Epoidesuchus Temporal range: | |
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Life Restoration | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Clade: | Crocodylomorpha |
Clade: | Crocodyliformes |
Clade: | † Notosuchia |
Clade: | † Sebecosuchia |
Clade: | † Sebecia |
Family: | † Peirosauridae |
Subfamily: | † Pepesuchinae |
Genus: | † Epoidesuchus Ruiz et al., 2024 |
Species: | †E. tavaresae |
Binomial name | |
†Epoidesuchus tavaresae Ruiz et al., 2024 | |
Epoidesuchus is an extinct pepesuchine peirosaurid from the Late Cretaceous (Campanian to Maastrichtian) Adamantina Formation of Brazil. Like other members of the Pepesuchinae and unlike the closely related members of Peirosaurinae, Epoidesuchus had long and gracile jaws, which may indicate that they were semi-aquatic animals more similar to modern crocodilians. The genus is monotypic, meaning it only contains a single species, Epoidesuchus tavaresae.
The holotype specimen of Epoidesuchus were found in 2011 in outcrops of the Adamantina Formation in the Brazilian municipality of Catanduva. The specimen; which consists of a nearly tubular snout, a partial skull roof, parts of the mandible and a single rib; was recovered from a road cut and subsequently sent to the Museu de Paleontologia Professor Antonio Celso de Arruda Campos. The material was first identified as a peirosaurid that same year by Fabiano Iori. The animal was eventually fully described in 2024 and recognized as a distinct species and genus by Juan V. Ruiz and colleagues, who coined the name Epoidesuchus tavaresae. [1]
The name Epoidesuchus is a combination of the Ancient Greek "epoide" meaning "enchanted" and "souchus", a commonly used suffix among crocodilians that harkons back to the Egyptian god Sobek. The use of "epoide" is a reference to the city of Catanduva, which is also known as the "Magic Spell City". The species name on the other hand honors Sandra Simionato Tavares, a paleontologist and the director of the museum that houses the type specimen. [1]
The skull of Epoidesuchus is known from various incomplete remains, including much of the skull table, a piece of a lacrimal and a skull fragment that preserves parts of the region that sits before the eyes. Unlike modern crocodiles, Epoidesuchus possessed a small opening in the skull before the eyes, known as the antorbital fossa. The fossa is separated from the eyesocket by the lacrimal bone, with the distance between it and the eyesocket being longer than the fossa itself is high. The skull table is dominated by two large openings, the supratemporal fenestra, the rims of which are on level with the rest of the skull roof rather than raised. The skull roof extends the furthest back with the squamosal bones, which project back and upward even after their lateral processes connect to the parietal bone. Notably, beyond this point the bones lack the ornamentation that is present across most the rest of the skull. [1]
Another largely complete element of Epoidesuchus is the left hemimandible, which preserves most of the dentary bone. The size and shape of the mandible suggests that Epoidesuchus had a long and narrow snout, with the preserved portion alone measuring 35 cm (14 in). It's roughly constant in height safe for a small increase towards the back of the skull. The dentaries are also not compressed mediolaterally but have a convex outer surface, which supports the placement of Epoidesuchus within Pepesuchinae. The edges of the lower jaw seem to run subparallel to the length of the skull as a whole and don't display noticeable narrowing towards the tip nor widening towards the back. However, there is a noticeable notch in the lower jaw close to the fourth dentary tooth, which might accommodate an enlarged maxillary tooth when the jaws were closed. [1]
The lower jaw preserves a total of 14 teeth of varying size and diameter, with the individual teeth and alveoli separated by prominent gaps that likely serve to allow the teeth of the upper and lower jaw to interlock. The distance between the individual teeth or alveoli is relatively consistent throughout the lower jaw, with the exception of the region between the sixth and ninth dentary teeth. The sixth and seventh tooth are very closely spaced, as are the eight and ninth, with prominent diastemas separating them from each other as well as the fifth and tenth dentary teeth. The eight tooth also happens to be the smallest tooth in the lower jaw, whereas the 11th and 13th are the largest. All teeth are laterally compressed, but unlike other peirosaurids they lack the serrations or crenulations that would make them ziphodont or pseudo-ziphodont, something that Epoidesuchus shares with the African Stolokrosuchus . [1]
