-3’) (Yu et al. 2019), and all sequences have been uploaded to GenBank (Table 4). Based on Tang et al. (2024), we selected species from the genera Rhacoph- orus, Buergeria, Nyctixalus, Chiromantis, Theloderma, Kurixalus, Zhangixalus, and Leptomantis as outgroups to reconstruct the phylogenetic relationships of the R. rhodopus and R. bipunctatus complexes; all Gen- Bank accession numbers are listed in Table 4. Sequences were aligned using the MUSCLE option in MEGA v. 7, and due to the absence of 12S rRNA and tRNA” sequences in some species, uncorrected pair- wise distances (p-distances) were calculated only for 16S rRNA sequences (500 bp) between species (Kumar et al. 2016). Prior to phylogenetic reconstruction, the best sub- stitution model was selected using the Akaike Informa- tion Criterion (AIC) in ;jMODELTEST v. 2.1.10 (Darriba et al. 2012). Bayesian inference (BI) was performed using MrBayes v.3.2.6 (Ronquist et al. 2012), with two runs performed simultaneously, each consisting of four Markov chains starting from a random tree. The chain was run for 6,000,000 generations, with sampling every Zoosyst. Evol. 101 (2) 2025, 437-447 Table 2. Measurements (mm) of adult specimens in the type series of R. medogensis sp. nov. 439 NO SWU SWU SWU SWU SWU SWU SWU SWU 0008699 0008599 0008600 0008601 0008602 0008603 0008604 0008701 SEX Adult male Adult male Adult male Adult male Adult male Adult male Adult male Subadult SVL 3335 36.2 34.1 35.4 33:5 34.0 34.6 27.4 HL DAA? 13.4 11.8 13:1 1230 12:3 12-7 10.8 HW qed ck 12.2 5 12.4 Le? 11.6 12.0 10.4 SL 5.0 5.4 aes. 5.4 533 52 53 4.7 IND 3.2 iy 3.4. 35 ioe) 3.4 333 27 lOD 4.6 4.5 4.6 4.6 4.8 4.5 4.7 3.6 UEW 2.5 vay 2.9 2.8 oof 25 Pay. 22 ED 4.2 4.6 4.3 4.4 4.4 4.2 4.4 3.6 TD ae oa anal eee 2.2 1.9 alk 1.9 DNE 26 Ph 2.4 2.4 2.4 2.6 225 222 FHL 16.3 16.5 16.5 ies | 16.8 16.9 18.3 13.9 aH 16.6 17.9 16.3 LF 16.2 Ly6 17.6 14.5 TFL 22.4 23.0 21.9 23.5 22.1 23.3 238 18.4 FL 14.0 14.8 13.2 14.9 14.5 15:2 1521 11.9 Table 3. Primer pairs for PCR amplification used in this study. Gene Primer Source 12S rRNA, tRNA, and partial 16S rRNA 1,000 generations. When the average standard deviation of the split frequency was less than 0.01, the first 25% of the sampled trees were discarded as burn-in, and the remaining trees were used to create a consensus tree and estimate the Bayesian posterior probabilities. Maximum likelihood (ML) analyses were performed using RAXML v7.0.3 (Stamatakis 2014) under the GTRGAMMA model with 1000 bootstrap replicates. Results Phylogenetic analyses The obtained sequence alignment was 1936 bp long. The phylogenetic trees inferred from BI and ML methods are generally consistent (Fig. 1). The phylogenetic results of this study show differences in the placement of certain species compared to Tang et al. (2024) but consistently indicate that the Rhacophorus bipunctatus complex is nested within the R. rhodopus complex. The R. rhodopus and R. bipunctatus complexes comprise 10 known spe- cies and | undescribed lineage (R. medogensis sp. nov., R. rhodopus, R. napoensis, R. giongica, R. borneensis, R. norhayatiae, R. reinwardtii, R. bipunctatus, R. kio, R. helenae, and R. sp.). The sequence of R. bipunctatus from the near-type locality clusters together with those from northeastern India and central-western Myanmar, representing the “true” R. bipunctatus. The new species R. medogensis sp. nov. clusters within the R. rhodopus and R. bipunctatus complexes and represents the sister group to R. borneensis, R. norhayatiae, and R. reinwardtii with strong support (BPP = 0.98, BS = 