JOURNAL OF MORPHOLOGY 265:102–119 (2005)
Morphology and Evolution of the Jaw Suspension in
Lamniform Sharks
C.D. Wilga*
Department of Biological Sciences, University of Rhode Island, Kingston, Rhode Island 02881
ABSTRACT The morphology of the jaw suspension and
jaw protrusion mechanism in lamniform sharks is de-
scribed and mapped onto a cladogram to investigate how
changes in jaw suspension and protrusion have evolved.
This has revealed that several evolutionary modifications
in the musculoskeletal apparatus of the jaws have taken
place among lamniform sharks. Galeomorph sharks (Car-
charhiniformes, Lamniformes, Orectolobiformes, and Het-
erodontiformes) have paired ethmopalatine ligaments
connecting the ethmoid process of the upper jaw to the
ethmoid region of the cranium. Basal lamniform sharks
also acquired a novel single palatonasal ligament connect-
ing the symphysis of the upper jaw to the cranium mid-
ventral to the nasal capsule. Sharks in the family Lamni-
dae subsequently lost the original paired ethmopalatine
ligament while retaining the novel palatonasal ligament.
Thus, basal lamniform taxa (Mitsukurina owstoni, Car-
charius taurus, Alopias vulpinnis) have increased liga-
mentous support of the lateral region of the upper jaw
while derived species (Lamnidae) have lost this lateral
support but gained anterior support. In previous studies
the morphology of the jaw suspension has been shown to
play a major role in the mechanism of upper jaw protru-
sion in elasmobranchs. The preorbitalis is the primary
muscle effecting upper jaw protrusion in squalean (sister
group to galeomorphs) and carcharhiniform (sister group
to lamniforms) sharks. The preorbitalis originates from
the quadratomandibularis muscle and inserts onto the
nasal capsule in squalean and carcharhiniform sharks.
Carcharhiniform sharks have evolved a subdivided preor-
bitalis muscle with the new division inserting near the
ethmoid process of the palatoquadrate (upper jaw). Alopid
sharks have also independently evolved a partially subdi-
vided preorbitalis with the new division inserting at the
base of the ethmoid process and surrounding connective
tissue. Lamnid sharks have retained the two preorbitalis
divisions but have modified both of the insertion points.
The original ventral preorbitalis division now inserts onto
the connective tissue surrounding the mid-region of the
upper jaw, while the new dorsal preorbitalis division in-
serts onto the surrounding connective tissue and skin at a
more posterior position on the upper jaw. The retractor
muscle of the jaws, the levator hyomandibularis, has also
been modified during the evolution of lamniform sharks.
In most sharks, including basal lamniforms, the levator
hyomandibularis inserts onto the hyomandibula and func-
tions to retract the jaws after protrusion. In alopid and
lamnid sharks the levator hyomandibularis inserts pri-
marily onto the upper and lower jaws around the jaw joint
and is a more direct route for retracting the jaws. Thus,
there has been at least one instance of character loss
(ethmopalatine ligament), acquisition (palatonasal liga-
ment), subdivision (preorbitalis), and modification (ven-
© 2005 WILEY-LISS, INC.
tral preorbitalis, dorsal preorbitalis, and levator hyoman-
dibularis) in the ligaments and muscles associated with
the jaw suspension and jaw protrusion mechanism in lam-
niform sharks. While derived lamniform sharks (Lamna
nasus, Carcharodon carcharius, and Isurus oxyrinchus)
lost the ancestral passive lateral support of the ethmoid
articulation of the upper jaw, they simultaneously ac-
quired muscular support by way of the levator hyoman-
dibularis, which provides a dynamic mechanism for lat-
eral support. The evolution of multiple divisions of
preorbitalis insertions onto the palatoquadrate and mod-
ification of the levator hyomandibularis insertion directly
onto the jaws provides an active mechanism for multiple
protractions and retractions of the upper jaw, which is
advantageous in those sharks that gouge or saw pieces
from large oversized prey items. J. Morphol. 265:102–119,
2005. © 2005 Wiley-Liss, Inc.
KEY WORDS: anatomy; jaw protrusion; Mitsukurina;
Carcharius; Alopias; Carcharodon; Isurus; Lamna
Understanding the sequence of modification in
musculoskeletal systems over the course of evolu-
tion is a central theme in functional morphology and
biomechanics. The upper jaw protrusion mechanism
in elasmobranchs provides an ideal system for
studying the evolution of a musculoskeletal system
because it is a central component of the feeding
mechanism in most sharks and rays. Upper jaw
protrusion in elasmobranchs occurs when the pala-
toquadrate cartilage (upper jaw) moves ventrally
away from the cranium while the lower jaw elevates
to surround the prey. This is in contrast to that of
bony fishes in which upper jaw protrusion is coupled
to lower jaw depression. Numerous functions for
protrusion of the upper jaw have been proposed:
more efficient biting of prey, gouging of the upper
jaw into large prey, a versatile yet hydrodynamic
subterminal mouth, reorienting of the teeth for in-
creased grasping ability, simultaneous closing of the
*Correspondence to: Cheryl Wilga, Department of Biological Sci-
ences, University of Rhode Island, 100 Flagg Road, Kingston, RI
02881-0816. E-mail: cwilga@uri.edu
Published online 6 May 2005 in
Wiley InterScience (www.interscience.wiley.com)
DOI: 10.1002/jmor.10342
 JAW MORPHOLOGY OF LAMNIFORM SHARKS 103

Fig. 1. Representative jaw suspension types in extant elasmobranchs. Amphistyly as in sevengill sharks, Notorynchus cepedianus
(after Daniel, 1922), orbitostyly as in spiny dogfish Squalus acanthias (after Wilga and Motta, 1998a), hyostyly as in carcharhiniform
sharks, lemon Negaprion brevirostris (after Motta and Wilga, 1995) and bonnethead Sphyrna tiburo, euhyostyly as in batoids, Atlantic
guitarfish Rhinobatos lentiginosus (after Wilga and Motta, 1998b). CH, ceratohyal (ventral hyoid element); CR, cranial; ET, ethmo-
palatine articulation; EM, ethmoid process; HM, hyomandibula (dorsal hyoid element); HY, hyoid (composed of hyomandibula and
ceratohyal); MC, Meckel’s cartilage (lower jaw); OP, orbital process; OR, orbital articulation; PO, postorbital articulation; PQ,
palatoquadrate (upper jaw); UJ, upper jaw. black, ethmopalatine and postorbital ligaments; dark gray, hyomandibula; light gray,
ceratohyal; white, cranium, palatoquadrate, and Meckel’s cartilage.

upper and lower jaws, and decreasing the time to
jaw closure (Springer, 1961; Alexander, 1967; Moss,
1972, 1977; Tricas and McCosker, 1984; Frazzetta
and Prange, 1987; Frazzetta, 1994). Only one func-
tion for upper jaw protrusion has been supported
experimentally, that of significantly reducing the
time to jaw closure (Motta and Wilga, 1995; Wilga,
2002).
