Golden eagle

For other uses, see Golden Eagle (disambiguation).

Golden eagle
Adult of the North American subspecies Aquila chrysaetos canadensis
call of a Golden Eagle
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Accipitriformes
Family: Accipitridae
Genus: Aquila
Species: A. chrysaetos
Binomial name
Aquila chrysaetos
(Linnaeus, 1758)
Range of A. chrysaetos      Breeding range     Year-round range     Wintering range
Synonyms

Falco chrysaëtos Linnaeus, 1758

The golden eagle (Aquila chrysaetos) is one of the best-known birds of prey in the Northern Hemisphere. It is the most widely distributed species of eagle. Like all eagles, it belongs to the family Accipitridae. These birds are dark brown, with lighter golden-brown plumage on their napes. Immature eagles of this species typically have white on the tail and often have white markings on the wings. Golden eagles use their agility and speed combined with powerful feet and massive, sharp talons to snatch up a variety of prey (mainly hares, rabbits, marmots and other ground squirrels).[2]

Golden eagles maintain home ranges or territories that may be as large as 200 km2 (77 sq mi). They build large nests in high places (mainly cliffs) to which they may return for several breeding years. Most breeding activities take place in the spring; they are monogamous and may remain together for several years or possibly for life. Females lay up to four eggs, and then incubate them for six weeks. Typically, one or two young survive to fledge in about three months. These juvenile golden eagles usually attain full independence in the fall, after which they wander widely until establishing a territory for themselves in four to five years.

Once widespread across the Holarctic, it has disappeared from many areas which are now more heavily populated by humans. Despite being extirpated from or uncommon in some of its former range, the species is still fairly ubiquitous, being present in sizeable stretches of Eurasia, North America, and parts of North Africa. It is the largest and least populous of the five species of true accipitrid to occur as a breeding species in both the Palearctic and the Nearctic.[3]

For centuries, this species has been one of the most highly regarded birds used in falconry, with the Eurasian subspecies having been used to hunt and kill prey such as gray wolves (Canis lupus) in some native communities. Due to its hunting prowess, the golden eagle is regarded with great mystic reverence in some ancient, tribal cultures. The golden eagle is one of the most extensively studied species of raptor in the world in some parts of its range, such as the Western United States and the Western Palearctic.

Description

In flight
Subadult, note white in tail and dark neck

The golden eagle is a very large, dark brown raptor with broad wings, ranging from 66 to 102 cm (26 to 40 in) in length and from 1.8 to 2.34 m (5 ft 11 in to 7 ft 8 in) in wingspan.[4][5][6][7] This species' wingspan is the fifth largest amongst extant eagle species.[5] In the largest race (A. c. daphanea) males and females weigh typically 4.05 kg (8.9 lb) and 6.35 kg (14.0 lb). In the smallest subspecies, A. c. japonica, males weigh 2.5 kg (5.5 lb) and females 3.25 kg (7.2 lb).[2] In the species overall, males may average around 3.6 kg (7.9 lb) and females around 5.1 kg (11 lb).[8] The maximum size of this species is a matter of some debate. Large races are the heaviest representatives of the Aquila genus and this species is on average the seventh-heaviest living eagle species. The golden eagle ranks as the second heaviest breeding eagle in North America, Europe and Africa but the fourth heaviest in Asia.[3][5] For some time, the largest known mass authenticated for a wild female was the specimen from the nominate race which weighed around 6.7 kg (15 lb) and spanned 2.55 m (8 ft 4 in) across the wings.[9] American golden eagles are typically somewhat smaller than the large Eurasian races, but a massive female that was banded and released in 2006 around Wyoming’s Bridger-Teton National Forest became the heaviest wild golden eagle on record, at 7.7 kg (17 lb).[10] No comprehensive range of weights are known for the largest subspecies (A. c. daphanea).[9] Captive birds have been measured up to a wingspan of 2.81 m (9 ft 3 in) and a mass of 12.1 kg (27 lb) (the latter figure was for an eagle bred for the purposes of falconry which tend to be unnaturally heavy), respectively.[9] The standard measurements of the species include a wing chord length of 52–72 cm (20–28 in), a tail length of 26.5–38 cm (10.4–15.0 in) and a tarsus length of 9.4–12.2 cm (3.7–4.8 in).[5] The culmen reportedly averages around 4.5 cm (1.8 in), with a range of 3.6 to 5 cm (1.4 to 2.0 in) and the bill length from the gape measures around 6 cm (2.4 in).[11][12] The long, straight and powerful hallux-claw (or hind claw, the equivalent to the big toe) can range from 4.5 to 6.34 cm (1.77 to 2.50 in), being about one centimeter more than the hallux-claw of a bald eagle (Haliaeetus leucocephalus) and a little more than one cm less than a harpy eagle (Harpia harpyja).[13][14][15] The sexes are similar in plumage but are considerably dimorphic in size. Females are rather larger than males with the differences increasing as the body size increases across the races. The large Himalayan golden eagles females are about 37% heavier and nearly 9% longer in wing length than the males of the race compared with the small Japanese golden eagles where females are a relatively modest 26% heavier and around 6% longer in wing length than males.[2][16]

The formidable foot and talons of a golden eagle

Adults are primarily dark brown in color, with a paler, typically golden color (the source of the species’ common name) on the back of the crown and nape, and some grey on the inner-wing and tail.[17] There are subtle differences in coloration among the races, described below. Unlike in other Aquila species, where the tarsal feathers are typically of a similar color to the rest of the plumage, the tarsal feathers of golden eagles tend to be paler, ranging from light golden to white.[2] In addition, some full-grown birds (especially in North America) have white "epaulettes" on the upper part of each scapular feather tract.[3][18] The bill is dark at the tip, fading to a lighter horn color, with a yellow cere.[19] As in many acciptrids, the bare portion of the feet is yellow.[5] This species moults gradually beginning in March or April until September or October each year. Moulting usually decreases in winter. Moult of the contour feathers begins on the head and neck region and process along the feather tracts in a general anterior-posterior direction. Feathers on head, neck, back and scapulars may be replaced annually. With large feathers of the wing and tail, moult beginning with innermost feather and proceeds outwards in a straightforward manner known as "descendant" moult.[18]

