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Many bird species migrate to take advantage of global differences of seasonal temperatures, therefore optimising availability of food sources and breeding habitat. These migrations vary among the different groups. Many landbirds, shorebirds, and waterbirds undertake annual long distance migrations, usually triggered by the length of daylight as well as weather conditions. These birds are characterised by a breeding season spent in the temperate or polar regions and a non-breeding season in the tropical regions or opposite hemisphere. Before migration, birds substantially increase body fats and reserves and reduce the size of some of their organs.[72][128] Migration is highly demanding energetically, particularly as birds need to cross deserts and oceans without refuelling. Landbirds have a flight range of around 2,500 km (1,600 mi) and shorebirds can fly up to 4,000 km (2,500 mi),[129] although the bar-tailed godwit is capable of non-stop flights of up to 10,200 km (6,300 mi).[130] Seabirds also undertake long migrations, the longest annual migration being those of sooty shearwaters, which nest in New Zealand and Chile and spend the northern summer feeding in the North Pacific off Japan, Alaska and California, an annual round trip of 64,000 km (39,800 mi).[131] Other seabirds disperse after breeding, travelling widely but having no set migration route. Albatrosses nesting in the Southern Ocean often undertake circumpolar trips between breeding seasons.[132] A map of the Pacific Ocean with several coloured lines representing bird routes running from New Zealand to Korea The routes of satellite-tagged bar-tailed godwits migrating north from New Zealand. This species has the longest known non-stop migration of any species, up to 10,200 km (6,300 mi). Some bird species undertake shorter migrations, travelling only as far as is required to avoid bad weather or obtain food. Irruptive species such as the boreal finches are one such group and can commonly be found at a location in one year and absent the next. This type of migration is normally associated with food availability.[133] Species may also travel shorter distances over part of their range, with individuals from higher latitudes travelling into the existing range of conspecifics; others undertake partial migrations, where only a fraction of the population, usually females and subdominant males, migrates.[134] Partial migration can form a large percentage of the migration behaviour of birds in some regions; in Australia, surveys found that 44% of non-passerine birds and 32% of passerines were partially migratory.[135] Altitudinal migration is a form of short distance migration in which birds spend the breeding season at higher altitudes elevations and move to lower ones during suboptimal conditions. It is most often triggered by temperature changes and usually occurs when the normal territories also become inhospitable due to lack of food.[136] Some species may also be nomadic, holding no fixed territory and moving according to weather and food availability. Parrots as a family are overwhelmingly neither migratory nor sedentary but considered to either be dispersive, irruptive, nomadic or undertake small and irregular migrations.[137] The ability of birds to return to precise locations across vast distances has been known for some time; in an experiment conducted in the 1950s a Manx shearwater released in Boston returned to its colony in Skomer, Wales, within 13 days, a distance of 5,150 km (3,200 mi).[138] Birds navigate during migration using a variety of methods. For diurnal migrants, the sun is used to navigate by day, and a stellar compass is used at night. Birds that use the sun compensate for the changing position of the sun during the day by the use of an internal clock.[57] Orientation with the stellar compass depends on the position of the constellations surrounding Polaris.[139] These are backed up in some species by their ability to sense the Earth's geomagnetism through specialised photoreceptors.[140] Communication See also: Bird vocalization Large brown patterned ground bird with outstretched wings each with a large spot in the centre The startling display of the sunbittern mimics a large predator. Birds communicate using primarily visual and auditory signals. Signals can be interspecific (between species) and intraspecific (within species). Birds sometimes use plumage to assess and assert social dominance,[141] to display breeding condition in sexually selected species, or to make threatening displays, as in the sunbittern's mimicry of a large predator to ward off hawks and protect young chicks.[142] Variation in plumage also allows for the identification of birds, particularly between species. Visual communication among birds may also involve ritualised displays, which have developed from non-signalling actions such as preening, the adjustments of feather position, pecking, or other behaviour. These displays may signal aggression or submission or may contribute to the formation of pair-bonds.[57] The most elaborate displays occur during courtship, where "dances" are often formed from complex combinations of many possible component movements;[143] males' breeding success may depend on the quality of such displays.