J . Zool., Lond.

( A ) (1985) 207,125-136

The mammalian postorbital bar as a torsion-resisting helical strut

W. S. GREAVES
Departments of Oral Anatomy and Anatomy, University of Illinois at Chicago,
801 South Paulina Street, Chicago, Illinois 60680 U.S.A.

(Accepted I 1 December 1984)

(With 4 figures in the text)

The mammalian skull is asymmetrically loaded during mastication because most of these animals
chew on only one side at a time. This loading regime tends to twist the braincase relative to the
rostral, tooth bearing part of the skull at the zone of potential weakness between the orbits. This
torsional effect is exaggerated, and a postorbital bar is present, in those animals with very large
masseter and pterygoid muscles. The lines of action of these muscles are oriented at large angles to
the long axis of the skull in lateral view, providing large components of force that twist the skull
segments relative to one another. When the temporalis is the dominant muscle, the torsional effect
is usually less important, and the bar is absent, because this muscle acts at a smaller angle to the
skull axis. The postorbital bar exhibits the predicted three dimensional spatial orientation
required to resist torsional forces: it is a segment of an imaginary 45" helix that is wound around
the skull, if the skull is idealized as a cylinder. This orientation i s significant because, in general,
maximum compressive and tensile shear stresses lie along 45" helices on a cylinder loaded in
torsion; to resist torsion, material should be placed far from the axis of torsion and along a helix
oriented at 45" to the deforming forces. Each half of a supraorbital ridge is also a segment of
a 45" helix that is perpendicular to the helix passing through the postorbital bar. This model
suggests that the postorbital bar is loaded in compression on the chewing side and in tension
on the non-chewing side; the supraorbital ridge is loaded in tension on the chewing side and in
compression on the non-chewing side.

Contents
Page
Introduction ...................................... 125
Methods and assumptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126
An a 1y sis ........................................ 127
Skull loading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
Analysis of the skull as a cylinder . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
Shear strains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
The orientation ofthe postorbital bar ........................ 130
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
General . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
Tests of the hypothesis ................................ 133
Apparent exceptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Further considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136

Introduction
The postorbital bar is a bony arch that extends from the frontal bone to the zygomatic arch in
the skulls of some groups of mammals. Most ungulates, in particular the selenodont artiodactyls,
125
0022-5460/85/090125+ 12 $03.00/0 @ 1985 The Zoological Society of London
126 W. S. G R E A V E S

have well-developed, bony postorbital bars. It has been postulated that this structure may, for
example, serve as a support or protector of the eye, may prevent the jaw muscles from impinging
on the eye, or may transmit or resist forces applied to the skull by horns, antlers, or by the jaw
muscles and lower teeth (e.g. Marinelli, 1933; Willemse, 1950; Cachel, 1979; Cartmill, 1980). While
all of the above suggestions seem intuitively reasonable, they are all so general that they tend not to
be explanations. Perhaps resisting tooth forces has the most appeal as a primary function. Yet
there are two major difficulties with the view that the postorbital bar serves as a force-resisting
buttress. Firstly, a tension-resisting ligament seems more appropriate in many cases, and secondly,
if the bar resists compressional forces generated a t the posterior end of the tooth row, why is it
absent in so many mammals where one would reasonably expect to find it?
The very presence, in a mammalian skull, of a postorbital bar formed of bone immediately
suggests that it functions as a strut resisting compression or bending a t least part of the time, since
a ligament is expected if only tensile forces are encountered (Oxnard, 1971). In mammalian
carnivores, where the bar is usually absent, the thickened fascia that spans the distance between
the postorbital processes forms a postorbital ligament which is thought to resist tensile forces
(Buckland-Wright, 1978). In addition, the cross-section of the zygomatic arch has its longest
dimension oriented dorsoventrally in carnivores, and the zygoma itself is curved into an arch
opening downward. These features are expected in a zygomatic arch resisting ventrally directed
forces from the masseter muscle (Hildebrand, 1974: fig. 18-7). In ungulates, it would seem even
more reasonable to expect strong tensile forces in the region of the postorbital bar, for the masseter
muscle, which is very large in this group, exerts a strong ventral force on the maxilla and jugal,
which form the anterior portion of the zygomatic arch. This force from below would tend to bend
the arch downward and the postorbital bar, connecting the upper edge of the zygoma to the top of
the skull, would be expected to be in tension. Why then is a bony bar present in ungulates?

