| Citation: | Yumeng Zhang, Xianren Shan, Xianghong Lin, Zhikun Gai, Philip C. J. Donoghue. A New Dayongaspid Galeaspid from the Silurian of the Lower Yangtze Region: Implications for Biogeography and the Evolution of Key Adaptations in Galeaspids. Journal of Earth Science, 2026, 37(1): 303-316. doi: 10.1007/s12583-026-0504-6 |
Galeaspids are an extinct group of jawless armored fishes that are integral to understanding the origin of anatomical innovations of jawed vertebrates. Resolving the nature of the jawless ancestor of jawed vertebrates requires resolution of the anatomy of the earliest galeaspids, which is currently poorly known. Here we describe a new dayongaspid,
| Chen, Y., Li, Q., Zhou, Z. D., et al., 2024. A New Genus of Galeaspids (Jawless Stem- Gnathostomata) from the Early Silurian Chongqing Lagerstätte, China. Vertebrata PalAsiatica, 62(4): 245–261. https://doi.org/10.19615/j.cnki.2096-9899.240820 |
| Donoghue, P. C. J., Forey, P. L., Aldridge, R. J., 2000. Conodont Affinity and Chordate Phylogeny. Biological Reviews, 75(2): 191–251. https://doi.org/10.1111/j.1469-185x.1999.tb00045.x |
| Gai, Z. K., Donoghue, P. C. J., Zhu, M., et al., 2011. Fossil Jawless Fish from China Foreshadows Early Jawed Vertebrate Anatomy. Nature, 476(7360): 324–327. https://doi.org/10.1038/nature10276 |
| Gai, Z. K., Li, Q., Ferrón, H. G., et al., 2022. Galeaspid Anatomy and the Origin of Vertebrate Paired Appendages. Nature, 609(7929): 959–963. https://doi.org/10.1038/s41586-022-04897-6 |
| Gai, Z. K., Lin, X. H., Shan, X. R., et al., 2023. Postcranial Disparity of Galeaspids and the Evolution of Swimming Speeds in Stem-Gnathostomes. National Science Review, 10(7): nwad050. https://doi.org/10.1093/nsr/nwad050 |
| Gai, Z. K., Lu, L. W., Zhao, W. J., et al., 2018. New Polybranchiaspiform Fishes (Agnatha: Galeaspida) from the Middle Palaeozoic of China and Their Ecomorphological Implications. PLoS One, 13(9): e0202217. https://doi.org/10.1371/journal.pone.0202217 |
| Gai, Z. K., Zhu, M., 2012. The Origin of the Vertebrate Jaw: Intersection between Developmental Biology-Based Model and Fossil Evidence. Chinese Science Bulletin, 57(30): 3819–3828. https://doi.org/10.1007/s11434-012-5372-z |
| Gai, Z. K., Zhu, M., 2017. Evolutionary History of Agnathans and Their Fossil Records in China. Shanghai Scientific and Technical Publishers, Shanghai. 1–314 (in Chinese with English Abstract) |
| Goloboff, P. A., Catalano, S. A., 2016. TNT Version 1.5, Including a Full Implementation of Phylogenetic Morphometrics. Cladistics, 32(3): 221–238. https://doi.org/10.1111/cla.12160 |
| Hughes, C. P., Ingham, J. K., Addison, R., 1975. The Morphology, Classification and Evolution of the Trinucleidae (Trilobita). Philosophical Transactions of the Royal Society of London Series B, Biological Sciences, 272(920): 537–604. https://doi.org/10.1098/rstb.1975.0100 |
|
Janvier, P., 1990. La Structure de l'exosquelette des Galeaspida (Vertebrata). Comptes Rendus de l'Académie des Sciences. Série Ⅱ. Mechanics. Physics. Chemistry. Space Sciences. Earth Sciences, 310(5): 655–659. |
