Allodaposuchus

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Subsectio: Pseudosuchia
Cladus: Suchia
Cladus: Paracrocodylomorpha
Cladus: Loricata
Superordo: Crocodylomorpha
Ordo: Crocodilia
Cladus: Mesoeucrocodylia
Cladus: Neosuchia
Subordo: Eusuchia

Familia: †Allodaposuchidae
Genus: †Allodaposuchus
Name

Allodaposuchus Puértolas, Canudo & Moreno-Azanza, 2013
References

ZooBank
Puértolas-Pascual, E.; Canudo, J.I.; Moreno-Azanza, M. 2013 (Online) 2014 (Print): The eusuchian crocodylomorph Allodaposuchus subjuniperus sp. nov., a new species from the latest Cretaceous (upper Maastrichtian) of Spain. Historical biology 26(1): 91-109. DOI: 10.1080/08912963.2012.763034 Reference page.

Allodaposuchus is an extinct genus of crocodyliforms that lived in what is now southern Europe during the Campanian and Maastrichtian stages of the Late Cretaceous. Although generally classified as a non-crocodylian eusuchian crocodylomorph, it is sometimes placed as one of the earliest true crocodylians. Allodaposuchus is one of the most common Late Cretaceous crocodylomorphs from Europe, with fossils known from Romania, Spain, and France.
Description
Restoration of A. precedens

Like many other Cretaceous crocodylomorphs, Allodaposuchus has a relatively small body size compared to living crocodylians. The largest known specimen of Allodaposuchus belongs to an individual that was probably around 3 metres (9.8 ft) long.[1] Although the shape varies between species, in general Allodaposuchus has a short, flattened, and rounded skull. Allodaposuchus precedens has a brevirostrine or "short-snouted" skull with a snout about the same length as the skull table (the region of the skull behind the eye sockets) and A. subjuniperus has a mesorostrine or "middle-snouted" skull with a snout that is longer than the skull table.[1][2] The main feature that distinguishes Allodaposuchus species 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 of Allodaposuchus is visible when the skull is viewed from the side.[3]

At least one species of Allodaposuchus, A. hulki, may have adaptations that would have allowed it to live on land for extended periods of time. A. hulki has large sinuses in 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. hulki has well-developed muscle attachments on its scapula, humerus, and ulna bones that would have allowed the forelimbs to have been held in a semi-erect stance suitable for walking over land. Remains of A. hulki come from interbedded sandstones and marls that, based on the presence of charophyte algae, likely formed in ephemeral ponds in a large floodplain far from permanent bodies of water like lakes or rivers. A. hulki may therefore have spent much of its time out of water, travelling between these ponds for food.[4]
History of study

While there are several described species of Allodaposuchus, the precise membership of the group is currently disputed.
Allodaposuchus precedens

The type species of Allodaposuchus, A. precedens, was named by Hungarian paleontologist Franz Nopcsa in 1928 from Vălioara, Romania.[5] Nopcsa found bone fragments in a deposit of the Hațeg Basin that dates back to the late Maastrichtian stage – the very end of the Late Cretaceous. Several partial skulls from Spain and France were attributed to A. precedens in 2001.[6] Some of these skulls came from Campanian-age deposits slightly older than those in Romania, meaning that the species must have persisted for about 5 million years.[1]

A 2013 study proposed that the French and Spanish fossils assigned to A. precedens in 2001 might actually represent a new unnamed species of Allodaposuchus currently identified as Allodaposuchus sp.[1] 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, making Allodaposuchus a nomen dubium or "dubious name".[7] However, the 2013 study reaffirmed the Romanian material's distinctiveness from other European Cretaceous crocodylomorphs and therefore reaffirmed the validity of Allodaposuchus as a genus.[1]
Allodaposuchus (Agaresuchus) subjuniperus ?

In 2013, a second species of Allodaposuchus, A. subjuniperus, was named on the basis of a skull from the late-Maastrichtian Conquès Formation, part of the Tremp Group, in the province of Huesca, Spain. The skull was found underneath a juniper tree whose roots had grown between the bones, hence the species name subjuniperus or "under juniper" in Latin.[1] However, in 2016, A. subjuniperus was moved to a new genus, Agaresuchus along with the type species of that genus, A. fontisensis, on the grounds that the Spanish species were sufficiently distinct from A. precedens.[8] In 2021, a phylogenetic analysis by Blanco disputed this result, suggesting that both A. fontisensis and A. subjuniperus belong within the genus Allodaposuchus proper, alongside the two species of Lohuecosuchus: L. megadontos and L. mechinorum.[9]
Allodaposuchus palustris

In 2014, A. palustris was described from a partial skull and other skeletal fragments found in Maastrichtian age sediments of the Tremp Formation in a fossil locality called Fumanya Sud in the southern Pyrenees.[10] These remains allowed for the first detailed description of the postcranial (non-skull) anatomy of Allodaposuchus.
Allodaposuchus hulki

A fourth species of Allodaposuchus, 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 the Hulk from Marvel Comics, in reference to features on the bones that suggest it had strong muscles.[4]
Allodaposuchus iberoarmoricanus

A. palustris was described by Blanco in 2021 based on fossils discovered in Late Campanian-aged fluvial deposits in Velaux-La Bastide Neuve, in Bouches-du-Rhône Department of southern France.[9] The species name is in reference to the Ibero-Armorican island of the Cretaceous European Archipelago.[9]
Allodaposuchus (Agaresuchus) fontisensis ?

