Abstract
The karst ecosystem in southwestern China is a global hotspot for cavefish diversity research, yet the origins and evolutionary history of this diversity remain poorly understood. To elucidate their evolutionary origins and history, we analyzed 183 cavefish species and their close relatives from ten clades. Our findings indicate that freshwater fishes began colonizing caves ~44 million years ago (Ma). Speciation was driven by in situ diversification, starting around 43.2 Ma, increasing sharply by ~35 Ma and 18 Ma, and peaking at ~8.3 Ma, 2.5 Ma, and 1.5 Ma. Distinct hydrological basins exhibited divergent diversification patterns. Dispersal between the Pearl and Yangtze River basins began by ~24.1 Ma, accelerated around 21 Ma, 13 Ma, and 9.6 Ma, and peaked at ~13 Ma, 5 Ma, and 2 Ma. We propose that river drainages in southwestern China developed stepwise from the late Eocene to early Pleistocene, with connectivity between the Pearl and Yangtze basins established by the late Oligocene ( ~ 26 Ma). The origin and diversification of cavefishes are closely linked to the evolution of karst landscapes, shaped by orogeny and monsoon-driven climate changes since the late Eocene. These insights are crucial for informing conservation strategies for these unique habitats under ongoing climate change.

Similar content being viewed by others
Data availability
The sequences reported in this paper have been deposited in GenBank (accession nos. ON116493–ON116532, ON148332–ON148334, OP473612–OP473909, OQ754141–OQ754145, OQ754148–OQ754150, OQ754152, OQ754154, OQ754155, OQ754157–OQ754159, PQ222367–PQ222376, PX715913–PX715924). All publicly available sequence data (e.g., GenBank accessions) used in this study are detailed in Supplementary Tables 1–8 and Supplementary Data 2–4. Source data underlying Fig. 1 are provided in Supplementary Data 1. Source data for Fig. 2 have been deposited in the Science Data Bank (https://doi.org/10.57760/sciencedb.35098)88. Source data for Figs. 3A, 3B, 3C, 3D, 3E, and 3F are provided in Supplementary Data 5, Supplementary Data 6, Supplementary Table 14, Supplementary Table 15, Supplementary Table 16, and PANGAEA39,97 (Supplementary Table 17), respectively. Source data underlying Fig. 4 are available in PANGAEA39,97 and included in Supplementary Table 17. All other datasets are provided in the supplementary information and data files associated with this publication. Source data for sequence alignments, time‑calibrated phylogenies, diversification analyses, and paleoenvironmental reconstructions are available from the Science Data Bank. (https://doi.org/10.57760/sciencedb.35098).
Code availability
All analyses were conducted using open-source software tools, with detailed parameters specified in the corresponding methods sections. Scripts used for generating the associated figures have been archived at the Science Data Bank (https://doi.org/10.57760/sciencedb.35098).
References
Lu, H. et al. Formation and evolution of the Asian landscape during the Cenozoic. Innov. Geosci. 1, 100020 (2023).
Ding, L. et al. Timing and mechanisms of Tibetan Plateau uplift. Nat. Rev. Earth Environ. 3, 652–667 (2022).
Yuan, J. et al. Rapid drift of the Tethyan Himalaya terrane before two-stage India-Asia collision. Natl. Sci. Rev. 8, nwaa173 (2021).
Luo, Y. & Li, S. The stepwise Indian–Eurasian collision and uplift of the Himalayan-Tibetan plateau drove the diversification of high-elevation Scytodes spiders. Cladistics 38, 582–594 (2022).
Favre, A. et al. The role of the uplift of the Qinghai-Tibetan Plateau for the evolution of Tibetan biotas. Biol. Rev. 90, 236–253 (2015).
Wu, F. et al. Reorganization of Asian climate in relation to Tibetan Plateau uplift. Nat. Rev. Earth Environ. 3, 684–700 (2022).
Zheng, H. et al. Pre-Miocene birth of the Yangtze River. Proc. Natl. Acad. Sci. USA 110, 7556–7561 (2013).
Zhang, Z. et al. Late Oligocene formation of the Pearl River triggered by the opening of the South China Sea. Geophys. Res. Lett. 50, e2023GL103049 (2023).
Spicer, R. A. Tibet, the Himalaya, Asian monsoons and biodiversity–In what ways are they related?. Plant Divers. 39, 233–244 (2017).
Ding, W. N., Ree, R. H., Spicer, R. A. & Xing, Y. W. Ancient orogenic and monsoon-driven assembly of the world’s richest temperate alpine flora. Science 369, 578–581 (2020).
Che, Y. & Yu, J. Karst in China 81–95 (Science Press, 1985).
