Vol. 20 No. 1 (2025)
Articles

Genetically informed range extension for Kurixalus inexpectatus (Anura: Rhacophoridae) in Fujian, with ecological niche modeling to guide further searches

Dallin B. Kohler
Laboratory of Animal Behaviour and Conservation, College of Ecology and the Environment, Nanjing Forestry University, Nanjing 210037, China
Siti N. Othman
College of Life Sciences, Nanjing Forestry University, Nanjing 210037, China
Zhenqi Wang
Yunxiao County Forestry Bureau, Zhangzhou 363300, China
Yiyao Zhu
Fujian Forestry Prospect and Design Institute, Fuzhou 350001, China
Yiyun Wu
Department of Life Sciences, National Chung Hsing University, Taichung, 402202, Taiwan
Ming-Feng Chuang
Global Change Biology Research Center, National Chung Hsing University, Taichung, 402202, Taiwan
Amaël Borzée
College of Life Sciences, Nanjing Forestry University, Nanjing 210037, China

Published 2025-05-12

Keywords

  • Maxent,
  • Cryptic species,
  • tree frog,
  • Kurixalus

How to Cite

Kohler, D. B., Othman, S. N., Wang, Z., Zhu, Y., Wu, Y., Chuang, M.-F., & Borzée, A. (2025). Genetically informed range extension for Kurixalus inexpectatus (Anura: Rhacophoridae) in Fujian, with ecological niche modeling to guide further searches. Acta Herpetologica, 20(1), 3–16. https://doi.org/10.36253/a_h-17227

Abstract

Undocumented and unnamed biodiversity is difficult to conserve effectively. We encountered several Kurixalus (Anura: Rhacophoridae) individuals of an unknown species identity in Fujian, China, a province with no previous records of Kurixalus. This genus of frogs, of which many new species have been described in recent years, has been the subject of much historical taxonomic confusion, largely due to conserved morphology. We sequenced two mitochondrial and one nuclear genes (1748 total bp) for three individuals from Fujian and reconstructed their phylogeny including all known lineages of Kurixalus from mainland China, and other East Asia lineages for which data were available. Both Bayesian Inference and Maximum Likelihood trees consistently showed that the individuals from Fujian were most closely related to K. inexpectatus, which previously was only known from northern Zhejiang, over 830 km north of this new location. Our results help clarify the identity of the unknown Kurixalus in Fujian and the distribution of Kurixalus, particularly regarding the distribution of K. hainanus, which has previously been confused with other members of the K. odontotarsus species complex. Additionally, we created ecological niche models using Maxent for the clade including both K. inexpectatus and its sister species K. idiootocus to guide future survey efforts.

References

  1. Ali, J.R. (2020): Geological data indicate that the interpretation for the age-calibrated phylogeny for the Kurixalus-genus frogs of South, South-east and East Asia (Lv et al., 2018) needs to be rethought. Mol. Phylogenet. Evol. 145: 106053. DOI: https://doi.org/10.1016/j.ympev.2018.02.011
  2. Ananjeva, N.B., Golynsky, E.E., Lin, S.-M., Orlov, N.L., Tseng, H.-Y. (2015): Modeling habitat suitability to predict the potential distribution of the Kelung cat snake Boiga kraepelini Steineger, 1902. Russ. J. Herpetol. 22: 197-205.
