Euoplocephalus

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Euoplocephalus
Temporal range: Late Cretaceous, 76.4–75.6Ma
Reconstruction of specimen ROM 1930, Senckenberg Museum
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Ornithischia
Family: Ankylosauridae
Subfamily: Ankylosaurinae
Genus: Euoplocephalus
Lambe, 1910
Type species
Stereocephalus tutus
Lambe, 1902
Species

Euoplocephalus tutus (Lambe, 1902)

Synonyms[1]

Stereocephalus tutus
Lambe, 1902 (preoccupied)

Euoplocephalus (/juːˌɒplɵˈsɛfələs/ ew-OP-lo-SEF-ə-ləs) is one of the largest genera of herbivorous ankylosaurian dinosaurs, living during the Late Cretaceous of Canada.

The first fossil of Euoplocephalus was found in 1897 in Alberta. In 1902, it was named Stereocephalus, but that name had already been given to an insect, so it was changed in 1910. Later, many more ankylosaurid remains were found from the Campanian of North America and often made separate genera. In 1971, Walter Coombs concluded that they all belonged to Euoplocephalus which then would be one of the best-known dinosaurs. Recently however, experts have come to the opposite conclusion, limiting the authentic finds of Euoplocephalus to about a dozen specimens. These include a number of almost complete skeletons, so much is nevertheless known about the build of the animal.

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.

Etymology[edit]

The genus name Euoplocephalus, meaning "well-armed head", is derived from the Greek words eu (εὖ) meaning "well", hoplo~ (ὁπλο~) meaning "armed", and kephale (κεφαλή) meaning "head".[2] This name has been misspelled more than a dozen different ways in formal scientific literature. The specific name tutus means "safely protected" in Latin. The only valid species known today is Euoplocephalus tutus.

Description[edit]

Size comparison between Euoplocephalus and a human

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).[3] 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.

The skull of Euoplocephalus can be distinguished from most other ankylosaurids by several anatomical details, including: the pattern of bony sculpturing in the region in front of the eyes; the form of the palpebral bones (small bones over the eyes),[4] which may have served as bony eyelids;[5] the shallowness of the nasal vestibule at the entrance of the nasal cavity;[4] the medial curve of the tooth rows in the upper jaw; and the teeth, which are relatively small, lacking true cingula, and having variable fluting of the denticles.[4] However, these traits are shared with a number of closely related forms, some of which in the past have been included in the genus.[6]

Skulls AMNH 5405 in ventral (B) and anterior (C) views, ROM 1930 in ventral (E) and posterior (F) views

The skull resembles a truncated equilateral triangle when viewed from above, and is slightly wider than it is long. The largest known skull, that of specimen AMNH 5403, has a length of 411 millimetres and a width of 478 millimetres. Most skull elements are completely fused and two skull openings normally present with dinosaurs, the antorbital fenestra and the upper temporal fenestra, have closed. The skull has nineteen to twenty-four teeth in each upper jaw. The frontmost snout bone, the premaxilla, is toothless. The teeth are very small, with a maximal height and width of just 7.5 millimetres. The strongly drooping snout is blunt, wide and high, and filled with very complex air passages and sinuses, the form and function of which are not yet completely understood. Each side has two external nostrils. The lower jaw has a very concave upper rim with twenty-one teeth. At its front a short low extension is present, to provide contact with the wide predentary, the bony core of the lower horny beak, that fitted within the upper beak of the snout.[4]

Mandibles of three specimens

As in most quadrupedal ornithischians, its neck is moderately short. The scapula is massive and robust, and the very robust forelimbs are shorter than the hindlimbs.[7] The tail is long and ends in a bony club. Old restorations of Euoplocephalus and rejected synonyms (Dyoplosaurus, Scolosaurus) often show a club with two large vertical spikes. This is an error based on a restoration of Scolosaurus by Franz Nopcsa; the specimen he used had an incomplete tail that stopped just beyond the pair of conical spikes now known to have been positioned halfway along its length. He restored the tail as ending just after the spines. Other artists combined the spikes with the tail club, compounding the inaccuracy.[7] The narrow distal half of the tail is stiffened by bundles of ossified tendons.

