Bird anatomy

Bird anatomy, or the physiological structure of birds' bodies, shows many unique adaptations, mostly aiding flight. Birds have evolved a light skeletal system and light but powerful muculature which, along with circulatory and respiratory systems capable of very high metabolic rates and oxygen supply, permit the bird to fly. The development of a beak has led to evolution of a specially adapted digestive system. These anatomical specializations have earned birds their own class in the vertebrate phylum.

Due to the high metabolic rate required for flight, birds have a high oxygen demand. Development of an efficient respiratory system enabled the evolution of flight in birds. Birds ventilate their lungs by means of air sacs, structures unique to birds and dinosaurs. These sacs do not play a direct role in gas exchange, but rather store air and act like bellows, allowing the lungs to maintain a fixed volume with fresh air constantly flowing through them. ^[1]

Three distinct sets of organs perform respiration—the anterior air sacs (interclavicular, cervicals, and anterior thoracics), the lungs, and the posterior air sacs (posterior thoracics and abdominals). The posterior and anterior air sacs, typically nine, expand during inhalation. Air enters the bird via the trachea. Half of the inhaled air enters the posterior air sacs, the other half passes through the lungs and into the anterior air sacs. The sacs contract during exhalation. Air from the anterior air sacs empties directly into the trachea and out the birds mouth or nares. The posterior air sacs empty their air into the lungs. Air passing through the lungs as the bird exhales is expelled via the trachea.

Since fresh air flows through the lungs in only one direction, there is no mixing of oxygen-rich air and oxygen-poor, carbon dioxide-rich air as in mammalian lungs. Thus, the partial pressure of oxygen in a bird's lungs is the same as the environment, and so birds have more efficient gas-exchange of both oxygen and carbon dioxide than do mammals.

Avian lungs do not have alveoli, as mammalian lungs do, but instead contain millions of tiny passages known as parabronchi, connected at either ends by the dorsobronchi and ventrobronchi. Air flows through the honeycombed walls of the parabronchi and into air capillaries, where oxygen and carbon dioxide are traded with cross-flowing blood capillaries by diffusion.

Birds also lack a diaphragm. The entire body cavity acts as a bellows to move air through the lungs. The active phase of respiration in birds is exhalation, requiring muscular contraction.

Parabronchi are millions of tiny passages in the lungs of birds that constitute the primary sites of gas exchange between air and blood for birds. This function in mammals is performed by the alveoli, which is an air sac.

Air flows from the parabronchi into air vesicles, called atria, which project radially from the parabronchi. These atria give rise to air capillaries, where oxygen and carbon dioxide are traded with cross-flowing blood capillaries by diffusion. [2]

The syrinx is the sound-producing vocal organ of birds, located at the base of a bird's trachea. As with the mammalian larynx, sound is produced by the vibration of air flowing through the organ. The syrinx enables some species of birds to produce extremely complex vocalizations, even mimicking human speech. In some songbirds, the syrinx can produce more than one sound at a time.

Birds have four chambered hearts, in common with humans, most mammals and some reptiles. This adaptation allows for efficient nutrient and oxygen transport throughout the body, providing birds with energy to fly and maintain high levels of activity. A Ruby-throated Hummingbird's heart beats up to 1200 times per minute (about 20 beats per second).^[2]

Many birds possess a muscular pouch along the esophagus called a crop. The crop functions to both soften food and regulate its flow through the system by storing it temporarily. The size and shape of the crop is quite variable among the birds. Members of the order Columbiformes, such as pigeons, produce a nutritious crop milk which is fed to their young by regurgitation. Birds possess a ventriculus, or gizzard, composed of four muscular bands that rotate and crush food by shifting the food from one area to the next within the gizzard. The gizzard of some species contains small pieces of grit or stone swallowed by the bird to aid in the grinding process of digestion, serving the function of mammalian or reptilian teeth. The use of gizzard stones is a similarity between birds and dinosaurs, which left gizzard stones called gastroliths as trace fossils.

The bird skeleton is highly adapted for flight. It is extremely lightweight but strong enough to withstand the stresses of taking off, flying, and landing. One key adaptation is the fusing of bones into single ossifications, such as the pygostyle. Because of this, birds usually have a smaller number of bones than other terrestrial vertebrates. Birds also lack teeth or even a true jaw, instead having evolved a beak, which is far more lightweight. The beaks of many baby birds have a projection called an egg tooth, which facilitates their exit from the egg.

Birds have many bones that are hollow with criss-crossing struts or trusses for structural strength. The number of hollow bones varies among species, though large gliding and soaring birds tend to have the most. Respiratory air sacs often form air pockets within the semi-hollow bones of the bird's skeleton.^[1] Some flightless birds like penguins and ostriches have only solid bones, further evidencing the link between flight and the adaptation of hollow bones.

Birds also have more cervical (neck) vertebrae than many other animals; most have a highly flexible neck consisting of 13-25 vertebrae. Birds are the only vertebrate animals to have a fused collarbone (the furcula or wishbone) or a keeled sternum or breastbone. The keel of the sternum serves as an attachment site for the muscles used for flight, or similarly for swimming in penguins. Again, flightless birds, such as ostriches, which do not have highly developed pectoral muscles, lack a pronounced keel on the sternum.

