Paranasuchus
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''Paranasuchus'' is an extinct genus of large
caiman A caiman ( (also spelled cayman) from Taíno language, Taíno ''kaiman'') is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family (biology), family, the other being alligators. ...
from the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
Ituzaingó Formation The Ituzaingó Formation (), in older literature also described as Entre Ríos or Entrerriana Formation, is an extensive formation (geology), geological formation of Late Miocene (Tortonian, or Huayquerian in the South American land mammal age, SA ...
of
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
was originally described as a new species of ''Caiman'', ''C. gasparinae'', in 2013, but a study from 2024 concluded that it differed significantly enough to represent its own genus. ''Paranasuchus'' was a large caiman with a broad skull and was only one of several generalized caimans that inhabited
Paraná Paraná, Paranã or Parana may refer to: Geology * Paraná Basin, a sedimentary basin in South America Places In Argentina *Paraná, Entre Ríos, a city * Paraná Department, a part of Entre Ríos Province In Brazil *Paraná (state), a state ...
during the Late Miocene.


History and naming

The holotype of ''Paranasuchus'' consists of a partial snout (including a
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
,
maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
an other adjacent bones) as well as an associated skull table from Argentinas
Ituzaingó Formation The Ituzaingó Formation (), in older literature also described as Entre Ríos or Entrerriana Formation, is an extensive formation (geology), geological formation of Late Miocene (Tortonian, or Huayquerian in the South American land mammal age, SA ...
. Though the material had been considered to be referrable to the extant
broad-snouted caiman The broad-snouted caiman (''Caiman latirostris'') is a crocodilian in the family Alligatoridae found in eastern and central South America, including the Pantanal habitat of Bolivia, Southeast Brazil, and Paraguay, as well as northern Argentina an ...
in an unpublished
doctoral thesis A thesis (: theses), or dissertation (abbreviated diss.), is a document submitted in support of candidature for an academic degree or professional qualification presenting the author's research and findings.International Standard ISO 7144: D ...
by Zulma Brandoni de Gasparini, a much later study from 2013 highlighted that this assumption was seemingly proposed without any actual justification or evidence. The study by Paula Bona & Ariana Paulina Carabajal instead proposed that the material represented a distinct species, which they dubbed ''Caiman gasparinae''. In addition to the type material, they also assigned a fragment of a premaxilla to the species that had previously been known under the names ''Alligator? ameghinoi'' and ''Xenosuchus paranesis ameghinoi''. Nearly a decade later, ''Caiman gasparinae'' was once again reanalyzed and compared to extensive caiman material as well as other fossil crocodilians from the Ituzaingó Formation. In addition to assigning even more specimens to the species, this study showed that it was not only distinct on a species level, but actually represented an entirely new genus, which was named ''Paranasuchus gasparinae''. The name ''Paranasuchus'' translates to "Crocodile from Paraná", named so for the banks of the
Paraná River The Paraná River ( ; ; ) is a river in south-central South America, running through Brazil, Paraguay, and Argentina for some ."Parana River". Encyclopædia Britannica. Encyclopædia Britannica Online. Encyclopædia Britannica Inc., 2012. Web. ...
where the fossils had been found. This parallels the name '' Paranacaiman'', known from the same deposits and published in the same study. The species name meanwhile honors paleontologist Zulma Brandoni de Gasparini, chosen as she spent much of her life studying extinct crocodyliforms.


Description

In general morphology ''Paranasuchus'' bears close resemblance to modern caimans with a broad snout, though the bones that form the very tip are noticeably lower than in modern forms. Looking at the skull from above shows that it possesses sinuous margins, with well-defined curves creating a wavy outline. This is especially prominent in the region where the premaxillary bones contact the
maxillae In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxillar ...
, which serves to set ''Paranasuchus'' apart from taxa like ''
Mourasuchus ''Mourasuchus'' is an extinct genus of giant, aberrant caiman from the Miocene of South America. Its skull has been described as duck-like, being broad, flat, and very elongate, superficially resembling '' Stomatosuchus'' from the Late Cretace ...
'', ''
Purussaurus ''Purussaurus'' is an extinct genus of giant caiman that lived in South America during the Miocene epoch, from the Friasian to the Huayquerian in the SALMA classification. It is known from skull material found in the Brazilian and Peruvian Am ...
'', '' Acresuchus'' and even the extant
broad-snouted caiman The broad-snouted caiman (''Caiman latirostris'') is a crocodilian in the family Alligatoridae found in eastern and central South America, including the Pantanal habitat of Bolivia, Southeast Brazil, and Paraguay, as well as northern Argentina an ...
, all of which have much less pronounced "festooning". The external nares face upward and are described as rounded, set apart from those of ''Mourasuchus'' by being not as wide and from those of ''Purussaurus'' due to not being as elongated. They also differ from those of modern caimans, in which the nares are shifted further forward. The nares are entirely surrounded by the premaxillary bones, preventing the
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
s from extending into the opening as they do in modern caimans and '' Acresuchus''. A major feature that separates ''Paranasuchus'' from '' Paranacaiman'' can be observed in the region where the prefrontals contact the
frontal bone In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
. In ''Paranacaiman'' this region bears a prominent V-shaped shelf which is absent in ''Paranasuchus''. The part of the frontal bone known as the interorbital bridge, the section located between the two eyesockets, is wider than it is long, something otherwise only seen in species of ''Purussaurus'' amongst caimans. Unlike in ''Paranacaiman'', the edges of the
skull table The skull roof or the roofing bones of the skull are a set of bones covering the brain, eyes and nostrils in bony fishes, including land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...
converge with each other toward the front of the skull and the supraoccipital bone is not nearly as sunken in, but rather aligns well with the inclination of the neighbouring squamosals. The
supratemporal fenestra Temporal fenestrae are openings in the temporal region of the skull of some amniotes, behind the orbit (eye socket). These openings have historically been used to track the evolution and affinities of reptiles. Temporal fenestrae are commonly (al ...
, two openings in the skull roof, are not quite circular in shape rather than elongated. The ornamentation of the skull roof, which is the pattern of pits in the bone's surface, is another feature that distinguishes ''Paranasuchus'' from contemporary caimans. Specifically, in ''Paranasuchus'' the ornamentation primarily consists of interconnected ridges that create a pattern of irregular and mostly incomplete cells, whereas in ''Paranacaiman'' the ornamentation is best described as a pattern of very well-defined cells and rounded pits.


