Herrerasaurids Temporal range: Late Triassic (Carnian), Possible Late Norian records if Chindesaurus and Caseosaurus are part of the family.
Skeleton of Herrerasaurus
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Dinosauria (?)
Clade:	Saurischia (?)
Clade:	†Herrerasauria
Family:	†Herrerasauridae Benedetto, 1973
Genera[1]
†Caseosaurus? †Chindesaurus? †Gnathovorax †Herrerasaurus †Sanjuansaurus †Staurikosaurus
Synonyms
Staurikosauridae Galton, 1977 Herrerasauria? Galton, 1985 Herreravia Paul, 1988

Herrerasauridae is a family of carnivorous dinosaurs, 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 Herrerasaurus ischigualastensis) 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.

Classification

Reconstruction of Staurikosaurus

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]

Other proposed members of the clade have included Sanjuansaurus^[8] from the Ischigualasto Formation of Argentina, Staurikosaurus and Gnathovorax from the Santa Maria Formation of southern Brazil,^[9] Chindesaurus from the Petrified Forest (Chinle Formation) of Arizona,^[10] and possibly Caseosaurus from the Dockum Formation of Texas,^[11][12] although the relationships of the North American animals are not fully understood, and not all paleontologists agree. Grzegorz Niedźwiedzki, Stephen L. Brusatte et al. (2014) described a European putative member of the group on the basis of Norian age fossils discovered in Poland.^[13] An unnamed herresaurid was reported from the Carnian Pebbly Arkose Formation in Zimbabwe by Griffin et al. (2022).^[14] Other possible basal saurischians include Alwalkeria from the Late Triassic Maleri Formation of southern India,^[15] and Teyuwasu (recently considered synonym of Staurikosaurus),^[16] known from very fragmentary remains from the Late Triassic of Brazil.^[17]

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-eusaurischian saurischians.^[20]

Timeline of genera

References

Wikispecies has information related to Herrerasauridae.