| Janvier, P., Thanh, T. D., Phuong, T. H., et al., 2009. Occurrence of Sanqiaspis, Liu, 1975 (Vertebrata, Galeaspida) in the Lower Devonian of Vietnam, with Remarks on the Anatomy and Systematics of the Sanqiaspididae. Comptes Rendus Palevol, 8(1): 59–65. https://doi.org/10.1016/j.crpv.2008.10.008 |
|
Janvier, P., Tông-Dzuy, T., Ta-Hoa, P., 1993. A New Early Devonian Galeaspid from Bac Thai Province, Vietnam. Palaeontology, 36(2): 297–309. |
| Jiang, W. Y., Zhu, M., Shi, X. D., et al., 2021. Qushiaspis, A New Genus of Gantarostrataspid Fish (Galeaspida, Stem-Gnathostomata) from the Lower Devonian of Yunnan, China. Historical Biology, 33(12): 3714–3722. https://doi.org/10.1080/08912963.2021.1888086 |
| Keating, J. N., Donoghue, P. C. J., 2016. Histology and Affinity of Anaspids, and the Early Evolution of the Vertebrate Dermal Skeleton. Proceedings Biological Sciences, 283(1826): 20152917. https://doi.org/10.1098/rspb.2015.2917 |
| Li, X. T., Zhang, Y. M., Lin, X. H., et al., 2024. New Findings of Changxingaspis (Xiushuiaspidae, Galeaspida) from the Silurian of Tarim Basin and Zhejiang Province, China. Acta Geologica Sinica - English Edition, 98(3): 531–540. https://doi.org/10.1111/1755-6724.15168 |
| Lin, X. H., Zan, C. Y., Gai, Z. K., et al., 2025. Deanaspis, A New Genus of Galeaspida (Jawless Stem Gnathostomata) from the Silurian of Jiangxi, China, and Its Evolutionary Implications. Journal of Systematic Palaeontology, 23(1): 2460479. https://doi.org/10.1080/14772019.2025.2460479 |
| Liu, Y. H., Zhu, M., Gai, Z. K., et al., 2015. Subclass Galeaspida. In: Zhu, M., ed. Palaeovertebrata Sinica, Volume Ⅰ, Fishes, Fascile, Agnathans. Science Press, Beijing. 141–272 (in Chinese) |
| Liu, Y. H., Zhu, M., Lin, X., et al., 2019. A Reappraisal of the Silurian Galeaspids (Stem-Gnathostomata) from Tarim Basin, Xinjiang. Vertebrata PalAsiatica, 57(4): 253–273. https://doi.org/10.19615/j.cnki.1000-3118.190524 (in Chinese with English Abstract) |
| Liu, Y. L., Huang, L. B., Zong, R. W., et al., 2021a. New Material of Galeaspids from the Silurian Llandovery from Wuhan in South China. Earth Science, 46(9): 3307–3320. https://doi.org/10.3799/dqkx.2020.343 (in Chinese with English Abstract) |
| Liu, Y. L., Huang, L. B., Zong, R. W., et al., 2021b. The Oldest Eugaleaspiform (Galeaspida) from the Silurian Fentou Formation (Telychian, Llandovery) of Wuhan, South China. Journal of Systematic Palaeontology, 19(4): 253–264. https://doi.org/10.1080/14772019.2021.1883755 |
| Liu, Y. L., Yin, J. Y., Fan, R. Y., et al., 2024. New Data on Silurian (Llandovery) Sinacanthids from Wuhan, South China and Their Biostratigraphic and Paleobiogeographic Implications. Palaeoworld, 33(5): 1242–1255. https://doi.org/10.1016/j.palwor.2023.11.001 |
|
Maddison, W. P., Maddison, D. R., 2019. Mesquite: A Modular System for Phylogenetic Workflows and Analyses. [2025-12-30]. |
