Ankylosauria
Ankylosauria consists of
three main
groups, Ankylosauridae, Nodosauridae, and Polacanthidae, plus some basal forms. All ankylosaurians
had armor over much of their bodies, mostly scutes and nodules, with large spines in some
cases. The scutes, or plates, are rectangular to oval objects organized in transverse
(side to side) rows, often with keels on the upper surface. Smaller nodules and
plates filled in the open spaces between large plates. In all three groups the first two rows of plates tend to form a sort of half-ring around the
neck; in nodosaurids, this comes from adjacent plates fusing with each other
(and there is a third row as well), while
ankylosaurids usually have the plates fused to the top of another band of bone. The
skull has armor plastered on to it, including a distinctive piece on the outside-rear of
the lower jaw. Feeding was generally accomplished similarly to how Scelidosaurus
did it, with precise
meeting of the teeth leading to a puncture-crush style like a mortar and pestle.
Besides body armor, which is also known lower
in Thyreophora, all ankylosaurians (except Mymoorapelta)
rewalled the acetabulum (the socket for the femur; it
is open in most dinosaurs) for strength and flared the upper surface of the ilium
outward. Ankylosaurians were very tubby beasts, suggesting that their somewhat
feeble teeth were backed up by an impressive internal digestion system.
Ankylosaurians are a mostly K group from North America and Asia, although
isolated finds put them in a number of odd places, including the Campanian (LK)
of Antarctica, the late Maastrichtian (LK) and the Toarcian (EJ) of India, and the
Campanian-Maastrichtian (LK) of Argentina. Interestingly, ankylosaurians
as a group are also the second-most common dinosaurs, next to hadrosaurids,
found in marine sediments; this likely is due to their solid construction and
(inferred) spacious digestive tracts and other viscera, allowing whole bodies to float out
to sea with surprising buoyancy despite the armor.
These classifications of Ankylosauria and Ankylosauridae
presented here are very tentative; much new material has been found in recent
years, and these classifications will doubtless be altered. For one new
look at these armored beasts, see The
Tree of Life's ankylosaur pages, and for anyone wishing to read more about
armored dinosaurs in general, and who isn't daunted by the technical literature,
there is The Armored Dinosaurs, 2001, Kenneth Carpenter (ed), which is
where I got much of the material for the reclassifications I've undertaken here.
<--Ankylosauria
|--Minmi
`--Ankylosauroidea
|-->Nodosauridae
|-->Polacanthidae
`-->Ankylosauridae
Ankylosauria:
| Taxon or Taxa: | Time\Place: | Comments: |
| Minmi paravertebra Molnar, 1980 | Aptian-Albian (EK) of Australia | This small ankylosaurian was first thought to be a nodosaurid, but is too generalized. It has unusual structures along its spine, called paravertebrae, whose function and purpose are still unclear; they may have strengthened the back and allowed for faster locomotion than other ankylosaurians. It is known from the partial type specimen and a nearly complete skeleton referred to M. sp.; the armor of the latter shows large scutes on the neck and flanking the shoulders and hips, with additional plate-like scutes on the tail (uncertain orientation). Possible gut contents show short lengths of vegetation not ground by gastroliths. |
Ankylosauria i.s.:
| Taxon or Taxa: | Time\Place: | Comments: |
| ?"Acanthopholis" eucercus (N.D.) Seeley, 1869 | late Albian (EK) of England | This animal is doubtfully ankylosaurian. |
| Aletopelta coombsi Ford and Kirkland, 2001 | mid Campanian (LK) of "California" (paleo-Mexico) | California's first unique named dinosaur, Aletopelta is based on a partial skeleton, originally thought to belong to a nodosaurid. It was found on a tectonic block that appears to have been rafted north from Mexico, making Aletopelta (whose name means "wandering shield" referring to this change in location) a representative of a heretofore-unknown ankylosaurid radiation, according to the authors. Unlike most ankylosaurids, it had some respectable plates and spines, and the pelvis was covered with locking scutes. It may have been a "stegopeltine" or a traditional ankylosaurid, but nodosaurid is most likely. |
