Taxonomic Status of Three Recently Proposed Species of the Genus Kurixalus (Anura, Rhacophoridae), with Discussion on the ZooBank Registrations for Electronical Publication

Abstract

In the present study, we propose the synonymization of Kurixalus silvaenaias with Kurixalus qionglaiensis. This conclusion is based on morphological examination and phylogenetic analysis of a series of newly collected specimens, as well as the type series of both nomenclatures. The publication dates of the two species were determined according to the related articles of the International Code of Zoological Nomenclature. Of note, the ZooBank registrations for both species are invalid. Consequently, the publication of K silvaenaias does not qualify as a published work and the correct publication date of K qionglaiensis should be the date on which it was physically printed. Furthermore, based on the results presented in this study and its original description, the proposal of K inexpectatus is deemed untenable and should, therefore, be considered a junior synonym of K idiootocus. We further provide suggestions for the authors, editors, and publishers who are working on taxonomic publications, as well as for ZooBank to improve the registering service.

Headnote

Abstract In the present study, we propose the synonymization of Kurixalus silvaenaias with Kurixalus qionglaiensis. This conclusion is based on morphological examination and phylogenetic analysis of a series of newly collected specimens, as well as the type series of both nomenclatures. The publication dates of the two species were determined according to the related articles of the International Code of Zoological Nomenclature. Of note, the ZooBank registrations for both species are invalid. Consequently, the publication of K silvaenaias does not qualify as a published work and the correct publication date of K qionglaiensis should be the date on which it was physically printed. Furthermore, based on the results presented in this study and its original description, the proposal of K inexpectatus is deemed untenable and should, therefore, be considered a junior synonym of K idiootocus. We further provide suggestions for the authors, editors, and publishers who are working on taxonomic publications, as well as for ZooBank to improve the registering service.

Keywords Kurixalus inexpectatus, Kurixalus qionglaiensis, Kurixalus silvaenaias, nomenclature, the Code

1. Introduction

The genus Kurixalus Ye, Fei and Dubois, 1999, commonly referred to as frilled treefrogs, currently contains 24 species. Recent phylogenetic analyses have revealed three distinct clades within the genus (Nguyen et al., 2020; Guo et al., 2022). The basal clade consists of three congeners from the Malay Archipelago and Peninsular Malaysia, suggesting an ancient Sundaland origin. The second clade, previously classified under the genus Aquixalus Delorme, Dubois, Grosjean and Ohler, 2005 and currently synonymized with Kurixalus (Li et al., 2009), includes 10 species from mainland Southeast Asia and southernmost China. The third clade, representing Kurixalus sensu stricto, contains the remaining species predominantly distributed in subtropical regions of China and Japan, except for K. gracilloides Nguyen, Duong, Luu and Poyarkov, 2020 which is found from Vietnam.

Within the Kurixalus sensu stricto clade, three recently described congeners K. silvaenaias Hou, Peng, Miao, Liu, Li and Orlov, 2021, K. qionglaiensis Guo, Zhong, Leung, Wang and Hu, 2022, and K. inexpectatus Messenger, Yang, Börzéé, Chuang and Othman, 2022 have significantly expanded the geographical range of the genus to approximately 30° N (Hou et al., 2021; Guo et al., 2022; Messenger et al., 2022), representing the northernmost distribution of this tropical-origin genus. However, the taxonomic status of these species remains problematic. Kurixalus qionglaiensis was identified based on specimens collected from "Lugou Bamboo Sea ††^), Pingle Town, Qionglai City, Sichuan Province, southwestern China (30°2Г5Г N, 103° 18'30" E; elevation 615 m a.s.l.)", while K. silvaenaias was identified based on specimens collected from Tingle Town (30°36'16" N, 103°29' 69" E, 623 m elevation), Qionglai County [in error, should be Qionglai City], Chengdu Prefecture [in error, should be Chengdu City], Sichuan Province, China", which is in close proximity to the locality of K. qionglaiensis (Hou et al., 2021; Guo et al., 2022). Additionally, the phylogenetic placement of both as the sister taxon to K. idiootocus (Kuramoto and Wang, 1987) suggests that these two species likely refer to the same population. Moreover, although K. inexpectatus was phylogenetically identified as the closest relative to K. idiootocus, neither morphological nor molecular comparisons among K. inexpectatus, K. qionglaiensis, and K. silvaenaias were provided in the original publication of K. inexpectatus, even though the authors have acknowledged the description of these two congeners (Messenger et al., 2022).

