Published September 2, 2025
| Version v1
Journal article
Open
Mitogenomic analysis reveals two cryptic lineages and provides insights into the historical biogeography of the genus Pseudohynobius (Caudata, Hynobiidae)
Authors/Creators
- 1. Yunnan University, Kunming, China|Guizhou Normal University, Guiyang, China|Southwest United Graduate School, Kunming, China
- 2. Guizhou Normal University, Guiyang, China
- 3. Guiyang Healthcare Vocational University, Guiyang, China
Description
The genus Pseudohynobius exhibits a west-to-east distribution across southwestern China, spanning diverse mountain ranges and elevations. However, knowledge regarding the diversity, phylogeny, and evolutionary history of this genus remains limited. In this study, based on the concept of phylogenetic species, we identified eight phylogenetic species, including two cryptic lineages. Divergence time estimation revealed that Pseudohynobius originated approximately 14.57 million years ago (Ma), with interspecific divergence mainly occurring between 8.0 and 1.4 Ma. Biogeographic analysis indicated that its most recent common ancestor likely inhabited the Hengduan–Dalou Mountains region in southwestern China around 8.62 Ma and that the Dalou Mountains served as a source of outward dispersal and a key region for speciation between 11 and 4 Ma. The lineage divergence dynamics of this genus are coupled with orogenic movements and paleoclimatic shifts, which may have been the primary drivers of its historical diversification. This study underscores the urgency of conducting further surveys and taxonomic studies to avoid underestimating the diversity of this critically endangered genus.
Files
ZSE_article_153576.pdf
Files
(5.1 MB)
| Name | Size | Download all |
|---|---|---|
|
md5:b8ef53d9e47da33c1b9f59b275e083ac
|
4.9 MB | Preview Download |
|
md5:ffa5491bce4b2c64bac3394c5e766195
|
225.5 kB | Preview Download |
Linked records
Additional details
References
- AmphibiaChina (2025) The database of Chinese amphibians. Kunming Institute of Zoology (CAS), Kunming, Yunnan, China. http://www.amphibiachina.org/ [November 25th, 2024]
- Antonelli A, Kissling WD, Flantua SGA, Bermúdez MA, Mulch A, Muellner-Riehl AN, Kreft H, Linder HP, Badgley C, Fjeldså J, Fritz SA, Rahbek C, Herman F, Hooghiemstra H, Hoorn C (2018) Geological and climatic influences on mountain biodiversity. Nature Geoscience 11(10): 718–725. https://doi.org/10.1038/s41561-018-0236-z
- Bouckaert R, Heled J, Kühnert D, Vaughan T, Wu C-H, Xie D, Suchard MA, Rambaut A, Drummond AJ (2014) BEAST 2: A software platform for Bayesian evolutionary analysis. PLoS Computational Biology 10(4): e1003537. https://doi.org/10.1371/journal.pcbi.1003537
- Che J, Zhou WW, Hu JS, Yan F, Papenfuss TJ, Wake DB, Zhang YP (2010) Spiny frogs (Paini) illuminate the history of the Himalayan region and Southeast Asia. Proceedings of the National Academy of Sciences of the United States of America 107(31): 13765–13770. https://doi.org/10.1073/pnas.1008415107
- Chen S (2023) Ultrafast one‐pass FASTQ data preprocessing, quality control, and deduplication using fastp. iMeta 2(2): e107. https://doi.org/10.1002/imt2.107
- Chen G, Xiao Y, Chen X, Gu X (2011) Genetic diversity analysis of D-loop region in populations of Paramesotriton caudopunctatus. Dongwuxue Zazhi 46(3): 55–63.