The juvenile golden eagle is similar to the adult but tends to be darker, appearing black on the back especially in East Asia. Compared to adults, juveniles have a more unfaded color. Young birds are white for about two-thirds of their tail length ending with a broad, black terminal band.[16] Occasionally, juvenile eagles have white patches on the remiges at the bases of the inner primaries and the outer secondaries, forming a crescent marking on the wings which tend to be divided by darker feathers.[2][20] Rarely, juvenile birds may have only trace amounts of white on the tail. Compared to the relatively consistently white tail, the white patches on the wing are extremely variable and some juveniles have almost no white visible. Juveniles of less than 12 months of age tend to have the most extensive amount of white to the plumage.[18] By their second summer, the white underwing coverts are usually replaced by a characteristic rusty-brown color. By the third summer, the upper-wing coverts are largely replaced by dark brown feathers, however not all feathers moult at once giving many juvenile birds a grizzled pattern. The tail also follows a similar pattern of maturation.[18][20] Due to the amount of variability in different individuals, juvenile eagles cannot be reliably aged on sight alone.[21] Many golden eagles still have white on the tail during their first attempt at nesting.[22] The final adult plumage is not fully attained until the birds are between 5 and a half and 6 and a half years old.[21]

Voice

Calls of a Golden Eagle, recorded at Kinlochewe, Scotland, July 1969

While many accipitrids are not known for their strong voices, golden eagles have a particular tendency for silence, even while breeding.[23] Some vocalization, however, has been recorded, and these normally are centering on the nesting period. The voice of the golden eagle is considered weak, high and shrill, even being emphatically described as “quite pathetic” and “puppy-like”, and as somewhat incongruous considering the formidable size and nature of the species.[5] Most known vocalizations seem to function as contact calls between eagles, sometimes adults to their offspring, occasionally territorial birds to intruders and rarely between a breeding pair. In Western Montana, nine distinct calls were noted: a chirp, a seeir, a pssa, a skonk, a cluck, a wonk, a honk and a hiss.[24]

Flight physiology

Golden eagle flying in dihedral with food

Golden eagles are sometimes considered the most superlative fliers among eagles and perhaps among all raptorial birds.[23] They are equipped with broad, long wings with somewhat finger-like indentations on the tips of the wing.[5] Golden eagles are unique among their genus in that they often fly in a slight dihedral, which means the wings are often held in a slight, upturned V.[5] When they must engage in flapping flight, golden eagles appear at their most labored but this flight method is generally less common than soaring or gliding flights.[2] Flapping flight usually consists of 6–8 deep wing-beats, interspersed with 2 to 3 second glides.[2] While soaring the wings and tail are held in one plane with the primary tips often spread.[25] A typical, unhurried soaring speed in golden eagles is around 45–52 kilometers per hour (28–32 mph).[26] When hunting or displaying, the golden eagle is capable of very fast gliding, attaining speeds of up to 190 km/h (120 mph).[2][25] When diving (or stooping) in the direction of prey or during territorial displays, the eagle holds its wings tight and partially closed against its body and the legs up against its tail. In a full stoop, a golden eagle can reach spectacular speeds of up to 240 to 320 kilometers per hour (150 to 200 mph) when diving after prey. Although less agile and maneuverable, the golden eagle is apparently quite the equal and possibly even the superior of the peregrine falcon’s stooping and gliding speeds.[3][27] This places the golden eagle as the one of the two fastest moving living animals on earth.[9] Although most flight in golden eagles has a purpose (e.g., territoriality, hunting, etc.), some flights (such as those by solitary birds or between well-established breeding pairs) seems to function merely as acts of playfulness.[3][23][28]

Confusion with other species

Golden eagles are readily distinguished by their brown plumage, paler than most other Aquila, and pale nape patch
Close-up of head

Size readily distinguishes this species from most other raptors when it is seen well. Most other raptors are considerably smaller. Buteo hawks, which are perhaps most similar to the golden eagle in structure among the species outside of the “booted eagle” group, are often amongst the larger very common raptors. However, a mid-sized Buteo is dwarfed by a golden eagle, as an adult female eagle has a wingspan of about twice the width and weighs around five times more. Buteos are also usually distinctly paler below, although some species occur in “dark morphs” which can be even darker than a golden eagle.[5][29] Only some Old World vultures and the California condor (Gymnogyps californianus) (among the other raptorial birds this eagle co-exists with) are distinctly larger than the golden eagle, with longer, broader wings, typically held more evenly in a slower, less forceful flight and often have dramatically different color patterns. The turkey vulture (Cathartes aura) is a potential confusion species in North America from a great distance, as it is a large species that (like the golden eagle) often flies with a pronounced dihedral but is easily separated by its less controlled, forceful flying style and its smaller, thinner body, much smaller head and, at closer range, its slaty black-brown color and silvery wing secondaries.[30] Compared to Haliaeetus eagles, the golden eagle has wings that are only somewhat more slender but are more hawk-like and lack the flat, plank-like wing positioning seen in the other genus.[30] Large northern Haliaeetus usually have a larger bill and larger head which more distinctly protrudes than that of a golden eagle’s in flight. The tail of the golden eagle is longer on average than those of Haliaeetus eagles, appearing to be two or three times the length of the head in soaring flight as opposed to the other eagles where the head is often more than twice the length of the tail.[30] Confusion is most likely between juvenile Haliaeetus and golden eagles since the adult golden has a more solidly golden-brown coloration and all Haliaeetus eagles have obvious distinctive plumages as adults. Haliaeetus eagles are often heavily streaked in their juvenile phase. Juvenile golden eagles can show large patches of white to the wings and tail that are quite different than the random, sometimes large and splotchy-looking distribution of white typical of juvenile Haliaeetus.[5][30]

Distinguishing the golden eagle from other Aquila eagles in Eurasia is potentially a greater identification problem. This identification may rely on the golden's relatively long tail and patterns of white or gray on the wings and tail. Other Aquila eagles do not generally fly in a pronounced dihedral as do golden eagles. At close range, the golden to rufous nape-shawl of the golden eagle is distinctive from other Aquila. Most other Aquila eagles are darker looking in plumage, although the smaller tawny eagle (A. rapax) is often paler than the golden eagle (overlap in range verified only in Bale Mountains, Ethiopia). Among Eurasian Aquila, the adult eastern imperial (A. heliaca) and Spanish imperial eagle (A. adalberti) come closest to attaining similar sizes as golden eagles but this species pair are distinguished by their relatively longer neck, flatter wings in flight, white markings on their shoulder forewing-coverts, paler cream-straw colored nape patch and generally darker coloration. Juvenile imperial eagles are much paler overall (caramel-cream in the Spanish; cream and tawny streaks in the Eastern) and are not likely to be confused.[5] Steppe eagles (A. nipalensis) can also be nearly golden eagle-sized but are more compact and smaller headed than a golden eagle with little color variation to their dark earth-brown plumage but for juvenile birds which have distinctive cream-colored bands running through their coverts and secondaries.[3] Verreaux's eagle (A. verreauxii) are most similar in size and body shape to the golden, with the Verreaux's being slightly longer overall but marginally less heavy and long-winged than the golden eagle.[5] The plumage is very distinctly different, however, as Verreaux's eagles are almost entirely jet-black but for some striking, contrasting white on the wing primaries, shoulders and upper-wing.[5] This closely related species is known to co-occur with the golden eagle only in the Bale Mountains of Ethiopia.[2] Other booted eagles in the golden eagle’s range are unlikely to be confused, due to the differences in size and form. Among the Aquila genus, only the long-winged and tailed wedge-tailed eagle (A. audax) of Australasia notably exceeds the golden eagle in average wingspan and length. However, the wedge-tailed eagle is a slightly less heavy bird.[9]

