Edmontonia

Edmontonia was an armoured dinosaur, part of the nodosaur family from the Late
Cretaceous Period. It is named after the Edmonton Formation (now the Horseshoe
Edmontonia
Canyon Formation in Canada), the unit of rock it was found in. Temporal range: Late Cretaceous ,
76.5–69 Ma

PreЄ Є O S D C P T J K PgN

Contents
Description
Size and general build
Distinguishing traits
Skeleton
Osteoderms
Discovery and species
Phylogeny
Paleobiology
Mounted skeleton of E. rugosidens,
Function of the armour
Paleoecology specimen AMNH 5665

See also Scientific classification

References Kingdom: Animalia
Phylum: Chordata
Description Clade: Dinosauria
Order: †Ornithischia
Size and general build Family: †Nodosauridae
Edmontonia was bulky, broad Genus: †Edmontonia
and tank-like. Its length has been Sternberg , 1928
estimated at about 6.6 m
(22 ft).[1] In 2010, Gregory S.
Species

Size comparison Paul considered both main

Edmontonia species, E.
E. longiceps Sternberg, 1928
longiceps and E. rugosidens, to (type)
[2]
be equally long at six metres and weigh three tonnes. E. rugosidens (Gilmore, 1930
Edmontonia had small, oval ridged bony plates on its back and head and many [originally Palaeoscincus
sharp spikes along its sides. The four largest spikes jutted out from the shoulders rugosidens ])
on each side, the second of which was split into subspines in E. rugosidens
Synonyms
specimens. Its skull had a pear-like shape when viewed from above.[1] Its neck
and shoulders were protected by three halfrings made of lar
ge keeled plates.
Chassternbergia Bakker, 1988

Distinguishing traits
In 1990, Kenneth Carpenter established some diagnostic traits for the genus as a whole, mainly comparing it with its close relative
Panoplosaurus. In top view, the snout has more parallel sides. The skull armour has a smooth surface. In the palate, the vomer is
keeled. The neural arches and neural spines are shorter than those of Panoplosaurus. The sacrum proper consists of three sacral
 vertebrae. In the shoulder girdle, the scapula andcoracoid are not fused.[3]

Carpenter also indicated in which way the main species differed from each other.
The type species, Edmontonia longiceps, is distinguished from E. rugosidens in
lacking sideways projecting osteoderms behind the eye sockets; having tooth rows
that are less divergent; possessing a more narrow palate; having a sacrum that is
wider than long and more robust; and in having shorter spikes at the sides. Also, an
Restoration of E. rugosidens
ossified cheek plate, known from E. rugosidens specimens, has not been found with
Edmontonia longiceps.[3]

Skeleton
The skull of Edmontonia, up to half a metre long, is somewhat elongated with a
protruding truncated snout. The snout carried a horny upper beak and the front snout
bones, the premaxillae, were toothless. The cutting edge of the upper beak continued
into the maxillary tooth rows, each containing fourteen to seventeen small teeth. In
each dentary of the lower jaws, eighteen to twenty-one teeth were present. In the
sides of the snout large depressions were present, "nasal vestibules", that each
possessed two smaller openings. The top of these was a horizontal oval and
represented the bony external nostril, the entrance to the nasal cavity, the normal air
passage. The more rounded second opening below and obliquely in front, was the
entrance to a "paranasal" tract, running along the outer side of the nasal cavity, in a
Restored E. rugosidens skeleton somewhat lower position. A study by Matthew Vickaryous in 2006 proved for the
without back armour first time the presence of multiple openings in a nodosaurid; such structures had
already been well established in ankylosaurids. The air tracts are however, much
simpler than in the typical ankylosaurid condition, and are not convoluted while
lacking bony turbinate bones. The nasal cavity is separated into two halves along the midline by a bone wall. This septum is
continued to below by the vomers, which are keeled, the keel featuring a pendulum-shaped appendage.[4] Another similarity with
Ankylosauridae is the presence of a secondary bone palate, a possible case of parallel evolution. This has been shown too for
Panoplosaurus.[3]

