|Distribution map of the Corvidae.
Extinct (post-1500) Extinct (pre-1500)
Corvidae is a cosmopolitan family of oscine passerine birds that contains the crows, ravens, rooks, jackdaws, jays, magpies, treepies, choughs, and nutcrackers. In colloquial English, they are known as the crow family, or, in jargon, corvids. Currently 133 species are included in this family. The genus Corvus, including the jackdaws, crows, rooks, and ravens, makes up over a third of the entire family. Corvids are the largest passerines.
Corvids display remarkable intelligence for animals of their size and are among the most intelligent birds thus far studied. Specifically, members of the family have demonstrated self-awareness in mirror tests (European magpies) and tool-making ability (e.g. crows and rooks), skills which until recently were thought to be possessed only by humans and a few other higher mammals. Their total brain-to-body mass ratio is equal to that of non-human great apes and cetaceans, and only slightly lower than that of humans.
They 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 southern tip of South America and the polar ice caps. The majority of the species are found in tropical South and Central America and in southern Asia, with fewer than 10 species each in Africa and Australasia. The genus Corvus has re-entered Australia in relatively recent geological prehistory, with five species and one subspecies there. Several species of raven have reached oceanic islands, and some of these species are now highly threatened with extinction or have already become extinct.
Systematics, taxonomy, and evolution
The name Corvidae for the family was introduced by the English zoologist William Elford Leach in a guide to the contents of the British Museum published in 1820. Over the years, much disagreement has arisen 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. In the late 1970s and throughout the 1980s, Sibley and Ahlquist united the corvids with other taxa in the Corvida, based on DNA–DNA hybridization. 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 Old World 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 of undistinguished lineage, is less clear than previously thought.
The crested jayshrike (Platylophus galericulatus) is traditionally included in the Corvidae, but is not a true member of this family, being closer to the helmetshrikes (Malaconotidae) or shrikes (Laniidae). Likewise, the Hume's ground "jay" (Pseudopodoces humilis) is in fact a member of the tit family Paridae. The following tree represents current insights in the phylogeny of the Crow family according to J. Boyd.
The earliest corvid fossils date to mid-Miocene Europe, 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, US)
- Corvidae gen. et sp. indet. (Edson Early Pliocene of Sherman County, US)
- Protocitta (Early Pleistocene of Reddick, US)
- Corvidae gen. et sp. indet. (Early/Middle Pleistocene of Sicily) - probably belongs in an extant genus
- Henocitta (Arredondo Clay Middle Pleistocene of Williston, US)
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 41 g (1.4 oz) and 21.5 cm (8.5 in). The largest corvids are the common raven (Corvus corax) and the thick-billed raven (Corvus crassirostris), both of which regularly exceed 1,400 grams (3.1 pounds) and 65 cm (26 in).
Corvids occur in most climatic zones. Most are sedentary and do not migrate significantly. However, during a shortage of food, irruptive 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.
Since crows and magpies have benefited and even increased in numbers due to human development, it was suggested that this might cause increased rates of nest predation of smaller bird species, leading to declines. Several studies have shown this concern to be unfounded. One study examined American crows, which had increased in numbers, were a suspect in nest predation of threatened marbled murrelets. 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. Similarly a study examining the decline of British songbirds found no link between Eurasian magpie numbers and population changes of 23 songbird species.
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 "king of the mountain", or "follow the leader", patterns. 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.
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 human food sources. 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 human 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.
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; jackdaws are known to breed in buildings and in rabbit warrens. 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 use several different forms of parental care, including biparental care and cooperative breeding. Cooperative breeding takes place when parents are helped in raising their offspring, usually by relatives but also sometimes by non-related adults. Such helpers at the nest in most cooperatively breeding birds are males, while females join other groups. White-throated magpie-jays are cooperatively breeding corvids where the helpers are mostly female.
Jerison (1973) has suggested that the degree of brain encephalization (the ratio of brain size to body size, EQ) may correlate with an animal's intelligence and cognitive skills. Corvids and psittacids have higher EQ than other bird families, similar to that of the apes. While among the Corvidae, ravens possess the largest brain to body size ratio. In addition to the high EQ, the Corvid's intelligence is boosted by their living environment. First, Corvids were found in some of the harshest environments on Earth, where surviving requires higher intelligence and better adaptations. Secondly, most of the Corvids are omnivorous, suggesting that they are exposed to more different stimuli and environments. Furthermore, many Corvids species live in a large family group and demonstrate high social complexities.