[144] Bird song Song of the house wren, a common North American songbird Problems playing this file? See media help. Bird calls and songs, which are produced in the syrinx, are the major means by which birds communicate with sound. This communication can be very complex; some species can operate the two sides of the syrinx independently, allowing the simultaneous production of two different songs.[74] Calls are used for a variety of purposes, including mate attraction,[57] evaluation of potential mates,[145] bond formation, the claiming and maintenance of territories,[57] the identification of other individuals (such as when parents look for chicks in colonies or when mates reunite at the start of breeding season),[146] and the warning of other birds of potential predators, sometimes with specific information about the nature of the threat.[147] Some birds also use mechanical sounds for auditory communication. The Coenocorypha snipes of New Zealand drive air through their feathers,[148] woodpeckers drum territorially,[71] and palm cockatoos use tools to drum.[149] massive flock of tiny birds seen from distance so that birds appear as specks Red-billed queleas, the most numerous species of bird,[150] form enormous flocks—sometimes tens of thousands strong. Flocking and other associations While some birds are essentially territorial or live in small family groups, other birds may form large flocks. The principal benefits of flocking are safety in numbers and increased foraging efficiency.[57] Defence against predators is particularly important in closed habitats like forests, where ambush predation is common and multiple eyes can provide a valuable early warning system. This has led to the development of many mixed-species feeding flocks, which are usually composed of small numbers of many species; these flocks provide safety in numbers but increase potential competition for resources.[151] Costs of flocking include bullying of socially subordinate birds by more dominant birds and the reduction of feeding efficiency in certain cases.[152] Birds sometimes also form associations with non-avian species. Plunge-diving seabirds associate with dolphins and tuna, which push shoaling fish towards the surface.[153] Hornbills have a mutualistic relationship with dwarf mongooses, in which they forage together and warn each other of nearby birds of prey and other predators.[154] Resting and roosting Pink flamingo with grey legs and long neck pressed against body and head tucked under wings Many birds, like this American flamingo, tuck their head into their back when sleeping The high metabolic rates of birds during the active part of the day is supplemented by rest at other times. Sleeping birds often use a type of sleep known as vigilant sleep, where periods of rest are interspersed with quick eye-opening "peeks", allowing them to be sensitive to disturbances and enable rapid escape from threats.[155] Swifts are believed to be able to sleep in flight and radar observations suggest that they orient themselves to face the wind in their roosting flight.[156] It has been suggested that there may be certain kinds of sleep which are possible even when in flight.[157] Some birds have also demonstrated the capacity to fall into slow-wave sleep one hemisphere of the brain at a time. The birds tend to exercise this ability depending upon its position relative to the outside of the flock. This may allow the eye opposite the sleeping hemisphere to remain vigilant for predators by viewing the outer margins of the flock. This adaptation is also known from marine mammals.[158] Communal roosting is common because it lowers the loss of body heat and decreases the risks associated with predators.[159] Roosting sites are often chosen with regard to thermoregulation and safety.[160] Many sleeping birds bend their heads over their backs and tuck their bills in their back feathers, although others place their beaks among their breast feathers. Many birds rest on one leg, while some may pull up their legs into their feathers, especially in cold weather. Perching birds have a tendon locking mechanism that helps them hold on to the perch when they are asleep. Many ground birds, such as quails and pheasants, roost in trees. A few parrots of the genus Loriculus roost hanging upside down.[161] Some hummingbirds go into a nightly state of torpor accompanied with a reduction of their metabolic rates.[162] This physiological adaptation shows in nearly a hundred other species, including owlet-nightjars, nightjars, and woodswallows. One species, the common poorwill, even enters a state of hibernation.[163] Birds do not have sweat glands, but they may cool themselves by moving to shade, standing in water, panting, increasing their surface area, fluttering their throat or by using special behaviours like urohidrosis to cool themselves. Breeding Main category: Avian sexuality See also: Animal sexual behaviour § Birds, Seabird breeding behavior and Sexual selection in birds Social systems Bird faces up with green face, black breast and pink lower body. Elaborate long feathers on the wings and tail. Like others of its family the male Raggiana bird-of-paradise has elaborate breeding plumage used to impress females.