Abstract
Dinosaurs and pterosaurs have remarkable diversity and disparity through most of the Mesozoic Era1,2,3. Soon after their origins, these reptiles diversified into a number of long-lived lineages, evolved unprecedented ecologies (for example, flying, large herbivorous forms) and spread across Pangaea4,5. Recent discoveries of dinosaur and pterosaur precursors6,7,8,9,10 demonstrated that these animals were also speciose and widespread, but those precursors have few if any well-preserved skulls, hands and associated skeletons11,12. Here we present a well-preserved partial skeleton (Upper Triassic, Brazil) of the new lagerpetid Venetoraptor gassenae gen. et sp. nov. that offers a more comprehensive look into the skull and ecology of one of these precursors. Its skull has a sharp, raptorial-like beak, preceding that of dinosaurs by around 80 million years, and a large hand with long, trenchant claws that firmly establishes the loss of obligatory quadrupedalism in these precursor lineages. Combining anatomical information of the new species with other dinosaur and pterosaur precursors shows that morphological disparity of precursors resembles that of Triassic pterosaurs and exceeds that of Triassic dinosaurs. Thus, the ‘success’ of pterosaurs and dinosaurs was a result of differential survival among a broader pool of ecomorphological variation. Our results show that the morphological diversity of ornithodirans started to flourish among early-diverging lineages and not only after the origins of dinosaurs and pterosaurs.
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Data availability
All data generated or analysed during this study are included in this published article and its Supplementary Information files. The datasets for phylogenetic, disparity and biogeographic analyses are deposited in figshare (https://doi.org/10.6084/m9.figshare.23508870, https://doi.org/10.6084/m9.figshare.23508858 and https://doi.org/10.6084/m9.figshare.23508849). This publication and associated nomenclatural acts have been registered in ZooBank as urn:lsid:zoobank.org:pub:922F8C41-84E3-46D7-9325-74BCC0D7A7B4.
Code availability
The custom codes used for disparity and biogeographic analyses are deposited in figshare (https://doi.org/10.6084/m9.figshare.23508858 and https://doi.org/10.6084/m9.figshare.23508849).
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Acknowledgements
This study used computational resources from Universidad Nacional de Córdoba (https://ccad.unc.edu.ar/), which are part of SNCAD – MinCyT, Argentina. We thank the Willi Hennig Society for supporting the free use of TNT software. We thank C. Fantini and M. Fernandes for reconstructions of V. gassenae. This study was supported by the Fundação de Amparo à Pesquisa do Estado do Rio Grande do Sul (FAPERGS no. 21/2551-0000680-3 to R.T.M); Conselho Nacional de Desenvolvimento Cientıfico e Tecnológico (CNPq nos. 404095/2021-6 and 303034/2022-0 to R.T.M., 406902/2022-4 to R.T.M., M.B.S. and A.W.A.K. and 313461/2018-0 and 406779/2021-0 to A.W.A.K.); Sepkoski Grant of the Paleontological Society (to M.D.E.); Agencia Nacional de Promoción Científica y Técnica (PICT nos. 2018-01186 to M.D.E. and 2018-01390 to F.L.A.); Coordenação de Aperfeiçoamento de Pessoal de Nível Superior scholarship (CAPES no. 88887.608076/2021-00 to M.S.G.); and Fundação Carlos Chagas Filho de Amparo à Pesquisa do Rio de Janeiro (FAPERJ no. E-26/201.095/2022).
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R.T.M., M.D.E. and S.J.N. designed the project. R.T.M. conducted specimen preparation and curation. R.T.M., M.D.E., M.S.G., F.L.A. and S.J.N. scored phylogenetic matrices. M.D.E. conducted phylogenetic, disparity and biogeographical analyses. R.T.M., M.D.E., M.S.G., F.L.A., F.E.N. and S.J.N. assembled the differential diagnosis. R.T.M. and M.S.G. created the figures with inputs from M.D.E., F.L.A., M.R.S., F.E.N., M.B.S., A.W.A.K. and S.J.N. All authors described the material, collected data and contributed to writing, discussion and conclusion.