  3. Blair, C., Bryson, R.W. (2017): Cryptic diversity and discordance in single‐locus species delimitation methods within horned lizards (Phrynosomatidae: Phrynosoma). Mol. Ecol. Resour. 17: 1168-1182. DOI: https://doi.org/10.1111/1755-0998.12658
  4. Bossuyt, F., Milinkovitch, M.C. (2000): Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proc. Natl. Acad. Sci. 97: 6585-6590. DOI: https://doi.org/10.1073/pnas.97.12.6585
  5. Breiner, F.T., Guisan, A., Bergamini, A., Nobis, M.P. (2015): Overcoming limitations of modelling rare species by using ensembles of small models. Methods Ecol. Evol. 6: 1210-1218. DOI: https://doi.org/10.1111/2041-210X.12403
  6. Chan, K.O., Hertwig, S.T., Neokleous, D.N., Flury, J.M., Brown, R.M. (2022): Widely used, short 16S rRNA mitochondrial gene fragments yield poor and erratic results in phylogenetic estimation and species delimitation of amphibians. BMC Ecol. Evol. 22: 37. DOI: https://doi.org/10.1186/s12862-022-01994-y
  7. Che, J., Chen, H., Yang, J., Jin, J., Jiang, K., Yuan, Z., Murphy, R.W., Zhang, Y. (2012): Universal COI primers for DNA barcoding amphibians. Mol. Ecol. Resour. 12: 247-258. DOI: https://doi.org/10.1111/j.1755-0998.2011.03090.x
  8. Chernomor, O., Von Haeseler, A., Minh, B.Q. (2016): Terrace aware data structure for phylogenomic inference from supermatrices. Syst. Biol. 65: 997-1008. DOI: https://doi.org/10.1093/sysbio/syw037
  9. Deng, K., He, Y.-X., Wang, X.-P., Wang, T.-L., Wang, J.-C., Chen, Y.-H., Cui, J.-G. (2024): Hainan frilled treefrogs’ calls partially conform to Menzerath–Altmann’s law, but oppose Zipf’s law of abbreviation. Anim. Behav. 213: 51-59. DOI: https://doi.org/10.1016/j.anbehav.2024.04.011
  10. Dufresnes, C., Litvinchuk, S.N. (2022): Diversity, distribution and molecular species delimitation in frogs and toads from the Eastern Palaearctic. Zool. J. Linn. Soc. 195: 695-760. DOI: https://doi.org/10.1093/zoolinnean/zlab083
  11. Edgar, R.C. (2004): MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Res. 32: 1792-1797. DOI: https://doi.org/10.1093/nar/gkh340
  12. Elith, J., Phillips, S.J., Hastie, T., Dudík, M., Chee, Y.E., Yates, C.J. (2011): A statistical explanation of MaxEnt for ecologists: Statistical explanation of MaxEnt. Divers. Distrib. 17: 43-57. DOI: https://doi.org/10.1111/j.1472-4642.2010.00725.x
  13. Entiauspe‐Neto, O.M., Dervanoski, D., Abegg, A.D. (2024): Can fieldwork driven by predictive species distribution models yield new rare or relevant geographic records? A case study with Neotropical snakes. Austral Ecol. 49: e70013. DOI: https://doi.org/10.1111/aec.70013
  14. Farr, T.G., Rosen, P.A., Caro, E., Crippen, R., Duren, R., Hensley, S., Kobrick, M., Paller, M., Rodriguez, E., Roth, L., Seal, D., Shaffer, S., Shimada, J., Umland, J., Werner, M., Oskin, M., Burbank, D., Alsdorf, D. (2007): The shuttle radar topography mission. Rev. Geophys. 45: 2005RG000183. DOI: https://doi.org/10.1029/2005RG000183
  15. Feng, X., Park, D.S., Liang, Y., Pandey, R., Papeş, M. (2019): Collinearity in ecological niche modeling: Confusions and challenges. Ecol. Evol. 9: 10365-10376. DOI: https://doi.org/10.1002/ece3.5555
  16. Fick, S.E., Hijmans, R.J. (2017): WorldClim 2: new 1‐km spatial resolution climate surfaces for global land areas. Int. J. Climatol. 37: 4302-4315. DOI: https://doi.org/10.1002/joc.5086
  17. Frost, D.R. (2024a): Amphibian species of the world 6.2, an online reference. American Museum of Natural History, New York. Available online at: https://amphibiansoftheworld.amnh.org/. [Accessed 13 September 2024].
