The fossil record demonstrates that birds are modern feathered dinosaurs, having evolved from earlier feathered dinosaurs within the theropod group, which are traditionally placed within the saurischian dinosaurs. The closest living relatives of birds are the crocodilians. Primitive bird-like dinosaurs that lie outside class Aves proper, in the broader group Avialae, have been found dating back to the mid-Jurassic period, around 170 million years ago. Many of these early "stem-birds", such as Archaeopteryx, were not yet capable of fully powered flight, and many retained primitive characteristics like toothy jaws in place of beaks, and long bony tails. DNA-based evidence finds that birds diversified dramatically around the time of the Cretaceous–Palaeogene extinction event 66 million years ago, which killed off the pterosaurs and all the non-avian dinosaur lineages. But birds, especially those in the southern continents, survived this event and then migrated to other parts of the world while diversifying during periods of global cooling. This makes them the sole surviving dinosaurs according to cladistics.
Some birds, especially corvids and parrots, are among the most intelligent animals; several bird species make and use tools, and many social species pass on knowledge across generations, which is considered a form of culture. Many species annually migrate great distances. Birds are social, communicating with visual signals, calls, and bird songs, and participating in such social behaviours as cooperative breeding and hunting, flocking, and mobbing of predators. The vast majority of bird species are socially monogamous (referring to social living arrangement, distinct from genetic monogamy), usually for one breeding season at a time, sometimes for years, but rarely for life. Other species have breeding systems that are polygynous (arrangement of one male with many females) or, rarely, polyandrous (arrangement of 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. Some birds, such as hens, lay eggs even when not fertilised, though unfertilised eggs do not produce offspring.
Many species of birds are economically important as food for human consumption and raw material in manufacturing, with domesticated and undomesticated birds (poultry and game) 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 prominently figure throughout human culture. About 120–130 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.
Aves and a sister group, the clade 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. However, an earlier definition proposed by Jacques Gauthier gained wide currency in the 21st century, and is used by many scientists including adherents of the Phylocode system. 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 Avialae, in part to avoid the uncertainties about the placement of Archaeopteryx in relation to animals traditionally thought of as theropod dinosaurs.
Gauthier identified four different definitions for the same biological name "Aves", which is a problem. Gauthier 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 those advanced archosaurs with feathers (alternately Avifilopluma)
Aves can mean those feathered dinosaurs that fly (alternately Avialae)
Aves can mean the last common ancestor of all the currently living birds and all of its descendants (a "crown group", in this sense synonymous with Neornithes)
Under the fourth definition Archaeopteryx is an avialan, and not a member of Aves. Gauthier's 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.
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." 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.
Cladogram following the results of a phylogenetic study by Cau et al., 2015.
Based on fossil and biological evidence, most scientists accept that birds are a specialised subgroup of theropoddinosaurs, and more specifically, they are members of Maniraptora, a group of theropods which includes dromaeosaurs and oviraptorids, among others. As scientists have discovered more theropods closely related to birds, the previously clear distinction between non-birds and birds has become blurred. Recent discoveries in the Liaoning Province of northeast China, which demonstrate many small theropod feathered dinosaurs, contribute to this ambiguity.
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. Together, these form a group called Paraves. Some basal members of this group, 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 omnivores.
The Late JurassicArchaeopteryx 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.
The well-known 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.
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). While the earliest forms, such as Archaeopteryx and Jeholornis, retained the long bony tails of their ancestors, the tails of more advanced avialans were shortened with the advent of the pygostyle bone in the group Pygostylia. 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.
Early diversity of bird ancestors
Ichthyornis, which lived 93 million years ago, was the first known prehistoric bird relative preserved with teeth.
Mesozoic bird phylogeny simplified after Wang et al., 2015's phylogenetic analysis.
The first large, diverse lineage of short-tailed avialans to evolve were the enantiornithes, or "opposite birds", so named because the construction of their shoulder bones was in reverse to that of modern birds. Enantiornithes occupied a wide array of ecological 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, the Hesperornithiformes, which became so well adapted to hunting fish in marine environments that they lost the ability to fly and became primarily aquatic. The early euornithes also saw the development of many traits associated with modern birds, like strongly keeled breastbones, toothless, beaked portions of their jaws (though most non-avian euornithes retained teeth in other parts of the jaws). Euornithes also included the first avialans to develop true pygostyle and a fully mobile fan of tail feathers, which may have replaced the "hind wing" as the primary mode of aerial maneuverability and braking in flight.
The discovery of Vegavis, a late Cretaceous member of the Anatidae, proved that the diversification of modern birds started before the Cenozoic. 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.
Most studies agree on a Cretaceous age for the most recent common ancestor of modern birds but estimates range from the Middle Cretaceous 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 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 modern birds originated early in the Late Cretaceous, a pulse of diversification in all major groups occurred around the Cretaceous–Palaeogene extinction event.
The classification of birds is a contentious issue. Sibley and Ahlquist's Phylogeny and Classification of Birds (1990) is a landmark work on the classification of birds, although it is frequently debated and constantly revised. Most evidence seems to suggest the assignment of orders is accurate, but scientists disagree about the relationships between 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.