- Chen MY, Mao RL, Liang D, Kuro-o M, Zeng XM, Zhang P (2015) A reinvestigation of phylogeny and divergence times of Hynobiidae (Amphibia, Caudata) based on 29 nuclear genes. Molecular Phylogenetics and Evolution 83: 1–6. https://doi.org/10.1016/j.ympev.2014.10.010
- Ding WN, Ree RH, Spicer RA, Xing YW (2020) Ancient orogenic and monsoon-driven assembly of the world's richest temperate alpine flora. science 369(6503): 578–581. https://doi.org/10.1126/science.abb4484
- Ding L, Kapp P, Cai F, Garzione CN, Xiong Z, Wang H, Wang C (2022) Timing and mechanisms of Tibetan Plateau uplift. Nature Reviews. Earth & Environment 3(10): 652–667. https://doi.org/10.1038/s43017-022-00318-4
- Dubois A, Ohler A, Pyron RA (2021) New concepts and methods for phylogenetic taxonomy and nomenclature in zoology, exemplified by a new ranked cladonomy of recent amphibians (Lissamphibia). Megataxa 5(1): 1–738. https://doi.org/10.11646/megataxa.5.1.1
- Farnsworth A, Lunt DJ, Robinson SA, Valdes PJ, Roberts WH, Clift PD, Markwick P, Su T, Wrobel N, Bragg F (2019) Past East Asian monsoon evolution controlled by paleogeography, not CO2. Science Advances 5(10): eaax1697. https://doi.org/10.1126/sciadv.aax1697
- Favre A, Päckert M, Pauls SU, Jähnig SC, Uhl D, Michalak I, Muellner-Riehl AN (2015) The role of the uplift of the Qinghai-Tibetan Plateau for the evolution of Tibetan biotas. Biological Reviews of the Cambridge Philosophical Society 90(1): 236–253. https://doi.org/10.1111/brv.12107
- Fei L, Ye C (2000) A new hynobiid subfamily with a new genus and new species of Hynobiidae from West China. Cultum Herpetologica Sinica 8: 64–70.
- Fei L, Ye CY (2016) Amphibians of China. Vol. 1, Science Press, Beijing, 349–368.
- Fei L, Hu SQ, Ye CY, Huang GP (2006) Fauna Sinica: Amphibia. Vol. 1, Science Press, 294–313.
- Feijó A, Ge D, Wen Z, Cheng J, Xia L, Patterson BD, Yang Q (2022) Mammalian diversification bursts and biotic turnovers are synchronous with Cenozoic geoclimatic events in Asia. Proceedings of the National Academy of Sciences of the United States of America 119(49): e2207845119. https://doi.org/10.1073/pnas.2207845119
- Frost DR (2025) Amphibian Species of the World: an Online Reference. Version 6.2. American Museum of Natural History, New York, USA. https://doi.org/10.5531/db.vz.0001 [July 15, 2025]
- Fu J, Wang Y, Zeng X, Liu Z, Zheng Y (2001) Genetic diversity of eastern Batrachuperus (Caudata, Hynobiidae). Ichthyology & Herpetology 2001(4): 1100–1107. https://doi.org/10.1643/0045-8511(2001)001[1100:GDOEBC]2.0.CO;2
- Fu X, Zhu W, Geng J, Yang S, Zhong K, Huang X, Zhang L, Xu X (2021) The present-day Yangtze River was established in the late Miocene: Evidence from detrital zircon ages. Journal of Asian Earth Sciences 205: 104600. https://doi.org/10.1016/j.jseaes.2020.104600
- Fu TT, Sun YB, Gao W, Long CB, Yang CH, Yang XW, Zhang Y, Lan XQ, Huang S, Jin JQ, Murphy RW, Zhang Y, Lai R, Hillis DM, Zhang YP, Che J (2022) The highest-elevation frog provides insights into mechanisms and evolution of defenses against high UV radiation. Proceedings of the National Academy of Sciences of the United States of America 119(46): e2212406119. https://doi.org/10.1073/pnas.2212406119
