Allodaposuchus
| Allodaposuchus | |
|---|---|
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| A. precedensskull | |
Scientific classification
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| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Clade: | Archosauria |
| Clade: | Pseudosuchia |
| Clade: | Crocodylomorpha |
| Clade: | Crocodyliformes |
| Clade: | Eusuchia |
| Clade: | †Allodaposuchidae |
| Genus: | †Allodaposuchus Nopcsa,1928 |
| Type species | |
| †Allodaposuchus precedens Nopcsa, 1928
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| Species | |
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| Synonyms | |
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Allodaposuchusis anextinctgenusofcrocodyliformsthat lived in what is now southern Europe during theCampanianandMaastrichtianstages, and possibly theSantonianstage, of theLate Cretaceous.Although generally classified as a non-crocodylianeusuchiancrocodylomorph,it is sometimes placed as one of the earliest true crocodylians.Allodaposuchusis one of the most common Late Cretaceous crocodylomorphs from Europe, with fossils known fromRomania,Spain,andFrance.
Description
[edit]
Like many other Cretaceous crocodylomorphs,Allodaposuchushas a relatively small body size compared to living crocodylians. The largest known specimen ofAllodaposuchusbelongs to an individual that was probably around 3 metres (9.8 ft) long.[2]Although the shape varies between species, in generalAllodaposuchushas a short, flattened, and rounded skull.Allodaposuchus precedenshas a brevirostrine or "short-snouted" skull with a snout about the same length as theskull table(the region of the skull behind the eye sockets) andA. subjuniperushas a mesorostrine or "middle-snouted" skull with a snout that is longer than the skull table.[2][3]The main feature that distinguishesAllodaposuchusspecies from other related crocodylomorphs is the orientation of a groove at the back of the skull called the cranioquadrate passage; unlike the cranioquadrate passages of other crocodylomorphs, which are only visible at the back of the skull, the cranioquadrate passage ofAllodaposuchusis visible when the skull is viewed from the side.[4]
At least one species ofAllodaposuchus,A. hulki,may have adaptations that would have allowed it to live on land for extended periods of time.A. hulkihas largesinusesin its skull that are not seen in any other crocodylian living or extinct and may have aided it in hearing out of water, as well as lightening the skull. Moreover,A. hulkihas well-developed muscle attachments on itsscapula,humerus,andulnabones that would have allowed the forelimbs to have been held in a semi-erect stance suitable for walking over land. Remains ofA. hulkicome from interbeddedsandstonesandmarlsthat, based on the presence ofcharophytealgae, likely formed in ephemeralpondsin a largefloodplainfar from permanent bodies of water like lakes or rivers.A. hulkimay therefore have spent much of its time out of water, travelling between these ponds for food.[5]
History of study
[edit]While there are severaldescribedspecies ofAllodaposuchus,the precise membership of the group is currently disputed.
Allodaposuchus precedens
[edit]Thetype speciesofAllodaposuchus,A. precedens,was named by Hungarian paleontologistFranz Nopcsain 1928 fromVălioara,Romania.[6]Nopcsa found bone fragments in a deposit of theHațeg Basinthat dates back to the lateMaastrichtianstage – the very end of theLate Cretaceous.Several partial skulls from Spain and France were attributed toA. precedensin 2001.[7]Some of these skulls came fromCampanian-age deposits slightly older than those in Romania, meaning that the species must have persisted for about 5 million years.[2]
A 2013 study proposed that the French and Spanish fossils assigned toA. precedensin 2001 might actually represent a new unnamed species ofAllodaposuchuscurrently identified asAllodaposuchussp.[2]A study published in 2005 had suggested that these fossils belong to several different genera of crocodylomorphs and that the original Romanian material is too fragmentary to assign to its own genus, makingAllodaposuchusanomen dubiumor "dubious name".[8]However, the 2013 study reaffirmed the Romanian material's distinctiveness from other European Cretaceous crocodylomorphs and therefore reaffirmed the validity ofAllodaposuchusas a genus.[2]
Allodaposuchus (Agaresuchus) subjuniperus?
