Piatnitzkysaurus
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''Piatnitzkysaurus'' ( meaning "Piatnitzky's lizard") is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
megalosauroid Megalosauroidea (meaning 'great/big lizard forms') is a Taxonomic rank, superfamily (or clade) of Tetanurae, tetanuran theropod dinosaurs that lived from the Middle Jurassic to the Late Cretaceous period. The group is defined as ''Megalosaurus, M ...
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
that lived approximately 179 to 177
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago) ...
during the lower part of the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
in what is now
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. ''Piatnitzkysaurus'' was a moderately large, lightly built,
bipedal Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear 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'' 'double' ...
, ground-dwelling
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
that could grow up to long.


Discovery and naming

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
specimen of ''Piatnitzkysaurus'', PVL 4073, was collected during expeditions in 1977, 1982, and 1983 at the Cañadón Asfalto Formation in sediments that were deposited during the Middle-Late
Toarcian The Toarcian is, in the ICS' geologic timescale, an age and stage in the Early or Lower Jurassic. It spans the time between 182.7 Ma (million years ago) and 174.1 Ma. It follows the Pliensbachian and is followed by the Aalenian. The Toarcian ...
stage Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * Sta ...
of the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
period, approximately 179 to 177 million years ago.Bonaparte, J. F. (1979)
Dinosaurs: a Jurassic assemblage from Patagonia.
''Science'', ''205''(4413), 1377-1379.
The specimen was very complete and is one of the best known from a Megalosaur, including a partial skull and partial anterior postcranial skeleton of a subadult individual preserved in semi-articulation.Bonaparte, J. F., & LANGE-BADRE, B. (1986)
Les Dinosaures (Carnosaures, Allosaurides, Sauropodes, Cetiosaurides) du Jurassique moyen Condor (Chubut, Argentine).
''Annales de paléontologie (1982)'', ''72''(3), 247-289.
Later in 1986, several bones of an adult individual were referred to the taxon and are the only described material besides the type. The type species, ''Piatnitzkysaurus floresi'', was described by Jose Bonaparte in 1979. It was named to honor Alejandro Matveievich Piatnitzky (1879–1959), a Russian-born Argentine geologist. In Bonaparte’s 1979 paper on the taxon, he briefly described it along with the sauropods '' Patagosaurus fariasi'' (a cetiosaurid)and '' Volkheimeria chubutensis'' (an early
eusauropod Eusauropoda (meaning "true sauropods") is a derived clade of sauropod dinosaurs. Eusauropods represent the node-based group that includes all descendant sauropods starting with the basal eusauropods of ''Shunosaurus'', and possibly ''Barapasaurus ...
) from the same deposits. It wasn’t until 1986 that Bonaparte fully described the theropod, classifying it in Allosauridae along with ''Allosaurus,
Acrocanthosaurus ''Acrocanthosaurus'' ( ; ) is a genus of carcharodontosaurid dinosaur that existed in what is now North America during the Aptian and early Albian stages of the Early Cretaceous, from 113 to 110 million years ago. Like most dinosaur genera, ' ...
,
Eustreptospondylus ''Eustreptospondylus'' ( ; meaning "true ''Streptospondylus''") is a genus of megalosaurid Theropoda, theropod dinosaur, from the Oxfordian stage of the Late Jurassic period (some time between 163 and 154 million years ago) in southern Eng ...
,'' and ''
Dilophosaurus ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 193 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserve ...
.'' ''Piatnitzkysaurus'' was later reclassified as an Abelisaurid, basal Carnosaur, basal Tetanuran, and non-Megalosaurid Megalosauroid in phylogenetic analyses, but was classified in its own family in 2012, the Piatnitzkysauridae.


Description

In total, two partial skeletons are known (a fractured skull and parts of two postcranial skeletons, among which
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...
MACN CH 895) and is the most completely known theropod from the Middle-Late Jurassic Period of the Southern Hemisphere. ''Piatnitzkysaurus'' was a relatively lightly built medium-sized bipedal carnivore that was around long and around in mass, though such estimates apply to the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
, which is a
subadult A juvenile is an individual organism that has not yet reached its adult form, sexual maturity or size. Juveniles can look very different from the adult form, particularly in colour, and may not fill the same niche as the adult form. In many o ...
. It had robust arms and powerful hind legs with four toes on each foot. Its ischium is long. Its braincase resembles that of another megalosauroid, the megalosaurid ''
Piveteausaurus ''Piveteausaurus'' (meaning " Jean Piveteau's lizard") is a genus of theropod dinosaur known from a partial skull discovered in the Middle Jurassic Marnes de Dives formation of Calvados, in northern France and lived about 164.7-161.2 million yea ...
'' from France. A general resemblance to the theropod ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
'' was noted by Benton (1992); however there are key differences between the two.Benton, M. (1992). Dinosaur and other prehistoric animal factfinder. 1st American ed. New York: Kingfisher Books. Chicago / Turabian - Author Date Citation The scapular blade of ''Piatnitzkysaurus'' is shorter and wider than more derived tenaurans. The humerus represents 50 percent of the length of the femur, which is also a primitive condition present among basal
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
s. The relative lengths of the ulna in respect to the humerus and femur are also similar to those of basal theropods, which means the forelimbs of ''Piatnitzkysaurus'' are proportionally longer than in ''Allosaurus''. Also primitive is the wide contact between the pubis and ischium, a condition that is much different than that of ''Allosaurus'' and more derived tetanurans, in which contact is reduced. The pubis of ''Piatnitzkysaurus'' also has a distal foot that is more modestly developed than in ''Allosaurus''. Due to the strong resemblance to ''Piatnitzkysaurus'', it has been suggested that the sister taxa '' Condorraptor'' could be better interpreted as the result of individual variation within the species, and not as separate taxa. The main noted differences between the two dinosaurs include both a less well developed enemial crest and a first sacral vertebra with a shallower fossa in ''Condorraptor''.