| Antarctopelta oliveroi Salgado and Gasparini, 2006 | late Campanian (LK) of Antarctica | First published on in 1991, Anarctopelta becomes the second-named classic dinosaur from Antarctica (well, technically James Ross Island; I don't know if that should count). The skeleton took several years to excavate, what with the frozen ground and all, and consists of part of a dentary, teeth, cranial armor, two cervicals and casts of three others, fragmentary dorsal ribs, a couple of dorsals from the presacral rod, three sacrals, eight caudals, part of a scapula, ilium, and femur, five metapodials, and assorted armor. The owner was about 4 m long, with features of both Nodosauridae and Ankylosauridae. |
| Bienosaurus lufengensis Dong, 2001 | Sinemurian (EJ) of China | This animal is based on a partial skull (well, mostly the lower jaw) first discovered in 1938. It may be related to Scelidosaurus, or just be a regular basal ankylosaurian. |
| Cryptosaurus eumerus (N.D.) Seeley, 1869 | Oxfordian (LJ) of England | Based upon an unusual femur, this animal is better known in the literature as Cryptodraco (a replacement name that turned out to be unnecessary). Cryptosaurus is the true name. |
| Danubiosaurus anceps (N.D.) Bunzel, 1871 (?Struthiosaurus) | early Campanian (LK) of Austria | This is one of a host of indeterminate LK European nodosaurids that may or may not be the same as Struthiosaurus. |
| Dracopelta zybszewskii Galton, 1980 | Kimmeridgian (LJ) of Portugal | Dracopelta is one of the better-known early ankylosaurians, based on material including part of a rib cage and some armor in situ. |
| "Hanwulosaurus" (N.N.) Anonymous, 2001 | ?LK of China | This unofficially-named animal is apparently a very large (~9m long) ankylosaurian, known from many remains including a large flat skull, verts, ribs, scapula, ulna, femorae, fibulae, and armor. It may belong to a new clade within Ankylosauria. |
| Hierosaurus sternbergi (N.D.) Wieland, 1909 | late Coniacian (LK) of Kansas | Sometimes synonymized with Nodosaurus, this animal is rather late in the fossil record to be so. It is known from armor. |
| "Iguanodon" phillipsii (N.D.) Seeley, 1869 | Oxfordian (LJ) of England | Better known as Priodontognathus Seeley, 1875, this is a poorly-known early ankylosaurian based on a partial upper jaw. It was once thought to be a stegosaurian. |
| Liaoningosaurus paradoxus Xu, Wang, and You, 2001 | mid Barremian-early Aptian (EK) of China | Liaoningosaurus is based on the almost-complete remains of a juvenile ankylosaurian (?nodosaurid) from the Yixian Formation. It is incredibly small (femora are 2.8 cm long, for example; body length maybe 34 cm, or a little over a foot)) and includes an interesting and unprecedented shell-like bony plate shielding the abdomen. |
| Onychosaurus hungaricus (N.D.) Nopcsa, 1902 (?Struthiosaurus) | Campanian (LK) of Romania | This is an indeterminate ankylosaurian based on armor. |
| Palaeoscincus costatus (N.D.) Leidy, 1856 (?Edmontonia) | late middle Campanian (LK) of Montana and Alberta | Palaeoscincus is a (in)famous ankylosaurian from the early days of American dinosaur paleo based on teeth. At one time, it was commonly restored with a tail club like an ankylosaurine. |
| Rhodanosaurus ludgunensis (N.D.) Nopcsa, 1929 (?Struthiosaurus) | Maastrichtian (LK) of France | Based on armor plates, this animal is an obscure indeterminate ankylosaurian, and yes, it may be the same as Struthiosaurus. |
| Sarcolestes leedsi Lydekker, 1893 | Callovian (MJ) of England | One of the earliest nodosaurids (?or polacanthid; well, ankylosaurian at any rate), Sarcolestes is based on a jaw once thought to have belonged to a theropod. |
| Sauroplites scutiger (N.D.) Bohlin, 1953 | Barremian-early Aptian (EK) of China | Sauroplites may be the same as the better-known Shamosaurus. |
| Stegosaurides excavatus (N.D.) Bohlin, 1953 | LK of China | This is just an obscure, indeterminate ankylosaurian. |
| Tianchiasaurus nedegoatpeferima (?N.D.) Dong, 1993 | Bathonian (MJ) of China | This is one of the most basal known ankylosaurians. Its odd species name comes from the cast of Jurassic Park (the name for a while was going to be Jurassosaurus nede...), wherein the first two letters of the last names of the actors were combined. Unfortunately, its remains are rather fragmentary. |
Nodosauridae, Polacanthidae, and Ankylosauridae: Next come the good old nodosaurids, the spiky, nodosaurid-like polacanthids, and two varieties of ankylosaurids: the long-skulled basal shamosaurines; and the ankylosaurines, with squat skulls and bodies.
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