A New Species of the Genus Gracixalus (Anura, Rhacophoridae) from Central Western Yunnan, China †
by
, ,
Shuo Liu
1,2
,
Zhongfu Yang
3,
Chunhua Wang
3,
Nengping Li
3,
Mian Hou
4,
Zengyang Luo
5,* and
Dingqi Rao
6,* 1
Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650223, China
2
Yunnan Key Laboratory of Biodiversity Information, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650201, China
3
Jingdong Management and Protection Bureau of Yunnan Wuliangshan and Ailaoshan National Nature Reserves, Jingdong 676200, China
4
College of Continuing (Online) Education, Sichuan Normal University, Chengdu 610068, China
5
Nanjian Management and Protection Bureau of Yunnan Wuliangshan National Nature Reserve, Nanjian 675700, China
6
Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650201, China
*
Authors to whom correspondence should be addressed.
†
Zoobank: urn:lsid:zoobank.org:pub:5F35C014-165F-4AE6-A3AA-32A4F48A95E8; urn:lsid:zoobank.org:act:86AA38CB-3CE6-4F26-AE2C-11A5DFC70F0C.
Taxonomy 2025, 5(4), 66; https://doi.org/10.3390/taxonomy5040066
Submission received: 26 August 2025
/
Revised: 4 November 2025
/
Accepted: 4 November 2025
/
Published: 26 November 2025
(This article belongs to the Special Issue Taxonomy in the 21st Century: Celebrating a New Chapter—First Impact Factor Received)
Abstract
A new species of the genus Gracixalus from Yunnan Wuliangshan National Nature Reserve, China, is described based on morphological and molecular evidence. Morphologically, the new species can be distinguished from other species of the genus by a combination of the following characters: body size relatively large, dorsal surface brownish yellow, dorsal skin rough with dense, small flatten tubercles, tibiotarsal projection absent, heels distinctly overlapping, tibiotarsal articulation reaching anterior corner of eye, and nuptial pad absent in adult male. The genetic distance (uncorrected p-distance) between the new species and other species of the genus ranged from 2.3% to 14.3% in the mitochondrial 16S rRNA gene. This study brings the total number of recognized species within the genus Gracixalus to 24 with 13 of which occur in China and four in Yunnan Province.
1. Introduction
The genus Gracixalus Delorme, Dubois, Grosjean & Ohler, 2005 is a group of small- to medium-sized arboreal frogs [1,2]. Their reproduction usually relies on tree holes or open or half open bamboo tubes with accumulated water [1,3,4]. The genus has a wide distribution from India, Myanmar, Thailand, Laos, and Vietnam, to southern China [5]. It currently contains 23 species, 12 of which are distributed in China and three have been recorded from Yunnan Province [4,5,6,7,8,9].
Initially, Gracixalus was established as a subgenus with the type species Philautus gracilipes Bourret, 1937 under the genus Aquixalus Delorme, Dubois, Grosjean & Ohler, 2005 by Delorme et al. [10]. Subsequently, Li et al. [11] elevated Gracixalus to a full genus. Later on, Dubois et al. [12] split Gracixalus into two genera, namely Gracixalus sensu stricto and Orixalus Dubois, Ohler & Pyron, 2021. However, this taxonomic change was rejected by Mahony et al. [13] and Luo et al. [4]. Due to their relatively small body size, conserved morphology, and difficulty in specimen collection, the species diversity of Gracixalus has long been severely underestimated. In recent years, new species have been continuously described within this genus [2,4,8,9,14].
Yunnan Wuliangshan National Nature Reserve located in the central west part of Yunnan Province, China, it stretches for approximately 418 km2 and marks the junction between the Hengduan Mountains and the Yunnan Plateau [15]. This nature reserve encompasses typical ecological systems of China’s subtropical plateau region, which includes evergreen broadleaf forests, monsoon forests, wetlands, and other ecosystems, and serves as a transitional zone in terms of flora and fauna [10]. Yunnan Wuliangshan National Nature Reserve is renowned for its rich biodiversity and is home to numerous rare and endangered wildlife species [15,16].
During our recent fieldwork in Yunnan Wuliangshan National Nature Reserve, we collected two specimens of the genus Gracixalus. However, these two specimens are distinct from all known species of the genus in both phylogenetic analyses and morphological comparison. Herein, we describe these two specimens as a new species of Gracixalus.
2. Materials and Methods
Field surveys were conducted under the permission of Jingdong Management and Protection Bureau of Yunnan Wuliangshan and Ailaoshan National Nature Reserves and specimen collection was performed in accordance with the Wild Animals Protection Law of the People’s Republic of China. The animal study was approved by the Ethics Committee of Kunming Institute of Zoology, Chinese Academy of Sciences (protocol code IACUC-OE-2024-10-001). Specimens were collected by hand at night, after being euthanized and fixed, and then were stored in 75% ethanol and deposited at Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences (KIZ).