During our survey in Qionglai City from 2022 to 2023, we collected a series of Kunxalus specimens from Pingle Town. Based on morphological and phylogenetic analyses of the newly collected specimens and type series of K. qionglaiensis and K. silvaenaias, we confirm that these individuals are conspecific. We also discuss the issue of nomenclature priority in the current era of electronic publication, emphasizing the importance of valid ZooBank registration. Additionally, based on the results presented in this study and its original description, we evaluate the taxonomic validity of K. inexpectatus.

2. Materials and Methods

2.1. Specimens and morphological measurements Nine newly collected Kunxalus specimens and the type series of K. qionglaiensis and K. silvaenaias were examined and measured. All specimens were deposited in the Herpetological Museum, Chengdu Institute of Biology (CIB), Chinese Academy of Sciences, China. External measurements were made using digital calipers (Neiko 01407A Stainless Steel 6-Inch Digital Caliper) to the nearest 0.1 mm. These measurements followed Jiang and Li (2021) and included snout-vent length (SVL); head length (HL); head width (HW); snout length (SL); internasal distance (INS); interorbital distance (IOS); horizontal diameter of eye (ED); horizontal diameter of tympanum (TD); hand length (HAL); forearm length (FOL); length of tarsus and foot (TFL); tibial length (TBL); finger III disc width (FDW3); and toe IV disc width (TDW4). The webbing formula followed Jiang and Li (2021).

2.2. Phylogenetic sampling and analyses Liver tissue from eight newly collected Kunxalus individuals was sampled for phylogenetic analysis. All samples obtained from euthanized specimens were preserved in 95% ethanol and stored at -20 °C. Genomic DNA was extracted using a DNA extraction kit (Sangon Biotech Co., Ltd., Shanghai, China). Three mitochondrial genes, including partial 12S ribosomal RNA gene (12S), partial 16S ribosomal RNA gene (16S) and partial cytochrome C oxidase 1 gene (COI), were amplified with primers following previous studies (Wilkinson et al., 2002; Lyu et al., 2023). Polymerase chain reaction (PCR) amplifications were processed under the following cycling conditions: denaturing at 95 °C for 4 min, 35 cycles of denaturing at 95 °C for 40 s, annealing at 53 °C (for 12S and 16S) / 48 °C (for COI) for 40 s, and extension at 72 °C for 60 s, and a final extension step at 72 °C for 10 min. The resulting PCR products were sequenced with both forward and reverse primers by Sangon Biotech Co., Ltd. (Shanghai, China). All sequences were deposited in GenBank (Table 1). For phylogenetic analyses, 37 sequences from additional Kunxalus congeners were obtained from GenBank (Table 1) and incorporated into our dataset. Three species from the Aquixalus clade were used as the out-groups.

DNA sequences were aligned using the Clustal W algorithm with default parameters (Thompson et al., 1997). Sequence data were analyzed using Bayesian inference (BI) in MrBayes v3.2.4 (Ronquist et al., 2012) and maximum-likelihood (ML) in RaxmlGUI vl.3 (Silvestro and Michalak, 2012). Two independent runs were conducted in BI analysis, each of which was performed for 20 000 000 generations and sampled every 1 000 generations with the first 25% of samples discarded as burn-in. For ML analysis, a bootstrap consensus tree inferred from 1 000 replicates was used to represent the evolutionary history of the taxa analyzed.

3. Results

The BI and ML analyses resulted in identical phylogenetic topologies (Figure 1). Results showed that 11 Kunxalus sensu stricto species formed a monophyletic group (Bayesian posterior probability (BPP) = 1.00, bootstrap support (BS) = 100), with three distinct clades further revealed within this group (BPP = 1.00, BS > 89, respectively). Clade A was monotypic and contained only K. gracilloides from Vietnam. Clade В was composed of four insular congeners from Taiwan Island and the Ryukyu Islands.