  18. Frost, D.R. (2024b): Kurixalus odontotarsus. American Museum of Natural History, New York. Available online at: https://amphibiansoftheworld.amnh.org/Amphibia/Anura/Rhacophoridae/Rhacophorinae/Kurixalus/Kurixalus-odontotarsus. [Accessed 19 November 2024].
  19. Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., De Sá, R.O., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M., Wheeler, W.C. (2006): The amphibian tree of life. Bull. Am. Mus. Nat. Hist. 297: 1-291. DOI: https://doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2
  20. GBIF.org (2024a): Occurrence Download: Anura. The Global Biodiversity Information Facility. https://doi.org/10.15468/DL.CXHN8Z.
  21. GBIF.org (2024b): Occurrence Download: Kurixalus idiootocus. The Global Biodiversity Information Facility. https://doi.org/10.15468/DL.XPSDR7.
  22. Guindon, S., Dufayard, J.-F., Lefort, V., Anisimova, M., Hordijk, W., Gascuel, O. (2010): New algorithms and methods to estimate Maximum-Likelihood Phylogenies: Assessing the Performance of PhyML 3.0. Syst. Biol. 59: 307-321. DOI: https://doi.org/10.1093/sysbio/syq010
  23. 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. DOI: https://doi.org/10.24272/j.issn.2095-8137.2021.293
  24. Hijmans, R.J. (2024): raster: Geographic data analysis and modeling.
  25. Hijmans, R.J., Barbosa, M., Ghosh, A., Mandel, A. (2024): ‘geodata’: Download geographic data.
  26. Hoang, D.T., Chernomor, O., Von Haeseler, A., Minh, B.Q., Vinh, L.S. (2018): UFBoot2: Improving the ultrafast bootstrap approximation. Mol. Biol. Evol. 35: 518-522. DOI: https://doi.org/10.1093/molbev/msx281
  27. Hofmann, E.P., Nicholson, K.E., Luque-Montes, I.R., Köhler, G., Cerrato-Mendoza, C.A., Medina-Flores, M., Wilson, L.D., Townsend, J.H. (2019): Cryptic diversity, but to what extent? Discordance between single-locus species delimitation methods within mainland anoles (Squamata: Dactyloidae) of Northern Central America. Front. Genet. 10: 11. DOI: https://doi.org/10.3389/fgene.2019.00011
  28. 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. DOI: https://doi.org/10.5376/amb.2021.11.0002
  29. Hou, J., Xiang, J., Li, D., Liu, X. (2023): Prediction of potential suitable distribution areas of Quasipaa spinosa in China based on MaxEnt optimization model. Biology 12: 366. DOI: https://doi.org/10.3390/biology12030366
  30. IUCN Standards and Petitions Committee (2024): Guidelines for using the IUCN Red List categories and criteria. Gland, Switzerland.