The morphology of the jaw suspension has been
found to play a major role in determining the extent
of upper jaw protrusion in elasmobranchs (Wilga,
2002). The type of jaw suspension allows some pre-
dictions to be made about the mobility of the man-
dibular arch as a whole on the cranium, but it is the
extent of the ligamentous and muscular connections
between the palatoquadrate and the cranium that
determines the extent that the upper jaw can pro-
trude. Several jaw suspension types are found
within extant chondrichthyans (Wilga, 2002) (Fig.
1). In all extant sharks the hyomandibula links the
jaws to the cranium, and thus is always functionally
suspensory, and therefore hyostylic, to some degree
(Maisey, 1980, 1984). These jaw suspension types
are defined by the amount of support that the hyo-
mandibular element of the hyoid arch gives to the
mandibular arch and whether it is ancestral or sec-
ondarily derived (Huxley, 1876; Gregory, 1904), the
state (intact or broken up) of the hyoid arch ele-
ments (hyomandibula and ceratohyal) (Gregory,
1904), and the presence of postorbital (Goodrich,
1909), orbital, or ethmoid articulations (Maisey,
1980).
Inclusion of the morphological state of the palato-
quadrate articulations and hyoid arch has clarified
our understanding of the jaw suspension types, both
phylogenetically and functionally (Wilga, 2002). Ho-
lostyly is characterized as fusion of the palatoquad-
rate to the cranium with a nonsuspensorial intact
hyoid arch (Holocephali) (Huxley, 1876; Gregory,
1904). In contrast, fusion of the palatoquadrate to
the cranium with a nonsuspensorial modified hyoid
arch (hyoid arch elements broken up and intercon-
nected by other skeletal elements) is retained in
autostyly (Dipnoi, Tetrapoda) (Huxley, 1876; Greg-
ory, 1904). A palatoquadrate with a postorbital ar-
ticulation that is independent of and suspended
from the cranium primarily by ligaments anteriorly
and by a hyomandibula posteriorly, which is pre-
sumed to contribute little to jaw support, is said to
be amphistylic (Xenacanthida, Palaeospinax, Hex-
anchiformes) (Huxley, 1876; Gregory, 1904). Amphi-
stylic sharks not only have a postorbital articula-
tion, but they also have an orbital process, which
also makes them orbitostylic (see below) (Parker,
1878; Gregory, 1904; Goodrich, 1909). Correspond-
ingly, in hyostyly, an ethmoid articulation is present
and the hyomandibula is thought to contribute more to
jaw support than the anterior ligaments (Hybodon-
tida, Heterodontiformes, Carcharhiniformes, Lamni-
formes) (Huxley, 1876; Gregory, 1904). In orbitostyly,
an orbital process articulates with the orbital wall
while the hyomandibula contributes significantly to
the support of the jaws (Chlamydoselachiformes,
Hexanchiformes, Squaliformes) (Maisey, 1980,
1984). Note that only squalean sharks possess an
orbital process, which is similar to but developmen-
tally distinct from the ethmoid process of galean
sharks (Maisey, 1980). Batoids are euhyostylic
(truly hyostylic), in which the only articulation of
104 C.D. WILGA
the palatoquadrate and cranium is with the hyo-
mandibula (Huxley, 1876; Gregory, 1904).
Most lamniform sharks are large, oceanic swim-
mers that must continuously swim in order to ven-
tilate the gills. These characteristics make them
difficult to study experimentally in the laboratory.
Probably the most thrilling event in the feeding
behavior of white, Carcharodon carcharias, and
sand tiger, Carcharius taurus, sharks is the protru-
sion of the upper jaw a remarkable distance from the
cranium. Experimental support is hardly necessary
to convince us that these species are capable of ex-
tensive protrusion of the upper jaw. However, there
are some lamniform species that have only moderate
protrusion, such as thresher, Alopias vulpinnis, and
porbeagle, Lamna nasus, sharks. Previous findings
on the musculoskeletal elements of the jaw appara-
tus (Wilga, 2002; Wilga et al., 2002) can be used to
formulate several predictions regarding the function
of the jaw suspension and upper jaw protrusion in
sharks.
The goals of this study are to: 1) describe the
morphology of the jaw suspension and jaw protru-
sion mechanism in representative species from all
major groups of lamniform sharks; 2) to map these
characters onto a cladogram to hypothesize how
changes in jaw suspension and protrusion have Fig. 2. Lamniform phylogeny after Martin (1995) and Shirai
(1996) with Carcharhiniformes (Galea) as the outgroup. Black
evolved within lamniform sharks; and 3) compare circle indicates sharks in the Order Lamniformes. Species of
these findings to previous research to evaluate the specimens dissected are indicated.
evolution of jaw suspension and protrusion in elas-
mobranchs.
rectly related to the jaw suspension, other muscles of interest
such as the hypobranchial muscles are described here that may
MATERIALS AND METHODS not be illustrated.
Specimens
Six species representing the three major and one of the four Character Mapping
minor families of lamniform sharks were used in this study (Fig.
2) (goblin, Mitsukurinidae, one of one species; sand tiger, Carcha- Morphological characteristics of the jaw suspension and upper
ridae, one of four species; thresher, Alopidae, one of three species; jaw from the six lamniform species were mapped onto an existing
mackeral sharks, Lamnidae, three of five species). Representa- cladogram by Shirai (1996) to better understand how the mor-
tives from the remaining minor lamniform families were not phological characters may have evolved. Note that these charac-
available (crocodile, Pseudocarcharidae; megamouth, Megachas- ters were not used to build the cladogram (Shirai, 1996); rather,
midae; basking, Cetorhinidae, each contains only one species). an existing cladogram was used to investigate how the morphol-
Seven specimens were collected from local shark tournaments ogy of the jaw suspension may have evolved. The methods of
and dissected fresh: two female Alopias vulpinnis 150 and 200 cm Wiley et al. (1991) were used to infer morphological states at the
FL (fork length); four Isurus oxyrinchus ranging from 156 to 181 nodes. The outgroup for the lamnid comparison is the sister group
cm FL; and one Lamna nasus NMFS #40 (ceratohyal removed). Carcharhiniformes, from which several species have been studied
The following preserved specimens were dissected at the Museum in relevant detail: Negaprion brevirostris, Sphyrna tiburo, Car-
of Comparative Zoology at Harvard University: Carcharius tau- charhinus limbatus, C. acronotus, C. plumbeus (Motta and Wilga,
rus MCZ 164335; Carcharodon carcharius MCZ 36470 male; and 1995; Wilga and Motta, 2000; unpubl. data). The outgroup for the
Mitsukurina owstoni MCZ 16073. elasmobranch comparison is the sister group Holocephali, from
which the species Hydrolagus collei has been studied in relevant
detail by the author and others (Edgeworth, 1935; Didier, 1995).
