Birds are a group of
warm-blooded vertebrates constituting the
class Aves (), characterised by
feather
Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier ...
s, 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 sma ...
of
hard-shelled eggs, a high
metabolic
Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
rate, a four-chambered
heart, and a strong yet lightweight
skeleton
A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
. Birds live worldwide and range in size from the
bee hummingbird to the
ostrich. There are about ten thousand living species, more than half of which are
passerine, 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
forelimb
A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...
s, gave birds the ability to fly, although further evolution has led to the
loss of flight in some birds, including
ratite
A ratite () is any of a diverse group of flightless, large, long-necked, and long-legged birds of the infraclass Palaeognathae. Kiwi, the exception, are much smaller and shorter-legged and are the only nocturnal extant ratites.
The systematics ...
s,
penguin
Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...
s, and diverse
endemic island species. The digestive and respiratory systems of birds are also uniquely adapted for flight. Some bird species of aquatic environments, particularly
seabirds and some
waterbirds, have further evolved for swimming.
Birds are
feathered theropod dinosaurs and constitute the
only known living dinosaurs. Likewise, birds are considered
reptile
Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s in the modern
cladistic sense of the term, and their closest living relatives are the
crocodilians. Birds are descendants of the primitive
avialans (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
Middle or The Middle may refer to:
* Centre (geometry), the point equally distant from the outer limits.
Places
* Middle (sheading), a subdivision of the Isle of Man
* Middle Bay (disambiguation)
* Middle Brook (disambiguation)
* Middle Creek (d ...
to
Late Cretaceous, and diversified dramatically around the time of the
Cretaceous–Paleogene extinction event 66 mya, which killed off the
pterosaurs and all non-avian dinosaurs.
Many
social species
Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies.
Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
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
Monogamy ( ) is a form of Dyad (sociology), dyadic Intimate relationship, relationship in which an individual has only one Significant other, partner during their lifetime. Alternately, only one partner at any one time (Monogamy#Serial monogamy, ...
, usually for one breeding season at a time, sometimes for years, and rarely for life. Other species have breeding systems that are
polygynous
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 ...
(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. 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
Wildlife refers to undomesticated animal species, but has come to include all organisms that grow or live wild in an area without being introduced by humans. Wildlife was also synonymous to game: those birds and mammals that were hunted fo ...
birds being important sources of eggs, meat, and feathers.
Songbirds, parrots, and other species are popular as pets.
Guano
Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
(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 industry.
Evolution and classification
The first
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood.
Classification is the grouping of related facts into classes.
It may also refer to:
Business, organizat ...
of birds was developed by
Francis Willughby and
John Ray
John Ray FRS (29 November 1627 – 17 January 1705) was a Christian English naturalist widely regarded as one of the earliest of the English parson-naturalists. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after ...
in their 1676 volume ''Ornithologiae''.
Carl Linnaeus 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.
Phylogenetic taxonomy
Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...
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 term, ...
Theropoda
Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally ca ...
.
Definition
Aves and a sister group, the order
Crocodilia, contain the only living representatives of the reptile clade
Archosauria. During the late 1990s, Aves was most commonly defined
phylogenetically
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
as all descendants of the
most recent common ancestor of modern birds and ''
Archaeopteryx lithographica''. However, an earlier definition proposed by
Jacques Gauthier
Jacques Armand Gauthier (born June 7, 1948 in New York City) is an American vertebrate paleontologist, comparative morphologist, and systematist, and one of the founders of the use of cladistics in biology.
Life and career
Gauthier is the so ...
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
In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
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 in ...
, 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-avian d ...
s closer to birds than to
crocodile
Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
s (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 phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
", 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 s ...
, 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
Jacques Armand Gauthier (born June 7, 1948 in New York City) is an American vertebrate paleontologist, comparative morphologist, and systematist, and one of the founders of the use of cladistics in biology.
Life and career
Gauthier is the so ...
, 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 dinosaurs and, more specifically, members of
Maniraptora, a group of theropods which includes
dromaeosaurids 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
Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmost ...
Province of northeast China, which demonstrate many small theropod
feathered dinosaurs, contribute to this ambiguity.
