What is a bird?

Birds

Everyone recognizes birds. They have feathers, wings, two legs, and a bill. Less uniquely, they have a backbone, are warm-blooded, and lay eggs. All but a few birds can fly. Birds have much in common with reptiles, from which they have evolved. They share several skeletal characteristics, nucleated red blood cells, and their young develop in cleidoic eggs. The main difference is feathers, which are modified scales. Not only do feathers allow flight, they are insulated, more so than mammalian hair, enabling birds to maintain steady internal temperatures and stay active even in extreme climates. The acquisition of flight and homeothermia has influenced the evolution of other anatomical and physiological changes in birds and led to increased cerebral and sensory development. It has freed them to travel the globe, colonizing most environments and diversifying to fill many ecological niches. Consequently, it is not surprising that birds are the most successful of the vertebrates, outnumbering the number of mammal species twofold.

Evolution and systematics

The fossil record of birds is patchy and their evolutionary history is poorly known. The first feathered animal, Archaeopteryx, has been identified in Upper Jurassic deposits, from 150 million years ago (mya). However, while it does appear intermediate between early reptiles and birds, there is some disagreement over whether it is a direct ancestor of present day birds. Fossils unequivocally of birds do not appear until the Cretaceous period, 80–120 mya, although the number of species suggests that they radiated earlier. The earliest remains are of large flightless diving birds, Hesperornis spp., with primitive teeth. Other toothed sea birds also lived during the Cretaceous, including the flighted ichthyosaurs. Also appearing in the Early Cretaceous were the Enantiornithes, a little understood group of seemingly primitive birds. At the end of the period, the toothed birds disappeared with the dinosaurs. Since then, only toothless birds have been found in the record and it is not clear how or when they arose, though it is thought that it was during the Cretaceous. By the Eocene (c. 50 mya), many modern forms were recognizable. These are non-passerines, including ostriches, penguins, storks, ducks, hawks, cuckoos, and kingfishers. The passerines (small songbirds) appear to have diversified 36–45 mya, along with flowering plants and insects. Several other forms, mostly large birds, were also present in the Eocene but died out. Other giant birds such as the larger moas of New Zealand and the elephant birds of Africa and Madagascar survived until about 10,000 years ago when they were exterminated by humans.

The evolutionary success of birds is evidenced by the wide variety of present-day forms. They have long been popular subjects of study for taxonomists. Traditional classifications are based mainly on morphological and anatomical differences in structure, plumage, and so forth. More recently, behavioral traits, song, and biochemical techniques (including DNA) have been employed. Yet, while there is general agreement as to the families to which the 9,000 or so extant bird species belong, a variety of opinions exists on the relationships within and between families.

Structure and function

General structure

Birds have adapted to a multitude of situations. For this reason, they occur in a wide diversity of shapes, sizes, and colors. Weighing up to 285 lbs (130 kg), and reaching 9 ft (2.75m), the flightless ostrich is the largest of the living birds. At a mere 0.7 oz (2 g) and around 2.4 in (6 cm), the tiny bee hummingbird is the smallest. Even closely related forms can look very different (adaptive radiation). A famous example is the enormous range of bill shapes and sizes in Charles Darwin's Galápagos finches; a single over-water colonizing species is thought to have undergone repeated evolutionary divergence to produce the 14 or so contemporary species on the different islands. Conversely, unrelated species can closely resemble each other (convergent evolution) because they have evolved for the same lifestyle. Examples of this are the Old World and New World vultures, which belong to the diurnal birds of prey and storks, respectively.

Body shapes vary enormously, from the flexible, long-necked form of the cranes and ibises to short-necked, stiff-backed falcons and penguins. These latter species, the speedy, predatory hunters of the air and the seas, have torpedo-shaped bodies to minimize drag. Bills and beaks take a variety of forms that generally reflect their major function in feeding: from the sturdy, seed-cracking bills of finches to the long, soil-probing

bill of a kiwi, the delicate curve of a nectar feeder's bill and the massive bone-shearing beak of a large vulture. In a few species, bills also serve as signals of breeding condition and sexual ornaments to attract the opposite sex. For example, the bills of cattle egrets turn from yellow to orange-yellow in the breeding season and the huge gorgeously rainbow-hued bills of the sulphur-breasted toucans may separate species. Similarly, birds' feet and legs suit their lifestyle: webbed for swimming; short and flat for ground dwellers; longer and grasping for perching species; powerful and heavily taloned for raptorial species. Stilt-like legs and spider-like toes with a span the length of the bird's body are a feature of the lily-pad walking jacanas. The legs are almost nonexistent in swifts and other birds that spend much of their lives on the wing, and long and muscular in ostriches and emus that stride and run across the plains. The ostrich has two toes, and a few species such as those that run on hard surfaces have lost the first (hind) toe or it is very small. Most species have four toes but their arrangement differs: in most perching birds, toes two, three, and four point forward and the hind toe opposes them; some species have two toes pointing forward, two back; others can move a toe to have either arrangement; swifts have all four toes pointing forward.