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 meta-analysis testing how often birds invented new ways to acquire food in the wild found corvids to be the most innovative birds. A 2004 review suggests that their cognitive abilities are on par with those of non-human great apes. Despite structural differences, the brains of corvids and great apes both evolved the ability to make geometrical measurements.
Ravens are found to show bystander affiliation and solicited bystander affiliation after aggressive conflicts. Most of the time, bystanders sharing a valuable relationship with the victim are more likely to affiliate with the victim to alleviate the victim's distress ("consolation") as a representation of empathy. Ravens are believed to be able to be sensitive to other's emotions.
Emotion contagion refers to the emotional state matching between individuals. Adriaense et al. (2018) used a bias paradigm to quantify emotional valence, which along with emotional arousal, define emotions. They manipulated the positive and negative affective states in the demonstrator ravens, which showed significantly different responses to the two states: behaving pessimism to the negative states and optimism to the positive states. Then the researchers trained another observer raven to observe the demonstrator's responses first and then present the observer with ambiguous stimuli. The experiment results confirmed the existence of negative emotional contagions in ravens, while the positive emotional contagion remained unclear. Therefore, ravens are capable of discerning the negative emotions in their conspecifics and showing signs of empathy.
Interspecific communications are evolutionarily beneficial for species living in the same environment. Facial expressions are the most used ways by humans to express emotions. Tate et al.(2006) explored the issue of non-human mammals processing the visual cues from face to achieve interspecific communication with humans. Researchers also examined the avian species' capabilities to interpret this non-verbal communication and be sensitive to human emotions. Based on the experimental subject of American Crows' behavioral changes to varying human gazes and facial expressions, Clucas et al.(2013) identified that crows are able to change their behaviors to human emotions. They further suggested that the high intelligence of the crows enables them to adapt well to human-dominated environments.
It is considered difficult to study emotions in animals when humans could not communicate with them. One way to identify animal personality traits is to observe the consistency of the individual's behavior over time and circumstances. For group-living species, there are two opposing hypotheses regarding the assortment of personality within a group: the social niche specialization hypothesis and the conformity hypothesis. To test these two hypotheses, McCune et al. (2018) performed an experiment on the boldness of two species in Corvidae: the Mexican Jay and California Scrub-Jay. Their results confirmed the conformity hypothesis, supported by the significant differences in the group effects.
The individual personality is both determined by genetics and shaped by social contexts. Miller et al. (2016) examined the role of the developmental and social environment in personality formation in common ravens and carrion crows, which are highly social corvids. The researchers highlighted the correlation between social contexts and an individual's consistent behavior over time (personality) by showing that conspecific presence promoted the behavioral similarities between individuals. Therefore, the researchers demonstrated that social contexts had a significant impact on the development of the raven's and crow's personalities.
The social complexity hypothesis is that living in a social group enhances the cognitive abilities of animals. 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, as well as contributing to the spread of information between members of the group.
Consciousness, culture-rudiments, and neurology
The Eurasian magpie is the only non-mammal species known to be able to recognize itself in a mirror test although late research could not replicate this finding. Studies using very similar setups could not find such behaviour in other corvids (e.g., Carrion crows). Magpies have been observed taking part in elaborate grieving rituals, which have been likened to human funerals, including laying grass wreaths. Marc Bekoff, at the University of Colorado, argues that it shows that they are capable of feeling complex emotions, including grief. Furthermore, carrion crows show a neuronal response that correlates with their perception of a stimulus, which some scientists have argued to be an empirical marker of (avian/corvid) sensory consciousness – the conscious perception of sensory input – in the crows which do not have a cerebral cortex. A related study shows that the birds' pallium's neuroarchitecture is reminiscent of the mammalian cortex.
Tool use, memory and complex rational thought
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 (that is, 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 California 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. Researchers have discovered that New Caledonian crows don't just use single objects as tools; they can also construct novel compound tools through assemblage of otherwise non-functional elements. 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.