Methods and assumptions

The general presence of a postorbital bar in ungulates but not in carnivores prompts one to consider other
features that correlate with the presence or absence of a bony bar. One of the more obvious differences
between these 2 groups is the dominance of the masseter and pterygoid muscles in ungulates, and the
temporalis muscle in carnivores. This difference appears significant in the present case because the large
temporalis in carnivores attaches to the brain case and may not have a major effect on the zygomatic arch
(Buckland-Wright, 1978). On the other hand, the large masseter muscle in ungulates, even though much of it
attaches to the jugal and maxilla, might have a significanteffect because it acts directly on the zygomatic arch.
Essentially, all carnivores and ungulates, both fossil and recent, with large temporal fossae, and therefore
presumably large temporalis muscles, do not generally have postorbital bars; those ungulates with small
temporal fossae and indications of large masseter muscles usually have a postorbital bar.
The purpose of this study is to present a theoretical mechanical analysis that offers an explanation for (1)
the presence of a bony postorbital bar in most ungulates, primates, and a few other mammals with large
masseter and pterygoid muscles, and (2) the absence of this element in most other mammals such as
carnivores with dominant temporalis muscles. The analysis, concentrating on ungulates and carnivores,
suggests that this bony bar does, indeed, resist compressional forces. These forces are present because the
skull is asymmetrically loaded during mastication in animals with a large masseter/pterygoid muscle
complex. A bony postorbital bar is necessary in these forms because strong torsional forces result that tend to
twist the braincase relative to the rest of the skull, necessitating a compressional strut to resist this tendency.
The skull is also loaded asymmetrically,but usually less so, when the temporalis is the dominant muscle and
the masseter and pterygoid are relatively small. Because of the orientation of the temporalis, twisting is
 P 0 S T 0 R B I T A I. B A R I27

minimized in this case and a bony postorbital bar is absent. Offering this hypothesis to account for a major
function of the postorbital bar is not meant to suggest that the bar might not perform other functions as well.
Moreover, the analysis in its simplest form is not likely to be applicable to mammalian groups, such as
rodents, that differ from ungulates and primates in having relatively large orbits and different jaw muscle
placement.

Analysis
Skull loading
Ungulates chew with the teeth on only one side at a time but the muscles and jaw joints on both
sides of the head are active during mastication (De Vree & Gans, 1976; Greaves, 1978; Walker,
1978; Weijs & Dantuma, 1981). This is also true of carnivores (Gorniak & Gans, 1980; Greaves,
1983). Therefore, muscle forces bringing the jaws together are dissipated, not only at the teeth but
also at both jaw joints (Greaves, 1978; Weijs, 1981). As a consequence of this mechanical
arrangement, as well as the activity of the jaw muscles on each side of the head, the jaw joint on the
non-chewing or balancing side is typically loaded more heavily than the joint on the chewing or
working side (Greaves, 1978; Hylander, 1979; Weijs, 1981; Weijs & Dantuma, 1981).
The important characteristic of this system for the analysis presented here is that the skull is
loaded asymmetrically. The jaw muscles are usually more active on the chewing side than on the
balancing side, but they both apply significant ventral forces to the skull. The reaction forces
applied to the skull by the lower jaw act at the bite point on one side of the skull as well as the two
jaw joints. But, the joint on the same side of the head as the bite point is not as heavily loaded as is
the opposite balancing side joint, and the teeth on the balancing side are not loaded at all.
Therefore, the lower jaw tends to push up on the tooth at the bite point and up on the jaw joint on
the opposite side, while the muscles on both sides of the head pull the sides of the skull down. The
skull is loaded in torsion as a result of this force regime.