- Fyhn MB, Phach PV (2015) Late Neogene structural inversion around the northern Gulf of Tonkin, Vietnam: Effects from right‐lateral displacement across the Red River fault zone. Tectonics 34(2): 290–312. https://doi.org/10.1002/2014TC003674
- Guo P, Zhu F, Liu Q, Wang P, Che J, Nguyen TQ (2020) Out of the Hengduan Mountains: Molecular phylogeny and historical biogeography of the Asian water snake genus Trimerodytes (Squamata, Colubridae). Molecular Phylogenetics and Evolution 152: 106927. https://doi.org/10.1016/j.ympev.2020.106927
- He J, Garzanti E, Jiang T, Barbarano M, Liu E, Chen S, Liao Y, Li X, Wang H (2023) Evolution of eastern Asia river systems reconstructed by the mineralogy and detrital-zircon geochronology of modern Red River and coastal Vietnam river sand. Earth-Science Reviews 245: 104572. https://doi.org/10.1016/j.earscirev.2023.104572
- Hoang DT, Chernomor O, Von Haeseler A, Minh BQ, Vinh LS (2018) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35(2): 518–522. https://doi.org/10.1093/molbev/msx281
- Hoorn C, Mosbrugger V, Mulch A, Antonelli A (2013) Biodiversity from mountain building. Nature Geoscience 6(3): 154–154. https://doi.org/10.1038/ngeo1742
- Huang J, Su Y, Zhao J, Liu X, Wang Z (2016) The complete mitogenome sequence of Pseudohynobius jinfo (Urodela: Hynobiidae). Mitochondrial DNA Part A DNA 27(5): 3141–3142. https://doi.org/10.3109/19401736.2015.1007303
- IUCN (2025) The IUCN Red List of Threatened Species. Version 2025-1. International Union for Conservation of Nature and Natural Resources, Gran, Switzerland. https://www.iucnredlist.org [July 15, 2025]
- Jia J, Jiang JP, Zhang MH, Gao KQ (2019) Osteology of Batrachuperus yenyuanensis (Urodela, Hynobiidae), a high-altitude mountain stream salamander from western China. PLoS ONE 14(1): e0211069. https://doi.org/10.1371/journal.pone.0211069
- Jia J, Gao KQ, Jiang JP, Bever GS, Xiong R, Wei G (2021) Comparative osteology of the hynobiid complex Liua-Protohynobius-Pseudohynobius (Amphibia, Urodela): I. Cranial anatomy of Pseudohynobius. Journal of Anatomy 238(2): 219–248. https://doi.org/10.1111/joa.13311
- Jiang J, Cai B, Wang B, Chen W, Wen Z, Zhang D, Wang W, Sui L, Ma S (2024) New vertebrate species discovered in China in 2023. Biodiversity Science 32(11): 24327. https://doi.org/10.17520/biods.2024327
- Jing H, Shu Z, Xia W, Yan S, Ya-Qiu L, Zhi-Jian W (2017) Skin Microstructure of Larval Pseudohynobius jinfo. Dongwuxue Zazhi 52(4): 620–627. https://doi.org/10.13859/j.cjz.201704009
- Kapli P, Lutteropp S, Zhang J, Kobert K, Pavlidis P, Stamatakis A, Flouri T (2017) Multi-rate Poisson tree processes for single-locus species delimitation under maximum likelihood and Markov chain Monte Carlo. Bioinformatics 33(11): 1630–1638. https://doi.org/10.1093/bioinformatics/btx025
- Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780. https://doi.org/10.1093/molbev/mst010
- Kumar S, Stecher G, Tamura K (2016) MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33(7): 1870–1874. https://doi.org/10.1093/molbev/msw054
- Lanfear R, Frandsen PB, Wright AM, Senfeld T, Calcott B (2017) PartitionFinder 2: New methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution 34(3): 772–773. https://doi.org/10.1093/molbev/msw260
- Li S, Tian Y, Gu X (2010) A new species of the genus Pseudohynobius (Caudata, Hynobiidae) from Guizhou, China. Acta Zootaxonomica Sinica 35(2): 407–412.