Clade C contained four lineages, with K. lenquanensis Yu, Wang, Hou, Rao and Yang, 2017 and K. raoi Zeng, Wang, Yu and Du, 2021 from southwestern China forming the two basal lineages (BPP > 0.99, BS > 99, respectively). All Kunxalus samples collected from Qionglai City, Sichuan, including those from the type series of K. qionglaiensis and K. silvaenaias, clustered together in a strongly supported lineage (BPP = 1.00, BS = 100) and almost without genetic divergence, indicating that these samples belonged to the same species. Morphological examination of these specimens from Qionglai confirmed that these individuals were conspecific. A supplementary description of this species is given below. The remaining lineage within clade C was composed of K. idiootocus from Taiwan Island and K. inexpectatus from eastern mainland of China, with small genetic divergence but relatively low support (BPP = 0.53, BS = 94). The taxonomic status of K. inexpectatus is also discussed below.

4. Taxonomic Account

Kurixalus qionglaiensis Guo, Zhong, Leung, Wang and Hu, 2022

Qionglai Frilled Tree Frog / Qióng Lái Yuan Zhï Shù Wā (Tß (ProQuest: ... denotes non-USASCII text omitted.) (Figure 2)

Kurixalus silvaenaias Hou, Peng, Miao, Liu, Li and Orlov, 2021, Animal Mol Breeding, 11: 7. Holotype: GIB 118049, by original designation. Unavailable nomenclatural act.

Kurixalus qionglaiensis Guo, Zhong, Leung, Wang and Hu, 2022, Zool Res, 43: 92. Holotype: GIB 118031, by original designation.

Specimens examined Twenty-four adult males.

Holotype GIB 118031 (field number PL2021060522), adult male, collected on 5 June 2021 from Lugou Bamboo Sea (30° 2Г51" N, 103° 18'30" E; elevation 615ma.sl), Pingle Town, Qionglai City, Chengdu City, Sichuan Province, China.

Paratypes CIB 118032-118037 (field number PL2021060 503-06, 14-15), six adult males, collected 5 June 2021 from the same location as the holotype.

Other materials CIB 118049-118056 (field number HM 202004001-008) (type series of K. silvaenaias), eight adult males, collected in April 2020 from Pingle Town (30°36'16" N, 103°29'69" E, 623 m elevation); CIB 119692-119697 (field number LJT-LAB2022591, 617-621), six adult males, collected on 28 June 2022 from Pingle Town (30.3696° N, 103.3055° E; 700 m a.sl); CIB 119698-119700 (field number LJT-LAB2023219-220, 223), three adult males, collected on 8 July 2023 from Lugou Bamboo Sea (30.3599° N, 103.2997° E; 740 m a.sl).

Diagnosis Medium body size, SVL 28.6-33.4 mm in adult males (n = 24); vomerine teeth distinct; dorsal surface overall rough with numerous irregular tubercles and granules; flanks relatively smooth with scattered warts; dorsal surfaces commonly dark brown, light brown or greenish brown, with large darker saddle-shaped patch on central back; irregular black stripes or patches on chin; pair of dark patches at armpits; single vocal sac present in males; single weak nuptial pad present on base of finger I in breeding males.

Description Body slender and slightly flattened. Head moderate and depressed, HW/HL 0.86-1.05 (n = 24); snout short, distinctly protruding in profile, projecting beyond lower jaw; canthus rostralis prominent and blunt, loreal region slightly concave; nostril oval, slightly prominent, closer to tip of snout than eye; internasal distance smaller than interorbital distance; top of head flat, pineal ocellus distinct or indistinct; eye large, convex; pupil oval, horizontal; tympanum small but distinct, rounded; supratympanic fold distinct, curving from posterior edge of eye to insertion of arm; vomerine teeth distinct; tongue large, margin distinctly notched.

Forelimbs slender; relative finger lengths I < II ~ IV < III; tips of all fingers dilated into oval and flattened discs with circummarginal grooves; fingers with rudimentary webs; outer margin of finger IV with distinct serrated fringes; subarticular tubercles prominent, rounded, formula 1, 1, 2, 1; supernumerary tubercle below base of finger IV distinct and prominent, those below other fingers indistinct in preserved specimens; inner metacarpal tubercle prominent and oval, two outer metacarpal tubercles indistinct in preserved specimens; series of large tubercles scattered along outer edge of hand and forearm.

