104 urn:lsid:arphahub.com:pub:f2cd1fff-21e4-581f-a7fa-850997197b7f urn:lsid:zoobank.org:pub:B1C81912-2D17-4CD8-8D2C-EFEAAAB2EF75 Vertebrate Zoology VZ 1864-5755 2625-8498 Senckenberg Gesellschaft für Naturforschung 10.3897/vz.76.e195550 195550 Research Article Amphibia Anura Megophryidae Systematics Taxonomy Survival in fragmented landscapes: Two new Boulenophrys species (Anura: Megophryidae) from the coastal mountains of eastern Guangdong, China Wang Jian https://orcid.org/0000-0003-4249-7767 1 Conceptualization Writing - original draft Zhan Bin-Bin 1 Writing - review and editing Xiao Wei-Wen 1 Investigation Tan Lin 2 Funding acquisition Chen Hong-Hui 1 Investigation Li Yuan-Hang https://orcid.org/0009-0009-6543-9245 1 Investigation Lin Shi-Shi lionrei@163.com 1 Funding acquisition Lyu Zhi-Tong lvzht@foxmail.com https://orcid.org/0000-0002-4891-3859 3 Data curation Zeng Zhao-Chi zhaochizeng@outlook.com 1 Data curation Guangdong Polytechnic of Environmental Protection Engineering, Foshan 528216, Guangdong, China Mountain Ecological Restoration and Biodiversity Conservation Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences Chengdu China https://ror.org/034t30j35 Guangdong Wildlife Monitoring Rescue and Conservation Center, Guangzhou 510520, Guangdong, China Guangdong Polytechnic of Environmental Protection Engineering Foshan China https://ror.org/04nat8y64 Mountain Ecological Restoration and Biodiversity Conservation Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610213, Sichuan, China Guangdong Wildlife Monitoring Rescue and Conservation Center Guangzhou China

Corresponding authors: Shi-Shi Lin (lionrei@163.com), Zhi-Tong Lyu (lvzt@cib.ac.cn), Zhao-Chi Zeng (zhaochizeng@outlook.com)

Academic editor Deepak Veerappan

 2026 02 07 2026 76 381 399 83421564-B92D-51C6-8AAC-537C2F836AB9 DE3C0022-A895-41F3-B1B8-72958DE590BB 14 04 2026 01 06 2026 Jian Wang, Bin-Bin Zhan, Wei-Wen Xiao, Lin Tan, Hong-Hui Chen, Yuan-Hang Li, Shi-Shi Lin, Zhi-Tong Lyu, Zhao-Chi Zeng 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. https://zoobank.org/DE3C0022-A895-41F3-B1B8-72958DE590BB

Abstract

Two new species, Boulenophrys donglisp. nov. and Boulenophrys raopingsp. nov., are described from the coastal mountains of Raoping County, Chaozhou City, eastern Guangdong, China, based on morphological and molecular evidence. Currently, these two new species are known only from low-altitude mountainous areas (below 500 m a.s.l.), where they inhabit fragmented patches of secondary forest interspersed with bamboo plantations. Given their extremely limited distribution, ongoing habitat loss, and isolated populations, we recommend listing these two new species as Critically Endangered (CR) in accordance with IUCN Red List criteria B1ab(i,ii,iii)c(i,ii)+B2ab(i,ii,iii)c(i,ii). Continued research is critical to fill existing knowledge gaps and to guide coordinated conservation efforts aimed at protecting these unique amphibians within the critical montane refugia of eastern Guangdong.

 Conservation highly threatened horned toads morphology mountain refugia phylogeny Introduction

The Chinese horned toads of the genus Boulenophrys Fei, Ye & Jiang, 2016, currently comprising 82 recognized species, are widespread in the subtropical and tropical regions of mainland East Asia (as of 17 April in Frost 2026; see our Table 1). The genus is primarily distributed in southern China, extending southward into the northernmost parts of Vietnam, Laos, Myanmar, and Thailand (Fei et al. 2012; Fei and Ye 2016; Lyu et al. 2023; Frost 2026). Species of Boulenophrys exhibit limited dispersal ability and narrow ecological niches, resulting in restricted distribution ranges for many taxa (Wang et al. 2019a; Lyu et al. 2023). The conservation of species in this genus faces considerable challenges, driven by the combined effects of multiple threatening factors. Additionally, high morphological conservatism and insufficient follow-up surveys have led to inadequate protection, largely due to misidentifications and data deficiencies (Lin et al. 2024). Therefore, understanding the true distribution pattern and surveying the current “gaps” of distribution for the genus Boulenophrys is necessary for informed conservation action. This is particularly true for regions with high human impacts, limited protection, and high biodiversity such as southeastern China (Wang et al. 2022, 2025c; Xu et al. 2024). Despite significant habitat degradation, remnant habitat patches in these areas may harbor overlooked populations or undescribed species. Given the genus’ limited dispersal and high habitat specificity, these fragmented landscapes could serve as the last refuges for genetically distinct lineages. Consequently, accurate extinction risk assessments for Boulenophrys species still require extensive field surveys and continuous data updates.

Literature for morphological characters of 82 recognized species of Boulenophrys.