The consensus view in contemporary
palaeontology
Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
is that the flying theropods, or
avialans, are the closest relatives of the
deinonychosaur
Deinonychosauria is a clade of paravian dinosaurs which lived from the Late Jurassic to the Late Cretaceous periods. Fossils have been found across the globe in North America, Europe, Africa, Asia, South America, and Antarctica,Case, J.A. ...
s, which include dromaeosaurids and
troodontids.
Together, these form a group called
Paraves. Some
basal
Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''.
Science
* Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure
* Basal (medicine), a minimal level that is nec ...
members of Deinonychosauria, such as ''
Microraptor'', 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
omnivore
An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
s.
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 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 (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 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. 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 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
The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teeth and cla ...
, 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 niches, 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 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-like ''
Ichthyornis'' and the
Hesperornithiformes
Hesperornithes is an extinct and highly specialized group of aquatic avialans closely related to the ancestors of modern birds. They inhabited both marine and freshwater habitats in the Northern Hemisphere, and include genera such as ''Hesperorni ...
, 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
In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
Aves (alternately Neornithes), which has two subdivisions: the Palaeognathae
Palaeognathae (; ) is a infraclass of birds, called paleognaths, within the class Aves of the clade Archosauria. It is one of the two extant infraclasses of birds, the other being Neognathae, both of which form Neornithes. Palaeognathae contai ...
, which includes the flightless ratite
A ratite () is any of a diverse group of flightless, large, long-necked, and long-legged birds of the infraclass Palaeognathae. Kiwi, the exception, are much smaller and shorter-legged and are the only nocturnal extant ratites.
The systematics ...
s (such as the ostriches) and the weak-flying tinamous, and the extremely diverse Neognathae
Neognathae (; ) is a infraclass of birds, called neognaths, within the class Aves of the clade Archosauria. Neognathae includes the majority of living birds; the exceptions being the tinamous and the flightless ratites, which belong instead to ...
, containing all other birds. These two subdivisions have variously been given the rank of superorder
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
, cohort, or infraclass. Depending on the taxonomic
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
viewpoint, the number of known living bird species varies anywhere from 9,800 to 10,758.
The discovery of '' Vegavis'' from the Maastrichtian, 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 configura ...
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
The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma.
Geology
Pro ...
to the latest Late Cretaceous. 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 stages ...
, 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 had been sequenced for only two birds, the chicken 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's breeding colonies up to inland in Antarctica. The highest bird diversity occurs in tropical regions. It was earlier thought that this high diversity was the result of higher speciation
Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
rates in the tropics; however recent studies found higher speciation rates in the high latitudes that were offset by greater extinction 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 species come ashore only to breed, while some penguin
Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...
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, yellow-headed caracara and galah, 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 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
In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate.
Examples
Human anatomy
* Interatri ...
(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 (zoolog ...
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
The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton).
The ...
to form the synsacrum
The synsacrum is a skeletal structure of birds and other dinosaurs, in which the sacrum is extended by incorporation of additional fused or partially fused caudal or lumbar vertebrae and it can only be seen in birds. Some posterior thoracic vert ...
. 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 reptile
Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s, birds are primarily uricotelic, that is, their kidneys extract nitrogenous waste from their bloodstream and excrete it as uric acid, instead of urea or ammonia, through the ureters into the intestine. Birds do not have a urinary bladder or external urethral opening and (with exception of the ostrich) 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
In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
. 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 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
Palaeognathae (; ) is a infraclass of birds, called paleognaths, within the class Aves of the clade Archosauria. It is one of the two extant infraclasses of birds, the other being Neognathae, both of which form Neornithes. Palaeognathae contai ...
(with the exception of the kiwis), the Anseriformes
Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which in ...
(with the exception of screamer
The screamers are three South American bird species placed in Family (biology), family Anhimidae. They were thought to be related to the Galliformes because of similar beak, bills, but are more closely related to ducks (family Anatidae),Todd, F. ...
s), and in rudimentary forms in Galliformes (but fully developed in Cracidae) possess a penis, which is never present in Neoaves
Neoaves is a clade that consists of all modern birds (Neornithes or Aves) with the exception of Paleognathae (ratites and kin) and Galloanserae (ducks, chickens and kin). Almost 95% of the roughly 10,000 known species of extant birds belong to ...