Li, X. Q. et al. Biotic colonization of subtropical East Asian caves through time. Proc. Natl. Acad. Sci. USA 119, e2207199119 (2022).
Wen, H. et al. Molecular phylogeny and historical biogeography of the cave fish genus Sinocyclocheilus (Cypriniformes: Cyprinidae) in southwest China. Integr. Zool. 17, 311–325 (2022).
Stokes, M. F. et al. Erosion of heterogeneous rock drives diversification of Appalachian fishes. Science 380, 855–859 (2023).
Hoorn, C., Mosbrugger, V., Mulch, A. & Antonelli, A. Biodiversity from mountain building. Nat. Geosci. 6, 154–154 (2013).
Rahbek, C. et al. Humboldt’s enigma: what causes global patterns of mountain biodiversity?. Science 365, 1108–1113 (2019).
Rahbek, C. et al. Building mountain biodiversity: geological and evolutionary processes. Science 365, 1114–1119 (2019).
Klaus, S., Morley, R. J., Plath, M., Zhang, Y.-P. & Li, J.-T. Biotic interchange between the Indian subcontinent and mainland Asia through time. Nat. Commun. 7, 12132 (2016).
Xu, W. et al. Herpetological phylogeographic analyses support a Miocene focal point of Himalayan uplift and biological diversification. Natl. Sci. Rev. 8, nwaa263 (2021).
White, W. B. & Culver, D. C. in Encyclopedia of Caves (Third Edition). Cave, Definition of (eds White, W. B., Culver, C. D. & Pipan, T.) Ch. 29, 255–259 (Academic Press, 2019).
Borowsky, R. Cavefishes. Curr. Biol. 28, R60–R64 (2018).
Zhao, Y. H., Gozlan, R. E. & Zhang, C. G. Out of sight out of mind: current knowledge of Chinese cave fishes. J. Fish Biol. 79, 1545–1562 (2011).
Ma, L., Zhao, Y. H. & Yang, J. X. in Encyclopedia of Caves (Third Edition). Cavefish of China (eds White, W. B., Culver, C. D. & Pipan, T.) Ch. 28, 237–254 (Academic Press, 2019).
Mao, T. R. et al. Evolution in Sinocyclocheilus cavefish is marked by rate shifts, reversals, and origin of novel traits. BMC Ecol. Evol. 21, 1–14 (2021).
Mao, T. et al. Evolving in the darkness: phylogenomics of Sinocyclocheilus cavefishes highlights recent diversification and cryptic diversity. Mol. Phylogenet. Evol. 168, 107400 (2022).
Shu, S. S., Jiang, W. S., Whitten, T., Yang, J. X. & Chen, X. Y. Drought and China’s cave species. Science 340, 272–272 (2013).
Duan, Y. F., Li, M., Xu, K. W., Zhang, L. & Zhang, L. B. Protect China’s karst cave habitats. Science 374, 699–699 (2021).
Wu, L. et al. A long-ignored unique ecosystem of cavefishes in the southern karst: achievements, challenges, and prospects. Environ. Sci. Pollut. Res. 30, 90489–90499 (2023).
Mao, T. et al. Caves as species pumps: key innovations, isolation and periodic introgression drive the world’s largest cavefish radiation in a dynamic karstic landscape. Mol. Ecol. 34, e70038 (2025).
Lee, T. Y. & Lawver, L. A. Cenozoic plate reconstruction of Southeast Asia. Tectonophysics 251, 85–138 (1995).
Chung, S. L. et al. Tibetan tectonic evolution inferred from spatial and temporal variations in post-collisional magmatism. Earth Sci. Rev. 68, 173–196 (2005).
Leloup, P. H. et al. New constraints on the structure, thermochronology, and timing of the Ailao Shan-Red River shear zone, SE Asia. J. Geophys. Res. Solid Earth 106, 6683–6732 (2001).
Schärer, U., Lian-Sheng, Z. & Tapponnier, P. Duration of strike-slip movements in large shear zones: The Red River belt, China. Earth Planet. Sci. Lett. 126, 379–397 (1994).
Li, S. et al. Oligocene deformation of the Chuandian Terrane in the SE margin of the Tibetan Plateau related to the extrusion of Indochina. Tectonics 39, e2019TC005974 (2020).
Li, X. Q. et al. Phylogenetic evidence clarifies the history of the extrusion of Indochina. Proc. Natl. Acad. Sci. USA 121, e2322527121 (2024).
Liu, X., Hou, J. & You, X. Neotectonic motion of Youjiang fault zone. J. Seismol. Res. 10, 175–183 (1987).