ID Boulenophrys spp. References
1 B. acuta (Wang, Li & Jin, 2014) Lyu et al. (2023)
2 B. angka (Wu et al., 2019) Wu et al. (2019)
3 B. anlongensis (Li, Lu, Liu & Wang, 2020) Li et al. (2020), Lyu et al. (2023)
4 B. baishanzuensis (Wu et al., 2020) Wu et al. (2020)
5 B. baolongensis (Ye, Fei & Xie, 2007) Ye et al. (2007), Fei and Ye (2016)
6 B. binchuanensis (Ye & Fei, 1995) Ye and Fei (1995), Lyu et al. (2023)
7 B. binlingensis (Jiang, Fei & Ye, 2009) Fei et al. (2009), Lyu et al. (2023)
8 B. boettgeri (Boulenger, 1899) Lyu et al. (2023)
9 B. brachykolos (Inger & Romer, 1961) Lyu et al. (2023)
10 B. caobangensis (Nguyen et al., 2020) Nguyen et al. (2020)
11 B. caudoprocta (Shen, 1994) Shen (1994), Lyu et al. (2023)
12 B. congjiangensis (Luo et al., 2021) Lyu et al. (2023)
13 B. changyangensis Li et al., 2025 Li et al. (2025)
14 B. cheni (Wang & Liu, 2014) Lyu et al. (2023)
15 B. chishuiensis (Xu et al., 2020) Lyu et al. (2023)
16 B. daiyunensis (Lyu, Wang & Wang, 2021) Lyu et al. (2023)
17 B. dalaolingensis Wang, Chen & Liu, 2025 Wang et al. (2025b)
18 B. daoji (Lyu, Zeng, Wang & Wang, 2021) Lyu et al. (2023)
19 B. daweimontis (Rao & Yang, 1997) Rao and Yang (1997), Lyu et al. (2023)
20 B. daxuemontis Liu et al., 2025 Liu et al. (2025b)
21 B. dongguanensis (Wang & Wang, 2019) Lyu et al. (2023)
22 B. dupanglingensis Xiao & Mo, 2025 Xiao et al. (2025)
23 B. elongata Zeng et al., 2024 Zeng et al. (2024)
24 B. fengshunensis Wang, Zeng, Lyu & Wang, 2022 Lyu et al. (2023)
25 B. fanjingmontis (Zhang, Liang, Ran & Shen, 2012) Lyu et al. (2023)
26 B. fansipanensis (Tapley et al., 2018) Tapley et al. (2018a)
27 B. frigida (Tapley et al., 2021) Tapley et al. (2021)
28 B. gaolanensis Song et al., 2024 Song et al. (2024)
29 B. gutu Kuang, Wei & Shi, 2025 Kuang et al. (2025)
30 B. hoanglienensis (Tapley et al., 2018) Tapley et al. (2018a)
31 B. huangniushiensis Wang et al., 2025 Wang et al. (2025b)
32 B. hungtai Wang et al., 2022 Lyu et al. (2023)
33 B. hengshanensis Qian et al., 2023 Qian et al. (2023)
34 B. insularis (Wang et al., 2017) Lyu et al. (2023), Wang et al. (2017a)
35 B. jiangi (Liu et al., 2020) Lyu et al. (2023)
36 B. jingdongensis (Fei & Ye, 1983) Fei et al. (1983), Lyu et al. (2023)
37 B. jinggangensis (Wang, 2012) Lyu et al. (2023)
38 B. jiulianensis (Wang, Zeng, Lyu & Wang, 2019) Lyu et al. (2023)
39 B. kuatunensis (Pope, 1929) Lyu et al. (2023)
40 B. leishanensis (Li et al., 2018) Lyu et al. (2023)
41 B. lushanensis Shen, Zhou & Li, 2025 Shen et al. (2025a)
42 B. lushuiensis (Shi et al., 2021) Lyu et al. (2023)
43 B. lichun Lin et al., 2024 Lin et al. (2024)
44 B. liboensis (Zhang et al., 2017) Zhang et al. (2017)
45 B. lini (Wang & Yang, 2014) Lyu et al. (2023)
46 B. lishuiensis (Wang, Liu & Jiang, 2017) Lyu et al. (2023), Wang et al. (2017b)
47 B. meihuamontis Luo & Huang, 2025 Luo et al. (2025)
48 B. minor (Stejneger, 1926) Lyu et al. (2023)
49 B. mirabilis (Lyu, Wang & Zhao, 2020) Lyu et al. (2023)
50 B. mufumontana (Wang, Lyu & Wang, 2019) Lyu et al. (2023)
51 B. nankunensis (Wang, Zeng & Wang, 2019) Lyu et al. (2023)
52 B. nanlingensis (Lyu, Wang, Liu & Wang, 2019) Lyu et al. (2023)
53 B. nebulosa Shen & Li, 2025 Shen et al. (2025b)
54 B. obesa (Wang, Li & Zhao, 2014) Lyu et al. (2023)
55 B. ombrophila (Messenger & Dahn, 2019) Lyu et al. (2023)
56 B. omeimontis (Liu, 1950) Liu (1950), Lyu et al. (2023)
57 B. palpebralespinosa (Bourret, 1937) Fei et al. (2009), Lyu et al. (2023)
58 B. pepe Wang & Zeng, 2024 Wang et al. (2024)
59 B. puningensis Wang et al., 2022 Lyu et al. (2023)
60 B. qianbeiensis (Su et al., 2020) Lyu et al. (2023)
61 B. rubrimera (Tapley et al., 2017) Tapley et al. (2017, 2018b)
62 B. sangzhiensis (Jiang, Ye & Fei, 2008) Lyu et al. (2023)
63 B. sanmingensis (Lyu & Wang, 2021) Lyu et al. (2023)
64 B. shimentaina (Lyu, Liu & Wang, 2020) Lyu et al. (2023)
65 B. shuichengensis (Tian & Sun, 1995) Tian and Sun (1995), Tian et al. (2000), Fei and Ye (2016)
66 B. shunhuangensis (Wang et al., 2019) Lyu et al. (2023)
67 B. spinata (Liu & Hu, 1973) Hu et al. (1973), Lyu et al. (2023)
68 B. tongboensis (Wang & Lyu, 2021) Lyu et al. (2023)
69 B. tuberogranulata (Shen, Mo & Li, 2010) Mo et al. (2010), Fei and Ye (2016), Lyu et al. (2023)
70 B. wugongensis (Wang, Lyu & Wang, 2019) Lyu et al. (2023)
71 B. wuliangshanensis (Ye & Fei, 1995) Ye and Fei (1995), Lyu et al. (2023)
72 B. wushanensis (Ye & Fei, 1995) Ye and Fei (1995), Fei and Ye (2016), Lyu et al. (2023)
73 B. xiangnanensis (Lyu, Zeng & Wang, 2020) Lyu et al. (2023)
74 B. xianjuensis (Wang et al., 2020) Lyu et al. (2023)
75 B. xuefengmontis Lyu & Wang, 2023 Lyu et al. (2023)
76 B. yangchunensis Zhao et al., 2025 Zhao et al. (2025)
77 B. yangmingensis (Lyu, Zeng & Wang, 2020) Lyu et al. (2023)
78 B. yaoshanensis Qi et al., 2021 Lyu et al. (2023)
79 B. yezhongensis Liu et al., 2025 Liu et al. (2025a)
80 B. yingdeensis Qi, Lyu, Wang & Wang, 2021 Lyu et al. (2023)
81 B. youran Xiao, Shi & Mo, 2025 Xiao et al. (2025)
82 B. yunkaiensis Qi, Wang, Lyu & Wang, 2021 Lyu et al. (2023)

Following the discoveries of Boulenophrys insularis in 2017, B. sanmingensis in 2021, B. puningensis, B. hungtai, and B. fengshunensis in 2022, and B. elongata in 2024 (Wang et al. 2017a, 2022; Lyu et al. 2021; Zeng et al. 2024), we continued systematic field surveys across mountainous and hilly habitats in eastern Guangdong, China. These efforts yielded new distribution records for B. puningensis and B. hungtai, extending their known ranges, and resulted in the discovery of two new species from the coastal mountains of Raoping County (Fig. 1). Detailed accounts of these findings are presented in the following sections.

10.3897/vz.76.e195550.figure1 C72ADA1D-4040-5806-AD4C-568BD3F0D60F

Geographic distribution of Boulenophrys species in eastern Guangdong and B. meihuamontis in western Fujian, China. Shape and color indicate differential species status (pentagon: new species; circle: recognized species), whereas number corresponds to different localities.

https://binary.pensoft.net/fig/1703702 Materials and Methods Morphology

Thirteen newly collected specimens of two unnamed species from Raoping County, Chaozhou City, Guangdong, China, were examined in this study. All specimens examined were fixed in 10% buffered formalin and later transferred to 70% ethanol. All studied specimens have been deposited at the Herpetological Museum at the Chengdu Institute of Biology, Chinese Academy of Sciences (CIB), in Chengdu, Sichuan Province; the Guangdong Polytechnic of Environmental Protection Engineering (GEP), in Foshan, Guangdong Province; and the Museum of Biology, Sun Yat-sen University (SYS), in Guangzhou, Guangdong Province, China.