Anatomical Description
Illustrations were prepared by photographing dissected speci- RESULTS
mens and tracing the outlines of muscles and cartilages. The
muscle fiber direction was sketched in by hand. The nomencla-
Jaw Suspension
ture for the chondrocranium, connective tissue, and musculature Ethmopalatine ligament. The more basal spe-
follows that of Wilga (2002) and Motta and Wilga (1995). Muscu- cies, Mitsukurina owstoni, Carcharius taurus, and
lar anatomy of taxa not dissected by the author relies on the Alopias vulpinnis have two ethmopalatine liga-
works of Gegenbauer (1865), Marion (1905), Luther (1909a,b),
Goodey (1910), Allis (1913, 1914, 1923), Daniel (1915, 1922),
ments, one on each side of the palatoquadrate sym-
Edgeworth (1935), Lightoller (1939), Moss (1972), Nobiling physis, while the more derived species, Lamna nasus,
(1977), Gadow (1988), and Compagno (1988). Although not di- Carcharodon carcharius, and Isurus oxyrinchus, lack
 JAW MORPHOLOGY OF LAMNIFORM SHARKS
them (Fig. 3). The ethmopalatine ligament is solid
and rope-like in all of the lamniform specimens ex-
amined here; however, the morphology of the eth-
moid process differs among them. In M. owstoni a
small ethmoid process projects dorsally from the
mid-dorsal surface of the palatoquadrate cartilage,
from which an ethmopalatine ligament extends to
the suborbital plate at the anterior edge of the eye
(Fig. 3). In C. taurus the ethmoid process is large
and projects dorsally into the anterior end of the
ventromedial surface of the orbit. The ethmoid pro-
cess has a wide cartilaginous base from which the
ethmopalatine ligament arises. The ligament ex-
tends first dorsally, then anteriorly to insert onto
the ethmoid cartilage posterior to the nasal capsule.
In A. vulpinnis the ethmoid process is of moderate
size and also has a cartilaginous base capped by a
ligamentous flap that extends from the base of the
process to its attachment in the ethmopalatine
groove of the orbit (Fig. 3B). This is very similar to
the condition observed in lemon sharks Negaprion
brevirostris (Carcharhiniformes) and other carchar-
hiniform sharks, which have solid rope-like ethmo-
palatine ligaments (Motta and Wilga, 1995; Wilga,
2002). The shark species examined from the more
distantly related shark orders, Orectolobiformes,
Heterodontiformes, and Squaliformes, have hollow
sleeve-like ligaments that fold over the ethmoid or
orbital process when the upper jaw is retracted
(Wilga, 2002).
Palatonasal ligament. A single solid palatona-
sal ligament emanates from the mid-ventral surface
of the cranium between the nasal capsules and ex-
tends to the symphysis of the palatoquadrate in all
of the species (Fig. 3). In Mitsukurina owstoni the
ligament is thin, solid, and strap-like (Fig. 3a). In
Carcharius taurus the palatonasal ligament is solid,
rope-like, and emanates from a low flat cartilagi-
nous base. In Alopias vulpinnis, Lamna nasus, Car-
charodon carcharius, and Isurus oxyrinchus the pal-
atonasal ligament is also large and rope-like and
contains a small vertically oriented rod-like carti-
lage extending four centimeters from the palato-
quadrate symphysis (Fig. 3B–D). The rod-like carti-
lage is embedded within the ligament and therefore
not visible in the figures. In L. nasus there is a small
process on the dorsal surface of the palatoquadrate
on either side of the symphysis from which the small
rod-like cartilage emanates. When the jaw is in the
retracted or resting position, the rod-like cartilage is
flipped forward over the dorsal surface of the pala-
toquadrate. When the jaw is manually protracted to
simulate upper jaw protrusion, the rod-like cartilage
is pulled vertically upright between the cranium and
palatoquadrate.
Axial Muscle
Epaxialis. The epaxialis inserts onto the poste-
rior and dorsal surface of the otic capsule (Fig.
105
4D,F). The epaxialis splits to circle around the pari-
etal fossa and then extends anteriorly onto the cra-
nium at various distances among the species. In
Mitsukurina owstoni, Carcharius taurus, and Isurus
oxyrinchus the epaxialis inserts as far anterior as
the postorbital process of the cranium. In Alopias
vulpinnis the epaxialis is massive and extends an-
teriorly as far as the anterior edge of the eye. In
Lamna nasus and Carcharodon carcharius the ep-
axialis extends anteriorly nearly to the mid-orbital
region. In C. carcharius the epaxialis also inserts
onto the postorbital process as well as in the region
between the hyomandibula and orbit (not visible in
lateral view).
Muscles of the Mandibular Muscle Plate
Levator palatoquadrati. The levator palato-
quadrati is vertically oriented and lies between the
postorbital process and the levator hyomandibularis
in all of the lamniform species examined (Fig. 4).
The origin may be from the lateral surface of the otic
capsule such as in Mitsukurina owstoni and Alopias
vulpinnis, from the dorsolateral edges of the epax-
ialis as in Carcharius taurus, Carcharodon car-
charius, and Isurus oxyrinchus, or from both as in
Lamna nasus. In A. vulpinnis, C. carcharius, and L.
nasus the levator palatoquadrati also originates
from the posterior surface of the postorbital process.
The levator palatoquadrati extends ventrally be-
hind the orbit to insert onto the ascending process of
the palatoquadrate, although details of the specific
region and area of the insertion varies among the
species. Unlike the other lamniforms described here,
the insertion of the levator palatoquadrati in Mit-
sukurina owstoni is relatively small. It extends ven-
trally over the hyomandibula to insert onto the dor-
sal surface of the palatoquadrate just anterior to and
on a small region of the ascending process (Fig. 4A).
The levator palatoquadrati is massive in Carcharius
taurus, with a broad insertion on the dorsal and
dorsolateral edges of the anterior half of the ascend-
ing processes of the palatoquadrate (Fig. 4B). In
Alopias vulpinnis the levator palatoquadrati is of
moderate size and inserts onto the dorsal and dor-
somedial surfaces of the ascending and anterior
crest regions of the ascending process of the palato-
quadrate (Fig. 4C). It does not extend past the crest
nor does it insert anterior to the ascending process.
In Lamna nasus the levator palatoquadrati inserts
onto the dorsal surface of the palatoquadrate imme-
diately anterior to as well as on the ascending pro-
cess (Fig. 4D). In Carcharodon carcharius the leva-
tor palatoquadrati inserts onto the dorsal surface of
the palatoquadrate immediately anterior to and
onto the ascending process up to the crest (Fig. 4E).
