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15.01.2016 00:08 - Encyclopedia Largest prehistoric animals Vol.1 Vertebrates part3 Dinosaurs ch.3 Ornithischia - Armoured dinosaurs
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Ornithischia is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure similar to that of birds.
Ornithischians with well known anatomical adaptations include the ceratopsians or "horn-faced" dinosaurs (e.g. Triceratops), armored dinosaurs (Thyreophora) such as stegosaurs and ankylosaurs, pachycephalosaurids and the ornithopods. Ornithischians shifted from bipedal to quadrupedal posture at least three times in their evolutionary history and it has been shown primitive members may have been capable of both forms of movement.

Most ornithischians were herbivorous.In fact, most of the unifying characters of Ornithischia are thought to be related to this herbivory.For example, the shift to an opisthopubic pelvis is thought to be related to the development of a large stomach or stomachs and gut which would allow ornithischians to digest plant matter better.The smallest known ornithischian is Fruitadens haagarorum.The largest Fruitadens individuals reached just 65–75 cm. Previously, only carnivorous, saurischian theropods were known to reach such small sizes.At the other end of the spectrum, the largest known ornithischians reach about 15 meters (smaller than the largest saurischians).
However, not all ornithischians were strictly herbivorous. Some groups, like the heterodontosaurids, were likely omnivores.At least one species of ankylosaurian, Liaoningosaurus paradoxus, appears to have been at least partially carnivorous, with hooked claws, fork-like teeth, and stomach contents suggesting that it may have fed on fish.
There is strong evidence that some ornithischians lived in herds.This evidence consists of multiple bone beds where large numbers of individuals of the same species and of different age groups died simultaneously.
The early ornithopods were only about 1 metre (3 feet) long, but probably very fast. They had a stiff tail, like the theropods, to help them balance as they ran on their hind legs. Later ornithopods became more adapted to grazing on all fours; their spines curved, and came to resemble the spines of modern ground-feeders such as the bison. As they became more adapted to eating while bent over, they became facultative quadrupeds; still running on two legs, and comfortable reaching up into trees; but spending most of their time walking or grazing while on all fours. The taxonomy of dinosaurs previously ascribed to the Hypsilophodontidae is problematic. The group previously consisted of all non-iguanodontian bipedal ornithischians, but a phylogenetic reappraisal has shown such species to be paraphyletic. As such, the hypsilophodont family is currently represented only by Hypsilophodon.
Later ornithopods became larger, but never rivalled the incredible size of the long-necked, long-tailed sauropods that they partially supplanted. The very largest, such as Shantungosaurus, were as heavy as medium-sized sauropods at up to 23 metric tons (25 short tons) but never grew much beyond 15 metres (50 feet).

Armoured dinosaurs (


Thyreophora is a group of armored ornithischian dinosaurs that lived from the early Jurassic Period until the end of the Cretaceous.
Thyreophorans are characterized by the presence of body armor lined up in longitudinal rows along the body. Primitive forms had simple, low, keeled scutes or osteoderms, whereas more derived forms developed more elaborate structures including spikes and plates. Most thyreophorans were herbivorous and had relatively small brains for their body size.
Thyreophora includes various subgroups, including the suborders Ankylosauria and Stegosauria. In both the suborders, the forelimbs were much shorter than the hindlimbs, particularly in stegosaurs. The clade has been defined as the group consisting of all species more closely related to Ankylosaurus than to Triceratops. Thyreophora is the sister group of Cerapoda within Genasauria.