  31. Kass, J.M., Muscarella, R., Galante, P.J., Bohl, C.L., Pinilla‐Buitrago, G.E., Boria, R.A., Soley‐Guardia, M., Anderson, R.P. (2021): ENMeval 2.0: Redesigned for customizable and reproducible modeling of species’ niches and distributions. Methods Ecol. Evol. 12: 1602-1608. DOI: https://doi.org/10.1111/2041-210X.13628
  32. Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P., Drummond, A. (2012): Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28: 1647-1649. DOI: https://doi.org/10.1093/bioinformatics/bts199
  33. Kidov, A.A., Litvinchuk, S.N. (2021): Distribution and conservation status of the Hyrcanian Wood Frog (Rana pseudodalmatina) in Azerbaijan. Russ. J. Herpetol. 28: 97-107. DOI: https://doi.org/10.30906/1026-2296-2021-28-2-97-107
  34. Kohler, D.B., Zhang, X., Messenger, K.R., Chin Yu An, K., Ghosh, D., Othman, S.N., Wang, Z., Amin, H., Prasad, V.K., Wu, Z., Borzée, A. (2024): At home in Jiangsu: Environmental niche modeling and new records for five species of amphibian and reptile in Jiangsu, China. Herpetozoa 37: 85-93. DOI: https://doi.org/10.3897/herpetozoa.37.e117370
  35. Kramer-Schadt, S., Niedballa, J., Pilgrim, J.D., Schröder, B., Lindenborn, J., Reinfelder, V., Stillfried, M., Heckmann, I., Scharf, A.K., Augeri, D.M., Cheyne, S.M., Hearn, A.J., Ross, J., Macdonald, D.W., Mathai, J., Eaton, J., Marshall, A.J., Semiadi, G., Rustam, R., Bernard, H., Alfred, R., Samejima, H., Duckworth, J.W., Breitenmoser-Wuersten, C., Belant, J.L., Hofer, H., Wilting, A. (2013): The importance of correcting for sampling bias in MaxEnt species distribution models. Divers. Distrib. 19: 1366-1379. DOI: https://doi.org/10.1111/ddi.12096
  36. Lanfear, R., Calcott, B., Ho, S.Y.W., Guindon, S. (2012): PartitionFinder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Mol. Biol. Evol. 29: 1695-1701. DOI: https://doi.org/10.1093/molbev/mss020
  37. Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T., Calcott, B. (2017): PartitionFinder 2: New methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Mol. Biol. Evol. msw260. DOI: https://doi.org/10.1093/molbev/msw260
  38. Li, J. (2023): iNaturalist observation 186300362. iNaturalist.org. Available online at: https://www.inaturalist.org/observations/186300362.
  39. 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. DOI: https://doi.org/10.1016/j.ympev.2008.03.023
  40. Li, J.-T., Li, Y., Klaus, S., Rao, D.-Q., Hillis, D.M., Zhang, Y.-P. (2013): Diversification of rhacophorid frogs provides evidence for accelerated faunal exchange between India and Eurasia during the Oligocene. Proc. Natl. Acad. Sci. 110: 3441-3446. DOI: https://doi.org/10.1073/pnas.1300881110
  41. Li, S., Dai, W., Wang, Z., Wu, Z., Wang, J. (2024): Detecting range shrinking from historical amphibian species occurrences under influence of human impacts: A case study using the Chinese giant salamander, Andrias davidianus. Ecol. Evol. 14: e70595. DOI: https://doi.org/10.1002/ece3.70595
  42. Lin, Y.-S., Kam, Y.-C. (2008): Nest choice and breeding phenology of an arboreal-breeding frog, Kurixalus eifingeri (Rhacophoridae), in a bamboo forest. Zool. Stud. 47: 129-137.
  43. Liu, C., Newell, G., White, M., Machunter, J. (2025): Improving the estimation of the Boyce index using statistical smoothing methods for evaluating species distribution models with presence‐only data. Ecography 2025: e07218. DOI: https://doi.org/10.1111/ecog.07218
