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The skull roof, or the roofing bones of the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
, are a set of
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
s covering the
brain A brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as Visual perception, vision. I ...
, eyes and nostrils in
bony fishes Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilage ...
and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparative anatomy the term is used on the full dermatocranium. Romer, A.S. & T.S. Parsons. 1977. ''The Vertebrate Body.'' 5th ed. Saunders, Philadelphia. (6th ed. 1985) In general anatomy, the roofing bones may refer specifically to the bones that form above and alongside the brain and neurocranium (i.e., excluding the marginal upper jaw bones such as the maxilla and premaxilla), and in human anatomy, the skull roof often refers specifically to the skullcap.


Origin

Early armoured fish did not have a skull in the common understanding of the word, but had an endocranium that was partially open above, topped by dermal bones forming armour. The dermal bones gradually
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variati ...
into a fixed unit overlaying the endocranium like a heavy "lid", protecting the animal's head and brain from above.
Cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
whose skeleton is formed from cartilage lack a continuous dermal armour and thus have no proper skull roof. A more or less full shield of fused dermal bones was common in early
bony fishes Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilage ...
of the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
, and particularly well developed in shallow water species.


Bony fishes and amphibians

In early
sarcopterygian Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includ ...
s the skull roof was composed of numerous bony plates, particularly around the nostrils and behind each eye. The skull proper was joined by the bones of the operculum. The skull itself was composed rather loosely, with a joint between the bones covering the brain (the
parietal bone The parietal bones () are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is n ...
s and the bones behind them) and the snout (the
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, ...
,
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Ea ...
and the bones in front and to the side of them). This joint disappeared in the evolving labyrinthodonts, at the same time the number of bones were reduced and the operculum disappeared. In frogs and
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
s the skull roof is further reduced and has large openings. Only in caecilians can a full covering skull roof be found, an adaption for burrowing. The skull roof in
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
is composed of a number of bony plates that are not readily compared to those found in early amphibians. Carroll, R.L. (1988): Vertebrate Paleontology and Evolution,'' W.H. Freeman & Co'', New York NY, p.148 In most ray-finned fishes the skull is often reduced to a series of loose elements, and a skull roof as such is not found.


Labyrinthodonts and early reptiles

The pattern of plates of the labyrinthodonts formed that basis for that seen in all land-living vertebrates. The roof itself formed a continuous cover over the whole of the head, leaving only openings for nostrils, eyes and a parietal eye between the
parietal bone The parietal bones () are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is n ...
s. This type of skull was inherited by the first reptiles as they evolved from labyrinthodont stock in the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
. This type of skull roof without any above openings behind the eyes is called ''
anapsid An anapsid is an amniote whose skull lacks one or more skull openings (fenestra, or fossae) near the temples. Traditionally, the Anapsida are the most primitive subclass of amniotes, the ancestral stock from which Synapsida and Diapsida evolv ...
''. Today anapsid skulls are only found in
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
s, though this may be a case of secondary loss of the post orbital openings.deBraga, M. and Rieppel, O. (1997). "Reptile phylogeny and the interrelationships of turtles." ''Zoological Journal of the Linnean Society'', 120: 281-354.


Diapsids and synapsids

In two groups of early reptiles the skull roof evolved post orbital openings to allow for greater movement of the jaw muscles. The two groups evolved the openings independently: * The Synapsids having one opening on each side, fairly low on the side of the skull, between the zygomatic bone and the elements above. * The Diapsids having two openings on each side, the two openings separated by an arch formed from processes of the postorbital and squamosal bones. The synapsids are the mammal-like reptiles and the
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s. In mammals, the side opening is closed by the sphenoid bone, so that the skull roof appear whole, despite the temporal opening. All other reptiles (with the possible exception of turtles) and the
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s are diapsids.


References

{{Reflist Vertebrate anatomy Human head and neck Flat bones