Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The Lin-

nean Society of London, 2004? 2004

140?
207221
Original Article

M. ANTÓN
ET AL.
MASTOID ANATOMY IN FELIDS

Zoological Journal of the Linnean Society, 2004, 140, 207–221. With 7 figures

Implications of the mastoid anatomy of larger extant

felids for the evolution and predatory behaviour of
sabretoothed cats (Mammalia, Carnivora, Felidae)
MAURICIO ANTÓN1*, MANUEL J. SALESA1, JUAN FRANCISCO PASTOR2,
ISRAEL M. SÁNCHEZ1, SUSANA FRAILE1 and JORGE MORALES1

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1
Departamento de Palaeobiología, Museo Nacional de Ciencias Naturales. C. José Gutiérrez Abascal, 2. 28006
Madrid, Spain
2
Museo Anatómico, departamento de Anatomía Humana, Universidad de Valladolid, C. Ramón y Cajal 7, 47005
Valladolid, Spain

Received November 2002; accepted for publication July 2003

Muscle attachments in the mastoid region of the skull of extant felids are studied through dissection of two adult
tigers Panthera tigris (Linnaeus, 1758) Pocock, 1930, a lion Panthera leo (Linnaeus, 1758) Pocock, 1930 and a puma
Puma concolor (Linnaeus, 1771) Jardine, 1834, providing for the first time an adequate reference for the study of the
evolution of that region in sabretoothed felids. Our study supports the inference by W. Akersten that the main mus-
cles inserting in the mastoid process in sabretooths were those originating in the atlas, rather than those from the
posterior neck, sternum and forelimb. Those inferences were based on the anatomy of the giant panda, Ailuropoda
melanoleuca (David, 1869) Milne-Edwards, 1870, raising uncertainties about homology, which were founded, as
revealed by our results. The mastoid muscle insertions in extant felids differ in important details from those
described for Ailuropoda, but agree with those described for domestic cats, hyenas and dogs. The large, antero-
ventrally projected mastoid process of pantherines allows a moderate implication of the m. obliquus capitis anterior
in head-flexion. This contradicts the widespread notion that the function of this muscle in carnivores is to extend the
atlanto-cranial joint and to flex it laterally, but supports previous inferences about the head-flexing function of
atlanto-mastoid muscles in machairodontines. Sabretooth mastoid morphology implies larger and longer-fibred
atlanto-mastoid muscles than in pantherines, and that most of their fibres ran inferior to the axis of rotation of the
atlanto-occipital joint, emphasizing head-flexing action. © 2004 The Linnean Society of London, Zoological Journal
of the Linnean Society, 2004, 140, 207–221.

ADDITIONAL KEYWORDS: basicranial morphology – Felinae – hunting behaviour – Machairodontinae – neck

muscles – skull.

INTRODUCTION The most recent group to evolve sabretooth adapta-

tions were the machairodontines, members of the sub-
The predatory behaviour of sabretoothed carnivorous
family Machairodontinae within the true-cat family,
mammals has been a matter of debate for nearly a cen-
Felidae. Like all derived sabretoothed mammals, the
tury, and it remains one of the most intriguing issues
felid sabretooths differed from their non-sabretoothed
in modern palaeontology. With no extant carnivore
relatives in showing modified mastoid regions, and
displaying such adaptations, we are left with the fossil
these modifications were recognized by early workers
remains, and our knowledge of functional anatomy, to
as a key feature for understanding the animal’s mode
propose credible hypotheses about the way those pred-
of attack.
ators used their elongated upper canines for prey
The precise interpretation of the meaning of the
procurement.
sabretooths’ mastoid morphology has varied consider-
ably. In the early 20th century, Matthew (1910) pro-
*Corresponding author. E-mail: mfanton@terra.es posed his ‘stabbing’ theory, according to which the

© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 207–221 207
208 M. ANTÓN ET AL.

sabretooths used the neck-flexing and head-flexing

muscles to power a downward stroke or stab, thus
allowing the upper canines to penetrate into the flesh
of prey. The mandible would have virtually no role in
this kind of attack, and the clear adaptations for gape
seen in sabretooths would be meant to keep the lower
jaw out of the way of the stabbing motion. In Mat-
thew’s scheme, the supposed hypertrophy of several
muscles that depress the head and neck was the key to
the success of the stabbing motion. As evidence for
that adaptation, he mentioned the enlarged trans-
verse processes of cervical vertebrae, which house the
insertions for a group of neck-flexing muscles called

