Nesbitt, S. J. et al. Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira. Nature 464, 95–98 (2010).
Martínez, R. N. et al. A basal dinosaur from the dawn of the dinosaur era in southwestern Pangaea. Science 331, 206–210 (2011).
Nesbitt, S. J., Barrett, P. M., Werning, S., Sidor, C. A. & Charig, A. J. The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania. Biol. Lett. 9, 20120949 (2013).
Heath, J. A., Cooper, N., Upchurch, P. & Mannion, P. D. Accounting for sampling heterogeneity suggests a low paleolatitude origin for dinosaurs. Curr. Biol. 35, 941–953 (2025).
Brusatte, S. L. et al. The origin and early radiation of dinosaurs. Earth Sci. Rev. 101, 68–100 (2010).
Langer, M. C., Ezcurra, M. D., Bittencourt, J. S. & Novas, F. E. The origin and early evolution of dinosaurs. Biol. Rev. Camb. Philos. Soc. 85, 55–110 (2010).
Langer, M. C. L., Ramezani, J. & Da Rosa, A. A. S. U-Pb age constraints on dinosaur rise from south Brazil. Gondw. Res. 57, 133–140 (2018).
Griffin, C. T. et al. Africa’s oldest dinosaurs reveal early suppression of dinosaur distribution. Nature 609, 313–319 (2022).
Lovelace, D. M. et al. Rethinking dinosaur origins: oldest known equatorial dinosaur-bearing assemblage (mid-late Carnian Popo Agie FM, Wyoming, USA). Zool. J. Linn. Soc. 203, zlae153 (2025).
Bakker, R. Anatomical and ecological evidence of endothermy in dinosaurs. Nature 238, 81–85 (1972).
Charig, A. J. Competition between therapsids and archosaurs during the Triassic period: a review and synthesis of current theories. Symp. Zool. Soc. Lond. 52, 597–628 (1984).
Benton, M. J. Late Triassic extinctions and the origin of the dinosaurs. Science 260, 769–770 (1993).
Benton, M. J. Dinosaur success in the Triassic: a noncompetitive ecological model. Quart. Rev. Biol. 58, 29 (1983).
Irmis, R. B. et al. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science 317, 358–361 (2007).
Ezcurra, M. D. A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: a reassessment of dinosaur origin and phylogeny. Syst. Palaeontol. 8, 371–424 (2010).
Bernardi, M., Gianolla, P., Petti, F. M., Mietto, P. & Benton, M. J. Dinosaur diversification linked with the Carnian Pluvial Episode. Nat. Commun. 9, 1499 (2018).
Benton, M. J., Bernardi, M. & Kinsella, C. The Carnian Pluvial Episode and the origin of dinosaurs. J. Geol. Soc. 175, 1019–1026 (2018).
Schoepfer, S. D., Algeo, T. J., van de Schootbrugge, B. & Whiteside, J. H. The Triassic–Jurassic transition—a review of environmental change at the dawn of modern life. Earth Sci. Rev. 232, 104099 (2022).
Qvarnström, M. et al. Digestive contents and food webs record the advent of dinosaur supremacy. Nature 636, 397–403 (2024).
Corecco, L., Kohn, M. J. & Schultz, C. L. Triassic climate and the rise of the dinosaur empire in South America. J. S. Am. Earth Sci. 142, 104977 (2024).
Novas, F. E., Ezcurra, M. D., Chatterjee, S. & Kutty, T. S. New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram formations of central India. Earth Environ. Sci. Trans. R. Soc. Edinb. 101, 333–349 (2010).
Martínez, R. N. et al. Vertebrate succession in the Ischigualasto Formation. J. Vert. Paleontol. 32, 10–30 (2013).
Sereno, P. C. & Novas, F. E. The complete skull and skeleton of an early dinosaur. Science 258, 1137–1140 (1992).
