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Piatnitzkysauridae is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
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 ...
or basal
Allosauroid Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroidea, were among th ...
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 ...
s. It only consists of three known dinosaur genera: '' 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 ...
'', and ''
Piatnitzkysaurus ''Piatnitzkysaurus'' ( meaning "Piatnitzky's lizard") is a genus of megalosauroid theropod dinosaur that lived approximately 179 to 177 million years ago during the lower part of the Jurassic Period in what is now Argentina. ''Piatnitzkysaurus'' ...
.'' The most complete and well known member of this family is ''Piatnitzkysaurus'', which also gives the family its name.


Description

So far all known piatnitzkysaurids have only been found in
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), 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 J ...
deposits of the
western hemisphere The Western Hemisphere is the half of the planet Earth that lies west of the prime meridian (which crosses Greenwich, London, United Kingdom) and east of the antimeridian. The other half is called the Eastern Hemisphere. Politically, the term We ...
/
New world The term ''New World'' is often used to mean the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxford University Press, p. 3 ...
. ''Piatnitzkysaurus'' and ''Condorraptor'' hail from the
Cañadón Asfalto Formation The Cañadón Asfalto Formation is a Lower Jurassic to Late Jurassic geologic formation, from the Jurassic period of the Mesozoic Era. Its age is controversial, uranium-lead dating of the volcanic tuff beds having given various different ages. A ...
of
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 ...
, which has been dated from the
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 ...
epoch of the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
to the
Bathonian In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age. Strat ...
epoch of the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
(approximately 179 to 168 million years ago). ''Marshosaurus'' was found in the
Morrison Formation The Morrison Formation is a distinctive sequence of Late Jurassic, Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandsto ...
of the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
, which was dated to the
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
epoch of the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
. Piatnitzkysaurids were among the first large theropods present in South America, and are evidence for a radiation of basal tetanurans in the middle Jurassic. Although there is support for basal megalosauroid affinities, piatnitzkysaurids also share some traits with
allosaurids Allosauridae is a family of medium to large bipedal, carnivorous allosauroid theropod dinosaurs from the Late Jurassic. Allosauridae is a fairly old taxonomic group, having been first named by the American paleontologist Othniel Charles Marsh i ...
, such as a 'sigmoidal' humerus. It has been proposed that they are basal Allosaurs or Non Megalosaurian Megalosauroids. If Piatnitzkysaurids are Megalosauroids, they can be distinguished from other megalosauroids and placed into their own category due to the following
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
(distinguishing features): *
Nutrient foramina The nutrient artery (arteria nutricia, or medullary), usually accompanied by one or two veins, enters the bone through the nutrient foramen, runs obliquely through the cortex, sends branches upward and downward to the bone marrow, which ramify in ...
on the surface 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 ...
which are arranged in two parallel rows (also found in ''
Eocarcharia ''Eocarcharia'' (meaning "dawn shark") is a genus of carcharodontosaurid theropod dinosaur from the Early Cretaceous Elrhaz Formation that lived in the Sahara 112 million years ago, in what today is the country of Niger. It was discovered in 2 ...
'' and ''
Shaochilong ''Shaochilong'' (meaning "shark toothed dragon") is an extinct genus of carcharodontosaurid dinosaur from the mid-Cretaceous (Late Cenomanian to the end of the Turonian stage) Ulansuhai Formation of China (about 92 million years ago). The type ...
''). * Vertically striated or ridged paradental plates (also found in
Abelisauridae Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are fou ...
, ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years ...
'' and ''
Proceratosaurus ''Proceratosaurus'' is a genus of carnivorous theropod dinosaur from the Middle Jurassic ( Bathonian) of England. ''Proceratosaurus'' was a small dinosaur, measuring in length and in body mass.Holtz, Thomas R. Jr. (2008) ''Dinosaurs: The Mos ...
).'' * Reduced parapophyses on the
axis An axis (plural ''axes'') is an imaginary line around which an object rotates or is symmetrical. Axis may also refer to: Mathematics * Axis of rotation: see rotation around a fixed axis * Axis (mathematics), a designator for a Cartesian-coordinat ...
(also found in
Coelophysoidea Coelophysoidea were common dinosaurs of the Late Triassic and Early Jurassic periods. They were widespread geographically, probably living on all continents. Coelophysoids were all slender, carnivorous forms with a superficial similarity to the ...
, ''
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 ''
Afrovenator ''Afrovenator'' (; "African hunter") is a genus of megalosaurid theropod dinosaur from the Middle Jurassic Period of northern Africa. Discovery and naming The remains of ''Afrovenator'' were discovered in 1993 in the Tiourarén Formation ...
''). * Anteriorly inclined posterior dorsal neural spines (in parallel with
Allosauroidea Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four family (biology), families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroid ...
). * A
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
with a canted (bent forward, away from the body) distal condyle (also in ''
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 ...
'', ''
Poekilopleuron ''Poekilopleuron'' (meaning "varied ribs") is a genus of tetanuran dinosaur, which lived during the middle Bathonian of the Jurassic, about 168 to 166 million years ago. The genus has been used under many different spelling variants, although on ...
'',
Allosauridae Allosauridae is a family of medium to large bipedal, carnivorous allosauroid theropod dinosaurs from the Late Jurassic. Allosauridae is a fairly old taxonomic group, having been first named by the American paleontologist Othniel Charles Marsh in ...
and ''
Fukuiraptor ''Fukuiraptor'' ("thief of Fukui") was a medium-sized megaraptoran theropod dinosaur of the Early Cretaceous epoch (either Barremian or Aptian) that lived in what is now Japan. ''Fukuiraptor'' is known from the Kitadani Formation and possibly ...
''). This form has also been described as 'sigmoidal' (S-shaped). In addition, this family shows several reversals to basal
Tetanuran 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 ...
traits. These include: * A short or absent anterior process of the maxilla (also in
Metriacanthosauridae Metriacanthosauridae is an extinct family of allosauroid theropod dinosaurs that lived from the Middle Jurassic to the Early Cretaceous. When broken down into its Greek roots, it means "moderately-spined lizards". The family is split into two su ...
and
Carcharodontosauridae Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
). * Moderately developed (rather than reduced or absent) diapophyses on the axis. * No
pleurocoels Skeletal pneumaticity is the presence of air spaces within bones. It is generally produced during development by excavation of bone by pneumatic diverticula (air sacs) from an air-filled space, such as the lungs or nasal cavity. Pneumatization is h ...
on the axis.


