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Revueltosaurus

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Revueltosaurus
Temporal range:Late Triassic,225–209.5Ma
Skull
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Aetosauriformes
Genus: Revueltosaurus
Hunt, 1989
Species
  • R. callenderiHunt, 1989 (type)
  • "R". olseni?(Hunt & Lucas, 1994)
  • "R". hunti?Heckert, 2002
Synonyms

Revueltosaurus( "Revuelto lizard" ) is anextinctgenusofsuchianpseudosuchian[1]fromLate Triassic(lateCarnianto middleNorianstage) deposits ofNew Mexico,ArizonaandNorth Carolina,United States. Many specimens, mostly teeth, have been assigned toRevueltosaurusover the years. Currently, three species are included in this genus, all of which were originally thought to represent monospecific genera of basalornithischiandinosaurs.[2][3][4]Revueltosaurus was about 1 meter (3.3 feet) long.

Species

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Revueltosaurus callenderi

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Teeth ofRevueltosaurus callendericollected in Petrified Forest National Park

R. callenderiwas named by Adrian P. Hunt in1989and it is thetype speciesof the genus. Thegeneric namehonors its type locality, Revuelto Creek,Quay Countyof New Mexico. Revuelto is derived from Spanishrevuelta,"revolution", in reference to the importance of the Late Triassic period for terrestrialvertebrateevolution. Thespecific namehonours the director of theNew Mexico Museum of Natural HistoryJonathan F. Callender.R. callenderiwas originally described as a basal ornithischian on the basis of 32 teeth: theholotypeNMMNHP-4957 a nearly complete premaxillary (incisiform) tooth, theparatypes(NMMNH P-4958-9, a nearly complete maxillary or dentary tooth crown with root and a nearly complete premaxillary tooth crown) and 28 referred specimens. All specimens were collected from the type locality, from theBull Canyon Formation,dating to the Norian stage.[5]

Later, other teeth were described by Padian (1990), Kaye and Padian (1994) and Hunt and Lucas (1994), fromChinle FormationofArizona.Andrew B. Heckert (2002) redescribed the genus in detail and named a second species,R. huntion the basis of teeth described by Hunt and Lucas (1994). He referredMNAV3690, an isolated tooth assigned by Kaye and Padian (1994) toR. callenderifrom thePlacerias/Downs quarry near St Johns, Arizona, toTecovasaurus.Heckert listed the referred specimens CMNH PR1697-1699 originally described by Padian (1990) and the topotypes NMMNH P-4960, P-16573, P-33783-798. Four teeth were referred to aff.Revueltosaurus callenderi(NMMNH P-17362, P-17382 and P-17187).R. callenderiis known to date only from the Revueltian (early-mid Norian, the type locality dates back to the early Norian) of Bull Canyon Formation (Dockum Group,New Mexico) and the Painted Desert of the Petrified Forest Member (Chinle Formation, Arizona).[6]Parkeret al.(2005) described several partially completeskeletonsofR. callenderi(PEFO 33787-95), including a nearly complete and a partialskulls,from theRevueltosaurusQuarry,Petrified Forest National Park.[7]Other skeletons were discovered from the Painted Desert, including the nearly complete specimen PEFO 34561 which have been fully reconstructed recently.[8]According to Sterling J. Nesbitt (2011), much of theRevueltosauruscranialandpostcranialmaterial originates from a monotypic bonebed. Some specimens occur as isolated bones, while others are completely associated and even fully articulated. Thus, nearly the entire skeleton ofR. callenderiis known and a full description of it is being prepared.[1]

