Old Earth Ministries Online Dinosaur Curriculum

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Lesson 12 - Majungasaurus

Majungasaurus ("Mahajanga lizard") is a genus of abelisaurid theropod dinosaur that lived in Madagascar from 70 to 65 million years ago, at the end of the Cretaceous Period. Only one species (M. crenatissimus) has been identified. This dinosaur was briefly called Majungatholus, a name which is now considered a junior synonym of Majungasaurus.

Like other abelisaurids, Majungasaurus was a bipedal predator with a short snout. Although the forelimbs are not completely known, they were very short, while the hindlimbs were longer and very stocky.

It can be distinguished from other abelisaurids by its wider skull, the very rough texture and thickened bone on the top of its snout, and the single rounded horn on the roof of its skull, which was originally mistaken for the dome of a pachycephalosaur. It also had more teeth in both upper and lower jaws than most abelisaurids.

Known from several well-preserved skulls and abundant skeletal material, Majungasaurus has recently become one of the best-studied theropod dinosaurs from the Southern Hemisphere. It appears to be most closely related to abelisaurids from India rather than South America or continental Africa, a fact which has important biogeographical implications. Majungasaurus was the apex predator in its ecosystem, mainly preying on sauropods like Rapetosaurus, and is also the only dinosaur for which direct evidence of cannibalism is known.

Description

Majungasaurus was a medium-sized theropod that typically measured 6–7 meters (20–23 ft) in length, including its tail. Fragmentary remains of larger individuals indicate that some adults reached lengths of more than 8 meters (26 ft). Scientists estimate that an average adult Majungasaurus weighed more than 1100 kilograms (2400 lb), although the largest animals would have weighed more. Its 8–9 meter (26–30 ft) relative Carnotaurus has been estimated to weigh 1500 kilograms (3300 lb).

The skull of Majungasaurus is exceptionally well-known compared to most theropods and generally similar to that of other abelisaurids. Like other abelisaurid skulls, its length was proportionally short for its height, although not as short as in Carnotaurus. The skulls of large individuals measured 60–70 centimeters (24–28 in) long.  However, the skull of Majungasaurus was markedly wider than in other abelisaurids. All abelisaurids had a rough, sculptured texture on the outside faces of the skull bones, and Majungasaurus was no exception. This was carried to an extreme on the nasal bones of Majungasaurus, which were extremely thick and fused together, with a low central ridge running along the half of the bone closest to the nostrils. A distinctive dome-like horn protruded from the fused frontal bones on top of the skull as well. In life, these structures would have been covered with some sort of integument, possibly made of keratin.

The postcranial skeleton of Majungasaurus closely resembles those of Carnotaurus and Aucasaurus, the only other abelisaurid genera for which complete skeletal material is known. Majungasaurus was bipedal, with a long tail to balance out the head and torso, putting the center of gravity over the hips.  The humerus (upper arm bone) was short and curved, closely resembling those of Aucasaurus and Carnotaurus. Also like related dinosaurs, Majungasaurus had very short forelimbs with four extremely reduced digits, with only two very short external fingers and no claws. The hand and finger bones of Majungasaurus, like other carnotaurines, lacked the characteristic pits and grooves where claws and tendons would normally attach, and its finger bones were fused together, indicating that the hand was immobile.

Discovery and Naming

French paleontologist Charles Depéret described the first theropod remains from northwestern Madagascar in 1896. These included two teeth, a claw, and some vertebrae discovered along the Betsiboka River by a French army officer and deposited in the collection of what is now the Université Claude Bernard Lyon 1. Depéret referred these fossils to the genus Megalosaurus, which at the time was a wastebasket taxon containing any number of unrelated large theropods, as the new species M. crenatissimus.

Numerous fragmentary remains from Mahajanga Province in northwestern Madagascar were recovered by French collectors over the next 100 years, many of which were deposited in the Muséum National d'Histoire Naturelle in Paris. In 1955, René Lavocat described a theropod dentary with teeth from the Maevarano Formation in the same region where the original material was found. The teeth matched those first described by Depéret, but the strongly curved jaw bone was very different from both Megalosaurus and Dryptosaurus. Lavocat renamed the genus Majungasaurus, using an older spelling of Mahajanga as well as the Greek word meaning "lizard", and made this jaw bone the type specimen. In the 1990s, more specimins were found which helped describe Majungasaurus to a greater extent.

Paleobiology

Skull ornamentation

Majungasaurus is perhaps most distinctive for its skull ornamentation, including the swollen and fused nasals and the frontal horn. Other ceratosaurs, including Carnotaurus, Rajasaurus, and Ceratosaurus itself bore crests on the head. These structures are likely to have played a role in intraspecific competition, although their exact function within that context is unknown. The hollow cavity inside the frontal horn of Majungasaurus would have weakened the structure and probably precluded its use in direct physical combat, although the horn may have served a display purpose. While there is variation in the ornamentation of Majungasaurus individuals, there is no evidence for sexual dimorphism.