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the scalenes, and the enlarged mastoid process, which
in turn houses the insertions for the sterno-cleido-
mastoid group, a set of muscles that depress the head,
pulling from their origins at the sternum and upper
arm (Fig. 1). Matthew’s theory was refined by Simpson
(1941), who proposed that the inertia of the predator’s
rush toward prey would be incorporated into the stab-
bing motion, and it was accepted by many specialists
for several decades. However, other workers (Bohlin,
1947) argued that, among other problems, the long,
flattened upper canines of sabretoothed cats were too
fragile to withstand such violent use (Biknevicius &
van Valkenburgh, 1996).
Most problems of the stabbing theory were satisfac-
torily overcome with the ‘canine shear-bite’ hypothesis
(Akersten, 1985), according to which the sabretooths
killed prey by means of a modified type of bite. In
Akersten’s scenario, the jaw-closing muscles have a
supporting role during the bite, but the main force for
penetration of the upper canines is provided by the
head-flexing action of the atlanto-mastoid muscula-
ture. In the canine shear-bite, the mandible has a
Figure 1. Schematic drawing of the skull and cervical ver-
clear part to play as support for the opposing action of
tebrae of the scimitar-toothed cat Homotherium latidens
the upper canines, and the modifications for wide gape
showing the hypothetical motions of the stabbing bite (top)
are seen as adaptations to allow the lower jaw to play
and the canine shear bite (bottom). In the first case the
such a role while allowing clearance between the main rotation is around a point behind the thoraco-cervical
upper and lower canines. In addition, a recent study joint (white circle) and the posterior cervicals, whereas in
(Antón & Galobart, 1999) has shown that the cervical the second case, the main rotation occurs at the atlanto-
morphology of sabretooths reflects adaptations for occipital joint (white circle). In the stabbing model (top),
strong muscular control of motions of the neck includ- the pull of the brachiocephalic muscles (single headed
ing extension and lateral rotation, as would be neces- arrow) and of the scalenes (two headed arrow) provides the
sary to provide support for a canine shear-bite, rather main force for the strike. In the canine shear-bite model
than the exclusive emphasis on efficient neck flexion (bottom), the pull of the atlanto-mastoid muscles (short
required by the ‘stabbing’ theory. two-headed arrow) is the most important force for the pen-
A central difference between Matthew’s hypothesis etration of the upper canines.
and Akersten’s is that the former considers that the
modifications in the mastoid region of sabretooths can
be explained as related to changes in the insertion occipital joint, whereas advocates of the stabbing the-
area, and function, of the brachiocephalic and sterno- ory emphasized the importance of flexion of the whole
mastoid muscles, whereas in Akersten’s view, the neck and head turning around the thoraco-cervical
atlanto-mastoids were the most important muscles joint and posterior cervical intervertebral joints
involved. In other words, Akersten gave more impor- (Fig. 1). But when Akersten needed to support his
tance to rotation of the head around the atlanto- ideas on anatomical grounds, he found a paucity of

© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 207–221

detailed studies about the muscle insertions in the
mastoid region of large cats. In his own words,
‘Descriptions of modern felid and canid anatomy often
ignore, or are not consistent in the terminology and
interpretation of those muscles’ (Akersten, 1985: 5).
As a result, he based his hypothesis on a published
study on the anatomy of the giant panda (Davis,
1964). Akersten’s choice was supported by the com-
mon-sense assumption of similarity in the disposition
of muscle insertions across different families and even
suborders within the Carnivora, but it gave rise to
uncertainties about homology, creating an unsatisfac-
tory situation, which has lasted to this day.
Over 15 years later, we still lack detailed compara-
tive data from the anatomy of the animals that would
provide the most adequate comparison, i.e. the living,
large members of the family Felidae, to which the
extinct machairodontines belong. The most useful
study of the muscles of a large felid is Barone’s (1967)
paper on the myology of the lion, in which the neck
musculature is described in detail, but insertion areas
in the mastoid region are only mentioned and not
described or figured. Similarly, for the domestic cat,
anatomical studies abound, but again the insertion
areas of muscles have been described (Reinhard &
Jennings, 1935; Barone, 1989) but not figured in suf-
ficient detail; regardless, the differences in morphol-
ogy between small cats and pantherines go beyond
mere size and, as commented below, may imply differ-
ences in muscle function.
A further complication lies in the fact that all anat-
omy treatises state that the action of the muscles most
relevant to Akersten’s analysis, the obliquus capitis
cranialis, is to extend and to rotate laterally the
atlanto-cranial joint, rather than flexing the head as
was assumed by Akersten (Evans & Christensen,
1979; Barone, 1989). A more recent, specific study of
the suboccipital muscles in the neck of the domestic
cat shows that the atlanto-mastoids generate little
torque in the saggital plane (Selbie, Thomson & Rich-
mond, 1993). With these data in mind, Akersten’s
hypothesis would imply a case of function shift in a
muscle group for the machairodontines (Bryant,
1996). But it is also feasible that these muscles have a
head-flexing action in some modern carnivores,
reflected in mastoid morphology.
This study intends to provide detailed information
about the muscle insertions in the mastoid area of
modern pantherines, and then to discuss the likely
implications of our findings in terms of the evolution
and functional anatomy of sabretoothed cats.
MATERIAL AND METHODS
An adult lioness Panthera leo, two adult tigers
Panthera tigris, one male and one female, an adult
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 207–221
MASTOID ANATOMY IN FELIDS 209
female puma Puma concolor and an adult female
genet Genetta genetta (Linnaeus, 1758) Cuvier,
1816 were dissected by the authors at the Anat-
omy Department of the University of Valladolid. All
specimens were captive animals coming from the
Zoological Park of Matapozuelos, Valladolid prov-
ince. We also incorporated information from a pre-
vious dissection of an adult female striped hyaena
Hyaena hyaena (Linnaeus, 1758) Brünnich, 1771
and an adult male binturong Arctictis bingturong
(Raffles, 1821) Temminck, 1824 from the Madrid
Zoological Park. We also studied osteological mate-
rial of extant and extinct carnivores to compare the
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morphology of the mastoid region. The skulls of
extant carnivores studied (besides those of the dis-
sected species) include specimens of domestic cat
Felis catus Linnaeus, 1758, spotted hyena Crocuta
crocuta (Erxleben, 1777) Kaup, 1828 and wolf
Canis lupus Linnaeus, 1758. Reference is also
made of published material of extinct machairodon-
tine felids of the genera Homotherium Fabrini,
1890 and Smilodon Lund, 1842, as well as descrip-
tions of giant panda Ailuropoda melanoleuca
(David, 1869) Milne-Edwards, 1870 and other
ursids.
The dissected specimens were fresh and were not
prepared with any preservation products, so the
observation of muscle fibres was excellent but dis-
section had to proceed quickly before the tissues
deteriorated. Each specimen was dissected in a sin-
gle session, except in the case of the tigress and
the puma, which were frozen and subjected to a
second session. In all specimens, dissection pro-
ceeded from the most external muscle layer, except
in the lioness, which had been partly defleshed
before our dissection, so that muscles external
to the splenius were damaged or missing, but
the deeper layers relevant for this study were
intact.
In each specimen, we dissected the neck muscles
that insert on the mastoid region and on adjacent
areas of the skull and we identified origin and inser-
tion areas of each muscle. We made both digital and
conventional photographs of each subsequent mus-
cle layer in each specimen, and relevant details of
muscle structure and insertion were sketched in
pencil directly from the dissected specimens. Skulls
of large pantherines were available for comparison
and identification of skull features that emerged
during dissection, and the skulls and skeletons of
each dissected specimen were subsequently pre-
pared for detailed observation of the relevant osteo-
logical details.
We followed the muscular nomenclature used by
(Barone, 1989) in his treatise on the anatomy of
domestic mammals.
210 M. ANTÓN ET AL.
RESULTS
MUSCLE ATTACHMENTS IN THE MASTOID AREA OF
EXTANT LARGER FELIDS
The most external muscle inserting in the mastoid
area is the brachiocephalicus, which originates on the