Birds have uncinate processes on the ribs, a feature also found in the tuatara Sphenodon. They also have a greatly elongate tetradiate pelvis as in some reptiles. The hindlimb has an intra-tarsal joint found also in some reptiles. There is extensive fusion of the trunk vertebrea as well as fusion with the pectoral girdle. They have a diapsid skull as in reptiles with a pre-lachrymal fossa (present in some reptiles). The skull has a single occipital condyle.^[3]

The skull consists of five major bones: the frontal (top of head), parietal (back of head), premaxillary and nasal (top beak), and the mandible (bottom beak). The skull of a normal bird usually weighs about 1% of the birds total bodyweight.

The vertebral column consists of vertebrae, and is divided into three sections: cervical (13-16) (neck), Synsacrum (fused vertebrae of the back, also fused to the hips (pelvis)), and pygostyle (tail).

The chest consists of the furcula (wishbone) and coracoid (collar bone), which two bones, together with the scapula (see below), form the pectoral girdle. The side of the chest is formed by the ribs, which meet at the sternum (mid-line of the chest).

The shoulder consists of the scapula (shoulder blade), coracoid (see The Chest), and humerus (upper arm). The humerus joins the radius and ulna (forearm) to form the elbow. The carpus and metacarpus form the "wrist" and "hand" of the bird, and the digits (fingers) are fused together. The bones in the wing are extremely light so that the bird can fly more easily.

The hips consist of the pelvis which includes three major bones: Illium (top of the hip), Ischium (sides of hip), and Pubis (front of the hip). These are fused into one (the innominate bone). They meet at the acetabulum (the hip socket) and articulate with the femur, which is the first bone of the hind limb.

The upper leg consists of the femur. At the knee joint, the femur connects to the tibiotarsus (shin) and fibula (side of lower leg). The tarsometatarsus forms the upper part of the foot, digits make up the toes. The leg bones of birds are the heaviest, contributing to a low center of gravity. This aids in flight.

Most birds have approximately 175 different muscles, mainly controlling the wings, skin, and legs. The largest muscles in the bird are the pectorals, or the breast muscles, which control the wings and make up about 15 - 25% of a flighted bird’s body weight. They provide the powerful wing stroke essential for flight. The muscle ventral (underneath) to the pectorals is the supracoracoideus. It raises the wing between wingbeats. The supracoracoideus and the pectorals together make up about 25 – 35% of the bird's full body weight.

The skin muscles help a bird in its flight by adjusting the feathers, which are attached to the skin muscle and help the bird in its flight maneuvers.

There are only a few muscles in the trunk and the tail, but they are very strong and are essential for the bird. The pygostyle controls all the movement in the tail and controls the feathers in the tail. This gives the tail a larger surface area which helps keep the bird in the air.

See also: Avian pallium Birds have acute eyesight - raptors have vision eight times sharper than humans - thanks to higher densities of photoreceptors in the retina (up to 1,000,000 per square mm in Buteos, compared to 200,000 for humans), a high number of optic nerves, a second set of eye muscles not found in other animals, and, in some cases, an indented fovea which magnifies the central part of the visual field. Many species, including hummingbirds and albatrosses, have two foveas in each eye. Many birds can detect polarised light. The eye occupies a considerable part of the skull and is surrounded by a sclerotic eye-ring, a ring of tiny bones that surround the eye. This character is also see in the reptiles.

Birds have a large brain to body mass ratio. This is reflected in the advanced and complex bird intelligence.

The region between the eye and bill on the side of a bird's head is called the lore. This region is sometimes featherless, and the skin may be tinted, as in many species of the cormorant family.

Although most male birds have no external sex organs, the male does have two testes which become hundreds of times larger during the breeding season to produce sperm. The female's ovaries also become larger, although only the left ovary usually functions.

In the males of species without a phallus (see below), sperm is stored in the seminal glomera within the cloacal protuberance prior to copulation. During copulation, the female moves her tail to the side and the male either mounts the female from behind or in front (in the stitchbird), or moves very close to her. The cloacae then touch, so that the sperm can enter the female's reproductive tract. This can happen very fast, sometimes in less than half a second.

The sperm is stored in the female's sperm storage tubules for a week to a year, depending on the species. Then, eggs will be fertilised individually as they leave the ovaries, before being laid by the female. The eggs continue their development outside the female body.

Many waterfowl and some other birds, such as the ostrich and turkey, possess a phallus. When not copulating, it is hidden within the proctodeum compartment within the cloaca, just inside the vent.

After the eggs hatch, parents provide varying degrees of care in terms of food and protection. Precocial birds can care for themselves independently within minutes of hatching; altricial hatchlings are helpless, blind, and naked, and require extended parental care. The chicks of many ground-nesting birds such as partridges and waders are often able to run virtually immediately after hatching; such birds are referred to as nidifugous. The young of hole-nesters, on the other hand, are often totally incapable of unassisted survival. The process whereby a chick acquires feathers until it can fly is called "fledging".

Some birds, such as pigeons, geese, and Red-crowned Cranes, remain with their mates for life and may produce offspring on a regular basis.

• Peter Grant & Killian Mullarny The New Approach to Identification, in Birding World, Vols. 1&2 ISSN 0969-6024