Implications and specific comparisons with other animals
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.
Contrary to earlier teleological classifications in which they were seen as "highest" songbirds due to their intelligence, current systematics might place corvids, based on their total number of physical characteristics instead of just their brains (which are the most developed of birds), in the lower middle of the passerine evolutionary tree, dependent on which subgroup is chosen as the most derived. 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 [...].
A study found that four-months-old ravens can have physical and social cognitive skills similar to that of adult great apes and concludes that "dynamic of the different influences that, during ontogeny, contributes to adult cognition" is required for the study of cognition.
Corvids are reservoirs (carriers) for the West Nile virus in the United States. 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 US in 1999 was the death of crows in New York.
Relationship with humans
Humans have been able to coexist with many members of the Corvidae family throughout history, most notably crows and ravens (see: Role in myth and culture). These positive interactions have extended into modern times.
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 considered responsible for placing the Sun in the sky.
Due to their carrion diet, the Celtic peoples strongly associated corvids with war, death and the battlefield – their great intelligence meant that they were often considered messengers, or manifestations of the gods such as Bendigeidfran Blessed raven or the Irish Morrigan, underworld deities that may be related to the later Arthurian Fisher King. The Welsh Dream of Rhonabwy illustrates well the association of ravens with war. In many parts of Britain, gatherings of crows, or more often magpies, are counted using the divination rhyme: one for sorrow, two for joy, three for a girl, four for a boy, five for silver, six for gold, seven for a secret never to be told. Another rhyme is: one for sorrow, two for mirth, three for a funeral, four for a birth, five for heaven, six for hell, and seven for the Devil, his own sel'. Cornish superstition holds that when a lone magpie is encountered, it must be loudly greeted with respect.
Various Germanic peoples highly revered the raven and the raven often features as a motif on shields or other war gear in Anglo-Saxon art such as the Sutton Hoo burial and Vendel period era art. The major deity Odin was so associated with ravens throughout history that he gained the kenning "raven god"[b] 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 Sutton Hoo treasure features stylised corvids with scrolled beaks in the decorative enamel work on the shield and purse lid reflecting their common totemic status to the Anglo-Saxons, whose pre-Christian indigenous beliefs were of the same origin as that of the aforementioned Vikings.
The sixth 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 forest.
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 rainforest 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 cause Northwestern crows and fish crows to decline.
- Genus Crypsirina
- Genus Dendrocitta
- Genus Platysmurus
- Genus Temnurus
- Ratchet-tailed treepie, Temnurus temnurus
- Oriental magpies
- Genus Cissa
- Genus Urocissa
- Old World jays
- Stresemann's bushcrow
- Holarctic magpies
- 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
- Tropical Asian species
- Daurian jackdaw, Corvus dauuricus
- Slender-billed crow, Corvus enca
- Palawan crow, Corvus pusillus
- Flores crow, Corvus florensis
- Large-billed crow, Corvus macrorhynchos
- Eastern jungle crow, Corvus levaillantii
- Indian jungle crow, Corvus culminatus
- 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
- Western jackdaw, Corvus monedula
- Fan-tailed raven, Corvus rhipidurus
- Brown-necked raven, Corvus ruficollis
- Hooded crow, Corvus cornix
- Holarctic species
- North and Central American species
- American crow, Corvus brachyrhynchos
- Northwestern crow, Corvus brachyrhynchos 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
- Sinaloa crow, Corvus sinaloae
- Western raven, Corvus (corax) sinuatus
- American crow, Corvus brachyrhynchos
- Tropical African species
- Australian and Melanesian species
- Genus Corvus
- Boreal jays
- New World jays
- Genus Aphelocoma – scrub-jays
- Genus Calocitta – magpie-jays
- Genus Cyanocitta
- 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
- White-tailed jay, Cyanocorax mystacalis
- San Blas jay, Cyanocorax sanblasianus
- Violaceous jay, Cyanocorax violaceus
- Green jay, Cyanocorax ynca
- Yucatan jay, Cyanocorax yucatanicus
- Genus Psilorhinus
- Brown jay, Psilorhinus morio
- Genus Cyanolyca
- Silvery-throated jay, Cyanolyca argentigula
- Black-collared jay, Cyanolyca armillata
- Azure-hooded jay, Cyanolyca cucullata
- White-throated jay, Cyanolyca mirabilis
- Dwarf jay, Cyanolyca nana
- Beautiful jay, Cyanolyca pulchra
- Black-throated jay, Cyanolyca pumilo
- Turquoise jay, Cyanolyca turcosa
- White-collared jay, Cyanolyca viridicyana
- Genus Gymnorhinus
- Pinyon jay, Gymnorhinus cyanocephalus
- Madge, S.; Burn, H. (1993). Crows and Jays. Helm. ISBN 978-1-873403-18-1.
