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Birds are a group of
warm-blooded Warm-blooded is an informal term referring to animal species which can maintain a body temperature higher than their environment. In particular, homeothermic species maintain a stable body temperature by regulating metabolic processes. The on ...
vertebrates constituting the
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differently ...
Aves (), characterised by feathers, toothless beaked jaws, the
laying Laying is the act of making equipment level. It usually involves moving equipment in small motions so that spirit levels are centralised in all planes. Movement is usually done by small worm gears or other fine setting devices for accurate s ...
of hard-shelled eggs, a high metabolic rate, a four-chambered
heart The heart is a muscular Organ (biology), organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as ca ...
, and a strong yet lightweight skeleton. Birds live worldwide and range in size from the bee hummingbird to the
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There ...
. There are about ten thousand living species, more than half of which are
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
, or "perching" birds. Birds have whose development varies according to species; the only known groups without wings are the extinct moa and elephant birds. Wings, which are modified forelimbs, gave birds the ability to fly, although further evolution has led to the loss of flight in some birds, including ratites,
penguin Penguins (order Sphenisciformes , family Spheniscidae ) are a group of aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: only one species, the Galápagos penguin, is found north of the Equator. Highly adap ...
s, and diverse
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
island species. The digestive and respiratory systems of birds are also uniquely adapted for flight. Some bird species of aquatic environments, particularly
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same envir ...
s and some waterbirds, have further evolved for swimming. Birds are feathered theropod
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s and constitute the only known living dinosaurs. Likewise, birds are considered reptiles in the modern cladistic sense of the term, and their closest living relatives are the
crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest livi ...
ns. Birds are descendants of the primitive
avialans Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally used ...
(whose members include ''
Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
'') which first appeared about 160 million years ago (mya) in China. According to DNA evidence, modern birds (Neornithes) evolved in the Middle to
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
, and diversified dramatically around the time of the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
66 mya, which killed off the
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
s and all non-avian dinosaurs. Many social species pass on knowledge across generations, which is considered a form of culture. Birds are social, communicating with visual signals, calls, and songs, and participating in such behaviours as cooperative breeding and hunting, flocking, and mobbing of predators. The vast majority of bird species are socially (but not necessarily sexually) monogamous, usually for one breeding season at a time, sometimes for years, and rarely for life. Other species have breeding systems that are polygynous (one male with many females) or, rarely, polyandrous (one female with many males). Birds produce offspring by laying eggs which are fertilised through
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
. They are usually laid in a nest and incubated by the parents. Most birds have an extended period of parental care after hatching. Many species of birds are economically important as food for human consumption and raw material in manufacturing, with domesticated and undomesticated birds being important sources of eggs, meat, and feathers. Songbirds, parrots, and other species are popular as pets. Guano (bird excrement) is harvested for use as a fertiliser. Birds figure throughout human culture. About 120 to 130 species have become
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
due to human activity since the 17th century, and hundreds more before then. Human activity threatens about 1,200 bird species with extinction, though efforts are underway to protect them. Recreational birdwatching is an important part of the
ecotourism Ecotourism is a form of tourism involving responsible travel (using sustainable transport) to natural areas, conserving the environment, and improving the well-being of the local people. Its purpose may be to educate the traveler, to provide fund ...
industry.


Evolution and classification

The first classification of birds was developed by Francis Willughby and John Ray in their 1676 volume ''Ornithologiae''.
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
modified that work in 1758 to devise the taxonomic classification system currently in use. Birds are categorised as the biological class Aves in
Linnaean taxonomy Linnaean taxonomy can mean either of two related concepts: # The particular form of biological classification (taxonomy) set up by Carl Linnaeus, as set forth in his ''Systema Naturae'' (1735) and subsequent works. In the taxonomy of Linnaeus t ...
. Phylogenetic taxonomy places Aves in the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
Theropoda.


Definition

Aves and a sister group, the order
Crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest livi ...
, contain the only living representatives of the reptile clade Archosauria. During the late 1990s, Aves was most commonly defined phylogenetically as all descendants of the most recent common ancestor of modern birds and '' Archaeopteryx lithographica''. However, an earlier definition proposed by Jacques Gauthier gained wide currency in the 21st century, and is used by many scientists including adherents to the '' PhyloCode''. Gauthier defined Aves to include only the crown group of the set of modern birds. This was done by excluding most groups known only from
fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
, and assigning them, instead, to the broader group Avialae, in part to avoid the uncertainties about the placement of ''Archaeopteryx'' in relation to animals traditionally thought of as theropod dinosaurs. Gauthier and de Queiroz identified four different definitions for the same biological name "Aves", which is a problem. The authors proposed to reserve the term Aves only for the crown group consisting of the last common ancestor of all living birds and all of its descendants, which corresponds to meaning number 4 below. He assigned other names to the other groups. # Aves can mean all
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
s closer to birds than to crocodiles (alternately Avemetatarsalia) # Aves can mean those advanced archosaurs with feathers (alternately Avifilopluma) # Aves can mean those feathered dinosaurs that fly (alternately Avialae) # Aves can mean the last common ancestor of all the currently living birds and all of its descendants (a " crown group", in this sense synonymous with Neornithes) Under the fourth definition ''Archaeopteryx'', traditionally considered one of the earliest members of Aves, is removed from this group, becoming a non-avian dinosaur instead. These proposals have been adopted by many researchers in the field of palaeontology and
bird evolution Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweight ...
, though the exact definitions applied have been inconsistent. Avialae, initially proposed to replace the traditional fossil content of Aves, is often used synonymously with the vernacular term "bird" by these researchers. Most researchers define Avialae as branch-based clade, though definitions vary. Many authors have used a definition similar to "all theropods closer to birds than to '' Deinonychus''", with '' Troodon'' being sometimes added as a second external specifier in case it is closer to birds than to ''Deinonychus''. Avialae is also occasionally defined as an apomorphy-based clade (that is, one based on physical characteristics). Jacques Gauthier, who named Avialae in 1986, re-defined it in 2001 as all dinosaurs that possessed feathered wings used in flapping flight, and the birds that descended from them. Despite being currently one of the most widely used, the crown-group definition of Aves has been criticised by some researchers. Lee and Spencer (1997) argued that, contrary to what Gauthier defended, this definition would not increase the stability of the clade and the exact content of Aves will always be uncertain because any defined clade (either crown or not) will have few synapomorphies distinguishing it from its closest relatives. Their alternative definition is synonymous to Avifilopluma.


Dinosaurs and the origin of birds

Based on fossil and biological evidence, most scientists accept that birds are a specialised subgroup of theropod
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s and, more specifically, members of Maniraptora, a group of theropods which includes
dromaeosaurids Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
and oviraptorosaurs, among others. As scientists have discovered more theropods closely related to birds, the previously clear distinction between non-birds and birds has become blurred. Recent discoveries in the Liaoning Province of northeast China, which demonstrate many small theropod
feathered dinosaur A feathered dinosaur is any species of dinosaur possessing feathers. While this includes all species of birds, there is a hypothesis that many, if not all non-avian dinosaur species also possessed feathers in some shape or form. It has been s ...
s, contribute to this ambiguity. The consensus view in contemporary palaeontology is that the flying theropods, or
avialans Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally used ...
, are the closest relatives of the deinonychosaurs, which include dromaeosaurids and troodontids. Together, these form a group called Paraves. Some basal members of Deinonychosauria, such as ''
Microraptor ''Microraptor'' ( Greek, μικρός, ''mīkros'': "small"; Latin, ''raptor'': "one who seizes") is a genus of small, four-winged dromaeosaurid dinosaurs. Numerous well-preserved fossil specimens have been recovered from Liaoning, China. ...
'', have features which may have enabled them to glide or fly. The most basal deinonychosaurs were very small. This evidence raises the possibility that the ancestor of all paravians may have been arboreal, have been able to glide, or both. Unlike ''Archaeopteryx'' and the non-avialan feathered dinosaurs, who primarily ate meat, recent studies suggest that the first avialans were omnivores. The Late Jurassic ''Archaeopteryx'' is well known as one of the first transitional fossils to be found, and it provided support for the
theory of evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
in the late 19th century. ''Archaeopteryx'' was the first fossil to display both clearly traditional reptilian characteristics—teeth, clawed fingers, and a long, lizard-like tail—as well as wings with flight feathers similar to those of modern birds. It is not considered a direct ancestor of birds, though it is possibly closely related to the true ancestor.


Early evolution

Over 40% of key traits found in modern birds evolved during the 60 million year transition from the earliest bird-line archosaurs to the first maniraptoromorphs, i.e. the first dinosaurs closer to living birds than to '' Tyrannosaurus rex''. The loss of osteoderms otherwise common in archosaurs and acquisition of primitive feathers might have occurred early during this phase. After the appearance of Maniraptoromorpha, the next 40 million years marked a continuous reduction of body size and the accumulation of
neotenic Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compare ...
(juvenile-like) characteristics. Hypercarnivory became increasingly less common while braincases enlarged and forelimbs became longer. The integument evolved into complex, pennaceous feathers. The oldest known paravian (and probably the earliest avialan) fossils come from the Tiaojishan Formation of China, which has been dated to the late
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
period ( Oxfordian stage), about 160 million years ago. The avialan species from this time period include '' Anchiornis huxleyi'', ''
Xiaotingia zhengi ''Xiaotingia'' is a genus of anchiornithid theropod dinosaur from Middle Jurassic or early Late Jurassic deposits of western Liaoning, China, containing a single species, ''Xiaotingia zhengi''. Discovery ''Xiaotingia'' is known from the holoty ...
'', and '' Aurornis xui''. The well-known probable early avialan, ''Archaeopteryx'', dates from slightly later Jurassic rocks (about 155 million years old) from
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwee ...
. Many of these early avialans shared unusual anatomical features that may be ancestral to modern birds but were later lost during bird evolution. These features include enlarged claws on the second toe which may have been held clear of the ground in life, and long feathers or "hind wings" covering the hind limbs and feet, which may have been used in aerial manoeuvreing. Avialans diversified into a wide variety of forms during the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
period. Many groups retained primitive characteristics, such as clawed wings and teeth, though the latter were lost independently in a number of avialan groups, including modern birds (Aves). Increasingly stiff tails (especially the outermost half) can be seen in the evolution of maniraptoromorphs, and this process culminated in the appearance of the pygostyle, an ossification of fused tail vertebrae. In the late Cretaceous, about 100 million years ago, the ancestors of all modern birds evolved a more open pelvis, allowing them to lay larger eggs compared to body size. Around 95 million years ago, they evolved a better sense of smell. A third stage of bird evolution starting with Ornithothoraces (the "bird-chested" avialans) can be associated with the refining of aerodynamics and flight capabilities, and the loss or co-ossification of several skeletal features. Particularly significant are the development of an enlarged, keeled sternum and the alula, and the loss of grasping hands.


