Introduction
Although incompletely prepared, this specimen provides an
excellent view of the relationships between the bones of the skull and jaws
when they are in occlusion. It is worth noting that the caudal region of the
skull – including the braincase and suspensorium – has been displaced
rostrally. For example, this has shifted the basisphenoid unnaturally ahead
relative to the pterygoids, and the basioccipital ahead of the otoccipitals.
In this specimen, the relationship between the skull and
closed jaws can clearly be seen. For example, the region of the greatest depth
of the jaws (where the prearticular and splenial meet) is positioned below the
medial edge of the midlength of the palatine. It is seen that the palatal
process of the maxilla extends medial to the mandible, which is consistent with
the position of depressions that received the teeth along the medial alveolar
process of the maxilla. Also, the dorsal edge of the mandible extended within a
wide slot formed by the jugal laterally and the ectopterygoid medially along
its course to the jaw joint. Finally, the slip of bone lateral to the
surangular and angular is the ventral edge of the jugal.
Description
SKULL BONES
Basioccipital: In
ventral view, the basioccipital is seen situated below the otoccipital and
caudal to the basisphenoid. It forms the caudoventral region of the occipital
condyle, the ventral part of the neck of the occipital condyle, the caudoventral
region of the basicranium, the caudoventral edge of the basicranium, and the
midregion of the caudal wall of the basisphenoid recess. The basioccipital is
positioned between the otoccipital and basisphenoid, except for the caudodorsal
region that culminates in the occipital condyle. The basioccipital does not
contribute to the caudoventrolateral corner of the basicranium; instead, the
otoccipital and basisphenoid form the corner where they contact each other
distal to the basioccipital. The external surface of the basioccipital occupies
four planes: caudoventral at the occipital condyle, caudoventral below the
occipital condyle, ventral between the basal tubera, and rostral at the basisphenoid
recess.
In ventral view, the basal tuber can be seen, and the
prominent ascending scar blocks the region of the subcondylar fossa from view. The
extensive muscle attachment surface below the occipital condyle dominates the
basioccipital. It can also be seen that the funnel-like ceiling of the
basisphenoid recess produces a concave surface in the rostral surface of the
bone.
Basisphenoid: In
ventral view, the basisphenoid is broadly exposed to view, especially the
basisphenoid recess and its contributions to the basal tuber complex. The bone
contacts the basioccipital caudally, the otoccipital caudodorsolaterally, and
the pterygoid rostroventrolaterally. With the otoccipital the bone forms the
caudoventrolateral corner of the basicranium. Although somewhat difficult to
determine from this image, the basisphenoid extends steeply rostroventrally
from the basal tuber complex to the basipterygoid process. As noted, the caudal
end of this skull is displaced rostrally; in its natural position, the basipterygoid
process would be situated above – not rostral to – the stout, caudally-
extending process from the pterygoid.
Ectopterygoid: Qlthough the ectopterygoid is often pictured
lying flat, in
ventral view its true orientation is seen, where it extends steeply rostrodorsally. This position accounts for its appearance in lateral view below
the jugal as a prominent hook-like process. Although concealed by matrix, the distal end of the thin
and rostrodorsally extending jugal process is apposed to the medial surface of the
jugal at the caudal end of the ventral cornual process of that bone. The caudomedial and caudal edges of the ectopterygoid have an extensive
overlapping contact with the pterygoid. The bone is in
apposition medially with a stout lateral process of the pterygoid.
Maxilla: In this
view, only the rostral part of the maxilla is in view, where it extends lateral to the
dentary; tts contacts with other facial bones cannot be seen. With the dorsal
processes of the palatines and the vomer concealed by matrix, it can be seen that the maxilla can be considered as a part of the palate. The palatal process (=shelf) of the
maxilla is completed caudomedially by the palatine, which continues this flat and
ventromedially facing surface caudally to the pterygoid.
Otoccipital: In
ventral view, the otoccipital is tilted rostroventrally at a steep
angle. The bone articulates ventrally with the basioccipital for most of
its width medially, and with a slip of the basisphenoid laterally. The
otoccipital forms the dorsolateral part of the neck of the occipital condyle,
and that of the condyle itself. In this specimen, the basioccipital is
displaced the rostrally relative to the otoccipital, exposing the joint surface
for the basioccipital to view, especially on the right side. The otoccipital
extends caudodorsolaterally, until the articular bone of the lower jaw conceals its lateral end from view. As the otoccipital extends laterally, it becomes
progressively vertically (caudally) oriented such that it appears to taper along
its course. In contrast, the ventromedial corner of the bone does taper to a
point as it extends toward the basal tuber.
