Gojirasaurus

Gojirasaurus quayi was described and named by Kenneth Carpenter in 1997 based on a partial skeleton, the holotype specimen UCM 47221, from Quay County, New Mexico. The holotype is an assortment of various postcranial bones, including a right scapula, right pubis, left tibia, left metatarsal V, four vertebral centra, a neural arch, and fragments of ribs and gastralia. In addition, a single large serrated tooth is associated with the postcranial material. The holotype is housed in the collections of the University of Colorado Museum of Natural History, in Boulder, Colorado.

The specimen hails from purplish-grey mudstones of the Bull Canyon Formation (sometimes called the Cooper Canyon Formation), a major fossiliferous component of the Dockum Group in eastern New Mexico. In particular, it was found at a site in the vicinity of Revuelto Creek, the type locality for the Revueltian "faunachron".{{Cite journal |last=Lucas |first=Spencer G |date=1998-11-01 |title=Global Triassic tetrapod biostratigraphy and biochronology |url=http://www.sciencedirect.com/science/article/pii/S0031018298001175 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=143 |issue=4 |pages=347–384 |bibcode=1998PPP...143..347L |doi=10.1016/S0031-0182(98)00117-5 |issn=0031-0182}} The Revueltian is a biostratigraphic unit roughly equivalent to the mid-late Norian stage of the Triassic Period, approximately 215–207 million years ago.{{Citation |last1=Martz |first1=J. W. |title=Revised Formulation of the Late Triassic Land Vertebrate "Faunachrons" of Western North America: Recommendations for Codifying Nascent Systems of Vertebrate Biochronology |date=2017-01-01 |work=Terrestrial Depositional Systems |pages=39–125 |editor-last=Zeigler |editor-first=Kate E. |url=https://www.sciencedirect.com/science/article/pii/B9780128032435000029 |access-date=2022-08-28 |publisher=Elsevier |language=en |isbn=978-0-12-803243-5 |last2=Parker |first2=W. G. |editor2-last=Parker |editor2-first=William G.}} The Revuelto Creek fossils were deposited in the earlier part of the Revueltian, close to around 212 million years ago.{{Cite journal |last=Marsh |first=Adam D. |last2=Parker |first2=William G. |date=2020-11-12 |title=New dinosauromorph specimens from Petrified Forest National Park and a global biostratigraphic review of Triassic dinosauromorph body fossils |url=https://escholarship.org/content/qt01w536hs/qt01w536hs.pdf |journal=PaleoBios |volume=37 |pages=1–56 |doi=10.5070/p9371050859 |issn=2373-8189}}

Though it would not be formally named until 1997, the fossils of Gojirasaurus were frequently mentioned through the 1980s and 1990s, as a robust Coelophysis-like theropod from Revuelto Creek.{{Cite journal |last1=Carpenter |first1=Kenneth |last2=Parrish |first2=Michael |date=1985 |title=Late Triassic Vertebrates from Revuelto Creek, Quay County, New Mexico |url=https://nmgs.nmt.edu/publications/guidebooks/downloads/36/36_p0197_p0198.pdf |journal=New Mexico Geological Society Guidebook, 36th Field Conference, Santa Rosa |pages=197–198}}{{Cite journal |last1=Long |first1=Robert A. |last2=Murry |first2=Phillip A. |date=1995 |title=Late Triassic (Carnian and Norian) tetrapods from the Southwestern United States |url=https://books.google.com/books?id=wbv9CQAAQBAJ |journal=New Mexico Museum of Natural History and Science Bulletin |volume=4 |pages=1–254}} Among these preliminary accounts is a short description by Parrish & Carpenter (1986).{{Cite book |last1=Parrish |first1=Michael |chapter-url=https://archive.org/details/beginningofageof0000unse/page/150/mode/2up |title=The Beginning of the Age of Dinosaurs |last2=Carpenter |first2=Kenneth |publisher=Cambridge University Press |year=1986 |isbn=0521303281 |editor-last=Padian |editor-first=Kevin |location=New York |pages=151–160 |chapter=A new vertebrate fauna from the Dockum Formation (Late Triassic) of eastern New Mexico}} In 1994, an unpublished thesis by Adrian Hunt attempted to name the Revuelto Creek theropod as "Revueltoraptor lucasi". Hunt's conception of the species included not just UCM 47221, but also numerous theropod-like fossils stored at the NMMNH (New Mexico Museum of Natural History and Science). As a name, "Revueltoraptor lucasi" was never formally published, and later evaluations recognized that most of Hunt's additional fossils are likely from Shuvosaurus rather than theropods.

