Megalosauridae is a monophyletic family of carnivorous theropod dinosaurs within the order Megalosauroidea, closely related to the family Spinosauridae. Some members of this family include Megalosaurus, Torvosaurus, Eustreptospondylus, and Afrovenator. Appearing in the Middle Jurassic, megalosaurids were among the first major radiation of large theropod dinosaurs, although they became extinct by the end of the Jurassic period. They were a relatively primitive group of basal tetanurans containing two main subfamilies, Megalosaurinae and Afrovenatorinae, along with the basal genus Eustreptospondylus, an unresolved taxon which differs from both subfamilies.
The defining megalosaurid is Megalosaurus bucklandii, first named and described in 1824 by William Buckland after multiple finds in Stonesfield, Oxfordshire, UK. Megalosaurus was the first formally described dinosaur and was the basis for the establishment of the clade Dinosauria. It is also one of the largest known Middle Jurassic carnivorous dinosaurs, with the best-preserved femur at 805 mm and a proposed body mass of around 943 kg. Megalosauridae is recognized as a mainly European group of dinosaurs, based on fossils found in France and the UK. However, recent discoveries in Niger have led some to consider the range of the family. Megalosaurids appeared right before the split of the supercontinent Pangaea into Gondwana and Laurasia. These large theropods therefore may have dominated both halves of the world during the Jurassic.
The family Megalosauridae was first defined by Thomas Huxley in 1869, yet it has been contested throughout history due to its role as a ‘waste-basket’ for many partially described dinosaurs or unidentified remains. In the early years of paleontology, most large theropods were grouped together and up to 48 species were included in the clade Megalosauria, the basal clade of Megalosauridae. Over time, most of these taxa were placed in other clades and the parameters of Megalosauridae were narrowed significantly. However, some controversy remains over whether Megalosauridae should be considered its own distinct group, and dinosaurs in this family remain some of the most problematic taxa in all Dinosauria. Some paleontologists, such as Paul Sereno in 2005, have disregarded the group due to its shaky foundation and lack of clarified phylogeny. However, recent research by Carrano, Benson, and Sampson has systematically analyzed all basal tetanurans and determined that Megalosauridae should exist as its own family.
|Skeletal mount of Torvosaurus tanneri, Museum of Ancient Life|
Torvosauridae Jensen, 1985
Like other tetanurans, megalosaurids are carnivorous theropods characterized by large size and bipedalism. Specifically, megalosaurids exhibit especially giant size, with some members of the family weighing more than one tonne. Over time, there is evidence of size increase within the family. Basal megalosaurids from the Early Jurassic had smaller body size than those appearing in the late Middle Jurassic. Due to this size increase over time, Megalosauridae appear to follow a size increase pattern similar to that of other giant sized theropods like Spinosauridae. This pattern follows Cope's Rule, the postulation by paleontologist Edward Cope about evolutionary increase in body size.
One unambiguous synapomorphy of Megalosauridae is a lower and longer skull with a length to height ratio of 3:1. In addition, the typical skull roof tends to be much less ornamented than that of other tetanurans, and crests or horns are either very small or absent entirely. Megalosaurids also have femoral heads with an orientation 45 degrees between anteromedial and fully medial. Megalosauridae are also defined by the following unique unambiguous synapomorphies:
Megalosaurinae (all megalosaurids more closely related to Megalosaurus than Afrovenator) are characterized by a moderate (0.5-2.0) height/length ratio of the premaxilla below the level of the nares, compared to other megalosaurids which have a lower ratio and thus less tall snout tip.
Afrovenatorinae (all megalosaurids more closely related to Afrovenator than Megalosaurus) are characterized by a squared anterior margin of the antorbital fossa and the puboischiadic plate being broadly open along the midline.
Dental findings are frequently used to differentiate between various theropods and to further inform cladistic phylogeny. Tooth morphology and dental evolutionary markers are prone to homoplasy and disappear or reappear throughout history. However, megalosaurids have several specific denture conditions that differentiate them from other basal theropods. One dental condition present in Megalosauridae is multiple enamel wrinkles near the carinae, the sharp edge or serration row of the tooth. Ornamented teeth and a well-marked enamel surface also characterize basal megalosaurids. The ornamentation and well-marked surface appears in early megalosaurids but disappears in derived megalosaurids, suggesting that the condition was lost over time as megalosaurids grew in size.