Hindlimbs moderately robust, long; tibiotarsal articulation reaching between anterior and posterior corners of eye when hindlimb adpressed alongside body; heels just meeting when hindlimbs flexed at right angles to body axis; relative toe lengths I < II < III ~ V < IV; tips of all toes expanded into welldeveloped oval and flattened discs with distinct circummarginal grooves; toes half webbed, webbing formula I 1-1 II У2-2 III ^-2 IV 1 Уз-Уз V; distinct lateral fringes present on inner and outer margins of all toes; subarticular tubercles prominent, rounded, formula 1, 1, 2, 3, 2; inner metatarsal tubercle distinct and oval, outer metatarsal tubercle absent; series of large tubercles scattered along outer edge of tarsus forming serrated fringes.

Dorsal skin overall rough with numerous irregular tubercles and granules; dorsolateral folds absent; flanks relatively smooth with scattered warts. Ventral skin wrinkled with flattened tubercles; large warts present on posterior of limbs and near cloaca.

Coloration Coloration in life varies among individuals. Dorsal surfaces commonly dark brown, light brown, or greenish brown; large darker saddle-shaped patch on central back, extending from upper eyelids to sacral area, but varying distinctly in shape and shade among individuals; light band on lip below eye; limbs with darker patches and indistinct bands. Ventral surfaces commonly creamy white; irregular black stripes or patches on chin; pair of dark patches at armpits; belly dark. All specimens faded in preservative, dorsal surfaces grayish or brownish; ventral surfaces yellowish or reddish; dorsal and ventral marking clearer.

Variations and sexual dimorphism Morphometric differences among specimens are presented in Table 2. Several measurements of the same individual varied from those reported by Guo et al. (2022) and Hou et al. (2021), attributed to differences in measurement methodologies. Hou et al. (2021) reported pronounced sexual dimorphism in this species, noting that the SVL of females is twice as long as that of males. However, no female specimens were captured for accurate measurements. According to Hou et al. (2021), females also display a more prominent snout tip than males. The specimens examined in this study were all breeding males, characterized by a single grayish white nuptial pad on the base of finger I and the presence of a single vocal sac.

Distribution and habitat Currently, Kunxalus qionglaiensis is known only from a single locality in Pingle Town within Qionglai City, located at the western edge of the Sichuan Basin. This treefrog inhabits bamboo and scrub forests between 600-740 m a.s.l, where it is observed sympatrically with Zhangixalus omeimontis (Stejneger, 1924). Guo et al. (2022) initially discovered this species in early June, while Hou et al. (2021) documented its breeding season as spanning from April to May. During our survey, conducted from late June to early July, we observed active singing males, but did not encounter any mating pairs.

Conservation recommendation Combining the survey data from 2020 to 2023 reported by Hou et al. (2021), Guo et al. (2022), and our study, we suggest that Kunxalus qionglaiensis is eligible for listing as Endangered (Blab(iii, iv) + 2ab(iii, iv)) in the IUCN Red List of Threatened Species.

5. Discussion

5.1. Naming of the Kurixalus population in Qionglai The principle of publication priority is fundamental in zoological nomenclature. In the era of electronic publication, determining the publication date of a nomenclature can be challenging, particularly for works issued in both electronic and print formats, which may present two different dates. The International Commission on Zoological Nomenclature (ICZN) has introduced a series of amendments to the International Code of Zoological Nomenclature (the Code; available on https://www.iczn.org/the-code/the-codeonline/) to clarify the electronic publication of new scientific names and nomenclatural acts (ICZN, 2012; Kreil and Pape, 2015). As per Article 21.9 of the Code, the publication date for a nomenclature issued in both print and electronic formats is determined by the edition that first fulfils the criteria set out in Article 8 and is not excluded by Article 9. Article 8 stipulates that currently an electronic publication satisfies the publication criteria if it includes statements of the publication date and the registration information in the Official Register of Zoological Nomenclature (ZooBank).