The generic name Gojirasaurus is derived from the name of the giant Japanese movie monster "Gojira" (Godzilla) and the Greek word "{{lang|grc-Latn|sauros}}" ({{lang|grc|σαυρος}}) meaning "lizard";{{cite book | author = Liddell, Henry George and Robert Scott | year = 1980 | title = A Greek-English Lexicon | publisher = Oxford University Press | location = United Kingdom | isbn = 978-0-19-910207-5| title-link = A Greek-English Lexicon | edition = Abridged }} thus, "Godzilla lizard". Carpenter (1997) selected "Gojira" in reference to the relatively large size of this theropod, which exceeded that of its Triassic counterparts. The specific name quayi, is a reference to Quay County.

File:Gojirasaurus BW.jpg

Gojirasaurus is one of the largest theropods known from the Triassic Period, with an estimate total length of {{convert|5.5|-|6|m|ft}} long and a weight of at least {{convert|150|kg|lb}}.{{cite book |last=Paul |first=Gregory S. |year=2010 |title=The Princeton Field Guide to Dinosaurs |url=https://archive.org/details/princetonfieldgu0000paul |url-access=registration |publisher=Princeton University Press |location=Princeton, NJ |page=[https://archive.org/details/princetonfieldgu0000paul/page/74 74] |isbn=978-0-691-13720-9}} Benson et al. (2018) estimated that it weighed around {{convert|190|kg|lb}}, only exceeded by Herrerasaurus among Triassic carnivorous dinosaurs.{{Cite journal |last1=Benson |first1=Roger B. J. |last2=Hunt |first2=Gene |last3=Carrano |first3=Matthew T. |last4=Campione |first4=Nicolás |date=2018 |editor-last=Mannion |editor-first=Philip |title=Cope's rule and the adaptive landscape of dinosaur body size evolution |url=https://onlinelibrary.wiley.com/doi/10.1111/pala.12329 |journal=Palaeontology |language=en |volume=61 |issue=1 |pages=13–48 |doi=10.1111/pala.12329 |bibcode=2018Palgy..61...13B |issn=0031-0239}} Carpenter (1997) pointed to features of the pelvis and ankle suggesting that this was an immature individual, and could therefore have grown to even a larger size in maturity. Christopher T. Griffin (2019) and Griffin and Nesbitt (2020) confirmed that Gojirasaurus possessed features indicative of ontogenetic immaturity.{{Cite journal|last=Griffin|first=Christopher T.|date=September 2019|title=Large neotheropods from the Upper Triassic of North America and the early evolution of large theropod body sizes|journal=Journal of Paleontology|language=en|volume=93|issue=5|pages=1010–1030|doi=10.1017/jpa.2019.13|issn=0022-3360|doi-access=free|bibcode=2019JPal...93.1010G }}

According to Nesbitt et al. (2007), Gojirasaurus can be distinguished based on the fact that its tibia is more robust than that of its relative Coelophysis. Rauhut (2003) attempted to diagnose this genus based on the fact that the mid/posterior dorsal vertebrae had taller neural spines than those observed in other coelophysoids.{{Cite journal |last=Rauhut |first=Oliver W.M. |date=2003 |title=The interrelationships and evolution of basal theropod dinosaurs |url=https://www.palass.org/sites/default/files/media/publications/special_papers_in_palaeontology/number_69/spp69_pp1-213.pdf |journal=Special Papers in Palaeontology |volume=69 |pages=1–213}}

Griffin (2019) followed Nesbitt (2007)'s diagnosis on Gojirasaurus and distinguishes it from Megapnosaurus and Dilophosaurus based on several characteristics of metatarsal V.