External measurements were recorded with a digital caliper (Neiko 01407A stainless steel 6-inch digital caliper) to the nearest 0.1 mm. These measurements are as follows: ED (eye diameter, from the anterior corner of the eye to posterior corner of the eye); FTL (foot length, from distal end of shank to the tip of digit IV); HDL (head length, from tip of snout to the articulation of the jaw); HDW (head width, head width at the commissure of the jaws); HND (hand length, from the proximal border of the outer palmar tubercle to the tip of digit III); IND (internasal distance, distance between nares); IOD (interorbital distance, minimum distance between upper eyelids); RAD (forearm or radiulna length, from the flexed elbow to the proximal border of the outer palmar tubercle); SNT (snout length, from tip of snout to the anterior corner of the eye); SVL (snout-vent length, from tip of snout to posterior margin of vent); TD (tympanum diameter, horizontal diameter of tympanum); TED (tympanum-eye distance, from anterior edge of tympanum to posterior corner of the eye); TIB (crus or tibiofibula length, from the outer surface of the flexed knee to the heel). Sex was determined by external secondary sexual characters, such as the presence of vocal sacs, nuptial pads, or nuptial spines in males and their absence in females (Fei et al. 2009). Morphological characters of all recognized species of the genus Boulenophrys used for comparisons were based on an examination of museum specimens (Table SS1) and information available in the literature (Table 1).

We used R v4.0.0 to conduct statistical analyses of morphometric measurements following the methods described in Lyu et al. (2025) for all relevant species. One-way analysis of variance (ANOVA) as well as principal component analysis (PCA) were included in the statistical analyses of morphometric measurements, with all measurements adjusted for allometric size correction prior to analysis (Chan and Grismer 2022). The PCA was followed by a permutational multivariate analysis of variances (PERMANOVA; Anderson 2001) and a permutational analysis of multivariate dispersions (PERMDISP; Anderson 2005) as implemented in the vegan R package (Oksanen et al. 2019), which test the null hypothesis of no differences in the position of cluster centroids (Anderson 2017). Both PERMANOVA and PERMDISP were run with 9.999 permutations, and their significance was tested through Monte Carlo simulations. Only male specimens (six from Dacheng Town, four from Qiandong Town, 13 B. hungtai, three B. meihuamontis and five B. insularis) were included in statistical analyses due to the limited availability of female specimens.

 Phylogeny

Nineteen newly collected samples were included in the phylogenetic analysis, including six from Dacheng Town, seven from Qiandong Town, four B. hungtai and two B. puningensis. DNA extraction, PCR amplification, and sequencing protocols follow that of Liu et al. (2018). Finally, 105 samples were used for the phylogenetic analyses, with the others downloaded from GenBank (https://www.ncbi.nlm.nih.gov/genbank). Two samples of the genus Xenophrys Günther, 1864 were used as outgroups (Table SS2). Two partial mitochondrial genes, the 16S ribosomal RNA (16S) and the cytochrome c oxidase 1 (COI), were used for the analysis.

Clustal X 2.0 (Thompson et al. 1997) was used to align the sequences with default parameters. Then we used PartitionFinder (Lanfear et al. 2012) to determine the optimal partitioning schemes, and the results showed that partitioning by gene was optimal for 16S, while partitioning by codon position was optimal for COI, with GTR+G+I identified as the best-fit nucleotide substitution model for all partitions. We constructed the phylogenetic trees using Bayesian inference (BI) using MrBayes v.3.2.4 (Ronquist et al. 2012), and maximum likelihood (ML) implemented in RaxmlGUI v.1.3 (Silvestro and Michalak 2012). For the ML analysis, an optimal tree was obtained and branch supports were evaluated with 1000 rapid bootstrapping replicates. For the BI analysis, two independent runs were conducted with each running for 10,000,000 generations and sampled every 1000 generations with the first 25% of samples discarded as burn-in, resulting in a potential scale reduction factor (PSRF) of < 0.01. Then, the remaining trees were used to create a consensus tree. Nodes having ML bootstrap values (BS) ≥ 70 and BI posterior probabilities (BPP) ≥ 0.90 were considered strongly supported. Genetic distances between all samples were calculated in MEGA 11 using uncorrected p distances based on the 16S rRNA gene, with all gaps removed.

 Results Molecular phylogenetic analysis

The BI and ML phylogenetic trees resulted in essentially identical topologies (ML phylogenetic topology in Fig. 2). The Boulenophrys samples from Liuhuang Town, Hongtuzhang of Fengshun (localities 8 and 9), and Liwangzhang of Wuhua (locality 6) were clustered with B. hungtai with strong support (BS = 99; BPP = 1.00), while the samples from Beiyang Town and Xintian Town of Luhe (localities 11 and 12) clustered with B. puningensis with strong support (BS = 100; BPP = 1.00). These results expand the distribution range of these two known species. Six samples from Dacheng Town of Raoping (locality 1) and seven samples from Qiandong Town of Raoping (locality 2) formed two monophyletic lineages with considerable branch length and strong support (BS = 100; BPP = 1.00, respectively). These two new lineages were clustered with B. hungtai, B. meihuamontis and B. insularis to form a major clade with strong support (BS = 97; BPP = 0.99), but relationship among these species remain unresolved. Genetic distance calculations further reveal that the interspecific divergences of these two unnamed lineages and known congeners are comparable to those of recognized species (Table SS3).

10.3897/vz.76.e195550.figure2 C5B5C858-F89F-5E51-AA26-444D8B9B2549

Maximum Likelihood tree using partial DNA sequences of the mitochondrial 16S rRNA and COI genes. Numbers at nodes before slashes indicate bootstrap support and numbers after slashes indicate Bayesian posterior probabilities.

https://binary.pensoft.net/fig/1703703 Morphological Analysis

Morphological analyses further supported significant differences among the five lineages in this major clade (including the two new lineages from Raoping, B. hungtai, B. meihuamontis and B. insularis). PCA Scatter plots demonstrated that specimens of this major clade were divided into five independent clusters (Fig. 3A). These five clusters corresponding to the five lineages were significantly different from each other as indicated by PERMANOVA (F = 75.38; R2 = 0.92; p < 0.001) and PERMDISP (F = 0.434; p = 0.784). ANOVA and boxplot results also revealed significant differences between the two new lineages and their closely related species (Fig. 3B, C). The specimens from Dacheng Town distinctly differ from B. hungtai in all morphological variables except IND and IOD; from B. meihuamontis in all morphological variables except IND, ED, and TD; and from B. insularis in all morphological variables except SNT and TD. The specimens from Qiandong Town obviously differ from B. hungtai in all morphological variables; from B. meihuamontis in all morphological variables; and from B. insularis in all morphological variables except TD. These two new lineages also showed significant differences from each other in HDL, SNT, IND, IOD, and ED. Further morphological examination also showed that the corresponding specimens of the two new lineages can be distinguished from all recognized congeners (see comparisons below). Thus, we describe the corresponding specimens from Dacheng Town and Qiandong Town, Raoping, as two new species, respectively.

