Neovenator | |
---|---|
Reconstructed skeleton in Japan | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Clade: | † Allosauria |
Clade: | † Carcharodontosauria |
Family: | † Neovenatoridae |
Genus: | † Neovenator Hutt, Martill & Barker, 1996 |
Species: | †N. salerii |
Binomial name | |
†Neovenator salerii Hutt, Martill & Barker, 1996 | |
Neovenator (nee-o-ven-a-tor meaning "new hunter") is a genus of carcharodontosaurian theropod dinosaur. It is known from several skeletons found in the Early Cretaceous (Hauterivian-Barremian) Wessex Formation on the south coast of the Isle of Wight, southern England. It is one of the best known theropod dinosaurs from the Early Cretaceous of Europe.
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 plant debris bed L9 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 (now Dinosaur Isle) 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 a theropod. The "Iguanodon", later referred to Mantellisaurus and ultimately made the separate genus Brighstoneus , 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 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 found by Nick Oliver is represented by specimen IWCMS 2002.186, [1] consisting of a lower jaw, parts of the cervical 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. [2]
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. [3]
In 1999, Hutt dedicated his (unpublished) master thesis to Neovenator. [4]
In 2008, Stephen Louis Brusatte, Roger Benson and Hutt redescribed the species in great detail. [5]
In 2012, teeth indistinguishable from those of the holotype of Neovenator were found in the Angeac lignitic bone bed, France, dating to the Barremian. [6]
Neovenator measured approximately 7 metres (23 ft) in length, and was of a gracile build, weighing 1 metric ton (1.1 short tons). [7] 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. [8] [9]
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. [3] Both praemaxillae are connected by an extra pen-in-socket connection. [10] 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 ilium in Neovenator is highly pneumatized [11] . 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. [5]
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. [5]
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. [12]
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. [11] Other studies have supported Neovenator being a carcharodontosaurid, and megaraptorans being tyrannosauroids.
The cladogram below follows the 2010 analysis by Benson, Carrano and Brusatte. [11]
Neovenatoridae |
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Cladogram after Novas et al., 2013 [13]
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 the 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 precision feeding. In support of this, the tooth wear for Neovenator seems to indicate that it avoided eating or biting into bone while it fed. Additionally, Neovenator might have used these integumentary sensory organs in courtship and sensing nest conditions, 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 yet discovered. [14] [15] However, a more recent study reviewing the evolution of the trigeminal canals among sauropsids notes that a much denser network of neurovascular canals in the snout and lower jaw is more commonly encountered in aquatic or semiaquatic taxa (e.g., Spinosaurus , Halszkaraptor , Plesiosaurus ), and taxa that developed a rhamphotheca (e.g., Caenagnathasia ), while terrestrial taxa such as tyrannosaurids and Neovenator may have had average facial sensitivity for non-edentulous terrestrial theropods, although further research is needed. [16]
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]
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 Ceratosuchops, Riparovenator, Baryonyx , Polacanthus , Iguanodon and Eotyrannus . The holotype bones were mixed with those of the herbivorous iguanodontian Brighstoneus and in the dig site also remains of fishes, amphibians, lizards, pterosaurs and Goniopholididae were present. Neovenator was likely the apex predator of its ecosystem.
Dinosaur Isle is a purpose-built dinosaur museum located in Sandown on the Isle of Wight in southern England.
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 sediments of the Weald Clay Formation, and became the holotype specimen of Baryonyx walkeri, named by palaeontologists Alan J. Charig and Angela C. Milner in 1986. The generic name, Baryonyx, means "heavy claw" and alludes to the animal's very large claw on the first finger; the specific name, walkeri, refers to its discoverer, amateur fossil collector William J. Walker. The holotype specimen is one of the most complete theropod skeletons from the UK, and its discovery attracted media attention. Specimens later discovered in other parts of the United Kingdom and Iberia have also been assigned to the genus, though many have since been moved to new genera.
Suchomimus is a genus of spinosaur dinosaur that lived between 125 and 112 million years ago in what is now Niger, north Africa, during the Aptian to early Albian stages of the Early Cretaceous period. It was named and described by paleontologist Paul Sereno and colleagues in 1998, based on a partial skeleton from the Elrhaz Formation. Suchomimus's long and shallow skull, similar to that of a crocodile, earns it its generic name, while the specific name Suchomimus tenerensis alludes to the locality of its first remains, the Ténéré Dese.