Revueltosaurus hunti

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R. huntiwas named by Andrew B. Heckert in2002as a basal ornithischian. Thespecific namehonours the paleontologist Adrian P. Hunt for his contributions to the Triassic paleontology. It is known from theholotypeNMMNHP-29357, a nearly complete tooth crown, the paratypes NMMNH P-29358-9 and topotypes P-29347-54 (and aff.R. huntispecimens NMMNH P-29355, P-29359) which were collected in the NMMNH locality 1171,Santa Fe County,New Mexico from the Los Esteros Member of theSanta Rosa Formation,Chinle Group, dating to the latestCarnianstage (Adamanian). Other isolated teeth (which Long and Murry (1995) assigned toR. callenderi) were collected two localities in the Blue Hills, east-central Arizona, from the Blue Mesa Member of the Chinle Formation, dating also to the latest Carnian.UCMPlocality V92048 have yielded the topotypes UCMPV 173839-41, and the referred teeth UCMPV 139563-75 are from UCMP locality V7307.[6]

Parkeret al.(2005) confirmed thatR. huntimay be assignable toRevueltosaurus,on the basis of undescribedsquamosalcollected from its referred locality that is nearly identical to that seen in the Petrified Forest material ofR. callenderi,thus suggesting thatR. huntiis a pseudosuchian.[7]In response to the report thatR. callenderiwas not an ornithischian, Heckert (2005) erected a new genus forR. hunti,Krzyzanowskisaurus.He suggested that the denticulated shelf on the teeth represented acingulum.Irmiset al.(2007) and Nesbittet al.(2007) synonymizedKrzyzanowskisauruswithRevueltosaurus,and argued that this condition of the dentation is anautapomorphyofR. huntiand is not homologous to the ornithischian asymmetric swelling or a mammalian cingulum.[2][3]

Revueltosaurus olseni

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R. olseniwas named by Hunt and Lucas in1994as a basal ornithischian, under the generic namePekinosaurus.Thegeneric namehonorsPekin, North Carolina,where its type series was collected. Thespecific namehonours the paleontologist Paul E. Olsen. Hunt and Lucas named the genus on the basis of the teeth seriesYPM7666-9. Note that they designed theholotypeto be YPM 8545, but the correct designation is YPM 7666 following Heckertet al.(2012). The type series ofR. olseniwas thought to originate from thePekin FormationofMontgomery County.Heckertet al.(2012) noted that recent field work has shown that strata formerly assigned to that formation, including the type locality ofR. olseni,actually pertain to the overlyingCumnock Formation(located in the Sanford Subbasin, Deep River Basin). Hence, the type locality and the newly discovered Moncure locality are from the same stratigraphic interval. The Cumnock Formation belongs to theChathan Group(of theNewark Supergroup), which is recognized as Late Triassic in age. Most workers have assigned the Cumnock Formation a Carnian age. Most recent work, however, demonstrates that much of it is in fact Norian in age. Therefore, the strata exposed at the Moncure locality are probably early Norian in age. The Moncure locality have yielded many referred specimens ofR. olseniincluding NCSM 21647, NCSM 23539, 41-46, 48-50, 52-57, NCSM 24722, 58-62 and NCSM 25194-227. NCSM 24728, 25192-3 and 25228 were referred to?Revueltosaurus.Someosteodermswere referred to it to, as they very similar to those seen in the Petrified Forest material ofR. callenderi.R. olseniis thought to be one of the most common tetrapods from its assemblage.[4]

Irmiset al.(2007) and Nesbittet al.(2007) tentatively synonymizedPekinosaurus(andGaltonia) withRevueltosaurus,and refer its type series toRevueltosaurus sp.They noted that Hunt and Lucas (1994) did not provide any differential diagnosis, autapomorphies, or unique combination of characters forPekinosaurus.Furthermore, they argued that it cannot be differentiated fromR. callenderi.[2][3]Heckertet al.(2012) agreed thatR. olseniis extremely similar toR. callenderi,however they found thatPekinosaurusteeth have slightly different proportions and outline thanR. callenderi.They provided a differential diagnosis for that species and referred it toRevueltosaurus,under the new combinationR. olseni.[4]