. 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 compartment within the cloaca, just inside the vent. Female birds have sperm storage
Female sperm storage is a biological process and often a type of sexual selection in which sperm cells transferred to a female during mating are temporarily retained within a specific part of the reproductive tract before the oocyte, or egg, is f ...
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
The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
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, 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 is not unknown in birds and eggs can be diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, automictic and results in male offspring.
Birds are solely gonochoric. Meaning they have two sexes: either female or male. 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 or ...
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. 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 systems 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
In classical Greek mythology, Syrinx (Greek Σύριγξ) was a nymph and a follower of Artemis, known for her chastity. Pursued by the amorous god Pan, she ran to a river's edge and asked for assistance from the river nymphs. In answer, sh ...
, 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 (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
Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek language, Greek ''erythros'' for "red" and ''k ...
in birds retain their nucleus.
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 of ...
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 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, 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 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 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
The cerebellum (Latin for "little brain") is a major feature of the hindbrain of all vertebrates. Although usually smaller than the cerebrum, in some animals such as the mormyrid fishes it may be as large as or even larger. In humans, the cerebel ...
coordinates movement and the cerebrum controls behaviour patterns, navigation, mating and nest building. Most birds have a poor sense of smell with notable exceptions including kiwis, New World vultures 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 f ...
. Birds are tetrachromatic
Tetrachromacy (from Greek ''tetra'', meaning "four" and ''chromo'', meaning "color") is the condition of possessing four independent channels for conveying color information, or possessing four types of cone cell in the eye. Organisms with tetr ...
, possessing ultraviolet (UV) sensitive cone cell
Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Cone ...
s 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 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
The nictitating membrane (from Latin '' nictare'', to blink) is a transparent or translucent third eyelid present in some animals that can be drawn across the eye from the medial canthus to protect and moisten it while maintaining vision. All ...
, a third eyelid that moves horizontally. The nictitating membrane also covers the eye and acts as a contact lens
Contact lenses, or simply contacts, are thin lenses placed directly on the surface of the eyes. Contact lenses are ocular prosthetic devices used by over 150 million people worldwide, and they can be worn to correct vision or for cosmetic ...
in many aquatic birds. The bird retina has a fan shaped blood supply system called the pecten
Pecten or pectin may refer to:
Pecten Biology
* Pecten (biology), any comb like structure in animals
* ''Pecten'' (bivalve), a genus of scallops
* Pecten (company), a subsidiary of Sinopec
* Pecten oculi, a structure in the bird retina whic ...
.
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, while birds with eyes on the front of their heads, such as owls, have binocular vision and can estimate the depth of field
The depth of field (DOF) is the distance between the nearest and the furthest objects that are in acceptably sharp focus in an image captured with a camera.
Factors affecting depth of field
For cameras that can only focus on one object dist ...
.[ The avian ear lacks external pinnae but is covered by feathers, although in some birds, such as the '' Asio'', '' Bubo'' and '' Otus'' owls, these feathers form tufts which resemble ears. The ]inner ear
The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...
has a cochlea, 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 can eject an unpleasant stomach oil
Stomach oil is the light oil composed of neutral dietary lipids found in the proventriculus (fore-gut) of birds in the order Procellariiformes. All albatrosses, procellarids (gadfly petrels and shearwaters) and northern and austral storm petrel ...
against an aggressor, and some species of pitohui
The pitohuis are bird species endemic to New Guinea. The onomatopoeic name is thought to be derived from that used by New Guineans from nearby Dorey (Manokwari), but it is also used as the name of a genus '' Pitohui'' which was established by the ...
s from New Guinea have a powerful neurotoxin
Neurotoxins are toxins that are destructive to nerve tissue (causing neurotoxicity). Neurotoxins are an extensive class of exogenous chemical neurological insultsSpencer 2000 that can adversely affect function in both developing and mature ner ...
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 solitaire ('' Pezophaps''), sheathbills ('' Chionis''), some guans (''Crax
''Crax'' is a genus of curassows in the order Galliformes, a clade of large, heavy-bodied, ground-feeding birds. They are known from tropical South America with one species, the great curassow, ranging northwards through Central America as far ...
'') 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, 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
Plumage ( "feather") is a layer of feathers that covers a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between species and subspecies and may vary with age classes. Within species, ...