The head armour tiles, or caputegulae, are smooth. Details differ between the
various specimens but all share a large central nasal tile on the snout, bend large
"loreal" tiles at the rear snout edges and a large central caputegula on the skull roof.
The tiles behind the upper eye socket rim in Edmontonia longiceps do not stick out
as much as in E. rugosidens, combined with a more narrow, pointed snout in the
former. Some E. rugosidens specimens are known that possess a "cheek plate" above
the lower jaw. Contrary to that discovered with Panoplosaurus, it is "free-floating",
not fused with the lower jaw bone.[5]

The vertebral column contains about eight neck vertebrae, about twelve "free" back
The AMNH 5381 specimen ofE.
vertebrae, a "sacral rod" of four fused rear dorsal vertebrae, three sacral vertebrae,
rugosidens, 1915 (first referred to
two caudosacrals and at least twenty, but probably about forty, tail vertebrae. In the
Palaeoscincus by Matthew in 1922),
neck the first two vertebrae, the atlas and axis, are fused. In the shoulder girdle, the showing the position of the dermal
coracoid has a rectangular profile, in contrast to the more rounded shape with armour
Panoplosaurus. Two sternal plates are present, connected to sternal ribs. The
forelimb is robust but relatively long. In Edmontonia longiceps and E. rugosidens
the deltopectoral crest of the humerus is gradually rounded. The metacarpus is robust compared to that of Panoplosaurus. The hand
very likely was tetradactyl, having four fingers.[3] The exact number of phalanges is unknown but the formula was by W.P. Coombs
suggested to be 2-3-3-4-?.[6]
Osteoderms
Apart from the head armour, the body was covered with osteoderms, skin
ossifications. The configuration of the armour of Edmontonia is relatively well
known, much of it having been discovered in articulation. The neck and shoulder
region was protected by three cervical halfrings, each consisting of fused rounded
rectangular, asymmetrically keeled, bone plates. These halfrings did not have a
continuous underlying bone band. The first and second halfrings each had three pairs
of segments. Below each lower end of the second halfring a side spike was present, a
separate triangular osteoderm pointing obliquely forward. In the third halfring over
Skull and neck armor
the shoulders, the two pairs of central segments are bordered on each side by a very
large forward-pointing spike that is bifurcated, featuring a secondary point above the
main one. A third large spike behind it points more sideways; a smaller fourth one, often connected to the third at the base, is directed
obliquely to behind. The row of side spikes is continued to the rear but there the osteoderms are much lower, curving strongly to
behind, with the point overhanging the rear edge. Gilmore had trouble believing that the shoulder spikes really pointed to the front as
this would have greatly hampered the animal while moving through vegetation. He suggested that the points had shifted during the
burial of the carcass. However, Carpenter and G.S. Paul, trying to reposition the spikes, found that it was impossible to rotate them
without losing conformity with the remainder of the armour. The side spikes have solid, not hollow, bases. The spikes differ in size
between E. rugosidens individuals; those of theE. longiceps holotype are relatively small.[3]

Behind the third halfring the back and hip are covered by numerous transverse rows of much smaller oval keeled osteoderms. These
are not ordered in longitudinal rows. The front rows have plates oriented along the length of the body, but to the rear the long axis of
these osteoderms gradually rotates sideways, their keels ultimately running transversely. Rosettes are lacking. The configuration of
the tail armour is unknown. The larger plates of all body parts were connected by small ossicles.[3] Such small round scutes also
covered the throat.[4]

