Pneumatized bones
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Skeletal pneumaticity is the presence of air spaces within bones. It is generally produced during development by excavation of bone by pneumatic
diverticula In medicine or biology, a diverticulum is an outpouching of a hollow (or a fluid-filled) structure in the body. Depending upon which layers of the structure are involved, diverticula are described as being either true or false. In medicine, t ...
(air sacs) from an air-filled space, such as the lungs or nasal cavity. Pneumatization is highly variable between individuals, and bones not normally pneumatized can become pneumatized in pathological development.


Cranial pneumaticity

Pneumatization occurs in the skulls of
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,
crocodilian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest livin ...
s and
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 among extant
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
s. Pneumatization has been documented in extinct
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
s including
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s and
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
s. Pneumatic spaces include the
paranasal sinuses Paranasal sinuses are a group of four paired air-filled spaces that surround the nasal cavity. The maxillary sinuses are located under the eyes; the frontal sinuses are above the eyes; the ethmoidal sinuses are between the eyes and the sph ...
and some of the
mastoid cells The mastoid cells (also called air cells of Lenoir or mastoid cells of Lenoir) are air-filled cavities within the mastoid process of the temporal bone of the cranium. The mastoid cells are a form of skeletal pneumaticity. Infection in these cells ...
.


Postcranial pneumaticity

Postcranial pneumaticity is found largely in certain
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
groups, namely
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s,
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
s, and
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. Vertebral pneumatization is widespread among
saurischian Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithis ...
dinosaurs, and some theropods have quite widespread pneumatization, for example '' Aerosteon riocoloradensis'' has pneumatization of the ilium,
furcula The (Latin for "little fork") or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is formed by the fusion of the two pink clavicles. In birds, its primary function is in the strengthening of the thoracic ...
, and
gastralia Gastralia (singular gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In thes ...
. Many modern birds are extensively pneumatized. The air pockets of the bones are connected to the pulmonary air sacs: However the extent of pneumaticity depends on species. For example it is slight in diving birds,
loon Loons ( North American English) or divers ( British / Irish English) are a group of aquatic birds found in much of North America and northern Eurasia. All living species of loons are members of the genus ''Gavia'', family Gaviidae and order ...
s lack pneumatic bones at all. Postcranial pneumatization is rarer outside of Archosauria. Examples include the hyoid in howler monkeys
Alouatta Howler monkeys (genus ''Alouatta'', monotypic in subfamily Alouattinae) are the most widespread primate genus in the Neotropics and are among the largest of the platyrrhines along with the muriquis (''Brachyteles''), the spider monkeys (''Atele ...
, and the dorsal vertebrae in the osteoglossiform fish '' Pantodon''. Slight pneumatization of the centra and rib heads by dorsal diverticula in the lungs of land tortoises has also been documented. In addition, pathological pneumatization has been known to occur in the human
atlas vertebra In anatomy, the atlas (C1) is the most superior (first) cervical vertebra of the spine and is located in the neck. It is named for Atlas of Greek mythology because, just as Atlas supported the globe, it supports the entire head. The atlas is t ...
.


Function of skeletal pneumaticity

The exact function of skeletal pneumaticity is not definitively known, but there are several working hypotheses concerning the role of skeletal pneumaticity in an organism.


Reduce body mass

By invading the bones, the pneumatic diverticula would replace marrow with air, reducing the overall body mass. Reducing the body mass would make it easier for pterosaurs and birds to fly as there is less mass to keep aloft with the same amount of muscle powering the flight strokes. Pneumatizing the vertebral column of
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', ' lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their ...
s would reduce the weight of these organisms, and make it easier to support and move the massive neck.


Alter skeletal mass distribution

Skeletal pneumaticity allows animals to redistribute the skeletal mass within their body. The skeletal mass of a bird (pneumatized) and a mammal (not pneumatized) with similar body size is roughly the same, yet the bones of birds were found to be denser than the bones of mammals. This suggests that pneumatization of bird bones does not affect the overall mass but allows for a better balance of weight within the body to allow for greater balance, agility and ease of flight.


Balance

In theropods, the head and neck are greatly pneumatized, and the forearms are reduced. This would help reduce the mass further away from the center of balance. This adjustment to the center of mass would allow the animal to reduce its rotational inertia, thereby increasing its agility. The sacral pneumaticity would lower its center of mass to a more ventral position, allowing it more stabilization.


Adaptation for high altitude

Screamer The screamers are three South American bird species placed in family Anhimidae. They were thought to be related to the Galliformes because of similar bills, but are more closely related to ducks (family Anatidae),Todd, F. (1991) and most closel ...
s are highly pneumatized birds with pneumatic diverticula traveling throughout the bones and into the skin. As screamers fly at high altitudes, it is hypothesized that the extreme pneumaticity in these birds is indicative of an adaptation for flying in high altitudes.


References

{{Reflist, 2 Vertebrate anatomy