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Extended data figures and tables
Extended Data Fig. 1 Provenance of Venetoraptor gassenae gen. et sp. nov. (CAPPA/UFSM 0356).
a, General view of the Buriol/Pivetta complex. b, Buriol site. The map was adapted from The Paleobiology Database (CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/).
Extended Data Fig. 2 Life and skull reconstruction of Venetoraptor gassenae gen. et sp. nov. (CAPPA/UFSM 0356).
a, Skull reconstruction in left lateral view according to the preserved bones of the holotype. b, Head reconstruction in left lateral view. c, Head reconstruction in left anterolateral view. Life reconstruction by Caio Fantini.
Extended Data Fig. 3 Additional bone elements of Venetoraptor gassenae (CAPPA/UFSM 0356).
a, Left orbitotemporal region of the skull in dorsal view. b, Braincase in dorsal view. c, Left manual digit III in medial view. Right femur in (d). anterior, (e) posteromedial, (f) proximal, (g) anterolateral, and (h) posterior views. 4t, fourth trochanter; alr, anterolateral ridge; at, anterior trochanter; clp, collateral ligament pit; crtf, crista tibiofibularis; ef, extensor fossa; f, frontal; fm, foramen magnum; ft, flexor tubercle; lc, lateral condyle; ltf, laterotemporal fenestra; mc, medial condyle; mc III, metacarpal III; o, orbit; p, parietal; pf, popliteal fossa; pmt, posteromedial tuber; po, postorbital; pof, postfrontal; pp, paroccipital process; prf, prefrontal; q, quadrate; so, supraoccipital; sq, squamosal; stf, supratemporal fenestra; ts, trochanteric shelf; ve, ventral emargination. Scale bars: 1 cm.
Extended Data Fig. 4 Additional bone elements of Venetoraptor gassenae (CAPPA/UFSM 0356).
a, Right manus in lateral view. Proximal portion of the right fibula in (b) lateral, (c) anterior, and (d) proximal views. e, Right metatarsal IV in anterior view. f, Right metatarsal III in anterior view. g, Digit III of the right pes in medial view. ef, extensor fossa; mc, metacarpal; ph, phalanx; uph, ungual phalanx. Scale bars: 1 cm.
Extended Data Fig. 5 Reduced strict consensus tree depicting the phylogenetic position of Venetoraptor gassenae gen. et sp. nov.
Absolute (left) and GC (group present/contradicted) (right) bootstrap frequencies and Bremer support values are shown above each branch. The silesaurid silhouette has been adapted from Müller & Garcia43 (CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/).
Extended Data Fig. 6 Majority rule tree recovered from the unconstrained Bayesian phylogenetic analysis depicting the position of Venetoraptor gassenae gen. et sp. nov.
Numbers at nodes indicate posterior probabilities and dotted red vertical lines indicate the boundaries between the Permian and Triassic, Triassic and Jurassic, and Cretaceus and Paleogene geological periods. Life reconstruction of Venetoraptor gassenae gen. et sp. nov. by Caio Fantini.
Extended Data Fig. 7 Ancestral geographic areas reconstructed by the Dispersal-Extinction-Cladogenesis model in the eucrocopodan region of the tree.
Abbreviations: AR, Argentina; BR, Brazil, Uruguay, Namibia; CH, China, Thailand, Kyrgyzstan; eNA, eastern USA, Eastern Canada, Morocco and Algeria; EU, Europe, Russia and Greenland; INT, India, Tanzania, Zambia, Madagascar, Israel and Saudi Arabia; sAF, South Africa, Lesotho, Zimbabwe; wNA, western USA, British Columbia, Mexico and Venezuela. Life reconstruction of Venetoraptor gassenae gen. et sp. nov. by Caio Fantini.
Extended Data Fig. 8 Results of the morphological disparity analyses.
