Education and training
2009–2013 Bachelor of Science, Geology (with focus in Palaeontology), School of Earth Science and Engineering, Nanjing University.
2013–2018 Doctor of Philosophy, Paleontology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, University of Chinese Academy of Sciences.
2018 Participant, Analytical Paleobiology Workshop, University of Florida. Organizing Chair: Michał Kowalewski.
Career
2016–2018 Smithsonian Fellow (Pre-doctoral), Department of Paleobiology, National Museum of Natural History, Smithsonian Institution.
2018–2019 Smithsonian Fellow (Post-doctoral), Department of Paleobiology, National Museum of Natural History, Smithsonian Institution.
2018–2021 Assistant Professor, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (NIGPAS).
2021–present Associate Professor, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (NIGPAS).
2022–present Specially Appointed Core Research Member, Chinese Academy of Sciences.
2024–present Head of the Chengjiang Palaeontological Station, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (NIGPAS).
My research is focused on the Cambrian explosion, and more specifically on the origin and early evolution of animal phyla and the transformations of early marine animal ecosystems. Much of the fossil evidence on these evolutionary and ecological processes has come from the Cambrian Konservat-Lagerstätten (exceptionally preserved fossil deposits). For years, I have been organizing fossil excavations on the Cambrian fossil deposits of China and collecting soft-bodied fossils from the Chengjiang, Huayuan, and other biotas. For a decade, I have been sorting out the tens of thousands of Burgess Shale-type fossil specimens reposited at the NIGPAS. More recently, I have been compiling and analyzing a global dataset on the Cambrian soft-bodied fossils. This reveals a major transformation in the composition of marine ecosystems at the Sinsk extinction event, which interrupted the Cambrian explosion.
Among the over 30 phyla of the animal kingdom, I am particularly interested in arthropods, the most diverse animal group since the Cambrian period. Extremely abundant arthropod fossils preserved with soft parts have been found from Cambrian Konservat-Lagerstätten, providing direct evidence for understanding the origin and early evolution of arthropods. I have been working on the important and major groups of Cambrian arthropods, with findings on the chimeric Kylinxia, radiodonts (Anomalocaris and its allies), soft-bodied trilobitomorphs, trilobites with appendages, and more. Many fossil arthropod taxa remain to be described, especially those from the recently discovered Huayuan biota. To extract the morpho-anatomical information out of the fossils, I have been using photography under a variety of lighting conditions, microtomography (micro-CT), scanning electron microscope (SEM), energy dispersive spectroscopy (EDS), X-ray fluorescence (XRF), Raman spectroscopy, and other advanced techniques. With new taxa and new data from these endeavors, I have been trying to recover a more reliable and stable phylogenetic framework in the stem part of the arthropod tree of life and to reconstruct the origins of novelties and innovations in the early history of arthropod evolution.
ORCID:https://orcid.org/0000-0002-5728-7896
Google Scholar: https://scholar.google.com/citations?user=YZS5XjYAAAAJ&hl=en
Papers
These include two first-author papers inNature.
1. Zeng H, Liu Q, Zhao FC, Luo C, Wang DZ, Zhu YY, Liu Y, Chen K, Sun ZX, Hong YJ, Miao LY, Hu CL, Sun HJ, Pan B, Zhao JL, Yin ZJ, Li GX, Yang XL, Yang AH, Hu SX, Zhu MY. 2026. A Cambrian soft-bodied biota after the first Phanerozoic mass extinction. Nature, 651(8106): 679–687.
2. Sun ZX, Zeng H, Zhao FC. 2026. Middle Cambrian large bivalved arthropods from Shandong, North China and their paleogeographic implications. Palaeogeography, Palaeoclimatology, Palaeoecology, 687: 113594.
3. Liu Y, Zeng H*, Zhao FC, Zhu YY, Li YM, Yin ZJ, Zhu MY. 2025. A tiny Cambrian stem-mandibulate reveals independent evolution of limb tagmatization and specialization in early euarthropods. Scientific Reports, 15: 19115.
4. Sun ZX, Zhao FC, Zeng H, Erwin DH, Zhu MY. 2025. Episodic body size variations of early Paleozoic trilobites associated with marine redox changes. Science Advances, 11(18): adt7572.