[edit]In 2013, a second species ofAllodaposuchus,A. subjuniperus,was named on the basis of a skull from the late-MaastrichtianConquès Formation,part of theTremp Group,in theprovince of Huesca,Spain. The skull was found underneath ajunipertree whose roots had grown between the bones, hence the species namesubjuniperusor "under juniper" inLatin.[2]However, in 2016,A. subjuniperuswas moved to a new genus,Agaresuchusalong with the type species of that genus,A. fontisensis,on the grounds that the Spanish species were sufficiently distinct fromA. precedens.[9]In 2021, a phylogenetic analysis by Blanco disputed this result, suggesting that bothA. fontisensisandA. subjuniperusbelong within the genusAllodaposuchusproper, alongside the two species ofLohuecosuchus:L. megadontosandL. mechinorum.[10]
Allodaposuchus palustris
[edit]In 2014,A. palustriswas described from a partial skull and other skeletal fragments found in Maastrichtian age sediments of theTremp Formationin a fossil locality called Fumanya Sud in the southernPyrenees.[11]These remains allowed for the first detailed description of thepostcranial(non-skull) anatomy ofAllodaposuchus.
Allodaposuchus hulki
[edit]A fourth species ofAllodaposuchus,A. hulki,was named in 2015 and also came from the Tremp Formation, although this time in a locality called Casa Fabà. The species is named after theHulkfromMarvel Comics,in reference to features on the bones that suggest it had strong muscles.[5]
Allodaposuchus iberoarmoricanus
[edit]A. palustriswas described by Blanco in 2021 based on fossils discovered inLate Campanian-agedfluvialdeposits inVelaux-La Bastide Neuve,inBouches-du-RhôneDepartmentofsouthern France.[10]Thespecies nameis in reference to theIbero-Armorican islandof theCretaceous European Archipelago.[10]
Allodaposuchus (Agaresuchus) fontisensis?
[edit]In 2016, the new genus and speciesAgaresuchus fontisensiswas discovered anddescribed.It was named from theLo Huecofossil site inFuentes,Cuenca,Spain;fontisis the Latin name of Fuentes.[9]A. subjuniperuswas then also placed into the newgenusAgaresuchus.[9]However, Blanco's 2021 study has called this into question, and instead proposed that they should both be considered members ofAllodaposuchus,withAgaresuchusas a junior synonym.[10]
Allodaposuchus (Lohuecosuchus) megadontosandAllodaposuchus (Lohuecosuchus) mechinorum?
[edit]The genusLohuecosuchuswas named in 2015 and contained two species,L. megadontosandL. mechinorum,from Spain and southern France.[12]However, Blanco's 2021 study has called this into question, and instead proposed that they should both be considered members ofAllodaposuchus,withLohuecosuchusas a junior synonym.[10]
Classification
[edit]
Allodaposuchusbelongs to thecladeAllodaposuchidae.The exact placement of Allodaposuchidae is still in dispute. Narváezet al.considered it thesister grouptoHylaeochampsidae,which together form acladethat is sister toCrocodylia.[12]Other studies have alternatively recovered them not as sister taxon, but rather as anevolutionary gradetowards Crocodylia, with Hylaeochampsidae morebasalthan Allodaposuchidae.[13][14]Alternatively, a 2021 analysis incorporatingpostcranialinformation recovered Allodaposuchidae within Crocodylia.[15]
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Cladogram 1:Narváezet al.,2015 |
Cladogram 2:Rio & Mannion, 2021 |
Cladogram 3:Blanco, 2021
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The internalphylogenyofAllodaposuchidaecan be shown in thecladogrambelow from the 2021 Blanco study:[15]
| Allodaposuchidae |
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In the 2021 study, Blanco recoveredAllodaposuchusasparaphyletic,withAgaresuchusandLohuecosuchus.Accordingly, Blanco proposed thatAgaresuchusandLohuecosuchusshould be consideredjunior synonymsofAllodaposuchus.[15]
References
[edit]- ^Hristova, L. (2020). "Description of Crocodylomorph Teeth from the Late Cretaceous Locality near Tran, Western Bulgaria".Comptes Rendus de l'Académie Bulgare des Sciences.73(3): 379–385.doi:10.7546/CRABS.2020.03.11.
- ^abcdefPuértolas-Pascual, E.; Canudo, J.I.; Moreno-Azanza, M. (2014). "The eusuchian crocodylomorphAllodaposuchus subjuniperussp. nov., a new species from the latest Cretaceous (upper Maastrichtian) of Spain ".Historical Biology.26(1): 91–109.Bibcode:2014HBio...26...91P.doi:10.1080/08912963.2012.763034.S2CID85004774.