The vertebral column of Euoplocephalus is made up of at least seven cervical vertebrae, at least eleven "free" dorsal vertebrae, typically four sacrodorsals forming a fused "sacral rod" in front of the sacrum proper, three "true" sacral vertebrae, and between one and four caudosacrals.[6] Like with other ankylosaurians,[1] the last four dorsal vertebrae and the first caudal vertebra are thus fused to the sacrum, forming a reinforced synsacrum of at least eight vertebrae,[7] the spines of which form a fused supraneural plate, also incorporating the zygapophyses. There are at least twenty-one caudal vertebrae; the total number of caudal vertebrae is uncertain because approximately ten are fused to form part of the tail club,[7] bringing the total to about thirty. This fusion is also seen in other ankylosaurids; it is possible that the extent of fusion is an age-related feature.[8][9]

Pelvis of specimen AMNH 5337

The humerus is very robust with strongly expanded upper and lower joints, combined with a narrow shaft. On the upper shaft an enormous deltopectoral crest is present of which the lower part does not gradually merge with the shaft but is warped to the front, forming a thick knob or lip. All this indicates a very heavy musculature. In the lower arm the robust ulna has a well-developed olecranon process. The wrist and hand bones are not well known. In the pelvis, the front blade of the ilium splayed out to the front, reaching all the way to the widest point of the belly to support the gut. This blade also forms a bone shelf at the rear side of the body. The rear blade of the ilium is shorter than the diameter of the hip socket it was located behind, meaning the leg is located at the rear end of the pelvis, near the tail base and much closer to the midline than the belly sides. The pubic bone is unknown. The ischium is a short, curved, vertically positioned bone strap. The thighbone is short, robust and straight with a low fourth trochanter positioned below the midpoint of the shaft. The robust shinbone is shorter than the thighbone. The foot is not well known but functionally tridactyl with hoof-shaped instead of sharp claws.[6]

Armor[edit]

Cervical half rings and osteoderms

The head and body of Euoplocephalus were covered with bony armor, except for parts of the limbs and possibly the distal tail. The armor consisted of osteoderms, skin ossifications that are not part of the skeleton proper. This armor was in 1982 extensively described by Kenneth Carpenter, who however, largely based himself on the very complete specimen NHMUK R5161, the holotype of Scolosaurus,[7] which genus no longer is seen as a synonym of Euoplocephalus. When limited to the certain material of the latter genus, little is known of the exact configuration of the armor, with the exception of the head and the neck. The most informative specimen in this respect would then be ROM 1930, having conserved some osteoderms of the torso in their original position.[6]

Restoration

In any case, much of the armor was made up of small ossicles,[10] bony round scutes with a diameter of less than five millimetres, of which often hundreds have been found with a single specimen. If the armor was configured in an identical way to that of Scolosaurus, many of these small ossicles had fused into a kind of pavement, forming transverse bands on the body. The banded arrangement is thought to have permitted some freedom of movement.[7] Four of these bands might have been present on the anterior half of the tail, three on the pelvis, perhaps fused into a single "sacral shield", and four across the front part of the torso. Inset in these bands were horizontal rows of larger oval, flat or keeled, scutes. Types of large scutes varied by body region. It might be that the scutes on the shoulder, near the midline of the body, were largest and tallest; ROM 1930 includes some osteoderms with a base length of fifteen centimeters. Little is known about the armor of the limbs. Large keeled plates were present on the upper arms as shown by specimen TMP 1997.132.01 conserving a round osteoderm near the humerus with a diameter of twenty centimeters and narrower spikes associated with the lower arm. The neck was protected by two bone rings, open at the underside, that are called "cervical half-rings". Earlier seen as a fusion of osteoderms,[7] this was doubted by Arbour et al. in 2013, who pointed out that they formed a lower layer, possibly consisting of ossified cartilage, as indicated by a smooth surface and a woven bone texture. Each half-ring is constructed out of six rectangular concave plates, three per side. Each plate has a large keeled osteoderm on top, often not fused with it. With Euoplocephalus, these neck osteoderms do not have smaller osteoderms at their bases, and their keels do not overhang their posterior edges.[6]

Skull AMNH 5404, with arrow pointing at eyelid armor

The armor of the skull consists of a large number of ossicles, called caputegulae ("head tiles"), that have fused with the normal skull elements, largely fading their sutures. On the snout they form a chaotic and asymmetric mosaic. On the rear nasal region, at the midline a single hexagonal larger plate is present. A keeled plate per side, somewhat more to the back forms the snout rim. Behind the level of the eye sockets the caputegulae fuse into a single bone surface. The upper rim of the eye sockets is formed by two pyramid-shaped osteoderms pointing to the sides and rear. In addition, Euoplocephalus had two pyramid-shaped squamosal "horns" growing from the back corners of its head. Between them the nuchal crest is covered by two osteoderms per side. At the lower rear side of the skull, a quadratojugal horn is present, in the form of an enormous tongue-shaped osteoderm projecting below.[6]