Nie, G., Yang, S., Zhang, P. & Li, B. Characteristics and tectonic significance of cenozoic strike-slip faults in Youjiang Region. Geotecton. Metallog. 43, 1094–1105 (2019).
Judd, E. J. et al. A 485-million-year history of Earth’s surface temperature. Science 385, eadk3705 (2024).
Westerhold, T. et al. An astronomically dated record of Earth’s climate and its predictability over the last 66 million years. Science 369, 1383–1387 (2020).
Luo, T. et al. Rapid speciation of Chinese hypogean fishes driven by paleogeoclimatic and morphological adaptations. Curr. Zool. 71, 715–732 (2025).
Ma, Z. et al. Head horn enhances hydrodynamic perception in eyeless cavefish. Adv. Sci. 11, 2406707 (2024).
Zhang, M. et al. A splicing mutation in mitfa is involved in the depigmentation of Cavefish Triplophysa rosa. Mol. Biol. Evol. 42, msaf175 (2025).
Lam, S. M. et al. Quantitative lipidomics and spatial ms-imaging uncovered neurological and systemic lipid metabolic pathways underlying troglomorphic adaptations in cave-dwelling fish. Mol. Biol. Evol. 39, msac050 (2022).
Ding, W. et al. Palaeovegetation variation in response to the late Oligocene-early Miocene East Asian summer monsoon in the Ying-Qiong Basin, South China Sea. Palaeogeogr. Palaeoclimatol. Palaeoecol. 567, 110205 (2021).
Ren, J., Schubert, B. A., Lukens, W. E. & Quan, C. Low oxygen isotope values of fossil cellulose indicate an intense monsoon in East Asia during the late Oligocene. Palaeogeogr. Palaeoclimatol. Palaeoecol. 577, 110556 (2021).
Vornlocher, J. R., Lukens, W. E., Schubert, B. A. & Quan, C. Late Oligocene precipitation seasonality in East Asia based on δ13C profiles in fossil wood. Paleoceanogr. Paleoclimatol. 36, e2021PA004229 (2021).
Chen, C. et al. Lower-altitude of the Himalayas before the mid-Pliocene as constrained by hydrological and thermal conditions. Earth Planet. Sci. Lett. 545, 116422 (2020).
Leloup, P. H. et al. The Ailao Shan-Red river shear zone (Yunnan, China), tertiary transform boundary of Indochina. Tectonophysics 251, 3–84 (1995).
Tapponnier, P. et al. The Ailao Shan/Red River metamorphic belt: tertiary left-lateral shear between Indochina and South China. Nature 343, 431–437 (1990).
Farnsworth, A. et al. Past East Asian monsoon evolution controlled by paleogeography, not CO2. Sci. Adv. 5, eaax1697 (2019).
Zhang, Q. et al. East Asian winter monsoon intensification over the Northwest Pacific Ocean driven by late Miocene atmospheric CO2 decline. Sci. Adv. 10, eadm8270 (2024).
Wan, S. et al. Interactive forces of temperature and topographic uplift shaped the East Asian monsoon rainfall evolution since the Oligocene. Innov. Geosci. 3, 100141 (2025).
Wang, Y. et al. Cenozoic exhumation of the Ailaoshan-Red River shear zone: New insights from low-temperature thermochronology. Tectonics 39, e2020TC006151 (2020).
Zhang, J., Zhong, D., Sang, H. & Zhou, Y. Structural and Geochronological Evidence for Multiple Episodes of Tertiary Deformation along the Ailaoshan-Red River Shear Zone, Southeastern Asia, Since the Paleocene. Acta Geol. Sin. Engl. Ed. 80, 79–96 (2006).
Jolivet, M. & Boulvais, P. Global significance of oxygen and carbon isotope compositions of pedogenic carbonates since the Cretaceous. Geosci. Front. 12, 101132 (2021).
Liu, Y. et al. New chronological constraints on the Plio-Pleistocene uplift of the Guizhou Plateau, SE margin of the Tibetan Plateau. Quat. Geochronol. 67, 101237 (2022).
Luo, T. et al. Phylogenetic relationships of Nemacheilidae cavefish (Heminoemacheilus, Oreonectes, Yunnanilus, Paranemachilus, and Troglonectes) revealed by analysis of mitochondrial genome and seven nuclear genes. Zool. Res. 44, 693–697 (2023).
Ding, L. et al. Quantifying the rise of the Himalaya orogen and implications for the South Asian monsoon. Geology 45, 215–218 (2017).
Zhang, B. et al. Comparative genomics reveals the hybrid origin of a macaque group. Sci. Adv. 9, eadd3580 (2023).