Taxonomy and systematics

This species was first described by Linnaeus in his 1758 Systema naturae as Falco chrysaetos.[31] Since birds were grouped largely on superficial characteristics at that time, many species were grouped by Linnaeus in the Falco genus. The type locality was given simply as "Europa"; it was later fixed to Sweden. It was moved to the new genus Aquila by French ornithologist Mathurin Jacques Brisson in 1760.[32] Aquila is Latin for "eagle", possibly derived from aquilus, "dark in colour" and chrysaetos is Ancient Greek for the golden eagle from khrusos, "gold" and aetos, "eagle".[33]

The golden eagle is part of a broad group of raptors called “booted eagles” which are defined by the feature that all species have feathering over their tarsus, unlike many other accipitrids which have bare legs. Included in this group are all species described as “hawk eagles” including the genera Spizaetus and Nisaetus, as well as assorted monotypical genera such as Oroaetus, Lophaetus, Stephanoaetus, Polemaetus, Lophotriorchis and Ictinaetus. The genus Aquila is distributed across every continent but for South America and Antarctica. Up to 20 species have been classified in the genus but the taxonomic placement of some of the traditional species has been questioned as of late. Traditionally, the Aquila eagles have been grouped superficially as largish, mainly brownish or dark-colored booted eagles that vary little in transition from their juvenile to their adult plumages. Genetic research has recently indicated the golden eagle is included in a clade with Verreaux's eagle in Africa as well as the Gurney's eagle (A. gurneyi) and the wedge-tailed eagle (clearly part of an Australasian radiation of the lineage). This identification of this particular clade has long been suspected based on similar morphological characteristics amongst these large-bodied species.[3] More surprisingly, the smaller, much paler-bellied sister species Bonelli's eagle (A. fasciatus) and African hawk-eagle (A. spilogaster), previously included in the Hieraaetus genus, have been revealed to be genetically much closer to the Verreaux's and golden eagle lineage than to other species traditionally included in the Aquila genus.[2][34][35] Other largish Aquila species, the eastern imperial, the Spanish imperial, the tawny and the steppe eagles, are now thought to be separate, close-knit clade, which attained some similar characteristics to the prior clade via convergent evolution.[34][35] Genetically, the “spotted eagles” (A. pomarina, hasata & clanga), have been discovered to be more closely related to the long-crested eagle (Lophaetus occipitalis) and the black eagle (Ictinaetus malayensis), and many generic reassignments have been advocated.[34][36] The Hieraaetus genus, including the booted eagle (H. pennatus), little eagle (H. morphnoides) and Ayres's hawk-eagle (H. ayresii), consists of much smaller species, that are in fact smallest birds called eagles outside of the unrelated Spilornis serpent-eagle genus. This genus has recently been eliminated by many authorities and are now occasionally also included in Aquila, although not all ornithological unions have followed this suit in this re-classification.[2][35][37] The small-bodied Wahlberg's eagle (H. wahlbergi) has been traditionally considered a Aquila species due to its lack of change from juvenile to adult plumage and brownish color but it is actually genetically aligned to the Hieraaetus lineage.[34][38] Cassin's hawk-eagle (H. africanus) is also probably closely related to the Hieraaetus group rather than the Spizaetus/Nisaetus “hawk-eagle” group (in which it was previously classified) which is not known to have radiated to Africa.[39]

Subspecies and distribution

A captive Aquila chrysaetos canadensis shows the typical rusty coloration of the race

There are six extant subspecies of golden eagle that differ slightly in size and plumage. Individuals of any race are somewhat variable and the differences between subspecies are clinal, especially in terms of body size. Other than these characteristics, there is little variation across the range of the species.[5] Some recent studies have gone so far as to propose that only two subspecies be recognized based on genetic markers: Aquila chrysaetos chrysaetos (including A. c. homeyeri) and A. c. canadensis (including the races A. c. japonica, A. c. daphanea and A. c. kamtschatica).[40]

The larger Middle Pleistocene golden eagles of France (and possibly elsewhere) are referred to a paleosubspecies Aquila chrysaetos bonifacti, and the huge specimens of the Late Pleistocene of Liko Cave (Crete) have been named Aquila chrysaetos simurgh (Weesie, 1988).[48][49][50] Similarly, an ancestral golden eagle, with a heavier, broader skull, larger wings and shorter legs when compared to modern birds, has been found in the La Brea Tar Pits of southern California.[51]

Habitat

Beinn Mhor on the Isle of Mull, Scotland is typical golden eagle habitat: rugged and mountainous.

Golden eagles are fairly adaptable in habitat but often reside in areas with a few shared ecological characteristics. They are best suited to hunting in open or semi-open areas and search them out year-around. Native vegetation seems to be attractive to them and they typically avoid developed areas of any type from urban to agricultural as well as heavily forested regions. In desolate areas (e.g., the southern Yukon), they can occur regularly at roadkills and garbage dumps.[52] The largest numbers of golden eagles are found in mountainous regions today, with many eagles doing a majority of their hunting and nesting on rock formations. However, they are not solely tied to high elevations and can breed in lowlands if the local habitats are suitable. Below are more detailed description of habitats occupied by golden eagles in both continents where they occur.[2]