Genomic DNA was extracted from liver tissue samples using a commercial tissue DNA isolation kit (Tiangen Biotech Co., Ltd., Beijing, China). A partial fragment of the mitochondrial 16S ribosomal RNA gene (16S) was amplified and sequenced using the primer pair L2188 [17] and 16H1 [18]. Amplification and sequencing were completed by Sangon Biotech Co., Ltd. (Shanghai, China). PCR amplification reaction conditions followed Liu et al. [8]. All new sequences have been deposited in GenBank and homologous sequences of other species of Gracixalus were retrieved from GenBank (Table 1). Kurixalus idiootocus (Kuramoto & Wang, 1987) was used as outgroup according to Luo et al. [4].
Sequences were aligned using ClustalW [19] integrated in MEGA 11.0.13 [20] with default parameters. Uncorrected pairwise distances were calculated in MEGA 11.0.13 [20]. The best substitution model GTR+F+I+G4 was selected using the Akaike Information Criterion in ModelFinder [21]. Bayesian inference and maximum likelihood were used to construct the phylogenetic tree. Bayesian inference was performed in MrBayes 3.2.7 [22] with the Markov chains were run for 5,000,000 generations and sampled every 100 generations. Maximum likelihood analysis was performed in IQ-TREE 1.6.12 [23]. with 1000 ultrafast bootstrap replicates.
Morphological measurements were taken using electronic digital vernier calipers to the nearest 0.1 mm followed Matsui et al. [24] and Le et al. [25]: SVL, snout-vent length; HL, head length, from the back of the mandible to the tip of the snout; HW, maximum head width, across the angle of the jaws; SNL, snout length, from the anterior corner of the eye to the tip of the snout; IN, internarial distance; IOD, interorbital distance; ED, eye diameter; UEW, maximum width of the upper eyelid; NEL, distance from the anterior corner of the eye to the nostril; TD, tympanum diameter; TEL, distance from the anterior margin of the tympanum to the posterior corner of the eye; DAE, distance between the anterior corners of the eyes; DPE, distance between the posterior corners of the eyes; MN, distance from the back of the mandible to the nostril; FLL, forelimb length, from the axilla to the tip of the third finger; LAL, lower arm and hand length, from the elbow to the tip of the third finger; 3FD, width of the disk of the third finger; HLL, hindlimb length, from the groin to the tip of the fourth toe; TL, tibia length, from the knee to the tarsus; TW, maximum tibia width; TFL, from the tarsus to the tip of the fourth toe; FL, foot length, from the inner metatarsal tubercle to the tip of the fourth toe; 3TD, width of the disk of the third toe; and IMT, inner metatarsal tubercle length.
3. Results
3.1. Phylogenetic Relationship
Bayesian inference and maximum likelihood analysis resulted in a consistent topology. The samples from Yunnan Wuliangshan National Nature Reserve formed a distinct lineage sister to Gracixalus sapaensis Matsui, Ohler, Eto & Nguyen, 2017 with strong support by Bayesian inference and weak support by maximum likelihood analysis (Figure 1). The genetic distance (uncorrected pairwise distance) between the novel linage and G. sapaensis was 2.5% and the minimum genetic distance (uncorrected pairwise distance) between the novel linage and its congeners was 2.3% (vs. G. tianlinensis Chen, Bei, Liao, Zhou & Mo, 2018), both larger than that (1.8%) between G. liusanjieae Wu, Pan, Chen, Ye, Yu & Zou, 2025 and G. weii Liu, Peng, Wang, Feng, Shen, Li, Chen, Su & Tang, 2025 and that (2.1%) between G. guangdongensis Wang, Zeng, Liu & Wang, 2018 and G. yunnanensis Yu, Li, Wang, Rao, Wu & Yang, 2019, and approximately equal to that (2.4%) between G. tianlinensis and G. ziegleri Le, Do, Tran, Nguyen, Orlov, Ninh & Nguyen, 2021 (Table 2).
3.2. Taxonomic Account
Gracixalus wuliangshan sp. nov.
urn:lsid:zoobank.org:act:86AA38CB-3CE6-4F26-AE2C-11A5DFC70F0C
Holotype. KIZ2025067, adult female, collected on 20 July 2025 by Shuo Liu from Caijia Village, Wenlong Town, Jingdong County, Yunnan Province, China (24°34′40″ N, 100°38′18″ E, 2250 m elevation).
Paratype. KIZ 036308, adult male, collected on 18 September 2016 by locals from the same locality as the holotype.
3.3. Etymology
The specific epithet is named for Yunnan Wuliangshan National Nature Reserve, in which the species was collected. We suggested “Wuliangshan small tree frog” for the common English name and “无量山纤树蛙 (wú liàng shān xiān shù wā)” for the common Chinese name.
3.4. Diagnosis
The new species can be distinguished from its congeners by a combination of the following morphological characters: body size relatively large, SVL 39.1 mm in male and 44.1 mm in female; dorsal surface brownish yellow with a distinct X-shaped pattern composed of wide dark brown stripes on dorsum; iris light yellowish gray; axilla and groin with light yellow spots; dorsal skin rough with dense small flatten tubercles; snout round; tibiotarsal projection absent; finger webbing rudimentary; heels distinctly overlapping when legs at right angle to body; tibiotarsal articulation reaching anterior corner of eye; internal vocal sac in male; nuptial pad absent in male.