The largest
thyreophoran was Ankylosaurus at 9 metres (30 ft) in length and 6 tonnes in weight.
Ankylosaurus was a large animal by modern standards. The skull of the largest known individual has been estimated at 64.5 cm (25.4 inches) long and 74.5 cm (29.3 inches) wide, the whole animal was estimated at 6.25 m (20.5 feet) long, 1.5 m (4.9 feet) wide and 1.7 m (5.6 feet) tall at the hip.
The body shape was low-slung and quite wide. It was quadrupedal, with the hind limbs longer than the forelimbs. Although its feet are still unknown, comparisons with other ankylosaurs suggest Ankylosaurus probably had five toes on each foot. The skull was low and triangular in shape, wider than it was long. Like other ankylosaurs, Ankylosaurus had small, leaf-shaped teeth, which were the smallest relative to the body size, than in any other ankylosaurid species,and lacked the tooth batteries of the contemporaneous ceratopsid and hadrosaurid dinosaurs. Bones in the skull and other parts of the body were fused, increasing their strength.
The most obvious feature of Ankylosaurus is its armor, consisting of massive knobs and plates of bone, known as osteoderms or scutes, embedded in the skin. Osteoderms are also found in the skin of crocodiles, armadillos and some lizards. The bone was probably overlain by a tough, horny layer of keratin. These osteoderms ranged greatly in size, from wide, flat plates to small, round nodules. The plates were aligned in regular horizontal rows down the animal"s neck, back, and hips, with the many smaller nodules protecting the areas between the large plates. Smaller plates may have been arranged on the limbs and tail. Compared to the slightly more ancient ankylosaurid Euoplocephalus, the plates of Ankylosaurus were smooth in texture, without the high keels found on the armor of the contemporaneous nodosaurid Edmontonia. A row of flat, triangular spikes may have protruded laterally along each side of the tail. Tough, rounded scales protected the top of the skull, while four large pyramidal horns projected outwards from its rear corners.
The tail club of Ankylosaurus was also composed of several large osteoderms, which were fused to the last few tail vertebrae. It was heavy and supported by the last seven tail vertebrae, which interlocked to form a stiff rod at the base of the club. Thick tendons have been preserved, which attached to these vertebrae.
Ankylosaurus was named as the type genus of the family Ankylosauridae.Ankylosaurids are members of the larger taxon Ankylosauria, which also contains the nodosaurids. Ankylosaur phylogeny is a contentious topic, with several mutually exclusive analyses presented in recent years, so the exact position of Ankylosaurus within Ankylosauridae is unknown. Ankylosaurus and Euoplocephalus are often thought to be sister taxa. However, other analyses have found these genera in different positions. Further discoveries or research may clarify the situation.
Like other ankylosaurs, Ankylosaurus was herbivorous. It"s wide muzzle was adapted for non-selective low-browse cropping. The absence of tooth batteries indicates that it did very little chewing. It may have had a hindgut fermentation system like modern herbivorous lizard. This is based on the features of the ribcage.
The tendons of the tail were partially ossified (or bony) and were not very elastic, allowing great force to be transmitted to the end of the tail when it was swung. It seems to have been an active defensive weapon, capable of producing enough of a devastating impact to break the bones of an assailant. A 2009 study showed that "large tail knobs could generate sufficient force to break bone during impacts, but average and small knobs could not", and that "tail swinging behavior is feasible in ankylosaurids, but it remains unknown whether the tail was used for interspecific defense, intraspecific combat, or both". It has also been proposed that the tail club acted as a decoy for the head, although this idea is now largely discredited.
Ankylosaurus magniventris existed between 68 to 66 million years ago, in the final Maastrichtian stage of the Late Cretaceous Period, and was among the last dinosaur species that appeared before the Cretaceous–Paleogene extinction event. The type specimen is from the Hell Creek Formation of Montana, while other specimens have been found in the Lance Formation of Wyoming and the Scollard Formation in Alberta, Canada, all of which date to the end of the Cretaceous.
The Hell Creek, Lance and Scollard Formations represent different sections of the western shore of the shallow sea that divided western and eastern North America during the Cretaceous. They represent a broad coastal plain, extending westward from the seaway to the newly formed Rocky Mountains. These formations are composed largely of sandstone and mudstone, which have been attributed to floodplain environments. The Hell Creek is the best studied of these ancient environments. At the time, this region was subtropical, with a warm and humid climate. Many plant species were supported, primarily angiosperms, with less common conifers, ferns and cycads. An abundance of fossil leaves found at dozens of different sites indicates that the area was largely forested by small trees.Ankylosaurus shared its environment with dinosaurs including the ceratopsids Triceratops and Torosaurus, hypsilophodont Thescelosaurus, hadrosaurid Edmontosaurus, nodosaurid Edmontonia, pachycephalosaurian Pachycephalosaurus, and the theropods Struthiomimus, Ornithomimus, Troodon, and Tyrannosaurus.
Fossils of Ankylosaurus are considerably rare in these sediments, compared to Edmontosaurus and the super-abundant Triceratops, which make up most of the large herbivore fauna. Another ankylosaur, Edmontonia, is also found in the same formations. However, Ankylosaurus and Edmontonia seem to have been separated both geographically and ecologically. The non-selective low browsing of Ankylosaurus may have limited it to the upland regions, away from the coast, while Edmontonia, which had a narrower muzzle for a more selective diet, seems to have lived at lower elevations, closer to the coast.

