Masiakasaurus Temporal range: Late Cretaceous (Maastrichtian) | |
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Reconstructed skeleton, Royal Ontario Museum, Toronto, Canada | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Clade: | † Abelisauria |
Family: | † Noasauridae |
Subfamily: | † Noasaurinae |
Genus: | † Masiakasaurus Sampson et al., 2001 |
Species: | †M. knopfleri |
Binomial name | |
†Masiakasaurus knopfleri Sampson et al., 2001 | |
Masiakasaurus is a genus of small predatory noasaurid theropod dinosaurs from the Late Cretaceous of Madagascar. In Malagasy, masiaka means "vicious"; thus, the genus name means "vicious lizard". The type species, Masiakasaurus knopfleri, was named after the musician Mark Knopfler, whose music inspired the expedition crew. It was named in 2001 by Scott D. Sampson, Matthew Carrano, and Catherine A. Forster. Unlike most theropods, the front teeth of M. knopfleri projected forward instead of straight down. This unique dentition suggests that they had a specialized diet, perhaps including fish and other small prey. Other bones of the skeleton indicate that Masiakasaurus were bipedal, with much shorter forelimbs than hindlimbs. M. knopfleri was a small theropod, reaching 1.8–2.1 m (5.9–6.9 ft) long and weighing 20 kg (44 lb). [1] [2] [3]
Masiakasaurus lived from 72.1 to 66 million years ago, along with animals such as Majungasaurus , Rapetosaurus , and Rahonavis . Masiakasaurus was a member of the group Noasauridae, small predatory ceratosaurs found primarily in South America.
Remains of Masiakasaurus have been found in the Late Cretaceous Maevarano Formation in northwestern Madagascar and were first described in the journal Nature in 2001. Fragmentary bones comprising around 40% of the skeleton were collected near the village of Berivotra. Several parts of the skull, including the distinctive teeth, were found. The humerus (upper arm bone), pubis, hindlimbs, and several vertebrae were also collected. [1]
In 2011, additional specimens of Masiakasaurus were described. The braincase, premaxilla, facial bones, ribcage, portions of the hands and pectoral girdle (coracoid), and much of the cervical and dorsal vertebral column were described for the first time. The discovery of this new material clarified many aspects of noasaurid anatomy and made the genus among the best known dinosaurs. [4] The new finds did however not allow for a detailed study of its evolutionary relationships among ceratosaurs. With the new material, around 65% of the skeleton is currently known. [5]
The most distinctive characteristic of Masiakasaurus is the forward-projecting, or procumbent, front teeth. The teeth are heterodont, meaning that they have different shapes along the jaw. [1] The first four dentary teeth of the lower jaw project forward, with the first tooth angled only 10° above the horizontal. These teeth are long and spoon-shaped with hooked edges. They have carinae, or sharp edges, that are weakly serrated. Serrations are more evident along the rear edge the posterior teeth in the back of the jaw, which are also recurved and laterally compressed (flattened from the side), resembling the less unusual teeth of other carnivorous dinosaurs. The margin of the dentary curves downward so that the alveoli (tooth sockets) of the front teeth are directed forward. In fact, the alveolus of the first tooth is actually situated lower than the bottom edge of the rest of the lower jaw. [6] The lower part of the rear edge of the dentary has a long prong, known as a ventral process. This differs from the situation in abelisaurids, which have a much shorter ventral process. On the other hand, the upper part of the rear edge of the dentary is very similar to that of abelisaurids such as Majungasaurus and Carnotaurus . This part of the bone possesses an array of four small structures, three of which line a socket which connects to the surangular bone at the back of the lower jaw. Although the surangular bone is not preserved, several other bones of the lower jaw are, including a triangular angular bone, a gently curving prearticular bone, and a damaged yet notably concave articular bone. The angular and prearticular formed the lower edge of a large and rounded in the lower jaw (known as a mandibular fenestra) while the articular bone formed the lower part of the jaw joint. A long and tapering hyoid (tongue bone) has also been preserved. [5] The front teeth of the upper jaw are also procumbent, and the margin of the premaxilla curves slightly upward to direct them outward. Unlike the skulls of abelisaurids, which are very deep, the skull of Masiakasaurus is long and low. The lacrimal and postorbital bones around the eye are textured with bumpy projections. Not including the highly modified jaws and teeth, the skull of Masiakasaurus possesses many general ceratosaurian characteristics. Overall, its morphology is intermediate between abelisaurids and more basal ceratosaurs. [5]
The neck is relatively narrow in comparison to abelisaurids and bear stout neck ribs. While many theropods have s-shaped necks, the ribs would make the neck rather stiff in Masiakasaurus, and the back of the neck is positioned almost horizontally, giving it only a slighter curve. Like those of other abelisauroids, the vertebrae are heavily pneumaticized, or hollowed, and have relatively short neural spines. Pneumaticity is limited to the neck and foremost back vertebrae, however. Pneumatic cavities are also present in the braincase. [5]