3.5. Description of Holotype
Body robust, SVL 44.1 mm; head slightly shorter than wide (HL/HW 0.93); snout round, projecting beyond margin of lower jaw in ventral view, length greater than eye diameter (SNL/ED 1.34); nostril small, almost round, closer to tip of snout than to eye; internarial distance nearly equal to interorbital distance (IN/IOD 0.98); canthus rostralis distinct, blunt; loreal region oblique, slightly concave; eye moderate (ED/HL 0.34), pupil oval, horizontal; upper eyelid width smaller than interorbital distance (UEW/IOD 0.82); tympanum distinct, slightly smaller than half of eye diameter (TD/ED 0.46), separated from eye by approximately half of tympanum diameter (TEL/TD 0.52); supratympanic fold distinct, extending from posterior corner of eye to above insertion of arm; vomerine teeth absent; choanae round; tongue oval, notched posteriorly.
Forelimbs relatively long (FLL/SVL 0.71); relative finger length III > IV > II > I; webbing between fingers rudimentary; tips of all fingers expanded into disks with circum-marginal grooves; subarticular tubercles prominent, round, formula 1, 1, 2, 2, distal one greater than proximal one on fingers III and IV; supernumerary tubercles absent; two metacarpal tubercles, oval.
Hindlimbs moderately long (HLL/SVL 1.57); relative toe length IV > V > III > II > I; tip of all toes expanded into disks with circum-marginal grooves; webbing between toes moderately developed, formula I13/2–2II11/3–21/3III11/2–22/3IV21/2–11/2V; subarticular tubercles prominent, round, formula 1, 1, 2, 3, 2; supernumerary tubercles absent; inner metatarsal tubercle oval, outer metatarsal tubercle absent; heels overlapping when legs at right angle to body; tibiotarsal articulation reaching anterior corner of eye when hindlimb adpressed forward to body.
Dorsal surface rough, covered with dense small tubercles; lateral surface of head and flank relatively smooth, scattered with some indistinct small tubercles; ventral surface of head smooth; ventral surface of body and limbs granular.
3.6. Coloration of Holotype in Life
Dorsal surface brownish yellow; a distinct X-shaped pattern composed of wide dark brown stripes on dorsum, upper half short, interior filled with dark brown, lower half long, extending almost to crotch; dorsal surface of limbs with distinct dark brown bands; lateral surface of head light brown, a wide dark brown stripe from anterior corner of eye to snout tip under canthus rostralis, tympanum region dark brown, iris light yellowish gray; flank brownish yellow, a few small light yellow spots on axilla, some large light yellow spots on groin; throat beige; belly grayish white; ventral surface of forelimbs light flesh color; ventral surface of hindlimbs light orange.
3.7. Coloration of Holotype in Preservative
Dorsal surface faded to brownish gray, X-shaped pattern on dorsum and bands on dorsal limbs still visible, ventral surface turned to yellowish white.
3.8. Variations
The male paratype has a smaller body size, an internal subgular vocal sac, and dark blotches on ventral surface. Apart from these, there is no significant difference between the paratype and the holotype (Table 3).
3.9. Distribution
The new species is currently known only from Jingdong County, Yunnan Province, China (Figure 4).
3.10. Natural History Notes
The advertisement calls of the new species were all heard from tall trees, and during our active search for this species, we did not find any individuals on the low plants, indicating that they are likely to inhabit the canopy layer. However, the type specimens of the new species were accidentally discovered on the leaves of small herbaceous plants while we were collecting specimens of other species, indicating that this species occasionally moves to areas near the ground. No eggs and tadpoles of the new species were found at the type locality.
3.11. Comparisons
Gracixalus wuliangshan sp. nov. differs from G. ananjevae (Matsui & Orlov, 2004) by snout length greater than eye diameter (vs. snout length equal to eye diameter) and nuptial pad absent (vs. present on finger I in males); differs from G. carinensis (Boulenger, 1893) by larger body size (SVL 39.1 mm in male and 44.1 mm in female vs. 30.2–38.1 mm in males and 40.3–41.8 mm in females) and dorsal surface rough with tubercles (vs. nearly smooth); differs from G. gracilipes (Bourret, 1937) by larger body size (SVL 39.1 mm in male and 44.1 mm in female vs. 20–24 mm in males and ca. 30 mm in females), snout round (vs. triangularly pointed), and dorsal surface brownish yellow (vs. greenish); differs from G. guangdongensis by larger body size in females (SVL 44.1 mm vs. 34.9–35.4 mm) and nuptial pad absent (vs. present on finger I in males); differs from G. huaping Luo, Zhang, Pan & Yu, 2025 by larger body size (SVL 39.1 mm in male and 44.1 mm in female vs. SVL 26.6–28.8 mm in males and 