, may vary within species by age, social status
Social status is the level of social value a person is considered to possess. More specifically, it refers to the relative level of respect, honour, assumed competence, and deference accorded to people, groups, and organizations in a society. Stat ...
, 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—"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 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 feather
Flight feathers (''Pennae volatus'') are the long, stiff, asymmetrically shaped, but symmetrically paired pennaceous feathers on the wings or tail of a bird; those on the wings are called remiges (), singular remex (), while those on the tail ...
s 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 Phasianidae are a family (biology), family of heavy, ground-living birds, which includes pheasants, partridges, junglefowl, chickens, Turkey bird, turkeys, Old World quail, and peafowl. The family includes many of the most popular Game (hunti ...
. 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 passerines 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 patc ...
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, inhibiting the growth of feather-degrading bacteria. This may be supplemented with the secretions of formic acid
Formic acid (), systematically named methanoic acid, is the simplest carboxylic acid, and has the chemical formula HCOOH and structure . It is an important intermediate in chemical synthesis and occurs naturally, most notably in some ants. Es ...
from ants, which birds receive through a behaviour known as anting, to remove feather parasites.
The scales 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
Homology may refer to:
Sciences
Biology
*Homology (biology), any characteristic of biological organisms that is derived from a common ancestor
*Sequence homology, biological homology between DNA, RNA, or protein sequences
* Homologous chrom ...
to those of reptiles and mammals.
Flight
Most birds can fly
Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...
, 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) 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
Flightless birds are birds that through evolution lost the ability to fly. There are over 60 extant species, including the well known ratites ( ostriches, emu, cassowaries, rheas, and kiwi) and penguins. The smallest flightless bird is ...
, 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 or ...
ian land predators. Flightlessness is almost exclusively correlated with gigantism
Gigantism ( el, γίγας, ''gígas'', "giant", plural γίγαντες, ''gígantes''), also known as giantism, is a condition characterized by excessive growth and height significantly above average. In humans, this condition is caused by ove ...
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 auks, shearwaters and dippers.
Behaviour
Most birds are diurnal, but some birds, such as many species of owls and nightjars, are nocturnal
Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite.
Nocturnal creatures generally have highly developed sens ...
or crepuscular
In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
(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
Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
, fruit, plants, seeds, carrion
Carrion () is the decaying flesh of dead animals, including human flesh.
Overview
Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
, 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 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
Pest or The Pest may refer to:
Science and medicine
* Pest (organism), an animal or plant deemed to be detrimental to humans or human concerns
** Weed, a plant considered undesirable
* Infectious disease, an illness resulting from an infection
** ...
insects are considered beneficial 'biological control agents' and their presence encouraged in biological pest control
Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
programmes. Combined, insectivorous birds eat 400–500 million metric tons of arthropods annually.
Nectar feeders such as hummingbird
Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics aro ...
s, 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 shorebirds with long bills probe for invertebrates; shorebirds' varied bill lengths and feeding methods result in the separation of ecological niches. Loon
Loons (North American English) or divers (British / Irish English) are a group of aquatic birds found in much of North America and northern Eurasia. All living species of loons are members of the genus ''Gavia'', family Gaviidae and order Gavi ...
s, diving ducks, penguin
Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...
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
Prions are misfolded proteins that have the ability to transmit their misfolded shape onto normal variants of the same protein. They characterize several fatal and transmissible neurodegenerative diseases in humans and many other animals. It ...
, and some ducks are filter feeder
Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s. Geese and dabbling duck
The Anatinae are a subfamily of the family Anatidae ( swans, geese and ducks). Its surviving members are the dabbling ducks, which feed mainly at the surface rather than by diving. The other members of the Anatinae are the extinct moa-nalo, a yo ...
s are primarily grazers.
Some species, including frigatebirds, gulls, and skuas, engage in kleptoparasitism, 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 frigatebird
The great frigatebird (''Fregata minor'') is a large seabird in the frigatebird family. There are major nesting populations in the tropical Pacific (including the Galapagos Islands) and Indian Oceans, as well as a tiny population in the South At ...
s stealing from masked boobies estimated that the frigatebirds stole at most 40% of their food and on average stole only 5%. Other birds are scavenger
Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
s; 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 sur ...