Analysis of the skull as a cylinder

If the skull is imagined as a long cylinder whose axis approximates the anteroposterior axis of
the skull, then at one end the cylinder is analogous to the anterior, tooth-bearing part of the skull,
while the other end represents the braincase (basicranial flexion is ignored). If the same forces that
are applied to skulls are applied to the cylinder (Fig. l), then the masseter and pterygoid muscles
pull the skull down at each side, but the occluding teeth push up only on (say) the right side. The
anterior part of the skull cylinder thus tends to rotate clockwise about its long axis as a result of the
bite force and contralateral muscle force. But the more heavily loaded condyle on the left side
of the jaw, together with the muscles of the right side, rotates the diagonally opposite glenoid
cavity, at the braincase, counterclockwise. Therefore, the anterior and posterior units of the skull
cylinder tend to rotate relative to one another. This rotation or twisting presumably takes place at
the relatively weak region between the orbits.
An obvious difficulty with the analysis to this point is that both ungulate and carnivore skulls
are likely to be stressed asymmetrically, but only a few mammalian groups, such as the ungulates
and primates, have an ossified postorbital bar. However, different jaw muscles have different
effects in this system. The temporalis and masseterlpterygoid muscle groups attach at different
locations and have different lines of action. The large masseter and pterygoid muscles in
ungulates attach to the maxilla and, in lateral view, tend to be approximately perpendicular to the
axis of the cylinder representing the skull. This orientation contributes large muscle force
128 W. S. G R E A V E S

F I G .1. Superimposed drawings of a skull of Odocoileus and a transparent cylinder. The arrows pointing down represent
the force of the masseter/pterygoid muscle groups; the stippled arrow represents the force at the left side while the solid
arrow represents the muscle force on the right side. The straight arrows pointing up indicate the reaction force at the teeth
on the right side (stippled arrow) and the reaction force at the left jaw joint (solid arrow). The solid arrows considered by
themselves tend to rotate the skull (cylinder) counter-clockwise in the direction indicated by the solid curved arrow on the
surface of the cylinder. The stippled arrows, in the same fashion, tend to produce clockwise rotation.

components that rotate the skull, as in the idealized skull cylinder above. Imagine a vertical loop of
rope (muscle) hanging over a horizontal cylinder (skull). Assuming friction between the rope and
the cylinder, a vertical force applied to the rope is well oriented to rotate the cylinder around a
horizontal axis. On the other hand, in carnivores, the large temporalis muscles arise on the
braincase and tend to form a smaller angle with the basicranial axis (again, the axis of the
cylinder). Here the rope (muscle) is not vertical to the horizontal cylinder (skull) but lies at an
angle, say 30°, to the horizontal. A force along the rope in this case provides a smaller vertical
component to rotate the cylinder. Therefore, only small components of the muscle force are
available to rotate the braincase, as in the idealized skull cylinder above; the largest component of
the temporalis force is parallel to the axis of the cylinder in lateral view, and would merely pull the
anterior and posterior segments of the skull together, or perhaps bend them relative to one
another. In general, masseter and pterygoid muscles are oriented in such a way that they apply
larger torsional forces to the skull than does the temporalis muscle, assuming that the total force
generated by the temporalis muscle is not so great as to create very large vertical components.