In the matter of the conspecificity between Kurixalus qionglaiensis and K. silvaenaias, establishing correct publication dates of these two species is crucial. Although the paper version of K. qionglaiensis was printed on 18 January 2022, it became available online on 10 December 2021, complete with the statements of electronic publication date and ZooBank registration (Guo et al., 2022), seemingly meeting Article 8 criteria. However, under Article 8.5.3, a valid ZooBank entry must include the name of an electronic archive intended to preserve the work, along with the associated ISSN or ISBN. Although the journal in which K. qionglaiensis was published, Zoological Research, is archived in Zenodo and PubMed Central (Figure ЗА), the ZooBank registration lacks the requisite archive statement for K. qionglaiensis (Figure 3C), rendering this registration invalid. Consequently, the correct publication date of K. qionglaiensis should be the date when it was physically printed (18 January 2022). Regarding K. silvaenaias, it was purportedly published online on 27 December 2021 (Hou et al, 2021). However, its ZooBank registration also fails to mention an electronic archive (Figure 3D). The publishing journal, Animal Molecular Breeding, exists only in an online format without a printed version (Figure 3B; available on https://animalscipublisher.com/index.php/amb). Therefore, the publication of K. silvaenaias by Hou et al. (2021) does not constitute a published work under Article 8 and should be regarded as an unavailable nomenclatural act according to Article 11. Hence, Kurixalus qionglaiensis Guo, Zhong, Leung, Wang and Hu, 2022 is the valid nomenclature for the frilled treefrog population in Pingle Town, Qionglai City.

A comparable debate regarding publication priority occurred between Gracixalus nonggangensis Mo, Zhang, Luo, Zhou, and Chen, 2013 and G. waza Nguyen, Le, Pham, Nguyen, Bonkowski, and Ziegler, 2013. This debate also centered on the challenge of determining the publication date in scenarios where both electronic and print editions are available, with the legitimacy of the ZooBank registration being a critical factor (Matsui et al., 2015; Wang et al., 2018; Nguyen et al., 2020). We also noticed numerous instances of invalid ZooBank registrations, suggesting such errors are not uncommon. One contributing factor may be that the requirement for an online archive, which is mandatory according to the Code, is not a required field in the ZooBank registration process (Figure 3E), leading to its frequent omission, especially among those unfamiliar with the stipulations set on in the Code. An example of valid ZooBank registration is shown in Figure 3F (Wang et al., 2022).

Particularly, an exception exists under Article 9.9 and Article 21.8.3 for works that are available online in preliminary versions prior to the official publication of their final version. Therefore, if a work is labeled as "just accepted", "pre-proof", or other similar status by a journal, it is typically regarded as a preliminary version, as it is not yet finalized and may undergo further changes. The Article 8.5 criteria become applicable only when a work is categorized as "in press", serving as "the version of record" following final proof corrections and accompanied by a clear statement of its online publication date (Kreil and Pape, 2015). We strongly recommend authors, editors, and publishers working with taxonomic publications should carefully read and understand detailed Articles of the Code on electronic publications and related in-depth explanations (ICZN, 2012; Kreil and Pape, 2015), to avoid causing a series of controversial nomenclatures (Doan et al., 2023; Dubois and Frétey, 2023) or taking the possibility of immoral or unexpected scooping actions. Furthermore, we suggest ZooBank to make all mandatory items for electronic publications defined by the Code as required fields when user registering (Figure 3E), to prevent further occurrences of the case discussed in this work.

5.2. Taxonomic status of Kurixalus inexpectatus In this study, we conducted a phylogenetic analysis including both Kurixalus inexpectatus and K. qionglaiensis, which was absent in the original description of K. inexpectatus (Messenger et al., 2022). Nevertheless, the results indicated that the classification of K. inexpectatus as a distinct species is not supported and it should be reclassified as a junior synonym of K. idiootocus. In the original study, phylogenetic analysis based on 12S-TYR gene fragments placed K. inexpectatus within the lineage of K. idiootocus (BPP = 0.97), resulting in a paraphyletic status for K. idiootocus following the erecting of K. inexpectatus (Messenger et al., 2022). Our phylogenetic analysis based on 12S-16S-COI gene fragments also suggested a distinct lineage of K. idiootocus, including all K. inexpectatus samples (Figure 1). The lineage received robust BS support (94), but relatively weak BPP support (0.53), primarily due to the limited molecular data available for K. inexpectatus (Table 1). Morphologically, no superficial anatomical characteristics distinguished these two species. The only reported differences were minor and included the location of the tibiotarsal articulation when extended (beyond the anterior corner of the eye in K. inexpectatus vs. at the center of the eye in K. idiootocus) and several morphometric ratios. However, the position of the tibiotarsal articulation when extended usually varies among individuals (e.g. between the anterior and posterior corners of the eye in K. qionglaiensis, as described above), and differences in morphometric ratios are typical among geographical populations. Although the original study analyzed acoustic differences between the two species, the methodology employed was overly simplistic. Extensive research has shown that the vocalizations of the genus Kurixalus are complex, with multiple call types that vary under different environmental conditions (Tan et al., 2014; Zhu et al., 2017; Yi and Sheridan, 2019; Zhang et al., 2021). This complexity underscores the need for more detailed and comprehensive analyses of acoustic data within this genus. Hence, the data currently available fails to distinctly differentiate K. inexpectatus from K. idiootocus, and thus they should be considered conspecific in spite of their isolated geographic distribution.