| Meng, X. Y., Gai, Z. K., 2022. Falxcornus, A New Genus of Tridensaspidae (Galeaspida, Stem-Gnathostomata) from the Lower Devonian in Qujing, Yunnan, China. Historical Biology, 34(5): 897–906 |
|
Pan, J., Wang, S. T., 1983. Xiushuiaspidae, A New family of Polybranchiaspformes from Xiushui of Jiangxi Province. Acta Palaeontologica Sinica, 22(5): 505–509. |
| Pan, J., Zeng, X. Y., 1985. Dayongaspidae, A New Family of Polybranchiaspiformes (Agnatha) from Early Silurian of Hunan, China. Vertebrata PalAsiatica, 23(3): 207–213. https://doi.org/10.19615/j.cnki.1000-3118.1985.03.003 (in Chinese with English Abstract) |
| Rong, J. Y., Wang, Y., Tang, P., et al., 2023. Age of the Silurian Tangchiawu, Kangshan and Jukeng Formations (with Marine Red Beds) in Eastern South China. Journal of Stratigraphy, 47(3): 237–267. https://doi.org/10.19839/j.cnki.dcxzz.2023.0015 (in Chinese with English Abstract) |
| Rong, J. Y., Wang, Y., Zhan, R. B., et al., 2019. Silurian Integrative Stratigraphy and Timescale of China. Science China Earth Sciences, 62(1): 89–111. https://doi.org/10.1007/s11430-017-9258-0 |
| Sansom, R. S., Freedman, K., Gabbott, S. E., et al., 2010. Taphonomy and Affinity of an Enigmatic Silurian Vertebrate, Jamoytius kerwoodi White. Palaeontology, 53(6): 1393–1409. https://doi.org/10.1111/j.1475-4983.2010.01019.x |
| Shan, X. R., Gai, Z. K., Lin, X. H., et al., 2022a. The Oldest Eugaleaspiform Fishes from the Silurian Red Beds in Jiangxi, South China and Their Stratigraphic Significance. Journal of Asian Earth Sciences, 229: 105187. https://doi.org/10.1016/j.jseaes.2022.105187 |
| Shan, X. R., Zhao, W. J., Lin, X. H., et al., 2022b. The Correlations of the Lower Red Beds of Early Telychian (Llandovery, Silurian) in China from the Palaeoichthyological Evidence. Journal of Stratigraphy, 46(2): 138–153. https://doi.org/10.19839/j.cnki.dcxzz.2022.0011 (in Chinese with English Abstract) |
| Shan, X. R., Lin, X. H., Zhang, Y. M., et al., 2023a. New Findings of Xiyuichthys (Xiushuiaspidae, Galeaspida) from the Silurian of Jiangxi Province and Tarim Basin. Vertebrata PalAsiatica, 61(4): 245–260. https://doi.org/10.19615/j.cnki.2096-9899.230904 |
| Shan, X. R., Zhao, W. J., Gai, Z. K., 2023b. A New Species of Jiangxialepis (Galeaspida) from the Lower Telychian (Silurian) of Jiangxi and Its Biostratigraphic Significance. Acta Geologica Sinica: English Edition, 97(2): 393–403. https://doi.org/10.1111/1755-6724.15009 |
| Shan, X. R., Zhu, M., Zhao, W. J., et al., 2023c. Fossil Fishes from the Silurian Lower Red Beds in Wuhan, Hubei and Their Biogeographic Significances. Journal of Palaegeography, 25(2): 327–340. https://doi.org/10.7605/gdlxb.2023.02.020 (in Chinese with English Abstract) |
| Shan, X. R., Zhu, M., Zhao, W. J., et al., 2020. A New Genus of Sinogaleaspids (Galeaspida, Stem-Gnathostomata) from the Silurian Period in Jiangxi, China. PeerJ, 8: e9008 https://doi.org/10.7717/peerj.9008 |
| Tarlo, L. H., 1967. Chapter 25 Agnatha. Geological Society, London, Special Publications, 2(1): 629–636. https://doi.org/10.1144/gsl.sp.1967.002.01.44 |
|