Early diversity of bird ancestors

The first large, diverse lineage of short-tailed avialans to evolve were the Enantiornithes, or "opposite birds", so named because the construction of their shoulder bones was in reverse to that of modern birds. Enantiornithes occupied a wide array of
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s, from sand-probing shorebirds and fish-eaters to tree-dwelling forms and seed-eaters. While they were the dominant group of avialans during the Cretaceous period, enantiornithes became extinct along with many other dinosaur groups at the end of the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
era. Many species of the second major avialan lineage to diversify, the Euornithes (meaning "true birds", because they include the ancestors of modern birds), were semi-aquatic and specialised in eating fish and other small aquatic organisms. Unlike the Enantiornithes, which dominated land-based and arboreal habitats, most early euornithes lacked perching adaptations and seem to have included shorebird-like species, waders, and swimming and diving species. The latter included the superficially
gull Gulls, or colloquially seagulls, are seabirds of the family Laridae in the suborder Lari. They are most closely related to the terns and skimmers and only distantly related to auks, and even more distantly to waders. Until the 21st century ...
-like '' Ichthyornis'' and the Hesperornithiformes, which became so well adapted to hunting fish in marine environments that they lost the ability to fly and became primarily aquatic. The early euornithes also saw the development of many traits associated with modern birds, like strongly keeled breastbones, toothless, beaked portions of their jaws (though most non-avian euornithes retained teeth in other parts of the jaws). Euornithes also included the first avialans to develop true pygostyle and a fully mobile fan of tail feathers, which may have replaced the "hind wing" as the primary mode of aerial maneuverability and braking in flight. A study on mosaic evolution in the avian skull found that the last common ancestor of all Neornithes might have had a beak similar to that of the modern hook-billed vanga and a skull similar to that of the Eurasian golden oriole. As both species are small aerial and canopy foraging omnivores, a similar ecological niche was inferred for this hypothetical ancestor.


Diversification of modern birds

All modern birds lie within the crown group Aves (alternately Neornithes), which has two subdivisions: the Palaeognathae, which includes the flightless ratites (such as the
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There ...
es) and the weak-flying
tinamou Tinamous () form an order of birds called Tinamiformes (), comprising a single family called Tinamidae (), divided into two distinct subfamilies, containing 46 species found in Mexico, Central America, and South America. The word "tinamou" co ...
s, and the extremely diverse Neognathae, containing all other birds. These two subdivisions have variously been given the rank of superorder, cohort, or infraclass. Depending on the taxonomic viewpoint, the number of known living bird species varies anywhere from 9,800 to 10,758. The discovery of '' Vegavis'' from the
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the inte ...
, the last stage of the Late Cretaceous proved that the diversification of modern birds started before the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configu ...
era. The affinities of an earlier fossil, the possible galliform '' Austinornis lentus'', dated to about 85 million years ago, are still too controversial to provide a fossil evidence of modern bird diversification. In 2020, '' Asteriornis'' from the Maastrichtian was described, it appears to be a close relative of Galloanserae, the earliest diverging lineage within Neognathae. Most studies agree on a Cretaceous age for the most recent common ancestor of modern birds but estimates range from the Early Cretaceous to the latest
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
. Similarly, there is no agreement on whether most of the early diversification of modern birds occurred before or after the Cretaceous–Palaeogene extinction event. This disagreement is in part caused by a divergence in the evidence; most molecular dating studies suggests a Cretaceous evolutionary radiation, while fossil evidence points to a Cenozoic radiation (the so-called 'rocks' versus 'clocks' controversy). Previous attempts to reconcile molecular and fossil evidence have proved controversial, but more recent estimates, using a more comprehensive sample of fossils and a new way of calibrating molecular clocks, showed that while according to some studies, modern birds originated early in the Late Cretaceous in Western
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final sta ...
, a pulse of diversification in all major groups occurred around the Cretaceous–Palaeogene extinction event. Modern birds expanded from West Gondwana to the Laurasia through two routes. One route was an Antarctic interchange in the Paleogene. This can be confirmed with the presence of multiple avian groups in Australia and New Zealand. The other route was probably through North America, via land bridges, during the Paleocene. This allowed the expansion and diversification of Neornithes into the Holarctic and Paleotropics. On the other hand, the occurrence of ''Asteriornis'' in the Northern Hemisphere challenges biogeographical hypotheses of a Gondwanan origin of crown birds.


Classification of bird orders

Cladogram of modern bird relationships based on Braun & Kimball (2021) The classification of birds is a contentious issue. Sibley and Ahlquist's ''Phylogeny and Classification of Birds'' (1990) is a landmark work on the subject. Most evidence seems to suggest the assignment of orders is accurate, but scientists disagree about the relationships among the orders themselves; evidence from modern bird anatomy, fossils and DNA have all been brought to bear on the problem, but no strong consensus has emerged. More recently, new fossil and molecular evidence is providing an increasingly clear picture of the evolution of modern bird orders.


Genomics

, the
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
had been sequenced for only two birds, the
chicken The chicken (''Gallus gallus domesticus'') is a domesticated junglefowl species, with attributes of wild species such as the grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster or cock is a term for an adu ...
and the zebra finch. the genomes of 542 species of birds had been completed. At least one genome has been sequenced from every order. These include at least one species in about 90% of extant avian families (218 out of 236 families recognised by the ''Howard and Moore Checklist''). Being able to sequence and compare whole genomes gives researchers many types of information, about genes, the DNA that regulates the genes, and their evolutionary history. This has led to reconsideration of some of the classifications that were based solely on the identification of protein-coding genes. Waterbirds such as pelicans and flamingos, for example, may have in common specific adaptations suited to their environment that were developed independently.


Distribution

Birds live and breed in most terrestrial habitats and on all seven continents, reaching their southern extreme in the
snow petrel The snow petrel (''Pagodroma nivea'') is the only member of the genus ''Pagodroma.'' It is one of only three birds that has been seen at the Geographic South Pole, along with the Antarctic petrel and the south polar skua, which have the most s ...
's breeding colonies up to inland in
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
. The highest bird diversity occurs in tropical regions. It was earlier thought that this high diversity was the result of higher speciation rates in the tropics; however recent studies found higher speciation rates in the high latitudes that were offset by greater
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the Endling, last individual of the species, although the Functional ext ...
rates than in the tropics. Many species migrate annually over great distances and across oceans; several families of birds have adapted to life both on the world's oceans and in them, and some
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same envir ...
species come ashore only to breed, while some
penguin Penguins (order Sphenisciformes , family Spheniscidae ) are a group of aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: only one species, the Galápagos penguin, is found north of the Equator. Highly adap ...
s have been recorded diving up to deep. Many bird species have established breeding populations in areas to which they have been introduced by humans. Some of these introductions have been deliberate; the ring-necked pheasant, for example, has been introduced around the world as a game bird. Others have been accidental, such as the establishment of wild monk parakeets in several North American cities after their escape from captivity. Some species, including
cattle egret The cattle egret (''Bubulcus ibis'') is a cosmopolitan species of heron ( family Ardeidae) found in the tropics, subtropics, and warm-temperate zones. It is the only member of the monotypic genus ''Bubulcus'', although some authorities regard ...
, yellow-headed caracara and
galah The galah (; ''Eolophus roseicapilla''), also known as the pink and grey cockatoo or rose-breasted cockatoo, is the only species within genus ''Eolophus'' of the cockatoo family. Found throughout Australia, it is among the most common of the ...
, have spread naturally far beyond their original ranges as agricultural expansion created alternative habitats although modern practices of intensive agriculture have negatively impacted farmland bird populations.


Anatomy and physiology

Compared with other vertebrates, birds have a body plan that shows many unusual adaptations, mostly to facilitate flight.


Skeletal system

The skeleton consists of very lightweight bones. They have large air-filled cavities (called pneumatic cavities) which connect with the
respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies g ...
. The skull bones in adults are fused and do not show cranial sutures. The orbital cavities that house the eyeballs are large and separated from each other by a bony septum (partition). The
spine Spine or spinal may refer to: Science Biology * Vertebral column, also known as the backbone * Dendritic spine, a small membranous protrusion from a neuron's dendrite * Thorns, spines, and prickles, needle-like structures in plants * Spine (zoolo ...
has cervical, thoracic, lumbar and caudal regions with the number of cervical (neck) vertebrae highly variable and especially flexible, but movement is reduced in the anterior thoracic vertebrae and absent in the later vertebrae. The last few are fused with the pelvis to form the synsacrum. The ribs are flattened and the sternum is keeled for the attachment of flight muscles except in the flightless bird orders. The forelimbs are modified into wings. The wings are more or less developed depending on the species; the only known groups that lost their wings are the
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
moa and elephant birds.


Excretory system

Like the reptiles, birds are primarily uricotelic, that is, their
kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blo ...
s extract nitrogenous waste from their bloodstream and excrete it as uric acid, instead of urea or
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous ...
, through the ureters into the intestine. Birds do not have a urinary bladder or external urethral opening and (with exception of the
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There ...
) uric acid is excreted along with faeces as a semisolid waste. However, birds such as hummingbirds can be facultatively ammonotelic, excreting most of the nitrogenous wastes as ammonia. They also excrete creatine, rather than creatinine like mammals. This material, as well as the output of the intestines, emerges from the bird's cloaca. The cloaca is a multi-purpose opening: waste is expelled through it, most birds mate by joining cloaca, and females lay eggs from it. In addition, many species of birds regurgitate pellets. It is a common but not universal feature of altricial
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
nestlings (born helpless, under constant parental care) that instead of excreting directly into the nest, they produce a fecal sac. This is a mucus-covered pouch that allows parents to either dispose of the waste outside the nest or to recycle the waste through their own digestive system.