29.8–32.8 mm in females) and nuptial pad absent (vs. present on finger I in males); differs from G. jinggangensis Zeng, Zhao, Chen, Chen, Zhang & Wang, 2017 by larger body size in females (SVL 44.1 mm vs. 31.6 mm), heels overlapping (vs. just meeting), and nuptial pad absent (vs. present on finger I and II in males); differs from G. jinxiuensis (Hu, 1978) by larger body size in females (SVL 44.1 mm vs. ca. 30 mm), heels overlapping (vs. just meeting), and tibiotarsal articulation reaching anterior corner of eye (vs. reaching posterior corner of eye); differs from G. liusanjieae by larger body size in females (SVL 44.1 mm vs. 38.1–39.7 mm), heels distinctly overlapping (vs. slightly overlapping), and tibiotarsal articulation reaching anterior corner of eye (vs. reaching center of eye); differs from G. lumarius Rowley, Le, Dau, Hoang & Cao, 2014 by larger size in females (SVL 44.1 mm vs. 36.3 mm), dorsal surface covered with small flatten tubercles (vs. dorsum with distinctive white conical tubercles in adult males), supratympanic fold distinct (vs. indistinct), and nuptial pad absent (vs. present on finger I in males); differs from G. medogensis (Ye & Hu, 1984) by larger body size in males (SVL 39.1 mm vs. 26.5 mm), dorsal surface brownish yellow (vs. grass green), dorsal surface rough with tubercles (vs. smooth), and finger webbing rudimentary (vs. absent); differs from G. nonggangensis Mo, Zhang, Luo, Zhou & Chen, 2013 by dorsal surface brownish yellow (vs. yellowish olive), and iris light yellowish gray (vs. olive); differs from G. patkaiensis Boruah, Deepak, Patel, Jithin, Yomcha & Das, 2023 by larger body size in males (SVL 31.9 mm vs. 23.6–26.5 mm), dorsal surface brownish yellow (vs. green), vocal sac internal (vs. external), and nuptial pad absent (vs. present on finger I in males); differs from G. quangi Rowley, Dau, Nguyen, Cao & Nguyen, 2011 by tibiotarsal projection absent (vs. present), dorsal surface brownish yellow (vs. olive-green), vocal sac internal (vs. external), snout round (vs. triangularly pointed), and nuptial pad absent (vs. present on finger I in males); differs from G. quyeti (Nguyen, Hendrix, Böhme, Vu & Ziegler, 2008) by dorsal surface brownish yellow (vs. brownish green to moss green), head wider than long (vs. head longer than wide), tibiotarsal articulation reaching anterior corner eye (vs. reaching snout), and supratympanic fold distinct (vs. indistinct); differs from G. seesom Matsui, Khonsue, Panha & Eto, 2015 by larger body size (SVL 39.1 mm in male and 44.1 mm in female vs. 21.6–23.0 mm in males and 23.2–25.4 mm in females), snout round (vs. triangularly pointed), and dorsal surface rough with tubercles (vs. smooth); differs from G. supercornutus (Orlov, Ho & Nguyen, 2004) by dorsal surface brownish yellow (vs. green), snout round (vs. triangularly pointed), dorsal surface with small flatten tubercles (vs. considerable bigger horn-like projections in supraorbital area, around cloaca, and on dorsum, forelimbs and hindlimbs), and tibiotarsal projection absent (vs. present); differs from G. trieng Rowley, Le, Hoang, Cao & Dau, 2020 by smaller body size in males (SVL 31.9 mm vs. 37.2–41.4 mm) and nuptial pad absent (vs. present on finger I in males); differs from G. truongi Tran, Pham, Le, Nguyen, Ziegler & Pham, 2023 by larger body size in females (SVL 44.1 mm vs. 37.6–39.3 mm), dorsal surface brownish yellow (vs. moss green), and nuptial pad absent (vs. present on finger I in males); differs from G. weii by larger body size in females (SVL 44.1 mm vs. 35.2–36.0 mm), articulation reaching anterior corner eye (vs. reaching center of eye), and nuptial pad absent (vs. present on finger I in males); differs from G. yunnanensis by dorsal surface with small flatten tubercles (vs. distinctive conical tubercles), vocal sac internal (vs. external), and nuptial pad absent (vs. present on finger I in males).
Gracixalus wuliangshan sp. nov. is phylogenetically closely related to G. sapaensis, G. tianlinensis, and G. ziegleri. However, Gracixalus wuliangshan sp. nov. can be distinguished from G. sapaensis by larger body size in females (SVL 44.1 mm vs. 27.2–39.5 mm), relatively smaller eye (ED/SVL 0.13 in male and 0.11 in female vs. 0.14–0.18 in males and 0.13–0.16 in females), and relatively longer foot (FL/SVL 0.51 in mele and 0.52 in female vs. 0.42–0.49 in males and 0.46–0.50 in females); can be distinguished from G. tianlinensis by larger body size in females (SVL 44.1 mm vs. 35.6–38.7 mm), vocal sac internal (vs. external), finger webbing rudimentary (vs. absent), and nuptial pad absent (vs. present on finger I and II in males); can be distinguished from G. ziegleri by larger body size in males (SVL 31.9 mm vs. 28.1–30.5 mm), tibiotarsal articulation reaching anterior corner eye (vs. reaching snout tip), and nuptial pad absent (vs. present on finger I in males).