[164] Ninety-five percent of bird species are socially monogamous. These species pair for at least the length of the breeding season or—in some cases—for several years or until the death of one mate.[165] Monogamy allows for both paternal care and biparental care, which is especially important for species in which females require males' assistance for successful brood-rearing.[166] Among many socially monogamous species, extra-pair copulation (infidelity) is common.[167] Such behaviour typically occurs between dominant males and females paired with subordinate males, but may also be the result of forced copulation in ducks and other anatids.[168] Female birds have sperm storage mechanisms that allow sperm from to remain viable long after copulation, a hundred days in some species.[169] Sperm from multiple males may compete through this mechanism. For females, possible benefits of extra-pair copulation include getting better genes for her offspring and insuring against the possibility of infertility in her mate.[170] Males of species that engage in extra-pair copulations will closely guard their mates to ensure the parentage of the offspring that they raise.[171] Other mating systems, including polygyny, polyandry, polygamy, polygynandry, and promiscuity, also occur.[57] Polygamous breeding systems arise when females are able to raise broods without the help of males.[57] Some species may use more than one system depending on the circumstances. Breeding usually involves some form of courtship display, typically performed by the male.[172] Most displays are rather simple and involve some type of song. Some displays, however, are quite elaborate. Depending on the species, these may include wing or tail drumming, dancing, aerial flights, or communal lekking. Females are generally the ones that drive partner selection,[173] although in the polyandrous phalaropes, this is reversed: plainer males choose brightly coloured females.[174] Courtship feeding, billing and allopreening are commonly performed between partners, generally after the birds have paired and mated.[71] Homosexual behaviour has been observed in males or females in numerous species of birds, including copulation, pair-bonding, and joint parenting of chicks.[175] Territories, nesting and incubation See also: Bird nest Many birds actively defend a territory from others of the same species during the breeding season; maintenance of territories protects the food source for their chicks. Species that are unable to defend feeding territories, such as seabirds and swifts, often breed in colonies instead; this is thought to offer protection from predators. Colonial breeders defend small nesting sites, and competition between and within species for nesting sites can be intense.[176] All birds lay amniotic eggs with hard shells made mostly of calcium carbonate.[57] Hole and burrow nesting species tend to lay white or pale eggs, while open nesters lay camouflaged eggs. There are many exceptions to this pattern, however; the ground-nesting nightjars have pale eggs, and camouflage is instead provided by their plumage. Species that are victims of brood parasites have varying egg colours to improve the chances of spotting a parasite's egg, which forces female parasites to match their eggs to those of their hosts.[177] Yellow weaver (bird) with black head hangs an upside-down nest woven out of grass fronds. Male golden-backed weavers construct elaborate suspended nests out of grass. Bird eggs are usually laid in a nest. Most species create somewhat elaborate nests, which can be cups, domes, plates, beds scrapes, mounds, or burrows.[178] Some bird nests, however, are extremely primitive; albatross nests are no more than a scrape on the ground. Most birds build nests in sheltered, hidden areas to avoid predation, but large or colonial birds—which are more capable of defence—may build more open nests. During nest construction, some species seek out plant matter from plants with parasite-reducing toxins to improve chick survival,[179] and feathers are often used for nest insulation.[178] Some bird species have no nests; the cliff-nesting common guillemot lays its eggs on bare rock, and male emperor penguins keep eggs between their body and feet. The absence of nests is especially prevalent in ground-nesting species where the newly hatched young are precocial. Nest made of straw with five white eggs and one grey speckled egg Nest of an eastern phoebe that has been parasitised by a brown-headed cowbird. Incubation, which optimises temperature for chick development, usually begins after the last egg has been laid.[57] In monogamous species incubation duties are often shared, whereas in polygamous species one parent is wholly responsible for incubation. Warmth from parents passes to the eggs through brood patches, areas of bare skin on the abdomen or breast of the incubating birds. Incubation can be an energetically demanding process; adult albatrosses, for instance, lose as much as 83 grams (2.9 oz) of body weight per day of incubation.[180] The warmth for the incubation of the eggs of megapodes comes from the sun, decaying vegetation or volcanic sources.[181] Incubation periods range from 10 days (in woodpeckers, cuckoos and passerine birds) to over 80 days (in albatrosses and kiwis).