From the family's inception, many specimens found in the field have been wrongly classified as megalosaurids. For example, most large carnivores found for about a century after the naming of Megalosaurus bucklandii were placed in Megalosauridae. Megalosaurus was the first paleontological finding of its kind when William Buckland discovered a giant femur and named it in 1824, predating even the term Dinosauria. When initially defined, the species M. bucklandii was anatomically based on various dissociated bones found in quarries around the village of Stonesfield, UK. Some of these early findings included a right dentary with a well-preserved tooth, ribs, pelvic bones, and sacral vertebrae. As early paleontologists and researchers found more dinosaur bones in the surrounding area, they attributed them all to M. bucklandii since it was the only named and described dinosaur at this point in history. Therefore, the species was initially described and classified by a mass of possibly unrelated characteristics.
Modern paleontology first began to approach the problematic cladistic separation of Megalosauridae during the early 20th century. Fredrich von Huene separated carnivorous theropods, which had all been grouped into the broad category of megalosaurids, into two distinct families of larger, more giant sized and smaller, more lightly built theropods. These two groups were named Coelurosauria and Pachypodosauria respectively. Later on, Huene distinguished between carnivorous and herbivorous dinosaurs in Pachypodosauria, placing the meat-eaters in a new group Carnosauria.
As more information was uncovered about basal theropods and phylogenetic characteristics, modern paleontologists began to question the proper naming for this group. In 2005 paleontologist Paul Sereno rejected the use of the clade Megalosauridae due to its ambiguous early history in favor of the name Torvosauridae. Today, it is accepted that megalosaurids existed at least as a group of basal tetanurans, due to the fact that they have more derived taxa than ceratosaurs and that the name Megalosauridae should represent this group. Megalosauridae also has priority over Torvosauridae under ICZN rules governing family names.
Megalosauridae was first phylogenetically defined in 1869 by Thomas Huxley, yet was used as a ‘waste-basket’ clade for many years. In 2002, Ronan Allain redefined the clade after he discovered a complete megalosaurid skull in northwestern France of species Poekilopleuron. Using the characters described in this study, Allain defined Megalosauridae as dinosaurs including Poekilopleuron valesdunesis, now known as Dubreuillosaurus, Torvosaurus, Afrovenator, and all descendants of their common ancestor. Allain also defined two taxa within Megalosauridae: Torvosaurinae was defined as all Megalosauridae more closely related to Torvosaurus than to Poekilopleuron and Afrovenator, and Megalosaurinae was defined as all those that are more closely related to Poekilopleuron. Megalosauridae also falls under the basal clade Megalosauroidea, which also contains Spinosauridae. However, many taxa are still quite unstable and cannot be placed in one clade with absolute certainty. For example, Eustreptospondylus and Streptospondylus, while they are both defined as Megalosauridae, are often excluded to make more stable cladograms since they are not defined to a certain subgroup. The cladogram presented here follows Benson (2010) and Benson et al. (2010).
Then, in 2012, Carrano, Benson, and Sampson did a much larger analysis of tetanurans and defined Megalosauria more broadly as the clade containing Megalosaurus, Spinosaurus, and all its descendants. In other words, Megalosauria is the group that contains the two families Megalosauridae and its close relative Spinosauridae. Within this new cladogram, Megalosauridae was given a new subfamily Afrovenatorinae, which included all megalosaurids more closely related to Afrovenator than Megalosaurus.
Carrano, Benson, and Sampson also included various megalosaurids that had previously been excluded from cladograms in their 2012 study, such as Duriavenator and Wiehenvenator in Megalosaurinae and Magnosaurus, Leshansaurus, and Piveteausaurus in Afrovenatorinae.
Scuirumimus albersodoerferi, a small theropod described in 2012 which preserved protofeathers, was initially believed to be a juvenile megalosauroid. This led to the belief that megalosaurids may have had feathers. However, subsequent analyses have placed Sciurimimus as a basal coelurosaur.
Megalosaurids have been suggested to be predators or scavengers inhabiting coastal environments. Middle Jurassic-era tracks believed to have left by megalosaurids have been found at Vale de Meios in Portugal. During the middle Jurassic, this site would have been a tidal flat exposed at low tide on the edge of a lagoon. Unlike most coastal tracks, which are parallel to the coastline and probably left by migrating animals, the Vale de Meios tracks were perpendicular to the coast, with the vast majority oriented towards the lagoon. This indicates that the megalosaurids which would have left these tracks approached the tidal flat once the tide retreated.