Classification

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Restoration

All species ofRevueltosauruswere thought to represent monospecific genera of basalornithischiandinosaursbased on theirteeth.New discoveries of skeletal material ofR. callenderihave suggested thatRevueltosaurusis a pseudosuchian. According to William Parker, "we have pretty much erased the record of Triassic ornithischian dinosaurs from North America, Europe and worldwide, except for South America."[9]His co-author Randall Irmis said, "because the teeth look like those we know from herbivorous ornithischians, people assigned them to the dinosaurs. We think we've shown that you can't rely on the dentition to determine what is an early dinosaur, which casts doubt on all the ornithischians from the Triassic of North America".[10]Nearly complete specimens ofR. callenderiwere tested in a broad phylogenetic analysis ofbasalarchosaurs for the first time when Sterling J. Nesbitt included the specimens PEFO 34561 and PEFO 34269 (a complete and a nearly complete skeleton, respectively) in his 2011 phylogenetic analysis.Revueltosauruswas placed at the base of the cladeSuchiaas thesister taxonto the armored and herbivorousaetosaurs.However,Revueltosaurusitself is not an aetosaur, since Aetosauria was redefined to exclude it. The analysis foundGracilisuchus,Turfanosuchusand theRevueltosaurus+Aetosauriaclade to nest in a largepolytomywithTicinosuchus+Paracrocodylomorpha.Thecladogrambelow follows Nesbitt (2011).[1]

Archosauria

References

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  1. ^abcSterling J. Nesbitt (2011)."The Early Evolution of Archosaurs: Relationships and the Origin of Major Clades".Bulletin of the American Museum of Natural History.352:1–292.doi:10.1206/352.1.hdl:2246/6112.S2CID83493714.
  2. ^abcRandall B. Irmis; William G. Parker; Sterling J. Nesbitt; Jun Liu (2007)."Early ornithischian dinosaurs: the Triassic record"(PDF).Historical Biology.19(1): 3–22.CiteSeerX10.1.1.539.8311.doi:10.1080/08912960600719988.S2CID11006994.
  3. ^abcSterling J. Nesbitt; Randall B. Irmis; William G. Parker (2007)."A critical re-evaluation of the Late Triassic dinosaur taxa of North America"(PDF).Journal of Systematic Palaeontology.5(2): 209–243.doi:10.1017/S1477201907002040.S2CID28782207.
  4. ^abcAndrew B. Heckert; Jonathan S. Mitchell; Vincent P. Schneider; Paul E. Olsen (2012)."Diverse new microvertebrate assemblage from the Upper Triassic Cumnock Formation, Sanford Subbasin, North Carolina, USA".Journal of Paleontology.86(2): 368–390.doi:10.1666/11-098.1.S2CID18239160.
  5. ^Adrian P. Hunt (1989)."A new?ornithischian dinosaur from the Bull Canyon Formation (Upper Triassic) of east-central New Mexico"(PDF).In Lucas, S. G. And A. P. Hunt (Eds.), Dawn of the Age of Dinosaurs in the American Southwest:355–358. Archived fromthe original(PDF)on 2013-05-16.
  6. ^abAndrew B. Heckert (2002)."A revision of the Upper Triassic ornithischian dinosaurRevueltosaurus,with a description of a new species "(PDF).New Mexico Museum of Natural History & Science Bulletin(21): 253–268.
  7. ^abWilliam G. Parker; Randall B. Irmis; Sterling J. Nesbitt; Jeffrey W. Martz; Lori S. Browne (2005)."The Late Triassic pseudosuchianRevueltosaurus callenderiand its implications for the diversity of early ornithischian dinosaurs ".Proceedings of the Royal Society B.272(1566): 963–969.doi:10.1098/rspb.2004.3047.PMC1564089.PMID16024353.
  8. ^[1][permanent dead link]
  9. ^Quoted in Sanders 2005, para. 6.
  10. ^Quoted in Sanders 2005, para. 4.
  • Sanders, Robert. June 24, 2005.Revueltosaurusskeleton unearthed at Petrified Forest upsets dinosaur tale.University of California Berkeley News.Retrieved March 15, 2006.