Parrish and Carpenter (1986) drew similarities between UCM 47221 and "Halticosaurus liliensterni", a large German theropod now known as Liliensternus. Both were assigned to the family "Procompsognathidae", though the authors acknowledged that family names in use at the time were provisional and likely to be obsolete in the future. Conversely, Hunt (1994) and Hunt et al. (1998){{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. |date=1998 |title=Late Triassic dinosaurs from the western United States |url=http://www.miketaylor.org.uk/dino/nm/HuntEtAl1998-lt-dinosaurs-western-us.pdf |journal=Geobios |language=en |volume=31 |issue=4 |pages=511–531 |doi=10.1016/S0016-6995(98)80123-X|bibcode=1998Geobi..31..511H }} argued that the specimen is a herrerasaurid, alongside various other fossils from the Late Triassic of North America.{{cite web |last=Mortimer |first=Mickey |year=2012 |title=Coelophysoidea |url=https://www.theropoddatabase.com/Coelophysoidea.htm#Gojirasaurusquayi |url-status= |archive-url= |archive-date= |website=The Theropod Database}} The formal naming and description of Gojirasaurus by Carpenter (1997) firmly classified it within Coelophysoidea. Many traits were comparable to Coelophysis, Dilophosaurus, and particularly Liliensternus.

Various phylogenetic analyses in the 2000s supported coelophysoid affinities for Gojirasaurus, close to or among the smaller Coelophysidae rather than with Dilophosaurus.Tykoski, R.S. & Rowe, T. (2004). "Ceratosauria". In: Weishampel, D.B., Dodson, P., & Osmolska, H. (Eds.) The Dinosauria (2nd edition). Berkeley: University of California Press. pp. 47–70 {{ISBN|0-520-24209-2}}{{Cite journal |last=Yates |first=Adam M. |date=2005 |title=A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods |url=https://core.ac.uk/download/pdf/39674608.pdf |journal=Palaeontologia Africana |volume=41 |pages=105–122}}{{Cite journal |last1=Carrano |first1=Matthew T. |last2=Hutchinson |first2=John R. |last3=Sampson |first3=Scott D. |date=2005-12-30 |title=New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona |url=https://www.researchgate.net/publication/230808545 |journal=Journal of Vertebrate Paleontology |volume=25 |issue=4 |pages=835–849 |doi=10.1671/0272-4634(2005)025[0835:niosha]2.0.co;2 |issn=0272-4634}}{{Cite journal |last1=Ezcurra |first1=Martin D. |last2=Novas |first2=Fernando E. |date=2007 |title=Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina |url=https://www.researchgate.net/publication/247525624 |journal=Historical Biology |language=en |volume=19 |issue=1 |pages=35–72 |doi=10.1080/08912960600845791 |bibcode=2007HBio...19...35E |issn=0891-2963}} Many studies noted that its remains are too fragmentary for further elaboration. Starting with Yates (2005), the monophyly of Coelophysoidea has been brought into question, with Dilophosaurus resolving as more closely related to averostrans (non-coelophysoid theropods). In some analyses, Gojirasaurus maintains its position among the coelophysids, but other studies instead bring it over to the lineage leading to Dilophosaurus and averostrans.{{Cite journal |last1=Ezcurra |first1=Martín D |last2=Butler |first2=Richard J |last3=Maidment |first3=Susannah C R |last4=Sansom |first4=Ivan J |last5=Meade |first5=Luke E |last6=Radley |first6=Jonathan D |date=2021-01-01 |title=A revision of the early neotheropod genus Sarcosaurus from the Early Jurassic (Hettangian–Sinemurian) of central England |url=https://academic.oup.com/zoolinnean/article/191/1/113/5861188 |journal=Zoological Journal of the Linnean Society |language=en |volume=191 |issue=1 |pages=113–149 |doi=10.1093/zoolinnean/zlaa054 |issn=0024-4082|hdl=11336/160038 |hdl-access=free }}{{Cite journal |last1=Spiekman |first1=Stephan N. F. |last2=Ezcurra |first2=Martín D. |last3=Butler |first3=Richard J. |last4=Fraser |first4=Nicholas C. |last5=Maidment |first5=Susannah C. R. |date=2021 |title=Pendraig milnerae, a new small-sized coelophysoid theropod from the Late Triassic of Wales |journal=Royal Society Open Science |language=en |volume=8 |issue=10 |doi=10.1098/rsos.210915 |issn=2054-5703 |pmc=8493203 |pmid=34754500|bibcode=2021RSOS....810915S }}{{Cite journal |last1=Ezcurra |first1=Martín D. |last2=Marke |first2=Daniel |last3=Walsh |first3=Stig A. |last4=Brusatte |first4=Stephen L. |date=2023-11-20 |title=A revision of the 'coelophysoid-grade' theropod specimen from the Lower Jurassic of the Isle of Skye (Scotland ) |url=https://www.researchgate.net/publication/374794630 |journal=Scottish Journal of Geology |language=en |volume=59 |issue=1–2 |page=012 |doi=10.1144/sjg2023-012 |bibcode=2023ScJG...59...12E |issn=0036-9276}}