Wings are less variable than lower limbs, although their different forms can be extreme: much reduced in the flightless ratites, put to good use as fins in penguins, and at their most extended in gliding species that spend much of their lives riding air currents. Not only are there differences between species, variation can be quite marked within species, either geographically or between the sexes.

Species often vary in size clinally (with environmental or geographic change), usually increasing in size between from hotter to cooler parts of their range (Bergman's rule); races at either end of the cline can be remarkably different. A few species even have different forms, for example, the large- and small-beaked snail kites. Males are often larger (sexual dimorphism), but in some species, including birds of prey, some seabirds, and game birds, the female is larger.

The senses

Birds' active lifestyles require highly developed senses. For the vast majority of species, sight is the dominant sense and the eyes are relatively large. The eyes are generally set to the sides of the head, allowing a wide field of view, (about 300°), presumably useful for detecting approaching predators. For predatory birds (insectivores and raptors), the eyes are set

more forward to give a greater overlap in the field of vision of the two eyes. This increase in binocular vision is important for depth perception. Compared with mammals' eyes, birds' eyes are relatively immobile. They compensate by being able to rotate the head by as much as 270° in species such as owls, which have the most forward facing eyes. Their eyes are protected by a nictitating membrane, which closes from the inside to the outside corner, and a top and bottom eyelid. Birds can focus their eyes rapidly, which is important in flight and when diving underwater. In general, they may not have exceptional visual acuity compared with humans. However, birds have a larger field of sharp vision, good color perception, and can also discriminate in the ultraviolet part of spectrum and in polarized light. Nocturnal species have more rods than cones in their retinas to enhance their vision in dim light.

The ear of birds is simpler than that of mammals, but their sense of hearing appears to be at least as sensitive. Some species, such as some of the owls, have a disc of stiff feathers around the face that directs sound to the ears, and asymmetrically placed ear openings and enlarged inner ears to enhance discrimination of direction and distance of the source of the sound. Oilbirds and some swiftlets that live in caves use echolocation. They emit audible clicks to help them navigate and locate prey in the dark.

The great number of sensory receptors and nerve endings distributed about the body indicate that birds' sense of touch,

pain, and temperature is keen. By contrast, the olfactory system is poorly developed and few birds seem to make great use of smell. Exceptions include the New World vultures and the kiwi, which can detect prey by its scent.

Plumage

Feathers distinguish birds from all other living animals (there is recent evidence that some dinosaurs were feathered but this remains controversial). Light, strong, and colorful, feathers are extraordinarily multifunctional. They provide warmth, protection from the elements, decoration and camouflage, and are specialized for aerodynamics and flight (most birds), hydrodynamics and diving (e.g., penguins), or to cope with both elements (e.g., cormorants). A few species use them to make sound (e.g., snipe) or carry water to their young (e.g., sandgrouse). Not least, they identify species and subspecies, may vary with age, sex and breeding condition, and signal emotion.

Feathers are made of keratin and, once grown, are entirely dead tissue. They are of six main types. The most obvious are the long, stiff feathers of the wings and tail that provide the flight surfaces; more flexible, contour feathers make the sculpted outer covering for the body; and down makes a soft insulative underlayer. Semi-plumes, which are between down and contour feathers in form, help to provide insulation and fill out body contours so that air (or water) flows easily over the body. Two types of feathers are mainly sensory in function: stiff bristles that are usually found around the face (around the feet in the Tyto owls) like the whiskers of a cat or a net around the gape of some insect-eating species, and filoplumes, which are fine, hair-like feathers with a tuft of barbs at the tip that lie beside contour feathers and monitor whether the plumage is in place. In some species, modified feathers form features such as crests, ornamental bristles, cheek tufts, plumes, and tail flags and trains.