The size of the lower jaw indicates that Epoidesuchus was at least 50% larger than other peirosaurids that inhabited the Adamantina Formation, which includes Itasuchus , Pepesuchus , Roxochampsa and Montealtosuchus . [1]
The phylogenetic analysis conducted during the description of Epoidesuchus recovers a tree that alligns with the so-called "Sebecia hypothesis", meaning that peirosaurids are found to clade with the families Sebecidae and Mahajangasuchidae. This hypothesis stands opposed to the "Sebecosuchia hypothesis", in which sebecids are most closely related to baurusuchids while peirosaurids clade with mahajangasuchids and uruguaysuchids. Within Peirosauridae, Ruiz and colleagues recovered a clear split between two distinct lineages, peirosaurines and pepesuchines. While the former are described as semi-aquatic to terrestrial animals with oreinirostral snouts (meaning their snouts appeared tall and domed) and includes taxa like Hamadasuchus , Uberabasuchus and Montealtosuchus , the later consists primarily of animals that are hypothesized to be more aquatic, possessing elongated or flattened snouts as seen in Caririsuchus , Roxochampsa and Pepesuchus . [1]
Though the name Pepesuchinae is used by Ruiz and colleagues to describe the clade, they do note that following the terminology of Pinheiro et al. 2018 Peirosauridae would be restricted to the clade they dub Peirosaurinae. Their version of Pepesuchinae meanwhile would be separated into the clade Itasuchidae (which would be composed of Pepesuchus and the recovered polytomy) and various more basal taxa. Subsequently, following this taxonomy Epoidesuchus would not be regarded as a pepesuchine nor an itasuchid, eventhough its position would not change. [1] Furthermore, the description of Epoidesuchus does not take into account the 2024 nomenclature published by Leardi and colleagues, in which they coin the clade Peirosauria, which consists of peirosaurids, itasuchids and mahajangasuchids. Within Peirosauria, Peirosauridae and Itasuchidae are roughly equivalent to the clades Peirosaurinae and Pepesuchinae as recovered by Ruiz and colleagues. [2]
Like several other members of Pepesuchinae, Epoidesuchus had elongated and narrow jaws that are wildly different from the tall, deep skulls seen in peirosaurines. Such a skull shape is unique among notosuchians, but much more widespread among neosuchians like goniopholidids and modern crocodilians, which may suggest a similar semi-aquatic lifestyle. Pepesuchines are however not perfect anatomical analogues to neosuchians, as they still have some adaptations seen in more terrestrial groups. Rather than having eyes located atop the head, as would be useful for laying in ambush with the body submerged, the eyes of Epoidesuchus faced towards the side of the skull. Considering both these similarities and differences might suggest that though more semi-aquatic than their relatives, pepesuchines such as Epoidesuchus were still less dependent on water than modern crocodilians. Semi-aquatic habits have also been reported from mahajangasuchine notosuchians, which includes Mahajangasuchus and Kaprosuchus , as well as the sebecid Lorosuchus , with the latter being much closer in anatomy to pepesuchines. This might suggest some degree of convergence between pepesuchines and Lorosuchus specifically. [1] Ruiz and colleagues note that pepesuchines often occur in units were neosuchians are rare or even absent. Epoidesuchus for example was native to Brazil's Adamantina Formation, which is part of the Bauru Group. Though the Bauru Group is renown for its diverse notosuchian fauna, only a single neosuchian has been recovered from any of the formations that are part of the unit. [1] Said neosuchian is Titanochampsa , which has so far only been found within the sediments of the Marilia Formation. [3]
Doratodon is an extinct genus of Late Cretaceous crocodylomorph that may have been a member of the Sebecosuchia. Doratodon was a relatively small animal with ziphodont teeth, meaning the teeth had flattened sides and serrated edges. Two species of Doratodon are known to science: D. carcharidens from Austria and Hungary, the type species; and D. ibericus from Spain. Teeth similar to those of Doratodon are also known from Italy and Romania, though they cannot be confidently assigned to this genus.