Eurasia

In the Arctic fringe of the great continent, golden eagles occur along the edge of the tundra and the taiga from the Kola peninsula to Anadyr in eastern Siberia, nesting in forests and hunting over nearby arctic heathland. Typical vegetation is stunted, fragmented larch woodland merging into low birch-willow scrub and various heathland. In the rocky, wet, windy maritime countries of the British Isles and western Scandinavia, the golden eagle is a mountain-dwelling bird. These areas include upland grasslands, blanket bog and sub-Arctic heaths but also fragmented woodland and woodland edge, including boreal forests. In Western Europe, golden eagle habitat is dominated by open, rough grassland, heath and bogs, in places enlivened by rocky ridges, spurs, crags, scree, slopes and grand plateaux. In Sweden, Finland, the Baltic States, Belarus and almost the entire distribution in Russia all the way to the Pacific Ocean, golden eagles occur sparsely in lowland taiga forest. These areas are dominated by stands of evergreens such as pine, larch and spruce, occasionally supplemented by birch and alder stands in southern Scandinavia and the Baltic States. This is largely marginal country for golden eagles and they occur where tree cover is thin and abuts open habitat. Golden eagle taiga habitat usually consists of extensive peatland formations caused by poorly drained soils. In central Europe, golden eagles today occur almost exclusively in the grand mountain ranges, such as Pyrenees, Alps, Carpathians and the Caucasus. Here, the species nests near the tree line and hunt subalpine and alpine pastures, grassland and heath above. Golden eagles also occur in moderately mountainous habitat along the Mediterranean Sea, from Iberia and the Atlas Mountains in Morocco, to Greece, Turkey and Kurdistan. This area is characterized by low mountains, Mediterranean maquis vegetation and sub-temperate open woodland in various stages of degradation. The local pine-oak vegetation, with a huge variety of Sclerophyllous shrubs are well-adapted to prolonged summer draughts. From Kurdistan and the southern Caspian Sea to the foothills of the Hindu Kush Mountains in Afghanistan, the typical golden eagle habitat is temperate desert-like mountain ranges surrounded by steppe landscapes interspersed with forest. Here the climate is colder and more continental than around the Mediterranean. Golden eagles occupy the alpine ranges from the Altai Mountains and the Pamir Mountains to Tibet, in the great Himalayan massif, and northwestern China, where they occupy the Tien Shan range. In these mountain ranges, the species often lives at very high elevations, living above tree line at more than 2,500 m (8,200 ft), often nesting in rocky scree and hunting in adjacent meadows. In Tibet, golden eagles inhabit high ridges and passes in the Lhasa River watershed, where it regularly joins groups of soaring Himalayan vultures (Gyps himalayensis).[53] One golden eagle was recorded circling at 6,190 m (20,310 ft) above sea-level in Khumbu in May 1975.[54] In the mountains of Japan and Korea, the golden eagle occupies deciduous scrub woodland and carpet-like stands of Siberian dwarf pine (Pinus pumila) that merge into grasslands and alpine heathland. The golden eagle occurs in mountains from the Adrar Plateau in Mauritania to northern Yemen and Oman where the desert habitat is largely bereft of vegetation but offers many rocky plateaus to support both the eagles and their prey. In Israel, their habitat is mainly rocky slopes and wide wadi areas, chiefly in desert and to a lesser extent in semi-desert and Mediterranean climates, extending to open areas.[55] In Northeastern Africa, the habitat is often of a sparse, desert-like character and is quite similar to the habitat in Middle East and the Arabian peninsula. In Ethiopia's Bale Mountains, where the vegetation is more lush and the climate is clearly less arid than in Northeastern Africa, the golden eagle occupies verdant mountains.[2]

North America

The ecozones occupied by golden eagles are roughly concurrent with those of Eurasia. In western and northern Alaska and northern Canada to the Ungava Peninsula in Quebec, the eagles occupy the Arctic fringe of North America (the species does not range into the true high Arctic tundra), where open canopy gives way to dwarf-shrub heathland with cottongrass and tussock tundra. In land-locked areas of the sub-Arctic, golden eagles are by far the largest raptor. From the Alaska Range to Washington and Oregon, it is often found in high mountains above the tree line or on bluffs and cliffs along river valleys below the tree line.[56][57] In Washington state, golden eagles can be found in clear-cut sections of otherwise dense coniferous forest zones with relatively little annual precipitation.[58] From east of the Canadian Rocky Mountains to the mountains of Labrador, the golden eagle is found in small numbers in boreal forest peatlands and similar mixed woodland areas. In the foothills of the Rocky Mountains in the United States are plains and prairies where golden eagles are widespread, especially where there's a low human presence. Here, grassland on low rolling hills and flat plains are typical, interrupted only by cottonwood stands around river valleys and wetlands where the eagles may build their nests. Golden eagles also occupy the desert-like Great Basin from southern Idaho to northern Arizona and New Mexico. In this habitat, trees are generally absent other than junipers with vegetation being dominated by sagebrush (Artemisia) and other low shrub species. Although the vegetation varies a bit more, similar habitat is occupied by golden eagles in Mexico.[59][60][61] However, golden eagles are typically absent in North America from true deserts, like the Sonora Desert, where annual precipitation is less than 20 cm (7.9 in).[62] Golden eagles occupy the mountains and coastal areas of California and Baja California in Mexico where hot, dry summers and moist winters are typical. The golden eagles here often nest in chaparral and oak woodland, oak savanna and grassland amongst low rolling hill typified by diverse vegetation.[63] In the Eastern United States, the species once bred widely in the Appalachian Plateau near burns, open marshes, meadows, bogs and lakes.[64][65] In Eastern North America, the species still breeds on the Gaspe Peninsula, Quebec. Until 1999, a pair of golden eagles were still known to nest in Maine but they are now believed to be absent as a breeding bird from the Eastern United States.[2] The golden eagles who breed in eastern Canada winter on montane grass and heath fields in the Appalachian Plateau region, especially in Pennsylvania, New York, West Virginia, Maryland and Virginia. Most sightings in the Eastern United States recently are concentrated within or along southwestern border of the Appalachian Plateau (30% of records) and within the Coastal Plain physiographic region (33% of records).[66]

Though they do regularly nest in the marsh-like peatland of the boreal forest, golden eagles are not generally associated with wetlands and, in fact, they can be found near some of the most arid spots on earth. In the wintering population of Eastern United States, however, they are often associated with steep river valleys, reservoirs, and marshes in inland areas as well as estuarine marshlands, barrier islands, managed wetlands, sounds, and mouths of major river systems in coastal areas. These wetlands are attractive due to a dominance of open vegetation, large concentrations of prey, and the general absence of human disturbance.[66] In the midwestern United States, they are not uncommon during winter near reservoirs and wildlife refuges that provide foraging opportunities at waterfowl concentrations.[67]

Dietary biology

Golden eagles usually hunt during daylight hours, but were recorded hunting from one hour before sunrise to one hour after sunset during the breeding season in southwestern Idaho.[68] The hunting success rate of golden eagles was calculated in Idaho, showing that, out of 115 hunting attempts, 20% were successful in procuring prey.[69] A fully-grown golden eagle requires about 230 to 250 g (8.1 to 8.8 oz) of food per day but in the life of most eagles there are cycles of feast and famine, and eagles have been known to go without food for up to a week and then gorge on up to 900 g (2.0 lb) at one sitting.[3]

Activity and movements

In Spain, golden eagles such as this one in the Province of Ávila are sedentary.