Bivariate plots using: a, Whole skeleton; b, Skull; c, Rostrum; d, Forelimb; e, Anterior zeugopodium and autopodium; f, Hindlimb. Sum of variances: g, Whole skeleton; h, Skull; i, Rostrum; j, Forelimb; k, Anterior zeugopodium and autopodium; l, Hindlimb. In the Sum of Variances the dots are means and the 95% confidence intervals were generated using the two tails of values recovered from 9,999 bootstrap technical replicates of a dataset composed of n = 11 (Whole skeleton, Forelimb), n = 12 (Skull, Rostrum), n = 7 (Anterior zeugopodium and autopodium) and n = 17 (Hindlimb) species of Ornithodiran precursors, n = 18 (Whole skeleton, Skull, Rostrum), n = 16 (Forelimb), n = 14 (Anterior zeugopodium and autopodium) and n = 19 (Hindlimb) species of Dinosauria, and n = 10 (Whole skeleton, Skull, Rostrum), n = 7 (Forelimb, Anterior zeugopodium and autopodium) and n = 6 (Hindlimb) species of Pterosauria. The pterosauromorph silhouette has been adapted from Kellner et al. 10 (CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/).
Extended Data Fig. 9 Additional results of the morphological disparity analyses.
Bivariate plots using: a, Whole skeleton; b, Skull; c, Rostrum; d, Forelimb; e, Anterior zeugopodium and autopodium; f, Hindlimb. Sum of variances: g, Whole skeleton; h, Skull; i, Rostrum; j, Forelimb; k, Anterior zeugopodium and autopodium; l, Hindlimb. In the Sum of Variances the dots are means and the 95% confidence intervals were generated using the two tails of values recovered from 9,999 bootstrap technical replicates of a dataset composed of n = 5 (Whole skeleton, Skull), n = 4 (Rostrum, Forelimb), n = 3 (Anterior zeugopodium and autopodium) and n = 9 (Hindlimb) species of Lagerpetidae, n = 5 (Whole skeleton), n = 6 (Skull, Forelimb), n = 7 (Rostrum, Hindlimb) and n = 3 (Anterior zeugopodium and autopodium) species of Silesauridae, n = 18 (Whole skeleton, Skull, Rostrum), n = 16 (Forelimb), n = 14 (Anterior zeugopodium and autopodium) and n = 19 (Hindlimb) species of Dinosauria, and n = 10 (Whole skeleton, Skull, Rostrum), n = 7 (Forelimb, Anterior zeugopodium and autopodium) and n = 6 (Hindlimb) species of Pterosauria. The pterosauromorph silhouette has been adapted from Kellner et al. 10 (CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/). The silesaurid silhouette has been adapted from Müller & Garcia43 (CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/).
Extended Data Fig. 10 Evolutionary tree of archosauromorphs (above) and dinosaurs (below) depicting distinct episodes of edentulism.
a, Langobadisaurus pandolfii. b, Trilophosaurus buettneri. c, Teyumbaita sulcognathus. d, Aetosauroides scagliai. e, Effigia okeeffeae. f, Venetoraptor gassenae gen. et sp. nov. g, Seazzadactylus venieri. h, Asilisaurus kongwe. i, Heterodontosaurus tucki. j, Stegosaurus stenops. k, Protoceratops andrewsi. l, Iguanodon bernissartensis. m, Limusaurus inextricabilis. n, Deinocheirus mirificus. o, Erlikosaurus andrewsi. p, Citipati osmolskae. q, Gobipteryx minuta. r, Gallus gallus.
Supplementary information
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This file contains information on fossil provenance, a differential diagnosis for the new taxon, details of skeletal reconstruction, detailed descriptions of the methods and Supplementary Table 1.
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Müller, R.T., Ezcurra, M.D., Garcia, M.S. et al. New reptile shows dinosaurs and pterosaurs evolved among diverse precursors. Nature 620, 589–594 (2023). https://doi.org/10.1038/s41586-023-06359-z
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DOI: https://doi.org/10.1038/s41586-023-06359-z
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