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''Vivaron'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
rauisuchid Rauisuchidae is a group of large (up to or more) predatory Triassic archosaurs. There is some disagreement over which genera should be included in Rauisuchidae and which should be in the related Prestosuchidae and Poposauridae, and indeed whethe ...
known from the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
(middle
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
)
Chinle Formation The Chinle Formation is an Upper Triassic continental geological formation of fluvial, lacustrine, and palustrine to eolian deposits spread across the U.S. states of Nevada, Utah, northern Arizona, western New Mexico, and western Colorado. In Ne ...
in
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
. It is the second rauisuchid known from the
southwestern United States The Southwestern United States, also known as the American Southwest or simply the Southwest, is a geographic and cultural region of the United States that generally includes Arizona, New Mexico, and adjacent portions of California, Colorado, Ne ...
, and it highlights the wide biogeographic range similar rauisuchid taxa occupied during the Late Triassic across
Pangaea Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
, despite the varied faunal assemblages at different latitudes.


Discovery

''Vivaron'' was named in 2016 from material collected at the Hayden Quarry at
Ghost Ranch Ghost Ranch is a retreat and education center located close to the village of Abiquiú in Rio Arriba County in north central New Mexico, United States. It was the home and studio of Georgia O'Keeffe, as well as the subject of many of her painti ...
, New Mexico in 2009. The locality is part of the Chinle Formation in the Petrified Forest Member, and dates to the middle Norian ~212 Ma, possibly representing one of the youngest known rauisuchids. Prior to its description, all Late Triassic rauisuchid material from Texas, Arizona and New Mexico had been referred to ''
Postosuchus kirkpatricki ''Postosuchus'', meaning "Crocodile from Post, Texas, Post", is an extinct genus of rauisuchidae, rauisuchid reptiles comprising two species, ''P. kirkpatricki'' and ''P. alisonae'', that lived in what is now North America during the Late Triass ...
''. However, the rauisuchid remains from Hayden Quarry could be clearly distinguished from ''Postosuchus'', and was erected as a new taxon ''Vivaron haydeni''. The generic epithet was named for Vivaron, a mythical 30 foot rattlesnake spirit that was said to live under Mesa Huerfano (Orphan Mesa) at Ghost Ranch, and the specific name in honour of John Hayden, a hiker who discovered the Hayden Quarry in 2002 where the material for ''Vivaron'' was collected.


Description

''Vivaron'' is known only from three jaw bones, associated skull fragments and referred hip bones. Comparisons with related taxa allow for an estimated length of between , similar in size and appearance to other rauisuchids. ''Vivaron'' can be distinguished from other rauisuchids by the presence of two prongs on the posterior portion of the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates 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 t ...
where it articulates with the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
. It is also unique among rauisuchids by possessing five
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * ...
in the
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 mammal has b ...
, similar to the state in early
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...
but in contrast with other rauisuchids, which only have four alveoli (however, a reconstruction of a ''Postosuchus'' skull (UCMP A269) figured in Long & Murry 1995 (Fig. 121) possibly shows a fifth alveolus in the left premaxilla). ''Vivaron'' was noted to share a number of characteristics with ''
Teratosaurus ''Teratosaurus'' is a genus of rauisuchians known from the Triassic Stubensandstein (Löwenstein Formation - Norian stage) of Germany. It is estimated to be 6 meters (19.5 ft) long. Discovery In 1860, Sixt Friedrich Jakob von Kapff at th ...
'', to which it is believed to be closely related to, particularly features of the maxilla and
antorbital fossa An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, birds ...
. The referred hip elements also share similarities to the referred hips of ''Teratosaurus'', potentially supporting their referral to each taxon, despite neither being directly associated with the holotype.


Classification

''Vivaron'' was found to be a member of Rauisuchidae by Lessner ''et al.'' 2016 in their phylogenetic analysis, with no changes to the relationships of
pseudosuchians Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
found in a previous analysis (Nesbitt, 2011). The analysis recovered Rauisuchidae as monophyletic but completely unresolved, with all six species in a polytomy. Surveys of the interrelationships within Rauisuchidae found ''P. kirkpatricki'' as the sister taxon of '' Polonosuchus'', and ''
Rauisuchus ''Rauisuchus'' (meaning "Wilhelm Rau's crocodile") is a genus of extinct archosaurs which lived in what is now the Geopark of Paleorrota (Santa María Formation), Brazil, during the Late Triassic period (235–228 million years ago). It conta ...
'' as the sister taxon to the rest of Rauisuchidae. The presence of ''Vivaron'' in the southwestern United States in addition to ''Postosuchus'' has implications for the classification of numerous other rauisuchid material from these locations, as almost all of it had been referred to the single species ''Postosuchus kirkpatricki'' based on their geographic location. ''Vivaron'' demonstrates the need for more detailed morphological analysis of all rauisuchid remains in the southwestern United States when determining their assignment within the group, and not refer them to a taxon based only on geographic distribution. The inferred close relationship between ''Vivaron'' and ''Teratosaurus'' also has implications for the biogeography of middle Norian Pangaea. During the late
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...
to early Norian, very similar faunal associations were present across Pangaea, all of which included rauisuchid taxa. However, by the middle Norian faunal assemblages were becoming disparate, particularly evident between high-latitude locations in Europe (including
sauropodomorphs Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lon ...
,
phytosaurs Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalen ...
, aetosaurs and
turtles Turtles are an order (biology), order of reptiles known as Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) an ...
) and the lower-latitude southern United States (including dinosauromorphs, metoposaurs,
phytosaurs Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalen ...
, aetosaurs and non-
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
archosauromorphs Archosauromorpha (Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, liz ...
). The close relationship between ''Vivaron'' and ''Teratosaurus'' demonstrates that despite being widely geographically separated and belonging to distinct faunal assemblages, there was a clear biogeographic link between the Chinle Formation and high-latitude Pangaea, and that rauisuchids were able to maintain a widespread distribution into the middle Norian.


References

{{Taxonbar, from=Q26902505 Rauisuchids Late Triassic archosaurs of North America Chinle fauna Fossil taxa described in 2016 Late Triassic pseudosuchians Paleontology in New Mexico Prehistoric pseudosuchian genera