humerus and has an aponeurotic insertion extending
from near the tip of the mastoid up the mastoid crest
and nuchal crest to the midline (Fig. 2). Immediately
internal to this muscle appear the fibres of the rhom-
boideus and splenius, which originate at the anterior
thoracic spinous processes and terminate in a common
aponeurosis extending just internal to that of the bra-
chiocephalic, and with broadly the same extension
(Fig. 3). At this same level, the fibres of the sterno-
mastoid muscle are seen, extending from their origin
at the sternum and terminating in a tendinous inser-
tion on the tip of the mastoid process. Some fibres of
the sternomastoid extend upwards, sharing with the
brachiocephalic a common aponeurotic insertion along
the mastoid crest (Fig. 3)
Internal to the m. splenius is the muscle mass of the
extensors of the neck, including the biventer cervicis,
semispinossus capitis, complexus and longissimus
capitis, which have fleshy insertions on the occipital
area, above the mastoid region. Inferior to these can
be observed the obliquus muscles: the obliquus capitis
caudalis extending from the spine of the axis to the
dorsal surface of the atlas wings, and the obliquus
capitis cranialis, extending from the ventral side of the
atlas wings to the mastoid process. The mastoid inser-
tion of the latter muscle is fleshy and extensive, cov-
ering much of the area of the process (Fig. 4A, B). In
the atlas, the origin area of the obliquus capitis crani-
alis occupies most of the ventral surface of the wing.
The superior fibres of this muscle are short, owing to
the small distance between their origin and insertion,
but the inferior fibres span a considerable distance
between the atlas wing and the lower part of the mas-
toid process. The insertion area in the skull is broadest
at its inferior tip. Superiorly, the insertion area is not
restricted to the mastoid process, and continues in a
narrowing band up the nuchal crest. In the felid spec-
imens dissected by us, we did not find evidence of an
accessory portion of this muscle extending dorsally
from the lip of the atlas wing, as described for the dog
(Evans & Christensen, 1979; Done et al., 1995). In the
puma specimen there were some additional fibres
extending dorsal to the atlas wing, in a position sim-
ilar to the mentioned ‘accessory portion’. Unlike the
main portion of the muscle, these fibres form a thin
layer, superficial to the m. obliquus capitis caudalis
and covered by the m. splenius. Cranially, these fibres
mingle with the latter muscle before it inserts on the
nuchal crest (Fig. 4B).
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 207–221
The origin of the m. digastricus on the paroccipital
process is found to be variable in extent. In all cases
the main fibres of the muscle attach to the paroccipital
process tip, but whereas in the male tiger the insertion
was apparently restricted to that process, on the
tigress some fibres of the muscle went to the tip of the
mastoid, and in the lioness the insertion extended to
the area around the base of the process, even into the
adjacent part of the tympanic bulla (Fig. 5A) In the
mandibular ramus, the digastric muscle insertion
extends from the level of the first molar anteriorly,
reaching the level of the symphisis.
The muscle rectus capitis lateralis originates on a
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small area of the ventral side of the atlas wing, medial
to the attachment of the m. obliquus capitis cranialis,
and inserts on the medial side of the paroccipital pro-
cess, medial to the insertion of the m. obliquus capitis
cranialis. In the puma, the insertion reaches the tip of
the paroccipital process, adjacent to the insertion of
the m. digastricus (Fig. 5B). In the genet, some fibres
attach to the tympanic bulla.
MUSCLE ATTACHMENTS IN MACHAIRODONTINES AND IN
OTHER CARNIVORES
The mastoid area in machairodont felids differs from
that of extant larger felids in several ways (Fig. 6).
The mastoid process itself is more developed, and it
projects in an antero-ventral direction to varying
extent, almost touching the postglenoid process in the
more derived machairodont genera such as Smilodon
and Homotherium. The paroccipital process is vari-
ably reduced, especially in some of the more derived
machairodont genera, and as a result the tip of the
mastoid process has a greater ventral projection than
the tip of the paroccipital, whereas the reverse is true
for the extant felids. The texture of the mastoid pro-
cess is often more rugose than in extant felids, and
more so in the more derived taxa. The orientation of
the tip of the process is variable, facing anteriorly in
some taxa, just as in extant felids, whereas in other
taxa it faces inferiorly. In most cases the process tip is
proportionally broader and more distinctly sculpted
than in extant felids. The mastoid crest is usually
more developed with a greater lateral expansion, and
as a result the surface of the mastoid process itself
tends to face more inferiorly and posteriorly, and less
laterally, than in extant felids (Fig. 7).
Compared with machairodontine and extant felids,
the mastoid area of canids shows a relatively small
mastoid process and a relatively large paroccipital
process. The mastoid process tip in canids is not placed
inferior but actually level with, or dorsal to, the axis of
rotation of the atlanto-occipital joint, whereas the
paroccipital process tip projects considerably more
ventrally. In bears and in the giant panda, the mastoid
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Figure 2. Photograph and schematic representation of superficial layer of head and neck muscles of male puma. 1, m.
brachiocephalicus.