- ^Martin, J.E. (2010). "AllodaposuchusNopsca, 1928 (Crocodylia, Eusuchia), from the Late Cretaceous of southern France and its relationships to Alligatoroidea ".Journal of Vertebrate Paleontology.30(3): 756–767.Bibcode:2010JVPal..30..756M.doi:10.1080/02724631003758318.
- ^Delfino, M.; Codrea, V.; Folie, A.; Dica, P.; Godefroit, P.; Smith, T. (2008)."A complete skull ofAllodaposuchus precedensNopcsa, 1928 (Eusuchia) and a reassessment of the morphology of the taxon based on the Romanian remains ".Journal of Vertebrate Paleontology.28:111–122.doi:10.1671/0272-4634(2008)28[111:ACSOAP]2.0.CO;2.
- ^abBlanco, Alejandro; Fortuny, Josep; Vicente, Alba; Luján, Angel H.; García Marçà, Jordi Alexis; Sellés, Albert G. (2015)."A new species ofAllodaposuchus(Eusuchia, Crocodylia) from the Maastrichtian (Late Cretaceous) of Spain: phylogenetic and paleobiological implications ".PeerJ.3:e1171: 1–35.doi:10.7717/peerj.1171.PMC4558081.PMID26339549.
- ^Nopcsa, F (1928). "Paleontological notes on Reptilia. 7. Classification of the Crocodilia".Geologica Hungarica, Series Palaeontologica.1:75–84.
- ^Buscalioni, A.D.; Ortega, F.; Weishampel, D.B.; Jianu, C.M. (2001). "A revision of the crocodyliformAllodaposuchus precedensfrom the Upper Cretaceous of the Hateg Basin, Romania. Its relevance in the phylogeny of Eusuchia ".Journal of Vertebrate Paleontology.21:74–86.doi:10.1671/0272-4634(2001)021[0074:AROTCA]2.0.CO;2.
- ^Martin, J.E.; Buffetaut, E. (2005). "An overview of the Late Cretaceous crocodilian assemblage from Cruzy, southern France".Kaupia.14:33–40.
- ^abcNarváez, I.; Brochu, C.A.; Escaso, F.; Pérez-García, A.; Ortega, F. (2016)."New Spanish Late Cretaceous eusuchian reveals the synchronic and sympatric presence of two allodaposuchids".Cretaceous Research.65:112–125.Bibcode:2016CrRes..65..112N.doi:10.1016/j.cretres.2016.04.018.
- ^abcdeBlanco, A. (2021)."Importance of the postcranial skeleton in eusuchian phylogeny: Reassessing the systematics of allodaposuchid crocodylians".PLOS ONE.16(6): e0251900.Bibcode:2021PLoSO..1651900B.doi:10.1371/journal.pone.0251900.PMC8189472.PMID34106925.
- ^Blanco, Alejandro; Puértolas Pascual, Eduardo; Marmi, Josep; Vila, Bernat; Sellés, Albert G. (2014)."Allodaposuchus palustrissp. nov. from the Upper Cretaceous of Fumanya (South Eastern Pyrenees, Iberian Peninsula): Systematics, Palaeoecology and Palaeobiogeography of the Enigmatic Allodaposuchian Crocodylians ".PLoS One.9(12): 1–34.Bibcode:2014PLoSO...9k5837B.doi:10.1371/journal.pone.0115837.PMC4281157.PMID25551630.
- ^abNarváez, I.; Brochu, C.A.; Escaso, F.; Pérez-García, A.; Ortega, F. (2015)."New crocodyliforms from southwestern Europe and definition of a diverse clade of european Late Cretaceous basal eusuchians".PLOS ONE.10(11): e0140679.Bibcode:2015PLoSO..1040679N.doi:10.1371/journal.pone.0140679.PMC4633049.PMID26535893.
- ^Michael S. Y. Lee; Adam M. Yates (27 June 2018)."Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil".Proceedings of the Royal Society B.285(1881).doi:10.1098/rspb.2018.1071.PMC6030529.PMID30051855.
- ^Rio, Jonathan P.; Mannion, Philip D. (6 September 2021)."Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem".PeerJ.9:e12094.doi:10.7717/peerj.12094.PMC8428266.PMID34567843.
- ^abcBlanco, Alejandro (2021-06-09)."Importance of the postcranial skeleton in eusuchian phylogeny: Reassessing the systematics of allodaposuchid crocodylians".PLOS ONE.16(6): e0251900.Bibcode:2021PLoSO..1651900B.doi:10.1371/journal.pone.0251900.ISSN1932-6203.PMC8189472.PMID34106925.
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