- Robertson, Don (30 January 2000): Bird Families of the World: Corvidae. Retrieved 2007-NOV-10.
- Clayton, Nicola; Emery, Nathan (2005). "Corvid cognition". Current Biology. 15 (3): R80–R81. doi:10.1016/j.cub.2005.01.020. PMID 15694292.
- "IOC World Bird List - Crows, mudnesters, melampittas, Ifrit, birds-of-paradise".
- Emery, N. J.; Clayton, Nicola S. (2004). "The mentality of crows: Convergent evolution of Intelligence in corvids and apes". Science. 306 (5703): 1903–1907. doi:10.1126/science.1098410. ISSN 0036-8075. PMID 15591194. S2CID 9828891.
- "Rooks reveal remarkable tool-use". BBC News. 26 May 2009. Retrieved 2 April 2010.
- Birding in India and South Asia: Corvidae. Retrieved 2007-NOV-10
- Leach, William Elford (1820). "Eleventh Room". Synopsis of the Contents of the British Museum. Vol. 17 (17th ed.). London: British Museum. pp. 67–68. The name of the author is not specified in the document.
- Bock, Walter J. (1994). History and Nomenclature of Avian Family-Group Names. Bulletin of the American Museum of Natural History. Vol. Number 222. New York: American Museum of Natural History. pp. 118, 222. hdl:2246/830.
- Jønsson, Knud A.; Fabre, Pierre-Henri; Ricklefs, Robert E.; Fjeldså, Jon (8 February 2011). "Major global radiation of corvoid birds originated in the proto-Papuan archipelago". Proceedings of the National Academy of Sciences. 108 (6): 2328–2333. Bibcode:2011PNAS..108.2328J. doi:10.1073/pnas.1018956108. ISSN 0027-8424. PMC 3038755. PMID 21262814.
- Jønsson, Knud A.; Fjeldså, Jon (2006). "A phylogenetic supertree of oscine passerine birds (Aves: Passeri)". Zoologica Scripta. 35 (2): 149–186. doi:10.1111/j.1463-6409.2006.00221.x. S2CID 85317440.
- Ericson, Per G. P.; Jansén, Anna-Lee; Johansson, Ulf S.; Ekman, Jan (2005). "Inter-generic relationships of the crows, jays, magpies and allied groups (Aves: Corvidae) based on nucleotide sequence data" (PDF). Journal of Avian Biology. 36 (3): 222–234. CiteSeerX 10.1.1.493.5531. doi:10.1111/j.0908-8857.2001.03409.x.
- Goodwin, D. (1986) Crows of the world. (2nd edition). British Museum of Natural History. ISBN 0-565-00979-6
- James, Helen F.; Ericson, Per G.P.; Slikas, Beth; Lei, Fu-min; Olson, Storrs L. (2003). "Pseudopodoces humilis, a misclassified terrestrial tit (Aves: Paridae) of the Tibetan Plateau: evolutionary consequences of shifting adaptive zones" (PDF). Ibis. 145 (2): 185–202. doi:10.1046/j.1474-919X.2003.00170.x.
- Boyd, J. "Corvoidea genus tree" (PDF). jboyd.net. Retrieved 19 June 2017.
- Mourer-Chauviré, C. C. (2004). "Cenozoic Birds of the World, Part 1: Europe". The Auk. 121 (2): 623. doi:10.1642/0004-8038(2004)121[0623:CBOTWP]2.0.CO;2. S2CID 86482094.
