THE ANATOMICAL RECORD 292:1246–1265 (2009)

New Insights Into Dinosaur Jaw

Muscle Anatomy
CASEY M. HOLLIDAY*
Department of Pathology and Anatomical Sciences, School of Medicine,
University of Missouri, Columbia, Missouri

ABSTRACT
Jaw muscles are key components of the head and critical to testing
hypotheses of soft-tissue homology, skull function, and evolution. Dinosaurs
evolved an extraordinary diversity of cranial forms adapted to a variety of
feeding behaviors. However, disparate evolutionary transformations in head
shape and function among dinosaurs and their living relatives, birds and croc-
odylians, impair straightforward reconstructions of muscles, and other impor-
tant cephalic soft tissues. This study presents the osteological correlates and
inferred soft tissue anatomy of the jaw muscles and relevant neurovascula-
ture in the temporal region of the dinosaur head. Hypotheses of jaw muscle
homology were tested across a broad range archosaur and sauropsid taxa to
more accurately infer muscle attachments in the adductor chambers of non-
avian dinosaurs. Many dinosaurs likely possessed m. levator pterygoideus, a
trait shared with lepidosaurs but not extant archosaurs. Several major clades
of dinosaurs (e.g., Ornithopoda, Ceratopsidae, Sauropoda) eliminated the epi-
pterygoid, thus impacting interpretations of m. pseudotemporalis profundus.
M. pseudotemporalis superficialis most likely attached to the caudoventral
surface of the laterosphenoid, a trait shared with extant archosaurs. Although
mm. adductor mandibulae externus profundus and medialis likely attached
to the caudal half of the dorsotemporal fossa and coronoid process, clear osteo-
logical correlates separating the individual bellies are rare. Most dinosaur
clades possess osteological correlates indicative of a pterygoideus ventralis
muscle that attaches to the lateral surface of the mandible, although the mus-
cle may have extended as far as the jugal in some taxa (e.g., hadrosaurs,
tyrannosaurs). The cranial and mandibular attachments of mm adductor
mandibulae externus superficialis and adductor mandibulae posterior were
consistent across all taxa studied. These new data greatly increase the
interpretive resolution of head anatomy in dinosaurs and provide the anatom-
ical foundation necessary for future analyses of skull function and evolution
in an important vertebrate clade. Anat Rec, 292:1246–1265, 2009. V C 2009

Wiley-Liss, Inc.

Key words: Dinosaur; functional morphology; jaw muscles

The dinosaur feeding apparatus comprised a compli-
cated network of jaw muscles that span an intricate
assemblage of intracranial joints that link the bones of
the skull together. Data from jaw muscles, such as their
homologies, attachments, and sizes, are important to
test hypotheses of cranial function and feeding behaviors
such as powered cranial kinesis, bite force estimation,
chewing behavior, and mechanical optimization. Not sur-
prisingly, jaw muscle reconstruction is a common prac-
tice among paleobiologists. However, despite a long
history of muscle reconstruction and soft-tissue infer-
Grant sponsor: NSF; Grant number: IBN-0407735; Grant
sponsors: Marshall University School of Medicine, The Jurassic
Foundation, Ohio University Student Enhancement Award.
*Correspondence to: Casey M. Holliday, Department of Pa-
thology and Anatomical Sciences, School of Medicine, Univer-
sity of Missouri, Columbia, MO 65211. Fax: 573-882-4612.
E-mail: hollidayca@missouri.edu
Received 9 June 2009; Accepted 9 June 2009
DOI 10.1002/ar.20982
Published online in Wiley InterScience (www.interscience.wiley.
com).