In 2016, the new genus and species Agaresuchus fontisensis was discovered and described. It was named from the Lo Hueco fossil site in Fuentes, Cuenca, Spain; fontis is the Latin name of Fuentes.[8] A. subjuniperus was then also placed into the new genus Agaresuchus.[8] However, Blanco's 2021 study has called this into question, and instead proposed that they should both be considered members of Allodaposuchus, with Agaresuchus as a junior synonym.[9]
Allodaposuchus (Lohuecosuchus) megadontos and Allodaposuchus (Lohuecosuchus) mechinorum ?

The genus Lohuecosuchus was named in 2015 and contained two species, L. megadontos and L. mechinorum, from Spain and southern France.[11] However, Blanco's 2021 study has called this into question, and instead proposed that they should both be considered members of Allodaposuchus, with Lohuecosuchus as a junior synonym.[9]
Classification
jaw fragments and teeth of A. palustris
Skeletal diagram showing known remains of A. palustris

Allodaposuchus belongs to the clade Allodaposuchidae. The exact placement of Allodaposuchidae is still in dispute. Narváez et al. considered it the sister group to Hylaeochampsidae, which together form a clade that is sister to Crocodylia.[11] Other studies have alternatively recovered them not as sister taxon, but rather as an evolutionary grade towards Crocodylia, with Hylaeochampsidae more basal than Allodaposuchidae.[12][13] Alternatively, a 2021 analysis incorporating postcranial information recovered Allodaposuchidae within Crocodylia.[14]

The internal phylogeny of Allodaposuchidae can be shown in the cladogram below from the 2021 Blanco study:^[14]

Allodaposuchidae

Arenysuchus gascabadiolorum Allodaposuchus precedens Allodaposuchus iberoarmoricanus Allodaposuchus subjuniperus Allodaposuchus palustris Allodaposuchus hulki Agaresuchus fontisensis Lohuecosuchus mechinorum Lohuecosuchus megadontos

In the 2021 study, Blanco recovered Allodaposuchus as paraphyletic, with Agaresuchus and Lohuecosuchus. Accordingly, Blanco proposed that Agaresuchus and Lohuecosuchus should be considered junior synonyms of Allodaposuchus.[14]
References

Puértolas-Pascual, E.; Canudo, J.I.; Moreno-Azanza, M. (2014). "The eusuchian crocodylomorph Allodaposuchus subjuniperus sp. nov., a new species from the latest Cretaceous (upper Maastrichtian) of Spain". Historical Biology. 26 (1): 91–109. doi:10.1080/08912963.2012.763034. S2CID 85004774.
Martin, J.E. (2010). "Allodaposuchus Nopsca, 1928 (Crocodylia, Eusuchia), from the Late Cretaceous of southern France and its relationships to Alligatoroidea". Journal of Vertebrate Paleontology. 30: 756–767. doi:10.1080/02724631003758318.
Delfino, M.; Codrea, V.; Folie, A.; Dica, P.; Godefroit, P.; Smith, T. (2008). "A complete skull of Allodaposuchus precedens Nopcsa, 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.
Blanco, Alejandro; Fortuny, Josep; Vicente, Alba; Luján, Angel H.; García Marçà, Jordi Alexis; Sellés, Albert G. (2015). "A new species of Allodaposuchus (Eusuchia, Crocodylia) from the Maastrichtian (Late Cretaceous) of Spain: phylogenetic and paleobiological implications". PeerJ. 3:e1171: 1–35. doi:10.7717/peerj.1171. PMC 4558081. PMID 26339549.
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 crocodyliform Allodaposuchus precedens from 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.
Narváez, 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. doi:10.1016/j.cretres.2016.04.018.
Blanco, 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. PMC 8189472. PMID 34106925.
Blanco, Alejandro; Puértolas Pascual, Eduardo; Marmi, Josep; Vila, Bernat; Sellés, Albert G. (2014). "Allodaposuchus palustris sp. nov. from the Upper Cretaceous of Fumanya (South Eastern Pyrenees, Iberian Peninsula): Systematics, Palaeoecology and Palaeobiogeography of the Enigmatic Allodaposuchian Crocodylians". PLoS One. 9: 1–34. doi:10.1371/journal.pone.0115837.
Narváez, 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. PMC 4633049. PMID 26535893.
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. PMC 6030529. PMID 30051855.
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. PMC 8428266. PMID 34567843.
Blanco, 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. ISSN 1932-6203. PMC 8189472. PMID 34106925.