K. Carpenter (ed.) 2001. The Armored Dinosaurs.

" Indiana
University Press. Bloomino-ton
b , IN
_._-_ .. _._-

9. Skull of the Polacanthid

Ankylosaur Hylaeosaurus
armatus Mantell, 1833,
from the Lower
Cretaceous of England
KE~'NETH CARPENTER

Abstract
The holotype of the polacanthid ankylosaur Hylaeosaurus armatus
includes a partial, highly damaged skull that has never been described
in detail. It consists of the rear portion, which is exposed ventrally, and
includes the left quadrate articulated to its paroccipital process, part of
the left quadratojugal, righf~postorbital horn, and possibly the rear
portions of both lower jaws. The postorbital horn is triangular and
small and is comparable with that of the polacanthid Gargoyleosaurus.
The quadrate has a prominent rim along its anterior lateral edge, a
feature also seen in the primitive nodosaurid Animantarx.

Introduction
The armored dinosaur Hylaeosaurus armatus was one of the earli-
est named dinosaurs. It was named by Gideon Mantell in 1833, only
eight years after the naming of Iguanodon and nine years after the
naming of Megalosaurus. The holotype, BMNH R3775, consisted of
the anterior portion of an articulated skeleton and was briefly described
by Mantell (1833) and in more detail by Owen (1857) and Pereda-
Suberbiola (1993). To date, however, the fragmentary skull associated
with the specimen has not been described.

169

Figure 9.1. Photograph of the
skull of Hylaeosaurus in close-up
(left), and an outline drawing of
the skull with parts labeled
(right). The postorbital horn is
not seen very well in the
photograph because of parallax.
170 • Kenneth Carpenter
axis
articular?
postorbital horn
Institutional Abbreviation. BMNH: Natural History Museum (for-
merly British Museum [Natural History]), London.
Description
Only the rear portion of the skull remains and is seen in ventral
view (Fig. 9.1). The fragment is 18.5 em wide and 12.2 em long. The
surface of some of the bones is damaged, making it difficult to delineate
their margins, especially because the matrix and bones are presently
nearly the same color. The axis and left quadrate provide key anatomi-
cal reference points to aid in the identification of the surrounding ele-
ments.
The left quadrate is visible in lateral view, with the exposed portion
10.7 em long. The quadrate is partially twisted so that a portion of the
posterior side faces laterally. This surface is damaged, but it can be seen
to articulate with the paroccipital process. No suture is seen separating
the quadrate head from the paroccipital process, implying that the two
elements are coossified. The distal, or condylar, end is overlain by ex-
tensively damaged bone or bones that may be the rear portion of the
right lower jaw. The quadrate is arched laterally, as it is in the pol a-
canthid Gargoyleosaurus (see Carpenter 2001 for a definition of the
Polacanthidae). A rim along the lateral edge of the quadrate, near the
quadratojugal, makes the shaft appear to have a groove along its back
side. The extent of this rim, however, cannot be determined because of
damage. This feature is seen in Animantarx ramaljonesi (Carpenter et
al. 1999) and Stegosaurus, and it probably marks the anterior edge of
the tympanic membrane. The quadrate shaft is 2.8 em in lateromedial
width, where it is least damaged near the quadratojugal.
 The quadratojugal is identified on the basis of its anatomical posi-
tion relative to the quadrate. It is short, measuring 1.4 cm long and 2.4
cm deep. The quadratojugal is small in most ankylosaurs and braces the
quadrate anterolaterally with the jugal. The anterior edge is damaged,
but only a little is probably missing on the basis of the proportions of
Gargoyleosaurus (personal observation). The quadratojugal is coossi-
fied higher on the quadrate shaft than it is in the nodosaurid Edmon-
tonia, were it is located near the distal condyle. The high position is
characteristic of the polacanthid Gargoyleosaurus and the primitive
nodosaurids Pawpawsaurus and Animantarx, but not the advanced
nodosaurid Edmontonia.
The only part of the paroccipital process that is visible is its distal
end, where it articulates with the quadrate head. It is at least 2.8 cm
deep. The quadrate head is damaged, but no suture is visible, suggesting
that the head was fused, at least in part, to the paroccipital process. The
medial part of the paroccipital process is covered by damaged bone or
bones that may be part of the left lower jaw. The damage is so severe,
however, that this identification is questionable. This damaged bone is
in contact with another bone anteriorly and is separated from it poste-
riorly by the axis. This bone is also severely damaged, but it may be the
rear portion of the right mandible on the basis of what appears to be the
medial process of the articular. If this object is the right mandible, it has
shifted back almost 10 cm from the estimated position of the distal end
of the quadrate.
Anterior to the paroccipital process is a somewhat bulbous bone
protruding posterior to the quadrate and lateral to the questionable left
mandible. Little detail can be seen, but on the basis of its topographic
position, is probably the left basitubera. If this identification is correct,
it is large for an ankylosaur.
The right postorbital horn is seen in lateral view. The surface is
damaged, but it still retains its triangular shape. It is 5.5 cm wide at its
base and projects laterally about 3.2 cm. The horn is proportional to
that seen in the polacanthids Gargoyleosaurus and Gastonia. There are
two small depressions on the surface near where the horn joins the rest
of the skull. Unfortunately, because the surrounding bone is damaged,
it is not certain if these depressions are pathologic or not.

Discussion
The skull to the holotype of Hylaeosaurus armatus is incomplete
and much of the surface is damaged, making comparisons with other,
more complete ankylosaur skulls difficult. The identifications offered
here, with the exception of the quadrate, are tentative and may be
changed if the skull fragment is prepared free of the block containing
the skeleton.
The similarities between the skulls of Gargoyleosaurus and Hylae-
osaurus enumerated above indicate a close affinity between the two
taxa and support the inclusion of Hylaeosaurus into the Polacanthidae
(see Carpenter 2001).
Acknowledgments. My thanks to Sandra Chapman and Angela

Skull of Hylaeosaurus armatus Mantell • 171

 Milner (Natural History Museum, London) for access to the anky-
losaur specimens in their care. Thanks also to the review comments by
Xabier Pereda Suberbiola.

References Cited
Carpenter, K. 2001. Phylogenetic analysis of the Ankylosauria. In K. Car-
penter (ed.), The Armored Dinosaurs. Bloomington: Indiana Univer-
sity Press [this volume, Chapter 21].
Carpenter, K.,]. Kirkland, D. Burge, and]. Bird. 1999. Ankylosaurs (Dino-
sauria: Ornithischia) of the Cedar Mountain Formation, Utah, and
their stratigraphic distribution. In D. Gillette (ed.), Vertebrate Paleon-
tology in Utah. Utah Geological Survey Miscellaneous Publication
99-1: 244-251.
Mantell, G. 1833. The Geology of the South-East of England. London:
Longman.
Owen, R. 1857. Monograph of the fossil Reptilia of the Wealden and
Purbeck Formations. Part IV. Hylaeosaurus. Palaeontological Society
of London Monograph 10: 8-26.
Pereda-Suberbiola, J. 1993. Hylaeosaurus, Polacanthus, and the systemat-
ics and stratigraphy of Wealden armored dinosaurs. Geological Maga-
zine 130: 767-781.

172 • Kenneth Carpenter