Despite the dramatic ways in which they attain food and interact with raptors of their own and other species, the daily life of golden eagles is often rather uneventful. In Idaho, adult male golden eagles were observed to sit awake on a perch for an average of 78% of daylight, whereas adult females sat on nest or perched for an average of 85% of the day.[70] During the peak of summer in Utah, hunting and territorial flights occurred mostly between 9:00 and 11:00 am and 4:00 and 6:00 pm, with the remaining 15 or so hours of daylight spent perching or resting.[23] When conditions are heavily anticyclonic, there is less soaring during the day.[2] During winter in Scotland, golden eagles soar frequently in order to scan the environment for carrion.[2] In the more wooded environments of Norway during autumn and winter, much less aerial activity is reported, since the eagles tend to avoid detection by actively contour-hunting rather than looking for carrion.[71] Golden eagles are believed to sleep through much of the night. Although usually highly solitary outside of the bond between breeding pairs, exceptionally cold weather in winter may cause eagles to put their usual guard down and perch together. The largest known congregation of golden eagles was observed on an extremely cold winter’s night in eastern Idaho when 124 individuals were observed perched closely along a line of 85 power poles.[72]

Migration

Most populations of golden eagles are sedentary, but the species is actually a partial migrant. Golden eagles are very hardy species, being well adapted to cold climates, however they cannot abide declining available food sources in the northern stretches of their range. Eagles raised at latitudes greater than 60° N are usually migratory, though a short migration may be untaken by those who breed or hatch at about 50° N.[73] During migration, they often use soaring-gliding flight, rather than powered flight.[73] In Finland, most banded juveniles move between 1,000 and 2,000 km (620 and 1,240 mi) due south, whereas adults stay locally through winter. Further east, conditions are too harsh for even wintering territorial adults.[2] Golden eagles that breed from the Kola peninsula to Anadyr in the Russian Far East migrate south to winter on the Russian and Mongolian steppes, and the North China Plains. The flat, relatively open landscapes in these regions hold relatively few resident breeding golden eagles.[74] Similarly the entire population of golden eagles from northern and central Alaska and northern Canada migrates south. At Mount Lorette in Alberta, approximately 4,000 golden eagles may pass during the fall, the largest recorded migration of golden eagles on earth.[75] Here the mountain ranges are relatively moderate and consistent, thus being reliable for thermals and updrafts which made long-distance migrating feasible.[75] Birds hatched in Denali National Park in Alaska traveled from 818 to 4,815 km (508 to 2,992 mi) to their winter ranges in western North America.[73] These western migrants may winter anywhere from southern Alberta and Montana to New Mexico and Arizona and from inland California to Nebraska. Adults who bred in northeastern Hudson Bay area of Canada reached their wintering grounds, which range from central Michigan to southern Pennsylvania to northeastern Alabama, in 26 to 40 days, with arrival dates from November to early December.[76] The departure dates from wintering grounds are variable. In southwestern Canada, they leave their wintering grounds by April 6 to May 8 (the mean being April 21); in southwestern Idaho, wintering birds leave from March 20 to April 13 (mean of March 29); and in the Southwestern United States, wintering birds may depart by early March.[2][73][77] Elsewhere in the species' breeding range, golden eagles (i.e., those who breed in the contiguous Western United States, all of Europe but for Northern Scandinavia, North Africa and all of Asia but for Northern Russia) are non-migratory and tend to remain within striking distance of their breeding territories throughout the year.[5] In Scotland, among all recovered, banded golden eagles (36 out of 1000, the rest mostly died or disappeared) the average distance between ringing and recovery was 44 km (27 mi), averaging 63 km (39 mi) in juveniles and 36 km (22 mi) in older birds.[2] In the dry Southwestern United States, golden eagles tend to move to higher elevations once the breeding season is complete.[28] In North Africa, populations breeding at lower latitudes, like Morocco, are mostly sedentary, although some occasionally disperse after breeding to areas outside of the normal breeding range.[78]

Territoriality

Two golden eagles in an aerial conflict over their home ranges, the upper bird clearly a juvenile.

Territoriality is believed to be the primary cause of interactions and confrontations between non-paired golden eagles. Golden eagles maintain some of the largest known home ranges (or territories) of any bird species but there is much variation of home range size across the range, possibly dictated by food abundance and habitat preference. Home ranges in most of the range can vary from 20 to 200 km2 (7.7 to 77.2 sq mi).[79] In San Diego County in California, the home ranges varied from 49 to 137 km2 (19 to 53 sq mi), with an average of 93 km2 (36 sq mi).[80] However, some home ranges have been much smaller, such as in southwestern Idaho where, possibly due to an abundance of jackrabbits, home ranges as small as 4.85 km2 (1.87 sq mi) are maintained.[60] The smallest known home ranges on record for golden eagles are in the Bale Mountains of Ethiopia, where they range from 1.5 to 9 km2 (0.58 to 3.47 sq mi).[81] 46% of undulating displays in Montana occurred shortly after the juvenile eagles left their parents range, suggesting that some residents defend and maintain territories year-round.[82] Elsewhere it is stated that home ranges are less strictly maintained during winter but hunting grounds are basically exclusive.[2] In Israel and Scotland, aggressive encounters peaked from winter until just before egg-laying and were less common during the nesting season.[2][83] Threat displays include undulating flight and aggressive direct flapping flight with exaggerated downstrokes.[24] Most displays by mature golden eagles (67% for males and 76% for females) occur, rather than around the nest, at the edge of their home ranges.[70] In Western Norway, most recorded undulating flight displays occur during the pre-laying period in late winter/early spring.[71] Display flights seem to be triggered by the presence of other golden eagles.[2] The use of display flights has a clear benefit in that it lessens the need for physical confrontations, which can be fatal.[70] Usually, non-breeding birds are treated aggressively by the golden eagle maintaining their home range, normally being chased to the apparent limit of the range but with no actual physical contact.[2] The territorial flight of the adult golden eagle is sometimes preceded or followed by intense bouts of undulating displays. The invader often responds by rolling over and presenting talons to the aggressor. Rarely, the two eagles will lock talons and tumble through the air; sometimes fall several revolutions and in some cases even tumble to the ground before releasing their grip.[71][84] In some parts of the Alps, the golden eagle population has reached the saturation point in appropriate habitat and apparently violent confrontations are more common than in other parts of the range.[85] Golden eagles may express their aggression via body language while perched, typically the adult female when confronted by an intruding eagle: the head and body are upright, feathers on head and neck are erect; the wings may be slightly spread and beak open; often accompanied by intense gaze. They then often engage in a similar posture with wings spread wide and oriented toward the threat; sometimes rocking back on tail and even flopping over onto the back with talons extended upward as defense. Such behavior may be accompanied by wing slap against the threatening intruder.[24] When approached by an intruder, the defending eagle turns away, partially spreads tail, lowers head, and remains still; adults on the nest may lower head and “freeze” when approached by a person or a helicopter.[24] Females in Israel displayed more than males and mostly against interspecific intruders; males apparently displayed primarily as part of courtship.[83] Five of 7 aggressive encounters at carcasses during winter in Norway were won by females; in 15 of 21 conflicts, the younger bird dominated an older conspecific.[86] However, obvious juvenile eagles (apparent to the adult eagles due to the amount of white on their wings and tail) are sometimes allowed to penetrate deeply into a pair’s home range and all parties commonly ignore each other.[87][88] In North Dakota, it was verified that parent eagles were not aggressive towards their own young after the nesting period and some juveniles stayed on their parents territory until their 2nd spring and then left by their own accord.[89]