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
Buttonquail or hemipodes are members of a small family of birds, Turnicidae, which resemble, but are unrelated to, the quails of Phasianidae. They inhabit warm grasslands in Asia, Africa, Europe, and Australia. There are 18 species in two genera ...
and bustard families are capable of sucking up water without the need to tilt back their heads. Some desert birds depend on water sources and sandgrouse 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. 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 seasonal temperatures, therefore optimising availability of food sources and breeding habitat. These migrations vary among the different groups. Many landbirds, shorebirds, and waterbirds undertake annual long-distance migrations, usually triggered by the length of daylight as well as weather conditions. These birds are characterised by a breeding season spent in the temperate or polar region
The polar regions, also called the frigid geographical zone, zones or polar zones, of Earth are the regions of the planet that surround its geographical poles (the North Pole, North and South Poles), lying within the polar circles. These high l ...
s and a non-breeding season in the tropical regions or opposite hemisphere. Before migration, birds substantially increase body fats and reserves and reduce the size of some of their organs.[ (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 . Seabirds also undertake long migrations, the longest annual migration being those of sooty shearwaters, which nest in New Zealand and Chile and spend the northern summer feeding in the North Pacific off Japan, Alaska and California, an annual round trip of . Other seabirds disperse after breeding, travelling widely but having no set migration route. Albatross
Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Pacifi ...
es nesting in the Southern Ocean 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 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 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
The Manx shearwater (''Puffinus puffinus'') is a medium-sized shearwater in the seabird family Procellariidae. The scientific name of this species records a name shift: Manx shearwaters were called Manks puffins in the 17th century. Puffin is an ...
released in Boston 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 constellation
A constellation is an area on the celestial sphere in which a group of visible stars forms Asterism (astronomy), a perceived pattern or outline, typically representing an animal, mythological subject, or inanimate object.
The origins of the e ...
s surrounding Polaris
Polaris is a star in the northern circumpolar constellation of Ursa Minor. It is designated α Ursae Minoris ( Latinized to ''Alpha Ursae Minoris'') and is commonly called the North Star or Pole Star. With an apparent magnitude that ...
. These are backed up in some species by their ability to sense the Earth's geomagnetism through specialised photoreceptors.
Communication
Birds communicate 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 hawk
Hawks are bird of prey, birds of prey of the family Accipitridae. They are widely distributed and are found on all continents except Antarctica.
* The subfamily Accipitrinae includes goshawks, sparrowhawks, sharp-shinned hawks and others. Th ...
s 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
In classical Greek mythology, Syrinx (Greek Σύριγξ) was a nymph and a follower of Artemis, known for her chastity. Pursued by the amorous god Pan, she ran to a river's edge and asked for assistance from the river nymphs. In answer, sh ...
, are the major means by which birds communicate with sound. This communication can be very complex; some species can operate the two sides of the syrinx independently, allowing the simultaneous production of two different songs.
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'' snipes of New Zealand drive air through their feathers, woodpeckers drum for long-distance communication, and palm cockatoo
The palm cockatoo (''Probosciger aterrimus''), also known as the goliath cockatoo or great black cockatoo, is a large smoky-grey or black parrot of the cockatoo family native to New Guinea, Aru Islands, and Cape York Peninsula. It has a very l ...
s 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. The principal benefits of flocking are safety in numbers and increased foraging efficiency. Defence against predators is particularly important in closed habitats like forests, where ambush predation is common and multiple eyes can provide a valuable early warning system. This has led to the development of many mixed-species feeding flocks, which are usually composed of small numbers of many species; these flocks provide safety in numbers but increase potential competition for resources. 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 seabirds associate with dolphins and tuna, which push shoaling fish towards the surface. Some species of hornbill
Hornbills (Bucerotidae) are a family (biology), family of bird found in tropical and subtropical Africa, Asia and Melanesia. They are characterized by a long, down-curved bill which is frequently brightly coloured and sometimes has a Casque (an ...
s have a mutualistic relationship with dwarf mongoose
The common dwarf mongoose (''Helogale parvula'') is a mongoose species native to Angola, northern Namibia, KwaZulu-Natal in South Africa, Zambia and East Africa. It is part of the genus ''Helogale'', along with the Ethiopian dwarf mongoose.
