beak beak the beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. a beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for food, courtship, and feeding young. the terms beak and rostrum are also used to refer to a similar mouth part in some ornithischians, pterosaurs, cetaceans, dicynodonts, anuran tadpoles, monotremes (i.e. echidnas and platypuses, which have a beak-like structure), sirens, pufferfish, billfishes and cephalopods. comparison of bird beaks, displaying different shapes adapted to different feeding methods. not to scale. although beaks vary significantly in size, shape, color and texture, they share a similar underlying structure. two bony projections – the upper and lower mandibles – are covered with a thin keratinized layer of epidermis known as the rhamphotheca. in most species, two holes called nares lead to the respiratory system. etymology although the word "beak" was, in the past, generally restricted to the sharpened bills of birds of prey, in modern ornithology, the terms beak and bill are generally considered to be synonymous. the word, which dates from the 13th century, comes from the middle english bec, which itself comes from the latin beccus. anatomy the bony core of the beak is a lightweight framework, like that seen on this barn owl's skull. although beaks vary significantly in size and shape from species to species, their underlying structures have a similar pattern. all beaks are composed of two jaws, generally known as the maxilla (upper) and mandible (lower).(p147) the upper, and in some cases the lower, mandibles are strengthened internally by a complex three-dimensional network of bony spicules (or trabeculae) seated in soft connective tissue and surrounded by the hard outer layers of the beak.(p149) the avian jaw apparatus is made up of two units: one four-bar linkage mechanism and one five-bar linkage mechanism. mandibles a gull's upper mandible can flex upwards because it is supported by small bones which can move slightly backwards and forwards. the upper mandible is supported by a three-pronged bone called the intermaxillary. the upper prong of this bone is embedded into the forehead, while the two lower prongs attach to the sides of the skull. at the base of the upper mandible a thin sheet of nasal bones is attached to the skull at the nasofrontal hinge, which gives mobility to the upper mandible, allowing it to move upwards and downwards. position of vomer (shaded red) in neognathae (left) and paleognathae (right) the base of the upper mandible, or the roof when seen from the mouth, is the palate, the structure of which differs greatly in the ratites. here, the vomer is large and connects with premaxillae and maxillopalatine bones in a condition termed as a "paleognathous palate". all other extant birds have a narrow forked vomer that does not connect with other bones and is then termed as neognathous. the shape of these bones varies across the bird families. the lower mandible is supported by a bone known as the inferior maxillary bone—a compound bone composed of two distinct ossified pieces. these ossified plates (or rami), which can be u-shaped or v-shaped,(p147) join distally (the exact location of the joint depends on the species) but are separated proximally, attaching on either side of the head to the quadrate bone. the jaw muscles, which allow the bird to close its beak, attach to the proximal end of the lower mandible and to the bird's skull.(p148) the muscles that depress the lower mandible are usually weak, except in a few birds such as the starlings and the extinct huia, which have well-developed digastric muscles that aid in foraging by prying or gaping actions. in most birds, these muscles are relatively small as compared to the jaw muscles of similarly sized mammals. rhamphotheca the outer surface of the beak consists of a thin sheath of keratin called the rhamphotheca,(p148) which can be subdivided into the rhinotheca of the upper mandible and the gnathotheca of the lower mandible.(p47) this covering arises from the malpighian layer of the bird's epidermis,(p47) growing from plates at the base of each mandible. there is a vascular layer between the rhamphotheca and the deeper layers of the dermis, which is attached directly to the periosteum of the bones of the beak. the rhamphotheca grows continuously in most birds, and in some species, the color varies seasonally. in some alcids, such as the puffins, parts of the rhamphotheca are shed each year after the breeding season, while some pelicans shed a part of the bill called a "bill horn" that develops in the breeding season. while most extant birds have a single seamless rhamphotheca, species in a few families, including the albatrosses(p47) and the emu, have compound rhamphothecae that consist of several pieces separated and defined by softer keratinous grooves. studies have shown that this was the primitive ancestral state of the rhamphotheca, and that the modern simple rhamphotheca resulted from the gradual loss of the defining grooves through evolution. tomia the sawtooth serrations on a common merganser's bill help it to hold tight to its fish prey. the tomia (singular tomium) are the cutting edges of the two mandibles.(p598) in most birds, these range from rounded to slightly sharp, but some species have evolved structural modifications that allow them to handle their typical food sources better. granivorous (seed-eating) birds, for example, have ridges in their tomia, which help the bird to slice through a seed's outer hull. most falcons have a sharp projection along the upper mandible, with a corresponding notch on the lower mandible. they use this "tooth" to sever their prey's vertebrae fatally or to rip insects apart. some kites, principally those that prey on insects or lizards, also have one or more of these sharp projections, as do the shrikes. the tomial teeth of falcons are underlain by bone, while the shrike tomial teeth are entirely keratinous. some fish-eating species, e.g., the mergansers, have sawtooth serrations along their tomia, which help them to keep hold of their slippery, wriggling prey.