This indicates that megalosaurids could have scavenged for the carcasses of marine creatures left by the receding tides. Another possibility is that megalosaurids were piscivorous, approaching the coast to hunt for fish. Spinosaurids, which were close relatives of megalosaurids, had numerous adaptations for piscivory and semiaquatic life, so such a lifestyle is supported by phylogenetic data. Shark teeth, cartilage fragments, and gastroliths have been documented as stomach contents in Poekilopleuron. Both this genus and Dubreillosaurus were discovered in sediments also preserving mangrove roots, providing further evidence for a coastal habitat. Nevertheless, this does not rule out the possibility that megalosaurids also fed on terrestrial prey.
Species included in Megalosauridae have been found on every modern continent, split relatively equally between sites on the Gondwana and Laurasia supercontinents. Paleogeography findings show that Megalosauridae was mainly restricted to the Middle to Late Jurassic, suggesting they went extinct at the Jurassic-Cretaceous boundary 145 million years ago.
The global radiation of these carnivorous theropods occurred in two steps. First, radiation occurred during Pangaea's breakup during the Early Jurassic, about 200 million years ago. When the Tethys Sea emerged between the supercontinent, megalosauroids radiated to the two halves of Pangaea. The second step of radiation occurred during the Middle and Late Jurassic, 174 to 145 million years ago, in allosauroids and coelurosaurs. Megalosauridae appears to have gone extinct at the end of this time period.
Megalosaurid remains have been found in various areas of the world throughout history. For example, Megalosauridae contains the most primitive theropod embryo ever found, from Early Tithonian Portugal 152 million years ago (mya). In addition, various megalosaurid fossil discoveries have been dated to Bajocian-Callovian England and France 168 to 163 mya, Middle Jurassic Africa about 170 mya, Late Jurassic China 163 to 145 mya, and Tithonian North America about 150 mya. Most recently, megalosaurids have been found in the Tiourarén Formation in Niger, proving again that these basal tetanurans have experienced global radiation.
Afrovenator (; "African hunter") is a genus of megalosaurid theropod dinosaur from the middle Jurassic Period of northern Africa.Dinosaur classification
Dinosaur classification began in 1842 when Sir Richard Owen placed Iguanodon, Megalosaurus, and Hylaeosaurus in "a distinct tribe or suborder of Saurian Reptiles, for which I would propose the name of Dinosauria." In 1887 and 1888 Harry Seeley divided dinosaurs into the two orders Saurischia and Ornithischia, based on their hip structure. These divisions have proved remarkably enduring, even through several seismic changes in the taxonomy of dinosaurs.
The largest change was prompted by entomologist Willi Hennig's work in the 1950s, which evolved into modern cladistics. For specimens known only from fossils, the rigorous analysis of characters to determine evolutionary relationships between different groups of animals (clades) proved incredibly useful. When computer-based analysis using cladistics came into its own in the 1990s, paleontologists became among the first zoologists to almost wholeheartedly adopt the system. Progressive scrutiny and work upon dinosaurian interrelationships, with the aid of new discoveries that have shed light on previously uncertain relationships between taxa, have begun to yield a stabilizing classification since the mid-2000s. While cladistics is the predominant classificatory system among paleontology professionals, the Linnean system is still in use, especially in works intended for popular distribution.Dubreuillosaurus
Dubreuillosaurus is a genus of carnivorous dinosaur from the middle Jurassic Period. It is a megalosaurid theropod. Its fossils were found in France. The only named species, Dubreuillosaurus valesdunensis, was originally described as a species of Poekilopleuron, Poekilopleuron? valesdunensis, which is still formally the type species of the genus. It was later renamed Dubreuillosaurus valesdunensis when, in 2005, Allain came to the conclusion that it was not part of the genus Poekilopleuron. Its type specimen, MNHN 1998-13, is only rivalled in the number of preserved elements in this group by that of Eustreptospondylus. Dubreuillosaurus is considered to be the sister species of Magnosaurus. It did not show signs of insular dwarfism even though it was uncovered on an island.Duriavenator