Acknowledgements We thank Bicheng ZHU, Junjie HUANG, Dihao WU, Jingyi XU, and Ziehen QIAO for their help in field and laboratory work. We thank two anonymous reviewers for their constructive comments on the manuscript. This work was supported by the Biological Resources Programme, Chinese Academy of Sciences (KFJ-BRP-017-086, KFJ-BRP-017-65), Youth Innovation Promotion Association of CAS (2021370), National Natural Science Foundation of China (32200363, 32370498), and Chengdu Municipal Park City Construction and Management Bureau.

Sidebar

* Corresponding authors: Mr. Ke JIANG, from Chengdu Institute of Biology (CIB), Chinese Academy of Sciences (CAS), Chengdu, Sichuan, China, with his research focusing on amphibian and reptile taxonomy; Dr. Dechun JIANG, from GIB, CAS, Chengdu, Sichuan, China, with her research focusing on treefrog taxonomy and biogeography.

E-mail: [email protected] (Ke JIANG); [email protected] (Dechun JIANG)

Received: 7 December 2023 Accepted: 17 January 2024

Published Online: 31 January 2024

2095-0357/© 2024 Asian Herpetological Research Editorial Office and Science Press. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

How to cite this article:

Lyu Z. T., Li M. L., Wang G. Q., Liu G., Liu M., Jiang K., Jiang D. C., Li. J. T. 2024. Taxonomic status of three recently proposed species of the genus Kunxalus (Anura, Rhacophoridae), with discussion on the ZooBank registrations for electronical publication. Asian Herpetol Res, 15(1): 31-40. DOI: 10.3724/ahr.2095-0357.2023.0049

References

References

Delorme M., Dubois A., Grosjean S., Ohler A. 2005. Une nouvelle classification générique et subgénérique de la tribu des Philautini (Amphibia, Anura, Rhacophorinae). Bull Mens Soc Linn Lyon, 74: 165-171

Doan T. M., Rowe J. W., Fitzgerald L. A. 2023. Journal of Herpetology is compliant with the international code of zoological nomenclature. J Herpetol, 57: 447-447

Dubois A., Frétey T. 2023. Problems with the availability of zoological nomenclatural acts published online: The case of the Journal of Herpetology. Bionomina, 34: 9-34

Fei L. 1999. Atlas of Amphibians of China. Zhengzhou, China: Henan Press of Science and Technology

Guo C. P., Zhong M. J., Wah Leung K., Wang X. Y, Hu J. H. 2022. A new species of the genus Kurixalus (Anura, Rhacophoridae) from Sichuan Province, southwestern China. Zool Res, 43: 90-94

Hou M., Peng X., Miao J., Liu S., Li P., Orlov N. L. 2021. Description a new species of genus Kurixalus (Amphibia: Anura: Rhacophoridae) from Chengdu Prefecture, Sichuan Province, China. Anim Mol Breed, 11: 1-16

ICZN. 2012. Amendment of Articles 8, 9, 10, 21 and 78 of the International Code of Zoological Nomenclature to expand and refine methods of publication. Zootaxa, 3450: 1-7

Jiang K., Li J. T. 2021. Method for External Measurement of Adult Anuran Specimens. Bio-101, e 1010675. https://doi.org/10.21769/ BioProtoc. 1010675

Kreil F. T., Pape T. 2015. Electronic publications need registration in ZooBank to be available. Bull Zool Nomenci, 72: 245-251

Kuramoto M., Wang C. S. 1987. A new rhacophorid treefrog from Taiwan, with comparisons to Chirixalus eiffingeri (Anura, Rhacophoridae). Copeia, 1987: 931-942