Reproductive system

Males within Palaeognathae (with the exception of the kiwis), the Anseriformes (with the exception of screamers), and in rudimentary forms in
Galliformes Galliformes is an order of heavy-bodied ground-feeding birds that includes turkeys, chickens, quail, and other landfowl. Gallinaceous birds, as they are called, are important in their ecosystems as seed dispersers and predators, and are ofte ...
(but fully developed in Cracidae) possess a
penis A penis (plural ''penises'' or ''penes'' () is the primary sexual organ that male animals use to inseminate females (or hermaphrodites) during copulation. Such organs occur in many animals, both vertebrate and invertebrate, but males d ...
, which is never present in Neoaves. The length is thought to be related to sperm competition. For male birds to get an erection, they depends on lymphatic fluid instead of blood. When not copulating, it is hidden within the
proctodeum A proctodeum is the back ectodermal part of an alimentary canal. It is created during embryogenesis by a folding of the outer body wall. It will form the lower part of the anal canal, below the pectinate line The pectinate line (dentate line) i ...
compartment within the cloaca, just inside the vent. Female birds have sperm storage tubules that allow sperm to remain viable long after copulation, a hundred days in some species. Sperm from multiple males may compete through this mechanism. Most female birds have a single ovary and a single oviduct, both on the left side, but there are exceptions: species in at least 16 different orders of birds have two ovaries. Even these species, however, tend to have a single oviduct. It has been speculated that this might be an adaptation to flight, but males have two testes, and it is also observed that the gonads in both sexes decrease dramatically in size outside the breeding season. Also terrestrial birds generally have a single ovary, as does the
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or mono ...
, an egg-laying mammal. A more likely explanation is that the egg develops a shell while passing through the oviduct over a period of about a day, so that if two eggs were to develop at the same time, there would be a risk to survival. While rare, mostly abortive,
parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and developmen ...
is not unknown in birds and eggs can be diploid, automictic and results in male offspring. Birds are solely gonochoric. Meaning they have two sexes: either
female Female ( symbol: ♀) is the sex of an organism that produces the large non-motile ova (egg cells), the type of gamete (sex cell) that fuses with the male gamete during sexual reproduction. A female has larger gametes than a male. Fema ...
or
male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
. The sex of birds is determined by the Z and W sex chromosomes, rather than by the X and Y chromosomes present in
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s. Male birds have two Z chromosomes (ZZ), and female birds have a W chromosome and a Z chromosome (WZ). In nearly all species of birds, an individual's sex is determined at fertilisation. However, one 2007 study claimed to demonstrate temperature-dependent sex determination among the Australian brushturkey, for which higher temperatures during incubation resulted in a higher female-to-male
sex ratio The sex ratio (or gender ratio) is usually defined as the ratio of males to females in a population. As explained by Fisher's principle, for evolutionary reasons this is typically about 1:1 in species which reproduce sexually. Many species dev ...
. This, however, was later proven to not be the case. These birds do not exhibit temperature-dependent sex determination, but temperature-dependent sex mortality.


Respiratory and circulatory systems

Birds have one of the most complex
respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies g ...
s of all animal groups. Upon inhalation, 75% of the fresh air bypasses the lungs and flows directly into a posterior air sac which extends from the lungs and connects with air spaces in the bones and fills them with air. The other 25% of the air goes directly into the lungs. When the bird exhales, the used air flows out of the lungs and the stored fresh air from the posterior air sac is simultaneously forced into the lungs. Thus, a bird's lungs receive a constant supply of fresh air during both inhalation and exhalation. Sound production is achieved using the syrinx, a muscular chamber incorporating multiple tympanic membranes which diverges from the lower end of the trachea; the trachea being elongated in some species, increasing the volume of vocalisations and the perception of the bird's size. In birds, the main arteries taking blood away from the heart originate from the right
aortic arch The aortic arch, arch of the aorta, or transverse aortic arch () is the part of the aorta between the ascending and descending aorta. The arch travels backward, so that it ultimately runs to the left of the trachea. Structure The aorta begins ...
(or pharyngeal arch), unlike in the mammals where the left aortic arch forms this part of the aorta. The postcava receives blood from the limbs via the renal portal system. Unlike in mammals, the circulating red blood cells in birds retain their
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: * Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucl ...
.


Heart type and features

The avian circulatory system is driven by a four-chambered, myogenic heart contained in a fibrous pericardial sac. This pericardial sac is filled with a serous fluid for lubrication. The heart itself is divided into a right and left half, each with an atrium and ventricle. The atrium and ventricles of each side are separated by atrioventricular valves which prevent back flow from one chamber to the next during contraction. Being myogenic, the heart's pace is maintained by pacemaker cells found in the sinoatrial node, located on the right atrium. The sinoatrial node uses calcium to cause a depolarising signal transduction pathway from the atrium through right and left atrioventricular bundle which communicates contraction to the ventricles. The avian heart also consists of muscular arches that are made up of thick bundles of muscular layers. Much like a mammalian heart, the avian heart is composed of endocardial, myocardial and
epicardial The pericardium, also called pericardial sac, is a double-walled sac containing the heart and the roots of the great vessels. It has two layers, an outer layer made of strong connective tissue (fibrous pericardium), and an inner layer made o ...
layers. The atrium walls tend to be thinner than the ventricle walls, due to the intense ventricular contraction used to pump oxygenated blood throughout the body. Avian hearts are generally larger than mammalian hearts when compared to body mass. This adaptation allows more blood to be pumped to meet the high metabolic need associated with flight.


Organisation

Birds have a very efficient system for diffusing oxygen into the blood; birds have a ten times greater surface area to
gas exchange Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a ...
volume than mammals. As a result, birds have more blood in their capillaries per unit of volume of lung than a mammal. The arteries are composed of thick elastic muscles to withstand the pressure of the ventricular contractions, and become more rigid as they move away from the heart. Blood moves through the arteries, which undergo
vasoconstriction Vasoconstriction is the narrowing of the blood vessels resulting from contraction of the muscular wall of the vessels, in particular the large arteries and small arterioles. The process is the opposite of vasodilation, the widening of blood vess ...
, and into arterioles which act as a transportation system to distribute primarily oxygen as well as nutrients to all tissues of the body. As the arterioles move away from the heart and into individual organs and tissues they are further divided to increase surface area and slow blood flow. Blood travels through the arterioles and moves into the capillaries where gas exchange can occur. Capillaries are organised into capillary beds in tissues; it is here that blood exchanges oxygen for carbon dioxide waste. In the capillary beds, blood flow is slowed to allow maximum
diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical ...
of oxygen into the tissues. Once the blood has become deoxygenated, it travels through venules then veins and back to the heart. Veins, unlike arteries, are thin and rigid as they do not need to withstand extreme pressure. As blood travels through the venules to the veins a funneling occurs called vasodilation bringing blood back to the heart. Once the blood reaches the heart, it moves first into the right atrium, then the right ventricle to be pumped through the lungs for further gas exchange of carbon dioxide waste for oxygen. Oxygenated blood then flows from the lungs through the left atrium to the left ventricle where it is pumped out to the body.


Nervous system

The
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes ...
is large relative to the bird's size. The most developed part of the brain is the one that controls the flight-related functions, while the cerebellum coordinates movement and the
cerebrum The cerebrum, telencephalon or endbrain is the largest part of the brain containing the cerebral cortex (of the two cerebral hemispheres), as well as several subcortical structures, including the hippocampus, basal ganglia, and olfactory bulb. ...
controls behaviour patterns, navigation, mating and nest building. Most birds have a poor sense of smell with notable exceptions including kiwis,
New World vulture The New World vulture or condor family, Cathartidae, contains seven extant species in five genera. It includes five extant vultures and two extant condors found in warm and temperate areas of the Americas. The "New World" vultures were widespr ...
s and
tubenoses Procellariiformes is an order of seabirds that comprises four families: the albatrosses, the petrels and shearwaters, and two families of storm petrels. Formerly called Tubinares and still called tubenoses in English, procellariiforms are oft ...
. The avian visual system is usually highly developed. Water birds have special flexible lenses, allowing accommodation for vision in air and water. Some species also have dual
fovea Fovea () (Latin for "pit"; plural foveae ) is a term in anatomy. It refers to a pit or depression in a structure. Human anatomy *Fovea centralis of the retina * Fovea buccalis or Dimple * Fovea of the femoral head * Trochlear fovea of the fr ...
. Birds are tetrachromatic, possessing
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiation ...
(UV) sensitive cone cells in the eye as well as green, red and blue ones. They also have double cones, likely to mediate achromatic vision. Many birds show plumage patterns in
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiation ...
that are invisible to the human eye; some birds whose sexes appear similar to the naked eye are distinguished by the presence of ultraviolet reflective patches on their feathers. Male blue tits have an ultraviolet reflective crown patch which is displayed in courtship by posturing and raising of their nape feathers. Ultraviolet light is also used in foraging— kestrels have been shown to search for prey by detecting the UV reflective urine trail marks left on the ground by rodents. With the exception of pigeons and a few other species, the eyelids of birds are not used in blinking. Instead the eye is lubricated by the nictitating membrane, a third eyelid that moves horizontally. The nictitating membrane also covers the eye and acts as a contact lens in many aquatic birds. The bird
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which the ...
has a fan shaped blood supply system called the pecten. Eyes of most birds are large, not very round and capable of only limited movement in the orbits, typically 10–20°. Birds with eyes on the sides of their heads have a wide
visual field The visual field is the "spatial array of visual sensations available to observation in introspectionist psychological experiments". Or simply, visual field can be defined as the entire area that can be seen when an eye is fixed straight at a poin ...
, while birds with eyes on the front of their heads, such as owls, have binocular vision and can estimate the depth of field. The avian ear lacks external pinnae but is covered by feathers, although in some birds, such as the '' Asio'', '' Bubo'' and ''
Otus Otus may refer to: * Otus (education), a K-12 educational technology company * HMS ''Otus'', two ships in the Royal Navy * ''Otus'' (bird), genus of owls * Otus (mythology), giant in Greek mythology, brother of Ephialtes, one of Aloadae * Otus ...
'' owls, these feathers form tufts which resemble ears. The inner ear has a
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus. A core component of the cochlea is the Organ of Corti, the sensory o ...
, but it is not spiral as in mammals.