4. Discussion
Rowley et al. [26] constructed a preliminary phylogenetic tree of the genus Gracixalus using 16S rRNA sequences and found that there are two clades within the genus, and they suggested the separation of the genus into two distinct genera in their brief discussion. However, this proposal was later refuted and not widely accepted [27]. In our phylogenetic analyses, the genus was divided into two distinct clades, although not highly supported, with the first clade containing G. gracilipes, G. patkaiensis, G. quangi, G. quyeti, G. seesom, and G. supercornutus, and the second clade containing G. ananjevae, G. guangdongensis, G. huaping, G. jinggangensis, G. jinxiuensis, G. liusanjieae, G. lumarius, G. medogensis, G. nonggangensis, G. sapaensis, G. tianlinensis, G. trieng, G. truongi, G. weii, G. yunnanensis, G. ziegleri, and Gracixalus sp. Most species of the first clade have a relatively slender habitus and a green dorsum, no obvious X-shaped pattern or an X-shaped pattern composed of narrow stripes presents on the dorsum with blank upper half of the X-pattern, their skin is usually semitransparent, especially on the ventral surface, with faintly visible bones and internal organs, they usually inhabit low trees or shrubs at low to medium altitudes, and lay small number of eggs on leaves overhanging small pools of water or moist areas in the vicinity of streams [2,26,28,29,30,31], while most species of the second clade have a relatively robust habitus and a brown dorsum, an X-shaped pattern composed of wide stripes presents on the dorsum with solid upper half of the X-pattern, their skin is usually opaque, they usually inhabit tall tree crowns at medium to high altitudes, and lay eggs in tree holes or open or semi open bamboo tubes with water inside [3,4,9,32,33]. In addition, there are differences in the advertisement calls between these two clades. The calls are non-stereotypical, highly variable in structure, duration, amplitude and frequency in the same species even in the same individual of the first clade, while the calls are relatively stereotypical, composed of a whistle plus none or several clicks in the same species of the second clade [2,3,26,34,35,36]. Although there is no available molecular data for G. carinensis, we consider that this species belongs to the second clade due to the morphological similarity between it and G. sapaensis. In summary, we agree with Rowley et al. [20] that the two clades within Gracixalus represent two different genera. However, we do not formally split this genus here, pending the addition of more molecular data, such as nuclear gene fragments, to obtain more stable and reliable conclusions.
The species diversity of this genus has been severely underestimated before; however, the taxonomy of this genus is a challenging task. Species of this genus often inhabit tall trees and rarely move to lower areas, making it relatively difficult to collect specimens of this genus. Therefore, the number of specimens of some species of this genus are very limited. This is not conducive to morphological statistical analysis, as the sample size is too small to have significant statistical significance. Furthermore, the available gene fragments for species of this genus are also limited. Recent phylogenetic analyses of this genus mostly focused on the mitochondrial 16S ribosomal RNA gene [2,4,8,9]. Although the genetic distances between some species of this genus were small in 16S, being less than 3.0% or even less than 2.0%, since this gene is highly conserved, these small genetic distances do not indicate that they are necessarily intraspecific. In addition, for some species of this genus, the available gene fragments are very short. Perhaps these small genetic distances are caused by the short length of gene fragments. Moreover, there have been few studies on the nuclear genes of this genus. Tran et al. [14] performed the split network analysis based on the nuclear gene Rag1; however, they did not conduct haplotype analysis. For the subsequent work on the taxonomy of this genus, we suggest strengthening the collection of specimens, including both known and unknown species, as well as obtaining long mitochondrial gene fragments and nuclear genes for phylogenetic analysis. This would facilitate obtaining more stable morphological analyses and more credible phylogenetic trees as well as more clear haplotype networks.
Previously, there were three species of Gracixalus recorded in Yunnan Province, namely G. gracilipes, distributed in Hekou, Pingbian, Xichou, and Malipo counties of southeastern Yunnan; G. yunnanensis, distributed in Cangyuan, Lancang, Menghai, Lüchun, Jinping, and Yuanyang counties of southwestern and southern Yunnan; and G. quangi, only distributed in Mengla County of southern Yunnan [5]. Gracixalus wuliangshan sp. nov. is currently known only in Jingdong County of central western Yunnan. At present, no other species of Gracixalus have been found sympatric with the new species.
Although no eggs and tadpoles of the new species were observed, we found many natural tree holes in its habitat, some of which have accumulated water inside (Figure 5). Therefore, we speculate that the new species has a similar breeding habit as other members of the same clade, laying eggs in tree holes.
As the new species usually inhabits tall trees, it is difficult to collect, so we only obtained two specimens of this species. However, based on the abundance of advertisement calls of this species at the type locality, we conclude that its population is not low. In addition, this species is distributed within the legally protected nature reserve, and therefore, we consider that the new species is currently not endangered and we do not suggest the threatened assessment of it at present.
Author Contributions
Conceptualization, D.R., Z.L. and S.L.; methodology, Z.Y., C.W., N.L. and M.H.; formal analysis, S.L.; investigation, S.L., Z.Y., C.W. and N.L.; resources, Z.L.; writing—original draft preparation, S.L.; writing—review and editing, S.L. and D.R. All authors have read and agreed to the published version of the manuscript.