- Proximal right coracoid of a jay-sized bird, perhaps an Holarctic magpie distinct from Pica: Wetmore, Alexander (1937). "The Eared Grebe and other Birds from the Pliocene of Kansas" (PDF). Condor. 39 (1): 36–44. doi:10.2307/1363487. JSTOR 1363487.
- Perrins, Christopher (2003): The New Encyclopedia of Birds Oxford University Press: Oxford ISBN 0-19-852506-0
- Shades of Night: The Aviary Archived 15 April 2006 at the Wayback Machine. Version of 2004-JUL-21. Retrieved 2007-NOV-10.
- Marzluff, John M.; Neatherlin, Eric (2006). "Corvid response to human settlements and campgrounds: Causes, consequences, and challenges for conservation". Biological Conservation. 130 (2): 301–314. doi:10.1016/j.biocon.2005.12.026.
- Thompson, D. L.; Green, R. E.; Gregory, R. D.; Baillie, S. R. (1998). "The widespread declines of songbirds in rural Britain do not correlate with the spread of their avian predators". Proceedings of the Royal Society B. 265 (1410): 2057–2062. doi:10.1098/rspb.1998.0540. PMC 1689492.
- Verhulst, Sion; Salomons, H. Martijn (2004). "Why fight? Socially dominant jackdaws, Corvus monedula, have low fitness". Animal Behaviour. 68 (4): 777–783. doi:10.1016/j.anbehav.2003.12.020. hdl:11370/c1816821-aa56-410e-a222-03b19875501f. S2CID 53199764.
- Gill, F.B. (2003) Ornithology (2nd edition). W.H. Freeman and Company, New York. ISBN 0-7167-2415-4
- Patterson, I. J.; Dunnet, G. M.; Fordham, R. A. (1971). "Ecological studies of the Rook Corvus frugilegus L. in northeast Scotland. Dispersion". J. Appl. Ecol. 8 (3): 815–833. doi:10.2307/2402685. JSTOR 2402685.
- Li, Shou-Hsien; Brown, Jerram L. (2000). "High frequency of extrapair fertilization in a plural breeding bird, the Mexican jay, revealed by DNA microsatellites". Animal Behaviour. 60 (6): 867–877. doi:10.1006/anbe.2000.1554. PMID 11124886. S2CID 34041075.
- Encyclopædia Britannica Online: Corvidae. Free subscription required.
- Cockburn, Andrew (7 June 2006). "Prevalence of different modes of parental care in birds". Proceedings of the Royal Society B: Biological Sciences. 273 (1592): 1375–1383. doi:10.1098/rspb.2005.3458. PMC 1560291. PMID 16777726.
- Bourke, Andrew F. G. (19 May 2014). "Hamilton's rule and the causes of social evolution". Philosophical Transactions of the Royal Society B: Biological Sciences. 369 (1642): 20130362. doi:10.1098/rstb.2013.0362. ISSN 0962-8436. PMC 3982664. PMID 24686934.
- Riehl, Christina (7 December 2013). "Evolutionary routes to non-kin cooperative breeding in birds". Proceedings of the Royal Society B: Biological Sciences. 280 (1772): 20132245. doi:10.1098/rspb.2013.2245. PMC 3813341. PMID 24132311.
- Jerison, Harry J. (1973), "Evolution of the Brain in Birds", Evolution of the Brain and Intelligence, Elsevier, pp. 177–199, doi:10.1016/b978-0-12-385250-2.50018-3, ISBN 978-0-12-385250-2, retrieved 11 March 2021
- Emery, Nathan J.; Clayton, Nicola S. (2004), Rogers, Lesley J.; Kaplan, Gisela (eds.), "Comparing the Complex Cognition of Birds and Primates", Comparative Vertebrate Cognition, Boston, MA: Springer US, pp. 3–55, doi:10.1007/978-1-4419-8913-0_1, ISBN 978-1-4613-4717-0, retrieved 11 March 2021
- Emery, Nathan J (29 January 2006). "Cognitive ornithology: the evolution of avian intelligence". Philosophical Transactions of the Royal Society B: Biological Sciences. 361 (1465): 23–43. doi:10.1098/rstb.2005.1736. ISSN 0962-8436. PMC 1626540. PMID 16553307.
- "Long childhoods and extended parenting help young crows grow smarter". phys.org. Retrieved 2 July 2020.