A persistent question regarding Gojirasaurus quayi is how much of its fossil material actually belongs to one species.{{cite journal |last1=Nesbitt |first1=Sterling J. |last2=Irmis |first2=Randall B. |last3=Parker |first3=William G. |date=June 2007 |title=A critical re-evaluation of the Late Triassic dinosaur taxa of North America |url=http://www.dinochecker.com/papers/triassic_dinos_revisited_Nesbitt_et_al_2007.pdf |journal=Journal of Systematic Palaeontology |volume=5 |issue=2 |pages=209–243 |doi=10.1017/S1477201907002040 |bibcode=2007JSPal...5..209N |s2cid=28782207}} Gojirasaurus coexisted with Shuvosaurus, an unusual reptile with a toothless beak and bipedal stance. Though previously regarded as an aberrant dinosaur, by 2007 most specialists agreed that Shuvosaurus was actually more closely related to crocodilians, and that its dinosaur-like traits are merely convergent.{{Cite journal |last1=Lucas |first1=Spencer G. |last2=Spielmann |first2=Justin A. |last3=Hunt |first3=Adrian P. |date=2007 |title=Taxonomy of Shuvosaurus, a Late Triassic archosaur from the Chinle Group, American Southwest |url=https://books.google.com/books?id=EGDmCQAAQBAJ&pg=259 |journal=New Mexico Museum of Natural History and Science Bulletin |volume=41 |pages=259–261}} According to Nesbitt et al. (2007), some components of UCM 47221, such as the tibia and pubis, are clearly coelophysoid in form and distinct from Shuvosaurus. However, other parts of the skeleton, such as the scapula and vertebrae, are not readily distinguishable from Shuvosaurus. The serrated tooth could be from any number of large carnivorous archosaurs which inhabited the area.