77). The undescribed L1091 (5'-AAAAAGCTTCAAACTGGGATTAGATACCCCACTAT-3') 16H1 (5'-CTCCGGTCTGAACTCAGATCACGTAGG-3') Kocher et al. 1989 Hedges 1994 lineage (R. sp.) represents the sister group to R. rhodopus with strong support (BPP = 0.99, BS = 95). P-distances of 16S rRNA between the sequences of R. medogensis sp. nov. and the other species of R. rhodopus and R. bipunc- tatus complexes varied from 7.5% to 11.6%, while the p-distances of 16S rRNA between R. sp. and R. rhodopus is 7% (Table 5). Taxonomic account Rhacophorus medogensis sp. nov. https://zoobank.org/4COCEFC4-09D5-4229-B2A 9-C0655D43E452 Figs 3, 4, Table 2 Chresonymy. Rhacophorus rhodopus—Hu 1987; Fei et al. 2004; Fei et al. 2009; Li et al. 2011; Fei et al. 2012; Li et al. 2022. Rhacophorus bipunctatus—Che et al. 2020; Tang et al. 2024 Holotype. SWU 0008699, adult male, collected in May 2024 by Xiaolong Liu, Renda Ai, and Xianqi Li from Didong Village, Medog, Xizang Tibetan Autono- mous Region, China (29.2205°N, 95.1293°E, elevation 771 m; Fig. 2A). Paratypes. Six adult males (SWU 0008599, SWU 0008600, SWU 0008601, SWU 0008602, SWU 0008603, and SWU 0008604) were collected at Buqun (Xigong) Lake, Medog, Xizang Tibetan Autonomous Re- gion, China (29.25241°N, 95.225759°E, elevation 1361 m). One subadult (SWU 0008701) was collected at the same locality as the holotype in May 2024 by Xiaolong Liu, Renda Aj, and Xianqi Li. zse.pensoft.net 440 Weng, S. et al.: One new species of the genus Rhacophorus from China Table 4. Species used in phylogenetic analyses of this study. No. CON DODO WN FE Etymology. The specific epithet “medogensis” is named after the type locality, Medog, Xizang, China. We suggest “Xizang flying frog” as its English com- mon name and “Mo Tudo Shu Wa” (2ehii py) as its Species R. medogensis sp. R. medogensis sp. R. medogensis sp. R. medogensis sp. R. medogensis sp. R. medogensis sp. R. medogensis sp. R. medogensis sp. R. medogensis sp. R. bipunctatus R. bipunctatus R. bipunctatus R. bipunctatus R. bipunctatus R. napoensis R. napoensis R. napoensis R. napoensis R. napoensis R. napoensis R. giongica R. giongica R. giongica R. rhodopus R. rhodopus R. rhodopus R. rhodopus R. rhodopus R. rhodopus R. rhodopus Sp Sp Sp Sp Sp Sp Sp Sp Sp R. borneensis R. borneensis R. helenae R. kio R. lateralis R. norhayatiae R. norhayatiae R. nigropalmatus R. reinwardtii Buergeria buergeri VADAAAAAADA nov. nov. nov. nov. nov. nov. nov. nov. nov. Chiromantis rufescens Kurixalus idiootocus Leptomantis gauni Nyctixalus pictus Theloderma albopunctatum Zhangixalus smaragdinus Chinese common name. zse.pensoft.net Voucher KIZO16380 YPX40427 L06245 L062456 SWU 0008599 SWU 0008600 SWU 0008601 SWU 0008603 SWU 0008699 CAS229913 CAS235303 PUCZM/1X/SL360 PUCZM/IX/SL612 / GXNU YU000171 GXNU YU000173 VNMN:4118 VNMN:4120 VNMN:4121 AMNH-A 161418 SN 030035 VNMN:4117 FMNH253114 clone 5 SCUM 060692L KIZ060821229 2004.04.09 2006.2519 K3046 K3085_1 clone 4 KIZ060821248 KIZ060821175 clone 2 KIZ587 KIZ589 USNM:Herp:587063 0937Y4 KUHE:53375 NMBE 1056517 BORN 22411 UNS 00451 KUHE 55165 f Rao081205 NNRn Ra0081204 ENS 16179 (UTA) TTU-R-11759 CAS 207601 CAS 211366 FMNH 273928 FMNH 231094 ROM 30246 HM05292 Locality Accession No Medog, Xizang, China MW111517 Medog, Xizang, China MW111518 Medog, Xizang, China JX219441 Medog, Xizang, China JX219442 Medog, Xizang, China PQ963460 Medog, Xizang, China PQ963459 Medog, Xizang, China PQ963458 Medog, Xizang, China PQ963457 Medog, Xizang, China PQ963456 Nagmung Township, Putao District, Kachin State, Myanmar JX219445 Mindat Township, Mindat District, Chin