Here the insertion narrows diagonally to insert only
onto the dorsomedial surface of crest. In Isurus
oxyrinchus the levator palatoquadrati inserts onto
106 C.D. WILGA

Fig. 3. Representative jaw suspension types in lamniform sharks. A: Goblin, Mitsukurina owstoni. B: Thresher, Alopias vulpinnis
(jaw suspension similar to sandtiger Carcharius taurus). C: Porbeagle, Lamna nasus. D: Mako, Isurus oxyrinchus (jaw suspension
similar to white Carcharodon carcharius). CR, cranium; EP, ethmopalatine ligament; HY, hyomandibula; MC, Meckel’s cartilage; NC,
nasal capsule; OP, ethmoid process; PN, palatonasal ligament; PQ, palatoquadrate or upper jaw; RO, rostrum.
JAW MORPHOLOGY OF LAMNIFORM SHARKS 107

Figure 3 (continued)
108 C.D. WILGA

Fig. 4. Lateral view of superficial muscles in lamniform sharks. A: Goblin, Mitsukurina owstoni. B: Sandtiger, Carcharius taurus.
C: Thresher, Alopias vulpinnis. D: Porbeagle, Lamna nasus. E: White, Carcharodon carcharius. F: Mako, Isurus oxyrinchus. BC,
branchial constrictors; CR, cranium; CY, ceratohyal; EP, epaxialis; EY, eye; HY, hyomandibula; IM, intermandibularis; LH, levator
hyomandibularis; LP, levator palatoquadrati; MC, Meckel’s cartilage; NC, nasal capsule; POD, dorsal preorbitalis; PO, preorbitalis
(homologous to POV); POV, ventral preorbitalis; PQ, palatoquadrate; PQP, ascending process of palatoquadrate; QMD, dorsal
quadratomandibularis superficial division; QMDm, dorsal quadratomandibularis medial division; QMV, ventral quadratomandibu-
laris; RO, rostrum.

the anterior portion of the ascending process of the
palatoquadrate (Fig. 4F).
Mid-lateral raphe of the quadratomandibu-
laris. The mid-lateral raphe is a broad tendinous
sheet that separates and provides attachment points
for the various divisions of the quadratomandibu-
laris (Fig. 4). In Mitsukurina owstoni and Car-
charius taurus the mid-lateral raphe extends poste-
riorly and medially from the anterior jaw joint to the
posterior jaw joint to separate the dorsal and ventral
divisions of the quadratomandibularis (Fig. 4A,B).
In Alopias vulpinnis and Lamna nasus the superfi-
cial portion of the raphe extends posteriorly from the
connective tissue at corner of the mouth but fans out
medially diagonally and posterodorsally to the re-
gion of the jaw joint (Fig. 4C,D). In Carcharodon
JAW MORPHOLOGY OF LAMNIFORM SHARKS 109

Figure 4 (continued)
110 C.D. WILGA

Figure 4 (continued)
 JAW MORPHOLOGY OF LAMNIFORM SHARKS
carcharius and Isurus oxyrinchus the raphe is a thin
broad connective tissue sheet that extends superfi-
cially dorsoposteriorly from the connective tissue at
the corner of the mouth to the palatoquadrate about
one-third of the way from its posterior end. The
raphe is not apparent superficially in C. carcharius.
The orientation of the raphe rotates medially from
more transverse at the corner of the mouth to more
sagittal at the palatoquadrate end to separate the
dorsal and ventral quadratomandibularis.
Preorbitalis. The posterior end of the preor-
bitalis muscle originates from the anterior surface of
the raphe at the corner of the mouth in all of the
lamniform species dissected here and from the skin
laterally in Isurus oxyrinchus (Fig. 4). Mitsukurina
owstoni, Carcharius taurus, and Carcharodon car-
charius possess only the original ventral preorbitalis
muscle. Alopias vulpinnis, Lamna nasus, and I.
oxyrinchus have two divisions of the preorbitalis, the
original ventral head and a new dorsal head that are
distinct from each other and the quadratomandibu-
laris at the insertion end (Fig. 4C,D,F). In C. taurus,
L. nasus, C. carcharius, and I. oxyrinchus the ven-
tral preorbitalis muscle extends anterodorsally to
insert onto the palatoquadrate-mandibular connec-
tive tissue sheath that overlies the teeth and jaws at
the anterior region of the palatoquadrate (Fig.
4B,D–F). In M. owstoni and A. vulpinnis the preor-
bitalis inserts onto the posteroventral surface of the
nasal capsule and the overlying skin. In A. vulpinnis
a second, smaller medial division of the preorbitalis
inserts onto the connective tissue overlying the jaws
and the base of the orbital process. In L. nasus and
I. oxyrinchus the dorsal preorbitalis extends anteri-
orly to insert onto the skin just anterior to the qua-
dratomandibularis muscle mass (Fig. 4D,F).
Quadratomandibularis dorsal superficial di-
vision. The superficial division of the dorsal qua-
dratomandibularis originates from the dorsal sur-
face of the mid-lateral raphe and fans out
anterodorsally to insert along the lateral edges of
the ascending process of the palatoquadrate (Fig. 4).
Mitsukurina owstoni has a relatively simple, undif-
ferentiated dorsal quadratomandibularis that is
most similar to the superficial division of the dorsal
quadratomandibularis of the remaining species dis-
cussed here (Fig. 4). In Carcharodon carcharius the
superficial fibers extend from the palatoquadrate
directly to the mandible without an intervening ra-
phe (Fig. 4E). The mandibular nerve partially sepa-
rates the superficial and medial divisions of the dor-
sal quadratomandibularis as in carcharhiniform
species (Motta and Wilga, 1995).
Quadratomandibularis dorsal medial divi-
sion. In Carcharius taurus the medial division of
the dorsal quadratomandibularis is very large and
originates from the mid-lateral raphe (Fig. 4B). It
inserts onto the anterodorsal edge of the ascending
process between the superficial quadratomandibu-
laris division and the preorbitalis. In Alopias vulp-
111
innis, Lamna nasus, Carcharodon carcharius, and
Isurus oxyrinchus the medial division of the quadra-
tomandibularis dorsal originates from the mid-
lateral raphe and fans out dorsally to insert onto the
lateral surface of the palatoquadrate ventral to the
superficial division. In L. nasus, C. taurus, C. car-
charius, and I. oxyrinchus the medial division ex-
tends more anterior than the superficial division to
insert onto the palatoquadrate anterior to the as-
cending process up to the insertion of the levator
palatoquadrati. However, the preorbitalis muscle
completely overlies it in L. nasus, C. carcharius, and
I. oxyrinchus, but not in C. taurus, where it is clearly
visible in superficial views.
Quadratomandibularis dorsal deep division.
The deep division is a small triangular division that
is located in the angle of the jaws in all of the
lamniform species dissected here. It originates from
a shallow shelf of cartilage on the mandible in the
corner of the mouth and extends dorsally to the
palatoquadrate under the rim at the jaw joint.
