Corvid

Corvidae is a cosmopolitan family of oscine passerine birds that contains the crows, ravens, rooks, jackdaws, jays, magpies, treepies, choughs and nutcrackers. The common English names used are corvids (more technically) or the crow family (more informally), and there are over 120 species. The genus Corvus, including the jackdaws, crows and ravens, makes up over a third of the entire family.

Systematics, taxonomy and evolution


Over the years there has been much disagreement on the exact evolutionary relationships of the corvid family and their relatives. What eventually seemed clear was that corvids are derived from Australasian ancestors and from there spread throughout the world. Other lineages derived from these ancestors evolved into ecologically diverse, but often Australasian groups. Sibley and Ahlquist united the corvids with other taxa in the Corvida. The presumed corvid relatives included currawongs, birds of paradise, whipbirds, quail-thrushes, whistlers, monarch flycatchers and drongos, shrikes, vireos and vangas, but current research favors the theory that this grouping is partly artificial. The corvids constitute the core group of the Corvoidea, together with their closest relatives (the birds of paradise, Australian mud-nesters and shrikes). They are also the core group of the Corvida, which includes the related groups, such as orioles and vireos.

Clarification of the interrelationships of the corvids has been achieved based on cladistic analysis of several DNA sequences. The jays and magpies do not constitute monophyletic lineages, but rather seem to split up into an American and Old World lineage, and an Holarctic and Oriental lineage, respectively. These are not closely related among each other. The position of the Azure-winged Magpie, which has always been a major enigma, is even less clear than before.

Genus Pyrrhocorax

 * Alpine Chough, Pyrrhocorax graculus
 * Red-billed Chough, Pyrrhocorax pyrrhocorax

Genus Temnurus

 * Ratchet-tailed Treepie, Temnurus temnurus

Genus Crypsirina

 * Hooded Treepie, Crypsirina cucullata
 * Black Racket-tailed Treepie, Crypsirina temia

Genus Platysmurus

 * Black Magpie, Platysmurus leucopterus

Genus Dendrocitta

 * Rufous Treepie, Dendrocitta vagabunda
 * Sumatran Treepie, Dendrocitta occipitalis
 * Bornean Treepie, Dendrocitta cinerascens
 * Grey Treepie, Dendrocitta formosae
 * White-bellied Treepie, Dendrocitta leucogastra
 * Collared Treepie, Dendrocitta frontalis
 * Andaman Treepie, Dendrocitta bayleyi

Genus Cissa

 * Common Green-Magpie, Cissa chinensis
 * Indochinese Green-Magpie, Cissa hypoleuca
 * Bornean Green-Magpie, Cissa jefferyi
 * Javan Green-Magpie, Cissa thalassina

Genus Urocissa

 * Sri Lanka Blue-Magpie, Urocissa ornata
 * Taiwan Blue-Magpie, Urocissa caerulea
 * Red-billed Blue-Magpie, Urocissa erythrorhyncha
 * Yellow-billed Blue-Magpie, Urocissa flavirostris
 * White-winged Magpie, Urocissa whiteheadi

Genus Cyanopica

 * Azure-winged Magpie, Cyanopica cyana
 * Iberian Magpie, Cyanopica cooki

Genus Perisoreus

 * Siberian Jay, Perisoreus infaustus
 * Sichuan Jay, Perisoreus internigrans
 * Grey Jay, Perisoreus canadensis

Genus Cyanolyca

 * White-throated Jay, Cyanolyca mirabilis
 * Dwarf Jay, Cyanolyca nana
 * Black-throated Jay, Cyanolyca pumilo
 * Silvery-throated Jay, Cyanolyca argentigula
 * Azure-hooded Jay, Cyanolyca cucullata
 * Beautiful Jay, Cyanolyca pulchra
 * Black-collared Jay, Cyanolyca armillata
 * Turquoise Jay, Cyanolyca turcosa
 * White-collared Jay, Cyanolyca viridicyana

Genus Gymnorhinus

 * Pinyon Jay, Gymnorhinus cyanocephalus

Genus Cyanocitta

 * Steller's Jay, Cyanocitta stelleri
 * Blue Jay, Cyanocitta cristata

Genus Aphelocoma

 * Unicoloured Jay, Aphelocoma unicolor
 * Transvolcanic Jay, Aphelocoma ultramarina
 * Mexican Jay, Aphelocoma wollweberi
 * Florida Scrub-Jay, Aphelocoma coerulescens
 * Woodhouse's Scrub-Jay, Aphelocoma woodhouseii
 * California Scrub-Jay, Aphelocoma californica
 * Island Scrub-Jay, Aphelocoma insularis