Mahajangasuchus is an extinct genus of crocodyliform which had blunt, conical teeth. The type species, M. insignis, lived during the Late Cretaceous; its fossils have been found in the Maevarano Formation in northern Madagascar. It was a fairly large predator, measuring up to 4 metres (13 ft) long.
Razanandrongobe is a genus of carnivorous ziphosuchian crocodyliform from the Middle Jurassic of Madagascar. It contains the type and only species Razanandrongobe sakalavae, named in 2004 by Simone Maganuco and colleagues based on isolated bones found in 2003. The remains, which included a fragment of maxilla and teeth, originated from the Bathonian-aged Sakaraha Formation of Mahajanga, Madagascar. While they clearly belonged to a member of the Archosauria, Maganuco and colleagues refrained from assigning the genus to a specific group because the fragmentary remains resembled lineages among both the theropod dinosaurs and crocodylomorphs.
Baurusuchidae is a Gondwanan family of mesoeucrocodylians that lived during the Late Cretaceous. It is a group of terrestrial hypercarnivorous crocodilians from South America and possibly Pakistan. Baurusuchidae has been, in accordance with the PhyloCode, officially defined as the least inclusive clade containing Cynodontosuchus rothi, Pissarrachampsa sera, and Baurusuchus pachecoi. Baurusuchids have been placed in the suborder Baurusuchia, and two subfamilies have been proposed: Baurusuchinae and Pissarrachampsinae.
Armadillosuchus is an extinct genus of sphagesaurid crocodylomorph. It was described in February 2009 from the late Campanian to early Maastrichtian Adamantina Formation of the Bauru Basin in Brazil, dating to approximately 70 Ma. Armadillosuchus was among the larger and more robust sphagesaurids, with a total length of approximately 2 metres (6.6 ft).
Bergisuchus is an extinct genus of small sebecosuchian mesoeucrocodylian known primarily from the Eocene Messel Pit in Germany. Few fossils of Bergisuchus have been discovered, only a single incomplete snout, a few partial lower jaws and some teeth. Despite being fragmentary, the jaw bones are enough to indicate that Bergisuchus had a short, deep, narrow snout and serrated teeth, quite unlike the broad flat snouts of modern crocodylians.
Peirosaurus is an extinct genus of peirosaurid crocodylomorph known from the Late Cretaceous period of Minas Gerais, southern Brazil. It contains a single species, Peirosaurus torminni. It is the type genus of the family Peirosauridae.
Kaprosuchus is an extinct genus of mahajangasuchid crocodyliform. It is known from a single nearly complete skull collected from the Upper Cretaceous Echkar Formation of Niger. The name means "boar crocodile" from the Greek κάπρος, kapros ("boar") and σοῦχος, soukhos ("crocodile") in reference to its unusually large caniniform teeth which resemble those of a boar. It has been nicknamed "BoarCroc" by Paul Sereno and Hans Larsson, who first described the genus in a monograph published in ZooKeys in 2009 along with other Saharan crocodyliformes such as Anatosuchus and Laganosuchus. The type species is K. saharicus.
Sebecus is an extinct genus of sebecid crocodylomorph from Eocene of South America. Like other sebecosuchians, it was entirely terrestrial and carnivorous. The genus is currently represented by two species, the type S. icaeorhinus and S. ayrampu. Several other species have been referred to Sebecus, but were later reclassified as their own genera.
Morrinhosuchus is an extinct genus of notosuchian crocodyliform from the Late Cretaceous Adamantina Formation of Brazil. It is known from a mandible and a portion of the front of the skull collected from the municipality of Monte Alto in São Paulo state. Morrinhosuchus refers to Morrinho de Santa Luzia, a hill nearby the collection site of the holotype, while luziae refers to the chapel of Santa Luzia, which is located on top of the hill.