Defence and intraspecific combat

A few species are able to use chemical defences against predators; some
Procellariiformes Procellariiformes is an order of seabirds that comprises four families: the albatrosses, the petrels and shearwaters, and two families of storm petrels. Formerly called Tubinares and still called tubenoses in English, procellariiforms are oft ...
can eject an unpleasant stomach oil against an aggressor, and some species of pitohuis from
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; id, Papua, or , historically ) is the world's second-largest island with an area of . Located in Oceania in the southwestern Pacific Ocean, the island is separated from Australia by the wide Torres ...
have a powerful neurotoxin in their skin and feathers. A lack of field observations limit our knowledge, but intraspecific conflicts are known to sometimes result in injury or death. The screamers ( Anhimidae), some jacanas ('' Jacana'', '' Hydrophasianus''), the spur-winged goose ('' Plectropterus''), the torrent duck ('' Merganetta'') and nine species of lapwing ('' Vanellus'') use a sharp spur on the wing as a weapon. The steamer ducks ('' Tachyeres''), geese and swans (''
Anserinae The Anserinae are a subfamily in the waterfowl family Anatidae. It includes the swans and true geese. Under alternative systematical concepts (see e.g., Terres & NAS, 1991), it is split into two subfamilies, the Anserinae contain the geese ...
''), the solitaire ('' Pezophaps''), sheathbills ('' Chionis''), some guans ('' Crax'') and stone curlews ('' Burhinus'') use a bony knob on the alular metacarpal to punch and hammer opponents. The jacanas '' Actophilornis'' and '' Irediparra'' have an expanded, blade-like radius. The extinct '' Xenicibis'' was unique in having an elongate forelimb and massive hand which likely functioned in combat or defence as a jointed club or flail. Swans, for instance, may strike with the bony spurs and bite when defending eggs or young.


Feathers, plumage, and scales

Feathers are a feature characteristic of birds (though also present in some dinosaurs not currently considered to be true birds). They facilitate flight, provide insulation that aids in
thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperatur ...
, and are used in display, camouflage, and signalling. There are several types of feathers, each serving its own set of purposes. Feathers are epidermal growths attached to the skin and arise only in specific tracts of skin called pterylae. The distribution pattern of these feather tracts (pterylosis) is used in taxonomy and systematics. The arrangement and appearance of feathers on the body, called plumage, may vary within species by age, social status, and sex. Plumage is regularly moulted; the standard plumage of a bird that has moulted after breeding is known as the "" plumage, or—in the
Humphrey–Parkes terminology Humphrey–Parkes terminology is a system of nomenclature for the plumage of birds. Before the Humphrey–Parkes system, plumage was named after the belief that a certain plumage was breeding plumage and others were not. However, as this system d ...
—"basic" plumage; breeding plumages or variations of the basic plumage are known under the Humphrey–Parkes system as "" plumages. Moulting is annual in most species, although
some Some may refer to: *''some'', an English word used as a determiner and pronoun; see use of ''some'' *The term associated with the existential quantifier *"Some", a song by Built to Spill from their 1994 album ''There's Nothing Wrong with Love'' * ...
may have two moults a year, and large birds of prey may moult only once every few years. Moulting patterns vary across species. In passerines, flight feathers are replaced one at a time with the innermost being the first. When the fifth of sixth primary is replaced, the outermost begin to drop. After the innermost tertiaries are moulted, the starting from the innermost begin to drop and this proceeds to the outer feathers (centrifugal moult). The greater primary are moulted in synchrony with the primary that they overlap. A small number of species, such as ducks and geese, lose all of their flight feathers at once, temporarily becoming flightless. As a general rule, the tail feathers are moulted and replaced starting with the innermost pair. Centripetal moults of tail feathers are however seen in the Phasianidae. The centrifugal moult is modified in the tail feathers of woodpeckers and treecreepers, in that it begins with the second innermost pair of feathers and finishes with the central pair of feathers so that the bird maintains a functional climbing tail. The general pattern seen in
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
s is that the primaries are replaced outward, secondaries inward, and the tail from centre outward. Before nesting, the females of most bird species gain a bare
brood patch A brood patch is a patch of featherless skin on the underside of birds during the nesting season. Feathers act as inherent insulators, and prevent efficient incubation. Birds have solved this evolutionary dilemma by developing dedicated brood pat ...
by losing feathers close to the belly. The skin there is well supplied with blood vessels and helps the bird in incubation. Feathers require maintenance and birds preen or groom them daily, spending an average of around 9% of their daily time on this. The bill is used to brush away foreign particles and to apply waxy secretions from the uropygial gland; these secretions protect the feathers' flexibility and act as an
antimicrobial agent An antimicrobial is an agent that kills microorganisms or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ...
, inhibiting the growth of feather-degrading
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
. This may be supplemented with the secretions of formic acid from ants, which birds receive through a behaviour known as anting, to remove feather parasites. The
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
of birds are composed of the same keratin as beaks, claws, and spurs. They are found mainly on the toes and metatarsus, but may be found further up on the ankle in some birds. Most bird scales do not overlap significantly, except in the cases of kingfishers and woodpeckers. The scales of birds are thought to be homologous to those of reptiles and mammals.


Flight

Most birds can
fly Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
, which distinguishes them from almost all other vertebrate classes. Flight is the primary means of locomotion for most bird species and is used for searching for food and for escaping from predators. Birds have various adaptations for flight, including a lightweight skeleton, two large flight muscles, the pectoralis (which accounts for 15% of the total mass of the bird) and the supracoracoideus, as well as a modified forelimb (
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is e ...
) that serves as an aerofoil. Wing shape and size generally determine a bird's flight style and performance; many birds combine powered, flapping flight with less energy-intensive soaring flight. About 60 extant bird species are flightless, as were many extinct birds. Flightlessness often arises in birds on isolated islands, most likely due to limited resources and the absence of
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
ian land predators. Flightlessness is almost exclusively correlated with gigantism due to an island's inheren condition of isolation. Although flightless, penguins use similar musculature and movements to "fly" through the water, as do some flight-capable birds such as
auk An auk or alcid is a bird of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The word "auk" is derived from Icelandic ''álka'', from Old Norse ''alka'' (a ...
s, shearwaters and dippers.


Behaviour

Most birds are diurnal, but some birds, such as many species of owls and nightjars, are nocturnal or crepuscular (active during twilight hours), and many coastal waders feed when the tides are appropriate, by day or night.


Diet and feeding

are varied and often include nectar, fruit, plants, seeds, carrion, and various small animals, including other birds. The
digestive system of birds Bird anatomy, or the physiological structure of birds' bodies, shows many unique adaptations, mostly aiding flight. Birds have a light skeletal system and light but powerful musculature which, along with circulatory and respiratory systems capabl ...
is unique, with a
crop A crop is a plant that can be grown and harvested extensively for profit or subsistence. When the plants of the same kind are cultivated at one place on a large scale, it is called a crop. Most crops are cultivated in agriculture or hydropon ...
for storage and a gizzard that contains swallowed stones for grinding food to compensate for the lack of teeth. Some species such as pigeons and some psittacine species do not have a gallbladder. Most birds are highly adapted for rapid digestion to aid with flight. Some migratory birds have adapted to use protein stored in many parts of their bodies, including protein from the intestines, as additional energy during migration. (Erratum in ''Proceedings of the Royal Society B'' 267(1461):2567.) Birds that employ many strategies to obtain food or feed on a variety of food items are called generalists, while others that concentrate time and effort on specific food items or have a single strategy to obtain food are considered specialists.
Avian foraging Avian foraging refers to the range of activities and behaviours exhibited by birds in their quest for food. In addition to their unique body adaptations, birds have a range of described behaviours that differ from the foraging behaviours of other a ...
strategies can vary widely by species. Many birds glean for insects, invertebrates, fruit, or seeds. Some hunt insects by suddenly attacking from a branch. Those species that seek pest
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s are considered beneficial 'biological control agents' and their presence encouraged in biological pest control programmes. Combined, insectivorous birds eat 400–500 million metric tons of arthropods annually. Nectar feeders such as hummingbirds, sunbirds, lories, and lorikeets amongst others have specially adapted brushy tongues and in many cases bills designed to fit
co-adapted In biology, co-adaptation is the process by which two or more species, genes or phenotypic traits undergo adaptation as a pair or group. This occurs when two or more interacting characteristics undergo natural selection together in response to the s ...
flowers. Kiwis and
shorebird 245px, A flock of Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflat ...
s with long bills probe for invertebrates; shorebirds' varied bill lengths and feeding methods result in the separation of
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s. Loons, diving ducks,
penguin Penguins (order Sphenisciformes , family Spheniscidae ) are a group of aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: only one species, the Galápagos penguin, is found north of the Equator. Highly adap ...
s and auks pursue their prey underwater, using their wings or feet for propulsion, while aerial predators such as sulids, kingfishers and terns plunge dive after their prey. Flamingos, three species of prion, and some ducks are filter feeders. Geese and dabbling ducks are primarily grazers. Some species, including
frigatebird Frigatebirds are a family of seabirds called Fregatidae which are found across all tropical and subtropical oceans. The five extant species are classified in a single genus, ''Fregata''. All have predominantly black plumage, long, deeply forke ...
s,
gull Gulls, or colloquially seagulls, are seabirds of the family Laridae in the suborder Lari. They are most closely related to the terns and skimmers and only distantly related to auks, and even more distantly to waders. Until the 21st century ...
s, and
skua The skuas are a group of predatory seabirds with seven species forming the genus ''Stercorarius'', the only genus in the family Stercorariidae. The three smaller skuas, the long-tailed skua, the Arctic skua, and the pomarine skua are calle ...
s, engage in
kleptoparasitism Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when ...
, stealing food items from other birds. Kleptoparasitism is thought to be a supplement to food obtained by hunting, rather than a significant part of any species' diet; a study of great frigatebirds stealing from masked boobies estimated that the frigatebirds stole at most 40% of their food and on average stole only 5%. Other birds are scavengers; some of these, like vultures, are specialised carrion eaters, while others, like gulls, corvids, or other birds of prey, are opportunists.