The armor of Euoplocephalus may have had a keratinous covering, or it may have floated in the skin, as is seen in modern crocodiles. In addition to protection, the heavily-vascularized armor may have had a role in thermoregulation.[7] The palpebral bones over the eyes may have provided additional protection for the eyes. Such bones have with Euoplocephalus been discovered in the upper part of the eye socket, instead of in front of the upper socket rim which is the more common position. Coombs explained this by assuming that these bones were located in the eyelid musculature and were probably mobile enough to be moved over the eyes.[5]

Discovery and species[edit]

Skull of the holotype CMN 0210

Canadian paleontologist Lawrence Morris Lambe discovered the first specimen on 18 August 1897 in the area of the present Dinosaur Provincial Park, in the valley of the Red Deer River. In 1902, this fossil, CMN 210 (also NMC 210), which consists of the upper part of a cranium and a transverse series of five scutes, was designated as the holotype specimen of the type species Stereocephalus tutus. The generic name was derived from Greek στερεός, stereos, "solid", and κεφαλή, kephalè, "head", which refers to the formidable armour.[11] However, the genus name was already preoccupied — the name had already been given to an insect, the beetle Stereocephalus Lynch 1884 — so Lambe changed it to Euoplocephalus in 1910, with as combinatio nova (new combination name) Euoplocephalus tutus.[12] The type species remains Stereocephalus tutus. In 1915, Edwin Hennig classified E. tutus under the genus Palaeoscincus Leidy 1856, coining a Palaeoscincus tutus.[13] Today however, Palaeoscincus is considered to be a nomen dubium based on indeterminate ankylosaurian teeth. In 1964, Euoplocepohalus was by Oskar Kuhn referred to Ankylosaurus, as a Ankylosaurus tutus.[14]

Referred material[edit]

During the early twentieth century many more ankylosaurid fossils were uncovered in North America. Some were referred to Euoplocephalus, others named as separate genera. In 1971 however, Walter Coombs submitted a dissertation containing a landmark re-appraisal of North American ankylosaurs. He noted that, among the many specimens similar to Euoplocephalus, their skulls varied so much that either every known specimen must be a new species, or they all represented individual variation within a single species: Euoplocephalus tutus.[8] Starting from this assumption that there was only one species of ankylosaur during the Campanian stage of the Upper Cretaceous, Coombs synonymized the genera Anodontosaurus, Dyoplosaurus, and Scolosaurus with Euoplocephalus and the species A. lambei, D. acutosquameus, and S. cutleri with E. tutus, creating a species that spanned nearly ten million years, or the entire Campanian.[15] The fossils now referred to this species contained more than forty individuals discovered in Alberta, Canada and Montana in the United States, which would have made Euoplocephalus the best known ankylosaurid. This included fifteen skulls, teeth, and a few almost-complete skeletons, found with the armor still attached.[1] Individual armor plates are the most commonly found element from them. In 1978, Coombs even included the Asian ankylosaurid Tarchia in the genus, renaming it as Euoplocephalus giganteus.[16]

Skull of referred specimen AMNH 5405

The synonymy of all Campanian North-American ankylosaurids was followed for several decades,[1] until scientists from the University of Alberta began to re-examine the fossils. A recent study found that Dyoplosaurus is in fact a valid taxon, and identified unique characteristics that differentiated it from Euoplocephalus, including its triangular claws.[15] Victoria Arbour (2010) argued that Anodontosaurus (known from the Horseshoe Canyon Formation) is distinct from Euoplocephalus and is also a valid taxon; according to Arbour, Anodontosaurus differs from Euoplocephalus in distinctive skull and cervical half ring ornamentation, as well as tail club morphology, including the presence of pointed, triangular knob osteoderms in Anodontosaurus. Furthermore, Arbour (2010) suggested to re-assign all Horseshoe Canyon Formation ankylosaurine specimens from Euoplocephalus to Anodontosaurus.[17] The validity of Anodontosaurus was accepted in two subsequent studies. The first, published by Paul Penkalski and William T. Blows in 2013, re-validated Scolosaurus as well.[18] The second study, by Penkalski (2013), named and described Oohkotokia from Montana on the basis of remains that were originally thought to be referable to Euoplocephalus.[19]

