Neovenator

Neovenator (nee-o-ven-a-tor) which means "new hunter" is a genus of allosauroid dinosaur. At the time of its discovery on the Isle of Wight, United Kingdom, it was the best-known large carnivorous dinosaur from the Early Cretaceous (Hauterivian-Barremian) of Europe.[1]

Neovenator
Temporal range: Hauterivian-Barremian
~130–125 Ma
Megalosaurus, World Museum Liverpool (2)
Reconstructed skeleton, World Museum Liverpool
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Family: Neovenatoridae
Genus: Neovenator
Hutt, Martill & Barker, 1996
Species:
N. salerii
Binomial name
Neovenator salerii
Hutt, Martill & Barker, 1996

Description

Neovenator
Restoration

Neovenator measured approximately 7.6 metres (25 ft) in length, and was of a gracile build, weighing 1,000 to 2,000 kilograms (2,200 to 4,400 lb).[2] Specimen MIWG 4199 indicates an individual with a possible length of about 10 metres (33 ft), but it only consists of a toe phalanx and its position in Neovenator is dubious.[3][4]

The various scientific descriptions of Neovenator have indicated some distinguishing traits. The nostril is twice as long as it is high. The praemaxilla in the snout bears five teeth. The maxilla is pierced by a large maxillary fenestra, the diameter of which equals a sixth of the length of the tooth row. The tooth crown equals a quarter of the tooth length, thus including the root. The toe claws have a groove on top.[5] Both praemaxillae are connected by an extra pen-in-socket connection.[6] The front joint surface of the intercentrum of the axis, the second neck vertebra, is transversely widened. The odontoid process of the axis has small openings along the side edge of the front facet. The neural process of the axis has a single small opening in the side. The rear neck vertebrae are fused with their neck ribs. On the eighth and ninth neck vertebrae, at the parapophysis, the lower rib joint facet, the internal camellate structure of the bone is visible. At the front neck vertebrae the undersides are formed as sharp keels which are not inset from the lateral sides. At the front back vertebrae, the hypapophyses, the lower swellings of the front facet edges, are formed like low mounds. On the rear back vertebrae the facets of the joint processes are continued sideways as curved flanges. The shoulder joint is wider transversely than long, measured from the front to the rear. The notch on the underside of the front blade of the ilium has a shelf at the inner side. The "feet" of the ischia are connected at their fronts but diverge at their rears. The head of the thighbone is obliquely directed to the front, to above and to the inside. On the thighbone the lesser trochanter has a robust ridge on its outer side. On the thighbone the fourth trochanter has a depression in the form of a thumbprint located to the outside of its upper limit. The front underside of the thighbone is nearly flat, only showing a short vertical groove between the lower condyles. The lower shinbone shows an oval rough area at the inner side. The top of the outer malleolus of the shinbone is pinched from the front to the rear. The outer front bulge of the top surface of the shinbone has a spur pointing to below. In the foot, the outer side of the second metatarsal has a hollow surface to contact the third metatarsal.[7]

Neovenator Scale
Estimated size based on the holotype

Several traits that once were thought to be unique or apomorphic for Neovenator, subsequent research showed to have been shared by other theropods. The nostrils are large but not uncommonly so. Having pneumatised rear back vertebrae is normal for carcharodontosaurids. Elevated paired nasal crests are shared with Allosaurus. Denticles continuing over the tooth apex are today known from other species.[7]

In 2015, it was reported that the front of the snout of Neovenator contains a complex system of neurovascular canals, functioning as sensory organs. This trait is also known from Spinosauridae and was there explained as an adaptation for searching prey in water. It was doubted, however, whether Neovenator used its system for the same purpose.[8]

Discovery and species

Neovenator salerii dorsal vertebra
First back vertebra

The first bones of Neovenator were discovered in the summer of 1978, when a storm made part of the Grange Chine collapse. Rocks containing fossils fell to the beach of Brighstone Bay on the southwestern coast of the Isle of Wight. The rocks consisted of a plant debris bed within the variegated clays and marls of the Wessex Formation dating from the Barremian stage of the Early Cretaceous, about 125 million years ago. They were first collected by the Henwood family and shortly afterwards by geology student David Richards. Richards sent the remains to the Museum of Isle of Wight Geology and the British Museum of Natural History. In the latter institution paleontologist Alan Jack Charig determined that the bones belonged to two kinds of animal: Iguanodon and some theropod. The "Iguanodon", later referred to Mantellisaurus, generated the most interest and in the early 1980s a team was sent by the BMNH to secure more of its bones. On that occasion an additional theropod tail vertebra was discovered.