Li J., Che J., Bain R H., Zhao E., Zhang Y. 2008. Molecular phylogeny of Rhacophoridae (Anura): A framework of taxonomic reassignment of species within the genera Aquixalus, Chiromantis, Rhacophorus, and Philautus. Mol Phylogenet Evol, 48: 302-312

Lyu Z. T., Qi S., Wang J., Zhang S. Y, Zhao J., Zeng Z. G, Wan H., Yang J. H., Mo Y. M., Wang Y. Y. 2023. Generic classification of Asian horned toads (Anura: Megophryidae: Megophryinae) and monograph of Chinese species. Zool Res, 44: 380-450

Matsui M., Khonsue W., Panha S., Eto K. 2015. A new tree frog of the genus Gracixalus from Thailand (Amphibia: Rhacophoridae). Zool Sei, 32: 204-210

Messenger K. R, Othman S. N., Chuang M. F., Yang Y, Börzéé A. 2022. Description of a new Kurixalus species (Rhacophoridae, Anura) and a northwards range extension of the genus. ZooKeys, 1108: 15-49

Nguyen T. V., Duong T. V., Luu K. T., Poyarkov N. A. 2020. A new species of Kurixalus (Anura: Rhacophoridae) from northern Vietnam with comments on the biogeography of the genus. J Nat Hist, 54: 195-223

Ronquist F., Teslenko M., van der Mark P., Ayres D. L, Darling A., Höhna S., Larget B., Liu L, Suchard M. A., Huelsenbeck J. P. 2012. MrBayes 3.2: Efficient bayesian phylogenetic inference and model choice across a large model space. Syst Biol, 61: 539-542

Silvestro D., Michalak I. 2012. raxmlGUI: A graphical front-end for RAxML. Org Divers Evol, 12: 335-337

Stejneger L. 1924. Herpetological novelties from China. Occas Pap Boston Soc Nat Hist, 5: 119-121

Tan W. H., Tsai C. G., Lin G, Lin Y. K. 2014. Urban canyon effect: Storm drains enhance call characteristics of the Mientien tree frog. J Zool, 294: 77-84

Thompson}. D., Gibson T. J., Plewniak F., Jeanmougin E, Higgins D. G. 1997. The CLUSTAL_X windows interface: Flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Res, 25: 4876-4882

Wang J., Zeng Z. G., Lyu Z. T., Liu Z. Y, Wang Y Y. 2018. Description of a new species of Gracixalus (Amphibia: Anura: Rhacophoridae) from Guangdong Province, southeastern China. Zootaxa, 4420: 251-269

Wang J., Zeng Z. C., Lyu Z. T., Qi S., Liu Z. Y, Chen H. H., Lu Y H., Xiao H. W., Lin C. R, Chen K., Wang Y. Y. 2022. Description of three new Boulenophrys species from eastern Guangdong, China, emphasizing the urgency of ecological conservation in this region (Anura, Megophryidae). Zootaxa, 5099: 91-119

Wilkinson J. A., Drewes R G, Tatum O. L. 2002. A molecular phylogenetic analysis of the family Rhacophoridae with an emphasis on the Asian and African genera. Mol Phylogenet Evol, 24: 265-273

Yi Y. Z., Sheridan J. A. 2019. Effects of traffic noise on vocalisations of the rhacophorid tree frog Kunxalus chaseni (Anura: Rhacophoridae) in Borneo. Raffles Bull Zool, 67: 77-82

Yu G., Wang J., Hou M., Rao D., Yang J. 2017. A new species of the genus Kunxalus from Yunnan, China (Anura, Rhacophoridae). ZooKeys, 694: 71-93

Zengj., Wang J. S., Yu G. H., Du L. N. 2021. A new species of Kunxalus (Anura, Rhacophoridae) from Guizhou, China. Zool Res, 42: 227-233

Zhang H., Zhu B., Zhou Y., He Q., Sun X., Wang J., Cui J. 2021. Females and males respond differently to calls impaired by noise in a tree frog. Ecol Evol, 11: 9159-9167

Zhu B., Wang J., Sun Z., Yang Y., Wang T., Brauth S. E., Tang Y., Cui J. 2017. Competitive pressures affect sexual signal complexity in Kunxalus odontotarsus: Insights into the evolution of compound calls. Biol Open, 6: 1913-1918

Scientific Editor: Bin WANG