Classification

Piatnitzkysauridae is defined as all members of
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 sha ...
more closely related to ''
Piatnitzkysaurus ''Piatnitzkysaurus'' ( meaning "Piatnitzky's lizard") is a genus of megalosauroid theropod dinosaur that lived approximately 179 to 177 million years ago during the lower part of the Jurassic Period in what is now Argentina. ''Piatnitzkysaurus'' ...
'' than to either ''
Spinosaurus ''Spinosaurus'' (; ) is a genus of spinosaurid dinosaur that lived in what now is North Africa during the Cenomanian to upper Turonian stages of the Late Cretaceous period, about 99 to 93.5 million years ago. The genus was known first fro ...
'' or ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years ...
''. ''Condorraptor, Marshosaurus, and Piatnitzkysaurus'' were all placed in a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
by Benson in 2010 during a redescription of ''Megalosaurus''. This clade was defined as a family by Carrano ''et al'' in 2012''.'' Piatnitzkysauridae has been recovered as the sister taxa to
Megalosauria 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 ...
, a clade containing the megalosaurids (such as ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years ...
'' and ''
Torvosaurus ''Torvosaurus'' () is a genus of carnivorous megalosaurid theropod dinosaur that lived approximately 165 to 148 million years ago during the late Middle and Late Jurassic period (Callovian to Tithonian stages) in what is now Colorado, Portuga ...
'') and the
spinosaurids The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
(such as ''
Spinosaurus ''Spinosaurus'' (; ) is a genus of spinosaurid dinosaur that lived in what now is North Africa during the Cenomanian to upper Turonian stages of the Late Cretaceous period, about 99 to 93.5 million years ago. The genus was known first fro ...
'' and ''
Baryonyx ''Baryonyx'' () is a genus of theropod dinosaur which lived in the Barremian stage of the Early Cretaceous period, about 130–125 million years ago. The first skeleton was discovered in 1983 in the Smokejack Clay Pit, of Surrey, England, in se ...
''). ''Condorraptor'' and ''Piatnitzkysaurus'' are generally grouped together within this family in a clade excluding ''Marshosaurus''. This is not only due to ''Condorraptor'' and ''Piatnitzkysaurus'' being from the same locality and time period, but also due to them having flat anterior faces of presacral vertebrae as well as a pronounced ventral keel on the anterior dorsal vertebrae. 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''. '' Xuanhanosaurus'', a problematic tetanuran with uncertain affinities, was placed as the sister taxon of these three genera when their relation was first observed in 2010. However, due to the incompleteness of ''Xuanhanosaurus's'' remains, this placement is considered uncertain. Other studies have considered ''Xuanhanosaurus'' a basal tetanuran. Xuanhanosaurus is still sometimes found as a Piatnitzkysauridae. In 2019, Rauhut and Pol described ''
Asfaltovenator vialidadi ''Asfaltovenator'' (meaning "Cañadón Asfalto Formation hunter" after the fossil formation in which its fossils were found) is a genus of possibly allosauroid dinosaur from the Lower Jurassic Cañadón Asfalto Formation from Chubut Province, Arge ...
'', a basal allosauroid displaying a mosaic of primitive and derived features seen within
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 ...
. Their phylogenetic analysis found traditional Megalosauroidea to represent a basal
grade Grade most commonly refers to: * Grade (education), a measurement of a student's performance * Grade, the number of the year a student has reached in a given educational stage * Grade (slope), the steepness of a slope Grade or grading may also ref ...
of carnosaurs,
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
with respect to
Allosauroidea Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four family (biology), families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroid ...
. Because the authors amended the definition of Allosauroidea to include all theropods that are closer to '' Allosaurus fragilis'' than to either '' Megalosaurus bucklandii'' or
Neornithes Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
, the Piatnitzkysauridae was found to fall within Allosauroidea. A cladogram displaying the relationships they recovered is shown below:


References

{{Taxonbar, from=Q5837348 Prehistoric tetanurans Late Jurassic first appearances Late Jurassic extinctions Prehistoric dinosaur families