Shear strains
Engineers have long known that a cylindrical bar that is twisted by forces perpendicular to its
long axis exhibits maximum tensile and compressive shear strains that lie along 45" helices on its
surface (e.g., Perry & Lissner, 1962). If the twisting forces are large enough, the bar fractures and
the broken surface has a helical contour. The edge of this contoured fracture surface is represented
on the surface of the cylinder as a 45" helix. This phenomenon can be demonstrated by twisting a
stick of chalk until it separates into two pieces.
Therefore, drawing a 45" helix on the surface of a cylinder, in effect, maps the orientation of
maximum tensile shear strains when the cylinder is put under torsion; maximum compressive
 POSTORBITAL BAR 129

F I G .2. Superimposed drawings of a skull of Odocoileus and a transparent cylinder. The pair of vertical arrows at each
end of the cylinder (skull) represent forces that rotate each end of the cylinder (skull) in the direction of the curved arrows.
The two helical lines drawn on the surface of the cylinder indicate the orientation of the maximum compressive and tensile
shear strains in a cylinder under torsion. These helices make an angle of 45" with the long axis of the cylinder and the
orientation of the deforming forces, and are mutually perpendicular. One helix is parallel to the postorbital bar on one side
of the head while the perpendicular helix is parallel to the postorbital bar on the opposite side. The axis of the cylinder is
parallel to the projection of the effectivejaw lever (i.e. a line from the bite point to the contralateral jaw joint) in lateral view.

shear strains lie along perpendicular 45" helices (Fig. 2). Since these strains are greatest at the
surface, torsion can be best resisted by buttressing material helically arranged at this location. A
segment of the surface helix appears as a line making an angle of 45" with the edges of the cylinder
in top and side views. In an end view of the cylinder, the maximum shear strains also lie at 45" to
the deforming forces.
In a skull under torsion, the maximum compressive and tensile shear strains should likewise lie
along 45" helices, even though the skull is a more complicated structure. However, the exact
orientation of the muscle and bite forces that twist the skull is often not obvious. Nevertheless, the
orientation of these forces can be estimated. The occlusal surfaces of the cheek teeth, on one side of
the head, and the jaw joint, on the opposite side, describe a plane (Gingerich, 1972;Greaves, 1980).
The other cheek teeth and the other joint define a second plane, These planes are perhaps best
visualized in a frontal view (Fig. 3). The lines connecting a jaw joint to any bite point lie within
one of these planes and represent the effective jaw levers (Greaves, 1980). In an ungulate with a
relatively small temporalis muscle, the major muscles of mastication are the masseters and medial
pterygoids. The lines of action of these muscle masses on either side are approximately
perpendicular to the projection of the jaw levers (within either plane) in a lateral view. The action
130 W. S. G R E A V E S

lines are not perpendicular to either of the above planes, in a frontal view, but each muscle has a
strong force component that is perpendicular to both of these planes (Greaves, 1980). Since the
major muscles closing the jaws have large force components that are perpendicular to the planes
and the jaw levers within them, the plane can thus be used as a reference from which to measure the
orientation of the postorbital bar in a number of living ungulates.