10.3897/vz.76.e195550.figure3 550C3F29-3940-53AA-8194-F69136E8A059

Morphometric comparisons of male specimens of Boulenophrys donglisp. nov. (n = 6), Boulenophrys raopingsp. nov. (n = 4), B. hungtai (n = 13), B. meihuamontis (n = 3) and B. insularis (n = 5). A Scatter plot of PC1 (76.2%) and PC2 (14.4%) from Principal Component Analysis. Arrows indicate loadings of morphological variables. B Pairwise ANOVA result for all ten morphological variables. C Boxplots for all ten morphological variables.

https://binary.pensoft.net/fig/1703704 Taxonomic accounts Animalia Anura Megophryidae D19E20CA-31B0-59FC-8F86-0D7E2F914569 Boulenophrys dongli https://zoobank.org/203B7648-85F5-4F6F-A12C-0A2BBBD744F6 sp. nov. Figures 4 , 5A , 6 , 10A Holotype.

CHINA • ♂; Guangdong Province, Chaozhou City, Raoping County, Dacheng Town, Mt Dabo; 23.647000°N, 117.137122°E; ca. 405 m a.s.l.; 16 January 2026; Jian Wang and Bin-Bin Zhan leg.; GEP a514.

10.3897/vz.76.e195550.figure4 59CD060C-D58E-5019-968B-95C494F2D3E5

Life aspect of the holotype (GEP a514) of Boulenophrys donglisp. nov.: A dorsolateral view, B dorsal view, C ventral view, D ventral view of hand, E ventral view of foot.

https://binary.pensoft.net/fig/1703705 10.3897/vz.76.e195550.figure5 2C3C793E-C5E6-534A-9BE1-C421C77D6C56

Morphological comparison of Boulenophrys donglisp. nov. and B. insularis.

https://binary.pensoft.net/fig/1703706 Paratypes.

CHINA • 5 ♂; same data as for holotype; CIB 121752 (field number: GEP a509), SYS a009611 (field number: GEP a510), GEP a511–513.

 Etymology.

The specific epithet dongli is derived from the ancient name “东里 (dōng lǐ)” of the region where the new species is distributed, meaning the eastern boundary of Guangdong, indicating the distribution area of the new species.

 Diagnosis.

(1) Small size (SVL 30.3–35.6 mm in six adult males); (2) snout rounded in dorsal view, canthus rostralis well developed, tongue not notched distally; (3) tympanum visible, tympanic margin indistinct, temporal region bearing conical spines except tympanum; (4) vomerine ridges and vomerine teeth present; (5) dorsal skin rough and highly granular, discontinuous X-shaped or V-shaped ridge on center of dorsum, dorsolateral ridges absent, sparse large tubercles on flanks, dorsal limbs with tubercles bearing conical spines, sparse rounded tubercles on belly, dense tubercles bearing conical spines on ventral thighs and around cloaca; (6) outer margin of upper eyelid with a small horn-like prominent tubercle, supratympanic fold distinct and narrow; (7) two metacarpal tubercles distinct, inner one observably enlarged, relative finger lengths I < II < IV < III, distinct subarticular tubercle at base of each finger; (8) heels not meeting when hindlimbs folded, tibio-tarsal articulation reaching posterior margin of tympanum; (9) toes without webbing and lateral fringes, inner metatarsal tubercle long ovoid, outer one absent, relative toe length I < II < V < III < IV; (10) dorsal surface yellowish-brown to dark brown with indistinct irregular dark-brown patches and small orange dots, a dark-brown triangular marking present between eyes, dorsal limbs and digits light brown with dark-brown transverse bands and orange dots; and (11) nuptial pad bearing dense nuptial spines on dorsal bases of fingers I and II in breeding adult males, subgular vocal sac present in males.

 Comparisons.

Boulenophrys donglisp. nov. is phylogenetically most closely related to three recognized congeners, i.e., B. insularis, B. meihuamontis, and B. hungtai (Fig. 2). Detailed comparative data of the new species with its phylogenetically close congeners are listed in Table 2. The new species can be easily distinguished from B. insularis (Fig. 5) by its tongue not notched distally (vs tongue weakly notched distally), tympanic margin indistinct (vs tympanic margin distinct and raised), toes without webbing (vs toes with rudimentary webbing), temporal region bearing conical spines (vs spines absent), and dense tubercles bearing conical spines present on ventral thighs (vs spines absent). The new species can be easily distinguished from B. meihuamontis by its larger body size, with SVL 30.3–35.6 mm in six adult males (vs 26.9–28.3 mm in three adult males), vomerine ridges and vomerine teeth present (vs absent), dorsolateral ridges absent (vs present), tibio-tarsal articulation reaching posterior margin of tympanum (vs reaching posterior corner of eye), and toes without webbing (vs toes with rudimentary webbing). The new species can be easily distinguished from B. hungtai by its tympanic margin indistinct (vs tympanic margin distinct and raised), vomerine ridges and vomerine teeth present (vs absent), dorsolateral ridges absent (vs present), and tibio-tarsal articulation reaching posterior margin of tympanum (vs reaching the region between middle of tympanum to posterior corner of eye).

Diagnostic characters separating Boulenophrys donglisp. nov., B. raopingsp. nov., and their phylogenetic congeners.

ID Boulenophrys spp. B. dongli sp. nov. B. raoping sp. nov. B. insularis B. meihuamontis B. hungtai
1 SVL in mm (males) 30.3–35.6 34.1–36.6 36.8–41.2 26.9–28.3 25.8–33.3
(n = 6) (n = 4) (n = 5) (n = 3) (n = 13)
2 SVL in mm (females) / 39.2–41.8 47.1 / /
(n = 3) (n = 1)
3 Tongue rounded rounded weakly notched rounded rounded
4 Vomerine ridges present present present absent absent
5 Vomerine teeth present absent present absent absent
6 Tympanic margin indistinct distinct distinct indistinct distinct
7 Dorsolateral ridges absent absent absent present present
8 Conical spines on ventral thighs present present absent / present
9 Tibio-tarsal articulation stretched forward posterior margin of tympanum anterior margin of tympanum posterior margin of tympanum posterior corner of the eye middle of tympanum to posterior corner of eye
10 Webbing on toes absent rudimentary rudimentary rudimentary absent

Boulenophrys donglisp. nov. can be distinguished by its heels not meeting when hindlimbs folded, from B. anlongensis, B. baishanzuensis, B. binlingensis, B. congjiangensis, B. changyangensis, B. cheni, B. chishuiensis, B. dalaolingensis, B. daxuemontis, B. daweimontis, B. dupanglingensis, B. jiangi, B. jingdongensis, B. jinggangensis, B. jiulianensis, B. leishanensis, B. liboensis, B. lini, B. mirabilis, B. mufumontana, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. sangzhiensis, B. sanmingensis, B. shimentaina, B. shunhuangensis, B. spinata, B. tongboensis, B. wuliangshanensis, B. xianjuensis, B. yangmingensis, B. yezhongensis, and B. youran (vs heels overlapping); from B. baolongensis, B. binchuanensis, B. caudoprocta, B. elongata, B. fanjingmontis, B. gutu, B. nebulosa, B. xiangnanensis, and B. yangchunensis (vs heels just meeting); and from B. angka, B. daiyunensis, B. huangniushiensis, B. minor, B. xuefengmontis, B. yaoshanensis, B. yingdeensis, and B. yunkaiensis (vs heels just meeting or overlapping).