V
C 2009 WILEY-LISS, INC.
 DINOSAUR JAW MUSCLE ANATOMY 1247
ences in dinosaur science, several challenges still impede TABLE 1. List of anatomical abbreviations
the development of precise hypotheses of structure, func- aIC Internal carotid artery
tion, and evolution of the jaw muscles in the group. aMN Mandibular artery
Major cranial structures including the antorbital cavity asp Ascending process of pterygoid
(Witmer, 1997; Molnar, 2008), ceratopsid frills (Lull, aST Stapedial artery
1908; Dodson, 1996), and buccal emarginations (Galton cot Cotyle
1973; Knoll, 2008) have only relatively recently been cr Crest
identified as non-muscular structures. Regardless, the ect Ectopterygoid
basic anatomy of the adductor chamber of dinosaurs has ept Epipterygoid
lig Ligament
not been thoroughly reviewed in a comparative context ls Laterosphenoid;
and numerous issues still impact the interpretations of mAMEM m. adductor mandibulae externus medialis
the region’s evolution. mAMEP m. adductor mandibulae externus profundus
Many previous analyses of dinosaur jaw muscle anat- mAMES m. adductor mandibulae externus superficialis
omy framed their hypotheses on a broadly comparative mAMP m. adductor mandibulae posterior
framework (e.g., Dollo, 1884; Lull, 1908; Versluys, 1910; mDM m. depressor mandibulae
Russell, 1935; Janensch, 1936; Ostrom, 1961, 1966; mLPt m. levator pterygoideus
Haas, 1963). With the advent of cladistic methodologies, mPPt m. protractor pterygoideus
it is possible to bring increased rigor permitted by phylo- mPSTp m. pseudotemporalis profundus
mPSTs m. pseudotemporalis superficialis
genetically constrained methods such as the Extant Phy- mPTd m. pterygoideus dorsalis
logenetic Bracket approach (Bryant and Russell, 1992; mPTv m. pterygoideus ventralis
Witmer, 1995a, 1997). In the past, workers reconstructed n,aIM Internal mandibular nerve and artery
muscles based on mammalian anatomy (Dollo, 1884; nCID Nerve to constrictor internus dorsalis muscles
Russell, 1935) and even included a muscular cheek in pal Palatine
their reconstructions (Lull, 1908). Haas (1955), Ostrom pt Pterygoid
(1961, 1964), and later Norman (1984) re-evaluated qj Quadratojugal
many earlier descriptions and included their own new qu Quadrate
insights on several clades of dinosaurs whereas drawing ri Rictus
sh Surangular shelf
more from lepidosaur anatomy. tcr Tensor crest
Soft-tissue inferences in fossils carry their own pitfalls V1 Ophthalmic nerve
and caution is necessary when reconstructing the mus- V2 Maxillary nerve
culature of extinct animals, particularly when their feed- V3 Mandibular nerve
ing behaviors may differ substantially from their extant VIIhy Hyomandibular ramus of facial nerve
relatives (Haas, 1969; Bryant and Seymour, 1990; VIIpal Palatine ramus of facial nerve
Witmer, 1995a,b; Carrano and Hutchinson, 2002;
Holliday and Witmer, 2008). Parsimony necessitates con-
servative inferences of jaw muscle anatomy. However,
because archosaurs are so morphologically diverse and bones of lizards seemingly offer fewer tendinous corre-
variation in jaw muscle morphology certainly exists lates despite the presence of large aponeuroses, such as
among sauropsids (Haas, 1973; Schumacher, 1973; the quadrate aponeurosis and the bodenaponeurosis.
Buhler, 1981; Busbey, 1989; Vandenberge and Zweers, Osteological correlates are highly susceptible to the
1993; Iordansky, 1973; Wu, 2003; Holliday and Witmer, vagaries of ontogeny and it is to be expected that juve-
2007), phylogenetic bracketing may still be inadequate nile individuals have fewer osteological correlates than
for all soft-tissue inferences and in some cases, compel- more mature individuals do. In addition to these more
ling evidence may override conservative interpretations. discrete correlates, Hutchinson (2001a) introduced the
Interpretations of muscle scars can be problematic concept of bone surface homology to refer to the corre-
(McGowan, 1979; Bryant and Seymour, 1990; Bryant spondence of general osteological regions that are contin-
and Russell, 1992; Rieppel, 2002) and correspondence of uous through evolution. In the head, examples of
the muscle-bone relationship in the head has only been homologous bone surfaces may include the lateral sur-
tested in a few cases (e.g., Montanucci, 1989, Witmer, face of the parietal and dorsal surfaces of the pterygoid,
1995b, 1997; Hieronymus, 2002). Osteological correlates which m. adductor mandibulae externus profundus and
can indicate the presence, attachment site, and to some m. pterygoideus dorsalis consistently attach to, respec-
extent, the size of a jaw muscle’s attachment. Tendinous tively. However, homologous bony surfaces and their cor-
or aponeurotic attachments leave more robust muscle responding muscles are more difficult to identify on the
scars than fleshy attachments, though fleshy attach- quadrate, which underwent significant morphological
ments can be equally informative (Bryant and Sey- and myological transformations in different sauropsid
mour, 1990; Bryant and Russell, 1992, Carrano and clades (Holliday and Witmer, 2007). These evolutionary
Hutchinson, 2002). In general, osteological correlates of changes can prove to be problematic in soft-tissue
jaw muscles are common among extant sauropsids, reconstruction.
although particular clades appear prone to having more Caveats aside, reconstructing jaw muscle anatomy
well-developed correlates than others have. For example, remains an important task necessary to better under-
the temporal bones of extant crocodylians offer a rich stand cranial function and behavior in extinct animals.
tapestry of fossae and ridges that consistently reflect the Osteological correlates have proven useful in recon-
attachments of particular muscle bellies, their aponeuro- structing soft tissues and tracking their evolution in the
ses, and adjacent neurovasculature (Iordansky, 1973; cephalic (Witmer 1995a,b, 1997; Snively and Russell;
Holliday and Witmer, in press). However, the temporal 2007) and postcranial anatomy (e.g., Hutchinson,
1248 HOLLIDAY
TABLE 2. List of institutional abbreviations
AMNH American Museum of Natural History
ANS Academy of Natural Sciences
BMNH British Museum of Natural History
BP Bernard Price Institute
CM Carnegie Museum
CMN Canadian Museum of Nature
CMNH Cleveland Museum of Natural History
DMNH Denver Museum of Natural History
DNM Dinosaur National Monument
FMNH Field Museum of Natural History
IGM Mongolian Geological Institute
MCZ Museum of Comparative Zoology
MOR Museum of the Rockies
MNN Musée National du Niger
ROM Royal Ontario Museum
SGM Ministère de l’Energie et des Mines
TMP Royal Tyrell Museum of Paleontology
UMNH Utah Museum of Natural History
USNM United States National Museum
ZG Zigong Dinosaur Museum
2001a,b; Carrano and Hutchinson, 2002; O’Connor,
2006) of extinct archosaurs. Although many species may
share numerous features with extant taxa, other derived
groups, such as hadrosaurs and oviraptorosaurs, present
a number of unique challenges. Numerous studies have
focused on the jaw muscle anatomy of single taxa though
no study has surveyed jaw muscle anatomy across the
entire clade. To allay this gap, this article reviews the
homologies and phylogenetic support of the different jaw
muscles with a focus on those of non-avian dinosaurs. It
uses the Extant Phylogenetic Bracket approach (Witmer,
1995a) to constrain inferences of soft tissues in fossil
taxa and relies on the archosaur soft-tissue homologies
provided by Holliday and Witmer (2007). It presents
phylogenetically constrained rules of construction for
jaw muscle anatomy and a generalized atlas of dinosaur
adductor chamber anatomy, including the osteological
correlates and their inferred soft-tissues. Tables 1 and 2
list anatomical and institutional abbreviations.
MATERIALS AND METHODS
Anatomical data were analyzed within the framework
of the Extant Phylogenetic Bracket approach (EPB),
which uses the principle of parsimony to infer soft tis-
sues in extinct taxa (see Witmer 1995a,b, 1997; Carrano
and Hutchinson, 2002). First, the adductor chambers of
extant archosaur and reptile taxa were examined to
identify patterns of topological similarity in jaw muscles
and surrounding soft-tissues and to suggest hypotheses
of soft tissue homology (Fig. 1). These patterns were
identified from dissections, computed tomographic (CT)
data, and observations of over 100 extant fresh and ske-
letonized crocodylian, avian, lepidosaurian, and testu-
dine taxa described in Holliday and Witmer (2007).
Second, causally associated osteological correlates of
these soft tissues were identified in the skeletons of
extant taxa (Fig. 1B–D). For example, in mammals, par-
ticular muscles (e.g., the temporalis muscle) consistently
leave a fossa or crest on the skull (e.g., temporal fossa)
that is used to infer the presence of that muscle when
cadaveric specimens are unavailable. Last, presumably
homologous osteological correlates of jaw muscles and
relevant soft tissues were identified across a large
assemblage of fossil archosaur material including basal
archosaurs, crurotarsans, and non-avian dinosaurs (Fig.
1A, Table 3). These osteological data, which are the focus
of this article, serve as proxies for the soft tissues in the
fossil taxa and thereby form the congruence test of
homology (Patterson, 1982; Witmer, 1997) used to infer
the positions and homologies of jaw muscles and rele-
vant neurovasculature using the phylogenetically based
scoring procedure of the EPB.
The EPB relies on drawing anatomical inferences from
not only fossil taxa, in this case non-avian dinosaurs,
but also that clade’s closest-related, extant ‘‘bracketing’’
taxa (birds and crocodylians) and finally outgroup taxa
(lepidosaurs and testudines) (Fig. 1A). Given the above
assumption that causally associated osteological corre-
lates are homologous among extant and fossil taxa (i.e.,
the congruence test is sound), soft-tissue inferences can
be refined further based on the phylogenetic distribution
of the osteological correlates among fossil and extant
clades. A soft tissue, in this case a muscle, can be inter-
preted as a level I, II, or III inference depending on the
presence of the muscle’s correlate in: both extant bracket
taxa and the fossil taxon; one extant taxon plus the fos-
sil taxon; or only the fossil taxon, respectively. Addition-
ally, the reconstruction of the correlated soft tissue,
along with consistent topological patterns recognized in
the extant groups, can subsequently dictate where
neighboring structures should have been (i.e., the maxil-
lary nerve, neighboring muscle) and vice versa. If no
clear osteological correlates are present among the
extant bracket taxa despite the presence of the soft tis-
sue, the levels of inference can then be assigned a
‘‘prime’’ designation (e.g., I0 , II0 , III0 ) in which each sub-
ordinate prime level is a weaker inference (i.e., it draws
less phylogenetic support) than its predecessor in the hi-
erarchy. Following Carrano and Hutchinson (2002), indi-
vidual muscle attachments may vary in their
distribution, therefore, whereas the cranial attachment
of a muscle may have a Level I inference, its mandibular
attachment may only be a Level 2 inference.
RESULTS
Overview of Adductor Chamber
Osteological Correlates
Despite the relative ubiquity of fossae, crests, tuberos-
ities, spurs, flanges, and other muscle-related bony
structures in the skulls of dinosaurs and other reptiles
(Figs. 2–4), these correlates are quite plastic and can
vary among individuals of the same taxon, let alone
among different taxa. This phenomenon is in part likely
due to ontogenetic differences among individuals of a
well-represented taxon. In juvenile Brachylophosaurus,
the cranial correlate of m. levator pterygoideus is a fossa
(Fig. 2A), whereas in an adult specimen (Fig. 2B), the
correlate is a long, striated spur. As in Brachylophosau-
rus, this spur for m. levator pterygoideus may slightly
overhang the ophthalmic groove, or as in Lambeosaurus
(Figs. 2C and 5B), it may drape ventrally to then fuse
onto the basisphenoid, forming a separate ophthalmic fo-
ramen. However, many adult individuals may also have
different osteological correlates. Among Tyrannosaurus
individuals, only one observed individual (AMNH 5117)
possesses a crest in the temporal fossa (Fig. 5C) that
 DINOSAUR JAW MUSCLE ANATOMY 1249

Fig. 1. Anatomy and phylogeny of dinosaurs and their extant archo-
saur and outgroup relatives.(A) Phylogenetic relationships of dinosaurs,
extant bracketing taxa, and outgroups. Extant clades in bold. Numbered
nodes: 1, Sauropsida; 2, Archosauria; 3, Dinosauria; 4, Ornithischia; 5,
Thyreophora; 6, Ornithopoda; 7, Ceratopsia; 8, Saurischia; 9, Sauropoda;
10, Theropoda; 11, Coelurosauria; 12, Maniraptora; 13, Aves. (B) Muscle
attachment surfaces in the skull and mandible of the monitor lizard (Vara-
nus exanthematicus); (C) Muscle attachment surfaces in the skull and
mandible of the chicken (Gallus gallus); (D) Muscle attachment surfaces in
the skull and mandible of the alligator (Alligator mississippiensis). Homolo-
gous muscles are similarly color coded throughout figures.
1250 HOLLIDAY