Quadratomandibularis ventral. The quadrato-
mandibularis ventral originates from the ventral
surface of the mid-lateral raphe and fans out to
insert along the mandible approximately as far an-
terior as the dorsal division of the quadratoman-
dibularis in all of the species studied here (Fig. 4). In
Lamna nasus a few superficial fibers pass over the
raphe to merge directly with the superficial fibers of
the quadratomandibularis dorsal (Fig. 4D). Also in
L. nasus, some fibers from the dorsal edge of the
palatoquadrate anterior to the jaw joint extend di-
rectly posterior to insert onto the mandible just pos-
terior to the jaw joint. In Carcharodon carcharius
and Isurus oxyrinchus some superficial fibers of the
quadratomandibularis ventral merge with superfi-
cial fibers of the quadratomandibularis dorsal (Fig.
4E,F). These fibers extend between the outermost
lateral edges of the ascending process to the lateral
edges of the posterior region of the mandible. In C.
carcharius a deep ventral quadratomandibularis di-
vision originates from the mid-lateral raphe to in-
sert onto the lateral fossa of the ascending process
and lies between the mandibular nerve and the
palatoquadrate.
Intermandibularis (not shown). The inter-
mandibularis extends from a mid-ventral raphe and
inserts onto the outer edges of the quadratoman-
dibularis ventral and mandible. In Isurus oxyrin-
chus the intermandibularis also extends between
the lateral edges of the palatoquadrate. No distinct
mid-ventral raphe is present in Carcharias taurus
or Lamna nasus. In L. nasus the insertion is broad
anteriorly and extends onto the ventral surface of a
large process on either side of the symphysis of the
mandible.
Intermandibular plate (not shown). A pair of
cartilaginous elements are attached to the mid-
ventral raphe between the mandibles, of which the
morphology varies greatly among the species. The
112 C.D. WILGA
intermandibular plates are located medial to the
ventral edge of the mandible and anterior to the
insertion of the quadratomandibularis. The inter-
mandibular plates are located deep to the interman-
dibularis and overlap when the jaws are retracted.
In Mitsukurina owstoni there is a large interman-
dibular plate consisting of a triangular-shaped car-
tilage with the apex pointed posteriorly. In Car-
charius taurus the intermandibular plate is more
teardrop-shaped with the apex directed anteriorly.
In Alopias vulpinnis the intermandibular plate is
small and lies near the edge of the mandible. In
Lamna nasus the intermandibular plate is small
and rod-like and is located at the posteroventral
edge of the coracomandibularis insertion onto the
mandible. In Carcharodon carcharius the interman-
dibular plate is also small and rod-like and is located
posterior to the process of the mandibular symphy-
sis.
Muscles of the Hyoid Muscle Plate
Levator hyomandibularis. The levator hyo-
mandibularis originates from the dorsolateral edge
of the epaxialis and extends ventrally to various
insertions among the species (Fig. 4). In Mitsuku-
rina owstoni the levator hyomandibularis inserts
onto the posterodorsal edge of the hyomandibula
(Fig. 4A). In Alopias vulpinnis and Lamna nasus the
levator hyomandibularis also originates from the
otic crest (Fig. 4C,D). In Carcharius taurus the le-
vator hyomandibularis is very thick and inserts onto
the entire dorsal surface of the hyomandibula (Fig.
4B). In C. taurus the anterior edge of the levator
hyomandibularis is overlapped by the levator pala-
toquadrate.
In some species the levator hyomandibularis has
multiple insertions onto the palatoquadrate and
hyomandibula, with some fibers inserting onto the
mandible and jaw joint, and a nonfunctional spi-
racular canal anterior to its insertion (Fig. 4). In
Alopias vulpinnis, Carcharodon carcharius, and Isu-
rus oxyrinchus the bulk of the fibers insert onto the
dorsal surface of the posterior third of the palato-
quadrate, while in Lamna nasus only superficial
fibers do (Fig. 4C–F). In A. vulpinnis, C. carcharius,
and I. oxyrinchus some of the medial fibers insert
onto a dorsal ridge along the anterior surface of the
hyomandibula as well as on the dorsal surface of its
proximal end. In L. nasus the bulk of the fibers
insert onto the dorsal surface of a ridge along the
anterior edge of the distal three-quarters of the hyo-
mandibula. In A. vulpinnis, L. nasus, C. carcharius,
and I. oxyrinchus some fibers insert onto the dorsal
surface of the mandible adjacent to the jaw joint as
well as the capsule of the jaw joint.
Interhyoideus (not shown). The interhyoideus
extends from the ceratohyal and posterior edge of
the mandible to the mid-ventral raphe. It lies deep
to the intermandibularis and begins just posterior to
the intermandibularis plate and ends with Meckel’s
cartilage.
Muscles of the Hypobranchial Muscle Plate
Coracomandibularis (not shown). The origin
of the coracomandibularis was present in only two of
the species. In Alopias vulpinnis the coracoman-
dibularis is very long and narrow and originates
from the coracoid bar. In Isurus oxyrinchus the cora-
comandibularis originates from the anteroventral
end of the coracoarcualis in the mid-region of the
coracoid bar, posterior to the coracohyoideus. It does
not originate from the coracoid bar. In Carcharius
taurus, A. vulpinnis, Lamna nasus, Carcharodon
carcharius, and I. oxyrinchus the coracomandibu-
laris extends anteriorly and splits just prior to the
insertion onto the posterior surface of Meckel’s car-
tilage on a large process on either side of the sym-
physis. A coracomandibularis was not present or
was indistinct from the intermandibularis in Mit-
sukurina owstoni.
Coracohyoideus (not shown). The origin of the
coracohyoideus was present in only four of the spe-
cies. In Mitsukurina owstoni, Carcharodon car-
charius, and Isurus oxyrinchus the coracohyoideus
originates from the ventral surface of the anterior
end of the coracoarcualis. In I. oxyrinchus the cora-
cohyoideus is much larger than the coracoarcualis.
In Alopias vulpinnis the coracohyoideus originates
primarily from the first basibranchial, hypo-
branchial, and ceratobranchial, with few fibers orig-
inating from the ventral surface of the anterior end
of the coracoarcualis as in most sharks. In all of the
species the coracohyoideus extends anteriorly to in-
sert onto the ventral surface of the basihyal.
Coracoarcualis (not shown). The origin of the
coracoarcualis was present in only four of the spe-
cies. In Mitsukurina owstoni, Carcharodon car-
charius, and Isurus oxyrinchus the coracoarcualis
originates from the anterodorsal edge of the coracoid
bar and inserts onto the dorsal surface of the poste-
rior end of the coracohyoideus. In Alopias vulpinnis
the coracoarcualis originates from the hypaxialis
musculature and the mid-ventral surface of the cor-
acoid bar (the pericardium is attached to the ante-
rior surface). The origin of the coracoarcualis over-
lies the hypaxialis, which inserts onto the posterior
surface of the coracoid bar.
