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''Riojasuchus'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of ornithosuchid
archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
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 a ...
(
Norian The Norian is a division of the Triassic geological period, Period. It has the rank of an age (geology), age (geochronology) or stage (stratigraphy), stage (chronostratigraphy). It lasted from ~227.3 to Mya (unit), million years ago. It was prec ...
) 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 ...
. Ornithosuchidae was a widespread
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of facultatively
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
pseudosuchians Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
(
crocodilian Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
-line archosaurs) with adaptations for scavenging. ''Riojasuchus'' is notable as one of the youngest and most complete members of the family. The
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...
and only known species, ''Riojasuchus tenuisceps'', was named and described by
José Bonaparte José Fernando Bonaparte (14 June 1928–18 February 2020) was an Argentine paleontologist who discovered a plethora of South American dinosaurs and mentored a new generation of Argentine paleontologists. He has been described by paleontologist ...
in 1967. It was one of the first of many well-preserved Triassic archosaurs to be discovered in Argentina. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
specimen, PVL 3827, was found in the
Los Colorados Formation The Los Colorados Formation is a sedimentary geological formation, rock formation of the Ischigualasto-Villa Unión Basin, found in the provinces of San Juan Province, Argentina, San Juan and La Rioja Province (Argentina), La Rioja in Argentina. ...
of the
Ischigualasto-Villa Unión Basin The Ischigualasto-Villa Unión Basin () is a small sedimentary basin located in the Argentine Northwest, Argentina. It is located in the southwestern part of La Rioja Province and the northeastern part of San Juan Province. The basin borders the ...
in northwestern Argentina.


Discovery and naming

The holotype specimen is PVL 3827, consisting of a complete skull and mandibles, as well as a nearly complete postcranial skeleton, and it was collected during the late 1960s. Three other specimens are known. PVL 3828 is almost as complete as the holotype, including a nearly complete skull. PVL 3826 consists of articulated vertebrae and limb fragments. PVL 3814 only involves a few fragments of vertebrae, limb bones, and osteoderms. All four specimens were found close together in the upper section of the Los Colorados Formation. The formation's diverse fossil beds have been
radiometrically dated Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares t ...
to the mid-Norian stage of the Late Triassic. The fossils are stored in the vertebrate paleontology collection (PVL) at Instituto Miguel Lillo in
San Miguel de Tucumán San Miguel de Tucumán (), usually called simply Tucumán, is the capital and largest city of Tucumán Province, located in northern Argentina from Buenos Aires. It is the fifth-largest city of Argentina after Buenos Aires, Córdoba, Argentin ...
. ''Riojasuchus'' and its fossils were initially described in papers by José Bonaparte in 1967 and 1972. The skull was redescribed by M. Belén von Baczko, Julia B. Desojo, and Denis Ponce in 2016. The postcranial skeleton was redescribed by von Baczko and Desojo in 2019.


Description

The skull of the type specimen is long, and has a large, curved snout and short mandibles.


Distinguishing characteristics

Many characteristics were identified by Bonaparte in 1969. They are listed below: *a pre
orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
al vacuity well bordered by protruding edges of the smaller preorbital opening, found in ''
Ornithosuchus ''Ornithosuchus'' (from , "bird" and , "crocodile") is an extinct genus of pseudosuchians from the Late Triassic (Carnian) Lossiemouth Sandstone of Scotland. It was originally thought to be the ancestor to the carnosaurian dinosaurs (such as ''Al ...
''; *an outgoing lateral lacrimal edge; *an infratemporal fenestra present in the shortest adult skull with mandibular fenestra; *the top of the surangular laterally pointed, and with a small prearticular process; *short atlas and cervical vertebrae, all with a ventral keel; *an ilium, pubis and femur, with the talus and calcaneus of the type of ''Ornithosuchus'' *and median orbits with a higher bottom than in ''Ornithosuchus''.


Classification

''Riojasuchus'' is a member of Ornithosuchidae, a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of facultatively
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
carnivores that were geographically widespread during the Late Triassic. Two other genera, ''
Ornithosuchus ''Ornithosuchus'' (from , "bird" and , "crocodile") is an extinct genus of pseudosuchians from the Late Triassic (Carnian) Lossiemouth Sandstone of Scotland. It was originally thought to be the ancestor to the carnosaurian dinosaurs (such as ''Al ...
'' and ''
Venaticosuchus ''Venaticosuchus'' is a genus of pseudosuchian archosaurs from the family Ornithosuchidae. Known from a single species, ''Venaticosuchus rusconii'', this genus is described based on an incomplete skull and jaw (as well as a lost partial forelimb ...
'', are currently known. The group was originally considered to be related to dinosaurs, before many phylogenetical analyses. Below is a phylogenetic cladogram by Butler ''et al.'' in 2011 showing the cladistics of
Archosauriformes Archosauriformes (Ancient Greek, Greek for 'ruling lizards', and Latin for 'form') is a clade of diapsid reptiles encompassing Archosaur, archosaurs and some of their close relatives. It was defined by Jacques Gauthier (1994) as the clade stemmin ...
, focusing mostly on
Pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
: Clade names follow Nesbitt 2011.


Paleoecology

Based on its skull morphology, ''R. tenuisceps'' is believed to have been a wading animal that used its jaws to capture amphibians, fish, and other aquatic prey. Fossils of ''Riojasuchas'' are found in the Los Colorados Formation, a Late Triassic unit in
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 ...
, and date to approximately 217 to 215 million years ago. At that age, it is the youngest ornithosuchid known. It lived alongside the
protosuchid Protosuchidae is a family of basal crocodyliform reptiles that lived from the Late Triassic to the Late Cretaceous. This family should not be confused with Protosuchia, which is an informal evolutionary grade of primitive crocodiliforms Distribu ...
''
Hemiprotosuchus ''Hemiprotosuchus'' is an extinct genus of protosuchid crocodyliform from the Late Triassic (Norian stage) Los Colorados Formation of the Ischigualasto-Villa Unión Basin in northwestern Argentina, South America. It was named in 1969 by paleontol ...
''; the sphenosuchid '' Pseudohesperosuchus''; the
stagonolepidid Aetosaurs () are heavily armored reptiles belonging to the extinct order Aetosauria (; from Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs ...
'' Neoaetosauroides''; the melanorosaurid '' Strenusaurus''; and the riojasaurid ''
Riojasaurus ''Riojasaurus'' (meaning "lizard from La Rioja") was a herbivorous sauropodomorph dinosaur named after La Rioja Province in Argentina where it was found in the Los Colorados Formation in the Ischigualasto-Villa Unión Basin by José Bonaparte. ...
''.


References

{{Portal bar, Paleontology, Reptiles, Argentina Ornithosuchidae Triassic archosaurs Norian life Late Triassic reptiles of South America Triassic Argentina Fossils of Argentina Los Colorados Formation Fossil taxa described in 1969 Taxa named by José Bonaparte Prehistoric pseudosuchian genera