TABLE 3. List of fossil non-avian dinosaur taxa Finally, different taxa have arguably homologous, but
studied and their identification numbers incredibly morphologically dissimilar osteological corre-
lates that preclude straightforward ranking systems.
Taxon Specimen identification
This problem is best represented by the coronoid attach-
Thyreophora ments of m. adductor mandibulae externus profundus on
Scelidosaurus BMNH R1111 the mandible. Among theropods, the correlate varies
Stegosaurus CM 41681, DMNH 2818, between a shallow depression, a smooth, flat area, or a
USNM 4932, USNM 6645, small, rugose tuberosity, referred to here as the coronoid
USNM 2274
Panoplosaurus ROM 20892, ROM 1215 eminence (Fig. 4J,K). On the other hand, the osteological
Euplocephalus TMP 91.127.1, TMP 97.132.01 correlate takes the form of a huge boss in some taxa,
Edmontonia NMC 8531 such as Panopolosaurus (Fig. 4G). Finally, the coronoid
Ceratopsia processes of derived ceratopsids and hadrosaurs are
Psittacosaurus IGM 100/1132 characteristically dorsally elongate, may or may not
Leptoceratops CMN 8887, CMN 8889 have striations on them (Fig. 4). The loss of bony ele-
Protoceratops IGM 100/1246 ments also influences interpretations of muscles and
Montanaceratops AMNH 5244 their osteological correlates. Several clades of dinosaurs
Arrhinoceratops USNM 15583 lost the epipterygoid (Fig. 6), which is the origin of m.
Triceratops CMN 8528, MOR 699, MOR 1120,
MOR 1157, MOR 1194, pseudotemporalis profundus. Just because the element
UCMP 113697, UCMP was lost, does not necessarily mean the muscle was too,
137266, USNM 5740, because birds also lost the epipterygoid but still have
USNM 4286, USNM 24216 the muscle. However, epipterygoid loss creates a greater
Chasmosaurus TMP 79.11.156, TMP 83. 25.1, challenge when inferring whether the muscle shifted as
TMP 85.56.27, TMP in Brachylophosaurus (Fig. 2A,B) or if it is absent. Thus,
87.20.247, TMP 80.18.199, these issues make scoring osteological correlates of
TMP 86.18.11, TMP 86.18.82, muscles as a means of character mapping a complicated,
TMP 98.68.08, TMP 81.18.86, if not largely subjective procedure.
TMP 96.12.295, TMP 80.16.866,
TMP 90.36.390, TMP 89.18.86,
TMP 65.12.03, TMP 80.16.866, Overview of Adductor Chamber
TMP 90.36.390 Soft Tissue Anatomy
Centrosaurus CMN 348, ROM 43214, ROM 767
Brachyceratops USNM 7951 In general, jaw muscles are divided into groups based
Einosaurus MOR 456, MOR 492 on their topological relationships with the branches of the
Pachyrhinosaurus TMP 89.55.188, TMP 89.55.1072, trigeminal nerve (Lakjer, 1926; Holliday and Witmer,
TMP 87.55.101 2007). These muscle groups include m. constrictor inter-
Albertaceratops TMP 2002.69.01 nus dorsalis, and mm. adductor mandibulae internus,
Pachycephalosauria
Homalocephale Cast of IGM 100/51 externus, and posterior, most of which are then further di-
Stegoceras AMNH 6788 vided into smaller muscle bellies. With a few exceptions,
Stenotholus AMNH 25375 such as extant crocodylians, the muscles maintain the
Pachycephalosaurus USNM 12031 same topological relationship with numerous other soft
Ornithopoda tissues among tetrapods (Holliday and Witmer, 2007;
Lesothosaurus BMNH R2477 Holliday and Witmer, in press). It is expected that these
Hypsilophodon BMNH R197 same topological patterns were present in dinosaurs as
Thescelosaurus ROM 8537 well and there are no data to suggest otherwise. There-
Parksosaurus ROM 804 fore, if the grooves or foramina for the ophthalmic nerve
Orodromeus MOR 1141
Zephyrosaurus MCZ 4392 (V1) and stapedial (aST) and internal carotid (aIC)
Tenontosaurus MOR 682 arteries, which, for example, circumscribe the dorsal and
Camptosaurus USNM 5669 caudal margins of the origin of m. protractor pterygoideus
Brachylophosaurus CMN 8893, MOR 1071 on the basisphenoid, reasonable inferences of the muscle’s
Maiasaura MOR 693 attachment location can be made, regardless of the pres-
Edmontosaurus CMN 2288, CMN 2289, ence of a clear osteological correlate (Fig. 5). Similarly, the
MOR 003, ROM 658, mandibular nerve (V3) and artery (aMN), always pass
USNM 4737, UCMP 130156 between mm. adductor mandibulae externus and poste-
Prosaurolophus ROM 667, MOR 454 rior in the temporal region, and then once they are in the
Kritosaurus AMNH 5204
Gryposaurus ROM 873, ROM 44770 mandibular fossa, they pass lateral and rostral to m.
Tsaagan IGM 100/1015 adductor mandibulae posterior. The mandibular neuro-
Velociraptor AMNH 6515 vascular bundle then gives off lateral and medial (i.e.,
mylohyoid nerve) mandibular branches. Thus, if the posi-
tion of the muscle can be inferred (which is consistently
marks the separation between m. pseudotemporalis the medial mandibular fossa), the location of the neuro-
superficialis rostrally, from m. adductor mandibulae vascular bundle can also be inferred regardless if they
externus profundus, caudally. Homologous crests are leave a groove or foramen. Finally, in all non-crocodylian
absent in MOR 555, MOR 1125, CMN 11841, and TMP sauropsids, the maxillary nerve (V2) passes between mm.
83.30.1. On the other hand, related taxa such as Nano- pseudotemporalis superficialis and adductor mandibulae
tyrannus, individuals of Daspletosaurus (Fig. 3I), as well externus profundus before turning rostrally toward the
as other theropods (Fig. 3C,D) possess these crests. suborbital region. In the Triceratops braincase, MOR 699,
 DINOSAUR JAW MUSCLE ANATOMY 1251

Fig. 2. Osteological correlates of the orbitotemporal and temporal
regions. Photos have been standardized to left, lateral views for simplic-
ity. White arrows: correlate for m. levator pterygoideus; black arrows,
correlate for m. protractor pterygoideus. Scale bar equals 1 cm. (A) Bra-
chylophosaurus (juvenile; MOR 1071); (B) Brachylophosaurus (adult;
MOR 1071); (C) Lambeosaurus (CMN 2869); (D) Protoceratops (IGM
100/1246); (E) Pachyrhinosaurus (TMP 89.55.188); (F) Triceratops (MOR
1120); (G) Plateosaurus (AMNH 6810); (H) Camarasaurus, CM 11378; I,
Nigersaurus, MNN GAD 512; (J) Majungasaurus (FMNH PR2100); (K)
Daspletosaurus (TMP 94.143.1); (L) Tsaagan, IGM 100/1015.
1252 HOLLIDAY

Fig. 3. Osteological correlates of the temporal region and palate.
Scale bar equals 1 cm. Braincases in dorsal view: (A) Edmontosaurus
(CMN 2289); (B) Leptoceratops (CMN 889); (C) Troodon (TMP
82.19.23); (D) Carcharodontosaurus (SGM Din 1; left, lateral view); (E)
Herrarasaurus (MCZ 7064); (F) Daspletosaurus (CMN 8506). Palates:
(G) Triceratops (MOR 699; left, lateral view) (H) Same as (G) close up
of left, medial view; (I) Brachylophosaurus (MOR 1071; left, lateral
view); J, Plateosaurus (AMNH 6810, left, lateral view); (K) Camarasau-
rus (CMN 11378 left, lateral view of CT-scan-rendered image); (L) Das-
pletosaurus (TMP 2001.36.1; left, lateral view).
 DINOSAUR JAW MUSCLE ANATOMY
Fig. 4. Osteological correlates of the mandible. Scale bar equals 1
cm. (A) Thescelosaurus (ROM 3587; left, lateral view); (B) Edmontosau-
rus (CMN 2289; right, medial view); (C) indet. hadrosaur (ROM 1949;
left, lateral view); (D) Leptoceratops (CMN 8889; left, lateral view); (E)
Centrosaurus (ROM 767; left, lateral view); (F) Stegosaurus (CM 41681;
the maxillomandibular foramen is situated directly ven-
tral to the laterosphenoid buttress, which is rather rostral
to where the foramen is in many dinosaurs (Fig. 5A).
However, the foramen is angled caudally and has a long
groove on its caudal edge, suggesting that the maxillary
(and mandibular) nerves are pushed caudally by m. pseu-
1253
right, medial view); (G) Panoplosaurus (ROM 1215; right, medial view);
(H) Plateosaurus (AMNH 6810; right, medial view); (I) Camarasaurus
(CM 11378; right, medial view); (J) Dromaeosaurus (AMNH 5356; left,
lateral view); (K) Tyrannosaurus (MOR 008, right, medial view); (L) Cae-
nognathus (ROM 8776; left, lateral view).
dotemporalis superficialis, which otherwise has no clear
osteological correlate on the braincase. Thus, nerve fora-
men morphology can help identify the presence of particu-
lar muscles. These examples illustrate how neurovascular
tissues, muscles, and their osteological correlates can
reciprocally illuminate one another.
1254 HOLLIDAY

In addition to topological patterns, the regions of

attachment can also be used to categorize the different
muscle groups into the temporal muscles (e.g., mm.
pseudotemporalis superficialis and adductor mandibulae
externus profundus), palatal muscles (e.g., mm. pterygoi-
deus and adductor mandibulae posterior), and the jaw
opening muscle m. depressor mandibulae. Many saurop-
sids, including most dinosaurs, also maintain m. con-
strictor internus dorsalis muscles that span the
orbitotemporal region between the braincase and the
palate (Haas, 1973; Rieppel, 2002; Holliday and Witmer,
2008). This organization will be used in the ‘‘Results’’
section later in which each muscle name will be followed
by its abbreviation and general levels of inference for its
origin and insertion in the format: Muscle name (abbre-
viation-origin/insertion). Table 4 lists the hypotheses of
muscle homologies from Holliday and Witmer (2007), the
muscles’ general attachments and their respective EPB
inference level for their origin and insertion.