The orientation of the yostorbital bar

In lateral view, the postorbital bar of many selenodont artiodactyls lies at approximately 45" to
the projection of the effective levers lying within the planes described above (Fig. 3). In a dorsal
view of the skull, the postorbital bars form angles of 45" with the sagittal suture (90° with each
other). In a frontal view, each bar forms an angle of 45" with the effectivejaw lever lying within the
plane defined by the occlusal surfaces of the cheek teeth, on the same side, and the opposite jaw
joint (Fig. 3). Since the masseter/pterygoid complex on each side has a large force component
perpendicular to an effective jaw lever, this component must also lie at approximately 45" to the
postorbital bar.
The lines of maximum shear strains in a piece of chalk and the postorbital bar on an artiodactyl
skull have the same general orientation relative to the twisting forces. An obvious interpretation is
that the bar is functioning as a torsion-resisting strut. Actually, the postorbital bar is only a
segment of an imaginary 45" helix wound around a skull idealized as a cylinder (Figs 2 and 3). It is
therefore positioned along a line of maximum tensile or compressive shear strain. The bar is
required near the orbit to span the interval between the braincase and the rostral part of the skull.
Generally, the postorbital bar is not continued as a rib-like strut on to the braincase, presumably
because this curved surface is sufficiently strong. The posterior aspect of the jugal and maxilla
serves as a stout anterior extension of the postorbital bar.
The rostral and braincase regions of the mammalian skull are rigid structures. The orbits lie
between these two skull segments and interrupt the continuity of the skull. The conception of the
skull as a simple cylinder is now modified to one where the skull is modelled as a cylinder with
a narrow waist in the middle of its length, like an hourglass. In those cases where large twisting
forces act, such as in species with large masseter/pterygoid complexes, the connection between
the anterior and posterior ends of the cylinder is strengthened. To best resist torsion, material
should be placed as far from the axis of rotation as possible and oriented at an angle of 45" to
the twisting forces. As a further constraint, in a real skull the orbit must be left unobstructed.
A postorbital bar meets these criteria admirably and supports the waist of the skull. It is located
far from the axis of rotation and extends from the dorsal aspect of the braincase to the jugal,
which is the ventral portion of the anterior skull segment. It does not interfere with the eye
and, in fact, may help support it. Finally, it is oriented along the theoretical maximum tensile
and compressive shear stresses, i.e. a segment of a 45" helix imagined to extend around the
skull.
On the chewing side, compressive forces would be found in the postorbital bar because, as the
muscle forces on each side pull the middle of the skull ventrally, the upper teeth are pushed
dorsally by the lower teeth, and the jaw joint on the diagonally opposite side of the head is also
being pushed dorsally by the condyle of the lower jaw. The anterior and posterior segments of the
skull tend to rotate relative to one another because the other tooth row is unloaded and the
remaining jaw joint is lightly loaded. The postorbital bar on the non-chewing side would then be
under tension. This loading regime would reverse when chewing shifted to the other side.
POSTORBITAL B A R 131

I
132 W. S. G R E A V E S

According to the hypothesis, the postorbital bar must therefore resist both tensile and compressive
forces, but not at the same time.
An additional adjustment to resist torsion is the shape of the medial wall of the orbit and the
anterior wall of the braincase on the diagonally opposite side. These surfaces together form a
single warped surface interrupted at the interorbital constriction. The shape of this surface,
extending from one side of the head to the other, is remarkably like that of the three dimensional
helical fracture surface on the piece of broken chalk. Since this surface represents the location of
the maximum shear forces, one is tempted to suggest that this warped bony surface also resists
maximum shear forces in the skull.
The assumption that, in some cases, the line of action of the masseterlpterygoid muscle group is
approximately perpendicular to the projection of thejaw lever (bite point to contralateral joint) in
lateral view is reasonable (Greaves, 1980). Of course, the major line of action of the jaw muscles
will not always be perpendicular to the projection of the jaw lever in lateral view. In the camel,
where the temporalis muscle is very large, the resultant line of action of the jaw muscles, again in
lateral view, is clearly not perpendicular to the lever extending from the joint to the opposite bite
point. While the camel is an extreme case, specimens of each species would nevertheless have to be
examined to secure an estimate of the orientation, in lateral view, of the line of action of the
resultant jaw muscle force. The point of the above discussion is that the postorbital bar is oriented
at 45" to the line of action of major components of the jaw muscle forces (in approximately the
same orientation as the maximum tensile and compressive shear strains in a cylinder under
torsion), regardless of the line of action of individual muscles relative to the effective jaw lever.