Funding
This research was funded by Biological Resources Programme, Chinese Academy of Sciences, the Position of Bioclassonomist of Chinese Academy of Sciences (grant no. CAS-TAX-24) and Foundation of Yunnan Key Laboratory of Biodiversity Information, Kunming Institute of Zoology, Chinese Academy of Sciences.
Data Availability Statement
All data are presented in this article and in GenBank (https://www.ncbi.nlm.nih.gov (accessed on 3 November 2025)).
Acknowledgments
We thank the forest ranger of Yunnan Wuliangshan National Nature Reserve Dazhi Wang for his help in collecting specimens. We are grateful to the editors and reviewers for their efforts on the manuscript.
Conflicts of Interest
The authors declare no conflicts of interest.
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Figure 1.
Bayesian phylogenetic tree of the genus Gracixalus inferred from 16S rRNA gene sequences. Numbers before and after “/” are Bayesian posterior probabilities and maximum likelihood bootstrap values (only values above 0.95/70 are shown), respectively.
Figure 1.
Bayesian phylogenetic tree of the genus Gracixalus inferred from 16S rRNA gene sequences. Numbers before and after “/” are Bayesian posterior probabilities and maximum likelihood bootstrap values (only values above 0.95/70 are shown), respectively.
Figure 2.
The type specimens of Gracixalus wuliangshan sp. nov. in preservative. (A) Dorsal view; (B) ventral view.
Figure 2.
The type specimens of Gracixalus wuliangshan sp. nov. in preservative. (A) Dorsal view; (B) ventral view.
Figure 3.
The holotype (KIZ2025067) of Gracixalus wuliangshan sp. nov. in life. (A) Dorsal view; (B) ventral view; (C) left view; (D) right view; (E) close-up view of the axilla, groin, and inner side of the thigh; (F) close-up view of the outer side of the thighs; (G) close-up view of the left palm; (H) close-up view of the left sole.
Figure 3.
The holotype (KIZ2025067) of Gracixalus wuliangshan sp. nov. in life. (A) Dorsal view; (B) ventral view; (C) left view; (D) right view; (E) close-up view of the axilla, groin, and inner side of the thigh; (F) close-up view of the outer side of the thighs; (G) close-up view of the left palm; (H) close-up view of the left sole.
Figure 4.
Map showing the type localities of the new species and closely related congeners. Star: Gracixalus wuliangshan sp. nov., square: G. tianlinensis, dot: G. sapaensis, and triangle: G. ziegleri. The elevation data were obtained from Geospatial Data Cloud (http://www.gscloud.cn/).
Figure 4.
Map showing the type localities of the new species and closely related congeners. Star: Gracixalus wuliangshan sp. nov., square: G. tianlinensis, dot: G. sapaensis, and triangle: G. ziegleri. The elevation data were obtained from Geospatial Data Cloud (http://www.gscloud.cn/).
Figure 5.
Some tree holes with water inside observed at the type locality of the new species.
Table 1.
Samples used in the phylogenetic analyses of this study.
| Species | Locality | Voucher no. | GenBank no. |
|---|---|---|---|
| Gracixalus wuliangshan sp. nov. | Jingdong, Yunnan, China | KIZ2025067 | PX600316 |
| Gracixalus wuliangshan sp. nov. | Jingdong, Yunnan, China | KIZ 036308 | PX600317 |
| Gracixalus ananjevae | Nghe An, Vietnam | VNMN 03012 | JN862546 |
| Gracixalus gracilipes | Lao Cai, Vietnam | AMS R 177672 | KT374014 |
| Gracixalus guangdongensis | Hunan, China | CIB HN201108200 | LC011936 |
| Gracixalus guangdongensis | Guangdong, China | SYS a005750 | MG520197 |