Feeding

Scientists have suggested that the unique skull shape of Majungasaurus and other abelisaurids indicate different predatory habits than other theropods. Whereas most theropods were characterized by long, low skulls of narrow width, abelisaurid skulls were taller and wider, and often shorter in length as well. The narrow skulls of other theropods were well-equipped to withstand the vertical stress of a powerful bite, but not as good at withstanding torsion (twisting). In comparison to modern mammalian predators, most theropods may have used a strategy similar in some ways to that of long- and narrow-snouted canids, with the delivery of many bites weakening the prey animal.

Abelisaurids, especially Majungasaurus, may instead have been adapted for a feeding strategy more similar to modern felids, with short and broad snouts, that bite once and hold on until the prey is subdued. Majungasaurus had an even broader snout than other abelisaurids, and other aspects of its anatomy may also support the bite-and-hold hypothesis. The neck was strengthened, with robust vertebrae, interlocking ribs and ossified tendons, as well as reinforced muscle attachment sites on the vertebrae and the back of the skull. These muscles would have been able to hold the head steady despite the struggles of its prey. Abelisaurid skulls were also strengthened in many areas by bone mineralized out of the skin, creating the characteristic rough texture of the bones. This is particularly true of Majungasaurus, where the nasal bones were fused and thickened for strength. On the other hand, the lower jaw of Majungasaurus sported a large fenestra (opening) on each side, as seen in other ceratosaurs, as well as synovial joints between certain bones that allowed a high degree of flexibility in the lower jaw, although not to the extent seen in snakes. This may have been an adaptation to prevent the fracture of the lower jaw when holding onto a struggling prey animal. The front teeth of the upper jaw were more robust than the rest, to provide an anchor point for the bite, while the low crown height of Majungasaurus teeth prevented them from breaking off during a struggle. Finally, unlike the teeth of Allosaurus and most other theropods, which were curved on both the front and back, abelisaurids like Majungasaurus had teeth curved on the front edge but straighter on the back (cutting) edge. This structure may have served to prevent slicing, and instead holding the teeth in place when biting.

Majungasaurus was the largest predator in its environment, while the only known large herbivores at the time were sauropods like Rapetosaurus. Scientists have suggested that Majungasaurus, and perhaps other abelisaurids, specialized on hunting sauropods.  Majungasaurus tooth marks on Rapetosaurus bones confirm that it at least fed on these sauropods, whether or not it actually killed them.

Cannibalism

Although sauropods may have been the prey of choice for Majungasaurus, recent discoveries in Madagascar indicate another surprising component of its diet: other Majungasaurus. Numerous bones of Majungasaurus have been discovered bearing tooth marks identical to those found on sauropod bones from the same localities. These marks have the same spacing as teeth in Majungasaurus jaws, are of the same size as Majungasaurus teeth, and contain smaller notches consistent with the serrations on those teeth. As Majungasaurus is the only large theropod known from the area, the simplest explanation is that it was feeding on other members of its own species. Suggestions that the Triassic Coelophysis was a cannibal have been recently disproven, leaving Majungasaurus as the only non-avian theropod with confirmed cannibalistic tendencies, although there is some evidence that cannibalism may have occurred in other species as well.

It is unknown if Majungasaurus actively hunted their own kind or only scavenged their carcasses.

Respiratory System

Scientists have reconstructed the respiratory system of Majungasaurus based on a superbly preserved series of vertebrae (UA 8678) recovered from the Maevarano Formation. Most of these vertebrae and some of the ribs contained cavities (pneumatic foramina) that may have resulted from the infiltration of avian-style lungs and air sacs. In birds, the neck vertebrae and ribs are hollowed out by the cervical air sac, the upper back vertebrae by the lung, and the lower back and sacral (hip) vertebrae by the abdominal air sac. Similar features in Majungasaurus vertebrae imply the presence of these air sacs. These air sacs may have allowed for a basic form of avian-style 'flow-through ventilation,' where air flow through the lungs is one-way, so that oxygen-rich air inhaled from outside the body is never mixed with exhaled air laden with carbon dioxide. This method of respiration, while complicated, is highly efficient.

The recognition of pneumatic foramina in Majungasaurus, besides providing an understanding of its respiratory biology, also has larger-scale implications for evolutionary biology. The split between the ceratosaur line, which led to Majungasaurus, and the tetanuran line, to which birds belong, occurred very early in the history of theropods. The avian respiratory system, present in both lines, must therefore have evolved before the split, and well before the evolution of birds themselves. This provides further evidence of the dinosaurian origin of birds.