Several amateur paleontologists, among them Keith and Jenny Simmonds, now began to search for additional remains of the predator. Ultimately, the total of secured bones included the snout, teeth, a front lower jaw, most of the vertebral column, ribs, belly ribs, chevrons, the left shoulder girdle, pelvis bones and a hindlimb. These were accessioned under numbers BMNH R10001 and MIWG 6348. They equalled approximately 70% of the skeleton. In 1985, excavations undertaken by Dr Steve Hutt of the MIWG revealed two vertebrae of a second individual, specimen MIWG.5470. In 1987, Jenny Simmonds found a third skeleton, containing vertebra and pelvic bones, specimen MIWG.6352. A fourth individual is represented by specimen IWCMS 2002.186, consisting of a lower jaw, vertebrae and limb elements. In 1990 the material, then considered a possible new species of Megalosaurus, was provisionally described by Hutt. Having mistaken the ischium of MIWG 6352 for a pubic bone, Hutt suggested this specimen represented a separate species.[9]

Megalosaurus
Reconstructed skull

In 1996, Steve Hutt, David Martill and Michael Barker named and described the type species Neovenator salerii. The generic name Neovenator means "new hunter" from the Greek neo~, "new" and Latin venator, "hunter". The specific name salerii honours the land owners of the site, the Salero family. In view of the large number of individuals involved in the discovery process, it was considered improper to single out one of them as discoverer. The holotype is the skeleton accessioned as BMNH R10001 and MIWG 6348.[5]

In 1999, Hutt dedicated his (unpublished) master thesis to Neovenator.[10]

In 2008, Stephen Louis Brusatte, Roger Benson and Hutt redescribed the species in great detail.[7]

In 2014, teeth indistinguishable from those of the holotype of Neovenator were found in the Angeac lignitic bone bed, France, dating to the Barremian.[11]

Classification

At the time that it was described, by Steve Hutt, Martill and Barker in 1996, it was considered the only known allosaurid in Europe. However, further studies suggested it had more in common with the advanced carcharodontosaurid group of allosaurs, and several studies including a detailed examination of the species by Benson, Carrano and Brusatte in 2010 suggest that it is closely related to the Carcharodontosauridae (in a group called Carcharodontosauria), but is actually closer to the megaraptorans, together with them forming the family Neovenatoridae.[12] Other studies have supported Neovenator being a carcharodontosaurid, and megaraptorans being tyrannosauroids.

The cladogram below follows the 2010 analysis by Benson, Carrano and Brusatte.[12]

Neovenatoridae

Neovenator Neovenator

unnamed

Chilantaisaurus

Megaraptora
unnamed

Siats meekerorum[13]

Australovenator Australovenator reconstruction

?Rapator

Fukuiraptor

unnamed

?Orkoraptor

Aerosteon

Megaraptor

Cladogram after Novas et al., 2013[14]

Allosaurus Allosaurus Revised

Carcharodontosauridae

Neovenator Neovenator

Eocarcharia

Concavenator

Acrocanthosaurus Acrocanthosaurus BW

Shaochilong Shaochilong

Carcharodontosaurinae

Carcharodontosaurus Carcharodontosaurus

Giganotosaurini

Tyrannotitan

Mapusaurus Mapusaurus BW

Giganotosaurus Giganotosaurus BW

Palaeobiology

Senses

Premaxilla and maxilla of Neovenator
Premaxilla and maxilla, with neurovascular network and CT sections