- Li S, Xu N, Liu J, Lyu J, Wang B, Wei G (2020) A revised species list of amphibians in Guizhou Province. Sichuan Journal of Zoology 39(6): 694–710. https://doi.org/10.11984/j.issn.1000-7083.20200049
- Li SF, Valdes PJ, Farnsworth A, Davies-Barnard T, Su T, Lunt DJ, Spicer RA, Liu J, Deng WYD, Huang J, Tang H, Ridgwell A, Chen LL, Zhou ZK (2021) Orographic evolution of northern Tibet shaped vegetation and plant diversity in eastern Asia. Science Advances 7(5): eabc7741. https://doi.org/10.1126/sciadv.abc7741
- Li XQ, Peng HW, Xiang KL, Xiang XG, Jabbour F, Ortiz RdC, Soltis PS, Soltis DE, Wang W (2024) Phylogenetic evidence clarifies the history of the extrusion of Indochina. Proceedings of the National academy of Sciences of the United States of America 121(35): e2322527121. https://doi.org/10.1073/pnas.232252712
- Luo T, Wen H, Gao K, Zhou J, Zhou J (2021) Phylogeography and cryptic species diversity of Paramesotriton caudopunctatus species group (Salamandridae, Paramesotriton) in Guizhou, China. Asian Herpetological Research 12(2): 188–200. https://doi.org/10.16373/j.cnki.ahr.200025
- Luo T, Yan SS, Xiao N, Zhou JJ, Wang XL, Chen WC, Deng HQ, Zhang BW, Zhou J (2022) Phylogenetic analysis of combined mitochondrial genome and 32 nuclear genes provides key insights into molecular systematics and historical biogeography of Asian warty newts of the genus Paramesotriton (Caudata, Salamandridae). Zoological Research 43(5): 787–804. https://doi.org/10.24272/j.issn.2095-8137.2022.081
- Luo T, Zhao XR, Lan CT, Li W, Deng HQ, Xiao N, Zhou J (2023) Integrated phylogenetic analyses reveal the evolutionary, biogeographic, and diversification history of Asian warty treefrog genus Theloderma (Anura, Rhacophoridae). Ecology and Evolution 13(12): e10829. https://doi.org/10.1002/ece3.10829
- Luo T, Luo FW, Lan CT, Xiao MY, Zhou JJ, Liao M, Xiao N, Zhou J (2024) Evolutionary history of Chinese karst loaches (Nemacheilidae, Karstsinnectes): New insights from mitochondrial-based genomes and description of a new species from Guangxi, China. Zoosystematics and Evolution 100(4): 1473–1486. https://doi.org/10.3897/zse.100.133964
- Luo T, Lan CT, Yu J, Song LX, Fan C, Wang JJ, Zhou JJ, Xiao N, Zhou J (2025) Rapid speciation of Chinese hypogean fishes driven by paleogeoclimatic and morphological adaptations. Current Zoology XX: zoaf010. https://doi.org/10.1093/cz/zoaf010
- Matzke NJ (2013) Probabilistic historical biogeography: New models for founder-event speciation, imperfect detection, and fossils allow improved accuracy and model-testing. Frontiers of Biogeography 5(4): 242–248.
- Matzke NJ (2014) Model selection in historical biogeography reveals that founder-event speciation is a crucial process in island clades. Systematic Biology 63(6): 951–970. https://doi.org/10.1093/sysbio/syu056
- Meleshko O, Martin MD, Korneliussen TS, Schröck C, Lamkowski P, Schmutz J, Healey A, Piatkowski BT, Shaw AJ, Weston DJ, Flatberg KI, Szövényi P, Hassel K, Stenøien HK (2021) Extensive genome-wide phylogenetic discordance is due to incomplete lineage Sorting and not ongoing introgression in a rapidly radiated bryophyte genus. Molecular Biology and Evolution 38(7): 2750–2766. https://doi.org/10.1093/molbev/msab063
- Meng G, Li Y, Yang C, Liu S (2019) MitoZ: A toolkit for animal mitochondrial genome assembly, annotation and visualization. Nucleic Acids Research 47(11): e63. https://doi.org/10.1093/nar/gkz173
- Nguyen LT, Schmidt HA, Von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274. https://doi.org/10.1093/molbev/msu300
- Peng R, Zhang P, Xiong JL, Gu HJ, Zeng XM, Zou FD (2010) Rediscovery of Protohynobius puxiongensis (Caudata, Hynobiidae) and its phylogenetic position based on complete mitochondrial genomes. Molecular Phylogenetics and Evolution 56(1): 252–258. https://doi.org/10.1016/j.ympev.2009.12.011
- Puillandre N, Brouillet S, Achaz G (2021) ASAP: Assemble species by automatic partitioning. Molecular Ecology Resources 21(2): 609–620. https://doi.org/10.1111/1755-0998.13281