 * Genus Calocitta - magpie-jays
 * Black-throated Magpie-jay, Calocitta colliei
 * White-throated Magpie-jay, Calocitta formosa
 * Genus Cyanocorax
 * Black-chested Jay, Cyanocorax affinis
 * Purplish-backed Jay, Cyanocorax beecheii
 * Azure Jay, Cyanocorax caeruleus
 * Cayenne Jay, Cyanocorax cayanus
 * Plush-crested Jay, Cyanocorax chrysops
 * Curl-crested Jay, Cyanocorax cristatellus
 * Purplish Jay, Cyanocorax cyanomelas
 * White-naped Jay, Cyanocorax cyanopogon
 * Tufted Jay, Cyanocorax dickeyi
 * Azure-naped Jay, Cyanocorax heilprini
 * Bushy-crested Jay, Cyanocorax melanocyaneus
 * Brown Jay, Cyanocorax morio
 * White-tailed Jay, Cyanocorax mystacalis
 * San Blas Jay, Cyanocorax sanblasianus
 * Violaceous Jay, Cyanocorax violaceus
 * Green Jay, Cyanocorax ynca
 * Yucatan Jay, Cyanocorax yucatanicus

Genus Garrulus

 * Eurasian Jay, Garrulus glandarius
 * Lanceolated Jay, Garrulus lanceolatus
 * Lidth's Jay, Garrulus lidthi


 * Genus Podoces - ground jays
 * Biddulph's Ground Jay, Podoces biddulphi
 * Henderson's Ground Jay, Podoces hendersoni
 * Grey Ground Jay, Podoces panderi
 * Persian Ground Jay, Podoces pleskei


 * Genus Ptilostomus
 * Piapiac, Ptilostomus afer


 * Stresemann's Bush Crow
 * Genus Zavattariornis
 * Stresemann's Bush Crow, Zavattariornis stresemanni


 * Nutcrackers
 * Genus Nucifraga
 * Spotted Nutcracker, Nucifraga caryocatactes
 * Clark's Nutcracker, Nucifraga columbiana


 * Holarctic magpies
 * Genus Pica
 * Black-billed Magpie, Pica hudsonia
 * Yellow-billed Magpie, Pica nuttalli
 * European Magpie, Pica pica
 * Korean Magpie, Pica (pica) sericea


 * True crows (crows, ravens, jackdaws and rooks)
 * Genus Corvus
 * Australian and Melanesian species
 * Little Crow, Corvus bennetti
 * Australian Raven, Corvus coronoides
 * Bismarck Crow, Corvus insularis
 * Brown-headed Crow, Corvus fuscicapillus
 * Bougainville Crow, Corvus meeki
 * Little Raven, Corvus mellori
 * New Caledonian Crow, Corvus moneduloides
 * Torresian Crow, Corvus orru
 * Forest Raven, Corvus tasmanicus
 * Relict Raven, Corvus (tasmanicus) boreus
 * Grey Crow, Corvus tristis
 * Long-billed Crow, Corvus validus
 * White-billed Crow, Corvus woodfordi
 * Pacific island species
 * Alalā (Hawaiian Crow), Corvus hawaiiensis (formerly Corvus tropicus) (extinct in the wild)
 * Mariana Crow, Corvus kubaryi
 * Tropical Asian species
 * Daurian Jackdaw, Corvus dauuricus
 * Slender-billed Crow, Corvus enca
 * Flores Crow, Corvus florensis
 * Large-billed Crow, Corvus macrorhynchos
 * Jungle Crow, Corvus (macrorhynchos) levaillantii
 * House Crow, Corvus splendens
 * Collared Crow, Corvus torquatus
 * Piping Crow, Corvus typicus
 * Banggai Crow, Corvus unicolor
 * Eurasian and North African species
 * Hooded Crow, Corvus cornix
 * Mesopotamian Crow, Corvus (cornix) capellanus
 * Carrion Crow (Western Carrion Crow), Corvus corone
 * Eastern Carrion Crow, Corvus (corone) orientalis
 * Rook, Corvus frugilegus
 * Jackdaw, Corvus monedula
 * Fan-tailed Raven, Corvus rhipidurus
 * Brown-necked Raven, Corvus ruficollis
 * Holarctic species
 * Common Raven, Corvus corax (see also next section)
 * Pied Raven, Corvus corax varius morpha leucophaeus (an extinct color variant)
 * North and Central American species
 * American Crow, Corvus brachyrhynchos
 * Northwestern Crow, Corvus caurinus
 * Chihuahuan Raven, Corvus cryptoleucus
 * Tamaulipas Crow, Corvus imparatus
 * Jamaican Crow, Corvus jamaicensis
 * White-necked Crow, Corvus leucognaphalus
 * Cuban Crow, Corvus nasicus
 * Fish Crow, Corvus ossifragus
 * Palm Crow, Corvus palmarum
 * Sinaloan Crow, Corvus sinaloae
 * Western Raven, Corvus (corax) sinuatus
 * Tropical African species
 * White-necked Raven, Corvus albicollis
 * Pied Crow, Corvus albus
 * Cape Crow, Corvus capensis
 * Thick-billed Raven, Corvus crassirostris
 * Somali Crow (Dwarf Raven), Corvus edithae