Water and drinking

Water is needed by many birds although their mode of excretion and lack of
sweat gland Sweat glands, also known as sudoriferous or sudoriparous glands, , are small tubular structures of the skin that produce sweat. Sweat glands are a type of exocrine gland, which are glands that produce and secrete substances onto an epithelial ...
s reduces the physiological demands. Some desert birds can obtain their water needs entirely from moisture in their food. They may also have other adaptations such as allowing their body temperature to rise, saving on moisture loss from evaporative cooling or panting. Seabirds can drink seawater and have salt glands inside the head that eliminate excess salt out of the nostrils. Most birds scoop water in their beaks and raise their head to let water run down the throat. Some species, especially of arid zones, belonging to the pigeon, finch, mousebird, button-quail and
bustard Bustards, including floricans and korhaans, are large, terrestrial birds living mainly in dry grassland areas and on the steppes of the Old World. They range in length from . They make up the family Otididae (, formerly known as Otidae). Bust ...
families are capable of sucking up water without the need to tilt back their heads. Some desert birds depend on water sources and
sandgrouse Sandgrouse is the common name for Pteroclidae , a family of sixteen species of bird, members of the order Pterocliformes . They are traditionally placed in two genera. The two central Asian species are classified as ''Syrrhaptes'' and the oth ...
are particularly well known for their daily congregations at waterholes. Nesting sandgrouse and many plovers carry water to their young by wetting their belly feathers. Some birds carry water for chicks at the nest in their crop or regurgitate it along with food. The pigeon family, flamingos and penguins have adaptations to produce a nutritive fluid called crop milk that they provide to their chicks.


Feather care

Feathers, being critical to the survival of a bird, require maintenance. Apart from physical wear and tear, feathers face the onslaught of fungi, ectoparasitic feather mites and
bird lice A bird louse is any chewing louse (small, biting insects) of order Phthiraptera which parasitizes warm-blooded animals, especially birds. Bird lice may feed on feathers, skin, or blood. They have no wings, and their biting mouth parts distin ...
. The physical condition of feathers are maintained by often with the application of secretions from the . Birds also bathe in water or dust themselves. While some birds dip into shallow water, more aerial species may make aerial dips into water and arboreal species often make use of dew or rain that collect on leaves. Birds of arid regions make use of loose soil to dust-bathe. A behaviour termed as anting in which the bird encourages ants to run through their plumage is also thought to help them reduce the ectoparasite load in feathers. Many species will spread out their wings and expose them to direct sunlight and this too is thought to help in reducing fungal and ectoparasitic activity that may lead to feather damage.


Migration

Many bird species migrate to take advantage of global differences of
season A season is a division of the year based on changes in weather, ecology, and the number of daylight hours in a given region. On Earth, seasons are the result of the axial parallelism of Earth's tilted orbit around the Sun. In temperate and ...
al temperatures, therefore optimising availability of food sources and breeding habitat. These migrations vary among the different groups. Many landbirds,
shorebird 245px, A flock of Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflat ...
s, and
waterbird A water bird, alternatively waterbird or aquatic bird, is a bird that lives on or around water. In some definitions, the term ''water bird'' is especially applied to birds in freshwater ecosystems, although others make no distinction from sea ...
s 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 The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
regions or opposite hemisphere. Before migration, birds substantially increase body fats and reserves and reduce the size of some of their organs. (Erratum in ''Proceedings of the Royal Society B'' 267(1461):2567.) Migration is highly demanding energetically, particularly as birds need to cross deserts and oceans without refuelling. Landbirds have a flight range of around and shorebirds can fly up to , although the bar-tailed godwit is capable of non-stop flights of up to .
Seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same envir ...
s also undertake long migrations, the longest annual migration being those of sooty shearwaters, which nest in
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island coun ...
and
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the eas ...
and spend the northern summer feeding in the North Pacific off Japan,
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U ...
and
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
, an annual round trip of . Other seabirds disperse after breeding, travelling widely but having no set migration route. Albatrosses nesting in the
Southern Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the World Ocean, generally taken to be south of 60° S latitude and encircling Antarctica. With a size of , it is regarded as the second-smal ...
often undertake circumpolar trips between breeding seasons. 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. 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. 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.
Altitudinal migration Altitudinal migration is a short-distance animal migration from lower altitudes to higher altitudes and back. Altitudinal migrants change their elevation with the seasons making this form of animal migration seasonal. Altitudinal migration can be m ...
is a form of short-distance migration in which birds spend the breeding season at higher altitudes 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. Some species may also be nomadic, holding no fixed territory and moving according to weather and food availability. Parrots as a
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
are overwhelmingly neither migratory nor sedentary but considered to either be dispersive, irruptive, nomadic or undertake small and irregular migrations. 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 Boston (), officially the City of Boston, is the state capital and most populous city of the Commonwealth of Massachusetts, as well as the cultural and financial center of the New England region of the United States. It is the 24th- mo ...
in the United States returned to its colony in Skomer, in Wales within 13 days, a distance of . 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. Orientation with the stellar compass depends on the position of the constellations surrounding Polaris. These are backed up in some species by their ability to sense the Earth's geomagnetism through specialised photoreceptors.


Communication

Birds
communicate Communication (from la, communicare, meaning "to share" or "to be in relation with") is usually defined as the transmission of information. The term may also refer to the message communicated through such transmissions or the field of inqu ...
primarily using visual and auditory signals. Signals can be interspecific (between species) and intraspecific (within species). Birds sometimes use plumage to assess and assert social dominance, 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. 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. The most elaborate displays occur during courtship, where "dances" are often formed from complex combinations of many possible component movements; males' breeding success may depend on the quality of such displays. Bird calls and songs, which are produced in the syrinx, are the major means by which birds communicate with
sound In physics, sound is a vibration that propagates as an acoustic wave, through a transmission medium such as a gas, liquid or solid. In human physiology and psychology, sound is the ''reception'' of such waves and their ''perception'' by ...
. 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. Calls are used for a variety of purposes, including mate attraction, evaluation of potential mates, bond formation, the claiming and maintenance of territories, the identification of other individuals (such as when parents look for chicks in colonies or when mates reunite at the start of breeding season), and the warning of other birds of potential predators, sometimes with specific information about the nature of the threat. Some birds also use mechanical sounds for auditory communication. The ''
Coenocorypha The austral snipes, also known as the New Zealand snipes or tutukiwi, are a genus, ''Coenocorypha'', of tiny birds in the sandpiper family, which are now only found on New Zealand's outlying islands. There are currently three living species an ...
''
snipe A snipe is any of about 26 wading bird species in three genera in the family Scolopacidae. They are characterized by a very long, slender bill, eyes placed high on the head, and cryptic/ camouflaging plumage. The '' Gallinago'' snipes have ...
s of
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island coun ...
drive air through their feathers, woodpeckers drum for long-distance communication, and palm cockatoos use tools to drum.


Flocking and other associations

While some birds are essentially territorial or live in small family groups, other birds may form large
flocks Flocking is the behaviour exhibited when a group of birds, called a flock, are foraging or in flight. Computer simulations and mathematical models that have been developed to emulate the flocking behaviours of birds can also generally be applie ...
. The principal benefits of flocking are
safety in numbers Safety in numbers is the hypothesis that, by being part of a large physical group or mass, an individual is less likely to be the victim of a mishap, accident, attack, or other bad event. Some related theories also argue (and can show statistica ...
and increased foraging efficiency. Defence against predators is particularly important in closed habitats like forests, where
ambush predation Ambush predators or sit-and-wait predators are carnivorous animals that capture or trap prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey ...
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. Costs of flocking include bullying of socially subordinate birds by more dominant birds and the reduction of feeding efficiency in certain cases. Birds sometimes also form associations with non-avian species. Plunge-diving
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same envir ...
s associate with
dolphin A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae (the b ...
s and
tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max length: ...
, which push shoaling fish towards the surface. Some species of 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.


Resting and roosting

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. 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. It has been suggested that there may be certain kinds of sleep which are possible even when in flight. Some birds have also demonstrated the capacity to fall into slow-wave sleep one
hemisphere Hemisphere refers to: * A half of a sphere As half of the Earth * A hemisphere of Earth ** Northern Hemisphere ** Southern Hemisphere ** Eastern Hemisphere ** Western Hemisphere ** Land and water hemispheres * A half of the (geocentric) celes ...
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
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s by viewing the outer margins of the flock. This adaptation is also known from
marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their ...
s.
Communal roosting Communal roosting is an animal behavior where a group of individuals, typically of the same species, congregate in an area for a few hours based on an external signal and will return to the same site with the reappearance of the signal. Environment ...
is common because it lowers the loss of body heat and decreases the risks associated with predators. Roosting sites are often chosen with regard to thermoregulation and safety. Unusual mobile roost sites include large herbivores on the African savanna that are used by oxpeckers. 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. Some hummingbirds go into a nightly state of
torpor Torpor is a state of decreased physiological activity in an animal, usually marked by a reduced body temperature and metabolic rate. Torpor enables animals to survive periods of reduced food availability. The term "torpor" can refer to the time ...
accompanied with a reduction of their metabolic rates. 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. Birds do not have sweat glands, but can lose water directly through the skin, and they may cool themselves by moving to shade, standing in water, panting, increasing their surface area, fluttering their throat or using special behaviours like
urohidrosis Urohidrosis (sometimes misspelled "urohydrosis") is the habit in some birds of defecating onto the scaly portions of the legs as a cooling mechanism, using evaporative cooling of the fluids. Several species of storks and New World vultures exhibit ...
to cool themselves.