Boulenophrys donglisp. nov. can be distinguished by its vomerine teeth present, from B. acuta, B. boettgeri, B. brachykolos, B. daoji, B. gaolanensis, B. hengshanensis, B. kuatunensis, B. lishuiensis, B. lushuiensis, B. obesa, B. ombrophila, B. pepe, B. shuichengensis, B. tuberogranulata, B. wugongensis, and B. wushanensis (vs vomerine teeth absent). Boulenophrys donglisp. nov. can be distinguished by its toes without lateral fringes, from B. lushanensis and B. rubrimera (vs toes with narrow lateral fringes). Boulenophrys donglisp. nov. can be distinguished by its toes without webbing, from B. brachykolos, B. dongguanensis, B. fengshunensis, B. nankunensis, and B. puningensis (vs toes with rudimentary webbing). Boulenophrys donglisp. nov. can be distinguished from the remaining B. fansipanensis, B. frigida, B. hoanglienensis, and B. lichun by its indistinct margin of the tympanum (vs distinct) and absent dorsolateral ridges (vs present).

 Description of holotype.

Adult male. Body size small, SVL 34.6 mm. Head width larger than head length, HDW/HDL 1.14; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes moderate in size, ED/HDL 0.31, pupil vertical, near diamond-shaped; nostril obliquely ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanic region oblique, tympanum distinct, tympanic margin indistinct; tympanum moderate in size, upper margin in contact with supratympanic fold, lower margin in contact with upper lip, TD/ED 0.71; large ovoid choanae at base of maxilla; vomerine ridges and vomerine teeth present, maxillary teeth present; margin of tongue rounded, not notched distally; single subgular vocal sac present.

Forearm length 0.21 of SVL, hand length 0.26 of SVL; webbing absent between fingers, lateral fringes absent, relative finger length I < II < IV < III; tips of fingers slightly dilated, round; subarticular tubercles on base of fingers present, distinct; inner metacarpal tubercle observably enlarged, outer one slightly smaller; single nuptial pad bearing dense nuptial spines present on dorsal surface of first and second fingers, respectively. Hindlimbs short, tibio-tarsal articulation reaching forward to posterior margin of tympanum when hindlimb stretched along body; heels not meeting when flexed hindlimbs held at right angles to body axis; crus length 0.41 of SVL and foot length 0.51 of SVL; relative toe length I < II < V < III < IV; tips of toes round and slightly dilated; toes without lateral fringes and webbing; subarticular tubercles on base of toes present and distinct; inner metatarsal tubercle long ovoid and lacking outer metatarsal tubercle.

Dorsal skin rough and highly granular; sparse large tubercles on flanks; single horn-like prominent tubercle on edge of upper eyelid; dorsal limbs with tubercles bearing conical spines; supratympanic fold distinct and narrow; discontinuous V-shaped ridge on center of dorsum, dorsolateral ridges absent; ventral surface smooth, sparse rounded tubercles on belly; dense tubercles bearing conical spines on ventral thighs and around cloaca; distinct pectoral gland closer to axilla; single femoral gland positioned on posterior surface of thigh at midpoint between knee and cloaca.

 Coloration of holotype.

In life, dorsal surface grayish-brown with indistinct irregular dark-brown patches and small orange dots, with a dark-brown triangular marking between eyes. An indistinct vertical dark-brown band present below the eye. Tympanum light brown. Dorsal surface of limbs with dark-brown transverse bands, orange dots and grayish-white tubercles. Tubercles on edge of upper eyelids orange. Supratympanic fold light brown with orange dots and irregular dark brown patches. Ventral surface pinkish-brown with grayish-white mottling and orange dots, a black longitudinal band on surface of throat; digits, subarticular tubercles, inner and outer metacarpal tubercles and inner metatarsal tubercle grayish-white. Pectoral glands and femoral glands grayish-white. Conical spines on temporal region, ventral thighs and around cloaca black. Iris silver with orange and dark brown mottling.

In preservative, dorsal surface fades to dark brown, with markings and patches more distinct, and orange dots absent. Ventral surface fades to dark brown, with the longitudinal band on surface of throat indistinct, grayish-white mottling more distinct, orange dots absent.

 Variation.

Morphometric variations listed in Table 3. Most of the paratypes are similar to the holotype in morphology and color pattern, except for the following (Fig. 6): dorsal coloration varies from yellowish-brown to dark brown; discontinuous X-shaped ridge on center of dorsum in the paratypes CIB 121752, SYS a009611, GEP a511; ventral surface with more distinct and denser grayish-white mottling and orange dots in the paratypes CIB 121752, SYS a009611, GEP a511; belly with dark brown dots in the paratypes SYS a009611, GEP a511; iris brown with dark brown mottling in the paratype GEP a511, coppery with dark brown mottling in the paratype GEP a512.

10.3897/vz.76.e195550.figure6 ACCB59C8-61AD-5924-8898-4C020CA9864C

Life aspect of the paratypes of Boulenophrys donglisp. nov.: A1–A3CIB 121752, B1–B3SYS a009611, C1–C3GEP a511, D1–D3GEP a512, E1–E3GEP a513.

https://binary.pensoft.net/fig/1703707

Measurements (in mm) of voucher specimens of Boulenophrys donglisp. nov. The asterisk indicates the holotype.

Voucher CIB 121752 SYS a009611 GEP a511 GEP a512 GEP a513 GEP a514*
Sex ♂ ♂ ♂ ♂ ♂ ♂
SVL 34.6 35.6 30.3 31.7 35.5 34.6
HDL 12.1 12.3 10.6 11.6 11.6 12.2
HDW 14.3 14.2 12.4 13.4 14.3 13.9
SNT 4.3 4.5 3.7 4.2 4.5 4.1
IND 3.9 4.0 3.2 3.3 4.2 3.5
IOD 3.7 3.6 2.9 3.1 3.5 3.4
ED 4.1 3.7 3.4 3.8 3.9 3.7
TD 3.3 3.0 2.1 2.4 2.7 2.6
TED 1.9 1.7 1.7 1.6 2.1 1.7
HND 8.9 9.6 7.7 8.5 8.3 8.9
RAD 7.4 7.8 6.8 7.1 7.6 7.3
TIB 13.7 14.0 12.5 13.3 13.3 14.0
FTL 19.2 19.9 18.0 18.5 20.6 19.8
Natural history, distribution, and threats.

Boulenophrys donglisp. nov. inhabits flowing montane streams and the nearby forest floor and leaf litter. The habitat consists of secondary forest interspersed with bamboo plantations (Fig. 7). Advertisement calls of males were heard from December to February, with calling individuals found in crevices under large rocks.

10.3897/vz.76.e195550.figure7 EB8CBF4A-ADD2-5B34-BF25-007D8E4BDB63

Aerial perspective view and microhabitat of the type locality of Boulenophrys donglisp. nov. in Mt Dabo, Dacheng Town, Raoping County, Chaozhou City, Guangdong Province, China.

https://binary.pensoft.net/fig/1703708

Comprehensive field surveys were conducted repeatedly across multiple elevational gradients in the surrounding regions and the mountainous areas of adjacent Fujian Province. However, no populations of B. donglisp. nov. were found outside its type locality (Fig. 1, locality 1) at elevations of 120–500 m a.s.l., suggesting that its distribution is highly restricted.