[57] The diversity of characteristics of birds is great, sometimes even in closely related species. Several avian characteristics are compared in the table below.[107][108] Species Adult weight (grams) Incubation (days) Clutches (per year) Clutch size Ruby-throated hummingbird (Archilochus colubris) 3 13 2.0 2 House sparrow (Passer domesticus) 25 11 4.5 5 Greater roadrunner (Geococcyx californianus) 376 20 1.5 4 Turkey vulture (Cathartes aura) 2,200 39 1.0 2 Laysan albatross (Diomedea immutabilis) 3,150 64 1.0 1 Magellanic penguin (Spheniscus magellanicus) 4,000 40 1.0 1 Golden eagle (Aquila chrysaetos) 4,800 40 1.0 2 Wild turkey (Meleagris gallopavo) 6,050 28 1.0 11 Columns are sortable Parental care and fledging At the time of their hatching, chicks range in development from helpless to independent, depending on their species. Helpless chicks are termed altricial, and tend to be born small, blind, immobile and naked; chicks that are mobile and feathered upon hatching are termed precocial. Altricial chicks need help thermoregulating and must be brooded for longer than precocial chicks. Chicks at neither of these extremes can be semi-precocial or semi-altricial. Hummingbird perched on edge of tiny nest places food into mouth of one of two chicks A female Calliope hummingbird feeding fully grown chicks. Looking down on three helpless blind chicks in a nest within the hollow of a dead tree trunk Altricial chicks of a white-breasted woodswallow. The length and nature of parental care varies widely amongst different orders and species. At one extreme, parental care in megapodes ends at hatching; the newly hatched chick digs itself out of the nest mound without parental assistance and can fend for itself immediately.[182] At the other extreme, many seabirds have extended periods of parental care, the longest being that of the great frigatebird, whose chicks take up to six months to fledge and are fed by the parents for up to an additional 14 months.[183] The chick guard stage describes the period of breeding during which one of the adult birds is permanently present at the nest after chicks have hatched. The main purpose of the guard stage is to aid offspring to thermoregulate and protect them from predation.[184] In some species, both parents care for nestlings and fledglings; in others, such care is the responsibility of only one sex. In some species, other members of the same species—usually close relatives of the breeding pair, such as offspring from previous broods—will help with the raising of the young.[185] Such alloparenting is particularly common among the Corvida, which includes such birds as the true crows, Australian magpie and fairy-wrens,[186] but has been observed in species as different as the rifleman and red kite. Among most groups of animals, male parental care is rare. In birds, however, it is quite common—more so than in any other vertebrate class.[57] Though territory and nest site defence, incubation, and chick feeding are often shared tasks, there is sometimes a division of labour in which one mate undertakes all or most of a particular duty.[187] The point at which chicks fledge varies dramatically. The chicks of the Synthliboramphus murrelets, like the ancient murrelet, leave the nest the night after they hatch, following their parents out to sea, where they are raised away from terrestrial predators.[188] Some other species, such as ducks, move their chicks away from the nest at an early age. In most species, chicks leave the nest just before, or soon after, they are able to fly. The amount of parental care after fledging varies; albatross chicks leave the nest on their own and receive no further help, while other species continue some supplementary feeding after fledging.[189] Chicks may also follow their parents during their first migration.[190] Brood parasites Main article: Brood parasite Small brown bird places an insect in the bill of much larger grey bird in nest Reed warbler raising a common cuckoo, a brood parasite. Brood parasitism, in which an egg-layer leaves her eggs with another individual's brood, is more common among birds than any other type of organism.[191] After a parasitic bird lays her eggs in another bird's nest, they are often accepted and raised by the host at the expense of the host's own brood. Brood parasites may be either obligate brood parasites, which must lay their eggs in the nests of other species because they are incapable of raising their own young, or non-obligate brood parasites, which sometimes lay eggs in the nests of conspecifics to increase their reproductive output even though they could have raised their own young.[192] One hundred bird species, including honeyguides, icterids, estrildid finches and ducks, are obligate parasites, though the most famous are the cuckoos.[191] Some brood parasites are adapted to hatch before their host's young, which allows them to destroy the host's eggs by pushing them out of the nest or to kill the host's chicks; this ensures that all food brought to the nest will be fed to the parasitic chicks.[193] Ecology Birds occupy a wide range of ecological positions.