Herrerasauridae
{{Short description|Extinct family of basal saurischian dinosaurs}}
{{Automatic taxobox
| name = Herrerasaurids
| fossil_range = Late Triassic (Carnian), {{fossilrange|233.23|228.91|latest=210}} Possible Late Norian records if Chindesaurus and Caseosaurus are part of the family.
| image = Herrerasaurus_ischigualastensis_DSC_2929.jpg
| image_alt = Herrerasaurus schigualastensis
| image_caption = Skeleton of Herrerasaurus
| taxon = Herrerasauridae
| authority = Benedetto, 1973
| subdivision_ranks = Genera
| subdivision = *{{extinct}}Caseosaurus?
- {{extinct}}Chindesaurus?
- {{extinct}}Daemonosaurus?
- {{extinct}}Gnathovorax
- {{extinct}}Herrerasaurus
- {{extinct}}Sanjuansaurus
- {{extinct}}Staurikosaurus
- {{extinct}}Tawa?
| 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),{{cite journal|last1=Langer|first1=M.C.|last2=Ramezani|first2=J.|last3=Da Rosa|first3=Á.A.S.|year=2018|title=U-Pb age constraints on dinosaur rise from south Brazil|journal=Gondwana Research|volume=X|issue=18|pages=133–140|doi=10.1016/j.gr.2018.01.005|bibcode=2018GondR..57..133L}} before becoming extinct by the end of the Carnian stage. Herrerasaurids were relatively small-sized dinosaurs, normally no more than {{convert|4|m|ft}} long,{{Cite book |title=The Dinosauria |last=Langer |first=Max C. |publisher=University of California Press |year=2004 |isbn=978-0-520-24209-8 |editor-last=Weishampel |editor-first=David B. |editor-link=David B. Weishampel |edition=2nd |location=Berkeley |pages=[https://archive.org/details/dinosauriandedit00weis/page/n43 25]–46 |chapter=Basal Saurischia |oclc=55000644 |ol=3305845M |editor-last2=Dodson |editor-first2=Peter |editor-link2=Peter Dodson |editor-last3=Osmólska |editor-first3=Halszka |editor-link3=Halszka Osmólska|title-link=The Dinosauria }}{{Cite journal |last1=Langer |first1=Max C |last2=Benton |first2=Michael J. |year=2006 |title=Early dinosaurs: a phylogenetic study |journal=Journal of Systematic Palaeontology |volume=4 |issue=4 |pages=309–358 |doi=10.1017/S1477201906001970|bibcode=2006JSPal...4..309L |s2cid=55723635 }} 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
File:Staurikosaurus-mingau.png]]
It is not clear where herrerasaurids lie on the early dinosaur evolutionary tree. They are possibly basal theropods or basal saurischians.{{Cite journal |last1=Brinkman |first1=Donald B. |last2=Sues |first2=Hans-Dieter |year=1987 |title=A staurikosaurid dinosaur from the Upper Triassic Ischigualasto Formation of Argentina and the relationships of the Staurikosauridae |url=http://www.palass.org/publications/palaeontology-journal/archive/30/3/article_pp493-503 |journal=Palaeontology |volume=30 |pages=493–503}} 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,{{Cite journal |last1=Nesbitt |first1=Sterling J. |last2=Smith |first2=Nathan D. |last3=Irmis |first3=Randall B. |last4=Turner |first4=Alan H. |last5=Downs |first5=Alex |last6=Norell |first6=Mark A. |name-list-style=amp |year=2009 |title=A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs |journal=Science |volume=326 |issue=5959 |pages=1530–1533 |doi=10.1126/science.1180350 |pmid=20007898 |bibcode=2009Sci...326.1530N|s2cid=8349110 }} 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.{{Cite journal |last=Ezcurra |first=Martin D. |year=2010 |title=A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: a reassessment of dinosaur origin and phylogeny |journal=Journal of Systematic Palaeontology |volume=8 |issue=3 |pages=371–425 |doi=10.1080/14772019.2010.484650|bibcode=2010JSPal...8..371E |s2cid=129244872 }} The situation is further complicated by uncertainties in correlating the ages of late Triassic beds bearing land animals.