(p48) birds in roughly 30 families have tomia lined with tight bunches of very short bristles along their entire length. most of these species are either insectivores (preferring hard-shelled prey) or snail eaters, and the brush-like projections may help to increase the coefficient of friction between the mandibles, thereby improving the bird's ability to hold hard prey items. serrations on hummingbird bills, found in 23% of all hummingbird genera, may perform a similar function, allowing the birds to effectively hold insect prey. they may also allow shorter-billed hummingbirds to function as nectar thieves, as they can more effectively hold and cut through long or waxy flower corollas. in some cases, the color of a bird's tomia can help to distinguish between similar species. the snow goose, for example, has a reddish-pink bill with black tomia, while the whole beak of the similar ross's goose is pinkish-red, without darker tomia. culmen a bird's culmen is measured in a straight line from the tip of the beak to a set point — here, where the feathering starts on the bird's forehead. the culmen is the dorsal ridge of the upper mandible.(p127) likened by ornithologist e. coues to the ridge line of a roof, it is the "highest middle lengthwise line of the bill" and runs from the point where the upper mandible emerges from the forehead's feathers to its tip.(p152) the bill's length along the culmen is one of the regular measurements made during bird banding (ringing) and is particularly useful in feeding studies. there are several standard measurements that can be made—from the beak's tip to the point where feathering starts on the forehead, from the tip to the anterior edge of the nostrils, from the tip to the base of the skull, or from the tip to the cere (for raptors and owls)(p342) — and scientists from various parts of the world generally favor one method over another. in all cases, these are chord measurements (measured in a straight line from point to point, ignoring any curve in the culmen) taken with calipers. the shape or color of the culmen can also help with the identification of birds in the field. for example, the culmen of the parrot crossbill is strongly decurved, while that of the very similar red crossbill is more moderately curved. the culmen of a juvenile common loon is all dark, while that of the very similarly plumaged juvenile yellow-billed loon is pale towards the tip. gonys the gonys is the ventral ridge of the lower mandible, created by the junction of the bone's two rami, or lateral plates.(p254) the proximal end of that junction—where the two plates separate—is known as the gonydeal angle or gonydeal expansion. in some gull species, the plates expand slightly at that point, creating a noticeable bulge; the size and shape of the gonydeal angle can be useful in identifying between otherwise similar species. adults of many species of large gulls have a reddish or orangish gonydeal spot near the gonydeal expansion. this spot triggers begging behavior in gull chicks. the chick pecks at the spot on its parent's bill, which in turn stimulates the parent to regurgitate food. commissure depending on its use, commissure may refer to the junction of the upper and lower mandibles,(p155) or alternately, to the full-length apposition of the closed mandibles, from the corners of the mouth to the tip of the beak.(p105) gape the gapes of juvenile altricial birds are often brightly coloured, as in this common starling. in bird anatomy, the gape is the interior of the open mouth of a bird, and the gape flange is the region where the two mandibles join together at the base of the beak. the width of the gape can be a factor in the choice of food. the gape flange on this juvenile house sparrow is the yellowish region at the base of the beak. gapes of juvenile altricial birds are often brightly coloured, sometimes with contrasting spots or other patterns, and these are believed to be an indication of their health, fitness and competitive ability. based on this, the parents decide how to distribute food among the chicks in the nest. some species, especially in the families viduidae and estrildidae, have bright spots on the gape known as gape tubercles or gape papillae. these nodular spots are conspicuous even in low light. a study examining the nestling gapes of eight passerine species found that the gapes were conspicuous in the ultraviolet spectrum (visible to birds but not to humans). parents may, however, not rely solely on the gape coloration, and other factors influencing their decision remain unknown. red gape color has been shown in several experiments to induce feeding. an experiment in manipulating brood size and immune system with barn swallow nestlings showed the vividness of the gape was positively correlated with t-cell–mediated immunocompetence, and that larger brood size and injection with an antigen led to a less vivid gape. conversely, the red gape of the common cuckoo (cuculus canorus) did not induce extra feeding in host parents. some brood parasites, such as the hodgson's hawk-cuckoo (c. fugax), have colored patches on the wing that mimic the gape color of the parasitized species. when born, the chick's gape flanges are fleshy. as it grows into a fledgling, the gape flanges remain somewhat swollen and can thus be used to recognize that a particular bird is young. by the time it reaches adulthood, the gape flanges will no longer be visible. nares falcons have a small tubercule within each nare. most species of birds have external nares (nostrils) located somewhere on their beak. the nares are two holes—circular, oval or slit-like in shape—which lead to the nasal cavities within the bird's skull, and thus to the rest of the respiratory system.(p375) in most bird species, the nares are located in the basal third of the upper mandible. kiwi are a notable exception; their nares are located at the tip of their bills. a handful of species have no external nares. cormorants and darters have primitive external nares as nestlings, but these close soon after the birds fledge; adults of these species (and gannets and boobies of all ages, which also lack external nostrils) breathe through their