DISCUSSION
Evolution of the Jaw Suspension in
Lamniformes
Anatomical dissection of the ligaments and mus-
culoskeletal apparatus of the jaw suspension of six
lamniform species from the four major groups has
revealed that several morphological modifications
have occurred during the evolution of lamniform
sharks (Fig. 5). Several ligamentous and musculo-
 JAW MORPHOLOGY OF LAMNIFORM SHARKS
Fig. 5. Evolution of the ligamentous and musculoskeletal el-
ements of the upper jaw in lamniform sharks, with Carcharhini-
formes as an outgroup. Phylogeny after Martin (1995) and Shirai
(1996).
skeletal elements connect the upper jaw to the cra-
nium and therefore contribute to support of the
jaws. Basal lamniform species have paired rope-like
ethmopalatine ligaments connecting the ethmoid
process of the upper jaw to the ethmoid region of the
cranium (Figs. 3A,B, 5-1), similar to their sister
group the Carcharhiniformes (Compagno, 1988;
Motta and Wilga, 1995; Wilga et al., 2002; Wilga,
2002). In addition, all lamniform species acquired a
novel single rope-like palatonasal ligament attach-
ing the symphysis of the upper jaw to the nasal
region of the cranium (Figs. 3, 5-2). Inside the novel
ligament of alopid and lamnid species there is a
small rod-like cartilage that bends anteriorly over
the upper jaw when retracted and is pulled verti-
cally during upper jaw protrusion (Fig. 5-3). The
rod-like palatonasal cartilage and the interman-
dibular plates may be analogous to the sesamoid
bones of bony vertebrates. Sesamoid bones can form
in tendons, often near articulations, and are thought
to modify pressure or diminish friction (Gray, 1918).
More derived sharks, family Lamnidae, subse-
quently lost the ancestral paired ethmopalatine lig-
aments while retaining the novel palatonasal liga-
ment (Figs. 3C,D, 5-4). Consequently, basal
lamniform species have increased the anterior liga-
mentous support of the upper jaw, while derived
lamniform species have decreased this support.
113
A number of evolutionary changes in the muscu-
loskeletal apparatus of the jaw suspension among
elasmobranchs have taken place, as revealed by
plotting these morphological characters onto an ex-
isting cladogram (Shirai, 1996) (Fig. 6). The out-
group to Elasmobranchi is the Holocephali, or chi-
meras, in which the upper jaw is fused to the
cranium and thus lacks cranial-palatoquadrate lig-
aments (Fig. 6-1) (Huxley, 1876; Gregory, 1904). The
basal morphology of the ethmopalatine ligament in
elasmobranchs appears to be a tubular sleeve-like
ligament that overlies and contains the orbital or
ethmoid process of the palatoquadrate and extends
to the outer edges of the ethmoid groove in the
cranium (Fig. 6-2) (Wilga, 2002). Thus far, basal and
derived squalean (Chlamydoselachiformes: Chlamy-
doselachus anguineus; Hexanchiformes: Notoryn-
chus cepedianus, Heptranchus perlo; Squaliformes:
Squalus acanthias), orectolobiform (Ginglymostoma
cirratum, Chiloscyllium plagiosum), and heterodon-
tiform (Heterodontus francisci) sharks that have
been examined have the sleeve-like ethmopalatine
ligament condition (Daniel, 1915; Allis, 1923; Motta
and Wilga, 1995, 1999; Wilga and Motta, 1998a,
2000). Lamniformes and Carcharhiniformes evolved
solid rope-like ethmopalatine ligaments connecting
the ethmoid process of the upper jaw to the ethmoid
region of the cranium (Fig. 6-3). All Lamniformes
evolved an additional single rope-like palatonasal
ligament (Fig. 6-4), while the ethmopalatine liga-
ments were lost in derived lamniform species, Fam-
ily Lamnidae (Fig. 6-5). The condition of the cranial-
palatine ligaments in the less well-known
lamniform families, Psudeocarchariidae, Megachas-
midae, and Ceteorhinidae, is unknown, and thus it
is possible that the loss of the palatonasal ligament
may be more widespread or there may be multiple
independent losses. Hexanchiformes evolved an ad-
ditional postorbital articulation (Fig. 6-6), while ba-
toids lost the ethmopalatine articulation (Fig. 6-7).
Evolution of the Jaw Suspension of
Elasmobranchii
A hypothetical ancestral condition for gnathos-
tomes called autodiastyly was proposed because it
presented a nonfused autostylic state from which all
other jaw suspension types could be derived (De
Beer and Moy-Thomas, 1935; Miles and Westoll,
1968; Miles, 1969; Moy-Thomas and Miles, 1971;
Lund and Grogan, 1997; Grogan et al., 1999; Grogan
and Lund, 2000). In autodiastyly, the hyoid arch is
nonsuspensorial and is similar in morphology to the
branchial arches but articulates with the palato-
quadrate (upper jaw) (De Beer and Moy-Thomas,
1935). In this state the palatoquadrate is not fused
to the cranium and has ethmoidal and orbital artic-
ulations with the cranium, which are considered
basal gnathostome characters (Janvier, 1996). Chi-
maeroid and selachian jaw suspension states are all
114 C.D. WILGA

Fig. 6. Evolution of cranial-palatoquadrate connections in elasmobranchs, with holocephalans as the outgroup (phylogeny after
Shirai, 1996). Jaw suspension types are in parentheses. 1: Holostylic condition illustrated by Callorhinchus milli (after Jollie, 1962).
2: Basal elasmobranch condition illustrated by Squalus acanthias (after Wilga and Motta, 1998a). 3: Carcharhiniform and lamniform
condition illustrated by Negaprion brevirostris (after Motta and Wilga, 1995). 4: Basal lamniform condition illustrated by Mitsukurina
owstoni. 5: Lamnid condition illustrated by Isurus oxyrinchus. 6: Hexanchiform condition illustrated by Heptranchias perlo (after
Daniel, 1922). 7: Batoid condition illustrated by Rhinobatos lentingosus (after Wilga and Motta, 1998b). Black, ethmopalatine and
postorbital ligaments; dark gray, hyomandibula; light gray, ceratohyal; white, cranium, palatoquadrate, and Meckel’s cartilage.

derivable from an autodiastylic plesiomorphic con-
dition (Fig. 7) (Grogan and Lund, 2000). Fusion of
the palatoquadrate to the cranium evolved in holo-
cephalans, while elaboration of a postorbital articu-
lation and modification of the epihyal to form a
suspensory hyomandibula in amphistyly evolved in
selachians (Lund and Grogan, 1997; Grogan et al.,
1999). Hexanchiform sharks retained the postor-
bital articulation as it acquired the orbitostylic ar-
ticulation, and therefore possesses two jaw suspen-
sion types, both of which contain sleeve-like
ligaments. The orbital articulation became promi-
nent and the postorbital articulation was lost in
orbitostyly as it spread throughout squalean sharks.
