Stegosauria

Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe and China, though one species (Kentrosaurus aethiopicus) is known to have lived in Africa. Their geographical origins are unclear; the earliest unequivocal stegosaurian, Huayangosaurus taibaii, lived in China. Stegosaurians were armored dinosaurs (thyreophorans). Originally, they did not differ much from more primitive members of that group, being small, low-slung, running animals protected by armored scutes. An early evolutionary innovation was the development of tail spikes, or 'thagomizers', as defensive weapons. Later species, belonging to a subgroup called the Stegosauridae, became larger, and developed long hindlimbs that no longer allowed them to run. This increased the importance of active defence by the thagomizer, which could ward off even large predators because the tail was in a higher position, pointing horizontally to the rear from the broad pelvis. Stegosaurids had complex arrays of spikes and plates running along their backs, hips and tails. Their necks became longer and their small heads became narrow, able to selectively bite off the best parts of cycads with their beaks. When these plant types declined in diversity, so did the stegosaurians, which became extinct during the first half of the Cretaceous period. The first stegosaurian finds in the early 19th century were fragmentary. Better fossil material, of the genus Dacentrurus, was discovered in 1874 in England. Soon after, in 1877, the first nearly-complete skeleton was discovered in the United States. Professor Othniel Charles Marsh that year classified such specimens in the new genus Stegosaurus, from which the group acquired its name, and which is still by far the most famous stegosaurian. During the latter half of the twentieth century, many important Chinese finds were made, representing about half of the presently known diversity of stegosaurians. Stegosaurians had characteristic small, long, flat, narrow heads and a horn-covered beak or rhamphotheca, which covered the front of the snout (two premaxillaries) and lower jaw (a single predentary) bones. Similar structures are seen in turtles and birds. Apart from Huayangosaurus, stegosaurians subsequently lost all premaxillary teeth within the upper beak. Huayangosaurus still had seven per side. The upper and lower jaws are equipped with rows of small teeth. Later species have a vertical bone plate covering the outer side of the lower jaw teeth. The structure of the upper jaw, with a low ridge above, and running parallel to, the tooth row, indicates the presence of a fleshy cheek. In stegosaurians, the typical archosaurian skull opening, the antorbital fenestra in front of the eye socket, is small, sometimes reduced to a narrow horizontal slit. All stegosaurians are quadrupedal, with hoof-like toes on all four limbs. All stegosaurians after Huayangosaurus have forelimbs much shorter than their hindlimbs. Their hindlimbs are long and straight, designed to carry the weight of the animal while stepping. The condyles of the lower thighbone are short from the front to the rear. This would have limited the supported rotation of the knee joint, making running impossible. Huayangosaurus had a thighbone like a running animal. The upper leg was always longer than the lower leg. Huayangosaurus had relatively long and slender arms. The forelimbs of later forms are very robust, with a massive humerus and ulna. The wrist bones were reinforced by a fusion into two blocks, an ulnar and a radial. The front feet of stegosaurians are commonly depicted in art and in museum displays with fingers splayed out and slanted downward. However, in this position, most bones in the hand would be disarticulated. In reality, the hand bones of stegosaurians were arranged into vertical columns, with the main fingers, orientated outwards, forming a tube-like structure. This is similar to the hands of sauropod dinosaurs, and is also supported by evidence from stegosaurian footprints and fossils found in a lifelike pose. The long hindlimbs elevated the tail base, such that the tail pointed out behind the animal almost horizontally from that high position. While walking, the tail would not have sloped downwards as this would have impeded the function of the tail base retractor muscles, to pull the thighbones backwards. However, it has been suggested by Robert Thomas Bakker that stegosaurians could rear on their hind legs to reach higher layers of plants, the tail then being used as a 'third leg'. The mobility of the tail was increased by a reduction or absence of ossified tendons, that with many Ornithischia stiffen the hip region. Huayangosaurus still possessed them. In species that had short forelimbs, the relatively short torso towards the front curved strongly downwards. The dorsal vertebrae typically were very high, with very tall neural arches and transverse processes pointing obliquely upwards to almost the level of the neural spine top. Stegosaurian back vertebrae can easily be identified by this unique configuration. The tall neural arches often house deep neural canals; enlarged canals in the sacral vertebrae have given rise to the incorrect notion of a 'second brain'. Despite the downwards curvature of the rump, the neck base was not very low and the head was held a considerable distance off the ground. The neck was flexible and moderately long. Huayangosaurus still had the probably original number of nine cervical vertebrae; Miragaia has an elongated neck with seventeen. The stegosaurian shoulder girdle was very robust. In Huayangosaurus, the acromion, a process on the lower front edge of the shoulderblade, was moderately developed; the coracoid was about as wide as the lower end of the scapula, with which it formed the shoulder joint. Later forms tend to have a strongly expanded acromion, while the coracoid, largely attached to the acromion, no longer extends to the rear lower corner of the scapula. Ossified sternal plates have never been found with Stegosauria and perhaps the sternum was completely absent.