The earliest jaws appeared in now extinct placoderms and spiny sharks during the Silurian, about 430 million years ago.
These linkages can be especially common and complex in the head of bony fishes, such as wrasses, which have evolved many specialized feeding mechanisms.
For suction feeding a system of linked four-bar linkages is responsible for the coordinated opening of the mouth and the three-dimensional expansion of the buccal cavity.
The simpler structure is found in jawless fish, in which the cranium is represented by a trough-like basket of cartilaginous elements only partially enclosing the brain, and associated with the capsules for the inner ears and the single nostril.
The cranium is a single structure forming a case around the brain, enclosing the lower surface and the sides, but always at least partially open at the top as a large fontanelle.
The most anterior part of the cranium includes a forward plate of cartilage, the rostrum, and capsules to enclose the olfactory organs.
Finally, the skull tapers towards the rear, where the foramen magnum lies immediately above a single condyle, articulating with the first vertebra.
The roof of the skull is generally well formed, and although the exact relationship of its bones to those of tetrapods is unclear, they are usually given similar names for convenience.
The skull roof is not fully formed, and consists of multiple, somewhat irregularly shaped bones with no direct relationship to those of tetrapods.
[11] This complex primitive pattern has, however, been simplified to various degrees in the great majority of vertebrates, as bones have either fused or vanished entirely.
In most vertebrates, the foremost part of the upper jaw, to which the incisors are attached in mammals consists of a separate pair of bones, the premaxillae.
[16] The premaxilla is unattached to the neurocranium (braincase); it plays a role in protruding the mouth and creating a circular opening.
To open the mouth, an adductor muscle pulls back the top of the maxilla, pushing the lower jaw forward.
[20] Many species can be readily recognized by their thick lips, the inside of which is sometimes curiously folded, a peculiarity which gave rise the German name of "lip-fishes" (Lippfische).
This "four-bar linkage" has the property of allowing numerous arrangements to achieve a given mechanical result (fast jaw protrusion or a forceful bite), thus decoupling morphology from function.
They are believed to have originated, in a similar way to oral jaws, as a modification of the fifth gill arch which no longer has a respiratory function.
The pharyngeal jaws of the moray are highly mobile, perhaps as an adaptation to the constricted nature of the burrows they inhabit which inhibits their ability to swallow as other fishes do by creating a negative pressure in the mouth.
The pharynx arises during development through a series of six or more outpocketings called pharyngeal arches on the lateral sides of the head.
The pharyngeal arches give rise to a number of different structures in the skeletal, muscular and circulatory systems in a manner which varies across the vertebrates.
However, the vertebrate pharynx is unique in that it gives rise to endoskeletal support through the contribution of neural crest cells.
The jaw's surface (in comparison to the vertebrae and gill arches) needs extra strength due to its heavy exposure to physical stress.
It has a layer of tiny hexagonal plates called "tesserae", which are crystal blocks of calcium salts arranged as a mosaic.
The hyoid articulates with the mandibular arch posteriorly, but it appears to provide little support to the upper and lower jaws.
[37] Shark teeth form within the jaw move outward in rows until they are eventually dislodged in a manner similar to a conveyor belt.
The purpose of the kype is not altogether clear, though they can be used to establish dominance by clamping them around the base of the tail (caudal peduncle) of an opponent.
[48] The pharyngeal jaw apparatus consists of two upper and one single lower plate, all of which have dentations that differ in size and type.
However studies of the cyclostomes, the jawless hagfishes and lampreys that did survive, have yielded little insight into the deep remodelling of the vertebrate skull that must have taken place as early jaws evolved.
[62] It is now accepted that the precursors of the jawed vertebrates are the long extinct bony (armoured) jawless fish, the so-called ostracoderms.
[66] Placoderms were a class of fish, heavily armoured at the front of their body, which first appeared in the fossil records during the Silurian about 430 million years ago.
Spiny sharks did not diversify as much as placoderms, but survived much longer into the Early Permian about 290 million years ago.