The contour, flight feathers, and semi-plumes have a central shaft, and a vane made up of barbs and barbules that interlock with each other and sometimes with neighboring feathers. The bird carefully maintains these links by nibbling and pulling the feathers through its bill. Many birds bathe regularly, most in water, but a few in dust. Some such as herons and elanine kites have powder down that grows continuously and crumbles into a fine powder that is spread through the feathers for cleaning and water resistance. Other

birds have oil glands at the base of the tail for the same purpose. Sunbathing also helps to maintain the health and curvature of feathers. Some birds, including many passerines, appear to use biting ants in feather maintenance, perhaps to control ectoparasites, either by wallowing among the swarm or by wiping individual ants through their plumage.

Over time, feathers become worn and bleached, and damaged by parasites. They are completely replaced annually in most except very large birds such as eagles and albatrosses, which spread the molt over two or so years. Some species, notably those that change from dull winter plumage into bright breeding colors (e.g., American goldfinch), have two molts a year: a full molt after breeding, and a partial body molt into breeding plumage. Most species shed their feathers sequentially to maintain their powers of flight. However, a few, particularly waterbirds that can find food in the relative safety of open water, replace all their flight feathers at once and are grounded for about five weeks.

Spectacular colors are a feature of birds, from the soft, mottled leaf patterns of nightjars (cryptic) to the gaudy, ornate plumes of a peacock (conspicuous). Cryptic colors conceal the bird from predators or rivals; conspicuous colors are used in courtship or threat. The colors themselves are produced by pigments in the feathers themselves or by structural features that interact with the pigment and the light to produce iridescent color, which can only be seen from certain angles, or non-iridescent color, that can be seen from any angle. In many species, the sexes are similar in color. In others, the sexes differ, and usually the male is showier and the female resembles a juvenile bird. In these species, sexual selection is thought to have favored dichromatism (and dimorphism) through female preference for partners with bright colors (and extravagant ornamentation). In the few polyandrous species, the reverse is that case and the females are more vibrant. Plumage may also vary geographically, with races from warm, humid climates tending to be more heavily pigmented than those from cool, dry regions (Gloger's rule).

Anatomy and physiology

The skeleto-muscular system of birds combines light weight with high power for flight. Muscle mass is concentrated near the center of gravity—around the breast and bases of the wings and legs—which gives a compact, aerodynamic form. Long tendons control movements at the ends of the limbs. Flighted birds have more massive breasts and wing muscles; in terrestrial birds, much of the muscle mass is in the upper legs. In perching birds, the tendon from the flexor muscle loops behind the ankle; when the ankle bends on landing, the toes automatically close around the perch and maintain the grip without effort, anchoring the bird even in sleep. In many species, the toe tendons have ridges, which also help to lock the feet around the perch.

In contrast to the mammalian skeleton, birds' bones are hollow and less massive and several have fused to form a strong, light frame. A bird's skeleton constitutes only about 5% of its mass. Another distinctive feature is that the bones, including the skull, are pneumatized: their core is filled with air via a system of interconnecting passages that connect with the air sacs of the respiratory system and nasal/tympanic cavities. Flighted species tend to have extensive pneumatization but it is reduced or lacking in diving birds, which would be hindered by such buoyancy. Even birds' bills are light—the horny equivalent of the heavy, toothed muzzle of mammals.

Respiration, circulation, and body temperature

Unlike mammals, birds lack a diaphragm. Instead, air is drawn into the rigid lungs by bellow-like expansion and contraction of the air sacs surrounding the lungs and another group in the head, which is driven by muscles that move the ribs and sternum up and out and back again. Features of birds' circulatory and respiratory systems make their respiration more efficient than that of most mammals, allowing them to use 25% more oxygen from each breath. This enables them to sustain a high metabolic rate and, among other benefits, assists

those high-flying migrants that cross the world's tallest mountains and reach altitudes up to 29,500 ft (9,000 m), where the oxygen content of the air is low.

Birds and mammals both generate their own body heat, but birds' high metabolic rate helps to maintain theirs at around 100°F (38–42°C), depending on species; 5–7°F (3–4°C) hotter than most mammals. When it becomes difficult to obtain enough energy to stay warm and active, a few species become torpid (lower their body temperature and become inactive) on the coldest days or during bad weather, usually for a few days or overnight. Other birds cope with cold by increasing their metabolic rate slightly, growing denser feathers, having a layer of fat, or by behavioral means such as huddling with others, tucking up a leg to decrease the heat loss surface, and fluffing out the feathers to trap more air. Lacking sweat glands to shed body heat, they may pant, lower their metabolic rate, seek shade, or raise their feathers to catch the breeze in hot weather.