| Gracixalus huaping | Huaping, Guangxi, China | GXNU YU000480 | PP083976 |
| Gracixalus huaping | Huaping, Guangxi, China | GXNU YU000511 | PP083977 |
| Gracixalus jinggangensis | Jinggangshan, Jiangxi, China | SYS a003186 | KY624587 |
| Gracixalus jinggangensis | Jinggangshan, Jiangxi, China | SYS a003170 | KY624586 |
| Gracixalus jinxiuensis | Jinxiu, Guangxi, China | SYS a002183 | KY624585 |
| Gracixalus jinxiuensis | Jinxiu, Guangxi, China | SYS a002182 | KY624584 |
| Gracixalus liusanjieae | Guilin, Guangxi, China | GXNU YU000796 | PV746480 |
| Gracixalus liusanjieae | Guilin, Guangxi, China | GXNU YU000962 | PV746481 |
| Gracixalus lumarius | Kon Tum, Vietnam | AMS R 176202 | KF918412 |
| Gracixalus medogensis | Medog, Xizang, China | KIZ 010956 | MW023606 |
| Gracixalus nonggangensis | Longzhou, Guangxi, China | NHMG20091009 | JX841319 |
| Gracixalus nonggangensis | Longzhou, Guangxi, China | NHMG200910010 | JX841320 |
| Gracixalus patkaiensis | Northeast India | WII-ADA1352 | OQ940026 |
| Gracixalus quangi | Nghe An, Vietnam | AMS R173417 | JN862539 |
| Gracixalus quyeti | Cha Noi, Vietnam | VNUH 160706 | EU871428 |
| Gracixalus sapaensis | Lao Cai, Vietnam | ROM 39660 | GQ285670 |
| Gracixalus sapaensis | Lao Cai, Vietnam | KUHE 46401 | LC011938 |
| Gracixalus seesom | Kanchanaburi, Thailand | KUHE 35084 | LC011932 |
| Gracixalus supercornutus | Kon Tum, Vietnam | AMS R173887 | JN862545 |
| Gracixalus tianlinensis | Tianlin, Guangxi, China | NHMG 1705016 | MH117961 |
| Gracixalus tianlinensis | Tianlin, Guangxi, China | NHMG 1705015 | MH117960 |
| Gracixalus trieng | Kon Tum, Vietnam | AMS R176206 | MT328246 |
| Gracixalus trieng | Kon Tum, Vietnam | UNS 00342/AMS R176205 | MT328245 |
| Gracixalus truongi | Tuan Giao, Dien Bien, Vietnam | IEBR A.5006 | OP750514 |
| Gracixalus truongi | Tuan Giao, Dien Bien, Vietnam | IEBR A.5005 | OP750513 |
| Gracixalus weii | Leigongshan, Guizhou, China | MT LS20240518009 | PP852212 |
| Gracixalus weii | Leigongshan, Guizhou, China | MT LS20240518010 | PP852213 |
| Gracixalus yunnanensis | Menghai, Yunnan, China | KIZ 20160216 | MK234877 |
| Gracixalus yunnanensis | Lancang, Yunnan, China | KIZ 20160222 | MK234878 |
| Gracixalus ziegleri | Yen Bai, Vietnam | MCC.2018.35 | LC642813 |
| Gracixalus ziegleri | Yen Bai, Vietnam | MCC.2018.15 | LC642812 |
| Gracixalus sp. | Wenshan, Yunnan, China | 03320Rao | GQ285669 |
| Kurixalus idiootocus | Taiwan, China | KUHE 12979 | AB933306 |
Table 2.
Uncorrected pairwise genetic distances (%) based on 16S rRNA gene sequences.
| 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 1 Gracixalus wuliangshan sp. nov. | |||||||||||||||||||||||
| 2 Gracixalus ananjevae | 5.6 | ||||||||||||||||||||||
| 3 Gracixalus gracilipes | 10.6 | 10.8 | |||||||||||||||||||||
| 4 Gracixalus guangdongensis | 4.5 | 4.4 | 9.7 | ||||||||||||||||||||
| 5 Gracixalus huaping | 6.7 | 7.4 | 10.1 | 5.8 | |||||||||||||||||||
| 6 Gracixalus jinggangensis | 5.8 | 7.3 | 10.3 | 5.4 | 6.3 | ||||||||||||||||||
| 7 Gracixalus jinxiuensis | 6.9 | 7.1 | 10.5 | 5.5 | 2.9 | 6.9 | |||||||||||||||||
| 8 Gracixalus liusanjieae | 5.5 | 7.0 | 9.4 | 4.1 | 3.9 | 5.2 | 3.1 | ||||||||||||||||
| 9 Gracixalus lumarius | 14.3 | 13.5 | 13.0 | 13.9 | 15.0 | 14.3 | 15.9 | 13.9 | |||||||||||||||
| 10 Gracixalus medogensis | 9.4 | 10.7 | 12.1 | 10.5 | 11.0 | 10.0 | 10.6 | 10.7 | 16.1 | ||||||||||||||
| 11 Gracixalus nonggangensis | 6.8 | 7.6 | 12.4 | 6.2 | 9.1 | 7.1 | 8.1 | 7.8 | 15.5 | 10.5 | |||||||||||||