Orbitotemporal Muscles
M. tensor periorbitae (mTP–Level I/II0 ). M. ten-
sor periorbitae (i.e., m. levator bulbi) attaches to the ros-
tral edge of the prootic in lepidosaurs and a prominent
crest on the lateral surface of the laterosphenoid in croc-
odylians and birds (Haas, 1973; Elzanowski, 1987). The
muscle then attaches to the rostral portion of the orbital
septum in birds and lepidosaurs and to the preotic pil-
lars in crocodylians, forming a sling under the orbital
contents in all taxa. All dinosaurs possess the caudal
osteological correlate of m. tensor periorbitae: the later-
osphenoid buttress, or antotic crest, making its recon-
struction a Level 1 inference. This sharp edge of the
laterosphenoid is the attachment of the muscle and
clearly separates the temporal region from the orbital
region (e.g., Fig. 2). In some cases, the muscle creeps
onto the rostral surface of the laterosphenoid, as in
Majungasaurus (Fig. 2J). On the other hand, correlates
for the muscle’s rostral attachment are yet to be found.

M. levator pterygoideus (mLPt–III/III). In lepido-

saurs, m. levator pterygoideus originates on the lateral
surfaces of the parietal and prootic (e.g., Lakjer, 1926;
Haas, 1973). This small muscle passes ventrally to insert
on the ventromedial surface of the epipterygoid and is
thought to help modulate movements of the palate dur-
ing feeding, particularly in taxa that express cranial
kinesis (Holliday and Witmer, 2008). The muscle is
absent in birds and crocodilians, although likely because
of different phenomena. Both extant archosaur clades
lost the epipterygoid, but crocodilians sutured the palate
to the braincase, thereby eliminating the orbitotemporal
space altogether. Some avian taxa occasionally possess a
short ligament between the braincase and the
quadrate that may be a rudiment of m. levator pterygoi-
deus or the epipterygoid (Dzerzhinsky and Yudin, 1982).
M. levator pterygoideus does not leave any consistent
osteological correlates on the cranium or palate in
lepidosaurs.
Reconstructions of m. levator pterygoideus’s cranial
and palatal attachments are Level III or III0 inferences,
the weakest possible soft-tissue inference (Table 4). Fig. 5. Muscle attachments, trigeminal nerves, and arteries of the
Regardless, osteological and topological data, and sup- braincases of select dinosaurs in left, lateral view. (A) Triceratops; (B)
Brachylophosaurus; (C) Tyrannosaurus.
port from lepidosaurian outgroups, indicate many non-
 DINOSAUR JAW MUSCLE ANATOMY 1255

Fig. 6. Palate anatomy and evolution in non-avian dinosaurs. (A) Palate morphology and homoplastic
epipterygoid loss in non-avian dinosaurs; Palates with muscle attachments in left, lateral view of: B, Tri-
ceratops; C, Brachylophosaurus; D, Tyrannosaurus.
1256 HOLLIDAY

TABLE 4. The jaw muscles of extant and extinct sauropsids coupled with each muscle attachment’s
general level of inference in non-avian dinosaurs
Level of inference
Abbreviation Full name Origin Insertion
mTP m. tensor periorbitae Laterosphenoid buttress I Rostral border of orbit II0
MLPt m. levator pterygoideus Laterosphenoid dorsal to III Medial surface of III
ophthalmic pterygoid and epipterygoid
foramen/groove
MPPt m. protractor Ala basisphenoid ventral to II Medial surface of II0
pterygoideus ophthalmic foramen/groove pterygoid and quadrate
MPTd m. pterygoideus Dorsal surface of I Medial surface of articular I
dorsalis rostral portion
of pterygoid and palatine
MPTv m. pterygoideus Caudoventral I Lateral surface of I
ventralis surface of pterygoid articular and surangular
mPSTp m. pseudotemporalis Lateral surface of I0 Medial surface of I0
profundus epipterygoid surangular/medial
mandibular fossa
mPSTs m. pseudotemporalis Rostromedial portion of I Medial surface of coronoid II0
superficialis temporal fossa eminence/rostral medial
mandibular fossa
mAMEP m. adductor mandibulae Caudomedial portion I Coronoid eminence I
externus profundus of temporal fossa
mAMEM m. adductor mandibulae Caudolateral portion I Coronoid eminence/dorsomedial I0
externus medialis of temporal fossa surface of surangular
mAMES m. adductor mandibulae Medial surface of I Dorsolateral surface I
externus superficialis upper temporal bar of surangular
mAMP m. adductor mandibulae Lateral surface I Medial mandibular fossa I
posterior of quadrate

avian dinosaurs likely possessed the muscle. Numerous
dinosaur taxa possess clear osteological correlates attrib-
utable to this muscle on the laterosphenoid. Cranial
osteological correlates of m. levator pterygoideus can be
identified in ceratopsids, ornithopods, some neosauro-
pods, and tyrannosaurs (Fig. 3). However, clear corre-
lates cannot be easily identified in thyreophorans and
deinonychosaurs (Fig. 2L). The muscle’s scars are mor-
phologically diverse ranging from small fossae (Fig. 2A)
to large tuberosities (Fig. 2K). Despite these differences
in shape, the correlates are always found just dorsal to
the ophthalmic groove or lateral to the ophthalmic fora-
men when present, and ventral to the epipterygoid
cotyle on the laterosphenoid. The only soft tissue that
occupies this region of the braincase in extant taxa is m.
levator pterygoideus. Thus, despite weak phylogenetic
support from extant bracketing taxa, the topological and
osteological support for inferring the presence of the
muscle is strong. The levator pterygoideus muscle does
not leave any robust osteological correlates in most dino-
saur taxa. However, in ceratopsids and hadrosaurs, a
large, striated flange arises dorsally from the pterygoid
that suggests the presence of a large, tendinous levator
pterygoideus attachment (Fig. 3G–I).
M. protractor pterygoideus (mPPt–II/II0 ). M.
protractor pterygoideus attaches to the ventrolateral
surface of the basisphenoid in lepidosaurs or parabasi-
sphenoid in birds ventral to the ophthalmic groove and
rostrolateral to the internal carotid foramen. The muscle
only occasionally leaves a small crest or spur on the
braincase of birds, but in lepidosaurs, the muscle occu-
pies the triangular ala basisphenoid, whose border is
excavated by the surrounding neurovasculature such as
the stapedial and internal carotid arteries (Fig. 5). Croc-
odyliforms lost m. protractor pterygoideus once the pal-
ate was firmly sutured to the braincase (Fig. 1D).
However, rauisuchians, such as Saurosuchus, also have
an ala basisphenoid suggesting these taxa possessed the
muscle (Holliday and Witmer, in press). Reconstructions
of the protractor pterygoideus musculature are Level II
inferences for the braincase origin. Like that of lizards,
the dinosaur ala basisphenoid is roughly triangular but
varies in morphology ranging from small tubercles or
flanges in ankylosaurs and deinonychosaurs (Fig. 2L) to
large rugose bosses such as those found in tyrannosaurs,
hadrosaurs, and ceratopsids (Fig. 2). During manirap-
toran evolution, the ala basisphenoid regresses leaving
no easily discernable cranial correlate for m. protractor
pterygoideus on the braincase, shifting the cranial
attachment site inference from a Level II to II0 . Whereas
the dromaeosaurs Velociraptor and Tsaagan (Fig. 2L)
have a small ala basisphenoid, the troodontid Saurorni-
thoides, and basal bird Archaeopteryx both lack the fea-
ture. Because extant birds still possess this muscle and
some taxa do have small bony spurs indicative of the
muscle’s attachment, it is reasonable to infer that their
closest theropod ancestors also possessed the muscle.
The protractor muscle attaches along the medial surfa-
ces of the pterygoid and quadrate in extant taxa and
likely does the same in dinosaurs. However, the muscle
does not leave any marked scars indicative of its attach-
ment to the pterygoid or quadrate making the palatal
insertion a Level II0 . The lack of a palatal correlate of m.
protractor pterygoideus suggests a fleshy attachment
was present. But because the middle ear cavity, which
directly abuts the protractor muscle, also leaves similar
 DINOSAUR JAW MUSCLE ANATOMY 1257
correlates on the medial surface of the quadrate and
pterygoid, it remains difficult to determine the expanse
and size of either the air sac or muscle. Finally, the ala
basisphenoid is often excavated by a neurovascular
groove and is particularly prominent in large taxa such
as Triceratops, Brachylophosaurus, and Tyrannosaurus
(Fig. 4). Ostrom (1961) inferred this groove in hadro-
saurs to be for the maxillary and mandibular nerves as
they coursed ventrally from the maxillomandibular fora-
men. However, when the palate is articulated with the
braincase, this groove lies well medial and ventral to the
pterygoid, a position quite different from the position
where the nerve is found in other sauropsids. These
grooves on the ala basisphenoid are most likely for the
neurovascular bundle (nCID) that supplies the protrac-
tor muscles and other constrictor internus dorsalis
muscles.

