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A new avialan theropod from an emerging Jurassic terrestrial fauna


Birds are descended from non-avialan theropod dinosaurs of the Late Jurassic period, but the earliest phase of this evolutionary process remains unclear owing to the exceedingly sparse and spatio-temporally restricted fossil record1,2,3,4,5. Information about the early-diverging species along the avialan line is crucial to understand the evolution of the characteristic bird bauplan, and to reconcile phylogenetic controversies over the origin of birds3,4. Here we describe one of the stratigraphically youngest and geographically southernmost Jurassic avialans, Fujianvenator prodigiosus gen. et sp. nov., from the Tithonian age of China. This specimen exhibits an unusual set of morphological features that are shared with other stem avialans, troodontids and dromaeosaurids, showing the effects of evolutionary mosaicism in deep avialan phylogeny. F. prodigiosus is distinct from all other Mesozoic avialan and non-avialan theropods in having a particularly elongated hindlimb, suggestive of a terrestrial or wading lifestyle—in contrast with other early avialans, which exhibit morphological adaptations to arboreal or aerial environments. During our fieldwork in Zhenghe where F. prodigiosus was found, we discovered a diverse assemblage of vertebrates dominated by aquatic and semi-aquatic species, including teleosts, testudines and choristoderes. Using in situ radioisotopic dating and stratigraphic surveys, we were able to date the fossil-containing horizons in this locality—which we name the Zhenghe Fauna—to 148–150 million years ago. The diversity of the Zhenghe Fauna and its precise chronological framework will provide key insights into terrestrial ecosystems of the Late Jurassic.

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Fig. 1: Morphology, phylogenetic and palaeogeographical position of F. prodigiosus, IVPP V31985.
Fig. 2: Anatomy of F. prodigiosus, IVPP V31985.
Fig. 3: Three-dimensional morphometric space of limb bones and cursoriality of Mesozoic theropods.
Fig. 4: Stratigraphic log and vertebrate fossil assemblage discovered from the Late Jurassic Zhenghe Fauna.

Data availability

The specimen (IVPP V31985) described in this study is archived and available on request from the Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences. Phylogenetic data matrices that support the findings of this research are included as Supplementary Information. The raw data used in morphometric analyses are available at figshare ( The Life Science Identifier for Fujianvenator is

Code availability

The R code used in morphometric analyses is available at figshare (


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We thank W.-Q. Feng, Y. Li, S. Miao, J.-T. Feng and X. Lin for helping with fieldwork; Y. Li for specimen preparation; W. Gao for photography; and S. Miao and J.-T. Feng for laser-stimulated fluorescence imaging. This research was supported by the National Natural Science Foundation of China (42225201 and 42288201); the Key Research Program of Frontier Sciences, CAS (ZDBS-LY-DQC002); the New Cornerstone Science Foundation through the XPLORER PRIZE; Fujian Provincial Department of Natural Resources under the program ‘Investigation of the Geological Relics and Fossil Resources of the Late Mesozoic Basins in Western Fujian’ (GY20220108); and Fujian Provincial Bureau of Geology and Mineral Exploration and Development under the program ‘Study on the Sedimentary Environment of Dinosaur Fossils in Fujian Province’ (MDDR2021-32).

Author information

Authors and Affiliations



M.W. designed the project. M.W. and L.X. supervised the fieldwork. M.W. performed the phylogenetic and comparative analyses. L.X., R.C., M.L., J.T., G.Z., L.W., W.H. and Y. L. collected samples and supervised the radioisotopic dating experiments. M.W. and C.Z. performed the tip-dating analyses. L.D., H.Y., and X.X. studied the vertebrate assemblage and paleoenvironment. M.W. and Z.Z. wrote the manuscript with input from all of the authors.

Corresponding author

Correspondence to Min Wang.

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Nature thanks Steve Brusatte, Daniel Field and Fernando Novas for their contribution to the peer review of this work.

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Extended data figures and tables

Extended Data Fig. 1 Counter slab of holotype of F. prodigiosus, IVPP V31985.

a, Photograph. b, Line drawing. ch, chevron; cv, caudal vertebra; dv, dorsal vertebra; gs, gastralia; lf, left femur; lfi, left fibula; lh, left humerus; li, left ilium; is, ischium; lr, left radius; ls, left scapula; lt, left tibia; lu, left ulna; mI to mV, metatarsal I to V; mcI to III, metacarpal I to III; pt, proximal tarsal; pu, pubis; rh, right humerus; ra, radiale; rc, right coracoid; rf, right femur; rfi, right fibula; ri, right ilium; rs, right scapula; rt, right tibia; st, sternum; un, ulnare; ?sv, possible sacral vertebra; I-1 to I-2, manual phalanx I-1 and I-2; II-1 to II-3, manual phalanx II-1 to II-3; III-1 to III-4, manual phalanx III-1 to III-4. Scale bars, 20 mm.