Reproduction

Eyrie (in hollow at left center) in the Valley of the Siagne de la Pare, Alpes-Maritimes, France

Golden eagles usually mate for life. A breeding pair is formed in a courtship display. This courtship includes undulating displays by both in the pair, with the male bird picking up a piece of rock or a small stick, and dropping it only to enter into a steep dive and catch it in mid-air, repeating the maneuver 3 or more times. The female takes a clump of earth and drops and catches it in the same fashion.[2][46] Golden eagles typically build several eyries within their territory (preferring cliffs) and use them alternately for several years. Their nesting areas are characterized by the extreme regularity of the nest spacing.[90] Mating and egg-laying timing for golden eagle is variable depending on the locality. Copulation normally lasts 10–20 seconds. Mating seems to occur around 40–46 days before the initial egg-laying.[2] The golden eagle chick may be heard from within the egg 15 hours before it begins hatching. After the first chip is broken off of the egg, there is no activity for around 27 hours. After this period, the hatching activity accelerates and the shell is broken apart in 35 hours. The chick is completely free in 37 hours.[24] In the first 10 days, chicks mainly lie down on the nest substrate.[24] The eagles are capable of preening on their second day but are continually thermoregulated via brooding by their parents until around 20 days.[2] Within 10 days, the hatchlings grow considerably, weighing around 500 g (1.1 lb).[91] Around this age, they also start sitting up more.[24] Around 20 days of age, the chicks generally start standing, which becomes the main position over the course of the next 40 days.[24] The whitish down continues until around 25 days of age, at which point it is gradually replaced by dark contour feathers that eclipse the down and the birds attain a general piebald appearance.[2] After hatching, 80% of food items and 90% of food biomass is captured and brought to the nest by the adult male. Fledging occurs at 66 to 75 days of age in Idaho and 70 to 81 days in Scotland. The first attempted flight departure after fledging can be abrupt, with the young jumping off and using a series of short, stiff wing-beats to glide downward or being blown out of nest while wing-flapping. 18 to 20 days after first fledging, the young eagles will take their first circling flight, but they cannot gain height as efficiently as their parents until approximately 60 days after fledging. In Cumbria, young golden eagles were first seen hunting large prey 59 days after fledging. 75 to 85 days after fledging, the young were largely independent of parents. Generally, breeding success seems to be greatest where prey is available in abundance.[2]

Longevity

Golden eagles are fairly long-living birds in natural conditions. The survival rate of raptorial birds tends to increase with larger body size, with a 30–50% annual loss of population rate in small falcons/accipiters, a 15–25% loss of population rate in medium-sized hawks (e.g., Buteos or kites) and a 5% or less rate of loss in eagles and vultures. The oldest known wild golden eagle was a bird banded in Sweden which was recovered 32 years later.[2] The longest-lived known wild golden eagle in North America was 23 years and 10 months.[92] The long-lived known captive golden eagle, a specimen in Europe, survived to 46 years of age.[93] The estimated adult annual survival rate on the Isle of Skye in Scotland is around 97.5%.[94] When this extrapolated into an estimated lifespan this results in 39 and half years as the average for adult golden eagles in this area, which is probably far too high an estimate.[2] Survival rates are usually much lower in juvenile eagles than in adult eagles. In the western Rocky Mountains, 50% of golden eagles banded in the nest died by the time they were 2 and a half years and an estimated 75% died by the time they were 5 years old.[95] Near a wind turbine facility in west-central California, estimated survival rates, based on conventional telemetry of 257 individuals, were 84% for first-year eagles, 79% for 1- to 3-year-olds and adult floaters and 91% for breeders; with no difference in survival rates between sexes.[63] Survival rates may be lower for migrating populations of golden eagles.[96] A 19–34% survival rate was estimated for juvenile eagles from Denali National Park in their first 11 months.[96] The average life expectancy of golden eagles in Germany is 13 years, extrapolated from a reported mere 92.5% survival rate.[97]

Natural mortality

Natural sources of mortality are largely reported in anecdotes. On rare occasions, golden eagles have been killed by competing predators or by hunting mammalian carnivores, including the aforementioned wolverine, snow leopard, cougar, brown bear and white-tailed eagle attacks. Most competitive attacks resulting in death probably occur at the talons of other golden eagles. Nestlings and fledglings are more likely to be killed by another predator than free-flying juveniles and adults. It has been suspected that golden eagle nests may be predated more frequently by other predators (especially birds, which are often the only other large animals that can access a golden eagle nest without the assistance of man-made climbing equipment) in areas where golden eagles are regularly disturbed at the nest by humans. Jeff Watson believed that common raven occasionally eats golden eagle eggs but only in situations where the parent eagles have abandoned their nesting attempt.[2] However, there are no confirmed accounts of predation by other bird species on golden eagle nests.[2] Occasionally, golden eagles may be killed by their prey in self-defense. There is an account of a golden eagle dying from the quills of a North American porcupine (Erethizon dorsatum) it had attempted to hunt.[98] On the Isle of Rùm in Scotland, there are few cases of red deer trampling golden eagles to death, probably the result of a hind having intercepted a bird that was trying to kill a fawn.[99] Although usually well out-matched by the predator, occasionally other large birds can put up a formidable fight against a golden eagle. An attempted capture of a great blue heron by a golden eagle resulted in the death of both birds from wounds sustained in the ensuing fight.[100] There is at least one case in Scotland of a golden eagle dying after being “oiled” by a northern fulmar, a bird whose primary defense against predators is to disgorge an oily secretion which may inhibit the predator's ability to fly.[101] Of natural sources of death, starvation is probably under-reported. 11 of 16 dead juvenile eagles which had hatched in Denali National Park had died of starvation.[96] Of 36 deaths of golden eagles in Idaho, 55% were possibly attributable to natural causes, specifically 8 (26%) from unknown trauma, 3 (10%) from disease and 6 (19%) from unknown causes.[102] Of 266 golden eagle deaths in Spain, only 6% were from unknown causes that could not directly attributed to human activities.[103] Avian cholera caused by bacteria (Pasteurella multocida) infects eagles that eat waterfowl that have died from the disease. The protozoan Trichomonas sp. caused the deaths of 4 fledglings in a study of wild golden eagles in Idaho.[104] Several further diseases that contribute to golden eagle deaths have been examined in Japan.[105] A captive eagle died from two malignant tumors – one in the liver and one in the kidney.[106]

In human culture

See main article : Golden eagles in human culture.