Palatine: In
ventral view, most of the pterygoid process and the rostral end of its maxillary
process of the palatine can be seen although the bone is largely concealed by the mandibular
ramus and matrix. The palatine contacts the pterygoid caudomedially and it
extends along the maxilla rostromedially. The pterygoid process is wide,
rodlike, and it extends caudomedially, where it is underlapped by the pterygoid.
The palatine sends a small slip along the rostrolateral edge of the pterygoid; otherwise it is completely covered by that bone. The pterygoid is positioned
distally on the process; i.e., the pterygoid does not reach the intersection
between the three primary processes (vomeropterygoid, maxillary, pterygoid) of the palatine.
In contrast, the maxillay process in this view is a narrow
and rostromedially tapering slip that extends along the medial edge of the
palatal process of the maxilla. Therefore, the palatine contributes to the
caudomedial edge of the palatal shelf.
On the midline, the complementary pterygoid and maxillary
processes surround a relatively wide space that separates the two bones. This
stands out here because the dorsally extending vomeropterygoid processes of the bone are not prepared to
view. If they were, it would be seen that they would have extended toward each
other and made contact dorsally, with their rostral margins defining the caudal
edge of the bony choana.
Pterygoid: In
ventral view, the pterygoid is a complex bone that divides from a common point
into four processes, including the quadrate process that extends
caudodorsolaterally, the basisphenoid process that extends caudolaterally, the
ectopterygoid process that extends laterally, and the main body of the bone
that extends rostrally.
The complementary bodies of the bone are separated by a long
interpterygoid vacuity, which extends rostrally between the palatines and
maxillae. The body of the bone is a long structure that extends ahead of the
lateral process to underlap the caudoventral surface of the pterygoid process
of the palatine.
The lateral process underlaps the caudoventral surface of
the ectopterygoid, and twists along its long axis into a steep caudoventral to
rostrodorsal orientation. The ventral surface between the lateral process and the
body is deeply concave; it is possible that this fossa is pneumatic and
associated with the large ectopterygoid pneumatic recess.
The basisphenoid process is the shortest of the processes,
which extends slightly caudolaterally to underlap the basipterygoid process of
the basisphenoid at a very loose basal joint. The lateral process extends
ventrolaterally relative to this process. In contrast, the quadrate process
extends steeply rostrodorsolaterally relative to the short process. In this
view, only the ventral edge of the dorsal process can be seen, extending along
the ventral surface of the orbital process of the quadrate.
Quadrate: In
ventral view, the quadrate is a narrow rostromedially extending strut that
overlaps the tip of the dorsal process of the quadrate. In part, this is an
artifact of the sediment-filled spaces in the skull.
MANDIBULAR BONES
Angular: In
ventral view, the angular is a dominant bone of the caudal half of the
ventrolateral and ventral surfaces of the mandibular ramus. It contacts the
surangular caudomedially, the dentary rostrolaterally, the prearticular
dorsally and medially, and the splenial rostromedially.
Caudally it overlaps the lateral surface of the surangular,
which separates the angular from the ventral margin of the bone in this region;
i.e., the caudal tip of the angular extends caudodorsally onto the lateral
surface of the surangular. Rostrally the angular extends along the prearticular
before it continues medially onto the ventral surface of that bone, where it
forms the ventral margin of the mandibular ramus. The angular then extends
rostrodorsally out of view between the dentary and splenial. The angular curves laterally along
its course.
Articular: The
articular forms the caudomedial end of the mandibular ramus, and it is the
widest part of the structure. In ventral view, the articular contacts the
prearticular rostromedially and the surangular rostrolaterally. The suture with
the prearticular is not easily seen in this specimen, so the placement and
course shown here is approximate. The articular is relatively widely exposed in
ventral view and it extends the mandibular ramus medially, where it constricts
the caudal region of the nasopharynx down to the width of the snout. Together,
the articular and prearticular form a prominent medial bar that extends below
the Meckelian fossa.
Dentary: In
ventral view, the dentary is the longest bone of the mandibular ramus, where it
contacts the angular caudomedially and the splenial medially. The dentary
follows the curvature of the snout, where its caudal two thirds extends
caudolaterally and its rostral third extends nearly directly rostrally to the
symphysis. The change in direction
occurs far rostrally, approximately at the midlength of the maxillary tooth
row. The row of neurovascular foramina that penetrates the ventrolateral
surface of the bone is in view along the rostral half of the bone; the nerve branches that extended from these openings to innervate the skin would have sent tactile information to the brain.
