Corvidae

Corvidae
Temporal range: Middle Miocene to present
Blue jay
Cyanocitta cristata
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Superfamily: Corvoidea
Family: Corvidae
Vigors, 1825
Genera

see text

Distribution map of the Corvidae.
  Native   (Re)Introduced
  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.[1][2][3] In common English, they are known as the crow family, or, more technically, corvids. Over 120 species are described. The genus Corvus, including the jackdaws, crows, and ravens, makes up over a third of the entire family.

They are considered the most intelligent of the birds, and among the most intelligent of all animals,[4] having demonstrated self-awareness in mirror tests (European magpies) and tool-making ability (crows, rooks[5])—skills until recently regarded as solely the province of humans and a few other higher mammals. Their total brain-to-body mass ratio is equal to that of great apes and cetaceans, and only slightly lower than in humans.[6]

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 tip of South America and the polar ice caps.[3] 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 and Australasia. The genus Corvus has re-entered Australia in relatively recent geological prehistory, with five species and one subspecies there.

Systematics, taxonomy, and evolution

Rufous treepie, Dendrocitta vagabunda
Yellow-billed blue magpie, Urocissa flavirostris
Eurasian jay (Garrulus glandarius)
Eurasian magpie, Pica pica
Plush-crested jay, Cyanocorax chrysops
Common raven, Corvus corax
Hooded crow, Corvus cornix
Thick-billed raven, Corvus crassirostris

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,[2] 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.[7]

Clarification of the interrelationships of the corvids has been achieved based on cladistic analysis of several DNA sequences.[7][8] 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.

FAMILY CORVIDAE

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 helmetshrikes (Malaconotidae) or shrikes (Laniidae); it is best considered Corvidae incertae sedis for the time being.[1][9] Likewise, the Hume's ground "jay" (Pseudopodoces humilis) is in fact a member of the tit family Paridae.[10]

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:

In addition, there are numerous fossil species of extant genera since the Mio–Pliocene, mainly European Corvus.[lower-alpha 1]

Biology

Morphology

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.[12] 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).

Species can be identified based on size, shape, and geography; however, some, especially the Australian crows, are best identified by their raucous calls.[2]

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.[2] When species are migratory, they will form large flocks in the fall (around August in the Northern Hemisphere) and travel south.[13]

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.[14]

Food and feeding

Corvids are highly opportunistic foragers. Here a jungle crow feeds on a shark carcass.

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.[14]

Some corvids are predators of other birds. During the wintering months, corvids typically form foraging flocks.[2] However, some crows also eat many agricultural pests including cutworms, wireworms, grasshoppers, and harmful weeds.[13] 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.[14]

Reproduction

A gray jay pair feeding their chicks.

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.[15] 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.[16] Males and females build large nests together in trees or on ledges. The male will also feed the female during incubation.[17] 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.[18] 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.[19]

Social life

Some corvids have strong organization and community groups. Jackdaws, for example, have a strong social hierarchy, and are facultatively colonial during breeding.[18] 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.[20]

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.[13]

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.[13]

Intelligence

The brain-to-body weight ratios of corvid brains are among the largest in birds, equal to that of most great apes and cetaceans, and only slightly lower than a human.[6] 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.[3]

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.[21] A meta-analysis testing how often birds invented new ways to acquire food in the wild found corvids to be the most innovative birds.[22] A 2004 review suggests that their cognitive abilities are on par with those of great apes.[23] 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.[24]

The Eurasian magpie is the only non-mammal species known to be able to recognize itself in a mirror test.[4] Magpies have been observed taking part in elaborate grieving rituals, which have been likened to human funerals, including laying grass wreaths.[25][26] Marc Bekoff, at the University of Colorado, argues that it shows that they are capable of feeling complex emotions, including grief.[26]

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.[27] 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.[28][29] 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.[30]

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.[3]

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.[31] 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.[3]

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.[32]

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.[13]

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.[7] 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 [...].[2]

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 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.[33]

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 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 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. Cornish superstition holds that when a magpie is encountered, it must be loudly greeted with respect.

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"[lower-alpha 2] 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.[34] 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 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.

The Hawaiian crow is extinct in the wild as a result of habitat loss and other factors.

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.[14]

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.[14]

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.[35] Also, since its semiarid scrubland habitat is an endangered ecosystem, the Florida scrub jay has a small and declining population.[36][37] 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 crows and fish crows.[38]

Notes

  1. ↑ See the genus accounts for more.
  2. ↑ e.g. Icelandic: hrafnaguð, as per the Gylfaginning.