If batoids are derived squaloids (Carvalho, 1996;
Shirai, 1996), then loss of the orbital articulation
created a modified suspension that is “truly” hyo-
stylic or euhyostylic, with the hyomandibula func-
tioning as the sole means of jaw support. If batoids
are the sister group to sharks (Douady et al., 2003;
Winchell et al., 2004), then they lost the ethmopala-
tine ligaments from the autodiastylic ancestor and
sharks evolved them independently. Deeberius ellef-
seni, an autodiastylic chondrichthyan, shows the
typical hypermineralization reinforcement of the
cartilage at the articulation sites in response to lo-
calized stress, such as that at a joint (Grogan and
Lund, 2000), which supports the supposition that
autodiastylic chondrichthyans had an ethmopala-
tine articulation. Only the ethmoid articulation,
which became solid and rope-like, was retained in
the hyostyly of galean sharks (Maisey and Carvalho,
1997). The ethmoid articulation moved medially on
the palatoquadrate in Heterodontiformes and Orec-
tolobiformes. Lamniforms evolved a single solid pal-
atonasal ligament, while lamnids subsequently lost
the paired ethmopalatine ligaments.
Evolution of Upper Jaw Mechanisms
Two cranial-palatoquadrate muscles, the preor-
bitalis and levator palatoquadrate, act to protrude
or retract the kinetic upper jaw in elasmobranchs.
Holocephalans appear to have lost these muscles
 JAW MORPHOLOGY OF LAMNIFORM SHARKS 115

Fig. 7. Possible evolutionary sequence for jaw suspension types in chondrichthyes. See text for description (autodiastylic after
Grogan and Lund, 2000; chimera after Jollie, 1962; amphistylic, sevengill, and heterodontiform after Schaefer and Rosen, 1961;
dogfish after Wilga and Motta, 1998a; batoid after Wilga and Motta, 1998b). Black, ethmopalatine and postorbital ligament; blue,
Meckel’s cartilage; gray, cranium; green, ceratohyal; red, hyomandibula, yellow, palatoquadrate or upper jaw.

when the upper jaw fused to the cranium (Edge-
worth, 1935; Didier, 1995). The ancestral condition
in elasmobranchs appears to be a horizontally di-
rected preorbitalis muscle that protrudes the upper
jaw (Fig. 8-1a), as represented by Squalus acanthias
(Wilga and Motta, 1998; Wilga et al., 2002). In Het-
erodontiformes and Orectolobiformes the preor-
bitalis muscle is subdivided, vertically aligned, and
inserts onto Meckel’s cartilage rather than onto the
quadratomandibularis as in other sharks (Fig. 8-1b)
(Smith, 1942; Compagno, 1977; Motta and Wilga,
1999; Wilga et al., 2002) and probably functions to
adduct the jaws rather than protrude the upper jaw.
In basal carcharhiniform and lamniform taxa the
preorbitalis has undergone a reversal back to the
horizontal orientation, but still functions to protrude
the upper jaw (Fig. 1c) (Luther, 1909b; Edgeworth,
1935; Compagno, 1977, 1988; Motta and Wilga,
1995; Wilga and Motta, 2000; Wilga et al., 2002).
Basal lamniform species have lost the new preor-
bitalis division in yet another reversal (Fig. 8-1d),
while the more derived Lamnidae retained the sub-
divided preorbitalis muscle (Fig. 8-1e) (Luther,
1909b; Edgeworth, 1935) that presumably still func-
tions to protrude the upper jaw. The only squalean
taxa to evolve multiple subdivided preorbitalis mus-
cles are batoids and their sister group, the saw
sharks Pristiophoridae (Fig. 8-1f) (Marion, 1905;
Luther, 1909b; Shirai, 1992; Wilga and Motta,
1998). One of the new preorbitalis divisions in ba-
toids is vertically aligned and inserts onto the lower
jaw (Marion, 1905; Luther, 1909; Wilga and Motta,
1998b), indicating that it would function to elevate
the lower jaw rather than protrude it. If batoids are
the sister group to sharks (Douady et al., 2003;
Winchell et al., 2004), rather than modified
squaleans, as shown in the figures, then they were
the first elasmobranch group to evolve duplicated
preorbitalis muscles instead of the ancestor to the
heterodontiform-orectolobiform lineage. If so, then
the other preorbitalis division became vertically
aligned in heterodontids and orectolobids. In gen-
eral for elasmobranch species, the preorbitalis divi-
sions that insert onto the palatoquadrate function to
protrude the upper jaw, while those divisions that
have been modified to insert onto Meckel’s cartilage
116 C.D. WILGA

Fig. 8. Evolution of upper jaw muscles in elasmobranchs (phylogeny after Shirai, 1996). 1a,2a: Basal elasmobranch condition,
illustrated by Squalus acanthias (after Wilga and Motta, 1998a). 1b,2b: Basal galeomorph condition illustrated by Ginglymostoma
cirratum (after Motta and Wilga, 1999). 1d: Basal lamniform condition illustrated by Carcharius taurus. 2d: Derived Carcharhiniform
condition illustrated by Sphyrna tiburo Carcharhinidae (after Wilga and Motta, 2000). *Note that some place sphyrnids in their own
family as the sister group to Carcharhinidae. 1e,2e: Derived Lamniform condition illustrated by Isurus oxyrinchus Lamnidae. 1f:
Batoid condition illustrated by Rhinobatos lentiginosus (after Wilga and Motta, 1998b). Color of muscular figures: blue, quadrato-
mandibularis; green, levator palatoquadrati; red, levator hyomandibularis; yellow, preorbitalis. Dotted line indicates location and
orientation of the levator palatoquadrati muscles when overlaid by quadratomandibularis and preorbitalis muscles. *Note that not all
families are shown in order for clarity.

have a modified function as well in elevating the
lower jaw (Wilga and Motta, 1998). Duplication by
subdivision of the ancestral single preorbitalis mus-
cle appears to have evolved in most elasmobranch
groups, apparently independently in some (Carchar-
hiniformes, Lamnidae, Pristiophoridae, and Ba-
toidea) and with several different morphologies.
Multiple preorbitalis muscle insertions will increase
the bite force and spread it over a larger area of the
upper jaw as well as provide finer control of upper
jaw protrusion.