Char ...
s, 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
Slow-wave sleep (SWS), often referred to as deep sleep, consists of stage three of non-rapid eye movement sleep. It usually lasts between 70 and 90 minutes and takes place during the first hours of the night. Initially, SWS consisted of both St ...
one hemisphere of the brain at a time. The birds tend to exercise this ability depending upon its position relative to the outside of the flock. This may allow the eye opposite the sleeping hemisphere to remain vigilant for predators by viewing the outer margins of the flock. This adaptation is also known from marine mammals. Communal roosting 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 hummingbird
Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics aro ...
s go into a nightly state of torpor 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
Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
. 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 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, polyandry, polygamy, polygynandry
Polygynandry is a mating system in which both males and females have multiple mating partners during a breeding season. In sexually reproducing diploid animals, different mating strategies are employed by males and females, because the cost of game ...
, and promiscuity
Promiscuity is the practice of engaging in sexual activity frequently with different Sexual partner, partners or being indiscriminate in the choice of sexual partners. The term can carry a moral judgment. A common example of behavior viewed as pro ...
, 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. 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, 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 simp ...
, 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 seabirds and swifts, often breed in colonies instead; this is thought to offer protection from predators. Colonial breeders defend small nesting sites, and competition between and within species for nesting sites can be intense.
All birds lay amniotic egg
Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are ...
s with hard shells made mostly of calcium carbonate. Hole and burrow nesting species tend to lay white or pale eggs, while open nesters lay camouflaged eggs. There are many exceptions to this pattern, however; the ground-nesting nightjars have pale eggs, and camouflage is instead provided by their plumage. Species that are victims of brood parasites have varying egg colours to improve the chances of spotting a parasite's egg, which forces female parasites to match their eggs to those of their hosts.
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 penguin
The emperor penguin (''Aptenodytes forsteri'') is the tallest and heaviest of all living penguin species and is endemic to Antarctica. The male and female are similar in plumage and size, reaching in length and weighing from . Feathers of th ...
s 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 w ...
) 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 patc ...
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, cuckoos and passerine 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
Blind may refer to:
* The state of blindness, being unable to see
* A window blind, a covering for a window
Blind may also refer to:
Arts, entertainment, and media Films
* ''Blind'' (2007 film), a Dutch drama by Tamar van den Dop
* ''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
The great frigatebird (''Fregata minor'') is a large seabird in the frigatebird family. There are major nesting populations in the tropical Pacific (including the Galapagos Islands) and Indian Oceans, as well as a tiny population in the South At ...
, whose chicks take up to six months to fledge
Fledging is the stage in a flying animal's life between hatching or birth and becoming capable of flight.
This term is most frequently applied to birds, but is also used for bats. For altricial birds, those that spend more time in vulnerable c ...
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 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
The division of labour is the separation of the tasks in any economic system or organisation so that participants may specialise (specialisation). Individuals, organizations, and nations are endowed with, or acquire specialised capabilities, and ...
in which one mate undertakes all or most of a particular duty.
The point at which chicks fledge
Fledging is the stage in a flying animal's life between hatching or birth and becoming capable of flight.
This term is most frequently applied to birds, but is also used for bats. For altricial birds, those that spend more time in vulnerable c ...
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 dimi ...
'' 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'' ...
s, icterid
Icterids () or New World blackbirds make up a family, the Icteridae (), of small to medium-sized, often colorful, New World passerine birds. Most species have black as a predominant plumage color, often enlivened by yellow, orange, or red. The ...
s, and ducks
Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form t ...
, are obligate parasites, though the most famous are the cuckoos. 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 a variety of mating behaviours, with the peacock tail being perhaps the most famous example of sexual selection and the Fisherian runaway. Commonly occurring sexual dimorphisms such as size and colour differences are energetically costly attributes that signal competitive breeding situations. Many types of avian sexual selection 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) 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 (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 (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) 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 insectivore
A robber fly eating a hoverfly
An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects.
The first vertebrate insectivores wer ...
s, frugivore
A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance an ...
s, or nectarivore
In zoology, a nectarivore is an animal which derives its energy and nutrient requirements from a diet consisting mainly or exclusively of the sugar-rich nectar produced by flowering plants.