Sereno, P. C., Forster, C. A., Rogers, R. R. & Monetta, A. M. Primitive dinosaur skeleton from Argentina and the early evolution of Dinosauria. Nature 361, 64–66 (1993).
Alcober, O. & Martínez, R. N. A new herrerasaurid (Dinosauria, Saurischia) from the Upper Triassic Ischigualasto Formation of northwestern Argentina. ZooKeys 63, 55–81 (2010).
Martínez, R. N. & Alcober, O. A. A basal sauropodomorph (Dinosauria: Saurischia) from the Ischigualasto Formation (Triassic, Carnian) and the early evolution of Sauropodomorpha. PLoS ONE 4, e4397 (2009).
Sereno, P. C., Martínez, R. N. & Alcober, O. A. Osteology of Eoraptor lunensis (Dinosauria, Sauropodomorpha). J. Vert. Paleontol. 32, 83–179 (2013).
Rogers, R. R. et al. The Ischigualasto tetrapod assemblage (Late Triassic, Argentina) and 40Ar/39Ar dating of dinosaur origins. Science 260, 794–797 (1993).
Desojo, J. B. et al. The Late Triassic Ischigualasto Formation at Cerro Las Lajas (La Rioja, Argentina): fossil tetrapods, high-resolution chronostratigraphy, and faunal correlations. Sci. Rep. 10, 12782 (2020).
Colombi, C. et al. A high-precision U-Pb zircon age constraints the timing of the faunistic and palynofloristic events of the Carnian Ischigualasto Formation, San Juan, Argentina. J. S. Am. Earth Sci. 111, 103433 (2021).
Tabor, N. et al. in Paleoenvironmental Record and Applications of Calcretes and Palustrine Carbonates Vol. 416 (eds Alonso-Zarza, A. & Tanner, L.) 17–41 (Geological Society of America, 2006).
Currie, B., Colombi, C., Tabor, N., Shipman, T. & Montañez, I. Stratigraphy and architecture of the Upper Triassic Ischigualasto Formation, Ischigualasto Provincial Park, San Juan, Argentina. J. S. Am. Earth Sci. 27, 74–87 (2009).
Colombi, C. E., Limarino, C. O. & Alcober, O. A. Allogenic controls on the fluvial architecture and fossil preservation of the Upper Triassic Ischigualasto Formation, NW Argentina. Sediment. Geol. 362, 1–16 (2017).
Mancuso, A. C. et al. Paleoenvironmental and biotic changes in the Late Triassic of Argentina: testing hypotheses of abiotic forcing at the basin scale. Front. Earth Sci. 10, 883788 (2022).
Colbert, E. H. The Triassic dinosaur Coelophysis. Mus. N. Ariz. Bull. 57, 1–160 (1989).
Raath, M. A. The Anatomy of the Triassic Theropod Syntarsus rhodesiensis (Saurischia: Podokesauridae) and a Consideration of its Biology (Rhodes University, 1977).
Langer, M. C., McPhee, B. W., Marsola, J. C. A., Roberto-da-Silva, L. & Cabreira, S. F. Anatomy of the dinosaur Pampadromaeus barberenai (Saurischia—Sauropodomorpha) from the Late Triassic Santa Maria Formation of southern Brazil. PLoS ONE 14, e0212543 (2019).
Nesbitt, S. J. et al. A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs. Science 326, 1530–1533 (2009).
Sereno, P. C. The pectoral girdle and forelimb of the basal theropod Herrerasaurus ischigualastensis. J. Vert. Paleontol. 13, 425–450 (1994).
Rowe, T. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. J. Vert. Paleontol. 9, 125–136 (1989).
Nesbitt, S. J. & Ezcurra, M. D. The early fossil record of dinosaurs in North America: a new neotheropod from the base of the Upper Triassic Dockum Group of Texas. Act. Palaeontol. Pol. 60, 513–526 (2015).
Mancuso, A. C., Benavente, C. A., Irmis, R. B. & Mundil, R. Evidence for the Carnian Pluvial Episode in Gondwana: new multiproxy climate records and their bearing on early dinosaur diversification. Gond. Res. 86, 104–125 (2020).
