Herrerasauridae is a family of carnivorousdinosaurs, possibly basal to either theropods or even all of saurischians, or even their own branching from Dracohors, separate from Dinosauria altogether. They are among the oldest known dinosaurs, first appearing in the fossil record around 233.23 million years ago (the Carnian stage of the Late Triassic),[2] before becoming extinct by the end of the Carnian stage. Herrerasaurids were relatively small-sized dinosaurs, normally no more than 4 metres (13 ft) long,[3][4] although the holotype specimen of "Frenguellisaurus ischigualastensis" (nowadays considered a synonym of Herrerasaurusischigualastensis) is thought to have reached around 6 meters (20 ft) long. The best known representatives of this group are from South America (Brazil, Argentina), where they were first discovered in the 1930s in relation to Staurikosaurus and 1960s in relation to Herrerasaurus. A nearly complete skeleton of Herrerasaurus ischigualastensis was discovered in the Ischigualasto Formation in San Juan, Argentina, in 1988. Less complete possible herrerasaurids have been found in North America and Africa, and they may have inhabited other continents as well.
Herrerasaurid anatomy is unusual and specialized, and they are not considered to be ancestral to any later dinosaur group. They only superficially resemble theropods and often present a mixture of very primitive and derived traits. The acetabulum is only partly open, and there are only two sacral vertebrae, the lowest number among dinosaurs. The pubic bone has a derived structure, being rotated somewhat posteriorly and folded to create a superficially tetanuran-like terminal expansion, especially prominent in H. ischigulastensis. The hand is primitive in having five metacarpals and the third finger longer than the second, but resembles those of theropods in having only three long fingers, with curved claws. Herrerasaurids also have a hinged mandible, which is also found in theropods. They may be the most basal theropods.
It is not clear where herrerasaurids lie on the early dinosaur evolutionary tree. They are possibly basal theropods or basal saurischians.[5] Early researchers even proposed that they represented an early lineage of sauropodomorphs. Some analyses, such as Nesbitt et al. 2009, have found Herrerasaurus and its relatives in Herrerasauridae to be very basal theropods,[6] while others (such as Ezcurra 2010) have found them to be basal to the clade Eusaurischia, that is, closer to the base of the saurischian tree than either theropods or sauropodomorphs, but not true members of either.[7] The situation is further complicated by uncertainties in correlating the ages of late Triassic beds bearing land animals.[3]
The discovery of the Herrerasaurid Gnathovorax indicates that the family falls outside the Theropoda and Sauropodomorpha in the cladistic analysis undertaken on the genus when it was described, but remains squarely within Saurischia as basal members of the order.[18][19] An unnamed herrerasaurid from the Carnian of Brazil was described and possibly belongs to a new morphotype of relatively large proportions, informally known as the "Big Saturnalia". In the phylogenetic analysis within this study, herrerasaurids are recovered as non-eusaurischiansaurischians.[20]
Fernando Novas (1992) defined Herrerasauridae as Herrerasaurus, Staurikosaurus, and their most recent common ancestor.[21]Paul Sereno (1998) defined the group as the most inclusive clade including H. ischigualastensis but not Passer domesticus.[22] Langer (2004) provided first phylogenetic definition of a higher level taxon, Herrerasauria, as Herrerasaurus but not Liliensternus or Plateosaurus.[3] According to current phylogenetic studies, all of these definitions describe the same clade.
The first cladogram presented follows one proposed analysis by Novas et al. in May 2011. In this review, Herrerasaurus is found to be a basal saurischian, but not a theropod.[23] The second cladogram is based on an analysis by Sues et al. in April 2011. This review classified Herrerasaurus as a basal theropod.[24]
Baron & Williams (2018) found Herrerasauria (including Daemonosaurus, Caseosaurus and Saltopus) outside Dinosauria.[12] A similar result was provided by the phylogenetic analysis by Cau, 2018:[27]
Novas et al., 2021 revised the fossil record of South American early dinosaurs and supported that Herrerasauria is part of Saurischia but diverging earlier than both Sauropodomorpha or Theropoda, and further corroborated with the hypothesis that Chindesaurus, Daemonosaurus and Tawa are members of the clade.[28] In 2024, Andrea Cau reclassified Herrerasauria within Theropoda.[29]
^Griffin, C. T.; Wynd, B. M.; Munyikwa, D.; Broderick, T. J.; Zondo, M.; Tolan, S.; Langer, M. C.; Nesbitt, S.J.; Taruvinga, H. R. (2022). "Africa's oldest dinosaurs reveal early suppression of dinosaur distribution". Nature. 609 (7926): 313–319. Bibcode:2022Natur.609..313G. doi:10.1038/s41586-022-05133-x. PMID36045297. S2CID251977824.
^Garcia, Maurício S.; Müller, Rodrigo T.; Dias-Da-Silva, Sérgio (2019-07-04). "On the taxonomic status of Teyuwasu barberenai Kischlat, 1999 (Archosauria: Dinosauriformes), a challenging taxon from the Upper Triassic of southern Brazil". Zootaxa. 4629 (1): 146–150. doi:10.11646/zootaxa.4629.1.12. ISSN1175-5334. PMID31712541. S2CID198274900.
^Kischlat, E.-E. (1999). "A new dinosaurian "rescued" from the Brazilian Triassic: Teyuwasu barberenai, new taxon". Paleontologia Em Destaque, Boletim Informativo da Sociedade Brasileira de Paleontologia. 14 (26): 58.
^Novas, F. E. (1992). "Phylogenetic relationships of the basal dinosaurs, the Herrerasauridae". Palaeontology. 35: 51–62.
^Sereno, P. C. (1998). "A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 210 (1): 41–83. doi:10.1127/njgpa/210/1998/41.
^Novas, Fernando E.; Ezcurra, Martin D.; Chatterjee, Sankar; Kutty, T. S. (2011). "New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram formations of central India". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 101 (3–4): 333–349. Bibcode:2010EESTR.101..333N. doi:10.1017/S1755691011020093. S2CID128620874.
^Garcia, M. S.; Müller, R. T.; Pretto, F. A.; Da-Rosa, Á. A. S.; Dias-Da-Silva, S. (2021). "Taxonomic and phylogenetic reassessment of a large-bodied dinosaur from the earliest dinosaur-bearing beds (Carnian, Upper Triassic) from southern Brazil". Journal of Systematic Palaeontology. 19: 1–37. Bibcode:2021JSPal..19....1G. doi:10.1080/14772019.2021.1873433. S2CID232313141.