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Cheryl Wilga.
Morphology and Evolution of the Jaw Suspension in Lamniform Sharks. Journal of Morphology. 265:102–119.
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ABSTRACT
The morphology of the
jaw suspension and jaw protrusion mechanism in lamniform sharks is described 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 (Carcharhiniformes, Lamniformes,
Orectolobiformes, and Heterodontiformes) 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 connecting the symphysis of the upper jaw to the cranium midventral to
the nasal capsule. Sharks in the family Lamnidae subsequently lost the original
paired ethmopalatine ligament while retaining the novel palatonasal ligament.
Thus, basal lamniform taxa (Mitsukurina owstoni, Carcharius taurus, Alopias
vulpinnis) have increased ligamentous 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
protrusion 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 preorbitalis muscle with the new division inserting near the ethmoid
process of the palatoquadrate (upper jaw). Alopid sharks have also independently
evolved a partially subdivided 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 inserts 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 functions to retract the jaws
after protrusion. In alopid and lamnid sharks the levator hyomandibularis
inserts primarily 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
ligament), subdivision (preorbitalis), and modification (ventral preorbitalis,
dorsal preorbitalis, and levator hyomandibularis) in the ligaments and muscles
associated with the jaw suspension and jaw protrusion mechanism in lamniform
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 acquired muscular
support by way of the levator hyomandibularis, which provides a dynamic
mechanism for lateral support. The evolution of multiple divisions of
preorbitalis insertions onto the palatoquadrate and modification 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.