Even the assuredly coelophysoid components of the skeleton have few unambiguous unique features. For example, the robust tibia is similar to Coelophysis-like fossils which Padian (1986) described from Petrified Forest National Park in Arizona, only differing in size.{{Cite book |last=Padian |first=Kevin |chapter-url=https://archive.org/details/beginningofageof0000unse/page/44/mode/2up |title=The Beginning of the Age of Dinosaurs |publisher=Cambridge University Press |year=1986 |isbn=0521303281 |editor-last=Padian |editor-first=Kevin |location=New York |pages=45–60 |chapter=On the type material of Coelophysis Cope (Saurischia: Theropoda) and a new specimen from the Petrified Forest of Arizona (Late Triassic: Chinle Formation)}} Several studies regard Gojirasaurus as a "metataxon": a collection of fossils for which assignment to a single species can neither be proven nor disproven. Nevertheless, Gojirasaurus persists in the scientific literature as a useful example of a large Triassic coelophysoid, validity notwithstanding.{{Cite journal |last1=Griffin |first1=Christopher T. |last2=Nesbitt |first2=Sterling J. |date=2020 |title=Does the Maximum Body Size of Theropods Increase across the Triassic–Jurassic Boundary? Integrating Ontogeny, Phylogeny, and Body Size |url=https://anatomypubs.onlinelibrary.wiley.com/doi/10.1002/ar.24130 |journal=The Anatomical Record |language=en |volume=303 |issue=4 |pages=1158–1169 |doi=10.1002/ar.24130 |pmid=30968581 |issn=1932-8486}}

{{Main|Bull Canyon Formation}}

Gojirasaurus's assignment to the Coelophysoidea would suggest that it was a bipedal, terrestrial, actively mobile carnivore.

The Revuelto Creek area preserves a diverse fauna of both terrestrial and aquatic animals from the Bull Canyon Formation.{{Cite journal |last=Hunt |first=Adrian P. |date=2001 |title=The vertebrate fauna, biostratigraphy and biochronology of the type Revultian land vertebrate faunachron, Bull Canyon Formation (Upper Triassic), east-central New Mexico |url=https://nmgs.nmt.edu/publications/guidebooks/downloads/52/52_p0123_p0151.pdf |journal=New Mexico Geological Society Guidebook, 52nd Field Conference, Geology of the Llano Estacado |language=en |publisher=New Mexico Geological Society |pages=123–151 |doi=10.56577/FFC-52.123 |isbn=978-1-58546-087-8}} On land, herbivorous pseudosuchians are quite common, including the shuvosaurid Shuvosaurus inexpectatus, the aetosaurs Typothorax coccinarum, Paratypothorax, and Rioarribasuchus,{{cite journal |last=Parker |first=W. G. |year=2007 |title=Reassessment of the aetosaur 'Desmatosuchus' chamaensis with a reanalysis of the phylogeny of the Aetosauria (Archosauria:Pseudosuchia) |url=http://www.miketaylor.org.uk/dino/nm/Parker2007-Heliocanthus.pdf |journal=Journal of Systematic Palaeontology |volume=5 |issue=1 |pages=41–68 |bibcode=2007JSPal...5...41P |doi=10.1017/S1477201906001994 |s2cid=85826683}} and the small aetosauriform Revueltosaurus callenderi.{{Cite journal |last=Parker |first=William G. |last2=Nesbitt |first2=Sterling J. |last3=Irmis |first3=Randall B. |last4=Martz |first4=Jeffrey W. |last5=Marsh |first5=Adam D. |last6=Brown |first6=Matthew A. |last7=Stocker |first7=Michelle R. |last8=Werning |first8=Sarah |date=2022 |title=Osteology and relationships of Revueltosaurus callenderi (Archosauria: Suchia) from the Upper Triassic (Norian) Chinle Formation of Petrified Forest National Park, Arizona, United States |url=https://anatomypubs.onlinelibrary.wiley.com/doi/10.1002/ar.24757 |journal=The Anatomical Record |language=en |volume=305 |issue=10 |pages=2353–2414 |doi=10.1002/ar.24757 |issn=1932-8486 |pmc=9544919 |pmid=34585850}} Both Shuvosaurus and Revueltosaurus have previously been misidentified as dinosaurs.