Wang, J. Q., Wang, N. Z., Zhang, G. R., et al., 2002. Agnathans from Llandovery (Silurian) of Kalpin, Xinjiang, China. Vertebrata PalAsiatica, 40(4): 245-256. |
|
Wang, N. Z., 1986. Notes on two genera of Middle Silurian Agnatha (Hanyangaspis and Latirostraspis) of China. In: Selected Papers of 13th and 14th Annual Conventions of the Paleontological Society of China, Hefei. Anhui Sci Tech Publ House, Hefei. 49–57. |
| Wang, N. Z., 1991. Two New Silurian Galeaspids (Jawless Craniates) from Zhejiang Province, China, with A Discussion of Galeaspid-Gnathostome Relationships. In: Chang, M. M., Liu, Y. H., Zhang, G. R., eds. Early Vertebrates and Related Problems of Evolutionary Biology, Science Press: Beijing. 41-66 |
| Wang, N. Z., Donoghue, P. C. J., Smith, M. M., et al., 2005. Histology of the Galeaspid Dermoskeleton and Endoskeleton, and the Origin and Early Evolution of the Vertebrate Cranial Endoskeleton. Journal of Vertebrate Paleontology, 25(4), 745–756. https://doi.org/10.1671/0272-4634(2005)025[0745:HOTGDA]2.0.CO;2 |
| Zhang, Y. M., Li, X. T., Shan, X. R., et al., 2024. The First Galeaspid Fish (Stem-Gnathostomata) from the Silurian Xiushan Formation of Hunan Province, China. Historical Biology, 36(9): 1685–1696. https://doi.org/10.1080/08912963.2023.2225083 |
| Zhao, W. J., Wang, S. T., Wang, J. Q., et al., 2009. The Subdivision and Correlation of the Silurian Fish-Bearing Strata and Caledonian Movement in Kalpin and Bachu regions, the Tarim Basin, Xinjiang. Journal of stratigraphy, 33(3): 225-240. https://doi.org/10.3969/j.issn.0253-4959.2009.03.001 (in Chinese with English Abstract) |
| Zhao, W. J., Zhu, M., 2010. Siluro-Devonian Vertebrate Biostratigraphy and Biogeography of China. Palaeoworld, 19(1/2): 4–26. https://doi.org/10.1016/j.palwor.2009.11.007 |
|
Zhu, M., 1992. Two New Eugaleaspids, with A Discussion on Eugaleaspid Phylogeny. Vertebrata PalAsiatica, 30(3): 169–184. |
| Zhu, M., Gai, Z. K., 2007. Phylogenetic Relationships of Galeaspids (Agnatha). Frontiers of Biology in China, 2(2): 151–169. https://doi.org/10.1007/s11515-007-0022-6 |
| Zhu, M., Janvier, P., 1998. The Histological Structure of the Endoskeleton in Galeaspids (Galeaspida, Vertebrata). Journal of Vertebrate Paleontology, 18(3): 650–654. https://doi.org/10.1080/02724634.1998.10011091 |
| Zong, R. W., Liu, Y. L., 2023. On the Age of Silurian Hanyangaspis-Sinacanthus Assemblage in Wuhan, Hubei Province. Journal of Stratigraphy, 47(3): 285–294. https://doi.org/10.19839/j.cnki.dcxzz.2023.0017 (in Chinese with English Abstract) |
| Zong, R. W., Liu, Y. L., Huang, L. B., et al., 2022b. Trilobites from the Silurian "Lower Red Beds" of Wuhan, South China: Stratigraphic and Paleogeographic Implications. Palaeoworld, 31(2): 239–248. https://doi.org/10.1016/j.palwor.2021.05.004 |
| Zong, R. W., Liu, Y. L., Yin, J. Y., et al., 2022a. Discovery of the Silurian Red Beds and Its Geological Significance in Wuhan Area, Eastern Hubei. South China Geology, 38(1): 110–119. https://doi.org/10.3969/j.issn.2097-0013.2022.01.008 (in Chinese with English Abstract) |