Discovery and species

In 1915, the American Museum of Natural History obtained the nearly complete,
articulated front half of an armoured dinosaur, found the same year by Barnum
Brown in Alberta, Canada. In 1922, William Diller Matthew referred this specimen,
AMNH 5381, to Palaeoscincus in a popular-science article, not indicating any
particular species.[7] It had been intended to name a new Palaeoscincus species in
cooperation with Brown but their article was never published.[3] Matthew also
referred specimen AMNH 5665, the front of a skeleton found by Levi Sternberg in
1917. In 1930 Charles Whitney Gilmore referred both specimens to Palaeoscincus

Life restoration of two E. rugosidens rugosidens.[8] This species was based on type specimen USNM 11868, a skeleton
from 1922, based on the 1915 AMNH found by George Fryer Sternberg in June 1928. The specific name is derived from
specimen Latin rugosus, "rough", and dens, "tooth". In 1940, Loris Shano Russell referred all
three specimens to Edmontonia, as an Edmontonia rugosidens.[9]

Meanwhile, the type species of Edmontonia, Edmontonia longiceps, had been named by Charles Mortram Sternberg in 1928. The
generic name Edmontonia refers to Edmonton or the Edmonton Formation. The specific name longiceps means "long-headed" in
Latin. Its holotype is specimen NMC 8531, consisting of a skull, right lower jaw and much of the postcranial skeleton, including the
armour. It was discovered nearMorrin in 1924 by George Paterson, the teamster of the expedition led by C.M. Sternberg.[10]

Edmontonia species include:

E. longiceps, the type, known from a complete skull, is known from the middleHorseshoe Canyon Formation(Unit 2)
which used to be dated to 71.5-71 million years ago.[11] This unit, which straddles theCampanian-Maastrichtian
boundary, has since been recalibrated to an age of about 72 million years. Isolated bones and shed teeth from E.
longiceps are also known from the upper Judith River Formation in Montana.
 E. rugosidens. This species has been given its own genus,
Chassternbergia, first coined as a subgenus by Dr. Robert Thomas
Bakker in 1988, as Edmontonia (Chassternbergia) rugosidensand is
based on differences in skull proportion fromE. longiceps and its earlier
time period.[5][12] It was given its full generic name in 1991 byGeorge
Olshevsky.[13] The name Chassternbergia honours Charles, "Chas", M.
Sternberg. This subgenus or genus name is rarely applied. [14][15] E.
rugosidens is found in the Campanian lower Dinosaur Park Formation,
dating from about 76.5-75 million years ago.[11] Many later finds have
been referred to E. rugosidens, among them CMN 8879, the top of a
skull found in 1937 by Harold D'acre Robinson Lowe; ROM 433, a Left side of E. rugosidens specimen
forked spine found by Jack Horner in 1986 among Oohkotokia material;
AMNH 5665
ROM 5340, paired medial plates; ROM 1215, a skeleton; R TMP
91.36.507, a skull; RTMP 98.74.1, a possibleEdmontonia skull; RTMP
98.71.1, a skeleton;[16] RTMP 98.98.01, a skull and right lower jaw;and
RTMP 2001.12.158, a skull.[4]
Edmontonia schlessmani was a renaming in 1992 of Denversaurus schlessmani("Schlessman's Denver lizard") by Adrian Hunt and
Spencer Lucas.[17] This taxon was erected by Bakker in 1988 for a skull from the Late Maastrichtian Upper Cretaceous Lance
Formation of South Dakota, specimen DMNH 468 found byPhilip Reinheimer in 1922. Thistype specimen of Denversaurus is in the
collections of the Denver Museum of Natural History (now the Denver Museum of Nature and Science), Denver, Colorado for which
the genus was named. The specific name honours Lee E. Schlessman, whose Schlessman Family Foundation sponsored the museum.
Bakker described the skull as being much wider at the rear than Edmontonia specimens.[5] However, later workers explained this by
its being crushed,[3] and considered the taxon a junior synonym of Edmontonia longiceps.[14] The Black Hills Institute has referred a
skeleton from the Lance Formation to Denversaurus, nicknamed "Tank". It has the inventory number BHI 127327.[18] New research
indicates that it is closely related toPanoplosaurus.[19]