Palaeoscincus asper, "the rough one", is now considered to be Euoplocephalus. It is a dubious tooth taxon from the late Campanian Dinosaur Park Formation of Alberta, Canada, named by Lawrence Morris Lambe in 1902.[20] It consists of a single tooth, specimen NMC 1349.[21]

In 2013 Arbour limited the specimens that could be reliably referred to Euoplocephalus to the lowest thirty metres of the Dinosaur Park Formation. The material would in that case, apart from the holotype, consist of partial skeletons with skull AMNH 5337, AMNH 5403, AMNH 5404, AMNH 5405, ROM 1930 and UALVP 31; partial skeleton lacking the skull AMNH 5406; CMN 842, a cervical half-ring; CMN 8876, a skull, TMP 1979.14.74, a fragmentary skull; and UALVP 47977, a skull roof piece. The hands, feet and tail, including the club, are therefore not completely known.[6]

Phylogeny[edit]

Skull of specimen UALVP 31
Skull of specimen ROM1930
Skull of specimen AMNH 5403

In 1910, Lambe assigned Euoplocephalus to the Stegosauria, a group then encompassing all armoured dinosaur forms and thus having a much wider range than the present concept. In 1917, Charles Whitney Gilmore assigned it to the Ankylosauridae.[22] Today, Euoplocephalus is still seen as an ankylosaurid, but as a member of the Ankylosauria, not the Stegosauria. It is likely also a member of the derived subgroup Ankylosaurinae. The recent splitting of the ankylosaurid Campanian material of North America has complicated the issue of the direct affinities of Euoplocephalus. Penkalski (2013) performed a small phylogenetic analysis of some ankylosaurine specimens. The only Anodontosaurus specimen that was included in this analysis was its holotype. Anodontosaurus was placed in a polytomy with the holotype of Euoplocephalus and some specimens that are referred to it, while Oohkotokia was placed in a clade with Dyoplosaurus, and specimens that are thought to represent either Dyoplosaurus or Scolosaurus.[19] According to the 2013 study of Arbour, Euoplocephalus has a basal position in the Ankylosauridae, perhaps forming a clade with Anodontosaurus that is the sister taxon of Ankylosaurus.[6]

The following cladogram is based on a phylogenetic analysis conducted by Arbour and Currie (2013):

Ankylosaurinae

Scolosaurus (=Oohkotokia horneri)





Ankylosaurus




Anodontosaurus



Euoplocephalus







Dyoplosaurus



Talarurus



Tsagantegia





Minotaurasaurus



Nodocephalosaurus



Tianzhenosaurus





Pinacosaurus grangeri



Pinacosaurus mephistocephalus





Tarchia



Saichania










The results of an earlier analysis of the ankylosaurid tree by Thompson et al. (2011), is shown by this cladogram.[23]

Ankylosauridae

Minmi




Liaoningosaurus




Cedarpelta




Gobisaurus




Shamosaurus


Ankylosaurinae

Tsagantegia



Zhongyuansaurus





Shanxia




Crichtonsaurus




Dyoplosaurus



Pinacosaurus mephistocephalus








Ankylosaurus



Euoplocephalus






Minotaurasaurus



Pinacosaurus





Nodocephalosaurus





Talarurus



Tianzhenosaurus





Tarchia



Saichania














Distinguishing anatomical features[edit]

Skull of specimen TMP 1991.127.1

A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism.

According to Coombs and Maryanska (1990), Euoplocephalus can be distinguished based on the following characteristics:

  • the premaxillae are not covered by dermal ossifications
  • The external nostrils are slit-like, face rostrally and are divided by the vertical septum
  • The beak has a width equal to or greater than the distance between the caudal-most maxillary teeth
  • The pes has three digits

In 2013 Victoria Arbour and Phil Currie provided a differential diagnosis, setting Euoplocephalus apart from its nearest relatives. When compared with Anodontosaurus and Scolosaurus, Euoplocephalus lacks round osteoderms at the base of the squamosal and quadratojugal horns. Compared with Anodontosaurus it lacks small osteoderms at the bases of the large osteoderms on the first cervical half-ring, but, contrary to that genus, does in top view have large rounded osteoderms at the sides of the tail club. It differs from Dyoplosaurus in possessing sacral ribs that perpendicularly point outwards. It differs from Scolosaurus in possessing keeled osteoderms with a round or oval base on the top and sides of the first cervical half-ring and having a shorter rear blade of the ilium. Euoplocephalus differs from Ankylosaurus in possessing anteriorly directed external nostrils and in lacking a continuous keel between the squamosal horn and the supraorbitals.[6]