5. Sun ZX, Wang DZ, Zeng H, Zhao FC. 2025. Biostratigraphy and regional correlation of trilobite zones from the Mingxinsi Formation of the Cambrian Series 2, Guizhou Province. Journal of Stratigraphy, 49(2): 142–159.
6. Zhu YY, Zeng H*, Sun ZX, Liu Y, Zhao FC. 2024. Soft-body structures of the trilobite Malongocephalus yunnanensis from the lower Cambrian Hongjingshao Formation in Yunnan, China. Acta Palaeontologica Sinica, 63(2): 207–220.
7. Sun ZX, Zeng H, Zhao FC. 2024. Unique Cambrian trilobite Eodontopleura: an example of strong convergence in trilobite evolution. Historical Biology, 36(2): 253–260.
8. Sun HJ, Zhao FC, Wu RL, Zeng H, Sun ZX. 2024. Spatiotemporal distribution and morphological diversity of the Cambrian Wiwaxia: New insights from South China. Global and Planetary Change, 239(2024): 104507.
9. Wang DZ, Nohejlová M, Sun ZX, Zeng H, Lefebvre B, Yang XL, Zhao FC. 2024. First report of lepidocystid echinoderm in the Cambrian of North China: Evolutionary and Palaeobiogeographic implications. Palaeogeography, Palaeoclimatology, Palaeoecology, 644(2024): 112194.
10. Yang AH, Luo C, Han J, Zhuravlev AY, Reitner J, Sun HJ, Zeng H, Zhao FC, Hu SX. 2024. Niche expansion of archaeocyaths during their palaeogeographic migration: Evidence from the Chengjiang Biota. Palaeogeography, Palaeoclimatology, Palaeoecology, 653(2024): 112419.
11. Zhu YY, Zeng H*, Liu Y, Zhao FC. 2023. New artiopodan euarthropods from the Chengjiang fauna (Cambrian, Stage 3) at Malong, Yunnan, China. Acta Palaeontologica Polonica, 68(3): 427–440.
12. Tian QY, Zhao FC, Zeng H, Zhu MY, Jiang BY. 2023. Response to Comments on “Ultrastructure reveals ancestral vertebrate pharyngeal skeleton in yunnanozoans”. Science, 381(6656): eadf3363.
13. Zeng H, Zhao FC, Zhu MY. 2022. Innovatiocaris, a complete radiodont from the early Cambrian Chengjiang Lagerstätte and its implications for the phylogeny of Radiodonta. Journal of the Geological Society, 180(1): jgs2021-164.
14. Zeng H. 2022. Resolving the problem of arthropod origin. Frontier Science, 16(63): 16–21.
15. Zeng H, Chen XY, Liu Y, Zhu MY, Zhao FC, Yang AH. 2022. A Cambrian soft-bodied conical animal illustrates the origins of lophophorate phyla. bioRxiv, DOI: 10.1101/2022.03.19.485005
16. Tian QY, Zhao FC, Zeng H, Zhu MY, Jiang BY. 2022. Ultrastructure reveals ancestral vertebrate pharyngeal skeleton in yunnanozoans. Science, 377(6602): 218–222.
17. Sun ZX, Zhao FC, Zeng H, Luo C, Van Iten H, Zhu MY. 2022. The middle Cambrian Linyi Lagerstätte from the North China Craton: a new window on Cambrian evolutionary fauna. National Science Review, 9(7): nwac069.
18. Wang DZ, Zhao FC, Zeng H, Li GX, Zhu MY. 2022. A new rhynchonelliform brachiopod Longtancunella with soft-part preservation from the Hongjingshao Formation (Cambrian Stage 3) in Yunnan, South China. Palaeoworld, 31(4): 570–581.
19. Zeng H, Zhao FC, Yin ZJ, Zhu MY. 2021. A new early Cambrian bivalved euarthropod from Yunnan, China and general interspecific morphological and size variations in Cambrian hymenocarines. Palaeoworld, 30(3): 387–397.