Duriavenator is a genus of theropod dinosaur described in 2008 by Roger Benson; its finds were excavated in England. The type species is D. hesperis, formerly known as Megalosaurus hesperis. Duriavenator lived during the Bajocian stage, around 170 million years ago, making it one of the oldest-known tetanurans. The genus name combines the Latin name of Dorset, Duria, with Latin for "hunter", venator. This genus has also gone under the unofficial name "Walkersaurus", which is a nomen nudum.Eustreptospondylus
Eustreptospondylus ( yoo-STREPT-o-spon-DY-ləs; meaning "true Streptospondylus") is a genus of megalosaurid theropod dinosaur, from the Oxfordian stage of the Late Jurassic period (some time between 163 and 154 million years ago) in southern England, at a time when Europe was a series of scattered islands (due to tectonic movement at the time which raised the sea-bed and flooded the lowland).Kaijiangosaurus
Kaijiangosaurus (meaning "Kiijiang lizard") is a genus of carnivorous tetanuran theropod dinosaur from the Middle Jurassic of China. In 1984 He Xinlu named and described the type species Kaijiangosaurus lini. The generic name refers to the River (jiang) Kai. The specific name honours the paleontologist Lin Wenqiu.Leshansaurus
Leshansaurus is a genus of theropod dinosaur from the Mid to Late Jurassic Dashanpu Formation of what is now China. It was described in 2009 by a team of Chinese paleontologists. The type species is Leshansaurus qianweiensis. Fossils of Leshansaurus were discovered in strata from the Shangshaximiao Formation, a formation rich in dinosaur fossils. Li et al. referred this taxon to Sinraptoridae – a group of carnosaurian theropods, but it may it belong to Megalosauridae instead.Loncosaurus
Loncosaurus (meaning uncertain; either Araucanian "chief" or Greek "lance" "lizard") was a genus of ornithopod dinosaur from the Upper Cretaceous of Provincia de Santa Cruz, Argentina. The type (and only known) species is Loncosaurus argentinus, described by the famous Argentinian paleontologist Florentino Ameghino, but is considered a dubious name. Details on this animal are often contradictory, befitting a genus that was long confused for a theropod.Magnosaurus
Magnosaurus (meaning 'large lizard') was a genus of basal tetanuran theropod dinosaur from the Middle Jurassic of England. It is based on fragmentary remains and has often been confused with or included in Megalosaurus.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 sharing a more recent common ancestor with it than with Allosaurus fragilis or Passer domesticus. Members of the group include Spinosaurus, Megalosaurus, and Torvosaurus.Megalosaurus
Megalosaurus (meaning "Great Lizard", from Greek μέγας, megas, meaning 'big', 'tall' or 'great' and σαῦρος, sauros, meaning 'lizard') is an extinct genus of large meat-eating theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years ago) of Southern England. Although fossils from other areas have been assigned to the genus, the only certain remains of Megalosaurus come from Oxfordshire and date to the late Middle Jurassic.
The earliest possible fossils of the genus came from the Taynton Limestone Formation. One of these was the lower part of a femur, discovered in the 17th century. It was originally described by Robert Plot as a thighbone of a Roman war elephant, and then as a biblical giant. The first scientific name given for it, in the 18th century, was Scrotum humanum, created by Richard Brookes as a caption; however, this is not considered valid today.
Megalosaurus was, in 1824, the first genus of non-avian dinosaur to be validly named. The type species is Megalosaurus bucklandii, named in 1827. In 1842, Megalosaurus was one of three genera on which Richard Owen based his Dinosauria. On Owen's directions a model was made as one of the Crystal Palace Dinosaurs, which greatly increased the public interest for prehistoric reptiles. Subsequently, over fifty other species would be classified under the genus, originally because dinosaurs were not well known, but even during the 20th century after many dinosaurs had been discovered. Today it is understood these additional species were not directly related to M. bucklandii, which is the only true Megalosaurus species. Because a complete skeleton of it has never been found, much is still unclear about its build.