[150] While some birds are generalists, others are highly specialised in their habitat or food requirements. Even within a single habitat, such as a forest, the niches occupied by different species of birds vary, with some species feeding in the forest canopy, others beneath the canopy, and still others on the forest floor. Forest birds may be insectivores, frugivores, and nectarivores. Aquatic birds generally feed by fishing, plant eating, and piracy or kleptoparasitism. Birds of prey specialise in hunting mammals or other birds, while vultures are specialised scavengers. Avivores are animals that are specialized at predating birds. Some nectar-feeding birds are important pollinators, and many frugivores play a key role in seed dispersal.[194] Plants and pollinating birds often coevolve,[195] and in some cases a flower's primary pollinator is the only species capable of reaching its nectar.[196] Birds are often important to island ecology. Birds have frequently reached islands that mammals have not; on those islands, birds may fulfill ecological roles typically played by larger animals. For example, in New Zealand the moas were important browsers, as are the kereru and kokako today.[194] Today the plants of New Zealand retain the defensive adaptations evolved to protect them from the extinct moa.[197] Nesting seabirds may also affect the ecology of islands and surrounding seas, principally through the concentration of large quantities of guano, which may enrich the local soil[198] and the surrounding seas.[199] A wide variety of Avian ecology field methods, including counts, nest monitoring, and capturing and marking, are used for researching avian ecology. Relationship with humans Two rows of cages in a dark barn with many white chickens in each cage Industrial farming of chickens Since birds are highly visible and common animals, humans have had a relationship with them since the dawn of man.[200] Sometimes, these relationships are mutualistic, like the cooperative honey-gathering among honeyguides and African peoples such as the Borana.[201] Other times, they may be commensal, as when species such as the house sparrow[202] have benefited from human activities. Several bird species have become commercially significant agricultural pests,[203] and some pose an aviation hazard.[204] Human activities can also be detrimental, and have threatened numerous bird species with extinction (hunting, avian lead poisoning, pesticides, roadkill, and predation by pet cats and dogs are common sources of death for birds). Birds can act as vectors for spreading diseases such as psittacosis, salmonellosis, campylobacteriosis, mycobacteriosis (avian tuberculosis), avian influenza (bird flu), giardiasis, and cryptosporidiosis over long distances. Some of these are zoonotic diseases that can also be transmitted to humans.[205] Economic importance See also: Pet § Birds Domesticated birds raised for meat and eggs, called poultry, are the largest source of animal protein eaten by humans; in 2003, 76 million tons of poultry and 61 million tons of eggs were produced worldwide.[206] Chickens account for much of human poultry consumption, though domesticated turkeys, ducks, and geese are also relatively common. Many species of birds are also hunted for meat. Bird hunting is primarily a recreational activity except in extremely undeveloped areas. The most important birds hunted in North and South America are waterfowl; other widely hunted birds include pheasants, wild turkeys, quail, doves, partridge, grouse, snipe, and woodcock.[207] Muttonbirding is also popular in Australia and New Zealand.[208] Though some hunting, such as that of muttonbirds, may be sustainable, hunting has led to the extinction or endangerment of dozens of species.[209] Illustration of fisherman on raft with pole for punting and numerous black birds on raft The use of cormorants by Asian fishermen is in steep decline but survives in some areas as a tourist attraction. Other commercially valuable products from birds include feathers (especially the down of geese and ducks), which are used as insulation in clothing and bedding, and seabird feces (guano), which is a valuable source of phosphorus and nitrogen. The War of the Pacific, sometimes called the Guano War, was fought in part over the control of guano deposits.[210] Birds have been domesticated by humans both as pets and for practical purposes. Colourful birds, such as parrots and mynas, are bred in captivity or kept as pets, a practice that has led to the illegal trafficking of some endangered species.[211] Falcons and cormorants have long been used for hunting and fishing, respectively. Messenger pigeons, used since at least 1 AD, remained important as recently as World War II. Today, such activities are more common either as hobbies, for entertainment and tourism,[212] or for sports such as pigeon racing. Amateur bird enthusiasts (called birdwatchers, twitchers or, more commonly, birders) number in the millions.[213] Many homeowners erect bird feeders near their homes to attract various species. Bird feeding has grown into a multimillion dollar industry; for example, an estimated 75% of households in Britain provide food for birds at some point during the winter.