Chris Barker and colleagues suggested that Neovenator may have possessed integumentary sensory organs on its snout, much as modern waterfowl and crocodilians use to find food in muddy water, based on neurovascular structures found on the skull. As Neovenator is believed to be completely terrestrial, unlike these aforementioned modern species, it is assumed that these sensory organs were used for other purposes, such as sensitivity to pressure and temperature, controlling jaw pressure and for precise feeding. Indeed, the tooth wear for Neovenator seems to indicate that it readily avoided eating or biting into bone while it fed, further vindicating the theory. Additionally, Neovenator may have used these integumentary sensory organs in courtship and quite possibly in nurturing its young, a technique seen today in most species of crocodilians and megapode birds. Though such structures are known for another theropod, the tyrannosaurid Daspletosaurus horneri, Neovenator's neurovascular structures that likely supported these organs are the best preserved and most complete in any known theropod thus far discovered.[15][16]

Palaeopathology

The holotype of Neovenator salerii had many pathologies. The authors of the genus list them as "midcaudal vertebrae fusions, healed fractures of mid-caudal vertebra transverse processes; osteophytes affecting pedal phalanges, healed gastralia rib fractures, some forming false joints... [and] scapula fracture."[17]

Palaeoecology

Fossil remains of Neovenator have been found on the Isle of Wight off southern England, and were first discovered in the 20th century. Neovenator perhaps existed alongside other dinosaurs found in the Wessex Formation of the early Cretaceous period, such as Baryonyx, Polacanthus and Iguanodon. The holotype bones were mixed with those of the plant-eater Mantellisaurus and in the dig site also remains of fishes, amphibians, lizards, pterosaurs and Goniopholididae were present. Neovenator was one of the apex predators of its day.

References

  1. ^ Neovenator at Fossilworks.org
  2. ^ "Neovenator - paleofiles.com".
  3. ^ Dodson P., Weishampel D. B. & Osmólska H., The Dinosauria, 2nd edition (2004), University of North Carolina Press, p. 104.
  4. ^ Brusatte, S. L. and Benson, R. B. J. and Hutt, S. (2008) The osteology of Neovenator salerii (Dinosauria: Theropoda) from the Wealden Group (Barremian) of the Isle of Wight. Other. Palaeontographical Society, Palaeontographical Society Monographs 162 (631).
  5. ^ a b S. Hutt, D.M. Martill, and M.J. Barker, 1996, "The first European allosauroid dinosaur (Lower Cretaceous, Wealden Group, England)", Neues Jahrbuch für Geologie und Paläontologie Monatshefte 1996(10): 635-644
  6. ^ Naish, D., Hutt S. and Martill, D., 2001, "Saurischian dinosaurs 2: Theropods". In: Martill D. and Naish D. (eds.), Dinosaurs of the Isle of Wight The Palaeontological Association, pp. 242-309
  7. ^ a b c Brusatte S.L., Benson R.B.J., Hutt S., 2008, The osteology of Neovenator salerii (Dinosauria: Theropoda) from the Wealden Group (Barremian) of the Isle of Wight, Monograph of the Palaeontographical Society 162(631) 166 pp
  8. ^ Barker, C., Dyke, G., Naish, D., Newham, E. and Katsamenis, O., 2015, "Complex neurovascular network in the rostrum of Neovenator salerii", SVPCA 2015 abstracts, 78
  9. ^ Hutt, S., Simmonds, K. & Hullman, G., 1990, "Predatory dinosaurs from the Isle of Wight", Proceedings of the Isle of Wight Natural History and Archaeological Society, 9: 137-146
  10. ^ Hutt, S.C. 1999. Neovenator salerii: A new theropod dinosaur from the Wealden of the Isle of Wight: its status and significance for Theropod evolution. A thesis submitted for the award of degree of Master of Philosophy (unpublished). University of Portsmouth
  11. ^ Néraudeau, Didier & Allain, Ronan & Ballèvre, Michel & Batten, David & Buffetaut, Eric & Colin, Jean-Paul & Dabard, Marie Pierre & Daviero-Gomez, Véronique & El Albani, Abderrazak & Gomez, Bernard & Grosheny, D & Le Loeuff, Jean & Leprince, A & Martín-Closas, Carles & Masure, Edwige & Mazin, J.-M & Philippe, Marc & Pouech, Joane & Tong, Haiyan & Vullo, Romain. (2012). The Hauterivian-Barremian lignitic bone bed of Angeac (Charente, south-west France): Stratigraphical, palaeobiological and palaeogeographical implications. Cretaceous Research. 10.1016/j.cretres.2012.01.006.
  12. ^ a b Benson, R.B.J., Carrano, M.T and Brusatte, S.L. (2010). "A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic." Naturwissenschaften, 97:71-78 . doi:10.1007/s00114-009-0614-x
  13. ^ Zanno, L. E.; Makovicky, P. J. (2013). "Neovenatorid theropods are apex predators in the Late Cretaceous of North America". Nature Communications. 4: 2827. Bibcode:2013NatCo...4E2827Z. doi:10.1038/ncomms3827. PMID 24264527.
  14. ^ Novas, Fernando E. (2013). "Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia". Cretaceous Research. 45: 174–215. doi:10.1016/j.cretres.2013.04.001.
  15. ^ University of Southampton. "Sensitive faces helped dinosaurs eat, woo and take temperature." ScienceDaily. ScienceDaily, 27 June 2017
  16. ^ Barker, Chris Tijani; Naish, Darren; Newham, Elis; Katsamenis, Orestis L.; Dyke, Gareth (2017). "Complex neuroanatomy in the rostrum of the Isle of Wight theropod Neovenator salerii". Scientific Reports. 7 (1). Bibcode:2017NatSR...7.3749B. doi:10.1038/s41598-017-03671-3.
  17. ^ Molnar, R. E., 2001, Theropod paleopathology: a literature survey: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, p. 337-363.