Other proposed members of the clade have included Sanjuansaurus{{Cite journal |last1=Alcober |first1=Oscar A. |last2=Martinez |first2=Ricardo N. |year=2010 |title=A new herrerasaurid (Dinosauria, Saurischia) from the Upper Triassic Ischigualasto Formation of northwestern Argentina |journal=ZooKeys |issue=63 |pages=55–81 |doi=10.3897/zookeys.63.550 |pmc=3088398 |pmid=21594020|doi-access=free |bibcode=2010ZooK...63...55A }} from the Ischigualasto Formation of Argentina, Staurikosaurus and Gnathovorax from the Santa Maria Formation of southern Brazil,{{cite journal|last=Colbert|first=E.H.|year=1970|title=A saurischian dinosaur from the Triassic of Brazil|journal=American Museum Novitates|issue=2405|pages=1–39}} Chindesaurus from the Petrified Forest (Chinle Formation) of Arizona,{{Cite journal |last1=Long |first1=R. A. |last2=Murry |first2=P. A. |year=1995 |title=Late Triassic (Carnian and Norian) Tetrapods from the Southwestern United States |publisher=New Mexico Museum of Natural History and Science|journal= Bulletin |volume=4 |pages=1–254}} and possibly Caseosaurus from the Dockum Formation of Texas,{{Cite journal |last1=Hunt |first1=Adrian P. |last2=Lucas |first2=Spencer G. |last3=Heckert |first3=Andrew B. |last4=Sullivan |first4=Robert M. |last5=Lockley |first5=Martin G. |year=1998 |title=Late Triassic Dinosaurs from the Western United States |journal=Geobios |volume=31 |issue=4 |pages=511–531 |doi=10.1016/S0016-6995(98)80123-X|bibcode=1998Geobi..31..511H }}{{cite journal |author1=Matthew G. Baron |author2=Megan E. Williams |year=2018 |title=A re-evaluation of the enigmatic dinosauriform Caseosaurus crosbyensis from the Late Triassic of Texas, USA and its implications for early dinosaur evolution |journal=Acta Palaeontologica Polonica |volume= 63|doi=10.4202/app.00372.2017 |doi-access=free }} 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.{{Cite journal |last1=Niedźwiedzki |first1=Grzegorz |last2= Brusatte |first2=Stephen L. |last3=Sulej |first3=Tomasz |last4=Butler |first4=Richard J. |year=2014 |title=Basal dinosauriform and theropod dinosaurs from the mid–late Norian (Late Triassic) of Poland: implications for Triassic dinosaur evolution and distribution |journal=Palaeontology |volume= 57|issue=6 |pages=1121–1142 |doi=10.1111/pala.12107|s2cid=129114548 |doi-access=free |bibcode=2014Palgy..57.1121N }} An unnamed herrerasaurid was reported from the Carnian Pebbly Arkose Formation in Zimbabwe by Griffin et al. (2022).{{Cite journal|last1=Griffin |first1=C. T. |last2=Wynd |first2=B. M. |last3=Munyikwa |first3=D. |last4=Broderick |first4=T. J. |last5=Zondo |first5=M. |last6=Tolan |first6=S. |last7=Langer |first7=M. C. |last8=Nesbitt |first8=S.J. |last9=Taruvinga |first9=H. R. |year=2022 |title=Africa's oldest dinosaurs reveal early suppression of dinosaur distribution |journal=Nature |volume=609 |issue=7926 |pages=313–319 |doi=10.1038/s41586-022-05133-x |pmid=36045297 |bibcode=2022Natur.609..313G |s2cid=251977824 }} Other possible basal saurischians include Alwalkeria from the Late Triassic Maleri Formation of southern India,{{Cite journal |last1=Chatterjee |first1=Sankar |last2=Creisler |first2=Benjamin S. |year=1994 |title=Alwalkeria (Theropoda) and Morturneria (Plesiosauria), new names for preoccupied Walkeria Chatterjee, 1987 and Turneria Chatterjee and Small, 1989 |journal=Journal of Vertebrate Paleontology |volume=14 |issue=1 |pages=142 |doi=10.1080/02724634.1994.10011546|bibcode=1994JVPal..14..142C }} and Teyuwasu (recently considered synonym of Staurikosaurus),{{Cite journal|last1=Garcia|first1=Maurício S.|last2=Müller|first2=Rodrigo T.|last3=Dias-Da-Silva|first3=Sérgio|date=2019-07-04|title=On the taxonomic status of Teyuwasu barberenai Kischlat, 1999 (Archosauria: Dinosauriformes), a challenging taxon from the Upper Triassic of southern Brazil|journal=Zootaxa|volume=4629|issue=1|pages=146–150|doi=10.11646/zootaxa.4629.1.12|issn=1175-5334|pmid=31712541|s2cid=198274900}} known from very fragmentary remains from the Late Triassic of Brazil.{{cite journal|last=Kischlat|first=E.-E.|year=1999|title=A new dinosaurian "rescued" from the Brazilian Triassic: Teyuwasu barberenai, new taxon|journal=Paleontologia Em Destaque, Boletim Informativo da Sociedade Brasileira de Paleontologia|volume=14|issue=26|pages=58}}