- Heidt, Amanda (8 June 2020). "Like humans, these big-brained birds may owe their smarts to long childhoods". Science. doi:10.1126/science.abd2209. S2CID 225766325.
- Uomini, Natalie; Fairlie, Joanna; Gray, Russell D.; Griesser, Michael (20 July 2020). "Extended parenting and the evolution of cognition". Philosophical Transactions of the Royal Society B: Biological Sciences. 375 (1803): 20190495. doi:10.1098/rstb.2019.0495. PMC 7293161. PMID 32475334.
- Krushinskii, L. V.; Zorina, Z. A.; Dashevskiy, B. A. (1979). "Ability of birds of the Corvidae family to operate by the empirical dimensions of figures". Zhurnal Vysshei Nervnoi Deiatelnosti Imeni I P Pavlova. 29 (3): 590–7. PMID 112801.
- Rincon, Paul (22 February 2005) Crows and jays top bird IQ scale. BBC.
- Emery, Nathan; Clayton, Nicola (2004). "The Mentality of Crows: Convergent Evolution of Intelligence in Corvids and Apes". Science. 306 (5703): 1903–7. Bibcode:2004Sci...306.1903E. CiteSeerX 10.1.1.299.6596. doi:10.1126/science.1098410. PMID 15591194. S2CID 9828891.
- Fraser, Orlaith N.; Bugnyar, Thomas (12 May 2010). Brosnan, Sarah Frances (ed.). "Do Ravens Show Consolation? Responses to Distressed Others". PLOS ONE. 5 (5): e10605. Bibcode:2010PLoSO...510605F. doi:10.1371/journal.pone.0010605. ISSN 1932-6203. PMC 2868892. PMID 20485685.
- Edgar, Joanne L.; Nicol, Christine J. (December 2018). "Socially-mediated arousal and contagion within domestic chick broods". Scientific Reports. 8 (1): 10509. Bibcode:2018NatSR...810509E. doi:10.1038/s41598-018-28923-8. ISSN 2045-2322. PMC 6043517. PMID 30002482.
- Adriaense, Jessie E. C.; Martin, Jordan S.; Schiestl, Martina; Lamm, Claus; Bugnyar, Thomas (4 June 2019). "Negative emotional contagion and cognitive bias in common ravens (Corvus corax)". Proceedings of the National Academy of Sciences. 116 (23): 11547–11552. doi:10.1073/pnas.1817066116. ISSN 0027-8424. PMC 6561263. PMID 31110007.
- Tate, Andrew J; Fischer, Hanno; Leigh, Andrea E; Kendrick, Keith M (29 December 2006). "Behavioural and neurophysiological evidence for face identity and face emotion processing in animals". Philosophical Transactions of the Royal Society B: Biological Sciences. 361 (1476): 2155–2172. doi:10.1098/rstb.2006.1937. ISSN 0962-8436. PMC 1764842. PMID 17118930.
- Clucas, Barbara; Marzluff, John M.; Mackovjak, David; Palmquist, Ila (April 2013). Ebensperger, L. (ed.). "Do American Crows Pay Attention to Human Gaze and Facial Expressions?". Ethology. 119 (4): 296–302. doi:10.1111/eth.12064.
- Both, Christiaan; Dingemanse, Niels J.; Drent, Piet J.; Tinbergen, Joost M. (July 2005). "Pairs of extreme avian personalities have highest reproductive success". Journal of Animal Ecology. 74 (4): 667–674. doi:10.1111/j.1365-2656.2005.00962.x. ISSN 0021-8790.
- Gosling, Samuel D. (2001). "From mice to men: What can we learn about personality from animal research?". Psychological Bulletin. 127 (1): 45–86. doi:10.1037/0033-2909.127.1.45. ISSN 1939-1455. PMID 11271756.
- Bergmüller, Ralph; Taborsky, Michael (September 2010). "Animal personality due to social niche specialisation". Trends in Ecology & Evolution. 25 (9): 504–511. doi:10.1016/j.tree.2010.06.012. PMID 20638151.