The main threats to B. donglisp. nov. at Mount Dabo include: (1) Natural environmental vulnerability, as the species occurs in low-lying coastal hills with hydrologically variable streams and a sensitive subtropical monsoon climate; (2) habitat degradation and land use, represented by extensive deforestation, conversion to monoculture plantations such as eucalyptus, and the prevalence of low-biodiversity secondary forests; (3) water resource diversion, whereby streams and small water sources in secondary forests are commonly diverted for agricultural and domestic use, reducing suitable aquatic and riparian habitat; (4) fragmented land tenure, including collective forests, private hills, and contracted lands, which complicates coordinated habitat conservation and restoration; and (5) ecological instability, whereby human disturbance interacts with naturally variable hydrology and coastal climatic conditions to exacerbate habitat instability and impede ecosystem recovery.

 Animalia Anura Megophryidae E8BC028D-7F35-52EA-AF18-5C1AA855C36A Boulenophrys raoping https://zoobank.org/50C97F34-D4E3-409F-927A-2BD1A3EDB1C5 sp. nov. Figures 8 , 9 , 10B Holotype.

CHINA • ♂; Guangdong Province, Chaozhou City, Raoping County, Qiandong Town, Mt Dabei; 23.662520°N, 116.867378°E; ca. 210 m a.s.l.; 24 January 2026; Jian Wang, Bin-Bin Zhan and Wei-Wen Xiao leg.; GEP a519.

10.3897/vz.76.e195550.figure8 336F9F95-C2F3-568C-9102-B242930451E3

Life aspect of the holotype (GEP a519) of Boulenophrys raopingsp. nov.: A1 dorsolateral view, A2 dorsal view, A3 ventral view, A4 ventral view of hand, A5 ventral view of foot; female paratype (GEP a523): B1 dorsolateral view, B2 dorsal view, B3 ventral view, B4 ventral view of hand, B5 ventral view of foot.

https://binary.pensoft.net/fig/1703709 10.3897/vz.76.e195550.figure9 628F40E5-D220-5357-9401-4BEE92945B4A

Life aspect of the paratypes of Boulenophrys raopingsp. nov.: A1–A3GEP a521, female; B1–B3CIB 121754, female; C1–C3GEP a517, male; D1–D3SYS a009628, male; E1–E3CIB 121753, male.

https://binary.pensoft.net/fig/1703710 10.3897/vz.76.e195550.figure10 7EE0A790-E6DC-5813-97D9-13DCEFBB2326

Holotypes in preservative of Boulenophrys donglisp. nov.: A1 dorsal view, A2 ventral view, A3 lateral view of head showing indistinct margin of tympanum, A4 ventral view of foot; and Boulenophrys raopingsp. nov.: B1 dorsal view, B2 ventral view, B3 lateral view of head showing distinct margin of tympanum, B4 ventral view of foot.

https://binary.pensoft.net/fig/1703711 Paratypes.

CHINA • 3 ♂; same data as for holotype; CIB 121753 (field number: GEP a520), SYS a009628 (field number: GEP a 518), GEP a517 • 3 ♀; same data as for holotype; CIB 121754 (field number: GEP a522), GEP a521, GEP a 523.

 Etymology.

The specific epithet raoping is derived from its type locality, Raoping County. The name “Raoping (饶平)” combines the characters “饶” (ráo), meaning “rich” or “abundant,” symbolizing prosperous harvests and natural resources, and “平” (píng), meaning “peace” or “stability,” reflecting aspirations for social harmony. Together, they convey the traditional Chinese ideal of material prosperity and enduring peace.

 Diagnosis.

(1) Small size (SVL 34.1–36.6 mm in four adult males, 39.2–41.8 mm in three adult females); (2) snout rounded in dorsal view, canthus rostralis well developed, tongue not notched distally; (3) tympanum distinct, tympanic margin distinct and raised, temporal region bearing conical spines except tympanum; (4) vomerine ridges present, vomerine teeth absent; (5) dorsal skin rough and highly granular, discontinuous X-shaped or V-shaped ridge on center of dorsum, dorsolateral ridges absent, sparse large tubercles on flanks, dorsal limbs with tubercles bearing conical spines, dense rounded tubercles on belly, dense tubercles bearing conical spines on ventral thighs and around cloaca; (6) outer margin of upper eyelid with a small horn-like prominent tubercle, supratympanic fold distinct and narrow; (7) two metacarpal tubercles distinct, inner one observably enlarged, relative finger lengths I < II < IV < III, distinct subarticular tubercle at base of each finger; (8) heels not meeting when hindlimbs folded, tibio-tarsal articulation reaching anterior margin of tympanum; (9) toes with rudimentary webbing and without lateral fringes, inner metatarsal tubercle long ovoid, outer one absent, relative toe length I < II < V < III < IV; (10) dorsal surface yellowish-brown to dark brown, with indistinct irregular dark-brown patches and small orange dots, with a dark-brown triangular marking between eyes, dorsal limbs and digits light brown with dark-brown transverse bands and orange dots; and (11) a single nuptial pad bearing dense nuptial spines on dorsal bases of fingers I and II in breeding adult males, subgular vocal sac present in males.

 Comparisons.

Boulenophrys raopingsp. nov. differs from B. donglisp. nov. (Fig. 10) by its vomerine teeth absent (vs present), rudimentary webbing on toes present (vs absent), tympanic margin distinct and raised (vs indistinct), and tibio-tarsal articulation reaching anterior margin of tympanum (vs reaching posterior margin of tympanum).

Boulenophrys raopingsp. nov. is phylogenetically most closely related to three recognized congeners, i.e., B. meihuamontis, B. insularis, and B. hungtai (Fig. 2). Detailed comparative data of the new species with its phylogenetically close congeners are listed in Table 2. The new species can be easily distinguished from B. meihuamontis by its larger body size, with SVL 34.1–36.6 mm in four adult males (vs 26.9–28.3 mm in three adult males), vomerine ridges present (vs absent), and margin of tympanum distinct and raised (vs indistinct), dorsolateral ridges absent (vs present). The new species can be easily distinguished from B. insularis by having no vomerine teeth (vs present), distally rounded tongue (vs distally weakly notched), tibio-tarsal articulation reaching anterior margin of tympanum (vs reaching posterior margin of tympanum), and dense tubercles bearing conical spines present on thighs and around cloaca (vs absent). The new species can be easily distinguished from B. hungtai by its vomerine ridges present (vs absent), absent dorsolateral ridges (vs present), and rudimentary webbing on toes (vs absent).