Digestion and excretion

To provide the energy needed for flight, and so that they are not weighed down for a long time by the food they have

consumed, birds have a high metabolic rate and digest food rapidly and efficiently. Their digestive tract is modified accordingly, and many species have the second part of their two-part stomach modified into a muscular gizzard where hard food is physically ground down so that gastric juices can penetrate easily. Some species swallow pebbles to assist with this breakdown. The digestive tract tends to be long in grazers, fisheaters, and seed-eaters, and short in meat- and insect-eaters.

Birds have three ways to rid themselves of excess water, salts, and waste products: through breathing and the skin; the renal system; and salt glands. The salt glands are located in the orbit of each eye. They secrete sodium chloride and, therefore, are well developed in seabirds that have a salty diet, and non-functional in some other groups. Birds' kidneys are more complex than those of mammals: they produce concentrated urine with nitrogen waste in the form of insoluble uric acid, rather than urea. Such a water-efficient system does not require a (heavy) bladder, with obvious advantages for flight. It also enables many species, especially those with a moist diet (carnivores, insectivores, and frugivores), to drink seldom or not at all. Both the white urine and dark fecal matter are voided through the anus, and bird droppings often contain both.

Life history and reproduction

Life history features

To a large extent, body size determines life history. Compared with smaller species, larger species tend to live longer, breed at a later age, have a longer breeding cycle, and, at each breeding attempt, produce fewer young with a greater chance of survival. There are always exceptions, and climate and risk of predation and other factors impinge on this overlying pattern. For example, some small temperate zone Australian passerines live up to 18 years and have small clutches, whereas ground-nesting grouse may live a few years and have large clutches. For convenience, species may be classed as fast-breeders (r-selected), which have many large clutches and short periods of nestling care, and slow-breeders (K-selected) that have a few small clutches and extended periods of offspring care. In reality, there is a continuum between the two extremes.

Mating systems

The majority of bird species are at least socially monogamous, that is, a pair of birds cooperates to raise young. They may stay together for the breeding attempt or mate for life. However, many other arrangements exist. Some birds have a polygynous mating system, particularly species that use rich resources that are clumped, so that a male can support more than one female (e.g., New World blackbirds, some harriers). Successive polygyny is less common, mainly practiced by species in which the female alone raises the chicks and visits a lek where males display. The female may mate with several (e.g., black grouse and some birds of paradise) males. Less frequent again is polyandry, where the females mate with various males and leave them to care for the eggs and chicks (e.g., emus, buttonquail, and jacanas). Within these systems, there is also scope for cheating, and the advent of DNA fingerprinting has revealed that in many monogamous species, there are broods of mixed paternity. At its extreme are species such as the superb fairy-wren in which about three-quarters of the chicks are raised by males that are not the biological father.

Most birds nest as solitary pairs, but some 13% of species are colonial, particularly seabirds. Colonies may be a few pairs (e.g., king penguin) or millions (e.g., queleas). In between are species that nest in loose colonies, either regularly or when conditions are favorable. Nest spacing varies enormously from a few inches/centimeters in some colonies to several miles/kilometers in species that have large territories. Spacing is linked to food and nest sites; nests are closer together where resources are plentiful.

Breeding seasons and nests

Most birds have a regular breeding season, timed to coincide with the most abundant season of the year, but some, particularly those adapted to unpredictable climates, are opportunistic, only breeding when conditions allow. The largest species may breed every two years, but most attempt to breed at least annually, some raising several broods over the breeding season.

Birds build their nests of several substrates; the most important issues are protection from predators and the elements. Therefore, species that nest on predator-free islands often build close to the ground but, on the mainland, species build in higher locations. The nest itself must hold and shelter the eggs; in form, they vary enormously from simple scrapes in the dirt to large complex stick nests and hanging structures, woven together or glued with cobwebs or mud, and lined with soft material. Tree holes and holes in banks or cliffs also make good nests but competition for them may be fierce. The megapodes construct a mound in which they bury their eggs and maintain the temperature at about 90–95°F (32–35°C) by scratching soil on or off. Many species use the same nest area, nest site, or actual nest year after year.