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212 M. ANTÓN ET AL.

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Figure 3. Photograph and schematic representation of intermediate plane of neck muscles of male puma. 2, m. trapezius;
3, m. splenius; 4, m. sternomastoideus.

process is much larger and projects far more ventrally mastoid process, and the insertion area for the m.
than in the wolf, and indeed more than in extant obliquus capitis cranialis is contained in the vertically
felids. In hyaenids, the paroccipital is ventrally pro- orientated band between the paroccipital and the mas-
jected and anteriorly placed and thus very close to the toid crest.

© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 207–221

In his description of the giant panda, Davis (1964)
figures a bundle of muscle fibres similar in position to
the accessory portion of the m. obliquus capitis crani-
alis described in the dog (Evans & Christensen, 1979;
Done et al., 1995). As noted above, we did not find evi-
dence of such a portion in the studied pantherines,
although a thin band of muscle fibres was observed in
the puma. In a binturong and a striped hyaena previ-
ously dissected by us there was no accessory portion,
but in the genet, as in the puma, we observed a thin
band of muscle stemming from a small aponeurosis
that envelopes the lateral border of the atlas wing.
These fibres appear to be continuous with the m. obliq-
uus capitis cranialis but they join cranially with the
m. splenius and they lack a separate insertion area in
the skull.
DISCUSSION
Our observations of the muscle insertions in the mas-
toid region of modern felids support most of Akersten’s
interpretations about the myology of extinct sabre-
tooths, but also allow us to update and revise his
discussion of the head-flexing musculature. As he cor-
rectly extrapolated from Davis’s (1964) study of the
giant panda, the muscle fibres having a most exten-
sive attachment area in the mastoid process are those
originating on the ventral side of the atlas wings.
Given the distant relationship between felids and the
giant panda, Akersten cautioned that the muscles
extending between the ventral surfaces of the atlas
wings to the postero-lateral margins of the mastoid
process in Smilodon ‘may or may not be homologous
with those of Ailuropoda’ (Akersten, 1985: 5). Our
results show that those concerns about homology were
founded, because there are important differences in
detail between the positions of the muscle insertions
in extant felids and those described by Davis (1964) for
Ailuropoda. In figure 19 of Davis (1964) the insertion
of the m. obliquus capitis cranialis is seen to occupy
much of the surface of the mastoid and paroccipital
processes, extending to an area medial to the latter,
whereas the m. rectus capitis lateralis is seen to insert
on a portion of the mastoid process lateral to the inser-
tion of the previous muscle. Thus there appear to be
two atlanto-mastoid muscles in the giant panda. By
contrast, in extant felids dissected by us, the m. obliq-
uus capitis cranialis insertion occupies most of the
mastoid process, whereas the m. rectus capitis later-
alis inserts outside the mastoid process, in and near
the medial surface of the paroccipital process and
medial to the insertion of the previous muscle. The lat-
ter condition agrees with that described for the domes-
tic cat (Reinhard & Jenning, 1935; Barone, 1989), lion
(Barone, 1967), and with that described and figured
for the domestic dog (Barone, 1989), striped hyena
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 207–221
MASTOID ANATOMY IN FELIDS 213
(Spoor & Badoux, 1986) and genet (this paper). Con-
sequently, it can be said that in all these animals the
only atlanto-mastoid muscle is the m. obliquus capitis
cranialis. These differences suggest an apomorphic
condition for the panda or for the whole Ursidae, or
alternatively Davis (1964) may have been wrong about
the identity of the muscles in the specimen available
to him. In any event, dissection of additional speci-
mens of Ailuropoda would be desirable in order to
clarify this subject.
Some of the observed differences among the
reviewed carnivores in the disposition of some rele-
vant muscles may have a functional significance: the