  44. Luedtke, J.A., Chanson, J., Neam, K., Hobin, L., Maciel, A.O., Catenazzi, A., Borzée, A., Hamidy, A., Aowphol, A., Jean, A., Sosa-Bartuano, Á., Fong G., A., De Silva, A., Fouquet, A., Angulo, A., Kidov, A.A., Muñoz Saravia, A., Diesmos, A.C., Tominaga, A., Shrestha, B., Gratwicke, B., Tjaturadi, B., Martínez Rivera, C.C., Vásquez Almazán, C.R., Señaris, C., Chandramouli, S.R., Strüssmann, C., Cortez Fernández, C.F., Azat, C., Hoskin, C.J., Hilton-Taylor, C., Whyte, D.L., Gower, D.J., Olson, D.H., Cisneros-Heredia, D.F., Santana, D.J., Nagombi, E., Najafi-Majd, E., Quah, E.S.H., Bolaños, F., Xie, F., Brusquetti, F., Álvarez, F.S., Andreone, F., Glaw, F., Castañeda, F.E., Kraus, F., Parra-Olea, G., Chaves, G., Medina-Rangel, G.F., González-Durán, G., Ortega-Andrade, H.M., Machado, I.F., Das, I., Dias, I.R., Urbina-Cardona, J.N., Crnobrnja-Isailović, J., Yang, J.-H., Jianping, J., Wangyal, J.T., Rowley, J.J.L., Measey, J., Vasudevan, K., Chan, K.O., Gururaja, K.V., Ovaska, K., Warr, L.C., Canseco-Márquez, L., Toledo, L.F., Díaz, L.M., Khan, M.M.H., Meegaskumbura, M., Acevedo, M.E., Napoli, M.F., Ponce, M.A., Vaira, M., Lampo, M., Yánez-Muñoz, M.H., Scherz, M.D., Rödel, M.-O., Matsui, M., Fildor, M., Kusrini, M.D., Ahmed, M.F., Rais, M., Kouamé, N.G., García, N., Gonwouo, N.L., Burrowes, P.A., Imbun, P.Y., Wagner, P., Kok, P.J.R., Joglar, R.L., Auguste, R.J., Brandão, R.A., Ibáñez, R., Von May, R., Hedges, S.B., Biju, S.D., Ganesh, S.R., Wren, S., Das, S., Flechas, S.V., Ashpole, S.L., Robleto-Hernández, S.J., Loader, S.P., Incháustegui, S.J., Garg, S., Phimmachak, S., Richards, S.J., Slimani, T., Osborne-Naikatini, T., Abreu-Jardim, T.P.F., Condez, T.H., De Carvalho, T.R., Cutajar, T.P., Pierson, T.W., Nguyen, T.Q., Kaya, U., Yuan, Z., Long, B., Langhammer, P., Stuart, S.N. (2023): Ongoing declines for the world’s amphibians in the face of emerging threats. Nature 622: 308-314. DOI: https://doi.org/10.1038/s41586-023-06578-4
  45. Luo, T., Zhao, X., Lan, C., Li, W., Deng, H., Xiao, N., Zhou, J. (2023): Integrated phylogenetic analyses reveal the evolutionary, biogeographic, and diversification history of Asian warty treefrog genus Theloderma (Anura, Rhacophoridae). Ecol. Evol. 13: e10829. DOI: https://doi.org/10.1002/ece3.10829
  46. Lv, Y.-Y., He, K., Klaus, S., Brown, R.M., Li, J.-T. (2018): A comprehensive phylogeny of the genus Kurixalus (Rhacophoridae, Anura) sheds light on the geographical range evolution of frilled swamp treefrogs. Mol. Phylogenet. Evol. 121: 224-232. DOI: https://doi.org/10.1016/j.ympev.2017.09.019
  47. Lyu, Z., Li, M., Wang, G., Liu, G., Liu, M., Jiang, K., Jiang, D., Li, J. (2024): Taxonomic status of three recently proposed species of the genus Kurixalus (Anura, Rhacophoridae), with discussion on the ZooBank registrations for electronical publication. Asian Herpetol. Res. 15: 31-40. DOI: https://doi.org/10.3724/ahr.2095-0357.2023.0049
  48. Madhushanka, S., Manamendra-Arachchi, K. (2021): Morphometry and habitat selection of the mountain hourglass treefrog [Taruga eques (Günther, 1858)] (Amphibia, Rhacophoridae) in the Horton Plains National Park, Sri Lanka. Univ. Colombo Rev. 3: 64. DOI: https://doi.org/10.4038/ucr.v3i2.73