- Pyron RA, Wiens JJ (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution 61(2): 543–583. https://doi.org/10.1016/j.ympev.2011.06.012
- Rahbek C, Borregaard MK, Antonelli A, Colwell RK, Holt BG, Nogues-Bravo D, Rasmussen CMØ, Richardson K, Rosing MT, Whittaker RJ, Fjeldså J (2019a) Building mountain biodiversity: Geological and evolutionary processes. Science 365(6458): 1114–1119. https://doi.org/10.1126/science.aax0151
- Rahbek C, Borregaard MK, Colwell RK, Dalsgaard B, Holt BG, Morueta-Holme N, Nogues-Bravo D, Whittaker RJ, Fjeldså J (2019b) Humboldt's enigma: What causes global patterns of mountain biodiversity? Science 365(6458): 1108–1113. https://doi.org/10.1126/science.aax0149
- Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Posterior Summarization in Bayesian Phylogenetics Using Tracer 1.7. Systematic Biology 67(5): 901–904. https://doi.org/10.1093/sysbio/syy032
- Rivas-González I, Rousselle M, Li F, Zhou L, Dutheil JY, Munch K, Shao Y, Wu D, Schierup MH, Zhang G (2023) Pervasive incomplete lineage sorting illuminates speciation and selection in primates. Science 380(6648): eabn4409. https://doi.org/10.1126/science.abn4409
- Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029
- Sun BN, Wu JY, Liu YS, Ding ST, Li XC, Xie SP, Yan DF, Lin ZC (2011) Reconstructing Neogene vegetation and climates to infer tectonic uplift in western Yunnan, China. Palaeogeography, Palaeoclimatology, Palaeoecology 304(3): 328–336. https://doi.org/10.1016/j.palaeo.2010.09.023
- Tian Y, Li S, Gu X (2006) A new species of genus Pseudohynobius (Caudata, Hynobiidae)—Pseudohynobius shuichengensis. Acta Zoologica Sinica 52(3): 522–527.
- van Elst T, Sgarlata GM, Schüßler D, Tiley GP, Poelstra JW, Scheumann M, Blanco MB, Aleixo-Pais IG, Rina Evasoa M, Ganzhorn JU, Goodman SM, Hasiniaina AF, Hending D, Hohenlohe PA, Ibouroi MT, Iribar A, Jan F, Kappeler PM, Le Pors B, Manzi S, Olivieri G, Rakotonanahary AN, Rakotondranary SJ, Rakotondravony R, Ralison JM, Ranaivoarisoa JF, Randrianambinina B, Rasoloarison RM, Rasoloharijaona S, Rasolondraibe E, Teixeira H, Zaonarivelo JR, Louis EE, Yoder AD, Chikhi L, Radespiel U, Salmona J (2025) Integrative taxonomy clarifies the evolution of a cryptic primate clade. Nature Ecology & Evolution 9(1): 57–72. https://doi.org/10.1038/s41559-024-02547-w
- Wang Y, Zheng J, Zhang W, Li S, Liu X, Yang X, Liu Y (2012) Cenozoic uplift of the Tibetan Plateau: Evidence from the tectonic–sedimentary evolution of the western Qaidam Basin. Geoscience Frontiers 3(2): 175–187. https://doi.org/10.1016/j.gsf.2011.11.005
- Wang X, Liang D, Wang X, Tang M, Liu Y, Liu S, Zhang P (2022) Phylogenomics reveals the evolution, biogeography, and diversification history of voles in the Hengduan Mountains. Communications Biology 5(1): 1124. https://doi.org/10.1038/s42003-022-04108-y
- Wang Z, Othman SN, Qiu Z, Lu Y, Prasad VK, Dong Y, Lu CH, Borzée A (2023) An Isolated and Deeply Divergent Hynobius Species from Fujian, China. Animals 13: 1661. https://doi.org/10.3390/ani13101661
- Wen H, Luo T, Wang Y, Wang S, Liu T, Xiao N, Zhou J (2022) Molecular phylogeny and historical biogeography of the cave fish genus Sinocyclocheilus (Cypriniformes, Cyprinidae) in southwest China. Integrative Zoology 17(2): 311–325. https://doi.org/10.1111/1749-4877.12624
- Xia Y, Gu HF, Peng R, Chen Q, Zheng YC, Murphy RW, Zeng XM (2012) COI is better than 16S rRNA for DNA barcoding Asiatic salamanders (Amphibia, Caudata, Hynobiidae). Molecular Ecology Resources 12(1): 48–56. https://doi.org/10.1111/j.1755-0998.2011.03055.x
- Xing Y, Ree RH (2017) Uplift-driven diversification in the Hengduan Mountains, a temperate biodiversity hotspot. Proceedings of the National Academy of Sciences of the United States of America 114(17): E3444–E3451. https://doi.org/10.1073/pnas.1616063114
- Xu N, Zeng X, Fu J (2007) A new species of the genus Pseudohynobius (Caudata, Hynobiidae) from China. Acta Zootaxonomica Sinica 31(1): 230–233.