As in other ceratosaurs, the shoulder blade (scapula) and shoulder girdle fuse into a single bone, the scapulocoracoid. This bone is very large and broad, even compared to the condition in other ceratosaurs. The scapula portion (above the glenoid, or arm socket) tapers towards the back while the coracoid portion (below the glenoid) is expanded into a curved blade-like structure. While abelisaurids have arms that are extremely reduced in size, Masiakasaurus and other noasaurids had longer forelimbs. The humerus is slender and known bones of the hand are relatively short. The related genus Noasaurus has a large and curved raptorial ungual (claw) which was originally interpreted as a sickle-like foot claw as in dromaeosaurids such as Velociraptor . More recently, this has been re-evaluated as a claw of the hand. The penultimate phalanx, the finger bone that immediately precedes the raptorial ungual in Noasaurus, is also known in Masiakasaurus and has a similar appearance. The enlarged ungual, however, is unknown in Masiakasaurus. [5] It is assumed that members of this genus had four fingers, with the middle two fingers being the longest as in other ceratosaurians.
In its initial 2001 description, Masiakasaurus was classified as a basal abelisauroid related to Laevisuchus and Noasaurus, two poorly known genera named in 1933 and 1980, respectively. [1] In the following year, Carrano et al. (2002) placed Masiakasaurus along with Laevisuchus and Noasaurus in the family Noasauridae. They conducted a phylogenetic analysis of abelisauroids using characteristics from Masiakasaurus. Below is a cladogram from an updated version of their analysis showing the phylogenetic placement of Masiakasaurus. [7]
Abelisauroidea |
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Carrano et al. (2002) distinguished two forms of Masiakasaurus, a robust form and a gracile form. The robust morph includes specimens with thicker bones and more pronounced projections for the attachment of ligaments and muscles. The gracile form includes specimens that are more slender and have less pronounced muscle attachments. It also has unfused tibiae, unlike the fused tibiae of the robust form. These two varieties may be an indication of sexual dimorphism in Masiakasaurus, but they may also represent two distinct populations. [6]
One specimen of Masiakasaurus, a right scapulocoracoid, bears holes that may be puncture marks from predation or scavenging. Majungasaurus , a large abelisaurid from the Maevarano Formation, may have preyed upon Masiakasaurus. [8] The holes may also have been the result of an infection. [5]
The procument front teeth of Masiakasaurus were likely an adaptation for grasping small prey. They would have been unsuitable for tearing larger food apart. In the front of the jaws, carinae are restricted to the base of the teeth and would not have been used to tear prey. The back teeth, however, share the same general characteristics as those of most other theropods, suggesting that they served a similar function in Masiakasaurus such as cutting and slicing. [6]
Several feeding behaviors have been proposed for Masiakasaurus on the basis of its unusual dentition. Because the front teeth would have been well suited for grasping, Masiakasaurus may have consumed small vertebrates, invertebrates, and possibly even fruits. [6]
In 2013, Lee and O'Connor observed that Masiakasaurus would be a good subject for an analysis of theropod growth, considering that there is an abundance of fossil material to examine from a broad range of ontogenetic stages. The study showed that Masiakasaurus grew determinately, and reached full maturity at a small body size. Competing theories that Masiakasaurus specimens represent the juvenile form of a larger-bodied theropod were not supported by the data. Masiakasaurus took 8 to 10 years to grow the size of a large dog. This indicates a rate of growth that is 40% slower than that of comparably sized non-avian theropods, a finding that is supported by the unusual prominence of parallel-fibered bone which is known to be associated with relatively slow growth. However, individuals in this genus grew 40% faster than crocodylians. Lee and O'Connor noted that the evolution of slow growth gave this dinosaur the advantage of minimizing the nutritional investment allocated toward structural growth while living in a semiarid and seasonally stressful environment. [9]
Carnotaurus is a genus of theropod dinosaur that lived in South America during the Late Cretaceous period, probably sometime between 72 and 69 million years ago. The only species is Carnotaurus sastrei. Known from a single well-preserved skeleton, it is one of the best-understood theropods from the Southern Hemisphere. The skeleton, found in 1984, was uncovered in the Chubut Province of Argentina from rocks of the La Colonia Formation. Carnotaurus is a derived member of the Abelisauridae, a group of large theropods that occupied the large predatorial niche in the southern landmasses of Gondwana during the late Cretaceous. Within the Abelisauridae, the genus is often considered a member of the Brachyrostra, a clade of short-snouted forms restricted to South America.
Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with Ceratosaurus than with birds. The oldest known ceratosaur, Saltriovenator, dates to the earliest part of the Jurassic, around 199 million years ago. Ceratosauria includes three major clades: Ceratosauridae, Noasauridae, and Abelisauridae, found primarily in the Southern Hemisphere. Originally, Ceratosauria included the above dinosaurs plus the Late Triassic to Early Jurassic Coelophysoidea and Dilophosauridae, implying a much earlier divergence of ceratosaurs from other theropods. However, most recent studies have shown that coelophysoids and dilophosaurids do not form a natural group with other ceratosaurs, and are excluded from this group.
Rajasaurus is a genus of carnivorous abelisaurid theropod dinosaur from the Late Cretaceous of India, containing one species: Rajasaurus narmadensis. The bones were excavated from the Lameta Formation in the Gujarat state of Western India, probably inhabiting what is now the Narmada River Valley. It was formally described by palaeontologist Jeffrey A. Wilson and colleagues in 2003 based on a partial skeleton comprising the braincase, spine, hip bone, legs, and tail–a first for an Indian theropod. The dinosaur likely measured 6.6 metres (22 ft), and had a single horn on the forehead which was probably used for display and head-butting. Like other abelisaurids, Rajasaurus was probably an ambush predator.
Afrovenator is a genus of megalosaurid theropod dinosaur from the Middle or Late Jurassic Period on the Tiourarén Formation and maybe the Irhazer II Formation of the Niger Sahara region in northern Africa. Afrovenator represents the only properly identified Gondwanan megalosaur, with proposed material of the group present in the Late Jurassic on Tacuarembó Formation of Uruguay and the Tendaguru Formation of Tanzania.
Abelisauridae is a family of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found on the modern continents of Africa and South America, as well as on the Indian subcontinent and the island of Madagascar. Isolated teeth were found in the Late Jurassic of Portugal, and the Late Cretaceous genera Tarascosaurus, Arcovenator and Caletodraco have been described in France. Abelisaurids possibly first appeared during the Jurassic period based on fossil records, and some genera survived until the end of the Mesozoic era, around 66 million years ago.
Deltadromeus is a genus of theropod dinosaur from the Aoufous Formation of Morocco.
Ligabueino is a genus of noasaurid dinosaur named after its discoverer, Italian doctor Giancarlo Ligabue. It is known only from an extremely fragmentary specimen, measuring 79 cm (2.6 ft) long, found in the La Amarga Formation. In spite of initial reports that it was an adult, the unfused vertebrae indicate that the specimen was a juvenile. It was a theropod and lived during the Early Cretaceous Period, in what is now Patagonia. Contrary to initial classifications that placed it as a member of the Noasauridae, Carrano and colleagues found in 2011 that it could only be placed with any confidence in the group Abelisauroidea.
Elaphrosaurus is a genus of ceratosaurian theropod dinosaur that lived approximately 154 to 150 million years ago during the Late Jurassic Period in what is now Tanzania in Africa. Elaphrosaurus was a medium-sized but lightly built member of the group that could grow up to 6.2 m (20 ft) long. Morphologically, this dinosaur is significant in two ways. Firstly, it has a relatively long body but is very shallow-chested for a theropod of its size. Secondly, it has very short hindlimbs in comparison with its body. Phylogenetic analyses indicate that this genus is likely a ceratosaur. Earlier suggestions that it is a late surviving coelophysoid have been examined but generally dismissed. Elaphrosaurus is currently believed to be a very close relative of Limusaurus, an unusual beaked ceratosaurian which may have been either herbivorous or omnivorous.