[214] Religion, folklore and culture Woodcut of three long-legged and long-necked birds "The 3 of Birds" by the Master of the Playing Cards, 15th-century Germany A Swiss singing bird box, ca. 1810. An example of the interests of human beings in birds, in this case emulating nature with an automaton version. Birds play prominent and diverse roles in folklore, religion, and popular culture. In religion, birds may serve as either messengers or priests and leaders for a deity, such as in the Cult of Makemake, in which the Tangata manu of Easter Island served as chiefs;[215] or as the rooster(cock) serves as a tangible vessel of Christ as in the gospel of –Matthew, Mark and Luke in the New Testament with Christ speaking through the cock;[216] or as attendants, as in the case of Hugin and Munin, two common ravens who whispered news into the ears of the Norse god Odin.[217] In several civilizations of ancient Italy, particularly Etruscan and Roman religion, priests were involved in augury, or interpreting the words of birds while the "auspex" (from which the word "auspicious" is derived) watched their activities to foretell events.[218] They may also serve as religious symbols, as when Jonah (Hebrew: יוֹנָה, dove) embodied the fright, passivity, mourning, and beauty traditionally associated with doves.[219] Birds have themselves been deified, as in the case of the common peacock, which is perceived as Mother Earth by the Dravidians of India.[220] Some birds have also been perceived as monsters, including the mythological Roc and the Māori's legendary Pouākai, a giant bird capable of snatching humans.[221] painted tiles with design of birds from Qajar dynasty Birds have been featured in culture and art since prehistoric times, when they were represented in early cave paintings.[222] Birds were later used in religious or symbolic art and design, such as the magnificent Peacock Throne of the Mughal and Persian emperors.[223] With the advent of scientific interest in birds, many paintings of birds were commissioned for books. Among the most famous of these bird artists was John James Audubon, whose paintings of North American birds were a great commercial success in Europe and who later lent his name to the National Audubon Society.[224] Birds are also important figures in poetry; for example, Homer incorporated nightingales into his Odyssey, and Catullus used a sparrow as an erotic symbol in his Catullus 2.[225] The relationship between an albatross and a sailor is the central theme of Samuel Taylor Coleridge's The Rime of the Ancient Mariner, which led to the use of the term as a metaphor for a 'burden'.[226] Other English metaphors derive from birds; vulture funds and vulture investors, for instance, take their name from the scavenging vulture.[227] Perceptions of various bird species often vary across cultures. Owls are associated with bad luck, witchcraft, and death in parts of Africa,[228] but are regarded as wise across much of Europe.[229] Hoopoes were considered sacred in Ancient Egypt and symbols of virtue in Persia, but were thought of as thieves across much of Europe and harbingers of war in Scandinavia.[230] In religious images preserved from the Inca and Tiwanaku empires, birds are depicted in the process of transgressing boundaries between earthly and underground spiritual realms.[231] Indigenous peoples of the central Andes maintain legends of birds passing to and from metaphysical worlds.[231] The mythical chullumpi bird is said to mark the existence of a portal between such worlds, and to transform itself into a llama.[231][232] Conservation Bargraph showing estimates of the main causes of bird mortality in the US Large black bird with featherless head and hooked bill The California condor once numbered only 22 birds, but conservation measures have raised that to over 300 today. Main article: Bird conservation See also: Late Quaternary prehistoric birds, List of extinct birds and Raptor conservation Though human activities have allowed the expansion of a few species, such as the barn swallow and European starling, they have caused population decreases or extinction in many other species. Over a hundred bird species have gone extinct in historical times,[233] although the most dramatic human-caused avian extinctions, eradicating an estimated 750–1800 species, occurred during the human colonisation of Melanesian, Polynesian, and Micronesian islands.[234] Many bird populations are declining worldwide, with 1,227 species listed as threatened by Birdlife International and the IUCN in 2009.[235][236] The most commonly cited human threat to birds is habitat loss.[237] Other threats include overhunting, accidental mortality due to structural collisions or long-line fishing bycatch,[238] pollution (including oil spills and pesticide use),[239] competition and predation from nonnative invasive species,[240] and climate change. Governments and conservation groups work to protect birds, either by passing laws that preserve and restore bird habitat or by establishing captive populations for reintroductions. Such projects have produced some successes; one study estimated that conservation efforts saved 16 species of bird that would otherwise have gone extinct between 1994 and 2004, including the California condor and Norfolk parakeet.[241] |
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