External links

1996 in paleontology

Paleontology or palaeontology (from Greek: paleo, "ancient"; ontos, "being"; and logos, "knowledge") is the study of prehistoric life forms on Earth through the examination of plant and animal fossils. This includes the study of body fossils, tracks (ichnites), burrows, cast-off parts, fossilised feces (coprolites), palynomorphs and chemical residues. Because humans have encountered fossils for millennia, paleontology has a long history both before and after becoming formalized as a science. This article records significant discoveries and events related to paleontology that occurred or were published in the year 1996.

Aptian

The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous epoch or series and encompasses the time from 125.0 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), approximately. The Aptian succeeds the Barremian and precedes the Albian, all part of the Lower/Early Cretaceous.The Aptian partly overlaps the upper part of the regionally used (in Western Europe) stage Urgonian.

The Selli Event, also known as OAE1a, was one of two oceanic Anoxic events in the Cretaceous period, which occurred around 120 Ma and lasted approximately 1 to 1.3 million years. The Aptian extinction was a minor extinction event hypothesized to have occurred around 116 to 117 Ma.

Barremian

The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 125.0 ± 1.0 Ma). It is a subdivision of the Early Cretaceous epoch (or Lower Cretaceous series). It is preceded by the Hauterivian and followed by the Aptian stage.

Carcharodontosauridae

Carcharodontosaurids (from the Greek καρχαροδοντόσαυρος, carcharodontósauros: "shark-toothed lizards") were a group of carnivorous theropod dinosaurs. In 1931 Ernst Stromer named Carcharodontosauridae as a family, which, in modern paleontology, indicates a clade within Carnosauria. Carcharodontosaurids included some of the largest land predators ever known: Giganotosaurus, Mapusaurus, Carcharodontosaurus, and Tyrannotitan all rivaled or slightly exceeded Tyrannosaurus in length. A 2015 paper by Christophe Hendrickx and colleagues gives a maximum length estimate of 14 meters (46 feet) for the largest carcharodontosaurids, while the smallest carcharodontosaurids were estimated to have been at least 6 meters (20 feet) long.

Deltadromeus

Deltadromeus (meaning "delta runner") is a genus of theropod dinosaur from Northern Africa. It had long, unusually slender hind limbs for its size, suggesting that it was a swift runner. The skull is not known. One fossil specimen of a single species (D. agilis, or "agile delta runner") has been described, found in the Kem Kem Beds, which date to the mid Cretaceous Period (mid Cenomanian age), about 95 million years ago. It may be a junior synonym of the contemporary Bahariasaurus. Deltadromeus has often been considered a ceratosaurian, more specifically a member of the family Noasauridae. In 2016, a South American theropod known as Gualicho shinyae was found to possess many similarities with Deltadromeus. Depending on the phylogenetic position of Gualicho, Deltadromeus may have been a neovenatorid carnosaur, a tyrannosauroid, or a basal coelurosaur if its close relation to Gualicho is legitimate.