Eggs and incubation

Birds' eggs are beautiful in their variety. They may be plain or colored, marked or unmarked, oval or round. All are slightly more pointed at one end so that the egg tends to roll in a circle. Species that lay in the open where eggs may roll tend to have long-oval, cryptically colored eggs, and those that lay in holes tend to have rounder, unmarked eggs. Egg laying can be energetically costly, especially for small birds: for a hummingbird,

each egg (0.01 oz/0.3g) represents 25% of the bird's mass; for an ostrich, the 50 oz (1,500 g) egg is 1% of the hen's weight. Birds that have precocial chicks tend to have large eggs (about 35% yolk compared with 20% in altricial [more helpless] species) because the chick must be advanced and well-developed when it hatches. Clutch size varies enormously both within and between species. Nevertheless, most species have a typical number of eggs; one in the kiwi, perhaps 20 in some ducks. Larger species tend to have fewer eggs. In some species, two are laid but only one ever hatches. Across species, there tends to be a trade-off between egg-size and clutch-size, some species lay a few large eggs, others many smaller eggs. In some species, the clutch size is fixed (determinate layers), in others, if the eggs are lost or removed within the breeding season, the bird will go on laying eggs (indeterminate layers). The majority of species lay every second day until the clutch is complete; in a few of the largest species, the interval is four days. In the vast majority of species incubation is carried out by one or both of the parents, but a few species, such as some ducks, nest-dump (lay some of their eggs in a neighbor's nest), and some such as the cuckoos are parasites and lay their eggs in the nest of another species. Incubation varies little within species. Among species, it ranges in length from 10 days for some woodpeckers to about 80 days for albatross and the very large-egged kiwi.

Egg formation and embryo development

To most people, eggs and birds go together. Certainly all bird species lay shelled eggs, but so too do some reptiles. In

most bird species, only the left ovary develops. The ovary holds a large number of oocytes. During the breeding season, a few oocytes (immature eggs) start to develop. They are covered in a follicle that lays down yolk, which is manufactured in the liver and carried in the blood. The one with the most yolk is shed first; the follicle ruptures and the oocyte moves into the first part of the oviduct where it may be fertilized from sperm stored in the sperm storage glands. During about 20 hours it passes down the oviduct where it is covered in albumin and, towards the end, the outer layer calcifies to form the shell and any markings are laid down from blood (red-brown) or bile (blue-green) pigments. About this time, if another egg is to be laid, another oocyte is released. In a few large species, the process may take longer.

The egg contains nutrients for the developing embryo. There is some exchange of gases and water across the shell, and waste from the embryo is stored in a sac that develops outside of the embryo, but within the shell (the allantois). Another sac develops into an air sac for ventilation. Once incubation begins, development is rapid, triggered by heat either from a brooding parent, as in most birds, or the environment, as in mound builders, which bury their eggs to capture heat from the sun. Within days, the embryo has large eyes and rudimentary organs. As it develops, the yolk sac is absorbed and the embryo fills more of the shell. By hatching, the yolk is fully absorbed and the embryo has moved its bill into the air sac and begins to breathe air. By this time, mainly through water loss, the egg is about 12–15% lighter than at the start of incubation. The embryo uses an egg tooth on the tip of its bill to chip away a ring around one end of the shell (already thinned by loss of calcium to the embryo) and hatch. If it is altricial (e.g., songbirds and seabirds), the chick will be naked or downy, and helpless; the chick of a precocial species (e.g., ostriches, ducks, and game birds) will be feathered, able to regulate its own body temperature, and can follow its parent and feed itself almost immediately. The chicks of precocial species hatch synchronously (at the same time), those of other species are sometimes asynchronous, resulting in a mixed-age brood.

Growth and care of young

Most birds grow remarkably fast and, in many altricial species, are more or less fully grown by the time they leave the nest. At the extremes, chicks stay in the nest about 10–20 days

in passerines and 150–250 days in albatrosses. Precocial species, which are free of the nest, are slower growing—their rate of growth is approximately one-third that of altricial species.