Breeding


Social systems

Ninety-five per cent 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. Monogamy allows for both paternal care and biparental care, which is especially important for species in which care from both the female and the male parent is required in order to successfully rear a brood. Among many socially monogamous species, extra-pair copulation (infidelity) is common. 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. 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. Males of species that engage in extra-pair copulations will closely guard their mates to ensure the parentage of the offspring that they raise. Other mating systems, including
polygyny Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
,
polyandry Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives" ...
,
polygamy Crimes Polygamy (from Late Greek (') "state of marriage to many spouses") is the practice of marrying multiple spouses. When a man is married to more than one wife at the same time, sociologists call this polygyny. When a woman is marr ...
, polygynandry, and promiscuity, also occur. Polygamous breeding systems arise when females are able to raise broods without the help of males. Mating systems vary across bird families but variations within species are thought to be driven by environmental conditions. Breeding usually involves some form of courtship display, typically performed by the male. Most displays are rather simple and involve some type of
song A song is a musical composition intended to be performed by the human voice. This is often done at distinct and fixed pitches (melodies) using patterns of sound and silence. Songs contain various forms, such as those including the repetiti ...
. Some displays, however, are quite elaborate. Depending on the species, these may include wing or tail drumming, dancing, aerial flights, or communal
lekking A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate. A lek can also indicate an avail ...
. Females are generally the ones that drive partner selection, although in the polyandrous
phalaropes __NOTOC__ A phalarope is any of three living species of slender-necked shorebirds in the genus ''Phalaropus'' of the bird family Scolopacidae. Phalaropes are close relatives of the shanks and tattlers, the ''Actitis'' and Terek sandpipers, a ...
, this is reversed: plainer males choose brightly coloured females.
Courtship feeding A nuptial gift is a nutritional gift given by one partner in some animals' sexual reproduction practices. Formally, a nuptial gift is a material presentation to a recipient by a donor during or in relation to sexual intercourse that is not simpl ...
, billing and are commonly performed between partners, generally after the birds have paired and mated. Homosexual behaviour has been observed in males or females in numerous species of birds, including copulation, pair-bonding, and joint parenting of chicks. Over 130 avian species around the world engage in sexual interactions between the same sex or homosexual behaviours. "Same-sex courtship activities may involve elaborate displays, synchronized dances, gift-giving ceremonies, or behaviors at specific display areas including bowers, arenas, or leks."


Territories, nesting and incubation

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
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same envir ...
s and swifts, often breed in
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
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. All birds lay amniotic eggs with hard shells made mostly of calcium carbonate. Hole and burrow nesting species tend to lay white or pale eggs, while open nesters lay
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
d 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 Brood parasites are animals that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its o ...
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. Bird eggs are usually laid in a nest. Most species create somewhat elaborate nests, which can be cups, domes, plates, mounds, or burrows. Some bird nests can be a simple scrape, with minimal or no lining; most seabird and wader 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, and feathers are often used for nest insulation. 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 open habitat ground-nesting species where any addition of nest material would make the nest more conspicuous. Many ground nesting birds lay a clutch of eggs that hatch synchronously, with precocial chicks led away from the nests (
nidifugous In biology, nidifugous ( , ) organisms are those that leave the nest shortly after hatching or birth. The term is derived from Latin ''nidus'' for "nest" and ''fugere'', meaning "to flee". The terminology is most often used to describe birds and ...
) by their parents soon after hatching. Incubation, which regulates temperature for chick development, usually begins after the last egg has been laid. 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 patch A brood patch is a patch of featherless skin on the underside of birds during the nesting season. Feathers act as inherent insulators, and prevent efficient incubation. Birds have solved this evolutionary dilemma by developing dedicated brood pat ...
es, 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 of body weight per day of incubation. The warmth for the incubation of the eggs of megapodes comes from the sun, decaying vegetation or volcanic sources. Incubation periods range from 10 days (in woodpeckers,
cuckoo Cuckoos are birds in the Cuculidae family, the sole taxon in the order Cuculiformes . The cuckoo family includes the common or European cuckoo, roadrunners, koels, malkohas, couas, coucals and anis. The coucals and anis are sometimes separ ...
s and
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
birds) to over 80 days (in albatrosses and kiwis). The diversity of characteristics of birds is great, sometimes even in closely related species. Several avian characteristics are compared in the table below.


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. The young of many bird species do not precisely fit into either the precocial or altricial category, having some aspects of each and thus fall somewhere on an "altricial-precocial spectrum". Chicks at neither extreme but favouring one or the other may be termed or . 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. 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. 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.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 Other often refers to: * Other (philosophy), a concept in psychology and philosophy Other or The Other may also refer to: Film and television * ''The Other'' (1913 film), a German silent film directed by Max Mack * ''The Other'' (1930 film), a ...
of the same species—usually close relatives of the
breeding pair Breeding pair is a pair of animals which cooperate over time to produce offspring with some form of a bond between the individuals.Gaston, A. J.The evolution of group territorial behavior and cooperative breeding" The American Naturalist 112.988 ...
, such as offspring from previous broods—will help with the raising of the young. Such alloparenting is particularly common among the Corvida, which includes such birds as the true crows, Australian magpie and fairy-wrens, 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. Although 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. The point at which chicks fledge varies dramatically. The chicks of the ''
Synthliboramphus ''Synthliboramphus'' is a small genus of seabirds in the auk family from the North Pacific. The genus name ''Synthliboramphus'' is from Ancient Greek ''sunthlibo'', "to compress", and ''rhamphos'', "bill". The English name "Murrelet" is a dim ...
'' 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. 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. Chicks may also follow their parents during their first migration.


Brood parasites

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. 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. One hundred bird species, including
honeyguide Honeyguides (family Indicatoridae) are near passerine birds in the order Piciformes. They are also known as indicator birds, or honey birds, although the latter term is also used more narrowly to refer to species of the genus ''Prodotiscus''. Th ...
s, icterids, and ducks, are obligate parasites, though the most famous are the
cuckoo Cuckoos are birds in the Cuculidae family, the sole taxon in the order Cuculiformes . The cuckoo family includes the common or European cuckoo, roadrunners, koels, malkohas, couas, coucals and anis. The coucals and anis are sometimes separ ...
s. 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.


Sexual selection

Birds have
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variati ...
a variety of mating behaviours, with the
peacock Peafowl is a common name for three bird species in the genera '' Pavo'' and '' Afropavo'' within the tribe Pavonini of the family Phasianidae, the pheasants and their allies. Male peafowl are referred to as peacocks, and female peafowl are r ...
tail being perhaps the most famous example of
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...
and the Fisherian runaway. Commonly occurring
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
s such as size and colour differences are energetically costly attributes that signal competitive breeding situations. Many types of avian
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...
have been identified; intersexual selection, also known as female choice; and intrasexual competition, where individuals of the more abundant sex compete with each other for the privilege to mate. Sexually selected traits often evolve to become more pronounced in competitive breeding situations until the trait begins to limit the individual's fitness. Conflicts between an individual fitness and signalling adaptations ensure that sexually selected ornaments such as plumage colouration and courtship behaviour are "honest" traits. Signals must be costly to ensure that only good-quality individuals can present these exaggerated sexual ornaments and behaviours.


Inbreeding depression

Inbreeding causes early death (
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
) in the zebra finch ''Taeniopygia guttata''. Embryo survival (that is, hatching success of fertile eggs) was significantly lower for sib-sib mating pairs than for unrelated pairs. Darwin's finch ''Geospiza scandens'' experiences
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
(reduced survival of offspring) and the magnitude of this effect is influenced by environmental conditions such as low food availability.


Inbreeding avoidance

Incestuous matings by the purple-crowned fairy wren ''Malurus coronatus'' result in severe fitness costs due to
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
(greater than 30% reduction in hatchability of eggs). Females paired with related males may undertake extra pair matings (see Promiscuity#Other animals for 90% frequency in avian species) that can reduce the negative effects of inbreeding. However, there are ecological and demographic constraints on extra pair matings. Nevertheless, 43% of broods produced by incestuously paired females contained extra pair young. Inbreeding depression occurs in the great tit (''Parus major'') when the offspring produced as a result of a mating between close relatives show reduced fitness. In natural populations of ''Parus major'', inbreeding is avoided by dispersal of individuals from their birthplace, which reduces the chance of mating with a close relative.
Southern pied babbler The southern pied babbler (''Turdoides bicolor'') is a species of bird in the family Leiothrichidae, found in dry savannah of Botswana, Namibia, South Africa, and Zimbabwe. Description The southern pied babbler is a medium-sized cooperatively b ...
s ''Turdoides bicolor'' appear to avoid inbreeding in two ways. The first is through dispersal, and the second is by avoiding familiar group members as mates. Cooperative breeding in birds typically occurs when offspring, usually males, delay dispersal from their natal group in order to remain with the family to help rear younger kin. Female offspring rarely stay at home, dispersing over distances that allow them to breed independently, or to join unrelated groups. In general, inbreeding is avoided because it leads to a reduction in progeny fitness (
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
) due largely to the homozygous expression of deleterious recessive alleles. Cross-fertilisation between unrelated individuals ordinarily leads to the masking of deleterious recessive alleles in progeny.


Ecology

Birds occupy a wide range of ecological positions. 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, or nectarivores. Aquatic birds generally feed by fishing, plant eating, and piracy or
kleptoparasitism Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when ...
. Many grassland birds are granivores. Birds of prey specialise in hunting mammals or other birds, while vultures are specialised scavengers. Birds are also preyed upon by a range of mammals including a few avivorous bats. A wide range of endo- and ectoparasites depend on birds and some parasites that are transmitted from parent to young have co-evolved and show host-specificity. Some nectar-feeding birds are important pollinators, and many frugivores play a key role in seed dispersal. Plants and pollinating birds often coevolve, and in some cases a flower's primary pollinator is the only species capable of reaching its nectar. Birds are often important to island ecology. Birds have frequently reached islands that mammals have not; on those islands, birds may fulfil ecological roles typically played by larger animals. For example, in New Zealand nine species of moa were important browsers, as are the kererū and kokako today. Today the plants of New Zealand retain the defensive adaptations evolved to protect them from the extinct moa. Many birds act as ecosystem engineers through the construction of nests, which provide important microhabitats and food for hundreds of species of invertebrates. Nesting
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same envir ...
s may affect the ecology of islands and surrounding seas, principally through the concentration of large quantities of guano, which may enrich the local soil and the surrounding seas. A wide variety of
avian ecology field methods There are many field methods available for conducting avian ecological research. They can be divided into three types: counts, nest monitoring, and capturing and marking. Basic counts Basic bird counts are a good way to estimate population size, ...
, including counts, nest monitoring, and capturing and marking, are used for researching avian ecology.