- Judith, University of Helsinki, Department of Finnish, Finno-Ugrian and Scandinavian Studies Koski, Sonja Burkart (6 March 2015). Common marmosets show social plasticity and group-level similarity in personality. Nature Publishing Group. OCLC 951545397.
- Miller, Rachael; Laskowski, Kate L.; Schiestl, Martina; Bugnyar, Thomas; Schwab, Christine (5 February 2016). Hemelrijk, Charlotte K (ed.). "Socially Driven Consistent Behavioural Differences during Development in Common Ravens and Carrion Crows". PLOS ONE. 11 (2): e0148822. Bibcode:2016PLoSO..1148822M. doi:10.1371/journal.pone.0148822. ISSN 1932-6203. PMC 4746062. PMID 26848954.
- Bond, Alan B.; Kamil, Alan C.; Balda, Russell P. (2003). "Social complexity and transitive inference in corvids" (PDF). Animal Behaviour. 65 (3): 479–487. doi:10.1006/anbe.2003.2101. S2CID 9836564. Archived from the original (PDF) on 16 February 2008. Retrieved 10 November 2007.
- Prior, Helmut; Schwarz, Ariane; Güntürkün, Onur (2008). De Waal, Frans (ed.). "Mirror-Induced Behavior in the Magpie (Pica pica): Evidence of Self-Recognition". PLOS Biology. 6 (8): e202. doi:10.1371/journal.pbio.0060202. PMC 2517622. PMID 18715117.
- Soler, M.; Colmero, J. M.; Pérez-Contreras, T.; Peralta-Sánchez, J. M. (2020). "Replication of the mirror mark test experiment in the magpie (Pica pica) does not provide evidence of self-recognition". Journal of Comparative Psychology. 134 (4): 363–371. doi:10.1037/com0000223. PMID 32406720. S2CID 218636079.
- Brecht, Katharina F.; Müller, Jan; Nieder, Andreas (November 2020). "Carrion crows (Corvus corone corone) fail the mirror mark test yet again". Journal of Comparative Psychology. 134 (4): 372–378. doi:10.1037/com0000231. ISSN 1939-2087. PMID 32463251. S2CID 218976227.
- Vanhooland, Lisa-Claire; Bugnyar, Thomas; Massen, Jorg J. M. (May 2020). "Crows (Corvus corone ssp.) check contingency in a mirror yet fail the mirror-mark test". Journal of Comparative Psychology. 134 (2): 158–169. doi:10.1037/com0000195. ISSN 1939-2087. PMID 31589059. S2CID 203850271.
- Animal emotions, wild justice and why they matter: Grieving magpies, a pissy baboon, and empathic elephants Emotion, Space and Society xxx (2009) 1–4, Marc Bekoff
- "Researchers show conscious processes in birds' brains for the first time". phys.org. Retrieved 9 October 2020.
- Nieder, Andreas; Wagener, Lysann; Rinnert, Paul (25 September 2020). "A neural correlate of sensory consciousness in a corvid bird". Science. 369 (6511): 1626–1629. Bibcode:2020Sci...369.1626N. doi:10.1126/science.abb1447. ISSN 0036-8075. PMID 32973028. S2CID 221881862. Retrieved 9 October 2020.
- Stetka, Bret. "Bird Brains Are Far More Humanlike Than Once Thought". Scientific American. Retrieved 23 October 2020.
- Stacho, Martin; Herold, Christina; Rook, Noemi; Wagner, Hermann; Axer, Markus; Amunts, Katrin; Güntürkün, Onur (25 September 2020). "A cortex-like canonical circuit in the avian forebrain". Science. 369 (6511): eabc5534. doi:10.1126/science.abc5534. ISSN 0036-8075. PMID 32973004. S2CID 221882087. Retrieved 16 October 2020.
- "Attenborough – Crows in the City". YouTube.com. 12 February 2007. Archived from the original on 7 November 2021. Retrieved 9 March 2013.
- Burnell, Kristi L.; Tomback, Diane F. (1985). "Steller's jays steal Grey Jay caches: field and laboratory observation". Auk. 102 (2): 417–419. doi:10.2307/4086793. JSTOR 4086793.
- Waite, Thomas A. (1992). "Social hoarding and a load size-distance relationship in Gray Jays". The Condor. 94 (4): 995–998. doi:10.2307/1369297. JSTOR 1369297. S2CID 86971311.