- Xu W, Wu YH, Zhou WW, Chen HM, Zhang BL, Chen JM, Xu W, Rao DQ, Zhao H, Yan F (2024) Hidden hotspots of amphibian biodiversity in China. Proceedings of the National Academy of Sciences of the United States of America 121(20): e2320674121. https://doi.org/10.1073/pnas.2320674121
- Yang C, Wang C, Tian Y, Li S, Gu X (2013) Phylogenetic Relationships of Pseudohynobius (Caudata, Hynobiidae) Based on Mitochondrial cyt b Sequences. Journal of Liupanshui Normal University 25(1): 39–43.
- Yang C, Fu T, Lan X, Zhang Y, Nneji LM, Murphy RW, Sun Y, Che J (2019) Comparative skin histology of frogs reveals high-elevation adaptation of the Tibetan Nanorana parkeri. Asian Herpetological Research 10(2): 79–85. https://doi.org/10.16373/j.cnki.ahr.180093
- Zachos JC, Dickens GR, Zeebe RE (2008) An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature 451(7176): 279–283. https://doi.org/10.1038/nature06588
- Zeng X, Fu J, Chen L, Tian Y, Chen X (2006) Cryptic species and systematics of the hynobiid salamanders of the Liua–Pseudohynobius complex: Molecular and phylogenetic perspectives. Biochemical Systematics and Ecology 34(6): 467–477. https://doi.org/10.1016/j.bse.2006.01.006
- Zhang P, Chen YQ, Zhou H, Liu YF, Wang XL, Papenfuss TJ, Wake DB, Qu LH (2006) Phylogeny, evolution, and biogeography of Asiatic Salamanders (Hynobiidae). Proceedings of the National Academy of Sciences of the United States of America 103(19): 7360–7365. https://doi.org/10.1073/pnas.0602325103
- Zhang B, Liu R, Xiang H, Wan J, Huang X (2009) FT dating of fault rocks in the central‐southern section of the Red River Fault zone and its geological implications. Seismology and Geology 31(1): 44–56.
- Zhang J, Kapli P, Pavlidis P, Stamatakis A (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics 29(22): 2869–2876. https://doi.org/10.1093/bioinformatics/btt499
- Zhang D, Gao F, Jakovlić I, Zou H, Zhang J, Li WX, Wang GT (2020a) PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20(1): 348–355. https://doi.org/10.1111/1755-0998.13096
- Zhang DR, Hui H, Yu GH, Song XQ, Liu S, Yuan SQ, Xiao H, Rao DQ (2020b) Shared response to changes in drainage basin: Phylogeography of the Yunnan small narrow‐mouthed frog, Glyphoglossus yunnanensis (Anura: Microhylidae). Ecology and Evolution 10(3): 1567–1580. https://doi.org/10.1002/ece3.6011
- Zhang Y, Wang M, Cheng R, Luo Y, Li Y, Liu Z, Chen Q, Shen Y (2022) Mitochondrial characteristics of Pseudohynobius flavomaculatus a protected salamander in China, and biogeographical implications for the family Hynobiidae (Amphibia, Caudata). Zoosystematics and Evolution 98(2): 263–274. https://doi.org/10.3897/zse.98.66578
- Zhao Y, Su L, Zhang Z, Wang X (2016) Phylogenetic relationships of Pseudohynobius (Urodela, Hynobiidae) inferred from DNA barcoding analysis. Genetics and Molecular Research 15(2): gmr.15028155. http://dx.doi.org/10.4238/gmr.15028155.
- Zheng Y, Peng R, Kuro-o M, Zeng X (2011) Exploring patterns and extent of bias in estimating divergence time from mitochondrial DNA sequence data in a particular lineage: A case study of Salamanders (Order Caudata). Molecular Biology and Evolution 28(9): 2521–2535. https://doi.org/10.1093/molbev/msr072
- Zhou Q, Chen P (1993) Lithofacies change and palaeogeographical evolution during Late Cenozoic in Guizhou and its vicinity. Geology of Guizhou 10(3): 201–207.