The ancestral condition of the levator palatoqua-
drati in elasmobranchs appears to be a vertically
directed muscle that functions to retract the upper
jaw (Fig. 8-2a) as represented by Squalus acanthias
(Luther, 1909b; Edgeworth, 1935; Shirai, 1992;
Wilga and Motta, 1998a). Due to shifting of the jaw
joint under the cranium in Heterodontiformes and
Orectolobiformes, the orientation of the levator pala-
toquadrati is now horizontal (Fig. 8-2b), but is still
in a position to retract the upper jaw due to the
anteroventral location of the insertion relative to the
origin (Luther, 1909b; Edgeworth, 1935; Smith,
1942; Compagno, 1977; Motta and Wilga, 1999;
Wilga et al., 2002). The levator palatoquadrati has
reverted to the ancestral vertical orientation in car-
charhiniform and lamniform species (Fig. 8-2c)
(Luther, 1909b; Edgeworth, 1935; Compagno, 1977,
1988). The origin of the levator palatoquadrati ex-
panded and moved more anteriorly on the cranium
in sharks of the Family Carcharhinidae, resulting in
a more horizontal orientation that now functions to
assist the two preorbitalis divisions in protruding
the upper jaw (Fig. 8-2d) (Compagno, 1988; Motta
 JAW MORPHOLOGY OF LAMNIFORM SHARKS
and Wilga, 1995; Wilga and Motta, 2000; Wilga et
al., 2002).
The ancestral state of the levator hyomandibu-
laris in elasmobranchs is an insertion onto the hyo-
mandibula from an origin on the cranium or epaxial
musculature (Fig. 8-3a) (Luther, 1909b; Edgeworth,
1935; Compagno, 1977; Motta and Wilga, 2001;
Wilga et al., 2002). In this state the levator hyoman-
dibularis functions to elevate the hyomandibula,
which in turn retracts the jaws after they have been
protruded (Motta et al., 1997; Wilga and Motta,
1998). In basal lamniform species the levator hyo-
mandibularis is similar to that of other elasmo-
branchs and most likely also retracts the jaws after
protrusion. However, the insertion of the levator
hyomandibularis has been altered in alopid and
lamnid sharks to insert primarily onto the palato-
quadrate and Meckel’s cartilage surrounding the
jaw joint, although it does retain a partial insertion
onto the hyomandibular cartilage as well (Fig. 8-3b).
Thus, Alopias vulpinnis represents an intermediate
condition between basal and derived lamniforms in
having both ethmopalatine and palatonasal liga-
ments, a partially subdivided preorbitalis, and the
modified insertion of the levator hyomandibularis
muscle. Therefore, the loss of the ligaments does not
appear to have driven the derived muscular condi-
tion. The function of the levator hyomandibularis
probably has not been greatly affected by the new
insertion; elevating the jaws in the region of the jaw
joint is simply a more direct way of retracting the
jaws compared to the more indirect mechanism
through the hyomandibular cartilage of other shark
species. However, the onset of levator hyomandibu-
laris muscle activation relative to the onset of jaw
retraction has probably been shortened, and thus
the gape cycle, due to the more direct connection. A
reduced duration of the bite cycle will increase prey
capture performance in those species that prey on
elusive fishes, as do mako Isurus oxyrinchus,
thresher A. vulpinnis, and porbeagle Lamna nasus
sharks (Bigelow and Schroeder, 1948; Compagno,
1984).
Note that an alternative scenario may have taken
place in the evolution of the preorbitalis and levator
palatoquadrati muscles at the base of the galeo-
morph radiation. Heterodontiform and orectolobi-
form sharks may have independently acquired the
vertical orientation of the preorbitalis and the hori-
zontal orientation of the levator palatoquadrati
muscles as the jaws shifted more anteriorly under
the cranium (Fig. 8-1b,2b). If so, then the orientation
of the preorbitalis and levator palatoquadrati mus-
cles in carcharhiniform and lamniform sharks re-
flects the basal condition rather than a reversal back
to the basal condition (Fig. 8-1c,2c). Both scenarios
(this paragraph with two independent character ac-
quisitions in sister groups and the ones shown in the
figures with two independent reversals) require two
117
evolutionary changes occurring as mapped onto the
existing cladogram.
The addition of a direct insertion of the levator
hyomandibularis muscle onto the jaws in lamnids
resulted in greater active support of the jaws com-
pared to the situation in most shark species. A mus-
cular connection provides increased flexibility in ac-
tively preventing overextension of the jaws at
continuously variable lengths compared to a liga-
mentous connection that prevents overextension
only when fully extended. In other shark taxa the
levator hyomandibularis inserts only onto the hyo-
mandibular cartilage and when activated retracts
the hyomandibula, which then retracts the jaw ap-
paratus back under the cranium through the
hyomandibula-mandibular joint. This leaves only
the relatively small levator palatoquadrati muscle
to retract the upper jaw during the recovery phase
in most shark species (Motta et al., 1997; Wilga et
al., 2002). However, alopid and lamnid sharks
have a distinct advantage in that the relatively
larger levator hyomandibularis, as well as the le-
vator palatoquadrati, are in a position to retract
the upper jaw.
Possession of the direct levator hyomandibularis
insertion and duplicated preorbitalis muscle inser-
tions onto the upper jaw in lamnid species should
permit more forceful and controlled movements of
the upper jaw. This is advantageous in those species
that engage in gouging chunks from oversized prey,
such as white sharks, Carcharodon carcharius
(Tricas and McCosker, 1984; Klimley and Ainley,
1998). Indeed, white sharks are notable for the abil-
ity to protrude and retract the upper jaw several
times during a gape cycle (Tricas and McCosker,
1984; Klimley and Ainley, 1998), something that has
not yet been observed in carcharhiniform and
squaliform sharks (Motta and Wilga, 2001; Motta,
2004). Thus, lamnid sharks appear to be well
equipped for the repeated protrusion and retraction
cycles of the upper jaw required to gouge bites from
large or elusive prey.
In summary, there has been at least one instance
of character loss, acquisition, subdivision, and mod-
ification in the jaw suspension and muscles associ-
ated with jaw protrusion and retraction in lamni-
form sharks. It is of special significance that as
lamnid sharks lost the ancestral support of the eth-
moid articulation to the upper jaw anteriorly, they
gained new anterior muscular support to the jaws
through the duplicated preorbitalis muscle. This,
coupled with the new muscular connection of the
levator hyomandibularis posteriorly provides flexi-
bility in actively preventing overextension of the
jaws at various lengths and by allowing repeated
protrusion–retraction cycles of the upper jaw within
one gape cycle without first having to retract the
entire jaw apparatus. Finally, although evolution
has provided multiple insertions of the preorbitalis
muscles, and thus anterior support of the upper jaw,
118 C.D. WILGA
in both carcharhiniform and lamnid sharks, car-
charhiniform sharks have acquired more muscular
support for protrusion of the jaws by way of the
levator palatoquadrati, while lamnid sharks ac-
quired more muscular support for retraction of the
jaws by way of the levator hyomandibularis.
ACKNOWLEDGMENTS
Thanks to Karsten Hartel, Karel Liem, and
George Lauder from the Museum of Comparative
Zoology at Harvard University for providing speci-
mens for dissection. Tony Wood and Mike Smith
collected fresh specimens at local shark tourna-
ments. Also, sincere thanks to Jason Ramsay for
drawing the anatomical figures. Supported by NSF
IBN 0137912 and NOAA/CMER NA97FE0363.
LITERATURE CITED
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trusible upper jaws of some acanthopterygian fish. J Zool Lond
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