Compsosuchus is a dubious genus of abelisauroid dinosaur from the Late Cretaceous Lameta Formation of India.
Majungasaurus is a genus of abelisaurid theropod dinosaur that lived in Madagascar from 70 to 66 million years ago, at the end of the Cretaceous Period, making it one of the last-known non-avian dinosaurs that went extinct during the Cretaceous–Paleogene extinction event. The genus contains a single species, Majungasaurus crenatissimus. This dinosaur is also called Majungatholus, a name which is considered a junior synonym of Majungasaurus.
Genusaurus is a genus of abelisaurid theropod from the Early Cretaceous. Its fossils were found in France. Genusaurus is believed to have lived during the Albian stage, around 112-100 million years ago.
Noasaurus is a genus of ceratosaurian theropod dinosaur genus from the late Campanian-Maastrichtian of Argentina. The type and only species is N. leali.
Laevisuchus is a genus of theropod dinosaur from the Late Cretaceous. Its remains were discovered by Charles Alfred Matley near Jabalpur in Maastrichtian "Carnosaur Bed" deposits in the Lameta Formation in Madhya Pradesh, central India, and were named and described by paleontologists Friedrich von Huene and Matley in 1933.
Noasauridae is an extinct family of theropod dinosaurs belonging to the group Ceratosauria. They were closely related to the short-armed abelisaurids, although most noasaurids had much more traditional body types generally similar to other theropods. Their heads, on the other hand, had unusual adaptations depending on the subfamily. 'Traditional' noasaurids, sometimes grouped in the subfamily Noasaurinae, had sharp teeth which splayed outwards from a downturned lower jaw.
Limusaurus is a genus of theropod dinosaur that lived in what is now China during the Late Jurassic, around 161 to 157 million years ago. The type and only species Limusaurus inextricabilis was described in 2009 from specimens found in the Upper Shishugou Formation in the Junggar Basin of China. The genus name consists of the Latin words for "mud" and "lizard", and the species name means "impossible to extricate", both referring to these specimens possibly dying after being mired. Limusaurus was a small, slender animal, about 1.7 m in length and 15 kg (33 lb) in weight, which had a long neck and legs but very small forelimbs. It underwent a drastic morphological transformation as it aged: while juveniles were toothed, these teeth were completely lost and replaced by a beak with age. Several of these features were convergently similar to the later ornithomimid theropods as well as the earlier non-dinosaurian shuvosaurids.
Dahalokely is an extinct genus of carnivorous abelisauroid theropod dinosaur from the Late Cretaceous (Turonian) of Madagascar.
Arcovenator is an extinct genus of abelisaurid theropod dinosaurs hailing from the Late Cretaceous of France and possibly Spain. The type and only described species is Arcovenator escotae.
Majungasaurinae is a subfamily of large carnivorous theropods from the Upper Cretaceous, found in Madagascar, India, and France. It is a subgroup within the theropod family Abelisauridae, a Gondwanan clade known for their thick and often horned skulls and vestigial arms. The two subfamilies of Abelisauridae are Carnotaurinae, best known from the South American Carnotaurus, and Majungasaurinae, consisting of Madagascar’s Majungasaurus and its closest relatives. Their ancestors emerged in the Middle Jurassic, and the clade lasted until the Upper Cretaceous.
Afromimus is a genus of theropod dinosaur from the Early Cretaceous Elrhaz Formation of Niger. It contains a single species, A. tenerensis, named in 2017 by Paul Sereno from parts of the right leg, vertebrae, and ribs found in the Ténéré Desert. It was originally classified as an ornithomimosaurian, but subsequently it was argued to be an abelisauroid.
Tralkasaurus is a genus of abelisaurid dinosaur from the Huincul Formation from Río Negro Province in Argentina. The type and only species is Tralkasaurus cuyi, named in 2020 by Mauricio Cerroni and colleagues based on an incomplete skeleton. A medium-sized abelisaurid, Tralkasaurus exhibits a conflicting blend of characteristics found among the early-diverging abelisauroids with others that characterize the highly specialized clade Brachyrostra, and thus its position within the clade is poorly-resolved.