Extended Data Fig. 2 Additional anatomy of F. prodigiosus, IVPP V31985.

a, Right hand. b, Close-up of the left metacarpals. c, Pelvis. d, Proximal ends of metatarsals. is, ischium; lc, lateral condyle; lct, lateral condyle of tibiotarsus; li, left ilium; mc, medial condyle; mct, medial condyle of tibiotarsus; mcI to III, metacarpal I to III; mtI to mtV, metatarsal I to V; ob, obturator process; pa, pubic apron; pb, pubic boot; pp, posterior distal process; ri, right ilium; I-1 to I-2, manual phalanx I-1 and I-2; II-1, manual phalanx II-1; III-1 to III-4, manual phalanx III-1 to III-4. Scale bars, 10 mm.

Extended Data Fig. 3 Comparison of the pelvic anatomy of F. prodigiosus with that of other selected paravians.

a,b, Photograph (a) and reconstruction (b) of the pelvis of F. prodigiosus. cf, Reconstructed pelvis: avialan Anchiornis (c; modified from refs. 7 and 87), troodontid Sinovenator changii (d; modified from ref. 4), avialan Archaeopteryx (e; modified from ref. 4) and dromaeosaurid Microraptor (f; modified from ref. 4). ob, obturator process; pdp, posterior distal process; ppp, posterior proximal process. The arrowheads denote the constriction at the base of the obturator process. Line drawings are not to scale.

Extended Data Fig. 4 Morphometric analyses of limb-bone length across the Mesozoic theropod phylogeny.

af, Binary plots of the first three principal components (PCs) of the phylogenetic principal components analyses of the six limb segments (a,b), forelimb (c,d), and hindlimb (e,f).

Extended Data Fig. 5 Time-scaled phylogeny showing the position of F. prodigiosus through parsimony analyses.

The phylogeny is the strict consensus resulting from maximum parsimony analyses. The bootstrap and Bremer values are denoted in normal and bold italic fonts, respectively. Thick lines represent the first and last appearance datum of the geological stages or epochs in which a given species was discovered.

Extended Data Fig. 6 Dated phylogeny showing the position of F. prodigiosus through Bayesian tip-dating analysis.

The phylogeny is the majority-rule consensus obtained from the posterior trees of Bayesian tip-dating analysis using the fossilized birth–death model. The error bars at the nodes represent the 95% highest posterior density intervals. The shaded circles at the nodes represent the posterior probability of the corresponding clade.

Extended Data Fig. 7 Comparison of the hindlimb proportions of F. prodigiosus with those of stem and crown groups of theropods.

a,b, Ternary plot of the length proportion of the hindlimb segments (femur, tibia and metatarsal III) of Mesozoic theropods (a), and with the inclusion of crown birds (b).

Extended Data Fig. 8 Additional vertebrate fossils discovered in the Zhenghe Fauna during the 2022 fieldwork.

a,b, Teleostei indet. (Actinopterygii: Neopterygii). ch, Testudines indet. (Reptilia: Pantestudines). ik, Allochoristodera indet. (Reptilia: Choristodera). All photographs were taken in the field shortly after the discovery of the corresponding specimens.

Extended Data Fig. 9 Photograph of the 2022 fieldwork in Zhenghe.

Aerial shot showing the excavation area.

Extended Data Fig. 10 LA-ICP-MS concordial age plots of the fossil-bearing horizons of the Zhenghe Fauna.

a,b, Samples from the tuffite layers deposited just below (c, PM702-1) and above (d, PM701-4) the IVPP V31985-bearing horizon. c,d, Samples from the ignimbrites that underlie (a, Bdx03) and overlie (b, Bdx04) the fossil-bearing sediments.

Supplementary information

Supplementary Information

This file contains Supplementary Figures 1-2, Supplementary Tables 1, 4-7, character description, dataset used in phylogenetic analysis and supplementary references

Reporting Summary

Supplementary Table 2

Supplementary Table 3

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Xu, L., Wang, M., Chen, R. et al. A new avialan theropod from an emerging Jurassic terrestrial fauna. Nature 621, 336–343 (2023).

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