20. Sun ZX, Zeng H, Zhao FC. 2021. Digestive structures in Cambrian Miaolingian trilobites from Shandong, North China. Acta Palaeontologica Sinica, 60(1): 166–175.
21. Liu Y, Sun ZX, Zeng Han, Zhao FC. 2021. Brachiopods from the upper part of the Mantou Formation (Cambrian, Miaolingian, Wuliuan) in Weifang, Shandong, North China. Acta Palaeontologica Sinica, 60(1): 2020051.
22. Zeng H, Zhao FC, Niu KC, Zhu MY, Huang DY. 2020. An early Cambrian euarthropod with radiodont-like raptorial appendages. Nature, 588(7836): 101–105.
23. Sun ZX, Zeng H, Zhao FC. 2020. A new middle Cambrian radiodont from North China: Implications for morphological disparity and spatial distribution of hurdiids. Palaeogeography, Palaeoclimatology, Palaeoecology, 558: 109947.
24. Sun ZX, Zeng H, Zhao FC. 2020. Occurrence of the hurdiid radiodont Cambroraster in the middle Cambrian (Wuliuan) Mantou Formation of North China. Journal of Paleontology 94(5): 881–886.
25. Sun ZX, Zeng H, Zhao FC. 2020. First occurrence of the Cambrian arthropod Sidneyia Walcott, 1911 outside of Laurentia. Geological Magazine 157(3): 405–410.
26. Sun ZX, Zeng H, Zhao FC. 2020. A new middle Cambrian trilobite with a specialized cephalon from Shandong Province, North China. Acta Palaeontologica Polonica 65(4): 709–718.
27. Sun HJ, Yin ZJ, Li GX, Zhao FC, Zeng H, Zhu MY. 2020. Periodic shell decollation as an ecology-driven strategy in the early Cambrian Cupitheca. Palaeontology, 63(3): 431–442.
28. Sun HJ, Zhao FC, Steiner M, Li GX, Na L, Pan B, Yin ZJ, Zeng H, Van Iten H, Zhu MY. 2020. Skeletal faunas of the lower Cambrian Yu'anshan Formation, eastern Yunnan, China: Metazoan diversity and community structure during the Cambrian Age 3. Palaeogeography, Palaeoclimatology, Palaeoecology, 542: 109580.
29. Aria C, Zhao FC, Zeng H, Guo J, Zhu MY. 2020. Fossils from South China redefine the ancestral euarthropod body plan. BMC Evolutionary Biology 20(4): 1–17.
30. Luo C, Zhao FC, Zeng H, 2020. The first report of a vauxiid sponge from the Cambrian Chengjiang Biota. Journal of Paleontology, 94(1): 28–33.
31. Zhu MY, Zhao FC, Yin ZJ, Zeng H, Li GX. 2019. The Cambrian explosion: Advances and perspectives from China. Science China: Earth Sciences, 49(10): 1455–1490.
32. Zeng H, Zhao FC, Yin ZJ, Zhu MY. 2018. Morphology of diverse radiodontan head sclerites from the early Cambrian Chengjiang Lagerstätte, south-west China. Journal of Systematic Palaeontology, 16(1): 1–37.
33. Zeng H, Zhao FC, Yin ZJ, Zhu MY. 2018. A new radiodontan oral cone with a unique combination of anatomical features from the early Cambrian Guanshan Lagerstätte, eastern Yunnan, South China. Journal of Paleontology, 92(1): 40–48. (Cover story)
34. Sun HJ, Smith M, Zeng H, Zhao FC, Li GX, Zhu MY. 2018. Hyoliths with pedicles illuminate the origin of the brachiopod body plan. Proceedings of the Royal Society B: Biological Sciences, 285(1887): 20181780.
35. Sun HJ, Zhao FC, Wen RQ, Zeng H, Peng J. 2018. Feeding strategy and locomotion of Cambrian hyolithides. Palaeoworld, 27(3): 334–342.
36. Yin ZJ, Zhao DD, Pan B, Zhao FC, Zeng H, Li GX, Bottjer DJ, Zhu MY. 2018. Early Cambrian animal diapause embryos revealed by X-ray tomography. Geology, 46(5): 387–390.