Dal Corso, J. et al. Extinction and dawn of the modern world in the Carnian (Late Triassic). Sci. Adv. 6, eaba0099 (2020).
Colombi, C. E., Rogers, R. R. & Alcober, O. A. Vertebrate taphonomy of the Ischigualasto Formation. J. Vert. Paleontol. 32, 31–50 (2012).
Valdes, P. J., Scotese, C. R. & Lunt, D. J. Deep ocean temperatures through time. Clim. Past 17, 1483–1506 (2021).
Santi Malnis, P., Colombi, C. E., Rothis, L. M. & Alcober, O. Fluvial architecture and paleoenvironmental evolution of the Los Colorados Formation (Norian): postrift stage of the Ischigualasto–Villa Unión Basin, NW Argentina. J. Sediment. Res. 90, 1436–1462 (2020).
Apaldetti, C., Martínez, R. N., Cerda, I. A., Pol, D. & Alcober, O. An early trend towards gigantism in Triassic sauropodomorph dinosaurs. Nat. Ecol. Evol. 2, 1227–1232 (2018).
Barrett, P. M., Chapelle, K. E. J., Staunton, C. K., Botha, J. & Choiniere, J. N. Postcranial osteology of the neotype specimen of Massospondylus carinatus Owen, 1854 (Dinosauria: Sauropodomorpha) from the upper Elliot formation of South Africa. Palaeontol. Afr. 53, 114–178 (2019).
Waskow, K. & Sander, P. M. Growth record and histological variation in the dorsal ribs of Camarasaurus sp. (Sauropoda). J. Vert. Paleontol. 4, 852–869 (2014).
Waskow, K. & Mateus, O. Dorsal rib histology of dinosaurs and a crocodylomorph from western Portugal: skeletochronological implications on age determination and life history traits. C. R. Palevol 16, 425–439 (2017).
Cerda, I. A. et al. A basic guide for sampling and preparation of extant and fossil bones for histological studies. Publ. Electr. Asoc. Paleontol. Arg. 20, 15–28 (2020).
Francillon-Vieillot, H. et al. in Skeletal Biomineralization: Patterns, Processes and Evolutionary Trends (ed. Carter, J. G.) 471–548 (Van Nestrand Reinhold, 1990).
de Buffrénil, V. & Quilhac, A. in Comparative Skeletal Histology and Palaeohistology (eds de Buffrénil, V. et al.) 147–190 (CRC Press, 2021).
Ezcurra, M. D., Marke, D., Walsh, S. A. & Brusatte, S. L. A revision of the ‘coelophysoid-grade’ theropod specimen from the Lower Jurassic of the Isle of Skye (Scotland). Scott. J. Geol. 59, sjg2023–012 (2023).
Garcia, M. S., Cabreira, S. F., da Silva, L. R., Pretto, F. A. & Müller, R. T. A saurischian (Archosauria, Dinosauria) ilium from the Upper Triassic of southern Brazil and the rise of Herrerasauria. Anat. Rec. 307, 1011–1024 (2024).
Norman, D. B., Baron, M. G., Garcia, M. S. & Müller, R. T. Taxonomic, palaeobiological and evolutionary implications of a phylogenetic hypothesis for Ornithischia (Archosauria: Dinosauria). Zool. J. Linn. Soc. 196, 1273–1309 (2022).
Goloboff, P. A. & Morales, M. E. TNT version 1.6, with a graphical interface for MacOS and Linux, including new routines in parallel. Cladistics 39, 144–153 (2023).
Goloboff, P. A., Torres, A. & Arias, J. S. Weighted parsimony outperforms other methods of phylogenetic inference under models appropriate for morphology. Cladistics 34, 407–437 (2018).
Ezcurra, M. D. Exploring the effects of weighting against homoplasy in genealogies of palaeontological phylogenetic matrices. Cladistics 40, 242–281 (2024).
