Birds are a group of warm-blooded vertebrates constituting the class Aves (/ˈeɪviːz/),

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 skeleton. Birds live
worldwide and range in size from the 5.5 cm (2.2 in) bee hummingbird to the 2.8 m (9 ft 2 in)
common ostrich. There are over 11,000 living species, more than half of which are passerine, or
"perching" birds. Birds have wings whose development varies according to species; the only
known groups without wings are the extinct moa and elephant birds. Wings, which are modified
forelimbs, gave birds the ability to fly, although further evolution has led to the loss of flight in
some birds, including ratites, penguins, 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.
The study of birds is called ornithology.

Birds are feathered theropod dinosaurs and constitute the only known living dinosaurs. Likewise,
birds are considered reptiles in the modern cladistic sense of the term, and their closest living
relatives are the crocodilians. Birds are descendants of the primitive avialans (whose members
include Archaeopteryx) which first appeared during the Late Jurassic. According to recent
estimates, modern birds (Neornithes) evolved in the Late Cretaceous and diversified
dramatically around the time of the Cretaceous–Paleogene extinction event 66 million years ago,
which killed off the pterosaurs and all non-avian dinosaurs.[7]

Many social species pass on knowledge across generations, which is considered a form of
culture. Birds are social, communicating with visual signals, calls, and songs, and participating in
such behaviours as cooperative breeding and hunting, flocking, and mobbing of predators. The
vast majority of bird species are socially (but not necessarily sexually) monogamous, usually for
one breeding season at a time, sometimes for years, and rarely for life. Other species have
breeding systems that are polygynous (one male with many females) or, rarely, polyandrous (one
female with many males). Birds produce offspring by laying eggs which are fertilised through
sexual reproduction. They are usually laid in a nest and incubated by the parents. Most birds
have an extended period of parental care after hatching.

Many species of birds are economically important as food for human consumption and raw
material in manufacturing, with domesticated and undomesticated birds being important sources
of eggs, meat, and feathers. Songbirds, parrots, and other species are popular as pets. Guano
(bird excrement) is harvested for use as a fertiliser. Birds figure throughout human culture.
About 120 to 130 species have become extinct 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
Main article: Evolution of birds

Archaeopteryx is often considered the oldest known true bird.

The first classification of birds was developed by Francis Willughby and John Ray in their 1676
volume Ornithologiae.[8] Carl Linnaeus modified that work in 1758 to devise the taxonomic
classification system currently in use.[9] Birds are categorised as the biological class Aves in
Linnaean taxonomy. Phylogenetic taxonomy places Aves in the clade Theropoda.[10]

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 as all descendants of the most recent common ancestor of modern birds and
Archaeopteryx lithographica.[11] However, an earlier definition proposed by Jacques Gauthier
gained wide currency in the 21st century, and is used by many scientists including adherents to
the PhyloCode. Gauthier defined Aves to include only the crown group of the set of modern
birds. This was done by excluding most groups known only from fossils, and assigning them,
instead, to the broader group Avialae,[12] on the principle that a clade based on extant species
should be limited to those extant species and their closest extinct relatives.[12]

Gauthier and de Queiroz identified four different definitions for the same biological name
"Aves", which is a problem.[13] 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,[13]
which corresponds to meaning number 4 below. They assigned other names to the other groups.
[13]

Reptiles
 Squamat
es Lizards &
snakes

P
a
n
t
e
st
u Turtles
d
i
n
e
s

A
r Cro
c codi
h les
o
s
a
u Bir
r ds
s

The birds' phylogenetic relationships to major

living reptile groups. (The turtles' position is
uncertain: Some authorities embed them inside the
Archosaurs, with birds and crocodiles.)
1. Aves can mean all archosaurs closer to birds than to crocodiles (alternately
Avemetatarsalia)
2. Aves can mean those advanced archosaurs with feathers (alternately Avifilopluma)
3. Aves can mean those feathered dinosaurs that fly (alternately Avialae)
4. Aves can mean the last common ancestor of all the currently living birds and all of its
descendants (a "crown group", in this sense synonymous with Neornithes)
Under the fourth definition Archaeopteryx, traditionally considered one of the earliest members
of Aves, is removed from this group, becoming a non-avian dinosaur instead. These proposals
have been adopted by many researchers in the field of palaeontology and bird evolution, 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.[14]

Maniraptoromorpha
†Coelurus
 †Ornitholestes

M
a
n †Ornithomimosauria
ir
a
p
t
o Manirapto
ri ra
f
o †Alvarezsauridae
r
Cladogram showing the results of a phylogenetic study by Cau, 2018.[15]

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",[16][17] with
Troodon being sometimes added as a second external specifier in case it is closer to birds than to
Deinonychus.[18] Avialae is also occasionally defined as an apomorphy-based clade (that is, one
based on physical characteristics). Jacques Gauthier, who named Avialae in 1986, re-defined it in
2001 as all dinosaurs that possessed feathered wings used in flapping flight, and the birds that
descended from them.[13][19]

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.[20]

Dinosaurs and the origin of birds

Main article: Origin of birds
Parave
s
†Scansoriopterygidae
 †Eosinopteryx

E
u
m
a †Jinfengo
n pteryx
ir
a
p
t
o
r
a †Aurornis

Cladogram following the results of a

phylogenetic study by Cau et al., 2015[21]

Simplified phylogenetic tree showing the relationship

between modern birds and other dinosaurs[22]

Based on fossil and biological evidence, most scientists accept that birds are a specialised
subgroup of theropod dinosaurs[23] and, more specifically, members of Maniraptora, a group of
theropods which includes dromaeosaurids and oviraptorosaurs, among others.[24] As scientists
have discovered more theropods closely related to birds, the previously clear distinction between
non-birds and birds has become blurred. By the 2000s, discoveries in the Liaoning Province of
northeast China, which demonstrated many small theropod feathered dinosaurs, contributed to
this ambiguity.[25][26][27]
 Anchiornis huxleyi is an important source of information on the
early evolution of birds in the Late Jurassic period.[28]

The consensus view in contemporary palaeontology is that the flying theropods, or avialans, are
the closest relatives of the deinonychosaurs, which include dromaeosaurids and troodontids.[29]
Together, these form a group called Paraves. Some basal 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.[30][31] Unlike Archaeopteryx
and the non-avialan feathered dinosaurs, who primarily ate meat, studies suggest that the first
avialans were omnivores.[32]

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.[33]

Early evolution
See also: List of fossil bird genera

Confuciusornis sanctus, a Cretaceous bird from China that lived

125 million years ago, is the oldest known bird to have a beak.[34]

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.[15][35] 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. [15] The
integument evolved into complex, pennaceous feathers.[35]

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, and Aurornis xui.[14]

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 maneuvering.[36]

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).[37] 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.[15]
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.[38] Around 95 million
years ago, they evolved a better sense of smell.[39]

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. [15]

Avia