Relationship with humans

Since birds are highly visible and common animals, humans have had a relationship with them since the dawn of man. Sometimes, these relationships are mutualistic, like the cooperative honey-gathering among
honeyguide Honeyguides (family Indicatoridae) are near passerine birds in the order Piciformes. They are also known as indicator birds, or honey birds, although the latter term is also used more narrowly to refer to species of the genus ''Prodotiscus''. Th ...
s and African peoples such as the Borana. Other times, they may be commensal, as when species such as the
house sparrow The house sparrow (''Passer domesticus'') is a bird of the Old World sparrow, sparrow family Passeridae, found in most parts of the world. It is a small bird that has a typical length of and a mass of . Females and young birds are coloured pale ...
have benefited from human activities. Several bird species have become commercially significant agricultural pests, and some pose an aviation hazard. Human activities can also be detrimental, and have threatened numerous bird species with extinction (
hunting Hunting is the human activity, human practice of seeking, pursuing, capturing, or killing wildlife or feral animals. The most common reasons for humans to hunt are to harvest food (i.e. meat) and useful animal products (fur/hide (skin), hide, ...
,
avian lead poisoning Animal lead poisoning (also known as avian plumbism, or avian saturnism for birds) is a veterinary condition and pathology caused by increased levels of the heavy metal lead in animal's body. Lead interferes with a variety of body and natural proc ...
, pesticides, roadkill,
wind turbine A wind turbine is a device that converts the kinetic energy of wind into electrical energy. Hundreds of thousands of large turbines, in installations known as wind farms, now generate over 650 gigawatts of power, with 60 GW added each year. ...
kills and predation by pet cats and dogs are common causes of death for birds). Birds can act as vectors for spreading diseases such as psittacosis, salmonellosis, campylobacteriosis, mycobacteriosis (avian
tuberculosis Tuberculosis (TB) is an infectious disease usually caused by '' Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can also affect other parts of the body. Most infections show no symptoms, ...
), avian influenza (bird flu), giardiasis, and
cryptosporidiosis Cryptosporidiosis, sometimes informally called crypto, is a parasitic disease caused by '' Cryptosporidium'', a genus of protozoan parasites in the phylum Apicomplexa. It affects the distal small intestine and can affect the respiratory tra ...
over long distances. Some of these are
zoonotic diseases A zoonosis (; plural zoonoses) or zoonotic disease is an infectious disease of humans caused by a pathogen (an infectious agent, such as a bacterium, virus, parasite or prion) that has jumped from a non-human (usually a vertebrate) to a ...
that can also be transmitted to humans.


Economic importance

Domesticated birds raised for meat and eggs, called
poultry Poultry () are domesticated birds kept by humans for their eggs, their meat or their feathers. These birds are most typically members of the superorder Galloanserae (fowl), especially the order Galliformes (which includes chickens, qu ...
, are the largest source of animal protein eaten by humans; in 2003, tons of poultry and tons of eggs were produced worldwide.
Chicken The chicken (''Gallus gallus domesticus'') is a domesticated junglefowl species, with attributes of wild species such as the grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster or cock is a term for an adu ...
s 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 A snipe is any of about 26 wading bird species in three genera in the family Scolopacidae. They are characterized by a very long, slender bill, eyes placed high on the head, and cryptic/ camouflaging plumage. The '' Gallinago'' snipes have ...
, and
woodcock The woodcocks are a group of seven or eight very similar living species of wading birds in the genus ''Scolopax''. The genus name is Latin for a snipe or woodcock, and until around 1800 was used to refer to a variety of waders. The English ...
. Muttonbirding is also popular in Australia and New Zealand. Although some hunting, such as that of muttonbirds, may be sustainable, hunting has led to the extinction or endangerment of dozens of species. 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 faeces ( 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. Birds have been domesticated by humans both as pets and for practical purposes. Colourful birds, such as parrots and
myna The myna (; also spelled mynah) is a bird of the starling family (Sturnidae). This is a group of passerine birds which are native to southern Asia, especially India, Pakistan and Bangladesh. Several species have been introduced to areas lik ...
s, are bred in
captivity Captivity, or being held captive, is a state wherein humans or other animals are confined to a particular space and prevented from leaving or moving freely. An example in humans is imprisonment. Prisoners of war are usually held in captivity by a ...
or kept as pets, a practice that has led to the illegal trafficking of some endangered species.
Falcon Falcons () are birds of prey in the genus ''Falco'', which includes about 40 species. Falcons are widely distributed on all continents of the world except Antarctica, though closely related raptors did occur there in the Eocene. Adult falcons ...
s and cormorants have long been used for
hunting Hunting is the human activity, human practice of seeking, pursuing, capturing, or killing wildlife or feral animals. The most common reasons for humans to hunt are to harvest food (i.e. meat) and useful animal products (fur/hide (skin), hide, ...
and
fishing Fishing is the activity of trying to catch fish. Fish are often caught as wildlife from the natural environment, but may also be caught from fish stocking, stocked bodies of water such as fish pond, ponds, canals, park wetlands and reservoirs. ...
, respectively. Messenger pigeons, used since at least 1 AD, remained important as recently as
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the World War II by country, vast majority of the world's countries—including all of the great power ...
. Today, such activities are more common either as hobbies, for entertainment and tourism, Amateur bird enthusiasts (called birdwatchers, twitchers or, more commonly, birders) number in the millions. 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.


In religion and mythology

Birds play prominent and diverse roles in religion and mythology. In religion, birds may serve as either messengers or priests and leaders for a
deity A deity or god is a supernatural being who is considered divine or sacred. The ''Oxford Dictionary of English'' defines deity as a god or goddess, or anything revered as divine. C. Scott Littleton defines a deity as "a being with powers greate ...
, such as in the Cult of Makemake, in which the Tangata manu of
Easter Island Easter Island ( rap, Rapa Nui; es, Isla de Pascua) is an island and special territory of Chile in the southeastern Pacific Ocean, at the southeasternmost point of the Polynesian Triangle in Oceania. The island is most famous for its nearl ...
served as chiefs or as attendants, as in the case of Hugin and Munin, the two
common raven The common raven (''Corvus corax'') is a large all-black passerine bird. It is the most widely distributed of all corvids, found across the Northern Hemisphere. It is a raven known by many names at the subspecies level; there are at least ...
s who whispered news into the ears of the Norse god Odin. In several civilisations of
ancient Italy The history of Italy covers the ancient period, the Middle Ages, and the modern era. Since classical antiquity, ancient Etruscans, various Italic peoples (such as the Latins, Samnites, and Umbri), Celts, ''Magna Graecia'' colonists, and other a ...
, particularly Etruscan and
Roman Roman or Romans most often refers to: * Rome, the capital city of Italy * Ancient Rome, Roman civilization from 8th century BC to 5th century AD *Roman people, the people of ancient Rome *''Epistle to the Romans'', shortened to ''Romans'', a lett ...
religion Religion is usually defined as a social- cultural system of designated behaviors and practices, morals, beliefs, worldviews, texts, sanctified places, prophecies, ethics, or organizations, that generally relates humanity to supernatur ...
, 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. They may also serve as religious symbols, as when Jonah (, dove) embodied the fright, passivity, mourning, and beauty traditionally associated with doves. Birds have themselves been deified, as in the case of the
common peacock The Indian peafowl (''Pavo cristatus''), also known as the common peafowl, and blue peafowl, is a peafowl species native to the Indian subcontinent. It has been introduced to many other countries. Male peafowl are referred to as peacocks, and ...
, which is perceived as Mother Earth by the people of southern India. In the ancient world, doves were used as symbols of the Mesopotamian goddess Inanna (later known as Ishtar), the Canaanite mother goddess Asherah, and the Greek goddess
Aphrodite Aphrodite ( ; grc-gre, Ἀφροδίτη, Aphrodítē; , , ) is an ancient Greek goddess associated with love, lust, beauty, pleasure, passion, and procreation. She was syncretized with the Roman goddess . Aphrodite's major symbols incl ...
. In
ancient Greece Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cu ...
,
Athena Athena or Athene, often given the epithet Pallas, is an ancient Greek goddess associated with wisdom, warfare, and handicraft who was later syncretized with the Roman goddess Minerva. Athena was regarded as the patron and protectress of v ...
, the goddess of wisdom and patron deity of the city of
Athens Athens ( ; el, Αθήνα, Athína ; grc, Ἀθῆναι, Athênai (pl.) ) is both the capital and largest city of Greece. With a population close to four million, it is also the seventh largest city in the European Union. Athens dominates a ...
, had a little owl as her symbol. 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. Indigenous peoples of the central Andes maintain legends of birds passing to and from metaphysical worlds.