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presence of an accessory dorsal portion of the m. obliq-
uus capitis cranialis in the giant panda (as figured but
not mentioned by Davis, 1964: fig. 92) and in the dog
(where it is described as having a separate insertion in
the skull; Evans & Christensen, 1979) would point to
the existence of an extensor function of the muscle
besides that of head-flexion. By contrast, its reduction
or absence among the reviewed members of the Feli-
formia points to an almost exclusive flexor action.
The lack of comparative data led Akersten to say
that ‘modern felids and canids possess tiny mastoid
processes with only very minute muscles extending
between them and the atlas wings’ (Akersten, 1985:
5). Our results show that generalization to be wrong.
Although it is true that modern canids have relatively
small mastoid processes (Fig. 7), those of tigers and
lions are relatively much larger, providing attachment
areas for powerful atlanto-mastoid muscles. A further
difference between large canids and larger felids is
that the mastoid process of the latter projects further
downwards than in the former, not only enlarging the
muscle attachment area but also changing the action
of the atlanto-mastoid muscles. Effectively, the fibres
of the m. obliquus capitis cranialis in larger felids are
located more clearly below the level of the occipital
condyles, and as a result of this change, the contrac-
tion of that muscle is more effective at flexing the
head, whereas in canids the head-flexing action would
be less efficient and the muscle would contribute to
stabilize the atlanto-cranial articulation when con-
tracting bilaterally, while unilateral contraction
rotates the head laterally (Barone, 1989). In the
domestic cat, there is little ventral projection of the
mastoid process, and the atlanto-mastoid muscles are
said to generate little torque on the sagittal plane (Sel-
bie et al., 1993). So we can say that modern larger
felids show an incipient stage of the implication of the
atlanto-mastoid musculature in head flexion during
the bite, which would become far more emphasized in
machairodontines. Contribution of the neck muscula-
ture to vertical motions of the head associated with
mastication is already known in domestic cats
(Gorniak & Gans, 1980), so that the adaptation of
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Figure 4. (A) Photograph and schematic representation of deep muscles of the neck in male tiger. 5, m. obliquus capitis
caudalis; 6, m. obliquus capitis cranialis; 8, m. digastricus; At, lateral border of the atlas wings; Ax, dorsal border of axis. (B)
Photograph and schematic representation of deep muscles in a male puma. f, additional superficial fibres of m. obliquus
capitis cranialis, dorsal to the atlas wing.

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 MASTOID ANATOMY IN FELIDS 215

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B
Figure 4. Continued

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216 M. ANTÓN ET AL.

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A
Figure 5. (A) Photograph and schematic representation of deep muscles of the neck in a lioness. The posterior portion of
the temporalis muscle has been removed to make visible the nuchal region of skull and neck muscles attaching to it. 7, deep
extensors of the neck, including rectus capitis dorsalis major and minor; Am, auditory meatus; Mp, mastoid process; Nc,
nuchal crest. (B) Photograph and schematic representation of deep muscles of the neck of a male puma in ventral view. 9,
m. rectus capitis lateralis.

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 MASTOID ANATOMY IN FELIDS 217

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B
Figure 5. Continued

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218 M. ANTÓN ET AL.

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P
M

P
M
Figure 6. Photographs of the mastoid region of skull in female lion, Panthera leo (top) and scimitar-toothed cat, Homoth-
erium latidens (bottom) from Incarcal, Spain (IN-I 929). Note that the back of the skull is broken in the fossil. Muscle inser-
tion areas are marked; muscle numbering as in Figs 1–5. M, mastoid process; P, paroccipital process.