Boulenophrys raopingsp. nov. can be distinguished by its heels not meeting when hindlimbs folded, from B. anlongensis, B. baishanzuensis, B. binlingensis, B. congjiangensis, B. changyangensis, B. cheni, B. chishuiensis, B. dalaolingensis, B. daxuemontis, B. daweimontis, B. dupanglingensis, B. jiangi, B. jingdongensis, B. jinggangensis, B. jiulianensis, B. leishanensis, B. liboensis, B. lini, B. mirabilis, B. mufumontana, B. nanlingensis, B. omeimontis, B. palpebralespinosa, B. qianbeiensis, B. sangzhiensis, B. sanmingensis, B. shimentaina, B. shunhuangensis, B. spinata, B. tongboensis, B. wuliangshanensis, B. xianjuensis, B. yangmingensis, B. yezhongensis, and B. youran (vs heels overlapping); from B. baolongensis, B. binchuanensis, B. caudoprocta, B. elongata, B. fanjingmontis, B. gutu, B. nebulosa, B. xiangnanensis, and B. yangchunensis (vs heels just meeting); and from B. angka, B. daiyunensis, B. huangniushiensis, B. minor, B. xuefengmontis, B. yaoshanensis, B. yingdeensis, and B. yunkaiensis (vs heels just meeting or overlapping).

Boulenophrys raopingsp. nov. can be distinguished by its vomerine teeth absent, from B. brachykolos, B. daweimontis, B. dongguanensis, B. fansipanensis, B. fengshunensis, B. frigida, B. hoanglienensis, B. lichun, B. lushanensis, B. nankunensis, B. puningensis, and B. rubrimera (vs vomerine teeth present). Boulenophrys raopingsp. nov. can be distinguished by its toes without lateral fringes, from B. daoji and B. lushuiensis (vs toes with narrow lateral fringes); from B. shuichengensis (vs toes with wide lateral fringes). Boulenophrys raopingsp. nov. can be distinguished by its toes with rudimentary webbing, from B. gaolanensis, B. hengshanensis, B. kuatunensis, B. lishuiensis, B. ombrophila, and B. pepe (vs toes without webbing); from B. wushanensis (vs toes with more than one-fourth webbing). Boulenophrys raopingsp. nov. can be distinguished by its vomerine ridges present, from B. tuberogranulata and B. wushanensis (vs absent). Boulenophrys raopingsp. nov. can be distinguished from the remaining B. acuta, B. caobangensis, B. obesa, and B. wugongensis by its absent dorsolateral ridges (vs present).

 Description of holotype.

Adult male. Body size small, SVL 36.6 mm. Head width larger than head length, HDW/HDL 1.10; snout rounded in dorsal view, projecting, sloping backward to mouth in profile, protruding well beyond margin of lower jaw; top of head flat; eyes moderate in size, ED/HDL 0.32, pupil vertical, near diamond-shaped; nostril obliquely ovoid; canthus rostralis well developed; loreal region slightly oblique; internasal distance slightly larger than interorbital distance; tympanic region oblique, tympanum distinct, tympanic margin distinct and raised; tympanum moderate in size, upper margin in contact with supratympanic fold, lower margin in contact with upper lip, TD/ED 0.60; large ovoid choanae at base of maxilla; vomerine ridges present, vomerine teeth absent, maxillary teeth present; margin of tongue rounded, not notched distally; single subgular vocal sac present.

Forearm length 0.19 of SVL, hand length 0.24 of SVL; webbing absent between fingers, lateral fringes absent, relative finger length I < II < IV < III; tips of fingers slightly dilated, round; subarticular tubercles on base of fingers present, distinct; inner metacarpal tubercle observably enlarged, outer one slightly smaller; single nuptial pad bearing dense nuptial spines present on dorsal surface of first and second fingers, respectively. Hindlimbs short, tibio-tarsal articulation reaching forward to anterior margin of tympanum when hindlimb stretched along body; heels not meeting when flexed hindlimbs held at right angles to body axis; crus length 0.36 of SVL and foot length 0.52 of SVL; relative toe length I < II < V < III < IV; tips of toes round and slightly dilated; toes with rudimentary webbing and without lateral fringes; subarticular tubercles on base of toes present and distinct; inner metatarsal tubercle long ovoid and lacking outer metatarsal tubercle.

Dorsal skin rough and highly granular; sparse large tubercles on flanks; single horn-like prominent tubercle on edge of upper eyelid; dorsal limbs with tubercles bearing conical spines; supratympanic fold distinct and narrow; discontinuous V-shaped ridge on center of dorsum, dorsolateral ridges absent; ventral surface smooth, large rounded tubercles on belly; dense tubercles bearing conical spines on ventral thighs and around cloaca; distinct pectoral gland closer to axilla; single femoral gland positioned on posterior surface of thigh at midpoint between knee and cloaca.

 Coloration of holotype.

In life, dorsal surface orange-brown with indistinct irregular dark-brown patches and small orange dots, with a dark-brown triangular marking between eyes. An indistinct vertical dark-brown band present below the eye. Tympanum grayish-brown. Dorsal surface of limbs with dark-brown transverse bands, orange dots and grayish-white tubercles. Tubercles on edge of upper eyelids grayish-white. Supratympanic fold light brown with orange dots and irregular dark brown patches. Iris orange-brown with dark brown mottling. Ventral surface dark brown with grayish-white mottling and orange dots, a black longitudinal band on surface of throat; digits, subarticular tubercles, inner and outer metacarpal tubercles and inner metatarsal tubercle grayish-white. Pectoral glands and femoral glands grayish-white. Conical spines on temporal region grayish-white, on ventral thighs and around cloaca black.

In preservative, dorsal surface fades to dark brown, with markings and patches more distinct, and orange dots absent. Ventral surface fades to dark brown, with the longitudinal band on surface of throat and grayish-white mottling more distinct, orange dots absent.

 Variation.

Morphometric variations listed in Table 4. Most of the paratypes are similar to the holotype in morphology and color pattern, except for the following (Figs 8, 9): dorsal coloration varies from yellowish-brown to dark brown; discontinuous X-shaped ridge on center of dorsum in the female paratypes GEP a521 and CIB 121754; ventral surface of throat and chest with reddish orange patches in the female paratypes GEP a521 and CIB 121754; ventral markings and patches rather irregular in the male paratype GEP a517, and indistinct in the female paratype GEP a523. Nuptial pads, nuptial spines and vocal sac absent in the female paratypes GEP a521, 523 and CIB 121754.

Measurements (in mm) of voucher specimens of Boulenophrys raopingsp. nov. The asterisk indicates the holotype.

Voucher GEP a517 GEP a519* SYS a009628 CIB 121753 CIB 121754 GEP a521 GEP a523
Sex ♂ ♂ ♂ ♂ ♀ ♀ ♀
SVL 36.3 36.6 34.1 34.8 39.2 41.8 40.8
HDL 12.2 13.1 12.1 12.0 14.0 13.9 14.3
HDW 14.6 14.4 13.7 13.5 15.3 16.5 16.0
SNT 4.8 4.6 4.4 4.5 5.1 4.5 5.1
IND 4.0 4.2 4.0 4.0 4.0 4.2 4.2
IOD 3.6 3.9 3.6 3.6 3.9 4.0 4.0
ED 4.0 4.2 4.0 4.0 4.0 4.4 4.5
TD 2.6 2.5 2.4 2.7 2.8 2.8 3.2
TED 1.6 1.9 1.6 2.0 2.4 2.1 2.1
HND 8.8 8.9 8.4 8.1 9.5 10.3 10.3
RAD 7.3 7.0 7.0 7.4 8.3 9.1 8.5
TIB 13.7 13.2 13.0 13.0 14.6 15.9 15.4
FTL 19.7 19.0 19.4 18.8 21.6 23.6 22.6
Natural history, distribution, and threats.