 * New World jays

The Crested Jay (Platylophus galericulatus) is traditionally included in the Corvidae, but might not be a true member of this family, possibly being closer to the helmet-shrikes (Malaconotidae) or shrikes (Laniidae); it is best considered Corvidae incertae sedis for the time being. Likewise, the Hume's Ground "Jay" (Pseudopodoces humilis) is in fact a member of the tit family Paridae.

Fossil record
The earliest corvid fossils date to the mid-Miocene, about 17 million years ago; Miocorvus and Miopica may be ancestral to crows and some of the magpie lineage, respectively, or similar to the living forms due to convergent evolution. The known prehistoric corvid genera appear to be mainly of the New World and Old World jay and Holarctic magpie lineages:


 * Miocorvus (Middle Miocene of Sansan, France)
 * Miopica (Middle Miocene of SW Ukraine)
 * Miocitta (Pawnee Creek Late Miocene of Logan County, USA)
 * Corvidae gen. et sp. indet. (Edson Early Pliocene of Sherman County, USA)
 * Protocitta (Early Pleistocene of Reddick, USA)
 * Corvidae gen. et sp. indet. (Early/Middle Pleistocene of Sicily) - probably belongs in an extant genus
 * Henocitta (Arredondo Clay Middle Pleistocene of Williston, USA)

In addition, there are numerous fossil species of extant genera since the Mio–Pliocene, mainly European Corvus.

Morphology
Corvids are medium to large in size, with strong feet and bills, rictal bristles and a single moult each year (most passerines moult twice). Corvids are found worldwide except for the tip of South America and the polar ice caps. The majority of the species are found in tropical South and Central America, southern Asia and Eurasia, with fewer than 10 species each in Africa, Australasia and North America. The genus Corvus has re-entered Australia in relatively recent geological prehistory, with five species and one subspecies there.

Corvids are large to very large passerines with a robust build, strong legs and all species except the Pinyon Jay have nostrils covered by bristle-like feathers. Many corvids of temperate zones have mainly black or blue coloured plumage; however, some are pied black and white, some have a blue-purple iridescence and many tropical species are brightly coloured. The sexes are very similar in color and size. Corvids have strong, stout bills and large wingspans. The family includes the largest members of the passerine order.

The smallest corvid is the Dwarf Jay (Aphelocoma nana), at 40 g (1.4 oz) and 21.5 cm (8.5 inches). The largest corvids are the Common Raven (Corvus corax) and the Thick-billed Raven (Corvus crassirostris), both of which regularly exceed 1400 grams (3 lbs) and 65 cm (26 inches).

Species can be identified based on size, shape, and geography; however, some, especially the Australian crows, are best identified by their raucous calls.

Ecology
Corvids occur in most climatic zones. Most are sedentary and do not migrate significantly. However, during a shortage of food, eruptive migration can occur. When species are migratory, they will form large flocks in the fall (around August in the northern hemisphere) and travel south.

One reason for the success of crows, compared to ravens, is their ability to overlap breeding territory. During breeding season, crows were shown to overlap breeding territory six times as much as ravens. This invasion of breeding ranges allowed a related increase in local population density.

Food and feeding
The natural diet of many corvid species is omnivorous, consisting of invertebrates, nestlings, small mammals, berries, fruits, seeds, and carrion. However, some corvids, especially the crows, have adapted well to human conditions and have come to rely on anthropogenic foods. In a US study of American Crows, Common Ravens and Steller's Jays around campgrounds and human settlements, the crows appeared to have the most diverse diet of all, taking anthropogenic foods such as bread, spaghetti, fried potatoes, dog food, sandwiches, and livestock feed. The increase in available anthropogenic food sources is contributing to population increase in some corvid species.

Some corvids are predators of other birds. During the wintering months, corvids typically form foraging flocks. However, some crows also eat many agricultural pests including cutworms, wireworms, grasshoppers, and harmful weeds Some corvids will eat carrion, and since they lack a specialized beak for tearing into flesh, they must wait until animals are opened, whether by other predators or as roadkill.