Edmontonia australis was named by Tracy Lee Ford in 2000 on the basis of cervical scutes, the holotype NMMNH P-25063, a pair
of medial keeled neck osteoderms from the Maastrichtian Kirtland Formation of New Mexico and the paratype NMMNH P-27450, a
right middle neck plate.[12] Although later considered to a dubious name,[15] it is now considered a junior synonym of
Glyptodontopelta mimus.[20]

The naming history was further complicated in 1971, when Walter Preston Coombs Jr renamed both Edmontonia species, into
Panoplosaurus longiceps and Panoplosaurus rugosidens respectively.[21] The latter species, which due to its much more complete
material has determined the image of Edmontonia, until 1940 thus appeared under the name of Palaeoscincus, and during the 1970s
and 1980s was shown as "Panoplosaurus" until newer research revived the name
Edmontonia.

In 2010, G.S. Paul suggested that E. rugosidens was the direct ancestor of Edmontonia longiceps and the latter was again the direct
ancestor of E. schlessmani.[2]

Phylogeny
C.M. Sternberg originally did not provide a classification of Edmontonia. In 1930, L.S. Russell placed the genus in the Nodosauridae,
which has been confirmed by subsequent analyses. Edmontonia was generally shown to be a derived nodosaurid, closely related to
Panoplosaurus. Russell in 1940 named a separate Edmontoniinae. In 1988 Bakker proposed that the Edmontoniinae with the
Panoplosaurinae should be joined into Edmontoniidae, the presumed sister group of the Nodosauridae within Nodosauroidea which
he assumed not be ankylosaurians but the last surviving stegosaurians.[5] Exact cladistic analysis has not confirmed these hypotheses
however, and the concepts of Edmontoniinae and Edmontoniidae are not in modern use.

The following cladogram shows the position of Edmontonia in the nodosaurid evolutionary tree according to an analysis in 2011 by
[22]
paleontologists Richard Stephen Thompson, Jolyon C. Parish, Susannah C. R. Maidment and Paul M. Barrett.

Nodosauridae
Antarctopelta

Mymoorapelta
 Hylaeosaurus

Anoplosaurus

Tatankacephalus

Horshamosaurus

Gargoyleosaurus

Hoplitosaurus

Polacanthinae Gastonia

Peloroplites

Polacanthus

Struthiosaurus

Zhejiangosaurus

Hungarosaurus

Animantarx

Niobrarasaurus

Nodosaurus

Pawpawsaurus

Sauropelta

Silvisaurus

Stegopelta

Texasetes

Edmontonia

Panoplosaurus

Paleobiology

Function of the armour

The large spikes were probably used between males in contests of strength to defend territory or gain mates.[1] The spikes would also
have been useful for intimidating predators or rival males, passive protection, or for active self-defense.[1] The large forward pointing
shoulder spikes could have been used to run through attacking theropods.[2] Carpenter suggested that the larger spikes of AMNH
5665 indicated this was a male specimen, a case of sexual dimorphism. However, he
admitted the possibility of ontogeny, older individuals having longer spikes, as the
specimen was relatively large.[3] Traditionally it had been assumed that to protect
themselves from predators, nodosaurids likeEdmontonia might have crouched down
on the ground to minimize the possibility of attack to their defenseless underbelly,
.[2]
trying to prevent being flipped over by a predator

Paleoecology Close up of the shoulder spikes of

AMNH 5665
Rings in the petrified wood of trees contemporary with Edmontonia show evidence
of strong seasonal changes in precipitation and temperature;[1] this may hold an
explanation for why so many specimens have been found with their armor plating and spikes in the same position they were in life.[1]
The Edmontonia could have died due to drought, dried up, and then rapidly became covered in sediment when the rainy season
began.[1]