  49. Messenger, K.R., Othman, S.N., Chuang, M.-F., Yang, Y., Borzée, A. (2022): Description of a new Kurixalus species (Rhacophoridae, Anura) and a northwards range extension of the genus. ZooKeys 1108: 15-49. DOI: https://doi.org/10.3897/zookeys.1108.81725
  50. Mo, Q., Sun, T., Chen, H., Yu, G., Du, L. (2023): Biogeographic origin of Kurixalus (Anura, Rhacophoridae) on the east Asian islands and tempo of diversification within Kurixalus. Animals 13: 2754. DOI: https://doi.org/10.3390/ani13172754
  51. Nguyen, T.T., Matsui, M., Duc, H.M. (2014): A new tree frog of the genus Kurixalus (Anura: Rhacophoridae) from Vietnam. Curr. Herpetol. 33: 101-111. DOI: https://doi.org/10.5358/hsj.33.101
  52. 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. DOI: https://doi.org/10.1080/00222933.2020.1728411
  53. Pan, T., Wang, H., Duan, S., Ali, I., Yan, P., Cai, R., Wang, M., Zhang, J., Zhang, H., Zhang, B., Wu, X. (2019): Historical population decline and habitat loss in a critically endangered species, the Chinese alligator (Alligator sinensis). Glob. Ecol. Conserv. 20: e00692. DOI: https://doi.org/10.1016/j.gecco.2019.e00692
  54. Pang, C., Tang, S., Yu, G., Zhou, J.-J. (2024): First description of the female and morphological variations with range extension of Kurixalus lenquanensis (Anura, Rhacophoridae). Biodivers. Data J. 12: e130411. DOI: https://doi.org/10.3897/BDJ.12.e130411
  55. Phillips, S.J., Anderson, R.P., Dudík, M., Schapire, R.E., Blair, M.E. (2017): Opening the black box: an open-source release of Maxent. Ecography 40: 887-893. DOI: https://doi.org/10.1111/ecog.03049
  56. Proosdij, A.S.J. van, Sosef, M.S.M., Wieringa, J.J., Raes, N. (2016): Minimum required number of specimen records to develop accurate species distribution models. Ecography 39: 542-552. DOI: https://doi.org/10.1111/ecog.01509
  57. QGIS.org (2023): QGIS. QGIS Association.
  58. R Core Team (2022): R: A Language and environment for statistical computing. Vienna, Austria, Foundation for Statistical Computing.
  59. Rhoden, C.M., Peterman, W.E., Taylor, C.A. (2017): Maxent-directed field surveys identify new populations of narrowly endemic habitat specialists. PeerJ 5: e3632. DOI: https://doi.org/10.7717/peerj.3632
  60. 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. DOI: https://doi.org/10.1093/sysbio/sys029
  61. Sarker, G.C., Wostl, E., Thammachoti, P., Sidik, I., Hamidy, A., Kurniawan, N., Smith, E.N. (2019): New species, diversity, systematics, and conservation assessment of the puppet toads of Sumatra (Anura: Bufonidae: Sigalegalephrynus). Zootaxa 4679: 365-391. DOI: https://doi.org/10.11646/zootaxa.4679.2.9
  62. Shin, Y., Min, M., Borzée, A. (2021): Driven to the edge: Species distribution modeling of a clawed salamander (Hynobiidae: Onychodactylus koreanus) predicts range shifts and drastic decrease of suitable habitats in response to climate change. Ecol. Evol. 11: 14669-14688. DOI: https://doi.org/10.1002/ece3.8155
  63. Trifinopoulos, J., Nguyen, L.-T., von Haeseler, A., Minh, B.Q. (2016): W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Res. 44: W232–W235. DOI: https://doi.org/10.1093/nar/gkw256
  64. Vignali, S., Barras, A.G., Arlettaz, R., Braunisch, V. (2020): SDMtune: An R package to tune and evaluate species distribution models. Ecol. Evol. 10: 11488-11506. DOI: https://doi.org/10.1002/ece3.6786