Nectar as a food source presents a number of benefits a ...
s. Aquatic birds generally feed by fishing, plant eating, and piracy or kleptoparasitism. Many grassland birds are granivores. Birds of prey specialise in hunting mammals or other birds, while vultures are specialised scavenger
Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
s. 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
In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
, 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 engineer
An ecosystem engineer is any species that creates, significantly modifies, maintains or destroys a habitat. These organisms can have a large impact on species richness and landscape-level heterogeneity of an area. As a result, ecosystem enginee ...
s through the construction of nests, which provide important microhabitats and food for hundreds of species of invertebrates. Nesting seabirds may affect the ecology of islands and surrounding seas, principally through the concentration of large quantities of guano
Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
, which may enrich the local soil and the surrounding seas.
A wide variety of avian ecology field methods, including counts, nest monitoring, and capturing and marking, are used for researching avian ecology.
Relationship with humans
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'' ...
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 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 brown and grey, a ...
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 ...
, pesticide
Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and lampri ...
s, roadkill
Roadkill is an animal or animals that have been struck and killed by drivers of motor vehicles on highways. Wildlife-vehicle collisions (WVC) have increasingly been the topic of academic research to understand the causes, and how it can be mi ...
, wind turbine kills and predation by pet cat
The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s and dogs are common causes of death for birds).
Birds can act as vectors for spreading diseases such as psittacosis
Psittacosis—also known as parrot fever, and ornithosis—is a zoonotic infectious disease in humans caused by a bacterium called ''Chlamydia psittaci'' and contracted from infected parrots, such as macaws, cockatiels, and budgerigars, and from ...
, salmonellosis, campylobacteriosis, mycobacteriosis (avian tuberculosis), avian influenza (bird flu), giardiasis, and cryptosporidiosis over long distances. Some of these are zoonotic diseases that can also be transmitted to humans.
Economic importance
Domesticated birds raised for meat and eggs, called poultry, are the largest source of animal protein eaten by humans; in 2003, tons of poultry and tons of eggs were produced worldwide. Chickens account for much of human poultry consumption, though domesticated turkeys, ducks
Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form t ...
, and geese are also relatively common. Many species of birds are also hunted for meat. Bird hunting is primarily a recreational activity except in extremely undeveloped areas. The most important birds hunted in North and South America are waterfowl; other widely hunted birds include pheasants, wild turkeys, quail, doves, partridge, grouse, snipe, and woodcock. Muttonbirding
Muttonbirding is the seasonal harvesting of the chicks of petrels, especially shearwater species, for food, oil and feathers by recreational or commercial hunters. Such hunting of petrels and other seabirds has occurred in various locations since p ...
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
Down most often refers to:
* Down, the relative direction opposed to up
* Down (gridiron football), in American/Canadian football, a period when one play takes place
* Down feather, a soft bird feather used in bedding and clothing
* Downland, a ty ...
of geese and ducks), which are used as insulation in clothing and bedding, and seabird faeces (guano
Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
), 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 mynas, 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
An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inv ...
. Falcons and cormorant
Phalacrocoracidae is a family of approximately 40 species of aquatic birds commonly known as cormorants and shags. Several different classifications of the family have been proposed, but in 2021 the IOC adopted a consensus taxonomy of seven ge ...
s 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, respectively. Messenger pigeons, used since at least 1 AD, remained important as recently as World War II. Today, such activities are more common either as hobbies, for entertainment and tourism,
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 served as chiefs or as attendants, as in the case of Hugin and Munin, the two common ravens who whispered news into the ears of the Norse god Odin
Odin (; from non, Óðinn, ) is a widely revered Æsir, god in Germanic paganism. Norse mythology, the source of most surviving information about him, associates him with wisdom, healing, death, royalty, the gallows, knowledge, war, battle, v ...
. 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 oth ...
, particularly Etruscan and Roman religion, priests were involved in augur
An augur was a priest and official in the classical Roman world. His main role was the practice of augury, the interpretation of the will of the gods by studying the flight of birds. Determinations were based upon whether they were flying i ...
y, 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, 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
Inanna, also sux, 𒀭𒊩𒌆𒀭𒈾, nin-an-na, label=none is an List of Mesopotamian deities, ancient Mesopotamian goddess of love, war, and fertility. She is also associated with beauty, sex, Divine law, divine justice, and political p ...