Tarchia  gigantea
Size estimates of Tarchia have been largely based on Dyoplosaurus giganteus, the holotype of which is one of the largest ankylosaurian individuals known. This would make Tarchia the longest known Asian ankylosaur, with an estimated body length of 8 metres (26 ft).
Estimated at over eight meters long, Tarchia is one of the largest ankylosaurids currently known, rivalling even the more famous Ankylosaurus. In fact given that Ankylosaurus is still known only from partial remains, Tarchia may one day actually prove to be the biggest. Tarchia was named along with another large, but slightly smaller ankylosaurid called Saichania, and although quite similar to one another, there are a number of identifiable differences between the two, particularly differences associated with the skull proportions. Despite these differences however, both Tarchia and Saichania both share bulbous bone growths that are present across the tops of their skulls. A North American ankylosaurid called Nodocephalosaurus also has these bumps, strongly suggesting a possible relationship with Tarchia and Saichania.
Tarchia possessed a wide cropping beak across its mouth that allowed large amounts of vegetation to be indiscriminately pulled into the mouth. These plants would have likely been quite tough considering that Tarchia lived in an arid climate that was near desert in places, and would have required a large degree of processing in the mouth. Evidence for this comes from the teeth which show occlusion wear, basically meaning that the teeth of the upper and lower jaws regularly made contact. Like other ankylosaurids Tarchia had teeth more suited to chopping, and with every up and down movement of the jaw, the food in the mouth would be chopped into smaller and smaller pieces. This was not just to help swallowing but to increase the efficiency of digestion as the teeth chopping the food would provide a greater surface area to be exposed to the digestive acids of the stomach, greatly enhancing the nutritional gain.
Tarchia also had a hard palate and a network of air passages in the snout which would have helped to moisten the dry air of its ecosystem before it reached its lungs. This would greatly reduce the amount of water lost through the process respiration, a vital adaptation considered the climate that Tarchia lived in. Additionally the presence of the hard palate (unknown in most dinosaurs, but seemingly common in ankylosaurids) meant that Tarchia could still breathe while it processed food in its mouth.

Edmontonia was one of the larger nodasaurids, dinosaurs that were similar to the ankylosaurids but lacked their tail clubs and had narrower mouths. Still, Edmontonia did possess armour plates called osteoderms along its back, and is particularly noted for having large spikes that point out from the sides of its body, the four largest of which are above the shoulder. It seems unlikely that these spines were just for defence as they only cover a relatively small part of the body, and they may not have provided all that much protection against large theropod dinosaurs such as tyrannosaurs like Daspletosaurus and Albertosaurus.
Edmontonia was bulky, broad and tank-like. Its length has been estimated at about 6.6 m (22 ft).In 2010, Gregory S. Paul considered both main Edmontonia species, E. longiceps and E. rugosidens, to be equally long at six metres and weigh three tonnes.
Edmontonia had small, oval ridged bony plates on its back and head and many sharp spikes along its sides. The four largest spikes jutted out from the shoulders on each side, the second of which was split into subspines in E. rugosidens specimens. Its skull had a pear-like shape when viewed from above.Its neck and shoulders were protected by three halfrings made of large keeled plates.
It’s plausible that these shoulder spikes may have been more like deer antlers, with the larger and more developed spikes belonging to the more mature individuals. Additionally it is possible that two Edmontonia may have walked up to one another and engaged in a pushing contest for dominance, the horns locking so that the Edmontonia could get a grip of one another. Again larger spikes would have been preferable in such a contest as they would have allowed the more mature individuals a reach advantage.
Study of other fossils found in association with Edmontonia, specifically petrified trees, has come to the conclusion that Edmontonia lived in an environment that saw extended wet and dry periods throughout the year. This would suggest that like other animals in similar climates, Edmontonia would lay eggs to hatch in time for the wet season so that the newly hatched young had a plentiful supply of fresh vegetation. Additionally the adults would have spent much of their time feeding upon as many plants as they could so that they could build up fat reserves to better survive the lean periods of the dry seasons. The oncoming of the wet season is also thought to be the reason why some Edmontonia remains have the spikes and armour in exactly the same position as they were in life. This is simply a case of flood water washing a large amount of sediment over an individual that had died during the dry season, burying and protecting it from carnivorous animals that may have scavenged and pulled the body to pieces had it not been buried from them.

Panoplosaurus is one of the more famous of the North American nodosaurs, and a genus that lived during the later stages of the Cretaceous. Like with other nodosaurs, Panoplosaurus would have looked similar to an ankylosaur, though with a few key differences. The most obvious difference at a glance would have been the lack of a tail club; instead the tail of Panoplosaurus would have terminated in a tapered point like so many other dinosaurs. Also like other nodosaurs, and unlike ankylosaurs, the snout and mouth of Panoplosaurus would have been relatively narrow, possibly indicating that Panoplosaurus was a selective browser of low growing plants.
Like its nodosaur relatives, Panoplosaurus would have had heavy bony scutes running down the back and sides of the body. These scutes, technically known as osteoderms, would have acted like mail armour against the teeth and claws of predatory dinosaurs. However, one of the more common armoured choices of nodosaurs, the large shoulder spikes seen in some other genera, were absent in Panoplosaurus. This might suggest that the presence and arrangement of shoulder spikes in nodosaurs may have also served a display as well as a defensive purpose. With the absence of spikes, Panoplosaurus seem to have favoured the development of very large osteoderms that grew very close together on the neck.