State, Myanmar JX219444 Mizoram, Inida MH087073 Mizoram, Inida MH087076 Mawblang, Cherapunji, Southern Khasi Hills, India* OL988889 Napo, Guangxi, China ON217796 Napo, Guangxi, China ON217798 Yen Tu, Bac Giang, Vietnam LC010605 Pu Huong, Nghe An, Vietnam LCO10609 Thanh Hoa, Vietnam LC010608 Huon Son Reserve, Ha Tinh, Vietnam AY843750 Hainan, China EU215529 kK’ Bang, Gia Lai, Vietnam LC010604 Ankhe Dist, Gia Lai, Vietnam GQ204716 Mengyang, Yunnan, China EF646366 Mengyang, Yunnan, China EU2Z15531 Lvchun, Yunnan, China EF564574 Long Nai Khao, Phongsali, Laos KR828049 Ban Vang Thong, Louangphrabang, Laos KR828069 Doi Chiang Dao, Chiang Mai, Thailand KR828066 Mae Lao-Mae Sae Wildlife Sanctuary, Chiang Mai, Thailand KR828067 Jingdong, Yunnan, China EF646365 Jingdong, Yunnan, China EF564575 Yongde, Yunnan, China EF564573 Yongde, Yunnan, China EF646363 Longling, Yunnan, China EF564577 Longling, Yunnan, China EF564578 Kandawgyi National Gardens, Mandalay, Myanmar MG935991 Kui Buri NP, Prachuap Khiri Khan, Thailand KR828058 Genting, Pahang, Malaysia LCO010569 Batang Ai NP, Sarawak, Malaysia JN377366 Sabah, Maliau Basin, Malaysia AB781694 Binh Thuan, Vietnam JQ288090 Xuan Lien, Than Hoa, Vietnam AB781695 Mudigere, India AB530548 Malaysia JX219443 Johor, Endau Rompin, Malaysia AB728191 Malaysia JX219437 Java, Patuha, Indonesia KY886328 Japan AF458122 Bioko Norte Province, Equatorial Guinea AF458126 Taipei, Taiwan, China AF458129 Bintulu, Sarawak, Malaysia JX219456 Lahad Datu, Sabah, Malaysia AF458135 Vietnam AF458148 Xima, Yingjiang, Yunnan, China MN613221 Diagnosis. The genus Rhacophorus, also known as fly- ing frogs, is characterized by the following features: 1) body size relatively moderate or large (SVL 30-100 mm, above 40 mm in most species); 2) presence of intercalary cartilage between terminal and penultimate phalanges of digits; 3) 441 Zoosyst. Evol. 101 (2) 2025, 437-447 EF564578 EF564577 EF564573 EF646365 EF646363 EF564575 MG935991 KR828058 Lco10569 KR828069 0.99/95 KR828049 EF646366 EF564574 400 KR828066 ones £U215531 KR828067 MH087076 MH087073 +JX219444 1/100 JX219445 OLg8s889 MW111518 +JX219442 MW111517 0.98/81 +JX219441 PQ963457 PQ963459 4100 Pag63456 Pa963460 1/100 PQ963458 JX219443 AB728191 aimed 0.99/91 KY886328 ‘ Ly 0.99/99 JN377366 AB781694 ON217798 0.98/7 R. rhodopus and R. bipunctatus complexes ON217796 LC010605 41/100 LC010609 LC010608 AY843750 1198 JQ288090 0.95/99 AB781695 Rh LC010604 f 4100 1/100 GQ204716 EU215529 100 JX219437 AB530548 MN613221 AF458126 0.96/-) JX219456 AF458148 R._napoensis ATT Ar] 1/96 AF458135 AF458129 - AF458122 Figure 1. Bayesian phylogenetic tree of R. rhodopus and R. bipunctatus complexes and related species inferred from 12S rRNA, tRNA- Val and 16S rRNA genes. The numbers above and below the branches are Bayesian posterior probabilities (BPP) and maximum likeli- hood bootstrap values (BS), “-” denotes a BPP < 0.95 and BS < 70. The scale bar represents 0.04 nucleotide substitutions per site. The “black star” means this sequence from from near type locality of R. bipunctatus, R. medogensis sp. nov., R. rhodopus and R. sp photoed by Xiaolong Liu, R. bipunctatus from Bordoloi et al. (2007), R. giongica photoed by Chenxi Liao, R. napoensis by Junkai Huang. Table 5. Mean uncorrected pairwise distances (%) between clades of R. rhodopus and R. bipunctatus complexes and related species based on 16S rRNA sequences. No Species 1 2 3 4 5 6 7 8 9 10 11 1 R. medogensis sp. nov. 2 FR. bipunctatus LO 3. RFR. napoensis 8.4 8.8 4 R.sp 9.7 9.7 10.4 5 R. giongica 9.1 13.1 LOF 13.9 6 R. reinwardtii 8.6 O2 