Spiekman, S. N. F., Ezcurra, M. D., Butler, R. J., Fraser, N. C. & Maidment, S. C. R. Pendraig milnerae, a new small-sized coelophysoid theropod from the Late Triassic of Wales. R. Soc. Open Sci. 8, 210915 (2021).
Martínez, R. N. et al. Supplementary Information of A Carnian theropod with unexpectedly derived features during the first dinosaur radiation. figshare https://doi.org/10.6084/m9.figshare.29539004 (2025).
Ronquist, F., van der Mark, P. & Huelsenbeck, J. P. in The Phylogenetic Handbook: a Practical Approach to Phylogenetic Analysis and Hypothesis Testing (eds Lemey, P. et al.) 210−266 (Cambridge Univ. Press, 2009).
Ezcurra, M. D. & Butler, R. J. The rise of the ruling reptiles and ecosystem recovery from the Permo-Triassic mass extinction. Proc. R. Soc. B 285, 20180361 (2018).
Ezcurra, M. D., Scheyer, T. M. & Butler, R. J. The origin and early evolution of Sauria: reassessing the Permian saurian fossil record and the timing of the crocodile-lizard divergence. PLoS ONE 9, e89165 (2014).
Rambaut, A., Drummond, A. J., Xie, D., Baele, G. & Suchard, M. A. Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Syst. Biol. 67, 901–904 (2018).
Benson, R. B. et al. Rates of dinosaur body mass evolution indicate 170 million years of sustained ecological innovation on the avian stem lineage. PLoS Biol. 12, e1001853 (2014).
Apaldetti, C., Pol, D., Ezcurra, M. D. & Martínez, R. N. Sauropodomorph evolution across the Triassic–Jurassic boundary: body size, locomotion, and their influence on morphological disparity. Sci. Rep. 11, 22534 (2021).
Ezcurra, M. D. A new early coelophysoid neotheropod from the Late Triassic of northwestern Argentina. Ameghiniana 54, 506–538 (2017).
Campione, N. E. MASSTIMATE: body mass estimation equations for vertebrates. R package version 2.0-1 (CRAN, 2020); https://CRAN.R-project.org/package=MASSTIMATE
Campione, N. E., Evans, D. C., Brown, C. M. & Carrano, M. T. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol. Evol. 5, 913–923 (2014).
Revell, L. J. phytools 2.0: an updated R ecosystem for phylogenetic comparative methods (and other things). PeerJ 12, e16505 (2024).
Bapst, D. W. paleotree: an R package for paleontological and phylogenetic analyses of evolution. Methods Ecol. Evol. 3, 803–807 (2012).
Laurin, M. The evolution of body size, Cope’s rule and the origin of amniotes. Syst. Biol. 53, 594–622 (2004).
Oksanen, J. vegan: community ecology package. R package version 2.6-4 (CRAN, 2022); https://CRAN.R-project.org/package=vegan
Kent, D. V., Santi Malnis, P., Colombi, C. E., Alcober, O. A. & Mart¡nez, R. N. Age constraints on the dispersal of dinosaurs in the Late Triassic from magnetochronology of the Los Colorados Formation (Argentina). Proc. Natl Acad. Sci. USA 111, 7958–7963 (2014).
Marsicano, C., Irmis, R., Mancuso, A., Mundil, R. & Chemale, F. The precise temporal calibration of dinosaur origins. Proc.Natl Acad. Sci. USA 113, 509–513 (2016).
Ezcurra, M. D. et al. Deep faunistic turnovers preceded the rise of dinosaurs in southwestern Pangaea. Nat. Ecol. Evol. 1, 1477–1483 (2017).
Aguirre Palafox, L. E. et al. U-Pb geochronology of paleosol carbonate cements by LA-ICP-MS: A proof of concept and strategy for dating the terrestrial record. Geochem. Geophys. Geosyst. 25, e2024GC011488 (2024).