1870s illustration of burkut falconers in Kazakhstan

As early as recorded history, mankind was fascinated by the eagle. Most early recorded cultures regarded the golden eagle with reverence. It was only after the Industrial Revolution, when sport-hunting became widespread and commercial stock farming became internationally common, that humans started to widely regard golden eagles as a threat to their livelihoods. This period also brought about the firearm and industrialized poisons, which made it easy for humans to kill the evasive and powerful birds. The following are various reportages of the significance of eagles, many likely pertaining to the golden eagles, in early cultures and older religions as well as national and military insignias.

Status and conservation

See main article: Status and conservation of the golden eagle

In flight in Czech Republic

At one time, the golden eagle lived in a great majority of temperate Europe, North Asia, North America, North Africa, and Japan. Although widespread and quite secure in some areas, in many parts of the range golden eagles have experienced sharp population declines and have even been extirpated from some areas. The total number of individual golden eagles from around the range is estimated to range somewhere between 170,000 and 250,000 while the estimated total number of breeding pairs ranges from 60,000 to 100,000.[2][107] Few other eagle species are as numerous, though some species like tawny eagle, wedge-tailed eagle and bald eagle have total estimated populations of a similar size to the golden eagle’s despite having distributions which are more restricted.[3][5] The world’s most populous eagle may be the African fish eagle (Haliaeetus vocifer), which has a stable total population estimated at 300,000 individuals and is found solely in Africa.[3][108] On a global scale, the golden eagle is not considered threatened by the IUCN.[1]