The amount and type of care given is diverse. Nidicolous species, in which the chicks stay in the nest, put in considerable effort bringing food to the nest, either bringing many small items (e.g., insectivores) or a few large ones (raptors). Some species regurgitate food for the chicks (e.g., seabirds). These species must also keep the chicks warm and dry and protect them from predators, and may also keep the nest free of droppings. Nudifugous species, in which the chicks follow adult(s), are either fed by the adult (some seabirds) or, more commonly, feed themselves on plant material (e.g., many waterfowl and game birds). Parental care varies accordingly. As protection against predators, some waterfowl carry their young on their backs and, in colonial species, several neighbors unite to try to drive off an intruder. In some species, the breeding pair is accompanied by "helpers," which may be offspring from earlier breeding attempts or unrelated individuals, males or females or both sexes, juveniles or adults. Helpers help particularly in those species in which prey is difficult to find or catch.

Ecology

All species have a niche—the range of environmental conditions under which they can survive and reproduce. The

niche may be broad and unspecialized or narrow and highly specialized. Coexisting species tend not to overlap much in their niche requirements. This is particularly obvious with food: the type and where, when, and how it is collected. For example, three hawks coexist in some Australian woodlands and they roughly segregate by species and sex: the small male sparrow hawk takes passerines in the canopy; the medium-sized brown goshawk hunts birds and small mammals in the air and on the ground in more open spaces; and the largest, the female gray goshawk, captures medium-sized birds and mammals on or near the ground.

To provide energy for flight, birds need highly nutritive food. For this reason, most birds eat at least some arthropods, especially insects. They may do this incidentally, for example, mixed in with nectar, but mostly they are actively captured. In addition to this, there are four basic diet groups: the carnivores (those that eat other vertebrates, including the fish-eaters); berry- and seedeaters; eaters of non-flowering plants (fungi, mosses, algae, and so forth); eaters of flowering plants (roots, tubers, leaves, nectar). Grasses and herbage are low in nutritive value and must be consumed in too-large quantities to be of major importance to many species, and few birds have the ability to digest cellulose. Not surprisingly, birds are pollinators and seed dispersers for a great variety of plants. Some species specialize, others are more varied in their diet. Methods of collecting food are numerous, but most involve sight.

Birds are infected by a variety of diseases, both internal and external, caused by bacteria, viruses, and parasites. Some of these can also affect humans and their livestock. In general, healthy birds can carry both internal and external parasites without obvious harm. Nevertheless, disease outbreaks occur; for example, following floods, outbreaks of insect-borne pox can occur in wild birds. Birds can also suffer from exposure to toxic substances of both natural (e.g., toxic algal blooms) and human-made origin (e.g., oil spills, pesticides, lead-shot, and other pollutants). Predation is a fact of life for the majority of bird species, particularly for their eggs and young. Their life history has evolved to allow for losses, which are naturally high. For example, in passerines, perhaps 5% of eggs result in adult birds and annual mortality of adults may be as high as 60%. For larger species, the rate of mortality tends to be lower. Provided that they are not too prolonged or severe, starvation, predation, disease, and other causes of

mortality are compensatory and, in general, bird populations recover quickly.

Distribution and biogeography

Birds are distributed across all continents and on most islands, and in all major habitats from caves to mountaintops, deserts to rainforests. Many factors limit the distribution of species, all relating to their ecology: climate, habitat availability, and the presence of predators, competitors, and food. There may also be physical barriers such as mountain ranges, oceans, and impassable expanses of unsuitable habitat. The breeding range often differs from the nonbreeding range because of seasonal or other movements to remain in the most favorable conditions. There are broad patterns to general distribution; for example, woodpeckers are found in many regions of the world but are absent from Australia, New Zealand, and Madagascar. The ratites are southern in distribution, pointing to their early evolution in Gondwana (the huge southern super-continent that split into the southern continents). These patterns reveal six major biogeographical realms: Neotropical, Nearctic, Ethiopian, Palearctic, Oriental, and Australian. The greatest number of species is found in the Neotropics (South and Central America, the West Indies, and southern Mexico) with roughly 3,000 species, and 31 endemic families. The Nearctic (North America, Greenland, Iceland), with about 1,000 breeding species and no endemic bird families, is among those areas with the fewest species.

Bird populations vary enormously in their abundance and density. Some species are widely scattered across the landscape (e.g., the solitary eagles), others live in crowded colonies of millions (queleas). There are several general patterns: population density is related to body size (smaller species tend to be more numerous); the number of species and of individuals tends to be greatest in complex habitats (e.g., forest compared with grassland) and where several habitats meet (ecotones); the number of species increases with the size of the habitat patch; and, the number of species and overall densities tend to increase from the poles to the equator.