The dentary is widest along the rostral third of the bone,
which corresponds to the dentigerous region of the bone. This approximately
matches the region of the largest maxillary teeth; the rostral end of the bone
attenuates at a blunt tip that is flattened medially by the symphysis. The
caudal half of the bone is much narrower than the part below the teeth; this region does not
continuously reduce in width, but it is abruptly attenuated where it extends
onto the lateral surface of the angular.
The rostral end of the dentary abuts the lingual surface of
the premaxillary teeth, indicating an overbite when the mouth is closed. It is
also seen that the maxillary teeth are positioned lateral to the dentary,
indicating that contact between the dentary and maxillary teeth does not occur
in tyrannosaurids; likewise, there is no occlusion between the dentary and
premaxillary teeth in tyrannosaurids. The medial edge of the dentary is
continuous except caudally, where it deviates caudolaterally at the angular.
Intercoronoid:
Nearly the entire caudal extent of the intercoronoid is seen in ventral view at
the rostrodorsal limit of the Meckelian fossa. Put another way, the Meckelian
fossa separates the caudoventral edge of the intercoronoid from the caudodorsal edge
of the prearticular. This small part of the bone is overlapped rostromedially
by the prearticular and it is apposed to the medial surface of the surangular.
This part of the intercoronoid is triangular in shape, flat, and faces
ventromedially, in plane with the rest of the mandibular bones in the midregion
of the skull. When the jaws are closed, the caudal end of the intercoronoid is
positioned lateral to the ectopterygoid, below its jugal process and above its
caudolateral process.
The rostral part of the intercoronoid can be seen above the
splenial; the suture between them was difficult to trace rostrally, so in the
illustration a dashed line marks its approximate course. The rostral part of
the bone completes the upper part of the medial wall of the dental alveoli (tooth
sockets) in the form of a long strap that is dorsoventrally shallow and
mediolaterally thin. The ventral edge of the intercoronoid extends along the
rostrodorsal edge of the splenial; its rostral end is blocked from view by the
dentary as it twists along its long axis into a vertical orientation.
Prearticular: In
ventral view, the prearticular tilts medioventrally, showing it is the longest
bone of the caudal region of the mandibular ramus, which extends for half of
its length. The prearticular contacts the articular caudally, the surangular
caudolaterally, the angular laterally and ventrally, the splenial
rostroventrally, and the intercoronoid caudodorsally. The prearticular forms the ventrla margin of the ramus along its course between the surangular and the region where it is undrlapped by the angular.
The prearticulosplenial
contact is separated for a short distance rostrodorsally by the internal
mandibular fenestra. The prearticular is mediolaterally wide caudally, whereas
it is thin rostrally. With the jaws closed, the rostrodorsal end of the bone is
positioned below the base of the pterygoid process of the palatine.
Caudally the prearticular overlaps the rostroventromedial
surface of the articular, whereas it is overlapped caudoventrolaterally by the
surangular, and ventrally by the angular. The rostroventral edge of the bone
extends along the edge of the splenial. Rostrodorsally the bone tapers toward
the dorsal tip of the splenial.
Splenial: In
ventral view, the medial surface of the splenial tilts mediodorsally into view,
showing it is the longest mandibular bone in the midregion of the ramus. The
splenial contacts the prearticular caudodorsally, the angular caudoventrally,
the dentary ventrally, and the intercoronoid rostrodorsally. The caudodorsal
margin of the bone is deeply notched by the internal mandibular fenestra, which
separates it from the prearticular. At no point does the splenial does not
reach the ventral margin of the mandibular ramus. The medial surface of the
bone is essentially flat, except caudally where a ridge can be seen.
The splenial starts below the rostral end of the Meckelian
fossa between the prearticular and the angular, and it extends rostrally where
it stops below the midlength of the dentigerous region of the snout. The bone
tapers rostrally below the intercoronoid and above the dentary. It is
perforated rostrally by the large rostral mylohyoid foramen. With the mouth
closed, the foramen is positioned below the palatal process of the maxilla,
ahead of the palatine bone. The dorsal apex of the splenial is positioned between
the pterygoid and maxillary processes of the palatine, ahead of the
prearticular.
Surangular: In
ventral view, only the caudoventral corner of the surangular can be seen. The
bone contacts the articular caudoventromedially, the prearticular
ventromedially, and the angular ventrolaterally. The lateral surface of the
surangular is deeply creased at the caudal end of the surangular shelf. Despite
its small size, the surangular forms the caudoventral corner of the mandibular
ramus, and virtually prevents the articular from lateral exposure. A small tab
from the surangular extends medially to underlap the region where the articular
and prearticular join each other; in this region the surangular manages to form
the ventral surface of the ramus.
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