Since crows do not seem to mind human development, it was suggested that the crow population increase would cause increased rates of nest predation. However, Steller's Jays, which are successful independently of human development, are more efficient in plundering small birds' nests than American Crows and Common Ravens. Therefore, the human relationship with crows and ravens did not significantly increase nest predation, compared to other factors such as habitat destruction.

Reproduction
Many species of corvid are territorial, protecting territories throughout the year or simply during the breeding season. In some cases territories may only be guarded during the day, with the pair joining off-territory roosts at night. Some corvids are well known communal roosters. Some groups of roosting corvids can be very large, with a roost of 65,000 Rooks counted in Scotland. Some, including the Rook and the Jackdaw, are also communal nesters.

The partner bond in corvids is extremely strong and even lifelong in some species. This monogamous lifestyle, however, can still contain extra-pair copulations. Males and females build large nests together in trees or on ledges. The male will also feed the female during incubation. The nests are constructed of a mass of bulky twigs lined with grass and bark. Corvids can lay between 3 and 10 eggs, typically ranging between 4 and 7. The eggs are usually greenish in colour with brown blotches. Once hatched, the young remain in the nests for up to 6–10 weeks depending on the species. Corvids provide biparental care.

Jackdaws can breed in buildings or in rabbit warrens. White-throated Magpie-jays are cooperatively breeding corvids where the helpers are mostly female. Cooperative breeding takes place when additional adults help raise the nestlings. Such helpers at the nest in most cooperatively breeding birds are males, while females join other groups.

Social life
Some corvids have strong organization and community groups. Jackdaws, for example, have a strong social hierarchy, and are facultatively colonial during breeding. Providing mutual aid has also been recorded within many of the corvid species.

Young corvids have been known to play and take part in elaborate social games. Documented group games follow a "king of the mountain"- and "follow the leader"-type pattern. Other play involves the manipulation, passing, and balancing of sticks. Corvids also take part in other activities, such as sliding down smooth surfaces. These games are understood to play a large role in the adaptive and survival ability of the birds.

Mate selection is quite complex and accompanied with much social play in the Corvidae. Youngsters of social corvid species undergo a series of tests, including aerobatic feats, before being accepted as a mate by the opposite sex.

Some corvids can be aggressive. Blue Jays, for example, are well known to attack anything that threatens their nest. Crows have been known to attack dogs, cats, ravens, and birds of prey. Most of the time these assaults take place as a distraction long enough to allow an opportunity for stealing food.

Intelligence
They are considered the most intelligent of the birds, and among the most intelligent of all animals  having demonstrated self-awareness in mirror tests (European Magpies) and tool making ability (Crows, Rooks )—skills until recently regarded as solely the province of humans and a few other higher mammals. Their total brain to body ratio is equal to that of great apes and cetaceans, and only slightly lower than in humans.

The brain-to-body weight ratios of corvid brains are among the largest in birds, equal to that of great apes and cetaceans, and only slightly lower than a human. Their intelligence is boosted by the long growing period of the young. By remaining with the parents, the young have more opportunities to learn necessary skills. Since most corvids are cooperative brooders, their young can learn from different members of the group.

When compared to dogs and cats in an experiment testing the ability to seek out food according to three-dimensional clues, corvids out-performed the mammals. A metaanalysis testing how often birds invented new ways to acquire food in the wild found corvids the most innovative birds. A 2004 review suggests that their cognitive abilities are on par with those of great apes. Despite structural differences, the brains of corvids and great apes both evolved the ability to make geometrical measurements.

Corvid ingenuity is represented through their feeding skills, memorization abilities, use of tools, and group behaviour. Living in large social groups has long been connected with high cognitive ability. To live in a large group, a member must be able to recognize individuals and track the social position and foraging of other members over time. Members must also be able to distinguish between sex, age, reproductive status, and dominance, and to update this information constantly. It might be that social complexity corresponds to their high cognition.

The European Magpie is one of the few non-mammal species known to be able to recognize itself in a mirror test.

There are also specific examples of corvid cleverness. One Carrion Crow was documented to crack nuts by placing them on a crosswalk, letting the passing cars crack the shell, waiting for the light to turn red, and then safely retrieving the contents. A group of crows in England took turns lifting garbage bin lids while their companions collected food.

Members of the corvid family have been known to watch other birds, remember where they hide their food, then return once the owner leaves. Corvids also move their food around between hiding places to avoid thievery, but only if they have previously been thieves themselves i.e., they remember previous relevant social contexts, use their own experience of having been a thief to predict the behavior of a pilferer, and can determine the safest course to protect their caches from being pilfered. Studies to assess similar cognitive abilities in apes have been inconclusive.