Braincase

The braincase of ''Piatnitzkysaurus'' has been reviewed in detail by Oliver Rauhut; the review constitutes one of the few detailed accounts of braincase morphology in basal theropods. ''Piatnitzkysaurus'' is the only member of
Piatnitzkysauridae Piatnitzkysauridae is an extinct family of megalosauroid or basal Allosauroid dinosaurs. It only consists of three known dinosaur genera: '' Condorraptor'', ''Marshosaurus'', and '' Piatnitzkysaurus.'' The most complete and well known member of ...
with cranial material preserved, for which two maxillae, a frontal, a braincase, and a partial dentary are known. ''Piatnitzkysaurus'' is among the most basal members of the tetanurans and is important for understanding not only Middle Jurassic
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
evolution in the Southern Hemisphere, but also for knowledge of character evolution at the base of
tetanurae Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans (including birds). Tetanurans ar ...
. The braincase of the holotype of ''Piatnitzkysaurus floresi'' (PVL 4073) is rather well preserved and shows no signs of deformation. The sutures between the individual bones of the skull are only partially visible, which indicates that the holotype represents a subadult individual. This is also consistent with the state of ossification in the post-cranial skeleton. An unusual feature of the braincase is the parasphenoid recess, which has only been described in two other non-avian theropods, ''
Sinovenator ''Sinovenator'' (meaning "Chinese hunter") is a genus of troodontid dinosaur from China. It is from the early Cretaceous Period. Discovery and naming Two specimens of a troodontid were described in 2002. They are both housed in the Institu ...
'' and ''
Sinornithosaurus ''Sinornithosaurus'' (derived from a combination of Latin and Greek, meaning 'Chinese bird-lizard') is a genus of feathered dromaeosaurid dinosaur from the early Cretaceous Period (late Barremian) of the Yixian Formation in what is now China. ...
''. Given the distant phylogenetic position of the basal tetanuran ''Piatnitzkysaurus'' and the advanced maniraptoran and deinonychosaurian taxa ''Sinovenator'' and ''Sinorntihosaurus'', the presence of this recess represents a convergence and can be considered an
autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...
of the former genus. Another noteworthy aspect of the braincase is the prominent hook-like basisphenoidal wing, a feature that is also found in other theropods such as ''
Ceratosaurus ''Ceratosaurus'' (from Greek κέρας/κέρατος, ' meaning "horn" and σαῦρος ' meaning "lizard") was a carnivorous theropod dinosaur in the Late Jurassic period ( Kimmeridgian to Tithonian). The genus was first described in 1 ...
'', ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
'', and ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosaurus'' live ...
''.


Classification

The most basal clade within
Megalosauroidea Megalosauroidea (meaning 'great/big lizard forms') is a superfamily (or clade) of tetanuran theropod dinosaurs that lived from the Middle Jurassic to the Late Cretaceous period. The group is defined as '' Megalosaurus bucklandii'' and all taxa s ...
contains '' Condorraptor'', ''
Marshosaurus ''Marshosaurus'' is a genus of medium-sized carnivorous theropod dinosaur, belonging to the Megalosauroidea, from the Late Jurassic Morrison Formation of Utah and possibly Colorado. Description ''Marshosaurus'' was medium-sized for a thero ...
'', ''Piatnitzkysaurus'' and '' Xuanhanosaurus''. The next most basal clade comprises '' Chuandongocoelurus'' and '' Monolophosaurus''. However, the affiliation of these clades with Megalosauroidea is poorly supported by tree support metrics, and it is possible that they will be classified outside of Megalosauroidea by future analyses.


References

{{Portal bar, Dinosaurs Megalosaurs Dinosaur genera Middle Jurassic dinosaurs of South America Jurassic Argentina Fossils of Argentina Cañadón Asfalto Formation Fossil taxa described in 1979 Taxa named by José Bonaparte