Euoplocephalus  is one of the largest genera of herbivorous ankylosaurian dinosaurs, living during the Late Cretaceous of Canada. It has only one named species, Euoplocephalus tutus. Euoplocephalus was about five to six meters long and weighed over two tons. Its body was low-slung and very flat and wide, standing on four sturdy legs. Its head had a short drooping snout with a horny beak to bite off plants that were digested in the large gut. Like other ankylosaurids, Euoplocephalus was largely covered by bony armor plates, among them rows of large high-ridged oval scutes. The neck was protected by two bone rings. It could also actively defend itself against predators like Gorgosaurus using a heavy club-like tail end.
Among the ankylosaurids, Euoplocephalus was exceeded in size only by Ankylosaurus, and perhaps Tarchia and Cedarpelta. Euoplocephalus was 6 metres (20 ft) long and weighed about 2 tonnes (2.2 short tons). It was also 2.4 metres (7.9 ft) wide. Like other ankylosaurids, it had a very broad and flat low-slung torso, about four feet high, positioned on four short legs.
Whereas Ankylosaurus is the most famous ankylosaurid dinosaur, Euoplocephalus is one of the most important because of the huge number of remains that have been attributed to the genus. These include remains of over forty individual Euoplocephalus, comprising fifteen known skulls and some post cranial skeletons that are almost complete. This wealth of material has not only helped to increase our understanding of Euoplocephalus and the Ankylosauridae as a group, but has also revealed avenues of research that were not previously considered by palaeontologists.

Cedarpelta is an extinct genus of herbivorous basal ankylosaurid ankylosaur, based on material recovered from the Lower Cretaceous of North America. The skull lacks extensive cranial ornamentation, a trait which has been interpreted as plesiomorphic for ankylosaurs.
Cedarpelta was a large ankylosaurian. In 2010, Gregory S. Paul indicated a body length of seven metres and a weight of five tonnes. Kenneth Carpenter estimated it smaller at five metres. Carpenter et al. (2001) established several distinguishing traits of Cedarpelta. The body of the praemaxilla, the front snout bone, is short in front of its nasal branch. The outer sides of the two praemaxillae run more parallel compared to the snouts of later forms which are strongly diverging to behind. The cutting edge of the bone core of the upper beak is limited to the front of the praemaxilla. Each praemaxilla has six (conical) teeth. The quadrate, and with it the entire back of the skull, is inclined to the front. The head of the quadrate is not fused with the paroccipital process, contrary to the situation in Shamosaurus. The neck of the occipital condyle is long and sticking out to behind, like with nodosaurids, not obliquely to below as in typical ankylosaurids. The tubera basilaria, appending processes of the rear lower braincase, form a large wedge directed to below. The pterygoid is elongated from the front to the rear and has a saddle-shaped process on its outer edge oriented to behind and sideways. The coronoid process of the rear lower jaw has an oval process at the inside. The straight ischium has a knob-shaped boss at the inside near the pubic pedicle.
Cedarpelta shows a mix of basal ("primitive") and derived traits. The presence of premaxillary teeth is a plesiomorphic character because it is inherited from earlier Ornithischia. In contrast, closure of the opening on the side of the skull behind the orbit, the lateral temporal fenestra, is an advanced, derived (apomorphic) character only known in ankylosaurid ankylosaurians.
Two skulls are known, and the skull length for Cedarpelta is estimated to have been roughly 60 centimetres (24 in). One of the Cedarpelta skulls was found disarticulated, a first for an ankylosaur skull, allowing paleontologists a unique opportunity to examine the individual bones instead of being limited to an ossified unit. The skull is relatively elongated and does not show a strongly appending beak. Of the conical premaxillary teeth, the first is the largest. The maxilla bears eighteen teeth. The eye socket is surrounded by the lacrimal, a single supraorbital and a large postorbital, excluding the prefrontal and the jugal from the orbital rim. The postcranial skeleton was in 2001 not described in any detail.
The skulls, though of large and thus not juvenile individuals, do not show a distinctive pattern of fused caputegulae, head tiles. This inspired Carpenter to propose an alternative hypothesis of ankylosaur skull osteoderm formation. Formerly, it had been assumed that such armour plates were either formed by direct skin ossification into distinct scutes which later fused to the skull (the more popular theory), or by a reaction of the skull bones to the pattern of overlying scales. The lack of a clear pattern in Cedarpelta suggested to Carpenter that the ossification took place in an intermediate layer between the scales and the skull roof itself, which he surmised to have been the periosteum.





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