Edmontonia rugosidens existed in the upper section of the Dinosaur Park Formation, about 76.5–75 million years ago. It lived
alongside numerous other giant herbivores, such as the hadrosaurids Gryposaurus, Corythosaurus and Parasaurolophus, the
ceratopsids Centrosaurus and Chasmosaurus, and ankylosaurids Scolosaurus[11] and Dyoplosaurus[11] Studies of the jaw anatomy
and mechanics of these dinosaurs suggests they probably all occupied slightly different ecological niches in order to avoid direct
competition for food in such a crowded eco-space.[23] The only large predators known from the same levels of the formation as
Edmontonia are the tyrannosauridsGorgosaurus libratus and an unnamed species ofDaspletosaurus.[11]

Edmontonia longiceps is known from the Horseshoe Canyon Formation, from the middle unit, which was dated to 71.5-71 million
years ago in 2009.[11] The fauna of the Horseshoe Canyon Formation is well-known, as vertebrate fossils, including those of
dinosaurs, are quite common. Sharks, rays, sturgeons, bowfins, gars and the gar-like Aspidorhynchus made up the fish fauna. The
saltwater plesiosaur Leurospondylus has been found in marine sediments in the Horseshoe Canyon, while freshwater environments
were populated by turtles, Champsosaurus, and crocodilians like Leidyosuchus and Stangerochampsa. Dinosaurs dominate the fauna,
especially hadrosaurs, which make up half of all dinosaurs known, including the genera Edmontosaurus, Saurolophus and
Hypacrosaurus. Ceratopsians and ornithomimids were also very common, together making up another third of the known fauna.
Along with much rarer ankylosaurians and pachycephalosaurs, all of these animals would have been prey for a diverse array of
carnivorous theropods, including troodontids, dromaeosaurids, and caenagnathids.[24][25] Adult Albertosaurus was the apex predator
[24]
in this environment, with intermediate niches possibly filled by juvenile albertosaurs.

See also
Timeline of ankylosaur research

References
1. "Edmontonia." In: Dodson, Peter & Britt, Brooks & Carpenter, Kenneth & Forster, Catherine A. & Gillette, David D. &
Norell, Mark A. & Olshevsky, George & Parrish, J. Michael & Weishampel, David B. The Age of Dinosaurs.
Publications International, LTD. p. 141. ISBN 0-7853-0443-6.
2. Paul, G.S., 2010, The Princeton Field Guide to Dinosaurs, Princeton University Press p. 238
3. Carpenter, K. 1990. "Ankylosaur systematics: example using Panoplosaurus and Edmontonia (Ankylosauria:
Nodosauridae)", In: Carpenter, K. & Currie, P.J. (eds) Dinosaur Systematics: Approaches and Perspectives
,
Cambridge University Press, Cambridge, pp. 281-298
4. Vickaryous, Matthew K. (2006). "New information on the cranial anatomy ofEdmontonia rugosidensGilmore, a Late
Cretaceous nodosaurid dinosaur from Dinosaur Provincial Park, Alberta".Journal of Vertebrate Paleontology. 26 (4):
1011–1013. doi:10.1671/0272-4634(2006)26[1011:niotca]2.0.co;2(https://doi.org/10.1671%2F0272-4634%28200
6%2926%5B1011%3Aniotca%5D2.0.co%3B2) .
 5. Bakker, R.T. (1988). Review of the Late Cretaceous nodosauroid Dinosauria:Denversaurus schlessmani, a new
armor-plated dinosaur from the Latest Cretaceous of South Dakota, the last survivor of the nodosaurians, with
comments on Stegosaur-Nodosaur relationships.Hunteria 1(3):1-23.(1988).
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Dinosauria. University of California Press. pp. 456–483.
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9. Russell, L.S. (1940). "Edmontonia rugosidens (Gilmore), an armored dinosaur from the Belly River Series of
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24. Eberth, D.A., 1997, "Edmonton group". In: Currie, .PJ., Padian, K. (Eds.), Encyclopedia of Dinosaurs. Academic
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