Palatal Muscles
M. pseudotemporalis profundus (mPSTp–I/I’).
M. pseudotemporalis profundus attaches to the lateral
surface of the epipterygoid in lepidosaurs (Fig. 1B), the
lateral bridge of the laterosphenoid in crocodylians, and
the orbital process of the quadrate in birds (Vandenberge
and Zweers, 1993; Holliday and Witmer, 2007). The mus-
cle then attaches along the dorsomedial surface of the
mandible in lepidosaurs and birds. In crocodylians, the
muscle is vestigial and variably merges with the fibers
of the medial surfaces of the temporal muscles. There
are no clear osteological correlates of the muscle’s
attachment to the mandible, whereas those to the cra-
nium are fairly robust. Reconstructions of m. pseudotem-
poralis profundus are level I inferences in dinosaurs
that have epipterygoids. However, because crocodylians
and birds eliminated the epipterygoid and shifted the
position of the muscle (for reviews see Holliday and
Witmer, 2007, 2008, in press) interpretations of dinosaur
anatomy may not be clear. Among dinosaurs, the muscle
likely attached to the lateral surface of the epipterygoid,
as in lizards, and the surface of the bony element in
large theropod taxa occasionally has a fossa suggesting
the attachment of this muscle. However, several clades
of dinosaurs also lost the epipterygoid including hadro-
saurs, ceratopsids, and sauropods (Fig. 6). The position
Fig. 7. Mandibular anatomy and m. pterygoideus ventralis attach-
of the origin of m. pseudotemporalis profundus in cera- ment hypotheses in two derived non-avian dinosaurs. (A) Brachylo-
topsids and sauropods is unclear and it is possible that phosaurus: right, conservative, ventral mandibular attachment of m.
they lost the muscle. However, ornithopods (e.g., Figs. pterygoideus ventralis; left, osteological correlates suggest a jugal
2A,B and 5B) often have a fossa present between the attachment of the muscle; (B) soft-tissue anatomy of the medial por-
inferred origins of mm. levator pterygoideus and pseudo- tion of the mandible of the hadrosaur Edmontosaurus; (C) Nanotyran-
temporalis superficialis that may be for m. pseudotem- nus: right, conservative, ventral mandibular attachment of m.
poralis profundus. Mandibular attachments of the pterygoideus ventralis; left, osteological correlates suggest a jugal
muscle are ill-defined and it is inferred that, like those attachment of the muscle; (D) soft-tissue anatomy of the medial por-
attachments of extant taxa, the muscle likely attached tion of the mandible of the theropod Tyrannosaurus.
along the medial surface of the coronoid process or
surangular. ces left by the nasal passages and paranasal air sinuses
in dinosaur taxa (Witmer, 1997). Occasionally, a small
M. pterygoideus dorsalis (mPTd–I/I). M. ptery- crest on the inner surface of the maxilla or lateral sur-
goideus dorsalis originates along the dorsal surfaces of face of the palatine may demarcate the shift from a mus-
the pterygoid and palatine bones in lepidosaurs, crocody- cle attachment caudally to the caviconchal recess and
lians, and birds (Witmer, 1995b, 1997). The muscle then antorbital cavity rostrally. However, these structures are
inserts onto the medial surface of the articular and ret- rare or often distorted in fossil taxa. The mandibular
roarticular process. The rostral extent of the palatal attachments of m. pterygoideus dorsalis are well sup-
attachments of m. pterygoideus dorsalis can be difficult ported by the common presence of a smooth excavation
to discern from the equally smooth and excavated surfa- along the medial surface of the retroarticular process of
1258 HOLLIDAY