References

  1. 1 2 BirdLife International (2013). "Aquila chrysaetos". IUCN Red List of Threatened Species. Version 2013.2. International Union for Conservation of Nature. Retrieved 26 November 2013.
  2. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 Watson, Jeff (2010). The Golden Eagle. A&C Black. ISBN 978-1-4081-1420-9.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 Brown, Leslie; Amadon, Dean (1986). Eagles, Hawks and Falcons of the World. Wellfleet. ISBN 978-1555214722.
  4. ↑ "Golden Eagle, Life History". All About Birds. Cornell Lab of Ornithology. 2009. Retrieved 26 December 2010.
  5. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 Ferguson-Lees, James; Christie, David A. (2001). Raptors of the World. Houghton Mifflin Harcourt. ISBN 0-618-12762-3.
  6. ↑ Peterson, Roger Tory (1998). A Field Guide to Western Birds: A Completely New Guide to Field Marks of All Species Found in North America West of the 100th Meridian and North of Mexico. Houghton Mifflin Harcourt. pp. 180–. ISBN 978-0-395-91174-7.
  7. ↑ National Geographic Field Guide To The Birds Of North America (4th ed.). 2002. ISBN 978-0-79226-877-2.
  8. ↑ del Hoyo, J; Elliot, A; Sargatal, J (1996). Handbook of the Birds of the World. 3. Barcelona: Lynx Edicions. ISBN 84-87334-20-2.
  9. 1 2 3 4 5 6 7 8 Wood, Gerald (1983). The Guinness Book of Animal Facts and Feats. ISBN 978-0-85112-235-9.
  10. ↑ "Golden Eagle May Set Record". Newwest. Retrieved June 26, 2015.
  11. ↑ Sagip Eagle Archived September 28, 2013, at the Wayback Machine.. Gbgm-umc.org. Retrieved on 2012–08–22.
  12. ↑ "Birds of North America Online—Birds of North America Online". Bna.birds.cornell.edu. Retrieved 2012-11-08.
  13. 1 2 Bortolotti, G.R. (1984). "Age and sex size variation in Golden Eagles". Journal of Field Ornithology. 55 (1): 54–66. JSTOR 4512857.
  14. ↑ Fowler, J.M.; Cope, J.B. (1964). "Notes on the Harpy Eagle in British Guiana". Auk. 81 (3): 257–273. doi:10.2307/4082683. JSTOR 4082683.
  15. ↑ Nye, Peter. "Journey North Bald Eagles". Journey North. Retrieved 2013-04-22.
  16. 1 2 Terres, John K. (1980). "Golden Eagle". The Audubon Society Encyclopedia of North American Birds. New York: Alfred A. Knopf. p. 478. ISBN 0-517-03288-0.
  17. ↑ Clark, William S.; Wheeler, Brian K. (2001). A field guide to hawks of North America. New York: Houghton Mifflin Company. p. 241. ISBN 0-395-67067-5.
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  19. ↑ Clark, William S (1983). "The field identification of North American eagles" (PDF). North American Birds. 37 (5): 822–826. Retrieved 27 December 2010.
  20. 1 2 Liguori, Jerry (2004). "How to Age Golden Eagles" (PDF). Birding: 278–283. Retrieved 27 December 2010.
  21. 1 2 Tjernberg, M. (1988). "Age determination of Golden Eagles, Aquila chrysaetos". Vår fågelvärld. 47: 321–334. ISSN 0042-2649.
  22. ↑ Ellis, D.H.; Lish, J.W. (2006). "Thinking about feathers: adaptations of golden eagle retrices". Journal of Raptor Research. 40 (1): 1–28. doi:10.3356/0892-1016(2006)40[1:TAFAOG]2.0.CO;2.
  23. 1 2 3 4 5 6 Brown, L. (1976). Eagles of the World. David and Charles. ISBN 0360003184.
  24. 1 2 3 4 5 6 7 8 Ellis, D.H. (1979). Development of behaviour in the Golden Eagle. Wildlife Monographs. 70. Wiley. JSTOR 3830587.
  25. 1 2 "Golden Eagle Fact Sheet". San Diego Zoo. Retrieved 2013-05-01.
  26. ↑ ADW: Aquila chrysaetos: INFORMATION. Animaldiversity.ummz.umich.edu. Retrieved on 2012–08–22.
  27. ↑ Darling, F.F. (1934). "Speed of a Golden Eagle's flight". Nature. 134 (3383): 325–6. doi:10.1038/134325c0.
  28. 1 2 Palmer, R.S. (1988). "Golden eagle: Aquila chrysaetos". Family Accipitridae (concluded) Buteos, Golden Eagle, Family Falconidae, Crested Caracara, Falcons. Handbook of North American birds. 5. Yale University Press. pp. 180–231. ISBN 0300040601.
  29. ↑ Cornell University. Bna.birds.cornell.edu. Retrieved on 2012–08–22.
  30. 1 2 3 4 Sutton, C.; Dunne, P.; Sibley, D. (1989). Hawks in Flight: The Flight Identification of North American Migrant Raptors. Boston: Houghton Mifflin Harcourt. ISBN 0-3955-1022-8.
  31. ↑ Linnaeus, C. (1758). Systema Naturæ per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Volume 1 (in Latin) (10th ed.). Holmiae:Laurentii Salvii. p. 88. [Falco] cera lutea, pedibus lanatis, corpore fusco ferrugineo vario, cauda nigra basi cinereo-undulata.– (A [diurnal raptor] with yellow cere, [feathered tarsometatarsus], body dusky brown variegated with rusty, tail black with ashy-waved base.)
  32. ↑ Brisson, Mathurin-Jacques; Martinet, François Nicolas (1760). Ornithologie; ou, Méthode contenant la division des oiseaux en ordres, sections, genres, espéces & leurs variétés. &c. Paris: C.J.B. Bauche. pp. 28, 419.
  33. ↑ Jobling, James A (2010). The Helm Dictionary of Scientific Bird Names. London: Christopher Helm. pp. 52, 104. ISBN 978-1-4081-2501-4.
  34. 1 2 3 4 Helbig, A.J.; Kocum, A.; Seibold, I.; Braun, M.J. "A multi-gene phylogeny of aquiline eagles (Aves: Accipitriformes) reveals extensive paraphyly at the genus level" (PDF). Molecular Phylogenetics and Evolution. 35: 147–164. doi:10.1016/j.ympev.2004.10.003. Archived from the original (PDF) on August 13, 2012.
  35. 1 2 3 Lerner, H.R.; Mindell, D.P. (November 2005). "Phylogeny of eagles, Old World vultures, and other Accipitridae based on nuclear and mitochondrial DNA" (PDF). Mol. Phylogenet. Evol. 37: 327–46. doi:10.1016/j.ympev.2005.04.010. PMID 15925523.
  36. ↑ "Lophaetus pomarinus". The Peregrine Fund. Retrieved 2013-04-22.
  37. ↑ "Aquila morphnoides". The Peregrine Fund. Retrieved 2013-04-22.
  38. ↑ "Aquila wahlbergi". The Peregrine Fund. Retrieved 2013-04-23.
  39. ↑ "Aquila africana". The Peregrine Fund. Retrieved 2013-04-23.
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  41. ↑ "Birds of Kazakhstan. Golden Eagle (Aquila chrysaetos chrysaetos).". birds.kz. Retrieved 2013-04-23.
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  48. ↑ Simurgh is a mythical bird. Some say the Persian word simurgh or its Middle Persian form sēn-murw also meant golden eagle; see "Simorğ" in Encyclopædia Iranica.
  49. ↑ Weesie, P.D.M. (1988). "The Quaternary avifauna of. Crete, Greece". Palaeovertebrata. 18 (1): 1–94.
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  51. ↑ Howard, H. (1947). "An ancestral Golden Eagles a question in taxonomy". Auk. 64 (2): 287–291. doi:10.2307/4080550. JSTOR 4080550.
  52. ↑ Burles, D. W. & Frey, R. (1981). Raptor monitoring program, 1980 report. Kluane National Park, YT.
  53. ↑ Lang, A.; Bishop, M.A.; Le Seuer, A. (2007). "An annotated list of birds wintering in the Lhasa river watershed and Yamzho Yumco, Tibet Autonomous Region, China" (PDF). Forktail. 23.
  54. ↑ Inskipp, C.; Inskipp, T. (1991). A guide to the birds of Nepal (2nd ed.). Smithsonian Institution Press. ISBN 0713681098.
  55. ↑ Shirihai, H.; Roselaar, C.S. (1996). The Birds of Israel: A Complete Avifauna and Bird Atlas of Israel. Academic Press. ISBN 0126402558.
  56. ↑ Ritchie, R.J.; Curatolo, J.A. (1982). "Notes on Golden Eagle productivity and nest site characteristics, Porcupine River, Alaska, 1979–1982" (PDF). Raptor Research. 16 (4): 123–7.
  57. ↑ Petersen, M.R.; Weir, D.N.; Dick, M.H. (1991). "Birds of the Kilbuck and Ahklun Mountain Region, Alaska". North American Fauna. 76: 1–158. doi:10.3996/nafa.76.0001.
  58. ↑ Marr, N.V.; Knight, R.L. (1983). "Food Habits of Golden Eagles in Eastern Washington". Murrelet. 64 (3): 73–77. doi:10.2307/3535265. JSTOR 3535265.
  59. ↑ Craig, T.H.; Craig, E.H. (1984). "Results of a helicopter survey of cliff nesting raptors in a deep canyon in southern Idaho" (PDF). Journal of Raptor Research. 18 (1): 20–25.
  60. 1 2 Marzluff, J.M.; Knick, S.T.; Vekasy, M.S.; Schueck, L.S.; Zarriello, T.J. (1997). "Spatial use and habitat selection of Golden Eagles in southwestern Idaho" (PDF). Auk. 114 (4): 673–687. doi:10.2307/4089287. JSTOR 4089287.
  61. ↑ Millsap, B.A. (1981). Distributional status of Falconiformes in west central Arizona-with notes on ecology, reproductive success and management. Technical Note. 355. U.S. Department of the Interior, Bureau of Land Management.
  62. ↑ Root, T.L. (1988). Atlas of wintering North American birds; an analysis of Christmas Bird Count data. University of Chicago Press. ISBN 0226725391.
  63. 1 2 Hunt, W. G., Jackman, R.E. Brown, T. L., Gilardi, J. G., Driscoll, D. E. & Culp, L. 1995. A pilot Golden Eagle population study in the Altamont Pass Wind Resource Area, California. Predatory Bird Research Group, University of California, Santa Cruz.
  64. ↑ Spofford, W.R. (1971). "The breeding status of the Golden Eagle in the Appalachians". American Birds. 25: 3–7.
  65. ↑ Singer, F.J. (1974). "Status of the Osprey, Bald Eagle, and Golden Eagle in the Adirondacks". New York Fish & Game Commission Journal. 21: 18–31.
  66. 1 2 Millsap, B.A.; Vana, S.L. (1984). "Distribution of wintering Golden Eagles in the eastern United States". Wilson Bulletin. 96 (4): 692–701. JSTOR 4162004.
  67. ↑ Wingfield, G.A. (1991). "Central plains buteos and Golden Eagle". In Pendleton, B.G. Proceedings of the Midwest raptor management symposium and workshop. National Wildlife Federation. pp. 60–68. ISBN 0945051506.
  68. ↑ Dunstan, T. C., Harper, J.H. & Phipps, K.B. 1978. Habitat use and hunting strategies of Prairie Falcons, Red-tailed Hawks, and Golden Eagles. Fin. Rep. Western Illinois University, Macomb.
  69. ↑ Collopy, M.W. "Foraging Behavior and Success of Golden Eagles" (PDF). Auk. Retrieved 2013-04-24.
  70. 1 2 3 Collopy, M.W.; Edwards Jr., T.C. (1989). "Territory size, activity budget, and role of undulating flight in nesting Golden Eagles" (PDF). Journal of Field Ornithology. 60 (1): 43–51.
  71. 1 2 3 Bergo, G. (1987). "Territorial behaviour of Golden Eagles in western Norway" (PDF). British Birds. 80 (8): 361–376.
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