Boulenophrys raopingsp. nov. inhabits flowing montane streams and the nearby forest floor and leaf litter. The habitat consists of secondary forest interspersed with bamboo plantations (Fig. 11). Advertisement calls of males were heard from December to February, with calling males found in crevices beneath large rocks. Two female paratypes, GEP a521 and CIB 121754, had completed oviposition, whereas specimen GEP a523 possessed mature internal oocytes.

10.3897/vz.76.e195550.figure11 E4BBC53F-6DF5-5A04-8C78-EF517094E744

Aerial perspective view and microhabitat of the type locality of Boulenophrys raopingsp. nov. in Mt Dabei, Qiandong Town, Raoping County, Chaozhou City, Guangdong Province, China.

https://binary.pensoft.net/fig/1703712

Comprehensive field surveys were conducted repeatedly across multiple elevational gradients in the surrounding regions and the mountainous areas of adjacent Fujian Province. However, no populations of B. raopingsp. nov. were found outside its type locality (Fig. 1, locality 2) at elevations of 150–350 m a.s.l., suggesting that its distribution is highly restricted.

The main threats to B. raopingsp. nov. at Mt Dabei include: (1) Impacts of eucalyptus plantations, where expanding monocultures intensify water scarcity, degrade soil conditions, and reduce understory biodiversity; these changes alter microclimates and increase resource competition, with no effective local policy protections in place; (2) fragmented land tenure, including collective forests, private hills, and contracted lands, which complicates coordinated habitat conservation and restoration; (3) water resource diversion, in which streams and water sources within secondary forests are often diverted for agricultural and domestic irrigation, thereby reducing suitable habitat for the species; and (4) ecological instability, whereby human activities interact with naturally variable hydrology and coastal climatic conditions to amplify habitat instability and impede ecosystem recovery.

 Discussion

As emphasized by multiple recent studies, genetic distance is now considered inappropriate as key evidence for species delimitation, particularly in amphibian groups (e.g., Hillis 2019; Chan et al. 2022; Lyu et al. 2023). Therefore, the recognition of B. donglisp. nov. and B. raopingsp. nov. in this work are primarily based on their independent phylogenetic placements and distinct morphological diagnoses compared with all other known congeners, and the p distances are used only for comparison with its congeners. The genetic distance in 16S rRNA gene between B. donglisp. nov. and its closely relate species range from 1.0–1.8%, and those between B. raopingsp. nov. and its closely related species range from 1.6–2.5%. Such values are greater than or comparable to the interspecific distances among several recognized species, including 1.0% between B. wugongensis and B. hengshanensis, 0.8% between B. binlingensis and B. qianbeiensis, 1.1% between B. fanjingmontis and B. sangzhiensis, 1.3% between B. xiangnanensis and B. yingdeensis, 1.3% between B. tuberogranulata and B. wushanensis, 1.5% between B. leishanensis and B. changyangensis, and only 1.7% between B. boettgeri and B. liboensis (two species with obviously distinguishable morphological characteristics). Other similar cases can be found in other congeners (Table SS3).

Inadequate field surveys significantly impede accurate conservation assessments for threatened Boulenophrys species. Such assessments are essential for understanding the impacts of habitat loss on endemic herpetofauna and for informing the designation of protected areas (Lin et al. 2024; Zeng et al. 2024; Zhao et al. 2025). Recent supplementary surveys in eastern Guangdong Province, China highlight this challenge and its broader conservation implications. For instance, although the discovery of the two new Boulenophrys species has enhanced regional biodiversity records, it has also refined the known distribution of B. insularis, confirming that this species is restricted to the mountainous western part of Nan’ao Island (Wang et al. 2017a; Lyu et al. 2023; this study). Furthermore, despite minor expansions in the documented ranges of B. hungtai and B. puningensis, their distributions remain highly localized and fragmented, a pattern consistent with strong habitat specialization and high vulnerability to environmental change (this study). In particular, for the two new species described in this work, repeated and multi-elevation field surveys were carried out across surrounding areas and mountainous terrains of adjacent Fujian Province. Nevertheless, each species is presently known exclusively from a single locality, both situated within coastal mountain ranges. The two localities are close, with a direct distance of less than 50 km, but are separated by the Huanggang River Delta, which might have acted as an isolation barrier during the speciation process of these two congeners. Given their extremely narrow geographic range, sole known occurrence at the type locality, and multiple ongoing anthropogenic and natural threatening factors, these species face severe survival risks. Accordingly, we recommend them to be listed as Critically Endangered (CR) based on the IUCN Red List criteria B1ab(i,ii,iii)c(i,ii)+B2ab(i,ii,iii)c(i,ii) (IUCN Standards and Petitions Committee 2024).

The hilly and mountainous landscapes of eastern Guangdong function as crucial refuges for numerous threatened amphibians, including several urodeles and anurans of conservation priority (Wang et al. 2022, 2024; Zeng et al. 2024). Nevertheless, long-term monitoring has revealed continuous habitat degradation driven by anthropogenic pressures, including land-use conversion, infrastructure development, and climate variability (Figs 7, 11). This ongoing decline underscores the urgent need for integrated, evidence-based conservation strategies. To resolve these challenges, filling critical knowledge gaps through continued field research is indispensable. Translating scientific insights into coordinated on-the-ground actions, including habitat restoration, targeted protection of key sites, and landscape connectivity planning, will enable effective mitigation of further habitat loss and support the recovery of eastern Guangdong’s unique amphibian fauna.

 Acknowledgements

We sincerely thank the editor and reviewers for their constructive comments and revisions. This work was supported by the DFGP Project of Fauna of Guangdong-202115, the Youth Innovative Talents Program of Guangdong Colleges and Universities (2024KQNCX316), the Project of Guangdong Innovation Team for the Higher-educational Institution (2024KCXTD078), Biological Resources Programme, Chinese Academy of Sciences (CAS-TAX-24-052), the Project of resource monitoring, epidemic diseases monitoring and rescue capability of wildlife (monitoring and conservation of wildlife under special state protection) in 2025 (ZDGZ25-ZFC08013), and the Integrated Project for Monitoring, Epidemic Pre-warning and Rescue of Key Protected Wildlife (Survey and Monitoring of Amphibians, Reptiles and Other Protected Wildlife in Eastern Guangdong) in 2026 (ZDGZ26-ZFC08007).

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Jian Wang and Bin-Bin Zhan contributed equally to this work.

 Supplementary materials 10.3897/vz.76.e195550.suppl1 A4CB08C3-3597-5CC3-964E-C27244512C31

Tables S1–S3

: .zip

Table SS1. Specimens of phylogenetically close congeners examined in this study. Asterisks indicate the holotype [.xlsx file]. — Table SS2. Collection localities, voucher data, and GenBank accession numbers for all samples used in this study[.xlsx file]. — Table SS3. Uncorrected p distances (percentage) between Boulenophrys species based on the 16S ribosomal RNA gene[.xlsx file].

 https://binary.pensoft.net/file/1703713 This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Wang J, Zhan BB, Xiao WW, Tan L, Chen HH, Li YH, Lin SS, Lyu ZT, Zeng ZC (2026)