Fig. 8. Jaw muscle anatomy in three dinosaurs in lateral view. Right, superficial muscles; left, deeper
muscles. (A) Edmontosaurus (CMN, 2289; modified with permission from Rybczynski et al., 2008) with ju-
gal removed; (B) Diplodocus (CM 3452); (C) Majungasaurus (FMNH PR2100).
 DINOSAUR JAW MUSCLE ANATOMY
the articular in dinosaurs. Therefore, both cranial and
mandibular reconstructions of the m. pterygoideus dor-
salis are Level I inferences.
M. pterygoideus ventralis (mPTv–I/I). M. ptery-
goideus ventralis attaches along the caudoventral edge
of the pterygoid in crocodylians and birds. Among croco-
dylians, the muscle conspicuously wraps around the ret-
roarticular process to attach on the ventrolateral surface
of the retroarticular process and surangular. The muscle
often attaches along the ventral edge of the retroarticu-
lar process but also wraps around the retroarticular pro-
cess in some birds to attach to the lateral surface of the
mandible and even attaches to the jugal in some parrots
(Hofer, 1950). It remains unclear if lepidosaurs possess a
clear, separate ventral belly of m. pterygoideus. Regard-
less, the muscle occasionally also wraps around to the
lateral surface of the mandible and in some taxa (e.g.,
Uromastix) it attaches to the lower temporal bar.
Among dinosaurs, the attachment of m. pterygoideus
ventralis to the ventral edge of the palate is a phyloge-
netically supported inference although clear palatal ana-
tomical correlates are rare (e.g., Fig. 3G) making the
origin a Level I inference. On the other hand, the man-
dibular insertion of m. pterygoideus ventralis is well-
supported by a smooth fossa on the ventrolateral surface
of the mandible making it a clear Level I inference. Like
the muscle’s attachment in crocodilians and many birds,
the muscle also likely wrapped around the retroarticular
process to attach to the lateral surface of the mandible
(Fig. 4). However, the extent of the muscle’s attachment
across the mandible is difficult to determine in most di-
nosaur taxa. Among most ornithischians and sauropods,
there is no clear demarcation between the attachment of
m. adductor mandibulae externus superficialis and m.
pterygoideus ventralis, though a few individuals have a
fossa for the former muscle (Fig. 4C) suggesting it does
not extend very far ventrally. Among theropods, the ven-
tral surface of the mandibular shelf likely marks the
dorsal extent of the pterygoid muscle (Figs. 4J and 7C).
Interestingly, both derived hadrosaurs and tyrannosaurs
often possess flanges and bony spurs that descend off of
the jugal and tend to point toward the retroarticular
process. In some cases (e.g., Brachylophosaurus), the
spurs are rugose and striated suggesting a tendinous
attachment. These morphologies suggest that m. ptery-
goideus ventralis may actually have attached to the ju-
gal, rather than simply the mandible (Fig. 7).
M. adductor mandibulae posterior (mAMP–I/
I). M. adductor mandibulae posterior is the most phyloge-
netically and anatomically consistent muscle in the
adductor chamber. Among extant taxa, the muscle
attaches across the lateral surface of the quadrate. In
crocodylians, the muscle leaves a number of crests and
fossae (Iordansky, 1973; Holliday and Witmer, 2007)
marking its aponeurotic and fleshy attachments whereas
among birds and lizards, correlates are rare (Fig. 1).
Osteological correlates of the muscle on the quadrate are
also rare among dinosaurs (Fig. 3). Regardless, data from
extant bracketing and outgroup taxa suggest that the
muscle also attached to the quadrate in these fossil taxa.
M. adductor mandibulae posterior consistently attaches
to Meckel’s cartilage within the medial mandibular fossa
in lizards, crocodylians, and birds, other than anseri-
1259
forms, where it attaches to the lateral surface of the man-
dible. All data from dinosaurs suggest that m. adductor
mandibulae posterior also filled the medial mandibular
fossa making it a Level I inference (Fig. 4).
Temporal Muscles
The temporal region is dominated by the vertically ori-
ented jaw closing muscles including m. pseudotemporalis
superficialis, which is a division of m. adductor mandi-
bulae internus and bellies of m. adductor mandibulae
externus including mm. adductor mandibulae externus
profundus, medialis, and superficialis. Significant
changes occurred in the organization of the temporal
muscles during archosaur evolution and the muscles’
relationships with the skull in crocodilians, birds, and
non-avian dinosaurs are not necessarily similar to those
of lepidosaurs. As will be discussed thoroughly later, the
organization of the temporal muscles changed in the
skull roof as well as on the mandible in archosaurs.
These changes create challenges when identifying
homologies as well as determining the attachments,
function, and evolution of the muscles in fossil taxa. The
following passages will document the temporal muscles
from superficial (i.e., m. adductor mandibulae externus
superficialis) to deep (i.e., m. pseudotemporalis superfi-
cialis; Figs. 2, 3, 8).
M. adductor mandibulae externus superficialis
(mAMES–I/I). M. adductor mandibulae externus super-
ficialis invariably attaches across the upper temporal
bar in lepidosaurs via a fleshy attachment that does not
leave any specific osteological correlate. The muscle then
attaches to the dorsolateral surface of the mandible and
only occasionally leaves a smooth, ovate region that may
be bordered laterally by a faint ridge marking the tran-
sition from muscle to integument attachment. In crocodi-
lians, the muscle attaches along the ventral surface of
the quadratojugal by means of a fleshy attachment later-
ally and aponeurotic attachment medially, which leaves
a rostroventrally oriented ridge that it shares with m.
adductor mandibulae posterior (Iordansky, 1973; Schu-
macher, 1973). The mandibular attachment of the croco-
dylian m. adductor mandibulae externus superficialis is
well-emarginated by a smooth region on the dorsal sur-
face of the surangular that is bounded rostrally by the
coronoid eminence left by m. adductor mandibulae exter-
nus profundus. Because birds lost the upper temporal
bar, m. adductor mandibulae superficialis generally
attaches across the lateral surface of the squamosal and
when present, the ventral surface of the postorbital pro-
cess (e.g., Anas), but also across the suprameatal shelf
in many others (e.g., Gallus, Larus, Phalacrocorax; Fig.
1B). In ratites (e.g., Struthio), m. adductor mandibulae
externus superficialis is greatly reduced and attaches
along the temporal fascia and only slightly to the lateral
edge of the postorbital process. The muscle often leaves
small crests marking its aponeurotic attachments on the
cranial surface in most birds.
As in extant taxa, the temporal bar is arguably the
best indicator for the attachment of m. adductor mandi-
bulae externus superficialis. Taxa with long upper tem-
poral bars likely had rostrocaudally broader muscles;
those with shorter bars have smaller muscles. Despite
this basic anatomical relationship with the skull, the
1260 HOLLIDAY
muscle does not leave clear osteological correlates. In
taxa that have rugose bone textures on the skull surface
(e.g., tyrannosaurs, abelisaurids), the muscle attachment
is easily recognizable because of its contrasting smooth
texture. Thus, its reconstruction is a well-supported,
Level I inference. However, the medial, or deep, extent
of the muscle is difficult to estimate because it does not
usually attach to bony structures that mark its bounda-
ries with deeper muscles such as mm. adductor mandi-
bulae externus medialis or profundus caudomedially, or
m. pseudotemporalis superficialis rostromedially.
The most problematic concern of reconstructions of m.
adductor mandibulae externus superficialis’s origin is
that despite the muscle’s likely attachment across the
medial surface of the squamosal and postorbital, basi-
cally all of the medial surfaces of the postorbital, post-
frontal, squamosal, jugal, and quadratojugal are smooth
because of the numerous soft-tissues that emarginate
the region including fascia, neurovascular bundles, peri-
orbital structures, and pneumatic diverticulae. Haas
(1963) interpreted m. adductor mandibulae externus
superficialis to attach along the postorbital bar, rather
than the upper temporal bar in Diplodocus based on his
evaluation of the smooth, slightly grooved medial surface
of the bone. However, this groove is more likely exca-
vated by postorbital or jugal vessels from the stapedial
artery that pass along the medial surfaces of the postor-
bital and jugal. Among theropod dinosaurs that have a
prominently excavated squamosal recess, the interface
between the pneumatic sinus and muscle on the squa-
mosal may mark the muscle’s caudal boundary (Witmer
and Ridgely, 2008). However, it is also possible that m.
adductor mandibulae externus superficialis simply
enclosed the medial portions of the sinus, which likely
derives from the suborbital diverticulum (Witmer and
Ridgely, 2008). In taxa that do not have well-developed
diverticular correlates, although their phylogenetic rela-
tionships support the structure’s reconstruction, the in-
ference of a muscle attachment versus pneumatic
structure is equivocal. Finally, many lepidosaurs also
have a small superficial muscle belly, m. levator anguli
oris, which is often associated with m. adductor mandi-
bulae externus superficialis (Haas, 1973). The muscle
originates on the pretemporal and lateral temporal fas-
ciae and then inserts on the rictus (i.e., corner of the
mouth; Haas, 1973). Extant archosaurs do not possess
this muscle nor does the muscle leave osteological corre-
lates on the skeleton. Thus, the reconstruction of m. le-
vator anguli oris is a Level III0 inference in dinosaurs.
In most lepidosaurs, the m. adductor mandibulae
externus superficialis attaches along the dorsolateral
surface of the surangular, caudal to the coronoid process.
The muscle is typically a fleshy, parallel-fibered belly
and occasionally leaves a fossa or a slight bony ridge
(Fig. 1B). In crocodylians, the muscle occupies the major-
ity of the dorsal surface of the surangular, caudal to the
coronoid eminence and leaves a faint ridge marking its
extent laterally. The muscle’s mandibular attachment
varies among birds and generally attaches along the lat-
eral surface of the mandible between the jaw joint cau-
dally, the attachment of m. adductor mandibulae
externus profundus rostrally and the attachment of m.
pterygoideus ventralis ventrally.
As in extant taxa, the dorsolateral surface of the sur-
angular is the most likely attachment of m. adductor
mandibulae externus superficialis among non-avian di-
nosaur taxa and the muscle does not lend many specific
identifiable correlates other than a smooth region of the
surangular. Haas (1955) reconstructed the muscle in
Protoceratops attaching across the lateral surface of the
surangular based on the smooth, shallow fossa on the
element and this is a reasonable inference. In basal
(e.g., Leptoceratops, CMN 8889) and derived ceratopsids
(e.g., Styracosaurus CMN 344), the smooth surangular
becomes incorporated into the caudal portion of the
large, rugose coronoid process suggesting a marked dif-
ference in muscle attachment type between the two
structures (Fig. 4G). The caudal, smooth surface is most
likely the attachment for m. adductor mandibulae exter-
nus superficialis (e.g., Ostrom, 1964). The dorsal surface
of the surangular forms the caudal portion of the coro-
noid process in basal ornithopods such as Thescelosaurus
(ROM 3587; Fig. 4A) and Dryosaurus (CM 3392), and
there is occasionally a slight shallow fossa long the dor-
sal surface of the element that is the correlate for m.
adductor mandibulae externus superficialis. However, in
