Proceratosaurus
Encyclopedia
Proceratosaurus is a genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of small-sized (~3 metres (9.8 ft) long) carnivorous
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...

 theropod dinosaur
Dinosaur
Dinosaurs are a diverse group of animals of the clade and superorder Dinosauria. They were the dominant terrestrial vertebrates for over 160 million years, from the late Triassic period until the end of the Cretaceous , when the Cretaceous–Paleogene extinction event led to the extinction of...

 from the Middle Jurassic
Middle Jurassic
The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from 176-161 million years ago. In European lithostratigraphy, rocks of this Middle Jurassic age are called the Dogger....

 (Bathonian
Bathonian
In the geologic timescale the Bathonian is an age or stage of the Middle Jurassic. It lasted from approximately 167.7 Ma to around 164.7 Ma...

) of England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...

. It was originally thought to be an ancestor of Ceratosaurus
Ceratosaurus
Ceratosaurus meaning "horned lizard", in reference to the horn on its nose , was a large predatory theropod dinosaur from the Late Jurassic Period , found in the Morrison Formation of North America, in Tanzania and Portugal...

, due to the similar small crest on its snout. Now, however, it is considered a coelurosaur, specifically one of the earliest known members of Tyrannosauroidea
Tyrannosauroidea
Tyrannosauroidea is a superfamily of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent beginning in the Jurassic Period...

, the clade of basal relatives of the tyrannosaurs.

The type specimen is held in the London Museum of Natural History
Natural History Museum
The Natural History Museum is one of three large museums on Exhibition Road, South Kensington, London, England . Its main frontage is on Cromwell Road...

 and was recovered in 1910
1910 in paleontology
-Dinosaurs:* AMNH 5244, a ceratopsian braincase, was found isolated during an American Museum of Natural History Barnum Brown-led expedition.-Newly named dinosaurs:...

 at Minchinhampton
Minchinhampton
Minchinhampton is an ancient market town, located on a hilltop south-south-east of Stroud, Gloucestershire, England, in the Cotswolds. The town is twinned with Nkokoto, in Tanzania....

 while excavating for a reservoir.

Classification

Arthur Smith Woodward
Arthur Smith Woodward
Sir Arthur Smith Woodward was an English palaeontologist.-Biography:Woodward was born in Macclesfield, Cheshire, England and was educated there and at Owens College, Manchester. He joined the staff of the Department of Geology at the Natural History Museum in 1882. He became assistant Keeper of...

, who initially studied Proceratosaurus, originally thought it to be an ancestor of the Late Jurassic Ceratosaurus, due to the similarity of their nasal crests. Later study during the 1930s by Friedrich von Huene
Friedrich von Huene
Friedrich von Huene was a German paleontologist who named more dinosaurs in the early 20th century than anyone else in Europe.-Biography:...

 supported this interpretation, and Huene thought both dinosaurs represented members of the group Coelurosauria
Coelurosauria
Coelurosauria is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. In the past, it was used to refer to all small theropods, although this classification has been abolished...

.

It was not until the late 1980s, after Ceratosaurus had been shown to be a much more primitive theropod and not a coelurosaur, that the classification of Proceratosaurus was again re-examined. Gregory S. Paul
Gregory S. Paul
Gregory Scott Paul is a freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dinosaurs and his detailed illustrations, both live and skeletal...

 suggested that it was a close relative of Ornitholestes
Ornitholestes
Ornitholestes was a small theropod dinosaur of the late Jurassic of Western Laurasia . To date, it is known only from a single partial skeleton, and badly crushed skull found at the Bone Cabin Quarry near Medicine Bow, Wyoming, in 1900...

, again mainly due to the crest on the nose (though the idea that Ornitholestes bore a nasal crest was later disproved). Paul considered both Proceratosaurus and Ornitholestes to be neither ceratosaurs nor coelurosaurs, but instead primitive allosauroids
Allosauroidea
Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four families — the Sinraptoridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae...

. Furthermore, Paul considered the much larger dinosaur Piveteausaurus
Piveteausaurus
Piveteausaurus 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.-History and description:...

to be the same genus as Proceratosaurus, making Piveteausaurus a junior synonym. However, no overlapping bones between the two had yet been exposed from the rock around their fossils, and future study showed that they were indeed distinct.

Several phylogenetic studies in the early 21st century finally found Proceratosaurus (as well as Ornitholestes) to be a coelurosaur, only distantly related to the ceratosaurids and allosauroids, though one opinion published in 2000 considered Proceratosaurus a ceratosaurid without presenting supporting evidence. Phylogenetic analyses by Thomas R. Holtz, Jr. in 2004 also placed Proceratosaurus among the coelurosaurs, though with only weak support, and again found an (also weakly supported) close relationship with Ornitholestes.

The first major re-evaluation of Proceratosaurus and its relationships was published in 2010 by Oliver Rauhut and colleagues. Their study concluded that Proceratosaurus was in fact a coelurosaur, and moreover a tyrannosauroid
Tyrannosauroidea
Tyrannosauroidea is a superfamily of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent beginning in the Jurassic Period...

, a member of the lineage leading to the giant tyrannosaurs of the Late Cretaceous. Furthermore, they found that Proceratosaurus was most closely related to the Chinese tyrannosauroid Guanlong
Guanlong
Guanlong was a genus of proceratosaurid tyrannosauroid dinosaur, one of the earliest known examples of the lineage.-Description and discovery:...

. They named the clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 containing these two dinosaurs the Proceratosauridae
Proceratosauridae
Proceratosauridae is a family or clade of theropod dinosaurs, probably belonging to the tyrannosaur lineage. It was first named in 2010 by Oliver Rauhut and colleagues in their re-evaluation of the type genus, Proceratosaurus...

, defined as all theropods closer to Proceratosaurus than to Tyrannosaurus, Allosaurus
Allosaurus
Allosaurus is a genus of large theropod dinosaur that lived 155 to 150 million years ago during the late Jurassic period . The name Allosaurus means "different lizard". It is derived from the Greek /allos and /sauros...

, Compsognathus
Compsognathus
Compsognathus was a small, bipedal, carnivorous theropod dinosaur. The animal was the size of a turkey and lived around 150 million years ago, the early Tithonian stage of the late Jurassic Period, in what is now Europe. Paleontologists have found two well-preserved fossils, one in Germany...

, Coelurus
Coelurus
Coelurus is a genus of coelurosaur dinosaur from the Late Jurassic period . The name means "hollow tail", referring to its hollow tail vertebrae...

, Ornithomimus
Ornithomimus
Ornithomimus is a genus of ornithomimid dinosaur from the Late Cretaceous Period of what is now North America.In 1890 Ornithomimus velox was named by Othniel Charles Marsh on the basis of a foot and partial hand from the Maastrichtian Denver Formation. Another seventeen species have been named since...

, or Deinonychus
Deinonychus
Deinonychus was a genus of carnivorous dromaeosaurid dinosaur. There is one described species, Deinonychus antirrhopus. This 3.4 meter long dinosaur lived during the early Cretaceous Period, about 115–108 million years ago . Fossils have been recovered from the U.S...

.

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