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.{{Cite web|url=https://blogs.scientificamerican.com/laelaps/stunning-skeleton-reveals-early-carnivorous-dinosaur/|title = Stunning Skeleton Reveals Early Carnivorous Dinosaur}}{{Cite journal|doi = 10.7717/peerj.7963|title = Gnathovorax cabreirai: A new early dinosaur and the origin and initial radiation of predatory dinosaurs|year = 2019|last1 = Pacheco|first1 = Cristian|last2 = Müller|first2 = Rodrigo T.|last3 = Langer|first3 = Max|last4 = Pretto|first4 = Flávio A.|last5 = Kerber|first5 = Leonardo|last6 = Dias Da Silva|first6 = Sérgio|journal = PeerJ|volume = 7|pages = e7963|pmid = 31720108|pmc = 6844243 | doi-access=free }} 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.{{Cite journal|last1=Garcia|first1=Maurício S.|last2=Müller|first2=Rodrigo T.|last3=Pretto|first3=Flávio A.|last4=Da-Rosa|first4=Átila A. S.|last5=Dias-Da-Silva|first5=Sérgio|date=2021-01-02|title=Taxonomic and phylogenetic reassessment of a large-bodied dinosaur from the earliest dinosaur-bearing beds (Carnian, Upper Triassic) from southern Brazil|url=https://www.tandfonline.com/doi/full/10.1080/14772019.2021.1873433|journal=Journal of Systematic Palaeontology|language=en|volume=19|issue=1|pages=1–37|doi=10.1080/14772019.2021.1873433|bibcode=2021JSPal..19....1G |s2cid=232313141|issn=1477-2019}}
=Phylogeny=
Fernando Novas (1992) defined Herrerasauridae as Herrerasaurus, Staurikosaurus, and their most recent common ancestor.{{Cite journal |last=Novas |first=F. E. |year=1992 |title=Phylogenetic relationships of the basal dinosaurs, the Herrerasauridae |journal=Palaeontology |volume=35 |pages=51–62}} Paul Sereno (1998) defined the group as the most inclusive clade including H. ischigualastensis but not Passer domesticus.{{Cite journal |last=Sereno |first=P. C. |year=1998 |title=A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria |journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen |volume=210 |issue=1 |pages=41–83|doi=10.1127/njgpa/210/1998/41 }} Langer (2004) provided first phylogenetic definition of a higher level taxon, Herrerasauria, as Herrerasaurus but not Liliensternus or Plateosaurus. 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.{{Cite journal |last1= Novas |first1=Fernando E. |last2= Ezcurra |first2=Martin D. |last3=Chatterjee |first3=Sankar |last4=Kutty |first4=T. S. |year=2011 |title=New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram formations of central India |journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh |volume=101 |issue=3–4 |pages=333–349 |doi=10.1017/S1755691011020093|bibcode=2010EESTR.101..333N |s2cid=128620874 }} The second cladogram is based on an analysis by Sues et al. in April 2011. This review classified Herrerasaurus as a basal theropod.{{Cite journal |last1= Sues |first1=Hans-Dieter |last2= Nesbitt |first2=Sterling J. |last3= Berman |first3=David S. |last4= Henrici |first4=Amy C. |year=2011 |title=A late-surviving basal theropod dinosaur from the latest Triassic of North America |journal=Proceedings of the Royal Society B |volume=278 |issue=1723 |pages=3459–64 |doi=10.1098/rspb.2011.0410 |pmc=3177637 |pmid=21490016}}
{{clade| style=font-size:80%;line-height:80%
|label1=Dinosauria
|1={{clade
|1=Ornithischia
|label2=Saurischia
|2={{clade
|label1=Herrerasauridae
|1={{clade
|label2=Eusaurischia
|2={{clade
|label2=Theropoda
|2={{clade
|1=Chindesaurus
|2={{clade
|1=Tawa
|2={{clade
|1=Eoraptor
|2=Neotheropoda }} }} }}
}} }} }} }}
{{clade| style=font-size:80%;line-height:80%
|label1=Dinosauria
|1={{clade
|1=Ornithischia
|label2=Saurischia
|2={{clade
|label2=Theropoda
|2={{clade
|label1=Herrerasauridae
|1={{clade
|2={{clade
|2=Chindesaurus }} }}
|2={{clade
|1=Eoraptor
|2={{clade
|2={{clade
|1=Tawa
|2=Neotheropoda }} }} }} }} }} }} }}