derived ornithopods, the attachment of m. adductor
mandibulae externus superficialis is rarely demarcated
(e.g., Fig. 4C), but still most likely attaches along caudo-
dorsal surface of the surangular, just rostral to the jaw
joint (e.g., Ostrom, 1961; Rybczynski et al., 2008). In
sauropods, the dorsal edge of the surangular is often
smooth and acutely angled suggesting a very thin man-
dibular attachment of m. adductor mandibulae externus
superficialis. There are no clear, consistent osteological
correlates of the muscle on the lateral surface of the
mandibulae in Camarasaurus or Diplodocus (Fig. 5B).
Non-avian theropods have the most clearly defined man-
dibular attachments for m. adductor mandibulae exter-
nus superficialis because the surangular has a
prominent shelf on its lateral edge marking the lateral
extent of the muscle (Figs. 5 and 8).
M. adductor mandibulae externus medialis
(mAMEM–I/I0 ). M. adductor mandibulae externus
medialis is the most problematic of the temporal muscles
in several ways. For reasons discussed later in ‘‘Discus-
sion’’ section, it is difficult to interpret in the adductor
chamber of extant reptiles and its reconstruction in fos-
sil taxa is equally ambiguous. The muscle is large and
well-differentiated in lepidosaurs and attaches to the lat-
eral surface of the large temporal aponeurosis, the bode-
naponeurosis (Haas, 1973). The muscle attaches along
the caudal surface of the dorsotemporal fossa and may
leave a shallow fossa on the surface of the posttemporal
bar, lateral to the spur often left by the bodenaponeuro-
sis. However, its attachment to the mandible is unclear
because it may share a common attachment with mm.
pseudotemporalis superficialis and adductor mandibulae
externus profundus. In birds and crocodylians, the mus-
cle is often anatomically and always topologically indis-
tinguishable from mm. adductor mandibulae externus
profundus and superficialis (Holliday and Witmer, 2007).
In crocodylians, m. adductor mandibulae externus medi-
alis is a small, quadrangular muscle that occupies a
smooth region of the quadrate between the trigeminal
foramen and m. adductor mandibulae posterior, with
which it shares aponeurotic attachments. However, the
muscle melds with parts of mm. adductor mandibulae
 DINOSAUR JAW MUSCLE ANATOMY
externus profundus and superficialis as it attaches to
the mandible leaving no specific osteological correlate.
Among birds, many of which have divided m. adductor
mandibulae externus into more than three, simple bel-
lies, a discrete m. adductor mandibulae externus medi-
alis is difficult to separate from deep or superficial
portions without corroborating developmental evidence.
That said, these intermediate bellies of m. adductor
mandibulae externus typically attach to the temporal
fossa, or to fasciae or aponeuroses of the other temporal
muscles cranially and then to the coronoid region of the
mandible.
Reconstructions of the cranial and mandibular attach-
ments of m. adductor mandibulae externus medialis are
Level I0 inferences among non-avian dinosaurs. However,
these inferences are problematic, because interpretations
of attachments of m. adductor mandibulae externus
medialis depend on interpretations of where the sur-
rounding muscles attach, which typically leave corre-
lates that are more consistent. The mandibles of
hadrosaurs and ceratopsids (Fig. 4B, D, E) occasionally
have shallow fossae that pass between the coronoid pro-
cess and the jaw joint, a region that was certainly
bounded by mm. adductor mandibulae externus profun-
dus and superficialis, respectively. It could be expected
that m. adductor mandibulae externus medialis occupied
this intermediate position. However, this inference is
equal to the hypothesis that the muscle attached to the
coronoid process with m. adductor mandibulae externus
profundus.
M. adductor mandibulae externus profundus
(mAMEP–I/I). Musculus adductor mandibulae exter-
nus profundus is the deepest portion of the adductor
mandibulae externus muscle group. In lepidosaurs, it is
relatively small compared with the other temporal
muscles and attaches to the caudomedial corner of the
dorsotemporal fossa and a portion of the prootic, some-
what deep to mm. pseudotemporalis superficialis and
adductor mandibulae externus medialis. It attaches lat-
erally to the bodenaponeurosis and then prominently
attaches to the coronoid process. In crocodylians, m.
adductor mandibulae externus profundus is the only
muscle that occupies the dorsotemporal fossa, though
temporal osteological correlates of basal crocodylomorphs
show evidence of there being multiple muscles in the
fossa (Holliday and Witmer, 2007, in press). In crocodyli-
ans, the muscle attaches to the caudomedial corner of
the dorsotemporal fossa and then attaches as a tendon
to a characteristic rugosity on the dorsal surface of the
surangular rostral to m. adductor mandibulae externus
superficialis. In ratites and anseriforms, the muscle
attaches to the postorbital process and in galliforms and
most other birds, the muscle attaches to the temporal
fossa. The muscle then consistently attaches to the coro-
noid process in all birds.
Among dinosaurs, m. adductor mandibulae externus
profundus most likely occupied most of the caudomedial
portion of the dorsotemporal fossa and likely attached to
the sagittal and nuchal crests when present. However,
as noted earlier, its caudolateral border, which would
have been shared with mm. adductor mandibulae exter-
nus medialis or superficialis, is unclear. The muscle’s
rostral bony attachment is occasionally marked by a
1261
small spur or vertical crest in a few taxa (e.g., Carcharo-
dontosaurus, Daspletosaurus; Fig. 3D,F), or a subtle
shift in curvature of the dorsal edge of the fossa (e.g.,
Herrerasaurus, Hypsilophodon, Brachylophosaurus;
Figs. 2 and 3). However, in most taxa, there are no de-
finable osteological correlates other than the dorsotem-
poral fossa itself that indicate the rostral extent of m.
adductor mandibulae externus profundus. In Edmonto-
saurus (CMN 2289; Fig. 3A), there is a shallow fossa on
in the ventral portion of the temporal fossa that may
correspond to this muscle, which would then suggest
that m. adductor mandibulae externus medialis may
extend across the dorsal edge of the dorsotemporal fossa,
as in lizards. However, this is the only specimen
observed that bears this feature. That said, cranial
reconstructions of the m. adductor mandibulae externus
profundus are Level I inferences. Most non-avian dino-
saurs also possess mandibular osteological correlates of
the muscle including coronoid processes in most ornithi-
schians and smaller coronoid eminences in theropods
and thus, these the muscle’s mandibular attachments
are also Level I inferences.
M. pseudotemporalis superficialis (mPTs–I/I0 ).
The deepest and most rostral temporal muscle is m.
pseudotemporalis superficialis. In lepidosaurs, m. pseu-
dotemporalis superficialis attaches to the medial surface
of the dorsotemporal fossa. The muscle then attaches to
the medial portion of the coronoid region. During croco-
dyliform evolution, m. pseudotemporalis superficialis
shifted from a position similar to that present in lepido-
saurs to the caudal surface of the laterosphenoid and
does not attach within the dorsotemporal fossa proper in
extant crocodylians (Holliday and Witmer, 2007, in
press). The muscle then inserts on the rostral portion of
the medial mandibular fossa where it is compressed by
the pterygoid buttress and develops a sesamoid carti-
lage, the cartilago transiliens. In ratites other than
Apteryx, m. pseudotemporalis superficialis solely occu-
pies the dorsotemporal fossa and also attaches in the
medial mandibular fossa via an intertendon. In virtually
all other birds, the muscle is miniscule relative to other
jaw muscles and attaches along caudoventral edge of the
laterosphenoid buttress of the temporal region and then
to the medial surface of the coronoid region of the
mandible.
Reconstructions of m. pseudotemporalis superficialis
are Level I or I0 inferences (Table 4). Despite this strong
phylogenetic support, the position of m. pseudotempora-
lis superficialis is difficult to clearly identify among
many dinosaurs. Drawing from the muscle’s position in
lepidosaurs, many early studies considered it to be the
dominant muscle of the dorsotemporal fossa. Lull (1908)
reconstructed the muscle on the rostral surface of the
laterosphenoid, inside of the orbit, in Protoceratops.
However, as noted by Haas (1955) this was a novel, and
unsupported interpretation and the muscle likely occu-
pied part of the dorsotemporal fossa. Several authors
have suggested that m. pseudotemporalis superficialis
attached to a large excavation on the dorsal surface of
the frontal in many theropod taxa (e.g., Coria and Cur-
rie, 2002; Molnar, 2008). However, the shape, horizontal
orientation, surrounding bony structures, and the phylo-
genetic distribution of this structure suggest that a
1262 HOLLIDAY
muscle did not likely attach to it (Holliday, 2008; unpub-
lished data) and phylogenetic bracketing further sup-
ports that m. pseudotemporalis superficialis more likely
attached within the dorsotemporal fossa proper. How-
ever, like m. adductor mandibulae externus profundus,
m. pseudotemporalis superficialis does not commonly
leave specific osteological correlates on the skull other
than the characteristic smooth area of rostrolateral por-
tion of the dorsotemporal fossa (Fig. 3). As noted in the
earlier section, some individuals may have a crest or
curvature shift on the temporal fossa that indicates the
separation between m. adductor mandibulae externus
profundus and m. pseudotemporalis superficialis (Fig 3).
The rostral extent of the muscle is bounded by the later-
osphenoid buttress and the attachment of m. tensor
periorbitae.
The mandibular attachment of m. pseudotemporalis
superficialis is difficult to infer. The extant bracketing
taxa suggest that the muscle probably attached to the
rostral portion of the medial mandibular fossa in ankylo-
saurs, stegosaurs, pachycephalosaurs, and basal ceratop-
sids and iguanodontians. However, m. pseudotemporalis
superficialis likely attached to the medial surface of the
coronoid process in many derived iguanodontians and
ceratopsids. The derived ornithischian condition—a coro-
noid attachment—is hypothesized to be similar to that
present in lepidosaurs because in hadrosaurs and cera-
topsids, the medial mandibular fossa is very small and
significantly caudal to the coronoid process thereby giv-
ing the muscle a rostral moment, rather than the caudo-
vertical moment it provides in other taxa. Furthermore,
the coronoid processes of these taxa tend to have differ-
ent domains of striations on them, which suggest the
presence differently oriented muscle attachments (Fig.
4B,E). In theropods and sauropods, which have large,
rostrally extended medial mandibular fossae that are
very similar to those found in crocodylians and ratites,
inferring the muscle to attach in the rostral portion of
the fossa is a Level I inference.
DISCUSSION
Jaw muscle inferences offer a potential wealth of
phylogenetic, anatomical, and functional information.
However, there are a number of vagaries associated with
them and the distribution of osteological correlates on the
skull can differ among individuals as well as clades. On the
other hand, specific osteological correlates, and their soft-
tissue inferences, may be phylogenetically robust. For
example, with the exception of extant crocodylians, which
have significantly reorganized their jaw muscles, m. adduc-
tor mandibulae posterior is a relatively simple, parallel-
fibered muscle among extant sauropsids. Therefore, from a
phylogenetic standpoint, it is reasonable to infer that the
