Herrerasauridae

Herrerasaurids Temporal range: Late Triassic, 231.4–225 Ma PreЄ Є O S D C P T J K Pg N
Mounted Herrerasaurus ischigualastensis skeleton cast, at the Field Museum in Chicago
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Dinosauria
Order:	Saurischia
Clade:	Eusaurischia
Suborder:	Theropoda
Family:	†Herrerasauridae Reig, 1963
Genera[1]
†Caseosaurus †Chindesaurus †Herrerasaurus †Sanjuansaurus †Staurikosaurus
Synonyms
Staurikosauridae Galton, 1977

Herrerasaurids are among the oldest known dinosaurs, appearing in the fossil record 231.4 million years ago (Late Triassic). They became extinct by the end of the Triassic period. Herrerasaurids were small-sized (not more than 4 metres (13 ft) long) carnivorous theropods^[2] or basal saurischians.^[3][4] The best known representatives of this group are from South America (Brazil, Argentina), where they were first discovered in the 1960s. A nearly complete skeleton of Herrerasaurus ischigulastensis was discovered in the Ischigualasto Formation in San Juan, Argentina, in 1988. Less complete herrerasaurids have been found in North America, 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 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 clearly theropod in having only three long fingers, with curved claws. Herrerasaurids also have a hinged mandible like all theropods.

Classification

Where herrerasaurids lie on the early dinosaur evolutionary tree is unclear. They are possibly basal theropods or basal saurischians but may in fact predate the saurischian-ornithischian split.^[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,^[2] 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.^[6] 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^[7] from the same Ischigualasto Formation of Argentina as Herrerasaurus, Staurikosaurus from the Santa Maria Formation of southern Brazil,^[8] Chindesaurus from the Upper Petrified Forest (Chinle Formation) of Arizona,^[9] and possibly Caseosaurus from the Dockum Formation of Texas,^[10] although the relationships of these animals are not fully understood, and not all paleontologists agree. Niedźwiedzki et al. (2014) described an European member of the group on the basis of Norian fossils discovered in Poland.^[11] Other possible basal theropods, Alwalkeria from the Late Triassic Maleri Formation of India,^[12] and Teyuwasu, known from very fragmentary remains from the Late Triassic of Brazil, might be related.^[13]

Phylogeny

Novas (1992) defined Herrerasauridae as Herrerasaurus, Staurikosaurus, and their most recent common ancestor.^[14] Sereno (1998) defined the group as the most inclusive clade including H. ischigualastensis but not Passer domesticus.^[15] 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 here follows one proposed analysis by Fernando E. Novas, Martin D. Ezcurra, Sankar Chatterjee and T. S. Kutty in 2011. In this review, Herrerasaurus is a primitive saurischian, but not a theropod.^[16] The second cladogram is based on an analysis by Hans-Dieter Sues, Sterling J. Nesbitt, David S Berman and Amy C. Henrici, in 2011/12. This review indicated Herrerasaurus is a basal theropod.^[17]

Dinosauria	Ornithischia  Saurischia   Herrerasauridae  Herrerasaurus Staurikosaurus Unnamed herrerasaurid  Eusaurischia   Theropoda  Chindesaurus Tawa Eoraptor Neotheropoda Sauropodomorpha

Dinosauria	Ornithischia  Saurischia  Sauropodomorpha  Theropoda   Herrerasauridae  Staurikosaurus Herrerasaurus Chindesaurus Eoraptor Daemonosaurus Tawa Neotheropoda

References

1. Holtz, Thomas R. Jr. (2012) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2011 Appendix.
2. Nesbitt, S. J.; Smith, N. D.; Irmis, R. B.; Turner, A. H.; Downs, A. & Norell, M. A. (2009). "A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs". Science 326 (5959): 1530–1533. doi:10.1126/science.1180350. PMID 20007898.
3. Langer, Max C. (2004). "Basal Saurischia". In Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.). The Dinosauria (2nd ed.). Berkeley: University of California Press. pp. 25–46. ISBN 0-520-24209-2.
4. Langer, M.C; Benton, M.J. (2006). "Early dinosaurs: a phylogenetic study". Journal of Systematic Palaeontology 4 (4): 309–358. doi:10.1017/S1477201906001970.
5. Brinkman, D.B.; Sues, H.-D. (1987). "A staurikosaurid dinosaur from the Upper Triassic Ischigualasto Formation of Argentina and the relationships of the Staurikosauridae". Palaeontology 30: 493–503.
6. Ezcurra, M.D. (2010). "A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: a reassessment of dinosaur origin and phylogeny". Journal of Systematic Palaeontology 8 (3): 371–425. doi:10.1080/14772019.2010.484650.
7. Alcober, Oscar A.; Martinez, Ricardo N. (2010). "A new herrerasaurid (Dinosauria, Saurischia) from the Upper Triassic Ischigualasto Formation of northwestern Argentina". ZooKeys 63 (63): 55–81. doi:10.3897/zookeys.63.550. PMC 3088398. PMID 21594020.
8. Colbert, E.H. (1970). "A saurischian dinosaur from the Triassic of Brazil". American Museum Novitates 2405: 1–39.
9. Long, R.A.; Murry, P.A. (1995). "Late Triassic (Carnian and Norian) Tetrapods from the Southwestern United States". New Mexico Museum of Natural History and Science, Bulletin 4: 1–254.
10. Hunt, A.P.; Lucas, S.G.; Heckert, A.B.; Sullivan, R.M.; Lockley, M.G. (1998). "Late Triassic Dinosaurs from the Western United States". Geobios 31 (4): 511–531. doi:10.1016/S0016-6995(98)80123-X.
11. Grzegorz Niedźwiedzki, Stephen L. Brusatte, Tomasz Sulej and Richard J. Butler (2014). "Basal dinosauriform and theropod dinosaurs from the mid–late Norian (Late Triassic) of Poland: implications for Triassic dinosaur evolution and distribution". Palaeontology. in press. doi:10.1111/pala.12107.
12. Chatterjee, S.; Creisler, B.S. (1994). "Alwalkeria (Theropoda) and Morturneria (Plesiosauria), new names for preoccupied Walkeria Chatterjee, 1987 and Turneria Chatterjee and Small, 1989". Journal of Vertebrate Paleontology 14 (1): 142. doi:10.1080/02724634.1994.10011546.
13. 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.
14. Novas, F. E. (1992). "Phylogenetic relationships of the basal dinosaurs, the Herrerasauridae". Palaeontology 35: 51–62.
15. 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.
16. Fernando E. Novas, Martin D. Ezcurra, Sankar Chatterjee and T. S. Kutty (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. doi:10.1017/S1755691011020093.
17. Hans-Dieter Sues, Sterling J. Nesbitt, David S Berman and Amy C. Henrici (2011). "A late-surviving basal theropod dinosaur from the latest Triassic of North America". Proceedings of the Royal Society B 278 (1723): 3459–64. doi:10.1098/rspb.2011.0410. PMC 3177637. PMID 21490016.