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, some of which were first identified on the island, as well as the contemporary non-dinosaurian species of crocodile, turtle and pterosaur.
Valdosaurus is a genus of bipedal herbivorous iguanodont ornithopod dinosaur found on the Isle of Wight and elsewhere in England, Spain and possibly also Romania. It lived during the Early Cretaceous.
Eotyrannus is a genus of tyrannosauroid theropod dinosaur hailing from the Early Cretaceous Wessex Formation beds, included in Wealden Group, located in the southwest coast of the Isle of Wight, United Kingdom. The remains (MIWG1997.550), consisting of assorted skull, axial skeleton and appendicular skeleton elements, from a juvenile or subadult, found in a plant debris clay bed, were described by Hutt et al. in early 2001. The etymology of the generic name refers to the animal's classification as an early tyrannosaur or "tyrant lizard", while the specific name honors the discoverer of the fossil.
Aristosuchus is a genus of small coelurosaurian dinosaur whose name was derived from the Greek ἄριστος and σουχος. It shared many characteristics with birds.
Calamospondylus is a genus of theropod dinosaur. It lived during the Early Cretaceous and its fossils were found on the Isle of Wight in southern England. The type species is C. oweni.
Calamosaurus was a genus of small theropod dinosaur from the Barremian-age Lower Cretaceous Wessex Formation of the Isle of Wight, England. It is based on two cervical vertebrae, collected by Reverend William Fox.
Eucamerotus was a genus of sauropod dinosaur from the Barremian-age Lower Cretaceous Wessex Formation (Wealden) of the Isle of Wight, England.
Oplosaurus was a genus of sauropod dinosaur from the Barremian-age Lower Cretaceous Wessex Formation of the Isle of Wight, England. It is known from a single tooth usually referred to the contemporaneous "wastebasket taxon" Pelorosaurus, although there is no solid evidence for this.
Ornithodesmus is a genus of small, dromaeosaurid dinosaur from the Isle of Wight in England, dating to about 125 million years ago. The name was originally assigned to a bird-like sacrum, initially believed to come from a bird and subsequently identified as a pterosaur. More complete pterosaur remains were later assigned to Ornithodesmus, until recently a detailed analysis determined that the original specimen in fact came from a small theropod, specifically a dromaeosaur, making it one of the first dromaeosaurs known to science. All pterosaurian material previously assigned to this genus has been renamed Istiodactylus.
The Wessex Formation is a fossil-rich English geological formation that dates from the Berriasian to Barremian stages 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. It is part of the strata of the Wessex Basin, exposed in both the Isle of Purbeck and the Isle of Wight. While the Purbeck sections are largely barren of vertebrate remains, the Isle of Wight sections are well known for producing the richest and most diverse fauna in Early Cretaceous Europe.
Megaraptora is a clade of carnivorous theropod dinosaurs with controversial relationships to other tetanuran theropods. Its derived members, the Megaraptoridae are noted for their large hand claws and powerfully-built forelimbs, which are usually reduced in size in other large theropods.
Ichthyovenator is a genus of spinosaurid dinosaur that lived in what is now Laos, sometime between 125 and 113 million years ago, during the Aptian stage of the Early Cretaceous period. It is known from fossils collected from the Grès supérieurs Formation of the Savannakhet Basin, the first of which were found in 2010, consisting of a partial skeleton without the skull or limbs. This specimen became the holotype of the new genus and species Ichthyovenator laosensis, and was described by palaeontologist Ronan Allain and colleagues in 2012. The generic name, meaning "fish hunter", refers to its assumed piscivorous lifestyle, while the specific name alludes to the country of Laos. In 2014, it was announced that more remains from the dig site had been recovered; these fossils included teeth, more vertebrae (backbones) and a pubic bone from the same individual.
Vectidraco, is a genus of azhdarchoid pterosaur from the Lower Cretaceous of England.
Ceratosuchops is a genus of spinosaurid from the Early Cretaceous (Barremian) of Britain.
Riparovenator is a genus of baryonychine spinosaurid dinosaur from the Early Cretaceous (Barremian) period of Britain. The genus contains a single species, Riparovenator milnerae.
Brighstoneus is a genus of hadrosauriform dinosaur from the Early Cretaceous Wessex Formation of the Isle of Wight, England. The genus contains a single species, Brighstoneus simmondsi, known from a partial skeleton.
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