muscle was also parallel-fibered among non-avian dino-
saurs. This inference is anatomically supported by the lack
of crests and other osteological correlates that would sug-
gest tendinous attachments on the quadrates. The tempo-
ral muscles are more pinnate than m. adductor
mandibulae posterior, and thus, a pinnate m. adductor
mandibulae externus profundus or m. pseudotemporalis
superficialis is a relatively strong inference that is sup-
ported by the extant phylogenetic bracket as well as at
least a few individuals that have crests suggesting strong
aponeuroses. However, these inferences also rely on a con-
sistent distribution of correlates, which given the fragmen-
tary nature of skull specimens, is still a challenge. These
issues aside, there are several trends in dinosaur jaw mus-
cle anatomy and evolution that can be explored.
Protractor Muscles
Protractor muscles are key structures involved in cra-
nial kinesis, or the ability of an animal to move the pal-
ate independent of the braincase and mandible.
However, to what extent dinosaurs could display this
complicated behavior remains debatable (Holliday and
Witmer, 2008). Regardless as to whether the protractor
muscles are functional or evolutionary relics, they are
ubiquitous among dinosaurs. For example, compared
with those of basal ornithopods, the ala basisphenoid of
hadrosaurs have a greatly expanded tripartite morphol-
ogy suggesting an enlarged and modified muscle that
had pronounced tendinous attachments to the braincase
that radiated not only caudoventrally, but also rostrolat-
erally onto the palate. These data suggest m. protractor
pterygoideus may have become hypertrophied to resist
laterally oriented forces generated by the medially inset
dental batteries that derived ornithopods evolved
(Rybczynski et al., 2008). Similarly, the alae basisphe-
noid of many non-coleurosaurian theropods, such as Her-
rerasaurus, Allosaurus, and Majungasaurus are smooth,
triangular pendants that are attachments for a modest,
fleshy, pinnate m. protractor pterygoideus. However,
those of large tyrannosaurs become dorsoventrally elon-
gate, highly textured, and covered with numerous crests
that point ventrally or caudoventrally suggesting an
increase in aponeurotic attachment for m. protractor
pterygoideus.
These changes in morphology of the ala basisphenoid
may be functionally adaptive, but they may also simply
be the results of the evolution of large head size in
tyrannosaurs, ceratopsids and hadrosaurs. Similarly, the
disappearance of the ala basisphenoid during manirap-
toran evolution may be associated with trends in minia-
turization displayed in the clade [e.g., Sinosauropteryx,
(Currie and Chen, 2001); Bambiraptor, (Burnham,
2004); Sinovenator (Xu et al., 2002); Microraptor, (Xu et
al., 2003); Mei, (Xu and Norell, 2004), and Mahakala,
(Turner et al., 2007)]. Whereas adductor muscles are
known to scale positively with body size in lizards (Her-
rel and O’Reilly, 2005; Herrel et al., 2007), it remains to
be determined if protractor muscles scale similarly with
other parts of the feeding apparatus among extant and
fossil taxa. Neosauropods attained huge body sizes with-
out the accompanying increase in head size and reduced
their alae basisphenoid to thin flanges of bone that sug-
gest they had a simple soft-tissue septum that separated
the orbit from the ear. However, these taxa possess also
some of the most extreme head and feeding structures
found in dinosaurs (e.g., Nigersaurus, Diplodocus).
Thus, teasing out the differences between allometry and
functional significance in the system is challenging, but
it offers numerous new directions to explore.
Temporal Muscles
Muscles that directly attach to the bony surfaces of
the temporal region, such as mm. pseudotemporalis
superficialis and adductor mandibulae externus
 DINOSAUR JAW MUSCLE ANATOMY
profundus, are more likely to leave discernable osteologi-
cal correlates that support anatomical inferences. How-
ever, muscles that instead primarily attach to soft
tissues such as skin that covers the lateral temporal fen-
estrae, such as m. levator anguli oris, or large aponeuro-
ses, such as m. adductor mandibulae externus medialis,
will not leave bony evidence of their attachments. This
presents a major problem with reconstruction of struc-
ture in the temporal region because m. adductor mandi-
bulae externus medialis was likely an important muscle
in the temporal region of dinosaurs. Despite these rather
robust inferences, the anatomical and phylogenetic vaga-
ries of the muscle preclude estimations of the location
and potential size of the muscle in fossil archosaurs.
Whereas lepidosaurs have prominent m. adductor man-
dibulae externus medialis bellies, extant archosaurs do
not. This is problematic because, like all tetrapods, non-
avian dinosaurs probably had at least rudimentary ver-
sions of this muscle and phylogenetic bracketing is sup-
portive of the muscle’s reconstruction. However, from an
anatomical perspective, it is simply unclear where and
to what extent the muscle attached on the dorsotemporal
fossa or the coronoid region of the mandible.
The identification of evolutionary patterns of jaw
muscles is clouded by the difficulties in diagnosing
homologies of specific bellies among fossil taxa. Although
all dinosaurs had temporal muscles, exactly which
muscles occupied the dorsotemporal fossa are difficult
hypotheses to test. The most consistent inferences are
that m. adductor mandibulae externus superficialis
attached to the upper temporal bar and m. pseudotem-
poralis superficialis attached to the caudal surface of the
laterosphenoid. The region’s organization in extant taxa
brackets inferences to a point: lepidosaur temporal fos-
sae have three muscles (mm. pseudotemporalis superfi-
cialis, adductor mandibulae externus profundus and
medialis); crocodylians and neognath birds have one (m.
adductor mandibulae externus profundus). However, it
is clear that the extant conditions in crocodylians and
birds are highly derived and numerous data suggest fos-
sil crocodylomorphs and maniraptoran dinosaurs cer-
tainly had multiple muscles in the dorsotemporal fossa.
The few osteological correlates of temporal muscles that
do exist among non-avian dinosaurs indicate that the
mm. adductor mandibulae externus profundus/medialis
complex was the dominant muscle of the dorsotemporal
fossa whereas m. pseudotemporalis superficialis was lim-
ited to the caudal surface of the laterosphenoid. How-
ever, if one considers that ratites (e.g., Struthio,
Eudromia) may represent the primitive avian condition,
rather than a secondarily derived clade, then one would
infer that m. pseudotemporalis superficialis was the
dominant temporal muscle among, at least, theropod
dinosaurs. Clear data that indicate shifts in muscle
attachments among different clades of non-avian dino-
saurs, such as the envisioned muscle shifts between ba-
sal and derived ceratopsians and ornithopods, would
certainly benefit systematic analyses.
Inferences of jaw muscles reciprocally illuminate infer-
ences of other cranial soft tissues. As they exit the brain-
case, the mandibular, and more often, the maxillary
nerves excavate portions of the laterosphenoid and
prootic. It is assumed that non-avian dinosaur jaw
muscles and nerves follow the same topological patterns
as other sauropsids, thus if a groove for the maxillary
1263
nerve is identified on the laterosphenoid, marking its
course rostrally, it can be inferred that m. pseudotem-
poralis superficialis, which lies medial to the nerve was
small or attached rostrally on the braincase. In hadro-
saurs, the maxillomandibular nerves exit directly later-
ally and a bit caudally, often leaving a shallow groove or
lip on the prootic (Figs. 2 and 5). This suggests that m.
pseudotemporalis superficialis did attach to the rostral
portion of the dorsotemporal fossa even though there are
no direct correlates that indicate the muscle’s attach-
ment. Therefore, given adequate knowledge of jaw mus-
cle anatomy and using neurovascular topological rules, a
portion of the other soft tissues of the adductor chamber
can be reconstructed with relative confidence. Interest-
ingly, in hadrosaurs, when the temporal muscles are
reconstructed from the temporal fossa to the coronoid
process, around the laterosphenoid buttress, the muscles
pass through much of the orbit. Therefore, interpreta-
tions of muscle anatomy may directly impact inferences
of eyeball size. Thus, as in other vertebrates, orbit size
may not be a complete index of eyeball size (Ross and
Kirk, 2007). These packing issues can be further
explored not only in the orbit, but also the nasal cavity,
pharynx, and middle ear cavities, which are bounded by
large portions of the pterygoideus dorsalis and ventralis
muscles, and protractor muscles, respectively.
Finally, feeding behavior and connective tissue adapt-
ive plasticity are major factors involved in the structure
and function of jaw muscles and the skull (Ravosa et al.,
2007). These phenomena make interpreting jaw muscle
functional anatomy extremely difficult over ontogenies of
animals and may even manifest themselves among dif-
ferent populations of the same taxon (Erickson et al.,
2004). These anatomical features must be well-under-
stood before the testing of functional and systematic
hypotheses. The best solution to investigating feeding
behaviors, such as chewing or bite force, among non-
avian dinosaurs may be to test hypotheses using gross
as well as more exact muscle inferences (i.e., ‘‘temporal’’
muscles versus specific muscle bellies) while also testing
a variety of virtual physiological cross-sectional areas,
recruitment levels, and other behavioral or biomechani-
cal parameters (e.g., Rayfield, 2005; Wroe et al., 2005;
Rayfield and Milner, 2008). Anatomical inferences can
only form a general framework for functional analysis
when dealing with soft tissues such as jaw muscles.
In conclusion, jaw muscles offer a variety of bony
structures that support inferences of their reconstruction
in fossil taxa such as non-avian dinosaurs. Even individ-
uals of the same taxon may have more or fewer osteolog-
ical correlates that support one muscle’s inference
versus another. When anatomy does not offer necessary
insight for soft-tissue reconstruction, phylogenetic brack-
eting may lend inferential support. Among the muscles
described above, the orbitotemporal muscles have the
weakest phylogenetic support, but generally have the
strongest anatomical support: m. levator pterygoideus is
a Level III inference, but has excellent osteological cor-
relates that support its reconstruction. On the other
hand, temporal muscles such as m. adductor mandibulae
externus medialis or m. pseudotemporalis superficialis
have strong phylogenetic support, and it is likely all fos-
sil reptiles had the muscle bellies, but anatomical struc-
tures that explicitly support identifications of their
attachments are rare. It is relatively easy and
1264 HOLLIDAY
straightforward to make conservative interpretations of
jaw muscles and caution is suggested when using muscle
data in evolutionary and functional analyses. Regard-
less, jaw muscles are a critical component to under-
standing head anatomy, function, and evolution in non-
avian dinosaurs.
ACKNOWLEDGMENTS
Many thanks to Lawrence Witmer, Natalia Rybczyn-
ski, David Dufeau, Tobin Hieronymus, Ryan Ridgely,
Emily Rayfield, Mark Young, and others for providing
advice and assistance during the development of this
project. Thanks to numerous staff and curators at muse-
ums for access to specimens. Comments from Peter Dod-
son and two anonymous reviewers greatly improved the
manuscript.
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