(later known as Ishtar), the Canaanite mother goddess Asherah, and the Greek goddess Aphrodite. In ancient Greece, Athena, the goddess of wisdom and patron deity of the city of Athens, had a little owl
The little owl (''Athene noctua''), also known as the owl of Athena or owl of Minerva, is a bird that inhabits much of the temperate and warmer parts of Europe, the Palearctic east to Korea, and North Africa. It was introduced into Britain at ...
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 painting
In archaeology, Cave paintings are a type of parietal art (which category also includes petroglyphs, or engravings), found on the wall or ceilings of caves. The term usually implies prehistoric origin, and the oldest known are more than 40,000 ye ...
s. Some birds have been perceived as monsters, including the mythological Roc and the Māori'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
Mughal or Moghul may refer to:
Related to the Mughal Empire
* Mughal Empire of South Asia between the 16th and 19th centuries
* Mughal dynasty
* Mughal emperors
* Mughal people, a social group of Central and South Asia
* Mughal architecture
* Mug ...
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
John James Audubon (born Jean-Jacques Rabin; April 26, 1785 – January 27, 1851) was an American self-trained artist, naturalist, and ornithologist. His combined interests in art and ornithology turned into a plan to make a complete pictoria ...
, whose paintings of North American birds were a great commercial success in Europe and who later lent his name to the National Audubon Society
The National Audubon Society (Audubon; ) is an American non-profit environmental organization dedicated to conservation of birds and their habitats. Located in the United States and incorporated in 1905, Audubon is one of the oldest of such organ ...
. Birds are also important figures in poetry; for example, Homer incorporated nightingales into his '' Odyssey'', and Catullus 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 hedg ...
as an erotic symbol in his Catullus 2
Catullus 2 is a poem by Roman poet Gaius Valerius Catullus (''c''. 84 – ''c''. 54 BCE) that describes the affectionate relationship between an unnamed "puella" (possibly Catullus' lover, Lesbia), and her pet sparrow. As scholar and poet John ...
. The relationship between an albatross
Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Pacifi ...
and a sailor is the central theme of Samuel Taylor Coleridge's '' The Rime of the Ancient Mariner'', which led to the use of the term as a metaphor for a 'burden'. Other English metaphors derive from birds; vulture funds 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, 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, but were thought of as thieves across much of Europe and harbingers of war in Scandinavia. 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 branch ...
, 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, just ...
, 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 its wh ...
.
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, Messiaen, and Beethoven did, along with many later composers; they can incorporate recordings of birds into their works, as Ottorino Respighi
Ottorino Respighi ( , , ; 9 July 187918 April 1936) was an Italian composer, violinist, teacher, and musicologist and one of the leading Italian composers of the early 20th century. List of compositions by Ottorino Respighi, His compositions r ...
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
David Rothenberg (born 1962) is a professor of philosophy and music at the New Jersey Institute of Technology, with a special interest in animal sounds as music. He is also a composer and jazz musician whose books and recordings reflect a lon ...
, 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 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 Melanesian, Polynesian, and Micronesian islands. Many bird populations are declining worldwide, with 1,227 species listed as threatened 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. 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 funct ...
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 spill
An oil spill is the release of a liquid petroleum hydrocarbon into the environment, especially the marine ecosystem, due to human activity, and is a form of pollution. The term is usually given to marine oil spills, where oil is released into th ...
s and pesticide use), competition and predation from nonnative invasive species
An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
, and climate change.
Governments and conservation 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 media
...
and restore Restore may refer to:
* ReStore - Retail building supply stores run by local Habitat for Humanity affiliates
*"Restore", a single by Chris August
Christopher August Megert (born March 20, 1982) is an American Contemporary Christian musician who ...
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.
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
* Bat
* Climate change and birds
Bird, Birds are an animal group impacted by human-caused climate change. Changes to bird biology, Species distribution, distribution, and behaviour are among many effects of climate change, and will vary with the temperature reached over preindustr ...
* Glossary of bird terms
* List of individual birds
* Ornithology
* Paleocene dinosaurs
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutiona ...
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
The National Audubon Society (Audubon; ) is an American non-profit environmental organization dedicated to conservation of birds and their habitats. Located in the United States and incorporated in 1905, Audubon is one of the oldest of such organ ...
Cornell Lab of Ornithology
*
Ornithology
Sora
– Searchable on