37. Zeng H, Zhao FC, Yin ZJ, Zhu MY. 2017. Appendages of an early Cambrian metadoxidid trilobite from Yunnan, SW China support mandibulate affinities of trilobites and artiopods. Geological Magazine, 154(6): 1306–1328.
38. Smith EF, Nelson LL, Tweedt SM, Zeng H, Workman JB. 2017. A cosmopolitan late Ediacaran biotic assemblage: new fossils from Nevada and Namibia support a global biostratigraphic link. Proceedings of the Royal Society B: Biological Sciences, 284(1858): 20170934.
39. Zhao FC, Smith MR, Yin ZJ, Zeng H, Li GX, Zhu MY. 2017. Orthrozanclus elongata n. sp. and the significance of sclerite-covered taxa for early trochozoan evolution. Scientific Reports, 7(1): 16232.
40. Zhao FC, Smith MR, Yin ZJ, Zeng H, Hu SX, Li GX, Zhu MY. 2015. First report of Wiwaxia from the Cambrian Chengjiang Lagerstätte. Geological Magazine, 152(2): 378–382.
41. Zeng H, Zhao FC, Yin ZJ, Li GX, Zhu MY. 2014. A Chengjiang-type fossil assemblage from the Hongjingshao Formation (Cambrian Stage 3) at Chenggong, Kunming, Yunnan. Chinese Science Bulletin, 59(25): 3169–3175. (Cover story)
42. Zhao FC, Hu SX, Zeng H, Zhu MY. 2014. A new helmetiid arthropod from the early Cambrian Chengjiang Lagerstätte, southwest China. Journal of Paleontology, 88(2): 367–370.
Books
1. Zhu MY, Zhao FC, Zeng H, Huang DY, Li GX, Luo C. 2025. The Chengjiang fauna: A fossil natural world heritage witnessed the explosive evolution of life on Earth. Pheonix Science Press. 768 pp.
Grants
1. Principal investigator, Equipment Procurement and Maintenance Program, Chinese Academy of Sciences, 2026.
2. Principal investigator, Jiangsu Excellent Young Scientists Fund, Jiangsu Provincial Department of Science and Technology, 2023–2026.
3. Principal investigator, YIPA Membership Program, Youth Innovation Promotion Association, Chinese Academy of Sciences, 2023–2026.
4. Principal investigator, NSFC General Program, National Natural Science Foundation of China, 2023–2026.
5. Core member, Forward-looking Research Project, State Key Laboratory of Palaeobiology and Stratigraphy, 2023–2026.
6. Core member, National Key Research and Development Program, Ministry of Science and Technology, 2023–2026.
7. Principal investigator, Fossil Excavation Project, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 2025.
8. Principal investigator, NSFC Young Scientists Program, National Natural Science Foundation of China, 2020–2022.
9. Principal investigator, Fundamental Research Project, State Key Laboratory of Palaeobiology and Stratigraphy, 2020–2022.
10. Principal investigator, Jiangsu Young Scientists Fund, Jiangsu Provincial Department of Science and Technology, 2019–2022.
Talent program
1. Youth Innovation Promotion Association, Chinese Academy of Sciences (2023–2026)
2. Jiangsu Excellent Young Scientists Fund (2023–2026)
3. Jiangsu High-Level Talent Cultivation Program (2024–present)
Distinctions
1. President's Excellence Award, Chinese Academy of Sciences (2018)
2. Top 10 Annual Advances in Geoscience of China, Geological Society of China (2020)
3. Top 10 Annual Advances in Palaeontology of China, Palaeontological Society of China (2020)
4. Jiangsu Youth Science and Technology Innovation U35 Prize (2021)
Journal editorial board
1. Earth History and Biodiversity (2024–present)
2. Acta Geologica Sinica (2026–present)
Scientific committee
1. Palaeontological Society of China, Committee of Invertebrate Paleontology (2024–present)
2. Palaeontological Society of China, Committee of New Techniques and Methods (2024–present)
3. Palaeontological Society of Jiangsu Province (2024–2029)
4. Geological Society of Hunan Province, Committee of Palaeontology (2025–2029)