Discussion
General
As mentioned above, a large temporalis and small masseter and pterygoid are correlated with
the absence of a postorbital bar and a small temporalis, and large masseter and pterygoid with the
presence of the bar (cf. Turnbull, 1970). Postorbital bars are often absent in early members of a
group but present in later forms. Among the horses, Hyracotherium did not have a postorbital bar,
but Equus does. Intermediate forms with incipient postorbital bars are present (e.g. Parahippus,
Osborn, 1918). Presumably, a threshold had been approached in this genus such that occasionally
a few members would have a postorbital bar. The early oreodonts provide another example.
Postorbital bars are also sometimes observed in domestic cat skulls. Turnbull's (1970) muscle
weight data indicate that, of all the carnivores, felids tend to have the smallest temporalis muscles
as well as the largest masseter muscles. If a threshold is involved, then one expects to find examples
within the felids.
The primates make up the other major group of mammals where a postorbital bar is usually
found. My estimated line of action of the resultant of the forces of the jaw muscles again forms an
angle of 45" with the postorbital bar (cf. Cachel, 1979). The temporalis in primates is often larger
and forms a fairly large angle with the long axis of the braincase in lateral view, and so contributes
more torsional force than does the carnivore temporalis.
According to the model described above, the ossified postorbital septum, found in some
primates, can perhaps be considered the functional analogue of the medial wall of the orbit in
artiodactyls. The anterior surface of the braincase, and the medial wall of the orbit on the other
side in artiodactyls, is remarkably like the warped broken surface of the piece of chalk. In
 POSTORBITAL BAR 133

primates, the anterior braincase is similar to this region in artiodactyls, although generally more
globular. On the other hand, the medial wall of the orbit in primates is essentially on the mid-line
and the ossified postorbital septum, when present, has approximately the same position as the
orbital wall in artiodactyls. This is not to suggest that other functions of the postorbital septum in
primates are not important (cf. Cartmill, 1980).
Supraorbital ridges, found in some primates, continue to excite interest in the anthropological
community. The ridge above each orbit, considered as a separate structure, is oriented along a
segment of a helix making an angle of approximately 90" with the adjacent postorbital bar. Thus,
when the postorbital bar is resisting maximum compressive forces, that half of the supraorbital
ridge on the same side is resisting maximum tensile forces; when the bar is resisting tensile forces,
that half of the ridge is dealing with compressive forces.
Artiodactyls also have what amount to supraorbital ridges. Since the orbits in these animals face
about 30" toward the front, the helices that extend through the centre-lines of the ridges intersect at
an angle of about 60" at the mid-line (Fig. 3). The centre-lines of a postorbital bar and the adjacent
supraorbital ridge remain perpendicular. In primates, where the orbits generally face more
anteriorly, the helices passing through the individual supraorbital ridges meet at larger angles at
the mid-line (Fig. 4). In extreme cases such as GoriZZu, the ridges essentially form a single
prominence extending from side to side in almost a straight line. However, this ridge is composed
of two segments, one over each orbit, each of which is oriented at 90" to its adjacent postorbital
bar. The common assumption is that the supraorbital ridges resist forces due to mastication. The
analysis described here suggests that this is true, but at the same time stresses that the orientation
of the ridge conforms to the theoretically predicted orientation that allows it to resist the
maximum tensile or compressive shearing forces, depending upon whether the ridge is on the
chewing or non-chewing side.

Tests of the hypothesis

The following natural experiment has the potential to test the above explanation for the
presence of the postorbital bar. If one side of the head of an ungulate is injured or diseased such
that mastication can no longer take place on that side, then certain predictions are obvious. Since
biting on the affected side is no longer taking place, the postorbital bar no longer resists
compressional forces, but only tensional forces due to chewing on the contralateral side. If the
problem persists, the bone of the postorbital bar on the affected side is expected to atrophy owing
to the absence of compressional forces.
An artiodactyl skull from a diseased animal in the Department of Oral Anatomy's collection
tends to support the hypothesis. A severe fungus infection has essentially destroyed the maxilla
and most of the jugal in a domestic cow skull. The appearance of hypereruption on the affected
side, and the greater wear on the non-affected side, indicate that mastication had either ceased, or
at least been greatly reduced on the diseased side. The absence of compressive forces on the non-
chewing side has led to atrophy of the postorbital bar. What remains of the bar (its cross-sectional
area is less than half that of the non-affected side) is a much thinner structure than the bar on the
working side, which has continued to resist compressive forces. Moreover, the suture on the non-
chewing side is much shorter and not in contact all along its length. This single case can only be
suggestive but does indicate a possible experimental approach. Unfortunately, earlier studies
concerned with removing muscles to determine the effect on their bony attachments are of little
help (e.g. Washburn, 1947; Avis, 1959, 1961).
134 W. S. G R E A V E S

