The oldest articulated bony fish from the early Silurian period

All data analysed in this paper, including the phylogenetic datasets, are available as part of the Article, Extended Data Figs. 1–8 and the Supplementary Information. Supplementary Data 1–7 are available at Figshare (https://doi.org/10.6084/m9.figshare.28881827)53. The time ranges for each fossil taxa derived from the DEEPBONE database (www.deepbone.org). The nomenclature described in this publication has been registered at ZooBank (https://zoobank.org/References/4631593F-F077-484A-B7F6-32DFEB92E22C). All specimens described in this study are archived and available on request from the IVPP, Chinese Academy of Sciences, Beijing, China.

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Article ADS PubMed PubMed Central Google Scholar

Friedman, M. & Brazeau, M. D. A reappraisal of the origin and basal radiation of the Osteichthyes. J. Vertebr. Paleontol. 30, 36–56 (2010).

Friedman, M. & Sallan, L. C. Five hundred million years of extinction and recovery: a Phanerozoic survey of large-scale diversity patterns in fishes. Palaeontology 55, 707–742 (2012).

Botella, H. et al. Jaws and teeth of the earliest bony fishes. Nature 448, 583–586 (2007).

Article ADS CAS PubMed Google Scholar

Chen, D.-L. et al. The stem osteichthyan Andreolepis and the origin of tooth replacement. Nature 539, 237–241 (2016).

Zhu, M. et al. The oldest articulated osteichthyan reveals mosaic gnathostome characters. Nature 458, 469–474 (2009).

Lu, J. et al. Largest Silurian fish illuminates the origin of osteichthyan characters. Nature https://doi.org/10.1038/s41586-025-10008-y (2026).

Pearson, D. M. & Westoll, T. S. The Devonian actinopterygian Cheirolepis Agassiz. Earth Environ. Sci. Trans. R. Soc. Edinb. 70, 337–399 (1979).

Zhu, Y.-A. et al. The oldest complete jawed vertebrates from the early Silurian of China. Nature 609, 954–958 (2022).

Gai, Z.-K. et al. Galeaspid anatomy and the origin of vertebrate paired appendages. Nature 609, 959–963 (2022).

Chen, Y. et al. A new family of galeaspids (jawless stem-Gnathostomata) from the early Silurian of Chongqing, southwestern China. Acta Geol. Sin. En. Ed. 96, 430–439 (2022).

Brazeau, M. D. & Friedman, M. The characters of Palaeozoic jawed vertebrates. J. Zool. Lond. 170, 779–821 (2014).

Cui, X.-D. et al. Bony-fish-like scales in a Silurian maxillate placoderm. Nat. Commun. 14, 7622 (2023).

Article ADS CAS PubMed PubMed Central Google Scholar

Irisarri, I. et al. Phylotranscriptomic consolidation of the jawed vertebrate timetree. Nat. Ecol. Evol. 1, 1370–1378 (2017).

Article PubMed PubMed Central Google Scholar

Chen, D.-L. et al. Scale morphology and squamation of the late Silurian osteichthyan Andreolepis from Gotland, Sweden. Hist. Biol. 24, 411–423 (2012).

Janvier, P. Early Vertebrates (Clarendon, 1996).

Brazeau, M. D. The braincase and jaws of a Devonian ‘acanthodian’ and modern gnathostome origins. Nature 457, 305–308 (2009).

Davis, S. P., Finarelli, J. A. & Coates, M. I. Acanthodes and shark-like conditions in the last common ancestor of modern gnathostomes. Nature 486, 247–250 (2012).

Zhu, M. et al. A Silurian placoderm with osteichthyan-like marginal jaw bones. Nature 502, 188–193 (2013).

Lu, J. et al. The oldest actinopterygian highlights the cryptic early history of the hyperdiverse ray-finned fishes. Curr. Biol. 26, 1602–1608 (2016).

Basden, A. M. et al. The most primitive osteichthyan braincase? Nature 403, 185–188 (2000).

Basden, A. M. & Young, G. C. A primitive actinopterygian neurocranium from the Early Devonian of southeastern Australia. J. Vert. Paleont. 21, 754–766 (2001).

Clement, A. M. et al. Neurocranial anatomy of an enigmatic Early Devonian fish sheds light on early osteichthyan evolution. eLife https://doi.org/10.7554/eLife.34349 (2018).

Giles, S., Friedman, M. & Brazeau, M. D. Osteichthyan-like cranial conditions in an Early Devonian stem gnathostome. Nature 520, 82–85 (2015).

Cloutier, R. & Ahlberg, P. E. in Interrelationships of Fishes (eds Stiasnny, M. L. J. et al.) 445–479 (Academic, 1996).

Li, Q. et al. A new Silurian fish close to the common ancestor of modern gnathostomes. Curr. Biol. 31, 3613–3620 (2021).

Schultze, H.-P. & Cumbaa, S. L. in Major Events in Early Vertebrate Evolution: Palaeontology, Phylogeny, Genetics and Development (ed. Ahlberg, P. E.) 315–332 (Taylor & Francis, 2001).

Burrow, C. J., Jones, A. S. & Young, G. C. X-ray microtomography of 410 million-year-old optic capsules from placoderm fishes. Micron 36, 551–557 (2005).

Gardiner, B. G. The relationships of the palaeoniscid fishes, a review based on new specimens of Mimia and Moythomasia from the Upper Devonian of western Australia. Bull. Br. Mus. (Nat. Hist.) Geol. 37, 173–428 (1984).

Long, J. A. New palaeoniscoid fishes from the Late Devonian and Early Carboniferous of Victoria. Mem. Ass. Australas. Palaeontols. 7, 1–64 (1988).

Cui, X.-D., Qiao, T. & Zhu, M. Scale morphology and squamation pattern of Guiyu oneiros provide new insights into early osteichthyan body plan. Sci. Rep. 9, 4411 (2019).

Choo, B. et al. A new osteichthyan from the late Silurian of Yunnan, China. PLoS ONE 12, e0170929 (2017).

Watson, D. M. S. The acanthodian fishes. Philos. Trans. R. Soc. Lond. B Biol. Sci. 228, 49–146 (1937).

Choo, B. Revision of the actinopterygian genus Mimipiscis (=Mimia) from the Upper Devonian Gogo Formation of western Australia and the interrelationships of the early Actinopterygii. Earth Environ. Sci. Trans. R. Soc. 102, 77–104 (2012).

Zhu, M. et al. Fossil fishes from China provide first evidence of dermal pelvic girdles in osteichthyans. PLoS ONE 7, e35103 (2012).