References

  1. 1 2 Madge, S.; Burn, H. (1993). Crows and Jays. Helm. ISBN 1-873403-18-6.
  2. 1 2 3 4 5 6 Robertson, Don (30 January 2000): Bird Families of the World: Corvidae. Retrieved 2007-NOV-10.
  3. 1 2 3 4 5 Clayton, Nicola; Emery, Nathan (2005). "Corvid cognition". Current Biology. 15 (3): R80–R81. doi:10.1016/j.cub.2005.01.020. PMID 15694292.
  4. 1 2 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 2517622Freely accessible. PMID 18715117.
  5. ↑ "Rooks reveal remarkable tool-use". BBC News. 26 May 2009. Retrieved 2 April 2010.
  6. 1 2 Birding in India and South Asia: Corvidae. Retrieved 2007-NOV-10
  7. 1 2 3 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.
  8. ↑ 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. doi:10.1111/j.0908-8857.2001.03409.x.
  9. ↑ Goodwin, D. (1986) Crows of the world. (2nd edition). British Museum of Natural History. ISBN 0-565-00979-6
  10. ↑ 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.
  11. ↑ Proximal right coracoid of a jay-sized bird, perhaps an Holarctic magpie distinct from Pica: Wetmore, Alexander. "The Eared Grebe and other Birds from the Pliocene of Kansas" (PDF). Condor. 39 (1): 40. doi:10.2307/1363487.
  12. ↑ Perrins, Christopher (2003): The New Encyclopedia of Birds Oxford University Press: Oxford ISBN 0-19-852506-0
  13. 1 2 3 4 5 Shades of Night: The Aviary. Version of 2004-JUL-21. Retrieved 2007-NOV-10.
  14. 1 2 3 4 5 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.
  15. ↑ 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.
  16. ↑ 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.
  17. ↑ Encyclopædia Britannica Online: Corvidae. Free subscription required.
  18. 1 2 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.
  19. ↑ Berg, Elena C. (2005). "Parentage and reproductive success in the white-throated magpie-jay, Calocitta formosa, a cooperative breeder with female helpers". Animal Behaviour. 70 (2): 375–385. doi:10.1016/j.anbehav.2004.11.008.
  20. ↑ Gill, F.B. (2003) Ornithology (2nd edition). W.H. Freeman and Company, New York. ISBN 0-7167-2415-4
  21. ↑ 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.
  22. ↑ Rincon, Paul (22 February 2005) Crows and jays top bird IQ scale. BBC.
  23. ↑ Emery, Nathan; Clayton, Nicola (2004). "The Mentality of Crows: Convergent Evolution of Intelligence in Corvids and Apes". Science. 306 (5703): 1903–7. doi:10.1126/science.1098410. PMID 15591194.
  24. ↑ 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.
  25. ↑ Magpies grieve for their dead (and even turn up for funerals) By DAVID DERBYSHIRE FOR MAILONLINE, UPDATED: 01:57, 24 October 2009
  26. 1 2 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
  27. ↑ "Attenborough - Crows in the City". YouTube.com. 12 February 2007. Retrieved 9 March 2013.
  28. ↑ 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.
  29. ↑ 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.
  30. ↑ Owen, James (9 December 2004) Crows as Clever as Great Apes, Study Says. National Geographic News, Retrieved 2007-NOV-10.
  31. ↑ Morelle, Rebecca (20 April 2010). "BBC On-line: Clever New Caledonian crows can use three tools". BBC News. Retrieved 9 March 2013.
  32. ↑ 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.
  33. ↑ 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 2631775Freely accessible. PMID 11585521.
  34. ↑ Chappell, J. (2006). "Living with the Trickster: Crows, Ravens, and Human Culture". PLoS Biology. 4 (1): e14. doi:10.1371/journal.pbio.0040014. PMC 1326277Freely accessible.
  35. ↑ 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.
  36. ↑ BirdLife International (2004). "Aphelocoma coerulescens". IUCN Red List of Threatened Species. Version 2006. International Union for Conservation of Nature. Retrieved 11 May 2006.
  37. ↑ 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.
  38. ↑ Marzluff, John M.; Angell, T. (2005). In the Company of Crows and Ravens. New Haven, Connecticut: Yale University Press. ISBN 0-300-10076-0.

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