Dinosaur Isle

Dinosaur Isle is a purpose-built dinosaur museum located in Sandown on the Isle of Wight in southern England.The museum was designed by Isle of Wight architects Rainey Petrie Johns in the shape of a giant pterosaur. It claims to be the first custom-built dinosaur museum in Europe.

Dinosaurs of the Isle of Wight

The Isle of Wight is one of the richest dinosaur localities in Europe, with over 20 species of dinosaur having been recognised from the early Cretaceous Period (in particular between 132 and 110 million years ago), some of which were first identified on the island, as well as the contemporary non-dinosaurian species of crocodile, turtle and pterosaur.

Compton Bay, near Freshwater features dinosaur footprints which are visible at low tide.

Erectopus

Erectopus is a basal allosauroid theropod from the Lower Cretaceous of France.

Gastralium

Gastralia (singular gastralium) are dermal bones found in the ventral body wall of modern crocodilian and Sphenodon species. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In these reptiles, gastralia provide support for the abdomen and attachment sites for abdominal muscles.

Gastralia may have been derived from the ventral scales found in animals like rhipidistians, labyrinthodonts, and Acanthostega, and may be related to ventral elements of turtle plastrons.

Similar, but not homologous cartilagenous elements, are found in the ventral body walls of lizards and anurans. These structures have been referred to as inscriptional ribs, based on their alleged association with the inscriptiones tendinae (the tendons that form the six pack in humans). However, the terminology for these gastral-like structures remains confused. Both types, along with sternal ribs (ossified costal cartilages), have been referred to as abdominal ribs, a term with limited usefulness that should be avoided.

Gastralia are also present in a variety of extinct animals, including theropod and prosauropod dinosaurs, pterosaurs, plesiosaurs, champsosaurs and some primitive pelycosaurs. In dinosaurs, the elements articulate with each other in a sort of zig-zag along the midline and may have aided in respiration. Although they were thought to be present in some basal ornithischian dinosaurs, and sauropods (most notably Eobrontosaurus), the perceived occurrences have been shown to be mistaken.

Gualicho

Gualicho (named in reference to the gualichu) is a genus of theropod dinosaur. The type species is Gualicho shinyae. Gualicho lived in what is now northern Patagonia, on what was then a South American island continent split off from the supercontinent Gondwana. The fossils were found in the Huincul Formation, dating to the late Cenomanian-early Turonian age of the upper Cretaceous Period, around 93 million years ago.

List of creatures in Primeval

The following is a complete list of creatures from the universe of ITV science fiction television series Primeval and also any spin-off media, including Primeval: New World ("PNW"). The series includes various imaginary species which are not native to the series setting, with some being prehistoric and others being futuristic. Various creatures were designed with some artistic license, for dramatic effect. A number of creatures from the Walking with... series were also reimagined for dramatic effect.

In 2007 Character Options announced they would create Primeval action figures, including both a flying Rex and a large plush toy Rex, Future Predators, Hesperornis and dodos.