The ability to hide food requires highly accurate spatial memories. Corvids have been recorded to recall their food's hiding place up to nine months later. It is suggested that vertical landmarks (like trees) are used to remember locations. There has also been evidence that Western Scrub-Jays, which store perishable foods, not only remember where they stored their food, but for how long. This has been compared to episodic memory, previously thought unique to humans.

New Caledonian Crows (Corvus moneduloides) are notable for their highly developed tool fabrication. They make angling tools of twigs and leaves trimmed into hooks, then use the hooks to pull insect larvae from tree holes. Tools are engineered according to task and apparently also to learned preference. Recent studies revealed abilities to solve complicated problems, which suggests high level of innovation of a complex nature. Other corvids that have been observed using tools include the American Crow, Blue Jay and Green Jay. Diversity in tool design among corvids suggests cultural variation. Again, great apes are the only other animals known to use tools in such a fashion.

Clark's Nutcrackers and Jackdaws were compared in a 2002 study based on geometric rule learning. The corvids, along with a domestic pigeon, had to locate a target between two landmarks, while distances and landmarks were altered. The nutcrackers were more accurate in their searches than the jackdaws and pigeons.

The scarecrow is an archetypal scare tactic in the agricultural business. However, due to corvids' quick wit, scarecrows are soon ignored and used as perches. Despite farmers' efforts to rid themselves of corvid pests, their attempts have only expanded corvid territories and strengthened their numbers.

Current systematics places corvids, based on physical characteristics other than their brains (the most developed of birds), in the lower middle of the passerines, contrary to earlier teleological classifications as "highest" songbirds due to their intelligence. As per one observer:

"During the 19th century there arose the belief that these were the 'most advanced' birds, based upon the belief that Darwinian evolution brings 'progress'. In such a classification the 'most intelligent' of birds were listed last reflecting their position 'atop the pyramid'. Modern biologists reject the concept of hierarchical 'progress' in evolution [...]."

The other major group of highly intelligent birds of the order Psittaciformes (which includes 'true' parrots, cockatoos and New Zealand parrots) is not closely related to corvids.

Disease
Corvids are reservoirs (carriers) for the West Nile Virus in the USA. They are infected by mosquitoes (the vectors), primarily of the Culex species. Crows and Ravens are quickly killed by this disease, so their deaths are an early-warning system when West Nile Virus arrives in an area (as are horse and other bird species deaths). One of the first signs that West Nile Virus first arrived in the USA in 1999 was the death of crows in New York.

Relationship with humans
Several different corvids, particularly ravens, have occasionally served as pets, although they are not able to speak as readily as parrots and do not like being caged.

Role in myth and culture
Folklore often represents corvids as clever, and even mystical, animals. Some Native Americans, such as the Haida, believed that a raven created the earth and despite being a trickster spirit, ravens were popular on totems, credited with creating man, and responsible for placing the Sun in the sky.

Various Germanic peoples highly revered the raven. The major deity Odin was so associated with ravens throughout history that he gained the kenning "raven god" and the raven banner was the flag of various Viking Age Scandinavian chieftains. He was also attended by Hugin and Munin, two ravens who whispered news into his ears. The Valravn sometimes appears in modern Scandinavian folklore.

The 6th century BC Greek scribe Aesop featured corvids as intelligent antagonists in many fables. Later, in western literature, popularized by American poet Edgar Allan Poe's work "The Raven", the Common Raven becomes a symbol of the main character's descent into madness.

Status and conservation
Unlike many other bird families, corvid fitness and reproduction, especially with many crows, has increased due to human development. The survival and reproductive success of certain crows and ravens is assisted by their close relationship with humans.

Human development provides additional resources by clearing land, creating shrublands rich in berries and insects. When the cleared land naturally replenishes, jays and crows use the young dense trees for nesting sites. Ravens typically use larger trees in denser forests.

Despite the fact that most corvids are not threatened (many even increasing due to human activity) a few species are in danger. For example, the destruction of the Southeast Asian rainforests is endangering mixed-species feeding flocks with members from the family Corvidae. Also, since its semiarid scrubland habitat is an endangered ecosystem, the Florida Scrub-jay has a small and declining population. A number of island species, which are more vulnerable to introduced species and habitat loss, have been driven to extinction, such as the New Zealand Raven, or are threatened, like the Mariana Crow.

The American Crow population of the United States has grown over the years. It is possible that the American Crow, due to humans increasing suitable habitat, will drive out Northwestern and Fish Crows.