  65. Warren, D.L., Seifert, S.N. (2011): Ecological niche modeling in Maxent: the importance of model complexity and the performance of model selection criteria. Ecol. Appl. 21: 335-342. DOI: https://doi.org/10.1890/10-1171.1
  66. Wilkinson, J.A., Drewes, R.C., 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. DOI: https://doi.org/10.1016/S1055-7903(02)00212-9
  67. Wu, S.-P., Huang, C.-C., Tsai, C.-L., Lin, T.-E., Jhang, J.-J., Wu, S.-H. (2016): Systematic revision of the Taiwanese genus Kurixalus members with a description of two new endemic species (Anura, Rhacophoridae). ZooKeys 557: 121-153. DOI: https://doi.org/10.3897/zookeys.557.6131
  68. Xu, W., Wu, Y.-H., Zhou, W.-W., Chen, H.-M., Zhang, B.-L., Chen, J.-M., Xu, W., Rao, D.-Q., Zhao, H., Yan, F., Yuan, Z., Jiang, K., Jin, J.-Q., Hou, M., Zou, D., Wang, L.-J., Zheng, Y., Li, J.-T., Jiang, J., Zeng, X.-M., Chen, Y., Liao, Z.-Y., Li, C., Li, X.-Y., Gao, W., Wang, K., Zhang, D.-R., Lu, C., Yin, T., Ding, Z., Zhao, G.-G., Chai, J., Zhao, W.-G., Zhang, Y.-P., Wiens, J.J., Che, J. (2024): Hidden hotspots of amphibian biodiversity in China. Proc. Natl. Acad. Sci. 121: e2320674121. DOI: https://doi.org/10.1073/pnas.2320674121
  69. Yu, G., Hui, H., Rao, D., Yang, J. (2018): A new species of Kurixalus from western Yunnan, China (Anura, Rhacophoridae). ZooKeys 770: 211-226. DOI: https://doi.org/10.3897/zookeys.770.23526
  70. Yu, G., Rao, D., Matsui, M., Yang, J. (2017a): Coalescent-based delimitation outperforms distance-based methods for delineating less divergent species: the case of Kurixalus odontotarsus species group. Sci. Rep. 7: 16124. DOI: https://doi.org/10.1038/s41598-017-16309-1
  71. Yu, G., Wang, J., Hou, M., Rao, D., Yang, J. (2017b): A new species of the genus Kurixalus from Yunnan, China (Anura, Rhacophoridae). ZooKeys 694: 71-93. DOI: https://doi.org/10.3897/zookeys.694.12785
  72. Yu, G.-H., Du, L.-N., Wang, J.-S., Rao, D.-Q., Wu, Z.-J., Yang, J.-X. (2020): From mainland to islands: colonization history in the tree frog Kurixalus (Anura: Rhacophoridae). Curr. Zool. 66: 667-675. DOI: https://doi.org/10.1093/cz/zoaa023
  73. Zanaga, D., Van De Kerchove, R., De Keersmaecker, W., Souverijns, N., Brockmann, C., Quast, R., Wevers, J., Grosu, A., Paccini, A., Vergnaud, S., Cartus, O., Santoro, M., Fritz, S., Georgieva, I., Lesiv, M., Carter, S., Herold, M., Li, L., Tsendbazar, N.-E., Ramoino, F., Arino, O. (2021): ESA WorldCover 10 m 2020 v100. Zenodo. https://doi.org/10.5281/ZENODO.5571936.
  74. Zeng, J., Wang, J.-S., Yu, G.-H., Du, L.-N. (2021): A new species of Kurixalus (Anura, Rhacophoridae) from Guizhou, China. Zool. Res. 42: 227-233.
  75. Zhao, E., Wang, L., Shi, H., Wu, G., Zhao, H. (2005): Chinese rhacophorid frogs and description of a new species of Rhacophorus. Sichuan J. Zool. 24: 297-300.
  76. 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 Kurixalus odontotarsus: insights into the evolution of compound calls. Biol. Open 6: 1913-1918. DOI: https://doi.org/10.1242/bio.028928
  77. Zhu, G., Qiao, H. (2016): Effect of the Maxent model’s complexity on the prediction of species potential distributions. Biodivers. Sci. 24: 1189-1196. DOI: https://doi.org/10.17520/biods.2016265