Megaraptora

Megaraptora is a clade of carnivorous theropod dinosaurs with elongated hand claws and controversial relations to other theropods.Megaraptorans are incompletely known, and no complete megaraptoran skeleton has been found. However, they still possessed a number of unique features. Their forelimbs were large and strongly built, and the ulna bone had a unique shape in members of the family Megaraptoridae, a subset of megaraptorans which excludes Fukuiraptor. The first two fingers were elongated, with massive curved claws, while the third finger was small. Megaraptoran skull material is very incomplete, but a juvenile Megaraptor described in 2014 preserved a portion of the snout, which was long and slender. Leg bones referred to megaraptorans were also quite slender and similar to those of coelurosaurs adapted for running. Although megaraptorans were thick-bodied theropods, their bones were heavily pneumatized, or filled with air pockets. The vertebrae, ribs, and the ilium bone of the hip were pneumatized to an extent which was very rare among theropods, only seen elsewhere in taxa such as Neovenator. Other characteristic features include opisthocoelous neck vertebrae and compsognathid-like teeth.The clade was originally named in 2010 as a subset of the family Neovenatoridae, a group of lightly-built allosauroids related to the massive carcharodontosaurids such as Giganotosaurus and Carcharodontosaurus. A 2013 phylogenetic analysis by Fernando Novas and his colleagues disagreed with this classification scheme, and instead argued that the megaraptorans evolved deep within Tyrannosauroidea, a superfamily of basal coelurosaurs including the famous Tyrannosaurus. Subsequent refinements to Novas's data and methodologies have supported a third position for the group, at the base of Coelurosauria among other controversial theropods such as Gualicho, but not within the Tyrannosauroidea. Regardless of their position, it is clear that megaraptorans experienced a large amount of convergent evolution with either Neovenator-like allosauroids or basal coelurosaurs.Megaraptorans were most diverse in the early Late Cretaceous of South America, particularly Patagonia. However, they had a widespread distribution. Fukuiraptor, the most basal ("primitive") known member of the group, lived in Japan. Megaraptoran material is also common in Australia, and the largest known predatory dinosaur from the continent, Australovenator, was a megaraptoran.

Neovenatoridae

Neovenatoridae is a family of large carnivorous dinosaurs representing a branch of the allosauroids, a large group of carnosaurs that also includes the sinraptorids, carcharodontosaurids, and allosaurids. Compared to other allosauroids, neovenatorids had short, wide shoulder blades, and their ilia (upper hip bones) had many cavities.

Shaochilong

Shaochilong (meaning "shark toothed dragon") is a genus of carcharodontosaurid dinosaur from the mid Cretaceous (Turonian stage) Ulansuhai Formation of China (about 92 million years ago). The type species, S. maortuensis, was originally named Chilantaisaurus maortuensis, but was re-described and reclassified in 2009.

Siats

Siats is an extinct genus of large neovenatorid theropod dinosaur known from the Late Cretaceous Cedar Mountain Formation of Utah, US. It contains a single species, Siats meekerorum. S. meekerorum could be the first neovenatorid discovered in North America and the geologically youngest allosauroid yet discovered from the continent. It was initially classified as a megaraptoran, a clade of large theropods with very controversial relationships. This group may be examples of tyrannosauroids, neovenatorid allosauroids, or basal coelurosaurs.

Tratayenia

Tratayenia is an extinct genus of megaraptoran theropod dinosaurs known from remains found in the Santonian-age Bajo de la Carpa Formation of Argentina. The type and only species, Tratayenia rosalesi, was described in March 2018.Tratayenia can be distinguished from other megaraptorans on the basis of three autapomorphies (unique derived features) of the front portion of each dorsal vertebra, as well as a single autapomorphy of the sacrum. Tratayenia is the youngest known genus of megaraptoran, having lived only about 83 million years ago.

Tyrannotitan

Tyrannotitan (meaning "titanic tyrant") is a genus of huge bipedal carnivorous dinosaur of the carcharodontosaurid family from the Aptian stage of the early Cretaceous period, discovered in Argentina. It is closely related to other giant predators like Carcharodontosaurus and especially Giganotosaurus as well as Mapusaurus.

Valdoraptor

Valdoraptor (meaning "Wealden plunderer") is a genus of theropod dinosaur from the Early Cretaceous. Its fossils were found in England. It is known only from bones of the feet. The holotype, BMNH R2559 (incorrectly given by Owen as BMNH R2556), was found near Cuckfield in layers of the Tunbridge Wells Sand Formation dating from the late Valanginian. The specimen is damaged lacking parts of the upper and lower ends. It has a conserved length of 215 millimetres (8.5 in) and an estimated length of 240 millimetres (9.4 in). This genus is paleontologically significant for being the first ornithomimosaur specimen known from England and represents the earliest record of ornithomimosaurs.

Wessex Formation

The Wessex Formation is a fossil-rich English geological formation that dates from the Berriasian to Barremian stages (about 145–125 million years ago) of the Early Cretaceous. It forms part of the Wealden Group and underlies the younger Vectis Formation and overlies the Durlston Formation. The dominant lithology of this unit is mudstone with some interbedded sandstones.

Basal allosauroids
Metriacanthosauridae
Allosauria

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