A large phylogenetic analysis of early dinosaurs by Matthew Baron, David Norman and Paul Barrett (2017) found Herrerasauridae within the clade Saurischia, as the sister group to Sauropodomorpha. This was the result of the removal of Theropoda from Saurischia and its placement next to Ornithischia within the newly created clade Ornithoscelida.{{cite journal | last1 = Baron | first1 = M.G. | last2 = Norman | first2 = D.B. | last3 = Barrett | first3 = P.M. | year = 2017 | title = A new hypothesis of dinosaur relationships and early dinosaur evolution | journal = Nature | volume = 543 | issue = 7646 | pages = 501–506 | doi = 10.1038/nature21700 | pmid = 28332513 | bibcode = 2017Natur.543..501B | s2cid = 205254710 }}
{{clade| style=font-size:80%;line-height:80%
|label1=Dinosauria
|1={{clade
|label1=Saurischia
|1={{clade
|label1=Herrerasauridae
|1={{clade
|2={{clade
|2={{clade
|2=Chindesaurus
}} }} }}
|2=Sauropodomorpha }}
|label2=Ornithoscelida
|2={{clade
|1=Ornithischia
|label2=Theropoda
|2={{clade
|1=Eoraptor
|2={{clade
|1=Tawa
|2={{clade
|1=Eodromaeus
|2={{clade
|2=Neotheropoda }} }} }} }} }} }} }}
Baron & Williams (2018) found Herrerasauria (including Daemonosaurus, Caseosaurus and Saltopus) outside Dinosauria. A similar result was provided by the phylogenetic analysis by Cau, 2018:{{Cite journal |author=Andrea Cau |year=2018 |title=The assembly of the avian body plan: a 160-million-year long process |journal=Bollettino della Società Paleontologica Italiana |volume=57 |issue=1 |pages=1–25 |doi=10.4435/BSPI.2018.01 |doi-broken-date=2024-11-20 |url=http://paleoitalia.org/media/u/archives/01_Cau_2018_BSPI_571.pdf |access-date=2018-06-19 |archive-date=2018-12-21 |archive-url=https://web.archive.org/web/20181221094620/http://paleoitalia.org/media/u/archives/01_Cau_2018_BSPI_571.pdf |url-status=usurped }}
{{clade| style=font-size:80%; line-height:80%
|label1=Dracohors
|1={{clade
|1=†Silesauridae (including Pisanosaurus)
|2={{clade
|label1=†Herrerasauria
|1={{Clade
|1={{Clade
|1=†Staurikosaurus
|2={{Clade
|1=†Herrerasaurus
|2=†Sanjuansaurus }} }}
|2={{Clade
|1=†Daemonosaurus
|2=†Tawa }}
}}
|label2=Dinosauria
|2={{Clade
|1=†Sauropodomorpha
|2={{Clade
|1=†Eodromaeus
|label2=Ornithoscelida
|2={{Clade
|1=†Ornithischia
|2=Theropoda }} }} }} }} }} }}
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.{{Cite journal|date=2021-10-01|title=Review of the fossil record of early dinosaurs from South America, and its phylogenetic implications|url=https://www.sciencedirect.com/science/article/abs/pii/S0895981121001887|journal=Journal of South American Earth Sciences|language=en|volume=110|pages=103341|doi=10.1016/j.jsames.2021.103341|issn=0895-9811|last1=Novas|first1=Fernando E.|last2=Agnolin|first2=Federico L.|last3=Ezcurra|first3=Martín D.|last4=Temp Müller|first4=Rodrigo|last5=Martinelli|first5=Agustín G.|last6=Langer|first6=Max C.|bibcode=2021JSAES.11003341N }} In 2024, Andrea Cau reclassified Herrerasauria within Theropoda.{{Cite journal|last=Cau |first=A. |title=A Unified Framework for Predatory Dinosaur Macroevolution |year=2024 |journal=Bollettino della Società Paleontologica Italiana |volume=63 |issue=1 |doi=10.4435/BSPI.2024.08 |doi-broken-date=2024-11-20 |url=https://www.researchgate.net/publication/379902868 }}
Timeline of genera
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from: -251.2 till: -247.2 color:earlytriassic text:Olenekian
from: -247.2 till: -242 color:middletriassic text:Anisian
from: -242 till: -237 color:middletriassic text:Ladinian
from: -237 till: -227 color:latetriassic text:Carnian
from: -227 till: -208.5 color:latetriassic text:Norian
from: -208.5 till: -201.3 color:latetriassic text:Rhaetian
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from: -251.9 till: -247.2 color:earlytriassic text:Early Triassic
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color:middletriassic bar:NAM3 from:-231.6 till:-231.4 text:Sanjuansaurus
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from: -201.3 till: -201 color:earlyjurassic
References
{{Wikispecies|Herrerasauridae}}
{{Portal|Dinosaurs}}
{{Reflist}}
{{Avemetatarsalia}}
{{Taxonbar|from=Q134186}}
Category:Prehistoric dinosaur families