Brain and inner ear structure

Computed tomography, also known as CT scanning, of a complete Majungasaurus skull (FMNH PR 2100) allowed a rough reconstruction of its brain and inner ear structure. Overall, the brain was very small relative to body size, but otherwise similar to many other non-coelurosaurian theropods, with a very conservative form closer to modern crocodilians than to birds. One difference between Majungasaurus and other theropods was its smaller flocculus, a region of the cerebellum that helps to coordinate movements of the eye with movements of the head. This suggests that Majungasaurus and other abelisaurids like Indosaurus, which also had a small flocculus, did not rely on quick head movements to sight and capture prey.

Inferences about behavior can also be drawn from examination of the inner ear. The semicircular canals within the inner ear aid in balance, and the lateral semicircular canal is usually parallel to the ground when the animal holds its head in an alert posture. When the skull of Majungasaurus is rotated so that its lateral canal is parallel to the ground, the entire skull is nearly horizontal. This contrasts with many other theropods, where the head was more strongly downturned when in the alert position. The lateral canal is also significantly longer in Majungasaurus than in its more basal relative Ceratosaurus, indicating a greater sensitivity to side-to-side motions of the head.

Pathology

A 2007 report described pathologies in the bones of Majungasaurus. Scientists examined the remains of at least 21 individuals and discovered four with noticeable pathologies. While pathology had been studied in large tetanuran theropods like allosaurids and tyrannosaurids, this was the first time an abelisauroid had been examined in this manner. No wounds were found on any skull elements, in contrast to tyrannosaurids where sometimes gruesome facial bites were common. One of the specimens was a phalanx (toe bone) of the foot, which had apparently been broken and subsequently healed.

Most of the pathologies occurred on the vertebrae. For example, a dorsal (back) vertebra from a juvenile animal showed an exostosis (bony growth) on its underside. The growth probably resulted from the conversion of cartilage or a ligament to bone during development, but the cause of the ossification was not determined. Hypervitaminosis A and bone spurs were ruled out, and an osteoma (benign bone tumor) was deemed unlikely. Another specimen, a small caudal (tail) vertebra, was also found to have an abnormal growth, this time on the top of its neural spine, which projects upwards from the vertebrae, allowing muscle attachment. Similar growths from the neural spine have been found in specimens of Allosaurus and Masiakasaurus, probably resulting from the ossification of a ligament running either between the neural spines (interspinal ligament) or along their tops (supraspinal ligament).

The most serious pathology discovered was in a series of five large tail vertebrae. The first two vertebrae showed only minor abnormalities with the exception of a large groove that extended along the left side of both bones. However, the next three vertebrae were completely fused together at many different points, forming a solid bony mass. There is no sign of any other vertebrae after the fifth in the series, indicating that the tail ended there prematurely. From the size of the last vertebrae, scientists judged that about ten vertebrae were lost. One explanation for this pathology is severe physical trauma resulting in the loss of the tail tip, followed by osteomyelitis (infection) of the last remaining vertebrae. Alternatively, the infection may have come first and led to the end of the tail becoming necrotic and falling off. This is the first example of tail truncation known in a non-avian theropod dinosaur.

Paleoecology

All specimens of Majungasaurus have been recovered from the Maevarano Formation in the Mahajanga Province in northwestern Madagascar. Most of these, including all of the most complete material, came from the Anembalemba Member, although Majungasaurus teeth have also been found in the underlying Masorobe Member and the overlying Miadana Member. While these sediments have not been dated radiometrically, evidence from biostratigraphy and paleomagnetism suggest that they were deposited during the Maastrichtian stage, which lasted from 70 to 65 Ma (million years ago). Majungasaurus teeth are found up until the very end of the Maastrichtian, when all non-avian dinosaurs went extinct.

Then as now, Madagascar was an island, having separated from the Indian subcontinent less than 20 million years earlier. It was drifting northwards but still 10–15° more southerly in latitude than it is today. The prevailing climate of the time was semi-arid, with pronounced seasonality in temperature and rainfall. Majungasaurus inhabited a coastal flood plain cut by many sandy river channels. Strong geological evidence suggests the occurrence of periodic debris flows through these channels at the beginning of the wet season, burying the carcasses of organisms killed during the preceding dry season and providing for their exceptional preservation as fossils. Sea levels in the area were rising throughout the Maastrichtian, and would continue to do so into the Paleocene Epoch, so Majungasaurus may have roamed coastal environments like tidal flats as well. The neighboring Berivotra Formation represents the contemporaneous marine environment.

Besides Majungasaurus, fossil taxa recovered from the Maevarano include fish, frogs, lizards, snakes, seven distinct species of crocodylomorphs, five or six species of mammals, Vorona and several other birds, the possibly flighted dromaeosaurid Rahonavis, the noasaurid Masiakasaurus and two titanosaurian sauropods, including Rapetosaurus. Majungasaurus was by far the largest carnivore and probably the dominant predator on land, although large crocodylomorphs like Mahajangasuchus and Trematochampsa might have competed with it closer to water.

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