Piatnitzkysaurus
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''Piatnitzkysaurus'' ( ; meaning "Piatnitzky's lizard") is a
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
tetanuran Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, and coelurosaurs (which includes tyrannosauroids, ornithomimosaurs, compsognathids and maniraptoran ...
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
that lived approximately 179 to 177
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
during the lower part of the
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 143.1 Mya. ...
Period Period may refer to: Common uses * Period (punctuation) * Era, a length or span of time *Menstruation, commonly referred to as a "period" Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (o ...
in what is now
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 ...
. ''Piatnitzkysaurus'' was a moderately large, lightly built,
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'' ...
, 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 Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
that could grow over long.


Discovery and naming

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 of ''Piatnitzkysaurus'', PVL 4073, was collected during expeditions in 1977, 1982, and 1983 at the
Cañadón Asfalto Formation The Cañadón Asfalto Formation is a geological formation from the Lower Jurassic, with doubtful layers of Late Jurassic age previously referred to it. The Cañadón Asfalto Formation is located in the Cañadón Asfalto Basin, a rift basin in th ...
in sediments that were deposited during the Middle-Late
Toarcian The Toarcian is, in the International Commission on Stratigraphy, ICS' geologic timescale, an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic. It spans the time between 184.2 Megaannum, Ma (million ...
stage Stage, stages, or staging may refer to: Arts and media 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 Brit ...
of the
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 143.1 Mya. ...
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 Alejandro is the Spanish form of the name Alexander. Alejandro has multiple variations in different languages, including Aleksander (Czech, Polish), Alexandre ( French), Alexandros (Greek), Alsander ( Irish), Alessandro (Italian), Aleksandr (Rus ...
(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 ''Patagosaurus'' (meaning "Patagonia lizard") is an extinct genus of eusauropod dinosaur from the Middle-Late Toarcian of Patagonia, Argentina. It was first found in deposits of the Cañadón Asfalto Formation, which date to around 179 to 177&nb ...
'' (a cetiosaurid)and '' Volkheimeria chubutensis'' (an early eusauropod) 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 dinosaurs 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'' ( ;), from Ancient Greek εὖ (''eû''), meaning "well", στρεπτός (''streptós''), meaning "twisted", and σπόνδυλος (''spóndulos''), meaning "vertebra", is a genus of megalosaurid theropod dinosaur, ...
,'' and ''
Dilophosaurus ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 186 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserv ...
.'' ''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
postcrania The postcranium ("behind the cranium"; plural: postcrania) or postcranial skeleton in zoology and vertebrate paleontology is the skeleton apart from the skull. The postcranium encompasses the axial skeleton, which includes the entirety of the verte ...
l 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 (biology), isotype ...
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 (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 ...
, which is a
subadult A juvenile is an individual organism (especially an animal) 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 t ...
. It had robust arms and powerful hind legs with four toes on each foot. Its
ischium The ischium (; : is ...
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 y ...
'' from France. A general resemblance to the theropod ''
Allosaurus ''Allosaurus'' ( ) is an extinct genus of theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic period ( Kimmeridgian to late Tithonian ages). The first fossil remains that could definitively be ascribed to th ...
'' was noted by Benton (1992); however there are key differences between the two. The scapular blade of ''Piatnitzkysaurus'' is shorter and wider than more derived tetanurans. The humerus represents 50 percent of the length of the femur, which is also a primitive condition present among basal
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
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 The ischium (; : is ...
, 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''. However, detailed redescription of the appendicular anatomy of ''Piatnitzkysaurus'' revealed many differences between the two taxa.


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 to four known dinosaur genera: '' Condorraptor'', '' Marshosaurus'', '' Piatnitzkysaurus and'' possibly '' Xuanhanosaurus.'' The ...
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 (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
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 Theropoda, theropod dinosaurs, including Megalosauroidea, megalosauroids, Allosauroidea, allosauroids, and Coelurosauria, coelurosaurs (which includes Tyrannosauroi ...
. 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 The parasphenoid is a bone which can be found in the cranium of many vertebrates. It is an unpaired dermal bone which lies at the midline of the roof of the mouth. In many reptiles (including birds), it fuses to the endochondral (cartilage-derived ...
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 (geology), Period. Discovery and naming Two specimens of a troodontidae, troodontid were described in 2002 in paleo ...
'' and ''
Sinornithosaurus ''Sinornithosaurus'' (derived from a combination of Latin and Ancient Greek, Greek, meaning 'Chinese bird-lizard') is a genus of Feathered dinosaurs, feathered dromaeosaurid dinosaur from the early Cretaceous Period (geology), Period (late Barrem ...
''. Given the distant phylogenetic position of the basal tetanuran ''Piatnitzkysaurus'' and the advanced maniraptoran and deinonychosaurian taxa ''Sinovenator'' and ''Sinornithosaurus'', 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 Synapomorphy, 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 (cladistics), outgroup taxa, not ...
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 'horn' and 'lizard') is a genus of carnivorous theropod dinosaur that lived in the Late Jurassic period (Kimmeridgian to Tithonian ages). The genus was first described in 1884 by American paleontologist Othni ...
'', ''
Allosaurus ''Allosaurus'' ( ) is an extinct genus of theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic period ( Kimmeridgian to late Tithonian ages). The first fossil remains that could definitively be ascribed to th ...
'', and ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
''.


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 ...
contains '' Condorraptor'', ''
Marshosaurus ''Marshosaurus'' is a genus of medium-sized carnivorous Theropoda, theropod dinosaur, belonging to the family Piatnitzkysauridae, from the Late Jurassic Morrison Formation of Utah and possibly Colorado. Discovery and naming During the 1960s, o ...
'', ''Piatnitzkysaurus'' and '' Xuanhanosaurus''. The next most basal clade comprises ''
Chuandongocoelurus ''Chuandongocoelurus'' ( ) is a genus of carnivorous tetanuran theropod dinosaur from the Jurassic of China. Discovery and naming The type species ''Chuandongocoelurus primitivus'' was first described and named by Chinese paleontologist He ...
'' and ''
Monolophosaurus ''Monolophosaurus'' ( ; meaning "single-crested lizard") is an extinct genus of tetanuran theropod dinosaur from the Middle Jurassic Shishugou Formation in what is now Xinjiang, China.Holtz, Thomas R. Jr. (2011) ''Dinosaurs: The Most Complete, ...
''. 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


External links

{{Portal bar, Dinosaurs Piatnitzkysauridae Dinosaur genera Toarcian dinosaurs Cañadón Asfalto Formation Taxa named by José Bonaparte Fossil taxa described in 1979 Dinosaurs of Argentina