Ato G3 12.2 7 R. rhodopus 116 10.1 LE 7G) 123 10.3 8 R. norhayatiae 8.6 9.5 8.1 9.3 134 5.8 Aided 9 PR. helenae 10.3 10.1 9.9 10.4 12.4 8.9 13.2 10.1 10 R. borneensis HS 8.7 6.7 8.6 10.1 6.4 11.7 oa 6.7 11 R.kio 10.4 11.1 9.5 L..3 13.5 La 14.0 9.3 6.8 7.8 terminal phalanges of fingers and toes Y-shaped; 4) tips of the digits expanded into large disks bearing circummarginal grooves; 5) webbed fingers; 6) skin not co-ossified to the skull; 7) upper eyelid projections absent, tarsal projections present in most species; 8) dermal folds along the forearm or tarsus present; 9) pupil horizontal; 10) iris without “X”- shaped marking; 11) white foam nests or jelly-encapsulat- ed eggs produced by breeding pairs; and (12) distributed mainly in Indochina (Jiang et al. 2019). Rhacophorus me- dogensis sp. nov. is placed in the genus Rhacophorus due to the combination of the following features: 1) body size relatively moderate (adult males SVL 31.6—38.7 mm, n = 17; adult females SVL 50.1-55.7 mm, n = 2); 2) presence of intercalary cartilage between terminal and penultimate phalanges of digits; 3) terminal phalanges of fingers and toes Y-shaped; 4) tips of the digits expanded into large disks bearing circummarginal grooves; 5) webbed fingers; 6) tar- sal projections present; 7) pupil horizontal; 8) tris without “X”-shaped marking. Rhacophorus medogensis sp. nov. can be distinguished from other species in the R. rhodopus and R. bipunctatus complexes by the following combination of features: 1) medium adult males body size (adult males SVL 31.6—38.7 mm); 2) dorsal surface reddish brown, light green, light brown, or grayish green in life; 3) pineal ocellus obvi- ous; 4) toe webbing formula: 11—1TI1—1.5111—11V1—1V; 5) irregularly shaped large black spots, white pattern in black spots on flanks; 6) snout pointed with an appendage on the tip; 7) tongue pyriform, with a deep notch at the posterior tip; 8) throat rough; 9) palm rough with small tubercles; 10) tibiotarsal articulation reaching the eye. zse.pensoft.net 442 Figure 2. (A) Large habitat at the type locality of R. medogensis sp. nov., Didong Village, Medog County, Xizang Tibetan Autonomous Region, China (29.2205°N, 95.1293°E, elevation 771 m); (B) macrohabitat of R. medogensis sp. nov. (SWU 0008601), Buqun (Xigong) Lake, Medog County, Xizang Tibetan Autonomous Re- gion, China (29.25241°N, 95.225759°E, elevation 1361 m). Description of holotype. Adult male, medium body size (SVL 33.5 mm); head length (HL 12.1 mm) longer than head width (HW 11.1 mm); snout pointed with an ap- pendage on tip, sloping in profile, and protruding beyond the margin of lower jaw in ventral view; snout length (SL 5.0 mm) is longer than the diameter of the eye (ED 4.2 mm); the canthus rostralis is distinct and curved; loreal region oblique, concave; nostrils oval, lateral, slightly protuberant, and slightly closer to the tip of snout than the eye; the internarial space (IND 3.2 mm) ts slightly small- er than the interorbital distance OD 4.6 mm) and larger than the width of the upper eyelid (UEW 2.5 mm); the pu- pil is horizontal; pineal ocellus obvious; tympanum dis- tinct (TD 2.2 mm), rounded, and nearly about half of eye diameter (ED 4.2 mm); the supratympanic fold is narrow and flat; tongue pyriform, with a deep notch at the posteri- or tip; choanae oval; vomerine teeth present in two series; with an internal single subgular vocal sac; a vocal sac opening on the floor of the mouth at each corner (Fig. 3). Forelimbs strong, length of forearm and hand (FHL 16.3 mm); relative length of fingers I < II < IV