- Owen, James (9 December 2004) Crows as Clever as Great Apes, Study Says. National Geographic News, Retrieved 2007-NOV-10.
- Morelle, Rebecca (20 April 2010). "BBC On-line: Clever New Caledonian crows can use three tools". BBC News. Retrieved 9 March 2013.
- Baraniuk, Chris (12 December 2020). "Crows could be the smartest animal other than primates". BBC. Retrieved 31 August 2020.
- Bayern, A. M. P. von; Danel, S.; Auersperg, A. M. I.; Mioduszewska, B.; Kacelnik, A. (24 October 2018). "Compound tool construction by New Caledonian crows". Scientific Reports. 8 (1): 15676. Bibcode:2018NatSR...815676B. doi:10.1038/s41598-018-33458-z. ISSN 2045-2322. PMC 6200727. PMID 30356096.
- Starr, Michelle (25 October 2018). "Crows Can Build Compound Tools Out of Multiple Parts, And Are You Even Surprised". ScienceAlert. Retrieved 4 April 2020.
- Jones, Juli E.; Antoniadis, Elena; Shettleworth, Sara J.; Kamil, Alan C. (2002). "A Comparative Study of Geometric Rule Learning by Nutcrackers (Nucifraga columbiana), Pigeons (Columba livia), and Jackdaws (Corvus monedula)" (PDF). Journal of Comparative Psychology. 116 (4): 350–356. doi:10.1037/0735-7036.116.4.350. PMID 12539930.
- "Cognitive performance of four-months-old ravens may parallel adult apes". phys.org. Retrieved 17 January 2021.
- Eidson, M; Komar, N; Sorhage, F; Nelson, R; Talbot, T; Mostashari, F; McLean, R; West Nile Virus Avian Mortality Surveillance Group (2001). "Crow deaths as a sentinel surveillance system for West Nile virus in the northeastern United States, 1999". Emerging Infectious Diseases. 7 (4): 615–20. doi:10.3201/eid0704.010402. PMC 2631775. PMID 11585521.
- "Raven Releasing the Sun". Native Online. Native Online. Retrieved 30 July 2020.
- Chappell, J. (2006). "Living with the Trickster: Crows, Ravens, and Human Culture". PLOS Biology. 4 (1): e14. doi:10.1371/journal.pbio.0040014. PMC 1326277.
- Lee, T. M.; Soh, M. C. K.; Sodhi, N.; Koh, L. P.; Lim, S. L. H. (2005). "Effects of habitat disturbance on mixed species bird flocks in a tropical sub-montane rainforest". Biological Conservation. 122 (2): 193–204. doi:10.1016/j.biocon.2004.07.005.
- BirdLife International. (2016). Aphelocoma coerulescens. The IUCN Red List of Threatened Species. doi:10.2305/IUCN.UK.2016-3.RLTS.T22705629A94028132.en
- Breininger, D. R.; Toland, B.; Oddy, D. M.; Legare, M. L. (2006). "Landcover characterizations and Florida scrub-jay (Aphelocoma coerulescens) population dynamics" (PDF). Biological Conservation. 128 (2): 169–181. doi:10.1016/j.biocon.2005.09.026.
- Marzluff, John M.; Angell, T. (2005). In the Company of Crows and Ravens. New Haven, Connecticut: Yale University Press. ISBN 978-0-300-10076-1.
- "Species Updates – IOC World Bird List". Retrieved 29 May 2021.
- Charles Sibley & Jon Edward Ahlquist (1991): Phylogeny and Classification of Birds: A Study in Molecular Evolution. New Haven, Conn.: Yale University Press. ISBN 0-300-04085-7.
- Corvidae videos on the Internet Bird Collection
- corvids.de – Corvids-Literature-Database
- Corvidae sounds on xeno-canto.org
- Corvid Corner A site about the Corvidae
- AvesNoir A site about corvids in art, culture, and literature.
- Discovery of species-wide tool use in the Hawaiian crow
- Rooks reveal remarkable tool use
- Clever New Caledonian crows can use three tools
- Talking Eurasian magpie Pica pica
- Rare crow shows a talent for tool use