Paleoecology[edit]

Euoplocephalus, following the synonymizations proposed by Coombs (1971), was thought to exist for far longer, and was a member of more distinct faunas, than any of its contemporaries, as these fossils date to between 76.5 and 67 million years ago, in the Campanian-Maastrichtian ages of the late Cretaceous period, and came from the Dinosaur Park and Horseshoe Canyon Formations of Alberta, Two Medicine Formation of Montana, and possibly from the Oldman Formation of Montana.[15][19] Fossils that were initially believed to be from the Judith River Formation of Montana,[15] are actually from the Dinosaur Park Formation.[24] However, recent studies referred all Horseshoe Canyon Formation specimens to Anodontosaurus,[17] and all Two Medicine Formation specimens to Oohkotokia.[19] A specimen from the lowermost Dinosaur Park, or possibly from the underlying Oldman Formation, was reassigned back to Scolosaurus.[18] Although the stratigraphic range of the holotype of Euoplocephalus is uncertain, all specimens that can be reliably referred to E. tutus came from the lower 40 m and the upper >10 m of the Dinosaur Park Formation. There are no known ankylosaurids from the top 20–25 m of the Formation. Thus, all Euoplocephalus specimens date to between 76.4 (or less) and 75.6 million years ago, in the late Campanian stage.[19]

Paleobiology[edit]

Restored skeleton of ROM 1930, Royal Tyrrell Museum

According to Coombs, Euoplocephalus may have had cursorial abilities on par with those of the modern rhinoceros and hippopotamus.[25] Based on the form of the humerus-shoulder articulation and the arrangement of the protracting muscles of the upper arm, it appears that the upper arm sloped away from the body.[26] Coombs and Maryanska (1990) observed that Euoplocephalus specimens are usually discovered as isolated elements or partial skeletons, which suggested that this animal engaged in solitary habits and was usually either solitary or participated in small group clusters.

Tail club[edit]

The tail club of ankylosaurids has often been interpreted as a defensive weapon. In Euoplocephalus, the presence of ossified tendons only with the distal half of the tail may support such a function.[27] Because only the distal half of the tail was stiffened by tendons, the anterior half could still move freely from side to side. The ossified tendons would have transmitted the force of the swing to the club and reinforced the supporting vertebrae.[7] The club was likely held just above the ground, as there was not sufficient musculature to raise the tail very high.[28] A 2009 study concluded that "large ankylosaurian clubs could generate sufficient force to break bone during impacts, while average and small ones could not".[29] It has also concluded that "tail swinging behavior is feasible in ankylosaurids, but it remains unknown whether the tail was used for interspecific defense, intraspecific combat, or both".[29] The tail club could be swung low, toward the fragile metatarsals or shin bones of attacking theropods.

Coombs observed three different types of tail clubs, differentiated by their shape: round, elongate, or bluntly pointed. The variation can be explained either ontogenically, which means that the shape of the club changed as the animal proceeded through growth stages in its life, or may be attributable to individual variation. Coombs doubted that the differences in tail club morphology represented sexual dimorphism. Coombs described specimen ROM 7751, which is the smallest known ankylosaurid tail club, probably belonging to a subadult that lived in what is now the Dinosaur Park Formation in Alberta.

Senses[edit]

Skull of specimen AMNH 5337

Euoplocephalus had relatively small eyes, but this does not necessarily mean that it had restricted vision. The complex respiratory passages observed in the skull suggest that Euoplocephalus had a good sense of smell, although in 1978 an examination of casts of the endocranium did not show an enlarged olfactory region of the brain.[30] Teresa Maryanska, who has worked extensively on Mongolian ankylosaurids, suggested that the respiratory passages were primarily used to perform a mammal-like treatment of inhaled air, based on the presence and arrangement of specialized bones,[9] which are present in Euoplocephalus.[31] A 2011 study suggested that Euoplocephalus used its nasal passages for sound production and researchers discovered an enlarged and vascularised chamber at the back of the nasal tract, which was considered by the authors to be an adaptation to improve its sense of smell. The researchers also managed to reconstruct the dinosaur's inner ear and concluded that it was capable of hearing at low frequencies.[32]

Diet[edit]