Phylogeny

Although initially described as a species of ''Caiman'', later studies have repeatedly shown that ''Paranasuchus'' was only distantly related to the modern genus. One example of this can be found in the 2021 study by Jonathan P. Rio and Phillip D. Mannion, who recovered ''Paranasuchus'' (at that point still known as ''Caiman gasparinae'') outside of the clade
Jacarea Jacarea is a clade of caimans within the subfamily Caimaninae. Jacarea was first named by Norell in 1988 to include the extant species within the genera ''Caiman'' and ''Melanosuchus'', while excluding the dwarf caiman genus '' Paleosuchus''. I ...
and more closely related to the flat-snouted caiman ''Mourasuchus'' alongside the holotype of ''Paranacaiman'' (then under the name ''Caiman lutescens''). The 2024 study by Bona and colleagues is very similar in regards to the closest relatives of the animal, although the clade's position among caimans is shifted. Like in the study by Rio and Mannion, ''Paranasuchus'' was found to be most closely related to ''Paranacaiman'' as well as the various species of ''Mourasuchus'', with which they form a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
group. However, in the more recent study these animals further clade with other large-bodied caimans of the Miocene, ''
Purussaurus ''Purussaurus'' is an extinct genus of giant caiman that lived in South America during the Miocene epoch, from the Friasian to the Huayquerian in the SALMA classification. It is known from skull material found in the Brazilian and Peruvian Am ...
'' and '' Acresuchus'', with the latter two thus falling outside of Jacarea.


Paleobiology

Fossils of ''Paranasuchus'' have only been recovered from Argentina's
Ituzaingó Formation The Ituzaingó Formation (), in older literature also described as Entre Ríos or Entrerriana Formation, is an extensive formation (geology), geological formation of Late Miocene (Tortonian, or Huayquerian in the South American land mammal age, SA ...
. Like other localities that preserve a glimpse into the Miocene fauna of South America, the Ituzaingó Formation is rich in crocodilian fossils. In addition to ''Paranasuchus'' this includes the large-bodied caiman ''Paranacaiman'', the flat-headed ''Mourasuchus'', '' Caiman australis'' as well as the
gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family (biology), family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males ...
''
Gryposuchus ''Gryposuchus'' is an extinct genus of gavialid crocodilian. Fossils have been found from Argentina, Colombia, Venezuela, Brazil and the Peruvian Amazon. The genus existed during the Miocene epoch (Colhuehuapian to Huayquerian). One recently d ...
''. The presence of the enormous ''
Purussaurus ''Purussaurus'' is an extinct genus of giant caiman that lived in South America during the Miocene epoch, from the Friasian to the Huayquerian in the SALMA classification. It is known from skull material found in the Brazilian and Peruvian Am ...
'' has also been reported in older publications, but in their 2024 study Bona and colleagues cast doubt over this identification given the very limited material. What sets this southern crocodilian fauna apart from those found further north, such as those of the
Urumaco Formation The Urumaco Formation is a formation in Venezuela that includes deposits from the Late Miocene. It is the site of several "giant forms": the turtles, crocodiles, sloths and rodents of Urumaco are among the largest of their groups. Location The ...
in
Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...
and
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
's
Solimões Formation The Pebas Formation is a lithostratigraphic unit of Miocene age, found in western Amazonia. The formation extends over , including parts of Brazil, Peru, Ecuador and Colombia.Wesselingh et al., 2006 It is interpreted as representing the deposit ...
, is the fact that Ituzaingó preserves a much higher amount of generalist caimans (''Paranacaiman'', ''Paranasuchus'' and ''Caiman australis'') while the northern localities preserve a diverse assemblage of smaller hard-prey specialists.


References

{{Taxonbar, from=Q130045137 Caiman Miocene crocodylomorphs Miocene reptiles of South America Neogene Argentina Fossils of Argentina Fossil taxa described in 2024