F I G .4. Oblique views of the skulls and lower jaws of Pun (above) and Papio (below) demonstrating that the centre-lines
ofthe postorbital bar and the supraorbitdl ridge on the same side are approximately perpendicular. Because the orbits face
so far forward, extensions of the centre-lines of the supraorbital ridges are almost coincident; the two supraorbital ridges
form a single prominence extending across the skull. Compare with the artiodactyl in Fig. 3.
 POSTORBITAL BAR I35

Apparent exceptions
Apparent exceptions to the hypothesis, such as the fossil entelodonts and the marsupial
sabertooth, seem to have had large temporalis muscles, yet also had postorbital bars. The
temporalis muscle, in lateral view, usually forms a small angle with the long axis of the braincase,
and so has little rotating power compared to the masseter and pterygoid muscles. However,
judging from figures of these animals in Gregory ( I 95 l), the temporalis muscles were oriented so as
to have a more vertical pull, produced more torsion, and necessitated a postorbital bar. Generally,
temporalis size is a good predictor of the presence or absence of a postorbital bar, but size per se is
not enough, orientation is also critical.

Further considerations
This view of skull structure also suggests that the zygomatic arches tend to be compressed
dorsally and posteriorly on the chewing side, and tensed ventrally and anteriorly on the
balancing side. The suture between the jugal and the squamosal remains open in artiodactyls.
Given that this suture may resist tension as well as compression, one prediction is that the
orientation of the fibres spanning this suture will consist of two mutually perpendicular groups,
each resisting the appropriate stress.
Previous studies have postulated forces in various parts of the skull and demonstrated these
forces. For example, Buckland-Wright (1978) stressed both upper carnassial teeth at the same time
in cat skulls, and found strains in predicted locations. But since cats do not use both carnassials at
the same time, and since one suspects that almost any loading regime will produce some strain in
most parts of the skull, simply finding 'predicted' strains perhaps does not tell us a great deal.
Therefore, experiments mimicking the natural experiment mentioned above may be a more
productive approach.
Finally, while the rodents have yet to be studied in any detail, one may at least mention the many
cases where the infraorbital foramen is enlarged, and slips of the jaw-elevating musculature pass
through it on the way to their insertion on the mandible. In these forms, the outer rim of the
foramen forms a stout bar that might be considered to be a preorbital bar. This bar has the same
general 45" orientation as the postorbital bar in ungulates. Therefore, one may tentatively assume
that, where it is present, the enlarged outer rim of the infraorbital foramen in rodents is also a
torsion-resisting helical strut.

Summary
The key element in the present analysis is the tendency for the rostrum and braincase to rotate
relative to one another because of asymmetrical loading of the skull during mastication. Although
there are exceptions, in general, large masseter/pterygoid muscles are oriented in such a way that
they supply large twisting forces, while large temporalis muscles do not supply such forces. The
most appropriate way to resist shearing forces due to torsion is to place additional material as far
from the axis of rotation as possible and oriented at 45" to the twisting forces in a helical pattern.
The postorbital bar is spatially oriented on the skull so that it forms a segment of a helix that maps
the orientation of the maximum compressive and tensile shear stresses. This orientation suggests
that a primary function of the postorbital bar is to resist torsional forces in the skull. A major
practical prediction of the hypothesis outlined above is that, in many mammals, the line of action
of the resultant force of the masticatory musculature forms an angle of 45" with the postorbital bar
136 W. S. G R E A V E S

in lateral view. In a frontal view, large components of the jaw muscle force form an angle of 45"
with the postorbital bar. These general relationships may well be true in other vertebrates with a
postorbital bar as well.