Behavior

Bird species can be social or solitary; they may nest, roost, and feed in small or large groups for part of their lives (e.g., when young or in the nonbreeding season) or their entire lives. Some form feeding flocks with other bird species or nest near more powerful species for protection. Some associate loosely with other vertebrates, such as following monkey troops to catch the insects they flush. A few live more or less commensally, for example, by feeding on ticks from ungulates. Some species spend much of their lives on the ground (terrestrial species), others in the air, in water, or in various combinations. The majority of species are active by day (diurnal), but many are crepuscular (active in twilight) or truly nocturnal. During inactive periods, most retire to a safe roost where they socialize, preen, relax, or sleep. They appear to need to sleep several hours a day.

Birds have developed complex communication systems, including various calls and song and visual signals involving posture, movements, facial expressions, display of certain characteristics of plumage, and, in some species, flushing of the skin (e.g., vultures) or popping of the eyes (Australian choughs). Some behaviors, like the dance of brolgas and sychronized swimming of swans, is ritualized, others appear more spontaneous. Courtship is a highly ritualized sequence of displays, on the ground or in the air, and can involve courtship feeding, which may be a way for females to judge the quality of their partner.

As a group, birds are cerebrally advanced. Much of their behavior is innate but they also learn by experience throughout life. They have good memories, such as retrieving cached food items from several stores several weeks after they were hidden. Some species appear to be particularly adaptable if not intelligent. Certainly, their behavior can be complex and interesting. Individual rooks wait to cache nuts if a non-hoarding rook is passing by, yet are unconcerned by rooks that are also hoarding. Jays that steal food are more likely to move their caches to prevent theft than are jays that are not thieves. Parrots have a complex social system and enjoy playing with each other and with found objects. Black kites and green-backed herons have learned to place bread on water to attract fish. One of the Galápagos finches uses a cactus thorn to probe bark crevices for insects that it cannot reach with its short tongue.

Resources

Books

Brooke, M., and T. Birkhead. The Cambridge Encyclopedia of Ornithology. Cambridge: Cambridge University Press, 1991.

Campbell, B., and E. Lack, eds. A Dictionary of Birds. Calton: Poyser, 1985.

de Juana, E. Handbook of the Birds of the World. Vol. 1. Ostrich to Ducks, edited by J. A. del Hoyo, A. Elliot, and J. Sargatal. Barcelona: Lynx Edicions, 1992.

Farner, D. S., J. R. King, and K. C. Parkes, eds. Avian Biology. Vols. 6, 7, and 8. London: Academic Press, 1982 through 1985.

Farner, D. S., and J. R. King, eds. Avian Biology. Vols. 1, 2, 3, 4, and 5. London: Academic Press, 1971 through 1975.

Perrins, C. M. The Illustrated Encyclopedia of Birds. London: Headline, 1990.

Perrins, C. M., and T. R. Birkhead. Avian Ecology. Glasgow: Blackie, 1983.

Welty, J. C., and L. Baptista. The Life of Birds. 4th ed. London: Poyser, 1988.

Organizations

American Ornithologists' Union. Suite 402, 1313 Dolley Madison Blvd, McLean, VA 22101 USA. E-mail: [email protected] Web site: <www.aou.org>

BirdLife International. Wellbrook Court, Girton Road, Cambridge, Cambridgeshire CB3 ONA United Kingdom. Phone: +44 1 223 277 318. Fax: +44-1-223-277-200. E-mail: [email protected] Web site: <http://www.birdlife.net>

Birds Australia. 415 Riversdale Road, Hawthorn East, Victoria 3123 Australia. Phone: +61 3 9882 2622. Fax: +61 3 98822677. E-mail: [email protected] Web site: <http://www.birdsaustralia.com.au>

British Trust for Ornithology. The Nunnery, Thetford, Norfolk IP24 2PU United Kingdom. Phone: +44 0 1842750050. Fax: +44 0 1842 750030. E-mail: [email protected] Web site: <www.bto.org>

National Audubon Society. 700 Broadway, New York, NY 10003 USA. Phone: (212) 979-3000. Fax: (212) 978-3188. Web site: <www.Audubon.org>

Royal Society for the Protection of Birds. Admail 975 Freepost ANG 6335, The Lodge, Sandy, Bedfordshire SG19 2TN United Kingdom. Web site: <www.rspb.org.uk>

Penny Olsen, PhD