Sebecia is an extinct clade of mesoeucrocodylian crocodyliforms that includes peirosaurids and sebecids. It was first constructed in 2007 to include Hamadasuchus, Peirosauridae, and Sebecus. It was initially considered to be the sister taxon of the clade Neosuchia, which includes living crocodilians, although some later studies have placed it within Neosuchia as a basal clade. Sebecians were terrestrial crocodyliforms characterized by their deep snouts and ziphodont dentition. They first appeared in the Late Cretaceous, survived the Cretaceous–Paleogene extinction event, and became extinct in the Miocene epoch.
Stratiotosuchus is an extinct genus of baurusuchid mesoeucrocodylian from the Adamantina Formation in Brazil. It lived during the Late Cretaceous. The first fossils were found in the 1980s, and the type species Stratiotosuchus maxhechti was named in 2001. A hyperpredator, it and other baurusuchids may have filled niches occupied elsewhere by theropod dinosaurs.
Pepesuchus is an extinct genus of carnivorous metasuchian from the Late Cretaceous period. It is a peirosaurid which lived during the Campanian and Maastrichtian stages of the Late Cretaceous in what is now state of São Paulo, Brazil. It was a semiaquatic crocodylomorph.
Campinasuchus is an extinct genus of baurusuchid mesoeucrocodylian from Minas Gerais State of Brazil.
Pissarrachampsa is an extinct genus of baurusuchid mesoeucrocodylian from the Late Cretaceous of Brazil. It is based on a nearly complete skull and a referred partial skull and lower jaw from the ?Campanian - ?Maastrichtian-age Vale do Rio do Peixe Formation of the Bauru Group, found in the vicinity of Gurinhatã, Brazil.
Caipirasuchus is an extinct genus of sphagesaurid notosuchians known from the Late Cretaceous of northern São Paulo State and western Minas Gerais State, southeastern Brazil. The type species, C. paulistanus, was named in 2011. A second species, C. montealtensis, was referred to Caipirasuchus in 2013 after having been named in 2008 as a species of Sphagesaurus. A third species, C. stenognathus, was described in 2014. A fourth species, C. mineirus, was described in 2018. A fifth species, C. attenboroughi, was named in 2021 in honour of David Attenborough, and a sixth species, C. catanduvensis, was described in 2024, with a distinct chamber associated with the airways, possibly used in vocalization.
Aplestosuchus is an extinct genus of baurusuchid mesoeucrocodylian known from the Late Cretaceous Adamantina Formation of São Paulo, southern Brazil. It contains a single species, Aplestosuchus sordidus. A. sordidus is represented by a single articulated and nearly complete skeleton, preserving the remains of an unidentified sphagesaurid crocodyliform in its abdominal cavity. The specimen represents direct evidence of predation between different taxa of crocodyliforms in the fossil record.
Aphaurosuchus is an extinct genus of baurusuchid mesoeucrocodylian known from the Late Cretaceous Bauru Basin of São Paulo, southern Brazil. It contains two species, Aphaurosuchus escharafacies and Aphaurosuchus kaiju.
Titanochampsa is a genus of large mesoeucrocodylian from the Maastrichtian Marilia Formation of Brazil. Although only known from a single skull roof, the material shows that Titanochampsa was not a member of Notosuchia, which were previously believed to have been the only crocodyliforms present in the strata of the Bauru Group. Body size estimates vary greatly and range between 2.98–5.88 m due to the incomplete nature of the holotype fossil. The overall anatomy of the skull roof, alongside its size and possible affinities with Neosuchians, may suggest that it was a semi-aquatic ambush hunter similar to modern crocodilians.
Varanosuchus is an extinct genus of atoposaurid neosuchian from the Early Cretaceous Sao Khua Formation of Thailand. Varanosuchus is known from three individuals which preserve assorted postcranial material and a complete skull. The skull of Varanosuchus was altirostral, meaning it wasn't flattened like in modern crocodilians and instead much deeper, while the limbs were slender and straight, leading to it somewhat resembling a monitor lizard. Little is known about the ecology of atoposaurids, however, based on the slender, erect limbs, the altirostral skull and the well ornamented osteoderms it has been suggested that Varanosuchus was a terrestrial animal with some semi-aquatic affinities. Only a single species is recognized, Varanosuchus sakonnakhonensis.