Euoplocephalus, like other ankylosaurians, is thought to have been a herbivore.[1] It had a broad muzzle, which could indicate that it was a non-selective feeder, perhaps similar to a hippopotamus. This would provide niche separation from contemporaneous nodosaurids with narrower muzzles.[7] Ankylosaurians have historically been thought of as feeding using simple up-and-down movements of the jaws, but Euoplocephalus appears to have been able to make more complex movements. Tooth wear and jaw articulations (within the lower jaw and at the lower jaw-quadrate joint) suggest that the lower jaws were pulled back during feeding, and also slightly pivoted inward. This action would have sheared food.[33] Haas (1969) examined the evidence for the jaw muscles of two skulls (AMNH 5337 and 5405) and concluded that despite the large size of the skulls the associated musculature was relatively weak. Haas extrapolated that dinosaurs like Euoplocephalus likely ate relatively soft vegetation.[34]

References[edit]

  1. ^ a b c d e Vickaryous, M.K., Maryanska, T., and Weishampel, D.B. (2004). "Ankylosauria" In D. B. Weishampel, P. Dodson, and H. Osmolska (eds.), The Dinosauria (second edition). University of California Press, Berkeley 363-392.
  2. ^ Liddell, Henry George and Robert Scott (1980). A Greek-English Lexicon (Abridged Edition). United Kingdom: Oxford University Press. ISBN 0-19-910207-4. 
  3. ^ Bakker, R. T. (1980) "Dinosaur heresy-dinosaur renaissance". In D. K. R. Thomas, and E. C. Olson (eds.), A cold look at the warm blooded dinosaurs, pp. 351–462. Westview Press, Boulder, Colorado.
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  8. ^ a b Coombs W. (1971) The Ankylosauridae. Ph.D. thesis, Columbia University, New York, NY, 487 p.
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  10. ^ http://www.merriam-webster.com/dictionary/dermal%20ossicle
  11. ^ L. M. Lambe. (1902). "New genera and species from the Belly River Series (mid-Cretaceous)". Geological Survey of Canada Contributions to Canadian Palaeontology 3 (2): 25–81. 
  12. ^ L. M. Lambe. (1910). "Note on the parietal crest of Centrosaurus apertus and a proposed new generic name for Stereocephalus tutus". The Ottawa Naturalist 24: 149–51. 
  13. ^ Hennig, E., 1915, Stegosauria: Fossilium Catalogus I, Animalia pars 9, 16 pp
  14. ^ Kuhn, O., 1964, Ornithischia: Fossilium Catalogus, I: Animalia, Pars 105, 80 pp
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  16. ^ Coombs, W.P. Jr., 1978, "The families of the ornithischian dinosaur order Ankylosauria", Palaeontology 21: 143–170
  17. ^ a b Arbour, Victoria (2010). "A Cretaceous armoury: Multiple ankylosaurid taxa in the Late Cretaceous of Alberta, Canada and Montana, USA". Journal of Vertebrate Paleontology 30 (Supplement 2): 55A. doi:10.1080/02724634.2010.10411819. 
  18. ^ a b Penkalski, P.; Blows, W. T. (2013). "Scolosaurus cutleri (Ornithischia: Ankylosauria) from the Upper Cretaceous Dinosaur Park Formation of Alberta, Canada". Canadian Journal of Earth Sciences: 130110052638009. doi:10.1139/cjes-2012-0098.  edit
  19. ^ a b c d e Penkalski, P. (2013). "A new ankylosaurid from the late Cretaceous Two Medicine Formation of Montana, USA". Acta Palaeontologica Polonica. doi:10.4202/app.2012.0125.  edit
  20. ^ Lambe, L.M. (1902). "On Vertebrata of the mid-Cretaceous of the Northwest Territory. 2. New genera and species from the Belly River Series (mid-Cretaceous)". Contributions to Canadian Paleontology 3: 25–81. 
  21. ^ Coombs, Jr., W.P. (1990). Teeth and taxonomy in ankylosaurs. In: Carpenter, K., and Currie, P.J. (eds.). Dinosaur Systematics: Approaches and Perspectives. Cambridge University Press:Cambridge, 269-279. ISBN 0-521-36672-0
  22. ^ C.W. Gilmore, 1917, "Brachyceratops, a ceratopsian dinosaur from the Two Medicine Formation of Montana, with notes on associated fossil reptiles", United States Geological Survey Professional Paper 103: 1-45
  23. ^ Richard S. Thompson, Jolyon C. Parish, Susannah C. R. Maidment and Paul M. Barrett (2011). "Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora)". Journal of Systematic Palaeontology 10 (2): 301–312. doi:10.1080/14772019.2011.569091. 
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