In culture and folklore

Birds have featured in culture and art since prehistoric times, when they were represented in early cave paintings. Some birds have been perceived as monsters, including the mythological Roc and the
Māori Māori or Maori can refer to: Relating to the Māori people * Māori people of New Zealand, or members of that group * Māori language, the language of the Māori people of New Zealand * Māori culture * Cook Islanders, the Māori people of the Co ...
's legendary , a giant bird capable of snatching humans. Birds were later used as symbols of power, as in the magnificent Peacock Throne of the Mughal and Persian emperors. 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. Birds are also important figures in poetry; for example,
Homer Homer (; grc, Ὅμηρος , ''Hómēros'') (born ) was a Greek poet who is credited as the author of the ''Iliad'' and the ''Odyssey'', two epic poems that are foundational works of ancient Greek literature. Homer is considered one of the ...
incorporated nightingales into his ''
Odyssey The ''Odyssey'' (; grc, Ὀδύσσεια, Odýsseia, ) is one of two major ancient Greek epic poems attributed to Homer. It is one of the oldest extant works of literature still widely read by modern audiences. As with the '' Iliad'', ...
'', and
Catullus Gaius Valerius Catullus (; 84 - 54 BCE), often referred to simply as Catullus (, ), was a Latin poet of the late Roman Republic who wrote chiefly in the neoteric style of poetry, focusing on personal life rather than classical heroes. His ...
used a
sparrow Sparrow may refer to: Birds * Old World sparrows, family Passeridae * New World sparrows, family Passerellidae * two species in the Passerine family Estrildidae: ** Java sparrow ** Timor sparrow * Hedge sparrow, also known as the dunnock or hed ...
as an erotic symbol in his Catullus 2. The relationship between an albatross and a sailor is the central theme of
Samuel Taylor Coleridge Samuel Taylor Coleridge (; 21 October 177225 July 1834) was an English poet, literary critic, philosopher, and theologian who, with his friend William Wordsworth, was a founder of the Romantic Movement in England and a member of the Lak ...
's ''
The Rime of the Ancient Mariner ''The Rime of the Ancient Mariner'' (originally ''The Rime of the Ancyent Marinere'') is the longest major poem by the English poet Samuel Taylor Coleridge, written in 1797–1798 and published in 1798 in the first edition of '' Lyrical Ball ...
'', which led to the use of the term as a metaphor for a 'burden'. Other
English English usually refers to: * English language * English people English may also refer to: Peoples, culture, and language * ''English'', an adjective for something of, from, or related to England ** English national ...
metaphors derive from birds;
vulture fund A vulture fund is a hedge fund, private-equity fund or distressed debt fund, that invests in debt considered to be very weak or in default, known as distressed securities. Investors in the fund profit by buying debt at a discounted price on a ...
s and vulture investors, for instance, take their name from the scavenging vulture. Perceptions of bird species vary across cultures. Owls are associated with bad luck,
witchcraft Witchcraft traditionally means the use of magic or supernatural powers to harm others. A practitioner is a witch. In medieval and early modern Europe, where the term originated, accused witches were usually women who were believed to have ...
, and death in parts of Africa, but are regarded as wise across much of Europe. Hoopoes were considered sacred in Ancient Egypt and symbols of virtue in
Persia Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...
, but were thought of as thieves across much of Europe and harbingers of war in
Scandinavia Scandinavia; Sámi languages: /. ( ) is a subregion in Northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. In English usage, ''Scandinavia'' most commonly refers to Denmark, Norway, and S ...
. In
heraldry Heraldry is a discipline relating to the design, display and study of armorial bearings (known as armory), as well as related disciplines, such as vexillology, together with the study of ceremony, rank and pedigree. Armory, the best-known bran ...
, birds, especially
eagles Eagle is the common name for many large birds of prey of the family Accipitridae. Eagles belong to several groups of genera, some of which are closely related. Most of the 68 species of eagle are from Eurasia and Africa. Outside this area, jus ...
, often appear in
coats of arms A coat of arms is a heraldic visual design on an escutcheon (i.e., shield), surcoat, or tabard (the latter two being outer garments). The coat of arms on an escutcheon forms the central element of the full heraldic achievement, which in i ...
.


In music

In music, birdsong has influenced composers and musicians in several ways: they can be inspired by birdsong; they can intentionally imitate bird song in a composition, as
Vivaldi Antonio Lucio Vivaldi (4 March 1678 – 28 July 1741) was an Italian composer, virtuoso violinist and impresario of Baroque music. Regarded as one of the greatest Baroque composers, Vivaldi's influence during his lifetime was widesprea ...
,
Messiaen Olivier Eugène Prosper Charles Messiaen (, ; ; 10 December 1908 – 27 April 1992) was a French composer, organist, and ornithologist who was one of the major composers of the 20th century. His music is rhythmically complex; harmonically ...
, and
Beethoven Ludwig van Beethoven (baptised 17 December 177026 March 1827) was a German composer and pianist. Beethoven remains one of the most admired composers in the history of Western music; his works rank amongst the most performed of the classic ...
did, along with many later composers; they can incorporate recordings of birds into their works, as Ottorino Respighi first did; or like
Beatrice Harrison Beatrice Harrison (9 December 1892 – 10 March 1965) was a British cellist active in the first half of the 20th century. She gave first performances of several important English works, especially those of Frederick Delius, and made the first or ...
and David Rothenberg, they can duet with birds.


Conservation

Although 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 Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the Endling, last individual of the species, although the Functional ext ...
in many other species. Over a hundred bird species have gone extinct in historical times, although the most dramatic human-caused avian extinctions, eradicating an estimated 750–1800 species, occurred during the human colonisation of
Melanesia Melanesia (, ) is a subregion of Oceania in the southwestern Pacific Ocean. It extends from Indonesia's New Guinea in the west to Fiji in the east, and includes the Arafura Sea. The region includes the four independent countries of Fiji, V ...
n,
Polynesia Polynesia () "many" and νῆσος () "island"), to, Polinisia; mi, Porinihia; haw, Polenekia; fj, Polinisia; sm, Polenisia; rar, Porinetia; ty, Pōrīnetia; tvl, Polenisia; tkl, Polenihia (, ) is a subregion of Oceania, made up of ...
n, and
Micronesia Micronesia (, ) is a subregion of Oceania, consisting of about 2,000 small islands in the western Pacific Ocean. It has a close shared cultural history with three other island regions: the Philippines to the west, Polynesia to the east, ...
n islands. Many bird populations are declining worldwide, with 1,227 species listed as
threatened Threatened species are any species (including animals, plants and fungi) which are vulnerable to endangerment in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of '' critical depe ...
by
BirdLife International BirdLife International is a global partnership of non-governmental organizations that strives to conserve birds and their habitats. BirdLife International's priorities include preventing extinction of bird species, identifying and safeguarding ...
and the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
in 2009. The most commonly cited human threat to birds is
habitat loss Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
. Other threats include overhunting, accidental mortality due to collisions with
buildings A building, or edifice, is an enclosed structure with a roof and walls standing more or less permanently in one place, such as a house or factory (although there's also portable buildings). Buildings come in a variety of sizes, shapes, and func ...
or vehicles, long-line fishing
bycatch Bycatch (or by-catch), in the fishing industry, is a fish or other marine species that is caught unintentionally while fishing for specific species or sizes of wildlife. Bycatch is either the wrong species, the wrong sex, or is undersized or juve ...
, pollution (including oil spills and pesticide use), competition and predation from nonnative invasive species, and
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
. Governments and
conservation Conservation is the preservation or efficient use of resources, or the conservation of various quantities under physical laws. Conservation may also refer to: Environment and natural resources * Nature conservation, the protection and manageme ...
groups work to protect birds, either by passing laws that
preserve The word preserve may refer to: Common uses * Fruit preserves, a type of sweet spread or condiment * Nature reserve, an area of importance for wildlife, flora, fauna or other special interest, usually protected Arts, entertainment, and medi ...
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 The Norfolk parakeet (''Cyanoramphus cookii''), also called Tasman parakeet, Norfolk Island green parrot or Norfolk Island red-crowned parakeet, is a species of parrot in the family Psittaculidae. It is endemic to Norfolk Island (located betwe ...
.


See also

*
Animal track __notoc__ An animal track is an imprint left behind in soil, snow, or mud, or on some other ground surface, by an animal walking across it. Animal tracks are used by hunters in tracking their prey and by naturalists to identify animals living i ...
*
Avian sleep In birds, sleep consists of "periods of eye closure interrupted by short periods of eye-opening." During the short periods of eye-opening, electroencephalographic (EEG) studies indicate that the birds are still sleeping; the voltage level in the ...
* Bat *
Climate change and birds Birds are an animal group impacted by human-caused climate change. Changes to bird biology, distribution, and behaviour are among many effects of climate change, and will vary with the temperature reached over preindustrial levels. As many birds ...
* Glossary of bird terms *
List of individual birds This is a list of well-known real birds. For famous fictional birds, see list of fictional birds. * Águia Vitória, a bald eagle who serves as the mascot for Portuguese football club S.L. Benfica * Albert Ross, an albatross believed to have bee ...
*
Ornithology Ornithology is a branch of zoology that concerns the "methodological study and consequent knowledge of birds with all that relates to them." Several aspects of ornithology differ from related disciplines, due partly to the high visibility and t ...
* Paleocene dinosaurs


References


Further reading

* Roger Lederer und Carol Burr: ''Latein für Vogelbeobachter: über 3000 ornithologische Begriffe erklärt und erforscht'', aus dem Englischen übersetzt von Susanne Kuhlmannn-Krieg, Verlag DuMont, Köln 2014, . * del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi (eds.): ''Handbook of the Birds of the World'' (17-volume encyclopaedia), Lynx Edicions, Barcelona, 1992–2010. (''Vol. 1: Ostrich to Ducks'': , etc.). * ''All the Birds of the World'', Lynx Edicions, 2020. * ''National Geographic Field Guide to Birds of North America'', National Geographic, 7th edition, 2017. * ''National Audubon Society Field Guide to North American Birds: Eastern Region'', National Audubon Society, Knopf. * ''National Audubon Society Field Guide to North American Birds: Western Region'', National Audubon Society, Knopf. * Svensson, Lars: ''Birds of Europe'', Princeton University Press, second edition, 2010. * Svensson, Lars: ''Collins Bird Guide: The Most Complete Guide to the Birds of Britain and Europe'', Collins, 2nd edition, 2010.


External links


Birdlife International
– Dedicated to bird conservation worldwide; has a database with about 250,000 records on endangered bird species.



from the National Audubon Society
Cornell Lab of Ornithology
*




Ornithology

Sora
– Searchable online research archive; Archives of the following ornithological journals '' The Auk'', '' Condor'', ''Journal of Field Ornithology, ''North American Bird Bander'', ''Studies in Avian Biology'', ''Pacific Coast Avifauna'', and the '' Wilson Bulletin''.
The Internet Bird Collection
– A free library of videos of the world's birds
The Institute for Bird Populations, California

List of field guides to birds
from the International Field Guides database
RSPB bird identifier
– Interactive identification of all UK birds

— University of California Museum of Paleontology. {{Authority control Animal classes Dinosaurs Extant Late Cretaceous first appearances Feathered dinosaurs Santonian first appearances Taxa named by Carl Linnaeus