Jensen, A. et al. Complex evolutionary history with extensive ancestral gene flow in an African primate radiation. Mol. Biol. Evol. 40, msad247 (2023).
Wang, X. et al. Phylogenomics reveals the evolution, biogeography, and diversification history of voles in the Hengduan Mountains. Commun. Biol. 5, 1124 (2022).
Foley, N. M. et al. Karyotypic stasis and swarming influenced the evolution of viral tolerance in a species-rich bat radiation. Cell Genom. 4, 100482 (2024).
Qian, Y. et al. The role of introgression during the radiation of endemic fishes adapted to living at extreme altitudes in the Tibetan Plateau. Mol. Biol. Evol. 40, msad129 (2023).
Svardal, H. et al. Ancestral hybridization facilitated species diversification in the Lake Malawi Cichlid fish adaptive radiation. Mol. Biol. Evol. 37, 1100–1113 (2020).
He, J. et al. Evolution of eastern Asia river systems reconstructed by the mineralogy and detrital-zircon geochronology of modern Red River and coastal Vietnam river sand. Earth Sci. Rev. 245, 104572 (2023).
Gilley, L. D. et al. Direct dating of left-lateral deformation along the Red River shear zone, China and Vietnam. J. Geophys. Res. Solid Earth 108, 2127 (2003).
Fyhn, M. B. & Phach, P. V. Late Neogene structural inversion around the northern Gulf of Tonkin, Vietnam: effects from right-lateral displacement across the Red River fault zone. Tectonics 34, 290–312 (2015).
Zhang, B., Liu, R., Xiang, H., Wan, J. & Huang, X. FT dating of fault rocks in the central-southern section of the Red River Fault zone and its geological implications. Seismol. Geol. 31, 44–56 (2009).
Clift, P. D. et al. Correlation of Himalayan exhumation rates and Asian monsoon intensity. Nat. Geosci. 1, 875–880 (2008).
Replumaz, A., Lacassin, R., Tapponnier, P. & Leloup, P. Large river offsets and Plio-Quaternary dextral slip rate on the Red River fault (Yunnan, China). J. Geophys. Res. Solid Earth 106, 819–836 (2001).
Fu, X. et al. The present-day Yangtze River was established in the late Miocene: evidence from detrital zircon ages. J. Asian Earth Sci. 205, 104600 (2021).
Zheng, H. et al. Formation of the First Bend in the late Eocene gave birth to the modern Yangtze River, China. Geology 49, 35–39 (2021).
Chen, F. et al. Evolution of the Yangtze River and its biodiversity. Innovation 4, 100417 (2023).
Clift, P. D. et al. A Late Eocene-Oligocene Through-Flowing River Between the Upper Yangtze and South China Sea. Geochem. Geophys. Geosyst. 21, e2020GC009046 (2020).
Zhou, Q. & Chen, P. Lithofacies change and palaeogeographical evolution during Late Cenozoic in Guizhou and its vicinity. Geol. Guizhou 10, 201–207 (1993).
Luo, T. et al. Evolutionary history of Chinese karst loaches (Nemacheilidae, Karstsinnectes): new insights from mitochondrial-based genomes and description of a new species from Guangxi, China. Zoosyst. Evol. 100, 1473–1486 (2024).
Zhang, J. et al. Evolutionary history of the Arctic flora. Nat. Commun. 14, 4021 (2023).
Lu, C. et al. Contrasting evolutionary trajectories of terrestrial vertebrates in the Hengduan Mountains hotspot. Natl. Sci. Rev. 12, nwaf157 (2025).
Cao, G. L. et al. Cenozoic geoclimatic changes drove the evolutionary dynamics of floristic endemism on the Qinghai–Tibet Plateau. Proc. Natl. Acad. Sci. USA 122, e2426017122 (2025).
Jiang, D., Klaus, S., Zhang, Y.-P., Hillis, D. M. & Li, J.-T. Asymmetric biotic interchange across the Bering land bridge between Eurasia and North America. Natl. Sci. Rev. 6, 739–745 (2019).
Chen, S., Zhou, Y., Chen, Y. & Gu, J. fastp: an ultra-fast all-in-one FASTQ preprocessor. Bioinformatics 34, i884–i890 (2018).
Allio, R. et al. MitoFinder: efficient automated large-scale extraction of mitogenomic data in target enrichment phylogenomics. Mol. Ecol. Resour. 20, 892–905 (2020).
Zhang, D. et al. PhyloSuite: an integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Mol. Ecol. Resour. 20, 348–355 (2020).
Katoh, K. & Standley, D. M. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Mol. Biol. Evol. 30, 772–780 (2013).