Among terrestrial carnivores, Gojirasaurus was joined by at least a few other true dinosaurus. Bull Canyon dinosaur fragments are sometimes identified as coelophysids, herrerasaurids, and/or Chindesaurus, but most are too fragmentary to assess in great detail. Lepidosauromorphs, crocodylomorphs, and large carnivorous "rauisuchians" are also represented by rare fragments. Particularly robust archosauromorph limb fragments, previously thought to be from a late-surviving rhynchosaur ("Otischalkia"), are probably from malerisaurine azendohsaurids instead.{{Cite journal |last=Christian A. Sidor |last2=Marsh |first2=Adam D. |last3=Smith |first3=Elliot Armour |date=2024-05-30 |title=THE ALLOKOTOSAUR (REPTILIA: ARCHOSAUROMORPHA) ASSEMBLAGE FROM A MULTITAXIC BONEBED IN THE SONSELA MEMBER (JIM CAMP WASH BEDS, CHINLE FORMATION) AT PETRIFIED FOREST NATIONAL PARK, U.S.A. |url=https://petrifiedforestfieldinstitute.org/lithodendron/lithodendron/spring-2024/the-allokotosaur-assemblage-from-a-multitaxic-bonebed-at-petrified-forest/ |journal=Lithodendron: The Science and History Journal of Petrified Forest National Park |volume=1 |issue=Spring 2024 |pages=95–118 |doi=10.69575/FAIY7658|url-access=subscription }} The early turtle Chinlechelys tenertesta is a notable component of the terrestrial fauna.{{Cite journal |last1=Joyce |first1=W.G. |last2=Lucas |first2=S.G. |last3=Scheyer |first3=T.M. |last4=Heckert |first4=A.B. |last5=Hunt |first5=A.P. |year=2009 |title=A thin-shelled reptile from the Late Triassic of North America and the origin of the turtle shell |journal=Proceedings of the Royal Society B |volume=276 |issue=1656 |pages=507–513 |doi=10.1098/rspb.2008.1196 |issn=1471-2954 |jstor=30244885 |pmc=2664348 |pmid=18842543}}{{Cite journal |last1=Lichtig |first1=A. J. |last2=Lucas |first2=S. G. |year=2021 |title=Chinlechelys from the Upper Triassic of New Mexico, USA, and the origin of turtles |journal=Palaeontologia Electronica |volume=24 |issue=1 |pages=Article number 24.1.a13 |doi=10.26879/886 |doi-access=free}}

Phytosaur fossils are common at Revuelto Creek. One particularly impressive phytosaur skull was initially referred to "Rutiodon" (Smilosuchus) gregorii, and later to "Arribasuchus" buceros. It and other Bull Canyon phytosaur remains most likely belong to a species of Machaeroprosopus (Pseudopalatus).{{Cite journal |last=Hunt |first=Adrian P. |last2=Lucas |first2=Spencer G. |last3=Spielmann |first3=Justin A. |date=2006 |title=Sexual dimorphism in a large brachyrostral phytosaur (Archosauria: Crurotarsi) from the Late Triassic of western North America |url=https://www.researchgate.net/publication/281782930 |journal=New Mexico Museum of Natural History and Science Bulletin |volume=37 |pages=563–567}} Small metoposaurid amphibians, sometimes identified as "Apachesaurus gregorii", frequent the area alongside larger metoposaurids. Various fish inhabited the waterways: arganodontid lungfish, coelacanths (Quayia zideki), and actinopterygians.

{{Portal|Dinosaurs}}

• Timeline of coelophysoid research

{{Reflist}}

• [https://fossilvertebratesandtraces.colorado.edu/index.php/Detail/vertebrate/54346/subsite/vertebrate Low-resolution fossil images on The University of Colorado Museum of Natural History Fossil Vertebrate Collection database]

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Category:Coelophysoidea

Category:Dinosaur genera

Category:Norian dinosaurs

Category:Taxa named by Kenneth Carpenter

Category:Fossil taxa described in 1997

Category:Dinosaurs of the United States

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