| 12 Gracixalus patkaiensis | 10.6 | 10.6 | 3.8 | 8.9 | 9.2 | 9.9 | 9.6 | 8.6 | 13.1 | 12.1 | 11.7 | ||||||||||||
| 13 Gracixalus quangi | 9.3 | 9.7 | 4.8 | 8.1 | 8.8 | 7.6 | 9.5 | 8.2 | 12.8 | 11.9 | 11.0 | 3.8 | |||||||||||
| 14 Gracixalus quyeti | 10.8 | 11.3 | 7.2 | 10.0 | 10.0 | 11.3 | 10.3 | 9.3 | 13.5 | 12.5 | 12.3 | 7.2 | 5.9 | ||||||||||
| 15 Gracixalus sapaensis | 2.5 | 6.4 | 10.4 | 5.1 | 6.6 | 6.2 | 6.8 | 5.2 | 15.5 | 9.8 | 7.7 | 10.2 | 9.2 | 10.9 | |||||||||
| 16 Gracixalus seesom | 8.7 | 10.4 | 6.2 | 8.3 | 8.8 | 9.4 | 9.2 | 7.4 | 15.5 | 10.8 | 9.9 | 5.3 | 6.0 | 8.8 | 8.2 | ||||||||
| 17 Gracixalus supercornutus | 10.9 | 11.0 | 6.4 | 9.2 | 10.3 | 9.0 | 10.7 | 9.9 | 14.1 | 12.1 | 12.1 | 5.5 | 2.7 | 6.0 | 10.7 | 7.2 | |||||||
| 18 Gracixalus tianlinensis | 2.3 | 6.3 | 10.5 | 4.6 | 6.2 | 6.5 | 5.8 | 4.9 | 14.7 | 10.1 | 7.0 | 9.4 | 9.7 | 10.4 | 3.3 | 7.4 | 11.1 | ||||||
| 19 Gracixalus trieng | 3.5 | 5.3 | 10.1 | 3.5 | 5.7 | 5.0 | 5.2 | 4.2 | 13.7 | 9.1 | 6.6 | 9.0 | 8.2 | 8.8 | 4.6 | 7.4 | 9.4 | 4.0 | |||||
| 20 Gracixalus truongi | 6.1 | 6.2 | 11.0 | 5.0 | 6.7 | 6.5 | 7.5 | 6.3 | 14.8 | 10.8 | 8.2 | 9.7 | 8.8 | 10.7 | 6.6 | 9.9 | 9.7 | 5.8 | 4.5 | ||||
| 21 Gracixalus weii | 4.9 | 5.5 | 8.3 | 3.6 | 3.3 | 6.0 | 3.3 | 1.8 | 13.8 | 11.5 | 5.9 | 10.1 | 7.9 | 9.7 | 4.6 | 7.8 | 9.3 | 3.8 | 4.0 | 6.1 | |||
| 22 Gracixalus yunnanensis | 4.2 | 4.0 | 10.5 | 2.1 | 6.0 | 5.0 | 5.8 | 4.9 | 13.7 | 9.7 | 6.6 | 9.7 | 8.6 | 10.2 | 5.0 | 8.5 | 9.4 | 4.3 | 4.0 | 4.7 | 4.0 | ||
| 23 Gracixalus ziegleri | 2.4 | 5.3 | 10.5 | 4.5 | 5.8 | 5.7 | 6.1 | 5.2 | 14.9 | 9.6 | 6.9 | 11.0 | 9.7 | 11.2 | 3.1 | 8.1 | 10.9 | 2.4 | 3.7 | 5.4 | 4.1 | 4.0 | |
| 24 Gracixalus sp. | 9.0 | 2.3 | 11.6 | 5.3 | 7.9 | 7.7 | 7.3 | 6.5 | 14.0 | 10.7 | 8.5 | 11.6 | 10.1 | 11.0 | 8.4 | 9.7 | 10.8 | 6.5 | 5.1 | 6.7 | 5.7 | 5.0 | 9.7 |
Table 3.
Measurements (mm) of the type specimens of Gracixalus wuliangshan sp. nov.
| KIZ2025067 Female | KIZ 036308 Male | KIZ2025067 Female | KIZ 036308 Male | ||
|---|---|---|---|---|---|
| SVL | 44.1 | 31.9 | DPE | 13.6 | 10.7 |
| HL | 14.9 | 11.5 | MN | 12.3 | 9.5 |
| HW | 16.0 | 12.6 | FLL | 31.1 | 22.9 |
| SNL | 6.7 | 5.0 | LAL | 24.7 | 18.3 |
| IN | 4.3 | 3.7 | 3FD | 3.0 | 2.4 |
| IOD | 4.4 | 3.4 | HLL | 69.1 | 51.7 |
| ED | 5.0 | 4.1 | TL | 22.9 | 17.3 |
| UEW | 3.6 | 3.1 | TW | 5.7 | 4.3 |
| NEL | 3.5 | 2.7 | TFL | 33.1 | 24.3 |
| TD | 2.3 | 2.2 | FL | 22.8 | 16.4 |
| TEL | 1.2 | 0.8 | 3TD | 2.2 | 1.6 |
| DAE | 8.3 | 6.8 | IMT | 2.3 | 1.4 |
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MDPI and ACS Style
Liu, S.; Yang, Z.; Wang, C.; Li, N.; Hou, M.; Luo, Z.; Rao, D. A New Species of the Genus Gracixalus (Anura, Rhacophoridae) from Central Western Yunnan, China. Taxonomy 2025, 5, 66. https://doi.org/10.3390/taxonomy5040066
AMA Style
Liu S, Yang Z, Wang C, Li N, Hou M, Luo Z, Rao D. A New Species of the Genus Gracixalus (Anura, Rhacophoridae) from Central Western Yunnan, China. Taxonomy. 2025; 5(4):66. https://doi.org/10.3390/taxonomy5040066
Chicago/Turabian StyleLiu, Shuo, Zhongfu Yang, Chunhua Wang, Nengping Li, Mian Hou, Zengyang Luo, and Dingqi Rao. 2025. "A New Species of the Genus Gracixalus (Anura, Rhacophoridae) from Central Western Yunnan, China" Taxonomy 5, no. 4: 66. https://doi.org/10.3390/taxonomy5040066
APA StyleLiu, S., Yang, Z., Wang, C., Li, N., Hou, M., Luo, Z., & Rao, D. (2025). A New Species of the Genus Gracixalus (Anura, Rhacophoridae) from Central Western Yunnan, China. Taxonomy, 5(4), 66. https://doi.org/10.3390/taxonomy5040066