The first naturalists who investigated Megalosaurus mistook it for a gigantic lizard of twenty metres length. In 1842, Owen concluded that it was no longer than nine metres, standing on upright legs. He still thought it was a quadruped, though. Modern scientists, by comparing Megalosaurus with its direct relatives in the Megalosauridae, were able to obtain a more accurate picture. Megalosaurus was about six metres long, weighing about seven hundred kilogrammes. It was bipedal, walking on stout hindlimbs, its horizontal torso balanced by a horizontal tail. Its forelimbs were short, though very robust. Megalosaurus had a rather large head, equipped with long curved teeth. It was generally a robust and heavily muscled animal.Orionides
Orionides is a clade of tetanuran theropod dinosaurs from the Middle Jurassic to the Present. The clade includes most theropod dinosaurs, including birds.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.Poekilopleuron
Poekilopleuron (meaning "varied ribs") is an extinct genus of megalosauroid tetanuran theropod 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 only one, Poekilopleuron, is valid. The type species is P. bucklandii, named after William Buckland, and many junior synonyms of it have also been erected. Few material is currently known, as the holotype was destroyed in World War II, although many casts of the material still exist.Streptospondylus
Streptospondylus (meaning "reversed vertebra") is a genus of tetanuran theropod dinosaur known from the Late Jurassic period of France, 161 million years ago. It was a medium-sized predator.Tetanurae
Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, maniraptorans, and birds. Tetanurans are defined as all theropods more closely related to modern birds than to Ceratosaurus and contain the majority of predatory dinosaur diversity. Tetanurae likely diverged from its sister group, Ceratosauria, during the late Triassic. Tetanurae first appeared in the fossil record by the Early Jurassic about 190 mya and by the Middle Jurassic had become globally distributed.The group was named by Jacques Gauthier in 1986 and originally had two main subgroups: Carnosauria and Coelurosauria, the clade containing birds and related dinosaurs such as compsognathids, tyrannosaurids, ornithomimosaurs, and maniraptorans. The original Carnosauria was a polyphyletic group including any large carnivorous theropod. Many of Gauthier's carnosaurs, such as tyrannosaurids, have since been re-classified as coelurosaurs or primitive tetanurans. Carnosauria has been reclassified as a group containing allosaurids that split from the Coelurosauria at the Neotetanurae/Avetheropoda node. Members of Spinosauroidea are believed to represent basal tetanurans.Tetanuran evolution was characterized by parallel diversification of multiple lineages, repeatedly attaining large body size and similar locomotor morphology. Cryolophosaurus has been claimed as the first true member of the group, but subsequent studies have disagreed on whether it is a dilophosaurid or tetanuran. Arcucci and Coria (2003) classified Zupaysaurus as an early tetanuran, but it was later placed as a sister taxon to the clade containing dilophosaurids, ceratosaurs, and tetanurans.Shared tetanuran features include a ribcage indicating a sophisticated air-sac-ventilated lung system similar to that in modern birds. This character would have been accompanied by an advanced circulatory system. Other tetanuran characterizing features include the absence of the fourth digit of the hand, placement of the maxillary teeth anterior to the orbit, a strap-like scapula, maxillary fenestrae, and stiffened tails. During the Late Jurassic and Early Cretaceous, large spinosaurids and allosaurs flourished but possibly died out in the northern hemisphere before the end of the Cretaceous, and were replaced as apex predators by tyrannosauroid coelurosaurs. At least in South America, carcharodontosaurid allosaurs persisted until the end of the Mesozoic Era, and died out at the same time the non-avian coelurosaurs.Torvosaurus
Torvosaurus () is a genus of carnivorous megalosaurid theropod dinosaurs that lived approximately 153 to 148 million years ago during the Late Jurassic Period (Kimmeridgian to Tithonian) in what is now Colorado and Portugal. It contains two currently recognized species, Torvosaurus tanneri and Torvosaurus gurneyi.
In 1979 the type species Torvosaurus tanneri was named: it was a large, heavily built, bipedal carnivore, that could grow to a length of about 10 m (33 ft). T. tanneri was among the largest carnivores of its time, together with Epanterias and Saurophaganax (which could be both synonyms of Allosaurus). Specimens referred to Torvosaurus gurneyi were initially claimed to be up to twelve metres long, but later shown to be smaller. Based on bone morphology Torvosaurus is thought to have had short but very powerful arms.Wiehenvenator
Wiehenvenator is a genus of predatory megalosaurid theropod dinosaur from the Middle Jurassic (Callovian) of Germany.Xuanhanosaurus
Xuanhanosaurus (meaning "Xuanhan lizard") is a genus of theropod dinosaur that lived during the Middle Jurassic of China, around 167.7 to 161.2 million years ago.