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 MASTOID ANATOMY IN FELIDS 219

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Figure 7. Drawing of the skull and anterior cervicals of Panthera tigris (top) and Homotherium latidens (bottom) with
fibres of selected muscles. Muscle numbering as in Figs 1–5. A black circle in the condylar area represents the position of
the rotation centre of the atlanto-occipital articulation. Notice how, in Homotherium, most fibres of the obliquus capitis cra-
nialis extend well below that centre of rotation, and would therefore have a stronger head-flexing action. Notice also how
the greater distance between the posterior tip of the atlas wings and the tip of the mastoid process in Homotherium makes
for longer inferior fibres of the obliquus capitis cranialis muscle.

sabretooths would simply elaborate on an existing bio-
mechanical system (Bryant, 1996).
All this is in contrast with the interpretations of
Matthew, who considered that the most relevant mus-
cles for head flexion in machairodonts where the m.
brachiocephalicus (his cleidomastoids) and m. sterno-
mastoideus, and showed these muscles as occupying
with their attachments the full extent of the mastoid
process (Matthew, 1910: figure 9).
Our results clearly show that the insertion of the m.
brachiocephalicus and the more superior portions of
the sterno-mastoids, although broadly coinciding with
the position indicated in Matthew’s figure, are actu-
ally aponeurotic and restricted to a thin band in the
outer margin of the mastoid process, i.e. in the mas-
toid crest, whereas the attachment of the m. sterno-
mastoideus is restricted to the process tip. The
insertion of the m. obliquus capitis cranialis, by con-
trast, is fleshy and extensive, and the action of this
muscle is likely to have had a stronger effect on the
flexion of the head.
These findings support the interpretation that the
action of the atlanto-mastoid musculature (i.e. the m.
obliquus capitis cranialis) was the most relevant for
head-flexing motions aiding the penetration of the
upper canines into the flesh of prey. The immediate
implication of this is that the relevant action involved
was rotation of the head around the atlanto-occipital
joint (an action associated with the canine shear-bite),
rather than rotation of the whole neck and head

© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 207–221
220 M. ANTÓN ET AL.
around the thoraco-cervical joint (an action associated
with the ‘stabbing’ bite). In the canine shear-bite, mus-
cular action from a static start provides the main force
for penetration, whereas in the ‘stabbing’ interpreta-
tion, the momentum gathered during rotation of the
head along a wide arc, or even during the leap toward
prey, is invoked to explain canine penetration. It is
true nonetheless that the downward projection of the
tip of the mastoid process would improve the efficiency
of the sternomastoid muscles as head flexors, an
action which would be incorporated to the canine
shear-bite mechanics as discussed by Akersten (1985)
and Antón & Galobart (1999).
The reduction in size and in ventral projection of the
paroccipital process in sabretooths implies changes in
the action of the m. digastricus, which is to depress the
mandible. In spite of the variation observed in the
extent of the area of origin of this muscle in the extant
felids studied, the main attachment is always on the
tip of the paroccipital, and the lesser the ventral pro-
jection of the process tip, the greater the distance
between the origin and insertion of the fibres of the m.
digastricus. Such an increase in the vertical span of
the fibres would be advantageous in terms of the
increased gape required by the canine shear-bite
(Antón & Galobart, 1999).
CLOSING COMMENT: ANATOMICAL
DESCRIPTIONS, SOFT TISSUE
RECONSTRUCTION AND
PALAEOBIOLOGICAL INFERENCE
The importance of soft tissue reconstruction in fossil
vertebrates is being more and more widely recognized
among evolutionists. As summarized by Witmer (1995:
20) ‘(1) Soft tissues largely are responsible for the
existence, maintenance and form of bones, (2) judge-
ments about the form and actions of soft tissues are
(implicitly if not explicitly) the basis for a host of
palaeobiological inferences; and (3), with regard to
systematics, soft tissue relationships may provide
testable hypotheses on independence or nonindepen-
dence of phylogenetic characters’.
However, accurate soft tissue reconstruction is
impossible without a sound database about the rela-
tionships between bone and soft tissue in extant taxa.
Unfortunately, it appears that descriptive mono-
graphs about the gross anatomy of animals have lost
the favour of current zoological science, due at least in
part to a prejudice among modern scientists against
the descriptive aspect of natural history (Gould, 1989–
1991: 96–100). However, the field of palaeobiological
inference provides many occasions to regret the pau-
city of accurate descriptions of modern organisms, and
of the musculo-skeletal systems of modern vertebrates
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 207–221
in particular, the case of the mastoid region in carni-
vores being just one example of the problem. When-
ever palaeobiologists attempt to reconstruct soft
tissues in fossil mammals, the descriptive anatomical
references cited are often remarkably ancient (Spoor
& Badoux, 1986; Witmer, Sampson & Solounias, 1999;
Naples & Martin, 2000).
While preparing his monograph about the giant
panda, Davis commented about the lack of comparative
information in plaintive terms: ‘The muscles of the Car-
nivora are comparatively well known, but even for this
order our knowledge is at a primitive level. Descrip-
tions are incomplete and inaccurate, often doing little
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more than establish the fact that a given muscle is
present in species dissected. Even for domestic carni-
vores – the dog and the cat – the standard reference
works are full of inaccuracies and are inadequately
illustrated. Most of the genera of Carnivora have never
been dissected at all.’ (Davis, 1964: 146). He goes on to
comment on the importance of muscular differences for
the overall problem of evolutionary mechanisms, but
then asks: ‘How can such differences be detected and
evaluated?. Certainly not on the basis of existing
descriptions and illustrations.’ (Davis, 1964: 146).
Most of the descriptions that we used as reference to
discern the pattern of muscle insertions in the mastoid
region in carnivores simply did not exist when Davis
wrote his monograph, but even those references are
not enough, either in detail or range of species treated,
to solve the problem, and in general the situation
today remains much as it was in 1964. With our
results in hand, and given the insufficient references
available to Davis, there is now reason to question the
interpretations that he made of atlanto-mastoid mus-
culature in Ailuropoda, but in order to add to the dis-
cussion, it would be necessary to describe additional
specimens of the giant panda, and of other ursids.
Such investigations would be especially pertinent to
the subject of atlanto-mastoid muscle function in
sabretooths, because the giant panda and other ursids
share a mastoid morphology different from most mod-
ern carnivores and intriguingly similar to that of
machairodontines (Akersten, 1985).
As Witmer commented, ‘the accuracy of soft tissue
reconstructions is vital because mistakes in soft-tissue
inference cascade upwards through the “inverted pyr-
amid” of paleobiological inference, amplifying the
error.’ (Witmer, 1995: 22). If the soft tissue inferences
are based on an insufficient or inaccurate database
about the anatomy of modern taxa, then all the sub-
sequent steps will be questionable.
CONCLUSIONS
The results of our study show that the most extensive
muscular insertions in the mastoid process of modern

big cats are those of the obliquus capitis cranialis. The
sternomastoid muscle inserts on the tip of the mastoid
process, and the m. brachiocephalicus inserts through
a thin aponeurotic band along the mastoid crest. The
m. digastricus inserts on the tip of the paroccipital
process but can extend to adjacent areas depending on
individual variation.
These results allow interpretation, in terms of mus-
cle function, of the differences in mastoid morphology
between modern pantherines and machairodontines.
The enlarged, ventrally projecting mastoid process of
the latter implies that the insertion area of the atlan-
tomastoid muscles was larger and that the more infe-
rior fibres of these muscles had an enhanced action as
head flexors, being able to rotate the head along a
large arc around the atlanto-occipital articulation. The
head-flexing action of the portions of the sternomas-
toid inserting on the mastoid process tip would also be
improved by the ventral projection of the latter,
whereas the function of the m. brachiocephalicus
would be little altered. The fibres of the m. digastricus
were probably longer in machairodontines as a result
of the reduction of the paroccipital process, which
implied that the process tip was higher and thus fur-
ther from the insertion of the m. digastricus in the
mandibular ramus.
ACKNOWLEDGEMENTS
This paper was supported by project MCT-BTE2002-
00410. We are grateful to Angel Galobart (Instituto de
Palaeontología Miquel Crusafont) and Josep Tarrús
(Museu Arqueológic Comarcal de Banyoles) for the
opportunity to study the fossils of Homotherium lati-
dens from Incarcal.
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