Kumar, S., Stecher, G. & Tamura, K. MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Mol. Biol. Evol. 33, 1870–1874 (2016).
Lanfear, R., Frandsen, P. B., Wright, A. M., Senfeld, T. & Calcott, B. PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Mol. Biol. Evol. 34, 772–773 (2017).
Bouckaert, R. et al. BEAST 2: a software platform for Bayesian evolutionary analysis. PLoS Comput. Biol. 10, e1003537 (2014).
Luo, T. et al. Evolutionary history of Chinese cavefishes parallels paleogeoclimatic and river capture processes [Data set]. Sci. Data Bank https://doi.org/10.57760/sciencedb.35098 (2026).
Rambaut, A., Drummond, A. J., Xie, D., Baele, G. & Suchard, M. A. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Syst. Biol. 67, 901–904 (2018).
Matzke, N. J. Probabilistic historical biogeography: new models for founder-event speciation, imperfect detection, and fossils allow improved accuracy and model-testing. Front. Biogeogr. 5, 242–248 (2013).
Ree, R. H. & Smith, S. A. Maximum likelihood inference of geographic range evolution by dispersal, local extinction, and cladogenesis. Syst. Biol. 57, 4–14 (2008).
Ronquist, F. Dispersal-vicariance analysis: a new approach to the quantification of historical biogeography. Syst. Biol. 46, 195–203 (1997).
Landis, M. J., Matzke, N. J., Moore, B. R. & Huelsenbeck, J. P. Bayesian analysis of biogeography when the number of areas is large. Syst. Biol. 62, 789–804 (2013).
Yu, Y., Blair, C. & He, X. RASP 4: ancestral state reconstruction tool for multiple genes and characters. Mol. Biol. Evol. 37, 604–606 (2020).
Muggeo, V. M. Estimating regression models with unknown break-points. Stat. Med. 22, 3055–3071 (2003).
Muggeo, V. M. Segmented: an R package to fit regression models with broken-line relationships. R News 8, 20–25 (2008).
Westerhold, T. Cenozoic global reference benthic carbon and oxygen isotope dataset (CENOGRID) [Data set]. PANGAEA https://doi.org/10.1594/PANGAEA.917503 (2020).
Jiang, Z. C. et al. Calculation of atmospheric CO2 sink formed in karst progresses of the karst divided regions in China. Carsolog. Sin. 30, 363–367 (2011).
Clift, P. D. Controls on the erosion of Cenozoic Asia and the flux of clastic sediment to the ocean. Earth Planet. Sci. Lett. 241, 571–580 (2006).
Acknowledgements
This work was supported by the following programs: the Endangerment Processes and Mechanisms of Endangered Flagship Animals (2022YFF1301502), the Strategic Priority Research Program of the Chinese Academy of Sciences (CAS) (XDB31000000), and the Guizhou Normal University Academic Emerging Talent Fund Project (Qianshi Xin Miao [2021] 20). We are grateful to Prof. Li-Na Du for providing tissue samples of Karstsinnectes longzhouensis and Troglonectes hechiensis. We are grateful to Mr. Jia-Hu Lan for providing tissue samples of Karstsinnectes acridorsalis and Troglonectes barbatus used in this study. We also thank Chang-Ting Lan, Jing Yu, Ya-Li Wang, Xin-Rui Zhao, Xing-Liang Wang, Zhi-Xia Chen, and Ming-Le Mao for their assistance in field surveys and laboratory work. We thank LetPub (www.letpub.com.cn) for its linguistic assistance during the preparation of this manuscript.
Author information
Authors and Affiliations
Contributions
T.L.: Data curation, formal analysis, investigation, methodology, visualization, writing—original draft, writing—review and editing; M.Y.X.: Conceptualization, data curation, investigation, methodology, project administration, supervision, visualization, writing—review and editing; M.L. and J. J.W.: Data curation; N.X.: Writing—review and editing; J.Z.: Conceptualization, formal analysis, funding acquisition, investigation, methodology, project administration, resources, supervision, visualization, writing—review and editing. All authors gave final approval for publication and agreed to be held accountable for the work performed therein.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Peer review
Peer review information
Communications Biology thanks Xidong Mu and the other anonymous reviewer(s) for their contribution to the peer review of this work. Primary handling editor: George Inglis. [A peer review file is available].
Additional information
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by-nc-nd/4.0/.
About this article
Cite this article
Luo, T., Xiao, MY., Liao, M. et al. Evolutionary history of Chinese cavefishes parallels paleogeoclimatic and river capture processes. Commun Biol (2026). https://doi.org/10.1038/s42003-026-09881-8
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s42003-026-09881-8