The comments of H. R. Barghusen, M. L. Greaves, S. W. Herring and J. W. Osborn greatly improved

previous drafts of the manuscript. M. L. Greaves drafted the figures.

R E F E R E N C ES
Avis, V. (1959). The relation ofthe temporal muscle to the form of the coronoid process. Am. J . phys. Anthrop. 17: 99-104.
Avis, V. (1961). The significance of the angle of the mandible: an experimental and comparative study. Am. J. phys.
Anthrop. 1 9 55-61.
Buckland-Wright, J. C. (1978). Bone structure and the patterns of force transmission in the cat skull (Felis cutus).
J. Morph. 155 35-62.
Cachel, S. M. (1979). A functional analysis of the primate masticatory system and the origin of the anthropoid postorbital
septum. Am. J.phys. Anthrop. 5 0 1-18.
Cartmill, M. (1980). Morphology, function, and evolution of the anthropoid postorbital septum. In Evolutionary biology
of New Worldmonkeysandcontinentuldrift: 243-274. Ciochon, R. L. & Chiarelli, A. B. (Eds). New York: Plenum.
De Vree, F. & Gans, C. (1976). Mastication in pigmy goats (Capra hircus). Annls SOC.r . zool. Belg. 105 255-306.
Gingerich, Philip D. (1972). Molar occlusion and jaw mechanics of the Eocene primate Adapis. Am. J. phys. Anthrop. 36:
359-368.
Gorniak, G. C. & Gans, C. (1980). Quantitative assay of electromyograms during mastication in domestic cats (Felis
catus). J. Morph. 163 253-281.
Greaves, W. S. (1978). The jaw lever system in ungulates: a new model. J. Zool., Lond. 184 271-285.
Greaves, W. S. (1980). The mammalian jaw mechanism: the high glenoid cavity. Am. Nut. 116 432-440.
Greaves, W. S. (1983). A functional analysis of carnassial biting. Biol. J. Linn. SOC.2 0 353-363.
Gregory, W. K. (1951). Evolution emerging II. New York Macmillan.
Hildebrand, M. (1974). Analysis of vertebrate structure. New York: John Wiley.
Hylander, W. L. (1979). An experimental analysis of temporomandibular joint reaction force in macaques. Am. J .
phys. Anthrop. 51: 433-456.
Marinelli, W. (1933). Die functionelle Bedeutung der Postorbitalspange bei Giraffe und Pferd. Biologia gen. 9
201-222. [Cited in Willemse, 1950.1
Osborn, H. F. (1918). Equidae of the Oligocene, Miocene and Pliocene of North America, iconographic type revision.
Mem. Am. Mus. nut. Hist. 2 1-217.
Oxnard, C. E. (1971). Tensile forces in skeletal structures. J . Morph. 134 425-436.
Perry, C. C. & Lissner, H. R. (1962). The strain gage primer. (2nd edn). New York McGraw-Hill.
Turnbuil, W. D. (1970). Mammalian masticatory apparatus. Fieldiuna (Geol.) 18: 147-356.
Walker, A. (1978). Functional anatomy of oral tissues: mastication and deglutition. In Textbook of oral biology:
277-296. Shaw, J. H. et al. (Eds). Philadelphia: W. B. Saunders.
Washburn, S. L. (1947). The relation of the temporal muscle to the form of the skull. Anat. Rec. 99:239-248.
Weijs, W. A. (1981). Mechanical loading of the human jaw joint during unilateral biting. Acta morph. neerrl.-scand. 19:
261-262.
Weijs, W. A. & Dantuma, R. (1981). Functional anatomy of the masticatory apparatus in the rabbit (Oryctolugus
cuniculus L.). Neth. J. Zool. 31: 99-147.
Willemse, M. C. A. (1950). The shifting of the molar row with regard to the orbit in Equus and Girafla. Zool. Meded.,
Leiden 3 0 31 1-326.